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PROBOSCIDEA
 RESTORATION OF THE JEFFERSONIAN MAMMOTH {PARELEPHAS JEFFERSONII)
After a painting by Charles R. Knight in 1909, under the (Urertion of Henry Fairfield Osborn

The skeleton on which this painting was based is moulded in the American Museum of Natural Histuri/, Hull of
theAge of Man. It was found near Jonesboro, Indiana, on the farm of Dora E. Gift, in 1903; purchased for the
American Museum with the Jesup Fund in 1904; mounted in 1906; first described and figured by the present
author in 1907. As found the skeleton was embedded in a muck deposit of late Pleistocene age, fifteen feet below
the surface. This deposit is probably of post{?)-W i sconsi n age {according to the geologic time scale having been laid
doivn about 15,000 years ago). Representatives of the Parelephasphylum appeared in Europe in the early Pleistocene
and persisted into the Third Interglacial. This Third Inlerglacial period may mark the time of migration across
Asia into North America. In fact, it is suggested by the present author that such migration might hare occurred in
Second or eve^i in First Interglacial time.

The most striking features of this individual are the complete incurvation and cro.ssing of the tu.'iks, indicating
that is an old bull, and the relatively small size of the head.
it It is here represented with a hairy covering, as Parele-
phas is characteristic of the north temperate region both of Europe a nd the ( 'n ited States.
,
 PROBOSCIDEA
A MONOGRAPH OF THE DISCOVERY, EVOLUTION, MIGRATION
AND EXTINCTION OF THE MASTODONTS AND
ELEPHANTS OF THE WORLD

BY

HENRY FAIRFIELD OSBORN

A.B. Princeton, 1877; D.Sc. Princeton, 1880; Honorary LL.D. Trinity, 1901; LL.D. Princeton, 1902;
Sc.D. Cambridge, 1904; LL.D. Columbia, 1907; Ph.D. Christiania, 1911; D.Sc. Yale, 1923;
D.Sc. Oxford, 1926; D.Sc. New York, 1927; LL.D. Union, 1928; Doctor
Natural
OF THE University of Paris, 1931; Doctor of Science,
Johann Wolfgang Goethe University, 1934
Research Professor of Zoology, Columbia University; Honorary Curator-in-Chief of Vertebrate
Palaeontology, The American Museum of Natural History; Senior Palaeontologist,
United States Geological Survey; Honorary President, The American
Museum of Natural History; Honorary President,
The New York Zoological Society

EDITED BY MABEL RICE PERCY

VOLUME II

STEGODONTOIDEA
ELEPHANTOIDEA

PUBLISHED ON THE J. PIERPONT MORGAN FUND BY THE TRUSTEES

OF THE AMERICAN MUSEUM OF NATURAL HISTORY
THE AMERICAN MUSEUM PRESS
NEW YORK, 1942
 PUBLICATION NOTE
Volume I of this work was issued August 15, 1936. The present volume,
containing the Stegodontoidea and Elephantoidea as well as tables, conclusions,
and general index, has been prepared from the materials left by the late author
(see Publication Note to Volume I).

Copyright, 1942, by
The American Museum of Natural History
 CONTENTS
PAGE
LIST OF ILLUSTRATIONS xv

CHAPTER
XIV. THE ROOF-TOOTHED STEGODONTS, SUPERFAMILY STEGODONTOIDEA 805
Classification of the genera Stegolophodon and Stegodon 807
History of discovery of the subfamily Stegodontinae. Principles of type revision of species 815
The Stegodontinse and Mastodontinse of China 816
Pliocene to Pleistocene Proboscidea of Japan 818
Phylogenetic discussion of the thirty described species of Stegodon ts and Stegolophodonts 819
—
Probable European Asiatic origin and migration of the primitive Stegodonts 822
Type revision of the species in order of original discovery and description 822
Firsttwo Stegodonts discovered in Burma (1928) 825
Mastodon latidens Clift, 1828 = Stegolophodon
[ latidens] 827
Mastodon elephantoides Clift, 1828 = Stegodon
[ elephantoides] 828
Discoveries in India and Burma 829
Stegodonts of China, India, Java, the Philippine Islands, Austria, Japan, and Burma 831
Characters of the subfamily Stegodontinae 837
Stegolophodon Schlesinger, 1917, generic definition 839
cautleyi Lydekker, 1886 840
latidens Clift, 1828 842
sublatidens Schlesinger, 1917 846
stegodontoides Pilgrim, 1913 846
nathotensis Osborn, 1929 847
cautleyi progressus Osborn, 1929 848
lydekkeri Osborn, 1936 851
Stegodon Falconer and Cautley, 1847, 1857, generic definition 853
sinensis Owen, 1870 860
elephantoides Clift, 1828 861
bombifrans Falconer and Cautley, 1846 863
insignis Falconer and Cautley, 1845, 1846 866
ganesa Falconer and Cautley, 1845, 1846 869
insignis-ganesa 874
insignis birmanicus Osborn, 1929 874
orientalis grangeri Osborn, 1929 875
pinjorensis Osborn, 1929 883
orientalis Owen, 1870 884
airdwana Martin, 1890 885
ganesa var. javanicus Dubois, 1908 889
trigonocephalus Martin, 1887 890
mindanensis Naumann, 1890 892
auroi-B Matsumoto, 1915, 1918 892
Matsumoto, 1924
orientalis shodoensis 893
van der Maarel, 1932
bondolensis
894
trigonocephalus praecursor von Koenigswald, 1933 896
(Parastegodon?) kwantoensis Tokunaga, 1934 897
yiishensis Young, 1935
897
898
officinalis Hopwood, 1935
zdanskyi Hopwood, 1935
899
(Parastegodon) sugiyarnai Tokunaga, 1935 899
901
Matsumoto on the phylogeny and classification of the Japanese Mastodonts, Stegodonts, and Elephants
908
Osborn's comments (1929) on Matsumoto's phylogeny and classification of 1924-1927

XV. CLASSIFICATION OF THE ELEPHANTOIDEA BY THEIR DIVERGENT AND HARMONIC CRANIAL

911
AND DENTAL CHARACTERS
912
Elephantoidea Osborn, 1921, superfamily definition
912
Elephantidae Gray, 1821, family definition

vii
i
OSBORN: THE PROBOSCIDEA

AFTER XV — Continued page

Failure of previous dental classifications 914
Classification by cranial and dental ciiaracters 914
Cranial mechanics of Elephas (Weithofer, Osborn, Gregory) 915
Comparative cranial sections of elephant skulls 918
Ontogenetic cranial changes in Elephas indicus 919
Dental and cranial adaptation to prevailing feeding habits the key to phylogenetic classification 927
Ridge-plate formulae of primitive and progressive genera in adaptation to prevailing habits of feeding 927
Food of the Indian and African elephants and of the mammoth 927
Seasonal changes in food of the mammoth 929
Summary of progression from browsing to grazing dentition 929
Vertebral distinctions of Elephas, Loxodonta, Mammonteus, and Parelephas 930
Vertebral formulae 930
Synopsis of subfamily classification of the Elephantoidea 932

XVI. THE GENUS ARCHIDISKODON (SUPERFAMILY ELEPHANTOIDEA), MOST PRIMITIVE MEMBER OF

THE SUBFAMILY MAMMONTIN^ 935
History of the subfamily Mammontinae 935
Mammontinse Osborn, 1921, subfamily definition 937
History of the genus Archidiskodon 939
Older of discovery and description of twenty-two species of Archidiskodonts 942
Archidiskodonts of Eurasia and America 943
New Archidiskodonts and Loxodonts of Africa 944
Approximate phylogenetic order of succession of species of Archidiskodon and Parelephas (1928) 946
Archidiskodon Pohlig, 1885, 1888, generic definition 947
planifrons Falconer and Cautley, 1846 [1845] 950
Measurements of twenty-seven specimens collected by Barnura Brown in the Siwaliks 954
Leith-Adamsia siwalikiensis 959
planifrons rumanus Stefanescu, 1924 968
meiidionalis Nesti, 1825 969
lyrodon {
= A. meridionalis female] 977
Durfort skeleton 977
meridionalis cromerensis Deperet and Mayet, 1923 980
Archidiskodonts and Metarchidiskodonts of South Africa 983
Archidiskodon (continued)
proplanifrons Osborn, 1934 986
subplanifrons Osborn, 1928 987
broomi Osborn, 1928 989
vanalpheni Dart, 1929 990
milletti Dart, 1929 991
loxodontoides Dart, 1929 991
yorki Dart, 1929 992
M etarchidiskodon Osborn, 1934, generic definition 994
griqua Haughton, 1922 994
Archidiskodonts of the United States and Mexico 996
Archidiskodon (continued)
imperalor Leidy, 1858 998
imperator Freudenberg, 1922
silveslris 1015
imperalor falconeri Freudenberg, 1922 1016
El. Coluinbi var. imperator Freudenberg, 1922 1017
maibeni (skeletal characters) 1019
hayi Barbour, 1915 1023
imperator srolti Barbour, 1925 1025
imperator maibeni Barbour, 1925 1027
haroldcooki Hay, 1928 1029
eiilis Stock and Furlong, 1928 1031
sonoriensis Osborn, 1929 1033
meridionalis nebrascensis Osborn, 1932 1033
 CONTENTS ix

CHAPTER PAGE
XVII. THE GENUS PARELEPHAS (SUPERFAMILY ELEPHANTOIDEA), OF THE SUBFAMILY MAMMON-
TIN^, INTERMEDIATE BETWEEN ARCHIDISKODON AND MAMMONTEUS, DISTRIBUT-
ED IN THE NORTH TEMPERATE ZONE OF EURASIA AND NORTH AMERICA 1039

European north temperate origin. History of separation from other extinct proboscideans 1039
Order of discovery and description of species of Parelephas 1047
Parelephas Osborn, 1924, generic definition 1048
trogontherioides Zuffardi, 1913 1055
trogontherii Pohlig, 1885, 1888-1891 1056
Irogontherii nestii Pohlig, 1891 1059
armeniacus Falconer, 1857 1060
intermedius Jourdan, 1861 1062
wusti Pavlow, 1909 1065
North and South American species of Parelephas 1067
jacksoni Mather, 1838 1068
{1)rmnfiissippiensis Foster, 1872 1070
Columbian Mammoth (Parelephas columhi) 1070
columhi Falconer, 1857, 1863, 1868 1071
texlanus [ = coluvihi] Owen, 1859, Blake, 1861, 1862 1073
Cohen Collection (Phosphate Beds of South Carolina) 1075
Amherst skeleton 1079
columhi felicis Freudenberg, 1922 1082
columbi cayennensis Osborn, 1929 1083
jeffersonii Osborn, 1922
1083
Elephas roosevelti [
= Parelephas jeffersonii] Hay, 1922 1095
progressus Osborn, 1924 1097
washingtonii Osborn, 1923 HOI
eellsi Hay, 1926 1104
floridanus Osborn, 1929 1105

XVIII. THE GENUS MAMMONTEUS (SUPERFAMILY ELEPHANTOIDEA), OF THE SUBFAMILY MAM-

MONTINiE, THE TRUE NORTHERN WOOLLY MAMMOTH 1117

1118
Sceleto Elephantino Tonnse, 1695, confused with the mammoth, Blumenbach,
1799
description (1735) of the Elephas primigenius of Siberia 1119
Breyne's
apphed mammoth 1120
Names successively to the
word Mammut 1124
Native Siberian origin of the
1124
Ides (1706)
112^
Howorth (1882)
1788, Osborn, 1924, generic definition
1126
Mammonteus Camper,
1127
External characters and feeding habits
1129
Skeletal characters of Mammonteus primigenius
Historical order of naming of species of Mammonteus exclusive of species which are now known to belong
^^"^^
to Parelephas trogontherii, etc
1139
Aurignacian mammoth hunters of Moravia
1140
Typical progressive Eurasiatic stages of Mammonteus
1141
primigenius Blumenbach, 1799, 1803
1146
Elephas odontotyrannus [ = M. primigenius] Eichwald, 1835
1149
Primitive European stages of Mammonteus primigenius
^^^"
primigenius leith-adamsi Pohlig, 1888
^^
primigenius hydruntinus Botti, 1891 -^

primigenius fraasi Dietrich, 1912

11^4
primigenius astensis Dep6ret and Mayet, 1923
H^^
Forest Bed, or Cromerian, fauna
American stages of Mammonteus ^^
primigenius americaiius DeKay, 1842
X OSBORN: THE PROBOSCIDEA

CHAPTER XVIII —Continued page

primigenius compressus Osborn, 1924 1157
prirnigenius alaskensis sp. nov 1159
Frozen mammoth of Siberia 1162
Summary of the discovery and natural history of the woolly mammoth 1163

XIX. THE GENERA LOXODONTA, PALiEOLOXODON, AND HESPEROLOXODON OF THE SUPERFAMILY

ELEPHANTOIDEA, SUBFAMILY LOXODONTIN^ 1171

Classification and histoiy of discovery of the Loxodontinse 1173

Difficulties of generic nomenclature 1174
Order of discovery and description of the fifty-three type species of the extinct Loxodontinae 1187
Loxodontinae Osborn, 1918, subfamily definition 1191
Loxodonta F. Cuvier, 1825, 1827, generic definition 1191
Order of description of eighteen living African species and subspecies 1192
Loxodo7ita (continued)
africana Blumenbach, 1797 1 197
africana {?)cottoni Eales, 1926-1929 1202
comaliae Aradas, 1870 1204
Palxoloxodon Matsumoto, 1924, generic definition 1207
namadicus Falconer and Cautley, 1846, 1847 1211
Hesperoloxodon Osborn, 1931, generic definition 1217
aniiquus Falconer and Cautley, 1847, 1857 1217
Upnor skeleton of H. antiquus 1222
Coadaptation of the vertebral column with the superior incisive tusks of the elephants 1228
Hesperoloxodon (continued)
antiquus nanus Acconci, 1880 1230
antiquus plaiyrhynchus Graells, 1897 1231
antiquus ausonius Major, 1875, Verri, 1886, Dep6ret and Mayet, 1923 1232
antiquus germanicus StefSnescu, 1924 1233
Hesperoloxodon antiquus italicus of Southern Italy and H. antiquus germanicus of North Central Germany .... 1238
antiquus italicus Osborn, 1931 1245
aniiquus germanicus of Steinheim 1253
Extinct dwarfed species of the Mediterranean Islands 1257
Palseoloxodon (continued)
melitensis Falconer, 1862 1262
falconeri Busk, 1867 1263
mnaidriensis Adams, 1870 1265
lamarmorae Forsyth Major, 1883 1266
Cypriotes Bate, 1903 1266
creticus Bate, 1907 1267
The dwarfed elephants of the Mediterranean Islands and the question of Pleistocene isthmuses (Vaufrey, 1929) . 1268
Ancestral stages of Palasoloxodon in Africa 1273
atlanticus Pomel, 1879 1274
jolensis Pomel, 1895 1274
recki Dietrich, 1916 1275
Palxoloxodon and Loxodonta of South Africa 1277
Palxoloxodon (continued)
andrewsi Dart, 1929 1278
hanekami Dart, 1929 1279
yorki Dart, 1929 1280
wilmani Dart, 1929 1280
kuhni Dart, 1929 1281
urchidiskodonloides Haughton, 1932 1282
iransvaulensis Dart, 1927 1284
sheppardi Dart, 1927 1285
 CONTENTS xi

CHAPTER XIX — (Continued) page

Loxodonta (continued)
Zulu Scott, 1907 1286
prima Dart, 1929 1287
africana var. obliqua Dart, 1929 1287
subantiqua Haughton, 1932 1288
Loxodontines of Japan and Java 1289
Summary of Matsumoto'sobservations and theories of 1924 and 1929
final 1290
Osborn's summary (1930) of the observations of Makiyama (1924) and of Matsumoto (1924-1929) 1292
Two Japanese subspecies described by Makiyama (1924) 1293
Pabeoloxodon namadicus naumanni Makiyama, 1924 1295
namadi Makiyama, 1924
Palseoloxodon namadicus 1296
Five Japanese Loxodontines described by Matsumoto 1297
Palseoloxodon protomammonieus Matsumoto, 1924, 1926 1297
Palxoloxodon tokunagai Matsumoto, 1929 1298
Palseoloxodon protomammonieus proximus Matsumoto, 1926 1298
Palseoloxodon namadicus yabei Matsumoto, 1929 1299
Palseoloxodon (Archidiskodonf) tokunagai mut. junior Matsumoto, 1929 1299
Japanese species described by Saheki and Tokunaga (1931, 1934) 1300
Parelephas protomammonieus malsumotoi Saheki, 1931 1300
Palseoloxodon yokohamanus Tokunaga, 1934 1301
Javanese species described by Dubois 1302
Palseoloxodon hysudrindicus Dubois, 1908 1302
Geographic distribution along the eastern coast of Asia 1304

XX. THE SUBFAMILY ELEPHANTINE (SUPERFAMILY ELEPHANTOIDEA), OF EASTERN ASIA, IN-

CLUDING THE RECENT ELEPHAS AND RELATED LOWER PLEISTOCENE SPECIES 1307
Historical introduction and nomenclature (850? B.C. 1936) — 1308
Falconer (1868) on the specific unity and vertebral formulae of the Asiatic elephants 1312
Corse, de Blainville and Falconer on characters of the geographic varieties 1313
Elephas indicus Isodactylus Hodgson 1313
Elephas indicus Heterodactylus Hodgson 1313
Elephas indicus ceylanicus de Blainville 1313
Elephas indicus bengalensis de Blainville 1313
Elephas indicus (Dauntela var.) 1314
Elephas indicus (Mukna var.) 1314
Living specific or subspecific forms, continental and insular 1315
Fossil forms more or less closely related to Elephas indicus 1318
Names of species and subspecies of the subfamily Elephantinse in order of description 1319
Elephantinse Osborn, 1910, subfamily definition 1320
Elephas Linnaeus, 1735-1758, generic definition 1322
indicus Linnaeus, 1735-1754, collective species 1323
indicus ceylanicus de Blainville, 1845 1327
indicus bengalensis de Blainville, 1845 1327
indicus sumalranus Temminck, 1847 1329
indicus hirsutus Lydekker, 1914 1332
indicus Buski[=? Palseoloxodon buski] Matsumoto, 1927 1333
Distinctions and measurements of the Indian elephant 1334
Characters of the Upper Pliocene and Lower Pleistocene species Hypselephas hysudricus and Platelephas plat-
ycephalus 1339
Hypselephas Osborn, 1936, generic definition 1340
hysudricus Falconer and Cautley, 1845, 1846 1340
Observations of Osborn on fourteen specimens collected by Barnum Brown in the Siwaliks 1345
Cranial characters and affinities 1348
Platelephas Osborn, 1936, generic definition 1358
plaiycephalus Osborn, 1929 1359
Appendix to Chapter XX 1362
xii OSBORN: THE PROBOSCIDEA

CHAPTER PAGE
XXI. NOMENCLATURE OF THE PROBOSCIDEA 1363
The American Mastodon {Mastodon americanus) 1363
The Northern or Woolly Mammoth {Mammonteus primigenius) 1365
List of superfamilies 1367
List of famiUes 1368
List of subfamilies 1369
List of genera 1371
List of species, subspecies, and varieties 1382

XXII. THE GEOLOGIC SUCCESSION OF THE PROBOSCIDEA 1421

Africa 1422
Eocene and Oligocene of North Africa 1423
Miocene of North Africa 1426
Miocene of Central and East Africa 1428
Pleistocene of North Africa 1429
Pleistocene of Central and East Africa 1432
Pleistocene of South Africa 1437
The Orient 1439
Miocene of Baluchistan and Sind 1439
The Siwalik series (Miocene-Pleistocene) of North India 1442
Pleistocene of Central India 1447
Pleistocene of Ceylon 1450
Pleistocene of Burma 1450
Pleistocene of South China 1452
Pleistocene of Indo-China 1453
Pleistocene of the East Indies 1453
Europe 1456
Lower Miocene : Burdigahan 1457
Middle Miocene : Helvetian and Tortonian —Vindobonian 1460
Upper Miocene : Sarmatian 1464
Lower Pliocene : Pontian 1466
Middle Pliocene : Plaisancian 1469
Upper Pliocene : Astian 1470
Pleistocene 1472
Asia 1477
Miocene of MongoUa and Central Asia 1477
Miocene North China
of 1479
Pliocene of Mongolia 1481
Pliocene of North China 1481
Pleistocene of North China 1483
Miocene to Pleistocene of Japan 1490
North America 1490
Upper Miocene: Barstovian 1491
Lower Pliocene Clarendonian
: 1495
Middle Pliocene Hemphillian
: 1502
Upper Pliocene Blancan
: 1503
Pliocene of Mexico 1506
Proboscideans from undetermined levels in the Miocene and Pliocene of North America 1507
North American Tertiaiy horizons containing fragmentary proboscidean remains 1508
Pleistocene of North America 1510
Pleistocene of Mexico 1515
Central and South America 1516
Pliocene of Central America 1516
Pleistocene of Argentina 1516
Pleistocene of the Andean valleys 1519
Pleistocene of Brazil and French Guiana 1521
 CONTENTS xiii

CHAPTER PAGE
XXIII. AFFINITIES, MIGRATIONS, AND PHYLOGENY OF THE PROBOSCIDEA: A SUMMARY 1523
Five superfamilies 1524
Eight families 1525
Twenty-one subfamilies 1526
Forty-four genera 1526
Valid species (352) 1527
Osborn's final (1935) classification of the Mceritherioidea, Deinotherioidea, and Mastodontoidea 1529
Osborn's final (1935) classification of the Stegodontoidea and Elephantoidea 1539
Explanation of terms used throughout the text of the present Memoir 1545
Characters, affinities, and migrations of the Proboscidea 1552
Moeritheres 1552
Deinotheres 1553
Longirostrines (genera, Trilophodon, Megabelodon; subgenera, Genomastodon, Choerolophodon, Tatabelodon).. 1555
Gnathabelodonts (Gnathabelodon) 1557
Amebelodonts {Amebelodon, Phiomia) 1558
Tetralophodonts (genera, Tetralophodon, Morrillia; subgenus, Lydekkeria) 1559
Notorostrines {Cordillerion) 1560
Rhynchorostrines {Rhynchotherium, Blickotherium, Aybelodon) 1561
Brevirostrines {Anancus, Penlalophodon, Synconolophus) 1563
Humboldtines (Cuvieronius, Eubelodon, Stegomastodon) 1566
Serridentines (Serridentinus, Ocalientinus, Se.rbelodon, Trobelodon) 1568
Platybelodonts {Platybelodon, Torynobelodon) 1570
Notiomastodonts (Notiomastodon) 1572
Palifiomastodonts {Palxomaslodon) 1572
Mastodonts {Mastodon, Miomastodon, Pliomastodon) 1574
Zygolophodonts {Zygolophodon, Turicius) 1575
Stegolophodonts (Stegolophodon) 1578
Stegodonts (Stegodon) 1579
Mammontines {Archidiskodon, Metarchidiskodon, Parelephas, Mammonteus) 1582
Loxodontines (genera, Loxodonta, Palseoloxodon, Hesperoloxodon; Sivalikia, Pilgrimia, synonyms of
Palxoloxodon) 1590
Elephantines {Elephas, Hypselephas, Platelephas) 1595
Skeletal material 1600
Heights of proboscideans, estimated and actual 1604

APPENDIX TO VOLUME II. PROBOSCIDEAN DENTAL HISTOLOGY 1607

LIST OF CATALOGUE NUMBERS OF SPECIMENS MENTIONED IN THE PROBOSCIDEA MEMOIR,

VOLUME II 1609

SUPPLEMENTARY BIBLIOGRAPHY 1623

INDEX TO VOLUMES I AND II 1631

 .

LIST OF ILLUSTRATIONS
PLATE PAGE
Frontispiece. Restoration of the Jeffersonian Mammoth {Parelephas jeffersonii). After painting by Knight.
XIII. Stegolophodontinae Stegolophodon
:
850-851
XIV. Migration and evolution of Mceritherium and Deinotherium 1552-1553
XV. Migration and evolution of Trilophodon, Megabelodon, Gnaihabelodon, Phiomia, Amehelodon, Tetralophodon, and
Morrillia 1556-1557
XVI. Migration and evolution of Cordillerion, Rhynchotherium, Blickotherium, Aybelodon, Anancus, Pentalophodon, and
Sxjnconolophus 1560-1561
XVII. Migration and evolution of Eubelodon, Cuvieronius, and Stegomastodo7i 1566-1567
XVIII. Migration and evolution of Trobelodon, Serbelodon, Serrideniinus, Ocalientinus, Plalybelodon, Torynobelodon, and
Notiomastodon 1568-1569
XIX. Migration and evolution of Palxomastodo7i, Miomaslodon, Pliomastodon, Mastodon, Turicius, Zygolophodon, and
Stegolophodon 1572-1573
XX. Migration and evolution of Stegodon 1578-1579
XXI. Migration and evolution of Archidiskodon and Metarchidiskodon 1582-1583
XXII. Migration and evolution of Parelephas, Mammonteus, and Elephas 1584-1585
XXIII. Migration and evolution of Loxodonta, Palieoloxodon, and Hesperoloxodon 1590-1591
XXIV. Osborn-Reeds Correlation Table of 1922 and 1929 1606-1607
XXV. Northwestern India and adjacent territory, showing especially the Siwalik Hills where Falconer and Cautley made
their classic collections and Dr. Barnum Brown recovered in 1922 the fine Proboscidea collections for the
American Museum 1606-1607
XXVI. Elephas indicus: (Fig. 1) Complete transverse section near tip of unerupted tusk. (Fig. 2) Area of thin section
within circle on Fig. 1. (Fig. 3) Same as Fig. 2, photographed between crossed nicols 1630-1631
XXVII. Elephas indicus: (Fig. 1) Area within circle on PI. xxvi, Fig. 2. (Fig. 2) Transverse section of about half of a small,
mature tusk about 14 inches long. (Fig. 3) Part of section mthin circle on Fig. 2. (Fig. 4) Same as Fig. 3, with
crossed nicols. (Fig. 5) A small area near outer edge of cement band of section shown in Figs. 2-4. (Fig. 6)
Phiomia wintoni: Transverse thin section about one-half inch from tip of small, worn tusk 1630-1631
XXVIII. Phiomia wintoni: (Fig. 1) Same as PI. xxvii, Fig. 6, with crossed nicols. (Fig. 2) Area within circle on PI. xxvii,
Fig. 6 1630-1631
XXIX. Trilophodon obscurus: (Fig. 1) Transverse section of part of tusk. (Fig. 2) Area within circle on Fig. 1. Crossed
nicols. (Fig. 3) Same tusk as Figs. 1 and 2, thin section across enamel cut in a plane vertical to surface and
parallel to longitudinal axis of tooth. Crossed nicols. (Fig. 4) Trilophodon (Megabelodon) sp. Transverse thin
section of tusk near edge of enamel band. (Fig. 5) A different part of same thin section as Fig. 4. Crossed
nicols 1630-1631
XXX. Trilophodon (Megabelodon) sp.: (Fig. 1) Vertical thin section of broken molar tubercle.
Part of section in (Fig. 2)
circle on Fig. 1. Crossed nicols. (Fig. 3) Transverse thin section of a molar cusp, from same tooth as Figs. 1
and 2 but a different cusp. (Fig. 4) Part of section in inner circle on Fig. 3. (Fig. 5) Part of section in outer
circle on Fig. 3 1630-1631

FIGURE
681. Scene on the ancient Solo River, illustrating the Dietrich-Osborn theory of the Middle Pleistocene age of Pithecanthropus
erectus. Restoration by Flinsch 804
682. Family group of Stegodon orientalis grangeri. Restoration by Flinsch 804
683. Stegodon elephantoides ( = cliftii), cotype molar. After Lydekker 808
684. Stegodon orientalis grangeri, referred first right superior molar 808
685. Structural evolution of the cones, conelets, and ridge-crests in the Stegolophodon phylum, in comparison with Paheomastodon 810
686. Structural evolution of the cones, conelets, and ridge-crests in the Stegodon phylum 811
687. Gradual progressive hypsodontj' in superior grinders of Stegodon 813
688. Gradual progressive hpysodonty in superior and inferior grinders of Stegodon 813
689. Turicius compared with Trilophodon molars. After Mayet 819
690. Turicius and Stegolophodon form of grinding teeth 821
691. Map showing geographic distribution of types and referred specimens of Stegolophodon and Stegodon 823
692. Fossil-bearing horizons along the Irrawaddy River, Burma 824
693. Stegolophodon latidens, lectotype palate. After Clift 826
694. Stegolophodon latidens, cotype third right inferior molar. After Clift 826

XV
xvi OSBORN: THE PROBOSCIDEA

FIGtTRE PAGE
695. Elephas cliftii Falconer and Cautley equals Stegodon elephantoides ( = cliflii), cotype first left superior molar. After Clift . . . 826
696. Stegodon elephantaides, lectotype lower jaw. After Clift 826
697. Stegodon insignis, lectotype and cotype molars. After Falconer and Cautley 829
698. Stegodon ganesa, lectotype third superior molar. After Falconer and Cautley 829
699. Stegodon bombifrons, cotype skull. After Falconer and Cautley 830
700. Elephas cliftii, original type figure of first left superior molar, equals Stegodon elephantoides { = cliftii). After Clift 831
701. Elephas cliftii, new type figure, equals Stegodon elephantoides ( = cliftii). After photograph of type caSt 831
702. Stegodon sinensis, type third superior deciduous premolar. After Owen 831
703. Stegodon orientalis, type. Portion of true molar and posterior end of milk molar. After Owen 832
704. Stegolophodon cautleyi, lectotype third left superior molar. After Lydekker 832
705. Stegodon trigonocephalus, type immature skull. After Martin 833
706. Stegodon (Archidiskodon?) mindanensis, type molar. After Naumann 833
707. Stegodon airmmna, type lower jaw. After Martin 834
708. Stegolophodon stegodontoides, type right third superior molar. After Lydekker 834
709. Stegodon aurorse, type right second superior molar. After Matsumoto 834
710. Stegolophodon snblatidens, type molar. After photograph 835
711. Stegodon pinjorensis, type palate. After photograph 835
712. Stegolophodon cautleyi, lectotype left third superior molar. After Lydekker 841
713. Stegolophodon cautleyi, lectotype left third superior molar. After Falconer and Cautley 841
714. Stegolophodon cautleyi, cotype first superior molar. After Lydekker 842
715. Stegolophodon cautleyi, referred left second superior molar. After Lydekker 842
716. Stegolophodon latidens, lectotype (new figure) right second and third superior molars. Orthogonal projection after cast. . 843
717. Stegolophodo7i latidens, cotype third inferior molar. After Clift 843
718. Stegolophodon latidens, lectotype right second and third superior molars. Perspective. After Clift 844
719. Stegolophodon latidens, lectotype molars. Section after Falconer and Cautley, in comparison with Mastodon americanus,
third superior molar 844
720. Stegolophodon latidens, lectotj^je third superior molar. After Gaudry 844
721. Stegolophodon latidens, referred third right superior molar from Japan. Primitive stage. After Matsumoto 845
722. Stegolophodon sublatidens, type. Posterior half of a third right superior molar. After photograph 846
723. Stegolophodon stegodontoides, type third right superior molar. After Lydekker 847
724. Stegolophodon nathotensis, type fragmentary molars. American Museum (Barnum Brown) collection 848
725. Stegolophodon cautleyi pi-ogressus, type cranium. American Mu.seum (Barnum Brown) collection 849
726. Stegolophodon cautleyi progressus, type right superior dentition. American Museum (Barnum Brown) collection 849
727. Stegolophodon cautleyi progressus, type skull, left lateral, palatal, and right lateral aspects. After photograph 850
728. Stegolophodon lydekkeri, type third left superior molar. After Lydekker 851
729. Falconer's map of the geology of India 852
730. Map of chief Miocene and Pliocene fo.ssil mammal depo.sits of A.sia. After Osborn 853
731. Species of Stegodon from India, China, and Java. Restorations by Flinsch 855
732. Stegodon ganesa, S. ifisignis, and S. bombifrons crania. After plates by Falconer and Cautley 856
733. Stegodon ganesa, cranium with tusk extremities turned inward, in contrast to Falconer and Cautley's restoration (Fig. 732)
with tusks turned outward and closely appre.ssed 857
734. Stegodon bombifrons, lectotype. A generalized cranium. After Falconer and Cautley 858
735. Stegodon insig7ns, a specialized cranium. After Falconer and Cautley 858
736. Stegodon ganesa, a specialized male cranium. Restoration after Falconer and Cautley 858
737. Stegodon sinensis, type third right superior deciduous premolar. After Owen 860
738. Stegodon elephantoides, lectotype second and third left inferior molars. After Clift 861
739. Stegodon elephantoides Clift { = cliftii Falconer). Cotype first left superior molar. After cast 862
740. Stegodon elephantcndes C\iit { = cliftii Falconer). Cotype first left superior molar. After Falconer and Cautley 862
741. Stegodon elephantoides { = cliftii), referred third left inferior molar. After Falconer 863
742. Stegodon bombifro7is, lectotype, cotype, and referred crania. After Falconer and Cautley 864
743. Stegodon bombifrons. Restoration by Flinsch 864
744. Stegodon bombifrons, cotype .skull. After I'\alconer and Cautley 865
745. Stegodon bombifrons, referred third right superior molar. After Falconer and Cautley 866
746. Stegodon bombifrons, referred fragment of third right superior molar. American Museum (Barnum Brown) collection. . . 866
747. Stegodon insignis, lectotype and cotype molars. After Falconer and Cautley 867
748. Stegodon insignis, referred superior and inferior molars. American Museum (Barnum Brown) collection 868
 .

LIST OF ILLUSTRATIONS xvii

FIGURE PAGfJ
749. Slrgodun insignis, referred second right superior molar. American Museum (Barnum Brown) collection 869
750. Sk-godon insignis, referred inferior mandil)lc. American Museum (Barinnn Brown) collection 869
751. Stegodon insignis, referred juvenile and young adult lower jaws. American Museum (Barnum Brown) collection 870
752. Stegodon insignis, referred crania. After Falconer and Cautley 870
753. Stegodon insignis, referred third left inferior molar. American Museum (Barnum Brown) collection 871
754. Stegodon insignis, oft-reproduced referred skull and tusks. After Falconer and Cautley 871
755. Stegodon ganesa. Restoration by Flinsch 872
756. Stegodon insignis. Restoration by Flinsch 872
757. Sectioned molars (lectotypes) of Stegodon ganesa and Stegolophodon latidens. After Falconer and Cautley 874
758. Stegodon insignis binnanicus, type ramus with third left inferior molar. American Museum (Barmim Brown) collection. 875
759. Stegodon orientaUs grangeri, type and referred molars. American Museum (Walter Granger) collection 876
760. Stegodon insignis-ganesa ref., <S. insignis hirmanicus type, and S. orientaUs grangeri type. Comparison of left third inferior
molars 877
761. Stegodon orientaUs grangeri, referred superior and inferior molars 878
762. Stegodon orientaUs grangeri, type and referred superior and inferior molars. After photographs 879
763. Stegodon orientaUs grangeri, infantile, juvenile, young adult, and mature adult crania 880
764. Stegodon pinjorensis, S. insignis, S. airaivana, and S. orientaUs grangeri, sections of third left superior molars 881
765. Stegodon pinjorensis, type skull. American Museum (Barnimi Brown) collection 882
766. Stegodon insignis, referred lower jaw and right superior tusk (American Museum, l^arniun Brown, collection), compared
with S. ganesa, referred skull with tusks, after Falconer and Cautley 882
767. Stegodon pinjorensis, type skull, also same skull superimposed on referred skull of S. ganesa 883
768. Stegodon pinjorensis. Restoration by Flinsch 883
769. Stegodon orientaUs, type. Portion of true molar and posterior end of milk molar. After Owen 884
770. Stegodon insignis{?) = orientaUs{f)
and Serridentinus
ref. lydekkeri type. Molars, after Schlo.sscr 885
771. Stegodon airawana, type lower jaw. After Martin 886
772. Map of Kendeng horizon, Trinil, Java. After Dubois 887
773. Stegodon airawana, referred skull. After Janensch 888
774. Stegodon airehvann. Restoration by Flinsch 889
775. Stegodon trigonocephahis. Restoration by Flinsch 890
776. Stegodon trigonocephahis, type skull, after Martin, and single ridge-crest of molar, doubtfully referred by Naumann to S.
insignis or to <S'. ganesa 890
777. Stegodont crania: Stegodon pinjorensis type, S. bornbifrons cotype, S. orientaUs grangeri ref., S. trigonocephahis type, S.
airdwana ref 891
778. Stegodon (Archidiskodonf) mindnnensis, type inferior molar (incomplete). After Naumann 892
779. Stegodon airdwana, section of referred second molar, and section of ridge-crest of third superior molar. After Janensch. 892
780. Stegodon aiirorse, type second right superior molar. After Matsumoto 893
781. Stegodon aiirora\ type second right superior molar (vertical section). Aft(>r i)hotograph 893
782. Stegodon bondolensis, type mandible with third molars in siti(. After yiin d(>r Maarel 895
783. Stegodon trigonocephahis praecwsor, type third left inferior molar. After von Koenigswald 896
784. Parastegodon? kwantoensis, ty])e lower jaw with right second molar. After Tokunaga 897
785. Stegodon yiishensis, type third left superioi- molar. After Young 897
786. Stegodon offieinaUs, type molar fragment. After Hopwood 898
787. Stegodon oflkinaU's, referred molar fragment. After Hojiwood 898
788. Stegodon zdanskyi, type third right inferior molar. After Hopwood 899
789. Parastegodon [Stegodon?] sugiyamai, type, probably a left second superior molar. After Tokunaga 900
790. Thirteen fossil mammal-bearing formations of Japan. Matsumoto
After 902
791. Theoretic jjhylogeny of the Mastodontidse. After Matsumoto 903
792. Theoretic phylogeny of the Stegodonts. After Matsumoto 904
793. phylogeny of the Elephantidae of Asia and Europe. After Matsiunoto
Theor(>tic 905
794. Elephanti,d£ePrimitive, intermediate, and progressive mandibles and grinding teeth. After Falconer and Cautley
: 910
795. General climatic distribution of the subfamilies of the Elephantoidea and Stegodontoidea, including theoretic migration
lines (1938) 914
796. Asiatic elephant, juvenile cranium. After Osborn and Gregory 916
797. Asiatic elephant, juvenile cranium, also orbitosphenoidal region, left side. After Osborn and Gregory 917
798. Asiatic elephant, infantile cranium, basis cranii 917
799. Asiatic elephant, infantile cranium, occiput and jaws 918
xviii OSBORN: THE PROBOSCIDEA

FIGUKE PAGE
800. Asiatic cipphant {Ehphas indicus hengalen.sis), adult cranium, ])alatal view 918
801. Five wire section lines. Key to sections. Young Elephas indicus bengalensis cranium 919
802. Asiatic elephant, young. Interior view of skull, after Gregory 920
803. Elephas indicus cranium. Fronto-occipital growth curves of vertex 921
804. Elephas indicus cranium. Growth ciu'ves of vertex: midfrontal vertical section 921
805. Preliminary study of fronto-occipital-basilar planes: Elephas indicus, Loxodonta africana (adult skulls), and Parelephas
jeffersonii (young male skull) subsequently referred to Archidiskodon imperator 922
806. Later study of fronto-occipital-niolar-3 planes. Para-occipitofrontal section of Loxodonta africana, Parelephas jeffersonii
male and female (the latter subsequently made the type of Mammonteus primigenius compressus by Osborn), and
Elephas iyidicus 922
807. Elephas indicus, superior nasal growth stages (sections) 923
808. Elephas indicus, transfrontal growth stages (sections) 923
809. Elephas indicus, oceipitohorizontal growth stages (sections) 923
810. Mid-occipitofrontal sections, vertical longitudinal Crania of Loxodonta africana, Elephas indicus, and Parelephas jeffersonii
:

[
= Archidiskodon imperator] 924
811. Frontal .sections: Crania o( Loxodonta africana, Elephas indicus, and Parelephas jeffersonii = Archidiskodon imperator] [ . . . 924
812. Nasal contours (sections) Loxodonta africana and Elephas indicus compared with Parelephas jeffersonii and Archidiskodon
:

imperator 925
813. Midfrontal or intertemporal forehead (Loxodonta africana, Elephas indicus, and Parelephas jeffersonii) 925
814. Oceipitohorizontal sections through liack of occiput (Loxodonta africana, Elephas indicus, Parelephas jeffersonii) 925
815. Map. Successive habitats and world migration routes of the archaic-toothed mammoth Archidiskodon 934
816. Mid-cranial axes (I/jxodontines, Elephantines, Mammontines) 937
817. Archidiskodon and Stegodon, comparative profiles of crania 938
818. Hypsicephalic crania of the Mammontinse (Archidiskodon imperator, Mammonteus primigenius, Parelephas jeffersonii and,

P. xcashingtonii) 939
819. Comparative molars showing evolution of the ridges in the Elephantoidea and Stegodontoidea (Mam-
series of superior
monteus primigenius compressus, Archidiskodon planifrons, Stegodon aurorse, S. ganesa, S. insignis) 939
820. Map of central region of the Siwalik Hills, 200 miles south and north of Simla 940
821. Map of chief Lower andUpper Pleistocene localities of western Eurasia in which occur species of Archidiskodon, Parelephas,
Mammonteus, Loxodonta, and Palseoloxodon. After Osborn 941
822. Map showing geographic distribution of the principal species of Archidiskodonts 942
823. Vaal River gravel terraces, South Africa 945
824. Archidiskodon imperatw maibeni, family group along the Platte River, Nebraska. Restoration by Flinsch 947
825. Archidiskodon planifrons, lectotype right second superior molar, and cotype left third inferior molar. After Falconer and
Cautley 951
826. Map of favorable exposures, southwest of Simla, of the Archidiskodon planifrons life zone, U))iier Siwaliks, India, eliiefiy
Pinjor horizon 952
827. Archidiskodon planifrons, new lectotype molar (Miss Woodward's drawing)
figure of right second sujx'rior 952
828. Archidiskodon planifrons, lectotype right second superior molar. After Lydekker 953
829. Archidiskodon planifrons, referred right third superior molar. After Falconer and Cautley 953
830. Archidiskodon planifrons, ])rofile of skull. Ke])roduced from (laudry after Falconer and C'autley 953
831. Archidiskodon planifrons, referred first left superior molar. American Museum (Barnum Brown) collection 956
832. Archidiskodon planifrons, referred first left and right sujierior molars. American Museum (Barnum Brown) collec-tion. 956
833. Archidiskodon planifrons, referred second left sujjerior molar. American Museum (Barnum Brown) collection 956
834. Archidiskodon planifrons, referred third left superior molar. American Museum (Barnum Brown) collection 956
835. Archidiskodon planifrons, referred left third inferior molar. American Museum (Barnum Brown) collection 956
836. Archidiskodon. planifrons, portion of refei-red fiist and second right inferior molars. American Museum (Barnum Brown)
collection 957
837. Archidiskodon planifrons, ref(>rred fourth inferior deciduous premolar. American Museum (Barnum lirown) collection. . 957
838. Archidiskodon planifrons, fourth left inferior deciduous premolar and first left inferior molar. American Museum (liarnum
Brown) collection 957
839. Archidiskodon planifrons, portion of lower jaw with I'iglit third molar in situ. American Mu.seum (Barnum Blown) col-

lection 957
840. A rchidi.><kodon planifrons, port'iou of lower jaw with right third molar in situ. .Vmerican Museum (Barnum Brown)
collection 957
841. Archidiskodon planifrons, referred left third inferior molar. American Museum (Barnum Brown) collection 958
 .

LIST OF ILLUSTRATIONS xix

FIGURE l'A(;i';

842. AirhifliskodoN plunifrons, roforrod tliird liglil American Mvisoum (liarniiin Urown) eolleetion
inferior molar. 958
843. Archuliskodoii phinifrons, referred third left inferior molar. American Miuscum (Barnum Brown) collection 958
844. Archidiskodon planifrons, portion of referred lower jaw with left third molar in situ. American Museum (Bariumi Brown)
collection g.^g
845. Archidiskodon planifro?is, referred right third inferior molar. American Museum (Barnum Brown) collection 959
846. Archidiskodon planifrons, referred left third inferior molar, also transverse vertical section. American Museum (Barnum
Brown) collection 959
847. Leith-Adamsia sixmUkiensis, type third right superior molars = synonym of Archidiskodon planifrons].
[ After Falconer
and Cautley qqq
848. Archidiskodon planifrons, referred cranium (supposed female). Redrawn after Falconer and Cautley 960
849. Archidiskodon planifrons, three referred primitive mandibles from the Siwaliks, India, Chagny-Belleeroix and Seneze,
France. After Mayet and Roman 962
850. Archidiskodon planifrons of C'hagny, referred tusks, maxillae, condyle, and atlas. After Mayet and Roman 963
851. Map of type locality of Archidiskodon proplanifrons, and type and referred localities of .4. planifrons; also theoretic range
from supposed African center northward to France and Britain and eastward to India 964
852. Archidiskodon playiifrons of Piltdown, England. Molar fragments, after Smith Woodward 965
853. Archidiskodon planifrons, molar fragments from Piltdown, England. Redrawn by Miss G. M. Woodward 966
854. Archidiskodon planifrons, lectotype sixth, seventh, and eighth superior ridge-plates. After dra\ving by Miss G. M. Wood-
ward 9(56
855. Archidiskodon planifrons of Piltdown, England (sectioned molar), in compari.son with lectotype and referred molar sections 967
856. Scene on the ancient river Ouse, illustrating the Osborn theory of the Upper Pliocene age of Eoanthropus dawsoni. Restora-
tion l)y Flinsch 968
857. Archidiskodon planifrons rumaniis, type third left inferior molar and portion of referred third right inferior molar. After
Stefanescu 969
858. Archidiskodon mei-idionalis, lectotype cranium (C). After Nesti 970
859. Archidiskodon meridionalis, cotype cranium (A). After Nesti 971
860. Archidiskodon meridionalis, restored cotype cranium (A). After Weithofer 971
861. Archidiskodon meridionalis, lectotype cranium (C). After Weithofer 972
862. Archidiskodon meridionalis, referred third inferior molars from the Val d'Arno and Norwich Crag. After Falconer and
Cautley 973
863. Archidiskodon meridionalis, referred third superior molar from Chagny, France. After Gaudry 974
864. Elephas lyrodon [
= Archidiskodon meridionalis, female], type skull. After Weithofer 975
865. Crania (19) of the Mammontinse {Mammonteus primigeniiis, Parelephas trogontherii Archidiskodon meridionalis, A. impera-
,

tor, and A. planifrons). After Pohlig, Falconer, Weithofer 976

866. Archidiskodon meridionalis of Durfort, skeleton, largely restored. After photograph 978
867. Archidiskodon meridionalis of Durfort. Restoration by Flinsch 979
868. Archidiskodon meridionalis of Durfort. Skeleton redrawn after jjhotographic plate in Gaudry 979
869. Archidiskodon meridionalis, referred third left (?) superior molar from Essex, England. After Lydekker 980
870. Archidiskodon meridionalis crotnerensis, type third left superior molar. After Deperet and Mayet 980
871. Archidiskodon, Parelephas, Mammonteus: Primiti^•e grinding teeth from the Forest Bed and Red Crag, England. After
pencil sketches by the present author 981
872. Map showing fossil Proboscidea route from southern equatorial Africa 983
873. Archidiskodon proplanifrons, type right third superior molar. After original 987
874. Archidiskodon subplanifrons, Osborn's original type figure of third right inferior molar 987
875. Archidiskodon subplanifrons, type right third inferior molar (new figure) 988
876. Archidiskodon planifrons, referred third right inferior molar 988
877. Archidiskodon broomi, Osborn's original type figure of third right inferior molar 989
878. Archidiskodon vanalpheni, type third left superior molar. After Dart 990
879. Archidiskodon milletti, type third left superior molar. After Dart 991
880. Archidiskodon lo.rodonloides, type third left superior molar. After Dart 992
881. Archidiskodon yorki, type molar fragment. After Dart 993
882. Metarchidiskodon griqua, type third left superior molar. ( )riginal figure of Haughton and new figure after Osborn 995
883. Metarchidiskodon griqna, referred fragmentary molar, from Kaiso Bone-beds, Africa. After photograph 995
884. Archidiskodon imperalor, tyi:)C fragment of a third right superior molar. Leidy's original typo figure 999
885. Archidiskodon imperator, type third right superior molar. After Osborn 999
886. A rchidiskodon imperator, Leidy's type molar and Osborn's neotype combined. After Osborn 999
XX OSBORN: THE PROBOSCIDEA

FIGUHE PAGE
887. Archidiskoclon imperator, type molar compared with type molar of Parelephas rohinihi. After i)liotograpli 1000
888. Archidiskodon imperator, ncotypc third right siip(>rior molar. After Osborn 1000
889. Archidiskodon imperator, referred third superior and inferior molars of two individuals, showing mechanical reversal of the
convex and concave surfaces. After O.sborn 1001
890. Map showing distribution of Archidiskodon imperator west of the Mississippi River. After Hay 1003
891. Archidiskodon imperator, referred cranium of young male from Texas. American Museum Cope Collection 1004
892. Archidiskodon imperator, referred mandible, in comparison with mandibles of Parelephas jejfersonii 1006
893. Mandibles of Elephas indicus, Loxodonta africana, Parelephas irashingtonii, and Archidiskodon hayi 1006
894. Archidiskodon imperator, right superior tusk of record size, from Post, Texas, combined with a superb tusk formerly in the

National Museum of Mexico, now destroyed. After photograph 1007

895. Archidiskodon imperator, referred young male(?) skull from tar pools of Rancho La Brea, C'alifornia 1007
896. Archidiskodon imperator, referred skull from Victoria, Texas 1008
897. Archidiskodon imperator, referred skull from Victoria, Texas, as mounted in the American Museum 1009
898. Archidiskodon imperator, referred jaw from Tule Canon, Texas 1010
899. Archidiskodon imperator, crushed skull of aged male (Nebraska Museum) from Hay Springs, Nebraska 1011
900. Archidiskodon imperator, crushed skull of adult male in the American Museum, from Hay Springs, Nebraska 1011
901. Archidiskodon imperator, referred third right superior molar, from Zumpango, Mexico. After photograph 1013
902. Archidiskodon imperator, referred mature male cranium, from Tepexpan, Mexico. After photograph 1014
903. Archidiskodon hayi (?), referred mandible in Ceological Institute of City of Mexico. After jihotograph 1014
904. Archidiskodon imperator silvestris, type third left superior molar. After Freudenbcrg 1015
905. Archidiskodon imperator fakoneri cotype jaw after Freudenberg. Originally figured by \'illada
,
1016
906. Archidiskodon imperator, referred. Aged male. Reconstruction 1017
907. Archidiskodon imperator, referred right forelimb, as mounted in theAmerican Museum 1018
908. Archidiskodon imperator (young adult) and Loxodonta africana oxyotis ("Jumbo"), referred limb bones 1018
909. Archidiskodon imperator. After restoration by Osborn and Knight, 1908 1019
910. Archidiskodon imperator maibeni, forclimbs of type skeleton, also cranium and tusks of Parelephas jeffersonii 1020
911. Archidiskodon imperator maibeni, mounted type skeleton in Morrill Hall, University of Nebraska 1021
912. Shoulder heights of living and extinct elephants (Elephas indicus, Loxodonta africana oxyotis, Archidiskodon imperator, A.
imperator maibeni) 1022
913. Archidiskodon hayi, tyi)e jaw. After Barbour 1024
914. Archidiskodon planifrons, primitive mandibles from the Siwaliks, India, Chagny-Bellecroix and Seneze, France. After
Mayet and Roman 1024
915. Archidiskodon hayi, type mandible, compared with A. imperator ref 1025
916. Archidiskodon imperator, referred juvenile jaw, subsequently made by Barbour the type of Elephas [Archidiskodon] scotli.

After i)h()tograi)h 11*26

917. ComiJarisonof 1yp<> mandihlcs oi Archidiskodon imperator maibeni and A. imperator scotli; also enlarged views of right .second
inferior molar of the type of A. scotti 1*'27

918. Archidiskodon imperator maibeni, type superior and inferior dentition. After photograph 1028
919. Archidiskodon haroldcooki, type mandible with third right molar in situ. After Hay and Cook 1029
920. Archidiskodon eiilis, type, compared with A. imperator ref. After photograph 1030
921. Archidiskodon exilis, type. Facial portion of skull, with tusks and lower jaw. Restoration. After photograph 1031
922. Map showing location of some occurrences of fossil ele])hants on Channel Islands. After Stock 1032
923. Archidi.skodon .sonoriensis, anterior portion of type mandible and maxilla, showing third sup(>rior and inferior molars. 1033
. . .

924. Archidiskodon meridionalis nebrasce7wis, type mandible. After Osborn 1034

925. Archidiskodon meridionalis of Durfort, referred sujierior and inferior molars, found associated with sk(>leton. After casts. .
. 1035
926. Archidiskodon meridionalis of Durfort, second and third su])erior and inferior molars of skeleton. After casts 1035
927. Archidiskodon meridionalis nebra.scensis, type. Restoration by Flinseh 1036
928. Archidiskodon meridionalis of Durfort and A. meridionalis nebrascensis of Nebraska. Restorations by Flinseh 1037

929. Parelephas trogontherii of Mosbach. Restoration by Flinseh 1038

930. Parelephas jeffersonii, type. Restoration by Knight 1040
931. Parelephas jeffersonii, type skeleton. Second figure after Osborn 1041
932. Map of chief Lower to Upper Pleistocene localities in which occur species of Archidiskodon, I'urclcphas, Mammonteus,
Loxodonta, and Pala-oloxodon. After Osborn 1042
933. Map .showing geographic distribution of si)ecies of Parelephas, types and referred specimens 1047
934. Mammonteus primigenius and Parelephas trogontherii, cranial profiles. After Polilig and l''aleoner 1050
935. Left lateral profiles of seven species of Parelephas with progressive ridge formulae 1051
 . .

LIST OF ILLUSTRATIONS xxi

FIGURE PAGE
936. ParelepJms of Eiiropo and America in comparison with Elephas indicus bengnlensis. Restorations by Flinsch 1052
937. Cranial ]3rofiles oi Airhidiskodon imperator, Mammonteus pnmigenius, Parelephasjeffersonii, and /*. washinglonii 1053
938. Parelephas trogontherioides, lectotype and cotype molars. After Zuffardi 1054
939. Parelephas trogonlherii, type third superior and inferior molais. After Pohlig 1057
940. Parelephas trogonlherii, referred molars from Siissenborn and Weimar. After Wiist 1058
94 1 Elephas antiquus Neslii Pohlig = Parelephas{?) trogontherii nestii], cotype or syntype
[ left third superior and inferior molars.
After photographs 1059
942. Parelephas armeniacus, type left third superior molar. After Falconer 1061
943. Parelephas intermedius, referred molars. After photographs 1063
944. Parelephas intermedius, restored skeleton in Lyons Museum. After Lortct and Chantre 1064
945. Parelephas cotype molars, from Tiraspol, Russia. After Pavlow
wiisti, 1066
946. Parelephas jacksoni, type juvenile jaw, and referred Elephas [Mammonteus] primigenius jaw. After Mather 1068
947. Diagrammatic cross-.scction of type locality of Parelephas jacksoni. Aitcv Mather 1069
948. Parelephas rohnnbi, type third right lower molar (middle portion) longitudinally and vertically bisected. After Falconer. . . 1071
949. Parelephas columbi, restored type molar, redrawn for present Memoir 1072
950. Elephas lexiaiius [
= Parelephas cohimhi ref.], type right third inferior molar. After Blake 1073
951. Parelephas columbi, Falconer's type third right inferior molar and Osborn's neotyi)e third left inferior molar. After Usborn 1074
952. Parelephas columbi, key to superior and inferior grinding teeth selected from the Cohen Collection, phosphate beds, Charles-
ton, S. C 1076
953. Map showing distribution of Parelephas jejfersonii, Mammonteus primigenius, Parelephas columbi, and Archidiskodon
imperator in the United States and Canada. After Hay 1078
954. Parelephas columbi, superior and inferior molars found in incomplete skull of Amherst skeleton 1079
955. Parelephas columbi. Amherst skeleton 1081
956. Parelephas columbi felicis, type third right superior molar. After Freudenberg 1082
957. Parelephas columbi cayennensis, type fragmentary third right superior molar. After cast 1083
958. Parelephas jeffersonii, showing ridge-plate compression at three levels of third right inferior molar 1085
959. Parelephas jeffersonii, aged type third superior and inferior molars; also same superposed on type molars of Elephas
rooseveUi 1086
960. Parelephas jeffersonii, type and paratype [ideotype] grinding teeth and jaws 1089
961. Frontal views of crania of Parelephas jeffersonii and P. loashingtonii 1090
962. Profile views of type and referred crania of Parelephas jeffersonii, P. washingtonii, and Mammonteus primigenius; also
front view of M. primigenius 1091
963. Parelephas jeffersonii (Franklin County Mammoth) and Elephas indicus bengalensis crania 1092
964. Parelephas jeffersonii (Franklin County Mammoth) skull in Nebraska State Museum. After photograph 1093
965. Parelephas jeffersonii (Franklin County Mammoth), aged. Diagram showing ridge-plates of second and third superior
molars, and portion of crown of third inferior molar. After Barbour 1093
966. Parelephas jeffersonii, type skeleton in American Mu.seum. First published type figure by Osborn 1094
967. Parelephas jeffersonii, paratype [ideotype] jaw 1096
968. Elephas rooseveUi (syn. Parelephasjeffersonii), type third superior and inferior molars 1096
969. Parelephas progressus, type third superior and inferior molars from Zanes\-ille, Ohio, side \-iews, originally figured li.y

Osborn as paratypes of Elephas jeffersonii. After Osborn 1098

970. Parelephas progressus, type molars, crown views. After Osborn 1098
971. Parelephas washingtonii, referred cranium 1100
972. Parelephas washingtonii, type adult jaw 1 101
973. Parelephas washingtonii, referred young adult male skull, right lateral aspect 1102
974. Parelephas washingtonii, referred young adult male skull, left lateral aspect, combined with type jaw 1102
975. Parelephas washingtonii, type jaw compared with that of P. jeffersonii 1103
976. Parelephas xcashingtonii, referred cranium with second and third superior grinders in situ 1103
977. Parelephas ivashingtonii, referred second left inferior molar. After Peterson 1104
978. Parelephasil) type skull fragment. After Hay
eellsi, 1104
979. Parelephas jloridanus, type and paratype crania and jaws compared with type molar fragment of P. columbi cayennensis .... 1 105
980. Parelephas floridanus, type and paratype palates, with molars in situ 1106
981. Parelephas floridanus, type right and left superior and inferior molars. After Osborn 1108
982. Parelephas floridanus, type third left superior molar (detailed photograph) and drawing showing method of measuring length
of superior molar crown. After Osborn 1 109
983. Parelephas floridanus, type mandible. After Osborn 1110
xxii OSBORN: THE PROBOSCIDEA

FIGURE PAGE
984. Porelephas flon'danus, paiatype mandible. After O.sborii 1110
985. Parelephas floridanus, type inferior molar.s. After Osboni 1110
986. Parelephas flon'danus, reconstructed type cranium. After Osborn 1111
987. Parelephas floridanus, referred inferior mandible and milk dentition 1111
988. New .standard method (1930) of i)robo.scidean .skeletal mea.surcment, illustrated on type skeleton of Parelephas jeffersonii.
After Osborn 1112
989. Parelephas floridanus, referred right third inferior molar, showing method of ridge-plate mea.surement 1115
990. Maminonleus primigenius (Woolly Mammoth). After painting by Knight 1116
991. Messerschmidt cranium of the mammoth of Siberia. After Breyne 1119
992. Comparison of crania of Matnmonteus primigenius, Elephas indicus, and Loxodonta nfricana. After Cuvier 1121
993. Mannnonteus primigenius, lectotype molars. After casts 1122
994. Migrating Woolly Mammoth (Mammontevs primigenius) as it appeared on the river Somme, northern France. Restoration
by Knight 1126
995. ( omjiarison of the tip of the triuik of the mammoth {Mammonteus primigenius) with that of Elephas indicus and Loxodonta

afrirana. After Flerov 1128

996. Manimonteus primigenius, skeleton, from Kolyma-Beresowka River, Siberia. After Salensky 1130
997. Mummonkus primigenius Jraasi, skeleton, from Stcinheim on the Murr, Wurttemlx'rg. After Abel 1130
998. Mammo7iteus primigenius, skeleton, from Borna, Germany. After Abel 1130
999. Munimonieus primigenius, skeleton, fnmi Lierre, Belgium. After Dupont 1130
1000. Restoration of the Woolly Mammoth sketched on the wall of the cavern of Les C'ombarelles aux Eyzies (Dordogne),
France. After Capitan, Breuil, and Peyrony 1131
Outlines of the W^jolly Mammoth from the Oiotto of Oombarelles 1131
Charging mammoth incised on a tusk of Mammonteus primigenius discovered in the rock shelter of La Madeleine (Dor-
dogne), France. After Lartet and Chiisty 1132
Maji showing geographic distribution of Mammonteus and Parelephas in North America. After Hay 1133
Map showing location and principal disco^eIies of fossil manmialian fauna of Alaska-Yukon to the year 1929 1134
Diagram .showing geographic range of Mammonteus and Parelephas. Now superseded by figiu-e 795 above 1135
Map sliowing geographic distribution of principal species of Mammonteus, types and referred 1136
Mammonteus primigenius, skeleton from Moravia. After photograph 1139
Mannnonteus primigenius, referred molars of Alaska 1 142
Mammonteus pri7nigenius and Parelephas jeffersonii and P. irashingtonii crania compared 1144
Mammonteus primigenius, male cranium, from the Yukon. After photograph 1145
Mammonteus primigenius fiom Alaska, showing growth stages in the jaws and teeth 1145
Type of Elephas odontoti/raiuius = third superior molar of Mammonteus primigenius]. After Eichwald
[
1146
.Mammonteus primigenius, Elephas indicus bengalensis, and Loxodonta africana oxyotis. Restorations by Flinsch 1147
Mammonteus primigenius ("Adams skeleton") from the Lena River, Siberia. Subsequently made by Brandt the ty]x> of
Elephas brachyramphus. After Tile.sius 1148
Mammonteus primigenius leith-adamsi, type third left inferior molar. After Pohlig 1150
Mammonteus hydruntinus, type first left superior molar. After Botti 1 151
Mammonteus primigenius fraasi, type cranium. After Dietrich 1152
.Mammonteus primigenius fraasi, type mounted skeleton. After photograph 1153
Mammonteus primigenius asten»is, type and paratype molars. After Dep^ret and Mayet 1154
Mammonteus primigenius ('^)astensis, reierred molars, compared with (f) Parelephas, Archidiskodon or Hesperolo.rodon
molars. After direct scale tracings by the present author 1155
1021. Mammonteus primigenius americanus, type. Portion of upper molar from near Rochester, New York, .\fter DeKay 1156
1022. .Mamnio)iteus primigenius compressus, type second and tliird superior molars, .•\fter Osborn 1157
1023. .Mammonteus primigenius compressus, type female skull. After Osborn 1158
1024. Mammonteus primigenius compressus, paratype third right superior molar. XHcr ( )sh()rn 1159
1025. Mammonteus primigenius alaskensis, cotype crania, superposed outlines 1159
1026. .Mammonteus primigenius alaskensis, cotype crania 1160
1027. Discoveiy .sites of frozen carcasses of tlie Woolly JVLammoth and Rhinoceros. After Tolmachoff 1162
1028. Map .showing geographic distribution of mammoths in Upper Pliocene and Pleistocene times. After Osborn 1164
1029. .lefi'ersonian Mammoth of Indiana 1 165
1030. Imperial ]\Iammoth of Nebraska 1 165
1031. Arched tu.sks of the African elephant 1166
1032. Mammonteus primigenius, referred skull and half-grown tusks of male specimen found on the Yukon River, Alaska 1166
 .

LIST OF ILLUSTRATIONS xxiii

FIGURE PAGE
1033. Circular tusks of the Woolly Mammoth of Siberia 1 166
1034. Woolly Mammoth, Somme Kiver, France 1166
1035. Strata of the Pekarna Cave, Moravia. After photograph 1168
1036. Mammoth pit of V&tonice, Moravia. After photograph 1168
1037. Ciiant killing stone of the Moravian hunters. After photograph 1168
1038. Ivory figurine of a woman's head, from Brassempouy. After Pilloy in Piette's "L'Art pendant I'Age du Renne" 1168
1039. Equine ivory statuette from Lourdes. After Pilloy in Piette's "L'Art pendant I'Age du Renne" 1169
1040. Loiodonta africana, young adult bull of the Lake Paradise region, east central Africa. Photograph by Mr. and Mrs. Martin
Johnson, and shown in film "Simba." Courtesy of Mr. Daniel E. Pomeroy 1170
1041. Crania of Loxodonta africana, Palxoloxodon namadicus, and Hesperoloxodon antiquus. After Falconer and Cautley, Pilgrim,
Weithofer, and Pohlig 1 172
1042. Cuvier's figures and definitions of Elephas primigenius (Messerschmidt's cranium), E. indicus, and E. africanus 1173
1043. Loxodonta africana and Elephas indicits, third inferior molars. After Owen 1175
1044. Loxodonta africana and Elephas indicus, second .superior molars. After Lydekker 1175
1045. Loxodonta africana oxyotis ("Jumbo"), cranium and jaws 1 176
1046. Palseoloxodon namadicus, tyjie cranium, from the Nerbudda. After Falconer and Cautley 1176
1047. Pala'oloxodon and Hesperoloxodon as depicted by the cave men of North Africa and Spain. After Pomel and after Breuil . . . 1184
1048. Map showing geographic distribution of the principal species of the Loxodontinse 1186
1049. Habitat of the African elephant (Loxodonta), forest and savanna of the Uasin (Jishu Plateau, Kenya ( 'olony. ( 'alf of old fe-
male charging elephant. After photograph by Carl E. Akeley 1189
1050. Habitat of the African elephant (Loxodonta). Same region as previous figure. Females and young bulls in forest. After
photograph by Kermit Roosevelt 1 189
1051. Small herd of African elephants (Loxodonta). After film photograph by Martin Johnson 1189
1052. Akeley group of African elephants in the American Museum 1190
1053. "Khartum" (Loxodonta africana oxyotis), formerly living in the New York Zoological Park. Two growth stages. After
photograph 1 194
1054. Remarkable cave paintings of white rhinoceros and African elephant, discovered in South Africa. Courtesy of London
Illustrated News 1 194
1055. Map showing distribution of existing African elephant (Loxodonta) 1195
1056. Loxodonta africana oxyotis and L. africana pumilio. Restorations by Flinsch 1196
1057. Loxodonta africana, Blumenbach's original figure of type, a right second inferior molar, described as Elephas africanus 1 197
1058. Elephas priscus = Loxodonta africana]. Type molar after Goldfuss
[ 1197
1059. Loxodonta africana peeli, referred skull and tusks of adult male, from Mt. Kenya. After photograph 1198
1060. Loxodonta africana oxyotis ("Jumbo"), referred middle-aged skull (24 years) 1199
1061. Loxodonta africana oxyotis ("Jumbo"), superior and palatal views of cranium. Age twenty-four years 1200
1062. Comparison of tusks of Loxodonta africana oxyotis and Mammonteus primigenius. After photographs 1201
1063. Loxodonta africana albertensis and L. africana peeli, full-grown heads (male and female) 1202
1064. Loxodonta africana (?)cottoni, foetal cranium, jaw, and milk dentition. After Eales 1203
1065. Tusks of African elephant, believed to be the heaviest in the world, shown in front of a typical Arab door at Zanzibar. One
tu.sk now in British Mu.seum. After Kunz, courte.sy of executors of his estate 1204
1066. Loxodonta cornaliae, type right superior molar. After Aradas 1205
1067. Young Addobush elephant (Loxodonta) from Cape Colony. After photograph 1205
1068. Hesperoloxodon of Europe, Palieoloxodon of India and of the Mediterranean Islands, compared with drawing of Hesper-
oloxodon by cave men of northern Spain. Restorations by Flinsch 1206
1069. Comparison of crania of Palxoloxodon namadicus, Hesperoloxodon antiquus italicus, H. antiquus ausonius, and H. antiquus
platyrhynchus. After Falconer and Cautley, Pilgrim, Weithofer, Graells, and Pohlig 1208
1070. Palieoloxodon namadicus, type female(?) cranium. After Falconer and Cautley 1211
1071. Channel of the Godavari near Nandiir Madm^shwar, India. After Pilgrim 1213
1072. Comparison of Hesperoloxodon and Palseoloxodon (syn. Sivalikia) superior and inferior molars 1214
1073. Three progressive broadening stages in the Palseoloxodon (syn. Sivalikia) and Hesperoloxodon superior grinding teeth. . . 1215
1074. Hesperoloxodon antiquus (Upnor elephant). Restoration by Flinsch 1216
1075. Hesperoloxodon a?iiiquus, lectotype left second inferior molar. After Falconer and Cautley 1218
1076. Hesperoloxodon antiquus, referred left second inferior molar from firay's Thiurock. After Falconer and Cautley 1219
1077. Hesperoloxodon antiquus, referred third right superior molar. After Falconer and Cautley 1220
1078. Hesperoloxodon antiquus (Upnor elephant). Referred second superior and inferior molars. After photographs 1220
1079. Hesperoloxodon antiquus of Upnor. Mounted skeleton in British Museum. After photographs 1223
xxiv OSBORN: THE PROBOSCIDEA

FIGURE PAGE
1080. Hesperoloxodon antiquus of Upnor, skeleton in the British Musoum. Redrawn to show original and restored parts 1224
1081. Scapulae of Hesperoloxndon a7itiquus of Upnor, Loxodonla africana ("Jumbo") of the Sudan, and Elephas indicus. After
Andrews and Cooper 1225
Dorsolumbar vertebrae of Hesperoloxodon antiquus of Upnor and Elephas indicus 1225
Upnor elephant, partly restored skeleton and flesh outlines 1227
\'ertebral columns of Hesperoloxodon antiquus, Loxodonta africana oxyotis, Elephas indicus, Mammonteus primigenius, and
Parelephas jeffersonii 1229
Hesperoloxodon antiquus nanus, type, possibly a left second superior molar. After Acconci 1231
Hesperoloxodon antiquus plalyrhynchus, type premaxillaiy rostrum with tusk and portion of right superior maxillary with
second molar in situ. After Graells 1231
Hesperoloxodon antiquus ausonius, type third right and left inferior molars. After Dep^ret and Mayet 1232
Hesperoloxodon antiquus, H. antiquus germanicus, H. antiquus italicus, progressive stages in evolution of grinding teeth 1234
Hesperoloxodon antiquus germanicus, type. Portion of right second inferior molar. After Stefanescu 1235
Hesperoloxodon antiquus germanicus, right tusk in Field Museum of Natural History, from Steinheim, male and female
tusks in Ciotha Museum, from Tonna, right male tusk in Stuttgart Museum, from Steinheim 1236
Hesperoloxodon antiquus germanicus, right tusk excavated at Steinheim. Diagrammatic sketch reproduced through courtesy
of Dr. Henry Field 1237
Hesperoloxodon antiquus italicus. Restoration by Flinsch 1238
Loxodonta africana albertensis. Restoration by Flinsch 1239
Map of valley of the Liri, Italy, showing the location where Hesperoloxodon antiquus italicus was found, and other exposures.
After De
Lorenzo and D'Erasmo 1 239
Mammalian fossils associated with type cranium of Hesperoloxodon antiquus italicus. After Osborn 1240
Hesperoloxodon antiquus italicus of Pignataro Interamna, type cranium before removal 1241

:Map showing location of Pignataro Interamna, region of valley of the Liri. After Century Atlas, 1913 1241

Hesperoloxodon antiquus italicus, type cranium. After De Lorenzo's original sketch and measurements 1242
Hesperoloxodon antiquus italicus, type cranium. After De Lorenzo and D'Erasmo 1243
Hesperoloxodon antiquus italicus, front view of cranium. After De Lorenzo and D'Erasmo 1243
Hesperoloxodon atdiquus italicus, type .superior grinders. After photograph 1244
Hesperoloxodon antiquus italicus, type right second and third superior and inferior grinders. After Osborn 1244
Hesperoloxodon antiquus italicus, type mandible, right lateral view with second and third superior teeth superposed on cor-
responding inferior teeth, and superior view. After Osborn 1245
Hesperoloxodon antiquus italicus, type third right inferior molar. After Osborn 1245
Hesperoloxodon antiquus italicus, type cranium, three aspects 1246
Hesperoloxodon antiquus italicus, type cranium as reconstructed and mounted American Museum. After Osborn .... 1247
in the

Hesperoloxodon antiquus italicus type and Loxodonta africana peeli ref. crania. After Osborn 1248
Comparative bathycephaly of the Loxodontinae: Loxodonta africana, Palxoloxodon namadicus, and Hesperoloxodon
antiquus italicus. After Osborn 1249
Comparison of scapulae of Hesperoloxodon antiquus italicus, Loxodonta africana, and Elephas indicus 1249
Palxoloxodon namadicus, referred cranium, of the Godavari Alluvium at Nandiir Madm^shwar, India.
After Pilgrim 1250
Hesperoloxodon antiquus italicus, referred femur, with Dr. Pohlig standing beside it. After photograph 1251

Intracranial brain casts of Hesperoloxodon antiquus italicus, Loxodonta africana, and Elephas indicus. After Osborn 1252

W. Bauer Quarry at Steinheim on the Murr, showing the site of the 1928 discovery of the cranium of Hesperoloxodon
antiquus germanicus, ref. After photograph 1253
1114. Hesperoloxodon antiquus germanicus, referred crania of 1926 and 1928. After photographs 1254

1115. Hesperoloxodcm antiquus germanicus, referred third right superior molar from Steinheim 1255
1116. Elephas antiquitatis Kruger = Hesperoloxodon antiquus germanicus ref.], type molar. After Breislak
[
1256
1117. Bathymetric map of the Mediterranean Islands. By permission of Longmans, Green and Company, from map edited by
Chisholm and Leete ^^^'

1118. Dwarfed elephants of the Mediterranean Islands. Diagrammatic representation 1258

1119. Dwarfed elephants of the Mediterranean Islands. Restorations by Flinsch 1259
1 120. Loxodonta africana pumilio, young "pygmy" elephant pa.s.sing beneath adult Elephas indicus, female. After photograph ... 1259
1121. Pala-oloxodon mnaidriensis, referred, the "Elephas {antiquu.'^) Melita'" of Pohlig. Fully adult cranium from the (Jrotta di
Pontale, Carini, Sicily. After Pohlig 1260 •

1122. Palxoloxodon mnaidriensis, referred juvenile cranium, the "Elephas (antiquus) Melitie" of Pohlig. After Pohlig 1200
1123. Pala-oloxodon rnelitensis, type third left superior molar. After Falconer 1262
1124. Palwoloxodon melitensis, referred mandible from the Grotta di Pontale, Sicily 1263
 LIST OF ILLUSTRATIONS xxv

FIGURE PAGE
1125. Palseoloxodon 7nnmdriensis, third right inferior molar. After Leith Adams 1264
1126. Palseoloxodon mnaidriensts, type and para type molars. After Leith Adam.s 1264
1127. Denizens of ancient Malta. Restoration by Leith Adams as dwarfed African elephants 1265
1128. Palseoloxodon Cypriotes, cotype molars. After Bate 1266
1129. Palseoloxodon creticus, cotype molars. After Bate 1267
1130. Section of Grotto of Luparello, Palermo, Sicily. After \'aufrey 1268
11.31. Tusks in the three dwarfed species of the Mediterranean Islands: Palseoloxodon fakoneri, P. melilensis, P. mnaidriensis.
After Vaiifrey 1271
1132. Two types of molars belonging to Elephas [
= Palseoloxodon] mnaidriensis according to Vaiifrey, namely, 'tjqpe endioganai'
and 'type pachyganal,' from Shantiun and Puntali. After Vaufrey 1271
1133. Molars referred by ^aufrey to Elephas = Palieoloxodon] melitensis, from Luparello,
[ Sicily, and Benghi.sa, Malta. After
Vaufrey 1272
1134. Ulnae of Palseoloxodon mnaidriensis, P. melitensis, and P. fakoneri. After Vaufrey 1272
1135. Palseoloxodon allanticus, cotype right second inferior molar. After Pomel 1274
1136. Palseoloxodon atlaniicus, referred third left superior molar. After Pomel 1274
1137. Palseoloxodon jolensis, type left third inferior molar. After Pomel 1275
1138. Palseoloxodon recki, lectotype left second inferior molar. After Dietrich 1276
1139. Palseoloxodon{1) andrewsi (Dart's tjrpe of Archidiskodon andrewsi). Left third inferior molar, restored. After O.sborn .... 1278
1140. Palseoloxodon hanekomi, type ?third right superior molar. After Dart 1279
1141. Palseoloxodon yorki, type right ?third inferior molar. After Dart 1280
1142. Palseoloxodon vnlmani, type ?third inferior molar. After Dart 1281
1143. Palseoloxodon kuhni, type "?lower left molar." After Dart 1281
1144. Palseoloxodon archidiskodontoides, type ?second superior molar and part of right humerus. After Haughton 1282
1145. Palseoloxodon transvaalensis, type right third superior molar. Modified after Dart's photographs 1284
1146. Palseoloxodon sheppardi, type left third superior molar. Modified after Dart's photographs 1284
1147. Loxodonta zulu, type third left inferior molar. After Scott 1286
1148. Loxodonta zulu, referred third left inferior molar, from Kaiso Bone-beds, near Lake Albert, Africa. After photograph 1287
1149. Loxodonta prima, type third left inferior molar. After Dart 1287
1150. Loxodonta africana var. obliqua, type third right inferior molar. After Dart 1288
1151. Loxodonta subantiqua, type "possibly a right lower molar, probably the .second." After Haughton 1288
1152. Palseoloxodon najnadicus nanmanni, type, of Makiyama, in comparison with Hesperoloxodon antiquus germanicus ref. of
Pohlig. Diagrammatic outline sketch 1294
1153. Palseoloxodon namadicus namadi, type right third superior molar. After Makiyama 1296
1154. Palseoloxodon protomammonteus, type left third infeiior molar. After Matsumoto 1297
1155. Palseoloxodon protomammonteus proximus, type fragment of loft third inferior molar. After Mat.sumoto 1298
1156. Pals'oloxodon namadicus yabei, type right ramus of mandible, containing third molar. After Matsumoto 1299
1157. Palseoloxodon (Archidiskodonf) tokunagai mut. junior, type right second inferior molar. After Mat.sumoto 1300
1158. Parelephas protomammonteus matsumotoi, type. Portion of left mandibular ramus with third molar //( situ. After Saheki.
Not determined by the present author 1300
1159. Palieoloxodon yokohamanus, type second right superior molar. After Tokunaga 1301
1160. Palseoloxodon hysudrindiciis, cotype molars. After photographs 1302
1161. Map showing Japan as part of the Asiatic continent in Plio-Pleistocene time. After Longmans' New School Atlas 1304
1162. Map showing Japan as part of the Asiatic continent in Plio-Pleistocenc time. After Yabe 1305
1163. Referred Elephas indicus. Male and female Ceylon elephants. After photograph by Plate Ltd 1306
1164. First definition of the genus Elephas, bracketed with ?Rhinoceros. After first edition of Linnaeus' "Sy.stema Naturs,"
page 10 1309
1165. Facsimile of portion of page 11 of Linnaeus' Memoir of the Museum Adolphi Friderici Regis, Stockholm, 1754, in which first
appears the species name Elephas indicus 1309
1166. Facsimile of page 33 of Linnaeus' original tenth edition of the "Systema Naturae," 1758, in which Elephas maximus is sub-
stituted for Elephas indicus 1310
1167. Indian elephant group in the American Museum of Natural History. Specimens from the hills in the Province of Mysore,
shot in 1923 by Mr. Arthur S. Vernay 1311
1168. Elephas indicus sumatranus, pair of young elephants from Sumatra captive (1921) in the Zoological Park of Wa-shington.
.\fter photograph 1314
1169. The Sumatran elephant, apparently a female, living in the Am.stcrdam Zoological Gardens, August, 1913. After photo-
graph 1314
xxvi OSBORN: THE PROBOSCIDEA

FIGURE TAGE
1170. EhphoH indints rri/laniciis and E. indicus botgalrnsi.s, eraniti. Types. After dc IMaiiivillc 1816
1171. Crania of Elephus indicun (Dauntcla var.) and E. indicus (iMukna var.). After Falconer and Cautley 1317
1172. Elephas indicufs bengalensis, type, and E. indicus ceylanicus, referred, crania. After photographs 1317
1173. Map showing geograjihic distribution of the principal .species and subspecies (Hving and extnct) of Elephas, Hijpselephas,
and Plalelephas 1318
Platelephas platyrephalus, Hypselephas hysudricus, and Elephas indicus. Restorations of heads by Flinsch 1320
Elephas indicus hengalensis. A herd of wild elephants in a bamboo jungle of Mysore. After ])hotograph 1322
Elephas indicus, referred third inferior molar (vertical section) of an unusually large specimen from Assam. After Falconer
and Cautley 1324
Elephas asinlicvs. Blumenbach's original type figure of a first right superior molar 1325
Elephas indicus ref . Section of a partly worn third inferior molar. After Gaudry 1325
Elephas indicus hengalensis, known as the "giant tusker of Udiapur," showing abnormal length of tusks. After photographs. 1326
Comparison of crania of Elephas indicus ceylanicus and E. indicus hengalensis 1327
Elephas indicus sumalranus formerly living in the Rotterdam Zoological Gardens. After Lydekker 1329
Elephas sumalranus, cotype male and female crania from Palcmbang, in Leiden Museum. After photographs 1330
Elephas sumalranus crania (adult and infantile), in Munich Museum. After photographs 1330
Sumatra!! elephant from Batang Serangan. Mounted specimen in Munich Museum. After photograph 1331
Sumatran elephant (infantile) in Munich Museum. Mounted specimen. After photograph 1331
Mounted Burmese elephant. After photograjih 1332
Elephas indicus hirsulus, type, formerly living in the Gardens of the Zoological Society, London. Mounted specimen in the
British Museum (Natural History). After Lydekker 1333
Elephas indicus buski Matsumoto [=1Palseoloxodon huski], type first superior molar of the left side, from Japan. After
Matsumoto 1333
Second molar from between Kanagawa and Tokio (Yedo), Jajjan, referred to Palseoloxodon by the present
right superior
author. After Lydekker 1334
Superior view of heads of a young African elephant and of an adult Lidian elephant. After Geoffroy St.-Hilaire and
Frederic Cuvier 1335
Loxodonla africana and Elephas indicus, crown views of third right inferior molars. After Owen 1335
Comparison of low-browed African cranium {Loxodonla africana) and high-browed Indian cranium {Elephas indicus),
showing deeply embedded brain below cranial air cells 1335
Unguligradism. Radiograph of right foot of a young Indian elephant 1336
Estimated shoulder heights of Indian elephant, .skeletal and fle.sh 1337
"Elephas planifrons" and "Elephas hysudricus" life zones. After Pilgrim 1338
Map showing Upper Siwalik exposures of the Simla foothills, India 1339
Hypselephas hysudricus, type and jjaratype molars. Sections after Falconer and Cautley 1341
Hypselephas hysudricus, paratype third right inferior molar. After Falconer and Cautley 1342
Hijpselephas hysudricus, referred third left inferior molar. American Museum (Barnum Brown) collection 1342
Hypseliphas hysudricus, icferred second left inferior molar 1343
Hypselephas hysudricus, referred third left superior and inferior molar .sections. American Museum (Barnum Mrown)
collection 1344
Hypselephas hysudricus, referred third right superior molar, inner view, also transverse section, and photograph of occlusal
surface. American Mu.seum (Barnum Brown) collection 1345
Hi/pselephas hysudricus, referred superior and inferior molars from India. American Mu.seum (Barnum Brown) colhiction . . 1346
(
'ompari.soii of two crania of Hypselephas hysudricus with Mukna and Dauntcla varieties of Elephas indicus. After Falconer
and Cautley 1349
Hypselephas hysudricus, referred adult mal(> craniimi. After Falconer and Cautley 1350
Hypselephas hysudricus, portion of young jaw with greatly elongated rostrum. American Museum (Barnum Brown)
collection 1351
Plalelephas platycephalus, type cranium (palatal view). American Museum (Barnum Brown) collection 1352
Archidi.skodon planifrons, referred adult cranium of supposed female, with small tusks. After Falconer and Cautley 1352
Hypselephas hysudricus, male craniimi (palalal view). After Falconer and Cautley 1352
Hypselephas hysudricus, referred female cranium in Amherst Museum, collected near Kulhi, a district of the Punjab, by
M. M. Carieton 1353
1211. /////wefe/j/io.s/i.v.saf/c/rM.s, restored juvenile skull. .American Mu.seum (Barnum Brown) collection, .\fter jjliotograpli 1354
1212. Hypselephas hysudricus, referred cranium. After Falconer and Cautley 1354
1213. Hypselephas hysudricus, referred juvenile crania in British and American Museums 1355
 LIST OF ILLUSTRATIONS xxvii

FIGURE PAGE
1214. Ifypnelephas hysudricus, referred juvenile eniiiium and jaws. American IMufieuni (Barnuni Brown) collection 1356
1215. Fiypselephas hysudricus, referred right second superior molar, originally selected by Osl)orn as the type of Elephas platyce-
phalus angustidens. American Museum (Barnum Brown) collection 1357
1216. Upper Pliocene and Lower Pleistocene strata near Siswan, India, showing site where type cranium of Platelephas platyre-
phalus was found. After ])hotographs 1358
1217. Comparison of the types of Platelephas plaiycephalus and Stegodon pinjorensis. Cranial sections 1360
1218. Plalelephas platycephalus and Hypselephas hysudricus, right cranial profiles 1360
1219. Platelephas platycephalus, four aspects of type cranium. American Museum (Barnum Brown) collection 1361
1220. Geological relationships of African Proboscidea. Columnar section by Colbert 1422
1221. Geological relationships of Oriental Proboscidea. Columnar section by Colbert 1440
1222. Geological relationships of European Proboscidea. Columnar section by Colbert 1457
1223. Geological relationships of Asiatic Proboscidea.Columnar section by Colbert 1477
1224. Geological relationships of North American Proboscidea. Columnar section by Colbert 1491
1225. Geological relationships of South American Proboscidea. Columnar section by Colbert 1516
1226. Models of Recent and extinct Mammoths and Mastodons. After Knight 1522
1227. Map showing geographic distribution of the Moeritherioidea, Deinotherioidea, and Mastodontoidea 1528
1228. Map showing geographic distribution of the Stegodontoidea and Elephantoidea (including the Stegolophodontinae of the
Mastodontoidea) 1538
1229. Four-coned ancestral grinders of the Proboscidea (e.g., Mwritherium) compared with the six-coned Palitomastodon molars. . 1544
1230. Molar diagrams showing typical crown pattern (median sulcus, median conules, double trefoils, .serrate .spurs) in each of the
four families of the Mastodontoidea 1546
1231. Accelerated evolution of ridge-plates from Archidiskodon planifrotis into A. imperator; also Stegodon grangeri molar, with
enamel foldings, inserted to show \'-shaped valleys of the stegodontoid molar as compared with I'-shaped valleys of the
elephantoid molar 1547
1232. Brevirost rinse: Proversion of ridge-crests in Anancus 1548
1233. Humboldtinae: Retroversion and centroversion of superior ridge-crests in Cuvieronius and Stegomastodon 1548
1234. Crown view of third inferior molars of the right side of Loxodonta africana and Elephas indicus. After Owen 1549
1235. Archidiskodon subplanifrons, type third right inferior molar. Section showing cement, dentine, and enamel. Drawing by
D. F. Levett Bradley 1549
1236. Chief head and dental forms of four of the superfamilies (I H) of the Proboscidea (McEritherioidea, Deinotherioidea, Masto-
dontoidea, Elephantoidea) 1550
1237. Divergent adaptive radiation of crania and incisive tusks in six bunomastodont .subfamilies 1551
1238. Deinotherium giganteum, juvenile jaws showing replacement molars. After Lartet 1554
1239. Cencplasmie evolution of the archaic-toothed mammoths during a three-million-year period so far as known to April, 1935.
After Osborn 1581
1240. Alloiometrons: Adaptive speed and weight proportions. After Osborn 1581
1241. General climatic distribution of the subfamilies of the Ele])hantoidea and Stegodontoidea including theoretic migration
lines (1938). After American CJeographical Society North Polar Projection 1589
1242. Worldwide distribution of the Proboscidea in past and present time. Same as figure 6 of Volume wit h modifications
I , 1594
1243. Foot trail of Indian elephant "Gunda," formerly li\ing in the New York Zoological Park, taken in sand 1598
1244. Elephas indicus ref., showing contrast in proportions between adult and young. After photograph by Underwood aiul

Underwood 1599
 Chapter XIV

THE ROOF-TOOTHED STEGODONTS, SUPERFAMILY STEGODONTOIDEA

Possible ancestry in the miocene zygolophodonts of western europe. Primitive forest-browsing

ELEPHANTS OF THE ORIENTAL REGION. SlOWLY PROGRESSIVE FROM MIDDLE MIOCENE TO MIDDLE [upPEr]
pleistocene TIME. BrACHYODONT TO SUBHYPSODONT, BUT PROGRESSIVE RIDGE FORMULA. EXTINCTION IN
MIDDLE [upper] PLEISTOCENE TIME.

I. Introduction. Twenty-third species, Stegodon bondolensis, Java.

1. History of classification. Twenty-fourth species, Stegodon trigonocephalus prae-
cursor, Java.
2. Habits and general characters.
Twenty-fifth species, Parastegodon? hvantoensis, Japan.
3. Approximate descending ridge formulae, after Falconer,
Twenty-sixth species, Stegodon yiishensis, China.
Lydekker, Martin, and Osborn.
Twenty-seventh species, Stegodon officinalis, China.
4. Geologic and diphyletic order of the Stegodontinse. Twenty-eighth species, Stegodon zdanskyi, China.
5. History of discovery of the subfamily Stegodontinse. Twenty-ninth species, Parastegodon [Stegodon?] sugi-
Principles of type revision of the species. yamai, Japan.
6. The Stegodontinse and Mastodontinae of China. Thirtieth species, Stegolophodon lydekkeri, Borneo.
7. Pliocene to Pleistocene Proboscidea of Japan.
8. Phylogenetic discussion of the thirty described species III. Systematic Arrangement of the Stegolophodonts and
of Stegodonts and Stegolophodonts.
Stegodonts in Phyloge.netic Order.
Probable African-European-Asiatic origin and migra- 1. Characters of the subfamily Stegodontinse.
tion of the primitive Stegodonts. 2. History of the generic names assigned to the Stegolopho-
donts and to the Stegodonts.
II. Type Revision of the Species in Order of Original Generic characters of Stegolophodon Schlesinger.
Discovery and Description. Systematic description of species of Stegolophodon.
1. The first two Stegodonts, discovered in Burma, 1828. Stegolophodon cautleyi of the Upper Miocene
Mastodon laiidens Clift. [Middle Pliocene], Perim Island.
Mastodon elephantoides Clift. Stegolophodon latidens of the Lower Pliocene
2. Discoveries in India and Burma (1845, 1846). [?Lower Pleistocene] of Burma and [Middle
Third species, Elephas insignis, India. Pliocene] of India.
Fourth species, Elephas ganesa, India. Stegolophodon sublatidens of the Middle(?) Pliocene
Fifth species, Elephas hombifrons, India. of Austria.
Sixth species, Elephas cliftii, Burma. Stegolophodon stegodontoides of the Upper(?) Plio-
cene of India.
3. The Stegodonts of China, India, Java, the Philippine
Stegolophodo7i nathotensis of India.
Islands, Austria, Japan, and Burma.
Stegolophodon cautleyi progressus of India.
Seventh species, Stegodon sinensis, China.
Stegolophodon lydekkeri of Borneo.
Eighth species, Stegodon orientalis, China.
Ninth species, Mastodon cautleyi of Perim Island.
Tenth species, Stegodon trigonocephalus of Java. IV. Succession of Species of the Genus Stegodon.
Eleventh species, Stegodon mindanensis, Philippine Genus Stegodon Falconer and Cautley, 1847, 1857.
Islands. Skulls of Stegodonts in the British and Indian Museums.
Twelfth species, Stegodon airdwana of Java. Characters of referred skulls of Indian Stegodonts,
Thirteenth species, Stegodon ganesa var. javaniciis of after Falconer, 1868.
Trinil, Java. Stegodont crania of China and of the East Indies.
Fourteenth species. Mastodon stegodontoides of Lehri, Systematic description of species of Stegodon.
India. Stegodon sinensis of the Yangtze River, China.
Fifteenth species, Elephas (Prostegodon, Parastegodon) Stegodon elephantoides of Burma.
auroTBe of Japan. Stegodon cliftii of the Irrawaddy River, Burma, and
Sixteenth species. Mastodon (Bunolophodon) longi- of the Dhok Pathan horizon, India.
rostre Kaup forma siiblatidens of Austria. Falconer's Notes of 1868 on Stegodon cliftii.
Seventeenth species, Stegodon orientalis shodoensis of Stegodon bombifrons of the Lower [Middle] Pliocene,
Japan. Dhok Pathan horizon.
Eighteenth species, Stegolophodon nathotensis, India. Falconer's Notes of 1868 on Elephas [
= Stegodon]
Nineteenth species, Stegolophodon cautleyi progres- bombifrons.
sus, India. Lydekker's Notes of 1886 on Elephas [
= Stegodon]
Twentieth species, Stegodon orientalis grangeri, bombifrons.
China. Stegodon insignis of the Upper Pliocene [to Ujjper
Twenty-first species, Stegodon insignis birmanicus, Pleistocene] of India.
Burma. Stegodon ganesa of the Upper Pliocene [to Upper
'J'wenty-second species, Stegodon pinjorensis, India. Pleistocene] of India.

805
 '

806 OSBORN: THE PROBOSCIDEA

Falconer's Notes of 1868 on Stegodon insignis Middle Pleistocene, Java (synonym of .S'.

and S. ganesa. airdwana or S. irigonocephaliis).

Lydckker's (1886) comparison of Stegodon in- Stegodon (Archidiskodon?) mindariensis of Minda-
signis, S. ganesa, and S. bombifrons. nao, Philippine Islands.
Stegodon insignis-ganesa a collective sjjecies. Stegodon aurone of Mt. Tomuro, Japan.
Stegodon insignia birnianicus of Burma. Stegodon orientalis shodoensis of Japan.
Stegodon orientalis grangeri of China.
Cranial characters of .S. orientalis grangeri. Recently described Stegodonts fkom Java, China, and
Stegodon pinjorensis of India. Japan.
Stegodon orientalis of Szechuan, northwest China. Stegodon bondolensis of Java.
Lj'dekker's Notes of 1886 on Stegodon orientalis. Stegodon trigonoccphalus praecursor of Java.
Stegodon airdirana of Trinil, Pithecanthropus ereclus Parastegodon'^. kwantoensis of Japan.
zone, Java. Stegodon yilshensis of ("hina.
Stegodon airdwana fauna, Kendeng-Schichten Stegodon ojficinalis of China.
layer of Trinil, Java. Stegodon zdanskyi of China.
Comparison of Stegodon airdirana of Ja\-a with Parastegodon [Stegodon'!] sugiyawui of Japan.
Stegodon insignis-gane.sa of India.
Stegodon Irigonocephaliis of the vicinity of Surakarta, Ari'ENDix: Matsumoto on the phylogeny and classification
Java. of the Japanese Mastodonts, Stegodonts, and Elephants
Stegodon ganesa vav.javaiiiriisof theTrinil horizon, (1924-1927).

INTRODUCTION
I.

1. HISTORY OF CLASSIFICATION
[The name Stegodontoidea first appeared in a diagram by Professor Osborii in his article of June, 1935,

entitled, "The Ancestral Tree of the Proboscidea. Discovery, Evolution, Migration and Extinction over a
oO,0()0,()00 Year Period" (Osborn, 1935.937, p. 407, fig. 2), in which superfamily he included both Stegolophodon
and Stegodon under the family Stegodontidse of Young-Hopwood, thus removing them from the Elephantoidea.
In Volume I of the present Memoir (published Aug. 15, 1936) Professor Osborn confirmed his separation of
the Stegodontoidea from the Elephantoidea (pp. 22, 25) but he withdrew the Stegolophodonts, placing them in

the superfamily Mastodontoidea, family Mastodontidse, and creating a new subfamily, the Stegolophodontinse
(see pp. 700, 737, and PI. n ), to embrace the various species, owing to the "intermediate position of [the] molars

between the true Mastodontidse . . . and the true Stegodontoidea," and suggesting (p. 191) the "possible derivation
of the grinding teeth of the Stegodontoidea from those of Stegolophodon." The true Stegodonts he retained in
the superfamily Stegodontoidea, subfamily Stegodontinai.

As early as 1857 Falconer observed (1857.1, p. 314) that "The Stegodons constitute the intermediate group
of the Proboscidea from which the other species diverge through their dental characters, on the one side into the
Mastodons, and on the other into the typical Elephants." Later Falconer (in Murchison, Pal. Mem., 1868,
\'ol. II, footnote, p. 268) remarked that "The Indian fossil species, which have been ranged under th(> designation
of Stegodon, establish, through their molar teetli, a manifest and nearly unbroken passage from tlie Mastodons
into the tru(> Elephants." Also, as recently as 1932, van der Maarel (1932.1, p. 162) expressed the opinion that
"all the species of Stegolophodon . . . being all v(>ry primitive forms . . . may as well be reckoned to the family of
t he M asiodontidx .'

Therefore, while various scientific observers have regarded certain of the Stegodonts as transitional Ix'tween
the Mastodontida^ and the Elephantida^, it remained for Professor Osborn to assign the superfamily name Stego-
tlontoidea to the true Stegotlonts and to remove the Stegolophodonts to the sui)erfamily Mastodontoidea, under
the new subfamily name Stegolophodontinse, the members of which he designated (Vol. I, j). 690) as "pro-stego-
donts."

Tlie regrettable deatli of Professor Osborn in No\eniber of 1935 precludes the full treatment of lliese groups
as contemplated by him (see Vol. I, p. 197) ; it is deemed best, therefore, to allow the present chapter to remain as
 THESTEGODONTIN^: HISTORY 807

first written by him, making such changes (either in square brackets or in editorial notes) as are consistent with
the known opinions of the author, namely, as deduced from pubUshed statements, accumulated notes, and
interlined text.
The classification of the genera Stegolophodon and Stegodon, now to be described, would appear, therefore, to
be as follows:

SuPERFAMiLY : STEGODONTOIDEA Osborn, 1935, 1936

Separated by Osborn (Vol. I, 1936, pp. 22, 25, of the present Memoir) from the Elephantoidea Osborn, 1921,
as a distinct stock, but without diagnosis. However, according to his observations on the sectioned molars, the
valleys separating the adjacent ridges are closed or V-shaped at the bottom in the Stegodontoids (Fig. 764)

and open or U-shaped in the Elephantoids (Fig. 1231). He also considered that the extremely short face of the
Stegodontoids could not have given rise to the longer face of the Elephantoids.

Family: STEGODONTIDiE Young-Hopwood, 1935

This name appears in Young (1935.1, p. 5) but without definition. Later in 1935, Hopwood, in his Memoir
on the "Fossil Proboscidea from China," defines the family as follows (p. 71): "The animals included in this
family have skulls which resemble those of the true elephants but which are more primitive. They have very
long sockets for the tusks, and the grinding teeth and palate are well below the plane of the occipital condyles.
The grinding teeth remain brachyodont throughout the whole of their evolutionary history, but they parallel
the teeth of the true elephants in showing a progressive increase in the number of ridges. This is especially true
of the third molars. With this increase in the number of ridges, and its accompanying increase of length, there

is an ever increasing curvature of the occlusal surface which reaches its maximum in certain specimens referred to
S. airmmna and S. insignis. Owing to the short palate, it was impossible for the whole of the very long teeth to
be accommodated in the upper jaw at one time. The curvature of the crown allowed the tooth to follow a more or
less circular path which brought it from a position practically parallel to the plane of the occiput to the correct

position for mastication. There is an ever increasing amount of cement, and the ridges show an increasing number
of mammillae on their crests."

"This sub-family comprises two groups''' of animals. One, with compressed, tectiform, ridges is given the
generic name Stegodon Falconer & Cautley. The other, in which the ridges are blunter, and composed of round-
ed conules, is known as Stegolophodon Schlesinger. Both genera occur in India, but, so far as is known, Stegodon
is the only genus found in China."

Subfamily: Stegodontin^ Osborn, 1918, 1921

Original reference: Bull. Geol. Soc. Amer., 1918, Vol. XXIX, pp. 135, 136 (Osborn, 1918.468); Araer. Mus. Novitates, 1921,
No. 1, pp. 12, 13 (Osborn, 1921.515).

Subfamily Definition (Osborn, 1918.468, p. 136): "The Stegodontinse may be distinguished as

a phylum confined to Asia, in which the grinding teeth remain brachyodont, short-crowned, although
a very large number of cross crests evolve, especially on the posterior grinding teeth. From an early
member of this subfamily, perhaps of Middle Miocene time, were given off one or more branches of the
elephant and mammoth phyla."

(Osborn, 1921.515, pp. 12 and 13): "We observe that the Stegodonts are persistent browsers,
probably tropical, forest-living proboscideans. According to Pohlig, from the skeleton discovered in
'[Apcording to Osborn, 1936, Vol. I, p. 700, by removal of the Stegolophodonts to the family Mastodontidse, the Stegodontidae embrace the true Stego-
—
donts {Stegodon Falconer and Cautley) only. Editor.]
808 OSBORN: THE PROBOSCIDEA

Trinil, Java, they have short, massive bodies hke those of the Mastodontinse of the north temperate
forests. The skull and tusks do not lead into either the Elephantina^ or the
Mammontinae types The . . .

distinctive featiu'e of the grinding teeth is the rapid multiplication of transverse crests which rise from
the fornuila 4.5.6.6.7-8 in S. [Stegodon] ciiftii (Lower Pliocene) to 5.9.10.12.13 in S. insignis (Lower Pleis-
tocene). Jaw rapidly abbreviated. Upper tusks straight, parallel, slightly upcurved (adapted to dense
forests). Grinders brachyodont to subhypsodont, crests breaking up into small mammillae, valleys
filling with cement."
Li the first article, namely, "A Long-jawed Mastodon Skeleton from South Dakota and Phylogeny of the
Proboscidea," 1918, Osborn placed the Stegodontinae under the Elephantidae (p. 135), and in an accompanying
table mentioned the species Stegodon ganesa, S. ciiftii, S. bombifrons, and S. latidem. The last-mentioned species
was made by Schlesinger the genotype of Stegolophodon.
The above on both Stegoloplwdon and Stegodon. After the removal of the former
definitions were based
genus to the superfamily Alastodontoidea, family Mastodontidse, subfamily Stegolophodontinae, no revised
definition of the subfamily Stegodontinae was given by the author.

LATE Tl':inTARY (iEOLOGY OF INDIA

The geology of Lidia (the Siwalik Hills and Perim Island) will be found in Edwin
latest opinion regarding the

H. Colbert's Memoir of 1935 on "Siwalik Mammals in The American Museum of Natural History," pages 6 to 55;
in a summary on page 21 it will be noted that Doctor Matthew in 1929 assigned to the Siwalik beds a somewhat
liigher position in the geologic time scale than Doctor Pilgrim in 1927. also Chapter XXII Compare of the present

Memoir. "Geologic Succession of the Proboscidea," which has been written by Doctor Colbert.

Throughout the present Volume, therefore, the later determinations will be inserted in square brackets or
added in footnotes. It should be recalled that the chapters constituting this Volume were written about eight to
ten years ago and were awaiting final revision by Professor Osborn. Editor.] —
2. HABITS AND GENERAL CHARACTERS
We observe that this subfamily [Stegodontinae^] includes forest-hving browsers, which probably developed in a
Fig. 20. forested or semi-forested oriental region, ranging through
India into Burma, China, Japan, and southward into
Java, Borneo,^ and the Philippines. The ancestors of the
subfamily may Miocene deposits of western
be found in

Europe\ A single tooth, named Mastodon {Bunolopho-

don) longirustre Kaup forma nublalidem, has been de-

Hfjihas c!>f/i.— The first (?) lolt upper true molar : from (lie .'^Iwaliksof liuniia.

I. Tlie lowpr borflor of tlic fi;:;uro is t)ic inner border of tlie ppccimcn.
(KromGaudrTS '
Kncliainements.)

CoTYPE OK Stegodon ki.kpiiantoidks Glut (=cLii'rn Falconeu)

Type of Ekphas ciiftii Falponer and Cautloy, 1846, a first loft
Kig. 683. RbFERKED superior MOT.AII OF STE(i()DON OKIBNTAI.IS OKANOEKl
upper true molar, l.M', onc-lialf natural si:!e. The same six erestcd tooth I'^ig. 684. Referred first right .superior molar, r.M' (rev.) of Slnioilnn

api)ear,s in figures 686, 7()0, and 701. After Lydekker, 1S86.2, j). 81, fig. 20 oricnlalis grangcri (Amer. Mus. 18530- wrongly numbered 18.530). Acluiil

(taken from a woodcut m (laudry, 1878, p. 176, fij,'. 232). median length 181 mm.

'[At the time this chapter was written the subfamily Stegodontina; was thought to embrace both the .Stegolophoduuts (now removed to the Mnstod.mlnidea
subfamily Stegolophodontina;, p. 700) and the true Stegodonts. Editor.] —
•'[Type locality of Slegolophodon lydckkeri Osborn, 1936 (see Vol. I, p. 700). -Editor.;
'[Sec Vol. I, p. 197.— Editor.)
 THESTEGODONTIN.^: HISTORY 809

scribed by Schlesinger from near Teschen (Schlesien), Austria; this resembles Mastodon [
= Stegolophodon\
lalideiiH of Burma and is icferred in the present Memoir to Stegolophodon subldtidens (see Vol. I, p. 737; Vol. 11,

p. 846).

The skull is subelephantine in type, brachycephalic, brachyopic, the rostrum being elongated to support the
tusks; the grinding teeth and palate are depressed far below the occipital condyles (bathycephalic). As in the

elephants, the jaw is greatly abbreviated. The upper tusks are straight or slightly upcurved, elongating, without
trace of enamel band, to a length of about 10 ft. Considering the large size of the tusks the skull is relatively
small. The grinding teetli are brachyodont to subhypsodont, yet the ridge-crests in the posterior molars, M3,
multiply from five plus [Stegolophodon] to fifteen plus [Stegodon], each crest breaking up into small nipples, mam-
illae, or conelets. The lower incisors disappear very early.

As in certain of the Mastodontidee and as in all the Elephantidse, the grinding teeth increase the number of
their ridges by adding crests both in front and behind, as first observed by Falconer. Thus in M 1 of Stegodon
"""g"""
orienlalis grangeri (Fig. 684) the ridge formula may be written: . The descending scale of the ridge formulae

in the premolars and molars of the principal species of Stegodonts is appro.ximately as below.

3. APPROXIMATE DESCENDING RIDGE FORMUL.^, AFTER FALCONER, LYDEKKER, MARTIN,

AND OSBORN
Observe that (1) the maximum upper ridge-crests, rising from 21e to 51e, are, so far as known, less numerous
than the maximum lower ridge-crests which probably rise to more than 51 ; (2) this is compensated for by the fact
that the upper molars are throughout broader than the lower molars; (3) the differences in width and in the
number of ridge-crests of the upper molars in comparison with the lower molars are beautifully shown in figure 687,

also in figures 759 and 762, the type of Stegodon orientalis grangeri. See also details of progressive ridge-crest
formulae, talons and half ridge-crests, under each species.

Maximum Estimated maximum

conelets upper and lowek
 1

810 OSBORN: THE PROBOSCIDEA

The individual number of ridge-crests from Dp 2 to M3 is mainly assembled from a very careful collation of

the ridge-crest formulae given by Falconer (see below), supplemented by the observations of Lydekker, of Martin,
and of Osborn.
CoNELETS. —The conelets of Stegolophodon and of Stegodon arise chiefly by binary, rarely by ternary, fission
of the primary cones. Starting with the original loph of the Palseomastodon stage, consisting of two cones, normal
binary fission would produce:
2 4-t- — {S. cautleyi) —8 (*S'. elej)hantoides) — 20-1- i>^- airdwana), etc.

But the binary fission is not so regular as this. The newer anterior and posterior crests exhibit fewer conelets
tluin the older mid-crests, so that the highest number of conelets usually will be found in the third and fourth
crests.

6 J crests
STEQOLOPHODON 5TE60D0NT0IDE5

4--^ coTtelets 5TEQ0L0PH0D0N LATIDEN5

STEQOLOPHODON CAUTLEYI

PALAE0MA5T0D0N
Fig. 68.5. Structural Evolution of the Cone.s, Conelets, and Ridqe-crbsts in the Steoo-
LOPHODON Phylum, in comparison with Pal.«omastodon
(1) PAL.EOMA8TODON, PRIMARY Type. Four jjiimary cones; ridgc-ficsts, jjioto-, meta-, ami
—
rudimentary tritoloph. [See pp. 143 and (191 of Vol. I, for subfamily position of Valxomaslndon. Editor.
(2-4) Stegolophodon Phylum. Binary fission of primary cones into four to five coneUls, vestigial
conules in S. cautleyi and .S. lalidens (protoconule = p.l., metaconule = m.l.); addition of trito-, tetarto-,
pcnta-, and hexalophs. Gradual loss of median sulcus, presistent, however, in the first two anterior
crests.
Stegolophodon Uilidens (3, left) is a third lower molar, r.Ma, inserted for romparison, and has seven
phis (7J2) ridge-erests, four plus conelets, and median sulcus.
 )

Lrni
/5 crests 20 or r,,ore ConeZet 10 STEGODON AIRAWANA
-fissures

i-4 crests // conelets 9 STEQODON IN51GNIS- QANESA

/34 crests // or less conelets 8 STEGODON ORIENTALIS qPANQERI

7 STEQODON BOMBIFRONS

W crests 5 To 8 coneZets 5TEGOD0N E LEPHANT0IDE5

Iml

a^ crests JO -^ coneZets 5 STEQODON ELEPHANT0IDE5 (--CLIFTII

Fig. 686. Strdctdral Evolutiox of the Cones, Conelets, and Rxdge-crests in the Stegodon
Phylum, in Ascending Order (5-10).
(5-10) Stegodon Phylum. Binary or ternary fis.sion of the cones into conelets (5-20); addition of
anterior and po.sterior ridge-crests (10-15 inM 3); addition of cement.

(5) Stegodon ekphantmdes (=cliftii)

with six and a ([uarter ridge-crests and ten plus conelets (M').
Stegodon elephantoides with ten ridge-crests and five to eiglit conelets (M3).
(6)
(7) Stegodon bombifrons witli nine and a
half ridge-crests and eleven plus conelets (M^).

(8) Stegodon orientalis gnmgeri with thirteen

and a half ridge-crests and eleven or less conelets (M3).
(9) Stegodon in&ignis-ganesa
with fourteen ridge-crests and eleven conelets (M3).
(10) Stegodon airhwana with fifteen ridge-crests and
twenty plus conelets (M').

811
gl2 OSBORN: THE PROBOSCIDEA

—
Figures 685 and 686. Particularly interesting and significant in the Stegodontinae is the transformation of
the original cones by fission into conelets. Thus in the archetypal Palasomastodon molar there are two cones in
the protoloph, in Stegolophodon latidens and cautleiji each cone splits into two, making four plus conelets in the
»S'.

metaloph; in *S. stegodontoides each of these four conelets tends to split into two, tending to form from five to
eight conelets, but this splitting is not regular and no loph actually attains eight. In Stegodon elepkantoides the

equal splitting gives rise to from^we to eight conelets; in S. elepkantoides ( = cliftii), S. bombifrons, and *S'. insignis-

ganesa the fission gives rise to from eleven to twelve conelets, hence each ridge is finally surmounted by twelve cone-

lets which, when slightly worn, present eight loops. In Stegodon airdwana, the most progressive species, the
conelets range from thirteen to twenty plus. Thus the maximum number of cones and conelets in each crest runs

as follows:

Molar cones and conelets: Primitive (Palxomastodon) 2-4-6-8-12-20+ progressive {Stegodon airdwana).

Ridge-crest Evolution. —The Stegodontinae also furnish a beautiful example of the evolution through
w liich each ridge passes in turn, from the primitive submastodontoid type seen in Palseomastodon into the highly
progressive subelephantoid type seen in Stegodon insignis-ganesa and S. airdwana. The superior (-loph) and

inferior (-lophid) ridge-crests may receive a brief numerical terminology, namely:

Pro-protoloph— id = One-half, anterior rudimentary ridge Pentalopli—id = Fifth ridge

Post-metaloph— id = One-half, posterior rudimentary ridge Hexaloph—id = Sixth ridge
Protoloph—id = First primary ridge = protocone and Heptaloph— id = Seventh ridge
paracone of Ungulata Octaloph—id = Eighth ridge
Metaloph —id = Second primary ridge = hypocone and Ennealoph— id = Ninth ridge
= Tenth ridge
metacone of Ungulata Decaloph—id
Tritoloph— id = Third ridge Endecaloph—id = Eleventh ridge
Tetartoloph— id = Fourth ridge Dodecaloph— id = Twelfth ridge

Crest Addition. — In Stegolophodon and in the Mastodontidae the homology of the protoloph and of the
metaloph is simple as compared with other ungulates, but since the increment of ridge-crests in the elephantine
molar by addition to both the anterior and
is posterior ridges, namely, the pro-protoloph and the post-metaloph,
it soon becomes difficult to determine which ridge-crests correspond with the primary protoloph and metaloph
of Palseomastodon and of other ungulates.

Intermediate Molars Uniform. —A constant feature in the Proboscidea appears to be the uniformity of
the three 'intermediate molars,' namely. Dp 4, Ml, M 2, which tend to have the same ridge-crest formula in

each species, for example:

Stegodon insignis-ganesa Dp 4 7K> Ml 7}^ M2 7K-8 = intermediate molars

Stegolophodon latidens Dp 4 4 M 1 4^ M 2 4^-5 = intermediate molars
Trilophodon angastidens Dp 43 M13 M23 = intermediate molars

Consequently it may be difficult to distinguish these 'intermediate molars' from each other by the ridge formula
alone; whereas they may be distinguished by the character of wear, by the width of the crowns, and by the
condition of the fangs.

Ridge-crest Elevation.— The progres.sive elevation of the ridge-crests in the Stegodontinae is illustrated in
two diagrammatic figures (Figs. 687, 688), which demonstrate the constant progressive heightening of the ridge-
crests as we ascend from the Lower(?) Phocene Stegodon sinensis and [Middle Pliocene] S. bombifrons into the
Middle Pleistocene S. airdwana stage. The early phases of ridge-crest elevation (Fig. 687) may be compared with
the later phases (Figs. 688, 781) as follows:
Stegodon orienlalis Owen Stegodon aurorx Matsumoto
Stegodon Osborn
orienlalis grangeri Stegodon airdwana Martin
Stegodon bombifrons Falconer and C'autley Stegodon insiqnis l-'alconer and Caul ley
Stegodon sinensis Owen
 THE STEGODONTINiE: HISTORY 813

Descending Order of Species. — (1) At the summit of the known Stegodontinae is Stegodon airdwana, the
most progressive both in ridge-crests and in conelets —a Lower to Middle Pleistocene stage. The following extract
from a letter by Doctor Dietrich (March 10, 1924, and notes) confirms the Middle Pleistocene age of this species:

For a long time I have been trying to prove that the Stegodon [airdwana] species from the Pithecanthropus strata [Trinil,
Java] are the very youngest, that is, young Pleistocene. In detail: "Geologisches Alter. Aus morphologischen (Jriinden muss
die javanische Art jiinger sein als die chinesische und alle bekannten kontinentalen Stegodon-Arten. Dass St. Airawana im
Pleistocan lebte, wird \on fast alien Autoren angenonimen ihr pleistocdnes Alter diirfte gesichert sein. Erweist sich die chines-
;

ische Stegodontenfauna als alt-pleistociin, dann ist die Trinilfauna jiinger als alt-pleistociin. Ich halte es aus geologischen und
anderen Griinden sogar fiir wahrscheinlich, dass die Trinilfauna (und damit Pitheacanthropus) ju/i^-pleistociin ist; das wird sich
mit Hilfe des St. Airawana bei besserer Kenntnis der kontinentalen Stegodonformen vielleicht erweisen lassen."

znner yict^ All '/^ nat: si7ef

5/ \y ^-'r.c/pl

5TEQ0D0N OR1ENTALI5 %
Type
STEQODON AIRAWANA rs/.
Ca^t. A.M. 6 33S

STEQCDON ORIENTALIS QRANiqERl

A.M./aiOS •%-

r. m.-
AURORAE
Type

B0MBIFR0N5 %-"

5TEQ0DON SINENSIS
-Type % STEQCOON
-Type
IN5IQNIS

Fig. 687. Gradual Progressive Hypso-

DONTY IN Superior Grinders Fig. 688. Gradual Progressive Hypsodonty in Superior
AND Inferior Grinders
Stegodon orientalis Owen, type. Superior
deciduous premolar, r.Dp'. Stegodon airawana Martin. Middle Pleistocene of Java.
Stegodon orientalis grangeri Osborn. Superior Elevated and approximated ridge-crests (9-l.j}'>), right third
deciduous premolar, r.Dp'. inferior molar, r.Ms.

Stegodon bombifrons Falconer and Cautley. Stegodon aurorx Matsumoto, type. Middle Pleistocene
Third superior molar, M'. [?Upper Pliocene] of Japan. Elevated and closely approximated
Stegodon sinensis Owen, type. Superior ridge-crests, right second superior molar, r.M^.

deciduous premolar, r.Dp'. Stegodon insignis Falc. and Caut., type. Lower Pleistocene,
AH figures four-fifths natural size, e.\cepting Upper Siwaliks, Boulder Conglomerate, of India. Third superior
Stegodon t>omhifronswhich is four-fifteenths molar, M', with eleven-)- ridge-crests.
natural size.Observe that the ridge-crests (1-5) All figures to same scale, one-third natural size. Observe that
in transition from S. sinensis type to S. orientalis in Stegodon airawana and S. aurorx the ridge-crests are much
type are broad, progressively elevated, and more elevated and approximated than in S. insignis, but that S.
approximated. irisignis is less elevated than S. orientalis (Fig. 687).
Table III FSTEGOLOPHODON PHYT.UM RTECODON PHYLrM
 ^

THESTEGODONTIN^: HISTORY 815

(2) Next in descending order are referred Stegodon insignis and S. ganesa, of Lower to Middle [Upper] Pleisto-
cene age. (3) Third in descending order is Stegodon orientalis grangeri, slightly more primitive than S. insignis
both in ridge-crest and conelet progression —probably of Lower Pleistocene age. (4) Next is the [?Upper Pliocene]
Stegodon auroras of Mt. Tomuro, Japan. (5) Far more primitive both in ridge-crests and conelets are Stegodon
bombifrons [of the Middle Pliocene] and *S'. elephantoides ( = cliftii), known to be of Lower Phocene (type) age.'
It is probable that the referred *S. bombifrons of the Middle [Upper] Pliocene, Tatrot horizon, will prove to be some-
what more progressive than the lectotype of S. bombifrons of the Middle Pliocene, Dhok Pathan horizon.
(6) Representing the Stegolophodon phylum is the Upper(?) Pliocene Stegolophodon stegodontoides type, of
approximately the same age as Stegodon ganesa lectotype and S. orientalis grangeri type. (7) Stegolophodon
latidens ref. is recorded in the same Middle Pliocene (Dhok Pathan) geologic level as Stegodon bombifrons lecto-
type. (8)The Upper Miocene [Middle Pliocene] yields Stegolophodon cautleyi lectotype of Perim Island (Dhok
Pathan). (9) The Middle Miocene [Mio-Pliocene] yields Stegolophodon cautleyi progressus and S. nathotensis.

4. GEOLOGIC AND DIPHYLETIC ORDER OF THE STEGODONTIN^

The geologic level of Falconer's types of Elephas [ = Stegodon] ganesa and E. [
= S.] insignis, although unre-
corded, is probably Pinjor (see Vol. I, Fig. 413). Pilgrim writes (letter of January 26, 1927) : "I am now almost
convinced that the bulk of the fossils were collected from that zone, i. e.. Upper Pliocene [Pinjor zone, Moginand],
but that some came from the Boulder Conglomerate zone Falconer distinctly states." As observed by Barnum
Brown, the exact geologic horizon of the type and referred specimens of S. insignis and S. ganesa is uncertain,
because apparently in the Pinjor horizon are obtained jaws and skulls redeposited from the Boulder Conglomerate
above.

5. HISTORY OF DISCOVERY OF THE SUBFAMILY STEGODONTIN^. PRINCIPLES OF TYPE

REVISION OF THE SPECIES
Genera. — Up to 1924 sixteen species of [Stegolophodonts and] Stegodonts had been discovered and described
between the years 1828 and 1917, in the order shown in the list in Section II below. In 1924 Matsumoto described
a new subspecies from Japan, namely, Stegodon orientalis shodoensis. In 1929 Osborn described Stegodon orientalis
grangeri from the pits near Wanhsien, Szechuan, China, Stegodon insignis birmanicus from the Pliocene of Burma,
Irrawaddy River, and Stegodon pinjorensis from the Lower Pleistocene (?Pinjor) of India; also two Stegolopho-
donts (Stegolophodon nathotensis and S. cautleyi progressus). All the early species were described either as
Mastodon {e.g., Mastodon latidens, Mastodon elephantoides) or as Elephas (e.g., Elephas insignis, Elephas ganesa).
Although the name Stegodon appears as early as 1847 ("Fauna Antiqua Sivalensis," Falconer and Cautley, PI.

XLii), it was not until the year 1857 (pp. 314, 318, table opp. p. 319) that Falconer ventured to select Elephas

cliftii, E. bombifrons, E. ganesa{?), and E. insignis as representing a new subgenus, Stegodon.

It was in 1922 that Matsumoto in a letter to the present author announced his intention of making Mastodon
latidens the type of a new genus, Prostegodon. The name, however, is preoccupied by the Stegolophodon of Schle-
singer, 1917 (genotype Mastodon latidens Clift), in which genus Osborn united (1929) the four species M. latidens
Clift, M. cautleyi Lydekker, M. stegodontoides Pilgrim, and M. (Bunolophodon) longirostre Kaup /orma sublatidens
Schlesinger, also Stegolophodon nathotensis Osborn from the lower Middle Siwaliks and Stegolophodon cautleyi
progressus Osborn from 2,000 feet above the base of the Lower Siwaliks, India.
'[See footnote on preceding page (p. 814) regarding the Lower Pleistocene age of Stegodon elephantoides i=diftii). Editor.] —
^[To these six species should be added Stegolophodon lydekkeri Osborn, described in Vohime I of the present Memoir, page 700. — Editor.]
 —

816 OSBORN: THEPROBOSCIDEA

[Recently six additional species of Stegodon have been described, namely, Stegodon boiidolensis van der Maarel
1932, and *S'. trigonocephalus praecursor von Koenigswald, 1933, from Java, *S. officinalis, S. zdanskyi Hopwood,
1935, and S. yunhensis Young, 1935, from China, also Paraslegodon [Stegudon?\ sugiyamai Tokunaga, 1935, from
Japan. The generic determination of Tokvmaga's species Paraslegodon? kwantoensis, 1934, has not been given in
the present Memoir, owing to Professor Osborn's views regarding the genus Paraslegodon (see next paragraph).
Editor.]

The genus Paraslegodon of Matsumoto, 1924, belongs to the genus Archidiskodon or to a progressive
Slegodon, as the genotypic species, Elephas {Paraslegodon) aurorx, is slightly more primitive than Archidiskodon
planifrons.

Principles of Type Revision. —The consideration of these matters of descent and phylogeny, however,
must be preceded by a very rigid review of each species of Stegodont in the order of its original description,
following the standard methods of type analysis established throughout this Memoir, namely:

1) Determination of the actual type specimen or the specimen first mentioned among a series of cotypes.

2) Fixation of the original lype figttre, the one first published by the author or selected by the author
from other publications.

3) Enumeration of the lype characters observed in the type specimen by the author or by subsequent
observers.

4) Elimination from the type list of characters foimded on referred specimens which do not actually
belong to the same species as the type.

5) Determination of the type locality and distinction of the topotypes.

Original Descriptions. —The revision of these various species of Stegodonts, on the strict application of

the five rules above, has been a long and very difficult task. In order to establish absolutely the original author's
intention, the author's original description is cited in full. When too prolix, as in the case of many descriptions by
Owen and by Lydekker, excerpts are made in the author's own language.

This revision and establishment of type characters must be followed by a restudy and revision of referred speci-
mens, which in most cases can only be done in future by monographic research directly upon the specimens them-
selves, amphfied by knowledge afforded by fresh materials. Consequently the present Stegodont chapter lays no
claim to completeness or finality ; it leaves many questions wholly undecided, for example, the sexual or specific
dental or cranial distinctions between Slegodon insignis and S. ganesa.

In the meantime our present establishment of the types and of the type specific characters based upon the
type specimens themselves as well as of characters derived from properly referred specimens in the same geologic
horizon, as in the case of Slegodon bombifrons, may lay a firm foundation for future monographic research.

6. THE STEGODONTINJ^ AND MASTODONTIN/E OF CHINA

Two species of Stegodonts described by Owen in 1870, namely, Slegodon orienlalis and S. si^ietms, were
determined from (collections l)rought in by dealers for medicinal purposes and not procured in sitii by palieontolo-
gists.
 : :

THE STEGODONTIN^: HISTORY 817

a) Koken's Notes of 1885 and Schlosser's Notes of 1903

Koken, "Ueber fossile Siiugethiere aus China," 1885, pp. 31-44; Schlosser, "Die fossilen Saugethiere Chinas," 1903, pp. 43-49

Koken's specimens were also probably collected from the caverns of Yunnan by dealers for von Richthofen;
his list is as follows (Koken, 1885, p. 33)

Proboscidia.
1. Mastodon perimensis var. sinensis Yiinnan.
2. Mastodon aff Pandionis
. Yiinnan.
*3. Stegodon Clijtii Shanghai ; oberer Hoangho (West-Kansu).
*4. Stegodon insignis Yunnan ; Szechuen.
5. Stegodon aff. bombifrons Yunnan.
Schon friiher beschriebene oder erwahnte Arten sind durch *ausgezeichnet.

The type fragment.s all appear as though they had been brought in by native collectors, for medicine dealers,
since they consist of broken teeth only; the same is true of the Schlosser collection. Consequently we regard the
specific and generic determinations made from these imperfect materials as of provisional value. The species of
Proboscidea contained in Schlosser's article of 1903, following Koken's review of 1885, are as follows:

Reference in
Present Memoir
Fokien, Kansu Stegodon orientalis Owen, 1870 = Stegodon insignis Falconer (fide
Koken, 1885, fide Schlosser) Stegodon orientalis
?Shanghai Stegodon sinensis Owen, 1870 = Stegodon clifti Falconer (fide Koken,
1885), = Elephas clifti (fide Lydekker, 1886) Stegodon sinensis
Stegodon bombifrons Falconer, 1846 = Stegodon aff. bombifrons (fide
Koken, 1885), = Elephas bombifrons (fide
Lydekker, 1886) IStegodon bombifrons
Rothe Thone Mastodon latidens ("lift, 1828 = Mastodon aff. latidens (fide Schlosser) IStegolophodon latidens
= Shansi
Rothhche Sande = Mastodon Lydekkeri Schlosser, 1903, related to Mastodon latidens
Tientsin, Honan, etc. (fide Schlosser, 1903), to Serridentinus (fide
Osborn) 1 Serridentinus lydekkeri
Mastodon perimensis var. sinensis Koken, 1885= Tetralophodon (Ly-
dekkeria) sinensis (fide Osborn) Tetralophodon (Lydekkeria) sinensis
Rothe Thone Mastodon pandionis Falconer (fide Koken, 1885) = Incertse sedis (fide
= Shansi Osborn) Incertx sedis

b) American Museum Discovery of Stegodon orientalis grangeri near the

Yangtze River, China
The first scientific party to collect fossils in situ in China, with records of the actual locality and geologic level,
was that of the American Museum, under Walter Granger, in 1920-1921.

In a preliminary notice of a collection secured during the winter of 1920-1921 by Dr. Granger of the Central
Asiatic Expedition, Matthew and Granger (1923) described the material as occurring in a series of pits or fissures
at the village of Yenchingkou in the vicinity of Wanhsien, province of Szechuan, about one hundred and forty
miles distant in an air line from Chungkingfoo, the type locality of Stegodon orientalis Owen. This Yenchingkou
material includes a fairly complete adult skull, two young skulls, a series of palates and lower jaws, and many
teeth, which Matthew and Granger figured and partly described (pp. 567-571, figs. 3-6). They remark (p. 567)
"Stegodon orientalis Owen. Schlosser regards this species as identical with *S. insignis of India, basing the reference
upon the fragmentary teeth described by Owen. Matsumoto regards it as distinct, upon the evidence of the
referred material which he describes and figures. The Yen-ching-kao material includes a fairly complete adult
skull, two young skulls, a series of palates and lower jaws and many teeth. It should enable us to estimate the
affinities of the species more exactly when it has been cleaned up and studied." They deferred further description
of this fine material to Osborn (see below, pp. 875-881).
 818 OSBORN: THE PROBOSCIDEA

Geologic Age of Stegodon orientalis and S. orientalis grangeri Types. —Matthew and Granger
(1923, pp. 563, 565) observe:

The Chinese fossil mammals described by Owen in 1870 [Footnote: '(iuar. Journ. Geol. Soc, London, XXVI, jjp. 417-4.36,
Pis. xxvii-xxix.'] came from 'a cave near the city of ( 'hung-king-foo in the province of Sze-chuan.' Chung-king is on the
Yang-tse-kiang above Wan-hsien, about one hundred and forty miles distant [southwest from the ( iranger locality of Ycnching-
kouj in an air line. Possibly the Chinese informants of Consul Swinhoe, who sent the fossils to Owen, misled him, uninten-
. . .

tionally or deliberately, as to the locality. Owen regarded the [Chungkingfoo] fauna as Pliocene and described the follow-
. . .

ing species:

[Chungkingfoo] [Yenchingkou]
Stegodon orientalis [type].
Parts of molars. [S. orientalis grangeri type
Rhinoceros sinensis [type].
Parts of 4 upper and 4 lower molars. R. sinensis ref.
Tapiriis sinensis [type]. Parts of 3 upper and 4 lower teeth. Tapirus sinensis ref.
Chalicotheriimi sinense [type]. Part of an upper molar. Chalicotherium sinense ref.
Hyaena sijiensis [type]. Canine, 2 premolars. Hyxna sinensis ref.]
Owen's descriptions and figures accord very well with some of the species in our collection [the Yenchingkou collection
of the .American Museum], so that we have referred them to his species, whether or not later investigation proves them to be
exact topotypes.

7. PLIOCENE TO PLEISTOCENE PROBOSCIDEA OF JAPAN

Discovery. —Matsumoto's discoveries and descriptions of the Japanese Proboscidea up to 1924 are recited in
more detail at the close of this chapter (pp. 901 to 909). His Elephas {Prostegodon, Parastegodon) aurorse, 1915,
1918, 1924, from a compari.son with Stegodon airdwana of Java, appears to belong to Stegodon aurorse rather than
to represent a distinct genus, i. e., Parastegodon. Matsumoto (1918, pp. 51, 52) gives the history of discovery of
Stegodon and Elephas in Japan, beginning with Leith Adams
Brauns (1883), Lydekker (1868), Naumann (1881),
(1886), Martin (1886 [1887]), Tokunaga (1906), Sato (1914), Kato (1914), Matsimioto (1915, 1918). The Japanese
species of Stegodon referred by Naumann to S. cliftii and insignis have been transferred by Matsimioto to
.S'.

Owen's Chinese species Stegodon sinensis and S. orientalis. Matsumoto remarks as to sexual characters (op. cil.,
1918, p. 52):

Thus, the present writer's opinion [Footnote: 'This Vol., p. 10.'] that, St. sinensis, as well a,s St. orientalis, is geologically
younger than ,S7. diSlii, evidently holds true also in the .Japanese specimens. One evidence noticeable is that the Stegodont
species are usually found in couples. For example, St. cliftii and bombifrons are found associated with each other from the Dhok
Pathan to the Tatrot horizon, St. ganesa and insignis from the Boulder Conglomerate horizon and the Lower Pleistocene of
Narbada, .S7. orientalis and sinensis from the Uppermost Pliocene to Lower Pleistocene of China and .Japan, and .S7. airdwana
and trigonocephalus from the Lower [to Middle] Pleistocene of Java. One may imagine the po.ssibility, that each couple of
species represent sexual dimorphism of one and the same species.

Matsumoto (1924, 1926, 1927) continues the history of discovery of the Stegodonts and Elephants of
Japan up to the year 1927, as set forth in detail in this chapter, and presents his recent views as to the phylugeny
of these animals as shown in figures 791 792, and 793. , To the fossil fauna of Japan he adds the following five re-
ferred and new species {op. cit., 1926.1, p. 1):

k Cautlcy. 4. ,S'. clifti Falconer & Cautley, ibid. [.Journ. Ceol. Soc. Tokyo, Vol. XX.Xl], 1924
Cenus Stegodon Falconer
|1!)24.:^|, .\kira-mura, Kage District, Province of Isc— ijossibly Plaisancian-.\stian. 5. .S. sinensis Owen, ibid., 1924, p.
1). :327.
328. Island of Sliodo (Shodo-siiima or Shozu-siiima), Inland 8ea —
Milazzian-Tyrrhenian. 6. .S. orientalis Owen, ibid., 1924, p.
330. (Also as S. bombifrons, ibid., j). 329.) Nagahama, Minato Town, I'rovince of Kazusa; Togane Town, same province—
Cakbrian. Riuge, Ikadachi-mura, Province of Omi~Calabrian or possibly Cromerian. Okimisome, I'be Coal-Field, Province
of —
Suwo Cromerian. 7. S. orientalis shddoensi.s, nov., ibid., ]). 333. Islands of Mitsugo (.\Iitsugo-shima) and Island of Siiodo,
Inland Sea; off Nagasaki, Eastern Sea— Milazzian-Tyrrhenian. Kashiwazaki, Province of Echigo (?this form).

Of tlie al)()V(' the genus Parastegodon Matsumoto, 1924, is regardeil in tiie i)re.sent Memoir as belonging to
the genus Archidiskodon or to a progressive Stegodon.
 —

THE STEGODONTINiE: HISTORY 819

8. PHYLOGENETIC DISCUSSION OF THE THIRTY DESCRIBED SPECIES OF STEGODONTS

AND STEGOLOPHODONTS
Double or Multiple Phyla of the Stegodonts. Schlosser, 1903. — Schlosser (1903, p. 191), in his

revision of the Stegodonts of China, doubtfully suggested that west European species, originally described as

Mastodon turicensis [
= il/. M. pyrenaicus, etc.], separated as Zygolophodon by Vacek in 1877 and
tapiroides, (1926)

by Osborn as Turichis (M. turicensis, M. tapiroides) and Zygolophodon {M. pyrenaicvs), may have given rise to

such Upper Miocene [Middle Pliocene] Stegodonts of India as Stegolophodon cautleyi.

Schlosser was also the first to suggest {op. cit., p. 191) that Mastodon turicensis [=Turicius tapiroides] of
the Lower Miocene of Europe may have given rise to the Mastodon [=Stegolophodon] latidens of the Lower Pliocene^
of Asia from which in turn sprang off the true Stegodonts, such as Stegodon insignis.

More in detail, Schlosser, who was the first to discuss the double phylogeny of the Stegodonts {op. cit., p.

206), separated them into two divisions as follows:

Mastodon latidens Clift Slegodon ganesa Falconer and Cautley

Mastodon cautleyi Lydekker Stegodon insignis Falconer and Cautley
Stegodon cliftii Falconer and Cautley
Stegodon bombifrons Falconer and Cautley

Two Phyla Suggested by Pilgrim, 1913. —The second to discuss the phylogeny of the Stegodonts was
Pilgrim. The foundations of a diphyletic arrangement of the Stegodonts were laid by his observation (1913,

JjOWEK Miocene. Turicius (4,5) compared with

Trilophodon (2)

Fig. 689. Left third inferior molar, I.M3. Comparison of

Turicius turicensis [= tapiroides] (upper 4, .5) with Trilophodon
pontileviensis (lower 2), one-half natural .size. After Mayet,
1908, PI. XI, figs. 2, 4, and 5.

These molar teeth exhibit the profound difference between

the zygolophodont molar (above), resembling that of a primitive
Stegodont with uninterruiited, widely open valley.s, and the
bunolophodont molar (below), in which the valleys are closed by
the central collides.

(Upper) Figs. 4, 5. "Mastodon turicensis [ = T. tapiroides].

Pontlcvoy. Dcrniere molaire infcrieure. Paris Museum.
—
Grandeur naturolle." Fig. o. "Id. vue d'en liaut." See also
figure 138C, CI.

(Lower) "Mastodon angustidens. l''aluii de Pont-

l''ig. 2.-
Communiciuee par M. .Joan
levoy. Dernicre molaire supcrieure.
de Bodard, Pontlevoy. Grandeur naturelle." = Trilophodon [

pontileviensis.]

pp. 293, 294) that Mastodon [

= Stegolophodon] latidens occurs in the same Lower [Middle] Pliocene beds with
Stegodon bombifrons. Pilgrim's discussion of the phylogeny of the Stegodonts may be paraphrased as follows:

(a) In the lower deposits of Perim Island there appears a species Mastodon cautleyi representing a line of
evolution which in all its earlier stages is entirely unknown in Europe [see Turicius (?) nnd Zygolophodon (?)].

The lectotype of M. cautleyi is a last upper molar, figured by Lydekker in 1886 (1886.1).
'[See note on page 824 below where Stegolophodon latidens is given as of Lower Pleistocene age. — Editor.]
 ;

820 OSBORN: THE PROBOSCIDEA

(b) In the higher Perim levels, as well as in Lower [Middle] Pliocene age, is found
the Middle Siwaliks of
a further development of this type in the species Mastodon latidens, with an increased number of ridges. The
larger form of M. latidens is to be regarded as a direct descendant of Mastodon cautleyi. The last species of masto-

dont to be referred to this Une is represented by a tooth from Lehri, for which Pilgrim proposed the name Mastodon
[Stegolophodon] stegodontoides.

(c) The type tooth of M. stegodontoides, of which the horizon is uncertain but may possibly be Upper Siwalik,
is distinguished from that of M. latidens by the almost entire absence of accessory columns [conules of Osborn]
M. stegodontoides, like M. latidens, carries on each of its ridges four columns [
= conelets of Osborn], while the
anterior ridges of the next higher stage, Stegodon elephantoides {=cliftii),^ carries nine or ten mamillse [=conelets of
Osborn].

(d) So close is M. stegodontoides to S. elephantoides ( = cliftii) that it is hard to separate the two genera.

(e) The true Stegodon type represented by S. bombifrons appears in the Lower [Middle] Pliocene, Dhok
Patlian horizon, before the extinction of the Mastodon = Stegolophodon]
[ cautleyi-latidens-stegodontoides phylum
which is parallel with it.

Pilgrim accordingly divides the Stegodon tinse into two generic phyla: to the first he applies the name Masto-
don [
= Stegolophodon], to the second he applies the name Stegodon. This may be graphically represented as
follows:

Mastodon Series = Stegolophodon] [ Stegodon Series = Stegodon] [

Upper Pliocene Mastodon stegodontoides Stegodon bombifrons [Middle Pliocene]

Lower Pliocene' Mastodon latidens Stegodon cliftii = Stegodon elephantoides
[ {- cliftii),
Upper Miocene [Middle Pliocene] Mastodon cautleyi Lower Pliocene]'

SCHLESINGER, 1917. — Schlesinger recognized the distinctness of Stegolophodon, basing his type on Mastodon
latidens Clift.

Matsumoto, 1922. —The fact that Mastodon latidens appears in the same Lower [Middle] Pliocene geologic
horizon as S. bombifrons is very significant; it tends to support the idea that the Stegodonts were diphyletic.
This idea is perhaps carried a step further by Matsumoto who writes (letter, Nov. 20, 1922) : "In my report just
in preparation on the Japanese 'Mastodonts,' I follow you to refer 'Mastodon' latidens to the genus Stegodon
creating however a subgenus Prostegodon for it. Prostegodon is the primitive representative of the Stegodon-
phylum, representing half bunomastodontine and half stegodontine dental characters. Schlosser's opinion, that
Prostegodon might be ? a descendant of 'Mastodon' turicens, does not appear to be correct at all."

Osborn, 1923. — Osborn (1923.601, ji. 2) erroneously adopted the generic name Prostegodon Matsumoto,
based on the genotypic species Mastodon latideyis Clift. Prostegodon, however, is preoccupied by Stegolophodon
Schlesinger, 1917. Thus the diphyletic arrangement of the Asiatic and European species and genera would appear
as follows:

Stegolophodon stegodontoides Stegodon ganesa

Stegolophodon cautleyi Stegodon insignis
Stegolophodon latidens Stegodon bombifrons
Turicius (?) sp. Stegodon elephantoides ( = cliftii)
Zygolophodon (?) .sp.

Matsumoto, 1924, 1920. -A more recent stej) is lliat of Mntsutiiol-o in a dingrain enii)odied in a, letter
dated Sendai, November 20, 1924, in which the polyphyletic Stegodontinae are tlivitletl theoretically into five
'[See footnote on page 824 regarding the Lower Pleistocene age of Stegodon elephantoides (=cliftii). — Editor.]
 ,

THESTEGODONTIN^: HISTORY 821

phyla (Fig. 792), and Trilophodon [

= Zygolophodon] pyrenaicus (Fig. 791) is placed as ancestral to Tetralophodon
falconeri, Parastegodon [
= Stegolophodon] lalidens, P. [
= S.] stegodontoides, and Stegodon.

In other words: (1) The type of Mastodon {

= Zygolophodon\ pyrenaicus of the Middle Miocene or its an-
cestors Zygolophodon pyrenaicus aurelianensis Osborn of the Lower Miocene of western Europe may be ancestral
to the Stegolophodon and Stegodon species of southern Asia.

Ji^: S 00.
Rg:3.

Fig. 230. — l)i.Tniijre niobire inrunciiie du yiustodon luricenms (tujnroidei;)

aux 2/5 de grandeur. — Miocuiic moycii dc Siniorrc, C.crs. (D'aprus Lartet.)

TuRicius (C) AND Stegolophodon (A, B) Form of Grinding Teeth

Compare with Stegodon Molars (Fig. 686)

Fig. 690. (C) Type

third right inferior molar, r.Ma, of Turicius turicensis simorrensis Osborn, 1926, erroneously determined by Lartet (1859, PI. xv, fig. 3)
as Mastodon lapiraides, two-fifths natural size. Upper Middle Miocene of Simorre. After Gaudry, 1878, p. 174, fig. 230. Reversed in drawing. See also
Vol. I, pp. 207 and 220 of the present Memoir.

Cotype and lectotype of Mastodon [=:Stegohphodon] cautleyi Lydekker, 1886.

A (Cotype). First superior molar of the left side, l.M', one-third natural size. After Falconer and Cautlcy, 1846 [1847, PI. XL, fig-s. 3, 3a], as "Mastodon
lalidens." Length 4 inches, width 2.3 inches. Brit. Mus. M.2817. Cast Amer. Mus. 26965. Perim Island. See Lydekker, 1886.1, p.xv, fig. 5.

B (Lectotype). Third superior true molar of the left side, l.M', one-third natural size. After Falconer and Cautlcy, 1846 [1847, PI. xxxi, figs. 6, 6al as
"Mastodon latidcns." Length 8.5 inches, width 4.5 inches. Brit. Mus. M.2705. Cast Amer. Mus. 26966. Perim Island. See also Lydekker, 1886.1, p. xv, fig.
6, and 1886.2, p. 73, fig. 18. Same as figures 141 and 142 of Volume I of the present Memoir.

Matsumoto held (1924)

(2) that the Stegodonts are not merely diphyletic (e. g., Stegolophodon and Stegodon),
but polyphyletic by subdivision of the species of Stegodon into five distinct hnes of descent, as clearly displayed in
figure 792.

Osborn, 1927. —The present author (1927) takes the more conservative view that there are certainly two
distinct phyla, namely: (a) Stegolophodon cautlexji, S. lalidens, S. stegodontoides, and (b) Stegodon elephantoides
{ = cliftii) to S. airdwana. The latter appear to present a progressive series in the increasing number of conelets

and ridge-crests or lophs, but without very marked divergence, as shown in the geologic succession table (Table

III, p. 814).
822 OSBORN: THE PROBOSCIDEA

Suggested European-Asiatic Origin and Migration of the Primitive Stegodonts

The suggestion by Sehlosser (1903), rejected by Matsuinoto (1922), that the species of StegolopJwdon may
1)0 derived from mastodonts related to the Turicius or Zyyolophodon of western Europe, should not be lightly
dismissed. These west European animals have been grouped in the genus Zygolophodon by Vacck (1877), based
on the three species Mastodon borsoni, M. turicensis, and M. tapiroides ( = M. pyrenaicus). The genus Zygolo-
phodon embraces a type molar tooth fundamentally homologous and analogous, as shown in the accompanying
comparison of the lectotype of Mastodon [
= StegolopJwdon] cautleyi from the Middle Pliocene of Perim Island,
with a third inferior molar erroneously referred by Lartet (1859) to M. tapiroides [
= type of Turicius turicensis
simorrensis Osborn, 1926— Fig. 690] from the Middle Miocene of Simorre. Molars of the Turicius turicensis

[
= tapiroides] type occur in the Lower Miocene, Burdigalian, of the Falun de Pontlevoy, contemporaneous with
tlie referred Trilophodon angustidens [
= T. pontileviensis —Fig. 689, 2].

Actual relationship to primitive Stegodonts [Stegolopiiodontina^] of Asia is rei)rescnted by the Mastodon

{Bunolophodon) longirostre Kaup forma suhlatidens of Schlesinger from the Pliocene of Teschen (Schlesien),
Austria (Fig. 722). A striking analogy to the Stegodon type is seen in the molar teeth referred by von Meyer to
M. [Turicius] turicensis from the lignites of Elgg and Kapfnach, and finally in the strictly stegodont Mastodon [T.]

virgatidens of von Meyer (Fig. 168). The only way to test this theory, however, is to place models or casts of the
teeth of Zygolophodon, of Turicius, and of Stegolophodon side by side to see whether they compare in close detail, in

which case the genus Zygolophodon Vacek would replace the genus Stegolophodon Schlesinger.

Conclusion: Probable African-European-Asiatic Origin and Migration of the

Primitive Stegodonts
Osl:)orn, 1927' : If, as now appears probable, (1) the Trilophodon. pliylum first arrived in southern Europe and
migrated eastward into India, (2) it is also probable that certain primitive species of the forest-living Zygolopho-
don or Turicius phyla gave ri.se in Lower Miocene time to forest-living animals which spread into the forests of
southern Asia and developed into the Stegodon series, as first adumbrated by Schlosser (1903).

The European species actually resembling these animals is the Mastodon (Bunolophodon) longirostre Kaup
forma sublatidens Schlesinger from Schlesien, Austria (Fig. 722), as described below.

II. TYPE REVISION OF THE SPECIES IN ORDER OF ORIGINAL DISCOVERY

AND DESCRIPTION
In the previous Section I of the present Chapter XIV we have discussed the habits and general cluiracters, the
lidge formulae, the geologic order, the history of discovery, the principles of type revision of the species, the
Stegodonts of China, Japan, and the East Indies, the phylogcnetic succession, and, finally, the probable origin of

the Stegodontinse [and Stegolophodontinse] in western Europe, and hence more remotely in Afri(^a.

We now pa.ss in Section II to the very complicated subject of the type revision of the species on tlu> jjriiiciples

enumerated above (p. 816). For this purpose we will review the thirty species described in I he y{>ars between
1828 and 1936 in the order of their description, quoting extensively from the original type descriptions and re-
producing every available type or lectotype figure dir(M;(ly after the oiiginal author.

'[Cami>arc Vol. I, i>ii. 19-3, H)7, iilso Pis. ii lo iv.— Editor.)

 THESTEGODONTIN^: HISTORY 823

Fig. 691. Geograpliic distribution of the iwiiicipal species of Stcgolophodon and Slcgodon. Tlie white dots within tlie black areas represent the
approximate localitieswhere the types of these thirty species were discovered. Numbers 1, 9, 14, 16, 18, 19 and 30 are Stegolophodonts (see also Fig. 1228).
The white crosses represent referred specimens.

STEGODONTS [AND STEGOLOPHODONTS] IN ORDER OF THEIR DISCOVERY AND DESCRIPTION

See Figure 691

1.
 \ 56

——^ ^ i'alay

Fio. 692. FossiL-BEAKiNO Horizons along the Irkawaddv River, Burma

(l.<-ft) Map illustrating J.
Crawfurd's journey to Ava and Martaban in the years 1826 and 1827. After William Clift, 1828, Pi. xi.iv. (UiKlil) M:i|> illus-
trating the explorations of Barnum Brown for the American Museum
of Natural History in the year 1923. Based on the ofh.ial Indimi Survey Cazetteer.
3 -3. Ava, upper level.s of the Irravvaddy .Series (600 feet in thiekness) = Upper Pliocene, Upper .Siwaliks of India, containing SOijoilim binminicus Usl.orn
and undetermmed Hos. (Lower Pleistocene (cf. Colbert, chap. XXII,
pp. H.'.O, 14.51 of the present Memoir.-Kditor.]
2. Pondaung Clays (.50 feet in thickness) = Upper Eocene, containing the suillinc Anthracotheriida-; also of the Order Peris.sodactyla: Fam. Titano-
tl.enidic. Gen. Sivatilarwps, sp. ,S. cotlen, S. hirmanicum, S. rugosidrns, Gen. Eotitanotherium, sp. /?. lahirii; Fam. Amynodontiila-, Gen. Paramynodon, sp.
/ coitcn, P. birmaniciis; Fam. Tapiridic = I^phiodontidffl, Gen. Indolophus, sp.
. [ /. guplai, Gen. Chaamothmiim, sp. C. hirmanicum.. [For additional members
of the Pondaung fauna, see Colbert, 1938.1, pp. 2.).)-.398. -Editor.)
1. Yenangyaung (2.50 miles south of Ava), lowest levels of the Irrawaddy Series (estimated at 1.500 feet in thickruss), base of the
Middle Siwaliks, eon-
tanung Slrgohphodon Uitidem type, Skgodon deplmntoidcs type, S. cli/lii type [SIcgodon clephanloidcs (^cliflii)].

(Note by Edwin H. Colbert:— The total thi<knes.s of the Irrawaddy Scries is estimated at 5,000 foot, of which lie upper levels onlv (about 000 fec-t), so
I

far as known, are mammal bearing and are now reganled by recent geologists and
paheonlologists as of T-ower I'leislocerLe age, Stamp (1022, pp. WT, IDS),
for example, has shown that .Uaslndon [Slrgnhplwdo,,] Inlidnm
and llippopolamvs irrmmlicu.'i are probably limited to the upper Irrawaddy l)cds (Lower Pleisto-
cene). He considers that Pilgrim erred (1910,
p. 190) in pl.icing the.se species in the lower Irrawad.ly fauna (cf, Colbert, Chap. XXII, pp. 1.50, 14.51, of the1

present Memoir). Colbert places all the probosci<Ieans discovered

in Burma up to the present time, namely, SUgolophodon hl.idrns, Slajodoii dcplmnloidrs, S.
cleplianloidrsi^cliflu). and N. birmmiicu.',. in the upper Irrawad.ly .Series,
thus it-ssigning them to the I,ower Pleistocene. The lowest levels of the Series are
considered of Middle to Ui.pcr Pliocene .age. Likewise the Pondaung
clays are of a total thiekness of 6,.500 feet, the mammal-bearing portion apparently not
it j
exceeding oO feet.— Editor.]
524
 :

THE STEGODONTIN^: HISTORY 825

1. THE FIRST TWO STEGODONTS, DISCOVERED IN BURMA, 1828

—
Geology, Irrawaddy River, Burma. In the years 1826 and 1827, J. Crawfurd, F.R.S., while on an em-
bassy to Ava, Burma, discovered an extensive deposit of organic remains in that unknown and distant region.
On the Irrawaddy River, 250 miles below Ava, a gravel and sand deposit contained fossil bones, as mapped and
described by Buckland (1828). This is the type region of Mastodon latidens Clift, 1828, and of Mastodon elephant-
aides Clift, 1828, also of "Elephas diftti" Falconer and Cautley, 1846. According to Buckland (1828, p. 378) the
exposure is an extensive one:

These plants were found most abundantly in the same region with the fossil bones, but occur also along nearly the whole
course of the Irawadi from Ava to Prome. They were principally collected from a tract of country (Footnote: 'See annexed
map, Plate xliv.'] extending over a square of more than twenty miles on the east bank of the Irawadi, near the town of Wetma-
sut, about half-way between Ava and Prome, between lat. 20° and 21° N. The occurrence of bones was most abundant in
a small .space near the centre of this district, occupying about one third of the above-named area, the svuface of which is com-
I)osed chietiy of barren sand hills mixed with gravel; beneath these are strata containing shells and lignite, thnnigh which they
sink wells about two hundred feet to collect petroleum.

This indicates that the cotype specimens came from lower and higher geologic levels, a fact not realized in

Buckland's paper, nor in subsequent descriptions excepting those of Pilgrim.' Referring to Cliffs paper of 1828,
Buckland mentions (p. 380)

. . . two new and strongly characterized species, one of which, from its approximation to the elephant in the structure of the
teeth, Mr. Clift proposes to designate by the name of Mastodon elephantoides: to the other he has given the name of Mnstodon
latidens.

Lydekker (1886.2, p. 81) states that the type of E. cliftii, l.M\ "was obtained near Yenankhoung, on the left
bank of the Irawadi in Upper Burma, by Crawfurd in 1826, and is preserved in the Museum of the Geological
"2
Society.

D. N. Wadia (1919, p. 213) includes this Burmese [Yenangyaung] deposit in the "Irrawaddy system"
[series], which combines marine and fluviatile strata. The upper part = 2,000 feet), composed of "sands and clays(

with abundance of fossil wood and mammals," is of fluviatile origin and corresponds to the Manchhars of Balu-
chistan and to the Siwaliks of the sub-Himalayas.

Barnum Brown visited this region in 1923 for the American Museum of Natural History and prepared a new
map (Fig. 692, right) of this clas.sic collecting ground which should be compared with Crawfurd's original map
(Fig. 692, left). The three geologic levels, discovered along the Irrawaddy River, Burma, are shown in these maps.

HISTORY OF DISCOVERY OF BURMESE STEGODONTS

Clift, 1828. —The first proboscidean described from Burma was Mastodon latidens Clift (1828, p. 371),

the second species was Mastodon elephantoides Clift (1828, p. 372), both species founded on excellent cotypes
with the excellent figures which are reproduced herewith (Figs. 693, 694, 695, 696).

Mastodon latidens. —The cotypes of M. latidens (Figs. 693, 694) were found in 1826 on the Irrawaddy
River, 250 miles below Ava, near Yenangyaung, Burma. Cliffs figured specimens include: Plate xxxvi, upper
jaw (palate with anterior molar teeth, M^ much worn), which rightly (Osborn) belongs to Mastodon {
= Stegodon]
elephantoides; Plate xxxvii, upper molar teeth of the right side, M^, M^ (a younger animal), also Plate xxxviii,
fig. 1, anterior part of the lower jaw; these certainly are the cotypes, because they conform with Clift's descrip-
tion — all from the left bank of the Irrawaddy.

'[See this Volume, Chapter XXII, p. 1450, by EJwin H. Colbert.— Editor.]

2[Now in British Museum (Brit. Mus. M. 10520)].
 FOUR CLASSIC STEGODONT AND STEGOLOPHODONT TYPES OF CLIFT AND FALCONER AND CAUTLEY

Lectotype Palate of Stegolophodon latidens

693 (left). Lectotype palate (perspective of r.M') of
Fig.
Mastodon latidens Clift, 1828, PL xxxvii, fig. 1, with r.M^"' in
situ, about one-third natural size. Inverted to show natural
position of molars. From near Yenangyaung, Burma.
Observe beginnings of binary fission of cones in tritoloph.
.•\n accurate scale drawing of these teeth is shown in figure 71()

from a cast (Amer. Mus. 21978).

Fig. 093

CoTTPE Third Inferior Mol.^r of Steoolophodon latidens ^

Fig. 694 (right). Cotype third right inferior molar, r.Ma, of Mastodnu
latidens Clift, 1828, Pi. xxxviii, fig. 1, one-third natural size. Jaw omitted.
From near Yenangyaung, Burma.

Fig. 694

CoTYPE OF Stegodon elephantoides Clift (=cliftii Falconer)

Fig. 695 (left). Type first left superior molar, l.M', of Elephas cliftii Falconer and Cautley,
1846, after photograph of cast Amer. Mus. Warren Coll. 10382, one-half natural size. Inverted
to sliow the molar in natural position. From near Yenangyaung, Burma. See also figure
683. Compare Chft, 1828, PI. xxxix, fig. 6. Original in Briti.sh Museum (Natural History)
M. 10520.

Fig. 695

Lectotype Lower Jaw of Stegodon

elephantoides
Fig. 690 (right). Lectotype second
and tliird left inferior molars, I.M3-2, of
Mastodon lU pliantoides Clift, 1828.
PI. xxxviii, fig. 2, one-third natural
size. From M<';ir '^'enangyaurig, Hurma,

J^nidtj: fy iT^WirwwW
n.cUftU esju^iA,}

Fig. 696
826
 :

THE STEGODONTINiE: HISTORY 827

Mastodon latidens 370).— On comparing the teeth of our Mastodon latidens witli those of the Mastodon of
(op. cit., i>.

the < »hi() (M. Riganteum), we hnd the elevate<l points or ridges in the tooth of the former more numerous, le.ss distant, and
shall
the interstices less deeji tlian in those of the latter; in short, we shall obser\e that the teeth begin to assume the apiiearance of
those of the elephant. )n advancing to Mastodon elephantoides, we shall find all these features of similarity more strongly
(

—
developed; the points and ridges are still more numerous, and the structure, wei'e it not for the absence of crusta petrosa,
becomes almost that of the tooth of the elephant.

Mastodon elephantoides.— This species (Fig. 696) was found in the same locality as the cotypes of M.
latidem. Cliffs figiu-ed specimens include: Plate xxxviii, fig. 2 (lower jaw with M,, M;i in situ); Plate xxxvi
(palate figured by CUift as M. latidens), and Plate xxxix, fig. 6 (a first superior molar, l.M') afterward made the
type of Elephas cliftn by Falconer and Cautley, referred in the present Memoir to Stegodon elephantoides {=cliftii).

Mastodon elephantoides (op. cit., p. 372).— The tooth [Fig. 690], which is eleven inches long and three inches and a half
broad, has no less than ten denticules [i.e., ridges], and each of the.se denticules is mammillated with small points; five being
the smallest number, and eight the greatest on any one <lenticide. In front of this beautiful tooth we have a remnant of the
preceding one, . . .

Osborn, 1924: Pending final revision, which can only be made by a ree.xamination of the specimens and the
localities from which they came, we may designate these Stegodonts from Burma as follows:

Mastodon = Stegoloi)liu(hm]
[ latidens (
'lift, 1828, founded upon a palate from near Yenangyaung, Burma, with five and a half
ridge-crests in the third superior molar, four and a half to five in the second superior
molar, and with four to five mamillse (or conelets) on each crest. See (
'lift, 1828,
PI. XXXVII, fig. 1 (Fig. 693 of the present Memoir).

Also a lower jaw (.see Clift, PI. xxx\iii, fig. 1), r.Ms with seven ridge-crests
(Fig. 694 of the present Memoir).

Madodon [=,Slegodo)i] dephanloides Clift, 1828, founded upon a third inferior molar of the left side, I.M3, from near
Yenangyaung, Burma, with ten ridge-crests, five to eight conelets on each. See
('lift, 1828, PI. XXXVIII, fig. 2 CFig. 696 of the present Memoir).

Also a palate showing r.'SV- and l.M'-, with six and a quarter ridge-crests (see
Clift. 1828, PI. xxxvi), erroneously marked "Upper Jaw of Mastodon latidens."
(Not figured in the present Memoir.)
Stegodon elephnutoides {=diftii) Also a superior molar, l.M', with six and a quarter ridge-crests, from near
left first
Yenangyaung, Burma. See Clift, 1828, PI. xxxix, fig. 6, marked "Upper molar of
M. Elephantoides." Afterward made the type of Elephas cliftii by Falconer and
Cautley. (Figs. 683, 695, of the present Memoir.)

Original Cotypes of Mastodon latidens Clift (Figs. 693, 694)

First Species. — Cliffs designation (1828) of Mastodon latidens as including a lower jaw (PI. xxxviii, fig.

1 — Fig. 694 of the present Memoir) and a palate containing M^, M^ (PL xxxvii, fig. 1 — Fig. 693 of the present
Memoir) establishes these specimens as the cotypes. The same superior teeth, r.M", r.M^, were sectioned and
figured by Falconer and Cautley (1846, p. 48 [1845, PI. iii, fig. 8])

The shows five principal ridges with a posterior talon ridge and a subordinate ridge in front. The ridges are
last tooth
transverse, and divided by a longitudinal cleft into two pairs of princijjal points without intermediate mammillae in the hollows.
The enamel is very thick, and the cement is reduced to a thin layer which is only observable in the bottom of the hollows. . . .

The anterior tooth [M'^] had been a long time in use, and the ridges are nearly all worn out. They were four in number, in this
as well as in the two teeth which preceded it in the jaw. We believe this to be a small or dwarf variety of M. latidens. . . .

(Clift, 1828, p. 371) Dentition. : —

Each tooth of the lower jaw [Fig, 694] consists of seven denticules, which are elevated,
rounded, and mammillated: the mammillae being from three to four in number. The dentition both in this s])ecies and in M.
elephantoides, \'ery much resembles that of the ele])hant. We have the molar tooth gradually protruded forward, and rising
as the fangs are added, according to the demand made by the abrasion of the exposed crown, and the consequent absorption
of the anterior fang; the posterior part of the tooth not having cut the gum, while the anterior portion is completely worn away.
The relics of the preceding tooth, the place of which the tooth in use was progressively supplying, are plainly to be seen [Foot-
note: 'See Plate xxxvii. fig. 1. Plate xxxviii. fig. 2.'].
 —

828 OSBORN: THE PROBOSCIDEA

According to the above, the ridge formula is as follows:

Ridge formula of Mastodon [

= Stegolophodmi] latidens: Dp 4| M ly M 2t M 3-,^.

Osborn, 1927: The above is the main ridge formula given by Falconer, omitting the anterior and posterior
talon ridges (compare tables above, Section I, No. 3, and below Section IV of this chapter, also full description).

Original Cotypes of Mastodon elephantoides Clift (Fig. 696)

Second Species. — The second species is Mastodon elephantoides Clift.

Clift, 1828, p. 372. Mastodon elephantoides. —

M. dentibus molaribus latis, denticulis numerosis, compressis. This
species must have been smaller than the last; and though we have one fine example of the lower jaw, showing the tooth in the
highest degree of perfection, that is the onlj^ portion of the animal from which we can safely draw any inference as to its struc-
ture and habits. The tooth, which is eleven inches long and three inches and a half broad, has no less than ten denticules
[i. e., ridge-crests], and each of these denticules is mammillated with small points [conelets]; five being the smallest number,
and eight the greatest on any one denticule. The denticules of the tooth are much more compressed than those in the
. . .

species last described; they are closer together [Footnote: 'Eight denticules of M. elephantoides occupy the same space as five
denticules of M. latidens.'J, and the enamel appears to be not so thick. They form a series of plates mucronated with small
points. There is no apparent commissure, neither is there any central depression on the contrary, the plates rather rise in the
:

middle.

Clift uses the word denticules in the sense of ridges, or crests, or lophs; subsequently Falconer uses the word
denticles in the sense of "mammillae" or conelets. Osborn introduces the word conelets, because these small,
roimded "mammillae" appear on the summits of the primary cones, as e.xplained above (p. 812).
(Clift, 1828, "Explanation of Plates," PI. xxxix, fig. 6): "Upper molar tooth of Mastodon elephantoides."

[No mention of this tooth is found in the text.] As reproduced herewith (Fig. 695) this molar corresponds closely
in scale and structure with Cliffs type of M. elephantoides (Fig. 696).

Osborn, 1927 : From these two teeth, figured and described together by Clift, 1828, not improbably represent-
ing the same species, also from the palate with r.M^ and l.M^ in situ (Chft's figure, PL xxxvi), the following ridge

formula may be written:

Ridge formula of Mastodon [

= Stegodon] elephantoides: M 1^ M2^ M3ro.
The above ridge formula is similar to that of Stegodon bombifrons and much more primitive than that of S.
insignis-ganesa. Consequently Mastodon elephantoides Clift, 1828, of Lower Pliocene age,' based upon two figured
specimens (Figs. 695, 696), appears to be well established as the second species of Stegodon described from Burma
in 1828. The name elephantoides was dropped, however, by Falconer and Cautley in 1846 and a new name, Elephas

cliftii, was apphed to the second specimen (Fig. 695 of the present Memoir) figured by Clift, PI. xxxix, fig. 6.

Species M. elephantoides Dropped by Falconer and Lydekker. — In 1846, Falconer and Cautley
erroneously alleged that Clift had confused the remains of the two species Mastodon latidens and M. elephantoides

under the name M. elephantoides; they accordingly {op. cit., p. 47) dropped the name elephantoides and proposed
anew name for the species of "transitional Mastodons," which had been partly confused with Mastodon latidens.

Thus to the Stegodonts with six ridges on the intermediate molars they gave the name of Elephas cliftii (Figs.

683, 695),and to the Stegodonts with a greater number of ridges the name of E. insignis (Fig. 697). In a sub-
sequent paper (Falconer, 1857, p. 314), these and other species appeared under the subgeneric name of Stegodon
Falconer. Lydekker (Palaeontologia Indica, 1880, pp. 256, 257) also set aside the prior specific name elephantoides
Clift, 1828, and made it a synonym of Stegodon cliftii Falc. and Caut., 1846. In subsequent literature (e. g..
Pilgrim) the Falconer-Lydekker usage is followed, i.e., M. elephantoides is dropped.

M. elephantoides Revived. —In the present Memoir the specific name elephantoides is revived by Osborn
for reasons given in the .systematic revisions above and below.

'[See note on page 824 above regarding the Lower Pleistocene age of Siegudon elephantoides. — Editor.]
 : .

THE STEGODONTIN.E: HISTORY 829

2. DISCOVERIES IN INDIA AND BURMA (1845, 1846)

Third Species. —The third species of Stegodont {Elephas insignis) was described (Falconer and Cautley,
1846, p. 37) as follows (see Fig. 697)

. the four anterior ridges being affected by wear, and the six posterior ridges entire,
. . The white mass in the centre [dentine] . . .

represents the body of ivory, which is projected upwards in ten angular lobes terminating in a sharp edge. The interspaces . . .

of the five posterior ridges of enamel are completely filled up by a mass of cement, or 'cortical,' much exceeding the enamel in
thickness; and in cjuantity iu nearly as great an amount of development as the ivory core of the ridge.

^v fi

Kj <•

Fig. 697. Lectotype and Cotype of Steqodon insignis

(Left) Lectotype superior molar, l.M^ of Elephas insignis Falconer and Cautley, 1846 [1845, PI.ii, fig. ffa], one-third natural

size. Probably Pinjor horizon, Upper Pliocene [Lower Pleistocene], Upper Siwaliks, India. Inverted to show natural position
of molar.

(Right) Elephas [ =Siegodon] insignis cotype. Anterior portion of a thii-d inferior molar, M3. After Falconer and Cautley,
op. cit, PI. II, fig. 6h.

Falconer distinguished this species as possessing a ridge formula of M3 --, with valleys between acute ridges
deeply filled with cement, as compared with Stegodon elephanioides which has ten less acute ridges without cement
(Fig. 696). Lectotype ridge formula (Fig. 697) oi Elephas [
= Stegodon] insignis: M3—
Fourth Species.— The fourth species of Stegodont (described by Falconer and Cautley, 1846, p. 45) was
named Elephas ganesa:
Fig. 7a, pi. 3., represents a section of the last upper molar of an undescribed Indian fossil species, named E. Ganesa, in this
work. The crown consists of ten principal ridge.s, with a subordinate 'talon' ridge in front and behind [i.e., M3 ^^"^°"'''
]. The
anterior seven ridges have their summits worn, the two in front being ground down to the common
base of ivory, the tooth
having been a considerable time in use. A small portion is broken off at the anterior end. The disposition and relative propor-
tions of the ivory, enamel, and cement, bear the closest resemblance to
those of the corresponding tooth of E. insignis (pi. 2, fig. 6a), and the
nimiber of ridges agrees. The section presents the same chevron-formed
character in the ridges, but the interspaces are narrower, the cement is
in less quantity, and the layer of enamel is thicker.

Lydekker observes (Lydekker, 1880.1, p. 268): . . . "these

latter teeth, however, cannot be distinguished from those named
Stegodon insignis, and as we shall see subsequently, it is only the
Lectotype of Stegodon ganesa
Fig. 698. Lectotype M' of Elephas Ganesa Falconer and adult skulls of these two very closely allied species that can be
Cautley, 1846, [1845, Pl. in, fig. 7a],
Probably Pinjor horizon, Upper Pliocene [Lower Pleistocene].
one-third natural size.
distinguished." This has led to the opinion that S. insignis may
Upper Siwaliks, India. represent a female and S. ganesa a male of the same species.
 : .

830 OSBORN: THE PROBOSCIDEA

The ridge formula of the lectotyi)e third superior molar of Stegodun ganesa (Fig. 698) appears to he practi-
cally tli(» same as that of the lectotyjie of ;S'. insignis (Fig. 697). Other more progressive ridges foruuihr are
shown in the comparative ridge formulae tables above and below in this Memoir.
Ridge formula (Fig. 698) of Elephas [
= Stegodon] ganesa: M3 '^'=^^^.

Fifth Species. —Falconer described (Falconer and (Jautley, 1846, p. 46) Elephas bombifrons, the fifth species

of Stegodont, as follows:

This species, of the distinctness of which we are assured, by possessing several crania containing perfect teeth, belongs to the
same group [Stegodon] as the two species last described. The crown is divitled into similar transverse ridges, composed of numer-
ous mammillse, which yield a corresponding chevron-shaped section, and the interspaces are occupied by a thick coat of cement;
but they differ, in being broader and less elevated, with more open hollows. The principal ridges of the last molar [IVP] do not
exceed eight in the upper jaw, and nine in the lower [M3]; while in E. insigniti they amount to ten in the former [AP], and
reach as many as thirteen in the latter [M3]. The last tooth of the upper jaw measures eleven inches in length, by four and
a half in width.

Falconer and Cautley (1846 [1847, PL xxviii]) assign to M'* nine ridges and a heel.

Lectotype ridge formula of Stegodon bombifrons: M 3f

Sixth Species. —A sixth species of Stegodont (Figs. 683 and 695) was erroneously proposed by Falconer and
Cautley in 1846 (1846, p. 47) under the name of 'Elephas clij'tii.' They selected the type of this species as

follows

CoTYrE OK Stuqodon bomhikhons

Cotypc skull of Elephas bombifrons
Fig. 699.
FiilconiTand Cautley, 1846 [1847, Pis. xxvii,
xxviiil, ont'-sixtli natural size. Brit. Mus.
M.2979; cast Amcr. Mus. Warren Coll. 101578.
From the Siwalik Hills, India, probably the Dhok
Pathan horizon, fpper Siwaliks, Middle Pliocene.
As a view of the third molars was not given in the
loctotyiJC of Lydekker (Falconer and Cautley,
op. cit., PI. XXVI, Brit. Mus. M.2978), the cotypc
skull is figured here.
 THESTEGODONTINiE: HISTORY 831

In our view, the tooth represented in pi. 39, fig. 6, of Mr. CUft's memoir in tlie Geological Transactions [Clift, 1828],
under the name of Mastodon Elephantoides, and the palate specimen represented in pi. 36 of the same memoir, under the
name of M. latidens, belong to this species.

This was an unfortunate error. As explained above, the tooth in PI. 39, fig. 6, of CHft's Memoir, measuring
155 mm. in length, 83 mm. in breadth, has a ridge formula of M — 1 ; the palate specimen (PI. 36), erroneously
identified by Clift as "Mastodon latidens,'' also has a ridge formula of (?)M 2—. Consequently they belong to
the same species, namely, Stegodon elephantoides, but to preserve the name cliftii, which runs all through the
previous literature, they are designated in the present Memoir as Stegodon elephantoides = cliftii). {

Stegodon elephantoides (= cliftii)

Fig. 700. Original type figure of Elephas cliftii Falconer Fig. 701. New figure of type of Elephas cliflii Falconer and
and Cautley, 1846, a first superior molar of tlie left side, l.M', Cautley, 1846, a first superior molar of the left side, l.M', de-
one-half natural size. From near Yenangyaung, Burm;i. scribed and figured as M. Elephantoides by Clift, 1828, PI. xxxix,
After Clift, 1828, PI. xxxix, fig. G, figured as an "Upper fig. Subsequently seli^cted by Falconer and Cautley as the ty]>e
6.
molar of Af. EUphntdoides." Original in the British Museum of Elephas cliftii. After i^hotograph of type cast (Amer. Mus.
(Brit.Mus. M. 10520); cast Amcr. Mus. Warren Coll. 10382. Warren Coll. 10382). One-half natural size. Observe that the
Inverted to show natural position. molar is here placed in its natural position. Original in the
British Museum (Brit. Mus. M. 10520).

3. THE STEGODONTS OF CHINA, INDIA, JAVA, THE PHILIPPINE ISLANDS, AUSTRIA,

JAPAN, AND BURMA
Seventh Species. —The seventh species of Stegodont described (Fig. 702) was the Stegodon sinensis of
Richard Owen (Owen, 1870, p. 417), alleged to be (p. 421) "from marly beds in the vicinity of Shanghai," China,
which has been mistakenly regarded by some authors as close to, or as a synonym of, Stegodon cliftii (Owen, op.

Fig. 702. Type r.Dp^ of Stegodon sinensis Owen, 1870, PI. xxvii, figs. 1, 2, natural size. From "marly beds
in the vicinity of Shanghai," China, of Upper Miocene [Lower Pliocene] age.
 832 OSBORN: THE PROBOSCIDEA

cit., p. 418; Brauns, 1883, p. 44). This synonymy is not borne out by careful comparison with the type figures,
because S. sinensis is nuich more primitive than ti. cliftii.

Eighth Species. —In the same paper Owen {op. cit., 1870, p. 421) describes Stegodon orientalis, the eighth
species of Stegodont (Fig. 703), "from a cave, near the city of Chung-king-foo, in the province of Sze-chuen,"
based upon molar fragments which he rightly observes "more resemble the teeth of Stegodon Cliftii, St. insignis,

and St. ganesa of Falconer than does the St. sinensis; and in the apparent quantity of coronal cement . . . as
well as in the evidence of a hinder talon . . . they are more like *S^. insignis than St. Cliftii."

Fig. 703. Type of Stegodon orienlalis Owen, figs. 1-4.

1870, PI. xxviii, "Portion of true molar" (figs. 1, 2) .anil "hind enj
of milk-molar" (figs. 3, 4). From near Chungkingfoo, China, probably of Lower Pleistocene age.

Ninth Species.— il/astodow cautleyi Lydekker, 1886 (1886.1), was founded on five teeth of th(> upper jaw, of
whicli wo s(>]oct as the type (Fig. 704) the third left upper true molar, from Perim Island, India, a tooth first figured
Fie, 18.
by Falconer and Cautley and referred to the species
Mastodon latidens Clift (PI. xxxi, figs. 6, 6a —see
Vol. I, fig. 142 of present Memoir, the caption of
which is erroneous— corrected in present c^hapter,

p. 821). The four cotypes, obviously smaller and

inucli more primitive teeth than the type of Mas-
todon latidens, are also from Perim Island and are
recorded, like the lectotype, from an older geologic;
horizon, namely. Upper Miocene [now (1935-1938)
regarded as Middle Pliocene, Dhok Pathan]. Ac-
cording to these specimens, the ridge formula of
Mastodon [
= Stegolophodon] cautleyi is about the
—
Mastodon cautleyi. The third left upper true molar, io an unworn condition
from the Siwaliks of Perim Island. J. The lower border of the figure is
;

same as that of Mastodon = Stegolophodon] [ latidens.

the inner border of the specimen.

Lectotype of Stegolophodon cadtleyi Mastodon = Stegolophodon]

[ cautleyi: M 2^ M 3^^^.
Fig. 704. Third left superior molar of Mastodon cautleyi Lydckkor,
selected as the type (sec Pilgrim, 1913, [). 294). Roprod\iwd
18S(i,

after Lydekker, ISSO

Tenth Species. — In 1887 Martin described
(1886.2, p. 73, fig. 18), one-half natural .size. ("Fossile Siiugethier-reste von Java und Jajjan")
 :

THE STEGODONTIN^: HISTORY 833

from Java a species to whicli he gave the name Stegodon trigonocephalus (Fig. 705), in reference to the triangular
shape of the head. The geologic level is regarded by Matsumoto as equivalent to the Lower Pleistocene, Boulder
Conglomerate beds of India. It may be of the same geologic age as the type of Pithecanthropus erectus. The
juvenile type does not admit of giving the mature ridge formula; the ridges are closely compressed, buried in
cement, and each ridge is surmounted with ten to twelve conelets, the exact number being indeterminable from
the figures.

Fig. 705. Type of Stegodon trigonocephalus Martin, 1887, immature skull, one-eighth natural size. I'Vom vicinity of Surakarta,
Java, probably of Lower Pleistocene age. (Right figures) Tab. ii, figs. 1, la; (left figure) Tab. iii, fig. 1.

Eleventh Species. —The eleventh species (Fig. 706) was described by Naumann in 1890 from Mindanao,
Philippine Islands, as Stegodon mindanensis. It was first referred to Stegodon trigonocephalus by Naumann in
1887, but subsequently was made the type of a distinct species. The compressed cement-covered ridges with
multiple conelets indicate a Lower to Middle Pleistocene stage of evolution, similar to that of S. trigonocephalus.

Twelfth Species. —A twelfth species, Stegodon airdxvana (Fig. 707), was described by Martin in 1890 from
Alas-Tuwa, Java, based upon a type jaw containing the right and left third inferior molar teeth, M3. In its

high ridge formula the author compared it to Stegodon insignis and S. ganesa. In a later paper by Janensch
(1911, 1). 187), he distinguished S. airdwana from both S. insignis and S. ganesa and wrote the ridge formula
as follows, according to the present writer's understanding of the Janensch system [cf. Table V]

Ridge formula of Stegodon airdwana: Dp 3 '-^ Dp 4 *'-^ M 1 '-^^^

M2 li-0-!.i
M3 14-1 1-1 2-1,4
15-1 3-(i •

This species is important because it is highly characteristic of the

Trinil Pithecanthropus erectus beds.
l. %.

Type or Stegodon (Archidiskodon?) mindanensi.s

Fig. 706. Type of Strgodon mindanensis
Thirteenth Species. — In 1908 Dubois described from the Trinil
Naumann, 1890, molar tooth. Mindanao, Philip- Kendeng-Schichten, Java, Stegodon ganesa javanicus, as a variety of S.
pine Islands, Lower to Middle Pleistocene age.
ganesa [which proves to be a synonym of either S. airdwana or S. tri-
.\rtor Naumann, 1887, Taf. i, figs. and 2, under the
1

name Stegodon trigonocephalus. gonocephalus (see p. 889 below)].

834 OSBORN: THE PROBOSCIDEA

Fourteenth Species.- — Very important is the Mastodon stegodontoides of Pilgrim, 1913, from Lehri, Punjab,
India (Fig. 708), wliich Pilgrim described as follows (p. 294):
The last species of Mastodon whicli can be referred to this line [the M. cautleyi-M. lalidens Hne] is the tooth from Ix-hri,
of which the horizon is uncertain but ma,y be possibly Upper Siwahk, figured by Lj^dekker in Pal. Ind. ser. 10, Vol. 1, Plate 39,
as .1/. lalidens but which is better recognized as a new species, owing to the ahnost entire absence of accessory columns, for

which I propose the name Mastodon stegodontoides. So close is this to Stegodon difti that it is hard to separate the two genera.
It will be seen that Mastodon stegodontoides carries on none of its ridges more than the usual four columns while anterior ridges
of Stegodon difti carry nine or ten mammillae.

Fifteenth Species. —Very progressive is the

Elephas {Prostegodon, Parastegodon) mirorx of Mat-
sumoto, 1915-1924, from Mt. Tomuro, Kaga, Japan
(Fig. 709), which Matsimioto first regarded as a stage

Type of Stegolophodon steqodontoides

I'^if!. 70S. Type r.M' of Maslodon slegodontoidcs Pilgrim, 1913. .\ftcr Lydek-
k(_T, ISSO, PL xxxix: "Maslodon The third right
(Tetralophodon) lalidens, Clift.
upper true molar: from Lehri, in the Punjab. The specimen is drawn of the
natural size, and is viewed from the inner [outer] side." Ind. Mus. A.86. Reduced
to one-half natural size. Provisionally placed in the Upper Pliocene, Pinjor
formation (.see Fig. 413, also PI. xiii).

^i^V^ffft

l''ig. 707. Type of SIfyodon Airawiiiid .Martin, 1890, Tab. i, figs.

Kcndcng-Schichtcn horizon (Pilhccanlhropus ereclus zone), Middle Type of Stegodon auror.*;

1, 2, lower jaw, one-fourth natural size. From .\la.s-Tuwa, Triiiil, .lava. Fig. 709. Type r.M^ of Elephas (Proslegodon, Parastcyodon) aurora-
Pleistocene. Matsumoto, 1918, PI. xx, figs. 1 and 3, one-half natural size.
 THESTEGODONTIN^: HISTORY 835

in Elephas close to the Upper Pliocene Elephas [

= Archidiskodon\ planifrons of India, but which he subsequently

made the genotype of a new genus Paraslegodon. knowledge of Stegodon airdwana, the present
With our fuller

species may be placed in the true Stegodon phylum, distinguished by cranial characters from species of the Ele-
Ijhantidse, as Stegodon aurorx.

Sixteenth Species.— The subspecies Mastodon {Bunolophodon) longirostre Kaup forma sublatidem Schle-

singer, 1917, from near Teschen (Schlesien), Austria (Fig. 710), appears to be very close to the Mastodon [
= Stego-
lophodon] stegodontoides of Pilgrim ; consequently it is removed to this genus, namely, Stegolophodon snblatidens.

—
Seventeenth Species. The subspecies Stegodon orientalis sliodoensis
of Matsumoto, 1924, named from the Island of Shodo, Inland Sea, is
regarded by its author as a descendant of S. orientalis Owen, of China,
contemporaneous with the referred S. insignis of the Middle [Upper]
Pleistocene, Narbada of India. Not figured in present Memoir.
Eighteenth Species. — The type of Stegolophodon nathotensis Osborn,
1929, consists of fragmentary molars (Fig. 724) collected by Barnum
Brown in 1922 in the Lower Chinji horizon, near Nathot, India.
Type of Stegolophodon sublatidens
Fig. 710. Tyjjc of Mastodon {Bunolophodon)
Kaup forma sublatidens Schk'singcr,
Nineteenth Species.— Somewhat more advanced than Stegolophodon
longirostre
1917, one-half natural size. Compare full legend cautleyi of the Upper Miocene [Middle Phocene] is the Stegolophodon caut-
below of figure 722.
leyi progressus Osborn, 1929 [of the Mio-Pliocene], the type of which (Amer.

Mus. 19446) is a complete cranium (Figs. 725, 727) collected at the summit of the Lower Chinji horizon, 2,000

feet above the base of the Lower Siwaliks, India. This is a young individual in which the molar ridge-crests

are intermediate in formula and pattern between the 'Mastodon' cautleyi of Lydekker and the 'A/.' latidens of

Clift.

Twentieth Species. — Somewhat more primitive and an-

cient than the Upper Pliocene [Lower Pleistocene] Stegodon in-
signis Falconer type is the subspecific stage Stegodon orientalis
grangeri Osborn, 1929 (Fig. 762) of the Yangtze River region
near Wanhsien, Sezchuan, 140 miles northeast of the type
locality of S. orientalis Owen. This subspecies is now very
fully knowai and is amply illustrated and defined in the present

Memoir. See type figures on pages 876, 877, 879, and 881 below.

Twenty-first Species. —Stegodon insignis birmanicus Os-

born, 1929 (Fig. 758), from the Up])cr Pliocene of Burma. See
tjqje figure on page 875.
—
Twenty-second Species. Stegodon pinjurensis Osborn,
1929, from the Lower Pleistocene of India, upper levels of the
Pin j or horizon (Figs. 711, 765, 767).

|The following species have been descril)ed since the author's Fig. 711. Photographic reproau.ti.m ..f type palate of

intensive review of the true Stegodonts, consequently the de- stegodon idnjormsis Oshom, \<m (Amer. Mus. i'j772), .•olle<-te,l

by Barnuni Brown three miles north of Siswan, India. Alx^iil

terminations of Professor Osborn cannot be given and the spe- one-eighth natural size. See figure 765 below.
o
1
o
 .

THE STEGODONTINiE: HISTORY 837

cies are listed here according to the nomenclature of the various authors, without comment, with the exception
of Parastegodoni kwantoensis and P. sugiyamai.

—
Twenty-third Species. The type of Stegodon bondolensis van der Maarel, 1932, is a fragment of a mandible
with what is inferred to be the third molar of each side, found at Bondol, Java (Fig. 782 of the present Memoir).

Twenty-fourth Species. —In 1933 von Koenigswald described from Bumiaju, Java, a lower jaw with
third molar of both sides complete, which he named Stegodon trigonocephalus praecursor (Fig. 783 of the present
Memoir)

Twenty-fifth Species. —The species Parastegodon? kwantoensis Tokunaga, 1934, PI. ix, a portion of a jaw
with second right molar in situ, is indeterminate, owing to the fact that Professor Osborn regarded Parastegodon
as either a progressive Stegodon or a primitive Archidiskodon. vSee type figure 784 on page 897 below.

Twenty-sixth Species. —Slegodmi T/iishensis Young, 1935, from Yiishe, China. Type, a well preserved
upper left third molar, PL v, fig. 1 (Fig. 785 of the present Memoir).

Twenty-seventh Species.— The type of Stegodon officinalis Hopwood, 1935, is a fragment of an unworn
lower molar, said to have come from Szechuan, China. This is figured in Hopwood, 1935, PI. vii, fig. 3 (Fig.
786 of the present Memoir).

Twenty-eighth Species. —The type of Stegodon zdanskiji Hopwood, 1935, is a fragment of a right third
lower molar, consisting of the first four ridges, and figured in Hopwood, PI. vii, fig. 5 (Fig. 788 of the present
Memoir).

Twenty-ninth Species. —The type of Parastegodon sugiyamai Tokunaga, 1935 [Stegodon? sugiyamai] is

either a first or a second molar, probably of the upper left side, found in Shikoku, Japan (Fig. 789 of the present
Memoir) .
— Editor.]
Thirtieth Species. — In Volume I of the present Memoir (p. 700, fig. 660) will be found the type descrip-
tion of Stegolophodon lydekkeri, dedicated by the present author to his friend Richard Lydekker. The type is

a third left superior molar from Borneo, "much more progressive than the S. latidens type."

III. SYSTEMATIC ARRANGEMENT OF THE STEGOLOPHODONTS AND STEGODONTS

IN PHYLOGENETIC ORDER

. 1. CHARACTERS OF THE SUBFAMILY STEGODONTIN^^

—
Subfamily Characters. (1) Habits chiefly browsing; tropical forest living; crushing of coarse
leafage, herbage, and wood fiber; not progressing to the grazing type. (2) Cranium relatively abbre-
viated, mesocephalic, bathycephalic; grinding-tooth plane deeply depressed to occipital condyles;
occipitofrontal plane neither elevated nor expanded, non-acrocephalic, non-hypsicephalic. (3) Tusks
straight or slightly curved, horizontal or subhorizontal in direction, continuously serving in browsing
habits. (4) Grinding teeth brachyodont to subhypsodont, ridge-plates of
5W-6H
3 increasing from '/4-6-W M
[Stegolophodon] to "I'^J;' [Stegodon]

'(In Volume I (1936) of the present Memoir, the aiitlior si'i)aratecl the Stcgodimtoidea (true St(>gocloiits) from the Elepliaiitoidea (pp. 22, 25), also the
genus Stegodon from Stegolophodon, placing all the Stcgolophodonts in the superfamily Mastodontoidea, family Mastodontidae, new subfamily Stcgolophodon-
tinac (p. 700) and the true Stegodonts in the superfamily Stegodontoidea, family Stegodontidse, subfamily Stegodontinae (see pp. 806-808 above). — Editor.]
838 OSBORN: THE PROBOSCIDEA

Tliese cliariictors are observed in the species and subspecies discovered since 1828, many of which were origi-
nally referred to the genus Mastodon and the genus Elephas and are now referred to the primitive genus Stegolo-
phodon and to the more progressive genus Stegodon.

2. HISTORY OF THE GENERIC NAMES ASSIGNED TO THE STEGOLOPHODONTS AND TO THE

STEGODONTS
The history of the generic term Stegodon Falconer is fully given in Chapter XXI, also in the present Chapter
XI\', which may be sunnnarized as follows: (1) The name Stegodon applies chiefly to the more progressive
Stegodonts. (2) To the more primitive Stegodonts, e.g., the Upper Miocene [Middle Phocene] MastodoJi
cautleyi, the Lower M.
Pliocene' latidens, and the Pliocene M. stegodontoides, the name Stegolophodon Schle-
singer is applicable. The name Stegolophodon also applies to the Mastodon {Bunolophodon) longirostre Kaup
forma suhlaiidens Schlesinger The name Stegolophodon Schlesinger, 1917, preoccupies the
of Austria. name
Prostegodon Matsumoto, 1922-1924. (3) The name Parastegodon Matsumoto, 1924, based on the genotypic

species Elephas aurorse, appears to be in part a synonym of Archidiskodon. These interpretations of the generic
names ado])ted in the present Memoir are displayed in the following table.

i'i(vi,( )( ;i;\v. s and stegodonts arranged uy countries in approximate ascend-

ri;(;oi.()PHODONTs
ing ()I{Di:i{ OI' EVOLUTION, THE MOST PRIMITIVE FORMS BELOW, THE MOST PROGRE-SSIVE FORMS ABOVE
|Th(; species described by various authors since 1929 are omitted here owing to tiic fact that they had not been studied in
detail by Profe.ssor Osborn. — Editor.]
STEGODON
Jajiaii Stegodon orienlalis shodoensis Matsumoto, Island of Shodo, In-
land Sea, Japan
-1924 Elephas {Prostegodon, Parastegodon) aiirorw Matsumoto,
Mt. Tomuro, Kaga, .lajmn
Stegodon 7nindanensis Naumann, Mindanao, Phihppine Islands
Stegodon ganesa var. javanicus Dubois, Kendeng. Trinil, Java
Stegodon trigonorephalns Martin, vicinity of ?Surakarta, .Ia\-a
Stegodon Airdirana Martin, Alas-Tuwa, Kendeng, Trinil, Java
Stegodon orieiitolis Owen, Chungkingfoo, Szechuan, China
Stegodon orientalis gratigeri Osborn, Yenchingkou, Szechuan,
China
Stegodon pinjorensis Osborn, near Siswan, India
(184.5] Stegodon (ranesa Falc. and Caut., Siwahk Hills, India
]184.5] Stegodon insignis Falc. and Caut., Siwalik Hills, India
Stegodon insignis birmanicns Osborn, Mingoon opposite Man-
dalay, I^vu-ma
Elephas honibifrons I'alc. and Caut., Siwalik Hills, India
Mdtitodou elephanloides Clift, near Yenangyaung, Irrawaddy
River, Burma
Elephas cliftii Falc. and Caut., near Yenangyaung, Irrawaddy
River, Burma
Stegodon sinensis Owen, near Shanghai, China
 :

THESTEGOLOPHODONTINiE: STEGOLOPHODON . 839

Superfamily: MASTODONTOIDEA Osborn, 1921

Family: MASTODONTID^ Girard, 1852
Subfamily: Stegolophodontin^e Osborn, 1936

Genus: STEGOLOPHODON Schlesinger, 1917

Oiiginal reference:Denk. Xaturhist. Hofmus., 1917, I, p. 115.

Genotypic species: Mastodon latidens Clift, 1828.
Compare Stego (lopho) don Pohlig, 1888, p. 252.
Syn.: Prostegodon Matsumoto MS., in Osborn, 1923; Matsumoto, 1924, p. 325.

Generic Characters. — (Schlesinger, 1917, p. 115, footnote): "Ich schlage fur M. latidens,
das sich durch seine kurze Syraphyse von dem Subgenus Bimolophodon, durch seinen Molarenbau von
Dibunodon entfernt, den Untergattungsnamen Stegolophodon vor. Der Name bringt einerseits die
nahen Beziehungen zum Genus Stegodon, anderseits die Loslosung der Untergattung von Bimolophodon
und ihre Sonderstelhmg gegeniiber Dibunodon zum Ausdruclc."
Osborn, 1926: (1) Six^ species of iStegolophodon, namely. Mastodon latidens, the more primitive M.
cautleyi, and the more progressive M
siegodontoides; also S. cautleyi progressus, S. nathotensis, and
.

Mastodon {Bunolophodon) longirostre Kaup/orma snblatidens. As defined by the teeth, these species have
in common the following generic characters: (2) Lophs as in Mastodon and Zygolophodon, tendency to
form from four to six transversely arranged cones and conelets (conelets somewhat irregular) and to con-
solidate into ridge-crests; (3) molar pattern transitional between the Zygolophodon type and the
Stegodon type; (4) ridge-crest formula known in Stegolophodon latidens as follows. Dp 3- Dp 4^ 1 ^i^i^ M
M 2 |||4 M
3 li^. [Anterior ridge-crests with persistent median sulcus (see Vol. I, p. 700).— Editor.]

This genus resembles Trilophodon and Tetralophodon in the retention of a broad enamel band on the straight
superior incisive tusks (Figs. 725 and 727) ; it differs widely from Trilophodon in the presence oifour ridge-crests
on the intemediate molars (Fig. 726) ; it also differs from Tetralophodon in the absence of trefoils and in the

progressive tendency of the cones and conelets to form regular transverse ridge-crests surmounted by regular

conelets, as seen in the genotypic species Mastodon = Stegolophodon]

[ latidens, also in Mastodon = Stegolophodon]
[

stegodontoides.

Four of the known species are provisionally distinguished as follows

^Middle Pliocene Lower Pliocene- Middle(?) Pliocene Upper Pliocene
Stegolophodon cautleyi Stegolophodon latidens Stegolophodon sublatidens Stegolophodon stegodontoides
M2A M3^^ M2f|f| M3'^, (?)M3^' M3^*
4-5 conelets on each ridge- 4-5 conelets on each ridge- 4-5 conelets on fourth, 5-6 conelets on each ridge-
crest, very irregular. crest, more regular. fifth, and sixth ridge-crests, crest, very regular. Posterior
Posterior ridge-crests arched ridge-crests arched or convexo-
or slightly convexo-concave, concave.

Stego(lopho)don (see Pohlig, 1888, p. 252). By strict rules this term may be regarded as a nomen nudum and
should not stand in the way of Schlesinger's excellent name Stegolophodon.

Prostegodon. (1) For the first printed use of the name Prostegodon, see Osborn, 1923.601, p. 2: "Prostegodon,

new genus, Matsumoto. In a letter from Dr. H. Matsumoto, dated November 20, 1922, from Sendai, Japan, he
writes: Tn my report just in preparation on the Japanese 'Mastodonts,' I follow you to refer "Mastodon" latidens
to the genus Stegodon, creating however a subgenus Prostegodon for it. Prostegodon is the primitive representative
of the Stegodon-phylum, representing half bunomastodontine and half stegodontine dental characters. Schlosser's

opinion, that Prostegodon might be ? a descendant of "Mastodon" turicens, does not appear to be correct at all.'

Genotypic species Mastodon latidens Clift. Tliis genus should be credited to Doctor Matsumoto."

'[Seven species including Stegolophodon lydekkeri described in Vol. I, p. 700, of the present Memoir. Editor.] —
-[See page 824 above where it is stated that S. latidens is limited to the upper Irrawaddy beds (Lower Pleistocene).— Editor.]
840 . OSBORN: THE PROBOSCIDEA

(2) In 1924 Matsunioto's report on the Japanese Mastodonts, referred to above, appeared in the Journal of
the Geological Society of Tokyo, Volume XXXI, in which lie defined the genus Proslegodon (p. 325). This,
however, was published in the Japanese language.

(3) In 1926 Matsumoto published his English text on this genus ("On Two New Mastodonts and an Arche-
typal Stegodont of Japan," 1926, p. 9), from which the following is a direct quotation:

Skull and mandible onl.y inipcrfeptly known, brevirostral. Lower incisor-tusks might be absent, or abortive if present at
ail. Intermediate molars four- or five-ridged, last molars five- or six-ridged, (irinders essentially lophodont, though their first
and second ridges may show a slight tendency of bunodonty and of trefoil pattern of cusps; mesial longitudinal cleft evident;
inner and outer cusps opposite, instead of being alternate; valleys widely open, free of cement.

(4) Thus Proslegodon Matsumoto-Osborn, 1923, 1924, 1926, becomes a synonym of Stegolophodon Schlesinger,

1917.

The genus Parudegodon Matsumoto, 1924 (1924.2), founded upon the genotypic species Elephas {Prostego-
(.'))

dun) aurorx Matsumoto (Fig. 709), also probably [in part] becomes a synonym of Slegodon, because as shown below
the genotypic species E. (P.) aurorse. = Slegodon aurorx of the present Memoir] is somewhat more primitive than
[

Slegodon airdicana Martin of the Middle(?) Pleistocene and quite distinct from E. [Archidiskodon] plmiifrons.
The name Paraslegodon was originally published by Matsumoto in 1924, pp. 256, 257: Paraslegodon gen. nov.
= Slegodon mmdanensis-Elephas aurorse group type Elephas aurorse; it was subsequently cited by Matsumoto
;

(1926.1, p. 1). Elephas aurorse was originally figured in 1918, PI. xx, as reproduced in figure 709 of the present
Memoir. From the isolated type second superior grinder of the right side, r.M", it was difficult to determine
whether was a progressive Slegodon
this like S. airdwana or whether it was transitional to a primitive elephant

like Archidiskodon planifrons. We have finally referred it to the genus Slegodon.

SYSTEMATIC DESCRIPTION OF SPECIES OF STEGOLOPHODON

Stegolophodon cautleyi Lydekker, 1886 —
Geologic Horizon. According to Pilgrim-Osboru (see Fig.
Figures 142, 08.5, 090, 701, 712-715, PI. xiii 413 of Vol. I), the type of Stegolophodon cautleyi from the Perim
Lcctotype: Middle Pliocene, Perim Island, India; referred, Salt Range,
Inland formation is of Middle Pliocene Dhok Pathan age, equal
Simla Hills, India. approximately to the Plaisancian (of France) and Levantin (of
Falconer's very accurate figure of the lectotype (Fig. 713) Austria and Hungary). In the same formation occur the types of

displays clearly (1) the characteristic median fissure [or sulcus], Anancus perimensis, Deinothcrium indicum, and D. angiistideris,

a i)rimitive character (cf. Fig. 68.5, 2-4), (2) vestigial median alsoHipparion pernncnge type. Referred specimens of cautleyi ,S'.

conules, also observable in Stegolophodon latidens, and (3) internal occur again in the Salt Range, Simla Hills, in the same level with
conelets blocking the valleys. If N. cautleyi is not really ancestral specimens of Anancus perimensis, of Hipparion thcohaldi, and of
to S. lalidens, it is certainly a much more primitive and less perfect Aceratherimn perimense. In the Lower Pliocene' it is rci)laced by
Stegolojjhodont, retaining certain resemblances to the less regular Stegolophodon lalidens.

molars of primitive species of Zygolophodon. The species is repre- Mastodon cautleyi Lydekker, 1886. "Addenda to Synopsis of
sented in the present Memoir by the lectotyi)e (Figs. 712 and 713), Siwalik & Narbada Mammalia." Mem. Ool. Surv. Ind., Pal.
also by the cotype of Lydekker (Fig. 714), and by the referred Indica, 1886, Ser. X, \o\. Ill, pp. xiv xix. Lk( totype and
upi)cr molar (Fig. 715). Compare the superior grinding teeth with cotypes. — {Op. rit., j). xiv) : "The specimens on which this provi-
the somewhat more progres.sive species Stegolophodon caidlcyi sional species founded are five in ntmiber, and are all cheek-teeth
is

progressus (Figs. 72.5-727). of the ui)per jaw; four of them being in an unworn condition."
Specific Characters (Osborn).— Distinguished by median The third superior molar of the left side (Lydekker, 1886.1, p. xv,
formation of ridge-cre.sts .surmounted
fissure [or sulcus], irregular fig. 6) has been selected as the type (see Pilgrim, 1913, j). 294).

by from four to five 'conelets' each, with smaH'conules' or 'acres- Hoiuzon and Locamtv.- -Perim Lsland, India; .Middle Plio-
sory tubercles' in the valleys, as shown in the lcctotyi)c and cotype cenc. Lkctotvpe and Cotvpe Vigvhks.— (Op. cit., j). xv,
figures (Figs. 714 = cotyi)e, 712 = lec1otype). M'' with .">+ ridge- figs. .5 and 6) Fig. .5 (lirit. Mus. M. 2817, cast Amer. Mus. 2696.5),
:

crests; ap. 210 mm., tr. 112 mm. figured in the "Fauna Antiqua Sivalensis," PI. xl, figs. 3, 3a, as

'(See note on page 824 above regarding the Lower Pleistocene age of Stegolophodon lalidens. — Editor.l
 THE STEGOLOPHODONTIN.^: STEGOLOPHODON 841

M. kitidens (appears as M. cautleyi, Lydekker, 1S86.2, ]). 72, fig. distinction between typical molars of those two species, it has
17); V'lg. 6 (Brit. Mils. M.2705, cast Amcr. Miis. 26966), figured in been thought, after considerable hesitation, advisable to pro-
the "Fauna Autiqua Sivalen.sis," PI. xxxi, figs. 6, 6a, as M. latidens visionally apjiiy a distinct specific name to the aberrant form,
(appears as M. cautleyi, Lydekker, 1886.2, p. 73, fig. 18) chosen as which may be called M.
cautleyi. The specimens on which this
. . .

the type (by Pilgrim); Brit. Mus. M.2884 (no history), figured in provisional species founded are fi\e in number, and are all
is

the "Fauna Antiqua Sivalensis," PI. xl, figs. 2, 2a, as M. latidens cheek-teeth of the upper jaw; four of them being in an unworn
(appears as M. Lydekker, 1886.2, p. 71); Ind. Mus. A.48
cautleyi, condition. Three of these teeth, which are all from Perim Island,
(cast Brit. Mus. M.3428),' figured in Lydekker, 1880, PI. xl, as are in the British Museum, and are figured in the 'Fauna Antiqua
M. ])erimenf<is (appears as M. puujabiensis, Lydekker, 1886.2, p. Sivalensis,' under the name of M. latidens: the first (pi. xl. figs.
Ind. Mus. A. 437 (cast Brit. AIus. 2887), figured in Lydekker, 2, 2a [Brit. Mus. M.2884]) is the right
"'— the second (pi. xl.
60); ;

1884.3, PI. XVI, fig. 2, as M. perimensis (appears as AI. cautleyi, figs. 3, 3a [Brit. Mus. M.2817]) is the left"^, and is refigured of

Lydekker, 1886.2, p. 72). the natural size in woodcut fig. 5; while the third (]>!. xxxi. figs.

Fiif. 18.

MastQdo7i catUleyi. —
Tlie third lei't upper true molup, iu an unworn condition;
from the Siwaliks of Pcrim Island. J. The lower border of the figure ia
the inner border of the specimen.

Lectotype of Stegolophodon cautleyi

Fig. 712. Lectotype (see Pilgrim, 101.3, p. 294) of Mastodon cautleyi Lydekker, Fig. 713. Lectotype of Mastodon cautleyi Lydek-
1886, l.M', one-half natural Middle Pliocene, Perim Island, India. Brit. Mus.
.size. ker (.same lei-totype tooth as that shown in figure 712).
M.2705. After Lydekker, 1886.2, p. 73, fig. 18; .same as Lydekker, 1886.1, p. xv, Third upper true molar of the left side, I.M', from Perim
fig. 6. See also four cotype figures of Lydekker, namely, Lydekker, 1886.1, p. xv, fig. Island, one-third natural size. Originally figurotl by
5; PI. XVI, fig. 2 (as M. perimensis); Lydekker, 1880.1, PI. xl (as M. perimensis), Falconer and Cautley, 1846 (1847, PI. xxxi, figs. 6, 6a|
1886.2, p. 60 (as M. punjahiensis); Falconer and Cautley, 1846 [1847, PI. xxxi, figs. as Mastodon latidens, and described as follows (Falconer,
6, 6o (as M. latidens)]. 1868, Vol. I, p. 463): "Figs. 6 and 6o.— .W. latidens.
Observe the irregular character of the ridge-crests and the presence of rudimen- Upjjer true molar, very perfect. —
B.M. Length, 8..) in.
tary intermediate conules in the first and second valleys (Lydekker, 1884.3, PI. xvi, Width, 4.5 in." Brit. Mus. M.2705.
fig. 2, a, a, b, a).

Original Description (Lydekker, 1886.1, p. xiv). — 6, 6a [lectotype, Brit. Mus. M.2705]) is the left"- and is refigured ,

"Mastodon cautleyi, n. sp. Lyd. —

... In recently describing on a larger scale in woodcut fig. 6. The other two specimens are
a molar of Mastodon latidens from Borneo, the present writer in the Indian Museum, and are figured in the present work under
[Footnote: 'Proc. Zool. Soc. 1885, pl. xlviii.'] referred to certain the name of M. perimensis; the first'" [Footnote: '
"Cat. Siwalik
Siwalik specimens in the British and Indian Museums which Vert. Ind. Mus." No. A.48. (1885). M. perimensis.']
pt. I, p. 97.
appeared to indicate a more or less complete transition between (vol. I. pl. XL.) being the partially-worn left "— with the as.sociated ,

typical molars of that species on the one hand and those of M. ^^, the former [Footnote: 'When describing this specimen in
perimensis on the other. A subsequent examination of the speci- Calcutta the writer could not identify it with the one figured in
mens in question has however led to the conclusion that they the "F.A.S." pl. XL. fig. 3, owing to the small size of the figures in
cannot apjiarently be satisfactorily referred to either one of those that work, which renders them almost useless for comparison.']
species; and as there can be no question as to the strongly marked of which apparently agrees precisely with the homologous British

'[Lydekker subsequently (1886.2, p. 60) made this (cast Brit. Mus. M.3428) a cotype of Mastodon [
= Tetralophodon] punjabiensis (cf. Vol. I, p. 363, of the
ent Memoir).
present Mftmoir"^ —Editor.]
P^Hitnr 1
 ;

842 OSBORN: THE PROBOSCIDEA

Museum specimen, and the other'" [Footnote: '"Cat. SiwaUk alsofrom Perim Island. According to these specimens, the ridge
Vert. Ind. Mus." pt. I. p. 97. No. A.437 (1885). M. perimcn.v.s.'] formula of Mastodnn = Stegolophodon] caiitlci/! is as follows:
[

x\i. the imperfect right "", in an unworn

(vol. ITT. pi. fig. 2)
Ridge formula: Dp i'-^=^ M l*-^"^^ M 3^^.
condition."
According to P'alconer and Lydekker, (1) the same ridge
Specific Characters (Lydekker, 1886.1, p. xv). — "All formula is observed in specimens of Mastodon = Stegolophodon] [

these five specimens agree so exactly with one another that there cautleyi from Perim Island (type locality) and from the Dhok
can be no hesitation in referring them to one and the
little or Pathan, namely, 2- M M
3—; (2) the open and irregular ridge
same Their essential characters are that the ridges are
species. structure of the molar teeth in S. cautleyi is more primitive than
moderately tall, and inclined forwards; the valleys partially the close-set, regular ridge structure in the type of lalideitti, as is .S'.

blocked by accessory tubercles, of which there are none on the seen at once by comparison of the type figures of the two species;
outer side of the median longitudinal cleft. The first inner column (3) Lydekker separated the species Mastodon cautleyi from speci-

always has accessory tubercles on both and there are similar sides, mens in the Perim Lsland beds which had been referred by l''alconer
tubercles on the hinder side of both the second and third inner to Mastodon latidens; (4) Pilgrim (1913, p. 294) accepted this
columns: the hind talon of the 'intermediate' molars (woodcut species, regarding it as a direct ancestor of one of the larger forms
fig. 5) is relatively small; while in unworn e.xamples the hinder of M. latidens.

Fig. 14.

Fie;. 17.

Maiiodon penmcnsis.— Uhe second left upper triie molar, in nn unworn con-
Masiodon caullcT/i.— The Rrsl Mt upper true molar in an unworn contlition; dition from the Siwaliks of Perim Island, j.
;
a. Eiternal accessory

from the Siwaliks of Perim Island. }. The lower border of the figure is tubercles. The lower border of tlie figure is tlio inner border of the speci-
the inner border of the specimen. men. (From the Palajontologia Indica.')
'

CoTYPE First and Refebred Second Superior Molar.s op Stegolophodon cautleyi

Fig. 714. Cotypc of Mastodon cautleyi Lydekker, 1886 (Brit. Mu.s. Fig. 71.5. .1/astoiore pmmens^'s (Inil. Mus. A.355, cast Brit. Mu.s. M.2851).
M.2817), p. XV, fig. r,. Compare "Fiiuna .'\iiti(|iia Sivalensi-s," Fnle. and After Lydekker, 1886.2, pp. 57 and 58, fig. 14.
Caut., 184611847, Pi. xi-, figs. 3, 3a (;l/. Inlulcns)]. After Lydekker, 1886.2, Reierrrd by Oshorn to Stegnlopkmlon caiillfi/i. Compare type of ,S. mi/^r'i/J
p. 72, fig. 17. progressus (Fig. 726).
Compare type of S. cautleyi progressus (Fig. 726).

aspect of the outer column of the first ridge is deeply concave, and Stegolophodon latidens Clift, 1828
the arrangement of the tubercles on the inner column of the same Figures 685, 693, 694, 716-721, 757, PI. xiii

ridge forms a V. The third true molar (woodcut fig. 6) is very Lectotype: Lower Pliocene, lowest levels of the Irraw.addy Series
wide, tapers but little posteriorly, and carries five ridges and a (fluviatile)- near Yenangyaung, Burma; referred, Middle Pliocene of India
simple hind talon, the latter consisting of a narrow ridge with six (Dhok Pathan zone), and Ixnver (7) Pliocene of China and Japan.

small tubercles. All the teeth are relatively wide, with a well- This is the chai'acteristic Lower Pliocene- stage of the Irra-
marked median longitudinal cleft; they appear to have no a]3preci- waddy Series, Burma and cotypcs) and of the Middle
(lectotyi)c
able quantity of cement in the valleys, and when worn present . . . Pliocene stage of the Siwaliks and Perim Island, India (referred
trefoils on their inner columns." The cotypc figure reproduced specimens).
above (Fig. 714) is a first left superior true molar from Perim Islaiui Specific Cmauacters (Osborn). — Distinguished by more
the lectotype (Figs. 712 and 713) is a third left .superior true molar. regular formation of the ridge-crests than in Stegolophodon cautleyi,

'[Cast Brit. Mus. M.2887.1

'[See note on page 824 above regarding the Lower Pleistocene age of Stegolophodon latidens. — Editor.)
 THE STEGOLOPHODONTIN^: STEGOLOPHODON S4:{

siinuounted by four to five conelcts each, with two vestigial

'conules'; median fissure [or sulcus] in anterior ridge-crests; ridge-
crests more stegodontoid. Hidge-crest formula: M 2 HtjM 3 ^'.01.
This important stage, originally described as Mastodon by
( lift and as Slegodon by Falconer, has been aptly chosen as the

STEqOLOPHODON LATIDENS
844 OSBORN: THE PROBOSCIDEA

History of Discovery. — The history of discovery of this Tetralophodon punjabiensis, Hipparion pimjabiense, Tetraconodon
species (Stegolophodon latidens) in 1826 by CHft is very fully set magnus, and Tauroiragus latidens.
forth in this Memoir (pp. 825, 827 above) and need not be repeat- Referred specimens of Stcgolophodnn latidciif:, according to
ed here; also Cliffs original description and ridge formula as Pilgrim, also occur in the overlying Dhok Pathan beds (Salt Range)
abbreviated above. Attention should be concentrated on the associated with the types of Tetralophodon punjabiensis, Tetra-
characters of the cotype specimens as shown in the four figures of belodon = Synconolophus] cornigatus, Mastodon = Si/nconolophus]
[ [

hasnoti, and Stegodon bombifrons. It is probable that

specimens referred by Falconer to Stegolophodon lati-

dens from the Upper Miocene [Middle Pliocene] of
Perim Island belong rather to *S'. cautletji..

231. Premifere arrifere-molaire superieurc du Mastodon latidens,

a 1/2 grandeur, dicouverte par Crawfiird, dans I'Avu.

Lectotype Molars of Stegolophodon latidens Fig. 720. Lectotype M' of Mastodon [Stegolophodon] latidens

Fig. 718. [Perspective.] Lectotype of 3/o.5/o(ton iahticHsClift , 1828, reduced from Cliffs Clift, 1828, reproduced from an engraving by Gaudry (1878,
PI. XXXVII, fig. 1, Found on the left bank of the Irrawaddy River,
to one-half natural size. p. 17.5, fig. 231). The third superior molar discovered by Craw-
below Ava, Burma. (Clift, 1828, p. 371 and Explanation of Plate.s): "The i)alate, and furd below Ava.
molar teeth of the right side of Mastodon latidens (a younger animal than the last). The
anterior tot)tli is very much worn. The anterior part of the posterior tooth appears to have
Geologic and Geographic Distribution. —The
been just brought into use." These teeth include a right M- and M^. The ridge-crests arc: cotypes oi Stegolophodon hitidens are from the Irrawaddy
M 2^^^ M Z^iiSiA^, Cast Amer. Mus. 21978. See orthogonal projection (Fig. 71(i)- River, 250 miles below Ava, near Yenangyaung,

Fig. 719. (Left) Section of molars, M^ M^ (Jde Falconer, Pal. Mem., 1868, Vol. I, p. 424) of Ma.-ilodon Uilidens Clift, 1828 [Stego-
lophodon latidens, lectotype], reduced to one-third natural size. After Falconer and Cautley, 1846 [184.'3, PI. iii, fig. 8). Compare witli
tlie crown view of same molar reproduced above.
(Right) Section of M^ of Mastodon americanus, from Missouri, op. cil., PI. in, fig. 9, inserted by Falconer for comparison. Rrit.

Mus. 17420.

the superior molars reproduced above(Figs. 718,719,720,716), and Burma. In the Calcutta Museum are listed by Lydekker
(1)
in the lower jaw (see Fig. 717 for r.Ma). (1885, pp. 94-97) thirty-three specimens referred to M. latidens
Geologic Age. — The type of Stegolophodon latidens, according ('lift, as follows: (a) Burma— six specimens of grinding teeth; (b)

to Pilgrim (notes of April .3, 1923), probably occurs in the lowest Asnot, Punjab— five —
grinding teeth; (c) Sind five grinding teeth;
levels of the (fluviatile), near Yenangyaung,
Irrawaddy Series (d) the remainder are from the Punjab, Potwar district, Gadari,
Burma, of Lower Pliocene age.' In beds of the same age in the Lehri, Jabi, Niki. (2) In the British Museum collections are listed

Salt Range, Pabbi Hills, Jammu, and Kangra, are also found by Lydekker (1886.2, i)p. 74-78) eleven specimens from: (a) The
'[See note on page 824 above. ^Editor. 1
 : — —

THE STEGOLOPHODONTIN^: STEGOLOPHODON 845

Pliocene Siwaliks of the Punjab; (b) from near Yenangyaung, outer columns soon uniting. The hind
talons are very large, while
upper Burma; (c) Perim Island, Gulf of Cambay; (d) the Plio- the inner border of the crown concave and without a distinct
is

cene of Bruni, northwest Borneo [made by Osborn (1936, Vol. cingulum a fifth ridge is
; sometimes present in "' - and the . . . ;

I, p. 700, of this Memoir) the type of Stegolophodon lydekkeri]; hind talon of "^ is always large, and may sometimes be reckoned
(e) Lehri, Punjab. as a sixth ridge. The enamel is very thick, and in the hinder
Lower Pliocene' Age. — Pilgrim assigns (Ghap. XXII below) teeth quite smooth cement is usually absent, and premolars were
;

Stegolophodon latidens cotypes to Burma, Irrawaddy Series, lowest probably developed."

levels and referred specimens to the Jammu and
(fluviatile), Summary of Characters (Lydekker, 1880.1, p. 236).
Kangra beds Range and Pabbi Hills.
of India, also to the Salt "Having now passed in review the molar series of M. latidens, we
According to Pilgrim the species ranges upwards into the upper may sum up what is known regarding the species. The adult
Irrawaddy levels (fluviatile), Burma, and into the Salt Range, cranium is unfortunately quite unknown the palate is noticeable;

Hasnot, Dhok Pathan zone of India, equivalent to the Middle from the extent to which the molars converge anteriorly. The
Pliocene Plaisancian (of France) and Levantin (of Austria and mandible is known by a specimen of the greater part of the right
Hungary). Stegolophodon latidens may be regarded therefore as ramus, containing the two last molars, in the collection of the
a characteristic Lower Pliocene' stage of the Stegodonts; whereas Indian Museum. This mandible is very long and slender, and
the more primitive Stegolophodon cautleyi specific type belongs sub-circular in cross-section, in the middle its vertical diameter at
in the Middle Pliocene of Perim Island, also referred specimens the penultimate molar being 5.8 inches and its transverse diameter
of the Salt Range, Simla Hills. 5.6 inches. The lower border is nearly straight up to the symphy-

NOTES OF FALCONER, 1868, ON MASTODON [STEGOLOPHODONl

LATIDENS, ALSO OF LYDEKKER, 1886
Falconer, "Palaeontological Memoirs," VoL I, pp. 424, 463, 471, Pis. iii,

XXX, XL of the "Fauna Antiqua Sivalensis"; Lydekker, "Catalogue of the

Fo.s.sil Mammalia in the British Museum (Natural History)," Part IV, 1886,
p. 74, & Narbada Probosc-idia," 1880, p. 236.
and "Siwahk
Stegolophodon latidens. Lectotypc. Plate iii, fig. 8, IVP
ridges 4)2, M' ridges )^-5-)^, 4 conelets or denticles, enamel thick,
cement thin.
Upper Molars. —Plate xl, figs. 1, la, ?second upper milk
molar, Dp', with 2 ridges; upper milk molar.
figs. 2, 2a, ?third
Dp'', with 4 ridges; figs. 3, 3a, M'
and back and front heel
ridges 4
()4-4-'4 ridges), length 4 in. = 101 mm., width 2.3 in. = 59 mm.

—
Lower Molars. Plate xxx, fig. 6, lower jaw, I.M3, 5 ridges and
a double heel (5J2 ridges), no cement, length of molar 11.3 in. =
288 mm., width 4.5 in. = 114 mm.
From these observations of Falconer, and from those of
Lydekker below, we may write the following collective formula, Referred Stegolophodon latidens of Japan, a very Primitive Stage
jM-imitive and progressive Fig. 721. Referred 4}l'+ ridge-crested Stegolophodon latidens {cautleyi^)

Stegolophodon latidens: Dp 3^ Dp 4- M 1 ^^^^^ M 2^ from Sliiwogama, Miyagi District, Province of Rikuzen, Japan; "probably of
typical Pontian age [Lower Pliocene]." Cast Amer. Mus. 22610. Inner and
^^^ "^ i4-5-^i superior aspects of a third right upjier molar, r.M', with 4}^+ ridge-crests.
Osborn, 1927: Cliffs figures (Figs. 717 and 718) as well as (Upper) Internal aspect; (lower) crown aspect. After Matsumoto, 1926,
his description assign to M3 the following ridge-crests -^ which PI. V, figs. 1 and 3: "Fig. 1. Proslegodon latidens (Clift); right D* or M';

should be interpreted as ^fff^f. This is approximately in accordance inner view; natural size. Fig. 3
. . [crown] view; natural size." Reduced
. . .

in the present figure to one-half natural size. Length of r.M' along median
with Falconer and Lydekker's formula, although, as in all the Stego-
line 115 mm. Compare Mastodon (Banolophodon) = Stegolophodon] sublatidens
[
donts, it is difficult to count the new or rudimentary ridge-crests. Schlesinger (Fig. 710).
Characters (Falconer, 1868.1, Lydekker, 1886.2). (1) —
The presence of a median furrow between the primitive inner and sis, with a slight convexity in the middle. From the extremely
outer lobes, (2) of four conelets or denticles on each ridge, (3) of small size and circular section of the ramus where broken off at

thick enamel and thin cement, and (4) of conules in the two anterior the commencement of the symphysis, the latter must have been
valleys — the.se are the chief characters assigned by Falconer, 1868. short, and was probably unprovided with incisors."

Lydekker (1886.2, p. 74) characterizes the dentition as follows: "The ridge formula of the milk-molars is constant, and con-
"The mandibular symphysis is not certainly known, but it was forms to the normal tetralophodont order; in the true molars,
probably short and tuskless. The upper molars are \ery wide, with however, there is not an unfrequent tendency to assume a pentalo-
no cingulum and relatively low transverse ridges, the median longi- phodont type, the hind-talon of many of these teeth not un-

tudinal cleft being frequently indistinct, the accessory tubercles frequently, partly or entirely, taking the form of a fifth ridge. In
small and the valleys comparatively open, the trefoils on the worn the following ridge-formula these varieties are indicated :

ridges imperfect, and the worn dentine surfaces on the inner and Milk-molars. True molars.
2+3+4 (4-5)+(4-5)+(5-6)
'[See note on page 824 above. — Editor.) 2+3+4 4+(4-5)+5
 —

846 OSBORN: THE PROROSCIDEA

This tendency to the prfKluction of an additional ridge in tlie 102. Typk. — A third light superior molar, v.'SP. ('ollection
true molars of M. Jnlideiix will be siil)se(|uently shown to he (if the Royal Museum fif Natur.al History, \'ierma (alter Samm-
a character which it possesses in connnon with M. .s/ra/cz/.s/.s. lungs bestand). IIohizo.n axd Focamiv. — From near
This tendency to variation enables us easily to com])rehend how Teschen (Schlesien), Austria; MiddleC?) Pliocene. Tvi'k Fk;-
the passage from the tetralophodont Mastodons to the hexalopho- I'RE. Op. cit., Taf. XVII, fig. 2.

dont Stegodons (S. cliftii) was effected." Type Description.— (Schlesinger, 1917, i)p. 101, 102): "Es
ware unmoglich, diesen Zahn, den Typus e'lner forma sublati-
icli als
STEGOLOPHODON LATIDENS IN CHINA AND JAPAN dens n.f. auffasse, innerhalb der Variationsbreite von M. longi-
According to Schlosser (1903, p. 45), in tiie Munich collection. rostre anzufiihren, wenn nicht die vorbeschriebenen Molaren direkt
from an unknown locality, is a fragment of a lower molar tooth zu ihm iibei'leiten wiirden. PrakLisch ist der Zahn— stegodont, d.h.
referable to -1/. hilidens. Schlosser's figure of this fr.agnient re-
cr weist jene Form von Jochzithnigkeit auf, die im (iegensatze
i.s

produced in figure 770, 7.

zu den Verhaltnissen bei den tapiroiden Mastodonten durcli Um-
A primitive Japanese stage (Fig. 721) has been referred to this bildung der Sjierrhocker zu Xel3en])feilern und jMnordnung in die
species by Matsumoto; it apparently possesses but four and Flucht gleichgrosser. Mammillen erreicht wird, wahrend .jene
a half ridge-crests and thus approaches the »S'. caiitlcyi stage (cf.
.Vrten und Nebenhocker versehmelzen und die Spei-r-
Ilaupt-
Figs. 713 and 712). hoeker als Crista nach aussen drilngen und dann unterdriicken
[Footnote: 'Ich kann nicht umhin, auf die liestiitigung des "(ie-
Stegolophodon sublatidens Schlesinger, 1917 setzes der Nichtumkehrbarkeit der Entwicklung" (L. Dollo) hin-
Figures 710, 722, PI. xiii zuweisen, die darin liegt, dass eine funktionelle Zygodontie von
MiddleC?) Pliocene age. From near Te.sehen (Sehlesien), Au.stria. der Bunodontie aus nicht durch ofi'ene Riickkehr zum Urtypus
This species (Fig. 722) is founded ujion the posterior half of (M. pygmaeum), sondern durch die Rildung der ganz anderen
a superior molar quite closely comparable to Pilgrim's type (Fig. Stegodontie erreicht wird.']."
723) of Mastodon [ = Stegolophodon] stegodontoides; it is more History. — In defining the genus Stegolophodon, with the geno-
prinuti\'e in every way but the affinity is very striking; it thus typic species Mastodon latidens ('lift, of Burma, Schlesinger implies,
but does not so state with definiteness, the inclusion of his species
'Mastodon stihlatidens' within the genus Stegolophodon. His type
of Mastodon sublatidens (Fig. 722) is an imperfect third superioi-
molar of the right side, r.M', from Schlesien, Austria, of Pliocene
age. The Jive conelets, of which two of the crests are comi)oscd,
ai'e arched transversely; this arcuate character is exceptional and
does not appear in the two species of Zygolophodon (Z. borsoni,
Z. pyrenaieus) nor in the genotypic species of Stegolophodon, name-
ly, Mastodon latidens of Burma,
which the ridge-crests ai'C
in
directly transverse. The resemblance to Stegolophodon stegodon-
toides is much closer, although S. sublatidens is a smaller and more

Type of Stegolophodon sublatidens jirimitive form, with more open valleys and less distinct division of

Fig. 722. T\\)C of Mastodon (Bunolophodon) lungirostre Kaup funiia sub- the cones into five conelets, as admirably shown in the accompany-
latidens Sclilesingcr, 1917, posterior half of a tliird right superior mohir, ing figures 722 and 723, the types of these two .species.
r.M'. From near Teschen (Schlesien), Austria, Middle(?) Pliocene. Collec-
tion of the Royal Museum of Natural History, Vienna ("alter Sammlungs
bestand"). Photograph loaned by Doctor Schlesinger. Reproduced one-half Stegolophodon stegodontoides I'iliirim, 1913
natural size. Compare Schlesinger, 1917, Taf. xv^i, fig. 2.
Figures 68.5, 708, 723, PI. xiii
The five conelets, of which two of the ridge-erests are i-omposcd, are
Lehri, in tlic Piuijal), India, possibly tlpi)er Siwaliks. ITpperC?) Pliocene.
arched transversely, a character not observed in Stegolopliodon liUidcns, in
which the ridge-cre.sts are directly transverse, but seen in the type of Stego- This is the characteristic Upi)er(?) Pliocene stage of the Pin-
lophodon stegodontoides.
jor horizon. Upper Siwaliks.
Specific Characters (Osbohn). Larger and much more—
appears to represent the first true Slrgolnphodoti disco\-erc(l in |)rogressive than Stegolopfiodon latidens. M' measures ap. 210 mm.,
Europe. tr. 110 mm., approximately same dimensions as the type of
Spkcific —
Characters. Inferior in size to Stegoloitliddnii Mastodon = Stegolophodon] cautleyi. Ridge-crests 6)2 with five to
[

stegodontoides] valleys between ridge-crests more oi)en, four to six conelets. ^'alleys compressed or part ly closed, less ojjen han in t

five conelets on fourth, fifth, and sixth ridge-crests; each j)osterior S. latidens. \estige of median fissure on three anterior ritlgc-ciests
ridge-crest arched or convexo-concave; width of tetartoloph 82 (Fig. 723).
mm., width of tetartoloph in .S'. stegodontoides 107 mm.; rudimen- I'he tyi)e tooth of this species is transitional between the
tary cement in the valleys. Stegolophodon and the Stegudon pattern, yet the species belongs to
Mastodon (Bunolophodon) longirostre Kauj) fon/io sulilnliilens an independent line of descent contemporaneous in the rp])er
"Die Mastodonten des K. K. Xaturhistorischen
n.f. .Schlesinger. Pliocene with an advanced stage of Stegodon such as Stegodon
llofmuseums." Dcnk. Xatiuhist. Ilofmus., I, 1917, pp. 101, insignis. From Lehri, in the Punjab, India, Pilgrim (1913) selected
 THE STEGOLOPHODONTINJE: STECJOLOPHODON 847

a six and a half crested third right upper true molar, r.M', as the Osborn, 1924: While we recognize the usual four columns in
ty]>e of a new species Mastodon slegodoiitonles. lie remarked {op. this molar tooth, we observe in Lydekker's figure that the summits
ciL, p. 294): "The last species of Mastodon which can be referred of the two inner columns tend to subdi\ide thus in the tetartoloph ;

to this [cauUeyi-latidens] line is the tooth from l.ehri, of which the and the pentaloph there are five distinct conelets, as compared with
horizon is uncertain but may be possibly Upper Siwahk." The four plus observed in Stegolophodon latidens and in aS. cautleyi.
type specimen was originally figured natural size by Lydekker in Lydekker's Description (1880.1, pp. 235-237).— "77mf/
"Palseontologia Indica," ser. 10, Vol. I, 1880, PI. xxxix, as: —
upper true molar. The large tooth represented in Plate xxxix is
"Mastodon (Tetralophodon) lalidens, Clift. The third right upper a specimen of the last upper molar of the right side of M. latidens,
true molar: from Lehri, in the Punjab. The specimen is drawn collected by Mr. A. B. Wynne near the village of Lehri, in the
of the natural size and is viewed from the inner [outer] side." Punjab. This specimen is implanted in a fragment of the maxilla,
Mastodon stegodonloides Pilgrim, 1913. "The Correlation of which also contains the two last ridges of the preceding or penulti-
the Siwaliks with Mammal Horizons of Europe." Pec. fieol. mate tooth. The figured tooth is entirely unworn, and was still
Surv. India, Vol. XLIII, Pt. 4, p. 294. Type.^A third supe- covered by the gum at the death of the animal. The crown carries
rior molar of the right side, r.M^, Ind. Mus. A.86. Horizon six transverse ridges, the hindmost of which is considerably smaller

.\ND Locality. —
Possibly Upper Siwaliks, Pinjor formation, than the others, and probably rejiresents an ultra-developed talon,
Upper (?) PHocene; Lehri, Punjab, India. Type Figure. — as v,e saw to be the case in the penultimate molar represented in
fig. 1 of Plate xxxviii. The tooth consequently belongs to a 'pen-
talophodont' type of dentition. The ridges are low and simple,
and with the exception of the first, are slightly convex anteriorly,
and as concave posteriorly. Each ridge is divided by a longitudinal
cleft placed somewhat externally to the mesial antero-posterior
axis of the tooth. The internal moiety of each ridge, with the ex-
ception of the second and sixth, bears three mammillae or cusps,
while the external moiety bears only two on each ridge. The val-
leys are quite simple and uninterrupted. In this tooth, as also in
all the previously described specimens, there is no trace of cement.
The length of the specimen is 8.6 inches, and its greatest width 4.2
inches."
Lydekker's description and figure concur in giving five cone-
lets as characteristic of each ridge and as showing the three poste-
Type of Steoolophodon stegodontoides rior ridges more progressive than the three anterior.
Fig. 723. Type r.M' of Mastodon stegodonloides Pilgrim, 1913. After Also exhibiting this six+ ridged, five to six conelet condition,
Lydekker, 1880, PI. xxxix: "Mastodon {Tetralophodon) latidens, Clift. The is a third left upper true molar from Borneo erroneously referred
third right upper true molar: from Lehri, in the Punjab. Tlie specimen by Lydekker to Mastodon latidens.'^
is drawn of the natural size, and is viewed from the inner [outer] side." Ind.
Mus. A.86. Reduced to one-half natural size. Provisionally placed in the
Upper Pliocene, Pinjor formation (see Vol. I, fig. 413, also PI. xm). Stegolophodon nathotensis Osborn, 1929
Figure 724, PI. xiii
Lower Chinji horizon, near Nathot, India; Mio-Pliocene.
Lydekker, 1880.1, PI. xxxix, figured as Mastodon {Tetralophodon)
latidens. This type was discovered by Barnum Brown in 1922 in the
Type Figure. — This fine half-size drawing of the type tooth Lower Chinji horizon, near Nathot, India, the exact level being
measures: ap. 210 mm., tr. 110 mm. The ridge formula is 3—. M unrecorded. As shown in figure 413 the species is of approximately
In the anterior three ridges of the tooth there is still a trace of the same geologic age as Trilophodon chinjiensis type, Deinotherium
subdivision into inner and outer lobes, which disappears in the pentapotamise type, and Serridetdinus browni type. The Chinji
posterior three ridges. The conelet formula is: protoloph — six, horizon, of a total thickness of 2,400 feet, doubtless represents
metaloph— five to six, tritoloph —four to five, tetartoloph — five, a \-ery long period of geologic time.
pentaloph — five, hexaloph —four; the conelets are more or less Stegolophodon nathotensis Osborn, 1929. "New Eurasiatic and
discrete, the valleys apparently open. Consequently this molar American Proboscideans," Amer. Mus. Novitates, No. 393, Dec.
crown appears to be descended from that of Stegolophodon latidens 24, pp. 13-15 (Osborn, 1929.797). Type.— Amer. Mus.
and represents a marked advance towards the true Stegodoti type. 19455. Posterior half of a fragmentaiy right third suj^erior mo-
Specific Characters (Pilgrim). —
(1) Owing to the almost lar,r.M^ (Fig. 724, C, CI); posterior half of a right third inferior
entire absence of accessory columns the name Mastodon stegodon- molar, r.Ms (A, Al) anterior half of a left second superior molar,
;

loides is proposed. (2) It will be seen that the type molar carries l.M^ (B, Bl). Horizon and Locality. Found near Nathot, —
on none of its ridges more than the usual four columns [character- India, "Lower Chinji horizon, the exact level being unrecorded,
istic of Stegolophodon latidens], while the anterior ridges of Stegodon lower Middle Siwaliks Middle to Upper Miocene [Mio-Pliocene]."
;

elephantoides { = cliftii) carry nine or ten mammillae [i.e., conelets]. Type Figure.— Op. cit., 1929.797, p. 14, fig. 14.

[Made by Professer Osborn in 1936 the type of Slegoloptiodon lydekkeri (see Vol. I, p. 700, fig. 660). —Editor.]
 ,

848 OSBORN: THE PROBOSCIDEA

Type Characters. — Ridge-crests low, blunted, with four to No. 393, Dec. 24, pp. 13 15 (Osborn, 1929.797). Type.—
five blunted conelets (Fig. 724 B, A) on each crest; ridge-crests Amer. Mus. 19446. A juvenile cranium (Figs. 725 and 727),
transversely arcuate or arched, as in Stegolophodnn stegodonioides. containing right and left .superior tusks with broad enamel band,
Enamel thick with slightly indented border. Hugose externa! also in .situ third and fourth superior deciduous premolars and
cingulum. Ridge-crest formula unknown, probably M 3 Ij^; first molar of both sides (see Fig. 726, r.Dp'"'', r.M'). Horizon
characters of ridge-crests and conelets quite distinct from those
—
AND Locality. "Twelve miles east of C'hinji Bungalow, India . . .

of Stegolophodon latidens, ridge-crest formula probably lower than Collected by Barnum Brown in 1922 at summit of ]>ower Chinji
in S. latidens. Species at present known by the type specimen horizon, 2,000 feet above base of Lower Siwaliks. Middle Miocene
only, which belongs in a lower geologic level, namely, Mio-Pliocene, [Mio-Pliocene]." Type Figvre.— Op. cit., 1929.797, ]x 14,
than the type of S. latidens, which belongs in the Lower Pliocene.' fig. 15. See also figures 725 to 727 of present Memoir.
See measurements in legend (Fig. 724). Type Characters. — Superior tusks laterally compressed.

STEqOLOPHODON NATHOTENSIS

^j ouier vieiv^' i,
°-"'^ icppe

Fig. 724. Type Osborn, 1929 (Amor. Mils. 19455), Lower Chinji horizon, from near Nathot, India. One-half natural size.
of Stegolophodon nalhotensis
A and a half ridge-crests. Meas.: ap. 180e mm., tr. 88e mm.; height of nietalo|)li .38e mm.;
third superior molar of the right side, r.M'*, probably with four
I.M^ (protoloph) tr. 73 ram., (metaloph) 36e mm.; r.Ms, tetartolophid .and jjentalophid (talon only). Ridge-crest of tetartoloph of r.M' with five arcuate
conelets. Mjussive, brachyodont. Osborn, 1929.797, p. 14, fig. 14.

Stegolophodon cautleyi progressus Osborn, 1929 with broad external enamel band ; ridge-crest formula as compared
Figures 725-727, PI. xiii with that of Stegolophodon cautleyi, as follows:
Summit of Lower Chinji horizon, 2,000 feet above base of Lower Siwaliks, Stegolophodon cautleyi progressus: r.Dp 3^ r.Dp4 — - r.Ml^^^^
twelve miles east of Chinji Bungalow, India; Mio-Pliocene.
Stegoloj}hodon cautleyi (typical): r. Dp 3 — r. Dp 4
)i-3-ii

This type is of great importance and interest as yielding for r.M 1^^
the time a knowledge of the cranial structure of Stegolophodon
first Four ridge-crests in intermediate molars, r.Dl)^ r.M', with
in its Miocene stage of evolution. Distinguished as Stegolophodon four irregular conelets on each crest; rudimentary anterior and
by its four-crested intermediate molars, its superior tusks, and its posterior ridge-crests in r.M'; conelets less blunt and crowns less
cranial profile, which remind us strongly of the primitive type of brachyodont than in Stegolophodon nalhotensis; median fissure in
skull and tusks seen in Trilophodon and in Serridentijius; the four- r.M' wanting, as shown in comparison of figure 726 with figure 714,
crested intermediate molars parallel those observed in Telralopho- cotype of Stegolophodon cautleyi, or decidedly less distinct than in
don, a genus from which the present Stegolophodon is shar])ly re- from Perim Island (Fig. 712) and the referred
S. cautleyi lectotype
moved by the absence of trefoils, vestiges of which ai)i)car only in l.M- (Fig. 715); traces of irregular internal trefoil conelets on
the valleys between the ridge-crests. Altogether Stegolophodon r.Dp< and r.M'.
cautleyi progressus (probably of the summit of the Miocene or Of very great importance and interest is the i^re.sence of
lower levels of the Pliocene) is a progressive ascending mutation superior incisive tusks with lateral enamel band as well as other
of the typical Stegolophodon cautleyi of Perim Island, which we evidence in the structure of the superior grinding teeth of the
regard as of Middle Pliocene age. affinities of this tetraloijliodont tyjie with the much more i)riniiti\e
Stegolophodon cautleyi progressus Osborn, 1929. "New trilophodont mastodonts of the Lower Miocene of France (cf.

Eurasiatic and American Proboscideans," Amer. Mus. Novitates, also Figs. 725-727).

'[See notes on pages 814 and 824 above. — Editor.!

 STEQOLOPHODON CAUTLEYI PROqRESSUS

Amer AfU5. I9^^b '^yP^

~ 7ia.t. sije

Fig. 725. Type cranium of Stegolophodon cautkyi progressus Osborn, 1929 (Amer. Mus. 19446), same specimen as figure 727.
Al, Superior aspect, exhibiting cranium with anterior narial opening and prcmaxillaries.
A2, Palatal view, exhibiting lateral enamel band of tusks, much worn Dp' with remnants of 3+ ridge-crests, Dp^ with 4+ ridge-crests, M' with 4+ ridge-

crests, also posterior narial opening and basicranial foramina.

A3, Right lateral aspect, exhibiting enamel band on tusk.
.\4, Occii)ital aspect, slightly compressed by crushing.

Amer Mus /9^^6 T'ype

i nat szje
%^
Fig. 72(5. of SlignlophoJon caiilkiji progressus Osborn, 1929.
Type Detailed study of right
Dp'-M', combination drawing of two sides. Coronal ridge-crests intermediate in
sui)erior lientition,
formula and pattern between the typical Mastodon [=Slrgolopho<hm] cautleyi Lydekker and M.
1
= SlegohpfiodoH] lalidins Clift; similar in the regular disposition of the cones and conelcts.

849
 Fig. 727. Type skull (Amor. Mus. 19440) of Slcgohphodon catd-
progreasus Osborn, 1929, collected by Banuini Brdwii twelve miles
Iciji

east of Chinji Bungalow, summit of the Lower Cliinji horizon, 2,000

feet above base of the Lower Siwaliks, India. This skull, containing
tiie

the right and left superior tusks, third and fourth superior deciduous

premolars, and the fii'st superior molars (partly or\ipted), shows the
laterally compressed tusks with enamel band, 4+ ridgc-crests on
Dp"*, 4+ ridge-crests on M'; it is somewhat more progressive
than
the type of Mastodon [
= Stcgolophodon] caulleyi Lydekker.
Left lateral, palatal, and right lateral aspects, one-fifth natural size.

850
 STEGOLOPHODON

ExtLTnal
S STECODONTOIDES TYPE

S.LATIDENS TYPE
S.
Extfrmxt
CAUTLEYI TYPE
H
S.LATIDENS after CUft

5 CAUTLEYI PT^QOPIESSUS TYPE

S.SUBLATIDENS TYPE S.NATH0TEN51S

PLATE XIII
Stegolophodontin.e, Stegolophodon: Primitive Longitudinal Sulcus, Shaded (A, B), Persisting in I-IV Anterior Ridge-crests (E, El),
IN I-III Anterior Ridge-crests (F), Vestigial in Ridge-crests I-III (G, H, I). Conelets Rounded, Increasing from 4-5 (A, D),
5-7 (C, H, I, G). Posterior Ridge-crests IV-VI Progressive, with Conelets 5-7 (G), Conelets not Exceeding 5 (H, I)
Mio-Pliocene B, Stegolophodon cautleyi progressus type, r.M'. Amer. Mus. 19446, near Chinji Bungalow, India. Summit of Lower Cliiiiji

horizon, 2,000 feet above base of Lower Siwaliks.

C, Stegolophodon natholensis type, r.Ms. Conelets 5-6. Amor. Mas. 19455, near Nathot, Lower Middle Siwaliks, India. Lower
Chinji horizon.
Lower Pliocene' D, Stegolophodon latidens cotype, r.Ms. Ridge-erests I-VIII, conelets 4, sulcus on ridge-crests I-IV. After Clift, 1828,
PI. XXXVIII, near Yenangyaung, Burma.
fig. 1,

E, Stegolophodon latidens cotype, r.Ml Ridge-crests I-VI, sulcus on ridge-crests I-IV. E 1 (.section), ridge-crests coalescent
at base (I-III). After cast (Amer. Mus. 21978) of Cliffs type, 1828, PI. xxxvii, fig. 1, near Yenangyaung, Burma.
F, Stegolophodon latidensref., r.M'. Sulcus on ridge-crests I-III, conelets 4-6, ridge-crests 4)2- After Matsumoto, 1926.1, PI.
V, figs.and 3 (Prostegodon), Sliiwogama, Miyagi District, Province of Rikuzen, Japan.
1

Pliocene (?) G, Stegolophodon hjdekkeri type, l.M'. Ridge-crests I-VI, conelets 4-7, sulcus on ridge-crests I and II only. After Lydekker,
1886.2, fig. 19 (as M. latidens), Borneo.
Middle(?) Pliocene A, Stegolophodon sublatidens typo, r.Ml Conelets 4-5. After Schlesinger, 1917, Taf. xvii, fig. 2, Teschen (Schlesien), Austria.
Middle Pliocene H, Stegolophodon cautleyi loetotype, I.M^. Ridge-crests I-V, sulcus on ridge-crests I and II, conelets 5. After Lydekker,
1886.1, p. XV, fig. 6, Perim Island, India
Upper(?) Pliucone I, Stegolophodon stegodontoides Pilgrim, type, r.M'. Ridge-crests I-VI, sulcus on ridge-crests I-III, conelets 5. After Lydekker,
1880, PI. XXXIX, Lehri, Punjab, India, possibly Upper Siwaliks.
CoMPAR,\TivE Observations (1935)
Molars of Stegolophodon are readily distinguished from those of the Mastodon, Zygolophodon, and Turicius phyla by the following characters:
First, by the persistence of the median stdcns separating the inner and outer pairs of cones of all the crests (A, B, D), of the three to four anterior
crests (E, F, I), of the two anterior crests (G, H).
Second, by the rounded, bunoid conelets separated by median sulcus (AH).
Third, by the closure of the enamel in the base of the transverse valleys, as seen in section (El), very characteristic of Stegodon.
The second and third of the Stegolophodon characters enumerated above link this genus with the genus Stegodon. But we must remember that
Stegolophodon cautleyi is of Middle Pliocene age (Perim Island), contemporary with the true Middle Pliocene Stegodon bombifrons (Dhok Pathan).

'[See note on page 824 above regarding the Lower Pleistocene rather than Lower Pliocene age of Stegolophodon latidens. — Editor.]
 —

THE STEGOLOPHODONTIN^: STEGOLOPHODON 851

(n'ideuce in the structure of the superior grinding teeth of the tyjies of Stegolophodon sublatide/is Schlesinger, 1917 (PI. xiii, A)
affinities of this tetralophodont type with the much more primitive and S. lalidens Glift, 1828 (PI. xiii, E) and a molar tooth from
trilophodont mastodonts of the Lower Miocene of France (cf. also Borneo described and figured by Lydekker as 'Mastodon lalidens'
Figs. 725 and 727). Clift (1885.2, 1886.1, 1886.2). This molar Professor Osborn
selected as the type of a new Pliocene species, namely, Stegolo-
Stegolophodon lydekkeri Usborn, 193G phodon lydekkeri (Vol. I, p. 700).
Figure 728, PI. xin —
Type. Third superior molar of the left side, l.JVP, original
riom vicinity of Bruni, northwest coa.st of Borneo. in the Zoological Society of London; cast in the British Museum

I
In an intensive reexamination of \arious species of Stegolo- (M.2498). Horizon and Locality.— (Lydekker, 1885.2, p.
jjhodon, Professor Osborn discovered wide differences between tire 777): "The specimen forming the subject of the present notice
was forwarded from Borneo to the Secretary of this Society by
Fig. 19.
Mr. A. H. Everett, C.M.Z.S., who stated that it was found during
the early part of the present year [1885] by a Kadayan in the
jungle in the vicinity of Bruni, on the north-west coast of Borneo."
Pliocene (?). Type Figure.— Lydekker, 1885.2, PI. xlviii,
figs. 1 and 2; .see also Lydekker, 1886.1, fig. 7, 1886.2, fig. 19, as
well as PI. xiii and figures 660 and 728 of the present Memoir.
Description. — (Lydekker, 1885.2, pp. 777-779) : "The speci-
men is the crown of the last
upper true molar of a tetralopho-
left

dont Mastodon, and agrees so closely with Indian teeth of the

Siwalik Mastodon lalidens, Clift [Footnote: 'Trans. Geol. Soc.
ser. 2, vol. ii, pt. 3, p. 371 (1828).'], that it may be safely referred
to that species, although it indicates a very small individual. . . .

It will be seen that the Borneo specimen agrees [with .1/. lalidens
Mastodvii latidens. — The third upper true molar of a small
left iiuliviclual in a
of the Punjab — made by Pilgrim in 1913 the type of Mastodon
partially-worn condition ; from the Pliocene (?) of Borneo. J. The lower
border of the figure is the inner border of the specimen. (From the stegodont aides] in the number of ridges (although the hind talon
'
Palasontologia Indica.') is considerably smaller), but is of greatly inferior size, the dimen-
Fig. 728. Stegolophodon lydekkeri Usljorn, 1936, Vohime I, p. 700, fig. sions of the two specimens being as follows, in inches:
(itjO, of the present Memoir. Third left superior mohir exhibiting six ridge- Punjab Borneo.
i-rests and aFigured by Lydekker, 1886.2, fig. 19, as Mastodon lalidens.
talon.
Extreme length 8.6 6.3
Compare Lydekker, 188o.2, PI. xLviii, also PI. xni of the present Memoir.
Two-thirds natiwal size. Width of first ridge 4.2 2.95"]
 Fig. 729. Falconer's map of the geology of India (see "PahEontological Memoirs,"
1868, Vol. I, description of PI. ii). The red fossiliferous areas, here colored black, are de-
scribed by Falconer as follows;
"The red [black] stripe represents the Sewalik Hills, stretching from the Hydaspes to
the Gundiick River, 800 miles. The small red [black] patch behind the Himalayahs repre-
sents the ossiferous plain of Tibet about 16,000 feet above the sea. The other red [black]
patches represent the Nerbudda [Pleistocene], and the [black dot within the circle the] Gulf
of Cambay [Pcrim Island-Middle Pliocene] fos.sil tracts."
The following is Falconer's interpretation of the geology of India:
"The great mass of light shadin/] represents the sujjposed insular form of the contineni
of India at an early period of the Tertiary epoch, the island forming a sort of triangle, of
which the eastern and western Ghats formed the sides and the great Vindhya range the base,
with an irregular patch of mountainous country stretching north forming the Aravalli range."
"The dark shading rejjresents the ))lains of India, forming the valley systems of the
Ganges and Indus drainage, which were formerly narrow ocean straits. These straits were
the n'cipients of the silt and alluviiun washed out of the Himalayahs, and were at length
elevated above the ,sea, .so as to form th(^ existing continent. The Sewalik Fauna (hen spread
over the continent, from the mouth of the Irrawaddi to the Gulf of Cambay 2,000 miles,
and north to the .Iheluin 1,.')00 miles, .\fter the long establishim'nt of the Sewalik I'auna,
a great upheavemcnt took place along the line of the Himalayahs, elevating a narrow bell of
the plains into the Sewalik Hills, and adding many thousand feet to the height of the
Himalayahs."

852
 IV. SUCCESSION OF SPECIES OF THE GENUS STEGODON
Superfamily: STEGODONTOIDEA Osborn, 1935, 1936
Family: STE(iODONTID^ Young-Hop wood, 1935
Subfamily: Stegodontin^ Osborn, 1918, 1921

Genus: STEGODON Falconer and Cautley, 1847, 1857

Original reference: Falconer and Cautley, "Fauna Antiqua Sivalensis," 1846 [1847, PL xlii]; also Falconer, Quart. Journ.
Geol. Soc. London, 1857, Vol. XIII, pp. 314, 318, and Synoptical Table opposite page 319.
Genotypic species: Elephas Cliflii, E. bombifrons, E. IGanesa, E. insignis}
Syn.: i^j/iTMenodott Cope, 1889; Parasfejodon JNIatsumoto, 1924 (in part).

—
Generic Definition. (Falconer, 1857, p. 318): "Dentiuni molarium 3 utrinque intermedioruni
eoronis complicata colliculis hypisonieris {e.g. 7 + +
7 8), mammillatis, tectiformibus. Praemolares non-
dum observati."
—
Generic Characters. Ridge-crests intermediate between Stegolophodon and Archidiskodon
planifrons types, progressively multiplying from six to eleven in the intermediate molars, from nine to
fifteen and a half in the posterior molars. Cones rapidly subdividing by binary or ternary fission into
multiple conelets. Ridge-crests elevating from brachyodont {Stegodon sinensis) to subhypsodont {S.
airatrdna and S. aurorx stages). Cement developing in the valleys. Crania of mastodontoid {S.
bombifrons) to extremely abbreviated, female? {S. insignis), more elongated, male? (S. ganesa), more
triangular (*S. trigonocephalus) form. Tusks attaining" great dimensions {S. ganesa). Phylum parallel
to that of the true Archidiskodon and Elephas, not directly ancestral, readily distinguished by cranial
and dental characters.
Coniparc figuics 820
The generic name Stegodon was first prnited Fig. 730. After Osboni, 1910.346, i). 323, fig. 154. 72i) ;iihI

of present Memoir.
in tlie "Fauna Antiqua Sivalensis" of Falconer

and Cautley, 1846 [1847, PI. xlii]. It was ten

years later that it appeared as a subgenus (Fal-
coner, 1857, pp. 314, 318, table opp. p. 319) to
include the following species, Elephas cliftii, E.
bombifrons, E. ganesa{?), and E. insignis, and
has subsequently been used as a genus by all the
principal authors, except Lydekker who clung
to the Cuvierian division of the mastodonts and
elephants into the two genera, Mastodon and
Elephas. Pohlig in 1888, p. 252, wrote the genus
in this way: Stego(lopho)don. Schlesinger, 1917,

p. 115, separated the species M. latidens as the

type of a new genus Stegolophodon.

Stegodon (omitting the six^ species described

above under Stegolophodon) is readily definable _
chief Miocene and PUocene fossU mammal depoaits of Asia. 1. Maragha,
^<'^^'^- Perim Island. 3. Manchhar Beds of Sind. 4. Siwaliks of the Punjab. 5. Sub-
2.
ind ILdUay
auu di^tintriii^liihlp from
rpfldilv UlSliagUlSnaUie liom any
«inv genus nf
o-pnii'^ Ul
himalayan Siwallks (River Brahmaputra to River Jhelum). 6. Valley of the Lower Irawadi,
Burma. 7. Miocene and Pliocene deposits of China (Provinces of Shan-si, Shen-si, Sze-chuan,
thp Mn**tnflnntirlfP on me
ine iViaSlOUOmiUcB, nnp nana,
tliP one linnrl or or thp
nf tne
Kwang-Tung, Ho-nan, Hu-nan, Hu-peh). 8. Miocene and PUocene deposits of Japan.
Elephantidae, on the other (see Vol. I, p. 25), by
the brachyodonty to subhypsodonty of its ridge-crests, in contrast to the hypsodonty in all the species of the
Elephas and Loxodonta phyla. The relative height attained is shown in figures 687 and 688. While the ridge-
HHopwood in his Memoir of 1935 on the "Fossil Proboscidea from China," p. 72, remarks: "According to the modern school of priority-purists this
necessitates the selection of E. cliflii as genotyjie as a matter of course. Fortunately, we can do so and adhere to the author's original intention, for, in his
later writings Falconer expressly says that Stegodon corresponds 'with the forms collectively designated Mastodon elcphantoides by Cliff, (1857, p. 314)."
—Editor.]
-[To these six species should be added the Stegolophodon lydekkeri Osborn, described in Volume I of the present Memoir (p. 700). — Editor.]
853
854 OSBORN: THE PROBOSCIDEA

crests are composed exactly as we know that we shall find them hi the direct ancestors of Elephas, they

remain short crowned, even in the progressive species Stegodon insignis and ;S. ganesa, and in the still more pro-
gressive S. orientaUs and .S. airdwana. This is undoubtedly an adaptation to browsing on leaves and softer
kinds of food, which leads us to believe that the Stegodonts were persistent browsers rather than grazers, as in
all the i^liyla of Elcplnts and of Loxodonta.

Table V. Si>i;cn;s ix Approximate Ascending Order of Collective Maximum and Minimim l\iiKii:-ci(EsT«

l'"alconer (1868), Lydekker (1886), .Martin (1890), ^Nlatsumoto (1918), Osborn (1929)
 THE STEGODONTIN^: STEGODON 855

The order of ridge-crest addition and development (Table V) corresponds approximately with the phylogenetic
and geologic ascending order as shown in Table IV above of the present chapter, in which the species are grouped
partly by geographic distribution. The ridge-crest formulae of certain species, e.g., Stegodon sinensis, S. ganesa
javanicus, S. orientalis, and S. orientalis shodoensis, are too imperfectly known to determine precisely their phylo-
genetic position. In general, S. sinensis appears to be the most primitive, while S. airdwana appears to be the
most progressive and geologically recent.

STEGODON GANE5A 3050min..lO'e STEGODON PINJORENSIS 2745mm.. 9 e

INDIA SIWALIKS INDIA SIWALIKS

STEGODOrJ INSlCrjlS I925mni.,0'4''8"6

INDIA SIWALIKS
STEGODON ORANGERI 225&i;in;.,7'5"e
CHINA

STEGODON TR1G0N0CEPHA,LUS I428mm„4'8>i"e

JAVA

STEGODON BOMBIFRONS
INDIA
2148
SIWALIKS
mm.. 7'>t"e
^m
STEGODON AIRAWANA
JAVA
76min..3'lO/3"e

Fig. 731. Species of Stegodon from India, Chin.\, and Java

Restorations by Maboret Flinsch, under the direction of Henry Fairfield Osborn, One oNE-HU>fDRBDTH Natur.^l Size

Skulls of Stegodon bombifrons, S. insignis, and S.ganesa in the British and Indian Museums
Before considering in detail the succeeding species of Stegodon, it is necessary to examine and compare the
known crania of various species with each other, as assembled in figure 732 from Falconer and Cautley's beautiful
plates, with the crania of other Stegodonts from the East Indies, e.g., Stegodon airdwana (Fig. 773) and .S. trigono-
 ELtPHAS eOMBIFRONS B*»,
PI. XLIII, XIII

ELEPHAS GANESA R»f.

Pi, XXir, F.Q. 2

I'lK- 732. The Strgodonts of wliirh (ho cninia wcrt' known to Falconer iiro illiislratcd in tliis comparative plate in
tlircr sprrios of
such a manner may be readily contrasted. TlirouKliout the Stegmlon crania are of relatively small size,
tliat their distinctive eliaraeters

laekiiiK the cancellate structure characteristic of EUphas. All one-twentieth natural size. Compare figure 777.
Elii>lms = Slrgodon] bombifrons, a Middle [to Ujiper] Flioceni- stage of evolution, is illustrated in the lower line. [The cranium of E.
[

Immbijnms at extreme right should read: PI. xr,v, xui (rev.).|

Elrphns [=Strgo<ltm] in.iignis, a Ixnvpr to Middle |Up|)er] Pleistocene stage of evolution, is illustrated in the middle of the diagram.
Elephas =Stcgodon\ gancsa, a Lower to Middle [Upper] Pleistocene stage of evolution, is illustrated in the top lines. It is extraordi-
[

nary that tusks of such enormous length and width should be supported by a cranium of such small size. The extremities of the
tusks should probably be turned inward, instead of outward as figured by Falconer and Cautley; the left tusk is complete; a portion
of the right tuskis omitted in this drawing. [Compare figure 733 for revised ri'storation of the tusks.)

856
 THE STEGODONTIN^: STEGODON 85

cephalus (Fig. 776) of Java, also with the newly found *S'. orientaUs grangeri (Fig. 763) of China. These Stegodont
crania should be compared with the large craiiiuni of AirliidiHkddoii pUutiJron^ (Fig. 830), one of the most primitive
of the true elephants.

In general the cranium of Stegodon bombifrons (Fig. 732, bottom row, also Fig. 742 and Fig. 744) is more
generalized and subelephantine in character, whereas the crania of Stegodon ganesa and S. insignis are very highly
specialized, of relatively small size, and bear little resemblance to the crania of the true Elephantidae. Quite
different are the small, triangular crania of Stegodon trigonocephalus and S. airdunna of the East Indies.
Sexual Divergence very Marked. The referred — giant skull of Stegodon ganesa obviously belongs to
a full-grown male Stegodont (as shown in fom- as]>ects in Fig. 732). On the contrary, the crania referred to S.

Fig. 733. Cianium of Elephas [Stegodon] ganesa, after skull referred to this species by Falconer. Drawings made from original plates in Falconer and
Cautley's "Fauna Antiqua Sivalensis" reduced to one-sixteenth natural size. See figure 732 opposite for frontal, lateral, and palatal views, reproduced to a one-
twentieth scale. The frontal profile is seen to differ profoundly from that of Stegodon insignis and less profoundly from that of S. bombifrons.
In an attempted restoration of Stegodon ganesa from the tusks as originally represented by Falconer and Cautley, it was found impossible to lower the
it was also observed that the extremities of the tusks turned outward, unlike all other proboscideans.
proboscis between the closely appressed tnsks; Accordingly
on receipt of the gift of the cast from the British Museum (Jan. 23, 1931) the tusks were readjustctl in a position with the extremities turned inward, allowing
a sufficient space for the descent of the proboscis between the butts. The present figure rcjirescnts the specimen according to this conception of the position
of the tusks, in contrast (Fig. 732) with Falconer and Cautley's restoration.
[The most painstaking examination of the original sijecimen by Professor Osborn, Doctor Gregory, and Doctor Colbert failed to reveal any evidence
cither tliat the tusks have been transposed or tliat they are twisted in the alveolus by ijost-mortem changes. Nevertheless all felt that the space between them
is insufficient for the trunk as restored by Falconer and Cautley and that the whole arrangement looks abnormal. — Editor.]
858 OSBORN: THE PROBOSCIDEA

insignis (Fig. 732) obviously belong to small-tusked females, because the alveolar processes for the insertion of the
tusks are extremely small ami narrow.

If, as seems possible, all the crania referred by Falconer and Cautley to Elephas [
= Stegodon] insignis are
females, and the great cranium referred to Elephas [
= Stegodon] ganesa is a male, there is in Stegodon a far greater
sexual disparity and difference than prevails between the female and male crania of either Elephas indicus or
Loxodonta africana, as figured below in the present Memoir. If this be true, the sexual disparity in cranial charac-
ters constitutes an important specific distinction of Stegodon insignis and ;S. ganesa.

In the present Memoir we two species separately but agree with the theory suggested by more than
treat the

one author, especially Lydekker and Matsumoto, that the crania of S. insignis represent the females of the same
collective species as the referred male cranium of S. ganesa.

British Museum, W. D. Matthew, September, 1920. —The skull of Stegodon bombifrom is essentially
elephantine, the shortness of the enamel plates being the chief distinction; the skull is apparently shorter than in
Loxodonta; the jaw is deeper; the symphyseal process may be a little heavier. In contrast, the skull of S.
insignis has a supranarial region with a great thickening of the cellular tissue which appears to round back into
the occiput, the occipital crest being very little developed; a very curiously shaped head and very small tusks.

Characters of Referred Skulls of Indian Stegodonts, x

after Falconer, 1868 - -

(See Figs. 732-736, 754, 777)

Skull of Elephas [
= Stegodon] bombifrons (Falconer, 1868, Vol. I, p.

458, PI. xxvii): "Very fine and perfect skull, anterior view. Four other
views of same skull are given in Plate xxviii. This head is very marked;
it is convex from occiput to front and also across, and is very narrow
at the temporal contraction. The bounding ridges sweep round by a

a Generalized Cranium A Specialized Cranium A Specialized Male Cranium

Fig. 734. Elephas [= Stegodon] bombi- Fig. 735. Elephas [=Slegott(m\ insig- Fig. 736. Re.storation: Elephas
frons. Front view of cranium, oiic-twclfth nis, one-twelfth natural size. After [= Stegodon] gnnesn, one-twelfth natural
natural size. Brit. Mus. M.2979; cast Falconer and Cautley, 184611847, PI. xv). .size. Brit. Mus. M.3008. After Falconer
.Vmer. Mus. 1037S. .\fter Falconer and and Cautley, 1846 [1847, PI. xxiri].
Cautley, 1840 (1847, PI. xxvii].
 — —

THE STEGODONTIN^: STEGODON 859

l)old curve into the post-orbitary processes, as in E. [

= Archidiskodon] meridionalis. There is a deep furrow
between the tusks. The nasal opening for the trunk is above the Hne (or nearly so) of the post-orbitary processes

of the frontal bone. Above the infra-orbitary foramen on the right side there is another smaller opening."

Skulls of Elephas [
= Stegodon\ insignis (Falconer, 1868, Vol. I, p. 448, PI. xv) : "This is the most remarkable
of all the Indian fossil Elephants. The cranium is as singular and grotesque in construction as that of the Dino-
fherium giganteum. The cranium is seen to differ remarkably from that of E. Ganesa (Plates xxi. and xxii.)
notwithstanding that the molars of the two species agree so closely. That of E. insignis is flattened at the top,
elongated from side to side and singularly modified, so as to bear an analogy to the cranium of Dinotherium
giganteum, while that of E. Ganesa does not differ much from the ordinary type of the Elephants."

(Op. cit., p. 449, PI. XVI, fig. 1) : "This head is very cubical in form, is old, very concave in front and vertically;
teeth broken. Interval between incisive sheaths deep. No tusks."

{Op. cit., p. 449, PI. XVII, figs. 3 and 4): "Anterior and lateral view of another cranium. Both zygomatic
arches are missing, and the left side of the cranium is deficient. Shows the great length of the incisive sheaths."

Skulls oi Elephas [
= Stegodon] ganesa (Falconer, 1868, Vol. I, p. 453, PL xxi): "Large skull, with fragment
of left incisive in situ, and corresponding fragment of right incisive detached. The incisive alveoli are remark-
ably elongated, as in E. primigenius. The plane of the incisives is continuous with that of the frontal, but with a
tendency to obliquity forwards. The skull is very imperfect on right side."

{Op. cit., p. 454, PL xxii) : "Fig. 1.—Elephas Ganesa. Lateral view of large skull figured in PL xxi. —B.M.
Fig. 2. —E. Ganesa. Palate view of same skull. The right incisive is seen in section. The posterior true molar
is seen on either side of palate. It has ten plates and a heel behind, and a small talon in front; the hind heel has
few denticles; the foiu' front ridges are worn. The alveoli are parallel as in the Mammoth. —B.M. Fig. 3.

E. Ganesa. Sketch showing restoration of skull, with tusks, of iJ.Gawesa, profile view, one-thirteenth of natural size."

{Op. cit., p. 454, PL xxiii): Restoration. "Sketch showing restoration of skull, with tusks, of E. Ganesa,
oblique antero-lateral view, one-thirteenth of natural size."

{Op. cit., p. 455, PL XXIV. A. ) : "Figs. 1 and la. Elephas Ganesa. Fragment of skull with palate and back
molars on both sides. This is a most remarkable specimen. I have called it E. Ganesa (H. F.), and it much
resembles the molar of the big Ganesa specimen (Plate xxii. fig. 2) in form and in the compression of the ridges,

but the ridges are few. . . . B.M."

Stegodont Crania of China and of the East Indies

(See Figs. 763, 776, 773, 777)
A most fortunate discovery from the pits along the Yangtze River near Wanhsien, Province of Szechuan,
China, is the extensive series of Stegodont crania in all stages of development, infantile, juvenile, young adult,
and mature adult. This priceless collection made by Walter Granger of the Central Asiatic Expedition of the

American Museum illustrates in a perfect manner (Figs. 759, 761-763, 686) the complete ontogeny, dental
succession, and metamorphosis of the cranial form. The young adult and mature adult crania, together with
the inferior mandible (Fig. 763), apparently resemble the cranium of S. bombifrons more closely than the crania

of S. insignis and S. ganesa. This material was described by the present author as belonging to a new subspecies,
namely, Stegodon orientalis grangeri.

The cranium of the East Indian species Stegodon airdwana (Fig. 773) of Java belongs to a very advanced
mid-Pleistocene stage, resembling *S'. bombifrons and *S. orientalis grangeri in profile but differing in the frontal
aspect, which is flattened. The cranium of S. trigonocephalus (Fig. 776), as its specific name indicates, has a
 — — —

860 OSBORN: THE PROBOSCIDEA

triangular rather than a rounded superior profile, and, althougli of the same geologic age as S. insignis and *S.

ganesa, it is entirely different from the Siwalik species in its profile and proportions. Consequently it appears
that both in cranial and dental characters the East Indian species represent a distinct and somewhat dwarfed
side-brancli of tlie northerly continental species.

SYSTEAIATIC DESCRIPTION OF SPECIES OF STl'^GODON

Stegodon sinensis Owen, ISTO parison is strengthened by the fact that Koken (1885) referred
Figures 087, 702, 737 the Stegodon sinensis of Owen to the species Stegodon rliflii of

Alleged to be "from marly beds in the vieinity of Shangiiai," China. Falconer.

Tliis animal (Figs. 687, 702) is probably of Upper Miocene [Lower Pliocene) (3) Lydekker (1886.2, p. 79) also referred S. sinensis Owen to
age, for it is more primitive than Slegodun bombi/rons, and possibly ancestral Elephas cliftii Falc. {Op. cit., p. 80) : Brit. Mus. 41925. "A third
to the S. bomhijrons stage.
right upper milk-molar, provisionally referred to this species; from
—
Specific Characters. This little-known true Stegodont, the Pliocene near Shanghai, China. This specimen is the type of
Stegodon sinensis (Dp 3 ^^), is somewhat more primitive than Stegodon sineiisis, and is described and figured under that name by
S. homhifroiis (Dp 3-), since the ridge-crests are less elevated or Owen in the Quart. Journ. (!eol. Soc. vol. xxvi. p. 417, pi. xxvii.;
hypsodont as compared (Figs. 687, 688) with S. bombifrons, it is also figured and provisionally referred to the present species
S. orienialis graiigeri, and S. insignis. Estimated number of cone- [Elephas by the writer, in the 'Palseontologia Indica,' ser.
cliftii]

lets on fourth ridge thirteen to fifteen. Falconer, Lydekker, and 10, vol. i. p. 257, pi. xlv. fig. 2. There are four complete ridges, and
Martin ascribe (see Table \ above) four ridge-crests to Dp' of S. a large anterior talon, which is reckoned by Owen as a fifth ridge.
elephantoides { = cliftii), S. bombifrons, and S. trigonocephalus. The median longitudinal cleft is very indistinct. Presented by
—
HiSTOHY. (1) After comjiaring the type deciduous molar Prof. Sir R. Ou'en, K.C.B., 1870."
with all the Siwalik specimens in the British Museum, Owen (4) Osborn (1924) prefers to retain the name Stegodon sinensis
concluded (1870, p. 420) that the above Chinese tooth was most until further local material can be secured for comparison.

Fig. 737. Type of Stegodon sinensis Owen, 1870, PI. xxvil, figs. 1 and 2, natural size. Alleged to be from "marly
beds" near Shanghai, China. (Op. cit., p. 417) "The tooth in question is the second upper molar ((/3 of the type series)
:

from the right side. Its crown, in a length of three inches, is divided into five transverse ridges."
Falconer, Lydekker, and Martin (Taljle V above) ascribe four ridge-crests to Dp' of Stegodon elephantoides
(= cliftii), S. bombifrons, and S. trigonocephalus.

closely related to undetermined Siwalik specimens which he com- Stegodon sinensis Ow-en, 1870. "On Fossil Remains of
pared with the M. elephantoides of Clift. These two undeter- Mammals found in China." Quart. .lourn. Geol. Soc. London,
mined specimens are described by Falconer (1868, Vol. I, p. 460) Vol. XXVI, p. 417. Type.— (Op.
cit., p. 417): ". Second . .

as follows: Plate xxix. a, "Fig. 5. E. bombifron.sl Fragment of upper molar {d 3 from the right side [Dp']."
of the type series)
molar, from lower jaw, right side, with four ridges. — B.M. Length, Brit. Mus. 41925. Horizon and Locality. Alleged to be —
5.8 in. Width, 4.5 in. Fig. 6. E. bojtibifronsl Fragment of molar "from marly beds in the vicinity of Shanghai," China. Probably
with three ridges and a heel. 'Doubtful what figs. 5 and 6 are.' tapper Miocene [Lower? Pliocene]. Type Figure. Op. cit., —
H. F. Length, 4.4. in. Width, 4.5 in." This shows that Falconer PI. xxvii, figs. 1-3.
regarded these Siwalik teeth as doubtfully related to E. bombifrons Type Description. (Owen, op. cit.,— pp. 417, 418) : "Stegodon
and that Owen also indirectly comjjared f hem with E. bombifrons. sinensis, Ow. The tooth in question is the second upper molar
Stegodon bombifrons lectotype is of Middle Pliocene age. (d 3 of the type series) from the right side. Its crown, in a length of
(2) One should also compare Owen's type tooth with figuie 700 three inches, is divided into five transver.se ridges, the jiroportions
above of Mastodon [Stegodon] elephantoides (=cliftii}. Tiiis coni- of which, as to height and basal breadth, with the ridged and
 THE STEGODONTINiE: STEGODON 861

wrinkled character of the enamel, suffice for its reference to a Stegodon elephantoides ('lift, 1828
species of the group of Proboscidians discovered by C'rawfurd in Figures 683, 686, 695, 696, 700, 701, 738, 739 741
the Irrawadi Tcrtiaries of Ava, and described by ('lift in the Lower PlioceiU', lowest levels of the Irrawaddy Series (fluviatile),' near
second volume of the second series of the Transactions of the Yenangyaung, 250 miles below Ava, Burma.
Geological Society (p. 369, pis. 36-39, 1828). ... In the present Syn.: Elephas cliftii Falconer, 1846 [ = StegodoH elephantoides (=cliftii)
of the present Memoir.]
tooth the first or foremost ridge (PI. xxvii. figs. 1 & 2, 1) is defined
by a cleft on the outer side of the tooth, but not on the inner side, Specific Characters (Clift, 1828, Osborn, 1929). Third —
fig. 3; here the abraded surfaces or ridges and 2 are blended by1 inferior molar, M3, with nine complete ridge-crests and a well-
wear into a common hollow field of smooth dentine (fig. 1, a). developed half ridge, equaling ten; five to eight conelets on each
There is a slight constriction near the part where the worn surface ridge-crest; length 11 inches = 280 mm., est. breadth Sji inches =
of the first ridge blends with that of the second and this constric- ; 90 mm. no apparent cement. Superior molar, l.M^ ( = Falconer's
;

f ion, which may be detected in the succeeding ridges, I take to be type of Elephus cliftii) with six and a fourth ridge-crests, length
a tiace of that stronger one which more completely divides the 155 mm., breadth 83 mm., ten to twelve conelets on each; cement
in the bottom of the valleys. Palate with l.M^, r.M^, with six and
a fourth ridge-crests; length of l.M^, 186 mm., breadth 102 mm.;
riilge-crests worn; traces of cement. From the three specimens

iM a.ij,^ Uiktj Jktial fy

CS^Un-M-Jti

Mus
T^totJ. by /Cnyrfijirei ZsaJ SSi/ Mdt-; p^aw AwiJ.

Lectotype of Stegodon elephantoides

Fig. 738. Lectotype left Ms, M3, of Mastodon elephantoides Clift, 1828, PI. xxxvni, fig. 2, one-third natural size-
{Op. cit.,p. 372 and Explanation of Plates): "Left side of the lower jaw of Mastodon elephantoides. The remains of
the anterior molar tooth [l.Mj] are seen, and behind it, the posterior tooth which was advancing, and which, in
consequence of the jaw-bone being broken away, is seen through its whole length. This tooth is eleven inches long and
three and a half broad."

transverse coronal ridge in the molars of better Mastodons into enumerated below the lectotype and cotype ridge formula is

an inner and an outer part. A well marked tubercle (figs. 1 and compiled as follows: M 1^ M 2^ M 3rs-
2,/) projects at the outer side of the base of the first ridge, 1, Osborn, 1927: This Stegodont is siinilar to S. bonibifrons in
near the interspace between that and the second ridge. Never- . . . ridge-crest formula.
theless in the number of ridges in a given tract of the grinding- History. — As fully explained above (pp. 855, 856), Stegodon
surface, in their height and breadth of base, and in the absence of elephantoides Clift is the second species of Stegodont based by Clift
intervening cement, the conformity of the Chinese molar with the on a lower jaw (PI.xxxviii, fig. 2) and on an upper molar (PI.
grinders of the Mastodon elephantoides The enamel also is close. XXXIX, fig. 6). Unfortunately Falconer was led to abandon
shows the same vertical linear impressions and ridges, by which 'Mastodon elephantoides' and to substitute 'Elephas cliftii'; conse-
we may reckon that the summit (say, of the fourth ridge in the quently all the literature subsequent to Cliff's original description

tooth here described), if it were unworn, might be cleft into from and most of the reproductions of his illustrations appear under the
thirteen to fifteen small mamillse." Owen did not compare this specific name Elephas cliftii, which is actually a synonym of
specimen with Cliff's original type lower molar of Mastodon Stegodon elephantoides. Falconer's mistake partly arose through
elephantoides, but with specimens referred by Clift to that species. Cliffs error in entitling the palate of Mastodon [
= Stegodon]

'[See note on page 824 above regarding the Lower Pleistocene age of Stegodon elephanlnides. — Editor.)
862 OSBORX: THE PROBOSCIDEA

elephantoides (figured in PI. xxxvi) as 'Maatodon latidens.' cene, lowest levels of the Irrawaddy Series (fluviatile).'- LiccTO-
Consequently ('lift's species M. [
= Slpgodo)i] rlephnntnidcs rests TYPE Figure.— Clift, 1828, PI. xxxviii, fig. 2. Cotype
upon three specimens: Figure.— Clift, 1828, PI. xxxix, fig. 6 = !<t4'gudoH elephantoides [

{ = dijtii).\ Referred Palate.— Clift,

1828, PI. xxxvi.
Lectotype: "Left lower jaw, Mastodon Pllephanloides" (C'lift,
Clift's Original Description of 'Mastodon' elephan-
1828, PI. xxxviii, 2 = figures 696 and 738 of the
present Memoir).
fig.
toides. —
We refer to Clift's clear and consistent description
quoted in full above (p. 827) together with the beautiful lectotype
CotjTie: First left superior molar, l.M\ "Upper molar of M.
figure reproduced in our figure 738 herewith. Clift's specimen
Elephanloide.^" (C'lift, 1828, PI. xxxix, fig. 6 = figures
comes from Burma and his figureagrees exactly with his descrip-
700, 739, and 740 of the present Memoir), .sub-
sequently made the type of Elephas rliflii by
tion, namely, M 3to, with five to eight conelets on each ridge.

Falconer and Cautley.

Fig. 2a
Referred: Palate, with l.M- and r.lSP, "Upper jaw of Mastodon
latidens" (C'lift, 1828, PI. xxxvi), not figured in the
present Memoir.

Mastodon elephantoides Clift, 1828. "On the Fossil Remains

of two New Species of Mastodon, and of other vertebrated Animals,

Cotype of Stegodon elephantoides Clift (=cliftii Falconer)

Fig. 739. New figure of type (l.M') of Elephas cliflii Cotype l.M' of Stegodon elephantoides
Fig. 740.
Falconer and Cautley, 1846, first figured by Clift, 1828, ( = dijlii), and Cautley, 184(1 [IS47,
after Falconer
PL XXXIX,fig. 6, as an "t'pper molar of M. Elephantoides." PI. XXX, figs. 2, 2a], loft M', said to he the same

Reproduced herewith from cast (Amer. Mus. Warren molar (see Pal. Mem., 1868, Vol. I, j). 461) as that
Coll. 10382) one-half natural size. Original in British figured in Clift, 1828, PI. xxxix, fig. 6, although
Museum (Brit. Mus. M. 10520). From near Yenangj'aung, the drawing does not agree well with Clift's original
Burma. illustration nor with the reproduction from the
cast (Fig. 739 opposite).

Observe rudimentary anterior ridge (pro-protoloph) and rudimentary seventh ridge (heptaloph), also the six ridge-
and hexalophs), the ridge-crest summits each crowned with from ten to
crests (proto-, meta-, trito-, tetarto-, penta-,
twelve conelets.

found on the left Bank of the Irawadi." Trans, (ieol. Soc, COTYPE CH.\R.\CTERS OF STEGODON ELEPH.\NTOIDES
London, (2), II, Pt. Ill, 1828, pp. 372, 373. Lectotype.— (1) = CLIFTII) FALCONER AND CAUTLEY, 1846, PAGE 47,
(

Left side of lower jaw with Mj, M3; original in Museum of

BASED ON CLIFT'S SPECIMEN AND FIGURE
Geological Society of London; cast Brit. Mus. 7393 [referred by Falconer's designation of this type ('^left M') and of this
Falconer and Cautley in the "Fauna Antiqua Sivalensis," I'l. species is "The same group comprises a fourth e.xtinct
as follows:
XX, figs. 9, 9a, PI. XX. A, fig. 6, to E. insignis].' ("otype. — Indian species, named in this work, E. Cliflii, which furnishes the
(2) First upper molar of the left side, l.M' (original in the British next link in the chain of forms presented by the molars of the
Museum (Brit. Mus. M. 10520), cast Amer. Mus. Warren Coll. Elci)liantidie. ... In our view, the tooth represented in pi. 39,

10382) ;same tooth was selected by Falconer and Cautley as the fig. 6, of Mr. Clift's memoir in the Geological Transactions [Clift,

type of Elephas cliftii. Referred (Osborn). Palate with — 1828(, under the name of Mastodon Elephantoides, and the palate
l.M- and r.M- (figured as .Mastodon hUidens by Clift). llou- .specimen represented in pi. 36 of the same memoir, under the name
izoN and —
Locality. Left bank of the Irrawaddy River, of M. latidens, belong to this species. The penultimate and . . .

near Yanangj^iung, 250 miles below Ava, Burma. Lower Plio- antepenultimate molars in the upper jaw have onlj' six transverse

'[See Chakravaiti, D. K., Quart. .Tourn. Geol., Mining, Metalluig. .Soc. India, 1937.1, p. 34, who referred it to .S'. f;»7)/ia»i(oirfos.— Editor.]

-[See note on page 824 above regarding the Lower Pleistocene age of Stegodon elephantoides. — Editor.)
 :

THE STEGODONTINiE: STEGODON 863

ridges, continuous, and chevron shaped, with numerous mammil- PHYLETIC CONCLUSIONS
lae, as in K. insignis and E. Gnm-sn; hut the cement does not fill Osborn, 1927: Falconer considered Cliffs Lower Pliocene-
up the interspaces of the ridges, being reduced to a comparatively type of Mastodon [
= Stegodon] elephantoides as close to his Upper
inconsiderable quantity in the bottom of the hollows. E. Clijiii, Pliocene S. insignia, but S. elephantoides proves rather to be close
in the reduced number of the coronal ridges, and in the other to the Middle Pliocene S. bombifrons, as shown in the comparative
characters of the teeth, appears to constitute the dental link which ridge formulae table above, the two formulae abbreviated being as
forms the immediate passage from Elephas into Mastodon." follows
S. bombifrons: Dp 3i Dp 4 ^^ M 1 ^^^ M 2 i M 3" It
ADDITIONAL CHARACTERS OF THE COTYPE SPECIMEN OF S. elephantoides: Dp 3| Dp 4J M 1 ^* M 2 -p ^"
M "3 10- 1
STEGODON ELEPHANTOIDES = CLIFTII) (
Elephas = Stegodon elephantoides
cliftii [ (
=
cliftii)], fide Falc.
"But the detached tooth
(Falconer, 1868, Vol. II, p. 84):
and Caut. : Dp 3 } Dp 4 1 M 1 -^ M 2 ^- M 3 »«•

[Fig. 740] on the upper jaw is seen entire, and beautifully preserv-
ed, in the specimen fig. 2 of the same plate [i.e., PI. xxx], presenting Hg.l.
six ridges and a small hind talon. The same tooth is represented by
fig. 6 of PI. XXXIX. of Mr. Cliffs Memoir (Geol. Trans., vol. ii.

2nd series). It is there described as an upper molar tooth of

Mastodon Elephanto'ides, under which title Mr. Clift included
specimens that are referred in our arrangement to two distinct
forms. The Elephantine affinities of this tooth are indicated
. . .

by the absence of a longitudinal line of division along the crown,

and by the great number of points (about eleven in each) that enter
into the composition of the ridges." Of the same cotype molar,
Lydekker (1886, p. 81) observes: "7388. Cast of the first (?) left
upper true molar in an early stage of wear (woodcut, fig. 20).
The original was obtained near Yenankhoung, on the left bank of
the Irawadi in Upper Burma, by Crawfurd in 1826, and is pre-
served in the Museum of the Geological Society;'" There is . . .

scarcely any trace of the median cleft, the cement is slight, and
there are numerous cusps. Mantell Collection. Purchased, 1836."

FALCONER'S NOTES OF 1868 ON STEGODON ELEPHANTOIDES

( = CLIFTII) Referred Stegodon elephantoides (= cliftii)
Fig. 741. Referred I.M3 of Elephas cliftii, after
Falconer, "Palaeontological Memoirs," Vol. I, 1868, pp. 461, 462, Plate
Falconer, 1868, Vol. II, PI. v, figs. 1, 2, "Views in plan and
xxx of the "Fauna Antiqua Sivalensis."
profile of last true molar, lower jaw, left side, rather less
It is very important to observe that the four specimens
than one-fourth (two-ninths) of the natural size." See also
described by Falconer, including the cotype of .S. elephantoides
the "Fauna Antiqua Sivalensis," Falconer and Cautley,
from Burma, agree with each other both in the number and char- 1846 [1847, PI. xxx, figs. 5, 5o]. From Burma, presented to
acter of the ridges and in the number of conelets on each ridge. the British Museum by Colonel Burney (Brit. Mus. 14759).

—
Upper Jaws. Burma, left bank of the Irrawaddy River, 250
miles below Ava. Plate xxx, figs. 1, la, E. cliftii F. & C, palate. The ridge formula of I.M3, namely, i^, assigned by Falconer
Dp*, ? ridges; Burma, 250 mUes below Ava,
figs. 2, 2a, type,'" and Lydekker, actually occurs inan aged specimen from Burma,
l.^U, and a small
ridges &%, beautifully preserved, "six ridges Falconer's figure of which (Falconer and Cautley, 1846 [1847, PI.

hind talon ... as many as eleven to twelve denticles. ... Its xxx, figs. 5, reproduced herewith in our figure 741; this
5a] is

elephantine affinities are indicated by the great number of . . .

grinder is so old that the two anterior ridge-crests may have worn
points [denticles]"; superb palate, M^, ridges 6K, little
fig. 3, off, consequently the ridge formula is uncertain; the dimensions

cement (referred by Chft, 1828, PL xxxvi, to M. latidens); figs. (length 12.7 in. 4.5 in. = 115 mm.) considerably
=323 mm., breadth
4, 4a, 46, fragment of M-, right side, 5 ridges, cement moderate in exceed those of the type of Stegodon elephantoides given above.

quantity, from near Yenangyaung, upper Burma. Lower There is no substantial basis, therefore, for the assignment to the

Jaws.— Burma. Plate xxx, figs. 5, 5a, I.M3, "eight ridges and synonymous Elephas = Stegodon] cliftii of a ridge formula inferior
[

a talon," little cement. to that of Mastodon [

= Stegodon] elephantoides Clift.

Cotype Characters. —From the above it appears that the

four specimens described by Falconer from near Yenangyaung, Stegodon bombifrons Falconer and Cautley, 1846
250 miles below Ava, Burma, exhibit the following characters: (1) Figures 686, 687, 699, 731, 732, 734, 742-746, 777, PI. xx
Denticles very numerous, eleven to twelve on each ridge ; (2) ridges Lectotyi)e; Siwalik probably from the Dhok
Hills (Jide Falconer),
moderately high with little cement; (3) ridge formula as follows: Pathan horizon, Lower [Middle] Pliocene, India, in which this species is very
Stegodon elephantoides { = cliftii): M l^^^M 2^' i^^. M abundant and characteristic (Pilgrim-Brown).

'Now in the British Museum (Natural History), M. 10520= Stegodon elephantoides (=diftii).
-[See note on page 824 above regarding the Lower Pleistocene age of Stegodon elephantoides. —Editor.
864 OSBORN: THE PROBOSCIDEA

This an curly Lower [Middle] Pliocene stage of the true

is .S. elephantoides, namely, length 280 mm., breadth 90 nun. (7)

Stegaddn based on types from the Dhok Pathan zone which also The collective (Falconei-, T^ydekker, Osborn) ridge-crest formula

contains Slegoloiihodoii. latidena. From this species some of the (not including the uncxi)os('d ridge-crests) is as follows:

earlier authors believed that the genus Elephas originated. Re- Stegodon howbifrnns: Dp 3 t Dp 4 j^r^^^lj^ M 1 ^^YVi

ferred Stegodon bomhifronn occurs in the Tatrot horizon. Middle M 2 M3 7»t-8-8t>-9-9^t

8-9-9VS [1 1+1 •

[Upper] Phocene. A superior molar from Pathan

the DIkjU
horizon (Fig. 746) exhibits \ery clearly the cone and conelet FALCONER'S NOTES OF 1868 ON ELEPH.\S [=STEG0D0N1
structure of the crown. BOMBIFRONS
E. [Elephuii] hombifrons Falconer and Cautley, 1846. "Fauna Fiilconer, 'Tiilu'diitoloKic^al Memoirs," Vol. I, ISOS, pp. l.'iC), l.'jS^dl,

Antiqua Sivalensis" letterpress, 1846, pp. 46, 47.

. . .
Llxto- Pliitos XXV -XXIX. H of tlic "Fuunii .Antiiiua Sivalensis."
[Errors of lietenniiuitioM iiulii'ated hy Oshorii in scpiaR^ l)nu-kots.l
TYPK AND CoTVPKS. {Op. cit., p. 46): ". scvoral crania.con-
.

taining perfect teeth," of which the lectotype is figured in PI. Elephit.s boiNb/froiia. I'alconer and Cautley, 1846 [1847, Plates
XXVI. Horizon and LocALiTY.^Siwalik Hills, India, prob- xxvi, XXVII, xxviii]. Lectotype and Cotypes. Cranium —
ably from the Dhok Pathan horizon. Lower [Middle] Pliocene. represented in Plate xxvi (Lydekker's lectotype); Plate xxvii,
Lectotype and Cotype Figures.— Op. cit., 1846 [1847, Pis. xxvi very fine skull, M', ridges 9%, length of molar, 10.2 in. = 257 mm.,
(lectotype), xxvii, xxviii], figures 742 and 744 of the present width 3.7 in. = 93 mm., and Plate xxviii, fig. 2, same .skull.

Memoir.
Falconer and Cautley, 1846, p. 46 (see above p. 858) based
this species on "several crania containing perfect teeth," described
as "from the Sewalik Hills." Of these Lydekker (1886.2, p. 83)
designated as the lectotype P.rit. Mus. M.2978. The exact locality
is not given. According to Pilgrim (Vol. I, fig. 413, also Chap.
XX n abundantly in the Dhok Pathan
below) this species occurs
horizon of Lower [Middle] Pliocene age, a horizon which also
contains Teiralophodon punjabiensis, Synconolophus corriigalun,
and S. hasnoli. Falconer assigned to this species a low ridge
formula, namely : M 3 ^^
Characters (Falconer and Cautley, 1846, p. 46); see
OsBORN ABOVE, PP. 855-859.— (1) Type based on several crania Fig. 743. Stegodon bombifrons. Much more ))rimilive
containing perfect teeth. (2) Crown divided into transverse ridges, than S. insignis. Observe transverse ridges of trunk to
composed ofnumerous mammillae [conelets] of chevron-shaped elevated nasals, extreme broadening of sumnut of oeeijiut.

by a thick coat of cement. (4) Ears conjeetural. Limbs given the same proportions as
section. (3) Intersjiaces occupied
the Stegodonts throughout, without knowledge of skeletal
Principal ridges of third upper and lower molars, 3¥, in contrast M material. Female to right, direetly after lectotype skull;
to [S.] insignis. (5) Third upper molar measures 11 inches (279 no inferior tusks. Male to left. Restoration by Margret
mm.) in length by 4K inches (114 mm.) in width. (6) The lower Flinseli, 1930. One-fiftieth natural size.

third molar of the left side, with nine [9)2] ridge-crests, measures
13.4 inches in length ( = 340 mm.) by 4.2 inches in breadth (= 105 Upper Jaws. — Plate xxix, fig. 1, broken cranium, M', ridges
mm.), coasiderably exceeding the dimensions of the lectotype of 8K, length of molar 10 in. = 253 mm., width 4 in. = 101 mm.; figs.

ELEPHAS BOMBIFRONS Cotype

PI. XXVIII, Fig. 2
ELEPHAS BOMBIFRONS Lectotype Lyd
ELEPHAS BOMBIFRONS Ret. PI. XXVI
Pi. XLIII, XIII
ELEPHAS BOMBIFRONS Ret.
PI. XLV, XIII (rov.)

Steoodon Crania, after Falconer and Cautley's Illustrations in the "Fauna Antiqua Sivalensis"
Fig. 742. Crania of Stegodon (Elcphas) bombifrons, lectotype, cotype, and referred. Outlines assembled from original plate drawings
if this .sjieeies in Falconer and Cautley's "Fauna Antitiua Sivalen.sis," frontal, palatal, and lateral .aspects. All one-sixteenth natural size.
 THE STEGODONTIN.E: STEGODON 865

2, 2a, cranium, ]\P, 8 ridges; figs. 4, 4tt, r.lVP, 8+ ridges; figs. 5, conelets is approximately the same as in Stcyodon elephanloides
5a, upper jaw, r.M', 7K ridges; figs. 6, 6a, palate with r.M', 9K { = cliftn).
ridges, length 10.9 in. = 276 mm., width 3.8 in. =97 mm. (at ends), From the above observations we deduce the ridge formula of
4.3 in. = 112 mm. (in middle). Stegodon bombifrons practically as above under "Characters."
Lower Jaws. Plate xxv, — figs. 3, 3a, lower jaw, r.Mj, 9%
ridges, enamel very thick, scanty cement. Plate xxix.a, figs. 1,
la, lower jaw with Dp4, ridges 6K, "probably the third [fourth] LYDEKKER'S NOTES OF 1886 ON ELEPHAS (=STEGODONl
milk molar"; figs. 2, 2a, lower jaw, r.Mi, ridges 7'4, "certainly
BOMBIFRONS
the first true molar"; figs. 3, 3a, lower jaw, I.M2, ridges Iji; figs. "Catalogue of the Fossil Mammalia in the British Museum (Natural History),"
Part IV, 1886, pp. 82-88
4, 4o, lower jaw, M2, ridges 7}i; figs. 7, 7a, lower jaw, Mi, ridges 7.
Plate XXIX. B, figs. 5, 5a, lower jaw, r.Dp, "with 5 ridges and an Lydekker (1886.2, p. 83) designates as his lectotype Brit.
anterior and posterior talon"; figs. 6, 6a, lower jaw, r.Mi, ridges Mus. M.2978: "The cranium, showing the third true molars of
7}i; figs. 7, 7a, lower jaw, I.M3, ridges 9>2 [11 + ]. both sides in an early stage of wear. This specimen is the type,
In the two plates (xxix.a, xxix.b) of the "Fauna Antiqua and is figured by Falconer and Cautley in the 'Fauna Antiqua
Sivalensis" about sixteen specimens of inferior molar teeth are Sivalensis,' pi. xxvi"; reproduced in figure 742 of the present
beautifully figured; they exhibit six to ten conelets on the unworn Memoir.
crown and a maximum of eleven conelets on the worn crown; Lydekker's notes are based on forty-four specimens in the
five of the conelets double by dichotomy. Thus the number of British Museum referred to this species, chiefly from the Cautley

CoTYPE OF Stegodon Bombifrons

I''ig.Cotype of Elcphas bombifrons I'alconer and Cautley, 1846. After Falconer and Cautley, 1846 [1847, Pis. xxvii, xxvin], one-sixth natural .size.
744.
Brit Mus. M.2979; cast Amer. Mus. 10378. From the Siwalik Hills, India.
.

Falconer, Pal. Mem., 1868, Vol. I, p. 458: (PI. xxvii) "Very fine and perfect skull, anterior view." (PI. xxviii) Fig. 1. "Lateral view of same skull,
as figured in Plate xxvii. —
B. M. Fig. 2. Palate view of same skull, showing sections of tusks, and last ? true molar on either side, with 9 ridges and a heel;
. .

—
the 8 front ridges worn. The interval between the molars in front is very narrow; behind they are extremely divergent. -B.M." Fig. 5. Occipital view of
another skull.
 :

866 OSBORN: THE PROBOSCIDEA

Collection (1842); the specific references therefore are based on ridges, enable us to write the standard maximum formula as
Falconer's determinations. follows
Specific Chakacters.— (Lydekker, 1886.2, p. 82): "The ^Maximum ridge formula of Ekphas = Stegodon] bombifrons:
[

ridges are rather taller, some\\hat wider apart, and more numerous Dp 2j Dp 3i Dp 4 'i M 1 f M 2,\^ M 3 i
than in E. clifti [
= Stegodo7i clephantoides { = clifiii)], and the
valleys are generally completely filled with cement; it is, however, Stegodon insignis Falconer and Cautley, 1845, 1846
sometimes very difficult to distinguish between the hinder teeth of Figures 686, 688, 697, 731, 732, 735, 747-753, 756, 760, 764, 766, 776, 819, PI. xx
the two species, while in the opposite direction it is often difficult Siwalik HiUs, India, Upper Pliocene, Pinjor horizon, to Lower Pleistocene,
to distinguish between those of E. bombifrons and E. insignis. Boulder Conglomerate {fide Falconer and Pilgrim), to Upper Pleistocene. In
The teeth figured by Falconer and Cautley under the name of the present Memoir (see Fig. 413) the upper levels of the Pinjor horizon are
E. ganesa cannot be distinguished from those of the present species, of Lower Pleistocene age {fide Barnum Brown).
and are therefore provisionally classed under the same head. Falconer's types of this species agree in the ridge-crest foi mula
The teeth are frequently very large, and the ridges are often with the type which he subsequently described as Stegodon ganesa

Fig. 746. Referred fragment of superior molar of

Stegodon bombifrons collected by Barnum Brown in
1922 in the Dhok Pathan horizon. Lower [Middle]
Kekekked Right Third SupEnion Molar of Steqodon bombifrons Pliocene, two and a half miles northeast of Ha-snot,
Fig. 745. A
partly worn right superior molar, r.M', referred by Falconer and Cautley India. This specimen is wrongly numbered; it
to Elephaa bombifrons and corresponding very closely in all details to Falconer's description should read Araer. Mus. 20044.
of the cotypcs of this species, namely M
3 with nine and a half ridges; conelets six divid-
: — Three anterior ridge-crests of a right third superior
ing into eleven. After Falconer and Cautley, 1846 (1847, PI. xxix, fig. 6], one-third natural molar, r.M' (compare Fig. 759 of Stegodon orien-
size. talis grangeri type).

curved; a trace of the median longitudinal cleft can often be but they exhibit profoundly different characters in the cranium, as
observed in the two or three ridges, and the inner columns of
first shown in a comparison of figures 735 (S. insignis) and 736 (<S.
these ridges occasionally show accessory tubercles near the longi- ganesa), or in a comparison of figures 752 (S. insignis) and 732
tudinal cleft, where they assume a Maslodon-hke shape. The (S. ganesa). This profound difference, as explained above and
plane of wear of the teeth of this and the following species [E. below, is attributed to the fact that all the crania referred by
ganesa] is similar to that of the true F^lephants. The mandibular F'alconer and Cautley to Stegodon insignis represent small-tusked
symphysis is produced into a spout-like termination, as in E. and probably female individuals, while crania referred to S. ganesa
indicus. The cranium has the fronto-parietal region very convex, represent large-tusked and probably male individuals.
the constriction of the frontals by the temporal fossff being more ]''alconer originally described Elephus insignis in 1846, [). 37,
marked than in the other species. Ilab. India (Punjab to Siwalik and in the same communication (]>.45) he named a fourth species
Hills) and (?)China [Footnote: 'Koken, Pal. Abhand. vol. in. Elephas Ganesa, describing a third superior molar (which had
pt. 2, p. 12 (1885).']. The species may perhaps also occur in been figured in 1845, PI. in, fig. 7a) and remarking that the tooth
Java." bears the closest resemblance to the corresponding tooth in E.
The ridge formula of Lydekker (op. 1886.2, p. 82), namely, insignis. This doubt always remained in Falconer's mind, for
MnA
m. Dpi I
1 MC.i-e)
V ,(,,.7), M
TV r 8.(fl.7).(8-9) .

7.(7.8).(8-9)' 's less

,
cit.,

precise
.

m
.
^ r
not refernng to
• ^
in his notes of 1S67, ]). 4, and of 1S68, Vol. I, yi. 424, he remarked:

tlic half-ridges but does not otherwise differ excepting in the supe- "In f.'u't, there are no good characters by which the teeth of these
rior ridges (^) of Dp fiom
type ridge fornuila gathered
'.i tiie two can be satisfactorily distinguished, although tiie crania
sj)ecies
from Falconer's obs('r\ations abov(^. Lydekker's obser\ations fully are so remarkably different." fn I'^alconer's mind, flicrcfore,
substantiate Falconer's fornmla of 1868, and, omitting the half- Elephus = Stegodon] insignis po.ssessed a cranium of the type he
[
 A —

THESTEGODONTIN^: STEGODON 867

figured in PI. xv of the "Fauna Antiqua Sivalensis," reproduced —

Horizon and Locality. Siwalik Hills, India, probably Pinjor
herewith in our figure 735, in contrast to Elephas [
= Stegodon] horizon. Upper Pliocene or Lower Pleistocene. Lectotype
gaiwsd which possessed a cranium of the type figured in PI. Figure.— Op. cit., 1846 [1845, PI. ii, fig. 6a]. Cotype.— 0/).
XXIII, reproduced in our figure 736. cit., 1846 [1845, PI. ii, fig. 66]. —
Description. Falconer's
—
Sexual Disparity. As explained above in the discussion of original description of 1846, p. 37, quoted in part above in this

the crania. Falconer apparently selected the female crania as refer- Memoir (p. 829) was restated in 1868, Vol. I, p. 423, fig. 6a of

able to Elephas [ = Stegodo7)] insignis and the wio/e crania as refer- PI. II, as follows: "Elephasfrom the Sewalik Hills [Fig.
iiisignis,

able to E. [S.] ganesa. A detailed comparison of all the referred 747 of the present Memoir]. Vertical section of last upper molar.
specimens has failed to establish any true specific distinction be- The four anterior ridges are affected by wear; the six posterior
tween these two species. Consequently we may regard the lecto- ridges are entire, the fangs are fully developed, and their mode of
type and referred specimens from the Pinjor, Boulder Conglomer- implantation in the jaw is distinctly shown. The white mass in
ate, and Godavari, Narbada Alluvium, as 'collective species' the centre represents the body of ivory, which is projected upwards
including a number of ascending mutations or subspecies which in ten angular lobes, terminating in a sharp edge. The height of
will be recognizable by profound monographic research; it is not these lobes does not much exceed the width of their base, and closely

7u/ eh

Ft J t'li

Lectotype axd Cotvpe of Stegodon insignis

Fig. 747. Lectotype left superior molar, I.M', of Elephas inmgnis Falconer and Cautley, 1846, from the Siwalik Hills, India, after Falconer and
Cautley, 1846 [1845, PI. ii, fig. 6a], also cotype third inferior molar, M3, after Falconer and Cautley, 1846 [1845, PI. 11, fig. 66j; both figures one-third
natural size.
(Falconer and Cautley, 1846, p. 37): "Fig. 6a, pi. 2 [lectotype l.M' —
Brit. Mus. M.30151, represents a vertical and longitudinal section of the
last upper molar of an Indian which we have named Elephas insignis in this work." Length of this tooth 10.3 inches. Inverted to show
fossil species,
natural position. (Falconer, "Palseontological Memoirs," 1868, I, p. 423): "The four anterior ridges are affected by wear; the six posterior ridges are
entire, the fangs are full}' dcvelojied, and their mode of implantation in the jaw is distinctly shown." Fig. 66 (cotype M3 Brit. Mus. M.3039) is a —
vertical section of a third lower molar of E. insignis, one third natural size (cf. Falconer, 1867.1, p. 4; 1868.1, I, p. 424; also caption to figure 097 above).

probnble that a single true specific stage passed from the Pinjor applied over them is a thick layer of enamel reflected up and down
into the Narbada Alluvium horizon. in a continuous zig-zag plate. The interspaces of the five posterior
In the present Memoir we shall first treat Falconer's descrip- ridges of enamel are completely filled up by a mass of cement much
tions of the types and referred specimens of these two species exceeding the enamel in thickness (vide Plate vi. fig. 7). This is

separately and then unite them under the collective species name the best illustration of the intermediate type of a proboscidean
Stegodon insignis-ganesa. molar tooth, from which those of the other species diverge in
Cranium. — The lectotype cranium of the third species of opposite directions. It belongs to the Mastodon Elephanio'ides of
Stegodont, described as Elephas insignis by Falconer and Cautley, CUft. The dark granulated shade below the portion of the ivory
1846, from the Cautley Collection but is without record as to
is nucleus sustaining the five posterior ridges indicates the hollow of
its The specimen is now in the British Mu-
exact geologic level. their common fang, which in the fossil is occupied by a core of
seum. Lydekker (1886.2, p. 91) designates it as ".M. 3015. An sandstone. B.M. (Reproduced in PL iv. fig. 1.) Length of tooth,
imperfect cranium, showing the third true molar of either side." 10.3 in."
Elephas insignis Falconer and Cautley, 1846. "Fauna Anti- Characters of Elephas [= Stegodon] i\sifiNis. (Lydokkor, —
qua Si\alensis," letterpress, 1846, p. 37. Leciotype. — 1886.2, p. 89): "The apparent impossibility of distinguisliing (Ik;
third superior molar of the left side, l.M', in an imperfect cranium dentition of this species from that of E. ganesa has been already
(Brit. Mus. M.3015) containing the third true molar of either side. mentioned. . . . The ridges of the cheek-teeth are usually rather
 J not. 31^ e

I 777?
-foresJiortenecl

7i.Mj

Fig. 748. Stegodon insignis ref. (Amcr. Mvis. 19869) suijcrior and inferior molars proVjably belonging
to the same individual as the mandible shown in figure 7.50. Pinjor horizon, Upper Siwaliks, near Siswan,
India. One-third natural size.
A, Al, A2, Imperfect second superior molar, 1.M-, with +5^ ridge-crests much
worn; a complete
worn.
third superior molar, LM', with }j-8-}'j ridge-crests, of which the two anterior are slightly
three anterior are
B, Right third inferior molar, r.Mj, displaying K-9+ ridge-crests, of which the
total of 12 ridge-crests.
slightly worn (eon\parc Fig. 7.")0). The l.M.i, now completely expo.sed, displays a

868
 A

THE STEGODONTIN^: STEGODON 869

taller and narrower than in E. bombifrons, their average number

greater, and the cement still more abundant. It is, however, not
always easy to distinguish between the two. The third molar is
usually narrower posteriorly, and the enamel frequently thinner.
The taller and more numerous ridges indicate that the present
species is intermediate in respect of dental characters between E.
/t. M /3369
bombifrons and E. plmiifrons. The adult cranium is remarkable
for the great depression of the fronto-parietal region, although this
feature marked in some specimens than in others [Footnote:
is less

'Compare "Fauna Antiqua Sivalensis," pi. xliii. figs. 15, 156.'];

but in the young cranium the contour is indistinguishable from
that of the adult E. ganesa [Footnote: 'It is of course self-evident
that these young crania (like detached teeth) might equally well
have belonged to E. ganesa.']."
Referkkd Materials in the American Museum. —
typical individual (Amer. Mus. 19869— Figs. 748, 750), In the
primitive S. insigni.^-gancsa stage, was foimd by Barmun Brown
near Siswan (Pinjor horizon), India, in which the upper and
lower jaws are fortunately associated; this specimen exhibits the
following ridge-crest formula:

M2 ±5W=7) M3
Also recorded from Siswan are two other si)ccimens, namely,
a second sujierior molar, r.M" (Amer. Mus. 19804 — Fig. 749), in

Fig. 749. Rpferrcd Stegodon insignis (Amer. Mus. 19804),

a seeond superior mohir, r.M-, taken from palate found below
the conglomerates, three miles northeast of Si.swan, Up))er
Siwaliks, India. Ridge-erests, namely, }2~7-y>, agree elosely
with S. insignis, somewhat too progressive for S. bombifrons.

which the ridge-crests ( ^"^''' ) agree closely with the tyjjical

formula of Stegodon iiisignis, and a third left inferior molar, l.Mj
—
(Amer. Mus. 19859 Fig. 753) with ^r^^i ridge-crests. A juvenile Fig. 7.50. Referred inferior mandible, assoeiatod with two upper teeth,
(Amer. Mus. 19869), collected by Barn\nn Brown in the
right lower jaw (Amer. Mus. 19858 —
Fig. 751) exhibits Dp4 with
of Stegodon insignis
Pinjor horizon, Upper Siwaliks, near Siswan, India; it contains both second
7 ridge-crests; a young adult lower jaw (Amer. Mus. 19964 Fig. — inferior molars, M2, well worn, also right and left third inferior molars, M3,
752B) shows I.M2 in full u.se, I.M3 coming into jjlace; the ridge- little worn, with )^9-|- ridge-crests; the I.M3, after removal of the bone,

crestformula of I.M2 is 7. Still another specimen from the same displays 12 ridge-crcsts (compare Fig. 748, probably of the same individual).

Siswan is a lower jaw, also a right upi)er tusk complete

locality of
(Amer. Mus. 19773— Fig. 766). Stegodon ganesa Falconer and Cautley, 1845, 1846
The grinding teeth of these three individuals in the American Figures 686, 698, 731-733, 736, 7.54, 755, 757, 760, 766, 819, PI. xx
Museum collection (see Figs. 748, 749, 7.50, and 753), all recorded
Siwalik Hills, India, Upper Pliocene Pinjor horizon, to Lower Pleistocene,
from the Pinjor horizon. Upper Pliocene [or Lower Pleistocene], Boulder Conglomerate (fide Falconer and Pilgrim), to Upper Pleistocene.
exhibit i)rogressive variation in the ridge count, as indicated by In the present Memoir (Fig. 413) the upper levels of the Pinjor horizon are of
numerals in these figures; according to Doctor Brown, it is not Lower Pleistocene age (fide. Barnum Brown).
certain that they belong in the Pinjor horizon proper, but may This species (fide Falconer, Lydekker) exhibits the same deni al
have come in by erosion from the conglomerates above. characters (Fig. 757) as Stegodon insignis but is distingiushed by
 dp^ STEqODON INSIGNIS i?e/r

crown view j/^ not. 5ixe .rev.

mer Mus. 19858

j/aiv, rev.

Fig. 7ol. Referred lower jaw, left aspect, of Slegodon insignis (Ainer.
Mus. 19964), mature adult, colle<'ted by Barnum Brown in 1922 in the upper
clays below the conglomerates, three miles north of Sisw-an, Pinjor horizon.
Upper Pliocene, India. The juvenile right lower jaw of 5. insignis (Amer.
Mus. 19858), collected by Dr. Brown also in the upper clays but two miles
north of Siswan, summit of the Pliocene, is inserted for comparison.
The ramus (Amer. Mus. 19964) exliibits a fully adult form, with abbrevi- /•^ncit. szje
ated symphysis, I.M2 in I.M3 coming into place. The ridge-crest
full use,
formula is M 2-J,
M 3 |. One-sixth natural size. younq adult JCl^
Observe Dp4 of Amer. Mus. 19858 which exhibits seven ridge-crests as
compared with the typical Dp4 of S. insignis with seven to nine ridge-crests
(of. Table V, p. 854 above). One-fourth natural size.

ELEPHAS INSIGNIS Bef. ELEPHAS INSIGNIS Hof.

PI, XVI, Fig, 2 PI, XLV, XV, B'fev.)

I'ig. 752. Referred crania of Elephas [=Stegodon] insignis. Specimens

reproduced in outline from original plates in Falconer and Cautley's "Fauna
ELEPHAS INSIGNIS Ret. .Viitiqua Sivalensis," as indicated in the printed legends. Frontal a.spect of
PI. XVI. Fig. 3
four crania, lateral aspect of one cranium, and posterior aspect of one cranium.
ELEPHAS INSIGNIS Ref.
PI. XV,
.Ml one-twentieth natural siz(^ Compare figures 732 and 777.

870
 THE STEGODONTIN^: STEGODON 871

profoundly different cranial contours, profile and facial aspects molar, M': "Plate iii, Fig. 7a. —Elephas Ganem, a fossil Indian
(compare Figs. 732, 752). Falconer, immediately after describing species. Vertical section of last upper molar. The crown consists
Eleplnis insignis (1846, p. 37), described Ehphas ganesa (1846, p. of ten principal ridges, with a subordinate talon ridge in front and
45) ; ajjparently hewas very much puzzled by the exact similarity in behind. The anterior seven ridges have their summits worn.
the structure of the teeth as contrasted with the profound difference A small portion is broken off at the anterior end. The disposition

Fig. 75.3. Rcferrod Slcgodon in-

signis (.\mcr. Mus. 198.59), collected by
Barnum Brown in 1922 two and a half
miles south of Cliarnian, near Siswan,
India, below the conglomerates, probably
Pinjor horizon. Upper Pliocene.
A third left inferior molar, I.M3;
length 291 mm., width 93 mm.; ridge-
crests of M 3 4-1 2 j«. One-third natural
size.

^rf*

in the character of the skull, for in the "Palseontological Memoirs" and relative proportions of the ivory, enamel, and cement bear the
of 1868 (Vol. II, p. 84) appears the following statement: closest resemblance to those of the corresponding tooth of E. insig-
"Regarding the specific distinctness of E. (Sicg.) Ganesa I am nis, and the number of ridges agrees. In fact, there are no good
by no means so well assured; this species is chiefly founded on characters by which the teeth of these two species can be satis-
a huge cranium in the British Museum with long tusks, presented factorily distinguished, although the crania are so remarkably
by Colonel Baker. I ha\e not been able to reconcile the form of different.— B. M Length of tooth, 9.25 in."
this cranium with either that of E. (Steg.) insignis or E. {Steg.) Lydekker (1886.2, p. 89) erroneously Mus. M.3008
.selects Brit.

bombifrons; but at the same time I must confess that I have as the type of E. ganesa: "The imperfect cranium, showing the
failed in tracing its dentition satisfactorily as a distinct form l^artially-woin third true molar of either side, the base of the left
through different ages." and the greater portion of the right incisor; from the Pliocene of
E. [Elephas] Ganesa Falconer and Cautley. "Fauna Antiqua the Siwalik Hills. This specimen (the missing portions of which
Sivalensis," letterpress, 1846, p. 45. Lectotype. —A last upper have been restored in wood) is the type, and is figured by Falconer
molar, M'. Brit. Mus. 18489. Horizon and Locality. — and Cautley in the 'Fauna Antiqua Sivalensis.' pis. xxi., xxii.,
Siwahk Hills, India, Pinjor horizon. Upper Phocene or Lower xxiii., and xliii. fig. 14. It is remarkable for the enormous size of the
Pleistocene. —
Lectotype Figi he. Op. cit., 1846 [1845. PI. in, incisors. Presented bij Gen. Sir W. E. Baker, K.C.B., 1848."
fig. —
7a see figures 698 and 757 of the present Memoir].
—
Lectotype Description. {Op. cit., 1846, p. 45) "The crown :

consists of ten principal ridges, with a subordinate 'talon' ridge in

front and behind. The anterior seven ridges have their summits
worn, the two in front being ground down to the common base of
ivory, the tooth having been a considerable time in use."
Characters. —Falconer (1868, Vol. I, p. 424) remarks of this

Fig. 754. This is the

famous and oft-repro-
duced referred skull
and tusks of Elephas
I
= Slegodon] ganesa
^ X.'
Falconer and Cautley,
1845- 1S47, reproduced
from Plate xxii, fig. 3,

of the "Fauna An-

tiqua Sivalensis."
Same skull figures 732,
733, and 736. Original
in the British Museum Sca/e o/'Iixt
(M.3008).

ELEPHAS GANESA
 Fig. 755. Restoration (1930 1933) of Slcgodon ganesa, to a onc-thirty-fifth scale, by Margret I'Miiisuh, under the direction of Henry Fairfield Osborn.

Based on Falconer's figures of the skull and tusks [.see also Fig. 733.)

Fig. 7.')G. Restoration (1934) of Slcgodon insigni«, by Margret Flinsch, under the dhection of Henry Fairfield Osborn. All figures about one thirty-
sixth natural size. Restoration based on crania figured in tlic "Fauna .\nti(|ua Sivalensis" by Falconer and Cautley. Aged individual at left (PI. xvi,
fig. 1), center (PI. XLiii, fig. xv..\, cf. PI. xv), right (PI. XLiii, fig. xv.B, of. PI. xvii, figs. 1 and 2).

872
 — —

THE STEGODONTIN^: STEGODON 873

FALCONER'S NOTES OF 1868 ON STEGODON INSIGNIS AND S. GANESA

FalcDner, "Paltoontological Memoirs," Vol. I, 1868, pp. 423, 424, 452, Plates ii -xxv of the "Fauna Antiqua Sivalensis"

Slegodon i.nsigiiis. Lectotype of Slegodon insignis from the Siwalik Hills. Plate ii, fig. 6a, M^ with 10 ridges, the height of
the.se ridgesdocs not much exceed the width at the base, closely covered with a thick layer of enamel, interspaces of five posterior
ridges of enamel completely filled with cement. (Falconer and Cautley, 1846.1, p. 37) "This tooth belongs to one of the forms
:

which have been included under the name of Mast. Elephantoides, by Mr. Clift; and which Profe.ssor Owen names 'Transitional
"
Ma.stodons.'
Stegodon ganesa. Lectotype. Plate in, fig. 7a, M^ length 9.25 in. = 235 mm. ". ten principal ridges, with a subordinate
; . .

talon ridge in front and behind. The disposition and relative proportions of the ivory, enamel, and cement bear the closest
. . .

resemblance to those of the corresponding tooth of E. insignis, and the number of ridges agrees. In fact, there are no good
characters by which the teeth of these two species can be satisfactorily distinguished, although the crania are so remarkably
different.— B.M."

Upper Jaws and Grinding Teeth

Stegodon insignis. —Plate xix, upper jaw with Dp- and Dp', Dp' with 6 ridges; figs. 4, in, skull, M' with 7%
figs. 1, In,
ridges; figs. 6, 6a, skull with M-', J\F with ll}j ridges, abundant cement, length of M' 11 in. = 280 mm., width 3.8 in. =97 mm.
Plate xix.A, figs. 2, 2a, palate with M^, M'; 2 ridges 7, M M
3 ridges K-ll-}-- Plate xxiv, figs. 6, 6a, iVP ridges 7)2, M- ridges
}r-8, ridges very high and compressed, ten points or denticles on 5th ridge. Plate xxiv.a, fig. 2, skull fragment, M' ridges
Qji, M- ridges }2-7-};', little cement.
—
Stegodon ganesa. Plate xxii, fig. 2, fine large skull, AP ridges 10}^, small talon in front. Plate xxiv, fig. 1, upper jaw,
M' ridges 6)2; fig. 2, upper jaw, (?) M- ridges 7; fig. 3, upper jaw, M^ ridges 7)j; fig. 4, upper jaw, M' ridges &}i, last ridge
—
with 7 denticles; fig. 5, imperfect upper molar 6 ridges with 9 denticles on fourth ridge.

Lower Jaws and Grinding Teeth

Stegodon insignis. —Plate 3, 3a, lower jaw, M3 with 12)2 ridges, 9 denticles on the 7th ridge; figs. 4, 4a,
xviii.a, figs.
lower jaw with Mi, M2, Mi with 4 ridges, M2 with 7 ridges, plates very deej); figs. 5, 5a, lower jaw, I.M3 with 11}2 ridges.
Plate XX, fig. 6, 6a, lower jaw, AIj with 9 ridges and front and back heel; figs. 7, 7a, lower jaw, M3 with 12 or 13 ridges;
figs. 9, 9a, lower jaw, M2 with ridges. Plate xx.a, fig. 6, lower jaw, M2 7 to 8 ridges. Plate xxiv.a, fig. 3, M2 [M3] ridges 12)),
9,''2

length 11.5 in. =291 mm., width 4 in. = 101 mm. Plate xxv, fig. 4, lower jaw, Mi ridges 7)2.
—
Stegodon ganesa. Plate in, fig. 7b, M3 [?M2l "appears to have consisted of eight principal ridges, with a talon ridge
behind, and a .subordinate ridge in front. ... It bears a close resemblance to the corresponding inferior tooth of E. insignis
in the form of the ridges, thickness of enamel, and proportion of cement. —
B.M." Plate xx.a, figs. 1, la, lower jaw, Mj with '

5)2 ridges, M2 with 8 ridges; figs. 2, 2a, Mj with 7)2 ridges. Plate xxv, fig. 1, M3 ridges +8. Plate xxv. a, fig. 1, lower jaw,
M3 ridges 7)2+. Plate xxix.b, figs. 2, 2a, lower jaw, Dp^ ridges 7)2.
Skulls of S. insignis and S. ganesa
Slegodon insignis. Plate xv. "The cranium is seen to differ remarkably from that of E. Ganesa (Plates xxi. and xxii.)
notwithstanding that the molars of the two species agree so closely. That of E. insignis is flattened at the top, elongated from
side to side and singularly modified, so as to bear an analogy to the cranium of Dinotherium giganteum, while that of E. Ganesa
does not differ much from the ordinary type of the Elephants." Plate xvi, figs. 1, 2, 3. "Fig. 1. Elephas insignis. Broken
cranium, oblique antero-Iateral view. Left orbit, &c., gone. This head is very cubical in form, is old, very concave in front and
vertically; teeth broken. Interval between inci.sive sheaths deep. No tusks. A black .specimen in Cautley's collection.
B.M." Fig. 4, skull with M', number of ridges +10)2, length of M^ 9.4 in. =239 mm. Plate xvii, figs. 1, 2, same skull as Plate
XVI, fig. 3. Extreme length from occipital bulge to plane of molars 23 in. = 583 mm., extreme width of occiput 25.5 in. = 647 mm.,
width of naso-maxillary opening 11.3 in. =288 mm., occipital condyles to anterior end of palate 22 in. =560 mm. See also
Plate XVII, figs. 3, 4, and Plate xviii, figs. 1, 2, 3, 4.

Summary of Ridge Formula

The above observations on ridge formulae include teeth in different stages of development and of attrition, hence the many discrep-
ancies. The adult ridge formulae are indistinguishable in the two species. Consequently we may combine the ridge numbers observed by
Falconer (in Murchison) in the following collective ridge formula of Stegodon ijisignis-ganesa.
Stegodon insignis-ganesa: Dp 3^ Dp 47^ M Ull M 2 '^f^ M 3
-'fUrrT^-
This collective ridge formula includes the maxima numbers only; whereas that given below by Lydekker (1886.2, p. 89, under E.
insignis) includes both maxima and minima. In a collective ridge formula we include ascending mutations, i.e., successive geologic stag(!s
in the evolution of the ridges, some more primitive, some more progressive, all constituting a collective species.

Lvdekker's (1886) Comparison of Stegodon bombifuons, S. insignis, and S. Ganesa

"Catalogue of the Fossil Mammalia in the British Museum (Natural History)," Part IV, 1886, pp. 88, 89
—
Characters of E. = Stegodon] ganesa. (Lydekker, 1886.2, p. 88): "The third true molars in the type cranium of this species
[

contain ten ridges, and thereby agree with the corresponding teeth of E. insignis rather than of E. bombifrons, a conclusion confirmed by
a second cranium, in which there appear to be either ten or eleven ridges in the same tooth [Footnote: 'See Rec. Geol. Surv. Ind. vol. ix.
p. 48 (1876).']. This close resemblance between the last molar of this form and of E. insignis renders it apparently impossible to draw any
distinction between the earlier teeth of the two forms [Footnote: 'The majority of the teeth figured in the "Fauna Antiqua Sivalensis"
under the name of E. ganesa have the low ridge-formula of E. bombifrons {q. v.).'], and all such teeth are therefore referred to the latter.
Falconer [Footnote: 'See "Palseontological Memoirs," vol. ii. p. 84.'] had considerable doubts as to the specific distinctness of the present
form, and as the resemblance between the type cranium and the young cranium of E. insignis [Footnote: 'See "Fauna Antiqua Sivalen-
 .

874 OSBORN: THE PROBOSCIDEA

sis," xliii. figs. 14, 15.'] indicates that (lie two are closely relatetl, it is jiossiblo that E. gaiiem may be the male form of E. insigiiis.
])!. Phe
athiltcranium iloes not differ \ery witlely from the tyi)e of E. indiciis, although the frontal constriction is less marked."
Lydekker's ridge formula of Elephas = Slegodon] innlgniti-ganesa, namely, Mm. [Dp] fr-7:l^-i}y
[
(7.8).(7-8).C9-l 1)
(7rfoMi^2Mn:i3)> oinit** M
the half-ridges; it also gives a higher ridge formula to 1 and 2. M M
Consequently the following maximum ridge formula according
to l.ydekker is higher than that of Falconer; this ridge formula, how^ever, is not sustained by Jjydekker's own observations on fifty-nine
specimens of the ('autley Collection in the British Museum, many of which are described and figured by Falconer.
Lydekker's combined ridge formula of Elephas = Stegodon] insignis-ganesa may be written as follows:
[

Dp 2 t Dp 3 r Dp 4 i M 1 t% M 2 U 3 H.
Stegodon insignis-ganesa presence of Slegodon airdwana in the same Kendcng-Schichten or
Falc. ner and Cautley (1846)-Osborn (1928) Trinil horizon [Middle Pleistocene] of Java with Palxoloxodon
hi/siidrindieus, which, according to Stremme (1911) and Janensch
ria\ing now reviewed in detail the observations of Falconer
(1911) is comparable to Elephas anliquus (i.e., P. naniadkus).
and of Lydekker, we appear to be forced to the conclusion that
female and male See note on the fauna of the Kendeng-Schichten below.
Stegodon insignis and tS. gaiiesa are respectively
In summation, the phylum Stegodon insignis-ganesa of India
representatives of a single 'collective species' which first appears in
Pleistocene, Pinjor horizon, extending into the Boulder
and the phylum Stegodon airdwana {=javanicasy of Java repre-
the Lower
sent the last surviving Lower and Middle Pleistocene [to
Conglomerate, while specimens at present referred to the same
species occur in the Middle [LTpper] Pleistocene Godavari, Narbada
l^pper Pleistocene] members of the southern forest-browsing

Allu\ium,contem]ioraneous with Elephas (Hypselephas) hysudrictis Stegodonts. These two Stegodont phyla were geologically con-
and I'aUeoloxodoii iianiadicus of the Codavari. temporaneous with the forest- and plains-browsing Loxodonts,
Doubtless we are dealing with a series of ascending mutations and with the grazing ancestral true elephants, such as Archi-
which in time may be distinguished by valid constant specific or diskodon

Sectioned Molahs of Stegodon ganesa and Stegolophodon latidens

Fig. 757. (Left) Vertical section of loctotyiie left third sui>ei'ior molar, 1 .
M'', of Elephas [
= Stegodon] ganesa Falconer and Cautley,
1846 [1845, PI. HI, fig. 7a], less than one-half natural size. Brit. Mus. M. 18489 (erroneou.sly referred by Lydekker, 188G.2, p. 84, to E.
bombifrons). From Siwalik Hills, India. (Falconer and Cautley, 1846.1, p. 45): "The crown consists of ten principal ri<lgcs, with a
subordinate 'talon' ridge in front and behind. The anterior seven ridges have their summits worn, the two in front being ground
down to the common base of ivory, the tooth having been a considerable time in use." Molar inverted to show natural position.
(Right) Vertical section of two last upper molars of the right side, r.M^"', of Mastodon {= Stegolophodon] laiidens Clift, after
Falconer and Cautley, 1846 [1845, PI. ill, fig. 8], than one-half natural size.
le.ss Same molars as those figured by Clift in 1828, PI.
XXXVII, fig. 1. From Iriawaddy River, Burma. Compare figure 719 above.

subspecific characters; meanwhile we regard these ascending Stegodon insignis birmanicus Osborn, 1929
mutations as a collective sjiecies Stegodon insignis-ganesa. The Figures 758, 760
collective ridge formula is probably as follows, indicating as Mingoon, opposite Mandalaj', Burma; upper levels of the Irrawaddy
iiiiniiiKi the ridge-crests characteristic of the more primitive stages Series, Upper Pliocene."

and as maxima the ridge-crests characteri.stic of the more progres-

This left ramus (Amer. Mus. 20002), collected by Barnum
sive stages so far as known:
Brown ill 1922, agrcM's with Stegodon insignis in the number of the
S. insignis-ganesa : Dp 2 i Dp 3 Dp 4 ^^„ M 1
7
7H- S
W- 1 ridge-crcsis of the third molar hut greatly exceeds this typical
M2^^tir-M3 «"**
^" "M-9.V', w-1 i-H-iz-iBM-is- Siwalik species in size; the jaw and inferior grinder appear to be
The survival in India of referred Slegodon insignis-ganesa in one of the largest of the Stegodonts heretofore described (Figs. 758
the same Middle [Upper] Pleistocene le\cls with the true elejihants 760) the tooth indicates a higher degree of siiecialization than the
;

and Loxodonts, such as Elephas (Hypselephas) hysudrieiis and tyjiical S. insignis-ganesa and a specialization different from that of
Palseoloxodon namadieus, is j)aralleled in the East Indies by the >S. orientaiis grangeri; f he S. insign is birnian ints molar is of gigant ic
'[Synonym of Stegodon airawaiia or S. trigonncephalus (see footnote on page 889 below). Editor.] —
'[See note on page 824 above regarding the Lower Pleistocene age of the upper levels of the Irrawaddy Scries. — Editor.)
 THE STEGODONTIN^: STEGODON
size, there are many ridge-crests, far apait, with a mciUuni iiumtjer second pair, are partly worn, ridge-crests slightly open; this is
of conelets, and considerable cement; the ,S. nrieiitalifi (jranqeri followed Ijy ridge-crests pentalophid to octalophid (5th to 8th),
molar is of medium to large size, there are many ridgo-crests, jiartly worn; conelets still separate; the ninth to twelfth ridge-
ap])roximated, manj' conelets, and modei'ate de\elopment of crests are entirely unworn. The conelets are few and stout; they
cement. \ary in number from four to twehe on each ridge-crest. Cement
Stegodon insignis birmanicus Osborn, 1929. "New Eurasiatic is present all the way back.

Fig. 758. Type of Slegodon insignis birmanicus Osborn, 1929 (Amcr. Miis. 20002), a twelve aiul a lialf erested third left

inferior molar, l.Ms, in the same ridge-crest stage as Slegodon insigrtis-ganesa. One-tliird natural size, t^rom Mingoon, opposite
Mandalay, Burma.

and American Proboscideans." Amer. Mus. Novitates, No. 393, Stegodon orientalis grangeri ()sl)oni, 1929
Dec. 24, 1929, pp. 15, 16. Horizon and Locality. Upper — Figures 682, 684, 686, 687, 731, 759-764, 777, 1231, PI. xx
Pliocene [now regarded as of Lower Pleistocene age]. Mingoon, Upper Pliocene [Lower Pleistocene] of Yenchingkou, neai- Wanhsien,
opposite Mandalay, Burma. Type Figure. — Op. cil., 1929.797, Province of Szechuan, China.
p. 16, fig. 16.
The subspecies Stegodon orientalis gmngeri, as shown in fig-
—
Type. Amer. Mus. 20002, a very large and massive left
ure 687 is more primitive than the type of S. orientalis, also
inferior jaw containing the left third inferior molar, I.M3. The from a cave in Szechuan; the ridge-crests are less elevated and
ridge-crests of I.M3, namely, 7^775. are the same in number as in
wider apart at the base and seem to be even more primitive than
Slegodon insigtds, but the elongation of this inferior molar and the
those of the iS'. insignis type (Figs. 688, 747) the cranium is much
;

open character of the ridge-crests are quite distinctive from S. smaller and simpler than that of S. ins/gnis-ganesn (Figs. 735, 736)
insignis; the jaw is more massive and the inferior grinding teeth
and resembles in its contour rather that of .S'. bomhifrons (Figs.
surpass in length measurement those of any other Stegodont type
732, 734, 777).
known; the grinders are larger and exhibit fewer conelets. The
Stegodon orientalis grungeri (_)sborn, 1929. "New Eurasiatic
principal measurements (in millimeters) are as follows:
and American Proboscideans." Amer. Mus'. Novitates, No. 393,
Total length of third inferior molar, I.M3 Dec. 24, 1929, pp. 16 and 17. Type.— Amer. Mus. 18714,
a left third superior molar, l.M', and right and left third inferior
At the crown 333
At the base 362
molars, r.Ms, I.M3, of the same individual. Horizon and
Total maximum breadth of I.M3 97
Locality'. —Upper Pliocene [now regarded as of Lower Pleistocene
Breadth-length index of I.M3 29^27 age]. Yenchingkou, near Wanhsien, Province of Szechuan,
Length of mandibular ramus from base of M3 China. —
Type Figure. Op. cit., Osborn, 1929.797, p. 16, fig. 16.
—
1

to tip of symphysis 560 Type and Referred Figures. Aside from the type (Figs.
Horizontal thickness of mandibidar ramus 204 759, 760, 762) there is very abundant referred material from the

Height of ramus at symphysis 208 same locality, including the complete superior and inferior den-
tition, Dp 2-Dp 4, M
1-M 3, in various stages of attrition and
In detail, the entire jaw and posterior grinder are very mas- dental succession, as illustrated in figures 687, 759, 763, 761, 762 of
sive.Of the total of 12-)2 ridge-crests, the anterior pair are deeply the present Memoir, constituting the most complete material of
worn and widely open; the tritolophid and tetartolophid, or dental succession known in the fossil Proboscidea.
 STEQODON ORIENTALIS QRANQERI
PERMANENT MOLARS

y^ /VZ /S636

STEqODON 0RIENTALI5 QRANQERI

MILK DENTITION -^ naT. si'je

A.M. /8705
A./^. /a7/4- Type

i. dp2 I. c^pj

Fig. 7o9. Type (Amcr. Mus. 18714) and referred specimens (Amer. Mus. 18705, 18711, 18630a, 18636, 18642) of Slegodoii nrifnlnlis grangcri, part of the

collection from Yenchingkou, Province of Szechuan, China, made by Walter Granger during the winter of 1920-1921. All figures one-fourtli luitunil size. (Cf.

Figs. 761 and 702 for crown view.s, giving ridge-crests and conclets.) The specimens hearing numher 18642 have hccn sent to Peking, China, in cxcliiiMgc.

Left lateral aspect of: Crown view of:

R.Dp2, r.Dp' (rev.)— I.Dp2, l.Dp,, (Amer. Mus. 18705) Type third left inferior molar, I.M.i, strongly concavo-convex, out-

L.Dp', l.Dp" (Amer. Mus. 18711), and l.Dp4 (Amer. Mus. 18630n) wardly arched. Amer. Mus. 18714.

L.M', r.Mi (rev.) (Amer. Mus. 18636) Type third left superior niolai', l.M'', strongly convex, parallel sides.

L.M^, also r.Ma (rev.) (Amer. Mus. 18642) Amor. Mils. 18714.

L.M^ I.Ms (Amer. Mus. 18714)— type.

876
 THE STEGODONTIN^: STEGODON 877

Materials of Stegodon orientalis grangeri, Type and orientalisOwen (Fig. 769) which is recorded from a cave in the
—
Referred. Collected by Walter (iranger in 1920-1921, all from Province of Szechuan, northwest China, and much more progres-
the same pit; estimated specimens: sive thanOwen's type of Stegodon sinensis which appears to be
comparable to S. bombifrons. The ridge-crest formula, beautifully
Separate crania, more or complete less 6
shown in figure 759, is as follows:
Separate crania, more or fragmentary
less 7
Right and left mandibular rami and complete in- Dp2^^Dp3^, Dp4 5i-6-V» Ml M-6-^s
M2 V4-8-W
(7)-9» J^l 3
11-W
jj.f 3
ferior jaws 28
Separate superior and inferior grinding teeth in
Comparison with other Chinese Stegodonts. — On closely
comparing Owen's types of Stegodon sinensis (Figs. 687, 702) and
various stages of succession 49
S. orientalis (Figs. 687, 769) with the Stegodon orientalis grangeri
Skeletal bones, very few, not exceeding 3-4
teeth (Figs. 687, 759, 761) collected by Doctor Granger at Yen-
—
Characters. An ascending mutation or a subspecific stage chingkou, it is certain that we have to do with three distinct spe-

regarded as somewhat more primitive than the type of Stegodon cific and subspecific stages broadly distingui.shed as follows:

Upper Miocene [Pliocene] stage. Lower Pleistocene stage. (?) Lower Pleistocene stage.
Owen's type of Osborn's type of Owen's type of
Stegodon sinensis Stegodon orientalis grangeri Stegodon orientalis
Brachyodont. Subhypsodont (first stage). Subhypsodont (second stage).
Lophs compressed, far apart. Lophs more compressed, more elevated. Lophs still more compressed, still more
Compare Stegodon bombifrons. Compare the typical Stegodon orientalis. elevated.
Compare the referred Stegodon orien-
talis and <S. airdwana.

Tier y%e^
Amer. Mils /a7IA- 7y/>e I. mi
STEqODON ORIENTALIS QRAN5ERI /3 naZ. stje

Amer Mus 20002' Type'

STEqODON INSIQNIS BIRMANICUS

Arner. f^us. /SS69 J^e/r^^^^ ^ .^^; ;

J-" ITtner VLen/
'-""'^
STECODON INSIQNIS - GANESA ^~--<=s^^i,«^__»£...— ^. „<.^

COMPARLSON OF StEGODON INSIGNIS-GANESA, S. INSIQNIS BIRMANICUS, AND S. ORIENTALIS GRANGERI

Fig. 7t)0. Left third inferior molars, to a one-third scale, from the internal aspect, showing the eruption line of the grinder and exposing the six
anterior ridge-crests in Stegodon orientalis grangeri (ujiper) and S. insignis ganesa (lower). After Osborn, 1929.797, p. 16, fig. 16.
Stegodon orientalis grangeri, type (Amer. Mus. 18714). Large size with 13-|- ridge-crests of which 6+ anterior were erupted while 7-13 were
buried in the jaw.
Stegodon insignis birmanicus, type (Amer. Mus. 20002), with 5-6 ridge-crests erui)ted, 6-12 buried in the jaw and encased in cement (dotted).
Stegodon insignis-ganesa ref. (Amer. Mus. 19869), with -|-l-6 ridge-crests erupted, 7-11 buried in the jaw, mostly lacking cement.
This comparative figure illustrates the gigantic size attained; the largest {S. insignis birmanicus) slightly exceeds in length the correspond-
ing grinder of »S. orientalis grangeri.
 <5
mm
2— M Slrgmhn nriinlnlis gmiigeri (Amcr. Mus.
Fig. 761. Superior and inferior second, third, and fourtli premolars and first molar; (Dp 1 ) of

18636, 18711, 18630a, 1870.5), side and crown views, one-half natural size (ef. Fig. 7.59).
(Upper pair). First superior and inferior molars (Amer. Mus. 18636), l.M> with +6+ ridgc-erests, conelets 10 niaxinmni, r.M, (rev.)

with 8 ridge-crests, conelets 9.

conelets 12 13, 1.1)))4
(Middle pair). Fourth superior and inferior deciduous premolars, l.Dp^ {Xim-T. Mus. 18711) with }.,&-}: ridge-crests,
(.\mer. Mus. 18630a) with 7-)3 ridge-crests, conelets 12 maximum.
(Lower pair). .Second and third superior and inferior deciduous i)remolars (Amer. Mus. 1S70,')), r.Dp- (rev.) with +3+
^
_
ridge-crests, r. Dp
(rev.) with 6+ ridge-crests, conelets 15 maximum, l.Dp2 with 2 ridge-crests, l.Dps with 6 1- ridge-crests, conelets 16 maximum.

878
 Fig. 762. Type and referred .superior and inferior molars of Stegodon onentalis grangeri. Crown views (cf. Fig. 7.59). After

retouched pliotographs, one-half natural size.

(Upper pair) Type (Amer. Mus. 18714), left third superior molar, l.M^ with -I-IIJ2 ridge-crests. Left third inferior molar, I.M3,
with Yr-M ridge-crests; inner side strongly convex, outer side strongly concave, in contrast to the parallel sides of the upper molar.
(Lower pair) Amer. Mus. 18642. Left second superior molar, l.M^, with -f-8-f- ridge-crests. Right second inferior molar, r.M2 (rev.)
with 9-1- ridge-crests. These molars have been sent to Peking, China, in exchange.

879
 —

880 OSBORN: THE PROBOSCIDEA

CRANIAL CHARACTERS OF STEGODON ORIENTALIS GRANGERI to lower grinders, M1.2, with ridge-crests 8 and 9 respectively (cf.
(Figs. 7.')9, 701, 703, 777)
Figs. 759,762 lower); these ridge-crests, partly bathed in cement,
The iiifiiiililc craniuin of Slcgndoii oricidalis graiigcri (Amer. of moderate hypsodonty, exhibit the characteristic convex coronal
Mvis. 18638, 18702), as represented in the composition drawing curvatiue of the upper ridge-crests, the concave coronal curvature
(Fig. 763), exhibits the small deciduous superior incisors and the of the lower ridge-crests, a feature much more strongly marked in
deciduous premolars, Dp 2-4 (for crown views of these deciduous the third molars (Fig. 759). In the inalure adult cranium (Amer.

STEqOOON ORIENTALIS qPANCqERI

/)// -g nat.sije

Fig. 703. Infantile, juvenile, yoinig adult, mature adult crania showing eruption of Dp'— M', Dps — M3. Yenehingkou, Provinee of Szechuan, China.
Infantile —
cranium (Amer. Mus. 18038) showing succession of Dp" Dp'', also Di', together with j.aw (Amer. Mus. 18640) belonging to tliis sUull, showing
Dp2— Dpi.
Juvenile cranium (.\mer. Mus. 18702), jaw (Amer. Mus. 18711), showing eruption of Dp3, Dp4.
Young adult cranium (Amer. Mus. 18630) showing M' and M^. Jaw (Amer. Mus. 18630) with Mj in xilu, .sent to British Museum.
Mature adult cranium (Amer. Mus. 18708), jaw (Amer. Mus. 18029 — .sent abroad in exchange), showing M^, M', and M3 in silu.

premolars, compare Fig. 761); the rounded jirofile, of subtriangu- Mus. 18708, .skull, 18629 jaw) the second superior molar, AP, is

lar form, should be compared with the infantile cranium of Mas- disappearing, while the third superior and inferior molars, M',
todon acutidens (Fig. 131) and that oi Elephas indicus (Fig. 799). M3, are fully functional, their reciprocal convexo-concave relations
The juvenile cranium (.\mer. Mus. 18702, sktdl, 18711, jaw being beautifully shown in figure 763 (right). The upper jiortion
a composition drawing) has lost the deciduous incisors, also Dj) 2, of this mature adult cranium is fractured, leaving tlie exact
but retains the functional Dp 3-Dii 4 (cf. Figs. 759 and 761). profile contoui- in doubt.
The young adult cranium (Amer. Mus. 1S630, skidl, 18636, The third superior and inferior molars of this subspecies
jaw) exhibits the rounded and greatly elevated dome; of the l)rain grandly r('|)resented in the li/pc si)ocimen (Amer. Mus. 18714) seen
case; all the deciduous teeth, Di-, Dp 2-4, have disappeared and I lie in both lateral and crown \iews in figures 759 and 762, add a great
grinding function is now assumed by
and second truethe first deal to our knowledge of the Stegodont dentition and show
molars, M 1 -M 2 (cf. would a|)pear that
Figs. 761, 763). It \ery marked mechanical (-ontrasts between the supeiior and in-

the grinders M''-, with ridge-crests 6 and 8 respectively, are opposed ferioi' grinders as follows:
 THE STEGODONTINJ!;: STEGODON 881

Third SurEiuon Molaus A Acry careful study of these four ontogenetic stages of the
], literal profile strongly convex. cranium, namely, infantile, juvenile, young adult, and mature
Posterior ridge-plates elongated, partly <jr wholly bathed adult, reveals resemblances and contrasts with the Mastodon and
in cement. Elephas crania which will probably prove of phyletic or generic
Sides of crown parallel. value. Similarly, the remarkable ridge-crest succession, the
mechanical warping of the superior and inferior grinders, the
Third Ixferior Molars gradual increase in height (hypsodonty) of the superior ridge-crests
l.ateral profile horizontal. from Dp 2-M 3, indicate that the highest and most progressive
Anterior and posterior ridge-plates of uniform height; ridge-crests are reserved for the extremely adult stages in which the
cement chiefly in median plates. posterior ridge-crests of the eleven and a half crested third superior
Sides of crown strongly arched, convex inner side, concaK molar grind against the less hypsodont ridge-crests of the thirteen

outer side. crested third inferior molar.

STEGODON mSIQNlS

Br/t Mus. M.30/5 Type

STEGODON PINJORENSIS

STEGOOON AIRAWANA

Fig. 704. Cuni|)a.r!itivc sections of tliird left superior molars of .Stcgodonts of

India, Java, and China, all to a one-third scale.
A, Stegodnti pinjorcnsis Osborn, type. A 14-|- ridge-crested I.M^, from near
Siswan, India. Amer. Mus. 19772.
B, Sicgoclon insifjnis Falconer and Cautley, lectotype. All ridge-erest,etl
l.M^ from the Siwalik Hills, India. Alter Falconer and Cautley, 1840 [184."),
PI. 11, fig. Oal. Brit. Mus. M.3015.
C, Stcgodon airawana Martin, rcf. Portion of an l.M' of the Middle Pleisto-
cene of Trinil, Java, exiiibiting 7 of the 12-14 ridge-crests; much more recent in
structure than the molars of Stcgodon. insignis of the Lower Pleisto<^ene. This
tooth was presented to the Amei'ican Museum by the Geological-Palaeontologi-
cal Tnstitutc and Museum of the University of Berlin, Gi^nnany, through the courtesy of Geheimrat Pompeckj and Doctor Dietrich. Amer. Mus. 22G30.
D, Skgodon oriculalis grangcri Osborn, typ(^ An l.M^ with +11 }> ridge-(Te.sts, from near Wanhsien, Province of S?;echuaii, China. Amer. Mus. 1S7I 1.
STEGODON QANESA Ref. r Rev. >

A-fter Thlc.oner

Fig. 766. Referred lower jaw and right superior tusk (A) of Slcgodon ganesa (Amur. Mu.s. 19773), collected by Barnuni Brown in 1923 in the Upper
Siwaliks, below the conglomerates, three miles north of Siswan, India, compared with skull and tusks (B), after Falconer, 1846 [1817, PI. .xxii, fig. 3].
Original in British Museum (Brit. Mus. M.3008). Both figures one-sixteenth natural size.

882
 THE STEGODONTIN^: STEGODON 883

Stegodon pinjorensis Osborn, 1929 occipital crest, lofty and greatly abbreviated frontonasal surface,
Figures 711, 731, 704, 765, 707, 70S, 777, 817, 1217, PI. xx anterior nares correspondingly elevated, grinding surface of the
"Three miles nortli of Siswan, India." Upper levels of the Pinjor horizon, large molars very strongly arched, but the cranium is relatively
Lower Pleistocene. more depressed or bathycephalic than in S. ganesa."

Slegodon pinjure nsis Osborn, 1929. "New Eurasiatic and ^5 A/atural 5tj&
American Proboscideans." Amer. Mus. Novitates, No. 393, p. 18.
—
Type. "Amer. Mus. 19772. A male cranium, rostrum wanting;
portions of right inferior tusk preserved." Horizon and
Locality. — "Recorded by Barnum Brown as follows: 'Skull.
.Just below Conglomerate beds, Upper Siwaliks, three miles north

of Siswan, India.' This progi-essive cranium does not belong in the

Pinjor (I'pper Pliocene) horizon, as the specific name pinjorensis
was probably deposited from the overlying Boulder
suggests, but
Conglomerate beds of Lower Pleistocene age." Type
Figure. — Osborn, 1929.797, p. 17, fig. 17.
Type Description. — "Superior grinding teeth distinguished
from those of Slegodon insignis-ganesa bj' their superior size,
nuich more numerous ridge-crests, progressive hypsodonty; the
comparative ridge formula of M 3 are as follows: A. S. PINJORENSIS Type A. At /9772
S. PANES* Tfef /l^ter rbZconer
Slegodon pinjorensis: M3 ^^^"^^

Slegodon insignis birmanicus: M 3 YzTi Fig. 707. Tyiic skull (Amer. Mus. 19772) of Slegodon pinjorensis (A),
Slegodon insignis-ganesa: M3 lilsfj't
also skull of
skull of
same individual (A) superimposed (dotted
ganesa, after Falconer (B).
lines) on referred
<S. Onc-sixtecntli natural size.
Observe the extreme bathyroplialy in the skull of Slegodon ganesa in
This type male cranium resembles that of the male type of comparison with the more typical vStegodont form, as exemplified by the
Slegodon ganesa Falc, namely, with small rounded parieto- skull of S. pinjorensis.

Fig. 768. Front view of Slegodon pinjorensis before present arrangement

of the tusks, which now are regarded as possibly turning inward, as in new
restoration of Slegodon ganesa (see Fig. 733, p. 857 above); otherwise head
true to present proportions. Forelimbs entirely conjectural, drawn in pro-
portion to size of head. Ungues 5 and 5. Restoration by Margret Flinscli,
May, 1930. One-fiftieth natural size.
 "

884 OSBORN: THE PROBOSCIDEA

Stegodon orientalis Owen, 1870 city of Chung-king-foo, in the province of Sze-chuen.' Type
Figure 087, 703. 7ti9, 770 Figure. — Op. cit., PI. xxviii, figs. 1-4.

Szechuaii, iiortliwcst China, near the city of Clmngkiiigfoo. Probiiljly Type Desckiption. — {Op. cit., p. 421): "The dentine retains
Lower Pleistocene. Swinhoe Collection of 1870. its original white colour, . . . the enamel also has its recent pearly
The imperfectly known Stegodon orientalis, as shown in com- tint; a thick mass of cement appears to have been retained in the
parison of figures 687, 688, 697, and 769, is apparently more intervals of the coronal ridges. One of these ridges, with the con-
progressive, with more elevated
than the type of >S'.
ridge-crests, tiguous halves of two others, form a molar two inches nine lines in
insignis (Fig. 697); the ridge-crests (Fig. 769) are more acute and breadth (PI. xxviii. figs. 1 & 2); a portion of a posterior ridge
more compressed at the base, and the summits are more closely with a low basal heel, from the same, or the same-sized tooth, and
approximated; with numerous conelets and strong layers of the last two ridges, with a terminal half ridge or talon, of a milk-
cement in the valleys. It is, however, impossible clearly to define molar, one inch and a half in breadth (ib. figs. 3 & 4), represent the
and separate this species until further material is found in the present species. . . . The condition of the fragments agrees with
type locality. It is somewhat more progressive than S. orientalis the statement, viz. that they were from a cavern. . . . The ridge
gr anger i. (ib. fig. 1) a a runs straight, or nearly so, across the tooth; the
This species described by Owen from an imperfect type figure entire ridge is cleft at the summit into about a dozen mamillce
(Fig. 769) is regarded by Lydekker (1886.2, p. 97) as a synonym of by as many vertical grooves ; the dentine rises into the base of each
3

I'"ig. 769. Type oi Stegodon orientalis Owen, 1870, from a cavern in Szechuan, northwest China, .^fter Owen, 1870, PI. xxviii, figs. 14.
Original in the British Museum (41926-7). "Portion of true molar, grinding-surfacc." Fig. 2. Same, "side vi(-w."
(Op. cit., p. 433, fig. 1):
Fig. 3. "Hind end of milk-molar, d 3, grinding-surface." Fig. 4. Same, "side view."
—
Locality. (Op. cit., p. 421): "These fragments form part of the series of teeth obtained by Mr. Swinhoe, and said to be 'from a cave, near
the city of Chimg-king-foo, in the (jrovincc of Szc-chucn.' The condition of the fragments agrees witli tlie statement, viz. tliat they were from
a cavern. . . . [)). 434] Mr. H. Woodward stated tliat Mr. Swinhoe liad himself obtained a .series of these fossils from a cave many miles in-
land — he believed, on the course of the Yang-tse-kiang."
Stegodon insignis (.sec citation below). Koken (1885) also regarded mamilla. The enamel (e) averages two lines in thickness."
5. orientalis as a synonym of S. insignis, Schlosser (1903) agreeing "From the above-defined characters it is plain that we have
with Koken. In Osborn's opinion, the imperfect character of the here, also, parts of a 'transitional Mastodon,' in other words,
type molar (Fig. 769) and associated milk molar renders it tlifficult a species of Stegodon, Fr. In the straight, or nearly straight,
to determine whether these sjiecimens are in the Stegodon insignis direction of the coronal ridges, and the absence of any trace of
stage; in both specimens the lophs are elevated. From another mid cleft, these molar fragments more resemble Ihe (oeth of
cave locality, 140 miles distant, near Yenchingkou, comes the Stegodon Cliftii, St. insignis, and St. ganesa of Falconer than does
superb material collected by (iranger, which proves to be somewiiat the St. sinensis; and in the apparent quantity of coronal cement
more priniiti\e than Owen's type of Stegodon orientalis ant! is (ib. fig. 2r) as well as in the evidence of a hinder talon (ib. fig. 3 1),

described as Stegodon oricidalis grangeri (see p. 875 above). they are more like St. insignis than St. Cliflii. Yet the two hinder
Owen's lyi)e description is in jjart as follows: ridges, with the terminal talon of the tooth (ib. figs. 3 & 4),
Stegodon orientalis Owen, 1870. "On Fossil i{einains of Mam- which, in breadth, corresjionds with the second vi])pcr deciduous
mals found in China." (Juart. Journ. Cleol. Soc. ]>ondon, \'ol. molar of .S7. insignis and .S7. sinensis, clearly differ from both.
XXVI, Pt. 1, p. 421. Type.— Molar fragments (Brit. Mus. The two ridges run straighter across, are of the same extent,
last
41926-7). IIoiuzo.N' and L()c,\ijtv. — (Op. cil., ]i. 421): ". . . and are divider! by more ntmierous xerticnl grooves into smaller
obtained by Mr. Swinhoe, and said to be 'from a ca\c, near the and correspondingly luuncrous apical maiiiilhe. The second of
 THE STEGODONTIN^: STEGODON 885

these ridges is cleft in tho middle. From the alleged conditions of Ste^odon airawana Martin, 1890
discovery, and the little-altered condition of the above-described Figures 686, 688, 707, 731, 764, 771, 773, 774, 777, 779, PL xx
portions of proboscidian molars, one would be led to deem them Kendeng-Schichten, Pilhecanthropxis erectus zone, Middle(?) Pleistocene,
to be of as comparatively recent geological age as our ordinary Alas-Tuwa, Trinil, Java.
British Cave-fossils."
This is the most progressive Stegodont known, surpassing

ON STEGODON ORIENTALIS Stegodon orientalis and greatly surpassing S. insignis-gancsa in the

LYDEKKER'S NOTES OF 1886
"Catalogue of the Fossil Mammalia in the British Museum (Natural History)," elevation (hypsodonty) and number of the ridge-crests. As shown
Ft. IV, 1886, p. 97 in figures 687 and 688, the fourteen to fifteen ridge-crests of the
Lydekker (1886.2, p. 97) treated this species as follows: third right inferior molar, r.Mj, are almost columnar in side \iew
"[Brit.Mus.] 41926-7. The last two ridges and talon of an un- and much more elevated than in »S. orientalis (Figs. 687, 779, and
worn fourth lower milk-molar and portions of two other cheek- 769). In front view the cranium (Figs. 77.3, 777) resembles that of

10 +i

Fig. 770. Referred Stegodon insigxis(?) = orientalis(?) and Type of Serri-

DENTINUS LTDEKKERI. AFTER ScHLOSSER, 1903, TaF. XIV, FlGS. 7-10
Fig. 7. Tooth referred by Schlosser to Mastodon a£f. latidens Clift, from the
rothe Thone = Schansi; fig. 10, a left inferior molar, VMs, from ?Fokien, referred by
Schlosser to Stegodon insignis.
Fig. 8. Type of Mastodon = Scrridentinus] hjdekkeri Schlosser, from the rothliche
[

Sande = Tientsin, Honan, north China, presumably a left M' (cast Amer. Mus. 10374);
fig. 9, supposed inferior premolar referred by Schlosser to Mastodon lydetcheri =Serri- [

dentinus lydekkeri of the present Memoir).

teeth, provisionally referred to the present species. These S. trigonocephalus (Fig. 776) and differs widely from that of S.
specimens, which were obtained from a cavern in Sechuen, north- insignis-ganesa or >§. bombifrons. This is a dwarfed insular form,
west China, are the types of Owen's Elephas (Stegodon) orientalis, very progressive in molar structure.
and are described and figured by him under that name in the History. —This species was named by Martin in 1890 and
They show,
(^uart. .lourn. fieol. Soc. vol. xxvi. pi. x.xviii. figs. 1-4. regarded by Janensch (compare Janensch, 1911, p. 171, fig. 12,
however, no characters by which they can be distinguished from also Taf. xxiii, fig. 3) as differing in skull structure both from
the teeth of the present species [i.e., Stegodon insignis], as the Stegodon insignis and S. ganesa while similar in jaw structure.
writer has already observed in the 'Pateontologia Indica,' ser. 10, The assigned ridge formula is: Dp 3* Dp 4- M l-M2-^M3^y^.
vu\. i. p. 269. P urchased from R. Swinhoe, Esq., 1870." Dubois in his Trinil-I-'auna (1908, p. 1256) observes that this
886 OSBORN: THE PROBOSCIDEA

species of Stegodon is highly characteristic of the Trinil Pithecan- STEGODON AIRAWANA FAUNA OF THE KENDENG-SCHICHTEN
LAYER, JAVA, AS LISTED BY STREMME, 1911
thropus beds, also that the third lower molar rises to fourteen
ridges, i.e., M
3 tt, and is more progressive than that of insignis. This fauna was first regarded as of Lower Pleistocene age
This would tend to place Pithecanthropus erectus as of Middle ( = Boulder Conglomerate zone of Pilgrim), or transitional to
Pleistocene age. Dietrich (letter, March 10, 1924, and notes), Middle [Upper] Pleistocene ( = Godavari Alluvium, Nerbudda of
as cited above in this ^Memoir (p. 813), on morphological grounds, Pilgrim) which contains Stegodon insignis ref., S. ganesa ref., and
regards this Javanese species as more recent than any of the Palxoloxodon namadicus. Stremme remarks (1911, p. 144): "Ein
known continental sjiecies of Stegodonts. wichtiges Leitfossil wiire eventuell Elephas, dessen Zahnbruchstiick
Stegodon Airdwnna Martin, 1890. "Ueber Neue Stegodon- Janensch dem Elcphas antiquus am niichsten stellt. Das Stiick
Reste Aus Java," Verh. Kon. Akad. Wetensch. Afdeel Natuurk.,
.Amsterdam, Deel XXVIII, p. 4, Type. Incomplete man- —
dible with thirtl molar of either side in place. IIouizon and
—
Locality. Kendeng-Schichten, Pithecanthropus erectus zone,
:Middle(?) Pleistocene,Alas-Tuwa, Trinil, Java. Type
—
FiGiHE. Martin, 1890, Tab. i, figs. 1 and 2 (mandible), also
Tab. 11, figs. 3 and 4 (type Ms).
—
Type Description. {Op. cit., p. 4) "Nur von Java in einer:

unvollstiindigen Mandibel und darin steckenden ]\Iolaren bekannt.

Letztere mit 9 Jochen und 2 Talons. Durch den niastodonartigen
C'harakter der Kronenspalte und die geringen Cementmengen
schliesst sich die Art an .S7. Cliftii und St. bombifrons, durch die
hoheren Joche an St. in.signis und St. ganesa an; sie nimmt somit,
gleich dem ebenfalls auf Java beschriinkten St. trigonocephalus
[Footnote: 'Vg\. Sammlgn. Ser. I, Bd. 4, pag. 102.'], eine Mittel-
stellung zwischen beiden Gruppen ein."
Janensch, 1911 ("Die Proboscidier-Schiidel der Trinil-Expedi-
tions-Sammlung") concludes his detailed description of the skull,
dentition, and skeleton of Stegodon airdwana with the statement
(p. 192) that while Stegodon insignis and .S'. ganesa are closest in
their dentition to S. airdwana, yet in certain details they are some-
what more primitive; also in skull structure S. airdwana differs
both from S. insignis and S. ganesa, while in jaw structure the
three species are similar.
Stremme ("Die Saugetiere mit Ausnahme der Proboscidier,"
1911, J). 143) observes: "Stegodon. Airdwana Mart, ist iiach
Janensch mit Stegodon ganesa und Stegodon insignis am niichsten
vcrwandt, die im indischen PliocJin und Plcistociin vorkonunen."
Janensch (1911, p. 192) determines the Trinil Stegodon as Stegodon
airdwana Martin, and observes regarding the ridge formula (op.
cit.. 1). 187): "Die Jochforniel [Footnote: 'In dieser Formel be-
deutct X den Talon.'] lautet, sowcit bis jctzt bekannt:

ax
Mm^ M X 7XX1IXX1
XI 3X
1-1

l3ubois gibt iieucrdings (Trinil-Fauna S. 125t]) an, class die

Zahl der Joche dei- letzten untcrcn Molaren bei dem Stegodon von
Trinil bis zu wenig.stens 14 gehe, doch ist aus seiner Angabe nicht
zu ersehen, ob er die Talons etwa mitzahlt." He also observes that
this si)ecies of Stegodon is highly characteristic of the Trinil
Pithecanthropus beds. The other Javan species, Stegodon trigono-
cephalus Martin, does not occur at Trinil.
Osborn, 1922: Our knowledge of this very progressive Javan
type has been greatly extended by the researches of Janensch
Fifj;. 77L Typo of Stegodon Airawann Martin, ISiX), 'Pah. i, fifjs. 1 ami
(1911) on the rich collection discovered by the Sclcnka-l^lancken-
2, oiic-fuurth natural size. The type rif;lit inferior molar, r.Mj, is al.so figured
li(ini I'Apeditioii in the Trinil beds in which tiie type of Pithecan-
in Tab. ii, figs. 3 and 4, two-thirds natural size. (Martin, op. eil., p. 4):
thropus erectus occuis. The fauna of these i)eds, as desciibed by "Nur von Java in einer unvollstiindigen Mandibel und darin steckenden
Strenune (1911, |ip. 82-150) includes (pj). 141, 142) the following. Molaren bekaiuil. Letztere mit .loclien und 2 Talons."
 ::: :

THE STEGODONTIN^: STEGODON 887

stammt nicht von Trinil; auch Dubois hat keine Elephas-Heste The last word by Dietrich (1926.1, p. 1.39) makes it still more
von Trinil in seiner grossen Saiiimlung." recent, namely: "Selbst wenn das Entwicklungstempo rascher ge-
worden ist, kommen wir Airawana zu einem sehr viel jiingeren
fiir

Primates Pilhccanthropus crcdus Dubois. Alter als bishcr fiir ihn und damit fiir die Trinilschichten fest-
Alacacus ncincstrinua mradana, related to exist- gesetzt wurdc, niimlich zu .Jung- bis Jiingstpleistocan. Diese Auf-
ing Zati of Sumatra and Ncmcutriuus of fassung lies geologischen Alters der Pithecanthropusschichten
Borneo. bahnt der \'on Dubois 192.3 gewonnenen Erkenntnis, dass Pithe-
Ungulata Stcgodon gancsa javanicus Dubois = [>S7c(;ti(/(»/( canthropus ein tilled der Hominiden ist, den Weg zu dem weiteren
Airdwana or .S'. trigonocephalus Martin]. Schritt, das P. bereits zur Gattung Homo gehort."
Elephas hysiidrindicus Dubois = Elephas sp. an- Osborn, 1928: It now seems probable that Stegodon airdwana
iiquus [namadicus] Falconer (Jide Stremme, is of lower Middle Pleistocene age, somewhat more ancient than

1911, and Janensch, 1911). the Godavari, Narbada Alluvium, and with a more early fauna,
[It is important to note that E. hysudriridicus Tafcl XXX IX.
does not occur at the locality of Trinil.)

Rhinoceros sivasondaicus Dubois, intermediate

between R. sivalensis and R. sondaicus}^^
Rhinoceros kendengindicus Dubois.
Tapirus pandanicus Dubois.

Artiodactyla Sus hrachygnathus Dubois, related to recent

(Sms verrucosus of Java.
macrognalhus Dubois.
.S((.s

Hippopotamus (Hexaprotodon) sivajavanicus

Dubois, related to Hexaprotodon sivalensis.

Cervuhis kendengensis Stremme, related to

existing muntjac.
Cervus (Axis) Uriocerus Duhuis = Ccrvus {Axis)
Lydekkeri INlartin.

Cervus {Rusa) kendengensis Dubois.

Cervus (Rusa) palseomendjangan Dubois.
Duboisia (Tetraceros) Kroescnii Dubois, re-

lated to existing Boselaphus and Tetraceros.

Leptobos Grocnereldtii Dubois.
Leptobos depend irornis Dubois.
}?nii;;^ Jlrwchcn.tuJtrcrzr{c-Turrc VuJJccui.rn.arLtcC
Bibos palxosondaicus Dubois, related to the
existing Bihos sondaicus.^^^
Bibos protocavifrons Dubois. a. Kartcnsl;"uzc der Umgebung vun Trinil. M.isstal) I : I 500000.
Buffelus {Bubalus) palseokerabau Dubois, re-
lated to existing Javan Buffelus."' Fig. 772. Sketch of the Kendeng horizon (vertical shading), Trinil, Java,
containing the type of Pithecayilhropm ereclus Dubois and the type of Stegodon
Rodentia Hysirix. gancxa var. javanicus {=S. airawana or S. trigonocephatus), also referred specimens
of Stegodon airawana. After Dubois, 1908, Taf. xxxix. Middle Pleistocene.
Edentata Manis palseojavanicu Dubois.

Carnivora : Mccecyon trinilensis, related to recent Mececyon

javanicus. including the rhinoceroses related to the R. sivasondaicus and
Felis oxygnatha Dubois. lacking the 7?. unicornis and Equus namadicus forms as well as the
Felis trinilensis Dubois. Palseoloxudon namadicus of the Godavari, Narbada Alluvium.
Felis rnicrogale Dubois = Feliopsis palxojavdnica Comparison of the beautiful figures of Janensch (1911, figs.
Stremme. 1-16, Taf. xxi-xxv) together with casts (Amer. Mus. 6835) indi-
Hyxna bathygnatha Dubois. cates that Stegodon airdwana is a much more progressive species
Lul7-a palseoleptonyx Dubois. than the S. insignis type of the Upper Pliocene [to Lower Pleisto-
cene], and referred of the Lower and Middle [Upper] Pleistocene;

More 1924) Dietrich determined that the

recently (letter, it is nearly as progressive as the Stegodon aurorse type from Mt.

species Stegodon airdwana, which the Kendeng horizon contains, Tomuro, Japan, as shown in the comparative figure (Fig. 688).
is somewhat more progressi\-e and consequently more recent geo- Consequently we are justified in accepting Dietrich's recent con-

logically than S. insigni.i-ganesa. clusion that Stegodon airdwana and Pithecanthropus erectus are

.we do not doubt that Stremme would have come to a |)lcistoceiie age, had he -as we— arrived at the conclusion that
". the
'((Maarel, 1932, p. 193) .

Trinil fauna contains at least three still living species viz., Bibos sundaicus fossilis, Buffelus Ijubalus var. sondaicus fossilis, and Rliinoceros sondatcus
fOSS His. "\
 : :

888 OSBORN: THE PROBOSCIDEA

of Middle rather than of Lower Pleistocene age. While the vertical It is in these respects the most specialized Stegodont of all

heightening or hy])sodonty of the molar crown approaches that of known si)ccies, a terminal form based upon a Stegodont foundation.
the Archidiskodon planifrons type, the entire conformation of the The dental formula of S. airdwana runs as follows:
S. airawana cranium (Fig. 773) is totally different
strates afresh that the most progressive Stcgodontinw arc parallel
and demon- Stegodon airdicana : Dp 2 ? Dp 3^ Dp 4 ^M 1 -^ M2 ^
with, rather than ancestral to, the Elephantidse. The greater or lesser development of the ridge formula de-
Sfteiika-Triiiit'Erii^iliitH.
pends upon the anterior and posterior half-ridges or talons, as in
the reckoning of Falconer and Lydekker. In contrast [Osborn] the
maximal ridge formula of the best-known continental .species,
Stegodon insignis, is as follows
7»
[Stegodoyi insignis-ganesa : Dp 3 f Dp 4 714-9 "*^ • 7M-10
AI2 8-8H
M3
M-9-W -^'^ " 12W-X3J-

—
Length of Molaes. In the following table the characteristic
maximal and minimal molar tooth length is expressed in millimeters

LTpper Molars Dp-

Fig 1-2. S'jorfon Airawana Makt.

Vctbg von Wilhelm Engclmann id I.eipiig

I'"if;. 773. Frdiit, and side views of referred Stegodon airawana Martin, as
fi|i;uied by Jancnsch in his memoir of the Selenka-BIanckenhorn Trinil-
Expcdition, 1911, Taf. xxi, figs. 1 and 2. One-sixth natural size.

COMPARISON OF STEGODON AIRAWANA MARTIN WITH

STEGODON INSIGNIS-GANESA
Translated from the German of Dr. W. O. Dietrich with interpolations l)y

H. F. Osborn and oorrected by Doctor Dietrich, April 18, 1021

Stegodon airdwana Martin from the Pleistocene of middle

Java (Pithecanthropus beds of Trinil) manifests evidences of insular
dwarfing; it is in all its parts, both of teeth and skeleton, smaller
tlian the continental west Asiatic and Indian races and s])C('ies.
Si)ecial features of the .S. airawana teeth include the following char-
acteristics: (1) Shallow longitudinal cleavage of the crowns; (2)
relatively slender and narrow ridge-crests; (3) strong division of
the ridge-crests into numerous mamillae [conelets], relatively
limited cement covering; (4) small, thin, and strongly folded
enamel on the abrasion sui'faces. (5) While Stegodon airdivana is
a dwarfed lroi)ical form as compared with the giant S. insignis-
ganesa, it exhibits more numerous, more elevated, and more com-
pressed ridge-crests.
 THE STEGODONTIN.E: STEGODON 889

(leteiinine positively the progressive development within the Stego- No. 6, p. 1257. Tvi'K. — Material from Keudciiu;. TTom-
dont series. In order to measure the specialization of the molar ZON AND Locality. — Trinil, Kendeng Schichten, Middle Pleis-
teeth, including anterior addition and reduction and posterior tocene, Java.
addition and reduction, a relative proportion may be established Description {Op. cit., 1908, pp. 1256, 1257). -" Die Untersuch-
between the development of the two anterior teeth and the two ung eines sehr reichlichen Stegodonten-materiales aus dem ganzen
posterior teeth. It is noteworthy that the proportion between the Kendeng, unter welchem auch mehrere Schadel von Jungen und
first and second molars in the Chine.se and Javanese .species is the alten Tieren, hat mieh nun zu dem Ergebnisse gefuhrt, dass alle
same. From careful calculations it follows that M- in airdwana iS'. diese l^eberreste einer einzigen Art angehiJren. Diese unterscheidet
is more strongly reduced than in the corresponding tooth of the sich \on Stegodon ganesa kaum anders als durch ihre viel ger-
S. orientah's of Owen. ingere Grosse. Der Schadel besitzt dieselbe weite Temporal-
On all these structural grounds Doctor Dietrich concludes that grube, ist ebenso charakteristiscli stark brachycephal und auch
S. airdwana is more recent than either of the Asiatic species of the iibrigens sind beide Formen sehr iihnlich. Nur iiltere Schadel
continent and that its newer Pleistocene age is rendered certain. weichen in einigen Beziehungen etwas von der typischen Ganesa-
Form indem niimlich die Frontal- und Occipitalteile sich
ab,
gegeneinander abflachen und mehr oder weniger scharf von
STEGODON GANES.\ VAR. JAVANICUS DUBOIS, 1908
einander getrennt sind. Das braucht uns abcr nicht davon
Middle Pleistocene, Triiiil, Kendeng-Schichten, Java
abzuhalten diese Formen derselben Art zuzuschreiben, denn es
This subspecies, Stegodon ganesa javanicus, belonging in the kann ja bei den Elephanten die Schadelform einer und derselben
same Kendeng beds as .S. airdwana Martin, 1890, is to be regarded, Art, innerhalb gewissen Grenzen, betrachtliche Verschiedenheiten
as remarked by Stremme, as a synonym of airdwana.'- .S'. zeigen. Der Ganesa-Typus bleibt bei unserer Form doch immer
"Das Geologische
In his paper of 1908, entitled Alter der erkenbar."
Kendeng-Oder Trinil-Fauna," Dr. Eugen Dubois, the discoverer "Auch durch ihre Molaren ist sie von S. ganesa spezifisch
of Pithecanthropus erectus, assigns to Pithecanthropus and the nicht zu trennen, nur geht die Lamellenzahl im Unterkiefer bis
accompanying Kendeng-Trinil Fauna at the most a Lower Pleisto- zu wenigstens 14 anstatt 13; die Zahl der Lamellen hat aber,
cene age ("alt-diluvialen Alters")- The remains occur in the wie wir durch Pohlig wi.ssen, fiir die Unterscheidung der Elephan-
fossil-bearing tufa known as the "Kendeng-Schichten," in the tenarten nicht die grosse Bedeutung welche man ihr friiher zu-
center of which lies Trinil, as shown in Plate xxxix
772 of the (Fig. schrieb."
present Memoir). Recently Dietrich has assigned a Middle "Zu dieser Art gehoren auch die zwei von M.\rtin als
Pleistocene age to the Kendeng Trinil fauna. Stegodon trigonocephalus beschriebenen Schadel. Waren
. . .

Dubois (1908, j). 1257) erroneously remarks that the two diese gut erhalten gewesen, so hatten sie erkennen lassen, dass
skulls described by Martin as S. trigonocephalus really belong die Form keine urspriingliche und der Art eigen-
dreieckige
specifically to Stegodon ganesa in their cranial and dental characters, tiimliche Der darauf Bezug nehmende Name ist also zu
ist.

but in order to distinguish this smaller Kendeng species, as an kassieren. Ich schlage nun vor, diese kleinere Kendeng-Form
insular variety, from the giant continental form of ganesa, he nur als VarietJit von der riesigen Festlandsform, der sie deutlich
proposes the subspecific name Stegodon ganesa var. javanicus. sehr nahe steht, zu trennen, ftihre sie also als Stegodon ganesa var.
He considers it probable (1908) that this Javan subspecies be- javanicus ein."
longed to the Upper Pliocene fauna. Characters {Op. cit., 1908, p. 1257).
— "Wenn man nun auch
Stremme, "Die Siiugetiere mit Ausnahme der Proboscidier," nicht zugeben kann, dass Stegodon ganesa und Stegodon insignis
in "Die Pithecanthropus-Schichten auf Java," 1911, p. 142, regards nahe
einer Art angehoren, so sind beide jedenfalls einander sehr
this subspecies as closely related to Stegodon airdwana Martin, verwandt Molaren sind nicht oder kaum zu unterscheiden),
(die
1890. Stegodon insignis aber kann man, mit M. Schlosser ), . . .

According to Dubois (1908) the two skulls described by geradezu als das Leitfossil der jlingeren Pliociinfauna Ostasiens
Martin as Stegodon trigonocephalus belong to this species." It is ansehen. Es ist schon hierdurch wahrsclieinlich, dass unsere
important to Ciote that the true .S. trigonocephalus of Martin does javanische Stegodon ganesa der gleichen jungpliocanen Fauna ange-
not occur in the same level as Pithecanthropus erectus. hort. Unzweifelhafte Ueberreste von Stegodon ganesa sind nun
"Das Geolog-
Stegodon Ganesa var. javanicus Dubois, 1908. aber noch nicht in jlingeren als Pliociincn Schichten angetroffen
ische Alter der Kendeng-Oder Trinil-Fauna." Tijdsch. Konink. worden, denn der in den Narbada-Schichten gefuudene Stoss-
Neder. Aardrijks. Genoots. Amsterdam, Tweede Serie, Deel xxvb, zahn kann man dieser Art nicht mit Sicherheit zuschreiben ..."

'[The following note was prepared by Dr. George Gaylord Simpson (October, 1937): "Stegodon trigonocephalus Martin, 1887, pp. 27, 36, 41, was
founded i)iincipally on two skuHs, Professor Osborn selecting tlic younger as type. These were probably from the vicinity of Surakarta (Solo), Java (see
Martin, 1S87.1, i)p. 2.5, 27). Duljois briefly cliaracterized abundant material from Kendeng, referring it to Stegodon ganesa but a.s a new variety javanicus
(Dubois, 1908, pp. 1256, 12.57). He also referred S. trigonocephahis to S. ganesa. He probably considered trigonoceplialus as the same as his vari(>ty javanicus

but he did not explicitly say so and the clear implication is tliat they luivc different types. Von Koenigswald considers S. trigonocephalus as distinct from
S. ganesa but the same as S. airawana (1933, pp. 103-105). He implies that S. ganesa var. javanicus is the same as S. trigonocephalus and S. airawana and
applies to the species the oldest of the three names, S. trigonocephalus. Maarel, however, considers .S'. trigonocephalus to be distinct from S. airawana. and this
was Professor Osborn's opinion. It appears, in any case, that S. ganesa var. javanicus Dubois, 190S, is synonymous with S. trigonoceplialus, S. airawana,
or both, and as both names antedate it, Dubois' name is invalid." Editor.] —
890 OSHOKN: THE PROHOSCIDEA

Stegodon trigonocephalus Martin, IS.Si Stegodon trigonocephalus Martin, 1887. "Fossile Siiugethier-
figures 70."), 731, 77."), 77(1, 777, I'l. xx reste von Java imd ,l!i]);m," Samrnlung. (leolog. l{eichs-Museimis,
Probably vicinity of Surakarta, Java. Dubois notes that Stegodon Ijeiden, Beitriige z. (ieolog. Ost-Asiens und Australicns, f^er. I,

trigonocephalus docs not occur intlie Kcndeng-Schichten of Trinil; Matsu- Bd. IV, Heft 2, pp. 27, 36 (1887); also Jaarb. Mijnw. Neder.
moto regards the geologic level of »S. trigonocephalus as equivalent to the Oost-Indie, 1887, Wetensch. C!ed. 16, Palaeontologie van Neder-
Lower Pleistocene, Boulder Conglomerate bods of India. laudsch-Indie. Verhandeling No. 21, pp. 3, 12(1887). Type.—
Stegodon trigonocephalus is a dwarfed insular Stcgndont, based Young skull with third and foiu'th deciduous premolars in the
chieflyupon an immature cranium (Fig. 776) which has sub- CJoologicalMuseum of Leiden. Hokizon and Locality. —
stantially the same characteristic insular form as the immature Probably from vicinity of Surakarta, Java; said to be associated
cranium of S. airawana (Fig. 773), but which is very different from with remains of proboscideans referred to Stegodon bombifrons,
Eiielephas namadicus, and Eiielephas hysudricus. Ty'pe
4j?l.^r-^^^ —
Figure. (Op. cit., Martin, 1887, Sammlung. Geolog. Reichs-
Mu!5eums Leiden, Tab. ii, figs. 1, la, and Tab. iii, fig. 1.
Description. — The original description by Martin (1887, pp.
36-41, with figures) gives the characters of the immature type
skull and teeth in great detail, and on page 4.5 offers the follow-
ing distinctions from Stegodon and .S. bombifrons: "Zuniichst
cliftii

ist es von Bedeutung, dass die Ziihne von denjenigen des Stegodon
voUig verschieden gebaut sind, dass eine Vereinigung der
Cliftii so
Fig. 775. Stegodon trigonocepliolus. Rather young cranial profiles javanischen Art mit der genannten, von der ein Schiidel noch nicht
drawn directly from type. Frontal line much longer than in S. pinjorensis.
bekannt ist, von vornherein ausgeschlossen wird. Ebenso be-
Cranium extremely abbreviated. Skeleton conjectural, as in other Stegodonts.
Restoration by Margret Flinsch (1930), under the direction of Henry Fairfield stimmt unterscheiden sich aber (soweit unsere Kenntniss bis jetzt
Osborn. One-fiftieth natural size. reicht) audi die Zt'ihne von St. trigonocephalus durch die Jochzahl
der Praemolaren von alien anderen, bis jetzt bekannten Art.en von
the crania of »S'. bombifrons or »S. insignis-ganesa. This ju^•enile 2**^"
Stegodon, . . . Bezeichnend Praemolaren von St.
fiir den
type represents the immature condition of the Stegodont cranium
trigonocephalus ist Form, und ebenso fiir den
ferner seine ovale
and consequently is very interesting and important. It is now re- gten
pi<^pniolaren die kaum mcrkliche Convergenz seiner Seiten-
garded as in a Pleistocene stage of evolution, but the extremely
fliichen nach vorne zu, Merkmale, die von alien bekannten, ent-
immature condition of the teeth renders its specific determination
very difficult. F'rom the following description and figures of sprechenden Ziihnen der iibrigen Stegodontenarten abweichen.
Martin, it is difficult to distinguish this immature type from Stego- Durch die hohcn, schmalen Joche und vor allem durch die feine
don airawana; it is apparently a somewhat more primitive Lower Fiiltelung diirfte audi bei Bruchstiicken der ^Nlolaren von Si.

Pleistocene form. trigonocephalus eine LTnterscheidung von St. bombifrons bisweilen

..'^". '\.

^iiUsihi'^^

Fig. 776. Type of Stegodon trigonocephalus Martin, 1887, Tab. n, figs. 1, la, and Tab. in, fin. 1. The side and front views are reproduced one-eighth
natural size; the crown view of the tooth reproduced one-third natural size.
is

Taf. I, figs. 3 and 4 (Naumann, 1887). A single crest from .lava, distinct from ,S'. trigonocephalus and d()ul)lfully referred by Naumann to.S'. insignis Falc.
ami Caut. (op. cit., p. 9): "Das Zahnbruchsfiick kann nur zu Si. in.'iignis oder zu St. Gane.ia gehoren."
 THE STEGODONTIN^: STEGODON 891

Fig. 777. Front and side view outlines to a one-sixteenth scale of Stcgodont crania of:

A, Al, Stegodon pinjorensis type, mature adult (Amer. Mus. 19772), from near Siswan, India. See also figure 765.
B, Bl, Stegodon bombifrons cotype, mature adult (Brit. Mus. M. 2979, cast Aracr. Mus. 10378). See figure 734.
C, Stegodon orientalis grangeri ref., young adult (.\mer. Mus. 18(530) collected by Walter Ciranger, Wanhsien, Province of Szp<'huan, China. See figure 763.
D, Stegodon orientalis grangeri ref., infantile (Anicr. Mus. 18032) collected by Walter Granger, Wanhsien, Pnivince of Szechuan, China.
E, El, Stegodon trigonocephalus tyjie, juvenile, from vicinity of ?Surakarta, Java. Sammlung. Geolog. Reichs-Museums, Leiden. Sec figure 776.
F, Fl, Stegodon airhwana ref., juvenile, Java. See figure 773.

ermoglicht werden; dagcgen ist die Trennung solcher Reste \on immature type skull and to a second older skull which is a referred
St. insignis und ganesa mit sehr grossen Schwierigkeiten verbun- specimen, namely: {op. cil., p. 41) "6. Alter Schddel. (Tab. iv u.
den." V Fig. 1)."
Ridge formula of S. trigonocephalus: Dp 3- Dp 4^^ as compared A specimen from northwest Mindanao, Philippine Islands,
by the author with S. diftii: Dp 3^ Dp 4-; with .S. bombifrons: referred to this species is described in detail by Dr. Edmund Nau-
Dp3,^^Dp4^^^''; with.S. ^a/ifsa: Dp 2^ Dp 3- Dp 4^. The skull, mann of IMunich University in his "Fossile Elephantenreste \'on
however, is quite different in form from that of S. insignis. Martin Mindanao, Sumatra und Malakka," Abhand. u. Berichte des
inclines to compare S. trigonocephalus with S. insignis rather than Konigl. Zoolog. Anthrop.-Ethnog. Museums zu Dresden, 1887,
with the Lower [Middle] Pliocene stage of S. bombifrons. pp. 5-8. He subsequently (1890) made it the type of Stegodon
It is important to note that these characters relate both to the mindanensis.

StEGODONTS OF THE PHILIPPINE ISLANDS AND JaPAN

We owe to Naumaiin (1890) and to Matsumoto (1915, 1918) the discovery and description of three very
progressive species, originally referred to Stegodon, which in part or wholly seem transitional in structure to
ArcMdiskodon or to Elephas. The true affinity of these types rests upon (a) the still unknown structure of the
cranium and (b) the ridge formula and height and form of the ridge-crests as now revealed in the case of the species
Stegodon aurorse (Fig. 781). In brief, until the cranium is known, we cannot be certain whether these animals are
progressive Stegodonts or primitive Archidiskodonts. The history of discovery and description is as follows.
 :

892 OSBORN: THE PROBOSCIDEA

Stegodon (Archidiskodon?) mindanensis Naiiiu.imi, 1S9() was doubifuiiy referred by Naumann to S. iiisignis or .S. ijniicsa
•'iguics 700, 77S Falconer and C'autley. (Naumann, 1887, p. 9): "Das Zahn-
Mindanao, Philippine Ishiiuls, Lowci(?) to Middle Pleistocene. bruchstuck kann nur zu St. insignis oder zu St. Ganesa gehoren."
If a true Stegodont, Stegodon mindanensis (Fig. 778) is even
Stegodon Mindanensis Naumann, 1S90. "Stegodon Minda-
more progressive than S. airawana, because the valleys between
neue Art von Uebergangs-Mastodonten." Zeitschr.
nensin, eine
the ridge-crests are entirely closed up; it compares somewhat more
(ies., Bd. XLII, Heft 1, pp. 166 169.
deutsch. geol. Type.—
Fragment of a molar tooth. Horizon and Locality. — closely with Stegodon aurorx (Figs. 780, 781), also an imperfectlj'
known sjjecies possibly referable to Archidiskodon.
Mindanao, Philippine Islands, (?)Lower to Middle Pleisto-
very difficult to determine the characters
Osborn, 1924:
cene. —
Type FiGriu;. Naumann, 1887, Taf. i, figs. 1 and
It is
of this type (Fig. 778) without examination of the original speci-
2. —
Type Description. (Naumann, 1890, p. 167): ". . . so
men. The type lower molar is far more progressive in the direction
bleibt (loch das friiher erzielte Rcsultat, nach wclchem (lurch die
of Archidiskodon or Elephas than the types of either Stegodon
insignis or S. ganesa; it is, in fact, a true-crested tooth in which
the ridges are closely compressed and the valleys closed. Conse-
quently this tooth should be compared with that of a primitive
species of elephant or of Archidiskodon.

Stegodon auroras Matsimioto, 1915, 1918

Figures OSS, 709, 7S0, 781, S19
TvpE OF Stegodon (archidlskodon?) mindanensis
Uppci(?) Pliocene, Mt. Toniuro, Kaga, Japan.
Fig. 778. Type incomplete inferior molar of Stegodon 7Hi}idanettsis
Naumann, 1890, from Mindanao, Philippine Islands (originally figured by Like Stegodon mindanensis, Elephas (Prostegodon, Parastego-
Naumann in 1887, Taf. figs. 1 and 2, as Stegodon tiigonocephalus).
i, don) aurors- is either a highly progressive Stegodon or a primitive
.•\scompared with Stegodon airawana (Fig. 77S), the ridge-crests in Archidiskodon, a point to be determined positively l)y the discovery
S. mindanensis type are more vertically placed, as in the Stegodon aiirorx
of a cranium.
typo (Fig. 780).
Elephas (Prostegodon, Parastegodox) auuor.e Matsu-
MOTO. — In describing the type specimen, determined as a second
Untersuchung der beiden Zahnbruchstiicke von Mindanao 'die right superior molar, r.M^, Matsumoto remarked (1918, p. 52)
Verbreitung der Siwalikfauna iiber das Ciebiet der Philippinen ". . . this specimen represents a species new to science, being ... a
bewiesen und die enge \'erkniipfung einer wahrscheinlich jungter- transitional form from Stegodon to Elephas." Originally described
tijiren Saugethierfauna auf Java und den f^hilippinen durch eine as Elephas aurorsc Mats., 1915, also 1918, j). 52, Pl.xx, it was after-
in der Entwicklungsreihe der Stegodonten und EIe])hanten ward referred by Matsumoto (in Osborn, 1923.601) to Prostegodon,
hochwichtige Art' constatirt sein .sollte, zu liecht bestehen. . . the genotype of which is Ma.<itodon latidens, and finally (1924.2)
^lerkwiudig ist ferner ein medianer Einsclmitt der Krone, der was made the genotype of Parastegodon Matsumoto, to include
jederseits \'on einem secundiiren Einschnitt begleitet wird. Durch the Stegodon mindanensis-E. atirorx group.
diese Spaltungen werden die Mamillenreihen in (Iruppcn zerlegt." Elephas aurora: Matsumoto, 1915, 1918. Preliminary reports
The type of this species was originally figured by Naumann in of this species were published in the Scientifie (Jazette, Tokyo,
1887 as Stegodon trigonorephalus; it was regarded by him as an Vol. Ill, No. 5, 1915, pp. 308 315, and in the Journal of the
outlyer of the Siwalik proboscidean fauna. A single ridge-crest Geological Society, Tokyo, Vol. XXIII, No. 275, 1916, i). 294
fiom Java (see I'^ig. 776, lower), distinct fiom S. tiigonocephalus, (both in Japanese). Also (in English) "(^n a New Archetypal

Fig. 779. Kcferrcd by Jancnsch to S. airaivnnn. Nine ridge-crested second superior molar, M'-', after .laiicnscli, 191 1, and ridge-crest section
p. 171, fig. 12,

of a third superior molar, M^, after Janensch, op. cit., p. 174, fig. 13. One-half natural size. Both figures inverted to show the molars in natural jmsition.
(Left) Lateral view. (Right) Ridge-crest section.
It be observed that the convexity of the ridge-crests of the .second molar crown, resulting in the wide divergence of the cre.sts, together with the more
will
numerous ridge-crests, tends to relate this animal to Stegodon rather than to Archidiskodon, in which the ridges are more vertical and more closely compressed.
 - —

THE STEGODONTINi^^.: STECIODON 893

Fossil Elephant from Mt. Toimiio, Kuga." Sci. Uei)f. Tulioku

Imp. Univ., 1918, (2), 111, Xo. 2, p. 52. Typk.— A right AP
with ten and a half ridges, thus exceeding the ridge formula of
JM- in Archidiskodoit planifroits, namely, 2^^. Original in M
Geological Institute, Tokyo. Horizon and Locality. Mt. —
Tomuio, Kaga, Japan, (?)Upper Pliocene. Type FiorRE.
Op. cit., 1918, PI. XX, figs. 1-3. SCPPLEMENTARY DESCRIPTION
(Matsumoto, 1924.2, pp. 256, 257, 262).— Made the genotype of
the genus Parasiegodon (see p. 903 of the present Memoir).'
Type Description (1918, pp. 52-55).
unique type
— "The
specimen corresponds to an ujiper, probably penultimate, molar
of the right side. It measures 180 mm. in length, 75 mm. in
maximum width and 48 mm. in the maximum height of the crown."
The reasons why the autiior referred this new species to Elephas
and not to Strrindon arc as follows: The ridge formula of M-
corresponds to 10,'i>, while that of the known species of Slegodon
corresponds to 6 to 9. The author gi\es six additional reasons
for separating the animal from Stegodon, and six additional reasons
for separating it from E. planifro/is, to which he regards
specitically
it as allied generically. The author concludes (p. 55): "The
present species appears to the present writer to be more archety-
pal and more Stegodon-\ike than E. planifruns in the second distinc-
tive characteristic, but just the opposite in the third, fifth and
sixth distinctive characteristics. At any rate, the discovery of the
present species is worthy [to] be considered as an additional
datum to pro\e the very intimate alliance of Stegodon and Elephas."
Type of Stegodon auroRvE
I"ig. 7S0. Tyi)C of Elephas {Piostcgodon, Parasiegodon) aurorx Stegodon orientalis shodoensis ]\Iatsumoto, 1924
Matsumoto, after Matsumoto, 1918, PI. xx, figs. 1 and 3. One-half Middle Pleistocene. I.sland of Mitsugo (Mitsugo-shima) and Island of
natural size. Determined by Matsumoto (1918) as a ten and a half Shodo, Inland Sea, Japan.
crested second right superior molar, r.M". Compare figures 699 of S.
Slegodon orieidalis tshodocnsis Matsumoto, 1924. "Preliminary
bombifrons and figure 762 of 5. orientalis grangeri showing convex
external side aTid flat internal side; also figures 6S6, 7, and 6SS. Notes on the Species of Stegodon in Japan." Journ. Geol. Soc.
Tokyo, XXXI, Xo. 373, pp. 333-335. Type.— A fragment of
skull— left and right upper jaw bone with third molar attached.

Uyeno Museum of Tokyo, No. 2194, presented by Mr. Tomekichi

Ozaki. —
Horizon and Locality. ]Mitsugo-shima, Yoshima-
mura, Xakatadotsu-gun, Province of Sanuki, Japan. Milazzian-
Tyrrhenian. ]\Iiddle Pleistocene. Type Figure. Type —
figure not seen by the jiresent author.
Description in Japanese by Malsiiinolo; Iranslation by Messrs.
Shoichi Ichikawa and Ushinosuke Narahara. —
(Matsumoto, 1924.3,
pp. 333-335). "Materials: A fiagment of skull: left and right
upper jaw bone \vith third molar attached (found at Mitsuko-
shima, Yoshima-mura, Nakatadotsu-gun, Province of Sanuki.
Specimen No. 2194, Uyeno Museum of Tokyo, presented by Mr.
Tomekichi Ozaki). A small fragment of third molar (found at
Shodo-shima and the property of the Tokyo Imperial LTniversity).
Third upper left molar (found at Shodo-shima, the cast is kept at
Fig. 781. Vertical section of type molar, r.M-, of Stegodon aurone Kyoto Imperial University). A fragment of an upper palate with
(compaie Fig. 780, crown and side views of tyiie of Elephas (Prostegodon,
third molar of left and right attached (found at Shodo-shima, the
Parasiegodon) aurorse, also Fig. 82o, lectotype of E. = .Archidiskodon]
[

planifrons). One-half natural size. After ])hotograph sent by Doctor

property of Kyushu Imperial L'ni\'ersity and the cast kept at
Matsumoto. Kyoto Imperial University)."
HSee also Matsumoto's article (in English) "On Parastcgodon Mastumoto and its bearing on the Descent of Earlier Elephants," Sci. Kept. Tohokn Imp.
Univ., (2), XIII, No. 1 (Matsumoto, 1929.3).— Editor.!

-[Dr. Jiro Makiyama has recently reviewed the Japanese Proboscidea (see Makiyama, "Japonic Proboscidea," 1938, Mem. Coll. Sci. Kyoto Imp. Univ.,
(B), XIV, No. 1, pp. 1-59), in which he figures (p. 18, fig. 7) the holotype palate of Slegodon shodoctisis, with right an<l left third molars in situ. On page 19 he
states that "The material of Matsumoto was not enough to give full details of the species, but it was named before Parasiegodon akashieiiMS Takai, 1936, which
seems to me to be in the nearest relation, as it may have a new name form S. shodoensis akashien.fis." This subspecies he describes on pages 21 to 27, including
three figures (Figs. 10-12).— Editor.)
894 OSBORN: THE PROBOSCIDEA

"Ridge formula of molar: left, a+9X = }i-Q-}Q; right,

l
a + 8X. The exact measurements of ridges cannot be determined.
a+8X = ]4-8-}i\.
[ Tlie exact measurement of ridges cannot lie Length, 205 mm., right 20U mm. tireatest width, 85 mm. at
left

determined. The length as it is, left 226 mm., right 215 mm. The 6th ridge from back (left), 84 mm. at 5th ridge from back (right).
greatest width, left 97 mm. at 2nd ridge, right 96 mm. at 3rd Height, 45 mm. at 3rd ridge 50 mm. at 2nd ridge (right).
(left),

ridge to 6th ridge. The height of crown, left 52 mm. at 6th and There are four ridges within 100 mm. The interlocking occlusion
7th ridges, right 49 mm. at 7th ridge. The 6th ridge of left, and of the molar is elephant type."
7th ridge of right shows the beginning of wear. There are four
. . . "A small fragment of third molar, which shows three ridges.
ridges within 100 mm. The length of each ridge is 25 mm. and I cannot determine whether it belongs to the upper or lower jaw.

this is rather narrow as a width for third molar of Stegodon. The The length from inside to outside of each ridge is 90 to 96 mm.
valley is narrow, and the entrance of inside of valley has supple- The width of front to back is 24 to 25 mm."
mental cusps. The fold of enamel is very regular and fine. The "Among these specimens, the first has comparatively low
cement is well develoi)ed. The interlocking of opposing surfaces crown and the second and third have very high crowns. The
of themolar is of Mastodon type." fourth specimen is rather low as absolute value but as a ratio it is
"Second example: Ridge formula may probably be XllX. high. The molars are different in sizes, and this differentiation
Greatest width, 93K mm. at 6th ridge, height, 55 mm. at 7th ridge. (or varieties) is the one which occurs in the last period of the

There are four ridges within 100 mm. Judging by the measurement, existence of the species."
it looks like a second molar, but by the .shape of its being attached "These specimens resemble the Stegodon insignis which is

to the palate is a third molar. Ridge formula, left a+9X ; right found in the Narbada Valley of India."

V. RECENTLY DESCRIBED STEGODONTS FROM JAVA, CHINA, AND JAPAN

[The following species were described resi)ectively by Dr. Franciscus Hendricus van der Maarel ("Contribu-
tion to the Knowledge of tlie Fossil Mammalian Faima of Java," 1932), by Dr. G. H. Ralph von Koenigswald
("Beitrag zur Kenntnis der Fossilen Wirbeltiere Javas," 1933), by Dr. Shigeyasu Tokunaga ("Fossil Elephant
teeth foimd at Yokohama and Kakio, Kanagawa Prefecture," Journ. Geog. (Tokyo), XLVI, No. 546, 1934, and
"A New Elephant found in Shikoku, Japan," Proc. Imp. Acad. Tokyo, XI, 1935), by Dr. Chung-Chien
Fossil
Young("Miscellaneous Manmialian Fossils from Shansi and Honan," 1935), and by Dr. Arthur Tindell Hopwood
("Fossil Proboscidea front China," 1935). While the species of Stegodonts in these pul^lications (other than
Parastegodon? kwantoensis and Parastegodon sugiyamai of Tokunaga) were noted by Professor Osborn, they
were not studied intensively by him, consequently they are hsted here together with excerpts from the original
descriptions, including type figures, but without comment or determinations by the author of the present

Memoir.
Doctor Hopwood states on page 103 of his Memoir that "Insufficient is known of the Stegodontidie of China
to make a comparison with those of India very profitable. At present it seems as though they occur earUer in
India than in China, where they are a late invasion from the South. Palaeoloxodon, on the other hand, may be
more recent in India than in China." Editor.] —
Stegodon bondolensis van dor Maarel, 1932 "The diastema descends obliquely forwards at an acute angle
Figure 782 of about 35° with the inferior border of the ramus. The mandibu-
IJondoI nciir KiiwiiiiK, district RiuMiiiblatiiiif;, Regency niora, Residency lar symi)hysis is produced into a relatively broad, spoutlike
Remhaiig, Java. termination. Its lower border is damaged. The outer surface
. . .

van der Maarel, 1932. "Contribution to

Stegodon bondolensis of the horizontal ramus presents two foramina mentalia, the one
the Knowledge of the I'ossil Mammalian Fauna of .Ia\a," ])p. situated some 8 cm. below the anterior extremity of the alveolar
158-164.
—
TvPK. "Fine fragment of the mandible, com- margin, the other about 8 cm. in advance of the former and nearer
prising the whole of the horizontal ramus and the lower ])ortion of to the inferior bolder of the ramus."
the ascending ramus of either side; containing on either side "Only the lowci- portion of the coronoid jirocess has been
a molar which is inferred to be the M3 from the absence of the jtreserved."
indication ofany other tooth behind." Honizox and "Measurements of lower jaw in cm.
—
Locality. Bondol near Kuwung, district Kandublatimg, Regen- See text fig. 24.

cy Blora, Residency Rcmbaiig, .la\ a. T'i i'k Fkukk. Op. — greatest height = 16
In plane I
n't., PI. XIV, figs. 1, 4, 5, also text figiu'os 24 and 25. greatest width = 7.8
Dkscriptiox.— (Van der Maarel, 1932.1, pp. 158-164): "The greatest height = 12
In i)lane II
mandible is broken into two in the symphysis, but botii parts greatest width = 17
match exactly. 'I'he alveolar border of both molars is dninaged." Length of RC =47.5"
 THE STEGODONTIN^: STEGODON 895

"As the front .sideof both duinugeil,

moliii's is seems — ut it can, however, only be .seen in the last three ridge-crests. In this
first sight — impossible to ascertain the true number of ridge-crests connection attention may be drawn to the fact that in the mohirs
carried by each of them. Fortunately, however, in the I.M3 a small of the present sjiecimen enamel, cement ami the greater part of the
portion of the front surface of the fangs has been preserved. It is dentine had the same irregularly spotted appearance. As a result
situated below the anterior side of the foremost ridge-crest, and the crenulation of the enamel was very indistinct, and it was only
proves with absolute certainty, that this ridge-crest is really the possible to make it distinctly visible in the photograph by black-
first one. That is to say: not a single ridge-crest has been lost. ening the dentine of the tooth. Another result is, however, that

The ridge formula is, therefore, 8X." 'Stufenbildung' though present, is in general not clearly exhibited."

Fig. 782. Type mandiblo and enlargement of third left inferior molar of Slegndon bondolensis. After
van der Maarel, 1932, PI. xiv, figs. 1, 4, and 5. Molar about three-fifths natural size; mandible about
one-seventh natural size. From Bondol, Java.
(Upper) Crown view of third left inferior molar, I.M3, with 8 4- ridge-crests; same molar as in ac-
companying lower jaw.
(Lower, right) Superior view of fragmentary lower jaw with right and left third molars in situ.
(Lower, left) Right lateral view of same jaw.

"The tooth is distinctly curved outwards. The base of ridge- "There no indication of the presence of a median longitudi-
is

crests 1-7 are more or less of the same length from ridge-crest; nal cleft. Cementis present, though in small quantities. The
8 inner and outer side of the molar converge suddenly back- valleys between ridge-crests 1, 2, 3, 4 and 5 possess but little
wards." cement. More backwards the amount is somewhat less small, not
"The base of the crown and the grinding surface are strongly only the base of the valleys being filled to a greater extent, but
concave. The grinding surface slopes obliquely from the outside also the lingiuil and buccal sides of the ridge-crests, and the
inwards." posterior side of the hind talon being covered by a thin coat of
"The enamel is moderately thick and consists of two layers cement. Noteworthy is the stronger, local de\'elopment of cement
(see the buccal cusps of ridge-crest 7). A distinct 'Stufenbildung' near the buccal side of ridge-crest 7."
896 OSBORN: THE PROBOSCIDEA

"MeASUUEMKNTS of U. and L.M3 IN MM. schcinend weniger an Ihihe zunalun, als os bei der (ypischen Form
der Fall ist. Die Ictzteii drei .lochc und der Talon liegen noch
Total length of r. M3 190 mm.lnicasured along lucilian line
dick unter Zeraent."
Ditto " 1. Ms 197 "
J
of crown."
"Diesen letztaufgefuhrten Zahn bilde ich Taf. xxvii ab,
"As a matter of fact all the species of Stegolophodoti may be neben dem entsprechenden Zahn von St. t. trigonocephalus von
left out of consideration, being all very primitive forms which in Pitoe, nahe Trinil. Dieser letztgenannte Zahn ist ca. 270 lang mm
my opinion may as well be reckoned to the family of the Maslo- imd gehorte einem mittelgrossen Tier. Zwei letzte Molaren
donlidx." von Trinil sind (nach .lanonsch) 304 und 303 mm
lang, auch solche

"On the other hand, remaining species [of Stegodon]

most of the von Watoealang und Ngandong erreichen 300 mm, (da sie z. T.
are considerably moi-e progressive than the form to which the mit der Hinterhiilfte noch im Kiefer stecken, kann ihre Grosse nur
lower jaw in question belongs." geschiitzt werden.) Dem gegenliber miissen die Ziihne \'on

"In my opinion we may, therefore, conclude, that the present Boemiajoe mit ca. 240 mm
genannt werden. Dabei haben
klein

Javan form is distinct from all other species. I propose the sie dicselbe Breite und dieselbe Biegung, wie die anderen Ziihne.

specific name of bondolensis, indicating the locality from whence it Die Verkiirzung kommt daher, dass die Ziihne von Boemiajoe nur
has been procured." X X
X 11 Joche besitzen gegeniiber X 13 bei der jiingeren Form.

Fig. 783. Tj'i)e left third inferior molar, I.M3, of Stegodon trigonocephalus praecursor, after von Koenigsvvald, 1933, Taf. xxvii,
fig. 2, two-tliirds natural size. From Bumiaju, Java.

Stegodon trigonocephalus praecursor Dubois (1908, pg. 1256) gibt fur das Stegodon der Kendenglagen
\()ii Koonigswald, 1933 sogar 14 Joche an, aus seinen Ausfiihrungen (Abbildungen
doch ist

hat er bisher nicht vcroffentlicht) nicht zu ersehen ob er nicht

Figure 783
etwa einen Talon mitgerochnot hat. Die Exemplare der Selenka-
Untere Schiehton, Kali Glagah, Bumiaju, Java.
Expedition haben alio, und auch die mir vorliegenden bis auf eines,

Stegodon trigonocephalus praecursor \on Koenigswald, 1933. nur 13 Joche. Dieses eine von Lepen Alit bei Tingang zeigt nur
"Beitrag zur Kenntnis der Fossilen Wirbeltiere Javas." Wetens- X 11 X Joche (vergl. v. d. Maarel, pg. 143) und gehort aber
chappelijke Mededeelingen, Dienst Mijnbouvv Nederl. -Indie, I dennoch einem typischen St. t. trigonocephalus. Dieser Fund zeigt,
Teil,No. 23, pp. 104, 105. Type.— Lower jaw with third dass noch unter den Stegodonten der Kendengschichten als
molar of both sides complete. Horizon and Locality. — Ausnahme Formen mit weniger Jochen auftreten konnen. Es ist
Untere Schichten, Kali (ilagah, Bumiaju, Java. Type cine bekannte Erscheinung, dass oft bei spezialisierteren Arten
—
Figure. Op. cit., Taf. xxvii, fig. 2 (third lower molar of the left auch noch Riickschliige auftreten."
side, I.M3). "Ein woiteres Kennzoichcn der Ziihne von Boemiajoe ist,
DESCRiPTioN.^(von Koenig.swald, 1933.1, pp. 104, lOfiJ: dass der dicke Schmelz viel weniger intensiv gefaltelt ist als bei
"Es liegt ein Unterkiefer Nor, dessen beide M3 noch komplet der typischen Art (Taf. xxvii). Das liisst auch ein Vergleich mit
sind. M3 rechts ist isoliert, ein 'i'eil der starken Zenientlage isl den Abbildungen bei Mahiin und Janensch deutlich erkennen."
abgewittert, so dass auch die letzten noch nicht angekauten Joche "Von Boemiajoe stammt audi noch ein auffallend grosser
und der kleine Talon hervortreten. Der linke M3 sitzt noch auf Stosszahn von etwa 3,50 m Liinge, der seiner ganzen Form nach
dem leider etwas gequetschten Kieferast, der im vorderen Teil an- wohl nur einem Stegodon gehort haben durfte."
 THE STEGODONTIN^: STEGODON 897

Stegodon yushensis Yovuis, 1935

Figure 78.")

l''rom Yiishe, China.

Stcgodon yuahensis Young, 1935. "Miscellaneous

Mammalian Fossils from Shansi and Ilonan." Pal.
Sinica, (C), IX, Fasc. 2, 1935, pp. 26-28. Type.—
"A presened upper left third molar."
well Hori-
zon AND Locality. "Pontian —
'violet .sands' of the
Yiishe formation from Yiishe," China. Type
—
Figure. Op. cit., PI. v, figs. 1, la.
Description.— (Young, 1935.1, pp. 26-28):
"With the exception of the fourth ridge and of a
large part of the cingulum, the tooth is well pre-
served, including the roots. Crown composed of five
ridges, not including the posterior heel and the

I'^ig. 784. Typo lower jaw witli right .second molar, r.Mo, in situ, of Parastegodon'! anterior cingulum. . . . The ridges are rather closely
kwantocnsis Tokunaga, 1934, PI. ix, fig. 1, one-half natural size. set; the vallej's not very deep; only slight traces
of cement can be observed. Cingulum is \ery weakly indicated in
the preserved parts."

Parastegodon? kwantoensis Tokunaga, 1934

Figure 784

Kakio, Kanagawa Prefecture, Japan. Upper Pliocene (fide Tokunaga).

[The description in Japanese by Professor Shigeyasu Tokunaga

of his species Paradcgodon? kwantoensiti, 1934, was not reviewed by
Professor Osborn. However, as the i)resent author regarded
Parastegodon as either a progressive Stegodon or a primitive Archi-
diskodon, his generic reference cannot be given in the present
Memoir. The following resume in English is by Mr. Ushinosuke
Narahara of the American Museum of Natural History. — Editor.]
Parastegodoni kwantoensis Tokunaga, 1934. "Fossil Elephant
teeth found at Yokohama and Kakio, Kanagawa Prefecture,"
Journ. Geog. (Tokyo), Vol. XLVI, No. 546, .July, 1934, pp.
365-369. Type. — Portion of lower jaw with right second
molar, r.Mo, in situ. Horizon and Locat.ity. — Kakio,
Kanagawa Prefecture, .Japan. Ui)iior Pliocene (fide Toku-
naga). Type Figure. — Op. cit., PI. ix, figs. 1-3.

Description. — I'ortion of a hjwer jaw with right second

molar, r.M2, in situ; eight and a half ridge-crests, probably ten if

perfect; molar valleys deep, narrow, and filled witii cement.

Length of jaw bone 253 mm.

Second molar, r.Ma
Length 190
Breadth 76
Height (outside) 35
Height (inside) 31
5,4 ridge-crests in 100 mm.

The s])ecimen is primiti\e like Parastegodon \Stegodon] aurora'

Matsumoto, but with certain differences, which he [Tokunaga]
I ^- „„. ^, , , ^ ., , r^ -, ms c <,
'
' & J
Pjg YR-y. Third .superior molar of the , i i
left side, l.M'', ol biKjoitim i/iishenMs,
believes warrants its assignment to a new species, namely, after Young, 193.5, PI. v, figs. T, la, two-thirds natural size. From Yiishe, China.
Parastegodon'! kwantoensis. (Upper) Crown view. (Lower) External view.
 . "

898 OSBORN: THE PROBOSCIDEA

"With the exception which has three roots

of the first ridge —
Material. (Hopwood, 1935.1, p. 73): "The unworn last
(extcriKil, iiifomal, under which the
median) and of the last ridge, ridgeand talon of a third molar. Probably from the upper jaw."
roots build a continuous transversal lamella, each ridge has two Cast Amer. Mus. 21879.
distinct root s. The four jjosterior ones are laterally fused along the —
Description. (Hopwood, 1935.1, j))). 73-75): "A Stegodon
external side of the footli. and the three jjosterior ones along the with the ridge-crests widely spaced, with four to five maniilla? on

internal side." each ridge; and second ridges of lower molar divided by
first

a median cleft; conules of upper molars not united into ridges."

"Dimensions: — [with some omissions) "The anterior ridge is divided into two cusps by a deep
Maximiun length of the tooth 158 mm median cleft. The post-trite cusp has two large cones with a third,
Breadth at the third ridge [maximum] 83 mm.
smaller one, between them. The tips of the cones of the pretrite
Height of the crown above the ciiigulum at the
cusps have, with one exception, been broken off. The second ridge
3rd ridge 36 mm .

is also divided by a median cleft, but this is not so prominent as in

"Both Stegodoa officinalis Hopw. and zdanskyi Hopw. are the first ridge. The pretrite cusp has three cones, and the post-
distinctly larger, and they have a higher number of ridges at the trite has two. The ridges are much narrower at the top than at
third molar. In size Stcgodon orientalis Owen (1870) and Stcgodon the base, this is especially true of the second hence the labial and
orientalis graiigeri Osborn stand closer; but they also have one lingual surfaces of the tooth slope inwards to a very marked
ridge more at -\F and the cement of their teeth is more developed. extent. The
anterior surface of the second ridge is nearly perpen-
S. sinensis Owen and S. aff. bonibifrons are described from frag- dicular, whereas the posterior surface slopes away at a fairly
ments only. The former one is hardly separable from S. orientalis: steep angle. This difference in the slope of the two surfaces gives
and the latter one is too large." the ridge the appearance of being tilted forwards, a sign of deri-
"We therefore ha\e to ileal with a new species, for \\ liich the vation from a lower tooth. There are no lateral cingula, nor
. . .

name Stegodon yiisheiisis (sp. nov.) is proposed." are there cingules at the entrances to the valleys. No cement is
pre.sent."
"The second fragment consists of the last ridge and talon of
Stegodon officinalis Iloijwood, 1935
a large tooth, which is doubtfully regarded as a third upper molar.
I'lKiircs 786, 7S7
The ridge consists of five cones which are not fused to one mass,
Said to have come fri)ni .S/.ccluiaii(?), Chiiiii. Horiz(jii unkiiowii.
but are sei)arated by definite clefts. One cleft is deeper than the
Young in his memoir "Miscellaneous Mammalian Fossils others hence the cones are divided into two groups, one with three
;

from Shansi and Honan," 1935, p. 27, compares Stcgodon officinalis cones and the other with two. A large part of the talon is covered
Hopwood (ex iNIS.) with his new species S. yiishensis. by cement, above which there rise the tips of four cones. . . The
ridges are upright, with their anterior and posterior surfaces
sloping at about the same angle; the valley between them is
narrow, and partly filled with cement."
Discussion by Hopwood. — "There
is no jjroof that these two

s])ecimens are correctly associated.At the same time it is evident

from their preservation that they are both deri\'ed from the same
deposit, which a])pears to have been of a lignitic, or peaty nature.
They indicate the existence of a species which has hitherto been
unknown and it is preferable to asstime an unproven connexion,
instead of making two 'species', one of which might have to be
relegated to the synonymy at a later period."
"In placing the sjjecies with Stegodon rather than in Stcgolo-
phodoN. Im^e had rcgarfl to the characters of the ridg(!s of the
1

I'lg. 787. "I.a.st ridge and holotype. One of the chief dilferences between the two genera is
rig. 78f>. Type of Stegodon offici- talon of an unworn upper(?) that whereas in Stegodon the cones are united into ridges, in
nalis Hopwood, coiisi.stiiig of tlio "first molar," referred by Hopwood to
Stegolophodon they remain discrete. .Vdniittedly the cones in the
two ridges of ail unworii lower molar" Stegodon officinalifi, one-lialf nat-
fragment of an upper tooth are still divided by deep clefts, but
((•;vst Amer. Mils. 21878), one-half ural size. Cast Amer. Mils.
natural size, .\fter Ho|)wood, 1935.1, 21879. After Hopwood, 193.J.I, they show the first stages of union, and the cones in each half of
PI. VII, fig. 3. I'l. VII, fig. 4. the ridg(^ arc more closely joined to each other than the two halves
of the ridges."
Stegodon officinalis Hopwood, 1935. "Fossil Proboscidea from "This species is the most primitive yet recorded from China.
China." Pal. Sinica, (f), TX, Fasc. 3, 1935, pp. 73 75 (Hopwood. In the division of the lower ridges into two cusps, as well as in the
1935.1). Typk.- "The first two ridges of an unworn lower division of the u|)por ridges and tlu; slight dis])lacement of the
molar." C'a.stAmer. Mus. 21878. Hoiuzox anb Locality. — two halves, it retains characters which derive fioin a mastodont
"Bought in a meilicinc shop, ilanchow. Said to have come from ancestry. Traces of this inheritance are never entirely lost among
Szechuan." Horizon unknown. Typk l''iGT'iiK. Op. cit., the Stegodontida^ and they persist with more or less regularity

1935.1, PI. VII. fig. 3. among the elephants. There are several species in a similar stage
 THE STEGODONTIN^: STEGODON S99

of evolutionamong the fauna of the SiwaHk deposits of India, but with one or two mammillae at the summit. Between these cones
they have not yet been described. So far as is at present known, is a long, ro(.>f-like portion with a mammillated crest. The number
the Indian species have bhinter cones than the Chinese form, and of mammillae varies with the manner of counting. If tlio main
tiieir ridges are not so high." eminences alone are reckoned, there are from four to six; if each
"Tlie trivial name chance that these specimens
refers to the tiny sub-di\ision, (jr inilication of one, is counted, this number is

were purchased in the shop of a Chinese druggist." increased."

"The crown of the tooth is curved to the rigiit, and, .seen
from the sitle, it is very slightly conca\'e. The ridges have a for-
Stegodon zdanskyi Hojiwood, 1935
ward pitch that is to say, their anterior faces are steeper than the
;

Figure 788
posterior. All these are marks of a right lower molar."
Horizon unknown.
"Anteriorly there is a relati\ely slender cingulum, which
The name Stegodon zdanskiji Ilopwood first appeared (ex MS.) receives a nodular buttress passing downwards and inwards from
in Young's memoir on "Miscellaneous Mammalian Fossils from the inner cone of the pretrite cusp of the first ridge. There are no
Shansi and Honan," 1935, pp. 27 and 28; to this species Young lateral cingules at the entrances of the valleys. A small quantity
has referred "an ujjper left maxillary with M-, M^ in situ and. an . .
of cement is present."
isolated right M''' evidently belonging to the same individual, Discussion by Hopwood. — "This species is the largest Stegodon
both found in the Pontian sands of Yiishe. A fragment of molar yet recorded. Apparently it has nothing to do with the Indian
and some milk teeth may also belong to the same species." species hitherto described; they are all smaller, have the ridges
Stegodon zdanskyi Hopwood, 1935. "Fossil Proboscidea from closer and more copious cement.
together, An undescribed
China." Pal. Sinica, (C), IX, Fasc. 3, 1935, pp. 75, 76 (Hopwood, Indian species, in which the third upper molar is 243 mm long
1935.1). Type.— "The first four ridges of a right third and 181 mm
wide, is of about the same size, but its ridges are more
lower molar." Cast Amer. ^lus. IIokizon and
21872. of the type of StegolopJwdon stegodontoides (Pilgrim), and it is
Locality. — "Bought in a Medicine Shop, Shanghai." Horizon probably referable to the same genus as that sj^ecies."
unknown. —
Typk Figure. Op. cit., 1935.1, PI. vii, fig. 5. "Apart from its size, the most interesting feature of S.
zdanskyi is the curious mixture of elephantine and mastodontine
features in its ridges. In its composition of two cusps, each made
up of two cones, and in the relations between the pretrite cusp,
anterior buttress, and the cingulum, the first ridge is, fundamental-
ly, that of any mastodont of the bunolophodont, or i)riniiti\e

zygodont, type. All the other ridges are essentially elephantine in

their structure. They each have a large cone at either end, and
a long, mammillated, roof-like portion in the centre. If the two
fissures which divide the tooth in this manner are deep enough,
the partly worn tooth would show a tripartite enamel figure.
This tripartite division is characteristic of the elephants, though it
is also shown by some, at least of the Stegodon group (cf. Soergel,

1912 [1912.2], p. 8, fig. 2).In most cases it consists of two outer

rings with an ellipse between them; exactly the type of figure
which could arise during the wearing down of a lower tooth of S.
zdanskyi."
Fig. 788. Typi' of StrgDitoii zJnnski/i Ilupunod. "First four
ridges of a right tliird lower molar" (i-ast Amor. Mas. 21872), after "All the other remains of Stegodon from < 'iiiua liillicrto de-
Hopwood, 193.5.1, PI. vii, fig. 5, one-half natural size. scribed (Owen, 1870 [1870.1]; Koken, 1885 [1885.1]; Schlosser,
1903 [1903.1]; ct alii) belong to more advanced species, and, by
Description. — (Hopwood, 1935.1, pj). 75, 76): "A Stegodon a general consensus of opinion, the various writers, other than
of very large size, with eight to ten mammillae on each ridge-crest; Owen, have referred them to such species as S. homhifrons, S.
anterior ridge still preserving the two cusps of tiie earlier mastodont ganesa, or S. insignis."
ancestors, posterior ridges foreshadowing the structure of the plates "The trivial name is given in honour of Dr. ). Zdansky of the (

found in the teeth of the P^lephantida"." Egyptian University, Cairo, who is well known for his valuable
"The first is di\ide(l into two cu.sps by
ridge of this specimen researches on the fossil mammals of China."
a prominent mesial Each cusp, esjiecially the pretrite one,
cleft.
is divided into two cones by a cleft which is not so prominent as Parastegodon [Stegodon?] sugiyamai Tokuiiaga, 1935
that which separates the cusps. The cones themselves are again Figure 78(1

divided at the summit into two or three mammillae by grooves, At Iruhi, in Saida village, Sliikoku, Japan. I'liper Pliocene or I<ower

which do not jiroceed very far down the anterior and posterior Pleistocene (/((/< Tokunaga).
surfaces. None of the other ridges show this sub-division; they I'arastegodon sugiyanuii Tokunaga, 1935. "A Aew Fossil
arc all divided, but in a very different manner." Elephant Found in Shikoku, Japan." Proc. Imp. Acad. Tokyo,
"Each of the jjosterior ridges has a large cone at either end, \o\. XI, ]). 434. Type. — An up]ier UKjlar of the left side,
900 OSBORN: THE PROBOSCIDEA

probabl}' an M'^. Horizon axd Locality. — "Mr. 'I\ t.n'yunoci'phahts Mart, from Java; and is closest to the former.
Sugiyama recently sent the author [Tokunaga] a specimen of The following is the description of the specimen."
a molar-tooth, obtaineil during road construction at Iruhi in "In the crow'n view, one side of the tooth is nearly straight
Saida village, Shikoku. The stratum from which it was obtained and the other slightly conve.x. The grinding surface is oval and
consists of an alternation of loose shale and coarse-grained quartz- slightly convex, and its base is also slightly con\ex antero-poster-
sandstone, and probably referable to either the old Pleistocene or iorly, indicating that it is an upper left molar. The anterior part of

to the youngest Pliocene." Tvpk Figuke. — Op. cit., p. 433, the crown is broken off antl there [are] I3reser\-e(l a posterior talon,
crown and inner side view of molar. seven ridges and a part of another; most prt)bably the first ridge
—
Type Description. (Op. cil., pp. 432-434): "Since medi- and the anterior talon are lost. The foremost, namely, second
aeval times numerous sjiecimens of fossil bones and teeth of ele- ridge has its anterior portion somewhat damaged and its outer
phants have been scooped up from the sea-bottom by fishermen's portion broken off, and the next three ridges also lack their outer-
nets along the eastern part of the Inland Sea of Japan. How- most portions. All the ridges are distinctly exposed, not covered
ever, very seldom are sjieciniens discovered from strata exposed on by cement. The posterior talon was co\'ered by cement and
first showed itself by shaving off the cement; it is half as high as

the adjoining ridge. The last ridge is just touched by wear and
the other ridges were worn by grinding."
"The present specimens, when complete, probably had nine
ridges, an anterior and a posterior talon."
"The length of the crown of the in-esent specimen is 120
nun +20 mm, the latter term being the estimated length of the
lost part. The width of the grinding surface is 60 at the sixth mm
ridge (67 nun at its base). Tiie height of the crest is 34 mm at
the worn eightli and 38 mm at the unworn ninth. The eighth and
ninth ridge-crests decline forwards and are a little curved in side
view, while the sc\enth but slightly declines forwards and is flat

and not curved in the same view; all the other ridges stand up-
right. In well worn stage each ridge-crest is narrowly rectangular,
not strongly constricted, and has its enamel finely plicated; in
a less ad\anced stage of wearing, it shows a large luuuber of con-
strictions. In these features the present specimen resembles in
general S. airawana; however, the detailed enamel plication of
its crown-ridges is when coini)ared with the speci-
characteristic
mens of by Soergel, Janensch, Stehlin,
the latter illustrated
Maarel and Koenigswald. The unworn last or ninth ridge of ours
has seven mammillae on crest and the eighth has seven constric-
tions. In the seventh ridge the dentine area is expanded and there
are still while in the sixth and all the others
five constrictions,
forwards there no constriction. The enamel is finely and rather
is

regularly plicated in the seventh and anterior ridges and never

form rough irregular undulations exhibited on grinding surface by
Tvi'K OK I'AliA.STHCioDO.V [STEaODON?) SOlilVAMAl
Maarel's Javan specimen, as his photograph shows, which is
V'\^. 789. 'I'j'pc molur, proliably !iti l.M", of I'arnstegoihin fnigii/amai
similar to our molar in the size of crown and grinding degree. The
TokunaKa, 193."), text figure, p. 433, i)iic-lia!f natural .sizi-. From Shikoku.
.lapau.
width of enamel measures 4 mm at the thickest portion in grinding
surface."
"In this specimen all its ridges except the posterior talon
the land surface along tliat part of the sea. The author aware of
is rise higher above the cement and the la.st or ninth is the highest."
only several namely, a few found in situ
ca.ses, at Akashi on tlu; "The projjcrly of the enamel i)li(^alion of each ridge-crest of
north shore of Ihc sea, and at Sue village, Ayauta-gun, and the author's specimen agrees with no (iescrihcd .Japanese species
Saida village. Mitoyo-gun, Kagawa-kcn, both in the scnilh of the and differs in several i)oints from the ,)a\;ui sjx'cies. Moreover
sea." it presents several features different from fossil elephants of other
"The j)resent specimen ob\iously belongs to Stcgodo/Uinnr lands. The present specimen evidently needs a new name, and
inhaving a low crown, and in other ]K)ints, but in shape, size, is named, Parodegodon xiigiyamai in honor of the discoverer,
number of ridges, and esi)ecially in the nature of its enamel Mr. Tsurukichi Sugiyama. The author is at- jjresent unable to
plication the present fossil is referable to no known s|)ecies from affirm whether it represents the first or tiie second innl.n-, hut it
Japan. It closely resembles Stegodon airdwaita Mart., and Slegodoii evitlcntly belongs to one of them."
 APPENDIX TO CHAPTER XIV
MATSUMOTO ON THE PHYLOGENY AND CLASSIFICATION OF THE JAPANESE
MASTODONTS, STEGODONTS, AND ELEPHANTS (1924, 1925, 1926, 1927)'

The numerous and diversified Proboscidea of Japan prove that this region from Miocene to early Pleistocene
times had broad land connections with southern Asia, Burma, and India, with remote relations by migration to
Europe and Africa.

Subsequent to and independently of the conclusions reached by Osborn on the classification and phylogeny
of the mastodontine and elephantine Proboscidea, Hikoshichiro Matsumoto published the systematic and
theoretic results of his important revision of the Japanese Proboscidea in six papers, as follows:

"Preliminary Note on Fossil Elephants in Japan," Journal of the Geological Society of Tokyo, Vol.
XXXI, No. 371, September 20, 1924 (1924.2).
"Preliminary Notes on the Species of Stegodon in Japan," Journal of the Geological Society of Tokyo,
Vol. XXXI, No. 373, November 20, 1924 (1924.3).

"Preliminary Notes on Two New Species of Fossil Mastodon from Japan," Journal of the Geological
Society of Tokyo, Vol. XXXI, No. 375, December 20, 1924 (1924.4).
"On Two New Mastodonts and an Archetypal Stegodont Japan," Science Reports of the Tohoku
of
Imperial University, Second Series (Geology), Vol. X, No. 1, 1926 (1926.1).
"On the Archetypal Mammoths from the Province of Kazusa," Science Reports of the Tohoku Imperial
University, Second Series (Geology), Vol. X, No. 2, 1926 (1926.2).

"On a New Fossil Race of the Asiatic Elephant in Japan," Science Reports of the Tohoku Imperial
University, Second Series (Geology), Vol. X, No. 3, 1927 (1927.1).

His results are summed up in his three phylogenetic diagrams reproduced herewith (Figs. 791, 792, 793) and
in the following systematic summary or conspectus compiled from notes included in Doctor Matsumoto's letters
of July 14 and November 20, 1924, and from his two more recent papers of 1926 and 1927. Matsumoto's valuable
letters and notes, intercalated in the English text of his published report of 1926, give us a clear statement of
his observations and opinions, as compared with those of the present Memoir indicated in square brackets.

Osborn (1927) adopts certain of Matsumoto's generic and specific terms (e.g., Palasoloxodon Mats., which
antedates Sivalikia Osb.), but rejects others (e.g., Hemimastodon Pilgrim, Prostegodon and Parastegodon Mat-
sumoto, which are antedated by genera of other authors). A full synopsis of Matsumoto's work (1924-1927)' is
given herewith for monographic purposes, followed by Osborn's comments (p. 908 below).

1. Genus Hemimastodon Pilgrim = Suina, Volume [ I, Chapter VIII]

Hemimastodon Pilgrim.
1. Hemimastodon anmdens sp. nov., Journ. Geol. Soc. Tokyo, Vol. XXXI, 1924.4, pp. 401, 405. See present

Memoir, Chapter X, p. 457. [

= Serridentinus annedens.]

2. Genus Trilophodon Falconer [=Trilophodon of the Present Memoir, Chapter VIII]

Trilophodon Falconer.
2. Trilophodon sendaicus sp. nov., Jom-n. Geol. Soc. Tokyo, Vol. XXXI, 1924.4, pp. 402, 408. See present

Memoir, Chapter VIII, p. 280. [

= Triloplwdon ><cmlairus.\

'[For a continuaUon of Matsumoto's observations and theories, np to and including 1929, see Cliai). XIX, pp. 1289 to 1300 below.— Editor.]

901
 ) ,

('Monastirtan
Census cfr. nippon
^Tyrrheniaa Loom
JBison occidenfalis
Stefociou sinensis £ cfn
Milazrian. ?fjA
5. oricntalis

< ("MitteldUuviQlschotterl E^Zephas, iast mutati-on

( = 2.. naumanni )

h Cromerian -Sicilian. E Zepha.5

1
mu-iaizon of Th/ryo -BecZ
1= ? L. hysu-driTidica )
I/) ("Altdiluviolschotter)
Z) has, F7mj.ta.t10n
Or
( PHocdnschotter)
<
CO Calabrian SfeaocZon orien^ahs
F8
Tlephas, mutation of /'^inafo
I = 'Po.reiephas trogontkerii
/ Astian
VPlaiSanc^an 3tegodoT7 orientalis
... I . -,^71 Z'umefopias, n. sp.
(=Merced -Etchec|oin) Younqer Lignites ^
or ^
^ri^ia
'''^' so.

Pontian "Taisunokuchi T- /- f ^ygoZophodon n jp. ,

(
py rertat cu-5 type )
:

(= Santa Marqarita)
Lower Lignites
CD
< ^ ^ Hanareyama ^^ T^roste^odon latidem.
CL or
Upper Saboyama
Sarmatian
< (sCierljo) Taqa or Shirado
7^ 4 J?icrocerus n.sp ,

Tbrtonian
(=Briones Ibqari or
-^•^ KadonoSQwa /TJ J)e5mo5fyiu-:- /aponicu^, fype

IYicarya
JDesmostyZus
UJ Helvetian ITurhinodelphib
< (="Tembl or)
X
10 = TsuUvyoshi 'Vicarya
F2
JDeSTTiostylus
uJ
ct: 1 Jcliocetus, n. sp.
UJ
(- Burdiqauan
z Hiramaki
o (=Vaqueros)
i: ^ »[ f-Zemimastodon, n.sp.
Artchitherium -fauna
Thirteen Fossil Mammal-beauing Formations of Japan. Matsumoto, 1921
Fig. 790. Corivlatioii Lower Mioconc HunliKnliun to tlio summit of tlic Plristoi'oiic Montistiriati of the mariiir aiui fri'shwaliT bcils of .Iai)aii with
from tlio

till' marine ami coastal .stages of western Europe (HurdiKaliaii to Mona.Mtirian) and with the epicontinental and coastal »taK<s of
the California coast of North
Amcrioa (Monterey .shale, San Pablo, Saugus-Tulare, etc.). Kejiroduecd with slight additions after pen drawing by Doctor Matsumoto (letter, .July M, 1924).
be ob-servcd that Matsumoto '.s summary of 1929 (.sec Chap. XIX, pp. 1290-1292) also Osburn's
It will summary (p. 1292) alter both the above .sj)ecific

nomenclature and the geologic levels in the Japanese Tertiaries.

902
 MATSUMOTO ON JAPANESE MASTODONTS, STEGODONTS, AND ELEPHANTS 903

Older Pleistocene

Pliocene f. filegodontoidtii
(India)

T. scm/dicit'i
I
(S. & E. Asia) (Japan) Auanct/s
[Holarclio 6f India)
Pontian
P. Idfidcns T. injitiniicnfs pats.
(S. & E. Asia) (I (i^itro-fjrr man tens
p. cautleyi Tftrolophodon
(India) (Holantic & India)

T. jiijiciKticua |)iiry, T purriiiiH *rv pni"s, ^, , .^

Sai nmtiun T ffilroneri
stibtvfuroutcns
uuslrn-gernianicu-i
(Europe}

Tortoninn T. Jfitconeri
(India)

Helvetian

T. ptjfcnaicus pari, 7*. uniju^tidtina

BurdigaMan //. crcpusrnli H. »tnnirlcii\ Zyr/otopiifi>/o,i siiblupir<nd(u.<: (Pal«arctie A India)
(India) (Japan) (Enropo) (Europe)

Hypothet irni
p[0-a""cci'""'^
Aquitaninn Stampian
Hypothetical
pTO-crcpusculi

T'h iomia
Sannoisian
('^gypt)

Fig. 791. Theoretic puylogeny of the Mastodoxtid.e (after Matsumoto, 1920.1, p. 3)

It is observed in tliis phylum that: (1) Phiomia gives rise to Trilophodon {pvo-amiectens, He mi mastodon crepusculi [=Suinal, H. l=Serridentinas] annec-
tens); that it extends into Trilophodon anguslidens of the Burdigalian and Sarmatian (of Europe), also into Tetralophodon and Ariancus (all unknown in Japan).

(2) The second i)hylum includes: Trilophodon [ = Zygolophodon] ptjrenaicus theoretically giving rise to T. falconcri, Parastegodon latidens, P. sUgodontoides, and
^tegodou] the possible derivation of Parastegodon = Stegolophodon] latidens from Trilophodon [= Zygolophodon] pyrenaicus is in accord with Schlosser and
[

Osborn.

3. Genus Parastegodon Matsumoto, 1924 [may equal Archidiskodon Pohlig, 1885, 1888, of the
Present Memoir, Chapter XVI, or a Progressive Stegodon, this Chapter]^
Parastegodon gen. nov., Journ. Geol. Soc. Tokyo, Vol. XXXI, 1924, pp. 256, 257, 262(1924.2).
— Stegodon mindanensis-Elephas aurorse group of Matsumoto.
3. Genotype: Eleyhas aurorse,
5. mindanensis-axirorse group. Low crowned, hence l:iiegodon-\i\<.i^] ridge.s close set and valleys very
narrow, very acute toward the base of crown (contrast to Stegodon, even to those with rather high
and narrow ridges of the Pleistocene). No loxodont sinus.
Parastegodon [
= Stegodon] aurorse Matsumoto, 1924, op, cit.y p. 262.
Mt. Tomuro, Province of —
Kaga horizon unknown, possibly PUocene.
Archidiskodon Pohhg, 1885, 1888, Matsumoto, 1924, pp. 256, 259 (1924.2).
= Elephas plam'frons-meridionalis group. No known Japanese representative.
HSee Matsumoto, 1929, "On Parastegodon Matsumoto and its bearing on the Descent of Earlier Elephants," Sci. Rept. Tokoku Imp. Univ., (2), Geology,
XIII, No. 1, pp. 13-15 (in English).— Editor.]
904 OSBORN: THE PROBOSCIDEA

(^^U^f'tM^'^^T''^ 0'Cec<:r«''-o->-i^

"lXi/l^t**'*^^»-'<»'*^

nJU-}p^^^

[^Aytry^^-t^^

C/iUceJi'-'U.-'^.^i^

lAlWU^ fo-VA-LLe^^v-V^

4
L•^,>tA J(yv'-AA.!«,.w^

Fig. 792. Theoretic phylogeny of the Stegodonts (after M.^tsumoto, MS. of November 20, 1924)

Observe that Vrmlrgodon ineludcs five phyla, namely: (1) The phylum of latidens and slegodonloidcs [ = the Slegolophodon of Schle.'iinger]; that SUgudon
iiiclii(k'.s (2) the pliylum of ch/a' and smcnsis, (3) tlic phylum of 6owiJ^!/rons, orte/i/afo, and orienlalis sliodocnsis, (4) the phylum of bombifrons, insignis, and

ganesa, and (5) the i)hylum of airaivana and trigonncephalus. Thi.s .subdivision of tlie species of Stegodonts into several phyla, grouped under two genera,
namely, (a) I'roslcgodoti [= Skgolophodun] and (b) Stcgodoii, with their numerous branches, agrees substantially with the conclusions reached by Osborn.
This diagram is reproduced in Matsnmoto (1924.3, p. 339), with omission of text outside of black lines and with extension of diagonal line from boinhifmns
(India) into the difli (India) cohunn.

4. Genus Prostegodon Matsumoto, 1923, 1924 [

= Stegolophodon Schlesinger, 1917, of the Present
Memoir, this Chapter]
Prostegodon gen. nov., Matsumoto in Osborn, Amer. Mus. Novitates, No. 99, 1923, p. 2 (Osborn, 1923.601);
also Matsumoto, Journ. Geol. Soc. Tokyo, Vol. XXXI, p. 325 (1924.3).
4. Genotyi)o: Madodon latidens Clift. [This genus {Prostegodo7i = Stegolophodon^) includes also Mastodon
cautleyi Lydekker and M . stegodontoides Pilgrim, etc.]

Matsumoto, 1926.1, p. 9.
— "Skull and mandible only, imperfectly known, brevirostral. Lower incisur-tusks

might be absent, or al>ortiv(> if present at all. Intermediate molars four- or five-ridged, last molars five- or six-

ridged. Grinders essentially lophodont, though their first and second ridges may show a slight tendency of buno-
donty and of trefoil pattern of cusps; mesial longitudinal cleft evident; inner and outer cusps opposite, instead
of being alternate; valleys widely open, free of cement. ... As clearly pointed out by Lydekker and by Pilgrim,
the grouj) which now forms the present genus is very closely allied with the coiniiioii Stegodonts, tliough more
archetypal th;in the .same. Notwithstanding that, it is perfectly distinct phyletically from the other two phyla of

'[See page 839.— Editor.]

J
 MATSUMOTO ON JAPANESE MASTODONTS, STEflODONTS, AND ELEPHANTS 90.'^

iCiK*L^-ticrftZi»^^ ^,lc^
(jaM^fiamicrn 6 c-c-c-oe^/SJ-jtA^tl- (7t>ljtAcolu-(,e<^fyt^ fc3(^^><W ,H.
;/,

(JtLLctf<!ir>C(fii'Cn'i^ Z<?*»<W^ i

H.^e ^*^~<JU.*^u^
*s^i f
. w.i<u 3

]J*T-M*>lA.>^Vi

l^rVWX. *CoA<A. vx

fi^ 14.

^_^l^'>«t*-AA,*i->i

(?'r^)

CftXoJ***^*^^
Cfc-i^UiT-*.)

(A4*''>jCi^%JL*)

(oo*AX-iK.tA^

Fig. 793. Theoretic phylogeny ok the Elephantid* of A.sia and Eorope (afteb Matsumoto, MS. of November 20, 1924)

Oliscivc that: and mindanensis; (2) Euelephas includes the " Irogonlherii" phylum [=l'arfbphas
(1) /'n;a.?<cffO(ton ineUides "insigni>i" (pars), aurora,
of Osfxiiiil and tho "jjrimigenius" phylum = .Ua/Hwo«<<'((.s of Osborn], both derived from the specie.s prntnmammonleus of Japan; (3) the Arcliiilinkodon
[

phylum incliide.s the .speoies planifroiis, cnmnnnHis, meridional is, and hysudricus; (4) the Elephas phylum includes liy.sudriruHcus, nainadicua (in pari), and the
true indicus of Japan, China, and India; (.")) the I'alxohxodon i)hylum includes three divisions (a) typical Palxoloxodon namadicus and imunmnni [equivalent
to theSuaiifcia of Osborn], (b) the anhguus phylum of Europe, with ausonius and anliquus, (c) the narrow-toothed priscus, iokunagai, and atUinticus = Pil- [

grimia of Osborn, made by Osborn in 1934 a synonym of Palxoloxodon Matsumoto], and (6) Loxodonla including africanus.

Tetralophodonts, viz., Tetralophodon (longirostral) and Anancus { = Dibxmodon), being distinguished from the
former by being brevirostral, by the more distinct lophodonty, and by the single and feeble, but not doul)Ie and
prominent, trefoil pattern of cusps of grinders, if distinct at all, and from the latter by the more distinct lopho-

donty, by the simpler and feebler trefoil pattern of cusps and by the opposite, but not alternate, halves of ridges
of grinders. Further, it is distinguished from the common Stegodonts by the lower ridge-formula, by the rather
less distinct lophodonty, by the ridges of grinders consisting of smaller number of mammillae and by the entire
absence of cement."

Prostegodon latidens Clift. Mt. Hanare, Kuji District, Province of Hitachi; Kitayama, northern precinct
of Sendai. (Letter of Nov. 20, 1924.) Lower Pontian.

Prostegodon latidens Clift. Shiwogama, Miyagi District, Province of Rikuzen. Horizon: a bed of coarse-
grained sandy tuff, probably to be referred to the Sawoyama Formation proper (Matsumoto, 1926.1, p.

10). Probably of typical Pontian age.

Note: La connection with a long description of this latter specimen, measurements are given of the tooth,
a Dp* or an M\
 —

90f) OSHORN: THE rKDHOSCIDl'JA

5. Genus Stegodon Falconer and Cautley = Stegodon of the Present Memoir,

[ this Chapter]

Siegodon Falconer and Cautley.

5. Stegodon clifti Falconer and Cautley, Journ. Geo). 8oc. Tokyo, Vol. XXXI, 1924, p. 329 (1924.3).
Akira-mura, Kage District, Province of Ise — possibly Plaisancian-Astian.
6. Stegodon sinensis Owen, op. cit., 1924, p. 328 (1924.3).
Island of Shodo (Shodo-shima or Shozu-shima), Inland Sea —Milazzian-Tyrrhenian.
7. Stegodon orientalis Owen (typicus), op. cit., 1924, p. 330 (1924.3). (Also as S. bombifrons, op. cit., p. 329.)
Nagahama, Minato Town, Kimitsu District, Province of Kazusa; Euelephas protomammonteus
zone, i. e., Calabrian. —
Togane Town, same province Calabrian. Ikadachi-mura (formerly Riuge-
mura). Province of Onii: wvAy possibly be Calabrian or Cromerian. (This locality has not yet yielded
elephant remains. The specimen of 'E. namadicus' mistakably said to have come from this locality,

is realh' from Yokosuka.) Okimisono, Uhe coal-field, Province of Suwo— probably Cromerian (Sicilian).

8. Stegodon orientalis shodoensis subsp. nov., op. cit., 1924, p. 333 (1924.3).
Islands of Alitsugo (Alitsugo-shima) and Island of Shodo, Inland Sea; off Nagasaki, Eastern Sea
Milazzian-Tyrrhenian. Kashiwazaki (town), Province of Echigo (?this form).

9. Stegodon bombifrons Falconer and Cautley, 1924.3, p. 336.

Higashi-Kanamachi (town). Province of Kazusa. Upper part of Plaisancian-Astian.

(). Gknus Euelephas Falconer = Parelephas Osborn (Chap. XVII), Mammonteus Osborn (Chap.
[

XVIII), AND Pal.eoloxodon Matsumoto (Chap. XIX), in part, of the Present Memoir]

Euelephas Falconer.
= E. trogontherii-primigenius group. Matsumoto, 1924, pp. 256, 258 (1924.2).
The Upper Pliocene form of this phylum is ai/rora^-like in the shape of the basal parts of ridges and
valleys. No loxodont sinus.
10. EneJephas protomammonteus, sp.n., op. cit., p. 262, 265.

Nagahama, Minato Town, Kimitsu District, Province of Kazusa (type loc.) ; Seki-mura, same
district; Uehata, Akimoto-mura, same district; Kokubo, Onuki-mura, same district; Otomi, Mat-
suoka-mura, same district.
Horizon: Sanuki Bed, which is the lowest one of the gravel bed series (Narita Series, s. ext.) of

Calabrian-Pleistocene.
Smaller and more archetypal than E. irogontherii.
Newly recognized in the Japanese Upper Pliocene fauna. I mean the oldest Elephas from Minato.
Molars .small; crown low. Lower molars E. trogontherii-\\kc, while the upper molars are E. primigenius-
like. I became convinced, after examining a number of upper molars, that it is a mnmmotli but not
a Loxodont.
Parelephas protomammonteus (Matsumoto) typicus, 1926.2, p. 43, PI. xviii, figs. 1-4 (holotype); PI.

XIX, figs. 1-3 (paratype); Pis. xx-xxiii, figs. 1, 2 (referred).

Nagahama, Town of Minato, Kimitsu District, Province of Kazusa (locality of tyj^e and paratype
and one referred .specimen); Seki-mura, same district; Uehata, Akimoto-miua, same district; San-
kawa Valley, Hosono, Matsuoka-mura, same district (referred specimens).
 .

MATSUMOTO ON JAPANESE MASTODONTS, STEC.ODONTS, AND ELEPHANTS <JU7

10a. Parelephns protonmiamonteioi proximus, mut. nov., 1926, p. 48, PI. xxiv, figs. 1, 2 (1926.2).
Isone, Kokubo, Onuki-mura, Kimitsu District, Province of Kazusa. Horizon: Prol^ahly "from
a certain lower part of the Narita Series." Age: May belong to Lower Calabrian.

11. Euelephas trogontherii (PohUg), 1924.2, p. 265.

Lake Kasumiga-ura, Province of Hitachi; Hishiike, Hadsu District, Province of Mikawa (lost
specimen).
Horizon: Loxodonta {Palxoloxodon) namadicus naumanni zone, which corresponds to the Tokyo
Bed, as well as the middle bed of the Narita Series, s. ext., just mentioned.
Age: Cromerian ( = SiciUan).
12. Euelephas primigenius typicus. Japan, Siberia, Eastern Mongoha (1924.2).
The precise locahty of the Japanese specimen, which I examined, from the collection of the Kyoto
Imperial University, subsequently \\'ill be determined by one of the professors of tliat University.
{Euelephas primigenius sibiricus). Siberia, Eastern Mongolia; but not yet in Japan.

7. Genus Elephas Linn^us = Elephas of the Present Memoir, Chapter XX]

[

Elephas Linne, s. s. (1924.2, p. 256).

= E. hysudrindicus-indicus group.
Characterized by the total absence of loxodont sinus, and very fine, regular, and deep plication of
enamel.

13. Elephas indicus Linne, op. cit., 1924, p. 266 (1924.2).

Between Tokyo and Kanagawa (Leith Adams); Yedobashi, Tokyo; Ninohe District, Province of
Mutsu; Sapporo, Hokkaido; Province of Mino; Prefecture of Wakayama; besides several specimens
from unknown localities — probabl}' Post-Monastirian and Pre-Neolithie. This species lived also in
China.
This true Elephas phylum was doubtless present in Japan, though so long confused with 'E. namadi-
cus.^ Naumann's Yedobashi specimen; Busk-Lydekker's Kanagawa specimen; Province of
E.g.,

Mino specimen and Prof. Tokunaga's photograph at your hand (this specimen was lost by the great
disaster of last year of Japan). All the Japanese specimens are not at all or scarcely fossilized, occurring

from loam-soil or surface deposits. May possibly be of Post-Monastirian warm period.

13a. Elephas indicus Buski subsp. nov., 1927, p. 57, Pis. xxvii, figs. 2 and 3 (type); PI. xxviii, figs. 1, 2
(referred)

Ninohe District, Province of Mutsu (type locality) ;

precise locality of referred specimen unknown.
Age: May belong to a very late geological age, such as the Post-Monastirian.

8. Genus Loxodonta Cuvier = Pal^oloxodon Matsumoto of the Present Memoir, Chapter XIX]
[

Loxodonta Cuv. s. ext., 1924, pp. 257, 260 (1924.2).

Subgenus Palxoloxodon, subg. n., op. cit., pp. 257, 260 [antedates Sivalikia Osborn].
= E. anliquus-namadicus group.
Genotype: E. namadicus naumanni Makiyama. Almost similar to Cromerian stage of E. antiqiins.

Tokyo Bed, i.e., middle bed of Upper Musashino. Probably Cromerian.

908 OSBOIW: THE PROBOSCIDEA

Phylum A. K. melitensis-atlanticus phylum.

14. Loxodonta {Palseoloxodon) tokunagai, sp. n., 1924, p. 267 (1924.2).
Hira-mura, Higashi-Tonami District, Province of Etehu (type loc).
Horizon and age unknown, may be either Calabrian or older Pleistocene.

Phylum B. E. aniiqmis-namadicus phylum.

15. Loxodonta {Palseoloxodon) namadicus naumanni (Makiyama), 1924, p. 264 (1924.2).

Yokosuka, Province of Sagami (Naumann); Tabata, Tokyo (Tokunaga); Sahamma, Province
of Totomi (Makiyama); Kasumiga-ura (lake). Province of Hitachi; Imba-numa (lake). Province of
Shimosa; Hishiike, Hadzu District, Province of Mikawa; Nakao, Kiyokawa-mura, Kimitsu District,
Province of Kazusa; Yamawaki, Tamaki-mura, same district Miyata, Miura Penninsura, Province of
;

Sagami.
Horizon: Tokyo Bed, as well as middle bed of the Narita Series, s. ext., above mentioned.
Age: Cromerian ( = Sicilian).

16. Loxodonta {Palseoloxodon) namadicus {typicus) Falconer and Cautley, 1924, p. 269 (1924.2).
'E. namadicus,' younger type. Shozu-shima; and also younger terrace deposits all over Japan
including Hokkaido. Probably Milazzian, or also ?Tyrrhenian. Inland Sea; Tsukiyoshi, Minami-
Akita District, Province of Ugo (Matsumoto) ; Sagawa Town, Aki District, Province of Tosa. Hanno-
sura. Province of Noto (Matsumoto) ; Sorachi, Province of Ishikari, Hokkaido (Matsumoto) ; Itsukairhi,
Province of Musashi.

This form in Japan consists of two types, the larger-molared type corresponds well to the Narbada
type, while the smaller-molarerl type is commonest in Japan. Age: Milazzian-Tyrrhenian.

Few-ridged Palsearctic Loxodonta [priscus-melitensis group].

E. pmcMs-like form. M3, ca. X13X; crown low; enamel thick. Hira-mura, Province of Etchii
(horizon unknown) ; Shozu-shima, probably Milazzian.

Subgenus Loxodonta, s.s., op. cit., 1924, pp. 257, 261 (1924.2).

= E. africanus group. No known Japanese representative.

OSBORN COMMENTS (1929) ON MATSUMOTO'S PHYLOGENY AND

CLASSIFICATION (1924 1927)

The very important but debatable result of Matsumoto's researches and observations above cited is the
presence in Japan of the mastodontoid genus Trilophodon and of representatives of four' subfamilies, namely, the
Stegodontinso, Manunontina^, LoxodontiniE, and Elei)hantiniiB. Without monographic examination and com-
l)arison of these original materials and types of Asiatic and European species, it is difficult to form coi-reot

judgments as to the true phyletic and generic relationships of many of these species and subspecies.
'[Actually five subfamilies, as Professor Osborn described a new subfamily, the Stegolophodontin*, type Slegolophodon, in Volume 1 of the pip.seut
Memoir.— Editor.]
 .

MATSUMOTO ON JAPANESE MASTODONTS, STEGODONTS, AND ELEPHANTS 909

rKOBOSCIDlOA DlSCOVEUIOt) IN .IaI'AN, BuuMA, JnDIA, AND ('lIlNA

References by Matsumoto (1924-1927) References by Osborn in the Japan

Present Memoir
Genus Hemimaslodon Pilgrim = Suina
1. H . annectens Matsumoto = Serridentinus anneclens Matsumoto
Genus Trilophodon Falconer = Trilophodon Falconer
2. T. sendaicus Matsumoto = Trilophodon sendaicus Matsumoto
Genus Parastegodon Matsumoto -- Archidiskodon Pohlig or progressive Slego-
don
3. Genotypic species Parastegodon aurorx Matsumoto --
Slegodon aurorse Matsumoto
Genus Proslegodon Matsumoto = Stegolophodon Schlesinger
4. Genotypic species M. latidens Clift = Stegolophodon latidens Clift
Vjdxms Stegodon Falconer and Cautley = Slegodon Falconer and Cautley
5. Slegodon cliftii F. and C. = Slegodon elephanloides ( = cliflii) Falconer
and Cautley
6. Slegodon sinensis Owen Slegodon sinensis Owen
7. Slegodon orienUdis Owen -Slegodon orienlalis Owen
8. Slegodon orienlalis shodoensis Matsumoto = Slegodon orienlalis shodoensis Matsumoto

9. Slegodon bombifrons Falconer and Cautley = Slegodon bombifrons Falconer and Cautley

Genus Euelephas Falconer Parelephas Osborn (in part), Mammon-

teus Osborn (in part), Palxoloxodon
Matsumoto (in part)
10. Euelephas (Parelephas) protomammonteus Matsumoto = Palseoloxodon protomammonteus Matsu-
moto
10a. Parelephas protomammonleus proximus Matsumoto = Palxoloxodon protomammonteus proximus
Matsumoto
11 Euelephas Irogontherii Pohlig
12. Euelephas primigenius = Mammonleus primigenius Blumenbach
Genus Elephas Linnseus = Elephas Linnseus
13. Elephas indicus Linnseus = Elephas indicus Linnseus
13a. Elephas indicus Buski Matsumoto = Elephas [Palxoloxodonf] Buski
Matsumoto
< ienus LoxodonUi (
'uvier = I^oxodonta Cuvier
Subgenus Palaoloxodon ^Matsumoto = Palaoloxodon Matsumoto; Sivalikia
Osborn, a synonym
14. Lo.rodonta {Palxoloxodon) tokunagai Matsumoto = Palseoloxodon tokunagai Matsumoto
15. Loxodonla (Palaeoloxodon) namndica nainnanni
Makiyama = Palseoloxodon namadicus naumanni
Makiyama
16. Loxodonla (Pahroloxodon) namadica Falconer and
Cautley = Palseoloxodon namadicus Falconer
and Cautley
 ^

Blepliantid;c. After Falconer and Cautley,

Fig. 794. Primitive, intermofliato, and i)roKressive mandibles and grinding teeth of the
1846 [1847, PI. xiii.AJ. One-.sixth natural size.
ridge formula an,l elongate rostrum. =Lo.xodontin.«
Fig. 8. E. africanus[ = Loxodonla africana]. Least progressive, platycephalic mandible, with iirimitivo
Fig. 6. Elephas indicus. Slightly more progressive mandible, with somewhat more reduced ro.strum.
- Ei-ErH.*.NTiN.E
more abbreviated mandible, with reduced rostrum. =Eleph.\ntin*
Fig. 7. Elep}ms hijsudricus l=llypselephas hjsudricus]. Still
ro.strum. -Mammontin^
Fig. 3. Elephas primigenius [probably Parekphas trogontherii]. Very robust mandible, with vestigial
= Relatively elongate mandible, with reduced ro.strum. .\ppoars as E. mcndiowdis
Figs. 4,5. Elephas antiquus[ Hesperoloxodon anliquus].
- Loxodontin.b
on plate; changed by Falconer on original plate in British Museum.
molars in sUu. greatly reduced rostrum. =Mammontin«
Fig. 2. Ekphas primigenius = Mammonteus primigcniux]. Juvenile individual, with second
I
inferior

Fig 1. Elephas primigenius [^Mammonteus primigenius]. Fully adult mandible, with broadly arched ro.strum. Third mferior grmders
progressive, hypsicephalic, brachycephalic, and bathyccpl.alic type of mandible.
=Mammontin.b
well worn, indicating advanced age. The most
910
 ,

Chapter XV
CLASSIFICATION OF THE ELEPHANTOIDEA BY THEIR DIVERGENT AND
HARMONIC CRANIAL AND DENTAL CHARACTERS
Application of principles of phylogenetic classification. Distinction of parallel and convergent
FROM divergent AND ADAPTIVELY RADIATING CHARACTERS OBSERVABLE IN CRANIAL AND DENTAL MECHANISMS.
Recapitulation of ancestral characters in course of development. Correlation of brachycephaly and
HYPSICEPHALY WITH FEEDING HABITS AND HABITAT, WITH BRACHYODONTY AND HYPSODONTY. ThREE SUBFAMILY
i'Hyla: mammontin.^, loxodontin.e, and elephantin.e.

I. Introductory Section. 3. Seasonal changes in food of the mammoth.

1. Principles and methods of phylogenetic classification 4. Summary of progression from browsing to grazing
applied to the Elephantida?. dentition.
Failure of previous dental classifications.
L lassihcation by cranial and dental characters.
m Vertebral Distinctions of Elephas, Loxodonta, Mam-
monteus and Parelephas
Cranial mechanics of Elephas (Weithofer,
1, Vertebral formula; in the above genera.
^
Osborn, Ciregory).
Comparative cranial sections of elephant skulls. IV. Synopsis of Subfamily Classification of the Elephant-
Ontogenetic cranial changes in Elephns indicus. Oidea.
Generic contrasts in cranial sections: Loxodonta, 1. Subfamily Mammontinse, originally browsing, pro-
Parelephas, Mammonteus, Elephas, and Archi- gressive to extreme grazing type. Genera:
diskodon. Archidiskodon, Parelephas, Mammonleus.
^- ^"bfamily Loxodontinae, feeding habits chiefly
II. Dental and Cranial Adaptation to Prevailing Feeding
bi-owsing. Genera Loxodonta, PaLroloxodon

1.
...primitive
Habits the Key to Phylogenetic Classification.
,_,.,,.
Kidge-plate formulae ot
,

and progressive o r, i
'''"M^'^™
c
Hesperoloxodon.
•, ^i i ,
•
:

i a u ,

genera in adaptation to prevailing h.t^^its of

^- ^ ^^^'^''^l'^' '^'^^1^ browsing, second-
'^''
•^ grazing. Genera: Elephas, Hypselephas,
feedine:
J^lciiclcDhas
2. Food of the Indian and African elephants and of the
mammoth. V. Final Summary of Chapter.

I. INTRODUCTORY SECTION
In the years 1902-1903 the author, with the aid of his colleague William King Gregory, devoted many month.s
of research to the ontogenetic development of the elephantine cranium, also to the method of analyzing the
actual forms and proportions of the elephantine cranium by means of sections made with bent wire. Only by such
means can the cranial proportions be effectively studied and illustrated. Gregory pubhshed part of the results
of these ontogenetic studies in his paper of 1903, entitled, "Adaptive Significance of the Shortening of the Ele-
phant's Skull," as quoted below. The complete results obtained by the wire sectional method, as illustrated in
figures 801, 803 to 814, are here pubhshed for the first time.

1. PRINCIPLES AND METHODS OF PHYLOGENETIC CLASSIFICATION APPLIED TO THE

ELEPHANTID^
The secret of phylogenetic classification in the family ElephantidiB is to discover characters in which the
different branches diverge from each other. Hitherto students of phylogeny have been deceived by the more
numerous characters in which they converge, in other words, by their parallelism. In the present chapter it is

shown that whereas all the Elephantidae converge in their proboscis, in their tusks, and in certain progression in
their grinding teeth (e.g., addition of ridge-plates), they also diverge in the profound proportional changes in the
cranium, with which natiu-ally the tusks and the grinding teeth are harmonic. While outwardly similar, the crania
in different subfamilies of elephants are inirardly profoundly divergent. These divergences are in continuous adapta-
tion to the prevailing habitat and feeding habits.
911
 — .

912 OSRORN: THE PROBOSCIDEA

Step by step it has been observed by Osborn, in comparing the crania of the Hving and fossil elephants, that
all the dental and cranial characters are harmonic, that there is a close adaptive correlation between the form and
position of the grinding teeth and of the component parts of the cranium. Moreover, the lines of descent, which
are apparently dissimilar in superficial dental structure, are really profoundly related both in dental and cranial
harmony. From these comparative observations, which will now be described in detail, has arisen the present
phylogentic classification of the Elephantidse. After a review of the cranial and what is known of the vertebral

characters, the three subfamilies and included genera will be fully defined in Section IV of the present chapter
(cf. p. 932 below).

Suborder or Superfamily : ELEPHANTOIDEA Osborn, 1921

(See Volume I of the present Memoir, pp. 22-33, figs. 3, 4, 5, 7, 8, and PI. xi)

Original reference: "The Evolution, Phylogeny and Classification of the Proboscidea" (Osborn, 1921.515, pp. 2 and 4).

[The present chapter was written before Professor Osborn separated the Stegodontoidea from the Elephant-
oidea, as fully set forth in the introductory pages of the preceding chapter. Consequently in the following citations

the reader should disregard the inclusion of the Stegodonts in the Elephantoidea, retaining only the subfamilies
Mammontinae (Archidiskodon, Parelephas, Mammonteus), Loxodontinae {Loxodonta, Palxoloxodon, Hesperoloxo-
don), and Elephantinae (Elephas, Hypselephas, Platelephas)

It was in January, 1921, in his article on "The Evolution, Phylogeny and Classification of the Proboscidea"
that Professor Osborn first published the term Elephantoidea (Osborn, 1921.515, pp. 2 and 4 respectively):

Elephantoidea to include the Elephantinse, Loxodontinae, Stegodontinse, and Mammontinae.

IV.
. [p. 4]
. One prime distinction in this superfamily is the very early complete loss of the lower incisor
teeth, accompanied by the early development of the upper incisors into horizontal or upturned tusks
finally devoid of enamel except at the tips in the young stage. Vestigial enamel bands are recorded in
early stages of the stegodonts. A
second distinctive character is the absence of conule tlevelopment
into trefoils, so characteristic of the mastodontoids, and the early tendency to form evenly transverse,
more or less mammillate, crests which become in the highest degree hypsodont and polylophodont in
adaptation to chiefiy grazing habits.

Subsequently on sectioning certain of the molars of the Stegodontoidea and the Elephantoidea it was found
that the valleys separating the adjacent ridges were V-shaped in the former and U-shaped in the latter.

Family: ELEPHANTID^ Gray, 1821

Original reference: Gray, "On the Natural Arrangement of Vertebrose Animals," London Medical Repository, 1821, XV, No.
88, p. 305.
Syn. : Elepluisideie Lesson, 1842, p. 156; Elephantida; Bonaparte, 1838, p. 112, and 1850; Girard, 1852, p. 326; Zittel, 1891, p. 458.

While Gray in his classification of the Proboscidea (op. cit., p. 305) assigned the terms Elephantidse and
Mastodonadae respectively to the elephants and mastodonts, Girard (1852, p. 326) was the first to use the form
Mastodontidae.

Family Defi.nition (Gray, 1821, p. 305). -PROBOSCIDLE Fam. 1. ELEPIIANTIDyE.—

Teeth, two grinders in each jaw, composed of transverse vertical lamina>, enveloped in enamel, and
soldered together ))y a cortical substance.

Elephant, Elephas Lin. E. Indicus Cuv.

Osborn (1910.346, p. 558): Fam. Elephantidse. Dinotheres, Mastodons, and Elephants.
 CLASSIFICATION OF THE ELEPHANTOIDEA 913

Osborn (1925.662, p. 28): Family: ELEPHANTID/E, distinguished by plated grinding teeth

developing out of the more or less closely compressed, serrated ridges of Stegodon into the broadly plated
grinders of Arcfiidiskodon, the lozenge-shaped grinders of Loxodonta, and the compressed, finely plated
grinders of Parelephas, of Mammonteus, and of Elephas the type genus of the family.

It will be observed, however, by referring to page 25 of Volume I (1936) of the present Memoir, that Professor
Osborn altered his opinion as to the development of the grinding teeth of the Elephantidse (p. 942 below) "out
of the more or less closely compressed, serrated ridges of Stegodon": "It has been assumed by practically all

palseontologists that the Elephants were descended from the Stegodonts. This assumption now proves to be
erroneous, for neither tlie Stegodon grinding tooth with enamel valleys closed at the bottom, nor the Stegodon
cranium with its extremely short face, can give rise to the elephantoid molar or the face of the elephantoid
cranium." —Editor.]
Subfamily: Mammontin^ Osborn, 1921
The Mammontinse include the southern mammoth {Archidiskodon) , the north temperate mammoth {Parele-
phas), the true northern mammoth {Mammonteus)}

Genus Archidiskodon = the Southern Mammoth,

inchitling Archidiskodon proplanifrons, siibplanifrons, planifrons, A. meridionalis, A. imperator, and other
species, chiefly ranging in the south temperate zone.

Genus Parelephas =the North Temperate Mammoth,

including the species Parelephas trogontherii, P. jeffcrsonii, P. trogontherioidcs, P. columbi, usually inter-
mediate in geographic range and climatic life zone between the southern mammoth and the northern
mammoth.
Genus Mammonteus^ =the Northern or Woolly Mammoth,
typified by Mammonteus primigenius, including also M.
primigenius americanus, M. primigenius alasken-
sis, M. primigenius compressus, and other subspecies of the northern steppes and tundras.

Subfamily: Loxodontin.e Osborn, 1918

The Loxodontinae include the three genera Loxodonta, Palseoloxodon, and Hesperoloxodon.
Genus Loxodonta
including Loxodonta africana, and probably L. zulu, L. prima, L. africana var. obliqua, L. subanliqua, of Africa.
Genus Palxoloxodon
including especially Palxoloxodon namadicus of India; P. melitensis, P. mnaidriensis, P. falconer i, P.
Cypriotes,and P. creticus of the Mediterranean Islands, also the Palaeoloxodonts of Africa, of .Jajjan, and
of Java.

Genus Hesperoloxodon
including Hesperoloxodon antiquus, H. antiquus italicus, H. antiquus germanicus, etc. of Europe.

Subfamily Elephantin^e: Osborn, 1910

The Elephantinae include the typical or true elephants of India, consisting of the three genera Elephas,
Hypselephas, and Platelephas.

Genus Elephas
including the typical Elephas indicus and the geographical varieties of E. indicus bengalensis and E. indicus
ceylanicus, also E. sumatranus.
Genus Hypselephas
including the extinct Pleistocene species Elephas [Hi/psekphas] hysudricus of India.
Genus PlutcUphas
including Platelephas platycephalus of the Upper Phocene or Lower Pleistocene of India.

HThe validity of the generic term Mammonteus is doubtful— see Chapter XXI, pp. 1365-1367, below, on the nomenclature of the Proboscidea.— Editor.
914 OSBORN: THE PROBOSCIDEA

Fig. 79.5. Gener.\l Climatic Distribution op the Subfamilies of the Elephantoidea and Stegodontoidea, Including
Theoretic Migration Lines (1938)
noutii polak pliojection prepared by the american geographical society in 1924
1) The Stegodonts (Stegofhu): Soutlieni and E.astern A.sia, .Jiipaii, and the East Indies.
2) The Southern Mammoths (Archiriiskodon): Africa, Southern Europe, Eughmd, Southern A.sia, the United States, and Mexico.
3) The Northern Mammoth.s (^fammonl(us): Circumpohir distribution extending southward to the 40th parallel in late Pleistocene times.
4) Troooxtherian Mammoths (I'ardcphas): Southern and Eastern Europe, the United States, Mexico, and South America; elsewhere theoretic (?)
migration lines.

o) The Loxodontine Elephants {Loxodonla. PaUcaloxodun, JIcKpcroluxodon): Southern Europe, Asia, and the continent of Africa. See figure 1242 for
living African elephants.
6) Indian Elephants {EUiihus, Ili/jiseh ijlma, T'lntdc jihax) , recent and fossil: Southern Asia and the East Indies. See also figure 1242.

Failure of Puevious Dental Classifications

Falccjiirr's classification (1857, p. 318, and 1868, Vol. I, p. 82 et seq.) into three subgenera was based chiefly
on the grinding teeth: Subgcn. i. Stegodon. Hiibgen. 2. Loxodon. Subgcii. 3. Euelephas. Pohlig's classification
(1888, p. 138) was also based solely on the grinding teeth: Polydiskodon, Archidiskodon, and Loxo{disko)don.
We find that clas.sification of the Elephantidse by dental characters alone cannot be i)hylogenctic.

Classification by Cranial and Dental Characters

The phylogenetic classification in this Memoir is based on the evolution of the craiiiuin and jaws, harmonic
with the grinding teeth, especially with the thirtl superior and inferior molai's, AP-I\l.i, w iiich pro\es to afford an
absolute means of distinguishing generic phyla from each other.
 : ;

CLASSIFICATION OF THE ELEPHANTOIDEA 915

CuviER.— From the first by Cuvier of the African from the Indian elephant and from Elephas
distinction
primigenius,^ all authors have given more or less attention to the external characters of the crania of the Ele-
phantidaj, although they have not made the cranium the chief basis of classification.

Cranial Mechanics of Elephas (Weithofer, Osborn, Gregory)

Weithofer. — In 1890 Weithofer made a detailed comparative and functional study of the elephant's
skull, and concluded that the ontogenetic mechanical changes in the skull, observable during individual growth,
as affecting phylogeny, such as the fore-and-aft compression and vertical heightening and deepening of the skull,
the wide separation of the inner and outer tabulae of the bones and the cancellous condition of the diploe, the
forward shifting of the orbits from a point above the anterior grinders, etc., are primarily correlated with the
prodigious development of the tusks —weapons and crowbars whose effectiveness increased with and reciprocally
hastened the phyletic advance in body dimensions. Gregory (1903.1) did not accept Weithofer's conclusion that
the tusks were the chief cause of fore-and-aft compression, but showed that the proboscis is another and perhaps
the chief factor in the extraordinary process of fore-and-aft cranial compression, or cyrtocephaly.

Osborn-Gregory. —Studies of the elephant cranium as a whole were undertaken by Osborn (1902-1903)
with the cooperation of William K. Gregory. Gregory (1903) in his "Adaptive Significance of the Shortening of
the Elephant's Skull" attributed the fore-and-aft compression principle to the enlargement and backward shifting
both of the proboscis and of the tusks. His principal conclusions and explanatory figures may be freely summarized,
without quotations, as follows: (1) First factor. Lengthening of the proboscis correlated with shortening of
cranium and of neck pari passu with lengthening of limbs and increased height; also with downward shifting and
elongation of tusks (vide Weithofer). (2) Second factor. Backward shifting of weighty tusks and of proboscis

diminished the anteroposterior space (i.e., brachycephaly) for grinding teeth (Dp 2-M 3). (3) Third factor.
Rapidly heightening grinding teeth (i.e., hypsodonty), together with large backwardly and upwardly growing
molar-alveolar pouch.

Harmonic with the enlargement and backward shifting of the tusks, the elongation of the proboscis, the
fore-and-aft compression of the whole cranium, and the vertical elongation (hypsodonty) of the grinding teeth, are
the following proportional changes in the cranium of Elephas: (a) Hard palate tilted obliquely upwards; (b)

l)alatines reduced lateroposteriorly and widely divergent posteriorly; (c) posterior nares pushed very far back;
(d) large vertical pterygoid wing of ahsphenoid encircles and functionally replaces posterior molar-alveolar pouch
(e) foramen ovale of alisphenoid shifted backward and outward, becomes confluent with foramen lacerum medium;

(f) presphenoid, basisphenoid, basioccipital thicken in median plane, so that basis cranii point sharply downward

at an angle of 90°+ with occipital plane; (g) tympanic bullae flatten and become closely appressed to the skull.

The following is cited from Gregory (1903.1, p. 391)

. .the obliquely placed external portion [Fig. 797] of the orbito.sphenoid
. has been squeezed into a long, thin process;
. . .

internally [Figs. 798, 802] the anterior edges of the basisphenoid are directed outward and backward; both internally and
externally the optic foramen, foramen lacerum anterius, and foramen rotundum, in the order named, are obliciucly arranged
on descending levels from within outward and from in front backward, the whole region ha\-ing been thickened by the separation
[diploe] of the inner and outer tabula; of all the bones, and also sharing in the upward-and-backward tilting of the nasal region
and in the general fore-and-aft squeezing = cyrtocephaly] of the skull, the end result being that the foramina have been pulled
[

out into long tunnels running obliquely forward, outward, and downward; especially internally the fore-and-aft extent of the
alisphenoid proper is brief. Internally the skull has shortened up, one might almost say in bellows fashion, with the optic
foramen on each side at the apex of the internal transverse folding [Figs. 79S, 802], the ridge of the 'lesser wing' of the human
sphenoid. As the skull has also expanded transversely = brachycephaly], the general effect of the internal \iew of the skull is
[

thus that of compression around the center (represented by the basisphenoid) and increa.sing expansion toward the periphery-
somewhat recalling the conditions of the domehke hmnan skull.

'Cuvier's keen jxTception of the cranial distinctions of Elephas primigenius, E. nfricanus, and E. iiulicus arc shown clearly in his definitions of these three
species in the first edition of his "Ossemens Fossilcs" (1806.1, pp. 262 264), in which the three crania are figoi-ed side by side (cf. Fig. 992 of the present Memoir)
in anterior and lateral aspects. See further comment in Chapter XVIII {Mammonkas) on Cuvier's treatment of E. primigenius.
 .

916 OSBORN: THE PROBOSCIDEA

The above principles of the ontogenetic and phylogenetic adaptations in the Elephas indicus cranium cover
three chief cranio-mechanical factors in the order named:

(1) The lengthening of the proboscis or trunk of first importance as affecting aUke males and females.
(2) The lengthening and increasing weight of the tusks, as affecting chiefly the male, and to a much less

degree, the female cranium.

(3) The heightemng of the molar crowns, especially of M^-Mg, as affecting the male and female cranium aUke.

Juvenile Cranium of Asiatic Elephant

Fig. 790. Palatal view (left) of jiivcnilc iTaniiini of A.siatic elephant {Elephas i7uKcus); superior view (right) of same cranium. AftcT Osborn and Gregory
(see Gregory, 1903.1, figs. 1 and 2, ]>]>. 390 and 391).

— Premaxillary.
P.nix. p. As. — Pterygoid wing of .ali.sphenoid. Ip. — Tympanohyal.
h. i. e. c. — Canal for internal carotid
Mx.— M axillary ft.— Pterygoid. cu. — Eustachian opening of tympanic. artery.
— Maxillary iKimli for molars.
.V/x. p. Sq. —Sfiuamosal. a. p. f.— Anterior palatine foramina /. sL in. — Stylomastoid foramen.
.Ua.— Malar. Ex. — Exoeeipital.
o. (canals). /. I. p. — Foramen laeerum j)osterius.
— Postorbital ridge
Po.f. of frontal. pg. — Postglenoid ledge of sriuamosal. i. 0. — Infraorbital foramen.
f. c. /. — Notch, a vestige of condylar
Mx. — Palatine
pal. ledge of maxillary. p.ty. — Pcst-tympanic ledge of .squa- p. n. — Posterior nares. foramen (?) (confluent with/. I. p.).
PJ.— Palatine. mosal, which with forms a
pg. p. a. s. — Alisplienoid canal. i.-. —
Tusk,
Vo. —-Vomer. secondary external auditory meatus. /. I. — Foramen laeerum medium.
in. p' {dm 2). Third —
premolar (or
Ps. — Presphenoid. Ty. —Tympanic bulla. f.ov. — Foramen ovale (confluent with second deciduous molar of authors).
Bs. —Basisphenoid. ly. p. —-Anterior process of tymijanic. /. I. m.). p* {dm 3). —
Fourth premolar (or
Bo. —-Basioeeipital. third deciduous molar of authors).

Osborn (1904-1924), in comparative observations, is inclined to attribute cciually great, if not greater cranio-
mechanical influence to the vertical elongation ( = hypsodonty+bathycephaly) of tlie posterior grinding teeth in
relation toadajjtive radiation in feeding habits, as .shown in a (H)Mipnris()ii of (Fig. 806) llic |)iin'ly I Jiowsing African
elei)hant {Loxodonta) with the browsing and grazing Indian elei)hant (Klcphd^) and with the cliicfly grazing north-
ern mammoth (Mammonteus).
 Po.f.

Ex. 0.

Fig.
797. (Right figure) Orbitosphenoidal region, left side. From
Osborn and
Gregory (see Gregory, 1903.1, p. 393, fig. 4). Tlic view is
obliquely from the
side and from below the malar bone (ef. left figure—
Fig. 3 of Gregory, 1903. 1
).
(Left figure) Juvenile cranium of Asiatie Elephant (.see Gregory, 1903,1,
p. 392, fig. 3). Compare figure 796.
Po. /.— Postorbital ridge of frontal. Mx. p.— Maxillary i)oueh for molars.
Fr.— Frontal. op. f. —
Foramen optieum.
0. s. — External process of orbito- /. I. a. —Foramen laeerum anterius.
sphenoid.
As. —Alisphenoid. /. r. —
and a. a. s. Arcade leading to
Sq. — Squamosal, foramen rotundum and anterior
p. .4s.— Pterygoid wing of alisphenoid. opening of the alisphenoid canal.

Elephss indicus male juv,

Amer. Mus. 44 Ret.

InI'Antile Cu.\nium of AsrATic

Eleph.wt
Fig. 798. Infantile basis cranii.
Skull Elrphas indicu.s (.\mer.
of
Mus. Dept. Mam. 44); same indi-
vidual as that represented in figure
799. One-half natural size. Ob-
serve identification of basicranial
foramina by Gregory in 1903.
process for sttac/iment masfoid region of Penotic Compare infantile separation with
"/ tympgnohya/ the adult fore-and-aft <-ompres.sion
and confluence, cyrtoccphaly (Fig.
process o/tymp. tu//a SCO), of the liasicranial foramina.
918 OSBORX: THE PROBOSCIDEA

Assigning, therefore, a proportional influence to the three great cranio-mechanical factors, (1) tlie proboscis,

(2) the tusks, and (3) the grinders, relative to the frevaiUng feeding habits, whether purely browsing, browsing and
grazing, or purely grazing, we come very near a complete mechanical
shall interpretation of the cranium of the
elephants in the fiv^e generic phyla which we shall presently examine.

E. indicus '

^
Afner. Mus. 3819 Ref.

Pa.

Adult Cr.\niu.m of .\si.\tic Elephant

Fig. 800. Adult palate of Indian elephant {Klcphas indicus
IxFANTiLE Cranium of Asiatic Elephant beiigalensis), Amer. Mus. Dept. Mam. 3819. This is a middle-aged
Fig. 799. Rcfcrrod infantile occiput and jaws of Elcplias specimen with superior molar, ?M', showing eleven to twelve worn
indicus (.\mcr. Mu.s. Dcpt. Mam. 44). This is a much younger plates. One-eiglitli natural size.
si>ecimen than tliat figured (Fig. 797) by Gregory in 1903, p. Compare the fore-and-aft compression ami loiiflucnce (eyrto-
392, fig. 3, whicti shows a long shallow jaw. One-fourth natural cephaly) of the ba.sicranial foramina with the relatively primitive
size. condition in the infantile cranium (Fig. 798).

Comparative Cranial Sections of Elephant Skulls

OsBORN. —In order to ascertain the more profound changes which have taken place in the evolution of the
crania of the elephants, Osborn and Gregory, as stated above, employed in 1902-1903 a method of sectioning the
skull (Fig. SOI) in four different planes, by means of copper wires. A -similar method had been used (Osborn-
Gregory) in studying the cranial mechanics of the peri.s.sodactyl family of titanothercs.
 ::

CLASSIFICATION OF THE ELEPHANTOIDEA 919

These planes were established with reference to the basicranial axes, after the manner of Huxley, Flower,
Lankester, and others in their studies on the cranial axes of other ungulates ; also with reference to the horizontal
grinding surfaces of the molar teeth.

The Five Wire Section Lines (A-E)

Fig. 801. Kc}' to Sections. Young Elephas indicus bcngalensis cranium apparently of the bioail narial 'Dauntela' variety. The
nianner in which the cranial sections are recorded in the figures of this chapter is shown in the above key diagram to sections, also in
diagrammatic figures 803-814.

C, Midvertical, sagittal, nasals to vertex of the occiput. />, Para-occipitofrontal, sagittal, longitudinal rim of anterior narcs to
occipital condyles. A, Horizontal nasal. E, Occipitohorizontal through back of occiput. B, Vertical transverse frontals, intertemporal.

The intensive application of this method by Osborn in 1924 revealed the following Unes of phylogenetic

divergence: (1) Progressive brachycephaly of the skull involves fore-and-aft shortening of the individual bones
( = cyrtocephaly), also an expansion in the vertical planes ( = hypsicephaly). (2) Shortening ( = cyrtocephaly)
and deepening ( = bathycephaly) of the temporal fossae and upward expansion of the grinders (= hypsodonty)
and of the alveolar pouch profoundly alter the bones and foramina of the sphenoidal region. We may summarize
these proportional changes as follows:

Cyrtocephaly Fore-and-aft faciocranial abbreviation.

Cyrtodonty: Fore-and-aft molar-crown abbreviation.
Hypsicephaly: Vertical heightening of the cranium and jaws.
Hypsodonty: Vertical heightening of the molar crowns.
[Brachyodonty: Vertically low molar crowns.]
Acrocephaly: Vertical heightening of the occipitofrontal apex.
Bathycephaly: Vertical deepening of the basicranium, molar alveoli, and jaws.
I5rachycephaly: Broadening of (a) the occiput, or of (b) the zygomatic arches.
(
'yP^ocephaly Downward flexure of the facial to the basicranial axes.
[Orthocephaly: Lack of inclination of the basifacial to the basicranial axis (cf. p. 924 below)].

Ontogenetic Cranial Changes in Elephas indicus

Api^lying the methods described above, it is desirable to examine the ontogeny of the Elephas indicus cranium,
in accordance with the biogenetic law that youthful crania and jaws in all phyla of the Elephantidx more or less

closely resemble each other, whereas adult crania and jaws very widely differ frorn each other.
920 OSBORN: THE PROBOSCIDEA

The chief ontogenetic or growth and age changes are clearly displayed in the accompanying series of Jlgures

(Figs. 801, 807, 809, 799, 798, 796, 797, 802) illustrating the transformation from the infantile to the adult condition

in the cranium of Elephan indicus, which should be examined in connection with the comparative sections, also
with the adult crania figured in the present and succeeding chapters on Mammonteus, Loxodonta, and Elepltas.

Fig. 802. Interior view of skull of youiif^ Asiatic Elopliiint, after GrcRory, 1903, PI. xxiii. Same .skull as tliat rcpri'.si'iitwl in figures 7'J(i, 7'J7.

Crib. p). — Cribriform jilate. —

f.l.p. Foramen lacerumposterius.
orb.sph. ridge — Orbitosjihenoid ridge. — Internal carotid canal.
i.c.can.
sel.tur.— Sella turcica. tym. — Tympanic bulla.
Fr. — Frontal. Pa.int.tab. — Internal table of parietal.
f.r. —Foramen rolundum. Fr.ex.tab. — Outer table of frontal.
f.l.a. — Foramen laeenim anterius. Basisph. — Basisphenoid.
f.l.ni. — Foramen lacerum medium. Ali.sph. — Ali.sphenoiil.

Fronto-occipital and Frontal Growth Curves (Figs. 803, 804). —As shown in midsection D, the frontal
profile of the infantile cranium of Elephas indicus resembles that of the adult jirohle of Loxodonta africana. The
juvenile profile displays the rounded and expanded fronto-occipital curve characteristic of E. indicus, in contrast
to the angular occipitofrontal curve of L. africana. The adult fronto-occijiital profile is a uniformly rounded
dome. The old male pi'ofile exhibits an acrocejjhalic dome, which attains the same heiglit as in the highest
Parelephas jeffersonii (Fig. 810) and is distinguished by the rounded bulbous frontals.

The infantile frontals resemble those of L. africana. The juvenile frontal curve is elongate, (;onvex superiorly,
concave inferiorly or convexo-concave, in contrast to the uniformly convex frontals of L. africana (Fig. 811),
or the uniformly concave frontals of Parelephas and Mammonteus in section (Figs. 805, 800, 810, 811).
 CLASSIFICATION OF THE ELEPHANTOIDEA 921

Nasal Growth Stages (Fig. 807).— The nasal growth stages reflect the increasing aljl)reviation (cyrto-
cephaly) and broadening (brachycephaly) of the cranium. The infantile nasals are short, pointed, and narrow.
The juvenile nasals are broader and uniformly arched. The adult nasals are broad and truncated. The old male
nasals are extremely broad, truncated, and abbreviated.

growth staqe^

SinJicui, ijrowt^ stages

VrrTicaf sagittal
nasals To vsrtsx oeeiput

Fig. 803. Front o-occipital gniwtli ciirvt'S of vertex. Elei>has indicus Fig. 804. Growth curves of vertex. Midfrontal vertical
cranium, one-sixth natural size. This Section D passes between tlie oeeiijital Section of Elephas indicus skull, ono-sixth natural size. The
convexities. The increasing occipital convexities are shown in tlie passage from increasing concavity of the lower part, and convexity of the upper
the infantile to the old male stages of growth. The section is taken at the left part, of the forehead in the passage from the infantile to the old
of the medial line. male stages of growth are shown.

Fronto-intertemporal Growth Stages (Fig. 808).— Like the nasal growth, the fronto-intertemporal
growth reflects increasing abbreviation (cyrtocephaly) and broadening (brachycephaly) of the cranium, as
observed in the infantile, the juvenile, the adult, and the old male growth stages shown in Section B. As compared
with the convex profile of L. africana (Fig. 813), the midfrontal profile of E. indicus is plane. The angle between
the superior fronto- and laterotemporal surfaces of E. indicus (Figs. 808, 813) is sharply angular, whereas in L.
africana (Fig. 813) and Parelephas jeffersonii (Fig. 813) the frontotemporal union is rounded or gently convex.

OcciPiTOHORizoNTAL 809).— The extraordinary brachycephaly of the E. indicus

Growth Stages (Fig.

cranium is most clearly displayed in the rapid lateral expansion of the occiput and deepening of the interoccipital
space. As clearly displayed in figure 809, even the infantile cranium is relatively broad in horizontal section. In
the juvenile stage the bulbous backward growth of the occiput begins with the deepening of the interoccipital
space. In the adult the occipital expansion describes a uniformly convex rounded curve. In the old male the
winglike lateral expansion results in an acute angle between the occipital and temporal faces. This shows that the
extreme of brachycephaly is attained in the broadening of the occiput. In the comparative figure 814 it appears
that this extreme occipital brachycephaly is much greater in the old male of E. indicus than in the adult P. jeffer-
sonii; it also exceeds the occipital brachycephaly of the L. africana cranium.

Palatal Abbreviation 800).—The palate of the juvenile cranium (Fig. 796, left), as compared
(Figs. 796,
with the adult palate of E. indicus bengalensis (Fig. 800) and the adult palate of L. africana (Fig. 1061), proves
that in the transition from the juvenile to the adult stage there is a very marked abbreviation (cyrtocephaly) and
 .

anttrtar narta

longitudinal
Condifle to

middle of n'^fiT half ef paritfala

Fig. S05 ^.- _^-

Pkki.imixaky Stuuv of Fronto-occipital-basilar Planes •'
-^

Fig. Comparative para-opfipitofrontal section D (Fig. 107). Soction carried

80."). / / j

down through tlie bonis cranii to show the widely different occipitofrontal planes of the / / ,'

skull in the three great subfamily divisions of the Elephantidffi. Reduced to one-sixth ..--J^h
'

natiMiil .size. For basis cranii substitute basioccipital plane (see Fig. 1192 below). ..•' I __/
/• ',y'
EbErHANTiN.E = £'icp/ms indiciis, Indian elephant contour. Para-occipitofrontal _^-'

union a right angle, rounded.

LoxODONTiN.E = Loxorfon(a africana, African elephant contour. Occipitofrontal

union rectangular. -maU;
I
,
Tn. ^orimiyefiiu.s ^ai Mus
***"
MAM.\iONTix.E = PnrWcp/m.s
*
iVffcr.'ionii,'
.* ^ Mammoth contoin-. Occipitol'rontal union
_^ , mf? i ,
771^ /..africana
'

nciile.

Tliesc three sections are made from adult skulls in the American Mu.seum collection,
n;iniclv, of Eli/ilias iii<licu.'<, of Loxodania africana, and of a young male skull of Parcle- ^^^^
'"" """ ^'•""""''"^ .t.^/v,^.
l,h„s j'effersonii (Amcr. Mus. 1447.5).' Compare sections in figure 806. '

m ? T.jrfftrserxii TItf. Nat Mils.

I
. -—.. .
,...,..-^ ".-.TT -.V.l'
/026/
m? m? r indtcus

Fig. SOfi

Later Study ok Fkonto-occipital-Molau-3 Planes

Fig. SOO. Para-occipitofrontal soction /) (Fig. SOI) of: Loxodontina- {Loxndnrila africana), Mammontinie (Parelephas jcffcrsonii, Mammonleus
lirimi(iniius), Elephaiitinse (KUphas indicus). Eac-h .section (D) is taken as follows: (1) To the left of the median sagittal line; (2) the inferior surface of the

grinding teeth (M^-M') |ilaced horizontally; (3) the oci-ipital condyles superimposed. It will be ol)scrvcd that:

(1) I,().\<M)<)NTIN.K. In A«.iY»/r)H((i n/rtca/i« the fronto-occipitocondylar angles and the molar teeth (M--M') are profoundly different from those in the
Elephantina' and in th<' Mammontime.

(2) Mam.montix.k. I'areli-phas jeffersonii agrees with .Mammonleus primigenius, male and female-, in the concave frontal profile; /'. jejfer.':onii differs
from .U, primigenius in the greater anteroposterior extent, it is less compres.sed. .U. primigenius, male and female, is the most strongly compressed, i.e.,

hypsicephalic, acrocephalic.

Ei-ephantin.k. The adult KU-phas indicus skull differs widely from the loxodontine and inarnrnontine ty|)es in the frontal convexity as well as in tlie
(3)
more vertical position of llic occipital plane. Compare .sections in figure 805.

'[Subsequently Anu'r. Mus. 1447.5 was provisionally referred by Professor Osborn to Archidiskoitun impernlnr.—'EiWioT.]
-Amer. Mus. 14.5,59 was made the type of Mammonleus primigenius compressus by Oshiwn in 1924 (1924. ()33, )>. .5).

922
 '

CLASSIFICATION OF THE ELEPIIANTOIDEA 923

a moderate lateral expansion (brachycephaly) of the E. indicus cranium. As noted above, the bracliycephaly of

the E. indicus cranium is most marked in the transverse occipital expansion at the very back of the skull, whereas
in L. africona (P'ig. 1061) the transverse zygomatic expansion of the temporal arcades exceeds the occipital ex-
pansion.

Growth of the Jaw (Figs. 799, 797, left). —The elongate infantile jaw of E. indicus (Fig. 799) resembles

the adult jaw of the Upper Pliocene Archidiskodon planifrons (Fig. 849), also to a less degree the adult jaw of L.
africann. In brief, the juvenile jaw of E. indicus is harmonic with the mcsocephalic juvenile cranium (Fig. 797,
left). The growth stages in the jaw are harmonic with the increasing cyrtocephaly, brachycephaly, and bathy-
£.indicui. {/rowth stages Sfcno/v A cephaly of the cranium. Consequently the proportions of the adult jaws of
Horiiifntal rtasjls
E. indicus, as well shown in figure 1204, are abbreviated (cyrtocephalic),
deepened or depressed (bathycephalic) , and broadened (brachycephalic).

Fig. 807. Superior nasal growth stages.

Section A horizontal, showing increasing
breadth of the nasals in the i)assago from the
infantile to the old male growth stages of
Elephas indicu.'i. One-sixth natural size.

Fig. 808. Transfrontal growth stages

Section B. Expansion of the fronto-inter-
temporal space in infantile to old male growth Fig. 809. Occipitohorizontal growth stages, Section E. Complete brachycephalic trans-
stages in Elephas indicus. One-sixth natural formation of the horizontal contours of the back of the occiput of Elephas indicus in the passage
size. from the infantile to the old male growth stages. One-sixth natural size.

In the jaw evolution of every phylum of the Elephantidse, these three contemporaneous adaptations occur to fit

the jaw into its very confined space behind the vertically placed (cyptoceplialic) maxillopremaxillary sockets of
the tusks.

Downward Flexure of Basis Cranii (Figs. 805, 806). —The ontogenetic progressive downward flexure of

the basis cranii (cyptocephaly) is illustrated in the series of comparative figures (Figs. 805,806, 1192) which dem-
onstrate the transition between the relatively horizontal L. africana basis cranii (Fig. 1192) and the most sharply
deflected basis cranii of Parelephas and of Mammonteics (Figs. 805, 806).

Downward Flexure of Dental Alveoli (Fig. 806).— As first observed by Leith Adams, and as again
observed by Gregory and l)y Osborn, the downward growth of the dental alveoli is harmonic with the vertical
extension (hypsodonty) and fore-and-aft compression (cyrtodonty) of the grinding teeth, especially of M^-Ms.
This ontogenetic bathycephaly is not shown in the ontogenetic figures but is clearly displayed in the comparative
figure 806, in which it appears that the second and third molars of E. indicus are much more depressed or bathy-
cephaUc than the second and third molars of L. africana.
 —

924 OSBORN: THE PROBOSCIDEA

Generic Contrasts in Cranial Sections: Loxodonta, Parelephas, Mammonteus, and Elephas

The sectional contours described above were jilotted as in figure 805 and almost immediately revealed the
profound differences between the three chief phyla of elephants.

Encouraged by this preliminary observation, which gives a new clue to the classification and phylogeny of

the Elephantidae, a series of sections were taken in all crania available to the author and it was found that the
other cranial sections supported the inferences derived from the axial section, as shown in figures 805 and 806.

It is demonstrated that the crania exhibit three subfamily types, namely, loxodontine, elephantine, and
mammontine.

(1) Flexure (Cyrtocephaly) of Fronto-occipital-Basilar Planes (Fig. 805). — The midvertical section
(C) combined with the para-occipitofrontal section (D) is the most important, because, as shown in figures 805,
806, and 816, it reveals the angular flexure (cyrtocephaly) which the entire upper part of the cranium bears to
the basis cranii and especially the angle between the occipital 'plane and the basis cranii. It appears that in Loxo-
donta (tfricana the angle of the basis cranii to the occipital plane (100°) approaches a right angle; this is because

SCCTION D SfcriON C
YirTical longitudinal
Virticat layttfal
rim of onferior naraa To eondylfl
natali to vtrftn occifiut

•5-''
iV /

J;' -' -/

I
'

Fig. 8n. Frontal section.'!. Comparative vertical section

Fig. 810. Skull. Mid-occipitofrontal Section D, vertical longitudinal, from of the frontalbones and occiput of the three subfamily divi-
rim (if anterior nare.« to occii)ital condyles, with the ba.sis crnnii in a uniform plane. sions of the elephant family (('), vertical sagittal, nasals to
Reduced to one-sixth natural .size. vertex of occiput. Reduced to one-sixth natural .size.

I.oxoDONTiNE type {Loxodonta ofricann, male), fronto-occipital .angulate, (Loxodonta africnna, male), convex.
I.oxoDONTi.VE type
pitcliing forward. Elephantine type (Elephas indiciin, adult), concave.
Elei'ii.w'ti.ne tyi)e {KUphns imliciiK, male, adult), fronto-occipital rounded, Elephantine type (Elephas indicus, old male), con-
convex. cavo-convex.
M.\MM0NT1NE type (Parelephas jefersmiii, adult males, and a younfi male'), Mammontine type (I'arelephas jeffersonii} young m.ale),

fronto-occipital angle concave or slightly convex. imiformlv concave.

the cranium of the African elephant as compared with that of Elephas indicus is less brachycephalic, moi-e orthoce-
phalic, and less bathycephalic or vertically deepened. In E. indicus the occipital plane forms a more open angle
of 121° with the basis cranii, expressive of its greater bathycephaly. A still wiilei' angle (130°) however, , is attained
ill the bathycephalic Parelephas jeffersonii, while in Mammonteus primigenius the angle rises to 136°.

'[Subsequently .\mer. Mus. 1447.') was provisionally referred by Professor Osborn to Archidiskodmi itnpirntor. — Editor.)
 CLASSIFICATION OF THE ELEPHANTOIDEA 925

(2) Deepening (Bathycephaly) of the Molar-3 Plane

(Fig. 8()6.)~The horizontal grinding surface of
the superior molars below, coniiiared with the cranial profiles above (Fig. 806), reveals bathycephaly as the
most
distinctive cranio-mechanical adaptation.

In the LoxoDONTiN^: (a) The bathycephaly is less extreme: (b) the occipitofrontal union is less elevated
above the horizontal M'-^ grinding surface; (c) the fronto-occipital plane is at an oblique angle with the molar-3
plane; consequently the Loxodonta skull is less hypsicephalic.

In the Mammontine it is observed (Fig. 806) : ( a) That the

SecTiON A
HartjonTol no5aU bathycephaly of Mammonteus primigenius and of Parelephas jeffer-
sonii is practically the same as that of Elephas indicus, because the
molar-2-3 plane descends to the same degree below the occipital con-
dyle; (b) while the Mammontinae agree with the Elephantinae in
bathycephaly, they differ widely in the fronto-occipital profiles, as
shown in comparison of the line E. indicus with the line 'E.' jeffersonii;
Fig. 812.Nasal contour.s. Comparative horizon-
tal, Section A, of nasals showing the broad nasals of
(c) Mammontinae agree with each other in the fronto-occipital
the
Loxodonta africana and of Elephas indicus as compared profiles, as shown in the lines 'E.' jeffersonii 'E.' primigenius female
with the pointed mammontine nasals of Parelephas
jeffersoiiii, male, of P. jeffersonii,^ young male, and of P. [
= type of Mammonteus primigenius compressus Osborn, 1924], 'E.'
jeffersonii, adult male — all in the American M\i.seum primigenius male (d) in the Mammontinae the M. primigenius male
;
collections. Reduced to one-sixth natural size.
and female profiles seem to be relatively more hyjjsicephalic and
Sfcno/v B C, a/,ricanu.*. fjule
VerTirof Transverse^
frvriTal^^ ^^.-^""^
I
^-^~-^^ £2.''Ji::'i.^^°iylL

/\, _£ indtcus. old malt

/ Jl
Y
I '1 ^/.^j:Cfjj.?2.\'. /'

\C. - :^./?//}?j:^^ji:. _ _^\

// ... -^ /*//*'"**"?.'/„

Fig. 813. Midfrontal or intertemporal forehead.

Uniformly convex tran.sfrontal, Section B, of Loxo-
donta africana, male, as compared with the slightly con-
cave frontal section of an adult Elephas indicus and the
planoconcave frontal sections of three skulls of the
mammontine type of Parelephas jeffersonii all in the —
American Museum collections. Reduced to one-sixth Fig. 814. Oceipitohorizontal, Section E, through back of occiput at broadest portion in the
natural size. three subfamily divisions of the elephant family. Reduced to one-sixth natural size.
LoxoDONTiN.B = Loxorfon(a africann, uniformly LoxoDONTiN.« = Loxo(fon/n africana, broad convex flare of occiput; deep ligamentum nucha;
convex. pit. Indicated by
EhEPHANTiNX = Elephas indicus, planoconcave, Ei.EPH.\iiTi^M = Elephas indicus, very broad, acute flare of occiput; ligamentum nucluv pit
borders angulate. moderately deep. Indicated by - - -
Mammontin.e = Poreiep/tas jeffersonii, planocon- MAi,tMOiiTitix = Parelephas jeffersonii, convex flare of occiput, less expanded; ligamentum
cave, transfrontal borders rounded. nucha; pit moderately deep. Indicated by . . .

acrocephalic than those of P. jeffersonii; consequently the Mammonteus cranium suffers even greater fore-and-aft
compression than the Parelephas cranium.

The Elephantin/E profile is followed in the dotted line E. indicus: (a) The bathycephaly indicated by the
M^'^ depres.sion is the same as in Parelephas jeffersonii; (b) the fronto-occipital profile of E. indicus differs widely
from that of P. jeffersonii and still more widely from that of Mammonteus primigenius ; thus the E. indicus crani-
um is equally bathycephalic but it is less hypsicephalic and acrocephalic than either that of Parelephas or
Manunonteus.
'[See footnote on opposite page. — Editor.]
 .

92fi OSBORN: THE PROBOSCIDEA

Hypsicephaly and Acuocki'iialy in Fronto-occipital Phofiles (Fig. 810). In the Loxodontines, as

(3) —
.sliown also in figures 805 and 806: (a) The angular fronto-occipital profile is totally different from that of either

Parelephas or Elephas (b) in Elephas, both in the adult and old male stages of E. indicus, there is a rounded dome-
;

like fronto-occipital profile, which rises to a great height in the old male; (c) in the "Parelephas jeffersoyiii"
crania, shown in the line 'E.' jeffermnii, the cranium is equally hypsicephalic, but it is not acrocephalic, since it

reaches an acute apex through the characteristically mammontine concavity of the frontals.

In the frontal sections of the forehead profile we readily distinguish between the uniformly convex profile of
Loxodonta africana, the concavo-convex profile of Elephas indicus, and the concave profile of Parelephas jeffersonii.

The concave frontal profile of P. jeffersonii relates this animal to Mammonteus.

(4) Brachycephalic and Hypsicephalic Nasal Contours (Fig. 812). —The broad nasals of Loxodonta
africana male are harmonic with the brachycephaly of the cranium. In the Elephas indicus adult the nasals exhibit
a rounded profile, but in the E. indicus old male the end of the nasals is truncated. In Parelephas jeffersonii the
crania examined exhibit uniformly acute or pointed nasals clearly distinguishable from those of either Elephas
or Loxodonta.

(5) Convex or Concave Transfrontal Sections (Figs. 806, 801, 805, 813). —In Loxodonta the forehead or
frontal bone is convex in both planes, sagittal and transverse, as shown in figures 806, 805, and 813. In Elephas
indicus the transfrontal section (Fig. 813) is seen to be plane with sharply angulate lateral borders defining the
temporal fossae. In Parelephas jeffersonii the transfrontal section is rounded but less convex than that of L.
africana.

(6) Relative Brachycephaly of the Occipital Planes (Fig. 814). — Whereas the Loxodonta africana
cranium, as shown in the plain line 'E.' africanus male, is in general more brachycephalic than that of Elephas

indicus, the greatest lateral expansion of the occipital region is attained in the E. indicus old male, in which the
expansion far exceeds that of Parelephas jeffersonii, as indicated in the lines 'E.' jeffersonii. The pit for the liga-

mentum nuchse is similar in the E. indicus old male and in 'E.' jeffersonii, whereas the pit in Loxodonta africana
is very much deeper.

Summary. —In summing up the cranial and profile proportions of Loxodonta, of Parelephas, of Mammonteus,
and of Elephas in respect to the .six chief regions described above, we observe that Loxodonta differs profovmdly in
every character from Elephas, Parelephas, and Mammonteus; this is ])robably due to profound differences in
feeding habits. The crania of Parelephas and of Mammonteus agree in their bathycei^haly with the cranium of
Elephas, but differ widely in their acrocephaly. There appear to be clear lines of demarcation into three subfamily
groups which are summarized below (p. 932).

While the mechanical problem of the tusks and of the proboscis remains relatively the same in these three

generic stages of cranial evolution, the mechanical problem of the grinding teeth changes, as follows: (1) Tlio

relatively low-crowned grinding teeth of Loxodonta are correlated with a relatively less brachycephalic or meso-
cephalic cranium; (2) in the opposite extreme the many plated, high-crowned, extremely hypsodont grinding
teeth of Mammonteus are correlated with an extremely hypsicephalic and bathycephalic cranium; (3) extreme
fore-and-aft compression of the cranium of Mammonteus (i.e., cyrtocephaly, bathycephaly) is coordinated with
extreme shortening, heightening, and multiplication of the dental ridge-plates (i.e., cyrtodonty, hypsodonty,
polydiskodonty)
 CLASSIFKJATION OF THE KLRPHANTOrDRA 927

II. DENTAL AND CRANIAL ADAPTATION TO PREVAILING FEEDING HABITS THE

KEY TO PHYLOGENETIC CLASSIFICATION
As among other ungulates, the choicest elephantine diet abundant in favorable seasons of the year is that most
frequently resorted to and most dominant in cranial and dental mechanics. Seasonal or climatic and secular
changes may have enforced changes of prevailing diet. Persistence of dental type is favored by persistent climatic
conditions and flora, as we believe was the case in the true Stegodon, which continued to browse through the Upper
Tertiary and into early to Upper Pleistocene time in the tropical forests of India and the islands of the East
Indies, eastward into China and Japan. Accordingly the grinding teeth of Stegodon multiply their ridge-crests

but do not become sensibly hypsodont as in the elephants. The cranial evolution of the Stegodonts also appears
to be distinctive.

1 .
RIDGE-PLATE FORMULA OF PRIMITIVE AND PROGRESSIVE GENERA IN ADAPTATION TO
PREVAILING HABITS OF FEEDING
The ridge-crests of Stegodon are transformed into the ridge-plates of the Elephantoidea.'
Ridge-plate Formulae.— Falconer (1868, Vol. II, p. 13) in defining his three subgenera of Elephas laid great
stress on the number of ridge-plates in the intermediate molars, namely, Dp 4, M M1, 2, as follows:

Subgen. Stegodon, hypisomeris, e. g., Dp 4= 7, M 1 = 7, M 2= 8.

" Loxodon, hypisomeris, e. g., Dp 4= 7, M 1= 7, M 2= 8.
" Euelephas, anisomeris, e. g., Dp 4 = 12, M 1 = 14, M 2=18.
Throughout his masterly work "On the Species of Mastodon and Elephant Occurring in the Fossil State in
Great Britain," 1863, Parts I and II, and "On the American Fossil Elephant of the Regions Bordering the Gulf of
Mexico," Part III, Falconer makes a profoimd study of the ridge formulae, describing the formulae in various species
with an accuracy which cannot be challenged today. This great section of Falconer (1868, Vol. II, pp. 1-291) is

a masterpiece of accurate observation, including very appropriately a discussion of the unity or plurality of species
(pp. 254-271) based on the vertebral formulae, as cited below.

2. FOOD OF THE INDIAN AND AFRICAN ELEPHANTS AND OF THE MAMMOTH

Falconer concludes Section III of his "Palaeontological Memoirs" of 1868, Vol. II, with a philosophical
treatise (pp. 277-291) on the "Food of Living and Extinct Elephants," which he connects very closely with the
comparative structure of the ridge-plates in Elephas, Loxodonta, and Mammonteus.

Food of the Indian Elephant [Falconer, 1868, II, pp. 277-280]. —

The 'Sal,' or 'Taiai' Forests, which stretch at the foot
of the Himalayahs, from lat. 30°, where the Ganges and .Jumna esca])e from the mountains, to the Brahmapootra, embracing
a range of several hundred miles, are here selected to furnish the chief illustrations which I have to adduce. They everywhere
aboimd with Elephants, southwards from lat. 30°, which may be regarded as the extreme northern limit of the habitat of the spe-
cies at the present day. Forests presenting similar phy.sical characters extend along the continuation of the same range, through
Sylhet, Chittagong, Arracan, Pegu, and the Tenasserim pro\-inces, to the point of the Malay Peninsula; they become more and
more troi^ical in their vegetation, and, as a general rule, the IClephants improve in size, form, and vigour, according to their
more southern habitat. ... In fact, the range of his (Indian Elephant] arboreous .selection is restricted within a narrow circle, and
mainly to the foliage and branches of trees that abound in milky juice which is not acrid, belonging to the families of the
M areas, Artocarpex, and Sapotacex, such as species of Ficus, Balis, Artocarpus, Bassia, and Mimusops [Footnote: 'Also Mesiia

'[It will be noted from the following quotation (see Vokinic I, ]>. 25, of the present Memoir) that Professor Osboni subsequently altered his opinion: "It
lias been assimied by praetieally all palaeontologists that the Elephants were descended from the Stegodonts. This assumption now proves to be erroneous,
for neither the Stegodon gi-inding tooth with enamel valleys closed at the bottom, nor the Stegodon cranium with its extremely short face, can give rise to the

—
elephantoid molar or the face of the elephantoid cranium." Editor.]
 —

928 OSBORN: THE PROBOSCIDEA

ferrea (Nat. Onl. CliiKifwew), on the iiuthority of Tenuont's \at. Hist, of Coylon, p. 230.']. Of tlioso, by far the greater part of
his stai)lc food derived from the colossal fig-trees which abound in the forests of India; sucli as Finis Indica, the 'Bur,' or
is

Banyan-tree; F. religiom, 'Peepul,' or 'Bodhi-droonia' (Tree of knowledge); F. venom, 'Pilkhun'; F. ranlifolia, 'Gujeena,' or
'Assoud'; F. glomernta, 'C.oolur'; /''. Tsiela, 'Kuth-bur'; and in Assam, Ficiis eloxtira, or the 'India-rubber tree,' besides other
more southern species of similar habit and properties. The strong partiality of the Elejihant for the.se trees is so well known
to the natives, that the 'Obees,' or Pit-falls, for entrapi)ing the animal are invariably constructed in their neighbourhood, and
many of their old Sanscrit names connect them specially with the Elephant [Footnote: "Nagbhundoo," "Koonjurashun";
'

"Gujashun," and "Gujbhukshuk"; all being to the effect of "food of Elephants." {Vide Madden, Journ. Asiat. Soc. Beng., vol.
xvii. p. 380.)']. He tears down their branches, and crunches the twigs and leaves, stripping off the lactiferous bark of the larger
boughs. The Elephant of the 'Sal' Forests also derives occasional food from the foliage and fruit of Artocarpus Lackoocha,
'Dhao'; ... Of aliment derived from the roots of Dicotyledonous trees and shrubs, such as the African EIe])hant is said to
affect, I know of but one form in the 'Si'd Forests' which the Indian species is known to touch, namelj\ the huge tuberous
dilatation of the ligneous root of the Scanilent Piieraria tuberosa, 'Sural.'
, . Among the monocotyledonous families, a very large
. .

portion of his habitual fare is derived from the Graininex, and more si)aringly from Palms; of the former, he luxuriates on the
young shoots and tender foliage of various species of Bamboo, which occur in vast abundance, together with the fleshy albuminous
fruit of Beesha liheedii, found in the southern forests. The 'jhils,' or swamps, to which he resorts, are slieeted with the gigantic
reeds of Ariindo kiirka, 'Nul,' the young culms of which, together with the stems and leaves of Typha FJIephantina, 'Patela,' at
certain seasons, constitute a favourite food of the Indian Elephant. The open glades and prairie lands are covered with species
of Saccharum, forming what is called 'Grass Jungle,' compo.sed chiefly of i5l. sponlaneum, 'Kas,' interspersed with S.fuscum, 'Tat,'
S. Sara, 'Surkura,' or 'IMoonj,' S. exaltatum, 'Suroo,' &c. Clumps of these grasses are twisted up by his trunk, in his journeys
to and from the forests; they are beaten against his legs to free the roots from sand, and then subjected to mastication. The
sand which still adheres to these grasses, together with the large quantity of silica contained in the leaves and culms of .Sac-
chaniin spontdnciim, the most characteristic species of the grass jungle, performs an important duty in the economy of wear of the
Elephant's molar teeth [Footnote 'The excessive abundance of silica in the culms and leaves of S. sponlaneum is practically shown
:

when it is attempted to mow it with an English scythe. After a few sweeps, the edge of the implement is rounded off, as I have
repeatedly witnessed.'] ... It is difficult to conceive of a mechanism better adapted to the duty which they have to perform
than is presented by the molars of the Indian Elephant. Taking the three true molars, which serve during the adult stage of
the animal, they are composed successively of 12, 16, and 24 ridges. Each ridge has the core formed of a high wedge-shaped
plate of ivory; ... A constant equilibrium is maintained, in the normal state, between the nature of the food, the waste of the
crown-surface, the absorption of the fangs, the forward movement of the body of the tooth, and the replacement of the worn-
out portion by a succession of fresh plates, jjrotruded from behind.

—
Food of the African Elephant [Falconer, 1868, II, pp. 282-284]. Our knowledge of his food is, therefore, of a vague
and general character, being deri\ed from the cursory observation of travellers, whose attention was not specially directed to
the subject. The molar teeth of the African Elephant are intermediate, in construction and triturating characters, between
those of the Euelephnnles, or Elephants proper, and the fossil Slegodon.s. They present, in the three intermediate and last
molars for the ridge-formula, the successive ciphers 7: 7, 8, 10; while E. aniiquiis presents the ciphers 10: 10, 12, 16, and E.
piimigenius and E. Indicus, 12: 12, 16, 24. The aggregate of the series of ridges in the first amounts only to 32; in the second
to 48; and in the two last to 64 ; involving a great difference in the triturating mechanism of the teeth. In the African form the
molars are also shorter, narrower, and of less elevation, than in the Asiatic species. The discs of wear, instead of the narrow
trans\erse bands seen in the latter, exhibit the well-known rhomboidal expansion characteristic of the species. Instead,
therefore, of being adai)ted to contu.se and triturate the branches and twigs of trees, they are better suited for squeezing and
crushing leaves, and succulent stems or roots. The habits of the animal, as observed by travellers, are in accordance with these
indications. Besides browsing on the foliage of the Mimosas and Acacias, which abound in Southern Afiica, they tear up the
trees of certain si)ecies of these genera by the roots, aided, according to Pringle, by their tusk, used as a crow-bar (?), and they
devour the succulent i)arts of these roots in the inverted trees [Footnote: 'Cited in the "Library of Entertaining Knowledge."
Menageries, vol. ii. p. 36.']. Burchell mentions a small sjiecies of Prosopis, P. Elephnntnrhiza, as yielding a favourite food to
the lOlephant [Footnote: ^Acnciu Ehphuiilinu, Hurch. "Tra\els in South Africa," vol. i. ji. 236. Elephanlorhiza Burchellii,
Benth.'i; and the succulent 'Spekboom' Puittdacnria Afra, or 'Tree Purslane,' is noticed by most travellers as yielding another.
That the African Elephant, such as we now see it, formerly extended to the South of Eurojie, has been put beyond question
1st, by the researches of Lartet upon remains found in the neighbourhood of Madrid [I'ootnote: 'Comptes llendus. 22 fev. 1858.
Tom. xlvi.'l 2nd, by the remains discovered by Baron Anca in the cave of San Teodoro in Sicily [Footnote: 'Bullet. Soc. Geol.
;

de l'"rance. 2e Ser. t. x\ii. p. 684. PI. xi. figs. 5 & 6.']; 3rd, by a molar from Grotta Santa, near Syracu.se, described by the
(
'anon Alessi (Footnote: '.\tti dell' .\ccad. di Scienz. Xatur. tom. vii. p. 2'23.'], and identified by myself; and lastly, by a molar
exhumed by M. Charles laudin, in lSo8, in a cave near Palermo. ... Of the more ancient lOuropean fossil species, E. aidiquus
(

is that which most resembles the African I'llephant in the mesial expansion of the discs of its worn molars. But the character is
shown in a much less degree, and the great difference in the ridge-formula of the two si)ecies places them in two distinct sub-
genera.

—
Food of the Mammoth [Falconer, 1868, II, p. 284]. In order to estimate the force and value of the arguments which
have been raised on this head, it is necessary to institute a rigorous comi)arison between the mechanical conditions of the molar-
crowns of the Indian Elei)hant and of the fossil s])ecies. The ridge-formula is the same in both, being for the four last t(;eth of
the upper jaw 12: 12, 16, 24. The number of ridges in the three first of the.se is very constant; the last, as already stated, is
variable within certain limits, twenty-two being the most common number. Taking the penultinuite, as in the case of the
Indian Elei)hant, the worn surface of the crown would show sixty-four alternations of unequally hard materials.
 CLASSIFICATION OF THE ELEPHANTOIDEA 929

Falconer points out that the teeth of the mammoth are far less adapted to browsing and especially to crush-
ing the woody fiber and bark of treesthan the teeth of the Indian elephant, and he ingeniously argues a priori
that the food of the mammoth was widely different from that of the Indian elephant. This argument, based on
the structure of the molar crowns, is fully sustained by the discovery that, during certain seasons of the year at
least, the food of the mammoth consisted chiefly of grasses, as quoted in full from Felix in his description of Eleplias
primigenius (see Felix, 1912, p. 1127 of this Memoir).

3. SEASONAL CHANGES IN FOOD OF THE MAMMOTH

It proves that the extremely hypsodont finely plated teeth of the
, mammoth are principally adapted to the
northern grasses which j^rcvail on the tundras and Arctic prairies during the summer season. As recently described
by Stefansson, the summer tundra flora closely imitates that of the grasslands which we know as 'prairies' in
temperate latitudes. In this brief season the mammoth obtained its chief food supply for the year, and, like other
northern Herbivora, stored up large reservoirs of fat which were drawn upon during the long Arctic winter
season. In summer a grazer chiefly if not exclusively, in winter it became a browser, feeding upon twigs and the
branches of conifers and woody substances which contained materials of far less nutritional value. Thus its
habits directly reversed those of the African elephant (Loxodonta) which , is chiefly a browser and incidentally, and
for the sake of food variety, a grazer, or of the Indian elephant, which is both a browser and a grazer. Similar
seasonal habits are doubtless characteristic of the northern types of horses with extremely hypsodont teeth,
primarily adapted to grazing, secondarily to browsing.

Conclusion. — The number, compression, and elevation of the ridge-plates increase in all generic phyla, as the

primitive browsing habit develops into a grazing habit.

4. SUMMARY OF PROGRESSION FROM BROWSING TO GRAZING DENTITION

(1) The proboscis, the tusks, and the grinding teeth of the Elephantidse originate in the browsing and plant-
uprooting habits of Stegodon with elongate tusks.'

(2) As in other ungulates, the Elephantidse first passed through a purely browsing stage with brachyodont
grinders, into a browsing stage with hypsodont grinders, typified by primitive species of Archidiskodon like A.
proplanifrons and .4. planifrons.

(3) There also arose the browsing and crushing stage with hypsodont and loxodont grinders, typified by Loxo-
donta africana, in which the teeth were adapted to browsing not only on leafage but on the stems, twigs, branches,
and roots, in which the plant-uprooting functions of the tusks were useful. There is little or no record of grazing
in this stage.

(4) Adaptation to seasonal habits of browsing on the more tender foliage and of grazing on the coarser weeds
and grasses, as well as on tender shoots and buds, is typified in the still more hypsodont grinders of Elephas
indicus, which closely parallel in the elongation and multiplication of the ridge laminae species of the genus
Parelephas.

(5) Finally there arose the extremely hypsodont, thin-plated, reatively smooth-crowned grinding teeth of
Mammonteus primigenius, chiefly adapted to grazing, but dining certain seasons of the year forced into use for
browsing purposes.
'[But sec footnote on page 927 above. — Editor.]
930 OSBORN: THE PROBOSCIDEA

The primary ridgo forinulip and the primary cranial characters in the several subfamily and generic phyla
of the Elephantidae were probably similar. We have seen that the juvenile Elephan indicua cranium resembles in

many respects tlie adult Loxodonta ufricana cranium ; the only primitive true elephant cranium we know is that
of Archidiskodon planifrons, which it would be interesting to section and compare with Stegodon on the one hand
and with Elephas indicus on the other.

As clearly set forth in the introductory pages of this chapter, there is so much parallelism, and even conver-

gence, in the grinding teeth characters of different phyla of elephants as they pass from the chiefly browsing into
the browsing and grazing and into the chiefly grazing and browsing types, that it is only by the combined analysis
of the tusks, of the grinding teeth, of the cranial axes, and of the cranial planes and profiles, that we can distinguish
these great family and subfamily phyla from each other.

III. VERTEBRAL DISTINCTIONS OF ELEPHAS, LOXODONTA, MAMMONTEUS,

AND PARELEPHAS
Divergence in vertebral and rib formulae in the dorsal, lumbar, and caudal regions will undoubtedly aid us in

distinguishing genera, species, and subspecies of the elephants. We have not at present accurate and final observa-
tions on the distinctive characters of the ribs and of the backbone, as shown in the following review.

1. VERTEBRAL FORMULA IN THE ABOVE GENERA

Falconer in his invaluable Memoir of 1863, as reprinted in full in the "Palaeontological Memoirs" of 1868,

Vol. II, pp. 212-291, gives an excellent review of the existing and extinct species of elephants, including his own
observations (presented in more detail in Chap. XX, pp. 1312, 1313 of the present Memoir) on the vertebral as well
as on the ridge-plate formulae; lie sununarizes (p. 257) the conclusions of Schlegel, 1845-1867, and of Tomminck,

1862 (see Sclatcr's translation, 1862.1, of Schlegel's paper on "The Sumatran Elephant") in the following table:
 CLASSIFICATION OF THE ELEPHANTOIDEA 93I

Falconer's own observations and records differ from those of Schlegel and Temminck and may be summed up
as follows:

Elephas primigenius: In the mammoth there appear (Schlegel) to be 18 dorsal vertebra; and 18 ribs. According to Tilcsius, in the
niaiiinioth skeleton {E. primigenius) in St. Petersburg there are 19 dorsal vertebrae and 19 ribs.

Lo.rodonUi africana, African elci)hant:

Congo (Cuvier, de Blainville, 1844, Laurillard, Daubenton) an imported elephant: 20 dorsal vertebra and 20 pairs
of ribs; total
formula: 7 cervical, 20 dorsal, 3 lumbar, 3 sacral, 31 caudal vertebrae; 20 pairs of ribs.
Cape of Good Hope: 7 cervical, 20 dorsal, 3 lumbar, 4 sacral, 26-30 caudal vertebra; 20 pairs of ribs [typical capensis].
Algoa Bay' (Flower- Falconer) 7 cervical, 21 dorsal, 3 sacral, 30 caudal vertebra;; 21 pairs of ribs.
:

Sumatran elephant (Schlegel) 20 dorsal vertebrae and 20 pairs of ribs.

:

Indian Elephant:
Ceylon (Camper, de Blainville, Cuvier) 20 dorsal vertebrae and 20 pairs of ribs.
:

Bengal (( 'amper, do Blainville, Cuvier) 20 dorsal vertebrae and 20 pairs of ribs.

:

Bengal (Falconer) 7 cervical, 20 dorsal, 3 lumbar, 4 sacral vertebra;, 20 pairs of

: ribs; 20th dorsal small and unsymmetrical.

Falconer concludes that the continental elephant of northern India varies in the number of its dorsal verte-
brae from 19 to 20, as the African varies from 20 to 21.

Vektebkal Formulae According to Falconer's Summary

[According to Eales (1929)^
African Indian Sumatran African Foetus
Cervical vertebrae 7 7 7 7
Dorsal vertebra; 20-21 19-20 20 21
Pairs of ribs 20-21 19-20 20 21
Lumbar vertebrae 3 3 3 3
Sacral vertebrae 3 [-4] 3 [-4] 3 [-4] 6
Caudal vertebrae 26-31 26 [-34] 26]

Vertebral Formulae According to Flower's and Osboun's Summary of 1926^

Loxodonta africana Elephas inrlicns MammontcKS primigenius Parelephas jeffersonii

Flower, 1885 "Jumbo" Amer. Mus. Flower, 1885 Falconer and Amer. Felix, 1912 Type
Dept. Mam. 3283 Mus. 14559 { = M. M. primigenius Amer. Mus. 9950
primigenius com-
pressus Type)
Cervicals 7 7 7 7 7 7
Dorsals 19 20 19-20 18-19 19 19
Pairs of ribs 18-19 19 19
Lumbars 4 3 5-3 4-3 5 4
Sacrals 5 4 4 4-3 4 5
Caudals 24+ 21 24-30+ 21 21 (Salensky)^ 12+

'[Citod from letter, July 2.'), 1929, from Dr. Hubert Collar, Curator, The Mu-scum, SafTron Walden, E.ssex: "I liave verified the particulans you send
relating to the skeleton of the African Elephant in this Mu.seuni and findthem to be correct, except that a few of the caudal vertebra; are missing."
"The geographic locality is also correct according to our records, but I would irainttlie terra 'imported from Algoa Bay' does not necessarily
out that
mean that the animal was killed at that place. .'Actually, this s|)ecimen with many was sent from Port Elizabeth in 1833, and so far as I know, there
otliers
is no record showing exactly where it was obtained. It is a fair inference, however, from indirect evidence that the i)lace was somewhere in the locality."
—Editor.!

'Eales (1929, pp. 223, 224) observes as to the Congo elephant: "The number of fcetal vertebra; is: Cervical, 7; Thoracic, 21; Lumbar, 3; Sacral, 6;
Caudal, 26. .Six sacral vcrtcbnc bear facets for the ilium. There is, however, no fusion of vertebra; in this region in the foetus.
. . The difference between the . . .

African and the Indian Elephant lies in the thoracic and sacral regions. The African Eleijliaiit has a larger number of ribs but fewer sacral vertebra' than
the Indian Elephant (adult). In the foetus, however, there are six vertebral attachments on the ilium, a number greater than any given for the Indian
Elephant. . . .

Foetus.— [In addition to the 21 dorsal vertebra;]: sternal ribs, 6; indirectly sternal ribs, 11; free, -1.

—
Adult African (Oxford Museum). ti, 10, 't, giving a total of 21 in t^ach case.
—
Adult Indian. Various figures, e.g. Camper, 8, 12; Perrault, 7, 13; Hluir, 8, 1 1
— the details of indirectly sternal and free ribs not being given."
^Compare a more recent summary (1929) in Chapter XIX, p. 1227.

[See Zalensky, Vladimir Vladimirovich, in Bibliography of Volume I of the present Memoir. — Editor.)
 'X

^ 6
C <
I^ CO

X I

:=: c
r: <
p &-

2 o

o 3

— : X
 ;

IV. SYNOPSIS OF SUBFAMILY CLASSIFICATION OF THE ELEPHANTOIDEA

As shown in the three leading sections of the present chapter, the cranium affords the chief basis of subfamily
classification, secondly the tusks, and thirdly the grinding teeth. The three genera Archidiskodon, Parelephas,
and Mammonteus appear to show profound subfamily resemblances in the cranium and tusks, while they are
widely divergent in the grinding teeth.

In tlie succeeding chapters distinctions between these profound subfamily resemblances in the cranium,
tusks, and grinding teeth are pointed out in detail, in addition to the accompanying synopsis (p. 932).

V. FINAL SUMMARY OF CHAPTER

The present Chapter XV demonstrates that profound cranial characteristics separate generic phyla iMch
hitherto have been united by parallelism or convergence in characters of the grinding teeth.

The principles of adaptive radiation and of phylogenetic classification as applied to the Elephantidae, and
developed chiefly through the researches of Falconer, Weithofer, and Osborn, yield a number of very surprising
results and run counter to all previous systematic and phylogenetic conclusions. The systematic synopsis at the

close of the present chapter (p. 932) summarizes the conclusions arrived at in this and the following chapters,
resulting from the most intensive and difficult research incurred in the prei^aration of this entire Memoir.

The author hopes that the reader will suspend judgment as to this surprising divergence and polyphyly in

the Elephantidse until the succeeding chapters of this Memoir have been thoroughly examined, namely, C'hajiters

XVI (Archidiskodon), XVII {Parelephas), XVIII (Mammonteus) included within the subfamily Mammontinffi

Chapter XIX (Loxodontinae), and Chapter XX (Elephantinae).

933
 e

"3

J3

S
o
a
<
6h

*^ T3
X
 Chapter XVI

THE GENUS ARCHIDISKODON (SUPERFAMILY ELEPHANTOIDEA), MOST

PRIMITIVE MEMBER OF THE SUBFAMILY MAMMONTIN^

Archidiskodon, parelephas, and mammonteus, united in the subfamily mammontin^ by similar

PROFOUND cranial CHARACTERS. WiDELY SEPARATED IN THE RIDGE-PLATE STRUCTURE OF THEIR GRINDING TEETH,
ALSO IN THEIR HABITS AND GEOGRAPHIC DISTRIBUTION: ARCHIDISKODON IN SOUTHERN PARALLELS; PARELEPHAS
IN TEMPERATE PARALLELS; MAMMONTEUS IN NORTHERN PARALLELS. SPECIES, PHYLOGENY, AND DISTRIBUTION
OF ARCHIDISKODON.

I. Historical Introduction. Review of successive discoveries in Nebriiska, Kansas,

1. History of the subfamily IVIammontinae. Texas, CaHfornia, Florida, South CaroHna, Oklahoma,
Indiana, and Mexico.
2. History of the genus Archidiskodon.
3. Order of discovery and description of twenty-two Archidiskodon imperator Leidy, type characters, distinctions
species of Archidiskodonts. from Elephas [Parelephas] colmnbi, and comparison with
4. Archidiskodonts of Eurasia and America. A. nieridionalis and A. planifrons.

5. New Archidiskodonts and Loxodonts of Africa. Type dental, cranial, and skeletal characters.
6. Approximate ascending phylogenetic order of succes-
Observations on the characters and geographic distribution
sion of species of Archidiskodon and Parelephas
of ^. imperator in the United States and Mexico.
(1928).
Synopsis of A. imperator in the American, U. S. National,
II. Characters of Archidiskodon and Metakchidiskodon
Nebraska State, City of Mexico, and other museums.
AND Included Species.
1. Archidiskodonts of southern Eurasia. Cranial characters of Archidiskodon in comparison with all

2. Archidiskodonts and Metarchidiskodonts of South

the known American crania.

Africa. Skeletal characters of Archidiskodon from a comparison of

3. Archidiskodonts of the United States and Mexico. the various skeletal materials known.

I. HISTORICAL INTRODUCTION
Subfamily Classification. — In its profound cranial characters, as shown in figures 805, 806, and 816,
as well as in its cranial profile, we observe the surprising fact that Archidiskodon is far closer to Mammonteus
and Parelephas than to either Elephas or Loxodonta. Consequently the subfamily Mammontinae, as shown on
page 932, appears to include three great branches, Archidiskodon, Parelephas, and Mammonteus, which exhibit
three distinct modes of adaptation of the grinding teeth, independently developed. The author has been very
slow in reaching this conclusion, for, as shown below, his subfamily Euelephantinse = Mammontinae] at first
[

included the northern mammoths only.

1. HISTORY OF THE SUBFAMILY MAMMONTIN.E

In December, 1917, Osborn (1918.468) presented before the Palseontological Society of America his poly-
phyletic theory of the Proboscidea, a theory more or less fully anticipated by previous authors but more radical

935
936 OSBURN: THEPROBOSCIDEA

in its inclusion of numerous branches. These branches of the Elephantidae, as extended in successive years

between 1910 and 1921, are as follows:

Elephantine Osborn, 1910, p. 558. 'I'lie true elephants of Asia.

StegodontinvE' Osborn, 1918, pp. 135, 136. Primitive elephants of southern Asia.
LoxoDONTiNE Osborn, 1918, pp. 135, 136. The African, Eurasiatic, and insular loxodonts.
EuELEPHANTiNE- Osbom, 1918, p. 136. The mammoths of Eurasia and North America.
Mammontin* Osborn, 1921, p. 1, established "because the genus Eitelephas is invalid; the term Mammontinx
(i.e., les mammonts, the mammoths) may be substituted."

According to this foin-fold subfamily classification of 1918-1921, the synoptic arrangement of these branches
is as follows:
[Subfamily: Stegodontine,' includinK the Stegoloijhodonts^ antl true
ELEPIIANT01D1'1\, elephant-like probu- Stegodonts, from primitive member.s of which the elephant sub-
scideans. I
families may have branched off.
[Subfamily: Mammontine or mammoths,
united by very distinctive
Elephantide, the elephant-like probe- |
separeLtedhy several types of dental characters.
cranial characters;
scideans and their ancestors, in which j Subfamily: LoxoDONTiNyE, typified by the African elephants {Loxodon-
the ridge-crested grinders gradually I and the numerous living subspecies), by Pa/a?o?o.rodon
/a a/r/ra/io
transform into ridge-plated grind- antiquus* and P. nntnadicus, of southern Asia, and by the dwarfed
ers, the lower incisive tusks abort, elephants of the Mediterranean islands.
and the upper incisive tusks attain Subfamily: Elephantine, typified by Elephas indicus {majcimus),
\ery great size. such as the insular types of Ceylon and the continental types of
southern Asia.

We may now recall the first conception by the present author of the diversities which jjrevail in the subfamily
Manuiiontinse, by quoting from the author's paper (1921.515) on the "Evolution, Phylogeny, and Classification
of the Proboscidea," pages 14 and 15:

The MammontinsB. The Mammoths

a striking fact that the oldest geologic appearance of a member of the true Elephantoidea is the Elephas planifrons
It is
occurring in the Pinjor horizon. Upper Siwaliks, Middle to Upjier Pliocene, India. All the fauna of the great Siwalik deposits
underlying this geologic level, according to Pilgrim, contain only Stegodonts, Longirostrines, and Brevirostrines. This is signifi-
cant of a north Eurasiatic center of adaptive radiation of both the Mammontinse and the Elephantine. The chief distinction
between these two subfamilies lies in the flattened forehead of the Mammoths, to which the specific name planifrons refers, a
forehead which becomes increasingly concave and compressed anteroposteriorly until it reaches the high, narrow peak of
E. imperator.
.\gain, the succession of species is probably polyphyletic, awaiting analysis. In descending order the main geologic succes-
sion is as follows:

[Heferencc in Present Memoir

Elephas primigenius Bluiiienbach. Northern Eurasia and North Americ'a, ri)per Pleistocene = Mammonteiis primigenins
" columbi Falconer. Middle Pleistocene, North America = Parelephascolinnbi
" imperator Lower Pleistocene, North America
IjC'iAj. = Archidiskodon imperator
" trogontherii Pohlig. Lower Plei.stocene, Europe = Parelephas trogontherii
" hysudricus Falconer. Uppermost Pliocene, India
^'^^
= llypselephas hysndricus
" meridionalis Nesli. Upper I'liocenc, Val d'Arno, Italy =ArchidiskodoN mcridionalis
" planifrons Falconer. Pinjor horizon. Middle to Upper Pliocene, India,
also Austria and Bessarabia (Russia) = Archidiskodon planifrons]

The position of E. hysuilricus in this phylum is doubtful. The cranium referred to this species by Falconer is not of the
I'''

mannnontine type. In 1913 Pilgrim traced ba('k E. planifrons to the Upper Miocene [Middle Pliocene] Si.egodon cautleyi, but it
would appear at jjresent that none of the known Stegodonts gave rise to the Mammoths. lOxtreme cranial abbreviation, hyi)er-
brachycephaly, and acroco))haly are great characteristics of all the phyla in this subfamily (excepting possibly that to which E.

'[Removed by Prof('Ssor O.sborn, Vol. I, p. 22, of tli(^ pn'.scut Memoir to hi.s new .superfamiiy .Stegodoiitoidea. Kditor.]
'-[Subfamily Mammontinae .sub.stituted (see Csborn, 1921.515, p. 1). — Editor.)
'[Separated by Professor Osborn from the true Stegodonts under the new subfamily name Stegolophodnntinir and placed in the superfamily Mastodont-
oidea, family Mastodontida- (see Vol. I, p. 700). — Editor.]
'[Members of the 'Elephas' anliquus group subsequently regarded by Professor Osborn as belonging to his new genus llesperoloxodon (see Osborn, 1931. S46,
p. 21).— Editor.)
^[Included in the subfamily Elephantina; of the pre.sent Memoir. — Editor.)
 THE MAMMONTIN^: ARCHIDISKODON AND MI<:TAK('11IDISK0D0N 937

hjj.'^iidriciis belongs). There

is a wide range of divergence in the tliickness and MiultijiHcation of the hinielliB of the grinders.

Klephm iinperator may

be derived from the A,', laeridionaliti type, with very few lamellae, composed of thick enamel bands and
with a great coating of cement, or from the E. pUintfrom Falconer type. The E. primigeniti.s phylum presents the highest lamel-
lar formula known, with relatively little cement; this phylum is also tlistinguished by the loss of a digit in the pes, becoming
tetradactyl, a uniciue character among proboscideans. Very great shoulder height, estimated at thirteen feet, is attained by
K. imperaUir in the favorable environment of the southern United .States and Mexico, as compared with the height of nine feet
six inches attained by E. primigenius in the frigid north.

Superfamily: ELEPHANTOIDEA Osborn, 1921

Family: ELEPHANTID^ Gray, 1821

Subfamily: M AMMONTI Ny€ Osborn, 1921

Original reference: Amer. Mus. Novitates, No. 1, pp. 1 and 14 (Osborn, 1921.515).
Includes Archidiskodontinae and Parelephantinaj of Dietrich, 1927, p. 313 (in part).

—
Subfamily Characters. United by the common cranial characters of Archidiskodon, Par-
elephas,and Mammonteua, namely, cyrtocephaly, bathycephaly, hypsicei^haly, acrocejihaly, attain-
ing extreme cyrtocei)haly and acrocephaly in Mammonteus. Incisive tusk alveoli vertical, tusks elon-
gated, inciu'ved, crossing each other in old males, not serving in feeding habits. Grinding teeth pro-
gressive from a primitive, subhypsodont stage (Archidiakodon) to an excessively progressive, hyper-
hypsodont stage {Mammonteus). Habits and geographic distribution varying widely in the three generic
phyla.

The cranial unity of the Mammontinsp and the profound distinctions both from the Loxodontn nfricann and
Elephas indicus types are fully discussed in Chapter XV and clearly set forth in the diagrams (Figs. 805, 806, and

L-/00 /^-/36'

LOXODONTINE, ELEPHANTINE, AND MaMMONTINE MiD-CRANIAL AxES

Compare with figures 805 and 806 of Chapter XV
Fig. 816. The liorizontal plane of the worn siirfaoe of the .superior
grinders correspond.s with the hne from tlir niid-condyle to the anterior
nare.s in Elephas indicus.

Luxodonta africana, the most primitive, occipital

pUme (L) 100°. = Loxodontin.e

Elephas indicus, intermediate, witli rounded cranial

section, occipital plane (E) 121°. = Elephantin.b

Mammtmieus primigenius, the most progressive,

liypsicephalic, acrocephalic, bathycephalic, oc(i|)ital
jilane (M) 136\ =Mammontin.e
lo72fitudinal
Condyle to
midaie o/ right half of parietals
Clnierior'\

816); 816 represents sagittal sections taken through the center of the right half of the cranium. The
figure

sections reveal a most surprising resemblance between the three genera of the Mammontins to each other and
equally surprising contrasts between the mammontine and the Indian and African elephant crania. A summary
of these characters, quoted from page 932 above, is as follows:

(1) Subfamily Mammontine, including the three generic phyla Archidiskodon, Parelephas, and Mammonteus, which
exhibit three distinct modes of adaptation in the grinding teeth.
(2) Tusks and tusk alveoli progres.sively vertical; tusks elongate, incurved, crossing in old males, not serving in feeding
habits.
938 OSBORN: THE PROBOSCIDEA

(3) Grinding teeth subhypsodont to extremely hypsodont, typically

broad and typically short vertically; progressively
deepening ridge-plates; typical M
3 ["^ in Archidiskodon planifrons, multiplying to M3
^J in Mammoiiteus primigenius com-
pressus. [Parelephas progressus has the following ridge-plate count: 3 j^. Editor.] M —
(4) Cranium shortened (cyrtocephalic), tleepened (bathycephalic), heightened (hypsicephalic), pointed (acrocephalic),
reaching an extreme fore-and-aft abbreviation and elevation in Mainmoideus.
(5) Habits varying in the three generic phyla. Originally browsing, progressive to extreme grazing type.

A PLANIFRONS Tfe/: Q A. MERIDI0NALI5 7y/>e f S. PINJORENSIS 7y/>e n A, IMPETRATOR -Ref

Florence Mwz
^ Amer Mui /9772 Amer Mils. .'
British Mu^.
14476

1/20 Natural

Fig. 817. Comparative profiles of riaiiia of Archidiskodon {A, B, D) ami oraiiiuni of Slegmhn (C), all one-twpiitieth natural size.
A, Archidiskodim plnnifrons ref. (Brit. Mus. M.30G0). C, Hiegodon pinjorensis tyi)t' (Amer. Mas. 19772).
B, Archidiakodon meridionalis type (Florence Mils.). D, Archidiskodon imperalor ref. (Amer. Mils. 14476).
Observe in A, B, D the progres.-iive increase in size, abbreviation, heightening, and deepening of the cranium, heightening of the occipital crest, and con-
cavity of the lengthening forehead. Contrast with the still more abbreviated, hypsice])halfc cranium of .S'/c3(«/o7t pjVy'ocen.sn's (C).

While united by these profound cranial characters, the three genera, on the contrary, are widely distinguished
by the specialization of their grinding teeth. It seems hazardous, therefore, to unite them in a single subfamily.

The outstaiuling generic distinctions are as follows:

Archidiskodon Pohlig, typified by Parelephas Osborn, typified by Ele- Mammonteus Camper-Osborn, typi-
ElephuK meridionalis and E. planifrons. phas jeffersonii. fied by Elephas primigenius.
Arrhidiskoilon tusks and craniiun as Parelephas tusk and cranial charac- Mammonleus crania as in other
in other Maininontinae. Primitive ters as in other Mammontina;; cranium Mamniontina', but excessively acro-
grinding teeth, subloxodont, subhypso- less cyrtocephalic than in Archidisko- cephalic, hypsicephalic, bathycephalic,
dont to hyp.sodont. Ridge-plates ex- don or Mammonteus. Convergent with and cyrtocephalic. CJrinders with ex-
tremely broad plated, laminate, slowly the P>lephantinse in bathycephaly and treme fore-and-aft compression; ridge-
progressing in number from 3 xt+ M multiplication of ridge-plates; grind- ])lates ])rogre.ssively deepening and
(.4. planifrons) toj^'.h^ {A. imperalor). ers re.sembling those of Elephas indicus; broadening, the most elevated and at-
l']nainel thick. ridge-plates relatively fine, progres- tenuated known, multiplying from
sively hypsodont, multiplying from P. M 3 I gin*} {M. astensis), to f}- (M. M3
trogontherii (M 3 H|), to P. jeffer- primigenius), to M
3 fy {M. primi-
sonii (M 3 §i), to Parelephas progressus genius compressu.'i). Enamel thin.
(M30i). Enamel medium.
Mammoths of the south temj)erate Mammoths of the mid-temperate re- Mammoths of circumpolar regions;
regions; chiefly browsing. gions of Eurasia and North America; habits chiefly grazing, browsing in un-
browsing and grazing habits, adapta- fiivorable seasons.
tions similar to Elephas indicus.

Progressive in size to the gigantic Intermediate in size, progressive to Dwarfed in size, with digits in pes
Archidi.tkodon imperalor maibeni. Parelephas floridanus. reduced to four.
 THE MAMMONTIN.E: ARCHIDISKODON AND METARCHIDISKODON 939

2. HISTORY OF THE GENUS ARCHIDISKODON

Although long united with Elephas, the species which are grouped in this genus constitute a very distinct
generic phylum for which Pohlig's name Archidiskodon, referring to the archaic molar ridge-plates, seems very
appropriate. As shown in our "Classification of the Elephantoidea" (Chapter XV) %-nxit. siie
above, also in figures 805, 806, 816, and 817, Archidiskodon is profoundly related
in its cranial characters to Mammonteus and to Parelephas; consequently it belongs
in the subfamily Mammontinae, as defined on page 937 of this Memoir.

A. IMPERATOR Ref. M. PRfMIQENIUS Ref. P. JEFFERSONII Type P WASHrNQTONII

Ids AnqeZes A^u-S. A/ar. Mils Bsao Amer. Mus. 99S0 Arner. Mui- BC^

M. PRIMIQENIUS
COMPRESSUS

'y^o natural sije

Fig. 818. Hypsiceph.^lic Cbani.\ of the Mammontin«

A, Archidiskodon imperator, juvenile male cranium, from Rancho La Brea, California.
B, Mammonteus primigenius, young adult male cranium, from Siberia.
C, Parelephas jeffersonii, type cranium, aged male, from Indiana.
D, Parelephas washingtonii, adult male cranium, from tlie state of Washington.
Observe (1) the strong fore-and-aft cranial compres.sion and vertical elevation of the occiput (hypsicephaly);
(2) the slightly concave forehead; (3) the very deep depression of the mandibular rami (bathycephaly); (4) the
sharp downward flexure of the basicranial axis (cyptocephaly); (5) the disimrity in size of the M. primigenius
skull and jaws.

Fig. 819. Comparative Series of Superior Molars showing evolution of the hidgb in the
Elephantoidea and Stegodontoidea. One-sixth natural size.
E, Mammonteus primigenius compressus, tyi)e M^, showing extreme comjjression of 27 ridge-plates.
D, Archidiskodon planifrons, type M', with 9-10 ridge-plates.
C, Stegodon aurorie, type M", with 10 -|- ridge-crests (cf. .S'. airawana, Fig. 764C).
B, Stegodon ganesa, type M^, with 10 -|- ridge-crests.
A, Stegodon insignis, type M^, with 10 ridge-crests.
(cf. p. 892) "is either a liighly progressive Stegodon or a primitive Archidiskodon, a
Stegodon aurora; |«)int to be
determined positively by the discovery of a cranium."

Fig. 819

PoHLiG, 1885, 1888. —To our knowledge Pohlig was the first to separate the elephants with archetypal ridge-
plates as Archidiskodonten (1885, p. 1027): "8. Ich theile die Elephanten nach Kronenformen und Lamellen-
zahlen der Molaren ein in Archidiskodonten {E. planifrons, E. meridionalis) , Loxodonten {E. africanus, fE. anti-
quus) und Polydiskodonten {E. primigenius, E. indicus etc.), die Stegodonten mit Clift wieder zu Mastodon
zahlend." Three years later PoWig (1888, p. 138) defined the genus as follows: "1. Archidiskodonten. Typus:
E. meridionalis. Uebergang zu der folgenden Gruppe bildet E. planifrons. Tapinodiske, laticoronate, kurze und
 AMERICAN MUSEUM OF NATURAL HISTORY
ERWIN J RAISZ A H 1827

Central Region (see black squahb) of the Siwalik Hills, 200 milks .south
and north ok Simla India
Tertiary rauKe. 800 miles in lengtli, on the southwestern
base of the Himalaya Mts., arul northeast of the Punjal, Classic '

Sewalik"
.xnn.fr"'' T^\'^ ^v^"'^
""" "'^,"' .''''"''' ''"' ''"^'' "'^''^-^^ '"«'-"^-^"'' ''-^ '''<"'• ''"'''> '' C-"".v in .827, explore, l.v Dr Hugh Falconer
uTsai and detlh "T ""T t"'.''' k
XXV, inap of India,

940
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 941

pachyganale Molaren. Parsilamellat. (Meist nur 15 Lamellen an M. III.)" In the same Memoir (p. 252) he
introduced the generic terms: 4^ Archidiskodon 5. Polydiskodon 4''- Loxo(-disko-)don. From these citations
we have abstracted the above generic characters.

ScHLESiNGER, 1912, 1916. — Schlesinger (1912, pp. 98, 99, footnote) confirmed this generic description and
amphfied PohUg's definition as follows: "Ausdriicke halte ich fUr einen MissgrifT und gebe ihre Erklarung: Tapino-
disk = mit niedrigen Jochen (Gegensatz = hypselodisk), latikoronat = breitkronig (Gegensatz = angustikoronat),
pachyganal = mit dickem Schmelzblech (Gegensatz = endioganal), parsilamellat = mit wenigen Lamellen (Gegen-
satz = densilamellat)." In a later paper (1916,
Fig. 821. Chief Lower and Upper Pleistocene localities of western Eurasia in wliieh
pp. 102, 103) Schlesinger cites the term as a
occur species of Archidiskodon, Parelephas, Mammonteus, Loxodonta, and Pal^oloxodon.
subgenus: "Elephas {Archidiscodon) meridion- After Osborn, 1910..346, p. 391, fig. 176. See more detailed caption, figure 932, below.

alis Nesti," and (op. cit., pp. 112, ll3)"Elephas

(Archidiscodon) planifrons Falc." Schlesinger
also in his "Studien iiber die Stammesge-
schichte der Proboscidier" of 1912 recognized
a resemblance to the broad-plated American
species Elephas columbi and E. imperator, but
failed to separate these species into a distinct

generic phylum.

Osborn (1922.555) distinguished Elephas

columbi from E. imperator (p. 3) as follows:

"We thus find by the characters of the type

and neotype specimens that the real Elephas
columbi is not the animal we have been describ-
ing under this name; it is a dwarf form, per-
haps a dwarf female, of the animal which we
PLEISTOCENE —
EUROPE. 1 Forest Bed of Cromer (Norfolk). Sables de 2
have been describing under the name Elephas St. Prest near ChartresEure-et-Loire)
( . 3 Malbaitu (Puy-de-Dome). 4 Peyrotles (Bouchea-
du-Rhone). 5 Solhilac near Puy. Clay deposits of 6 Durfort (Card). 7 Cajarc (Lot-et-Ga-
ronne). 8 Val d'Arno (Tuscany). 9 Leffe near Bergamo (Lombardy). 10 Riidorf near Pots-
imperator."^
dam (Brandenburg). Gravels of 11 Sussenbom near Weimar. Sands of 12 Mosbach in
northern Baden. Freshwater deposits of 13 Clacton (Essex). Sands of Mauer near 14 Hei'
He clearly defined (op. cit., pp. 3 and 4) delberg (western Germany). 15 Chelles on the Mame, near Paris. 16 iS(. Acheul (Somme).
17 Ilford and Grays Thurrock (Essex). Lignites of 18 Dilmlen and of Utznach, near Zurich.
the characters of the grinding teeth of E. im- 19 Taubach near Weimar. 20 U'iWfcr'rcWi caiic on il/on( Sanfis (eastern Switzerland). Tuffs
of 21 the Tiber Valley, near Rome. Caves of 22 Neandertal, near Diisseldorf (western Ger-
perator, concluding (p. 5): "The cranial char- many), 23 Spy, near Amur (Belgium), 23a Krapina (Croatia), 24 Chapelle-aux~Sa ints (Cor-
r^ze). Caves and alluvial deposits of 25 Temifine (or PaXikao) near Gran (Algeria), 26 Pointe
acters observed in three more or less complete Pescade, near Algiers (Algeria). 27 Prince's Cave (Monaco). Sandy clays of 28 Vdklinshofen
(Alsace). 29 Saalfeld (Saxe-Meiningen). Travertines, etc., of 30 Gera, Jena (Saxe- Weimar).
skulls referred to Elephas imperator tend to 31 Leipzig (Saxony) .32 Solutre, north of Lyons. Loess of 33 Wtirzburff (Bavaria). 3i Thiede
near Braunschweig (Prussia). Cave of 35 Montmaurin (Haute-Garonne). 36 Chdteauneuf-
support the direct descent of this animal from sur-Chartnte (Charente). Caves of 37 Schweizersbild near Schaffhausen, and Kesslerlock near
W
Thayngen (northern Switzerland). Remains of lake dwellings at 38 auwyl (hucerne) 39 Ro' ,

the E. meridionalis of the Val d'Arno, Upper benhausen, south of Lake Pfaffikon. 40 Concise on Lake Neuchatel (Switzerland). Peatbogs of
41 Hassleben, near Weimar. Travertines of 42 Langensalza (Erfurt) in central Germany,
Pliocene of Italy." Caves of the 43 Island of Malta, 44 Island of Crete, 45 Island of Cyprus.

Subsequently Osborn (1924.633, p. 2) confirmed Pohlig: "We therefore confirm Pohlig's separation of the

southern mammoths Elephas planifrons, E. meridionalis, and E. imperator into the distinct generic phylum
Archidiskodon,'" and in the year 1925 (Osborn, 1925.662) he placed Archidiskodon among the Mammontinae,
as shown in the diagram of that year (Vol. I, Fig. 7, of the present Memoir), concluding (p. 28) with a redefinition
of the family Elephantidae and Race XIII, as follows:

'[Subsequently (see pp. 997, 1001 )Professor Osborn separated Elephas imperator and E. columbi, assigning the former to tlie genus Archidiskodon Pohlig
and the latter to Parelephas Osborn. —
Editor.]
 —

942 OSBORN: THE PROBOSCIDEA

Family: ELEPHANTID.'E, distinguished by plated grinding teeth developing out of the more or less closely compressed,
serrated ridges of Stegodon^ into the broadly plateil grinders of Archidiskodon. the lozenge-shaped
grinders of Loxodonla, antl the compressed, finely plated grinders of Parelephas, of Mammonteus,
and of Elephas the type genus of the family. . . .

Race XIII. The Southern Mammoths, or Archidiskodonts. Excessively broad-plated grinders with
abundant cement; first known in India, migrating westward into southern Europe, eastward into
America, where arriving in late Pliocene or early Pleistocene time they finally gave rise to the Im-
perial Mammoth, Archidiskodon imperator, the last of its race.

Fig. 822. Geographic distriljution of the principal .speoie.s of Arcliitiiskodonts. The white dots within tlie black areas represent tlie approximate locaHties
where the twenty-two species were discovered; these dots each carry a number in a circle representing the chronologi(; sequence of type de-
tyjies of the.se
scription. The white crosses represent some of the principal referred specimens mentioned in the present Memoir.

3. ORDER OF DISCOVERY AND DESCRIPTION OF TWENTY-TWO SPECIES

OF ARCHIDISKODONTS
See Figure 822
Reference in Present Memoir
1. 1825 E/ep/ifls 7nmrf{ona/7s Nesti, Val d'Arno, Italy = Archidiskodon meridionalis
2. 1846 [18451 Elepfuis plunifrons Falconer and ( 'autley, Siwalik Hills, northern India = A rch idiskodon pla n ifran s
1. 1855 E. giganlcii.s Ayiiiard (MS., also in l"'alconer, 1857, p. 321), France NouKUi nudum = Archidiskodon
meridianalisC!)
[1857-1868 Elephas columbi Falconer, CJeorgia = Parelephas columbi (see Chap. XVII)]
3. 1858 Elephas imperator Leidy, ?Seneca, Thomas Co., Nebraska {fide Hay, 1924,
p. 100)-^ = Archidiskodon imperator
[1859-1861 Elephas teaianus Owen, 1859, Blake, 1861, Texas = Parelephas columbi (see Chap. XVII)]
1. 1889-1890 Elephas lyrodon Weithofer, Val d'Arno, Italy = Archidiskodon meridionalis (female?)
[1890 Elephas mindaneiisis Naumann, Philippine Islands = Siegodon (Archidiskodon'.') inindanensis
(.see C:hap. XIV)]
4. 1915 Elephas hayi Barbour, Crete, Saline Co., Nebraska = Archidiskodon hayi
[1922 El. [Elephas] Columbi var. Felicis Freudenberg, Mexico = Parelephas columbi felicis (see Chap.
XVII)]
5. 1922 El. [Elepha.s]Columbi var. .vlveslris Freudenberg, Ejutla, State of
Oaxaca, Mexico = Archidiskodon imperator silvestris

'(See Chapter XIV, pp. 800, 807, where Professor Osborn's final views regarding tin iiiplcte separation of the Stegodontoidea from the Elephantoidea
are given. — Editor.]
-(Exact locality not cited. Lugn and Schultz (193t.l, p. 373) give Pawnee Loup Branch of Platte Hiver= Middle Loup, probably Hooker County.
Editor.l
 THE MAMMONTIN.^: ARCHIDISKODON AND METARCHIDISKODON 943

6. 1922 El. [Elephas] Columbi var. Falconeri Freudenberg, Tequixquiac,

Valley of Mexico
7. 1922 Loxodonta griqua Haughton, CJriqualand West, Transvaal, S. Africa
3. 1922 Elephas Columbi var. imperator Freudenberg, Spokam Bar, near
Helena, Montana
8. 1923 Elephas meridionalis, mutation cromerensis Deperet and Mayet,
Kessingland, Suffolk, England
9. 1924 Elephas antiquus rumanus S. Stefanescu, Tulucesti (Covurlui), Rumania
10. 1925 Elephas scotti Barbour, near Staplehurst, Seward Co., Nebraska

11. 1925 Elephas tnaiheni Barbour, near Curtis, Lincoln Co., Nebraska
[1927 Archidiskodon transvaalensis Dart, Lowest Terrace, Vaal River, S. Africa

[1927 Archidiskodon sheppardi Dart, Lowest Terrace, Vaal River, S. Africa

2. Leith-Adamsia siwalikiensis Matsumoto, Siwalik Hills, India

Elephas haroldcooki Hay, Frederick, Oklahoma
Elephas exilis Stock and Furlong, Santa Rosa Island, California
Archidiskodon subplanifrons Osborn, Sydney-on-Vaal, S. Africa
Archidiskodon broomi Osborn, near Kimberley, S. Africa
Archidiskodon sonoriensis Osborn, near Arizpe, northern Sonora, Mexico
Lox. (Pal.) Tokunagai junior mut. Matsumoto

Archidiskodon vanalpheni Dart, Sydney-on-Vaal, S. Africa

Archidiskodon millelti Dart, Sydney-on-Vaal, S. Africa

Archidiskodon loxodontoides Dart, Sydney-on-Vaal, S. Africa

Archidiskodon yorki Dart, near Christiana, S. Africa

Archidiskodon andrewsi Dart, ?Middle Terrace, Vaal River, S. Africa

[1929 Archidiskodon hanekomi Dart, Vaal River, S. Africa

21. 1932 Archidiskodon meridionalis nebrascensis Osborn, Angus, Nuckolls Co.,

Nebraska

22. 1934 Archidiskodon proplanifrons Osborn, Cong-Gong, near the Vaal River,
S. Africa
944 OSBORN: THE PROBOSCIDEA

In 1922 Freudenberg reviewed the elephants of Mexico and described foiu' subspecies of Elephas columbi
namely, El. Columbi var. Felicis, El. Columbi var. .s//res^m, El. Columbi var. Falconeri, and El. Columbi var.
imperator; the first {El. Columbi var. Felicis) we regard as more closely related to Falconer's species Elephas

columbi, the remaining three to Leidy's species Elephas imperator.

In 1923 Deperet and Mayet devoted a volume to their thorough researches of the species of eastern Eurasia,
Elephas planifrons and E. meridionalis, and added the Lower Pleistocene subspecies Elephas meridionalis, muta-
tion cromerensis, of Kessingland, Suffolk, England.

The wide geographic distribution of Archidiskodon is indicated by Stefanescu's description in 1924 of Elephas
antiquus rumanus of Rumania, by the discovery of two American species, Elephas scotti and E. maibeni,
in Nebraska, described by Barbour in the year 1925, and
by the most surprising and welcome discovery
finally

of all, the Archidiskodon subplanifrons and A. broomi in the Vaal River diggings of South Africa, described by

the present author in 1928, as well as Dart's new species, Archidiskodon transvaalenMs and .4. sheppardi from
the Vaal River gravel terraces, described in 1927.^

5. NEW ARCHIDISKODONTS AND LOXODONTS OF AFRICA

After the present author had described and figured, but not published,'- the two names above {Archidiskodon
subplanifrons and A. broomi), he received Prof. Raymond A. Dart's "Mammoths and Man in the Transvaal,"

Nature, December 10, 1927, with the welcome figures [Fig. 823 of the present Memoir] and description of the Vaal
River terraces at Windsorton and Bloemhof, of which the following is an abstract:

The Vaal River valley, near Bloemhof, in the southwestern Transvaal, belongs to a great watershed which has yielded
Australopithecus africanus Dart, 1925 [from Taungs district, Bechuanaland], and the Boskop man, besides stone implements,
rock engravings, and other evidences of primitive man. The watershed includes three gravel deposits of different geologic age
(Du Toit, Ann. Rept. (Jeol. C'omm., 1906) situated on a number of terraces.
Upper.—The highest and presumably the oldest [Pliocene?] terrace has an altitude of some 200 to 300 feet above the river
exposed at a distance of to 6 miles. No
3}i have been described from
fossils hitherto most ancient (Dart, op.
this terrace cit., p. 1).

[This possibly the level at

is which Archidiskodon subplanifrons Osborn, a very primitive form, was found — Fig. 875 of the pres-
ent Memoir. — Osborn.]

Middle. — The middle or second [Pleistocene or Mastodon] gravel terrace on a lower level at several points west of Barkly
is

West; from this terrace numerous palaeolithic implements have been described (Hodkinson, 1926); also (Beck, 1906) a frag-
mentary tooth of Mastodon {Bunolophodon) sp. Felix, at Waldeck's Plant, 60 to 80 feet above the river bed. Haughton described
—
Loxodonta griqua = Metarchidiskodon see figure 882 of the present Memoir], also a new giraffe {Griquatherium cinyulntiim)
[

from this 60 to 80 foot terrace. [Doctor Broom holds that the mammoth teeth are washed in from an older geologic deposit
(see Nature, March 3, 1928, p. 324) and are not truly associated with the flint implements, in geologic age.]

—
LowEH. A still lower gravel deposit [Pleistocene], near the level of the present river Vaal, contains Equus and Hippo-
potamus amphibius var. robuslus. From this low river bed gravel, \Yi miles below The Bend on the Vaal River, Haughton
determined as Elephas (cf. antiquus) a portion of a tooth recovered at a de])th of 5 feet, but which Dart regarded {op. cit., p. 42)
as resembling E. antiquus recki Dietrich. [Apparently this same specimen is now described by (Jsborn as Archidiskodon broomi.
the label of which bears the inscription, "3682 Mus. Kimb. The Bend. II. Else." Fig. 877 of the present Memoir.] From this
level are recently recorded Archidiskodon Iranspaalensi.s Dart and A. sheppardi Dart.l'l

^[Archidiskodon transvaalensis and .1. (.see Osborn, 1934.92."), pp. 2iinil 11)
s/icppanij prove to belong to the genus Palipoloxodon Since this section of the
Memoir was have been (iescribeii by Professor Dart (1929) and one by Professor Osborn (1931); jilso two
written, four species from South Africa
.species from North America have been described by Professor Osborn, one from Mexico (1929) and one from Nebraska (1932). A <'omplete list of the
species of Archidiskodon is given on pages 942 and 943 above. Editor.] —
^Subsequently published in Nature (Osborn, 1928.749, pp. 672, (173) under the title "Mammoths and Man in the Transvaal."

J
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 945

Gr^vcu te:rf*.acc Go - so feet

/ rlvcr bcd c.ravel terrace

DiA li-^SE

The Vaal River Gravel Terraces, South

Africa, the Scene of the Discovery of
Mammoths and Man in the Transvaal
After Dart, 1927
These terraces (upper at Windsorton,
Fig. 823.
middle and lower at Blocmliof) have yielded the
types of Boskop man (Homo capensis), of Aus-
tralopithecus africanus, of Loxodonta [
= Metarcki-
diskodon] griqua Haughton, of Archidiskodon
[
= Pal^oloxodon] transvaalensis and .4. [=P.\
sheppardi of Dart, and of A. subplanifrons and
A. broomi of Osborn, also of Mastodon {Buno-
hphodon) sp. Felix. After Dart, 1927, figs. 1 to 3.

Transvaal

Old Riven bed dicoincs Sand House oh island New BlutR BED
', Gravel bed or
, UNKNOWN DEPTH

Consequently a summary of the discoveries and descriptions of South African specimens (1928) is as follows:

Original Description Reference in Present Memoir

Loxodonta griqua Haughton, 1922, type, Griqualand West, Transvaal = M elarchidiskodon griqua
Elephas (Lo.rorfo«) zulu Scott, 1907, type, Zululand - Loxodonta zulu
Elephas zulu, referred by Hopwood, 1926, Kaiso Bone-beds, near Lake Albert - Loxodonta zulu
Elephas antiquus Necki Dietrich, 1916, type, Tanganyika Territory = Palseoloxodon recki
Elephas aff. meridionalts Nesti, 1825, referred by Hopwood, 1926, from Kaiso
Bone-beds, near Lake Albert - M elarchidiskodon griqua [or A. planifronsf)
Archidiskodon transvaalensis Dart, 1927, type, middle Vaal River gravel terrace - Palxoloxodon transvaalensis
Archidiskodon sheppardi Dart, 1927, type, lowest Vaal River gravel terrace - Palseoloxodon sheppardi
Archidiskodon subplnnifrons Osborn, 1928, type, middle(?) Vaal River gravel
terrace --
Archidiskodon subplanifrons
Archidiskodon broomi Osborn, 1928, type, middlc(?) Vaal River gravel terrace -Archidiskodon broomi
Mastodon (Bunolophodon) sp. Felix, referred by Beck, 1906, Waldeck's Plant --
Trilophodon{'!) sp. indet.

The above species of Archidiskodon from Africa are fully described below, following the description of the
Eurasiatic Archidiskodonts.
946 OSBORN: THE PROBOSCIDEA

6. APPROXIMATE ASCENDING PHYLOGENETIC ORDER OF SUCCESSION OF

SPECIES OF ARCHIDISKODON AND PARELEPHAS (1928)
Only an approximate phylogenetic order of succession of the species of Archidiskodon may be given at the
present time. In this phylogenetic list we may inchide with some certainty fifteen of the better-known species
of Archidiskodon and Parelephas, placed in ascending order as follows:

ARCHIDISKODON AND PARELEPHAS OF NORTH AMERICA AND EURASIA

listocene? level
 THE IVIAMMONTIN.^: ARCHIDISKODON AND METARGHIDISKODON 947

Family Group of Archidiskgdon imperator maibeni along the Platte River, Nebraska
Restoration by Margret Flinsch Buba in 1935, under the direction of Henry Fairfield Osborn
Bull in foreground l/40th natural size and bull at extreme left l/140th natural size. Cow and calf in middle foreground l/60th natural size

Fig. 824. The bull in the foreground and the one in the distance at the were based on mea.surements of the skeleton of Archidiskodon
extreme left

imperator tnaibeni in the Nebraska State Museum (Neb. Mus. 5-9-22), from Lincoln County, Nebraska (.see Figs. 910-912, 918 of the present Memoir),
supplemented by a fine .skull of A. imperator in the Geological Institute of Mexico (No. 212) from Tepexpan (see Fig. 902) and of a giant tusk of the same species
(.1. imperator) in the American Museum (Amcr. Mus. 22481), from Post, Texas (see Fig. 894). The height at the withers (toj) of shoulder blade) is 3826
mm.; adding the usual 6/3'yc gives a height in the flesh of 4068 mm. or 13 ft. 4% in.

II. CHARACTERS OF ARCHIDISKODON AND METARCHIDISKODON AND INCLUDED SPECIES

Superfamily: ELEPHANTOIDEA Osborn, 1921
Family: ELEPHANTID^ Gray, 1821
Subfamily: Mammontin^ Osborn, 1921

Genus: ARCHIDISKODON Pohlig, 1885, 1888

Original reference: Zeitschr. deutsch. geol. Ges., XXXVII, Heft IV, p. 1027 (Pohlig, 1885.1); Xova Acta Leop. Carol., LIII, No. 1,

pp. 138, 252 (Pohlig, 1888).

GenotjTDic species: Elephas meridionalis and E. planifrons.
Syn.: Parastegodon Matsumoto, 1924 (in part); genotype, Elephas aurorx Matsumoto, 1924. Leith-Adamsia Matsumoto, 1927;
genotype, Leith-Adamsia Siwalikiensis Matsumoto, 1927.

Generic Characters (translation after Pohlig, 1888, p. 138). Genotypes E. meridionalis, —

E. planifrons; transitional to higher groups. Molars with few ridge-plates, not exceeding fifteen [?]
lamellae; crowns broad, short, and with thick enamel (i.e., "laticoronate, kurze und pachyganale
Molaren") ridge-plates tapering to summit in sections.
;
 : I

948 OSBORN: THE PROBOSCIDEA

—
Generic Characters (Osborn, 1928). Superior tusks large, incurved, crossing in old males
as in other Mammontinse. Cranium extremely heightened (hypsicephalic), foreshortened and broadened
(brachycephalic), deepened (bathycephalic). Molar ridge-plates extremely broad, enamel border
thickened, cement usually very thick. Ridge-plate formula slowly progressing from minimum,
(^4. plamfrons), to maximum, 3 M ——
{A. imperator). Finally attaining gigantic size.
3 M ^
This genus is given first consideration in the history of the subfamily Mammon tina^ for the reason that

Elephas = Archidiskodon]
[ planifrons of the Upper Pliocene of India is geologically the earliest of the Elephantidse

at present known. It is a very striking circumstance that until we reach the Upper Pliocene, Pinjor horizon, of

India we discover Mastodonts and Stegodonts only, there being no trace of the Elephantidae; consequently
Archidiskodon planifrons appears to be a new arrival in southern Eurasia in Upper Pliocene time, probably a de-
scendant of ancestors coming from some region of southern Africa, such as Archidiskodon subplanifrons.^

Osborn, 1929: Comparing Falconer's measurements of thirty-nine Siwalik specimens (Table VI, opposite page)
with Barnum Brown's measurements of twenty-seven Siwalik specimens (p. 954 below), we find a closely similar

range of 'ascending mutations' rising from the most primitive ridge formula, M 3
»H
through intermediate
mutations, M3f j'^j, to the most progressive mutations, M3j K-l 2-W
Thus in sixty-six specimens collected in
the Pinjor horizon, of the Siwaliks, India, we observe gradual ascending mutations of the Archidiskodon planifrons
stage toward the A. meridionalis stage. The ridge-plates throughout in M^ are more numerous than those in M'*.
Summary. —The ascending mutations in the thirty-nine sjiecimens referred by Falconer (1868) to 'Elephas
planifrons' are seen to compare very closely with those in the twenty-seven specimens of the collection made by
Barnum Brown and referred by Osborn to Archidiskodon planifrons. This renders it probable that the Falconer
collection and the Brown collection were from the same geologic horizon. The extremes in both collections are

seen in the maximum and minimum heights of the ridge-plates

Uppkr Pliocene Archidiskodon planifrons

Measurements of Ridge-plates in Millimeters
Falconer (1868) Brown (1922)
Max. Min. Max. Min.
Third superior molars, M'
Height 123 64 102 88e
I>cngth 279 247 254 201
Third inferior molars, M3
Height 115 77 135 53
I.cngtli 323 224 313 203

Allowing for individual differences in size due to sex and geologic level, and for probable differences in mode
of measurement, especially of the height of the ridge-plates, it appears probable that in both collections we have
to do with a series of ascending mutations some of which attain the size and ridge-plate height of Archidiskodon meri-
dionalis. It is noteworthy that in the A. planifrons r.M-, the length (221 mm.) and the height of the 7th ridge-
plate (71e mm.) agree closely with the 7th ridge-plate height of M- of ^4. planifrons from the Piltdown gravels of
Sussex, England (see Figs. 852, 853, 854).

Falconer observes that while the ridge-plates in A. meridionalis increase in nimiber they do not increase
correspondingly in height. In two of his figiu'cs reproduced in the j^resent Memoir (Fig. 862) the measurements are:

i'liiMirivK .\rciiii)Iskodon meridionalis [or .v. I'LAMFRON.s]

Length lircadtli Ifoiglil
Val d'Arno, r.Ms 10 in. = 255 mm.]
[ 3.4 in. = 87 mm. [ 5 in. I
= 126 mm.
Norwich C'rag, r.M.i 11.25 = 287 mm.]
[ 3.8 = 97 mm.[ 4.8 [
= 123 mm.
*[0r .irchidiskodon proplanifrotis, an o<iuall\- primitive species sub.sequently described by Professor Osborn from Gong-Gong, Vaal River (Osborn, 1934.925.
p. 10).—Editor.]
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 949

UPPER SIWALIKS MUTATIONS

TYPICAL ARCHIDISKODON PLANIFRONS: M 3 fJt, ASCENDING TO TYPICAL A. MERIDIONALIS: M 1 3-1 4
3 11-14

Table VI. Summary of Measurements of Thirty-nine Specimens referred by Falconer,

1868, TO Elephas = Archidiskodon] planifrons
[
 950 OSBORN: THE PROBOSCIDEA

1-. ARCHIDISKODONTS OF SOUTHERN EURASIA

From our present knowledge it appears probable that Archidiskodon originated in Africa, because seven stages
in the specific evolution of this genus have been discovered on that continent. In India Archidiskodon planifrons
suddenly appears in the Upper Pliocene, Pinjor horizon,' while /I. meridionalis appears in the uppermost Pliocene
and Lower Pleistocene The subspecific name Elephas = Archidiskodon] meridimialis cromeren-
of western Europe. [

sis has been applied by Deperet and Mayet to rather primitive archidiskodont molars found in the Forest Bed of
East Anglia.

SYSTEMATIC DESCRIPTION OF SPECIES

Archidiskodon planifrons Falconer and Cautley, 1846 [1845] In the collection of 1922 made by Harnuni Brown, twenty-
Figures 81.5, 817, 819, 825, 827-850, 8.52-856, 865, 871, 876, 914, seven or more specimens are now recorded as, or referred to,
1208, 1231, 1239, PI. xxi Archidiskodon planifrons; this collection (see map. Fig. 826) is
'Sewalik
Hills' of Falconer, Pinjor horizon, Upper
northern India; recorded by Brown e.specially from near the towns of Kalka,
Pliocene, Upper Siwaliks (Pilgrim)'; Upper Siwaliks, below Boulder Conglom- Charnian, Siswan, Chandigarh, and Mirzapur, as shown in the
erate, Lower Pleistocene (Brown).
complete li.st on page 955 below.
Syn. Leilh-Adamsia Siwalikiensis Matsumoto, 1927.
:

None
of the twenty-seven or more s])ccimens of the Brown
Specific Characters (Falconer, 1846, 1863, 1868; Osborn,
collectionwas actually found embedded in the rock; all of them
1928). —
Species distinguished from Elephas = Stegodou] auroras [

were loosely embedded in sand, occurring in gullies or depressions.

by the more elevated or hypsodont ridge-plates distinguished from ;

Consequently the exact geologic age and horizon levels are inde-
Elephas = Archidiskodon] meridionalis by the much less elevated
[

ridge-plates; by the flattened forehead to which the specific name

terminable; yet they may all be safely classed as L^pper Pliocene.'

planifrons refers in distinction from the concave forehead of A.

—
History. Falconer and ( 'autley first published in the "Fauna
Antiqua Sivalensis" the name Elephas planifrons in the legend of
meridionalis; also by the more primitive ridge-plate formula,
In 1846, Falconer wrote:
M 3 JY+, and by the succession of premolars, this being the last
Plate
"The next
II, figs. 5a, 56, of date 1845. p. 38,
serial modification in the disposition of the three dental
species of the Elephantidse in which the true premolars, P 3 and
substances, and in the consequent form of the teeth, is exhibited in
P 4. are erupted. Rostrum of jaw elongated and depressed (see
fig. 5a of the same plate, which represents a section of the penulti-
Fig. 849).
which we have
Materials. — The
above comparison of measurements of mate
named
upjier molar of another Indian fossil sjjecies

thirty-nine specimens in the Falconer and Cautley collection

E. planifrons." As usual in his descriptions the locality is

(1830-1840) and of twenty-seven or more specimensinthe American

given simply Siwalik Hills, but we know from Pilgrim (1913, j).
294), as (luoted above, that this phylum first api)ears as the .species
Museum collection of 1922 made by Barnum Brown (see Summary
E. planifrons in the Pinjor horizon. Upper Pliocene,' some 2,000
of Mca.surements, Tables Vl and VII, pp. 949 and 954) renders it
probable that both collections, totaling sixty-six or more speci-
feetabove the base of the Tatrot horizon. It also occurs in Austria
and Bessarabia.
mens, came from the same general locality and represent ascending
mutations, from the more jirimitivc typical Archidiskodon plani- As early as 1863 Falconer remarked (]). SO): "The nearest
frons, M
3 YiX' toward the more jirogressivc ^4. meridionnlis affinity, and that a very close one, of the I'An-o|)ean E. meridionalis

stage, M
3j^iEj^. We ha\c to do, therefore, with nscrndinq mu- is with the Miocene E. (Lo.rod.) planifrons of India." I'alconer
tations. thus led the way for our present knowledge, namely, that Archidis-
Geologic Locality and Levels. All the chief specimens — kodon planifrons is directly ancestral to .4. meridionalis.
(numbering 39 in Table VI above) listed, described, or figured by E. [Elephas] planifrons Falconer and Cautley, 1846 [1845].
Falconer and Cautley as 'Elephas planifrons' are recorded as
"Fauna Antiqua Sivalensis," 1846, p. 38 [1845, PI. ii, figs. 5a,
simi)ly from the Siwalik Hills, since they did not distinguish be-
tween 'upper,' 'middle,' and 'lower' Siwaliks.
5&]. Lectotype. — A second superior molar with nine ritlges,

the three anterior ridges being in u.sc (Brit. .Mus. M.3068). Co-
Pilgrim (1913) attributes this species only to the 'Upper
Siwaliks,' Pinjor horizon, summit of the Pliocene; he writes (1913,
type. —Broken third molar of lower jaw, I.M,,, with nine ridges
p. 294): "There is absolutely no trace of Elephas either in the
remaining. Brit. Mus. M.2010. Horizon and Locality. —
IMiddle Siwalik or in the Tatrot zone of the Upper Siwalik. It Siwalik Hills, India, Upper Siwaliks, Pinjor horizon. Upper
first ajipears as the species Elephas planifrons some 2,000 feet Pliocene.' Lectotype and Cotype Figures. Falconer and —
above the base of the Tatrot zone." Cautley, 1846 (1845, PI. n, figs. Tm, .56].

'[Recent field studies in northern India l).v Dr. Hcllni\it dc Terra and Perc Teilhard dc Cli.-irdin liavc ofTcrecl strong evidence to support the conclusion
that the Tatrot and Pinjor horizons arc equivaUnit and are entirely of Lower Pleistocene age. The Boulder Conglomenitc ranges up to he Middh' Pleistocene
t

(see Chap. XXII below on the Geologic- Succession of the Probo.scidea for more detailed discussion). This note was prepared by Dr. E. H. Colbert,
January, 1937.— Editor.]
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 951

Description. — (Falconer and Cautley, 1846.1, p. 38): "The between molars of the upper and lower jaws, and of different ages
next serial modification in the disposition of the three dental in the same species."
substances, and in the consequent form of the teeth, is exhibited
in fig. 5a of the same plate, which represents a section of the FALCONER'S NOTES OF 1868 ON TYPE .\ND REFERRED SPECI-
penultimate upper molar of another Indian fossil species which we MENS OF ELEPHAS PLANIFRONS
have named E. planifrons. This tooth shows nine ridges, the three Falconer, "Palffiontological Memoirs," Vol. I, 1868, pp. 423-442, Plate.s
VI, VIII, XI, XII, and xiv of the "Fauna Antiqua SivalcnsLs."
anterior of which alone have been in use, the two first being worn
11,

down to a single disc of ivory. The common nucleus of this Falconer's observations on this species are to be found in his
substance is of less thickness than in the corresponding tooth of successive publications of 1845 (Plates), 1846 (Letterpress), 1857,
E. insignis, and the divisions which are continued upwards from it 1863, 1865, and 1868. In the latter year ("Palseontological
into the centre of the ridges are more elongated, with a narrower Memoirs" of 1868) a summary was pub-
of all his observations
base, forming irregular-shaped wedges. The layer of enamel is lishedby Murchison, covering in detail thirty-nine or more speci-
diminished in thickness and is less uniform in outline, and the mens referred to Elephas planifrons, all presiunably collected
surface in contact with the cement shows a feathered or ragged from the Pinjor horizon, Upper Pliocene.'
edge, indicating superficial inequalities for the firmer cohesion Elephas planifrons: [Lectotype (Burr. Mus. M.3068) and
of this latter substance. The enamel is reflected over the ridges of CoTYPE (Brit. Mus. M. 2010)]. —Plate ii, fig. 5a, M- with 9 ridges,
ivory, and down into the hollows zig-zag wise, exactly as in fig. 6a, cement filling valleys, enamel folds thick, length 8.7 in. = 221
the principal difference being that the ridges are narrower, with mm.; fig. 5b, Ms, vertical section, with 9 ridges.

Lectotype (R.M") 4ND Cotype (L.M3) of Archidiskodon planifrons

Fig. Lectotype and cotype of Elephas planifrons Falconer and Cautley, 1846 [1845, PI. 11, figs. 5a, 56]. From the Siwalik Hills India, Upper
82.'). ,

Siwaliks, Pinjor horizon, Upper Phocenc.' One-third natural size. See "Description of the Plates in the Fauna Antiqua Sivalensi.s" (Falconer, 1867, p. 3; also
Falconer [Murchison], 1868, Vol. I, p. 423):
"Fig. 5a. — Vertical section of penultimate u|)pcr molar [r.M-|, with nine ridges, the tliree anterior of which alone have been in use, the two first being
. . .

worn down to a single disc of ivory." Brit. Mus. M.3068. Molar inverted to show natural ])osition.

"Fig. 56.— "[Cotype.) Vertical section of portion of last molar of lower jaw [I.M3I, with nine ridges." Brit. Mus. M.2010. Osbor n (1924): The
cotype M3 is a broken tooth which typically presents ten ridges, sometimes eleven.
Observe in comparing the original figure (5a) with the new figure (Fig. 827) that Falconer omitted the seven ridge-plated r.M'. See also figure 828 after
Lydckker. M", ridge-plates pre-concave, post-convex. M3, ridge-plates pre-convex, post-concave.

a greater vertical height. The cement substance attains its maxi- Ridge Formulae in Upper and Lower Grinding Teeth
mum of development in this species, completely filling up the wide —
Upper Jaws. Plate vi, figs. 4, 5, 6. Skull with premolar,
interspaces of the ridges, over which it is continued in a thick mass. ?Pm, Dp^ M\ Left Pm with 3 ridges and 2 half-ridges. Dp< with
This tooth measures 8.7 inches in length." 6 ridges also a heel and front ridge, enamel thick. (Footnote, p.
"Fig. 5b, represents a corresponding section of a portion of the 427): Palate with M', 7 ridges and 2 half-ridges; length 6.5 in.

last molar of the lower jaw of the same species, comprising nine = 165 mm., width 3.4 in. = 87 mm., height at 4th ridge 3.5 in.

ridges. This tooth had been longer in use than that of the upper = 89 mm., fifth and sixth ridges with 6 conelets each. Plate xi,

jaw, and all the ridges are more or less worn except the two last. fig. aged palate with r. and l.M', 10 ridges including posterior
1,

same general characters exhibited by fig. 5a, in the

It presents the half-ridge,enamel very thick; fig. 4, M^ right side, remarkable
elongated cuneiform ivory ridges, unequal enamel, and abundant fragment, 6% ridge-plates, length of fragment 5.4 in. = 135 mm.,
cement, the differences being merely such as constantly hold width 2.5 in. = 63 mm. Plate xii, figs. 1, la. Dp- with 4 ridges;

'[See footnote on previous page. —Editor.]

952 OSBORN: THE PROBOSCIDEA

fig. 2, section of r.Dp', with 6 main ridges, a baciv talon and front on 4th ridge, length of Mi 6 in. = 153 mm., width 2.8 in. =71 mm.,
heel, total 6 and 2 half-ridges, length 3.8 in. =97 mm., width length of Mo 8 =204 mm., width 2.8 in. =71 mm.; fig. 7, left
in.

2.3 in. = 59 mm. ; figs. 4, 4a, M' with 5 ridges and heel remaining, lower jaw, M3 very aged, length of molar 10.2 in. = 259 mm.,
two gone, 7 probable total, length 5.2 in. = 132 mm., width 2.8 width 4 in. = 102 mm., three mental foramina; fig. 8, fine lower
in. = 71 mm.; fig. 5, M' [same as fig. 4, this plate], M-, the latter jaw fragment, M2 [M3, Osborn] complete, with 11,'^ ridges, enamel
with 8 ridges and front heel, length of M- 8 in. = 204 mm., width very thick, plates wide apart, much cement, few conelets, three
3 in. = 77 mm. ; fig. 5«, palate with and l.IVP, 8 distinct ridges and
r.

front and back heel, total of 8 ridges and 2 half-ridges, conelets few
Type
Brit. Mus. M.3068
3'\
Right Second
Superior Molar
5v rMI. I

2400
-
ChxuycUgarh'/f-

New Lectotype Figure of Archidiskodon planifrons, R.M-

Fig. 827. New lectotype figure of Elephas planifrons Falconer (Brit. Mus.
M.3068), 9+ ridge-plated second superior molar, of the right side, r.M^, with
worn crown of a ?7/i ridge-plated first superior molar of the right side, r.M'.
Redrawn by Miss G. M. Woodward for this Memoir; one-third natural size.
Compare figure 825.
Ob.serve in comparing this figure with figure 8.54 of the same molar, that
Vip,. 821). Fii\'oral)lo cxpo.surc.s, southwest of Simla, of the Archidiskodon the ridge-plates agree in size with those of Piltdown, England (Fig. 853).
planifrnns life zone, Ujipor Siwaliks, India, chiefly Pinjor horizon, from which
were collected by Barnum Brown in 1922 twenty-seven or more specimen.s
referable to thi.s .sjiecies, especially from Kalka, Cliarnian, Siswan, Chandigarh, mental foramina, length of M2 [M.i, Osliorii] 12.1 in. =308 mm.,
and Mirza|iur, also transitions to .1. mcridimmlis. Compare figures 820, and width 3.8 in. = 97 mm.; fig. fragment of I.M3 nuich worn; fig.
9,
720, PI. x.w.
10, lower jaw, M2 with 8}^ ridges, cement abundant, enamel thick,
no crimping, and no mesial expansion, length of M2 7 in. = 178
and large, enamel thick, length of l.M- 7.5 in. = 191 mm., width mm., width 3.7 in. = 95 mm., I.M3 height of 5tli imworn ridge
3.2 in. = 81 mm., fig. 6, M- with 8 main ridges. Plate xiv, fig. 8, 3.8 in. = 97 mm. Plate xii, fig. 7, right lower jaw, with Dpa with
r.M', showing 8 or 9 ridges and a heel, enamel thick, length 10 in. 6 main ridges and small half-ridge (same formula as in Loxodotila
= 255 mm., width 3.5 in. = 89 mm. (ifricniia); fig. 8, left lower jaw, with small \erticMlIy succeeding
I,oWKU .Iaws. — Plate viii, figs. 2, 2«, jjcrfcct lower jaw, M3 premolar (c), also D])^ with 7 main ridges, double
above it I)p3,

with a total of 911 ridges, enamel very thick, length of M.i 8.8 in. front heeland small half-ridge behind; figs. 10, 10a, first left true
= 224 mm., width 3.8 in. = 97 mm. Plate xi, fig. 2, sti])erl) lower molar, l.Mi, with 7 main ridges, a small ridge in front, no heel
jaw, two mental foramina, r. and l.M.i with 13 ridges and front and behind, length of Mi 6.7 in. = 171 mm., widlli 2.3 lo 2.6 in. = 59
back half-ridges, enamel very thick, length of r.Mj 11.8 in. =300 to 66 mm.; figs. 12, 12rt, Vr.Ma, "apparently of a small sized
mm., width 3 in. = 77 mm.; fig. 3, superb loft half jaw, only 7 indixidual," length 10 in. =255 nun., width in front 3.5 in. =89
remaining ridges in M3, enamel thick, cement abiuidant, "Proved nun., height of crown at 7th jilatc 4 in. = 102 nun., ten main plates
to be E. planifrons by the distance between the plates, the \ery and a front pl.-ilc and lieel; fig. 13, left lower jaw, I.M3 entire,
low crown, thick enamel, and two mental foramina," extreme with about 13 ridges and a heel, or |)ossil)ly 14, length of M3
length of jaw 24.2 in. = 014 nmi., lengtii of M.i 10 in. = 255 mm., 12.7 in. =323 nun., witHli 3.(5 in. = 91 mm., lieiglit at 10th ridge 4.5
width of M,i 3.6 in. =91 mm.; fig. 5, r.Ms, enormous tooth frag- in. = 114 nun. Plate x\\, fig. 9, last lower molar, right side, r.Ms,
ment, very thick enamel, low ridges, and mesial exi)ansion, 9 ridges length 9.5 in. = 241 nun., width 3.5 in. =89 mm., height at 6th
remaining, length of M., 10.5 in. =267 nun., width 4.2 in. = 105 ridge 3 in. = 77 nun., shows 8 ritlg(>K and a heel. Plate xviii.a,
mm., height of 9th ridge 3.5 in. = 89 mm.; fig. 6, left jaw with figs. 1, an I.M3, an enormous specimen; 8 plates, lengtii 10.4
1(7,

l.Mi, ridge-plates 6-I-, M2, ridge-i)latcs 9 and .«mall heel, 5 conelets in. = 265 mm., w idth 4.1 in. = 104 nun., hciglil 3.2 in. = 81 mm.
 :

THE MAMM0NTINJ5: ARCHIDISKODON AND METARCHIDISKODON 953

Collective Ridge Formula (Falconer, 1868). — (1) The regarded as the l.ypr [lectotype] (see Fig. 825). From I^ydekker's
]ieiiiuinent, foiiith premolar, P 4, i)ersists, with 3 ridges. (2) From descriptions the following characters may be summarized
the specimens of Elephas plaiiifrons, ag described by Falconer (39 (1) Both the third and fourth milk molars were vertically
ofwhich appear in Table VI "Summary of Measurements" above), succeeded by ])rem()lars, thus the dental formula is: D)V'"^ Dpo.4,
from tlie Upper Siwaliks, we deduce the collective ridge formula r^"* P3-4, M'"^ M1-3. This succession is an important character
below; (3) the minimum numbers represent partly worn or partly which must be looked for in the ancestors of this species; it does
developed teeth, while the maximum numbers represent fully worn not occur in any other species of the Elephantidae thus far known;
and fully developed teeth; (4) half-ridges develop both in front nor is it found in Slegodon bombifrons. (2) The correct collective

'/-^\

Elephas planifrons. — Vertical and longitudinal section of the second upper true
molar; from the Pliocene of the Siwalik Hills. J. Elephas planifrons.— The hinder half of the third right upper true molar
from the Pliocene of the Siwalik Hills. %. The lower border of the
rig. 828. Lectotype nine and a third ridged upper molar, r.M-, of Elephas figure is the inner border of the specimen.
[Archidiskodon] planifrotis. Reproduced from a wood engraving by Lydekker
(1886.2, p.l02, fig. 24). Brit. Mus. M.3068. One-third natural size. Inverted Fig. 829. Referred right M' of Elephas [= Archidiskodon] planifrons.
to .show natural position of molar. After Lydekker, 1886.2, p. 102, fig. 25. Brit. Mus. M.3070. Two-thirds
a,cement; 6, enamel; c, dentine. natural size. Figured by Falconer and Cautley, 1846 [1845, PI. xi, fig. 4|,
We observe that the cement (a) completely fills the valleys. The enamel and made one of the genotypes of Leilh-Adamsia Matsumoto, 1927, but
(6) is extremely thick, since the sides of the enamel ridge-plates diverge to- regarded by the present author as a synonym of Archidiskodon planifrons.
wards the apex and converge towards the base; the cement interspaces (See Fig. 847 below.)
between the ridge-plates become constantly narrower as the crown wears As compared with figure 828 this is a crown view of a much worn third
down, while the dentine within the enamel becomes constantly broader, as superior molar in which the enamel loops in the middle of the crown are
shown in the figure opposite. contiguous, the dental areas are expanded, the cement areas are contracted.
The cement extends beyond the outer borders of the plates, a highly char-
acteristic feature of Archidiskodon.

and behind the main ridges; (5) the collective formula below
indicates the half-ridges as well as the main ridges. ridge formula is that deduced from Falconer as his maximum-
Maximum and minimum collective ridge formula [of ascending minimum or typical, as given above. The ridge formula of
mutations] of Elephas planifrons: Lydekker (1886.2, p. 99) is incorrect. (3) Crowns of molars broad;
cement in the interspaces frequently very great; enamel usually
Dp 2- Dp 3 H-6-H^P4«^)
6 45
Dp4*^^M 1
i4-7
M
•IVl 2
^
^''—*—
844-9-W thick, frequently devoid of plication in the middle and near the
M3 8-9-VS-K-lg4i
8-9-M-l 0-Vi-V5-l 2-W-l 4-W root of the crown plication near the summit of the crown relative-
;

ly coarse. (4) Ridge-plates subeUipsoidal, frequently with a mid-

(6) It is interesting to observe the half-ridges arising both in
front and behind the main which seems to be a char-
ridges,
compared with Elephas hysudri-
acteristic feature of this species, as Fig. 830. Skull of Elephas
cus. (7) A primitive or simplified formula from Falconer of the
[= Archidiskodon] planifrons from the
Siwalik Hills, India, reproduced from
minimum ridges in E. planifrons from the Upper Siwaliks of India
Gaudry, 1878, p. 185, fig. 246, after Fal-
would be: coner 1846 [1845, PI. x). One-.sixteenth
natural size. This .skull (Brit. Mus.
Dp 21 Dp 3 ^^ (P 4 ±^) Dp 4 ?± M 1 '4 M 2 f M 3 |f±. M. 3060) was erroneously selected by
Lydekker as the type of Elephas
planifrons, and is figured in his

SPECIFIC CHARACTERS OF ARCHIDISKODON PLANIFRONS "Catalogue of the Fossil Mammalia in

(LYDEKKER, 1886; OSBORN, 1924) the British Museum (Natural HLstory)"

of 1886 (1886.2, woodcut fig. 23, p.
The next review of Archidiskodon planifrons is that of Lydek-
100). Compare figures 848 and 817 of
ker (1886.2, pp. 98-107) in which the cranium (Brit. Mus. M. 3060 the same cranium.
—our Fig. 830) was selected as the type ; in the present Memoir the Deperet and Mayet regard this cranium as that of a female, which, if

specimen first described and figured by Falconer and Cautley is true, partly accounts for its small size (.see p. 962 of the present Memoir).
954 OSBORN: THE PROBOSCIDEA

expansion, sometimes imperfectly lozenge shaped, in slightly cephaly), by smallness of the nasal ajierture, by wide di\-ergence of
worn teeth the middle portion of each ridge frequently forms an the incisive alveoli, as in ^4. meridionalis; vertex of skull flattened
isolated disk. (5) Cranium characterized by flatness of the fronto- almost at right angles to occiput, occiput pitching forward 55°
jiarietal region, by a small incision of the temporal fosss on the when the grinding teeth are horizontal; tusks very stout and
frontals, by a comparatively slight elevation of the vertex (hypsi- incurved.

REVISION OF ,\RCHIDISKODON PLANIFRONS BY OSBORN (1927-1928), BASED ON THE SIWALIK COLLECTION

MADE BY BARNUM BROWN IN 1922

The twenty-seven specimens referable to Archidiskodon planijrons and its ascending mutations were found in the 'Upper
Siwaliks,' below the Boulder Conglomerate formation, and recorded as follows:

Table VH. Mea.surements of Twenty-seven Specimens Collected by Brown and Referred by Osborn to Archidiskodon
planifrons, ascending to a. meridionalis

All recorded by Barnum Brown from near Kalka, Charnian, Siswan, Chandigarh and Mirzapur (see Figs. 820, 826)

Superior
 ;

THE MAMMONTINiE: ARCHIDISKODON AND METARCHIDLSKODON 955

As remarked abo\-e, the large American Museum collection of localities of the Pinjor horizon level, namely, from deposits around
Archidiskodon -planifrons, including examples of the entire den- Kalka (see Figs. 826, 820). The following 29 si)ecimens, belonging
tition, but lacking a skull, renders it highly i)robable that the to 24 individuals, are given with measurements above (Table VII)
Falconer and C'autley collection of A. phuiifroiis, figured and tliey are also illustrated in great detail in the accompanying
listed in detail, as above described, came from the same geologic figures (Figs. 831-844).

Near Kalka 9 miles west Amer. Mus. 19778 Fragment of right jaw with Mi and M2 in situ (Fig. 836 for M,).
6 miles west Amer. Mus. 19798 Fragment of jaw with r.Ms in situ (Fig. 855 FOR
r.Mj).
Near Charnian 2 miles south Amer. Mus. 19819 Fragment of left jaw with I.M3 in situ (Fig. 844).
Near Siswan Amer. Mus. 19821 Fragment of maxilla with l.M- in situ (Fig. 833).
Amer. Mus. 19870 Fragment of jaw with I.M2 and I.M3 in situ, M2 incomplete, M3 partly
erupted (not figured in ])resent Memoir).
Amer. Mus. 19879 Fragment of jaw with l.M 3 in situ (Fig. 846).
Amer. Mus. 19880 Left AF (not figured in present Memoir).
Amer. Mus. 19881 Left M'
(Fig. 831).
Amer. Mus. 19882 Fragment of maxilla with r.M'' (not figured in ]iresent Memoir).
3 miles north Amer. Mus. 19871 Fragment of jaw with l.Mi and M2 in situ (not figured in pre.sent Memoir).
3 miles north Amer. Mus. 19873 Fragment of jaw with r.Dp4 in situ (Fig. 837).
3 miles north Amer. Mus. 19968 Fragment of jaw with I.M3 in situ (Fig. 843).
Amer. Mus. 19965 Fragment jaw with r.Ms in situ (Fig. 840).
of
Amer. Mus. 19862 Fragment jaw with l.Dp4 and l.Mi (Fig. 838).
of
Near Chandigarh 3 miles west Amer. Mus. 19861 Fragment of jaw with I.M3 in situ (Fig. 841).
3 miles west Amer. Mus. 19916 ?Right M' (not figured in present Memoir).
3 miles west Amer. Mus. 19950 Fragment of maxilla with l.M' in situ (Fig. 834).
1 mile west Amer. Mus. 19951 Right M3 (Fig. 842).
1 mile west Amer. Mus. 19952 Left M3 (Fig. 835).
3 miles west Amer. Mus. 19955 Right Ml and left M' (Fig. 832).
1 mile west Amer. Mus. 19961 ?LeftMl (not figured in present Memoir).
3 miles west Amer. Mus. 19917 Right Ml (not figured in pre.sent Memoir).
Probably near Chandi-
garh, record incomplete Amer. Mus. 19967 Lower jaw, left ramus.
Near Mirzapur 3 miles north- Amer. Mus. 19864 Fragment of jaw with r.Ms in situ (Fig. 839).
east

SUMMARY OF DENTAL MEASUREMENTS OF THE AMERICAN MUSEUM COLLECTION MADE BY BARNUM BROWN IN 1922

These measurements accord in general with those of the First Superior Molar, M' Minimum
Falconer and Cautley Collection in Table VI above and exhibit Length
variations in size, length, breadth, and proportions, partly attrib- Breadth
utable to male or female sex, partly to progressive ascending muta- Breadth-length index
tions. The increase in number of the ridge-plates and half ridge- Height
plates is probably attributable to progressive evolution or to Ridge-plates
ascending mutations ranging into higher geologic levels. In Third Inferior Molar, M3
general, the smaller animals, with fewer ridge-plates, probably Length
occurred in lower geologic levels than the larger animals, with more Breadth
numerous ridge-plates and other progressive characters. Breadth-length index
Height
Third Superior Molar, M' Minimum Maximum
Ridge-plates
Length 201 254
Breadth 88 109 Second Inferior Molar, M2
Breadth-length index 38 47 Length
Height 66e 102 Breadth
Ridge-plates 9-1- 12 Breadth-length index

Second Superior Molar, M^ Height

Ridge-plates
(Only specimen available in Amer.
Mus. Coll., No.19821) First Inferior Molar Mi
Length 217 Length
Breadth 94 Breadth
Breadth-length index 43 Breadth-length index
Height 66e Height
Ridge-plates 10 Ridge-plates
 'A A/at. sij

^A/dTsije

L.M- ARCHIDISKODON PLANIFRONS Ri-/ Amer. /^U5. /982/

ARCHIDISKODON PLANIFRONS «»/.

ARCHIDISKODON PLANIFRONS Ktf.

Amer. Mus. fSBBJ L.M

Amer. Afuj. /S>$SS
3 _4- J

Fig. 832
Fig. 833
Fig. 831

ARCHIDISKODON PLANIFRONS «./

American Museum Collection

Upper Pliocene Abchidiskodon plani-
frons of the plnjor horizon
(see Fig. 826)
Figs. 831-835. Archidiskodon planifrons,
referred superior and inferior grinding teeth of
the Barnum Brown Siwalik Collection, listed
with measurements in Table VII. All figures
drawn to the same one-fourth scale. Cement
(dotted), dentine (horizontal lining). Com-
pare Falconer's measurements in Table VI.
Figures: (831) First left superior molar,
l.M'. (832) First left and right superior
molars, I.M', r.M'.
(833) Second left .superior molar, l.M^;
a primitive stage as shown in midsection
(Fig. 855); this M- agrees closely with that of
Piltdown, Sussex (Fig. 853). (834) Third
left superior molar, 1.M^ 12 plated.

(835) Third left inferior molar, I.M3.

i^ A/at. sije

Fig. 834

956
 Fig. 836 Fig. 837 Fig. 838

Fig. 839 Fig. 840

American Museum. Upper Pliocene Archidiskodon planifrons of the Pinjor Horizon (see Fig. 826)
Fig,';. 836-840. Inferior grinders of Archidiskodon planifrons (Figs. 836-840), in the Barnum Brown Siwalik CoUeRtion, fully listed with measurements in
Table VII. All figures one-fourth natural size, excepting 837 which is one-third natural size. Cement (dotted), dentine (horizontal lining). Compare Falconer's
measurements, Table VI.

Figures: (836) Portion of first and .second right molars, r.Mi, r.M^. (837) Fourth deciduous iiremolar, Dp4, with 9 ridge-plates. (838)Worn crown of
left fourth premolar, l.Dp^, and first molar, l.Mj. (839) Portion of lower jaw with third right molar, r.Ms. (840) Portion of lower jaw with third right
ranlar, r.M.i.

957
 jL=6=s^g=S:

Fig. 843 Fig. 844

American Museum Archidiskodon planifrons of the Pinjor Horizon (see Fig. 826). Inferior Molars
Figs. 841-844. Archidiskodon planifrons referred infcuor grmdiu^ teeth and jaws in the Siwalik CoUoction of the American Museum made by narniini
Brown, as fully listed with measurements in Table VII. Ail figures one-fourth natural size. Cement (dotted), dentine (horizontal lining). Comi)are
Falconer's measurements, Table VI.
Figures: (841) 12 plated left third molar, I.M3. (842) 12 plated right third molar, r.Mj, of a slightly older individual; a progressive stage as shown in

midsection (Fig. 85oB). (843) 9+ plated left third molar, I.M3, imperfect anteriorly. (844) II plated ieft third molar, I.M3.

958
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 959

LEITH-ADAMSIA SIWALIKIENSIS
Figure 847
Lp:ith-Adamsia siwalikiensis' Matsumoto, 1927. "On —
Leith-Adamsia siwalikiensis, a New Ceneric and Specific Name of
Archetypal Elephants." Japanese Journ. Geol. and Geog., V,
No. 4, Art. 12, 1 page. —
Type. Two superior molars, both of
the right side, r.M', in the collections of the British Museum
(Bfit. Mus. M.3070 and 36695). From India. Type
—
Figure. Falconer, "Fauna Antiqua Sivalensis," Pis. xi, fig. 4,
and XIV, fig. 8. Type Description.— (Matsumoto, 1927.2):
"In the course of a study of fos.sil Elephants, the writer has come
to be faced with the serious need for a ]>roper name for a certain
type of archetypal Elephants of India. He means the small and
narrow-molared form recorded under the name of Elephas plani-
frons.This form appears, in all likelihood, to stand at the starting
point of the entirephylum of the Loxodontine Elephants. As it
would appear occupy too important a position among the
to
Elephantidse to be left unnamed, the writer proposes here to call

/Imer: Mus. /995/

DON PLANIFRONS Rtj

Upper Pliocene Archidiskodon planifrons of the Pinjor Horizon

(see Figs. 826, 820)
Fig. 845. Arclndiskodon planifrons ref., third right inferior molar, r.Mj
(Amer. Mus. 19951), in the Barnum Brown Siwalik Collection, one-third
natural size. Compared with the specimen is
collective figures above, this
very close to the typical ridge formula of Archidiskodon planifrons, namely,
M 3TTm ridge-plates. I..ength 313 mm., width 101 mm., height of seventh
ridge-plate 124 mm. (see Table VII, p. 9.';4).

^.Af./S^S7S ysA/oTstje
Upper Pliocene Archidiskodon planifrons of the Pinjor Horizon (see Fig. 826)
Fig. 846 (right). Archidiskodon planifrons ref., a third inferior molar of the left side, I.M3 (Amer. Mus. 19879) in the Barnum Brown Siwalik Collection,
listed with measurements in Table VII; portions of ten ridge-plates are preserved, as displayed in the transverse vertical section of
the same tooth (left).
All figures one-third natural size. Compare Falconer's measurements Table VI, p. 949.
'[Regarded by Professor Osborn as a synonym of Elephas [Archidiskodon] planifrons. —Editor.]
 7

9G0 OSBORN: THE PllOBOSCIDEA

itby a generic name as follows, in honour of the late Professor profile of A. meridionalis (Fig. 817). Meanwhile the cranium
Doctor Leith Adams, the eminent i)akpontologist and specialist on heightens (hypsice])haly) and dee])ens (bathyceplialy), in adapta-
fossil Elephants." tion to the enlarging and deepening of the third superior molars,
also, as fully explained in Chapter XV, j). 915, on the mechanics of
" Leith- Adamsia, gen. no\-. A genus of archetypal Elephants.
the proboscidean cranium, the heightening of the occipital crest
Cheek-teeth subhypsodont, narrow-crowned, with a low ridge-
(acrocephaly) is in adaptation to the elongating tusks and the
formula; lo.xodont sinus present, and of an obtuse ty])e; disks
strengthening of the cervical ligaments and muscles which sway
of well-worn ridges may be more or less lozenge-shaped."
the great tusks and proboscis. Nothing is known of the limb
skeleton of A. planifrons, but it is inferred that the animal was
greatly inferior in shoulder height, in length, and in i)roboscis
/i,/ t development to its giant successors A. meridionalis and A. impe-
rator. The completely preserved tusks in members of this species
discovered in southern France (Fig. 850) lack the strong outward
curvature and incurvature characteristic of A. meridionalis
(lyrodon), see figure 864, reaching a supreme stage in A. imperator.

Referred Archidiskodon planifrons

Fig. 847. Type molars of Leith-Adamsia siwalikiensis Matsumoto,
1927. After Falconer and Cautley, 1846 [1847, PI. xi, fig. 4, and Pi. xiv, fig. 8), ..) i.
..V ARCHIDISKODON PLANIFRONS /?e/
Xz'VoTsije A/ter Falconer Platt$ S \ 10
one-third natural size, reduced to one-fourth natural size. i

(PI. XI, fig. 4) Brit. Mus. M.3070. An r.M', posterior half of crown,
exhibiting 6^ ridge-plates (same molar as in Fig. 829, above). Supposed Female Cranium
Fig. 848. Cranium of Archidislcodon planifrons in the British Museum
(PI. XIV, fig. 8) Brit. Mus. 36695. An r.M^ with 6^ posterior ridge-plates
Mus. M.3060) redrawn after Falconer and Cautley, 1846
(Brit. [1845, PI. ix,
preserved of a probable total of 10 ridge-plates.
front view of skull, PI. x, side view of skull). Five views of this unique cranium
are shown in Falconer and Cautley's plate.s ix and x; a brief descrijjtion in
the 'Talaeontological Memoirs" of 1868, Vol. I, p. 430, is as follows:
CRANIAL CHARACTERS OF ARCHIDISKODON PLANIFRONS
"Plate IX. Elephas planifrons (Falc. and Caut), from the Sewalik Hills.
The single cranium known in the British Museum collection Front view of skull, one-third of natural size. The forehead of this species is
(Brit. Mus. M.3060— .see Figs. 830, 848, and 817) is extremely very flat; the naso-maxillary opening very small, and the occipital fissure
very low. . Plate x.
. . The last true molar is seen in germ and intact on the
. . .

primitive, somewhat resembling that of certain species of Stegodon,

right side, and well worn on the other, so that the corresponding tooth on the
for example, Stegodon pinjorensis type (Amer. Mus. 19772 see — right side of the lower jaw had probably been wanting. It has eleven ridges

Figs. 711, 765, and 817C), except that Archidiskodon planifrons is and a heel. The pterygoids are very low."
much less elevated (acrocephalic) than S. pinjorensis, as clearly The principal measurements given by Falconer (cf. op. oil., 1868, p. 430)

shown in the comimrative profiles (Fig. 817). This is in adaptation arc the following:

to the relatively abbreviate 10 -|- ridge-plated crown of M' in A.

Extreme length from occiput to incisive alveoli.. .25 in. =635 mm.
planifrons, as compared with the extremely elongate 15 ridge-
Extreme width of occiput 21 .7 =551
Ijlated crown of M' in .S. pinjorensis. Also compare the flattened
Height of occiput 13.7 =348
forehead of .4. planifrons with the abbreviated and extremely Occiput to anterior border of orbits 20 .
= 526
elevated forehead of S. pinjorensis. Length of right M' 9.7 =247
This flat-faced condition, to which Falconer assigned tlie Width of crown 3.5 =89
specific name planifrons, develojis into tlie concave-forehead Height of crown plates 4.0 =102
 ;

THE MAMMONTINiE: ARCHIDISKODON AND METARCHIDISKODON 961

ARCHIDISKODON PLANIFRONS IN WESTERN EURASIA, AUSTRIA, RUMANIA, AND RUSSIA

The species of southern mammoth known as Archidiskodon discovered in ferruginous sands near the village of Farladani
meridionalis has long been known in western Eurasia. The dis- (Bessarabia), a locality which had previously yielded remains
covery of the more primitive ancestral A. plam'frons in Austria, of Mastodon (Zygolophodon) and thus regarded as of
borsoni,
Bessarabia (southern Russia), southern France, and England, is Upper Pliocene age. Madam Pavlow concludes (p. 27) "C'est :

relatively recent, dating from Schlesinger's announcement of the une dent (m"?) tres massive (fig. 23), surtout dans sa partie
discovery of 'Elephas' plam'frons in Lower Austria in 1912. superieure. Elle est tres jeune, a peine usee, sur une surface de 15
Austria. —Schlesinger in his "Studien fiber die Stammes- cm., elle n'a que 5 lames aux contours tres irreguliers, a I'email tres
geschichte der Proboscidier," of 1912, p. 89, announced the evi- epais et avec de larges espaces de cement. Les quatre premieres
dence of Elephas planifrons in Lower Austria: "Um so iiber- lames sont us6es, la 5-e tres peu; la 6-e presque intacte, ce ne sont
raschender war es, als dem niederosterreichischen Landesmuseum que des rondelettes liees entre elles. Par derriere se trouvent
in Wien ein Elefantenmahlzahn von ungemein primitivem Charak- encore 9 plaques intactes; elles sont toutes tres robustes et
ter zukam. Es war sehr naheliegend, das Stiick mit einer der mesurent sur le cote 2 cm. entre les enfoncements qui les bornent.
beiden im Jungtertiiir Europas nicht seltenen Arten zu identifi- Je n'ai pu trouver aucune forme en Europe qui pourrait corre-
zieren. Ein nur oberflachUcher Vergleich riickte den LTrelefanten spondre avec cette dent par le dessin de I'email, ainsi que par le
{E. antiqmis Fnlc.) giinzlich ausser Betracht, eingehende Studien nombre des lames, et c'est parmi des figures donnee par Falconer
aber sprachen zufolge eben der Merkmale gegen eine Bestimmung pour I'El. planifrons que j'ai trouve ses semblables. T. XI, f. 57.
als E. meridionalis Nesii, welche den Zahn dem E. planifrons Falc, T. XVIII, f. 12. T. XIV, f. 8, 9 (Fauna Antiq. Sivalensis)."
einer typischen Form der indischen Sewalik-Hills [Footnote: 'Ich —
Summary. Mayet (1920, p. 310) remarks: "Une revision des
bemerke, dass dies der von H. Falconer und P. Cautley (Palaeon- Elephants pliocenes d'Europe devient n^cessaire, a la suite de
tological Memoirs Vol. I, pag. 31) zum erstenmal gebrauchte laquelle une partie des molaires regardees comme appartenant a des
richtige Name fiir die dem Himalaya siidlich vorgelagerte Hiigel- formes archaiques d'E. jneridionalis seront vraisemblablement con-
kette ist.'] nahe brachten. siderees comme provenant de E. planifrons. La necessite de cette
Die Annahme des Vorkommens einer so ausschliesslich revision a ete entrevue par d'eminents paleontologistes. La mise
sewalischen Art in unserem Gebiete mag vorerst befremdend und au jour dans les sables de Chagny d'un E. planifrons I'impose.
gewagt erscheinen. Doch schwinden derartige Zweifel alsbald, Deja M™'' Pavlow [Footnote: 'Marie Pavlow, Les Elephants
wenn wir bedenken, dass sich die Verbreitung der Riisseltiere, fossiles de la Russie (Nouveaux Memoires de la Societe imperiale
wie die etiicher Siiugerstamme, ohne die Annahme ausgedehnter des Naturalistes de Moscou, t. 17, p. 2.')] a signale son e.xistence
Wanderungen nicht begreifen lasst [Footnote: 'Vgl. Ch. Deperet, dans le Pliocene recent de la Bessarabie et le Dr. Schlesinger
Die Umbildung der Tierwelt (deutsch von R. N. Wegner), pag. [Footnote: 'Giinther Schlesinger, Studien iiber die Stammesge-
260 ff., Stuttgart 1909.']." schichte der Proboscidier (Jahrbuch der k. k. Geolog. Reich-
This specific determination as well as the geologic age of the sanstalt, Vienne, 1912, p. 87).'] dans celui de la Basse-Autriche,
specimen very carefully described in great detail by Schlesinger a Dobermannsdorf."
(op. cit., pp. 94-111) were challenged by Soergel in articles entitled
"Die Stammesgeschichte der Elephanten," 1915, and "Die Plani- ARCHIDISKODON PLANIFRONS IN FRANCE
frons-Frage," 1921, with the following conclusion, namely, that An indubitable discovery of Elephas planifrons Falc' is that
the teeth from Dobermannsdorf, Krems, and Laaerberg are recorded by Mayet in his"Decouverte d'un squelette d'Elephas
actually referable to Elephas meridionalis and that the geologic age planifrons Falconer dans les sables de Chagny, a Bellecroix pres
is not Middle Phocene but of much more recent date. Schlesinger ("hagny (Saone-et-Loire)," 1920, in the Comptes Rendus (Paris),
replied in great detail in his "Meinc Antwort in der Planifrons- pp. 308 311.
frage" of 1916. —
Geologic Age. (Mayet, 1920, j). 308): "Une faune abon-
The molar referred to A . planifrons from Laaerberg, said to be dante les date parfaitement les sables de Chagny se parallelisent
:

a third inferior molar of the left side (see Schlesinger, 1916.2, p. avec les sables a Mastodontes de la region du Puy, avec les alluvions
119, fig.s. 6, 7) does not in Osborn's opinion sustain the specific tie Perrier et probableinent avec le Crag de Norwich. lis sont de

reference of this specimen to "Elephas {Archidiscodon) planifrons" I'extreme debut du Pliocene superieur. Les depots marecageux
it istoo high and narrow, the ridge formula (IO/2-II), length (280 du cirque de Seneze sont plus recents; les gisements classiques du
mm.), width (87 mm.), index (31), and height of the 4th ridge-plate Val d'Arno superieur, du bassin de Florence, des niveaux fluvio-
(106 mm.) seem to rule out this third inferior molar from A. plani- lacustres de I'Astesan, sont du meme age. Cet ensemble differe de
frons and to relate it to Palseoloxodon [Hesperoloxodon] antiquus. celui des gisements plus recents qui appartiennent au Saint-
—
Bessarabia, Russia. In the valuable Memoir by Marie Pav- Prestien: sables a E. meridionalis de Saint-Prest, tufs volcaniques
low, "Les Elephants fossiles de la Russie" (1910), cited by Mayet, de Saint-Martial (Hcrault), limons de Durfort, etc. Dans ces
she fully discusses in her description of the elephants of Tiraspol gra\-iers de Chagny-Bellecroix, fin mai dernier, a etc decouvert un

(p. 4) the milk and permanent dentition of Elephas planifrons and squelette incomplete d'elcjjhant: base du crane (celui-ci tres
on page 27 she describes "El. id. planifrons Falc, PI. i, fig. 23," fragmente, dcjjourvu de sa moitic antero-superieure) a\'ec deux

'[Professor Osborn regarded this specimen as a Lower Pleistocene form (see Fig. 1239 of present Memoir) and intended to make it the type of a new species
of Archidiskodon. The description, however, —
was never written. For restoration, see figure 815. Editor.]
962 OSBORN: THE PROBOSCIDEA

molaires, M'; mandibule avec deux niolaires en place, Mj, et une

apophyse nientonniere bien developpce, deux defenses, atlas,
onioplate, femurs, cotes, etc. Ces ossements ne justifieraient qu'-
imparfaitement la presente Note s'ils ne se rajiportaient a une
espece non encore identifi(5e parmi les elephants pliocenes de
1
'Europe occidentale: Elephas planifrons Falconer."
In the original and subsequent papers the authors compare this
important specimen both with the typical Elephas planifrons of the
Pinjor horizon and with Elephas meridionalis. Mayet and Roman
(1923) devote to this subject an exhaustive review in their "Les
Elephants Pliocenes" (Premiere Partie), in which by far the most
profound study is made of the dentition of the Upper Pliocene
Proboscidea of France, and Depi^ret and Mayet {pp. cit., Deuxieme
Partie) of the chief specimens referable to this species from the
Siwaliks (pp. 95-97), from Bessarabia (pp. 101, 102), from Austria
(the Schlesinger discoveries, pp. 102-104), as well as specimens
possibly referable to this species from England (p. 104), from other
II
parts of France, including especially Chagny (pp. 106-110), from
Italy (pp. 110-120), and possibly also from Africa (p. 120). From
all these materials the following specific characters are deduced.
Characteres Specifiques de L'Elephas planifrons (De-
PEKET AND Mayet, 1923, PP. 121-123).— "[1] Le crd^e male adulte
est encore inconnu. Le seul crane qui ait ete dccouvert est le
crane des Siwaliks figure par Falconer (pi. ix et x), et reproduit
dans tous les ou\rages. im crane femelle de taille relati\-e-
C'est
ment faible, au \ertex peu eleve, au front i)lat et non excave, peu
echancrc sur le cote par la fosse temporale; le plan fronto-nasal
est oblique en avant, tres peu redresse. Ce sont la des caractcres
qui se retrouvent dans les cranes jeunes et dans les cranes femelles
des autres especes d'elephants vivants et fossiles. [2] Les defenses,
III bien conservees dans le sujet de Chagny (1" partie, p. 77), sont
tres rapprochees Tune de I'autre a la base, peu divergentes; elles

sont longues (plus de trois metres), mais surtout tres epaisses

(grand diametre, maximum 22 centimetres) ; leur incurvation est
faible vers le haut et un peu en dehors, avec a peine une tendance
spiralee. [3] La mandihule bien conservee des pieces de Chagny et
Primitive Lowek Jaws of Auchidiskouon de Seneze presente un caractere distinctif importavt: la symphyse
Fig. 849. Siipcrpo.sitioii of three mandibles of Elepkas pUiniJroms, Imliii se jirolonge en avant par une sorte de hec allonge ct recnurbe en bas.
ami France, reduced to approximatrcly the same one-tenth scale, after Mayet Ce caractere existe egalement dans la mandibule des Siwaliks
and Roman, 1923, p. 81, fig. 13. See figure 914 below. figuree par Falconer (pi. viii, fig. 2) sous le nom erronc d'E.
hysudricus. On voit egalement ce bee un peu brise au bout dans la
I, Elcfihas planifrons of the Siwaliks. After Falconer.
mandibule du Serre, pres Peccioli (Musee gdologique Florence).
IT, Elr/ihas pUmifrnn.'^ of Chagny-Bellecroix, France. See also figure S5(). [4] La dentition est complete: 3 molaires de lait et 3 arriere-

Ill, Elcphas planifrons of Senezc, France.

molaires. On a observ6 dans les pieces de I'lnde la presence de
deux premolaires inferieures de seconde dentition (fig. 15, p. 122),
We observe the uniform prolongation and beaklike depression of the caractere unique dans tout le groupe des Elephants. Ces pre-
symphysis. In describing these jaws Mayet and Roman remark (op. cil., p. molaires n'ont pas encore etc observees dans les sujets d'Europc
79): "Mandibule (pi. i, fig. 1 et 2). Courte, a branches horizontalcs ba.s.ses, et c'est la seule objection qui puisse subsister sur leur assimilation
opaisscs, la mandibule donne rimpression d'etre massive en arriere, amineie, specifique avec I'espece des Siwaliks. Nous pensons qu'on ob-
effiI6c dans sa partie ant6rieurc a comme projet6e en avant. Cette impression servera ces premolaires lorsqu'on aura des mandibules a l'6tat
tiont il la direction dc la sympliyse et, pour >me grande partie, i\ rcxistcnee d'un
d'evolution individuelle convenable. [5] Parmi les molaires, les
'bee',

182.'),
d'une ,sort,e d'apophy.se mentonnidre, qui plonge obliciuement.
Nesti disait quo I'K. meridionaliis etait un Elephant a bee et,
Doja, en
dans Fauna
Mm sont les plus caracteristiques: la formule dentaire est de 9
ii un peu inferieurs
10 lames en haut, et de 10 a 11 en bas, chiffres
anliqxia sivaleMsis, Falconer insiste sur le bee d'E. planifrons. II faut toutefois
a ceux de V K. meridionalis. La frequence laminaire est de 3, 5,
rcmarquor, des maintenant (voir aussi 2° partie), que Ic bee de I'E. mcridion-
alis so dirigc en avant pre-sipie horizontalement et quo colui d'^. planifrons a 4 ])our 10 centimetres de longueur de couronne, chitTre toujours
jjlonge vers Ic bas on tondant a .so rapprochor do la vertiealo. Ce caractere est inferieur a. celui de VE. meridionalis (4, 5, k 5). La couronne est
particuli^rement net sur unc mandibule de Scneze (v. pi. n)." notablcment plus basse que dans cette derniere espcice; elle est

'[See footnote on previous page. — Editor.]

 —

THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 963

d'un type remarquablcment brachyodonte. Les lames sont larges, ARCHIDISKODON MERIDIONALIS [AND ?PLANIFR0NS1 IN THE
I'email epais et generalement
peu festonne, la plicature n'interes- PLIOCENE RED CRAG AND NORWICH CRAG OF SUFFOLK
sant qu'exceptionnellement la totalite du cordon d'email. On AND NORFOLK, ENGLAND
observe un sinus loxodonie inconstant, plus frequent sur le cote Compare page 972; see figure 871 G, H
posterieur des lames, mais existant parfois des deux cotes. [6] La The Red Crag is of greater antiquity as a whole than the
taille de hauteur du sujet de
\'E. planifrons est considerable: la Norwich Crag (Judd, 1911, p. 185; Osborn, 1922..563, p. 436);
Chagny au garrot devait etre d'environ 3 m. 85 metres: c'est une in it are recorded both Archidiskodon and A. meridion-
(?) planifrons
dimension a peine un peu inferieure a celle d'E. meridionalis." alis. The less richly mammaliferous Norwich Crag is partly more
recent and yields ^4. meridionalis (Osborn, op. cil., p. 437). To-
gether these horizons represent a very long period of closing Pli-
ocene time, in which A. planifrons may have evolved into .4.
meridionalis. Compare similar ascending mutations in Europe
(Deperet) and in India (Osborn).
Possibly belonging to Archidiskodon planifrons are the Red
Crag teeth, 'broad plated with very thick enamel,' and the femur,
referred to "E. antiquus" and "E. meridionalis" by Leith Adams
(1877-1881, PL XXVI, figs. 2, 3, 4).

In the Ipswich Museum (Osborn, 1922.563) is to be found

a typical Upper Pliocene collection recorded from the Red Crag
horizon; it contains si)ccimens referred to Castor {Trogontherium},
Orca (Trichecodon) huxleyi, Hyxnarctos, Felis pardoides, Ursus
arvernensis, Mastodon [Anancus] arvernensis (27 specimens),
Mastodon [Zygolophodon] borsoni (20 specimens), also two 10-11
ridge-plated molars (?)erroneously referred to E. [
= Hesperolo.ro-
Fit.. 12. — l.leph.\s pUnifrons de Ilellccroix-Chapny. don] antiquus and E. [
= Archidiskodon] meridionalis. Compare
Primitive Tusks and Jaws ok Archidiskodon of Chagny figure 871 G, H, Archidi.skodon (f) planifrons.
Fig. 850. Tu.sks, ma.xillie, condyle, and atlas of Elephas planifrons of In three British collections and in several faunal lists (Osborn,
Chagny. After Mayet and Roman, 1923, jj. 77, fig. 12. From the sables de 1922.563, pp. 436, 437) six grinding teeth referred to 'Elephas
Chagny at Bellecroix near Chagny, France. Upper Pliocene. About one-
meridionali.'i' are Red Crag or to the Norwich
attributed to the
thirtieth natural .size. This .si)ecimen is carefully described (op. oil., 1923, pp.
The
Crag of East Anglia, an Upper Pliocene stage; it is possible that
7.5-80). when viewed from above, with
tusks are massive, nearly straight
The tusks measure: right 2.15 m.,
a very feeble torsion from within outwards. certain of these grinders are referable rather to the more primitive
left 2.18 m. The third sujierior molar measures: ap. 264 mm., tr. 117 mm.; Archidiskodon planifrons. See Falconer's notes (1868, Vol. II, pp.
length of median lamina 26 mm. The lower molars measure: ap. 305 ram., 130-132, and PI. viii, figs. 1 and 4) on "Elephas {Loxodon) meri-
tr. 108 mm., height 44 mm.; laminar formula 10 + sujjerior molars 9+, ;
dionalis.'"Falconer repeatedly compares these teeth to those of
inferior molars 10+. For restoration, sec figure 815 above.
"E. (Loxodon) planifrons"; see full abstract below (pp. 972-974)
of Falconer's notes of 1868.
Description of Chagny Specimen. An animal from the — The finely preserved 10-11 ridge-plated molar in the Ipswich
Upper Pliocene' of Chagny, France, is characterized by Mayet
IMuscum (Fig. 8710 below) from the 'Norfolk Bone Bed,' re-
(1920, i)p. 309, 310) as follows: "Nous pouvons ra])])rochcr notre
ferred to 'Elephas meridionalis,' appears to Osborn to resemble
clcjjhant de Bellecroix de Elephas plniiifrons du Pliocene moyen et
Archidi.'ikodon planifrons, as indicated in the figure; the approximate
superieur de I'Inde (Pinjor horizon, Siwalik-Hills) meme formule :

measurements are, length 190 mm., breadth 77 mm., index 41;

^Yoif meme couronne trcs basse, meme caractcre de I'email,
it exhibits ten to eleven ridge-plates only (cf. Amer. Mus. 19864^
meme aspect mastodonto'idc de la mandibule, du menton, des
Fig. 839) ; it is greatly inferior in breadth to the more recent
defenses, il y a identite des caracteres specifiques."
Forest Bed molar (Fig. 871 F) attributed to A. meridionalis
Frequence Formule (Savin Mus., Cromer, No. 197), to be described below under
"Es])cces. Eapportf-. laminaire. dentaire. Archidiskodon meridionalis cromeren.sis, also to the Forest Bed
x-l 0-J
E. planifrons, Chagny 2,28 4 x-l O-x molar (Fig. 871 E). The measurements and other characters of
E. planifrons, Siwalik- these molars resemble very closely those of the molars described
__x-l_o-x_,,
Hills 2 a 2, 5 4 x-l 0-1 1-x by Deperet and Mayet in their Memoir of 1923 and figured in
The number of plates in 10 cm. ("frequence laminaire") is PL IX.

four as compared with five to six in E. meridionalis. The x of Lydekker (1886.2, p. 113) provisionally refers to yl;v7(«//sfcorfo«
Mayct's dcscrijjtion refers to the rudimentary plates at the front meridionalis a mandibular symi^hysis from the Norwich Crag of
and back of the crown. It would ai)i)ear that the ridge formula of Thorpe Mus. M.2009); four ridges of a true molar, trans-
(Brit.
E. planifrons in comparison with the Chagny specimen may be versely cut and polished, from the Red Crag of Felixstowe, Suffolk
written: (Brit. Mus. 44895) also fragment of a molar, vertically and
;

M 2f M 3f^ longitudinally cut and polished, from the Red Crag of Fakenham,
'[See footnote on page 961 above. —Editor.]
 :

964 OSBORN: THE PROBOSCIDEA

Suffolk (Brit. Mus. 44140). These specimens are probably attrib-

utable to A. planifrons.
Osborn, 1929: Certain of the 'Norfolk Bone Bed' or Red
Crag molars agree precisely in measurement (length, breadth, in-
dex) and in the number of ridge-plates with certain of the Archi-
diskodoti planifrons molars of the American Museum collection
made by Barnum Brown, e.g., figure 839, found three mOes north-
east of Mirzapur, Pinjor horizon, India, indicating a common
eastward to westward range of A. planifrons from the Siwalik
Hills to East Anglia in Upper Pliocene time.

GEOGRAPHIC DISTRIBUTION OF ARCHIDISKODON PLANIFRONS

(COMPARE SCHLESINGER, 1911, 1912, 1913, DEPERET AND
MAYET, 1923, SABBA STEFANESCU, 1927,
HOPWOOD, 1935)
The migration area of Archidiskodon planifrons extends from
India to western Europe, passing through southern Russia (the
Caucasus, Crimea, Bessarabia) the basin of the Danube, near
Vienna, Italy (Astesan, the superior and inferior Val d'Arno),
France (Chagny, Seneze, le Puy, Randan) to England (Crag of
Suffolk and Norfolk, Piltdown grav^els of Sussex) finally in northern ,

Africa (Algeria). In many of these localities the geologic and

stratigraphic level is difficult to determine precisely. The type
horizon of the Pinjor, Siwaliks, is regarded as between Middle and
Upper Pliocene.' In Russia and Austria the formations containing
A. planifrons are Pliocene, but it is difficult to exactly determine
the level. In France and Italy it is possible to determine with great
Fig. 851. Type localities of Archidislcodon proplanifrons (22) and of A.
precision the horizon of Puy, of Seneze, and of Chagny, which are
planifrons (2) in circles. In solid black (2), Upper Pliocene' localities of A.
the exact equivalents of the classic horizon of Perrier (Auvergne)
planifrons referred specimens. In oblique lines, theoretic range from the sup-
and constitute the most ancient phase of recent Pliocene time posed African center (22) northward to France and Britain, eastward to India.
(Calabrian = Villafranchian) at the limit of the 'Pliocene ancien' Compare Dcporet and Mayet (1923) who give the route of migration in the

(Astian). We may therefore consider that A . planifrons character- ojjposite direction.

izes the horizon bounded by the 'Pliocene ancien' and the 'Pliocene
recent,' a horizon somewhat in advance of that which contains
A. nicrididrialis, a species In which A. planifrons is related by a con- ARCHIDISKODON PLANIFRONS OF THE PILTDOWN GRAVELS
(Figs. 852 and 853)
tiniial series of inler mediate mutations.
Lower Pleistocene of
\\ third suju'rior molar found in the The teeth figured (Fig. 852) and determined by Smith Wood-
Shansi, China, has been compared by Dr. A. Tindell Ilopwood ward as "Stegodon," as pointed out by Freudcnberg (1915) and
(1935.1, p. 88, PI. viii) to Archidiskodon planifrons. — Editor.] Matsumoto (1918), probably belong to Elephas [
= Archidiskodon]
Ascending Mutations of A. planifrons in India and in planifrons. These and other fossil mammal remains in the Pilt-

Wkstkkn Eurasia (Osboun, 1928). The above conclusion of— down gravels may be divided geologically as follows
Deperet and Mayet that Archidiskodon planifrons passes by a con-
Pleistocene(?) Cervus elaphus ref.
tinuous scries of ascending mutations into A. nieridionalis agrees
Ca.^lnr (?) fiber rof.
entirely with that independently reached above by Osborn in the
II ippopotamus an>phibius (?) ref.
measurement and analysis (Tables VI and Vll) of the sixty-six
specimens in the Falconer and Cautley and Barnum Brf)wn col- Pliocene(?) Koanthropus dawsoni tyjjc.

lections of the Pinjor horizon. Upper Siwaliks, India. The A. Archidiskodon planifrons ref., "Slegodon sp." of
planifrons ridge formula; of the specimens of C'hagny, France, Smith Woodward.
agree closely with the tyjiical A. planifrons ridge formula* of the Ananciis arvrrnensis ref., "Mastodon" of Smith
sjjecimens from the Pinjor horizon, India. This was undoubtedly Woodward.
a case of indejjcndent contemporaneous progression in India and Egiiiis slenonis (?) ref.

in France. Rhinoceros etruscus ref.

'[Upper Pliocene to Lower Pleistocene (see Vol. I, fig. 413). Sec also footnote on p. 950 above. — Editor.]
 : : —

THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 965

Authorities differ as to whether the Piltdown gravels, contain- PI. 26, Fig. 3 und 4, kiirzlich aus dem Belvedereschotter von Wien
ing Ennuthropus dawsoni, are of Upper Pliocene or of Pleistocene durch Schlesinger abgebildet wurden. Ferner fanden sich mehrere
age, as shown in the following citations from Smith Woodward and Ziihne von Castor fiber, der nach Newton (The Vertebrata of the
Freudenberg Pliocene deposits of Britain, PI. v fig. 16) gleichfalls im Red Crag
Smith Woodward (1913, p. 123).^"It is clear that this vorkommt. Ferner ein Fragment von Mastodon arverncn.sisf, von
stratified gravel at Piltdown is of Pleistocene age, but that it con- Hippopotamus major und von Rhinoceros etruscus?"
tains, in its lowest stratum, animal remains derived from some —
History. (1) Freudenberg (1915, p. 420) pointed out that
destroyed Pliocene deposit probably situated not far away, and the molar crest fragments (Fig. 852) found in the Piltdown gravels
with the type of Eoanthropus dawsoni resemble Archidiskodon
planifrons rather than the "Stegodon sp." to which Smith Wood-
ward referred them (Woodward, 1913, pp. 139, 142, 144, PI. xxi,
figs. 2, 2a, 3, 3a). (2) Matsumoto observed (1918, p. 55): "But,
judging from his [Woodward's] figures, the real reference of his
'Stegodon sp.' appears to the present writer, as well as to Freuden-
berg [Footnote: 'Neu. Jahrb. f. Min. Geol. u. Pal., Bd. I., Heft 3,
1915: —Freudenberg, Woodward's papers on Eoanthro-
referating
pus dawsoni, has pointed out that. Woodward's 'Stegodon sp.' is not
referred to genuine Stegodon but to Elephas cf. planifrons.'], to be
otherwise than that stated by Woodward. Woodward's material
which the ridges appear
consists of several fragments of molars, of
to be too high and too narrow and the valleys too deep and too
narrow to be referred to the Stegodonts. Woodward has compared
it with molars of E. meridionalis but not of E. planifrons. The
present writer suspects that, one compare Woodward's material
if

with molars of E. planifrons and of the Stegodonts, one may easily

recognise its closer resemblance to the former rather than to the
latter." (3) As remarked by Matsumoto (op. cit., 1918, p. 56):
"As to Woodward, his Mastodon sp. and 'Stegodon sp.,' of which
association with thesehuman remains may perhaps be secondary,
have very probably been due to earlier Tertiary strata; and — —
the former species is probably to be referred to M. arvernen.'iis.
Now, the age of European E. cf. planifrons is included in that of
M. ai-vernensis; so that it is highly possible, that these two species
are found from one and the same deposit either primary or second-
ary." (4) This determination by Freudenberg and Matsumoto is
in accord with the discovery of Keid Moir that the flint imjilements
found in the Upper Pliocene Red Crag of Foxhall are similar to
those found in the Piltdown gravels.
AnCHIUISKODON PLANIKKONS OF PiLTDOWN (SuSSEx)
Conclusions as to Geologic Age (Osbohn, .Ii'ne, 1928).
Fig. 852. Fragments of a molar tooth probably referable to Archidiskodon
planifrnns. After Smith Woodward, 1913, PI. xxi, figs. 2, 2a and 3, 3a, Smith Woodward and other high English authorities (1913) re-
natural size. Found with the remain.s of Eoaiilhropus dawsoni in the Piltdown garded the two probable Pliocene species, Archidiskodon planifrons
gravel.s, Piltdown Common, Fletohing (Sussex), England. ref. and Anancus arvernensis ref., as washed in from an older
Vertical section of supposed portion of molar representing the valleys Pliocene deposit; whereas the Piltdown skull, Eoanthropus
between the superior (2, 2a) sixth and seventh or seventh and eiglit ridge-
dawsoni, was regarded as of the same geologic age as the Piltdown
plates and inferior (3, 3a) third and fourth ridge-plates (sec Figs. 853 to 855 C).
gra\'els, an age which has not yet been positively determined, and

which may also include Castor fiber ref.. Hippopotamus amphibius

consisting of worn and broken fragments. These were mixed with and Cervus elaphus
ref., ref., properly belonging in the Pleistocene.
fragments of early Pleistocene mammalia in a better state of pres- Osborn observes
ervation, and both forms were associated with the human skull (1) The dark-colored skull fragments of Eoanthropus dawsoni
and mandible, which show no more wear and tear than they might are of Pliocene age, if washed in from the same geologic stratum as

have received in situ." Arrhidi.^kodon planifrons and A. arvernensis. (2) On the other
—
Freudenberg (1915, p. 420). "An der Basis fandcn sich hand, these skull fragments are of Pleistocene age, if washed in
Schiidelund Unterkiefer von Eoanthropus Dawsoni in unzusam- from the same stratum as Hippopotamus amphibius and other
menhiingenden Fragmenten. Im selben Lager zwei Zahnfrag- Pleistocene fossils. On this point Hopwood inclines to Pliocene
mente eines etwas gerollten Molaren von Elephas cf. planifrons age; he writes (letter, June 4, 1928):
(wohl nicht Stegodonl), wie solche schon friiher aus Red Crag von "Secondly, I am inclined to put the skull with the older fauna,
Leith Adams in A monograph on the British fossil Elephants, and the Eolithic culture. To put the skull with the older objects
960 OSBORN: THE PROBOSCIDEA

presents some difficulties, but I is much to be said

think that there stream entire, and that for some reason or other the facial skeleton

in favour of such a step. remains are of the same dark

First, the became detached. The brain-case was lying apart from the face
chocolate colour'" as the other fossils in the same group, whereas the and the whole lot was smashed [by the workmen] when the gravel
younger fauna is represented by fragments of a light ochraceous was being dug. It is noteworthy that the nasals were found in the
colour. This is not very strong evidence I admit, but it would be disturbed gravel, and this would account for the fact that the
strange to find one dark specimen where all the others are light maxillse and other facial bones were missing. I do not pretend that
The find of the nasal-bones and turbinals i)resents the most serious this is the whole story, but it seems to me that these are little

difficulty to be met. They were found lying together in the same circumstantial details which help to strengthen the case for putting
spot. My own idea is that the skull had been rolling along the the skull with the older set of fossils."
[The foregoing text on Archidixkodon jilauifrons of the Pilf-

Longitudinal sec tion down gravels was written by Professor Usborn in June of 1928.
In the autumn Henri
of 1933, at the request of Prof. Dr.
Crown yiew of two ridqe plates
du Professeur Begoucn, Professor Osborn
Delsol, Secretaire, Jubilo
j)repared an article for the volume in celebration of the 70tli
birthday of Comte Henri Begouen entitled "The Geologic Age of
the Piltdown (Eoanthrnpus) and of the Trinil {Pilhecanlhropus)
Man," (?publishcd)'- in which he advanced certain evidence in
support of the new hypothesis that Eoanthropus is of closing

Cross
section
AnCHlDISKODON PI-ANIKRONS OK PiLTOWN, EnGI.AND, AND Ol' THE PlNJOR HoUIZON, InDFA, EXIUBITINO AI'IMIO.XIMATELY THE SAME EnAMEL lilUGE-I'LATE
Height, namely, 71 mm. Natural size.
Fig. 853. Eneland. lldllcd frjiKiiK'nts of Archidiskmlon molars (Brit. Fig. 854. India. Sixth, .seventh, an<l eighth superior ridge-plates of the
Mils. E595 lower, E(i'22 upper), i)re,sumably portion.s of molar repre.sontiiiK Siwalik lectotype of lilrphns [— Arrhidiskodon] idanifrons reproduced natural
llic between the .superior (B, Bl) sixth and seventh or .seventh and
valley.s size from drawing by Miss O. M. Woodward (Fig. S.").")D), for direct eonipari.son
eiKlith ridge-platesand inferior (A, Al) third an<l fourth ridge-plates, from with the corresponding ridge-plates (Fig. 853) of A. phinifroitJS from the
the Piltdown gravels, Sussex. Redrawn natural size by Mi.ss (!. M. Woodward, Piltdown gravels.
July, 1928. Longitudinal and ero.ss-seetions; erown vicw.s. Compare figures
852 and 855.

'[Doctor Hopwood in a later palter ("Fo,ssil Elephants and Man," I, 193.'), |). J8) mak(\s the following statement:
Proc. Ccol. Assoc, XIA'I, Pt. "Con-
cerning the colour, reference should here be made to a recent paper by Smith Woodward (1933) in which he .states that Mr. Dawson soaked the
Sir Arthur
first pieces in a solution of bichromate of pota.sh to harden them! The remaining jiieces were not so treated and retain heir original colour. This explains the
I

very dark ehoi-olatc tone of parts of the brain-ca.se in contrast with the lighter, .slightly more grcyi.sh colour of the remainder, but it di)i's not afTect the statement
that the colour of the human remains as a whole agrees with tliat of the fossils in Group A rather tlian with those in Group B. The fragments of Piltdown II.
confirm this statement." — Editor.)
-[Sec Bibliography under Osborn (193G.951), at close of this Volume. — Editor.)
 —

THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 967

Tertiary age, while Pithecanthropus is of middle Quaternary age The following is a citation from Professor Osborii's article
evidence based chiefly on the ritige-plate and ganonietric method, (forwarded to Dr. Delsol in October, 1933), which, to the best of
which consists of counting the number of ridge-plates and esti- our knowledge, gives his final ojjinion on the subject in ciuestiou:
mating the enamel length of proboscidean molars. While Dr. "The conclusions which may now be drawn from the ridge
Eugen Dubois adhered to his original opinion that Pitheranthropus ]jlate and ganometric and from other palseontological methoils
is of Upper Pliocene or Lower Pleistocene age. Dr. W. O. Dietrich of com]jarison are as follows:
and Professor Osborn independently reached the conclusion that "First, Pithecanthropus crectus, by association with Stegodun
there is strong e\'idence for assigning to Pithecanthropus a more airdwana with a ganometric measure of 482 mm., also with an
recent geologic age, namely. Middle Pleistocene, owing partly to
the associated remains of Stegodoii airdwana and Elephas [Palscolox-
ARCHIDISKODON PLANIFRONS Amer. Mu5. f982l Ref.
odon] hysudrindicus in the Trinil beds. As to the Piltdown fossils
and flints, Professor Osborn regarded these remanics, or worked
over siiecimens, as having been washed in from older horizons,
belonging in two groups, as clearly shown in the coloring of the
\ery fragmentary fossils.

Type
Brif. Mas. M.3068
-
__, Ri^hf Second
"^ 1 1 Superior Molar
enamel

Line of wear
.'® 6 P e e e 6/-r'i

'"XTpll il^;' ARCHIDISKODON PLANIFRONS

Amer Mas /995I Ref

Fig.855. Comparative sections of the lectotype and referred Archidistcodoti planifrons molars, with the A. planifrons molar fragments of Piltdown (solid black).
Reduced to a uniform one-third scale for comparison with Falconer's figure (Fig. 825).
\, Referred eleven ridge-plated r.Mj (Amer. Mus. 19798), 203 mm. in length, maximum 10th ridge-platc height 61 mm., a small and very primitive
molar.
B, Referred third right inferior molar, r.Ms (.\mer. Mas. 199-51), 313 mm. in length, maximum 9th ridge-plate height 113 mm., consequently a large and
progressive lower molar stage. Exterior of the same tooth (Fig. 84.5).
C, Second left superior liiolar, l.M'' (.\mer. Mus. 19821), 220 mm. in length, maximum 7th ridgr-plalc licight 74 mm., somewhat exceeding the 71 nun.

Piltdown specimen.
ridge-jilate height in the
D, Falconer's type of Elephas [Archidiskodon] planifro7is (Brit. Mus. M.3068), an r.M', 221 mm. in length, maximum 7th ridge-plate height 71 mm.
Redrawn by Miss G. M. Woodward for this Memoir (cf. Figs. 828 and 825).
968 OSBORN: THE PROBOSCIDEA

Elephas hijsudritidicus Rrinder of an estimated measure of 5459 Sci., Paris, Tome 179, p. 1418, December 15, 1924. Type.—
mm., proves to be of Middle Pleistocene age." Fragment of a third inferior molar of the left side, I.M3, in the
"Second, by association with Arrhidiskodon planifrotis with Laboratory of Geology, University of Bucharest. Hohizon
eleven ridge jjlates and an enamel length of 1113 mm., Eoanlhropus AND Locality. — Upper Pliocene. Tulucesti (Covurlui), Rumania.
dawsoni of the Piltdown gra\els may pro\e to be of Upper Pliocene Type Figure. — Sabba Stefanescu, 1927; originally figured by
age." Sava Athanasiu in 1912 [1915], PI. xvii, fig. 4, as Elephas cfr.

"Third, thus, by the application of this ridge plate and gano- mfridionalis.
metric method these two famous fossil men change places, as Type Description.— (S. Stefanescu, 1924, p. 1418, and 1927):
follows: "A I'exception de Maria Pavlow . . qui a rapporte a VElephas
"The supposed oldest fossilman, Pithecanthropus, is dethroned af. plauifrnns une molaire qu'elle a regue de Ferladany (Farladeni)
and becomes a survival; Eoanlhropus is enthroned as the oldest en Bessarabie, personne a ma connaissance n'a indique jusqu'a
fossil man knoivn up to the present time." — Editor.] present cette espece dans les couches geologiques de Roumanie. Or

Fio. 856. Scene on the Ancient Riveh Ouse Illu.stkating the Osborn Theory of the Upper Pliocene Age ok Eo.^nthropus uawsoni
Restoration by Margrct Flin.sch in 1934, under the direction of Henry Fairfield Osborn
Eoanthrnpna (/(/»woni Woodward, a.s restored from the cranium and jaw discovered in the Piltdown gravels dej)ositcd in tlu^ cluiniic!l of tJie ancient rivoi'
Ou.se. Bodily proportion.s and outlines from a Bushman hunter (.j feet) of tlie Kalahari Desert, We.st Africa, reduced in present restoration to a scale of
1 "JOth natural size. The erect figure is holding tlic sliar|)ened femur of Anaricus or of Arckidinkixloii. discovered in flic Piltdown gravels. Osborn does not
agree with his friend Henri Breuil tliat this sharpened bone is the work of a rodent.
The mammals are (left foreground) ArchiiHxknilim plntiijrons, l/;59tli natural size, (left rear across the riv<'r) the straight-tusked mastodon, Anancus
arvernensis, and Archidiskodon planifrons, 1 /100th luitural size, (right bank) herd o! JCqims stoionis, 1/lGOth natural size.

ARCHTDISKODOX PLANIFRONS OF RUMANIA l)lusieurs molaires et ([uelques nioitios de mandibules d'elephants

Archidiskodon planifrons rumanus S. Stefanescu, 1924 de ma collection jin'senfeiit d(>s cnraclcn^s tjui rMi)])elI('iit de tres
Figure ,S.")7 |)r('s cetix de V Elephas planifrons. Par consctiuent, je puis afhrnier
From Tulucesti (Covurlui), Rumania. Upper Plioceue. (|ue cette espece est non seulement rei)resentcc mais (lu'clle est
Elephas antiqiuis rumanus Stefanescu, 1924. "Sin- la i)rcsence I'utie (les plus rcpandues dans les couches pliocenes et pleistocenes
de VElephas planifrons et de trois mutations de VElephas uutiquus de notre pays. Mais n'ayant jias rcHUieilli nioi-iuenie ces fossiles, il

dans les couches geologiques de Roumanie." ( 'ompt Hend. Acad. . lue ])arait hasardeux de pr^ciser leur age geologique. . . . D'autre
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 969

part, vu grande variabilite des caracteres specifiques de VElephas

la a une mutation ancestrale de VElephits (uili<iuus, que j'ai ilononuiiee
planifrons, il est absolument necessaire de ie\iser ces caracteres rinnanus."
afin lie conclure, s'il y a lieu, a I'existence d'une seule ou de "Je n'insiste pas pour le moment sur les caracteres de cette
plusieurs mutations, ou meme a I'existence d'une seule ou de plu- mutation qui, a mon avis, est la plus rapprochee de I'origine
sieurs especes, confondus ensemble par les paleontologistes qui mastodontide de I'espece anliquus. J'ajoute seulement que la mu-
m'ont precede. Dans ce but je i^ratiquerai ?na methode de recherchefi tation ausonius de I'ltalie lui succede et (jue la mutation de Weimar
basee sur I'organisation des lames isolees des molaires,methode et Taubach (Allemagne), que je designe sous le nom germanicus,
naturelle ciui m'a permis de reconnaitre que la
tout recemment est la plus recente."
molaire provenant de Tulueesti (C'ovurlui) et rapportee par Sava "Toutes ces trois mutations ont ete trouvees en Koumanie:
Athanasiu [Footnote: 'Mamifere pliocene de la Tiducesii, districtul rumanus a Tulueesti (C'ovurlui), avsonius a Colintina (Ilfov),
Covurhii, 1915.'] k VElephas cf. meridionaUs, doit etre attribuee germanicus a Tanganu (Ilfov)."

Type and Referred Inferior Molars of Archidiskodon planifrons rumanus

Fig. 857. (Left) Type third left inferior molar, I.M3, of Elephas anliquus rumanus S. Stef3,nescu, 1924;
originally figured by Sava Athanasiu, in 1912 [1915], PI. xvii, fig. 4, as "i'fepfeis efr. »nen'(HonaZis"; refigured
by StefSnescu in 1927 as Elephas anliquus rumanus. Original in the Laboratory of Geology, University of
Bucharest. (Right) Last two ridge-plates and talon of a referred third right inferior molar, r.Ms, stripped of
its very thick cement (ColL of the Laboratory of Palaeontology, University of Bucharest). After Stefanescu,
1927.
The present author considers that tliis type is more closely related to Archidiskodon owing to tlie com-
paratively low ridge-plates and the excessively wide cement areas between them. StefSnescu rightly attributes
this specimen to Elephas = .-irchidiskodon] planifrons.
[ He states (1924, p. 1418): "Or plusieurs molaires et
quelques moitiiis de mandibules d'alephants de ma collection pr(5sentent des caracteres qui rappellent de tres
pros ceu.x de VElephas planifrons. Par consequent, je puis affirmer que cette espece est non seulement repre-
sentee mais qu'elle est I'une des plus repandues dans les couches pliocenes et pleistocenes de notre pays."

ARCHIDISKODON MERIDIONALIS AND A. MERIDIONALIS CROMERENSIS

Archidiskodon meridionalis Nesti, 1825 attributable to the fluviatOe waters of the Arno, which at some
Figures 815, 817, 858-869, 925-928, 1239, PI. xxi ancient period poured forth from natural cataracts having their

Upper Lower Pleistocene of Val d'Arno suporieure, northern

Pliocene' to source in the mountains between Incisa and Rignano. If it is true,

Italy, of Saint-Prest, France, and of England (Forest Bed). howe\'er, that ribs of Fiseter [Physeler] were found two miles from
Syn.: Elephas giganteus Aymard (MS. in Lartet, 1857); Elephas lyrodon Arezzo, then there is reason to believe that the deposits of the Val
Weithofer, 1890, ba.sed on a female type skull. d'Arno superieure were not of fluviatile origin.
Early Discoveries of Elephas meridionalis by Tozzetti Falconer in the "Palaontological Memoirs" of 1868, Vol. II,
IN —
THE Val d'Arno. Giovanni Targioni Tozzetti, who in 1725- p. 104, remarks that "The 'Val d'Arno Superiore' has, from remote

1745 traveled through the greater part of Tuscany, described ages, been celebrated for the vast abundance of fossil remains
especially the geology of the valley of the Arno, making a sharp found there. Huge bones and teeth of Elephants were especially
distinction between the Lower (Val d'Arno inferieure) and the numerous. A large collection of these was formed by Targioni
Upper (Val d'Arno superieure). The 'Val d'Arno inferieure' he Toretti [Tozzetti], which ultimately found its way into the Grand
regarded as a sea-water deposit, which formed the hills and filled Ducal Museum at Florence; and numerous additions were made
them with marine fossils; these fossils occur in abundance from by Nesti, who, in 1808, soon after the publication of Cuvier's
Capraja to the sea. The 'Val d'Arno superieure,' with its hills of 'Memoir of the Mammoth' (Annales du Museum, torn, viii.), ex-
chalk and ochreous clay, weathered by sand and ice, shows no amined the Tuscan Elephantine remains, and was so satisfied of
trace of marine animals; these deposits, Tozzetti believes, were their difference from those of the Mammoth, that he proposed for

'Possibly Lower Pleistocene (see footnote on p. 1049). — Editor.

 970 OSBORN: THE PROBOSCIDEA

tliem two specific flesignalions, namely, Elephas meridionalis and of sjiecific distinction. This singular conclusion is, in some
. . .

E. minutus (or mininuts].^ by the fact that Ciuier luid

Influcnceil measure, explained by the fact that hardly a specimen of a molar
laid so much stress ujion the peculiar form of the lower jaw, and of the true Mammoth exists in the Florentine Museum for com-
guttered beak of the symphysis, as distinctive marks of E. prinii- liarison. It is, perhajts, still more remarkable that the ex])prienced
genius, Nesti (not a professed anatomist) was naturally led to eye of ("uvier should have glanced o\er the multifarious evidence
direct his attention, in the first instance, chiefly to the same parts sujiphed by the Tuscan collections, without being convinced that
in the Val d'Arno remains. Unluckily the specimen that presented E. meridionalis was a well-founded species, considering the ra])idity
the most pronounced beak had lost molar teeth [Nesti, 1808,
its with which he seized, anil the logical precision with which he char-
Tav. I, figs. I and ii]; Nesti assumed to be of an Elephant. But
it acterized, the distinctive marks of the Mammoth from the existing
this selected 'piece justificative' for his Elephas meridionalis was Indian Eleijhant."
l)roved by (
'uvier to be the lower jaw of Mastodon Anwrncnsifi, . . . "Failing the teeth, Nesti drew his specific distinctions from the
and E. iiinndiis to be merely a young Elephant." form of tlic ci'aiiiuin and lower jaw. Ample evitlence is afforded by

Lectotype Cii.iNinM (C) of Archidiskodon meridionalis, Museum of I'lukencb

Fig. 858. Lectotype figure after Nesti of the first described mid figured cranium (Cranium C) of Elephas meridional^ Nesti, 182.5, Tav. i, figs. 1 and 2;
cited by Falconer (1868, Vol. II, p. 122) and selected as the Ujpe by Deporct and Mayet (1923, p. 12G). Oiie-twelftli natural size. Compare figure 861 of the
present Memoir, also figure 865 (13).

"After a long interval, during which ('uvier had visited the them for establishing E. meridionalis as an independent form."
Tuscan brought out another memoir upon the
collections, Nesti "The Abbe Croizet, to whom palaeontology is indebted for so
subject |1825|, in which, ujjon greatly extended obser\ations on much \-aluable research on the fossil fauna of Velay, was the first
s|)ecimens of all ages, from the foetus upwards, including crania, who had the courage to question the decision of ("uvier against
lower jaws, molars, tusks, and bones of the extremities, he upheld E. meridionalis. In his work upon Puy-de-D6rae, he has figured
the soundness of his first inference in regard to the distinctness of and described a fragment of an ui)per (?) molar (lower left of
E. meridionalis, while he admits tacitly the force of Cuvier's Croizet and .lobert) discovered at Malbattu. ... He refers to
criticism upon his second species, E. minulus. . . . Another circum- Nesti's researches, and sums u]) by inferring that, as there are
stance, which materially damaged the authority of Nesti upon two living Elephants, so there were two fossil species the one with —
a question of such difficulty and imi)ortance, is that he states that, attenuated plates, being the Mammoth of Siberia, the other with
after examining a vast number of molars of all ages, he had found thick plates, as seen in specimens from . . . the \'al d'.\rno. He
—
them to vary so much some having thick i)lates, others thin, considered the facts sufficient, but a.ssigned no other name to the
"
and the same tooth presenting such different {jatterns, according to second species than that of 'Ek'])hant de Malbattu.'
its age and degree of wear —
that he had abandoned the characters Geologic Level, Fouest Bed oh ("komerian (compake
yielded by the molar teeth as worthless ( !) for any reliable marks OsBORN, 1922.563, pp. 439, 440.— (1) The survival in the Forest
'[Re-searches of the present author failed to substantiate the assignment by Nesti of the names Elephas minutus or E. minimus in either of his articles of
1808 or 1825. Weithofer (1890.1, p. 134) attributes both these names to Falconer, i.e., Elephas minulus (Pal. Mem., 1868, V'ol. II, p. 104) and E. minimus
(1846, letterpress, "Fauna .Anticjua Sivalensis," p. 13). —Editor.]
 ;

THE MAMMONTINiE: ARCHIDISKODON AND METARCHIDISKODON 971

Bed of East Anglia, above the Red and Norwich Crags {Archidisko- Specific 'haracters of Archidiskodon meridionalis. As
( —
don planifrons level) ofwarm temperate types, such as specimens compared with Elephati = Archidisko(l(in\ pUuiifnnts, cranium of
[

referred to Arrhidiskodon meridionalis, Pala'olojodon [Hesperoloxo- Arrhidiskodon meridionalis larger, more lofty or acrocephalic;
don] (uiti(/uus, Hyxi/a tttn'otci, Dicerorhinus etrufscuH, Equus stenotiis, frontonasal contour ty])ically concave instead of phiiw (Fig. 861)
Machxrodus, is exactly paralleled by the same genera and species fronto-oceipital crest broadly truncated (Fig. 865) not rising to an
occurring in southern France and northern Italy (Val d'Arno) acute a])ex as in ^4. imperator. Maximum ridgc-jjlate formula:
Dp 4| M 1/+ M 2|f| M 3j-j.T3 a^^ compared (p. 933)
with the typical ridge-plate formula of A. planifrons
(after Falconer) of Dp 3 §+ Dp 4 y| 1?^ 2f M M
M 3 ij^. Crinding teeth broad, heavily plated with
cement; superior incisive tusks large and lyrate.
Another progressive character in A. nwridionalis is the
loss of true premolar (P 3-4) eruption. In brief, the

Young Male Cotype Cranium (A) of Archidiskodon meridionalis Ne.sti, Museum of Florence
Fig. 859. Cranium A of Nesti, cotyjic of Elephas meridionalis Nesti, Fig. 860. Side view of the restored cotype skull of Elephas niendionalis
182.5, Tav. i, fig. 3, one-eighth natural size. Figure reproduced from a plate in the Florence Museum. After Weithofer, 1890, Taf. i, fig. 2: "Elephas
which also gives the profile (fig. 2) and front view (fig. 1) of the lectotype meridionalis Nesti; Cranium A; oberes Arnothal; von links." Reproduced
specimen (Cranium C). about one-tenth natural comparison with figure 859, which
size, for direct is

After this cranium (A) was figured, as above, the incisive alveoli were one-eighth natural size, .showing that the peak or apex of the cranium is

added, as shown in figure 860 oppo.site. Falconer (1868, Vol. II, p. 122) greatly elevated. Consequently the frontojiarietal plane is much more
observed that enormous tusks have also been atlded, yielding a diameter of elongate than that in E. planifrons.
0.26 m. or 10.2 inches.

during the long warm 1st Inlerglacial period. (2) During this typical A. meridionalis of the Ipper Pliocene, lowermost Pleis-
period there also appear for the time in Great Britain certain
first tocene, and Lower Pleistocene is much more progressive than the
African types, like the Hippopotamus, and there became more typical A. planifrons.
abundant in Great Britain the loxodontinc type, Palseoloxodon This is the great 'southern mammoth,' well named E. meri-
[Hesperoloxodon] antiquus, as well as the Hyxna. (3) The Forest dionalis by Nesti connected with Elephas planifrons
in 1825. It is
Bed arri\'al of tundra and northern forest types, such as Mani- by a series of ascending mutations, completely confirming Fal-
monleus primigenius, Ovibos inoschatus, Aires latifrons, is a dis- coner's observations of 1863, p. 80 "The nearest affinity, and that
:

tinctive feature of the northern latitude and cold climate of East a very close one, of the European Elephas meridionalis is with the
Anglia during the period of the first Scandinavian glaciation, Miocene E. (Loxod.) planifrons of India." Nesti's type is a fine
which has no ])arallel in southern France or in northern Italy. skull still preserved in the Florence Museum.
 972 OSBORN: THE PROBOSCIDEA

El. [Elfphns] meri(li.»ial/s Nesti, 1808, 1825. "Di Alcvine FALCONER'S NOTES OF I8C8 ON TYPE AND REFERRED SPECI-
Ossa Fossili di Maiiiinifeii chc s'incontnvno nel Valdarno," Ann. MENS OF ELEPHAS [ARCHIDISKODON] MERIDIONALIS [AND
Mils. Imp. di Fisica e Storia Nat. Firenze, 1808,
?PLANIFRONS] OF THE NORWICH CRAG AND VAL D'ARNO
I (description
Falconer, "PaliEontological Memoirs," Vol. I, 1868, pp. 443-447,
without name); "Sulla nuova specie di elefante fnssile del
Plate xiv.b' of the "Fauna Antiqua Sivalonsis"
\'aldaino" (letter from Nesti to Dott. Prof. Ottaviano Targioni
Falconer's observations collected in his "Palaeontological
Tozzetti). Nuov. Giorn. Lett., 1825, XI, No.
24, p. 211. Lectotype. Cranium with — Memoirs" extended from the year 1863, in

third superior grinding teeth, the rostrum and tusks which he published a ridge formula of the
first

species Elephas mer/dioiialis, up to the year

being broiven away ( = Cranium C of Nesti) Cotype .

1868, his final observations being embodieil

( = Cranium \ of Nesti). Horizon and Locali-
ty. — Sui)eri()r Val d'Arno, Upper Pliocene, northern by Murchison in the "Palaeontological
Italy. Lectotype Figure. — Op.
rit., 1825, Tav.
Memoirs" of that date. He closely
I, and 2. Cotype, Tav. i, fig. 3.
figs. 1
i compared specimens from the Nor-
—
Description. In his description of 1825 Nesti wich Crag (in the Norwich Mu-
compares the skull with that of "Mastodonte" and seum), and from the Val
of "primigenia," concluding (p. 211): "Potrebbe a
questa Specie imjjorsi nome di Elephans Valdar-
il

nen.si.s, o Etruscus, anco Italicus, ma poiche le

o
regioni, nelle quali questo animale viveva, non sono
note, e d'altronde pare che fosse destinato a climi piu
temperati e meridionali della Specie primigenia, pre-
ferisco di appellarlo El. tneridionalis." There is an
earlier reference to this fossil which does not in-
clude the name. It is to be found in the Annali
del Museo Imperiale di Fisica e Storia Naturale,
Firenze, Tome I, 1808, mentioned above with title
of the article.
Lectotype Skull. Cranium C' — of Nesti's description was
selected by Deperet and Mayet (1923, p. 126) as the type; see also
Falconer (1868, Vol. It is here reproduced from Nesti's
II, p. 122).

original figures, frontand side views, figs. 1 and 2 of his Tav. i

(Fig. 858 of the present Memoir). Falconer {op. cit., p. 122) de-
scribed Cranium C as follows: "It is nearly perfect in the frontal
and occipital regions, condyles, maxillaries, and molars, but im-
perfect in the facial portion, the border of the nasal opening being
broken, together with the terminal portion of the incisive alveoli
Aged Male. Lectotype Skull (C) ok Archidiskodon mehidionalis
and the zygomatic arches. Since Nesti's figures were taken, this Nesti, Museum of Florence
specimen has suffered considerable damage, the upper lamina of Fig. 861. Lectotype .skull of Elephas meridionalis Nesti, as refiguretl by
the right incisive alveolus having disappeared, together with the Weithofer in 1890, Taf. ii, fig. 1: "Elephax mrridionalis Nesti; Cranium C;
salient tip of the nasals and the lateral margin including the left
oberes Arnothal; von rechts." One-seventh natural size. Compare figures
8.58 and present Memoir.
86.T (7, 13, 14) of the
orbit. The last molar is present on either side, far advanced in
An same skull, after Falconer's plate of 1847, appears in
outline of the
wear. (See PI. i. fig. 11, and PL ii. fig. 16.)" figure 86.5 of the present Memoir. All these figures agree in the peculiarly
Cotype Skull. — Falconer also described the cotype Cranium flattened appearance of the summit of the aged occiput, which is quite unlike
A as follows (op. cit., p. 122): "4. The cranium A of Nesti's the acrocephalic occiput of the juvenile Archidiskodon imperalor.
references, fig. 3, comjjrising the ])alatine, maxillary, and temporal
regions, the inferior part of the occiput,
and the zygomatic arches, d'Arno (in the Museum of Florence). In his ridge formula he
the only deficiency being in the facial region. The specimen, undoubtedly confused ascending mutations of specimens belong-
which is highly ferruginous, has now joined on to it the entire ing to Arrhidiiihodon pkniifrmif: with specimens truly representa-
incisive sheaths (not rei)resented in Nesti's figure) and two enor- tive of ])rimiti\-e Archidiskodon meridionali.'i.
mous which are spread out horizontally in the Theristocau-
tusks, Compare above with the ridge formulae and measurements of
/of/o/i-manner above noticed. Nesti, in his memoir, cites the tusks Archidiskodon plniiifro?!.^. The ridge formula; below gi\-en by
of this specimen as yielding a diameter of 0.26'", or 10.2 inches. Falconer and others embrace the A. planifrons stage as well as the
The last molar, much worn, is present on either side." A . meridionalis stage.

'[A footnote by the Editor of the Palffiontological Memoirs on pp. 443, 444 of Volume 1, states that according to the corrected copy by Falconer in the
Briti.sh Museum, all the figures in PI. xiv.b, except 10, 17,and 18, should belong to E. antiquus; that the correction, however, is incompatible with the descrip-
tion and identification of every figure in pi. xiv.H, given in a subsequent part of the same memoir, according to which every figure in the ])late, with the except-
ion of 16, belongs to E. meridionalis. — Editor.]
 — —

THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 973

Elephas meridionalis Nesti, 1825. Lectotype. Cranium C, — front and back talons, discs of first three wide and open, tendency
Tav. I, figs. 1 —
and 2. Cotype. Cranium A, Tav. i, fig. 3. to mesial expansion, crown low in reference to breadth; figs. 6,
—
Upper Jaws. Plate xiv.b, figs. 1, la, Dp^, ridge-plates K-6-K, 6a, lower jaw, l.Mi, ridge-plates %-8, the crown presents a front
"the crown is composed of six principal ridges, besides front and talon and eight ridges, all of them worn, discs wide and open,
back talons. compared with the corresponding tooth of E.
. . . Norwich Museum; figs. 7, 7a, lower jaw, r.Ms, ridge-plates 7 + ,

(Loxodon) planifrons, which it resembles very closely, ... it has Norwich Museum; figs. 10, 10a, lower jaw from Val d'Arno,
a broader crown"; figs. 2, 2a, r. Dp', ridge-plates 6, Norwich Crag. demonstrating how exactly the English specimens agree with the
—
Lower Jaws. Plate xiv.b,' figs. 3, 3o, lower jaw, r.Dps, Italian, long symphysis, gradual incUnation into a beak; figs.
ridge-plates YrQ-Yi, very broad in the crown relative to the length, 17, 17a, r.Ms, from Val d'Arno, showing 13 ridge-plates, length
discs of ridges very wide as in Italian specimens, Norwich Museum; 10 in. =255 mm., width 3.4 in. =87 mm.; figs. 18, 18a, Norwich,
figs. 4, 4a, lower jaw, l.Dp4, 8 ridge-plates, Norwich Museum, the lower jaw, r.Ms, ridge-plates K^ll-K, "showing eleven principal
ridge formula in these specimens agrees with the Italian, also the ridges, an anterior talon, and [?] a back talon," cement decomposed
broad crown, the low ridges, and thick plates of enamel; figs. 5, or denuded, 4 thick denticles on 6th to 8th ridges, length 11.25 in.

5a, lower jaw, l.Mi, ridge-plates K-8-/2, eight principal ridges, with = 287 mm.

-/^ /<fa^.

FCg:M.

Fig. 862. Molars from the Val d'Arno (left) and the Norwich Crag (right) referred to 'Elephas' [Archidiskodon] meridionalis by Falconer
"Fauna Antiqua Sivalensis," Plate xiv.b, figs. 17, 17a, 18, I80, one-third natural size
Compare "Palsontological Memoirs," Vol. I, 1868, pp. 443, 444, 447, 448
Figs. 17, 17a. Third inferior molar of the right .side, r.Ms, Val d'Arno. Figs. 18, 18a. Third inferior molar of the right side, r.Ma, "Mam-
Original in Oxford Museum, one-third natural size. maliferous Crag" (Norwich). Norwich Museum 1570, one-third natural size.
(Op. cit., p. 447): "Figs. 17 and 17a. Elephas meridionalis. A Val {Op. cit., p. 447): "Figs. 18 and 18a. Elephas meridio7ialis. The finest
d'Arno lower molar of the same age, from Dr. Buckland's collection in the detached molar of this species that has come under my ob.servation is a speci-
Oxford Museum, crown side. (Reproduced in Plate viii of vol. ii.) Length of men which was discovered in the 'Mammaliferous Crag' on the Thorpe Road,
crown, 10. in. Width of crown, 3.4 in. Height of crown, 5. in." near Norwich, by Mr. Prestwich. ... It is now lodged in the Museum at
Norwich, and is the specimen which first convinced me many years ago that
Note I. — Falconer (as explained in the "Palseontological Memoirs,"
the 'Crag' yielded a species of Elephant entirely distinct from the Mammoth
1868, Vol. I, pp. 443, 444), in lettering the plates of the "Fauna Antiqua Extreme length of crown, 11.25 in. Width of crown
and from E. aniiquus. . . .

Sivalensis," became time the plates were lettered that he had

satisfied at the
in front, 3.3 in. Width at fifth ridge, where the crown is broadest, 3.8 in.
interchanged Elephas meridionalis and E. aniiquus. In his Memoir of 1857
Extreme height of ridge, 4.8 in. Width of ninth ridge, 3.5 in. Height of ninth
(not published until after his death) he concluded [Pt. II of his Memoir
ridge, 4.6 in." Reproduced in the 'Pateontological Memoirs,' Vol. II, PI.
publi.shed in 1865, p. 281]: "I beg leave to explain now, that all the plates
VIII.
bearing the name of E. meridionalis in the 'Fauna Antiqua Sivalensis,' includ-
ing the outline-figures of crania in plate 42, belong to E. aniiquus, while those Note II. — Compare these i)rimitive .4. [planifrons] meridionalis molars
that bear the latter name belong to E. (Loxodon) meridionalis. In the descrip- of Italy and England with the more progressive A. planifrons molars of the
tions which follow, they will be cited as such." Siwaliks, India.

'[See footnote on opposite page. —Editor.]

 : :

974 OSRORN: THE PROBOSCIDEA

Falconer, 1868, E. meridionalis: Dp 3^^^" Dp 4^ Weithofer in his collective formula, like Falconer (1868), un-
M 1 vi-e-ti M2 7 4-9 M3 is-i i-vi-
doubtedly erred by including specimens having the typical Arcki-
The above ridge formula is too low, obviously because di.^kodon planifrons ridge formula with those of A. meridionalis;

Falconer included within his 'Elephas' tneridimialis grinding teeth thus this collective formula ranges from the formulae of Archidis-
belonging to Archidiskodon plaiiifrons. IMore distinctive of the kodon planifrons, e.g., M
3 tt, to A. meridionalis, e.g., 3 tI. M
species A. meridionalis, in Osborn's opinion, is the ridge formula Lydekker, 1886. — Lydekker in his "Catalogue of the Fossil
which we may deduce from Falconer's own observations (1868, Mammalia in the British Museum (Natural History)," 1886.2, p.
Vol. II, p. 1 18) on nine Itahan crania in the Florence Museum 107, adopts the Leith Adams formula of 1877-1881 as follows:
(1) The molars of E. meridionalis so closely resemble those of E.
Falconer (Italy), E. meridionalis: Dp 2f Dp 3f Dp 4t
planifrons, that if they both occurred in the same area it is more
M 1 /^ M 2 If: M 3 T^J^. than doubtful they could be specifically distinguished; (2) both
if
Weithofer (1890, p. 173) observes that Falconer's high figure,
frequently exhibit partial denudation of the enamel ridges; (3)
M 3 Yi, rests on a single indi\idual. there is variation in the thickness of the enamel and in the breadth
of the ridges, in some molars the enamel being relatively thin ami
Leith Adams, 1877-1881 considerably plicated; (4) the cranium is characterized by large,

Formula. — Leith Adams reviewed this important species in slightly curved tusks and widely diverging alveoli; (5) in general
contour is intermediate between the cranium of E. planifrons
his "British Fossil Elephants," 1877-1881, and while in general
it

confirming Falconer's observations by excluding E. plaiiifrons, he and E. hysudricus, although nearer the latter; (6) it agrees with the
reached a nuich more accurate dental formula which became the cranium of E. planifrons in the relative distance between the
nasals and the vertex, but has the vertex more vaulted, the
standard
frontal profile concave, the temporal fossae intruding largely on the
Leith Adams, 1877-1881, E. meridionalis: Dp 2f Dp 3t frontal aspect; (7) the species attained an enormous size, the
Dp4fMlf^M2|i}iM3|Hi. height of some individuals being estimated at upwards of fifteen
feet.
This formula of 1877-1881, together with Leith Adams' sum-
mary of the points distinguishing the molars of Elephas ineridionalis
from those of the narrow-toothed Elephas antiquus (p. 232), are to
be found in his important Memoir (p. 208). He assigns the follow-
ing characters to Elephas meridionalis.
Characters. — (Cf. op. cit., p. 30): M^ in E. meridionalis,
ridges nearly as broad as they are long, thick plates, grosser masses
of intervening cement, machaerides uncrimped, X 10 X (Cf p. . .

44) : ]\P massive, enamel and plates very thick, machaerides

scarcely plaited, great breadth and low ridge formula, which rarely
if ever exceeds that of M' of E. antiquus. (Cf. p. 48) : In M= the
highest ridge formula of E. meridionalis equals lowest of E. anti-
quus; so that in number of ridges we find E. primigenius, E.
antiquus, E. meridionalis, and E. namadicus meeting at their
extremes. Leith Adams (cf. pp. 129 to 144) compares the cranium
Archidiskodon meridionalis Ref.
of E. meridionalis in great detail with that of E. planifrons, E.
Fig. 863. Rpferred Elephas meridionalis of Cliagny (Cote-d'Or), one-
hysudricus, E. bombifrons, E. africanus, etc., concluding that the
fourtli natural size, after Gaudry, 1878, p. 178, fig. 237. Tlii.s fourteen ridged
skull and dentition of E. planifrons make the nearest approach to
tliird .superior molar, M^, of Chagny, is to be compared with tlie tliirteeii
E. meridionalis (see pp. 186, 208-210, 239, 244). ridged molar stage of Eh'phas ineridionalis of the Val d'.\rno, i.e., M 3 ,^J -, a.s

—
Error. Originally Leith Adams also made the error of dedueed by Falconer (see Falconer, 18G8, Vol. II, p. 118). Observe superior
including the low ridge-plate formula (M 3 \^) of E. planifrons ridge-j)late.s ])rc-concave, post-convex (ef. Fig. 827, .1. planifrons).

with the high ridge-plate formula (M 3 Hrfj) of E. meridionalis,

in describing the range of evolution and variation in the ridge
formula of the British specimens of E. meridionalis.
Cranial and Dental Characters. — Weithofer (1890, jip.

136, 137) refigurcs Nesti's type skull (Fig. 861 of the present
Weithofer, 1890, Lydekker, 1886 Memoir) showing the right side (Taf. ii, fig. 1): "Elephas meri-
Collective Ridge Formula. The collective ridge-plate — dionalis Nesti; Cranium C; oberes Arnothal; von rechts."
formula [of A. planifrons and A. meridionalis] may also be cited Falconer also rofigures this skull (1846 [1847, figs, xix of Pis.

from Weithofer (1890, p. 172) : "Als (lesammtformel fiir die Ziihne XLii and XLiv|), as well as Depi'ret and Mayet (1923, p. 128, fig.

des El. meridionalis, soweit sie hauptsiichlich aus dem Material 16). The three skulls in the Florence Museum exhibit the follow-
des Museums zu Florenz resultirt, ergabe sich demnach": ing characters: (1) concave frontals, (3) high
Pointed nasals, (2)
Weithofer, 1890, E. meridionalis: Dp 2 7,^3 Dp 3
'.'""
o('cii)ilal crest flattened anteroposteriorly, (4) extreme hrachy-

Dp4^MlHM2f^jM3ii^^ cephalic cranium shortened anteroposteriorly, (5) broad narial

 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 975

openings, (6) parallel sides of the premaxillary sockets of the tusks, SKULLS AND SKELETONS OF ARCHIDISKODON MERIDIONALIS
as contrasted with the broadly flaring sides of the sockets similar In figure 865 are displayed outlines of the lectotype (Nos.
to those in the contemporary Hesperohu-odon antiquus. All 7, 13, 14), cotype (Nos. 8, 9, 11, 11a, 15), and other skulls of Archi-
these characters point to the cranial relationship of the Italian diskodon meridionalis, male and female, of which we ha\e been able
Archidiskodon meridionalis and its ancestor A. planifrons. to find figures in the literature.

They include the following:

1) Male skull figured in error by Falconer and Cautley in 1847

as Elephas antiquus: front view, PI. xlii, fig. xix; side
view, PI. XLiv, fig. XIX. Cranium C of Nesti [lectotype of
Elephas meridionalis Nesti, 1825, Tav. i, figs. 1, 2].

2) Cranium of Elephas meridionalis, side view, after Weit-

hofer. Cranium C of Nesti.

3) Cranium A of Nesti [cotype], palatal view after Nesti,

1825, Tav. i, fig. 3; front and side views after Weithofer,
1890, Taf. I, figs. 1, 2.

4) Front and side views of female cranium, type of Elephas

lyrodon Weithofer, 1890, Taf. v, fig. 1, Taf. iv, fig. 2.

Characters (Osborn, 1924). —

These crania agree in the
following points: (1) Extremely broad and vertically shallow
anterior narial openings, a feature shared by Archidiskodon im-
perator (Fig. 896); (2) extreme cranial abbreviation (hyper-
brachycephaly) and depth (bathycephaly) resulting in hypsicephaly
and acrocephaly (3) concave forehead or frontoparietal profile;
;

(4) i)arieto-occipita] crest rising high in profile (cf. Weithofer,

1890, Taf. I, fig. 2, E. meridionalis cotype, with E. imperalor ref.,

Los Angeles, (5) through hypsicephaly, orbits and occipi-

Calif.) ;

tal condyles approximated, vertical diameters greatly exceeding

anteroposterior diameters; there seems to be little doubt of the
phylogenetic kinship of the Archidiskodon meridionalis with the A.
imperalor crania; (6) cranial profiles of A. meridionalis and A.
imperator analogous to the hypsicephalic and acrocephalic profile
Weithofer's Type Skull of Elephas lyrodon = Archidiskodon Mammonteus
of primigenius; (7) in comparison with the more
meridionalis female
primitive, more platycephalic A. planifrons, with smaller narial
Fig. 864. Types of two individuals of Elephas lyrodon Weithofer,
openings (Fig. 848), the known crania of A. meridionalis are much
Florence Museum, upper Val d'Arno deposits (Weithofer, 1889, pp. 79 and
80): "Hier sei beziiglich der zum erstenmal genannten Species Elephas lyrodon larger and more progressive than the known crania oiA. planifrons.
nov. sp. nui bemerkt, dass sie auf zwei vollstandige Schiidel sammt Stoss-
ziihnen, sowie Schadelfragmenten mit Stossziihnen und mehreren Unterkiefern CRANIA IN THE FLORENCE MUSEUM
und isolirten Stoss- und BackenzJihnen des Museums von Florenz basirt ist."
After Weithofer, 1890, Taf. in, fig. 2 (right); iv, fig. 2, Schadel o (left); v,
(W. D. Matthew, September, 1920). Here are found be- —
sides the tyi^e skull of Elephas meridionalis Nesti [Cranium C]
fig. 1, Schadel a (middle). All one-twentieth natural size. Compare figure
865 (11, 11a).
several more or less incomplete skulls, many jaws and teeth, in-
cluding those from the Upper Pliocene near Magello. A fine
.series of E. meridionalis crania published by Weithofer (1890)
MS. Elephas giganteus Aymard (in Falconer, 1857, p. includes the type of Elephas lyrodon Weithofer, which is obviously
321).— (Lucien Mayet, letter, December 11, 1922): "Elephas a female of Elephas [
= Archidiskodon] meridionalis in which the
giganteus Aymard n'est qu'une designation portee sur des etiquettes tusks are of a lyrate arrangement drawn together at the points,
de sa collection par ce paleontologiste. II y aurait lieu de retrouver more after the manner of Hesperoloxodon antiquus but more slen-
les pieces et de les determiner ; ce serait peut-etre E. meridional- der. In the same beds with these crania of E. [A.] meridionalis was
?'s(?) ou E. lrogontherii{1) ou E. antiquus{?) II s'agit . tres probable- found part of a cranium of the straight-tusked elephant//, antiquus,
ment la comme — pour beaucoup des ossements recueillis par which exhibits a wide spreading of the tusks at the base, a broaden-
Aymard dans le Pliocene superieur et le Pleistocene du bassin du ed rostrum not less than 2}i feet apart at the rim of the socket.
—
Puy de designation inscrite sur des etiquettes, sans que les Contemporary with the above Archidiskodon and Hesperoloxodon
pieces correspondantes aient jamais ete d^terminees ou aient fait are numerous remains of Anancus arvernensis, including a fairly
I'objet d'une revision ulterieure. C'est un nom — celui d'Elephas preserved skull with tusks found in the environs of Florence, one
giganteus —a faire disparaitre purement et simplement." tusk broken off and repointed during life.
 /--,..-.'--,--A
E, PLANIFRONS He'
E. PLANIFRONS Re'-
1846, Pi. X, Fifl. 1 'w*
XVI («v.i F»lc
PLANIFRONS
,

f Ref F«lo. 1M7, PI. XUV, F.o

E. PLANIFRONS Rel
.

F.ic. 1845, Pi. IX

File, 1B47, PI. XUl. FiQ. XVI

the IMIESENT AUTHOR IN THE EXPLANATORY LEGEND OPPOSITE.

Fig. 8G5. Crania of the MAMMONTm;E (1-19) as determined uy
976
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 977

Elephas lyrodon Weithofer, 1889, 1890. — (Weithofer, schen Form ihrcr Stosszahne Elephas lyrodon nov. spec, benannte."
1890, p. 173) "Hier in Florenz befindet sich ein vcillig ausge-
: Matthew, 1920, Osborn, 1924: The type cranium of Elephas
wachsener, leider aber nicht besonders gut erhaltener Schadel lyrodon may be regarded, from the very slender character of the
sammt den beiden Incisiven in situ (Falconer's Nr. 6), ein gleich- tusks, as a female of Archidiskodon meridionalis.
falls sehr altes Pramaxillarfragment mit dem rechten Stosszahn

vollstiindig, dem linken zum grossten Theil erhalten (Falconer's SKELETON OF ARCHIDISKODON MERIDIONALIS IN THE PARIS
Nr. 9), ferner ein j lingerer Schadel sammt beiden Stosszahnen und MUSEUM
dem Unterkiefer, womit weiter auch die Wirbel, Rippen, Schulter- The affinity of Archidiskodon meridionalis to A. irnperator is
blatt und Becken im Zusammenhang gefunden vvorden waren clearly displayed in a comparison of figures 866 (.4. meridionalis)
(Falconer's Nr. 8), ein Oberkieferfragment mit dem linken and 896 {A. The superb skeleton discovered in 1869
irnperator).
Stosszahn und jederseits zwei Backenziihnen in situ, endlich mehre- near the village of Durfort, Gard, is shown herewith (Fig. 866)

re mehr oder weniger vollstjindige Unterkiefer oder Unterkiefer- through a photograph kindly furnished by Dr. Marcellin Boule and
halften; einige Stosszahne, sowie offenbar auch einigc der i.solirten retouched by our own artist.
Backenzahne werden hieher geziihlt wcrden konnen." Weithofer —
Skeletal Characters. Gaudry (1893, p. 12) observes of
(op. cit., pp. 191, 192)observes very close kinship to E . meridionalis this specimen: "L'filcphant de Durfort n'appartient pas a la
in the dentition and ridge formula: M 3 j"]^| . He points out race primitive de VElephas meridionalis, ou les molaires ont un
numerous and skeleton, and
differences in the structure of the skull petit nombre de collines basses, enduites d'un email epais, mais a
then concludes (p. 193) "Ich glaubte daher, durch die angefiihrten
: la race modifice de cette espece, c'est-a-dire au type du Val d'Arno
Umstiinde gezwungen, diese neue Form als eine distincte Species [Footnote: y a des dents du Val d'Arno qui, par leur allonge-
'II

betrachten zu mussen, die ich nach der so iiberaus charakteristi- ment et leurs lames nombreuses, etroites, a email mince, ressem-

EXPLANATORY LEGEND OF MaMMONTEUS, PaRELEPHAS, AND ArCHIDISKODON CrANIA

Key to Osborn's dptprminations in the present Memoir of the crania
illustrated in figure 865.

Fig. 865. The inscriptions on this figure were written before the phylogenetie relationships of these crania were fully understood and before Elephas
was clearly separated from E. primigcnius. The outline figures taken from various authors are arranged in three sections: Upper, including Elephas
Irogontherii
primigenius and E. trogontherii; middle, including E. meridionalis, and lower, including E. planifrons.

(A) Skulls of True Mammonteus primigenius (a); of Parelephas trogontherii (6)

f
2. Typical MammonUus primigenius. E. primigenius ref., after Falconer, 1846 [1847, PI. XLiii, fig. xxiv|.
(a) 'I
1. Mammonteus primigenius. E. primigenius rci., after Pohlig, 1891, p. 384, fig. 120.

[ 5. " " E. primigenius veL, after Pohlig, 1891, p. 384, fig. 120 (rev.).

[ 3. Typical Parelephas Irogontherii. E. primigenius trogontherii (female), after Pohlig, 1891, p. 386, fig. 121.

(6) \ 4. " " " Side view of same cranium as above.

0. " " " E. primigenius rcf., after Falconer, 1846 [1847, PI. xlv, fig. xxiv (rev.)].

(B) Skulls of Archidiskodon meridionalis; (C) Skull of ,\. imperator

7. A rchidiskodon meridionalis. E. meridionalis Icctotype, after Falconer, 1846 [1847, PI. xLii, fig. xixj, Val d'Arno, Italy = Cranium C of Nesti.

i:i. E. meridionalis Icctotype, after Falconer, 1846 [1847, PI. XLiv, fig. xix (rev.)]. Side view of same skull as above.
9. E. meridionalis Nesti, eotype, 1825, Tav. i, fig. 3, Val d'Arno, Italy = Cranium A of Nesti.
8. E. meridionalis eotype, after Weithofer, 1890, Taf. I, fig. 1 = Cranium A of Nesti, Florence Museum.
15. E. meridionalis eotype, after Weithofer, 1890, Taf. i, fig. 2 = Cranium A of Nesti, Florence Museum.
14. E. meridionalis Icctotype, after Weithofer, 1890, Taf. ii, fig. 1 (rev.), Val d'Arno, Italy = Cranium C of Nesti.

Side view of same skull as that figured by Falconer above.

11,11a. VFemale of Archidiskodon meridionalis. E. lyrodon Weithofer, type, 1890, Taf. v, fig. 1, Taf. iv, fig. 2 (rev.).
12. ?Female of Archidiskodon meridioruilis. E. lyrodon ref., after Weithofer, 1890, Taf. vi, figs. 1, 2.

(C) 10. Archidiskodon imperator. E. imperator ref.. Museum of History, Science and Art, Los Angeles, Calif. I'Vom Rancho La Brea.

(D) Skulls of Archidiskodon planifrons, Upper Pliocene of India

16. Archidiskodon planifrons. E. planifrons ref., after Falconer, 1846 [1847, PI. xlii, fig. xvi], Pinjor horizon, India.
" " after Falconer, 1846 [1845, PL x, fig. 1 (rev.)].
17. E. planifrons ref.,

18. " " E. planifrons rcf., after Falconer, 1846 [1845, PI. ix (skull); PI. xi, fig. 3 (jaw)|.
" " after Falconer, 1846 [1847, PI. xliv, xvi (rev.)].
19. E. planifrons ref., fig.

We observe that in its front and side cranial aspects Archidiskodon planifrons is very primitive. In the two crania of .1. mrridionalis shown in side view, we
observe a transition from the yl. /jfam/rons type to the more typical .4. meridionalis, namely, E. meridionalis Weithofer, 1890, Taf. i, fig. 2; this skull is
decidedly hypsicephalic; it strongly resembles in this feature the skull of -4. imperator, also the skull of the typical Mammonteus primigenius; it differs pro-
foundly from the skull of Parelephas trogontherii which has a profile more like that of Elephas indicus.
 Abchidiskodon mbbidionalis of Durfobt, of Cromerian or Nokfolkian, Lower Pleistocene,
Age
de Durfort," as restored and inount,ed in the tialei'ic
I.'ig 866 Skolotoii (largplv restored) of Elephas = Archidiskodon] meridionalis, known as
[
"I'Elcphant
(Gaudry, discovered in 1869 near the village of Durfort, Gard, between Ntmes and Vigan;
de I'al.V.ntologio, Mus.mn, d'Histoirc Naturelle, Paris 1893);
under the direction of Gervais; installed in 1885; fully compared, measured, and figured by
Gaudry m 1893.
exeavati<.nr„m|)letedJune21, 1873; restored
One-thirtietli natural size. Photograph by courtesy of Dr. Mareellin Boule.

December 12, 1921): 'Me n'ai j.as voulu doteriorer ••etle

Doctor Boule writes a,s to the parts restored, or "artifieiels," in this Durfort skeleton (letter of
marquant sur parties restaurfies, mai.s voiei la nomenclature des principaux elements de squelette qu'un cxamen assez somraairc et
photographic en elle Ics
cchafaudages, nous a montro etre arlificicls: Crane.-Parties supencure ct
superficid Ic scul quV.n puisse faire en cettc saison .sans avoir recours a des
Alveoles des defenses, en partie. Extromite ant6rieure de la defense droite. Mandibide droite.-Branehcs montantcs, extr6nut6 de la symphysc.
posterieurc
sont restaurees ,.u en platre. Membres ant6ricurs.-Un
Colomie vertebrale. ^uclques vertebres eervieales, lombaires et caudalcs. Cbtes.-La plupart
3 ou 4 i)halanges. Membres posterieurs.— T^te supiSrieure du tibia droit et p6ron6 droit."
scaphoidc et

Compare figure 868, lateral view of the same skeleton, modified after Gaudry, 1893, Plate.

Ilidgc-platcs (Gaudry, 1893, p. 13): Durfort ont des lames ph.s nombreuscs ct plus n.inccs ,|uc ,-cllcs dc Scniur, dc
"I>es molaires de I'ftlophant de
(pic nous pos.scdon.- au Museum.
ous pi)s,s(Klons
du Monte Verde, pres dc Rome, lOllcs vcsscniliicnt pn'S(iiic
Chagny, dc Perols, prcs de Montpellier, dc Randan
. .
(Allier),
Referred by Gaudry to a progressive stage of Ekphas [=Archi-
autant aux dents de V Elephas (inliquus appelces inlcrmedius tpi'aux dents dc \'E. meridiimalia."
diskodon] meridionalis.

978
 Archidiskodon meridionalis of Durfort, France
Rfsldration by Margrct Flinsfh Buba, May 23, 1930, oiu'-fifticMi natural size
Fig. Primitive staKc, drawn diroctly from skeleton in Paris Museum, represcntin;; an animal of young adult ago. Obs(-rv(' tlic lialf-grown tusk.^,
867.
the proljoscis elevated in order to expo.se the long slender rostrum of the lower jaw (haraeteri.sti<- of this species. Ear drawn .small, of somewhat primitive
contour. Ratlier drooping posterior quarters, draw n from supposed mammontiiic affinities. Digits o in the manus and 4 in the pes, ungues 5 and 4.

Durfort Skeleton of Archidiskodon meridionalis. Lower Pleistocene Age. About one-fiftieth natural .size. Compare Figure 866
Fig. 868. Museum d'Histoire Natureile, Paris (Gaudry, 1893); for details
Lateral view of "I'Elephant dc Durfort" in the Galerie de Paleontologie,
see legend of figure 866, for description see pages 977, 980 of the present Memoir.
This classic skeleton, redrawn after photographic plate in Gaudry (1893), exhibits the right lateral view as compared with the oblique anterior view
introduced in figure 866 above, and enables us to estimate clearly the height of this animal in the restoration (Fig. 867).
According to Gaudry's measurements, the skeletal shoulder height from the ground to the top of the anterior dor.sal spine is 3830 mm. (12 ft. 6^4 in.);
summit of occiput to the ground 4150 mm. (13 ft. 7% in.), as mounted.
According to Osborn's measurements, tlie skeletal shoulder height is 3499 mm. (11 ft. 5?^ in.).
The method of the present author of estimating the skeletal shoulder height is to take the standing height of the forelimb to the top of the scapula. This
measurement was arrived at through the observation of the author, when sitting upon a living elephant, that by placing his thumbs together on the tallest
spine between the scapulse with the extended small fingers resting on the top of each scapula, the spines were found to be on a level with the scapulae, the
latter rising and lowering a few inches above and below the spines in the process of walking. This accounts (in the present instance) for the smaller measure-
ment given by Osborn (3499 mm.) in the skeleton and, after adding the usual 6^ per cent, of 3721 mm. or 12 ft. 2}i in. in the flesh (see restoration to the
same one-fiftieth scale, Fig. 867).

979
 —

980 OSBORN: THE PROBOSCIDEA

blent plus a pelles de VEIcphas auliquits qu'a certaines dents d'Ele- The associated fauna is described by Gaudry (op. cit., p. 14)
phas meridionalis.'] et du Forest-bed ou les eollines commencent as follows: "En realite, tous les os de mammiferes de Durfort que
a se multiplier, a diminuer d'epaisseur, a augmenter de hauteur. nous avons dans le Museum se rapportent seulement a 4 Elephants,
EUes ressemblent presque autant aux dents de VElephas anliquus 4 Hippopotames, 5 Bisons, 4 Cerfs, 1 Rhinoceros, 1 Cheval." The
appelees intermedius qu'aux dents de VE. meridionalis. II me associated florais rich in oak trees (op. cit., p. 17) ". il y a eu : . .

semble en outre que les defenses sont plus courbees et que les os la une veritable foret de ces arbres, comprenant au moins quatre
des pattes sont moins opais que dans les E. meridionalis les plus especes." There were also beeches.
anciens; en cela I'animal de Durfort marque encore une tendance
vers les Elephants quaternaires. Les molaires de 1' Elephant de Archidiskodon meridionalis cromerensis
Durfort ont des lames plus nombreuses et plus minces que celles Deperet and Mayot, 1923
de Semur, de Chagny, de Perols, pres de IMontpellier, de Randan Figures 870, 87 IF
(Allier), du Monte Verde, pres de Rome, que nous poss6dons au
Kessingland, Suffolk, England. Age: Cromer Forest Bed of Norfolk
Museum." East Anglia = Cromerian or Norfo!kian stage, 1st Irilrrglacinl of authors.
"J'ai dernierement, avec M. Marcellin Boule, travaille a See geologic note above on the Forest Bed or Cromerian (p. 970). Lower
degager un enorme Elephas meridionalis qui a etc decouvert dans Pleistoeene.

les sables voleaniques de Sencze, pres de Brioude, par un savant Specific Chahacteus (compare Depehet and Mavet, 1923,
archeologue, M. Le Blanc. Ses dents contrastent singuliorement p. 153).— M' with -I-IOK (i.e., 13?) ridge-plates; very broad,
avec celles de VElephas meridionalis de Durfort par leurs eollines diameters 180 mm. X 80 mm.; laminar frequency 6-5 in 10 cm.,
basses, tres grosses, a email epais. Elles annoncent un animal greatly exceeding the typical E. meridionalis (5); enamel of
encore i)lus gigantesque." moderate thickness, much plicated, with salient loxodont sinus on
The dimensions given by Oaudry (]). 19) are as follows: posterior border.
According to Deperet and Mayet this subspecies is regarded
"Hauteur du squelette, a la tete 4™15
as a final phase of the Lower Pleistocene Forest Bed age of Eng-
du squelette, au garrot 3 83
land, a horizon which Osborn (1922.570) regards as corresponding
Longueur du squelette avec les de-
with the Isl Interglacial period, the period at which the true
fenses, la queue n'etant
Archidiskodon meridionalis makes its last a]:)pearance in southern
pas allongee et etant
Europe.
placee dans sa po.sition
Giant grinders of the southern mammoth, Archidiskodon meri-
naturelle 6 80"
dionalis, occur in the Forest Bed fauna (Osborn, 1922.563, p. 440,
listof vertebrata, etc.). The 12-|- ridge-plated tooth (Savin Mus.
—
No. 197 Fig. 871 F of the present Memoir) is 109 mm. in breadth,
as compared with 80 mm. in Deperet and Mayet's type (see Fig.

EUphas anfiquus. — The first left upper true molar ; from the Pleistocene of
Grays, Eteex. J. The lower border of the figure is the inner border
of the specimen.

ReKEUIIED .\rCHIDISKODON MEIllDION.\LIS FHOM THE I,OWEIt PLEISTOCENE OI' Essex (I^ekt), .\ni) Tvi-e op Archidiskodon meiudio.nalis cuomkiiensis
OK NoHFOL K (Right)
Eleven plated grinder, Vl.M', of Archidiskodon meridionalis
Fig. 869. Fig. 870. Ty|ie l.M' of Elephas meridionalis cromerensis Dep6ret and
illustratedand erroneously referred by Lydckker to "Elephas anliquus." Mayet, 1923, about one-half natural size:
PI. ix, fig. 1, p. 220,

After I.ydekker, 1886.2, ii. 12.5, fig. 26 (Brit. Mus. M.2004). Three-fourths "Fig. Elephas meridionalis mutation cromerensis, du Forest-Bed,
1.

natural size. a Kessingland. M' gauche avec fossili.sation et patine caracteristifiues (Iron-
lA'dekkcr described (p. 126) this "The first left upper
tooth as follows: pan). (Voir, p. 1.52.) Ce trc^s beau document i)al(5ontologique qui ressemble —
true molar in a half-worn eoudition; probably from Grays. This specimen de fayon frappante a dent n° 33,334 du British Museum- fait partie de la
la

(woodcut, fig. 26) agrees very closely in general characters with the .Japanese collection du Dr. Pontier. Photographic obligcanuncnt c(iiunuini(|uee par
molar of E. nninadicus figured on page 168 |Fig. 1189 of present Memoir]; notre .savant confrere."
it is noticed by I.eith-Adams, op.[1877 1881], p. 22.
cit. ,Vo histori/." Com- (Osborn) Mrit. Mus, 33,331 from the I''i>rcst Bed of the Norfolk coiist (,se(>

pare figure 870 op()ositc, type of the Elephas meridionalis cromerensis of Leith Adams, 1S77 1881, PI. xxiv, fig. 2, p. 198) agrees closely with the above
Dcpdrct and Mayet. Deperet and Mayet type.
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 981

871 as compared with Fig. 870, Dcperet and Mayet's type). Also ASCENDING MUTATIONS OF ARCHIDISKODON MERIDIONALLS
very numerous A. meridionalis grinders in the British Museum are Revision by Deperet and Mayet (1923)
catalogued by Lydekker (1886.2, pp. 108-113), mostly entered as Deporet and Mayet give a detailed description (op. ril., pp.
"dredged Happisburgh" or from the Forest Bed of Cromer,
off 125-160) of all the principal known remains referable to this great
Norfolk. Two grinders (cf. Fig. 862) figured by Falconer agree species, showing that the archaic forms of skull, jaw, and molar
in width with the measurements of Depcret and Mayet's type (80
mm.). The superior grinders of A. planifrons from the Siwaliks MAMMONTEUS PRIMIGENIUS 'ASTENSIS
M*MMONTEUS PRIMIGENIUS?ASTENSIS
vary in width from 88 to 100 mm. the inferior grinders vary in
;

width from 78 to 109 mm. (see Table VII above). There is

therefore no appreciable change in the breadth of the grinding
*13 +
teeth between A. planifrons and these specimens of ^. meridionalis /ost./c/' Mai Forest Bed F.Tfsf Bed of Crome'', Norfolk

(see Osborn's remarks above, page 970, on the Forest Bed fauna). PARELEPHUS > TROCONTMERH SP? PAPELEPHAS ''
TffOGONTMERII SP?
Elephas meridionalis, mutation cromerensis Deperet and
?r
Mayet, 1923. "Monographie des Elephants Pliocenes d'Europe et
de I'Afrique du Nord," Deuxieme Partie of "Les Elephants
Pliocenes." Ann. Univ. de Lyon, Nou\'elle Serie, I. Sciences, —
Medecine. Fasc. 43, pp. 150, 152, 157. Type.— A third 5ai.n Mui No 1240 Forest Bed

Horizon and Locality. — Ip5.,ch Mas Forest Be. — Dri.<iaea

....,.« A. MERlDrON Alis CROMERENS-S
superior molar of the left side, l.M^ —
ARCMIDISKOOON MERIDIONALIS CROMERENSIS
f^Vi /%"/% ""^ \x3
Forest Bed at Kessingland, England, Lower Pleistocene. Type
Figure.— Op. cit., PL ix, fig. 1.

Type Description. — (Op. cit., p. 153): "Cette derniere piece

(pi. IX, fig. 1) est une M' tres usee en avant ou manquent sans doute
plusieurs lames. La couronne est large (80 mm.) pour une longueur
conservee de 180 millimetres. On n'observe plus que 10 lames plus
le talon posterieur. L'email est peu cpais et tres plisse sur toute
I'epaisseur des bandelettes; il existe de sinus loxodontes assez
saillants du cote posterieur. La frequence laminaire atteint 6, 5
pour 10 centimetres, chiffre tres superieur a celui de I'E. meri-
dionalis type (5) et meme a celui de I'animal de Saint-Prest (5, 5).
C 'est le caractere essentiel qui permettra toujours de reconnaitre
la mutation cromerensis de I'horizon du Forest-bed."
Rase of Red Crgq Fglkenha.

19 28 H. F. O.

Primitive Grinding Teeth of the Mammontin.e; Archidiskodon, Parelephas, M.uhmonteus

Fig. 871. Molar ridge-plate structure and laminar frequency in the Forest Bed (Lower Pleistocene) and Red Crag (Upper Pliocene) eleiihants, after pencil
sketches by the author in 1926 of specimens in the Ipswich and Savin (Cromer) Museums. All figures reproduced one-fourth natural size. Compare Deporet
and Mayet, 1923, Pis. ix and xi.
Pleistocene A, Cromer Forest Bed (Ipswich Mammonleus primigenius(l) astensis Dep6rct and Mayet, M" (?), 11 ridge-plates or laminie in 10 cm.; total
Mas.) of 15 -f- ridge-plates.
B, Cromer Forest Bed Mammonleus primigeniusC?) asicyisisC?) Depcret and Mayet, I.Mj, 12 ridgc-platcs or lamina- in 10 cm.; total
of 13 -|- ridgc-plates.
C, Cromer Forest Bed (Ijiswich Parelephas (?) Irogonlherii Pohlig, sp.(?),r.M3, 8 ridge-plates or lamina? in 10 cm.; total of l.j-l- ridge-plates.
Mus., dredged)
D, Cromer Forest Bed (Savin Parelephas (?) trogontherii Pohlig. sp.(?),' f)% ridge-plates or lamina? in 10 cm.; total of 21+ ridgc-plates.
Mus. 1240) The determination of Parelephas (?) (C, D), corresponding to P. Irogontherii, is somewhat doubtful.
In the Savin Museum also occurs the typical llesperoloxodon aniiquus ausonius (?) with 12 ridge-plates.
E, Cromer Forest Bed Archidiskodon meridionalis cromerensis Deporet and Mayet, r.M', 6 ridge-plates or lamina? in 10 cm.; total
(Trimingham) of 12-|- ridge-plates.
F, Cromer Forest Bed (Savin Archidiskodon meridionalis cromerensis Dep6ret and Mayet, r.M', only 5 ridge-plates or lamina; in 10 cm.;
Mus. 197) total of 12+ ridge-plates.

Pliocene G, Red Crag (Falkenham) Archidiskodon planifrons (?) Falconer and Cautley, or E. [Hesperoloxodon] aniiquus, M3, 5/2 ridge-platcs or
laminae in 10 cm.; total of +11 ridgc-plates.
H, Red Crag (Ipswich Mus.) Archidiskodon planifrons (?) Falconer and Cautley, or E. [Hesperoloxodon] aniiquus, 0/2 ridge-platcs or lamina;
in 10 cm.; total of 10+ ridge-plates.

Observe that .Mammonleus has very compact ridgc-plates and fine enamel, that in the ParelcphasC?) ridgc-platcs the enamel is somewhat thicker, that
Archidiskodon has extremely broad and widely separated ridge-platcs of coarse enamel. Compare Depcret and Mayet (1923, "Les Elephants Pliocenes," Deux-
ieme Partie, pp. 98-160) for molar diagrams of A. planifrons and A. meridionalis, type ancien, which correspond very closely with the present diagram (E, F, G, H).

'Teeth similar to those of Parelephas Irogonlherii were selected by Pohlig, 1891 (1892), pp. 303, 304, as the cotypos of his Elephas aniiquus Nestii, as shown
in figure94 1 below (Chap. XVII) referred by the present author to ParelephasC!) Irogonlherii neslii.
;
982 OSBORN: THE PROBOSCIDEA

teeth approach A. plaiu'froiis, while the lii{j;hly progressive forms "IV. Elephas meridionulis, mutation ilu Suinl-I'rcslitn (ttndu

{op. ril., p. 153) approach the Lower Pleistocene stage of Archidis- Pliocene). Couronne moins basse (H = 90 a 95 milli-
large et

kodon meridionalis cromerensis. metres) 1 ou 2 lames de plus aux

; Mm
frequence laminaire 5, 5;
;

—
History. The southern mammoth is represented by superb email plus mince a plis serres affectant toute I'cpaisseur de la
specimens in the museums of France, which have recently been bandelette sinus loxodontes faibles et irreguliers.
monographed by Deperct and Mayet (1923), reciting the studies
"V. Elephas meridionalis, mutationcTOfnerensis {iormo (|uater-
of Nesti (1808, 1825), of Falconer (1868), of Weithofer (1890), of
naire du Forest-bed). Couronne moins large et plus haute (H = 90
(
'uvier, of de Blainville, of Owen, and of Gervais. The southern
a 95 millimetres); 12 a 15 lames aux M'', de 12 a 16 aux M3
mammoth (Elephas meridionalis) was long confused with the
sur les molaires entieres. Frequence laminaire 6 k 6, 5; email
northern mammoth {Elephan primigenius). Deperet and Mayet
beaucoup plus mince, a plis nombreux et serres affectant toute
remark {op. cil., p. 125): "("est seulement apres les voyages de
I'epaisseur de la bandelette; sinus loxodontes peu accuses et tres
I'alconcr en Italic et la publication des observations de ce savant en
irreguliers."
1868 que les caracteres \'K. meridionalis furent enfin reconnus et
I'espece definitivement admise par les paleontologistes." "Extinction du rameau."
DisTKiBiTiON. — As to geographic distribution, De]ieret and
Specific Characters of Archidiskodon meridionalis
Mayet conclude (op. rit., )). 144): "Ij' Elephas meridionalis, dont
(Deperet and Mayet, 1923, p. 156). "1° Crane. Le crane
— —
le centre de dispersion principal est repandu sur pres-
I'ltalie, s'est
connu surtout par les belles pieces de Florence, est caracterise
ijue toute la surface de TEuropc temperee: Russie du Sud, bassin
par: une boite cerebrale arrondie tres large en haut et a bords
tlu Danube, Allemagne du Sud, France, Angleterre, peninsule
presque paralleles; un vertex peu elevc arrondi, mais non j)ro-
Iberique, et enfin dans I'Afrique du Nord. Les gisements sont tres
longe en dome; une region fronto-parietale excavee chez I'adulte
iiombreux; aussi notre etude descriptive se limitera-t-elle aux
une ouverture nasale reculee tres en arriere, etroite et tres etendue
l)ipces les plus import antes, notamment les Mm, et surtout a celles
en travers, surmontee d'une epine nasale saillante; des arcades
dont nous avons pu obtenir des photographies directes, grace a
zygomatiques dejetees vers le bas; des alveoles des defenses rap-
I'obligeance de divers de nos confreres, que nous citerons au cours
proches et subparalleles."
(le cet expose. Comme en Italic, nous pourrons y reconnaitre trois
mutations successives de I'espece: A. Forme archaique. B. Forme "2° Mandibule. — La mandibule offre des particularites interes-

type. Forme evoliiee oit rwetde."

('. santes: chez le jeune et la femelle, la symjihyse .se jirolonge par

—
Mutations. The chronological mutations close as follows un bee presque horizontal qui continue le bord inferieur de I'os.
{op. cil., p. 150): Chez le male adulte, au contraire, la mandibule est depourvue de
bee et presente une terminaison mousse et obtuse, avec i)arfois
"2° Mutation de I'etage Cromerien (Sicilien) qui represente le
une toute petite pointe mediane insignifiante (scjuelette de Dur-
debut du (^uaternaire et que nous designerons sous le nom de
fort). On a deja dit que VE. planifrons possede dans les deux
niutalion cromerensis."
sexes un bee mandibulaire beaucoup plus fort et dejeto presque
"1° Mutation de I'etage Saint-Prestien, cjue nous rapportons a verticalement vers le bas."
du Pliocene."
I'extreme fin
"3° Defenses. — Les defenses sont, chez le jeune et la femelle,

di\ergentes des la base et iissez fortement spiralees. Chez le

Ascending Skhies of STUATHiiiAPHir Mitations of male adulte, elles sont presque paralleles a leur base, .«e dirigent en
A. .MICRiniONALIS, BEGINNING OF EXTINCTION OF THE PhVLIM bas et en dehors, decrivant ensuite une legere spirale dont les
IN Entoi'E (Deperet and Mayet, 1923, v. 157) pointes reviennent un peu en dedans. (Jhez VE. planifron.'<, les
defenses sont encore [ilus paralleles et out une courbure concaNC
"I. Elephas planifrons. Couronne tres basse (hauteur d'uiie
moins prononcee."
lame mediane moyennement usee, 50 a 60 millimetres) et large;
lames transverses peu nombreuses (10 chez M' et 10 11 chcz Ms); "4° Molaires. — Les molaires de \'E. ineridionulis sont de
grand ecartement des lames (frequence laminaire 3, 5 a 4 lames par iiieme type general que celles de VE. planifrons: couronne large et
10 centimetres de longueur de la couronne); email tres cpais, a basse, lames tran.sverses i)e.u nombreuses, email cpais et generale-
iarges ondulations limitees a la parol externe des bandelettes; sinus ment peu plissc; sinus loxodontes assez accu.ses mais inconstants."
loxodo.ntcs tres saillants et assez reguliers, surtout en avant."
THE ARCHIDISKODON MERIDIONALIS I'HYMM MKIR.VrKS TO
"II. Elephas meridionalis, mutatioji archaique. Couronne AMERICA
large et bas.se (H =60 a 75 millimetres); frequence laminaire 4, 5;
Osborn (1924) welcomes this splendid resume by his friends
email dpais a Iarges plis i)ou profonds; .sinus loxodontes plus
Deperet and Mayet of specific characters and mutations of A.
irr^guliers."
planifrons and A. meridionalis, but does not agree that there is
"in. Elephas meridionalis, mutation ttjpe. Couronne large et adequate evidence that this phylum became extinct, for it apjiears
iin |)eu inoins ba.sse (H=75 a 90 millimetres); I\P a\ec 11 a 13 quite certain that the Archidiskodon meridionalis tyi)e migrated
lames, M.I avec 11 a 14; freciuence laminaire 5. lOmail moiiis cpais from Asia into North America and became ancestral to Archidis-
et a plis plus .serres qui int<>ressent presque toute l'e])aisseur (l(!s kodon impcralor of Nebraska. Nor does Osborn believe that the
bandelettes; sinus loxodontes peu accuses et irrcguliers." meridionalis of Italy and France is in any way related to the
 THE MAMMONTIN.E: ARCHIDISKODON AND METARCHIDISKODON 983

Elephas hysudricus of the Upper Siwaliks treated in Chapter XX. it may indicate that members of the generic piiyluin Archidinkodon
The cranial types in these two animals are fundamentally distinct. originated in Africa and migrated northward into Europe and east-
If Africa should prove to be the home of Archidiskodon, as ward into southern Asia rather than having followed the generally
appears probable through the discovery of two new primiti\e accepted reverse line of migration from southern Asia westward
species in southern Africa, namely, A. subplanifrons and A. broomi, into Europe and Africa.

[It willbe observed that Professor Osborn wrote this portion of the Memoir in 1924. From that time onward he
constantly sought new evidence to test his hypothesis of an African center of dispersal of the Proboscidea (see Fig. 815).
A decade of study (1924-1934) only confirmed him in his belief and in 1934 he wrote an article in American Museum
Novitates (Osborn, 1934.925) entitled "Primiti\^e Archidiskodon and Palaeoloxodon of South Africa," on the basis of
which the following section has been revised but in no sense have the views of the author been changed otherwise than
—
appear in his own writings. Editor.]

I'lG. 872. Fo.SSIL PUOBOSCIDEA RoUTE FROM SoUTHEnN EQUATORIAL AFRICA

Compare Andrews (1911), Dietrich (1916), Haughton (1922), Hopwood
(1926),Dart (1927), and Broom (1928). [Since 1928 other species have been
described (see pp. 986-993 below).— Editor.]

(2) Archidiskodon subplanifrons, Sydney-on-Vaal, Vaal River diggings,

near Kimberley.

(7) M elarchidiskmlon griqua, Vaal River diggings, Griqnaland West.

(10) Archidiskodon broomi, The Bend, Vaal River, near Kimberley.

(11) Palieoloxodon transvaalensis, Bloemhof, southwestern Transvaal.

(12) Palsoloxodon sheppardi, Bloemhof, southwestern Transvaal.

(3) Loxodonta zulti, Zululand.

(9) Mastodon s|j. (?), northwest of Lake Nyassa, near Uraha Hill.

(4) Deinotherium hobleyi, Karungu, near Victoria Nyanza, Tanganyika

Territory.

(6) Palseoloxodon recki, near Oldoway, Serrengctisteppe, northern

Tanganyika Territory.

(8) Metarchidiskodon grigtia, Kaiso Bone-beds, near Albert Nyanza.

(.5) Mastodon sp. (?), near Khartum, Sudan.

(1) Deinotherium, ref., Elephas ref., Lake Rudolf.

2. ARCHIDISKODONTS AND METARCHIDISKODONTS OF SOUTH AFRICA

(Osborn, 1934.925, pp. 1 — 10): "Every year brings fresh proof that Africa was the center of the origin and
adaptive radiation of the Proboscidea. Since 1907 numerous more or less primitive superior and inferior grinding

teeth have been discovered from the Vaal River terraces and other localities of the Transvaal, South Africa. The
geologic level and localities are chiefly on the (1) higher and most ancient terrace (200-300 feet) of the Vaal River;
(2) middle terrace (60-80 feet) of the Vaal River; (3) lowest and most recent terrace (40 feet) of the Vaal River.
 . : — . : :

984 OSBORN: THE PROBOSCIDEA

Theoretically the lowest terrace may be as old as the lower levels of the middle terrace. Flint iun)lements occur
in the middle and lower terraces only (Dart, 1929). In Osborn's opinion the Archidiskodon subplanijrons, and
Archidiskodon pro planifruns, . . types found in the middle terrace were washed in from an older Pliocene horizon."

"Certain of the Transvaal grinding teeth surely belong to very primitive stages of Archidiskodon; others
probably belong to primitive stages of Palaeoloxodon and were originally referred to Loxodo7ita, to Archidiskodon
and to PUgrimiay^ Only by careful comparison and analysis is it possible to separate the species belonging to
these several genera from each other."

"Up to the present time [1934-] the nineteen species described by W. B. 8cott (1907), Raymond A. Dart
(1927, 1929), S. H. Haughton (1922, 1932) and H. F. Osborn (1928) are provisionally referred as follows:

Locality Original Reference Present Generic Reference

Zululand Loxodon Zulu Scott, 1907 = Loxodorda
Vaal River
?]\Iiddle terrace, Loxodonta griqua Haughton, 1922 = Mctarchidiskodon, n.g.
Lowest terrace, Vaal River Archidiskodon transvaalensis Dart, 1927 = Palaeoloxodon
Lowest terrace, Vaal River Archidiskodon sheppardi Dart, 1927 = Palaeoloxodon
?Middle terrace (lower), Vaal Ri\er Archidiskodon subplanifrons Osborn, 1928 = Archidiskodon
Lowest terrace, Vaal River Archidiskodon broomi Osborn, 1928 = A rchidiskodon
Middle terrace (lower), Sydney-on-Vaal Archidiskodon vanalpheni Dart, 1929 = Archidiskodon
Middle terrace (lower), Sydney-on-Vaal Archidiskodon loxodontoidcs Dart, 1929 = A rch idiskodo n
Middle terrace (lower), Sydney-on-Vaal Archidiskodon milletti Dart, 1929 = Archidiskodon
?Middle terrace, Vaal River Archidiskodon andrewsi Dart, 1929 = f Palaeoloxodon
Lowest terrace, Vaal River Archidiskodon hanekomi Dart, 1929 = Palaeoloxodon
Middle terrace, Vaal River Archidiskodon yorki Dart, 1929 = A rchidiskodon
Lowest terrace, Vaal River Pilgrimia yorki Dart, 1929 = Palaeoloxodon
Lowest terrace, Vaal River Pilgrivtia wilmani Dart, 1929 = Palaeoloxodon
Pniel Estate, ? River Pilgrimia kuhni Dart, 1929 = Palaeoloxodon
?Recent, Limpopo River Loxodonta prima Dart, 1929 = Loxodonta
?Recent, Steelpoort River Loxodonta nfricana, var. obliqua Dart, 1932 = Loxodonta
[1929]
?Middle Terrace, Vaal River Pilgrimia archidiskodontoides Haughton,
1932 = Palaeoloxodon
Higher terrace, Vaal River Pilgrimia stibantiqua Haughton, 1932 = Loxodonta"

"Sununing up these species, the ascending geologic level records (Dart, Haughton) are as follows' [Table
VIII of the present Memoir]

Table VIII. "Faunal Distribution on the River Terraces of the Transvaal"

"Middle Terrace Middle terrace, 60-80 feet. Lowest terrace, 40 feet. Recent —4 feet.
Sy d ney-o n- Vaal With flint implements. Pleistocene
?Lower Pleistocene ?Middle Pleistocene. Bloemhof

Upper levels: Upper levels: Levels unknown

?A. subplanifrons* ^4. yorki Bubalis baini '/P. hinickoini
?A. proplanifrons** A. broomi Equus capensis fL. subantiqua
Older than the Upper Pliocene ^. AL griqua fP. archidiskodontoides
planifrons of the SiwaUks, India
Lower levels: Lowest levels: Recent
*actuallv recorded from a depth of /-". andreivsi P. transvaalensis L. prima
50-60 feet, middle terrace (60-80 ft.) A. vanalpheni P. sheppardi L. africana var. obliqua"
A. 7nilletti P. yorki
**at a depth of .56 feet, middle A . loxodontoides P. wilmani
terrace, Vaal River Bunolophodon ?gen. ?sp. P. kuhni
''' Pilgrimia Osborn (December 20, 1924) is antedated by Palaeoloxodon Matxiimoto (September 20, 1924).
-[To tlie.se might be added Archidiskodon proplanifrons described in this article (Osborn, 1934.92.'), p. 10), see page 986, figure 873, below.— Editor.]
 : : .

THE MAMMONTINiE: ARCHIDISKODON AND METARCHIDISKODON 985

"By their outstanding characters these species divide into four groups as follows" [Table IX of the present

Memoir]

Table IX. "Provisional Grouping of Species Referred to Four Transvaal Genera"

"A. subplaiiifrons group M. griqua group P. transvaalensi s group Loxodonta prima group

Crowns very broad, 101 to 114 Crowns relatively narrow, 86- Crowns M^ relatively nar-
of Crowns relatively narrow, 74
mm. Enamel very thick. 94 mm. Enamel thick. row, 70 mm. {P. wilmani) to mm. (L. prima) to 92 mm.
Transverse conelets 4-6 (A. Transverse crests 6-8. Ce- 110 mm. {P. transvaalensis) (L. subaniiqua). Enamel
pruplanifronn) to 22-24 {A. ment areas narrower, not Indices = 41 to 51. Enamel relatively thin, coarsely
broonii). Cement enveloping enveloping crown. Valleys relatively thin ; conelets fine- crimped conelets numerous.
;

crown. V-shaped cemented V-shaped {M. griqua). Post- ly crimped, i.e. numerous. Cement thin in middle,
valleys at smnmits broader sinus fold very prominent. Cement areas progressively thick at edge. Ridge jjlates

than dentinal areas. Lo.xo- Total enamel length un- narrower t han dentinal areas. jier 100 mm. = 4 (L. afrirana

dont sinus foldings double, known. The mass of cement Cemented valleys greatly re- obliqua) to B'i (L. subanti-
less prominent, irregular. exceeds the mass of dentine. duced. Ridge plates narrow qua). Broad typical loxo-
The mass of cement exceeds and increasingly lofty, 128 dont sinus expansion, double
the mass of dentine. mm. {P. wilmani), 259 mm. sinus foldings in contact.
{P. hanekomi). Ridge plates Total ridge 9 (L. plates

Cf paratype of A meridionalis
. . Cf. A. planifrons r)nnnn)is
per 100 mm. 4-6. Sinus fold- prima) to 12-13 (L. zulu).
ings extremely reduced or
Nesti of Val d'Arno, also Stefanescu.
progressi\ely wanting. \a\-
Brit. Mus. M 1264 M 1 , 1 2642.
leys V-shaped {P. andrewsi).

In this group are the following In this group may be the In this group are the following: In this group are the following;
species following species:
P. = Pilg.]
[ kuhni, P. L. zulu, L. [
= Pilg.] subanii-
A. proplanifrons, A. sub- .1/. griqua, P. andrewsi, A. [
= Pilg.] yorki, P. [
= Pilg.] qua, L. africana obliqua, L.
planifrons, A. milletli, A. lo.rodontoides. wilmani, P. archidiskodont- prima.
yorki, A. vanalpheni, A. oides, P. sheppardi, P. trans-
broomi. The generic relationships of vaalensis and P. hanekomi.
thisgroup are doubtful the These occur only on the more
;
Also possibly ?P. andrewsi.
The ridge plate height increases narrow crowns separate the recent levels and are clearly
These seven or eight types
from 55 in .4. proplanifrons types of M. griqua and of P. related to the existing Afri-
are much more uniform in
to 62e. in A. subplanifrons, andrewsi from the broad can elephant, distinguished
character than members of
to 118 in A. milletli, to 129 crowns of the typical Archi- by the above characters."
the M. griqua group, P.
in A. vanalpheni and 110+ diskodon. The U-shaped sheppardi and P. transvaal-
in A. broomi. Meanwhile valley of M. griqua of Fig. 3 ensis formerly being referred
the number of ridge plates is quite distinct from the by Dart to Archidisko-
in 100 mm.
remains con- V-shaped valley of P. an- don. P. kuhni, P. yorki
stant, namely, 3 in ^. pro- drewsi; similar teeth hsLve and P. wilmani were referred
planifrons, ^Yi in A. plani- been discovered in Europe. by Dart to Pilgrimia; the
frons of India and 3 in ^4. It is probable that these prevailing characters relate
broomi. teeth represent a genus dis- them more closely to Palaeo-
tinct eitherfrom Archidisko- loxodon.
don or Palaeoloxodon, name-
ly Metarchidiskodon.

"The above phylogenetic arrangement is provisional. Only by the longitudinal sectioning method of Falconer
is it possible to ascertain the true structural relationships of these Proboscidean molars. This is illustrated in the
wide difference between sections of M. griqua (Fig. 3 [
= Fig. 882]) and P. andrewsi (Fig. 5 [=Fig. 1139]), also in the
wide difference between the sections of A. subplanifrons (Fig. 1 [
= Fig. 875] and A. proplanifrons (Fig. 2 [
= Fig.
 —

986 OSBORN: THE PROBOSCIDEA

873]). A very interesting comparison is that of the 'planijrom series of Africa and the SiwaHks of India analysed
as follows" [Table X of the pre.sent Menioii'|:

Table X. "Comparative Measurements of the Primitive Species of Ahchidiskooon FiiOM the Transvaal
AND FROM the UpPER SiWALIKS OF InDIa"

o
o

A. prnplanifrons KM''

A. subplanifrofis RM3

A. plamfrons* mu
 — —

THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 987

Archidiskodon subplanifrons Osboni, 192S folhjws (Osborn, 1928.749, p. 672) "The first typo (re|)resented in
:

Figuies 81.j, 874, 875, 123.'-), 1239, PI. xxi Fig. 1 [

= Fig.
874 of the present Memoir]) I name Archidinkodoit
Upper (?) Pliocene. Sydney-on-Vaal, Vaal River diggings, South Afriea, fiuhplanifrons; it is a low-crowned, broad-plated, heavily cemented
on the banks of the riv(>r, ata depth of from M
to 60 feet [Middle Plioeene tooth, apparently a third inferior molar of the right side (McGregor
see Fig. 1239]. The
Mus. 3920). specific name subplanifrons refers to the fact
—
History. The tyjie tooth was riiscovered by Mr. W. Millett that the crown height — from 2 to 2)1 inches — is about equal to that
in the Vaal River diggings at Sydney-on-Vaal, South Africa, on the of the low-crowned types of Elephas planifrons Falconer of the
banks of the river, at a depth of from 50 to 60 feet the geologic ; Siwalik Hills, India; in some Upper Siwalik speci-
of Falconer's
age is probably r])i)er Pliocene. Through the courtesy of Miss mens the crown rises from SJi to
4% inches. The present specimen
Wilnian, Curator of the McCiregor Museum at Kiniberley, and of accordingly is believed to be of Upper Pliocene age."
Archidiskodon subplanifrons Osborn, 1928. "Mammoths and
Ce/iti nitti
,

Man in the Transvaal." Nature, Vol. GXXI, No. 3052, April 28,
1928, pp. 672, 673. Supplementary description: "Primitive
Archidiskodon and Palaeoloxodon South Africa" (Osborn,
of
1934.925, p. 10, fig. 1 = Fig. 875 of the present Memoir). Horizon
.\Ni) Locality. —
Upper(?) Pliocene. [Middle Pliocene See Fig. —
—
1239 below. F^ditor.] Sydney-on-Vaal, Vaal River diggings, on
the banks of the river, at a depth of from 50 to 60 feet i?Middle
Terrace), South Africa. —
Type Figure. Op. cil., p. 672, fig. 1
—
Type. Posterior half of a third right inferior molar, r.Ms
(McGregor Mus. 3920, cast Amer. Mus. 21924), exhibiting four
complete posterior ridge-plates and half of another ridge-plate,
deeply surrounded with cement on crown and sides, also buried in
cement is a rudimentary plate; extremely low crowned with broad,

/^iQreqor Mui. J920 Ximier.y 3. Africa.

r. TO. 5

/^ nat. size- McGregor Mas. 45i>-4-

Type. Arc/iLdiskodon propLanlfrons. Osborn. 1954-

Fig. 873.Type, right third superior molar, r.M', of Archidiskodon pro-
McGregor Museum 433-1, Kimberley, South Africa;
planifrons, after original.
cast Amer. Mus. 26969. One-half natvu-al size. Originally referred by
Haughton (1932.1, p. 2) to Archidiskodon subplanifrons.
B, Crown view showing 5% ridge-plates, very broad cement areas c, c, c;

sixth plate represented by two conelets. Bl, section of the same showing
cement-filled ridge-plates more widely open than in A. subplanifrons.

our correspondent Dr. R. Broom, the molar was forwarded January

20, 1927, to the American Museum for comparison and type de-
scription; in return for this courtesy, a number of facsimile casts Original Type Figure of Archidiskodon subpl.^nifrons
were made, so that each specimen bears the inscription: Archidis- Fig. 874. Ty|)e of Archidiskodon subplanifrons Osborn, 1928, from the
kodon mbplanifron.% McGregor Mus. 3920, type, cast A. M. 21924. Upper (?) Pliocene [Middle Plioeene —
see Fig. 1239], Sydney-on-Vaal, South

This species was finally described and figured by Osborn in Africa, one-half natural size. Third inferior molar of the right side, r.Ms
(McGregor Mus. 3920, Kiniberley, South Africa; cast Amer. Mus. 21924). After
"Nature," April 28, 1928, pp. 672, 673, in comparison with A.
original type specimen kindly loaned to the American Museum for figuring
hroomi. and description (cf. Osborn, 1928.749, fig. 1, p. 672). Compare figures 840,
Type Description. —The original description by Osborn is as 876, also an r.Mj (Amer. Mus. 19965) of ^. planifrons from the Siwalik Hills.
988 OSBORN: THE PROBOSCIDEA

heavy enamel, ridse-])lates set wide apart, two of the anterior external aspect the ridge-plates are less elevated.
Length of type
ridge-phxtes expanding into a 'loxodont sinus' (4th and 5th); molar 153 mm., breadth lUl mm., index 66; enamel length, re-
strongly resembUng the Archidiskodon planifrons type (Figs. 828, stored, 650 mm., enamel area 2,600 sq. mm.; average enamel
829), hence the specific name Airhidiskodon ftuhplariifwns. thickness 4 mm. Ridge-plates (6) postconvex, jireconcave, 5-6
Specific Characters. —
The concavely worn superior surface subo\al to round conelets with double central folds in the ridge-
(Fig. 874, lower) enables us to determine this as an inferior tooth plate; ridge-plates per 100 mm. =4.
of the right side, apparently an r.Ms, in view of the posterior con- Specific Compariso.n with Archidiskodon planifrons in
vexity (Fig. 874, upper) which forbids its reference to an r.M2. —
THE Brown Siwalik Collection. The South African type of
The crown is excessively broad and short, with convex cement- Archidiskodon subplanifrons (Figs. 874, 875) is very similar to
covered sides and ridge-plates which gradually increase in height
posteriorly, iis follows (reading backwards) 4th, imperfect, with
:

central loxodont sinus; 5th, tr. 92 mm., height 53e mm., with
anterior and posterior loxodont sinus; 6th, with cement, tr. 97
mm., height 53e mm., loxodont sinus less prominent; 7th, tr. 101
mm., height 63 mm., loxodont sinus less prominent, five to .six
conelets; 8th, with cement, tr. 95 mm., height 61 mm., summit
of ridge-plate narrow with six conelets; after this a rudimentary
plate buried in cement. The thick, non-crenulated enamel of the
ridge-plates strongly reminds the observer of the ridge-plates oi A.
planifrons (Fig. 829). Without the aid of a section it is difficult to
determine whether the ridge-plates are lofty as in Falconer's type of
A. planifrons (Falconer and Cautley, "Fauna Antiqua Sivalensis,"
1846 [1845, PI. II, fig. 5al— Figs. 825, 828 of present Memoir); in

//// // // %'); JvV 5,.-; 6. 7. 5

1996.5

Archidiskodon
pLaniFrons. Ref.
Falconer. 1845
Enamel tengtti '810 mm. est.
Ridge plate httght = 55m

A I 1/2 not size ' McQregor Mas. 3920

Type. Arctitdiskodon subplanifrons. Osborn.i928
Fig. 876. Referred third right inferior molar, r.M.-i, of Archidiskodon
Fig. 875. Now figure of type riglit thinl inferior molsir, r.M.i, of Archidis- idanifnms Falconer, 184.'), from three miles north of Siswan, India. Sj;, [91
kodon xubplanifronfi O.slmrn, 1928, from tli<! middle terrace, Sydiiey-on-Vaal, ridge-plat.es. Iiitenial aspect (below), exhihitiiin the gently folded section of
South Africa. McOrcKor Museum 3920, Kiml)erley, South Africa; cast the entire molar with relatively low enamel ridge-plat<'s and estimated total
Amer. Mus, 21924. One-half natural size. enamel length of 810 mm. One-half natural size.
—
A, crown view, c cement, rf -dentine. Line of mid.section. At, the Compare this molar of Archidiskodon planifrons from India with type
samp in midsection, exiiibiting .six ridge-plates. molar of .1. subplanifrons (Fig. 874) from South Africa.
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 989

a referred specimen of A. planifrons (Amer. Mus. 19965) from the —

Characters. (1) The ridge-plates present an
Specific
Upper Pliocene Pinjor horizon, Upper SiwaUks, India. For com- anterior concavity and a posterior convexity as in ^. imperalor
parison both teeth are carefully figured to a uniform scale (Figs. (Fig. 889A) (2) the
;
ridge-plates converge towards each other on
875, 876). Observe in these teeth: (1) The similar height of the the external side and are somewhat farther apart on the internal
ridge-plates, e.g., A. planifrons 55e mm., A. subplanifrons 53e side, consequently we are inclined to consider this molar as an
mm.; (2) A. planifrons with 8% [9] ridge-plates, A. subplanifrons r.]\l2 by comparison with Archidiskodon imperalor (Amer. Mus.

with 8)26 ridge-plates; (3) similar constitution of the enamel and 14558 — Fig. 889 A), but the exact placing of this tooth is uncertain;
cement; (4) similar obhque, outwardly facing, slightly concave (3) the fact that the fifth ridge-plate preserved is broader (tr.
ridge-plates; (5) the median loops, 'loxodont sinus,' are somewhat 107 mm.) than the sixth and seventh ridge-plates preserved
more prominent in A. subplanifrons than in ^4. planifrons. (tr. 105 mm.) favors the determination of this tooth as a ten or

eleven plated r.M2 rather than a twelve to fourteen plated r.Ma;

Archidiskodon broomi Osborn, 1928 (4) the maximum width of the fifth ridge-plate or pentalophid is
Figure 877 107 mm., practically the same as that of the corresponding penta-
Lower or Middle Pleistocene. Discovered in 1920 at The Bend, on the
Va.al River, near Kimberley, South Africa.

History. —Aside
from the uncertainty as to the exact loca-
most interesting type molar of Archidiskodon
tion in the jaw of this
broomi, there is no doubt as to its relationship to the gigantic,
broad-plated genus Archidiskodon and as to its affinity to A. meri-
dionalis, the southern mammoth of Europe, as well as to its
American relative A. imperalor, the imperial mammoth of the
southern United States and of Mexico. Consequently this is

a most important and interesting case of geographic distribution

and of independent parallel evolution of species of Archidiskodon in
South Africa and southern Europe and in the southern United
States.
Archidiskodon broomi Osborn, 1928. "Mammoths and Man in
the Transvaal." Nature, Vol. CXXI, No. 3052, April 28, 1928, pp. ARCHI0I5K0D0N BROOMI
672, 673. Horizon and Locality. — Lower or Middle Mus. Ktmk 3682 Tyfle //. M cast 2/90/
Pleistocene. The Bend, on the Vaal River, near Kimberley, South
Africa. —
Type Figure. Op. cil., fig. 2, p. 672.
Original Type Description of Archidiskodon broomi
(Osborn, 1928.749, p. 672).— "In the second type (Fig. 2 = Fig. [

877 of the present Memoir]), the anterior half of a third superior

[inferior] molar, probably of Pleistocene age, we observe a far more

progressive stage, with lofty ridge-plates, the sixth attaining

a height of 5 inches, equal to that of certain specimens of Archi-
diskodon meridionalis in which the ridge-plates equal or exceed 5
inches. This relatively high-crowned type (McGregor Mus., 3682)
I name Archidiskodon brooyni, in honour of Dr. Robert Broom, who,
after the specimen was named in MS. and figured by myself, re-
quested that one of these molars should be named after Mr. W.
Millett, who discovered the type of A. subplanifrons at a depth of
from 50 to 60 feet in the Vaal River diggings near Sydney-on-Vaal."
2 nat. sue
—
Type. Seven-plated anterior portion of an imperfect third
inferior molar of the right side, bearing the inscription: "3682
Mus. Kimb. The Bend. H. Else." Cast Amer. Mus. 21907. Re-
Osbgrn's Original Type Figure of Archidlskodon broomi
produced from original specimen (Fig. 877 of the present Memoir).
By comparison with the superior and inferior molars (Amer. Mus. 877.
Fig. Type of Archidiskodon broomi Osborn, 1928. Original in
the McGregor Mu.seuni, Kimberley, South .Africa (No. 3682), castAmer.
14476, 14558) referred to Archidiskodon imperalor (as illustrated in
Mus. 21907. Third inferior molar of the right side, r.Ms. From The Bend,
Fig. 889 B, A, of the present Memoir), this type grinding tooth of Vaal River, near Kimberley, South Africa. After original specimen kindly
Archidiskodon broomi agrees approximately in size with A. im- loaned to the American Museum for figuring and description. Compare
peralor in the maximum width of the fifth ridge-plate, namely, 107 Osborn, 1928.749, p. 672, fig. 2.
Seven ridge-plates partly preserved, broadest portion of the crown at
mm. The ridge-plates of A broomi also agree exactly in width (107
.

fifth ridge-plate, indicating that three or four posterior ridge-]jlates are

mm.) with those of M^ in Brit. Mus. 7436 (see Falconer and missing and that this may represent a ten or eleven plated second molar, r.M2,
Cautley, "Fauna Antiqua Sivalensis," 1846 [1847, PL xiv.b, fig.
according to the formula; of Leith Adams (1877-1881) and of Weithofer (1890),
13a]), a true Elephas [
= Hesperoloxodon] antiquus molar. in wliich Archidiskodon meridionalis is clearly distinguished from .1 . planifrons.
990 OSBORN: THE PROBOSCIDEA

lophid in A. iniperdlor(tr. 108 mm.) and the corresponding penta- tion of the plates is not less marked in their transverse length which
l(tpli A. men'dioiKtlis (AP); (5) in ,1. iinpenilor (Amer. ]\Ius.
in at the ba.se is in the region of 4 inches, hut at the grinding eilge is

—
14476 Fig. 889B) the convexity of the ridge-plates is anterior, the not more than 2}i mm.
These features lend to the
inches or 63
—
concavity is posterior therefore, by comparison, the A. broomi tooth as a whole a pyramidal appearance which is in sharp con-
type is an inferior molar; (6) against this interpretation, however, trast to the squat semicuboidal form and exuberant cementum

isthe fact that in profile (Fig. 877, lower) the wearing surface of the found in .4. subplanifrons. The enamel in thickness (3-4 mm.) is
ridge-plates is slightly convex (as in superior molars generally), comparable with that of A. subplanifrons but is somewhat more
whereas, according to rule, the profile of the inferior molars in folded than in that form, definite longitudinal depressions running
Archidiskodon .should be slightly concave (.\mer. Mus. 10598 and e\'en along the exposed lateral and medial sides of the plates. It

14558— Fig. 892 B, A). is these enamel foldings which more jiarticularly give the tooth its

The detailed specific and generic characters and measurements rugged character."
are as follows: and second ridge-plates narrow and com-
(1) First "Owing to the early age of the tooth, not more than three
pletely worn into confluence, third ridge-plate (tr. 86e mm.), plates (excluding the most anterior talon) being in wear, a clear
fourth ridge-plate (tr. 102 mm.), fifth ridge-plate (tr. 107 mm.),
.sixth ridge-plate (tr. 105 mm.), seventh ridge-plate (tr. 105 mm.),

height of seventh ridge-plate (110 mm.); (2) enamel borders of

each ridge-plate folded or ptychoid, slightly expanded in the fourth
ridge-plate on wear into a faint 'loxodont sinus'; (3) the ptychoid
enamel borders dLstingui.sh the sides of the ridge-plates (Fig. 877,
lower); (4) broad cement deposit between the ridge-plates, not,
however, covering the plates at the sides as in /I. imperaior.

Archidiskodon vanalpheni Dart, 1929

Figure 878
From Sydney-on-V'aal, South Africa. Middle terrace (lower levels).

?Lower Pleistocene.

Archidiskodon vanalpheni Dart, 1929. "Mammoths and

Other Fossil Elephants of the Vaal and Limpopo Watersheds."
So. Afr. Journ. Sci., XXVI, p. 704. Type.— Third superior
molar of the left side, l.Ml McCiregor Museum (Kimberley)
4086 (cast Amer. Mus. 22723). Horizon and Locality. —
"Middle terrace at a depth of 80 feet at Sydney-on-Vaal," South
Africa. ?Pliocene [?Lower Pleistocene {fide Osborn, 1934.925, pp.
2, 3)]. Type Figure.— Dart, op. cil., p. 704, figs. 8 and 9.
Type Description.— (Dart, 1929.1, pp. 704-706): "This
tooth (greatest length 234 mm., greatest width 112 mm., greatest
height 129 mm.) has 8 plates and is incomplete posteriorly, there

having been originally probably 9 plates present (see Figs. 8 and

9 (Fig. 878 of present Memoir])." Fig. 878. Type left third superior molar, l.M' (McGregor Mu.s. 4086,
"The tooth in A. vanalpheni presents a characteristically ea.st Amer. Mu.s. 22723), of Archidiskodon vanalpheni Dart, 1929, figs. 8 and 9,

rugged appearance, the massive plates being separated laterally p. 704, one-third natural size. From Sydney-oii-Vaal, South Africa.

l)y great valleculae approaching an inch in depth. These huge

furrows are continued across the grinding surface of the tooth to
a depth of % inch or more (save in the case of the space between the jiicture of the grinding surface in full wear is not a\ailable. In
3rd and 4th plates only) and give to the organ an appearance the fourth plate, 6 mammillated ])oints are in slight wear, in the
simulating that characteristic of the Stegodonts. Owing to the third there are 2 islets and 3 mammillated proce.sses — indicating
fragility of thespecimen, the fangs of the tooth are absent, as in all apparently 7 mammillated processes to each plate, a number
also are most of the mammillated processes of the posterior four which appears to be ratified in the posterior plates which ha\'e
plates. The first or anterior plate is talon-like in appearance, lost their processes by fracture —
while the second j)late shows four
being aborted and only half the height of the succeeding one, to islets. A tendency is demonstrated especially by the anterior
which it is firmly adherent, with the interposition between them of talon and the second and third plates towards their centre to
very little cementum." throw out a posterior buttress. There is no evidence whate\er of
"The massive plates are markedly cuneiform, as opposed to the an anterior buttress."
chevron-patterned plates of Stegodonts on the one hand, and the "The tooth is distinguished also by the great degree of sep-
pectinated arrangement exhibited by more atlvanced elephants on aration between the plates owing to the presence of great wedges
the other hand, their antero-posterior witlth tapering from 1% of interlamellar cementum desjiite their valleculisation. This
inches at their base to %-'/i inch at the grinding edge. The contrac- .separation, which in A. subplanifrons does not exceed %-% in., is
 —

THE MAMMONTIN.E: ARCHIDISKODON AND METARCHIDISKODON 991

%-Va in. in A. vanalpheni. There can be little doubt that we generally only 'i inch apart, so that in .4. tnilletli a length of 5)^
possess here another very ancient Archidiskodont species, which inches includes 5 plates while in A. vanalpheni it includes but 4
despite its greater number of ridge plates (A. siibplcinifrons shows plates."
only 5 plates but may have had more) illustrates in its cement "When in addition it is pointed out that, while there is here,
characters and the ridge plate form an exceedingly primitive type. too, an appreciable vallecuHsation of the interlamellar cementum
I have named it after Mr. van Alphen who was responsible for which is continued across the tooth between the posterior plates
securing it for scientific investigation." there is, nevertheless, in A. milletti a far greater relative bulk of
cementum and a far greater tendency towards the reduction of the
Archidiskodon milletti Dart, 1929 valleys, it is clear that we are confronted with an advancing and
Figure 879 specificall}^ distinct though closely related form of Archidiskodont.
From Sydney-on-Vaal, South Africa. Middle torrace (lower levels). I have named it after Mr. Millett who has been responsible for the
?TyOwer Plei.stoocno. finding of the three mammoth teeth reco\ei'ed at Sydney-on-Vaal."
Archidiskodon Dart, 1929.
milleili "Mammoths and Other "There remains, of course, the possibility that one of these
Fossil Elephants of the Vaal and Limpopo Watersheds." So. forms with a well-marked posterior buttress is closely related to
Afr. Journ. Sci., XXVI, p. 706. Type. Third superior—
molar of the left side, l.M^. McGregor Museum 4085; cast
Amer. Mus. 22722. Horizon and Locality. "Middle ter- —
race at a depth of 80 feet at Sydney-on-Vaal," South Africa.
?Pliocene [?Lower Pleistocene {fide Osborn, 1934.925, p. 3)]. Type
—
Figure. Dart, o-p. cit., p. 706, figs. 10 and 11.
Type Description.— (Dart, 1929.1, pp. 707-708): "In
general measurements (greatest length 222 mm., greatest width
108 mm., greatest height 118 mm.) it corresponds fairly closely
[with A. vanalpheni] being, however, considerably .shorter, wider
and low'cr than the foregoing. These measurements are fairly
comparable with those given ior A. vanalpheni, seeing that here, too,
the fangs are absent; here, too, eight plates are present including
a similar anterior talon, which in this case, however, more closely
approximates in height that of the second plate, and is actually in
wear. Here also one or more posterior plates are absent by loss.
(Figs. 10, 11 [Fig. 879 of present Memoir])."
"But it is on still closer examination that distinctions are to
be drawn between the teeth. While in this tooth, too, the indi\i-
dual plates are cuneiform, the disparity between their thickness
at base and edge is not so great, in that the plates are not more
than % inch thick at their bases and are %-}2 inch thick at their
edges. The degree of contraction in transvei-se length of plates
from base to edge, is, however, virtually identical with that in
A. vanalpheni." Fig 879. Ty|x> left third superior molar, l.M^ (McGregor Mus. 4085,
"The enamel is just as thick as that of A. vanalpheni and while cast Amer. Mus. 22722), of Archidiskodon milleili Dart, 1929, figs. 10and U,
it displays some crimping is not nearly so rugged in appearance as p. 706, one-third natural size. From Sydney-on-Vaal, South Afri<'a.

that of A. vanalpheni, while on the exposed sides of the plates

there is no trace whatever of longitudinal grooving. Owing to the if ^1. griqua of Haughton.
not identical with, I have had the
early age of this tooth also, which is practically identical in this examining that well-worn and fragmentary
privilege personally of
respect with that of the preceding, there is no clear picture of its specimen, and, with a view to making our knowledge of the South
surface in wear. The anterior talon is in wear and shows a median African mammoths as complete as possible, have taken the
posterior buttress as also do the second, third and fourth plates. liberty of making some additional remarks upon the material."
The fifth, seventh and eighth plates show five digitations, the
sixth plate six, the fourth plate six digitations and a posterior
buttress digitation, the third plate seven worn digitations and Archidiskodon loxodontoides Dart, 1929
a posterior buttress digitation, and the second plate two islets, the Figure 880
posterior buttress protruding from the medial one of the two From Sydney-on-Vaal, South Africa. Middle terrace (lower levels).

islets. There is no evidence of any anterior buttress in any of the ? Lower Pleistocene.

plates." Archidiskodon loxodontoides Dart, 1929. "Mammoths and

"The degree of separation between the second and third Other Fossil Elephants of the Vaal and Limpopo Watersheds."
plates, 'A-V/i in., is identical with what is typical for A. vanalpheni, So. Afr. Journ. Sci., XXVI, p. 709. Type.— Third superior
but the other plates are not nearly so widely separated being molar of the left .side, l.M'. McGregor Mu.seum (Kimberley)
992 OSBORN: THE PROBOSCIDEA

4087; cast. Amer. Miis. 22724. Horizon and Locality. — is evident, due to the presence of distinct buttresses here on both
"Middle terrace at a depth of 80 feet(?) at Sydney-on-Vaal," the anterior and posterior aspects of the plate. In the second and
South Africa. ?PHoceiie [?Lower Pleistocene, fide Osborn, 1934. first plates the islets have completely disappeared, and in the
925, p. 3]. Type Figure.— Dart, op. dt., p. 709, fig. 13. second plate especially there is shown a virtually median widening
Type Description.— (Dart, 1929.1, pp. 709-711): "This out with the full development of the buttresses."
tooth, also apparently a third upper molar (greatest length 149 "The plates, which are roughly 19 mm. wide in their more or
mm., greatest width 94 mm., greatest height 112 mm.), is less less parallel portions, expand to 31 mm. in the regions of the
complete than the previous two new types described, presenting buttresses with the result that the buttresses of adjacent plates are
only 4% in. plates and lacking in addition one or more plates both in direct contact with one another and together separate the inter-
anteriorly and posteriorly. It is impossible to state whether it lamellar stratum of cementum, 10-17 mm. wide, on one side from
j:)ossessed originally a ridge plate number as great as that of the that on the other side of adjacent buttresses, as occurs in the true
—
two preceding it probably did not. All the plates present were Loxodonts, a condition not previously encountered to my knowl-
in wear and the specimen was probably from an old adult. For an edge in any Archidiskodont, and justifying the name given to this
upper molar in early wear it is not higher than one would e.xpect in peculiar species."
A. subplanifrons, but it is narrower than that specimen and, "The cementum is abundant and almost encases the ridges
indeed, than all the aforementioned forms. Here also the fangs completely on the medial aspect of the tooth. Laterally, however,
are absent and the measurements of height consequently com- there are distinct and deep trenches ]{-% inches in depth between
jiarable (see Fig. 13 [Fig. 880 of present Memoir])." adjacent plates, indicating the probable origin of this species from
a form comparable with those previously described, especially
A. vanalpheni. Indeed, if it were not for the greatly reduced
width of the tooth and reduced width of interlamellar cementum
as well as the presence of the anterior buttress in A . luxodonloides,
I should have been considerably more reluctant to separate it

specifically from A. vanalpheni. It must be closely related to that

speciesand the recovery of further remains of both types, especially
a well worn molar of A. vanalpheni, will be awaited with great
interest in this connection. It may be repeated, however, that
there no evidence of such an unusual anterior buttress in A.
is

vanalpheni and the resultant loxodontoid form of plate in surface

wear as exists in A. loxodo7itoides. The same feature serves to
distinguish it equally from the A. griqua of Haughton."

Archidiskodon yorki Dart, 1929

Figure 881

Vanasswegen.shoek-Bloemheuvel, near Christiana, South .Africa. Middle

Fig. 880. Type left third superior molar, l.M^ (McGregor Mus. 4087, terrace (upper levels), ?Lower Pleistocene.
cast Amer. Mus. 22724), of Archidiskodnn Inxodontoides Dart, 1929, fig. 13, [).
Archidiskodon yorki Dart, 1929. "Mammoths and Other
709, oiic-tliird luttura! size. From Sydncy-oii-V'aal, South Africa.
Fossil Elephants of the Vaal and Limpopo Watersheds." So. Afr.
.lourn. Sci., XXVI, p. 717. Type. ". —
two very frag-
. .

"The plates are not so distinctively cuneiform as in the two mented enamel plates an Archidiskodont mammoth" {op. cit.,
of
previously described forms, nor do they present so distinct a de- p. 715). McGregor Museum (Kimberley) 4073. Horizon
gree of contraction in length toward the grinding surface where
—
AND Locality. "Middle gravel stratum of river bed gravels at
they show a transverse length of 80 mm. The enamel is thick Vanasswegenshoek-Bloemheuvel, near Christiana," South Africa.
(3-4 mm.) and isvery markedly plicated or crenated, despite its TMiddle Pleistocene [?Lower Pleistocene, ^rfe Osborn, 1934.925, p.
thickness. Longitudinal grooving of the medial and lateral 3]. Type Figure.— Dart, op. cit., p. 717, fig. 19.
aspects of the ridge plates is also strongly in evidence. Owing to Type Description.- (Dart, 1929.1, pp. 717, 718): "The
the well worn nature of the tooth the pattern produced on the plates of this tooth are of great size (fragment is 114 mm. broad
grinding surface is very clear. The most posterior or fifth plate and 193 mm. high, but the widest diameter of the tooth was at
shows 7 mamniillated processes which have all been broken off, least 120 mm. and the height at least 200 mm.). In all their
the fourth shows eight pedunculated processes in wear, the third mea,surements these plates (see Fig. 19 [Fig. 881 of ])resent
and fourth from the medial side having fused to form a small Memoir]) exceed considerably those of A. hroojni The enamel . . .

islet. In the third plate there are three islets, one formed by the plates measure 16-23 mm. across, that is, slightly narrower than
fusion of the medial two mamillae and one islet by the lateral two. in A. broomi, and ajjproach cio.sely in form the type presented by
The four central mamillae are fused to form a large central islet A. broomi. Laterally \iewed, howe\'er, there is a distance from
and in the region corresponding to the medial two of the four mid-point to mid-point of succeeding enamel ridge plates in A.
central mamillae a pronounced anterior and posterior dilatation broomi of at least 32 mm. whereas in this specimen the .same
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 993

measurement does not exceed 24 mm. thus demonstrating the SPECIES PROVISIONALLY REFERRED BY THE PRESENT
relatively decreased amount in it of the interlamellar cementum. AUTHOR TO THE GENUS PAL.a;OLOXODON
We are, therefore, dealing here with a broader and higher tooth Archidiskodon transvaalensis Dart, 1927
than in the type of A. broonii and one with slightly narrower
enamel plates and decreased interlamellar cementum. It must be From Bloemhof, Vaal River, South Africa. Pleistocene.

specifically different but closely related thereto and I will name it, A third superior molar, r.M^, described by Doctor Dart in
despite the paucity of the remains, Archidiskodon yorki in honour 1927 as Archidiskodon transvaalensis, was removed by Professor
of its discoverer." Osborn to the genus Palseoloxodon (see Osborn, 1934.925, pp. 2
"The finding of A. hroomi and A. yorki, which are advanced and 14). This species is treated fully in Chapter XIX, p. 1284,
Archidiskodont types in gravel stratawhich appear to be directly below (the Loxodontinse).
capable of correlation with the advanced Archidiskodont gravel
at Bloemhof, is of great interest and importance. Both of these
mammoths are admittedly more i)rimiti\'e than the Bloemhof Archidiskodon sheppardi Dart, 1927
mammoths, although more advanced than A. subplanifronti, which From Bloemhof, Vaal River, South Africa. Pleistocene.
came from the 50-60 feet dejjth at Sydney-on-Vaal." The species Archidiskodon sheppardi, type an l.M^, also de-
"There is no question in my mind, from the state of fossilisa- scribed by Doctor Dart at the same time as his species A. trans-
tion of A. yorki, from its fragmentation and from its unnatural vaalensis, was regarded by Professor Osborn as belonging to the
stratigraphical jjosition as compared with the Bloemhof mam- genus PalivoloTodon and as a consequence is treated in full in
moths, that it belongs to the more superficial part of the middle Chapter XIX, p. 1278 below (the Loxodontinse).
terrace, as I have already suggested is true also of .4. hroomi."

Archidiskodon andrewsi Dart, 1929

From Gong-Gong, ? Middle terrace (lower levels), Vaal River, South
Africa. ?Lower Pleistocene.
The type of Archidiskodon andrewsiDart is a fragmentary
third lower molar of the left side, I.M3 (McGregor Mus. 435; cast
Amer. Mus. 26968). It was first thought by Professor Osborn to be
referable to Archidiskodon subplanifrons, but on further study of
the original specimen kindly loaned by Curator Wilman of the
McGregor Museum he was inclined to regard it as belonging to the
genus Palseoloxodon (see Osborn, 1934.925). The description and
figure, therefore, will be found below on page 1278 of Chapter XIX
(the Loxodontinse).

Archidiskodon hanekomi Dart, 1929

From the old river bed of the Vaal River at a depth of 20 feet at Deli)oort's
Hope, South Africa.
In 1929 Dart described a ?third right upper molar, r.M',
in the McGregor Museum (No. 2930), which he named Archidisko-
don hanekomi. Professor Osborn, however, in his article "Primitive
Archidiskodon and Palaeolo.rodon of South Africa" (Osborn,

Fig. 881. Type molar fragment of Archidiskodim yorki Dart, 1929, fig.
1934.925) referred this species to the genus Palxolo.rodon. The
19 (McGregor Mus. 4073), found in river bed gravels at Vanasswegenshoek- description and figure will be found on page 1279, figure 1140 of
Bloemheuvel, near Christiana, South Africa. One-half natural size. Chapter XIX (the Loxodontinse).
 Genus: METARCHIDISKODON Osborn, 1934
Original reference: "Primitive Archidiskodon and I'alaeoloxoilon of Soutii Africa," Anier. Mils. Novitates, No. 741, August 24,
1934 (Osborn, 1934.925, p. 12).
Genotypic species: Loxodonta griqua Haughton, 1922.

—
Generic Characters. (Osborn, 1934.925, p. 12): "This group [Metarchidiskodon griqua group]
includes the fragmentary type (Fig. 3 [ = Fig. 882 of present Memoir]) of 'Loxodonta' griqua Haughton,
1922. This specimen appears to belong to a distinct form of grinding tooth to which the new generic
. . .

M
name elarchidiskodon may be applied, and distinguished from Archidiskodon as follows: (1) 3 with M
a relatively long narrow crown; index cannot be estimated at present. (2) Deep U-shaped valleys filled
with cement. (3) Enamel ridge plates extending to the bottom of the crown. (4) Very prominent
post-sinus folds instead of median sinus expansion of the typical Archidiskodon. Type species: Loxo- . . .

donta griqua Haughton, 1922." See Table IX above (p. 985) under "M. griqua group."

Specific Characters of Metarchidiskodon griqua, by Osborn, 1934. — (Osborn, 1934.925, p. 12): "(1)
Cement areas equal or exceed dentine areas; (2) pre-sinus folds absent or inconspicuous; very prominent post-
sinus folds; (3) very deep U-shaped valleys extending to the bottom of the crown; this is a very important point.
(4) These valleys are filled to the summit with cement. (5) Enamel ridge plates very deep, extending to the bottom
of the crown, closely compressed with very narrow dentinal areas between."

"Type figure 3 [
= Fig. 882 of present Memoir] to be compared with the relatively narrow grinding teeth of
similar molars observed in the Val d'Arno specimens and in British Mu-seum M12641, M12642."

Metarchidiskodon griqua Haughton, 1922 Rept. Geol. Gomm., 1907, pp. 171, 172) : "The gravels are situated
I'igurcs 882, 883 at \'arious levels as well as at varying distances from the river,

\'aal Rivor gravels, Griqualand West, Soutli Africa. ?Lo\ver Pleistocene. and though they form a number of fairly distinct terraces, it

From these gravels Fraas (1907) described Equun cf. zebra, is not always easy to determine their relati\'e age. ... On the west
of the Vaal the terrace finely developed at Klipdam, where it has
is
Hippopotamus amphibius var. rohuslus, Damaliscus sp. and Masto-
an altitude of 200 feet above the river and a distance from it of
don sp. Felix (see Beck, 1906.1, p.
; 49) described Mastodon
3}2 to 6 miles." (Osborn, 1934.925, pp. 2, 3) ?Middle terrace
(Biinohphodon) sp., and Haughton (1922) described a new giraffe
{Griquatherium cingulatum).
(upper levels), 60-80 feet, ?Lower Pleistocene. Type
History. — Originally described by Haughton in 1922 as
—
Figure. Haughton, op. cit., 1922, PI. i, figs. 1 and 2.

Loxodonta griqua this extremely fragmentary superior molar proves

—
Type Description. (Haughton, op. cit., 1922, p. 12): ". . .

a portion of an Elephant molar. The fragment consists of

belong to a new genus, Metarchidiskodon. The new type . . .

to
three plates of which one side is imperfect and show.s a vertical
figure (Fig. 882, right) is reconstructed from the original specimen
(McCiregor Mus. 3686) kindly loaned by Miss Wilman, Director
longitudinal section. . Although the molar is incomplete, it was
. .

obviously large and broad and fairly low. The enamel is thick, the
of the Museum. It apparently represents three central ridge-
borders show no strong crimping but a considerable amount of
plates ["Supposed third, fourth and fifth" (fide Osborn 1934.925,
unevenness. The cement wedges are large, the plates are wide
p. 8. fig. 3)] of a third left upper true molar (?1.M^), as compared
apart and were certainly few in number. In vertical section the
with .'i(c/iK//.sA-of/o/i planifrons (Fig. 829); the elevation of the ridge-
tooth has an appearance intermediate between those of E. africanus
plates and approximation of the valleys are less extreme than in the
and E. planifron>< figured by Falconer and somewhat similar to that
type of A. planifrons fFig. 825); the more worn anterior plate is
to the left, the less worn posterior plate is to the right; the concave
from the Red ( 'rag of Suffolk assigned by Leith Adams to E. anti-
quus (Brit. F'oss. F^lephants, PI. xxvi., fig. 3 [2|). The cement
side of each plate opens backward as in A. planifrons; the back-
wedges between the plates are as thick as the plates; and their
ward enamel loops are much more prominent than in A. subplani-
bases are more rounded than wedge-shaped. The sides of the
frons; width of central ridge-plate 90e mm., height of same ridge-
plates are more parallel than in planifronn, thus ap])roximating to
plate 90e mm. those of africanu.s. The most characteristic feature of the tooth,
Loxodonta (iriqna Haughton, 1922. "A Note on Some Fossils
however, is the shape of the wearing surface of the plates. ... [p.
from the Vaal Ri\cr (iravels." Trans. (Jeol. Soc. S. Africa, 1922,
13] The present tooth seems to differ considerably from that of
\'ol. .\XIV, ])p. 11 13. Typk.- -I'Vagmentary molar. Hoht- E. Zulu and also from that of E. antiqutts. These show a central
ZON AND Locality. — From the river gravels of the Vaal Kivcr, angular notch on the enamel, but never to the same extent; while
Griqualand West {op. vit., p. 11, (|Uoted from du Tdit, llth .\nii. the strong crimi)ing of the enamel in both these forms is a distin-

994
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 995

guishing feature, as well as the comparative lowness and width of

the tooth in this fragmentary specimen."
Specific Characters of Archidiskodon = Metarchi- [

diskodon] griqua, by Osborn, 1928. (1) Superior ridge-plates —

much more elevated, i.e., 3Ke in. =90e mm., than in the type of
A. subplaiiifrons, i.e., 2}2-2% in. = 64-70 mm. (2) Central enamel
loops much more prominent than in A. subplanifrons. (3) Width
of centralridge-plate 90e mm. ; height of central ridge-])late
90e mm.
Kaiso Bonp>beds (Hopwood, 1926, pp. 33, 34). — From the
Kaiso Bone-beds, near Albert Nyanza, Hojjwood describes a frag-
mentary tooth (Brit. Mus. I\I12641), see figure 883, as follows:
"The chief specimen consists of three lamellae and the loxodont
sinus of a fourth. At the anterior end is part of the posterior wall
of a fifth plate which has been deejily abraded. The one behind it
has been worn in such a manner as to reveal the great depth of the Srzt.Mtis M./Z6^f
digitations. This sjjecimen has been cut across in order to show
the enamel figures [Fig. 883, lower]. . . . These characters, taken
together with the lamellar frequency of four, indicate a very
primitive form, which might be compared with E. pla/iifrons
Falconer."
External LM^
Y2 nat. size

FIG. \i.—Elephaa afl. mcridionalU Nesti. Enamel pattern of larger

frnnrneiit. Rcc'i. Ml'26-ll. 1/-2 iml. .?i/,c.

FIG. H.—Elephas aff. meridioyxalis Nesti. Development of the enamel

pattern in an isolated plate. A is 13.5 mm. from the top; B

McQregorMus. 5686 6 mm. from A: C, 8 mm. from B. und D 12 5 from C mm

Begd. Mia642. 1/2 nat. size.
Type. Loxodonta griqua
Haughton. 1922
Fig. 883. Referred Metarchidiskodon griqua (non meridionalis)
Type Third Left Superior Mol.\r ok Metarchidiskodon griqua (Upper) Afragmentary molar from the Kaiso Bone-beds of Africa, doubt-
Fig. 882. Original (upper,},4 lowor,}^ nat. size) and new (riglit pair,K nat. fully referred by Hopwood to Elephas = A rchidiskodoyi] meridionalis but which
[

size)type figures of Loxodonta griqua Haughton, 1922, PI. i, figs. 1 and 2; both proves to be closer to Metarchidiskodon griqua. After photograph kindly fur-
drawings after fractured type specimen (?1.M') in the McGregor Museum at nished the present author by Dr. A. Tindell Hopwood (cf. Hopwood, 1926, PI.
Kinil)erley (McGregor Mus. 3686). From the river gravels of the Vaal, Gri- Ill, fig. 2). Brit. Mus. M12641. Natural size.
qualand West, South .\frica. Original type figures (left): ''Fig. 1.— Cirindiiig (Middle) Mid-section of the worn coronal surface. Aft(-r Hopwood, 1926,
surface of fragmentary molar. . . . Fig. 2. — Profile view of same." fig. 13, p. 34. One-half natural .size. Brit. Mus. M 12641. Ob.serve the very
(Right) New figure after Osborn, 1934.92.5, p. 8, fig. 3: Suppo.scd third, prominent median 'loxodont sinus.'
fourth, and an I.M^. Observe very deep U-shaped valleys,
fifth ridge-plates of (Lower) Summit of the worn coronal surface. After Hopwood, 1926,
thick simply folded enamel with very prominent looped post-sinus folds. fig. 14, p. 34. One-half natural size. Brit. Mus. M12642.
 .

996 OSBORN: THE PROBOSCIDEA

This tooth, the enamel ])Iate characters of which arc best, lameHae 100 nun., breadth 72 nun., height 105 mm., as compared
shown in figure 883 (lower), appears to resemble Metarchidi.skodon with the following corresponding measurements in the type of
griquamore closely than Archidiskodon subplanifrons, because the M. griqua, breadth of center ridge-plate 90e mm., height 90e
enamel ridge-plates are more elevated, the measurements given mm. This ridge-j)latc height exceeds that of the fifth ridge-plate
by Hopwood being {op. cit., 1926, table, p. 35), length of four of /I. AM67J?nM7/70«s, namely, 53e mm., with a breadth of 92 mm.

3. ARCHIDISKODONTS OF THE UNITED STATES AND MEXICO

This noble genus appears to have been given origin in Africa, then to have migrated to Eurasia along the
south temperate parallels, and finally to have reached its climax of evolution in the United States from Nebraska
southward to the high plateau of Mexico^

Two quite distinct lines of descent, referable to Parelephas and Archidiskodon respectively, are found in the
United States and Mexico, in which the low ridge formulae are very similar, namely:

Elephas [
= Parelephas] coluinbi Falconer, 1857-1868. Upper Pleistocene of southern United States and of Mexico.
Smaller animal than Archidiskodon, witii narrower grinders (Fig. 887), thin cement outer coating; maximum ridge-
plate formula,
1 8-1 9
3 ^-grj i+- M
Elephas [
= Archidiskodon] imperalor, Leidy, 1858. Lower- Pleistocene of southern United States and of Mexico.
Larger animal in size, with broader grinders (Fig. 887), very broad enamel plates, and heavy cement outer coating;
ridge-plate formula, M3 J^rH

—
Geologic Age. Falconer erred in considering these animals of the same species; they are really very
distinct. They were not geologically contemporaneous; to our knowledge the true Archidiskodon imperator and

true Parelephas columbi are not found in the same horizons. A. imperator is a late Pliocene and early Pleistocene
species (Nebraskan and Aftonian glacial stages, cf. Hay, 1923, p. 15), while P. columbi is probably a late Pleisto-
cene species (during lowan to Wisconsin stages, abundant in the phosphate beds of South Carolina and Florida,
cf. Hay, op. cit., p. 431, map 12, also pp. 430 and 155). The true P. columbi of Georgia, South Carolina, and
Florida are not to be confused with the Parelephas jeffersonii of the northern states (cf. Hay, op. cit., map 12, p.

431, error). Gidley held that A. imperator and P. columbi were contemporaneous.

GiDLEY, 1928. — Gidley observes

December 6) "There can be no question that remains of Archidisko-
(letter, :

don imperator and Parelephas columbi, as these two species are now defined by Osborn [in the present Memoir],
are found associated in the No. 2 beds of Sellards at Melbourne and Vero, and it is equally certain, I think, that
these species were contemporaneous in Florida. From my work at Melbourne and Vero last winter I conclude
that the 'No. 2' deposit is almost entirely of wind-blown sand origin, slowly accumulated, and that stream action
had little or nothing to do with its formation. On this theory mixtures of material of different periods would not
be possible. ... I believe it . . . probable that descendants of A. imperator existed in Florida as well as in Mexico
and the south, until Upper Pleistocene times." Recently described from Melbourne and Vero, Florida, by
Gidley, are:

Parelephas floridanus: Nat. Mus. 118U6, 118U8, and 11810 (sec Parelephas, Chap. XVII, footnote, p.
1078).
Archidiskodon imperator: Nat. Mus. 11805, 11814, and 11G20 (see below, p. 1005).

'[For Professor Osborii's final opinion on the migration of Archidiskodon from the V'aal Rivor of South Africa to the Niobrara Kivor of Nebraska, see
Chap. XXIII, p. 1580.— Editor.]
-[Early and Middle Pleistocene (Jide Lugn and .Suhullz, 1934.1, pp. 373-37(5). — Editor.)
 :

THE MAMMONTIN.E: ARCHIDISKODON AND METARCHIDISKODON 997

ORDER OF DISCOVERY IN AMERICA OF ARCHIDISKODON AND FARELEPHAS

Since the opening sections of the present chapter were written several years ago, our knowledge of the two
distinct lines of Mammontine descent found in the United States and Mexico has been greatly enriched by three
more or less complete fossil skeletons of Parelephas discovered in Florida, also by intense and more extended
observation on the differences between the Parelephas and the Archidiskodon crania as well as by careful compari-
son of the Parelephas columbi grinding teeth with those of Archidiskodon imperator.

Falconer's type of 'Elephas' columbi is the middle portion of a third inferior grinder in which the ridge-plates
are very far apart, thus resembling the widely separated ridge-plates of Archidiskodon imperator; he concluded,
therefore, that Leidy's 'Elephas^ imperator was the same species as his own 'Elephas' columbi. Osborn was long
misled by these widely separated ridge-plates and erroneously concluded that the type of 'E.' columbi belonged in
the genus Archidiskodon; whereas the newly discovered materials above mentioned prove that 'E.' columbi
belongs within the phylum Parelephas. But, as will be shown in a subsequent chapter (Chap. XVII), the species
Parelephas columbi Falc, 1857, is clearly distinct from the species Parelephas jeffersonii Osborn, 1922. Thus the
two quite distinct lines of descent found chiefly in the southern United States' and in Mexico appear as follows in
the order of discovery

Archidiskodon Phylum Parelephas Phylum

1858 Elephas imperator Leidy, ^ehTaska. = Archidiskodun im- 1838 Elephas jacksoni Mather, Ohio = Parelephas jarksoni{'!)
perator
1857 Elephas columbi Falconer, Georgia. = Parelephas columbi
1915 Elephas hayi Barbour, Nebraska = 4 rt'/i?f//sA'odoH hayi
1922 Elephas Columbi var. silvestris Freudenberg, Mexico 1859-1861 Elephas texianus Owen-Blake, Texas = Parelephas
= Archidiskodon imperator silvestris columbi
1922 Elephas Columbi var. Falconeri Freudenberg, Mexico
1922 Elephas Columbi var. Felicis Freudenberg, Mexico = Parele-
= Archidiskodon imperator falconeri
phas columbi felicis
1925 Elephas scotti Barbour, Nebraska = Archidiskodon impe-
rator scotti (or juvenile A. imperator) 1922 Elephas jeffersonii Osborn, Indiana = Parelephas jeffersonii
1925 Elephas (Archidiskodon) Barbour, niaibeyii Nebraska
1927 Elephas roosevelli Hay, Il\iDois = Parelephas jeffersonii
= Archidiskodon imperator maibeni
1928 Elephas haroldcooki Hay, Ok\&homa. = Archidiskodon har- 1929 Parelephas floridanus Osborn, Florida = Pa?-ctep/ias Jlori-

oldcooki danus
1928 Elephas exilis Stock and Furlong, California = .4 ;y/u'-
1929 Parelephas columbi cayennensis Osborn, French Cuiana,
diskodon e.rilts
South America = Parefep/ios columbi cayennensis
1929 Archidiskodon sonoriensis Osborn, Mexico = Archidiskodon
sonoriensis
Narrower grinding teeth; thinner enamel ridge-plates; thin
Very broad grinding teeth; thick enamel ridge-plates; cement outer coating; enamel ridge-plates widely .separated or
heavy cement outer coating enamel ridge-plates widely separated
; arcuate at the base (as in type of 'E.' columbi), more closely con-
from the wearing surface of the crown downwards to the face of the vergent at the summit; ridge-plates increasing from 3 j^ M
plates. Ridge-plate formula not known to exceed M 3 iv. (P. columbi) to M3 Iff (P. progressus).

It would be natural to suppose that from the progressive A. meridionalis stage these southern manmioths
migrated across Asia and through the United States southward, but the present palseontologic and geologic evi-
dence does not support this direct derivation of the imperial mammoth from the progressive A. meridionalis type,-

'[In Edmunds of the Department of Geology of the Univeraity of Saskatchewan forwarded to the American Museum for identi-
1932 Professor F. H.
fication a proboscidean molar from the vicinity of Wiscton, Saskatchewan, which proved upon examination by Dr. Edwin H. Colbert of the Museum staff to
be referable to Archidiskodon imperator. This is important as establishing the most northerly range of the genus in North America. Editor.] —
'-(Subsequently Professor Osborn became convinced that the "Elephas meridionalis" of France was the direct ancestor of the "Elephas imperator" of
North .America, a conclusion arrived at through the discovery of a skeleton (named by Osborn Archidiskodon meridiorMlis nebrascensis), foimd one mile north-
west of Angus, Nuckolls County, Nebraska (see Osborn, 1932.893, p. 1): "New and positive evidence of the correctness of this theory is now afforded by the
discovery of the complete skeleton which forms the subject of the present paper. This skeleton with the lower jaw in a complete state of preservation proves to
resemble very closely indeed in every detail the 'Elephas meridionalis' of Durfort, France, as fully described by .-Mbcrt Gaudry." Editor.] —
998 OSBORN: THE PROBOSCIDEA

because since the year 1915 there have been discovered at least two specific stages very much more primitive than
A. imperator. These are the Archidiskodon hayi Barbour, from near Crete, Saline County, Nebraska, also the
Archidiskodon sonoriensis from Sonora, Mexico, with its prominent elongate protuberance of the rostrum (Fig. 923,
cf. Fig. 903). Of these relatively primitive forms, A. hayi (Fig. 913, lower) has an elongate mandibular ramus,
with a third inferior molar exhibiting from ten to eleven ridge-plates set very far apart. This not only appears
to be the most primitive species of proboscidean thus far discovered in America, but it seems to resemble very
closely the primitive lower jaws (Fig. 849) of A. planifrons of southern France and of India.

The Archidiskodon sonoriensis is a far more progressive species than A. hayi, but it exhibits an elongated
mandibular rostrum of a much more primitive character (Fig. 923) than the brevirostral mandibular symphysis
of A. imperator (Fig. 892 A, B, 898 A, Al).

These discoveries render it highly probable that the first migration from Eurasia to America occurred during
the Upper Pliocene Archidiskodon planifrons phase; they also render it possible that the typical A. planifrons
of the Pinjor horizon of India migrated to America at about the same period, and perhaps as a traveling companion

of the ancestors of Stegomastodon mirijicus.

SYSTEMATIC DESCRIPTION OF SPECIES

Archidiskodon imperator Leidy, 1858 Specific Charactp:rs, Osborn, 1928. —
(1) Relatively short

Figures 80o, 810. 812, 815, 817, 818, 805, 884-889, 891, 892, 894- and broad superior and inferior grinding teeth completely sur-
902, 906-909, 912, 915, 916, 920, 937, 1030, 1226, 1231, PI. xxi rounded by a heavy layer of cement. (2) Ridge-plate formula:
Lower Pleistocene, ('i)lsl Interglacial, or (?)Aftonian age. Central and M 3 1 8-V9-2 ^^ superior molars observed in which the ridge-
i

Southern United States, 40th parallel southward into Mexico. Osborn, 1921-
plates exceed eighteen. (3) Brachycephalic and hypsicephalic
1924: The unrecorded level is probably equivalent to the Equus beds of Cope,
portions of cranium correlated with the excessively short, broad,
thv Equui excelsus-Elephas imperator zone of Osborn (Osborn, 1918.473, p. 34).
Hay, 1924, p. 100: "Seneca, Thomas County (6 [see map, Fig. 8901).— and elevated superior molar teeth (Figs. 889B, 888), also with the
Somewhere along Middle Loup Fork River, probably in Thomas County, was relatively broad and short inferior molar teeth (Figs. 889A, 892B,
found, by F. V. Hayden's party, the tooth which forms the type of Elcphas left, 898). (4) Lower .jaw short, deep, excessively broadened or
imperator. A more exact locality has not been determined. In the region swollen to accommodate the broad inferior grinding teeth (Figs.
about Seneca, as reported to the writer by Dr. W. D. Matthew, were found, in
898, 892); normally 16-18, may include im-
inferior ridge-plates
1916, remains of Equus, Camelops, Platygnnus, Canis, etc. Here, too, Hayden
probably foimd tlie ty|ie of Stegomastodon mirificus."^ perfect ridge-plates 19-20 (Fig. 892A). Jaws much more abbrevi-
Synonyms or related forms (Mexico): El. Volumbi var. silvestris Frcud- ate than in Archidiskodon planifrons (Fig. 865), much more mas-
enb., 1922, Oaxaca, Mexico; El. Cnlumbi var. Falrxmeri Freudenb., 1922, sive and swollen than in A. meridionalis or any other species of
Tecpiixquiac, Mexico; El. Columbi var. imperator Freudenb., 1922.
proboscidean; rostrum deep and short; symphysis rounded; the
Synonyms or related forms (United States): Elfphas scnili Barbour, 1925;
superior border of the coronoid process greatly raised above the
('!)Elepha.s (Archidiskodon) maibeni Barbour 1925, 1926, Elcphas haroldcooki
Hay, 1928. See type descriptions of these synonymous or related forms below. grinding surface (Fig. 898).
Elephas imperator Leidy, 1858. [On new species of Mastodon
This imperial species, well named Elephas imperator by Leidy
and Elephant from Nebraska, Mastodon mirijicus, Elephas im-
because of its commanding size, appears to be a direct descendant-
perator.] Proc. Acad. Nat. Sci. Phila., March, 1858, Vol. X, p.
of Archidiskodon meridionalis and of A. planifrons of southern
10 (Leidy, 1858.2) also "Notice of Remains of Extinct Verte-
;

luirasia. It progresses beyond the A. meridionalis stage, the

brata. from the Valley of the Niobrara River," np. cit., p. 29
. . .

number of plates in its grinding teeth rising to eighteen (ma.\.

(Leidy, 1858.4). Type.— (Op. cit., 1858.2, p. 10): ". . . part
twenty), but it enamel and the wide borders
retains the very thiclv
of an upper molar tooth of an Elephant from the Niobrara; . . . The
of cement. Inasmuch as A. meridionalis extended from Upper
Pliocene into Lower Pleistocene time in western Kuroijc, it may by
name Elephas imperator."
species he proposed lo distinguish by the

migration have given rise to A. imperator- which seems to char- {Op. cit., "The fragment of the tooth has been
1858.4, p. 29):

acterize Lower Pleistocene (?Aftonian) time, especially in the west- assumed to belong to an unnamed species from the fact that it
ern and southwestern United States and in Mexico; then this was found in association witli a fauna very distinct from any
imperial mammoth appears to liave become extinct.^ previously noticed." Nat. Mus. 185; cast Amer. Mus. 2568.

'(Lugn and Schultz (1934.1, p. 373) give the type locality of Eleplias [Archidiskodon] imperator as: "Pawnee. Loup Briuicli of Plaltc River = Middle
Loup, probably Hooki^ Co. [Nebraska], 1858." Editor.] —
'-[See footnote 2 on previous page — Editor.]
'[See footnote 2 on page 9% above.^Editor.]
 :

THE MAMMONTINyE: ARCHIDISKODON AND METARCHIDISKODON 999

Horizon and Locality. — (Leidy, 1858.2): "Valley of the a peculiar fauna, an associate of the Mastodon mirificus, as the
Niobrara river, Nebraska." (Leidy, 1869, p. 254) "The fragment : ordinary E. americanus was of the M. americanus. The specimen . . .

of a molar tooth originally referred to a .species with the name of assigned to E. imperator is represented in figure 3, |)late xxv, one-
Elephas imperator was obtained by Prof. Hayden on the Loup third the diameter of nature. It exhibits the characters attributetl
Fork of the Platte River." (Hay, 1914, pp. 421, 422) Type col- by Dr. Falconer to E. Columbi, compared with the supposed
lected from Loup Fork of Platte River, now in the National American variety of E. primigcnins. The specimen is the fore part
Maseum, designated Nat. Mus. 185. (As quoted above from Hay, of an upper molar, probably the fifth. The triturating surface,
1924, p. 100) Probably Seneca, Thomas County, Nebraska. extending the breadth of the fragment, is nearly five inches at its
(Lugn and Schultz, 1934.1, p. 373) "Pawnee Loup Branch of widest part. The breadth on the less broken side is about seven
Platte River = Middle Loup, probably Hooker Co. [Nebraska], and a half inches, and contains only as many ridges, that is to —
1858." Type Figure.— Leidy, 1869, PI. xxv, fig. 3. say, one ridge to an inch of breadth. There is, however, a thick

E, imperator
Nal Mus. 185. Type (shaded!
Amer, Mus. I187t. rev. foutlin

Type Third Right Superior Molar of Archidiskodon imperator, and Reconstruction

Fig. 884. Type figure of Elephas imperator Fig. 885. Type molar, r.M', of Elephas Fig. 886. Leidy's type molar (shaded) and
Leidy, 1858. After Leidy, 1869, PI. xxv, fig. 3. imperator, crown view. After Osborn, 1922.555, Osborn's neotype (outline) combined, both be-
One-third natural size. Fragment of a third I). 4, fig. 4. longing to M^ of the right side, .\fter Osborn,
right superior molar, r.M',from Nebraska (Nat. Tliis tootli is in the same position as in 1922.555, p. 4, fig. 4. Neotype (Amer. Mus.
Mus. 185; cast Amer. Mus. 2568). Inverted. Leidy's type drawing (Fig. 884), namely, anterior 11871) from Guadalajara, Jalisco, Mexico. See
The anterior surface is to the left; ridge-crests surface to the left, posterior surface to the right. figure 888, showing eighteen ridge-plates.
1-8 are preserved (cf. Fig. 886), ridge-plates
concave posteriorly.

Type Description, 1858.— (Leidy, 1858.4, p. 29): "The talon in front, and the ridges curve considerably backward on the
Niobrara collection also contains the anterior portion of an upper less broken side of the tooth. The four more perfect ridges of the
molar tooth [Figs. 884-886] of an Elephant of larger proportions specimen at the middle of the triturating surface occupy a space of
than any which are known to us. The triturating surface is a little over three inches, including the three intermediate plates of
within a line or two of five inches in breadth, and within a space of cementum. The widest ridges measure four and a half inches. The
seven inches only eight enamel folds or double plates exist. In the enamel is thick and strongly crimped, and the dentinal tracts are
most thick plated variety of teeth of the Elephas americanus which slightly dilated at their middle. Elephas imperator, if not regarded
we have seen, in the same space ten folds were counted. As in the as a peculiar species of a peculiar fauna, may be viewed, together
latter, E. primigenius, and the recent Elephant of India, the enamel with those teeth which have been referred to E. Cohimhi, as belong-
plates become worn on the triturating surface into transverse, ing to the Elephas americanus."
strongly crenulated ellipses." (Osborn, 1924: (1) In his second description Leidy correctly
—
Second Description, 1869. (Leidy, 1869, pp. 254, 255) designated the type horizon as "the Loup Fork of the Platte
"The fragment of a molar tooth originally referred to a species with River," Nebraska. Doubtless influenced by the high authority
(2)
the name of Elephas imperator was obtained by Prof. Hayden on of Falconer (1863, pp. 58, 59 cited below), Leidy was inclined to
the Loup Fork of the Platte River. I was led to refer it to a abandon the name Elephas imperator and to refer this type to
species different from the more ordinary American Elephant, from Elephas columbi, and finally (see Leidy, 1869, pp. 251, 252, 255;
its greater size, the comparative coarseness of the constituent 1871, p. 359; 1877, p. 213) to E. americanus De Kay, a species
elements, together with the fact that it appeared to be a member of representing the American variety of Elephas primigenius, namely.
1000 USBORN: THE PROBOSCIDEA

Mnmmonteun priinigctn'us nmcricnnuf: of the jiresent Memoir (see p. T.,eidy adds, 'that the fragment of (he tooth has been assumed to be-
1156). This was de.arly a scries of unfortunate errors on Leidy's long to an unnamed species, from the
it was found in as- fact that
part, due to lack of material. any previously noticed.' "
sociation with a fauna very distinct from
In a subse(iuent passage in the same ])aper (1863, p. 67),
FALCONER ERRONEOUSLY UNITES (1863) ELEPHAS IMPERATOR Falconer observes that in his opinion there arc "but two well-
WITH ELEPHAS COLUMBI determined species of fossil Elephant known in North America,"
(Falconer, 1863, pp. 58, 59, 67): "The second case is more namely: (1) E. pn'migenius (syn. E. americaniis). (2) E. cohimbi
remarkable and important, being that of the Elephant of the
fossil (syn. E. primigenius, pro parte; E. lexianus). He concludes (p.
Pliocene Fauna of Niobrara, an affluent of the Missouri River, in 67): "The same, with our present knowledge, must be said of the
Nebraska, the account of which, by Dr. Leidy, has excited much

E, Imp^ratoi
Neotype

'34 567
E. imperatop
Amer. Mus. 11B71
Neotype

1/4 not. size

Types of Archidiskodon imperatou (upper) and Parelepiias
COLUMBI (lower)
lis- 887. Elephas columbi (lower figure), fragment, of a third right inferior Neotype of Archidiskodon imperator
molar, r.M.^, two-sevenths natural size, and Elephas imprrnlor (ujiper figure), Fig. 888. From Guadalajara, .laliseo, Mexieo. B 2, Neotype superior
fragment of a third right superior molar, r.M', two-fifths natural size, photo- molar of Archidiskodon iiripcrnlor, M'' of the right, side (Amer. Mu.s. 11871),
graphed directly from the t.yp(^ easts of PJ. columhi {Kmov. Mus. 1747) and inner view, eighteen ridge-|)lates. B, The jsame, crown view. One-fourth
E. imperator (Anier. Mua. 2568). Compare figures 918 and 881. natural size. After Osborn, 1922..').').'), j). 5, tig. o. See figure 886 Al, in outline.

interestand surprise among Palseontologists. [Footnote: 'Proceed. E. imperator of Leidy, from Niobrara. Until a i)erfect molar is
Acad. Nat. Scien. Philadelp., 1858, p. 20, et .seg.'] ... 1 Mastodon figured and described, no satisfactory opinion can be formed as to
of the sub-genus Telralophodon, M. mirificus, Leidy, and a huge what the species is. Dr. Leidy, as already stated, iissumed it to be
Flephant, E. [Eueleph.) imperator, Leidy. The published descrip- distinct,and gave it the name upon the asstuuption."
tive details of this Elephant are as yet but very meagre. One speci- That Falconer erred as to synonymy is clearly demonstrated in
men only is mentioned, being the anterior i)ortion of an upper the accomi)anying figure (Fig. 887) in which we observe that
molar, of larger dimensions than any known to the author. The while the enamel ridges are equidistant in Archidiskodon imperator
crown is stated to be 'within a line or two of five inches in breadth, anil in Parelepha.s colnmbi, antl that while the riilge formula
and within a space of seven inches, only eight enamel folds or is aijproximately similar in the two species, namely, 3 J^tH (*4- M
double i)lates exist.' This would give an a\crage of nearly ninc- imperator), ^r^.'^^j^ (P- columbi), we also observe that the molar of
tenths of an inch to each ridge, corresponding closely with the A. imperator (u])pcr) is much broader and is surrounded with
proportions yielded by E. Columbi. The ridges are described as a broad layer of cement, while the molar of P. columbi (lower) is
becoming worn into transverse strongly crenelated ellipses. Dr. much narrower and is lacking in cement.
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1001

OSBORN SEPARATES (1922) ELEPHAS IMPERATOR LEIDY FROM us to determine exactly to what portion of the complete neotype
ELEPHAS COLUMBI FALCONER tooth (Fig. 5 = Fig.
888 of the present Memoir]) this ancient and
[

In reexamining and comparing the fragmentary molar types of much battered type belonged; the eight ridge-plates of the type
Elephas columbi and E. imperator, Osborn (1922.555, pp. 1-7) [Nat. Mus. 185] which are jireserved, in comparison with those of
determined with great care the exact position and character of both the neotype (Amer. Mus. 11871), constitute the anteroposterior
type fragments as shown in figures 949, 951A, 951B (for E. co- portion of a much-worn molar, M^ of the right side, in which thir-
lumbi) and in figure 886 (for E. imperator) of the jjresent Memoir.
, teen ridge-phites were in use out of an estimated total of seventeen
The E. imperator tyjje iiro\'es to be the anterior portion of a third [eighteen]. Of these plates five occupy a line 100 mm. long; this is
right superior molar containing portions of the first to the eighth because the ridge-plates are arcuate and widest apart in the middle
ridge-plates greatly worn, anil broken away from ridge-plates nine portion of the crown. The neotype tooth (Amer. Mus. 11871),

E. imperator

Amor. Mus. 14558 Rel

1/4 nat. sizo

Fig. 889. Superior (B) and inforior (A) molars of reforred Amerioan Museum .speeimcns, one-fourth natural size.
.Irc/iirfisAvK/iwf i'mpprator, Tlie.se two
individuals are believed to be of e(jrr('spoiiding age. They exhibit, (M^, M3) meclianieal reversal of the convex and concave .surfaces both in the crown
contours, crown surfaces, and ridge-plates. After Osborn, 1922.555, p. 0, fig. 6.

A, Inferior molars (Amer. Mus. 14558), from Ness County, Kan.sas, with 14-15 ridge-plates in u.se, a total of 19 vi.sible in left molar. See also figure 892A,
where a total of 20 ridge-plates are visible owing to artificial exposure.

B, Superior molars (Amer. Mus. 14476), from Victoria, Victoria County, Texas, with 14-15 ridge-plates in use, a total of 154-- Inferior view of this skull
shown in figure 897. See also figure 896.

to seventeen, as clearly shown in outline in the accompanying from Guadalajara, Jalisco, Mexico, appears to attain the full size of

figure 886 Al. At the same time Osborn selected as a neotype the superior grinders of this species of mammoth ; the ridge formida
the third superior molar tooth (Amer. Mus. 11871) from Guadala- may be written M 3 1 8-1 9 [M
L-^ -^ 3
'-'
1 8-1 9-20J This accords with the
jara, Jalisco, Mexico, from which the outline in figure 886 Al is actual average count of the ridge-plates in E. imperator by Hay
drawn. (1914) and by Osborn (1921-1922) in individuals which can with-
Characters of the Archidiskodon imperator Type and out question be referred to E. imperator. ... In the neotype (Fig.
Neotype.— (Cf. Osborn, 1922.555, pp. 3-5) "We are indebted to: 5 =Fig. 888 of the present Memoir]) thirteen plates were in use;
[

the National Museum for the loan of the Elephas imperator type in the referred skull (Amer. Mus. 14476) fifteen plates were in use
specimen (Fig. 4 [=Fig. 886 of the pre.sent Memoir]), enabling (Fig. 6B = Fig.
[ 889 B of the present Memoir]); in the referred
1002 OSBORN: THEPROBOSCIDEA

lower jaw (Anier. Mus. 14558) fifteen plates were in use (Fig. 6A Meanwhile Hay (1914) made a series of observations on the
[
= Fig. 889 of the present Memoir)). The total ridge-plates in M3 type and other grinding teeth truly referable to E. = A.] imptrator [

attain nineteen [twenty], as clearly shown in Fig. 6A = Figs. 889A

[ in various museums, from which the following citations may be
and 892A of the jiresent Memoir]." We have, therefore, the fol- made,
lowing ritlge formulae (cf. specific definition above).
Upper Pleistocene. Elephas [
= Parelephas] columbi of the HAY'S (1914, 1923, 1924) OBSERVATIONS ON ELEPHAS IMPERATOR
southern United States: M3 r^.\-a'+- Osborn summarizes below (pp. 1087, 1088) Hay's observations
Lower Pleistocene'. Elephas [
= Archidiskodon] imperatnr of on Elephas = Parelephas] columbi and geographic distribution
[

the southeastern and western United Slates: 3 isMa-l o- M records, as well as of E. [

= Arrhidlsk(Hlon] imperator in the region
Upper Pliocene to Lower Pleistocene. Elephas [=Archidisko- west of the Mississippi River (Fig. 890); also Osborn points out
do>i\ men'diunalis, ancestral, of western Europe: 3 titH- M (P- 699) the clear distinction between the true E. = P.] colwmbi [

(Osborn, 1922.555, p. 5): "The cranial characters observed (as limited by Osborn) and the true i?. [.4.] /7«pe?-a<or of Leidy.
in three more or less complete skulls referred to Elephas imperator Dkntal Characters (Hay, 1914, p. 421). It now remains —
tend to support the direct descent of this animal from the E. to summarize Hay's observations of 1914 on the grinding teeth
meridionalis of the ^'al d'Arno, Upper Pliocene of Italy." discovered near the type locality of Nebraska and on those found
Osborn, 1924: Thus £. [
= Archidiskodon] imperator appears in Oklahoma and Kan.sas (see Fig. 890) "Tooth formula not well
:

to be firndy established in its dental characters and ridge-plate known; the hindermost molars having apparently from sixteen
formula as a probable successor of £. = ^4.] meridionalis, whereas [ to twenty ridge-plates; the teeth large, with thick ridge-i)lates and
E. = Parelephas] columbi is clearly distinguished as a smaller form
{ thick enamel; the ridge-plates often concave on their hinder face
with narrow molar teeth which lack the very broadly encircling and more or less warped those of the hinder half of the lower teeth
;

layer of cement but exhibit practically the same ridge formula, leaning strongly forward. This is a species not yet well known,
Freudenberg (1922) has suggested that the true E. = P.] columbi [ although various parts of its skeleton have been collected." The
is geologically a successor of the true E. [
= .4.] imperator stage. chief materials observed and enumerated by Hay are:

Xat. Mus. 185 M^ fragment (Loup Fork of Platte River, Neb.), eight anterior ridge-plates. {Op. cit., p. 422): "The breadth of the
Type grinding face is 125 mm., but of this about 5 mm. on
each side belong to the cement. There are hardly five ridge-
plates in a line 100 mm. long and crossing the plates at right angles. The enamel plates are each nearly as thick
as the layer of dentine enclosed by them. The plates of cement intervening between the ridge-plates are somewhat,
but not greatly thicker than the plates of dentine. The face of each enamel plate which is directed toward the
l)late of cement is moderately striated from the base to the summit, but this striation is not deep enough to be
called crimping. At their inner and outer margins the ridge-plates are turned backward so as to make each one
deeply concave on the hinder face, convex on the front face."

Nat. Mus. 2216 M^ (Afton, Okla.). {Op. cit., p. 422): "The surface of wear extends back to the ninth plate. There are counted
sixteen plates. The anterior talon is mi.ssing on account of wear, and the posterior one broken off. It seems not
is

unlikely that at least two ridge-plates are missing, either in front or behind. The plates are very thick, there
being hardlj' five of them in a 100 mm. line."
{Op. cit., p. "In another tooth, which is broken, it is seen that the hinder faces of the plates are considerably
422):
dish-shaped. be ob-served likewise, that the plates converge toward their summits. The base of the tooth
It will
is very convex. The length of the tooth, from the base of the first ridge-plate to that of the hindermost, is 350 mm.

The thickness is 126 mm., more than one-third the length of the tooth. The height of the ninth plate is 250 mm.
As will be noted, the plane of wear, at the stage represented by this tooth, strikes the summits of the plates very
obliquely."

Nat. Mus. 2217 M3 (Afton, Okla.). {Op. cit., p. 423) : "The length from the base of the anterior plate present to that of the hinder one,
is 297 mm. The greatest width, taken at one-half the height of the tenth jilate, is 125 mm.; the height of the
ninth ])Iate is, in a straight line, 170mm. At the middle of the inner and outer faces there are only three and
a half ridge-plates crossed by a line 100 mm. long. At the rear there are five plates in such a line. There are to be
counted sixteen ridge-plates and a posterior talon. These are about as thick as the cement plates. The enamel is

thick and considerably crimped."

Phil. Acad. M3 (Wellington, Kan.?). {Op. cit., p. 424): "It is i)retty certain that one ridge-plate is missing in front, and one or
more at the rear. Eighteen plates are present. From the front of this tooth to the base of the hindermost plate
is 435 mm., a little more than seventeen inches. The height of the tenth plate, in a straight line, is 180 mm. . . .

On the hinder end of the tooth, on the grinding surface, six plates outcrop in a distanc^e of 100 mm.; but on the
side of the tooth, at one-half of the height, there are only three and one-half plates in 100 mm."
'[See footnote 2 on page 996 above regarding the Early and Middle Pleistocene age of A. imperator. — Editor.]
 )

THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1003

Nat. Mus. 6662 Dps (Afton, Okla.), length 135 mm., width 69 mm.; ridge-plates 7+; 5% or 6 ridge-plates in 100 mm.; enamel thick
(see Hay, op. cit., Pi. lxi, figs. 7, 8).

Nat. Mus. 6663 Dp'' or M' (Afton, Okla.), doubtfully referred. 12 estimated ridge-plates (Hay, op. cit., p. 414).

WiKonain lowan lUinoian lUnsM Pre4Visconsin

CD
Orifliut

Fig. 890. Distribution of .\rchidiskodon imperator west of THE Mi.ssissippi River. .After Hay, 1924, Map 10, p. 337,

AND EXPLAN.\TI0N OF MaP, P 336 (Texas omitted).

Nebraska. (6) Near type locality "Loup Fork of Platte River" [probably Kansas. Localities 1 to 3, Wellington, Ness City, Russell (Russell not
Seneca (fide Hay)), al.so eight other localities, Holmes- on map).
ville,Fairbury, Reynolds, Wauneta, Cody, Grayson,
Colorado. Localities 1 to 6, Laveta, Colorado Springs, Denver, Boulder,
Hebron, and Sutton. [Lugn and Scliultz (1934.1, p.
Fort Collins, Glenwood Springs.
373) give "Pawnee Loup Branch of Platte River =
Middle Loup, probably Hooker Co.," as the type Iowa. Localities 1 and 2, Mt. Pi.sgah and Mapleton.
locality. —Editor. Wyoming. (1) Powder River.
Oklahoma. (1) Afton, Ottawa County; type region of the Aftonian or
Montana. Localities 1 and 2, Helena and Valier.
Ist Interglacial stage, in which age A. imperator was
very abundant. Also three other localities, Kinlock, These grinding teeth are recorded as exhibiting the specific characters of

Okeene, and Kingfisher. Kingfisher not on map. .1 rchidislcodon imperator or of A . tiayi.

 1 4 nat. size

Cranium of a Young Male of ARCHiorsKODON imperator of Texas in the American Museum

I''ig. 891. Rilfncd young male skull of Archidiskodon imperator Leidy (Amer. Mus. Cojie Coll. 14475), discovered near Dallas, Texas. Compare figures
895, 896 (restored), also figure 805.

.\, Frontal aspect. One-eighth natural size. Al, Left lateral aspect. One-eighth natural .size. A2, Partly worn left M-. One-fourth natural size.

Tliis young male skull was described by Cope in 1889 (1889.2) as "Elephas pritnigenius coliunbi Fale." and figured by him in PI. xiv, also on p. 208, fig. 9.
It was erroneou.sly referred by Osborn (1922..5.55, p. \!i) to Elephas = Parelephas] jeffersonii. Both the jirofile and front views as well as the wire mid-.section
[

(Fig. 80.'), Chapter XV) of this cranium in<licate that it is more elo.sely related to Arcliidiskodoii than to Parelephns. The single molar tooth preserved, l.M",
enables us to determine this individual as probably belonging to A rchidiskoilon imperator.

Second left superior molar, I.M", length 140e

Ridge-plates estimated (? 12); expo.sed S+
Width at broadest ridge-plate 80
Ridge-plates in 10 em. 6
Tip of left tusk to occipital condyle l,400e
Alveolus of left tu.sk to occipital condyle 960e
Breadth acro.ss cranium, posterior portion of occiput 032
Breadth acro.ss jugals tilOe
Breadth between orbits 484
Breadth between temporal fenestrje 330

We observe that this Texas skull (Amer. Mus. Cope Coll. 14475) belongs to a young male mammoth in which the superior tusks are partly erupt<'d; it was
at first regarded as a female .skull and was so labeled, but the diameter of the tu.sks forbids such a reference. In vertical .section (Fig. 810) this cranium closely
corresponds with the two other crania in the Ami^ican Museum collection certainly referable to th(! s]«;cics Parilephas jefferxonii. The young grinding
tooth shown herewith (.-^2), a left .second superior molar, I.M^, does not .so clearly display the characters of /'. jefft-monii as to preclude the possibility
of tliis cranium belonging to a juvenile Archidiskodon imperator; con.sequently the reference is somewhat doubtful. The skull was found in the Texa.s
geographi<' region characteri.stic of A. imperator in early Pleistocene times, a region which was also traversed by the migrations of I'.jeffersottii in late Pleistocene
times.

1004
 :

THE MAMMONTIN^: ARCHIDISKOnON AND METARCHIDISKODON 1005

Geographic Distribution and Nomenclature (Hay, 1914, anterior worn off, length 228 mm., breadth 97 mm.
1923, 1924).— Hay (1923, 1924) has made interesting and important Gidley (letter, Oct. 27, 1928) adds that the specimen from
obser\'ations on the geographic distribution of the remains of Venice has an extremely deep jaw but otherwise is like .4. impera-
Elephas [
= Archidiskodon] imperator in the central region of the tor; that both A. = Parelephas] columbi and ^4. imperator occur
[

United States which are summarized on pages 434 and 435, map at Melbourne and at Vero. "I think, therefore, there can be no
and legend, 1923, and pages 336 and 337, map and legend, 1924 doubt that these two species were contemporaneous in Florida,
[
= Fig. 890 of the present Memoir]. and perhaps also in other localities of the southern and south-
Unfortunately (1) some of the specimens referred in Hay's western United States."
work (1914, 1923, 1924) to E. imperator may belong to the true Osborn, 1922-1928: This is a very important case of the ap-
Elephas \
= Parelephas] columbi and owing to the con-
vice versa, parent association of Archidi.skodon imperator ref. and of Parelephas
fusion which has existed in the minds American
of practically all columbi ref. in the same geologic locality, if not actually from the
writers up to July 8, 1922 (Osborn, 1922.555), when Osborn clearly same level.
distinguished the true E. columbi from the true E. imperator. (2)
In Hay's volume of October, 1924, the northerly mammoths ARCHIDISKODON IMPERATOR IN THE AMERICAN MUSEUM
previously (1914-1923) named by him "Elephas columbi," as COLLECTION
plotted on map 5, p. 327, and map 6, p. 329, are partly named by See figures 885, 886, 888, 889, 891, 892, 896-898, 900, 906-908, 912, 920
of the present Memoir.
him "Elephas boreus," a name preoccupied by Parelephas jeffersonii
Osborn; also, on maps 7 to 9, are partly named Elephas columbi. The type cast, the neotype, and the finest referred specimens
As above noted. Hay (1924, pp. 57-84, "Finds of Elephas columbi referable to Archidiskodon in the American Museum collection are
in the Middle Region of North America") fails to recognize the listed herewith (bottom of this page). These superb materials are
now well-determined typical ridge formula of Parelephas columbi, illustrated in various figures of the present Memoir and add
namely, I\I 3 riTe+- greatly to our knowledge of this imperial species of southern mam-
moth. They display clearly the specific characters enumerated
ARCHIDISKODON IMPERATOR IN THE U. S. NATIONAL MUSEUM above.
FLORIDA COLLECTION
Specimens collected for the U. S. National Museum by James COMPARISON OF THE JAW AND DENTITION WITH OTHER
W. Gidley in the locality of Melbourne, Florida, and presumably all PROBOSCIDEANS
from the "No. 2" bed of Sellards, are as follows: A great deal of study has been given to comparison of the jaws
Nat. Mus. 11805. A third inferior molar, M3, with 15-18 of various types of the Proboscidea, especially in correlation with
ridge-plates; laminar frequency 5 ridge-plates in (1)Abbreviation (brachycephaly) and elevation (hypsicephaly) of
10 cm. the cranium, also with (2) insertion of the third inferior molar
Nat. Mus. 11814. A much worn third inferior molar, M3; tooth which naturally is the chief functional organ of the body.
laminar frecjuency 5 ridge-plates in 10 cm. Total It is seen in the accompanying diagram (Figs. 892, 893) that the
ridge-plates 12+
(tooth worn to base). progressive jaws of Archidiskodon imperator (Fig. 892 A, B) are
Nat. Mus. 11620. The Venice mammoth from near Mel- profoundly different from the relatively elongate jaws represented
bourne, Florida. Aged individual, with third superi- in figures 893, A, B, C (Archidiskodon hayi, Parelephas washing-

or and inferior molars much worn, also jaw, por- tonii, Loiodonta africana). The jaws of Archidiskodon imperator
tions of skull, right tusk, and hind Third
foot. (Fig. 892 A, B) are also readily distinguished both from those of
superior molar, l.M', total ridge-plates estimated at Parelephas jeffersonii (Fig. 892 C, D) and of Elephas indicus (Fig.
18)2-19, 3 to 4 anterior worn off, length 200 mm., 893, D, E).
breadth 106 mm., 5}i ridge-plates in 10 cm.; maxi- It is observed that in the Texas jaw (Amer. Mus. 10598 —see
mum breadth 106 mm. as compared with 125 mm. Fig. 892 B) M3 exhibits sixteen ridge-plates, while in the Kansas
in Leidy's Nebraska type (Fig. 884). Third inferior —
jaw (Amer. Mus. 14558 see Fig. 892A) M3 exhibits five accessory
molar, r.Ms, estimated ridge-plates 16-17, 3 to 4 ridge-plates, namely, 16, 17, 18, 19, 20; this difference cannot be due

Amer. Mus. 2568 Type r.M^ of Elephas imperator, cast (Amer. Mus. 2568) after original in U. S. National Museum (Nat. Mus. 185).
United States See figures 884, 886, 887.
Amer. Mus. 11871 Neotype (Osborn) of Elephas imperator, right third superior molar, r.M^ from Guadalajara, Jalisco, Me.xico.
Mexico See figures 886 Al, 888.
Amer. Mus. 14476 Lower portion of cranium with tusks, here represented in figures 889B, 896, 897, 906. From Victoria, Victoria
Texas Gounty, Texas.
Amer. Mus. 14558 Imperfect lower jaw, right and left third inferior molars. Crown view (Fig. 889A); side view (Figs. 892A, Al).
Kansas Total of 19-20 ridge-plates, of which the 19th and 20th are extremely short and rudimentary (see Fig. 892).
From Ness County, Kansas.
Amer. Mus. 10598 Lower jaw with partly worn M3 (see Figs. 892, 898), also right forelimb (see Figs. 906, 907, 908, 912). From near
Texas Tule Cafion, Briscoe County, Texas.
Amer. Mus. Gope Referred young male skull of Archidiskodon imperator, with tusks and l.M'-, discovered near Dallas, Texas, and
Goll. 14475 described by Cope in 1889 (1889.2) as "Elephas primigenius columbi Falc." (see Fig. 891).
Texas
Elepnas jetlersonn
Amer Mus 13225

Jaws of Archidiskodon imperator in comparison with Parelephas je FFERSONII, P. WASHINOTONII, LoXODONTA AFRICANA, AND ElEPHAS INDICUS
Fig. 892. and M3 of Elephns =Parele-
Internal aspect of the lower jaw.s [ Fig. 893. Internal views of jaws of Elephns indicus, Loxodonta africana,
phas\ jfffersonii (paratype and type) and Elephns = Archvliskodojt] imperator,
[ Elephas [= Parelephas] washinglonii, and E. = Archidiskodon] hnyi, for com-
\

al.so sections Dl, Cl, Bl, Al, cut at point indicated by the dotted line S. parison with figure 892. One-eighth natural size.
( )iie-eighth natural size. E, Elephns indicus, a fully developed M3 of the right side with twenty-
D, Elephtts [
= Parelephas] jeffersonii ])aratype (Amer. Mus. 13225); M3 seven plates. Drawn after de Blainville, 1839 1804, PI. ix, fig. ti'' (reversed).
in situ with fifteen worn plates and eight unworn plates, a total of twenty-three D, Elephas indicus, right jaw (Amer. Mus. Dept. Mam. .54201); M3 with
plates. [See Fig. 960 for final count of twenty-four ridge-platcs in this |)ara- nineteen plates fully formed and five imperfect plates in the jaw, a total of
type specimen.] twenty-four plates.
C, Type of Elephas [
= Parelephas] jeffersonii (Amer. Mus. 99.J0), a very C, Loxodonta africana jaw (Amer. Mus, Dept. Mam. 39083, Akeley Coll.);
aged individual, M3 with seventeen worn jilates and three additional unworn ?M2 with twelve plates developed.
plates in situ, a total of twenty plates. This, as exi)lained elsewhere, is a B, Type jaw of Elephas = Parelephas] washingtonii Osborn (Amer. Mus.
(

very aged individual and it is probable that one or two of the cxtreine an- S081A), with twenty-one plates developed in M^. Compare figure 909, type of
terior i)lates have been worn off. Parelephas progressus.
B, Jaw [= Archidiskodim] imperator ref. (.\iner. Mus. l()r)98),
of Elephas A, Type jaw of Elephas [
= A rchidiskodon] haiji Barbour (Neb. Mus.
M3 with fourteen worn plates and a total of sixteen plates in situ; a very 23 0-14)."
robust individual. From Tule Canon, Texas.
A, Elephas = Archidiskodon] imperator ref. (Amer. Mus. 14558), from
[

Ness County, Kansas, M3 with fourteen to fifteen (est.) worn i)lates and five
additional, a total of twenty plates. See also figure 889A, crown view of same
tooth.

1000
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1007

to age or to sex; more probably due to the progressive addi-

it is Incisive Tusks. —The characters of the tusks are superbly
tion of ridge-plates in theKansas specimen. Majestic as are these disi)layed in the cranium (Amer. Mus. 14476) as shown in figure
imperial mammoths from Texas the largest pair of tusks known 896; these tusks curve downwards, outwards, upwards, inwards,
is from Mexico and is described below, with change of name, by until finally they cross each other on the median line, forming
Osborn (Fig. 894).' a complete superior arch, as in the aged type specimen of Elephas
Apposition of the Grinders. In the American Museum — [
= Parelephas] jeffersonii. The tusks are parallel but not closely
collection there is no case of skull and jaws being found together, opposed to each other where they issue from the alveoli, as in the
so that we cannot positively determine the apposition of the specimens referred to this species in the Nebraska State Museum
grinders it is almost certain that an equal number of ridges are in
; (Fig. 899) found at Hay Springs, Sheridan County, Nebraska.
use at the same time in animals of the same age. Consequently This comparative study of the superior and inferior teeth and
the superior and inferior molars represented in figure 889, in each of jaws enables us to define the species A. imperator much more
which fifteen ridge-plates are in use above and below, probably closely than before.
represent animals of the same age, although they may not be ex-
actly in the same stage of dental progression. The teeth from Record Proboscidean Tusk
Kansas (Amer. Mus. 14558— Fig. 889A, 892A) have 14-15 ridge- A record proboscidean tusk (Amer. Mus. 22481) was presented
plates in use, also the maxillary teeth from Texas (Amer. Mus. to theAmerican Museum in 1934 by Mr. (Jeorge D. Doughty of
—
14476 see Fig. 889B) have fifteen ridge-plates in use. These Post, Texas. This tusk was found in the vicinity of Mr. Doughty's
grinders (Fig. 889) are examples of typical A. imperator molars home, namely, Post, Garza County, Texas, in which region the
exhibiting the following characters: (1) Inferior ridge-plates con- Imperial Mammoth (Archidiskodon imperator) seems to have
cave anteriorly, partly worn summits of ridge-plates exhibiting flourished in Pleistocene time. This gift was first noted in the
January, 1935, number of Natural History, page 84, followed in
the April number, page 357, by a brief description with compara-
tive estimated measurements. Since that time the tusk has been
restored conservatively to conform to the measurements given by
Mr. Doughty, who was unable to save the anterior end; in life it

Fig. 894. A right superior tusk of

record size (16+ ft.) of Archidiskodon
imperator (Amer. Mus. 22481), from
Post, Texas, one-twentieth natural size,
in comparison with the superb left tu.sk

(13 9Kft. in.) formerly in the National

Museum, Mexico, after photograph of

original taken by Barnum Brown in

1910. A letter from Sr. Francesco Con-
treras of the Universidad Nacional de
Mexico, May, 1935, states that the orig-
inal tusk w.is unfortunately destroyed.

-i'mf»m>'-»«^-

7-8 conelets; (2) superior molars, \M\ ap. 290 mm. (restored),
tr. Ill mm., index 38; r.M', ap. 295 mm. (restored), tr. 120 mm.,
index 41 M', ridge-plates concave
; (3) posteriorly, thus following
the mechanical principle of reversal; (4) breadth-length index,
M' = 41.
 Skull of ArchidUkodon imperator (Amer. M.is. 14476), found at Victoria,
Texas. The upper portion of tins skull is entire-
Fig 890
grinding teeth are perfectly preserved. The palate and grinding teeth of
ly restored. The prema.xillaries, the palate, and the .superior
nat,ural, without restorat,ion; they measure 4 m.
thisspecimen are also represented in figures 897 and 889. The tusks are complete and
m. 20 cm., or 13 the measunnK'nt of the tusks of A. imperator in the Geologi-
21 5 cm. or 13 ft. 10 in., as compared with 4 est., ft. 9>i in.,

cal Institute of the City of Mexico (Fig. 894), a.s remeasured by

Barnum Brown. [See footnote on preceding page which give.s the
length of the record tusk of this species (Amer. Mus. 22481) as 16+ ft.— Editor.)
1008
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1009

was evidently longer than now restored, namely,

16+ feet. Associated with this giant tusk were
two superior molars (M-), probably belonging to
the same individual.

CRANIAL CHARACTERS OF ARCHIDISKODON

Compare Chapter XV, pp. 915-926, also figurrs 86.5,

891, 895, 896, 897, 902, and 906 of the present Memoir.
In the progress from youth to maturity, the
occipital summit and profile of the Archidiskodon
cranium changes profoundly with growth, with sex,
and with the development of the enormous' superior
tusks; these progressive stages are dis]3layed in a
comparison of the following eight more or less com-
plete crania in the Los Angeles, American, and
Nebraska State museums, and the Geological In-
stitute of Mexico:
Fig. 895. California. Primitive stage. Young
male(?) skull from Elephant Pit No. 9, Rancho La
Brea tar pool (Los Angeles Mus. Nos. 3800-1 (skull)
and 3801-1 (jaws). = Archidiskodon imperator.]
[

Fig. 891. Texas. Young male stage. Skull

from near Dallas (Amer. Mus. 14475).
[= Archidiskodon imperator.]
Fig. 902. Mexico. Adult male stage. Skull
from Tepexpan (Geol. Inst. 212).
[
= Archidiskodon imperator.]
Fig. 897. Texas. Aged stage. Skull (summit
restored) from Victoria, Victoria County (Amer.
Mus. 14476). [
= Archidiskodon imperator.]
Fig. 918. Nebraska. 'Lincoln County Mam-
moth.' Type mandible, and tusks from near
skull,
Curtis, Lincoln County (Neb. Mus. 5-9-22).
[
= Archidiskodon imperator maibeni.]
Nebraska. 'Adams County Mam-
moth.' Not figured in present Memoir.
Skull, mandible, and teeth from sandpit 6
miles due south of Hastings (Neb. Mus.
11-3-13). [
= Archidiskodon imperator.]
Nebraska. 'Howard County Mam-
moth.' Skull, mandible, teeth, and tusks,
from near Dannebrog (Neb. Mus. 2-7-17B).
= Archidiskodon imperator.]
[

Nebraska. 'Custer County Mammoth.'

and teeth from Callaway (Neb.
Skull, tusk,
Mus. 16-6-16). = Archidiskodon imperator.]
[

Texas, Victoria, Skull (Amer. Mus.

14476— Figs. 896, 897, 906).— Only the
lower portion of the cranium, including the
palate, condyles, and paired grinding teeth,
is preserved, the upper portion, as shown
in figure 896, being entirely restored
above the white dotted line; as the summit
Fig. 897. Skull of Archidiskodon imfjcralnr from Texa.s, as largely restored and mounted in llie
of the Victoria, Texas, skull (Amer. Mus.
.\merioan Museum.
14476) is restored, we have not ventured to
upper part of cranium
Skull (Amer. Mus. 14476) found near Victoria, Victoria County, Texas;
give this hypsicephalic character full expres-
entirely restored. Side view one twenty-fourth natural size; palatal view one-sixth natural size.
sion; it characterizes all the young speci- See also figure 889 for view of palate; figure 896 for front view of same skull.
1010 OSBORN: THE PROBOSCIDEA

mens of the species /I. imperator; but an aged Mexican skull (Fig. closely analogous if not genetically related to those of the true
902) has a broad, massive occiput. mammoth (Mannnonleun primigenius), while they arc widely
The hypsicephaly or peaked contour of the superior crest is distinct from those of the trogontherian elephant {Parelephas
based upon a beautiful cranium of a young male (?) in the Los trogontherii) or from the typical elephant (Elephns indirus).
Angeles Museum, from Rancho La Brea (Figs. 865, No. 10, and Comparing the profile and palate of this cranium with thai of
895); we observe a remarkable similarity between the ])rofile Archidiskodon planifrons and of A. meridionalis, we observe the
of this Los Angeles cranium and that of the true Mammonleus close similarity in fore-and-aft compression (cyrtocephaly) and
primigenius, namely: (1) Forehead concave; (2) parieto-occipital corresponding vertical elevation (hypsicephaly, acrocephaly).
union acute, hypsicephalic ; (3) occipital condyles and orbits
approximate, i.e., brachycranial (4) depth extreme from peak
;
FLATTENED CRANIA OF ARCHIDISKODON IMPERATOR FROM
of cranium to base of lower jaw. Reverting to figure 865, these HAY SPRINGS, NEBRASKA, IN THE NEBRASKA AND
distinctive cranial characters of Archidiskodon imperator are AMERICAN MUSEUMS. AFTONIAN AGE.
Asprovisionally determined by Matthew (1902.1, pp. 317,
318, and 1918.1, pp. 226, 227) from the American Museum col-
Amer. Mus. 10598 Ref. lections of 1893 and 1897 (modified by Hay and by Osborn), also
by Frick (1929.1, p. 107, and 1930.1, p. 79), [and finally by Barbour
and Schultz who have published a preliminary list of the Hay
Springs fauna (1937.1, pp. 3-6)], the following includes the species
so far recordedby the above-mentioned authors:

Edentata
Mylodon garmani Allen
Mylodon nebrnscensis (Brown)
Megalonyx leidyi Lindahl
RODENTIA
Cynomys niobrarius Hay
Geomys sp.
Thomamys sp.
Castoroides ohioensia nebraskennis Barbour
Castor sp.
Ondatra nebrascensis (Hollister)
Microtus?

Carnivora
Canis latrans? Say
Canis (Aenocyon) dirus nebrascensis Frick
Arctodus simus nebrascensis Frick
Mustela vison? Schreber
Smilodon nebrascensis Matthew
PnOBOSCIDEA
Archidiskodon imperator (Leidy)

Perissodactvla
Equus excelsus Leidy
Equus excelsus niobrarensis Hay
Equus calabatiis nebrascensis Frick
Artiodactyla
1/B nat. size
Leidy
Platygoiitis vrtus
Camelops kansnnus Leidy
Fig. 898. Jaw of Archidiskodon imperator rof. (Amor. Mils. 10.598), found Camelops vitakerianus? (Cope)
at TiilcCanon, Texas, by the American Museum Exi)edition of 1899. One- Tanupolama amcricanus (Wortman)
eighth natural size. Section and inside view of the same jaw, completely
Odocoileus sheridanus Frick
exposing M3, shown in figure 892B, Bl.
\, Higlit rsunus, outer aspect jaw uptilted. Capromeryx furcifer Matthew
;

k\, Top view; tooth gn-atly shortened by perspective. Telrnmcryx (Ilayoccros) falkcnbarhi l*'rick

A2, Same tooth in perpendicular view of crown, sliowing sixteeii ridge- Bovid
platcs, fourteen of which arc worn, indicating that tliis was a fully adult
animal. It is a striking fact that two extremely flattened crania have
See figure 907 for forelimb of same individual. been discovered in the deposits of Hay Springs, Sheridan County,
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1011

Nebraska, which are considered of Lower Pleistocene (Aftonian) exposed ridge-plates, also three on the deeply worn anterior
age. The first is the flattened cranium (Fig. 899) in thc^ Nebraska portion of the tooth, thirteen worn ridge-plates in all; the total
Museum; the second is the flat cranium (Fig. 900) in the Ameri- number of ridge-plates unknown. The tusks are very massive in
is

can Museum of Natural History. The flattening of these crania transverse section and closely approximated where they issue from
is a proof of the heavy geologic pressure to which these beds were the alveoli; when found the left tusk was 9 feet long, but it was
formerly subjected. partly destroyed by careless collecting.
Nebraska Museum. — In figure 899 is represented the palate American Museum Crushed Skull (Fig. 900). — This
by Prof. E. H. Barbour to Elephas imperator
of a flat skull referred
(Neb. Mus. 1-11-8-17E); it was found in 1917 at Hay Springs,
Sheridan County, Nebraska, in a bed of diatomaceous earth,
flattened to four inches in thickness. The broad-plated grinding
teeth enable us to confirm this reference, the molars presenting
marked resemblance to those of the typical Archidiskodon im-
perator. The geologic age of this specimen is very important,
since the Hay Springs fauna is now regarded as of Lower Pleistocene
(Aftonian) age.' In describing this skull, Barbour observed ten

Fig. 899. ArckiiliskodoH imperator cranium of aged male (Neb. Mus. Fig. 900. Archidiskodon imperator cranium of adult male (,\mer. Mus.
1-11-8-17E) foimd a bed of dJatomaceou.s earth, flattened to four inches in
in 17355), extremely flattened, exhibiting small posterior nares and cranial
thickness; discovered in 1917 at Hay Springs, Sheridan County, Nebraska. foramina. Discovered in 1916 at Hay Springs, Sheridan County, Nebraska.
'[Middle Pleistocene (see Barbour and Schultz, 1937.1, p. 3).— Editor.]
 \ —

1012 OSBORN: THE PROBOSCIDEA

crushed cranivim (Ainer. Mus. 17355) closely resembles in size and 5-9-22 Skull, mandible, tusk, forelimbs, six cervicals,
character that in the Nebraska Museum (Neb. Mus. 1-11-8-17E). several dorsals, four lumbars, sacrum, femur,
It was discovered in 1916 by Albert Thomson in the famous Hay Lincoln Co., Nebraska.
part of pelvis, ribs, etc.
Springs quarry, Sheridan (
'ounty, Nebra-ska.The cranium belongs Described as Elephas {Archidiskodon) maibeni.
to a fully adult male, somewhat less aged than the Nebraska Lincoln County Mammoth.
Museum cranium (Fig. 899), because the third superior molars, [
= Archidiskodon imperalor 7naibeni.]
r.M', I.M', are less worn, displaying anterior ridge-plates. Besides 1-4-26 Skull, teeth, and tusksform apparently identi-
of a
the anomalous crushing, this specimen finely displays the char- cal with A. maibeni; the same type of teeth and

acters of the palate, especially the 'backward' posterior nares, the same size and curvature of tusk (see Barbour,
which contrii-sts with the very large 'forward' posterior nares of 1926.1, 1). 122). From Lingle, Wyoming.
the superb Parelephas jeffersonii cranium (the Franklin County 1
= Archidiskodon imperalor maibeni.]
Mammoth) in the Nebraska Museum (Neb. Mus. 1-4-15 see Fig. — [1-11-8-17E Palate of a flat skull referred by Barbour to
963), which for a time was erroneously referred to E. [
= Parelephas] Elephas imperalor, Hay Springs, Sheridan Coun-
rnlitmbi. This palate of Archidiskodon imperalor (Amer. Mus. ty, Nebraska. Sheridan County Mammoth.
17355 — Fig. 900) should be compared with that of Parelephas [
= Archidiskodon imperalor.]
jc_lfersonii, which more nearly resembles that of Elephas indicus
Mandibles
bengalensis (Fig. 800). The backwardly placed opening of the
19-9-17 Mandible, left half, with teeth. Powell, Jefferson
posterior nares appears to be a very distinctive character of
Co., Nebraska. [
= Archidiskodon imperalor.]
Archidiskodon.
29-25-11-18 Mandible and teeth. Inland, Clay Co.,
HaySprings, Nebuaska, Fhick Collection. The follow- — Nebraska. [
= Archidiskodon imperalcrr.]
ing specimens, collected under the direction of Mr. Childs Frick,
2K-3-8-19 Mandible and teeth. Republican City, Harlan
serve to confirm Archidiskodon imperalor as the characteristic
Co., Nebraska. = Archidiskodon imperalor.]
[

proboscidean of the Hay Springs horizon.

8-7-08 Mandible and teeth. Young. Benkleman, Dundy
Amer. Mus. numbers: 25501 A, fragment of superior molar,
Co., Nebraska. [
= Archidiskodon imperalor.]
riflgc-iilate maximum height 244 mm.; 25501, posterior jiortion
18-2-22 Mandible with teeth. In Aftonian gravels. Staple-
of inferior molar, ridge-plate maximum height 164 mm.; 25501B,
hurst, Seward Co., Nebraska. Type of Elephas
anterior portion of extremely worn superior molar, maximum
scotli Barbour, 1925. [
= Archidiskodon im-
width 111 mm. 25500A, middle portion of an aged inferior molar,
;

peralor scotli, or juvenile A. imperalor.]

width 97 mm., laminar frequency 6K in 10 cm. 25500, portion of ;

23-6-14 Mandible with teeth. From Crete, Saline Co.,

half-worn superior molar, width 107 mm., laminar frequency 6 in
Nebraska. Type of Elephas hayi Barbour, 1915.
10 cm.; 25505 A-D, portions of inferior milk molars; 25506, por-
[
= Archidiskodon harji.]
tion of first milk molar.
Grinding Teeth
CRANIAL MATERIAL IN THE NEBRASKA STATE MUSEUM Largely as plotted by Hay (1924) under Elephas [
= Archi-
The finest is in the Nebraska State
Archidiskodon collection diskodon] imperalor, thirty-two or more upper and lower grinding
Museum by Director Barbour (1925.3, pp. 117,
as recently listed teeth, usually associated, from the following counties in Nebraska,
118) in connection with his description of the "Columbian Mam- Kansas, Indiana, and Wyoming:
moth Elephas Maibeni." This list is entitled "Columbi Material Dawes, Furnas, Jefferson, Platte, Richardson, (lagc, Cass,
in the State Museum"; as revised by Barbour to July, 1925, it is Thayer, Howard, Adams, Fillmore, Buffalo, Cheyenne,
as follows: Valley, Harrison, Clay, Butler of Nebraska; also —
Skulls — —
Herndon of Kansas, Tipton of Indiana, and (loshen
11-3-13 Skull, mandible, and teeth from sandpit 6 miles of Wyoming.
due south of Hastings, Adams ('o., Nebraska.
Neb. Mus. 1-4-26 Archidiskodon imperalor maibeni ref. of
Adams County Mammoth.
Lingle, (Joslien County, Wyoming.
[
= Archidiskodon imperalor.]
Walls of cranium \ery sloping; maxi-
1-4-15 Skull, mandible, and tusks complete, of extraor-
mum transverse measurement of pal-
dinary size. Campbell, Franklin Co., Nebraska.
ate 30 in., breadth of nasal opening
Franklin County Mammoth (compare Figs.
20 in.; circumference of tusk 25 in.;
963 and 964 of present Memoir).
ridge-plates as thick as those in A.
[
= Parelephas jeffersonii.]
imperalor.
16-6-16 Skull, tusk, teeth, and femur. Callaway, Custer Co.,
Nebraska. Cluster (Bounty Mammoth. 5-11-20 Arrhidi.skodon imperalor ref., from Bell-

[
= .4 rchidiskodon intpcrator. wood, Butler County, Nebraska (cast
2-7-17B Skull, mandible, teeth, tusks, two tibise, scapula, Amer. Mus. 20069). Posterior half
and ribs. Found 7 miles south of I''arwell, near of a third right inferior molar, r.Ms,
Dannebrog, Howard Co., Nebraska. Howard with eight broad ridgc-i)lates; six

County Mammoth. = ^4 rchidiskodon imperalor.]

[ arched ridge-plates in 20 cm. as com-
 .

THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1013

pared with nine arched ridge-plates in form, Elephas imperator. In the collection of the Geological Survey
20 cm. in the corresponding tooth of of Mexico in the new survey building are the skull and tusks of an
Parelephascolumbi (Amer. Mus. 13707, E. imperator of magnificent proportions, the tusks measuring 5 m.
I.M3); breadth 4)-^ in. = 112 mm. This 10 cm., or 16 feet 10 inches [4 m. 20 cm. est., or 13ft. 9% in., as re-

is the most broadly ridge-plated molar measured by Barnum Brown] in length ; this specimen was secured
on record. during the excavations for the great drainage canal of the Mexican
4-12-13 Portions of inferior molars and femur. Valley."
Arched laminiE of inferior molar, four In the collection of the Geological Institute of Mexico, under
in 10 cm., as compared with three the direction of Dr. Jose G. Aguilera, there are also several single
arched laminse in 10 cm. (Neb. Mus. teeth of Archidiskodon imperator, molars of the Parelephas columbi
5-11-20). Femur 56 in. long or 1422 type from the \'illage of Zacapii in Michoacan and of A imperator .

mm.; same measurement as that of from the valley of Puebla (cf. Osborn, op. cit., p. 931).
Neb. Mus. 13-24-10-14 (see Fig. 908). —
Reyes, 1923. The specimens described and figured (1923) by
_ Seiiorita Reyes, are as follows:

Elephas imperator
Escuela de Ingenieros No. 1. Jaw containing
M 3, with 6 ridge-plates e.xposed. From Tequix-
quiac. Figs. 1 and 2, p. 228.

Geol. Inst. 207. Palate with r.M^ 13 ridge-

plates exposed; length 220 mm., breadth 82 mm.
From Zumjjango. Fig. 3, p. 229 (see Fig. 901 of
the present Memoir).
Geol. Inst. 210. R.M», length 300 mm.,
breadth 115 mm., 12)i: ridge-plates in 25 cm. From
Tepexpan. Fig. 4, p. 230.

Geol. Inst. 212. Cranium of young [adult]

individual, a beautiful specimen well conserved.
It is without tusks. Enormous prominence of
occipital crest. Associated lower jaw. Dimen.sions
of second superior molar, M- length of worn
: area
168 mm., breadth 108 mm. From Tepexpan. Fig.

7, p. 233, Fig. 8, p. 234, and Fig. 9, p. 235 (see

Fig. 902 of the present Memoir).
Posterior view (Fig. 7).
Anterior view (Fig. 8), forehead concave.
Summit of occipital crest broadly rugose.
Fig. 901. Referred superior molar, r.M', of Archidiskodon imperator in the Geological Crown view of lower jaw (Fig. 9).
Institute of the City of Mexico (No. 207). After photograph kindly furnished by Senorita
Reyes. One-half natural size. Compare Reyes, 1923, fig. 3. This tooth was described Geol. Inst. 211. Left ramus of jaw' with I.M3,

by Senorita Reyes (1923, p. 229) as follows: "Fig. 3. Escala: Vt. Ejemplar num. 207 del I. 12;^ ridge-plates in 25 cm., length 220 mm., breadth
Geologico. Elephas imperator, Leidy. Molar derecho." Locality: Zumpango, Mexico. 98 mm.; length of femur 1360 mm. From Tepex-
Observe that there are b% ridge-plates in 10 cm. as compared with 9}i in P. colurnhi pan. Fig. 5, p. 231, Fig. 6, p. 232 (see Fig. 903 of
felicis. Length of worn surface 220 mm., breadth 82 mm.
the present Memoir)
Archidiskodon (?)hayi Ref. in Mexico. A mandible' (Fig. —
ARCHIDISKODON IMPERATOR IN THE MUSEUMS 903) from Mexico (Geol. Inst, v-211), referred by Seiiorita Reyes
OF THE CITY OF MEXICO to Elephas hayi, supports the evidence afforded by Barbour's type
See Parelephas colurnhi below (Chap. XVII) of E. hayi that an Upper Pliocene stage of Archidiskodon, apjjarent-
We owe Osborn (1905.270), to Freudenberg (1922), and
to ly similar in progression to the 4. planifrons of Asia and of southern
finally and most
fully to Senorita Reyes (1923) descriptions of the France, entered North America.
superb materials referable to Archidiskodon imperator and to Osborn, 1924: This mandible from Mexico is intermediate
Parelephas colurnhi in the museums of the City of Mexico. between the E. hayi type of Barbour and the E. imperator type of
Progressive Stages (Mexico). —
As described by Osborn Leidy; it shows a prolonged rostrum (see Fig. 903) ; it appears to
(1905.270, p. 931): "The elephant remains in the National Mu- us like a progressively modified rostrum derived from an ancestor
seum have usually been ascribed to Elephas columbi; but they in- with a jaw like that of .4 . planifrons. While this mandible has been
clude molar teeth not only of this species, but of the much larger referred to Elephas hayi by Seiiorita Reyes, it seems to Osborn to

'Osborn, 1929: .\ similar mandible is described below (p. 1033) as Archidiskodon sonoriensis, to which species tliis specimen may also be referable.
1014 OSBORN: THE I'ROBOSCIDEA

Ckanium ok an Adult Male ok Auchidiskodon impehatou is the Geological Museum ok Mexico

rig. 902. Keferrei mature cranium of Archiili.'ikoihii impcrator in the Geological Institute of the City of Mexico (Geol. Inst. 212), reduced to one-
twelfth natural size. After photograph kindly fiu-nished the i)resent author by Scnorita Reyes Locality: Tepexpan, Mexico. Compare Reyes, 1923, figs.
7, 8, and 9.

This robust male cranium lacks the sharply peaked, acrocephalic, occipital crest structure seen in the young male(?) cranium from Rancho La Brca (Fig.
89.>). Judging by the elongated and massive alveoli and rugose exoccipital muscular attachments this cranium supported an enormnus pair of tusks. The
adult male tusks are estimated at 13H 16+ feet in length (see Fig. 894)

represent rather an intermediate and distinct sj^ecies in which there

is a much larger number of ridge-plates than in the t yi)e of E. hniji.
It another proof of how much we have still to learn regarding
is

the characters and migrations of Arrhidhhodon in North America.

Doubtless future excavation will re\eal additional material
of great value. We turn to Frcudenberg (1922) for his views
regarding the relationshi]) of the true Elcphas coliiinbi Falconer to
the true E. imperator of Leidy (see j). 1017 below); also for his
remarks on the relationship of A. impcrator to the A. meridimiaUs
ofEurope and to the A. planifrons of Asia and southern France.
AnCESTKV of ElKPHAS IMPKHATOll (FnKUDIONBEKG, 1922,
SoERGEL, 1915').— (Freudenberg, 1922, p. 171) "Die Mastodonten :

wanderten fri'iher nach Amerika iiber, die Elefanten s])ater. Die

innerasiatischcn Hochliinder, die den meisten Saugetierstiimmen
den Ursprung gaben, sind auch liier in diesem Fall als Ileimat der

mexikanischen Arten anzusehen. [I'Dotnote: 'V'gl. W. D. Matthew,

(
'limatc and I'A-olution. Annals of th(! New York Acad, of Science.
\'ol. 24. pag. 171-318.']. Fiir die Elefanten gilt das mit ziemlicher
Sicherheit. Eine Ableitung des El. iinprraior von El. nicridioruilis
Euroi^is, wie Soergel |l'"ootnote: '\V. Soergei, Die Stammesge- Fig. 903. Mandible of Elc/ikax [
= ArchUIUkodon] hayi{l) ref. in the collec-
schichfe der I'^lefanten. I\'. Die anierikanischen Elefanten. tion of the Geological Institute of the City of Photograph
Mexico (v-211).
Cenfralhl. f. Mineralog., Oeolog. u. Paliiontolog. 1915. No. 9. through the courtesy of Scnorita Reyes (conipaie Reyes, 1923, figs. and (i). .">

pag. 27S 2S:}.'| (las will, ist gcsiicht. El. plnii/frons isl weit eher About ono.seventli natural size. The prolongation of the .symphy.seal rostrum
suggests a remote resemblance to the type of Arrliiiliskniliiii hiii/i Barbour
der Stammvater alter spiilerer Eiefaiil idcii als der /s7. in<riili(iii(ilis,
(l'"ig. 913 of th(; present Memoir); this feature, however, is more character-
'Sec citation below, page 101."). istic of A. sonoriensis. [See footnote on p. 1013. —
Editor.)
 —

THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1015

der auch sicher niit Unrecht als der Ahne des EI. antiquus von Second [third] superior molar of the left side, M- [l.M'']. Pal. Coll.
gcnanntem Autor angesprochen wird." Univ. Leipzig 4402. Horizon and Locality. Ejutla, —
Ancestry of Elephas imperator, Soergel, 1915, p. 281. Estado de Oaxaca, Mexico. —
Type Figure. (Op. cit., p. 146,
"Wir iniissen annehmen, dass die nach Anierika iiberwandernden fig. 19): "Fig. 19. a. El. Columbi va,r. silvestris. Kronenansicht
Fornien der Meridionalis-Trogontherii-Reihe schon beim Uber- des zweiten oberen Molaren, in % nat. Gr. b. Seitenan,sicht in
wandern resp. kurz vorher besondere Charaktere gegeniiber dem etwa Js nat. Grosse. Original in Leii)zig, von Ejutla, Estado de
europaischen El. trogontherii merid7onali.s ausgebildet hatten, Oaxaca in Mexiko. Coll. Felix, wohl aus jiingerom Diluvium.
Charaktere, die in der weiteren Entwicklung sich zum 'Inipemtor- Subtropische Waldform."
Typus' steigerten. Es war also im altesten Diluvium im Kreis der
kontinentalen Elephantenformen eine Variationsbreite mit 2 Polen,
El. trogontherii ineridionalis im Weston und dem tlirekten Vor-
fahren des El. imperator im Osten \orhanden. Es ist klar, dass
als direkter Vorfahr beider Pole die nachstJiltere Mutation in der
kontinentalen Reihe zu gelten hat, das ist El. meridionalis des
altesten Diluviums, vielleicht auch des Oberpliociin. Als Vorfahr
des El. imperator Leidy hat also jedenfalls EI. meridionalis Nesti
zu gelten."
In our judgment Freudenberg was less fortunate in his treat-
ment of EIepha.s imperator as a subspecies of E. rolumbi. For
reasons originally stated by Osborn (1922.555) the type of Elephas
columbi Falc. is readily distinguishable from the type of Elephas
imperator Leidy; consequently the treatment of E. imperator as
a subspecies of E. rolumbi is invalid; it should stand E. = Archi- [

diskodon] imperator.
Synonyms of Archidiskodon imperator. — The four sub-
species of Elephas rolumbi proposed
by Freudenberg (1922) in this
manner are redetermined by Osborn as follows:

El. Columbi var. Felieis Freudenberg, 1922, p. 147, Taf. xvi

(viii), fig. 4, Tecamachalco, Puebla, Mexico
= Parelephas rolumbi felieis.
El.Columbi var. silvestris Freudenberg, 1922, p. 152, fig. 19,
Ejutla, Oaxaca, Mexico =Arrhidi.'<kndon imperator silve.'ttris.
EI. Columbi var. Fakoneri Freudenberg, 1922, p. 153, fig. 21,
Tequixquiac, Mexico = Archidiskodon imperator fakoneri.
El. Columbi var. imperator Freudenberg, 1922, pp. 160-171 B
= Archidiskodon imperator.
The subspecific value of Archidiskodon imperator silvestris
and of .4. imperator falroneri remains to be determined. We may
here quote in full Freudenberg's description of these subspecies and
reproduce his type figures. 1'reudenberg's Type of Archidiskodon imperator silvestris
Fig. 904.Type, left M', of Eleplias Columbi var. silvcslris Freudenberg,
1922, I). 146, reduced to one-third natural size. From Ejutla, Estado de
fig. 19,
Archidiskodon imperator silvestris Freudenberg, 1922 Oaxaca, Mexico. After photograjjlis sent by Doctor Freudenberg. Original in
Figure 904 Leipzig (Pal. Coll. Univ. Leipzig, 4402), Coll. Felix. B, Crown view; A, side

Ejutla, vState of Oaxaca, Mexico. view.

Probably Lo\ver(?) Pleistocene.
The type of this subspecies, related by Freudenberg to the
species Elephas [
= Parelepha.s] columbi, is obviously referable as Type Description. — (Op. ril., p. 152): "3. Dor Zahn von
a synonym or subspecies of Archidiskodon imperator. The type Ejutla {El. Columbi var. silvestris wir uns
Freudenberg). Wenden
(Fig. 904), with at least 16 ridge-plates, is too elevated to be re- genauer dem oberen Molaren von Ejutla (Estado de (_)axaca) zu,
garded as an M-; it appears rather to be a left M'; the six ridge- welchen Felix als 'El. Columbi' etikettiert und nur mit ^'orbe-
plates exposed (Fig. 904 B) resemble those of A. imperator rather halt wegen der relativ enggestellten Lamellen unter El. primi-
than of P. columbi; this Ejutla type has the very broad outer genius Blumenbach var. angefiihrt hat. Die.ser Zahn triigt jctzt
coating of cement characteristic of A. imperator and lacking in P. die No. 4402 der jialiiontologischen Sammlung der UniversitJit
columbi. Leipzig. Es ist cin zweiter obercr Molar der linken Seite. Ich
El. Columbi var. silve.slri.'i "Die Siiugetier-
Freudenberg, 1922. bilde ihn ab von der Seite als Textfig. 19b und a und von der Kau-
fauna des PliocJins und Postpliociins von Mexiko." (!eol. und Hiiche aus. Das Hinterende weist eine merkliche Einwiirtsdrehung
Palaeont. Abhand., N. F., Band XIV, Heft 3, pp. 152, 153. Type. iler Lamellen auf und zugleich eine \'erticfung in 5 cm Entfernung
 :

1016 OSBORN: THE PROBOSCIDEA

unter der letzten angekauten Lamelle. Diese Grube halte ich Soergelschen Nomenklatur) ist median lamellar, lateral anular.

trotz ihrer Rauhigkeit fur eine Pressionsmarke, hervorgebracht Besonders an den drei ersten Jochen ist dieser Bau der Schmelz-
voin nachdrangenden M^ Die vordersten 4 Lamellen sind abge- pfeiler zu beobachten. Ein mir vorliegender M'' von El. trogon-
brochen; einige davon diirften abgekaut sein. Dadurch ist die therii Pohlig [Footnote: '1) Abgebildet in meiner Arbeit: Die
Lamellenzahl im Minimum = 16." The author rightly contrasts von Mitteleuropa etc. Geolog. u.
Siiugetiere des iilteren Quartiirs

this species as identical in the ridgeformula of M' with that of the palaont. Abh. N. F. Bd. 12. Heft 4/5. Jena 1914. t. 3. f. 5. Siehe
subspecies Parelephas columbi felicis. He remarks (op. cil. p. 153) ,
auch unsere Taf. ix [xvii].'] von .Jockgrim in der Pfalz hat eine
"In der Art der Abkauung gleicht dieser Zahn sehr dem von Kaufliiche von 195 mm bei einer maximalen Breite von 82 mm. Das
Pohlig— Nova Acta Acad. C. L. C. G. Nat. Cur. Vol. 57 abgebil- Verhilltnis von Liinge zu Breite ist also iiber 2: 1 (Breite=l).

deten M^ des El. Americae Pohlig aus Chihuahua, Mexiko. Es Wilhrend bei El. Columbi die Verhaltniszahl f iir die Lange (Breite =
wird auch einM'von dort an gleicher Stelle abgebildet mit dersel- 1) unter 2 ist."

ben schiefen Stellung der Lamellen zur Liingsachse des Zahns wie Characters. — (Op. cil., p. 154): "Als altquartiir haben sich
an den Molaren von Ejutla und ganz verschieden von den Hoch- jene Formen herausgestellt, welche wir nach Falconer als Elephas
plateauformen der Mesa central von Mexiko oder Puebla. Es Columbi var. Falconeri bezeichnen miissen. Ihre Lamellen sind
empfiehlt sich aus oben genannten Griinden nicht, Pohligs Namen kiirzer und stehen isolierter (wie z. B. an dem Taf. viii [xvi], Fig.

El. Americae [^Elephas americanus De Kay, 1842] fur die Wald- 1 abgebildeten Molaren) verglichen mit der var. silvestris und erst

form des Cnlumbi anzuvvenden, dessen Verbreitungsgebiet die

El. recht mit der var. Felicis. Bei der typischen (altciuartiiren)
siidwestlichen Randgebirge des mexikanischen Hochplateaus
warcn. Ihnen gegeniiber stehen die als El. Columbi var. Felicis
aus der Mesa central bezeichneten Steppenformen.

Archidiskodon imperator falconer! Freudenberg, 1922

Figure 905
Tequixquiac, Valley of Mexico; probably Lower(?) Pleistocene.

This Tequixquiac jaw (Fig. 905) represents the cotype and

is undoubtedly related to Archidiskodon imperator rather than
to Parelephas columbi. From the figure it is difficult to give the
characters of this jaw, but appears to be somewhat longer and
it

more i)rimitivc than the type of A. imperator. The lower grinding

teeth apparently present the following formula: 3igTT7- In M
this jaw the 16+ ridge-plates exposed in the I.M3 are too widely
separated to be related to P. columbi; the I.M3 measures ap. 175+
mm., 95 mm., the dimensions a])parently equal those of P.
tr.

columbi, namely, neotype I.M3 ap. 298 mm., tr. 91 mm. The
locality of Tequixquiac yields both A. imperator and P. columbi,
according to Reyes.
El. Columbi Falconeri Freudenberg, 1922. "Die Siiugetier-
va,T.

und Postpliocans von Mexiko." Geol. und Pala;-

fauiia des Pliociins
oiit. Abhand., N. F., Band XIV, Heft 3, pp. 153-160. Lecto-
—
TYPE. First left lower molar, l.Mi, Mus. Royal Coll. Surg.
741a.' —
Cotype. Lower jaw with both third molars M3 in
situ. —
Horizon and Locality. Tequixquiac, Mexico. Lec-
—
TOTYPE Figure. After Falconer, 1863.1, PI. 11, fig. 1. Cotype
I'iGURE. — Freudenberg, 1922, p. 154, fig. 21.
Description. — {Op. cil., p. 153): "Als Typus der Form gilt
mir der von Falconer abgebildet* Mi von Mexiko (London) und
ein M3, den ich auf S. 54 [154], Fig. 21 abbilde. Dieser M' [M3]
gehrirt wohl der linken Seite an. Er stammt vermutlich aus dem
CLCPHis PRiaicemus.
\'alle Die Hinterseite des Zahnes ist abgebrochen. Die
de Mexico.
Zahl der fehlendcn Disken lasst sich nicht mehr bestimmen.
l'-hcns()\vcnig weiss viele Lamellen vorn durch Abkauung
man, wie
verschwunden Die Kaufliiche ist intakt und wurde in nattir-
.sind.

lichcr CJrosse abgebildet. Die Liinge der Kaufliiche misst 175 mm,
CoTYi'E OK Ahchidi.skodon imperatok falconehi Fhemdenbeho
die Breite in der Mitte, einschliesslich des Zements, misst 95 mm.
I'^ig. Cotype jaw (one-sixth natural size) of Eleplias Columbi var.
905.
Die Kaufliiche ist also etwa doi)pelt so lang wie breit. Der Ver- fig. 21; crown view one-half natural size.
Falconeri I'Veudenberg, 1922, p. 154,
schmelzungstypus der eben erst angekauten Disken (im Sinne der From Tequixquiac, Mexico. Originally figured by Villada, 1903, lAm. vm.
'[By inference this is Professor Osborn's IcctotyiJC. It is not figured, however, in tlie present Memoir -Editor.]
 —

THE MAMMONTIN.E: ARCHIDISKODON AND METARCHIDISKODON 1017

Columbi-Form. der gerade auch das Original Pohligs aiis Mexiko The principal partly or fully associated materials hitherto described
zugehort,kommen am oberen INP vier Lamelien auf 53 mm, are the following:
wiihrend deren 6 sich auf die gleiche Streeke verteilen bei dem jung- Figs. 907 and 906. Forelimb from Tule Canon, Briscoe
diluvialen Zahn von Mexiko Taf. viii [xvi], Fig. 4. Es mag nun County, Texas (Amer. Mus. 10598).
dieser weniger Waldform als Steppenform gewesen sein." Fig. 908. Femur from near Reynolds, Jefferson
County, Nebraska (Neb. Mus. 13-24-
EI. Columbi var. imperator Leidy 10-14).

= Archidiskodon Fig. 912. Forelimbs associated with numerous ver-

[ imperator] Freudenberg, 1922
tebrae, skull, jaw, one tusk, and parts
Spokam Bar near Helena, Montana.

Under the true specific name Archidiskodon imperator should

of hindlimbs — the finest material thus
far discovered [
= Archidiskodon im-
apparently be included the El. Cohimbi var. imperator of Freuden-
perator from near Curtis,
maibeni]
berg, 1922, in the Geological Mu.seum of Bonn.
Lincoln County, Nebraska (Neb. Mus.
El. Columbi var. imperator Leidy Freudenberg, 1922. "Die
5-9-22).
Saugetierfauna des Pliociins und PostpliocJins von Mexiko." Geol.
und Palaeont. Abhand., N. F., Band XIV, Heft 3, pp. 160-171.
Freudenberg frequently introduces the subspecific name
Elepha.^; columbi var. imperator Leidy (op. cit.,andpp. 155, 156)
finally definitely employs
on page 160.
it Type Description. —
{Op. cit., p. 160): "5. El. Columbi var. imperator Leidy. Wegen
der vielfachen Verwechslungen und wegen der mutmasslichen Ab-
stammung des El. Columbi von El. imperator ist es niitig, naher auf
diese Form einzugehen. Es ist das um so notwendiger, als H. F.
Osborn [Footnote: 'Recent Vertebrate Palaeontology. Fossil
mammals of Mexico. Science. Vol. 21. 1908 [1905]'.] diese Art aus
Mexiko anfiihrt. Osborn griindete seine Bestimmung wohl in
erster Linie auf den gewaltig langen Stosszahn, der 16 Fu.ss misst
und somit der langste Stosszahn ist, von dem eine Kunde vorliegt.
Aber warum sollte nicht ein El. Columbi gelegentlich riesige Dimen-
sionen erreichen, zumal da diese gute Art in eine ganze Anzahl von
Unterarten einmal zerlegt werden muss, sowie Matschie 4 Arten des
afrikanischen Elefanten heute unterscheidet."
—
Characters. After quoting Leidy, Osborn, Lull, Lucas,
Holmes, Pohlig, Soergel, and Cope {op. cit., pp. 160-169), Freuden-
berg concludes (p. 170): "'El. imperator' gewinnt mit seinem
Archidiskodon imperator. Aqed Male
Aufsteigen ins jiingere Diluvium immer mehr Aehnlichkeit mit den
Fig. 906. Outline reeon.struotion from right forelimb in the Ameriean
englamelligen, schmalkronigen, eigentlichen 'Columbi-Rnssen.'
Museum —
(Amer. Mus. 10598 Fig. 907) and from skull (Amer. Mus. 14476
Vielleicht losen sich die letztenHerde des waldlebenden Kaiser- Fig. 896), one-sixtieth natural size.
elefanten auf im Co/uw6/-Hauptstamm als sogenannte var. The forelimb (Amer. Mus. 10598) measures 348 2mm. or 11 ft. .')
in., as
silvetitris. Das hiingt doch wohl mit dem immer weiter ziiruck- shown in figure 912. This gives a shoulder height in the flesh of .3702 mm.,
weichenden Seen- und Waldklima zusammen, das offenbar der whirh may be taken as a conservative estimat<^ of the shoulder height in the
flesh of a nine-tenths grown Archidiskodon imperator. Compare figure 912,
Existenz der Imperator-Rasi^e in der Sonorischen Faunenprovinz
also figure 907, lateral and front views of the same forelimb, one twenty-fourth
giinstig war. Die zunehmende Austrocknung der diluvialen Seen-
natural The diagram of the forelimb, found in 1899 near Tule Canon,
size.
gebiete forderte die Entstehung von manmiutahnlichen Columbi- Texas, taken from the mounted forelimb in the Ameriean Museum (Amer.
is

Rassen, die wohl vorwiegend auf die sparliche Koniferenkost und Mus 10598). The diagram of the skull is taken from Archidixliodon imperator

auf harte Steppenpflanzen angewiesen waren. Die im tropischen (Amer. Mus. 14476), found at Victoria, Texas.

Laubwald iisenden Imperator-Hassen, die, wie El. indicus auf

Ceylon gute Bergsteiger gewesen sein mochten, haben bei Af ton wie Fore- and Hindlimbs of A. imperator (Fig. 907). — For-
im Becken von Puebla, Chihuahua und Mexiko als deutlichen tunately the complete right forelimb (Amer. Mus. 10598), dis-

Hinweis auf einstige Holzasung tief ausgehohlte Zementintervalle covered in 1899 near Tule Canon, Briscoe County, Texas, along
zwischen den ausserst kraftig gebauten Dentinpfeilern und ihrer with other parts of the skeleton belonging to a single individual,
oft stark gefalteten Schmelzhiille." gives us the means of estimating the height of Archidiskodon
imperator, namely, 12 ft. Wa in. =3702 mm. at the withers, 13 ft. =
SKELETAL CHARACTERS OF ARCHIDISKODON 3960 mm. at the top of the head when elevated. This elevation is
Much remains to be done in establishing the skeletal characters shown diagrammatically in figure 906 which combines the skull
of Archidi.'ikodon imperator. In the flu\'iatile sand and gravel (Amer. Mus. 14476) from Victoria, Victoria County, Texiis, with
deposits of early Pleistocene time, in which these imperial mam- the forelimb (Amer. Mus. 10598) from Tule Canon, Briscoe
moths occur, skulls, jaws, and skeletal parts are widely scattered. County, Texas.
1018 OSBORN: THE PROROSCIDEA

In the licli collections made by Professor Barbour for tiie

State .Miiseiini, I'liiversity of XclHaska, are a femur (Fis. 908) and

a humerus which may be comiJaretl with corresjjondinf^ limb ele-
ments in the skeleton of "Jumbo" {Loxorlonta afrirana oxyotis)
in the American Museum. The specific reference is somewhat
uncertain. The fenun- probaby does not represent a full-grown
A>-chidisko(Ion imperator, which would greatly exceed the specimen
here represented in the pliotoKraph (Fifi. 90S) kindly furnished by
Professor Harbour.

I'oltKT.lMB OK ArCHIDISKODON IMI'EIIATOK. A.VIEHirAN MusEUM

Compare Figure 912
rin- iH)7. Rislit forclimb roforrod to ArclwHskudon imperator, a.s moiintcil •ig. 908. l-iMH Bones of Ahchiuiskodo.n impek.^toh (Vouncj Adui.t) and
in tlio .AmiTicaii Museum (.Amer. Mus. 10.")98). one twenty-fourth natural l.OXODONTA AFRICAN A OXYOTIS COMPARED
-.ize, found l)y .\ll)an Stewart in 1899 nearTule Canon, Bri.seoe Covnity, Texas, (bower) Three views (E) of femur of referred Arrhidiskmlon imperalor
;il()nKwith other parts of the skeU'ton belonging to a singh' in(livi<luah (Neb. Mus. 13 24 10 14) from Reynolds, .leffer.son County, Nebraska, one-
The same forelimb is represented, vertically extended, in figure 012. The tenth natural size. Height of fenuu- 1422 mm. =.")G in. (Barbour, 1925.3, p.
vertieal measurements of the separate .segments are: Scapula 1017 mm., llfi), .same measurement as the fenuw (Neb. Mus. 4-12-13), namely, 4.8 feet.

humerus 109.') mm., radius 890 mm., manus 480 mm., sum of total vertical (Upper) The same femur (Neb. Mus. 13-24- 10 14), one-twentieth
height 3482 mm. or II ft. in. .Xs the limb is always somewhat flexed, the
.'>
natural size, compared with (.\, B) femur and humerus of Loxndonta tijri-
cartilages, foot pads, and the flesh and skin above the shoulder give this cana oxyotis (skeleton of ".Jumbo") in the American Museum; also with
animal a total height of 3702 mm., or 12 ft. ]\ in., a conservative estimate of (C, D) humerus and femur (latter computed) referred to Arc)niHifkn(Um im-
the height of this animal. peralor (Amer. Mus. IO.jOS). Compare figure 907 oi)posite.
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1019

HAUHonirS DESCRIPTION (1925) OF THE SKELETON OF flocks of itoultry on the Karriger farm, before it was realized
ARCHIDISKODON MAIBENI that they were out of the orilinary, after which the remaining
(Soo type description of skull, jaws, and dentition below.)
parts were dug out and bared for with unusual appreciation and
As above noted, this is the most nearly complete skeleton of discernment. . . . The bones of Elephas maibeni were found pro-
Archidiskodon thus far discovered. The following is a free rather jecting from a loessial wall at the bottom of a small canyon. The
than a literal citation from Professor Barbour's \aluable paper of is about 100 feet.
general thickness of the loess at this point In an
August, 1925 (Barbour, 1925.3), entitled, "Skeletal Parts of the attempt to find additional material the writer, aided by Mr.
Columbian Mammoth, Elephas maibeni, sp. nov." For reasons William Hall and Mr. H.8. Karrigcr, blasted out many cubic yards
shown in the systematic treatment of E. (Archidiskodon) maibeni of the loessial wall."
below, there appears to be no doubt that this skeleton is properly "The skeletal parts preserved are the skull, mandible, one

Fig. !MHI. Im|ii'iiul Mammoth (Archvlukodon iiii/xralor) of Nebraska and Texas. After restoration by Osborn and KniKJit, 190S. .\bout one-fiftieth
natural size.

referable to Archidiskodon rather than to Elephas; also that it is tusk, the atlas, axis, and four other cervicals, several thoracics,
closely related to Archidiskodon imperator rather than to Parelephas lumbars, and the sacrum, ribs, and double ribs, both fore limbs and

colmnbi. The narrative of discovery is as follows: parts of the hind limbs. Both fore limbs are practically complete
"The and most remarkable specimen of the columbi
last and are essentially ])erfect save that but one foot bone was found,
[imperator] type was found in IJncoln County, about 16 miles hence the feet must be supplied. The hind quarters are represented
north of Curtis, on the Karriger farm. It was discovered by Mr. by parts of the peh'is, the shaft of a femur, and the major portion of
and Mrs. H. S. Karriger, and was dug out antl preserved by them. a fibula. The dentition is perfect. The molars have 14 ridges
Later it was procured of them for the palaeontological collections of bonded together by an uncommon thickness of cement, which is
Mr. Hector Maiben, who, next to Mr. Charles H. Morrill has been a character of Columbian elephants. Even the great Columbian
the most generous contributor of funds for the purchase and pres- elephant from Franklin Count}' [referred to Parelephas jeffersonii
ervation of choice Nebraska specimens. ... It should be recorded in the present Memoir] seems surpassed in size. Heretofore, the
in connection with this specimen that an unknown number of bones tusks of the Franklin County elephant [Neb. Mus. 1-4-15] have
and jjurts of bones were poimded uj) to furnish lime for the large been considered the largest repoi'ted, namely about ISli feet long
1020 OSBORN: THE PROBOSCIDEA

with a maximum circumference, near the incisive sheath, of 29 to note that the cancellous portionis uniform and continuous and

inches. The skull and mandible of Klephas maibeni is noticeably iswithout partitions or vestiges of the dual origin. From this it
larger. The incisive sheath shows that the tusk had a diameter of may be inferred that they had been in coalescence for ages and
lO'i inches and the incredible circumference of 33 inches." that the character may have become fixed. At any rate it is not
"The outstanding characters of Elephas = Aickidiskodon\ [ a ca.se of pathology. The great fore limbs, several vertebrae, and
maibeni are size, extreme curvature and divergence of tusks and their corresponding ribs, have been mounted as an arch, a palaeo-
incisive sheaths, unusual shortness of centra coupled with great zoologic arch [Fig. 910 of the present Memoir], through which all

width. The tusk must have lain in a plane or nearly so and must students and visitors must on entering the main floor. It is
ptiss

have de.scribed a circle, the radius being 28 inches (711 mm.). but a temporary mount which must be dismantled and moved into
The diameter of the tusk i.s 6^i inches (165 mm.) at the tip, 7U the new museum sometime in 1926, where the complete skeleton
inches (190 mm.) four feet back of the tip, and IOJ2 inches (297 will be carefully articulated and properly installed. So many
mm. [267]) at the incisive sheath. Originally it was a magnificent skeletal parts are at hand that this huge elephant when ready for
jiiece fif ivory." exhibition will seem complete. Its proper installation demands
"The fore quarters have bones unexpectedly large and mas- a ceiling 18 feet high." [See Fig. 911 for present moimt. Eflitor.] —

Fig. 910. I'Virolimbs of typo .skolefun of

ArcMdinkoilon imperator maibeni as moimted
in the Nebraska State Museum (Neb. Mus.
.")
9-22) in the year 192.'). Professor Barbour
is .standing in the background. The dimen-
sions of this sii|)erb skeleton are shown (Fig.
912) in direet comparison with the forclimbs
of .1. imperator from Te.xas, of Loxmlontn
and of Elephas indiciis.
(ifricana,
The cranium and tu.sks in the back-
ground to the left, from Franklin County,
Nebraska, belong to Parelephas jeffersoiiii.

Barbour notes (192.').3, legend to Fig. .')8):

'The great .skull background

and tusks in the
are tho.se of Elcjjhas columbi from .Jefferson
|Franklin| County."
[For |)resent mount, see figure 911 on
following page. — Editor.]

sive, especially the humerus. The humerus is huge beyond the "Mammoth scapulae are large, heavy, and very thin in portions,
visualization of those who have hence must judge of it
not .seen it, so it seems the more remarkable that the two huge shoulder blades
from figures and measurements. In the hind quarters the bones should have been preserved practically without blemish. The
are, if anything, less massive than might be exi)ected. They seem right humerus is likewise perfectly preserved; in the left the head
in contrast to those of the fore limbs, .lutlgiiig from the very short is wanting but has been modeled on from the right humerus. The
vertebriE the body must have been unduly foreshortened." right ulna is perfect save that the distal epiphysis is missing. This

"The centra of the vertebrae are very short, comi>ared with has been modeled after the left ulna in which the ej)iphysis is
their width, ('onsecjuently certain ribs, presumably the fifth and l)resent but the shaft missing. . . . The fore liivibs, four vertebrae,
sixth pairs, came
contact and became completely fused into one
in two pairs of single ribs, and one i)air of double ribs of this ex-
shaft with a double head and double tubercle, making huge and cej)tional mammoth are mounted in approximate position and
peculiar mammal ribs. A slight longitudinal depression is a vestige make an impressive arch, the height of which is 13 feet from the tip
of the original boundary between the two shafts. It is interesting of the toe to the top of the spine, see fig. 58 [Fig. 910 of the present
 :

THE MAMMONTINiE: ARCHIDISKODON AND METARCHIDISKODON 1021

Memoir; see also Fig. 912]". The tallest living African elephant stands 11 feet high and the
"In the flesh the height of Elephas [
= Archidiskodon] maibeni average elephant of the menagerie and circus 8 to 9 feet."
at the shoulder must have been about 13 feet, and the top of the Referring to figure 912 of the present Memoir, we observe
head of this magnificent beast must have been about 14 feet above that A 1 (right) is foreshortened, while A, B, C, D represent ortho-

the ground." gonal full length projections of each limb segment, with the actual
measurement of each segment in millimeters. This affords an
"From the tip of the toes to the top of the scapula is 11 feet, 6
absolutely reliable comparison of the ascending height of these
inches."
four animals.
"This specimen is believed to hold the record for size amongst
The following entirely consistent comparative measurements
the Columbian group of mammoths." appear to demonstrate that Archidi.ikodon rmperntor and the more
"In point of .size, Elephas [
= Archidiskodon] maibeni was primitive giant species A. maibeni towered in height far above the
a rival of the Imperial elephant itself, which stood 13Ji feet high. largest existing elephants

Fig. 912, A, Archidiskodon imperator maibeni

B, Archidiskodon imperator
C, Loxodonta africaiiri oxyoiis (".lumbo")
D, Elephas indicus
Fig. 1083, Hesperoloxodon antiqiiiis (Upnor)
 Est. height
r-^OOO Imperial Mammoth (mailreni)
Imperial
'^ Mammoth Est —3702
height
African Elephant Max. height
-3568
Indian flephant
Max. height .3/94-
-3200
300O '3007

-2000 V/

o
\ 2

1000

INDICUS
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1023

The principal measurements of the type skull and skelet

 .

1024 OSBORN: THE PROBOSCIDEA

process is conspicuously robust, boing 2?^ inches (70""") through there are but 10 in one tooth, and 11 in the other, with no plates
near its base, and an inch (25"'"') near the summit. It stands 4 missing. . In E. hayi, there are 4 and a fraction transverse
. .

inches (102""") above the superior mandibular border, and 2 enamel ridges to the decimeter. The valleys are deep and bordered
inches (51 """) above the crown of the teeth. It is set more by highly crenulated enamel ridges. The great anterior prong
obliquely than in other mammoths. Its inner surface is deeply branches widely and carries 3 plates. The teeth lack the sym-
and extends from the outer to the inner alveolar border.
pitted, . . metrical development common to mammoths. They are notice-
The teeth are those of a mature individual, with the crowns well ably constricted back of the anterior prong, and taper posteriorly
to 1)2 inches (38"'™) there are but 11 transverse ridges at most
. . .

[to each molar tooth], the last being small, perhaps a heel. This
form seems to be an earlier and more primitive type of mammoth
than any other known to the State [Nebraska]. The inferior dental
foramen is small, and has a circular border, while in E. imperator
it is very large and deeply notched, as shown in the accompanying

/^// to ya /Vat. sije

J From 5iwaZi7<5 \

Barbour's Type .Iaw of Archidiskodon hayi

Fig. 913. Ty|)e of Ekphas haiji Barbour, 1915, p. l.W, fig. 1 (Nob. Mus.
23-G-14), one-eighth natural .size. Ramus perfect to summit, condyle only
restorpd.
Compare jaw of Archidixkoilon jiltinifrona of Chagiiy, France (I'ig. 914).
Profe.s.sor Barbour (letter of .January 17, 1923) remarks: ". . . we both From 5 en 676
examined it [the type] as well as we know how, and count it an old not a young
individual. I feel quite .sure of this. [E.\ haiji is very distinctive."
ARCHIDISKODON PLANIFRONS
worn. Though well cemented and strong, the teeth of E. hayi are
noticeably small. The postero-anterior diameter is but 9 inche.s Fig. 914. Primitive mandibles of Ehphas [Archidiskodon] planifrons of
"'"'). the Siwaliks, India, Chagny-Bellecroix and Seneze of France, after Mayet
(229 "'"'), and the greatest transverse diameter 3 inches (76
and Roman, 1923, p. 81, fig. 13, in.serted for compari.son with type figure of
. . . The dimensions of these teeth agree more closely with those See caption to figure 849,
E. [Archidiskodon] hayi (Fig. 913 opposite). p.
of our earlier Nebraska mastodons than with those of our mam- 962 above. One-eighth natural size. Ob.serve .similarity to th<! Chagny
moths. The number of transverse plates is noticeably reduced, for mandible.
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1025

figures. Although inferior dental foramina differ in individuals, Archidiskodon imperator scotti Barbour, 1925
and even between opposite sides of the jaw, the differences shown =A. imperator ?juvenile]
[

by the cuts are significant. The ascending rami of our probo- Figures 916, 917, PI. xxi
scideans also vary between wide limits." Five miles south of Staplehurst, Seward County, Nebraska. Lower
Specific Characters. —
(1) Mandibular ramus elongate, de- Pleistocene, Aftonian gravels.

pressed, coronoid process low. (2) Third inferior molars broad in Awaiting further evidence, Osborn is inclined to regard the

center, with ten to eleven ridge-plates set very far apart; laminar type of 'Elephas scolti' as representing a young individual of

frequency i/i in 10 cm. molars narrowing posteriorly. (3) Ramus

;
A rchidiskodon imperator.

of jaw shallow, moderately expanded; rostrum relatively promi- This Archidiskodont, discovered in 1922 in the Aftonian
nent. (4) Agreeing with Archidiskodon planijrons in ridge formula, gravels, 5 miles south of Staplehurst, Seward County, Nebraska,
differing in greater width of M3. (5) Dimensions of M3, length 229 and 20 miles south of Crete, Saline County, where the type of
mm., maximum breadth 76 mm., index 33.
Comparison with Archidiskodon planifrons (Osborn,
1924). —
It is remarkable that the type of Archidiskodon hayi re-
sembles closely in the profile of the jaw the Archidiskodon plani-
frons of India and of southern France, apparently justifying Bar-
bour's statement that A. hayi seems to be an earlier and more
l)rimiti\e type of mammoth
than any other known to the state
[Nebraska], and suggesting the possibility that we have to do with

Fig. 915. Barbour's Type of Archidiskodon hayi compared with A. imperator Ree.

(Left lower) Type mandible of Archidiskodon hayi, sectioned at three points, grinding teeth with lO-U ridge-plates.
right
(Left upper) Archidiskodon imperatorref. Same aspect of right mandibular ramus, grinding teeth with 18 ridge-plates.
(Right upper and lower) Archidiskodon imperator with 18 ridge-plates, lateral and superior aspects of mandible.
Both specimens in the Nebraska State Museum, Morrill Collection. After Barbour, 1915.2, figs. 2, 3, and 4.

the arrival in North America in the late Pliocene or early Pleisto- Elephas = Archidiskodon] hayi was found (1914), is regarded by
[

cene of a primitive proboscidean, displaying some of the chief A. Barbour (1925.1, p. 22) as a mature individual, as primitive as
planifrons characters, namely: (1) Jaw long and shallow; (2) A. hayi, if not more so. This statement rests upon the identifi-
rostrum prominent; coronoid relatively depressed; (4) grind-
(3) cation of the small 8 ridge-plated molars as representing third
ing teeth broad with 10-11 transverse ridge-plates. Supplementary inferior molars, M3, which appears to Osborn very doubtful,
figuresand sketches of the jaw and teeth of A. hayi (jaw. Fig. especially as the jaws of the type of 'Elephas scotti' (Fig. 916) are
893 A) serve to emphasize the wide contrast between this type very robust in section with greatly abbreviated symphysis, alto-
jaw and the adult jaw of ^. imperator (Fig. 892 B, A). gether different from the relatively slender, elongated mandibular
 —

1026 OSBORN: THE PKOBOSCIDEA

rami of A. hayi. Consequently it seems probable that they are FiGUKE. Op. cii., text figures 7-10.
second inferior molars, M2. Type Description.— (Barbour, 1925.1, pj). 21-24): "On
Elephas srolli Barbour, 1925. "Elephas scotti, A New- February 18, 1922, the mandible of an unusually primitive mam-
Primitive Mammoth from Nebraska." The Nebraska State moth was secured for the palaeontological collections of IMr. Hector
Museum, Bull. 2, Vol. I, April, 1925, pp. 21-24 (Barbour, 1925.1). Maiben by Mr. E. T. Engle. The peculiarities of this mam-
. . .

Type. —
Mandible with last [second?] lower molar of each side moth seem to entitle it to a position as a di&tinct species, for which
(Neb. Mus. 18-2-22, the Maiben Palaeontological Collections. we are proposing the name, Elephas scotti, named for Profes.sor
Cast of type right M3 [r.:M2], Amer. Mus. 14610). Horizon William D. [B.] Scott. It cannot be compared with the later and
.\ND Locality. —
Discovered on farm of Mr. E. J. Hartman, five more advanced mammoths, such as imperator, jeffersoni (columbi),
miles south of Staplehurst, Seward County, Nebraska. Type or jjrimigenius. comparable instead, with the earlier and more
It is

conservative mammoth, Elephas hayi. The new mammoth is

. . .

as primitive as Elephas hayi, if not more so. Like hayi, it is un-

doubtedly a mature individual. Its teeth are taken to be last
[second] molars. The enamel plates, which are highly crenulatcd,
incline noticeably backward, and are worn with extreme obliciuity.
At the same time the valleys, or dental spaces, are so deeply in-
dented, as to still further heighten and exaggerate the effect. There
are but five pronounced ridges, and in all, but eight and a cone.
Two of the anterior ridges are so confluent that the count is ren-
dered somewhat uncertain, as shown in figure 10 [our figure 917A,
Al, A2]. The transverse ridges in E. hayi are eleven. The molars
of scotti measure 219 mm. (8\ in.) in length, by 117 mm. (4!'8 in.)
in extreme width. They are short and abrujjtly expanded in the
middle. In the mammoths the number of enamel ridges to the
decimeter serves, in a general way, in the recognition of s])ecies.
In E. scotti there are three and a fraction, transverse, grinding
ridges to the decimeter; in E. hayi four and a fraction; in E.
imperator five to six; in E. jeffersoni (columbi) six to eight; and
in E. primigenius nine to ten. It is a noteworthy feature that the

robust jaws of Elephas scotti come within three-fourths of an inch

of meeting on the middle line, as is plainly shown in the figures
[see Figs. 916 and 917A of the present Memoir]. This is not due to
crushing, as far as can be learned, for the specimen in hand is
essentially perfect. The coronoids of the earlier and the later
mammoths differ widely and are worthy of notice. Those of scotti
and hayi are much more robust, thick, and heavy, and flare out-
wardly, and are posterior to the molars. The inner wall is broader
and more heax'ily roughened and pitted for ligamentous attach-
ment. Each ramus, measured back of the molar, has a width of
185 mm., (7X6 in.) and a depth of 180 mm., (7's in.). On the middle
line the jaws are but 19 mm., (:'.| in.) apart."
Osborn, 1928: The Seward (^ounty jaw (Neb. Mus. 18-2 22
— see Fig. 916, and as seen from abo\c. Fig. 917.\) in Osborn's
opinion resembles a juvenile jaw of .4. impcrnlor rather than the
type jaw of E. hayi Barbour. In superior view (Fig. 916) the jaw
appears fairly robust, less swollen than that of A. iinpcralor, with
highly characteristic outwardly flaring coionoid i)r(>cesscs. In
lateral view (Fig. 916) it is relatively short and deej), the rostrum
is and dejiressed, in wide contrast to the long, shallow jaw of
short
the tyjje of A. hai/i. The single grinding toolii, in which 8 9 ridge-
plates appear, may represent M2 of a young A. imperator; the
ric- I'I'i. .Juvenile jaw of Archiilinkndnn impiralur ref. (Neb. ^I^l.>^. tooth is obli(|uoly worn and conseciuenfly the dental space between
IS 2-22), friini Sew;ir(l Coiinly, Ni'bniskji. After plKitonnnih by Prof. the broad enamel ridges a|)i)e;ns to be much greater than it actually
E. II. Hiirbiiiir wlm in l!(2.') niiuie lliis tlie type of Eliplmx xmlli. ('ompMre is; in the actual distance between th(^ ridge-plates
the disparity
figure !(I7. Seiile about one-sixth natural size.
(Fig. 917 A2) and the a|)pai-eiit distance due to obliriuity of wear
Tlio.se 8 ridge-platfd griiulinK teetli, deseribed as third ii)f<'rior molars,

M3, are regardi^d by Osborn as seeond inferior molars, Mj (ettst of r.M'j, .\nier. (Fig. 917 Al) are clearly shown in this diagrammatic represen-
Mus. 14610). tation.
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1027

Archidiskodon imperator maibeni Barbour, 1925 thoracics, lumbars, and the sacrum, ribs, and double ribs, both fore
(Sec full (l('SiTij)tion above, page 1019, of the skeleton of .1. maibeni) limbs and parts of the hind limbs. . . . The hind quarters are
Figures 815, 824, 910-912, 917, 918, 1239, PI. xxi represented by parts of the pelvis, the shaft of a femur, and the
Tiineolii County, about sixteen miles north of Curtis, Nebraska. major portion of a fibula. The dentition is perfect." Neb. Mus.
Aftoniaii,' loes.s 100 feet in general tliiekness. Upper Pleistocene (see 5-9-22. —
Horizon and Locality. Discovered by Mr. and
I'ig. 1239, also PI. viii, Vol. I). Mrs. H. S. Karriger about sixteen miles north of Curtis, Lincoln
The type an unusually complete skeleton,
of this species is County, Nebraska, on the Karriger farm. Type Figure. —
entitled the 'Lincoln County Mammoth,' in the Nebraska State Op. ciL, Figs. 58-60, 63-70, 72, 74, 76-87.
Museum, described by Barbour in 1925 (1925.3) as the "Columbian Type Description.— (Barbour, "Con- 1925.3, pp. 97-111):
Mammoth Elephas maibeni, sp. nov." and subsequently (June, fusion has long and jeffersonian
surrounded the Columbian
1926) transferred by him to Archidiskodon maibeni, now mounted mammoths. But the one under consideration is undoubtedly of
in the new Museum, Morrill Hall, University of Nebraska. To the true Columbian type. The bones of Elephas maibeni were

Neb. Mus. 18-2-22

I"ig. 917. Comparison of : = A rrhidiskodon iiiiperalor maibeni] (Neb. Mus. 5-9-22), with
(B) Mandible of the t yp<' of Elephas (Archidiskodo7i) maibeni [

third inferior molars, M3, in plaee, exhibiting 1.')+ ridge-plates, 14 exposed; type jaw of Elephas scolli [= Archidiskodon imperator scolti or ?juvenile
and (.\)

A. imperator] (Neb. Mu.s. 18-2-22), containing second inferior molars, M2, exhibiting 8 ridge-plates, and measuring ap. 219 mm., tr. 117 mm., i.e., short and
abru])tly expanded in the middle; \l, A2, enlarged views of right second inferior molar of the type of 'E. scotii' (cast Amer. Mus. 14610).

the above full description of the skeleton (p. 1019) by Barbour found projecting from a loessial wall at the bottom of a small
(1925.3) may now be added the type description and characters of canyon. The general thickness of the loess at this point is about
the skull and dentition (Barbour, 1926.1), as follows: 100 feet. The outstanding characters of Elephas maibeni are
. . .

Archidiskodon maibeni Barbour, 1926. Professor Barbour in size, extreme curvature and divergence of tusks and incisive

his supplementary description ("Archidiskodon maibeni," Nebras- sheaths, unusual shortness of centra coupled with great width.
ka State Museum, Bull. 11, Vol. I, June, 1926, pp. 119-122) states: The tusk must have lain in a plane or nearly so and must have
"Archidiskodon maibeni was first described in Bulletin 10 of the described a circle, the radius being 28 inches (711 mm.) The
Nebraska State Museum under the title 'Skeletal Parts of the diameter of the tusk is 6)2 inches (165 mm.) at the tip, 7'; inches
Columbian Mammoth, Elephas Maibeni, sp. nov.'" Under this (190 mm.) and 10).^ inches (297 [267] mm.)
four feet back of the tip,
designation I^arbour gives additional measurements and comments at the incisive sheath. it was a magnificent piece of
Originally
on the skeleton. He also mentions the discovery of another skull i^'ory. The humerus is huge beyond the visualization of those
. . .

(Neb. Mus. 1-4-26) referable to A. maibeni. who have not seen it, hence must judge of it from figures and
Elephas maibeni Barbour, 1925. "Skeletal Parts of the measurements. In the hind quarters the bones are, if anything,
Columbian Mammoth, Elephas Maibeni, sp. nov.," Nebraska less massive than might be expected Judging from the very
State Museum, Bull. 10, Vol. I, August, pp. 95-1 18. Type.— short vertebrae the body must have been unduly foreshortened.
(Op. cit., p. 98): "The skeletal jjarts ])reserved are the skull, . .The skull is broken into several large, and numerous small
.

mandible, one tusk, the atlas, axis, and four other cervicals, several pieces, which have not been set permanently in place. The . . .

'[Lugn and Schultz (1934.1, p. 376, also Tabic A) regard it as of lowan, late Pleistocene age. — Editor.)
1028 OSBORN: THE PROBOSCIDEA

accompanying free hand sketches were made with the pieces set apart. . The anterior borders of the coronoids are excessively
. .

approximately in position, and give our impressions of the skull. roughened; they are even nodular, and we find no parallel. . . .

. .The head must have presented a bull-dog effect or aspect.

. The lachrymal process, which is uncommonly large, prominent,
It might be called the bull-dog mammoth. The mandible is . . . and acorn-shaped, is dissimilar to all others available for study and
well preserved, and in point of size passes the largest in our col- comparison. The fore limbs, four vertebrae, two pairs of single
. . .

lections. The rami are widely divergent and the condyles far and one pair of double ribs of this exceptional mammoth are
ribs,
mounted in approximate position and make an impressive arch,
the height of which is 13 feet from the tip of the toe to the top of
the spine, see fig. 58 [Fig. 910 of the present Memoir]. In the flesh
the height of Elephas maibeni at the shoulder must have been
about 13 feet, and the top of the head of this magnificent beast

Bl y.

BaKBOUr's TyI'E of AnCHIDIKKODON IMI'EUATOU MAIBENI

Ji'lg. 918. .Irc/iidwfcodontmperatorjKaibem', tho 'Lincoln County Mammoth,' .superior and inferior dentition of (lie type skeleton (Neb. Mus. 5-9-22)-
.\fter pliotograph.s kindly furnished the present author by Prof. E. H. Barbour (compare Barbour, 1925.3, figs. 64, 65, 67, 68).

.\, MandibU' of .1. imprrator maibeni from above, about one-eiglith natural size.

Right and left inferior molars, r.Ms, I.M3, with 15-|- ridge-plates, 14 exposed, more or less worn, broad < inieiit, somewhat .sinuous, but slightly
concave posteriorly, crown externally plane, internally convex.

Al, The same in lateral view. One-eighth natural size.

B, Palate containing right and left third ,su|«Tior molars, r.M'', I.M''. One-eighth natural size.

Bl, Right third suijcrior molar, r.M'. One-fourth natural size.

C, Extremity of tusk 4 feet, in lengtli measured mi outer curve. About (ine-tcnth natural size.

Obs<'rve in B, Bl, 16+ ridge-plates, of which I 13 show wear, (|uite .strongly concave posteriorly with heavy border of cement; crown externally
.signs of
convex, internally slightly (concave. Very similar in —
contour and ridge formula to A. imperalor (.\mer. Mus. 14470 Fig. SSOB), from Victoria, Texas.
Observe in A, Al, 15-f- ridge-plates, 14 ex|«jsed.
 THE MAMMONTIN^: ARCHIDISKODON AND METAIICHIDISKODON 1029

must have been abuut 14 feet above the ground. From the tip of at Frederick, Oklahoma. Type Figure. — Hay and Cook,
the toes to the top of the scapula is 1 1 feet, 6 inches. This specimen 1930.1, Pis. Ill, fig. 1, v, fig. 1, XIII and xiv.
is believed to hold the record for sizeamongst the Columbian Type Description.— (Op. cit., 1928, p. 33): "Elcphan harold-
group of mammoths. In point of size, Elephas maibeni was a rival cooki Hay. Based on a nearly complete lower jaw containing the
of the Imperial elephant itself, which stood l.S}^ feet high. The right and left last molars. Anterior fang and 2 ridge-plates absent
tallest living African elephant stands 11 feet high and the average through wear. 12 ridge-plates and the rear talon present. 4.4
elephant of the menagerie and circus 8 to 9 feet." ridge-plates in 100 Enamel thick, moderately folded. A
mm.
loxodont expansion present in some of the ridge-plates. The crown
very high. Found in Holloman's gravel quarry at Frederick,
Oklahoma. No. 1057, Colorado Museum of Natural History,
Denver."
Supplementary Description (Hay and Cook, 1930.1, pp.
32, 33).
—
"This jaw was embedded in the cemented gravel very
close to the Permian red clay ... It will be seen that nearly all of
the jaw behind the teeth These teeth are in fine con-
is lacking.
dition and show that the animal was somewhat beyond middle age.
The height of the jaw where the ascending ramus arises is a little
more than 200 mm. Its thickness, where greatest, at the middle of
the tooth is 155 mm.; the width of the bone, taken at the rise of
the ascending rami, is about 390 mm."
"As will be observed from the views of the jaw, it is wholly
without a beak in front; and does not turn downward, also the
symphysis is short, about 60 mm. in length. The teeth present are
])arts of the hindmost molars. This is shown by the arrangement
of the rear ridge-plates, only partly shown in the figures. In front
of each tooth (plate iii, [=Fig. 919 of present Memoir]),
figure 1
especially of the one of the right side, is seen a cavity in which was
lodged the anterior fang. The tooth had been worn down to the
bar of bone separating this fang from the part behind it and the
fang had fallen out. With it went three worn-out ridge-plates.
Behind the bar of bone may be counted on the grinding face 11
ridge-plates, the tenth being represented by a small circle of enamel,
the eleventh by a dot of enamel. The length of the abraded surface
of the molar (plate v) is 182 mm. its width, with the cement, is
;

85 mm. without it, 80 mm."

;

"Behind this eleventh ridge-plate is a mass which may be

regarded as a talon, a ridge-plate as yet undeveloped. Adding
now to these eleven ridge-plates the two supported by the anterior
fang and the one by the bar of bone behind it, we have 14 ridge-
plates for the hindmost molar of the species. From plate xiv it is
seen that the ridge-plates are very thick. Measured at one-half
their height three are spanned by a line just a little less than 75 mm.
Type of Archidiskodon hakoldcooki
Measured at one-half their height 4.44 plates are crossed by a 100
Fig. 919. Typo mandibk' and tliird inferior molar of tlii' right .side

in silu of Elephas haroldcooki Hay, 1928. After Hay and Cook, 1930, PI. iii,
mm. line. The hind end of the crown is almost 6 inches high,
fig. 1. One-fifth natural .size.
nearly as high as the grinding surface was long."
"The view (plate xiv) of the inner face of the tooth shows
that the ridge-plates are at first directed strongly forward, then
Archidiskodon haroldcooki Iluy, 1928 arc turned abruptly upward and (in relation to the grinding face)
Figu"' 919 somewhat backward. It is due to this oblic|uity that the front
Found in Holloman's gravel quarry, Frederick, Oklahoma. Aftonian? enamel plate of the hinder ridge-plates is more exposed to view
gravels. than they are farther forward. It will be seen that the outer ends
Elephas haroldcooki Hay, 1928. "Preliminary Descriptions of of the loops of enamel are rather strongly turned forward. The
Fossil Mammals Recently Discovered in Oklahoma, Texas and enamel, as usual in primitive elephants, is thick, here about
New Mexico." Proc. Colo. Mus. Nat. Hist., Vol. VIII, No. 2, 3 mm. It is moderately crimped. A feature of interest is the
Pt. February 2, 1928, p. 33.
1, Type.— Nearly complete presence of a lozenge-shaped expansion at the middle of each
lower jaw containing last right and left molars in situ (Colo. Mus. ridge-plate. This is a characteristic of the earlier elephants, as
1057). HouizoN AND Locality. — Holloman's gravel quarry, E. planifrons, E. meridionalis, and E. imperator."
 ARCHIDISKODON /MPERATOR /?e/"
DW.,.,.- I.vs.,.,. ,S...OKS
OP AkCH.„,SKODON- ..OM,.U,0„ „,..„
A. ,M,.K,UT„K

1030
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1031

Archidiskodon exilis Stock and Furlong, 1928 recently summarized the available information and recognizes the
Figures 815, 920, 921, PI. xxi presence of Elephas imperator and of an undetermined species.
Santa Rosa Island, California. Plcistocrno. During the past year Dr. Spencer Atkinson and Mr. J. A. Barbieri,
This dwarfed insular form of imperial mammoth, estimated of Pasadena, secured a fragmentary elephant skull on Santa Rosa
at 6-8 feet in height measured at the shoulder, as compared with Island and presented the specimen to the California Institute of
12 ft. 1% in., distinctive height of Airhidiskorlon imperator. is of the Technology. Through the courtesy of the ^'ail Company of Los
greatest interest. There doubt from the drawings recei\ed
is little Angeles, owners of the island, the California Institute, with the
(Fig. 920) that this is a diminutive insular form related either to cooperation of the Carnegie Institution of Washington, have been
A. imperator or to Parelephas coliimhi. given the oi)]iortunity to investigate the occurrence, and facilities
Elephas exilis Stock and Furlong, 1928. "The Pleistocene were kindly made a\ailable to collect further remains. . The
. .

Santa Rosa Island, California," Science, Vol.

l^lci)hants of LXVIII, collections secured by the California Institute include a number of
No. 1754, p. 140.
—
Type. "A skull and mandible including teeth, parts of skulls and skeletal material. Occasionally several
four cheek-teeth and two tusks" (Calif. Inst. Tech. Coll. Vert. skeletal elements and teeth are found associated in the deposits.
Pal. 14). Horizon and Locality. Santa Rosa Island, — Usually the remains are scattered. One curious feature of the
California, the second largest of four islands separated from the occurrence is the ai)parent total absence of associated mammalian
mainland by the Santa Barbara Channel. Pleistocene. Type types. The proboscidean remains are referable to the genus

Restoration of Archidiskodon exilis of Santa Rosa Island

Fig. 921. with tusks and lower jaw, of Elephas exilis Stock and Furlong, 1928. From
Fai-ial [lortion of tlio skull,

Quaternary deposits, Santa Rosa Island, California. After unpublished photograph kindly furnished by Dr. Cliester
Stock.

Figure.— Stock, 1935.1, p. 210, fig. 6. Supplementary De- Elephas. The individuals exhibit considerable variation in size,
scription. —Stock, "Exiled Elephants of the Channel Islands, and this is undoubtedly to be a.scribed in part to differences in age.

California," Scientific Monthly, 1935, XLI, September, pp. 205- A survey of the collection as a whole yields the impression rather
214, text figs. 1-10. strongly that the elephant types were of relati\-ely small size.
—
Type Description. (Stock and Furlong, 1928.1, pp. 140, Some of the forms may have a height of six to eight feet as measur-
141): "W. G. Blunt's discovery of fossil teeth of an elephant on ed at the shoulder. The larger individuals are perhaps comparable
Santa Rosa Island, one of the Channel Islands off the coast of in size American mastodon and are certainly smaller,
to the
southern California, was recorded by Stearns [Footnote: 'Stearns, possibly considerably smaller, than the Pleistocene mammoths of
R. E. C., Proc. Calif. Acad. Sci., Vol. 5, p. 152, 1873.'] in 1873. the southwestern United States. While tlie Santa Rosa Island
Since that time this interesting and significant occurrence has been elephant has been determined as representing the species Elephas
referred to by several authors. Hay [Footnote: 'Hay, O. P., primigcniiis Blumenbach and E. (Archidiskodon) imperator Leidy,
Carnegie Inst. Wash. Pub. 322B, pp. 42, 43 and 51, 1927.'] has the difference in size, coui)led with differences noted in the skull and
1032 OSBORN: THE PROBOSCIDEA

dentition, seem quite clearly to distinguish the island form as Conclusions (pp. 212-214).
—
"Whether or not more than one
a distinct species for which the name Elephas exilis is here pro- species of elephant is present among the island forms remains to be

posed." definitely determined. In this connection, it should be recognized

Supplementary Description (Stock, 1935.1, pp. 206, 207, that an interesting and perhaps significant difference may exist

212-214). —
"Remains of extinct elephants are now known to between forms on Santa Rosa and the types of San Miguel."
tho.sc

occur on three of the Channel Islands, namely, on San Miguel, "Numerous cheek-teeth and tusks, fragmentary jaws and
Santa Rosa and Santa Cruz (see Fig. 1 = Fig. 922 of present [
skeletal elements comprise the bulk of the collections obtained on

Memoir]). The first material was found on Santa Rosa more than Santa Rosa. Individuals of all ages are preserved, from an unborn
sixty years ago, and this island has furnished by far the largest type to fully grown adults. The youngest specimen, evidently
collection of fossil specimens representing the.se types. Similar belonging to a foetus, is represented by a lower jaw (Fig. 6 [not
material has been brought to light on San Miguel. In contrast to figured in present Memoir]) in which the enamel plates had not firm-
the rather numerous finds of elephant remains in Quaternary de- ly consolidated to form the lower cheek-teeth and had not erupted
l)osits of Santa Rosa, the presence of elephants on Santa Cruz is through the gums. One fairly complete skull repre.sents an adult in-
known thus far by only two fragmentary enamel plates of a cheek- dividual and furnishes valuable information as to the specific char-
tooth." acters of the island elephants. Wlien found in Quaternary strata,
"San Miguel: Although this island is wind-swept and shifting exposed in the sea-cliff near the mouth of the Caiiada Corral, only
sand dunes mantle much of the area underlain by sediments of the weathered cranial portion was visible. Excavation revealed
. . .

Tertiary and Quaternary age, the incision of the present land sur- the rest of the skull and upper tusks with the lower jaw in
. . .

face by ravines and gullies and the constant though gradual re- position below the palate. [This is the type.] Illustrations of this
specimen and of a young adult skull of the im-
perial mammoth (Airhidiskodon impemlor), drawn
to the same scale, are shown in Figure 9 [not fig-
Scale
ured in ])resent Memoir]."
Submarine depths -n fathoms
"Comparison of fossil remains of elephants
found on Santa Rosa with comparable materials
nta Barbara
occurring on the mainland establishes clearly the
Carpinterja
fact that the island forms were smaller in stature
than their relatives of the mainland. Consider-
Ventura
able variation in size exists among the island types,
but the difference in stature between island and
mainland forms remains a notable feature. . . .

While the elephants of the mainland ranged in

height from a])i)roximately IO/2 feet to 13/2 feet
as measured at the shoulders, those of the islands
presumably never exceeded 8 or 9 feet in height
and the smaller individuals were probably no
taller than 6 feet. Thus, the smaller size of these
elephants presents a character wherein they re-
semble the fossil or subfossil, dwarfed elephants
described from the Maltese Islands of the Medi-
terranean. The diminution in size, however, has
"Map of coastal province of southern California in vicinity of .Santa Barbara.
not been carried so far in the Channel Island
Fig. 922.
I><)iation of some occurrences of fossil elephants on Channel Islands shown by x. Dotted elephants as in the Maltese species."
line indicates hypothetical border of land during Pleistocene time, after Chancy and Mason." "As mentioned before, the elephants of San
Reproduced from Stock, 193.J.1, p. 206, fig. 1.
Miguel are among the largest types to be recorded
from the island region. Tusks of these forms have
been found which measure 5 feet in length and
cession of the ,sea-cliffs develop exposures on which occasionally the 6 inches in diameter at the base. While some of the fossil materials
weathered-out materials of fossil mammals have been discovered. on Santa Rosa likewise indicate the former presence of relatively
Several tusks and cheek-teeth of elephants were found in a thin large individuals, it is possible that the average size of the San
series of (Quaternary alluvial deposits lying beneath a table-like Miguel elephants was larger than that of the Santa Rosa types.
surface and exposed in the sides of gullies near the northwest end
Were this ultimately established to be the case, on the basis of
of San Miguel. Scattered proboscidean teeth have been found
a comparison with more extensive collections than are now avail-
from time to time elsewhere on this island. Among tlie fossil
materials are specimens which clearly point to the fact that the
able from San Miguel, it is interesting to speculate whether the

San Miguel elephants are among the largest types to be obtained difference may not have been the result of an earlier extinction of
in the insular region." elephants on the smaller of the two islands."
 :

THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODON 1033

Archidiskodon sonoriensis Osborn, 1929 Archidiskodon meridionaHs nebrascensis Osborn, 1932

Figure 923 Figures 815, 924, 927, 928, 1239, PI. xxi

One mile east of Arizpe, northern Sonora, Mexico. Lower Pleistocene. One mile northwest of Angus, Nuckolls County, Ncbrask.a. Lower to
Middle Pleistocene.
Archidiskodnn sonoriensis Osborn, 1929. "New Eurasiatic and
American Proboscideans," Amer. Mus. Novitates, No. 393, Dec. Archidiskodon meridionnlis nebrascensis Osborn, 1932. "The
24, 1929, p. 18. Type.— "Nearly complete skeleton,' of 'Elephas meridionalis' Stage Arrives in America," Proc. Colo.
which the palate with third superior molar, M', of both sides, Mus. Nat. Hist., XI, No. 1, Sept. 7, 1932, pp. 1-3 (Osborn,
right lower jaw (lacking ascending ramus), with third inferior 1932.893). Type. —Skeleton, lacking cranium also tusks, ex-
molar, r.Ms, in situ, also .symphysis, are in the American Museum." cepting mid-portion of tusk, with lower jaw
left comjilete state in

Amer. Mus. 22637 (Osborn, 1929.797). Horizon and of preservation. Colo. Mus. 1359. Horizon and Locality. —
Locality. —
"One mile east of Arizpe, northern Sonora, Mexico, on Found "one mile northwest of Angus, Nuckolls County, Nebras-
the Sonora River, 60 miles southeast of Cananea and approximate- ka, some fourteen or fifteen years ago." Lugn and Schultz (1934.1,

ly 100 miles north of La Prietas and San Jose de Pimas. . . . The Table A) regard this species as of Yarmouth (Upland) age, equiv-

Fig. 923. Archidiskodon sonorietisis, anterior portion of type mandible and maxilla showing r.Ms, rM^ (Amer. Mus.
22637), one-sixth natural .size. Compare O.sborn, 1929.797, p. 18, fig. 18.

Arizpe horizon is regarded by Barnum Brown as Lower Pleistocene alent to Lower to Middle Pleistocene. Type Figure. —
(lake deposit)." Type Figure. —Op. cii., p. 18, fig. 18. Osborn, 1932.893, figs. 1 and 2.

Specific Characters. — "Mandibular rostrum prolonged Characters. — "The inferior grinding teeth similar in charac-
obliquely downwards, with downturned beak, as seen both in ter and in ridge formula to the 'Elephas meridionalis' of Durfort,
front and from symphysial groove to tip of
side views; length but somewhat broader with much thicker surrounding layer of
rostrum 230 mm., exposed length of M' 246 mm., of Ms 346 mm.; cement. . . . Mandible: (1) A very prominent rostrum. (2) A rel-

depth from third unbroken plate to bottom of jaw 244 mm. A atively elongate and shallow ramus. (3) Measurements as follows

total of ll + 2(?) exposed ridge-plates in M=, of 2(?) + lH-3 in Length mandibular condyle to symphysis 943 mm.
Mo." Depth below M3 of mandibular ramus 220 mm.
'[Remainder of skeleton unintentionally destroyed by discoverer. — Editor.]
Fig. 924. Type Mandible of Archidiskodon meridionalis nebrascensis (Colo. Mns. 1359). Both figures one-fifth natukal size

(UpiM'r) Cro-.vn view of mandihlc with nglit and left third molurs in silu. After Osborn, 1932.893, fifj. 1.

(Lower) Lateral view of mandible and ro.struin. -Vfter Osborn, 1932.893, fig. 2.

1031
 THE MAMMONTIN.E: ARCHIDISKODON AND METARCHIDISKODON 1035

Drawn from cast Amer Mas 2!Q95 I.pnfjth of r.M3, Third Inferior Mohir 289 mm.
Width of r.M3, Third Inferior Molar 84 mm.
R.M3 total ridge-erest.s + 13 +
L.M3 total ridge-crests 'a 13 }i

Ridge-crests in 10 cm b]i

Widest ridge-crest 73 mm.

The above ridge-crest formula, together w ith the dimensions of the

thin! grinders, agree closely with those which ]irevail among most
yARCHIOISKOljON MERI0I0NALI5 Ref. of DurfcrT France
of the specimens referred to 'Elephas tneridionalis' in the British
Museum as described and figured by Falconer."

"Unfortunately, the cranium was the first part of this animal

to be exposed and was completely weathered out. Only the e.\-
tremity of one of the superior incisive tusks remained; this was
lost; a mid-portion of the left incisive tusk remains. Fortunately,
every was preserved in absolutely com-
othei' part of the skeleton
plete condition on one side of the animal or on both sides so that
the skeleton is now superbly mounted and becomes a classic in
Drawn from cast Amer Mu'^ 21394 all its dimensions as follows:

Fig. 925. Referred superior and inferior molar.s found as.sooiated witli Vertebral column:
Durfort skeleton (Archidiskodon meridionalis) Mu.seum d'Histoire
in the
Nafurelle, Paris, after casts kindly furnished the present author by Dr. 7 cervical vertebrae measuring 543 mm.
Marcellin Boule in January, 1930. Both figures one-third natural size. 19 dorsal vertebrae measuring 1640 mm.
(rp|)cr) Right second superior molar, r.M- (cast Amer. Mus. 2189')), 4 hunbar vertebrae measuring 400 mm.
with +8 worn ridge-plates; ^Yi ridge-plates in 10 cm.; coronal surfa(^e length
Sacrals not preserved.
162 mm., maximum breadth 81 mm.
7 caudals only preserved.
(Lower) Summit of crown of left third inferior molar, I.M3 (cast Amer.
Mus. 21894), with HM ridge-plates, 6 partly worn; o anterior ridge-plates Height dorsal spine to ground (as
in 10 cm. ; maximum length 276 mm., maximum breadth 83 mm. mounted) 3695 mm.

R.M-

L.M ARCHIDISKODON MERIDIONALIS ffcf of Hur/jrr Trance

J)T-ah'n from cast Amer A/us 2/89/
ARCMIOlSKOOOh MERIDIONAUS ^e/ of J,urfo»^ rr^,^ L.M '/^ /Vafural si'73
Mrot^n ^rom cast AmerMuj 2/gs/

Fig. 926. Superior and inferior molars of Durfort skeleton {Archidiskodon mmdionalis) in the Museum d'Histoire Naturellc, Pari.s, after ca.sts kindly
furnished the present author by Dr. Marcellin Boule in April, 1930. All figures one-third natural .size. Compare with figure
924, showing type mandible (if
.4. meridionalis nebrascensis of Nebraska, with third inferior molars in situ.

(Ix'ft) Left and right second and third superior molars (M-"'), after cast (Amer. Mus. 21891).
(Right) Right second and third inferior molars (r.M2.3), same as opposite figure, and left .second and lliird inferior molars (l.M-.-a), after cast (Amer Mus
21891).
1036 OSBORN: THE PROBOSCIDEA
Fore and Hind Limbs:
Height fore-limb scapula to ground 3454 mm. Pelvis, length of os innominatum 1350 mm.
Scapula, length of 1020 mm. Pelvis, width of os innominatum 1754 mm.
Humerus, length of 1220 mm. Femur, length of 1390 mm.
Ulna, length of 910 mm. Tibia, articular length of 840 mm.
Radius, length of 980 mm. Pes: astragalus to tip of Mts. Ill 475 mm.
jMedian metacari)us HI 200 mm. Pes: depth of Mts. Ill 150 E."

Fig. 927. Restoration of Archidiskodon meridionalis nehrascensi.i Osborn, 1932, from the complete skeleton and m.andible with lower portion of the tu.sk.s,

lacking only the cranium. One-fiftieth natural size.

The grinding teeth of thi.s |>riceless specimen are in the .same stage of evolution as those of the famous 'FAeplmn' niiridionalis of Durfort, France, a.s

established by casts of the Durfort grinding teeth kindly sent to the American Museum by JJirector Marcellin Boule (see I'^igs. 92."), 92()).
 THE MAMMONTIN^: ARCHIDISKODON AND METARCHIDISKODOX 1037

TABLE XI. Measurements of Molars of Archidiskodon meridionalis of Durfort (cf. Figs. 925,926, also Fig. 924,
Mandible and Third Molars of A. meridionalis nebrascensis)

Cast Amer. Mus. 21891

Associated molars
Cast Amer.Mus. 21894
Cast Amer. Mus. 21895
 —

Chapter XVII

THE GENUS PARELEPHAS (SUPERFAMILY ELEPHANTOIDEA). OF THE

SUBFAMILY MAMMONTINyE, INTERMEDIATE BETWEEN ARCHIDISKODON
AND MAMMONTEUS, DISTRIBUTED IN THE NORTH TEMPERATE ZONE
OF EURASIA AND NORTH AMERICA
Profound cranial and incisive tusk resemblances to archidiskodon and mammonteus. Clear dis-
tinctions FROM these genera IN GRINDING TOOTH STRUCTURE. FiRST APPEARANCE IN THE UPPER PLIOCENE^
AND LOWER PLEISTOCENE OF EUROPE. MIGRATION THROUGH ASIA MINOR AND ASIA TO THE UNITED STATES
WASHINGTON, NEBRASKA, TEXAS, ILLINOIS, INDIANA, OHIO, SOUTH CAROLINA, GEORGIA, FLORIDA AND FRENCH —
GUIANA, SOUTH AMERICA. EXTINCTION IN UPPER PLEISTOCENE OR P0STGLACIAL(?) TIMES.

1. European north temperate origin. History of separation from 3. North and South American species of Parelephas.
other extinct proboscideans. Parelephas jacksoni, Ohio, e.xact locality unrecorded.
Complete separation by Osborn of the phylum Parelephas. ParelephasC^) mississippiensis(1), Indiana.
Order of discovery and description of species of Parelephas.
Parelephas culumbi, Georgia.
Phylogenetic order of succession of Parelephas.
Distinctive cranial characters of Parelephas. Parelephas columbi felicis, Mexico.
2. Systematic description of European and Asiatic species in Parelephas columbi cayennensis, French Guiana,
ascending progressive order. South America.
Parelephas Irogonlherioides, Italy. Parelephas jeffersonii, Indiana.
Parelephas trogontherii, Germany. Elephas roosevelti (synonym of P. jeffersonii), Illinois.
Parelephasil) Irogonlherii neslii, England.
Parelephas progressus, Ohio.
Parelephas armeniacus, Asia Minor.
Parelephas ivashinglonii, Washington.
Parelephas unrecorded in China and Japan.
Parelephas intermedins, France. Parelephas eellsi, Washington.
Parelephas wiisli, South Russia. Parelephas floridanus, Florida.

1. EUROPEAN NORTH TEMPERATE ORIGIN. HISTORY OF SEPARATION FROM OTHER

EXTINCT PROBOSCIDEANS
The generic i)hylum separated as Parelephas by Osborn in the year 1924 is briefly mentioned in Chapter II of

the present Memoir and distinguished especially in its cranial characters in Chapter XV, in which it appears
that Parelephas is linked with Archidiskodon and Mammonteus in its cranial resemblances and great incurved
incisive tusks, while in its grinding teeth and ridge-plate formulae it is so nearly intermediate between these two
genera as to have been mistaken for an actual connecting link. In the present chapter it is shown to be an entirely
distinct genericphylum which during the more temperate interglacial periods (Fig. 795) occupied the same geo-
graphic range as that of the true woolly mammoth (Mammonteus) during the glacial periods.

History. — The sixteen species which are grouped in the genus Parelephas constitute a very ancient and
distinct generic phylum for which Osborn's name {Parelephas) seems appropriate, in reference to the convergence
or parallelism of the grinding teeth in this phylum with those of the true Elephas. For more than half a century,
owing to the similarity in appearance of the grinding teeth to those of the true mammoth (Mammonteus), all

English and American authors, including Falconer, Leith Adams, and Lydekker, confused the grinding teeth of
specimens which Osborn now refers to Parelephas with those of the northern mammoth Elephas [Mammonteus]
primigenius, and this accounts for the great discrepancies in the collective ridge formulae attributed to the species
E. primigenius, i.e., M 3 \^ as given by Leith Adams (1877-1881, p. 127) and by Lydekker (1886.2, p. 175), and
copied by Hay (1914, p. 395). This unfortunate confusion in the ridge formula arose despite the fact that Falconer
'[See footnote 1 on page 1049 below. — Editor.)

1039
1040 OSBORN: THE PROBOSC'IDEA

had correctly defined the ridge-plate formula of E. primigenius as

as early as 1863 (p. 65) 3 ff, a formula which M
is fully confirmed by Osborn's researches for this Memoir (see Chapter XVIII). The lower ridge-plate count
(M 3 tI) attributed by Adams and Lydekker to Elephas primigenius really belongs to grinding teeth of primitive
Middle Pleistocene species of Parelephas, such as P. trogonlherii, which exhibits 3 g+ ridge-plates. As described M J

in detail below, Jourdan (1861) was the first to separate from E. primigenius one of these species of Parelephas

under the name Elephas intermedius, a stage which exhibits M3 |f ridge-plates; he was followed by Pohlig (1885)
who clearly defined the more primitive Elephas trogontherii.

I-
it

i-

Restoration of the Type of Parelephas jeffersonii

One-thirtieth natural size

Fig. 9.30. This painting hy Charles R. Knight, in the year 1909, was taken direetly from the type skeleton of Parelephas jeffemonii in the
American Museum of Natural History. The characters of the typical /'a;f/cp/i(i,s cranium and tusks are particularly well shown, with short concave
foreiiead an<l prominent convex occipitofrontal ere.st. The sli.ape of the ears i.s entirely conjectural. The hairy covering, unlike the hairy and
woolly covering of Mainmoidius /jriiiiigeuius, is a wholly conjectural character, because no remains of the iiair have been discovered, hut the presence
of this Jeffersonian mammoth in north temperate regions, appearing in post(?)-Wisconsin times, furnishes indirect evidence of a hairy if not of
a woolly coat.

In the present chapter, chiefly from the researches of Deperet, Mayet, and Osborn, it is shown that the
phylum Parelephas constitutes a long line of progressive ascent wholly distinct from that of Mammonteus primigenius.
Its first appearance is in Upper Phocene time in Italy. Its final appearance in IV Glacial and Postglacial times,
principally on or near the 40th parallel of the United States, in the species Parelephas jeffersonii (M 3 H), is

followed by the closing stage Parelephas progressus (M 3 M).

 THE MAMMONTINiE: PARELEPHAS 1041

It has also been a long and difficult matter both in Europe and America to clearly separate the members of
the generic phylum Parelephas from the newer phylum of the broad-toothed, narrow-plated true mammoth
(E. = Mammonteus] primigenius) on the one hand, and from the older phylum of the broad-toothed, broad-plated
[

southern {Elephas meridionalis) and imperial {E. imperator) mammoths of the genus Archidiskodon on the other.
The chief grounds of separation are as follows: (1) The cranium in Elephas intermedius, E. irogontherii, and E.
jeffersonii is now known to be readily distinguishable from the crania of either E. = Mammonteus] primigenius or [

E. = Archidiskodon] imperator; (2) the grinding teeth are intermediate in form and in the number of plates, as
[

Second Figure of the Type Skeleton of Parelephas jeffersonii

One-thirtieth natural size

Fig. 931. Second figure of the aged type skeleton of Parelephas jeffersonii Osborn, 1922, p. 11, fig. 10, as mounted in the American Museum
(Amer. Mus. 9950). For further information about this type skeleton, see legend of figure 966, also the description below.

observed by Jourdan in 1861 in applying the name Elephas intermedius; (3) it is noteworthy that both in France
and Germany the grinding teeth of Parelephas have been independently described as intermediate in structure
between those of E. [
= Archidiskodon] meridionalis and E. [=Mammonteus] primigenius, as shown in the full

historic notes below. Elephas columbi proves to be Parelephas columbi.

France: Jourdan, 1861. — Despite the early separation by Jourdan of Elephas intermedius from E. primi-
genius and description of the grinding teeth as intermediate between E. primigenius and E. meridionalis (1861),
1042 OSBORN: THE PROBOSCIDEA

Gaudry, who was very conservative in the matter of applying new names to species, continued to describe (1876, p.
40, also PI. ix) these animals as "Elephas primigenius a lames ecartees." The steps in the gradual separation of
this phylum by palaeontologists of France are clearly described by Deperet and Mayet (1923, pp. 176-190).

Fig. 032. Chief Lower to Upper Pleistocene localities in wliieh occur species of
Archidiskodon, Parelephas, Mammonteus, Loxodonla, and Palseoloxodon (syn. Pilgrimia),
Italy: Falconer, 1868. —As early as 1868
after Osborn, 1910.346, p. 391, fig. 176. (see below under Parelephas armeniacus) , Fal-
Fossils attributed to the ijhylum Parclephas, and of large size, are especially abun-
coner entertained a strong suspicion that a
dant Middle Pleistocene
in the of Siis-senborn (Fig. 932,11), of Mosbaeh (12), and of
Taubach (19). At Mosbaeh (in Isl! Inlcrghcial times) they occur in the same layers, form closely related to E. armeniacus occurred
but with much greater frequency, than those of E. [Hesperoloxodon] antiquum, namely,
three to one (Soergel, 1912, p. 32); more recently it is estimated (Schmidtgen, 1926)
at St. Paolo, Italy, which he identified (1868,
that in the Mosbaeh sands E. [Parclephas] trogonthtrii is about ten times as abundant as II, pp. 249, 250) as resembling E. armeniacus,
E. [Hesperoloxodon] antiquus; in Mosbaeh also this elephant attains its greatest size.
but which we now know belonged to Parelephas
irogontherioides (see p. 1055). This is a fine ex-
ample of Falconer's unerring sense of form.

Germany: Pohlig, 1885.— In 1885 Pohlig,

on discovering two grinding teeth of one of
these mammoths in the Interglacial sands of

SUssenborn near Weimar, proposed the specific

name Elephas irogontherii; he maintained that

both geologically and zoologically this species
was a link, or intermediate, between Elephas
primigenius [i.e., Mammonteus] and Elephas
meridionalis [i.e., Archidiskodon]. To quote
Pohhg's own language {op. cit., 1885, p. 1027):
"Unter der Bezeichnung 'Elephas Irogontherii

Pohl.' fUhre ich in meiner Monographic eine

PLBISTOCBNX —
EUROPE. 1 Forrst Bed of Cromer (Norfolk). Sables de 2 europaische Molarenform auf, welche zwischen
St. Prest near Chiirtres (Eure-et- Loire). 3 Maihnitu (Puy-de-Dorae). 4 Peyrolles (Bouches-
du-Rhonc). 5 .Holhilac near Puy. Clay deposits of 6 Uurforl (Card). 7 Cnjarc (Lot-et-Ga- denjenigen des E. primigenius und E. meridi-
ronne). 8 Vol d'.irno (Tuscany). 9 Leffe near Bergamo (Lombardy). 10 Rixtlor/ near Pots-
dam (Brandenburg). Gravels of 11 SUssenborn near Weimar. Sands of 12 Mosbaeh in onalis zoologisch, wie ihrer geologischen Lager-
northern Baden. Freshwater deposits of 13 CUicton (Essex). Sands of Mauer near 14 Hei-
delberg (western Germany). 15 Chelles on the Mame, near Paris. 16 SI. Acheul (Sonime). stiitte nach, in der Mitte steht."
17 Il/ord and Grays Thurrock (Essex). Lignites of 18 Diimlen and of Utznach, near Zurich.
19 Tauhach near Weimar. 20 WtWirrcWi cooe on A/on( 5an(is (eastern Switzerland). Tuffs
of 21 the Tiber Valley, near Rome. Caves of 22 Neandertal, near Dusseldorf (western Ger- Pohlig also observed the relationship of E.
many), 23 .Spy. near .\mur (Belgium), 23a Krapina (Croatia), 24 Chapelle-aux-Soints (Cor-
r^zc). Caves and alluvial deposits of 25 Temifine (or Palikao) near Gran (Algeria). 26 Pointe Irogontherii to the species which Falconer named
Peacade, near Algiers (Algeria). 27 Prince's Cave (Monaco). Sandy clays of 28 Vdklinshofen
(Alsace). 29 Saalfcld (.Saxe-.Meiningen). Travertines, etc., of 30 Gera, Jena (.Saxe-Weimar). Elephas armeniacus in 1857. At the same time
31 L«ipii'fl (Saxony). 38 .So/u(rc, north of Lyons. Loess of 33 Wurjfcurff (Bavaria). H
Thicde
Pohlig erred in suggesting the relationship of E.
near Braunschweig (Prussia). Cave of 35 Montmaurin (Haute-Garonne). 36 Chdteauneuf-
sur-Chartnle (Charente). Caves of 37 .Schweizershild near Schaffhausen, and Kesslerloch near
Thayngen (northern Switzerland). Remains of lake dwellings at 38 Wauwyl (Lucerne), 39 Ro- trogonlherii to Elephas [Palxoloxodon] natnadicus
benhausen, south of Lake Pfaffikon. 40 Concise on Lake Neuchatcl (Switzerland). Peatbogs of
41 Hassleben. near Weimar. Travertines of 42 Langenaalza (Erfurt) in central Germany. Falc. ; he also erred in suggesting that both in
Caves of the 43 Island ol Malta, 44 Island of Crete, 45 Island of Cyprus.
craniology and dentition there was a direct
INot on map: 46 San Paolo Je Villafranca (Piedmont), Italy, 47 Eizcruni, .\rnienia. 48 Plateau
loess.Lyon, France. 49 Tira.spol, .S. Kuasia)
phylogenetic or ancestral succession between
E. meridionalis, E. Irogontherii, and E. primigenius (i.e., "directer Verwandtschaft").

"Under the name of Elephas trogontherii, Pohlig, I have described European

In 1886, p. 181, Pohlig remarks:
molars which hold a middle place, both zoologically and geologically, between those of E. primigenius and E.
meridionalis, most closely apprf)ucliiiig those of PJ. antiquus in the ridge-fonnula, but differing more from them than
from the other two in the form of the crown. The position of E. Irogontherii with regard to E. armeniacus, Falc,
and E. namadicus, Falc, still remains to be investigated."
 : .

THE MAMMONTIN^: PARELEPHAS 1043

France: Deperet, 1923. — In 1923, Deperet and Mayet in their invaluable review of "Les Elephants
Pliocenes," Deuxieme Partie, out of respect for Pohlig's clear distinction and definition of the species E. trogon-
therii, decided not to adopt the term Elephas intermedius of Jourdan, but to name this phylum "Rameau de
VElephas They
trogontJierii." recognized its remote kinship to Elephas primigenius (see "III Groupc des Elephas

irogontherii et E. primigenius (Mammouths)") but established it (p. 176) as a distinct branch of the "groupe des

Mammouths." Osborn believes this to be its true phyletic position, resting on two chief characters, namely:
(a) The intermediate character of the grinding teeth, observed by Jourdan, by Gaudry, by Pohlig, and by Deperet

and Mayet; (b) the dome-shaped cranium, figured and observed by Pohlig and by Deperet and Mayet.
In their analytical treatment of the geologic succession of species, Deperet and Mayet clearly pointed out that
the type locality {j3d Interglacial plateau loess near Lyons) of Elephas intermedius Jourd. is of more recent geologic
age than the type locality {Sd rnterglacial sands) of Elephas trogontherii Pohl.; moreover in seeking a Pliocene
ancestor of this phylum, they believe they have discovered it in the Upper Pliocene subspecies of Italy, named by
Zuffardi in 1913 Elephas antiquus var. frogontherioides. They thereby confirmed Falconer's observation of 1868
that an elephant closely similar to E. armcniacus occurred in northern Italy (see Parelephas armeniacus descrip-
tion, p. 1060 below).

Consequently the actual geologic succession of the three types of Parelephas thus far discovered in France and
Italy is as follows

Pleistocene {Sd Inkrglaciul plateau loess), Lyons, France Elephus intermedius Jourdan, type
Pleistocene (2d Interglacial sands), Siissenborn near Weimar, (lermany Elephas trogontherii Polilig, type
Upper Pliocene (Villafranchian), Italy [see footnote, p. 1049 below. Ed.] — Elephas antiquus var. trogontherioides Zuffardi, type

America: Osborn, 1922. —Among the abundant remains of members of thisphylum in America, a precisely
similar confusion between specimens belonging to Archidiskodon, Parelephas, and Mammonleus arose in the minds
of American palaeontologists (Cope, Osborn, and Hay). Cope (1889.2, pp. 208, 209) referred the remains of a very
fine skull of Archidiskodon (Fig. 891) from Texas to "Elephas primigenius columbi Falc." Osborn also saw only
resemblances to "Elephas columbi" in the fine skeleton found in Indiana (type of Parelephas jeffersonii Osborn,
Figs. 961, 931); whereas Hay (1914) referred the same skeleton to Elephas primigenius. No stronger proof
could be given of the truly intermediate character of members of this phylum in America than this alternate
reference of the same skeleton by Osborn to Elephas columbi and by Hay to Elephas primigenius. In Hay's
exhaustive summary (1914) of grinding teeth and other remains from various parts of the United States (Alaska
to Florida) which he referred to "Elephas columbi," he includes teeth which certainly belong to Parelephas columbi
as well as to the truly intermediate form (P. jeffersonii)

Osborn (1922.555) was the first to separate these intermediate animals, previously described by Cope, Hay,
and himself as "E. columbi" and "E. primigenius," under the new specific name of Elephas jeffersonii, a species

which he subsequently (1924.633, p. 4) made the genotype of Parelephas. This separation was based principally
upon cranial characters (Osborn, 1922.555, p. 15): "Still more obvious are the differences between the relatively
long, broad, and shallow crania of E. jeffersonii and the relatively short, narrow, and deep crania of E. primigenius^
proportions which are correlated respectively with the corresponding proportions just described and figured in
the teeth."

Phylogenetic Relationships (Pohlig, Soergel). — Inasmuch as the same species of animal had previously
been erroneously described by Cope and Osborn under the name of "Elephas columbi," we must credit Soergel
(1921, p. 60) with the prior observation that this supposed "Elephas columbi" of America { = Elephas [Parelephas]
jeffersonii Osborn) shows phyletic relationships to the Elephas [
= Parelephas] trogontherii of Pohlig.
1044 OSBORN: THE PROBOSCIDEA

Schmidtgen and Freudenberg (1926, pp. 62, 68) describe and discuss Elephas [Parelephas] Irogontherii of
Mosbacli, especially the supposed phylum ("Stammreihe") Elephas meridionalis E. Irogontherii E. primi- — —
genius, which in unbroken succession lived in the region of Wiesbaden.
Osborn, 1924-1928: It has remained, however, for Osborn in the present Memoir to institute a close compari-

son between all the known characters of the Elephas Irogontherii phylum of Europe and the Elephas jeffersonii of

America and to establish the fact that there is a close phyletic relationship which justifies the linking of the
European and American species in the new and distinct genus Parelephas. This aflfinity is most clearly indicated
in the cranium, as shown in the resemblance of several crania, erroneously figured as "Elephas primigenius" by
Falconer, to those of E. trogontherii as figured by Pohlig, and to the cranium of E. jeffersonii as figured in the

present Memoir; there can be no doubt that this relatively broad, elongate, and rounded cranium is entirely
distinct from the extremely short, compressed, and peaked cranium of the true mammoth Elephas primigenius.
Habitat of Parelp:phas trogontherii (Soergel, 1912). In his masterly monograph of 1912, entitled —
"Elephas trogontherii Pohl. und Elephas antiquus Falc," Soergel observes (pp. 105, 106) that the Steppe [plains]
and \^'ald [forest] faunas lived contemporaneously in central Europe; as observed in the Id Interglacial period,
they are found in the lower sands of Mosbach, the "kiese" [gravels] of Mauer, the sands of Petersdorf bei Gleiwitz
in Schlesien, the "Tone" [clays] and "grauen Rheinsande" [gray sands of the Rhine] near Jockgrim in Pfalz, and

the gravels of Sussenborn (which persist into II Glacial time).' He divides the plains or "Steppe" and forest
or "Wald" faunas of the 1st Interglacial period as follows:

Chiefly Inland Steppes or Plains Chiefly Ocean Borders and Forests

E. [Parelephas] Irogontherii Pohl. E. [Hesperoloxodon] anliquus Falc.
Very abundant in Mosbach; absent in Mauer. Very abundant in Mauer.
Dicerorhinus etruscus F'alc. Dicerorhinus etruscus Falc.
Equus stenonis Cocchi, E. siissenbornensis Wlist, Equus stenonis Cocchi, E. mosbachensis v. Reich.
E. mosba£hcnsis v. Reich.
Leptobos etruscus Falc.
Bison priscus Boj. Bison priscus Boj.
Cervus elaphus Linn., C. elaphus trogontherii Pohl., Cervus elaphus antiqui Pohl., C. capreolus Linn.,
C. capreolus Linn., Alces lalifrons .lohns. Aloes lalifrons .Johns.
Ursus arvernensis Croiz. and Job., U. Deningeri v. Ursus arvernensis Croiz. and Job., [/. Deningeri v.
Reich. Reich.
Felis lea fossilis Goldf. Felis leo fossilis Goldf.
Hysena arvernensis Croiz. and Job.
Canis neschersensis Croiz. and Job. Canis neschcrsensis Croiz. and Job.
[Homo heidelbergensis Schoet. (lower sands of Mauer)].

Certainly Hesperoloxodon antiquus, a forest loving form with coarser teeth, occurred nearer the ocean borders,
whereas Parelephas trogontherii, with finer teeth, frequented both the steppes (plains) and forests. Since animals
of both habitats are buried together in river sands, gravels, and clays, there is an intermingling of plains and forest
faunas, as shown in Mosbach and Mauer (Osborn).
Of the habitat and geographic distribution, Soergel {op. cit., p. 110) observes that whereas Hesperoloxodon
antiquus preferred a warmer ocean bordering climate, not under the direct influence of the northern inland ice
masses, Parelephas trogontherii sought the cooler northern-northeastern continental localities. Accordingly in
Italy, Spain, and Greece it is almost entirely wanting; in France P. trogo7itherii is less abundant than H. antiquus,
at least, it appears in less characteristic forms. The homeland of the P. Irogontherii type appears to be confined to
England and Germany, perhaps al.so to Russia. During the arid 1st Interglacial peroid we find P. Irogontherii

distributed from southern England over middle Germany and eastward to southern Russia, as shown in the
Forest Bed of Cromer, the sands of Mosbach, the gravels of SUssenborn, the sands of Petersdorf in Schlesien, and
the clays of Jockgrim in Pfalz. Already in this Isl Interglacial period migration towards the east may have begun.
'[See p. 1056 below, where Parekphas Irogontherii type is placed by Professor Osborn in the 2d Interglacial. — Editor.]
 : . :

THE MAMMONTINiE: PARELEPHAS 1045

In the 3nd Interglacial period we observe practically the same geographic distribution of Parelephas tro-

yontherii; its southernmost appearance in Germany at this time is at Steinheim on the river Murr; its northern-
most appearance is at Rixdorf near Berlin. In Thuringia it is found in the Ilm gravels below the older travertines
of Taubach, also in the stream gravels at Vieselbach near Erfurt, etc.

For the better understanding of the geologic as well as the geographic distribution of the Parelephas trogon-
therii phylum in Germany may be cited the following geologic correlation by Soergel (op. cit., p. 106)

Dem I. 'Interglazial' gehoren an: Dem II. 'Interglazial' gehoren an: Dem III. Interglazial gehoren an:
die Sande von Mosbach [n. die Bachkiese bei Vieselbach, der Travertin von Taubach-
Baden], die Saalekiese von Uichteritz Ehringsdorf- Weimar,
die Kiese von Mauer [b. Heidel- b. Weissenfels, dcr Tra\ertin von Burg-Cirae-
berg], die Ilmkiese unter dem alteren fentonna,
die Sande von Petersdorf b. Travertin von Taubach- der Travertin von Bilzingslcbcn
Gleiwitz in Schlesien, Ehringsdorf [b. Weimar], und verschiedene andere Tra\--
die Tone und grauen Rhein- die Schotter von Steinheim ertinvorkommen Thiiringcns.
sande b. Jockgrim i. d. (II. Glazial-Interglazial).
Pfalz,
die Kiese von Siissenborn [bei
Weimar], die allerdings die Sande von Rixdorf b. Berlin
bis in die \l. Eiszeit hin- fiir eine Ablagerung des II.
eingehen. 'Interglazial.'
[Plains ("Steppe") and forest and forest faunas not clearly
[Plains [Plains and forest faunas again api^ear
("Wald") faunas, as listed above.) separated Parelephas trogontherii and
; partly separated, as in Mauer and
The Siissenborn deposits extend into Hesperoloxodon antiquus occur in the Mosbach.
H Glacial time {fide Soergel, 1912.) same horizon.] Hesperoloxodoiiantiquussurvives Par-
elephas trogonlherii, which disappears.
In the tundra fauna appears Mam-
monteus pr imige nius.]

Complete Separation by Osborn of the Generic Phylum Parelephas

Osborn finally (1924.633, p. 2) concluded to cut the Gordian knot and terminate this confusion by distinguish-
ing the generic phylum Parelephas throughout from both Mammonteus and Archidiskodon, as shown in the follow-

ing citation

[p. 2] Much more difficult has been the separation of the third generic series of the Mammontinae, which hitherto has been
referred to the genus Elephas but which we now remove to the new generic phylum Parelephas.
[p. 4] The eight or ten species included within this genus constitute a very ancient and wholly distinct generic phylum, for
which the name Parelephas seems appropriate, because in certain characters these animals parallel the true Elephas, although in
profound cranial and dental structure they are closely related to and convergent with the mammoths Archidiskodon and
Mammonteus.
Generic Characters. —A
phylum of the Mammontinae. Elephas jeffersonii, genotypic species, E. armeniacus, E.
intermedins, E. trogontherii, E. trogontherioides. Craniimi intermediate in form between that of Archidiskodon and that of
Mammonteus, namely, brachycephalic, acrocephalic. Frontals concave, occipital crest elevated; occiput more or less convex.
Molars in the Upper Pliocene and Lower Pleistocene stages with relatively few ridge-plates, i.e., 3 -}-f+; progressive Upper M
Pleistocene stages with multiple ridge-plates, i.e., 3 f§. M
Ridge-plates compressed to 7-8-9 in 100 mm. Alolar crowns broad,
RP short, with enamel of intermediate thickness, more or less crimped or sinuous.
The progressive ridge formulae in Parelephas are distinct throughout from those of Mammonteus, and the final ridge formulae
in the two generic phyla are different, namely:
Mammonteus primigenius comprcssus, M
3 Tf = final stage.
Parelephas jeffersonii progressus, M
3 ff = final stage.
The crania of Parelephas throughout are readily distinguishable both in frontal and lateral aspects, and especially in vertical
section, from those of Mammonteus, as can be seen in all of Falconer's beautiful plates of E. primigenius and in Pohlig's excellent
figures of P. trogontherii.
The jaws of Parelephas and of Mammonteus are less readily distinguishable, but by more profound study they can also be
separated from those of Mammonteus. The contrasts in the crania of the two genera may be summed up as follows:
Mammonteus. — Cranium
and jaws extremely compressed fore-and-aft (cyrtocephalic) extremely elevated and pointed ;

above (hypsicephaHc) extremely depressed and foreshortened below (bathycephalic).

;

Parelephas. —
Cranium moderately compressed fore-and-aft (cyrtocephalic); moderately elevated occipitofrontal borders
(acrocephahc) moderately depressed molar-grinding area (bathycephalic)
;

Thus, while the intermediate forms of crania and teeth of Parelephas and of Mammonteus may prove difficult to separate,
the two finally progressive forms are readily separable, namely, Parelephas jeffersonii progressus and Mammonteus primigenius
compressus.
1046 OSBORN: THE PROBOSCIDEA

To this phylum certainly belongs the Elephas armeniacus of northern Asia Minor, as first observed by Fal-
coner, as well as many specimens to the eastward described under other names. The connection of this jihylum

with the Parelephas jeffersonii of late Glacial times in the United States seems highly probable; the reseml)lance
both of the teeth and cranium has been repeatedly observed between E. trogontherii and the northerly tyi)c form-
erly known as "Elephas columbi," now known as Parelephas jeffersonii.

The southerly type (Georgia, South Carolina, Florida) of Elephas [

= Parelephas] culumbl, with rather
primitive ridge formula (M 3 rirp), has recently been reinforced by the large and more progressive Parelephas
Jloridanus 3 ^l), apparently a late Pleistocene stage distinguished by its larger size,
(M more robust tusks, and
coarser ridge-plates from the northerly (Indiana) Parelephas jeffersonii type.

A branch of the Parelephas cohnnbi stage appears to have migrated to South America, as mentioned Ijy Lull

1908, p. 204) and Freudenbcrg (1922, p. 159) on the authority of Lartct (1859, pp. 500, 505).

I.AiriKT, \H'->9. VI'. .'iOO' and 505. —

L'existcncc des Ck'phants n'cst encore indicnit'e dans rAnierifiue nieridionalc ((ue ])ar un
fragment de molaire a lames epaisses, rapporte de Cayenne i)ar le capitaine Ferret et donne par lui au Miisec de Marseille. . . .

Dans rAniprique dii Sud, deux formes si)ecifiques du genre Mastodonte se montrent dans des depots posl-pliocenes; mais ]ieiit-
etre se sont-elles aussi retrouvees dans des formations plus anciennes et raijportables a I'age precedent on pliocene, (^uant au
type Elephant, il n'y est encore indique que par le seul fragment deja cite d'une molaire a lames epaisses, rapports de Cayenne
par le capitaine Ferret.
—
iMiErDENBERC, 1922, PP. 159, 160. Wir haben Lartets Autoritat fiir den Fund eines Zahnes von Elephas im unteren
Fleistociinvon Cayenne (Franzosisch-Cniayana). Nach der Reschreibung der dickriickigen Schmelzplatten ist das Excnijilar
offcnbar von El. impcrator. Wahrscheinlich vollzog sich eine Wanderung nach dem Siiden, bevor die Redingungen geschaffen
«aren. wclche spater die Wanderung der Proboscidier nach Siidamerika ermoglichten. Ks ist das einzige mir bekannt . . .

gewordene Beispiel eines wahren Elefanten im Siiden des Hochlands von Mexiko.

Osborn, 1929: Through the extreme courtesy of Mm. Laurent and Repelin of the Faculte des Sciences de
Mar.seille, this priceless molar fragment from French Guiana has been located in the Museum d' Histoire Naturelle

fie Marseille, photographs taken, and casts made, especially for the purpose of specific and generic identification

for the present Memoir. As shown in figure 957, on a one-half scale, this specimen consists of three ridge-plates
of coarse enamel, as described by Lartet, remotely resembling in crown and side views the posterior ridge-plates

of a superior grinding tooth of Parelephas columbi (Fig. 952). In 1929 the present writer (Osborn, 1929.797, p. 20)
made this the type of a new subspecies, namely, Parelephas columbi cayennensis.

Th(> rcH-entlj- disco\-('red Parelephas Jloridanus includes parts of three full skeletons collected by the Holmes
Florida Expedition of the year 1929. In this stage, intermediate in ridge-plate fornuila (M 3 ff^) between Elephas
[Parelephas\ columbi (M 3 \j+) and P. jeffersonii (M 3 H), the ridge-plates are broad at the base and compressed
at the summit ; the inci.sive tusks extremely massive and relatively short. The males attain very large size. The
femora measure, according to age, 1230 mm. to 1393 mm., as compared with 1250 mm. in the adult Indiana type
of P. jeffersonii.

Summary of Migration
The restudy of long-known specimens like Elephas jacksoni (1838), Elephas columbi (1857), Elephas armen-
iacus (1857), and Elephas intermedius (1861), together with the recent recognition of Parelephas columbi cayen-
nensis in French Guiana, builds up a Parelephas phylogeny and migration second only to that of Archidiskodon
(Chapter XVI). We await the discovery (probably in northern Africa) of an ancestral stage more i)rimitive than
Parelephas trogontherioides of the Uppei- Pliocene- of Italy.

'Dr. Richard S. Lull (letter, Sept. 21, 1928) wrilt's: "rruin this (icworiptiori, inentioiiinE the fragment of ji molar with 'thick plates,' I drew my own
conclusion as to the genus and species of that elephant. It seemed to me that that it would be Elephas —Archidiskodon] iiuj/cralor."
[

-[See footnote 1 on page 1019 below.— Editor.)

 THE MAMMONTIN^: PARELEPHAS 1047

Fig. 933. Geographic distribution (aecording to tlie numbers in the accompanying hst) of species of Parelephas. The white dots within the black
areas represent the approximate localities where the types of these species were discovered. The white crosses represent referred specimens. It will be
noted that Professor Osborn regarded EUpkas texianus Owen, Blake (1859-1861) as a synonym of Parelephas columbi, and Elephas roosevelli Hay (1922) as
a synonym of Parelephas jeffcrsonii.

ORDER OF DISCOVERY AND DESCRIPTION OF SPECIES OF PARELEPHAS

See Figure 933

1838 1. Jackson County, Ohio

1857-1868 2. Brunswick Canal, Georgia
1857 3. Armenia, Asia Minor
1859-1861 4. San Felipe de Austen on Brazos
River, Texas
1861 5. France, 3d Interglacial loess
1872 6. Indiana

1872

1885

1891

1910
1048 OSBORN: THE PROBOSCIDEA

Sui>erfamily: ELEPHANTOIDEA Osborn, 1921

Family: ELEPHANTID^ Gray, 1821

Subfamily: Mammontin^e Osborn, 1921
Genus: PARELEPHAS Osborn, 1924
Original reference: Osborn, Amer. Mus. Novitates, No. 152, pp. 4 and 5 (Osborn, 1924.633).

Generic Characters (cf. Osborn, 1924.633, p. 4). — Genotypic species, Elephan jeffersonii Osborn,
of Indiana; species referred to this genus, Elephas armeniacus, E. intermedius, E. trogonlherii, and E.
trogontherioides. Cranium intermediate in form between that of Archidiskodon and that of Mammonteus,
namely, bathycephalic, acrocephalic. Frontals concave, occipital crest elevated; occiput more or less
convex. in the Upper Pliocene to Middle Pleistocene stages with relatively few ridge-plates,
Molars
i.e., M progressive Upper Pleistocene stages (P. progressus) with multiple ridge-plates, i.e.,
3 14+;
M 3 26. Ridge-plates compressed to 6-6.5-7-8-9 in 100 mm. Molar crowns broad, M^ short, with enamel
of intermediate thickness, more or less crimped or sinuous. Adapted to continental plains or steppe
environment, grazing and browsing.

I Pl)er Pleistocene
 '

THE MAMMONTIN^.: PARELEPHAS 1049

Only an api)roximate phylogenetic order of succession may be given at the present time. In this phylogenetic
list we some doubt the imperfectly known Elephas jacksoni Mather, also the recently described
include with
Elephns washingtonii Osborn. In phylogenetic order it would seem that this Parelephas phylum first appears in
Elephas trogontherioides of the Upper Pliocene (Villafranchian)' of Italy. Doubtless there were many intermediate
forms or ascending mutations between this Pliocene stage and the Mid-Pleistocene Elephas trogontherii of Pohlig
at the close of the 2d Interglacial period when this animal was quite abundant. The type of E. trogontherii of
Germany is about the same stage of evolution of the third upper molar as the type molar of the E.
in fact in

armeniaais of Asia Minor.

The phylum is next represented in the 3rd Interglacial in the plateau loess of southern France in the form of
Elephas intermedins of Jourdan. Deperet and Mayet consider the type of Elephas intermedins as more progressive
than the type of E. trogontherii. Pohlig also states that a form related to E. trogontherii survived into 3rd Inter-
glacial times in Germany and was then replaced by Elephas primigenius; whereas Deperet and Mayet consider
that the Elephas trogontherii-intermedius phylum of Europe became extinct before 3d Interglacial time. This
3d Interglacial period may possibly mark (Fig. 795) its time of migration across Asia into North America.^

Priority of Deperet in separating the E. trogontherii Phylum.— To Charles Deperet and his colleague
Lucien Mayet belongs the credit of clearly distinguishing the origin and geographic distribution of the "Rameau
de V Elephas trogo7itherii," to which phylum Osborn has assigned the generic name Parelephas. Deperet and Mayet
(1923, pp. 176-183) first distinguished the "A. Rameau de VElephas trogontherii Pohlig" from "B. Rameau de
VE. primigenius Blumenbach," both of which they included within the "Groupe des Elephas trogontherii et E.
primigenius (Mammouths)." They credit Jourdan with the recognition of E. intermedins as distinguishable from
E. primigenius, followed by Gaudry (1876) and by Leith Adams (1877) who traced this form back to the Forest
Bed of England, then by Pohlig (1889 [1888]) who described E. trogontherii of Sussenborn near Weimar, but which
Deperet and Mayet remark is absolutely identical with the E. intermedius of Jourdan. Finally they trace the
"Rameau de VElephas trogontherii" down into the Pliocene superieur (Villafranchian of Italy), confirming a note
by Pohlig (1889 [1888], p. 208) as to the tooth subsequently made by Zuffardi (1913, p. 174, Tav. xii [vi], figs. 3a, 36)

the type of Elephas primigenius var. trogontherii. Deperet and Mayet observe (p. 177) that the molars of this
E. trogontherii var. are readily distinguished from those of the contemporary E. [Archidiskodon] meridioncdis
Nesti and E. [Hesperoloxodon] ausonius Major-Deperet. Zuffardi also described and figured another molar under
the name oi Elephas antiquus var. trogontherioides (Zuffardi, Tav. ix [iii], fig. 6), and two other molars of the
E. trogontherii form (PI. ix [in], figs. 4 and 5a, 56). They state (p. 179) that these four molars of E. trogontherii

described from the "Villafranchien de I'Astesan" are the only ones thus far observed in Italian Museums, although
there are probably others, and that they are clearly distinguishable from 'E. meridionalis' and 'E. antiquus,

concluding (p. 180) as follows:

On peut done dire seulement que la forme pliocene de VE. trogontherii possede des molaires de dimensions plutot un peu
rcduites, sans qu'il puisse etre question d'une veritable mutation de petite taille de I'espece. On doit s'attendre a decouvrir un
jour dans le Pliocene ancien de quelque region eloignee les types ancestraux nains du rameau de l'^^. trogontherii.

'[There is a tendency among many Chap. XXII of this Memoir by Dr. Edwin H. Colbert, where the views of Haug, Hopwood, Pinkley, et al are
geologists (see
cited) to regard the Villafranchian as of Lowest Pleistocene (or Quaternary) rather than of Upper Pliocene age. Apparently one of the most reliable factors
in the determination of the boundaries between the Pliocene and the Pleistocene, aside from the geological evidence, is a change in fauna, for example, the
presence of Bos (including Bison), Elephas, or Equus, would indicate the Pleistocene age of the deposit in which they were found. Compare also Haug, 1911.1,
pp. 1701 and 1767, Hopwood, 1935.2, pp. 46 and 47.— Editor.]

^[See page 1071 below where Professor Osborn expresses the opinion that migration to America may have taken place during Sd Interglacial, and possibly
as early as 1st Interglacial time. — Editor.)
1050 OSBORN: THE PROBOSCIDEA

They also recognize (p. 180) the "Rameaii de VElephas trogontherii" in the Lower Pleistocene Forest Bed of

Cromer, but find no trace of it in the Pliocene superieur Crags of England.

From the Forest Bed they trace (p. 181) the phylum E. trogontherii, with gradual increase in size, through
the terraces of 30 meters (Chellean or Tyrrhenian stage) in the valley of the Thames, also at Gray's and at Ilford

where Lydekker (1886.2, pp. 127, 134) mistook E. trogontherii for E. antiquus.

In Germany observations fail to indicate the Pliocene age of E. trogontherii, but this species appears in a 1st

Interglacinl stage corresponding to the Forest Bed (Sicilien, Lower Pleistocene), also according to Wiist (1901, p.

61) in the ancient gravels of the Ilm at Sussenborn near Weimar, where it was very abundant. This level they
regard as contemporaneous with the 1st Interglacial sands of Mosbach near Wiesbaden and of Mauer near Heidel-
berg (Chellean). They also state (p. 182) that Pohlig records E. trogontherii from SUssenborn, Heinthurm and
Denstedt, near Weimar; Brucksdorf near Halle; Suiza, Angelhausen, near Armstadt; Apolda, Mosbach, near
Wiesbaden; the lower gravels of Taubach, and of Rixdorf near Berlin; the gravels of the terrace of 30 meters at
Mauer near Heidelberg, and the travertines of Taubach and of Weimar. It became extinct, without extending
liigher, in the Quaternary.

Osborn, 1929: Observe the constantly ascending ridge formula in M 3 ^il (in the Upper Pliocene) to M 3 M
(in the Upper Pleistocene), a period of perhaps 1,250,000 years. Observe also that during 3d Interglacial times
Parelephas disappears in western Europe and in early Pleistocene times it appears in North America.

PRIMIGENIUS Rtr
falo ,
1847. Pi, XLV, F.g XXIViftv."

r. PH!MIGCNIUS Atf
P&'.-B. I89I p 30*. Fifl t20
E PRIMIGENIUS Rel
Fi'c. IW, Pi. Xtlll, Ftg. XXIV
E. PRIMIGENIUS TROGONTHERI
Polil'9. 1691. p 368. Fig 121

Fig. 934. Cranial Profiles of Mammonteus primioenius (1,2,5) .\nd Parelephas troqontherii (3,4,0) of Europe
One-twentieth natural size. The plate lettering is incorrect (compare Fig. Sfi."), ii[)i)pr row, ami koy, p. 977)
3. Parelephas trogontherii (female); front view of cranium. After Pohlig, 1891.
4. Parelephas trogontherii (female); side view of same eraiiium. After Pohlig, 1891.
G. Parelephas trogontherii; adult male cranium in .side view. After Falconer, 1847.
i). Mammonteus primigenius; adult cranium. After Pohlig, 1891.
1. Mammonteus primigenius; front view of cranium (after Pohlig, 1891), wliiili resembles Parelephas in certain characters.
2. Mammonteus primigenius, front view (if cranium (after Falconer, 1847), which apjjears to be tyi)ical of 'E. primigenius.'

Observe that the Parelephas trogontherii cranium, both in the male (6) and in the female (3, 4), is relatively lower, broader, and less compressed fore and aft
than the Mammonteus primigenius male {'i); in other words, it is less hy[).sicephali<' and less bathycephalie, with broader space between condyle and orbit.
Compare esjiecially (o) M
primigenius and ((1) /'. trogontherii, both males.
.
 ;.

THE MAMMONTIN^.: PARELEPHAS 1051

Distinctive Cranial Characters of Parelephas

Up to the time that Pohlig separated the grinding teeth of Parelephas trogontheru from tlie grinding teeth of
Mammonteus primigenius the crania in these two distinct lines of generic descent were confused, as well as the
grinding teeth, yet, as shown in figures 865, 934, 935, and 937, the crania of Parelephas are readily distinguishable
from those of Mammonteus.

The principal characters by which we distinguish the Parelephas crania are the following: (1) In frontal
aspect the crania of Parelephas are relatively broader, more spreading, and more brae hy cephalic than those of
Mammonteus, which are deeper and more bathycephalic ; (2) in lateral aspect (a) the orbit is more widely separated
from the occipital condyle, (b) the occiput is much more convex, thus throwing the occipitoparietal apex farther
forward, (c) the height from the occipital apex to the superior molar crowns is less deep, i.e., less bathycephalic,
(d) the apex formed at the summit of the cranium is less acute, (e) the facial front is shorter and more deeply
concave, (f) the maxillo-premaxillary sockets are less vertical and the tusks emerge in a less vertical plane; (3) in

frontal aspect (a) the premaxillary sockets are more expanded at the extremities, whereas in Mammonteus they
are more elongate and more closely compressed, (b) the transverse diameter of the frontals is relatively broader

than in Mammonteus, (c) the anterior nares are proportionately broader transversely and less deepened vertically
(4) in brief, the jirojiortions of the cranium of Parelephas throughout are harmonious with those of the grinding
teeth, i.e., less compressed anteroposteriorly, less bathycephalic and less hyp.sicephalic than those of Mammonteus

Profiles (Fig. 935) of juvenile, adult, and middle-aged crania of Parelephas reveal a contour which is readily
distinguishable from that of the larger Archidiskodon cranium, on the one hand, and that of the smaller and much
more compressed Mammonteus cranium, on the other; but it is not at all surprising that Falconer confused the
Parelephas and Mammonteus crania, because they present so many points of subfamily (i.e., Mammontinae)

Left Lateral Profiles of Seven Species of Parelephas with Prooressive Ridge Formula
One-thirtieth natural size

Fig. 93.5. This series is remarkably uniform in its progressive characters:

1) M 3 x^ to M 3 l-f ; superior ridge-plates exceeding the inferior ridge-plates in number.

2) The orbits lie just below the level of the occipital condyles, with relatively wide separation.
3) The restored in P. inlermedius (C), P. columbi (E), and /'. floridanus (F), but is perfectly preserved in P. trogonlherii (A, B),
occipitofrontal profile is

female and male of Mid-Pleistocene time, al.so in P. washinglonii (D), P. jeffersonii (G), aged type, and P.jeffersonii (H), giant male in the Nebraska Museum.

4) All the profiles present a concave forehead, a moderately elevated summit, and a rather angulate occipital contour.
5) As shown by comparison with figure 834, the hypsicephaly, acrocephaly, and bathyeephaly of the cranium are much less acute than in Mammonteus.
ior)2 OSBORN: THE PROBOSCIDEA

resemblance. The relation (Figs. 806, 816) of the basis cranii to the fronto-occipital contours reveals a very

jirofound difference between the fully adult Parelephas jeffersonii (Nat. Mus. 10261) and the adult Elephas
iitdicus profile, on the one hand, and the Loxodonta africana profile, on the other. In all stages Parelephas agrees

both with Arc/iidiskodon and Mammonteus in the concave frontal plane or forehead.

PARELEPHAS TROGONTHERIl 4500mni..l4'9/a" e PARELEPHAS FLORIDANUS 272] mm.,\2'2>/t" e

GERMANY MOSBACH FLORIDA

PARELEPHAS TROGONTHERII 4054mm..l3'3>^" e PARELEPHAS JEFFERSONII 3402mm.,ll'lV

GERMANY TAUBACH INDIANA

ELEPHAS INDICU3 BENOALENSIS 3200min„10'6" PARELEPHAS COLUMBl 3b4omm.,l I'l l/e"

INDIA GEORGIA

Fig. 936. Parelephas of Europe and America in comparison with Elephas indicus benoalensis
Rostoration.s to a one-fiftieth scale by Margret Flin.scli Bul)a, under tlic diieetion of Henry Fairfield Osborn.

The crania conform in size with the three skeletons more or less fully known, namely, of Parelephas inter-
medhis of France (Fig. 944), .said to attain a skeletal height of 3750 nun., or 12 ft. 3^^ in., of P. jeffersonii in the
American Museum (Fig. 966), with an estimated height of 3200 nun., or 10 ft. 6 in., of the giant P. jeffersonii of
the Nebraska Mu.seum (Neb. Mus. 1-4-15), the skeleton of which is unknown, and of P. columhi of the Amherst
Museum (Fig. 955), with an estimated height of 3430 imu., or 1 1 ft. 3 in., finally of the giant P.Jloridanus, known
 J

THE MAMMONTIN^: PARELEPHAS 1053

from an associated humerus and femur in the American Museum to attain an estimated height of 3584 mm., or
11 ft. 9 in., probably influenced by the favorable climatic conditions of Florida in contrast to the more severe
northern conditions which produced P. jeffersonii. For estimated height in the flesh, compare figure 936 on
opposite page.

A. IMPERATOR Kef. M. PRIMiqENIUS Fief. P. JEFFERSONII 7")/oe .- WAaHIN^TONII

/.OS AngeZes Mu-s. Var. /^u5 SS30 Amsr. Mu:,. 9 9 SO Araer. /Vtts. 66

^20 natural sijS

Fig. 937. Cranial Profiles of the Mammontin.« of America

One-twentieth natural size
(A) Archidiskodon imperator, young male in the Los Angeles Museum.
(B) Mamnwnieus primigenius, adult male, dwarfed, in the U. S. National Museum.
(C) Parelephas jeffersonii, type aged male in the American Museum of Natural History.

(D) Parelephns washingionii, referred young adult male in the American Museum of Natural History, the least progressive of the series.

Observe extreme progressive hypsicephaly, cyrtocephaly, and bathycephaly as we pass from right (D) to left (B, A); thereby the occipital crest
is elevated, the grinders are depressed, the orbits are approximated to the condyles (B), the face is flattened or concave.
 LeCTOTYPE and CoTYPES of PaRELEPHAS TROQONTHERIOIDBS
About two-fifths natural size
Fig. 938. Lpctotype and cotypes of Elephas anliquus Falc. var. trogoniherioides Zuffardi, 1913, Tav. ix (iii), figs. 3a-G6, namoly: Lectotype from Nizza
dolla Paglia (Astiisan), third left superior molar, l.M' (Ha, 6'6), with 14+ ridgc-platcs; cotypes from near Sail Paohi de Villafranea, third riglit inferior molar,
r.M3(4), with +14+ ridKe-i)latcs, third left inferior molar, I.M3 (50,56), withWH '; ridge-plates and talon, and.seeond left inferior molar, I.M2 (3(i, 36), withal least
10 ridge-plates. Comjjare Zuffardi, op. cil., pp. 130, 155-158, also Dep6ret and Mayct, 1923, pp. 177 170. These figures arc designated by the author as follows:

"|I'ig.] 3a.— Molare probabilmonte quinto inferiorc sinistro — pag. 155

(G), [35].
" 36. — Lo stes.sovisto dal fianco osterno.
" 4 — Molare ultimo inferiore destro
.
— pag. 156
('8'), [Mus. Palais Carignan, Turin.]
[36).
" 00. — Molare ultimo inferiorc sinistro — pag. 156
('9'), [36|.
" 56. — Lo stesso visto dal fianco intcrno.
6a.-— Molare ultimo supcriore sinistro —
('5'), [Geol. Mus. Turin]
p.ag. 1.58 [381.
06. — Lo stes.sovisto dal fianco csterno. [Geol. Mus. Turin.]"

1054
 —

THE MAMMONTIN^: PARELEPHAS 1055

2. SYSTEMATIC DESCRIPTION OF EUROPEAN AND ASIATIC SPECIES IN ASCENDING

PROGRESSIVE ORDER

Parelephas trogontherioides Zuffardi, 1913 concava e di forma Lunghezza della corona m. 0,156.
ovale. . . .

Figure 938 T>arghezza massima m. 0,105. Altezza massima della

della corona,
Type: Upper Pliocene (Villafranchian stage),' Piedmont, northern Italy, corona m. 0,094. Indice dentale 0,015. Frequenza laminare 6^.
Nizza della Paglia and San Paolo de Villafranoa (Astesan). Referred: Red Simbolo [ridge-plate formula] a- 10 ." Zuffardi states on page—
and Norwich Crags of East Anglia. 156 that the two third inferior molars [cotyi)es] (Tav. ix, figs.
Third superior and inferior molars, M', M3, described as 4, 5a, 56) were [erroneously] referred to Elephas antiquus
a mutation of small size of Elephas trogontherii; grinding teeth of Falconer and Cautley, from the Pliocene lacustrine of Astigiana
reduced dimensions; ridge-plates of similar character to E. 5. Paolo, according to the label attached to the specimens in the
laminar frequence 6 to 6.5 in 10 cm.; total esti-
trogonlherii; Turin Museum, numbered 8 and 9 respectively. No. 5 [lectotype]
mated ridge-plates in M
3 *tili*? as preserved in the lectotype of the collection (Tav. ix, figs. 6a, 66), also [erroneously] referred
(Fig. 938, 6a, 66), in the cotype M 3 ^^tttzjs (Fig. 938,4), in the to Elephas antiquus, is from Nizza della Paglia (cf. p. 158);
cotype M3 ^^rnr^j (Fig. 938, 5a, 56) ; estimated ridge-plate formula length of crown 210 mm., maximum breadth of crown 110 mm.,
M 3 virTT^fl somewhat inferior to that of E. trogontherii, as we maximum height of crown 150 mm., laminar frequence 6M to 7,
should anticipate. We may depend upon the observations of ridge-plate formula 15 x. —
Deperet and Mayet (1923, p. 178)
Falconer, Pohlig, Zuffardi, and Deperet as to the resemblance of estimate the total ridge-plates of this type specimen at about 20.
these cotypes to E. [= Parelephas] trogontherii, which, if cor- Zutt'ardi's full type description is as follows:
roborated by future disco^'ery, make this the most primitive (Zuffardi, 1913, p. 158) "La stessa discussione puo farsi in
species. riguardo di un altro grosso molare superiore (Tav. ix [iii], fig.

As to size, Osborn observes that whereas the typical molars of 6a, 66). Euelephas antiquus Cautl.
Esso porta I'indicazione '5'

Parelephas trogontherioides e.xhibit fewer ridge-plates (M 3 qij^), et Falc. Nizza della Paglia —
Dono del sig. cav. Pio Corso Bosen-
they are about equal in size (length, ^P, 2104-mm., breadth 110 asco' ed e accompagnato da un biglietto cosi concepito 'n.'' 20
mm., height 150 mm.) to Pohlig's type (Fig. 939) of P. trogon- Rinvenuto nel 1819 in un podere della famiglia Corso Bosenasco
therii mm., breadth 101 mm., height 138 mm.),
(M*, length 225 dal sig. Pio Corso Bosenasco Sindaco di Nizza Lapaglia, a Nizza
AF of P. trogontherii from
also superior in size to the smallest Lapaglia.' fi un dente alto e massiccio, troncato davanti, arro-
Siissenborn near Weimar (length 213 mm., breadth 74 mm., tondato e assottigliato in dietro. I fianchi della corona, ancora
height 118 mm.); consequently P. trogontherioides cannot be rivestiti dal cemento, sono verticalmente quasi diritti, e longitu-
described as a "mutation de petite taille"; it is, however, inferior dinalmente convesso I'interno, concavo I'esterno. La superficie
one of the largest superior molars of Siissenborn, namely,
in size to di abrasione, quasi rettangolare-arrotondata, e composta di dodici
length 317 mm., breadth 76 mm., height 183 mm. (see Soergel, lamine, tortuose, larghe come gli interspazi di cemento e tendono
1912, PI. viii). ad allargarsi bruscamente nel mezzo in modo da originare una
E. [Elephas] antiquus var. trogontherioides Zuffardi, 1913. espansione ovolare. Le coma piegano piii meno sentitamente
"Elefanti Fossili del Piemonte." Palaeont.-Italica, XIX, pp. air indietro, le lamelle sono leggermente pachiganali e fittamente
130, 155-158. Lectotype and Cotypes. — Lectotype, third increspate. Della 1.* lamina e presente solo la meta interna della
left superior molar, l.M' (Fig. 938, 6a, 66) cotypes, third right ; lamella di smalto posteriore, e della 2." manca la meta esterna della
inferior molar, r.Mj (4), third left inferior molar, I.M3 (5a, 56), lamella anteriore. Sino all'8.^ compresa, le lamine hanno figura
and second left inferior molar, I.M2 (3a, 36). Hokizon and completa; la 9", 10", 11." hanno tre elementi laminaroidi di cui il

Locality. —Upper Pliocene (Villafranchian stage),' Nizza della mediano piu sviluppato, e assai meno I'estremo interno ridotto
Paglia and San Paolo de Villaf ranca (Astesan) Piedmont, northern , quasi a un semplice anulo. La 12." e composta di quattro cercini
Italy. Lectotype and Cotype Figures. — Zuffardi, op. cit., appena usati. Segue I'altra parte della superficie di abrasione non
1913, Tav. IX (ill), figs. 3a-66. usata che forma con la precedente un angolo di poco superiore al
—
Description. (Zuffardi, 1913, p. 130): "Pero pensando che
retto. Essa quantunque ricoperta ancora da cemento, dalle costo-
il Jourdan considerava la sua specie E. intermedius come specie
lature lateral! puo ritenersi constituita da cinque lamine delle quali
di passaggio dall' E. antiquus all' E. primigenius, per evitare
I'ultima, strettissima e arrotondata, puo considerarsi come tallone
dannose confusioni, proporrei ora il nome di E. antiquus var.
prossimale."
trogontherioides alia varietii in questione, in ricordo del nome
"Le radici, per la massima parte perdute, si iniziano in un
specifico dapprima dato al materiale attualmente ad essa attribuito.
... 155] Ricordo prima un dente [cotype I.M2] (Tav. ix [iii],
brusco restringimento laterale della corona e vi si vedono i tronconi
[p.
fig. 3a, 36) che
trovava in raccolta senza indicazione alcuna e che
si
basali di un paio di robusti rami anteriori probabilmente liberi, e i

distinguo colla lettera G. La corona larga con fianchi non molto resti dell'ammasso delle altre radici. Dall'andamento di queste e

convessi, termina anteriormente con una faccia ristretta e arroton- dalla forma della corona sembra manchino al massimo altre due
data, e posteriormente e tronca. La superficie di abrasione e amine."

'[See footnote 1 on p. 1049 above regarding the possible Lowest Pleistocene age of the Villafranchian. —Editor.)
 , —

1056 OSBORN: THE PROBOSCIDEA

"Valori metrici: earth colored by fire ("rotgebrannte Erde"), also bones belonging
^ , , ,, „r.,r> an ancient canipina; place of priniiti\e man which inieht well
to
Lunehezza della corona m. 0,210 , , , ,, ,, e ,i , , , •

have beioneed to Uunui fiei<ltlberqeii.sis; oi the same geologic age

, i

, . , 11 i< ,> ,,,A

Larghozza niassima della corona 0,110 .. /, * r, j u- u -^ ^u j -^
.
Altezza massima della corona
-,11 ,< ,>
0,150
, r,.
, /t^
i^

, i M\f\n\ t-.u
ii
iis the Cromer I'ore.st Bed which is quite as old as the deposits of
^ u ^ i ^•
r •

^ ,. , , , ,1 ,^,^-,r Jockgrim (1* reudenberg, 1909). 1 he very ancient characteristics

Indice dentale 0,015 , ^/!
of this small c Irogoidneni
u E. <j. •• . . /
mutation at once eonhrm c ^u <
. .
„,^ „ the great
'

IK" antiquity of this basal Mosbach sands horizon, which also contains
the Etruscan rhinoceros (Dicerorh-inus eiruscus), Trogontheriuin
"fi d'uopo notare pero che il Falconer ([40), vol. II, pag. cuvieri, and Hippopotamus. In this lower Mosbach stage of this
249-250) anche a questo dente quando parlando
si riferiva for.se small E. trogontherii mutation, the third inferior molar, Ms, ex-
del suo E. nrmeiiiacus, che credeva ravvisare in un grosso molare hibits 8 ridge-plates in 10 cm., e.\actly as in the primitive Irogoii-
segnato col n.° 7 nella nostra raccolta, e che ci occupera ])iu avanti Iherii molars of Jockgrim, whereas the true Elephos primigeiiiiis M3
(pag. 174 [54]), affermava che altri ve n'erano tra molari di S. i exhibits 7 ridge-plates in 10 cm. These basal sands of Mosbach
I'.aolo." cannot be more recent than the 1st Interglacial (Giinz-Mindel)
„ T /i~v ^ AT ,rvr>n -/. Stage of thc Alps, or the Elephant Cromer (Forest Bed Bed of
,„„,
Pliocene of Italy (Deperet and Mayet, 1923, pp. 1<6-
,. , ,. .
/,D/.Dx Ti 11- /,oo,A r,oooiN
,

SBrics).
„^.,,. , ^ V, \ t,,
Still uiorc aucicnt arc the Underlying laycrs at Mosbach
. , , . , > <

183).— According to lalconer (1868), Pohhg (1889 1888), and

, I

„, ^ .•.•,. , ,
an tt
, . .

, /,r^,oN containing tlephas = AirnidisKodon\ mcndionahs and Upper

, 1 , ,
r, «-

,
.. r.

,
L

,
..
Zuffardi (1913), this rarelephas trogontnerioides branch
r-
i.e.,
, c ,
I .

•
il TT
• •
I

ni-
I 1

/TT-ii
„,.
1 liocene launa with
. .,,
[

.,,,,/
=
Mastodon \ Aiiancus\ nrrernensis and the
,
• , !,
,

I)hyluiii]
,

although rare hrst appears in the L pper Pliocene (Villa- „ , ,,

fninchian) of Italv. As discovered near .San Paolo de Villafranca ,, .

,„„'o' n^, , i t-v ^i-il t^ j i
Osborn, 1928: Ihe above note by Dr. Wilhelm I'reudenberg
., 1

„ ,," ,,,
,

1-
.

/ ^L r n.
., .,

L
,

.
,

.L
,, ,,
this lectotypej M-" exhibits so closely all the characters of the cor-
-.i . i
.
1 , ,

11
.

IS
, 1
deeply interesting as tending to eonhrm
.
, ,. . i- . ^ ^^ ..-,,/•
the great antiquity of
,1
.

responding tooth ot A/emflas <7'ooon</!e?» that it can only be separat- ,1 r. , , , r^ ,, ..1

,' . ,, ,
,,n^ox .• 1 r. Ti 1 \. 1 the Parelepnas phylum in western Europe and the po.ssible co-
lalconer (1808) mentioned ban Paolo as one of the
ed as a variety, ., , T .,. , r. /o „ j
,...,,,
m^ , ,, 1,. ,1 , r, . .
,
l
existence of a primitive species of f'amepna.s
.
>
( i^sp.
.
<rono«//(e)7o/ae.s)
\

localities Italy containing teeth resembling the type 01 E. .,1 „ n /w

.,.,,. 1- f
armemacus of Asia Minor, which he accordingly referred to the ILL 1 1 . 1
with Heidelberg man
1
(Homo 1 1 .1
heidetberqensts).
.

Armenian species (seep. 1060 below). ,oor 1000 lom

,, ,,.' ,\,
ror this and other T,
,,,.
I pper Pliocene teeth found at or near San
c. ,,,,,, Parelephas trogontherii r,,,-
Pohlig, 1885, 1888-1891

ni ij 11 ic lutuu'/o-j- luT^-..
Paolo and described and hgured both by Zurfardi and by Deperet
Figures 794, 865, 871, 929, 934-936, 939, 940, PI. XXII

^, ,,1 I -n
and Mavet, the subspecihc m L a?i^(/«MS
name A/ep/io.s
,
J- .1
var. //ooonZ/jeri-
i

,,,
Type:
' ,
2d Interglacial deposits at Siissenborn near Weimar (Fig. 932,
^ „, j ^ ^, , , •, t> ,•./Toot <>
'

, ,
11)> northern Germany. Referred: Isl-M. ,
Interglacial. Pohhg (1887.2,
oides was given, the characters being summed up as follows
1 1

,,.274). "vom mir zuerst in den alten Flusskiesen ThUringens zu Sii.s.senborn

(Deperet and Mayet, 1923, p. 179): "Ces molaires presentent tous etc. bei Weimar in grosserer Menge aufgefundene altdiluviale Ele|)hantpn-
les caracteres essentiels de VE. trogontherii quaternaire, savoir: zahnform beschrieben."
une couronne tres large et tres haute, avec des lames ^paisses et This very important species, widely distributed in 2d Inter-
a.ssez ecartees (frequence laminaire de 6 a 6, 5) un email tortueux ; glacial times in western Europe, is, with the more progressive
et fortement plisse, orne sur sa longueur de quelques sinus aigus Elephas intermedius of Jourdan, a member of the distinct phyletic
mais ne ressemblant en rien au sinus loxodonte median
irreguliers, series Parelephas. Parelephas trogontherii belongs to a phylum by
des E. meridionalis et antiquus. ... [p. 180] On pent done dire itself which appears sparsely in Europe at the close of Pliocene
seulement que la forme pliocene de VE. trogontherii possede des time and reappears in the /.s/ and .^rf/Htec^/arm/ periods. We do not
molaires de dimensions plutot un peu n'duites, sans qu'il puisse agree with Pohlig's opinion that the species i?. ^ro^ow/Zier// is a con-
etre question d'une veritable mutation de petite taille de I'espece. necting ancestral link between Elephas meridionalis and E. primi-
t)n doit s'attendre a d^couvrir un jour dans le Pliocene ancien de genius. With Deperet we regard it rather as a distinct phylum,
(juelque region eloignee les types ancestraux nains du rameau de The estimated ridge-plate count of the type, M^ (Fig. 939, upper)
VE. trogontherii." is {-;, somewhat exceeding the ridge-plate count of E. [P.] trogon-

FiRST Interglacial of Germany (Freudenberg, 1926). //ienoirfe.s Zuffardi.

In the basal layers of the Mosbach sands, near Wiesbaden (lug. Soergel (1912.2, Tab. vii, viii) records three inferior and
932, 12), Freudenberg records small mutations ("kleinere Vari- five superior molars from the type locality of Sussenborn {1st to
ante") which he refers to Elephas trogontherii, associated with 2d Interglacial deposits), in the liebling and Weimar Museum
artificially broken off ("kiinstlich abgetrennte") portions of the collections, which vary widely in length, breadth, and height, as
horns of Cervus verticornis, charred wood ("Holzkohlen"), and follows:

Sussenborn: Reeling and Weimar Mus. Ridge formula Length Breadth Height
R.M3 No. 46 (Weimar) xl8x(xl7.x) 270 62 160
17 (Weimar) 16 x 255 83 105
10 (Weimar) x21x 316 73 137
R.M» No. 80 (Rebling) xl8x 302 91 192
69 (Hebhng) 20 x 291 88 160
32 (Weimar) x20x 296 66 137
89 (RebUng) xl8x 264 71 162
87 (Rebling) x20x 274 85 171
 ;

THE MAMMONTIN/E: PARELEPHAS 1057

Elephas lnty<intherii Pohlig, 1885. "Ueber eine Hipparionen- in ridge formula, in the structure of the
crowns of the molar teeth
Fauna von Maragha in Nordpersien, iiber fossile Elephantenreste it issharply separated, because the grinders of E. irogontherii are
Kaukasiens und Persiens und iiber die Resultate einer Mono- different in proportion from those of E. antiqu us; the enamel
graphie der fossilen Elephanten Deutschlands und Italiens."
Zeitschr. deutseh. gaol. Ges., 1885, XXXVII, Heft IV, p.
1027. —
Type. Apparently a last superior molar, M', of the
right side, also a last inferior molar, M3, of the same side. Hori-
—
zon AND Locality. 2d Inlerglacial deposits, SiJssenborn near
Weimar, northern Germany. —
Type Figure. Pohlig, 1888,
p. 193, fig. 79, and p. 195, fig. 82 (Fig. 939 of the present Memoir).
Schwabe Coll., Weimar. Referred Specimens. The Siis- —
senborn stage of the upper and lower grinding teeth of Elephaa
trogontherii, illustrated by Wiist, 1901, Taf. 11, figs. 4-12 (Fig. 940),
appears in comparison with the stage erroneously referred to
'Elephas' [Hesperoloxodon] antiquus in figures 3 and 18.

RELATIONSHIPS OF ELEPHAS TROGONTHERII POHLIG (1885,

1887, 1888-1891, 1907, 1912) AND SOERGEL (1921)

Relationships.— (Pohlig, 1885, p. 1027): "6. Unter der

Bezeichnung 'Elephas trogontherii Pohl.' fiihre ich in meiner
Monographie eine europiiische Molarenform auf, welche zwischen
denjenigen des E. primigenius und E. meridionalis zoologisch,
wie ihrer geologischen Lagerstiitte naeh, in der Mitte steht, dem Fig. "3. Letzter Maxillenmolar von Elephas trogontherii. Original zu Weimar
E. antiquus in der Lamellenformel am nachsten kommt, aber durch (Dr. Schwabe), '/a

die Kronenform und geologische Lagerstatte von dieser Art weit

scharfer gesondert ist alsvon den anderen beiden Species. Das
Verhaltniss von E. trogontherii zu E. armeniacns Falc. und audi
E. namadicus Falc.-Cautl. bleibt noch genauer zu untersuchen. —
E. meridionalis und E. primigenius stehen nach Kranologie und
Dentition iiber E. trogontherii hin in director Verwandtschaft."
Description (Pohlig, 1887.2, p. 274) "Unter diesem Namen
:

Elephas trogontherii habe ich in meiner Monographie eine, wie es

scheint, in Europa allgemein verbreitete, von mir zuerst in den
alten Flusskiesen Thiiringens zu Siissenbom etc. bei Weimar in
grosserer Menge aufgefundene altdiluviale Elephantenzahnform
beschrieben, welche geologisch sowohl als zoologisch die beiden
Arten E. meridionalis Nesti aus dem Pliocaen und E. primigenius
Blum, aus dem Plistocaen vollig mit einander verkettet: in der
Form der Krone und Kaufliiche und indem Charakter der ein-
zelnen Schmelzfiguren auf letztercr stimmt jene altdiluviale
Rasse mit den zuletzt genanntcn beiden Arten iiberein, steht
jedoch,was die Anzahl der Schmelzlaniellen und die Dicke der
Schmelzwande anbetrifft, zwischen diesen beiden Species in der
—
Mitte, in den unterplistocaenen Schichten mehr dem E. meri-
dionalis, inden mittelplistocaenen Lagerstatten mehr dem Mam- Fig. 82. Letzter Mandibelmolar von Elephas trogontherii. Original zu Weimar
muth Beziehung angenahert."
in dieser
(Dr. Schwabe), '/,.
Pohlig (1885-1887) from the first expressed the opinion that
Elephas trogontherii forms a connecting ancestral link between Types or Parelephas trogontherii. Compare Soeroei,, 1912.2, Tab. vii
Onr-third natural size
Elephas meridionalis and Elephas primigenius, its earlier represen-
Fig. 939. Type third suijorior and inferior niolar.s of Elephas trogontherii
tatives resembling the former, its more recent representatives re-
Pohlig, 1885. After Pohlig, 1888-1891, p. 193, fig. 79, and p. 195, fig. 82.
sembling the latter, accordingly he wrote the specific name at the These arc the specimens first mentioned by Pohlig (1885, p. 1027) and figured
time E. primigenius trogontherii. He held that this species entered by him in his Memoir of 1888-1891: "Fig. 79. Letzter Maxillenmolar von
Europe during the second glaciation (Saxonian-Mindel) and sur- Eleplias Irogonlhrrii. Original zu Weimar (Dr. Schwabe),
):)." This is apparent-

vived during the ,^r/ Inlerglacial period (Helvetian, Mindel-Riss) ly a molar of the right side, r.M'; it is eomijlcte and displays 15
last superior
ridges. "Fig. 82. Letzter Mandibelmolar von Elfplms trngntdhrrii. Original
thus E. trogontherii was contemporary with the appearance of the
zu Weimar (Dr. Schwabe), }:1." This is a last inferior molar of the right side,
Chellean industry in France (Penck). Pohlig (1888-1891, p. 458)
r.Ms; it is broken in front and displays 16+ ridges. The incomplete ridge
pointed out that while Elephas trogontherii resembles E. antiquus formula of these types of Elephas trogontherii is: M
^\%X- See text below.
 :

1058 OSBORN: THE PKOROSCIDEA

bands rise above the cement not vertically but obliquely, after the eit., pp. 69-97) concurs with Pohlig in the opinion (hat Elephas
manner of (he Asia<icelci)hant. PJ. /lof/o/f/An// hesi)caks !i rolatixo- irogontherii is ancestral to Elephas primigenius.
ly moist climate but is also occasionally minj^lcd w itli remains of Haditai'. — In a letter of ,lune 30, 1922, Soergel states:
the reindeer (linngifrr) where (he forest bordered on the open According to my interi)retation E. trogontherii lived during a period
country, as a((os(c(l by (he appearance of bo(h animals in Siisson- of limited rainfall, in a half-arid climate, not migral-ing during
born andS(einhoim. Pohlig (18S8-1891, p. 380, fig. 121) figures the glacial times inio the colder ice-dad regions of middle lOurope
female skull as Elephas {primigenius) Irogontherii (reproduced in (compare also Soergel: 'Die Ursachen der diluvialen Aufschot-
Fig. 865, Nos, .3, 4 above), in which the superoccipital region is terung und Erosion,' Verlag Porntrager, 1921). E. trogontherii
imjierfect and restored. lV)hlig's opinion as to the (mnxitiomd therefore occupied the region la(er traversed by the mammoth.
character of this animal between E. mcridionalts and E. primi- We still regard E. trogontherii as the direct ancestor of E. primi-
genius is indicaied in (he names he has successively assigned to it, genius. We possess in the museinns of I'^urope all the transition
namely forms numbers and are unable to make any sharp dis-
in large
Elephas Irogontherii I'ohlig, 1885, p. 1027. tinction between the two species. If one seeks a complete under-
Elephas (primigenius bczw. mericlin)mlis) irogontherii, 1887, standing of the chief literature of value on this subject, it may be
p. 274. found in the following works: M. Pavlow, 'Les elephants fossiles
Elephas (primigenius) irogontherii Pohlig, 1887 11888'J, i). 799. de la Husse,' Nouv. M<'>m. d. 1. Soc. Imp. des Nat. de Moscou,
Soergel (1921.2, \). 68) in his comi)rehensive review eniided 'J\ XVII, 1910; W. Soergel, 'Elephas trogontherii vmd Elephas
'Elephas rulumbi I'alconer" likewise adopts J'ohlig's opinion and antiquus etc' Palieonlographica, 1912; 1']. Wiist, 'Das Plioziin
places Elephas meridionalis-trogontherii-primigenius as successive und das iilteste Pleistoziin Thuringens. Abhand. d. nalurf.
jjhyletic stages in the same west iMiropean phylum, fie assigns (iesellsch. zu Halle, Bd. XXIII, 1900. 1 rejoic'C that in tlie iirinci-

(op. oil., J). 55) the following collective ridge formula to (he Elephas pal points we agree as to the diluvial elephanls and their phylogeny.
Irogontherii specimens of Sussenborn and Mosbach: PHVLOciENKTK; (
"oNcusioN (OsBOHN, 1924). Osborn does —
16-2^1 not agree wi(h either Pohlig or Soergel that the typical Elephas
Dp2i Dp 3^'- Dp4-^V Mllfffj M2H-:f= M3f trogontherii is a descendant of A'. [Archidiskodon] meridional is;
Soergel is of the opinion that Elephas trogoDtherii (western nor does he agree that it should be regarded as transitional or
luu-ope) and the tyjjical Elephas eolumbi (southern U.S.A.) are in ancestral to the typical E. [Mammonleus] primigenius. The pro-
similar stages of e\olution. He points out that the typical Elephas found difference between the broad-plated molars of the Aixhi-

Nr. 18 Nr. 6

Fig. 040. Hcfcrrcd molars (r.M.T with 18+ ridRc-pliitc's) of 'Kltphns Irogoittlirrii' Pohlig from tho 2<l I tilrrglacinl of Sii.sscnborii aii<l Woiinar, Oormany.
After Wiist, 1901, Taf. ii. Wiist. orronroiisly refers r.M'' (Nr. 3) and r.M" (Nr. 18) to Elrphas antiquus; they are Parvlcphas trngonllicrii molars from the
type locality (Siissenhorn near Weimar).

"Nr. 3. Klephas antiquus Falc. M.III max. dextr. Siissenborn. . . .

" 6. Desgl. M.III max. dextr. Siis.senborn. . . .

" 18. Elcplias antiquus Fate. M.II max. dextr. Weimar."

meridional is may hv regarded as the direet aneestor of the Elephas phylum widely disl-in-
di.slcodon planifron.s-mcridionalis-imperalor

eolnmbi-imperator phylum: (Op. eit., 1921, p. 68) "Diese Tatsache guishes this phylum from Elephas trogontherii; Osborn accepts
lj<'reclitigt zu der .\nnahme, 'd;iss die nach .Vmerika iiberwandern- rather (he view of Depi'ret and Mayet (Iiat E. trogontherii repre-
den I'ormen der .Meridionalis-trogontherii-llcihc schon beim sents a distinct branch of the mamniolhs. !\Ioreo\'cr, the extreme
Ueberwandcrn resp. kurz vorher besondere Charaktere gegeniiber hypsicephalic, acroccphalic, bathycephalic cranium of Elephas
dem europiiischen El. trogontherii meridionaJis ausgebildet batten, primigenius, with corresponding hypsodon(. finely compressed,
Charaktere, die in der weiteren ICntwicklung sich zum 'Imperaior- and laminated molars, cannot be derived from (he cranium and
Typus' 8teiger(en. Es war also im iiltesten Diluviimi im Krcis der molars of the Elephas Irogontherii type. Osborn agrees entirely
knntincnt;ileii l'',lefantenf()rmen eine ^'aria(i^Ilsbrcilc iiii( zwci \\i(h Dep(''r('( aii<l Maye( (ha( we ha\e (o do wi(li throe (lis(incl

Polen, El. trogontherii mrriilionalis im Wcsicii und ilem direktcn |)hyla, certainly separate from (he clo.sc of Pliocene (im(\ .\gaiii

N'orfahren des El. imperalor im t)s(en vorhanden." Soergel (op. (Jsborn does not accept the ojiinion of Soergel that E. trogontherii

'In the Bibliography of the present Memoir (p. 795), it is possible that 1887.1 should road 1888.
 THE MAMMONTIN^: PARELEPHAS 10o9

is related to the true E. {

= Parelephas] colutnbi of the southern race; he then separates this race as E. Nedii. I'ohlig's lypes or
United States, although it may well prove to be related to tht; (•(jlj/pcs of E. Ne.slii are therefore molars of Forest Ji(!d age.
anunal described by Cope, Hay, Osborn, and others as "Ekphas CoTYPES. —
As shown above, the subspecies Elephas anliquns
cohtmbi" but which now proves to represent a distinct species, Xeslii was erroneously based on cotype specimens selected from
namely, the Parelephas jejfernonii of Osborn. Consequently, Leith .\danis' .Monograph of 1877 1881, pp. 37 and 38, namely,
Osborn, as cited above, has placed E. Irogontherii in the distinct Prit. Mus. 33327, a dredged lower molar from Happisborough with
phylum Parelephas, the generic characters of which are defined bj' + 16 + ridge-plates in 11. o inches [catalogued by Lyrlekker, 1886.2,
cranial and dental characters in the chapter on the "C^lassification I). upper true molar"]; P.rit. .\lus. 27915
128, as a "right third
of the Elephantoidea" (Chap. XV). [given as No. 27515 Adams], a 14+ ridge-plated third left
by I>eith
upper true molar from Walton (lOssex), with 13+ ridge-jjlates in
Parelephas '?j trogontherii nestii I'uhlig, 1891 8.5 inches, and Hrit. Mus. 39463, a half-worn third left lower true
Figure 911 molar from Southwold (Suffolk), compare Lydekker's "Catalogue
Typo loc-alitj': ".JunKplior'aoiicii," Forest Bed (Norfolk), Walton (Essex), of the Fossil Mammalia in the Pritish Museum," 1886.2, pp. 128,
and Soutliwold (Suffolk), England. Lower Pleistocene. 129, and 135. Lower Pleistocene Cromeri-
.Ml these cotyjies are of
Regarded by Ueporet and Mayet (192.'j, j). 103) as a synonj-in of EU/jlut.s an or Forest Jled age; their generic and specific reference to
[JlesperiiUjiodon] aumniua F. Major. Cotype.s regarded by C. W. .\ndrews as
'E. aidiquus' is doubtful; C. W. Andrews refers Prit. Mus. 27915
belonging to E. [= Parelephas] Irogontherii.
and 39463 to 'E. tro(ji)tilherii; Ilopwood (letter, August 6, 1929)
Pohlig (op. cil., ]). 303) in reviewing I>citli .Vdams' Monograj)h
kindly furnished i)hotograi)hs and detailed measurements wliich
on Elephas aniif/iiiis ob.served that certain teetii of the 'riianie.s
tend to confirm ,\ndrews' reference to 'E. Irogonlherii.'
Valley (Adams, 1877-1881, p. 33), of Norfolk (p. 40) and of South-
Hai)i)isborough (Forest Ped), England. Prit. Mus. 33327,
wold (p. 38), ilo not agree with the continental molars [of < Icnnany]
r..M', length 268 mm., breadth 92 mm., height 196 mm.; ridge-
referred t(j E. aidiqims. Pohlig continues, that j)art of the "jung-
plates + 16+, 6.3 in 10 cm.
l)liocaenen . . . I'ore.stbed stammenden" molars do belong to the
Walton (Essex), England. Prit Mus. 27915, \.W, length
typical E. aiiUquus, but that the majority l>elong to an inferior
302 mm., breadth 80 mm., height 167 mm.; ridge-plates 18, G'A
in 10 cm. (referred to 'E. trogontherii' by C. W. .\ndrews).

/j/Vaiu; Southwold (Suffolk), England. Prit. Mus. 39463, I..M3,

length 243+ mm., breadth 83 mm., height 103 uim., ridge-plates

^ A/atu^roJ szje

„ 10.3 8 7
j3 ^
,2
j: / f

Bntfpus. 39463

COTYPKH OF Ei.EPHAS AXTIQUUS nestii PoHMO now TR.\NSI-EUIIKD TO pAHEI-fil'HAsf?) TItOGONTHEKlI .N'KSTII

I'ig. (Ml.
Cotype.s or syntypes of A^cp/iu* art(t9«u« ,N>«(u Pohlig, 1891, after photographs kindly furnished by Dr. A. Tindell Ilopwood of the Uritish
.Museum, August 6, 1929. From the Lower Pleistocene of eastern England. One-third natural size.
(f>eft) Brit. Mus. 2791.J. Left third superior molar, I.M', from Walton (Essex), England, exliibiting I 18 ridge-plates; also crown view of same.
(Right) Brit. Mus. 3946.3. Incomplete left third inferior molar, I.M3, from Southwold (.Suffolk), England, exhibiting 1-14 posterior ridge-plates; anterior
ridge-plates broken off. Lateral and crown views of same.
1060 OSBORN: THE PROBOSCIDEA

14+, 7H to 8 in 10 cm. (referred to 'E. trogontherii' by C. W. gesprochene Vermuthung, dass Loxodon von unhckannten Masto-
Andrews). dontenformen sich ableite, fallen gelassen werden; denn ganz wie
Unfortunately these specimens were not figured by Lcith E. Nestii zum typischen E. antiquus verhiilt sich nach Mitgetheil-
Adams or Pohlig, but they were described by Leith Adams as tem das Wenige, was man vom E. priscus weiss zum modernen . .
.
,

belonging to his 'Ekphas antiquus.' It is to be regretted that the —

E. africanus, es zeigt den Charakter des letzteren minder markirt,
name Elephas antiquus Nestii has been adopted and widely quoted mit Eigenthiimlichkeiten archidiskodonten Geprages, wenn auch
in the literature, for we cannot agree with Deperet and Mayet in geringem Grade, vermengt. ... In der lamellenformel wiirde

(1923) that 'Elephas antiquus Nestii' Pohlig, erroneously based on E. Nestii sowohl mit E. antiquus lypus, als auch mit E. trogontherii
Lower Pleistocene cotypes, synonymous with 'Elephas auso7iius'
is iibereinstimmen; und da an mehreren Punkten der englischen
Major-Depcret-Mayet, correctly based on an Upper Pliocene type Ostkiiste, wie bei Mundesley-Cromer, C'lacton etc. interglaciale

of the Villafranchian stage.' Schichten mit E. antiquus und E. primigenius — an erstercm sogar
History. — In 1891 Pohlig imperfectly established this sub- mit ganz iihnlichem Erhaltungszustand —die jungpliocaenen mit
species (named after Filippo Nesti) as of Upper Pliocene age, E. meridionalis, E. Nestii und E. trogontherii iiberlagern, also

whereas the cotypes selected by him, as mentioned by Leith Adams Molaren aller dieser Formen in die Brandung und von da in die
in his Monograph of 1877-1881, are said to be from Walton, South- Museen gelangen: so war die Schwierigkeit, alles das nunmehr
wold, and Norfolk, the latter from the Cromer Forest Bed level Vermengte in der rechten Art wieder voneinander zu sondern,
which is of Lower Pleistocene age. This subspecies, founded on anfangs begreiflicher Weise selbst fiir den Geiibten zu gross. Und
a misconception as to its Pliocene age, is invalid it is certainly not ;
doch sind eben grade diese Verhaltnisse von grosstem Interessc."
in the same geologic stage as the Upper Pliocene 'Elephas' ausonius Osborn, 1930: We are therefore obliged to regard Elephas
Major-Deperet-Mayet, from the Villafranchian of Italy.' Although antiquus Nestii as comparable to the species 'Elephas' [
= Par-
not figured, the ridge formula was said by Pohlig (1891, p. 305) to elephasi"!)] trogontherii Pohlig (cf. Parelephas trogontherioides

agree with that of E. antiquus: "In der Lamellenformel wiirde E. Zuffardi).

Nestii sowohl mit E. antiquus lypus, als auch mit E. trogontherii

iibereinstimmen."
Parelephas armeniacus Falconer, 1857
E. Nestii Pohlig, 1891. "Dentition und Kranologie des
Figure 942
Elephas antiquus Falc. mit Beitragen iiber Elephas primigenius
Pleistocene, near Khanoos, Province of Erzcrum, Armenia.
Blum, und Elephas mcridioiialis Nesti. Nachtriige," pp. 303,
. . .

304, 465. On page 465 the name appears as Elephas antiquus We note that Falconer in 1868 (Vol. II, pp. 187, 192, 193, 248,
Nestii n.f . Type.— (Letter, "As type
Pohlig, Sept. 10, 1924) : 249, 250) observed a strong resemblance between his type of E.
specimens of E. Nestii from the forest bed are named in my mono- (Euelephas) armeniacus and Upper Pliocene molars from the
graph (p. 304) those of L. Adams' [monograph], p. 38, from Walton locality of San Paolo, or near it, Nizza della Pagha, Italy, recently
and Southwold, and the Nr. 33327 of the British Museum, from chosen by Zuffardi as the types of Elephas antiquus var. trogon-
Norfolk." Locality and Horizon. Forest Bed (Norfolk), — therioides.It is probable, therefore, that the species Elephas

Walton, and Southwold, England; Lower Pleistocene. Type armeniacus of Falconer belongs in the generic phylum Parelephas,
FiciiRE. —
Not figured. See figure 941 of the present Memoir after but we cannot be certain of this reference until the cranial charac-
photographs. Referred skull of northern Italy figured by Pohlig, ters of the species become known. The specific distinctions are:
(ip. cit., 1891, p. 350 = ('raiiiTmi B of Nesti (Elephas vicridionalis); Ridge-plates of M3 "^^^-^
(Fig. 942) , worn anteriorly. This api)ears
referred to Elephas antiquus by Weithofer 1890, and erroneously to be a more progressive stage than Parelephas trogontherii and
by Pohlig to his subspecies E. antiquus Nestii.
referred much more progressive than P. trogontherioides.
Type Description.— (Pohlig, 1891, pi). 303 305): "b. E. (Eueleph.) Armeniacus I'alconer, 1857. "( )n the S|)ecies <if
—
Ein Theil der letzteren die aus dem jungpliocaenen [Footnote: Mastodon and Elephant occurring in the fossil state in Great
'Vgl. u. (sub E. meridionalis) .'] Forestbed stannnenden — lassen Britain. Part I. Mastodon." Quart. Journ. Geol. Soc. London,
(lie Zugehorigkeit wcnigstens zu der Species E. antiquus zu, Vol. XIII, Synoptical Table op]). p. 319. Type. — (Falconer,
nianche mcigen selbst als typische Molaren derselben gelten; die 1863, p. 74): ". . . two last upper molars in fine preservation, and
meistcn haben jcdoch Abweichungcn von der typischen Gleich- a portion of a lower, apparently of the same individual."
all

miissigkeit der Kronenbreite niclit nur, sondern auch von dem Brit. Mus. 32250, 32251, and 32252; also fragment of tusk, head
charakteristisclicn Lamellenbavi der diluvialen Ausgangsform, be- of femur,and portion of tibia (Brit. Mus. 32256, 32254, 32253),
sondcrs von dem ausgcpriigten Loxodontismus, der complicirteren which Lydekker (1886.2, p. 174) regards as belonging to the same
I'estonirung Dicke des Schmelzes Ix^i jener. Da
und der grtisseren individual as the teeth. Horizon an» Locality. — Pleisto-
nun diesc Eigenthiimlichkeiten zum Theil mit den im ersten Ab- cene. Near Khanoos, Province of Erzerum, Armenia. Type
schnitt erorterten, ebenfalls jungpliocaenen Erfunden von der Figure.— Falconer, 1863, PI. ii, fig. 2.

S|jecies aus dem .\rnothal iibereinstimmen, so scheint es mir nun- History. —I'^alconer first named thissi)cciesin his "Syn(>|)(ical
mehr docli geboten, diesen pliocaenen E. antiquus von dem dilu- Table" of 1857, facing i)age 319: "Spec. 14. E. (Eueleph.)
vialen durch cine Rassenbczei(;hnung, ctwa E. Nestii, vorltiufig zu Armeniacus Armenia: Erzeroom ... In the Brit. Mus. Coll.
. . .

(rennen. . . . Es mijsste danach die in dem ersten Abschnitt aus- Discovered between Erzeroom and Moosh in 1856. The molar

'[See footnote 1 on page 1049 above regarding tlie possible Lowest Pleistocene age of the Villafranchian. — Editor.)
 THE MAMMONTIN^: PARELEPHAS 1061

plates closely aijproxiinatctl, and the cnainel-edges very undulat- monly seen in the existing species, less (jpen and less parallel
ed." In 1863, pp. 74-76, he renewed his description as follows: across.The crown is broad, and the enamel plates are high. To
"These Khanoos molars are intermediate in character, between render these descriiitive details more appreciable and available for
the Mammoth and the Indian Elephant, but more nearly allied to comparison, I append the princiiml dimensions [l.M']:^
the latter. ... In the synoptical table appended to my Memoir
on the 'Species of Mastodon and Elephant, &c.,' the Khanoos [In.] [Mm.
fossil form is ranged between E. indicus and E. primigenius, under 11.75 299
the provisional name of E. Armeniacus. [Footnote: 'Quart.
Journ. Geol. Society, 1857, vol. xiii, p. 319.'] . . . [p. 76] while E. 9.5 242
Armeniacus, as stated above, approaches nearer to the existing
Indian species." Falconer also speaks of the remains of fossil
5.7
elephants on the banks of the Bosphorus and the northern shores of
the Black Sea and of the Sea of Azof {op. cit. , p. 75) . On page 74 he
giveswhat may be called the type description.
Type Description (Falconer, 1863, p. 74). "The speci- —
mens presented by Colonel Giels to the national collection, consist
of two last upper molars in fine preservation, and a portion of
a lower, all apparently of the same individual. These molars
strike a practised eye, at the first glance, as presenting something
intermediate between the Mammoth and the existing Indian
 1062 OSBORN: THE PROBOSCIDEA

Euelephn.s armoiianis, auct. This species is definitely known only of a fossilmolar tooth from China, which, on comi)arison with the
by the following [six] specimens, which belong to an animal of teeth presented by Colonel (nils [(Jiels] from .\rmenia, so exactly

\ery large size. The structure of the molars indicates a species resembles them in every respect, that no doubt can be entertained
intermediate between E. pn'inigeniiin and E. indicus, which is as to its belonging to the same species, namely, E. Armeniacus of
probably an ancestral form closely connected with both [Footnote: Dr. Falconer; and as such, I believe, it was regarded by that
76/V/. vol. xlii, p. 174 (1886).']. The crowns of the molars are eminent palaeontologist. The occurrence, therefore, of the same
extremely wide, the ridges api)roximated, with their extremities form in .lapan would not have been very sur])rising; but, so far as
curxing backwards, the enamel is moderately thick and much I am able to judge, it is impossible to identify Dr. Duggan's
l)licated, and the wear of the crown irregular. In their great width specimen with E. Armeniacus."
these teeth agree with those of E. priniigeniun, but in the other .Japan. —
The .Japanese tooth found by Dr. Duggan, in com-
characters with those of E. indicus [Footnote: 'See Leith- Adams, pany with Mr. Hodgson, in 18.59, is referred to Pabeoloxodon (see
British Fossil Elephants(Mon. Pal. Soc), p. 241.']." figure by Leith Adams, 1868.1, p. 497).'
Lydekkcr (1886.3, p. 174) remarks: "From strata of un-
(.5)

known age at Erzerum, in Armenia, Dr. Falconer many years ago Parelephas intermedius .Jourdan, 1861
described some elephant molars under the name of Elephan Figures 935, 943, 944
arnieiiiaciis; and as Erzerum is comjiaratively near to Tabriz, it Pleistocene plateau loess, near Lyons, Rlioue Valley, France. Type
may be suggested that some of the Maragha elephants' teeth may locality l)elieved to be more recent (i.e., fSd I ritcrglacial) than the 3d Inler-

not imjjrobably belong to this species; but be this as it may, the

gladal deposits of Siissenborn, containing the type of Elephas [= Parelephas]
trogontherii.
Erzerum and Maragha faunas may be geographically grouped
.lourdan applied this name to the species of mammoth which
together."
Phylogenetic Conclusion (Osboun): The extremely able he regarded as intermediate or transitional between Elephas primi-
and acute observations of Falconer (18.57-1868) on this species genius and E. indicus. The name is appropriate today for the
from Armenia and on allied forms from Italy, confirmed by specific more progressive and geologically younger than
stage
Parelephas trogontherii. The history of this stage is very interest-
ZufTardi's observations on the resemblance of this animal to
Elephna trogontherii of Pohlig, as signified by the specific name ing, as detailed below. In referred grinding teeth from the same

trogonlheiioides, also the recent observations by Deperet and

locality (Fig. 943), the ridge-plate formula is: M 1 ^2^^ ^^ 3""-.

Mayet (1923), all emphasize the intermediate character of this Deperet (letter, Sept. 10, 1924) assigns the following ridge formula:

species and its phyletic relationship to the Elephafs inlermedius

M3 20-2 1-

of .Jourdan and to the E. trogontherii of Pohlig.

In his original description of this species, Jourdan remarked

The ridge formula, M 3 ±ili*, indicates that E. armeniacus that of all fossil elephants this approached most nearly to Elephas

I'alconer is in about the same stage of evolution as the type of E. indicus. He ap{)lied the name intermedius to certain broad molars

trogonlhrrii Pohlig, in which the ridge formula is said to be M 3 Yi+- resembling in breadth the molars of E. primigenius but with thicker
\\ hilo I'alconer repeatedly described the type of E. armeniacus as
and more widely separated lamellse, in distinction from the narrow

iiitennediate in character l)etween E. primigenina and E. indirns, molars of E. antiguus. Unfortunately the type molars of E. inter-
hut approacliiiig the latter species more clo.sely, Osborn regards it 7nedius have been misplaced and no type figure was i)ublisheil;

rather as belonging to the same phylum as E. inlermedius and E. conseciuently the specific name Elephas intermedius .Jourdan, 1861,

trogontherii and refers it to the indei^endent genus Parchphas. although repeatedly cited, has not gained acceptance; none the
less it is certainly valid. The species is related to but not identical
PARELEPHAS ERRONEOUSLY RECORDED IN CHINA with the widely known Elephas \
= Parrlcpha.^] trogontherii of
AND JAPAN Pohlig, which in turn resembles the Parelephas jeffer.sonii of Osborn.
I'>roneous arc the records of Farelephos in (
"hina and Japan. Deperet and Mayet (1923) describe this animal throughout as
The supi)osed Elcphns nrmeniarus (Brit. Mus. 29007) is related Elephas trogontherii.
(Lydekker, 1886.2, p. 169, Hopwood, letter, 1928) to Elephas Elephas intermedius Jourdan, 1861. "Des terrains siderolit-
[Pula'oloxodon] namadicus. Matsumoto's types of Parelephas iques," Compt. Rend. Acad. Sci., Tome LIII, 1861, p. 1013. Type
protomnmmonleus and P. protomammontcns pro.rirnu.i of .Japan also Description. {Op. — cit., p. 1013): "La faune du .sidcrolititiuo du
iielong to Palnvloxodon. The following literature may be (luotcd: neocene ou etages les plus superieures des terrains tertiaircs sc
LvDKKXKK (1886.3, p. 174).^"There is in the British Mu- caracterise done dans ses couches infericures par VElephas m,eri-
seum an molar from China (No. 29007), which has been
elejihant's dionatis,dans les couches moyennes par VElephas antiquus, ct
suggested to belong to this siiecies [Elephan armcniacux]; and if dans les superieures par VElephas intermedius, (|ui, dc tous les
this were correct it would seem that the range of E. armeniacuH Elephants fossiles, est celui qui presente le plus de rapport avec
was somewhat the .same as that of Rhinoceros Hlonfordi, i.e. that rElephant actuel des Indes."
itextended from western Asia through the regions lying to the IlisTOitY.- Lortet and Chantre (1872, p. 79) oljscrve:
north of India and China; I am, however, disposed to refer the "Elephas inlermedius (.Jourdan, mss.). Cette csi)ece, (;reec par —
specimen to E. nnnwdicus." M. .Jourdan sur ime forme de VElephas untiquus de I'";dr-on('r, est
1'akki,ki'Has AUMENiACi's KKK. (Lcitli Aiiauis and (1. Busk, c(!lle (jui est la |)lus commune tlans Ic bassin du Rhone lyonnais;
1868, pp. 496 499): "In the British Museum there is a large part elle jjarait meme y avoir etc cantonnee. Les caractercs transitoires
'[See page 1334 below.— Editor.)
 THE MAMMONTIN.E: PARELEPHAS 1063

entrel 'Elcphas primigenius et VElcphas uidiquus, faciles a constater It appears best also to conserve the name Elephas Irogonlherii
sur de nombreuses pieces, ont engage M. Jourdan elever au rang
t\ Pohlig for less progressive specimens from Pohlig's type locality,
d'espoce cette forme, et de lui assigner le nom six'cifique cVinter- especially since according to Deperet, Mayet, and Pohlig this
medins. Ces earacteres sont tres-apparents sur certaines dents; phylum (i.e., "rameau de VEIephas Irogontherii Pohlig") is of long
comme appartenant
cependant, parmi celles qui ont ete considerees duration.
a VEIephas inlennedius, un grand nonibre de pieces peuvent etre TvPK Locality of E. intermedus. In the environs of —
rangces dans la categorie de VEIephas antiquus." Lyons remains of this species are found abundantly in the loess
Validity of the Species Elephas intermedius (Osborn, covering the plateaus (Deperet and Mayet, 1923, p. 182): "Aux
1924). —
Inasmuch as Jourdan's type description (1861, p. 1013) environs de Lyon, il abonde dans le kess des plateaux (Saint-Cyr,
characterizes the species as being closely related to Elephas indiciis, .Saint-Romain-au-]\Iont-d'Or, la Duchere, Fourviere, C'aluire,
and inasmuch as Lortet and Chantre (1872, p. 79) cited the name, Margniole, Croix-Rousse, Saint-Clair, le Vernay, etc.), ou Jourdan
amplified the specific characters, and referred to this species the I'a decrit sous le nom (VE. intermedins, (''est a cette espece qu'ap-
important skeleton of Lyons (Fig. 944), beau sciuelette entier reconstitue au Museum de Lyon,
jjartient le
it appears best to retain the specific et provenant du loess de la montee de Choulans, a Lyon. Les
name Elephas intermedius, atleast molaires du loess de la region lyonnaise representent une \ariete
for the more progressive speci- ou peut-etre une tnutation plus evoluee, a lames un peu plus serrees
mens which occur in (frequence laminaire entre 6, 5 et 7) que dans les molaires d'Alle-
is region of France, magne et d'Angleterre (frequence laminaire 6 Ti 6,5)."
This is the \e\e\ of the type locality of Elephas intermedius
Jourdan, which Deperet states may represent a variety, or perhaps
a more ad\anced mutation with laminse somewhat more con-
tracted, namely, between 6, o, and 7, than the (ierman and English
varieties with laminae 6 to 6, 5. Consequently Deperet regards
Elephas intermedius as more progressive than the typical Elephas
trogontherii of Pohlig.

ReI'EHRED MOL.MIS OF P.\BELEPHAS INTEKMEDIUS, I,VON.S MuSEUM

One-half natural size
I'^ig. 913. Superior molar teeth referred to Elephas inlennedius Jourdan from the collection of the Musevmi de la Ville a J.,yon. MivT |ihot,ographs,

prepared under tlio direction of Doctors Deperet and Mayet (see letter of September 10, 1924).

(Left) This beautifully preserved r.M' exhibits 22 ridge-plates, of which the 12 anterior are worn.
(Right) This first inferior molar, I. Mi, exhibits 12 worn ridge-plates, but the formula appears to l)e (with an additional unworn ridge-plate): l.M 1 fi-
 1064 OSBORN: THE PROBOSCIDEA

Deperet (Translation of a Letter dated Lyons, August .served to bevery distinct from the narrow molars of the Elephas
26, —
The Elephas intennedius was created by Jourdan,
1921). aiilujUHS type,molars resembling in breadth those of Elephas
a former palaeontologist of Lyons, who applied the name, unfor- primigemus, but with thicker and more widely separated lamellEe.
tunately without a figure, to certain broad molars, which he ob- Lortet and Chantre in their memoir on the Mastodons and the
Elephants (Archiv. Mus. Lyon, T. I., 1872, et T. II, 1879) cited
this speciesand figured other pieces which could be related to it.
I believe that E. intermedinsis probably the same animal as that

named by M. Gaudry Elephas prittiigenius var. a lames eearlets and

by Pohlig Elephas trogontherii. We have at Lyons in the University
Mu.seum also in the Municipal Museum numerous molars of this
species and even a complete skeleton discovered in the loess of the
hillof St. Foy at Lyons. It is this complete skeleton [Fig. 944]
which has been figured by Lortet and ("hantre, but in the absence
of figures of the type itself I fear that it may be necessary for us to
adopt the name of trogontherii which appears to have priority.
Deperet and Mayet, 1923, p. 176. Deperet and Mayet, —
\\hile regarding E. intermedius as conspecific with E. trogontherii,
treat this animal as follows: "III Groupe des Elephas tro-
gontherii ET E. PRiMiGENius (Mammouths). Le groupe des
Mammouths comjjrend des Elephants de taille moyenne, au crane
allonge, au vertex tres eleve et etroit en haut (cranes en dome), aux
defenses fortement spiralees, aux molaires gencralement larges et
hautes (type hypselodonte), avec lames nombreuses plus ou moins
Restored Skeleton of Parelephas intermedtus, Lyons Museum serr^es dont les bandes d'email sont paralleles et ne presentent pas
One-sixtieth natural size de sinus loxodontes medians. Nous y distinguerons deux rameaux
Fig. 944. Restored skeleton referred by Jourdan to his species Elephas ayant vecu parallelement depuis le Pliocene superieur j usque dans
inlermedius. The plate was prepared under Jourdan 's direction but was first le Quaternaire: A. Le rameau de VE. trogontherii Pohlig. B.
printed by Lortet andChantre in the frontispiece of their Memoir of 1872[1876J,
Le rameau de VE. primigenius Blumenbach."
from which it is here reproduced one-sixtieth natural size. The height is said
to be 3 m. 75 cm. = 12 ft. 3^ in. Deperet and Mayet continue: "A. Rameau de l'Elephas
(Lortet and Chantre, op. cil., "Elephas intermedtus (Jourdan,
p. 79):
—
trogontherii. Jourdan avait reconnu le premier, voici plus d'un
mss.). — Cette e.spece, cr6ce par M. Jourdan
sur une forme de V Elephas demi-siecle, parmi les molaires de Mammouth de la region lyon-
antiquus de Falconer, est celle qui est la plus commune dans Ic ba-^sin du nai.se, une forme aux lames plus cpaisses, plus ccartees et moins
Rhone lyonnais; elle paralt mpme y avoir oto cantonn6e. Les caracteres nombreuses que dans le Mammouth normal et lui avait donne le
transitoircs cntre V Elephas primigeyiim et V Elephas antiquus, faciles a constater
sur de nombreuses pieces, ont cngag6 M. Jourdan a clever au rang d'especc
nom d'E. intermedius. Lortet et Chantre (1876) n'ont figure de
cette forme, ct de lui a.s.signer le nom spccifique A' inlermedius. Ces caracteres cette espcce que le squclette reconstitue du Iress de Choulans
sont tres-apparents sur certaines dents; cependant, parmi celles qui ont etc qui se trouve au Museum de Lyon et forme le frontispice de leur
coM.sidorces comme appartenaiit a I'Elephas inlermedius, un grand nombre de ouvrage; mais ils n'ont donne malheureusement aucune figure
])i(^cespeuvent ctrc rangces dans la catogorir de I'Elephas antiquus. Dans la
des molaires de VE. intermedius, devenu ainsi une esi)ece purement
vallee de la Saone on a trouvc des ossemeiits d'Elephas inlermeilius, dans Ic
nominale, qui ne saurait etre retenue."
Ichni ou a la Hmite des alluvions, a Lyon et aux environs. (Saint-Rambcrt-
I'llc-Barbr, U: Viriiay, Culuire, la Duchere, Vaisc, la Dcmi-T^une, Saint-Just, Gaudry (1876, p. 40 also PI. ix) and Leith Adams (1877, p. 31)
Fourvicre ct la Quarantaine.) C'cst a la (Juarantainc que M. Jourdan a trouv(^, observesimilar intermediate forms of teeth in Louverne (May(Uine),
en 1862, le squclette dc I'individu qui a pu etre remont6 dans les galeries du France, and in England in the Forest-bed, to the middle of the
Museum de Lyon par les .soins de M. Charles Rf'ivil, I'un de nos proparateurs. Pleistocene.
Cc squelettc, un des plus grands et des plus complets que I'on |)uisse voir en
Europe, mesure 3 metres 75 centimetres de hauteur au garrot. II est surtout
and Mayet, 1923, p. 177): "C'e.st a Pohlig
Finally (Deperet
rcmar(|uable i)ar la courburc tres-prononcec des scs onornies defenses et par (jue nous devons d'avoir distingud, parmi les Elephants quater-
les formes trapucs et ma.ssivcs de ses mcmbres." naires d'AUemagne, sous le nom d'PJ. trogontherii (1889, p. 189),
This animal, discovered in the Pleistocene loe.ss of the hill of St. Foy, une forme particuliere tlu groupe des Mammouths, dont les molaires
I,yons, I'ratice, is regarded by Dei)6ret and Mayct as similar to Elephas ont une couronne large, comme chez les E. meridionalis et primi-
Plateau loess ("provenant du Icess de la niontoe de
trogontherii Pohlig.
genius, mais a\e(^ un nombre total de lames (14 a 22 aux Mm)
Choulans, a Lyon." Deperet ct Mayct). Dcp6ret and Mayet remark (1923,
sujM^rieur a la moyenne de VE. meridionalis (12 a 14), mais in-
p. 176): "Ix)rtet ct Chantre (1876) n'ont figure de cette esp^ce (jue le squclette
reconstitue du loess dc Choulans qui se trouvc au Musrtum de Lyon et forme le fcrieur a celle de VE. primigenius (18 a 27). Cet E. trogontherii
fri)ntis|)irc dc leur ouvrage; mais ils n'ont donno malhcureu.semcnt aucune est d'ailleurs absohiment identicjue a VE. inlermedius dc Jourdan,
figure des molaires de I'A'. inlermeilius, dcvciiu ainsi une cspece purement comme nous avons pu nous en assurer sur les nombrcux sujets du
tmmittale, qvii nc saurait ctrc rctenuc." The skull is said to hv. erroneoit.sli/
hess de la region lyonnaise."
restored at the summit of the occijmt. The rounded summit of the cranium
in the above figure agrees fairly w<-ll with the rounded summit <haracteri.stic
Thus Dei)eret and Mayet fii-st concluded that the Elephas
of Parelephas trngonllierii; the tusks aiJ|)ear to be recurved rather than in- Irogonlherii of Pohlig is absolutely identical with the Elephas inter-
curved as in I'arelcphas jeffersonii. medius of Jourdan, but subsequently stated (p. 183) that it belongs
 . : —

THE MAMMONTIN^: PARELEPHAS 1065

to a more progressive stage, thus agreeing witli Osborn's opinion the Elephas intermedius of Jourdan, namely, M3 If. It is the most
in the present Memoir. phylum, with the possible exception of the
easterly record of this
Geographic Distribution of E. intermedius-trogontherii tooth erroneously recorded from China, as noted above, page 1062.
—
Phylum. Deperet and Mayet {of. cit., p. 180) declare that with This specific stage of Parelephas is certainly more progressive
the exception of the rare molars of the Upper Pliocene of A.stesan, than the P. trogontherii of the 1st Interglacial gravels of Siissenborn,
Elephas trongontherii represents a very distinctive Quaternary a level containing Dicerorhinus etruscus and Equus siissenbornensis
branch or phylum (see Table, p. 1044 above) and now regarded as of Lower Pleisto-

(1) "La distrihulion geographique .s'ctend de I'Est a rOue.st cene age, and the ridge-plate formula agrees closely with that of
depuis la Ru.ssie jusqu'a 1' Europe occidentale et du Sud au Nord P. intermedius of the plateau loess near Lyons. The upper level of
depuis ritalie jusqu'aux lies Britanniques. Laissant de cote les Siissenborn, containing Elephas trogontherii ty]je, is attributed to

details de cette dispersion geographique, nous essayerons seule- the £d Interglacial stage, that is, more recent than II Glacial.
ment de preciser I'etendue de sa repartition stratigraphique dans Pavlow (1910, pp. 6-18), in describing the dentition of

le terrain quaternaire." Elephas wiisti of Tiraspol, bases her description on a series of

In England the species phylum] appears very abun-

[i.e.,
superior and inferior molars (Dp 3-M 3) from the Tiraspol
(2)
dantly at the beginning of the Quaternary period in the Forest-bed
collection of Moscow, excellently figured (as reproduced in our Fig.
945) from photographs in PI. i, figs. 1-12 inclusive. In the same
of Cromer, or Sicilian stage, where it has been described and figured
plate (fig. 23) appears a third inferior molar from Tiraspol in the
under different names by Leith Adams (1877); it persisted in
Geological Collection of the University of Moscow which she
England to higher Quaternary levels, namely, to the gravels of the
refers to Elephas hysudricus.
terrace of 30 meters (Tyrrhenian stage of Deperet, Chellean
Pavlow at first states (p. 7) that the second molars = Dp 3] [
industry)
approach those of Elephas meridionalis of the Val d'Arno (Weit-
(3) In Germany the species E. trogontherii was first' observed
hofer, 1890, PI. xii) and of Elephas antiquus of Weimar (Pohlig,
in the ancient gravels at Siissenborn near Weimar, contemporane-
1891, Taf. 2 bis, fig. 8), but on the final page (p. 18) she con-
ous with the deposits of Mosbach, and of Mauer near Heidelberg;
cludes: "Prenant en consideration que les dents de Siissenborn,
this is 3d Interglarial time of the Osborn-Reeds Table (PI. xxiv)
decrites par Wiist sous le nom de VEl. trogontherii Pohl., se dis-
of the present Memoir; at this time the species appears to be very
tinguent des formes pour lesquelles Pohlig a cree ce nom, nous pro-
abundant in Europe.
posons un nouveau nom specifique de VEl. Wiisti pour nos formes,
(4) In Europe the species [i.e., phylum] E. trogontherii be- celles de Siissenborn, decrites par Wiist, et les formes de Tiraspol,
comes extinct in 3d Interglacial time and is replaced by the true decrites par Mr. Sinzow." It is therein suggested that Elephas
mammoth Elephas primigenius during the period of the Fourth wiisti, a species dedicated to Doctor Wiist, is not remote from the
Glaciation. So far as we can judge
Elephas trogontherii Pohl. of Siissenborn.
(5) In southern France E. trogontherii does not occur in the from Pavlow's figures (PI. i) and description (pp. 6-18) the cotype
Sicilian stage (Durfort, Solilhac, Malbattu, Rozieres) in which molars of either side have the following ridge formula:
Elephas meridionalis is abundant. Parelephas ivilsti: Dp 3 %f^ Dp 4 i& 1 j|^
1 6-1 8
2 15-16 M M
!0-23
RiDGE Formula (Letter, Deperet, September 10, 1924). M3 1 9-20'

"Ces jours derniers, nous avons pu enfin faire avec M. ]\Iayet Parelephas trogontherii: Dp 3 ^ Dp 4 rs M 1 ~r| M 2]~
quelques recherches au Museum de la ville oii se trouvent environ M3
une trentaine de molaires de cette espece. Malheureusement il y —
Geologic Level. (Pavlow, op. cit., 1910, pp. 1 and 4):
en a fort peu qui soient des M' intactes et susceptibles de fournir "Une grande collection de dents et d'ossements d' Elephants fossiles,
des mensurations. provenant du gravier de Tiraspol (gouv. Kherson) et se trouvant
De I'ensemble de mes mesures vous pouvez en toute securite dans le Cabinet geologique de I'Universite de Moscou, a dfl servir
inscrire le chiffre Xoilf^ pour la formule des I\P de VElephas inter- de materiaux pour I'ouvrage present. Le fait est que les dents
. . .

me.dius: done une mutation de passage entre le trogontherii

c'est sont bien representees, elles forment presque une serie complete
et votre E. roosevelti [P. jeffersonii]. J'ai deja eu cette impression et peuvent donner une bonne idee des caracteres generaux du
dans mon Memoire sur les Elephants pliocenes et j'ai ete tres systeme dentaire; tandis que jusqu'a present il n'y a que quelques
sensible a I'eloge que vous voulez bien faire de ce travail." dents isolees de ces depots qui ont ete decrites par Mr. Sinzow sous
le nom d'Elephas trogontherii Pohl. Je commencerai mon
. . .

Parelephas wusti Pavlow, 1909 travail par la description des Elephants de Tiraspol, autant qu'ils
Figure 94.")
m'ont servi de base dans mon etude; apres quoi j'en ferai la com-
Upper Pleistocene gravels of Tiraspol (gouv. Kherson), southern Russia. paraison avec autres formes d'elephants pour determiner leur
les
This most welcome discovery by Pavlow (1909.1- - supple- position dans la ligne du developpement. J'ai deja signale que les
mented by letters of May 31 and August 22, 1924) probably repre- dents d'elephants provenant de cette localite ont ete citees par
sents an Upper Pleistocene stage about as progressive (M3 2-5) as plusieurs savants, Mrs Sokolow, Sinzow, Laskarew, sous les noms
'[See page 1044 above where Soergel lists Parelephas trogontherii as abundant in Mosbach in Isi Interglacial times. —Editor.]
^Original description in Ru.ssian (French translation) "Les elephants posttertiaires de diverges localites en Russie (Annuaire g6ol. et mineral. Russie,
1909, Vol. XI, Livr. 6-7, pp. 171-174, PI. v, figs. 1,2); supplementary de.scription "Les Elephants Fossiles de la Russie" (Nouv. M6m. Soc. Imp. Naturalistes,
Moscou, 1910, Tome XVII, Livr. 2, pp. 6-18, PI. i, figs. 1-12).
1060. OSBORN: THE PROROSriDEA

(U\ers: tantot c'<'tait EL antiquus, mais

El. meridionalis, tantot 6, fig. 1) : "Je commence par les dents qui doivent etre rapportees
dans ces dernicrs temps on est arrive lesrapporter
:'i a 1'^/. irogon- a la meme esjjece. Nous avons dejii indiquo que les premieres
Iherii Polil. et c'est sous ce noni que Mr. J. V. Sinzow les a decrites molaires sujK'rieuros et inierieures (les ;/;', tiii |
= Dp-, DpsI) man-
et figurt'es dans son ouvrage [I''ootnote: 'J. Sinzow. (Jeol.-l'alae- ciuent dans la collection de Tiras|)ol. Pour les deuxiemes niolaires
onth. Peohaeht.in.Siidrussland. 19()0. Odessa. PI. iv, v.']." I
= Dp 3) nous avons deux exemplaires presque identiques, jirivos
de leur derniere lame (PI. i, f. 1).

{Op. cit., p. 13, PI. I, figs. 10, 11, 12): "La w« [M^] fig. 10,
quoique deja bien usee possede encore 7 lames intactes et 13
entamoes par la mastication. Un caraetere assez different nous
. . .

parait presenter une autre molaire, m^ [iVP], f.ll, dans laquelle les
lames sont usees en biais, ce qui les rend plus rapprochees. C'est
une jeune dent; le cement qui devrait
la recouvrir est peu de-
de ses lames sur son cote.
veloping, ce qui jiermet de voir I'epaisseur
Nous trouvons 10 lames sur la surface masticatrice longue de 16
cm., large de 8,5 cm. Le nombre total de lames est 22."
(Op. cit., p. 14, fig. 12): "Sa description [M., fig. 8 of PI. i]

sera faite plus tard, c'est la dent qui nous occupe pour le moment
fig. 12, PL I. C'est une derniere molaire gauche /«6 [Mj], longue de
29 cm. sur sa surface superieure, et de 40 cm. sur le milieu de son
cote interne, ce qui prouve que les lames qui la forment sont rap-
prochees j)ar leiu's parties suijerieures et s'<''cartent en descendant.

CoTYPES OF PaRELEPHAS Wi'STI

One-fourth natural size
Fig. 94,). Cotypo.s (M'^) of Elephas misti Pavlow, 1910, PI. i, fig. 10 (lower right), same as Pavlow, 1909.1, PI. v, fig. 1; figs. II (lower left), and Ua
( upper left). From Tiraspol, southern Ru.ssia. Originals in the Tiraspol Colleetion, llniversity of Moscow.

Elephnit Wii.^ti Pavlow, 1910. "Les Elephants Fossiles de la Les 6 lames anterieures sont les .seules usees, quoique faihlement;
Hussie," Nouveaux Memoires de la Societe Imperiale des Natural- les deux suivantes sont a peine entamees; les 12 qui suivent sont
istes de Moscou, Tome XVII, Livr. 2, pp. 6-18. Cotypes. — intactes."
Series of sujierior and inferior molars (Dp3 -M 3), indicated by CoNCLTisioN, OsBOUN, 1928: would appear that the molars
It

Pavlow as M 2-M 6, in the Cabinet geologique de I'Universite formula

of Purelephas wusti are closely relatetl in their ridge-plale
lie Mo.scou. Horizon and Locality. —Gravels of Tiraspol to the collective formula of P. trogontherii and P. intermedius.
(gouv. Kherson), South Russia. I'pi.er Pleistocene. Cotvpe From a comparison, howe\'er, of the above formula;, it would also
FIGUKE.S.- Pavlow, 1909.1, PL v, figs. 1 and 2; 1910.1, PL i, figs. ai)i)ear that P. wiitsti is a somewhat more progressive species, w ith
1-12 inclusive. Two of the cotypes are shown in figure 945 of the a slightly higher dental ridge-plate formula, than that of P. trogon-
present Memoir. therii. Conseciuently it api)ears 1o be closest to the P. intermedius
SiPPLK.ME.NTAKV Desckiption. — (Pavlow, op. cH., 1910.1, p. stage of .hjurdan.
 :

THE MAMMONTIN^: PARELEPHAS 1067

3. NORTH AND SOUTH AMERICAN SPECIES OF PARELEPHAS

Migration from Eurasia.— As shown above, the Parelephas phyhim becomes known in the Upj^er PHocene'
of Italy in Parelephas trogontherioides (M 3 [^tI^); it reappears in the 1st I iiterglacial stage of the Cromer
Forest Bed, East Angha, in P. trogontherii ref.? (M 3 vr|—see Fig. 871); it is very abundant in the 2d Interglacial
of Slissenborn near Weimar (M 3 \^, type —see p. 1056) ; it continues in P. intermedius of southern France
(M 3 -see p. 1062); again it is found in P. wiisU of .southern Russia (M 3 M—see p. 1065 above).
During the 3d Interglacial period it apparently migrated from western Eurasia, but future discoveries may reveal
its presence in eastern Eurasia in northerly latitudes. The next record is in the state of Washington in Parelephas
washingtomi (M 3 ^), with the least progressive ridge formula thus far observed in America; it then becomes very
abundant, under favorable conditions of the Middle United States, in Parelephas jeffersonii (M 3 It), appearing to
culminate in the highly progressive P. progressus (M 3 M). Meanwhile the giant Parelephas Jloridanus (M 3 f-fl)

appeared in Florida.

we have briefly described the steps taken by Osborn toward the separa-
In the opening section of this chapter
tion of species of the genus Parelephas from Mammonteus primigenius, on the one hand, and from the true Elephas
columbi of the southern Unitetl States, on the other. To oiu- jiresent knowledge the order of description of the
American species has been as follows (see p. 1047 and fig. 933)

1838
1857-
loas OSBORN: THE PROBOSCIDEA

The third species, Elephas texianus, first named (1859) by Owen and hitor (1801, 1802) described by Rlake, is

treated in the present Memoir as a synonym of Parelephas columbi.

The fourth species named, but neither described nor figured, is the 'Elephas' Indianapolis or inississippiensia
of Foster, 1872, an indeterminate species probably referable to Parelephas.

Fortunately the fifth species described, namely, Elephas {Parelephas) jeffersonii Osborn, 1922, is based upon
a superb type specimen (Fig. 900) which had previously been erroneously described by Osborn (1907) as Elephas
colu)7ibi, and redescribed by Hay (1914) as Elephas primigenius. This genotypic species is now the most com-
pletely known of all the extinct proboscideans, for the aged grinding teeth (Fig. 959) of the senescent type specimen
arc sui)plemented by superb paratype' specimens (Figs. 900, 907, 892), also by very large numbers of teeth and
jaws collected from the great geographic belt of the 40th parallel northward occupied by species of this genus.

The type ridge formula of Parelephas jeffersonii, in comparison with that of other American species, is given
on ])age 1070 below.

Osborn would gladly surrender priority in specific nomenclature to Mather, if Mather's type should be
located and compared. It appears from the above ridge-plate formulae, actual and estimated, that if we are able

to locate the type of Parelephas jacksoni (indirectly named after President Andrew Jackson), we may be able to
determine that it is conspecific with Parelephas jeffersonii (named after President Jefferson). In the same presi-
dential line of nomenclature is also Elephas = Parelephas]
[
roosevelti Hay, 1922 (named after President Roosevelt),

herein shown to be a synonym of Parelephas jeffersonii. Finally, Parelephas ivashingtonii was named after the

first President of the United States. It will be observed that P. waskingtonii is apparently the least progressive
in its ridge formula, namely, M 3 7,,, which is similar to that of P. intermedius of southern France, namely,

M 3 1^1, also to that of P. wilsti of southern Russia, M3 ^o, as shown in the comparative table above (p. 1048).

Parelephas jacksoni Matlior, 1838

Figure 946

Ploistocene, Jackson County, Ohio.

Compare Elephas intlianapolis or tnississippiensis Foster, 1872; also
Elejthas (Parelephas) jeffersonii Osborn, 1922-1924.
Compare Leidy, 18(19, p. 399, who suggested that if E. jacksoni were
adopted, to be correct, it should read E. jacksonensis.

Mather, 1838, is interesting as the earliest

Elepluis jncksoni
A. Lower jaw of the Elephant found B. Lower jaw of the fossil
name given any species of American mammoth, even antedating
to at Jackson. Elephant.
EleTphas americamis De Kay (1842) and E. columbi Falconer (1857).
The figure and description indicate that this is a valid species,
which will be recognized if the type can be located; Hay states
(letter dated March 14, 1922) that in his examination of the
Pleistocene mammal collections of the United States he has not
been able to locate this type jaw. Mather's description of E.
jack.suni was reviewed by Falconer (1863, p. 57) and the species
regarded as indeterminate. Hay (1923, p. 147) also regarded the Type of Parelephas jacksoni
si)ecies as "wlujlly indeterminable." Juvenile ty|)e jaw (A) of Elephas jacksoni Mather, 1838, from
Fig. 94().
E. [Elephas] jacksoni Mather, 1838. "First Annual Report on .laekson County, Ohio, as distinguished from a lower jaw (B) referred by
the (leological Survey of the State of Ohio," embodying Report of Mather to Elephas primigenius. Type specimen not located at present.
AKso (C) supi)lementary drawing of r.Mj exhibiting sixteen worn ridge-plates.
('. 15riggs, .Jr., I'ourth .\ssistant (ieologist, pp. 90, 97, with geologic
Reproduced after Mather's type figure (1838.3, p. .363).
.section (1838.1 — Fig.
947 of present Memoir); review of First
A young jaw exhibiting the riglit and left second inferior molars and
Annual Report on the (leological Survey of Ohio, quoting portions a few ridge-i)lates of tlie third inferior molars, unc^vcniy worn on the opposite
of Ueijort by ('. Hriggs, .Jr., Amer. .Journ. Sci. and Arts, Vol. sides, consequentlj' exhibiting an unetpial number of worn ri(lK<'-plates.

'Teehnieally ideotype (see Sehuehert, 190.'j.2, p. 1.')), beeau.se strictly .speaking a paratype must be one of the specimens mentioned in the original descrij)-
tioii.
 —

THE MAMMONTIN^: PARELEPHAS 1069

XXXIV, p. 358 (1838.2) "Remarks ; in addition to and explanation

of the Review of the Rejiort of the Geological —
Survey of Ohio in
a letter to the Editor," Amer. Journ. Sci. and Arts, Vol. XXXIV,
pp. 362-364 (de.scription, name, and figure — 1838.3). Type. —
Lower jaw from Jackson County, Ohio. Type Figure. —
Op. cit., 1838.3, p. 363, fig. A, also supplementary figure on the
same page of the crown of r.M2, exhibiting 16 worn ridge-plates.
This type jaw, according to the following citation from Mather
(op. cit., 1838.2, p. 358) appears to be part of the skeleton found on
"one of the branches of Salt creek, in the northwest part of Jackson
county," Ohio:
"Among the most interesting details of this report [Report of
C. Briggs, Jr., to the (leological Survey of Ohio, on the work of
J. W. Foster and him.self], are those respecting the fossil elephant

discovered during the past season. As there is some doubt as to

the geological position of the deposits in which these bones are
found, we will extract the description."
'About two years ago, some bones, so large as to attract the
attention of the inhabitants, became exposed in the bank of one of
the branches of Salt creek, in the northwest part of Jackson county.
They were dug out by individuals in the vicinity, from whom we
obtained a tooth, a part of the lower jaw, and some ribs. In the
examinations at this place, during the past season, it was concluded
to make further explorations, not only with the hope of finding
other bones, but with a view of ascertaining the situation, and the
nature of the materials, in which they were found. The explora-
tions were successful. There were found some mutilated and de-
cayed fragments of the skull, two grinders, two patellae, seven or
eight ribs, as many vertebrae, and a tusk. Most of these are nearly
perfect, except the bones of the head. The tusk, though it retained
its natural shape as it lay in the ground, yet, being very frail, it

was necessary to saw it into four pieces in order to remove it.'

'The following are the dimen.sions of the tusk, taken before it
was removed from the place in which it was found:

Length on the outer curve,

1070 OSBORN: THE PROBOSCIDEA

(Amer. Jour. Sri., vol. XXXIV, p. 362, figures) for a lower jaw of Calvin, 1911, p. 213 (footnote) states: " Elephas indianapolis,
an elephant, found in Jackson County, Ohio. This jaw is, however, a new species of fo.ssil elephant, by .1. W. Foster. Proceedings of
from the deserijition anil the tifjure, wholly indeterniinable." the American Association for the .\dvancement of Science, twenty-
Referrkd to P.^kelkphas (l)sborn, 1924): Osborn observes meeting, held at Dubucjue, August, 1872, p. 259. The title
first

that the type figure (Fig. 946 A, C) is sufficiently characteristic, onlyis printed in the Proceedings. After the reading of the paper,
accompanied by Mather's quaint description, to distinguish this and at the suggestion of some of the naturalists present, Foster
jaw from that of Mammonteus primigenius and to relate it rather proposed to change the name to Elephas mississippiensis, and
to Paielephas. Allowing for the primitive wood engraving, the under this designation there is a short reference to the paper and
rostrum (a) of Klephasjacksoni resembles quite closely the rostrum the proposed species in "Nature," Vol. VI, ]). 443.
of Parelephas jeffersonii paratype (Amer. Mus. 13225) shown in "Nature," 1872, September 26, p. 443: "Prof. ,1. W. Foster
figure 960 B of the present Memoir; this rostrum would be more of Chicago, read a paper on what lie considers a new species of
l)rominent in an immature jaw like the type of Elephas jacksoni
fossil elephant, called by him Elephas Missi.ssippiensis. He
than in a mature jaw.
presented to the association a fossil tooth found in Indiana, and
The second feature of note is the fact that M2 is still in use and
which he regards as differing specifically from that of any other
that M.I is coming into use; this individual is therefore younger
fossil elephant found in America or on the C/ontinent. The differ-
than either of the paratypes of P. jeffersonii (cf. Fig. 960). The
M ences are so great that he holds them sufficient to constitute the
numberof ridge-plates .shown in this wood engraving is 2,:i^7no+
M3r7;ir6+ (incomplete) which agrees fairly well with the ridge- new species."

plate formula of M 2I'^J" assigned to Parelephas jeffersonii.

' Calvin, 1911, p. 213:"At the meeting of the American Asso-
Consequently it appears probable that Elephas jacksoni is related ciation for the Advancement of Science, in Dubuque, in 1872, the
to the genus l^irelephas rather than to the genus Mammonteus. writer had the privilege of seeing the tooth, which Foster regarded
It would be of the utmost interest to discover this young type jaw as the type of a new species of fossil elephant, Elephas indianapolis,
and to determine more closely its actual relationships. The tusk and of hearing his description of it, and one at least of the distin-
measurements cited above from Mather (outer curve 10 ft. 9 in.) guishing characters of the supposed new species was the looping of
accord in size with those of the type of Parelephas jeffersonii the enamel from the middle of the grinding surface to the sides,
(outer curve 11 ft. 4% in.), since the type of P. jacksoni belongs to
instead of the usual arrangement of this tissue in elongated ellipses
a much younger animal than the type of P. jeffersonii, an aged bull.
surrounding plates of dentine. In the bibliography and catalogue

Parelephas mississippiensis (?) Foster, 1872 of the fossil vertebrata of North America, by Hay, Bulletin of the
(?)

From Indiana, exact locality not recorded. United States Geological Survey, No. 179, page 714, the tooth de-

Elephas indianapolis Foster, 1872. "Elephas Indianapolis. scribed by Foster is assumed to be one of the forms of molars of

A New Species of Fo.ssil Elephant." Proc. Amer. Assn. Adv. Sci., Elephas pri migenius."
August, 1872, p. 259. Osborn, 1929 : Genus and species indeterminate.

THE COLUMBIAN MAMMOTH (PARELEPHAS COLUMBI)

The second member of the Parelephas phykim to be described, figured, and named was the 'Elephas columbi'
of Falconer, 1857, representing a specific stage which has become widely Icnown among tdl palaeontologists as the

Columbian Mammoth. As narrated in the early part of Chapter XVI, Falconer beheved this animal to l)e iden-

tical with Leitly's 'Elephas [

= Archidiskodon\ imperator.' Osborn for several years treated it under the genus
Archidiskodon but as a species quite distinct from A. imperator; finally Osborn, with the accession of the new and
rich materials now to be described, perceived its true phyletic relationship to the phylum Parelephas. It is, how-
ever, clearly separable from Parelephas jeffersonii, being much more primitive in its ridge formula, as shown in the

following comparison of the five known outstanding American species of the genus Parelephas.

Parelephas progressus M 3 f^ Parelephas jloridanus M3ii-±

Parelephas jeffersonii M2 M3lf Parelephas columbi M3 ^
Parelephas washingtonii M3 IfT

The low ridge formula of the true Parelephas columbi of Georgia tmd South Carolina (M 3 —;) is very surpris-
ing and significant; this low ridge formula of P. columbi appears to prove that its ancestors migratetl into America
early in Pleistocene time. This theory of the early geologic entrance into America of rclat ivel y primitive species of
 THE MAMMONTINiE: PARELEPHAS 1071

Parelephas is supported by tlic primitive character of the lower jaw of P. wasMngtorm (Fig. 972), in which the
type and referred ridge fornuda is M 3 M; as pointed out below, the ridge formula of P. washingtonii agrees quite
closely with that of P. intermedius (M
§^") of southern France. Taken altogether this evidence appears to
3
indicate that the ancestors of P. columhi and of P. imshingtonn may have passed across Europe and Asia and
migrated far southward in North America during the M
and even possibly during the 1st Interglacial period,
following the migration wave into America of Archidiskodon.

Parelephas columbi Falconer, 1857, 1863, 1868 characters. "Subgen. 3. Ei'elephas Spec. 12. E. (Eueleph.)
. . .

Figures 887, 935, 936, 948-902, 954, 955, PI. xxii Columbi . . . Post-Pliocene? . Mexico; tieorgia; Alabama.
. . . . .

Type: Upper(?) Pleistocene, Brunswick Canal, near Darien, Georgia. A Syn. E. Jacksoni? (Sillim. Journ. 1838, vol. xxxiv. p. 363)."
Of this description Falconer observes (Nat. Hi.st. Rev., 1863,
Referred: Upper(?) Pleistocene, South Carolina, Florida, and Mexico.
p. 45) "Thus, the leading points of the dental characters, and the
:

SjT).: Elephas texianus Owen, 1859 (name); Blake, 1861 (name), 1862
(definition and figure).
precise place in the natural series occupied by the species, were
distinctly indicated, together with its range of habitat along a
Specific Characters (Osborn, 1924). — (1) Type and neo- stretch of nearly 20" of longitude in the regions bordering the CJulf
type ridge-plates: 3 M (2) Ridge-plates typically 6J2 in of Me.xico." Type and Locality. — (Falconer, op. cit.,

100 mm. (3) Third superior and inferior molars relatively short 1863, p. 43) : "My first
knowledge of this form dates from the year
anteroposteriorly and deep vertically, corresponding with a hypsi- 1846, when Sir Charles (then Mr.) Lyell submitted to me, for ex-
cephalic or acrocephalic cranium. (4) Ridge-plate height of third amination, some fossil mammalian remains, which he had brought
superior molar, M^, 145 mm. (min.), 207 mm. (max.); of third with him on his return from his second visit to America [Footnote:
inferior molar, M3, 147 mm. (min.), 166 mm. (max.). 'Second Visit to N. America, 3rd edit. 1855. Vol. i. p. 347.']. They
(5) The ridge-plate formula of the third molars, compared formed part of a collection which had been exhumed in 1838-39,
with that of primiti\'e species and of more progressi\e species in digging the Brunswick Canal, near Darien in Georgia. A
of Arrhidi.skodon. may be written as follows: selected series of these remains was presented by Mr. Hamilton
Couper, the discoverer, to the Academy of Natural Sciences of
Upper Pleisto- Parelephas columhi M 3 1 5-16+' thin ce-
Philadelphia, where they were identified by Dr. Harlan, some of
cene ment outer coating.
Lower Pleisto- Archidiskodon iniperator ^I 3 .ll'llo heavy ,

cene cement outer coat-

ing.
Upper Pliocene Archidiskodon meridionalis M 3 Y^fn. moderate
to Lower cement outer coat-
Pleistocene ing.
Upper Pliocene Archidiskodon planifrons M 3 Y^, no cement
outer coating.

This species, clearly established by Falconer in 1857 and sub-

sequently described in great detail (1863) from a type molar found
in 1838-1839 while digging the Brunswick Canal, near Darien,
Georgia, has had a most checkered history and synonymy. There
is no question as to the validity of Falconer's type, type locality or
region, and type description; 'Elephas columbi' is a clearly distin-
guished species of the southern L^nited States. With this species
other specimens have been sadly confused by Cope, Osborn, Hay,
and others; in fact, 'E. columbi' is one of the most sadly misused
names in the American proboscidean literature. Let us firet turn
to Falconer's type description and to his application of the type
characters in a number of referred specimens from Texas and Type Figdre of Parelephas columbi
Mexico. Fig. 948. lypeoi Elephas colu7nhi¥aXconfiV, 1857. After Falconer, 1868,
Vol. II, PI. X, Middle portion of the third right lower molar, r.Ms,
fig. 1.

HISTORY: LYELL, FALCONER, OWEN, BLAKE, longitudinally and vertically bisected. From the Brunswick Canal, near
Darien, Georgia. Charles Lyell Collection, 1846. Brit. Mus. 40769. Cast
OSBORN (1846-1924)
Amer. Mus. 1747. (Lydekker, 1886, p. 173) "The greater part of the second
:

[third] right lower true molar which has been longitudinally and vertically
E. (Eueleph.) Columhi Falconer, 1857. "On the Species of
bisected; from the Pleistocene of the Brunswick canal, Darien, Georgia.
Mastodon and Elephant occurring in the fossil state in Great
Described and figured in the 'Nat. Hist. Rev.,' 1863, p. 52, PI. i. and in the
Britain. Part I. Mastodon." Quart. Journ. Geol. Soc. London, 'Pateontological Memoirs,' vol. ii. pp. 221, 222, PI. x. fig. 1. Presented 61/
Vol. XHI, 1857, synoptic table opposite p. 319, including dental C. Falconer, Esq., 1867."
1072 OSBORN: THE PROBOSCIDEA

whose determinations were corrected by Professor Owen, and those spondingly open interspaces filled with thick masses of cement.

of the latter more recently by Dr. I.eidy." These characters were strongly in contrast with the attenuated,
(Lydekker, 1886.2, Pt. IV, ]). 173): "The greater part of the parallel, and pectiniform disposition of the materials seen in molar
second lower true molar which has been longitudinally
[third] right .sections of E. primigenius; combined with the dilated outline of
and vertically bisected; from the Pleistocene of the Brunswick the 'discs of wear,' and the decided crimping in the i)lates of enamel,
canal, Darien, Georgia." Brit. Mus. 40769. Cast Amer. Mus. they led me to regard the form as occujjyiiig a j^lace in the series
1747. Type Figure. — Falconer, op. cit., 1863, pp. 52, 114, between E. antiquus and E. Indicus, and as differing more from the
PI. reproduced in the "Palaeontological Memoirs," Vol. II,
i; Mammoth than does the latter from the existing Indian Elephant."
1868, PI. X, fig. 1. (Explanation PI. i, 1863) "Section of the middle [6] These fact.s were epitomized in the table of Ajiril 8, 1857, in

portion of an adult lower molar, of Elephas Columbi, from the which E. columbi is included in the same group as E. indicus.
post-pliocene deposit of the Brunswick Canal, near Darien, in [7] "Sir Charles Lyell's Georgian specimen, from the Brunswick

Georgia (i). 52); showing the disposition and relative j^roportions Canal, upon which my first knowledge of E. Columbi was founded,
of the ivory, enamel, and cement, as compared with corresponding consists of the middle portion of the penultimate or last true molar,
sections of E. Indicus, and E. prirnigenius, contained in the Fauna probably the latter (m.3) lower jaw right side, broken off, both at
Antiqua Sivalen.sis, PI. i. (Nat. size)." the anterior and posterior ends. The fragment comprises ten
complete ridges, with part of two others, of which the anterior
seven are more or less worn. All the fangs are broken off, together
with the basal mass of ivory. The summit of the crown is concave
from back to front, and the tooth is also concave with a little
torsion on the outside, and convex inwards, showing that it was
considerably arcuated laterally, like the specimen last described.
The discs of wear are of moderate width, as in the Indian Elephant,
with a tendency in some of them to expansion in the middle. This
is most pronounced in the second, where the expansion nearly

attains half an inch. The plates of enamel are thicker than in the
1/4 ml tui
Mammoth, and about equal to those of the Indian Elephant;
they present a considerable amount of parallel shallow plaiting,
Restored Tvpe of Parelephas coldmbi which is prominently shown where they rise above the level of the
Fig. 949. Type r.Ms of Elephas columbi, internal aspect, redrawn for the cement. The wear of the crown takes place in a succession of steps,
l)resent Memoir from Briti.sh Mu.seum cast of type (Amer. Mus. 1747).
from the front backwards, which importance to notice with
it is of
.Anterior and posterior plates restored from neotype (Amer. Mus. 13707).
reference to the inferred food of the species. These steps rise like
Compare Osborn, 1922.505, p. 1, fig. 1.

This fractured tyjie agrees exactly in size with a complete thini inferior a flight of stairs, each being composed of the whole mass of cement
molar found in the phosphate beds near Charleston, South Carolina, which of one of the valleys, and the combined enamel plates and ivory
contains seventeen ridge-plates, three of which belong in front and four bcliind of the ridge immediately behind it. There are fi\'e of these steps in
the ridges 4 to 13 contained in the type specimen, showing that the full number The
the (ieorgian specimen, the posterior ridges being intact.
iif ridges in M3 is probably seventeen.
[type] dimensions are as follows:

Type Desckiption (Falconer, 1863, pp. 43-52).— "[1] The [Falconer] [( )sborn
specimens brought by the latter [Sir Charles Lyell] included some
fragments of the molars of a fossil Elephant, which, after careful 9.5
examination, I satisfied my.self belonged to a .si)ecies wholly distinct
from the prevailing fossil form of North America, namely, E.
priuiigt'itins; .[2] I apjilied to it, in my notes of a systematic
. . .

classification of the Proboscidia [referring to his description of

1857], the designation of E. Columbi, after the great discoverer;
.
[3] proving the co-existence of a distinct species of Ele])hant
. .

with the extinct Edentate Fauna of the Southern States of the

Union." [4] Confirmed by other specimens from Mexico, Texas,
and other of the Southern States of North .\merica. [5] "I resorted
to the crucial test of sawing u]) the principal molar of the Brunswick
Canal series longitudinally and vertically, in the manner figured
in the plates devoted to the Elephants, in the 'I'auna Anticjua
Sivalensis,' a procedure which commonly quashes at a glance all
doubts as to the specific distinctness, or otherwise, of Elephant
molars, in critical cases. The section yielded roUiculi, showing
rather thick plates of enamel folded upon cimeiform cores of ivory,
of very considerable width at their base, and separated by corre-
 : —

THE MAMMONTIN^: PARELEPHAS 1073

near Ihc btise, (he excessive tliickness of six-tenths of an inch, vegetable diet." (Blake, The Geologist, Vol. IV, 1861, i). 470):
being about twice as much as what is ordinarily seen in the section "South of the 30th degree of N. latitude it [(the Mammoth) E.
of the Mammoth. For the contrasted diiTcrcnce, I refer to the primigenius], however, gives place to a totally different species of
"
sections, pi. i, fig. 1, of the 'Fauna Antiqua Sivalensis.' true Elephant {Elephas Texianus, Owen, E. Columbi.' Falconer),
—
Falconer, 1863. In the introductory remarks to his invalu- the molars of which, by their less degree of complexity, were more
able paper of 1863 cited above. Falconer treats most fully the adapted to triturate the soft succulent herbage of Texas and
dentition, habits, and range of Elephas columbi and associated Mexico."
fossil mammals. He then considers the origin and range in time of Blake, 1862.— "Molar tooth of Elephas texianus (N.S.),"
the mammoth {Elephas primigenius) and the persistence of its after figure by Charles Carter Blake, in The Geologist, 1862, PI. iv,
distinctive characters, followed by a discussion of the unity or ]). 57, in his article "On a Fossil Elephant from Texas {E.
plurality of the species of the existing Indian elephant {Elephas Texianus)." This tooth appears to be the one referred to by
indiciis). Finally he gives the fullest exposition of the food of Owen in September, 1858, and designated by Blake as follows
living and extinct species, E. indicus, E. africanus, E. primigenius. (Blake, 1862, p. 58) : was a distinct species
"This opinion [that it

In this paper P'alconer classifies the species of elephants by ridge of elephant, closely allied to the Indian type] was confirmed by
formulse, grouping those with a high ridge formula in Euelephas, Professor Owen, and after the name of Elephas Texianus had been
those with a low ridge formula in Loxodon. given to the species, the specimen was deposited in the British
Estimated Ridge Formula. — In this species Falconer finally Museum, and now forms one of the most conspicuous objects in
(1863, p. 56) estimates the ridge formula, based on referred speci- the gallery devoted to Proboscidea. Professor Owen, in September,
mens some of which certainly belonged to Elephas imperator, as 1858, thought fit to adopt the name of E. Texianus for the species,
follows in his eloquent address to the British Association (and also in the
second edition of 'Palaeontology,' p. 395). This the only
E. columbi: Dp 2| Dp 3f Dp 4ft M lit M 2-ff M 3§™. specimen which have seen of this type, as Dr. Falconer has not
I
. . . is

Falconer's formula (1863) was shown by Osborn (1922.555, p. stated where the specimens are on which he described his species.
2) to be jDrobably too high. Osborn's formula of Ms
(1929) is given above in the definition of the species and is

discussed below in the analysis of the Cohen Collection.

—
Synonymy. Falconer {op. cit., 1863, p. 67) finally
concludes: (1) That his Elephas columbi of 1857 includes
in part specimens described by American palaeontologists
as Elephas primigenius; (2) that the Elephas le.rianus
Owen is a synonym of it; (3) that Elephas imperator
Leidy is synonym, and that until a perfect molar is
also a
figured and described, no satisfactory opinion can be
formed as to what E. imperator is. Owen-Blake Elephas texianus Type [ = Parelephas columbi ref.]

Elephas TEXiANUs Owen, 1859, Blake, 1861, 1862. Fig. 9.50. Type right third inferior molar, r.Ms, of Elephas texianus Owen, 1859,
Blake, 1861, after Blake, 1862, PI. iv. Scale not given by Blake, but estimated at ap-
Elephas tej-ianus <)wen-Blake, given as a name only by
proximately one-third natural size from dimensions given by Falconer, 1868, Vol. II, p.
Owen in 1859, was defined by Blake in 1862 (1862, p. 58),
223, namely, exteme length of crown 12..j in. or 318 mm., width at eighth disc 3.8 in. or
as follows: "The figure by Mr. Mackie [Fig. 950 of the 99 mm. These dimensions somewhat exceed those of Osborn's neotypc (Amer. Mus.
present Memoir] gives a better idea of its appearance than 13707), length 298 mm., breadth 91 mm.
any mere \erbal description. I however define it as Ele- (Lydekker, 1886, p. 172): Brit. Mus. 33218, "The well-worn third right lower true
molar; from [San Felipe de Austin, on the Brazos River, Texas]. Described and figured
phas texianus, denlium molarium (m.6), colliculi undulati,
by Blake in the 'Geologist,' vol. v. p. .57, j)!. iv. (as E. texianus), and also described
magis remoti quam in E. Indico." It is antedated, how-
by Falconer in the 'Nat. Hist. Rev.' 1863, p. 52, and in the 'Palsontological Memoirs,'
ever, by Elephas columbi Falconer, and was the subject Vol. if. p. 222. Purcliased. About 1858." Known as the "Bollaert molar." Estimated
of an animated controversy between Falconer, Owen, and ridge-plates 17-I-.
Blake.
Elephas texianus Owen. "Address of the President." Brit. He appends as a doubtful synonym, 'E. jacksoni.', Silliman's
Assn., 1859, p. Ixxxvi. "Geology tells us that at least two kinds Journal, 1838, vol. xxxiv. page 363'; but after examination of the
of Elephant {Mastodon Andium and Ma.st. Humboldtii) formerly did very bad drawings contained in that page, I cannot make any
derive their subsistence, along with the great Megatherioid beasts,
distinction between them and E. primigenius. The tooth of E.
from that abundant source [tropical America]. Nay more; at
Texianus (ra.6, lower jaw) has enamel-folds much wider and much
least two other kinds of Elephant {Mastodon ohioticus and Elephas
more waved and undidated than that of the E. Jacksoni. The
texianus) existed in the warm and temperate latitudes of North
canals of cement are consequently of much greater width, and the
America." Name only. (Owen, 'Palaeontology,' 2d Edit., 1861, p.
whole aspect of the tooth is much more like E. Indicus."
395, (juotcd by Falconer, Nat. Hist. Rev., 1863, p. 46) ". where : . .

it [the Manuuoth] existed not only with the gigantic Mastodon Osborn, 1924: This figure and description appear to confirm
Ohioticus, but also with a second species of true Elephant {Elephas Falconer's opinion that Elephas texianus Owen, 1859, Blake, 1861,
texianus, Blake) the teeth of which were adapted to a succulent 1862, is a synonym of E. columbi Falc.
 I

1074 OSBORN: THE PROBOSCIDEA

OSBORN (1922) SEPARATES PARELEPHAS COLUMBI present communication is to clear up this confusion and to propose

FROM PARELEPHAS JEFFERSONII' Elcphas jeffcrsonii as a new species of American Pleistocene

Consistent with the researches on which the present Memoir mammoth. .We thus find by the characters of the type
. .

throughout has been based. Osborn reexamined Falconer's type and neotype specimens [see Fig. 951 of the present Memoir]
sjxicimen and description and realized that in common with Cope, that the real Elcphas columbi is not the animal we have been de-
I lay, and other American palaeontologists, he had previously erred scribing under this name; it is a dwarf form, perhaps a dwarf

in confu.sing this with an entirely diiTerent species of fossil elephant, female, of the animal which we ha\e been describing under the
which, as a consequence, he (Osborn, 1922.55.5. jip. 1-3) separated name Elephas impcralor."

E. columbi
O.axn (.Dm caM o, lyp* Am«r. Mu^ 1747
MiowJ Ifom *™». Mu» 13707 tiwj

E. columbi
An.*.. Mu» 13707

N«o(yD*

Neotype
Amer. Mus. 13707 (rev.
<outer view'

I"iG. 951. Falconer's Type (A) .\nd Osborn's Neotype (B, C) of Elephas [Parelephas] columbi
One-fourth natural size
(I.,oft) Crown view of tlie tyi)e, r.M.^ (upper figure) and neotype, I.M3 (lower figure) of Elephas columbi (east Amer. Mus. 1747). (.\) Shaded portions
represent ridge-plates 4 13 wliieh are jireservcd in the type, ridge-plates 1-3 being restored from the neotype. (B) Crown view of neotype^ a complete
third lower molar of the left side, I.M3, .showing ridge-plates l-l.i and rudiment of ridge-plate 16 [observe that in the restored type specimen (Fig. 949) the
estimated ridge-plate eount is 16-t- (17).] .\fter Osborn, 1922.555, p. 2, fig. 2.

(Right) Neotype molars of Klrphns columbi, one-fourth natural size, from the phosphate beds of South Carolina (Amer. Mus. 13707). Two indi-
viduals. (C) External view of r.M', eighteen ridge-plates plus a probable posterior one and a half ridge-plate. Bl, External view of I.M3; same tooth as fig-
ure (B) op|K)site, with fifteen complete ridge-plates and an incipient sixteenth plate. After Osborn, 1922.555, p. 3, fig. 3.

as Elrphan jcjfer.sonii iind subsequently (Osborn, 1924.6.'i.3, p. 4) as The affinity between Parelephas columbi and Archidiskodon
Parelephas. imperator he found by no means so close as Falconer belie\ed; the
(Osborn, (1922. .555. pi). 13) : "The present article relates
di.stinctions have been pointed out in the full description of A.
explicitly to the type characters of Elephas roli(mhr. of E. impcralor,
imperator above. Prior to, and independently of, Osborn's obser-
and of the .\merican sjjecimens referred to E. primigeiiius, three
sjxicies which have become more or loss confused in all the previous
vations, Freudcnberg had revived Falconer's oiiinion as to the

literature liecauso the characters of the tyiK' specimens ha\o not- do.se relationship of the true Elephas columbi of Mexico and the
been precisely detcrminetl and compared. The object of the true E. imperator.

'At this time (1922) Osborn placed 'EUphas columbi' within the phylum .irchidiskodon: in the present Memoir it is placed witliin the phylum
PareUplias.
 THE MAMMONTIN.^: PARELEPHAS 1075

TYPE CHARACTERS The variations in the ridge-phite count in 10 cm. and tlie
Consequently the reestablishment of the type and neotype compression depend ujion the level of the crown from which the
characters of E. = Parelephas] coltimbi is based upon the follow-
[
ridge-])late count is made and approximate Parclephas columbi to
ing materials: Arrhidiskodon imperator with which it was long confused. P.
Falconer's type of Elephas [
= Parelephas] columbi, r.Ms (Brit, columbi is much more distinct from P. jeffersonii, which has a higher
Mus. 40769, cast Amer. Mus. 1747). Cieorgia. ridge-plate formula and a higher ridge-plate compression, namely,
Owen-Blake type of Elcphas lexianus, r.Mj (Brit. Mus. ridge-plate formula M3ft, ridge-plate compression 7 (min.), 9 [11X>|
33218). Te.xas. (max.), according to the level of the crown from which the com-
Osborn's neotypes of Elephas [
= Parclephas] columbi, r.M^, pression is counted.
I.M3 (Amer. Mus. 13707). Phosphate beds of Charleston, S. C. They show, as illustrated in hgure 949, that in Falconer's type

Number of
Superior Molars Specimens
R.M'-L.M' — one-third worn, one-half worn, two-thirds worn, greatly worn, very aged, complete or fragmentary,
preser^'ing from 8-18K ridge-plates 14
R.M--L.M- — one-third worn, one-half worn, complete or fragmentary, preserving 10-13}^ ridge-plates 6
R.M'-L.M' — one-third to one-half worn, preserving 10)^-11}^ ridge-plates 3

Inferior Molars
R.M3-L.M3 — one-third to one-half worn, adult, aged, young, preserving from 6-16 ridge-plates 7
R.M2-L.M2 —one-third worn, two-thirds worn, aged, fractured, preserving from 9-12}2 ridge-plates 5
R.Mi-L.Mi —one-fourth worn, one-half worn, preserving from 10^-12)4 ridge-plates 2
R.Dp4 —one-fourth worn, preserving ridge-plates lO)^ 1

Total number of grinding teeth 38

Thirty-six inferior and superior grinding teeth of Elephas molar three ridge-plates are missing in front and four to five ridge-
[
= Parelephas] columbi (Amer. Mus. 13708 a-z and 13709). plates are missing behind. Thus the type and neotype ridge
The type and neotype molar characters of Parelephas columbi formula of Parelephas columbi is as follows: M ^Y^rri+-
may be summarized as follows:
-p g. OSBORN ESTABLISHES THE CHARACTERS OF PARELEPHAS
T^AT ,- 1. COLUMBI BY THIRTY-EIGHT NEOTYPE AND REFERRED
R.M3 l.e -I
rKlge-i)lates,
,1 a
11 worn, 6 unworn, 6 c m
•
i„
10 cm.,
MOLARS FROM THE PHOSPHATE BEDS OF
breadth 82e mm., length 241e mm., index 34e. SOUTH CAROLINA
Neotypes: See Figure 952 of the present Memoir
R.M^ 19;^ ridge-plates, 8 worn, 10-K unworn, o}i-(i% in 10 -pj^^ ^,^^y Cohen Collection from the phosphate beds
extensive
cm., breadth 96 mm., length 249 mm., index 39.
„e^r Charleston, South CaroHna, presented to the American
L.M3 15-16 ridge-plates, 9 worn, 6 unworn, 4)2-5)2 in Museum in 1908 by Mrs. Morris K. Jesup, includes a complete
10 cm., brcadtli 91 mm., length 298 mm., index 31. series of thirty-eight upper and lower grinding teeth to which the

Cohen Collection: Number of Ridge-plates

Amer. Mus. No. Total Worn Unworn in 10 cm. Breadth Length Index Height Remarks
13707 (Neotype) R.M^ 19);e 8 10+ 5^2-6)^ 96 249 39 207 9tii pi. One-fourth worn, pre-
serving 18 }2 ridge-
plates. Finely pre-
served.
13708-e L.M2 -|-12K 11 6-7 92 197K 47 106 8th pi. One-half worn, preserv-
ing 11?4 ridge-plates.
Finely preserved.
13708-m R.M' +11K +10 }i 7-8 77 169 45 One-half worn, preserv-
ing 10>o ridge-plates.
1747 (Type cast) R.M3 17e 11 6 6 82e 241e 34e 152e 10th pi. One-third worn, preserv-
ing lOJi ridge-plates.
13708-i L.M2 +12\e 11 1)4 5}i-6}i 74 210 35 Two-thirds worn, pre-
serving 12)2 ridge-
plates. Very aged.
13708-k K.Mi 13 12 1 7)2-8 67 167 40 One-half worn, preserv-
ing 12)^ ridge-plates.
Finely preserved.
13708-n IJ.Dpi )2-ll-)2 9 4 8-10 49 133 37 91 0th pi. One-fourth worn, pre-
serving )j~ll-)^ ridge-
plates. Finely pre-
served.
1076 OSBORN: THE PllOBOSClDEA

following numbers have been assigned: Anicr. INhis. 13708 a-z the i)hylogenetic table above (p. 1048); (3) on the contrary, in
(35 speeiniens in all); and Anicr. IMus. 13709; also Amer. Mus. the Arehidiskodon phylum the inferior ridge-i)lates exceed the
13707, a third inferior molar anil a third superior molar chosen by superior ridge-plates in number, e.g., A. imperator (M 3 iJ), A.
the present author (1922.555, pp. 2, 3) as the ncotyjjes. planifrons (M 3 y1+)i ^- meridiotmUs (M 3 Jivfj); (4) as shown
These thirty-eight specimens agree entirely in their generic in the section of Falconer's type (Figs. 948, 949, 951), the arcuate
and specific characters, thus positively establishing the grinding ridge-plates are w'idely separated in mid-section, thus, in extremely
tooth characters of Parelephas columbi; the largest specimens are worn grinding teeth of Parelephas columbi the ridge-plates appear

the males, the smaller specimens are the females or dwarfed males. to be as far apart as in ^4 . imperator.
The summary of the Cohen Collection is as above (p. 1075). From these exceptionally fine materials subjected to the most
—
Constant Ridge Formula. From an examination of these careful study and comparison by the author and Dr. C. C. Mook,
fourteen third superior molars, the prevailing ridge-plate formula tiie ridge formula of Parelephas columbi may now be written as
(M 3 i'/."iV+) agrees very closely with that of the molars in the follows:
Amherst specimens (Fig. 954), namely, M
3 ff^, and appears to Dp 4 js:YT4i M
»-ll-'/i ^" 1
' -,% M" -!^,^
12M "^ 2 M
3 15-16 +
+12»5 ^" "
1 8-1 ft
-

firmly establish four facts: (1) The low ridge-plate formula of the The maximum and minimum measurements of the larger

third superior molars is typical and constant; (2) there are more (male) and the smaller (female) grinding teeth are probably due to
(
ridge-jilates in the third superior than in the third inferior molars sexual differences; certain of the "ohen specimens agree exactly in
— a characteristic of all known species of Parelephas, as shown in size with Falconer's type. The details are as follows:

^Iin Min.

M'
M2
Ml
Ms
Ma
M,
Dp4
 — :

THE MAMMONTINJi;: PARELEPHAS 10 I i

Laminah Frequency. — The ridge-plates are typically diver- Simon's Island the canal pa,ssed through Six-mile Swamp. This is
gent at the unworn summit of the crown (5/2-6)2 laminae in 10 connected at its northern end with Altamaha River, at the southern
cm.), more contiguous as we descend toward the base of the crown with Turtle River. The swamp has thus the appearance of a lake
(6-7K) laminse in 10 cm.); in mid-crown the ridge-plates are set which has become filled with alluvial deposits. These consist of
widely apart as in Archidiskodon imperator, more widely apart a compact clay, usually yellow- and impregnated with iron. There
than in Parelephas jeffersonii. In all degrees of age and wear are thin strata of soft, chalky marl and many fragments of petrified
the minimum laminar frequency is 5}i in 10 cm.; the typical wood. At the bottom of this deposit were found the bones of
laminar frequency is 6]4 in 10 cm., the maximum laminar frequency Megatherium, Elephas, Mammut, Equus, and Bi.wn. Beneath the
is 8^2 in 10 cm. The above variations accord with the point at clay stratum was sand with marine shells."
which the measurement is taken. This explains why certain of From Hay's faunistic tables of 1914.1, 1923.1, and 1924.1,
these fractured, broad-plated grinders have been mistakenly re- the following species may be selected as characteristic of the
ferred to Archidiskodon imperator by Hay and others, while certain Upper Pleistocene life zone, to which Parelephas columbi probably
of the narrow-plated molars have been referred to Parelephas belongs
jeffersonii. The grinders (Fig. 954) of the Amherst skeleton belong
to the narrow-plated variety. See also Florida specimens of P. Fauna OF THE Type Locality Fauna of the Phosphate
floridanus, (Nat. Mus. 11808, 11806, 11810), page 1079 below. OF Elephas [= Parelephas] Beds near Charleston, S.
COLUMBI NEAR DaRIEN, C, where Elephas [= Par-
PARELEPHAS COLUMBI ABUNDANT IN THE SOUTHEASTERN Brunswick Canal, elephas] columbi OCCURS in
UNITED STATES Georgia abundance
This Hay's interesting account of the geology and fauna of
is
Parelephas columbi type Parelephas columbi ref.

the important and distinct type of Elephas columbi Falconer. Megatherium mirabile type Ocalientinus (Serridentinus)

(Hay, 1923.1, p. 157): "1. Brunswick, Glynn County [Georgia]. obliquidens type

This is the type locality of Elephas columbi. This species was based Mastodon americanus ref. Mastodon americanus ref.

by Falconer (Quart. Jour. Geol. Soc. Lond., XIII, 1857, table Equus Ifrateimus] leidyi type Bison latifrons ref.
opposite p. 219 [319]) on a part of a tooth received from the Bison latifrons (?) ref. Tapirus haysii ref.

geologist Charles Lyell and which had been found in the Brunswick
Canal. The specimen consisted of 10 median plates of a lower Upper Pleistocene Age. —Besides the above and numerous
second or third molar. Falconer figured it in 1868 (Palaeont. Mem., other species, Hay many specimens referred
(1923, p. 363) includes
vol. II, pp. 214, 221, plate x). Lyell (Second Visit, etc. vol. i, p. found in South Carolina but not definitely
to several other species
348) noted that an elephant had been found in excavating the recorded from the "Charleston phosphate beds," for example,
canal. Richard Harlan, in 1842 (Proc. Acad. Nat. Sci. Phila., Archidiskodon imperator ref., which may have been washed in or
vol. I, 189), stated that a large collection of bones of various
p. dredged from an older horizon.
aniiiials had been presented to the Academy by J. Hamilton Hay, 1914, 1923.— In his Memoir of 1914 and again in 1923
Couper, of Darien, Georgia. Among these were teeth of E. primi- Hay grouped together as one species, namely, "Elephas columbi"
genius. Couper, in 1848 ([1846.1] Hodgson's Memoir, etc., p. 45), (Hay, 1914, pp. 410-421, also 1923, pp. 430, 431, map 12), the
stated that two lower jawbones with teeth, several loose teeth, mammoths now separated by Osborn into Parelephas columbi
two tusks, and several vertebrae of Elephas primigenius had been [southeasterly range in Fig. 953] and Parelephas jeffersonii [norther-
collected in the canal during 1838 and 1839. These remains quite ly range insame figure]. Those in the northerly range probablj'
certainly belonged to Elephas columbi unless possibly some belong- includesome specimens referable to Mammonteus.
ed to E. imperator. Leidy (Jour. Acad. Nat. Sci. Phila., vol. vii, Osborn, 1928: Osborn determines as Parelephas columbi two
1869, p. 254) records the presence in the collection of the Academy of the grinding teeth from the Charleston phosphate beds of

of a lower molar of E. columbi. The present writer has seen in this South Carolina and from Ocala, Florida, described and figured by
collection parts of four teeth of this species which had been sent Hay, 1914, p. 413, PI. lxi, as follows:

from the Brunswick Canal, doubtless parts of the Couper collection.

The species are listed on page 369. . .
[p. 369] The most striking was Dp2 (Ocala, Florida), see Leidy, Trans. Wag-
the great ground-sloth, of the genus Megatherium, and which ner Inst. Sci., Vol. II, p. 17, PI. iii, figs. 6
I.«idy afterwards called Megatherium mirabile ... .[p. 370] Elephas and 7, also Hay, 1914, p. 413, PI. lxi, figs.

columbi (p. 157). Mammut americanum (p. 120). Bison [laii- 2 and 3.

Jrom?\ sp. indet. (p. 261). . . . E. [Equus fraternus] leidyi (p. Up' (Ocala, Florida, fide Leidy), length 110
193). . Megatherium mirabile (p. 36)."
. . mm. ; ridge-plates 8-9 (see Hay, op. cit., PI.

{Op. cit., p. 370): "J. Hamilton Couper (Hodgison's Memoir, LXI, figs. 5, 6).

))p. 37-40) has given an account of the topography and geology of Nat. Mus. 1614 Dp' (phosphate beds. South Carolina), length
the region through which the Brunswick Canal was being con- 101 mm., height fourth ridge-plate 94 mm.;
structed (map 40). On one of the plates of the work is a section ridge-plates 8-|- (see Hay, op. cit., PI. lxi,
from the ocean westward 21 miles. About 10 miles west of St. fig. 4).
1078 OSBORN: THE PROBOSCIDEA

PARELEPHAS COLUMBI' OF FLORIDA Mus. 11620), an aged individual recently described by Gidley (sec

Flokiim. — Parelephas cohimbi is fairly abundant in Florida; ]iage1005 above). The true P. colnmbi, relatively abundant, agrees
few specimens of Archidiskodon imperator arc recorded
relati\-ely very closely in size and ridge-j)latc formula with Falconer's type
from Florida, the most perfect being the \'enice mammoth (Nat. from Georgia (Fig. 948) and with Osborn's neotype from the South

Kansan
77 ? 7

Doubtful

DriftlesB

NORTHETILT CIRCLES = DISTRIBUTION OF PaRBLEPHAS JEFFERSONII AND MaMMONTEUS

piiiMiQENius. Southeasterly circles = Distribution of Parelephas
COLtrMBI AND ArCHIDISKODON IMPERATOR

FiR. !).")3. RiitiRo of I'nrdrphns jrffrrsonii and P. cnhimbi, iiicludiiig the type lociility
(I) of /'. cohimbi in ('irorniii, .'ilsd Wxc. nrotypc loi'iility (3) fniiii tlio i)lio.spliato IhmIs
of South Carolina. Modified from llay (l()2;j, map 12, j). llil), wlio erroneously
treated I 'arclc phas jcffrnnmii and /'. cobinihi as belonging to the same .species.

'Since this Florida .section was written (I92S) the more |)rogre.ssive Parrlepkasfloridanussiasr, M3 has been described (O.sborn, 1930.837).
The grinding teeth (Nat. Mu.s. ll.SOG, 11808, 11810) belong to I'.Jloridanus (op. cit., p. 17).
 THE MAMMONTIN^: PARELEPHAS 1079

Carolina phosphates (Fig. 951). Recently described by Gidley,

from Melbourne and V'ero, Florida, presumably from the "No. 2"
bed of Sellards, are three P. rolumbi^ molars:

Nat. Mus. 11808. Third superior molar, r.M', adult; laminar

frequency 6-6K ridge-plates in 10 cm.
Total ridge-plates 19-20, 13 preserved
(tooth not complete).

Nat. Mus. 11806. A second superior molar, 1.M-; laminar

frequency 8 ridge-plates in 10 cm., +12 +
ridge-plates preserved.

Nat. Mus. 11810. A third superior molar, r.M', aged individ-

ual ; laminar frequency 7-8 ridge-plates
in 10 cm. Total ridge-plates 18 to 20,
15 preserved, 3 to 5 missing.

Ciidley (.June 14, 1928) observes that while in Parelephas

columbi ref. (Nat. Mus. 11808) the laminar frequency is only 6
measured on the outer side and 6% measured on the inner side of the
crown, in 10 cm., at the same level, Nat. Mus. 11806 (P. rolumbi
ref.) exhibits a laminar freciuency at the base of nearly 8 ridge-
plates in 10 cm., the compression being much greater. Can such
a difference be accounted for on the ground of individual variation?
(iidley (letter, December 6, 1928) observes in Nat. Mus.
11806 {P. columbi) 12 ridge-plates plus 7 to 10 missing, a total of
19 to 22, hence an M^
in Nat. Mus. 11810 (P. columbi) he ob-
serves 15 worn ridge-plates ])lus 3 to 5 missing, a total of 18 to 20.
Loomis (1923-1928) collected in the No. 2 horizon, latest
Pleistocene, near Melbourne, Brevard County, Florida, for the
Amherst Museum, the invaluable specimens now assembled in the
mounted skeleton (Amherst Mus. 25-1) of the typical Parelephas
columbi represented in our figures 954 and 955 and fully described
below.

AMHERST SKELETON OF PARELEPHAS COLUMBI

One of the most fortunate tliscovcries in the recent history of
mammalian palseontology is that of the Amherst skeleton referred
to Parelephas columbi, found in 1923 and exhumed in December
of that year. It is recorded from the Sellards "No. 2" horizon,
latest Pleistocene, Brevard ('ounty, Florida, near Melbourne.
Through Prof. Frederic B. Loomis of Amherst College it came
into the possession of the Amher.st Museum; it bears the number
Amherst ^lus. 25-1. The materials were assembled under the
direction of Professor Loomis, and mounted, as shown in figure 955,
about one fifty-sixth natural size.

—
Typical of P. columbi. The beautifully preserved third
superior and inferior grinding teeth in the Amherst specimen
(Fig. 954) agree very closely in their ridge-plate formula (M 3 J-|^)
with the typical Parelephas columbi of Georgia and the phosphate Molars of the Amherst Skeleton (cf. Fig. 955)
beds of South Carolina, in which the prevailing ridge-plate formula Fig. 954.Superior and inferior molars found in incomplete skull of
isM3,J^^. Parelephas columbi (Amherst Mus. 25-1). About one-third natural size.

Skeletal Chauactkrs. — Professor Loomis writes (cf. letter,

(Upper
displaying
figures)
18+
Third left

ridge-plates, of
superior molar, l.M', crown and internal views,
which 15+ are worn. Height 206 mm.
July 28, 1928): In regartl to our skeleton of Archidiskodon
(Lower figures) Third left inferior molar, I.M3, crown and external views,
[Parelephas] columbi the four posterior grinding teeth [Fig. 954] Length 318 mm. =
displaying 10+ ridge-plates, of which 12+ are worn.
and about 4 feet of each tusk are original, but most of the skull is 12)^ in.

'ISee footnote on opposite page. — Editor.]

 :

1080 OSBORN: THE PROBOSCIDEA

restored. column, excelling four dorsals, and all

All the vertebral var. silvestris, El. Colundii var. Falconcri. and KL Citliindji \ar,

of the ribs, excei)ting a few gaps, were found together. Of the based on lyiie grintling teeth. ()sl>orn conse(|uentlv
inijirndor.

forelinib, the right .'icapula is complete and half of the left scapula; describes these four subspecies under I'arclephas tolumbi [El.
both humeri were i)resent, the right humerus lacking the head. Colunibi var. Felicis) and Archidiskodon imperator, giving facsim-
The left radius and ulna were complete and fortunately the carpals, ile rejiroductions of the broad-plated type figures directlj- after
inetacar})als, and phalanges of the forefeet, excepting three digits. Freudenberg. We also reproduce herewith (Fig. 901) the figure of

()f the iielvic region, the ischium and pubes, and the lower portion a 13-plated grinding tooth after a jihotograph loaned by Senorita
of the ilium, including some 12 inches above the acetabulum, are Reyes, referable to A. imperator.
])resent but the upper part of the ilium is restored. The hindhmbs,
,
Osborn, 1928: Probably members of this piiylum migrated
including the femora, tibiae, and left fibula, are complete, also the into Mexico in Upper Pleistocene time, long after A. imperator
lower end of the right fibula. had become extinct.
very interesting to give the principal measurements in
It is —
Reyes, 1923. The Proboscidea collections from the Valley of
comparison with those of Archidiskodon im-peralor (Amer. Mus. Mexico were described by Senorita Alicia E. Reyes in her paper
10598) and of .4. imperator maibem (Neb. Mus. 5-9-22): entitled "Los Elefantes de la Cuenca de Mexico," 1923, p. 227

NEW FORELIMB ESTIMATES OF PARELEPHAS AND ARCHIDISKODON

P. columbi
Amherst ]\Ius. 25-1

Ft. In. Mm.

Complete heiglit at shoulder from sunnnit of scapula
to base of manus, as mounted 11 3430
With fully extended manus
—
I'oRELiMB Vertical diameter
Right scapula 3
Right humerus
Left radius
Left ulna
Manus and carpus fully extended
Manus and carpus, as mounted
—
IIiNDLiMB Vertical diameter
Right femur
Right tibia
I'ibula
( '()nii)letc length of himllindj, acetabulum to base of
mounted
pes, as
The above comiiarative measurements
scapula and noiu'al spines of tiuterior dorsals

Skeletal slioulder height

Hciglit in the flesh
 :

THE MAMMONTIN^: PARELEPHAS 1081

Amherst Skeleton of Parelephas coldmbi from near Melbourne, Florida

About one fifty-sixth natural size

Fig. 955.Materials for the mounted skeleton of Parelephas columbi, bearing tlie number Amherst, Mus. 25-1, were collected by Prof. Frederic B.
Loomi.s of Amherst College in the No. 2 horizon, latest Pleistocene, Brevard County, Florida, near Melbourne. While the specimen wa.s found in 1923 and
Taken up that December, it did not come into the possession of the Amherst Mu.seum until 1925.
Skull mostly restored, also portions of both tusks, four dorsals, parts of the limb bones, ilium, manus, and pes; otherwise largely original (see below).
Superior and inferior molars shown in figure 954.

of Chiconautla, and in Villa de Guadalupe. Her observations relat- and a discussion of the observations of Leidy, Osborn, and Falconer.
ing to localities and to E. columbi are as follows, those relating to She dismisses the observations of Barcena and Castillo (1882-1884)
E. impeialnr are presented in abstract above. as to the human remains being contemporaneous with the mam-
moth at Tequixquiac, from which locality both P. columbi and
Elephas [
= Parelephas] columbi.
.4. imperator are recorded. It is not .stated whether P. columbi is
Escuela de Ingenieros No. 2. R.M' with 18 ridge-plates, 14 in geologically more recent than .4. imperator, as probably the case.
is
25 cm. ; length 280 mm., breadth 110 mm., height 200 mm. From Reyes writes (June 12, 1924) : "The httle data which we have
Tequixquiac. Fig. 10, p. 236. concerning the stratigraphy of the Valley [of Mexico] seem to
Instituto Ceologico (without number). R.Ms with 15K ridge- confirm your hypothesis that E. columbi is more recent than E.
plates in 25 cm.; length 290 mm., breadth 102 mm., height 180 imperator; but they are insufficient. It is not possible to correlate
mm. From Zumpango. Quaternary strata of the Valley with "las Nordicas," because
Instituto tieologico 213. Fragment and erroneously
of skull glacial periods or extremely cold periods do not exist. Diaz
restored left tusk; also both third molars, length of r.M^ 350 mm., Lozano informs me that the flora is constantly tropical. All the
breadth 115 mm., height 190 mm. Total ridge-plates 19)21 16 in examples which I have examined seem to me to belong to the two
25 cm. From Tequixquiac. Fig. 11, p. 237. species columbi and imperator. More intensive study would
Instituto Geologieo 214. Inferior mandible with I.M3; 16 to perhaps permit a subdivision into varieties."
17 ridge-plates in 25 cm. ; length 268 mm., breadth 90 mm., height —
Faunal Succession. A brief summary of our present knowl-
190 mm. From Guadalupe. Fig. 12, p. 238.
Villa de edge of the Proboscidea and other mammals of this region in
Escuela Ingenieros No. 3. L.jVP, with 21 ridge-plates, 20 in Upper Pliocene and in Pleistocene times is as follows
25 cm.; length 275 mm., breadth 88 mm., height 190 mm. From
Tequixquiac. Fig. 13, p. 239. Pleistocene Parelephas columbi Tequixquiac, Zumpango,
:

Villa de Guadalupe.
GEOLOGIC AGE OF THE PROBOSCIDEA FAUNA OF MEXICO Archidiskodon imperator: Tepexpan, Tequix-
See Cope, 1884.2, "The E.\tinct Mammalia of the Valley of Mexico," also quiac, Zumpango, Villa de Guadalupe.
Furlong, 1925, "Notes on the Occurrence of Mammalian Remains in the
Pleistocene of Mexico," etc., and Villada, 1903, "Apuntes Acerca de la Fauna Pliocene or Cordillerion tropicus (cf. Mastodon humboldtii).
F6sil del Valle de Mexico." Pleistocene Cordillerion oligobunis Cope, Tequixquiac.
Reyes, 1923.— This author concludes (1923, p. 239j with Cordillerion oligobunis progressus Freudenberg,
a comparison of Parelephas columbi and Archidiskodon imperator Canyon of Aculcingo.
 :

1082 OSBORN: THE PROBOSCIDEA

Pliocene Cordillerion oligobunis aiiiiquissimus Freuden- El. Columbi var. Felicis Freudenberg, 1922. "Die Siiugetier-
berg, Hidalgo, Valley of Amajaque. fauna des Pliociins und Posti^liociins von Mexiko," Ceol. und
Pliocene Cor(llllei-ioii{'!) uligobunix felicis Freudenberg, Palaeont. Abhand., N. F., 1922, Band XIV, Heft 3, pp. 147-
Puebla. 152. Type. —Third right superior molar, r.Ml Original in
Pliocene or Cordillei-ion{1) oligobunis i7iter7nediiis Freuden- Leipzig (Pal. Coll. Univ. Leipzig 4403), Steppenform.
('oil. Felix.

Pleistocene berg, Mexico. Horizon and Locality. — Tecamachaico, Puebla, Mexico. Pleis-
Hipparion caslilli [Pi'otohippus] Cope. tocene. Type Figure. —
Op. cit., Taf. viii (xvi), fig. 4:
Pliocene Rhynchoiherium. tlascalx Osborn, type from "Elephas Columbi var. Felicis nov. forma."
Valley of Mexico, neotype from Sonora. —
Type Description. (1925.1, p. 148): "Ein von Tecama-
chaico . . . im Staate Puebla stammender Molar, No. 4403 der
Other Plei.stoecne and Upper Pliocene faunas are the following paliiontologischen Sammlung der Universitat Leipzig, gleicht eben-
(Cope, 1884, Villada, 1903. Furlong, 1925) falls sehr einem solchen \on El. primigenius, unterscheidet sich
jedoch durch eine etwas bedeutendere Dicke der Schmelzlagen
Tpfiuixfiuiac: Equns Owen], E. tau Owen, E.
[lurvidens und ist vielleicht schon der folgenden Art zuzurechnen; doch
crenidens Cope, E. barcensei Cope, E. occi- Das gleiche
stehen die Schmelzbi'ich.sen ausserordentlich gedrangt.
deritalis Feidy, ref. gilt fiir einen ebenfalls in unserer Sammlung
Zahn von befindlichen
Bison laiifrons Harlan, ref. Ejutla im Staat Oaxaca [Footnote: 'Unsere Textfigur 19a und
CameJops hestermts Leidy, ref. 19b. El. Columbi var. Herr Prof. Dr. Felix hatte die
silveslris.'].

Platygomis {'!)conipresstis Le Conte, ref. Liebenswiirdigkeit, mir aus dem

^Museum der
paliiontologischen
Caprovwyx mexicana Furlong. Universitat Leipzig diesen Zahn neben einigen anderen aus IMexiko
jEnocyon dims.
Teleoceras {'!)fossiger Cope, ref. [Pliocene].
Near Monte- Glyplodon [mexicanus].
rey: Nothrotherium.
Neotoma, near occidentalis, ref.

Furlong concludes (1925.1, p. 152): "[1] The mammalian

remains obtained at the four localities, Tequixquiac, Zumpango,
Saltillo,and Monterey, in central and north-central Mexico, are
indicative of the largeand varied mammalian faunas that will be
found in the Pleistocene deposits of Mexico when intensi\e work is
carried on. [2] The two types of deposits recognized in the quarry Type ok Parelephas coldmbi felicis
at El Tajo, near Tequixquiac, and the mammalian forms found in Fig. 956. Type, r.M', of Elephas Columbi var. Felicis Freudenberg, 1922,
each of these accumulations, indicate apparently two distinct Taf. VIII (xvi), fig. 4, one-half natural size. From Tecamachaico, Puebla,
stages in the Pleistocene history of this region. [3] The presence of Mexico. Original in Leipzig (Pal. Coll. Univ. Leipzig 4403), Coll. Felix.

Mnocyon dims, a coyote-like form, and of Capromeryx mexicana Observe that of the 14+ ridge-plates exposed by wear there are 9}i in
10 cm., slightly exceeding the maximum number attributable to this species as
in the brecciated fissure deposits suggests a faunal horizon compar-
noted in the specimens of the Cohen Collection from the Charleston phosphate
able perhaps to that found at Rancho La Brea, California. The beds.
assemblage of mammals thus far found in the beds overlying the
fissure deposits indicates a somewhat later horizon in the Pleisto-
cene. [4] The paucity of mammalian remains found in the cave-
zur Untersuchung zu senden. Es handelt sich bei dem Elefanten-

Cerro de la Silla offers little opportunity to add

fissure deposits in
zahn von Tecamachaico um einen M^
Er des rechten Oberkiefers.

to our knowledge of the upland mammalian fauna of the Pleis- tragt die Katalognummer 4403 des dortigen Museums.
Der Zahn
tocene or to make comparisons with the faunas from the Cali- hat aussen weisse Farbe, welche auch das Innere aufweist, und
fornia caves. The presence of Nothrotherium sp., Equus sp., and
Diese Art der Erhaltung
oberfliichlich einzelne Hiimatitflecken.

Neotoma, near occidentalis, a form related to the recent wood rat, begegnete mir nur bei einigen mitteldiluvialen Fossilien, z. B.

are close to the types found in Samwel Ca\e, Shasta County, von San Luis."
California."
—
Characters. Freudenberg {op. cit., pp. 147-152) points out
the confusion by Cope and Felix of the races of Elephas columbi
Falc. with those of E. primigenius; he also points out the charac-
Parelephas columbi felicis Froudcnberg, 1922
ters of Falconer's type of E. columbi; following P'alconer's estimate
Figure 9.5(j
formula of E. columbi as:
of 1863, p. 56, he gives (p. 141) the ridge
Pleistocene, Tecamachaico, Puebla, Mexico M 1 T#, M 2Tf, M 3f-oi|. He concludes (p. 145) that E. columbi

Freudenberg's excellent figure and description of the type, Vsxlc. is he then proceeds to describe
related to E. imperator Leidy;
a number of races or varieties of these species of "Elephas columbi,"
an r.M'',which he named El. Columbi var. Felicis, enables us to
relate this type to the genus Parelephas, a species probably more of which E. columbi var. Felicis is the first mentioned he distin- ;

progressive than the typical P. columbi. guishes this subspecies by its closely compressed lamellae as most
 :

THE MAMMONTINiE: PARELEPHAS 1083

nearly resembling E. primigenuis, in contrast to a second subspecies coarse enamel, deejjly grooved or crenulated on the sides; the
El. Cloumbi var. Falroneri (cf. A. iinperntnr) in which the lamellae external cement, formerly pre.sent, has been dissolved or worn
are less com])ressed. away; the apices of the three ridge-jilates j^resent a convex
Osborn, 1930: The compressed ridge-plates (see
relatively hence .supporting the superior molar reference indicated by
profile,
legend of Fig. 956) together with the estimated ridge formula of the posterior concavity. They correspond broadly with ridge-
M 3 VoT^ niay range this subspecies Parelephas columbi felicis be- and 18 of an r.M' of Parelephas columbi; in size they
plates 16, 17,
yond the limit of the typical P. columbi (M 3 j^) into the ridge- correspond with ridge-plates 19, 20, and 21 of Jloridanus; they
plate estimates of the more progressive species P. Jloridanus seem relatively narrow, thus agreeing with the narrow ridge-
(M 3 H^) or P. jeffersonii (M 3 f|). plated P. columbi rather than with the broad-plated Archidiskodon
imperator.
Geographically the nearest representative of Parelephas
Parelephas columbi cayennensis Osborn, 1929
columbi cayennensis to the north is the Parelephas columbi felicis
Figures 957, 979
Freudenberg, 1922, from Puebla, described on the opposite page
Cayenne, French Guiana, South America. Probably Upper Pleistocene.
1082 and illustrated in figure 956 of the present Memoir, in which
As on page 1046 of the present chapter, the sole
set forth the posterior ridge-plates are broader, with le-ss coarse enamel, than
elephant so far known
to have reached South America is a sub- in the present type specimen.
species of Parelephas columbi reported by Captain Perret and
Type of Parelephas columbi cayennensis Osborn
successively mentioned by Lartet (1859), Lull (1908), and Freuden-
berg (1922), located in the Marseille Museum by Doctors Laurent AZ.
and Repelin in 1929, and finally described by Osborn (1929.797, p. £
20) as Parelephas columbi cayennensis.
characterized this specimen as
Lartet (1859,
"un fragment de molaire a lames
p. 500)
Vf
epaisses," which led Freudenberg (1922, pp. 159, 160) to refer it to
El. [Archidiskodon] imperator, from which it jiroves to be positi\'ely «^ « f
POST,

distinguished by the very narrow molar crown as described below.

Parelephas columbi cayennensis Osborn, 1929. "New Eurasi-
atic and American Proboscideans." Amer. Mus. Novitates No.
393, December 24, 1929, pp. 20, 21. Type.— "Three and
a half ridge-plates of a third right superior molar, r.M', collected
by Captain Perret in Cayenne (French Guiana), South America,
Fig. 957. Three a.spects (Al, A2, B)
and now preserved in the Museum d'Histoire Naturelle, Marseille,
of the fragmentary type, r.M', after cast
France, as No. 8449 (cast Amer. Mus. 21933)." Horizon (Amer. Mus. 21933) taken from type speci-
—
AND Locality. Probably Upper Pleistocene. Cayenne, French men in the Marseille Museum No. 8449.
Guiana, South America. Type Figure. Op. cit., p. 21, fig. — One-half natural size. Compare Osborn, POST

20. 1929.797, fig. 20 (Fig. 979 of the present

—
Specific Characters. The superior type fragments, photo-
Memoir).

graphs and casts of which have been kindly furnished the present Al, A2, Lateral and inferior or root
aspect of supposed ridge-plates 16, 17, and
writer through the courtesy of Director W. Laurent of the Museum
18-K.
d'Histoire Naturelle of Marseille, and Prof. W. Repelin, Conserva-
B, Crown view of ridge-plate, partly
teur, is characterized by Professor Repelin (translation of letter of
worn, exhibiting thick enamel.
March 30, 1929) as follows: "They were in rather bad condition
and so badly cemented together that I had to take away the larger
part of the cement. One of these fragments, A-1, is represented in
side view on one of these photographs. The lamellae are very worn
and they show the plate of blackish or brownish enamel rather in Parelephas jeffersonii Osborn, 1922
relief and the ivory forms a small median depression. The cement
Figures 806, 810-814, 818, 892, 930, 931, 93.5-937, 958-968, 975,
which separates the lamellae is not very thick in this specimen. In
988, 1009, 1029, 1084, PI. xxii
the other photograph, A-2, the specimen is shown as seen from
Upper Pleistocene, post(?)-Wisconsin, IV Glacial till {fide Levcrett and
above. Finally another fragment, B, is also represented as seen Hay, 1923); Jone.sboro, Indiana.
from abo\e. It has been worn in the direction of the lamellae, but Syn.: Elephas roosevelli Hay, 1922. Compare also Elephas [Parelephas]
this worn part shows only a very irregular section of ivory and jacksoni Mather, 1838, which may prove to antedate Parelephas jeffersonii.
enamel."
This well-known and
\'ery important specific stage, genotypic
The fragment consisting of three and a half lamellae or ridge-
of Parelephas, widespread throughout the Middle LTnited States
is
plates is made the type of the new subspecies Parelephas columbi
and gives us a com])lete knowledge of the skeletal and dental
cayennensis. These ridge-plates appear to belong to the posterior
characters. The ridge-plate formula is
portion of the crown of a third superior molar of the right side,
r.M'; they are strongly concave posteriorly and are composed of Dp 3 ^ Dp 4 +1 M 1 M 2 ^^ M 3 ft
1084 OSBORN: THE PROBOSCIDEA

History of the XaiME.— This species is very abundant in the part of the skeleton (Fig. 10), found near .lonesboro, Indiana, on
latter part of the Pleistocene period in the northern United States the farm of Dora S. [E.] Gift; purchased for the ^luseum with the
and southward into Mexico; the type may
even be of Postglacial Jesup Fund in 1904, restored and mounted in 1906; described by
age; it is the animal erroneously described by Cope and Osborn as Osborn in 1907 as Elephas columbi, determined [and described] by
"Elephas roltimbi," and by Hay as "E. priinigenius," and finally Hay (1914) as Elephas primigenius."
separated by Osborn as Elephas jeffersonii. It now becomes the (Op. cit., p. 15) : "Cranial Characters. — Still more obvious
most fully known species of American mammoth. This species are the differences between the relatively long, broad, and shallow
was described by Osborn July 8, 1922. Hay subsequently (Sept. crania of E. jeffersonii and the relatively short, narrow, and deep
;5(). 1922) proposed the name Elephas roosevelli. Mather in 1838 crania of E. primigenius, proportions which are correlated respec-
gave the name Elephan juckf<uni to a similar animal, the type of tively with the corresponding proportions just described and figur-
\\ hicli has been lost rendering the species indeterminate at jiresent. ed in the teeth."
In 1924 Osborn made Elephas jeffersonii the genotypic species "The four complete skulls of this species known to the writer
of the new genus Farelephas. are those of (1) in the type mounted skeleton (Amer. Mus. 9950);
Elephas jeffersonii Osborn, 1922. "Species of American Pleis- (2) the fine male skull associated with the jaws and a large part of
tocene Mammoths. Elephas jeffersonii, New Species." Amer. the skeleton (Amer. Mus. Cope Coll. 8681) from Whitman County,
Mus. Novitates No. 41, July 8, 1922, pp. 11-16. Type.— Wa.shington, and labeled 'Elephas columbi' [.since referred to Par-
Skull, jaws, and greater part of skeleton (Amer. Mus. 9950). elephas washingtonii]; (3) the young male skull (Amer. Mus. Cope
—
Horizon and Locality. Upper Pleistocene, post(?)-Wisconsin, Coll. 14475) from Dallas, Texas, also labeled 'Elephas columbi' [now
IV Glacial till; Jonesboro, Indiana, on the farm of Dora S. [E.] referred to Archidiskodon imperator]. (4) To these should be
Gift. Type Figure. — Osborn, op. rit., 1922.555, p. 11, fig. added the very large male skull (Nat. Mus. 10261) collected near
10. CoNFt-siON AS TO Paratypes. (1) A pair of upper — Cincinnati, Ohio; in this specimen the ridge formula is 3 *^; M
and lower grinding teeth of both sides (Amer. Mus. Warren Coll. seventeen plates were in use the compression of the grinding teeth
;

10457) fnjiii Zanesville, Ohio, erroneously selected by Osborn is greater, namely:

(1922.555, pp. 11, 12, also figs. 11 and 12) as paratypes; (2) 9 ridge-plates in 100 mm. on the outer [convex] side, at the
subsequently (1923.601, p. 4) referred by Osborn to Elephas worn edge,
washingtonii; (3) finally (1924.633, pp. 4 and 7) made by Osborn 9 ridge-plates in 100 mm. on the worn mid-coronal surface.
the type of a new subspecies, Parelephas jeffersonii progressus The cranial characters of this specimen are entirely similar to
{ = Parelephas progressus);(4) new paratype [ideotype, see foot- those of the three skulls in the American Museum collections,
note on p. 1068] jaws (Amer. Mus. 13225, 21892) selected by except that it is larger and more robust."
Osborn in the present Memoir. {Op. cit., p. 12): "This typical American species is named in
IIlstory of Specific .\nd Generic Separ.\tion by Osborn. — honor of the third president of the United States, Thomas Jefferson,
The following type description is reprinted in part from Osborn's in commemoration of his long-continued devotion to mammalian
paper cited above (Osborn, 1922.555). Osborn introduced the palaeontology."
subject as follows (p. 1): "The present article relates explicitly to Type Locality and post(?)-Wisconsin Age (IV Glacial)
the type characters of Elephas colitmbi, of E. imperator, and of the —
of Parelephas jeffersonii Type. The geologic age of the type
American specimens referred to E. primigenius, three species which of Parelephas jeffersonii is a very important matter; according to
have become more or less confused in all the previous literature Leverett and Hay, the Glacial or Postglacial deposit where it was
because the characters of the type specimens have not been precise- found is of post (?) -Wisconsin age, i.e., IV (iLacial.
ly determined and compared. The object of the present com- (Hay, 1923, p. 139): "6. Fairmount, CJrant County [Indiana].
munication is to clear up this confusion and to propose Elephas Here was found, in 1904, the nearly complete skeleton of the
jeffersonii as a new species of American Pleistocene mammoth. . . . mammoth mounted in the American Museum of Natural History
[Op. cit., p. 15] The American elephant heretofore widely known as in New York City. It has been described and figured [as Elephas
'Elephas columbi,' the Columbian Mammoth, will hereafter be primigenius] by the writer (36th Ann. Rej). Geol. Surv. Indiana, p.
known as Elephas jeffersonii, the Jeffersonian Mammoth." 718, figs. 63, 64 [1912.2]; Iowa Geol. Surv., vol. XXIII, p. 396,
fig. 133) [1914.1]. was found on the farm of Mrs. Dora C. [E.]
It
OSBORN'S SPECIFIC (1922) TYPE DESCRIPTION AND TYPE Gift, about 4 miles east of Fairmount. The location is in the
FIGURES OF PARELEPHAS JEFFERSONII
southeast quarter of section 23, township 23 north, range 8 east.
(Osborn, 1922.555, p. 11): "The above diagnoses of the real This information has been furnished by Mr. George Swisher,
specific characters of the grinding teeth of (the true] Elephas surveyor of Grant County."
columbi. . and of the true E. primigenius leaves without a name
. . "This whole region is mapped by Leverett as being occupied by
the animal which jjreviously has been described in all the literature ground moraine of till plains, and the animal must have lived after
(excepting Soergel's recent Memoir) as Elephas 'columbi.' This the Wisconsin ice cleared away. A tract more or less morainic, an
animal is better known than either of the others; it is represented extension of the Union ('ity moraine, is indicated by Leverett on
in all the collections of the i)rincii)al mu.seums of the United States, his latest map as passing further south than Fairmount. At the
as described by Hay (1914), and particularly in the .American earliest it must have been after the withdrawal of the ice from the
Museum by four especially fine sjjecimens. Of these we select as Union City moraine that the animal lived. Considering the
the type Amer. Mus. 9950, including the skull, jaws, and greater character of the surrounding country, the nature of the deposit
 — " ;

THE MAMMONTIN^: PARELEPHAS 1085

inclosing the skeleton, and the depth at which it was buried, it and described by Warren in 1855 (p. 163, PI. xxviii, fig. C) as
might be supposed that it was not long after the formation of the Elephas 'Primigenius.^
Union City moraine that this elephant existed." It appears (Osborn, 1924.633, pp. 4, 7) that these Zanesville
Genotypic Species of Parelephas (Osborn, 1924.633). 'paratype' molars are typical of a progressive mutation of Parele-
After the establishment (Osborn, 1922.555) of Elephas jeffersonii phas with a higher ridge formula, namely, M
3||. To this sub-
as a species distinct from Elephas columbi and from Elephas species the name Parelephas jeffersonii progressus was assigned.
primigenius, Osborn finallj' (1924.633) reached the conclusion that It becomes Parelephas progressus in the present Memoir.
Elephas jeffersonii could be placed neither in the phylum of
Archidiskodon nor in the phylum of Mammonteus (e.g., Mam-
monteus primigenius) and for the reasons above recited he selected
DENTAL CHARACTERS OF PARELEPHAS JEFFERSONII
it as the genotypic species of the new genus Parelephas. —
Ridge Formula. It has been a very difficult matter to
determine the ridge formula of the species P. jeffersonii; as noted
SPECIFIC CHARACTERS OF PARELEPHAS JEFFERSONII (M
above, the high ridge formula 31^) originally given by Osborn,
Osborn's Error (1922.555) as to Ridge Formula based on as based on the original paratypes from Zanesville, t)hio, was
Zanesville Paratype (since made the Type of Parelephas erroneous; the finally corrected ridge formula of the type, new
—
JEFFERSONII PKOGREssus OsBORN, 1924). In the above original paratypes, and referred specimens of Parelephas jeffersonii is:

description of Elephas (Parelephas) jeffersonii, Osborn erroneously M 3M.

Parelephas jeffersonii Ridge-plate compression at three levels of R.Ms

One-third natural size
Fig. 958. Right third lower molar, r.Ms, of Parelephas jeffersonii new paratype
[ideotype] (Amer. Mus. 13225). Observe: (1) The arcuate arrangement of the
typical twenty-three ridge-plates of this partly worn molar. By counting the ridge-
plates at different levels, by the horizontal lines A, B, C, D, we
as indicated
observe that the upper plates are much more closely compressed than the lower
plates, as follows:

A, Summit of crown, 11'^ plates in 10 cm.

B, One-sixth below summit, 10 plates in 10 cm.
C, Four-sixths below summit, 7}^ plates in 10 cm.
D, Near base of crown, OX plates in 10 cm.

selected as a paratype two upper and lower grinders from Zanesville, The type dentition (Amer.Mus. 9950) is that of a very aged
Ohio, contained in the Warren Collection (Osborn, 1922.555, male by the remarkable length ( = 11 ft.
(Fig. 959), as indicated yiz-s/
p. 11): "As the paratype of this species we select a pair of upper 4J^in. or 3.47 m.), incurvature, and crossing of the tusks (Fig. 966)
and lower grinding teeth of both sides (Amer. Mus. 10457) acquired second by the fact that in the third superior and inferior molars
with the Warren Collection in 1906 and described by Warren in the anterior ridge-plates are worn off. Consequently Osborn was
1855 (p. 163, PI. XXVIII, fig. C) as Elephas 'Primigenius." In 1923, unable to gi\e the type ridge formula correctly in his type descrip-
Osborn corrected this error (1923.601, p. 4), but remained in doubt tion (1922.555) and made the error of depending upon the errone-
as to the true relationship of the Zanesville teeth: "Related to this ously associated Zanesville paratype; in his second description,
species [Elephas washingtonii] may be a pair of upper and lower Osborn (1923.601, p. 4) wrote: "Since the original description of
grinding teeth of both sides from Zanesville, Ohio (Amer. Mus. Elephas jeffersonii was written the molars [Fig. 959] in the aged type
Warren Coll. 10457) acquired with the Warren Collection in 1906 specimen (Amer. Mus. 9950) have been cut out of the jaw of the
 1086 OSBORN: THE PROBOSCIDEA

skull and carefully sectioned; it has thus been found that they are namely: M 3^^ [^]. There can be no question that the type
relati\ely short and deep and present a different ridge formula, of E. jeffersonii has a shorter jaw and shorter and ilee]ier molar
teeth than those of the erroneously associated i)aratype which
may now be referred to E. waxhiiigtonii."
The molar teeth in the aged type speci-
result of sectioning the
men proved that the anterior ridge-jjlates were entirely worn off;
E. jeffersonii Osb. the aged type ridge formula may therefore be written: + 17
3 + 2 0'" M
Amer. Mus. 9950 Type
louter vtew) The adult ridge formula, however, is 3 ^~. M

£ jeffersonii Osb.
Amer. f^us. 9950 Typo

E. jeffersonii Osb
Amer. Mus. 9950 Type
(outer viewl

E. jeffersonii Osb, E, jeffersonii Osb

Amer. Mu< 9'*^^'''vpe
Amer Mus 9950 Type
(inner) view

All U nat. size

Fig. 959. (LEfT) Aged Type Thikd Superior and Inferior Molars of Parelbphas jeffersonii
(Right) The same superposed on Elephas roosevelti [=Syx. Parelephas jeffersonii]
(lioft) Aged type superior and inferior molars of Parelephas jeffersonii (Amer. Mus. 9950), viewed upon the outer surface with the anterior
portion of the crown to the left.

Superior molar, M', partly encased in the maxilla, consequently the fangs are not displayed. Inferior molar, M3, completely removed from the jaw
and sectioned to show the incomplete posterior ridge-plates.

AI, Outer view of left sui)erior molar, l.M^, 1^ anterior ridge-plates worn off; remaining ridge jjlates (5-17) exposed; 17 ridge-plates in use. Total
estimated ridge-plates 25.

Bl, Right inferior molar, r.Mj (rev.), 4 anterior ridge-plates worn off; 18 ridge-plates in use (1-18); incomplete ridge-plates 19-20 demonstrated by
vertical and horizontal sections of this tooth. Total estimated ridge-plates 24.
B2, Inner aspect of right M3, showing 18 ridge-plates in use (1-18); incomplete ridge-plates 19 and 20, at posterior border. Total estimated ridge-
plates 24.

(Right) Diagram showing type superior and inferior molars of Parelephas jeffersonii projected upon the type superior and inferior molars
in heavy lines
of Elephas roosevelti (light lines). This diagram was constructed by superposing type figure 959, left, upon type figure 9(18.

This diagram shows: (1) That the young adult molar teeth of E. roosevelti (light lines) exhibit a ridge formula of M 3 -B^, whereas the aged molar
teeth of P. jeffersonii (heavy lines) exhibit a ridge formula of M
the anterior ridge-plates in this s|)ecies are worn off before the posterior
3 Xirti (2)
ridge-plates attain their full length; (3) in the type of E. roosevelli 10 13 ridge-plate.s are in u.se, in the type of P. jeffersonii 17-18 ridge-plates are in use;
(4) the long diameter of M' is oblique, while the long diameter of M3 is horizontal; (5) following the rule in the Ele|)liantiila', approximately the same number
of ridge-plates is in use at the same time in the superior and inferior molars, namely, in the young E. moseveUi {-3^; in the aged /'. jeffersonii xt'~Vf- C^) '^''^
worn ridge-plate surface of M' is convex; that of Ms is concave.
 : :

THE MAMMONTIN^: PARELEPHAS 1087

It is proven by comparison of the aged type teeth (Fig. 959) Ridge-plate Count at Different Crown Levels (Fig.
of Parelephos jeffersonii with the adult tyi^e teeth (Fig. 968) of its 958. —
(1) As observed in the diagram and figure (Figs. 958, 959)
synonym Elephas ronsevelli, that in the aged type of P. jeffersonii the ridge-plates are arcuate and the ridge-plate arches converge
fwe to six ridge-plates have been worn off in front of the upper toward the summit, the greatest nunaber of ridge-plates may be
molars and four ridge-plates in front of the lower molars, by ex- counted at the summit of the unworn crown, e.g., ll^-i ridge-plates
treme use. This is shown diagrammatically in figure 959, in which in 100 mm., a ridge-plate compression approaching that of Mani-
the aged teeth of the type of P. jeffersonii are superposed upon the monteus primigenius. (2) As we descend down the side of the
adult teeth of the type of E. roosevelti. This diagram also illustrates crown (B), we may find 10 ridge-plates in 100 mm. (3) As we
worn off while the posterior
that the anterior ridge-plates begin to be reach the point of maximum expansion of the ridge-plate arches
ridge-plates have not attained their full depth and are still (C), we may count 7}^ ridge-plates in 100 mm. In general the
growing. compression is greater in the short superior molars than it is in the
Corrected Ridge Formula. —Fortunately it is now possible long inferior molars ; consequently we get the highest count in the
to verify this interpretation of the ridge formulae of these two superior teeth. (4) This difference in the compression of the
types and positively to establish the adult ridge formula of Par- arcuate ridges explains the discrepancy in the records of the
elephas jeffersonii (E. roosevelti syn.) by comparison oijlve different "laminaire frequence" of Deperet and the ridge-plate counts of
specimens as follows: Hay. Similar discrepancies are observed in the ritlge- plate com-

Indiana Type skull and jaws of P. jeffersonii (Amer. Mus. 9950), aged ridge formula: M 3 tM- In Ms, 17 ridge-plates
in use, 3 rudimentary.

Kansas New paratype^ jaw of P. jV^ersonu (Amer. Mus. 21892), a young jaw. Ridge formula: M 3^^. In Ms, 12 ridge-
plates in use.

Kentucky New paratype' jaw of P. jeffersonii (Amer. Mus. 13225), a middle-aged jaw. Ridge formula: M 3 wz+. In Ms,
14-15 ridge-plates in use.

Ohio Referred skull of P. jeffersonii (Nat. Mus. 10261), adult male. Ridge formula: M 3^. In M^, 17 ridge-plates
in use.

Illinois Type grinding teeth of £'Zep/(n.5rooseyeZ<j Hay (Nat. Mus. 2195). Ridge formula: M Sff.

Thus from the recurrent evidence of five different specimens, pression of Archidiskodon and of Mamynonieus. According to the
young, middle-aged, adult, and e.xtremely aged (i.e., type), we observations of Hay (1914), as accepted or modified by Osborn,
conclude as follows: the ridge-plate compression characteristic of different specimens of

Typical adult ridge formula of Parelephas jeffersonii type and

Parelephas jeffersonii may be summarized as follows

four referred specimens: M 3 ff. In the inferior molai-s: 7 ridge-plates (at base) to 11)4 (at
Typical Ridge-plate Compression. —Osborn (1922.555, p. summit of crown) in 100 mm.
12) carefully described the ridge-plate compression in the type In the superior molars: 7 ridge-plates (at ba.se) to 10 (at sum-
inferior molars of Parelephas jeffersonii (Amer. Mus. 9950) as mit of crown) in 100 mm.
follows
REFERRED RIDGE FORMUL.E OBSERVED BY HAY AND OSBORN
7 ridge-plates in 100 mm. on the convex internal surface. The majority of the teeth from the middle and northern
7-7/2 ridge-plates in 100 mm., obliquely worn on mid-coronal United States described as "Elephas columbi" by Hay (1914, pp.
surface; 17 ridge-plates in use. 410-421) actually belong to Parelephas jeffersonii and are readily
distinguishable from the teeth of the true type of Elephas columbi
8 ridge-plates in 100 mm. on the concave external surface.
Falc. of the Southern States as well as from the true teeth of
Thus the number of the ridge-plates in the type of Parelephas Mammonteus primigenius and of Archidiskodon imperator. Elimi-
jeffersonii is: 7, or 8, or 9 in 100 mm., depending upon where the nating six of Hay's specimens from Afton, Oklahoma, which may
count is taken. belong to A. imperator Leidy, also three specimens from the phos-

'[See footnote on page 10G8 above. —Editor.]

1088 OSBORN: THE PROBOSCIDEA

phate beds of South Carolina, which may belong to the true Other Spkcific Characters of the Teeth (Hay, 1914, p.

^7/epAas co/«m?)/ Falc,Hay's detailed observations on teeth prob- 395, Osborn, 1922-1924). — (1) Ridge-plates more widespread
ably or possibly (Alaska?) referable to Parelephas jeffersonii or to than those of E. primigenius. (2) Enamel plates thicker and more
P. M'a.s7ii«(7<o?(// (Washington, Oregon, Alaska) may be summarized channeled and crimped, {'.i) Superior ridge-plates transverse or
as follows: concave posteriorly; inferior ridge-plates transverse or conca\e

Kentucky Phil. Acad. Dp^ (Big Bone Lick, Ky.), length 145 mm., width 75 mm., height of first plate 143 mm.; ridge-
plates 12, front and rear talons. "A line 100 mm. long passes across eight of these plates,
a number greater than one might expect in this species;"

Alaska Nat. Mus. 6669 Dp4 (Alaska, Yukon), length 180 mm., width 67 mm., height 135 mm.; 13 ridge-plates. "There
are seven ridge-plates in a 100 mm. line."

Montana Nat. Mus. 6052 Dp4 (Glendive, Mont.), length 155 mm., width 75 mm.; ridge-plates 8 + .

? Nat. Mus. 287 M- (Loc. unrec), length 300 mm., width 78 mm., height of thirteenth plate 195 mm.; ridge-
plates 18 or 19; mm. line; amount of cement between the plates
eight plates in a 100 small.

Iowa Univ. Iowa 167 M- (Logan, Iowa), length 192 mm., wiilth 70 mm., height of eighth plate 160 mm.; eight plates
in a 100 mm. line.

Alaska Nat. Mus. 6668 Mi (Alaska), length 277 mm., width 90 mm., height of sixth plate 140 mm.; ridge-plates 19;
seven ridge-plates in a 100 mm. line.

Oregon M^ (Mt. Angel, Clack. Co., Ore.), length 355 mm., height of thirteenth plate 193 mm.; ridge-
plates 21 + 5/2 ridge-plates in a line 100 mm. long.
;

Indiana Neb. Mus. Ms (Tipton Co., Ind.), length 280 mm., height of fourth plate 152 mm.; ridge-plates 22 + .

To the above etg^i specimens observed by Hay from Kentucky, anteriorly, i.e., disc shaped or bent. (4) Rear or last superior

Iowa, Indiana, Montana, Oregon, and Alaska, we may add the molars, M', usually strongly arched. (5) Superior ridge-plates,
following six specimens observed and included by Osborn in the M^ not known to exceed 25 to 26; inferior ridge-plates of M3 not
species I'arelephats jeffersonii or P. washingtonii: known to exceed 23 to 24. (6) Bony sheaths of superior tusks

Indiana Amer. Mus. 9950 (type skull) M^ M3 (Jonesboro, Ind.), type aged male, anterior ridge-plates worn ofT. MS+M-
Washington Amer. Mus. 8681 (referred ParelepJias washingtonii skull) M^'' (Whitman Co., southwest Washington), male:
M^ 10+ ridge-plates; M\ 23 ridge-plates. (See Fig. 976.)

Kentucky Amer. Mus. 13225 (paratype jaw) M3 (Twin Cr. near Sanders, Ky., Big Bone Lick region), middle aged, paratype
of P. jeffersonii: M3, 24 ridge-plates.

Kansas Amer. Mus. 21892 (paratype jaw) M3 (near Pendennis, Lane Co., Kan.), young adult, paratype of P. jeffersonii:
Ms, 24 ridge-plates.

Ohio Nat. Mus. 10261 (referred skull) M' (Cincinnati, 0.): M', 26 ridge-plates.

Illinois Nat. Mus. 2195 (type teeth) M^ M3 (Ashland, Cass Co., 111.). Type oi Elephasroosevelli: M', 25 ridge-plates;
M,, 24 ridge-plates. (See l'"ig. 968.)

The type and referred ridge formula of Parelephas jeffersonii shorter and more divergent than those of Elephas primigenius.

is: Dp 2' Dp 3'^ Dp 4+1 M 1

'
'+ M 2'\^ M 3 M. (7) Superior tusks in young males diverging rajiidly, in middle age
According to these fourteen specimens, as described by Hay beginning to rotate, in atlult males converging, so that in aged
and Osborn, the averagespecific ridge-plate frequency of Parelephas males the tips cross.

jeffersonii in 10 cm., as compared with that in Mnmnionteus Detailed Strcctiihe ok the Teeth. — The detailed structure
primigenius, Archidiskodon imperntor, and Parelephas coluinhi, is of the su])erior and inferior teeth of Parelephas jeffersonii is beauti-
as follows: fully shown in the following figures of this Memoir which have been
prepared with the utmost care and precision:
Mammonteus primigenius and M. primigenius compressus:
Fig. 959. Vertical view of the aged superior and inferior
8-10-11-12-13
molars of the tyi)e of Parelephas jeffersonii (Amer. Mus. 9950).
Parelephas jeffersonii 7-8-9 :

Archidiskodon imperalor: 5-6-7 Fig. 960. Superior view of the type and paratype [ideotype]
Parelephas columbi: 5-6 inferior grinding teeth and jaws of Parelephas jeffersonii.
 THE MAMMONTIN.E: PARELEPHAS 1089

Fig. 968. Exterior and crown views of superior and inferior the type jaw (Amer. Mus. 9950) also the internal aspect of the
grinding teetii of the type of Elephas roosevcUi (Nat. Miis. 2195). middle-aged paratype [ideotype] jaw (Amer. Mus. 13225).

Fig. 959. Diagram showing superior and inferior grinding

—
CoMPAUisoNS. These comparative figures, which have been
prepared with the greatest care to a uniform one-eighth and one-
teeth of type of Parelephas jeffersonii and of Elephas rooscvelli.
fourth scale, will enable the student to distinguish these jaws of
Fig. 967. Middle-aged paratype [ideo type] jaw oi Parelephas Parelephas jeffersonii very readily from those of Archidiskodoit
jeffersonii (Amer. Mus. 13225). imperator and somewhat less readily from those of Mammonteus

Jaws of Parelephas jeffersonii from Indiana, Kentucky, and Kansas

One-eighth natural size

Fig. 900. Superior view of type and paratype [ideotype] grinding teeth and jaws of Elephas jeffersonii. Compare (B) with Mather's type of Elephas
[= Parelephas] jacksoHt (Fig. 946).
A, Ideotyijc young adull jaw (Amer. Mus. 21892) with M2 in situ, eight plates in use; M3 with twelve plates in use; total number of ridge-plates in Ms
twenty-four. From near Pendennis, Lane Co., Kansas. Compare Amer. Mus. 14558 (Archidiskodon imperator ref. —Fig. 889).
B, Ideotype middle-aged jaw (Amer. Mus. 13225), Mj shed, M3 with fourteen to fifteen plates in use; total number of plates in M3 twenty-four; jaws
somewhat more massive. From Twin Creek, near Sanders, Kentucky, Big Bone Lick region.
C, Tj'pe of aged Parelephas jeffersonii Osborn (Amer. Mus. 9950); jaw with M3 worn to the edge of the anterior fang; one or possibly two plates have
been worn off, seventeen plates in use, three posterior plates shown in section (Fig. 959, right). Jaws stiil more massive. From Jonesboro, Indiana.

Fig. 892. Comparative figure of jaws of Parelephas jeffer- primigenius. (1) The jaws in all these species of Mammontims are
sonii and other species. correlated with the brachycephali/ and aerocephaly of the cranium,
that is, the jaws are excessively short and deep. (2) The ramus in
Fig. 958. Right third lower molar paratype [ideotype] of
P. jeffersonii (see Fig. 892 D, Dl, C, CI) is far less robust than the
Parelephas jejfersoiiii(Amer. Mus. 13225).
ramus in .4. imperator (Fig. 892 B, Bl, A, Al). (3) The ramus of
P. jeffersonii with rounded inferior border differs from that of
JAWS OF PARELEPHAS JEFFERSONII Elephas indicus (Fig. 893 D, Dl) in which the inferior border is
The first to distinguish the jaw of Elephas [
= Parelephas] more pointed. (4) The deeply depressed ramus of P. jeffersonii is
jacksoni from the jaw of Elephas [
= Mammonteus] primigenius was readily distinguished from the horizontal ramus of Loxodonta
Mather in his description and figure (Fig. 946) of 1838. When all africana (Fig. 893 C, CI) or from the relatively shallow ramus of
the jaws figured in the present chapter, both in superior and lateral the Elephas = Parelephas] Washington ii type (Fig. 893 B, Bl).
[

view, are compared with the jaws figured in the Mammoiiteus —

Rostrum. The most characteristic feature is the prominent
chapter (Chap. XVIII), the latter, it will be observed, are shorter, rostrum perfectly preserved in the paratype [ideotype] of P.
deeper, and more comi)ressed anteroposteriorly, while relati\ely jeffersonii (Fig. 960 B) which projects conspicuously beyond the line
broader, with a broader groove above the rostrum. of the chin. 1 his prominent rostrum apparently distinguishes the
—
Specific Characters of the Jaws. Certainly belonging to Parelephas jeffersonii jaw from the Mammonteus primigenius jaw.
this species are the three specimens shown from above in figure 960, This rostrum is the feature especially shown in Mather's type figure
namely, the aged type (Amer. Mus. 9950), the paratype [ideotype] (Fig. 946) of it is also pointed out in Mather's
Elephas jacksoni;
middle-aged jaw (Amer. JMus. 13225) from the Big Bone Lick type description which leads us to believe that E. jacksoni is more
region, Kentucky, and the paratyijc [ideotype] young adult jaw closely related to Parelephas jeffersonii than to the true Mam-
(Amer. Mus. 21892) from Lane County, Kansas. The lateral and monteus primigenius, although in certain specimens referred to
superior views of the middle-aged paratype [ideotype] are shown in E. primigenius the rostrum is quite prominent, so that we cannot
detail in figure 967. In figure 892 is shown the internal aspect of place too great reliance on this character.
1090 OSBORN: THE PROBOSCIDEA

TYPE AND REFERRED SKULLS OF PARELEPHAS JEFFERSONII shown on the right side of the profile in comparison with a single
We may fortunately compare five finely preserved crania cranium of M. primigenius (C,C'). In this comparison it is ap-
of different ages referable to this species in American collections parent that M. primigenius is even more brachycephalic and
with a number of European crania referred to Elephas inlermedius acrocephalic than P. jeffersonii or P. washingtonii.
(Fig. 935) and to E. irogontherii (Fig. 934). Thus we may establish
the fundamental similarity of cranial structure which distinguishes COMPARISON OF PARELEPHAS AND MAMMONTEUS CRANIA
all the species referable to the genus Parelephas. We observe (Fig. 961) that in the frontal aspect these three
Cranial Characters (Figs. 934, 935, 961-963).— As i)oint- crania of Parelephas are remarkably similar and readily distinguish-
ed out in the introduction to the Mammontinse, superficially the able from crania of Manimonteus (Fig. 962 C). In frontal aspect:
profile aspect of the cranium of Parelephas jeffersonii (Fig. 962A) (1) The occipital crest, which is perfectly preserved in the aged
resembles that of Elephas tndicus, but the midcranial section shows type skull from Indiana (A) is broad and swelling at the summit;

that it is fundamentally distinct. The

form of P. jeffersoiiii
cranial it is on the right side in the adult skull of the
also well preserved

is actually intermediate between the cranial form of Archidiskodon specimen from the state of Washington (B); it is erroneously
iiHperator and that of Ma m mordeus primigcn ius. Close comparison restored in the specimen from Ohio (C). (2) The anterior narial
with M. priwigenius is afforded in figure 962, in which two crania, openings are ^•el•y broad and widely open, whereas in Manimonteus
one of P. jeffersonii (A) and the other of P. washingtonii (B) are they are smaller and more contracted. (3) The orbital sockets are

Frontal Views of Three PAnELEi-HAs Crania; (A, C) P. jeffersonii, (B) P. washingtonii

All figures one-twelfth natural size

Fig. 961. Front view of the (A) type skull of Parelephas jeffersonii Osborn (Amer. Mus. 9950), from Jonesboro, Indiana, coini)are(l with one skull (B) in the
.\meriean Museum and one (C) in the U. S. National Museum, as referred to in the original type description.
A, Front view of the aged tyix' .skull of Parelephas jeffensntiii (Amer. Mus. 9950); actual mea.'<>ircment 28.8 inches across the outside of the orbits.

B, Hef.Trcd skull of Parelephas wnshiiiglnnii (Amer. Mus. Cojjc Coll. 8081) from Whitman County, Washington, heretofore dcsiKiiatcd as "EIrphns
aAumbi," and subse(|ucntly referred by Osborn (1922. .5.55, p. 15) to Parelephas jeffersonii. This skull was never figured or described by Cope, it was found in tin
unopened box in his collection. For full description of this specimen, see pages 1101, 1 103, and figures 937, 973, 974, 971, and 976 of the present Memoir.
C, A very large male skull in the National Museum (Nat. Mus. 10261), collected near Cincinnati, Ohio, referred to P. Jc^frsorwi in 0.sborn's type descri))-
tion (1922.555, p. 15): "The cranial characters of this specimen are entirely similar to those of the three skulls in tlie .\merican Mu.scum collections, except
that it is larger and more robust." Occiput incorrectly restored.
 THE MAMMONTIN^: PARELEPHAS 1091

much more prominent than in .1/. priniigrnius, as perfectly shown to add the description and figures of the superb skull and tusks
in all three skulls (A,B,C). (4) The maxillo-premaxillary sockets of from Nebraska preserved in the Museum of the University of
the superior tusks are relatively less elongate and less parallel than Nebraska (Neb. Mus. 1-4-15), photographs of which (Figs. 963
in Mammonteus; they diverge widely where the tusks issue from and 964) we are able to reproduce through the courtesy of Prof.
the skull. In profile view (Fig. 962 B) the orbits are more
(5) E. H. Barbour. Like the two crania from Indiana and Ohio
widely separated from the occipital condyles than in M. primigenius previously described, this Nebraska skull was originally referred
(Fig. 962 C). In profile view the occiput is more prominent
(6) by Professor Barbour to "Elephas columbi," but it agrees in all
and bulging 962 B) than the relatively vertical occiput of
(Fig. particulars with the two crania above described as Parclephas
M. primigenius (Fig. 962 C). (7) In its proportions the M. jeffersonii and is the largest and in many respects one of the finest
primigenius cranium is broader, shorter, higher, deeper, i.e., more representatives of this type of cranium, the tusks exceeding in
brachycephalic and hypsicephalic, than the P. jejjersonii cranium. length and in circumference those of the type skull of P. jeffersonii
(8) In all these characteristics the cranium of P. jeffersonii appears from Indiana.
to agree more closely with the crania of the European species The left profile view of this Nebraska cranium (Neb. Mus.
Parclephas intermedius (Fig. 944), and P. trogoniherii as shown in 1-4-15) is shown in figure963 the profile is closely similar to that
;

profile in figure 934, than it does with either profile or front views of the type of Parelephas jeffersonii, except that the concave plane
of the cranium of Mammonteus primigenius. of the forehead is relatively more elongate; we observe that the

SKULL OF PARELEPHAS JEFFERSONII (KNOWN AS THE orbital region is very prominent, the summit of the occiput is
FRANKLIN COUNTY MAMMOTH) IN THE rounded, the postoccipital profile is very convex, the occipital
NEBRASKA STATE MUSEUM condyle is elevated ; the entire profile is similar to that of the male
Discovered in the loess of Wisconsin or late Pleistocene time. skull of Parelephas figured by Falconer (Fig. 934). The palatal
—
Nebraska Skull. Since the crania above described come view (Fig. 963) displays the third superior grinding teeth in. situ
from the states of Indiana, Ohio, and Washington, it is interesting and exhibits (Fig. 965) nineteen ridge-plates, of which the anterior

Mammonteus primigenius (C, C), Parelephas jeffersonii (A), P. washingtonii (B)

All figures one-twelfth natural size

I'ig. 962. Profile views of type and referred skulls of Parc/ep/ias jej'crsojiu (A), P. washinglonii (B), and Maminontcus primiqcidus (C) in the American
Museum and United States National Museum, also front view of M. primigenius (C).
A, Type of Parclephas jeffersonii (Amer. Mus. 9950), reversed. An aged individual. Main portion of the tusks not included. From .loncsboro, Indiana.
B, Referred skull of Parelephas washinglonii (Amer. Mus. Cope Coll. 8681). From Wliitman County, Wasliington.
Mammonteus primigenius
C, Referred skull of (Nat. Mus. 8580) from Siberia, with jaws belonging to another individual (Nat. Mus. 8579), from Alaska.
C, Front view of same skull and jaws.
OS
o
 THE MAMMONTIN^: PARELEPHAS 1093

thirteen are in use. Behind these thirteen more or less worn ridge- thorough soaking, and reinforcing with an internal steel rod in
plates (Fig. 963) are observed seven to eight unworn ridge-plates plaster to prevent them from crumbling away. Vigorous efforts
making an estimated total of twenty to twenty-three ridge-plates. were made to find the missing lower bones of the limb and the feet
The widely divergent maxillo-premaxillary region presents the but without success.
usual contrast with the parallel and convergent bony sockets of the Mounting of Parelephas jeffersonii Type. The missing —
Mammonteus primigenius type of cranium. Comparison is afforded parts are the radius and ulna of both sides, the right tibia and
in figures 899 and 900 with the palatal view of two crushed skulls of fibula, the fore- and hindfeet. These parts are restored from the
Archidiskodon imperator from Hay Springs, Sheridan County, more massive limbs of two specimens of Archidiskodon and it is
Nebraska (see Chap. XVI). doubtful whether they are of the proper proportions for Parelephas.
Locality and Geologic Level. This — skull was discovered Thanks to the kind cooperation of Dr. Marcellin Boule of the
from 11 to 12 feet underground while digging for the foundation of Museum of Palaeontology, Paris, casts were secured of the lower
the new high school of Camjibell, Franklin County, Nebraska; portions of the limbs and of the feet of the great skeleton of Elephas
the date of discovery was April 1, 1915. The geologic deposit was meridionaUs (Fig. 866) in the Paris Museum; on arrival in the
a bed of loess which we may infer belongs to late Pleistocene times. American Museum these casts were remodeled, recourse being
had
As restored under Professor Barbour's direction and mounted in for comparison to remains of the feet and limbs of Archidiskodon

A/SB. MUS /-.*-/J

The Franklin County Mammoth of Nebraska

\M^^
Fig. 964. Nebraska Skull. — Complete skull, jaws, and tusks of
Parelephas jeffersonii (Neb. Mus. 1-4-15) discovered in the loess near Camp-
bell,Franklin County, Nebraska. After photograph by E. H. Barbour. Skull
Molar Ridge-plates of Franklin County Mammoth of Nebraska.
essentially without restoration.
Part of the coronoid region of the jaw restored Aged Parelephas jeffersonii
in plaster; symphysis of the jaw perfect; both mandibular rami broken.
Fig. 965. Franklin County Mammoth. Diagram drawn from casts
Tips of the tusks weathered off and restored with characteristic curvature.
showing the (13) superior and (16) inferior worn ridge-plates of Parelephas
One-tenth natural size.
jeffersonii ref. (Morrill Coll., Neb. Mus. 1-4-15). Outlines furnished by
Professor E. H. Barbour, August 29, 1924.
Crown view of M-, M'. (Lower) Portion of the crown of M3
(LTpper)
the University of Nebraska Museum, the skull and tusks are shown
of the same specimen. See figure 963 for cranium of tliis specimen before
in side view in figure 964. The tusks are 12K feet in length and 29 restoration, figure 964 after restoration.
inches in circumference at the thickest point; the cranium and
jaws are essentially without restoration; the enormous dejjth of
imperator;all this work of restoring and remodeling was done by
the skull from the occipital condyle to the inferior border of the
Otto Falkenbach under the direction of head preparator Adam
jaw is shown by the seated figure of a student whose forehead rests
Hermann. Mr. Hermann himself restored the missing surfaces of
just opposite the occipital condyle.
the upper parts of the skeleton which were, however, in exception-
ally complete condition.
TYPE SKELETON OF PARELEPHAS JEFFERSONII —
Pose. The animal is represented with uplifted head and as
See figures 931 and 966 of mounted skeleton if just starting to walk. For the pose of the skeleton the coopera-
The type skeleton was found near Jones-
of Elephas jeffersonii tion of Director Hornaday of the Zoological Park was secured;
boro, Indiana, in 1903. was purchased for the Museum with the
It the tracks of the Indian elephant "Gunda" slowly walking over
Jesup Fund in .\ugust, 1904, from Dora E. Gift and others. As a sandy surface fifty feet long were carefully plotted by park
found the skeleton was imbedded in a muck deposit of late Pleisto- engineer Beerbower (Fig. 1243 of this Memoir) through this means ;

cene age, fifteen feet below the surface; this deposit is considered it is believed that the fore- and hindfeet are correctly placed. The
of IV Glacial, post(?)-Wisconsin by Leverett and Hay. The
age, author also made a special study of the position of the bones of all
whole upper portion of the skeleton was complete and intact, parts in the living elephant. The result of all this work is a pose
including the remarkably incurved tusks, which were preserved for and gait which are believed to be true to life.

their entire length although requiring surface restoration and The most striking features of this skeleton of an aged in-
 Type Skeleton of Pahelephas jepfersonii in the American Museum
One twenty-fourth natural size
Fig. 966. First published tyjie figure of the aged skeleton of Elephas jeffersonii Osborn, 1922 (Amor.Mus. 9950), as mounted in the Ameriean Museum.
I'liis siH-cimrn, including the skull, jaws, and greater part was found near Jonesboro, Indiana, on the farm of Dora E. Gift; i)inihased for the
of the skeleton,
American Museum with the Jesup Fund in 1904; parts of limbs restored and mounted in 1906; described and figured by Osborn in 1907 (1907.29.J) as
Elephas columbi; redescribed by Hay (1914) as "Elephas primigenius," and again in February, 1923, p. 139, as K. primigenius of post(?) -Wisconsin
age.
This skeleton was finally selected and figured by Osborn in 1922 (1922.555, p. 11, fig. 10) as the type of Elephas jeffersonii new species.
(Cf. Osborn, 1907.295, p. 256): As found the skeleton was imbedded in a muck deposit of late Pleistocene age, fifteen feet below the surface. The whole
upper portion of the skeleton was comijlete and intact, including the remarkably incurved tusks, which were preserved for their entire length although requiring
surface restoration and thorough soaking, and reinforcing with an internal steel rod in plaster to prevent them from crumbling away. Vigorous efforts were
made to find the missing [restored] lower bones of the limb and the feet but without success.
The missing parts [restored] are the radius and ulna of both sides, the right tibia and fibula, the [both] fore-and hindfeet. Thanks to the kind cooperation
of Dr. Marcellin Boule of the Museum of Pala-ontology, Paris, casts were secured of the lower portions of the limbs and of the feet of
the Elephas meridinnalis
(I'ig. 8ti6) in the Paris Mu.scum; on arrival in the .\merican Museum these were remodeled, recourse being
had for comparison to remaitis of the feet and
limbs of Elephax imperator; all this work of restoring and remodeling was done by Otto Falkenbach imder the direction of head preparator Adam Hermann. The
animal is represented with ujjlifted head and as if just starting to walk. For the po.se of the skeleton the cooi>eration of Director Hornaday of the Zoological
Park was secured; the tracks of the Indian elephant "Gunda" (Chap. XXHI, p.I598) .slowly walking over a.sandy surface fifty feet long (Fig. 1243 of the present
Memoir) were carefully plotted by park engineer Beerbower; througli this means it is believed that the fore- and hindfeet arc correctly i)laccd. The author
also made a special study of the position of the bones of all parts in the living elephant.

1094
 THE MAMMONTIN^: PARELEPHAS 1095

dividual of Parelephas jeffersonii are the following: (1) The com-

plete incurvature and crossing of the tusks, indicating that this
animal is an old bull which had retired from the herd. (2) The
relatively small size of the skull. (3) The abbreviation of the back
and body in contrast to the vertical elongation of the limbs.
Dietrich (1916, p. 76, 1924, p. 1.3 — p. 1276 below), in describ-
ing Elcphas anliquus recki, discusses the size of the humerus of
recent and fossil proboscideans and gives a number of measure-
ments which we may compare with those of the type of Parelephas
jeffersonii as follows:

Length of Estimated Shoulder

Humerus Height

Parelephas trogontherii
of Mosbach 1480 mm.' 4500 mm. [Schmidtgen,
1926, p. 64]
[Parelephas jeffersonii
1096
OSBORN: THE PROBOSCIDEA

"Type specimen. Upper and lower hindermost molars, No.

2195, U. S. National Museum.Type locality. Ashland, Cass
Type formation. Pleistocene. Diagnosis.
County, Illinois.

Hindermost molars long and low, the base and the summit
approxi-

mately parallel, consisting apparently of 25 plates; of these 8 in

Parelephas jeffersonii (synonym Elephas roosevelti)

Third Superior and Inferior Molars
Fig 968 Figured for the present Memoir. Type superior and inf.'rior
Mus. from Ashland, Cass
molars of Eleptias roosevdli Hay (Nat. 2195),
Paratype ok PaHELEI'HAS jbffeusonii
County, Illinois. One-fourth natural size.
I'ig. 9G7. Jaw of midillc-agcd paratyix; [ideotypol of Parelcplias jeffcrmnii
Big Bone the right superior molar, r.M^
(.\iiicr. Mu;*. 1322.-)) from Twin Creek, near .Sanders, Kentueky, A, Al, Crown and external views of
allowing twenty-five ridge-plates (1-25 in the figure).
l.ick region.
M2 been rceently slied; M3 exhibits twenty-three
Tlie rudiment of lias r.Ma, showing
B, Bl, Crown and side views of the right inferior molar,
ridge-plates (sec Fig. 9G0B, showing 24 ridgc-platcs) of which the anterior being broken away.
ridge-plate's 3-24, the anterior ridge-plates
fourteen are in use.
 THE MAMMONTIN^: PARELEPHAS 1097

a 100 mm. line; enamel thin, delicate, and little folded. [Type The diameters, hdge-]ilate measurements, and indices of thc^fe
figure. None. See figure 968 of the present Memoir]." young molars clo.sely resemble those of the aged molars of the type
Type Description of Elephas koosevelti. The type — of Elephas jeffersonii (Fig. 959, left) as clearly shown in the diagram
description was introduced by the following notes: (Hay, Sept. (Fig. 959, right), in which the superior and inferior molars of the ag-
30, 1922, p. 100): "Professor Osborn in his paper of July 8 ed E. jeffersonii type are projected to the same scale upon the type
[Osborn 1922.555] has published an interesting figure of upper teeth superior and inferior molars of the young adult E. roosevelti. Con-
of an elephant (his fig. 8) ["Fig. 8. Fourth and third superior sequently we have been able to amplify the specific characters of
molars of .Elephas primigenius"] found in Indiana. On plate
. . E. jeffersonii by a study of these grinding teeth of E. roosevelti,
Lix of the twenty-third volume of the Iowa Geological Survey, I which compare so closely in all respects except in age and wear.
published a figure of very similar teeth found at Milwaukee, Wis- The ridge-plate compression is as follows:
consin, and preserved in the Public Museum of that city. The
hindermost molar had just begun to suffer wear. In the U. S. Left Right
National Museum are right and left hindermost molars (No. 2195) M^ ridge-plates in 100 mm. on convex
of similar form, found at Ashland, Cass County, Illinois; also an external surface 9K. 9
upjier left hindermost molar (No. 4761) hardly different, discovered ridge-plates in 100 mm. on mid-
in Wayne Township, Darke County, Ohio. The peculiarity of all coronal surface 10 9)i

these teeth is the low elongated form and the approximate parallel- ridge-plates in 100 mm. on mid-
ism of the upper and the lower borders. Inasmuch as the molar coronal conca\e internal surface. 8 9
descends at a nearly right angle with the grinding face of the tooth Ms ridge-jjlates in 100 mm. on mid-
in front it seems probable that the skull was short. Professor coronal concave external surface, 7 7-8K
Osborn has referred his specimen to Elephas primigenius; but I ridge-plates on worn mid-coronal
find no teeth from Alaska or the Old World which present similar surface, that is, in 100 mm incomplete
characters. I believe that a hitherto unrecognized species is in- ridge-plates in 100 mm. on con\'ex
dicated. This I propo.se to call Elephas roosevelti in honor of internal surface 6-7K
another statesman and naturalist, one whose multifarious interest
led him to pursue living elephants in their African wilds."
"The Ashland teeth are chosen because with them came the Parelephas progressus Osborn, 1924

nearly complete lower right hindermost molar. The length of the Figures 969, 970

molars is 300 mm., the height 170 mm., the width of the
close to Upper Plei-stocene, post(?)-Wiscon.sin (IV Glacial) age, Zanesville,
upper teeth 90 mm., of the lower 85 mm." Muskingum County, Ohio.

Osborn, 1924 Hay's type description, in which he compares

:
based on a jiair of superior and
This very progressive species is

this animal with E. primigenius, was not accompanied by a figure inferior grinding teeth describedby Warren in 1855 as Elephas
and for the purposes of this Memoir the type teeth of Elephas ^Primigenius'; erroneously selected by Osborn (1922.555, pp. 11,
roosevelti (Nat. Mus. 2195) were kindly loaned to the American 12) as a paratype of Elephas jeffersonii. The type teeth belong
Museum by Curator Merrill for examination and execution of the to a much larger animal than the type and referred specimens of
type figure which is reproduced herewith (Fig. 968). Elephas jeffersonii and present a more progressive ridge-plate
This type figure, including its very careful enumeration of the formula, namely : M
3 M. Its descent from the P. jeffersonii stage
ridge-plates as preserved, namely, 24-25 ridge-plates preserved is indicated by the similarity in the ridge-plate compression, indi-
above, 21 ridge-plates preserved below, agrees with the type de- cating that the cranium of this species will jirove to have the same
scription of Hay. The mid-coronal ridge-plate count is 8 in a 100 distinctive proportions as the cranium of P. jeffersonii.

mm. line. The diameters of the molars are as follows: Parelephas jeffersonii progressus Osborn, 1924. "Parelephas
in Relation to Phyla and Genera of the Family Elephantidae."
Left Right Amer. Mus. Novitates, No. 152, December 20, 1924, i)p. 1, 4,

Third superior molars, length across and 7 (Osborn, 1924.633). Type.— Amer. Mus. Warren Coll.

25 ridge-plates 227 mm. 282 mm. 10457. A pair of superior and inferior grinding teeth of both
Third superior molars, height of sides. Horizon and Locality. — Zanesville, Muskingum
tallest ridge-plate 202 192 County, Ohio; LTpper Pleistocene, Wisconsin or post(?)-Wisconsin
Third superior molars, max. width 95 98 (IV Glacial) age. Type Figure.— Osborn, 1922.555, p. 13,
Third superior molars, breadth- fig. 11, and p. 14, fig. 12.

length index tooth incomplete Type Description.— (Osborn, 1924.633, pp. 4, 7): "These
Third inferior molars, length, esti- progressive grinders, described by Warren in 1855 as Elephas
mated, of 22 ridge-plates 304e primigenius, by Osborn in 1922 as a paratype of Elephas jeffersonii,
Third inferior molars, height of tall- belong to a much more progressive stage than the type of Elephas
est ridge-plate 154 jeffersonii, and referred specimens, presenting a progressive ridge-
Third inferior molars, max. width 92 plate formula of M 3 M, as compared with the typical ridge-plate
Third inferior molars, breadth-length formula of Parelephas jeffersonii, namely M 3 ¥x. interest- It is

index tooth incomplete ing to observe that these type molars (Osboin, 1922.555, p. 14,
1098 OSBORN: THE PROBOSCIDEA

fig. show eighteen ridge-plates

12) in use in both ]\P and M3; the ing teeth of both sides (Amer. Mus. 10457) acquired by the
superior molars show from four to six ridge-plates in excess of the American IMuseuni with the Warren collection in 1906. It was
inferior molars; thus they are readily distinguishable from those of described by Warren in 1855 (p. 163, PI. xxviii, fig. V) as Elephas
Mammonleus primigenius compressus, as described above. In all 'Primigenius' and described and figured by Osborn in 1922
other characters these type grinding teeth (Osborn, 1922.555, fig. (1922.555, p. 13, fig. 11) as Elephas jeffersonii. This type affords
12) are related to those of Parelephas jeffersonii Consequently, we . a complete comparison with the species Parelephas jeffersonii (as
assign the subspecific name progressus to denote this extreme stage defined above with a ridge-plate formula of M 3 f^), the ridge-plate
in the long Parelephas series." formula in P. progressus being M
S .^t^tie the right Ms presenting y

Type, Zanesvillj:, Ohio. —A pair of upper and lower grind- the last lower ridge-plates more fully developed than in the left
lower M.i.
Characters of Type. — It is interesting to observe that these

Amer. Mut. 10457

Amer. Mus 10457

(outer view)

Amer. Mus. 10457

'outer wiewi

Amer. Mus. 10457

Amer. Mus. 10457

(inner view)

Type Molar.s of Parelephas phooressus (Side and Crown View.s) Originally fiqured by Osbohn a.s Paratvpbs of Elephas jeffersonii
One-fourth natural .size

Fig. 969. Side view.s of type molars of Parelephas jeffersonii Fig. 970. Crown views of type molars of Parelephas jeffersonii
progressus (Amer. Mus. Warren Coll. 104.57), Zanesvillc, Ohio, one- progressus (Amer. Mus. Warren Coll. 104.')7), the .same individual
fourth natural size. After Osborn, 1922..Jo.">, j). 13, fig. 1 1. as in figure 969, one-foiirtli natural size. Aftt^r Osborn, 1922.5.5.5,

Al, External aspect of left superior molar, l.M', .showing 30 p. 14, fig. 12,

ridge-plates, of which 18 are in use.

A, Crown view of left superior molar, l.M'', showing 30 ridgc-
Bl, External aspect of left inferior molar, l.M.i, showing 24 ridge-
plates, of which 18 are in use.
plates, of which 17 are in use.

B2, Internal aspect of right inferior molar, r.Ms, showing 2fi B, Crown view of right and left third inferior molars containing
ridge-plates, of which 18 are in use. respectively 26 and 24/2 ridge-plates, of which 17 are in use.
 : :

THE MAMMONTIN^: PARELEPHAS 1099

type molars show 18 ridge-plates in use in both M^ and M3, but 8ci., ser. 2, vol. XV, pji. 146-147) is found a brief account of
that the superior molars show from/o(/r to .six ridge-plates in excess the disco\ery of elephant remains at Zanesville. One tusk and four
of the inferior molars, the ridge formula being \l ^^-t'l-Ta- The mol
molars were found. Two of the latter weighe<l (probably while
ridge-plate compression, however, corresjjonils with that of Par- wet) 20 pounds each and two others 14 pounds each. They had
elephas jeffersonii and is between that of
directly intermediate been found on the line of what was then called the Ohio Central
Parelephas coluinbi-Airhidiskodoii iniperator and that of Mam- Railroad and in the eastern part of the city. At about the same
monleus primigeniiis, the internal, external, and coronal ridge- time (Proc. Boston Soc. Nat. Hist., vol. IV, p. 377) Warren ex-
plate measurements being as follows hibited a tooth of an elephant, one of three received by him from
Zanesville (misprinted Lanesville). In the .second edition of his
M^ 7-8 ridge-plates in 100 mm. on the external convex surface
monograph on "Mastodon giganteus" Warren figured one of these
8-8)l> ridge-plates in 100 mm. on the oblique mid-coronal
teeth (his plate xxviii). It was stated that he had four of the
grinding surface
teeth, all belonging to Elephas primigenius. These are now in the
9-10 ridge-plates in 100 mm. on the internal concave
American Museum of Natural History, New York [Amer. Mus.
surface
10457). ^rhe right u])per hindermost molar is a fine large tooth.
Ms, 6)^-7 ridge-plates in 100 mm. on the external concave
The large front root is missing, as are quite certainly about 3
surface
plates. There are now 28 present. The length along the nearly
7-7}) ridge-plates in 100 mm. on the oblic|ue mid-coronal
straight base is 335 mm. The rear is high and arched. There are
grinding surface
9 plates in a 100-mm. line and the enamel is little festooned.
6-6K ridge-plates in 100 mm. on the internal convex
Foster, in 1857 (Proc. Amer. Assoc. Atlv. Sci., 10th meeting, p.
surface
156), described the discovery and exhumation of these remains,
This compression with averages of 6-7-8-8'i-9-10 ridge-plates publishing a geological section illustrated by a figure. The elephant
in 100 nun., according to the point of measurement of the ridge- bed is 37 feet above the river and over 20 feet from the surface."
plates, concurs with the measurements given by Osborn in the six —
Measurements of Indices. It is interesting to make a close
type and referred specimens of Parelephas jcffermnii. comparison of the measurements and indices of this historic pair
Comparing the ridge-plate formula of J^arelephax progressus of grinding teeth with the type grinding teeth of Elephas jeffersonii
(M 3 „/ii72i) with that of the most progressive form of Mam- and of Elephas rooseveUi.
montens prrmigenius from Alaska ami Indiana, namely, M. primi- From the following ccjmparative measurements of the grinding
geniiis rompressus (M 3;-.,^,-|^), we obserx'e that /-". progressus has teeth it appears that Parelephas progre.ssus was undoubtedly
a higher ridge-plate formula, but M. primigenius compressus (Al- an animal of larger size than the type of P. jeffersonii; in the

aska) has a much clo.ser ridge compression. grinding teeth the ridge-plates, while more numerous, are much
As shown in the type figures of Parelephas progressus, here- more compressed, so that the total length of AP mm.) is longer
(205
with reproduced (Fig. 969), the ridge-plates in external and internal than the total length of M' in P. jeffersonii (203-|- mm.) the grind-;

have the sinuous ami arcuate forms, such as are observed in

as])ects ing teeth are relatively broader (109 mm.) as comi)ared with 108
Parelephas jeffersonii, more pronounced than the simple, arcuate mm. ; the average thickness of the superior enamelled ridge-plates
ridge-plates of Mammonteus primigenius of Alaska (Fig. 1008). in the two species is the same, namely, 12.5 mm. In the lower
(Ikologic Location after Hay. — (Hay, 1923, p. 134) grinding teeth the ridge-plates of P. progre.^sus occupy 221 mm.
"2. Zanesville, Muskingum County, [Ohio). — . . . Zanesville is as compared with 208 mm. in P. jeffer.sonii (cf. table below).
situated in the unglaciated part of the State [of Ohio)but outwash ; Progressive Specific Characters. From these compari- —
from both the Illinoian and the Wisconsin glaciers has been de- sons we observe that this Zanesville mammoth may be regarded as
])osited along the river. For a knowledge of the Pleistocene epoch a distinct species, namely, Parelephas progre.ssus, i)0.s.sibly the last
in that region, Leverett's work may be consulted (Monogr. U. S. member of its race, lingering probably into pf)st-Wisconsin or early
(".eol. Surv., vol. XI^I, p. 158, plate 11). ... In 1853 (Amer. .Jour. Postglacial times.

Parelephas

M' Total mid-length of crown including all the ridge-plates.

Maximum height of crown, height of tallest ridge-plate.
Maximum width of crown at the broadest portion
Breadth-length index
Height-length index
Average thickness of superior ridge-plates

Ms Total mid-length of crown including all the ridge-plates.

Maximum height of crown, height of tallest ridge-plate.
Maximum width of crown at the broadest portion
Breadth-length index
Height-length index
Average thickness of inferior ridge-plates
 Amer Mus. 8681 Ref,

Referred Cranium of Parelephas washingtonii from the State of Washington

One-eighth natural size

Fig. 971. Referred skull of Parelephas washingtonii (Amer. Mus. Cope Coll. 8681). Pleistocene beds near Pine Creek, Whitman County, state of
Washington. From the same locality is recorded the type jaw of Elephas [= Parelephas] ivashirigtonii Osborn (Amer. Mus. 8681 A). This superb skull was
dis<'overed by Charles H. Sternberg in 1876 and purchased by Prof. E. D. Cope, but the box containing it was never opened. Observe that M^ exiiibits 10 +
worn ridge-plates, the anterior plates having disappeared; M^ exhibits 6 worn ridge-plates, the posterior i)Iates not having come into use.
A front view of the .same skull is shown in figure 961 B; a perfect right lateral aspect is shown in figure 962 B; the l.M'' is also shown in section in
figure 976.This beautiful drawing and figure 973 were executed by Mrs. L. M. Sterling in 1902. The determination and lettering of the bones and foramina
were under the direction of William K. Gregory.

1100
 THE MAMMONTIN^: PARELEPHAS 1101

Parelephas washingtonii Osborn, 1923 region as well as a long and shallow ramus, quite different in
Figures 818, 893, 935, 937, 961, 962, 971-977, 1009 proportions from that of the true E. jeffersonii type. This species
Pleistocene, pre(?)-Wisconsin age, Pine Creek, Whitman County, is named honor of President Washington, in contrast to the
in
Washington. species previously named in honor of President Jefferson."
The type of the species Parelephas washingtonii (Amer. Mus. This type jaw is specifically distinct from that of any known
8681A) is one of the most primitive jaws thus far found in America; member of the Parelephas phylum. Comparison with the Loxo-
it comes from Pine Creek, Whitman County, state of Washington. donla jaw (893, C, CI) demonstrates that this species (P. washing-
The jaw has a very prominent rostrum, a long and shallow ramus, tonii) had a relatively long jaw but not quite so elongate as that
and a depressed sinus between the coronoid and the condyle, differ- of Loxodonta.
ing in every respect from the mandibular ramus referable to any The measurements of the lower left grinding tooth, Ms, are
of the known species of Parelephas and resembling only in the form as follows:
of the mandibular ramus the primitive species Elephas [Archidis-
kodoii] hayi of Barbour. Consequently the reference to the genus M3 maximum length of crown, including 21 ridge-
Parelephas (?) is largely provisional and we are uncertain as to its plates 348 mm.
phylogenetic position; the relatively high ridge-plate formula maximum width of crown across the broadest
(M 3 21 ) clearly separates the lower grinders of this species from ridge-plate 88
the type of Archidiskodon hayi (M 37-5777), as shown in the com- ridge-plates in 100 mm 5
parative figures 915, 893. It is noteworthy that the ridge formula breadth-length index 25
of P. washingtonii (M 3 |-^) agrees with that of P. intermedius Specific Characters. — Ramus long and shallow with very
(M 3 ll'.ll ) of southern France. prominent rostrum, depressed coronoid and depressed sinus be-
Elephas washingtonii Osborn, 1923. "New Subfamily, tween coronoid and angle. Ridge formula: 3 ut. M
Breadth-
Generic, and Specific Stages in the Evolution of the Proboscidea." length index lower than in other species of Parelephas. Probably
Amer. Mus. Novitates, No. 99, Dec. 27, 1923, p. 4. Type.— a jjrimitive member of the Parelephas phylum.
Lower jaw containing two third inferior molars, with the ascending —
State of Washington Skull. The superb referred skull
rami missing. Horizon and Locality. Pine Creek, Whit- — (Fig. 971) oi Parelephas washingtonii (Amer. Mus. Cope Coll. 8681)
man County, state of Washington. Pleistocene, pre(?)-Wisconsin is stated by Mr. Charles Sternberg to have been collected by him-

age. Type Figure.— Figures 972, 975, 893, B, Bl, of the self in Pleistocene beds near Pine Creek, Whitman County, state of
present Memoir. Washington. It had never been described or figured by Professor
Type Description.— (Osborn, 1923.601, p. 4): "Elephas Cope, but was found in an unopened box in the Cope Collection
washingtonii, new species. As the type of the species Elephas when acquired by the American Museum. In previous literature
washingtonii we select a lower jaw (Amer. Mus. 8681A) containing by the present author, this skull has been referred to as "Elephas
two third inferior molars, with the ascending rami missing, from columbi." The brachycephalic proportions of the Washington skull
Pine Creek, Whitman County, Washington. The ridge formula in which relate it to Parelephas are shown both in the anterior aspect
these two teeth is: M 3 21. The jaw has a depressed coronoid (Fig. 971A) and in the palatal aspect (Al). All three aspects

E. washingtonii
Amer. Mus 8681A Type

Fig. 972. New figure of the type adult jaw of Parelephas washingtonii Osborn (Amer. Mus. 8681A), one-eighth natural size, exhibiting twenty-one
ridge-plates in M3. Median (left), coronal (center), and internal (right) aspects. Compare with figures 974, 97.5 B, 893, B, Bl.
As fully characterized in the type description and by the comparative figures, this jaw is readily distinguished by the greatly depressed coronoid
region which gives it a wholly different aspect from the type jaw of Parelephas jeffersonii.
1102 OSBORN: THE PROBOSCIDEA

(Figs. 961 B. 962 B, 971 A) also palcatal asi»ct (971 Alj slioukl be 'mudsprings,' in the swamps of Pine Creek valley, Whitman
compared with the corresponding aspects of the cranium of Elephas County, Washington, about 100 miles north of Walla Walla. Skull
I'ndicu.i (Fig. 963) to convince the student that the crania of Pnr- dislodged and brought to the surface by means of a long iron rod
and P. washingtonii are far more brachycephalic
eleyhas jeffersonii with grajipjing hooks, on Mr. Copeland's farm.' First e.xhibited
and bathycephalic than the cranium of E. indicus. by a showman, then through Charles H. Sternberg jjurchased by
Geology .\nd Fauna (compare Sternberg, 1903, pp. 511, E. D. Cojje. I^he bed e.\]ilored by Sternberg early in 1878 yielded
512). —We owe to Charles H. Sternberg an interesting description recent buffalo crania, bones of the skeleton, and arrow-points.
of the beds and the conditions under which he discovered this very Copeland reported nine other mammoth specimens from this
important skull. The following is a brief resume of his description: swamp which he had deposited in a college at Forest Grove,
From a deposit of e.xtinct animals, discovered in 1876 in Oregon: "He discovered a flint spear-j)oint in the gravel above
the mammoth bones, associated with charred and partly petrified
wood that bore the marks of tools upon it, also deer, buffalo and
bird bones. ... I ne\'er doubted, from what I saw and hearil at the
other excavations in the immediate neighborhood, and where the
collectors went through the same kind of ])eat, clay and gravel as
we had gone through, that man, the buffalo, elei^hant and many
existing sf^ecies once lived together in eastern Washington."
Characters of the Skull (P^icis. 971, 973, 974, 976). This —
superb skull is certainly referable to Parelephas ivdnhiiigtonii,
and the specific reference is correct, because the exposure of the

V'6 nat size

Fig. 974. Young adult male skull (Amer. Mus. Cojic Coll. 8681) referred
to Parelephas roashinglimii Osborn. Left lateral aspect of the .same skull as
Fig. 973. Young adult male skull rpferrod to Pareleplms washingtonii that shown in figure 973 oi)posite. Also type jaw (Amer. Mus. 8681 A).
Osborn (Amer. Mus. Cope. Coll. 8681), from the Ivower Pleistocene of tlie The type jaw of P. wnsldiigliinii (.\mer. Mus. 8681A), found in the .same
state of Wa-shington. Right lateral aspect. One-eighth natural size. Same locality, namely, in the swam|)s of Pine Creek, Whitman Covnity, Washington,
skull as that shown in figures 901, 9()2, and 971
; also figure 974 opposite from about 100 miles from Walla Walla, belongs to a much older individual than the
photograph of left lateral as|M'ct. skull (Amer. Mus. Cope Coll. 8681). One-twelfth natural size.

'A large part of the skeleton, beautifully pre.served, was also recovered from the bed of gravel below (Science, N. S., 1903, Vol. X\"II, p. ,")12).
 THE MAMMONTINiE: PARELEPHAS 1103

third superior molar (Pig. 976) reveals the complete ridge formula. Catalogue) compares very closely with PareJephats washingtonii
Although found in the same deposits as the type, it belongs to Osborn. The posterior portion of the tooth is broken, but there
a much younger animal, because the third superior grinding teeth, are sixteen cross-crests present and there were probably from three
M^, are just coming into use, exhibiting only five or six worn ridge- to five additional crests in the complete tooth. The size of the
plates (Fig. 971 Al), whereas in the type jaw (Fig. 972) the second tooth, the number of the cross-crests as well as theii- thickness and
molars are completely erupted and the third molars exhibit degree of crenulation is quite closely identical with the type in the
seventeen to eighteen ridge-plates of the total of twenty-one. American Museum of Natural History, of which an illustration
is herewith published through the courtesy of Professor Henry
Fairfield Osborn, President of the American Museum of Natural
History, New York."

E. jetfersonii Osb.
Amer. Mus 9950 Type

Fig. 975. Type jaw (B) of Elephas [Parelephas] washinglonii (Amer.

Mus. 8681A), compareii with tyjie jaw (A) of Elephas [Parelephas] jeffersonii
(Amer. Mus. 9950).
Thi.s comparative figure clearly show.s the profoundly different form of the Amer. Mus,. 86SJ
rostrum and of the coronoid region. The measurements of the jaw may
readily be taken by multiplying the drawings in this figure and in figure 893
PARELEPHAS WASHINQTONII Pef.
eight diameters; both figures are drawn to a one-eighth scale.

Referred Second and Third Superior Grinders ok P.\relbphas

REFERRED PARELEPHAS WASHINGTONIt, PETERSON, 1928 washinqtonii, in situ

Peterson published the following description of a second left Contrast witli l.M^ of Parelephas floridanus type (Fig. 982)

inferior molar, I.M2, from Colorado, which he refers to Parelephas Fig. 976. Referred cranium of Parelephas washinglonii (Amer. Mus.
Cope Coll. 8681), from Pleistocene beds near Pine Creek, Whitman County,
washingtonii.
—
Peterson, 1928, pp. 118-120. "In the Pleistocene formation
state of Washington.
views in figure 971.
Same cranium as that shown in jjalatal and frontal

on Lay Creek, Moffat County, Colorado, about one half mile from Observe that the premaxillaries of the left side have been removed to
Lay Post Office, Mr. M. A. Langley found a left lower molar of completely expose the left third superior molar, l.M^, which exhibits 23 ridge-
plates, of which only the 5 anterior are worn or in use (as shown in palatal
a Proboscidean, which was later presented to the Denver Museum
view. Fig. 971). The second superior molar, 1.M-, is shown in profile, exhibiting
of Natural History by Mr. A. G. AVallihan of Lay, Clolorado. 10+ ridge-plates, the anterior plates having disappeared. This figure shows
Upon comparison this lower molar (No. 472, Denver Museum admirably the enormous alveolus for the reception of this grinding tooth.
1104 OSBORN: THE PROBOSCIDEA

Parelephas? eellsi Hay, 1926 and a lower, whicli are referred to Elephas coluinbi, a fragment of
Figure 978 a skull, which is here described, and tusks, one small and some
Port Williams, Clallam County, Washington. othei's of large size. These tusks the writer has not seen. Thej'
Elephas eellsi Hay, 1926. "Description of remains of an ele-
may belong to E. columbi or to the elephant forming the subject of
this paper. I regard the skull as belonging to a hitherto undescrib-
phant found at Port Williams, Washington." Journ. Washington
Acad. Sci., Vol. XVI, No. 6, March 19, 1926, pp. 154-159. ed species, and, with the intention of honoring the finder [Rev.
—
Type. Fragment of a skull. Horizon and Locality. "Port — Myron Eells, a missionary among the Twana and Clallam Indians
for over 30 years], I name it Elephas eellsi. At first view the speci-
Williams, Clallam County, Washington. This place is on the
southern shore of the eastern end of the Strait of San Juan de men is an unpromising one. It consists of a part of the left maxilla

Fuca." —
Type Figure. Op. tit., p. 156, figs. 1 and 2. and a smaller part of the right."
Type Description.— (Hay, op. cit., 1926, p. 154): "The col- Osborn, 1929: Genus and species indeterminate (cf. Parele-

lection made by Mr. Eells consists of two third molars, an u])per

phas washinglonii referred cranium. Figs. 973, 974).

Second left inferior molar of Parelephas washingtonii Ref.

Fig. 977. Left interior molar, I.M2, from Lay Creek, Moffat County, Fig. 978. Tyi)e figure of Elephas eellsi Hay, 192(1, figs. 1 and 2, one-fourth
Colorado (Denver Mus. 472), referred by Peterson, 1928, fig. 23, to Parelephas natural .size, found at Port Williams, Clallam County, Washington. Fragment
washingtonii. Oh.serve fifteen broad ridgo-plates with a maximum of seventeen, of a skull. An indeterminate sjieeies (eomjiare Parelephas washingtonii.
as compared with twenty-one in the type, I.M3 (Fig. 972), of P. washingUinii. Figs. 973, 974).
 THE MAMMONTINtE: PARELEPHAS 1105

PARELEPHA8 FL0RIDANU8 OF FLORIDA

A recent discovery of great interest a deposit near Bradenton, Florida, apparently of Upper Pleistocene
is

age, yielding the more or less complete remains of at least seven individual elephants, which represent a hitherto
unrecognized specific stage described by Osborn (1930.837) as Parelephas floridanus. The importance of this
discovery rests in the fact that it enables us to distinguish clearly this very large and progressive animal of
Florida from the typical Parelephas columbi, a specific name which, as shown on pages 1043, 1046, 1067, 1070-1083,
has embraced many quite distinct ascending mutations and specific stages. Near to this more primitive, true
Parelephas columbi the French Guiana species {Parelephas columbi cayennensis) appears to be ranked.

Parelephas floridanus compared with P. columbi and P. jeffersonii

The fossil mannnoths hitherto discovered in Florida have been determined by Hay, Gidley, and Osborn as
belonging either to the relatively rare Archidiskodon imperator or to a stage very near to the typical Parelephas
columbi from the Darien canal, Georgia, or contiguous phosphate beds of South Carolina. A third stage was
distinguished by Osborn in 1930 (1930.837) as Parelephas floridanus.

PARELEPHAS COLUMBI CAYENNENSIS ( S^axied ) Type

Skulls //S Natural sije
Tooth /4. •

PARELEPHAS TLORIDANUS (orthoqonaD AM 26820 Type

Fig. 979. Type (A) and Paratype (B) Crania and Jaws op Parelephas floridanus compared with Type (C) of P. columbi cayennensis
Crania one-sixteenth natural size; molar one-fourth natural size

A, Type (Amor. Mus. 26820), a middle-aged male with relatively short, obtuse tusks, exhibiting the second superior molar, M-, still in use, the third
superior molar, M'^, just coming into function. Compare attritional surfaces displayed in figure 981. The horizontal line x x indicates the attritional
level attained in the more aged paratype bull (B), Amer. Mus. 26821.

B, Aged bull with maximum full-grown tusks; second superior molar, M-, disappearing, third superior molar, M'', worn down to the very aged attritional
level (x x), as indicated in comparison with the younger bull (A).

Observe other age and growth differences in the depth of the jaw in A (cf. Figs. 983, 984). As detailed in the legend to figure 986, the outlines of the summit
of the occiput were restored from the more complete crania of Parelephas jeffersonii and P. washiyiglonii in the American and Nebraska Museum collections.

C, Ridge-plates 16-18 of type of Parelephas columbi cayennensis superposed on outline of complete r. M' of Parelephas columbi rcf. (Amer. Mus. 13708a),
from the phosphate beds near Charleston, S. C. Compare figure 957.
1106 OSBORN: THE PROBOSCIDEA

A.M.26820 Type

Fig. 980. Parelephas floridands Type and Paratvpe, Palatal Aspect

One -SIXTH natural size
Gift of Mr. Walter W. Holmes, 1929
(Right) Type molar crowns of the middle-aged adult (.\mer. Mus. 26820) with a total of 14-1.5 ridge-plates in use in the closely conjoined M" and M',
« hich appear to function like a single grinder.

(Left) .\ged paratypc (Amer. Mus. 26821) in which the 16-17 ridge-plates of the single third superior grinders, M', are simultaneously in function.
Compare figure 979 showing the attrition level of the grinders in the middle-aged adult and aged specimens. Observe also that in the type younger
bull14-lo ridgc-plates are in trituration, while in the aged bull paratvpe 16-17 ridge-plates are in use in correlation with the larger food requirements of the
more aged specimen.

Parelephas columbi. —As shown on pages 1075-1077 of the present Memoir, in the typical Parelephas
columbi the ridge formula is M 3 y^; to this typical stage, in which the grinding teeth are relatively small, belong
all the thirty-eight specimens from the phosphate beds of South Carolina fully described on the above-mentioned
|)ages, also the Amherst skeleton of P. columbi described by Loomis (1923-1928) on pages 1079-1081. In the
Amherst specimen the ridge-plate formula (M 3 j^) agrees closely with that of the South Carohna phosphate
collection, in which the prevailing ridge-plate formula is M3 iV-"iV+ -

Parelephas floridanus. —The type and referred individuals of Parelephas Jloridanus, varying in sex, size,

and by the ridge-plate formula

age, are all distinguished M
3 l-'^, indicating a much more advanced stage of
evolution; moreover, the type and paratype skulls of P. floridanus indicate an animal of very large size, with
crania and massive tusks differing in proportion from those of any Parelephas hitherto discovered, excepting
perhaps the "Franklin County Mammoth" (Neb. Mus. 1-4-15), referred to Parelephas jeffersonii.

This fine type collection of P. floridanus was the gift in the year 1929 of Mr. Walter W. Holmes of St. Peters-
burg, Florida, who with enthusiasm and generosity has been promoting the American Museum explorations in
Florida since the year 1923.

Materials. —The deposit near Bradenton, Florida, yielding the type, paratype, and other specimens, was
found in February of 1929 by Mr. J. E. Moore of Sarasota, who discovered the palate of the paratype specimen
 .

THE MAMMONTIN^: PARELEPHAS 1107

(Amer. Mas. 26821) ])rotruding from the side of the bank of a drainage canal. The whole deposit is 27 feet broad
and 10 feet dee]), the fine white river sand and black soil filling the interstices of the more or less broken crania;
this deposit was thoroughly worked by Carl Sorensen during the period March 2 where he re-
to AjmuI 2, 1929,
covered the remains of at least seven individual elephants. All the crania and jaws had been fractured to a greater
or less degree, transported a considerable distance, and collected in what may have been a deep marginal pool of
a low gradient river. These seven individuals, including type and paratype, are represented by parts of crania,
jaws, and the included grinding teeth.

The chief materials in this Holmes-Moore collection are as follows:

Amcr. IMus. 26820. Tyjic. MidcIle-aRod male. Skull and jaws (Figs. 979, 980, 981, with associated ribs, 1. scapula, 1. fibula,
r. ulna, 1. radius, two humeri, r. femur, two tibiae, parts of pelvis, and certain foot bones of the skele-
ton. Smaller right femur, length 12.50 mm. (4 ft. l)i in.). Smaller right humerus, length 1100 mm.
(3 ft. 1\, in.).

Amer. Mus. 26821. Paratype. Palate and jaws of an old male individual, third superior and inferior molars only. Right side
of cranium (Figs. 981, 979, 980), palate, lower jaw, very large tusk, also associated large vertebrse
(268336 in measurements below). Also larger left femur (1393 mm.). Larger left humerus complete,
length 1200 mm. (3 ft. II', in.).

Amer. Mus. 26822. Paratype. Palate with r.M' and liNP, of smaller size, supposed female.
Amer. Mus. 26823. Portion of right palate, with r.M^ of larger size, suppo.sed male.
Amer. Mus. 26824. Half of right palate, with r.M^ of small size, supposed female.
Amer. Mus. 26825. Third left upper, l.]\P, and left lower grinder, I.M3, complete, broad plated, of largest size, supposed male.

Amer. Mus. 26826. Right lower grinder, r.Ms, incomplete, (?)large size, supposed male.
Amer. Mus. 26833 a, h, c, d, e, /. Five vertebral series, scattered. See description and Table XIV on page 1114.

—
Geologic Age. Important as bearing on the relatively recent geologic age of this Bradenton deposit are the
horn and part of the cranium of a very large bison (perhaps Bison regius Hay), also part of the cranium of Castoroi-
des. Dr. G. G. Simpson (1930), who surveyed the mammalian deposits of Florida with the aid of Chief Geologist
Herman Gunter, considers this deposit as probably of late Pleistocene age.

Ridge-plate Distinctions. —The specific constancy both of the ridge-plates and of the molar measurements
as a whole estal)lish beyond question the clear separation of Pareleplias floridanus from the more primitive typical
/•. columbi, on the one hand, and from the more progressive P. jeffersonii and P. progressm, on the other, as shown
in the following comparison of the ridge-plate formulae and of the maximum or male diameters of the molar crowns
of M3:

Ridge-plates Length lireadth Height

Pareleplui, pwgressus |-|

205 mm. 109 mm. 203 mm.
Elephas roosevcUi[ = Parelephas jeffersonii ref.]. .
^
24
282 98 202

Parelephas floridanus ^
^^
^20
288
88
106
235
207
Parelephas columbi 1 6 +

In the "Franklin County Mammoth" of Nebraska, referred to Parelephas jeffersonii (pp. 1091-1093, Fig. 964
of the present Memoir), the tusks arc 12>2 ft. in length and strongly incurved; the third superior molar (M^) ex-
hibits 19 ridge-i)lates; with the possible exception of this Nebraska specimen, P. floridanus is the largest known
member of the Parelephas phylum in America.
1108 OSBORN: THE PROBOSCIDEA

Parelephas floridanus Osborn, 1929 and inferior ridge-plate fonnula: M3 §1+, max. |^^, intermediate
Figures 935, 936, 979-987, 989, PI. xxii between Parelephas colunibi (rif+) ^^'^ P- jeffersonii (f |) ridgc- ;

Manatee County, Florida, two miles south of Bradenton. Uppcr(?) j)lates broad and widely separated at base, more compressed at

Pleistocene. summit. Incisi\e tusks extremely massive and relatively short.

Parelephas floridanus Osborn, 1929. "New Eurasiatic and Males attain very large size. Femora measure 1250 to 1410 [1393]
American Proboscideans." Anier. Mus. Novitates, No. 393, mm."
December 24, 1929, p. 20. Type. — "Anterior portion of Supplementary Description (Osborn, 1930.837): "Parelephas
cranium, maxilla with M^, M' of both sides, and tusks, together floridanus from the Upper Pleistocene of Florida compared with
with lower jaw, Mo of both sides in situ, of a middle-aged indi- P. jeffersonii," Amer. Mus. Novitates, No. 443, December 18, 1930,
vidual, [Ms partly erupted]; also associated (?) right femur and pp. 1-17.
other slveletal parts." Amer. Mus. 26820. Paratypes. — General Characters (cf. Osborn, 1930.837). Comparison —
"Amer. IMus. 26821, adult jaw with M2, M3 in place, and Amer. is made with the full-grown type skeleton of Parelephas
naturally
jeffersonii (Fig. 988) and with the type dentition of Elephas
roosevelti [
= P. jeffersonii ref.j (Fig. 968). The incisive tusks are
shorter and more robust; the grinding teeth, while exhibiting
fewer ridge-plates, are of larger size and the ridge-plates are more
widely separated (see Figs. 980 and 985), with thick enamel, yet
clearly distinguishable from the broader crowns and still thicker
enamel of Arckidiskodon imperator of Florida. The type skeleton
and limbs of this middle-aged bull of Parelephas floridanus type
(Amer. Mus. 26820) slightly exceed in size those of the very aged
bull (Fig. 988) of Parelephas jeffersonii type (Amer. Mus. 9950).
In brief, the middle-aged Parelephas floridanus type agrees with
the aged P. jeffersonii in size, whereas the aged male paratype of
P. floridanus (Amer. Mus. 26821) greatly exceeds in size the aged
male type of P. jeffersonii, and is not far inferior to the giant
Archidiskodon imperator maibeni in the Nebraska Museum (Figs.
910, 911).
These general characters are very important in considering
Parelephas floridanus as a most welcome new stage or ascending
mutation in the long history of the Parelephas phylum first known
in the tapper Pliocene' Parelephas trogontherioides of western
Elurope.

SEPARATION OF PARELEPHAS FLORIDANUS FROM P. COLUMBI

By the older method of ridge-plate count and molar crown
proiJortions we may estimate the ascending mutation stage of three
specimens listed as P. columbi in the National Museum and
described on page 1079, namely:

Nat. Mus. 11810, r.M^, with 18 to 20 ridge-plates, length 258 mm.,

Fig. 981. Tyju' right and left superior and inferior molars of I'anicpltas breadth 90 mm., height 167-|-mm.
fluridanun (Amer. Mus. 26820), one-sixth natural size. After Osborn, 1929.797, Nat. Mus. 11808, r.M', with -fl3 ridge-plates preser\ed, length
fig. 19. 200 mm., breadth 89 mm., height 195 mm.;
Observe cspeciallj' the equal attritional areas above and below; fifteen 7 to 8 anterior ridge-plates missing.
combined superior ridge-plates in l.M-', opposed to 13-14 ridge-plates in Nat. Mus. 11806, l.M=, with -f- 1 2 -|- ridge-plates.
l.Mi-s, a perfect mechanical balance. Compare l.M-'' (Figs. 980, 982, and
legend), also I.M2.3 (Fig. 985, and legend).
Ridge-plate Heights. —By the newer height measurement
Parelephas floridanus (M 3 1y+) ""^ "^^Y exceeds P. columhi
Mus. 26822, fragment of palate with M'' of both sides; also as- (M 3 X6+) in the number of ridge-plates but in the height of the
sociated (?) left femur and other members of vertebral skeleton ridge-plates of the grinding teeth ; M' is much taller or more hypso-
of an individual of larger size (.\iiier. Mus. 26821)." Hohizon dont in P. flaridnniis than in P. rolinnhi; in the lypeof P. floridanus
.VXD Locality. -Manatee Coimty, Morida, two miles south of the eight superior ridgc-platcs (12-19) lueasure 1309 nun. in col-
Bradenton. Flu viatile fine sand, (?) Upper Pleistocene. Type lective height, whereas in the neotype of /'. columbi the correspond-
FiGUKE.— 0/>. Osborn, 1929.797, p. 19, fig.
(77., 19. ing ridgp-i)lates (12-19) measure 1117 mm. in their collective
Specific CfiAHAiPEKs. (0/>. ciL, 1029.797, p. 20): "Superior height. In Nat. Mus. 11808 the collecti\'e height of the eight

'[See footnote 1 on page 1049 above. — Editor.)

 THE MAMMONTINtE: PARELEPHAS 1109

ridge-plates (13-20) is 1196 mm.; it is therefore intermediate in COMPARATIVE DENTAL MEASUREMENTS OF PARELEPHAS
collective height measurement between P. floridanus (1309 mm.) JEFFERSONII AND P. FLORIDANUS
and P. columbi (1117 mm.) Consequently the National Museum Parelephas floridanus Type and Referred Molars (see
specimens above are nearer to Parelephas floridanus in measure- Table XII).— The total number of ridge-plates in M' o( Parelephas
ment than to the typical P. columbi, to which they were first floridanus is 22+, in Ms it is 21-)-, as compared with M 3|f in
referred by Osborn (see p. 1078). Elephas roosevelH [
= Parelephas jeffersonii ref.] (Fig. 968).
The ganometrir length (Osborn-Colbert, 1931.858) of the —
Ridge-plate Count. According to the ridge-plate count of
superior and inferior ridge-plates of an elephant grinder can only the type and paratype molars of Parelephas floridanus taken along
be measured in longitudinal section measurements on unsectioned
; the central line of the crown, in smaller males and fefnales there
teeth are liable to be erroneous up to as much as twenty-fi\e per- are 6/5 ridge-plates in 100 mm.; in the male type TA in 100 mm.,
cent. as compared with 8 ridge-plates in a 100 mm. line in P. jeffersonii.
1110 OSBORN: THE FROBOSCIDEA

Type Mandible of Parelephas kloridanus Paratype Mandible of Parelephas flohidanus

Fig. 983. Type mandible of Parelephas Jloridanus (Amer. Mus. 26820). Fig. 984. Deeply depressed aged mandible of the |)aratype of /'rtrc/f p/wis
Oue-sixth natural .size, .\fter Osborn, 1930.837, fig. 4. Jloridanus (AmrT. Mus. 26821), exhibiting only M.i in situ, Mo having been
Observe the very robust proportions, strongly abbreviated or truncated .shed. One-sixth natural size. After O.sborn, 1930.837, fig. 5.
rostrum, wholly worn r.M2, and partly worn r.Ms. Compare figure 981. Observe the very prominent, peaked rostrum, also the relatively greater
lower border of the mandible below the
balhtjceplialy or depression of the
level of the condyle, as compared with the more youthful type mandible
opposite.

.A,s newly restored (l''i{i. 986), tlie tyi)e tusk (Amer. Mus.
2GS2()) is much loiifior than shown in figure 979. The ))ariitype
tusk (Amer. I\Ius. 26S21) is correctly reiiresented in figin-e 979;
it is apparently from the extreme wear of M'';
full f;ro\vn, jiidfjiing

it measures 2320 mm. as compared with 3500 mm., length of the

lypc aged male tusk of P. jctfernonii (Amer. Mus. 9950). In brief,
in the aged males of P. Jloiidaiiiis the tusks are more massive, less

incurved, and relatively sliorter (=2320 mm.) than the aged male
tusks ( =3500 mm.) of /'. jrlfcrsniiii.

COMPARATIVE CRANIAL AND SKELETAL MEASUREMENTS OF

LM2 PARELEPHAS I'LORIDANUS AND P. JEFFERSONII
As shownTable XIII and figm-es 979 and 9S4, the crania
in

:uid jaws of P. Jloridanus greatly exceed in size and in massiveiiess

the aged type cranium of P. jeffersonii. The ratio of increase in
size in the jaw across the premaxillaries is about 10 per cent. The
estimated bathycei)halic measurement from the summit of the
Type Inferior Molars of the Lfft SroK ok I'auki.ephas floridanus lOOOe mm. us
occii)ut to the occlusal surface of the grinding teeth is
Fig. 98.'>. Type (Amer. Mus. 2tj820) internal and crown views of comiiared 880 mm. in /'. jrffcrKoiiii
with The longitiidinai .

second and third inferior grinding teeth, l.Mo, l.M.i; I.M3 exhibits 21 +
nieasurenient from the occipital condyle to the anterior rim of the
ridge-plates. One-sixth natural size. Compare figure 981, coronal aspect
of ty|x; suijcrior molars of both sides, and inferior of left side. After orbit is 770e mm. as compared with 720 mm. in P. jcfferisoiiii. It
Osborn, 1930.837, fig. 7. thus api)cars that P.fluridanus is much more bathycei)halic (lOOOe
 THE MAMMONTIN^: PARELEPHAS nil

The aged bull paratype (Amer. Mus. 26821) of P. flori-

danus (left humerus, 1185 mm.; left femur, 1393 mm.) ex-
ceeds in size the aged male type (Amer. Mus. 9950) of /-".

jeffersonii. In fact, the mea.surenicnt of the humerus some-

what exceeds that of the Amherst skelet(jn of Parelephas
columbi (Amherst Mus. 25-1) and is not far inferior to the
measurements of both the humerus and femur of the giant
.4.imperalor maiheni of the Nebraska Museum (Neb. Mus.
5-9-22).

Reconstructed Ttpe Cranium of Parelephas floridanus beneath

Restoration of Parelephas jeffersonii
Fig. 986. Type male cranium of Parelephas floridanus (Amer. Mus. 26820) as
now exhibited in the American Museum, Hall of the Age of Man. Compare figure
979. After Osborn, 1930.837, fig. 8.
In this reconstruction, made under the direction of the author and Mr. Charles
Lang, the height of the orbit is determined from the more complete paratype
(Amer. Mus. 26821); the frontal profile and occipital region are determined partly
from the cranium (Amer. Mus. Cope Coll. 8681) of the more primitive P. washirig-
tonii, partly from the type (Amer. Mus. 9950) of P. jeffersonii, and partly from

the giant P. jeffersonii in the Nebraska Museum known as the "Franklin County
Mammoth" (Neb. Mus. 1-4-15).

mm.) than the relatively primitive referred cranium of P. washiiuj-

tonii (Amer. Mus. 8681) in which the bathycephaly is: height
880 mm., length 700 mm.
Increasing bathycephaly is also indicated in the mandible by
comparison of the type jaw of the adult bull of middle age (Amer.
Mus. 26820— Fig. 983), exhibiting the wholly worn M, and partly
worn M3, with strongly abbreviated rostrum. Much deeper or more
bathycephalic is the aged paratype jaw (Amer. Mus. 26821 Fig. —
984) retaining only the much worn third inferior molar, M3.
Comparison of Shoulder Height. —As shown in Table XIII,
the skeleton and limbs of the type middle-aged bull of P. floridanus
(Amer. Mus. 26820) exactly equal or slightly exceed in size the
very aged bull type of P. jeffersonii (Amer. Mus. 9950) measured
as shown in figure 988. This statement is proved by the compari-
sons below.
1112 OSBORN: THE PllOBOSCIDEA

A few coiujjarative iiieasiireiaents witli Parelephax culumbi Right humerus of /'. jcjjcrnoiiii, aged Ijull typo 1 120
and Archidiskodon imperalor are also significant, as follows: Right humerus of P. coliimbi, Amherst skeleton 1030

Observe especially from the relative proportions of the

Hight liuiiierus of ,1. imperalor maihcni of tlie
humerus that the aged bull of J'arclcphasfloridaitiis greatly exceeds
Nebraska Museum 1251 mm.
in shoulder height, size, and all other dimensions the aged bull of
Left humerus of aged male P. Jloridanim \nu-d- P. jeffersonii. The comparative measurements of the humerus
ty])e 1185 are very important, because the humerus always forms the most
Right humerus of tyi)e younger male P. reliable method of estimating the shoulder height of any member
floridanus 1140 of the elephant family.

l'"ig. 988. New .sUiiihuil iiictliod (1930) of skeletal mea.surement in I'arckpkas ami other extinct and living pruboseideiiiis, illii,sliate<l on the
type skeleton (Amcr. Mus. 9950) of Parekplias jeffersonii in the American Museum collection, Hall of the Age of Man. After Osborn, 1930.837,
fig- 9-

Bathycephaiy of cranium
vertical 880 mm.
horizontal 720
Jaw
length, condyle to tip of ro.strum 675
Incisive tu.sks
in socket 480 mm., exposed 3020 mm., total 3500
Scapula
vortical 900
horizontal 617
Humerus, articular length 1085
Femur, articular length 1255

This mclliod is uniform witli that prc'sent Memoir for tlu; measurement of the (Taiiium, tusks, and fore- and liind-
adopted throughout the
iimbs (cf. caption to Fig. 868). marked falling away of the posterior part of tlie v(!rt(!bral (column and the relative abbreviation of the
Observ(? the
hindlimbs, also characteristic of Mainmonlcus primigeniiia and in a less degree of Archidiskodon mcridionalis.
THE MAMMONTIN^: PARELEPHAS 1113
1114 OSBORN: THE PROBOSCIDEA

FIVE VERTEBRAL SERIES PROBABLY ASSOCIATED WITH FIVE OF THE CRANIA OR JAWS. COMPARE TABLE XIV.

Several scattered series of vertebrae ha\e been reassembled

according to size which represent five, or at the most six, indi-

viduals, numbered as follows: Amer. Mus. 268.33a, 268336,

268336', 26833rf, 26833e, 26833/. Of these, one vertebral series
(Amer. Mus. 26833a) may with some probability be associated
with the type (Amer. ]Mus. 26820) cranium and jaws; while
another series (Amer. Mus. 26833b) may be associated with the
paratype cranium and jaws (Amer. Mus. 26821).
In the assembhng, partly by order of size, these vertebra]
 THE MAMMONTIN^: PARELEPHAS 1115

Whereas, as appears in Table XI V,the trans\erse measurements measurements of ]'. Jlnridnnus display great irregularity, owing to
of Parelephao jefferftnnii are fairly constant, similar transverse irregular disposition or distortion and to possible errors in our
assemblage.

Refeered Right Third Inferior Molar of Paeelephas ploridanus

Method of Ridge-plate Measurement
Fig. 989. Total length 274 mm., maximum liciglit of fourteenth ridge-plate, 18.0 mm.
A finely preserved right third inferior molar, r.Mj (Fla. Surv. V-4.')29) from Fig I.sland,
Itchatueknee River, Florida, referred to Parelephas floridanus. Courtesy of the Florida
Geologieal Survey.
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 Chapter XVIII

THE GENUS MAMMONTEUS' (SUPERFAMILY ELEPHANTOIDEA), OF THE

SUBFAMILY MAMMONTIN/E, THE TRUE NORTHERN
WOOLLY MAMMOTH
Early descriptions of the mammoth during the seventeenth and eighteenth centuries (1696-1788)
IN EURASIA. History of eurasiatic and north American discovery. Characters of mammonteus primi-
genius and of supposed ancestral stages leading back to upper pliocene time. Extreme stages of
evolution in AMERICA. FEEDING HABITS AND GEOGRAPHIC DISTRIBUTION.

I. Historical Introduction (1696-1788). Mammonteus primigenius.

Names successively applied to the mammoth. Typical ridge formula.
Native Siberian origin of the word Mammut. Skulls and jaws of the true Mammonteus primigenius.
'Adams skeleton.'
II. Characters of the Sitbfamily Mammontin^, including
2. Primitive European stages of Mammonteus primigenius.
THE Genus Mammonteus.
Mammonteus primigenius leith-adamsi.
1. External characters and feeding habits. Mammonteus primigenius hydruntinus.
2. Skeletal characters of Mammonteus primigenius. Mammonteus primigenius fraasi.
3. Historical order of naming of species of Mammonteus. Mammonteus primigenius astensis.
4. Geologic and provisionally ascending phyletic order of .3. American stages of Mammonteus.
species and subspecies of Mammonteus. Eurasia and
Mammonteus primigenius americanus.
North America.
Mammonteus primigenius compressus.
III. Systematic Description of Species of Mammonteus. Mammonteus primigenius alaskensis.
1. Typical progressive Eurasiatic stages of Mammonteus. 4. The frozen Mammoth of Siberia.

I. HISTORICAL INTRODUCTION (1696-1788)

The subfamily name Mammontinae Osborn, 1921, replaces the name Euelephantinse Osborn, 1918, because
the subgenus Euelephas Falconer is invalid (see Chapter XIX, p. 1177); the term Mammontinae, signifying 'les

mammonts,' the 'mammoths,' was substituted in 1921. As set forth in the two preceding chapters, the Mam-
montinae embrace three genera, namely, Archidiskodon of the southern and .south temperate zones, Parelephas
of the intermediate and north temperate zones, and Mammonteus^ of the northerly and circumpolar zones.

It is probable that these northerly or woolly mammoths were the first mammalian fossils of northern Eurasia
to be discovered and recognized as extinct. The earliest descriptions are naturally lost in obscurity. Of this
early period Falconer (1846, pp. 11 and 12) writes:

—
Next, in regard to the establishment of the species. The fossil remains of the Mammoth had, during ages, attracted more or
every country in Europe, having been found in England and in all parts of the Continent, from Italy to Siberia.
less attention in
But it was only towards the close of the last century that definite notions as to the species, were arrived at. Pallas, who had
better opportunities for determining the point than any of his contemporaries, upon the perfect remains so commonly met with
in Russia, erroneously considered the fossil teeth to be identical with those of the Indian species. A great advance was made in
the inquiry through the disco\'ery, by Peter Camper, of the specific difference between the teeth of the Asiatic and African Ele-
phants [Footnote: 'P. [A]. Camper, Descript. Anatom. d'un Elephant male, p. 16.'], when Blumenbach and Cuvier almost

HThe generic name Mammonteus (Mammonteum Camper, 1788, pp. 251, 259; Osborn, 1924.633, p. 2) for the northern Mammoth of doubtful validityi.s

as Camper's description has reference to an animal in .\mcrica and not to the Mammoth, moreover lie u.sed the word "mammonteum" in an adjectival sense.
Dr. A, Tindell Hojiwood in his recent (1935) memoir on the "Fossil Proboscidea from China," p. 97, adopted the generic term Mammulhus Burnett, 1830,
genotype Mammuthun borealis. For a historical account of the names applied to the Mammoth and the Mastodon, see Chapter XXI below on Nomencla-
ture. — Editor.]
1117
 :

1118 OSBORN: THE PROBOSCIDEA

simultaneously entered upon the investigation, and arrived at the same result, \'iz., that the Mammoth was an extinct form, dif-
fering from both of the existing species. Struck with the length of the cranium, and of the incisive sheaths in the Mammoth, as
represented in the figures of Messerschmidt's specimen attached to Breyne's excellent remarks in the Philosijihical Transactions
[Footnote: 'Phil. Trans, vol. and connecting these peculiarities with the great width of the crown, and
xl. 1738 [1741], p. 124.'],

the narrowness and number was conducted to his first happy conclusion. The prob-
of the plates in the fossil grinders, Cuvier
ability of a similar difference characterizing the species in other fossil genera, Hashed across his mind, and opened to him new
views respecting the theory of the earth. Great and important were the results; and after they had been achieved, the illustri-
ous Anatomist reverted, in terms of the liveliest acknowledgment, to the long neglected figures of Messerschmidt, which had
helped him to the first idea [Footnote: 'Cuvier, Oss. Fossil, tom. i. p. 178.'].

ScELETO Elephantino Tonn^e, 1695, CONFUSED WITH THE Mammoth, Blumenbach, 1799
In 1695 Liidolf described the mammoth of Siberia. The earliest account of the fossil elephant in Germany
is that of Wilhelmus Ernestus Tentzelius' (Phil. Trans. Roy. Soc, London, 1698, Vol. XIX, pp. 757-776):

—
Tk.ntzelius (Tentzel), 1698, pp. 757, 758. III. Wilhelnii Ernesti Tenlzelii Historiographi Ducniis Saxonici Epislola de
Sceleto Elephaidino Tonme. Tonna inter Thuringiae Dinastias baud po.strema,
. . . ((uorum alter appellatur Burg-Tonna, . . .

. .quam effodientes mense Decembri sujierioris anni ossa qusedam maxima reperiunt,
. Prior sententia mihi cumprimis . . .

placet, quam ita defendere aggredior, ut primo ostendam, sceleto nostro omnia convenire, quae ad vera elephanti ossa requirun-
tur; deinde evincam, non esse minerale fossile, sed animale {letrefactum; denique inquiram, rjuomodo in has terras ipsuuKiue
locum pervenerit Elephantus.

This classic or original description of the 'elephant' discovered in a sand-pit at Burgtonna near Gotha,
Thuringia, is of the skeleton referred to by Blumenbach in his type description of Elephas primigenius cited
below, but this is not to be regarded as the type specimen.- It is addressed "ad Virum toto orbe celeberrimum
Antonium Magliabechnim, Serenissimi Magni Hetruriae Duds Bihliolhecarium & Consiliarium.." Following the
dedication Tentzelius gives a description of the discovery in the month of December, 1695; he fully describes

l)arts of the teeth, skull, and skeleton found, and discusses local opinion as to whether they were the remains of
a petrified elephant or of a fossil unicorn (p. 758) ; he cites the descriptions of John Ray, of Pliny, and of Aristotle

to prove that the specimen from Tonna was an elephant ; he assures the most learned and celebrated Magliabechi
that the bones and teeth correspond point by point with those of the elephant; he proceeds with a comparison
with the Ceylon elephant (p. 771) and with Pigafetta's descriptions of the African elephant; he cites Steno as to
the habits and distribution of the Indian and African elephants, and concludes that the Burgtonna remains
(p. 774) constitute proofs of the Flood, of the rise of the waters even to the height of the Thuringian mountains.

The Editor of this classic contribution to the Philosophical Transactions, (Gothae, 1696, p. 776), concludes:

The Author of this Letter has favour' d the Royal Society with some pieces of Bones of the Sceleton of this Elephant, viz. part
the

of the Skull, wherein appear its Cells, some of the Teeth both of those that grind, and such as are called Elephants Teeth or Ivory,
with some other pieces of Bones, all which they found agreable to his Description, and ordered (hey should be carefully preserv'd in
their Repository.

Thirty-one years later Sir Hans Sloane (Phil. Trans., 1729, pp. 457-494, 497-514) refers to this discovery as
follows (p. 508)

The Skeleton of an Elephant which was dug up in a Sand-pit near Tonna in Thuringcn, in 1095, is one of the most curious,
and most compleat
also the in its Kind, forasmuch as they found the whole Head, with four (irinders, and the two denies exerti,
or Tusks, the Bones of the fore and Hind-legs, one of the Shoulder-bones, the Back-bones, with the Ribs, and several of the

'Karl von Zittcl ("HLstory of Geology and Palaeontology," 1901, p. 134) refers to thi.s discovery as follows: "The skeleton found at Burgtonna in 1090 was
one of the most famous discoveries, as it gave rise to a dispute between Ernst Tentzel and the medical faculty in Gotha. The other professors saw in the large
hones only sports of nature, hut Tentzel proved to their discomfiture tliut the bones were real, and h.id belonged to elephants."

-The Burgtonna skeleton belongs (Dietrich, 19.30) to Eliphas antiquus; while first mentioned, it is not th(' type. In defining and describing EUphas
primigenius (1799), Blumenbach had in mind the true mammoth of Siberia and Germany.
 THE MAMMONTIN.E: MAMMONTEUS 1119

Vertebrae of the Neck. But the whole hath been so accurately described by Wilhelmus Ernestus Tenlzelius, Historiographer
to the Dukes of Sai-ony, in a Letter to the learned MagUabechi, printed in the Philosophical Transactions [Footnote: 'No. 234,
pag. 737.'], that it is needless to add any thing, the rather, as that Gentleman was pleased to oblige the Royal Society with some
Pieces of the Bones of this Elephant, with Part of the Skull, wherein appeared its Cells, some of the Grinders, and Part of the
denies exerti; all which being produced at a Meeting of the Royal Society, were found exactly agreeable to his Description, and
ordered to be carefully preserved in their Repo.sitory. [See footnote 1 below.]

Breyne's Description (1735) of the Elephas primigenius of Siberia

A second scientific description of the mammoth, from the river Indigirka, Siberia, with figure of the skull
(Fig. 991), was a communication to Sir Hans Sloane, President of the Royal Society, by John Phil. Breyne, in
a letter dated "Dantzick, Sept. 28, 1735," from which the following excerpts are made (Phil. Trans. Roy. Soc,
London, 1741, Vol. XL, pp. 124-138).
A Letter from John Phil. Breyne, M.D.F.R.S. to Sir Hans Sloane, Bart. Pres. R.S. with Observations, and a Description of
some Mammoth's Bones dug up in Siberia, proving them to have belonged to Elephants.

Sir,
Your very learned and instructive Accounts of Elephants Teeth and Bones found under Ground, I saw with great
Pleasure in the Philosophical Transactions, N° 403. and 404. In the same Year, to wit 1728, I was busied about the very
same Matter, especially to prove, that the extraordinary large Teeth and Bones found under (Jround, and digged up
in several Places of Siberia, by the Name of Mammoth's, or Mammut's, Teeth and Bones, were.

I. True Bones and Teeth of some

large Animals once living; and,

Fig. I . n . sl .
II. That those Animals were Ele-
phants, by the Analogy of the Teeth and
2 94 jnrn Bones, with the known ones of Elephants.

11 ^ginches III. That they were brought and left

there by the universal Deluge. I made hke-
wise several useful Inferences about This
Matter.
At the same time there flourished in our
City a Society of some learned and ingen-
ious Gentlemen, who met once a Week in a
certain Place: In one of those Meetings in
the Month of March, I had the Honour to
read and communicate my Thoughts and
Observations about this Subject; which,
as I believe, they will not be disagreeable
to you, I have translated into the English
Tongue, and joined to this present Letter.
After that, viz. in the Year 1730, Dr.
Messerschmidt returned to Dantzick, from
his Travels thro' Siberia, and was pleased
to communicate to me some curious
Draughts of a Part of a Skeleton, to wit,
jx..^r,. of a very large Skull, Dens exsertus & mo-
laris, with the Os femoris, belonging to the
The Messerschmidt Cranium of Siberia. Copied by Ccvier Animal commonly called Mammoth, found
Fig. 991. Cranium of the mammoth, and figured by Dr. John Phil. Breyne in the
as described in Siberia; by which our Assertion, that
Philosophical Transactions of the Royal Society of London, 1737 (printed in 1741, Vol. XL, No. 446, the Teeth and Bones, called in Rusland
pp. 124-138, PI. I, figs. I, ii). One-sixteenth natural size. This classical Breyne figure of the mam- Mammoths Bones, are the true Teeth and
moth cranium was reproduced by Cuvier in his "Ossemens Fossiles" of 1806 and used as the basis Bones of Elephants, is not only, as you
of comparison of the E. primigenius cranium with the crania of E. africanus and E. indicus (Cuvier, wished in your first Account, put in a
1806.1, PI. XXXIX, fig. 1, and PI. xli, figs. 9.1, lO.A, ll.F.— our Fig. 992). Made by Brandt (1833.1,- greater Light, but, if I am not mistaken,
p. XII) the type of Elephas giganteus. clemonstrated beyond all Doubt.

'Hopwood (letter, May Royal Society records (November 3, 1697) reveal that the Tentzel specimens were assigned the number
20, 1930) finds that the
120B in various lists uj) to the "It would seem, then, as though the specimens were lost between 1733 and 1763."
year 1712: Fragments of the Burgtonna
—
skeleton of Elephas antiquus, incisive tusks and other remains, are still preserved in the Gotha Museum (fide Dietrich, letter, April 14, 1930 see Chap. XIX, the
Loxodontinae, p. 1181).
^[See footnote on page 1136 regarding the date of description as 1832. —Editor.]
 —

1120 OSBORN: THE PROBOSCIDEA

Observations on the Mammoth's Bones and Teeth found in Siberia: Read in a Meeting of some learned Gentlemen at
Dantzick in the Year 1728. by J. P. B.

That learned and curious Gentleman Dr. Daniel Gottlieb Messerschmidt, who was sent some Years ago, by his late
Cznrish Majesty, Peter the Great, into Siberia, to search after the Products of Nature in this iminhabited and cold
Country, was pleased to send me in the Year 1722, amongst some other Samples of Natural Things out of Siberia,
two very large Teeth, called there, Mammoth or Mammut's Teeth, with
the following Inscription: Dens molaris,
ut videtur, diluvianus, Bellux cujusdam hactenus
pro Elephantino habendus sit, cujus jam penes Te
incognitse, nisi
esto arbitrium, Russis Mammoth, repertus in Montium altissimis jitgis ad Thomam fluvium. Alterum est frustum.
aliud Eboris Denti exerto Elephantis non absimile, ab aliis repertum in Thomse Montibus.

[p. 137] The bones of this Skeleton, with the Ribs, Vertebrx, and others thereto belonging, were found in the sandy
Side of a steep Hill, on the Eastern Bank of the River Indigirska [Indigirka], which falls into the Northern Ocean, not far
from the Mouth of the Rivulet Wolorkowoi rurzei.

NAMES SUCCESSIVELY APPLIED TO THE MAMMOTH

Among the first descriptions of species, to which Latin names were appended, are the following:

1696 Mammontova Kost [Mammotovoi host in Ludolf, 1696.1, p. 92] (probably derived from the Tatar word mamxi signifying
earth and A:o.s/ signifying ivory = tusks) Ludloff [Ludolf]. Cited by Fischer de Waldheim, "Oryctographie," III,

Fossiles du Gouvern. de Mo.scou, 1830-1837, p. 111.

"I. Elephant. 1. Le Mammont . . . Elephas mammonteus. m. [Fischer, 1830-1837] .Je conserve la denomi-
. . .

nation la plus ancienne, et je n'ecris point Mammouth parceque ce nom n'a pris origine que par corruption ou une fausse
lecture du mot Mammont [Footnote: MaMMOHiiiTi en rus.se; le Ver (t.) terminal a etc change par les anglais en
h et le n a ete pris pour un u. D'ailleurs Ludloff est le premier qui en parle (1696) et appelle ces ossemens Manl-
montova Kost et justifie ainsi ma denomination .systematique.']."
Cited by Cuvier (Ann. Miis. VIII, 1806, p. 45): "C'est sous le nom de comes de mammont, mximmontova-kost,
qu'ils designent les defenses."

1788 Mammonteum Camper (Nova Acta Acad. Sci. Imp. Petropol. Communicanda, II, 1787, p. 251).
". OS humanum petrifactum, aut fossile, etiamsi Mammonteorum, Elephantorum,
. . Adserere ex eodem . . .

principio audeo Mammonteum animal extinctum non modo esse, sed nullam omnino habuisse cum Elephanto
.similitudineml"

1796 Mammoth, Cuvier [MS.] "Mi'moire sur les especes d'Elephans tant vivantes que fossiles," etc.
"Whatever may have been the approximation previously made by
(See I'alconer, 1868, Vol. II, pp. 158, 159):
Merk or Blumenbach towards a Mammoth from the two living species, Cuvier was undoubtedly
distinction of the
the first to characterize the extinct species with exactness, in his joint memoir with Geoffroy, under the name of
Elephas Mammoth, in the year 1796 [Footnote: 'Mem. de ITn.stitut, 1" C'lasse, tom. ii.']. In the same year, he read
a memoir at the first public meeting of the 'Institute,' but which was not published until 1806 [1799], in which the
diagnostic marks are very pointedly expressed under the designation of Elephas Mammonteus: 'Maxilla obtusiore,
lamellis molarium tenuibus, rectis,' as distinguished from Elephas Indicus: Fronte plano-concava, lamellis molarium
arcuatis, undatis.' ('u\ier connected these dental and manilibular distinctions with others yielded by Me.sser
Schmidt's [Daniel fiottlieb Messerschmidt —see Breyne, 1737 (1741), Pis. i, ii, iii] figure of the skull of the Mammoth,
and combined th<' whole in the extended specific definition of the extinct form, which appeared memoir of 1806 in his

'L'J^k'phant k crane allonge, a front concave, a tres longues alveoles des defenses, a machoire inferieure obtuse, a ma-
chelieres plus larges, paralleles, marqu('>es des rubans plus serres.''^' He abandoned the name E. Mammonteus of his
memoir of 1796 [1799], and adopted the designation of Elephas primigenius, proposed by Blumenbach [Footnote:

'Cuvier's definition (1800.1, p. 2f;4) "A'ci('^/ia7j/ d cra^tc aHojtgi', ... que noii.s nommon.s iUphanl fosaiU' (elephas primigenius, Blunipiib.), est le mammonl
des Riis.ses," in contrast to (op. cit., p. 2()'2) "L'elephant a crime arrondi, . . . (elephas africanus)," also to (op. cit., p. 2(13) "L'Hfphanl a crane alhnge . . .

(ele phas indicus) ," i.s clearly illustrated in his Plates xxxix and xi.i, in which he places Breyne's fiKure of Elephas primigenius with his own figures of Elephas
indicus and Elephas africanus. This definition was literally repeated in the three successive editions of the "Os.semens Fossiles" (1812, 182."), 1834) and exert-
ed a profound influence on other |)aheontologists, for it was not until 182.') that the mammoth of the south {Elephas meridionalis) was named by Ncsti, in
contra.st to this elephant of the north (Elephas primigenius) with curved tu.sks, also to Elephas antiquus I'''alconer (1847), with straight tusks.
 :

THE MAMMONTIN^: MAMMONTEUS 1121

'Voigt's Mag. 1803, Band v. p. 16.'], in 1803, which is that now generally accepted among palaeontologists [see
Blumenbach, 1799, p. 697, for first use of name Elephas To
normal form, as already stated,
primigenius]. this
Cuvier referred all the fossil remains of Elephants found over the whole of Europe, in Northern Asia, and in North
America, however much at variance with the terms of his definition and to the last he clung to the specific unity of
;

the 'Elephant fossile' with the jealous partiality of a discoverer for the earliest results of his most cherished labours.

The Mammoth, as compared with those of the existing Indian Ele-

distinctive characters in the molars of the
phant, upon which Cuiver reUed, may —
be expressed in the following terms: 1. Great narrowness or compression and
approximation of the crown-ridges, involving both a larger number in the same length of crown and in triturating use
at the same time. 2. Tenuity of, and absence of crimping in, the enamel-plates. 3. Greater width of the molar-
crowns, both absolutely and relatively to their length."

1796 Elephas mammonteus Cuvier (1796 [MS.], published August-September, 1799).

(Cuvier, 1799, p. 21) "les caracteres distinctifs des quatre especes que ja'i decrites dans
: le cours de ce memoire.
Les voici
Especes qu'on sail exister.
Elephas capensis [Cuv. =Loxodonta africana Blum.], /ronie convexd, lamellis molarium rhombo'idalibus.
Elephas indicus [Linn. = Elephas indicus Linn.], froute plano-concava, lamellis molarium arcuatis undalis.

Especes qu'oji ne con/wit que fossiles.

Elephas mammonteus [Cuv. = Elephas primigenius Blum.], maxilld obtusiore, lamellis molarium tenuibus rectis.
Elephas americanus [KerT = Masiodun americanus of the present ]VIemoir], molaribus muUi-cuspidibus, lamellis
post detritionem quadri-lobatis."

L' elephant it cnine allonge'., a front concave, a tres-

longues ah^ules des defenses, h mdchoire inferieure ob-

tuse, a mdchelieres plus larges, paralleles, marquees de
rubans plus serre's, que nous nommons elephant f ssih ( eU-
phas primigenius , Blunienb. ) , est le mammont des Russes.

B Fi^. 2. /7y. ^. I.
L' elephant ii crane allonge , ti front concave , it petites
oreilles , ii mdchelieres marquees de rubans ondojans que
nous appelons elephant des Indes ( elephas indicus ) , est un
quatliupcde qu'on u'a observe d'une maniere cerlaine qu'au-
dcladc rindus.

C /z>. J. /i>. M A.
U Aiphanl it crane arrondi it larges oreilles
, it mdche- ,

lieresmarquees de losanges sur lew couronne, que nous

appelons elephant d Afrique ( elephas africanus ) est un ,

quadrupede dont la seule patrie connue est jusqu'a present

I'Afrique.

Fig. 992. Comparison of crania of Elephas \Mammonleus\ primigenius, E. indicus, and E. [Loxodonia] africanus, after Cuvier, 1806.1, PI. 39 (ii), figs. 1, 2, 3,

and PL 41 (iv), figs. 9.1, One thirty-seventh to one-twentieth natural size.

lO.A, and U.F.
Figures 1 above and 11. F below arc two views of the same skull (Messerschmidt, see Fig. 991), first figured by John Philip Breync (1741.1, PI. i), subse-
quently figured by Cuvier (1806.1, PI. 39, fig. 1, and PI. 41, fig. U.F), and finally made the type of Ekphas giganteus by Brandt (1833.1,' p. XII).
The descriptive captions at the right are taken from Cuvier {op. eil., 1806.1, p. 262 {E. africanus), p. 263 (E. indicus), and p. 264 (E. primigenius).

Cuvier's designations were: "le Mammouth" (Review, Cuvier, an 3 [1795], p. 90'); ElephoH mammonteus
Cuvier, 1796, MS., published 1799, Fructidor, an VII [August-September, 1799], p. 21; not until 1806 did he
adopt the specific name of Elephas primigenius Blumenbach.
'[See footnote on page 1136 regarding the date of description as 1832. — Editor.]
1122 OSBORN: THE PROBOSCIDEA

1799 Elephas primigenius Blumenbach ("Handbuch der Naturgeschichte," Sechste auflage, p. 697).'"
3) Von einem ungeheuer grossen Elephanten (Elephas primigeniusf) [die vermeinten Riesenknochen** unsrer
ehrlichen Alten]; unter andcm aiich in Menge in Dcutschland***). So z. B. das boriichtigte Elephantengerippo
das 1695 bey Burg-Tonna ini (iothaischen ausgcgraben worden etc.

s. Voigts Magazin. V. B. I. St S. 16 u. f.

•(Kriegsr. Merk) letlres sur les os fossiles d'elephans cl de rhinoceros qui sc Irouvent en Allcmagnc ttc. I-III. St. Darmst. 1783 u. f. 4.

i^A^aTuTnl size JJrawn from cost

A.M. 26981

"'^>r.^
20
Zl
^2\2i

7 8 S JO „ ji ,j uiJi-

)6\l7\!a\!3-.20\^' ,^^ \

LM
Drawn from
A M.Z69ao
^^v \: ^'
From Siberia from Osterode
Lectotypei o/ ELEPHAS [= MAMMONTEUS] PRIMIQENIUS ^/z^to. Originals m Blumenlroch Coll Qpttin^en Mus.

Fig. 993. Lectotypes of Elephas [Mammgnteus] primigenius Blumenbach

After casts furnished by Director R. W. Hoffmann, Zoologische Institut der Universitat Gottingen

Siberian lectotype (I.M3) from the Bhimenbacli Collection of the Zoological Institute of the University, Gottingen (cast Amer. Mus. 26980).
(Left)
Described by Dietrich (letter, July 4, 1930) as follows: "Es handelt sicli bei dem Zahn aus Sibirien M3 sin., dist. niihcre Provenienz und Acquisition unbekannt."

Interpreted by Osborn a.s a portion of a third inferior molar of the left side, namely, I.M3, with 16 of the anterior ridge-plates preserved, of which at
least 1.5 are more or less fully worn; 8 posterior ridge-plates are missing; 7/2 ridge-plates in 100 mm. on the convex internal side. Height of 14th ridge-plate
(tallest) 112 mm. Drawn in crown and internal aspects.

(Right) Osterode lectotyiw (I.D])^) from the Blumenbach Collection of the Zoological Institute of the University, Gottingen (<'ast .\mcr. Mvis. 26981).
Determined by Dietrich (letter, .hily 4, 1930) as follows:"Der Zahn aus Osterode ist ein dritter oberer Milchmolar (m' [sin.]); er ist Kurz beschrieben in
.\nnalen der Physik, hersg. v. Gilbert 15, (4o|, 1813, p. 427/28. Er ist typisch fiir unserc spiitglacialen Mammute. Blumenbach diirfte ihn vor dem sibirischen
Zahn erworben haben, so da,ss dieserm'aus Osterode als Type specimen des Elephas primigenius Blumenbach, 1799, zu betrac-hten ist, wenn man nicht
vorzieht, das .\dams-Tilesius'.sche Skelet zum Typus der Spezies zu machen."

Interpreted by Osborn as belonging on the left side, namely, as an l.Dp'', with 12^3 ridgc-plates; 10-1 1 ridge-plates on the convex external side in 100 mm.

The above passage in which Rhinienhach, the pioneer of niaininaHan palaeontology in Europe, refers to the
discovery near "Bvng-Tonna" in the year 1695, is regarded as the tyi)e description of Elephas primigenius Blum.;
unfortunately the "Burg-Tonna" skeleton, now in the Gotha Museum, belongs to 'Elephas antiquus.' On the
advice of Dietrich (1930),- we may select as lectotypes of E. primigenius: (1) A grinder from Siberia in the Blumen-

"The priority of publication of Bhmienbach's spcu'ies Elephas primigenius (spring of 1799) is established over Cuvicr's Elephas mammonleus (August-
September, 1799). Sherborn writes (letter. May 16,1930): "It is rare that one finds .so jjcrfectly clear a statement in these old reviews. This comes from
Gottingische Anzeiger von gelclirten Saclien, 18 Dec. 1799, \i. 20.")7. 'Von <lcni RlunKwibarliisclicii Handbuch der Naturgeschichte ist schon vorige Ostern
. . .

die 6te .\ufiage auf 708 S. herausgekommen.'"

'See letter of February 8, 1930, from W. O. Dietrich, Geologisch-Palaontologisches Institut und Museum der Universitat Berlin: "Das zoologische Institut
der Universitat Gottingen besitzt nach Auskunft durch den Diri'ktor Prof. Dr. R. W. Hoffmann nur zwei Stiieke fossiler Elefanten aiis der Zeit Blumen-
bachs 1. Eincn Molar mit der Bezcichnung Elephas primigenius. Diluvium, Sibirien. Coll. Blum. 2. Eincn Molar mit der Bezeichnung Osterode (Harz)
1808."
 THE MAMMONTINiE: MAMMONTEUS 1123

bach Collection of the Gottingeii Museum, cast Amer. Mus. 26980; also (2) a molar from Osterode (Harz),
Germany, original in same Museum; cast Amer. Mus. 26981 (see Fig. 993 on opposite page.)

Blumenbach's type description is in a rare document; no figure accompanies it; the sixth edition (1799) of
his"Handbuch der Naturgeschichte," in which the above type description appears, attracted the attention of the
French naturalist Soulange Artaud, who visited Blumenbach and translated it under the title "Manuel d'Histoire
Naturelle," 1803. In the manuscript of the French volume, Blumenbach inserted the name and definition of
Elephas primigenius, consequently the species Elephas primigenius is of date 1799, 1803.

1799 Elephas mamnionteus Cuvier (MS., 1796, published August-September, 1799).

(Letter, Messieurs Eniile Picard and A. Lacroix, April 15, 1930): "Nous n'avons pas trouve trace d'unc telle
publication, niais il est evidentmemoire etait eerit avant le 21 Janvier 1796.
que le Ce memoire a etc lu a la classe
. . .

des sciences de I'Institut national, le pluviose, an IV (21 Janvier 1796). Cette lecture est mentionnee au proces-
verbal de la seance (Proccs-verbaux des seances de 1' Academic tenues depuis la fondation de I'Institut jusqu'au niois
d'aoilt 1835, t.I page 6). Reserve pour etre imprime, il a etc jniblic dans le tome II des Memoires de I'Institut
national, en PVuctidor, an VII (aout-scptcnibre 1799)."

Cited by Tilesius (1815, p. 470).

1806- Elephas primigenius Blum., Cuvier ("Oss. Fos.").

1834
1807 Elephas pmnxvus Blumenbach (in M. Adams; translation (1808) from the French by Sir Joseph Banks, Phil Mag.,
XXIX, London, p. 152; cited by Tilesius, 1815, p. 452.
1807 Elephas Mammouth Link, Beschr. Nat. Samml. Univ. Rostock (4) 1807,3 (fide Sherborn, 1928, Pt. XV, p. 3845).
1815 Elephas primordialis Blumenbach (in Tilesius, 1815, p. 470).

1820 Elephas jubatus Schlotheim.

1829 Elephas manimonteus Fischer de Waldheim.
1829 Elephas paniseus Fischer de Waldheim.
1829 Elephas periboletes Fischer de Waldheim.
1829 Elephas pygmaeus Fischer de Waldheim.
1829 Elephas campylotes Fischer de Waldheim.
1829 Elephas Kamenskii Fischer de Waldheim.
1830 Mammuthus Borealis Burnett.
1832 Mammul Sibiricum (in von Meyer, Palaeologica, p. 64).
1832 Elephas brachyratnphus Brandt ('Adams skeleton').
1832 [Elephas] homotaphnis Brandt.
1832 Elephas giganleus Brandt.
1832 Elephas commutatus Brandt.
1832 Elephas slenotoerhus Brandt.
1832 Elephas plalytaphrus Brandt.
1832 Elephas affinis Brandt.
1834 Elephas macrorynchus Morren.
1835 Elephas odontolyrannus Eichwald.
1837 DicYCLOTHERiuM E. Geoffroy St. Hilaire, Compte Rend. Acad. Sci., I\^, No. 4, pp. 119, 120, fig. 1.
1839- E. [Elephas] primigenius sibiricits de Blainville (Atlas, "Ost^ographie," PI. iii, 1845), also Deperet and Mayet,
1864 1923, pp. 183-201.
1839- E. [Elephas] primigenius germanicus de Blainville (Atlas, "Osteographie," PI. in, 1845).
1864
1848 Cheirolites von Meyer (in Bronn's Handb. einer Gesch. d. Natur., Ill, Index Pal., p. 286).
1850 Synodontherium Costa, "Palaeont. del Regno di Napoli," Atti Acad. Pontaniana, V, Pt. I, p. 271, Tav. ni, figs. 1-4.
1885 POLYDISKODONTEN Pohlig, p. 1027.
1888 PoLYDisKODON Pohlig, Nova Acta K. Leop.-Carol. Deutsch. Akad., LI II, p. 252.
1913 Elephas primigenius comune Issel, 1879 (in Zuffardi, 1913, p. 136)'.

'[Zuffardi (1913, p. 136) attributes this subspecies to Issel (1879) who, in describing two molars from Camporosso near Ventimiglia, in the region of S.
Andrea, Italy, states (p. 160) that these two teeth belong to the "comune Elephas primigenius," using comune in the adjectival sense. — Editor.]
1124 OSBORN: THE PROBOSCIDEA

Out of this host of names, barbaric, seiuibarbaric, non-technically Latinized or technically Latinized, applied
to the northern mammoth between the years 1696 and 1888, the species Elephas primigenius of Blumenbach alone
survives and is accepted in the scientific literature of the entire world.

As to the genus, nearly a century and a half of research since 1799, when Blumenbach assigned the name Elephas
primigenius, demonstrates that the woolly mammoth belongs not to Elephas, but to a genus of its own, distinct by
all the canons of nomenclature from the true modern Elephas. Technically the choice of a generic name rests

between the inappropriate Dicyclotherium von Meyer, 1848, Synodontherium Costa,

(k'offroy, 1837, Cheirolites

1850, and the first really appropriate generic name Polydiskodon Pohlig, 1885-1888. As Linnseus chose many of
his generic names, i.e., from those previously ill defined or not technically proposed, so in the present Memoir
we may go back to the eminently appropriate and distinctive generic term Mammonteum, 1788^ (changing the

orthography to Mammonteus).

Native Siberian Origin of the Word Mammut

Ides, 1706.— The following is cited (pp. 25, 26) from the book of travels "Three Years Travels from Moscow
over-land to China: thro' Great Ustiga, Siriania, Permia, Sibiria, Daour, Great Tartary &c. to Peking" of Eber-
hard Ysbrant Ides, who was sent ("by the most Serene and most Potent Soveraign Czar, and Great Prince,
Peter Alexewitz") on an embassy to the King of China in 1692, returning in the year 1695.

AinoDgst the Hills which are situate North-East of, and not far from hence [village of Makofskoi near the river Keta],

the Mammuts Tongues and Legs are found: as they are also particularly on the Shoars of the Rivers Jenize, Trugan, Motigamsea,
Lena, and near Jakutskoi, to as far as the Frozen Sea. In the Spring when the Ice of this River breaks, it is driven in such vast
quantities, and with such force by the high swollen Waters, that it frequently carries very high Banks before it, and breaks off

the tops of Hills, which falling down, discover these Animals whole, or their Teeth only, almost frozen to the Earth, which thaw
by degrees. had a Person with me to China, who annually went out in search of these Bones; he told me as a certain truth,
I

that he and Companions found a Head of one of these Animals, which was discovered by the fall of such a frozen piece of
his
Earth. As soon as he opened it he found the greatest part of the Flesh rotten, but it was not without difficulty that they broke
out his Teeth, which were placed before his Mouth as those of the Elephants are; they also took some Bones out of his head,
and afterwards came to his Fore-foot, which they cut off, and carried part of it to the City of Trugan, the Circumference of it be-
ing as largo as that of the wast of an ordinary Man. The Bones of the Head appeared somewhat red, as tho' they were
tinctured with Blood.

Concerning this .\nimal tlicrc arc \cry different reports. The Heathens of Jakuti, TungtiM, and Odiucki, say that they
continually, or at least by reason of the very hard I'Yosts, mostly live under ground, where they go backwards and forwards; to
confirm which they tell us, 'I'hat they ha\e often seen the Earth heaved up when one of these Beasts was on the March, and
after he was past the place sink in, and thereby make a deep Pit. They further beUeve, that if this Animal comes so near to
the surface of the frozen Earth as to smell or discern the Air, he immediately dies, which they say is the reason that several of
them are found dead, on the high Banks of the River, where they unawares came out of the (!round. This is the Opinion
of the Infidels concerning these Beasts, which are never seen.
Hut the old Sibfrian Huiisia7iti affirm that the Mammidh is very like the Elephant; with this only difference, that the
Teeth of the former are firmer, and not so straight as those of the latter. They also are of Opinion, that there were Elephants in

this Countrj- before the Deluge, when this Climate was warmer, and that their drowned bodies floating on the surface of the
Water of that Flood, were and forced into Subterranean Cavities: But that after this Noachian. Deluge, the Air
at last wa.sli'd

which was before warm was changed to cold, and that these bones have lain frozen in the Earth ever since, and so arc jireserved
from iMitrefaction till Ihey thaw, and come to light; which is no very unreasonable conjecture. Tho' it is not absolutely
necessary that this Climate shoulil have been warmer before the Flood, since the ( 'arkasses of the drowned Elcphantx were
very likely to float from other places several hundred Miles distant, to this Country, in the great Deluge which covered the
surface of the whole Earth. Some of these Teeth, which doubtless have lain the whole Sununer on the Shoar, are intirely
black and broken, and can never be restored to their former condition; but those which are found in good case, arc as good as

'[Sec footnote on page 1117 above rcgarJing the doubtful validity of the genu.s Mammonleus. — Editor].
 — —

THE MAMMONTIN^: MAMMONTEUS 1125

Ivory, and are accordingly transported to all parts of Muscovy, when they arc used to make Combs, and all other such like things,
instead of Ivory. The above mentioned Person also told me, that he once found two Teeth in one Head that weighed above 12
Russian Pounds, which amount to 400 German Pounds; So that these Animals must of necessity be very large, tho' a great
many lesser Teeth are found. By all that 1 could gather from the Heathens, there's no Person ever saw one of these Beasts
alive, or can give any account of it's shape; so that the most that is said on this subject arises from bare conjecture only.

—
HowoRTH, 1882.1, PP. 30-32. Howorth has given an excellent account of the name "Mammoth" in his article

of 1882 in "The Field Naturalist," page 30, from which we quote the following:

In the year 1666, a learned Dutchman, named Cornelius Witzen, who became burgomaster of Amsterdam, visited Moscow.
From the materials he there collected, and from various other sources, he compiled a famous work which he afterwards published
at Amsterdam, entitled Noord en Oost Tatarye [Witsen, Nicolaes, 1692.1]. ... It has never been translated into any other
western language, and remains still buried to a large part of the scientific world in the original Dutch.
In this work the name Mammoth appears for the first time, and is there written Mammout. Witzen describes how numbers
of elephants' teeth are found on the banks of the Siberian rivers, and says 'by the Inlanders,' i. e. the Russian settlers in
Siberia, 'these teeth are called Mammouttekoos, while the animal itself is called Mammout.' {Op. cit. ed. 1785, 742). In the
Grammatica Russica of Ludolf, jjublished in 1696, p. 96, these bones are called Mammotovoi Kost. Among the Russians the
name is invariably written Mamont — Bull. St. Pet. Acad. x. 258. The insertion of the n in this form is no doubt a corruption,
although an old corruption, since Captain J. B. Miiller, in 1716, says 'the inhabitants (i. e. the Russians) call the beast Mamant.'
Schiefier explains it as due to a confusion made by the peasants between the name Mammoth and that of Saint Mamas, called
Mamant by the Russians, to whom the second of September is dedicated in the Russian calendar {id. note 2). Pallas derived
Mammoth from the word Mama, which he says means earth in the Tartar tongue, but as Baer says, in the great Polyglot, edited
by Pallas himself, in which the various dialects of Tartar are illustrated, no such name as Mama occurs as a synonym for earth.

Baer himself suggests that the name is derived from ma, which means the earth in various Fin dialects, while mut or muit is the
name by which the mole is still known to the Esthonians, whence he explains Mammoth as meaning the earth-mole (Bull. —
St. Pet. Acad. x. 258). Not only is this an inconsequential etymology, since all moles live under ground, but the notion of
finding an explanation of the name among a people so far removed from Siberia as the Esthonians is quite fanciful. Klaproth
was told by the Buriat and Mongol Lamas whom he consulted in 1806, that the name was of Tibetan origin. This again is
a most unlikely quarter to go to for our etymology. A much more reasonable explanation of the name was given long ago by
the Swedish exile to Siberia, Strahlenberg, who wrote his well-known description of the north and eastern parts of Europe at
the beginning of the last century. In the 13th chapter of this work, under the heading Mamatova Kost, we read 'As to the
name it doubtless had from the Hebrew and Arabick; this word denoting Behemot, of which Job speaks (in the xi.
its origin

chapter), and which the Arabs pronounce Mehemot**** This is certain, that they ((. e. the Arabs) brought the word into Great
Tartary for the Ostiacks near the river Oby call the Mammuth Khosar, and the Tartars call it Khir; and although the Arabian
;

name of an elephant is Fyhl, yet if very large they add the adjective Mehemodi to it; and these Arabs coming into Tartary, and
finding there the relicks of some monstrous great beasts, not certain of what kind they might be, they called these teeth Me-
hemot, which afterwards became a Proper Name among the Tartars, and by the Russians is corruptly pronounced Mammoth.***
The Russian Mammoth came from the word Behemot; in which opinion I am confirmed by the testimony of an
certainly
ancient Russian Priest, Gregory by name, Father-Confessor to Princess Sophia, who was many years an exile in Siberia, from
whom I was told, that formerly the name for these bones in Siberia was not Mammoth, but Memolh, and that the Russian
dialect had made that alteration.' ^{Strahlenberg Eng. ed. 403). This view is curiously confirmed by the fact that Father
Avril, a Jesuit, who went overland to China in 1685, and who is quoted by Witzen in the notices he gives of the Mammoth,
never calls it Mammoth, but always Behemot.— (^mPs Travels, Eng. ed., 175, 177). Witzen himself also says that these teeth
belong to the beast Behemoth, called Mammout, otherwise Mammona by the Russians.— {Op. cit., 742). The Turkish dialects
habitually interchange the letters B and M, which are, in fact, used indifferently by them, so that the change from Behemoth to
Mehemoth is and natural, and we can hardly doubt that Mammoth is in fact a mere form of Behemoth. At
perfectly regular
first it looks strange that the Arabs should have given a name to this beast which has become current all over the world; but

this is easily explained when we remember the immense enterprise and energy of the Arabs in the ninth and two succeeding
centuries, when, as we know, their traders and emissaries frequented the borderlands of Siberia, and probably first initiated the
trade in fossil ivory in the west. Father Avril's narrative shows that this trade still survived in his day, and he tells us the
. . .

Persians and Turks put a great value on elephants' teeth from Siberia, and preferred a scimitar or a dagger haft made of this
precious ivory before a handle of massy gold or silver. There is, therefore, every reason to believe that the name Mammoth,
as well as Mammoth ivory itself, were first introduced to the notice of the western world as an article of commerce by the Arabs,
who were familiar with it probably as early as the ninth century.
1126 OSBORN: THE PROBOSCIDEA

I'ig. 91)4. Rostoratiim of the migrating \\<^()lly mammoth (Elephas [Mammonteus] primigenius Blum.) a.s it appeared on the river .Somme, northen
France. Details after the Upper PaliPolithic etching.-* and paintings of Magdalenian time, especially in the caverns of Font-de-Gaume and Conibarelles
Painted in 1919 by Charles R. Knight, under the direction of Henry Fairfield Osborn, for the Hall of the Age of Man, American Museum of Natural History.

II. CHARACTERS OF THE SUBFAMILY MAMMONTIN^^, INCLUDING THE GENUS

MAMMONTEUS
Superfamily: ELEPHANTOIDEA Osborn, 1921
Family: ELEPHANTIDiE Gray, 1821
Subfamily: Mammontin^ Osborn, 1921 (as defined above in Chapter XVI, Arclndiskodon)

Genus: MAMMONTEUS Camper, 1788'

OriKJnal reference: ('ami)cr, 1788, pp. 2.51,259; Osborn, 1924.(3:«, p. 2.

Pleistoeene of I'Ain)i)e, northern Asiii, and North America.

Svn. (partial list): Dici/clvlheriuni Geoffroy, 1837; CheiroliUs \-on Meyer, 1848; Sijuoihint.hcriiiiii ('os(a, 1850; I'ob/dinko-
donlcn Pohlis, 1885, PohjdiHkodon Pohlig, 1888.

Mammoxteum Camper, ". 1788, p.

—
os humantini i)etrifactvnn, aut fossilo, etiaiiisi Mannnonteoruin,
251. . .

j!]l('l)haiitoruni, Adserere ex eodeni principio audeo Mauimontcuni animal extinctum non mudo e.sse, «ed
. . .

nullam omnino haljiiisse cum Elephanto similitudinem!"

Generic Characters in Cranium and Teeth (Osborn, 1928). —The Mammonteum of Camper,
1788 [Mammonteus of Osborn, 1924], inchides Elephas primigenius Blum., also Elephas americanus
DeKay, Elephas primigenius astensis Deperet and Mayet, Elephas primigenius fraasi Dietrich, and
Mammonteus primigenius compressus 0,sborn.- (1) Cranium related to that of Archidiskodon and of
Parelephas, but extremely acrocephalic, hypsicephalic, bathycephalic. Frontal.s concave, occipital
cre.st greatly elevated, occii)ut .slightly convex. (2) Molans, in Upi)er Pliocene to Ujiper Pleistocene stages,
with relatively numerous ridge-plates; Upper Pliocene stag(>, M. primigenius astensis, 3 jfr^l; M
typical Upper Pleistocene stage, Mammonteus primigenius, 3 M; final progres.sive stage, M. primi- M
genius compressus, 3 (^f^y. M
Ridge-plates compressed in typical superior molars to 10-11-12 in 100

'[Sec footnote on page 1117 above for remarks on the doubtful validity of the genus Manwionteua. Editor.] —
'[To these should be added Mammonteus primigenius alaskcnsis described below (pp. ll.')9-1161), this chapter. — Editor.]
 : :

THE MAMMONTIN^: MAMMONTEUS 1127

mm., molars 13 in 100 mm.; in progressive inferior molars 8-9-10 in 100

in highly progressive superior
mm. Molar crowns broad, M'' extremely short with enamel ridge-plates of minimum thickness, ridge-plates
more or less crimped or sinuous. (3) Manus pentadactyl with five digital nails (fide Herz, 1902), manus
and pes tetradactyl, not pentadactyl (fide Salensky, 1904, p. 86— see citation from Salensky' on page 1129
below); four digits {fide Pfizenmayer, 1926, p. 239), total phalanges in each foot reduced to nine in
number, e.g., " n m it v d^IT'' (4) Dietrich (1912) records five digits in the manus and pes of the large
and more primitive Upper(?) Pleistocene M. primigemus fraasi. (5) Tail abbreviate, caudals 21. (6)
Habitat, northern tundras and steppes, grassy meadows; summer diet chiefly of grasses.

All authors have included E. primigenius within the genus Elephas, the genotypic species of which is Elephas
indicus Linnaeus, 1754 ( = Elephas maximus Linnseus, 1758) . It has now been demonstrated that the true northern
mammoth {Elephas primigenius) is one of the final members of a long series of species and of ascending mutations
extending back through the entire period of Pleistocene time and first recognized in the Upper Pliocene of Italy
as Elephas primigenius mut. astensis, described by Deperet and Mayet in 1923. Throughout this long geologic
period these ascending mutations and species exhibit fundamental dental or cranial characters and proportions
which clearly distinguish them from Elephas indicus ;
consequently we have to do with a distinct generic phylum,
comparable to the phyla of Archidiskodon and of Parelephas. For this 'northern' or 'woolly' mammoth the
appropriate name Mammonteum Camper { = Mammonteus) was revived by the present author in 1924 (Osborn,
1924.633, p. 2).

1. EXTERNAL CHARACTERS AND FEEDING HABITS

The true Mammoth is the only extinct proboscidean of which the characters of the soft parts and of the
hairy and woolly covering, as well as the nature of the food both of the Mammoth and Mastodon, are fully known.
The student is referred to the reports of Herz (1902) on the mammoth discovered on the Beresowka River; also
to Salensky' (1903) who described in the greatest detail the external characters of the same individual. The
most recent descriptions, however, are those Quackenbush (1909) noting the discovery in 1907 at "Historic
of
Bluff," Eschscholtz Bay, Alaska, of part of a skeleton of the Mammoth, together with some skin and hair (Amer.

—
Mus. 13749 paratype of M. primigenius compressus) and of Felix (1912) in his "Das Mammuth von Borna,"
,

from which we may cite by translation

—
Summer Food of Grasses (Felix, 1912, p. 11). On uncovering the skull a portion of the animal's food was
found in the form of a wad lying between the upper and lower teeth. Its death, therefore, must have been so
sudden that it did not have time to swallow this food. In addition, the entire stomach was discovered containing
about 12 kilograms (24 lbs.) of undigested food. On examination this material proved to contain a flora of no
great variety, but of exceeding interest, because it consisted of plants that are still native to the place, i.e., Bere-
sowka River, northern Siberia. They are almost exclusively grasses. The needles of conifers occur very rarely.
Here is the list

1. Alopecurus alpinus Sm. = alpine foxtail grass].

[ 6. Care.x glareosa Wg. = clustered
[ sedge].
2. Hordeum jubatum L. = squirrel-tail grass].
[ 7. Carex incurva Lightf. [
= curved sedge].
3. Agrostis borealis Hartm. = red bent-grass]. [ 8. Thymus serpyllum L. [
= wild thyme].
4. Atropis distans Griseb. = sweet-grass].
[ 9. O.xytropis campestris DC. = field oxytrope].
[

5. Beckmannia crucaeformis Host. = slough-grass]. [ 10. Papaver alpinum L. [

= alpine poppy].
11. Ranunculus acer L. var. borealis [
= common buttercup].

'[See Zalensky, Vladimir Vladimirovieh, in Bibliography of Volume I of the present Memoir. — Editor.]
 :

128 OSBORN: THE PROBOSCIDEA

{Op. cil., p. 12) : All these kinds of plants are found in the same locality at the present day and form a char-
acteristic meadow flora. With the exception of Alopecurus alpiims Sni. and Papaver alpinum L. which are also
found in the tundra, there were no typical tundra plants. From other discoverieswe know that the mammoth
fed also on: (1) Betula nana L. [
= dwarfed birch]; (2) Salix polaris Wahlnbrg. = arctic willow];
[ (3) Cladonia
rangiferina Hoffm. = reindeer
[ moss].

In addition to these the manmioth ate (probably in winter) the bark and twigs of conifers, chiefly larches.

Proportions. — (P^elix, 1913, pp. 13, 14, translation) : (1) This discovery not only gave us complete knowledge
of the skeleton of E. primigenius but showed conclusively the proper position of the tusk in the jaw, and its inclina-

CoMI'ArUSON OK THE TiP OF THE TrUNK OF THE MaMMOTH (MaMMONTEUS PRIMIGENIUs) WITH THAT OF THE INDIAN
ELEPHANT (ElEPHAS INDICUs) AND OF THE AFRICAN ELEPHANT (LoXODONTA AFRICANA)
After Flcrov, 1931, Figs. 1, 2, and 3
Fig. 995. The following is cited from Doctor Flerov's interesting description (1931, pp. 863 and 869) of the
MaMMONTEUS PKIMIGENIUS first discovery of the tip of a mammoth's trunk
Rkconstkuction
In 1924 a well preserved mammoth trunk was found by an unknown Tungus in the everfrozcn .soil on the banks of
the Bolshaya Baranikha River in the Kolyma district. This region is very distant from any inhabited place, devoid of
regular meanscommunication and peopled with half-wild tribes. Five years elapsed before the news of a discovery,
of
so exceptionally interesting for the Science, reachedby a good chance some of our geologists working in the Kolyma
district. The trunk in this time passed from person to person, its tip was cut off, then dried and the rest was thrown
away and lost. Only in 1929 the dried tip of the trunk was handed over by Mrs. Kondratiev, a resident of Sredne-
Kolymsk, a little town on the Kolyma River, to Mr. K. J. Pjatovsky, a geologist, who sent it to the Zoological Museum
of the Academy of Sciences through Mr. M. J. Tkatchenko, an assistant of the Yakutsk Museum.

Table of Measurements
(Cf. op. cit., p. 869)

ElEI'HAS I.NDICI s mm ram

Length of the tij) of trunk on the external side 280 Length on the exterior border
of the 'lip' 36
Maximal width 114 Width of the right nasal opening 23
U idtli above the base of the 'lip' 74 Width of the left nasal opening 17
Width of the 'lip' 71 Thickness of the wall dividing both nasal op('nings I'l

l.ciigtii of the finger-like |)rocess 95 Thickness of the skin on the exterior siile 2
\\ idtli of the finger-like process on tlic base 57 Thickness of the skin on the interior .side 0.5

IjOXODONTA ai-ukana

tion. (2) As to the animal's outward appearance, it involved a number of corrections in even the best reconstruc-
tions that had been made. The proportion
(3) of the length of skull and trunk in the mammoth is quite different
from that of existing elephants. The length of the mammoth's skull is more than half that of the trunk while the
elephant's skull [E. indicm] is always less than half the trunk length. The mammoth's head, therefore, was larger

in ])rop()rtion to the body than that of the recent elephants, and in con.sequence the tusks could attain enormous
proportions. The largest of the tusks in the St. Petersburg Museum mea.sures no less than 4, 17 m. (13'8")
and ill the Franzens-Museum of Briinn there is a tusk that actually exceeds 5 m. (16'o") in length!
 THE MAMMONTIN^.: MAMMONTEUS 1129

The tnink of the Beresowka mammoth was entirely missing, but probably differed little from that of recent
elephants. The many representations of the mammoth found in the Palaeolithic caves of France all show a strong-
ly developed trunk. The ear was somewhat smaller than that of the Indian elephant. Its length was 38 cm.
(15"), its breadth, 17 cm. (6%"). The ears, as well as the whole body, were covered with a thick coat consisting of
short wool and longer hair. The massive body, rather short in proportion to its height, is joined to the mighty
head by a short neck which seems still shorter on account of the great muscles.

The tail — almost unknown until this discovery — is (1) conical in form, sharply pointed at the end and about
36 cm. (14") broad at the root. It ended in a bunch of bristles. (2) Salensky' gives the length — measured from
the back— as 60 cm. Pfizenmayer writes [1905, pp. 524, 525] 'The
{2S'A"). : tail of this specimen — measured from

the under side — 35 cm. long, decidedly shorter than that of living elephants.
is The number of [caudal] vertebrae

is only 21.' (3) It would appear from make it much too long. The difference in
this that the older reconstructions

the measurement of Salensky and of Pfizenmayer can hardly be attributed to the mode of measurement. Proba-
bly Salensky included the bunch of bristles at the end in his length, while Pfizenmayer did not.

The skill was extraordinarily thick and underneath it was a layer of fat from 1 to 9 cm. (%-3'/i in.) deep. The
whole body was thickly covered with hair, even the legs down to the horny ends of the toes. The mammoth
was thus particularly well fitted to withstand cold, while its outward appearance was quite different from that of
its living relatives. The covering of the body consisted of three elements: (1) Fine, soft, woolly hair about an
inch (20-25 mm.) long, in color varying from faded blond to yellow brown, and covering the entire body; (2)

coarser and longer hair up to 20 inches (50 cm.) in length, of a dark rust colored brown, covering the entire neck

and trunk (of the body) perhaps forming a fringe of hair still heavier and thicker from the cheeks along the shoul-

ders and sides to the rump, similar to that of the yak. (3) "Bristles" —so called on account of their stiff ness —from
8 to 14 inches (20-35 cm.) in length, much darker than the other kinds of hair, found only in the "brush" at the
end of the tail.

2. SKELETAL CHARACTERS OF MAMMONTEUS PRIMIGENIUS

These authors do not give a complete description of the osteological characters of the Beresowka mammoth.
The most important note is that of Salensky' (1903, p. 86) : "An osteological study of the Beresowka mammoth
led to the unexpected conclusion that considerable differences exist between the Elephas primigenius and the
recent elephants [E. indicus], notably in the foot, which is tetradactyl and not pentadactyl. These differences
absolutely exclude the possibility of the direct descent of the elephant from the mammoth."

One of the fullest comparative discussions of the mammoth skeleton is that of Dietrich (1912), in which the

vertebral formula, covering our whole knowledge, is summarized below.

Abbreviated Vertebral Column. — Since the abbreviated backbone shares the extreme fore-and-aft com-
pression of the cranium, E. primigenius has relatively the shortest vertebral column of any of the Proboscidea.

The reader is here referred to pages 930, 931, Chapter XV for a full treatment of vertebral distinctions in the genera
of the Proboscidea. This abbreviation is indicated, however, in the shortening of the vertebral centra as well as in

the number of dorso-lumbar vertebra? (dorso-lumbars 24-23 in Elephas indicus, 23 in Loxodonta africana, 22-24 in

'See footnote on page 1 127 above.

 Skeletons of Mammonteus primigenitjs Mounted in the Museums of Russia, Germany, and Belgium
Moiinted skeletons of Kolyma-Bcrpsowka, of Borna, of Steinheim, ami of Licrrc. Reduced to a unifomi scale of about one-fiftieth natural size

Fig. 996. Skeleton of Mammonteus primigenius from the Kolyma-

Beresowka River, .Siberia, as mounted in the Museum at Leningrad, Russia. Fig. 998. Skeleton of Mo )HHjo?((ei/.spnmifff/iiH.s from Borna, Germany, as
After Abel, 192."), p. .'iti, fig. 36, as reconstructed by W. Salensky |see footnote, mounted in the Museum fiir Volkerkunde, Leipzig (Gra-ssi-Museum). After
p. 1127 above]. Abel, 192"), p. 279, fig. 204 (.see Felix, 1912, Taf. viii, for original).

Fig. 997. Skeleton of Mammonteus primigenius ( = Elephas

Mammonteus] primigenius fraasi Dietrich, 1912) from Steinheim
[.

on the Murr, Wiirttemberg, Germany, as mounted in the Stuttgarter

Naturalienkabiiiett. .4fter Abel, 1925, p. 57, fig. 37, as reconstructed
by E. Frmis and W. O. Dietrich. About one-fiftieth natural size. Fig. 999. Mammouth du Musee de Hruxelles decouvert a Lierre (Province d'Anvers)
en 1860. After Dupont "L'Homme pendant les Ages de La Pierre dans les Environs
de Dinant-sur-Meuse," Brussels, 1873, PI. n. One-fiftieth natural size.

It will be observed that the acrocephalic, hypsieephalic, and bathycephalic proportions of the cranium and the harmonic abbreviation of the vertebral
column are manifest in each of these reconstructions. The sharp notch behind the peaked skull in the Combarelles restoration (Fig. 1000) is a feature of every
drawing of Palaeolithic age. Compare figure 994.
'.\dam.s Skeleto.n',' 1807 (see Fia. 1014). — The first complete remains, known as the 'Adams skeleton' as mounted (Fig. 1014) in the Zoological Museum
of the .\cademy of Sciences of the U. S. S. R., Leningrad, were discovered in 1799 on the banks of the Lena River at the threshold of the Polar Sea (cf. Lang,
1925, p. 28):
"Imbedded had been for thou.sands of years, its meat was still in such condition as to be eagerly devoured by polar bears, wolves, and other
in ice, as it
carnivores attracted from great distances. As time went on every warm season bared more of the body; only the natives contested the booty by securing some
of the meat for thcnr dog.s through the following years of exposure. It was then that the intrepid explorer and botanist Adams happened to arrive in the
neighborhood and, hearing of the famed monster, lost no time in reaching it. Most of the soft parts were gone, one limb had been carried away, and a native
had .sawed off both tasks and .sold them for about fifty rubles. Through -\dams' energy and foresight practically all remaining bones were collected. He also
took to Petrograd a piece of the hide with the hair in place. It was from the still frozen side upon which the mammoth lay, and so heavy as to tax the strength
of ten men to <lrag it along the shore. \ large amount of loose, coarse hair, evidently trampled into the snow by feasting polar bears, was long enough to be
considered as having formed a mane." This '.Adams skeleton' as described and figured by Tilesius in 1815 was further described by Cuvier as cited below
(see Fig. 1014). [Made by Brandt (1832, pp. XI, XII) the type of Elephas brachyrainphus.— Editor.]

1130
 THE MAMMONTIN.E: MAMMONTEUS 1131

Mammonteus primigenius, and 23 in Parelephan J€JJersonii)^Q,s shown in l^he following comparative table —see also
Chapters XV and XIX:

LoxodonUi africana Elephas indicus Parelephafs Mainmonleus primigenius

Flower, "Jumbo" Flower? • li.rType rj'i;, Falconer and Feli.x, 'Adams Beresovvka
1885 Amer. Mus. 1885 Amer. Mus. Araer. Mus. 1912 Mammoth, Mammoth
Dept. Mam. 9950' ,S- -'^^14559 (M. p. Tilesius Pfizenmayer
3283 compressus,] .
^^
1815 in Dietrich
^^y
1912

Cervicals. . . i i 7
Dorsals 19 20 19-20 19 18-19 17-19? IS
Pairs of ribs. 19 18-19 ,19, ,. 17^19?
Lumbars. . . 4 3 5-3 4 4 3 5 4? 5
Saerals 5 4 4 5 4-3 4 4
Claudals. . . . 24 + 21 24-30 + 12+ 21 21 (Sa- 8+ - 21
lensky)'

Fig. lOOV.' The above outlines of the woolly mammotli are from the
Grotto of Combarelles, in which representations of the mammoth are next in
order of frequency to those of the horse. Compare Capitan, Breuil, and
Peyrony, 1924, pp. 136, 137, figs. 120, 121. MacCurdy (1924.1, p. 275) records
twenty or more caverns in France, Spain, and Germany in which drawings of
Fig. 1000. Restoration of the woolly mammoth sketched on the wall of the the mammoth occur, mostly in outline, while the famous painted murals of
eavern of Les Combarclles aux Eyzies (Dordogne), Franco. After Capitan, Font-de-Gaimie are of a dark brownish tint. The mammoth was the only
Breuil, and Peyrony, 1924, p. 137, fig. 121. Observe the very small feet, the member of the elephant family in northern Europe during the period of the
short tail, the drooping hind quarters, the upcurved backbone, and the neck Fourth Glaciation or close of the Stone Age. In Spain, however, the straight-
sharply incurved behind tlie summit of the occiput, characters whi(,h agree tusked Elephas [Hesperoloxodon] dntiquus lingered to the very end of the Stone
closely with those of tlie skeletons of Steinheira (Fig. 997) and of Borna (Fig. Age and its outline in the Cavern of Pindal, northern Spain, is shown in figure
998). 1047 of the present Memoir (Chap. XIX, the Loxodontinse).

Pfizenmayer, 1926, p. 239. While the skeleton of manus and pes in elephants has five digits, that of the

mammoth has only four in both fore- and hindfeet [cf. Herz, 1920, five digits' in manus]. The first digit is com-
pletely lacking, and the other digits even in older animals show three phalanges only in the third digit, while

digits II, IV and V have but two phalanges in fore- and hindfeet.

All four extremities (feet) of the Beresowka mammoth were complete, and the left hindfoot from the knee
joint down was preserved in alcohol with the skin and soft parts. This formula of four digits for the Siberian mam-
moth was demonstrated when three extremities of the Beresowka mammoth were macerated by Professor Salen-
sky,i director of the Museum, and myself as his assistant. There can be no doubt about this fact, as we also
proved it to be the case in three older fore- and hindfeet which had been preserved with the soft parts intact.
They belonged to the Maydell mammoth (1869) in the St. Petersburg Museum collection, and showed the same
feature. Also the perfectly preserved fore- and hindfeet of the mammoth of Sanga-jurach agreed with the former

'See footnote on page 1 127 above.

 :

1132 OSBORN: THE PROBOSCIDEA

discoveries in this respect. The phalangeal formula of the adult Siberian mammoth for both manus and pes is
2 PhuluOKca.
therefore II III IV V DiiiU

Restoration. — It will be observed in the Palaeolithic restoration (Fig. 1002) that the apex of the cranium
is extremely short and pointed (cyrtocephalic, acrocephalic), while the facial profile of the cranium (Fig. 1000)
is rounded, a feature probably due to the storage of fat during the winter beneath the woolly dome. All these
drawings indicate also a sharp depression between the head and the back, due to the extremely sharp elevation of
the cranium; they also indicate the rapid downward slope (usually exaggerated) of the back toward the pelvis.
Consequently the drawings fully confirm the evidence afforded by the fossilized skeletons, that the mammoth
was wholly different in bodily contour from either of the existing elephants, Elephas indicus or Loxodonta africana.

Another characteristic is the rapidly sloping hind quarters, as observed in all the numerous drawings and
paintings of Mammonleus primigenius by the Cro-Magnon artists of Upper Palaeolithic age, the most striking and

Fig. 1002. Charging mammoth incised on a tusk of Elephas primigenius discovered in 1865 in the Magdalenian cviltural level of the great rock shelter of
La Madeleine (Dordogne), France, and first described by Lartet in the Comptes Rendus des S6anees do I'Academie des Sciences, Vol. LXI, 1865, pp. 309-311,
subsequently figured by Lartet and Christy in the Rcliquise Aquitaniese, 1875, B. PI. xxviii, and amplified by them in the Descriptions of the Plates, p. 168, as
follows

"It is a thin oblong piece, convex from side to side with the roundness of the tusk, and somewhat concave in the longitudinal direction, owing to its
curvature. The outer surface presents what at first appears to be a medley of faintly scratched lines; but, on closer and more careful inspection, they re-
solve them.selves into a characteristic outline of a
hairy Elephant, with some of the lines doubled and
redoubled apparently by the old artist's repeated
attempts to sketch out the main features of his sub-
ject. The lofty skull and hollow forehead are recog-
nizable as striking features, characteristic of the
Siberian Mammoth at St. Petersburg (Footnote:
'See Le Hon's "L'Homme fos.sile," 1867, p. 70,
woodcut. 'I, of the skull of the Mammoth from Ilford,
Mu.seum (Footnote:
Essex, preserved in the British
'Much modern information on the features, distri-
bution, and general natural history of the Mam-
moth, together with references to other authors, will
be found in the Memoirs by Mr. Henry Woodward,
F.R.S., of the British Mu.seum, in the Geological
Magazine, 1864, vol. i. p. 241, and woodcut; 1868,
vol. V. p. 540, plates 22 and 23; 1869, vol. vi. p. 58;
and 1871, vol. viii. p. 193, pi. 4.'], and of the Belgian
Mammoth at Bru.ssels (Footnote: 'See Dr. E. Du-
pont's "L'Homme pendant les ages de la Pierre
dans les Environs de Dinant-sur-Meuse" (pi. 2),
8vo, 2nd edit. 1872.'].The small eye and long trunk
of the Elephant, and the great curved tusks and
shaggy hair peculiar to the Mammoth, are easily
recognized. The upper and more convex sketch-
lines of the back agree with the high withers of the
Mammoth; and the lower and sloping dorsal lines
probably had reference, in the draughtsman's mind,
to some siM'cial attitude of the animal, with which also the outstretched portion of the hind leg, and the elevated tail, may be as.sociated.
"Mr. H. Woodward, F.R.S., F.G.S., has kindly communicated the interesting suggestion that the attitude of the animal, together with the vertical position
of the trunk, would well accord with the idea of one of a herd of Elephants coming down by moonlight to drink, and that the confusing doiibk lines might then be
explained as an attempt, on the part of the artist, to represent the rest of the herd. In running, or when alarmed, the trunk of the Elephant is always raised.
And he adds that there can be little or no doubt that the sketch, rude as it is, was the result of a life-sludu of the animal, and is consequently of the highest
importance as attesting the actual presence of the living Mammoth in France when the Caves of Perigord were occupied by Man."
In the present figure the other outlines, i)robably designed to indicate a charging herd of mammoths, are represented by dotted lines only (cf. Osborn's
"Men of the Old Stone .\ge," This classic engraving, now preserved in the Mu.seum d'Hi.stoire Naturcllc, Paris, is one of the most realistic
p. 384, and Fig. 199):
pieces of Pala-olithic engraving which has ever been found; observe especially the outline of the ear, the elevation of the highly peaked (acroi'epahlic) head, also
the remarkably lifelike action of the limbs and body. There are indications that the artist used this relatively small piece of ivory for the n-presentation of
three mammoths the tusks and trunks of two other elephants appear in the distance.
;
 THE MAMMONTIN^: MAMMONTEUS 1133

probably the most accurate representation of the mammoth being the famous charging mammoth as engraved on

an ivory tusk (Fig. 1002), the original of which is now in the Museum d'Histoire Naturelle, Paris.

—
Geogeaphic Distribution. (1) The geographic distribution of Mammonteus primigenius in the United
States, in the recent valuable memoirs by Hay (1923, 1924), is unfortunately marred by the fact that many of the
recorded specimens attributed by him to "Elephas primigenius" really belong to the animal described in the
present Memoir as Parelephas jeffersonii. (2) The same confusion of the remains of Parelephas with those of
Mammonteus doubtless applies to the plotted distribution' by Osborn (Fig. 1005) in the Northern Hemisphere,
designed to show the former distribution of the true mammoth; the European plotting (solid black) is largely
after Boule and certainly includes both the 'Elephas primigenius' and the 'Elephas trogontherii' range. (3) The

Asiatic plotting (deep bars) is largely conjectural, and the same doubt exists as to the distinction between the
'E. primigenius' and the 'E. trogontherii.' The American plotting (solid black) is also after Hay and, as above
noted, doubtless includes the range of species of both Mammonteus and Parelephas. (4) The distribution of

MAPS AND THEIR EXPLANATIONS. MAPS AND THEIR EXPLANATIONS.

Mai- 12.
Map 11.

Wiiconsin lewin lllinoiao

[K3
Kantan
^.
Pr&wisconsin DriftUss

Geographic Distribution op Mammonteus and Parelephas. After Hat

Fig. 1003. Geographic distribution of Mammonteus primigenius in the central (left) and eastern (right) region.s of North America, after plotting by
O. P. Hay: Hay, 1924, map 12, pp. 340, 341; (right) 1923, map 11, pp. 428, 429. A point of great significance is this indubitable occurrence of Mam-
(Left)
monteus primigenius in the Wisconsin and pre-Wisconsin 'drift' (IV Glacial), in the areas of the 'middle drift' (Illinoian, III Glacial), and lowan?, IV
Glacial), and in the 'old drift' (Kansan, II Glacial). See Plate viii opposite page 348 of Volume I of the present Memoir.
This plotting by Hay (1924) also includes finds of Parelephas jeffersonii and of P. progressus (listed under the names 'Elephas boreus,' 1924, pp. 47-.'J6, and
'Elephas columbi,' 1924, pp. 57-84). .^s shown in the present Memoir, the crania and grinding teeth of the various species of Mammonteus and Parelephas,

after years of confusionby all paleontologists including the pre.sent author, are now readily distinguishable.

'[The plotted distribution (Fig. lOOo) is entirely out of date (1938); it is now superseded by figure 795 in Chapter XV above, in which the range of the
—
southern 'trogontherians' and northern 'mammoths' is theoretically plotted. Editor.]
11.34 OSBORN: THE PROBOSCIDEA

Pleistocene life in the very important geographic area of Alaska, on the confines of North America and Asia, has
been treated in recent years by Maddren (1905), by Gilmore (1908), by Quackenbush (1909), by Osborn (1910.
346), and l)y Frick (1930). A synthesis of their observations is recorded in the accompanying map of 1930 (Fig.

1004.)

Alaska, broadly connected with, the Asiatic mainland, was a peninsula for free migration between Eurasia
and America; the fossil fauna thus far discovered, chiefly of Pleistocene times, is of boreal or circumpolar char-

acter but includes mammals of the north temperate latitude, such as the horse, the bison, a single foot bone of

Fig. 1001. Fossil mammalian fauna o{ Alaska-Yukon and location of principal discoveries to the year 1929. Modified after Qi.ackenbusli (1909, see full

bibliography, pp. 128 130, and map), including discoveries and records of Maddren (1905) and identifications of Gerrit S. Miller, Gidley, Hay, Lambe, also
of Frick (1930).
As shown by the symbols, Mamnionteus priinigenius was extremely abundant; M. prim. cnnipresKus, represented by the single paratype specimen from
lOsclischoltz Bay; Equux .surprisingly abundant, there being recorded thirty-three loi-alities, including (Wainwright) the north coast; Ma.ftodon americanus
very rare,' being recorded only in the upper Canadian Yukon, on the Indian River, Long. 140, I.al. 04; and six records of Ow'bo.s. In the very rich explorations,
made through the i-ooperation of the Fairbanks Exploration Company, Frick, Bunnell, and Kaisen (1929) discovered the two new types Felin alrox alitskensis
tmAMnocijondiruH alaskeyisis, resembling like forms from the Raneho La Brea, .southern California, intermingled with very numerous remains of Hifion cras-
simrnis, Equus, Mamnionteus,- etc.

'[Compare Mastodon americanus alaskensis Frick, 1933, on page 176 of Volume I of the present Memoir. — Editor.]
-[Mammonteus etjuals Mammonteus primigenius alaskensis sp. nov., from near Fairbanks (Long. 148.10 W., Lat. 64.59 N.) the description of which will be
found on page 1159 below. — Editor.]
 THE MAMMONTINiE: MAMMONTEUS 1135

a camel, and the recently discovered lion^ or tiger {Felts atrox alaskensis Frick). Alaska was free from glaciers,
except in its central mountainous belt. Scattered remains of mammals occur: (1) In the frozen tundras along
the northern coast, the black muck accumulated in gulches and valleys of the smaller streams, (3) in the
(2) in
'Yukon silts' and the 'Kowak clays,' (4) in the more recent fluvial and alluvial deposits.
fine elevated clays of the
In the northerly clays of Kotzebue Sound, Eschscholtz Bay (1907-1908), Quackenbush discovered a female
mammoth skeleton (Amer. Mus. 13749) with portions of hair and wool perfectly preserved. Recent stripping
operations of the U. S. Smelting and Alining Company, on Gold Stream and Clery Creek near Fairbanks (Fig.
1004), yielded the following (Frick, 1930, p. 73): ". during the four summer months some twenty-eight large
. .

cases of skulls, jaws, and bones — rare and important evidence on the prehistory of Alaska which otherwise
would have been lost to science. The great percentage of this material, interestingly enough,
came from three
restricted areas, 'bone pits,' scattered between several widely separated operations of the Company, the re-
mainder of the worked areas being, for the purposes of the bone hunter, nearly barren."
Pre-Tundra or Fossil Fauna of Alaska-Yukon
Mammonteus primigenius Blum. Symbos tyrelli Osgood Bison alleni Marsh Equus lambei Hay
M ammonteus -primigenius compressus Osb. Bootherium sargenti Gidley Ovis sp. Camelops sp. Gidley
Mastodon americanus Kerr Ovibos yukonensis Gidley Oreamnos sp. Arctodus yukonensis Lambe
Rangifer sp. Ovibos moschatus Zimm. Equus alaskas Hay .Enocyon dirus alaskensis Frick
Bison crassicornis Rich. Felts atrox alaskensis Frick

Am er Geoijr Soc ATKI

Geographic Range of Mammonteus and of Parelephas

Superseded by figure 79.5, Chapter XV, above
Fig. 1005. This diagram shows that these two mammoths, confused in all previous descriptions, were north temperate and circumpolar in distribution.
Closest to the Pole and to the northern ice-sheets during the Fourth Glaciation (Wisconsin-Wtirm) was doubtless the range of Mammonleug primigenius;
closest to the 40th parallel was doubtless the chief range of Parelephas trogontherii, P. intermedius, and P. jeffersonii. It is possible that Parelephas Irogon-
iherii and its successor P. intermedius migrated (diagonal) at the close of 1st or 2d Interglacial time across northern Asia, thence across Bering Strait into

Alaska and southward to the 40th parallel of the United States.

'According to Frick's researches (letter, March, 1930), the cranium of Felis atrox alaskensis resembles that of the lion {Felis Leo) rather than that of the
tiger (Felis tigris).
1136 O.SBORN: THE PROBOSCIDEA

Fig. lOOG. Geographic distribution of the principal species of il/a/n»wn(eus. Tlic white dots within the black areas represent the approximate localities
where the types of these twenty-nine species and subspecies were discovered. The white cros.ses represent referred specimens.

3. HISTORICAL ORDER OF NAMING OF SPECIES OF MAMMONTEUS EXCLUSIVE OF SPECIES

WHICH ARE NOW KNOWN TO BELONG TO PARELEPHAS TROGONTHERII, ETC.

See Figure 1006

Late Pleistocene 1.
 :

THE MAMMONTIN^: MAMMONTEUS 113/

20. 1835 Elephas odontolyrannus Eichwald, banks of the Nicnian, (list, of

Novogrodek, gouv. Vilna, Russia = Mammonteus primigenius
21. 1845 E. [Elephas] primigenius sibiricus^ de Blainville (Atlas, "Osteographie,"
PI. in) = Mammonteus primigenius
22. 1879 Elephas primigenius comune Issel [in Zuffardi, 1913, p. 136], Campo-
rosso, near Ventimiglia, Italy Mammonteus primigenius
(fide Issel, 1879, p. 160)
Upper to Lower 23. 1888 Elephas (primigenius) Leith-Adamsi Pohlig, Dornap (Fuhlrott), = M ammonteus C.) primi-
Pleistocene Thuringia, Germany genius leith-adamsi
24. 1891 Elephas priinigenius Blum. var. hijdrunliyms'Botii, Olranto, Italy = Mammonteus primigenius
hydrunlinus
25. 1912 Elephas primigenius Fraasi Dietrich, Steinheim, Germany = Mammonteus primigenius
fraasi
1913 Elephas primigenius Blum. var. Irogontherii Pohl. Zuffardi = Mammonteus primigenius
mutation astensis (see

No. 26 below)
Ujjper Pliocene'- 26. 1923 Elephas primigenius mutation astensis Deperet and Mayet, San Paolo = Mammonteus primigenius
de Villafranca, Italy astensis

2. SPECIES AS NAMED IN NORTH AMERICA

Upper Pleistocene 27. 1842 Elephas americanus DeKay, Monroe County, near Rochester, = Mammonteus primigenius
New York americatrus
28. 1924 Mammonteus primigenius compressus (Jsborn, Rochester, Indiana = Mammonteus primigenius
compressus
29. 1933 Elephas primigenius alaskensis Osborn (in Frick, notncn nudnm), = Mammonteus primigenius
vicinity of Fairbanks, Alaska alaskoisis sp. nov.

Possibly twenty or more of the above species are synonyms of Mammonteus primigenius and probably belong
to the Upper Pleistocene drift and Postglacial deposits of the Fourth Glaciation. Four of the above, more primi-
tive, of greater geologic age, and with probably lower ridge formulae, are as follows

1888 Elephas primigenius (?) Leith-Adamsi Pohlig, Nova Acta Leop. Carol., LIII, Nr. 1, i^p. 229, 232.

1891 Elephas primigenius var. hydrunlinus Botti, Bol. Soc. geol. Ital., IX, p. 709.

1912 Elephas primigenius fraasi Dietrich, Jahresh. Ver. vaterl. Naturk. Wiirttemberg, LXVIII. This species is fully annotat-
ed below.
1923 Elephas primigenius nuit. astensis Deperet and Mayet, "Les Elephants Pliocenes," Deuxieme Partie, pp. 183, 184. This
species is fully annotated below.

4. GEOLOGIC AND PROVISIONALLY ASCENDING PHYLETIC ORDER OF SPECIES AND SUB-

SPECIES OF MAMMONTEUS. EURASIA AND NORTH AMERICA
Falconer and Hay have referred to Elephas primigenius the American specimens of the true woolly mammoth,
the ridge formulae of which remain to be more precisely determined. Falconer attributed to the Ainerican speci-
mens the same ridge formula, namely, M 3 M, but remarked that the ridge-plates were more compressed or closer
together; this compression reaches the highest possible stage in Mammonteus primigenius compressus, in which

'[See footnote on p. 1391 below « here it is stated that E. primigenius sibiricus and other .suljspetics "perliai)s may be regarded as geographic designations
rather than as .subspecies." —Editor.]
"[Possibly Lo«er Pleistocene (see footnote 1 on p. 1049 above). — Editor.]
1138 OSBORN: THE PROBOSCIDEA

the ridge-plates increase to M3 tr. Altogether, including Upper to Lower Pleistocene and Upper Phocene stages,
six' species and subspecies have been described as follows:

Upper Pleistocene Alaska, Indiana Maniniontcus primigenius compressus = The most extreme or progressive
IV Glacial and Postglacial Osborn subspecies known

Ujjper Pleistocene Alaska ant! the Mammonteus primigenius americanus --

Slightly progressive American
IV Glacial and Postglacial United States DeKay stages

Upper Pleistocene Siberia and west- Mammonteus primigenius primigenius = Typical or true mammoth
IV Glacial and Postglacial ern Europe Blumenbach M3fl
Srf(?) Interghcial Western Europe Mammonteus pritnigcnius fraasi Dietrich = Ancestral Upper(?) Pleistocene
subspecies of Mammonteus
primigenius
M3^
3rf(?) Interglacial Thuringia, Mammonteus primigenius Leith-Adamsi = Small primitive branch of the
Germany Pohlig main stem of Mammonteus
primigenius
Lower Pleistocene Forest Bed, East Mammonteus primigenius astensis {"f) ref. = (Of. Falconer, 1868, II, p.
Anglia Deperet and Mayet 170): Small "slightly thick"
. pre-glacial variety of
. .

Elephas primigenius from

the Norwich coast (cf Fig. .

1020).

Upper Pliocene- Italy Mammonteus pri7nigemus astensis type = Ridge formula: M 3 -ff^^
Deperet and Mayet (cf. Fig. 1019)

As shown above, the typical or true mammoth was found in Siberia long before it was recognized in western
Euroj^e. Blumenbach had in mind both Sil^erian and North German specimens in defining Elephas primigenius
in 1799 (1799.1, p. 697), also Cuvier had the Siberian mammoth in mind in defining Elephas mammonteus in

1796 (MS., published in 1799). Unfortunately Blumenbach in his type description mentioned as an example the
Burgtonna skeleton, which we now know belongs to 'Elephas antiquus' and is not the type. The exact evolutionary
stage of 'Elephas primigenius' was finally defined and determined by Falconer in 1863 (pp. 64,65), with a typical

ridge formula of M 3M. Attributed, or referred, to the typical species Elephas primigenius have been all the
Pleistocene mammoths of western Europe and North America including, as we have seen, all the species of Parele-

phas also all the primitive and geologically ancient species and subspecies of the true mammoth. The American
subspecies Mammonteus primigenius americanus has the same ridge formula but the plates are more compressed
{fide Falconer). Consequently the recognition of the geologically ancient forms with relatively low ridge fornuilse,

as listed above, is a matter of the utmost importance and interest.

We somewhat problematic whether the Upper Pliocene Mammonteus primigenius astensis

regard it as
Deperet and Mayet, 1923, is ancestral to Mammonteus; certainly it shares characters of Mammonteus and of
Parelephas; its ridge formula of M 3 {—, is higher than that of the contemporary Parelephas species P. trogon-
therioides.

The Forest Bed stage, previously determined by all authors as the true Matnmonteus primigenius, more
probably belongs to a primitive stage with f(nver ridge-plates, perhaps M 3 lliff, similar to M. primigenius
astensis. From among the considerable number of Forest Bed specimens probably a correct ridge-plate count
can be made.
'[To these should be added the new subspc(;ics Mammonteus primigenius alaskensis (sec pji. 1 U>9 to 1 101 of this chapter for description). — Editor.)
^[See footnote I on page 1049.— Editor.)
 . .

THE MAMM0NTINJ5: MAMMONTEUS 1139

AuRiGNACiAN Mammoth Hunters OF MoRAViA.— The following passages may be freely cited from Doctor
Absolon's recent description in the Illustrated London News (Nov. 23, 1929) of his remarkable discoveries in the
years 1924-1929 of mammoth-hunting stations of Moravia:

of these stations we now know about one hundred in Moravia,

but only a small part of their area is explored as yet. ... It is
because Moravia was a kind of passage, by way of which the fossil
mankind of the increasing Aurignacian tribe penetrated from Asia
through Russia to the West of Europe. They are: Vestonice, . . .

Pfedmosti, Pctfkovice, Pekarna, and Ondratice. Peknrna is a cave

(Fig. 1) ; the rest of them are in loess on the slopes of hills. Pekarna
is the most important, Vestonice the largest of them. . . . This was
the first culture of the Old Aurignacian invasion of mammoth-
hunters coming frcm Asia, the cradle of mankind. . . .

One thing is certain —

namely, that the mammoth-hunters
killed these huge pachyderms in hundreds, and that in diluvial
Moravia, a great tragedy, like the destruction of elephants in
Africa, took place. . .

In the refuse-heap discovered in 192.5, no tusks were found in

the whole area of 45 square metres; but in that encountered in
1926, on the other hand, three heaps of tusks were piled one o\er
another, between which was left a narrow path (Figure 1.3). In
the same year an imposing sight opened before our astonished eyes,
when we discovered a field of huge pelvic bones (Fig. 14) of adult
mammoths. The skulls are usually broken to ]iieces, because mam-
moth-brain was appreciated by primitive hunters as a delicacy.
Nevertheless, we found an intact skull of strikingly large propor-
tions in 1928 (Figure 15). Long bones (femurs, tibiae) were found
also in strange po.sition forming a half-circle, so that their broken
ends all pointed in one direction: evidently the fire was kept
alight by the fat which flowed out of the ends of the burning bones
into the flames. . .

Lower jaw-bones lie generally apart, and the teeth have often
keen knocked out and jiiled up in heaps. In 1927 we found at
Pfedmosti a jaw-bone within which was a red-])ainted stone club
(Fig. 9) which might have been used for striking out the teeth from
—
the jaw-bones a unitpie discovery. From 1924 to 1929 we count-
ed as many mammoths, all of thcni caught and killed by
as sixty
man, on the area of 1600 square metres.
There cannot be the least doubt that Ihe hunters did not
attack these powerful animals 'face to face,' but caught them by
cunning, enticing or driving them into large jntfalls. The picture
(Figure 16) shows a stratum dipping abrujjtly downwards. It
must have been purposely made; dug in diluvial times. We
intend to try to open this pitfall, for such I take it to be. Mam-
moths trapjied and caught were kflled by large stones, trimmed to Fig. 1007. Skeleton of the Mammoth {Mammonleus primigenius) from
serve such a purpose. These stones might have been suspended in the dihivium of Moravia, now in theMoravian Government Museum of Brno
strong leather straps and thus let down on the animals by the (Briinn), Czechoslovakia. After photograph furnished by Prof. Karel Absolon

united efforts of .several men, in the same way that navvies drive of the University of Prague and Curator of the Museum at Briinn. This
skeleton was found nearly complete at Piedmont in Moravia, whereas all the
piles into river-beds by means of rams. 1 have found one such
skeletons in the kitehen-middens of the mammoth hunters of Moravia were
stone, trimmed like a big pear, or bomb, 1 metre long, and weighing scattered and partly destroyed for a footl supply. It measures 3 m. or 9 ft.
over 120 pounds (Fig. 11). [8ee Fig. 1037, tiiis chapter. Editor.] — 10 in. in height and -!.12 m. or 14 ft. 6 in. in length.
 III. SYSTEMATIC DESCRIPTION OF SPECIES OF MAMMONTEUS
Specific Revision. — (1) All the earlier writers even up to the time of Lydekker's "C'atalogue of the Fossil

Mammalia in the British Museum" (1886.2) included within the typical species Elephas [
= Mammonteus] primi-
genius all the progressive and primitive maumioths with fine-plated molars discovered in Pleistocene deposits,

from the typical horizon of the Fourth Glaciation downwards to the Forest Bed stages of the Lower Pleistocene.
(2) They also included within E. primigenius, as we have seen above in the Parelephas chapter, many stages
properly belonging to Parelephas trogontherii and its ascending and descending mutations. (3) Within the last

half centurj^, however, Pohlig (1888) and Dietrich (1912) have separated the stages of 2d and 3d Interglacial times
in Germany as Elephas primigenius leith-adamsi Pohlig, 1888, and as Elephas primigenius fraasi Dietrich, 1912;

(4) also Deperet and Mayet (1923) believe they have recognized an Upper Pliocene (Villafranchian) stage, to which
they have given the name Elephas primigenius mut. astensis. Thus there have been four great steps in specific
revision, concluding with the present revision by Osborn.

As shown in the table above, the highly specialized Mammonteus conforms with the principle of ridge-plate
addition as we pass from Upper Pliocene to the close of Pleistocene times, so beautifully manifested in the two
more generalized mammontines Archidiskodon and Parelephas. We may select the third superior and inferior

molars to show this parallel ridge-plate progression of M 3 in two of these three great lines of ascent.
 .:

THE MAMMONTINiE: MAMMONTEUS 1141

SYSTEMATIC DESCRIPTION OF SPECIES

Mammonteus priniigenius Bhimenbach, 1799, 1803 plementum, 1904-1905, p. 600) remarks: "Le nom barbare de
Figures 794, 81G, 818, 865, 934, 937, 962, 990-999, 1000-1002, 'Mammout,' base, par Blumenbach, sur une erreur grossiere
1007-1014, 1032-1034, 1042, 1062, 1084, 1226, PI. xxii (ridentitc du Maslodonte de I'Ohio avec V Elephas primigenius),
Upper Pleistocene, IV Glacial drift(?), Siberia and northern Germany. n'a aucun droit, malgre sa priorite, a etre substitue a celui de 'Mas-
For synonyms, see page 1 136. todon,' genre bien caracterise par Cuvier."
Type Description of Elephas primigenius, 1799. (Bhimen- — Osborn, 1924 The above paragraphs from the quaint writings
:

bach, 1799, p. 697) "3) Von einem ungeheuer grossen Elephanten

:
of Blumenbach and of Artaud firmly establish the specific name
(Elephas primigenius?) [die vermeinten Riesenknochen** unsrer Elephas primigenius as of date 1799, 1803; they show that
ehrhchen Ahen] unter andern auch in Menge in Deutschland***)
;
Blumenbach erroneously selected the Burgtonna skeleton' as an
So z. B. das berijchtigte Elephantengerippe das 1695 bey Burg- example of E. primigenius; they show that there was no type
Tonna im Gothaischen ausgegraben worden etc." figure. Finally, as pointed out by Trouessart, Blumenbach applied
—
Lectotypes. Blumenbach's type description (1799, p. 697) the barbaric name "Mammut" to the American Mastodon of Ohio
is in a rare document; no type figure accompanies it. Blumen- under the erroneous idea that the latter was related to Elephas
bach had the Siberian and German mammoth in mind. The 'Ele- primigenius. ,

phantengerippe' of Burgtonna is not the type. Dietrich and

Osborn select (1930) as lectotypes: (1) A Siberian grinder in the
Blumenbach Collection of the Zoological Institute of the Uni- TYPICAL UNIFORM RIDGE FORMULA OF MAMMONTEUS
versity of Gottingen, and (2) a molar from Osterode (Harz), PRIMIGENIUS OF EUROPE
Germany. The sixth edition of his "Handbuch der Natur- —
(Falconer, 1863). As to the ridge formula of the mammoth.
geschichte," in which the type description appears, attracted the Falconer (1863, p. 64) remarks that in the European and American
attention of the French naturalist Soulange Artaud, who visited specimens the characters are constant, as defined by Cuvier and as
Blumenbach and translated his "Handbuch" into French under witnessed in the British Museum collections from Siberia and in the
the title "Manuel d'Histoire Naturelle." In the manuscript of the American collections from Eschscholtz Bay. He concludes (op.
French \olume of 1803, Blumenbach inserted the name and defini- cit., p. 65): "They aU present, in the main, the same characters:
tion of Elephas primigenius, consequently the species Elephas a uniform ridge-formula; the same obtuse form of the lower jaw,
primigenius is of date 1799, 1803. and the same broad crowned molars, composed of closely com-
Second Description, 1803 (Blumenbach-Artaud). — Many pressed cplUculi, with numerous digitations and attenuated un-
authorities, namely, Trouessart (1897, p. 711), Lydekker (1886.2, crimped .enamel-plates. One of the most essential points, is to
. . .

p. 175), and Hay (1902, p. 713), erroneously cite 1803 as the date of determine the constancy of the ridge-formula [italics Osborn], which,
the type description of this species, i.e., "Manuel d'Histoire after the examination of a very large quantity of materials, I
Naturelle, traduit de I'Allemand, de J. Fr. Blumenbach, . . . par believe in the Manimoth to be thus:
Soulange Artaud," 1803, Vol. II. The explanation of the definition
of a new species in the translation is found in the "Preface du Tra- [Dp 2) i, [Dp 3] f [Dp
,
4] H, [M 1] 41, [M 2] if, [M 3] M."
ducteur," p. xvi: "C'est que j'ai travaille sous les yeux de M.
Blumenbach lui-rneme; qu'il a eu la complaisance de revoir mon
Of this ridge formula he observes {op. cit., p. 65) "The plates :

manuscrit, et qu'ainsi je puis presque repondre de la fidelite de la

advance by quaternary increments in each series, bearing in mind,
traduction." This shows that Blumenbach inserted the name and
that the first commonly
true molar, although of larger dimensions,
definition of E. primigenius in Artaud's MS.
repeats the number of by the milk-molar,
ridges presented
last and
In Volume II, p. 407, of Artaud, we find the following definition Mastodons more
that the last true molar in all the Elephants and is
"3.° D'un elephant enormement grand {elephas primigenius) (ce
composite than the others. The formula in the North American
. . .

sont pretendus os de geants de nos bons aieux); il se trouve

les
Mammoth is identical with that of the Siberian and European
des OS fossiles de cet animal, en grande quantitc, en Allemagne . . .

forms. Exceptions are occasionally met, in which an unusual

(Le squelette d'elephant, par exemple, qui a ete trouve, en 1695,
number is presented."
of plates
pres de Burgtonna, dans le pays de Gotha)."
In Artaud's translation no figure of Elephas primigenius oc-
Comparison with Elephas (Euelephas) indicus. — Fal-
coner (1863) considers and repeatedly states that the ridge formula
curs,although in the explanation of the figures appears the Ameri-
of E. primigenius is closely similar to that of the subgeneric group
can Mastodon designated as "28. Fossile Ohio incognitum" and
on page 408 designated as "le mammouth de I'Ohio {mammout as typified by E. {Euelephas) indicus in which he corrects his
formula of 1857, p. 315, and substitutes {op. cit., p. 65) for E.
Ohioticum)," also the "Table Alphabetique des Noms Latins, de
indicus the following collective formula:
Genres et d'Especes" of Artaud's translation, p. 443, gives the
orthography as "Mammut Ohioticum."
Trouessart (1904-1905).— Trouessart (Quinquennale Sup- Dp 2 1, Dp 3 f Dp 4 «, , M H M 2 if M 3 ^Ht-
1 , ,

**f. Voigts Magazin. V. B. I. St S. 16 u. f. .

***(Kriegsr. Merk) kllres sur les os fossiles d'elephans el dc rhi7ioceros qui sctrouvent en Allemagne Ac. I-IU. St. DuTmst. 1783 u. f.i,

'See footnote 2 on page 1118.

 —

1142 OSBORN: THE PROBOSCIDEA

Errohs of Lkith Adams, I.ydkkkeh, Hay. — Falconer, with dental and cranial characters of Elephuti prunigenliis. He did not
unerring eye and logical mind, dwelt upon the constancy of the confuse with this true species of mammoth (E. priniigcniuK) the
ridge formulse of other species of elephants, as was done by sub-
sequent writers, i.e., Leith Adams, Lydekker, Felix, Soergel, and
Air 1/4 nit. sit«
Hay, who assigned to E. primigenius the collective ridge fornuda
derived from a considerable number of species belonging, as shown
in this Memoir, to two distinct phyla, namely, to Mammonieus
1^'
E. primiooniua and to Parelephas. As to the ridge formula of Elephas primigenivs,
23 Am.r Mus. 13752 Re(.
Felix (1912) erroneously follows Leith Adams in giving the ridge
formula of E. primigenius as M
18-27 (?29)- This erroneous col-
3 ji
lective ridge formula first appears in Leith Adams (1879),' then in
Lydekker (1886.2, p. 175), and finally reappears in Hay (1914, p.
395), as follows:

Collective ridge formula: Dp 2 t Dp 3 Dp 4 1^;^ Ml

E, prrmiflenius

Am»r. Mui. 13752 Ref. The above partly erroneous formula is imdoubtedly based on
the ridge-plate formulse of species of Parelephas combined with the
ridge-plate formulae of species of Mammonteus. This confusion
throughout Hay's observations on the grinding teeth of Elephas
primigenius (1914, pp. 395-410) imfortunately renders his observa-
tions of Uttle systematic or phylogenetic value. Hay's obser-\-ations
had been summarized for extensive citation in the present Memoir,
but for the reasons given we are obliged to omit them.

Osborn's Observations (1922-1924) on American Museum

Specimens Referred to Mammonteus primigenius
E. primigenlus Osborn (1922) supports Falconer's observations of 1863.
Am>r. Mui. U371 Rit. Compare Osborn, 1922.555, pp. 7-11. Elephas primigenius
(Innir vllw)

is the name assigned to a series of species which ranged through the

entire Pleistocene epoch, from the Lower Pleistocene Forest Red'^

deposits of East Anglia to the southerly range of this animal in the
middle United States. It is a collective species embracing an
undoubted progressive evolution and intensification of specific
characters extending over a very long period of time. It exhibits

\arious extremes of fore-and-aft comjjression of the craniimi

E. prifniginiua
with related fore-and-aft compression of the grinding teeth {hgp-
Ainw. Mui. I«37i Rat sicephali/and balhycephaly). The cranium is high, jiointed at the
summit, relati\ely narrow, and relatively ileej). The forehead
from the peak of the craniiun to the extremity of the na.sals is rela-
tively elongate and sUghtly concave.
Kkfkhked M.mvjmontfvs rRiMiGBNius Molars of Ala.ska with Tvi-ical
M. PRiMiGENiu.s Ridge Formula
l{iD(iE-PLATE Compression. —
As comi)ared with E. cohnttbi
and E. imperalor, 10 ridge-plates of M. primigenius are compressed
l'"ig. 1008. 'I'hinI .superior and inferior molars (twcnty-tlircc to twenty-
four ri(lKe-i)lates) of Mammoiileus pritnignnux, Ala.ska, of less
into a line 100 mm. in length in the typical M. primigenius of
(•omi>re.s.se(i type

(Am.r. Mils. After ()sl)orn, 1922..wo, p. 10, fig. 9. The ridge

]37.')2, 14:i71). (iermany; the most highly compressed tooth ohserxed by Osborn
formula of these typical s|)ecimfns of M. primigenius porrespond.s very closely is an M^ from Alaska (Amer. Mus. 13749 [paratype of M. primi-
with tliat given by Falconer ( 1863) as characteristic of western Europe, namely, genius compressus]) in which 13 ridge-plates are compressed into
a line 100 mm. in length (Fig. 1024) ; a similar condition pre\'ails in
A, Crown view, leftM', twentj-three ridge-plates. A 2, Outer view of
a female skull from Indiana (Amer. Mus. 14559 [since made the
same tooth. It is apparent that one or )«)ssibly two of the anterior ridge-plates
type of M. primigenius compressus]), in which 13 ridge-plates arc
have been worn off, consequently the formula should be written: M .3 '„^-.

H 2, Inner view of right M3 of another individual, twenty-four ridge- compressed into a 100 mm. space, the total number of ridge-plates

platcs. B, Crown view of .same. rising to 27 (Fig. 1022).

'See Bibliograi)liy of the present Memoir, Vol. I, p. 762, under Leith Adams, 1877-1881.
"The Forest Bed Mammonteus primigenius as(ensts(?) is diagrammatically figured (Fig. 871, Archidiskodon, Chapter XVI, the present Chapter, Fig. 1020)
in comparison with the contemporaneous molars of Parelephas (C, D) and with Archidiskodon (E-H). This figure by the author shows (.\) the thin, finc-platcd
or "slightly thick" (fide Falconer, 1868, Vol. II, p. 170) enamel characteristic of Mammonteus at the beginning of Pleistocene time.
 : : : —

THE MAMMONTIN^: MAMMONTEUS 1143

The is of the kind shown in

prevailing condition, however, COMPARISON WITH PARELEPHAS AND MAMMONTEUS
figure 1008,namely, Amer. Mus. 13752, from Alaska, in which the PRIMIGENIUS COMPRESSUS
ridge formula is M
3 ^-^, here figured with a lower molar (Amer. By the above observations of Osborn, the typical ridge fornuila
Mus. 14371), in which the ridge formula is 3 ^; in these speci- M of M 3 of Mammonteus primigenius agrees precisely with that es-
mens there are 10 plates in a 100 mm. line; these less compressed tablished by Falconer in 1863, namely: M 3 W. This relatively
molars are arcuate, thus the count of the ridge-plates is greater on low ridge formula agrees approximately with that of Parelephas
the concave side of the tooth and smaller on the convex side. jeffersonii,namely, M
3 M, but the M^ of M. primigenius is
relatively shorter and deeper than the M' of P. jeffersonii; conse-
For example, in a superior molar, M' (Amer. Mus. 10656),
quently the ridge-plate compression is much closer in M. primi-
from Germany, the compression is as follows
genius. The ridge-plate compression is still closer in Mammonteus
M' 10 ridge-plates in 100 mm. on the external convex surface,
primigenius compressus, in which M3 is the broadest and deepest

11 ridge-plates on the horizontal mid-coronal surface,

proboscidean molar known, the formula being M 3 i^^, the ridge-
plate compression rising to 13 in 100 mm.
Undoubtedly inter-
12 ridge-plates on the internal concave surface.
mediate ascending mutations will be found between these two
stages of ridge-plate evolution; meanwhile they may be distin-
In the lower molars, these M' conditions are reversed in count- guished as follows
ing the ridge-plates in which the external surface is concave and Typical Mammonteus primigenius primigenius of northern
the internal surface is convex. For example, in an M3 from Alaska Europe, Alaska, and the United States: ridge formula 3 fr; M
(Amer. Mus. 14343) the count is as follows: ridge-plate compression in the superior molars 10-11-12 in 100 mm.
Mammonteus primigenius compressus of Alaska and Indiana:
Ms 10 ridge-plates in 100 mm. on the external concave surface, ridge formula M
3 (7)^; ridge-plate compression 13 in 100 mm.
10 ridge-plates in 100 mm. on the horizontal mid-coronal A fresh survey of the true Mammonteus primigenius grinding

surface, teeth, of Alaska and the northern United States, will probably
8-9 ridge-plates in 100 mm. on the internal convex surface. reveal a series of intermediate ascending mutations between these
two extremes. When the grinding teeth which have been errone-
ously referred to Mammonteus primigenius are eliminated, namely,
In the highly compressed superior tooth, M' (Amer. Mus.
all of those molars actually belonging to Parelephas, we shall prob-
13749 [paratype of M. primigenius compressus]) from Alaska (Fig.
,

ably discover these transitional mutations.

1024) the count on
, all three measurements is the same, namely
SKULLS AND JAWS OF MAMMONTEUS PRIMIGENIUS
Compare Figures 86.5, 962, 1009, 1010, 1023
M' 13 ridge-plates in 100 mm. on the external convex surface,
The true cranium of Mammonteus primigenius (Figs. 865, 962
13 ridge-plates in 100 mm. on the horizontal mid-coronal
C, C) is readily distinguishable from that of Parelephas trogontherii
surface,
(Fig. 865) and of P. jeffersonii (Fig. 962 A) with which it has been
13 ridge-plates in 100 mm. on the internal concave surface.
confused except by Fohlig. The jaws of M. primigenius (Fig. 962
C, C', and Fig. 1011 C,B, A) are also clearly distinguishable. In
from the above observations that the horizontal
It follows this connection observe the characters pointed out by Falconer,
mid-coronal section gives, as a rule, an average between the internal Leith Adams, Pohlig, and others enumerated above.
and external sides the average in M. primigenius is 10 ridge-plates
;
Cranial Materials Figured Herewith (Figure 962).
in 100 mm., the minimum observed is 8 in 100 mm., the maximum Crania of the true Mammonteus primigenius, as distinguished from
observed is 13 in 100 mm.' This variation in compression applies crania of Parelephas jeffersonii, are very rare in the United States.
to fifteen specimens ranging through England, Germany, Siberia, Fortunately there is a finely preserved cranium from Siberia in the
Alaska, Ohio, and Indiana, and characterizes the typical M. primi- United States National Museum (Nat. Mus. 8580), photographs
genius M' of Thuringia, Germany (cf. Soergel, 1912.2, Tab. vii, ofwhich are reproduced in figure 962; the lateral aspect of this
VIIl). cranium (Fig. 962 C) is perfect, but the frontal aspect (Fig. 962 C)
The worn mid-coronal surface is sometimes fully horizontal, is foreshortened and thus gives an erroneous impression of the
i.e., at right angles to the perpendicular ridge-plates; in such case height of this cranium.
it registers the exact distance between the plates. In other cases —
Cranial Characters. All the distinctive characters of the
the wear is obliquely horizontal; in which case it increases the true Mammonteus primigenius skull arise from the maximum /oce-
actual distance between the plates. important to note also It is and-aft compression, resulting in bathycephaly and hypsicephaly,
that the ridge-plates are arcuate and closely compressed more which exceeds by far that ofany other proboscidean or other
towards their summits; thus more ridge-plates may be counted mammalian skull thus far known. (1) This compression brings the
in 100 mm. at the summit of the crown than at the base of the anterior rim of the orbits much closer to the occipital condyles
crown, and as a rule the ridge-plate count should be taken midway (Fig. 962 C) than in either P. jeffersonii (Fig. 962 A), or P. wash-
between the summit and the base, both on the internal and external ingtonii (B) ; (2) it elevates the occipital crest, which is relatively
sides. higher, more elevated, and more pointed, i.e., acrocephalic, in M.

'[In Mammonteus primigenius compressus Osborn. —Editor.;

1144 OSBORN: THE PROBOSCIDEA

CoMrAIilSON OF MaMMONTEUS PRIMIGENIUS (C, C') AND OF PaKELEPHAS (A, B) CuANIA

See also comparative figure (Fig. 865) of the European Crania

Fig. 1009. Profile views of type and referred .skulls of Farpicp/ias Je^ersonii (A), P. washingtonii (B), and Manimonleiis primigcnius (C) in the American
Museum and United States National Museum, also front view of M. primigenius (C). All figures one-twelfth natural size.

A, Type of Parelephas jeffersonii (Amer. Mus. 99.50), reversed. An aged individual. Main jjortion of the tusks not in<luded. From Jonesboro, Indiana.

B, Referred skull of Parclcphnx imxhinglotni (.Xnier. Mus. Coi)e Coll. 8081). From Whitman County, Washington.
C, Referred skull of Manunonkus primigcnius (Nat. Mus. 8.380), from Siberia, with jaws belonging to another individual (Nat. Mus. 8.079), from .Maska.

C, Front view of same skull and jaws.

pniniijcnius (I'ig. 9(52 (") tliuii it is in /'. ji'Jhrsoiiii (i'ig. 962 A) Figures 865, 934, 937. — Comparison with crania referable to
or P. ivashinglonii (Fig. 962 B). (3) This fore-and-aft coni])rcs.sion the typical M. pritnigeniiis of western l']urope
is afforded in figure

brings tlic aiulitory foramen of ^f. priiniqciifiift clone to tlie occipital 865, esijccially the frontal view (No. 2) inscribed " E. primigcnius
condyles; it whereby tlie apex of the
increases the hijpsirephulij, lief., Falc, 1847, PI. xliii, Fig. xxiv"; this west European
occipital crest is above the grinding surface
raised relatively higher cranium may be considered a "typical Mammnnteus primigenius."
of M'; and (4) it accounts for the extreme hypsodonty of M', As far as we can
jiidgc from l''alconer's figure (1847, PI. xi.iii, fig.
which is the highest and the shortest of all ])roboscidean molar XXI v) cranium shows exactly the same characters throughout
this
teeth. (5) This compression or bathycephaiy extends to the lower as the crania above described from Alaska and from Indiana; we
jaw which has an excessively short, deejjly deiiressed ramus, observe esi)ecially the small anterior nares, the tubular tusk
terminating in a deflected and extremely narrow rostrum (Fig. insertions, the extremely ele\ated occiput.
962 C, C). (6) This vertical compression or hypsicephaly extends Another "typical Mammonteus primigenius" cranium (Figs.
to the vertically placed tubular maxillo-premaxillary in.sertions of 865, 934) is that figured after Pohlig (Pohlig, 1S91. j). 3S4, fig.

the incisor tusks (Fig. 1023), which are relatively longer, narrower, 120 — reversed); this cranium in profile view may be compared
and deeper than those of any species of Parelephan. (7) The with that of the female skull {Mammonteus primigenius compressus)
frontal or facial as])ec( of MnmmontriiK primigenius romprrstnis from Indiana (l''ig. 1023) in its extreme acrocephaly, hypsico)ihal.y,
(Fig. 1023) is (luite distinct from the frontal or facial aspect of for(!-and-aft comjiression, deep INT' inserti(Jii, and relatixcly plano-
Parclephas (Fig. 961); the arches supporting the orbits appear frontal and occipital surfaces. In brief, the west European crania
relati\ely broader, because the frontals above and the maxillaries of the true M . primigenius exhibit jireciscly the same characters as
below are laterally compressed. the true M . primigenius crania of Alaska and of Indiana. They
 THE MAMMONTIN.E: MAMMONTEUS 1145

completely confirm the specific and generic separation of the Mus. 8579--Fig. lUll B) is also from Alaska, and indicates twenty-
cranium of Mammotdeus from the cranium of Parelephott and still four ridge-plates; 4162— Fig. 1101 A) is
the third jaw (Nat. Mus.
more from the cranium of Elcphus. Doubtless other true crania of from Kk'ijhant PoinI Alaska. We owe to Gidley exact information
,

M. primigeniiis will be discovered in the United States and in regarding the American crania and jaws referred to Mammonteus
Alaska; it is of the utmost importance that these should be care- primigenius in the present Memoii-.
fully exhumed and that they should find their way permanently to
some museum.
Figure 1010. —
It is through the courtesy of Charles W. Gilmore
of the United States National Museum that we reproduce herewith
(Fig. 1010) a superb cranium and tusks of M. primigeuius discover-
ed in the Yukon Territory, Canada, and described by Gilmore
(1908, PI. vii); it is difficult to determine from the photograpli
whether this cranium belongs to the true MaiNnionlcds or to
Parelephas; we are inclined to relate it to Mammonteus. A profile E. primigenius
Nal. MiiS.6666 Ref.
photograph of this fine skull would at once determine the question
of its affinity.

E. primgerMu5
Nat. Mus. 8579 Ref.

1234 eg

E. primigenius
Nal. Mus. 4162 Ret
CR.'VNinM OF MALE MaMMONTEUS PRIMIGENIDS OF THE YuKON
Fig. 1010. Male skull and tusks probably referable to Mammonteus
primigenius, found in gravel 42 feet below the surface on Quartz Creek, near
Dawson, Yukon Territory, Canada, Maroh, 1904. After f)hotograph by
Gilmore appearing in his important paper entitled, "Smith.sonian Exploration
in Alaska in 1907 in Search of Pleistocene Fossil Vertebrates, Second Expedi-
tion," 1908, PI. vii.
As observed in the text, the generic and specific reference of this skull is
somewhat uncertain; the very prominent antorbital tuberosities resemble Growth Stages in the Jaws and Teeth of Mammonteits primigenius
those in the type female skull of Maimiionleus pninigeiiius cnmpressns of Fig. 1011. Internal aspect of three jaws of typical Mammonleus primi-
Indiana. genius from Alaska, in the United States National Museum. \l\ one-eighth
natural size. Sections CI, Bl, Al.

Figure 1023.— Fortunate also is the presentation of the skull of C, Juvenile jaw of Mammonteus primigenius (Nat. Mus. 6666) from
a female mammoth (M. pnmigeniv>i rornpressus type) in the Ala.ska, twenty-five miles above Anvik. Ridge formula: M
2 yjtjl, 3 Y2+- M
American Museum (Anier. Mus. 14559) from Rochester, Indiana, In M2 at least one plate ismissing in front of the alveolus of the anterior root,

Mam- therefore there may be four or more plates missing; total preserved thirteen,
which more correctly shows the frontal plane of the true
maximum seventeen ridge-plates in Mo. In M3 no ridge-plates in u.se; 22-)-
monteus primigenius cranial type. ridge-plates developed out of the typical twenty-four ridge-plates characteristic
of this species.
JAWS OF THE TRUE MAMMONTEUS PRIMIGENIUS B, Adult Mammonteus primigenius jaw (Nat. Mus. 8.579) from Alaska,
We have enumerated above (pp. 1141-1144) the chief obser- showing M3 in situ; seventeen plates in use, five additional plates, total
vations of Falconer, Leith Adams, Pohlig, Soergel, and other twenty-two plates; apparently the two anterior ridge-plates of the typical
number twenty-four have been worn off in this adult jaw.
writers on the jaws of the European specimens of the true Mam-
A, Aged Mammonleus primigenius (Nat. Mus. 4162), from Elephant
monteus primigenius. The characters observed in the xVmerican
Point, Alaska, M3 in situ; seventeen to eighteen plates in use, two additional
specimens are as follows: unworn, nineteen to twenty plates present, apparently four anterior
I)lates
The first jaw (Nat. Mus. 6666 —.shown in Fig. 1011 C) is from ridge-plates worn off out of the maximum twenty-four characteristic of this
Alaska, twenty-five miles north of .\nvik. The second jaw (Nat. species.
1146 OSBORN: THE PROBOSCIDEA

Jaw Characters. —
As in the case of the skull, the jaws of the u des differences, qu'on ne saurait admettre dans une et la meme
true Mammonteus primigenius have doubtless been partly confused espece. Voici le resultat de mes recherches sur beaucoup de dents

by Falconer and by all subsequent writers with those properly be- molaires que j 'ai pu examiner et qui me fait distinguer les especes

longing to Parekphas both in Eurasia and in North America. suivantes:

As shown in figures 962 and 1011 of this Memoir, the true

Mammonleus jaws, with their true Mammonteus inferior grinding 1. Elephas mammonteus, dentibus molaribus rectis, laminis

teeth, are harmonic with the extreme hypsicephaly and bathycephaly numerosis angustis, parum elevatis, anguste fim-

of the cranium. In proportion they are shorter anteroposteriorly, briatis.

ami deeper from the mandibular condyle to the symphysis of the 2. Elephas paniscus, dentibus molaribus rectis, laminis
mandibuli; the rostrum is deeper and more deflected; the section latis elevatis, parum fimbriatis, latere longe distinctis.
of the ramus is more rounded.
3. Elephas periholeles, dentibus molaribus rectis, laminis
These hypsicephalic proportions of the jaw are clearly dis- elevatis, profunde fimbriatis, oblique projectis.
played in the referred aduU jaw of Mammonleus primigenius (Nat.
4. Elephas pygmaeus, dentibus molaribus similibus mam-
.Mus. 8579) from Alaska, which is shown in side view in figure 962C
monteo; sed magnitudine, plus quam dimidio minori-
and in mid-section in figure 1011 Bl; this adult jaw, both in ex-
bus.
ternal aspect (Fig. 962) and in section (Fig. 1011 B, Bl) displays the
deeply depressed rostrum, also the relatively abbreviated third 5. Elephas campylotes, dentibus molaribus subarcuatis,
inferior molar, M3. laminis angustis, numerosis, arcuatis, parum elevatis.

In the young M. primigenius jaw (Nat. Mus. 6666) the

rostmm is less depressed and as a whole is relatively longer and Later in the same year in a Bulletin of the Soci^t^ Imperiale
partakes more of the character of the young jaw of Elephas indicus; des Naturalistes de Moscou, Fischer (1829.2) reviews the above
in fact, all young jaws of Elephas, of Mammonteus, and of Lo.ro- five species to which he adds a sixth (p. 276) under the designation
lionta are relatively elongate, and the deep, hypsicephalic pro- of Elephas Kamenskii, as follows: "D'apr<?s cette maniere de voir,
portion is acquired only in the adult. il faudrait necessairement indiquer comme espece distincte celle,

Fortunately three growth stages of the typical Mammonteus

primigenius jaws aie well displayed (Fig. 1011) in three specimens r,^ r Ti^ jr
in the United States National Museum, as carefully drawn to
a uniform one-eighth scale.

'oMPARisoN OF Mammonteus with Parelephas and Archi-

(

—
DisKODON. FZxactly similar mid-sections of the jaws of Archidis-
kodon and of Parelephas are shown in figures 892 and 893, by
which it appears that the adult jaw- of Mammonteus primigenius
may readily be distinguished from adult jaws of Archidiskodon
and of Parelephas in mid-section; also from the adult jaw of
Elephas indicus (Fig. 893, D, Dl) by the following bathycephalic
characters: (1) Depression of the rostrum; (2) elevation of the
coronoid and of the mandibular condyle; (3) narrowness of the
space between the anterior border of the condyle and the posterior
border of the angle; (4) relative abbreviation and depth of Ms,
which compressed into a much smaller, shorter space antero-
is

posteriorly than that of Parelephas jeffersonii or of Archidiskodon

imperatur. This confirms the statement above, that the jaws and
inferior grinders (jf the true Mammonteus are shorter, deeper, that
is, more bathycejihalic, in proportion, than the jaws of Elephas,
Parelephas, ox Archidiskodon.

Fischer de Waldheim, 1829.1, p. 285.— "C'est ^ Mr. le Baron

Cuvier que nous devons le d^veloppement des caract^res anatom-

iques (jui font distinguer le Mammont ou I'espece fossile d'Elcphant,

de celles qui existent encore. Je me suis porte longtemps avec
I'idre, que j'ai cnoncee dans ma Zoognosie [Footnote: 'Fischer,
Zoognosia tabulis synopticis illustrata. Mosquae. Vol. Ill, p.
320.'], que parmi les ossemens fossiles d'Elc'phant pourraient etre
A Typical Third SiiPEHion Molar of Mammonteus primigenius
cachoes plusieurs especes, que nous confondons dans une seule,
Fig. 1012. Type of Elephas odnntoti/rannus Eichwald, 1835, Pi. Lxiii, figs.
savoir celle du Mammont. La forme de la m&choire inf(?rieure,
1 and 2. One-third natural size. IaiM .superior molar of tlie "right" [left]
surtout celle des dents molaires et de leur lames nous conduisent side, l.M^. From Russia.
 : .

THE MAMMONTIN^: MAMMONTEUS 1147

a la ciuelle a appaitcnue la rnachoire infi'Tieuie tic Sibciie que nous 17-19? rib-bearing dorsals, and 4 lumbars, or 28-30 presacral
devons a S. E. Mr. Bantich-Kamensky." vertebrae].
Adams, in describing this specimen, used the f(uaint measure-
"6. Elephas Kamenskii, molaribus subarcuatis utrinque
ments and language of the time (Adams, 1807, "Relation d'un
attenuatis; laminis paruni elevatis, numerosis, medio
Voyage a la Mer Glaciale et Decouverte des restes d'un Mam-
annulatis."
mouth," Journ. du Nord, St. Petersburg, translated by Sir
History and Synonymy. — Following the early descriptions of Joseph Banks, Philo.sophical Magazine, 1808, Vol. XXIX, p.
Ludolf (1696) and of Ides (1706), the mammoth waited nearly 148): "The i^arts least damaged are a fore foot and a hind one;
a century for the generic name Mammnnleus Camper (1788)' and they are covered with skin, and have still the sole attached.
another decade (1799) for the specific name primigenius Blumen- According to the assertion of the Toungouse chief, the animal had
bach, after which it was fairly deluged with the host of generic and been so large and well fed, that its belly hung down below the
specific names listed above, which rank with those apjjlied to the knee joints. This mammoth is a male, with a long mane at his
MaModoti among the curiosities of scientific literature. Cuvier neck, but it and no trunk. The skin, three-fourths of
has no tail
(1806.1) traces the discovery of fossil elephants back to Theo- which are in my is of a deep gray, and covered with
possession,
phrastus (born 372 B. f'., died 287 B. ('.), a pupil of Aristotle, and a reddish hair and black bristles. The humidity of the soil where
follows {op. with discoveries in (Jreece, Crete, and
cit., ])p. 4, 5) the animal has lain so long, has made the bristles lo.se some part of
North Africa, continuing with a long account of the introduction of their elasticity. The entire carcase, the bones of which I collected
the domestication of the recent elephant in Europe and Africa and upon the spot, is 4 archines and a half high by 7 long, from the tip
closing with a summing up of all the previous fossil discoveries of
elephants of Upper Pliocene and Miocene times in all parts of
Europe and northern Asia, as well as of the Mastodon in America.
In the first and all subsequent editions of the "Ossemens Fossiles"
he recognizes only three species of elephants, living and fossil,
namely, Elephas ajricanui^, Elephas iiidicufi. and Elephas primi-
genius, which he clearly separates by cranial and dental characters,
as cited above (caption to Fig. 992)
—
Adams (1807), Tilesius (1815). Adams described the animal
as the mammoth; he states on page 152 that Blumenbach actually
LOXQDONTA AFRlCANA OXYOTIS 3568nini,.ll'8/z"
called the animal Elephas primsevus. Cuvier assigned the name CENTRAL AFRICA
Elephas manirnonteus to the northern Mammoth (1799). Brandt
named it Elephas brachyrainphus (read 1831, i^ublished 1832).
From the time of Camper (1788) and of Cuvier to the time of
Tilesius (1815) a classical feature of the various descriptions is the
constant use of "mammonteum," "mammonteus," and "mam-
monteo" as the Latin designation of the mammoth, first as an
adjective and then as a specific name, i.e., Elephas nmmmonteus
Cuvier, as compared with Elephas indicus Linnaeus.
Among the outstanding points in Tilesius' 100-page descrip-
tion in Latin of the 'Adams skeleton' are the measurements given
in "pedum anglicum" = about 9 [ inches] and "pollicum anglicum" ELEPHAS INDICUS BENCALENSIS 3200mm.,l0'5"
INDIA BENGAL
[
= about 1 inch]. Tilesius gives the entire length of the skeleton,
from the curvature of the tusks to the tip of the tail, as "viginti
pedum" = about 15 feet in our measurement]. The vertebral
[

column, according to the Adams-Tilesius descriptions of 1807, 1815

(written immediately on Adams' return from the field), includes
28-30 presacrals, that is, from the first cervical to the last lumbar,
as follows

Cervicals 7
Dorsals 17-19? MAHMOHTEUS PRIHIGENIUS 2999mni..9'lO"
Lumbars 4? N. EURASIA
Caudals 8 + Fig. 1013. Restorations by Margrct Flinseli Buba, under the direction
of Henry Fairfield Osborn, of Loxodonta africaiui oiyotis, Eti-phas iiulicus
Only the eight anterior caudals are preserved [see Tilesius' bengalensis, and Mammonteus primigenius. AH figures to a one one-hundredth
figure reproduced herewith (Fig. 1014), which includes 7 cervicals. scale.

'[See footnote on page 1117 above regarding the validity of the genus Mammonteus. — Editor.]
 —

1148 OSBORN: THE PROBOSCIDEA

Fig. 1014. 'Adams skeleton' of Siep/ms prmiffeTMu* Blum, in the Zoological Museum of the Academy of Sciences of the U. S. S. R., Leningrad, Russia.
Reproduced after Tilcsius, 1815, Tab. x. About one twenty-sixth natural size. This was also reproduced by Cuvier in his "Rechcrches sur les Ossemens
Fossiles," Quatriemc Edition, Atlas, 1836, PI. 17 (xi), one-fiftieth natural size. (Cf. Cuvier, op. cit., 1836, Explanation of Plates, p. 9): "Planche 17. E16phans.
PI. XI. Fig. 1. Stjuelette entier d'elephant fossile, rapporte par M. Adams dc la mer Glaciale et copi6 d'apres Tilcsius. L'oreille aba conserve quelques
parties mollcs, et les pieds sont encore couverts de peau, et garnis de leurs semelles."

of the nose to the coccyx [Footnote: 'An archine is a little more Cuvier (op. cit., 1834, Vol. II, pp. 131, 133, 204, 208, 231).—
than two feet English measure.']; without however comprehend- Cuvier describes this famous skeleton as follows: "La deuxieme
ing thetwo horns, each of which is a toise and a half long, and both est celle de I'elephant rapporte a Pc'tersbourg par M. Adams, et
together weigh 10 pouds [Footnote: 'A poud is 40 pounds.']. dont la conservation allait presque jusqu'au merveilleux. Le fait
The head alone weighs eleven pouds and a half." fut annoncp d'abord en octobre 1807, dans le Journal du Nord,

Length of vertebral column 12-14 "pedum anglicum" = 9-10 feet]

Neck short 17 "poUicum anglicum" = 17 inches]
Entire length of skeleton, from forward arch of tusks to tuber-
osities of ischium 20 "pedum anglicum" = 15 feetl
Height of skeleton 4 "archines" and a half = about9ft. 3 in.']
Length of tusks VA "toise" = about 9 feet]

Length of humerus 40 "poUicum anglicum" : about 3 ft. 4 in.]

Length of cubitus or ulna 35 "poUicum anglicum" = about 2 ft. 11 in.]
Length of femur 46/^ "pollicumanglicum" = about 3 ft. WA in.
Length of tibia 28 "pollicum anglicum" = about 2 ft. 4 in.]
Height of scapula 29 "pollicum anglicum" = about 2 ft. 5 in.

'[Lang in his article in Zoologica (1925.1, p. 28) gives a height of 9 ft. 11 in. at the shoulder, as mounted in the Leningrad Museum. Editor.)
 THE MAMMONTIN^: MAMMONTEUS 1149

recueil imprinie a Petersbourg, n° xxx, et ce morceau, qui a i-eparu Measurements

depuis en divers journaux allemands, a cte reimprime en 1815, (Compare op. cit., p. 208)
dans le tome V
des M^moires de I'Academie de Petersbourg. Crane de Mes- Crane du squel-
Nous en tirons les details qui suivent. En 1799, un pecheur serschmidt, ette de M.
Tongouse remarqua sur les bords de la mer Glaciale, pres de Transact, philos., Adams, PI. 17,
rembouehure de la Lena, au milieu des glagons, un bloc informe vol. xl, pi. I. [Fig. 1014]

qu'il ne put reconnaltre. L'annee d'apres il s'aperQut que cette PI. 8, fig. I

masses etait un peu plus degagee, mais ne devinait point encore ce [Fig. 991]

que ce pouvait ^tre. Vers la fin de I'et^ suivant, le flanc tout entier Depuis le sommet jusqu'au bord
de I'animal et une des defenses ^taient distinctement sortis des des alveoles 1,178 1,300

glagons. Ce ne fut que la cinquieme annee que les glaces ayant Depuis le sommet jusqu'aux
condyles occipitaux 0,663 0,770
fondu plus vite que de coutume, cette masse enorme vint echouer
Des condyles aux bords alvt'olaires 0,946
a la cote sur un banc de sable. Au mois de mars 1804, le pecheur
enleva les defenses, dont il se d^fit pour une valeur de cinquante (Cuvier, op. cit., p. 231): "La peau est semblable a celle de

roubles. On ex^cuta, a cette occasion, un dessin grossier de I'elephant vivant, mais on n'y distingue pas les points bruns qu'on

I'animal, dont j'ai une copie que je dois a I'amite de M. Blumenbach. remarque dans I'espece des Indes. M. Adams assure que la peau

L'animal etait male; .ses defenses ctaient longues de plus de dont il avait conserve les trois quarts etait d'un gris fonce. . . .

. . .

en suivant courbures, et sa tete, sans les defenses, [p. 232] M. Adams nous dit qu'une des oreilles de son individu
neuf jiieds les
etait bien conservee et garnie d'une touffe de crins; mais dans son
pesait plus de quatre cents livres. ... [p. 204] Encore plus recem-
c'tat actuel, comme on pent le voir, pi. 17 ab, fig.i, elle est fort
ment j'ai retrouv^ ces longs alveoles dans le crane des bords du
alteree et n'a i)lus Les pieds du squelette de Peters-
aucun poil.
Volga, grave par M. Tilesius, et dont je donne une copie pi. 15,
bourg sont encore converts de peau et garnis de leurs semelles.
fig. 7. II est vrai que ce caractere ne se montre pas sur le grand
M. que ces semelles sont arrondies, et comme dilatees
Tilesius dit
squelette de M. Adams, mais M. Tilesius nous dit expressement et foulees par le poids du corps; en sorte qu'elles remontent sur
que les Tongouses avaient mutile les bords des alveoles lorsqu'ils les bords du pied et les recouvrent. II y avait quelque chose de

en arracherent les defenses, et qu'ensuite M. Adams, pour les y semblable dans I'elephant de la menagerie de Versailles, decrit par
rajuster, fit encore rogner et egaliser ces bords [Footnote: Mem. Perrault. Ni M. Adams, ni M. Tilesius ne nous parlent du nombre
de I'Acad. de Petersb., t. v. (1815), p. 511.]." des ongles."

2. PRIMITIVE EUROPEAN STAGES OF MAMMONTEUS PRIMIGENIUS

As listed above (pp. 1136-1138), European stages of dwarfed dimensions, or supposedly more primitive than
the typical Mammonteus primigenius of late Glacial and Postglacial times, were described in the following order:

1888 Mammonteus primigenius leith-adamsi Pohlig, a dwarfed Sd(?) Interglacial race of Thuringia, Germany.
1891 Mammonteus primigenius hydruntinus Botti, a dwarfed race of the cavern of Cardamone, Otranto, southeastern Italy.
1912 Mammonteus primigenius fraasi Dietrich, an Upper(?) Pleistocene stage of Steinheim on the Murr, central Germany.
1923 Mammonteus primigenius astensis Dep^ret and Mayet, an Upper Pliocene stage of Piedmont, northern Italy.

Between the typical Mammonteus 'primigenius Blum, of northern Germany and the Upper Phocene' M.
primigenius astensis of northern Italy there are doubtless numerous intermediate ascending mutations which only
by very close monographic research can be clearly distinguished from each other. It must be remembered, how-
ever, that Mammonteus was not a permanent resident like the members of the Hesperoloxodon antiquus or Parelephas
trogontherii phyla, but a southerly and northerly migrant during the advances and retreats of the four great
glaciations. Pohlig regards his subspecies leith-adamsi (Fig. 1015) as a diminutive variety of 3d{?) Interglacial time
in the forests of Thuringia, while Botti describes also as a dwarfed subspecies his hydruntinus'- (Fig. 1016) from the

caverns bordering the Otranto Straits, southeastern Italy. Dietrich clearly distinguishes his subspecies fraasi
(Fig. 1017) as a full-sized race, with pentadactyl manus and pes, of Middle [Upper?] Pleistocene time, Uving on the

'[Possibly Lower Pleistocene (see footnote 1 on p. 1049 above). —Editor.]

^Bearing on the presence of in southeastern Italy is the unique note of James Smith (Proo. Geol. Soc, London, 1847,
Mammonteus primigenius hydruntinus
p. .')2): "The fragment which accompanied the specimen was given Mr. Smith by Mr. St. John of Valetta, who stated that he found it, encrusted with
to
stalagmite and adherent to the rock, in the island of Gozo. According to Dr. Falconer, it consists of two plates of a young grinder of the true Elephas primi-
genius. The occurrence of so large an animal in a locality of such limited extent seems to point to a period when it was connected with a continent."
 : —

1150 OSBORN: THE PROBOSCIDEA

borderland between the forests and steppes of central Germany. Of great phyletic significance, if confirmed, is the
very primitive stage A/, primigenius astensis (Fig. 1019) of the Upper Pliocene of northern Italy, described by
Deperet and Mayet, and also observed in the Forest Bed, Lower Pleistocene, of Anglia (Fig. 1020) ; this primitive

M. p. astensis is distinguishable by its thicker enamel and fewer ridge-plates (M 3 Trrrs) ^s compared with M 3 1^
of the typical Upper Pleistocene M. primigenius. The supposed phyletic order of descent of these primitive
species is given above on page 1138; the following are the type descriptions and figures in chronological order.

Mammonteus primigenius leith-adamsi Pohlig, ingia, Dornap (Fuhlrott), Germany. Type FicrnE. Op.
1888 al, p. 229, fig. 101,c, rf.

Figure 1015 Type Description — Op. cit., p. 232: ". . . Noch fremd-
artiger ist ein jiachyganaler Mandibelzahn aus den Steinbriichen
3(1(1) Interglacial Travertines of Thuringia, Dornap (Fuhlrott).

In 1888, Pohlig described this diminutive variety of Mammon-

von Dornap (Fuhlrott), welcher — 15 x in mindestens 0,22 ca. X
branch of the main stem of the true Mam-
0,08 m aufweist; . . . Den vorstehend angefiihrten Belegen der
teus primigenius as a
typischen, der angusticoronaten und der pachyganalen rheinischen
monteus primigenius. It was found in Thuringian deposits of Srf(?)
Mammuthform reihen sich in Folgendem einige hochst bemerkens-
is consequently more ancient than the typical
Interglacial time and
werthe Beispiele der diminutiven Primigeniusvarietdt in den Bonner
.Mammonteus primigenius, which belongs to IV Glacial and
^
Sammlungen an; es mag geeignet .sein, dieses Zwergmammuth
Postglacial times.
durch eine besondere Bezeichnung, etwa E. (primigenius) Leilh-
E. [Elephas] (primigenius) Leith-Adamsi Pohlig, 1888.
Adamsi n.f. von der Stammform zu unterscheiden."
"Dentition und Kranologie de.s Elephas antiquus Falc. mit Bei-
Troue.s.sart (1897) cites Elephas (primigenius) Leith-Adanuv ai^
tragen iiber Elephas primigenius Blum, und Elephas meridionalis
a subspecies of primigenius. His citation is as follows (op. cil.,
Xesti," Nova Acta Leop. Carol., LIII, Nr. 1. pp. 229, 232.
p. 711): "d. —Leith-Adamsi, Pohlig (Var. minor), 1889."
—
Type. Third left inferior molar, I.M3. Original in Bonn.
Pohlig's letter (Sept. 10, 1924) does not authenticate var. minor
loc. cil.,

Horizon and Locality. — Srf(?) Interglacial travertines of Thur-

as a subspecific term. Following the original description of the
species in 1888, Pohlig in 1892 refers again to this species as perhaps
one of the dwarf eleijhants which found its way to the mainland
(op. cit., 1892.1, pp. 260, 261): "Es ergibt sich also der inter-
essante Schluss, dass gleich dem Zwergmammuth, Elephas Leilh-
Adamsi, auch der mediterran insulare Zwerg-Urelephant Elephas
Melilse stellenweise wieder mit dem Festland communiciren konnte;
— dieser jedoch offenbar, nach der grossen Seltenheit seiner Ueber-
reste daselbst zu schliessen, nur auf ganz kurze Zeit."-

Mammonteus primigenius hydruntinus Botti, 1891

Figure 1016

La Grotta Cssifora di Cardamone in Terra d'Otranto, Italy.

This subspecies based upon a twelve ridge-plated fir.st su-
is

perior molar of the left side, l.M' (Fig. 1016); the validity of this
subspecies is dependent upon its geologic age.
The title of Botti's paper is "La Grotta Ossifera di Cardamone
in Terra d'Otranto" and consists of descriptions of fossil bones
found in this cave, including a molar and other jjortions of an
elephant to which he assigned the name E. primigenius Blum,
Ein maxillarer und zwei mandibulare letzte Molaren
Fig. 101.
var. hydruntinus. The si)ecific reference and type description are
—
von Elephas primigenius (in c il son E.pr. Leith-.iddin^i).
as follows
Originale in Bonn (Vo).
E. primigenius Blum. var. hydrunlinus Botti, "La Grotta
Fig. 101.5. Type figure (c,(l) of E. (primigenius) Leilh-Adamsi. .\ftor
O.ssifera di Cardamone in Terra d'Otranto," Boll. Soc. geol. Ital.,
Pohlig, 1888, p. 229, fig. 101.
Vol. IX, 1891, p. 709, Tav. xxvi.

'The type figure (Fig. lOluc, d) and description of the type third left inferior molar do not convince Osborn (1930) that this tooth belongs in the true .Mam-
nmnteu.i primigenius phylum. It appears too long ami narrow, the ridge-i)late.s are too widely separated, and the enamel is too thick. A reexamination of the
type would settle this question.

-.As noted above, the type third left iiifiTior molar of 'I'', (iiriiiiigetiius) Leilh-.idamsi' in the Bonn Museum does not appear to clearly establish its re-

lationship to the Mammonteus phylum.

 THE MAMMONTINJi:: MAMMONTEUS 1151

—
Type Description. (Botti, o-p. cil., p. 705) "Uno di questi,
: lamine anteriori. Di tallone anteriore non scorgo traccia, sebbene
un 1° niolare vero, superiore, sinistro (tav. xxvi, fig. 1, la) presenta una certa depressione, la quale potrebbe dipendere dal contatto del
una corona alta 90""° e la lunghezza di 122'"™; la superficie dente che lo precedeva nella serie."
triturante, restremita posteriore non essendo ancora scoperta, si "[P. 706] L'altro,molto consumato dall'uso, alto 55"'™, (tav.
riduce alia lunghezza di 105 ""», la sua larghezza massima, misurata XXVI, fig. ha la superficie triturante lunga 97"^"', la quale
2, 2a),

nel terzo anteriore 60°""; questa superficie inclina leggermente presenta la maggior larghezza di 46™™ nel terzo posteriore, concava
verso I'interno, ma non e convessa, bensi plana, particolarita che dal davanti all'indietro, leggermente inclinata verso il lato interno.
ricordo avere osservata in tutti gli altri molari superiori di Carda- Ha parimenti dodici lame, manca affatto di radici e ritengo essere
mone, sebbene oppugnata da coloro che vogliono i molari
la quale, I'ultimo molare di latte o premolare, inferiore, sinistro, della stessa
superiori a superficie convessa, concorda coU'assioma dato da Fal- sunnominata specie."
"I dischi resultant! dal consumo delle lamine per efTetto della
masticazione, sono, in ambedue molari, quasi retti e stretti, niente
i

allargati nel centro: le lamine di smalto, leggermente flessuose,

corrono senza notevoli restringimenti, quasi parallele, ma note-
volmente increspate."
"[Pp. 708, 709]. In conclusione, questi piccoli molari di C'ar-
damone, che non trovano riscontro, che io sappia, se non in un
molare (modello) del Museo di Firenze, asserto proveniente da
Roma, ed in quello del Museo di Torino, proveniente dalle alluvioni
del Po, in Italia, e dipoi in due individui del Museo nazionale di

Budapest, hanno un fades affatto ditTerente da tutti gli altri in

''"•'"
Htm

Type op Mammonteus hydruntinus

Fig. 1016. Type l.M' of Elephas iprimigenius Blum. var. hydrunlinm Botti, 1891, Tav. xxvi, figs. 1, la, 2, 2a. From the GrottaCardamone near Otranto,
Italy. Scale (left figure) one-half natural size, height of crown 90 mm., length 122 mm.; (right figure) a milk tooth, height 55 mm., length 97 mm., breadth
46 mm.

coner: 'Constant character of mammoth's molars of all ages and grandissimo numero esistenti nei Musei coUa determinazione di E.
regions: worn surface nearly flat' [Footnote: 'Falconer H. Pakonto- primigenius Blum.; hanno lo smalto pivi fino e piil finamente in-
logical Memoirs and Notes. Vol. II, p. 285.']." crespato, le lame piu serrate e piu avvicinate, le dimension! della
"Ed infatti undici lame in uso, leultimequattro a piccoli dischi meta piu moderate."
non ancora divenuti confluenti, ed una non ancora scoperta mi "Nuove specie furono create con minor divario di forme e I'ele-
fanno ritenere per questo dente la formula di 12 lame, non com- fante di Cardamone differisce certamente dal primigenius tipico
preso il tallone, formula appartenente all'ultimo dente di latte piu di quello che il primigenius differisca dsdVindicus; ma con-
(secondo Owen e Falconer) o premolare (secondo Blainville) ovvero siderando che il molare di Roma e quello delle alluvioni del Po,
ad un antepenultimo (ossia primo) molare vero di E. primigenius non che quell! di Budapest, furono raccolti prima di quell! di Carda-
Blum." mone e nessuno penso a separarl! dalla specie primigenia, e riflet-
"Mi fanno inclinare a quest'ultima opinione la forma massiccia tendo che in sostanza e po! un solo e medesimo piano di struttura
del dente e la presenza di robuste radici, sebbene troncate nel- che si sviluppa, sebbene in grado diverso, nell' E. primigenius
I'esemplare in esame, in specie di una anteriore, isolata dalle suc- Blum, e negl! elefanti di Cardamone, cosi stimo miglior partito di
cessive, che sono fra loro saldate, la quale sostiene le due prime lasciar questi ultimi nella specie primigenia, tutt'al piu proponendo
 8

1152 OSRORN: THE PROBOSCIDEA

di flistingucrli, poicho sono in fiitto ilistinti, in ragionc della dimcn- 13. Lamelle ist nur an dor Lal)ialseite eiitwickelt.'l in 27 cm Liinge,

sione di una meta niinoie e della piu sensibile ristrettezza dei loro 22 cm Hohe [Footnote: '.\n der 1(5. Lamelle.'), 10,4 cm Breite,

elementi, come una varieta, die potrebbe chiamarsi, dalla regione —

,\bkauungsgiad x 17. l^amello. Distanz dei' Laniellen 4
. . . —
di provenienza: mm., Liinge tier —
Lamellen 7 8 mm (an tier labialen Seite), Form
des Lamellenumrisses, nach unten verschmalert, Dicke des
E. primigenius Blum var. hydruntinus." Schmelzes 1, 4 1, 5 — mm
(starke Kniuselung), Typus der Ver-
schmelzungsfigur der Lamellen, median lamellar, latei-al aimuliir.
(P. 106): "Diagnose: Grosse hoch-
"Questa distinzione d'altronde non mi appartiene del tutto."
"Gia fino dal 1873, faceva notare Leith Adams alia Zoological beinige, fiinfzehige Mammutrasse mit-

Society considerevolidiscrepanzenei molari a lame ordinariamente

kurzem schlankem Rumpf.
unil Im
Schiidel und den Molaren mit primiti-
sottili del Mammuth, ed allegava una autorita altamente compe-
M. Davies del British Museum, quale, dall'esame ven ^lerkniiden. Schadel verhaltnis-
tente, quella di il

Elephas primigenius, tro\ati in Inghilterra ed miissig iiiedrig, Molaren weitlaiumellig-

di numerosi molari di
er als hei den jiingsten Mammuten.
alt rove, era da assai tempo venuto nella opinione che esistano due

distinte varieta, facili a riconoscere, i molari dell'una essendo

format i di lame separate da ristretti strati di comento
sottili

interposti, I'altra composta di lame piu dense con intervalli piu

larpchi, soggiungendo quest'ultima forma essere piu frequente

deU'altra [Footnote: 'Leith Adams A. op. rit. p. 6, 7.']."

Mammonteus primigenius fraasi Diefrich, 1912

Figures 1017, lOlS

"Mitteldiluvialen Schottern in Steinheim a.d. Murr," Germany.

This subspecies, named in honor of Eberhard Fraas by Profes-

sor Dietrich in 1912, is based on a type skeleton mounted under the
direction of Professor Fraas in 1911 in the Museum of Stuttgart.

As displayed in the type figure (1912, Taf. i, ii), it appears to

represent a Middle [Upper?] Pleistocene race or subspecies of
Elephns primigenius clearly characterized by its large size, fewer
ridge-plates in lower molars, and five digits in the manus and pes,
probably a typical phalangeal formula.
Elephas primigenius Fraasi Dietrich, 1912. "Elepha.s primi-
genius Fraasi, eine schwabische Mammutrasse." Jahresh. Ver. f.
vaterl. Naturk. in Wiirttemberg, Jahrg. LXVIII, pp. 42-106.
Type. — Skeleton, adult male. Mounted under the direction of
Professor Fraas in the K. Naturalienkabinett, Stuttgart (No.
12837). HouizoN and Locality.— Found in 1910 in "Mittel-
diluvialen Schottern in Steinheim a. d. Murr." Typp: Fig-
URK.— Op. ciL, Taf. i and ii, also text figs. 2, 4, 1 1-14, 16-21, 24-26.

Type Desckiption. — {Op. cit., pp. 10.5, 106) : "E. primigenius

Fraasi, wie ich die neue Rasse Herrn Professor Fraas zu Ehren
neime, ist also keineswegs synonym mit der eben erwiihiiten Be-
zeichnung, worunter letliglich eine Anzahl bestimmter Molaren-
formen zu einer Mutationsreihe zusammengefasst werden. E.
primigenius Fraasi ist vielmehr innerhalb dieser Reihe eine
distinkte, osteologisch voUstiiiidig bekannte Form, in deren Skelett
eine Reihe von Merkmalen vereinigt sind wie es bisher an keinem
anderen Mammut beobachtet ist. Au.sserdem ist der E. primi- Fig. 1017. Tv|)c cranium of the svibspccios Elephas primigenius fraasi

genius Fraasi eine charakteristische Lokalform, denn die Stein- Dictrii-li, 1912, Taf. ii, figs. 1 and 2 (
= Mniiimonleits primigenius fraasi of the
prosont Mcnioir|.
heimer Funde zeigen immer wieder das gleiche gewaltige Ausmass
Tills fraiiiuni, associated witli tlic skeleton now mounted in the K.
der ("dieder." Stuttgart (No. 128:57) illustrates the extreme hypsi-
Naturalienkabinett in

Observe the relatively primitixe structure of M;, (cf. Dietricli, cephaly and aerocephaly, fore-and-aft compression, of the profile of the fronto-
1912, p. 76): occipital region and tlie maikcd concavity of tlic frontals, agreeing clo.sely
with the cranial characters of all species of Mammonteus and differing widely
Mj max. sin. F'ig. 12. Lamellenforinel x 22 x |l'"ootn()to: 'Die from the cranial profile of Parelephas.
 ;

THE MAMMONTIN^: MAMMONTEUS 1153

Carpus uikI Tarsus hochentwickelt, der crste aussen lein serial, even surface which is admirably designed for the comminution of
inncii iKihczu serial. Die Maniichen gehoren zu den Riesenfornicii the finer grasses. The mammoth is thus chiefly a grass-eater,
sic tragen starke. gcbogcnc, abcr mir wpiiig spiial gcdrehte Stoss- a fact which is also attested by the contents of the stomach. It
zahiic, dcreii Wachstuin einer Hemmuiig uuterliegt. Die iStoss- also is an open country animal, but we cannot of course deduce
zahne werden gebraucht." from the structure of its teeth whether this country was warm or
"Typus: Das Skclett No. 12837 (d' adult) iin K. Naturalien- cold, for while it is occasionally found mingled with an arctic
kabinett zu Stuttgart." fauna this is not invariably the case. The lemains at Steinheim
Habits of Elephas [Mammonteus] pkimigenius fkaasi prove that it lived on the borderland between the forests and the
(cf. —
Dietrich, 1912, p. 42). In E. [Mammonteus] primigenius the steppes because it is found in association with E. [Hesperoloxodon]
enamel folds barely rise above the cement, affording a relatively antiquus.

Fig. 1018. Mounted skeleton of Mammonteus primigenius fraasi (Naturalienkabinett, Stuttgart, No. 12837), after photograph kindly forwarded to tlic

present author by Dr. W. O. Dietrich. Compare Dietrich, 1912.1, Taf. i. The skeletal height, from the sole of the foot to the top of the cranium, is at
least 4 m. or 13 ft. 1% in. (.see Dietrich, op. cil., p. 64).
 — —

1154 OSBORN: THE PROBOSCIDEA

Mammonteus primigenius astensis sont celles de M' d'un sujet de petite taillc d'E. primigenius, M'
Depcret and Mayet, 1923 dont la longueur moycnne est, d'apres Pohlig, de 300 millimetres et
Figures 871, 1019, 1020 atteint parfois 385 millimetres. La frequence laminaire est de
pres de 8 lames pour 10 centimetres de couronne ; c'est le chiff re
Upper Pliocene (Villafraiichian stage),' San Paolo tie Villafranca, northern
Italy.
normiddeVE. primigenius. Les bandes d'email sont tres minces,

confirmed by more material, the recognition by Deperet

If
bien plus que chez VE. trogontherii du meme gisement; elles sont
tout aussi onduleuses, mais beaucoup moins plissees, I'^mail etant
and Mayet of an Upper Pliocene stage in the evolution of the genus
presque uni d'un bout a I'autre des bandes, sauf quelques plis
Mammonieus is a step of the utmost importance in the phylogeny
and phylogeny of the genus dans les lames anterieures tres unies. II n'existe aucune trace
of the Proboscidea especially in the
Mammonteus. The type grinders (Fig. 1019) appear to Osborn to du sinus loxodonte."
be relatively narrower than we should expect. If, however, we Falconer, 1868. Vol. II, p. 170. — Falconer described as
may depend upon the highly trained and acute obser\'ations of follows a perfect l.M' "of the pre-glacial variety of Elephas primi-
(
'harles Deperet and Lucien Mayet then Elephas primigenius mut. genius. . . . The matrix is indisputably of the forest-bed of the
nstensis actually belongs in the Mammonieus phylum and is quite Norfolk Coast." Length 11.5 in. (292 mm.), breadth 4.1 in. (104

Fig. 1019. Type and Paratype Grinders of Mammonteus primigenius astensis

(I>cft) Type of Elephas primigenius astensis from San Paolo de Villa- (Right) Paratype of Elephas primigenius astensis. After DepSret and
franca. After Depfirct and Mayet, 1923, PI. xi, fig. .5, p. 221: "Fig. ."j.— Mayet, 1923, 221 "Fig. 6.
PI. xi, fig. 6, p. Elephas primigenius, mut. astensis.
:

Elephas primigenius, mut. astensis de San Paolo de Villafranca. M^ droite. Pliocene fluvio-lacustre de I'Astesan, h San Paolo. Ma droite. (Voir p. 185).
(Voir p. 184). Musoe du Palais Carignan, a Turin, figuree par Zuffardi, 1913, Pi^ce du Mus(5e g6ologique de Turin, n° 15, figuroe par Zuffaidi, 1913, pi. v,
pi. VI, fig. 2a [as Elephas primigenius Blum. var. trogontlwrii Pohl.]." Less than fig. 8a [as Elephas primigenius Blum. var. trogontherii Pohl.l." Less than one-
one-half natural .size; actual length 216 mm., breadth 94 mm., height 186 third natural size; length 280 mm., breadth 84 mm., height 138 mm.; nineteen
mm.; nineteen ridge-plates present. ridge-plates plus talon.

separate from what these authors have named "Ciroui)e des Elephas mm.), height 7 in. (178 mm.); ridge-plates 18-1-, fourteen in a
Trogontherii." space of 7}2 in. (190 mm.); enamel slightly thick. This l.M' is

E. [Eleph(i.'<] primigenius mutatifin nstensis Dejjeret and cither of Mammonteus primigenius or of Parelephas trogontherii
Mayet. "Les Elephants Pliocenes," Deuxieme Partie, 1923, pp. (ef. Fig. 1020).
183, 184. —
Type. Kight third .superior molar, r.M'. Musee du Paratypk. — {Op.
Deperet and Mayet, p. 185): "Les Ms
cit.,

Palais Carignan, a Turin. Hohizon and Locality. San — fig. 8-9) ont une couronne rela-
figurces par Zufiardi (1913, pi. v,
Paolo de Villafranca. —
Type Figuke. Op. ril., PI. xi, fig. 5, tivement etroite et fortement arquce comme dans la derniere
]). 221; figured by Zuffardi, 1913, Tav. xii [vi], fig. 2a, as Elephas molaire inferieure des Elephants. Nous figurons (pi. xi, fig. 6) la

primigenius Blum. var. trogontherii Pohl. dent du cote droit qui est la mieux conservee. La longueur de la
—
Desciuption of Type. {Op. cit., p. 184): "Molaire superi- couronne est de 280 millimetres mesurce en ligne droite pour une
eure M'. —
II existe au Mu.see de Turin les deux M' d'un meme largeur de 84 millimetres et une hauteur de 138 millimetres. On
sujet. Nous figurons (pi. xi, fig. 5) la dent du cote droit qui est la compte 19 lames plus le talon et des traces de quelques lames us^es
mieux conservee. L'animal etait dejii assez age, car la couronne est a la partie anterieure. La frequence laminaire est de 7, 5a 8 comme
tres arasce en avant par la mastication et ne compte plus que 19 chez VE. primigenius. Les caracteres de I'email sont idcntiques
lames prcsentes, il doit en manqucr quclques-imcs, 2 ou 3 jieut-etre. a ceux des molaires prccedentes: email mince, uni et non plisse,
La longueur conservee de la Couronne est dc 216 millimetres et on sans sinus loxodontes."
peut estimer la longueur totale probable a 249 millimetres. La Osborn, 1924: The relatively high ridge formula, M 3 llril,
largeur de la couronne est de 94 millimetres et la hauteur au milieu is what we should expect in an ancestral stage of the phylum Mam-
atteint le chiffre considerable de 186 millimetres. Ces dimensions monteus, which in M . primigenius, at the close of the Pleistocene

'[Possibly Lower Pleistocene (see footnote 1 on p. 1049 above). Editor.)

 :

THE MAMMONTIN^: MAMMONTEUS 1155

age, rises to M 3 M, and finally in M. primigenius coinpressus to genius astensis: M3 1 9-20' This is a much higher ridge formula
M 3 ^^^ From the above very accurate description and admirable than that of the typical Parelephas trogontherii of Pohlig, namely,
figures may be deduced the ridge formula of Mammonteus primi- M 3 y|-|, although the type of P. trogontherii belongs to a more
recent geologic stage than the type of M. primigenius astetisis.
The ridge formula of M. primigenius astensis is thus higher than
MAMMONTEUS PRIMIGENiUS' ASTENSIS
MAMMONTEUS PRIMIGENIUS' ASTENSIS that of any contemporaiy species of mammoth or elephant, much
M2 exceeding the contemporary ridge formulse of species of Archidisko-
M2
don and of Parelephas (see Fig. 1020).

Mammonteus primigenius Forest Bed, Eng-

(?)astensis.
land ((Jsborn, 1922, 1924).— The Cromer Forest Bed of England
includes a sub-arctic faunal phase of the northern latitude of East
Anglia, Lat. 53, during the period of the Scandinavian glacia-
first

tion, which has no parallel in southern France or in northern Italy,

Lat. 44-46. The Forest Bed contains two distinct faunas, namely:
5Qiin Nc 1240 Fcesf Bed
'^^^'^
A.
t^lu3

MERIDiONALIS CROr^ERENSlS I. Survivals. — Upper Pliocene Red and Norwich Crags warm
ARCMIDISKODON MERIDiONALIS CROMERENSIS types which survived in the Forest Bed and belong to the 1st

Interglcaial stage.

II. Arrivals. — Pleistocene cold types which first appear in

the Forest Bed and may belong to the I Glacial stage.

P'rom the studies of Geikie, Prestwich, Clement Keid, and

SaiT ^fu5 A'o /97 Forest Bed from the collections of A. C. Savin, Osborn in 1922 (1922.563, p.
ARCHIDISKOOON 7PLAN1FR0NS 439) summarized these two faunas as given below.

In brackets are indicated the possible references of the Forest

Bed proboscideans to the north Italian species, but to determine
the exact specific and subspecific references of the Forest Bed
proboscideans, namely, "Elephas meridionalis'' "Elephas anti-
/^ivwc/j /'/(is Re J Cra^ NAT. SIZE quus," and "Elephas primigenius" very close examination and
Base of Red Crag Falkenna
comparison will be required
19 28 H. F, O.

(1) The "Elephas primigenius" listed below may prove to

Fig. 1020. Mammonteus primigenius Crowns
{'!)as(ensis (A, B). of correspond in its ridge formula with that of the Elephas primi-
referrpd superior (M-) and inferior (Mo) molars, from the Forest Bed of
genius mut. a.4ensis Depe'ret and Mayet (M 3 rfrls)' of Asti,
Cromer, Norfolk, England, comiiared with (?) Parelephas (C, D), Archidiskodon
northern Italy. (2) It is also possible that the "Elephas antiquus
(E, F), and Archidiskodon or Hesperoloxodon (G, H) molars. All figures one-
fourth natural size, reiwodueed from direct scale tracings by the author.
Nc-^tii type" of the Forest Bed may pro\-e to correspond with

I''or details sec figure 871, Chapter XVI. members of the Parelephas trogontherii phylum.'

FOREST BED, OR CROMERIAN, FAUNA OF EAST ANGLIA

Survivals
I. 11. Arrivals
Warm ITpper Pliocene Red Crag and Norwich Crag Types Cold and northern Pleistocene Types which Fir.d Appear
which Survive in the Forest Bed of Cromer in the Forest Bed of Cromer
Elephas [Archidi.'ikodon] meridionalis [cromerensis?] type Elephas [.Mammonteus] primigenius [
= astensi,s?\
Elephas antiquus Nestii type = Parelephas{'!) trogontherii
[
ue.stii] Equus caballus fossilis
Elephas [Ifcsperolo.rodon] antiquus (typical) Sus scroja
Hysena striata Bison bonasus
Hysena antiqua Caprovis Savinii
Rhinoceros etruscus Ovibos moschatus
Eguus stenotiis Alces latifrons
Cervus carnutorum Capreolus capreolus
Ma^.hserodus sp.? Cervus elaphus
Trogontherium Cuvieri Ursus spelseus
Macacus sp.? Felis spelsea
Rhinoceros megarhinus Varied forest rodent fauna
Hippopotamus amphibiu.s Northern forest small Carnivora

'[After this text was written. Professor Osborn referred 'Elephas' anliguus Neslii Pohlig, cotypcs, to Pareleplias{?) trogontherii nestii (sec Cliap. XVII,
above, p. 1059.— Editor.] .
 .

1156 OSBORN: THE PROBOSCIDEA

3. AMERICAN STAGES OF MAMMONTEUS

In the full description and discussion above of the typical species Mammonleus primigenius, with a ridge-plate

formula of M 3 M, as determined by Falconer, this species appears to range completely across Eurasia into Alaska,
with the same ridge-plate formula. In Alaska also occurs* the much more progressive species Mammonleus
primigenius compressus (M 3 ^^jh)) which ranges southward into Indiana, probably subsequent to the time of the
appearance of the typical M. primigenius. Unfortunately the type of M. primigenius americanus DeKay, Upper
Pleistocene of New York, has been lost and further research is necessary to determine precisely the ridge formula
of this stage, which is estimated in the present Memoir as M 3 If
Owing to the confusion by Hay and others of species of the true Mammonleus with species of Parelephas
jeffersonii, in which the ridge-plate formula ranges from M 3 If to M, much careful research remains to be done in
connection with the revision and determination of the species and subspecies of Mammonleus characteristic of the

northern regions of North America. The tentative arrangement is as follows:

Mammonleus primigenius: M 3 fr, typical of Alaska and ranging southward.

Mammonleus primigenius americanus: M 3 ffe, typical of New York.
Manimontciis primigenius compressus: M 3 ^f^ occurring in Alaska, also ranging southward into Indiana (typical).
So far as ovu' i)resent evidence goes, Mammonteus primigenius reached its final and most progressive stages
of evolution in North America, probably in Postglacial time.

Mammonteus primigenius americanus DeKay, 1842

Figure 1021

L'ppcr Pleistocene, IV Glacial (Wisconsin diluvium), Irondiquoit River,

Monroe County, near Rochester, New York.
Mi. M I .^kt. f

The type of this species was in the Cabinet of the Lyceum of

Natural Histoiy, New York, which was destroyed by fire. The
name was alluded to by I>eidy (1858.2, p. 29), also by Adams,
(
'ope, and Pohlig. As DeKay mentions thirteen plates in five
inches ( = 126 mm.), and also gives an excellent figure, it seems
desirable to retain this subspecific name as Mammonteus primi-
genius americanus DeKay.
E. [Elephas] americanus DeKay, 1842. "Natural History of
New York. Part I. Zoology: Zoology of New York, or the New
York Fauna," 1842, ]). 101. Typk. — Portion of an upper
molar tooth with thirteen ridge-plates. Hoiuzon and
LofALiTV. — Irondiquoit River, Monroe County, near Kochester,
New York. —
Typk Figukk. Op. cil., PI. xxxii, fig. 2. f^Airi SIX/'
—
Typk Descuiption. "It is with some hesitation that 1 ven-
ture to designate, under a new name, a species founded on speci-
mens of teeth, which appear to differ widely from any hitherto met
in this country. The specimens above alluded to were found in
. . .
Type op Mammonteus primigenius americanus
a diluvial formation near the Irondiquoit river in Monroe county, Fig. 1021. Type of Elephas americanus Dc Kay, 1842, PI. x.xxii, fig. 2,
tliree-fiftli.s natural size. an upper molar tooth with thirteen ridge-
I'ortion of
ten miles east of the city of Rochester. According to a wiiter in
platcs. From near the Irondicjuoit River, Monroe County, in the vicinity of
the American .Journal, \'ol. 32, p. 377, these remains consisted of
Rochester, New York. Type formerly in the Cabinet of the Lyceum of Natural
a tusk and two molars, one of which is in the Cabinet of the History, New York, destroyed by fire.

'[See new subspecies Mammonteus primigenius alaskensis from near Fairbanks, Alaska (pp. 1159 to 1161 of the present chapter). — Editor.)
 — A

THE MAMMONTIN^: MAMMONTEUS 1157

Lyceum, and is that figured in the plate. This is six inches in its coner observed (1863, p. 66) that a constant character of the North
greatest depth; and, as nearly as can be conjectured from the American mammoth is that the ridges and their constituent ele-
part which remains, it must have been about eight inches long, and ments are more attenuated and condensed. Thus, one tooth with
three in breadth on its grinding surface, which is, however, too 14 ridges shows an average of .36 in.; another presents 17 discs
much injured to exhibit the ends of the enamel. There are thirteen with an average of .46 in. Taken singly, the difference between the
plates in a space of five inches, and they are more compressed than higher ridge condensation in American as distinguished from
in any fossil species with which I am acquainted, being almost in European specimens seems inconsiderable, but when it extends
contact, with very little interstitial substance. It is altogether over a length of crown comprising 16 or 24 ridges, it is perceptible
different from any fossil elephant hitherto described, and merits the at a glance. It givesa certain amount of distinctive physiognomy
distinct appellation of E. americanus." to the molars of the North American mammoth. Falconer, how-
Falconer (1863). Elephas primigenius [americanus]. Fal- ever, does not )-egard this as indicating more than a slight geo-
graphical variety, as the other characters remain constant to the
true mammoth type.
Osborn, 1924: Falconer's observation that a constant char-
acter of the North American mammoth is that the ridges and their
Amer. Mus. 14559
(outer view) constituent elements are more attenuated and condensed conforms
with De Kay's type description that "there are thirteen plates in

a space of five inches, and they are more compressed than in any
fossil species with which I am acquainted, being almost in contact,
with very little interstitial Assuming that De Kay's
substance."
measurement of thirteen plates in a space of five inches was ac-
curately made, we find that the ridge-plate compression of the type
of Mammonteus primigenius americanus is as follows: 10-|- ridge-

plates in 100 mm. This would relate this subspecies more closely
to the typical Mammonteus primigenius than to M. primigenius
compressus, but while De Kay's specific name may be retained, we
can hardly consider that this can be raised to a higher rank than
a subspecies, namely, Mammonteus primigenius americanus.

Mammonteus primigenius compressus Osborn, 1924

Figures 806, 819, 1022-1024

Type from Rochester, Indiana; paratype from Alaska. Probably of the

Postglacial or retreat period,IV (Wisconsin) Glacial.
This subspecies is typified by the female skull (Amer. Mus.
14559), from Rochester, Indiana, and the paratype from Alaska
(Amer. Mus. 13749), fully figured and described in this Memoir.
It is readily distinguished from the typical Mammonteus primi-
genius of Eurasia (M 3 M) by the very high compression of its

dental ridge-plates, namely, M 3 j^. The original description by

Osborn (1924.633) is as follows:
Mammonteus primigenius compressus Osborn, 1924. "Par-
elephas in Relation to Phyla and Genera of the Family Elephanti-
dse." Amer. Mus. Novitates No. 152, Dec. 20, 1924, pp. 5-7.

Mammonteus primigenius comprbssus. Type. From Indiana Type. —Skull of a female mammoth including both tusks, also

Fig. 1022. Type second and third superior molars of female Mammonteus superior grinding teeth. Horizon and Locality. — Roches-
primigenius compressus (Amer. Mus. 145.59), from Indiana, one-fourth natural ter, Indiana; Upper Pleistocene of Alaska and of the Central
size. The teeth (Al) belong on the right side of the beautiful female skull also United States. Type Figure.— Op. cii., 1924.633, fig. 2, p.
represented in figure 1023. After Osborn, 1924.633, p.
as Ekphas primigenius, see Osborn, 1922.555, p. 8,
6, fig.

fig.
2 (originally figured
8). Ridge formula: 6; and figure 1023 of the present Memoir. Paratype. —
third right superior molar, r.M', from Alaska (Amer. Mus.

A External view showing twenty-seven ridge-plates of M^.

13749). Paratype Figure.— See Osborn, 1922.555, p. 7,
1,
fig. 7 (Fig. 1024 of the present Memoir).
A, Crown view showing M' with eight ridge-plates in use, closely com-
Type Description (Osborn, 1924.633, pp. 5-7).— The origi-
pressed to M" with eight ridge-plates in use.
1158 OSBORN: THE PROBOSCIDEA

nal description of this subspecies is in pait as follows: "(1) Ex- typical Mammonteus primigenius of Eurasia and of North America
treme fore-and-aft compression and vertical elevation (hypsiceph- displays a very constant ridge formula, as especially observed by
aly, bathycephaly), correlated with extreme hj'psodonty and Falconer, namely:
forc-and-iift compression of the twenty-seven lidge-plates which
compose iVl'. (2) Measurement across outside of orbits, 262 mm.;
Dp 2i Dp S^ Dp 4H \l 1 if M 2 U M3 2
rc
1

from top of skull to bottom of premaxillaries, 393 mm." Falcouer himself remarked that while American specmiens of the
"After careful and prolonged examination of the specimens true E. primigenius display a similar formula, namely, M 3 fr,
and the descriptions of Falconer (1863), we conclude that the the grinding teeth in general show more closely compacted ridges."

Mammonteus puimioiinius compukssiis. Type Female Skull kuom Indiana

Fit;. 1023. Tvpr .skiiU of fcmiilc mammotli (MnininonkiiK priiitigrniiis comi>rcssus), .Vmcr. Mu.s. 145.")9, from RochestiT, Indiana, acriuircd by punlia.M' in
I'J'Jt. Sec occipitofrontal section of same skull (Fig. 806); alsotcctli of same skull (Fig. 1022). This skull is tlic ly/jc of the species Mammoiilcnx primigenius
rompreasus Osborn. One-twelfth natural size.
Observe in the three aspects of the .skull, frontal, jialatal, and lateral, the following inijiortant points: (1) The extreme fore-and-aft compression (cyrto-
eephaly, hypsicephaly or acrocephaly) correlated with the extreme comi)res.sion of the twenty-seven ridge-plates which compose M' and are displayed in detail
in figure 1022; (2) the slemler parallel female tusk insertions and deep extension of the incisive alveolar plates; (3) this is the most extreme fore-and-aft com-
pression of the skviU, grinding teeth, and alveolar processes thus far recorded; (4) the measurement across outside of orbits is 262 mm.; from top of skull to
bottom of premaxillaries 393 mm.
The cranium of this type, iis beautifully displayed in the figure at the right, exhibits the utmost extreme of vertical elevation (hypsicephaly) of the o(^eiput
and of vertical depression (bathycephaly) of the grinding teeth. In harmony with the almost straight vertical line of the forehead and of the condylar pro-
cesses are the maxilhe and the sharply downturned tu.sks. The apex of the skull, half complete, would terminate in a sharp ])eak (acrocephaly). This harmonious
fore-and aft compres.-^ion and deepening of the entire cranium, which brings the o<'cipit.al condyles relatively close to the narcs, apparently exhibits he ab.solute-I

ly finest stage in the evolution of the Mammonteus cranium and dentition. Certainly this remarkable foreshortening, doubtless initiated in the .1/. primi-
genius aslcnsis of the Upper Pliocene [Lower Pleistocene?), could go no farther and we may regard this species as the final stage in the evolution of he phyhun t

Mammonteus.
 THE MAMMONTIN^: MAMMONTEUS 1159

"It is owing to the excessively high compression (hypsodonty) Remains of Trilophodont-Tetrabelodont Mastodons," Bull. Amer.
and multiphcation of the plates (polydiskodonty), amounting to Mus. Nat. Hist., LIX, pp. 506, 632. Cotypes.— Four crania
M 3 (7)^71 that the new subspecific name Mammonteus primigenius (infantile, —
with deciduous dentition, A. C. F: A. M. 26991; ju-
compressus is now defined and illustrated by figine 2 [Fig. 1022 of venile, with deciduous dentition, A. C.—F: A. M. 26990; adult
the present Memoir]." female, with left maxillary tusk, A. C.—F: A. M. 26989; and
The type and paratype superior grinding teeth are clearly aged male, Alaska College). Horizon and Locality. "Bone —
illustrated in figures 1022 and 1024 of the present Memoir, and the pits" scattered between several widely separated stripping opera-
cranial characters of the type are also shown in figure 1023, a female tions of the U. S. Smelting and Mining Company, vicinity of
cranium in which the summit of the occiput is unfortunately Fairbanks and the Tanana River, Alaska. Pleistocene. Cotype
broken away. We do not at present know of any jaw in which the —
Figures. Figures 1025 and 1026 of the present Memoir. Referred
grinders display this extreme stage of compression. Consequently figure, see Frick, 1933.1, PI.
12A.
the ridge formula of the lower teeth is not certainly known. The Fkick, 1933.1, p. 632.— " Elephas primigenius alaskensis Oshorn.
ridge formula of M3 is probably (^faV- Among the twenty-six tusks secured of the Northern Mammoth
E. primigenius there is one of unusualsize, the same weighing in the neighborhood
A2 Aftier. Mus. 13749 Het.
'inner view' of 300 pounds. The average number of laminae in the normal m'
is twenty-five and the length of the crown in use varies between
157-207 mm. A mandible of an aged cow is remarkable in that the
lastmolars have been extruded and the alveoli healed over.
Another mandible exhibits a somewhat produced symphysis."

/ -'

• 5 s > e 9

E. primigenius
Amer. Mus 13748 Ret

Mammonteus primigenius compre.ssus. P.*.ratype. From Alaska

Fig. 1024. Paratype figure of right third superior molar, r.M^, of Mammon-
teus primigenius compressus (Amer. Mus. 13749), from "Historic Bluff,"
Eschscholtz Bay, Alaska, showing the maximum compression of the ridge-
plates; ridge-formula 3 ^^M One-fourth natural size.
. After Osborn,
1922.555, p. 7, fig. 7.

Mammonteus primigenius alaskensis sp. no\'.'

Figures 1025, 1026

From vicinity of Fairbanks, Alaska. Pleistocene.

[In anticipation of Professor Osborn's description of a series of crania
of the northern Mammoth, discovered in 1929 near Fairbanks, Alaska,
Mr. Childs Frick, who generously accorded Professor Osborn the privilege
of first description, u.sed (1933.1, pp. 506, 632) the manuscript name
Elephas primigenius alaskensis. Mr. Frick not only listed the probos-
cidean material collected in the Fairbanks region by the joint Alaska
College-Flick American Museum parties, but he figured (PI. 12A) a
palatal specimen (A. C. —
F: A. M. 27010) which he referred to this
subspecies.
While it was Professor Osborn's intention to describe in Novitates
this new ascending mutational stage, prior to the article by Mr. Frick in
the Bulletin of the American Museum of Natural Historj', the text was
never completed; in order, however, that such portion of it as he had
prepared may be recorded, we are inserting it herewith, including certain
essential additions. — Editor. |

Elephas primigenius alaskensis Osborn (in Frick, 1933). "New

1160 OSBORN: THE PROBOSCIDEA

"List of Matehial p'uom the NiciNnv ok Faiubanks, Alaska 2 complete and 12 partial radii (largest 61 cm., smallest
26 tusks (largest, on curve 12 ft. 10 in., hiisc circumference 56 cm.)
24 in.) 8 complete and 8 partial fenuirs (largest 113 cm., smallest
Large bull skull with nv's and tusk (9 ft. fS in. x 18.5 in.), 96.5 cm.)
Alaska College Collection 12 complete and 14 partial tibiae (largest 68 cm., smallest
Large cow skull with m's and tusk (6 ft. 5 in. x 13 in.) 44 cm.)
4 smaller skulls, including 2 of calves 3 complete and 2 partial fibulae

6 partial palates 37 metapodials

Some 24 mandibles or paitial mandibles 17 astragali
204 detached molais or partial molars 16 calcanea
31 partial scapulae 60 carpal or tarsal bones
7 complete and 5 partial humeri (largest 98 cm., smallest 6 patellae
75 cm.) Nmnerous vertebrae
16 complete and 5 partial ulna> Several ribs, partial ilia"

CirrvrE Chanh op Mammonteus primioknius alaskensis

-Ml ono-twciitioth natural sizi:

FIr. 1026. Coty|)(! crania ofElcphas [^rammonteus] primiqniiii^ nlaskcnsis: (I) Infantile (.\. C.-F:A. M. 26991); juvenile (.\.C. F:A.M. 26990); adult
female (.\.C.-F:.\.M. 20989); aged male (Alaska College, through the eourteisy of President Charles E. Bunnell). Compare figure 1023 above.
 THE MAMMONTINiE: MAMMONTEUS 1161

ASSOCIATED FAUNA In 1924 Osborn described (1924.633, p. 5) the most progressive

(cf. Frick, 1930.1, p. 79): form Mammonteus primigenius compressus: M
3 ^f^. Recent
Pre-tundra Fauna of Alaska-Yukon explorations in Alaska have revealed to President Charles E.
Bunnell of the Alaska Agricultural College and School of Mines
Large herds of the super-bison, Bison crassicornis Rich, ref.
and to Mr. Childs Frick a more primitive a.scending mutation to
A new Alaskan lion, Felis atrox alaskensis, Frick
which the name Mammonteus primigenius alaskensis may be
A new wolf, Aenocyon dims alaskensis, Frick
assigned.
An arctothere, Arctodus yukonensts Lambe, ref.

A camel, Camelops (?)

The Alaska College and the American Museum are indebted
to the ITnited States Smelting and Mining Company for the
Three species of musk-oxen, Ovibos, Symhos lyrelli O.sgood,
privilege of making a very careful survey of the fo.ssils revealed
and Bootherium sargenti didley
during the recent stripping operations of the company on Cold
A horse, Equus alaskx Hay, ref.
A mastodon. Mastodon americanus Kerr, ref. Stieam and Clery Creek near Fairbanks; the survey was conduct-
A Mammoth, Mammonteus primigenius alaskejisis ed under the supeivision of Mr. Peter Ktiisen during the seasons of
The association of these mammals points to a Pleistocene 1929 and 1930. On this material Fiick (1930.1, p. 73) reports as

phase which Frick inclines to compare with the Sheridan fauna of follows: ". . . the jfiint Museum and College party succeeded in
the Aftonian or 1st I nterglacial stage of Nebraska. harxesting during the four summer months some twenty-eight
—
Osborn's Manuscript (1931). The chief result of intensive large cases of skulls, jaws, and bones —rare and important evidence
researches on the genus Mammonteus is to establish in the e\o- on the prehistory of Alaska which otherwise would have been lost
lution of the genus a number of successive ascending mutations to science. The great percentage of this material, interestingly
ancestral to, or descending from, a typical 'Elephas primigenins enough, came from three restricted areas, 'bone pits,' scattered
Blum.' (1799) of Siberia and north Germany to which Falconer between se\'eral widely separated operations of the Company, the
(1863.1, p. 65) rightly attributed a very constant ridge formula,
remainder of the worked areas being, for the purposes of the bone
namely:
Qp gi Dp3l Dp 4 i| M H M 2^ M 3 ^
1 hunter, nearly barren."

TABLE XV. Cranial and Dental Measurements

1162 OSBORN: THE PROBOSCIDEA

4. THE FROZEN MAMMOTH OF SIBERIA

Recent studies by Herbert Lang (1925) and by I. P. Tolmachoff (1928, 1929) add greatly to our knowledge
of the habits, structure, distribution, and causes of extinction of the mammoth and form a fitting conclusion to the
present chapter.

The 'Adams mammoth,' located by a Tungusian fisherman in 1799 on the banks of the Lena River, served
Tilesius (1815) as the basis of the first description of a complete skeleton, measuring 9 ft. 3 in. at the shoulder.'
This was followed by successive discoveries, described by Lang and in greater detail by Tolmachoff (1929).

The distinctive external characteristics besides those already mentioned above include its much shorter and
more massive body, its large bulky head, the relatively small size of the trunk, the closely spiraled tusks, the

small ears, the small size of the feet, the two-fingered tip of the trunk, the long fan-shaped, bristled tassel of the
tail, the layer of fat three or four inches in thickness on the under side of tlie belly, and the winter layer of fat on
the concave face of the cranium.

70
 —

THE MAMMONTINiE: MAMMONTEUS 1163

SUMMARY OF THE DISCOVERY AND NATURAL HISTORY OF THE WOOLLY MAMMOTH

Compare Osborn's "The Romance of the Woolly Mammoth" in Natural History, Vol. XXX, No. 3, 1930, May-June, pp. 227-241
Since 1907, when the explorations and researches for the present Memoir were begun, our knowledge of the
classification and palaeontology of the woolly mammoth has not only been greatly extended by exploration and
discovery but greatly modified by intensive and very laborious research. Especially surprising, perhaps, is the
evidence presented for the first time in this Memoir that Mammonteus is probably a branch of the same primary
stock as that which gave rise to the intermediate mammoth Parelephas and to the southern mammoth Archi-
diskodon. This subfamily interrelationship may not be absolutely demonstrated, but the evidence for it seems
to be very strong in the unique structure of the tusks, which are entirely relieved in later years of the mechanical
function of procuring food, with a corresponding feeble structure of the backbone. If this relationship between
the Mammontines proves to be the true one, it is certainly a paradox that the grinding teeth of the relatively
primitive Archidiskodon are so widely separated in character from those of the most liighly specialized Mam-
monteus, while ParelepJias is intermediate.

A recent popular summary of these contrasts (cf. Osborn, 1930.824) may appropriately conclude our previous
systematic and formal presentation of this subject.

The woolly mammoth i.s the classic of palaeontology; it is the sides of these animals, just as during the million-year Stone Age
first extinct mammal to he found by man ; it is the first to be used they contrasted with the hairy and woolly coxering of the mam-
as proof of a imiversal deluge; it is the first to be used as proof of moth of the North.
the existence of a long extinct world of mammalian life antecedent Upon close observation, however, a very important difference
to the tleluge; it is the first to receive a scientific description in the between all the mammoths and the living African and Indian
Latin language; it is the first to receive a scientific name Elephas elephants is thus revealed in the curvature and uses of the tusks,
primigenius or 'the first of all, or original, elephant.' It took namely, the tusks of the mammoth, while emerging from the
nearly two centuries thus to baptize the woolly mammoth as the upper jaw closely side by side, soon begin to spread apart anil then,
fossil mammalian world and to usher in the
honored primate of the slowly rotating on their axes, turn inwards toward each other to
elaborate and intricate science of mammalian palseontology finally form a huge ivoiy circle with tips actually crossing.
through which we now decipher the prehistory of the earth for the A similar ivory circle around the large proboscis is also ob-
past thirty million years almost as clearly as if we were able to served in the lelated extinct elephants to which for this and other
project ourselves back into these long corridors of time. reasons the name 'mammoth' may be applied. The Jeffersonian
The romance of the woolly mammoth stretches back foi- mammoth, Parelephas jejfersonii, was probably a hairy type lack-
—
thousands perhaps for hundreds of thousands of years to the — ing the heavy undercoating of wool of its northern relative, which
time when the men of the Old Stone Age discovered that the ivory roamed from the north temperate region to about N. Lat. 40°
of the tusk of the mammoth and of other elephants was superior over Fjance through Geiinany to the United States. The imperial
to bone for several utilitarian purposes, as well as for the expression msimmoth, Archidiskodon imperator [Fig. 1030] was probably hair-
of man's sculptural instinct. At all times, from the very beginning less, with great incurved ivory tusks that attained a gigantic size

of the Age of Man or close of the Pliocene period when, on the in its antecedents of the more southerly latitudes of Eiu'ope and
authority of Charles Deperet, its ancestors first appeared in the Asia. As shown on the map below, these three mammoths — the
warm forests of Italy, the woolly mammoth has kept its beautiful, 'woolly,' the 'Jeffersonian,' the 'imperial' — formed three great
brightly shining tusks sharpened at the tips, both for prowess in mammoth belts around the Northern Hemisphere which were as
combat mastery of the female herd, and to further the
for the sharply demarcated as are the reindeer, the moose, and the stag, or
efforts of the young mothers to ward off the many enemies that wapiti, belts today.
might surround and attack the baby mammoths straying for We know that Stone Age man hunted the bone of the elephant
a moment from the sheltering maternal side. It is only when the and mastodon for a million years, for we find that the Piltdown
old bull mammoths retire from their paternal duties to a quiet, Man of Upper Tertiary time fashioned one of the long bones of the
bucolic life on the northern tundras that the ivory tusks begin to mastodon for a tool; it is not improbable that he tried to fashion
curl inward, even to the crossing point, thus rendering them the ivoiy tusks as well, although we have no proof of this until
useless both for piu'poses of combat and for herd protection. This ivory carvings are discovered toward the close of the Old Stone
is true also of the Jeffersonian [Parelephas] and the imperial Age. Certainly ivory has been treasured by man for thousands of
[Archidiskodon] mammoths of North America. years. Primitive traders carried ivory from point to point. It is
Similar tusks glisten today in the African and Indian elephant, not surprising, therefore, that man knew about ivory long before
their beautiful 'ivory-white' contrasting with the somber gray he knew about its source.
 : .

1164 OSBORN: THE PROBOSCIDEA

Thus, again, in the early and wide-spread quest of ivory by discovered the ivory tusks by digging in the earth, the word
man for utihtarian and artistic purposes it is not siu'prising that the 'mammoth' is certainly derived from the alleged Tatar word
word 'ivory' is the actual soince of the scientific name Elephas, mamma, signifying 'earth.' As early as 1696 this was combined
a Greek term of vague linguistic origin, subsequently Latinized. with another Tatar word kost, signifying 'ivory,' and the two words
Homer used this Greek term not in leference to the elephant itself were Latinized by Ludolf into Mammotovoi Kost. The buried
but to its tusks or 'ivorv.' With both Homer and Hesiod the tusks were sometimes mistaken for horns; Cuvier alludes to these
words of Tatar origin as follows: "C'est sous
le nom de comes de rnammont, mam7nontova-kost,
qu'ils designent les Thus in the
defenses."
Latinization of the Tatar word mamma into
the Gallic mammont there was assigned by
Camper in 1788 the name Mammonteus, which
is now applied by Osborn to the woolly mam-

moth as a genus distinguished in many external

and internal characters from the genus Elephas
which is typified by the elephant of India.
Thus, as cited in full above (p. 1124) from
Ides, originated the word 'mammoth,' variously
spelled in different languages mammot, mamant
(Russian), mammouth (French), mammuth
(German), from the Tatar mamma because the
remains of these animals were found imbedded
in the earth, the nati\'es therefore belie\-ing
that the animals burrowed like moles. Un-
doubtedly the woolly mammoth was in its time
the colossus of living mammals of northern
or western Europe; in consequence the word
'mammoth' has become a convenient adjective
Geographic Distribution of Mammoths in Upper Pliocene and Pleistocene Times to signify gigantic, immense, of great com-
Fig. 1028. Partly theoretic geographic distribution (1930) of the southern, intermediate, and parative As a matter of fact, the woolly
size.
northern mammoths. See figure 795 for distribution map of 1938. mammoth is a dwarf in comparison with its
Southern mammoths: Archidiskodon planifrons, A. meridionalis, A. imperator. lelatives, the Jeffersonian and imperial ele-
Intermediate mammoths: Parelcphas trogantherii, P. jeffersonii.
phants, as is shown in the illustrations by
Northern mammoths: Mammonteus primigenius, M. primigenius compressus, M. primi-
genius alaskensis. Knight which have been brought to a rela-
Relatives of the southern imperial mammoth, Archidiskodon imperator, are now known to have tively uniform scale (Figs. 1029, 1030, 1034).
ranged from South Africa to India and from Nebraska to Mexico. [Since this was written, a speci- Long before the elephant was known in
men referable to Archidiskodon cf. planifrons has been described from Shansi, China, by Dr. Arthur Europe, Phoenician traders brought ivory from
Tindell Hopwood (1935.1, pp. 87-90), who states as follows: "It is clear that there a wide gap in
is the South to Greece; in the North ivory was
the known distribution of the genus, and that the forms from the Old and New Worlds are separated also procured in the tusks, the average weight
by most Hitherto no species of Archidiskodon has been described from the Far East, and
of Asia.
the sijccimen described below is the first evidence that this gap may be expected to close with the of which was 288 pounds, of the more or less
—
march of knowledge." Editor.] The temperate Trogontherian and Jeffersonian mammoths of the fresh or partly frozen bodies of the woolly mam-
genus Parelephas ranged along the 40th parallel from southern France to central United States. The moths. This northern source of the primitive
northern mammotli, of the genus Mammonteus, ranged about the Arctic Circle from western Europe
ivory trade was summarized bj' Herbert Lang
to eastern North America.
(1925); it has been thoroughly studied in re-
cent years by Basset Digby (1926), Pfizenmayer
word Elephas means ivoiy. Aristotle (384-322 B.C.) in his (1926), and Tohnachoff (1929). In Tolmachoff's recent and
History of Animals' used e\i(i)as in its modern generic sense as comprehensive memoir (1929) is found a map (Fig. 1027 above)
applying to the elephant of India, in referring to its courage in and a list of not less than thirty-nine localities where frozen car-
combat casses have been found, beginning with Ysbrant Ides (1692) and
". Elephants also fight fiercely with each other, and strike
. .
ending with Andrews' discovery of 1923. The author gives due
with their tusks; the conquered submits entirely, and cannot credit to the works of Basset Digby and of Pfizenmayer. He also
endure the voice of the victor: and elephants differ nuich in the gives a most detailed and interesting history of the progress of
courage they exhibit." discovery of the woolly mammoth, Elephas primigenius, and of
The word 'mammoth' has a similar non-classic and indirect its woolly companion, Rhinoceros antiquitatis, the 'rhinoceros of
origin. In allusion to the fact that the Tatars of Siberia first antiquity.' The export ivory industry of Siberia, dating back

'Translation by Richard Cresswell, London, 1887, p. 234

 THE MAMMONTINiE: MAMMONTEUS 1165

to very ancient times, furnishes a very good idea of tlie immense an extinct elephant extended well over a whole century; in
number of mammoths that have been discovered by exploring par- 1696 the Russian explorer Ludolf described the mammoth of
ties in the frozen ground of Siberia, estimated by Middendorf (1885) Siberia under the name Mammotovoi koxt; two years later the
at 20,000 during the past two centuries and by Nordenskiold Cierman scholar Tentzelius defended against all sceptics the
(1882) at a very much higher figure. The
highest estimate is 46,750 for the last two and '^-"

a half centuries, although one estimate even

puts the number at 250 specimens annually,
a total of 62,500 for the two hundred fifty
^
:ii^

years. The northern ivory trade to China is

now traced back as far as 500 B.C. Doubt-
less the \alue of northern ivory rose as the
Chinese gradually exterminated their own
native or imported breeds of elephants from
India.
Herbert Lang (op. cit., 1925) notes that
Ides, famous Dutch traveler and am-
the
bassador to China, seems to have been the
earliest to gather first-hand information re-

garding the frozen Siberian mammoths.' In

traversing northern Siberia between the years
1692 and 1695, Ides learned that many of
the Yakuts, Timguses, and Ostyaks stead-
fastly believed that these huge monsters spent
their lives deep underground, moving about
easily in though the
spacious tunnels even
earth was thoroughly frozen. Should they
become particularly active, the whole ground Imperial Mammoth of Nebraska. About one fifty-fifth Natural Size

might rise above them, caving in later as they Fig. 1030. This superb animal, first named Elephas imperator by Leidy, stands fully 13 feet at

passed on but should they come to the surface

;
the shoulder — 4 feet taller than the woolly mammoth but with similar curved tusks and sloping hind
([uarters. The best skeleton is in the Nebraska State Museum {Archidiskodon imperator tnaibeni);
and breathe the warm air, they instantly died. the finest cranium is in the Geological Institute of the City of Mexico (Fig. 902 of present Mem-
It seems as if the very gradual recog- oir). The animal ranged from Nebraska to Mexico. Like the Indian elephant, it ijrobably had
nition of the woolly mammoth as actually little hair and no wool.
discovery of a really fossil elephant at Burg-Tonna near
Gotha, but this classic skeleton has recently proved to belong
not to the mammoth family but to the straight-tusked
elephant family known as the 'elephant of antiquity'; in
1728 Sir Hans Sloane confirmed the finding of Tentzelius
by reference to the discovery; seven years later (1735)
John Phil. Breyne wrote to the Royal Society as follows:
"... I was busied ... to prove, that the extraordinary
large Teeth and Bones found under Groimd, and digged
up in several Places of Siberia, by the name of Mammoth's
or Mammut's, Teeth and Bones, were.

I. True Bones and Teeth of some large Animals

once living; and.

IT That those Animals were Elephants, by the Analo-

gy of the Teeth and Bones, with the known
ones of Elephants.

Jeffersonian Mammoth of Indiana. About one fortv-fipth Natur.4L Size III. That they were brought and left there bj' the

Fig. 1029. The Jeffersonian mammoth was probably hairy, with a fine universal Deluge. I made likewise several
undercoating of wool in the winter season. Its tusks are incurved exactly as in useful Inferences about this matter."
the remotely related woolly and imperial species. Its concave forehead is
quite distinct from that of the modern elephant. As found in Indiana, it
The first finely etched figure of the cranium of the mam-
measured 10)^ feet at the shoulder. moth brought by Messerschmidt from Siberia was published
'[See also Witsen, Nicolaes, 1692.— Editor.]
1166 OSBORN: THE PROBOSCIDEA

bj- Doctor Breyne in 1737 [1741?]. Thus step Ijy step the way County near Rochester, New York. Finally, in 1924, the present
was prepared for Bhimenbach to name the animal in 1799 antl for author described his Mammonteus primigeyiius compressus, the
the great Ciivier to thoroughly describe it in a series of memoirs most highly specialized and perfected mammoth thus far found, as

CuiCULAR TdSKS of THE WoOLLY M AMMOTH

Arched Tusks of the African Elephant OF Siberia
Fig. 1031. Giant arched tusks of tho African Fig. Contrast
1033. these circle-shaped
picphant, 11 feet in length, presented by Charles
tusks, 8 feet in length, of the male woolly mam-
D. Barney to the Heads and Horn.s Collection
moth of Siberia witli the arched tusks of the
of the New York Zoological Park.
African elephant shown in oiiposite figure.

known both near Rochester, Indiana, and in Alaska (see p. 1157

between 1796 and 18.34. Since that time our knowledge has
above).
advanced by leaps and bounds. The first bones of the mastodon
Beginning with the discovery of a complete frozen mammoth
found on the banks of the Hudson River in 1705 and of the Ohio
Uivcr in 1739 were naturally confused in pAirope with those of the
skeleton — known as the Adams' mammoth — nejir the mouth of the
mammoth when Blumenbach first labeled them Ohio incog iiiiuni rr
and then Mnminnt. A century later, in 1842, DeKay described his
Elephas americamis, a fo.ssil mammoth tooth founfl in Monroe

Woolly Mammoth, Somme River, France

Onk-fiftieth Nawhal Size
Fig. 1034. Thismammoth is a little more than 9 feet at the shoulder,
Skcll a.vd Tusks of the Woolly Ma.vimoth
wherea.s the .Icffcrsonian mammoth is 10)^ feet and the imiicrial mammoth
Fig. 1032. and half-grown tusk.s of a middle-aged male specimen of
Skull 13 feet. The painting closely follows the drawings in the cavern of Combarelles
Mamtnonleus primigeniun found on the Yukon River, .\laska. With advancing (p. 1169, tailpiece). The rapidly sloping hind quarters of the animal serve as
age the.se tusks would curve inward and cross in the mi(l<llc line a watershed for rain and sleet.
 THE MAMMONTINiE: MAMMONTEUS 1167

Lena River in 1799, other complete skeletons were unearthed and mammoth are given very fully because they enhance the reputation
mounted in the museums of St. Petersburg, Leipsie, Stuttgart, and of the Old Stone Age artists not only as close observers but as
Brussels, all closely similar in size, in the wheel-like curvature of the portrayers with marvelous fidelity of the external appearance of
great tusks, in the extraordinary foreshortening of the skull the woolly mammoth. Among the very numerous etchings,
(p. 1158 above), in the rapid falling away or sloping of the hind drawings, and paintings there is one which possesses high artistic
quarters, in the shortness of the backbone and of the tail, in the merit as truthfully depicting the characteristics of the charging
reduction or complete loss of one of the digits of the hand — briefly, mammoth (p. 1132 above). The majority of these etchings and
in the truly marvelous adaptation, in every part of the skeleton sculptures, as fully enmnerated' in MacCurdy's encyclopedic
as well as in the teeth, to the very severe conditions of boreal life. volumes, are of the modern comic supplement order; they give the
In 1912 Fehx described a mammoth found near Borna, whose impression that our Stone Age ancestors were struck only with the
death must have been so sudden that the animal did not have time humorous side of the mammoth as he appeared in the full panoply
to swallow a mouthful of food which lay in the form of a wad of his winter coat. But our increasing knowledge of these animals,
between the upper and lower teeth in the stomach of this animal
; derived from study of the frozen skeletons of Siberia and from the
were about twenty-four pounds of undigested plant food, exceed- extremely close studies made by the present author in the pre-
ingly interesting because it consisted of plants that are still native paration of this memoir on the Proboscidea, assures us that even the
to the place (Beresowka River, northern Siberia), the tundra crude outlines on the walls of the cavern of Combarelles (p. 1169
flora which the mammoths stored up during the short summer tailpiece) are not of the comic supplement order but are very truth-
season for the long winter. These plants are almost exclusively ful portrayals of the woolly mammoth as he actually appeared dur-
grasses and form the characteristic meadow flora; the needles of ing the winter season, rounded out with his huge thickness of fat
conifers occur very rarely. From other discoveries we know that (three and a half inches in places) over the entire body and with
the mammoth fed during the winter on the arctic willow, Sali.r a woolly covering extending down to his very hoofs and masking
polaris of Wahlenberg, and on other northerly dwarf plants. the muscular outlines of the limbs beneath. His extremely elevated
The description by Felix not only gives us complete knowledge head was followed by a deep nick or indenture of the neck, then by
of the skeleton of Elephas primigenius but shows conclusively the the rising hump of the back which sloped rapidly downward into the
proper position of the tusk in the jaw and also its inclination. As to depressed region of the pelvis and terminated suddenly in a short,
the animal's outward appearance, it involved a number of cor- blunt tail.

rections in even the best reconstructions that hail been made the ; Thus the apparent caricatures of the Stone Age artists are
proportion of the length of skull and trunk in the mammoth is realities; the mammoth in his winter pelage was even more dis-
quite different from that of existing elephants: in the mammoth guised than the yak of Tibet. The sloping hind quarters served
the skull is more than half the trunk length, in the elephant {E. admirably as a watershed for the torrents of sleet and rain and the
indicus) it is always less than half. The mammoth's head, there- whirlwinds of snow that raged during the northern blizzards. Not
fore, was higher in proportion to the body than that of recent only this, but the apparently weak and sloping hind quarters were
elephants and in consequence the tusks could attain enormous all that the mammoth needed for the forward propulsion of his

proportions. The largest of the tusks in the Leningrad Museum body, since, unlike all modern elephants, he never used his tusks
measures no less than 13 ft. 8 in. and in the Franzens-Museum of for digging or uprooting purposes; consequently the hind limbs
Briinn there is a tusk that actually exceeds 16 ft. 5 in. in length, were not propellers of the body as they are in the African and
including the long basal portion of the tusk which was incased Indian elephants of today. Another important feature is the
within the exceptionally long tube-like premaxillary bones shown bulbous or well-rounded forehead, which rises like a sloping dome
in our figure (p. 1158) of the female mammoth skull of Indiana, the at the top of the otherwise pointed head; this swollen forehead,
type of Osborn's M. primigenius cojnpressus. we are sure, was a food reservoir for the winter season which dis-
The trunk of the mammoth was extremely well developed. appeared as winter advanced into spring, when the deep fatty
The ear was somewhat smaller than that of the Indian elephant, covering all over the body was exhausted and the animal began to
being about 15 in. in length and 6.7 in. in breadth, and, like the assume the normal outlines and proportions of other elephants.
rest of the body, was covered with a thick coat consisting of short This interpretation is to our mind absolutely demonstrated by
wool and longer hair. The tail is conical in form, about 14 in. the contrast between the comic outlines of the Combarelles mam-
broad at the root, sharply pointed at the end where it terminates moths of midwinter and the wonderfully spirited charging mam-
in a bunch of bristles. The skin was extraordinarily thick and moth of midsummer engraved on a section of mammoth ivory
underneath it was a layer of fat up to 3/2 in. deep. The whole body tusk discovered beneath the Magdalenian shelter of La Madeleine
was thickly covered with fine soft hair about an inch long, varying along the Vezere River (p. 1 132 above). Here we see that the round
in color from faded blond to yellow brown; coarser and longer fatty outlines of winter give place to those which conform very
hair, sometinaes 20 in. in length, of a dark, rust-colored brown, closely to the lean profile of the summer season; the high, peaked
covered the entire neck and trunk, perhaps forming a fringe of hair skull especially, with its characteristic concave forehead, affords us
still heavie.r and thicker from the cheeks along the shoulders and a view of the actual summer profile of the mammoth which corre-
sides to the rump. sponds with that of the Indiana mammoth shown on page 1158.
The above details regarding the external appearance of the The classic Magdalenian engraving is one of the most realistic

'George Grant MacCurdy: Human Origins. Vol. I, 1924, p. 265.'

 1168 OSBORN: THE PROBOSCIDEA

pieces of Palaeolithic art that has ever been found; there are the cavern of Aurignac and hence known as man of the Aui ignacian
indications that the artist iisefl the relatively small piece of ivory stone culture. This race is known as the Brimn or Pfedmost;
for the representation of three mammoths, for the tusks and trunks they are long-headed, with a narrow, short face anil rather promi-
of two other elephants appear in the distance. Observe especially nent brow ridges, and with brain development inferioi- to the
the outline of the ear, the elevation of the
highly peaked, acrocephalic head, and the
remarkably lifelike action of the limbs and
body.
Toward the close of the Old Stone Age
there began the wide-spread custom of the
ceremonial burial of the dead to which we
owe our really remarkable knowledge of the
great hunting races that swarmed o\'er cen-
tral and western Europe during closing glacial Giant Killing Stone
The Mammoth Pit OF THE Moravian
time, succeeding the last of the Neanderthal oi-

Vestonice, Moravia Hunters

race which seems to have had full possession
Fig. 1036. View of
Fig. 10.37. Doctor
of the river shores and caverns at the height mammoth pit of Ab.solon interprets tlic
the
of the fourth or last descent of the Scandinavian hxrge stones found in
V'fstonioe. Mammoth
t he cave as the weapons
glaciers. Largely known by their flint imple- skeletoiLS are strewn
with which the mam-
over tlie floor. (Photo-
ments foimd in the burials of hunters and moths were killed after
graph by Doctor Ab-
warriors of Neanderthal time, we know little solon.)
being trap|)ed. Photo-
graph by Doctor .4b-
of their prowess; the Neanderthal flint imple-
solon.
ments are relatively small and clumsy and it
broad-faced, artistic Cro-Magnons who drew and painted the
would appear that the Neanderthal hunters were not equipped
mammoth. The Briinn people were apparently more interested in
to pursue the large and formidable woolly mammoths of their
eating the mammoth than in depicting it, but they have left some
time.
sculptures in bone and ivory of both animal and human figures.
Succeeding the Neanderthals, however, was another race,
This was one phase of the Briinn Aurignacian culture, but their
probably from the Far East of central Asia, first found buried in
culture is more largely known by their stone implements adapted
to the making of clothing and to the killing of smaller kinds of
animals — none of the Aurignacian implements so far found in

western Europe was adapted to the chase of larger game. Evidence

of the killing and consumption for food of a great number of the
wild horses of the period has recently been found near Solutr6 in
southern France, but, so far as I know, without the accompaniment
of very large flint implements of the chase.
The discovery of giant killing flints by Dr. Karel Ab.solon of

StHATA of THE Pekarna Cave, Moravia

Fig. New and Old .Stone Age .strata, from liistoric time downward
103.').

to Lower Aurignacian— altogether ten distinct layers, representing a |>eriod

which Doctor Absolon believes ext<!nded over 100,000 years. The mammoth Ivory Figurine from Brassempouy
layeris contemporaneous with the Lower Aurignacian. (.\ftcr photograph Fig. 1038. Back and profile views of a woman's head earved on ivory.
by Doctor Absolon.) (After ,1. Pilloy in Edouard Piette's L'AH pendant L'Age du Renne, 1907.)
 THE MAMMONTIN^: MAMMONTEUS 1169

the University of Prague and the Museum of Briinn, one of the entered the continent of America but reached even a higher point
most distinguished archieologists of central Europe, is therefore of specialization in his essential grinding tooth mechanism, just as
a revelation not only of the pursuit of the woolly mammoth for we Americans pride ourselves today on our mechanical achieve-
purposes of food, but of the killing methods employed, whereby the ments.
This grinding tooth is a perfect marvel of
adaptation and it is lodged below the very
highly peaked cranium with a correspondingly
deep depression and foreshortening of the jaw
and approximation of the front and back
planes of the skull which are so extreme as
to appear almost artificial. It is this high,
narrow, and deep skull to which the Greek ad-
jectives hypsicephaly, acrocephaly and bathy-
cephaly especially apply as absolutely unique
in the animal kingdom [See Fig. 1023 above];
in adaptation to grassy diet it is the extreme
and relatively flatheaded
antithesis to the long
cranium of the American mastodon which was
the contemporaneous forest dweller of Alaska
and as far south as Florida.
The woolly mammoth, however, like the
jjolar bear, arctic fox, ptarmigan, and arctic
reindeer, was at its best when in the severe
climate of the far North, defying with its
Eqtjine Ivory Statuette from Lourdes
woolly covering the coldest arctic blasts.
Fig. 1039. This .statuette, found in the Grottc dcs E.spcluguos at Lourdes, is carved from
mammoth ivory. Twice natural size. (After J. Pilloy in Edouard Piette's L'Arl pendant L'Aqc During the summer season it lost not a
du Renne, 1907). moment's time in laying in its grassy hoaid
for the coming winter, in comparison with its
animals were driven into great pits and then felled by giant stones less hardy relative, the Jeffersonian mammoth of the mid- tem-
let down by These stones appear (Fig. 1037) like
their captors. perate region, and its subtropical and more remote relative, the
greatly magnified coups de poings of the long bygone Chellean and imperial mammoth of the south temperate zone.
Acheulean age. Citations from the description of his remarkable Like many other fossil mammals the mammoths appear to have
discoveries in the years 1924-1929 of mammoth-hunting stations become rather suddenly extinct after the climax of the Old Stone
of Moravia are given on page 1139 above. Age, namelj', during the slow northward recession of the final
The woolly mammoth appears to have ranged almost ex- great glaciers of Scandinavia and North America. The cause of
clusively north of the 40th parallel; it loved the borders of the their disappearance at the very moment when they reached the
retreating glaciers of the close of the Glacial Age both in Europe highest degree of specialization and perfection of their grinding
and North America. In Alaska it was extremely abundant and teeth (Fig. 1022) is a mystery; it is attributed by Tolmachoff to
was occasionally foimd in frozen form along the ancient shores of further specialization and Ijy Howorth to the universal flooding
Eschscholtz Bay. Alaska, broadly connected with the Asiatic which accompanied the sidlen northward retreat of the great
mainland by a great isthmus bounded on the south by the present glaciers. A more likely explanation is that during this unfavorable
mountain peaks of the Aleutian Islands, yields abundant remains period the herds may ha\-e become numerically reduced by under-
not only of the true woolly mammoth closely similar to the E. feeding through lack of grassy food diu'ing the summer season.
primigenius of Blumenbach, but also an extremely rare and highly During these decades the underfed mothers were probably unable
specialized grinder (Fig. 1022) to which the specific terminal cow- to protect their yovmg from the attacks of wolves and other
pressus was recently applied by Osborn in descriptive relation to carnivorous mammals. Fortimately for us, during the height of
the exclusively compressed and fine-plated grinding teeth which their supremacy in western Europe they had been superbly drawn,
attain the very high number of 27 compressed ridge-plates above modeled, and painted by the artists of the closing period of the
and below, in comparison with the 24 ridge-plates above and below Old Stone Age. In fact, it was not very long after the recognition
in E. primigeyiius of Siberia and western Europe. This proves that of the woolly mammoth as a true fossil that man was also discover-
the marvelous adaptability of the mammoth did not cease when he ed in the fossil state.
 Fig. 1040. A young adult bull elephant {I^xodonla africana) in the bush of the Lake Paradise region, etist Central Africa, as photographed by Mr. and
Mrs. Martin Johnson in 1923-1924, and shown in the film "Simba." Reproduced through the courtesy of Mr. Daniel E. Pomeroy.

1170
 Chapter XIX
THE GENERA LOXODONTA, PAL/EOLOXODON, AND HESPEROLOXODON
OF THE SUPERFAMILY ELEPHANTOIDEA. SUBFAMILY LOXODONTIN/E
LoXODONTA, PAL^OLOXODON, AND HESPEROLOXODON, UNITED IN THE SUBFAMILY LOXODONTIN.E
BY SIMILAR CRANIAL AND DENTAL CHARACTERS. SIMILAR ESPECIALLY IN THE BROAD PREMAXILLARY
ROSTRUM AND THE WIDELY SPREADING SUPERIOR INCISIVE TUSKS WITH UPROOTING FUNCTIONS TO THE
END OF LIFE. AlSO RELATIVELY SIMPLE GRINDING TEETH AND MORE OR LESS PRONOUNCED MESIAL
'lOXODONT sinus' ADAPTED TO BROWSING RATHER THAN TO GRAZING. Of EXCLUSIVE AFRICAN, MEDITER-
RANEAN, AND EURASIATIC DISTRIBUTION. SURVIVING IN THE EXISTING LOXODONTA AFRICANA.

I. Classification and Histokv ok Discovery of the V. Les Elephants Nains des Iles Mediterraneennes et
LOXODONTIN.E. LA Que.stion des Isthmes Pleistocenes (Vau-
1. Difficulties of generic nomenclature. FREY, 1929).
2. History of discovery and separation of European,
Indian, Mediterranean, African, Japanese, and VI. Ancestral Stages of Pal.eoloxodon in Africa.
Javanese species. Palceoloxodon alia nlic us.
3. Order of discovery and description of the fifty-three Palspoloxodon jolensis.
type species of the extinct Loxodontinse. Palxoloxodon revki.

II. Systematic Revision of the Loxodontin^.

Subfamily characters. VII. Pal^oloxodon and Loxodonta of South Africa.
Lo.rodnnta: Cieneric characters. ^Palxoloxodon andrewsi.
1. Order of description of eighteen living African Palavloxodon hanekomi.
species and subspecies. Palit'oloxodon yorki.
2. Systematic description of species of Loxodonta: Palspoloxodon wilniani.
Loxodonta africana, L. rornaliae. Palspoloxodon kuhai.
Palseoloxodon archidiskodontoides.
III. EuRASiATic Species of Pal^oloxodon and Hesperoloxo- Palxoloxodon transvaalensis.
DON. Palseoloxodon sheppardi.
Palieoloxodon: Generic characters. Loxodonta zulu.
Palxoloxodon namadicus. Loxodonta. prima.
Hes-peroloxodon: Generic characters. Loxodonta africana vm'. obliqua.
Hesperoloxodon nntiqu us. Loxodonta subantiqua.
Upnor elephant (//. antiquus)
Hesperoloxodon antiquus nanus.
VIII. Loxodontines of .Japan and Java.
Ilesperolo.rodon antiquus platyrhynrhus.
Palseoloxodon namadicus naumanni.
He.'^perolo.rodon antiquus ausonius.
Palaroloxodon namadicus namadi.
Hesperoloxodon antiquus germanicus of Rumania.
Palseoloxodon protomauimontcus.
Hesperoloxodon antiquus italicus.
Hesperoloxodon antiquus germanicus of Steinheim.
Pa Ispolo.rodon loku naga i.
Palseoloxodon protoniammontcus proxinius.
IV. Extinct Dwarfed Species of the Mediterranean Palseoloxodon namadicus yabei.
Islands. PalsPolo.rodon tok u naga i junior mut.
Paheoloxodon melitensis. Parelephas protomammouteus matsurnotoi.
Palseoloxodonfalconeri. Palseolo.Todon i/okohama n us.
Palscoloxodon mnaidriensis. Pals'oloxodon hysudrindicus, Java.
Pakeolo.rodon la marmorae.
Palseoloxodon Cypriotes. IX. Summary of Geographic Distribution along the
Palscoloxodon creticus. Eastern Coast of Asia.

The cranial, dental, and incisive tusk resemblances of members of the Loxodontinae to each other and the
profoimd distinctions in cranial profiles, sections, and proportions from the Mammontinse (Archidiskodon,
Parelephas, Mammonteus) types are fully set forth in Chapter XV and synoptically in figure 1041, which illustrates
the profile and frontal aspects of the cliief specimens known of Palxoloxodon namadicus and Hesperoloxodon
antiquus in comparison with Loxodonta africana. The typical species of Palseoloxodon exhibits the broadly diver-
1171
 SKULLS OF LOXODONTS FROM ASIA, EUROPE, AND AFRICA
Alt fiawras oAe-t<*r«nu«in nAtwrai 9*i*

L AFRICANUS R*f. L AFRICANUS Ref.

L AFAlCANUS Rat.
F«l€..IM7. Pt. XUV. F19. XVII UvJ Faic . 1&47. PI. XIII A. F.g. 8
Fate . *B*T. PL XLII. Fie. XVM

L **fTK}uus n«>. I ANTIOUUS

WM»>«(«r, <«90. T»i 11. F.« 2 Pot»>e. <9S*. 9 3)0. F>« )0»

Fig. 1041. Crania of Loxodonta africana, Paueoloxodon n'am.u>icus, and Hesperoloxodon antiques
All to the same Assembled in the year 1922. Compare figure 1009 of the year 1929
scale, one-twontictli natural size.

Upper Row. Loxodonta africana, after Falconer and Cautley, 18J7, Pis. xui, xliv, xiu.a. •

skulls of P. namadicus in troni

f *

Second Row. PaUoloxodon namadicus Falconer and Cautley, type, 1S47, Pis. xii.a, xii.b, front and side views. Two referred
and side views, after Falconer and Cautley, Pis. xxiv..\, xui, xliv. .

of P. namadicus, after PUgnm, iwo.

Third Row. Referred skulls of FaUoloxodon mdUefisis, aft<^ Pohlig, 1893, front and side views, island of SieUy, and
front and side views. .

Fourth Row. (Left) Referred rostrum of Ilcsprroloxodon antiquiu ausonius?, after Weithofer, 1890.
(Middle and right) Rostriun of H, auhquus ausonius
erroneously referred by Pohlig (1891, p. 350, fig. 109) to Elephas {antiquus) Xeslii. Recorded as of Upper PUoeene age.

1172
 THE LOXODONTIN^: CLASSIFICATION AND HISTORY OF DISCOVERY 117.3

gent premaxillary rostrum similar to that of Loxodonta africana but differs very widely in the broadly rugose
parietofrontal crest which extends like a Phrygian cap down over the frontals almost to the nasals ; this is seen
even in the dwarfed species L. {antiquns) melitx^ [
= Palseoloxodon melitensis]. It is obvious that in Palseoloxodon
the cranium somewhat more hypsicephaUc and bathycephalic than in Loxodonta, but this cannot be shown in
is

longitudinal section at present. The function of the gigantic rugose Phrygian cap is doubtless for the support and
retraction of enormous divergent tusks and a greatly broadened and enlarged proboscis.

U elephant h crane allonge ^ a front concave, it tres-

longues alveoles des defenses, h mdchoire inferietire ob-

tuse, a mdcltelieres plus larges, paralliles, marquees de
ruhans plus serres, que nous nommons elephant f ssilc ( ele-
phas primigenius , Blumenb. )
est le mammont des Ru^ses.
,

B Fi^.2. Fi^.s-I.
L' elephant a crane allonge , a front concave , a petites
oreUles , ii mdchelieres marr/ue'es de rubans ondoyans que
nous appelons elephant des Indes ( elephas indicus ) , est un
quadiupede qu'on n'a observe d'une maniere certaine qii'au-
dcla de I'lndus.

C Fip. J. /}>. m A.
U ^Uphant a
crane arrondi, it larges oreilles ii mdche- ,

lieresmarquees de losanges sur leur couronne, que nous

appelons elephant dAfrique ( elephas africanus ) est un ,

quadrupede dont la seule palrie connue est jusqu'a present

I'Afriqne.

Fig. 1042. Cuvier'.s figures and definitions of Elephas primigenius (Messerschmidt's cranium, cf. Fig. 991), E. indicus, and E. africanus. After Cuvier

1806.1, PI. 39 (ii), figs. 1, 2, 3, and PI. 41 (iv), figs. 9.1, lO.A, and U.F. One thirty-seventh to one-twentieth natural size. See also figure 992 above.

I. CLASSIFICATION AND HISTORY OF DISCOVERY OF THE LOXODONTIN^

(Continued from Chapter II, p. 32, of Vol. I, and Chapter XV of the present Volume)
As repeatedly observed in the pre.sent Memoir (e.g., Chap. I, p. 5, Chap. II, pp. 17, 18), it is impractica-
ble in palaeontology to apply all the principles of nomenclature established in zoology and botany, because the
classification of the imperfectly known fossil forms is ever changing with our increasing knowledge of origins,
adaptive radiations, and phyletic successions. Such mutability does not disturb the nomenclature of living animals

and plants, in which priority of adequate description, figure, and definition is the chief concern of systematists.

Zoological principles of classification, as in the classic works of Cuvier and of Falconer, must therefore sur-

render to phylogenetic principles of classification names and classifications of the dawn

(cf. Chap. II, pp. 5-13) and
period of palaeontology must be reworded and rearranged on modern principles and knowledge (cf. Chap. XV,
pp. 911, 912). Even certain technical rules of nomenclature, of orthography, and of priority, as revised by
zoologists,- must give way to palaeontologic standards; otherwise we find ourselves repeatedly compelled to

'(The lettering in figure 1041 was executed at a time when all these species were united in the single genus Loxodonta, that is, before Professor Osborn had
adopted Palsoloxodon Matsumoto (1924) for members of the namadicus group and had created the new genus Hesperoloxodon for certain other species. Editor. ] —
"The International Commission on Zoological Nomenclature, Congress held at Monaco in 1910.
 :

1174 OSBORN: THE PROBOSCIDEA

sacrifice the spirit to the letter and to bury as nomina nuda the classic terms of vertebrate palaeontology proposed
by such great founders as Blumenbach,' Cuvier, and Falconer. In no subfamily is this historic recognition and
common-sense spirit more essential than in the Loxodontinae, as shown in the following revision.

1. DIFFICULTIES OF GENERIC NOMENCLATURE

The generic nomenclature of the Loxodontinae is full of confusion, which can only be cleared up by a common-
sense interpretation and reexamination of the original definitions and types. The history of nomenclature is as
follows:

LoxoDONTE, F. Cuvier. — Frederic Cuvier (1825) was the first to apply the appropriate term "Loxodonte"
(signifying slanthig toothed) to distinguish the African from the Indian elephants, but despite the profound differ-

ences in dental and cranial characters and in geographic distribution, the generic term Elephas continued to be
widely used as late as 1886, e.g., Lydekker, 'Elephas africanus,' and even in 1928, Andrews, Forster Cooper, and
Bather adhered to the generic term Elephas in describing the Upnor elephant as 'Elephas antiquus.'

In the case of the term "Mastodonte" of G. Cuvier, 1806, it was not until 1817 that G. Cuvier adopted the
Linnsean classic system of generic orthography: (1) Accordingly F. Cuvier's original term of 1825 was "Loxo-
donte," as G. Cuvier's original term of 1806 was "Mastodonte."" (2) In an unsigned review of F. Cuvier's
description in the Zoological Journal of London (1827, 1828, Vol. Ill, p. 140), the generic term Loxodonta was sub-
stituted for his "Loxodonte" of 1825; therefore the generic term Loxodonta should be credited to F. Cuvier as the
generic term Mastodon should be credited to G. Cuvier. (3) In 1843, p. 184, Gray, in his "List of the Specimens
of Mammalia in the Collection of the British Museum," employed the Latin spelling, as appears from the following
citation

Tlic African Elephant. J.o.xodonta Africana. Elephas Africanus, BLumciib. Abhild. t. 19. f. c. E. maximvis, part, Linn.
Mam. Lithog. t. Harris, W. A. Afrit: t. 22. Cimer, Oss. Fos. 1. 1. 2 to 8.

Again in 1869, p. 359, in his "Catalogue of Carnivorous, Pachydermatous, and Edentate Mammalia in the
British Museum," Gray employs the same spelling:

2. Loxodonta.

Lamina of the teeth with lozenge-shaped erown. Skull subglobular, forehead shelving, erowii rounded; fiont of lower jaw
aeutc, produced. Trunk (conical, thick at the base. Ears very large.
Loxodonta, F. Cuvier, Denis Mamm. [182.5,1827.]

Loxodonta africana. (African Elephant.) B.M.

Loxodonta africana, Oray, lAst Mamm. B.M 1843; Gerrard, Cat. Bones B.M.
.

Elephas africanus, Blumenh. Abbild. t. 19. f.c; Kirk, F.Z.S. 1864, p. 654; Gicbel, Sdugelh. p. 1.59; Blainr. Osleogr.
Cravigrades, t. 3 (skull), t. 7 & 9 (teeth).
Elopha.s maximus (part.), Linn.

(4) Falconer and Cautley ("Fauna Antiqua Sivalensis," 1847, PI. xlii) and Falconer (Quart. Journ. Geo!.
Soc. London, 1857, pp. 315, 318) employed the shorter masculine term Loxodon [preoccupied], which Falconer
(1857, p. 315) defined as follows:

For this subgcneric group, the name of Loxodon [Footnote: 'From Ao|6j obliquits, and 68ovs dens, having reference to the
rhomb-shaped discs of the worn molars; an adaptation of the term 'Lo.rudonln' projxjsed by red. Cuvier, Hist. Natur(>!le do.s
I"'

Mammiferes, tom iii., "Article i;i6phant d'Afrique." 1835 [1825].'], first indicated by Frederick Cuvier, has been adopted. It
comprise.s both extinct and living species.

'Blumenbarh'.s original description (1799, p. 697) of Elephas primigenius rites as an example the Burgtonna (Gotha) skeleton, which certainly belongs
to 'Elephas antiquus,' ax fully explained in Chapter XVIII above; see also the present chapter (p. 1 181 below).

"Compare Chap. V, p. 119, Chap. VI, pp. 135, 165, and Chap. XXI, pp. 1363 and 1372, for the history of the term Mastodonte.
 THE LOXODONTIN^: CLASSIFICATION AND HISTORY OF DISCOVERY 1175

This most appropriate term, however, is preoccupied by Loxodon Muller and Henle, 1841, for a genus of
sharks, e.g., Loxodon macrorhinus; consequently Osborn adheres to the orthography Loxodonta (F. Cuvier,
1825, 1827; Gray, 1843), although it involves the feminine termination of the species referable to this great genus.
(5) Lydekker (Ency. Brit. 11th Ed.) never admitted the generic distinction of the genotypic and other species
of African elephants, but adhered to Elephas africanus. (6) Heller and Roosevelt (1914, p. 739) erroneously em-
ployed the trinomial Loxodonta africana africana, as listed under the existing African species below.

Elasmodon Falc. (Preoccupied) and Euelephas Falc. (Nomen nudum). (1) In 1846, Falconer and —
Cautley first named (p. 45), and figured (1847,P1s.xii.a, xii.B)a newUpper SiwaUk species as 'Elephas namadicus,'

the specific name referring to the Nerbudda [or Narbada] River, the Namadus River of Ptolemy. (2) A year later
(Fauna Antiqua Sivalensis, 1847, caption to PI. xlii, figs, xix-xxiv), they proposed the new subgeneric term
Elasmodon [in allusion to the waving enamel ridges], to include the six species E. antiquus, E. hysudricus, E.
meridionalis [not figured], E. namadicus, E. indicus (Dauntela var., Mukna var., and young), and E. primigenius;
these six heterogeneous species are the 'genotypes' of Elasmodon and of its substitute Euelephas. (3) In this same
caption they erroneously applied the generic name Loxodon to E. planijrons, grouping it with E. africanus. (4)
In 1857, learning that the name Elasmodus had been used for a series of fossil fish. Falconer (p. 315) substituted
the generic term Euelephas for Elasmodon: "For this subgeneric group we propose the term of Euelephas [Foot-
note, 'From ev bene, and eXe^ay, having reference to the typical Elephants most familiarly known.']." (5) W. L.
Sclater (1900, p. 317) was in error in making Elephas planijrons the genotypic species of Euelephas, because this
species was not included in the original definition of Elasmodon referred to above. (6) Palmer (1904, p. 275)

Third (Inferior) and Second Superior molars of the African (Loxodonta) and Asiatic (Elephas) Elephants.

Fig. 88.

African Elephant. \ iiat. eizc. Asiatic Elephant.

Fig. 1044. Crown view second right superior molar of Loxodonta
of: [K)
Fig. 1043. Crown view of the third inferior molar of the right side of: Fig. africana, with eight lozenge-shaped ridge-plates; (B) second right (?) superior
88, Loxodonta africana; Fig. 89, Ekphas indicus, one-third natural size. molar of Elephas indicus, exliibiting eleven worn ridge plates. After Lydekker,
Compare Owen, "HLstory of British Fossil Mammals, and Birds," 1846, pp. "Catalogue of the Ungulate Mammals in the British Museum (Natural
230-232, figs. 88, 89. History)," Vol. V, 1916, p. 80, fig. 22.
 Crania or AruicAN (I-ki't) and ok Nekbudda, Inuia (Riuht), Elephants

Both oiio-oighth natural sizi-

I/OXODONTA (Ma1-k) I'al.koloxoddn (1''kmali;)

I'Ik- 10)5. Craiiiuin ami jaws of the typical Afrii'aii cli'pliaiit of llic I'ig. lOlb. ('raniiiiii of tlic typ<' of l'nl:ii>lii.rii(lini niiiiiiiilicux. In tlie palato
Suiiaiicsr or Abyssiiimii subspci-ics l.oxodonlii afrianin oiiiotix. In tlic jaw observe the closely parallel and relatively eonipresseil molar ridge-plates cliar-
observe the extremely lozeiiRe-shaped or eeiitrally expaiulinK molar ridne- aeteristie of all species of /'(i/a«(yj(«/o/i,. Compare ligurc 1070, also figure 1072.
platcs cliaraet(!ristie of all speeies of the genus Loxodonla. Compare figure 1013.

1170
 :

THE LOXODONTINiE: CLASSIFICATION AND HISTORY OF DISCOVERY 1177

simply fiuoted Sclater's erroneous statement. (7) Falconer in his Memoir of 1857 (p. 318) defined in Latin the
collective genus Elephas (Linn.), on the basis of premolar and molar tooth structure, as embracing three subgenera,
namely

Subgen. 1. Stegodon.— Dcntium molarium 3 utrinque intermediorum coronLs complicata hypisomoris colliculis {e.g.
7 + 7+8), mammillatis, tectiformibus. Prsemolares nondum observati.
Subgen. 2. Loxodon. — Dent, molar. 3 utrinque intermedior. coronLs lamello.sa hypisomeris 7+7+colliculis (e.g. 8),
cuneiformibus. Prsemolares raro utrinque 2.

Subgen. 3. Euelephas. — Dent, molar. 3 utrinque intermedior. coronis lamellosa deinceps numero auctis,
colliculis ani.s-
omeris {e.g. 12+14+18), attenuatis, compressis. Praemolares nulli.

(8) In Falconer's classification (1847) Elephas [Archidiskodon] meridionalis was erroneously grouped with
Elnsmodon [ = Euelephas), while the closely related E. [Arch.] planifrons was grouped with Loxodon; this obvious
discrepancy was corrected in 1857 (Synop. Tab. opp. p. 319), but Falconer fell into a fresh error by uniting E.
{Loxod.) planijrons and E. {Loxod.) meridionalis in the .same group with the African elejjhant, Elephas (Loxod.)
ajricamis, because of the possession of 'digitated' ridge-plates.

(9) From the above historic analysis it is clear that 'Elasmodon' Falc. is preoccupied and that the substitute
name 'Euelephas' Falc. is an invalid and unusable nomen nudum, l^ecause it was based on an artificial and hetero-
geneous assemblage of six species, which, as shown in the present Memoir, belong to five or si.\ entirely distinct
generic phyla, namely:

Elephas aniiquus = Hesperoloxodon^

Elephas namadicus =Pala-oto.rodoi> (Syii.: Sivalikia, Pilgrimia)'
Elephas hysudricus = Hypselephas
Elephas indicus = Elephas
Elephas meridionalis = Archidiskodon
Elephas priinigenius = Ma?nmonteus^

(10) These criticisms imply no disrespect of Falconer's genius, but rather the insuperable difficulties under
which he labored in disregarding cranial characters and in defining from ridge-plate characters only, as pointed out
above in Chapter XV, "Classification of the Elephantoidea." As repeatedly shown in the present Memoir, the
arrangement of the ridge-plates, whether separated ("colliculi grosse digitati") or compressed ("colliculi approxi-

mati") is an unreliable and misleading basis of phyletic and generic distinction, because it is subject to parallelism
or convergence; Cuvier (1806-1836) used both cranial and dental characters in separating the only three species
recognized by him, viz., Elephas africanus, E. indicus, and E. primigenius, as shown above in Cuvier's legends
beneath figure 1042.

Falconer's reliance, for identification and definition of genera, on ridge-plate formulae is shown in the entirely
unnatural arrangement of genera in his Note Book for August 25, 1862 (Pal. Mem., 1868, II, p. 176) as follows:

[Falconer, 1862,1868] Milk True [Totals] [Present Memoir]

Dinothcrium 1+2+ 3 3+ 2+ 2 =13 Deinotherium giganteum
Trilophodon Ohioticus 1+2+ 3 3+ 3+ 4 =16 Mastodon americanus
Tetralophodon Arvernensis 2+3+ 4 4+ 4+ 5 =22 Anancus arvernensis
Pentalophodon 3+4+ 3 5+ 5+ 6 =28[26] Pentalophodon sivalensis
Stegodon insignis 2+5+ 7 7+ 8 + 10-11 =40 Stegodon insignis-ganesa
Loxodon meridionalis 3+6+ 8 8+ 9 + 12 =46 Archidiskodon meridionalis
Euelephas antiquus 3 + 6+10 10+12+ 16 =57 Hesperoloxodon aniiquus
" primigenius 4+8+12 12 + 16 + 24 =76 Mammonteus primigenius
indicus 4+8+12 12+16 + 24 =76 Elephas indicus

'[See p. 1217below for definition of Hesperoloxodon. Editor.] —

^[See explanation —
on p. 1 179 below. Editor.)
'[For doubtful validity of the genus Mammonteus, see footnote on i>.
1117 above. — Editor.]
 : i

1178 OSBORN: THE PROBOSCIDEA

—
Separation of Antiquus (III) and Africana (V) Lines. Dep6ret and Mayet (1923, table, pp. 204 and
205) in their phyletic arrangement of the Proboscidea distinguished five absolutely separate phyla or lines of
elephants, all of which, however, they included within the genus Elephas.

I. Primigenius line: astensis, primigenius, trogontherii, sibiricus.

p, II. Moridionalis line: planifrons, hysudricus, meridionalis inut. cromerensis.
(jENUS
Yll_ Antiquus line: ausonius, antiquus: melitensis, creticus, Cypriotes, atlanticus.
ll/LEPHAS
jY inflicus line: indicus, namadicus.
V. Afiiranus line: nfriranus = Loxodonta F. Cuvier].
[

In the following year Matsunioto (1924.2, Sept. 20) raised Dep^ret's 'III. Antiquus line' into a subgenus of

the genus Loxodonta, namely, Palxoloxodon, cited below.

Falconer, 1857, Osborn, 1918. — Falconer himself was the first to recognize the mutual resemblances of his

British Pleistocene species Elephas antiquus to his Siwalik species E. namadicus (1857, Synop. Tab. opp. p. 319),

l)ecause they both exhibited "collicuU approximati" ; he did not perceive, however, their phyletic resemblance in
cranial characters to the African elephant {Loxodonta africana), but only the difference in the ridge-plates ("col-
liculi dilatati") of the latter. It remained for several subsequent writers (Pohlig, Pilgrim) to perceive this very
close resemblance, and Osborn (1918.468, table opp. p. 134) finally placed Elephas africanus, E. antiquus, and E.
namadicus in the generic phylum Loxodonta, subfamily Loxodontinse.

Osborn six years later (1924.633, p. 2, Dec. 20) selected Elephas namadicus Falc. and Caut. as the geno-
type of his new genus Sivalikia, to embrace also E. antiquus, both distinguished from Loxodonta by more numer-
ous and more compressed ridge-plates (i.e., "coUiculi approximati").

Sivalikid, new genus. Typified by Loxodonta namadica Falconer, type species, and distinguished by broad grinding teeth,
numerous ridgo-plates, and absence of 'lo.xodont sinus.' Unfortunately the name Falconer ia is preoccupied for a genus of reptiles,
con.sequcnt]y the name Sivalikia is proposed in honor of Dr. Hugh Falconer's great work on the Siwalik fauna.

In the same paper (1924) he proposed the genus Pilgrimia, typified by the narrow-toothed Elephas falconer
of the Mediterranean Islands (Osborn, 1924.633, p. 2) as follows:

Pilgrimia, new genus. Typified by Elephas falroneri Busk, type species, E. melitensis Falconer, E. rnnaidrse Adams, and E.
antiquus Hecki Dietrich; distinguished by narrow grinding teeth, numerous ridge-plates, ioxodont siims' vestigial or absent.
The name Pilgrimia is given in honor of Dr. Guy E. Pilgrim of the Geological Survey of India, to whom paleontology is in-
debted for the complete solution of the stratigraphy of the Siwaliks and of other mammaliferous horizons of India and Burma.

Pal^oloxodon Matsumoto, 1924.' —Independently of Osborn's proposal of the genus Sivalikia, and at
a somewhat same year (1924), Matsumoto (pp. 256, 260) selected a subspecies of
earlier date (Sept. 20) in the

Elephas namadic^is, namely, E. namadicus naumanni Makiyama (1924), as the type of a new subgenus Palxoloxo-
don, a name which now jjroves applicable to the narrow- to broad-toothed- lo.xodonts of Japan and the East
Indies as well as to similar loxodonts of Eurasia and Africa. An abstract of Matsumoto's paper is as follows

(pp. 257, 260)

Palxoloxodon corresponds to Dep6ret's group of E. antiquus, with molars consisting of a small number of ridge-plates, others
with numerous ridge-plates. When worn the molars show clearly the rhombic form [lo.xodont siiuis] and the enamel folds are
almost ])arallel in the anterior and post<>rior portions of tlu; unworn teeth. Tin; molars vary in width, some being narrow,
others wide. This subgenus is divided into two lines (a) E. melitensis-atlanticus and (b) E. antiquus-namadicus, according to
the classification of Dei)(!;ret. E. antiquus is not extremely narrow, and the number of the ridges is rather numerous. The species
of this group belong to a later period, and the ridge-plates of thi; molai's are less rhombic in form; easily confused with E.
trogontherii. E. ausonius equals the Calabrian in Europe, E. antiquus the middle and lower Diluvian in Europe, E. namadicus
naumanni the lower Diluvian in Japan, E. namadicus the middle Diluvian in Japan and India.
'Unfortunately this article of 1924 did not reach the present author until much of the text of tliis Memoir had been prepared. The original was probably
lost intransmission to America.
-In all elephantine phyla the primitive grinders are relatively narrow, they become progressively hroad as the cranium shortens, broadens, and deepens.
Hence the narrow to broad or wide adaptation.
 . • :

THE LOXODONTIN^: CLASSIFICATION AND HISTORY OF DISCOVERY 1179

OsBORN, 1924.— During these years Osborn was engaged in a very close comparison of the crania and denti-
tion of the loxodonts of Asia, Europe, and Africa, the results of whicli are fully set forth in the present chapter,
finally describing the entire structure of all the known Loxodontinae. Unaware of Matsumoto's publication of
September he independently separated the genus Sivnlikia, based upon the genotypic species 'Elephas
20, 1924,
namadicus' Falc. and Caut., publishing the same on December 20 of the same year (1924), three months later than
Matsumoto's publication. In the present Memoir it is shown that the E. namadicus naumanni of Makiyama
(genotype' of Palseoloxodon) very close in character to the 'Elephas namadicus' of Falconer and Cautley
is

(genotype of Sivalikia). Although Matsumoto proposed the name Palseoloxodon as a subgenus of Loxodonta,
and Osborn proposed the name Sivalikia as a genus entirely distinct from Loxodonta, Dr. T.
S. Palmer (letter of
March 12, 1929) rules that even if incorrectly conceived and defined, the subgeneric name Palseoloxodon (Sept. 20,
1924) preoccupies and anticipates the full generic name Sivalikia (Dec. 20, 1924). Osborn accordingly accepts
Palmer's ruling, although it appears to do injustice to his four years of research, in which his conclusions quite
differ from those of Matsumoto. Meanwhile the generic name Sivalikia runs throughout the text and many
of the illustrations ;
the reader, therefore, should make the following substitution: Palseoloxodon (syn. Sivalikia,
Pilgrimia)
Original Definition op Genera (1825-1924)
These four generic lines were originally separated and defined by the characters of the subgenera and species
grouped within them, as follows:
I. Cemis: Loxodonta F. Cuvier, II. Subgenus: Palseoloxodon III. Genus: Sivalikia' Osl)orn, IV. Genus: Pilgrimia' Osborn
1825, 1827. Matsumoto, September December 20, 1924. December 20, 1924.
20, 1924.
Genotypic species: Elephas Genotypic species: Elephas Genotypic species: Elephas Genotypic species: Elephas
africanus Bluraenbach. namadicus naumanni Makiya- namadicus Falconer and Caut- falconeri Busk.
ma.' ley.

Relatively narrow grinding Ridge-plates expanded some- Relatively broad grinding Very long and narrow grind-
teeth with relatively few ridge- what mesially, 'loxodont sinus' teeth with numerous ridge- ing teeth, with numerous and
plates : rudimentary, vestigial, or ab- plates : closely compressed ridge-plates
M
3 TT^T2 sent, enamel typically plicate; Max.: MBifilHf Max. : ?M 3 ff
relatively narrow grinding teeth,
with numerous ridge-plates:
Max.:M3K-
Conservative in cranial and Progressive in cranial and Dwarfed elephants of the
grinding tooth structure. grinding tooth structure. Mediterranean Islands.
Distinguished by very broad 'Loxodont sinus' absent or 'Loxodont sinus' vestigial or
'loxodont sinus'; enamel non- rudimentary; enamel plicate. absent.
plicate, or smooth.
Descended from unknown an- Large elephants of Japan. Including animals of gigantic Including animals of diminu-
cestors- to the modern Loxo- size. tive size.
donta africana.
Premaxillaries broad. Premaxillaries broad. Premaxillaries very broad. Premaxillaries broad.
tusks widely divergent.
Inci.sive

Parietofrontal cranial vertex Cranium [of genotype] un- Parietofrontal cranial vertex Parietofrontal vertex expand-
rounded, subacrocephalic. known. [Cf. P. namadicus fig- broadly expanded, with promi- ed, with frontal crest.
ured by Falconer and by Pil- nent frontal crest.
grim (Fig. 1041).— Editor.)

The genus Loxodonta is read- The subgenus Palseoloxodon The genus Sivalikia, typified The genus Pilgrimia, founded
ily distinguished tjy its primi- Matsumoto, 1924, founded on by 'Elephas namadicus,' is dis- on the narrow-toothed, dwarfed
tive cranium, primitive ridge- the species ^Elephas namadicus tinguished by its relatively species 'Elephas falconeri,' with
plate formula, and specialized naumanni' Makiyama, 1924, is broad giinding teeth, numerous less numerous ridge-plates, may
lozenge-shaped or 'loxcxlont also characterized as relatively parallel ridge-plates, and the ab- prove to be ancestral to the
sinus.' narrow toothed. sence of a 'loxodont sinus.' It subgenus Palseoloxodon of Mat-
proves to be the same as the sumoto, 1924.^
subgenus Palseoloxodon Mats.
'[See footnote by Dr. E. H. Colbert on page 1247 below. —
Editor.]
^[Compare Loxodonta -prima and L. africana var. obliqua of Dart, 1929, on page 1287, below Editor.]
'[Sivalikia and Pilgrimia finally determined by Professor Osborn as synonyms of Palxoloxodon Matsumoto.- -Editor.l
 : . —

1180 OSBORN: THE PROBOSCIDEA

Definitions and Distinctions by Osborn of the Two Genera

There can be no question of the clear generic distinction of Palxoloxodon^ (synonyms Sivalikia and Pilgrimia)
from Loxodonta. Subsequent research may reveal more than one generic phylum within the genus Palxoloxodon,
liut at present it would appear, as explained in the following paragraphs of this chapter, that the phylum Palxolox-
odon originated in Africa, migrated northward into Europe, possibly developing into the 'Elephas antiquus'
group, also northeastward through the Mediterranean Islands, leaving dwarfed descendants resembling either the
'E. antiquus' group or the '£'. namadicus' group, finally arriving in southern and southeastern Asia, developing
into the 'Elephas namadicus' group.

Meanwhile the animals composing the great generic phyla Palseoloxodon and Hesperoloxodon, as in parallel

M 3 \~\{
species of Archidiskodon, increased in size and multiplied their molar ridge-plates from the minimum,
(Palseoloxodon atlanticus), to the maximum, M 3 y? (P. namadicus nawmanm), and from the minimum, M 3 l^it^
{Hesperoloxodon antiquus typicus), to M 3 ^^^^ (H. antiquus germanicus), to M 3 -— {H. antiquus italicus).

The species in these two great phyla are progressively distinguished by the successive addition of ridge-plates,

iiy by the absence (in the typical 'E. [Hesperoloxodon] antiquus'

the increasing hypsodonty of the grinding teeth,
group) of the transverse bony ridge above the orbits, or by the presence {E. mnaidriensis, E. melitensis, E. nama-
dicus = Palseoloxodon]) of this broad bony ridge above the orbits. Consequently according to this interpretation
\

the subfamily Loxodontinse contains three chief generic phyla, as follows

I. (loims: Loxodonta F. Cuvicr, 182.5, 1827. II. (lenus (subgenus): Palxoloxodon Mat- III.- Genus: Hesperoloxodon Osborn, 1931.
sumoto, 1924; Subgen. or syn.:
Sivalikia Osborn, 1924. Pilgrimia
Osborn, 1924.

Goiiotypio spocios: Elephas nfriranus Genotypic species Elephas namadicus Genotypic species Palseoloxodon an-
Rlumenbach. naumanni Makiyama. tiquus italicus Osborn.'

Relatively con.servative and primitive Relatively progressive in cranial and Cranium domelike with flattened
in cranial and grinding tooth structure. grinding tooth structure. forehead, more hypsicephalic and
bathycephalic than that of Pala'oloxo-
don

Cirinders distinguished by broadly Grinders distinguished by rudimen- Grinders hyp.sodont, 'loxodont sinus'
oi)en 'loxodont sinus' (lozenge shaped), tary, vestigial, or absent 'loxodont vestigial or absent.
on wear; enamel non-plicate or .smooth. sinus' ; enamel typically plicate.

Grinders relatively narrow, low crown- Primitive grinders relatively narrow,

Ridge-plate formula:
ed, with comparatively few ridge-plates: with increasingly numerous ridge-plates:

Max, M Min.: M3f|^ Max. M 3

Min. : M 3 f§|^ Max. : M 3 ^+
: .3
nrf^. :

Premaxillarii's broad with widely di- Premaxillaries extremely broad, in- Premaxillaries extremely broad, in-
vergent incisive tusks, elongate, slightly cisive tusks widely divergent, straight ci.sive tusks widely divergent, slightly
iiiciM'ved. or slightly incurved. upcurved and incurved.

Parietofrontal cranial vertex low, Parietofrontal cranial vertex some- Lacking prominent parietofrontal
rounded, .subacrocephalic, of persistent what more progressive, acrocephalic, crest so distinctive of Palseoloxodon
primitive form. with smooth or with prominent frontal namadicus.
ridge-crest for attachmeni of the
muscles of the trunk.
'[Subsequently (19:}1) Professor Osborii's rcscarchr.s on the skull of PateoZozotion a/iii^uu.s' iVnh'cu.s led him to the <unclu.sion that this sub.species belonged
in phylum from that eontaining 'Elephas' namadicus; con.sequently he provisionally assigned to it and to other members of the 'E. antiquus'
a quite di.stinet
group the new generic name llesperolnxodon (see Osborn, 1931.846, p, 21), retaining the name Pal:i'olox<nii>n for members of the 'E. namadicus' group.
Editor.]
'[Compiled from statements in the text. — Editor.)
'[See footnote by Dr. K. H. Colbert on page 1247 below. —Editor.)
 THE LOXODONTIN.E: CLASSIFICATION AND HISTORY OF DISCOVERY 1181

2. HISTORY OF DISCOVERY AND SEPARATION OF EUROPEAN, INDIAN, MEDITERRANEAN,

AFRICAN, JAPANESE, AND JAVANESE SPECIES
Europe: Sceleto Elephantino ToNNiE, 1695 (Tentzelius, 1698). In 1695 occurred the discovery near —
Gotha, northern Germany, of the classic 'Burg-Tonna skeleton,' followed by the first scientific descriptions, as
fuUy set forth in Chapter XVIII above, pp. 1118-1119; this famous skeleton, which aroused the interest of all
the savants of western Europe, was referred to by Blumenbach (1799) in his type description of Elephas primi-
genhis (also cited in Chapter XVIII, p. 1141), but it now proves to belong to the 'Elephas antiquus' phylum,
judging by the few parts that remain of this originally fine specimen.

As shown above in Chapter XVIII (footnote, p. 1119) of the present Memoir, parts of the Burgtonna skeleton
were presented by Tentzelius to the Royal Society through Sir Hans Sloane and are now lost. Other parts are
still preserved in the Gotha Museum. Dietrich writes (April 14, 1930) that there is no doubt that the classic
Burgtonna skeleton belongs to Elephas antiquus; its geological horizon is the travertine sands of the last Inter-
glacial Period, that is, it belongs to the same deposits as those of Weimar-Taubach-Ehringsdorf, a more recent
phase of the last Interglacial. Amthor of Gotha confirms the fact that the skeleton, incisors, tusks, etc., are
present in the Gotha Museum, but illustrations are not available.

Other European remains now known to belong to 'E. antiquus' were for a long period confused with those
of the southern mammoth, 'Elephas meridionalis.' It is also a singular and generally unnoticed fact that the
generically related 'Elephas namadicus' of India was described by Falconer and Cautley in 1846, a year before
they figured (1847) the 'Elephas antiquus' of Europe, as shown in the following history of nomenclature.

In the present Memoir are reproduced the type figures of all the species of Loxodonta, Palseoloxodon (syn.:
Sivalikia, Pilgrimia) , emd Hesperoloxodon so far as published by the authors, also so far as the type specimens have
been available.

India: Elephas namadicus Falc. and Caut. (1846). — It was a long time before the relationship of the
Indian species Elephas iiamadicus to the African genus Loxodonta rather than to the Asiatic Elephas indicus was
recognized. In 1846 Falconer and Cautley (1846, p. 45) observed:

Another extinct Indian species E. Namadicus (to be described in the sequel), which is closely allied to the existing Indian
form [i.e., Elephas indicus], comes between it and E. Hysudricus, together with a European fossil specie.s [probably referring to
the as yet unnamed Elephas antiquus Falconer and Cautley, 1847], which we believe to be distinct from the Mammoth [i.e.,
Elephas primigenius Blimi.]; and the gap between the existing African Elephant and E. planifrons is filled up by another well-
marked European fossil species, E. priscus (?), pi. 13, fig. 7, which is closely allied to the former.

The type of 'E. namadicus'' (Fig. 1070) was designated (Falconer, 1867, p. 15; 1868, Vol. I, p. 435) as from
the "valley of the Nerbudda. Probably a female, from small size of tusks." Falconer writes {op. cit., pp. 15
and 435) : "I have named the species E. Namadicus, after the Nerbudda river, the Namadus of Ptolemy."

Falconer and Cautley, 1847. —Falconer became impressed with the close resemblance between this Middle
[to Upper] Pleistocene species of India, which he named E. Namadicus in 1846, and the British Lower Pleistocene
proboscidean, which he named Elephas antiquus in 1847.

Great Britain —
Elephas antiquus Falconer (1847). Up to 1844 all the British fossil elephants had been
:

referred to 'Elephas primigenius.' At that time Falconer was arranging and describing his rich collections from
the sub-Himalaya and river deposits of central India, wloich had been aided by the contributions of Sir Proby T.
Cautley, Mr. Eraser, and others. The superb plates of the "Fauna Antiqua Sivalensis," beginning in 1845, dis-
1182 OSBORN: THE PROBOSCIDEA

play Falconer's confusion of the Elephas meridionalis of Nesti with the animal which he first erroneously named
'Elephas priscus' [after Goldiuss = Loxodonta africana] and finally in 1857 clearly distinguished in his mind as
Elephas antiquus.

The first figure of ^Elephas antiquus,'' published in 1847, PI. xii.d, fig. 4 (Brit. Mus. M.2006), may be taken as
Osborn's lectotype, although designated in the plate as "Elephas meridionalis,'" but corrected by Falconer in
a copy of the "Fauna Antiqua Sivalensis" belonging to the British Museum. Falconer's first' printed description
of E. antiquus is that pubUshed by Murchison in 1867, p. 18, in the legend of figures 4, 4a, of PI. xii.d; this de-

scription applies to a first true molar {M.^, fide Lydekker, 1886, p. 130) with twelve to thirteen ridges. On a sub-
sequent page {op. cit., 1867, p. 21) he described (PI. xiv, fig. 7), as 'Elephas priscus,' an unmistakable specimen of
E. antiquus from Gray's Thurrock (Brit. Mus. 39,370), designating it as a "la.st lower molar, left side, containing
eight ridges, heel inclusive" [l.Mo, see Fig. 1076 below]. He adds: "Besides the great expansion,this tooth differs
from all true E. antiquus specimens in the lowness of the crown ridges. ... If not a separate species, [it] is a very
marked variety."

Leith Adams (1877-1881, p. 47) erroneously determined the ridge formula of the typical Elephas antiquus as:

-L-'P ^ +3 + -? -'-'P "J +6-8+ ^P^+9-ll+ '*^ -l

+11-12+ -^''^^+12-13+ •^''^ <J
+1619 + '

Mediterranean Islands: Malta, Sardinia, And Sicily (1862-1907)

The existence of pigmy elephants on the island of Malta {Elephas melitensis Falc), as first announced at the
Cambridge meeting of the British Association, October 6, 1862, astounded the palaeontologists. Besides two
dwarfed species {E. melitensis Falc, 1862, 1868, and E. falconeri Busk, 1867), both found in the Zebbug Cave
of ]Malta, there were traces of a third elephant of nearly normal size referred to Elephas antiquus.

In addition to reviewing this important discovery, Leith Adams (1870, p. 224) added a third species {E.
mnaidrse), a name amended by its author to E. mnaidriensis in 1874, p. 116). "This species [Lydekker, 1886, p.

138] is considered to have averaged between six and seven feet in height and to have been allied to the narrow-
crowned race of E. antiquus and also to E. africanus. The ridge-formula is given by Leith Adams [Footnote:
'Trans. Zool. Soc. vol. ix. p. 112.'] (exclusive of talons) as":

'Elephas mnaidrse' : Dp 2 f Dp 3 f Dp 4 |^ M 1 1^ M 2 +t M 3 \f^,.

This formula is much lower than that of Hesperoloxodon antiquus, as cited from Leith Adams above; it is

actually lower than that of H. antiquus as now determined, namely, M3 lUlrii; but it is higher than that of
Loxodonta africana, namely, M 3 -i~rn-
Sicily. — The fossil elephants discovered from early times in Sicily included forms which Falconer considered
identical with the full-sized 'Elephas antiquus' and with 'E. melitensis,' a dwarfed form. Pohlig (1893, p. 81)

described the latter, from the Elephant Cave of Carini near Syracuse, as Elephas {antiquus) Melitse Falc. Soergel
(1912.2, p. 1) applies the term El. antiquus var. insularis to the Sicilian stage from Carini. According to our record
no new species have been described from Sicily, in which large island also the nearly full-sized elephant recorded as

'Elephas antiquus' prevailed at a certain period before it became dwarfed.

'Compare Preface by F. A. Bather to Memoir by Andrews antl Cooper "On a Specimen of Elephas anliquus from Upnor," 1928, p. iii.
 :

THE LOXODONTIN^: CLASSIFICATION AND HISTORY OF DISCOVERY 1183

Raniiro Fabiani (1928, p. 34) records the distribution of the dwarfed elephants of Sicily as follows:

—
Elephas {antiquus) mnaidriensis Leitli Adams. Grotte di S. Ciro, Olivella, Benfratelli, Billiemi, -S'. Elia, dei Pimtali,
Maccagnonef, S. Teodoro. Corso Vittorio Emanuele, Teatro Massimo, Giardino Inglese, Cappucciui, Acqua dei Corsari, Fossa
della Garofala (Palermo), Presso Case Capizzi (Monreale). Montelepre, Alcamo, Marianopoli, regione 'La Romana' presso
Sciacca, Siracusa, Tabuna di Ragusa.

—
Elephas (antiquus) melitensis Falconer (incluso I'E. Falconeri Busk). Grotte S. Giro, Addaura, LupareUo e Stazione
di Boccadifalco, Cava dell' Arena, Carburanceli, Cava Muletta (Capaci), S. Teodoro, della Santa (Siracusa). Montelepre,
Tabuna di Ragusa.

— In 1883 Forsyth Major described the dwarfed

Sardinia. Elephas Lamarmorae, from species, this island.

Eastern Mediterranean. — In 1903 Dorothea Bate described the diminutive form Elephas ctjpriotes

from the island of Cyprus. In 1907 the same author described the diminutive form Elephas creticus from the island
of Crete.

The dwarfed elephants of the Mediterranean Islands were long considered as subspecies of 'Elephas antiquus,'
but the cranium of 'E. melitensis,' as figured by Pohlig (Fig. 1121), more closely resembles that of E. [Palxoloxodon]
namadicus of India.

In October of 1929 appeared Raymond Vaufrey's admirable Memoir "Les Elephants Nains des lies Mediter-
raneennes" (Arch. Inst. Pal. Humaine, Mem. VI). This Memoir, with its invaluable observations and detailed
studies, has been briefly abstracted and annotated in Section V of the present chapter. Osborn accepts many of
Vaufrey's conclusions but gives his reasons for hesitating or declining to accept others.

Extinct African Species and Subspecies, Discovery and Description (1875-1929)

About 1875 discoveries began in the Pleistocene of North Africa. The first fossil species described, Elephas
atlanticus Pomel, 1879, proves to be a primitive loxodont and in the present Memoir is treated as Palxoloxodon
atlanticus. To the same species may be assigned Elephas africanus foss. of Thomas, named in 1884. In 1895
Pomel also described another form from Algeria, namely, Elephas jolensis, which is also a loxodont and is here
treated as Palseoloxodon jolensis.

Ancestral forms of the true 'Elephas africanus' (cf. Pomel, 1895, pp. 66, 67, Pis. i-iii) suddenly occur in the
more recent Quaternary deposits of North Africa, as excellently described and figured by Pomel, who in his "Re-
sume General Stratigraphique" distinguishes the Quaternary Proboscidea of the North African littoral as follows

Most Recent
Elephas africanus [ = Loxodonta africana] not included in any of the prehistoric drawings of more recent age. Certainly-
living inBarbary at a prehistoric period, but not figured. Found in the geologic horizons of least antiquity. Ridge-plate and
dental formulae corresponding with Falconer's formulae of Loxodonta africana.

Elevated Marine Littoral, Pleistocene

Elephas [Palseolo.rodon] atlanticus Pomel most characteristic and abundant of this period, affording a complete dental
formula. More primitive than Elephas [Hespei-oloxodon] antiquus, a contemporary of Bubalus antiquus in southern Oran; figured
in wall drawings by contemporary prehistoric men, who showed the difference from the African species by the conformation
of the ears (cf. Fig. 1047).
Elephas [Palieoloxodon] melitensis{f) Falc. ref. Quaternary of Ternifine.
Elephas [ = Palseolo.rodon] jolensis Pomel, of Jol ou .Julia casaroea, Cherchel, related to antiquus and to innaidriensis of the
island of Malta, but with less numerous ridge-plates.

Pliocene Series
Mastodon = Zygolophodon] borsoni
[ ref.
Elephas [
= Archidiskodon] meridionalis ref., possibly referable to Archidiskodon planifrons.
1184 OSBORN: THE PROBOSCIDEA

It is to these six species of proboscideans that Pomel's Memoir of 1895, "Les Elephants (iuateniaires," is

devoted.

PAL.150L0X0D0N AND HeSPEROLOXODON AS DEPICTED BY THE CaVE MeN OP NoRTH APRICA AND SpAIN. WiTH SMALL EARS
The left-liand figure, after Pomel, probably relates to Palaioloxodon atlanticus; the right-hand and middle figures, after Breuil, probably relate to
Hesperoloxodon antiquus platyrhynchus
Fig. 1047. The leit-hand figure is designated by Pomel (1895, Explanation of PI. xiv, fig. de Ouebar-Reeliim." Algeria.
4) as follows: "Elt'ipliant atlantique
The right-hand and middle figures are designated by Breuil (1911, p. 61, text fig. 57, and p. 239, text fig. 245) as follows:(Right figure) "I'Jlfiphant
trace en rougi; (largeur o"' 44) et signes formes de lignes rouges verticales en faisceau." Cavern of Pindal, provinct! of Asturias, northern Spain. Middle
figure) "Figures d'Klephants graves dans le Sud Oranais .... d'apres Flamand." Algeria.
Observe that these figures, although drawn by different artists, present a striking general agreement, that is, in the general body profile, the elevation
of the limbs, thi; slight eoneavity of the forehead, the shortness of the inferior abdominal region, and especially, as noted by Pomel, in the contour of the rel-
atively small and dejiressed ears, which entirely differ from those of the living African elephant or even from those of the pigmy variety {Loxodonia africana
pumilio). CoiLseciuently these figures are used in the preparation of our re.storations of the Upnor elephant (Fig. 1074) and other representatives of Hespemloxodon
ajiiiquus. The contour of the ear resembles that cf the Indian elephant (Fig. 1120) rather than that of the African elephant (Fig. 1052).

The first fossil elephant to be described from South Africa was Scott's Elephas (Loxodon) zulu, 1907, which
we refer to Loxodonia zulu (see Osborn, 1934.925, p. 2).

This was followed in 1916 by Dietrich's description of Elephas antiquus Recki of northern Tanganyika
Territory, referred in the present Memoir to Palaeoloxodon recki.

The supposed Loxodonia griqua of Haughton, from South Africa, described in 1922, proves to belong to the
more ancient genus Archidiskodon, namely, A. griqua.^

Germany and Rumania

In 1924 Sabba Stefanescu described the two subspecies Elephas antiquiis rumanus from Tulucesti, Rumania,
and E. antiquus germanicus from Tanganu (Ilfov) Rumania, horizon Taubach and Weimar, Germany, as well
of
as a referred E. antiquus ausonius and a referred E. antiquus germanicus also from Ilfov, Rumania. It is somewhat
doubtful whether these moderately l)road-plated types (Figs. 857 and 1089) are truly referable to the Elephas
[Hesperoloxodon] antiquus group or to the extremely broad -plated Archidiskodon planifrons. Judging from
Stefanescu's figures it would appear that:

Elephas antiquus rumaiius^. ^id., oi Wumama. -Archidiskodon planifrons rumanus. Upper Pliocene
of Rumania.
Elephas antiquus germanicus S. Stof., of Ciormany and Rumania = Hesperoloxodon antiquus qermnninis. UiipcrPlei.stoconc
of Cicrniany and Rumania.
Elephas antiquus atisonius rof., of Ilfov, Riunania = Hesperoloxodon antiquus ausonius. ?Upper Pliocene.

'(Professor Osborn in 1934 (1934.925, p. 12) niad(! Lnindnnta griqua the tyiM' of a new genus, namely, Metarchidiskodon (aco Chap. XVI, p. 994).-
Editor.l
 : :

THE LOXODONTIN^: CLASSIFICATION AND HISTORY OF DISCOVERY 1185

Japan and Java

(Compare abstract of Matsumoto's articles, also letters of July 14 and November 20, 1924, pp. 901-908, Chap. XIV)

Japan. — For a long period Japan constituted the extreme eastern portion of the continent of Asia, and, as

described in recent papers and memoirs by Matsumoto and others, contained a stegodont and elephant fauna
closely related to that of Burma and India. Beginning in the year 1924 Matsumoto has given a complete synopsis
of this mastodont, stegodont, and elephantine fauna, which summarized above (Chap. XIV, pp. 901-908).
is

In the present chapter it is shown that all the elephants described by Matsumoto under different generic and
specific names probably belong to the Loxodontinse and are more or less closely related to Palxoloxodon (syn.
Sivalikia) namadicus ;
none of them appears to be related to the genus Loxodonta or to the genus Parelephas as
Matsumoto supposed. In the order of description, these Japanese species are as follows:

Original Name, Matsumoto, Makiyama Specific Reference in Present Memoir

Elephas namadicus naumanni Makiyama (English description

June 30, 1924, p. 264) = Palseoloxodon namadicus naumanni

Made the genotypic species of Palxoloxodon Matsumoto (Sept. 20,
from an admirable type which
1924, p. 257) is fully described and refigured
on pp. 1294-1296 of the present Memoir.

Elephas namadicus namadi Makiyama (English description

June 30, 1924, p. 264) = Palseoloxodon ?iamadicus namadi^
Loxodonta Cuvier
Tjrpe Elephas africanus.
Palseoloxodon, n. Matsumoto (Sept. 20, 1924, pp. 257,
Subgenus.
=E. antiquus-namadicus group
260) = Genus Pal^oloxodon
Euelephas protomammonteus, sp. n. Matsumoto (Sept. 20, 1924, p. 262) = Palseoloxodon protomammonteus
Loxodo7ita [Palseoloxodon) tokunagai, sp. n. Matsumoto (Sept. 20, 1924, p. 267) = Palseoloxodon tokunagai
Loxodonta {Palseoloxodon) namadicus naumanni (Mak.), Matsumoto (Sept. 20,
1924, p. 268) = Palseoloxodon namadicus naumanni
Loxodonta {Palwoloiodon) namadicus Falc. and Caut. (typicus), Matsumoto
(Sept. 20, 1924, p. 269) = Palseoloxodon namadicus
Parelephas protomammonteiix proximus Matsumoto (1926.2, pp. 48-50) = Palseoloxodon prolomaminonteus proximus
Loxodonta {Palst'oloxodon) namadica {Yahei) Matsumoto, 1929, i). 4 = Palseoloxodon namadicus yahei
Loxodonta (Palseoloxodon) Tokunagai J luiior mut. Matsumoto, 1929, p. 10 = Palseoloxodon tokunagai mut. junior

Of the aljove species six type figures have been published which are available for reproduction in the present
Memoir, namely, Makiyama's type figures of Elephas namadicus Naumanni (here reproduced as Fig. 1152
(1)

below) and (2) of Elephas namadicus namadi (Fig. 1153); (3) Matsumoto's type figures oi Euelephas (Parelephas)
protomammonteus (Fig. 1154) and (4) of Parelephas protomammonteus proximus (Fig. 1155); also (5) of Loxodonta
{Palseoloxodon) namadica (Yahei) (Fig. 1156), and (6) of Loxodonta (Palxoloxodon) Tokunagai junior mut.
(Fig. 1157).

In 1924 (Sept. 20) Matsumoto pubUshed in the Journal of the Geological Society of Tokyo, Vol. XXXI, his
article "Preliminary Note on Fossil Elephants in Japan" (pp. 255-261), in which were listed the species named
in the column above, with the exception of the subspecies mentioned in the preceding paragraph, namely, Nos.
2, 4, 5, and 6. This Japanese paper was not received by the present author until February, 1929; if sent, the first
copy was lost in transmission.

'[See "1924 Elephas namadicus namadi Makiyama," Chap. XXI, p. 1409, of the present Memoir. — Editor.]
 :

1186 OSBORN: THE PROBOSCIDEA

In 1926 (Matsumoto, 1926.1) the same treatment was repeated in the Enghsh language and the same series
of species was Usted with the locahties in which each was found, as reproduced in Chapter XIV, pp. 901-908, of the

present Memoir. This list includes the following (cf. Chapter XIV, pp. 906-908, for full citations from Matsu-
moto's printed and written synopses)

9. Euelephas protomammonteus Matsumoto, 1924, p. 262. Smaller than E. trogontherii. Of Calabrian age.
10. Euelephas trogontherii (Pohlig), 1924, p. 265. Of Cromerian age.
11. Elephas indicus Linn., 1924, p. 266. Probably of Post-Monastirian and Pre-Neolithic age.

12. Loxodonta (Pabeoloxodon) tokunagai Matsumoto, 1924, p. 267. Possibly older Pleistocene.

13. Loxodonta {Palxoloxodon) namadica naumanni (Makiyama), 1924, p. 264. Of Cromerian [Tvower Pleistocene] age.
14. Loxodonta {False oloxodon) namadica (Falc. and Caut.), 1924, p. 269. Of Milazzian-Tyrrhenian age.

As stated above the specific and generic reference of these elephants awaits further study and comparison.
With the possible exception of the specimen identified as Elephas indicus, Osborn is inclined to regard them all

as belonging to the Palseoloxodon (syn. Sivalikia) phylum.

Java. — In 1908 Dubois described from the Kendeng formation, Pithecanthropus zone, Lower or Middle
Pleistocene of Java, Elephas hysudrindicus as standing near the Elephas hysudricus Falconer of the Siwaliks, but
still more close to Elephas indicus, accordingly naming it hysudrindicus. Stremme (1911) related this species more
closely to 'Elephas antiqiius.' Accompanying the publication of figure 1160, this species may confidently be
placed with Palseoloxodon as P. hysudrindicus.

I M I . I II i.. t I. .. v . i
-
i . i^v;_ \ iii , 1 -1. . 11 1! I 111 i rl-i ir I I-

{ LOXODONTINAE: LOXODONTA- PALAEOLOXODON

Fig. 1048. Geographic distribution (according to tlie numbers in the following list, which represent the chronologic sequence of type description) of
the principal species of the Loxodontina;. See also figure lOr)."), map of distribution of existing African elephants.
 THE LOXODONTIN^: CLASSIFICATION AND HISTORY OF DISCOVERY 1187

3. ORDER OF DISCOVERY AND DESCRIPTION OF THE FIFTY-THREE TYPE SPECIES

OF THE EXTINCT LOXODONTIN^

1. 1797 South Africa

2. 1798 South Africa

3. 1821 Germany
4. 1823 Germany
5. 1846 India

1846, 1847 England

6. 1847 Europe
1188 OSBORN: THE PROBOSCIDEA

29.
 Habitat of the African Elephant (Loxodonta)
Forest and savanna of the Uasin Gishu Plateau, Kenya Colony, near the 'Nzoi River. After photographs by
Kermit Roosevelt (right) and Carl E. Akeley (left)

Fig. 1049. Calf of old female charging elephant. Open savanna country. Fig. 10.50. Females and young bulls in forest. Photograph by Kermit
Photograjjli by Carl E. Akeley. Roosevelt.

Last of the African Elephant (Loxodonta)

Fig. 1051. A small herd of elephants, containing two bulls, two cows with calves, and one young bull in the center, passing througli the grassy meadows
of an open savanna country in British East Africa. After a film photograph taken by Martin Johnson in the year 1923.

1189
 . .

C
Akeley Group of African Elephants in the American Museum
Fig. 1052. .\ .ipciial expedition (if the Ameriean Museum
Natural History was sent out in 1909 under Carl E. Akeley to eolleel materials for this group of
of
African el<M)lianls, male, female, and young, for the African Hall of the Museum. The cow elephant was shot by ex-President Theodore Roosevelt, and the calf
l)y Kerniit Roosevelt, in 1909, on the Uasin Gishu Plateau where the Roosevelt and Akeley expeditions met. The large bull was shot by Mr. Akeley.
The composition {.\, B) shows the bull scenting danger, silently feeling fcr scent with his trunk, ears fully extended to catch the least s<nuid, for h<' docs not
si'C the source of disturbance. The attitude of the cow indicates that she has seen the intruder and has "frozen," cars liack, trunk pendant, prepared for
any move she may decide on, whether attack or retreat. The calf, con,'<cious of the alarm, is snuggling \ip to its motlier for protection. The young bull to the
right (C), startled, has started forward to swing around and fa<-e I lie danger, his trunk thrown back to catch the scent, and his ears forward to catch tlie .sounds.

Measurements in Millimeters of Adult and Young

Anii'r. Mus. 32727 Amer. Mus. 32732 Amcr. Mus. 32731 Amcr. Mus. .)K)8.">
Calf Female Small Bull Large Bull
Height of >houlder 13.')8.9 2590.8 2711.45 3251 .2 = 10 ft. 8 in.
Height to top of head U09.7 2679.7 2628.9 3403.6 = 11 ft. 2^
1473 2 = 4 ft. 10
( "iroimference of forefoot (io 051 1171 75 1352 55
. .

Height above iK-lvis 1346.2 2151 1 2489.2 3048. = 10 ft.

Total spread of ears 1066 8 2100 3 2870.2 3149.6 = 10 ft. 2}^

Exposed length of task.s:
(a) left 38.1 .508 635. 1422.4= 4 ft. 8
(b) right 44.45 508 660.4 1447.8= 4 ft. 8%
Geogkaphic Data of the Four African Elephants in the Group
1 The large l)\ill (Ijiixtuliiiitn nfricnm nlbrrlntxi>;). 3. The old female (Loxod/mta ajricana pirli).
Locality: Biidongo P'orcsl, eastof Lake Albert, I'nyoro, northern Uganda. Locality: Near 'Nzoi R,iver, Uasin Gishu Plati-au, Kenya Colony.
Amcr. Mus. Dept. Mam. 5 1085. by Colonel Theodore Roosevelt
.Shot
(According to condit ion of molars about middle-age.) Amer. Mus. Dept. Mam. 32732.
2. The small bull (Loxodonta africana albertnisis). (.\ccording to condition of molars approaches senile condit ion.)
Locality:Budongo Forest, ea.st of Lake Albert, Unyoro, northern Uganda. 4. The bull calf (Lox.odonia africami prdi).
Amcr. Mus. Dei)t. Mam. 32734. Lottality:Near 'Nzoi River, Uasin Gishu Plateau, Kenya Colony. Shot
(.^ccording to condition of molars rather older, but certainly not younger by Captain Knrmit Roosevelt,
t han the large bull No. 54085.) Amer. Mus. Dept. Mam. 32727.

1190
 .

II. SYSTEMATIC REVISION OF THE LOXODONTIN/E

Superfamily: ELEPHANTOIDEA Osborn, 1921
Family: ELEPHANTID^ Gray, 1821

Subfamily: LOXODONTINyE Osborn, 1918

Original reference: Osborn, Bull. Geol. Soc. Amer., XXIX, table (opp. p. 134), p. 136 (Osborn, 1918.468); Amer. Mus. Novi-
tates. No. 1, 1921, p. 15 (Osborn, 1921.515).
Compare Pohlig, 1885-1888, Loxodonten, Loxo (disko) donten, Loxo (disko) don {Elephas africanus, E. priscus, E. aniiquus).

—
Subfamily Characters. United by the common dental and cranial characters of Loxodonta,
Palseoloxodon, and Hesperoloxodon. Cranium relatively primitive, platycephalic, brachycephalic
(Loxodonta) somewhat more elevated or hypsicephalic in Palxoloxodon and Hesperoloxodon. Pre-
;

maxillary rostrum broadened for insertion of widely divergent incisive tusks. Tusks relatively straight
or slightly incurved, continuously serving in uprooting habits, chiefly in forests. Grinding teeth moder-
ately hypsodont, typically narrow to broad; lozenge shaped (Loxodonta) or with 'loxodont sinus'
rudimentary or absent (Palxoloxodon, Hesperoloxodon) Habits chiefly browsing, crushing of coarse leaf-
.

age, herbage, and wood fiber. Ridge formula progressive from 3 (Loxodonta) to M n^
3 y| (Palxolox- , M
odon) to
, M
3 —+ (Hesperoloxodon)

The grinding teeth of Palxoloxodon, as shown in P. namadicus, with "colhculi approximati" of Falconer
(Figs. 1070 and 1189), certainly display no resemblance to the lozenge-shaped grinders of the existing species of
Loxodonta, all of which are of the typical 'losange' form (Figs. 1043, 1058) first observed by F. Cuvier. Of somewhat
intermediate ridge pattern are many of the species of Hesperoloxodon, such as are shown in figure 1076 in the
^Elephas priscus' of Falconer and Cautley, but less distinctly in the lectotype specimen Elephas [Hesperoloxodon]
antiquxis (Fig. 1075) . This narrow 'loxodont sinus,' although but a rudiment, is characteristic of many of the species
of Palxoloxodon in the Mediterranean Islands and in Africa; it is also observed in primitive Archidiskodon molars.

From the resemblances and contrasts enumerated above in the comparative figures and definition of Loxo-
donta, Palxoloxodon, and Hesperoloxodon, it is certain that we have to do with three distinct generic phyla, the
descent lines of which begin to be known only in Upper Pliocene and Lower Pleistocene times. One of the
strongest proofs of this profound phyletic cleavage is seen in the fact that all the modern living species of Loxodonta
have a more primitive cranial structure and a more primitive ridge formula, namely, M 3 iytt^ (Loxodonta afri-
cana), whereas the long extinct Palxoloxodon namadicus exhibits M 3 yf and the typical Hesperoloxodon antiquus

In brief, in all cranial and dental characters the surviving species and subspecies of Loxodonta africana are
more primitive than any of the known fossil species or subspecies of Palxoloxodon riamadicus or of Hesperoloxodon
antiquus.

Genus: LOXODONTA F. Cuvier, 1825, 1827

Compare Loxodonle F. Cuvier, 1825; unsigned Loxodonta, 1827; Loxodonta Gray, 1843; Loxodon Falconer, 1847-1857.
The South African form Elephas africanus Blumenbach, 1797, is probably the genotypic species. It is important to note
that in 1798 G. Cuvier distinguished the extreme South African form as Elephas capensis.

Generic Characters. —
Ridge-plates of grinding teeth expanding into broad 'losange' or 'lozenge-
shaped' median sinus. Relatively narrow superior and inferior grinding teeth, with relatively few ridge-
plates; total permanent ridges Dp 4 3: Ijr. —M
Enamel borders thick and simple without foldings or
plications. Superior tusks widely divergent as they issue from the broadened premaxillaries relatively ;

straight, slightly upcurved (Fig. 1059) and incurved (Fig. 1063); marked sexual disparity in the tusks
of females (Fig. 1063). Typical ridge formula:

Dp 2 I Dp 3 t Dp 4 I M 1 i M 2 ^ M 3 ^i^,.
1191
 : :

1192 OSBORN: THE PROBOSCIDEA

Premaxillaries broadened; parietofrontal cranial vertex rounded, platycephalic to subacrocephalic.

Cranial profile and section much more jirimitive, less hypsicephalic and bathycephalic than in the Mam-
montinje (cf. pp. 921-925) or Elephantinse (cf. p. 921). Vertebral formula: Cervicals 7, dorsals 20-21,
lumbars 3, sacrals 4, caudals 26-31.
Adaptation of the grinding teeth (cf. Falconer, p. 927 above). The grinders of the African elephant
are relatively primitive in construction the aggregate series of upper ridge-plates (Dp* M^) amounts
; —
to only 32 as compared to 48 in 'Elephas antiqmis' and 64 in Elephas indicus. The molars are shorter,
narrower, and of less elevation than in E. [Hesperoloxodon] antiquus or E. indicus. In E. [H.] antiquus the
numerous and closely set ridge-plates, without mesial expansion, indicate feeding habits similar to those of
E. indicus. In Loxodonta africana the discs of wear exhibit the well-known rhomboidal expansion or
'losange' characteristic of all the species. These low grinding teeth are best adapted to squeezing and
crushing leaves and succulent stems or roots. Besides browsing on the foliage of the Mimosas and
Acacias, these African elephants used their tusks like a crow-bar to tear up the trees of certain species by
the roots.
In Elephas [Hesperoloxodon] antiquus the numerous narrow transverse ridge-plates of Dp 4 3, —M
totaling 48, form a perfect triturating mechanism like that of the Indian elephant. This food adaptation
alone, in Falconer's opinion of 1868, would entitle his Elephas [Palseoloxodon] namadicus to subgeneric
distinction from his 'Loxodon' he accordingly grouped it under the unusable generic names of Elasnio-
;

don (preoccupied) and Euelephas (invalid).

Generic Description, 1825-1827.— Cuvier, Frederic, et Geoffroy Saint-Hilaire, Etienne, 1824-1829,

"Histoire Naturelle des Mammiferes," Vol. Ill, Livr. LI, LII, 1825, p. 2 (Loxodonte)

Je proposerai pour iioni generiquc de cette espece, Ic mot clc Lo-wdonlc, qui pent rappoler Ic caractere clc ises dents, les
lo.sangcs qu'on apercoit sur leur coupe.

A review (unsigned) of this work appeared in the Zoological Journal, London, 1827, 1828, III, p. 140, noticing

the
disincmbcrment of the genus Elephas, for the purpose of e.stablishing a new one under the name of Loxodonta. For the . . .

Klephant of Asia he [Cuvier] retains the original generic name Elephas. The surfaces of its molar teeth present fasciae of
enamel irregularly festooned; while in those of the African Elephant, the type of the new genus Loxodonta, the enamel is
disposed in lozenges. In addition to this striking distinction derived from the dentary system, M. F. Cuvier also ciumierates
tiie other characters which have hitherto been regarded as specific. The smaller, more elongated, and less irregular head of the
African animal when compared with the Asiatic the rounded forehead of the former, strongly contrasted with the deej) depres-
;

sion in the middle of that of the latter; the ear of the former also twice the extent, while the tail is only lialf the length, itc.

1. ORDER OF DESCRIPTION OF EIGHTEEN LIVING AFRICAN SPECIES AND SUSBPECIES'

Osborn does not express any oi)inion as to the validity of all these eighteen African geographic species and
subspecies, as listed below with the aid of Mr. Herbert Lang and Dr. Paul Matschie, but furnishes this geographic-
varietal list as a real contribution to palaeontology and as a picture of past conditions, namely: In past time each
extinct collective species, by local and continental adaptive radiation, doubtless embraced numerous geographic sub-
species and varieties adapted to local conditions and as clearhj distinguishable from each other in external and internal
characters as are the eighteen living African species and subspecies.

From Blumenbach's first notice in 1797 to Roosevelt and Heller's work- in 1914 and finally Frade's articles

of 1924 and 1928=, eighteen species and subspecies have been proposed led by the Elephas capensis of G. Cuvier,

a name which is probably a synonym of Loxodonta africana. This list is extremely valuable to the palseontologist
as demonstrating the very wide range of geographic-varietal characters in existing loxodonts of a single continent.
Undoubtedly a similar range of geographic-varietal characters distinguished the great herds of elephants belonging

'Prepared with the cooijoration of Mr. Herbert Lang of the Ameriean Mviseum and of Dr. Paul Matsehie of the Berlin Mu.se\im. Chronological list of the
eighteen names i)ropo.sed for the living ,\friean elephants, with the name as the author originally deseribf^d it (to the left) and the present proposed name (to
the right), together with referenees to the tyi)C figures, locality of the type specimens, and the museums in whii'h they are to be found.
-Roosevelt, Theodore, and Heller, Edmund. 1914. "Life-Histories of African Game Animals," Vols. 1, II, New York. Also Fradc, Fernando, 192-i,
—
"Notes de Mammalogie .^fricaine," Pt. 2 Les Elephants du .Jardin Zoologique do Lisbonne, Bull. >Soc. Portugaisc Sci. Nat., Tome IX, Fasc. 3, pp. 130-135,
text figs. 1 6, and 1928, "Titulos c trabalhos cientificos (Curriculum vitac)," pp. 15, 16, Lisboa.
 —:

THE LOXODONTIN^: CLASSIFICATION AND HISTORY OF DISCOVERY 1193

to the genera Archidiskodon, Parelephas, Mammonteiis, and Elephas. A similar range of geographic-varietal
characters is now observed in the four to five living species and subspecies of the true genus Elephas, as described
below (Chap. XX).
According to different observers, the vertebral formulae, as given in detail on pages 930 and 931 of the present
Memoir, undoubtedly differ among these numerous subspecies and geographic varieties. The food of the African
elephant, as observed by Falconer (p. 927), also probably differs in the various regions of Africa.

Stjbspeoies
Elephas africamis Blumenbach, 1797, No. 19, fig. C. Type loc: South Africa; according to
Matschio (letter, 1921) probably from the Cape. [Location of type molar unknown.
Editor.] =Loxodonta africana africana}
Elephas capensis G. Cuvier, 1798, p. 149. Type loc: ?Upper Orange River, eastern S. Africa.
Type fig. .skull, in Cuvier, 1799, pp. 1-22. Type probably in Museum, Paris
: = Loxodonta africana capensis
Elephas cyclotis Matschie, 1900, p. 194. Type loc: Mwelle clistrict northeast of Yaunde on
the left bank of the Sanaga, east of the Nachtigall Falls, S. ( "ameroon. Type fig. Heck, :

1899, p. 116, PI. cxLVi (live animal from the Berlin Zoological Garden). Type (.skull) in
Berlin Museum = Loxodonta africana cyclotis-
Elephas (Loxodonta) oxyotis Matschie, 1900, p. 196. Type loc: Upper Atbara, Sudan. Type
not specified; species founded on numerous examples brought back from the Upper
Atbara region, Sudan, by Casanova, Hagenbeck and Menges; according to Matschie
(letter, 1921) a skull, undoubtedly similar to oxyotis, is in the Berlin Museum. = Loxodonta africana oxyotis
Elephas (Loxodonta) knochenhaueri Matschie, 1900, p. 197. Type loc: Barikiwa, Tanganyika
Territory. Type (skull) in Berlin Museum = Loxodonta africana knochenhaueri
Elephas africanus albertensis Lydekker, 1906, p. 1089. Type loc: South end of Lake Albert.
Type fig.: Lydekker, 1907.1, text fig. 121 (skull). Type in British Mu.seum (Natural
History) = Loxodonta africana albertensis
Elephas africanus pumilioNosick,190Q, p. &31. Type loc: French Congo. Type fig. Horna- :

day, 1905, pp. 237, 238 (photographs of live animal). Type in the American Museum of
Natural History (Amer. Mus. 35591) = Loxodonta africana pumilio
Elephas africanus toxotis Lydekker, 1907, pp. 385, 388. Type loc. Mos.sel Bay, western Cape
:

Colony. Type fig.: 1907.1, text fig. 106 (head). Type (mounted female) in South
African Museum at Capetown = Loxodonta africana toxotis
£?ep/ias o/n'canws seZoMsi Lydekker, 1907, pp. 387-389. Type loc: Mashonaland, southern
Rhodesia. Type fig. 1907.1, text fig. 108 (head).
: Type in Imperial Institute, London = Loxodonta africana selousi
Elephas africanus peeli Lydekker, 1907, pp. 393, 394. Type loc: Aberdare Mts., Kenya
Colony. Type fig.: Lydekker, 1907.1, text fig. 114 (mounted head). Lectotype in
private collection of Mr. C. V. A. Peel, 12 Woodstock Road, Oxford. Cotype in Mr. Roth-
schild's Museum at Tring = Loxodonta africana peeli
Elephas africanus cavendishi Lydekker, 1907, p. 395. Type loc. Lake Rudolf district. Type
:

fig.: Lydekker, 1907.1, text fig. 115 (mounted head). Type in British Museum (Natural
History) = Loxodonta africana cavendishi
Elephas africanus orleansi Lydekker, 1907, p. 398. Type loc. North Somaliland. Type fig.
:

Lydekker, 1907.1, text fig. 118 (dried right ear). Type in collection of Due d'Orl^ans at
Wood Norton = Loxodonta africana orleansi
Elephas africanus rothschildi Lydekker, 1907, p. 399. Type loc: French Sudan ("probably
southward of Lake Chad"). Type fig.: Lydekker, 1907.1, text fig. 119 (head from life).
Type (skeleton) in the American Museum of Natural History. Lydekker states that he
takes as type the statuette of "Jumbo" in the British Museum = Loxodonta africana rothschildi^
Elephas africanus cottoni Lydekker, 1907 (1907.2), p. 783. Type loc: Northeastern Congo.
Type fig.: Lydekker, 1907.1, text fig. Ill (ear). Type probably in Powell-Cotton's
private collection = Loxodonta africana cottoni
Elephas africanus Fransseni Schouteden, 1914, p. 396. Type loc. M'Paa near Bongo, north-
:

west of Lake Leopold II. Type fig.: Schouteden, 1914, PI. xi, figs. 1, 2 (dead animal).
Type (skin and skull) in Tervueren Museum = Loxodonta africana fransseni
£/ep/iasa/)7'm?iMs mofam6/cws Frade, 1924, pp. 131, 133. Type loc: Maputo, Mozambique.
Type fig.: Frade, 1924, text fig. 5. Type: Female living at the time of description in
the Jardin de Zoologique de Lisbonne = Loxodonta africana moqambica
Loxodonta africana Zukowskyi (Strand in Zukowsky, 1924), p. 68. Type loc. Kaoko District, :

southwest Africa. = Loxodonta africana zukowskyi

Loxodonta africana angolensis Frade, 1928, p. 15. Type loc: Region of Cunene, southern
Angola at the time of description, living in the Jardin de Zoologique de Lisbonne
;
= Loxodonta africana angolensis
^Loxodonta africana africana Roosevelt and Heller, 1914, Vol. II, p. 739. Type loc: near Albert Nyanza. The typical L. africana africana i.s believed to be
Blumenbach's type from the Cape Colony region. Consequently the Roosevelt and Heller name cannot be used for the subspecies from the Albert Nyanza =
Loxodonta africana ?subsp.
"[Compare, however, "Captive pigmy elephants in America," Journ. Mam., 1934, Vol. XV, p. 248, by C. Emerson Brown. Editor.] —
^[Professor Osborn referred "Jumbo" to the subspecies Loxodonta africana oxyotis (Osborn, 1931.846, p. 21), which would make rothschildi a synonym of
oxyolis.—Editor.)
 1194 OSBORN: THE PROBOSCIDEA

Osborn, 1929: In the present Memoir the skull and skeleton of "Jumbo" (Amer. Mus. Dept. Mam. 3283) are
referred to the subspecies Loxodonta ajricana oxyotis rather than to L. a. rothschildi Lydekker. The eighteen
subspecies listed above are partly distinguished by size, partly by geographic distribution, and partly by the shape
of the ears. The names adopted were listed by Matschie and Lang, with the exception of Loxodonta ajricana
mocambica Frade, Loxodonta ajricana zukowskyi Strand, and Loxodonta africana angolensis Frade.

Fig. 1053. Two growth stagfs in the Sudanese elephant "Khartum" of the species Loxodonta
africana oxr/otis Matschie, trom the Upper Atbara, Sudan, formerly living in the New York Zoo-
logical Park. Both photographs taken by Elwin R. Sanborn and reproduced by courtesy of the
New York Zoological Society. Reduced to a uniform one-fiftieth scale. "Khartum" was born
about 1903 and was captured on tlie Blue Nile in 1906; collected
by Capt. Stanley S. Flower, Director of the Government Zoological
Gardens, Egypt.
(Left) Young "Khartum" photographed in 190S at the age
of 5 years and measuring o ft. 1% in. or l.")67 mm.
(Right) Young adult "Khartum" photographed in 1930 at
t he age of 27 years and measuring 10 ft. 8/4 in. or 3257 mm.
The measurements of the successive heights of this elephant,
abundantly fed on a varied diet, indicate a growth at the shoulder
of 2007 mm. or 6 ft. 7 in., in twenty-four years elapsing between
October, 1906, and January, 1930, namely, an increase from 1250
nun. (4 ft. 1)^ in.) to 3257 mm. (10 ft. 8)4 in.), or an annual growth
of 83.6 mm. (3/4 in.). The rate of growth is very rapid until the
fifth year, then it slows down gradually to 2 in., and finally to '4 in.

a year, namely, from •January, 1929 (10 ft. 7'2 in.) to January,
1930 (10ft. 8',4in.).

rm jn" .Mix .

Fig. 1054. Remarkable cave paintings recently discovered in South Africa

representing still living species of the white rhinoceros and Afii(-an elephant.
Observe especially the extremely accurate profile of the small-tu.sked female
elephant, herewith reproduced one thirty-seventh natural size; also the correct proportions of the long-headed white rhinoceros
reproduced to the same one th'rty-seventh scale; both by courtesy of the London Illustrated News.

In the great area north of the dotted line (Fig. 1055), comprising the Atlas Mts., the desert of the Sahara,
and the more or less mountainous and formerly forested regions Ijordering the Mediterranean Sea, other sub-
species of the African elephant formerly ranged in large numbers, as recorded by Pomel and as summarized by
 THE LOXODONTIN^: CLASSIFICATION AND HISTORY OF DISCOVERY 1195

Sir Harry Johnston from various classical historians. Pomel (1895) also records 'Elephas africanus' in the Upper
Pleistocene of Oran, northern Africa.

Fig. 1055. Distribution of the existing African elephant (Loxodonla);

location of the proposed species and subspecies as indicated by various
authors. The four known type marked by a circle. The
localities are
northern limit of distribution has been indicated by an interrupted line.

1. L. africana Blumenbach. Range unknown, probably South

Africa.

2. L. africana albertensis Lydekker. Lake Albert region.

3. L. capensis G. Cuvier. Upper Orange River District.

4. L. africana cavendiski Lydekker. Galla Country.

5. L. africana cottoni Lydekker. Congo.
6. L. capensis cyclolis Matschie. Southern Cameroon.
7. L. africana fransseni Schouteden. M'Paa near Bongo, region
of Lake Leopold II.

8. L. africana knochenhaueri Matschie. Barikiwa, Tanganyika

Territory.

9. L. africana orleansi Lydekker. North Somaliland.

10. L. africana oxyotis Matschie. Upper .\tbara, Sudan.

11. L. africana peeli Lydekker. Aberdare Mts., Kenya Colony.

12. L. africana pundlio Noack. French Congo.
13. L. africana rolhschildi Lydekker. French Sudan.
14. L. africana selousihydekker. Mashonaland, southern Rhodesia.
15. L. africana loxotis Lydekker. Mossel Bay, western Cape Colony.
16. L. africana t>w(ambica Frade. Maputo, Mozambique.
17. L. africana zukowskyi (Strand in Zukowsky), Kaoko District,
southwest Africa.
18. L. africana angolensis Frade. Cunenc, Angola.

From SirHarry Johnston'.s "A History of the Colonization of Africa by Alien Race.s," 1905, pp. 6, 7: "This Punic explorer
[Hanno the Carthaginian] .started from Cartilage some time in the sixth century before Christ (perhaps about 520 B.C.) ... on
a voyage of discovery mainly. ... In the account given of the journey it is stated that after pa.ssing the Straits of Hercules, and
stopping at the site of the modern Sebu, they rounded Cape Cantin and came to a marsh in which a large number of elephants
were disporting themselves [Footnote 'This is an interesting observation. Not only does the statement repeatedly occur in the
:

writings of ancient Greek and Roman geographers that the African elephant was found wild in Mauretania in the.se times, but
this animal is pictured in the remarkable rock sculptures in the Sus country in the extreme south of Morocco, and in the Roman
mosaics and frescos found in the interior of Tunis, and now to be seen at the Bardo Museum near Tunis. (See for this the
travels of the Moroccan Jewish Rabbi, Mordokhai.)'] . The Carthaginians do not seem to have tamed the indigenous African
. .

elephant (which was certainly still found in Mauretania), but they introduced and used the Indian elephant."

From Sir Harry "The Opening Up of Africa," 1911, pp. 103-105: "Jugurtha [King of Numidia (134 B.C.)]
John.ston's
adopted a plan of fighting learntfrom the Carthaginian armies. He had war elephants of the African species, which he placed
in the van of his attack; but somehow they did not make much impression on the dogged Roman infantry. After the
Roman conquest the African elephant disappears from the annals of North Africa and, no doubt, became extinct everywhere
north of the Sahara except in Morocco, where— in the country near the High Atlas— it seems to have lingered till the arrival
of the Arabs. The camel had been introduced into North Africa from Egypt about 200 B.C., and was rapidly adopted by the
nomad races of Mauretania as an animal very useful in war. About 45 A.D. another general of British fame, Suetonius
. . .

Paulinus, marched up the valley of the Muluya river in Morocco and reached the High Atlas range. He ascended these
mountains to the snows, and descended to the southern side of them into the valley of the Gir stream, and gave a vivid de-
scription of the burnt-looking rocks of the desert and the swarms of elephants in the Atlas forests."
1196 OSBORN: THE PROBOSCIDEA

[The question having arisen as to the existence in the same forest of the South Cameroon elephant and the
"pygmy" elephant, recourse was had to the article of Dr. Glover M. Allen on the "Zoological Results of the
George Vanderbilt African Expedition of 1934. . . . The Forest Elephant of Africa" (Proc. Acad. Nat. Sci. Phila.,

LXXXVIII, 1936, pp. 15-44), in which he comes to the following conclusion. The African elephants are repre-

sented by two general types —the large, more or less typical Bush elephant of the eastern and southern plains,

and the smaller Forest elephant of the Congo forests. The former is Loxodonta africana and its several (?)sub-

species or (?) varieties; the latter is L. ajclotis or L. africana cydotis, type from the South Cameroons (syn. L.

piimilio, etc.). He also provisionally relegates both cottoni and fransseni of the Congo region to the synonymy of
cijcloiis, giving his reasons therefor on pages 21-24. Furthermore he regards the term "pygmy" for the Forest

LOXODONTA AFRICANA PUMILIO 2032mm. ,b'8"

W. AFRICA

LOXODONTA AFRICANA 0XY0TI5 350B mm J I'BKz"

CENTRAL AFRICA

Fig. 1056. Restorations by Margret Flinsch Buba, under the direction

of Henry Fairfield Osborn. One one-hundredth natural size.
The tusks of Loxodonta africana oxyolis were restored after the longest
(Fig. 1062) and heaviest (Fig. 106.5) pairs of tusks recorded. See caption to
figure 1093, this chapter.

Elephant as a misnomer —it is a small variety or a small species if one wishes to so designate it, which may reach

a shoulder height of 9)^ feet as compared with perhaps lly> feet in the largest Bush Elephants. Thus there is no
tremendous disparity in size between the two. In the closing paragraph (p. 41) of his article. Doctor Allen remarks
as follows: "Very probably the Forest Elephant represents more nearly the older original stock from which the
larger Bush Elephants have been derived. The evolutionary development of the latter has doubtless taken place
slowly and over a long period of time with intergradation and interbreeding of long duration before they became
sufficiently differentiated to maintain complete segregation, in the absence of physical barriers. . . For this reason,

and because of the slight but apparently constant differences in carriage, form of ears, and size, the Forest Elephant
is given tentatively the rank of a separate species. Nor is there any ground as yet for believing that it breaks up
further into geographical races within its rather limited range in the rain forest." — Editor.]
 . :

THE LOXODONTIN^: LOXODONTA 1197

2. SYSTEMATIC DESCRIPTION OF SPECIES OF LOXODONTA

Loxodonta africana Blumenbach, 1797 applying the descriptive generic term 'Loxodonte.' Type
Figures 794, 805, 806, 810-814, 816, 893, 908, 912, 992, 995, —
Locality. Matschie (1900, p. 190) states that no exact locality
1013, 1031, 1040-1045, 1049-1054, 1056-1065, 1067, 1081, is known for the type of E. africanus Blumenbach. It is probable
1093, 1107-1109, 1112, 1120, 1167, 1190-1192, 1226, that the type came from the Cape region of South Africa, be-
1234, PI. XXIII
cause at the time the Central African elephants were inaccessible
Elephas africanus Blumenbach, 1797. to European naturalists, whereas the South African mammal fauna
Syn.: Elephas capensis Cuvier, 1798: Elephas priscus Goldfuss, 1821. was comparatively well known. Consequently it is probable that
Type locality. Probably Cape Colony, South Africa. Eighteen hving or the type of Elephas africanus Blumenbach came from the same
recently extinct subspecies described from various parts of Africa soutli of the region as the tyjie of Elephas capensis Cuvier (see note under E.
equator. Extinct forms of the true 'Elephas africanus' described by Pomel
capensis below)
(1895) from the Upper Pleistocene of northern Africa. Range of the species,
formerly abundant, the North African coast and Atlas Mountains in the time
In 1797 Blumenbach clearly defined these teeth, as in the

of Hannibal. See caption to figure 1055. legend of his type (Fig. 1057). In 1846 Owen summed up the
distinction between these teeth and those of the Indian elephant
Elephas africanus, 1797. Blumenbach's paper (1797.1), in (Owen, 1846, pp. 230-232— sec Fig. 1043 of the present Memoir)
which the name Elephas africanus was first used, was accessible to "Thus which the lozenge-
in the African Elephant, (fig. 88,) in
the present author by photostat only, through the courtesy of the shaped plates are always much fewer and thicker than the flattened
British Museum. The chief references are as follows: 1. (Matschie, ones in the Asiatic species, the variation which can be detected in
letter, 1921):"Not in the edition of the 'Handbuch der Natur- any number of the grinders of the same size is very slight. ... In
geschichte' of 1779 [First Edition] but in the Fifth Edition (1797); the molars of the Asiatic Elephant, (fig. 89,) which, besides the
in 'Abbild. Naturhist. Gegenstande,' Heft 2, No. 19, fig. C, the difference in the shape of the plates, have always thinner and more
name is also used. . . . The arrangement of the lamellse would numerous plates than those of the African species, a greater amount
indicate its Cape
can only be a question of the Cape
origin. It of variation in both these characters obtains; and the like . . .

Colony and the Congo, perhaps the French Congo. The confluence caution is still more requisite in the comparison of the molars of the
of the anterior lamellae is peculiar." 2. (Andrews, C. W., and Mammoth (Elephas prirnigenius) which, having normally more
,

Bather, F. A., June 6, 1922): "Handbuch der Naturgeschichte." numerous and thinner plates than in the existing Asiatic Elephant,
Fifth Edition. Elephas asiaticus, p. 124. Elephas africanus, p. present a much greater range of variety."
125. Mammontovaiakost, p. 703. 3. (Sherborn, "Index Animali- Elephas capensis Cuvier, 1798, "Tableau filementaii'e de
um," 1902) Sherborn lists E. africanus in the Fifth Edition of the
: I'Histoire Naturelle des Animaux," p. 149. Type. —
"Handbuch," but reference is not made to a figure in this edition. (Matschie, letter, 1921): "Cuvier had at his disposal one skeleton
—
Type Molar, R. Ma. Blumenbach's type figure (Fig. 1057), from the Senegal and one skull from the Cape [Cap de Bonne-
probably a second right inferior molar, r.Ma, serves to characterize Esp^rance, Cuvier, Mem. France) National des Sciences
Inst, (de
cleai'ly the species with very prominent lozenge-shaped ridge-plates et Arts, sometimes called the Acad^mie des Sciences, Vol. II, ann^e
to which also the generic name Loxodonta applies so aptly; this 7, (1799)]." Figure.— Cuvier, 1799, Pis. iii, iv, fig. 2.
was the feature which commanded the attention of F. Cuvier in Type Locality. —Cape of Good Hope, South Africa.
The Expanding 'Loxodont Sinus' of the African Molah Crown (Loxodonta africana)

Fig. 1057. Type r.M2 unworn crown of Elephas africanus Blumenbach,

1797. After Blumenbach, 1797.2, No. 19: "C. vom Africanischen
Nur bildcn jene beym Elephas asiaticus geschliingelte an beiden Enden paar-
weis zusammenlaufende Linien; hingegen beym africanus rautenformige Fig. 1058. Type of Elephas priscus Goldfuss, 1821, PI. xliv, one-half
Leisten. Diese Zahne der beiderlei Elephanten sind nach Originalen im natural size. Type of a worn molar tooth said to have been found in the
hiesigen academischen Museum gezeichnet." neighborhood of Cologne described by the author as follows: (op. cit., p. 485)
[According to Pohle (see Dr. Glover Allen, 1936, p. 16) tlic present location "Beschreibung eines fossilen Backenzahns vom afrikanischen Elephanten. . . .

of the original tooth of Elephas africanus is unknown: "The molar tootli on Daher verdicnt ein unbezweifelt fossiler Backenzahn, dessen Rhombcn auf der
w'hich Blumenbach founded hi.s Elephas africanus is not now known to be in Kaufliiche deiien des afrikanischen Eleplianten entspreclien, welchen das
existence, although Pohle (1926) searched for it in recent years among the Museum der hiesigen Universitiit kiirzlich als ein schiitzbares Geschenk emp-
collections of the Zoological Museum and of tlie Anatomical In.stitute of the Aufmerksamkeit der Naturforscher
fing, die land es ist daher zu vcrmuthen,
. . .

University at Gottingen."— Editor.) dass auch dieses seltene Stiick in der Gegend von Coin gefunden worden sey."
1198 OSBORN: THE PROBOSCIDEA

(1) According to the above note, the type of Elephas capensis Memoirs" of 1868, and in the plates accompanying that work,
is probably a skull from the Cape of Good Hope. Lydekker most
after precise study, he gave a description of the lower ridge

(1907.1, p. 384) states that Ehphas capensis may "really be in- formula, including the half ridge-crests or "talons," as follows:
.separable from Blumenbacli's E. afiicanus typicus { = Loxodonta (Falconer, "Pateontological Memoirs," Vol. I, 1868, pp. 422,

africana africana], based on teeth from a locality unknown." 440, 441, Plates ii, xiii.a, xiv):

(2) Matschie (1900. p. 190) .states that if it were possible to prove

Elephas africanus. Plate ii, fig. 4a, M2 with 9)2 ridges; fig.

that the elephant in different parts of Africa shows certain dis- 4b,M2 with 9 ridges. Plate xiii.a, fig. 8, lower jaw with Dp4,
tinctive characteristics, i.e., occurs as different subspecies, one Mi; Ml with }^-7-}2 ridges. fig. 4, lower jaw, r.Dps
Plate xiv,
must not readily synonymize E. capensis Cuvier with E. africanus with 6)^ ridges; r.Ma
figs. 5, with 5a,
8)2 ridges. From the above
Blumenbach; it would then be necessary to distinguish E. capensis may be deduced Falconer's lower ridge-plate formula of referred

as the elephant of the Orange River District and E. africanus as Elephas africanus, namely:
Osborn, 1924: In the as fully
some other .subspecies." (3) list,
Dp 3 ^i M 1 jT.^ M2 ^. [M 3
cited above,Ekphas capensis appears as follows "Elephas capensis :

G. Cuvier, 1798, p. 149. Type loc: ?Upper Orange River, eastern Leith Adams, 1879-1881, p. 48. Leith — Adams gives the ridge
S. Africa. Type fig.: .skull, in Cuvier, 1799, pp. 1-22. Type
formula of Loxodonta africana as follows:
probably in Mu.seum, Paris = Loa-orfo« to africana capensis."
Elephas prisons Goldfuss, 1821, 1823. The same lozenge-
shaped ridge-plates are observed in the type of Elephas priscus
Dp 2 fit Dp 3 til Dp 4 ^M 1 ^ M 2 ^M 3 ^^^_.

Goldfuss, 1821. This type undoubtedly belongs to the living Felix, 1912, p. 17. —Felix gives the following ridge formula
species of Loxodonia africana, but it misled Goldfuss into the im- of Loxodonta africana:
pression that he was describing a fossil tooth, as shown in the

legend to figure 1058. Cuvier considered the Goldfuss type as Dp2f Dp3| Dp4^ Mlf M 2 ,^ M 3 ^.
belonging to a recent African elephant. Falconer remarks (1868,
Vol. II, p. 95): "In the autumn of 1847 I had an opportunity of
examining the specimens above referred to, in company with Dr.
Goldfuss. . . the fracture and texture of the ivory yielded the . . .

appearance characteristic of recent teeth, and conveyed to my

mind a corresponding impression that the molar was probably of
modern origin." Nevertheless Falconer employed the name
Elephas (Loxod.) priscus for undoubted Pleistocene fossil teeth
from Gray's Thurrock and elsewhere. Subsequently (cf. Leith
Adams, 1877-1881, pp. 1, 2) Falconer abandoned the name
Elephas priscus Goldfuss, and it is now considered {fide Pohlig)
as a synonym
of Loxodonia africana.
Pomel, 1895, p. 20, revives the name, after a very careful re-
view: Elephas africanus priscus, Goldfuss. He says that the teeth
of the africanus priscus type are found in Europe and certainly in
northern Africa. In Plate ii, figure 1, he refers the latter, from the
mountains of I'Oued Bourkika, to E. africanus.

CONSTANT RIDGE FORMULA OF LOXODONTA AFRICANA AS

OBSERVED BY VARIOUS AUTHORS
De Blainville (1839-1864, ^de Falconer, 1865, p. 264): Dp 2 f
Dp 3 ^ Dp 4 I M 1 I M 2 'i^ M 3 j^,.
Owen ("Odontography," 1840-1845, p. 638): Dp 2 J Dp 3 ]

Dp 4 ; M 1 1 M2 , .^ M3 - I
2-

Falconer (1865, p. 265) on De Blainville's and Owen's

materials, "excluding the two talons": Dp 2 | Dp 3 g Dp 4
M 1 M 2 ,% M 3 J^. Fig. 10.')9. PhotoKrapli of the domc-shapi-d skull and of the tusks (Amor.
Falconer (1865, p. 265) observes that a left M'*, eleven inches in Mus. Dept. Mam. 21889) of adult male Loxodonia africana peeli ref., reduced
length, from Cape Colony [region typical of Loxodonta africana to about one-twentieth natural size. Represents the cranium characteristic of
elephants of eastern equatorial Africa. Specimen collected in 1911 by Mrs.
capen.'iis Cuv.], exhibits thirteen j)lates, "i.e., eleven principal
back talons," namely, 3 ^"
""" Carl E. Akeley on southern sloiws of Mt. Kenya. Tusk.s measure 8 ft. aYi in.
ridges, besides front and JVI .

and 8 ft. 9'^ in. respectively and weigli 112 and U.') lbs. each.
Falconer, 1857-1868. ^Although Falconer did not adopt the Now exhibited in the Age of Man Hall, .•Vmcrican Museum, beside the
term Loxndon as of full generic value, he referred to these animals skull of Hesperoloxodon antiquun italiciis. Compare figure 1107 of tin' .same
as Loxodon, a subgenus of Elephas. In his "Palaeontological crania.
 THE LOXODONTIN^: LOXODONTA 1199

Constant Ridge Formula. —According to the above, the similar to those in Palxoloxodon namadicus and Ilesperoloxodon
ridge formula of Loxodonta africana is constant and very conserva- antiquus; (6) the palatal a.spect of the Loxodonta cranium should
tive, since the M 3 ridge formula of the living African elephant 1)0 examined side by side with the palatal aspect of the Elephas
closely corresponds with that of the primitive Upper Pliocene
Archidiskodon planifrons.
The constancy of the ridge formula in the two existing species
of elephants,Loxodonta africana and Elephas indicus, is a very im-
portant fact in its bearing on the probable constancy of ridge
formula in extinct species of elephants of a single geologic time
period, for example, in Mammonteus primigenius of Upper Pleisto-
^"^^
5^ Loxodonta africana (Jumbo)
A|V xIa *""' '^'"- 3283
ill

cene time, as observed both by Falconer and Osborn.

CRANIAL CHARACTERS AND TUSKS OF LOXODONTA AFRICANA

(Continued from Chapter XV, pp. 915-927)
Cranial Axis. —The profound characters of the platycephalic
cranial axis of Loxodonta africana in comparison with the hypsi-
cephalic Elephas indicus and Archidiskodon imperalor are clearly
shown in figure 805, also in figures 806, 812, 811, 810, and 813. In
Chapter XV, the introductory .section on the cranium of the
Elephantoidea, these profound differences between the craniiun of
Loxodonta and the crania of other genera of elephants are much
more apparent than the superficial differences which are displayed
in the accompanying figures (Figs. 1041, 1059, 1060, 1061, 1062).
The summary of the sectional or axial characters of the Loxo-
donta africa7ia skull is as follows : (1) Occipital plane, perpendicular
to ba.sis cranii, extremely long; (2) frontal plane extremely short,
convex anteroposteriorly and transversely; (3) nasals broad and
rounded; (4) occipitohorizontal section extremely broad with
deep pit for ligamcntum nuchse; (5) cranial dome, i. e., occipito-
horizontal contour, uniformly rounded; (6) cranium more platy-
cephalic and brachycephalic and less hypsicephalic and bathy-
cephalicthanthatof Mawmo/iteus, of il rc/iirf/sA:odort, or of Pa?e/ep/(as.
These primitive characters of low, rounded cranial dome and
of relatively mesocephalic profile contour are best displayed in
figure 1059 and in figure 1060. The latter illustrates the classic mid-
dle-aged skull of "Jumbo," a Sudanese subspecies. We observe:
(1) That in the fully adult skull the dome is continuously rounded
from the occipital condyles to the broad extremities of the nasals,
presenting the widest contrast to the profiles of Archidiskodon, of
Parekphas, of Mammonteus, and of Elephas; (2) that while
actually brachycephalic, the cranium of Loxodonta compara-
is also
tively mesocephalic or elongate (shown in the relatively long mandi-
bular ramus. Fig. 1060), as compared with the deeply depres.sed
mandibular ramus of Elephas indicus, or with the extremely bathy-
cephalic and abbreviate mandibular ramus of Mammonteus primi-
genius; (3) that since both superior and inferior molars, M', Ms,
are shorter and less hypsodont, the maxillary and mandibular
dental cavities are much less deep, thus accounting for the less
hypsicephalic proportions of this part of the skull ; (4) the superior
aspect of the mandibular rami (Fig. 1060) also displays the relative African Elephant Cranium at the Age of Twenty-four, M^, M2 in Use
prominence and horizontal distinction of the rostrum, again Fig. 1060. Middle-aged skull of tlie Sudanese or Abyssinian subspecies,
presenting a very wide contrast to the deep, hypsicephahc rostrum Loxodonta africana oxyotis (Amer. Mu.s. Dept. Mam. 3283) with nine-plated
of the Mammontinse and of the Elephantinse ; (5) the superior or M2 in use and an unworn M3 still embedded in the jaw. One-eighth natural
frontal aspect of the size. Observe the angular vertex. For full details, see legend to figure 1061.
cranium is beautifully displayed in figure
This is the famous individual, named "Jumbo," purrha.sed in 1883 by
1061A, .showing (a) the short frontal bones, (b) the massive orbital
P. T. Barnum, the American showman, from the London Zoological Society.
prominences, (c) the broad narial openings, (d) the widely separate "Jumbo" died Sept. 1.5, 1885; skin at Tufts College, Medford, Massachusetts;
maxillo-premaxillary sockets for the enormous incisive tu.sks, skeleton in the American Museum.
1200 OSBORN: THE PROBOSCIDEA

indicus cranium (Fig. 800). It will be observed that the Loxodonta W. K. Gregory on E. africanus (Jitmbo Skull). The re- —
skull (1061 B) is relatively broader, more brachycephciHr in all its markable fore.shortening (cyrtocephaly) and deepening (bathy-
dimen.sions. Compare the measurements from the occipital cephaly) of the African elephant cranium, while less extreme than
condyles to the extremitie.s of the prcmaxillarles in Loxodonta that of other elephants, is nevertheless far more advanced than in
(ifricana (Fig. 1061 B) with the measurements in Elephas indicus any species of mastodont. (1) Note extreme compression of the
(Fig. 800); compare also the zygomatic breadth of LoxodniiUi hinder part of the palate, correlated with the very backward exten-

Akkican Elephant Cha.nidm at the Aoe ok Twenty-four. Same Ckanuim a.s in Fioubb lOflO

Fig. 1061. Supfirior and palatal views of cranium of "Jumbo" {Loxodonia africana oxyolis), Amor. Mus. Dept. Mam. 3283; same skull as that shown in
figuro 104.'). One-cightli natural size. Compare B with similar views of Elephas indicus (Fig. 800).

.\ male (twenty-four years) taken in 1861 at the Setit River, northwestern Abys.sinia; famous in history as having lived three years in the
miilillc-agcd
Jardin des Plante.s, Paris, and then transferred to the London Zoological Society, which, doubtful of his temjier, ac('epted in 1882 an offer of .$10,000 made by the
celebrated circus proprietor, P. T. Barnum, of the United States. Jumbo is sup|)osed to have be(^n twenty-four years of age at the time of his death.

africana with the zygomatic breadth of Elephas indicus. The sion and great vertical depth [bathycephaly] of the maxillary
com|)arison sliow.s that what the Loxodonta skull lo.ses in length it alveoli containing M^ Ml (2) Note anterior position of suture
gains in breadth, i.e., in brachycephaly, and in platycephaly. It separating malar from maxillary portion of zygoma. In the primi-
also shows that while relatively narrow and long, the skull of tive mastodont Trilophodon the malar extends forward over the
Elephns indicus gains greatly in vertical depth; it is more hyp- maxillary to form the ])ost.orl)ital i)rocess; very primitive. (3)

sicephalic, more acrocephalic. Note extremely robust [brachyccphalyj buccal portion of zygoma.
 THE LOXODONTIN^: LOXODONTA 1201

(4) Note confluence of anterior palatine canals and wide divergence The different views of the head of this full-grown male ele-
of incisive alveoli. (5) Note reduction in size of auditory bulla phant, as displayed in figures 1063, 1052 A, B,C, all show the same
which is squeezed down close against the basis cranii ; the anterior depressed, relatively flattened or platycephalic profile, correspond-
prolongation under the eustachian tube is less prominent. (6) ing with the short space between the eyes and the top of the head,
Note reduction in size and in-sinking of foramen for carotid artery due to the lesser development of the frontal siiuises, as compared
in the bulla. with Elephas indicus or with the extreme acrocephaly and hypsi-
External Characters and Tusks of Loxodonta africana cephaly of Archidiskodon and of Mammoiiteus.
C).— The external characters of the male
(Figs. 1062, 1052 A, R, Male and Female Crania. —The wide disparity between

Fig. 1062. Comparison op Tusks of Loxodonta and of Mammonteus

(I^cft). Tusks New York Zoological Society collection, one twenty-fourth natural size. Right tusk 11 ft. o'iin. on
of Loxodonta africana oxyotis in the
outer curve; left tusk 11Circumference of right tusk ISK in.; of left tusk ISjg in. Weight of the pair 293 lbs. The locality record of these tasks is East
ft.

Africa; it is reported that they were recently owned by Kuig Menelek of Abyssinia who ]iresented them to a European political officer, eventually they were
offered for sale in London, were purchased by Rowland Ward, and finally were jiresented to the New York Zoological Society by Mr. Charles T. Barney.

(Right). Tusks of Mammonteus primigenius from Alaska, showing the circular curvature and crossing in Mammonteus as comjiared with the lyre-
shaped arrangement in Loxodonta. From original photograph brought from Alaska. One twenty-fourth natural size.

and female and young African elephant are beautifully displayed male and female crania and tusks in species of Loxodonta is shown
in the great group (Fig. 1052 A) which includes two subspecies of in the accompanying figure (Fig. 1063) of two mounted heads from
Loxodonta africana (peeli and albertensis) collected and mounted by east central Africa. The male is the great bull collected and mounted
Carl E. Akeley between the years 1909 and 1922 for the African by Carl E. Akeley; the geographic locality is northern Uganda;
Hall of the American Museimi of Natural History. The bull is the specific inference Loxodonta africana albertensis Lydekker.
is

a typical example of the male African elephant, as carefully ob- The female is a specimen collected by Paul Rainey north of Mt.
ser\-ed and measured by Mr. Akeley in the field; it attains a Kenya; the species is Loxodonta africana peeli.
shoulder height of 10 ft. 8 in. [as mounted, height of head 11 ft. These two heads display the following generic and sexual char-
2^2 in.]; the expanse of the ears is 10 ft. 2}^ in. acters:(1) Uniform elongation or mesocephaly of the cranium, as
1202 OSBORN: THE PROBOSCIDEA

compared with that of Elephas indicus; (2) the male and female fronto-occipital crest in the male as compared with the female
crania are substantially of the same elongate, mesocephalic propor- cranium. (7) Similar disparity, between male and female tusks,
tions; (3) the gigantic eare are relatively of the same size; (4) to that shown in figures 1063, 1090, and 1106, is displayed in all

there a great disparity in size of the female as compared with

is the known members of the Elephantidse or true elephants. The
the male cranium; (5) the female tusks are extremely slender as disi)arity in the known members of the Mastodontidae is somewhat

compared with the male tusks, yet they constitute formidable less extreme between the; male and female tusks.
weapons for the defense of the young; (6) this sexual disparity of
size is equally apparent in extinct species (e.g., Hesperoloxodon I'XETAL SKELETON OF LOXODONTA AFRICANA (?)COTTONI
antiquus, cf. Fig. 1090), and it is important to note that there should Eales, 1926-1929. —The foetal cranium, jaw, and milk denti-
be ob.served a very marked difference in the development of the tion of Loxodonta africana {'i)coltoni (Fig. 1034), as fully described

...^.rsXT^

Male

KiR. 1063. Hkad i)v .\dult Male and Female Loxodonta africana
Compare figun^ 1090, male ami female tusks of Ekphas [Hesperoloxodon] antiquus [gcntumicuji'l], after Polilip, 1888, Taf. I, figs. 1, la, lb, 2, 2u.

male head of Loxodonta africana albertrtusis (Amer. Mus.

l''iill-grown Full-grown female head ol Loxodonta africana peeli eolleetoil north of Mt.
Dept. Mam. from the Biidongo Forest, ea.st of Lake Albert, northern
.'>408.')), Kenya by Paul Rainey and mounted under the direction of Carl E. Akclcy.
Uganda, as mounted by Carl E. Akeley in the .\meriean Mu.seum, 1922. It is now in the private eolleetion of Mr. Percy Madeira of Philadelphia.
 THE LOXODONTINvE: LOXODONTA 1203

in Miss Nellie B. Eales's Memoir of 1926-1929, should be compared elephant skull, such as appears in figures 1060 and 1061.
with the juvenile stages of the cranium of Mastodon americanus No trace of the permanent premolars is found, as might be
(Chap. VI, Fig. 1.31B, Vol. I), also with the juvenile cranium of expected in so young a specimen; the ridge formula, as shown in
Elephas indkus (Chap. XV, pp. 91.5-918, Figs. 796, 797). The figure 1064, is:

cranial contours and the elongated jaw, the occipital condyle on the
same plane as the dentition, the budding incisors with delicate tips, Dp2f Dp 3? Dp 4^
the primitively plated crowns of the lower milk molars, Dp2.4,
contain no foreca.st of the tremendous transformation which this the posterior plates in Dp 4 not being calcified; this formula
foetal cranium is destined to undergo into the typical African agrees with that of Owen, Flower, and Lydekker.

o.t.s

Primitive Form or the Loxodontine Cranium and Dentition, after Eales

I'^ig. Fatal tTaniuin, jaw, and milk dontition of Loxodonta africana (l)colloni of tlic Belgian Congo, locality not given. After ])liotogra|)lis kindly
1064.
sent the present author by Miss Nellie B. Eales. Compare Eales on "The Anatomy of a Fu'tal African Elephant, Elephas africanus (Loxodonta africana),"
Trans. Roy. Soc. Edinburgh, Vol. LIV, Pt. Ill (No. 1 1), 1926, PI. ix, fig. 29, PI. x, figs. 32 and 34 on the ".\natomy of the Head," etc.; see also Vol. LV, Pt.
Ill (No. 25), 1928, on the "Body Muscles," and Vol. LVI, Pt. 1 (No. 11), 1929, on the "Contents of the Thorax and Abdomen and the Skeleton." In Vol. LVI,
Pt. 1 (No. 11), 1929, of this invaluable Memoir, the author's conclusions as to the phylogenetie relationships of Loxodonta are fully set forth.
1204 OSBORN: THE PROBOSCIDEA

Loxodonta cornaliae Aradas, 1870

Figure 106(5

I'"rom Catania, near the monastery of 8anta Cliiara, Sicily.

Quaternary or Post-Pliocene.
Elephas Cornaliae Aradas, 1870. "Sopra un Molare
Elefantino Fossile Riferibile a Specie Distinta dalle Cono-
sciute." Atti Accad. Gioenia sci. nat. Catania, (3), IV,
pp. 233-235. Type. — Superior molar of the right
side. — From Catania, near
Horizon and Locality.
the monastery of Santa Chiara, Sicily. Quaternary or
Post-Pliocene. Type Figure. — Op. cil., figs. 1, 2.

Type Description. — (Arada-s, op. pp. 233-235): cit.,

"Esso e un molare vero, superiore, destro; fe rotto in

mezzo quasi c per hingo, talchfe manca di poco meno
della meta manca ; piu'e del tallone anteriore o del posteri-
ore, e conseguentemente della estrema lamina anteriore,
non che della posteriore; le radici si conservano in ottimo
stato, meno quelle che dovevano spettare alle lamine
mancanti. Sulla sua superficie triturante, qualora stato
fosse intiero, si avrebbero potuto contare otto a nove
lamine. I dischi di logoramento sono romboidali, in forma
di losanghe; lelamine leggermente increspate, non molto
avvicinate. Lc digitazioni della corona dal lato esteriore
son quasi porpendicolari alia superficie triturante e rego-
larmente arrotondite."
"Or, se fossero questi soltanto i caratteri che il molare
in esame presenta, certamente potrebbe per la configu-

razionc del dischi di logoramento, come si h detto, rom-

boidali, Elephas africamis di Blumenbach.
riferirsi all'

Ma e da considerare, che gli angoli mediani delle losanghe

son poco approssimati a quelli delle contigue, e che se la
specie africana offre da 9 a 10 lamine in esercizio sopra
0, "20 o 0, "24 di superficie triturante in hmghezza

secondo Lartet, nel nostro molare, come si disse, non piu

di 7 o 8 se ne possono contare sopra 0, "24 di superficie
triturante."
"Oltre a cio h da notarsi, che la corona del molare,
che descriviamo, calcolandola nella .sua totalita, dob,
comprendendovi cio che manca, presenta tak; ))ro])orzi-
one tra la lunghezza e da non trovarsi in
la larghezza,

altri I'ugualc, essendo, a meglio spiegarci, assai i)iu largo

piDliorzionalmeutc d(-gli altri molari conosciuti."
"Pure si potrebbe, non ostante le premesse consid-
erazioni, e qualora non si volesse tener conto di altri
caratteri del descritto molare, riconoscere in esso una certa
afSnita coU'Elephas africanus. Pero, confrontandolo coi
Fig. 1065. Tusks of tho African elephant, believed to be tlic heaviest (combined
weight 461 lbs.) in the world, .shown in front of a ty|iifal Arab door at Zanzibar. From molari a questa specie, ben de.scritti e figurati nella
riforiti

Kilimanjaro, East Africa, [nir< lia.scd in Zanzibar in 1900 and exhibited for some time by eccellente monografia dcgli Elefanti .siciliani del prof.
Tiffany & Company of New York. The length of the larger tusk is given as 10 ft. 1 in. on Gemmellaro e del Barone Anca, saremmo forzati a dire,
outer curve, circumference 23\ in. at hollow end. One of these tusks is now in the British
o che il nostro spetti veramente all'africano e queUi dei
Museum (weight 226 lbs.). Reproduced after "Ivory and the Eleiiliant in Art, in
sullodati autori appartenenti a specie distinta, oppure
Archaeology, and in Science," 1916, p. 411, by Dr. George F. Kunz, through the courtesy
of the Executors of his Estate. tutto al contrario, essendo che la configurazione dei loro
 THE LOXODONTIN^E: LOXODONTA 1205

dischi di logoramento e Ic altre proporzioni fra loro apertamente Elefanti ai Mastodonti. Per noi il descritto molare dee appartenere
differiHCono." ad una specie nuova, che, ove fosse per tale riconosciuta dai dotti
"Ma tiittc qiicstc riflessioni debbon eedere e caderc a fronte di che saranno per giudicare la nostra opinione, vorremmo portasse
nil carattere singolare, che il nostro molare appresenta, e che in il nomc dell'egregio Prof. Clav. Emilio Cornalia, che ,sui vertebrati
alcun altro non si osserva, ed il quale basterebbe solo a far riguar- ha fatto studii positivi ed utili alia scienza, chiamandolo Elephas
dare come una specie distinta I'Elefante cui esso appartenne. Le Cornaliae."
lamine sono tra loro separate da solchi lati e profondi, molto
Icvigati, i quali a mo di semi-coniche escavazioni, vanno elevandosi

man mano dagli orli esterni, laterali e ad archi intcrrotti della

corona sino ai punti nei quali si pongono in avvicinamento gli

angoli mediani delle lamine. Quest! solchi, che le figure possono

assai meglio mostrare di qualsiiisi descrizionc, non sono da attribuirsi
a cause accidentali, per esser tutti regolarmente architcttati, senza
che si possa rilevare alcuna alterazione nella materia che la corona
esternamente compone, ne a dislocamento delle lamine, perche,
oltre che questo sarebbe un fatto senza alcuno stento riconoscibile,
gli abbassamenti sarebbero solo di un lato ed in modo di dare alle

lamine una disposizione scalariforme. Questi solchi, che, come

Type of Loxodonta cornaliae
abbiamo detto, formano un carattere nuovo e molto singolare,
Fig. 10t)6. Type superior molar of the right side of Elephas Cornaliae
a nostro modo di vedere, potrebbero significare il passaggio degli Aradas, 1870, fig. 2 of plate, two-thirds natural size. From Catania, Sicily.

Fig. 1067. Young .\ddobu.sh Elephant (LoxodoiUa) from Cape

Colony. Probably six months old, with a height at shoulder of about
three feet. Reproduced through the courtesy of Mr. Henry C. Raven,
who took the photograph in 1919.
 HESPEROLOXODON ANTIQUUS GERMANICUS: 4074mm , I3'4y6"e PALAEOLOXODON NAMAD1CU3: 3736min„ I2'3"e
NORTH GERMANY INDIA

HESPEROLOXODON ANTIQUUS ITALICUS; 4068inm.. 13' 4/e"e P MNAIORIENSIS ITOOmm . b'2»?e P MELITENSI3. I400mm., 47>ie
ROME P FALCONER!, 900111 m„2'li/2"e

HESPEROLOXODON
HESPEROLOXODON ANTIQUUS ITALICUS; 3905mm l2'9Ve ,
ANTIQUUS PLATYRHYNCHUS:^—^ 3828mm , l2'6H"e
PIGNATAPO PINDAL N. SPAIN

HESPEROLOXODON ANTIQUUS; 3934inm , 12 lO^a HESPEROLOXODON ANTIQUUS PLATYRHYNCHUS. 362811101, 12'6y/e

UPNOR N SPAIN
OK
Fl(i. IfKiS. REfTOIIATlONS BV MaIIOKKT Kl.lNSCII Hlll.S (I'.KJl), LNDKUTHE DIUECTION OK HknUV I'aIKIIKI.I) OsiKIKN, OK MI'EClhs OF 1 KSPKIinl.oXODON
1

KUUOI-K AND OK PaL,K01.0XOI)ON OK InDIA AND OK THE MeDITEIIKANBAN ISLANDS, IN COMl'AUISON WITH A DRAWING OK HesPEKOLOXODON ItV THE
CaVE MeN OK
NoHTHEiiN Spain. One oNE-iuiNi)itEi>rii natuiiai, size.

1206
 —

III. EURASIATIC SPECIES OF PAL^OLOXODON AND HESPEROLOXODON

Superfamily: ELEPHANTOIDEA Osborn, 1921

Family: ELEPHANTID^ Gray, 1821
Subfamily: Loxodontin^e Osborn, 1918

Genus: PAL^OLOXODON Matsumoto, 1924

Original reference: Matsumoto, Joiirn. Geol. Soc. Tokyo, 1924, XXXI, No. 371, pp. 257, 260 (Matsumoto, 1924.2).
Compare Ekphas namadicus Falc. and Caut., 1846. Genotypic species Elephas na7nadicus naumanni Makiyama, 1924.
Syn. : Elasmodon (preoc.) Falconer, 1847 (in part) ; Euelephas Falc, 1857 (in part). Sivolikia Osborn, 1924, and Pilgrimia Osborn,
1924 (in part). Subgenus Palseoloxodon = 'E. anliqitus-iiamadkus group' of Matsumoto (1924-1926).

Generic Characters. —
Ridge-plates of grinding teeth parallel, closely compressed, waving or
plicate. 'Loxodont sinus' rudimentary or absent. Progressively narrow to broad superior grinding
teeth, with numerous ridge-plates composed of thin, plicate enamel foldings; twelve ridge-plates in
10 cm. total number of ridge-plates in lower grinding series (Dp 4-M 3) fifty-one. Ridge-plate formula
;

progressive from:

P. melitensis, YT^f to P. atlanticus, M3 f|r? H, to P. namadicus [l, to P. namadicus naumanni, ~.

Correlated with more hypsodont and numerous ridge-plated grinding teeth, cranium more hypsi-
ce])halic and bathycephalic than that of Loxodonta; broad, rugose parietofrontal crest (P. 7ia7nadicus)
overlapping the forehead, adapted to muscular insertion of the very broad proboscis. Premaxillaries
(Fig. 1069) broadening inferiorly for the insertion of widely divergent superior incisive tusks. Tusks
relatively straight, slightly upcurved and incurved toward the extremities.

The genotype of Palseoloxodon Matsumoto is 'Elephas namadicus naumanni' Makiyama, 1924. The geno-
typic species of Sivalikia Osborn is 'Elephas namadicus' Falconer and Cautley, as fully explained in the historical
introduction to the present chapter; shown that the generic name Sivalikia Osborn (Dec.
it is also 20, 1924) is

technically preoccupied by the subgeneric name Palseoloxodon Matsumoto (Sept. 20, 1924).

In 1924 (MS.) Osborn observed: "The gigantic elephants typified by the Elephas namadicus of Falconer are
believed by Osborn to constitute a distinct generic phylum, to which the name Sivalikia is applied in honor of Dr.
Hugh Falconer's great work on the Siwalik fauna. They agree with Loxodorita in certain characters of the cranium,
especially in the low occiput and the distally broadened premaxillaries; they differ from Loxodonta in the com-
pressed ridge-plates lacking the 'loxodont sinus,' also in the breadth of the grinding teeth. This generic phylum
may spring from the giant species of North Africa, e.g., Loxodonta allanlica [Palseoloxodon atlanticus], but it is

certainly distinct from the typical Loxodonta africana group."

Historic Period. — It is not impossible that some elephants of the Palseoloxodon type survived into early
historic times, but it is an open question whether the elephants described from Mesopotamia were of the ancient
'loxodontine' or of the modern 'elephantine' type, probably the latter; drawings and inscriptions will probably
be found some day which will determine these relations.

(Letter, A. H. Ciodbej^ Assyrian Royal Inscriptions mention elephant hunting in nortliern Sj'ria, Meso-
.July 19, 1927):
potamia, etc. But no portray the elephant. Have you any information as to the species that once existed there?
reliefs, .so far,
any skeletal evidence? Also, .losephus mentions monster bones exhibited as 'giant's bones' in Palestine.
. . . Benjamin of
Tudela, A.D. 1173, was shown a 'giant's rib' at Damascus, 9 cubits long. (Letter, Godbey, April 5, 1928) "Hunting elephants :

on upper Euphrates is a notable achievement of Assyrian kings. The only portrayal so far is on Black Obelisk Shalmaneser,
small ears like Indian type, tu.sks project up from lower jaw; like a boar's, or 'Babyroussa.' It may be an artist's blunder.
The ideogram for elephant is 'Big-Horned Bison' ('Mountain Ox'), compare Bos Lucas, from Lucania, the 'boot-toe' of Italy.
1207
 .

S. NAMAOICA Ril.
F.lo.. 1847. Pi. XXIVA.FI9.

S. NAMAOICA Typ« 5. NAMADICA Tyn

F»ie., t84T, PI. XII A Falc, 1647. Pi. XII B. Fip. I (r«*.)

S. NAMADiCA R«f.
Falc 1S47. PI. XXIV A. Fig. 4
.

S. NAMADICA Ref S NAMADICA R«f

F*1e. tS47. PI. XLll. F.g XXII F«lc 1847. PI ALIV, Fla XXII (f«».i

S ANTIOUA NESTtI Typt

Pohlig. 1891. P 360. Fig. 109

£. (ANTIOUA) MELITAE P«f S lANTIOUA) MELITAE Rv*.

f>ehit«. }$%3. Tar. I. Fig. 1 P«hi>s. 1893. Tar. I. Fig. 1«

Fig. 1069. Sec explanatory legend on opposite page.

1208
 —

THE LOXODONTIN.E: PAL^OLOXODON 1209

As shown below (Chap. XX) in comments of Murray on the etymology of the word Elephas, signifying ivory,
it would seem that the animal first became known from trade in its tusks originating in Africa, as mentioned by
Homer, Hesiod, and Heroditus, whereas Aristotle treats only of Elephas indicus. From this it does not appear
that the Mesopotamian elephants were known to the Greeks.

Cranial Characters of PALiEOLOxoDON namadicus and Hesperoloxodon antiquus compared with

LoxoDONTA AFRiCANA.— In figure 1041 above the cranial characters of the recent Loxodonta a/nmwa are shown in
comparison with those of the fossil Indian species Palxoloxodon namadicus and those of the European species
Hesperoloxodon antiquus. The relatively low, brachy cephalic, platycephalic, and mesocephalic form of the
Loxodonta cranium, as compared with the hypsicephalic crania of Elephas, of Parelephas, and of Mammonteus, is

more clearly shown in the preceding figures (Chap. XV, pp. 915-926) relating to the comparative structure of
the crania in these genera.

In the present comparison (Figs. 1041, 1069) of the crania of Loxodonta africana, Palseoloxodon namadicus, and
Hesperoloxodon antiquus, we are especially struck by the short and extremely broad rostrum and by other charac-
ters, as follows: (1) Rostrum short and extremely broad, the premaxillaries diverging to the point where the tusks
issue from the skull; (2) thus the bases of the tusks are very far apart instead of being close together as in Mam-
monteus primigenius or relatively close as in E. indicus; (3) whereas the premaxillary sockets are relatively of the
same length in all three species, the divergence of the sockets in P. namadicus is about the same as in L. africana;

(4) the premaxillary sockets are relatively longer and diverge still more widely in H. antiquus; (5) the crania of
both P. namadicus and H. antiquus are distingushed from the cranium of L. africana by the greater development
of the fronto-occipital crest which in P. namadicus engulfs the frontal bones so that there is a very short space
between the lower border of this crest and the extremities of the nasals and the narial openings ; (6) the narial
openings are extremely broad and shallow, they exhibit approximately the same hour-glass-shaped form in L.
africana, P. namadicus, and H. antiquus; (7) the skull of P. namadicus, moreover, is relatively broader and more
flattened or platycephalic than the skull of L. africana; (8) this broadening and flattening of the summit of the
cranium reaches an extreme in the gigantic P. namadicus cranium.

The phyletic affinities of these three types of crania, namely, Loxodonta africana, Palseoloxodon namadicus,
and Hesperoloxodon antiquus, to each other are obvious, while there is a wide separation from the cranial type of
Elephas and a still wider separation from the cranial type of Archidiskodon and of Mammonteus, which represent
the opposite extreme of hypsicephaly.

Fig. 1069. Comparison of Crania of Pal.boloxodon namadicus, Hesperoloxodon antiquus italicus, H. ant. ausonius, and H. ant. platybhynchus
All to the same one-twentieth scale. Compare figures 1041, 1121, 1096, 1105, 1106

Matsumoto had preceded him by three months in the description of Palwoloio-

[At the time this figure was prepared Professor Osborn was not aware that
don, thereby rendering liis own genus Sivalikia a Consequently the reader should disregard the letter "S" in this figure (-Sioalikia) and substitute
synonym.
Matsumoto's genus Pabeoloxodon and Osborn's Hesperoloxodon (a subsequent determination for the antiquus group), as indicated in the following caption.
Editor.]

(Upper row) Palseoloxodon namadicus: One juvenile and two adult crania, front and side views, .\fter Falconer and Cautley, 1846 [1847].

(Second row) Hesperoloxodon antiquus italicus (left), excavated at Pignataro Interamna, Italy, about fifty miles north of Naples. Drawn after photograi)h
of specimen as it lay in the quarry (Fig. 1096); subject to modification after restoration of the cranium (Fig. 1098 and especially Fig. 1106, an orthogonal
drawing from the mount). (Right) Aged Palseoloxodon namadicus, after Pilgrim, 1905, front and side views.

(Third row) Hesperoloxodon antiquus ausonius"? (left), from the Val d'Arno of Italy, after Weithofer, 1890; H. antiquus platyrhynchus (middle), from San
Isidro, Spain, after Graells, 1897; H. antiquus ausonius (right), from the Val d'Arno of Italy, after Pohlig, 1891, erroneously referred to Eleplias (antiquus)
Nestii.

(Lower row) Palseoloxodon melitensis (left), from the island of Sicily, front and side views, after Pohlig, 1893; Hesperoloxodon antiquus ausonius (right),
from the Val d'Arno of Italy, after Pohlig, 1891, erroneously referred to Elephas (antiquus) Nestii.
1210 OSBORN: THE PROBOSCIDEA

Geologic and Geographic Distribution of the Loxodontin^. Originally described by Falconer from —
deposits now determined as of the Upper Pleistocene of India {Palseoloxodon namadicus) and the Lower(?) Pleis-
tocene of England (Hesperoloxodon aniiquus), the geologic range has been extended from the relatively small
Upper Pliocene' H. ausonius to the great ascending Upper Pleistocene mutations (i.e., H. antiquus in Germany =
H. antiquus germanicus), also throughout western Europe, to the progressive Palseoloxodon namadicus of the
Middle to Upper Pleistocene of India, to smaller subspecies or varieties of Palxoloxodon in Japan, described by
Matsumoto, and in Java, described by Dubois, to the P. atlaniicus of northern Africa and a long series of probably
antecedent species in central and southern Africa, as well as to larger and smaller varieties in the Mediterranean
Islands.

The straight-tusked elephants of western and southern Europe have been treated, in succession to Falconer,
b}- Leith Adamsby Pohlig, Weithofer, Soergel, and Berckhemer in Germany and Italy, by Forsyth
in Italy,

Major and Deperet and Mayet in the Pliocene of Italy and of England, by Andrews and Forster Cooper in the
description of the Upnor elephant {Hesperoloxodon antiquus) of England, and finally by Osborn in his description
of Palseoloxodon [Hesperoloxodon] antiquus italicus of Italy.

As the whole Pleistocene period is now estimated at approximately 1,000,000 years, the straight-tusked

elephant Hesperoloxodon antiquus hved in western Europe for an enormously long period of time, it survived many
climatic phases, it doubtless passed through many ascending mutations which will be determined by final mono-
graphic comparison and description. First appearing in Upper Phocene time, it survived three successive glaci-
ations of northern Europe, but perished toward the close of the 3d Interglacial. Throughout this long period its

companions in southwestern Europe were the Parelephas trogontherii, the hippopotami and the rhinoceroses
{Rhinoceros etruscus and R. merckii). The provisional geologic succession and companionship of these four types

may be presented in the table after Osborn and Reeds, 1922-1929 (PI. xxiv).

Osborn, 1929: Pilgrim (1905) first notes that Elephas namadicus is entirely absent from all the Pliocene
Siwalik strata, in which there is no ancestral type from which it might arise; this suggests the probability that

the genus Palseoloxodon originated in Africa, migrated north into Europe, thence to India and the Oriental
regions. As soon as we begin to examine the Pleistocene deposits of the Godavari, the Nerbudda, and the Ganges,
Palseoloxodon namadicus occurs in great abundance; it is also found sparingly in Burma, China, Java, and Japan,
as described by Owen, Martin, Koken, Schlosser, Naumann, Makiyama, and Matsumoto.

SYSTEMATIC DESCRIPTION OF SPECIES OF PALSEOLOXODON AND HESPEROLOXODON

The by Osborn makes no attempt at finality, but is devoted to the establishment of the
present revision
characters and geologic age of the principal generic and specific tyi)es, awaiting fuller monographic revision based
only upon clo.se comparison and measurement of the rich materials which the museums of western Europe and of
India afford. To the winter's knowledge, no complete cranium of Hesperoloxodon antiquus has been figured, and it
is accordingly of interest to reproduce the photograph of a cranium" recently discovered at Pignataro Interamna,
in Valle del Liri, near Cassino, Italy, between Naples and Rome, which appears to be exceptionally complete
(see Figs. 1096, 1098, and 1101 below).
'[See footnote 1 on p. 101!) regarding the ixjssible I>ower Pleistocene age of the Villafranchian. —Editor.]
-'Tliis iTanium wa.s presented (May, 1929) by the author to the American Mu.seum of Natural History and beans the number .\mer. Mus. 22634. When
discovered it was in perfect condition (as sliown in Figs. 1096 and 1009), but it was seriously damaged in removal and in attempts to repair it by the owner.
Full description and figures are given below in the present chapter.
 THE LOXODONTIN.E: PAL.EOLOXODON 1211

Palseoloxodon namadicus Falconer and Cautley, 1846, 1847 recognized and pointed out that his species E. namadicus of India
Figures 1041, 104(>, 1068-1070, 1072, 1073, llOS, 1110 most closely resembled his .species E. aniiquus of western Europe.
Typo locality: Viillcy of tlic Nerhudda (or Narhada), Ciodavari formation Meanwhile he gave his usual thorough description and character-
(st'o I'^ig. 729). Narhada Alluvium horizon, conipai-e 'Elt-phax aniiquus ization of this very imijortant Indian species. The systematic
{namadicus)' Pilgrim, 190"), containing also referred Sifgoihn iiisignis, S. references are as follows:
ganesa, and Rhinoceros unicornis. Ujjper Pleistocene.
E. [Elephas] Namadicus Falconer and Cautley, 1846, 1847.
Specific Characters. —Transverse occipitofrontal rugosity "Fauna Antiqua Sivalensis," 1846, p. 45, and Atlas, 1847. {Op.
very prominent in males and females. Typical ridge formula a.s cit., p. "Another extinct Indian species E. Namadicus (to be
45) :

below (p. 1212). Third .superior molars with an estimated total of described in the .sequel)." Type. Skull showing portions of —
fifteen ridge-plates; 8e ridge-plates in 10 cm.; maxinumi breadth the third true molar of either side (Brit. Mus. M.3092). Hori-
—
zon AND Locality. Valley of the Nerbudda, Upper Pleistocene,
India. —
Type Figure. Op. cit., Pis. xii.a, xii.b, figs. 1 and 3.

Type Skull of Pal.eoloxodon namadicus

Fig. 1070. (?)Female type cranium of Elephas Namadicus Falconer and Cautley, 1846 [1847, PI. xii.b, figs. 1 and 3]; lateral and palatal views of the
skull, showing portions of the third true molar of either side; ten and a half ritlge-plates remain (there were possibly three or four anterior to these, making
fourte(^n or fifteen). One-sixth natural size. Brit. Mus. M.3092; east .\mer. Mus. Warren Coll. 10381. Valley of the Nerbudda (or Narbada), India (.see

(Fig. 729).

101 mm. Breadth of M2 84 mm. ; estimated length of M2 264 mm. —

Type Description. (Falconer in Murchison, 1867, p. 15,
Ridge-plates broad, close set, entirely lacking 'loxodont sinus'; Pis. XII.Aand xii.b "From the valley of the Nerbudda. Probably
:

enamel borders thin. Skeleton of gigantic size, height estimated at a female, from .small size of tu.sks. ... It was chiselled out by Dr.
12-13feet(cf. Fig. 1068). Falconer, and determined by him to be a new species. In a letter
In 1846, Falconer and Cautley in the "Fauna Antiqua Sival- to Lieut. -Colonel Ousely, Dr. F. writes thus: 'It is probably the
ensis," p. 45, first named the Upper Pleistocene species of the most perfect specimen of a fossil elephant's cranium in Europe.
"Valley of the Nerbudda," India, Elephas Namadicus, but did The species is especially interesting from the form of the cranium,
not figure the species until 1847 (Pis. xii.a and b of the "Fauna which is so grotesquely constructed that it looks the caricature of
Antiqua Sivalensis"). Sub.sequently, in the year 1847 {op. cit., an elephant's head in a periwig. I have named the species E.
they figured and named the Lower Pleistocene species of
PI. XII. d), Namadicus, after the Nerbudda river, the Namadus of Ptolemy.'
western Europe Elephas aniiquus. Consequently the name There is a very similar specimen in the Museum of the Asiatic
Elephas namadicus Falc. and Caut. antedates the name Elephas Society of Bengal. Length of remaining portion left molar,
. . .

aniiquus Falc. and Caut. by nearly a year. Later Falconer clearly 7.5 in. [12e ridge-plates in 7.5 in. or 191 mm.]. Width of remaining
 :

1212 OSBORN: THE PROBOSCIDEA

portion left molar, 3.7 in. N.B. —Twelve plates in this extent. able to determine with precision, the formula for the three inter-
Width of palate in front (between molars), 2.8 in. Width behind, mediate molars, and the last true molar, above and below, is
4.1 in." Falconer gives full measurements
cil., pp. 15 and 16) (op. 10: 10, 12, 16, being nearly intermediate between the Indian and
and comparison with E. and E. primigenius.
indicus, E. hysudricus, African Elephants."
Falconer (1868) Materials and Ridge Formula. The —
ridge formula of Palseoloxodon namadicus may be derived by Fal- COMPARISON OF PAL^OLOXODON NAMADICUS AND
coner's careful examination and comparison of the type skull (Fig.
HESPEROLOXODON ANTIQUUS
1070) containing iVP, an imperfect tooth, also of referred upper Pilgrim (1905) on the Godavari Specimens of Pal^o-
jaws (Pi. xiii) and of several referred lower jaws (PI. xii.c, xii.d), loxodon namadicus. —The fullest and most recent treatment of
as annotated herewith the species Palseoloxodon namadicus is that of Pilgrim (1905):
Falconer, "Palseoiitological Memoirs," 1868, Vol. I, pp. 435- (1) He erroneously adopts Pohlig's opinion that Elephas namadicus
438. Typeskull: Plates xii. a, b,c,d and xiii, iidge-platcsM3'- --. and E. anliquus are closely related if not identical species.' (2)
Referred Upper Jaws. — Plate xiii, figs. 1, la, 16, upper jaw, Consequently he describes the Nerbudda species as "Elephas
M^ ridge-plates U-h- Referred Lower Jaws. — Plate xii.c, anliquus (namadicus)," giving namadicus the rank of a subspecies;
figs. 3, 3a, lower jaw, Dp4, ridge-plates lOJs; figs. 4, 4a, lower jaw, inasmuch, however, as Falconer named the species E. namadicus
Ms, ridge-plates 20)^; Plate xii.d, figs. 1, la, lower jaw, Mi, prior to naming the species E. anliquus, this usage (Pilgrim) cannot
ridge-plates M~13-,'i 15 in Mi, ridge-
all; figs. 2, 2a, lower jaw. be adopted; nor are these species identical. (3) He regards the
plates 13; figs. 3, Ma, about 15 ridge-plates.
3a, lower jaw, Godavari river gravels (Lat. 20° 1', Long. 74° 11') as of the same
Falconer's type and referred ridge formula of Elephas = [ Lower [Upper] Pleistocene age as the typical Nerbudda deposits
Palxoloxodon] namadicus is: in which the type of E. namadicus was discovered, containing
alike E. namadicus, Stegodon insignis ref., S. ganesa ref., Equus
Dp 4 Tow M 1 >i.i3->> M 2 fT7 M 3 \^^. namadicus, Hippopolamus H. telraprolodon, also
palseindicus,
Rhinoceros unicornis. He regards E. namadicus as abundant also
Lydekker (1886.2). —Lydekker does not give the complete in the Pliocene of Burma, of China, of Java, and of Japan.
ridge formula of Palseoloxodon namadicus but states (p. 169) that Pilgrim's description of the skull and skeleton of Elephas
the third lower molar in specimens referred to Elephas namadicus namadicus may be freely cited as follows (Pilgrim, 1905, pp. 203-
from Japan has but .sixteen ridge-plates. He characterizes the 206): "The cranium and bones, which I am describing, and which
grinding teeth of E. namadicus as follows (p. 167): "The cheek- represent the species Elephas anliquus {namadicus) Falc. et Cautl.,
teeth of this species appear frequently almost or quite indistin- belonged to an individual of remarkable size. It cannot have
guishable from those of the broad-toothed variety of E. anliquus, stood much less than 16 feet at the shoulder. The cranium, as
although the ridge-formula on the whole, rather higher and the
is, found, is larger than any hitherto recorded. The cranium
. . .

ridges themselves are somewhat taller; some teeth, however, either on one or both sides possesses all the essential features of the
e.specially those from Burma, China, and Japan, show excessive portion above the maxillaries and the foramen magmun. Tlie . . .

plication of the enamel, and thereby approximate to E. indicus, accompanying plates exhibit all the more important characters of
although with a lower ridge-formula. The adult cranium is . . . the present cranium, and the most casual inspection of it can leave
characterized by the presence of a bold, overlapping, transverse no doubt as to its identity with the crania from the Narbada beds,
ridge on the frontals. wliich appears to be wanting in E.
. . . figured by Falconer in the Anliqua Fauna Sivalensis, Plates 12A,
anliquus. ... In India the species occurs in the Pleistocene of the 12B, figs. 1-3, and Plate 24A, figs. 4, 4a, a.s Elephas namadicus.
Narbada valley, and it is probable that the other specimens are These are the two most complete crania which have been known up
from strata of equivalent age." to now, and are preserved in the British Museum. One of them
—
China. From China is recorded a 9X>-f ridge-plated third has small tusks and probably belonged to an adult female. . . .

superior molar (Brit. Mus. 29007); length 166 mm., breadth 101 The present skull is that of a fully grown male. It seems that the
mm., height 180 mm.; laminar frequency 6 in 100 mm. (fide supra-orbital ridge grew forward with age, so that in young skulls
Hoinvood, letter, August 9, 1928). there is a considerable interval between its margin and the extreme
Osbom, 1928: It is important to observe: (1) That the type tip of the nasal process; in the large female skull in the British
ridge formula of PaUeoloxodon namadicus, as deduced above from Museum this interval is sensibly diminished, while in this latest
Falconer's observations, greatly exceeds the constant ridge formula specimen, which represents the largest and presumably the most
of Loxodonla africana; (2) while Falconer also includes an M3 aged type with which we are acquainted, the supra-orbital ridge
with twenty ridge-plates, probably an erroneous generic
it is almost overhangs the nasal fossa, and the interval is reduced to its
reference, the true M 3 formula appears to be 3 IsiH; (3) in M smallest dimensions. .Considering only the teeth and mandible
. .

P. namadicus the ridge-plates are relatively broader, more numer- of E. anliquusand of E. namadicus, Leith Adams remarked . . .

ous, more closely compressed, and entirely lacking the loxodonl that they seemed to him to be indistinguishable. This opinion of
sinus character. (4) In 1868 (Vol. II, p. 261) Falconer observes "In : Leith Adams has been endorsed by many subsequent writers,
the fossil E. aritiquus of Europe, the dentition of which I have been among whom I need only mention Naumann, Weithofer, Pohlig,
HProfessor Osbom (1931.846, p. 21) provisionally made his subspecies Palseoloxodon anliquus ilalicus the genotype of a new genus Hesperoloxodon, as
distinct from PaUcoloxodon, to inchidf^ also other members of the anliquus group (see P"ig. 1068 of the present Memoir), which he finally adopted (Osbom, 1934.
926, p. 285, 1935.937, fig. 2, p. 407, and PI. xi. Vol. I of the present Memoir).— Editor.)
3

53
s as

I- 03

tf

s o

- "2

d o
-5 Z!
CO

?3 6

to

2S

-a -

OS

—o o
o

M Cl,
1214 OSBORN: THE PROBOSCIDEA

and Lydekker. ... So far is this from being the case, however,
that all the .skulls of the dwarf forms which Pohlig . . . has
figured from the Grotto di Pontale von Carini in Sicily bear
a striking resemblance to Elephas namadicus, and leave us
no excuse for separating the two forms specifically. The
accompanying text figure, taken from one of Pohlig's plates
[Fig. 1041], brings out these resemblances in a remarkable
degree. There is no doubt that future discoveries will prove
that the original E. antiquus of Europe possesses the same TYPE
craniological peculiarities as its Indian variety."' LM': lm3 LM^
Pilgrun {op. cil., pp. 208, 209) gives detailed comparative

Fig. 1072. CoMPAKisoN OF He.speroloxodon and Pal.eoloxodon

(SYN. Sivalikia) Superior and Inferior Molars

Drjnvn tn I lie same one-.sixth .scale. Compare figures 1073 and 1 152

A, Hesperoloxndon antiquus after For.ster Cooper, 1924, Pis. ix, x.

'"LM2
Left M', 17 ridge-plates; left M", 12 ridge-plates; left Ms, 1.') ridge-
plates. Lower Pleistocene. From Harrington, England.
rrzolarS
lo/^

JTnfertor-
B, Hesperoloxodnn nuiioniuii after Dejieret and Mayet, 1923, Pis.
molars
x, XI. Left ridge-plates; left M2, 11 ridge-plates; left M3, type,

m
M''', 1.')

19 ridge-plates. Upper Pliocene (Lower Pleistocene?]. San Romano.

Val d'.\ino inf., Italy.
All //s
A/alural
C, Hesperoloxodon antiquus Falc. and Cant., 1847, Pis. xii.d, xiv.a,
Sije
and Leith .-Vdams, 1877-1881, Pi. iv. Left M', 18 ridge-plates; left M-,
14 ridge-plates; left Mo, type, 12-|- ridge-jjlates; left M.3, 17 ridge-
plates. Lower Pleistocene. From England.

D, I'alvfoloiodoH namadicus Falc. and Caut., 1847, Pis. xii.b, d, c.

Left M3, type, 144- ridge-plates; left M2, 14 ridge-plates; left M.i, 19)^

ridge-plates. Upper Pleistocene. Nerbudda Valley, India.

Observe the progressive increase in breadth both of the superior and

inferior molars and the corresponding decrease in length. The superior
and inferior molars of Hesperohxodon ausonius are relatively the longest

and the narrnwesl; th(! grinders of H antiquus (.\, C) are intermediate

.

in width and length; tlie grinders of Pahtohxodon namadicus are

relatively the shortest and the broadest, also the most elevated or hypso-
dont. This change of proportions from the extremely long and narrow
type, H. ausonius, to the relatively .short and broad type, P. namadicus,
is <-orrelated with the i)rogressive hypsicephaly, bathycephaly, and
brachycephaly of tlie cranium. The same principle of the relative
shortening and broadening of the grinders is displayed in figure 1073.
v"H^z
AUSONIA p. ANTIQUA P. NAMADICA

measurements of the crania of Elephas antiquufi {namadicus), E. fossa and of the interjugal sjmce, and in tlie obtusely anglcnl junc-
antiquus, and E. antiquus melilensis in the nuiseums of Calcutta, tion between the frontal and occipital siu'faces. It dilTcrs in (lie
London, Florence, and Palermo, and concludes (p. 210): "E. slope of the occipital surface, in the greater length and shallowness
antiquus [Ifesperoloxodon antiquus] approaches nearest to E. of the inlciTnaxiliary median fossa, in the convexity of th(> occiput,
africanus [Loxodonta africana] in the flattened shajie of the vertex, in the higher position of the maxillary zygomatic ])rocess, ami in
in the .shortness and breadth of l)r<)w, in the form of tlie tem])onil the shorter length and greater width of the sub-orbital foramen."

'[Professor Osborn regarded E. antiquus as referable to his genus Hesperohxodon (see footnote 1 on p. 1 180 above).— Editor.]
 THE L0X0D0NTINJ5: PAL^OLOXODON 1215

GENERIC CHARACTERISTICS OF PAL^OLOXODON NAMADICUS Narbada elephants exhibit in addition the following points of
CRANIUM likene.ss: —
4. The shortness and breadth of the brow and the

(Compare Figs. 1069 and 1041) widening out of the cranium from below upward. 5. The rhom-
boidal outline of the temporal fossa and its sharply-cut, acute-
(Pilgrim, 1905, pp. 206, 207): "The crania of £'. [^Hespero- angled upper margin. 6. The presence of protuberances on
loxodon] antiquus (stem, sp.), imperfect as they show the are, either .side of the occipital fosssa. 7. The almost rectangular
following points in common with the Indian variety [namadicus] bend by which the occipital passes into the parietal, and the ob-
and the pygmy types which serve to distinguish them from all other tusely-angled junction between the parietal and frontal surfaces.

R M3(rev.)Ref.

t-^

SIVALIKIA NAMADICA /Iffer Falconer //-# J^aTural stje

R M^C-ev.) Ref. L M3 Ref.

/s*

7 <S s to n 'i '3 ''"^ SIVALIKIA ANTIQUA After raUoner

s ^ 7 a
L M3 Type

kj^, L M^ Ref.

SIVALIKIA AUSONIA After Heperet Afayef "V ffoman.

,

Fig. 1073. Three Progressive Broadening Stages in the Pal^oloxgdon (syn. Sivalikia) and Hbsperoloxodon
Superior Grinding Teeth. All to the same one-fourth scale.
Compare Figures 1072, 1152, 107,5, 1078

(Upper) Palxoloxodon (syn. Sivalikia) namadicus. Right M', 12+ ridge-plates; left M3, 19)^ ridge-plates, after Falconer and Cautley, 1847, PI. xiii, fig.
la, and PI. xii.c, fig. 4, respectively.

(Middle) Hesperoloxodon antiquus. Right M^ 19+ ridge-plates, after Falconer and Cautley, 1847, PI. xii.d, fig. 5, Kent, England (Canterbury Mus.);
left Ms, 17 ridge-plates, PI. xiv.a, fig. 11, Saffron Walden, England. Of intermediate length and width.
(Lower) Hesperoloxodon ausonius. Left M', 15e ridge-plates, after Deperet and Mayet, 1923, PI. x, fig. 3, Malafrasca (Val d'Arno sup.), Italy; left M3, type,
19 ridge-plates; PI. x, fig. 1, San Romano (V^al d'Arno inf.). Both in the Inst. G60I., Florence. Relatively long and narrow grinders.

Observe as in figure 1072 the progressive broadening and shortening of the third superior and inferior grinders. Also observe as in figure 871 that the
superior ridge-plates are concave posteriorly, the inferior ridge-plates concave anteriorly. Cement areas dotted.

elephants: —
1. The extreme divergence of the inci-sive alveoli 8. The well-marked frontal projection [namadicus] of the crown,
and the broad shallow depression which occupies their centre. which must have given an exceedingly beetling aspect to the
2. The great distance of the occipital fossa from the foramen living animal. 9. The approximately transver.sely oval contour
magnum and the basal breadth and extreme depth of the fossa. of the cranium, when viewed in a direction at right angles to the
3. The strong convexity of the occiput in a horizontal direction, plane of the occiput. It is much broader than high. [10.] E.
which pushes the zygomatic process of the temporal to the front africanus also approaches them to some extent in regard to the 1st,
in an unusual degree. The crania of the Sicilian and of the 4th, 5th, and 9th of the above characters."
 Straight-tusked Elephant of Upnor (Hesperoloxodon antiqdus) in the Britimh Museum. Restoration, under the direction ok Henry Fair-
field OsBORN, BY Maroret Flinsch Buba (1935) TO A one-fiftieth scale (Bulls) and a one-sixtieth scale (Calf)

Skeleton and growth stage based upon the Upnor elephant (Fig. 1079); head or cranium ba.sed upon the Pignataro Interamna elephant (Fig.

1098); the one fragmentary tusk found at Upnor furnishc.'i the length of the ivories in the restoration; ears based upon the cave drawing.s of Africa
and Spain (Fig. 1047). Several months' re.searth and all the .studies involved in the jjreparation of figures 1079, 1080, 1084, 1083, anil 1081 were
also preliminary to the execution of this restoration.

Fig. 1074. This giant straight-tusked elephant, which formerly ranged over western Europe, is here shown in its relatively early Pleistocene stage of
evolution and in a stages of growth which corresponds to the twenty-fifth jear of living African and Indian cle|)hants, namely, in which the second superior and
inferior molars are still in use, while the third superior and inferior molars have not ccme into use (cf. Fig. 1061 of Loxodonla africana).

In other words, the Upnor straight-tusked elephant in the British Museum was about 25 to 30 years of age; it had not attained the full height character-
istic of this species nor the maximum length of the tusks; yet it was a giant in life, measuring from the summit of the scapula to the ground, as originally
estimated by Andrews and Forster Cooix-r, 12 ft. i% in. (3700 mm.), and, as estimated by Osborn, 12 ft. 10^^ in. (3934 mm.) in the flesh.

The views of the skull and tusks, drawn from the Pignataro skull (Amer. Mus. 22034), are extremely accurate and characteristic, with great
directly
breadth between the tusks and corresiMjndingly broad proboscis. The relatively level vertebral contour lacks the marked mill-depression and elevation above
the pelvis which are characteristic of the African elephant (Fig. 1084). The relatively small, low-set ears are restored from the admirable cave drawings re-
produced from Pomcl (1895) and Breuil (1911) in figure 1047 above. As observed in the legend of that figure, the ears resemble those of the Indian (Fig. 1 120)
rather tlian the enormously enlarged and elevated ears of the .\frican elephant (I'Mg. 10.')2).

121G
 Genus: HESPEROLOXODON Osborn, 1931

Original reference: Amer. Mus. Novitates, No. 460, 1931, p. 21.

Genotypic species: Palxoloxodon antiquus italicus Osborn, 1931.846, ji. 21.'

[The genus Hesperoloxodon was provisionally proposed by Professor Osborn in his article entitled, "Palxoloxo-
don antiquus italicus 'Bp. Nov., Final Stage in the 'Elephas antiquus' Phylum" (see Osborn, 1931.846, p. 21), to

include Palxoloxodon antiquus italicus from Pignataro Interamna, Italy. Subsequently in various ways he indicated
his opinion that the typical 'Elephas antiquus'' and other members of the antiquus group should be referred to his
genus Hesperoloxodon, and finally in 1934 (Osborn, 1934.926, p. 285) the name appears as follows: "Hesperoloxo-
don Osborn, a Loxodontine of western Eurasia and Africa, never reaching America." Also the name appears on
a chart between Loxodonta and Palxoloxodon (see Osborn, 1935.937, p. 407, fig. 2), and on page 12 as well as on
Plate XI of Volume I of the present Memoir.

The accompanying definition, therefore, has been compiled from several sources and embodies, as far as can
be determined, the distinctive characters on which Professor Osborn separated Hesperoloxodon from Palxoloxodon
Matsumoto.

—
Generic Definition. (Osborn, 1931.846, p. 21): "Comparison [of the cranium of Palxoloxodon
antiquus italicus] with the cranium of 'Elephas namadicus' shows a strong resemblance in the breadth of
the premaxillary rostrum but an extreme difference in the summit of the cranium, which in 'E. namadicus'
is relatively low and reinforced by the overhanging parieto-frontal crest. This points to Pal. ant. italicus
as a member of a phylum quite distinct from that of the Siwalik 'E. namadicus,' a phylum which if sup-
ported by other cranial and skeletal differences might well constitute a new genus to which the name
Hesperoloxodon, or 'loxodont of the west,' might be applied. This name is provisionally proposed, as
I would not like to be forestalled a second time, as in the case of Palxoloxodon, a generic name assigned

to 'E. namadicus naumanni' by Matsumoto but a few weeks prior to my description of Sivalikia."

Supposed diagnostic characters of Hesperoloxodon compiled from statements in the present Memoir : Cranium
domelike with flattened forehead, more hypsicephalic and bathycephahc than that of Palxoloxodon; the promi-
nent frontoparietal crest or "bold overlapping transverse ridge on the frontals" as mentioned by Lydekker in
his description of E. [Palxoloxodon] namadicus (1886.2, p. 167) is lacking. Occiput relatively narrow and high
(broad and low in namadicus). Grinders hypsodont, 'loxodont sinus' vestigial or absent; ridge formulae: M 3i||f^J^
(typical), to M3 Vrsr igermanicus) to , M3 iy+ {italicus). Premaxillaries extremely broad, incisive tusks widely
divergent, slightly upcurved and incurved. —Editor.]
Hesperoloxodon antiquus Falconer and Cautley, 1847, 1857 Bed, Norfolk). probably belong to the Lower
All these beds
Figures 794, 871, 1068, 1072-1084, 1088, 1109, PI. xxiii Pleistocene, 1st. and arc geologically older than
Interglacial stage,
Type locality: Not recorded, probably Lower Pleistocene. Also very the '2d I nterglacial heds of Mo.sbach, etc., Germany, which contain
numerous referred specimens of Lower and Middle Pleistocene age (see larger and more progressive stages of Hesperoloxodon antiquus, as
Falconer, "PateontologicalMcmoirs," Vol. II, pp. 176-188)
Syn.: 'Elephas pnscus' Falconer and Cautley, 1847; Elephas (Loxod.)
priscm Fal ne 18'7
"
published by Pohlig.
, . ,,, ,
. ,-,,,
They are much older than the beds of Tau-
bach and Weimar contaming H. antiquus germanicus.

Geologic Age.— While the locality of Falconer's type speci- History.— As noted above, the "Sceleto Elephantine Tonnaj
men of Tentzelius (1698) is the first scientific description of the '^fepAas
(Brit. Mus. M.2006) is not recorded, Falconer's referred
specimen, Elephas (Loxod.) priscus, is from Gray's Thurrock anUquus' of Europe; Blumenbach confused this skeleton with
(Essex) ; the same locality yields six of the British Museum speci- that of 'E. prirnigenius' of Europe. The species 'E. antiquus' was

mens described by Leith Adams (1877-1881), while one specimen then confused even by Falconer until 1847 with 'E. mendionalis,'
comes from Clacton (Essex) and another from Cromer (Forest owing to the wide separation of the ridge-plates.

^[See footnote on p. 1247 below. —Editor.]

1217
1218 OSBORN: THE PROBOSCIDEA

Specific Characters. — Promin(>iit triui.svcrsc fronto-occipi- .Viitiqua Sivalensis" belonging to British Mu.scum; also PI. xiv.b
tal crest known to be lacking. Typical ridge formula of Hesperolo.ro- (the first time the name was published).- E. (Eueleph.) antiquus
don antiquus .somewhat greater than that of Paheoloxodon namadi- Falconer. "On the Species of Mastodon and Elephant occurring in
cus, namely; the fossil state in Great Britain," Quart. Journ. Geol. Soc. London,
1857, Vol. XIII, table opposite page 319. Type.— Lower
Dp2 ^ Dp3 ^ Dp4 J^ M 1 ^^ M 2 [| M 3 \^^,. jaw with second lower molar, M2, of the left side (Brit. Mus.
Grinding teeth relatively longer and narrower than in P. namadi- M.2006). Type Locality. Unnkown, undoubtedly Eng-—
cus; height more than double the width of the crown
(Falc). 'Loxodont sinus' ab.sent in the type (Fig.
1075); ab.sent or vestigial in other specimens (Fig.
1076); "mesial rhomboidal expansion of the discs of
wear" (Falc, 1868. II, p. 176). Inferior ridgo-plates
concave anteriorly; superior ridge-plates concave
posteriorly; crowns with thicker cement and rela-
tively thicker and less plicate enamel than in P.
namadirus; "great crimping of the enamel-plates"
(op. cit., Falconer, 1868, p. 176). Extremely narrow
'loxodoiit sinus' indicated in worn ridge-plates of
Hesperoloiodon antiquus and P. namadirus (Fig.
1072). See also characters noted under H. antiquus
italicus and //. antiquus germanicus below (pp. 1238-
1256).
The above specific characters, including Fal-
coner's final definition ("Palseontological Memoirs,"
1868, Vol. II, p. 176) doubtless apply to a collective
species embracing many ascending mutations from
Upper Pliocene to Middle Pleistocene time, but the
tyi)ical Hesperoloiodon antiquus is probably of
Lower Pleistocene age.
Falconer and C'autley named this .species of
straight-tusked elephant in 1847, a year after naming
the Indian .species Elephas namadicus in 1846;
coiiser|uently if the two species are identical, as
allegedby Pohlig and by Pilgrim, the name Elephas
namadicus ha.s the technical priority. There are
many reasons, how(>ver, for treating the great
straight-tusked elei)hant of southern Eiu'ope as
a species distinct from, although nearly allied to, its
Indian relative Elephas yiatnadicus. All authors
Lectotype Second Left Inferior Molar of Hesperoloxodon (syn. Sivalikia) antiquds
agree as to the relationship of these two animals and Falconer and Cautley
there is doubt as to their affinity, but it is
little
One-third natural size
important obscrxc that in Ivilconer's type of
to Fid. 107."). Lcctotypr, I.M2, of Elephas antiquus, first figured a,s "E. mrridionnlis" (Falconer
Elephas namadirus (l'"ig. 1070) th(> grinding teeth arc and Cautley, 1847, PI. xild, figs. 4, 4a); Brit. Mu.s. M. 200(1, locality unrecorded. One-third
niucli broader than in P'alconer's type of
Elephas natural .size. Named E. (Eueleph.) antiquus (Falconer, 18.57, Syiiop. Tab. opp. p. 319), de-

antiqiiun 1075), which are relatively narrow.

(Fig. fined by Falconer, 1867, p. 18, and 1868, Vol.
I, p. 438, as follows: "Elephas antiquus. ]>o\ver . . .

jaw, left side, with first [?| This tooth is a beautiful si)ecimen; shows twelve to
true molar.
(
"onsequently E. antiquus cannot be the same species
thirteen ridges, with front ridge and heel. It narrows excessively in front and behind, like fig.
as E. namadicus, however much the.se animals re- Zo! E. Namadicus! The erimping, Ac, are al.so exactly alike. B. M. [Brit. Mus. M. 2006|. —
semble each other in cranial characters.' Length of molar, 8. in. Width at middle, 2.6 in. Width in front, 1.3 in."

CONFUSED HISTORY OF NAME AND TYPE land, probably of Lower Pleistocene age. Type Figure.—
Elephas antiquus Falconer and Cautley, 1847. "Favma Anti- Falconer and Cautley, 1847, PI. xii.n, figs. 4, Aa.
qua Sivalensis," 1847, Atlas, figs. 4, 4a of Pi. xii.n, figtu-ed as Falconer's Type.— There can be no question that this second
" E. meridional is"hu{ corrected by Dr. Falconer in copy of "P'auna inferior molar (Fig. 1075) should !» regarded as Falconer's type.
'[In 1931 Professor O.sborn pro|)o.sed the genus //espenilaxodmi to embrace members of the 'EUiilins niiliiiuus group, rctai?iing Matsumoto's genu.s I'alscoloxo-
don, 1924, for members of the 'E.' uammlicus group (see pp. 1212 and 1217 of this chapter).— Editor.]
'-[Compare Bather, in Andrews and Cooper, 1928.1, p. iii; also for descriptive legend, see Falconer, 1867.1, pp. 18 and 23, and 1868.1, II, pp. 1,3S, 441, and
443.— Editor.]
 THE LOXODONTIN^: HESPEROLOXODON 1219

(0^^bonl, 1924): This twelve-plated Ai2, of the left jaw, may be Falconer, "Palaeontological Memoirs," Vol. I, 1868, pp. 438-
regarded as the type, since it was the first specimen figured and 440, 442, 443, 447, legends to Plates xii.d, xiii.a, xiv, xiv.a, xiv.b.
named (1847), and first described in 1867 by Falconer. Lydekker Elephas antiquus. Lectotype. Plate xii.d, figs. 4, 4a, Mi [M2]
(1886.2, p. 130) designated it as follows: "M.2006. Part of the with 12-13 ridge-plates, and front ridge and heel, closely similar to
left ramus of the mandible, containing the half-worn ,^7^; locality E. namadicus, crimping, etc., exactly alike. Upper Jaws. —
unknown. Figured by Falconer and Cautley in the 'Fauna Plate XII.D, figs. 5, 5a, M^?, with I6V2 ridge-plates [19 + ]. Plate
Antiqua Sivalensis,' pi. xii.o, fig. 4, 4a. No history." At the time, XIV, figs. 2, 2a, 26, Dp^, ridge-plates &A- Plate xiv.a, figs. 1, la,
unfortunately, Falconer was misled by the spurious fossil type of Dp', ridge-plates 5; figs. 2, 2a, Dp"*, ridge-plates 10; figs. 3, 3a,
Elephas priscus Goldfuss, for in the same table (Falconer, 1857, Dp"*, ridge-plates lO}:, figs, 4, 4a, M', ridge-plates 8; figs. 5, 5a,
opp. p. 319) he cites "E. (Loxod.) priscus (Goldf.) . . . Pliocene r.M', ridge-plates 14)^ [16^2]. Plate xiv.b, figs. 16, 16a, entire upper
. . . England; Lombardy Imperfectly known. FossU remains
. . . molar, M^ ridge-plates 16-17, length 11 in. = 280 mm. Lower
rare." This citation agrees entirely with his previous reference Jaws. — Plate xiii.a, fig. 4, lower jaw, Ms, ridge-plates 6-|-, M3,
(Falconer, 1846, p. 15), in which he recognizes E. priscus Goldfuss, ridge-plates 17; fig. 5, lower jaw. Mi, ridge-plates 12J2. Plate xiv,
1821, as a valid fossil type, although disputed by Cuvier. We figs. 1, la, 16, lower jaw. Dps, ridge-plates }^Q-%, from Grays,
must therefore regard the tooth described as E. (Loxod.) priscus, E.s,sex. Plate xiv.a, figs. 8, 8a, r.M,, ridge-plates 12; figs. 10, lOa,
and reproduced in this Memoir (Fig. 1076), as referable to the true lower jaw, r.Ms, ridge-plates 12}^; figs. 11, 11a, l.M.,, ridge-plates
Elephas [Hesperoloxodon] antiquus of Falconer. 15-16K.

E. (Loxod. ) priscus (Goldf.) of Falconer, 1857

= Hesperoloxodon antiquus
E. (Loxod.) priscus (Goldf.) Falconer, 1857. "On the Species
of Mastodon and Elephant occurring in the fossil state in Great
Britain," Quart. Journ. Geol. Soc, London, Vol. XHI, pp. 345,
346, and table opposite page 319. Type. —A second lower
molar of the left side. Horizon and Locality. — Gray's
Thurrock, England, Pleistocene. Figure. — Falconer and
Cautley, 1847, PI. xiv, figs. 7, 7a, lb, under the name E. priscus? \
FiffJa-
(Brit. Mus. 39370).
Falconer's type tooth of E. antiquus (Fig. 1075) and the tooth
designated as E. (Loxod.) priscus (Fig. 1076) .supplement each
other in the fact that the loxodont character is more evident in the
priscus variety. This is the eight-crested grinding tooth from the
brick earth of Gray's Thurrock, Pleistocene, to which Falconer
applied (1865, pp. 276, 277) the preoccupied name Elephas (Loxod.)
priscus, in allu.sion to the lozenge-shaped crests, which remotely
suggest those of the type of Elephas priscus Goldfuss ( = Loxodonla
njricana). Falconer described this tooth (1865, pp. 270, and 1868,
p. 96) as a "last molar [an error], left side, of the lower jaw."
Lydekker described the tooth (1886.2, p. 133) as a "second left
lower true molar." In the same Lydekker (p. 122)publication,
treats under Elephas antiquus as follows: "Syn. Elephas (Loxo-
it
Rekerred Hesperoloxodon anthjuus
don) priscus, Falconer and Cautley." Fig. 1076. Aged .second LMj, from Gray'.s Thurrock
molar of the left .side,
(Essex), figured as E. [Elephas] priscus"! by Falconer and Cautley, 1847, PI.
XIV, figs. 7, 7a, and as Elephas (Loxod.) priscus by Falconer, 1868, Vol. II,
ELEPHAS .\NTIQUUS AND E. NAMADICUS, FALCONER'S NOTES
Pi. vn, fig. 1. One-third natural size. Brit. Mus. 39370.
OF 1867 AND 1868 RELATING TO SPECIMENS FIGURED IN
THE PLATES OF THE "FAUNA ANTIQUA SIVALENSIS,"
1845-1847
Falconer's type and referred ridge formula of E. antiquus:
Osborn, 1923: Future research will probably distinguish
between the lectotype ridge formula of Elephas antiquus from Dp3 ,|ff:^Dp4 ^^^Ml .,^:,^^M2 ,:^M3 'ttV.Wtr -

Essex, Norfolk, etc., and the collective ridge formula of specimens

from other localities. The above observations of Falconer (1867, 1868) and on
Collective Ridge Formula. — Falconer's conception of the Falconer's plates (by Osborn) are consistent with Falconer's less
dental characters of Elephas antiquus and their points of similarity detailed formula of 1863. We may regard the following as his
to the molars ofElephas namadicus are to be found in the beautiful collective ritlgc formula of Elephas [
= Hesperoloxodon] antiquus:
plates and legends of the "Fauna Antiqua Sivalen.sis" and in the
"Palaeontological Memoirs" of 1868, as follows: Dp 3 I
Dp 4^ Mltf M2i| M 3 ifj^.
1220 OSBORN: THE PROBOSCIDEA

FALCONER'S OBSERVATIONS SUMMARIZED BY MURCHISON

"Palseontological Memoirs," 1868, Vol. II, pp. 176-188

We owe to Murchison ("Palseontological Memoirs," 1868, Vol.

II,pp. 176-188) the complete enumeration of the numerous speci-
mens regarded by Falco!ior as belonging to the above species, as
observed in the museums of England, Italy, and Sicily; they
doubtless belong to many successive geologic horizons. Falconer
himself observed many variations in the 'lo.xodont sinus' or loop,
the thickness of the enamel, iho width and length of the crown,
and the greater or less crimping or undulation of the ridge-plates.
The typical 'Elephas aiitiquus' of Falconer appears to have been of
Lower Pleistocene, or Cromer Forest Bed age, as shown by the
fact that the collective ridge formida cited above agrees precisely
with that assigned to ("romer Forest Bed .superior and inferior
molars as cited below by Osborn.

-/?y 3 a

Fig. 1078. I'l'NOH Elephant (Hesperoloxodon antiquus). Second

SUPERIOB AND MoLABS OF SKELETON DESCRIBED BELOW
InI'BUIOII

One-third natural size

A, Crown view of twelve ridge-platcd .second right superior molar, r.M".

B, External view of twelve ridge-plated second left superior molar, I.M^;

anterior ridges i>artly worn.

C, Lateral view of second right inferior molar, r.M2, retaining ridge-plates

2-1 iK.
Thiki) Right Supekioh Moi.ak ok Hesi-kroloxodon ANTiguus D, Crown view of second right inferior niol;ir, r.Mj, retaining ridge-plates
Compare diagrammatic figuro 108S 2-11)^ estimated.

Fig. 1077. Rcfcrrt'd 16X> ridge-plat cd molar, r.M'^, of Elephas anliquus, After retouched original photographs by C. Forster Cooper.
after Falroner and Cautiey, 1846 [1847, PI. xiv.A,Erroneously
fig.s. 5, So). Observe that these molars wore identified as second superior and inferior
designated on Falconer's plate (xiv.a) a.s 'Elephas mrriiHonalis,' but corrected molars by ,\ndrews (191,"), p. 11), as third superior and inferior molars by
in his handwriting in copy of the "Fauna Antiqua Sivalensi.s" belonging to the Fonstcr Cooper (1028, (jp. 23, 24). The two upper molars, r.M", 1.M-, .show
Briti.sh Museum. Falconer defined this specimen as follows (18f>7, p. 22, 1868, twelve remaining ri(lg<'-plates with a ))osterior talon; they measure -|-221 mm.
Vol. I, p. 442, PI. xiv.a): "Figs. 5 and oo.— Last true molar, upper jaw, right in length, S3 mm. in breadth, l.')0 mm. in maximum depth; laminar frequency
side. Has j + l
fourteen plates and a heel, well crimped. From forest bed, .")
ridge-plates in 10 cm. The lower molar, r.Mj, measures +231 mm. in length,
Ostend, Norfolk. Green collection.— No. 16,229 B.M. Length, 10. in. Width, 70 mm. in breadth, 123 mm. indepth; laminar frefpiencj' in 10 cm. Loxo-
.")

3.4 in. Height, 6. .5 in." One-third natural size. Compare with the type of dont sinjis rudimentary. This r.Mo compares elo.sely with I'aleoner's tyjie
Elephas [Hesperoloxodon] anliquus (Fig. 107.5). I.M2 (Fig. 1075).
 : — — 6 : :

THE LOXODONTIN^: HESPEROLOXODON 1221

Falconer found no trace of successional premolars such as are CONFUSION WITH PARELEPHAS MOLARS, BY ADAMS,
observed in Archidiskodon pla?ufrons nor of the first lower milk LYDEKKER, AND MATSUMOTO
molar. The other priceless observations contained in Falconer's Leith Adams (1877-1881, p. 47), after careful consideration of
Note-book of 1862 are summarized by Murchison (op. cit., Pal. Falconer's work and ridge formulae, concludes as follows: "From

Mem., 1868, p. 176) as follows:

the foregoing details it seems to me that the ridge formula of
Elephas antiquus, as far as British specimens in particular demon-
"4. Elcphas (Euelephas) antiquus. . . . compiled from entries strate, is, [without talons], in upper and lower jaws, as follows:
. . .

in Dr. Falconer's Note-books. [Ed.]

Dp 2 lif Dp 3 lii Dp 4 M 1 M2 M3|ti
The distinctive characters of the teeth of Elephas antiquus
Leith Adams' error arose from his confusing the 'E. antiquus^
may be expressed in the following terms.
molars with those of the contemporary Parelephas.

1. Narrowness of the tooth in proportion to its length and Distinction from Parelephas trogontherii. —
Lydekkcr
height. (1886.2, p. 122), although making use of the same materials as
Falconer, erroneously assigns to E. antiquus a higher ridge formula;
2. Great height of the plates. The height is more than
because, like Leith Adams, he includes within the narrow-plated
double the width of the crown.
E. antiquus the 'broad-plated' molars belonging to Parelephas
3. Mesial rhomboidal expansion of the discs of wear. trongontherii. He observes: "The ridge-formula [Footnote:
'Slightly modified from Leith-Adams, "British
Elephants" Fossil
4. Great crimping of the enamel-plates.
(Mon. Pal. Soc), p. 176. If the talons be included the formula will
be higher, see Leith-Adams, op. cit., p. 231.'], excluding talons,
The dental formula of E. antiquus is as follows: may be represented as
(2-3)
Milk Molars. True Molars.
Mm. [Dp 2] [Dp3]-^ [Dp4]||^ M.[M1];;,
15-20)
3+6+10 10+1 2 + 16 )) [M2] [M3] (

(ie- 2 1)
:

3+6+1 10+1 2+1

The higher numerals in this Leith Adams and Lydekker formula

The comparative enumeration of the ridge-plate totals and probably pertain to Parelephas trogontherii, namely
of the characteristic formulae was also commented upon in Fal-
Ml 1 2
1 2
M2 M3
coner's posthumous Memoir of 1865. Observe throughout his
notes that the Similarly Matsumoto (1924-1926) erroneously considered that
the Palxoloxodon phylum represented by his 'Parelephas proto-
First upper milk molar = Dp^ of the present Memoir. mammonteus' was ancestral through the Parelephas trogontherii
phylum to the Elephas = Mammonteus] primigenius phylum, as
Second upper milk molar = Dp' of the present Memoir.
[

indicated by his choice of the specific name protomammonleus'

'

Third upper milk, molar = Dp* of the present Memoir. (cf. p. 1297 below).

PRIMITIVE AND PROGRESSIVE RIDGE-PLATE FORMULA

Research to the end of the year 1929, aided by the cranium from Pignataro Interamna, Italy (the type of Palxoloxodon {
= Hesperoloxo-
don] antiquus ilalicus), reveals (Fig. 1088) a progressive hypsodonty and ridge-plate addition from the Upper Pliocene to the 3d Inter-
glacial stage, when H. antiquus germanicus and H. antiquus italicus mark the extinction of this phylum in western Europe.

Progressive Ridge Formulae. —The progression from the typical Lower Pleistocene M 3 |f cited above, to the progressive ridge
formulae given by Zuffardi, Deperet, Soergel, and others, namely M
3 f^ cited below, does not represent contemporary variations; it
represents rather progressive or ascending mutations. As shown in the Osborn-Reeds diagram (PI. xxrv), Hesperoloxodon antiquus lived
foran enormously long period of time, perhaps hundreds of thousands of years, during which ridge-plates were constantly being added.
During the same long geologic period, as shown in figures 1072 and 1073, the ridge-plates were constantly broadening.

Forster Cooper cites the primitive and progressive ridge formulae (1924, p. 117), as given by more recent authorities, as follows:

Zuffardi, 1913: M 2^^ jyjQ ^'^ "^

l_5/20
16/21 ] Elephas antiquus
Deperet, 1923: M 2 f^Jf Q lyr
-"
15/20 +
16/21 \ [ = gertfianicus], M2 10/13 ^ -^ ''
14/lT Elephas antiquus mut. ausonius
Soergel, 1912: M 2 '"^'^ M 3 ff'^ I
mid-Pleistocene
1222 OSBORN: THE PROBOSCIDEA

He adds the following characters:

Elephas anliquun: "Thick, much folded plates regular rhombic or rectangular [S].

Fairly tliick, Loxodont sinuses always well marked [D].

always folded.
Folding not particularly noticeable, lozenges marked, a considerable difference in size between upper
and lower molars [Z].
Long straight high crowns [Z]."

Elephas anliquus As antiquus but more deeply folded, and with folds more numerous, a very regular feature.
mut. ausonius: Loxodont sinuses variable but on the whole stronger and more prominent [D],
Elongated straight crowns about % smaller than antiquus from Clacton, Saffron Walden, etc. [D]."

Osborn, 1930: The ridge formulae cited by Forster Cooper (1924, p. 117) and the specific references and identifications are super-
seded by the more precise ridge formulae and specific identifications in the present Memoir; the unique 'Elephas ausonius' (M 3 l-^)
does not occur in the Forest Bed, the mut. ausonius is probably 'Parclephas trogonlherii nestii'; the higher ridge formulae attributed to
Zuffardi, Dcpcret, and Soergel belong only with the mid-Pleistocene [Upper Pleistocene] progressive E. [Ilcsperoloxodan] antiquus
germanicus, as shown in figure 1088.

PiuMiTivE Ridge-Plate Foumul^ and Measurements

Ridge-plates Length Breadth
Cromer Forest Bed R. M' 16-17 280 mm. = 11 in. 90 mm. =3.50 in.

Happisburgh (Forest Bed) R. Ma -1-11 267 mm. = 10.5 in. 88 mm. =3.40 in.

Upnor Elephant (after

Forster Cooper) R. 83 mm. =3.25 in.

70 mm. =2.75 in.

E. antiquus type 66 mm. =2.60 in.

 :

THE LOXODONTIN^: HESPEROLOXODON 1223

and-white ilKistrations to Dr. Andrews' section are by Miss Gertrude M. Woodward; Mr. For.ster Cooper has made the
drawings for his own contributions.
From this Memoir the following citations (Andrews) may be paraphrased (pp. 2 and 3)

The fore-limbs are represented by a left scapula humerus, wanting most of the head, the tuberosities and the front of
; a left
the upper part of the shaft the upper articular end of the right humerus the left ulna, wanting part of the shaft a complete
; ; ;

right radius and the greater part of the left and most of the bones of the fore-foot on one side or the other except the cunei-
;

form and unciform.

[Comparison with African (Loxodonta africana) and Indian (Elephas indicus = maximus) Skeletons]
The scapula is preserved on the left side only. It is much crushed, the spine being bent backwards over the post-scapular
fossa. The upper angle is not quite complete, and a considerable portion of the posterior angle is wanting. The form of the
glenoid surface is most nearly like that of E. africanus; the coracoid tuberosity is rather more developed than in that species,

but than in E. maximus; it is much as in E. primigenius. Between this tuberosity and the anterior border of the glenoid
less
cavity there is a deep pit, such as also occurs in E. a.fricanus. A similar pit is also seen in the glenoid portion of a large scapula
(B.M. Geol. Dept. 21680) from Grays, probably belonging to E. anliquus. The humerus is an enormously massive bone,
. . .

The Upnor Straight-tusked Elephant, Hesperoloxodon antiquds, of the British Museum, as Mounted in 1927

After photographs kindly furnished by the British Museum (Natural History)

Fig. 1079. This skeleton was discovered in 1911-1912 in a trench near Upnor on the banks of the Medway in Kent, England; it was excavated in 1915
under the supervision of Dr. C. W. Andrews and Mr. L. E. Parsons; the restoration and mounting under the direction of Doctor Andrews and Mr. C. Forster
Cooper (made jiossible tlu-ough the generosity of Mr. Rushtcn Parker) were completed July 23, 1927. The above figures (side and front views) are repro-
duced, one-thirtieth natural size, from original photographs kindly sent the present author by Mr. Cooper (cf. Cooper and Andrews, 1928, frontispiece and
PI. I).

The restored backbone (as shown in Fig. 1084) is complete except for two vertebrae; the radius of the left fore leg is restored in plaster, the right radius
is complete; this radius, with some of the bones of the right foot, one of the tusks (too shattered for restoration), and a few grinding teeth are exhibited in
a case adjacent to this mount in the British Museum. Compare figures 1080, 1081, 1082, and 1084, also text pages 1222-1228 of the present Memoir.
1224 OSBORN: THE PROBOSCIDEA

being apparently stouter in iiroportion to its length than in E. maximus, E. ofricanus, and E. priniigeniiis; The upijer oncl . . .

of the mdins is much like that of the radius of E. niaximii.s; its anterior border is nearly straight, and the outer end of the
humeral surface terminates in a point on the outer edge of the outer lobe of the humeral surface of the ulna. Only the left . . .

ulna was found. This was nearly complete, wanting only a portion of the shaft immediately above the distal articulation, part
of which is likewise missing. Falconer and Lcith Adams have both commented on the relative stoutness of the limbs of
. . .

Elephas anlicjuus; probably it is the natural result of the enormous bulk attained by this species. On the whole, the radius . . .

and lUna together are very similartotho.se of £. o//-/rfl/i((sand also to those of £. ««/.;V7»(/s;e('A-/. The colk^ction at the British . . .

Mu.seum includes the upper half of an ulna from Grays as large as that now under description. ... Of the fore-foot the following
bones arc preserved; scaphoid (right and left), lunar (right and left), pisiform (left), trapezium (right and left), trapezoid
fright and left), metacarpal I (right and left), metacarpal II (right), metacarpal III (right and left), metacarpal IV (left
and part of right), phalanges and sesamoids. ... In the scaphoid the radial facet is nearly flat. and is more like the . . .

corresi)onding surface in E. africanus. In the lunar the facet for the radius (right) occupies nearly the whole upper surface;
. .

it is concave from before backwards posteriorly and concave in the same direction in front. The magnum is preserved on both . .

sides. Its upper articular surface is almost exactly as in E. africanus. The surfaces for articulation with the trapezoid
. . .

and unciform are almost exactly like those occurring in E. africanus, but the distal (metacarpal) surface is peculiar. The . . .

metacarpals are extraordinarily stout and massive. The third metacarpal is the largest; it is preserved on both sides.
. . . On . . .

the whole, the third and fourth metacarpals are very like those of the African elephant. Only the proximal halves of the
. . .

two femora are preserved, the remainder having been destroyed in digging the original trench. The head of the bone is
nearly hemispherical, and the neck is very short. There seems to be no trace of the lesser trochanter, and in this our
. . .

specimen resembles the femiu' of E. africanus rather than that of E. maximiis. The left libia is nearly complete, but the
. . .

I'rNOll .StiIAIOHT-TUSKKI) ElKTHANT, HeSPEBOLOXODON ANTICJUU8, OF THE BhITI.SH MuSKUM, IIFDKAWN to MHOW THK OKKilNAL AND nKSTORBI) I'AHTS
(oBwguE shading)
Fig. 1080. The scale of the skeletal liciEht to ttip of scapula (3700 mm. = 12 ft. l^j in.) asrees with the estimates of Bather, .\M(lnnvs, and Cooper. Tlie
obliciuc .shading indicates the restored portions of tlie .scapula, humerus, radiu.s, carpals, tarsals, and metatarsals, and hotli femora, for a new restoration of the
vertebral column by Osborn, sec figure 1084, UesperoloxotUm miliquun of Ilpnor. I'"or a new reconstruction of the skeleton and outline of the flesh of the Upnor
ele|)hant, see figure 1083. For a new restoration of the Ujmor straight-tu.sked (•lci)hant, sec figure 1074. For comparative details of the vertebra;, seapulsD,
and backbones, see figures 1082, 1081, and 1084. One-thirtieth natural size.
 THE LOXODONTIN^: HESPEROLOXODON 1225

right is by the lower end, which is much cracked and distorted.

represented only The astragalar surface is wider from before . . .

backwards than any specimen with which comparison has been made, but it comes nearest to what occurs in E. afriranus.
in . . .

Tho fibula of the left side is complete except for two or three inches towards the upper end of the shaft; The form of these . . .

surfaces is much as in E. afriranus. The astragalus ... is very low and broad as in E. antiquus recki; the tibial facet is scarcely
. .

perceptibly concave from side to side. On the whole, the bone is much like that of E. maximus and E. primigenius, and
differs from the astragalus of E. africanus in the greater development of what may be called the neck, which carries the facet
for the navicular; Tiie calcaneum is preserved on both sides; it is a very heavily built and massive bone.
. . . On the whole, . . .

oiu' specimen most nearly resembles the calcaneum of E. africanus, except that the fibular facet is smaller. The navicular is . . .

preserved on the right side only. It differs only in detail from

the navicular of E. africanus; The navicular of E. antiquus
. . .

., , , recki is very similar to that of the Upnor specimen, but the

^/4 Natural sije , i ,• .
r • i
•
. • •

astragalar lacet is wider trom side to side in proportion to

i i
-'

its antero-posterior diameter. The first metatarsal is not

. . .

preserved on either side, but judging from the distal facet on the
ectocuneiform it must have been of fairly large size. The . . .

third metatarsal is considerably larger than the second, being

A,B, SIVALIKIA ANTIQUA (UPNOR)

/t'.fl/ ELEPHA3 (MA,XIMUSI INOICUS

/5^ J)orsolumlar

Fig. 1081. Comparative Views of the Scapulae of Hesperoloxodon (syn.

Sivalikia), of Loxodonta, and of Elephas
After Andrews and Ct)oper, 1928, figs. 1 and 2
Hesperoloxodon antiquus of Upnor. Very large, broadly triangular, greatly
exceeding in .size the si^apula of 'Jumbo.'
Loxodonta africana ('Jumbo') of the Sudan (Abyssinia). Of smaller size
and of very similar proportions to the scapula of Hesperoloxodon antiquus of
Hesperoloxodon antiquus of Upnor
Upnor. Front and lateral aspects of (A) erect first dorsal vertebra, of
Fig. 1082.
Elephas indicus. Scapida elevated with relatively small pre- and [xist- (B) .sixteenth dorsolumbar vertebra of Hesperoloxodon antiquus, to the same
scapular fos.sa;; much less bro.adly triangular than either Loxodonta or scale as (A') first dorsal and (B') sixteenth dorsolumbar of Elephas indicus
Hesperoloxodon. ( =maximus), one-twelfth natural size. Compare Forster Cooper (1928, fig. 4).

especially wider at its upper end. The proximal surface has a considerable area of contact with the mesocuneiform in E. ;

africanus this contact is very small, and in E. maximus is wanting altogether. .In E. antiquus recki the third metacarpal . .

is said to have a distinct facet for contact with the mesocuneiform, and as in the Upnor specimen the fourth metatarsal

has a considerable articulation with the ectocuneiform. It thus appears that in that form, as in our specimen, there is a con-
siderable degree of alternation between the distal row of tarsals and the metatarsals; this alternation exists in a lesser degree
in E. africanus, but is not present in E. maximus.

Forster Cooper's Observations (1928, pp. 19-24). — In general contour and proportions the pelvis is similar to that of
Elephas africanus, except for its considerably greater size. The principal measurements of the pelvis as restored are:

mm.
Extreme width across the ilia at right angles to the vertebral axis 1830
Greatest width of ilium 1100
Width between the acetabula 945
Greatest diameter of acetabulum 250
Approximate length of symphysis 570
 ,

1226 OSBORN: THE PROBOSCIDEA

The vertebral column was found practically complete, but much weathered. The number of dorsal vertebrae actually
. . .

found was twenty; the specimen has been mounted so as to leave space for two more. The series runs in order from the atlas
to the twentieth dorso-lumbar, the missing ones belong, therefore, to the vertebral column just in front of the sacrum. [Dorso-
lumbars of E. africanus = 23-24, of E. indicus { = maxirnus) =22-23.] The alias .... which lacks the wing on each side, has
. . .

a somewhat low spine on the neural arch, whose pedicel on each side is pierced by a foramen through which the vertebral artery
was transmitted. The axis
. . . which likewise may be compared with a specimen belonging to E. ineridionalis (B. M.
. . .

27872, .
), shows again a .slight ditference in proportions, and, of the two, has a rather more slender neural arch.
. . While
both the atlas and axis are each of them a little larger than the corresponding bones of E. meridionalis, they are not so to the
extent that might be expected in an animal of such remarkable size in the rest of its skeleton. ... Of the other vertebrae the
most striking point is the unusual angle at which the neural spines are placed, both in respect to the individual vertebra and
to the backbone itself. There is, in various species, some variation in the curve along the back that is formed by the tips of the
neural spines. Dietrich has illustrated diagrammatically these curves for E. africanus, E. priniigenius, and E. maximus. From
lack of sufficient material the curve for E. antiquus has been so far unknown, nor does the present skeleton, owing to damage,
yield absolutely definite information, but, as far as can be .seen, it is not unlike that of E. maximus, where the top of the spine
of the first dorsal vertebra reaches approximately the same height as the top of the scapula, and from this point the spines grad-
ually ri.se in height to a maximum in the region of the eleventh or twelfth dorsal vertebrae, and thereafter fall away in height
gradually down to that of the sacral vertebrae. The Upnor specimen, however, shows one great difference, not only from
E. maximus, but apparently from other elephants, in that the neural spines of all the dorso-lumbar vertebrae stand bolt up-
right at right angles to the longitudinal plane of the vertebra. In other elephants all the dorso-lumbar spines except the most
anterior slope backwards at a considerable angle. The condition here is undoubtedly natural, is not due to postmortem changes,
and appears to have no parallel. The spines of the vertebrae in this region also show another peculiarity, in that those of the
posterior lumbar vertebrae are broad riglit up o the top (as seen in side view) instead of dwindling down to a point, which is the
I

more usual condition.

In the measurements and comparisons below of the Upnor skeleton assembled by Andrews (pp. 1-18) and by Forster Cooper (pp.
19-25) there is The resemblance of the parts of the Upnor specimen observed and measured to certain of the
repeatedly pointed out : (1)
referred specimens from Grays Thurrock, from Happisburg, and from other Lower Pleistocene localities; also to similar parts referred
by Dietrich to Elephas [Pala-oloxodoii] recki, from which it appears that we have to do in all these British specimens with a Lower
Pleistocene phase of Hesperoloxodon antiquus; (2) in comparison by Andrews and Forster Cooper, the resemblances of the scapula,
humerus, and other parts are observed to be closer to the African elephant (L. africana) than to the Indian elephant (E. indicus
in general
= maximus) : (Cooper, p. 24) "the Upnor specimen shows, when compared with other forms, a great mixture of specific characters. The
femur, for example, 'resembles E. africanus more than E. maximus' (p. 13), while the astragalus 'on the whole is much like that of E.
maximus and E. primigenius, and differs from the astragalus of E. africanus' (p. 15), and so on. The general result of Andrews' detailed
investigation is that in some characters the specimen resembles one form and other forms in other characters, with the result that a clear
diagnosis is not possible. The absence of any definite standard of comparison as well as the absence of any complete specimen of
. . .

E. antiquus prevents us therefore from estimating the true value of the two outstanding features of this LTpnor specimen, namely the
curious and apparently unique upright position of the spines of the vertebral column, and the unexpectedly small size of the teeth in
relation to the great bulk of the body. Should examination of further material eventually show that this form differs from the type
[ofElephas antiquus] so far as to warrant the erection of a sub-species, then it is to be hoped that the present specimen will be taken as
the holotype, and that the sub-species will be named after Dr. Charles W. Andrews, who did so much to elucidate the eariy history of the
l^roboscidea."

The vertebral measurements of Hesperoloxodon antiquus somewhat exceed those of Archidiskodon meridionalis. The mass of the
atlas and axis about equals that of A. meridionalis, while the entire vertebral column greatly exceeds that of E. indicus { = maximus).
Forster Cooper's descriptions and figures {op.
cit., 1928, pp. 19-25) show that the vertebral column of //. antiquus is unique. It differs
widely from both Loxodonta africana and Elephas indicus {=rnaximus). As illustrated in our new comparative figure 1084, it differs still
more widely from the backbone of the two mammontines Parelephas jeffersonii antl Mammonteus primigenius. The following are
Caudry's (1893.1, p. 19) measurements of A. meridionalis of Durfort:

Durfort (meridionalis)

Ground to top of head

Ground to withers [scapula]
Length from tu.sks to droj) of tail
Length without tusks
Humerus
Cubitus
Femur
Tibia
Third metacarpal
Third metatarsal
 THE LOXODONTIN^: HESPEROLOXODON 1227

Comparison of Vertebral Characters of Hesperoloxodon antiqiius of Upnor with Loxodonta afrioana, Elephas indicfs
( = MAXiMus), Parelephas jeffersonii, AND Mammontehs primigenius (Fig. 1084)

(Compare pp. 930-931 of the present Memoir on Vertebral Distinctions of Elephas, Loxodonta, Mammonteus, and Parelephas)

—
Comparison of Vertebral Column in the Loxodontin.e, Mammontin.e, and Elephantine. The vertebral columns of these
five types of living and extinct probo.scideans are carefully redrawn in figure 1084 from original materials; they exhibit very significant
resemblances between Hesperoloxodon and Loxodonta ( = Loxodontinae) also between Parelephas jeffersonii and Mammonteus primigenius
,

( = Mammontinaj), but the Elephantinae {Elephas indicus) differ widely both from the Loxodontinae and Mammontinse in the construction

of the vertebral column.

Hesperoloxodon antiquus of Upnor: Cervicals 1-7, dorsals 1-20, lumbars 1-3 est.; all dorso-vertebral spines vertically placed, very
much elevated. Orthogonal drawing after photographs and data by C. Forster Cooper.
Loxodonta africana: Cervicals 1-7, dorsals 1-20, lumbars 1-3; vertebral spines oblique, expansive; post-dorsal and lumbar spines
elevated. After 'Jumbo' {Loxodonta africana oxyotis), from the Setit River, Abyssinia.
Elephas indicus, Asiatic elephant: Cervicals 1-7, dorsals 1-19, lumbars 1-3. Based on the mounted skeleton in the American
Museum (Amor. Mus. Dept. Mam. 39082, known as "Samson" when alive) but enlarged to the record size of the Indian elephant.
Mammonteus primigenius, the woolly mammoth: Cervicals 1-7, dorsals 1-19, lumbars 1-4. Drawn after the Elephas primigenius
of Borna, dcrmany, as mounted by Felix, 1912, PI. viii.
Parelephas jeffersonii, the Jeffersonian mammoth: Cervicals 1-7, dorsals 1-19, lumbars 1-4. Drawn after mounted type skeleton
in the American Mu.seum (Amer. Mus. 9950).

Observe Parelephas the marked resemblance to Mammo7iteus primigenius in the extremely oblique and recumbent vertebral
in
we pass from the posterior dorso-lumbar region, in wide contrast to the Loxodontinae. Observe also
spines, the rapid decrease in size as
the general resemblance between Hesperoloxodon (the straight-tusked elephant of Upnor) and the typical African elephant of Abyssinia,
as well as the marked resemblances between the vertebral columns both in the formulae and the recumbent spines of Parelephas and
Mammonteus. For the vertebral formulae of the different genera, species, and subspecies of elephants, compare Chapter XV, pp. 930
931. The formulae shown above may be summarized as follows:

Cervicals
Hesperoloxodon (straight-tusked ele-
phant of Upnor)
Loxodonta africana, 'Jumbo' (Amer.
Mus. Dept. Mam. 3283)
Elephas indicus (Asiatic elephant)
Mammonteus primigenius (mammoth
of Borna)
Parelephas jeffersonii (Jeffersonian
mammoth)
1228 OSBORN: THE PROBOSCIDEA

COADAPTATION OF THK VERTEBRAL CoLUMN WITH THE SUPERIOR InCISIVE TuSKS OF THE ELEPHANTS
With Notes by Robert T. Hatt, 1931

As shown in figure 1084 there are very striking divergencies in the neural spines of the Elephantidae. A glance at figures 1084 and 868
shows that in the Mammontinae (including Archidiskodon meridionah's of Tiuriort, Parelephasjeffersoriii, and Ma)n7nonleuspr>?nigeniux)
the spines are relatively low and backwardly inclined, diminishing rapidly in size toward the pelvis, in wide contrast (as shown in Fig.
1084) to the Loxodontinae, in which the spines are relatively large and less backwardly inclined. In Loxodonta africana the posterior
dorsals (D^) rise to the same height as the anterior dorsals (D'); in the remotely related Hesperoloxodon antiqitus of Upnor the dorsal
spines are even larger and absolutely erect, as first observed by Forster Cooper. Between these two extremes (in the Elephantinse) we
observe Elephas indicus in which the spines are intermediate in size and elevated in the mid-dorsal region into a decided mid-hump,
conspicuous in aged Indian elephants.

It proves that this is a case of coadaptation of the neural spines with the divergent functions of the superior incisive tusks: (l)In Loxo-
donta the tusks are constantly used for digging and uprooting purposes; (2) in Elephas indicus they are less frequently used; (3) in

Mammonteits and Parelephas they are not used at all for uprooting purposes, because they soon become incurved and actually cross each
other in old age; thus the only strain put upon the incisive tusks is the weight of the ivory or dentine which they carry, whereas in the
Loxodontines the ivory tusks not only have an enormous weight but are subject to tremendous strains in uprooting large trees.

This interesting functional divergence has been analyzed by Dr. Robert T. Hatt as follows:

The spinal columns of the elephantoid Proboscidea differ chiefly in their height and in the horizontal or oblique angulation
of their neural spines. These spinal columns present three main patterns, which are:

(1) —
SuBFAM. Mammontinae. Spines of the interscapular region long and backwardly inclined. Caudad to this the length
of spines decreasing markedly to the middle of the back where and beyond which they are insignificant in size (Mammonteus).

(2) Subfam. Elephantine. —

Spines of the interscapular region long but less inclined than in the first type. Caudad to the
interscapular region the spines gradually decrease in length into the sacral region. The spines of the lumbar region are of greater
length and strength than those of the lumbar region of the Mammontinae. {Elephas indicus.)

(3) Subfam. Loxodontinae. —

Spines of the interscapular region nearly or quite vertical. The spines decreasing in length
to a minimum height near the middle of the back. Caudad to this the spines again become longer, reaching a secondary summit
on the latter third of the dorsal vertebrae {Loxodonta africana). The skeleton of the African elephant, "Jumbo," being very
badly mounted, the spinal column of this individual is in consequence misleading as to the spinal curvature in the African ele-
phant this error is corrected in figure 1084.
;

[Hesperoloxodon is evidently nearer to the Loxodonta type than to any of the others, but differs in the great vertical develop-
ment of the mid-dorsal spines. —
Editor.]

The following deductions by Dr. Hatt, concerning the observed differences in the spines with relation to muscular strains, are drawn
from a consideration of the mechanical differences that accompany such changed proportions.

(1) The height and massiveness of the spines in the scapular region are determined by the magnitude of the force acting
upon these bodies; this is in large part an age and sex difference, for strain increases with increased size of head, tusks, fore-
limbs, etc., and balanced or diminished strains in the caudad, pelvic, and hiiullimb region.

(2) The is dependent upon the development of the spines on the caudal third of the
inclination of these interscapular spines
dorsal vertehrw. Those animals with low spines in this region presumably have weak spinalis muscles. As a result the strain
imposed on the interscapular spines by head pull is not counteracted by equal muscular pull from the rear, and the spines are
inclined in order to throw the strain near the long axis of the spine rather than perpendicular to this axis. Such conformation
occurs in Archidiskodon, Parelephas, and Mammonteus. It is probable that the woolly manunoths {Mammonteus primigeniu.^)
were incapable of raising both forefeet from the ground simultaneously, for their skeletons present evidence that the chief ex-
tensor muscles of the back were weak.

(3) The modern elephants on the contrary have strong spinalis muscles. Strain upon the interscapular spines, as produced
by head pull, is counteracted by contraction of the spinalis muscles to the rear. These opposing muscular forces are best met
by vertical spines intercepting the forces at right angles.
 ail i/ao natural si^e

Fig. 1084. Vertebral columns of Hesperoloxodmi anliquus, Loxodonla africana, Elephas indicus, Mammonleus primigenius, and Parelephas jeffersonii, drawn
to a uniform one-twentieth scale.
Hesperoloxodon antiqiius of Upnor, modified after Forster Cooper, 1928, and from photographs and data subsequently forwarded to the jjresent author.
Loxodonta africana oxyotis ("Jumbo"), after original in the American Museum (Amor. Mu.s. Dept. Mam. 3283).
Elephas iiidiciis, after original in the American Museimi (Amer. Mus. Dept. Mam. 39082).
Mammonteus primigenius, after Felix, 1912, PI. vm. Original in the Leipzig Museum.
Parelephas jeffersanii, after original type skeleton in the American Museum (Amer. Mus. 99.00).
Observe the very marked differences in the curvatures of the back, indicated by the heights of the dorsal spines, which correspond closely with profile
photographs of the living Loxodonta and Elephas in the cave drawings and with our own restorations of Hesperoloxodon anliquus and Parelephas jeffersonii.

1229
1230 OSBORN: THE PROBOSCIDEA

Comparison of the Upnor and San Isidro Skeletons of Hesperoloxodon antiquus with the African and Indian
Elephant Skeletons
 .

THE LOXODONTIN^: HESPEROLOXODON 1231

cosi piccola mole; quindi propongo per essa la dcnominazione di Length of expo.sed incisive tusks, taper-
Ekphas (Euelephas) antiquus var. nana. ing, pointed, and somewhat curved
inwards towards the extremities .2350 . . mm. 7 ft. 8}i in.
Transverse breadth across premaxil-
laries [as compared with 857 mm.

(2 ft. 9/4 in.) in Hesperoloxodon anti-

quus italic us of Pignataro Interamna
(Fig. 1098)1 [860e] 2 10
Measurements of associated skeleton:
Maximum height of the scapula. . . 1020 mm. 3 ft. 4/4 in.

Maximum length of humerus 1170 3 10

Maximum length of ulna 980 3 2)^

Maximum length of right and left

femora 1440 4 8'4

Maximum transverse width across

the iliac bones of the pelvis. 2350 . . . [1350]' 7 [4] 8^ [5}^

Estimated height of the body in the

flesh (Graells) 3900 12 Vi
Estimated shoulder height in the
flesh (Osborn) 3828 12 6%e
Type of Hesperoloxodon antiquds nanus
Fig. 1085. Type figure of Elephas (Euelephas) antiquus var. nana Acconoi,
1880, Tav. IV, fig.s. Described by the author as "La corona del terzo
6 and 7. The combined vertical height of the scapula, humerus, and
molare." One-half natural size. This appears to be the po.sterior portion of ulna (3170 mm.) is somewhat inferior to that of the combined
a 12)2 ridge-plated second su|x>rior molar of the left side, l.M", with the three
anterior ridge-plates worn off.

Larghezza massima della corona .Mill' 52 . . [Length 2.1 in.

Altezza massima della medesima. " 49 . Height 1.9 "

Diametro' massimo della siiperficie

triturante della corona " 46 . Width 1.7 "

The associated fauna includes referred specimens of Bos

primi genius, Cams lupus, Meles taxus, Ursus spelxus.

Hesperoloxodon antiquus platyrhynchus

Graells, 1897
Figures 1047, 1068, 1069, 1086, PI. xxin

(?) Lower or Middle Pleistocene. San Isidro del Campo, near Madrid,
Spain.

The type of 'Elephas platyrhynchus' does not deserve specific

rank but is possibly a subspecies or geographic variety of the
typical Hesperoloxodon antiquus, as shown by the comparison of its
extremely broad premaxillary rostrum (Fig. 1086) with the ro.s-

trums (Figs. 1069, 1105) of H. antiquus of Germany and Italy.

Elephas platyrhynchus Graells, 1897. "Fauna Mastodologica
Ib^rica." Mem. Real. Acad. Cien. Exactas, Fis. Nat., Madrid,
Tome XVII, pp. 558-572. Type. A portion of the superior —
maxillary with molar in situ, right tusk, complete pelvis, humerus,
tibia, Horizon and Locality. Probably Lower or
etc. — Type of Hesperoloxodon antiquus platyrhynchus
Middle Pleistocene.San Isidro del Campo, near Madrid, Spain.
Compare Hesperoloxodon anliquus of Upnor (Figs. 1079, 1080,
Type Figure.—Graells, op. cit., Lam. xviii —see figure 1086 of the 1083, 1081, 1074)
present Memoir.
Fig. 1086. Type premaxillary rostrum with tusk, and portion of right
The following are the measurements obtained from the superior maxillary with M" in situ, of Elephas platyrhynchus Graells, 1897,
author's description (op. cit., p. 569): Ldm. XVIII, figs. 9a and 10, one-sixteenth and one-fifth natural size respectively.

'See footnote on page 1230.

1232 OSBORN: THE PROBOSCIDEA

scapula, humerus, and radius of the Upnor elephant (3450 mm.)- the Nerbudda, India, namely, l.M 3 jlw This ridge formula
The specific name platyrhynrhus is fittingly derived from the exceeds that typical of Lower Pleistocene Hesperoloxodon
the
Greek irXdrus, broad, and pvyxos, snout. Hesperoloxodon pkitij- antiquus (Fig. 1072A), namely, l.M 3 5. Thus the type of He.'<pero-
,

rhynchns is exceeded in width by the Pignataro Interamna speci- loxodon antiquus ausonius exhibits an exceptionally high ridge
men (Fig. 1098). formula (M 3 1 stVo) ^or an Upper Pliocene* stage.
This type was probably found in Lower Pleistocene levels of History.— In 1875, Forsyth Major (following Falconer, 1868)
San Isidro overlying older beds of Middle Miocene age, to which discovered in the Upper PUocene' of Italy extremely long molar
the horizon name San Isidro is appHed; it is fully described by teeth, supposedly related to E. [Hesperoloxodon] antiquus but
Graells on pages 558 to 572 and figured in Lam. xviii. In the which he gave the manuscript name of
specifically distinct, to

greatly broadened and flattened structure of its premaxillaries Elephas ausonius. Similarly in 1891, Pohlig ob.served teeth (in the
(.see Lam. xviii, figs. 8a. 9a) this animal is apparently related to LIpper Pliocene' of Italy) which he erroneously referred to his
Hesperoloxodon antiquus, as .shown in comparison with figure 1106. Forest Bed species Elephas (antiquus) Nestii.
Somewhat against this reference is the strong curvature of the This renders it certain that in the LTpper Pliocene' of Italy
superior tusks; in favor of this reference is the conclusion of there was a relatively small narrow-toothed variety of the larger
Graells (p. 568) that this species from the heights of San Isidro del Hesperoloxodon antiquus of the Forest Bed level, clearly distinguish-
Campo is either a synonym of antiquus or represents an indepen- ed by the name of Elephas [
= Hesperoloxodon] ausonius; with the
dent species. greater ridge formula:

'Elephas ausonius' : M3 ;3.„,,

Hesperoloxodon antiquus ausoniusMajor (MS.), 1875,
Verri, 1886, Dep^ret and Mayet, 1923
Geologic Level. — Dep6ret and Mayet (1923, p. 162) and
Falconer (1868, Vol. II, p. 187) also describe Elephas antiquus as
Figures 1041, 1069, 1072, 1073, 1087
found in the marine PUocene of Rignano and in the lacustrine
Upper PUocene of Italy, Villafranohian stage,* San Romano, Val d'Arno
inferieur, also San Paolo de Villafranca.
Compare Upper Pliocene molars referred by Falconer to Elephas antiquus,
by PohUg to Elephas (antiquus) Nestii, by Sabba Stefanescu to Elephas antiquus
ausonius from Colintina (Ilfov), Rumania, not to be confused with the referred
E. antiquus ausonius of the Forest Bed, England.

The extremely narrow nineteen to twenty ridge-plated type

molar (Fig. 1087) of this Upper Pliocene species is very distinct from
the sixteen and a half to seventeen ridge-plated typical lower
molars of the Lower Plei.stocene 'Elephas antiquus' of Falconer, as
fully defined above. It has been erroneously confused with the
doubtful Lower Pleistocene subspecies 'Elephas (antiquus) nestii'
from which it is also quite distinct [ = Parelephas? trogontherii
nestii]. Falconer's observatioas of 1868 and Major's observations
Type of IIesi'euoloxodon a.ntiquds ausonius
of 1875 on the distinctness of this Upper PUocene stage were
Fig. 1087. Type of Eleplms ausonius Major. After Deperet and Mayet,
confirmed and extended by Dep^ret and Mayet in 1923, who "Fig. 1 et 2.^Ekphas auso?iius, de San
1923, PI. X, figs. 1 and 2, p. 220:
adopted Major's manuscript name of ausonius. Romano (Val d'Arno inferieur), pieces types de I'espece. M3 droite et gauche.
Specific Characters (Osborn, 1928). An Upper Pliocene — (Voir p. 166.) Institut geologique de Florence. Photographic du professeur
stage of Italy, distinguished from Hesperoloxodon antiquus by its Stefanini." Third Iowit molars of the right and left side. About one-fourth
natural size.
smaller size and extremely long and narrow third inferior molars
with a large number of ridge-plates, namely, j-^; The MB .

comparative measurements of the type M3 in millimeters are as Pliocene of Ast^san, at San Paolo de Villafranca: ". . . two last
follows: upper and two last lower molars, also from St. Paolo: . . Each
.

H. ausonius H . antiquus of the upper teeth consists of nineteen plates, the rear part being
Type M3 (typicus) M^ broken off. ... Of the lower molars, the right shows twenty plates,
Villafranchian Forest Bed and is very narrow for its height."
Length of crown 240 254 Forsyth Major (1875) first observed in the upper layers of the
Width of crown 74 85 Val d'Arno inferieur at San Romano teeth related to E. antiquus
Height 160 166-174 but sufficiently distinct to be separated under the manu.script name
of "Elephas ausonius." Dep6ret and Mayet note that F. Major's
Laminar frequency of 5-6 in 100 mm. Enamel coarse and muncr- manuscript name, cited by Weithofer (1891 [1890), p. 194), quoted
ously crimped. Lo.xodont sinus generally .strong. by Verri (1886) from labels in different mu.'^cums, deserves to be re-
The specific ridge formula (M 3 r/^iVo) agrees with that of vived and retained. Pohlig (1891, pp. 303, 350), while also clearly
a referred specimen of Palseoloxodoii namadicus (Fig. 1072D) from discerning the differences between the Upper Pliocene form and the

'[See footnote 1 on page 1049 above regarding the possible Lower Pleistocene age of the Vilkfranchian. —Editor.]
 THE LOXODONTIN^: HESPEROLOXODON 1233

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 .

THE LOXODONTIN.^: HESPEROLOXODON 1235

Pohlig's monographic descriptions of Taubach grinding tcetli, Specific Characters. — Comparing this type specimen with
selecting the most characteristic specimens he has described and third superior and other molars of 'Elephas antiquus' of Taubach
figured (Fig. 1088) as neotypcs of Elephas [
= Hcsperolo.rodon] and Weimar figured by Pohlig (1888-1891, Taf. iii bis, fig. 7a
antiqu us german ic us (explanation of plate, p. 272), and Taf. vi, fig. l'— Fig. 1088) we
In brief, Stefanescu designated as 'Elephas aidiquus german- record the following dimensions, after Pohlig and Osborn, of the
icus' the Upper Pleistocene mutation of Weimar and Taubach. neotypes of germanicus.
He designated Weimar and Taubach (Allemagne) as corresponding
with the geological horizon of the type. The type itself, however,
comes from Tanganu (Ilfov), Rumania (Fig. 1089). In the .same
notice Stefanescu characterized this subspecies as follows:
Ekphas antiquus gernianicus S. StefSnescu, 1924. "Sur la
presence dc I'Elephas planifrons et de trois mutations de VElephas
antiquus dans les couches geologiques de Roumanie." Compt.
Rend. Acad. Sci., Paris, Tome 179, p. 1418, December 15, 1924.
—
Type. Fractured crown of a second inferior molar of the right
side, r.Mj, in the Laboratory of Geology, University of Bucharest.
—
Horizon and Locality. Type from Tanganu (Ilfov), Rumania.
Upper Pleistocene horizon of Weimar and Taubach, Germany.
Type Figure.— S. Stefanescu, 1927.
—
Type Description. (Sabba Stefanescu, 1924, p. 1418, also
1927) The brief description of the subspecies gernianicus is quoted
:

with the context imder Archidiskodon planifrons rumanus in

Type of Hesperoloxodon antiquus

germanicus
Fig. 1089. Type of Elephas antiqu-
us germanicus S. Stcfiincscii, 1924.
Aft<T SU-fanoscu, 1927, Fig.s. A and B of
plate.

(A) Po.storior half of a .socoiid in-

feriormolar of the right side, r.M2,

exposing 11 posterior ridgc-plates (L.
1-11); 7-8 anterior ridge-plates wanting.

(B) Six intermediate ridge-plates of the same inferior molar, r.Mj:

'Tace tritnratriee dc la eouronne dc M' (Mo] d'Elfphas antiquus germanicus
(A) et de ses .six lames antcricures (Li-Le) detaehees (B): talon postcrieiir
(Tp), deux lames (Lu-Lio) depouillees de cement, deux lames (Lo-Ls) en-
veloppocs de cement, sept lames (L7-L1) entamees par I'erosion."

Chapter XVI, pj). 968, 969; it is also fully embodied in the follow-
ing sentence "Je n'insiste pas pour le moment sur les caracteres de
:

cette mutation qui, a mon avis, est la plus rajiprochee de I'origine

mastodontide de I'espece antiquus. J'ajoute sculement que la
mutation ausonius de I'ltalic lui succede et que la mutation de
Weimar et Taubach (Allemagne), que jc de.signe sous le nom
germanicus, est la plus recente."
1236 OSBORN: THE PROBOSCIDEA

TUSK OF HESPEROLOXODON ANTIQUUS GERMANICUS IN FIELD MUSEUM OF

NATURAL HISTORY, CHICAGO
[A decade ago Dr. Henry I'ield acquired by purchase for size than a male tusk from Tonna, Germany, preserved in the
Field Museum of Natural History an 'Elephas antiquus' tusk Gotha Museum, which, according to Dr. Hans Pohlig (1888, p.
[ = Hespeioloxo(lon
antiquus gcrmanicus], found at Steinheim on the 49, Taf. 1 — Fig. 1090 of the i)resent Memoir), mea.sures 2950 mm.
Murr, Germany, which at that time was supposed to be the in length or 9 8/s ft.Furthermore, Dr. Fritz Berckhemer
in.

largest of the antiquus species in existence. While this tu.sk mentions (1930.1, Abb. 4) an isolated tusk in the Stuttgart
measures 2925 mm. or 9 ft. 7}g in. in length, it is slightly smaller in Museum of even larger prouortions, namely, 3390 mm. or 11 ft.

292.') ram. 29.50 mm. 1820 mm. 3.">00 mm.

Fig. 1090. Male and Female Tusks of Hespeboloxodon antiquus (Right) Right male tusk (Stuttgart Mus. 16274) found in 1929 at Steinheim
GERMANICUS on the Murr, Germany, measuring 3.500 mm. or 1 1 ft. !)% in. Photograph kindly
All figures one-twentieth natural size. Compare figure 1106, Hesperoloxodon sent by Dr. Fritz Berckhemer.
antiquus ilalicus, one twenty-fourtli natural size Doctor Berckhemer writes May 1.5, 1937) that the largest tusk of
(letter.

(Left) Riglit tusk (Field M>is. Nat. Hist. 20161()) excavated at Steinheim 'J'Jlcphas antiquus' in the Stuttgart Museum collections measures about 3750
on the Murr, Germany. I^engtli on inner eurve 2770 mm. or 9 ft. 1in., on mm. or 12 ft. S% in.; the longest tusk recorded by Pohlig (op. dt., p. .51),
outer curve 292.') mm. or 9 ft. 7% in. Diagrammatic sketcli fumislicd through excavated from the "schottcrn" (rubble) of the ancient river Arno at the
the courtesy of Dr. Henry Field of Field Museum of Natural History, Chicago. Porta S. Lorenzo, is in the University of Rome and measures nearly 3900 mm.
(Center) Male and female t\i,sks unearthed at Tonna, Cermany, preserved —
or 12 ft. 9)2 ill. [ = Hesperoloxodon antiquus ilalicus of Rome see Fig. 1068
in the Mu.seum of Gotha. After Pohlig, 1888, Taf. i, figs, la, \b, 2, 2a. Compare above). The male tusks of Hesperoloxodon antiquus italictts (Fig. 1106)
figure 1063 (Loxodonta africana). The male tusk (Taf. i, fig. 1) measures measure 3070e mm. or 10 ft. Jg hi., within the alveolus 840-845 njm., free

29.J0 mm. or 9 ft. sYs in.; the female tu.sk (Fig. 2) 1820 mm. or 5 ft. 11^2 in. length beyond alveolus 2230- 2270 mm., a total of about 10 ft.
 THE LOXODONTIN^: HESPEROLOXODON 1237

5% in., and in a letter of May 15, 1937, he states that the largest Rhinoceros merckii and Bison priscus. This specimen was pur-
tusk in the collection found at Steinheim measures about 3750 mm. cha-sed by Dr. Henry Field, leader of the Marshall Field Archaeo-
or 12 ft. 3% in. logical Expedition to Western Europe (1927-1928), from Dr. F.
Doctor Field very generously offered Professor Osborn the Krantz, Herwarthstrasse 36, Bonn, Germany, who very kindly
privilege of first description in the present Memoir of the Field supplied the above information.
Museum specimen, an opportunity of which he intended to avail length of the tusk on the inner curve is 2770 mm. or 9 ft.
The
himself, but, as in other instances, his regrettable death prevented 1 in.,on the outer curve 2925 mm. or 9 ft. 7% in., along the pro-
the consummation of his plan. The following description has been jection of the base 2850 mm. or 9 ft. iVi in., and the maximum
prepared with the kind assistance of Doctor Field, supplemented circumference is 520 mm. or 1 ft. 8% in. The curvature of the tusk
by the diagrammatic sketch of the tusk (Fig. 1091). — Editor.] is clearly shown in the accompanying sketch, which is reproduced

herewith one-twentieth natural size; "the dotted lines indicate

the probable terminal extension and the depth of the pulp cavity."
Right Tusk of Hesperoloxodon antiquus germanicus in the The alveolar portion cannot be determined. The ivoiy is in fair
Collections of Field Museum of Natural History, condition, although the tusk was broken into four sections. ".

Chicago The tusk has been repaired by Mr. L. L. Pray (who also made
F.M.N.H. specimen No. 201616. Right tusk excavated at the diagrammatic sketch, reversed in this figure) and is now on
Steinheim on the Murr, Wiirttemberg, by the late Professor E. exhibition in the Hall of the Stone Age of the Old World (Hall C)
Fraas, Stuttgart, from intcrglacial river-sands associated with in Field Museum of Natural History.

(A) "•:,

flfC-^

fs]

• Jh^^t ^i-^*** fr" lA-^**^ '

Pig. 1091. Lateral and vertical views of right tusk of £?iep/tas o»i(i5« H.s [=//esperoioio(iori. antiquus germanicus] exacavated at Steinheim on the Murr,
Wiirttemberg, by the late Professor E. Frax'*, Stuttgart, from interglacial river sands, associated with Rhinoceros merckii and Bison priscus. One-twentieth
natural size. Compare figure 1090 (left). Sketch by Mr. L. L. Pray, reproduced through the courtesy of Dr. Henry Field.
1238 OSBORN: THE PROBOSCIDEA

HESPEROLOXODON ANTIQUUS ITALICUS OF SOUTHERN ITALY AND HESPEROLOXODON

ANTIQUUS GERMANICUS OF NORTH CENTRAL GERMANY
A flood of light on the cranial structure of the 'ancient' or 'straight-tusked' elephant of Europe has resulted
from the discovery in 1911-1912 at Pignataro Interamna, near Cassino, Italy, of the skuU and jaws herein de-
scribed as Palaeoloxodon [
= Hesperoloxodon] antiquus italicus, a subspecies of the classic 'Elephas antiquus' of
Falconer.

During the years 1926 and 1928 excavation in the region of Steinheim a.d. Murr, Germany, yielded to the
Stuttgart Museum one exceptionally perfect cranium and another less perfect cranium which were finally restored

Restoration of Hespeuoloxodon antiquus italicus

Adult but not full Krown. Slioukl'T liciKlit (fully adult) estimated at 13 ft. (i in. (cf. p. 12.")() below). Oiie.-fiftietli Natural siii(\

V'iK H)i)2. Tlii.s re.stdratiou of I'dLroLixodmi, \=Uixi>rrtili)xn(lnn] aniiquua iVrifeiis is based entirely on the type erauium (.\mer. Mu.s. 22(J3t) combined with
the skeletal eharaetcr.s of the Upnor 'straight-tusked' elephant Etrphas [Ilcspcmloxodon] aniiqum. Compare figures 1079, 1080, 1096, 1098, 1105, 1099, 1100,
and 1083 of the present Memoir, with accompanying skeletal mea.suremcnts and comparisons (pp. 1245-1252). The relatively small ears are drawn from
outlines by Pateolithic artists of North Africa and of Spain represented in figure 1047. Both in frontal and lateral aspects this 'straight-tusked' elephant
is widely different from the recent African elephant. After crayon drawings by Margrct Flinsch, 1931.

and reconstructed by Dr. Fritz Berckheiii(>r in 1929-1930, and herein referred to llesperoloxodnn antiqaus gcrmun-
icus of Stefanescu and Pohlig.

Thus within a period of twenty-years (1911-1931) there came a most welcome mass of new cranial knowledge
with a very important bearing on the evolutionary history and relationsliips of Falcon(>r's species 'PJIephm
antiquus.' As shown in the preceding pages of this chapter, the 'ancient' or 'straight-tusked' elephant has previ-
ously been known by isolated portions of the cranium, jaws, grinding teeth, and tusks, which can now be con-
sidered in their mutual mechanical relations, especially in the Italian specimen.
 THE LOXODONTINiE: HESPEROLOXODON 1239

LOXODONTA AFRICANA ALBERTENSIS, AFTER DRAWING BY MaRGRET FlINSCH, UNDER DIRECTION OF AUTHOR. OnE-FIFTIETH NATURAL SIZE
This combination drawing of a Central African elephant is based upon the bull elephant in the Carl E. Akeley group (Amer. Mus. Dept.
Fig. 1093.
—
Mam. 54085 see Fig. 1052 above) collected in 1909 by Mr. Akeley in the Budongo Forest, east of Lake Albert, Unyoro, Northern Uganda, also upon the
—
Sudanese elephant "Khartum" (Loxodonta africana oxyotis Fig. 1053), formerly living in the New York Zoological Park, and upon a photograph by Marius
Maxwell showing an elephant drinking at the river border.
The tusks (measuring 8 ft. 5/2 in. and 8 ft. 9^ in. respectively and weighing 1 12 and 1 15 lbs. each) of a bull sliot by Mrs. Akeley on the slopes of Mt. Kenya
{Loxodonla africana peeli — Fig. 1059) furnished the basis for those in the present restoration. The longest tusks of the African elephant on record are the lyre-
shaped pair in —
the Heads and Horns Collection of the New York Zoological Park (L. a. oxyotis Fig. 1062), which measure 11 ft. 5^ in., circumference 18'^ in.
with a combined weight of 293 lbs.
(right), 18/8 in. (left), The heaviest pair (although not the longest) recorded may be seen in our figure 1065, with a com-
bined weight of 461 lbs. The tusks of L. a. oxyotis (Fig. 1056) were restored after these two record pairs.
The large ears are drawn in extended position for contrast with the relatively small, low-set ears of Hesperoloxodon antiquus italicus (Fig. 1092).
The height at the shoulder in the flesh, namely, 11 ft. 6^ in. or 3520 mm., is after Ward's skeletal record (1928) of 10 ft. Q}i in. or 3290 mm. (cf. caption to
Fig. 912, Chap. XVI, above).

HESPEROLOXODON ANTIQUUS ITALICUS OF

SOUTHERN ITALY
Twelve have been recorded in which discoveries by
localities
ItaUan paleontologists of more or less perfect remains have been

made in this region since the first note by Oronzio Costa in June,
1864.
In July, 1926, the cranium and jaws of the 'Pignataro In-
teramna' elephant were discovered and exposed by a farmer,
Saverio Tiseo, while excavating for building purposes, in the extra-
ordinarily perfect condition shown in figure 1096 and most for-

tunately reported to Professor Giuseppe Dc Lorenzo, Director of

the Institute of Geology Naples and a member
of the LTniversity of
of the R. Accademia Nazionale dei Lincei. De Lorenzo
Professor
promptly made a preliminary communication (1926) on this mo.st
important discovery, and in the following year (1927) published,
with the cooperation of Professor Geremia D'Erasmo, also of the
University of Naples, a superb memoir entitled "L'Elephas anti-
quus neiritalia Meridionale" this memoir (pp. 1-35) affords
;

a valuable review of the discoveries previously made in the valley

of the river Liri (see Fig. 1094) in the following localities:
Upper Pleistocene Horizon of Hesperoloxodon antiquus italicus
CastelUri Casalvieri Isoletta
Compare Osborn, 1931.846, p. 4, fig. 3, also figure 1097 of the present Memoir.
Arpino Roccasecca Pontecorvo
Fig. 1094. Valley of the Liri River displaying the principal exposures along the
Fregellae Cassino (grotto) Pignataro Interamna eastern and western banks and slopes of the bordering hills, varying (nmi 60 to 70
Ceprano Aquino Caianello meters above the present sea level, of a tliickness of about .50 meters, wlierc the re-
mains of Hesperoloxodon antiquus italicus, of Hippopotamus, of Cervus, and of other
Geologic Age. — According to recent estimates of Pleistocene Pleistocene animals have been found. After De Lorenzo and D'Erasmo, 1927, p. 7,
time, a 500,000-year interval elapsed between the typical Lower fig. 1. Upper portion only. .Scale 1:3,000,000.
 :

1240 OSBORN: THE PROBOSCIDEA

Plc'istoceiip 'Elephas aidicjuus' Falconer and the new subspecies DESCRIPTION BY DE LORENZO AND D'ERASMC)
herein described, which is even somewhat more progressive than De Lorenzo and D'Erasmo (1926, 1927). On pages 35 — to

Pohlig of Taubach-Weiniar. The more 39 (see also Tav. i) of the De Lorenzo and D'Erasmo Memoir of
tlio 'Elephas gei-manicris'
or less fragmentary mammalian remains found in proximity to the 1927 is given a complete description of this superb cranium in its

type include the following (see Fig. 1095)

original state (as cited in full from De Lorenzo's original contribu-
tion of 1926, pp. 185-188), of which the following is a literal
(1) Cervus elaphus Linn. Three antlers from the left side,
: (a)
translation in part':
(b) two antlers from the right side, and (c) right astragalus. Stag.
"In the past month of July [1926], the farmer Saverio Tiseo,
(2) Bos primigenius Boj.: (a) Portion of left Mi and (b) frag-
of Pignataro Interamna near Cassino, excavating, for building
ment of mandible. Primitive ox. jjurposes, a piece of ground on his farm situated on the southern

(3) Palasoloxodon [Hesperoloxodon] antiquus italicus: (a) Frag- slope of the hill which borders the village and is really in Fon-
ment of r.M:, (b) symphysis of mandible, (c) a nearly complete tanarosa, found, at a depth of about 8 metres, a large cranium of
female tu.sk, (d) right .scapula (probably belonging to the type), and a mammifer."
(c) central portion of a right humerus. Ancient elephant. "The which the fossil in question was found forms,
locality in
with the hill Interamna, part of this whole sy.stem of
of Pignataro
(4) Hippopotamus amphibius Linn. (Pleist. = //. major
gently rolling hills, which extend from Aquino and Pontecorvo
C'uvior): (a) Complete mandible, (b) cervical 5, (c) left meta-
along the left bank of the Liri as far as the river Rapido below
tarsal IV, and (d) left femiu- (juvenile). African hippopotamus.
Ca!5sino, and which are made up of large alluvial deposits of the
(5) Rhinoceros merckii Kaup: (a) Right radius and (b) cervi-
early quaternary, deposited first among the chain of the Aurunei
cal 7. Merck's rhinoceros. and that of the Mainarde, later moulded by backwaters, defluents
(6) Associated with the manunal fauna was the imperfect shell in lesser volume of the present and more confined course of the

of a freshwater mollusc, referable to the genus Unio, of the family river Liri, which still today, as in the day of Orazio, continues with
UnionidJB. its calm waters the taciturn corrosion of its plains."

.\l.\.MM.\i,i.\N Fossils Associ.vted with the Tvpe Cii.\.\iUM oi' Htsi'BHOLoxoDON ANTiguus italicus
About, one-thirteenth natural size
After Osborn, 1931.846, p. 14, fig. 11

Fig. 101)."). Type right, srapula i'Sd) of I'alsioloxodon [Ilcsprrnloxodon] antiquum italicus.
(3a, 6, c, e) Fragment, of riglitsecond inferior molar (r.Mj), symphysis of mandible, nearly complete female tusk, and central portion of right liumerus of
I'alseoluxodon [= Hesperoloxodon] antiquus italicus, (3d) right scapula (i)robably belonging to type).
(4a, b, Mandible, .'jth
c, (/) cervical, Mts. IV, juvenile femur of Hippopotamus amphibius major roX.
(Jm, h) Right radius and 7th cervical of RIniutceros merckii ref.

(2a, b) Portion of left inferior molar (Mi) and fragment of mandible of Hos primigenius.
(la, b, c) Portions of three antlers from the left side, two antlers from the riglit side, also right astragalus of the stag Cervus elaphus.

'Kindly prepared by Miss Francesca LaMonte of the Department of Ichthyology of the American Museum of Natural History.
 THE LOXODONTIN^: HESPEROLOXODON 1241

PiGNATARO InTEBAMNA CraNIDM (TyPB) BEFORE REMOVAL, SHOWING (lEFT) THE ORIGINAL OwNBR SaVBRIO TiSEO
After Osborn 1931.846, p. 2, fig. 1

Fig. 1096. Type craniumin situ of Palseoloxodon [Hesperoloxodon] antiquum italicus (Anier. Mus. 22634) as found and exposed by Saverio Tiseo at Pignataro
Interamna, near Cassino, Italy, and measured and described by Giuseppe De Lorenzo in 1926 and 1927. Compare Tav. i, figs. 1 and 2, of the Memoir of
1927 by De Lorenzo and D'Erasmo.

"The present plains, formod in recent times by the Liri, extend

below Pignataro Interamna to about twenty metres above sea
level; while the hills mentioned above, formed by the early
quaternary diluvium, vary between 60 and 70 metres, thus giving
a thickness of about 50 metres to the uncovered Pleistocene ground.
This territory, prevalently clayish in the lower parts, becomes on
—
top sandy, yellowish sand and gravel interspersed with layers of
clay and mud, and, on top, volcanic cinders."
"In this early quaternary district some time ago there were
already found remains of fossil mammifers, especially elephants.
These have been preserved in part in the Museum of Geology and
Palaeontology of the University of Naples. Oronzio Costa first
noted them in the Rendiconti della Reale Accademia di Scienze
fis. e mat. di Napoli for June, 1864. They were fully described by
Giustiniano Nicolucci in his memoir Su gli elefanti fossili della

Valle del Liri [Concerning the fossil elephants of the Valle del
Liri] (Memorie della Soc. ital. delle Scienze, detta dei XL, vol. IV,

1882). Cacciamali contributes further to this in the BoUettino

della Societa geologica italiana, 1890, describing some molars of Upper Pleistoce.n'e Horizon of Hesperoloxodon antiquus italicu.s

the elephants of Val di Coniino and of Aquino. Finally, they are Compare Osborn, 1931.846, p. 3, fig. 2, al.so figure 1094 for details

mentioned by Antonio Weithofer in his memoir on the fossil Fig. 1097. Pignataro Interamna is near Cassino (circle), southwestern
Italy, about fifty miles north of Naples. Region of the Valley of the Liri
Proboscideans of Valdarno (a memoir which serves as the descrip-
(Liris) occupied in Pleistocene time by large herds of the 'ancient' or 'straight-
tive matter for a geologic atlas of Italy, vol. IV, part 2, Firenze,
tusked' elephant now known as Hesperoloxodon antiquus italicus, also by
1893). Hans Pohlig also speaks of this in his big monograph on Hippopotamus, Cervus, Bos, Dicerorhinus, and other species of 3d I nterglacial
Elephas antiquus, published in the Nova Acta Academiae Caes. time. After PI. 94 of the Century Atlas, edition of 1913.
1242 OSBORN: THE PROBOSCIDEA

Leopold. Carol. Gei-manicae naturae curiosorum, vol. 53, Halle,

1889, and vol. 57, Hallo, 1892."
"My own and the observations of others are gathered together
in my 'Geologia e Geografia fisica dell'Italia meridionalc,' Bari
(Laterza), 1904. On page 157 I spoke of the certain existence of
Elephas (Euelephas) antiquus Falc. in the early quaternary deposits
of the Valle del Liri."
"The fact that remains of fossil elephants already existed in
the Valle del Liri does not diminish the importance of the present
discoveries at Pignataro Interamna; of really exceptional impor-
tance because of the completeness of the exhumed cranium and
because of its position, a position which leads to the deduction
that it was found in its original posture, not a secondary one caused
by transportation, and this leads to the hope that it may be con-
nected with the rest of the animal's skeleton."

[857J

Db Lorenzo's Original Sketch and Measurements of the Type Cranium

OF Hesperoloxodon antiquus italicus
Compare Osborn, 1931.846, p. 8, fig. 7

Fig. 1098. — Diagrammatic sketch, prepared in the American Museum to

aid in the eighteen month.s' process of reconstruction. Cranium of Palseoloxo-

don [Hesperoloxodon] antiquus italicus (Amer. Mus. 22634), based upon original
photographs (Fig. 1096), showing the exact measurements recorded in De
Ix)renzo's contribution of 1926 and f\illy quoted in his Memoir of 1927. The
American Museum reconstniction, completed November, 1930, in front view-
accords exactly with the 1926-1927 measurements of De Lorenzo, except as to
width of rostrum [857 mm., 520 mm.l, as follows:

Apex of right tusk to vertex of cranium 5500 mm.

Vertex of right tusk to border of i)remaxillary socket 2200
Lower border of premaxiDary socket to vertex of
cranium
Diameter of incisive tusk at exit from socket
Transverse breadth across premaxillary sockets
Space between inner sides of incisive tusks
Transverse space across rostrum just below orbits
Midline of premaxillary rostrum to midline of nasal
opening
Transverse across narrowest portion of frontals
 THE LOXODONTINiE: HESPEROLOXODON 1243

form of the dijese [framework?], show clearly that the

selves, the $1,000.00 by Henry Fairfield Osborn, mounted step liy step to
cranium from Pignataro Iiiteramna belongs to the species Elephaf; $4,375.34, almost nine times the amount originally set aside for
[Euelephas) aidiquus Falconer, characteristic of the interglacial the purpo.se.
l)hases of the early quaternary and the largest of the few species of On
opening the boxes containing the fossil, the extremely
elephants which have inhabited the earth. The fortunate dis- painful discovery was made that between 1927, when the negotia-
covery of this complete cranium further shows that, contrary to tioas of Ti.seo with De Lorenzo were concluded, and 1929, the
what Pohlig wrote, Falconer was right in maintaining that his owner Saverio Tiseo had irretrievably damaged the entire upper
Elephas antiquus was closely allied to the living Indian elephant,
Euelephas indirus."
The above from G. De I^orenza and G. D'Erasmo
citation
(1927), pages 35-39, figures 10and 11, and plate i, figures 1 and 2,
affords invaluable information as to the Pignataro Interamna
cranium in its original undisturbed condition partly buried in the
matrix. These drawings and photographs demonstrate the superb
condition of the cranium and tusks when first exposed and our
knowledge is fortunately amplified by two photographs (repro-
duced in our Fig. 1096) subsequently taken by Saverio Tiseo before
he attempted to remove this priceless fossil from its original bed in
the matrix. The precise knowledge of the specimen in its original
condition afforded by these four photographs, also by the measure-
ments and outline sketches by De Lorenzo reproduced in our
figures 1099 and 1100, is in close accord with the measurements in
De Lorenzo's original paper of 1926, pages 187 and 188, as re-

De Lorenzo's Original Sketches of the Type Craniu.\i of Hesperoloxodon italicus

One-fortieth natural size
Fig. 1099.Pignataro Interamna cranium (Amer. Mus. 22634) in situ. Fig. 1100. Front view of the Pignataro Interamna oranium (Amer. Mu.s-
After De Lorenzo and D'Era.smo, 1927, p. 37, tig. 11 "Cranio dell 'El. antiquus
: 22634) in situ.After De Lorenzo and D'Era.smo, 1927, p. 36, fig. 10: "Cranio
di Pignataro Interamna, vi.sto di fianco, ancora parzialmente immer.so nella dell 'El. antiquus di Pignataro Interamna, vLsto di fronteCjio della grand, nat.)."
sabbia Oio della grand, nat.)." Reproduced .same size. Compare Osborn, Reproduced .same size. Compare Osborn, 1931.846, p. o, fig. 4.
1931.846, p. o, fig. 5.

produced in our diagram (Fig. 1098). Had it not been for these portion of the cranium by attempting
to remove it for purpo.ses of
priceless measiu'ements, sketches, and figures, we should find exhibition. Thus theand unique cranium shown in Profes-
.superb
ourselves obliged to record one of the most tragic losses in the .sor De Lorenzo's description and in the figures and photographs

history of vertebrate palaeontology, namely, the characters of the above mentioned and reproduced in the present text no longer
cranium, jaws, and tusks of an adult Elephas antiquus in a perfect existed. The remaining parts of the specimen, namely, the rostrum,
condition of preservation. and the lower portion of the occiput were also
tusks, palate, jaws,
seriously damaged, while the entire upper portion was irrevocably
ACQUISITION BY THE AMERICAN MUSEUM IN THE YEAR 1929 to science with the exception of three small pieces extricated
lo.st

In the hope that the excavations would be continued and the with great difficulty from the hard cement in which Tiseo had
precious remains would become a part of the collection of the attempted to repair the terrible injury which he had inflicted on
State, Profe.ssor De Lorenzo, as Director of the Institute of Geology this priceless .specimen. At first the reconstruction of the skull
of the University of Naples, made every effort (1927, p. 39) to appeared hopeless, and the present author, who had donated it to
secure the specimen for the Naples Museum, but without success. the American Museum collection on the basis of the excellent
During the following year (September 4, 1928), the American photographs showing the specimen in its original condition, was
Museum of Natural History was informed of the desire of Saverio not even allowed to see it in the laboratory. After eighteen months
Tiseo to dispose of the specimen and began negotiations (No- of arduous labor on the part of Mr. Jeremiah Walsh, under the
vember, 1928) on condition that no step would be taken without direction of preparator Charles Lang, and of Curator Barnum
due permission from the authorities of the Italian Government. Brown, and Honorary Curator-in-Chief Osborn, the
finally of
On December 3, 1928, these terms were formulated in detail, and reconstruction entered its final stages in which the precise measure-
in May, 1929, the specimen was received in the American Museum. ments, figures, and photographs secured by Professor De Lorenzo
The total cost to the American Museum, including the donation of of the cranium in its original unfractured condition were of in-
1244 OSBORN: THE PROBOSCIDEA

antiquus germnnims), remove this great branch of the family

Elephantidae from its previous obscurity and uncertainty and
enable us to rank it as among the best known of the fossil elephants
of Eurasia.
Whereas Elephas [Hesperoloxodun] anlujuus belongs in the
Lower Pleistocene, or 1st Interglacial, the present specimen belongs
in the upper Middle Pleistocene,' or 3d rnterglacial it is somewhat ;

more progressive than the H. antiquus gennanicus of Weimar.

Hesperoloxodon antiquus italicus is far superior in size to
Palxoloxodon namadicus and greatly surpasses H. antiquus typicus
both in the number and height of the ridge-plates (see Figs. 1102
and 1088). It is a very progressive ascending mutation, equaling
in size, but exceeding in the number of its .superior ridge-plates,
the most progressive //. antiquus gennanicus of Weimar. How-
ever, from close comparison with all the numerous specimens de-
scribed from Lower Pleistocene deposits in England (by Falconer
and others) to 3d Interglacial deposits in Weimar (by Pohlig and
Soergel), H. antiquus italicus appears to be the largest and most
progressive member of the 'Elephas antiquus' phylum thus far
discovered.
/^ A/afura7 sz^e

Fig. 1101. Typp .superior grinders of Hesperoloxodon antiquus italicus

(Amer. Mus. 22634). One-fourth natural size. After original ijliotograph.*,
retoudied and numliered, displaying ridge-plate.s 1 to 20 in r.M^, ridge-platas
4 to 12 in r.M-. Compare Osborn, 1931.846, p. 10, fig. 8.
(rp[)er) Second and third right su|)erior molars, r.M-, r.M''.
(Lower) Palate exhibiting right and left superior molars, M-, M^.
znner "View Ml '"^
^J
HESPEROLOXODON ANTIQUUS ITALICUS

calculable value and importance. Thus, after almost continuous Amer. Mus. 22654 Type-
and very expensive labor between May, 1929, and November, Fig. 1102. —Type right superior and inferior grinders, M M 2, 3, of Hespe-
1930, the specimen was ready for complete description and exhibi- roloxodon antiquus italicus (Amer. Mus. 22634). Diagrammatic key to the

tion, altliough not open to the public until January 1, 1931. superior and inferior ridge-plates:

Despite; all these drawbacks and scientific disappointments,

M2\^' M3^.
The principal measurements of r.M'' are: Length of 20 ridge-plates 29.')
the restored cranium and tusks, jaws, and scapula still afford
mm.; height of 5th ridge-plate 190 mm.; 6 ridge-plates in 10 cm. The
a wealth of new knowledge regarding the relation.ships of the clas.sic principal measurements of r.Ms an;: Length 306 mm., height of 8th ridge-plate
'Elephas antiquus,' which, added to the equally priceless Upnor 128 nun., 4)^ ridge-plates in 10 cm.
skeleton, also the Steinheim crania (Elephas [Hesperoloxodon] After Osborn, 1931.846, p. 13, fig. 10.

Osborn in the present Memoir, as well as the general opinion today of geologists and paleon-
'[Careful consideration of the several statements by Profes.sor
tologists,has led to the conclusion that Profes.sor Osborn would have placed the 3rd Interglacial in the Upper Pleistocene; hence throughout it will be observed
that this determination has been adopted. —
Editor.]
 THE LOXODONTIN^: HESPEROLOXODON 1245

Fortunately the second and thirtl .superior and inferior molars

of both sides were preserved in situ and their characters are very
clearly displayed in the accomi^anying type figures, in which all

the ridge-plates are shown both in crown and lateral view, with

clear enumeration of the ridge-plate numbers in figures 1101, 1103,

and 1104, and in the diagrammatic figure 1102. Very important

is figure 1011 (lower) in which 15 ridge-plates are shown in simul-

taneous use, namely, r.M'-, ridge-plates 4 to 12}^, plus r.M', ridge-

plates Jo-l to 6. This stage of attrition represents a young adult
male, corresponding with the attrition of Elephas iiulicus estimated
to be about forty years of age. To the 20 ridge-plates actually
observed in r.M' (Fig. 1101) there may possibly be added ridge-
plates 21 and 22; whereas in a much older individual of the
typical ffesperolo.vorlnn antiqiius only 16%-n ridge-plates are
shown.

SUBSPECIFtC DESCRIPTION
Compare Osborn, 1931.846, pp. 17-24

Hesperoloxodon antiquus italicus Osborn, 1931

Figures 1068, 1069, 1088, 1092, 1095, 1096, 1098-1108, 1111, 1112, PI. xxni
Pignataro Interamna, near Cassino, Italy. Upper Pleistocene river
gravels of 3d Interglacial time.

Palseoloxodon antiquus italicus Osborn, 1931. "Palxoloxodon

antiquus italicus sp. nov., Final Stage in the 'Elephas antiquus'
Phylum." Amer. Mus. Novitates, No. 460, pp. 1-24. Type.—
Cranium and jaws with superior and inferior dentition (M 2-M 3)
of a young adult male; also right scapula (Amer. Mus.
22634). Locality and Horizon. "Pignataro Interamna, —
near Cassino, Italy. Found in upper Middle' Pleistocene river
gravels, of 3d Intei-glacial time, equivalent to, or slightly more
recent in geologic age than, the 3d Interglacial stage of Taubach-
Weimar of the Ilm River valley, Saxe-Weimar, Thuringia, northern
Germany, 40 kilometers east and a little south of Burgtonna, north
of Gotha, where the first .skeleton of 'Elephas antiquus' (cited by
Blumenbach as Elephas primigenius) was discovered in 1695."
Type Figure. — Op. cit., figs. 1, 4-16.

^ A/a.tu.TTa7 5ije

Fig. 1103. —Type mandible, right lateral and superior aspects, of Hespero-
loxodon antiquus italicus, one-sixth natural size.
A, Right lateral view, with second and third superior teeth, M", M', super-
posed on corresponding inferior teeth, M2, M3. After Osborn, 1931.846, p. 16,
fig. 12.

B, Superior view, with second inferior molar, Mj, exhibiting ridge-plates Fig. 1104. — Type of Hesperoloxodon antiquus Ualicus (Amer. Mus.
7-12, third inferior mclar, M3, exhibiting ridge-plates 1-8. After original 22634), Inner view of third right inferior molar, r.Ms, exhibiting +18 +
photograph. ridge-plates. One-fourth natural size. After Osborn, 1931.846, ]). 12, fig. 9.

'[See footnote on opposite page. — Editor.]

 PALAEOLOXODON ANTIC\UU5\; ITALICUS Type

Fig. 110.5. Hesperoloxodon antiquus italicds Type, of Pignatabo Interamna, Italy, Valley of the Libi, in the American Museum of Natural
History (Amer. Mus. 22634)
After photographs and indicated measurements (March 10, 1931) by the present autlior. Reduced to a uniform scale of one-sixteenth natural size. See
also type figure 1 100.

A, Left lateral view of cranium.

Al, Anterior view of cranium, exhibiting upturned portions of tusks in slight perspective.
A2, Posterior view of cranium and jaws, exhibiting exposed lower grinding teeth of the right side, tusks in strong receding perspective.
Observe the relatively high, narrow occiput exceeding in height that of Hesperoloxodon antiquus germanicus. The adult cranium (A) is about equal in
—
depth or bathycephaly (990e mm.) to that of //. ant. germanicus (980 mm. Fig. 1 14, B); this is owing to the greater depression of the tritural surface of M'
1

below the occipital condyle.

1246
 : ;

THE LOXODONTIN^: HESPEROLOXODON 1247

In the following description (Osborn, 1931.846, pp. 17-24)

references to the figures in the present Memoir have been substi-
tuted for the original figure numbens.
—
"SuBSPECiFic Characters. Displayed in the measurements,
ridge-plate formulae,and height of the ridge-plates, enumerated
below; also illustrated in the type figures 1101, 1102, and 1103.
Progressive evolution indicated as follows

Pal. n»t. typicus-}^^ M3 \^-. M', length 254 mm., height of

tallest ridge-plate 174 mm.; Ms,
length 315 mm., height of tallest
ridge-plate 126 mm.
Pal. ant. germanicus; M 3 Y^. M\ length 295 mm., height
of tallest ridge-plate 190 mm. ; Mo,
length 425 mm., height of tallest
ridge-plate 120 mm.
Pal. a7it. ilalicus (type M 3 ;x+. M', length 295 mm.,
height of tallest ridge-plate 190 mm.
M3, length 306 mm., height of tallest
ridge-plate 128 mm."

—
"Incisive Tusks. The incisive tusks of the type are not fully
grown [see Figs. 1100, 1099, 1106, 1107, 1108]. Total estimated length
3030-3070 mm., that is, 800e mm. within the alveolus plus 2230-
2270 mm. beyond the alveolar border. The longest tusk recorded
by Pohlig in the University of Rome collection measures 3900 mm.,
or 12 ft. 9/^ in., in comparison with 10 ft., length of the present
specimen. This indicates that a full-grown adult male of Pal. ant.
italicus attained gigantic size."
"Lower Jaws [Figs. 1103, 1106]. —The type inferior mandible
is entirely complete, as represented in figure [1103], requiring little
or no restoration. As compared with the more or less complete
mandibles figured by Falconer, from the Lower Pleistocene of
England, it closely resembles in profile aspect the typical 'Elephas HESPEROLOXODON ANTIQUTJS ITALICDS, TyPE
antiquiis' jaw but is very much larger and more massive; it differs
One twenty-fourth natural size. See figure 1 10.5, one-sixteent h natiiral size
widely in every aspect from the mandibles belonging to any
Fig. 1106. Type cranium of Palaeoloxodon anliquus ilalicus (Amer. Mus.
species o! Archidiskodon, Parelephas, or Mammonteus ; the rostrum 22634) as reconstructed and mounted in the American Museum during 1929
is abruptly truncated but less abbreviate than in 'Elephas primi- and 1930. One twenty-fourth natural size. Restored parts (oblique lines);
genius,' which is more of the extreme bathycephalic type." parts preserved (shading), namely, occipital condyles, portions of premaxil-

—
"Skull. The extremely broad rostrum, characteristic of all laries and
7, fig. 6.
raaxillaries, and complete mandible. After Osborn, 1931.846, p.

stages of the 'Elephas antiquiis' phylum, measuring 857 mm. or

The measurements, as seen from the front, with but two exceptions accord
2 ft. 9/^ in., is exactly the same width as that of the 'Elephas with those given by De Lorenzo. The few original fragments seen in frontal
platyrhynchus' of Graells, which measures 860e mm. or about 2 ft. aspect lie at the back of the narial chamber (c); along the border of the
10 in. transversely; this animal is from the Pleistocene, at San left temporal fossa ((), and along the temporal arch («'); in lateral aspect of tlie
left side, as shown in the fragments in the left temporal fossa (f, fi) just above
Isidro, near Madrid, Spain."
the fragment of the right temporal arch («'), transferred to the left side for
"Most novel and surprising is the dome-like, highly arched purposes of restoration. In posterior aspect, the occipital parts preserved are
occipito-parietofrontal contour [Figs. 1099, 1100, 1106, 1107, 1108] portion of the superior border (0'), and parts of the posterior occipital plate
which superficially resembles that of the Indian elephant more (0^, 0^) and the very broad condyles (0^).

'[On page 17 of American Museum Novitates, No. 460, Osborn presents the descriptions of three subspecies of Palseoloxodon anliquus, designated as P.
anliquus lypicus, P. anliquus germanicus, and P. anliquus ilalicus. According to current rules the name lypicus, designating the type of the species anliquus,
must be regarded as a still-born synonym of ajiliquus, since the name of the typical subspecies should be the same as the name of the siiecies. Consequently
the three subspecies would be:

Palxoloxodon anliquus anliquus (Falconer) Palseoloxodon anliquus germanicus (Stef£lnescu) Palseoloxodon anliquus ilalicus Osborn

On page 21 of this same publication, Osborn proposes the new genus Hesperoloxodon, with Palxoloxodon anliquus ilalicus as the generic type. But
a genus is based upon a species, so that the type of Hesperoloxodon should be anliquus and all of its subspecies. In other words, a species cannot be split
between two or more genera on the basis of its subspecies. (E. H. Colbert.)
Nevertheless the author's plain intention was to recognize the distinction of a "new loxodont phylum of the we.st" (as opposed to the Indian rmmadicus);
as a "type" of the "phylum" or "genus" (terms often used interchangeably by him) he designated Palseoloxodon anliquus ilalicus. Since this was his intention
the subspecies italicus should have been raised to the rank of a species to serve as a type of the genus Hesperoloxodon. (W. K. Greqoby.)]
1248 OSBORN: THE PROBOSCIDEA

closelythan that of either Loxodonta africana or Palxoloxodon 12 ft. 1/8 in. ; thus it appears from the cranial proportions only
namadicus. This lofty profile and corresponding liathyeephaly that the skeleton of Pal. ant. italicus was about fifteen per cent,
are represented correctly in figures [1106 and 1107 of the taller at the shoulder than that of the African elephant and
present Memoir], because they accord [Fig. 1098] with the closely similar in height to that of theUpnor elephant."
—
. . .

measurements and photographs taken by De Lorenzo and Saverio Cgmparison with P.\l^oloxodon namadicus. "Examined
Tiseo before this cranium was damaged. Moreover, beside the closely, theabove measurements prove that the cranium of Pal.
well preserved and extraordinarily broad occipital condyles (280 aril, profoundly different from that of ^E. namadicus'
italicus is

mm.), there are portions of the occiput ([Fig. 1106], o', o-, o^, o*) (the genotypic species I'l of Palseoloxodon) as well as from that of

Comparison with African Elephant

Loxod

(1) From front of orbit to back of occipital condyle

(2) From summit of parieto-occipital crest to attritional surface of
M' (bathycephaly)
(3) Occipital condyles, transverse
(4) Temporal arches, transverse width across
(5) Premaxillary rostrum, maximum width across
(6) Mandibular condyles height above angle of jaw
(7) Mandibular length, condyle to apex of rostrum
(8) Incisive tusks:
total length of
free length beyond rostnmi
maximum diameter at exit from rostrum
total circumference of

which prove that the occiput is forwardly inclined. The chief

comparative measurements between this restored cranium and the
cranium of the adult African bull elephant {Loxodonta africana
peeli — .see Fig. 1107) are as [above]."
May 5, 1929) gave a very high estimate of the
Pohlig (letter
length of the incisive tusks of 'Elephas aniiquus' of Germany,
namely, an average length of 5000 mm., or 16 ft. 4's in., as compared
with the incisive tusks of a Siberian 'Elephas primigenius' in the
St. Petersburg Museum (4300 mm., or 14 ft. 1!^' in.). He adds:
"The average incisor lengtii of a Siberian Mammoth at Peters-
burg, measured and figured by me (1891, p. 321, PI. B, fig. 3) is
4 m. 30, —so that the average length of the Elephas antiquus
must have been more than 5 meters."
inci.sor {op. cit., p. 51)

Comparison with Loxodonta africana peeli. "In — (4)

width across temporal arches, L. africana [peeli] exceeds Pal.

ant. italicus by six per cent. In percentages the cranium of Pal.
ant. italicus in all other measurements is from seven to twenty-
nine per cent, larger than that of L. africana, as follows:

(1) Orbit to occipital condyles 10%

(2) Cranial apex to grinding surface of M^ 24%
(3) Transverse occipital condyles 13%
(5) Width across premaxillary rostrum 29%
(7) Mandibular length 7%"
"On the average of the five principal measurements, the
cranium of Pal. ant. italicus is about fifteen per cent, larger than
that of L. africana. If this fifteen per cent, increase obtains
throughout the entire skeleton, the skeletal height of Pal. atil.
italicus should be about 3673 mm. or 12 ft., as compared with the
height iti the flesh of a large adult bull of L. africana, namely, 3450
mm. or 11 ft. 4 in., or with the skeletal height of the Palseoloxodon
antiqmis {andrewsif) [
= Hesperoloxodon antiquus] of Upnor, from
the top of the scapula to the ground, namely, 3700 mm. or
 THE LOXODONTIN.E: HESPEROLOXODON 1249

Loxodvnta africana, which is a relatively primitive cranium. the Indian elephant than of the African elephant. This bathy-
Coordinated with its very tall grinding teeth, the cranium of the cephaly however, a parallelism rather than a point of affinity,
is,

Italian specimen is much more bathycephalic (depth 990e mm. as because the very broad rostrum of Pal. ant. itah'cus presents an
compared with 800 mm. in L. africana) it is correspondingly less
; extreme difference from the very narrow rostrum of Elephas
brachycephalic (750e mm. as compared with 797 mm.); this is in indicus. Comparison with the cranium of 'Ekphas namadicus'
accord with the cranial proportions which are much nearer those of shows a strong resemblance in the breadth of the premaxillary
rostrimi but an extreme difference in the summit of the cranium,
which in 'E. namadicus' is relatively low and reinforced by the
overhanging parieto-frontal crest."
"This points to Pal. ant. italicus as a member of a phylum
quite distinct from that of the Siwalik 'E. namadicus,' a phylum
which supported by other cranial and skeletal differences might
if

well constitute a new genus to which the name Hesperoloxodon,

or 'loxodont of the west,' might be applied. This name is provision-
ally proposed, as I would not like to be forestalled a second time,
as in the case of Palseoloxodon, a generic name assigned to 'E.
namadicus naumanni' by Matsumoto but a few weeks prior to my
description of Sivalikia.
"Scapula. —The right scapula (1109A, Al) was found near
the type and may be considered as belonging to the same individual
(Amer. Mus. 22634) it greatly exceeds in size that of the African
;

and Indian elephants of the same age [Fig. 1109C, D, E]. On the

fyZB l^alliral lijf

L. AFRICANA
/4./V; 2/BB9 P. NAMADICUS
Tilgrim /905 P. ANT. ITALICUS
Pin A.M. 72634

Comparative Bathycephaly of the Loxodontin.e

Fig. 1108. Frontal and latoral views of three adult males, one-fortieth Comparison of Scapul.e
natural size. After Osborn, 1931.846, y. 20, fig. 14. One twenty-eighth natural size

A, Al, Lnindoiita africana. Batliyeephaly, 800: 710 mm. Male. Fig. 1109. Right scapula (A, Al in reversed outline) of the ty|)e of Pal.
[= Hesperoloxodon] anl. italicus (Amer. Mus. 22634) drawn to the same scale
B, I'alicoloxodon namadicus. Batliyeephaly, 728: .J92 mm. I'emale.
as the corresponding scapula of (B) Pal. ant. (anrfrcuisi'),^ of (C) Lox. africana,
Bl, Palxoloxodon namadicus. Male cranium of the Godiivari .\lhiviuni. and of (D) Ekphas indicus, juvenile (both after Andrews and Cooper, 1928,
After Pilgrim, 190.^. fig. 2), also of (E) Elephas indicus, adult (Amer. Mus. .544.")3), Vernay's middle-

aged male, of which the entire forelimb is shown in figure 1 194 of the i)resent
C, Cl. Pal;rohxodi)H [Hcsptruloxiidon] aidiijuus italicus type. Bathy-
Memoir.
cephaly, 990e.: 820emm.
Ohs(Tve that i'al. [IhsiJcroloxodnn] aidiqii.us ilalicus i.s nuicli more liathy-
mm.) than Loxodoida africana (800: 710 mm.), which is
cephalic (990e: 820e
oth(>r hand, it is slightly exceeded in size by the left scapula [Fig.
approximately the same as Pal. namadicus (728: 592 mm.). The female
1109B] preserved in the Upnor skeleton. The measurements of the
cranium (B) is much smaller than the large male cranium of the Godavari
Alluvium (Bl); compare the more accurate diagrammatic figure (Fig. 1110) of scapulae of Pal. ant. ilalicus and Pal. ant. (aiidreivsify are taken
the same cranium. with the restored border indicated in dotted lines:

Loj-. africana Pal. ant. italicus Pal. ant. {andrewsiiy

oxyotis(Jumbo) Pignataro Upnor
Scapula [Amer. Mus. 3283] Amer. Mus. 22634] [Brit. Mus.]
Height, superior border (restored) to center of glenoid border 925 mm. -H5% =1065 mm. +10% =1170 ram.
Width, median, across pre- and post-scapular borders 594 -1-29% = 770 +12% = 868e
Anteroposterior diameter of neck of scapula 240 +28% = 307 + 6% = 324
Anteroposterior diameter of glenoid border 187 +35% = 253 +15% = 290"
'[Equals Hesperoloxodon antiquus —see footnote on p. 1222 above. —Editor.]
 ; :

1250 OSBORN: THE PROBOSCIDEA

PALAEOLOXODON ANTIQUUS NAMADICUS

Pilgrim. /90S. Vo! XXXII. Pi's 10. II. /?

Large Male Cranium op Pal^oloxodon namadicus op the GodXvari Alluvium

One-twentieth natural size

FiR. 1110. Pateo^io(io7i namadicus, large male cranium discovered in the Godavari Alluvium, at Nandiir Madmoshwar, India (cf. Pilgrim, 1905, Pis.
10, 11, 12). Mea.surements after Pilgrim and Osborn.
P. namadicus H. ant. italicus

Fronto-vertical length 1270 mm. 1400 mm.

Transverse premaxillary rcstrum 920 857
Occiput, transverse width across 1075 875
This comparison shows that Palxohxodon Tuimadicus exceeds in breadth, while Hesperoloxodon anliguus italicus exceeds in dcptli.

"Estimates of Skeletal and Flesh Height. It is im- — "To this estimated skeletal height should be added about six

portant to compare the estimates of the shoulder height derivable and one-third per cent, to obtain the height in the flesh, giving us
from the scapula as well as from the cranium; they are found to an estimated height at the shoidder of 3905 mm. or 12 ft. 9?4 in."
agree exactly, as shown below. Neither the Upnor nor the Italian "Pal. ant. italicus of Pignataro
specimen is full-grown, yet combined they afford a priceless means Interamna, adult:
"
of estimating the height of the full-grown 'Elephas antiqtius.'
Estimated height in the flesh 3905 mm. = 12 ft. 9^4^ in.
"In each of the four above dimensions the scapula of Pal. ant.
italicus is from fifteen to thirty-five per cent, larger than that of Pal. ant. (andrewsify of Upnor, young
Lot. africana oxyotis ('Jumbo,' Amer. Mus. Dcpt. Mam. 3283) adult
the actual .skeletal height of "Jumbo" is 10 ft. 5% in. or 3194 mm.; Estimated height in the flesh 3934 mm. = 12 ft. 10?^ in.
consequently if we add fifteen per cent, (the difference in scapular
height) to the skeletal height of "Jumbo" we obtain 3673 mm. or Lot. africana, adult:
12 ft. % in. as the estimated skeletal height of the Pal. ant. italicus Height in the flesh 3450 mm. = 11 ft. 3% in."
type; this agrees with the height estimated from the proportions
of the cranium, namely, about 3673 mm. or 12 ft. Ya in." "The Pignataro Interamna specimen is several years older
"In height the scapula of Pal. ant. italicus is about ten per than the Upnor specimen, as indicated by the fact that the posteri-
cent, less than that of Pal. ant. (andrewsify of Upnor, the skeletal or ridge-plates of the second molar and the anterior ridge-plates of
height of which is 3700 mm. or 12 ft. \% in." the third molar [Fig. 1101] are in use, while in the Upnor specimen
Pal. ant. italicus: the ridge-plates of the .second molar only are in use. Comparison
Total skeletal height estimated with the growth of the large African elephant "Khartum" in the
from proportions of cranium 3673 mm. = 12 ft. % in. ca. New York Zoological Park shows that in captive conditions at the
Total skeletal height estimated age of twenty-.seven years the animal grows three-quarters of an
from proportions of scapula 3673 mm. = 12 ft. % in. ca. inch a year. By such an estimate the Pignataro Interamna adult is
Pal. ant. (andreicsifY: about five years older than the Upnor young adult; had it con-
Total skeletal height estimated tinued to increase in height, the fully adult bulls would measure
from proportions of ontin^ about 13 ft. 6 in. in height, or two feet above the shoulder lunght of
fore limb 3700 mm. = 12 ft. 1% in." a large fully adult African bull elephant."

'[Equals Hesperoloxodon anliguus — sec footnote on p. 1222 above. — Editor.]

 :

THE LOXODONTIN.E: HESPEROLOXODON 1251

FEMORAL LENGTHS AFTER POHLIG

The humerus bears a more constant ratio to the height of an
elephant tlian the femur, because the relative height of the fore
and hind quarters differs in each phylum. Yet the femoral lengths
where obtainable are very important as a key to pelvic heights.
Consequently the following notes as to femoral lengths recently
communicated by letter from Dr. Hans Pohlig, the leading his-
torian of the fossil proboscideans of western Europe (whose
portrait beside the femur of Rome appears in figure 1111 of the
present Memoir) are of interest.
Pohlig, 1929-1931.— (Letter, "Bonn, 5.III, 29"): "The . . .

anliquus femiu- (from Rome) is now at Lumbres, P. Calais (Dr.

Ponticr) [see Fig. 1111]. As my own height is 1 m. 70, the femur
will be about 1 m. 50, but it is not quite entire and not quite adult;

that of the large Taubach skeleton (the huge grinders of which are
figured in my Mon., plate vi, fig. 2a, lb, and the premaxillaries
(average breadth 1 meter! ibid. fig. 3), is 1 m. 60 at least. The . . .

largest [Archidiskodon] meridionalis femur at Florence is 1 m. 38.

The [Parelephas] trogontherii femur at Budapest 1 m. 43, according
to my Stcgodon Osteologic."
The length measurements of femora thus kindly recorded by
Doctor Pohlig for comparison with those recorded by others are
as follows

Hesperoloxodon anliquus italicus ref.,

femur from Rome now at Lumbres,

Pas de Calais (Dr. Pontier) partly ,

restored 1500e mm. 4 ft. 11 in.

Hesperoloxodon anliquus germanirus
ref., femur of large Taubach skele-
ton huge molars, Pohlig, 1889,
(cf.

PI. VI, figs. 2a, lb) 1600e mm. 5 ft. 3 in.

Archidiskodon meridionalis ref.,

Florence Museum 1380 mm. 4 ft. Q'a in.

Parelephas Irongontherii ref., femur,

Budapest 1430 mm. 4 ft. 8% in.

COMPARATIVE BRAIN CHARACTERS OF LOXODONTA AND

HESPEROLOXODON
Although the entire upper portion of the type cranium of
Hesperoloxodon anliquus italicus was practically destroyed, the
solid walls of the brain case fortunately were preserved, revealing
for the first time the brain characters of a member of the 'Elephas
anliquus' phylum. For immediate comparison intracranial casts
were taken from the relatively young crania of the African and
Indian elephants in the American Museum, affording the following
brain cube comparison, as displayed in figure 1112:

Brain and Volume

C, Elephas indicus (Amer. Mus.

54261), young adult male
B, Loxodonta africana (Amer. Mus.
51939), young adult male
Hesperoloxodon anliquus italicus
(Amer. Mus. 22634), adult male
not full grown
1252 OSBORN: THE PROBOSCIDEA

Max Weber (1896)

Elephas indicus, large male
Elephas indicus female (20 yrs. old)
Loxodonta africana, female
Amer. Mus. specimens
C, Elephas indicus, male, not fully adult
B, Loxodonta africana, male, about half grown
A, Hesperoloxodon ant. italicus, male, three-
fourths adult
 .

THE LOXODONTINiE: HESPEROLOXODON 1253

HESPEROLOXODON ANTIQUUS GERMANICUS OF 'Elephas antiquus,' of the urus and of Megaceros; the layer "e"
STEINHEIM containing the cranium and tusks of Elephas antiquus of Steinheim
Many years ago (Feb. 26, 1901) Dr. Eberhard Fraas ])ro.sented accordingly belongs toward the close of the 2d Interglacial period
to the American Museum from his Steinheim collection a finely [3d (Osborn)]; in the same layer was found (5) the remains of Leo
preserved superior molar, r.M' (Amer. Mus. 10655 — Fig. 1115) sp., of Ursus sp., and of Meles taxus (the badger).
which agrees approximately in its ridge formula (M 3-—) with
progressive molars (M 3 f|f ) from the Taubach-Weimar region of
Thuringia, described by Pohlig (1888-1891), as set forth in pages
1233-1235 of the present Memoir, and by Soergel (1912, Tab. i,
VII, VIIl).

During the years 1926 and 1928 the region of Steinheim on the
river Murr yielded to the Stuttgart Museum a number of invalu-
able remains of "Der Waldelefant" or 'forest elephant,' a pre-
liminary account of which has been given by Dr. Fritz Berckhemer,
Konservator, Naturaliensammlung, Stuttgart.
Wiirttemberg
Following his two articles of 1929 and 1930, Doctor Berckhemer
kindly furnished (April, 1931) excellent photographs (Fig. 1114),
with complete measurements, for inclusion in the present Memoir
in comparison with Hesperoloxodon antiquus italicus (Fig. 1105).
—
Geologic Age. Doctor Berckhemer describes (1929, p. 188)
the related geologic levels and fauna as of 2d Interglacial age.
The present author regards them rather as of 3d Interglacial age.

"[I. 1926] In Steinheim wurde zuerst die Kiesgrubc von

Sigrist aufgesucht, die 1926 einen ersten Schadel von Elephas
antiquus geliefert hat, und in der vor zwei Jahren ein prjich tiger
Riesenhirsch . . . zutage kam. Im SO-Tcil der Gi-ube sind die
Schotter zurzeit in einer Machtigkeit von rund 10 m freigelegt;
der anstehendc Untergrund soil nach Aussage des Besitzers noch Fig. 1113. W. Bauer Quarry, Steinheim a. d. Murr, showing the Site of
THE 1928 Discovery
3 m tiefer leigen. Uber dcm Schotter folgt hier eine Decke von
After original photograph kindly furnished by Dr. Berckhemer
rund 4 m Less und Losslehm. Der anh'^uus-Schadel [1926] lag in
(cf.

Berckhemer 1930, pp. 331, 332)

diesem Teil der Gnibe 8 m tief unter der Grenze Schotter/Loss-
lehm. Kurz vor der Tagung war noch ein Stirnschadel vom Wisent a. "Zu oberst haben wir noch Losslehm und Loss."
gemeldet worden, dessen Bergung in Gegenwart der Gesellschaft c. (c), die links im Bild
"Es folgen nach oben rotlichbraune Schotter
vorgenommen wurde. Das eindrucksvolle Stiick fand sich im auch an die Stelle der hier durch eine abtragende wasserstromung zuvor
entfernten Lettenbank treten (Abtragungszone durch weisse Linie bezeich-
tieferen Teil der iiber dem antiquus-hager folgenden Mammut-
net)." [Elephas primigenius fraasi.]
schotter. Auf der gegeniiberliegenden Seite der Strasse befindet
sich die neue Grube von Bauer, die durch den Fund des Buffelus d. "Daruber liegt eine feinsandige Lettenbank von graugriiner bis
dunkelbrauner Farbe."
murrensis Berckh. bekannt geworden ist."
. . .

c. [The layer consists of clear white sand of variable fineness, which

"[II. 1928] Sie lieferte im vergangenen Jahr einen zweiten toward the bottom is mingled with more or less gravel and sand.)
Schadel von Elephas antiquus, der soweit erhalten war, dass er X. Elephas antiquus [Hesperoloxodon antiquus gcrmanicus] cranium of
zur vollen Gestalt erganzt werden konnte. Er ermoglichte zu- 1928.
sammen mit einem im Juni d.J. im selben Lager gefundenen
antiquus-Stoaszahn (einen Rest der bei der Bergung entstandenen
"d" The fine-sanded loam bank is a still water formation in
Hohlung konnten die Toilnehmer in der Schotterwand noch be-
which are found remains of Bos, Megaceros, and the bear {Ursus
merken) erstmals eine Rekonstruktion des Elephas antiquus durch
arctos)
den Stuttgarter Oberpraparator Bock."
"c" The reddish brown gravels and sands mark a decided
In his article of 1930 (pp. 332, 333) Doctor Berckhemer adds change of climate, the Elephas primigenius fraasi replacing the
the fauna: 'forest elephant'; other remains, including the wisent {Bison

"e" In this layer were discovered remains of (1) Rhinoceros

priscus) and the wild horse {Equus sp.) which is quite frequent,
indicate a drier and colder climate of the Glacial Period, with
nierckii, (2) of the Murr water-buffalo {Buffelus murrensis),
restricted forests.
(3) a nearly perfect skull of the diluvial urus or aurochs {Bos
primigenius), (4) very numerous remains of the stag {Cervus Osborn: The Elephas antiquus horizon of Steinheim yields
elaphus) and of the royal stag {Megaceros). Obviously during this a fauna distinctive of the3d. Interglacial stage, consequently these

period of the deposition of the lower white sands "e", the surround- Steinheim remains may be provisionally referred to Hesperoloxodon
ing country was richly forested as the home of the forest elephant, antiquus germanicus.
M
 THE LOXODONTINiE: HESPEROLOXODON 1255

apparently enable us to establish the cranial characters of Hespero-

loxodon antiquus germanicus as quite distinct from those of H.
ant. italicKs above described.
In Cranium A of 1928 (Stutt. Mus. 15930) sixteen ridge-
])iates are exposed on the worn surface of the left superior molar,
l.M^; the right molar, r.M', may show
15-16 ridge-platos as com-
l)ared with the perfect superior molar (Fig. 1115) from Steinheim
exhibiting 19 ridge-plates as compared with 3 f~f in the M
Taubach-Weimar horizons described by Pohlig (1888-1891) and
Soergel (1912) mentioned above. The diameters of the crowns of
^^ ^ ^'*^-
A. Stutt. Amer. Mus.
Mus. 15930 10655
II. M' Length of 16 exposed ridge-
plates 245 mm.
Breadth 88
Length of 11 exposed ridge-plat('s 168 mm.
Maximum height of 11th ridge-
plate 195
L. M' Length of 16 exposed ridgc^-
plates (total ?+ 17) 245
Breadth 89
In the Stuttgart Museum l.M^ (No. 15344) exhibits a total of
19 ridge-plates; an r.M' (Stutt. Mus. 16515), length 300 mm.,
exhibits -f 18+ ridge-plates (total 20 ridge-plates).
In the right maxillary alveolus of A (Stutt. Mus. 15930) are
found the remains of a tusk (Berckhemer, 1930, Abb. 5) which
had been broken off during the lifetime of the animal, consequently
the right alveolus had been diminished by the formation of new
bony parts and the right premaxillary had correspondingly di-
Referred Molar of Hesperoloxodon ANXiqutis germanicds minished in breadth as compared with the normal width of the left
Fig. 1115. Third superior molar of the right side, r.M' (Amer. Mus. premaxillary (Fig. 1114, A2). The remaining left superior incisive
10655) found at Steinheim a. d. Murr and presented American Museum
to the
tusk was not preserved with the skull. [The isolated right tusk,
by Dr. Eberhard Fraas, February 26, 1901.
Observe 19 ridge-plates, 11 anterior worn; length 280 mm., height of 3500 mm. in length, does not belong to this skull; Doctor Berck-
nth ridge-plate 195 mm. Compare figure 1088B, Weimar molar, lengtii 295 hemer, however, does not hesitate to ascribe it to the same species.]
mm., height 190 mm.; also figure 1114 A2, Steinheim specimen. Total width of premaxillaries as seen from above 850 mm.
Width of diminished right premaxillary 400
STEINHEIM SKULLS OF 1928 (FIG. 1114A) AND OF 1926 (FIG. 1114B) Width of full-grown left premaxillary 450
Doctor Bcrckhemer's descriptioas refer to an exceptionally The cranial and dental characters of this remarkably well-
perfect cranium (Fig. 1114, A) of 1928 (Stutt. Mus. 15930) and to preserved individual are beautifully shown in the accompanying
a less perfect cranium (Fig. 1114, B) of 1926 (Stutt. Mus. 15344). photographs (Fig. 1114, Al-3). The principal measurements, in

These priceless new materials, with associated faunal remains, addition to those given above, are the following:

Hesperoloxodon antiquus germanicus Hesperoloxodon antiquus

A: Cranium of B: Cranium of italicus
1928 (Stutt. Mus. 1926 (Stutt. Mus. (Amer. Mus. 22634)
15930) 15344)
Summit of occiput to tip of right premaxillary 1375 mm. 1400 mm.
Transverse across widest portion of occiput 1095 875e
Mid-length of cranium, occipital condyles to tip of premaxil-
lary junction 1150 1130
Breadth across zygomatic arches 880 865 mm. 750e
Summit of parieto-occipital crest to attritional surface of r.M'
(bathycephaly) 860 980-1- 990e
Summit of parieto-occipital crest (partly restored) to attri-
tional surface of l.M' 885
Width of occipital condyles 295 307 280
Width of anterior nares 400 385e
Width across narrowest portion of maxillaries 545 480 530
Width across broadest portion of premaxillaries 850 890e 857
Occipital condyle to front border of orbit (left) 775 850
Occipital condyle to front border of orbit (right) 765 820e
1256 OSBORN: THE PROBOSCIDEA

The B Cranium of 1926 (Stutt. Mus. 15344), from the above

mciisuiTmcnts, appears to belong to a .somewhat older individual
More
(lAuQirQ/eitc4'^8
than the A Cranium of 1928 (Stutt. Mus. 15930). of tlie

dental ridge-plates in M' are exposed; the bathycephaly or depth

(from the summit of the occiput to triturating surface of M') is

980+ mm. compared with 860 mm. in the A Cranium of 1928;

a.s

the horizontal measurement from the occipital condyle to the front

border of the orbit in the B Cranium of 1926 is 850 mm. a.s com-
pared with 775 mm. in the A Cranium of 1928; the occipital
condyles as well as the orbit are more prominent and pediculated;
the premaxillaries are more elongated. In brief, the B Cranium of
1926 (Stutt. Mus. 15344) belongs to a fully mature and adult,
but not aged, liuU of H esperoloxodon antiquus germanicus.

HKSPEKOLOXODON ANTIQUUS CERMANICUS

FROM THURINGIA
Elephas antiquitatis Kruger, 1823 [ = Hesperoloxodon
—
ANTIQUUS germantcus]. (Kriiger, 1823, p. 832): "Eleph. anti-
quitatis. Europaischer Elephant. An einigen Orten in Europa
werden einzelne Spuren von eincr Elcphantenart angetroffen,
welche weniger dem asiatischen, luid mehr dem afrikanischen
t'lejihanten sich nahert. Sind auch die bei Thiede*) und bei
Teschen am Boberflu.ss im Jahre 1795 entdeckten Zahne**) achte
Mammuthsziihne mit abgenutzter Kaufliiche gewesen, und gehoren
sie nicht einer davon abweichenden Art an, so zeigen sich doch

bisweilen andere, welche den Zahnen des afrikanischen Elephanten

sehr ahnlich sind. Selbst Cuvier, der nur eine Art, die asiatischen
Mammuths, annimmt, muss doch zugestehen, dass sich bei
Eichstedt ein dem afrikani.schen Elephantenzahn ahnlicher
gefvmden habe***). So sind wahrscheinlich mehrere Ueberreste in
den Sammlungcn vorhanden, nur hat man sie bis jetzt iibersehen,
mid aus Vorliebe fiir den nordischen Elephanten auf Abweichungen
nicht geachtet."
Osborn, 1931 It is apparent from the above description, and
:

figure from Breislak reproduced herewith, that Kriiger had in

mind the 'ancient' or 'straight-tusked' elephant as distinct in tusk
structure from the modern and resembling rather
Asiatic elephant
the African elephant, since the first locality mentioned (Thiede)
is in the Thuringian region, northern Germany. The tooth which
we may select as Kriiger's tyi)e (after Breislak, Bd. II, s. 428) is not
improbably of the same 3d Interglarkil geologic age as Pohlig's and ^Lja^nf ro7i
Stefane.scu's 'Ekphas germanicus:' of the Taubach- Weimar horizon.
HesI'EROLOXODON ANTIQUUS OBRMAMCU.S ReF.
It is therefore for (ierman paijeontologists to determine whether
Fig. 111(5. Typo molar of Ekphas aiUiquilalin Kriigor, 1823, referred by
the specific name 'Elephas antiquitatis,' 1823, has priority over Hic present autlior to Hespcroloxodon atdiquus germanicus. After Breislak,
'Elephas germanicus' Pohlig — Stefaneseu. 1820, p. 428.

•Abgebildct in Breislak's Leiirb. d. Ci.jlogii'. 2t(r Bd. S 128.

"Arch. d. Urw. III., 2. S 396.
***J'ai vu line molaire donnfie eommc d'Aichstcdt, dans la collcetion d. M. Ebcl a Bremen; qiii)ii|iic (1 appari'iH'i' bic'M fdssilc, clle etoit remarquable par
sa resscmblance avec les molaires d'Afrique. Rcch. s. 1. O.ss. fo.s.s. Tli. I. S. 127.
 IV. EXTINCT DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS
(Continued from pp. 1182, 1183 of the present chapter, modified and extended see also observations of ;

Raymond Vaufrey, 1929, below, pp. 1268 to 1272.)

Of the greatest interest are the full-sized and dwarfed species of elephants found in the cavern deposits of the
different islands of the Mediterranean, successively described by Busk (1867), Falconer (1862, 1868), Leith Adams
(1870, 1874), Forsyth Major (1883), and Bate (1903 and 1907). To the latter author we are especially indebted for
the most recent discoveries and descriptions of these insular proboscidean species and subspecies. Falconer and
the older authorities related these dwarfed species to the 'Elephas antiquus' of the European continent, but subse-
quent discovery has shown that they are more probably derived from certain of the extinct ancestral African
species described below as Palxoloxodon (synonym Pilgrimia)}

Bathymetric Map of the Mediterranean Islands

Fig. 1117. Dwarfed Mediterranean species of elephants are found east to west on the islands of Cyprus, Crete, Malta, Sicily, and Sardinia. The white
area.s are within the 100 fathom line (600 feet); the depression of the sea or elevation of the land (in certain limited areas) was to the 216 fattiom line
maximum
(about 1300 feet) or 400 m. Reproduced by jiermission of Longmans, Green and Company, publishers, from map edited by Chisholm and Leete.
From a recent hydrographic chart (No. 4300 U. S. Navy, January, 1930), it is shown that the submerged land connections are now at the following depths:
(1) Cyprusto Turkish mainland (N.E. to Gulf of Alexandria near Adana, Turkey), shallowest depth 320 fathoms, 1920 feet; (2) Crete to Turkish main-
land (N.E. via island of Rhodes), shallowest depth 250 fathoms, 1500 feet; (3) Malta to Sicily, shallowest depth 74 fathoms, 444 feet; (4) Sicily to Italy
across strait of Messina, 109 fathoms, 657 feet; (5) Tunisia to Sicily, shallowest depth, 168 fathoms, 1008 feet; (6) Sardinia to Corsica, 46 fathoms,
276 feet; Corsica (northern end) to Italy, 140 fathoms, 840 feet.

Before discussing these questions of affinity, and the conclusions reached by Pohlig (1888, 1893), we may
follow the chronological order of systematic description.

1862 Malta, Zebbug Cave. Type molar tooth of Elephas (Loxodon) Melitensis Falconer, described in 1862.
1867 Malta, Zebbug Cave. Type skeleton of Elephas fale oneri Busk, 1867.
1870 Malta, rock-fissure, Mnaidra Gap. Type molar of Elephas mnaidrx Adams, 1870.
1883 Sardinia, sands of Morimentu. Elephas Lamarmorae Major, 1883. Carpal and tarsal bones.
1903 Cyprus, Kerynia Hills. Elephas Cypriotes Bate, 1903. Cotype molars.
1907 Crete, cave near Cape Maleka. Elephas creticus Bate, 1907. Nine imperfect molars, portion of an incisor, and
dorsal half of a vertebra.
1912 Sicily, Carini. Elephas antiquus var. insularis Soergel (name only).

Falconer was the first to describe before the British Association, October 6, 1862 (see "The Parthenon" for

October 18, 1862, p. 780), one of the pygmy elephants of Malta found in the Zebbug Cave, namely, Elephas

'[In a footnote on page 1 of Professor Osborn's Novitates article on the "Primitive Archidiskodon and Palaeoloxodon of South Africa" (Osborn, 1934.925)
appears the following statement: "Pilgrimia Osborn (December 20, 1924) is antedated by Pateo^oxodon Matsumoto (September 20, 1924)." Hence Pilgrimia
becomes a synonym of Palseoloxodon. — Editor.)
1257
 —

1258 OSBORN: THE PROBOSCIDEA

Melitensis, which is supplemented by more detailed description in the "Palaeontological Memoirs" of 1868 (Fal-

coner, 1868, Vol. II, pp. 292-308). According to Leith Adams (1870, p. 223), Elephasfalconeri, althougli found in
the same cave, represents a more diminutive animal than the Elephas melitensis of Falconer.

ELEPHAS MNAIDRIENSIS ELEPHAS FALCONERI ELEPHAS MELITENSIS

^iqCO-m-m. =qOO'mm =mOO-m-m'
Dwarfed Elephants [Pal^oloxodon (syn. Pilgrimia)] of the Mediterranean Island.s
One twenty-ninth natural size. Compare Figure 1119
Fig. 1118. Diagrammatic representation of the manner in which tlie three restorations of figure 1119 were calculated. As a rule the humerus affords the
most reliable estimate of the height of the shoulders. Comparison of the relative heights of these animals may best be made with the Upnor elephant (Hespero-
loxodon aniiquus).
Length of Humerus E.stimated Shoulder Height
//. aniiquus of Upnor 1290 mm. =4 ft. 2% in. 3700 mm. = 12 ft. \% in.

P. mnaidriensis of Malta 573 mm. = 1 ft. 10^ in. 1900 mm. = 6 ft. 2% in.

P. melitensis of Malta 478 mm. = 1 ft. 6% in. 1400 mm. = 4 ft. 7}^ in.
/'. falconeri of Malta 270 mm. = 10^ in. 900 mm. = 2 ft. 11% in.

Broadly .speaking, PaUeoloxodon mnaidriensis is one-half the height of Hesperoloxodon aniiquus of Upnor, P. melitensis is about two-fifths the height of H.
aniiquus, while /'. falconeri is only about one-fourth the height of the Upnor animal. As fully explained in the text, these dwarfed elephants apiH^ar to be
related to P. namadicus and to be descended from North African ancestral stages of Palseoloxodon.

The cranium (Fig. 1121)' and jaw (Fig. 1124) of Palseoloxodon melitensis, with broadly overhanging fronto-
parietal crest, more closely resemble the P. namadicus of India than the Hesperoloxodon antiquus of western
Europe.
Origin. — Comparison of the type grinding teeth of the dwarfed Mediterranean species with the type grinding
teeth of the extinct African species described below reveals a striking general resemblance in the narrow proportions
and in the rudiment or absence of the 'loxodont sinus,' characters which appear to relate these teeth to the phylum
Palseoloxodon of Africa rather than to the typical Loxodonta africana. In Europe some of the narrow-toothed
varieties of Hesperoloxodon antiquus may be related to the dwarfed insular elephants.- The characteristic body
height and progressive ridge formulae of these species were estimated by the authors as follows:
'(Referred by Professor Osborn to Palseoloxodon mnaidriensis. —Editor.)
-[See the opinion expressed by Professor Osborn on page 1252 above regarding the origin of th(! dwarfed elephants of the Mediterranean Islands.
Edtior.)
 THE LOXODONTIN^: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1259

Height Ridge Formula

Malta Elephas melitensis Falc, 1862, 1868 Height of Indian Tapir, i.e., 5 feet (1525 mm.) M 1

1
2
2

Elephas falconer/ Busk, 1867 2 ft. 6 in. (760 mm.) to 3 feet (915 mm.) M
Elephas mnaidrs' Adams, 1870 6 to 7 feet (1830 mm. to 2135 mm.) M 13K
1 3

Sardinia Elephas Lamarmorse Major, 1883 2 to 3 feet (610 mm. to 915 mm.) M
Cyprus Elephas Cypriotes Bate, 1903 ?2 to 3 feet (610 mm. to 915 mm.) M
Crete Elephas creticus Bate, 1907 SHghtly larger than E. Cypriotes, max. 5 feet (1525 mm.) M

r^f^ii-.^i«uV->,i')3

Dwarfed Eleph.\nt.s, Pal.eoloxodon (syn. Pilgrimia), of the Mediterranean Lsland.s

One twenty-fourth natural size. Compare Figure 1118

Fig. 1119. In desvendingsi'iile oS size, irom 'Elephas mnaidrierisis' {left), to 'E. melitensis' {right), to the diminutive 'E. falconeri' (center), the relative
heiglits are very carefully estimated from the respective lengtli of the limb bones (indicated in solid black in Fig. 1118). Drawn under the direction of the author
by Margret Flinsch, 1930.
Thelarger stage {E. mnaidriensis) is that of which the comphite cranium is known =the E. {antiguus) Melilse of Pohhg, 1893|, probably resembling tliat
[

of 'E.namadicus' the same form of cranium is attributed to E. melitensis and E. falconeri. The tusks are drawn from tlie recent comparative figures of
;

Vaufrey. This restoration is entirely diifcrent from tliat of Adams (Fig. 1127), wliich is based solely on the theory that these dwarfed elephants were related
to Loiodonta africana.

YouNQ African Pygmy' Elephant passing beneath Adult

Indian Female Elephant
Fig. 1120. The existing pygmy elephant Elephas afiicanus
pumilio Noack, 1906, from the French Congo, attaining a
height of 2 ft. 6 in. at the age of two and a half years and
corrcsijonding at this age in size with the most diminutive
fossil species P. lamarmora', and P.
PaUeoloxndon falconeri,
Cypriotes. Photograph by Mr. Elwin R. Sanborn
(1922)
through the courtesy of tiie New York Zoological Park. Tlie
intimate companionship and friendship of these two elephants
has been described (1928) in William T. Hornaday's volume
"Wild Animal Interviews."
At the age of four and a half years the same animal at-
tained a height of 4 ft. 02 in., corresponding more nearly with
PaUeoloxodirn melitensis and P. creticus. The full-grown Loxo-
donta africana pumilio (Amer. Mus. Dept. Mam. 3.'j')9l) attains
a height of 6 ft. 8 in., equaling that of the larger fossil Medi-
terranean species.

'[See editorial note on p. 1190 above.]

 1260 OSBORN: THE PROBOSCIDEA

In comparison, the dwarfed or small forest' elephant of the Congo, Loxodonta africana pumilio (Fig. 1120),
measured 2 ft. = 760 mm.) at the age of two and a half years; consequently at this age it agreed
6 in. in height (

with Palxoloxodon falconeri. The same animal at the age of four and a half years measured 4 ft. 4K inches in height
at the shoulders, and therefore attained the approximate height = 5 ft. or 1525 mm.) of P. melitensis or of P. (

creticus.

Ridge Formula. — In the following type and subsequent descriptions the ridge formulae of the species are
very carefully cited from the earlier and later publications by the authors who have successively treated the denti-
tion of these interesting animals, namely, Falconer, Leith Adams, Pohlig, Busk, and Bate. These type ridge
formulse constitute an important part of the type descriptions; they differ widely from the collective ridge formulse
cited from Raymond Vaufrey (1929) below; compare, for example, the ridge iormulaoVE.mnaidriensis' (M SJ^ttDj

Referred Pal«oloxodon mnaidriensis. Palermo Collection, Cranium I

One-t'ightli natural size.After Pohlig, 1893, who referred it to Elephas (anliquus) Melilar

Fig. 1121. Fully adult cranium of "Elephas (anliquus) Melita'" Fale., from the Interglacial layers
of the Grotta the most complete cranium in the collection of the Palermo Mu-
di Pontale, Carini, Sicily,
seum, exhibiting the characteristic prominent transverse frontal crest, comparable to that of 'Elephas
[
= PaUeoloxodon] namadicus' of India. After Pohlig, 1893, Taf. i, figs. 1, la, reduced to one-twelfth
n.atural .size.

For the i)r(jportion of this insular form to the full-grown Palaoloxodmi namadicus, see figure 1009, also the restoration (Fig. 1119). .Mthougli
these dwarf insular species of Malta usually have been referred to 'Elephas anliquus,' the present siwcimen is obviously more closely related
to P. -namadicus, and, while adult, belongs to an individual inferior in size to the juvenile P. namadicus (Fig. 1069), after Falconer and Cautley,
1847, PI. xxiv.A, fig. 4a.

Referred Pal.«oloxodon mnaidrien.sis. Palermo Collection, Chanidm VI

One-eighth natural size. After Pohlig, 1893, who referred it to Elephas (anliquus) Melitx

Fig. 1 122. Palate of juvenile cranium (Palermo Coll. VI), after Pohlig, 1893, Taf. ii, fig. 2, one-eighth natural size.

This juvenile cranium (Palermo Coll. VI), retaining the 8+ ridge-plated r.M", also the partly erupted r.M^ displaying only 9 plates, 3 jiartly
worn, is in a larger but younger stage than the adult cranium (Palermo Coll. I— Fig. 1 121 herewith). The measurenUMit from condyle to extremity
of maxillaries is approximately 744 mm., while in the adult cranium the same measurement is approximately 67.') mm. This ontogenetic short-
ening and deep(ming of the cranium as it grows older conforms with what we observe in the crania of all species of elephants.
(Pohlig, op. cit., p. 90) ".\ii Cranium VI, das, in basaler Aiisicht, auf Tafel II, in Fig. 2, abgebildet das Suborbitalforamen kaum
ist, ist m 02 lang,
obwohlder Schiidel, der grosstesicilische, mehralsOm 7 maximaler Liinge misst . . . . Die M. II. M. III. noch fast intact; die maximale
sind stark abradirt,
Breite der Schnauze, welche cxtrem divergente Lateralrilnder hat, betriigt ca. m 48. . . . die Condylen sind an dieseni Exemplar, ini Gcgensatze zu den
kleineren Cranien, nach Form, Stellung und relativer Grosso sohr iihnlieh wie bei dem typischen und dem namadischen E. anliquus, und folglich audi bci E.
africanus."

'[See editorial note on p. 1196 aliove.]

"Referred to 'Elephas mnaidriensis' by Leith .\dams and Raymond Vaufrey.
 :

THE LOXODONTIN^: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1261

given by Leith Adams, 1874, p. 112, with that (M

3 ?Kt) of Vaufrey (1929, pp. 124, 128). It appears that
Vaufrey (pp. 113-132, figs. 28-37) has intermingled the ridge formulae of the grinding teeth of more than one
species or mutation.

POHLIG ON THE MONOSPECIFIC 'ElEPHAS (aNTIQUUS) MeLIT^' OF SiCILY

Pohhg (1893, pp. 75-108) described and figured fully the referred 'Elephas {antiquus) MeLitx' Falc. from the
famous elephant grotto of Carini, Sicily, where were found in great numbers the skeletal parts of this dwarfed
species with numerous remains of the degenerate stag Cervus (elaphus) Sicilise and less abundant remains of both
Bos and Bison, Bos (taurus) primigenii and Bison prisons ; also traces of Hyxna spelxa and of partly worked flint

implements. Pohhg {op. cit., p. 100) maintains that the cranial structure of the six best preserved skulls from
Carini closely unites these insular pony-elephants with the European E. antiquus. The extraordinary transverse
frontal crest (Fig. 1121) observed in the adult Carini specimens Pohhg regards as the compelhng ground for the

specific union of the three forms "E. Namadi, E. Melitx und E. antiquus typus." Pohhg adheres (p. 101) to the
conclusions reached in his earlier (1891) monograph on E. antiquus of the specific identity of E. namadicus and
E. antiquus.

From these remains his geological inferences are as follows {op. cit., p. 82)

2. Landverbindung zwischen Sicilien und Italien einerseits, Afrika andererseits [error], und Einwanderung der grossen
Saugethiere, gegen das Ende der ersten diluvialen Glacialperiode.

3. Erneutes saculares Steigen des Meeresspiegels zu Beginn der diluvialen Interglacialzeit, erneute Isolation Siciliens, Aus-
bildung von diminutiven Formen grosser Siiugethierarten, Anhaufung von Skelettheilen soleher auch in der Grotta di Pontale,
einem Zufluchtsort namentlich flir Elephanten, Edelhirsche und Rinder.

Pohlig's palseontological observations are {op. cit., p. 83):

In meiner angefiihrten Monographie des Elephas antiquus habe ich zuerst naciigewiesen, dass die Angaben de Anca's und
Gemmellaro's von Elephas armeniacus und E. africanus aus Sicilien, ebenso die Artbezeichnungen E. Falconer i von Busk
[Footnote: 'Transactions of the zoolog. soc. London 1868, VI., p. 5, No. 10.'] und E. mnaidriensis von L. Adams [Footnote:
'Ihid. vol. IX, p. 1. 1877.'] aus Malta auf Irrthum beruhen, und dass der 'Elephas melilensis' von Falconer nichts anderes ist,
als eine insulare Diminutivrasse des Urelephanten, Elephas antiquus, fiir welche ich daher die Bezeichnung E. {antiquus)
Melitae Falc. vorschlug. Zugleich erbrachte ich die ersten Nachweise der Thatsache, dass die gleiche Zwergelephantenrasse,
wenn auch nicht bis zu gleich extremer Grossenreduction, wie auf dem kleinen Malta, auch auf Sicilien und in anderen Mit-
telmeergegenden gelebt hat.

From this Carini grotto Pohlig describes and figures the most completely preserved crania in the Palermo
Museum {op. cit., pp. 84-98) concluding as follows (p. 98):

Die Vereinigung der drei Speciesnamen Elephas melitensis Falc., E. Falconeri Busk und E. mnaidriensis L. Adams, die
lediglich aufden Dimensions verhiiltnissen des Malteser Materiales beruhten, unter der Rassenbezeichnung E. antiquus Melitx
Falc. wurde in meiner Elephantenmonographie vorzugsweise begriindet auf die wichtigste bis dahin bekannte Eigenthiimlich-
keit jener Diminutivformen, deren Dentition, —
wobei der iiberraschende Nachweis in der Gestaltung der fruhesten, in gleicher
Weise sonst bisher von keiner Species bekannten Milchdentition der Malteser Elephanten, auch fiir den typischen Taubacher
E. antiquus, au.sschlaggebend sein musste.

According to my observations, Pohlig continues {op. cit., p. 99), neither in North Africa nor in lower Italy
(in the west) does any fossU species occur excepting 'E. antiquus,' while in the Mediterranean islands not a single
trace has been found of a specimen of the normal continental size. In Sicily occur only the larger types of dwarfed
E. antiquus from one-half to two-thirds the size of the normal continental forms, corresponding in dimensions
1262 OSBORN: THE PROBOSCIDEA

with the larger E. mnaidriensis Leith Adams of Malta. This larger subspecies obviously lived in Malta at a time
when it was united with Sicily by a land bridge. There followed more severe living conditions on the smaller
rocky islands which brought about further degeneration and reduction in size. According to the dimensions of the
numerous Umb bones from the Grotta di Pontale of Carini, in comparison with those of the typical E. antiquus
the Sicilian Elephas {antiquus) Melitse in its full growth or adult condition was about the size of a middle-aged
Indian elephant; whereas on the continent the intermediate stages between the full-sized E. antiquus and the
smaller forms of the known E. antiquus remains from the Islands are without exception diminutive and all the
specimens from the very rich material of Carini belong to the diminutive race. On the other hand, the single
Sicilian molar found elsewhere corresponds with a small example of the true E. antiquus.

Osborn (1928) entirely dissents from Pohhg's opinion as to the specific union of Elephas antiquus Falc, 1847,
with the earlier described species E. namadicus Falc, 1846. It would appear that Pilgrim (1905) was influenced by
Pohlig in the same erroneous confusion of these two very distinct species of western Europe and the Siwahks.

Osborn, 1930: The cranium of E. [Palaeoloxodon] mnaidriensis (Fig. 1121) closely resembles that of E. [Palseo-
loxodon] namadicus of the Nerbudda, India, in the transverse crest; it differs widely from the cranium of
Palseoloxodon [Hesperoloxodon] antiquus italicus (Fig. 1106).

SYSTEMATIC DESCRIPTION OF MEDITERRANEAN ISLAND SPECIES

Compare the specific revision by Vaufrey (1929)

Palseoloxodon melitensis Falconer, 1862 292, 299, .307, 308. Typk.— Third upper molar of the left
Figures 1041, 1008, 1009, lllS, 1119, 1123, 1124, 1127, 1131, 1133, PI. xxiii side, Mus. 44312).
l.M' (Brit. Hokizon and Locality. —
Pleistocene, Zebbiig Cave, island of Malta. From the lower cave de- Pleistocene, Zebbug Cave, island of Malta. Type Figure. —
posits or levels. Op. cil., 1868, Vol. II, PI. XI, figs. 1, la.

Elephns melitensis Falconer, 1862, "The Parthenon," October, —

Original Description. This elephant was found in the
1862, p. 780; al.so "Palajontological Memoirs," 1868, Vol. II, pi). Zebbug Cave, and is the first described of the "pigmy" elephants of
Malta. Falconer ("Parthenon," Oct. 18, 1862, p. 780— see reprint
in "PalaeontologicalMemoirs," Vol. II, p. 308) states: "The
pigmy Elephant was an animal of remarkably small proportions;
an adult individual could not have exceeded the Indian Tapir in
height and bulk, a creature not much larger than a full-grown Hog.
Contrasted with the bones and teeth of an adult African Elephant
the difference in size of these portions of its frame exhibited were
most striking. But though .so small, the skeleton agreed in every
. . .

particular with the one of greatest bulk. A series of harmonies ran

through the two skeletons, one bone answering to another truly,
and without ordinal or generic difference. The author could refer it
unhesitatingly to his subgenus Loxodon, in the African group of
Elephants."
Supplementary Description. — Falconer described the teeth
of this species at the British A.ssociation, October 6, 1862 (an ab-
stract of which appeared in "The Parthenon," as stated above)
Type of Pal.holoxgdon MBLiTt;Nsis and in 1868, Vol. II, pp. 292, 299, he designated the type as fol-

Fig. 1123. Elephas Mrlitensis Falconer, 1802, 18«i8, type molar tootli lows: "One most characteristic of these [fossils] is an upper
of the
(Brit. Mas. 44312), one-half natural size. After Falconer, 1808, Vol. II, PI. molar of the left .side. ... As this specimen is about to be returned
.XI, la: "Views in plan and profile of last upper true molar, left side.
figs. 1,
to Malta, at Captain Spratt's request, it is necessary to make an
Described at page 292." l.ydekker (1880, p. l.')4) catalogues this tooth (Brit.
accurate description of it, to accompany the figures drawn by Mr.
Mus. 44312) as follows: "The third left upper true molar; from Zebbug Cave.
Dinkel. (PI. xi. figs. 1 and la.)" He al.so states on pages 283, 284,
Described and figured in 'Falconer's Palffontological Memoirs,' vol. ii. p. 292,
pi. xi. figs. 1, In; and noticed by Busk, o/). ril. 11807.11, p. 290, atid by Lcith- that Captain Sprat had "lately discovered in Malta numerous re-
t

Adams, op. cit. [1874.1], x. p. 28. Leith-AdaimCiMeclvm. P urchaned, IH73." mains of a surprisingly small fossil Elephant, of the sub-genus
 :

THE LOXODONTIN^: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1263

Loxodon, which I have named E. Melitensis," and on the "Descrip- RESUME BY DEPERET AND MAYET, 1923, PAGE 175

tion of Plate xi" he refers to this species as "Elephas (Loxodon)

" Nous resumons dans le tableau ci-dessous les donndes numeriques
Melitensis
I"' caractdriseat les especes ou mutations du rameau de 1'^. meli-
Falconer (op. cit., 1868, p. 298) gives the ridge formula of
tensis :
E. melitensis as follows
Dimensions des Mm ea millimetres

[Dp 2] i [Dp 3] f [Dp 4] f [M 1] I [M 2] f [M 3] if.

Bate (cf. 1904, p. 357) observes that the ridge formula of E.

'^.y/"''<"e*
Cypriotes is than that which Leith Adams, later than
slightly lower •

Falconer and after examining a further large amount of material,

gives for E. melitensis:

Dp 2 I Dp 3 f Dp 4 1^ M 1 1-^ M 2 {^ M 3 f|.

R.M2

L.M2

iy4 Natural sue

S. (ANTiqUA) MELITAE /2ilc. Ref.

Referred Mandible of Pal^eoloxodon melitensis, from the Grotta di

PoNTALE, Sicily
One-fourth natural size
Fig. 1 124. Juvenile jaw containing right and left second inferior molars,
r.Ma, I.M2 (Amer. Mus. 2011), acquired by the Museum in exchange with the
Padua University, Italy. Observe the sharply truncated vertical rostrum and
the broad forwardly pitched coronoid process. The grinders exhibit nine
complete elevated ridge-plates and two posterior half ridge-plates, or a total of
eleven ridge-plates, namely, M 2 '^444^.
 :

1264 OSBORN: THE PROBOSCIDEA

Of this animal, Adams (1870, p. 223) remarks: "The ..'/'»rz^ S£<c/y,/o^. "/M a^ K'/'

Lesser Pigmy Elephant. Elephas faUoneri, Busk. As be-

fore remarked, the presence of this species in the exuviae
of the Zebbug cave was determined by Mr. Busk after Dr.
Falconer's death, and seemingly more or less from fragments
of bones, which on comparison with similar portions of the
skeleton of Elephas melilensis and a footbone (astragalus)
discovered by me
Mnaidra gap, there appeared sucli
in
and configuration as to warrant the
differences both in size
assumption that they represented a still more diminutive
proboscidian, the average height of which at the withers
could not have exceeded two feet six to three feet."
Osborn, 1930: As Busk's type (1867) is a diminutive
skeleton without teeth, it remained for Bate (1903, 1904) to
give the ridge formula of 'Elephas Cypriotes' cited below,
without reference to the ridge formula of 'E. falconeri.' So
far as we can find, Vaufrey was the first to suggest that
'E. Cypriotes' is synonymous with 'E. faUoneri' and to giv(>

(1929, pp. 98, 99) the high collective ridge formula of E.

falconeri —E. Cypriotes as follows
M2^_,:^M3H=^.
This is a higher ridge-plate formula than that (M 3 ~i)
finally assigned to E. Cypriotes by Bate, it equals that

(M 3 jf 1^) of E. mtiaidriensis, fide Adams. Tins discrejmncy

partly depends upon the reckoning of the half crests or talons
omitted above.

•'D'Mt.lt, 11M|<4>

Type and Pakatype Molars op Pal.«oloxodon MNAn>KiENSis

Reproduced herewith one-half natiu-al size

Fig. 1126. Elephas nmaidriensis Adams, from Mnaidra Gap, Malta, .\ftcr Leilh
Adams, 1874, Pl. vii, figs. 1, 2 and 2a. Original figures (n.atural size) reproduced
iKTowith one-half natural size.
(Up|)er figure, 1) Side view of a 14 ridge-plated third superior molar Iparatype]
(if the right sidi>, r.M', described by Leith Adams (np. cil,, p. 33) as follows:
"2. Two beautiful and highly suggestive exami)les of what must be considered
last true molars, are represented by the entire specimens Nos. 64 & !\9 (PI. vii. figs.

1 & 2 & 2a). The former, an up|>er tooth, shows foiu-teen ridges, including the l)ygmy
digitated posterior talon a, in a space of 7 inches [178 mm.|. Attached in front,
although not shown in the figure, are two plates of the iM'nultimate molar. As the
crown is just being invaded, of course its pattern is not develojx'd; the macluerides
are therefore well crimiK'd, and the plates and enamel thick." Brit. Mus. 44306.
(Lower figures, 2, 2a) Crown and internal views of a right third inferior molar,
Type of Pal.imloxodon mnaidriensis
r.Ms. This is the type molar; th<' same tooth as that shown in Adams' original
Fig. 1125. Type of Elephas mnaidrx Adam.s, 1870, PI. ii, figs. 2,
figure (Fig. 112.') of the present Memoir). This b<'autif\Uly i)rc.served molar is de-
2a [ = Elephas nmaidriensis Adam.s, 1874, ]). 116], one-half natural .sine.
scribed by Adams {op. cil., 1874, p. 33) as follows:
Adams (1874, p. .33) dcsfribcs the .sani(^ tcK)tli and gives a much The next, No. .59 (figs. 2 & 2a), is a much arcuated lower molar; the last ridge,
more accurate figure (Fig. 1 12))) of the type (PI. vii, figs. 2, 2a). Lydek- although rounded and finger-like, rises like the others from the common base to the
ker (1886, p. 142) designated this molar (Brit. Mus. 44304) as follows: same level as the penultimate. There is a slight flattening on its base internally,
"The third right lower true molar, about one-third worn; from Mnaiilra but no trace of what could be called a jiressure-mark. The crown is broad in front,
gap. Described and figured, op. cil. p. 33, pi. vii, figs. 2, 2a. The tai)ering .steadily po.steriorly. The
anterior talon is large and .semilunar; and the
crown is remarkable for its excessive lateral curvature." See al.so anterior fang .seems to support and the succeeding plate only. Here we have four-
it

figure 1 126. teen ridges in 6..'> inches 166 nun.|." Brit. Mus. 44304.
1
 :

THE LOXODONTINiE: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1265

Palseoloxodon mnaidriensis Adams, 1870

Figures 1068, 1118, 1119, 1121, 1122, 112.5-1127, 1131, 1132, PI. xxiii

Pleistocene, Mnaidra Gap, island of Malta.

This is tho third species to be described from tlic island of

Malta. Adams (1870, p. 223) calls it the "Large Elephant of
Malta," estimating it.s stature as six to seven feet, as compared
with the Elephas melitensis of intermediate stature, and the Ekphas
fakoneri of extremely dwarfed stature, namely, two to three feet.
Ekphas vinaidrx Adams, 1870. "Notes of a Naturalist in the
Nile Valley and Malta," etc., 1870, p. 223. Type.—A last
lower molar of the right side, r.Mj (Brit. Mus. 44304). Para-
type.— A third superior molar of the right r.M' (Brit. Mus. side,

44306). Horizon and Locality. — Pleistocene, Mnaidra

Gap, island of Malta. Type Figure. — Op. PL cit., ii, figs.

2,2a; also 1874, PI. vii, figs. 2, 2a. Paratype, op. crt., 1874, PI. vii,
fig. 1.

Type Description. —Adams (1870, p. 223) distinguishes this

as "The Large Elephant Malta," estimating its average stature
of
(op. cit., p. 228) as six feet ten inches to seven feet as compared
with the average height of E. falconeri of two feet six inches to three
feet (op. cit., p. 223). As E. falconeri is distinguished by its

extremely dwarfed stature, E. mnaidrse is distinguished by its

relatively large stature, and E. melitensis by its intermediate

stature. Adams (op. cit., p. 228) concludes: "Thus, to all present
appearances, wo have represented by the remains hitherto col-
lected in the Maltese islands no less than three distinct species of
elephants of about the proportions indicated, and represented in
the spirited and well-executed drawing, page 161 [Fig. 1127 of the
present Memoir], for which I am indebted to the able pencil of
Mrs. Blackburn, whose admirable representations of animals have
obtained for her a high position as a delineator of natural objects."
Subsequently Adams (1874, p. 116) added: "I believe that the
bones of the Maltese fossil elephants are divisible into three
varieties and two well-marked species, viz. a large and a small
Elephant, the latter showing two forms represented by the
Elephas melitensis of Falconer and Busk, which may have seldom
attained a height of 5 feet, and a diminutive or pygmy form named
by Mr. Busk Elephas falconeri, the .smallest bones of which indicate
an elephant about 3 feet in height. But there arc intermediate-
Denizens of Ancif nt Malta.— 1. Large fossil Elephant (Elephas mnaulne). p. 223. 2. Pigmy

sized bones which easily bridge over the differences between the Maltese Elephant (Elephas wMtr-nsis), p. 216. 3. Smallest Dwarf Elephant (Elephas Jalconeri), p. 223.
4. Fossil Hippopotamus (ff. perUlnndi), p. 212. 6. Great Dormouse (Mycmts TittJiUnsis), p. 234.
latter and the Elephas melitensis; nevertheless Mr. Busk has 5. Great extinct Swan (Cyi^TMis/flZconen), p. 237. 7. Large extinct fresh-water Turtle, p. 237.

pointed out characters appertaining to the two, and is of opinion

that they are distinct species. ... I have named the largest Ele- Adams' Restoration a.s Dwarfed African Elephants
phant Elephas mnaidriensis, in consideration of the circumstance
Fig. 1127. Reproduced directly from Adams' figure in his book "Notes of
that the gap, or rock-rent, from which I obtained the most perfect a Naturalist in the Nile Valley and Malta," 1870, opposite page 161. In this
its bony structure is situated close to the ruins of the
specimens of very picturesque drawing we observe that Adams has restored the external
Mnaidra temple, a prehistoric and megalithic structure bearing cars of these three species in direct imitation of the very large ears of Loxodonta
africana, whereas it is probable that these dwarfed elephants had the smaller
evidences of the earliest human occupation of the Island of
ears which we have shown in figures 1047 and 1074 to be doubtless characteristic
Malta." Leith Adams in this .second contribution (1874, p. 116)
of HesperoloxodoH antiquum and all its descendants. It should be remembered,
changed the original name mnaidrx to mnaidriensis, the latter liowcver, that the external oars of Palaioloxodon immadicus, to wliich the type
Ix'ing the form cited by Lydekker (1886, p. 138), and gave an of P. mcliteiisis is more nearly related, arc still unknown.
excellent new figure of the type molar (PI. vii, figs. 2, 2a), also the
Compare new restoration (Fig. under the direction of present
the 1 1 19), tlie
following ridge formula (p. 112 and PI. vii) author, based on the obvious affinity of these dwarfed elephants to Palseoloxo-
don namadicus of India rather than to Hesperoloxodon anliquus of western
Dp 2 f Dp 3 f Dp 4 M 1 1:1 M2 ;^-M3 \i-'-
1 3-I1-11'
Europe, also with small ears.
 :

1266 OSBORN: THE PROBOSCIDEA

Palseoloxodon lamarmorae Forsyth Major, 1883 very apt to be over-estimated owing to the respective dwarf
Sardinia, Quaternary sands of Morimontu near Gonncsa. proportions of these island races. The Maltese jiigmy species have
been considered to be most closely allied to E. antiquus and E.
Elephas Lamarmorae Major, 1883. "Dio Tyrrlicnis," etc.,
africanus. ... On the other hand it seems more probable that E.
Kosmos, 1883, VII Jahig., Bd. XIII, p. 6. Type.— The
eypriotes, which so far as available material is (concerned shows no
author in describing this species designates a.s the types "die Ex-
close affinity to the African species, is rather connected with E.
tremitaten-Knorlien eines kloineii, aber vollstiiiidig ausgcwach-
senen Elephaiiteii ziim Vorsehein." Horizon and Locality. — antiquus and E. meridionalis, agreeing with this last in the lowness
of its ridge formula, though differing in wanting the persistence
. . .

(Op. cit., p. 6) : "Alls quatern. Sanden von Morimentu b. Gonnesa

of the strongly marked digitation of the plates which is usually
(Sardinieii)." Type Figi're. No figures published. — found in the molars of that elephant.
—
Type Description. The animal is said to agree in size with
E. mnaidrse [E. mnaidriensis] from Malta, but differs pretty shar])ly
in the carpal and tarsal bones which constitute the type. The
author's full description (Major, 1883, p. 7) is as follows: "Im
vergangenen Jahre kamen in dem von Lamarmora beschriebenen
quaternaren 'Gres' von Morimentu bei Gonnesa (Sardinien) die
Extremitaten-Knochen eines kleinen, aber voUstandig ausge-
wachsenen Elephanten zum Vorschein. Die von Malta bekannten
Zwerg-elephanten, mit deren grosstem, E. mnaidriensis, der sar-
dische in den Dimensionen ziemlich gut tibereinstimmt, luden in
erster Linie zur Vergleichung ein ; es stellte sich aber heraus, dass
die Carpal-und Tarsalknochen ziemhch bcdoutend abweichen."
—
Summary. Agrees in size with E. 7nnaidriensis but differs
pretty sharply in the carpal and tarsal bones which constitute
the type.

Palseoloxodon Cypriotes Bate, 1903

Figure 1128

PlfLstoccne, Kerynia Hills, island of Cyprus.

Elephas Cypriotes Bate, 1903. "Preliminary Note on the Dis-

covery of a Pigmy Elephant in the Pleistocene of Cyprus." Proc.
Roy. Soc, London, 1903, Vol. 71, No. 475, pp. 498-500; also Cotypes op Pal^oloxodon Cypriotes
"Further Note on the Remains of Elephas Cypriotes from a Cave- Fig. 1128. Elephas Cypriotes Bate, 1903, Kerynia Hills, island of Cyprus.
Deposit in Cyprus." Phil. Trans. Roy. Soc. London, 1904, Vol. Selection from the many cotype molars fully figured by Bate (1904, Pis.

197 (B), pp. 347-360. Cotypes. The following have been — XXI, xxii).
"Crown view of .second true molar in right man-
(Upijer) PI. XXI, fig. 3,
selected by the present author from the many cotypes: a second
dibular ramus. B.M., M.8588, p. 35.5." Reduced to one-half natural size.
true molar of the right side (Brit. Mus. M.8588), also a last lower "Crown view of last lower true molar. B.M.,
true molar (Brit. Mus. M,8591). Horizon and Locality. — (Ijower) PI. XXII,
M.8591, p. 355"; fig.
fig.

6a, "Side
6,
view of ditto." Both reduced to one-half
Pleistocene, a single cave deposit situated on the southern side of natural size.

the Kerynia Range, island of Cyprus. Cotype Figures. —

Op. cit., 1904, Pis. XXI, XXII.
from the char-
Bate (1903, p. 500). — In the original description the author
In this connection
acters of the carpal
it is

bones (the only remains known) of

and
interesting to note that
tarsal
observes: "Taking into consideration the several characters in
E. lamarmorae, the dwarf elephant of Sardinia, Dr. C. I. Forsyth
which the teeth of the Cyprus elephant differ from those of all the
Major considers it to have been most closely allied to E. meri-
hitherto described dwarf species (putting on one side E. lamarmorae
dionalis. Since writing the first notice of E. Cypriotes I have
. . .

. .from the Pleistocene of Sardinia, the teeth of which are un-

.

come to the conclusion that the average number of plates in the

known to science) as well as the distinct habitat of the animal, I

from true molars is somewhat less than was at first supposed, therefore
have come to the conclusion that it is specifically distinct
the corrected ridge-formula, exclusive of talons, would stand as
these other small forms, though possibly they were derived from
follows
a common ancestor, and I, therefore, propose to name it Elephas
Cypriotes."
[E. Cypriotes]: [Dp 2] -=-, [Dp 3] %, [Dp 4] Y.l, [M 1] ^t
Bate (1904, p. 357): In a supplementary description the
'-^
author states: "Undoubtedly there is a .strong resemblance be- [M 2] 01 I'^l 'j] 11 - 12-
tween the teeth of E. cypi-iotes and those of the Maltese and
Sicihan forms, more especially E. inelilensis, but this likeness is This is slightly lower than that of E. mclitensis which Dr.
 THE LOXODONTIN^: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1267

Lcitli Adams, later than Dr. Fak-onor and after examining a further Bate (1907, p. 239).— The author described the type as fol-
large amount of material, give.s as: lows: "The remains of the smallest of the Cretan Elephants were
all obtained from a much damaged and weathered cave-de])osit in
[E. melitensis]: [Dp 2] i, [Dp 3] i [Dp 4] [Ml]
the limestone cliffs near Cape Maleka in the west of the island,
[M 2) li, [M 3] li which has already been described [Footnote: 'Geol. Mag. n.s. dec.
v. vol. ii. (1905) ]). 195.'], and where only some te(>th and limb-
The marked compression of the tusks of E. rypriotes
lateral
bones of small rodents were found besides those under discussion.
(Plate 22, fig. 9), which is a constant character in all the sjjeci-
These latter include nine imperfect molars and a few fragments,
mens so far obtained, would in itself be almost sufficient to distin-
among which are a ])ortion of an inci.sor and the dorsal half of
guish this from the other pigmy elephants of the Mediterranean
a vertebra. As this small race differsfrom those of other Mediter-
region. . . . Presui>posing that the dwarf elojihants of the Mediter-
ranean islands, and its minute proportions being seemingly the
ranean region were all derived from a common ancestor, the simpler
result of specialisation (hie to isolation in Crete, it is suggested
construction of the molars of E. Cypriotes might be explained on the
that it may be known by the above specific name denoting its
assumption that this species was isolated and subsequently differ-
island habitat."
entiated from the parent stock, prior to a similar fate overtaking
Of the nine molars two lielong to the upper series, namely, M'
the Maltese and Sicilian races. This is l)orne out by the fact that
consisting of seven or .seven and a half plates, length 54 mm.,
E. Cypriotes lived contemporaneously with //. [Hippopotamus]
height about 32 mm. A referred M' includes eight ])lus plates. Of
minutus, a more generalized and primitive form . . . than eitlior
the cotyp(> lower molars, one regarded as M2 includ(>s nine ])lates;
H. pentlandi or H. melitensis, the associate.s of the pigmy elephants
other lower second molars include eight to nine plates. M3 in-
(if Malta and Sicily. The geological evidence, as distinguished from
cludes eleven plus plates.
the palseo- zoological, also supports this theory, for it appears prob-
(Op. cit., p. 243): "From this brief account of the remains
able that Cyprus became an island at an early period."
l)rocured of E. creticus, it will be seen that this pigmy Elephant
must have been of slightly larger proportions than E. Cypriotes
Palseoloxodon creticus 15ate, 1907
and approached in size more closely to E. nieliiensis; that is to
Figaro 1129
say, it would have attained a-s a maximum a height of five feet. . .

Pleistui Till', cave deposit near Ca|xi Maleka, island of Crete. All the molars obtained differ from those of the two last-named
Elephas creticus Bate, 1907. "On Elephant Remains from dwarf species in being much lower in the crown this is jierhaps the ;

Crete, with Description of Elephas creticus, sp. n." Proc. Zool. most noticeable feature of the .series. At the same time the teeth
Soc, London, 1907, pp. 238-250. Cotypes. —Nine imperfect are wide, the cement-areas broad, and the enamel simjjlc, though at
cotype molars. Two are figured in the present Memoir (Fig. times broken up into a number of rings. So far as can be ascer-
1129), namely, r.M^ (Brit .Mus. M.9381) and an M2 (Brit. Mus. tained from th(^ scanty amount of material the ridge-formula must
M.9378). Horizon and Locality. Near Cape Maleka, — have b(»en low."
island of Crete. Figures. Op. cit., Pis. xii, figs. 1-3, and—
XIII, figs. 1 and 2. E. creticus: [M 1 ^ M 2 ^ M 3 n+].

COTVPES OF PaL.KOLOXODON CRETICUS

Fig. 1129. Elephas creticus Bate, 1907, Pis. xn, fig. 3, xui, fig. 1. Nine imperfect cotype molars from a cave deposit near Cape
Maleka, Crete. Both molars two-thirds natural size.

(Left) PI. xn, fig. 3: "Crown view of right lower third molar of E. creticus. ([Brit. Miis.| M.9381.)" +13 ridge-crests.

(Right) PI. XIII, fig. 1: Crown view of second lower molar of E. creticus, Brit. Mus. M.9378. +7 ridge-crests.
 V. LES ELEPHANTS NAINS DES ILES MEDITERRANEENNES ET LA QUESTION
DES ISTHMES PLEISTOCENES (VAUFREY, 1929)

Eiifin, par plusicum beaux specimens de Palerme, nous con?iaissons le crane de la plus grande, Elephas mnaidricnsis, crane
qui presente tous les caracteres propres a l' Elephas na^nadicus, c'esl-d-dire a V Elephant antique, notamment le bourrelet suprafronlal
si special. II n'y a done pas de doiite que les formes naincs des ties appariiennent toutes irois au phylum de V Elephas antiquus et
doivent Hre considerees commc des races de cette espece" (Vaufrcy, 1929, p. 209).

The valuable Memoir of Dr. Raymond Vaufrey, "Les Elephants Nains des lies Mediterraneennes" (October,
1929), reached the Osborn Library in December, 1929. The present abstract of the observations and conclusions
of the author should be compared with the previous history and abstracts of the extensive literature cited more or
less fully above. The author, under the guidance of Dr. Marcellin Boule, had the advantage of visiting the various

caves and rock-fissures in Sicily and Malta, in which these fossils occur, and of examining and comparing all the
collections. Among the many geologic observations of value are those in the Grotto of Luparello near Palermo
(Fig. 1130) which show that the lower geologic level of 'Elephas melitensis^ is much more ancient than the higher
level containing 'Elephas falconer i,' from which Vaufrey rightly infers that the extremely dwarfed ' E . falconeri' is

geologically more recent than the middle-sized 'E. melitensis.'

Vaufrey throughout regards the species 'Elephas namadicus' and 'E. antiquus' as synonymous. On the con-
trary, it is shown above in the present Memoir, that 'E. namadicus' Falc. (1846) antedates and is widely different
specifically, if not generically, from 'E. antiquus' Falc. (1847) of western Europe.

We need only abstract those portions of this Memoir wliich bear directly on the following questions: (1)

The time of insulation or separation of these islands from the mainland ; (2) the respective evidence of the rela-

tionship of the dwarfed elephants to (a) the typical 'Elephas antiquus' of western Europe, or to (b) the 'Elephas
namadicus' of India, or to (c) the north-
wardly migrating elephants of Africa;
(3) the time of entry of these dwarfed
elephants into the Mediterranean Is-

lands; (4) the geographic region from

which they entered the islands.

GEOLOGIC AND GEOGRAPHIC

OBSERVATIONS
The elevations and depressions of the Medi-
terranean .'iea leveland the consequent con-
nections of its present islands with the sur-
rounding mainland of Africa and Eurasia are
fvilly treated in the Intrnduction (pp. 1-45), dis-
cussed throughout the descriptions of the
islandsand in the formal text (pp. 45-202),
and summarized in the Conclusions Generates
(pp. 203-216).
1" CiEOGRAPHIC AND FaUNAL RELATIONS OF
''.

PaL.KOI.OXODON KAI,C:f)NKRI MOBK RECENT THAN P. MBUTENSIS THK IsLANos.— Vaufrey (p. 2) adopts Boule's
I'"ig. 1 130. Section of Grotto of Luparello, Palermo, Sicily, .show iiig l)elo\v the "Couehe.s a Ele|)ha.s (1906-1919) theory that in closing Miocene
melitensi.s" at the bottom of the cavern and separated above by a broad from the
.stalagmitic layer
and early Pliocene time (Pontian) the western
"Couches a Elephas Faleoneri." After Vaufrey, 1929, tig. 7, p. 51. The underlying marine phase
l)ii,sin of the Mediterranean was by elevation
("Plage marine"), 136", 40, i.s shown in the darkly tinted horizon at the bottom of the cavern. This
reduced to large lakes between which, over
"Plage marine" may be geologically correlated with the Monastirian stage (Dep(5ret, 1918-1921,
named after Monastir, Tunis), Hrvntiim of the sea level, or insular depression. great isthmuses, freely migrated the Hipparion

1268
 :

THE LOXODONTIN^: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1269

fauna of Leberon, Eppelsheim, Pikermi, Samos, Maragha, etc. ment apparent^es: elles prcsentent toutes trois les memes varia-
After this early Miocene-Pliocene Pontian stage of elevation there tions de la morphologic dentaire, deja relevees par les auteurs
followed a great depression of the land, several hundred meters dans les molaires d'Elephas anliquus. Enfin, par plusieurs beaux
below the present level, or a rise of the seashore lines, which specimens de Palermo, nous connaissons le crane de la plus grande,
completely isolated Europe from the African continent as well as Elephas mnaidriensis, crane qui presente tons les caracteres propres
from parts of Asia. In closing Pliocene time [Villafranchian' stage a VElephas namndicus, c'est-a-dire a I'Elephant antique, notam-
of northern Italy, containing Archidiskodon meridionalis, Hespero- ment le bourrelet suprafrontal
si special. II n'y a done pas de

loxodon ausonius, Parelephas trogontherioides] there is evidence of doute que formes naines des iles appartiennent toutes troi.s au
les
renewed African connections and migrations, but certainly no phylum de VElephas antiquus et doivent etre considerees comme
evidence of a Tunisian-Sicilian land bridge (p. 203), for from the des races de cette espece."
latest hydrographic charts (1916) the Mediterranean Sea, while Synonymy (op. cit., p. 143).
— "Les notions ainsi acquises
400 m. below its present level, was too high to form an isthmus for nous ont permis, en meme temps, de verifier I'identite des especes
intermigrations between Africa and Europe by way of Tunis and nouvelles creees en Sardaigne par Forsyth Major, en C'rete et a
Sicily. Moreover, the absence of a Tunisian-Sicilian land bridge, Chypre par Bate. II devient evident que le squelette de La Mari-
or, in fact, of any land connection between Sicily and north Africa menta se compare exactement par la taille au squelette de Luparello
during late Pliocene and Pleistocene times, is proved by the et se rapporte done a VElephas melilensis, que VElephas crelicus
contrast (Pomel, 1895, Boule, 1899) between the Pleistocene fauna par la morphologic et les dimensions des molaires s'assimile par-

SlCILY Malta Sardinia Cyprus Crete

E. antiquus var. insularis
E. mnaidriensis ref. (crania; etc.). . . E. mnaidriensis type E. mnaidriensis ref.
E. melitensis ref E. melitensis type =E. lamarmorae type =E. crelicus type
E. falconeri ref E. falconeri type =E. Cypriotes type

of north Africa, of Algeria, and of Tunis and that of the Mediter- meme race, enfin que les diagnoses dentaires
faitement aussi a cette
ranean Islands, summarized by Vaufrey (pp. 204-206) as follows: de VElephas Cypriotes et de VE. Falconeri sont .si analogues qu'il
(a) The Pleistocene fauna of north Africa is totally different from n'y a aucune raison de les designer sous deux noms differents. II
that of Europe or of the Mediterranean Lslands; (b) the late convient done de parler d'Elephas melitensis de Sardaigne et de
Pleistocene entry ofEuropean forms into north Africa was not via Crete et non d'E. Lamarmorx et d'E. crelicus, d'Elephas Falconeri
a Mediterranean land bridge between Sicily and Tunis (p. 207) but de Chypre et non d'E. Cypriotes. Exception faite, sans doute, pour
by way of land connections along the eastern Mediterranean shore quelques individus de normale, auxquels est dH I'introduction
taille

of Syria and the Suez; (c) whereas we find certain European and du phylum, les Elephants ne sont done represents dans les iles
other Holarctic forms in the late Pleistocene of north Africa, we que par trois races de taille decroissante : E. mtiaidriensis, E.
do not find a single African form in the Mediterranean Islands, melitensis et E. Falconeri. D'une part, la morphologic dentaire,
more especially in the island of Sicily; (d) the fossil remains notamment en ce qui concerne les caracteres des figures d'abrasion,
formerly attributed to the African elephant {E. africanus) belong nous interdit de consid^rer les races naines comme appartenant a
rather to aberrant specimens of 'Elephas mnaidriensis' ;
(e) thus plus d'une espece; d'autre part, les caracteres craniens d'Elephas
rejecting (p. 208) all previous suggestions of north African relation- mnaidriensis, ainsi que la presence dans les trois races d'une
ship, Vaufrey summarizes his conclusions as to the geologic age, premiere molaire inf^rieure a racine unique, designent cette espece
characters, and relationships of the dwarfed elephants as below. comme 6tant indubitablement Elephas antiquus.
—
Insulation (op. cit., p. 181). "De meme qu'en Sicile, la
faune a Elephants nains nous apparait done, d'une maniere
[Three Dwarfed Species or Races Only]
gen^rale, comme contemporaine du dernier Interglaciaire, et son
"Cette demonstration, toutefois, ne pouvait se fairc qu'en extinction comme s'^tant produite pendant la derniere periode
reprenant sur de nouvelles bases la question des Elephants nains; glaciaire au moment de la formation des couches qui la renferment.
elle n'e<it done pas ete possible si le hasard ne m'avait fait d^cou- L'absence de toute trace de I'Homme dans ces couches confirme ce
vrir a Luparello de nouveaux documents. II ressort de cette etude que nous ont appris d^ja les grottes siciliennes; Sicile et Malte
que les Elephants nains appartiennent a trois formes de tallies nous apparaissent une fois de plus comme des finisterres."
diff^rentes: Elephas mnaidriensis (hauteur au garrot: environ Ic. As to geologic age, Vaufrey concludes as follows (p. 209)
i", 90),£. 7neh'te»sis (hauteur au garrot: i", 40) et E. Falconeri "Rappelons enfin que les couches a Elephants nains de Sicile et
(hauteur au garrot: environ 0™, 90), caracterisees par la presence de Malte datent vraisemblablement de I'epoque wurmienne,
a la mandibule d'une premifere molaire [Dp2] a racine unique, 6poque a laquelle I'El^phant antique avait, sauf en Italic meri-
particularity qui n'existe que chezl'filephant antique, a I'exclusion dionale (Romanelli), disparu d'Europe occidentale et que, dans le
de toutes les Par ce caractere comme
autres especes d'filephants. gisement de Luparello, la plus petite des trois races {Elephas
I)ar la morphologic des molaires, ainsi que par leur presence dans Falconeri) etait superposee a celle de taille intermediaire {E.
les memes gisements, ces trois formes apparaissent comme etroite- melitensis)."

'[See footnote 1 on page 1049 above regarding the possible Lower Pleistocene age of the Villafranchian. —Editor.]
 —

1270 OSBORN: THE PROBOSCIDEA

2. Relationships. — The traditional opinion, seemingly 108) 13-14, laminar freciuence 3,'.| (type pachygaiial) to 6 (tyi)e
shared by Vaufrey, is that the dwarfed elephants of the Mediter- endioganal). (
"lencTal characters (p. 110): Except for
specific
ranean Islands should be regarded as the stranded deseendants of a difference in dimensions, (a) the general characters are similar
the well-known early and late Pleistocene European six»eies 'Ele- to those of the typical 'Elepha.s anliquitu,' with more feeble develop-
phas aiitiquus.'which roamed over the Mediterranean lands before ment of theenamel foldings correlated with reduction in the size of
they were broken up into islands. Vaufrey does not distinguish the molars; (b) a.s in 'E.falconeri,' there are two divergent tenden-
between 'E. namadicus' Falc, 1846, of India, and the typical 'E. cies in the grinding teeth, one of the fine enamel type {email mince)
antiqitus' P'alc, 1847, of western Europe; his comparisons and with compres.sion of the lamina, the other of the thick enamel
measurements are chiefly with those of the late Pleistocene [3d type (email epais) w'ith spreading of the laminse, corresponding
I nterglacial] stages of Weimar and Taubach (see Fig. 1088 of the respectively with the broad and short molars and the long and
l)reseiit Memoir), which he designates as 'Elcphan antiquus normal,' narrow molars described by Leith Adams. (5) Skeleton: Portions
selecting mean measurements between the larger and the smaller of mandibles, vertebrae, femur, ulna, radius, attributable to
dimensions given by Pohlig (cf. Vaufrey, op. cit., p. 138 see Tables — 'Elephas melilensis' ; discovered also in Cyprus by Bate, in the
below). cavern of Mnaidra by Adams, and in La Marimenta, Sardinia

Comparative Measurements (cf. Vaufrey, 1929, p. 138)

Measurements in Millimeters I. [Dp 2] II. [Dp 3] III. [Dp 4] IV. [M 1] V. [M 2] VI. [M 3]
3
E. Falrnncri 10
.

1,

E. melilensis 1, 30
E. mnaidriensis 1, 60
E. anlir/uus normal 1, 82

E. Falconeri 3- 3X
f
E. melilensis 7

E. mnaidriensis 3 -3X
 9 : ,

THE LOXODONTIN^: DWARFED SPECIES OF THE MEDITERRANEAN ISLANDS 1271

email epais with wide compression (Fig. 37,5 Fig. 1132 of present — region, a considerable period after the disappearance, during Inter-
Memoir). (4) The summary of dental characters (p. 131) is as glacial Epoch 3 or Riss-Wiirm interval of the Alps, of the north
follows: (a) A certain number of 'E. mnaidriensis' molars from the German Hesperoloxodon aidiqHu.s germanicus, and probably after the
Sicilian horizonsapproaches the classic structure of tlie continental disappearance of the southerly H. antiquus italicus; (5) the ridge
'E. antiquus,' namely, "couronne elancee et etroite, lames d'epais- formulae which Vaufrey assigns to 'E. falconeri' and to 'E. meliten-
seiu' moderee, email pliss6, souvent avec une expansion losangique sis' are collective; these 'collective ridge formulae' may belong to

mediane moder6e"; (b) from this central type, however, arose one or more specific stages, whereas in a certain stage of evolution
the ridge formula is constant; thus Vaufrey may not be right in
regarding as the same species the typical 'E. falconeri' of Malta and
the typical 'E. Cypriotes' Bate of Cyprus, which has a lower ridge
formula; again Vaufrey's ridge for-
mula of 'E. melitensis' differs widely
from that which Leith Adams finally
assigned to the same species; the
ridge formula which Vaufrey assigns
to 'E. mnaidriensis' namely, ^^^yf
is much below that of the Upper
Pleistocene species of Taubach and
Weimar (Fig. 1088), i.e., germanicus,
Fig. 27. — Sch^mas montrant les grandeurs relatives des defenses des races naines insulaires et de 1 Ul^phant M3 +
-1
italicus,
B
M
3 Ys+; it is also
antique normal. below that of the Lower Pleistocene
I, Elephns antiquus Falconeri de Luparello 2. E. ; a. melilensis de Luparello ; 3, £. a. mnaidfiensis des 'E. antiquus' typical of the Forest
Puntali 4. £ antiquus de Tilloux (d'apr^s M. Boule). ~ 1/20 de la grandeur naturelle.
M
;
Bed, 3 ieH-17 (6) these facts seem
>

Rolative sizp, thickness, and curvature of the incisive tusks in the thre( dwarfed subsix"cies of to render highly improbable the
Fig. 1131.
tlie Mediterranean I.slands, one-twentieth natural size. After Vaufrey, 1929, fig. 27, p. 115. descent of the dwarfed elephants
The 'Elephas
antiquus' of Tilloux (4) is found with some individuals of this species. from the large and specialized west
European 'E. antiquus,' or from the
variations, some of which accent the median sinus, broaden the lower Upper Pliocene or the progressive Pleistocene stages, and
ridge-plates, thicken the enamel, while others narrow the ridge- render more probable the descent of these dwarfed elephants from
plates, diminish the median sinus, and thin the enamel. The east African forms, such as the 'Elephas antiquus recki' Dietrich,
latter reachan extreme degree of thinness and plate compression 1916, M
3 nrrfTi. or the Algerian 'E. jolensis' Pomel, 1895, 3 75, M
which remind us of the 'Elephas armeniacus' Falc. = Parelephas [ or the Algerian 'E. atlanticus' Pomel, 1879, 3 i"|nT- M
armeniacus of the present Memoir]. In brief, there exist in the Finally, it appears probable that at least two separate lines of
cavern of Puntali all intermediates between these extreme types, phyletic descent are represented in these dwarfed elephants of
but none approaches the loxodont type of 'Elephas africanus' or of
the 'Elephas priscus' of Falconer.

OSBORN (1930) SUBSTITUTES A THEORY OF AFRICAN

DESCENT BY WAY OF EASTERLY RATHER THAN
NORTHERLY LAND CONNECTIONS
While recognizing the valuable and important observations of
this Memoir, we are unable to accept many of Vaufrey's conclu-
sions, for the following rea.sons: (1) The cranium of the dwarfed
elephants cannot be derived from that of the typical 'Elephas
antiquus' of western Europe, from which it differs very widely; (2)
all the known Upper Pliocene and Pleistocene stages of 'E. anti-
quus' described in detail in the present Memoir, namely, 'ausonius,'
'antiquus' and 'italicus,' are too highly
(typicus), 'germanicus'
specialized in their ridge formulae and other characters to give rise to
these dwarfed Mediterranean species, which are primitive in ridge
formulae, and related to Palxoloxodon namadicus in cranial Fig. 1132. Two types of molars (A) and (B) figured by Vaufrey, 1929,
structure; (3) the traditional opinion of 'E. antiquus' origin is not fig. 37, Nos. 4, 5, 6, as belonging Ui 'Elephas mnaidriensis'

sustained by our present knowledge of the highly diverse characters (A) Nos. 4 and 6, from Shantiiin and Puntali respectively, of the 'type
endioganal' {email mince); (B) No. 5, from Puntali, of the 'type pachyganal'
of the African,west European, and south Asiatic members of the
(email ipais). In Osborn's opinion the 'type endioganal' (email mince)
Loxodontinae the dwarfed elephants date from the closing
; (4) belongs to a different species from the 'type pachyganal' (email epais). One-
stage of Glacial time, namely, IV GLACi,\L = Wiirm of the Alpine sixth natural size.
1272 OSBORN: THE PROBOSCIDEA

tlicMediterranean Islands, a.s illustrated by Vaufrey (p. 109, fig. dans le i)remier ca.s a des molaires larges et court(!S (variete A de
23), namely: (A) with coarse enamel and ridge-jilates widely Leith Adams), dans le .second a des molaires longues et etroites
separated, that is, with a low laminar frequence of 3'm in 10 cm., (variete B de Leith Adams)."
corresponding with ^'aufrey's 'type pachyganal' {email epais), Osborn's Conclusions (1930). (1) Whereas the progossive —
known west European stages neither of Hesperoloxodon antiquus
nor of the Indian Palnoloxodon namndicus api)ear to be ancestral to
the dwarfed Mediterranean species, we seem to find on the con-
tinent of Africa a number of wore primitive nX&gcsr, of Palseoloxodon,
with more primitive ridge formulae and long narrow grinding teeth,
which further knowledge may prove to be ancestral to the dwarfed
Mediterranean types. This will depend on the discovery of
a cranium ancestral to that of 'Elephas melitensis,' which, in turn,
resembles that of 'E. namadicus' of India.
(2) The true 'mnnidriensis,' 'melitensis,' and 'falconeri' seem
to constitute a single phylum of regressive degeneration or dwarf-
ing of the 'endioganal' or email mince type. But intermingled with
the most ancient and larger 'mnaidriensis' stage in certain of the
caverns seem to be members of other phyla of the 'pachyganal,'
or email epais type.

Fig. Molars referred by Vaufrey to Elcphas = I'al:Foloxu<liiH\

1133. [

and Benphisa, Malta, (No.6), sfiowiiig

mdilftisis from liUparello, Sicily, (No. 1),
respectively widely separated ridge-plates, i.e., low laminar frequence, with
coarse enamel ('type pachyganal') and close laminar frequence, with fine
enamel ('type endioganal'). After Vaufrey, 1929, fig. 23, Nos. 1 and 6, one-half
natural size. See also Fig. 1 132 of the present Memoir.

e.g., figure 23.1, and (15) with fine enamel and close laminar fre-

quence, corresponding with X'aufrey's 'type endioganal' {cimiil

mince), e.g., figure 23,6. This distinction of (A) and (B) is in
accordance with the failure of all earlier authors clearly to dis-
tinguish the phylum Parclrphrts and tlie i)hylum Arrhidiakodon
from file j)hylum Lo.rodunt(i, which in the present Memoir are
sliown to be absolutely di.stinct from each other even at the close of
Pliocene timi'. Tims w(> caimot su])port the author's union
(pp. 109, 110) of these (A) and (B) types in the single species 'E.
Fig. 26. — Figure montrant Ics grandeurs relatives
des cubitus des trois races naincs A'Elephas antiquus.
melitensis,' in which (p. 110) ho i.s erroneously .supported by Leith
De gauche a droite : E. a. mnaidriensis, E. a. meliten-
Adams (1874. )>. 35) as follows: "Comme chez r£lci)hanf antique, sis (cubitus gauches), H. a. Falconeri (cubitus droit, avec
il y a deux tendances dixcrgentes. Tune vers la realisation d'un un fragment du radius). — 1/6 de la grandeur naturelle.
type a email mince a\'ec re.s.serrement des lames, I'autre vers celle
d'un tyjje a email epais avec 6cartcment des lames, aboutissant Fig. 1 131. t'iiia' of (lie throe species. ,\fter Vaufrey, 1929, fig. 2(1, p. 113.
 - — —

VI. ANCESTRAL STAGES OF PAL/EOLOXODON IN AFRICA

Whereas most previous writers, including Vaufrey (1929), derive the Mediterranean dwarfed insular species
from Elephas antiquus of western Europe, we would be inclined to regard them as dwarfed insular derivatives of
those extinct African species of Palseoloxodon, with which the genus Pilgrimia is synonymous. On a large scale
'Elephas zulu' Scott' resembles the types of Elephas antiquus Recki Dietrich, 1916, of Elephas jolensis Pomel,
1895, and of Elephas atlanticus Pomel, 1879, also of the dwarfed species of Malta, Elephas melilensis Falconer,
1862, 1868, 'Elephas falconeri' Busk, 1867, and Elephas mnaidriensis Adams, 1870. From the subjoined table of
ridge formulae it appears that the dwarfed Mediterranean species differ from the west European species of 'Elephas

antiquus' [
= Hesperoloxodon antiquus] but are closely similar to the African species of Palseoloxodon.

Firstly. — Since the first studies for this Memoir were begun (1900) evidence has rapidly accumulated not
only to demonstrate that the Order Proboscidea originated in Africa, as recited in previous chapters of the present
Memoir, but to render it probable that most of the separate genera and subfamilies also originated in Africa
instead of Asia as formerly supposed.

This apparently is true of the Archidiskodon phylum and now begins to be apparent in the Palaeoloxodon

phylum, because Africa and the Mediterranean Islands reveal stages of Palseoloxodon more primitive in molar
ridge formulae at least than any hitherto found in Eurasia, as displayed in the following comparative table com-
piled from previous records and type fornuilae:

Middle to Upper Pleistocene of India Elephas [Palseoloxodon] namadicus Falc. and Caut., 1846, 1847 M2 .^p^ M3 1 6

Lower Pleistocene of England Elephas [Hesperoloxodon] antiquus Falc. and Caut., 1847, 1857 M 2 11
12
M 3 [|~ 17
^^_ _

Malta Elephas [Palseoloxodon] mnaidriensis Adams, 1870 M 2 yf

10
IJ)
M 33 12-13-14
1 g - 1 3 - 1 4

Elephas [Palxoloxodon] melilensis Falconer, 1862 M 2 |^q9-1

M 3Q \12
Tl/r

Elephas [Palseoloxodon] falconeri Busk, 1867 [riilge formula after

Vaufrey, 1929]
10-12
M 2 ^^^ M Q3 3-14
l\/r
14
1

East Africa Elephas [Palseoloxodon] recki Dietrich, 1916 M 2 j^ M3

Algeria Elephas [Palseoloxodon] jolensis Pome\, 1S95 M2 M3 1 3

Elephas [Palseoloxodon] atlanticus Pomel, 1879 M 2 y^^ M 3 j~

Zululand Elephas {Loxodon) [Loxodonta] zulu Scott, 1907 M2— M 3 ,^_ ,

East Central Africa Elephas [Loxodonta] zulu ref. from Kaiso Bone Beds M2— M 3 y^
Recent Loxodonta africana M 2 -^ M 3 jy^t
Secondly. —The above ridge formulae demonstrate that the recent Loxodonta africana is more primitive in

dental ridge structure than any of the known Upper Pliocene or the Pleistocene species of Palaeoloxodon conse- ;

quently L. africana cannot be descended from any known typical species of Palseoloxodon but may have sprung
from a more primitive ancestral form still to be discovered.

Thirdly. — It is a striking fact that the broad conspicuous 'loxodont sinus' of the recent Loxodonta africana is

rudimentary or absent in all [see footnotes below, also pp. 1286-1288, this chapter] the extinct species of elephants
thus far described from Africa or Eurasia. This absence or rudimentary condition of the 'loxodont sinus' removes
certain extinct Pliocene and Pleistocene species from close relationship to the typical recent Loxodonta africana

and relates them rather to the dwarfed species of the Mediterranean Islands which we also have grouped under
the name Palseoloxodon (syn. Pilgrimia), as distinguished from the typical Palseoloxodon namadicus and Hespero-
loxodon antiquus.

'^[Elephas zulu Scott referred to Loxodonta by Professor Osborn (see Osborn, 1934.925, p. 2). — Editor.]
^[See Loxodonta prima Dart, 1929, and L. africana var. obliqua Dart, 1929, below, this chapter, pp. 1287, 1288. Editor.)

1273
1274 OSBORN: THE PROBOSCIDEA

SYSTEMATIC DESCRIPTION OF AFRICAN SPECIES

Palaeoloxodon atlanticus Pomel, 1879 race d'hommes primitifs places par la mythologie dans le massif
Figures 1047, 1135, 1136 du Nord de I'Afriquc." He designated the locality in 1895 {op. cit.,
Pleistocene. Xorth Africa, Ternifine near Mascara, Algeria. p. 42): "J'ai d^sign^ .sous ce nom, daas diver.ses publications,
une espece tres particuliere d'elephant decouverte dans des stations
Specific Characters (Osborx, 1928). —The cotype second
de la pierre eclatee a Ternifine, pres de Mascara, retrouv^e pres
inferiormolar (Fig. 1135), length 240 mm., breadth 60 mm., broadly
du village de la S6nia, par M. F^ningre, dans des fouilles pour
resembles Falconer's type of Elephas antiquits (Fig. 1075) except
fondations et, plus r^cemment, dans la caverne aux hippopotames
that the median 'loxodont sinus' is more widely expanded, re-
de Pointe-Pescade." In the same publication {op. cit., p. 51) he
sembling the conditions shown in Falconer's aged referred 'Elephas
defines the animal: "U elephas atlanticus etait arme de defenses
priscus,' (Fig. 1076) the ridge formula,
; 2 \^, agrees with his M robustes."
identification of the cotype molar (Fig. 1135) as a "P6nultieme (5*^)
Osborn, 1930: Pomel's detailed description (1895, pp. 42-51,
molaire inferieure"; counting the two 'talons' as ridges, it agrees
Pis. vi-x) indicates a ridge formula (including 'les talons') for
with the ridge formula at present known of Hesperoloxodon aniiquus
this species of:
(typicus), namely, M2 j-f
M
3 ffjl^i^. The following is Pomel's
main ridge formula, omitting talons (1895, p. 51), of: M 2 »4. io-w=i2 (length 240 mm., breadth 60 mm.). See page 44 of

Pomel, also figure 1135 of the present Memoir.

E. atlanticus: Dp 2 | Dp 3 f Dp 4 I M 1 ^ M 2 f| M 3 f|. M 3 "-^J-f^ll^ (length
li - I 2 - m mm., breadth fl mm., height of middle
plate 5^ mm.). See pages 43 and 47 of Pome), also figure 1136
History. —Pomel first named this species in 1879 but did not
of the present Memoir.
figure it until 1895.
Thus Palseoloxodon atlanticus, while having a relatively low
ridge-plate formula, namely, M 2 ^^ . ^ q . ^^ M3 J^rrlrll, is a very
large elephant, equaling in size the typical Hesperoloxodon antiquus
of the Forest Bed (Fig. 1088), namely, 3 \V-, length of inferiorM
molar 315 mm., height of middle ridge-plate 126 mm.
Counting the large anterior and posterior talons (PI. viii),
the ridge-plate count of the cotype is 2 p^, of the referred M
superior molar from Ternifine 3 M —
Consequently the ridge-
.

plate count may be actually higher than that given by Pomel.

V
Cotype R.Mj of Pal.«oloxodon .\tl.\.\ticus. One-third Natural Size
Fig. 1135. Elephas allanlicus Pomel, 1879. One of the coty[)es of Pomel,
1895, PI. VIII, fig. 1: "Ponultieme (o") molaire inforieure, vue par la couronne,
J^,et montrant le canal sjinphysaire, tres obtus; Ternifine;" identified as
a right inferior molar, r.Mj. [Op. cit., p. 44): "J'ai fait figurer, PI. viii, fig. 1
et 2, une magnifiqiie molaire tres bien conservoe, en partie encore contenue
dans son os mandibulaire; die a 240°"° de longueur aver une largeur de 60""°
au milieu; elle compte dix lames avec deux talons."
Observe: (1) That the central evaginations of the loxodont sinus' Fig. 1136.Referred third superior molar of the left side, l.M^, of Elephas
resemble those of the Hesperoloxodon antiquus, figured as 'Ekphas priscus' by atlanticus Pomel. After Pomel, 1895, PI. Vlll, fig. 3: "Derniere molaire
Falconer in 1868 and reproduced in our figure 1076; (2) that the ridge formula suporieure, vue par la couronne; Ternifine; K. Musee d'Oran."
isji-lO-YiOT 12, instead of 10 as given in Pomel's description of 1895, p. 51;
This very large superior molar (op. cit., 1895, p. 47), length 324 mm.,
(3) that the measurements are, lengtli 240 nun., breadth 60 mm.
breadth 84 mm., exhibits a ridge formula of 3 M —
the anterior and posterior
;

ridge-plates rise to the surface of the crown and are described by Pomel as
"talons."
Elephas atlanticus Pomel, 1879. "Ossements d'El^phants et
d'Hippopotames decouverts dans une station pr^historique de la
plainc d'Eghis (province d'Oran)." Bull soc. geol. France, Vol.
Palaeoloxodon jolensis Pomel, 1895
VII, Ser. 3, 1879, p. 51. Cotype. —Second inferior molar,
r.M2. Musee d'Oran, Algeria. Horizon and Locality. — Figure 1137

Ternifine, Mascara, Algeria. —

Cotype Figire. Pomel, 1895, Pleistocene. Algerian seacoast. North Africa.

PI. VIII, figs. 1,2. This is the .second North African species described by Pomel,

Pomel figured this specimen as above in 1895, but he named it in the same year (1895) in which he published his type figure of
in1879 (1879, p. 51) "Je me crois des lors autoris6 a donner a cette
: Elephas [
= Palseoloxodon] atlanticus.
forme particuliere, actuellement disparue de la region atlantique, Osborn, 1924: Pomel's type figure of M3 of the left .side ex-
mais seulement depuis une ^poque trfes-rapproch^e des temps hibits the ridge formula M
3 y-^^, a formula inferior to that of
historiques, le nom d'Elephas atlanticus, rappelant le nom de la Hesperoloxodon aniiquus, but exceeding that of Palxoloxodon
 :

THE LOXODONTIN.'E: ANCESTRAL STAGES OF PAL.EOLOXODON IN AFRICA 1275

a//a7j//r(/.s, lamiiiiir froqiicncy 14 ridge-plates in 30 cm.; it may from Pomel's

inferior molar, l.Mj, clearly distinguishes this species

be an ascending pi'ogression above P. ailanticus. 'Elepha.i atlaitticu.'^.' It is of and very primitive,

very small size
Elephax jolensis Pomel, 1895. "Paleontologie Monographies. as shown by the comparative measurements given by Pomel (cf.
Les Elephants Quatornaires." Carte Cieol. de I'Algene, 1895, p. 1895, p. 38)
32. Type. —Third left inferior molar, I.M3. Ecole des
Sciences, Algiers. Horizon and Locality. — (Op. cit., ex- Length Breadth Ridge-plates Height
planation of PI. V, fig. 3): " . . . trouvee a la ferme Beausejour, l.M,. M3 Max.
dans I'ancienne plage emergee," Algeria, North Africa. Type Antiquus [Elephas
—
Figure. Op. cit., PL v, figs. 3 and 4.
—
Type Description. {Op. cit., p. 32): "La piece la plus im-
portante pour la caract^ristique de cette forme est representee PI.
V, fig. 3 et 4. Elle a ete trouvee a la ferme de Beausejour. en aval
du Krober-Roumia, dans la plage marine soulevee et a ete donnee
a I'Ecole des Sciences par M. Maupas, con.servateur de la Biblio-
theque nationale d'Alger. Elle est remarquable par sa longueur,
par son etroitessc et par .son incurvation. Elle compte treize lames
dont la derniere, tres etalee, peut a la rigueur passer pour un talon

Type Left Third Inferior Mol.\r of Pal.eoloxodon jolensis

Fig. 1137. Tvi>e LM3 Pomel, 1895, PI. v, figs. 3 and
of Eleplms jolmsis
4, one-half natural "Derniere molaire inferieure, vue de profil, )2i
.size: 3.

trouvee k la ferme Beausejour, dans I'aneienne plage emergee." 4. "La meme

dent, vue par la couronne."

(Pomel, 1895, p. 38): "Le fossile algerien qui a treize lames a .son arriere-
molaire inforieure, a eette dent longue de 140'"'" avee une largeur de couronne
de 36 et une hauteur, a la septieme lame, de 70°""."

ambigu. . . . Cette dont est assez particuliere par sa forme en long

ruban, dont la partie posterieure est comme flanquee par une
double rang^e de disques de detrition s^pares de la partie moyenne;
ceux du cote exterieur se dMoublant plus habit uellement que les
interieurs. Cette disposition est assez semblable a celle qu'offre la

derniere molaire inferieure de Velepha.^ antiquus; mais dans

dent est beaucoup plus grande; elle presente quatorze
celui-ci la
lames pour 30''°Me longueur (ou meme quinze lames pour une
14°""."
longueur de 27*"") au lieu de treize lames pour
Csborn, 1928: This narrow thirteen ridge-plated left third
 :

1276 OSBORN: THE PROBOSCIDEA

sondern im zoologischcn Sinn als Bezeichnung einer neiien geo- ers der Ms den Elefanten von Oldoway vom Urelefanten, wahrend
graphisfhcn Ra.ssc verstandon wissen."
. . . die Stosszahne beide Elefanten wiederum einander niihern (s.

Dietrich adds {op. cit., p. 75) : "Nach den vorhandenen Lang- unten)."
knochen durfen wir fiir die Oldovvayer Elefantenrasse Riesenmasse Dietrich's ridge formula of Elephas aidiquus recki is as follows:
annehmon, so die Hohe im Widerrist mit 4m, die in der Kruppe
iiicht weniger, sondern eher mehr, denn die Riickenlinie des E. M 2 ri M 3 Tyrrw.
antiquUs senkt sich nicht wie beim Mammut von der Schulter nach
hinten ab, sondern steigt wie beim afrikanischen Elefanten in OVER-ESTIMATED HEIGHTS (DiETRICH, 1916, P. 76, 1924, P.
der Kruppe wieder an. Der Riesenwuchs des E. antiquus Recki 24).— Dietrich concludes (1916, p. 76): "Als wichtigstes Ergebnis
bestatigt die Tatsache, dass E. antiquus neben (nicht hinter) E. hebe ich hervor, das.s die grossten Hohen aller fossilen Elefanten
um 4 m herum liegen, dass es keine 5 m hohen Elefanten gegeben
hat, dass nicht einmal Dinothcrium diese Hohe erreicht hat und

Lectotype op Pal/Eoloxodon recki

Fig. 1 138. Leetotype I.M2 of Elephax
antiquus Recki Dietrich, 1916, Taf. i, fig.

2, one-tliird natural size. Geologisch-

Paliiontologisches Institut der Universitat
Berlin, XVII 1384. From the Oldoway-
Tuffe, Serengetisteppe, northern Tangan-
yika Territory.

meridionalis zu den grossten Elefanten, ja zu deu grossten Land- dass schliesslich die starksten Elefanten der Gegenwart hinter
.sjiugern iiberhaupt, zu rechnen ist." denen der Vergangenheit an Mass und Gewicht zuriickstehen."
Reck (1914, pp. 306 and 307) makes the following observa- According to Dietrich's measurements and estimates of
tions: "Cber die Merkmale der Unterkiefermolaren (M2 und M3) height, the 'Elephas antiquus recki' is something over 4 m., namely,
macht Dr. Dietrich zur vorlaufigen Charakterisierung der neuen 4030 mm., in skeletal height, or 13 ft. 2% in., exceeding the tallest
Elefanten folgende Angaben living African elephant by 2 ft., or 610 mm. This estimated height
of 'E. antiquus recki' is inferior to that of the 'E. antiquus german-

1. Lamelle plattig mit gebogenen seitlichen Umrissen. icus' of Taubach, in which the humerus is about 1300 mm. in
length. According to this estimate both these animals exceed the
2. Schmale Seitenpfeiler, breiter Mittelpfeiler der Lamelle,
height of the tallest of the known imperial mammoths, namely,
ausgesprochener Mamillenbau. Archidiskodon maibeni (3826 mm. or 12 ft. 6/8 in.), as shown in the
3. Massig dicker, miissig gekrauselter Schmelz. comparative shoulder heights of living and extinct elephants
(Chap. XVI, fig. 912). Dietrich's shoulder height of the 'E.
4. Weite Distanzierung der Lamellen durch breite Zementin- antiquus germanicus' of Taubach, estimated from the humeral
tervalle; doch ist die Tiilerweite ein schwankendes
length (1300 mm.), is still greater, namely, 4000+ mm. He
Merkmal. assigns the same height (4000+mm.), estimated from the humeral
5. Verhaltnismassig geringe Lamellenzahl (14 bis 16 bei M3 length of 1300 mm., to the 'Elephas antiquus' (typicus) of the
11 bei Mj). Lower Pleistocene (Mauer) of western Europe.
The length of the humerus, namely, 1290 mm., of Hesperoloxo-
6. Breite Zahnkronen der letzten Molaren (M3)." don antiquus of Upnor, as described above, agrees very closely
with Dietrich's estimates and measurements, giving an estimated
". . . Die Unterschiede zum Elefa.s Zulu Scott endlich liegen shoulder height of 3700 mm. or 12 ft. 1% in. this accords well with ;

haupt.sachlich in den Punkten 1, 3 und 4 der Charakteristik des the fact that the Upnor skeleton is not full grown, because the
Oldowayelefanten. Einigc der Ziihne, niimlich die vorletzten second inferior and .superior molars, M
2, are still in full use.

Molaren erinnern stark an E. antiquus Falc. aus dcm europiiischen By the practical agreement of the Dietrich humeral measure-
Diluvium; doch entfernt die Gesamtheit aller Merkmale, besond- ments with those of Forster Cooper (1928), we may reach a nearly
 THE LOXODONTIN^: ANCESTRAL STAGES OF PAL^OLOXODON IN AFRICA 1277

correct estimate of the maximum height of 'Elephas antiquus In comparison with the estimates of Archidiskodon
brief, in
germanicus' of Taubach, namely: and taking
(imperator) maibeni, as given in the legend of figure 912,
Humeral length 1300 mm. advantage of Dietrich's excellent table of comparative measure-
Skeletal height at shoulder 3729 ments (1916, p. 76), we may estimate the shoulder height of the
Height in the flesh at shoulder 3965 skeleton (in millimeters) of the following species as below.

Comparative Skeletal Measurements and Heights with those of 'Elephas antiquus recki'
(Dif:TRicH, 1916; Osborn, 1930)

The length of the humerus affords one standard means of estimating the skeletal height at the shoulder. In Archidiskodon
maibeni the humerus is relatively shorter and the estimated shoulder height is made from the entire forelimb.

Millimeters
 -

1278 OSBORN: THE PROBOSCIDEA

Pal8eoloxodon(?) andrewsi Dart, 1929' attention has been frequently drawn in dealing with all these
Figure 1139 ])rimitive mammoths. Further, despite the worn aspect of these
Gong-Gong, Vaal River, South Africji. Middlr terrace, GO-80 feet. fragments, Ido not think that the true height was very greatly in
Lower? Pleistocene. excess of what the fragments portray."
(Osborn, 1934.925, p. 14): "A primitive or ancestral member of "In form, measurements and general appearance, the tooth

the Palaeoloxodon group may be tliis problematic A . andrewsi Dart approximates more closely to A. loxodonloides than to any of the

(Fig. 5 = Fig. 1139 of the present Memoir]), a type which on types hitherto discu.ssed, but the coarser folding of its enamel, its
[

sectioning and very careful reexamination by the present author,

proves to be distinct both from A. planifrons (Fig. 4 [ =Fig. 876 of

the present Memoir)) and A. siibplanifrons (Fig. 1 [=Fig. 875 of I Centimeter,

the present Memoir]). The fragmentary type, f.LMa, displays the
following characters: Ridge plate height =48 mm. est., e.stimated
number of ridge-plates = 6. Feeble pre-sinus fold ; very prominent
po.st-sinus fold. Valleys V-shaped. Estimated length = 164 mm.
Estimated breadth = 83 mm., estimated index = 50. Enamel thick,

crimped. It has been extremely difficult to restore this terribly

shattered type specimen and deduce its outstanding characters . . .

from the enamel folds which certainly belong at the front and back
of the third inferior grinding tooth."
Archidiskodon andrewsi Dart, 1929. "Mammoths and Other
Fossil Elephants of the Vaal and Limpopo Watersheds," So.
African Journ. Sci., Vol. XXVI, pp. 711-713. Type.—
McGregor Mus. by Mr. Luke Rademan and presented
435, found
to the Museum by Mr. II. Rees. Cast Amer. Mus. 26968. [Re-
garded by Professor Osborn as a left third inferior molar, I.M3.]
Horizon AND Locality.— "Gong-Gong, Vaal River 'Deep diggings
at a depth of 80 feet,' probably Middle Terrace. . . . ?Pliocene."
Lower? Pleistocene.- Type Figure. —Op. cil., 1929, fig. 14, p.
711 [
= Fig. 1139 of the present Memoir].
Type Description.— (Dart, op. cii., 1929, p. 712): "The
fragments together embrace a length of 110 mm., the greatest
width is approximately 80 mm. and the greatest height 60 mm. i/^nat.size McQregor Mus. 455
They include one fragment with two and a half plates and another Type. Archidiskodon andrciosi. Dart. 1929
with one and a half plates and a talon. The enamel is very thick
(4-5 mm.). The plates are 17-22 mm. wide and display anterior Type of ?Pal«oloxodon andrewsi
Fig. 1 139. R^'stored type of "Archidiskodon" andrewsi Dart, 1929, McGregor
and posterior buttresses as well developed as in A. [Archidiskodon]
Mu.seum 435, Kimberley, South Africa; ca.st Amer. Mus. 26968. Crown view
loxodonloides. Here the width of the plates is in the region of 25 mm.
restored witli estimated 6)1 ridge-plates. Observe subequal cement
and dentinal
and by virtue of the abutment of adjoining l)uttresses u]ion one areas; pre- and two anterior ridges in contact; sharply
and post-sinus folds
another the interlamellar cemcntiun on either side of the tooth is V-shaped valleys between enamel ridges which penetrate about half the crown,
separated into two parts as in A. loxodonloides." thus differing widely from the enamel ridge-plates of A. suhplanifrons. This
is i)rovisionally r(^ferred to Palxoloxodnn. One-lialf natural size. After
"The enamel is markedly folded despite its thickness as in
O.sborn, 1934.92.-., fig. :,, p. 13.
A. loxodonloides, but in coarse fashion. The distance between
mid-ixjints of adjacent ridge plates measured laterally is VA in.

The reduction (7-11 mm.) of interlamellar cementum on the

medial side of the tooth and its almost comjjlete absence on the reduced iieight, its lack and almost total absence of interlamellar
lateral side in one fragment and its total ab.sence in the other cementum demonstrate we have here
tlu; virtual certainty that
fragment together with the rapid contraction of the lamellar a still more iirimitive form not- very far removed from the Stego-
thickness (1 in. at base, y> in. at grinding surface) are witness to the donts and apparently ancestrally related to A. vanalpheni and A.
Stegodontine characteristics of the molar— a featm-e to which lo.rodonloides."

'[It will be noted that the present spi'cies has been <iuestionably referred by Professor Osborn to Pnholoxodon (see Chap. XVI, pp. flS t, O.S.j above, where it

is provisionally listed under both tlii^ Metarchidiskodon griqiia and I'id^oloTodon Iransvaaleiisis groups). Editor.) —
^Cf. Table VIII, p. 984, of the present volume.
 —

THE LOXODONTIN^: PAL^OLOXODON AND LOXODONTA OF SOUTH AFRICA 1279

Palffloloxodon hanekomi Dart, 1929 "The discovery of this exceedingly progressive form, whose
I'igure 1140 hypsodont analogies are to be sought in America, renders it also

Delpoort's Hope, Vaal River, Soutli Africa. Level unknown — Pleistocene. probable that further discoveries of a convergent evolutionary

"Mammoths and Other nature between American and African forms will yet be made. . .
Archidiskodon hanekomi Dart, 1929.
.

Fossil Elephants of the Vaal and Limpopo Watersheds," So.

African Journ. Sci., Vol. XXVI, pp. 713-715. Type.—
McGregor Mus. 2930. "?Third right upper molar." Horizon
—
AND Locality. "The old river bed of the Vaal River at a depth
of 20 feet at Delpoort's Hope [South Africa]. ?Upper Pleisto-
cene." Type Figure.— Op. cil., figs. 15 and 16, p. 713 =Fig. [

1140 of the present Memoir].

Type Description. — (Dart. o-p. at., 1929, pp. 714, 715):
"This tooth was found by Major C. J. Hanekom on 23 October,
1927. It differs entirely in form from any hitherto discovered in

South Africa being considerably shorter (200 mm.) than it is high

(259 mm.). All specimens previou.sly found have a length in

excess of their height except in one case (A. [Palseoloxodon] trajis-

vaalensis) in which both measurements are virtually identical.

The great height of this tooth (lOh inches) equals that of the mo.st

gigantic form of Archidiskodont known, namely, A. imperator

Leidy of America. The greatest breadth (102 mm.) is virtually
identical with that of A. [P.] sheppardi (100 mm.), but is not .so

great as that of A. [P.] transvaalensis (110 mm.). The number of

plates also visible is 17 (Fig. 15 [

= Fig. 1140 of the present Mem-
oir]), that is, approximately identical with that of A. [P.] trans-

vaalensis but in excess of A. [P.] sheppardi, where the number of

lamellae is thirteen."
"In virtue of the hypsodonty of the specimen (Fig. 2 [=Fig.
1140 of the present Memoir]), there are only eight and a half plates
in wear (Cf. figures of Mammonteus prigimius [primigenius] com-
pressus, Osborn 1924). The irregularity of plate number in the
worn area due to the curious nature of the enamel pattern —which
forms an especially distinctive attribute of this particular species Type of Pal.eoloxodon hanekomi
Fig. 140. Type ?third right superior molar of Archidiskodon hanekomi
is unparalleled in any South African mammoth type and recalls in
1

Dart, 1929, from Delpoort's Hope, Soutli Africa. McGregor Mus. 2930.
some respects the irregularities of plate arrangements seen in
After Dart, 1929, figs. If) ami 16, p. 713, side and crown views respectively.
Mammonteus (Elephas) primigenius (Cf. Zittel's 'Textbook of One-third natural size.
Palaeontology,' Vol. 3, Fig. 349, 1925)."
"The individual lamellae are as broad (11-15 mm.) as those of
A. [P.] sheppardi, but do not reach so great a breadth as those of Actually, being discovered at a depth of twenty feet in the river
A. [P.] transvaalensis and the interlamellar cementum, unlike that bed gravels at Delpoort's Hope, it would seem that it belonged to
in both those forms, is about as broad (10-12 mm.) as the lamellae. an older geological horizon than A. [P.] transvaalensis and A. [P.\
The lamellar walls are virtually parallel with one another, but only sheppardi, which were found at a depth of 4 to 5 feet in the river
in the two anterior lamellae do they run entirely across the tooth bed gravels higher up the river at Bloemhof. Mr. Lowe's dis-
and then only in a zig-zag fashion, demonstrating that each lamel- covery that these river bed gravels at Bloemhof present two dif-
lar plate is virtually bifid to about 3 inches from its root base. ferent gravel strata of very variable depth, separated from one
These characters, coupled with the positive though not excessive another by two different phases of stone implement culture,
fore and aft compression of the seventeen ridge plates and vertical point to the necessity for similar investigations at Delpoort's
elevation of the tooth or hypsodonty, show that we have in South Hope and the possibility, despite the depth at which it was found,

Africa a further phase of mammoth evolution beyond what has that A. [P.] hanekomi belongs to a still more recent phase of the
heretofore been recorded for this country." Pleistocene than does A. [P.] transvaalensis."
1280 OSBORN: THE PROBOSCIDEA

Palseoloxodon yorki Dart, 1929 p. 294) is characteristic of a milk tooth. Hence the tooth must
Figure 1141 have been an adult one. ... It approaches in number of ridge
Near Christiana, South Africa, Vaal River. Middle(?) Pleistocene. plates, enamel pattern, and tooth form to E. prisrus Falconer and
to E. Irogo/itherii Pohl. There Ls exhibited a very slight tendency
Pilgrimia yorki Dart, 1929. "Mammoths and Other Fo.s.'iil
especially in the anterior crescents to throw out an anterior and
Elephants of the Vaal and Limpopo Watersheds," So. African
posterior buttress, and hence to provide a vestigial loxodont sinus.
Journ. Sci., Vol. XXVI, pp. 719, 720. Type.— McGregor
The width of each lamella varies from the almost parallel regions
Mus. 4074, cast Amer. Mus. 22727. "Right lower (?third) molar."
where it is 12 mm., to 20 mm. in the region of the slight buttresses,
Foimd by Mr. Alf. York during diamond digging operations in
1927. Horizon and Locality. "Lowest stratum of the — but the lamellae are well separated by cementum across the whole
grinding surface; the whole tooth is well encased in the abundant
river bed gravels on the farm Vanasswegenshoek, O. F. S., below
cementum."
Christiana, [South Africa], at a depth of within 6 feet. . . . VPleistn-
"The numerical seriation of this grinder of the lower jaw is
eene." Type Figure.— Op. ciL, figs. 20 and 21, p. 719
a matter of question. It may be a second molar, but in any ca.se it
[Fig. 1141 of the pre.sent Memoir].
indicates by its reduced width, its symmetrical anteriorly-concave
crescents, its slight tendency to single fore and aft buttresses, its
more delicate enamel (2-3 mm. thick), and finer crimping a dis-
tinctive type, so far as South Africa is concerned, whose relation-
ships are with the most primitive Pilgrimia = Palseoloj:odo?i].
[ It
is comparable in simplicity, but not in size, with E. falconeri
Busk. ... Of known fo.ssil forms outside Africa, it seems to ap-
proximate most closely to Elephas (Loxodon) priscus Falconer, not
only in its dimensions and regularly crescentic plates with sinuous
outline as viewed laterally, but also in the actual number of plates
and the tendency to buttressing. Indeed, the appearances are
such that A. [Palaeoloxodon] yorki might well be ancestrally related
to the E. {Loxodon) priscus of Falconer, as in A. [P.] yorki the
processes leading to the production of such forms as E. priscus and
E. Irogontherii seem to be incipient."
"A. [P.] yorki is the simplest of the Pilgrimia [Palxoloxodon]
type yet recovered in Southern Africa."

Palseoloxodon wilmani Dart, 1929

Figure 1142

Below Christiana, Vaal River, South Africa. Middle(?) Pleistocene.

Pilgrimia tvilmani Dart, 1929. "Mammoths and Other

Fossil Elejihants of the Vaal and Limpopo Watsrsheds," So.
African Journ. Sci., Vol. XXVI, pp. 720-722. Type.—
McCJregor Mus. 4075, cast Amer. Mus. 22726. "Left lower
(?third) molar." Horizon and Locality. "Lowest stratum —
Type of Pal.koi.oxodon yokki
of river bed gravels between the farms Vanasswegenshoek and
Fin. 1141. Type of Pilgrimia yorki Dart, 1929, from near Christiana,
Bloemheuvel, Transvaal, below Christiana at a depth of within
South Africa. "Right lower (?thir(l) molar" (McGregor Mus. 4074; cast
Amer. Mus. 22727). After Dart, 1929, figs. 20 and 21, p. 719, crown and side 6 feet. .?Middle Pleistocene."
. Type Figure. Op. cil., —
views, one-half natural size. fig. 22, p. 721 [Fig. 1142 of the present Memoir).
Type Description.— (Dart, op. cit., 1929, pp. 721, 722):
"This tooth fragment attains only the very narrow width of 70 mm.
or 2% in., but the height of the largest plate is 128 mm. It is,
Type Description. — (Dart, op. cit., 1929, pp. 719, 720): therefore, both narrower and higher than Pilgrimia [Pala'oloxodon]
"This elephant tooth (Figs. 20, 21 = Fig. 1141 of the pre.sent
[ yorki. Unfortunately the tooth is incomplete, the length of the
Memoir]), like the other remains found, i.s stony in nature and fragment being 115 mm. and including seven plates only. There
completely fossili.sed. It is a relatively diminutive tooth being must have been in the original tooth at least one and probably two
180 mm. long, 79 mm. broad, 1 15 mm. high, ixjssessing 9 plates, all more plates jjostcriorly and another one, or two, more plates
of which are There is no trace on the posterior plate or
in wear. anteriorly. The ridge plate number was ... ±11 andapi)arently
talon of a depre.s.sion arising from the pressiu'c of a tooth advancing in excess of P. yorki (.see Fig. 22 [Fig. 1142 of the jircsent

behind it, as Falconer has pointed out {Pnlaeonl. Memoirs, WA. II, Memoir]). But Ihc^ most striking dissimilarity between the two
 .

THE LOXODONTIN.^: PAL^OLOXODON AND LOXODONTA OF SOUTH AFRICA 1281

teeth lies in the character of the enamel plates wliich in this tooth Palseoloxodon kuhni Dart, 1929
are narrower (6 mm. in narrowest parts to 11 mm. in the slightly Figure 1 143
wider central parts), and are not so widely separated by the
interlamellar cementum. Further, they run almost directly Piii(^l Estate, South Africa. ?River bed gravels. Middle(?) Pleistocene.

transversely across the grinding surface, displaying little if any Pilgrimia kuhni Dart, 1929. "Mammoths and Other Fossil
tendency to cresccntie outline on this aspect of the tooth. The Elephants of the Vaal and Limpopo Watersheds," So. African
lamallae are distinctly narrower laterally than medially (that is, Journ. Sci., Vol. XXVI, pp. 723, 724. Type.— McGregor
the reverse of found in the modern African elephant and in
what is
Mus. 4144, cast Amer. Mus. 22725. "?Lower left molar." Hori-
Loxodonta antiqua zvlu) where they are somewhat bulged, termi- zon AND Locality.— [P. 723] "Pniel Estate. ?River bed gravels. .

nating in an anteriorly curved expanded portion almost as wide ?Pleistocene." [P. 724] "The site of discovery at Pniel must be
(9 mm.) as the widest central portion of the lamella." fairly comparable with that at Vanasswegenshoek, and this indi-
"The same tendency towards a median buttress as was found cates the necessity for stratigraphical study in the Pniel region in
in P. yorki is also encountered here, but to still less extent. The order to establish the correlation which is likely between these
enamel also is more delicate (1J^2 mm. thick) and is more finely two widely separated sites along the Vaal valley.
is the more This
crimped than in P. yorki. In sharp contrast also with P. yorki, necessary in view of suggestions, which have been put forward
the tooth is not ensheathed in cementum but the ridge plates from time to time, that the gravels of the river bed and of the
stand out to a depth of M to J^ inch, except for the upper 1 to IK terraces higher upstream are of a different age from the same gravel
inches of the lateral aspect, nor are the ridge plates much less
obvious on the medial aspect of the tooth."

Type of Pal.kolo.xodon kuhni

Fig. 1143. Type of Pilgrimia kuhni Dart, 1929, Pniel Estate, South
Africa. "?Lo\vcr loft molar" (McGregor Mus. 4144; cast Amer. Mus.
22725). After Dart, 1929, fig. 24, p. 725, one-third natural size.

lower down stream. In the solution of this question the identifica-

tion of elephant teeth types with particular gravels promises to be
of premier importance." Type Figure. — Op. cit., fig. 24, p.
725 [Fig. 1143 of the present Memoir].
Type Description. — (Dart, op. cit., 1929, pp. 723, 724):
"The fragment consists of and the greater portion of
four plates
the anterior talon. Its dimensions (great(>st length 96 mm.,
greatest width 75 mm. and greatest height 100 mm.) indicate to
some extent its relationships. Its width and height are fairly
closely comparable with those of certain specimens of E. antiquus
Recki and in its general morphology it forms the clo.sest approach

to that form which I have hitherto seen in this country. In that

form, however, the plates, as seen from the grinding aspect, are
usually definitely erescentic, the horns of the crescents facing
Type ok Pal.eoloxodon wilm.\ni forwards. In this specimen they run transversely across the tooth.
Fig. 1142. Type of Pilgrimia loilmani Dart, 1929, from bolow Christiana, The anterior talon is vestigial, the most anterior true plate is

Transvaal,_;South Africa. "Left lower (?third) molar" (McGregor Mus. continuous across the grinding surface, the second has a smaller
4075; cast Amer. Mus. 22726). After Dart, 1929, 22, p. 721, one-half left and a larger right island, while the third and the fourth have
fig.

natural size.
a central large island and internal and external smaller islets.
These features indicate that the total length of the tooth could not
"There can be no question that this is another species of have been great and the total number of plates in the tooth is
Pilgrimia = Palseoloxodon], which I shall denominate Pilgrimia
[
unhkely to have exceeded seven or eight (see Fig. 24 [Fig. 1143
wilmani, in honour of Miss Wilman, Director of the McGregor of the present Memoir])."
Memorial Museum at Kimberley, who has been personally respon- "The numerical seriation of the tooth is very doubtful. The
sible for retrieving so many scientific treasures from the Vaal enamel is very thin (2 mm.) and is, on the whole, very finely
valley and permanently safeguarding them in that institution." crimped."
 ;

1282 OSBORN: THE PROBOSCIDEA

Palffioloxodon archidiskodontoides flaughton, 1932 both showing very worn grinding surfaces. From the fact f liat the
Figure 11 11 socket into which the right tooth bounded posteriorly by
fits is

Sydncy-on-Vaal Breakwater, bed of the Vaal River, Soutli Afriea. a wall of bone with a thickened and somewhat rugose ventral
Level unknown —Pleistooenc. surface, and further that posterior to this wall is another, unfilled,
Pilgrimia archidiskodoiituidcti Haughtoii, 1932. "On Some socket, it is concluded that the teeth are the second permanent
South African Fossil Proboscidea," Trans. Roy. Soc. So. Africa, molars."
Vol. XXI, pp. 4-8. CoTYPES. ". —
a fragmentary skull . . "The left molar has nine plates and a posterior talon preserved.
with two worn molars that can be fitted into thoir sockets, a The three anterior plates are worn down to the roots, so that the
humerus which lacks the proximal end, the glenoid portion of enamel of each has coalesced with that of the succeeding plate.
a scapula, and the fairly complete left side of a pelvis." McGregor The degree of wearing is greater on the inner than on the outer
Museum, Kimberley, South Africa. Horizon and Locality. — side of the tooth. As preserved, the length of the tooth is 148 mm.
"Sydney-on-Vaal Breakwater in the bed of the Vaal River, lOK the greatest breadth was probably 94 mm.; the maximum thick-
feet 'below the maiden ground.'" Type Figure. Op. cil., — ness of the plates (4th and 5th) is 16 mm.; the height of the
Pis. i-iii =Fig. 1144 of the present Memoir].
[ posterior plate is about 145 mm. The length of the posterior six
—
Type Description. (Haughton, op. cit., 1932, pp. 4-8): plates is 107 mm. ; and six and a half plates occur in a length of
"In June 1930, Miss Wilman, the Curator of the McGregor 100 mm."
Museum at Kimberley, forwarded to the writer for examination "There is no evidence of any tendency to the formation of
a collection of fragmentary mammalian remains which had been a median loxodont buttress."
found by Mr. J. du Preez, jun., at the Sydney-on-Vaal Break-
water in the bed of the Vaal River, 10}^ feet 'below the maiden
ground.' In February 1931, thanks to the mediation of Major
H. P. Tuckey, Miss Wilman received a further consignment,
from the same locality, of material which was in the posses-
sion of Mr. J. J. de Jong. Among the heterogeneous collec-
tion of bones which made up these two consignments, the
writer discovered some Elephantid remains a fragmentary —
skull with two worn molars that can be fitted into their
sockets, a humerus which lacks the proximal end, the glenoid
portion of a scapula, and the fairly complete left .side of a pel-
vis. These are the bones that are bi'iefly described and
figured here."
"Teeth.— Both the left and right upper molars are preserved.
1^

-4

-^
i''

r 4
TvPK OK Pai,.k()U)xoi)on archidiskodontoides

Fig. 1144. 'I'ypv iti I'ilgriinia archidixhuhmUniks Haughton, 1932, from Sydney-on-Vaal Breakwater, I)(m1 >f Vaal H.iver, South Africa.. McGregor
Museum, Kimberley, South Afriea. After Haughton, 1932, PI. i, fig. 1, and PI. ii, figs. 1 and 2.

PI. I, fig. 1. ?Seeoiid .superior molar. About one-half natural size.

PI. II. Part of right humerus, anterior and posterior vi(,'ws.

 ' — —

THE LOXODONTINyE: PAL^OLOXODON AND LOXODONTA OF SOUTH AFRICA 1283

"Tlic oiiamel Is of medium thickness (2.5-3 mm.), and is by no thicker vcntrally than dorsally. In front of the molar, the palatal
means strongly crimped." surface of the maxilla is strongly hollowed —a feature that is not
"In the right molar the posterior talon is missing. Five plates seen in the skulls of Loxodonta africnna that have been examined."
occupy 80 mm., and there is very little interlamellar cement. The "Humerus. — Part of a right humerus ispreserved, the bone
plates have their anterior and posterior faces roughly parallel, lacking the ]jroximal end. Its chief features can easily be discerned
there being no marked antero-posterior thickening of the plate from the The bone is
illustrations given. much more robust than
towards the root. Laterally there is a distance of 15 mm. from that of Loxodonta africana. The shaft is thicker, and the deltoid
mid-point to mid-point of succeeding enamel ridge plates." crest much stronger and more prominent. The supinator crest is,
"Although they are wider than any of the three species de- proportionately, of about the same length; but, as in P. antiqua
scribed by Dart, these teeth seem —
on account of their high lamel- recki, its border is straight and ha-s not the curved form with upper
lar frequency —
to fall within the limits of the genus Pilgrimia protuberance that is seen in L. africana."
[
= Palxoloiodon]. ... In Pilgrimia the frequency is 5-6 in P. yorki "In actual size the bone is smaller than that of P. anliqua
and 6.5 in P. wihnani; and the length-lamellae ratio is 18.2 in the recki and that of P. antiqua andrewsi}^' Its chief measurements
former and 16 in the latter. In the form under discussion the are:

II. III. IV.

mm.
Maximum width, distal end 265-|- 350 350 227
Width, trochlear articular surface 230 285 306
Height of top of supinator crest above distal end 267 410? 450 275
Minimum thickness of shaft 125 145 167 80

figures are 6.5 and 15.4-15.6 respectively. The greater width of Comparative measurements of other forms are given in parallel
the two teeth described here tends to remove them from Pilgrimia columns.
as hitherto known in South Africa. Width, however, is a somewhat II P. a7itiqua recki from Oldoway, East Africa.
varying quantity within a species, as is evidenced by a study of Ill P. antiqua andrewsi [
= Hesperoloxodo7i antiquus] from
a large number molars of Loxodonta africana where the width
of Upnor, England.
varied from 60 mm. to 84 mm., and is dependent on the position IV Loxodonta africana from Addo."
and seriation of the tooth. Teeth from the lower jaw are always "Although the head of the bone is not preserved, it can be
narrower than teeth from the upper jaw; and the upper molars of seen that the inner border is more strongly bowed than in L.
Pilgrimia yorki will certainly be found to be broader than 79 mm., africana, and it is concluded that the caput humeri was larger or
which is the measurement given by the type lower molar. A stood away further from the main axis of the bone than in the
breadth of 94 mm. is obviously not an impossibility for a Pilgrimia modern form."
upper tooth." "Pelvis. —
The left side of the pelvis is almost entire, lacking
"That the form dilTers from any of the teeth of Pilgrimia most of the ischium, the symphyseal region, and the upper iliac
hitherto described from South Africa is evident. In its breadth crest."
and in the tapering form of its plates it recalls certain forms of "In shape, the ilium differs considerably from that of P.
Archidiskodon such as A. yorki; and I propose, therefore, for the antiqua recki a» figured by Dietrich. The pre-acetabular portion of
sake of convenience, to designate it by the new name Pilgrimia the iliac plate is considerably longer; at the narrowest part of the
archidiskodon loides.' shaft the outer face of the bone is practically flat, and above this
"Skull. — A portion of the maxillae, into which the two molars the outer face is far less concave than in the East African form.
preserved. It shows that the palate was narrow and vaulted,
fit, is From the ilium of L. africana this bone differs very considerably."
and that the molars diverged posteriorly. In front the width "The pubis has a much stouter shaft than in either P. antiqua
between the molars is about 50 mm., and at the back it was about recki or L. africana. In view of the broken nature of the borders of
90 mm. The vault of the palate is 60 mm. above the grinding the specimen it is not possible to give many measurements; but
surface of the teeth. The alveoli of the tooth in use and the suc- the following show some of the differences between this specimen
ceeding molar are separated by a wall of spongy bone, which is and that of a male L. africana from Addo.

P. archidiskodontoides

Greatest length of iliac blade

Distance from supra-acetabular ridge to spina anterior.
Thickness of ilium at spina anterior
Height of acetabulum
Width of acetabulum
1284 OSBORN: THE PROBOSCIDEA

Palseoloxodon transvaalensis Dart, 1927 Apart from these points, the whole atmosphere of each tooth is
Figure 1145 from that of the other. Fro7n the lateral aspect, despite its
tlifferent

From near Blooinliof, lowest terrace, Vaal River, Soutli Africa. Pleistocene. great height, the larger tooth has a massive squarish appearance,
markedly different from the triangular form of this aspect in the
Archidiskodon transvaalensis Dart, 1927. "Mammoths and
smaller tooth. The individual plates are obscured in the larger
Man in the Transvaal." Nature (Supplement), December 10,
tooth by a dense covering of cement over approximately the entire
1927, No. 3032, i)p. 41-48. Type.— Right third superior
lower half (115 mm.) of this surface, and almost filling up the inter-
molar, r.M', length 246 mm., breadth 110 mm., posterior height
247 mm. —
Type Figure. Op. cii., p. 47, fig. 6 (right), and
lamellar clefts in the upper half, which are thus rendered broad and
shallow. . . . From the grinding aspect, the larger tooth has a more
p. 48, fig. 7 (left).
bulging ovoid appearance than the narrower and more ellipsoidal
appearance of the smaller tooth, as follows also from their re-

ji Nat. Size
% Nat. Size

Dart's Tytl ok 1'al.eoloxodon transvaalensis' Dart'.s Type ok Pal.'eoloxodon sheppardp

Fig. Archidisknihn trnnsvndli-nxis Diirl. Type right. liird superior
111."). I Fig. 1 I4t). Archidiskodon shrppardi Dart. Type left third superior molar,

molar, r.M', of the lowest Vaal River terrace gravels, near Bloemhof, Soiitli l.M', of the lowest Vaal River terrace gravels, near Bloemhof, Soutli Africa.
Africa. Pleistocene. Modified after Dart's pliotograplis (if. Dart, 1927, figs. Pleistocene. Modified after Dart's photographs (cf. Dart, 1927, figs. 6, left,

6, right, and 7, left), one-third natural size. Cast Amer. Mus. 27769. and 7, right), one-third natural size.

This molar displays from 1 14 ridge-platcs, the maximum height of the This molar displays from 1-13 ridge-plates, as numbered, the height of the
eleventh plate being 247 mm. eleventh ridge-jjlate as preserved being 202 mm.

Typk Description.— (Dart, np. cit., j). 47): ". . . the larger spective length and width measurements. Desi)it(> the fact that
tooth wa,s greater in every dimension. Although the anterior the total lengths of the two specimens are virtually idoiiticai, there
plates are missing, its present length equals that of the other tooth, are three (and perhaps more) additional lamellae in the larger

It is 10 mm. broader and it is 50 mm. higher in its posterior portion. than in the smaller tooth."

'[Original specimens, formerly in the Ethnology Museum, University of the Witwatersrand, destroyed by fire. — Editor.)
 : — ,

THE LOXODONTIN^: PAL^OLOXODON AND LOXODONTA OF SOUTH AFRICA 1285

Palseoloxodon sheppardi Dart, 1927 from this level and described by Hodkinson. The expectation of
Figure 1146 human remains there of great importance phylogenetically cannot
Near Blocmhof, lowest Vaal River terrace, South Africa. Pleistocene. be exaggerated, since this mastodon bed must reach back to a rather
Archidiskodon Sheppardi Dart, 1927. "Mammoths and Man early phase of the Pleistocene.
in the Transvaal." Nature (Supplement), December 10, 1927, The 200-300 ft. terrace and any fossil mammalian remains or
No. 3032, pp. 41-48. —
Type. Left third superior molar, evidences of human culture at that level, which would ajjpear to
l.M', length 246 mm., breadth 100 mm., posterior height 202 approximate if not actually to be situate entirely within the
mm. Type Figure. —Op. cil., p. 47, fig. 6 (left), and p. 48, Pliocene, must be of premier anthropological importance."
fig. 7 (right).
Type Description. — (Dart, op. cil., pp. 47 and 48): "In NOTES ON ARCHIDISKODON [=PAL^OLOXODONI TRANSVAAL-
the smaller tooth the lamellse are covered with cement over ap- ENSIS DART, 1927, AND ARCHIDISKODON = PAL^OLOXOD0NI [

proximately the lower third (60 mm.) only of this surface, and SHEPPARDI DART, 1927
above this point the interlamellar clefts are extremely deep (5-10 Referring to the site of discovery of two molars from the third
mm.) and narrow in appearance. The same features are repeated or youngest terrace, or river bed gravel, Vaal River, near
Bloemhof
on the medial aspects of the teeth. the individual lamellae are
. . . South Africa (see above, p. 944, also Fig. 823),
Dart continues
appreciably wider in the larger than in the .smaller tooth, so that (op. cit., 1927, p. 43) "The animal remains forwarded from
:

the interlamellar cement is more abundant in this tooth than in the Bloemhof consisted of two right [right and left] upper molars. . . .

former. There are also differences in form between the lamellae of [p. 45] The present teeth have nothing in common with either
both teeth, in that the narrower lamellae are more recurved poster- Loxodonta griqua Haughton or with Elephas {Loxodonta) Zulu
iorly at each end of the lamellae, and the laminae of each lamella Scott. They represent an entirely different category altogether. . .

possess a narrower or finer enamel and are more nearly parallel in [p. 46] They may, therefore, be included with those of E. meri-
the smaller than in the larger tooth. ... In the smaller tooth the dionalis, E. planifrons, and E. imperator amongst the southern
pattern presented by the digitations as they come into wear is mammoths in the generic phylum Archidiskodon of the subfamily
considerably different, there being only three plates showing trans- Mammontinae. ... [p. 45] They are respectively right and left
ition stages from separate digitations to full plates. The most upper molars, and both arc presumably third molars. Their
posterior of the three shows three small islets, the second shows measurements are as follows
four somewhat larger islets, and the third one very large medial Right. Left.
islet and one small lateral islet. The remainder of the lamellae form [P. transvaalensis] [P. sheppardi]
complete single islands across the grinding surface of the tooth. . . . Third upper molar length 246 mm. 246 mm.
Sufficient differential characteristics between the teeth have been " breadth 110 " 100 "
discussed to indicate that it is highly improbable that they belong- " " " height of posterior
ed to the same species. Even if we looked upon the smaller tooth as portion247 " 202 "
being a second molar from a female, it is scarcely likely that there The larger right molar (type of Archidiskodon
superior
would be so great a gap between the two. In view, therefore, of the [Palseoloxodon] transvaalensis) was found in a separate pit from the
Sheppard brothers' interest in securing the teeth and forwarding smaller left superior molar (type of A. [P.] sheppardi); they
them for examination, I will denote the type indicated by this certainly belong to different individuals. There are thirteen
smaller upper and presumably third molar as Archidiskodon ridge-plates in the left molar (sheppardi) and from sixteen to
Sheppardi, sp. nov." eighteen in the right molar (transvaalensis): "The lamellae
Dart concludes (p. 48) "It is evident, therefore, that the
: [i.e., ridge-plates] are not compres.sed but are broad (17-19 mm.

southern mammoths were represented in southern Africa by at in the central portions, 13-15 mm. near the margin, and 15-17
least two distinct species of the genus [Archidiskodon], and that the mm. at the medial margin in the larger tooth, i.e. right molar;
line of their southerly migration is shown by the recovery of portion 12-14 mm. in the central portion and 10-11 mm. near the inner
of a tooth of a nearly related species from the depth of 60-80 feet and the outer margins of the smaller tooth, i.e. left molar). The
below the Nile at Khartum. ... It has been shown that the lowest lamellae are considerably broader than the interlamellar discs of
or niamrnoth [Archidiskodon] gravels of the Vaal bed are replete cementum; the cementum is nevertheless abundant in quantity
with evidences of the lower palaeolithic type of culture. They are although not so abundant relatively as in E. planifrons and E.
therefore presumably pre-Bushman in orientation. The only pre- meridionalis types." The enamel is definitely crimped. There is
Bushman type known from extreme southern Africa so far is Bos- no tendency, as seen in E. = Loxodonta] zulu and Lo.rodonla
[

kop man. Containing, as they do, extinct forms of mammalian [

= Metarchidiskodon] griqua, to throw out a median posterior
life, presumptive evidence furnished that these gravels will
there is buttress or 'loxodont sinus.'
yet supply this, and perhaps other hitherto unidentified forms of The author rightly continues [p. 46] that these grinders belong
human-kind, and show them to have been responsible for that to a genus not previously described from South Africa.'
culture. It remains to be determined whether the left (smaller tooth)
The recognition of extinct forms of mammalian life in the represents a different species from the right (larger tooth) ; they
gravels of the river bed further enhances the age of the 60-80 ft. or present marked divergencies from one another in details, as shown
mastodon terrace, and the evidences of palaeolithic culture secured in the accompanying type photographs (Figs. 1145 and 1146).

'[Referred by Professor Osborn (Osborn, 1934.925, p. 2) to Palxoloxodon. — Editor.]

1286 OSBORN: THE PROBOSCIDEA

Loxodonta zulu Scott, 1907 served from the table of measurements that while the breadth of
crown equals the maximum recorded, even of the ui)]ier molars, for
Figures 1147, 1148
any individual of the existing African or Indian species, the length
Zuliilaiid, Simtli Africa, associated with tlic followinp extinct s|)eci<'s:
is considerably less than in these exceptionally gigantic individuals.
The Ilippnixildiniix i)oriik'rosus, Bubalus andersoni, and Upniccros simplicidcns
... It is plain, however, that the number of ridges, not including
of Scott. Pleistocene.
the talons, cannot have been less than 12, and may have been
Specific Characters. — This very largo twelve or thirteen 13. . More significant than the number of the ridges is their
. .

ridge-plated molar, identified by Scott as an I.M3, is relatively shape, and one is immediately struck by the fact that in the fossil
broail, length 265 mm., breadth 85 mm., height of po.sterior portion the pattern is decidedly less loxodont than in the recent African
81 mm.; it ha-s therefore the relatively broad dimensions of re- species; the median expansion of each lamina is less, and hence
ferred superior molars of Elephas antiquus, but the concave side of the successive ridges are more widely separated, while in the less
the iidge-]ilates (pointed forward) proves that thi.s is a third left
abraded ridges the loxodont pattern is not displayed at all, the
inferior molar, as identified by Scott. Consequently the ridge two enamel walls of each lamina being quite parallel The enamel is .

formula is ai)pai('ntly: very thick (pachyganal) and very strongly crimped. The lateral
terminations of the laminse are either rounded or trifoliate, and each
M3 1 2?V ridge has a feebly curved or cre.scentic shape, with the horns
directed forward. The three posterior ridges have only the points
In describing this l'pi)er Pliocene or Pleistocene species, 'Scott of the digit ations exposed, and of the.se there are four to each ridge,
observed that the type tootii does not resemble that of Loxodonta and those of successive plates are arranged in longitudinal rows in
afrirana, becau.se it lacks the characteristic sinus loxodonte. On a manner suggestive of E. meridionalis."
a large scale it resembles the types of Elephas antiquus Recki Diet- Ill .several respects these teeth of E. zulu resemble the curious
rich, 1916, of East Africa, and Elephas atlantiriis Pomel, 1879, of
molars of E. antiquus, which were originally described by Falconer
Algeria, also the small Elephas joknsis Pomel, 1895, of Algeria, as as E. priscus. . . .

well as the dwarfed species of Malta, Elephas melitensis Falc,

1862. 1868, and Elephas iiinaidrsp Adams, 1870. Measurements.
Elephas (Loxodon) zulu Scott, 1907. "A Collection of Fossil
Third lower molar, length 265
Mammals from the Coast of Zululand." Third Rept. Geol. Surv.
width 085
Natal and Zuhiland, 1907, pp. 259-262. Type.— Third right
and left inferior molars, r.M.i, I.M3. Horizon and Locality. — " " " height of posterior portion .. . .081"

Zuhiland, southeast coast of Africa. Type Figure. Op. —

cii., PL XVIII. fig. 1.
PI. XVII. fig. 6, Scott regards this species as closely related to Hesperoloxodon
Type Description. {Op. rit., — p. 261): "Indeed this tooth antiquus and as possibly ancestral to Loxodonta africana. He
might almost be described as intermediate in character between concludes {op. "Should E. zulu eventually prove to be
cit., p. 262) :

Pohlig's two groups, the Arrhidi.sirodonta and the Loxodonta. The the actual ancestor of E. africanus, it would tend to give the latter
tliird lower molar is very large, almost e(}ualling in size that of the a less isolated position, connecting it with Asiatic and Eiuopean
most gigantic modern African elephants, and in shape is elongate species. At all events, it is extremely interesting and important to
and rather narrow, thougli broader relatively than in E. afrirauus, find in South Africa an elephant with so many points of resem-
so much .so as to \('rge upon tiie laticoronate type. It will be ob- blance to species characteristic of the northern hemisphei-e."
Reck (Sitzungsb. Ges. Naturf. Freundc zu
Berlin, 1914, p. 307) remarks: "[1] Die Untcr-
schiede zum Elefas Zulu Scott endlich liegen

hauptsachlich in den Punkten 1, 3 untl 4 der

Charakteristik des Oldowayelefanten. [2] Einige
der Ziihne, namlieh die vorletzten Molaren erinnern
stark iui E. antiquus Falc. aus dem europiiisehen
Diluvium; tloch entfernt die (Jesamtheit aller Merk-
nialc, hesonders d(>r M3 den lOlefanten von Oldoway
\om Urelefanten, wahrend die Stosszahnc beide
't-^'OiCf^
Elefanten wiederum einander niihern (s. unten). [3]

E. Zulu ist der nur aus zwei zusammengehorigcn

Type I.ekt Thiud I>(fbriok Moi^ah ok I.oxf>r)oNTA /.ulu M3 Molaren bekannte ganz vereinzelt dastehende
Kig. 1 147. One of the two tliird inferior molars (left and type of
risht) con.stituting the
Fund eines dem neucn Elefanten noch am niiehsten
Elephas (Loiiiilon) zuln Seott, 1907, I'l. xvni, fig. 1: "I.eft :}rd lower molar, crown-view; . . .

Same tooth as I'l. xvil., fiK- 0." Less than one-half natural size. stehenden Tieres der Elefas hi/sudrieu.s-UcWw. Trot z-
I,en(;lh 2(1.") mm., Iircadth
85 mm. dcm er ])riniitiver uiid au<'h iilter (Altdiluvial?) ist

'(Professor Osborn first thoiiclit this s|>ecies to be referable to I'aLrdloxndim, but in his article of 1934 on the "Primitive' Arckidislcmlun. and I'aliicutdxoilon of

South Africa" (Osborn, 1934.9'2.>, p. 2) he definitely referred Elephas (Loxodon) zulu to Loxodonta. — Editor.)
 THE LOXODONTINiE: PALiEOLOXODON AND LOXODONTA OF SOUTH AFRICA 1287

als dcr Oklowayclefant, ist er doch .schon zu viellamellig, um als 254 mm., breadth 74 mm., and height 142 mm. opposite the 7th
direkte Ahnenform des Oldowayelefanten gelten zu konnen. Frei- plate, there being only 9 plates (6 plates in 6 inches), correspond-
lich ist das Material zu sicheren Schlussen zu diirftig." ing closely with L. africnua and all in wear."
"There can be no doubt that in this narrow-crowned, loxodont-
simulating, few-plated, mountain-inhabiting specimen here depict-
ed, we have a long-sought ancestral type from which the modern
African elephant tooth might reasonably be derived by a progres-
sive widening-out of the loxodont sinus, for which reason I have
named it Loxodonta prima. This fossil is of importance not only in
demonstrating an advancing, though still simple Loxodont tooth
pattern in the Transvaal, but also in revealing Africa, and possibly
South Africa as the evolutionary home of the true Loxodonta. Its

.wa^''

Fig. 1148. Referred Loxodonla zulu (Brit. Mils. 12639). A thirteen Fig. 1149. Tyjje left third inferior molar, I.M3, of Loxixhmla prima Dart,
ridge-plated third left inferior molar, I.M3, from the Kaiso Bone-beds, near 1929, 25 and 26, p. 725, crown and side views respectively. McGregor
figs.

Lake Albert, Afriea. After photogra|)h kindly furnished the present author by Mus. 4077; cast Amer. Mus. 26987. One-third natural size. Found near
Dr. A. Tindell Hopwood (of. Hopnood, 1926, PI. iii, fig. 1). One-fourth Pilandsberg, Transvaal, South Africa.
natural size. Length 272 mm., breadth 65 mm.
habitat in the elevated and relatively waterless Pilandsberg sug-
Loxodonta prima Dart , 1929 Loxodont persistence, namely, their
gests the possible reasons for
Figure 1149 becoming inured to more arid conditions and more active move-
Pilandsberg, Transvaal, bank of Rhenoster spruit tributary of the Limpo- ment, as compared with their more ponderous Archidiskodont
po River, South Afriea. ?Recent. relatives."

Loxodonta prima Dart, 1929. "Mammoths and Otlier Fossil

Elephants of the Vaal and Limpopo Watersheds," So. African
Journ. Sri., Vol. XXVI, pp. 724-726. Type.— McGregor L 0x0 dont a af r icana var. obliqua Dart, 1929
Figure 1150
Mus. 4077, cast Amer. Mus. 26987. "Left lower third molar
Valley of Steel poort River tributary of Oliphants River, Northeast
[I.M3I,fragment of right third molar, separated plates of upper
Transvaal, South Afriea. ?Reccnt.
molars." Discovered by John Mostert. Horizon and
Locality. — Bank Rhenoster spruit tributary of the Limpopo
of Loxodonta africana var. obliqua Dart, 1929. "Mammoths and
River, at a depth of 4 feet on the farm Nooitgcdacht, Pilandsberg, Other Fossil Elephants of the Vaal and Limpopo Watersheds," So.
Transvaal. ?Recent. —
Type Figure. Op. cit., fig.s. 25 and African Journ. Sci., Vol. XXVI, pp. 726-728. Type.—
26, p. 725 [Fig. 1149 of the present Memoir]. McGregor Mus. 4078, cast Amer. Mus. 26988. "Right lower third
Type Description.— (Dart, op. cil., 1929, pp. 725, 726): molar [r.Ms]." Horizon and Locality. "Valley of Steel- —
"The outstanding character of this tooth (Figs. 25, 26 [Fig. 1149 poort River tributary of Oliphants River, N.-E. Transvaal, on the
of the present Memoir]) is its comparability with the living farm Kranzkloof, at a depth of 17 feet. ?Recent." . Type
. .

African species. It provides the first indubitable fossil evidence of Figure.— 0/j. cit., figs. 27 and 28, p. 726 [Fig. 1150 of the present
a very close approximation to the distinct ive lozenge-shaped Memoir].
lamellae of the living Loxodont grinding tooth. The ab.sciicc of Type Description.— (Dart, op. ril., 1929, pp. 727, 728):
such a fossil type up to the present time according to Osborn 'is "This molar, forwarded to me in February, 1928, by Mr. C.
a striking circumstance.' The length of the reconstructed tooth is Howard, was brittle superficially but nevertheless in an excellent
1288 OSBORN: THE PKOBOSCIDEA

state of preservation. It is a woll worn tootli 256 nim. long, 76 mm. "The salient enamel (3-4 mm. thick) Is criniijed, but so
broad, and 137 mm. in hoight, possessing eleven complete and one slightly that the general apjiearance is one of lack of crimping.

rudimentary posterior ridge plates, 12 in all. Its measurements The lateral terminations of the rhombs are not flattened l)ut are

correspond veiy closely with those of Loxodonta prima just de- on the contrary pointed while the medial terminations tend to be
scribed, and also with those of the living form. But its corre- more rounded and flattened in contrast with the classical de-
spondence with the living African elephant is demonstrated
not only by its measurements but also by the number of enamel
plates and the definite Loxodont character of the enamel sur-
faces in wear. Inde(>d, its resemblances are so close as to render
its separation from the exivSting species a matter of question."

Loxodonta africana var. obliqua Dart, 1929. Typo Typb of Loxodonta subantiqu.\
Fig. 1150.
third right inferior molar, r.Ms, from the valley of the Stoelpoort Fig. 11.31. Type of Pilgrimia subantiqua Haughton, 1932, from Delport's
River, northeast Transvaal, South Africa. McGregor Mus. 4078; Hope, South Africa, "ixLssibly a right lower molar, probably the second."
Amcr. Mus. 26988. After Dart, 1929,
ea.st fig.s. 27 and 28, p. 726, McGregor Mus. 4286. After Haughton, 1932, PI. iv, figs. 1 and 2, crown and
crown and sido vicw.s. One-third natural size. side views. About two-fifths natural size.

"Certain features which perhaps are characteristics of variety serijjtion (Falconer, 1868, 'Palaeontologieal Memoirs') of the living
rank, may, however, he referred to. It may be noted that in no species."
case do the median buttresses impinge directly on one another "Whether these characteristics will i)r()ve in the long run
adequate for the i)\H'pose of si)ecies differentiation is diflficuil to
but rather ovcrla]) uniformly to such an extent that the anterior it

buttresses impinge anteriorly on the posterior asjiect of tlu; medial assess. ... In any case, 1 have regarded the featiu-es i)resented as

portion of the ridge plate immediately in front of it, while the sufficiently distinctive to look upon it as a variety and to draw
posterior buttres.ses impinge posteriorly on the anterior aspect of attention to these details in recording the presence of a fossil form
the lateral portion of the ridge plate immediately behind it. In extremely closely related to the living African elephant at great
this the ridge plate rhombs are not regular but markedly
way depth in the valley of the Steelpoort river in the Transvaal."
skew or oblique in outline and api)earance (I'lgs. 27, 28 [Ing. 1150
of the present Memoir)). I should not have regarded this feat Un-
as being of special significance, seeing that a similar degree of Loxodonta subantiqua Haughton, 1932
obliqiiity due to overlapping of the rhombs is depicted in standard
Figure 1151
illustrations of the molars of African elejihants (e.g.
inferior
Owen's 'Odontography,' 1840-1845, PI. 148, Zittel's 'Textbook of
Delport's Hope, near Vaal River, Africa. Level unknown —Pleistocene.
Palaeontology,' 1925, l'"ig. 348), if it were not for the fact that in Pilqrimin xuhnntiqua Haughton, 1932. "On Some South
other illustrations (e.g. r'aicoiicr's 'Paiaeonlologicai .Mcnioiis,' \'ol. African Fossil I'roboscidea," Trans. Roy. Soc. So. Africa, Vol.
II, Plate 0, I'ig. 1) of the same species, there is no cxidcnic of sucii XXI, pp. 8-10. TvPK.— McGregor Mus. 4286, i)resented by
overlapi)ing of the rlioiHl)s. in addition, there is pr("sciil in this Mr. (1. Barren. "The tooth is possibly a right lower molar,
variety the full number of ])lat('s for the living species, together probably the .second." Hokizon ani> Locality. — Deli)ort's
with a vestigial plate remnant posteriorly." Hope, half a mile from the Vaal River, South Africa. Found at
 :

THE LOXODONTIN.E: PAL^OLOXODON OF JAPAN 1289

a depth of 40 foft in the 'higher terrace.' Pleistocene. Type giving a length-lamellae quotient of 19.5. The lamellar frequency
Figure.— Op. cit., PI. iv, figs. 1 and 2 [Fig. 1151 of the present varies from 4.75 in the front part of the tooth to 5.75 in the back
Memoir]. jiart, so that the individual lamellae increase in thickness towards
Type Description. — (Haughton, op. cit., 1932, pp. 8-9): the root. They width upwards. The maximum
also taper in
"The tooth is possibly a right lower molar, probably the second,
thickness of a plate (4th) mm., and the minimum thickne.ss of
is 19
and its is strongly concave in an antero-posterior
grinding .surface
interlamellar eementum is 4 mm. The greatest width (including
direction. Eleven plates are preserved, of which the first has been
eementum) is 92.5 mm., and the maximum width of a plate at the
worn down to the roots, and the last is just coming into wear.
grinding surface 79 mm. The enamel is not thick and is coarsely
There was probably a posterior talon. The plates are fully covered
with eementum l)oth on their medial and lateral sides.
crimped."
The
greatest length of the tooth as preserved, including the posterior "The tooth is low. The greatest height of any plate preserved,
eementum, is 245 mm. Measured in a straight line, the ten and above the roots, is about 75 mm., and it is doubtful if the maximimi
a half plates visible on the grinding siu'face occupy 206 mm., height of any of the posterior plates exceeded 85-90 mm."

VIII. LOXODONTINES OF JAPAN AND JAVA

(C'ontimied from Chap. XIV, pp. 901 to 909, and from \^. 1185 of the present chapter)

In this concluding historical and systematic .section of the Loxodontinse we may review the original as well
as one of the most recent treatments of the far eastern loxodonts of Japan and of Java, at a time when these coimtries
constituted the eastern portion of the Asiatic continent and successively attracted the mastodonts, the stegodonts,
and the true elephants chiefly of the genus Palseoloxodon.

History. — During the years since Dubois described (1908) his 'Elephas hysudrindicus' from the Kendeng for-
mation of Java (now believed to be of Middle Pleistocene age), great changes in nomenclature have been made
which are thoroughly set forth in the following systematic revision of eight species and subspecies' originally
named and more or less fully described by Dubois, Makiyama, and Matsumoto, the total list to our present
knowledge being as follows
Probable Ridge
Original Reference Reference in Present Memoir Geologic Age Formula
Java
Elephas hysudrindicus Dubois, 1908 = Palseoloxodon hysudrindicus Middlc(?) Pleistocene

Japan
Elephas namadicus naumanni Maki-
yama, 1924 = Palseoloxodon namadicus naumanni Middle Pleistocene (?) M 3 j'
Elephas namadicus narnndi Maki-
yama, 1924 = Palseoloxodon namadicus namadi Middle Pleistocene (?) M3 '-^

Euelephas protomamm,onteus Matsu-

moto, 1924 = Palseoloxodon protomammonteus Upper Pliocene (?) M3 r^TTTT
Loxodonia (Palseoloxodon) tokunagai
Matsumoto, 1924 = Palseoloxodon tokunagai Upper Pliocene (?) M3^ 3 -»
Parelephas protomammonteus proxi-
mus Matsumoto, 1926 = Palseoloxodon protomammonteus proximus Upper Pliocene (?) M3^
Loxodonta (Palseoloxodon) namadica
(Yahei) Matsumoto, 1929 = Palseoloxodon namadicus yahei Middle Pleistocene
Jmx. (Pal.) Tokunagai junior, mut.
Matsumoto, 1929 = Palseoloxodon tokunagai mut. junior Upper Pliocene(?) or
Lower Pleistocene M2TTn:.
'[To these should possibly be addedtlie following species described since this text was written and not examined by Professor Osborn: Parelephas \fPalwo-
loxodon] protomammonleus malsumoloi Huheki, 1931, from Mishima, Province of Kazusa, Palseoloxodon yokohamanus Toliunaga, 1934, from Yokohama, and
Palsenloxodon aoiiioriensis Tokunaga, 193G, from Tenjinbayaslii, Aomoii Prefectnie. Editor.] —
1290 OSBOKN: THE PROBOSCIDEA

Whereas Dubois and Makiyama gave merely a preliminary description of the three species named by them
{E. hysudrindicus, E. naumanni, E. namadi), Matsumoto named five species and contributed a succession of short

articles and memoirs, fully listed in the Bibliogiaphy of the present Memoir; also in aid of the jniblication of the
present Memoir, he sent the author in 1924 a letter giving his views at that time (June 24, 1924). His more mature
views are expressed in his two memoirs entitled, "On Loxodonta (Palieoloxodon) namadica (Falconer and Cautley)
in Japan," with six plates (Matsumoto, 1929.1,) and "On Loxodonta {Palaeoluxodon) tokunagai Matsumoto, with
Remarks on the Descent of Loxodontine Elephants," with one plate (Matsumoto, 1929.2).

SUMMARY OF MATSUMOTO'S FINAL OBSERVATIONS AND THEORIES OF 1924 AND 1929

Tlic i)resent author differs widely from Matsumoto's opinions and theories as to phylogeny, geographic
distribution,and nomenclature; palaeontologists are nevertheless deeply indebted to him for his great labors in
setting forth the characteristics, geographic distribution, and geologic succession of tlie Japanese loxodonts.
The following gives a summary of Matsumoto's observations and theories.

Matsumoto, 1924. —We owe to Doctor Matsumoto (letter, June 24, 1924) his geologic section (Fig. 790) of

the fossil bearing formations of Japan and a series of valuable notes on the Japanese forms referable to Loxodonta
[
= Palseoloxodon] namadica. It appears that all varieties of L. namadica from Japan were distinguished by smaller
and narrower grinding teeth than those of the typical L. namadica of the Nerbudda Valley. This relatively small
and relatively narrow molar proportion is characteristic of all the insular species of the genus Loxodonta = Palseolox- [

odon] ; it is also a primitive character of all loxodontines.

According to the 1924 observations of Matsumoto: (1) The parent forms of Loxodonta [
= Palaeoloxodon]
migrated into Japan early in Pleistocene time, namely, in the Calabrian-Villafranchian age, in which occurs an
older type almost similar to Loxodonta [
= Hesperoloxodon] ausonia in its evolutionary stage ; this is perhaps of
Upper Pliocene Calabrian or Villafranchian age; (2) in the succeeding Cromerian-Sicilian stage there occurs an
ascending mutation, the Loxodonta namadica var. naumanni of Makiyama [since made the "subgenotype" of Palse-
oloxodon by Matsumoto], almost similar to the type of L. — Hesperoloxodon] antiqua which occurs in the Cromer
[

Forest Bed; (3) a still more modern type, approaching more nearly the typical L. = P.] namadica of India, [

occurs in the terrace gravels corresponding perhaps with the Monastirian or Tyrrhenian stage of Deperet. In
descending geologic order these species appear to Matsumoto (letter, June 24, 1924) as follows:

III. Loxodonta namadica ?mut. =a final mutation of the youngpr terrace gravels ?Monastirian-Tyrrhenian age. Narrow
of
toothed, like the species naumanni, but thin ridged and thin enameled; loxodoiity absent or very
slight in the upper grinders, and slight loxodonty in the lowc-r grinders; laminar frequency 7-8.

11. Ldj-odonta antiqua var. naumnnni Makiyama = an older type, mutation of the Tokyo beds of ?('romerian-Siciliaii age. Al-
most similar to the species of the Cromer Forest Bed Loxodonta antiqua, as well as to the L. hysudrin-
dicn of ,Ia\'a, in its evolutionary molar stage; laminar frcHjuenc^y 6-6.5.

I. Older type, almost similar to Loxodonta ausonia of the Villafranchian = mutation of Minato, its geograjihieal locality.
Almost similar to L. ausonia in its evolutionary stage; crown low, M.t about 120 mm.; laminar
frequency 6-6.5; perhajjs of ('alabrian-^illafranchian age.

According to the observations of Matsumoto, the narrow-toothed Japanese elephants differ from the typical
broad-toothed Loxodonta [
= Palseoloxodon] namadica types of India and resemble the typical narrow-toothed
L. [Hesperoloxodon] ausonia types of southern Europe, the Mediterranean Islands, and North Africa.
 :

THE LOXODONTIN^: PAL^OLOXODON OF JAPAN 1291

Matsumoto, 1929. — The following passages are taken from Matsumoto's Memoirs of 1929 (1929.1, 1929.2),
entitled

On Loxodonta {Palxoloxodon) namndica (Falconer and Cautley) in .Japan. Sci. Ropt. Tohoku Imp. Univ., Socond Series
(Geology), Vol. XIII, No. 1, (1929.1).
On Loxodonta {Palseoloxodon) lokunagai Matsumoto, with Remarks on the Descent of Loxodontine Elephants. Sci. Rept.
Tohoku Imp. Univ., Second Series (CJeology), Vol. XIII, No. 1, (1929.2).

(Matsumoto, 1929.1, p. 1): "Assistant Professor Makiyama of the Kyoto Imperial University has really

laid the cornerstone of further progress in the study of Elephas namadicus Falconer & Cautley in Japan, of which
two subspecies have been distinguished by him. Subsequently, his E. namadicus Naumanni was selected by the
present writer to be the subgenotype of Palseoloxodon, which is referred to Loxodonta. The writer has now come to
distinguish three races of the species in question in Japan. One of them, corresponding to the subspecies Nau-
manni Makiyama, represents the Lower Pleistocene mutation of the species, while the other two appear to be

characteristic of the Middle Pleistocene. Thus, the racial subdivision of the present species may play a part, the
writer hopes, in the geological correlation of the Japanese Pleistocene."

(Matsumoto, 1929.2, pp. 7, 10): "An interesting archetypal Loxodontine elephant of Japan, named Loxo-
donta (Palseoloxodon) Tokunagai by the writer, as well as the subgenus Palseoloxodon created by him, has not yet
been fully described in a European tongue. The writer here wishes to furnish a description of the elephant with
a few remarks concerning the descent of Lo.xodonts. On this occasion, the writer has the pleasure to express his
hearty thanks to the authorities of both the Imperial Museum of Ueno and the Geological Institute of the Kyoto
Imperial University, by whom he was permitted to study the specimens described in the present report."

"Rise of Pal^oloxodon"
"As .specially noticed by Lydekker [Footnote: 'Lydekker, Brit. Mus. Cat. Foss. Mamm., Pt. IV., 1886, pp
102, 103 & 106, text-fig. 25.'], a small form of very archetypal elephant is recorded from India under the name of

Elephas planifrons. This form appears to be deviant from Archidiscodon in being narrow-toothed, though it

appears to be closely related to the same in having the loxodont sinus of an obtuse type in the grinders. Again, by
the first mentioned characteristic, it appears to be close to and by the second to be deviant from Palseoloxodon.
Phylogenically, it is, in all probabiUty, ancestral to Palseoloxodon, and hence to the Loxodontine elephants as

a whole. Thus, the form appears to be stronger to the side of Palseoloxodon than to the side of
affinity of this

Archidiscodon. A generic and a specific name have already been proposed to receive it [Footnote 'Matsumoto, on :

Leiih-Adamsia Siivalikiensis, Jap. Journ. Geol. & Geogr., Vol. V., No. 4, 1926-1927.']. Palseoloxodon, of which the
most archetypal known form appears to be represented by Lox. (Pal.) Tokunagai, might have arisen from such an
earlier form as represented by the aforementioned species through the partial perfection of the loxodonty of the

grinders. No doubt, the Loxodonts as a whole were originally a narrow-toothed type, with a low ridge-formula.
The acquisition of loxodonty might mechanically be correlated with the combined facts of their being narrow-

toothed and long retaining a low ridge-formula. Palseoloxodon appears to have originated in the region extending
from India to Japan, and then to have been distributed over practically the entire southern Palsearctic."

[Leiih-Adamsia siwalikiensis Mats, is a synonym of Archidiskodon planifrons Falc. and Caut. (see Chap. XVI,
p. 959, of the present Memoir).]

"Pal^oloxodon and Loxodonta, s.s."

"The writer has never seen a molar of Palseoloxodon, which is more like those of Loxodonta, s.s., than the type-
specimen of Lox. (Pal.) Tokunagai, in being very narrow-crowned and in the very markedly lozenge-shaped disks
 :

1292 OSBORN: THE PROBOSCIDEA

of the well-worn ridges. If we suppose that the cheek-teeth of just this type had acquired hypsodonty of a very
high degree, without increasing their width and the number of ridges, then we may obtain cheek-teeth of a type
just characteristic of Luxodonta, s.s. The modern Loxodonts might have originated in such a way, probably in
Africa."

"Specific Groups in Pal^oloxodon"

"The most archetypal group in Palxoloxodon corresponds evidently to that represented by the phyla of Lox.
{Pal.) prisca in Europe and of Lox. (Pal.) Tokunagai in Japan, showing the distinctive characteristics already

stated. The question arises whether the group of pygmy Loxodonts of the Mediterranean islands and coasts,
typified by Lox. (Pal.) melUensis (Falconer), is a close ally of the group just mentioned. The answer must be
negative. Though the cheek-teeth of the group in question have a low ridge-formula, the disks of their ridges,
as well as their general shape, appear to display an unmistakable similarity with those of the later phases of the
phylum The low ridge-formula in this group may [be] due to degeneration as a result of
of Lox. (Pal.) antiqua.

having been dwarfed. Phylogenically, this group by itself is far from being a natural one. It can be considered
natural only when it is taken as a group subordinate to the phylum of Lox. {Pal.) antiqua. By far the greatest part
of Palxoloxodon is occupied by the group represented by the phyla of Lox. {Pal.) namadica in Southern to Eastern

Asia and of Lox. {Pal.) antiqua in Europe. As indicated by the evolutionary tendencies observed in the mutations
of the phylum of Lox. {Pal.) namadica in Japan, Lox. {Pal.) Tokunagai is probably ancestral to this phylum.
Again, it appears probable, that the phylum of Lox. {Pal), antiqua, might also have had an almost similar form for
its ancestor. It does not appear probable, however, that Lox. {Pal.) prisca was actually ancestral to that phylum,
as a younger aspect appears to be present in the molars of this species in having the opposite loxodont sinus of
two neighbouring ridges well-spaced."

OSBORN'S SUMMARY (1930) OF THE OBSERVATIONS OF MAKIYAMA (1924)'

AND OF MATSUMOTO (1924-1929)

From these descriptions and plates we conclude: (1) That Palxoloxodon is the only genus of true elephants
that penetrated Japan ; (2) that the primitive species P. tokunagai may have entered as early as Upper Pliocene
time, and (3) that in ascending geologic levels there occur more progressive species, either indigenous or migrants

from the P. namadicus stock of India. (4) In the present historic, geologic, and systematic revision, therefore, we
shall present in each case Matsumoto's opinions of 1924, and, in conclusion, give his opinions of 1929 under
each species.

Osborn, 1930: Osborn does not accept any of the phylogenetic or geographic theories expressed by Matsu-
moto above; he regards Palxoloxodon as an entirely independent phylum originally derived from Africa and
subsequently migrating through India to the Far East. He condenses from the invaluable observations of Maki-
yama and of Matsumoto the following synopsis of their observations upon the ascending geologic successions and
specific characteristics of the Japanese loxodonts

Palxoloxodon tokunagai Matsumoto, 1924, possibly of Upper Pliocene age, is regarded as the most archetypal
group of Palxoloxodon in Japan, probably as ancestral to Palxoloxodon namadicus of southern and eastern Asia
and to P. [Hesperoloxodon] antiquus of Euroi^e (1929.2, pp. 10, 1 1). Estimated ridge formula of 3 ,,., 3 ^j. Ridge M
frequency 5 in 100 mm. Length 295e nun., breadth 80 mm. ; relatively low crowned. (No figure.)

MFor the results of Makiyama'.s recent studies, see his article "Japonic Proboscidea," Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, Vol. XIV, No. 1, Art.
1, May, 1938.— Editor.)
 9

THE LOXODONTIN^: PAL^OLOXODON OF JAPAN 1293

Palaeoloxodon tokunagai mut. junior Matsumoto, 1929, a mutation of Loxodonta {Palxoloxodon) tokunagai.
Very primitive ; either of Upper Pliocene or Lower Pleistocene age. Similar to Loxodonta {Pal3eoloxodon) prisca
of Falconer. Differs from P. tokunagai which has molars of larger size, with slightly higher ridge formula and
slightly more perfect lozenge-shaped discs of the ridges. Low crowned. Enamel thick. (Matsumoto, 1929.2,
p. 10, also PL VII, fig. 1, type r.M2, our Fig. 1157.)

Palxoloxodon protomammonteus Matsumoto, 1924, 1926, regarded as of Upper PUocene (?), Basal Calabrian
age (Fig. 1154). Lower molars small and narrow crowned. Estimated ridge-plate formula (M 3 ^nTTi) similar
to that of the Lower Pleistocene Hesperoloxodon antiquus.

Palseoloxodon protomammonteus proximus Matsumoto, 1926. Upper Pliocene(?). Lower Calabrian age.

Molars rather large and moderately wide. Type fragment, I.M3, with 9+ ridge-plates. Ridge frequency 5 in

100 mm. (Fig. 1155.)

Palseoloxodon namadicus naumanni Makiyama, 1924, characteristic of the Lower Pleistocene, very widely
distributed. A narrow-toothed variety. Ridge frequency 5-6 in 100 mm. in M,, 6 in 100 mm. in M'. Relatively
broad crowned. Ridge formula of M 3 ^4f^' [K, Fig. 1152]. Length M^ 303 mm., breadth 76 mm., max. height
216 mm. ; length M3 270 mm., breadth 72 mm.
Palseoloxodon namadicus yabei Matsumoto, 1929, characteristic of the Middle Pleistocene, very abundant.
Estimated ridge formula of M3 s^n^', length 255 mm., width 74 mm.; ridge frequency 6.5-7 in 100 mm. (Fig.

1156.)

Palseoloxodon namadicus namadi Makiyama, 1924, probably of Middle Pleistocene age. The largest and most
numerous group of the Japanese loxodonts, representing a broad-toothed variety. Ridge frequency 5 in 100 mm.
in M3. The type upper molar, r.M' (Fig. 1153), exhibits +12+ ridge-plates indicating a larger total, as observed
by Makiyama (1924, p. 263) "There is a loss of several posterior ridges, leaving fourteen in a space of 211 mm.
:

The crown is very broad being 77 mm. in front, 90 mm. at the middle and 65 mm. posteriorly, measured on the
second, fifth and fourteenth ridges respectively."

According to the above summary, the genus Palseoloxodon from Japan probably appeared in Upper Pliocene
time as a narrow-toothed, coarse-enameled species, and ascended from Lower into Middle Pleistocene time,
terminating in the species Palseoloxodon namadicus yabei, characteristic of the Middle Pleistocene and so closely
resembling P, namadicus that it was first described by Matsumoto as Palseoloxodon namadicus typicus. This
ascending order of phylogenetic succession, with broadening molar crowns and ridge-plates multiplying from
M3 v^^tttm estimated (tokunagai) to M3 - estimated {naumanni) differs widely from that shown in figure 790,
which represents Matsumoto's earher observations of the year 1924.

The Japanese species of Palseoloxodon may all be derived from the same Upper Pliocene African stock, with
narrow and low ridge formula, progressive in breadth and in ridge formula as follows:

(Upper Pliocene) M 3 t^ to (Lower Pleistocene) M 3 y^t to (Middle Pleistocene) M3 1

1 T

TWO JAPANESE SUBSPECIES DESCRIBED BY MAKIYAMA (1924)

Makiyama in a paper entitled "Notes on a Fossil Elephant from Sahamma, Totomi" (1924.2, pp. 261, 262,
264) expresses the opinion that there are at least two varieties of Japanese fossil elephants, namely, the narrow
toothed (e.g., Palseoloxodon naumanni) and the broad toothed (e.g., P. namadi), which hitherto have been called
 Tusk
yl2 Nat.

^ A/aturaZ sije
ELEPHAS NAMADICUS NAUMANNI Type
Kyoto /mp. Mus.

3 4 5 6 7
SIVALIKIA ANTIQUA of Taulrach

Genotypic Species (A-C) of PaL/Goloxodon Matsumoto; (D) Hesperoloxodon antiquus oermanictjs of Taubach
All figures one-fourth natural size, same scale as figure 1073 excepting tusk (A) one-twelfth natural size

Fig. 1152. Diagrammatic outline sketch of the ixjpi of KUphas namadicus

nautiianni Makiyama, 1924 = Palnoloxodon namndicus naumanni], see
[

Pis. XII, XIII, XIV, XV, and XVI, fig. 1, in comparison with \llespemhxMtona>diquus germanicux], after Pohlig, 1888, PI. iii, fig. 7. PI. iv, fig. 3.
A^itp/iasaHa'^iiu.s

All figures one-fourth natural size, with the exception of the incisive tusk which is one-twelfth natural size.

A, Incisive tusk, length 1930 mm. (PI. xii).

Bl, Third right superior molar, cxhiljiting II worn and .'J unworn ridge-plates, total 19. Side view. (PI. xiii, fig. 1. Length 303 mm., breadth 7€, mm.,
max. height 21() mm.)
B2, Crown view cf same superior molar (PI. xiii, fig. 2).
CI, Type infc^rior mandible, with M 3 in silu exhibiting 14 worn and 3 unworn ridge-|)lates, total 17. Superior view. Length of right ramus 40.') mm.,
of left ramus 413 mm.; length of l.M.i, 270 nun.; max. breadth of .same 72 mm. at fifth ridge (PI. xv).

C2, Same mandible, left lateral view (PI. xvi, fig. 1).

D, Elephas [ = Hesperoloxodon]
antiquus [germanicus] from Taubach, after Pohlig, 1888, PI. iii, fig. 7, left Mi, crown view, and PI. iv, fig. 3, left Ma, crown
view. Compare figure 1075, type second left inferior molar, l.Mj, exhibiting 12-13 ridge-plates, of 'Elephas antiquus' I''alc.

Ob.serve that the ty\>c of 'Elephas namndicus naumanni' Makiyama is decidedly inferior in size to the /'«ia'()^»jo(/oK ?mm«(/ic».s- of the Ncrbudda, figure
1073, but that the grinders exceed in relative breadth tho.se of Hesperoloxodon antiquus germanicus i^i 'V-M\\rM'\\ (D, l.M;, l.Mj), as figured by Pohlig.

1294
 —

THE LOXODONTIN^: PAL^OLOXODON OF JAPAN 1295

Elephas namadicus by many authors. Osborn, on the contrary, interprets the 'narrow toothed' as more primitive
and geologically ancient, the 'broad toothed' as more progressive and geologically younger ;
progressive broadening

of the grinding teeth and multiplying of the ridge-plates distinguish each line of ascent.

Narrow toothed. — {Op. "The second variety, to which the elephant of Sahamma belongs
cit., pp. 263, 264:
[i.e., Palxoloxodon namadicus naumanni\, is diagnosed by its narrower [76-79 mm.] bandy crown and lozenge-

shaped discs. ... I give here, a new subspecific name Elephas namadicus Naumanni to the second variety, for
which type the lower teeth from Sahamma should be chosen. The second variety is apparently referable to. . .

E. antiquus in many respects. Therefore, it is not unreasonable that Brauns [Footnote: 'Ueber Japanische Dilu-
viale Sjiugetierer, Zeitsch. Deut. Geol. Ges., 1883, pp. 35-42.'] had supposed the possibility of direct migration of
E. antiquus from Europe to Japan through Central Asia in the early Pleistocene age. He also pointed out some
differences between his E. antiquus in Japan and the types of E. namadicus in India."

Broad toothed. — (Op. cit., pp. 263, 264) : "The first variety [Palseoloxodon namadicus namadi] is diagnosed
by the broad [90 mm.] elliptical crown and less-crowded bandy discs. . . . The first variety referred to as typical
E. namadicus should be called more strictly Elephas namadicus namadi. '"^

While Matsumoto (1929.1, p. 1) gives the original reference of these two subspecies as "Chikyu —The Globe,
Vol. I, 1924, p. 381, PI. VIII," this publication is not available to the present author and he is therefore citing from
Makiyama's supplementary descriptions in English (Makiyama, 1924.2) mentioned above.

Palseoloxodon namadicus naumanni Makiyama, 1924' 1921. Type Figure. — Makiyama, op. cit., Pis. xii (incisor),
Figures 1152, 1189 XIII (third right superior molar), xiv (third left xv
superior molar),
From Sahamma, Totomi Province, Japan. Recorded from the Lower (mandible), xvi, fig. 1 (left lateral aspect ofsame mandible).
Pleistocene by both Makiyama and Matsumoto. Probably Middle Pleisto- Specific Characters (cf. Makiyama, pp. 260-264).
cene. Genotypic species of Palssoloxodon Matsumoto, September 20, 1924. Superior incisor 1930 mm.upcurved (PI. xii).
in length, strongly
This subspecific name was applied by Professor Makiyama of Third molar (PL xiv), length 286 mm., breadth 79
left .superior
the Kyoto Imperial University to an e.xcellent medium-toothed mm. at fourth ridge-plate, maximum ridge-plate height 217 mm.,
type (Fig. 1152) rivahngin thedimensionsof the teeth the 'Elephas K'-16-/^ ridge-plates in 286 mm.; third right superior molar (PI.
antiquus' of Weimar (Fig. 1088B, M'), in which the referred formula xiii), length 303 mm., breadth 76 mm., maximum ridge-plate height
is M3 -Y^, the enamel thick and strongly crimped, the dentinal 216 mm., total ridge-plates 19, laminar frequency 17 in 303 mm.,
discs e.xpanding mesially and coming into contact with extreme worn anterior plates exhibiting six lens-shaped discs with pro-
wear; the superior tusks strongly curved, the frontocranial nounced plications. Mandible (Pis. xv, xvi, fig. 1) with both third
structure unknown — on the whole, resembling the large Upper inferior molars in situ; symphyseal rostrum abrupt, subvertical;
Pleistocene 'E. antiquus [germanicus]' of —
Weimar distinguished I.M3, length 270 mm., breadth 72 mm., laminar frequency 17 ridge-
both from the broad-toothed variety (maximum breadth 90 mm.) plates in a space of 270 mm.; 14 anterior ridge-plates well worn
and from the extremely narrow-crowned variety. "showing a lozenge-shaped complete figure of enamel with minute
Elephas namadicus Naumanni Makiyama, 1924. "Notes on but well-defined central angulations in touch with each other as
a Fossil Elephant from Sahamma, Totomi." Mem. College Sci. obtained in E. africanus, and regularly crimped, comparatively
Kyoto Imp. Univ., Ser. B, Vol. I, No. 2, June 30, 1924, pp. 255- thick enamel layers."
264. —
Type. Cranium broken to pieces, leaving complete Comparisons. —
In dimensions the upper right last molar,
mandible with third inferior molars, also third superior molars, and r.M^, is very like that of Elephas antiquus [germanicus] from
inferior incisor. (Op. cit., p. 264): "I give here, a new subspecific Weimar cited by Soergel (1913, Taf. viii). It also resembles the
name Elephas namadicus Naumanni to the second variety, for thick-plated variety of E. antiquus (cf. Leith Adams, 1877, Pt. I,

which type the lower teeth from Sahamma should be chosen." p. 31). An M- described but not figured, length 204 mm., breadth
—
Horizon and Locality. Excavated at Sahamma, about 12 72 mm., height 140 mm., differs in dimensions from those of E.
kilometers northea-st of Hamamatsu, Totomi Province, Japan, in antiquus, corresponding more nearly to those of E. meridionalis

'[Original description in Japanese: "Chikyii—The Globe," Vol. I, 1924, p. 381, PI. viii (fide Matsumoto, 1929.1, p. 1). See also Chapter XXI of the
present Memoir, p. 1408, under "1924 Elephas nainadicus naumanni," and p. 1413, under "1929 Elephas {Palxoloxodon) namadicus setoensis," for Doctor
Makiyama's recent (1938) conclusions.— Editor.]
 |

1296 OSBORN: THE PROBOSCIDEA

(Soergel, 1913, Taf, vi); the discs are feebly rhombic and never mm. in front, 90 mm. at the middle and 65 mm. posteriorly, meas-

touch at the central part. The mandible (PI. xvi, fig. 1) in many ured on the second, fifth and fourteenth ridges resix'ctively. The
respects resembles that of E. antiquus. grinding surface displays a characteristic oval outline and well-
Final Diagnosis (Matsumoto, 1929.1, h. 2).— "The race spaced bandy discs with minute median angulations. The enamel
now under consideration can be diagnosed as follows Cheek-teeth :
layer is less well-crimped and thinnei- than that of the Sahamma

moderate in size, narrow-crowned. Well worn surface of the

last upper molar shaped like an elongated ellipsoid, and
that of the last lower molar band-like, with the outer border
almost linear or concave. Frequency of ridges low, number-
ing about 6 or less in a length of 100 mm. in the last upper
molar and 5-6 or less in the lower. Mammillae of the sum-

mits of ridges stout and rather few, remaining proximally

widely spaced to some extent. Disks of much or moder-
ately worn ridges lozenge-shaped, without any marked
differentiation of mesial portion and lateral arms. Even the
lateral portions of disks are rather thick antero-posteriorly,
the opposite loxodont sinus of the two neighbouring disks in
the much or moderately worn portion of crown being in con-
tact with or very closely set to each other. Layer of enamel
Type Molar op Pal^oloxodon namadicus namadi
rather thick; its plication being rather coarse, irregular and
Fig. 1153. Type r.M' of Elephas Tiamadicus namadi Makiyama, 1924, exhib-
rather strong. . . . Age: The present race appears, in all like-
iting worn ridge-plates, one anterior ridge-plate missing, posterior ridge-plates
12
lihood, to be characteristic of the Lower Pleistocene, ranging not shown in pliotograpli. Breadtli 90 mm. After Makiyama, 1924.2, PI. xvi, fig.
in occurrence very probably from its base, as at Okine, up to 2. One-half natural size.

its very close, as at Tabata."

form. . . . The first variety referred to as typical E. 7iamadicus,

should be called more strictly Elephas namadicus namadi."
Final Diagnosis (Matsumoto, 1929.1, p. 3).
— "The present
PalBBOloxodon namadicus namadi Makiyama, 1924'
race is the largest group of the Japanese Loxodonts and represents
Figures 1I.')3, 1189
the broad-toothed variety of the present species in Japan. . . .

DrcdKfii off tlic island of ShiVlo, .Sanuki Province, .lapan. Probably This race can be diagnosed as follows. Cheek-teeth large and
Middle Pleistocene (Jide Matsumoto, 1929.1, p. 4).
last lower molar shaped
rather broad. Well worn surface of the
Elephas namadicus namadi Makiyama, 1924. "Notes on a like an elongated ellipsoid. Frequency of ridges low, numbering
Fossil Elephant from Sahamma, Totomi." Mem. College Sci., about 5 or less in a length of 100 mm. in the last lower molar.
Kyoto Imp. Univ., Ser. B, Vol. I, No. 2, June 30, 1924, p. 264. Mammillae summits of ridges stout and few. Disks of only
of the
—
Type. Last upper molar, right .side, r.M'. Horizon and the very strongly worn ridges lozenge-shaped as a whole; those of
—
Locality. Dredged off the i.sland of Shodo, Sanuki Province, the moderately worn ridges consisting of a lozenge-shaped mesial
Japan. Probably of Middle Pleistocene age {fide Matsumoto, portion and of nearly parallel-sided lateral arms. Interspaces
1929.1, p. 4). Type Figure.— Op. til., PI. xvi, fig. 2. between lateral arms of successive disks broad antero-jiosteriorly.
Type Description.— (Op. cil., 1924.2, pp. 263, 264): "An Opposite loxodont sinus of the two neighbouring disks of mod-
excellent examjjle of the first variety is furnished by the last upper erately worn ridges widely separated from each other. Layer of
true molar. . . . There is a loss of several posterior ridges, leaving enamel rather thick its plication being coarse, irregular and feebly
;

fourteen in a space of 211 mm. This crown is very broad being 77 displayed."

'[Original description in Japanese: "Chikya—The Globe," Vol. 1, 1924, p. 381, PI. viii {fideMatsumoto, 1929.1, p. 1). See al.so Chapter XXI of the
present Memoir, p. 1408, under "1924 Elephas namadicus naumanni," and p. 1413 under "1929 Elephas (Palseoloxodon) namadicus seloensis," for Doctor Maki-
yama's recent (1938) conclusions. Editor. —
 :

THE LOXODONTIN^: PALiEOLOXODON OF JAPAN 1297

FIVE'JAPANESE LOXODONTINES DESCRIBED BY MATSUMOTO (1924-1929)

(Continued from Chap. XIV, pp. 908-908 of the present Memoir)

During the period 1924-1929, Matsumoto described two species and three subspecies of the true elephants
of Japan under the following names: Euelephas protomammonteus, 1924, Loxodonta (Palxoloxodon) tokimagai,
1924, Parelephas protomammonteus proximus, 1926, Loxodonta (Palxoloxodon) namadicus yabei, 1929, and Loxo-
donta (Palxoloxodon) tokunagai junior, mut., 1929. Matsumoto's Memoir of 1924, "Preliminary Note on Fossil
Elephants in Japan" (September 20), which contains the original descriptions of his subgenus Palxoloxodon, and
of his species Euelephas protomammonteus and Loxodonta {Palxoloxodon) tokunagai, was published in the Japanese
language ; consequently the present author in the systematic description of the species and subspecies has cited
freely from Matsumoto's later Memoirs of 1926 and 1929 (which appeared in the English language) as given
below.

Osborn, 1929: Since the names Euelephas {Parelephas) protomammonteus (1924, 1926) and Parelephas
protomammonteus proximus (1926) were assigned by Matsumoto under the impression that the genus Parelephas
Osborn was represented in Japan, and since they were not included in the author's subsequent memoirs of 1929,
our conclusion is that both these specimens belong to Palxoloxodon. See also Elephas indicus Buski Mats.,
1927 [
= Palxoloxodon buski] below, p. 1333.

PalsBoloxodon protomammonteus Matsumoto, 1924, 1926'

Figure 11.54

Nagahama, Town of Miiiato, Kimitsu District, Province of Kazusa,

Japan. Upper Pliocene (?), Lower Pleistocene.

Euelephas protomammonteus Matsumoto, 1924. "PreHminary

Note on Fossil Elephants in Japan." Journ. Geol. Soc, Tokyo,
1924, Vol. XXXI, p. 262 (in Japanese language). Parelephas
protomammonteus (Matsumoto) typicus Matsumoto, 1926. "On
the Archetypal Mammoths from the Province of Kazusa." Sci.
Kept. Tohoku Imp. Univ., (2), Geology, Vol. X, No. 2, pp. 43-
50. —
Type. A third inferior molar of the left side, I.M3,
belonging to Mr. Natsume of the Town of Minato. Horizon
—
AND Locality. Probably from the very base of the Narita series.
Nagahama, Town of Minato, Kimitsu District, Province of
Kazusa, Jai)an. Type Figure. Matsumoto, 1926.2, PI. —
xviii, figs. 1, 2. —
Paratype. A third inferior molar of the right
side, r.Ms. From the same locality as the type figured on PI. xix, ;

figs. 1-3.
In view of the fact that the original type description was ••fs

pulilished in the Japanese language, we may cite from Matsumoto's

Memoir of 1926, p. 50 (published in the English language) Type ok Pal^koloxodon protomammonteus
"Stage I. Parel. protomammonteus (typicus). Basal Calabrian. Fig. llo4. Type figure of Euelephas (Parelephas) yrohimniiiinoiUeus
(typicus), a. third inferior niohir of t crown and internal aspects,
lie left side, I.M3,
Molars small and especially narrow-crowned. Inner and
one-third natural size. From Nagahama, Town of Minato, Kimitsu District,
outer sides of ridges not very convex. Bases of ridges prominent,
Province of Kazusa, Japan. After Matsumoto, 1926.2, PI. xvni, figs. 1 and 2,
and valleys acutely pointed proximally. Basal cingula, and some- one-half natural size. Observe -|-16}^ ridge-plates.
times also accessory columns, well developed." (Matsumoto, op. "The holotype, M3, lacking its most anterior
cit., p. 44):
Osborn, 1928: Matsumoto's type figure and description appear and a posterior talon. Its original ridge formula
part, consists of sixteen ridges

to establish lieyond question a strong resemblance to Palxoloxodon might i)robably be about X17X. Its length, as preserved, is 275 mm.; and
its original length might probably be .some 300 mm. The maximal width is
[H esperoloxodon] antiquus typical form of the Lower Pleistocene of
75 mm. at the seventh ridg(\ as estimated above, and the height of the crown
England; the ridge-plate formula is practically the same, i.e.,
is 134 mm. at the eleventh ridge, as estimated above, which had just com-
M 3 TeW-ll- Matsumoto observes (op. cit., p. 44): "This molar menced to wear. The frequency of ridges in a lengUi (if 100 mm. is 5)^-6 at
is decidedly of a narrow-crowned type, much resembling in general the sides and about 7 at the grinding surface."

'[See Chapter XXI of the present Memoir, p. 1408, note under "1924 Elephas namadicus naumanni Makiyama." — Editor.]
 :

1298 OSBORN: THE PROBOSCIDEA

shape tlic corresponding ones of the pliyla of Loxodonta (Palaso- thick, measuring 2.5-3 mm. across; and its plication is very coarse
loxo(lo)i) namodira and antiqua, in contrast to Parel. trogontherii and indistinct. There is a distinct, linear, transverse streak, or
and higher mammoths. Consequently, the sides of the ridges in e\-en fissure, in each area of cement between two disks at the

fore and aft views are only weakly convex, instead of being so grinding surface."
strongly so as in the two last-mentioned species, and are not so
strongly convergent toward the base as in the same. The ridges,
except the very posterior ones, are curved forward in lateral views, Palseoloxodon protomammonteus proximus
quite as those of the lower molars of Parel. trogontherii." Matsumoto, 1926
Osborn agrees in the close resemblance of Palieoloxodon proto- Figure 11.5.5
mammonteus to Hesperoloxodon antiquus, but does not agree that it
Lower part of the Narita Series. Kokubo, Onuki-niura, Kimitsu
Isone,
is related to the Parelephas trogontherii phylum.
District, Province of Kazusa, Japan. Upper Pliocene (?).

Referring to Matsumoto's definition of Eiielephas (Parelephas)

Palseoloxodon tokunagai Matsumoto, 1929
protomammonteus (typicus), that species is regarded as basal
Soyama, Gokayama, Hira-mura, Higaslii-Tonami District, Province of
Calabrian, whereas the following species belongs on a higher level,
EtchO, Japan. Recorded as of Upper Pliocene or Lower Pleistocene age.
namely, lower Calabrian (Matsumoto, 1926.2, j). 50)
Loxodonta (Palseoloxodon) tokunagai Matsumoto, 1924. "Pre- "Stage II. Parel. protomammonteus proximus. Lower
liminarj' Note on Fossil Elephants in Japan." Journ. Geol. Soc, Calabrian."
Tokyo, Vol. XXXI, September 20, 1924, p. 267 (Japanese lan- "Molars rather large and moderately wide. Inner and outer
guage). Supplementary Description (1929.2): "On Loxodonta sides of ridges not very convex. Bases of ridges more or less jjronii-
(Palseoloxodon) tokunagai Matsumoto, with Remarks on the De- nent, and valleys acutely pointed proximally. Basal cingula appar-
scent of Loxodontine Elephants." Sci. Rept. Tohoku Imp. Univ., ently not well developed."
(2), Geology. Vol. XIII, No. 1, pp. 7-10 (English language). Parelephas protomammonteus proximus, mut. nov. jMatsumoto,
Type. — A third inferior molar of the right side, r.Ms. Originally 1926. "On the Archetypal Mammoths from the Province of
belonging to the Imix-rial Ueno, numbered 2208 now
Museum of — Kazu.sa." Sci. Reiit. Tohoku Imp. Univ., (2), Geology, Vol. X,
in the Museum of Education at Ochanomizu, Tokyo. Hori- No. 2, pp. 48-50. —
Type. Fragment of a third inferior
—
zon AND LotALiTY. Soyania, Ciokayama, Hira-mura, Higashi- molar of the left side, I.M3, belonging to Mr. Kato of Kokubo.
Tonami District, Province of Etchu, Japan. Of Upper Pliocene —
Horizon and Locality. "This specimen appears to have been
or Lower Pleistocene age. Type Figure. — The present derived, very probably, from a certain lower part of the Narita
author has been unable to locate a type figure. Series." Isone, Kokubo, Onuki-mura, Kimitsu District, Province
—
Type Description. (Cited from Matsumoto's Memoir of of Kazusa, Japan. Type Figure. Matsumoto, op. cit.,—
1929.2, p. 8) "Description of the type-specimen This last molar
: : 1926, PI. XXIV, figs. 1,2.
consists, as preserved, of eleven ridges and the single-columned Type Description. —
In addition to Matsumoto's type
posterior talon. Judging from the general shape of the molar, two figureand type description quoted below in the legend, Matsumoto
nior(> ridges in all likelihood, might have originally been present compares this stage with his Euelephas (Parelephas) protomam-
anterior to the most anterior ridge as preserved. Thus, the ridge- monteus as follows: The type molar of Parelephas protomam-
formula appears to be nearly Xl3X. Its length as preserved
measures 258 mm.; its original length can be estimated as about
29o nun. Its full maximal width is 80 mm. at the ninth ridge as
preserved, which is quite abnormal in having a superfluous column
standing out on its inner side. Not including this sui)erfluous
column, however, the true maximal width of the molar nieasun-s
74 mm. at the sixth to eighth ridges as preserved."
"The frequency of ridges in a length of 100 mm. is about 5 on
hotli the inner and outer sides, a-s well as at the grinding surface.
From an inner or outer view, the ridges are seen gradually narrowed
though the proximal ends of the valleys, with the ex('eption
distally,
of the most posterior ones, are of a rather obtuse typ(>. The
curvature of the ridges varies from very to rather slight for a lower TypE ok PAL.BOLOXODON PROTOMAMMONTEUS PUOXIMUS
molar."
1 1.55. Type figure of Parelepltas prolomammnnleus proximus, mut.
Fig.
"The .smallness of the transverse width of tlic more anterior nov. Matsumoto, 1920.2, PI. xxiv, fig. 1, one-half natural .size, from Isone,
ridges is quite remarkable; and the disks of the well worn ridges Kokubo, Onuki-mura, Kimit.su Di.strict, provinc^^ of Kazusa, .lapan. Frag-
are very markedly lozenge-shajK-d, — to such a degree that they ment of a l(^ft inferior molar, I.M3, crown view.
remind us of those in Loxodonta, s.s. The tips of I he anterior (Matsumoto, I92().2, p. 48): "Tliis specimen, n^prcsc^nting thi' anterior
part of Ms, consi.sts of nine ridges and nK^astires 168 nun. iti length as preserved.
loxoilont sinus of one ridge and the posterior loxodoni sinus of he I

Its maximal width is 88 mm. at the s<^vcnth ridge as preservcnl, and its height
one immediately preceding in the well worn portion of tlu; molar is 128 mm. at the last ridge as preserved, which is slightly worn. The frequency
are in close contact with each other. The layer of enamel is very
. . . of ridges in 100 ram. is .5."
 THE LOXODONTINiE: PAL^OLOXODON OF JAPAN 1299

monteun proximus is distinguished from tlio type molar of Parcle- ellipsoid. Frequency of ridges rather high, that of the last lower
phas prolomamnionteus by its greater width and by the slightly less molar of the type-specimen counting 6.5-7 in a length of 100 mm.
prominent bases of its ridges ; as a whole P. pro.rim us is a mutation Mammillae of the summits of ridges slender, tending to be numer-
intermediate between P. prolomammonteus and P. trogontherii. ous and cloisely set; the clefts between the mammillae extending
This phylogcnetie opinion is expressed by Matsumoto on page 50 proximally to a considerable height. Disks of the much worn
(o?). (/<., 1926.2) as follows: ridges lozenge-shaped; those of the moderately worn ridges con-
sisting of an expanded mesial portion and of
Stage III. Parel. trogontherii. Calabrian to Tyrrhenian. nearly parallel-sided lateral arms, which are
characteristically thin antero-posteriorly.
Stage II. Parel. protomammonteus proxinius. Lower
Calabrian.
Opjjosite loxodont sinus of the two neigh-
bouring disks in the moderately worn portion
Stage I. Parel. protomammonteus {typicus). Basal of the crown moderately or very widely sep-
Calabrian. arated from each other. Layer of enamel
thin; its and
plication being fine, irregular
OsBOHN, 1930. -In the above deseription Matsvmioto ex- strong. The present race is by far the
. . .

presses the opinion, not shared by Osborn, that these specimens most common Proboscidca in Japan."
are referable to the Parelephas trogontherii phylum. Osborn re-
gards them rather as loxodonts referable to the Palxoloxodon
phylum and quite distinct from any of the phyla of the Mam-
montinse. Both in the ridge formulae, and in the proportions and
the structure of the ridge-plates these types appear to agree quite
closely with Hcfperoloxodon aiiiiquu.'i and Pols-oloxodon namadicus
and to be very distinct from the Parelephas trogontherii phylum.

Palaoloxodon namadicus yabei Matsumoto 1929'

Figure 11.56

Inland Sea, .Japan. Recorded as of Middle Pleistoeenc age.

Loxodonta namadica {Yabei) Matsumoto,

(Palseoloxodon)
1929. "On Loxodonta (Palseoloxodon) namadica (Falconer and
Cautley) in Japan." Sci. Rept. Tohoku Imp. ITniv., (2), Geology,
Vol. XIII, No. 1, pp. 4 and 5. Type. "Right ramus of —
mandible, bearing la.st molar in situ belonging to the Second High
;

School." HouizoN AND Locality. Inland Sea. "The —

present race appears to be characteristic of tiie Middle Pleisto-
cene." Type Figure.— 0/j. cil., 1929.1, Pi. iii, fig. 2, and
PI. IV.

Type Description.— (Op. cit., 1929.1, p. 4): "The molar of

the type-specimen is nearly complete, save that the most anterior
portion of the crown, corresponding to the most anterior root, is Type op Pai„eoloxodon namadicus yabei
liroken away. The mentioned very prol;)ably consisted
lost portion
Fig. ll.')G. Typo
right, ramu.s of mandible, containing r.Mj in situ, of
of the anterior talon and first two ridges. Since this molar, as Loxodonta (Palseoloxodon) namadica (Yabei) Mat.'sumoto, 1929. Ridgc-platcs
preserved, contains fifteen complete ridges and the posterior talon, of r.Ms (J.J 7 44, lengtli 2.")r) mm., breadth 74 mm., ridge frc(|uen('y ()..5 7

it is almost probable, that its original ridge-formula corresponds to in 100 mm. After Matsumoto, 1929.1, PI. iii, fig. 2, and PI. iv. Jaw one-
fourth natural .size; molar Iwo-fiftlis natural size.
X17X. Its total length, including the broken portion, above the
margin of the jaw, is about 255 mm. Its maximal width is 74 mm.
at the eighth ridge. Its frequency of ridges in 100 mm. counts
6.5-7." Palseoloxodon (Archidiskodon ?) tokunagai mut. junior
Referred Specimen. — "Penultimate upper molar of right Matsumoto, 1929
side; belonging to our Institute of Geology and Palaeontology." Figure 1157
The specimen
locality of the referred is Sorachi, Uryu District, Japan, precise locality unknown. Probably upper ])art of the Upper
Province of Ishikari, Hokkaido. Pliocene or Tjower Pleistocene.

Matsumoto (op. cit., 1929, p. 5) : "This i-ace can be diagnos- The thick enamel, the widely spread ridge-plales, and I lie

ed as follows. C'heek-teeth moderate in narrow-crowned. Well

.size, alleged 'Upper Pliocene' or 'Lower Pleistocene'(?) age suggest
worn surface of the last lower molar shaped like an elongated a possible reference to Archidi.'ilcodoii (cf. pp. 957-959, ^4. pkiiii-

'[See Chapter XXI of the present Memoir, p. 1408, note under "1924 Elephas namadicus nautnanni Makiyama." — Editor.]
 —

1300 OSBORN: THE PROBOSCIDEA

The side view of the type (Matsumoto, the specimen to the Lower Pleistocene mutation Naumanni
frons of the Siwaliks).
1929.2, PI. VII, fig. 2) displays a relatively low-crowned, widely Makiyama — of Lox. (Pal.) namadica in Japan, as well as from the
degree of fo.ssilisation, the. specimen can probably be referred to
ridge-plated molar.
. .

Lox. {Pal.) Tokunagai junior, mut. nov. Matsumoto, 1929. either an upper part of the Upper Pliocene or to the Lower Pleisto-
"On Loxodonta {Palseoloxodoit) tokunagai Matsumoto, with Re- cene." Type Figure.—Op. cit., 1929.2, PI. vii, figs. 1, 2.
marks on the Descent of Loxodontine Elephants." Sci. Kept. T6- Type Description.— (Op. cit., 1929.2, p. 10): In a comparison
Geology, Vol. XIII, No. p. 10. Type.— of the type of the present mutation, namely, Loxodonta {Pal9eolo.ro-
hoku Imp. Univ., (2), 1,

Second inferior molar of the right side, r.M2, belonging to the don) toku7iagai junior, with the type of Loxodonta {Palseoloxodon)
Geological lastitute of the Kyoto Imperial University. Horizon tokunagai, Matsumoto notes the following differences which can be

AND Locality.—Japan, precise locality unknown. ". judging . . observed between them: (1) The more anterior portion of the

from the less but still archetypal feature, and from a comparison of crown of the mutation JM/(jor is not so narrow as that of tokunagai,
on the whole, it appears to be less nar-
row crowned; (2) the disks of ridges
are less lozenge shaped; (3) the oppo-
site loxodont sinus of the two neigh-
bouring disks of the well worn ridges
are closely set in the junior mutation,
whereas they are in contact with each
other in tokunagai; (4) the plication of
enamel is stronger, and (5) the layer of
enamel appears to be thinner in the
junior mutation than in tokunagai,
though this difference in the absolute
thickness should be underestimated
Primitive Type Molar ok Pal.boloxodon (Archidiskodon?) tokunagai mut. junior
in the present case. "The writer pro-
Fig. 11.57. Second inferior molar of the right side, r.M2, of Loxoihnta (Palxohxodon) Tokunagai junior poses, in passing, to refer the muta-
mut. Matsumoto, 1929. Original in the Geological Institute of the Kyoto Imperial University. Upper Plio-
tion represented by the specimen
cene or Lower Plcistocene(?). .\fter Matsumoto, 1929.2, PI. vii, fig. 1. Two-thirds natural size. This molar
. . .

exhibits -j-ll)^ ridge-plates. Length as preserved 223 mm., estimated length about 240 mm., breadth 73 mm. at to Lox. {Pal.) Tokunagai junior, mut.
fifth ridge as preserved. nov."

JAPANESE SPECIES DESCRIBED BY SAHEKI AND TOKUNAGA (1931, 1934)

Parelephas protomammonteus matsumotoi

Saheki, 1931'
Figure 11.58

From Mishima, Kiraitsu District, Chiba Prefecture, Province of

Kazusa, Japan.
[The description and figure of tins subsi)ecies were not
studied by Professor ()si)()rn. Inasmuch, however, as \w.
referred Parelephas protomammonteus and P. protomam-
monteus proximus of Matsumoto to the genus Palseoloxodon,
the pre.sent subspecies has been inserted in this .section tnider
the generic designation of the original author, namely, Par-
elephas. — Editor.]
Parelephas protomammonteus (Matsumoto) matsumotoi Type op Parelephas protomammonteus matsumotoi
n. var. Saheki, 1931. "On Parelephas protomammonteus Fig. 11.58. Portion of left mandib\ilar ramus, with M.i in <ilu, of Parelephas
protomammonteus rnatsunwtoi Saheki, 1931, PI. XV, fig. 1, one-fourth natural size.
(Matsumoto) Recently Found in the Province of Kazusa."
From Mishima, Province of Kazusa, Japan.
Japanese Journ. Geol. and Geog., Vol. VIII, No. 3, pp.
125-129, PI. XV, 1 text fig. Type.—". fragment of . .

a left mandibular ramus, including and bearing

symphy.sis, base of the Sanuki bed of Dr. H. Matsmnoto, [l^'ootnote:
Ms, (71 .sv7//." Horizon an» Locality. From Mishima, — 'The name "Sanuki bed" here used is diffennit from what Mr.
Kimitsu Di.strict, Chiba Prefecture, Provinci; of Kazusa, F. Ueda calls by the same name, but may corres])()iid to his
Japan. {Op. cit., p]). 128, 129): "Lower Calabrian in age. "Umegase bed." '] which is considered to be the ba.sal horizon
The conglomerate of th(; region undoubtedly belongs to the of the Narita series; to the Upper Miura .series of Dr. H. Yabe and

'[See Chapter XXI of the present Memoir, p. 1 U6, note under "1931 Parelephas prolonuiminonlcm (Matsumoto) tnatsuimloi Saheki." — Editor.]
 THE LOXODONTIN^: PALyEOLOXODON OF JAPAN 1301

Mr. S. Nomura; to the Sasage bed of Mr. J. Makiyama; to the "Three anterior mental foramina are present, though Matsu-
Higashi-Higasa bed of Mr. Y. Otsuka, and to the Umegase bed of moto's specimen is reported to have four —
all along the ridge of the

Mr. F. Ueda. As to the age of the bed, the first proposes that it is —
diastemata the middle one being the largest."
Calabrian and correlates it to the Naganuma bed of the Miura "The Ma of this mandibular ramus measures 287 mm. in
Peninsula, while tlie last says it is Lower Pliocene and compares length as preserved, lacking only some lower part of the posterior
it to the Koshiba bed of Lower Pliocene in Miura. But accord- talon, consisting of eighteen ridges and a posterior talon. Its

ing to Dr. 0. Abel, the Elephas group first appeared in Middle maximal width is 76 mm. at the seventh ridge, and its height is
Pliocene, while nearly all the other paleontologists state that it ca. 130 mm. at the twelfth ridge, which just shows signs of wearing.
appeared in later Pliocene. According to Dr. H. Matsumoto,
first The frequency of ridges in 100 mm. is 6 on the inner side, 7 on the
'any well-established true Elephant is as yet unknown throughout outer side, and less than 6 at the grinding surface."
the world before the earlier boundary of Calabrian.' Moreover, "The worn surface of the molar is oval and more highly
Stegodon orientalis Owen
which is found associated with
(typicus) concaved than that of P. protomammonteus (Matsumoto) typicus
Parelephas protom.aninwnteus (Matsumoto) typicus Matsumoto in Matsumoto, as also the mid-ridges which are concaved considera-
this case, has never been found to be older than Upper Pliocene.
. . . bly to the anterior. The layer of enamel of the rather slightly worn
[Footnote: 'Mr. J. Makiyama stated that Stegodon orientalis in our ridges on the grinding surface is irregularly wavy; that of the
country, is Pleistocene, (Proceedings of the Third Pan-Pacific more worn ridges is irregularly and coarsely plicated; while that
Science Congress).']" of the still more worn ridges is gently waving and their enamel
"In view of the foregoing, I am convinced that the bed is layer shows regular, fine, and weak plication."
Calabrian in age, and is to be correlated to the Naganuma bed of —
"The layer of enamel is 1.8 2 mm. thick."
the Miura Peninsula, ... as Dr. H. Matsumoto does. As to its
stratigraphical position, however, I am inclined to favour Mr. F. Palseoloxodon yokohamanus Tokunaga, 1934'
Ueda's views, separating it from the so-called Sanuki bed of Narita Figure 1159
series as another unit of geologic formation [Footnote: 'This From Yokohama, Japan. Lower(?) Pleistocene.
concerns the problem of the boundary between Pliocene and [The following text has been prepared from a translation
Pleistocene. The conspicuous unconformity between Shimosuye- kindly furnished by Mr. Ushinosuke Narahara of the American
yoshi bed, i.e. the lower part of Narita series in the Miura Peninsula Museum. The original description by Doctor Tokunaga was not
site and the Naganuma bed has been observed recently by Mr. Y. seen by Professor Osborn. —Editor.]
Otsuka, as well as by Mr. J. Makiyama.'] the —
Umegase bed as it PalxoloTodon yokohamanus Tokunaga, 1934, pp. 363-371,
may be called." Type Figure. Op. cit., PI. —
xv, figs. 1 and 3 "Fossil Elephant Teeth found at Yokohama and Kakio, Kanagawa
[Fig. 1158 of the present Memoir].
Type Description.— (Saheki, op. cit., 1931, pp. 125-129):
"Since 1928, the conglomerate bed exposed at the river cliff of the
Koito, has been worked for grit in the construction of prefectural
roads newly opened in the village of Mishima, Kimitsu district,
Chiba Prefecture. In the course of the work in December, 1928,
and some
the left half of a mandibular ramus, bearing a cheek tooth
pieces ofbone of a certain mammal, was found in the conglomerate
bed at Higashi-Higasa of the village."
"The mandibular ramus of the specimen measures as follows
(in mm.):

"Mandibular angle seems to be conspicuously smaller than

P. protomammonteus (Matsumoto) typicus Matsumoto."
"Length from anterior end of symphysis to posterior
Fig. 1159. Type second right superior molar, r.M", of Palseoloxodon
end of Ms 455
yokohamanus Tokunaga, 1934, PI. vni, fig. 1. Length of molar as preserved
Length of symphysis 75-a 178 mm. From Yokohama, Japan.
Width of ramus just anterior to base of ascending bar 160
75" Prefecture," Journ. Geog., Vol. XLVI, No. 546, July, 1934 (in
Depth symphysis
of
Japanese). Type. —A second right superior molar, r.M-.
"The symphysis is very short-spouted. The ridges of diaste- Horizon and Locality. —Found at the mouth of the Tsurumi-
mata are rather concavely arched, instead of being linear as in gawa in 1931, Yokohama, Japan. Lower Pleistocene. Type
those of P. protomammonteus (Matsumoto) typicus Matsumoto, Figure. —Op. PI. viii, figs. 1 and 2.
cit.,

and sloping down obliquely from the anterior ends of the alveolar The author states that the molar is of a dark brown color and
margins to that of the symphysis more gently than those of P. very shiny. It has ten preserved ridge-plates, two probably lacking,
protomammonteus (Matsumoto) typicus Matsumoto. The anterior making a total of 12. Length of molar as preserved 178 mm., great-
end of the alveolar margin lies at a distance anterior to the posterior est breadth 76 mm., maximum height 177 mm., 5, 6 ridge-plates
end of the symphysis." in 100 mm. Regarded by the author as of Lower Pleistocene age.

'[See Chapter XXI of the present Memoir, p. 1408, note under "1924 Elephas namadieus naumanni Makiyama." —Editor
1302 OSBORN: THE PROBOSCIDEA

JAVANESE SPECIES DESCRIBED BY DUBOIS

Palaeoloxodoi? hysudrindicus Dubui.s, 1908 The long and narrow right and left third inferior molars

Figure IKiO (B 122 E3 and B 122 E 2C, Dubois Coll., Leiden) are beautifully

Ape Man, Fithecanthmpus, Middle(?)Ploisto- preserved and, coming from the same individual, firmly establish
Contemporary of tlic Trinil
ceiie.
the dimensions, the number
of the ridge-plates, and the low crown

KciulcMK-Sehieliteii, Pitliecanthropus zone, Ixjwer (or Middle?) Pleisto- of this progressi\'e mid- Pleistocene species; rMs measures 324
eeiio of Java. This zone, originally determined as Upper Pliocene, has recently mm., l.M.-i, 297 mm., as reproduced one-third natural size. The
(Dietrich) been regarded as Middle Pleistocene, because it also contains the ridge-plate formula (M 3 j'g'Jj) surpasses that of Palxoloxodon
very progressive species Stegndon airhivana. namadicus naumanni (M 3 \^), the genotypic species of Palu'v-
The cotj'pes below (Fig. 1160), reproduced from original loxodon Matsumoto.
l)lu)tographs kindly sent to the present author by Doctor Dubois^,
are of great interest as positively demonstrating that 'Elepha.s
hysudrindirus' is a very ]3rogressi\'e species of Paln'oloxodon with
a high ridge-plate formula:

I', hy.siiilniidiciis .\I 3 YsVi-

L.M3

L.M-

COTYI-ES OF PaI^^OLOXODON HYStlDBINDICUS. CoNTEMPORAUV WITH PlTHE-CANTHROP0S

Compare ly[M' of llrsprrnloxodon anliquus iiusonius (I'^ig. 1087).

Fig. 1100. Ekphas hysudrindicus Dubois, 1908, after photographs of the original eotypes kindly furnished by Dr. Eugen Dubois for this Memoir. Un-
iformly redviccd to one-third natural size.
L.M', a third left s\iperior molar (B 8.') Dubois Coll., L<'idcn) from the Kendeng deposits, .Java, of an aged individual with ridge-plates .5 46 more or less
worn, ridge-plates 1 4 compli'tcly worn away, ridge-plates 17 19+ still unworn. Length 209 mm., reduced to 70 mm. or one-third natural size. External and
crown views.
H.M.I, a third right inferior molar (H 122 E3 Dubois Coll.,Leiden) from the Kondeng deposits, ,Iava, of a iniddlc-.iged individual with ridgc-plales 1 S more
or less fully wcrn, ridge-))lates 9 1 9/v complctelj' unworn. This griixler measures 32) mm. in length; il is reduced to lOS mm. or one-third nalunil size. Crown
views.

L.Ma, a tliird inferior grinder (B 122 E 20 Dubois Coll., Leiden) from the Kendcng deposits, .Java, of a more aged individual with ridgc-plates 3 fully 1 1

or partly worn and ridge-plates 1 3 worn away or imperfect, ridge-plates 12-19 unworn. This grindiT measures 297 mm. in length; it is reduced to 99 mm. or
one-third natural size. Crown and lateral aspects.
 THE LOXODONTIN^: PAL^OLOXODON OF JAVA 1303

The cotypo superior molar (B85 Dubois (

'oil.) of Palxolo.rodan verfiihrt die Anwesenheit des E. indirus uiitcr der Kciidcii!^-
on ridge-plates 5 to
hijfiudrindirus (Fig. 1160) exhibits, especially Fauna anzunehmon, obgleich die Lamellen-Zahl der fossilcn

12, enamel foldings or plications more numerous and deeper than javanisehen Molaren nicht iibcr 19 geht, also \ipl gcringer ist als
those of the type of P. namadkus namadi (Fig. 1153); these bei E. indicus; der (spater bckannt gewordenc) Schiidel niihert
Ijlications resemble those of the 'Elephas namadiciis' (Fig. 1189) de- sich aber bedeutend mehr der Siwalik- als der lebenden Art,
by
scribed Lydekker, 1886.2, as "from the Pleistocene between namentlich durch sein Profil, durch die grtissere laterale Ent-
Kanagawa and Tokio (Yedo), Japan"; it was this crimping or wickelung der parietofrontalen Hocker, und diu'ch die geraden
jjlication andthe absence of the ioxodont sinas' which led Dubois Alveolen fiir die Stosszahne, doch ist die Breite zwischen den
to lelate these grinders to Elephas indicus. Schlafengruben bereits grosser geworden; hierin niihert sich die
Dubois (1908) named this species as among his ''Kendeng- javanische ausgestorbene betrachtlich der lebenden indischen .\rt,

fossilien"; Stremme (1911) remarked that Dubois had no Elephas als deren unmittelbaren Stammvater wir die erstere wohl un-
remains from Trinil in his large collection. Dubois (1908) distin- zweifelhaft anzu.sehen haben."
guished Elephas hysudrindicus as standing near Elephas hysudricus, (Stremme, 1911, p. 144): "Ein wichtiges Leitfossil wiire

but still more close to the living Elephas indicus consequently he

;
eventuell Elephas, dessen Zahnbruchstiick Janensch dem Elephas
named it Elephas hysudrindicus. Stremme (1911, p. 144), however, antiquus am iiiichsten stellt. Das Stuck stammt nicht von Trinil;
remarked on its closer relationship to Elephas = Hesperoloxodon]
[ auch Dubois hat keiiie Elephas-Reste von Trinil in seiner grossen
antiquus: "In bezug auf die Zahl der Joche steht dieser dem E. Sammlung. Dubois selbst ist nicht geneigt, an Altersunterschiede
a7itiquus naher als dem rezenten E. indicus." der Fundstellen bei die.sem eigenartigen Fehleii zu denken, sondern
Elephas hysudrindicus Dubois, 1908. "Das Cleologische Alter er halt die Lebensweise der Elefanteii in bezug auf die Verteilung

der Kendeng-Oder Trinil-Fauna." Tijdschr. Nederl. Aardr. ihrer Reste fiir mas.sgebend. Dubois hat nach seinem gr'isseren
Genoots. Amsterdam, Tweedie Serie, Deel XXV
B, No. 6, p. Material des Elefanten diesen als Elephas hy.'iurlrindicus n. s]).

1257. CoTYPEs. — Molar teeth with lamellae not exceeding bezeichnct und stellt ihn namentlich nach dem Vergleiche der
19)2; skull subsequently found (see description below), but type nahe an Elephas hysudricus aus den Siwalik-Schichten vom
Schiidel
specimen not clearly designated or figured. Horizon Pendschab und den subhimalajischen Bergen und aus dem Alt-
—
AND Locality. Kendeng-Schichten, Java, Middle? Pleistocene. pleistocjin des Narbadatales."
—
CoTYPE Figures. Not published by the author (see Fig. 1160 Stremme also 'observes {op.
cit., p. 143): "Der Elephanten-
of the present Memoir). zahn zeigt nach Janensch mehr loxodonten Charakter als Elephas
—
Original Description. (Dubois, 1908, p. 1257): "Auch von indicus imd erinnert mehr an Elephas antiquus. (Der Vertreter des
Elephas liegt unter meinen Kendengfossilien nur eine einzige Art E. antiquus in Indien, nach Leith Adams Elephas namadicus Falc.
vor, woriibcr mich genaue Durchsicht der vielen gesammelten aus dem Altpleistociin des Narbadatales, wird aber ausdriicklich
Molaren und besonders auch Bekanntwerden mit der Schiidelform von Janensch als verschieden bezeichnet.)" Janensch ("Die
belehrt hat. Es ist diese eine dem Elephas hysudricus sehr nahe Proboscidier-Schadel der Trinil-Expeditions-Sammlung," 1911, p.
stehende, jedoch noch mehr als letzterer sich dem jetzt lebenden 194) does not comment directly on the characters of the si>ecies
Elephas indicus annaherende Art, die ich deshalb als Elephas E. hysudrindicus Dubois beyond the observation quoted from
hysudrindicus n. sp. bezeichnen \vill. Hatte man die Art aus den Stremme above.
Siwalik-Schichten bereits als die vermutliche Stammform des Osborn, 1930: This progressive stage of Palxolo.rodon, like
lebenden asiatischen Elephanten erkannt, durch die neuc Art von that of Stegodon airdwana, is of great anthropological interest

Java wird diese verwandtschaftliche Beziehung noch klarer. Die as establishing the Middle Pleistocene age of Pithecanthropus
grossc Ahnlichkeit der Molaren hatte mich anfanglich sogar dazu erectus, first pointed out by Dietrich.
 :

IX. SUMMARY OF GEOGRAPHIC DISTRIBUTION ALONG THE EASTERN COAST OF ASIA

!»nn.T|L,«*JWiER»rA

Altogether, the monographic re-

searches of Matsumoto and Makiyania
supplement those of Falconer, of Owen,
and of Osborn and prove that primi-
tive species of the genus Palseoloxodon
spread eastward across all of southern
Asia during Upper Pliocene time until
their march was arrested by the an-
cient Pliocene shore lines of Japan
and of the East Indies, including
Burma, Java, and Sumatra. It is in the

Kendeng stratum of Java, now regard-

ed as of Middle Pleistocene age (as fully
treated above. Chap. XIV, the Stego-
dontinse, especially p. 887) that we find
another advanced stage of Palxoloxo-
don, to which Dubois applied the name
'Elephas hysudrindicus.'

The bathymetric maps of Chisholni

and Leete (Fig. 1161) andof Yabe (Fig.
1162) are extremely interesting and im-
portant in displaying three features,
namely

(1) The ancient coast of northern

and southern Asia and of the East In-
dies during late Pliocene and early to

mid-Pleistocene times; (2) the succes-

sive northerly range over the Bering
Strait region of Archidiskodon, {?)Par-
ekphas, and Mammonteus to North
America; (3) the southerly and south-
easterly range of Stegolophodon, Stego-
don, and Palseoloxodon, and finally of

Elephas.

Vip,. IKil. .Iiipaii as part, of tlir Asiatic continont

ill I'lio-Plcistoccnc timo, slioxviiiK in white tho broad

true continental l)order on tlio 100-faMiom or (iOO-foot
lino. Rcproiiiicod by permission after IjonKmaTis' New
School Atlas. This shows clearly the niiKration lines
through vSumatra, Borneo, the Philippine Islands,
.lapan, and Kamchatka. Compare with the more
recent figure after Yabe (Fig. 1U)2).

1304
 THE LOXODONTIN^: GEOGRAPHIC DIRTRfBUTION ALONG ASIATIC COAST 130r)

The three outstanding facts are:

The geologically oldest or Lower Pliocene' range of the Stegotlonts'- covers an area from the type locality of
Stegolophodon latidens on the Irrawaddy River, Burma, 1300 miles southeast, to the northern Bruni district of
Borneo, namely, Brit. Mus. M.2498, described by Lydekker (1886.2, pp. 75, 76, Fig. 19) as 'Mastodon latidens'
but which the present author made the type of Stegolophodon lydekkeri in Vol. I, p. 700, of the present Memoir.

Of Upper Pliocene and Lower Pleistocene age is the transcontinental range of descendants of North African
species of Palseoloxodon, such as Palseoloxodon jolensts and P. atlanticus of Pomel, and P. recti of Dietrich, to
Japan ; must have occurred in late Pliocene or early Pleistocene time, because the more primitive Japanese
this

stage described by Matsumoto, namely, P. tokunagai (M 3 (iTTiTvi), corresponds closely in ridge formula with the
more primitive North African species and those of the Mediterranean Islands.

Finally there is the Palseoloxodon namadictis stage (M 3 J|:) of the Middle to Upper Pleistocene of India,

which corresponds closely with the P. namadicus naumanni of Makiyama, although the latter has a higher ridge
formula, namely, M 3 H.
Of the true Stegodonts, the Stegodon elephantoides of the Irrawaddy River (M 3 j^rro); including Stegndon
cliftii of Falconer (l.M 1 with ,,^^ ridge-crests), may be ancestral to the Stegodon insignis birmanicus of Burma
(M 3 Y^ij) and to the S. orientalis grangeri (M 3 y^) of the Yangtze River, China.

Fig. 1 162. Japan as part of the Asiatic continent in Piio-Pleistooene time

and two landlocked internal basins. After Yabe, 1929, ma]), p. 169, who
describes the latest land connection of the Japanese Islands with the Asiatic
continent as follows (p. 168):

"The sea floor now encircling the Japanese Islands to the depth of some
720 m., in the average, was once a land surface and the land submerged below
the sea level in a time geologically not much remote from the present. Prior to
this great submergence of land, the Japanese Islands were some 720 m. (237.5 ft.)

more elevated than they are at present and directly connected with the
Asiatic continent, as the annexed map shows. The 720 m. line, which marks
the true continental border of eastern Asia more properly than the 160 m. line,
follows closely the present Pacific coasts of Taiwan, the Riukiu Group, Kyflshu,
Shikoku, Honshij Hokkaido, the Chishima Group and Kamchatka. The
entire region lying inside this line is regarded once to have been a dry land, with
the exception of two extensive basins —one occupying the greater part of the
Japan Sea and the other [the] southernmost part of the Okhotsk Sea — , and
a narrow, Unear one lying along the inner border of the island arc of the Riukiu
Group. All these basins, now more than 700 m. deep, are thought by the
writer as having existed there as entirely or almost land-locked basins during
the continental stage of the Japanese Islands; ... It was at the time of the
maximal extension of land in the continental stage of the Japanese Islands
that the land took the outline stated above. ... In other words, the continental
stage of the Japanese Islands continued from the time of land emergence to the
90 m. line to the time of land submergence to the same line."

'[Probably Lower Pleistocene —see note on page 824 above. — Editor.]

-(See footnote on p.age 837 above. — Editor.]
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 Chapter XX

THE SUBFAMILY ELEPHANTIN/E (SUPERFAMILY ELEPHANTOIDEA), OF

EASTERN ASIA, INCLUDING THE RECENT ELEPHAS AND RELATED
LOWER PLEISTOCENE SPECIES

Classical references to elephas. Establishment by linn^us of the genus and species elephas
iNDicus (1735-1758). Characters of the subfamily elephantin^e and included species and subspecies or
geographic varieties, by corse, de blainville, falconer, temminck, matsumoto, and others (1799-1927).
Fossil lower pleistocene and upper PLIOCENE species described BY falconer and by osborn (1845-1930).

I. Historical Inthodiction and Nomenclature (850? Elephas indicus sinnatramis Tommiiifk.

B.C. -19.36). Elephas indicus hirsutus Lydekkor.
1. Falconer (1868) on the specific unity and vertel)ral Elephas indicus Buski Matsumoto [
= 1Palseolo.vndnii
formulae of the Asiatic elephants. hushi].
2. Corse, de Blainville, and Falconer on characters of the 2. Distinctions and measurements of the Indian elephant.
geographic varieties.
III. Characters of the Upper Pliocene and Lower Pleisto-
.3. Living specific or .subspecific forms, continental and
cene Species Hypselephas hysudricus and Platele-
in.sular.
PHAS PLATYCEPHALI'S.
4. Fossil forms more or less closely related to Elephas
1. Falconer's original descriptions of Elephas [Hypselephas]
indicus.
hysudricus.
5. Names of species and subspecies of the subfamily
2. Observations of Osborn on the fourteen specimens col-
Elephantinae in order of description.
lected by Barnum Brown (1922) referred to Hypsele-
II. Characters of the Subfamily' Elephantin/E and Incu'd- phas hysudricus.
ED Genera and Species. 3. Cranial characters and affinities of Hypselephas hysudri-
Genus Elephas. cus.
1. Systematic description of species of Elephas. Juvenile crania with dentition.
Elephas indicus LiiiniEus. 4. Genus Platelephas.
Elephas indicus ceylanicus de Blainville. Platelephas platycephcdus.
Elephas indicus hengalensis de Blainville. Appendix.

It is a striking circumstance that we have not as yet discovered the direct ancestry of either of tlie typical
living elephants, namely, the Elephas indicus of India or the Loxodonta africana of Africa. Both of these typical

Uving species, around which center historical as well as zoological references from the time of Homer (850? B.C.)
to the present time, are still lacking the phylogenetic background of their history. All attempts both by zoologists
and palaeontologists to trace back either Elephas indicus or Loxodonta africana to previously living or fossil forms'

are shown in the present Memoir to be problematical, because neither in the cranium, the teeth, nor the skeleton do
any of the known fossil elephants meet all the phylogenetic conditions. Even the Platelephas platycephalus of the
Upper Phocene and the Hypselephas hysudricus of the Lower Pleistocene of India, very fully described below
in the present chapter, fail to meet all the conditions ancestral to the true E. indicus. The conclusion is inevitable,

that in some as yet unexplored region of Asia the direct ancestors of E. indicus were slowly evolving, while in some
unexplored area, probably west of central Africa, the very conservative ancestors of L. africana were also slowly
evolving. In brief, we now know the phylogeny of most of the other subfamilies of the Elephantidae better than
we know the phylogeny of the Elephantinae, typified by the genus Elephas.

'[According to Pompl (189.i) the true Loxodonta africana occurs in comparatively recent deposits of North Africa, and Dart (1929) refers to t.lie discovery
prima Dart, 1929, Chap. XIX above, p. 1287). Editor.!
in llie Pilandsl)erg (Transvaal) of a ])rimitive foi'm of L. africana (of. LoxotUinIa —
1307
 ;

1308 OSBORN: THE PROBOSCIDEA

The early historical references or allusions to this most majestic of all living quadrupeds of the Asiatic con-

tinent are full of fascination and interest.' The uses of these animals in industry, in transportation, in art, and in

war would fill volumes. But here, as a fitting conclusion to the previous nineteen chapters of the history of the

Proboscidea, we must confine ourselves rigidly to the slow emergence and clarification of zoological nomenclature

and anatomical analysis from the time of the Greeks onward to the present time.

As in the Loxodontinse (Chap. XIX) and in the Mammontin» (Chap. XVIII), the strict application of the

Rules of the International Commission on Zoological Nomenclature (Congress of Berlin, 1901, and of Monaco,
1913) to types and descriptions of the early part of the nineteenth century is beset with insuperable difficulties, as
exempUfied in the whether the Indian elephant should be called 'Elephas indicus' or 'Elephas
classic case of

maximus.' The grounds for the adoption of 'Elephas maximus' are herewith shown to be either precarious or
absolutely untenable.

I. HISTORICAL INTRODUCTION AND NOMENCLATURE (850? B.C.-1936)

The very ancient name 'elephant' has risen in rank, becoming the type of the subfamily Elephantinae, family

Elephantidse, superfamily Elephantoidea.

—
History of Nomenclature. Elephas is a Greek name of uncertain origin, subsequently Latinized. It is

treated by Liddell and Scott (1883) and by James A. H. Murray (1891) as follows:

(Liddell aiul Scott, 1883, p. 454) eXe^as, avros, 6 the elephant, first mentioned by Hdt. as a native of Africa, 3.114., 4.191
:

whereas Arist. H. A. 2.1, 45 treats only of Elephas Indicus, of. 9.1, 30, etc., though the African is mentioned by him in Gael.
2.14,19:^not generally known in Greece till the time of Alexander, Paus. 1.12,4. IL known to Horn, only as the name for the
elephant's task, ivory, II. 5. 582, and so Hes. and Find.; for ivory was brought by Phoenician traffic to Greece long before the
animal was known to Greek travellers; Hdt. calls the tusks more accurately JXe(^afTOs vbovr^s, 3. 971: Hom. brings false —
dreams through an ivory gate, v. sub tXe^aipo/iat, (Pott and others refer to the Hebr. Eleph {ox), and compare bos Lucas,
. . .

the old Lat. name of the elephant, Lucret. 5. 1301; as Paus. (9.21, 2) calls a rhinoceros ravpos KIBlottlkos. On the other
hand the Hebr. name for the animal, ihah, recalls the Skt. ihhas, which is identical with the latter part of €X-€</>as, and the
first part of the Lat. eh-ur, whence iv-oire, etc.)

(Murray, 1891, III, Pt. 2, p.84): Elephant Gr. t\e4>as (gen. k\k^avTo{).
. . . The refashioning of the word after Lat.
seems to have taken i)lace earlier in Eng. than in Fr., the Fr. forms with el- being cited only from 15th c. Of the ultimate . . .

etymology nothing is really known. As the (ir. word is found (though only in sense 'ivory') in Homer and Hesiod, it seems
unlikely that it can be, as some have supposed, of Indian origin. The resemblance in sound to Heb. eleph 'ox' has . . .

given rise to a .suggestion of derivation from some PhcEnician or Punic compound of that word others have conjectured that the ;

word may be African.

Elephas was named by Ray (1693, p. 131 = p. 123 of authors]), and was included in the "Systema
as a genus [

Naturae" of Linnaeus in 1735 in the same division of quadrupeds with the rhinoceros (see below facsimile of portion
of page 10 of the First Edition of the "Systema Naturae" of Linnaeus, 1735). The genus Elephas Linn, dates from

1735, the species indicus dates from 1754, the species maximus dates from 1758, as shown in the three facsimile
Linnaean definitions from (1) the "Systema Naturae," First Edition, 1735, p. 10; (2) Memoir of the Museum
Adolphi Friderici Regis, Holmise [Stockholm], 1754, p. 11, and finally in the Tenth Edition of the "Systema
Natura;," 1758, p. 33.

Dr. C. Davies Sherborn of the British Museum, the greatest living authority on generic and specific names,
believes that it is impossible to determine which elephant, African or Asiatic, Linnaeus had in mind in proposing

the term maximus, a name without standing in Sherborn's opinion:

'Kunz, Gt'orgi: V., lOlti. Ivory and the Elephant in .\rt, in Archseology and in Science.
 THE ELEPHANTINE: HISTORICAL INTRODUCTION AND NOMENCLATURE 1309

(Sherborn, 1929, letter of July 14) "In reply to your enquiry about Elephas maximus, Linnseus. Linnaeus' type seems to
have been Ray, Quadr. 131. Blainville, Elephants (p. 47) in his 'Ost^ographie,' says Ray saw a young elephant in Florence.
Now Ray gave no locality for his specimen but described it in much detail. In the Mus. Adolf. Frid. in 1754 Linnaeus says
'Indien' both in his Latin and his Swedish version and why or on what authority he puts the word Zeylansk in opposition to
'indicus' passes my understanding. So far as I know there is no authority for Zeylansk and in the S. N., ed. XII, 48, Linnaeus
clearly spatchcocks the whole lot into one species both for Asia and Africa and considers that there is only one Elephant. And
as that beast has been more or less of a domesticated animal since before Alexander the CJreat, I don't think Linnaeus was far
wrong in such a guess. In such a case it appears to me that you have no option but to say 'indeterminate' and proceed to the
next person who divided the subject, e.g. Blumenbach in 1797 for 'africanus' and Cuvier in 1798 for 'indicus.' Unless one can
definitely say what beast Linnaeus saw when he gave the name 'maximus,' which apparently is now a thing impossible to decide,
it appears to me that 'maximus' has no standing and must be swept aside." [See Lonnberg (letter of 1929) quoted below,
also footnote on page 1323].

In 1754 the specific name indicus appears and in our present language this was the original genotypic species
of Elephas; it was universally used by all writers on the elephant family until 1901. Unfortunately, and
as such
for some reason unknown to us, in the Tenth Edition of the "Systema Naturae" Linnaeus substituted the name

Elephas maximus Linn, for that of E. indicus Linn., which, with all other names in the Tenth Edition, was officially

b 1
 — : "

1310 OSBORN: THE PROBOSCIDEA

believed that the number of created species was hmited; mind the Indian and African elephants were
in his of
the same species. This is indicated in the following letter from Professor Einar Lonnberg (July 31, 1929)

LiNX.Eis' Type of IOlkphas indicis (LiiNNBEKC, letter, July "In leply to your letter coiicorning the typo
,31, 1929).
of LiiiiiiBUs' Elephas iitdiciis in Museum Adolphi Friderici Regis, llfA, may
say that it is not a skeleton nor a stuffed animal
I

but a foetus, which still is kept in this Museum. The King had bought it together with several other natural history speeimons

from Holland, where it once was kept in 'Museum Sebae.' It is of an African Elephant, probably from West Africa.
The reason why Linujeus speaks of India and Ceylon is probably that he quotes Rajus in the first rank. I have not looked
up what Rajus says, but you can easily find that yourself after the quotation: 'Raj. quadr. 123.
In Sj'stema Xaturie, ed. X., Rajus is also quoted in the first rank and Seba only in the third rank. But Seba's specimen
is the same as that of Mu.seum Adolphi Friderici Regis, 1754. Seba has figured his specimen in his 'Thesaurus,' Vol. I, tab.
CXI, fig. 1, and he says that it is 'Een ongeboren Olifant uit Africa.'
From the above is evident, that the name Elephas maxi»nif< L. 1758 refers as well to the Indian (Rajus quoted first) as the
African (Seba quoted as third) Elephant."

Genus (Linn.eus, 1735, 1754, 1758).— The genus Elephas Linnseus dates from 1735 ("Systema Naturae,
First Edition, p. 10) without mention of a genotypic species. In 1754 (Memoir, Mus. Adolphi I'riderici Reg.,
Stockholm, p. 11) reappears the generic name Elephas with the species indicus, habitat India (Zeylansk), indicat-
ing that Linnseus based the species on the Ceylon variety of elephant. In 1758 ("Systema Naturae," Tenth
Edition, p. 33) again appears the generic name Elephas, habitat India (Zeylonse), with the new specific name
Elephas maximus. This edition (known as the Editio Decima, Reformata) was reprinted in 1901 and adopted as
the standard by the Fifth International Congress of Zoology of 1901, held at Berlin.

MAMMALIA BRUTA. Elephas, 33 In 1795 the name Elephantus was used by Geoffroy
and Cuvier, and in 1801 was by Cuvier and Lace-
II. BRUTA. cited
pede as Elephantus indicus, as Cuvier continued to ig-
'Denies Tr'tmores nulli utrinque.
nore the terminology of the "Systema Naturae" of
Linnseus.
J. ELEPHAS. 'Dentei Trimores nulli.
Laniarif fuperiorcs elou-
—
Genotypic Species. The species indicus of Lin-
naeus, which we select as genotypic, is important, be-
Trobofc'ts iongimma, preheufilis.
Corpus nudiiiiculura. cause of the recent substitution by the Congress of 1901
tnaxiraus I. Elephas. Ri'u quadr. ix^. Syjl Hut.ii. Sd.muf.x. of Elephas maximus Linnaetis (1758) for the original
t. HI./. I.'
name Elephas indicus Linnseus (1754). Why did Lin-
ElcphantUS. Gtjn. qvadr. 577. Aldr. quadr. I. I. c. 9.
jfoti/l. quadr. 30 t. 7S. /. 9. naetis abandon the appropriate name Elephas indicus
Habitat in ZeylonsE paludofit ad /Imiit/ , edit RamHy and substitute the inappropriate name Elephas maxi-
Ctcos , GuiUttditta femirta , Frumenrum.
Jilaximum quadriipet. Oc\iV\ farzi. Dcntes Laniarii /*-
mus? Is not the explanation found in his behef that the
feriorei txj'erti {F.iur). Aurts amplijjim,t , fcndulte I ndian and the African elephant were of the same species?
dentalte; ull. angl. %ij. p. loii. Cutis craffiffitna,
catlofa. Mimmx
loltorum pedum.
2 jtijeta peiflui.
Genua flexiiu.
Ungues
CoUum
in apiciiui
hrevc.
First. — There is little doubt' that the genotypic
I'tohoi^ch exttnfilii , acute odoraiii , l»co ma-
lotigijjfirria
, species Elephas indicus Linn., 1754 { = Elephas maxi-
ntii ipji ixjcrviem; ea tibum fotumque haurit , hofltm-
mus Linn., 1758) was a domesticated elephant from the
que peliie ; ea pr.tcifa occiditur ; murem metuit ful>
foinna oh trachea iiiferiionem. Retro coH isf mittgit, island of Ceylon in which occurs the native or indigenous
Portat domos , rehire cello tK/tdente ; in hello arma-
tur falcibiij ; vlaufculo inter axin tsf atlantem furif small, typical Ceylonese variety named by de Blainville
fui occiditur cxterum prtidens doiilit. This subspecies of
j ,
(1845) Elephas indicus ceylanicus.
Fig. 116(). Kucsiiiiilc of page 33, in .saiiii' size, of Linnaeus' original Tenth Ceylon is clearly distinguished by its small, slender
EMition of the "Sy.stema Natura-," 17.58, in whiih Elephas maximus i.s substi-
tusks pointing obliquely downwards, by its narrow,
tuted for Eteplia-s iiulicus, and the germs Eli/ihas and specie.s maximus are
defined. Habitat in Zevlona\ elevated head, with narrow narial openings, by its gentle

'In 1758, as shown in the aecompanying facsimile of page 33 of the Tenth Edition of the "Systema Natura?," Linnams follow.s Ray, as of first rank:
"maximius. 1. Elephas. Raj. quadr Habitat in Zeylona"."
 :

THE ELEPHANTINE: HISTORICAL INTRODUCTION AND NOMENCLATURE 1311

disposition facilitating domestication, and by its typical 20 dorsal vertebrae —characters which would now entitle
it to distinct subspecific rank from the large, broad-headed, and otherwise different elephant of Bengal (see com-
parative figures 1171 and 1170). The subspecies ceylanicus, moreover (Osborn, 1930), resembles the small,
slender-tusked Elephas indicus var. Mukna of Falconer and Cautley (1847).

Second. —The characters of the Ceylonese genotypic species Elephas indicus = E. maximus Linn.) are, how-
i

ever, confused by the fact that the larger Bengal elephant was imported into Ceylon, and it may have been one
of these larger Bengalese types of India imported into Ceylon which Linnaeus had in mind in applying his second
specific name Elephas maximus. On this point Lydekker observed (1916, p. 82)

The name Elephas maximus by a Ceylon elephant, and at first sight it would seem that Ceylon should be taken
is typified
as the typical locality of the species. But there are two races of elephants in that island one probably indigenous in which ;
— —
the tusks of the males ai'e insignificant, and the other almost certainly introduced —
in which they are large. Now it seems —
almost certain that the big-tusked race formed the type of Linne's species; and if so, Ceylon will not be the typical locality.
Unfortunately, it is uncertain whether the tusker-race was imported from the Indian mainland ... or from Lower Burma. . . .

Under these circumstances the writer [Lydekker] has considered it advisable to take southern India (say Mysore) as the type
locality, and to regard the small tusked Ceylon form as a second race.

Third. — From the above historic resume it appears technically that Linnaeus named as the genotypic species
the Ceylonese variety Elephas indicus ceijlanicus ; that Linnaeus' type specimen, however, was according to
Lonnberg (see letter, July 31, 1929, on p. 1310) the foetus of an African elephant {Loxodonta africana) ;
that in the
substitution of Elephas maximus for Elephas indicus he had both the African and the Indian elephant in mind,
apparently in the beUef that they constituted a single created species. For these reasons it is preferable to retain

the name Elephas indicus.

, ^
ii::.^^~iM^ d»tRv!

Indian Elephant Gkoop

Fig. 1107. The specimens in this grou]) are from tlie liills in the Province of Mysore, India, about 3') miles south of the city of Mysore. They were shot in
the spring of 1923, by Mr. Arthur S. Vernay, who presented them to the American Museum of Natural History. While they are not record specimens (see
where one of the
Fig. 1194, tallest elephants is given an estimated height of 10 ft. 6 in.), they are very fine examples and form an imposing central group in the

Vernay-Faunthorpe Hall.

The male (Amer. Mus. .54453) is at the left ; the female (Amer. Mus. 54452) at the right.

Estimated Measurements
(See caption to Fig. 1194)

Male
Skeletal height at shoulder
Height in the flesh
Scapula
Humeri IS
Ulna
 :

1312 OSBORN: THE PROBOSCIDEA

1. FALCONER (1868) ON THE SPECIFIC UNITY AND VERTEBRAL FORMUL.^ OF

THE ASIATIC ELEPHANTS
(Continued from pp. 930, 931 of the jjiesent Memoir)

As to specific unity, Falconer ("Palseontological Memoirs," Vol. II, pp. 267-270), after prolonged residence
in Ceylon and India and very careful study, erroneously concluded that there were no constant specific dif-
ferences either in the vertebral forniulse or in the structure of the cranium between the Bengal, or continental
Indian elephant, and the insular elephant of Ceylon, as appears in the following summary of his observations and
his conclusion that there is but a single species of Asiatic elephant:

Sumatran Elephant: 20 dorsals and 20 pairs of ribs (Schlegel)— constant.

Ceylon Elephant: 20 dorsals and 20 pairs of ribs (Peter Camper, Cuvier, de Rlainville).
Brought to Paris in 1795 4 lumbars; total dorso-lumbars 24.

Indian Elephant [Bengal?]: 19 dorsals and 19 pairs of ribs (Schlegel); by no means certain that the
Indian [Bengalese]
Schlisgel states that "all the number is constantly limited to 19.
elephants which he had examined had, [See 20 dorsals cited below, 'Duvaucol' and 'Choonee.']
without exception, only 19 dorsal verte-
brae and 19 pairs of ribs."

Bengal Elephant 20 dorsals. Schlegel meets thi.s exceptional case by the hypothesis that the
Diivaucel male skeleton in Paris Mu.seum live animal may have been imported from Ceylon into Bengal.
[see de Blainville, 1839-1864, PI. in
—
Fig. 1170 of the present Memoir].

Bengal Elephant: 20 dorsals and 20 pairs of ribs.

"Choonee." Imported from Bengal in 1810,

on board E. I. C. ship "Astell" by Capt.
Hay. College of Surgeons.

"This case, coupled with the Duvaucel skeleton in the 'Jardin des Plantes,' seems to establish, without .searching for
others, that the Continental Elephant of Northern India varies in the number of its dorsal vertebrae from 19 to 20, as the
African varies from 20 to 21." Footnote: "The ingenious vi(>\v advanced by Prof. Schlegel regarding the inverse relation
between the number of laminx in the molars and the number of dorsal vertebrae in the different species (supra, p. 263), does
not appear to be tenable again.st the evidence adduced above, of the numerical variability in the living species."

Bengal Elephant: These two skulls agree in general form and proportions. In the Ceylon
Brit. Mus. and Indian elephants the crania are "so closely similar, that, in a museum.
Killed in jungles on banks of Ganges, at no J
without a record, the mere form will not instruct the observer whence the
great distance from Meerut, in May, 1833 1 —
specimen came whether contiiK^ntal or insular."
Ceylon Elephant (Falconer's statement is not supported by de Blainville's plate .showing
C'oUege Surgeons 2656 the Ceylon and Bengal crania in jjrofile, as reproduced in our Fig. 1170.]

"The hypothesis entertained by Professor Schlegel, upon the statement of Diard, that Ceylon Elephants are frequently
imported into Bengal is, I am satisfied, untenable. On a review, therefore, of the whole case, the evidence in every aspect
. . .

appears to fail in showing that the Elephant of Ceylon and Sumatra is of a species distinct from the Continental Indian form.
. The result of this range of observation, combined with long osteological study, has been to establish the conviction in my
. .

mind that there is but a single species of Asiatic i;ici)liant at present known, modified, doubtless, according to his more north-
ern or southern habitat, but not to an extent exceeding that of a slight geographical variety."

In Falconer's opinion the cases above cited establish the fact that Elephas indicus, including the continental
and insular varieties, varies in the number of dorsal vertebrae from 19 to 20, in contrast to the African elephant
which varies from 20 to 21.

Osborn, 1929: From the above detailed observations of Falconer, together with those cited from Falconer
on i)p. 930, 931 of Chapter XV of the present Memoir, it appears probable that: (1) In the insular Ceylon ele-
 :

THE ELEPHANTINE: HISTORICAL INTRODUCTION AND NOMENCLATURE 1313

phant, Elephas indicus ceylanicus de Blain., there is a constant number of 20 dorsal vertebrae and 20 pairs of ribs
together with certain constant differences in the cranium, tusks, etc., clearly establishing the subspecies; (2) in

the continental form the vertebral formula is not constant, 19 dorsals and 19 pairs of ribs are recorded in certain
skeletons of Bengal, while 20 dorsals and 20 pairs of ribs are recorded in other cases. (3) This would indicate that
on the continent there are two hereditary blood strains, one characterized by 19 dorsals, the other by 20, possibly
arising from the race indigenous to Bengal mingled with the race indigenous to Ceylon. (4) It would be interesting
to ascertain whether there is a correlation between these two vertebral strains and the two cranial strains respec-
tively known as 'Mukna' and 'Dauntela.' (5) The case may be parallel with that seen in the Equida?, namely:
(a) Pure Arab strain of horses {Equus caballus africanus), in which there are 23 dorso-lumbar vertebrae, (b) the

pure northern strain (Equus caballus nordicus), in which there are 24 dorso-lumbar vertebrae; there are 5 lumbar
vertebrae in the Arab strain and 6 in the Nordic strain, the rib-bearing dorsals being 18 in each subspecies.

2. CORSE, DE BLAINVILLE, AND FALCONER ON CHARACTERS OF THE

GEOGRAPHIC VARIETIES
(Continued more in detail on pages 1325-1333 of the present chapter)

It ai)pears that the subspecific forms of Elephas indicus are less numerous and somewhat less diverse than
those of Loxodonta africana described above in Chapter XIX. The obvious explanation is that there is far less

physiographic variation in the range of the Indian as compared with the African elephant.

The geographic varieties or subspecies of the Indian elephant which have been successively named are:

In 1841: Elephas indicus Isodactylus Hodgson, of Nepal, Tarai, N. India.

Elephas indicus Heterodaclylus Hodgson, of Nepal, Tarai, N. India.

While Hodgson did not name the above subspecies until 1841, he observed as early as 1832 the differences in

the Indian elephant inhabiting the Ceylon and Bengal regions:

(Hodgson, 1832, pp. 344, 345): "The elephant and rhinoceros abound in the forest and hills of the lower region of Nepal,
where they breed, and have their fixed abode; and whence, in the season of the rains, they constantly issue into the cultivated
parts of the Tarai to feed upon the rice crops. Both these genera are entirely unknown to the central and northern regions.
The elephant is that so well known as the Indian variety, and as such is contra-distinguished from the African variety. But it
may be questioned, if there be not two distinct varieties or species in India alone, viz. the Ceylonese, and that of the saul
forest [Bengal]. The former differs materially from the latter by having a smaller lighter head, which is carried more elevated,
and by higher forequarters. It is also said to be larger, and of a more generous and bold temper. The difference of size, how-
ever, is certainly a mistake. I cannot speak to the point of temper."

In 1845: Elephas indicus ceylanicus de Blainville, the elephant of the island of Ceylon.
Elephas indicus bengalensis de Blainville, the elephant of Bengal, continental.

De Blainville figures in PI. in [Fig. 1170 of the present Memoir] of his "Osteographie" of 1839-1864 the two
subspecies from Ceylon and Bengal respectively, describing them briefly in the legend as follows

E. Indicus Ceylanicus, a grandes defen.ses. De profil, avec la mandibule en place; sexe inconnu.
E. Indicus Bengalensis, a grandes defenses. De profil, avec la mandibule en place. D'un individu male dont le squelette, prepare
par M. Duvaucel, a ete envoye par lui au Museum.

Geographic distinctions between the two continental races or varieties in India, known as the 'Dauntela'
and 'Mukna' (as fully cited l^elow) had been observed and very clearly stated by Corse as early as 1799 and
 :

1314 OSBORN: THE PROBOSCIDEA

accepted and amplified by Geoffrey St. Hilaire and Frederic Cuvier in 1825 (also cited below). Falconer .states

(1868, Vol. II, p. 257):

Hut even ill tlic Western India, at the extreme northern limit of the species at the i)resent clay, the
Sal forests of Nortli
cliffereiicc of slciidcr-built is well known, being expressed by Corse, for the Bengal \ariety, under the
and squat-built Elephants
designation of 'mirghi.' or Cervine [merghee, slender built, of Corse] for the former, and 'Koomarea' [koomnirah, deep bodied,
squat built, of Corse] for the latter, or when the characters are combined 'Sunkarcab.'

Falconer and Cautley accordingly gave geographic varietal names to the.se two very distinct continental

forms of the Indian elephant, known among the natives as 'Damitela' and 'Mukna,' and published in his plates
the following designations of these varieties

In 1847: Elephas indicus (Dauntela var.), the Dauntela variety = deep bodied, koomareah of Corse].
[

Elephns indinm (Mukna var.), the Mukna variety [=slender built, merghee of Corse].

Temminck designated the Sumatran elephant imder the following specific name and held that by its verte-

bral and other characters it was thoroughly entitled to specific distinction from the Indian elephant, a point
disputed by Falconer in 1868:

In 1847 : Elephas sumatranus, the elephant of Sumatra.

Eleph.\s indicus sumatra.nus, the Scmatkan Subspecies

Fig. 1 168. Pair of young elephants from Sumatra {Elephas swnatranus) Fig. 1169. The Sumatran elephant, apparently a female, living in the
captive (1921) in the ZooloRic.il Park of Wa.shinRton. Photograph kindly .Amsterdam Zoological Gardens, August, 1913. After a photograph kindly
pre.sented by Secretary Charles D. Walcott. presented by Mr. Draham Renshaw, inscribed "Elephas ijuiicxis sumaireTms,
(Broili, letter, .August 12, 1929): In the Stnte Zodlojiiical Museum of .\msterdani Zoological (iardens, Augu.st, 1913."
Munich an adult nioiuited specimen of Elfplm.f siiindlrniiux 'remmiiick,
there is (Letter of transmittal, November 21, •The Sumatran rlcjilianl
1923):
the skeleton of which is not mounted hut thi' skull is on exhibition. Head wa,sremarkable for the les.sellation of you see it was not a large
the hide; as
Preparalor Kustliardt of th<' zoological collection writes (.\ugust 10, 1929) animal .Buttikofer
. . in the Guide to the Rotterdam Zoo figured another
mounted specimen came from Sumatra in 1907; that the young
that the adult .Sumatran ele))hant."
mounted specimen (six months of age) exhibits dark brown hair and red woolly
hair.
 THE ELEPHANTINE: HISTORICAL INTRODUCTION AND NOMENCLATURE 1315

3. LIVING SPECIFIC OR SUBSPECIFIC FORMS, CONTINENTAL AND INSULAR

(Continued more in detail on pages 1323 to 1333 of the present chapter).
Falconer (1863, p. 81), in addition to his analysis of the vertebral characters, discusses very fully the unity or
plurality of species among the existing Asiatic elephants, namely, types from Ceylon, Nepal, Bengal, and Sumatra,
and decides in favor of the unity of the species Elephas indicus. He points out (1868, Vol. II, p. 258) the geo-
graphic variations which distinguish not only the Ceylonese and the Bengalese elephants, but the effects of
climate and breeding in different regions, namely, Assam, Silhet, Chittagong, Tipperah, or Cuttack. He concludes
that the specific distinctions of Temminck and Schlegel founded upon external characters, as in the case of Elephas

sumatranus, completely fail. He also believes that E. sumatranus is barely distinguished by the characters of the
ridge laminae. He points out (p. 260) that while the typical ridge formula of E. indicus is:

Dp 21 Dp3f Dp4H MIH M 2}| M 3|^

the last true molar, Mg, never shows less than 20 ridge-plates, commonly about 22 ridge-plates, but sometimes
in the lower jaw attaining as many as 27 ridge-plates. He remarks (p. 269) "the evidence in every aspect appears
to fail in showing that the Elephant of Ceylon [E. indicus ceylanicus] and of Sumatra [E. sumatranus] is of a species

distinct from the Continental Indian form [E. indicus]. Again (p. 270) he observes: "The result of this range of

observation, combined with long osteological study, has been to establish the conviction in my mind that there is
but a single species of Asiatic Elephant at present known, modified, doubtless, according to his more northern or

southern habitat, but not to an extent exceeding that of a slight geographical variety." He admits that the
Ceylon elephants are occasionally imported into Bengal but does not admit, as claimed by Schlegel, that this is an
explanation of the variation in the vertebral and rib formulae of the Indian elephants. Falconer (1863), while
inclined to accept Darwin's theory of the evolution of species, does not recognize the very great importance of
geographic isolation and of insulation in causing real geographic variation and subspecific and varietal evolution
both in external and internal characters. He is incUned to maintain the Linnsean idea of species and to oppose the
more modern idea of geographic subspecies and geologic ascending mutations.

Osborn believes that the wide variations in cranial and vertebral characters as well as in dental and dermal
characters and in the shape of the external ear support the subdivision of Elephas indicus into at least four out of
the large number (12) of geographic varieties or subspecific forms successively named, as follows:

COLLECTIVE SPECIES ELEPHAS INDICUS LINN/EUS, 1754

Synonyms:
Elcphaii }naximus Linnaeus, 1758; Elephas anialicus Blumenbach, 1797; Elephantun indicus (Cuvier), 1801.
Varieties and subspecies:
Elephas indicus Isodaclylus Hodgson, 1841 (named, without definition). Nepal, Tarai, N. India.
Elephas indicus Heterodaclylus Hodgson, 1841 (named, without definition). Nepal, Tarai, N. India.
Elephas indicus ceylanicus dc Blainville, 1845 (figure and plate description), the Ceylonese variety.
Elephas indicus bengaletisis de Blainville, 1845 (figure and plate description), the Bengalese variety.
Elephas indicus var. Dauntela Falc. and Caut., 1847 (figure and plate description, without subspecific name). Compare
Elephas indicus bengalensis de Blainville.
Elephas indicus var. Mukna Falc. and Caut., 1847 (figure and plate description, without subspecific name). Compare
Elephas indicus ceylanicus de Blainville.
Elephas sumatranus Temminck, 1847. The Sumatran variety.
Elephas maximus zeylanirus (Lydekkcr), 1907, 1916. Identical with Elephas indicus ceylanicus de Blainville, 1845.
Elephas maximus maximus (Lydekker), 1916. Compare Elephas
indicus bengalensis de Blainville, 1845.
Elephas maximus sumatranus (Lydekker), 1900, 1916. Identical with Elephas indicus sumatranus Temminck.
Elephas maximus hirsutus Lydekker, 1914, 1916 { = Elephas indicus hirsutus). Malay variety.
Elephas indicus Buski Matsuraoto, 1927 ( = 1 Palxoloiodon buski).
 .

E irrmnEus
. ciETriLjirsninMS

"*Siij5^ -

E.-inMCUS BErtEAJLEPiSIS

De Blainville's Types of Male Elei-has indicus ckylanicus and E. indicus bengalensis

Bengal [ef. Dauntela var.) ra.es of the
[of. Mukna var.) and tho broader-skulled, largo-tuskod
Fig. 1170. Th,. iiarrowcr-skulU-d, slcnder-tu.«kod Ceylon
Indian elephant. After dc Blainville, 1839-1864 [1845], PI. in, one-eiglith natural size.
,, , . y^ Premaxil-
, v ,„vil
(liypsieepluilie); orbits relatively .-loser to eondyl.-s (..yrto<-,e,,hahe).
Subsi>eeies Elepha^ indicus cfylanicus. .Summit of eranium more ,.o.nted
the Museum National d'H.sto.re Naturelle. Pans; Doetor
lary sockets longer; mandible shallower and more slender. The original of
this eranium is not in
is not, tl,e .same as the f.'malc erannun m the 1
aris
Anthony reports (letter, July 16, 1930) that he does not know where this fine cranium is presi-rved; it
Museum shown in figure 1172 of the present Memoir. , u v v .„.vil
, , ,
to orbits relatively less cyrtocephalic. Pren.aMi-
Subspecies Elcphas indicus bfnijakmis. O.'cipitofrontal crestmore rounded, less elevated; condyka
laries shorter, tusks broader; inframaxillarics more powerful. Mus. d'Hist. Nat. A. 8016.

1316
 THE ELEPHANTINE: HISTORICAL INTRODUCTION AND NOMENCLATURE 1317

The discovery and naming of these geographic varieties and subspecies are recited above and continued
more in detail on pages 1327 to 1334 of the present Memoir.

Since all the above varietal and subspecific

names were given prior to the adoption of the modern system of
nomenclature, we await monographic research and comparison of the living varieties and subspecies of Elephas;
meanwhile those that seem best entitled to consideration at present are the following:

1845 Elephas indicus ccylanicus de Blainville

Syn. Elephas maximus zeylanicus hydekkcr, 1916
:

Compare Elephas indicus var. Miikna' Falc. and Caut., 1847

1845 Elephas indicus bengalensis de Blainville
Syn. : Elephas niaximus maximus Lydckker, 1916
Compare Elephas indicus var. Dauntela' Falc. and Caut., 1847
1847 Elephas [indicus] sumatranus Temminck
Syn.: Elephas maximus sumatranus hydckkcr, 1916
1916 Elephas maximus hirsulus Lydekker [
= E. ijidicus hirsulus]-

Fig. 1172. Elephas indicus bengalensis Type and E. indicus ceylanicus

Referred. Paris Museum
After special photographs kindly taken under the direction of Dr. R. Anthony
Fig. 1171. The broader, large-tusked Dauntela and narrower, small-
(Ijcft) Full facial view of the cranium of Elephas indicus bengalensis
tusked MuKNA varieties of Elephas indicus. After Falconer and Cautlcy,
1846 [1847], PI. XLii, figs, xxiii.a (Davmtela) and xxiii.B (Mukna), onc- (Mus. d'Hist. Nat. A.8016) described by Dr. R. Anthony (Paris, letter, July
sixt^enth natural size. Falconer's distinctions between the Dauntela and 16, 1930) as follows: Cranium of a male Bengal elephant, .same as that

Mukna varieties are still more clearly shown in figure 1204 below. figured in profile by de Blainville in his "Osteographie," 1839-1864, PI. iii
[Fig. 170 of the i)resent Memoir]. Found along the Ganges River by Duvaucel
1
Dauntela var. Cranium relatively broader (brachycephalic), and less
in 1824. In this cranium the left intermaxillary is broken away and missing.
elevated. Premaxillariessliorter, with deep median excavation, highly charac-
teristic of the Elephantine?. Compare flypselephas hysudricus.
The measurement h to h' is 92.5 mm. or 3 ft. % in. Reproduced one-sixteenth
natural size.
Mukna var. Cranium relatively narrower in proportion to its height;
(Right) Full facial view of the cranium of Elephas indicus ceylanicus
occiput more elevated, anterior nares narrower. Premaxillaries more elongate;
(Mus. d'Hist. Nat. A. 8014). Of this specimen Doctor Anthony writes (Paris,
tusks less divergent.
July 16, 1930): A female born in Ceylon, brought from Holland, lived in the
Jardin des Plantes 1797-1816; di.ssected by Cuvier and Rou.sseau; figured by
de Blainville in his "Ost6ographie," 1839-1864, Pis. i and ii. Total facial
The resemblance of Elephas indicus ceylanicus de
height of cranium h to h^ 97.5 mm. or 3 ft. 2\ in. Reproduced one-sixteenth
Blain. to the E. indicus var. Mukna Falc. and Caut. is natural size.

afTorded by the drawings of the cranium (side view) by de Blainville and Falconer and Cautley figured fully below.
A fine front view of the cranium of bengalensis is compared with the front aspect of the cranium of ceylanicus
(Fig. 1172), as described by Doctor Anthony in the legend above.

'[Referring to the use of the terms "mukna" and "dauntelah," Mr. Dunbar Brander commented (November, 1930) that he did not like a classification
which used these two native terms, since a tuskless elephant (mukna) was apt to appear in any region and the term had no diagnostic value for any character
other than tusklessness. —
Editor.)
"[See footnote under description of hirsulus on page 1332 below. — Editor.)
1318 OSBORN: THE PROROSCIDEA

The resemblance of the Elephas indicus bengalensis de Bhiin. to the E. indicuH var. DaunteUi Falc. and (
'aut.

is also supported by a comjjarison of the crania and figures (lateral view). Further conijiarison should 1)(> made
of the front views of the crania in these two subspecies and varieties.

Fig. 1173. Geographic di.«tributior\ of tho principal species and subspecies (living and extinct) of Elephas, also of Htjpseh'ph(vi and Platiiephas, accord-
ing to the numbers given in the list on the op])Osite page. The white dots within the black areas represent the approximate localities where tlie types were
discovered; these dots each carry a number in a circle, representing the chronologic sequence of type description. The + locates some of the principal
referred specimens.

4. FOSSIL FORMS MORE OR LESS CLOSELY RELATED TO ELEPHAS INDICUS

It is remarkable that no fossil Pliocene ancestors of the recent Indian elephant have as yet been discovered.
While the cotypes of Elephas hysudricus, attributed by Pilgrim to the Boulder Conglomerate zone, are in i)art

recorded as found near Moginand, Simla Hills (Fig. 1196), none of the specimens personally collected by Barninn
Brown in 1922 was actually found in the Boulder Conglomerate zone; they all appear as if washed or erodetl out

of this zone with more or less adherent gravel or concretionary material; they were found in hollows or ravines
cut into tlic underlying Piiijor liorizon.

184G 1184")]. Elephas hysudricus Falconer and Cautley, of tlie Lower Pleistocene, found 'below the con-
glomerates' of India, shows few resemljlances in the cranium to the E. indicus (Dauntela var.) of Falconer and
Cautley, and no very marked resemblances in the grinding teeth ; this species appears to be unique.

19()<S. Elephas hysudrindicus. Dul)ois descril)ed this species as theoretically intermediate betw(!en Elephas
hysudricus and E. indicus, but Stremme rightly regards this animal as more nearly related to 'E. namadicus.'
Osborn treats this animal (Chap XIX) as belonging to Palxoloxodon, namely, Palxoloxodon hysudrindicus.

1927. Elephas indicus Buski. Matsinnoto, having become convinced of the occurrence in Japan of the true

Asiatic elephant in the fossil state (first suggested by Busk in 1868), described a first superior molar from the
Ninohc District as the type of this subspecies. Osborn regards this tooth as referable to Palscoloxodon.

1929. Elephas plutycephalus. Osborn in Memoir redescribes this si)ecies from the Upper
the present
Pliocene or Lower Pleistocene of India, recorded by Barnum Brown as from 'below the conglomerates,' a level
 :

THE ELEPHANTINE: HISTORICAL INTRODUCTION AND NOMENCLATURE 1319

higher than that in which Archidiskodon planifrons occurs and lower than that in whicli E. {Hijpiiclephas) occurs.
As shown in figure 1 174, this is a very ancient and primitive animal. It is now made the genotyi)e of Platelephns.

1929. ElephuH plalycephalua (tngustidens. Based upon a single second sui)erior molar, r.M'-, originally
mistaken by Osborn for a third inferior molar, which now proves to belong to Elephas [
= Hypselephas] hysudricus.

5. NAMES OF SPECIES AND SUBSPECIES OF THE SUBFAMILY ELEPHANTINE

IN ORDER OF DESCRIPTION
See Figure 1173

A summary of the names applied to the above living and fossil forms actually or apparently related to
Elephas is as follows

LIVING SPECIES

Collective
1320 OSBORN: THE PROBOSCIDEA

Fio. 1174. Restoiiations of Platelethas i-latycephalus, Hypselephas hysudiucus, and Elephas indicus. April, I'JIJO.

All three heads reduced to a uniform one twenty-fourth scale

(l.ifl) I'UtUirplmx i>l(ili/C(i)lialus drawn directly (Center) llypsdephas hysudricus profile (Right) Elephas indicus, a middle-aged female.
on the cranium, slightly crushed downwards; ears drawn directly on I''alconer's outline of the cranium Observe the relatively liigli po.>iition of the rye
of supiioscd primitive size, occipitofrontal line (Fig. 1204). Observe the extremely low position of midway between the lip and the summit of the
drawn directly on top of cranium. Observe the the eye, corresponding with the oeei]>ital concavity, cranium; also the lack of the occipitofrontal crest
eye in normal position. namely, quite close to the preniaxillary socket. and the presence of tlic prominent occipitofrontal
The ear is given a slender elephantine outline, as in convexity and nuiscular ridge for the supratempo-
H. imlicus. The small tusks correspond with the rals.

relatively small alveoli in this specimen.

II. CHARACTERS OF THE SUBFAMILY ELEPHANTIN/E AND INCLUDED

GENERA AND SPECIES
Superfamily: ELEPHANTOIDEA Osborn, 1921
Family: ELEPHANTID^ Gray, 1821

Subfamily: ELEPHANT I N>E Osborn, 1910

Original reference: "The Age of Mammals in Europe, Asia and North America," Osborn (1910.346, p. 558); also Osborn,

1918.468, p. 135.
Compare: Elephantina Bonaparte, 1838; Elephantini Wingc, 1906.

Subfamily Characters.— (1) Skull brachy cephalic, progressive from platycephalic (Elephas [Platele-
phas] platycephalus) to hypsicephaUc (E. indicus). (2) Jaws, including rostrum, progressively abbrevi-
ating and deepening, comi^letely brevirostral in recent time (E. indicus). (3) Premaxillaries and tusk
alveoli relatively narrow, in contrast to the broad rostrum of the Loxodontinse. (4) Upper tusks upturned,
out-turned, finally inturned, attaining moderate length, as compared with Loxodonta. (5) Grinding
teeth progressively hypsodont. Ridge-plates increasingly compressed and enamel finely folded (E.
indicus). (6) Ridge-plate formula of third molar multiplying from 3 '-^' {platycephahis), to i,.'rgTr9 M
(hysudricus), to ^r^
(indicus). (7) Inferior ridge-plates exceeding superior ridge-plates in number
(E. indicus), in contrast to the Mammontinae (Parelephas, M3'il-.li). (8) Including the jH-ogressive
E. indicus, readily distinguished by its cranial axes and contours from all the known members of the
Loxodontinse (Chap. XIX), and Mammontinae (Chaps. XVI-XVIII). (9) Provisionally including three
extinct specific if not generic phyla, represented by E. [Plntelephns] platycephalus, E. [Hypselephas]
hysudricus, and E. indicus, not constituting a single ascending phylum but probably polyphyletic.

Doubtful Relationships. —The above definition of the subfamily Elephantinpp is continued from Chapter
11, i)p. 11-13 and 16, also from C'hapter XV, pp. 913, 915, and 918. It is provisional and heterogeneous, because

Elephas hysudricus Falc. and Caut. and E. platycephalus Osborn are at present known by cranial characters only

and appear to represent generic or subgeneric phyla distinct from the true Elephas. There are three separate
lines of cranial adaptation in the fossil and living species embraced within the subfamily Elephantina?. This is in
 THE ELEPHANTINE: HISTORICAL INTRODUCTION AND NOMENCLATURE 1321

contrast to the cranial uniformity which prevails in the Mammontinse and the clearly defined phylogenetic suc-
cession observable in the three included genera {Archidiskodon, Parelephas, Mammonteus). These three generic or
subgeneric lines may be compared and distinguished phyletically as follows:

Phylum I Phylum II Phylum III

Platelephas Osborn Hypselephas Osborn Elephas Linnaeus

Typified by Elephas -plalycephnhta. Typified by Elephas hysudricus and Typified by Elephas indicus. Extreme
Upper Plioconp or Lower Pleistocene. E. platyrephalus anguslidens. Lower Pleis- Upper Pleistocene and Recent.
tocene.
Cranium relatively elongate, dolicho- Cranium elevated or hypsice])halic, Cranium bathycephalic, cyrtocepahlic,
cephalic, and platycephalic, occipital condyles well raised abov(> grinding sur- hypsicephalic, occipitofrontal dome more
condyles somewhat above level of grind- face of molars, occiput elevated with or less rounded, not acute, with expanding
ing surface of molars. broadly transverse frontal crest, front- diploe, frontals gently concave.
als deeply concave.
Premaxillaries greatly elongated in Premaxillaries relatively narrow or Premaxillaries relatively narrow, sub-
front of molars, .somewhat divergent; laterally compressed, not deeply extend(>d vertical, borders relatively close to front
tusks unknown. b(>low front of molars; tusks relatively of molars; tusks relatively straight, in-
straight, incurved, somewhat divergent curved, used in feeding and uprooting
at base; rostrum of lowci- jaw elongate, habits, unlike the adult Mammonteus.
prominent, ramus shallow. Jaw with extremely abbreviated rostnmi.
Orbits large, elevated, near frontal Orbits large, depressed, near maxillary Orbits large, elevated.
profile. rostrum.
Grinding teeth relatively low, ridge- Grinding teeth with ridge-plates con- Grinding teeth with finely plicated
plates directly transverse, as in Elephas, vexo-concave, reversed above antl below, enamel, not rising to the extreme hyp-
no rudiment of 'lo.xodont sinus.' Ritlgc- trace of a median 'lo.xodont sinus,' of le.ss sodonty of Parelephas, Archidiskodon, or
plate formula: h(>ight but otherwise as in Elephas. Mammonteus. Ridge-plates multiplying:
Ridge-plate formula:
M3i^ M 3 —2i—
M 3 17-18-19
Habits unknown, ])robably like those
Habits unknown, but from the tasks Known habits chiefly browsing, crush-
of Elephafi indicus.
and grinding teeth probably similar to ing of coarse leafage and herbage,
those of Elephas indieus. secondarily grazing.
Very primitive in cranial structure and Progressive in cranial structure and in Highly jjrogressive and distinctive in
in the limited number of ridge-plates.
the somewhat largei' number of ridge- cranial structure, with maximum number
plates, although the molar crowns are of plicated ridge-plates.
still low.

As the Pleistocene is now recognized as extending over approximately a million years of geologic time,
I.

Platelephas platycephalus was nearly contemporary with Archidiskodon planifrons and with Hesperoloxodon
antiquus of far western Europe and was separated by an enormous interval of geologic time from the recent
Elephas indicus, but despite its very primitive cranial structure it would be rash to disbar it entirely from the
ancestry of the modern E. indicus.

II. Similarly Hypselephas hysudricus of the Lower Pleistocene is perhaps a million years older than Elephas
indicus but its cranium does not appear to be evolving in the direction of that of E. indicus.

III. Therefore the recent species Elephas indicus of southeastern Asia is left without a known ancestral form,
just as its living contemporary Loxodonta africana is left without a known ancestral form.' In both cases this
ancestral time gap will probably be filled by discoveries in northerly unexplored regions of Eurasia and of Africa
respectively.

In view, therefore, of the uncertainty regarding the ancestral relationships of Platelephas platycephalus and
of Hypselephas hysudricus to Elephas indicus, we may at present define the genus Elephas from the characters
preserved in the genotypic species E. indicus and in its geographic varieties and subspecies, as follows.
'[See Loxodonta -prima Dart, 1929, also L. ajricana var. obliqua Dart, 1929, Chai>ter XIX, pp. 1287, 1288 of the present Memoir. — Editor.]
1322 OSBORN: THE PROROSriDEA

A Herd ok Wild Bengal Elephants (Elephas indicuo benqalbnsis)

Fig. 1175. .\ herd of wild elephants in a bamboo jungle of Mysore. Photograph by Mr. Wiehle of Bangalore, originally jjublished in The Illustrated
London Xews of Januar\- 8, 1910, and reproduced by courtesy of Mr. Bruce S. Ingram, Editor. The title of this remarkable jihotograph is ''Quietly dreaming
in the jungle." The giant bull to the left of the center illustrates the extreme prominence of the occipitofrontal convexities.

Superfamily: ELEPHANTOIDEA Osborn, 1921

Family: ELEPHANTID^ Gray, 1821
Subfamily: Elephantine Osborn, 1910
Genus: ELEPHAS Linn^us, 1735-1758
Pleistocene and Kecent of Siani and India: Ceylon, Bengal, Burma, and Sumatra.
Syn.: Euelephas Falconer (in part), 1857; Elephantus Cuvier and E. (ieoffroy, 1795; Polydiskodon Pohlig (in part), 1888.
Genotypic species: Elephas indicus of Ceylon.
Geographic varieties and subspecies: Elephas indicus ceylanicus, Elephas indicus hengalensis, Elephas indicus siimatranns,
Elephas indicus hirsutus.

—
Generic Characters. (Linn.TUs, 1758, p. 33): "Maximum qnadmpes. Oculi parvi. Dentes
Laniarii nuperiores exserti (Ebur). Aures amplissimx, pendulae, dentatx; act. angl. 211 p. 1051. Cutis .

crassissima callosa, Msuninx 2 juxta pectus. JJngiies in apicibus loborum pedum. Genua fI exilia. Col-
.

lum breve.
Proboscis longissima, extensilis, acute odorans, loco manus ipsi inserviens; ea cibum potumque hauril,
hoHtemque pellit; ea prxcisa occidiiur; murem metuit sub somno ob tracheae insertionem. Retro coil &
mingit. Portal domos, rectore collo insidente; in bello armalur falcibus; vulnusculo inter axin A atlantem
furiosus occidiiur; cxterum priidens, docilis.
(Osborn, 1924) Genotypic species Elephas indicus ( = maximus). Cranium widely distinct in form
:

from that of Loxodonta aj'ricana, of Archidiskodon, of Parelephas, of Mammonleus, namely, relatively

acrocephalic, hypsicephalic, moderately bathycephalic. Frontals gently concave, occiput decidedly
convex (male), fronto-occipital crest uniformly convex, moderately elevated (female), i.e., acrocephalic.
Molars of intermediate breadtli, absence of 'loxodont sinus' in Elephas indicus, faintly indicated in Hyps-
elephas hysudricus; moderately compres.sed enamel ridges of intermediate thickness, extremely crimped
or sinuous in E. indicus; more ridge-plates in inferior than in superior molars; ridge-plate formula of
E. indicus, M
3 ,;.\.7- of ^- hysudricus (?) ref., 3 rrTrsrri- M
Dorsal or rib-bearing vertebra- 19-20.
Digits of manus with five horny sheaths or nails; digits of pes with either four or five horny sheaths or
nails. Digital formula: Manus 5, pes 4-5.

As shown in the cranial and dental sections of the present Memoir, the genus Elephas may be clearly defined
as readily distinguishable both in cranial and dental characters not only from Loxodonta but from Archidiskodon,
Parelephas, and Mammonleus.
 THE ELEPHANTINtE: ELEPHAS 1323

Ridge-Plate Comparison with Parelephas. — It is a striking fact that Falconer's ridge fonnulu {Elephas
indicus), namely, M 3 u/.V;, assigns a higher number of ridge-plates to the last inferior molar than to the last

superior molar; this reverses the condition observed in Parelephaa progressus in which the ridge formula is M 3.^1-7^5.

If this difference proves to be constant, it affords additional means of distinguishing species of Parelephas from
species of Elephas, as follows: In Parelephas there are more ridge-plates in the superior molars than in the inferior,
whereas in Elephas indicus there are more ridge-plates in the inferior molars than in the supprior.

1. SYSTEMATIC DESCRIPTION OF SPECIES OF ELEPHAS

ELEPHAS INDICUS Linnaeus, 1 735-1 754, Collective Species'
Figures 794, 79()-800, 802-814, 810, 893, 912, 992, 99.",, 1012-1014, 1081, 1082, 1081, 1109, U12, 1123, 111)3, 1174, U7d-1178, 1190-1194, 1234, 1243, 1244

Syn: Elephas manmus Linn., 1758; Elephas asiaticus Blnmonbach, 1797; Elephanins indicuti ("uvior, 1801 (in f'uvior
and l-aoept'dc); Elephd.s (jigas Pony, 1811; E. (Euelephas) //;f//c(^s Falconer, 1857.

The history, nomenclature, and general characters of this collective species are fully set forth in the present

chapter, pages 1308 to 1321 above.

shown that Linnaeus (1754, p. 11

It is —see facsimile, p. 1309 above) in naming £J/ep/(a.s indicus, while depend-
ing upon Ray, Seba, and others, was very indefinite as to the type locality, mentioning both the continent of Asia
and the island of Ceylon. When he changed the specific name to 'maximus' (1758, p. 33) he may have had in

mind the African elephant, of which he had a foetal specimen, but he subsequently states "Habitat in Zeylonae."
Consequently we are inclined to the technical opinion that Linnaeus' tijpe, both of his 1754 description of Elephas
indicus and of his 1758 description of Elephas maximus, was the Ceylon animal subsequently named by de Blain-
ville (1845) Elephas indicus ceylanicus. If this historic interpretation is correct, E. indicus ceylanicus is not only
the type of the collective species Elephas indic^is, but it is genotypic of the genus Elephas itself.

The main descriptions, figures, measurements, and characterizations by all other authors have been of the
better known continental variety, to which de Blainville (1845) applied the subspecific name Elephas indicus
bengalensis. It becomes necessary, therefore, to review the observations and history of opinion on the specific

forms of the collective species 'Elephas indicusJ

[Elephas gigas Perry, 1811 ( = synonym of E. indicus, fide Colbert). — This species is described (p. li)

and figured (plate opiiosite that page) in George Perry's article of 1811 in the "Arcana," a portion of which
descrii)tion is cited herewith: ''Natural Order — Mammalia. Species — Elephas gigas. Generic Character —No
fore-teeth in either jaw; the tusks of the upper are elongated and projecting, none in the lower; the proboscis
or trunk very long and prehensile; the body armed with a very thick skin, covered with a few scattered hairs."

"The Elephant may justly be considered as the largest and strongest animal at present known, and is plenti-

fully found in a wild state in the extensive regions of Africa and Asia."

"There is also found a second and different species, which is said to reside in the kingdom of Thibet, and
being much smaller and of an opposite form, is to be considered as a separate animal from the above, under the
title or Name of the Elephas socotrus [indeterminable from the description]." Neither species determined by
the present author. — Editor.]
'Ray, in his description of Elephas (1693, pp. 131, 132), speaks of the elephant of Sumatra in reference to its weight, and to the skeleton in Florence in ref-
erence to tlie number of ribs, characters of the tusks, etc.

Seba (1734, ]). 175, PI. CXI) gives a very full description of the Foetus Elephantis Africani ineditus, wldch Liinnberg refers to as the type. In this
early stage of zoology no one dreamed of selecting any jiarticular si)ecimen and designating it as the (y/if.
 —

1324 OSBORN: THE PROBOSCIDEA

FALCONER DESCRIBES THE CONTINENTAL BREEDS OF BENGAL AND ASSAM

The characters of those continental breeds of Indian elephant, Falconer considers and repeatedly states that the ridge
were more profoundly and thoroughly studied by Falconer than formula of Elephas [Mammonteus] primigenius is closely similar to
by any pre\aous or subseciuent writer. Falconer's discussion of that of his subgeneric group E. (Eueleph.) indicus (cf. the ridge-
the vertebral formulae as well as of the varietal and subspecific plate fornudse of the ascending mutations of M. primigenius above
characters is fully given above in the present chapter; he was (p. 1138), namely, M 3 H to M3( '(?)2 7/-

not inclined to admit the constancy of the subspecific characters

claimed by Schlegel, Temminck, and other observers. Conse-
quently the following citations from Falconer refer to Elephafi
indicus as a 'Linnsean collective species.'
Falconer (1868), Dental Characters. — In comparison --,.^J!
with the grinding teeth of Elephas [Hypselephas] hysudricus (Figs.
1197, 1198, of this Memoir), Falconer's description of the .superior
grinding teeth of Elephas indicus is as follows (Falconer, 1868, I, p.

78) : "The existing Asiatic elephant, E. Indicus, furnishes the next

modification represented in this plate. Fig. 2, PI. v. (or fig. 2a,

PI. I., F.A.S.), shows a section of the penultimate upper molar of

this species. The gradual attenuation of the plates, successively
exhibited from E. insignis to E. Hysudricus, is here carried to
eighteen of these divisions being comprised within the space
exce.ss, ''
'/f
occupied by about nine in the equivalent tooth of the African
species. They are produced vertically in the same proportion, the
height of the middle plate being about three-fourths of the entire Fig. 1170. (Falconer, 1868, Vol. I, p. 422): "Elephas indicus. Vortical
Itmgth of the tooth ; they, in fact, represent parallel perpendicular section of unusually large specimen of last lower molar of an Indian Elephant
lamellae, of nearly luiiform thickness from the base to the apex, from Assam, in India House [London] collection. The entire length of the
interstratified with layers of cement of nearly thesame thickness. crown is about fifteen inches, and it includes as many as twenty-seven ridge.s,

The layer of enamel is attenuated into a thin transversely un- of which the anterior thirteen are more or less abraded. The first five or six

dulated brittle plate, the .surface of which is deeply wrinkled with ridges incline a little forwards, while the posterior ridges incline so much in an
opposite direction, that the hindermost are nearly horizontal, producing the
striae, for the firm cohesion of the cement. The general character of
flabelliform character that so readily distinguishes in most instances the last
the section a pectinated arrangement of the lobes like the teeth
is
from the pi^nultimate lower molar. The same disposition and proijortions
of a comb, which contrasts strongly with the chevron-formed ridges
of the dental substances are observed as in the upper grinder." After Falconer
of E. insignis, and the cuneiform plates of E. planifrons. The mass
and Cautley, 18t6 [184.51, PI. i, fig. 26. One-fourth natural size.
of ivoiy at the base of the tooth is much thinner than in the cor-
responding molar of E. Hysudricus, bearing but a very slender
jiroportion to the height of the tooth; and numerous small and
distinct fangs are given off fromThis tooth had
its inferior edge. Inferior Molar, M3, Assam Breed. — (Falconer, 1846, p.

been some time in use, the anterior part of the crown being worn off 43) : "Fig. 2b represents the section of a very fine specimen of the

as far as the ninth plate. The plane of the truncated portion is very last inferior molar of the existing Indian Elephant of Assam, from
oblique, being inclined n(!arly at a right angle to the coronal surface the collection at the India House. It is an uiuisually large speci-

of the unworn jjortion. This specimen is 8.2 inches in length." men, showing as many as twenty-.seven plates, the anterior twelve
HiDGE Formula of Elephas indicus (Falconer, 1868). of which have been in use. Precisely the same disposition of the

Falconer appears to have finally based the ridge formula of M3 on dental substances is observed in this case as in the upper grinder,

an Indian elephant from Assam (Falconer, 1846, p. 43 [1845, fig. 2/)], and they are developed in the same relative proportions. The
probable, therefore, that Falconer's vertical height of the plates is still greater than in the correspond-
with 27 ridge-plates); it is

ob.servations on the grinding teeth refer chiefly to the northern ing lower molar of E. hysudricus. The upper surface is concave,

race, Elephas indicus bengalensis (cf. Falconer, "Palaeontological and the luider very convex. The anterior [)lates are nearly vertical,

Memoirs," Vol. I, p. 422, PI. i) : Elephas indicus. Plate i, fig. 2b, while the posterior gradually slope backwards till they become

Indian elephant from A.ssam. M3 with 27 ridges, k'ngth of crown almost horizontal in the hindmost portion, with a corresponding
The formula of Falconer (1868) consistent gradual diminution in their height. This is a mechanical arrange-
about 15 in. final is

with that of Falconer (1863), namely: E. indicus of Assam, ment ari.sing from the contracted diameter of the posterior part of
developed,
M 3 2i- Thus Falconer corrected his earliest ridge formula (1857, the dental canal, in which the back part of the tooth is

p. 315) of Elephas indicus and substituted (1863, p. 65) the follow- close under the condyle, the plates being disposed so as to occupy
ing collective ridge formula of E. (Euelephas) indicus: the least vc^rtical space. The basal mass of ivory between the plates
and the fangs is reduced to a small quantity. This tooth measures
Dp 21 Dp 3f Dp 4f^ M Ifl M 2 H M 3^,^ lHYi inches long in a straight line."
 " :

THE ELEPHANTINE.: ELEPHAS 1325

RIDGE FORMULA OF ASSAM, CEVLON, AND SUMATRAN FORMS instances prove, so far as they go, that the ridge-formula is the same
Falconer "Palseontological Memoirs," 1868, Vol. II, pp. 256, 260. in the Ceylon and Sumatran form as in the Indian." He also
contests the alleged differences in the width of the enamel bands
Since the posterior ridge-plates of M', M3
develop very late in
and shape of the discs, stating as regards Elephas sumatranus that
life, the ridge-plate formula is difficult to determine. Schlegel,
the supposed width of the enamel bands is due to the obliquity of
Temminck, and other authorities contend that the very high ridge-
the section (cf. p. 262).
plate formula of the Assam breed, M
3 g/.^,, is not obtained in the
Falconer's entire discu.ssion (op. cit., pp. 2.56-270) of the unity
insular breeds of Ceylon and Sumatra, to which they assign not
or plurality of .species confirms his conviction that there is but a
only a lesser ridge-plate formula but a greater thickness of the
single species of Asiatic elephant, modified only to the extent of
enamel ridge-plates, as quoted by Falconer (1868, II, pp. 256, 260)
a slight geographical variety.
"(B.) Molar teeth. —
Ribbons (discs of wear) in form like those
of the Indian species, i.e. the enamel-plates highly crimped, parallel,
and freefrom the rhomb-shaped expansion of the African Elephant;
but the ribbons wider (in the direction of the long axis), and con-
fiequently less numerous than in the Indian species; the difference

being in the ratio of 3 or 4: 1 in the Sumatran, and 4 or 6: 1 in the

Continental Indian form (Schlegel in Temminck). Ribbons of
enamel nearly [or] quite as wide as in the African Elephant.
(C. L. Buonaparte.)"
(Schlegel in Falconer, op. cit., p. 260) " 'The laminse of the
:

teeth afford another distinction which however is less apparent to

the eye than that taken from the number of vertebrae. These Fig. 238. — Coupe d'une derniere molaire inferieure d' Elephas indicus,
laminiE, oi' bands, in E. Sumatranus, are wider (or if one may so au 1/4 de grandeur.

say, broader in the direction of the long axis of the teeth) than in
Fig. 1178. Section of a partly worn third inferior molar of Elephas indicus.
E. Indirus.'
After Gaudry, 1878, p. 179, fig. 238, one-fourth natural size.
Observe that six or more anterior ritlge-plates have been worn off, the
total number being twenty-four or more.

DISTINCTIONS BETWEEN THE VARIETIES OF THE INDIAN

ELEPHANT, CORSE (1799), GEOFFROY SAINT-HILAIRE AND
FREDERIC CUVIER (182.5), TEMMINCK (1847), DE BLAINVILLE
(1839-1864), AND FALCONER (1867, 1868)

(Continued from p. 1313 of the present chapter)

Fig. 1177. Type r.M' of Elephas asiaticus Blumenbaoh, 1797 (Bhimenbach is not clear from the following descriptions by Corse
It
1797.2, No. 19): "B. vom Asiatischen Elephanten, . . . Nur bilden jene beym (1799) and by Geoffroy and Cuvier (1825) in what part of India
Elephas asiaticus geschliingelte an beiden Enden paarweis zusammenlaufende
these two varieties occur; probably they refer to the continental
Linien; hingegen bej'm africanus raiitenforniige Leisten. Diese Zahne der
varieties of Bengal.
beiderlei Elei)hanten .sind nach Originalen im hiesigen acadcmi.sohpn Museum
gezeichnet." Inverted to show natural position of molar.
The derivation of the words mooknah, dauntelah, and pullung
Blumenbach's type figure is apparently a first superior molar of the right daunt, as given by Corse, is as follows: "Probably from rnookh,
side, r.M',outer side convex, inner side plane or slightly concave, exhibiting the mouth or face . . . Dauntelah signifies toothy; having large or
twelve ridge-plates, as in Falconer's typical ridge-plate formula of Elephas Pullung signifies a bed or cot, and daunt, teeth; and,
fine teeth. . .

indicus.
from the tusks projecting so regularly, and being a little curved
and elevated at the extremities, the natives suppose a man might
Falconer, after the examination of a very large quantity of lie on them at his ease, as on a bed [i.e., Pullung]."

materials in India and Europe, concludes that the ridge formula (Corse, 1799, p. 208) : "After premising these general observa-
runs thus {op. cit., p. 260): tions, I may here observe, that elephants have two tusks, in the
upper jaw only; but those in some of the females are so small as
Milk molars. True molars. not to appear beyond the lip, whilst in others they are almost as
20-2 4
large as in one variety of the male, named mooknah.
1 6,
2 0-2 4 1-2 71
The . . .

largest tusks, from which the best ivory is supplied, are taken from
This he regards as typical, stating (cf. p. 261) that neither that species of male named dauntelah, ... in con.sequence of his
Schlegel nor any of the other advocates of distinct specific ridge large tusks, and whose countenance, from this circumstance, is the
formulae have proved that either the Ceylon or the Sumatran most opposite, in appearance, to that of the mooknah; which, as
species shows a lesser number than 2 M
He concludes: "These I have just ob.served, is hardly to be distinguished, by his head,
1326 OSBORN: THE PROBOSCIDEA

Elephas indicus bengalensis

Fig. 1179. A faniou.s inclividiial known Hi.s tusks grew to sui-h a Icngtii that ho was \inablf to lie down and they were
as the 'giant tuskfr of Udiapur.'
shortened at the extremitie.s and encased in metal. After ]>hotograph by F. D. Fayrer, originally imblished in Asia Magazine, Vol. 29, June, 1929. Rei)rQduced
by courtesy of "Mondiale" through Asia Magazine. Seale approximately oiic-fifticth natural size.

from a female elephant. Though there is a material difference in apporte pas dans Lorsqu'on ne connait |)as le caractfere
le prix.

the appearance of a mooknah and a dauntelah, as well as in the d'un Elephant, Europ6ens aiment mieux I'acheter sans grandes
les
value of the tusks, yet, if they are of the same cast, (zal), size, and defenses, parce qu'il aura moins de moyens de nuire s'il se trouve
disposition, and perfect, that is, free from any defect or blemish, mediant mais les Iiidiens pr6fferent assez les individus a longues
:

there is scarcely any difference in their price. There must be

. . . defenses, pour s'exposer a tous les risques. Lorsque le bon naturel
five nails on each of his fore feet, and four on each of the hind ones, de I'animal est connu, les deux nations I'aiment mieux ave(! de
making eighteen in all; his head well set on, and carried rather grandes defenses. II y a une infinite de vari6t6s parmi les Daunte-
high. The arch or curve of his back rising gradually from the lahs, par rapport a la direction et a la courbure de leurs defenses.
shoulder to the middle, and thence descending to the insertion of Les plus estimfe sont ceux ou elles approchent le plus de la direc-
the tail; and all his joints firm and strong. There are several tion horizontale. Les princes indiens ont aussi un respect super-
other points, of less consequence, which are taken notice of by the stitieux pour les Dauntelahs qui n'ont qu'une defense, comme cela
natives as well as Europeans. The dauntelah is generally more arrive quelquefois."
daring, and less manageable, than the mooknah; for this reason, Osborn, 1930: From the facts and comparisons cited in the
until the temper and disposition of the two sj)ecies are ascertained, early jiart of the present chapter and given more in detail above,
Europeans will prefer the mooknah; but the natives, who are fond it appears ]jrobable that the continental Indian varieties, as ob-
of show, generally take their chance, and ])refer the dauntelah; served by Corse, Falconer, Geoffroy, and others, represent the
which, when known to be of a mild and gentle disposition, will descendants of two wild indigenous races or subspecies, including
always be preferred, both by Europeans and natives. The varieties the larger, more vigorous northern form (var. Dauntela) and the
between the mooknah and dauntelah are considerable, and for smaller, more slender southern form (var. Mukna), which, freely
these there are ai)proi)riate names, according as the form of the interbreeding, according to Corse, have given rise to a large
tusks varies from the projecting horizontal, but rather elevated, number of intermediate forms. It also appears probable, as sug-
curve of the pullung daunt ... of the perfect dauntelah, to the gested by .Sclileg(>l, that the smaller, more slender continental
nearly straight tusks of the mooknah, which i)()iiit directly dow?!- form is relatetl to the Elepha.s indicus ceylanicus of de Hlainville,
wards." whil(> the larger, more vigorous form is related to the E. indiru.'i

(Geoffroy Saint-Hilaire and F. Cuvier, 1825, j). 7, filejihant bengalensis of de Blainville. On further research it is possible
d'Asie): "Nous avons d^ja vu que les femelles des Indes n'ont that the.se two subsix'cies may be clearly defined not only by the
jamais que de trfes-courtes d6fen.ses: il y a des males qui n'en ont cranial characters described below, but by the vertebral fornudae
pas de plus longues, sans qu'on en sache la rai.son. On les appellc enumerated above (p. 1312), as well as by the number of ridge-
Mookna. ("eux qui les ont longues se nomment Dauntelah, du mot ])lates in the grinding teeth. It seems i)rem:itnn', however, to
daunt qui est le meme que not re mot dent. C'ette ditf<irence n'en attempt subspecific definition at present.
 :

THE ELEPHANTINE: ELEPHAS 1327

DE BLAINVILLE (1839-1804) SEPARATES THE BENGAr>

AND CEYLON BREEDS
From the nipasiirements and characters assigned liy ("orse,
CJeoffroy and F. Cuvier, de Rlain\ille, and Falconer, may be
deduced the foUowing:

Elephas indicus ceylanicus de Blainville, 1845

Figurfs 1170-1172, 1180, 1204, 1226

Island of Ceylon and mainland of India.

Com]>ar(' Elephas indicus var. Mukna Falc. and Caut ., 1847

Var. Mt'kna Falc. (cf. E. indicts ceylanicus de Blain.)

See Falconer, "Palfeontological Memoirs," 1868, I, p. 477, and

1845, Pis. i-iii of de Blainville's "Osteographie," 1839-1864.

Small slender tusks, nearly straight and jiointing ohliciuely

downwards.
Head smaller, hard to l)e distinguished from female elephant;
premaxillary rostrum narrow, more elongate.
Cranial dome more pointed, less brachycephalic, more hypsi-
cephalic (see Figs. 1180, 1204).
Narial openings narrower and smaller (see Fig. 1171).
Five nails on each of the fore feet and four on each of the hind
feet.

Of gentle disposition ; more manageable.

Elephasindicusbengalensisde Blainville, 1845

Figures 801, 936, 963, 1013, 1170-1172, 1175, 1179, 1180, 1204

Chiefly Bengal, Assam.

Compare Elephas indicus var. Dauntela Falc. and Caut., 1847.

Var. Dauntela Falc. (cf. E. indicus bengalensis de Blain.)

Elephas indicus ceylanicus
See Falconer, "Palaeontological Memoirs," 1868, I, p. 477, and male adult

1845, Pis. i-iii of de Blainville's "Osteographie," 1839-1864.

Large and fine tusks, projecting horizontal, but rather elevat-

ed, curve of the perfect Dauntela, known as the "Pullung daunt."
Head larger, premaxillary rostrum broader, less elongate.
All 1,16 nat. size

Cranial dome more rounded, more brachycephalic; frontals

more convex (see Fig. 1204).
Narial openings broader Comparison of Elephas indicos ceylanicd.s (A, female, juvenile; C,
(.see P'ig. 1171).
male) with Adult Males (B, D) of E. indicus bengalensis
Five nails on each of the fore feet and four on each of the hind
Fig. 1 180. Male, female, and geograpliio characters in the skull of Elephas
feet.
indicus. All figures copied, witli identifications, from de Blainville's ''Osteo-
Of more daring disposition; less manageable. graphie," 1839-1864, reduced to one-sixteenth natural size. Compare frontal
aspect of crania (Fig. 1172).
Continental and Insular Subspecies (de Blainville, A, Female of Elephas indicus (?) ceylanicus, with tusks in place. The
1839-1864).— Preceding Falconer's distinction between the Elephas frontal curvature and extreme hypsicephaly remind us strongly of the profile
indicus (Mukna var.) and the E. indicus (Dauntela var.) were de of the juvenile Hypselephas hysudricus (Fig. 1213).
Blainville's observations and his superb plates of 1845. When we B, Male of Elephas iruiicus {'?)bengalensis, with adult tusks in jJace.
compare these plates and the subspecific descriptions of de Blain-
C, .Adult male skull of Elephas indicus ceylanicus, the geographic type of
ville with the descriptions of Corse, Geofifroy Saint-Hilaire and F.
Ceylon.
Cuvier, and Falconer, we are struck by an obvious cranial re-
D, Adult male skull of Elephas indicus bengalensis, the geographic type
semblance, either due to affinity or to analogy, namely
of the Bengal district.
Elephas indicus ceylanicus de Blainville, 1845, resembles
Observe the marked resemblance of the extremely hypsicephalic and
Elephas indicus ( = Mukna var.) Falc. and Caut., 1847.
cyrtocephalic cranium (A) to that of tlic juvenile Hypselephan hysudricus
Elephas indicus bengalensis de Blainville, 1845, resembles cranium (Figs. 1213, 1214), as well as to that of the Elephas iruiicus displayed
Elephas indicus ( = Dauntela var.) Falc. and Caut., 1847. in figure 797.
 : 5

132S OSBORN: THE PROBOSCIDEA

De —
Blainville's Priority. According to de Blainvillc's From condyles to tip of tusk-sheath 33.0 35.25
figures of the Ceylon and Bengal crania, which represent individu- From ditto to anterior margin of molar
als of aijproxiniatcly the same age, de Blainville (1845) separated alveolus 22 . 22 25
.

these two crania subspecifically as types of two subspecies, namely From anterior margin occipital hole to pos-
Elephas rndicus ceylanicuf; and Elephas indicus bengalensis; terior border palate 12.0 11.5
this subspecific separation is fully justified by the more or less Length of palate 8.0 9.5
hypsicephalic characters of the skull, and differences in the jaws Depth of head from condyles to frontal sur-
and tusks; all these characters are widely different in these two face at middle, opposite nasal opening .. . 23.5 23.5
geograjihic subspecies.As indicated above, these .subspecific char- Height from diastemal surface to bulge of
acters appear to be the same as those which separate Elephas occiput 30 75 . 31 25
.

indicus (Dauntela var.) from Elephas indicus (Mukna var.) of Length of condyloid surface 6.5 6.25
Falconer. From this comparison, it would appear that Falconer's From ear-hole to top of occiput 19.0 19.0
two varietal names ('Mukna' and 'Dauntela') propo.sed in 1847 may Length of anterior tooth, upper jaw 8.37]
prove to be synonymous with the two subspecific names of de Width ditto ditto 3.5 [dropped out
Blainville pro])osed in 1845. Number of plates, about ten J

Ontogenetic and Sexual Characters. —De Blainville's Length of anterior tooth, lower jaw 9. 1 ,
37|droppedout
observations (1839-1864) on the sexual characters of Elephas Width ditto ditto 3
indicus are clearly displayed in figure 1180, copied from his plates.
Falconer also stated in this same footnote that "The plates of
We observe in figure 1180A a female apparently of Elephas
teeth in the Mukna variety slope greatly backwards and are exces-
indicus ceylanicus, with extremely hypsicephalic head; in figure
sively and finely crimped; tho.se of Dauntela are much less
1180C we observe an adult male of the same subspecies, E. indicus
crimped."
ceylanicus. In figure 1180B we observe an adult male, apparently of
Elephas indicus bengalensis, with its rounded superior occipito-
Osborn, 1930: We observe that Falconer omits the most
distinctive bathycephalic measurement, namely, (a) from the sum-
frontal dome; in figure 1180D we observe another adult male of
mit of the occiput to the occlusal surface of the superior grinders,
E. indicus bengalensis, with the same rounded dome but with more
as compared with (6) occipital condyles to the orbital level of the
massive cranial projjortions.
frontals.These two measurements are shown in figures 805 and
The crania of these subspecies, both male and female, are
806 and give us what may be called the bathycephalic index of
distingui.shed: (1) By numerous differences in the profile both of
the skull.
the cranium and of the jaws; by the abbreviation of the jaw
(2)
Summary of Osborn, 1930: (1) Prior to Falconer's de-
(C), the oldest individual figured; (3) by the uniformly convex
scription (1847, 1867), de Blainville in 1845 separated the insular
fronto-occipital profile (B and D), growth stages of E. indicus
Ceylon animal as a subspecies, Elephas indicus ceylanicus, from
bengalensis; (4) by the more pointed and hypsicephalic fronto-
the mainland Bengal animal, subspecies Elephas indicus bengalen-
occipital profile (A and C), growth stages of E. indicus ceylanicus.
(2) Falconer (1847) distinguished the two varieties 'Dauntela'
—
Falconer's Two Varieties. It remained for Falconer also
sis.

and 'Mukna.' (3) Lydekker observes (Ency. Brit. 11th Ed., p.

to name and distinguish these two continental varieties, signi-
259) that the insular "Ceylon animal, which is generally tuskless,
fying that he considered them of permanent varietal or subspecific
may be the typical E. maximus [of Linnseus], in which case the
value, as follows (footnote, Falconer, 1867, p. 57, and 1868, I,
Indian race will be E. maximus indicus." (4) It appears probable
p. 477):
that the Ceylon animal (E. indicus ceylanicus) with slender tusks
"Comparison between Mukna and Dauntela varieties of was introduced into India l)y breeders, giving rise by crossing to the
Elephas Indicus. two varieties or hybrids designated by Falconer as Dauntela var.
Mukna Dauntela. and as Mukna var.' This theory was suggested by Schlegel, but
(big head). Falconer (1868), as quoted above, did not admit that the insular
Inches Inches Ceylon breed ever exerted any considerable influence on the north-
Extreme length of cranium 40.5 41 .5 ern Bengalese breeds, nor did he admit the specific distinction of
Width between zygomatics 29.5 29.75 the Ceylonese or Sumatran elephants. (5) As .shown by Osborn in
Ditto post-orbitary proce.s.ses 22.5 24.0 a close comparison of the crania. Falconer's Mukna var. has
Length from niche of occiput to tips of nasals . 22.0 21.0 a cranium closely similar to that of de Blainville's E. indicus
Greatest width of occiput 31 .0 30.5 ceylanicus, while Falconer's Dauntela var. has a cranium more
Width of nasal opening 14.0 15.0 similar to de Blainville's E. indicus beyigalensis. (6) Another ex-
Depth of ditto 5.75 5.75 planation is that the island of Ceylon did not separate from the
Width of tusk-sheaths 14 25
. 17.5 mainland until recent geologic time and that the E. indicus
Narrow width of brow 13,5 13 25
. ceylanicus ranged through continental southern India.
Depth of orbit 6.5 6.5 This problem of the continental and insular raci^s and sub-
Height from condyles to occiput 22.25 22.0 species,however, requires further investigation from the original
Across condyles 8.75 8.5 materials.

'[See footnote 1 on page 1317 above. — Editor.]

 . ,

THE ELEPHANTINE: ELEPHAS 1329

Elephas indicus sumatranus Tcmininck, 1847 namely, the number of dorsal vertebrse and the ridge-jjlato formula,
Figures 1168, 1169, 1181-1185 may not be constant or valid. Consequently we are inclined to
District of Palembang, Island of Sumatra. the opinion that Elephas sumalraiius is a case of geographic

The profound cranial differences which divide the Ceylon and

isolation or insulation with characters of not more than sub-
specific value.
Bengal elephants from each other as well a-s from the Sumatran
elephant in all probability will be found to differentiate the Suma-
tran and other still undiscovered extinct ty|x>s. By comparison
with the evolution of other Pleistocene ungulates it appears prob-
able that a very long period of time separated these continental
and insular subspecies and sfx-cies from each other, a period of time
equi\alent perhaps to nearly half of Pleistocene time or 500,000
years, during which through isolation and segregation the sub-
specificand specific characters were thoroughly founded. Here
again monographic research is essential before we can reach a final
conclusion.
It is interesting to note that this Sumatran species, which we
may now regard a-s a subspecies, was partly di.stinguished by the
Dutch naturalist Temminck as early as 1847. He pointed out
that Elephas sinnatranus has one more dorsal vertebra, i.e., 20.
than E. itidicus, i.e., 19; that the free portion of the intermaxil-
laries is shorter, the nasal cavities are shorter, the space between
the orbits is narrower, the posterior portion of the cranium is

broader than in E. indicus, the grinders have the narrower propor-

tions of Lorodonla africana, the ridge-plate formula is intermediate
between that of L. africana and that of E. indicus, and that por-
tions of the E. sumatranus grinders and ridge-plate formula are
intermediate to those of L. africana, while the shape of the plates
is like that of the plates of E. indicus.
Elephas sumatranus Temminck 1847. "Coup-d'Oil General
Fig. 1181. Elephas indicus sumatranus from Deli (Labuan Deli?], Sumatra.
sur les Possessions Neerlandaises dans L'Inde Archipelagique,"
II, 1847, p. 91. Description. — Temminck distinguishes
Oblique front view of head, showing small and peculiarly formed ears and small
tusks of a young male. Photographed in the Rotterdam Zoological Gardens.
Elephas sumatranus from the African and Indian species as .\fter Lydekker, 1916, Vol. V, fig. 24, p. 83, through the courtesy of the
follows: British Museum (Natural History), .\ugust 10, 1929.

Elephas sumatranus Elephas indicus Loxodonta africana

Characters distinguishing E. suma- Enamel ridges thick crenu- Enamel ridges thin, crenu- Enamel bands lozenge shaped
tranus from
E. indicus and lated, lamcllffi broad, less lated, lamellae narrow; broad as in E. sumati-anus;
Loxodonta africana (cf. Tem- numerous; ratio 3 or 4 ratio 4 or 6 to 1 of width of ratio 3 or 4 lamellae to 1 of
minck, 1847, pp. 91, 92): lamellffi to 1 of width of grinder; 6 to 8 lamellae in width 6 lamellae in 12 cen-
;

grinder; 6 lamellae in 12 12 centimeters timeters

centimeters

Vertebral formula:
Cervicals. . . . . 7.. 7
Dorsals .20..
Lumbars ..3..
Sacrals . 4..
Caudals .34111.

True ribs . . . . . 6..

Floating ribs .14..
 Fig. 1182. Elephas sumatranus Temminck, cotyije male and female crania from Palembang, Sumatra, in the T>eiilen Museum. Compare Jentink,
Cat. Osteol. des Mamm., Mu.';. d'Hist. Nat., Tome IX, 1887, p. 169. Regu en 1845 dc M
J. C. Baud (if. Temminck, Coup d'leil poss. Nderl. Inde arch

Tome II, 1847, p. 91). .\fter photographs kindly furnished by Director E. D. van Oort.
A, Male cranium. Cat. a. Height of occipital crest to tip of premaxillaries 973mm
B, Female cranium. Cat. 6. Height of occipital crest to ti|) of [)remaxillaries 832 mm
A

Fig. 1183. Khi>hax .tmnairmous crania from Sumatra in the Munich

Museum. After pliotograi)hs kindly furnished by Dr. Hermann Diirck and
Herr Gustav Kiisthardt, November 13, 1930.

A, Adult cranivmi from Batang Serangan, l'nt(^rlangkal, cast coast of

Simiatra, belonging to the mounted .specimen represented in figure 1184.
Observe the depn-s.scd and prominent position of the lower border of the orbits,
as com))ared with Klephati indicus brngalcnsU (Fig. 1 170), aiiproaehing rather
E. indieux ccijhniciix (l*"ig. 1 170), and widely contrasting witl \h I.oxodonlinir (Fig. UOSA-C); also the slender tusks, although a male, and extr(-m(! bathy-
cephaly (750 mm. x 550 mm.).
B, Infantile cranium from Sumatra (('Xact locality unknown to the present author), belonging to the mounted s|wcimen n^presented in figure 1185.

Compare the position of the external ear with that in the cranial profih> of the adult E. stimatratuis skull (I'ig. 1184).

1330
 THE ELEPHANTINE];: ELEPHAS 1331

SuMATRAN Elephant in the Munich Museum

After photographs and measuromcnts by Dr. Hermann Diirok and Herr Gustav Kiisthardt
(November 13, 1930)
Fig. 1184. Thi.s specimen was measured immediately after it was shot at Batang Serangan,
I'Mterhmgkat, east eoast of Sumatra. Cranium, right lateral view of same individual as that
shown in figure 1183A.
Total length, tip of tusk to tip of tail 790 cm 23 ft.

Height of shoulder above forelimb '

294
Height above hind quarters 290
Length of ear to point 70
Breadth of car 89
Total length of tusk 130
Observe slender tusks, although a male, and extreme bathycephaly (750 mm. x 550 mm.)
1332 OSBORN: THE PROBOSCIDEA

of the extraordinary elephant with tusks sweeping round in ;i huf^c triangular, form of the ear, the early date at wiiich its upper
curve, mammoth fashion, was in the Indian Pavihon at the margin isbent over, and the presence in the young condition at —
Wembley Exhibition, whore I saw it. You will remember it was least, in —
some cases of a thick coat of black and in j)art Itristly
shot by Theobald and figured some years ago in the Bulletin of the hair."
New York Zoological Society." (Lydekker, 1914.2, pp. 285-288) : "Thanks to the Trustees of
 :

THE ELEPHANTINE: ELEPHAS AND PALEOLOXODON 1333

ill the Fodorated Malay States, records that the Elephants

. . . 496-498, text-fig. ;— Busk, Additional Remarks, id., pp. 498, 499.'].
of the Negri Sembilaii are of ordinary size, although of late years According to Busk, a certain fossil elephant from Japan is referred
most of the big bulls have been killed off. He also mentions that to Elephas indicus, representing, however, a form with teeth some-
jirac'tically all the bulls are tuskers, and very generally have one what larger than the average of the existing one. Quite recently
tusk mueh smaller than the other. The tail is considerably. . . the writer has come to be fully convinced about this first record,
longer tlian in the Indian calf, but since, according to Sanderson, which appears to have been overlooked so long, and proposes here
there is considerable variation in this respect among Indian Ele- to call the Japanese form in question to the credit of Busk, as
phants — which may or may not eventually prove to be of racial follows. Elephas indicus Linn^ Buski, subsp. nov."
value, — do not for the present propose to take any account of this
I This interesting specimen, as described by Matsumoto, is as
feature. The great difference in the contour of the ear, coupled follows
with the excessive development (at least in some instances) of Elephas indicus Linn6 Buski Matsumoto, 1927. "(^n a New
black and in part bristly hair in the juvenile condition, seems Fossil Race of the Asiatic Elephant in Japan." Sci. Re])t. Tohoku
sufficient to justify the separation of the Elephant of the Malay Imp. Univ., Second Series (Geology), Vol. X, No. 3, pj). 57,
Peninsula as a distinct race, under the name of Elephas maximus 58. —
Type. "Antepenultimate upper molar of left side
hirsutus." [l.M']." Inst. Geol. and Palsont. 7266. Horizon and
Locality. — (Op. cit., "Judging from the exceedingly feeble
p. 58) :

fossilization of the remains, as far as examined by the writer, the

present form may probably belong to a \'ery late geological age,
such as the Post-Monastirian. The occurrence of the present
form may indicate a warmer climate, that being well in accord with
certain geological evidence already known." From Ninohe
District, Province of Mutsu, Japan. —
Type Figure. Op. cit.,
1927, PI. xxvii, figs. 2 and 3.

Type Description. — (Matsumoto, 1927, j). 57): "The type-

specimen consists of eleven ridges besides the anterior and posterior
talons, and measures 155 mm. in length, 60 mm. in the greatest
width at the third ridge, and 154 mm. in the greatest lunght of
crown at the seventh ridge, which was just ready to commence to
wear. The middle part of the crown is peculiarly narrowed, being
narrower than both anterior and posterior parts. In a palatal view,
the crown is more or less bent inwards. Its frequency of riilges in
a length of 100 mm. is about 8 on both the sides and about 7 on the
grinding surface. The ridges are only very weakly flexuous; their
inner and outer sides in a fore-and-aft view are not very mark(?dly
convex, being almost parallel at the greater middle part of the
Type of Elephas indicu.s hirsutds
height of the ridges. The anterior and the posterior side of the
Fig. 1187. Typo of Elephas tnaximiis hirsutus, from Negri Sembilan,
Malay Peninsula, formerly in the Gardens of the Zoologieal Society, Regent's moderately to strongly worn ridges at the grinding surface are
Park, London, .\fter I>ydekker, 1916, Vol. V, p. 84, fig. 2.'), reprodiieed through nearly parallel, without any loxodont sinus. The plication of
the courtesy of the British Museum (Natural History), .\ugust 10, 1929. enamel of tho.se ridges is almost uniform from end to end, and is
very fine, regular, and strong, as a very distinctive characteristic
of the present species. The figure of enamel seen in the rather
Elephas indicus Buski Matsumoto, 1927'

[=?Pal»oloxodon buski]
Figure 1188

Tyix- Ninohe District, Province of Mutsu, Japan. Post-Pleistocene to

Recent.
Referred localities (Matsumoto, 1927, p. .)8): "Ov<n- forty miles from the
sea-shore betweenKanagawa and Tokyo (Leith Adams) Yedobashi, Tokyo ;

(Naumann); Province of Mino (Tokunaga); Sapporo, Province of Ishikari,

Hokkaido (Tokunaga); Prefecture of Wakayama."
. . .

Matsumoto (1927, p. 57) remarks that the "occurrence of the = ?Pal^oloxodon buski
Elephas indicus Buski Matsumoto, 1927 [ j

Asiatic elephant in a fos.sil state in Japan was stated for the first Matsumoto, 1927, PI.
Fig. 1188. Type figure of Elephas indicus buski
time by Leith Adams and communicated by Busk [Footnote:
xxvii, fig. 3, one-half natural size, from Ninohe District, Province of Mutsu,
'Leith Adams: Has the Asiatic Elephant Been Found in a Fossil Japan. Original in Institute juf Geology and Palaeontology, Tohoku Imperial
State? Quart. Journ. Geol. Soc, London, Vol. XXIV, 1868, pp. University (No. 7266). \ first superior molar of the left side, l.M\ crown view.

(=asialicus) proves that such

'Osborn, 1930: .\t first disposed to place this type near Elephas indicus, comparison with figure 1177, an M' of E. indicus
hysudrindicus.
reference is doubtful. It appears rather to belong near Palseoloxodon namadicus nawmnni, P. mmadicus namadi, or P.
1334 OSBORN: THE PROBOSCIDEA

sliglitly worn ridges is mesially laminar and laterally aniuilaf. The than in Dubois' type superior molar (I'if;. 1160) of I'ahiolo.rodon
worn surface of the sixth ridge, which is very slightly worn, con- hysudrindicus.
sists of five mammillse arranged in a transverse row. Tlie layer of Osborn, 1930: From
the above comparisons Osborn is in-
enamel is considerably thin, measuring about 1-1.5 mm. in thick- clined to agree with Lydekker that the above grinding teeth,
ness on the anterior and the posterior side of the ridges." namely, the type of Elephas indicus buski (Fig. 1188) and E.
Leith Adams and BrsK (1868). — (1) In 1868, as mentioned namadicvs ref Lydekker (Fig. 1 189) belong either to E. [Palieoloxo-
. ,

above, page Adams (1868, p. 497) described and

1062, Leith don] namadicus naumanni, E. [P.] namadicus numadi, or to some
Tooth of Elephas Indicus from Japan," discovered
figured a "Fo.ssil even more progressive species of Pahcoloxodon e.g., E. [P.\ ,

forty miles inland between Kanagawa and Jeddo. He rightly hysudrindicus.

interpreted this tooth (Fig. 1189 of present Memoir) as an r.M'-,

describing it as follows: "This penultimate true molar, right side,

upper jaw, although partaking of the characters of E. Armeniacus,

Falc, differs from the latter in its thick plaits, their less approxi-
mation, the festooning being carried round the loops of the disks of
wear, and the total absence of any mesial expansion." (2) Rusk
{op. cit., p. 498), after comparing the tooth with E. [Farel^pha)<\
Armenicu-vs Falc. and E. [P.] columbi Falc, and finally with
a corresponding fossil tooth [in his opinion an l.M' of E. indicus],
concludes as follows: "But these differences [referring to his Second Right Superior Molar referred to Pal.eoloxodon
compari.son with E. indicns] do not appear to be of much im- Fig. 1189. A second right superior molar, r.M", probably from the Middle
portance, and there seems to be every reason to believe that the Pleistocene of .Tapan, between Kanagawa and Tokio (Yedo). .\fter Lydekker,
Japanese fossil tooth belonged to a form of E. indicus, with teeth
1886.2, p. 168, fig. 29, about one-half natural size. Original in the Museum
at St. John, New Brunswick, Canada.
somewhat larger than the average of the existing one." (3) The This molar, first figured by Leith Adams and Busk (1868, ]). 497), and
teeth described and figured by Adams and Busk (1868) and by interpreted as a "Fossil Tooth of Elephas Imlicius from Jai)an," was compared
Matsumoto (1927) are not dissimilar in respect to thickness and by the same authors {op. cit., p. 498) with the E. [Parclephas] armrniacus
the interspacing of the enamel ridge-plates, but in the type of Falc. of Armenia and of China. Lengtii 187 mm., width 76 mm., height
111 mm. The present author regards it as referable eitlier to Pahfoloxodon
Elephas indicus buski the enamel foldings or plications are much
namadicus naumanni, P. namadicus namadi, or P. hysudrindicus. Observe
clo.ser and more compact and consequently mon> numerous. The crimping or plication of the enamel and absence of 'loxodont sinus,' resem-
ridge-plates in the Adams-Busk grinder (Fig. 1189) also in the type blance to Elephas indicus grinders. Observe similar plication or crimping in
of E. indicus buski {¥'\g. 11S8) fire much more widely interspaced the type (Fig. 1160) of P. h)jsudrindicus.

2. AND MEASUREMENTS OP^ THE INDIAN ELEPHANT

DISTINCTIONS
Comparison has been made in Chapter XV between the Indian and the African elephants as regards dental,
cranial, and vertebral structure and prevailing feeding habits. As to dental structure, the very earliest observers
remarked the profound differences between the crown view of the molars of the African and of the Asiatic ele-

phants, finally figured and described (1846) by Richard Owen (Fig. 1191). It is not possible that the African ele-

phant, with its coarse-plated, relatively low-crowned grinders, can attain the same age as the Indian elephant,
Avith its very high and long-lived grinding teeth.

Detailed observations by Falconer (fully discussed in Chap. XV) on the food of the Indian and African ele-

phants indicate that the straight tusks of the African elephant, as in all other Loxodontines, are used as crowbars
in uprooting trees as well as in combat. It does not appear that the shorter and more slender tusks of the Indian
elephant are used to so great a degree for such purposes; they are invariably more slender and more strongly
upcurved, and those of the old Indian bulls may attain great length (Fig. 1179). Doubtless for a very long period
of time the Asiatic elephant was hunted for its ivory, even before it was domesticated, both in India and Ceylon.
The ivory tusks, both of the Indian and ('eylonese varieties, are capable of carrying very heavy logs, held in jilace
across the tusks by the trunk.

In longitudinal section (Fig. 1 192) it appears that the low-browed African (>ranium, in perfect liarniony with its

low-crowned grinders, is in very wide contrast to the high-browed Indian {'ranium, with its very liigli grinding
teeth. On the principle explained in Chapter XV, the Indian cranium (Fig. 800) is far more halhycephalic than
 African Elephant. \ nat. size. Asiatic Elephant.

Fig.1 190. ,Su]jerior view of tlie. head of a young African elephant (left) Crown view of the third inferior molar of the right side of: Fig.
Fig. 1191.
and ofan adult Indian elephant (right). After Geoffroy Saint-Hilaire and SS,Loxodonta africana; Fig. 89, Elephas imlicus, one-third natural size, .'^fter
Frodcrie Cuvier, 1825, Livr. LI, LII, one twenty-fourth natural size. Owen, "A History of British Fossil Mammals and Birds," 1840, pp. 2.30-232:
The su(K'rior view of the head of the adult ElepJias indicus should be "Thus in the African Eleijhant, (fig. 88), in which the lozenge-shaped plates
compared with the superior view of the heads of the adult male {Loxodonta are always much fewer and thicker than the flattened ones in the Asiatic
africana albertensis)and female {Loxodonta africana peeli), as mounted in the species, the variationwhich can be detected in any number of the grinders of
American Museum collection and photographed from above in figure 1063 of this the same size very slight. ... In the molars of the Asiatic Elephant, (fig. 89),
is

Memoir. The very marked differences between the male or female Loxodonta which, besides the difference in the shajw of the jilates, have always thinner
and the male or female Elephas are indicated in this comijarison. Loxodonta is and more numerous i)lates than those of the African species, a greater amount
distinguished by the rounder occiput, lacking the pronoimced air sinu.ses of of variation in both these characters obtains; and the like caution is still
. . .

E. indicus, also by the greater elongation of the antorljital or facial i-egion of more requisite in the comparison of the molars of the Mammoth {Elephas
the cranium. The differences between the cranium of Loxodonta and of primigenius), which, having normally more lumierous and thinner plates than

Elephas are quite as pronounced as the differences between tiie grinding teeth in the existing Asiatic Elephant, present a much gieater lange of variety."

displayed in figure 1191.

A.M. 5/939

ELEPHAS INDICUS LOXODONTA AFRICANA

Indian cranium (Amer.
Fig. 1192. Contrasts of (right) thelow-browed African cranium (Amer. Mus. Dept. Mam. 51939) with (left) the high-browed
The deeply embedded below the cranial air cells, is seen to be slightly larger
Mus. Dept. Mam. 54261). Compare figures 805, 806, 800, 1061, and 1053. brain,

in the Indian elephant (6686 ccm.) then in the African elephant (6651 ccm.). Compare figure 1112.

Bath., the vertical bathycephalic diameter from summit of occiput to Resp. —Resp., the respiratory axis from the anterior to the ijosterior

crown of grinding teeth. nares ( = respiratory plane).

S.oc. — .S'. oc, the supraoccipital axis (=occipital plane). Pal.—'Pal., palatal axis of the roof of the mouth.

B. oc. — B. oc, the basioccipital axis ( = basioccipital plane). —

M' M^, grinding surface of the superior molar teeth.
1335
1336 OSBORN: THE PROBOSCIDEA
the African cranium (Fig. 1061), yot the African elephant, as shown in the measurements (legend to Fig. 1053) of
the growing "Khartum" {Loxodonta africana oxyotis) attains its greater height very rapidly, namely, from a height
(jf 4 ft. I'l in. in October, 1906, to a height of 10 ft. 8% in. in January, 1930, or an average annual growth of 3% in.

in twentj^-four years. The extreme shoulder height of the African elephant, in the flesh, is estimated at 1 1 ft. 6'A in.

(see footnote 2 on page 1022, caption to Fig. 912), whereas, as shown in the observations below, there are only two
records (neither of which is absolutely reliable) of the attainment by the Indian elephant of a greater height than
10 ft. 6 in., the average being about 9 feet.

Shoulder Heights of the Indian Elephant

Throughout this Memoir the estimates of shoulder heights are based on the fully extended forelimb, with
certain allowance for the cartilages between the bones, for the foot pads, and for the great muscles of the neck.

As compared either with the African or with the great

extinct elephants of Pleistocene time, the Indian elephant is

generally inferior in stature; Mammonteus primi-

it exceeds
genius in height by a foot to eighteen inches; the somewhat
doubtful 'record' of the Indian shoulder height, as shown in
figure 1194, namely, 10 feet 6 inches, agrees with the highest
figure given by Corse below, while the average large Indian

elephant rarely exceeds 9 feet.

The observations of Corse (1799, p. 35) chiefly relate to

the heights of domesticated elephants and are very interesting

to cite in this connection:

(Cf. Corse, 1799, p. 35): "elephants attain their full size be-
tween eighteen and twenty-four years of age. ... In India, the
height of females is, in general, from seven to eight feet and that
;

of males, from eight to ten feet, measured at the shoulder." One

el('l)hant, on good authority, exceeding ten feet, was a male, meas-
uring as follows:

From the top of the shoulder, perpendicular height 10 ft. 6 in.

"
From the to]) of the head, when set up, 12 " 2
" "
From the front of the face to the insertion of the tail 15 11

Of 150 Bengal elephants only a few male.s attained 9 feet 6

inches, not one ofthem attained 10 feet. The Ceylon elephants arc
neither higher nor superior in any respect to those of Bengal.

Contrary to prevailing opinion, the elephant is in no sense

Unoui.iguadism. Radiograph of Right Foot of Young plantigrade; as shown in the radiograph reproduced herewith
Indian Elephant
(Fig. 1193) the foot is unguligradc; the phalanges are subver-
Fig. This beautiful radiograph of tlio pes of a young
1193.
oiophant taken from the inner side of t he foot exhibits clearly the tical in position, the terminal i)halanges are greatly reduced in
toraplete inner or including Ph. IJ, Ph. I, Mts. I, Cu.,
first digit,

Nav., Astrag., Cal., Ep., and Tihw. The remaining four digits
size and are encased in horny sheaths which protect the front
are faintly seen in outline. The pes is shown to be ungidigradr, l)art of the foot from injury, while a posterior elastic pad is
to rest upon the ungues surrounding the tips of the terminal
i.e.,

phalanges, in this case Ph. II; it proves that EUphnx is in no sense l)ressed down in a walking or running gait.
plantigrade. It also exhibits the cartilaginous interspa<'es, which

add considerably to the height of the skeleton. Thi' scale is

in the accompanying figiu'e (Fig. 1194) are given the esti-
approximately two-fifths natural size. Reproduced through tlic

courtesy of Dr. G. M. Vcvcrs of the London Zoological Society. mated shoulder heights of three elephants in the American
 THE ELEPHANTINE: ELEPHAS 1337

Museum collection, ranging from 8 4% inches to 8 feet 8% inches, as compared with that of a very large male
ft.

measuring 10 feet 6 inches in the flesh,' according to Rowland Ward's "Records of Big Game" and a measure-
ment taken from Dollman-Bather, 1927. This 10 feet 6 inches maximum agrees with the single exceptionally
large specimen reported by Corse.

ELEPHAS INDICUS
Max.Ti

2660
2566

\\s
3
 :

III. CHARACTERS OF THE UPPER PLIOCENE AND LOWER PLEISTOCENE SPECIES

HYPSELEPHAS HYSUDRICUS AND PLATELEPHAS PLATYCEPHALUS
First it is important to note the great geologic antiquity and wide separation in time of the two fossil species,

now to be described, from the living Elephas indicus and its geographic subspecies.

This great geologic time interval between E. indicus, Hypselephas hysudricus, and Platelephas platycephalus
practically spans the whole Pleistocene period or Glacial Age, now estimated at about 1,000,000 years, in contrast
to the relatively brief period of 400,000 years which may have been in the mind of Falconer from the estimates of
Pleistocene time by his contemporary Charles Lyell.

Among the fossil types of the Siwaliks and contemporary geologic horizons of India, as displayed in the
geologic table of Pilgrim-Brown-Osborn-Colbert (1910-1927-1935 — Fig. 413), also the more detailed correlation

table of Pilgrim (1926 Fig. 1195), it appears that Hypselephas hysudricus was extremely abundant in the Lower
Pleistocene fluviatile deposits of the Simla Hills (Fig. 1196) below or eroded from what is known as the "Boulder

Conglomerates."

Hypselephas hysudricus occurs as a companion

of the giant stegodont species Stegodon insignis-

ganesa, also of the bovine Buffelus palseindicus. Its

remains are very numerous and in the present Mem-

oir references are given to 29 specimens very care-
fully described and in many cases beautifully figured

by Falconer both "Fauna Antiqua Sivalensis"

in the

and in the "Palaeontological Memoirs" of 1868, also

to 14 specimens collected by Barnum Brown in 1922
and referred by O.sborn to Hypselephas hysudricus,
in addition to a specimen also collected by him from

the "base of conglomerates," described herewith as

Platelephas platycephalus. In brief, the collections
of Falconer and of Brown include upwards of 43 type
and other specimens referable to H. hysudricus, so
that the cranium and dentition of this species are

very well known, in fact, better known than of many Upper Siwalik Expo.sitkks of the Simla Foothills, India
Compare I'okting Map (Pi. xxv), also figure 119.5 opposite ani!
of the living subspecies of elephants. loealitios listed on p. 1347 below
Fig. 1196. In the very thick Plio-Pleistoiene horizons of this region oceur
Osborn's analysis proves that Elephas hysudricus the formations whieh have yielded in the Falconer and Barniim Brown collec-
tions not less than 66 specimens of Archidiskodon planijrom, in addition to
represents a distinct generic phylum which may be numerous specimens referred to Stegodon insignis-gamm from the overlying
named Hypselephas ;' in this phylum for the present variegated beds at the base of the Boulder Conglomerate formation. The
Falconer and Brown collections include also remains of Hypsekphas hysudricm,
are placed two species in addition to the new forms Stegodon pinjorensis, Platelephas platycep}ialux,

etc. This region therefore is the richest known in remains of the Plio-Pleisto-
Elephas hysudricus Falconer and Cautley, 1845, cene proboscideans of India.
Eleven specimens of the typical Hypselephas hysudricus are recorded by
1846 = Hypselephas hysudricus. Barnum Brown as from 'below conglomerates' or 'top ol variegated beds,'
apparently deiKisited by erosion from the "Boulder Conglomerates" and three
Elephas platycephalus angustidens Osborn, 1929
from the "Upper Siwaliks." Brown's localities are near Chandigarh, Si.swan,
= Hypselephas hysudricus. and Kalka, as shown in the present figure.

'[See Osborn, 1934.926, p. 285, iind Vol. I, p. 12, of the present Memoir. -Editor.]

1339
1340 OSBORN: THE PROBOSCIDEA

Wholly distinct is the very primitive, flat-headed Platelephas platycephalus Osb. Before defining and char-
acterizing these new forms, the classic 'Elephas hymdrictifi' of Falconer may be fully described, preceded by the
generic characters of Hypselephas, as follows.

Superfamily: ELEPHANTOIDEA Osborn, 1921

Family: ELEPHANTID^ Gray, 1821

Subfamily: Elephantine Osborn, 1910

Genus: HYPSELEPHAS Osborn, 1936

Original rofcrcnoo: Osborn, 1934.926, p. 285 (iiomen nudum), and Vol. I of the present Memoir (1936), p. 12.
(ienotypic species: Elephas hysudricus Falconer and C'autley, 1845, 1846, and Elephas platycephalus angustidens Osborn,
1929.

—
Generic Giiaracters. Primitive elephants of India, progressive in cranial structure; cranium
elevated (hypsicephalic) occiput elevated with l)roadly transverse frontal crest, frontals deeply con-
;

cave. Premaxillaries relatively narrow or laterally compressed; tusks relatively straight, incurved,
somewhat divergent at base; rostrum of lower jaw elongate, prominent, ramus shallow. Orbits large,
dej^ressed, near maxillary rostrum. Molar crowns low; ridge-plates convexo-concave, reversed above
and below, rudimentary 'loxodont sinus.' Ridge-plate formula: 3 n--rs^- M

1. FALCONER'S ORIGINAL DESCRIPTIONS OF ELEPHAS [HYPSELEPHAS] HYSUDRICUS

Hypselephas hysudricus Falconer and Cautley 1845, 1846 , hys^idricus as figured by Falconer and Cautley (Fig. 1213) also by
Figures 794, U74, 1197-1206, 1209-1215, 1218 0.sborn in the present Memoir (Fig. 1214) closely resembles the

Lower Pleistocene, apparently deposited by erosion from the Boulder juvenile cranium of Elephas indicus as figured by de Blainville

Conglomerate zone, India. In the same zone occur Slegndnn ganesa, IHcern- (Fig. 1180A). The.se tall, highly compressed crania differ widely
rhinu.1 plalijrhinus, the horse of India {Equus nvalensis), and the camel of India from the low, flattened juvenile crania of Loxodonta. Ob.serve the
(CameUis sivalensis). Localities: Near Siswan, Chandigarh, Charnian, Kalka, rudimentary loxodont -sinus on the grinders (Figs. 1198, 1199, 1203).
as recorded by Brown.
—
Type Description. (Falconer and Cautley, op. cit., 1846, p.
E. [Elephas] Hysudricus Falconer and Cautley, 1845, 1846. 41) "Fig. 3fl, of this plate [PI. i] shows a section of the jienultimate
:

"Fauna Antiqua Sivalensis," 1845, and 1846, p. 41. Type.— upper molar [Fig. 1197, 3a] of an undescribed Indian fossil species
Second superior molar, M-. Paratype.— Portion of third named E. Hysudricus in this work. The tooth is in the middle
inferior molar, M,^. Horizon and Locality. Siwalik Hills, — stage of wear, eleven of the thirteen plates of which it is composed,
India, Lower Pleistocene. Type and Paratype Figures. — having been in use, and the two anterior ridges being worn out.

Op. (77., PI. I, figs. 3n, 36 [Fig. 1197 of the present Memoir]. The .same vertical disposition of ivory, enamel, and cement, is

Relationships. — This
phase was abundant in the
specific presented as in the African Elephant, but the plates are thinner and
Lower Plei.stocene of India, for in the present Memoir comparative a greater number of them is included in the same length, nine or
measurements are given of 29 specimens described by Falconer and ten plates in the latter being developed in the space occupied by
of 14 specimens collected by Dr. Barnum Blown in 1922, in all 43 thirteen or fourteen plates in the equivalent teeth of E. Hysudricus.
specimens. The plates are also more vertical, the interspaces occupied by the
By Falconer Elephas hysudricus was believed to be related to cement are wider than the ivory plates which represent
in general
Elephas indicus. By I.eith Adams it was considered ancestral to very atteiuiated wedges. The layer of enamel is ])roi)ortionally
both Elephas and Loxodonta. By Pohlig it is considered ancest ral thicker than in the African Elephant, approaching, in this respect,
to Elephas = Palseoloxodon] namadicus. By O.sborn it is regarded,
[
the teeth of E. planifrons, fig. 5. The vertical height of the tooth

on the whole, as more clo.sely related to Elephas indicus although is comparatively less in this specimen than in the African species,

by no means ancestral. The cranium is widely distinct from that th(! compensated by a greater development of the
difference being
of Palifoloxodon namadicus or of Loxodonta africana; the grinding basal ma.ss of ivory. This specimen measures 7.7 inches in length.
teeth are relatively shorter than those of P. namadicus, they re- A portion of the last molar of the lower jaw of this species is shown
semble rather certain of the grinding teeth referred in this Memoir in vertical section in fig. 36, comprising about fifteen plates [Fig.

to s])ecies of Pala>oloxodon of East India and of .Japan. The ex- 1197, 36]. The entire tooth, which is seen in figs. 12 and 12a of pi.
treme hyp.sicephalic and compre.s.sed juvenile cranium of Elephas 7, in situ in the jaw [Fig. 1 198 of present Memoir], is more elongated,
 : ,

THE ELEPHANTINJ5: HYPSELEPHAS 1341

and incluclcs a greater number of divisions than is usual in the last of the"Fauna Antiqua Sivalensis" (1845-1847); from this Atlas
inferior grinder of E. Hysudricus." and from Falconer's te.xt in the "Palaeontological Memoirs" of
Lydekker (1886, p. 117) erroneously selects the adult cranium 1868, Vol. I, pp. 421-440, may be derived the following notes:
as the type, in the following sentence: Mus. "M.3109. The
Brit. Plate I.[Type].— Fig. 3a, M- (.section), 11 ridge-plates in u.se,
cranium of an adult, with the crowns of the molars broken off. 2 anterior being worn out fig. 36 [Paratype, Osborn] M3 (section)
;

Figured by Falconer and Cautley in the 'Fauna Antiqua Sivalensis,' about 15 ridge-plates. Plate iv. Front view of skull
pis. IV., and v. figs. 1, 2. This and the following specimens are the
types of the species. Purchased, 1838." Length of cranium from occipital pro-
This adult cranium (Brit. Mus. M.3109— Fig.
1205 of the tuberances to left incisive 45. in.
present Memoir) contains only the badly fractured crowns of M', Extreme width of head (restored on left
Ms, consequently it cannot be the type, although Lydekker's con- side)
fusion may have arisen from Falconer's description of Plate i, fig. Length of alveolus of M^
3a (1868, I, p. 422), to which i.s added "(Reproduced in PI. v, Width of alveolus of M^
fig. 1.)."
Falconer's Ridge Formula. — In addition to his clearly Plate V, figs. 1, 2, 3, 4. Different views of same skull as above.
recognizable type. Falconer examined twenty-nine specimens (from Plate VI, figs. 1, 2, perfect small skull, Dp^ with 5 ridge-plates.
the ?Boulder Conglomerate zone) which he referred to the same Dp-" with +7+ ridge-plates; fig. 3, under surface of young skull.
species. These specimens arc admirably figured in his great Atlas Dp' with 5 ridge-plates. Dp'' with +8-|- ridge-plates. [Footnote:

//y "!.;
 —

1342 OSBORN: THE PROBOSCIDEA

Small head, Dp^ with 5+ ridgp-platos, Dp^ with +7+ ridgo-platos; suDRicus. —From the foregoing descriptions of twenty-nine type
another imperfect head of young animal. Dp' with 5+ ridge-plates, and examined by Falconer
referred specimens of Elephas hysudricus
Dp^ with +8+ ridge-plates. Fragment of large cranium; l.Dp^ may be deduced the following ridge formula, in which the minimum
with +7+ M' with +8+ ridge-plates.] Plate vii,
ridge-plates; figures represent young or jiartly developed teeth, while the

fig. 1, fragmentupper jaw,

of Dp^ and Dp''; figs. 2, 2a, r.M' with maximum figures represent the specific and mutational stage
12 ridge-plates (see vertical section) figs. 3, 3a, IM- with 13 ridge-
; attained by this species at the beginning of Pleistocene time:
plates (see vertical section); fig. 5, l.Dps with 7 or 8 ridge-plates;
fig. 6, ?l.Dp3 with 9 ridge-plates figs. 7, 7a, l.Dpa with 7 or 8 ridge-
; Dp 3^ Dp4' Ml 9 + - 1 2
M 2. MSr-
plates; fig. 8, l.Dp4 with 9 ridge-plates; fig. 9, Dp4 with 9 ridge-
plates; figs. 10, 10a, Ml with 12 ridge-plates; figs. 11, 11a, ?M2 Affinities of Hypselephas hysudricus. —Falconer in his

with 12 ridge-plates; figs. 12, 12a, M3 with 17-18 ridge-plates, type description (1846, p. 41) of the molar teeth of Elephas hysu-
enamel thin, plicated. Plate viii, fig. 1, cranium with M'"- and dricus (Fig. 1197) intimates that these teeth are to be regarded as
tusks; M' with 5 remaining ridge-plates and a heel, M- with 10-11 more primitive than tho.se of E. indicus, that is, with fewer and
ridge-plates; fig. 3, lower jaw; fig. 4, entire lower jaw, Mi with shorter ridge-plates. In his c()m])arison of the last superior molars
-l-9-|- ridge-plates, Ms (in germ) with 9 exposed; fig. 5, extremely of E. indicus with those of E. hysudricus, as quoted in full below,
aged right jaw, M3 with 5 remaining ridge-])lates only. Plate xii, he again intimates that the grinding teeth of E. indicus may be
fig. 13a, r.Ms with 104- i-idge-plates, one or two ])lates gone d(>rived from those of E. hysudricus. It was not his wont to discuss
an enormous tooth. Plate xii.b, fig. 4, M' with 8 ridge-plates phylogeny.
(remarkable in having so few ridge-plates), M^ (in germ) showing Leith Adams. —The next author to discuss the relationships
11 ridge-plates. Plate xii.c, figs. 6, 60, l.Mj with -|-10-f- ridge- of E. [Hypselephas] hysudricus was Leith Adams in his "Mono-
plates (believed to have come from Nerbudda). graph on the British Fossil Elephants," 1877-1881, p. 244. Leith
Falconer's (1868) Ridge Formila of Hypselephas hy- Adams, as quoted by Lydekker (1886, p. 98) erroneously suggests
that Elephas hysudricus gave rise to two branches, one comprising

Hypsei.ephas iiYSUDRTcrs, Pakatype Crown- hi- Tiiirti) Rioht Ixferioi;

Molar
Compare section (Fig. 1197, right)

Fig. 1198. Eighteen ridged third inferior molar of the right side, r.Mrt
(of. Elephas hyxudricus) . After Falconer and Cautley, 1846 (1845|, PI. vii,

figs. 12 and 12n, described in 1867 (Falconer, 1867, p. 8, and 1868, Vol. I, p.
428) as follows: "Fragment of lower jaw, with last molar, entire, in situ. The
tooth is more elongated, and includes a greater nmnbcr of divisions (17 or 18)
than is usual in the last inferior grinder of E. Hy-iudriciix. The specimen is now
cut into sections.— B. M." (Lydekker, 1886, p. 121): Brit. Mus. "M.3146. Fig. 1199. Referred third left inferior molar, I.M3, with 13-f ridge-plates
Fragment of the right ramus of the, mandible, containing the third true molar, (typical 17-18), of 1 1 ]i jixdr i>hm hijxwiricua (Amer. Mus. 198(i9a). Collected by
which has been longitudinally and vertically bisected, and shows eighteen Barnum Brown in 1922 below Boulder Conglomenites, 'I'pper Siwaliks,' near
ridges. The enamel is very "thin and much i>licated." Reproduced two-
. . .
Siswan, India. One-third natural size. See section of same molar (Fig. 1201).
ninths natural size. Observe that the 12 ridge-plates exposed agree very closely in character,
Compare I.M3 (.\mer. Mus. 19869« -Fig. 1199) also I.M3 (.Vmer. Mus. with slightly expanded 'loxodont sinus,' with tho.sc- of Falconer's |i:iraly|)c
19867— Fig. 1203). (Brit. Mus. M.3146) as shown in figure 1198 opposite.
 THE ELEPHANTINE: HYPSELEPHAS 1343

lElephas priinigenius and E. indicus, the other comprising Elephas Fig. 1200. Second left in-
HYSUDRICUS Ref.
namadicus, E. anliquus, and E. africanus; this suggestion of Leith ferior molar, l.Mo, of Elephas
[Hypselephas] hysudricus ref.
A.M./Saje '/4 /Yafiije
Adams that E. hyi^i/drirus gave rise on the one hand to the Indian
(Amer. Mu.><.l.')878), acquired
elephant and to tlie mammoth, and on the other liand to the through exchange with the
African elephant and its relatives E. namadicus and E. antiquus, Briti-sh Museum in 1911.
arose from the very confused ideas and theories of phylogeny at One-fourth natural size.

time when Leith Adams wrote his valuable monograph. Observe 11 to 12 ridge-plates
the
as in Falconer's figure (184(i
Falconer's comparative measurements (1868, Vol. I, pp. 435,
[1845], PI. vri, figs. 11, Ua),
436) of the ridge-plates of Ms in Elephas hysudricus, E. namadicus, described in the "Palaeontologieal Memoirs," Vol I, p. 428, and noted in
E. indicus [bengalensis], and E. priinigenius are extremely valuable the table of measurements herewith.
for reference as well as an index of progressive evolution.
The comparison below (p. 1344) indicates that as the ridge- Specific Characters. —
(Osborn, 1929) Infantile cranium and
plates increase in height and in number they decrease in the thickness jaws extremely hypsicephalic, forehead plane (Figs. 1213, 1214).
nf the enamel and in the width of the intervening spaces. Adult cranium with very broad rugose fronto-occipital expansion,

TABLE XVI. Twenty-nine Specimens, Type and Referred by Falconer (1868, I, pp. 421-440), of Elephas hysudricus
Figures, Page References, Measurements, and Indices

Superior
1344 OSBORN: THE PROBOSCIDEA

forehead concave (Fig. 1205). Anterior narcs broadly depre.s.sed low, primitive, with very prominent rostrum (Figs. 1214, 1205).
(Fig. 1205). Premaxillarics and tu.sk insertions narrow, nearly Orbits very low and prominent, unworn ridge-plates (Fig. 1203)
parallel (Fig.s. 1213, 1205) as in Elephas. Inferior mandible shal- low, primiti\'e, composed of 4 S eonel(>ts, totally unlike the ridge-

Measurements in Inches and Millimeters

 :

THE ELEPHANTINE: HYPSELEPHAS 1345

])latc.s of Elephas iiulicus. Ridge-plato formuki (Osborn, 1930) which are also observed in certain Japanese and East Indian speci-
mens now referred in this Memoir to the genus Palseoloxodoii,
Dp 3„„ Dp 4f± M 1^ M 2't\f^i M S^ subfamily Loxodontinae. The cranial profile of E.
hysudricus, on
the contrary (see comparative figure 1204), approaches that of
Total niimlxn- of ridge-plates, including Dp 2, in successive Elephas indicus var. Daunlela, which, in turn, agrees with the
use: iXiOT ^7+- '•^^ compared with the total, including Dp 2, extreme E. indicus bengalensis or continental variety of Indian
Elephas indicus ridge-plates (Falconer, 1863, p. 65): fXi™ '^i-
elephant. The true relationship of E. hysudricus to the existing
Cranial Comparison with Elephas indicus. —The phyletic species and subspecies of E. indicus can only be determined by
relationship of Elephas hysitdn'cus to Elephas indicus is somewhat fundamental sections of the basicranial axes. Meanwhile Elephas
uncertain; E. hysudricus differs in its narrow molar teeth and in hysudricus is provisionally placed in the same subfamily (Elephant-
the presence of a faintly indicated 'loxodont sinus,' characters inae) with Elephas indicus.

ELEPHAS HYSUDRICUS
/iTner. Mus. /9a30 Ref.
r.-m ^
^ Tiat. 51JS

Fig. 1202. This specimen was found by Barnum Brown

in 1922 at the top of the variegated clay beds, 'below con-
glomerates,' nine miles west of Kalka, India.

Al, Referred third right superior molar of Hypselephas

h ysiidricui (Amer. Mus. 19S30), exhibiting 19 ridge-plates,
of whicli the anterior ridge-plates 1 to 5 onlj' are worn, as
shown in A3. Length 271 mm., breadth 84 mm.
A2, Transverse section of ridge-plate 13, as indicated by
tlie arrows.

A3, Pliotograph of occhi.sal surface showing more or

less fully worn anterior ridge-plates (1-5).

2. OBSERVATIONS OF OSBORN ON THE FOURTEEN SPECIMENS COLLECTED BY BARNUM

BROWN (1922) REFERRED TO HYPSELEPHAS HYSUDRICUS
The following tables, giving detailed measurements of the specimens in the American Museum (Barnum
Brown) collection, (namely, length, breadth, index, number of ridge-plates, and frequency of ridge-plates in

10 cm.), supplemented by lateral and crown views of seven specimens shown in figure 1203, afford for the first time
a complete knowledge of the superior and inferior grinding teeth of Elephas [Hypselephas] hysudricus Falc.
The ridge-plate formula, based on the above fourteen specimens in the American Museum, is somewhat higher
than that deduced from Falconer's Iwenty-nine specimens, as follows:

Falconer (1868)
Osborn (1930) :
: Dp 3 ^
Dp 4 +-^
Dp 3 Dp 4|| M 1
M i^^H M 2 ^^iV^M 3 ry-rs
1

^ M 2 ^^^\^ M 3 Vr.
,t.
 znTier vteif

Fig. 1203. Detailed caption will be found on opposite page. Sec also figure 1201 for seetioii of third sui)erior molar (Anicr. Mas. 19863).

1316
 THE ELEPHANTINE: HYPSELEPHAS 1347

Since the American Museum Banium Brown)

( collection is very precisely recorded as to geologic level and
locality (Fig. 1196, PI. xxv), that not m
but found redeposited "below conglomerates" or "base of con-
is, .si/w

glomerates," near Siswan, Chandigarh, Charnian, and Kalka, and since the detailed measurements and indices
taken by Osborn demonstrate the close agreement with the Elephas hysudricus of Falconer, it is important to
record this collection in the same detail as the American Museum (Barnum Brown) collection of Archidiskodon
planifrons (p. 955) from the Pinjor horizon of the same region.

Hypselephas hysudricus Ref. and Platelephas platycephalus Type, Collected by Barnum Brown (1922) below Boulder
Conglomerates, not in situ

Elephas [Hypselephas] hysudricus Falconer

Level below
Boulder Conglomerates Locality Amer. Mus. Number Materials

'Top of variegated beds, 3 miles west of Chandigarh Amer. Mus. 19866

below conglomerates'

'Top of variegated beds, 3 miles northwest of Amer. Mus. 19786

below conglomerates" Chandigarh
'Below conglomerates' 2}i miles south of Charnian Amer. Mus. 19809
'Below conglomerates' 2 miles south of Charnian Amer. Mus. 19799
'Upper Siwaliks' Near Siswan Amer. Mus. 19962

'Below conglomerates' 3 miles north of Siswan Amer. Mus. 19783

'Below conglomerates' 3 miles north of Siswan Amer. Mus. 19867

'Upper Siwaliks' Near Siswan Amer. Mus. 19863
'Upper Siwaliks' Near Siswan Amer. Mus. 19869a
'Top of variegated beds, 9 miles west of Kalka Amer. Mus. 19830
below conglomerates'
'Below conglomerates' 2J.2 miles south of Charnian Amer. Mus. 19809A
'Below conglomerates' 3 miles west of Chandigarh Amer. Mus. 19956
'Upper clays, below con- 2J-2 miles northeast of Amer. Mus. 19416
glomerates' Charnian
'Below conglomerates' 3 miles west of C'handigarh Amer. Mus. 19915
1348 OSBORN: THE PROBOSCIDEA

TABLE XVn. FovKTEEN Specimens Collected (1922) by Barnum Bhown (exclusive ok Ameii. Mus. 15878) and Referred

BY OSBOHN TO Hy'PSELEPHAS HYSCDIUCUS, ALSO TyPE OK PlATELEPHAS PLATYCEPHALUS, LoWEH PLEISTOCENE

Figure in
 THE ELEPHANTINE: HYPSELEPHAS 1349

the head, neck, and proboscis. We observe in E. indicus (Dauntela var.) somewhat similar profile and front views
(Fig. 1204) correlated with the very powerful muscular attachments and greatly enlarged tusks. Differing from
either Loxodonta or Elephas are the lowered orbits of Hypselephas.

With the exception of tliis distinctively low position of the orbits the front view of the cranium of Hypselephas
hysudricus is observed (Fig. 1204) to bear a closer resemblance to that of the broad-narial variety Elephas indicus

SKULLS OF ASIATIC ELEPHANTS (AFTER FALCONERI

All figures one-twentieth nature! siie

E. INDICUS Ref, NDICUS Ret.

PI. XIII A, Fig. 6 Falc, IB47. PI. XIII B, Fig. e

F.lc . 1847.

HYSUDRICUS Re E. HYSUDRICUS Ref.

Falo.. 1845. PI. IV Falc. 1845. PI. V, Fig. 1 (fev.>

Comparison of Two Crania ok Hypsklephas hysudricus with Mukna and Dauntela Varieties of Elephas indicus
The juvenile cranium 'E. hysudricus' ref. (Falconer, 1845, PI. vi, figs. 1 and 2) is regarded as belonging to Hypselephas hysudricus
Fig. 1204. Comparison of EUphas [Hypselephas] hysudricus (Lower Pleistocene of India) with Elephas indicus (Mukna var.) and E. indicus (Dauntela
var.) living today in India. After Falconer and Cautley, 1846 [1845 and 1847]. One-twentieth natural size.

The two adult crania of Elephas [Hypselephas] hysudricus reproduced in the above figure from Falconer and Cautley, 1845, PI. iv, PI. v, Fig. 1, and 1847,
PI. XLiii, Fig.XX. a, are distinguished from the crania of E. indicus (Mukna var.) and E. indicus (Dauntela var.) not only by much less depth and greater breadth
(bathycephaly), but by the extremely depressed position of the orbits which are placed barely above the roots of the grinding teeth, whereas in E. indicus the
orbits arc relatively elevated or directly opposite the occipital ('ondyle and in E. hysudricus entirely below the level of the occipital condyle.

This deeply depressed position of the orbits is also observed in the juvenile cranium, figured to the same scale above as 'E. hysudricus' ref., Falc, 1845,
PI. VI, Fig. 2, referred in this Memoir to Hypselephas hysudricus.
Observe that this marked lowering of the orbits also characterizes tiie juvenile cranium in the American Museum, Brown Collection (Amcr. Mus.
19866) reproduced in figures 1213 and 1214, giving Hypselephas hysudricus not only an entirely different aspect, as shown in the restoration (Fig. 1174), but
in our opinion precluding the possibility of the derivation of the cranium of either variety of E. iiulicus from the cranium of E. hysudricus.
 ^ : :

1350 OSBORN: THE PROBOSCIDEA

bengalensis (Dauutela var.) than to the narrow-narial variety Elephas indicus ceylanicus (Mukna var.). Both in

frontal and lateral aspect the two skulls referred to E. hysudricus by Falconer, and beautifully figured in his Atlas

of the "Fauna Antiqua Sivalensis," bear a superficial resemblance to the frontal and lateral aspect of the cranium
of E. indicus (Dauntela var.). This resemblance, however, is certainly a convergence, because the low, narrow

grinding teeth of the type of Hypselephas hysudricus are wholly different in proportion from those of E. indicus.
The molars exhibit some resemblance or analogy to the extinct Palxoloxodon of Africa in (1) their loxodont sinus,

and (2) their long, narrow proportions.

The juvenile cranium (Figs. 1213, 1214) of 'Elephas hysudricus' [

= Hypselephas hysudricus] moreover bears

a somewhat close resemblance to the juvenile cranium of E. indicus (Fig. 1180A) at the time when the fourth
upper and lower milk molars, l.Dp4, l.Dp*, are in use; the extreme elevation or hypsicephaly of the skull at this

stage seems to be an adaptation to the large, elongate, permanent grinders, M'"", Mi.2.

From cranial and juvenile characters alone we are thus inclined to regard Hypselephas hysudricus as an early
offshoot in Lower Pleistocene times of the main stem which gave rise to Elephas indicus of recent times. It is

interesting to note that the progression of the ridge formula in the case of these two species is similar to that which

separates other Upper Pleistocene and recent types, namely

Recent: Elephas indicus, M3 g //g . Total number of ridge-plates (Dp 2-M 3) :

f|.

Lower Pleistocene: Hypselephas hysudricus, M3 r-i^t^. Total number of ridge-plates (Dp 2-M 3)
i8 +
!! + •

ELEPHAS HYSUDRICUS /?e/

yiz Nat. sija /fftcr Faiconcr Piaies 4 "^i S
— -"v-

I''ig. 1205. Referred adult male cranium of Ekphas Hypselephas] hysudricus,

[ after Falroncr and Cautley, 1816 [ISl.J, PI. iv, and I'l v, lig. 1|

Brit. Mas. M.3I09. Reproduced one-twelfth natural size.

 THE ELEPHANTINE: HYPSELEPHAS 1351

Grinding Mechanism.— The relatively low-crowned grinding teeth in the upper and lower jaws, with maxi-
mum elevations of the ridge-plates of M^ (138-137) and of M3 (145) are correlated with the two very important
and distinctive characters of the cranium and jaws, which are clearly displayed in figure 1205, reproduced from
Falconer and Cautley, 1846 [1845, PI. iv, and PI. v, figs. 1 and 2], Brit. Mus. M.3109. The jaws seem to be
relatively shallow and primitive with prominent rostrum. Since the crowns of the superior grinding teeth are
relatively low, the very prominent orbits with their rugose borders are placed immediately above the roots. We
observe that this cranium an adult male, with primitive lower jaw associated; premaxillaries contracted, as
is of

in Elephas indicus; inframaxillary rostrum very prominent; molar ridge-plates probably ^^~ (compare Figs.
1204 and 1213).

Falconer's description (1867, p. 6, and 1868, Vol. I, pp. 425 and 426) is as follows:

Plate IV. Elephas Hysudricus (Fale. and Caut.),

from the Sewalik hills. Front view of skull, one-fifth
nat. size. This fine specimen was purchased from Con-
ductor Dawe.
Length
— B. M.
cranium from the protuberances of
of the
the occipital to the broken tip of left incisive, 45. in.
JB
Length from broken occipital condyles to anterior
border of alveolus, 28. in. Vertical height of head,
from broken condyles to the pyramidal bulge of sinci-
put, 26. in. Vertical height from surface of occipital
to the tip of the nasals, 27.75. Extreme width of the
head restored on left side, 38.5 in. Width at narrow-
est part of forehead between zygomatic fossae, 10.5 in.
Width of naso-maxillary fissure, 18.5 in. Depth from
tip of nasals to anterior margin of naso-maxillary fis-
sure, 3.5 in. Depth of rami of na.so-maxillary fissure,
4. in. Width between middle of the orbits, mesial,
26. in. Greatest width of zygomatic fo.ssa, 12. in.
Depth from hollow of frontal to condyles, 20. in.
Depth from posterior border alveolus to margin of
naso-maxillary fissure, 21.5 in. Length of alveolus
of last grinder, 10.5 in. Depth of hollow of frontal
below mesial plane, 4.5 in. Extreme width of alveo-
lus, 4.75 in. Width of incisive sheath in front of the
alveolus, 18.5 in. Tran.sverse diameter of the left tusk,
7.5 in. Antero-post. of the left tusk, 7.75 in. Depth
below mesial plane of the occipital hollow, 8.5 in.
Width of bottom of occipital hollow, 5.75 in. Depth
of posterior bulge of the cranium from the occipital
bone to surface of zygomatic fossa, 15.5 in. Least
width at back part of cranium behind the alveoli, 8.5
in. Depth from posterior broken surface of condyle
to the posterior border of the alveolus, 19. in. Depth
of infra-orbital foramen, 2.5 in. Transverse diameter
of foramen, 1.75 in. Length of infra-orbital canal, 6. in.
Depth of the left orbit, 6.75 in. From anterior mar-
gin auditory foramen to anterior border of the orbit,
20. in. Vertical diameter auditory foramen, 1.5 in.
Depth of the fossa between incisive sheaths at the
top of it, 6.5 in. Width across fossa, 3. in. Depth of
the naso-maxillary vault, 12. in. Depth of skull from
posterior end of socket to the orbit, 22.5 in.
o
 THE ELEPHANTINE: HYPSELEPHAS 1353

ELEPHAS HYSUDRICUS
AMHERST MUSEUM

Htpselephas HYSUDRICUS IN Amherst Museum, Right Lateral, Palatal, and Frontal Aspects
Compare with
Falconer's cranium (Fig. 1205), also with palate of Plalelephas platycephalus (Fig. 1219) below
Fig. Female cranium referred to Hijpselephas hysudricus collected near Kullu, a district of the Punjab, lying north and northwest
1210.
of Simla, !)>•
M. M. Carleton between 18."j4 and 1861 and presented by him to the .\mherst Museum. The skull was originally covered with very coarse
conglomerate,
possibly indicating its horizon as the Plei.stocene Boulder Conglomerate.
The third l<-ft superior grinder, I.M^ length ZOr, mm., width 95 nmi., displays 17+ ridge-plates, of which the anterior 6 are worn. It thus resembles
an
„.
I.M' of Hypselfphas hysudricus ref. (Amer. Mus. 19803- I'^ig. 1203 of the pre.sent Memoir) and establishes its specific
position as llypnelephas hymdricus rath7ir
than ArchidiskodoH planifions.

by relatively small tusks; the less rugose orbits, in a similarly depressed position, are seen to lie (Fig. 1210, left)
directly upon the premaxillary sockets of the tusks instead of being raised above the tusk sockets, as in E.

indicus (Mukna var.) and E. indicus (Dauntela var.). The agreement of the crania in the British Museum
with the imperfect cranium in the Amherst Museum in this very exceptional position of the orbits tends

further to demonstrate that Hypselephas hysudricus belongs in a totally distinct phylum or hne of descent
which is not to be regarded as ancestral to the collective Elephas indicus type.

In palatal view the extreme brachycephaly of the Hypselephas hysudricus cranium is clearly displayed (Fig.
1209) in contrast to the more elongate and less widely expanded palate of Archidiskodon planifrons (Fig. 1208) and
to the extremely primitive and elongate palate of Plalelephas plalycephalus (Fig. 1207). We observe that in
Hypselephas hysudricus (Falc, 1845, PI. v, fig. 3) the tusks are vertically crowded close to the second superior
molars, that the space between the third molars and the occipital condyles is extremely short, that the posterior
nares open close to the occipital condyles, that the occiput is enormously broadened.

In contrast the tusks oi Archidiskodon planifrons (Fig. 1208) point obliquely forwards and downwards instead
of directly downwards, there is a very wide space between the posterior grinding teeth with their widely set ridge-

plates and the occipital condyles, the paroccipital palate is much narrower than in Hypselephas hysudricus.
Finally, in Plalelephas platycephalus (Fig. 1207) we observe a wholly distinct type of cranium, much more primitive
than that of vl. planifrons and far more primitive than that of Hypselephas hysudricus.

Juvenile Crania with Dentition

Figure 1204 above (p. 1349) illustrates theextreme contrast between the juvenile and adult crania of Hypsele-
phas hysudricus as reproduced to a one-twentieth scale from Falconer and Cautley 's beautiful lithographic drawings.
Falconer's material (1868, Vol. I, p. 426) included a "Perfect small head from the Geol. Soc. Museum, with the
 —

1354 OSBORN: THE PROBOSCIDEA

second [Dp'] and third [Dp*] milk molars and first true molar in germ." This small head is reproduced herewith in
figure 1212, also in figure 1213 (after Falconer and Cautley, PI. vi, figs. 1, 2). Another figure (Falconer and Cautley,
1846 [1845, PI. VI, fig. 3]) is that of the under surface of a young skull (see Falconer, 1868, Vol. I, p. 426):
"Fig. 3.^ Elephas Hysudricus, under surface of young skull. This specimen agrees in age and characters with that
shown in figs. 1 and 2, except that the third [Dp"] milk molar has 8 principal ridges, with a front and back heel,
instead of 7 as in the other. —^B.M. Length of second [Dp^] milk molar, 2.2 in. Width of second [Dp^] milk molar,
1.6 in. No. of plates about 5. Length of third [Dp*] milk molar, 4.3 in. Width, 2. in. Interval between second
[third] teeth, 1.2 in. Between third [fourth], 2.1 in."

In the American Museum collection is a shghtly more mature juvenile cranium (Amer. Mus. 19866) shown
in our figures 1211, 1213, and 1214, with the 8-9 ridge-plated LDp" in full use, and the l.M' just coming into use
(Fig. 1214).

The American Museum also has another specimen, an isolated r.M^ (Amer. Mus. 19915), which from its

extreme narrowness was mistaken by Osborn (Osborn, 1929.797, pp. 22 and 23) for a third lower molar and
erroneously made the type of Elephas platycephalus angustidens; further examination proves that this is a right
second superior molar, r.M=, of Hypselephas hysudricus Falconer and Cautley, with }^-12->2 ridge-plates ( = 14
ridge-plates), agreeing with the typical M^ ridge-plate formula, as shown in our revised figure (Fig. 1215).

With these specimens we are now enabled to measure, describe, and portray completely the characters of
the palate, jaws, and dentition, including M' M^ of the juvenile Hypselephas hysudricus.
Molar Crown Characters. —The first striking pecuUarity of the young American Museum skull (Fig.
1214) is: 1) The extreme length and narrow width of the molar crowns, the successive indices of Amer. Mus.
19866 being:
Amer. Mus. 19866
 Af JS866

Fig. 1213. Juvenile Crania of Hypselephas hysudricus, British and American Museums
One-sixth natural size. Compare figures 1211, 1212, 1214

(Left and center) British Mu.seiim cranium, a younger stage with 4)2 ridge-plated Dp' and K-7-K ridge-plated Dp'' in use. Same cranium as figure 1212.

(Right) American Museum cranium (Amer. Mus. 19866), somewhat older, Dp'' shed, the 8)^-9 ridge-plated Dp'' and the anterior portion of the 10-11
ridge-plated M' in use; M- in germ.
(Left) After Falconer and Cautley, 1846 [1845, PI. vi, figs. 1, 2). A referred juvenilecranium presented to the British Museum by the Council of the
Geological iSociety (and now bearing the catalogue number, Brit. Mus. M.3114). This specimen is de.scribed by Falconer (1867, p. 6, and 1868, I, p. 426) as

a "Perfect small head from the Geol. Soc. Museum, with the .se<ond [third] and third [fourth] milk molars, and first true molar in germ," namely, Dp' with
4/4 ridges. Dp'' with Observe the extreme hyjisicephaly and fore-and-aft compression, the plane forehead, the occiput without the superior ex-
)'<r1-Yi ridges.
pansion .seen in the adult, and the extremely narrow premaxillary ro.strum.

(Right) Reconstructed juvenile cranium (Amer. Mus. 19866), somewhat older, because the 8)2-9 ridge-plated Dp'' is still in complete u.se, the 10-11
ridge-plated l.M' is partly in u.se, the l.M^ is in germ. Compare the Barniim Brown table of measurements (p. 1348), also figure 1214 of the same cranium
on a one-sixth scale, and figure 1211 on a smaller one-twelfth scale.
 ''""'^
ifiit HVSUDRICUS AV

orthogonal view

Juvenile Cranium and Jaws of Referred Hypsblephas hysudricus

Fig. 1214. This imixirtant juvenile .s[X!cimen (Amer. Mus. 1986()) was collected by Barnum Brown in 1922 in the Upper Siwaliks of India, not in situ but
in erosion material 'below the conglomerates,' 3 miles west of Chandigarh. R(^produeed h(^rewith one-sixth natural size.

(Upper left) Palatal view .showing Dp^ with 8)2 to 9 ridge-plates in full use; the 11 ridge-plated l.M' is obliquely placed showing the slight wear of the
apices of ridge-plates 1 to 6. Observe the extreme .simplicity of the five comlels composing the ridge-plates of l.M' —a very primitive condition. Compare
figure 1203.

(Lower left) Detailed view of crown of LDp"" with 8W to 9 ridge-plates, r.Dp4 with 9 ridge-plates, and r.Mi.

(Upper right) Left lateral view of cranium showing the germ of l.M^; the partly restored l.M' in situ with 11 ridge-plates (to be I'omparcd with the crown
view of .same in upper left) exhibiting the very simple (condition of 4 to 5 eonelets; and the worn 8'4 ridge-plated I.Di)''.

(Lower right) Left lateral aspect of mandibular ramus with very prominent rostrum.

(Lower center) Coronal a.spect of the same juvenile jaw (Amer. Mus. 19866); Dp4 with 8)4-9 ridge-plates, typical of llypselephas ItijsuUricii.i; r.Mi with
3 worn ridge-plates. Observe the elongation of the rostrum.

1356
 THE ELEPHANTINE: HYPSELEPHAS 1357

2) A second distinction, as clearly displayed in l.M'

(Fig. 1214), and in r.M= (Fig. 1215), is the compression

and simplicity of the conelets on the summits of the

crowns; in l.M> the three anterior ridge-plates bear 5

conelets each, the eight succeeding ridge-plates bear 4

conelets each; in r.M- the six anterior ridge-plates bear

5 conelets each, the succeeding seven to thirteen ridge-

plates bear 4 conelets each.

3) A third distinctive character (Fig. 1215) is the

wide separation by cement filled valleys of the seven

anterior ridge-plates.

4) A fourth distinction is the relatively uniform

height of the ridge-plates, the maximum at the 5th plate
in r.lVP (Fig. 1215) being 102 mm.

Elephas platycephalus angustidens ( = Syno-

nym OF Hypselephas hysudricus). The type of this —
subspecies was collected by Dr. Barnum Brown in 1922
three miles west of Chandigarh, Siwalik Hills, India,
'below conglomerates' (see Osborn, 1929. 797, p. 22, also 1
\^
ELEPHAS (hypselephas) ANGUSTIDENS, Type

fig. 22, p. 23). It was at first regarded by the present

author (cf. p. 1354 above) as a third inferior molar
of the left side, I.M3, because of its extreme narrow-
ness. On further study it proved to be a second supe-
rior molar of the right side, r.M=, referable to Hypsele-
phas hysudricus (Fig. 1215).

Hypselephas hysudricds Outer-

Fig. 1215. A second superior molar of the right side, r.M-, with 14 ridge-
plates, of which the three anterior (1-3) are partly worn. One-third natural
size. This is the molar tooth originally selected by Osborn (Osborn, 1929.797,
y^ Mit. sije.
pp. 22, 23) as the type of Elephas platycephalus angustidens; erroneously
interpreted as a third inferior molar; the simple ridge-plate structure cor-
responds exactly with that in the young cranium of Hypselephas hysudricus
ref. (Amer. Mus. 19866—Figs. 1213, 1214).
Superfamily: ELEPHANTOIDEA Osborn, 1921

Family: ELEPHANTID^E Gray, 1821

Subfamily: Elephantin^e Osborn, 1910

Genus: PLATELEPHAS Osborn, 1936

Original reference: Osborn, 1934.926, p. 285 {nomen nudum); Vol. I of present Memoir (1936), p. 12, and PI. xi.

Genotypic species: Elephas platycephalus Osborn, 1929.797, p. 21.

Generic Characters. — Cranium relatively elongate, dolichocephalic, and platycephaUc ; occipital

condyles not greatly elevated above level of grinding surface of molars deeply indented supra-occipital ;

border. Premaxillaries greatly elongated in front of molars, somewhat divergent; tusks unknown.
Orbits large, elevated, near frontal profile. Grinding teeth imperfectly known, relatively low, ridge-
plates directly transverse, as in Elephas, no rudiment of 'loxodont sinus.' Ridge-plate formula, as far as
known, M 3 ^^.
Habits unknown, probably Uke those of Elephas indicus. Very primitive in cranial
structure and in the broad, depressed ridge-plates.

This unique cranial type representing a hitherto unknown and very primitive stage was found in a separate
mass of consolidated gravel which had apparently been washed down from an original boulder conglomerate bed
and into a shallow region bordering Amilee Creek, near Siswan, Simla Hills, India. While not found in situ, it is

apparently of the same Lower Pleistocene age as the 'Boulder Conglomerate formation' above.

clay

tlay

c^t^s
cor.gTome'*

Fig. 1216. Upper Pliocene and Lower Pleistocene Strata near Siswan, India
After photographs by Bariuim Brown

(Left) Cranium of Platele jihaa plalt/rephalux as fovintl on the banks of (Right) Alternating Variegated clays' and 'conglomerates' typical of the
Amilee Creek, near Siswan, India, not in nitu. 'Conglomerate' matrix was at- region at Siswan, India, near which was found much of the material col-
tached to the skull, indicating that it had been deposited above the 'varii'gated lected by Dr. Fiarnum Brown in 1922.

clays,' as shown to the right. With the exception of a single skull of Ros- The upper levels, ('onsisting of 'variegated I'lay.s' alternating with 'con-
elaphns, no fos.sils were found in the 'conglomerates.' glomerates,' are below the true Boulder Conglomerate zone.

1358
 :

THE ELEPHANTINE: PLATELEPHAS 1359

In brief, the proportions of this cranium, as compared with those of the middle-aged Elephas indicus, may be
described as dolichocephalic rather than brachy cephalic, as platycephalic rather than hypsicephalic, as longirostral
rather than brevirostral — all of which are primitive characters. The present phyletic reference is to the subfamily
Elephantine.

Platelephas platycephalus Osborn, 1929 hypsicephalic, the adult cranium of Platelephas platycephalus is

Figures 1174, 1207, 1216-1219 relatively platycephalic. The breadth-length index, as measured
From near Siswan, bed of Amilee Creek, Simla Hills, India. Upper from the summit of the occipital crest to the extremity of the
Pliocene or Lower Pleistocene. premaxillaries, compared with the greatest breadth across the
Elephas platycephalus Osborn, 1929. "New Eurasiatic and orbits, is 60, indicating that this cranium is also relatively dolichoce-
phalic.
American Proboscideans," Amer. Mus. Novitates, No. 393, Dec.
24, 1929, pp. 21 and 22. Type.— Cranium with M' of both The principal measurements of the type are as follows
sides partly exposed. Amer. Mus. 19818. Collected by Barnum
Cranial Measurements of Platelephas platycephalus Type
Brown Horizon and Locality.
in 1922. — Upper Pliocene or
Lower Pleistocene. Near Siswan, bed of Amilee Creek, Simla Hills, Length
India. Type Figure.— Osborn, op. cit., 1929.797, p. 22,
Occipital condyles to extremity of premaxillaries 1085
fig. 21.
Postoccipital crest to extremity of premaxillaries 1140
Type Description. — "Specific Characters. —Cranium of

very primitive elephantine affinity, low, flattened; orbits widely Breadth across
separated from occiput ;
premaxillary rostrum somewhat broaden- Supra-orbital proces.ses 680
ed, resembling that of Elephas; posterior nares deeply indented; Occipitotemporal region (Fig. 1219 A3) 665
occipital condyles on relatively low plane, not greatly elevated Orbitofrontal region 332
above grinders; relatively long and narrow cranial proportions. Extremity of premaxillaries 522e
Cranium widely different from the elevated Elephas [Hypselephas] Widest portion of zygomatic arches 800e
hysudricus or the greatly elevated Elephas indicus crania. Ridge-
Height
plates of type molars fractured or absent. Estimated ridge-plate
formula: M3^-^." From level of supra-occipital prominence to maxil-

Discovery. —This animal appears to belong to a primitive lary borders of grinders 616
cannot at Occiput, condyles to occipital crest 425
stage in the evolution of the Elephantinse, although it

present be regarded as ancestral either to Hypselephas hysudriciis

Alveolar border of M' to top of frontals 555
Grinding surface of M' to midfrontal region 605e
possibly of the Boulder Conglomerate formation, or to Elephas
and Recent time.
indicus of closing Pleistocene
Length
Both the cranium and the grinding teeth arc profoundly
Occipital condyles to posterior borders of third
distinct from those of Archidiskodon planijrons, a species so very
grinder, r.M' 342
abundant in the lower levels of the Pinjor horizon; thus Platelephas
Facial length, front of M' to tips of premaxillaries 507
platycephalus cannot be related to the genus Archidiskodon, as
Alveolus of left superior grinder
clearly shown in comparative views (Figs. 1207, 1208). The
Lengtii 220e
relatively narrow premaxillaries and closely appressed superior
Width 90e
tusks forbid its relationship to Palseoloxodon namadicus of the
Alveolus and crown of right superior grinder
much more recent Middle [to LTpper] Pleistocene of India. Conse-
Length of r.M^ (preserved portion) 175
quently it seems best to place it temporarily in the true Elephan-
Width of r.M= 87
tinoe phylum.
As shown especially in figures 1217 and 1218, the cranium is Transverse diameter of incisive alveoli 120-135e
widely different from that of Stegodon pinjorensis or of S. insignis-
ganesa. The specific name platycephalus refers to the highly char- The first distinctive character is the height from the grinders
acteristic and primitive lowering of the fronto-occipital profile and to the midfrontal region, namely, 605e mm., as compared with
the placing of the occipital condyle only slightly above the hori- 645 mm. in the cranium of Elephas indicus. The second distinction
zontal level of the maxillary border of the superior grinders, as of importance is the primitive elongation of the cranium, the
shown in figure Whereas the juvenile and adult Hypsele-
1219A. basicranial region measuring from the occipital condyles to the tip
phas hysudricus crania (Figs. 1213, 1205, 1218) are markedly of the premaxillaries 1085 mm., as compared with the maximum
1360 OSBORN: THE PROBOSCIDEA

occipital or temporal width of 665 mm. The third distinction is tho The premaxillaries are distinguished from those of Palaeoloxo-
marked separation of the orbits and the width across the supra- don namadicus by less expansion, the width being 522e mm., as
orbital region, 680 mm., as compared with the relatively closely compared with 857 mm. in the Pignataro skull of Hesperoloxodon
placed orbits of E. indicux, 561 mm. nntiqutis italicus; they arc, however, somewhat broader and

Mid-cayitai sae77oi TLsA, TaSTh. condyle seittan ^> -^<

J^oruontal occifiitai sectia.

Tbrijn^n m^nn^'v.
Mlsu/ median

Ty^ntol section
A": B
f 1 All y,6 nafUra/ Slje
STEqoDON PINJORtNSIS Tyfie AmerMui. /977S
B. ELEPHAS PLATYCEPHALUS Typ. Amur. MU5./96/8

Comparison of the Types of Platblbphas platycephalus and Steqodon pinjorensis

Fig. Plntekphas plalt/cephalus (
1217. ) type oranium in vertical section (right) has a totally different fronto-paroccipital profile from Stegodon

pinjorensis ( ), when the oeoipital condyles are placed at the same level.

The midcranial section (center) differs much less, becau.se of the deep indentations in the fronto-occipital condyles, as shown in figure 1219 .\2.
The horizontal occipital section (left) of Htegodon pinjorensis ( ) is shown to be much broader than that of I'Uileleptms platyccplialiis; the na.sal
sections are about the same breadth; the frontal section of S. pinjorensis is much broader than that of P. platycepluilus.

Comparative Measurements of Crania of Platelephas relativelymore expanded than the cranium of Elephas indicus, in
PLATYCEPHALUS AND ElePHAS INDICUS which they mea.sure transversely 414 mm. A fourth distinction is
the elongation of the premaxillaries in front of the third upper
Elephas Elephas indicus
platycephalus (Amer. Mus.
(Amer. Mus. Dept. Mam.
19818) 54261) Male

Width across temporals 665 mm. 710 mm.

Maximum width of occiput 680 755
Maximum width across occipital
condyles 216 214
Height of occiput from condyles to
occipital crest 425 440
Width across supra-orbital processes 680 561
Width across extremities of pre-
maxillaries 522e 414
y^6 /yatura/ site
Width of narial opening (inside) . . . 435 323
Length, front of right molar to tip of
premaxillaries 507 325 A. £. PLATYCEPHALUS ry/>e A^M.ISB/B \ '".

'
B- E. HVSUDBICUS A'f/' y^/ter falcmer --,__ '',

Length of premaxillaries to occipital

condyles 1085 925
Fig. 1218. Right cranial profile of Platelephas platycephalus ( ) and
Height of cranium, grinders to mid- of Ilypselt'phas hysudricus one-sixteenth natural size.
( ),
frontal region 605c 645 This figure illu.slrales the primitive platyceplialy of the Platelephas
Width across midtemporal region . . 332 334 platycephalus type as compared with the relative hypsicephaly of the Ilypsele-
plias hysudricus type. Both crania may be regarded as adult and fully de-
Length of tusks (outer curve) 2245
veloped. Observe that /'. platycephalus does not in the least resemble the
Transverse diameter of tu.sk at juvenile profile of //. hysudricus (Kig. 12i;!); it is much more primitive (cf.
emergence, i.e., mdth of alveoli 120-135c 117 Figs. 1207, 1209).
 THE ELEPHANTINE: PLATELEPHAS 1361

grinders, the distance being 507 mm., as compared with 325 mm. the anterior nares of the space between the premaxillary sockets
in E. indie us. and the tusks; (4) and finally the great prominence of the orbital
Special features are: (1) Very deep excavation of the superior rims, indicating that the eyes were set very wide apart.
border of the occiput, totally unlike that of either Archidiskodon Platelephas plalycephalus, therefore, is totally different in
or Elephas; (2) the relative breadth of the anterior nares, measur- profile as well as in superior and palatal aspects from cither Archi-
ing transversely 435 mm. ; (3) the very deep excavation just below diskodon planifrons or Elephas indicus.

Type Cranium of Platelephas platycbphalus

Fig. 1219. Four aspects of the type iranium of Platelephas plalycephalus (Amer. Mus. 19818), discovered by Barnum Brown in 1922 above the Pinjor
horizon of the Upper Siwaliks, 'below conglomerates.' One-twelfth natural size. Compare Osborn, 1929.797, p. 22, fig. 21.

A, Left lateral aspect, exhibiting orbits, auditory openings, and occipital condyles on practically the same level.

Al, Palatal view, exhibiting slightly broadened premaxillary rostrum, maxillary prominence containing fractured M' with 16)2 ridge-plates, deeply in-
dented posterior nares, and relatively low plane of occipital condyles.

.\2, Superior aspect, exhibiting moderately broadened premaxillarien, flattened fronto-occipital plane with deeply indent<^d supra-occipital border w itliout
rugosity, and relatively long and narrow cranial proportions.

.\3, Posterior aspect, exhibiting moderately small, elevated occipital condyles when cranium is placed in exactly the same plane as (.\). Compare with
widely different cranial profile of Hypselephas hysudricus (Fig. 120.5) and of E. indicus ceylanicus or E. indicus bengalensis (Fig. 1170), in which the occiijital
condyles are elevated above the level of the orbits.
 APPENDIX TO CHAPTER XX
[In a recent Memoir (1936) by Pere Teilhard de Chardin and Dr. C. C. Young, "On the Mammalian Remains
from the Archaeological Site of Anyang," Pal. Sinica, (C), XII, Fasc. 1, pp. 1, 52, and 53) they have given their
views regarding the existence in historical times of the wild Indian elephant in China, from which the following has
been extracted. The reader is also referred to the map on page 1594 which gives the range of Elephas indicus,
its varieties and subspecies, as far as known. —Editor.]
Close to the modern city of Changteho (N. Honan) are buried under the mud of the Chinese maritime plain the remains
of Anyang, the latter city being the old capital of the —
Shang dynasty (Circa 1400 B.C. 1100 B.C.).
Remains Elephant are not uncommon in Anyang (fragmentary young skulls and limb-bones). We only have at our
of
disposal an incomplete lower molar of a young individual, probably a tooth number 3. Six lamellae are preserved, occupying
a length of 68 mm. Thickness of one lamella 5 ram. breadth 48 mm. The lamellae are widely separated and set rather oblique-
;

ly.
As much as we can judge from this unsatisfactory specimen, we most probably are dealing here with an Elephas indicus,
and not at all with E. primigenius, as supposed by Matsumoto (1916), basing on a centrum of vertebra and Hopwood (1934),
using fragmentary molars. This reference of the Anyang Elephant to the Mammoth group was only possible when the deposits
were still held as being of Pleistocene age.
The presenceof E. indicus in Honan once more raises the question as to how far north this form was living in China, and
as to how long it lasted there, in historical times. Of course, palseontologically, we have no evidence to .suggest as yet that the
species ever existed in China proper. Its closest ally, E. namadicus, has never been found in the post-Loessic deposits. And no
Elephas (but only Stegodon) occur in the Szechuan Pleistocene fissures. But historically it has been stated in some Chinese
texts which were recovered in Anyang that 'the king hunted and killed an Elephas(?)'. A critical discussion of those texts was
given by Dr. Chang (1926). Dr. Chang concluded that there are no historical evidence proving the presence of wild Elephant
and Rhinoceros in N. China since historical times. This view seems to be so far the most conservative and the best supported
by facts.
We shall therefore admit here that the Anyang Elephant, just like the Tapir of the same locality, represents a southern
type imported to the city as a tribute, possibly for hunting purposes. It has to be noted in this connection that these exotic
. .

animals were brought and kept alive. Both in the case of the Elephant and the Tapir, our fossils belong to young individuals
represented not only by some ivory but also by perfectly useless bones.

1362
 Chapter XXI

NOMENCLATURE OF THE PROBOSCIDEA

History of nomenclature of the mastodon and the mammoth. Chronologic lists of superfamilies
(1921-1935), FAMILIES (1821-1937), subfamilies (1838-1937), genera and species (1735-1939).

1. History of nomenclature of the Mastodon and the 4. List of subfamilies.

Mammoth. 5. Genera and species of the Proboscidea in the
2. List of superfamilies. order named by the authors in the original
3. List of families. descriptions.

Professor Osborn regarded nomenclature as "the tool rather than the master of palaeontologic thought" and
believed that "no technical principles should override the work of the early discoverers and naturalists." As
repeatedly observed in the present Memoir (see especially Vol. I, Chap. I, pp. 5-13, also Vol. II, p. 1173),
he thought it "impracticable in palaeontology to apply all the principles of nomenclature established in zoologij
and botany, because the classification of the imperfectly known fossil forms is ever changing with our increasing
knowledge of origins, adaptive radiations, and phyletic successions. Such mutabiUty does not disturb the nomen-
clature of hving animals and plants, in which priority of adequate description, figure, and definition is the chief
concern of systematists."

Doctor Hopwood in his Memoir of 1935 on the "Fossil Proboscidea from China" (p. 11) states that he has
"attempted to make of the Rules [International Rules of Zoological Nomenclature] a useful servant, rather than to
allow them become a bhnd, unreasoning, master." The extent of the difficulties encountered in Proboscidean
to
palaeontology may be judged from the accompanying nomenclature of the Mastodon and the Mammoth.

This chapter was not completed by the author, but has been compiled by the editor from materials left by
Professor Osborn, under the conditions mentioned in the "Publication Note" on page viii of Volume I.

1. HISTORY OF NOMENCLATURE OF THE MASTODON AND THE MAMMOTH

1788 The genus Mammonteus Osborn (1924.633, p. 2; Mammonteum Camper, 1788, p. 251) is of doubtful
validity ; in fact, it is possible that Professor Osborn would have abandoned it in his final revision of the

present Volume and adopted Mammuthus Burnett, 1830, as the reader may conclude from the follo\ving
account of the nomenclature of the northern or woolly mammoth and of the true mastodon {Mastodon
americanus) the descriptive literature of which
, is so involved that the history of one is inseparable from
that of the other. Consequently the subject is treated chronologically, dating from 1788 to the present
time.

The American Mastodon (Mastodon americanus)

1792 Kerr.— In 1792, p. 116, Robert Kerr proposed the name Elephas americanus for tusks and grinders
found at Big Bone Lick near the banks of the Ohio River, Boone County, Kentucky.

1797 Blumenbach. — (1) In 1797 Blumenbach in his "Abbildungen Naturhistorische Gegenstande," em-
ployed the term Ohio-Incognitum, which first occurs on the back of the title-page following other specific
names, i.e., "11. Simia troglodytes ... 19. Backenzahne von fossilen Ohio-Incognitum, und von den beiderlei
Gattungen des Elephantengeschlechts." The name again appears opposite figure 19A.

1363
 : :

1364 OSBORN: THE PROBOSCIDEA

In a letter from C. W. Andrews to Professor Osborn (June 6, 1922) he quotes C. Davies Sherborn as
follows: "The words 'Ohio incogniium' are not used in a generic or specific sense; they mean simply 'the

Ohio incognitum' as is easily seen by the second page of the description, and are quite invalid." (See
Article 25 of the International Rules of Zoological Nomenclature.)

1799 (2) Subsequently (1799.1, p. 698) Blumenbach in his sixth edition of the "Handbuch der Natur-
geschichte" assigned the name Mammut ohioticum to the same animal. In the French edition of the
1803 "Handbuch" (translated in 1803 by Soulange Artaud under the supervision of Blumenbach) mammout
Ohioticum appears on page 408, also Ohio-Incognitum occurs under the figure 19A, first used in the 1797
"Abbildungen" of Blumenbach (see Vol. I of the present Memoir, caption to fig. 113, where it is stated by
Professor Osborn that it is "technically a type figure").

Dr. Edwin H. Colbert in a note to the editor (1937) makes the point that as Kerr, the author of
Elephas [
= Mastodon] americanus, did not include a figure in his description, we cannot be sure that
figure 113 just referred to is a figure of the type.

1806 CuviER.— It was 1806 before Cuvier applied "Le Grand Mastodonte" (1806.2) and "Mastodonte de
rOhio (1806.3) to the American Mastodon, which in 1817 (see below) he redescribed under the specific

designation of Mastodon giganteum.

Fischer de Waldheim. —^The following generic names were applied to the Mastodon by Fischer de
Waldheim
1808 Harpagmotherium.

1814 Mastotherium.

1816 Oken. — The final form Mastodon was published by Oken in 1816, p. 789, citing Cuvier's five classic

species ("Lehrbuch der Naturgeschichte," Theil III, p. 789).

1817 CuviER.— Although anticipated by Oken by one year (see above, 1816) in the use of the term Mastodon
(see 1817, "Le Regne Animal," I, p. 233, for description of Mastodon giganteum), Cuvier has been regarded
as the author of this genus, since he first used the French form Mastodonte in his description of 1806.

1868 Leidy. —Leidy (1868, p. 175)" was the first to use the name Mastodon americanus, embracing the
Mastodon ohioticus or M. giganteum of authors.

1869 Leidy in 1869 (p. 392) Usts Mamonteum Camper among the synonyms of Mastodon americanus.
1902 Hay. — A/ammwi Blumenbach was revived by Hay and adopted by many subsequent
(1902, p. 707)
authors. In his last work (1930, p. 623) Hay names the new subfamily Mammutinse to include the Masto-
donts, although he places it under the Elcphantidse without recognition of the family Mastodontidse.
1904- Trouessart. —The term Mammui Blumenbach was regarded by Trouessart (p. 600) as a misno-
1905 mer: "Le nom barbare de 'Mammut,' base par Blumenbach sur uiic orreur grossiere (ridentit6 du Masto-
donte de rOhio avec V Elephas primigenius) n'a aucun droit, malgre sa priorite, a etre substitue a celui dc
,

'Mastodon,' genre bien caracterise par Cuvier."

1920 Allen (excerpts from letter of Dr. Joel A. Allen to Professor Osborn, dated November 16, 1920)

111 your letter (October 22, 1920) you ask whether I consider Blumcnbach'.s name Mammui entitled to recognition.

1 certainly do, alt hoMKh with regret that such name was .ever proposed as a generic designation. Neither its barba-
. .

rous form nor the fact of his misconception that it was the mammoth renders it invalid .... All modern codes or rules
of nomenclature d('clare that such names are not to be rejected on the ground tiiat they are nonclassical, and that incor-
rectly formed cla.ssical names are not subject to emendation.
 :

NOMENCLATURE OF THE PROBOSCIDEA 1365

1935 HopwooD. —Doctor Hopwood substitutes Mastodon for Mammut, as stated in his "Fossil Proboscidea
from China," 1935, "To retain Cuvier's generic name instead of Bkimenbach's, is to follow con-
p. 42:
venience rather than rule. The generic term Mammut and the vernacular Mammoth are so much alike
that one or other should be suppressed in the interests of clarity. It is very much easier to discourage the
use of an unfamihar name than a well-known popular one, so that, whilst acknowledging Blumenbach's
prior claim, I have given the preference to Cuvier."

1936 Osborn's Conclusions in Volume I, pp. 6 and 7, of the Present Memoir. —For the reason that
Mammut (signifying 'earth-burrower' and suggesting the mammoth) was employed to describe the Masto-
don ; that if a vernacular name were to be accepted Ohio-Incognitum had priority, and lastly that inasmuch
as Cuvier's "Mastodonte" (Mastodon) had been used throughout the Uterature of the past century,
Professor Osborn committed himself to the genus Mastodon in the following citation from his Memoir
(Vol. I, p. 6): "Consequently to rob Cuvier of his clear conception of grinding tooth structure, which he
termed Mastodonte, and to substitute the barbaric term Mammut, signifying 'earth-burrower,' would be
gross injustice to the founder of vertebrate palseontology."

The Northern or Woolly Mammoth (Mammonteus primigenius)

1799 —
Blumenbach. In the same article in which Blumenbach described Mammut ohioticum, namely, in
the "Handbuch der Naturgeschichte," sixth edition (1799.1, p. 698), he also described Elephas primigenius
(p. 697), stating that there are abundant remains in Germany, but he mistakenly gives as an example

a skeletal specimen from Burg-Tonna (now in the Gotha Museum), which proves to belong to Elephas
[
— Hesperoloxodon] antiquus Falconer and not to the northern Mammoth.

1799 Cuvier. —In the autumn of the same year that Blumenbach described Elephas primigenius (1799),
Cuvier assigned the name Elephas mammonteus to the mammoth; subsequently, however, he adopted
Blumenbach's name E. primigenius.

The following is a list of the names assigned to the Mammoth {Elephas primigenius)

1830 Mammuthus Burnett, Quart. Journ. Sci. Lit. & Art, XXVIII, p. 352.

1837 Dicyclotherium E. Geoffroy St. Hilaire, Compt. Rend. Acad. Sci., Paris, pp. 119, 120, tig. 1.

1845 Mammonthetim de Blainville, 1839-1864, "Osteographie," p. 237.

1848 Cheirolites von Meyer, (in Bronn's "Handbuch einer Gesch. d. Natur.," Ill, Index Pal., p. 286).

1850 Synodontherium Costa, Palaeont. del Regno di Napoli, Pt. I, pp. 271-275, Tav. iii, figs. 1-4.

1888 Polydiskodon PohUg, Nova Acta Leop. Carol. Deutsch Akad., LIII, pp. 138, 252.

With the removal by Professor Osborn of Mammut Blumenbach from the valid genera of the Probo-
scidea, and the adoption of Mastodon Cuvier, there arose the question of the correct genus to which Elephas
primigenius, the northern Mammoth, should be referred. Professor Osborn chose Camper's name Mam-
monteum, 1788, which, like Mammut, had reference to a specimen of the American Mastodon and not to
 : —

1366 OSBORN: THE PROBOSCIDEA

the Maninioth. The revival of Mammonteum {Mammonteus) by Professor Osboni occurs in his article
"Parelephas in Relation to Phyla and Genera of the Family Elephantidae" (Osborn, 1924.633, p. 2) as

follows

1924 l^y iiiHiiy autliors all the generic phyla of the mammoths are .still referred to the genus Elephas. Such reference,
from ovu' present knowledge, is inconsistent with the fact that none of the mammoths contains the ancestral characters
of Elephas. (1) We thus revive the ill-defined name Mammonteus Camper for the Elephas primigenius phylum, which
Deperet and Mayet have traced back to the tapper Pliocene Elephas primigenius astensis of northern Italy and into the
Lower Pleistocene E. mericlionalis rromerensis of the Forest Bed of Cromer. Here this line terminates in the typical
E. primigenius of western Europe and onward into the progressive new subspecies Mam.monteus primigenius com-
prcssiis of North America.

1934 Preparatory to the final revision of the present Volume of the Memoir, Professor Osborn, in 1934,

Hopwood to the Mammuthus of Burnett (1830) as having priority

called the attention of Dr. A. Tindell
over Dicydotherium Geoffroy (1837) Doctor Hopwood adopted the name Mammuthus in his Memoir on
.

1935 "The Fossil Proboscidea from China," 1935, p. 98, from which we cite:

Remarks. —
With the progress of research on the Proboscidea, it has become ever clearer that the Mammoth
is from the Indian Elephant, which typifies the genus Elephas. Professor Osborn has sought to give expression
distinct
to this result by reviving an alleged genus of Peter Camj^er's and emending the name. If I do not adopt the same name,
it i.s because Camper was not referring to the Mammoth, and because he was using the word 'Mamonteimi' a.s a

vernacular. In the section 'De ossibus Mamonteis' Camper (1788, p. 259) u.ses 'Mamonteum' in an adjectival .sense
throughout. Not only so, but he also makes it quite clear that he is referring to an animal from America, and Pallas
contributes a foot-note on p. 261 in which he explains that whereas in Russia the term 'Mammontean bones' is common-
ly applied to the bones of Elephants found in superficial deposits, it has suited Camper to apply the name to bones found
in America. With so much evidence as to Camper's meaning, it is not possible to argue that the phrases on p. 251 of
the same work refer to the Mammoth as distinct from the Ele]>hant.
Several names ha\'e been applied to the Mammoth, but the first that is valid, in so far as it possesses a genotype,
appears to be Mammuthus Burnett, 1830. Under Elephant idse, Elephant-kind' he groups the
' following,

(iencra. Species.
I']le]ihas. Indicus. Indian Elephant.
Africainis. African.
.Maniniullius. Borealis. l'"ossil .Mamniotii.

Meridionalis.
Mastodon. Ciganteum. (iigantic.
Angustidens. Les.ser.

from this table il is clear that the word Mammoth in the sen.se generally accepted today, and did not apply it
lie u.-«'d

to the American Mastodon after the manner of the majority of English and American authors at the begiiuiing of the
nineteenth century, (rf. Cuvier, Ossemens foss., Ed. 1, Vol. II, Art. Sur le grand mastodonte), and that his Mammuthus
borealis is the equivalent of Elephas primigenius Blvun.

Doctor Hopwood then described a molar from Mongolia (regd. M10941) and fragment of another from
Honan (regd. M14102) which he refers to Mammuthus primigenius (Blumenbach).
The following excerpts from the correspondence between Doctor Hopwood and Professor Osborn are
self explanatory:

September 10, 1935. Professor Osborn wrote as follows: "Among the lOlephantidae 1 regret that you substituted
Mammuthus Burnett for Mammonteus Camper ... If wo stood on technicalities, the specific name Elephas primigenius
is based on a species indubitably belonging to Hesperolo.rodon antiquus Technical methods would compel us to alter . . .

the entire nomenclature of the fossil Proboscidea. As in religion the spirit rather than the letter is important."
 NOMENCLATURE OF THE PROBOSCIDEA 1367

October 19, 1935. Doctor Hopwood replied to Professor Osborn "With regard to the substitution of Mainmuthus
:

for Marmnonteus, I should not have done this had it been possible to show that Camper was referring to a European
fossil, but the evidence that the reference is to the American Mastodon is so strong that I had no other alternative. The
case of E. primigenius and H. antiqims is hardly parallel Blimienbach was naming a new species, and later authors have
;

misinterpreted him. The error is theirs, not Blumenbach's."

October 25, 1935. Professor Osborn replied at once as follows: "Your point that Camper was referring to the
American Mastodon when he used the name Mammonteus is a very strong one and I shall immediately refresh my memory
on this point, although I fear it may be difficult to correct it now."
October 29, 1935. Five days later he added the following postscript to the foregoing letter. "I am looking up
Camper's use of the name Mammonteus. At the time, it was believed that there was only one extinct elephant, the
true Mammoth in the Old World and the Mastodon in the New. First allusions to the American Mastodon speak of it
as the Mammoth. It is interesting to note that Kerr gave the name Elephas ainericanus in 1792, several years before
Blumenbach gave the name Elephas primigenius in 1799. I already have the complete bibliography of names applied to
the European Mammoth and shall give them in my Monograph. Perhaps the first properly defined generic name was
Dicyclotherium. Palmer, 'Index Generum Animalium,' 1904, p. 397, gives Mammont as applied to Mastodon ameri-
canus, Mammuthus Burnett. Burnett simply gives the name and the species Mammuthus borealis, a species not listed
in Sherborn's Index nor anywhere defined. I doubt if Mammont, Mammuthus, or Mammut can be adopted on any rules
of nomenclature, since a genus must rest on a type species. At present I prefer the Latin name Mammonteum Camper,
which rests, according to Camper's description, both on the specimens found in Siberia and in North America, but I shall
keep an open mind and settle this question in Volume II."
Professor Osborn's death occurred November 6, 1935.

1937 Conclusions. — A recent note from Dr. Edwin H. Colbert to the editor reads as follows:
1830 Mammuthus Burnett Type designated as Mainmuthus borealis, which is a synonym of Elephas
primigenius Blumenbach. Therefore the designation becomes Mammuthus primigenius (Blumenbach). Hopwood's
arguments (1935) are valid.

1937 Dr. William Berryman Scott in his recent (1937) revision of "A History of Land Mammals in the
Western Hemisphere," adopts Mammuthus Burnett.

Owing to the fact that Mammonteus has appeared so frequently in the literature of the past decade and has
been used throughout the entire text of the present Memoir, it has been decided to retain the name herein, with
the explanatory notes above, despite the evidence in favor of its aboUshment. —Editor.]
LIST OF SUPERFAMILIES
Year Name Author Bibliographic Reference Reference in Present
Memoir
1921 MCERITHERIOIDEA Osborn, Amer. Mus. Novitates, No. 1, p. 2. See Md'RITHERIOIDEA
also Volume I, p. 24, of the present Memoir.
Includes the family Moeritheriidse of Andrews, 1906.

1921 DINOTHERIOIDEA Osborn, loc. cit. DEINOTHERIOIDEA

Includes the family Curtognati [
= Curtognathidae, this
Memoir] of Kaup, 1833.

1921 MASTODONTOIDEA Osborn, loc. cit. MASTODONTOIDEA

Includes the families Mastodontidae of Girard, 1852, and
the Bunomastodontidae of Osborn, 1921.

1921 ELEPHANTOIDEA Osborn, loc. cit. ELEPHANTOIDEA

Includes the family Elephantidae Gray, 1821.

1935 STEGODONTOIDEA Osborn, Proc. Nat. Acad. Sci., XXI, No. 6, p. 408, STEGODONTOIDEA
also 2; Vol. I, pp. 22, 25, and Vol. II, p. 807,
fig.
of the present Memoir.
Includes the family Stegodontidae Young-Hopwood, 1935.
1368 OSBORN: THE PROBOSCIDEA

3. LIST OF FAMILIES
Yeab Name Author Bibliographic Reference Reference in Present
Memoir
1821 ELEPHANTID^ Gray London Medical Repository, XV, No. 88, p. 305. ELEPHANTID.E
1838, Bonaparte, Nuov. Ann. Sci. Nat. Bologna, Anno I, Tom. ELEPHANTID.^5
II, p. 112.

1850, Bonaparte,"Conspectus Systematis Mastozoologiae. Mam- ELEPHANTID^

malia."

1852, Girard, Proc. Amer. Assoc. Adv. Sci., for the year 1851, ELEPHANTID.E
pp. 326, 328.

1891, Zittel, "Handbuch der Palacontologio," p. 458. ELEPHANTIDvE (in part),

also Mastodontidse, Buno-
mastodontidse, Serridenti-
dae, Humbokitidse, Stego-
dontidse

1910, Osborn, "The Age of Mammals," p. 558. ELEPHANTID^ (in part),

al.so Curtognathidae, Masto-
dontidse, Bunomastodonti-
dae, Stegodontidae

1821 MASTODONAD/E (Way London Medical Repository, XV, No. 88, p. 306. MASTODONTID^
1833 CURTOGNATI Kaup Neues Jahrb. Min., p. 516. CURTOGNATHID^
Dinothcriidse Bonaparte, 1850.

1842 ELEPHASIDE^E Lesson "Nouveaii Tableau du Rfegne Animal," p. 156. ELEPHANTID^

Embraces one Mastodon, "Elephas arvernensin" Croizet and
Jobert.

1845 DINOTHERID^ Bonaparte "Catalogo metodico dei Mammiferi Europei," p. 4 {fide Palmer, CURTOGNATHIDiE
1904, p. 738).

1850 DINOTHERIIDyE Bonaparte "Conspectus Systematis Mastozoologia;. Mammalia." CURTOGNATHIDiE

1918, Osborn, Bull. Gcol. Soc. Amer., XXIX, p. 134. CURTOGNATHID.E
In Vol. pp. 26, 82, 83, this Memoir, changed to Cur-
I,

tognathidse, based on family Curtognati Kaup, 1833.

1852 MASTODONTID.E Gir;in Proc. Amer. Assoc. Adv. Sci., for the year 1851, pp. 326, 328. MAST0D0NTID.1':
1918, Osborn, Bull. Geol. Soc. Amer., XXIX, p. 134 (in part). MASTODONTID.E
1906 M(ERITIIEHIID.E Andrews ".\ nes(Ti|)tiv(' ('atalogue of the Tertiary Vertebrata of the MffiRITIIERIID.E
Fayum, Egyiit," p. 99.

1906 PAL.EUMASTUUONTID.E .Vndrews Op. nl., ]). 130. BUNOMASTODON riU.E

Regarded by the present author as invalid, because founded
on the generic characters of Phiomia rather than Pala'omasludon.

1921 BUNUMASTODOXTID.K Osborn Amer. Mus. Novitates, No. 1, i)p. 2, 4. lU"NOM.\ST0DONTII)yE

Replaced the subfamily Bunomastudontinae Osborn, 1918,
J))).
134-136 (see \'o!. I, p. 27, tliis Memoir). Bunomastodonti-
dffi inadmissible, however, under the rules of zoological nomen-

clature (.see Scott, 1937, p. 287). See also Trilophodontida;

Simpson, 1931, p. 1369, and Gomphotheriida; Cabrera, 1929,
below, this list.

1927 AMEBELODONTID.'E Barbour Neb. State Mus., Bull. 13, I, p. 131. bunomastodontiixl:
Subsequently changed by Barboui' (1929.2, p. 139) to (subfam. AmEBELO-
Amebclodontinae. dontin.e)
1929 MAMMUTID.E Cabrera Rev. Mus. La Plata, XXXII, p. 74. MASTODONTID;]']
1929 GOMPHOTHERIID/E Cabrera Ibid., p. 75. BUNOMASTODONTIDiE
(in parf ), also Serridentidse,
Ilumboldtida;
 —

NOMENCLATURE OF THE PROBOSCIDEA 13G9

1931 TRILOPHODONTID^. Simpson Bull. Amcr. Mus. Nat. Hi.st., LIX, Art. V, p. 281. BUNOMASTODONTID^.
Substituted inadmissible name Bunoma.stodontidffi.
for
Adopted by Scott, 1937, pp. 267, 280, 287. [Doctor Simpson now
prefers Gomphotheriidte Cabrera, 1929 (see p. 1525 below).
Editor.]

1935 HUMBOLDTIDiE Osborn Proc. Nat. Acad. Sci., XXI, No. 6, hg. 2 (name only)- HUMBOLDTID.E
1936, Osborn, this Memoir, Vol. I, pp. 575, 722.
Without a type genus. Stegomastodontidae substituted by
Scott, 1937 (see below, this list).

1935 SERRIDENTID.E Osborn Proc. Nat. Acad. Sci., XXI, No. 6, fig. 2 (name only). SERRIDENTID.E
1936, Osborn, this Memoir, Vol. I, p. 729.

1935 STEGODONTID/E Young Pal. Sinica, Ser. C, IX, Fasc. 2, p. 5. STEGODONTID^

1935, Hopwood, Pal. Sinica, Ser. C, IX, Fasc. 3, p. 71.
Included both Stegolophodon and Stegodon. Restricted in
present Memoir to Stegodon.

1935 DIBUNODONTID.^^. Hopwood Pal. Sinica, Ser. C, IX, Fasc. 3, pp. 11, 55. BUNOMASTODONTID.E
See especially Hay, 1925, Journ. Wash. Acad. Sci., XV, p. (in part, i. e., only the
382. Brevirostrinae of Osborn)

1937 STEGOMASTODONTIDiE Scott "A History of Land Mammals in the Western Hemi- HUMBOLDTID^
sphere," pp. 267, 281, 294.
Sub.stituted for Humboldtidae, which is without a type
genus.

LIST OF SUBFAMILIES
Reference in Present
Year Name Author Bibliographic Reference
Memoir
1838 Elephantina Bonaparte Nuov. Ann. Sci. Nat. Bologna, Anno I, Tom. Elephantin.e
II, p. 112; 1850, "Conspectus Systematis
Mastozoologise. Mammalia."

1841 Dinotherina Bonaparte Trans. Linn. Soc. London, XVIII, p. 253. Deinotheriin.e
1850 DiNOTHERiiNA Bonapartc "Conspectus Systematis Mastozoologise. Mam- Deinotheriine
malia."

1869 Mastodontina Brandt Mem. Acad. Imp. Sci. St. Petersb., (VII), Mastodontina
XIV, No. 1, p. 35.

1906 McERiTHERiiNi Winge "Jordfundne og nulevende Hovdyr (Ungulata) Mceritheriina

fra Lagoa Santa, Minas Geraes, Brasilien,"
p. 172.

1906 DiNOTHERiiNi Winge Loc. cit. DEINOTHERIINiE

1906 Elephantini Winge Loc. cit. Elephantine
1910 DiNOTHERiiNiE Osborn "The Age of Mammals," p. 558. Deinotheriin.e
1910 Mastodontina Osborn Loc. cit. Mastodontina
1910 Elephantine Osborn Loc. cit.; 1918, Bull. Geol. Soc. Amer., XXIX, Elephantine
p. 135.

1918 Bunomastodontina Osborn Bull. Geol. Soc. Amer., XXIX, pp. 134-136. Replaced by
BUNOMASTODONTIDE
(in part)

1918 Stegodontine Osborn Ibid., pp. 135, 136. Stegodontina

1918 Loxodontine Osborn Ibid., pp. 135, 136. loxodontina
 .

1370 OSBORN: THE PROBOSCIDEA

Year Name Author Bibliographic Reference Reference in Present

Memoir
1918 Eueleph ANTING Osborn Ibid., p. 136. Invalid
Invalid because the genus Euekphas is

invalid (see Chap. XIX, p. 1177, of present

Memoir)

1918 LoNGiROSTRiNiE Osbom Ibid., p. 136. LongirostrinvE

Without type genu.s. See Trilophodon-
tinse Scott, 1937, p. 1371 below, this list.

1918 Rhynchorostrin^ Osborn Ibid., p. 136. Rhynchorostrin^

See Rhynchotheriinse Cabrera, 1929, on
this page below; Scott, 1937, pp. 267, 280,
292, also adopts the form Rhynchotheriinse.

1918 Brevirostrin^ Osborn Ibid., p. 136. Brevirostrin^

See Pentalophodontinae Scott, 1937, p. 1371
below, this list.

1921 Mammontin^e Osborn Amer. Mus. Novitates, No. 1, pp. 1, 14. Mammontin^
Should Mammonteus prove to be invalid,
thiswould leave the subfamily Mammontinae
without a type genus.

1921 Serridentin^ Osborn Bull. Geol. Soc. Amer., XXXII, p. 330. Serridentin^
1921 NoTOROSTRiNiE Osborn Ibid., p. 330. Notorostrin^
Without type genus. See Cordillerioninae
Scott, 1937, p. 1371 below, this list.

1923 Mceritheriinae Winge-Osborn Amer. Mus. Novitates, No. 99, p. 1. McERITHERIINvE

1923 Zygolophodontin^ Osborn hoc. cit. ZYGOLOPHODONTINyE

1927 Archidiskodontin^ Dietrich Neues Jahrb. Min., I, Abth. B (Referate), [Mammontin^ (in
p. 313. part)]

1927 Pareleph ANTING Dietrich Loc. cit. [Mammontin^ (in

part)]

1928 Platybelodontin^ Borissiak Ann. Soc. Pal6ont. Russie, VII, p. 119. Platybelodontin^e

1929 AiMEBELODONTiNiE Barbour Neb. State Mus., Bull. 16, 1, p. 139. Amebelodontin^
1929 Rhynchotheriin.e Cabrera Rev. Mus. La Plata, XXXII, p. 75. Rhynchorostrin^.

1929 Gomphotheriin^ Cabrera Loc. cit. Longirostrin^ (in

part)
tetralophodontinie
Serridentin.e
1929 CrviERONiiN^ Cabrera Ibid., p. 76. Notorostrin^e (in
part)
Notiomastodontin^
1929 Anancin^e Cabrera Ibid., p. 76. Notorostrinve, Hum-
boldtin^, and
Brevirostrin^;
(in part)
 NOMENCLATURE OF THE PROBOSCIDEA 1371

Year Name Author Bibliographic Reference Reference in Present

Memoir
1930 Mammutin.e Hay "Fossil Vertebrata of North America," II, p. [Mastodontoidea of
623. O.sbornl
Name without definition but indicates the
inclusion of the subfamilies of the Mastodon-
toidea of Osborn.

1932 TetraLOPHODONTIN.^ van der Maarel "Contribution to the Knowledge of the Fossil Tetralophodontin.e
Mammalian Fauna of Java," p. 108.

1934 HuMBOLDTiN^ Osborn Proc. Amer. Phil. Soc, LXXIV, No. 4, p. 277, humboldtin^
fig.3, and p. 283, also Vol. I, p. .57.'i, of
present Memoir.

1935 Gnathalodontin.e Barbour and Neb. State Mus., Hull. 42, I, p. .395. Gnathabelodontin^
Sternberg [emend. 0.sborn Gnatha-
belodontinse]

1936 Pal.*:omastodontin.e Osborn This Memoir, Vol. I, p. 091. Pal^omastodontin^

1936 Stegolophodontin.^ Osborn Op. cil., p. 700. Stegolophodontin^
1936 NOTIOMASTODONTIN^ Osbom Op. cil.. p. 730. Notiomastodontin^
1937 Trilophodontin^ Scott "A History of Land Mammals in the Western Longirostrin.e
Hemisphere," pp. 267, 280.
Substituted for Longirostrinse which is

without a type genus.

1937 CoRDILLERIONINiE Scott Loc. cit. notorostrin^

Substituted for Notorostrinae which is

without a type genus.

1937 Pentalophodontin^ Scott Op. cit., pp. 267, 292. Brevirostrin.e

Founded on Penlalophodon Falconer, 1857,
1865. Compare Anancinae Cabrera, 1929,
above, this list.

1937 Stegomastodontin.e Scott Op. cit., pp. 267, 281. Humboldtin.e

Replaces Humboldtinse which is without
a type genus.

5. GENERA AND SPECIES OF THE PROBOSCIDEA IN THE ORDER NAMED BY THE

AUTHORS IN THE ORIGINAL DESCRIPTIONS^
The following Tables present a revised chronologic list (1735-1939) of generic and specific names as originally
and subsequently spelled by their authors, which rest upon the authority of the authors mentioned, or cited (e.g.,

fide) in cases where the original references were not available to the present author.

LIST OF GENERA
Generic Reference
Tear Genus Author Bibliographic Reference Qenotypic Species in Present Memoir
1735-1758 Elephas Linnaeus, "Systema Naturae," 1st to 9th editions. Elephas indicus Elephas
Linnaeus
[In the lOtli edition of tlie "Systema Naturas" (known as the Decima Reformata),
17.58, p. 33, Linnffus substituted thename Elephas maximus Linn, for tliat of E. iitdicm
Linn., wliieh, with all other names in that edition, was oftioially adopted by the Fifth In-
ternational Congress of Zoology, held in Berlin in 1901. — Editor.]
For first use of Elephas, .see John Ray, 1693, p. 131 [p. 123 of authors).

HThe author of the present refused to recognize privately printed publications (unless on sale, bearing the
Memoir name of the publisher) as a source of

systematic names; consequently such references have been omitted from the present list. Editor.) —
1872 OSBORN: THE PROBOSCIDEA
Generic Reference
Year Genus Author Bibliographic Reference Genotypic Species Memoir
in Present

1788 Mammonleum ('am[)er, Nova Acta Acad. Sci. Imp. Pctropol., II, for [Refers to an animal Mammonleus^
the year 1784, p. 251; Mammonleus Osborii, 1924, from America (cf Hop- .

Amer. Mus. Novitates, No. 152, p. 2. wood, 1935, p. 98).]

Camper's designation of Mammonleum is cited above. Hay (1902, p. 708) and J. A.
Allen (letter, Nov. 23, 1920) do not regard it as valid. Allen writes: "the names of the
fossil mammalshe [Camper] discusses are vernaeular names rendered into Latin. They
liave no nomenolatural significance." Osborn nevertheless adopts Mammonteum Camper,
1788, in preference to LHcyclotiwrium GeofFroy, 1837, as a distinct generic name assigned to
E. primigenius. While Allen in the above letter objects to Mammonteum as a vernacular
name rendered into Latin, in a letter (Nov. 15, 1920) on the subject of barbarous names as
generic names, he nevertheless agrees with Palmer ("Index Generum Mammalium," 1904,
])1). 4.J, 46) and says: "AU modern codes of [or] rules of nomenclature declare that such
names are not to be rejected on ground that they are nonclassical, and that incorrectly
tlu>

formed classical names are not suliject to emendation." See also Allen's ivAv. on the word
"Matiiitiul."

1793 Elephantu.s Ctn ier and (leoffroy. Revue Encyclop^dique (or Mag. En- Genotype not given in Elephas
(ycl()i)Miqiie), II, (6), p. 189. 1795, but in 1801
(Cuvier and Lacepfede)
indicus is mentioned

1797 Ohio Blumenbacli, Abbild. naturhist. Gegens., Heft 2, No. 19, fig. A, Ohio-incog n it u m. Mastodon
and back of title-page. Blumenbach
Not a valid generic description. See introduction to this chapter

1799 Mammut Blumenbach, Handb. d. Natur., 6th edition, p. 698. Mammut ohioticum Mastodon
Blumenbach
A vernacular name preoccupied by Ohio
for E. primigenius. Allen, however, writes
(letter, Nov. "In your letter you ask whether I consider Blumenbach's name
15, 1920):
Mammut entitled to recognition. I certainly do, although with regret that such name was
not [sic] ever proposed as a generic designation. Neither its barbarous form nor the fact of
his misconception that it was the mammoth renders it invalid."
Murray, 1908,VI, Pt. 2, p. 98: "MAMMOTH
Also 8 mamrauth, mamant, maman,
mamont, mammon, mammot (mammoht), 8-9 mammouth. [a. Russian, MaMMOTb, . . .

mammot, whence matnmotovoi host, mammoth's bones (Ludolf, Gram. Russ. 1696, p. 92);
now MaMaHT'b, mamant. Hence also F. mammouth, mamant, mammont. The
. . . . . .

word is of obscure origin; the alleged Tartar word mamn 'earth' (usually cited as the ety-
mon) is not known to exist.]"

1805 [1806] Mastodonte G. (

'uvicr Ann. Mu.s. d'Hist. Nat., VHI, pp. 270, 272, 293. Le Grand Mastodonte Mastodon
Cuvier By courtesy to
Cuvier restricted
Falconer and Cautley, 1846, letterpress, p. 18: "But in his second extended and to M. americanus
elaborate memoir, |)ul)lishcd in 1805 [1806], which formed the groundwork of what he has
written on the subject in the 'Ossemens Fossiles,' Cuvier separated the Elephants with
manimiUated molars from the ordinary forms with lamelliform molars, and united the
former into a genus which he designated Mastodon, taking the North American s|)ecies,
under the name of M. giganteum, as the type (Footnote: 'Annales du Museum d'Histoirc
viii. "Sur le grand Ma.stodonte." ']."
Naturelle, torn.
See also de Blainville (1834-69, p. 245); and Leidy (1869, p. 393): "The earliest
date at which I have been able to find the name of Mastodon systematically expressed,
is in the work here quoted [Cuvier, 1817,
p. 233]. Previously, Cuvier api)ears only to
have u.sed the gallicized term
Mastodonte. Bronn, in the 3d edition of the Lethaea
of
Geognostica, page 820, credits Mastodon to Cuvier as early as 1805, but does not give the
reference." [See Oken, 1916, this chapter, p. 1364 above.)

1808 Harpagmotherium Fischer de Waldheim Prog. d'Invit. Stance, Pub. Hnrpagmolherium raiia- Mastodon
8oc. Imp. Nat. Mokcou, September, pp. 19, 20 {Jide dense Fischer de
Palmer, 1904, p. 311). Waldheim
Comjiarc Sherborn, "Index Animaliimi," who gives on page 1022 (Sect. 2, Pt. 5, 1924)
llarpagonotherium canadense, and on jKige 2915 (Sect. 2, Pt. 12, 1927) Harpagmotherium.

^{Mammuthus Burnett, 1830 (see below, this list) selected by Dr. A. Tindell Hopwood, 1935, p. 27. I'^or complete historical account of the names Mam-
monleus, Mammut, Mastodonte, Mastodon, up to the year 1935, see pp. 1363-1367 above. —Editor.]
 NOMENCLATURE OF THE PROBOSCIDEA 1373

Generic Reference
Year Genus Author Bibliographic Reference Genotypic Species in Present Memoir
1814 Mastotherium Fischer de Waldheim "Zoognosia," III, p. 337. Species cited : M. Mastodon
megalodon (Cuv.) americanus
leptodon (Cuv.) Tiilophodon angusti-
dens
microdon (Cuv.) Turicius tapiroides
hyodon (Cuv.) Cordillerion andium
humboldtii (Cuv.) Cuvieronius liumboldiii

1816 Mastodon Oken Lehrbuch Natur., Theil III, Abth. 2, p. 789. Mastodon Cuv.
First use of the term Mastodon for Cuvier's Maslodonte, citing Cuvier's five classio
species.

1817 Mastodon C. Cuvier "Le Regne Animal," I, pp. 232, 233. Mastodon giganteum Mastodon
Cuvier, Mastodon
angustidens Cuvier
1817 Mastodontum de Blainville Nouv. Diet. Hist. Nat., IX, p. 276. [Mastodon Cuv.]

1825-1827 Loxodonte F. Cuvier "Hist. Nat. Mamm.," Ill, Livr. LI, LII, with Elephas africanus Loxodonta
[=Loxodonta, 2 pp. text, 1827, 1828, Zool. Journ., London, III, p. Bhunenbacli
1827, 1828] 140 (unsigned review).
Geoffroy Saint-Hilaire, Etieiine, et Cuvier, Frederic, 1824-1829 [1825], (Loxodonte),
p. 2: "Je proposerai pour nom generique de cette espece, le mot de Loxodonte, qui peut
rappeler le caractere de ses dents, les losanges qu'on aperQoit sur leur coupe."

A
review (unsigned) of this work appeared in the Zool. Journ., London, 1827, 1828,
III, p. 140, noticing the "dismemberment of the genus Elephas, for the purpose of establish-
ing a new one under the name of Loxodonta. For the Elepliant of Asia he [Cuvier]
. . .

retains the original generic name Elephas. The surfaces of its molar teeth present fascise of
enamel irregularly festooned; while in those of the African Elephant, the type of the new
genus Loxodonta, tlic enamel is disposed in lozenges. In addition to this sti iking distinction
derived from the dentary system, M. F. Cuvier also enumerates the other characters which
have hitherto been regarded as specific. The smaller, more elongated, and less irregular
head of the African animal when compared with the Asiatic; the rounded forehead of the
former, strongly contrasted with the deep depression in the middle of that of the latter; the
ear of the former also twice the extent, while the tail is ordy half the length, &c."

1829 Deinotherium Kaup Isis, [XXII], Heft IV, p. 401, Taf. i. Deinotherium gi- Deinotherium
ganteum Kaup
1830 Mammuthiis Burnett Quart. Journ. Sci., London, July-December, Mammuthus borealis Mammonteus
1829, p. 352. Burnett
[See introduction to the present chapter, pp. 1366 and 1367.]

1830 Tetracaulodon Godman Trans. Amer. Phil. Soc, N. S., Ill, p. 484. Tetracaulodon masto- Mastodon
dontoideum Godman
1837 Gomphotherium Burmeister Handb. d. Natur., p. 795. Not cited. Mastodon
Burmeister, 1837, p. 79.5: "Mastodon. Wie Elephas, aber die Backzahne mit 2 Reihen
—
kegelformiger Hooker. Von mehreren untergegangenen Arten findet man Knochen in
Nordamerika, besonders am Ohio, daher Ohiothier. Stossziihne in beiden Kiefem besass die
gleichfalls untergegangene Gatt. Gomphotherium."
Osborn, 1922: The name Gomphotherium is invalid: (1) Because no genotypic species
except the Ohio mastodon is mentioned; (2) because several different genera of raasto-
donts display the same character, namely, "Stosszahne in beiden Kiefem," e. g., Tetracaulo-
don Godman, a four-tusked true Mastodon americanus. Matthew (unpublished manu-
script of 1918) states: "Apparently the author [Burmeister] had primarily in mind certain
specimens of the American mastodon which retain the lower tusks; but for these the name
Tetracaulodon Godman had been proposed in 1830." (3) Confusion as to mastodons with
four tusks is demonstrated further by the fact that Kaup fir.st used Godman's term in
describing his Eppelsheim species, viz., Tetracaulodon longirostre. (4) It is consequently
clear that we cannot be certain what animal Burmeister had in mind. (See note under
Gamphotherium Gloger below.) Hay adopted the name in 1917 (see Gomphotherium
gratum and G. elegans, p. 1405 below).

1837 Dicyclotherium E. Geoffroy Saint-Hilaire Compt. Rend. Acad. Sci. Elephas primigenius Mammonteus
Paris, IV, No. 4, pp. 119, 120, fig. i. Blumenbach
This name assigned in reference to the fact that the genus was supposed to pass through
two cycles of time. A.ssigned by Geoffroy in an important communication on the influence
of climate on evolution.
 — —

1374 OSBORN: THE PROBOSCIDEA

Generic Reference
rear G«nus Author Bibliographic Reference Genotypic Species in Present Memoir

1840? Missouritiiit Koch "Fossil Remains," pp. 1, 2. M itoiouriuin Kocliii Mdxtoddii

Koch
First described as Koch's Missourian in 1839 (American Journal of Science, XXXV'II,
pp. 191, 192). In 1841 Kocli renamed it the Missouri Leviathan, or Leviathan missourii,
and in 1843 ho described it as the Missourium Theristocaulodon or Leinathan missouriensis.

1841 Cymatotherium Kaiip "Aivtcn der Urwelt," pp. 11-14, Tab. iv. Cymalotherium SiRENIAN?
antiquum Kaup
[Professor Osborn did not regard tliis as a proboscidean molar. It was originally
listed by Kaui) as a Sire.nian, but neither Dr. G. G. Simpson nor Dr. E. H. Colbert considers
it as such; they think it po.ssible that it may be an embryonic tooth of a i)ro))Oscidcan.
Editor.]

1841 Len'athaii Koch 'Description of the Missourium, or Missonii Levia- Levialha n M /.s.s(»//-// Mastodo
than," etc., London, p. 17. ( = Leviathan inissuuri-

ensis = Missourium
theristnrnidndov)

1841 Gampholherium Gloger "Gemeinniitziges Hand- u. Hilfsbuch Natiu'- Mastodon angustidens Trilophodon
gesch.," I, pp. xxxii, 119. See also Oldfield Thomas, Cuvier
1895, Ann. Mag. Nat. Hist., (6), XV, pp. 191, 192.
Gloger, 1841, p. 119: "Letztere theilte mit ihm noch ein anderes, welches man fiiglich
Schnabel-Mammuth nennen kann, (Gamphotherium angustidens,) obwohl bei ihm die
unteren Stosszahne .selbst nicht bloss klein blieben, sondern auch niu' in der Jugend vorhand-
en waren imd dann bald fiir immer ausfielen."
Gatiipholherium is regarded as a misspelling of Gomphotherium Burmeister, 1837, by
most authors (Matthew, Hay, .411en); Oldfield Thomas, liowever, writes (letter, Oct. 20,
1920): "Gamphotherium Gloger has as its genotype, by monotypy, G. angustidens no —
other species being referred to. There ajjpears to be no reason to suppose the word has any
sjiecial connection with Gamphotherium, either as misspelling or correction. it may have
. .

been a misprint for Gomphotherium of Burmeister, but there is no evidence for this, & the a
occurs equally in Gloger's Systematic Inde.\, in the body of the work, and in the alphabetical
index at the end. The genotype of Trilophodon Falc. & Caut., 1846, is fpiite clearly
. .

ohiolicus, no other species being mentioned in the paragrajihs referring to it."

Hopwood, 193.'3, pp. 13, 14: "Burmeister's genus Gomphotherium was originally
diagnosed thus, 'Stosszahne in beiden Kiefern besass die gleichfalls untergegaiigenc Gatt.
Gumpholherium.' (Burmeister, loc. cii.), but \\v mentioned no species as belonging to this
genus. The late Dr. O. P. Hay (Hay, 1923, [1923.2] p. 109), regarded this as coming imdcr
Opinion 46 of the International Commission on Zoological Nomenclature and adopted it as
a valid genus with genolectotype M. angustidens Cuvier (cf. Cope & Matthew, 1915, expl.
to pi. cxx). Since the American Ma.stodon occasionally has mandibular tusks, it is clear
that Biu'meister's diagnosis does not distinguish Gomphotherium from Mastodon vmdcr
wliich he expressly mentions 'Ohiothier', i. P., Mastodon americanus. On this we may
regard Gomphotherium as invalid, because the diagnosis is inadc(|uate. Not only is it in-
valid, but it also antedates Gamphotherium Gloger. Oldfield Thomas (1895, p. 189) regards
this latter as a new generic name, and (op. dt., p. 191) italicises it as a name which is not
a simple synonym of an earlier name. The spelling, however, is quite clearly either an error
of transcription, or a lapsus calami, or even a misprint, and the word should be written
Gomphotherium, thus becoming a synonym of Gamphotherium BurmeisU^r. The next name,
'I'rilophodon, was originally pro|x)sed for the species M. ohiolicus and M. angustidens. Of
these the former is tlu^ genotype of .Mastodon, hence the latter only remains in Trilophodon
and may be regarded as the gen<)tyi>e of that genus. For these reasons Trilophodon is here
adopted as a valid genus to include all the liunolophodont jjroboscideans grouped round
Mastodon anguMidens."

lS4t) Trilophodon (Section name) Falconer and Cautley "Fauna Antiqua Trilophodon
Sivalensis," letterpress, p. 54: "Mastudun. Sect. Collective genus. By
Trilophodon. M. Ohiolicus. The next degree of — courtesy to Falconer
deviation from the ordinary dental rule is presented restricted to M. an-
l^y Mastodon Ohiulicus." In 1847 Falconer and gustidens (see Trilopho-
Cautley (Pis. xlii-xlv) use the same term, e. g., don Falc, 1857).
 NOMENCLATURE OF THE PROBOSCIDEA 1375

Generic Reference
Year Genus Author Bibliographic Reference Genotypic Species in Present Memoir
Mastodon Sect. Trilophodon, to include the species:
III. Mastodon tapiroides, IV. M. ohioticus, V. M.
angustidens, VI. M. andium; the same species are
Usted by Falconer (1867, p. 56) as "I. Trilopho-
dontes," with the addition of Deinotherimn grgan-
teum and D. indicum.
Tliis does not appear as a generic definition; .see also Tetralophodon Falconer and
Cautley below. Compare also Warren (18.52, p. 139) "The Mastodons are separated into
:

two groups; one called Trilophodon, and the other, not particularly named, which might
be called Tetralophodon." Defined by Falconer as a subgenus of Mastodon (Falconer, 18.57,
pp. 313, 316, and Synop. Tab. opp. p. 319 with addition of four species and transfer of
andium to Tetralophodon).

1847 Tetralophodon (Section name) Falconer and Cautley Op. cit., Pis. Tetralophodon
XLii-XLV, (as cUstinguished from Sect. Trilopho- Collective genus. By
don) applied as a section of Mastodon to include the courtesy to Falconer
species: VII. M. perimensis, VIII. M. sivalensis, restricted to M. longi-
IX. arvernensis, X. M. longirostris,
M. M. XL rostris (see Tetralopho-
latidens; these five species in the order named are don Falc, 1857).
again listed by Falconer (1867, p. 56) as: "II.
Tetralophodontes," as distinguished from "I. Trilo-
phodontes" and "III. Stegodontes."
This designation of the Section Tetralophodon is not a generic definition; see also
Trilophodon above. Defined by Falconer as a subgenus of Mastodon (Falconer, 1857, pp.
313, 316, and Synop. Tab. opp. p. 319 with addition of andium). See Vol. I, Chap. IX, p.
348, of the present Memoir.

1847-1857 Slegodon Falconer and Cautley Op. cit., PL xlii, figs, xii-xv. Also Elephas cliftii Stegodon
Falconer, 1857, Quart. Journ. Geol. Soc. London, Falc. &
Caut.,
XIII, pp. 314, 318, and Synop. Tab. opp. p. 319. E. bombifrons
Falc. & Caut.,
Falconer, 1857, p. 314: "To this group we have assigned E. ganesa (?)
the subgeneric name 'From oTtyri
of Slegodon [Footnote: Falc. & Caut.,
ledum, and bhovs dens, having reference to the gable-end E. insignis
form of the section of the ridges.']- It is limited to extinct Falc. & Caut.
forms confined at present to the Indian Tertiaries. The
Stegodons constitute the intermediate group of the Proboseidea from which the other
species diverge through their dental characters, on the one side into the Mastodons, and on
the other into the typical Elephants."
Slegodon is admirably characterized by Falconer by four species belonging in the same
phylum. It preoccupies Emmenodon Cope, 1889.

1847-1857 Loxodon Falconer and Cautley Op. cit., [1847, Pis. xlii, xliv] (name). Elephas planifrons Loxodonta
Falconer, 1857, Quart. Journ. Geol. Soc. London, Falconer & Cautley,
XIII, pp. 315, 318, Synop. Tab. opp. p. 319; 1865, E. africanus Blimien-
ibid., XXI, p. 263. bach, E. priscus{?)
Goldfuss
Loxodon Falconer, 1847, 1857, preoccupied by Loxodon Miiller and Henle, 1841, for
a genus of sharks. Compare Loxodonte F. Cuvier, 1825, and Loxodonta (unsigned review of
F. Cuvier, 1827, 1828—see above, this list).

1847 Elasmodon Falconer and Cautley Op. cit., PI. xlii. Invalid, preoccupied
A subgeneric name based upon species now known to belong to five different genera
(see p. 1177 of the present Memoir), namely, Elephas [Hesperoloxodon] antiquus, E. [Hypsele-
phas] hysudricus, E. [Archidiskodon] meridionalis, E. [Palseoloxodon] rmmadicus, E. indicus,
and E. [MammonUus] primigenius. Name, moreover, preoccupied by Elasmodus, replaced
by Euelephas Falconer, 1857, p. 315 (see Euelephas, p. 1376, below).

1848 Cheirolites von Meyer (In Bronn's Handb. einer Gesch. d. Natur, III, Elephas primigenius Mammonteus
Index Pal., p. 286). Blumenbach
Compare Owen, 1846, p. 228: "A .separate plate [of molar tooth], with its digital

processes, offers a rude resemblance to a hand, and such specimens have been figured by the
older collectors of petrifactions, under the name of 'Cheirolites,' as the fossilized hand of
a monkey or a child."
1376 OSBORN: THE PROBOSCIDEA

Generic Reference
Year Genus Author Bibliographic Reference Genotjrpic Species in Present Memoir
1850 Syiiofinnthen'Kni Costa Atti Accad. Pontaniaiui, \', Pt. 1, pp. 271-275, Tootli helonguig to Mniiniioiiieiis
Tav. Ill, figs. 1-4. Elephax prim igeii /(<.'•

(fide Leidy,
Marschall).

1852 Tetralophodon Warren "The Mastodon Ciiganteus of North America," Mastodon laiideiis Invaliu, preoccupied
p. 139. Clift, M. ai~ver-
nensis Croizet «&
Collective genus invalid because based upon specimens Jobert, M. siva-
now known to belong to three different genera. Moreover, lensifi Cautley
preocrapi(>d by Tftraliiplindon Falconer and Cautley, 1840
[18471.

1855 Anannis Aymard (In Dorlhac Ann. Soe. Agric. Puy, XIX, for 1854, A na nc. us macropl lis Anancus
p. 507). Aymard { = M.
arvernensis)
Aitancus is the first valid generic name applied to a member of this ])hylum. A preced-
ing invalid collective name is Teiralopliodon Warren. Succeeding synonymous terms are
Tetralophodon Falconer and Cautley (in ])art), Pentalophodon Falconer, Bunolnphndon
V'acek (in part), l^itegomaalodon Pohlig, Rhabdobunus Hay, Dibunodon Schlesinger.
The name Anancus first appearedDorlhac (1855, p. 507) which he says was
in a list in

borrowed from a verbal communication to the above Society by Aymard in 1855; later in
1859, p. 493, Lartet established its validity by making Mastodon arverneTisis the type.

1857 Trilophodon Falconer Quart. Journ. Geol. Soe. London, XIII, p. 313; Mastodon angustidens Trilophodon
defined as a subgenus, p. 316 and Synop. Tab. opp. and other trilopho- By courtesy to Fal-
p. 319. donts, as listed abo\'e coner restricted to M.
under Trilophodon, angustidens
[See above, this list, under Gainphotherium, 1841, for 1846 (Section mame)
conclusions of Dr. Hopwood regarding validity of Trilopho-
don.]

1857 Euelephas Falconer Quart. Journ. Ocol. Soe. London, XIII, pp. 315- Elephas Hysudricus Invalid subgeneric
318, Synop. Tab. opp. p. 319, subgenus of Elephas. Falconer & Cautley, NAME
antiquus Falconer &
Falconer, 1857, p. 315, footnote: Sub.stitution of Euele- Cautley, meridionalis
phas for Elasmodo7i. Sdater in 1900, p. 317, erroneously Nesti, Namadicus
s[)ecified Elephas planifrons as the genotype of Euelephas, Falconer & Cautley,
because this species was not included in the original defi- indicus Linneeus,
nition of Elasmodon (.see Chap. XIX, p. 1177 above). primigenius Blumen-
bach

1857 Tetrahphodnn Falconer Quart. Journ. Cieol. Soe. London, XIII, pp. Mastodon longirostris Tetralophodon
313, 317, and Synop. Tab. opp. p. 319, defined as and other species, as By courte.sy to Fal-
a subgeiuis of Mastodon, with the doulitful inclusion Hsted above under coner restricted to
of Audi am at this time. Tetralophodon, 1846 .1/. longirostris
[1847] (Section name).

1857-1865 Pentalophodon Falconer Quart. Journ. (Jeol. Soe. London, XIII, p. 314; No species cited in Pentalophodon
1865, ibid., XXI, p. 262. 1857 ; Mastodon Collective genus. By
sivalensis referred to courtesy to Falconer
Tetralophodon, 1857; and Cautley restrict-
As a subgenus of Maslodon, name without genotyi)(.' in made the genotype, ed to M. sivalensis
1857, name with genotype (A/, sivalensis) in 1865. Falconer, 1865.
18.57, p. 314: "For reasons which will be explained in the sequel, it would seem tliat there
has existed in nature anotlier subgeneric grou]) of Mastodon, of which only a single form is
at pre.sent known, in which the crowns of the 'intermediate molars' are divided upon
a ([uinary ridge-formula. This grou]) in our arrangement would l)e characterized, in
harmony with the others, as PerUalophodon."
In i'"alconer's paper of 1857, .1/. sivalensis is nipcatedly (ilaced in the subgenus Tetralo-
phodon; I'alconer states on pag(! 317 that: ".Mastodon Sivaletms, although with five-
ridged 'intermediate molars,' is provisionally included under Telrnhiphodon." (See also
Synop. Tab. opp. p. 319.) Finally, in 1805, p. 262, Falconer a<lopts the name I'enlnlopho-
don with the genotypic species M. (Tetralophodon) sivalensis.
 NOMENCLATURE OF THE PROBOSCIDEA 1377

Generic Reference
Year Genus Author Bibliographic Reference Genotypic Species in Present Memoir
1867-1883 Leptodon Guiin (!col.Mag., IV, p. 422 (name only) ; 1883. Gcol. :\Iag., Leptodon minor, L. Hesperoloxodon
Dec. II, N. S., X, p. 458. giganteus Gunn
Gunn, 1867, p. 422: new species, called by Mr.
"(6) the necessity for establi.shing a
Gunn Leptodon, from the fineness of the enamel."Gunn, 1883, p. 458: "submitted to
a meeting of the Geological Society at Somerset Hou.se in 1867, under the names of Lepto-
don minor and Leptodon giganteus." Type: E. (Leptodon) aniiquus; Forest bed stage,
England. Generic name Leptodon preoccupied by Gaudry, 1860, for a genus of chalico-
theres; by Sundevall, 1835, for a genus of birds.

1868 AnloleiheriumYaXQowcv "Palseont. Mem." [editor: Murchison], I, p. Deinotheriiun

416, PI. XXXIV, figs. 1, 2.

Footnote on p. 416: "It docs not appear from Dr. Falconer's notes that he had seen
the specimen, which is here described from a drawing made by Col. Baker. — [Ed.]"
1868 Rhy nchotheri inn Vahoncv (MS. 1856); "Palaeont. Mem.," II, pp. 74, Cast of lower jaw from Rhynchollierium
75. See also Falconer, 1863, pp. 44, 56, 60, and Mexico, in Geneva
Osborn, 1918.468, p. 136. Museum
Falconer, 1868, pp. 74, 75: "Extract of Letter from Dr. Falconer to M. Lartet,
II,

September 12, 1856. 'At Genoa [Geneva] I saw a cast of a large lower jaw of a Mastodon
from Mexico, with an enormous bee abruptly deflected downwards and containing one very
large lower incisor. The beak is much thicker than in M. (Trilophodon) angustidens and
larger than in M. (Tetralophodon) longiroslris. You know that every one (Laurillard,
Gervais, &c.) has insisted on the absence of the lower incisors from both of the South
American species. The outline of the jaw resembles very much the figure in Alcide
D'Orbigny's Voyage, described by Laurillard as .1/. Andium. The specimen is unpublished
material, and I was therefore only allowed to examine it very cursorily. The Genoese
Ijalseontologists had provisionally named it Rhyncholheritun, from the enormous develop-
ment of the beak, approaching Dinotherium.'"
Search for this cast was unavailing for many years, but finally (1931) it was located at
the Natural History Museum in Geneva, Switzerland (see Vol. I, p. 477). In the meantime
Osborn (1921.515, p. 5) validated this genus: "The original genotype may be termed
Rhynchotherium tlascalse, new species, from the locality Tlascala." Compare figure 448.

1877 Cxnobasileus Cope Proc. Amer. Phil. Soc, XVI, pp. 584, 585 (Pal. C. treynontigerus Cope Not a proboscidean
Bull. 24). { = M. americanus,
fide Hay, 1902, p.
Based on an artifact; name withdrawn by Cope in 1889 708).
(Amer. Nat., XXIII, p. 207).

1877 Zygolophodon Vacek Abhand. geol. Reichsanst., VII, Heft 1\, p. 45. Mastodon borsoni Hays, Zygolophodon
Wien. M. turicensis By courtesy to
Schinz, M. tapi- Vacek restricted
A dear and admirable designation of the four species roides Cuvier, M. to M. borsoni
whicli together appeared to constitute a single distinct genus. pyrenaicus Lartet
Osborn, however, in 1920 (1926.706) removed the .species M.
luricerms, making it the type of a new genus Turicim; in tlic i)resi'nt Memoir he aissigiis
.1/. tapiroides Cuvier to the genus Turicius (see Vol. I, pp. 203, 217) leaving ^f. borsoni and

M. pyrenaicus as typical of Zygolophodon (see Vol. I, Cliap. VII).

Hopwood, 1935, p. 42: "Vacek (1877) used the genus A/asto(fo(t strictly in the Cuvieri-
an sense, and sub-divided it into two groups, Zygolophodon and Bunolophodon, including
in the former, M. tapiroides Cuv., M. pyrenaicus Lartet, 'Form von Baltaviir', 'M. Turicen-
sis von Pikerrai', M. Borsoni Hays, and M. ohioticus. Of these species Matthew (1918, p.
200) selected XL tapiroides as the genotype. Osborn (1926 [1926.706]) has taken M. borsoni
as genotyi^e, but luider the rules of priority this selection is inadmissible. . . . The first

mention of M. tapiroides is in Desmarest (1822, p. 386) who latinised Cuvicr's vernacular,

and took as his type the specimen figured by Cuvier. Hence, the genotype of Zygolophodon
is M. tapiroides Desmarest, spec, indet., and thus Zygolophodon remains an indeterminate

genus." See also this Memoir, Vol. I, p. 203, footnote.

1877 Bunolophodon Vacek Loc. cit. Mastodon atticus Collective genus.

Wagner, M. longi- Invalid = A nancus,
roatris fKaup, M. = Trilophodon, etc.
angustidens Cuvier,
Af arvernensis
.

Croizet and Jobert,

M. pentelici Gaudry
1378 OSBORN: THE PROBOSCIDEA
Generic Reference
Year Genus Author Bibliographic Reference Genotypic Species in Present Memoir
1882 .\ olelephas ()v.cn Proc. Roy. Soc. London, XXXIII, p. 448; also Phil. Xotelephas aiintraliti DiphotodoiN, a
Trans. Roy. Soc. London, 1883, CLXXIII, Pt. Ill, Owen marsupial
p. 777.
See note under Noteleplias australis, 1882 (p. 1397 below), from Jack and Etheridge,
1892, p. 683.

1884 Dibelodon Cope Proc. Amer. Phil. Soc, XXII, pp. 2-8. Mastodon shepardi Invalid
Leidy
Invalid because preoccupied by Khynchothcrium (see tiii.s Memoir, I, ]). 52.')).

1884 Telrabelodon Vo\)c /6;V/., pp. 4-5. Mastodon augustidem Invalid

Cuvier
Invalid because preoccupied by Trihphodon (see this Memoir, I, pp. 249, 525).

1885 Archidiskodonten Pohlig Zeitschr. deutseh. geol. CJes., XXXVII, p. Elephas planifrons Archidiskodon
1027. Falconer and Cautley

1888 Archidiskodon Pohlig Nova Acta Leop. Carol., LIII, pp. 138, 252. Elephas meridionalis Archidiskodon
Nesti, E. planifrons
Falconer and Cautley
Pohlig, 1885, p. 1027: "8. Ich theile die Elephanten nach Kronenformen und Lamel-
lenzahlen der Molaren ein in .Archidiskodonten {E. planifrons, E. meridionalis), Loxodonten
(E. africanus, IE. anliquus) und Polydiskodonten [E. primigcnius, E. indicus, etc.), die
Stegodonten mit Clift wieder zu Mastodon ziihlend."
An excellent generic distinction of two very closely related species (E. planifrons, E.
mcridionalvi) to which may now be added Elephas imperalor.
,

1885 Loj-o(/o;(<e« Pohlig Zeitschr. deutseh. geol. Ges., XXXVII, p. 1027. Elephas africanus Loxodonta
Blumenbach, ?£.
antiquus Falc. & Caut.

1888 Loxo-{disko)-donlen Pohlig Nova Acta Leop. Clarol., LIII, pp. 138, 252. Elephas africanus
Loxo{-disko-)don Blumenbach, E.
priscus Goldfuss, E.
antiquus Falc. &
Caut.
Proposed as a group or sectional name, equivalent to the Loxodontinie of Osborn.
Sec note under Archidiskodon, 1885-1888, above, first paragraph.

1885 l'olijdisko(lo)itrn Pohlig Zeitschr. deutseh. geol. (Jes., XXXVII, i).

Elephas primigcnius Invalid
1027. Blumenbach, E.
indicus Linnaeus

1888 Polydiskodon Pohlig Nova Acta Leop. Carol., LIII, pp. 138, 2.52. Elephas primigenius
Blimienbach, E.
indicus Linnaeus, E.
(?) namadicus Falc.
& Caut.
Proposed in 1885 as a group or sectional name; in 1888 as a genus. See note under
Archidiskodon, 1885-1888, above, first ijaragraph.
Collective genus invalid, because based on species now known to belong to three dif-
ferent genera, Mamnwnleus, Elephas, and Palseoloxodon.

1885 Stegodonten Pohlig Zeitschr. deutseh. geol. Ges., XXXVII, p. 1027. No .species cited

1888 Stego{-lopho-)don Pohlig Nova Acta Leop. Carol., LIII, p. 252.

Proposed in 1885 as a group or sectional name. See note uudc'r Archidiskodon, 1885-
1888, above, first paragraph.

1889 Emmenodon Cope Amer. Nat., XXIII, p. 194. Elephas cliftii V-Av. & Invalid
Caut., Mastodon cle-
Invalid because based upon two s|x;cies « liich are phantoides Clift
typical members of the genus Siegodon.
 .

NOMENCLATURE OF THE PROBOSCIDEA 1379

Generic Reference
Year Genus Author Bibliogrraphic Reference Genotypic Species in Present Memoir
1901 Palxomastodon Andrews Zoologist (4), V, Aug. 15, pp. 318, 319 (name Falieomadodon beadneUi Palseomastodon
only); Tageblatt V Internat. Zoologen-Cong., Ber- Andrews
lin, No. 6, Aug. 16, p. 4 (published vol. 1902, p.

528) Geol. Mag., Dec. IV, N. S., VIII, pp. 400-409.

;

1901 Moeritheritan Andrews Tageblatt V Internat. Zoologen-Cong., Berlin, Mceritherium lyonsi Mceritherium
No. 6, Aug. 16, p. 4 (name only) Geol. Mag., Dec. ; Andrews
IV, N. S., VIII, pp. 400-409.

1902 Phiomia Andrew.s and Beadnell "A Preliminary Note on Some New Phiomia serridens Phiomia
Mammals from Upper Eocene of Egypt," Surv. Andrews and Bead-
Dept., Pub. Works Ministry, pp. 1-9. nell

1912 Promaslodon Pohlig Bull. Boc. beige G6ol., XXVI, Proces-verbaux, Puerco mammal Invalid
p. 192.
Pohlig, 1912, p. 192 :
". . .illustree aussi i)ar oolui de Puprco (prototype de Promaslodon
Pohlig)." Founded on a supposed missing link in the American Paleocene Puerco; possi-
bly Cope's Polymastodon, a niultituberculate. This invalid description of Promaslodon
precedes (p. 192) the valid generic definition of Stegomaslodon <•{. mirtficmn, page 193, as
cited below.

1912 Stegomastodon Pohlig Ibid., p. 193. Mastodon mirificus Stegomaslodon

Leidy
Pohlig, 1912, p. 193: .c'est prouvo, notamment, par une supcrbc dent de Slego-
mastodon of. mirificum Leidy (prototype de Slegomaslodon Pohlig)."

1912 Hemimastodon Pilgrim Mem. (Jeol. Surv. India (Pal. Indica), N. S., Tdmbehdon crepufictdi SUINA
IV, Mem. 2, pp. 17-21. Pilgrim
Type Tctrabehdon crepuscidi Pilgrim, 1908 (p. l.'J7) subsequently named Hemimasto-
don, 1912.

1914 Eubelodon Barbour Univ. Neb. Studies, XIV, p. 194. Eubelodon morriUi Eubelodon
Barbour
1914-1917 Megabelodon Barbour Neb. Geol. Surv., IV, Pt. 14, p. 217 (as a sub- Telrabelodon lulli Megabelodon
genus); 1917, ibid., IV, Pt. 30, p. 512 (raised to Barbour
higher rank).
Barbour, 1914.2, ]). 217: "For this new species, wc wish to propose the name Tetra-
belodon lulli, for Professor Lull. It is not unlikely that it is entitled to rank as a new
subgenus at least, and it might not be amiss to propose the name Megabelodon."
Barbour, 1917, p. 512: "In the case of Tetrabelodon lulli, the sub-generic title
Megabelodon was proposed, but it seems entitled to higher rank [i. e., genus)."
. . .

1914 Uhabdobunus Hay Iowa Geol. Surv. Ann. Rept. for 1912, XXIII, pp. Madodon mirificus Slegomastodon
59, 373. Leidy
Preoccupied by Stegomaslodon Pohlig, 1912, above (ef. also Ha\-, 1930, Publ. Carnegie
Inst. Wash., No. 390, p. 633).

1917 Stcgolophodon Sclile.singer Denk. Naturhist. Hofmus., I, p. 115 (as a Mastodon latidens Slegolophodon
subgenus) ( 'lif t

Schlesinger, 1917, ]>. 115, footnote: "Ich schlage ftir M. latidens, das sich durch seine
kurzc Symphyse von dem Subgenus Bunolophodon, durch seinen Molarenbau von Dibuno-
don entfernt, den Untergattimgsnamen Stegolophodon vor. Der Name bringt einerseits die
nahen Beziehungen zum Genus Stegodon, anderseits die Loslosung der Untergattung von
Bunolophodon und ihre Sonderstellung gegenilber Dibunodon zum Au.sdruck." See
Stego(-lopho-)don Pohlig, 1888, above, this list.

1917 Dibunodon Schlesinger Ibid., p. 124 (as a subgenus, with definition). Mastodon. {Dibunodon) Anancus
arvernense Croiz. &
Jobert
1917 Choerolophodon Schlesinger Ibid., p. 181 (as a subgenu.s). Mastodon pentelicus Trilophodon
Gaudry and Lartet {Choerolophodon)
Type: .Mastodon (Choerolophodon) pentelici Gaudry and Lartet. Schlesinger, 1917,
p. "Schadel niedrig, langgestreckt mit zwei aufwiirts und auswiirts geschwungenen
181:
schmelzbandlosen Stossziihnen. Unterkiefer mit miissig langer Symphyse ohne untere
Inzisoren. Molaren chocrodont (hochgradig suid)." See text and figures. Vol. I, Chap.
VIII, this Memoir.
1380 OSBORN: THE PROBOSCIDEA

Generic Reference
Year Genus Author Bibliographic Reference Genot3T)ic Species in Present Memoir
1917 (jcnomaslodon Barbour Nob. Geol. Surv., l\, Pt. 30, p. 512. Tetrabelorlou irillisloni Trilophodoii
Barbour, T. liiUi {Genomastodon)
Barbour, 1917, p. 512: "Longirostral mastodons seem Barbour, T. otsborni
to have reached their maximum in sueh forms as Tetrabelo- Barbour
don willistoni, lulli, osbomi, and the like. For the present
at least, the may
be groui)ed under a new and distinrt genus, Genomastodon.
above .-In . .

the case of Tetrabelodon lulli, the sub-generic title Megabelodon was proposed, but it. . .

seems entitled to higher rank [i. e., genus]." Tetrabelodon lulli, therefore, is preoccupied by
Megabelodon lulli; by the process of elimination, Genomastodon applies to T. mllistoni and
T. osbomi onlv.

1917 Mastelephas Barbour Loc. cit. No definition and no NOMEN NUDUM

genotypic species
given.

1922 Miomastodoii Osborn Amcr. Mu.'^. Xovitates, No. 49, p. 4. Mastodon merriami Miomastodon
Osborn

1923 Cuvieronius Osborn .\mcr. Mus. Novitates, No. 99, p. 1. Mastodon hinnboldtii Cutneronins
Cuvier (in Desmar-
Cabrera (1929) erroneously refers Mastodon humboldtii est)
to thegenus Stegomastodon Pohlig, which, as shown above, is
based on Leidy's 'Mastodon mirificus,' wholly distinct from
M. humboldtii.

1923 Serridenlinus Osborn Ibid., p. 2. Mastodon product us Serridentinus

• Cope, M. serridens
Osborn, this Memoir, I, |)|). 28.5, 286: "(December, Cope, M. floridanus
1932) On verj' clo.se reexamination of tlie cast (Amer. Mus. Leidy, M. obscurus
1909) of Leidy's classic type (Figs. 232, 233) of 'Mastodon Leidy, Serridentinus
obscurus, also of the original type molar fragments of 'Ser- simplicidens Osborn
ridentinus' simplicidens Osborn, these two species prove to
belong in the genus Trilophodon because of the presence of 'central conules' uniting as
central trefoils with the median conules. Osborn, 1933: Final study of the veritable
. . .

type of Mastodon obscurus Leidy proves that it does not belong in the genus Serridentinus
Osborn, but in the related though distinct genus Trilophodon Falconer. The phylogenetic
position of T. obscurus appears to be distinct from the species Mastodon (= Serridentinus)
productus of Cope and .1/. [ =Ocalienlimis (Ser.)] floridanus of Leidy."

1923-1926 Prostegodon Matsumoto (In Osborn, Amer. Mus. Novitates, No. 99, p. Mastodon latideus Stegolophodon
2). Clift
Subsequently proposed by Matsumoto (1924.3, pp. 324-327) as a subgenus of Stegodon;
jireoccujiied by Stegolophodon Schlesinger, 1917, a genus also based upon the yfaslodon
Intidens of Clift. Tliis description appeared in the .Japanese language. In 1926.1, ]>. 9,
Matsumoto published his English text on this genus.

1924 Ilarpagonolherium Fischer do VValdheim, 1808 {Fide Shorborii. 1924,

Pt. V, p. 1022).

1924 Parastegodon Matsumoto .Jotirn. Geol. Soc. Tokyo, XXXI, pp. 256, Elrphns aurorx Archidiskodon?
257, 262 (in Japanese); 1929, Sci. Kept. Tohoku .Mats., 1915, or a progres-
Imp. Univ., (2), Geology, XIII, No. 1, pp. L3-15 1918 sive Stegodon
(ill Engli.sh).
Elephas (Parastegodon) axirorse, well established on an excellent type, is, like Stegodon
mindanensis, either a highly progressive member of the Stegodon i>hylum or a primitive
member of the Archidiskoilon pliylum, a point whicii can only be determined positively by
the discovery of the cranium of Parastegodon aurorx.

1924 Pabeoloxodon Matsumoto Journ. Geol. Soc. Tokyo, XXXI, \>\>. 255- Elephas naniadicus Palxoloxodon
272 (in Japanese) 1929, Sci. Kept. Tohoku Imp.
; nau ma n n i .M akiyama
Univ., (2), Geology, XIII, No. 1, pp. 7, 10 (in
English).

1924 Sivalikia Osborn .\nie.r. Mus. Novitates, No. 152, p. 2. Elephas namadicus Palxoloxodon
I'alconer and Cautley
 NOMENCLATURE OF THE PROBOSCIDEA •
1381

Generic Reference
Year Genus Author Bibliogrraphic Reference Genotypic Species in Present Memoir
1924 P ilgrimia Otihorn hoc. cit. Elephasfalconeri Busk; Palaolo.rodon
other species cited,
E. melitensis Fal-
coner, E. mnaidrx
Adams, E. antiquus
Recki Dietrich

1924 Mammonteus Osborn hoc. cit. Mammonleus

Compare Mamnumteum Camper, 1788, above, this list.

1924 Parelephas OshoTH Ibid., p. A. Elephas jeffersonii Parelephas

Osborn

1924 Morrillia Osborn Ihid., No. 154, p. 1. Tetralophodon barbouri Morrillia

Osborn
Raised to rank of genus, this Memoir, I, pp. 690, 739.

1924 Lydekkeria Osborn Ibid., p. 2. .Mastodon (Trilopho- Tetralophodon

See Volume I, p. 353, where it is stated that possibly don) falconeri {Lydekkeria)
Lydekkeria may become a synonym of Tetralophodon. Lydekker

1926 Pliomastodon Osborn Ibid., No. 238, p. 1. Mastodon (Miotnasto- PUomastodon

don) matthewi Osborn

1926 Turicius Osborn Ibid., p. 3. Mastodon turicense Turicius

Schinz
Separated from Zygolophodon by Osborn.

1926 Cordillerion Osborn Ibid., p. 15. Mastodonte des Cor- Cordillerion

dilieres Cuvier =
The generic name Cordillerion replaces the preoccupied Mastodon cordil-
name Dibelodon, as used by Cope and Lull, and Mastodon lerarum Desmarest
hiimboldtii Cuv. In 1923 Osborn proposed Cuvieronius to = Mastodon atldium
embrace M. humholdtii. Cuvier
Cabrera (1929, p. 90) erroneously holds that the geno-
type of Cordillerion {Mastodon andium Cuv.) belongs to the same genus as the genotype
of Cuvieronius (.\f. humboUtii Cuv.), and hence are alike referable to Stegoinaslodon Poh-
lig based on M. mirificus Leidy. He accordingly substitutes Notiomastodon gen. nov.,
1929, p. 90, for Cordillerion 0.sborn.

1927 Leiih-Adamsia Matsumoto Japanese Journ. Geol. and Geog., V, No. 4, Leith-Adamsia siwalik- Archidiskodon
Art. XII. iensis Matsumoto

1927 Amebelodon Barbour Neb. State ]Mus., Bull. 13, I, pp. 131-134. Amebelodon Jricki Amebelodon
Barbour

1928 Platybelodon Borissiak Ann. See. Paleont., Russie, VII, for the year Platybelodon Danovi Plalybelodon
1927, pp. 105-120. Borissiak

1929 Synconolophus Osborn Anicr. Mus. Novitates, No. 393, pp. 9, 10. Synconolophus dhok- Sytwonolophus
pathanensis Osborn

1929 Notiomastodon Cabrera Rev. Mus. La Plata, XXXII, pp. 90, 91. Notiomastodon ornatus Notiomastodon
Cabrera

1929 Torynobelodon Barbour Neb. State Mus., Bull. 16, I, pp. 147-153. Torynobelodon loomisi Torynobelodon
Barbour

1930 Prodinotherium fihik Geol. Hungarica (Palaeont. Ser.), Fasc. 6, pp. Prodinotherium Deinotherium
3-21. hungaricum fihik

1931 Teleobunomasiodon Revilliod Mem. Soc. Paleont. Suisse, LI, pp. 20, Mastodon bolivianus Cordilkrion{'!)
21. Philippi

1931 Hesperoloxodon Osborn Amer. Mus. Novitates, No. 460, p. 21. Palxoloxodon antiquus Ilesperoloxodon
italicus Osborn
1382 OSBORN: THE PROBOSCIDEA

Generic Reference
Year Genus Author Bibliographic Reference Genotypic Species in Present Memoir
19.33 Hlkkotherium Frick Hull. Amer. .Mu8. Nat. Hist., LIX, Art. IX, pp. lilickolfuiium blirki lilickotheriiiiii
505, 509, 515, 527-531. Frick

1933 Ayhelodon Frick Thid., pp. 505, 527, 532. Aybelodon hondurensis Aybelodon
Frick

1933 Ocalientinus Frick Ibid., pi). 505, 576, 579. Ocalientinus ojo- Ocalieniinus
caliensis Frick

1933 Trobelodoii Frick Ibid., pp. 505, 576, 580. Trobelodoii taoensis Trubclodon
Frick

1933 Talabelodon Frick /6k/., pp. 505, 576, 581. Talabelodon rio- Trilophodon
graiidciisis Frick (Talabelodon)

1933 I'araplalybelodon Frick Ibid., p. 592. NOMEN NUDU.M

1933 Serbelodon Frick Ibid., pp. 506, 592, 594. Serbelodon barbour- Serbelodon
ensis Frick

1933 Crypiomastodon von Koeiiigswald U'etcn.schappclijko Mcdcdeclingcn, Cryptomadodon mar- ?SlI{ENIAX

Diciist Mijnbouw Nederl. -Indie, I Teil, No. 23, pp. lini \o\\ Koenigswald
Ill, 112.

1934 Metarchidiskodon Osborn Amer. Miis. Novitates, No. 741, pp. 2, 12. Loxodonta griqua Metarchidiskodon
Haughton
1934-1941 Hypselephas Osborn Proc. Amer. Phil. Soc, LXXH', p. 285 (name Elephas hysndricus Hypselephas
only); 1936, this Memoir, I, p. 12; 1941, op. Falconer and
ciL, II, p. 1340. Cautley and Elephas
platycephalus an-
gustidens Osborn

1934-1941 I'latelepha.'i 0.sborn Proc. Amer. Phil. Soc.. LXXH', p. 285 (name Elephas plalycephalus Plalelephas
only); 1936, this Memoir, I, p. 12, PI. xi; 1941, Osborn
op. cit., II, p. 1358.

1935 (inathabelodon Barbour and Sternberg Neb. State Mus., Bull. 42, 1, pp. (inalhabelodon thorpei Gnathabelodon
395, 396. Barbour and Stern-
berg

LIST OF SPECIES, SUBSPECIES, AND VARIETIES

Reference in
Year Name Author Present Memoir
1754 Elcphati indirii.s Ijinnasufi "Systcma Naturtje," ]). 11. llai).: India, Ceylon. Klephih'i indini.t
(jire-Linnaean)
Tlii.s name was used by I.in?ia>us in \7'A, but in the 1758 edition
first of the "Syst<-ma
Natunr," p. 33, he used Elephas maximus for the Ceylon elephant.

17.58 Elephas maximus Linnaeus "Systema Naturae," edition 10, p. .33. Hab.: India, Ceylon. Elephas indicus
Sherborn (letter, .luly 14, 1929) is of tlie opinion tliat inaximwi should be "swept aside."

1775(?) Elefantc Indiana (Jiovanni Targioni Tozzetti Elephas \A rrhidisko-

don\ meridionalis
Nesti, 1825 {fide
Weithofer, 1890,
p. 133)

1792 Elephas americanus Kerr "Animal Kingdom of Linnaeus," 1, p. 116. Type loc: Big Bone Mastodon ameriranus
Lick(?), Boone County, Kentucky, near Ohio River.
Leidy, 1869, p. .392, footnote: "Cuvier, in the works quoted (Tabl. Klem. Hist. Nat.
(an. 6) 1798, 149; Mem. Inst. Nat. Sei. {.\n VII 11799], 19, 21.], and De Blainville, in his
OstcoK- tion. Elepliants, 327, 24."), attribute this name to Pennant. Faleoner and Cautley,
in the Kauna Anticj. .Sival., 17, also observe, 'that Pennant first ventured in 1793 to desig-
nate the .\mcriean fossil animal, in a sy.stematie work, as a speeies of Elephant by the name
of A', americanus.' I have been unable to find the name thus expressed in any of the works
of Pennant, nearer than the words '.\meriean Elephant,' wliich oeeur in the .Synopsis of
Quadrupeds of 1771 and in both editions of the Hislon,- of Quadrupeds, that of 1781 and
1793."
 NOMENCLATURE OF THE PROBOSCIDEA 1383

Reference in
Tear Name Author Present Memoir
1796 Elephas Mammoth Ciivier [and Geoffroy] Mammont'eus primi-
geniiis
Falconer, 1868, II, p. 158: 'Cuvier was undoubtedly the first to characterize the ex-
tinct species with exactness, in his joint memoir with Geoffroy, under the name of Elephas
Mammoth, in the year 1796 [Footnote: 'Mem. de I'lnstitut, 1" Classe, torn, ii.']."

1797 Ohio-Incognitum B\umenhBch Abbild. naturhist. Gengens., No. 19. Hab. : Ohio River, North Mastodon americanus
America.

1797 Elephas asiaticus BlumenhsLch "Handb. d. Natur.," 5th edition, p. 124. Hab.: India. Type fig. Elephas indicus
Blumenbach, Abbild. naturhist. Gegens., No. 19, fig. B.

1797 Elephas africanus Blumenbach "Handb. d. Natur.," 5th edition, p. 125. Hab.: South Africa. Lo.rodonta africana
Type fig.: Bhimenbach, Abbild. naturhist. Gegens., No. 19, fig. C.
Not in the edition of the "Handbuch der Naturgeschichte" of 1779 [First Edition] but
in the Fifth Edition (1797); in ".\bbild. Naturhist. Gegenstande," Heft 2, No. 19, fig. C,
the name is already used. The arrangement of the lamellae would indicate its Cape origin.
. .

It can only be a question of the Cape Colony and the Congo, perhaps of the French Congo.
The confluence of the anterior lamella is peculiar. (Matschie, letter, 1921.)
Sherbom (1902, Pt. I, p. 22) lists E. africanus as in the Fifth Edition of the "Hand-
buch" (1797.1, p. 12.5), but reference is not made to a figure in this edition. In 1923, Pt. II,
p. 135, he cites E. africanus in Blumenbach, Man. Hi.st. Nat., I, 1803, p. 155.

1798 Elephas indicus Cuvier "Tabl. El^men. Hist. Nat. Animaux." p. 148. Hab.: India. Sec Name preoccupied by
Cuvier's figure, with description, 1799, JMeni. Inst. Nat. Sci. ct Arts. Sci., Linnaeus, 1754
Math^m. et Phys., II, Fructidor, an VII [1799], p. 21, and Plates (ii, figs.
1,2: IV, fig. 1; v,fig. 1; vi. fig. 2).

1798 Elephas capensis Cuviev "Tabl. Elemcn. Hist. Nat. ,\nimaux," p. 149. Hab.: South Africa. Loxodonta africana
Type fig.: Cuvier, 1799, Mem. Inst. Nat. Sci. et Arts. Sci., Mathem. et Phys., capensis
II, Fructidor, an VII [1799], PL in and PI. iv, fig. 2.
Elephas capensis G. Cuvier, "Tableau Elomen.," 179S. Cuviei liadat his disposal one
skeleton from the .Senegal and one skull from the Cape ["Cap de Bonne-Esporance,"
Cuvier, Mem. Inst, (de France) National des Sciences et Arts, sometimes called the Acade-
mie des Sciences, Vol. II, annee 7 (1799?)]. (Matschie, letter, 1921.)

1799 Elephas primtgenius Biumenhsich "Handb. d. Natur.," 6th edition, p. 697. Hab.: Siberia and Manimonteus primi-
northern Germany. No original type figure. genius

1799 M ammut ohioticum Blumenbsich Op. c/L, p. 698. Hab.: Ohio River, North America. Mastodon americanus

1799 Elephas manimonteus Cuvier (1796 MS.), published August-September, 1799, Mem. Inst. Nat. Manimonteus primi-
Sci. et Arts. Sci., Mathem. et Phys., II, Fnictidor, an VII [1799], p. 21, Pis. genius
V, fig. 2, and vi, fig. 1.

Falconer, 1868, "In the same year [1796] he [Cuvier] read a memoir [MS.]
II, p. 158:
at the first public meeting of the 'Institute,' butwhich was not published until 1806 [1799,
p. 21], in which the diagnostic marks are verj- pointedly expressed under the designation
of Elephas Manimonteus: [subsequently
. . . —
1806, .\nn. Mus. hist, nat., V'lII, p. 264] he
abandoned the name E. Manimonteus of his [MS.] memoir of 1796, and adopted the desig-
nation of Elephas primigenius, proposed by Blumenbach [Footnote: 'Voigt's Mag. 1803,
Band v, p. 16. '], in 1803 which is that now generally accepted among paleontologists." See,
however. Bibliography, Cuvier, 1799.1.

1801 Elephantus indicus Cuvier and Lac6pede "La Menagerie du Museum National d'Histoire Elephas indicus
naturelle ou Les Animaux Vivants," An X. Hab. : India.

1803 Elephas macrocephalus A. Camper Mastodon americanus

Original text of 1802 not available. Atlas of 1803 gives the name Elephas macrocepha-
lus under Fig. 18. Sherborn, 1922-1933,"Index .\nimalium" (1928, p. 3760) gives the follow-
ing reference: '"Elephas macrocephalus,' Camper, CEuvres de P. Camper, II, 1803, 18, f. n.
[non usu spec.]."
1384 OSBORN: THE PROBOSCIDEA

Reference in
Year Name Author Present Memoir
1806 Ja- drnnil .Udslodontr i'uvivv Ann. AIiis. hist, iiat., V'lII, j)p. 270, 293 (Cuvier, 1806.2). Hab.: Mastoilon amfriranus
Ohio l^iver, North America. Type fig.: Cuvier, ibid., PI. 49 [i], figs. 1-5.
Cuvier, ibid., jip. 270, 272: "Animal tros-voisin do Tclcphant, mais a maclicliores
herissees do gros tubercules, dont on trouve les os en diver-s endroits dcs deux ooiitinens, et
surtout pros des bords de i'Ohio, dan.s rAmerique Septentrionalc, improprenient nomme
Mammouth jmr habitans des Etats-Unis.
le.s Anglais et .Nous empnuitons le nom
jiar les . .

do mastodonte do deux mots grccs qui signifient dents mammelonnies, et qui cxpriment par
consequent son principal earaetere."
Cf Mastodonte de I'Ohio Cuvier, below,
. thi.s list.

1806 Mastodonte de I'Ohio Cuvier Ibid., p. 412 (Cuvier, 1806.3). Hab.: Ohio River, North America. Mastodon americanus
Cf. Le Grand Mastodonte Cuvier, 180fj, above, and Mastodon giganteum Cuvier, 1817,
below, this list.

1806 Mdstodoidc a dents etroites Cuv'in- hoc. cit. Type loc.: Simorre, France. Type fig. : Cuvier, ifow/., Trilophodon angusti-
V\. 66 [i], fig. 4. dens
Cf. .\[astodo>i anguxlidcus Cuvier, 1817, below, this list.

1806 I'elil ninslodonte Cuy'ivv Ibid., ]). 413. Type loc: Montabu.sard, France. Type fig. : Cuvier, Titricius tapiroides
ibid., PI. 68 [ill], fig. 6.

Loc. cit. Tyjje loc. : Saxony, Germany. Type fig. : Cuvier, ibid., F\. &7 Trilophodon angusti-
[ii], fig. 11. dens minulus
Cuvier applied the term "Petit mastodonte" to two distinct t.yj)es, from Montabusard
and Saxony, to which he sub.sequently assigned the names Mastodonte tapiroide, 1821, and
M. minutus, 1824, respectively (sec below, this list, and also M. parvus, 1834).

1806 Mastodonte des Cordilieres Cuvier Loc. cit. Type loe. : Near volcano of Imbaburra, Quito, Ecua- Cordillerion andium
dor. Type fig.: Cuvier, ibid., PI. 67 [ii], fig. 1.

This is the single type which Cuvier subsequently (1824) called Mastodon andium (see
below) and wliich Desmarest called M. cordillerarum. The first distinctive generic name
a|)i)lied to these animals was Mastotheriuyn Fischer (1814). Subsequent names: Dibelodon
Cope (1884.2), Cordillerion Osborn (1926). See List of Genera above.

1806 Mastodonte h II niboldieii Cuvier Loc. cit. Type loc: Near Concepcion, Chile. Type fig. : Cuvier, Cuvieronius humboldtii
ibid., PI. 67 [ii], fig. 5.

Cf. Mastodon humboldtii Cuvier, in Desmarest, 1818-1824, below, tliis list.

1807 lil< plui.s indnniiiiittli Link Beschr. Nat. Samnil. Univ. Rostock (4) 1807, 3 (fide Sherborn, 1928, Maynmotdeits primi-
p. 3845). genius

1808 FAephos prinnvi'us IMumenbach (In Adams, translation from the French by Sir Joseph lianks, Mammonleus primi-
Mag. (Tilloch), XXIX, London, ]>. 152) cited by Tilesius, 1815, p. 452.
I'hil. ; genius
Hab.: Lena Kiver, Siberia.
Cf. Elrplias hrachijrainphus Brandt, 1832, below, this list.

1808 Elephas minimus Nesti

.\llril)utcd by Falconer and Cautley, 1840, "Fauna Aiili(iua Sivalensis," letterpress, p.
13, to Nest i, but not found by the present author in the original reference. Name abandon-
ed by Nesti.

1808 llarpagmolherium cmindense Fischer de Waldlicim Prog. d'Invit. Seance, Pub. Soe. Imp. Nat. Mastodon americanus
.Moscou, September, p. 19. Hab.: Oliio River, North America.

1809 Mu.stotheiium ohiotieum (Blum.) Fi.sciicr de Waldheim Mem. Soc Imp. Nat. Moscou, II, ]>. [Mastodon americanus]
252.

1811 Elephas gigas (1. Perry ".\rcana," p. Ii and plate. [Elephas indicus]

1811 Elephas socotrus G. Perry Op. ciL, p. \i. [Indeterminate]

ISM .Uos<o</ieri»w »!<'(7a/o(/on Fischer de Waldlicim "Zoognosia," p. 340. Hab.: Ohio Hivcr, North Mastodon americanus
America.
Cf. Le Ciraiid .Mastodonte and Maslodontr ile I'Oliio Cuvier, 1806, al)ove, this list.

1814 .l/«.!><o</iermm feptorfo« Fischer de Waldheim Loc. cit. Type loc: Simorre, France. Trilophodon angusti-
dens
Cf. Mastodonte a dents etroites Cuvier, 1800, above, this list.
 NOMENCLATURE OF THE PROBOSCIDEA 1385

Reference in
Year Name Author Present Memoir

1814 MastotheriummirrodonV\^ich('r<.h'\Va,\d\\eim Loc.cii. Typeloc. : Montabiisard, France. Turicius lapiroides

Cf. I'elil maslodonle, a petites dents, Cuvier, 1806, above, also Mastodon lapiroidcs
Cuvier, in Desmarcst, 1822, below, this list.

1814 A/as/o</ier/wm /i?/orfon Fischer de Waldheim Op. ctl, p. 341. Hab.: Cordilleras. Cordillerion andium
Cf. Le Maslodonte des Cordilieres Cuvier, 1806, above, this list.

1814 Madoiheriiini hiijubohttii Fi.scher de Waldheim Loc. cit. Cuvieronius humboldtii

Cf. Maslodonle humboldien Cuvier, 1806, above, and Mastodon hitmholdtii Cuvier, in
De.smarest, 1818-1824, below, this list.

1814 Mastodon Mac rod mt Wafmcsquc Specchio Sci., II, No. 12, p. 182. Hal).: North America. Maslodon aincrican 11s

1814 Mastodon rhomboiden RafinQ^quo Lor. cit Hab.: Cordillcra.s. Cordillerion audi am

1814 Mastodon humboldianus Rafmoaqne Loc.cii. Hab.: South America. Cuvieroit ias h umboldtii

1814 M astodo7i Senodon Rafinciique Loc.cii. Hab.: France and Europe. Trilophudiiti a ngu.sli-
deiis

1814 Mastodon microdon Rafinesque Loc.cii. Hab.: Europe. (?) Turicius lapiroides

1815 FJephas primordialis Blumenbach (In Tilesiu.s, Mem. Acad. Imp. Sci. St. Petersb., (V), V, p. Mammonleits primi-
470). Hab.: (iermany and Siberia. See also A'. pr/worrf/a//s Brayley, 1831, geniiis
Phil. Mag., IX, pp. 411-418; Bronn's Neue.s .Jahrb., 1833, p. 372, and the
"Index palseontologieu.s" of von Meyer and Ciiippert, in Bronn's "Handbuch
eincr Geschichte der Natur," 1848, III, p. 455. Hab.: E.schscholtz Bay,
Alaska.

1817 Mastodon giganteum Q'uvier "Le Regne Animal," p. 233. Type loc: Big Bone Lick, Boone Mastodon americanus
County, Kentucky, North .Imerica.
Cf. Le Grand Maslodonle Cuvier, 1806, Maslodonle de I'Ohio Cuvier, 1806, .1/0.5/0-
therium megalodon Fischer, 1814, above, also Mastodon maximus Cuvier, 1824, Mastodon
ohioticum, 1832, and Mastodon americanus Leidy, 1868, below, this list.

1817 Mastodon angustidens Cuv'iei- Loc.cii. Typeloc: Simorre, France. Type fig.: Cuvier, 1806, Trilophodoii (lugiisli-

Ann. Mus. hist, nat., VIII, PI. 66 [i], fig. 4. deiis

Not until the year 1817 did Cuvier substitute the name Mastodon angustidens for his
"Mastodonte a dents etroites" of 1806. Cf. Mastotheriuiii leplodon I'ischer, 1814, above,
this list.

1818 Mastodon minor Cuvier (In Desmarest, Nouv. Diet. d'Hist. Nat., XIX, p. 446). Trilophodon angusti-
dens minulus
Cf. Petit Maslodonle Cuvier, 1806, above, and Mastodon minutus Cuvier, 1824, below,
this list.

1818-1824 Mastodon humboldlii Cuvier (In Desmarest, ibid., p. 447); Cuvier, 1821-1824, "Ossemens Fos- Cuvieronius humboldlii
siles," V, Pt. 2, p. 527. Type loc: Near Concepcion, Chile. Type de-
scription: Cuvier, 1806, Ann. Mus. hist, nat., VIII, p. 413. Type fig.:
Cuvier, ibid., PI. 67 [11], fig. 5.
Cf. Maslodonle humboldien Cuvier, 1806, also Maslolherium humboldlii Fischer, 1814,
above, this li.st.

1820 £'/ep/i as jftbaiMs Schlotheim "Die Petrefactenkunde," p. 4. Hab.: Germany. Mammonteus primi-
genius

1821 M astodonte tapirotde Cnvier "Ossemens Fossiles," I, p. 268. Type loc: Calcaire de Monta- Turicius lapiroides
busard, France.
Cf. Felit maslodonle Cuvier, 1806, above, this list. Not until the year 1821 did Cuvier
replace "Petit mastodonte" with the name tapirolde, subsequently written tapimides (Des-

marest, 1822, p. 386; Cuvier, 1821-1824, V, Pt. 2, p. 527).

1821 Elephas priscus Goldfuss Nova Acta Acad. Leop. Carol., X, Pt. II, PI. xliv. Type loc: Near Loxodonla africana
(?)Cologne, Germany.
Falconer (1868, II, p. 94) retained the name Elephas {Loxod.) priscus [=Hesperoloxo-
don anliquus of |)resent Memoir] for undoubted Pleistocene fossil teeth from Gray's Thur-
rock and elsewhere, although he states (p. 95) that the actual type of Goldfuss, 1821,
"conveyed to my mind a corresponding impression that the molar was probably of modern
origin." Subsequently Falconer (cf. Leith Adams, 1877-1881, pp. 1,2) abandoned the name
Elephas priscus Goldfuss, and it is now considered {fide Pohlig) as a synonym of Loxodonla
africana.
 . :

1386 OSBORN: THE PROBOSCIDEA

Reference in
Year Name Author Present Memoir

1822 Mastodon cordillfranim Desmiimst "Mammalogio," Seconde partie, p. 385. Cordillerioii nndium
Cf. Manloiionle des Cordili'eres Cvivicr, 1806, and .UtiKlolherium hijodon I'"i.schcr, 1814,
above, also M. andium Cuvier, 1824, l)clo\v, this list.

1822 Mastodon ininiis Desmarest Op. cit., p. 386. Trilophodon (uigudi-

dens miniitiis
Cf. Petit mastodonte Ciivicr, 1806, abovp, and Mastodon miniitiis Cuvier, 1824, below,

this list.

1822 Mastodon tnpiroides Cuvier (In Desmarest, loc. cit.). Also Cuvier, 1821-1824, "Oasemens Fos- Ttiririus tapiroides
.siles," V, Pt. 2, p. 527. Type loc: Montabusard, France. Type descrip-
tion: Cuvier, 1806, Ann. Mus. hist, nat., VTII, p. 411. Type fig.: Cuvier,
ibul., PI. 68 [ill], fig. 6.

Cf. Petit mastodonte Cuvier, 1806, Mastotherium microdon Fiseher, 1814, Mastodonte
lapiroide Cuvier, 1821, above, this list

1823 Klcplias anliquitdtis Kriigev "Geschichte der Urwelt," p. 832. Type loc: Tliiede, Germany. Hesperoloxodon
Type fig. : Breislak, 1820, "Lehrbuch der Geologie," II, p. 428. antiquus germanicus

1824 Mastmhii moj-iiniis Cuvier "Ossemens Fossiles," V, Pt. 2, p. 527. Type loc: Big Bone Lick, Ma.ftodon americanus
Boone County, Kentucky.
Cf. Mastodon giganteum Cuvier, 1817, above, this li.st.

1824 Mastodon And m Cuvier

ill Loc. cil. Type loc: Near Volcano of Imbaburra, Quito, Ecuador. Cordillerion andium
Type description: Cuvier, 1806, Aim. Mus. hist, nat., VIII, pp. 411, 413.
Type fig.: Cuvier, 1806, ibid., PI. 67 [ii|, fig. 1.

Cf. Mastodonte des Cordilihes Cuvier, 1806, above, this list.

The name Mastodon andium is erroneously dated by Trouessart and others as 1806.
We cannot find the name earlier than 1824. It is thus technically preoccupied by Masto-
therium tiijodon Fischer, 1814, and Mastodon cordillerarum Desmarest, 1822, but the name
Mastodon andium Cuvier is adopted, following de BlainviUe and Falconer (.see footnote on
p. 122 of Vol. I of the present Memoir).

1824 Mastodon humboldii Cuvier "Os.semens Fossiles," V, Pt. 2, p. 527. Cuvieronius humboldtii
Cf. Mastodon humboldtii Cuvier, in Desmarest, 1818-1824, above, this list.

1824 Ma.stodon minutiis Cuvier Loc. cit. Type loc: Saxony. Type description : Cuvier, 1806, Ann. Trilophodon angusti-
Mus. hist, nat., VIII, p. 411. Type fig.: Cuvier, ibid., PI. 67 [ii], fig. 11. dens minutus
Cf. Petit mastodonte Cuvier, 1806, M. minor Ctivier, in Desmarest, 1818, above, and M.
parvus Cuvier, in Hays, 1834, below, this list.

1824 M a.stod on tiiricense Schinz "Naturgesch. u. Abbild. d. Siiugethiere," p. 278. Type loc: Elgg, Turicius turirensis
Canton Zurich, Switzerland. Type fig.: Schinz, 1833, Denk. .schweiz. Ges.
Naturw., I, Abth. 2, Taf. i, fig. 1, and p. 59.

1825 Elephas meridionalis 'Ncsti Nuov. Giorn. Lett., XI, No. 24, p. 211. Type loc: Val d'Arno Archidiskodon meridion-
northern Italy. Type figs.: Nesti, I'bfrf., Tav.
sup<!rieure, i, figs. 1, 2 (lectotype alis
cranium C); Tav. i, fig. 3 (cotype cranium A).

1828 Mn.'itodon ari'prnensis Croizet and .]ohevt "Ossemens Fossiles ... Puy-de-Dome," p. 138. Type Anancus arvernensis
loc: Perrier, Auvergne, France. Cotype figs.: Croizet and .lobert, op. cit.,
PI. I, figs. 1-4, PI. II, fig. 7.

1828 Mastodon latidensVliit Trans. Geol. Soc London, (2), II, Pt. Ill, pp. 369-375. Type loc: Stegolophodon Inlidens
Near Yenangyaung, Irrawaddy River, Burma. Lectotype and cotype figs.
Clift, ibid., PI. xxxvii, fig. 1, and PI. xxxviii, fig. 1.

1828 }[n.'<lodon elephajitoides C\U{ Ibid., pp. 372, 373. Type loc: Near Yenangyaung, Irrawaddy Stegodon e.lephantoides
River, Biuma. Lectotype fig.: Clift, ibid., PI. xxxviii, fig. 2.

1829 Dei notheiiuni giganteum K&np Isis, [XXII], Heft IV, p. 401. Type loc: Eppelsheim, Germany. Deinotheriamgigan-
Typefig. : Kaup, ibid., T&i. i. teum

1829 Elephas nidnunnntcns Fischer de Waldheim Nouv. M6m. Soc. Imp. Nat. Moscou, I, pp. 285, 286. Mammnnteus primi-
Ilab.: Russia. genius
Cf. Elephas mammonteus Cuvier (MS. 1796, published 1799) above, this list.
 " '

NOMENCLATURE OF THE PROBOSCIDEA 1387

Reference in
Year Name Author Present Memoir
1829 FAephns pnnisvuft Fischer de Waldhoim Ibid., pp. 285. 289. Halx: Volga, Rii.ssia. Mawmoiilciis pn'ini-
Written jiUo panicus. 0*'" "•'*

1829 £'Zep/ias per/6ofe<es Fischer de Waldheim 76 /rf., pp. 285, 290. Typeloc: Podolia, Rus.sia. Type Mammonteus primt-
fig. : Fischer, ibid., Tab. xvii, fig. 1. geniu.s
Also written proboletes.

1829 £/e/)/ifl.s p(/(/wa'«« Fi.scher de Waldheim /6(d., pp. 285, 292. Typeloc: District of Calomna and Manimonteii.t primi-
of Zwenigorod, Ru.s.sia. Type fig. : Fischer, /bid., Tab. xvii, fig. 2. (leniu.s

1829 jE'/ep/ifls campj/Zotes Fischer de Waldheim /6/(/., pp. 285, 291. Typeloc: Borders of the Bug. MammonieuH primi-
Russia. genim
1829 7!J/ep/ias A'aw(e/(sA-;'( Fischer de Waldheim Ibid., p. 27Q. Ilab.: Siberia. Mammoideim primi-
geniux

1830 Mammuthus boreali.'i Burnett Quart. Journ. Sci., London, p. 352. [Maminoiileus{'?)
primigen iii.s]
1830 T. [Tetracaulodon] M astodontoideum Godman Trans. Amer. Phil. Soc, N. S., Ill, jij). 478-485. Mastodon americantix
Typeloc: Near Newburgh, Orange County, New York. Type figs. : Cod-
man, ibid.. Pis. XVII, x\in.

1831 M. [Madodon] inlermedius Eichwald "Zoologia Spec," HI, p. 361. Type loc: \'olhynia, Russia. Anancua iiUermediiiK
Type: Maxilla (no figure found). Paratype: .law (supplementary de-
.scription, 1835, Nova Acta .\cad. Leop. Carol.. XMI, ]). 737, Pis. lviii, lix).

1831 Dinotherium moximnm Kaup "P'ossil Saugeth. Rheinhessens." {Fide de Blainville. 1839-1864, Deinolheriinn gigon-
"Osteographie," p. 18, on Dinotherium; fide von Meyer, 1832, "Palaeologica tea in
z. Ce.schichte der Erde," p. 78). Ty])e loc: Epjielsheim, Germany.

1831 Deinolheriinn Bararirum von Moyor Neues Jahrb. Min., p. 297. Typeloc: Gmiind, Bavaria. Deinotherimn bnrnri-
Supplementary description: von Meyer, 1832, Nova Acta Acad. Leop. Carol., ckih
XVI, Pt. II, p. 487, Tab. xxxiv, figs. 12-15, Tab. xxxvi.

1831-1833 Eleplta.'^ primordial i.'t Bravley ( "f . Elephas primordialis Blumenbach, in Tilesius, 1815, above, this
list.

Elephas priinordialis Brayley, 1831 (without iiamo); in Broun, 18,3.3, who attribiitps
the name to Brayley.

1832 Mammul Sibiririan von Meyer "Palseologica z. Gcschichte der Erde," p. 64. Mammonteux primi-
gen ins
Von Meyer attributes this name to Schlotheim.

1832 Dinotherium maximum yon Meyer (Ex MS. Kaup), "Palseologica z. Geschichte der Erde," p. 78. Deinotherinm gigan-
Typeloc: Eppelsheim, Germany. teum

1832 Dinotherium Cuvieri Ksiup "Description d'Ossements Fossiles," Cahier I, i)i). 2, 14. Hab.: Deinolherium. cuvieri
Comminge, Carlat-le-Comte, Chevilly, France.

1832 Tetracaulodon longirostre Ka.up Lsis, [XXV], Heft VI, p. 628. Typeloc: Eppelsheim, Germany. Tetralophodon longi-
Type fig.: Kaup, ibid., Taf. xi, fig. A. rndris
At thought to he referable to Mastodon anguslidens. Kaup, however, .substitutetl
first

the name Tetracaulodon hmgirosire, which subsequently (1835, p. 6.3) he changed to Masto-
don longirostris (see below, this li.st) by which name it has since been known. Its reference
in the present Memoir is to Tetralophodon longirostris.

1832 Mastodon ohioticum (In Bronn, Neues Jahrb. Min., p. 355. In Gervais, 1848-1852, Zool. Pal. Mastodon americanus
Frangaises, I, p. 187.)

1832 Elephas brachyramphus Brandt Mem. Acad. Imp. Sci. St. Petersb., (6), II, Math, et Phys., Bull. Mammonteus primi-
Sci., No. 2, p. xi. Type loc: Mer glaciale, near mouth of Lena River, genius
Siberia. Type: Adams .skeleton in the Zoological Museum of the Academy
of Sciences, Leningrad, U. S. S. R. Figured by Tilesius, 1815, Mem. Acad.
Imp. Sci. St. Petersb., (V), V, Tab. x. Described by Adams, 1808, Phil. Mag.
(Tilloch), XXIX, and Tilesius, 1815.
Cf. Elephas prinisevus Blumenbach, in Adams, 1808, above, this list.

1832 Elephas homotaphrus Brandt Ibid., p. xii (name only).

 . ::

1388 OSBORN: THE PROBOSCIDEA

Reference in
Year Name Author Present Memoir

1832 Klfiilias gigiiiilrii.s Brandt hoc. cil. Typoloc: ludigiika River, Siberia. Type: Messerschmidt Mammonteus pritni-
cranium. Figured by Breyiie, 1741, Phil. Trans. Roy. Soe. London, XL, PI. i, genius
figs. I, II ; and by Ciivier, 1806, "Ossemens Fossiles," PI. 39, fig. 1, and PI.
41, fig. ll.F.

1832 FAe-phas commutatus Bxa.nAi Loc. cil. Typeloc.: "Bords du Volga." Type: "Crane du Comte Mammonleus primi-
Mussin Puschkin." Figured by Cuvier, 1825, "Recherches Ossemens Fos- genius
siles," 3d edition, I, PI. ix, fig. 7, and p. 179.

1832 Elephas stenotoechus Brandt Ibid., p. xiii. No locality given. Type in Zoological Museum of the Mammonteus primi-
Academy of Sciences, Leningrad, U. S. S. R. genius

1832 Elephas platytaphrus Brandt Ibid., p. xiv. No locality given. Type in Zoological Museum of Mammonteus primi-
the Academy of Sciences, Leningrad, U. S. S. R. Type fig. : Cuvier, 1825, genius
"Recherches Ossemens Fossiles," 3d edition, I, Pi. ix, figs. 5, 6.

1832 Elephas ajjUnis Brandt Ibid., p. xiv.

The history of this six>fimen (a skeleton in the Zoological Museum of the Academy of
Sciences, Leningrad, U. S. S. R.) given in Oken's Isis of 1832, XXV, Heft X, pp. Ull-
i.s

1 114, by a writer signing himself "Ein Natiirforscher in St. Petersburg." He states that
this skeleton together with other materials was originally discovered in a hole beneath an
oven of a house in St. Petersburg which was built upon ground formerly a swamp and which
was torn down in 1828. This anonymous reviewer refers to the author of the description as
"Herr B" and expresses doubt as to the specimen being a fossil.
See also Neues Jahrb. Min., 1833, p. 611.

1833 Dinotherium medium Kaup Neues Jahrb. Min. p. 419. Type loc: Eppelsheim, Germany. Deinotherium medium
Type fig.: reproduced in Kaup, 1835, "De-
Kaup, ibid., Taf. vii, fig. 1,
scription d'Ossements Fossiles," Cahier IV, Add. Tab. i. Supplementary
description: Kaup, 1833, Neues Jahrb. Min., p. 509.

1834 M. [Maslodon] parvus Cuvier (In Hays, Trans. Amer. Phil. Soc, N. S., IV, p. 333.)
Cf . Petit mastodonte Cuvier, 1806, of Saxony, above, this list

1834 M. [Mastodon] BorsoniHsLys Ibid., p. 334. Type loc: Near Villanova d'Astica, Piedmont, Zygolophodon borsoni
Italy. Type fig.: Borson, 1823, Mem. Accad. Sci. Torino, XXVII, Tav. ii

(as M. giganteum).

1834 M. [Maslodon] Cuvieri Hays Ibid., pp. 322, 323, 334. Fragment of lower jaw in cabinet of the Maslodon americanus
American Philosophical Society. Type fig. : Hays, ?6?rf., PI. xxiv.

1834 M. [.Mastodon] Jeffersoni Hays Ibid., pp. 323, 334. Fragment of right lower jaw and portion of Mastodon americanus
left ramus in cabinet of the American Philosophical Society. Type fig.
Hays, ibid., PI. xxv.

1834 T. [Telracatdodon] Collinsii Hays Ibid., pp. 326, 327, 334. Portion of right lower jaw in cabinet Mastodon americanus
of the American Philosophical Society. Type fig. : Hays, ibid., PI. xxviii.

1834 T. [Tetracaulodon] Godmani Hays Ibid., pp. 327, 334. Fragment of right lower jaw in cabinet of Mastodon americanus
the American Philo.sophical Society. Type fig.: Hays, ibid., PI. xxix.

1834-1843 Maslodon Chapmani Hays Ibid., figs. 3 and 4, p. 338 (without name);
explanation of PI. xxii, Stegomastodon chap-
name used by Hays, 1843, Proc. Amer. Phil. Soc, II, p. 270. Type loc: mani
Unrecorded locality in i\w United States. Type lost or misplaced. Type fig.
Hays, 1834, Trans. Amer. Phil. Soc, N. S., IV, PI. xxii, figs. 3 and 4.

1834 Elephas mnrrnrt/nrhus Movmn "M6ni. Ossemens Fossiles fil6phans Relg.," p. 23. Type loc: Mammonteus pri mi-
Tamise, Belgium. Type fig.: Morreii, op. cil., PI. ii, figs. 1-4. genius

1834 Maslodon dubius Kaup and Scholl "Verzeichniss der Gypsabgiisse von den ausgezeichnetsten Telralophodon longi-
urweltlichen Thierrcsten des Grossherzoglichen Mu.seum zu Darmstadt," p. rostris
22. Type loc: Eppelsheim, Germany.
Mastodon dubius ia a synonym of M. longirostris (fide Kaup, 1835, p. 77).

1834 Masiorfon granrfjs Kaup and Scholl Op. cil., p. 25. Type loc: Eppelsheim, Germany. Type Telralophodon longi-
fig. : Kaup, 1835, "Description d'Ossements Fossiles," PI. xviii, fig. 9. rostris
Mastodon grandis is a synonym of M. longirostris (fide Kaup, 1835, p. 77).
 :

NOMENCLATURE OF THE PROBOSCIDEA 1389

Reference in
Year Name Author Present Memoir
1835 Elephafi odontotyrannus Eichwsild Nova Acta Acad. Loop. Carol., XVH, p. 723. Type loc: Mammonieii'i priini-
Banks of Niemaii River, Di.st. of Novogrodek, gouv. Vilna, Russia. Type genius
fig.: Eichwald, ibid., PI. lxiii, fig.s. 1, 2.

1835 Mastodon podolicum Eichwald Ibid.,p.73Q. Type loc: Near Tult.schin, Podolia, Russia. Type Deinolherinm podoli-
figs. : Eichwald, ibid., Pis. lvi, lvii. cum
See Vol. I, p. 85 (footnote) of the present Memoir.

1835 Dinotherium proavuni Eichwald Ibid., p. 741. Hab.: Podolia, Russia. Type fig.: Eichwald, Deinolherium proavus
ibid., PI. LX, figs. 1-5.
First mentioned as T.(Tapirus) proavus by Eichwald in 1827 (Naturhistorische Skizze,
p. 239 — not available to the present author). Described as Tapirus proavus in "Zoologia
Specialis," 1831, III, pp. 3.'53, 360. See de Blainville, 1839-1864, "Du Dinotlierium," pp.
12, 19.

1835 Dinoiherium iimlense Eichwald Ibid., p. 742. Hab.: Ural Mts. Deinotherinm umlense
See Pallas, 1777, p. 213, Tab. ix, fig. 4; .also de Blainville, 1839-1864, p. 19; and Lar-
tet, 1859, p. 482.

1835 Mastodon longirosiris Kauj) "Description d'Ossemenis Fossiles," Cahier IV, p. 65. Tetralophodon lonqi-
rosiris

Cf. Teiracaulodon longirostre Kaup, 1832, Mastodon dubius Kaup, 1834, and Mastodon
grandis Kaup, 1834, above, this list.

1836 Dinotherium serundarium Kaup (In Lartet, Bull. Soc. geol. France, (1), VII, p. 218.) Type Deinotherium secun-
loc: Simorre, France. darium{?)
Cited also by de Blainville, 1839-1864, "Du Dinotherium," p. 19. Original description
by Kaup not found by the present author.

1836 M. tapiroides-minus Jjurtet Loc. cit. Type loc: Simorre, France. Turicius tapiroides-
minus

1836 M. angustidejis minus Lartet Loc. cit. Type loc: Simorre, France. Trilophodon angusti-
dens minutus

1836 M. [M astodon] Sivalensis Ca,nt\Gy Journ. Asiatic Soc. Bengal, V, p. 294. Type loc: Doab Pentalophodon siva-
Canal, vicinity of Nahun, India. Type fig. Cautley, /6(rf., PI. xi, figs. : 2, 3. lensis

1838 E. [Elephas] jacksoni Mather "First Annual Rept. Geol. Survey Ohio," pp. 96, 97 (notice of di.s- Parelephas jacksoni
covery, without name); Amer. Journ. Sci., (1), XXXIV, p. 358 (description,
without name) ibid., pp. 362-364 (final description, with name). Type loc.
;

Salt Creek, Jackson County, Ohio. Type fig.: Mather, Amer. Journ. Sci., (1),
XXXIV, p. 363, fig. A. Present location of type specimen unknown.
1840(?) Missourium kochii Koch "Fossil Remains," p. 2. Type loc: 22 miles south of St. Louis, Jeffer- Mastodon aniericanus
son County, vicinity of Sulphur Springs, Missouri.
The type skull was found in May, 1839, and
first described (Amer. Journ. Sci., (1),

XXXVII, a subsequent paper ("Fossil Remains,"

pp. 191, 192) as Koch's Mi.ssourian; in
supposedly of date 1840, p. 2) Koch published a supplementary description, assigning the
name Missourium kochii. See Horner, 1840, Proc. Amer. Phil. Soc, I, pp. 279-283, for
remarks on the collection of Koch.

1841 Leviathan Missourii Koch "Description of the Missourium, or Missouri Leviathan," p. 13. Type Mastodon aniericanus
loc: Near the shores of the river La Pomme de Terre, a tributary of the
Osage River, Benton County, Missouri.
Skeleton excavated in March, 1840, and regarded by Koch as belonging to the same
genus, namely, Missourium; he named it, however, Missouri Leviathan (1841, p. 13).

Changed in 1843 to Missourium Theristocaulodon (see below, this list).

1841 T. [Tetracaulodon] Osagii Koch "Description of the Missourium, or Missouri Leviathan," Mastodon americanus
another edition, p. 1. Hab.: Missouri.

1841 Teiracaulodon Tapyroides Koch Loc. cit. Hab.: Missouri. Mastodon americanus

1841 Elephas indicus Isodactylus Hodgson Journ. Asiatic Soc. Bengal, N. S., X, p. 907 (name without Indeterminate
definition).

1841 Elephas indicus Heterodactylus Hodgson Loc. cit. (name without definition). Indeterminate
1390 OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir

1841 Dinolherutm Koiiigii Kaup "Aktcn dor Urwelt," pp. 49, 50. Typploc: Eppelshcim, Gprmany.

1841 Dinotheriumminutum von Meyer Neues Jahrb. Min., p. 459. Typeloc: Mo.sskirch, Germany.

1842 T. [Tetrac.aulodo7i] kochii Koch Proc. Geol. Soc. London, III, p. 715. Hab.: Missouri. Mastodon americanus
Koph, 1842, p. 715: "It does not require a close examination. to perceive that the . .

animal to which these remains belonged was neither male, female, nor young Mastodon,
or Missourium, the whole inner and outer conformation of the upper tusks showing that
they were calculated tobeased in harmony with the lower tusk in grubbing and rooting."

1842 Tetiaraulodnn flaysii Grant Ibid., Ill, Ft. II, p. 771. Mastodon americanus
Cf. Kocli, 1845, p. 29.

1842 TetracaulodoH Hucklandi Grant Loc. cit. Mastodon americanus

Cf. Koch, 1845, p. 29.

1842 Mastodon Brasiliensis Lund (In Lesson, "Nouv. Tabl. Regne Animal," p. 157.) Type loc: Cuvieronius brasili-
Valley of the Velhas River, Province of Minas Geraes, Brazil. ensis
Mastodon Brasiliensis, the specific name of which is attributed to Lund by Lesson,
appears as "Mastodon sp." in Lund (1839, p. 133), compared with M. andium and M.
tmmboMlii but net figured bv Lund. See Lund, 1839, pp. 117, 129, 130, 133 ("12. Mastodon
sp.").

1842 E.[Elephas\ ameriranus Y)e Kay "Natural History of New York," p. lOL Type loc: Irondi- Mammonteus primi-
quoit River, Monroe C'ounty, near Rochester, New York. Was in cabinet of genius americanus
the Lyceimi of Natural History, New York, but was destroyed by fire. Type
fig.: De Kay, op. cit., PI. xxxii, fig. 2.

1843 Missourium Theristocaulodon Koch "Description of the Missourium Theristocaulodon," pp. 9, Mastodon americanus
15, frontispiece.
First named by Koch, 1841, Leviathan Missourii.

1843 Leviathan rnissouriensis Koch Op. cit. (in title). Mastodon americanus
Corrected form of Leviathan Missourii but changed to Missourium Theristocaulodon in
1843 (see preceding item).

1843 Dinotherium Australe Owen Ann. Mag. Nat. Hist., XI, pp. 329-332, figs. 1, 2. Type loc: Diprotodon
Darling Downs, Australia.
See Owen, 1843, ibid., pp. 7 12, espt^cially p. 9, figs. 2 and 3, for first description of
femur and molar but without name. See also Owen, 1844, ibid., XIV, p. 268 (Dinottierium
australe = Diprotodon australis).

1844 Mastodon au.'ftralis Owen Ibid., XIV, p. 271, figs. 1 and 2 on p. 269. Type loc: Specimen Diprotodon
brought by native to Count Strzlecki in Australia, from cave further in the
interior than the ossiferous caves of the Wellington valley.
Probably a South American specimen accidentally misplaced with Australian speci-
mens by this traveler (Count Strzlecki). See Falconer, 18.57, table opposite p. 319, and
1868, II, pp. 271 276; also Ja<'k and Etheridge, 1892, p. 683.

1845 Mastodon arborense Koch "Die Riesenthiere der Urwelt," p. 18. Mastodon americanus
Apparently erroneously ascribed to von Meyer. See von Meyer and Goppert, 1848, p.
705: "[.Mastodon] Arboreiu<e [ruis<iuam Myr.) Koch Riesenthiere 18" Also p. 706:
"Mastodon Arborense (Mey.) Koch [err. typ. ?pro| = Ma.stodon Arvernensis."

1845 Mas^todon rugatum Koch Op. cit., p. 20. Mastodon americanus

See also Giebel, 1847, p. 202.

1845 Dinotherium arigustidens Koch Op. cit., ]). 41. Type loc: ('()nii)ubay [rambay(?), cf. pp. 85 Deinothcrium Hp.{?)
and 90 of present Memoir], India.
See von Meyer and Goppert, 1848, pp. 424, 425: "Dinotlicrium. . . angustidens Koch
= Mastodon angustidens Cuv."
 NOMENCLATURE OF THE PROBOSCIDEA 1391

Reference in
Tear Name Author Present Memoir
1845 Elephas kamensis de Blainville "Osteographie," p. 202. Mammonteus primi-
genius
De Blainville cites this species as Elephas Kamenskii or kamensis (see Elephas Kamen-
skii Fischer, 1829, above, this list).

1845 Elephas africanus prisms de Blainville Op. cit., p. 205.

European specimens considered by some palaeontologists to be referable to Loxodonta
africana, by others as of doubtful determination. Reviewed by Pomel, 1895.1, p. 20.

1845 Elephas ohioticiis de B\&invi\\e Op. cd, p. 261. Hab.: Ohio River. Mastodon americanus

1845 Dinotherium intermedium de B\a,mvi\\e Op. Ci7., Atlas, PI. in. Hab.: France. Deinotherium inter-
medium
1845 *E. [Elephas] primigenius sibiricus de Blainville Op. oil., Atlas, PI. iii. Hab.: Siberia. Mammonteus primi-
See also Deperet and Mayet, 1923, pp. 183-201. genius

1845 *E. [Elephas] primigenius germanicus de Blainville Op. cit., Atlas, PI. in. Hab.: Germany.

1845 *E. [Elephas] indicus ceylanicus de BlsLUwiWe Op. cit., Atlas, Pi. iii. Hab.: India, Ceylon. Elephas indicus
Living form. ceylanicus

1845 *E. [Elephas] indictisbengalensis deBlsbinville Op. cit., Atlas, PI. in. Hab.: India, chiefly Elephas indicus
Bengal and Assam. Living form. hengalensis
Compare Falconer and Cautley's figures (1846 [1847, oiE. indicus (Dauntela
PI. XLiij)
var.) and E. indicus (Mukna var.), also P^alconer (1867,
p. 57) in which he presents a de-
tailed comparison of the measurements between the Mukna and Dauntela varieties of
Elephas indicus. He adds (p. 58) "The plates of teeth in the Mukna variety slope greatly
:

backwards and are excessively and finely crimijed; those of Dauntela are much less
crimped."

1845 *E. [Elephas] primigenius meridionalis de Blainville Op. cit., Atlas, PI. in.

1845 Dinotherium [gig.] majus de Blainville "Osteographie, Du Dinotherium," p. 60. Deinotherium gigan-
See Weinsheimer, 1883, p. 210. teum

1845 Dinotherium [gig.\ medium de Blainville hoc. cit. Deinotherium gigan-

See Weinsheimer, 1883, p. 210. teum

1845 Dinotherium [gig.] minus de Blainville Loc. cit. Deinotherium gigan-

See Weinsheimer, 1883, p. 210. teum

1845 Dinotherium indicumY aXcowev Quart. Journ. Geol. See. London, I, p. 361. Type loc: Perim Deinotherium indicum
Island, India. Type fig.: Falconer, ihid., PI. xiv, figs. 1, la.
Falconer, 1845, pp. 370, 371: "The Dinotherium of Eppelsheim is known to range
through a very wide difference of size, dependent on sexual or individual peculiarities, and
several nominal species, chiefly founded upon this character, have been described by
authors. But Dr. Kaup informs me, that he now admits but two species, D. giganteum and
D. Komigii, as he regards all the rest, such as D. Cuvieri, D. Bavaricum, D. proavum, &c., to
be merely dwarfed varieties, or females of D. giganteum. M. De Blainville has arrived
at nearly the same conclusion in his O.steographip. It would be unsafe, therefore, to found
any opinion regarding the Indian fo.ssil merely on a difiference of size."

1845 Elephas Hysudricus Falconer and Cautley "Fauna Antiqua Sivalensis," PI. i, fig. 3a (type), fig. Hypselephas hysudricus
1846 36 (paratype). Hab.: Siwalik Hills, India. Type description Falconer and :

Cautley, 1846, "Fauna Antiqua Sivalensis," letterpress, p. 41.

1845 E. [Elephas] planifrons Falconer and Cautley Op. cit., PI. n, fig. 5o (lectotype), fig. 56 (cotype). Archidiskodon plani-
1846 Hab.: Sivvahk Hills, India. Type description Falconer and Cautley, 1846, : frons
op. cit., letterpress, p. 38.

1845 Elephas insignis Falconer and Cautley Op. cit., PI. n, fig. 6a (lectotype), fig. 66 (cotype). Hab.: Stegodon insignis-
1846 Siwalik Hills, India. Type description: Falconer and Cautley, 1846, op. cit., gane.sa
letterpress, pp. 37, 38.

*These perhaps may be regarded as geographic designations rather than as subspecies."

1392 OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir
1845 Elephas gnnean Falconer and Cautloy Op. cit., PI. iii, fig. 7a. Hab.: Siwalik Hills, India. Type Stegodon insignls-
1846 description: Falconer and Cautley, 1846, op. r/7., letterpress, p. 45. ganesa

1846 Elephas minimus Nesti (In Falconer and Cautley, 1846, op. cit., letterpress, p. 13.)
Falconer attributed Elephas minimus to Nesti, 1808, but the present autlior was
unable to find this name in eitlier of Nesti's articles of 1808 or 1825.

1846 .fi^fep/ias .Vamarficus Falconer and Cautley Op. cit., letterpress, p. 45. Hab.: Valley of the Palxoloxodon namadi-
Nerbudda River, India. Type figs. Falconer and Cautley, 1847, op. cit., : cus
Pis. XII. A, and xii.b, figs. 1 and 3. Supplementary description: Falconer,
1867, "Description of the Plates in the Fauna Antiqua Sivalensis," p. 15.

1846 E. [Elephas] priscusi Falconer and Cautley Op. cit., letterpress, p. 45 (name only); also PI. xiv, Ilesperoloxodon
1847 figs. 7, 7a, 76. E. [Elephas] (Loxod.) priscus (Goldf.) Falconer, 1857, Quart. aidiqiius
1857 .Journ. Ceol. Soc. London, XIII, pp. 345, 346, and table opp. page 319. Type
1867 loc: Cray's Thurrock, England. Type description: Falconer, 1867, "De-
1868 .scription of the Plates in the Fauna Antiqua Sivalensis," p. 21. Type fig.:
Falconer and Cautley, 1847, op. cit., PI. xiv, figs. 7, 7a, 76. Supplementary
description: Falconer, 1868, "Palaeontological Memoii-s," II, ji. 94, and PI.
VII, as Elephas (Loxod.) priscus.

Name preoccupied by Goltlfuss, 1821 (see above, this list).

1846 E. [Elephas] bombifrons Falconer and ('autley Op. cit., letterpress, p. 46. Hab.: Siwalik Hills, Stegodon bombifrons
India. Lectotype fig. Falconer and Cautley, 1847, op. c('<., PI. xxvi; cotypc
:

figs. Pis. xxvii, xxviii.

:

1846 E. [Elephn.s] cliftii Falconer and Cautley Op. cit., letterpress, p. 47. Type loc: Near Yenang- Stegodon elephant-
yaung, Irrawaddy River, Burma. Type fig.: Clift, 1828, Trans. Geol. Soc. oides { = cliftii)
London, (2), II, Pt. Ill, PI. xxxix, fig. 6, as M. elephantoides.

1846 Mastodon hrevirostre Gervais and de Serres Ann. Sci. Nat., (3), V, p. 268. Type loc: Montpel- Anancus arvernensis
lier, H^rault, France. Cotype figs.: Gervais, 1859, "Zoologie et Pal^ontol- brevirostris
ogie Frangaises," Deuxieine ed., PI. i, fig. 3, PI. iii, fig. 7 (same as in First
Edition of 1848-1852).
Until it is positively determined by further research that the "M." brevirostris of Mont-
pellier is identical with the ".1/." arvernensis of Auvergne, it seems best to retain this form as
a subspecies of Mastodon [Anancus] arvernensis.

1846 Elephas minimus Giehel Neues Jahrb. Min., p. 459. Type loc: Seveckenberg near Quedlin-
burg, northern Germany.
See Giebel in Isis, 1845, Heft VII, p. 483, Heft XII, p. 905, also in Fauna der Vorwelt,
1847, 1, Abtli. 1, 1). 211, for description but without name.

1847 £Zep/ias anii^Mws Falconer and Cautley "Fauna Antiqua Sivalensis," PI. xii.d. E. (Eueleph.) Ilesperoloxodon
1857 antiquus Falconer, 1857, Quart Journ. Geol. Soc, London, XIII, Synop. Tab. antiquus
1867 opp. p. 319. Hab.: Locality not recorded, but undoubtedly England. Typo
description Falconer, 1867, "Description of the Plates in the Fauna Antiqua
:

Sivalensis," p. 18. Type fig.: Falconer and Cautley, 1847, "Fauna Antiqua
Sivalensis," PI. xii.d, figs. 4, 4a.
Elephas antiquus misnamed E. meriJionalis on i)late (see Falconer, 1867, |). 18, and
1868, I, p. 438, together with legend), but correctctd by Falconer in <opy of the "Fauna
Antiqua Sivalensis" belonging to the British Museum. See also Falconer, 1868, II, pp.
176-188.

1847 Mastodon perimensis F&\coni:v iiiid Vantlvy Op. cit., PI. xxxi, figs. 9, 9a. Type loc: Perim Anaitcus perimensis
Island, India. Ty])e descri])tion: Falconer, 1867, "Description of the Plates
in the Fauna Antiqua Sivalensis," p. 44. Type fig-: Falconer and Cautley,
1847, op. cit., PI. XXXI, figs. 9, 9a.
Lydekker chose as type Brit. Mus. M.2882 (Pis. xxxviii and xxxix of the "I'^iuiia
Antiqua Sivalensis"), regarded as the paratype in the present Memoir.

1847 £?ep/ias /ndiCMS (Dauntela var.j Falconer and Cautley Op. cit., PI. xlii, fig. xxiii.a. Hab.: Elephas indiciis var.
India. Type; descrii)tion: Falconer, 1867, op. cit., p. 57; 1868, op. cit., I, p. Dauntela
477. Living form.
 NOMENCLATURE OF THE PROBOSCIDEA 1393

Reference in
Tear Name Author Present Memoir
1847 E'/ep/jos /nrf/cMS (Mukna var.) Falconer and C'autlcy Op. n't., PI. xlii, fig. xxiii.b. Hab.: Elephas indicus v&r.
India. Type description: Falconer, 1867, loc. cit., 1868, loc. cit. Living Mukna
form.

1847 Mastodon vellavus Aymard Bull. Soc. geol. France, (2), IV, p. 414. Type loc: Velay, France. Zijgolophodon borsoni
No record of figure. vellavus
While the author stated on p. 193, Vol. I, of the present Memoir that he believed both
"M." vellaviii and "M." Vialelii to be referable to "M. " [Z ygohphodon] borsoni, he thought
it best nevertheless to retain these as subspecific names (p.632). Compare also Falconer,
1868, II, p. 20.

1847 Mastodon Vialetii Aymsird 76(rf., p. 415. Type loc: Vialette, France. No record of figure. Zygolophodon horsoni
See note above under Mastodon vellavus Aymard, 1847. Vtaletu

1847 Elephas simiatranus Tcmmmck "Coup d'ffiil General," II, p. 91. Type loc: District of Palem- Elephas indicus
bang, Sumatra. Type .skcleton.s in Leiden Museum. No record of original sumatranus
type figure (sec figure 1182 of present Memoir). Living form.

1848 Mastodon Cuvieri Vome\ Bull. Soc. g^ol. France, (2), V, p. 258. Hab.: Gers and I'Orleanais, Trilophodon angus-
France. tidens cuvieri
Synonym of Mastodon angustidens Cuv. (fide Trouessart, 1897, p. 700).

1848 Mastodon Biiffonis Pomel Loc. cit. Type loc: Auvergne, Perrier, Franre. Zygolophodon borsoni
Synonym of Mastodon borsoni Hays (fide Trouessart, 1897, \>. 70o). oujjoms

1850 Elephas affinis Eichwald "Palseont. Rossii," p. 179 {fide Sherborn, 1922, "Index Animalium,"
p. 124). See also Eichwald, 1853, "Lethaea Rossica," p. 350, PI. xi, fig. 36.
Type loc: Vicinity of Taganrogh, Russia, near river Mjousse. Original in
Mus. Inst, des Mines, Leningrad.

1851 M astodon Simorrense Lartet "Notice sur La Colline de Sansan," p. 24. Type loc: Simorre, Trilophodon angusti-
France. dens
Synonym of Mastodon anguslidens Cuv. (fide Trouessart, 1897, p. 700).

1851 Mastodon Gaujaci h&rtet Op. cit., p. 27. Type loc: Lombez, France. Trilophodon angusli-
dens gaujaci
Synonym of Mastodon angusiidens Cuv. (fide Trouessart, 1897, p. 700).

1852 Ma.dodon humboldlius Wavvcn "The Mastodon Giganteus of North America," p. 126. Hab.: Cuvieronius humboldtii
South America.
Warren states that Cuvier, de Blainvillc, and Owen regarded this species as referable to
angustidens.

1854 Elephas Rupertianus Hiclvdrdson "Zoology of Voyage of H.M.S. Herald," pp. 101, 102, 141. Mastodon americanus
Type loc: Swan River, basin of Lake Winnipeg, Canada. rupcrtianus

1855 Anancus macroplus Aymard (In Dorlhac, Ami. Soc. Agric Puy, XIX, for 1854, p. 507). Type Anancus arvernensis
loc. Mt. Coupet, near Puy, France.
:

Synonym of M. arvernensis (fide Lartet, 18.59, p. 493).

Deperet, 1885, p. 159: "Le nom d' Anancus macroplus donne jmr M. Aymard aux
molaires des sujets aduUes du bassin du Puy, tres distinctes des molaires de lait decrites par
Croizet et Jobert, doit egalement disparattre devant le nom plus ancien d' arvernensis."
See also Deperet, 1890, pp. 62, 66.

1856 Mastodon pentelicus Gsiudry and LartQt Compt. Rend. Acad. Sci., XLIII, p. 273 (name only). Trilophodon (Choero-
1862 Type loc: Pikermi, Greece. Type description: Lartet, 1859, Bull. Soc lophodon) pentelicus
1867 geol. France, (2), XVI, p. 497; Gaudry, 1862, "Animaux Fossiles," p. 142.
Type figs.: Gaudry, 1862-1867, op. cit., PI. xxii, figs. 1-3, also PI. xxiii.
The name Choerolophodon is provisionally retained as a subgenus of Trilophodon for the
species M. pentelicus, although the type of the species J\l pentelicus is close to Telralophodon
.

longirostris.
1394 OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir

1857 E. [Elephas] (jiganieus AymanI (In Falconer, t^uait. .louru. Geol. Soc. London, XIII, p. 321; and •4 rrh idiskodon mcridi-
Falconer, 1868, "Palseontological Memoirs," II, p. 20, footnote.) onalis{l)

1857 M. {Triloph.) Paiidionis Falconer Quart. Joum. Geol. Soc. London, XIII, p. 317 and Synop. Trilophodon pavdiovin
Tab. opp. p. 319. Type loc: Deccan, India [error, probably Larkana Dis-
trict of Sind {fide Hopwood, letter, Feb. 10, 1932)]. Supplementary de-
scription: Falconer, 1868, "Palaeontological Memoirs," I, p. 124. Type fig.:

Falconer, 1868, op. cit., PI. xxxiv, figs. 6, 7.

1857-1859 M. {Triloph.) Pyrenaicus Lartet (Ex MS. Lartet, in Falconer, Quart. Joum. Geol. Soc. London, Zygolophodon pyrenai-
XIII, Synop. Tab. opp. p. 319.) M. [Ma.stodon] pyrenaicus Lartet, 1859, cus
Bull. Soc. g6ol. France, (2), XVI, p. 513. Type loc: Near Ile-en-Dodon,
(Haute-Garonne), France. Type fig.: Lartet, 1859, ibid., PI. xv, fig. 4.

1857 E. [Elephas] {Eueleph.) Armeniacus Falconer Ibid., Synop. Tab. opp. p. 319. Type loc: Near Parelephas armeniacus
Khanoos, Province of Erzerum, Armenia. Supplementary description: Fal-
coner, 1863, Nat. Hist. Rev., Ill, pp. 74, 75. Type fig. : Falconer, 1863, ibid.,
PI. 11, fig. 2.

1857 E. [Elephas] {Loxod.) priscus {Gold}.) Falconer Ibid., pp. 345, 346, Synop. Tab. opp. p. 319; 1868, Hesperoloxodon an-
"Palaeontological Memoirs," II, p. 94, as Elephas {Loxod.) priscus. Type tiquus
loc. Gray's Thurrock, England. See above, E. [Elephas] priscus? Falconer
:

and Cautley, 1846, 1847.

Name preoccupied by Goldfuss, 1821 (see above, this list).

1857 E. [Elephas] {Eueleph.) antiquus Falconer See under Elephas antiquus Falconer, 1847, above,
this list.

1857-1868 E. [Elephas] {Eueleph.) Columbi Falconer Quart. Journ. Geol. Soc. London, XIII, Synop. Tab. Parelephas columbi
opp. p. 319. Type loc: Brunswick canal, near Darien, Georgia. Supple-
mentary description: Falconer, 1863, Nat. Hist. Rev., Ill, pp. 43-52, 114.
Type fig.: Falconer, 1863, ibid., p. 114, PL i; 1868, "Palaeontological
Memoirs," II, PI. x, fig. 1. Neotype loc: Phosphate Beds near Charleston,
S. C.

1857 Mastodon atiicus Gaudry and Lartet (In Wagner, Abh. bayer. Akad. Wiss., VIII, Abtli. I, CI. Turicius atiicus
II, p. 140.) Type loc: Pikermi, Greece.
Gaudrj', 1862, p. 142: "C'est par une inadvertance de copie que Wagner a employ^ le
nom A'atlicus; car il nous attribue ce nom, et le .scul que nous ayons propose est celui dc
penlelicus."

1858 [Mastodon] dissimilis Jourdan (MS. 1840) Ann. Soc. Imp. Agric. Lyon, (3), II, p. Ixxxiv. Type Aiwncus nrvernensis
loc: Saonc Basin, France. di.ssimilis

1858 Mastodon mirificus Leidy Proc Acad. Nat. Sci. Phila., X, p]). 10, 28. Type loc: "Loup Fork Slegotnaslodon mirificus
of Platte River,"Nebraska (Leidy, 1873, Rept. V. S. (^.eol. Surv. Terr., I, p.
330) possibly near Seneca, Thomas County, Nebraska {fide Hay, 1924, Publ.
;

Carnegie Inst. Wash., No. 322A, p. 100); "Pawnee Loup Branch of Platte
River, Middle Loup, probably Hooker Co." {fide Lugn and Schultz, 1934,
Neb. State Mus., Bull. 41, I, p. 372). Type fig.: Leidy, 1869, Journ. Acad.
Nat. Sci. Phila., (2), VII, PI. xxv, figs. 1, 2.

Cf. Rhabilobunu.H mirificus Hay, 1914, below, this list.

1858 Elephas imperator Leidy Ibid., pp. 10, 29. Type loc: Loup Fork of Platte River, Nebraska Archidiskodon impera-
(Leidy, 1869, Journ. Acad. Nat. Sci. Phila., (2), VII, p. 2.54; and Hay, 1914, tor
Iowa (ieol. Surv., Ann. Rept. for 1912, XXIII, pp. 421, 422), po.ssibly Seneca,
Thomas County, Nebraska (see Hay, 1924, Publ. Carnegie Inst. Wash., No.
322A, p. 100); "Pawnee Loup Branch of Platte River = Middle Loup, proba-
l)ly Hooker Co. [Nebraska]" (fide Lugn and Schultz, 1934, Neb. State Mus.,
liull. 41, I, p. 373). Typo fig.: Leidy, 1869, Journ. Acad. Nat. Sci. Phila.,
(2), VII, PI. xxv, fig. 3. Osborn's neotype: 1922, Amer. Mus. Novitates,
No. 41, p. 4, fig. 4, from Guadalajara, Mexico.

1859-62 Elephas texianus Owen Hept. Brit, .\ssoc. Adv. Sci., 28th meeting, j). Ixxxvi (name); and Blake, Parelephas columbi
1861, Geologist, IV, p. 470 (name). Type loc San Fc'lipe do Austin, Brazos
:

River, Texas. Type description: Blake, 1862, Geologist, V, p. 58. Type

tig.: Blake, 1862, ibid., PI. iv.
 NOMENCLATURE OF THE PROBOSCIDEA 1395

Reference in
Year Name Author Present Memoir

1861 Dinutheiiiim leviu.s Joyivd-dii Compt. Rend. Acud. .Sci., LIU, p. lUU. Typeloc: tihve Saint- Dcinothcrium leviua
Alban, Iserc, France.

1861 Ekphas intermedins 3omda.n Ibid., p. 1013. Type loc: Near Lyons, Rhone Valley, France. Parelephas intermedius
No figure recorded, but see Lortet and Chantre, 1872 [1876], Arch. Mus. hist,
nat. Lyon, I, frontispiece, plate of referred skeleton (Fig. 944 of present
Memoir), also referred molars in the Museum dc Ville, Lyons (Fig. 943 of
present Memoir).
Very similar to Elephas Irogontherii Pohlig (fide Deperet letter, August 26, 1921).

1862-1868 Elephas Melilensis Falconer "The Parthenon," p. 780. See Falconer, 1868, "Palseont. Mem., Palxoloxodon meliten-
II, pp. 292, 299, 307, 308. Type loc: Zebbug Cave, Malta. Type fig.: sis
Falconer, 1868, op. cit., PI. xi, figs. 1, la.

1867 Elephas fakoneri Buiik Trans. Zool. See. London, VI, Pt. V, p. 251. Typeloc: Zebbug Cave, Palseoloxodon falconeri
Malta. Type fig.: Busk, ibid., Pis. xlix, l, li (not figured in present
Memoir).
See Falconer, 1868, II, p. 292, footnote, where it is stated by the editor that this and
other Maltese specimens had been identified by Falconer as early as July, 1860.

1867 Mastodon virgatidens von Meyer Palaeontogr., XVII, p. 61. Type loc: Near Fulda, northeast Turicius virgatidens
of Frankfort, Germany. Type fig.: von Meyer, ibid., Taf. iv, figs. 1-5.

1868 Mastodon Andaranus iWconer "Palseont. Mem.," I, p. 124, footnote. Type loc: Deccan, Triloplwdon pandionis
India.
See Vol. 1, 11. 267, of the present Memoir.

1868 Dinotherium Perimense Falconer Op. cit., I, p. 415 (name only). Type loc. : Perim Island, India. Deinotherium indicum

1868 Dinotherium Pentapotamicum Falconer Op. cit., II, p. 5.

See Dinotherium pentapotdmise Lydekker, 1876, below.

1868 Elephas minutus Nesti (In Falconer, op. cit., II, pp. 104, 105).
See note under Elephas minimus Nesti, 1846, above, this list.

1868 Trilophodon Ohioticus Falconer Op. cit., II, pp. 176 (footnote), 204. Hab.: North America. Mastodon americanus

1868 Mastodon amcricanus\je\AY Proc Acad. Nat. Sci. Phila., XX, p. 175. Hab.: North America. Mastodon americanus
Sec Leidy, 1869, .Journ. Acad. Nat. Sci. Phila., (2), VII, pp. 240, 395.
Lt^idy first used the term Mastodon americanus in the 1868 article, as the equivalent of
M. ohioticus and M. giganleus of authors.

1869 Mastodon obscurus Leidy Journ. Acad. Nat. Sci. Phila., (2), VII, pp. 244, 396. Typeloc: Near Trilophodon obscur us
Greensburgh (Greensboro), CaroUne County, Maryland. Type fig.: Leidy,
ibid., PI. xxvii, fig. 13.
Merrill (1907, Bull. U. S. Nat. Mus., No. 53, p. 45) regarded a third superior molar,
r.M', from Tarboro, North Carolina, as the type of Mastodon obscurus. The present author
refers it to Ocalientinus (Serridentinus) obliquidens of this Memoir (see Vol. I, p. 286).

1870 Stegodon sinensis Owen Quart. .lourn. Geol. Soc London, XXVI, Pt. I, p. 417. Type loc: Steg ado n sinensis
Alleged to be from marly beds vicinity of Shanghai, China. Type fig.Owen,
:

ibid., PI. xxvii, figs. 1-3.

1870 Stegodon orientalis Ov/en Ibid., p. i2l. Typeloc: (?)Cave near city of Clumgkingfoo, Province Stegodon orientalis
of Szechuan, China. Type fig.: Owen, ibid., PI. xxviii, figs. 1-4.

1870 Elephas mnaidrs; Adams "Notes of a Naturalist in Nile Valley and Malta," p. 223. Type loc: Palxoloxodon mnaidri-
Mnaidra Gap, Malta. Type fig. Adams, op. cit., PI. ii, figs. 2, 2a. Supple-
: ensis
mentary description as Elephas mnaidricnsis: Adams, 1874, Trans. Zool.
Soc. London, IX, Pt. I, p. 116; with figures of the type, PI. vii, figs. 2, 2a; of
the paratype, PI. vii, fig. 1.
139(j OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir
1870 Ekphas Curmiliae Aradas Atti Accad. Giooiiia .sci. nat. Catania, (3), IV, p. 235. Type loc: Lo.rodonki rornalise
Catania, near the monastery of Santa Chiara, Sicily. Type figs. Aradas, :

ibid., figs. 1, 2.

1871 Madodun shepardi heidy Proc. Acad. Nat. Sci. Phila., XXII, p. 98. Type loc: Dry Creek, Khynehotherium
Stanislaus County, California. Type fig.: Leidy, 1873, Rept. U. S. Geol. shepardi
Surv. Terr., I, PI. xxi, figs. 3, 4.

1872 Elephas Indianapolis Foster Proc. Amer. Assoc. Adv. Sci., August, 1872, p. 259 (name only).
Same specimen from Indiana for which the name Elephas Mississippiensis was subse-
quently proposed. See next item.

1872 Elephas Mississippiensis Foster Nature, VI, p. 443. Hab. : Indiana, exact locality unrecorded. Parelephas{t) missis-
Name changed from Elephas Indianapolis (see preceding item), without description or Stpptensisy!)
figure.

1873 Mastodon proavus Cope New Vertebrata from Tertiary of Colorado," p. 10. Type
"Synopsis of Serridentinus proavus
loc: Pawnee Buttes, Pawnee Creek, Weld County, Colorado. Type fig.:
Cope, 1889, Amer. Naturalist, XXIII, No. 268, p. 202, fig. 6, as Tetrabelodon
angustidens proavus.

1875 Mastodon prod itdus Cope Proc Acad. Nat. Sci. Phila., XXVI, pp. 221, 222. Type loc: Santa Serridentinus prodxwtus
Fe marls. New Mexico. Type figs. Cope, 1877, Kept. U. S. Geogr. Surv.
:

West of 100th Meridian (Wheeler), IV, Pt. II, Pis. lxx, figs. 1-3, lxxi, fig. 3.
Ncotype fig. Frick, 1926, Bull. Amer. Mus. Nat. Hist., LVI, Art. II, fig. 7A.
:

1875-1923 Elephas Aitsonius Major (MS. labels). Name in Verri, 1886, Boll. Soc geol. Ital., V, p. 453. Uesperoloxodon antiqu-
Type loc: San Romano, Val d'Arno inf., Italy. Type fig.: Dep6ret and us ausonius
Mayet, 1923, "Les Elephants Pliocenes," PI. x, figs. 1, 2.
See Weithofer, 1890, p. 194, footnote, and p. 206; also Depcret and Mayet, 1923, p.
162.

1876 Dinoiheriunt pcntapotamisp Lydekker Mem. (Jcol. Surv. India, Palaeont. Indica, (X), I, Pt. II, Deinotheriuin indicum
pp. 72, 73. (Falc Ex MS., fide Lydekker, 1885, "Catalogue of the Remains
of Siwalik Vertebrata. Cieological Department of the Indian Museum,
. .

Calcutta," p. 104.) Type loc: Near Attock, Indus valley, India. Type
fig.: Lydekker, 1876, ibid., pi. ix, figs. 1, 2.

Lydekker, 1876, i)p. 72, 73: "Subsoquontly, the same naturalist [Falconer] identified
two other molar tc^rth from near Attock ('Pal. Mem.,' vol. I, p. 414) as belonging to the
same gciuis [Deinothirium]; in the note on these specimen.s in tlie 'Palseontological Memoirs'
no .specific name was assigned to them, though they were considered to be of too small a size
to have belonged to 1). {prrimense) indicmn; these specimens are now in the Indian
Museum; they are ticketed with labels in Falconer's handwriting bearing the name of D.
pnitapolamiw, which name I have accordingly adoi)ted. The specimen is distinguished by
. .

its niiich smaller size from any of the Eurojican species; from the first premolar of D.

giganteum and />. cuvieri, it is distinguished by the following points: the antero-posterior
valley is very much deeper and wider in the Indian form, rendering thereby the inner
tubercles more comiiletely conical; the posterior tubercle is mammilliform in the Indian
form, whereas it is elongated transversely in the European form."
Sec Falconer, 1868, II, p. .5, footnote by Editor, in which he states that "A specimen of
the third lower premolar of this species, from the 'Red Marl' at Noorpoor, found in Dr.
"
Falconer's collection, is labelled in his hand-writing, Dinotheriurn Fentapotamicum, Falc'
'

187 Mastodon {Trilophodon) Fnlc.oneri Lydekker Hec Geol. Surv. India, X, Pt. II, p. 83. Type loc: Tetralophodon (Lydek-
Potwar district, Punjab, India. Sujjplementary d(wcription: Lydekker, keria) falconeri
1880, Mem. Geol. Surv. India, Palsont. Indi(;a, (X), 1, Pt. \, p. 206. Type
fig.: Lydekker, 1880, ibid., PI. xxxiii, figs. 1, 4.

1877 C.\Cxnobasileus\lremontigerus Cope Proc. Amer. Phil. Soc, X\l, pp. 584, 585. Type loc:
Probably Texas. No figure recorded.
Genus and species withdrawn by Cope in 1889 (1889.2, p. 207).
 NOMENCLATURE OF THE PROBOSCIDEA 1397

Reference in
Year Name Author Present Memoir
1878 Tetralophodon campeMer Cope. Proc. Amer. Phil. Soc, XVII, p. 225. Type loc. Republican : Tetralophodon campes-
River beds, Sappa Creek, Rawlins County, Kansas. Type figs.: Cope, 1889, ter
Amer. Naturalist, XXIII, Pis. ix, x; also Cope and Matthew, 1915, "Hither-
to Unpublished Plates of Tertiary Mammalia and Permian Vertebrata," Pis.
cxx, cxxi, CXXII, CXXIII.

1879 Mastodon affinia Jourdan (Ex MS. 1859, labels in Lyons Museum), in Lortet and Chantre, Arch, Zygolophodon borsoni
Mus. hist. nat. Lyon, II, p. 308. Type loc: (?) affinis

1879 Elephas primigenius comune Is.sel Ann. Mus. Civ. Storia Nat. Genova, (1), XIV, p. 153 (as Mammonteus primi-
1913 Elephas primigenius). In Zuffardi, 1913, Palseont. ital., XIX, p. 136. Type genius
loc: Camporosso, near Ventimiglia, Italy. Type figs.: Issel, ibid., figs. 1-4.

1879 Elephas atlanticus Pomc'l Bull. Soc geol. France, (3), VII, p. 51. Type loc: Ternifine, near Palieoloxodo u atlaii li-
Mascara, Algeria. Cotype fig.: Pomel, 1895, "Palfentologie Monographies, cus
No. 6. Les Elephants Quaternaires," Carte Geol. L'Alg^rie, PI. viii, figs. 1, 2.

1880 Diitotherium sindiense Lydekker JNIem. Geol. Surv. India, Palaeont. Indica, (X), I, Pt. V, p. 196. Deinotherinm sindi-
Type loc : Sind, India. Type fig. Lydekker, ibid., PI. xxxi, fig. 4.
: ense{1)
See Lydekker, 1879, Rec. Geol. Surv. India, XII, Pt. 1, p. 43, for first notice, but with-
out name.

1880 Elephas (Euelephas) antiquus var. nana Acconci Atti Soc. Toscana Sci. Nat., V, Fasc. I, pp. 146- Hesperolox.odon antiqu-
150. Type loc: Cavern near Monti Pisani, Cucigliana, Tuscany, Italy. us nanus
Type fig.: Acconci, ihid., Tav. iv, figs. 6, 7.

1882 Notelephas aiislralis Owen Proc. Roy. Soc. London, XXXIII, p. 448; 1883, Phil. Trans. Roy. Soc. Diprotodon australis
London, CLXXIII, Pt. Ill, p. 777. Type loc: Drift deposit of ravine in
district of Darling Downs, about 60 miles from Moreton Bay, Queensland,
Australia. Type fig.: Owen, 1883, /bzrf., PI. li.
.Jack and Etheridge, 1892, p. 683: "Notoelephas [Notelephas of Owen, 1882] auslralis
Owen . and species were founded on portions of a tusk indicating a mammal
.This genus
larger than Diprotodon, in fact the largest fossil mammal yet foreshadowed amongst the
extinct Australian forms. Sir Richard Owen appears to regard it as a Proboscidian
Placental. . . He does not appear to apprehend any connection between this tusk and the
molar tooth [.Mastodon australis Owen, 1844] formerly described by him, and forming
a portion of the late Count P. E. de Strzelecki's Collection. Prof. Owen remarks that this
molar is too large to be associated with the tusk, supposing the latter to have come from the
upper jaw of a full-grown individual of its si)ecies. Grave doubts have been expressed by
. .

several Writers as to the probability of this tooth as an Australian fossil, and it would
perhaps be better to expimge it from the list."

1883 Mastodon Zaddachi Jeiitzsch Schrift. phys.-okonom. Ges. Konig.sb., Jahrg. XXIII, Abth. 2, p. Zygolophodon borsoni
202. Type loc: Thorn, West Pioissia. Type fig.: Jentzsch, ibid., Taf. v, zaddachi
figs. 6a, 66.

1883 Leptodon minor Gunn Geol. Mag., Dec. II, N.

S., X, p. 458. Type loc: Forest bed, Norfolk, Hesperoloxodo n
England. Type fig. : New
Edition of Gunn's "Sketch of Geology of Norfolk, antiquus
PI. I, letter I" (fide Gunn, 1883, loc. cit.). This edition does not seem to have
been published (cf. "Memorials of John Gunn," 1891, p. v).

1883 Leptodon giganteus Gunn Loc. cit. Type loc; Forest bed, Mundesley, Norfolk, England. De- Hesperolo.rodon
scription and type figure: Gunn, 1891, "Memorials of John Gunn," PI. iv, antiquus
fig. 2.

Same as E. Gunnii Lartet, 1883, and Elephas giganteus intermedins Gunn, 1891 (see
below, this list).

1883 E. Gunnii Lartet (InGunn, 1883, loc. cit.) bed, Mundesley, Norfolk. Type
Type loc: Forest Hesperoloxodon
ramus No. 361, Gunn Norwich Castle Mus. 1703. Description and
Coll., antiquus
type fig. : Ciunn, 1891, "Memorials of John Gunn," PI. iv, fig. 2.
It will be ob.served in this quaint description of 1883 that Gunn designates as the type
of E. Gunnii Lartet the very same specimen and figure selected as the tyi>e of Leptodon
giganteus. A revision of these types and specific names applied to the collection of Gunn
may be found in J'. Leney's "Type-specimens in the Norwich Castle Museum," 1902, Geol.
Mag., p. 169.
1398 OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir
1883 Elcphas lAuiKinnorae Major Kosmos, XIII, .lalirg. \ll, ]). 6. TyiK' loc. : Moriinctitu h. ( loniiesa, F<il;iolo.rodiiii lainur-
Sardinia. No figures published. morae

1884 Elephan afriraniis foxnilis Tliomas, Philippe \l6m. Soc. geol. France, (3j, III, No. 2, p. 4(5. Palu'oloxudoii alianli-
Cited by Trouessart, 1897-1899, p. 708, as found in Algeria. cus

1884 Mastodon (Tn'lophodon) angustidens C'uv. var. palxindicus Lydckker Mem. Geol. Surv. India, Trilophodon palxindi-
Palaeont. Indica, (X), III, Pt. I, p. 19. Type loc.: Near Kamlial, northern cus
Punjab, India. Type fig. Lydekker, ?6?V/., Pi. iv, fig. 3.
:

1884 M. [Mastodon] euhypodoti Cope Anier. Naturalist, XVIII, p. 525. Type loc: Trail Canon, south Bllrkotheriu m euhypo-
fork of Driftwood Creek, Hitchcock County, Nebraska. Type figs. Cope, : doti
1889, ibid., XXIII, p. 203, fig. 7, and PI. xiii (erroneous drawing) figured as
Tetrabclodon euhypodon.

1884 M. [Mastodon] serridens Cope Loc. n't. Tyi^e loc. : Texas, probably (
"iarcndon formation. Type Serridentinus serridens
fig. : Cope, 1889, ibid., p. 205, fig. 8, figured as Tetrabclodon serride7is.

1884 Dibelodon tropicus Cope Proc. Amer. Phil. Soc, XXII, p. 7. Type loc: State of Michoacan, Cordillerion tropicus
\'alley of Mexico. Type fig.: von Meyer, 1867, Palseontogr., XVII, PI. vi,
as M. humboldtii. Ulide Coil.
.species {Uibelodon tropicus Cope) by FreiuU^ibcrf;, 1922, pp.
Sec diseus.sion of this
128, 129, in which ho reproduces the original type figure of Cope from von Meyer and places
it at the summit of an ascending scries ending with Mastodon oligohimis progrrssa l'^reud(Mi-
berg.

1885 Mastodon perimensis var. sinensis Koken Geol. u. Pal. Abh., Ill, Heft II, p. 34. Type loc: Tetralophodon (Lydek-
Yunnan, China. Type fig. : Koken, ibid., PI. xii, fig. 1. keria) sinensis

1885 Elephas trogontherii Polilig Zeitschr. deutsch. geol. Ges., XXXVII, Heft IV, p. 1027. Parelephas trogontherii
1887 Elephas (primigenius bezw. meridionalis) trogontherii Pohlig, 1887, Sitz.-Ber.
niederrhein. Ges. Bonn, p. 274.
Elephas (primigenius) trogontherii Pohlig, 1887 [1888?], Zeitsclir. deutsch.
geol. CJes., XXXIX, Heft IV, p. 799.
Tvi)e loc: Sii.s.senborn, near Weimar, northern Germany. Type figs.:
Pohlig, 1888, Nova Acta Leop. Carol., LIII, p. 193, fig. 79, and p. 195, fig. 82.
Cf. Elephas intermedius Jourdan, 1861, above, this list.

1886 Mdsliidon coulUyi Lydckker Mem. Geol. Surv. India, Palaeont. Indica, (X), III, i)p. xiv-xix. Ste.golophodon <« utleyi
Type loc: Pcrim Island, India. Lectotype fig.: Lydckker, ibid., p. xv, fig.
6. Cotype figs.: Lydckker, 1880, ibid., (X), I, Pt. V, PI. xl [.subsequently
referred by the; present author to Tetrolophodon punjobiensi.s]; 1884, ibid.,
(X), III, Pt. V, PI. XVI, fig. 2; 1886, ibid., (X), HI, p. xv, fig. 5; Falconer and
Cautley, 1847, "Fauna Anticjua Sivalcn.sis," PI.XL, figs. 2, 2a, 3,3«.

1886 MatitodoK piinj(ibien.'^i.'< Lydckker "Cat. Foss. Manim. Brit. Mus.," p. 60. Tyjjc loc: Siwalik Tetralophodon piuijab-
Hills, Punjab, India. Lectotype fig.: Lydckker, 1880, Mem. Geol. Surv. iensis
India, Palaeont. Indica, (X), 1, Pt. V, PI. xlii.
Originally referred to Mastodon (Tetralophodon) perimensis.

1886 Mastodon (Trilophodon) floridanus Leidy Proc Acad. Nat. Sci. Phila., XXXVIII, p. 12. Type Ocalienti.nus (Serriden-
1896 loc: Alachua clays, Mix.son's bone bed, near Williston, Levy County, timis) floridanus
Florida. Type fig.: Leidy and Lucas, 1896, Trans. Wagner Free Inst. Sci.,
IV, PI. VI, figs. 1, 2.

1887 E. [Elephas] antir/uus var. minor Pohlig ^'erh. natur. N'ereins ])reuss. Uhein., Jahrg. 44, p. 115.
Type loc : Seville, Spain.

1887 Stegodon trigonocephalus Martin Sammiuiig. ( Ic^log. Reichs-Museums, Leiden, 1'"= Scrit;, Bcitriige Stegodon Irigonorcphu-
z. Geolog. O.st-Asiens u. Australicns, IV, Heft II, pp. 27, 36. Type loc: lus
Probably vicinity of Surakarta, Java. Type figs.: Martin, ibid., Tab. ii, figs.
1, la, and Tab. iii, fig. 1.
 NOMENCLATURE OF THE PROBOSCIDEA 1399

Reference in
Year Name Author Present Memoir
1887 [1888?] Elephas (primigenius) irogontherii Pohlig See Elephas trogontherii Pohlig, 1885, 1887, above,
this list.

1888 E. [Elephas] (primigenius) Leith-Adamsi Pohlig Nova Acta Leop. Carol., LIII, No. 1, pp. 229, Mammonteus{1) primi-
232. Type loc: Dornap (Fuhlrott), Thuringia, Germany. Type fig.: genius leith-adamsi
Pohlig, ibid., p. 229, fig. 101 c-d.

1888 Mastodon bonaerensis Moreno "Informe Prelim, progresos Mus. La Plata," pp. 17, 18 (nomen Cuvieroniusbonaeren-
nudum). See Ameghino, 1889, Acta Acad. Nac. Cien, Cordoba, VI, p. 641. sis { = superbus)
Type loc. Arrecifes, Province of Buenos Aires, Argentina.
:

Not figured by Moreno, but Eduardo Carette figured it in his article "Los Proboscideos
F6siles Argentinos," 1919, Ldm. xi, fig. 2, as D. \Dihehdon\ platensvi.

1888 Mastodon aj-geritinus Ameghino "Rapidas Diagnosis," p. 7. Type loc: Valley of Tarij a River, Notiomastodon argen-
northern part of Argentina, exact locality not recorded. Type fig. Ameghino, : iinus
1889, Actas Acad. Nac. Cien. Cordoba, VI, p. 644.

1888 Mastodon platensis Ameghino Loc. cil. Type loc: San Nicolas de los Arroyos, Province of Cuvieronius platensis
Buenos Aires, Argentina. Type fig.: Ameghino, 1889, ibid., p. 641.

1888 Mastodon superbus Ameghino Loc. cil. Type loc: Pergamino, Province of Buenos Aires, Cuvieronius superbus
Argentina. Type fig.: Ameghino, 1889, ibid., p. 647.

1889 Mastodon rectus Ameghino Actas Acad. Nac Cien. Cordoba, VI, p. 643. Type loc: From the Cuvieronius rectus
Ensenada, near La Plata, Argentina. Type fig. Ameghino, ibid., p. 643. :

1889 Elephas (primigenius) Leith-Adamsi Pohlig var. minor (In Trouessart, 1897-1899, "Cat. Mamm.
Viv. Foss.,"p. 711).
Search in Pohhg's contributions of 1889 does not disclose a definition of this variety,
and a letter from Doctor Pohlig (Sept. 10, 1924) does not authenticate var. minor as a sub-
specific term (cf. p. 1150 of the present Memoir).

1889 Tetrabelodon brevidens Cope Amer. Naturalist, XXIII, pp. 198-202. Type loc: Smith River, Rhynch other ium
Meagher County, Montana. Type fig. Cope, jfc/c?., p. 201, fig. 5. : brevidens

1889 E. [Elephas] primigenius americanus Cope Ibid., ytp. 207, 209. Hab. : North America.

1889 Elephas primigenius columbi Cope Ibid., pp. 208, 209. Type loc : "Orange sand," city of Dallas,
northeastern Texas. Type figs. Cope, i6?c/., : PI.xiv and text fig. 9.

1889 Elephas lyrodon Weithofer Jahrb. Geol. Reichsanst., XXXIX, pp. 79, 80. Type loc.: Val Archidiskodon meri-
d'Arno superieur, Italy. Type figs.: Weithofer, 1890, Beitr. Pal. Osterr.- dionalis (female)
Ung., VIII, Taf. ni, fig. 2, Taf. iv, fig. 2, Taf. v, fig. 1.

Compare observations of Pohlig (1891, pp. 314, 334).

1890 Slegodon Mindanensis Naumann Zeitschr. deutsch. geol. Ges., XLII, Heft I, pp. 166, 167. Type Stegodon (Archidisko-
loc: Mindanao, Philippine Islands. Type fig. Naumann, 1887, Abh. Ber. : don'?) mindanensis
k. Zool.-Anthrop.-Ethnog. Mus. Dresden, No. 6, Taf. I, figs. 1 and 2, as S.
trigonocephalus.

1890 Stegodon Airdwana Martin Verb. Kon. Akad. Wetensch. Afdeel. Natuurk., Amsterdam, XXVIII, Stegodon airdwana
p. 4. Type loc: Ala.s-Tu\va, Trinil, Java. Type figs.: Martin, ibid., Tab. i,
figs. 1 and 2, Tab. ii, figs. 3 and 4.

1890 Mastodon rugosidetis Lcidy Proc Acad. Nat. Sci. Phila., XLII, p. 184. Type loc: Santee beds, Mastodon americanus
Beaufort County, South Carolina. No type figure recorded (see Fig. 115 of rugosidens
present Memoir drawn from cast, Amer. Mus. 14445). Original in Academy
of Natural Sciences, Philadelphia.

1891 E. [Elephas] primigenius Blum. var. hydruntinus Botti Boll. Soc geol. Ital., IX, p. 709. Type Mammonteus primi-
loc: La Grotta Ossifera di Cardamone, Terra d'Otranto, Italy. Type fig.: genius hydruntinus
Botti, ibid., Tav.fxxvi, figs. 1, la, 2, 2a.
1400 OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir
1891 E. [Klephas] giganteiis intertnedius Gwnn Heyperolosodon an-
Samc as Leplodun giganlcus Uuiiii, 1SS3, subsequently named by Lartet (in Gunn, llf/uus

1883) E. gunnii (sec above, this list).

1891 E. [Elephas] Neslii Pohlig Nova Acta Acad. Loop. Carol., LMI, pp. 303, 304. Elephas anliquun Farelephas{'?) trogon-
Neslii II. f., Pohlig, ibid., p. 465. Type loo.: Happi.sborough, Forest bed thern neslii
(Norfolk), Walton (E.isex), Soiithwold (Suffolk), England. C'otypc figs.:
None. Sec Fig.s. 941, 1041 of present Memoir.

1891 Miislodon inmtcriaims Ameghiiio Rovista Argent. Hist. Nat., I, p. 243. Type loc: Puerto Cuvieronius {l)maderi-
Madero in lUieno.s Aires, Argentina. anus

1892 D. [Dinotherium] gigantissimumG. Stefanciicu Bull. Geol. Soc. Amcr., Ill, pp. 81, 82. Type Deinotherium gigantis-
loc: Gaieeana, Tecuci 11, Rumania. simum
See Slefancscii, 1878, pp. 101 404, giving an account of discovery, also 189.3, p. 173,
Tab. IV and v (referred molars).

1892 Mdxlodiiii .<urcf'HM>r Cope Proc. Acad. Nat. Sci. Phila., XLIV, pp. 227, 228. Type loc: Blanco Stegomastodon successor
beds, Texas. Type figs.: ("ope, 1893, 4th Ann. Rept. Geol. Surv. Texas, PI.
XVI, figs. 1, la, PI. XVII, fig. 1, as Dibelodon tropicus.

1892 D. \l)ihrl()(l<}ii\ prariirsor Cope Anicr. Naturalist, XXVI, p. 1059 (name only). Tyjjc loc: Mt. Serbelodon('!) prxcursor
1893 RUinco, Llano Estacado, Texas. Type description: C ope, 1893, 4th .\nn.
Rept. Geol. Surv. Texas, pj). 64, 65. Tj'pc figs.: Cope, 1893, op. cit., Pis.
XVIII, XIX.

1893 T. [Telrabelodon] >ierride.ns ciinarronis Cope 4th Ann. Rept. Geol. Surv. Texas, pp. 18-20. Type Serridenlinus serridens
loc. Ea.st of Llano Estacado, Texas, north of south fork of Red River.
: Type rimnrronix
fig. Cope, op. rit., PI. Ill, figs. 2 and 3.
:

1893 Mu.'itodori uligobiini.s Cope Op. cit.. ]). .59. Type loc: Tequixquiac, Valley of Mexico. Type Cordillerion oligobunis
fig. : Afti^r Villada, 1903, Ann. Mus. Nac Mexico, VII, Lam. vii (as Dibelodon
Shepardi, Cope).

1893 .1/. |.U'i.v/(«/()/i| r/)/7c//.s/.s Philippi Zeit.schr. deutsch. geol. (!es.,XLV, p. 88. Type loc: Near Cuvieroniu.'i rhilensis
Lake Tagua-Tagua, Chile. Type fig. : Philippi, ibid., p. 89, fig. 4.

1893 .U(;.s7(«/()/( /)<(//i7V/;;i(,v Philii)pi Ibid., p. 89. Tvjie loc: riloni.-i, Bolivia. Type fig. : Philipi>i, Cordillerion bolivianus
ibid., p. 89, fig. 2.

1894 .Md.slodon Sain ndi l':t\lo\v (MS. labels Ilofniuseuni in Wicn), Mem. .\cad. Imp. Sci. St. Petersb., Tctralopliudon sp.
(8), I, No. 3, p. 16. Type loc. : Maragha, Persia.
See Kittl, 1887, ]). 320, for associated fauna of Maragha.

1895 ElcplKL-i jiilcnsis Pomel Paleontologie Monographies, No. 6, Carte (ieol. L'.\lgerie, |)p. 32, 39. Palxoloiodon jolensis
Hab.: Algerian .seacoast, downward from Krober-Roumia. Type fig.:
Pom(!l, op. cit., PI. V, figs. 3 and 4.

1896 Elcplm.s niendionali'i antiquitati.^ Portis "( 'ontribuzioni alia Storia fisica del Bacino di Roma, .
."

1913 Torino, II, p. 272.

Portis' original description of 189{) was not seen by the jjicsent author. In the
Rivista Italiana di Palcontologia, II, of the same year, the name ajJiiears in a phylogenetic
chart on page 331. Zuffardi in PaliPontologica Italica, 1913, XIX, p. \5!i, includes this

subspecies in the synonymy of Klcphas antiquum Falc. var. trogonlhmoidfs.

1897 .Ma.slodon angustidens Ciiv. inut. a,sc. pygmseus Depdret Bull. Soc g^ol. France, (3), XXV, p. 519. I'hiomia pygnmus
Type loc: Kabylic, near Isscrvillc, Algeria. TyP<" fiR- Dcperct, ibid., PI.
XIX, figs. 1 -3.

1897 Elephas plalyrhynchus Graells Mem. Heal. .\cad. Cien. Exactas, Fis. nat., Madrid, XVII, p. .569. Jfe.sjieralo.nidan antiqn-
Typc loc: San Isidro del Campo, near Madrid, Spain. Type fig.: (!raells, an platyrhynchu.s
ibid.. Lam. xviii, figs. 9a, 10.
 NOMENCLATURE OF THE PROROSCIDEA 1401

Reference in
Year Name Author Present Memoir
1S97 Mndodofi ai(relie/if:f, (\\v. (In (Iraclls, ?'6/r/., p. 573.) Hah.: Marliid. [Not determined by the
j)resent author]

1899 Mastodon angustidens var. latidens Lankester Geol. Mag., Doo. IV, N. S., VI, pp. 289-292, PI. xi. Zygolophodnn .sp.

Typo loc. : Suffolk Crag, England.

Figured as a, trilophodont Mastodon in Quart. Journ. Geol. Soc., 1870, PI. 34, figs. 1-4.

1900 Elephan ma.rimns sKinatraiiti-'i Lydckker "Great and Small Game of India . . . ," p. 11. Living Elephas iiidiciin
form. sumatranus
See Elephas sumatranus Temminck, 1847, above, this list ; also Lydekker, 1916, p. 83.

1900 Ehphns ryclotis Mat^chie Sitz.-Ber. Ge-f. naturf. Freunde

Berlin, No. 8, p. 194. Ty])e loc.: Lo.rodonln rnpensis
Mwelle district, S.Tyj^e fig.: Heck, 1899, "Leheude Rilder
Cameroon. rychtlis
aus dem {{ciche dei' Ticre, Berlin," PI. cxlvi (original not seen by the present
author). l"\)rinerly in Berlin Zoological Garden. Li\ing form.

1900 Elephas (Loj-ndoida) oj-yotifi M-Mschu' Ibid., [). 196. Typeloe.: Upper Albarti, Sudan. Living Louodoidd (tfn'rana
form. oxyotis

1900 Elephas (Lo.vodonta) kiioehenhaueri Matschic Ibid., p. 197. Ty]ieloc.: Barikiwa, Tanganyika Loxodonia africaiia
Territory. Living form. knorhenhoueri

1901 Pnlspomastodon Beadnelli Audrcwf^ Zoologist, (4), V, August 15, pp. 318, 319 (name only); Paheoniastodon
Tagehlatt \ Interuat. Zool.-Congr., Heiiin, No. 6, August 16, ]). 4; (ieol. beadnelli
Mag., Dec. IV., N. S., VIII, September, p. 401. Type loc: P'ayum, Egypt.
Type fig.: Andrews, 1901, Geol. .Mag., Dec. IV, N. S., VIII, p. 401. fig. 1,
A, B.

1901 Mwritherium L(/ONSi' Andrews Tageblatt V

Internat. Zool.-Congr., Berlin, No. 6, August 16, p. Moeritheritimlijonsi
4; Geol. Mag., Dec. IV, N. S., VIII, September, pp. 403, 404. Type loc.:
Fayiim, Egypt. Type fig.: Andrews, 1901, Geol. Mag., Dec. IV, N. S.,
VIII,p.404, fig. 2.

1902 Phiomin .serridens ,\ndrews and Beadnell "A Preliminary Note on Some New Mammals from the Phiomia serridens
Upper Eocene of Egypt," p. 3. Type loc: Fayiim, Egypt. Type figs.:
Andrews and Beadnell, loc. cit., figs. 1 and 2.

1902 Maeritheiitim gracile Andrews Cieol. Mag., Dec. IV, N. S., IX, p. 292. Type loc: Fayum, Mcsrilheriion gracile
Egypt. Type figs. Andrews, 1906, "A Descriptive Catalogue of the Tertiary
:

Vertebrata of the Fayum, Egypt," p. 127, and PI. xvii, figs. 1 and 2.

1902 Mastodon tarijensis Ameghino An. Mas. Nac. Buenos Aires, VIII (Ser. 3', I), p. 2 (name only). Cordillerion tari.jensis
Hab. Valley of Tarija, Bolivia.
:

1903 Mastodon Lydekker i ScMosi^cr Abh. bayer. Akad., II CI., XXII, Abth. 1, pp. 46, 47. Typeloe: Semdentinuslydekkeri
North China. Type fig. Schlo.sser, ibid., Taf. xiv, : fig. 8.

1903 Elephas Cypriotes B&te Soc. London, LXXI, pp. 498-500. Typeloe: Cave, Kerynia
Proc Roy. Palseoloxodon Cypriotes
Cyprus. Cotype figs.: Bate, 1904, Phil. Trans. Roy. Soc. London,
Hills,
CXCVII, B, Pis. XXI, xxii.

1904 Mwritherium trigodon Andrews Geol. Mag., Dec V, N. S., I, p. 112. Typeloe: Fayiim, Egypt. Mirritherium trigodon
Type Andrews, 1906, "A Descriptive Catalogue of the Tertiary Verte-
fig.:
brata of the Fayiim, Egypt," PI. ix, fig. 5. Supplementary description:
Andrews, 1906, op. cit., p. 128, name changed to trigonodon.

1904 Palxomastodon minor Andrews Ibid., p. Wo. Type loc. Fayiim, Egypt. Type figs. Andrews,
: : Phiomia minor
1906, "A Descriptive Catalogue of the Tertiary Vertebrata of the Fayum,
Egypt," PI. XIV, figs. 1, 1a, text fig. 50D. Supplementary description:
Andrews, 1906, op. cit., p. 168.

1905 Pnlxomastodon minus Andrews Ibid., II, p. 562. Phiomia minor

Error, the author liad reference to Paheoniastodon [
= Phiomia] mirwr.

1905 Pal^omastodon panni s Andrews Loc. cit. Type loc: Fayum, Egypt. Type figs. Andrews, : Palseomastodon parvus
1906, "A Descriptive Catalogue of the Tertiary Vertebrata of the Fayum,
Egypt," p. 143, text fig. 50C, and p. 163, text fig. 55.
1402 OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir
1905 Palieom(mtodon winioni Andrcwi^ Ibid., p. 56^. Typeloc: Fayum, Egypt. Type fig. Andrews, : Phiomiawintoni
1906, "A Descriptive Catalogue of the Tertiary Vertebrata of the FayOm,
Egypt," p. 157, text fig. 53.
Originally referred to Palseomasiodon; transferred by Matsumoto (1922, p. 3) to
Phiomia.

1906 Mceritherium trigonodon Andrews Op. cit., p. 128.

See Motritherium trigodon Andrews, 1904, above.

1906 Elephas africamis albertensis Lydekker Field (London), CVII, p. 1089. Type loc. South end of : Loxodonta africana
Lake Albert, Africa. Type fig.: Lydekker, 1907, Proc. Zool. Soc. London, alberlensis
text fig. 121. Living form.

1906 Elephas africanus puTnilio 'Noa.ck Zool. Anz., XXIX, pp. 631-633. Typeloc.: French Congo. Loxodonta africana
Type fig.: Hornaday, 1905, Bull. New York Zool. Soc, October, pp. 237, -pumilio
238. Living form.

1907 Elephas africanus toxotis Lydekker Proc. Zool. Soc. London, p. 385. Typeloc: Mossel Bay, Loxodonta africana
western Cape Colony. Type fig.: Lydekker, ibid., text fig. 106. Living toxotis
form.

1907 Elephas africanus selousi Lydekker Ibid., p. 387. Typeloc: Mashonaland, Rhodesia. Type Loxodonta africana
fig. Lydekker,
: ibid., text fig. 108. Living form. selousi

1907 Elephas africanus peeli Lydekker Ibid., p. 393. Type loc: Aberdare Mts., Kenya Colony. Loxodonta africana
Type fig.: Lydekker, ibid., text fig. 114. Living form. peeli

1907 Elephas africanus cavendishi Lydekker Ibid., p. 395. Type loc: Lake Rudolf district. Type Loxodonta africana
fig.: Lydekker, ibid., text fig. 115. Living form. cavendishi

1907 Elephas africanus orleansi Lydekker Ibid., p. 398. Typeloc: North Somaliland. Type fig.: Loxodonta africana
Lydekker, ibid., text fig. 118. Living form. orleansi

1907 Elephas africanus rothschildi Lydekker Ibid., p. 399. Typeloc: French Sudan. Type fig. : Loxodonta africana
Lydekker, ibid., text fig. 119. Living form. oxyotis
Lydekker states that he takes as type the statuette of "Jumbo" in the British Musuem.
Professor Osbom (1931.846, p. 21) referred "Jumbo" to the subspecies Loxodonta africana
oxyotis, which would make rothschildi a synonym of oxyotis.

1907 E. [Elepha.'i] a. [africanus] cottoni Lydekker Ibid., II, p. 783. Type loc: Northeastern Congo. Loxodonta africana
Type fig.: Lydekker, /birf., text fig. 111. Living form. cottoni

1907 Elephas maximus zeylanicus Lydekker "CJame Animals of India," p. 15. Living form. Elephas indicus
ceylanicus

1907 Elephas rreticus Ba,te Proc Zool. Soc. London, pp. 238-250. Type loc: Near Cape Maleka, Paheoloxodon creticus
Crete. Cotypc figs. : Bate, ibid., PI. xii, figs. 1-3, PI. xiii, figs. 1, 2.

1907 Paleomaslodo7i Barroisi Fontier Ann. Soc g^ol. du Nord, XXXVI, pp. 150, 151. Type loc:
FaylJm, Egypt. Type figs.: Pontier, ibid., text figs. 1 and 2.
Synonym, in part (fide Matsumoto, 1922) of Phiomia wintoni and P. minor (cf. Vol.
I, p. 61, this Memoir, where the present author confirms Matsumoto's reference).

1907 Elephas [Loxodon) zulu Scott 3d Rept. Geol. Surv. (?)Natal and Zululand, pp. 259-262. Type Loxodonta zulu
loc: Zululand, southeast coast of Africa. Type figs. Scott, op. c(7., PI. xvii, :

fig. 6, and PI. xviii, fig. 1.

1908 Dinolherium ndricum Pilgrim Rec Geol. Surv. India, XXXVII, Pt. II, p. 156. Deinotherium indicum
Name abandoned by the author and the subspecific term gajetise substituted in 1912, gajense
pp. 16 and 17. See below under Dinolherium indicum Lyd. var. ffojVn.sv Pilgrim, 1912.

1908 Telrahelodon rrepusruli Pilgrim Ibid., p. 157.

See Ilemimasiodon crepusculi Pilgrim, 1912, below, this list.
 NOMENCLATURE OF THE PROBOSCIDEA 1403

Reference in
Year Name Author Present Memoir

1908 Stcflodon javnnoqnnesa Dubois Tijdschr. Noderl. Aardr. (Jonoots Amsterdam, (2), XXVB, No. 6,

p. 1245.
Von Kocnig.swald, 1933.1, p. 105: "Im grossen Ganzpn sctic'mi. Slegodon trigonocephalus
im Habitii.s von Stegodon ganesa nicht sehr verschieden gewpseii zu .sein. Dubois dachte .^ogar
daraii, die javauisfhe Art nur aLs eine Varietal der iiidiselien zu betrachten; ganz klar
wird man uber seine Auffa.ssung nicht, da er die Art einnial als Stegodon javanoganesa, ( 1908,

pg. 1245), einige 8eit«n weiter als Stegodon ganesa var. javanicus bezeiehnet (1908, pg.
1257).

1908 Stegodon Ganesa var. javanicus Duhoifi Ibid., p. 1257 Typeloc: . Kendong-SohirlitPii, Trinil, Stegodon a irdwana [or
.lava. Figuro not found by the pre.sent author. N. trigonocephalus]

1908 Elephas hysndrindicus Duhoi>i hoc. cit. Typo loo.: Kondeng-Schichtou, .lava. Typo fig.: I'al:eolo.rodon Iii/midrin-
First published figure of cotype.s is in the present Memoir (Fig. 1 160). diciis

1909 Elephns Wiisti Piiv\o\v Annuairc (!eol. Min. Russie, XI, pp. 171-174. Typo Un-.: Tiraspol Parclepha.s wusti
(gouv. Kherson), southern Russia. Cotype figs.: Pavlow, ibid., PI. v, figs.
1 and 2; 1910, Nouv. Mem. See. Imp. Nat. Mosoou, XVII, Livr. 2, PI. i,
figs. 1-12 (Figs. 10 and 12 being the .same as 1909, PI. v, figs. 1 and 2).

1909 Gomphotheriinn conodon Cook Amor. .Journ. XXVIII, p. 18.3.

Sci., Type loo.: Near Agate, Not a proboscidean, but
Sioux County, Nebraska. Type fig. Cook, /6/rf.,
: p. 18.3. Dinohyus peter.soni
Holland

1909 Tetrabelodon dinotherioides Andrews Cool. Mag., Deo. V, N. S., VI, p. 347. Type loo.: "Loup Trilophodon dinotheri-
Fork beds," northwestern Kan.sas. Type fig. Andrews, : ibid., p. 348. oides

1911 FAephas primus Sohlosinger Monat.sbl. \'er. Landeskunde v. Niederrstor., V, p. 244. (Not determined by the
present author]

1911 Mxritherium Andrewsi^cYAosi^ev Beitr. Pal. Geol. Osterr.-Ung., XXIV, p. 130. Type loc: Maeritheriumandrewsi
Fayum, Egypt. Type figs.: Andrews, 1906, "A Desoriptivo Catalogue of the
Tertiary Vertobrata of the Fayum, Egypt," Pis. viii, ix.

1911 Dinotherium hobleyi Andrews Abstract, Proc. Zool. Soc. London, (May), p. 35; Proo. Zool. Deinotheriam hobleiji
Soc. London, (December), p. 943. Typo loc: Near Karung>i, east .side of
Victoria Nyanza, Africa. Type fig. Andrews, ibid.. PI. xlviii, figs. 1, la.
:

1912 Dinotherium indicum Lyd. var. gajen.'ie Pilgrim Mem. Geol. Surv. India, Palaeont. Indica, N.S., Deinolherium indicum
IV, Mem. 2, p. 16. Typeloc: Gaj beds of Bugti Hills, Baluchistan. Type gajen.se
fig.: Pilgrim, ibid., PL iv, figs. 1-4.
Same tyi>e specimen as FMnotherium ndricum Pilgrim, 1908, above, this list.

1912 Hemim.astodon crepusculi Filgiim Ibid., p. 17. Type loc: Gaj zone of Kumbhi, Bugti Hills, Suina
Baluchistan. Type fig. : Pilgrim, ibid., PI. iv, fig. 5; see also PI. iv, figs. 6-9,
and PI. Ill, fig. 4.

Same as Tetrabelodon crepusculi Pilgrim, 1908, above, this list.

1912 FAephas primigenius Fraasi Dietrich Jahrosh. Ver. Naturk. Wiirttcmb., LXVIII, pp. 42-106. Mammonteus primi-
Type loc: Steinheim a.d. Murr, Germany. Type figs.: Dietrich, /6;V;., Taf. genius fraasi
I and II, also text figs. 2, 4, 11-14, 16-21, 24-26.

1912 Mastodon arvernensis Croiz. et Job. var. progressor Khomenko Amuiaire Geol. Min. Russie, XIV, Anancus arvernensi.'i

Livr. 6, pp. 159-165. Type loc Near Gavanosy, district of Ismail, southern
:
progre.^sor
Bes.sarabia, Ru.ssia. Type fig.: Khomenko, ibid., PI. i.

1912 Mastodon arvernensis Croiz. et Job. var. conservativus Khomenko Ibid., p. 165.
Synonym of Mastodon arvernensis Croiz. et Job. var. progressor Khomenko, 1912, above.

1912 Elephas aritiguns var. insularis Soerge\ Palseontogr., LX, p. 1. l>pe loc. : Carini, Sicily. Nomen nudum

1913 Ma.slodon angustidens vvlv. au.stro-germanica Wegnev Palaontogr. LX, pp. 255-263.
,
Typeloc: Trilophodon angusii-
NearOppeln, eastern Germany. Type fig.: Wagner, i'6/d., Taf. xv, fig. 2. dens var. austro-
germanicus
1404 OSBORN: THE PROBOSCIDEA
Reference in
Yeax Name Author Present Memoir

191:5 Telmhelodon corrugatus PWgrim Rcc. Cool. Surv. India, XLIII, Pt. IV, p. 293. Type loc: Syncnnoluphiis cor-
Ha.snot, India. Typo fig. Lydekker, 1880, Mem. (icol. Suiv. India, Palaeont.
: nigatus
Iiidica, (X), I, Pt. V, PI. XXXV, fig. 4, as Mastodon {Triluphodon) pandionis;
see also PI. xxxvi, fig. 1.

1913 Mastodon hasnotiVAgraa. Loc.cit. Type loc: Near Bhimbar, northwest of .lamniu, India. Type Synconolophus hasnoti
fig.: Lydekker, 1880, ibid., PI. xliv, fig. 3, as Madodon {Telralophodon)
sivalensis.

1913 Mastodon stegodontoides Pilgrim Ibid., p. 294. Type loc: Lehri, Punjab, India. Type fig.: Stegolophodon stegodon-
Lydekker, 1880, ibid., PI. xxxix, as Mastodon {Tetrulophodon) latidens. toides

1913 Telrabelodon tnacrogiialhusVilgnm Ibid., p. 309. Type loc: Near C'hinji, India. No published Trilophodon macro-
type figure found l)y the present author. See figure 219 of the jjresent Memoir. gnathuii

1913 Tetrabelodon angustidens var. chinjiensi.s Pilgrim Ibid., p. 316 (name). Holotype of Osborn Trilophodon chinjiensis
(Amer. Mus. 19421) found two miles west of Chinji Bungalow, India (see
Fig. 218 of the present Memoir).

1913 Elephas antiqum ¥sl]c. vsir. trogotUherioide.'i Zuffardi Palajont. Ital., XIX, pi). 130, 155. Type Farelephas Irogontheri-
loc: Piedmont, Italy; lectotype from Nizza della Paglia (Astesan) cotypes ; oides
from near San Paolo de Villafranca. Figures: ZufTardi, ibid., Tav. ix, figs.
3fl-66.

1913 Elephas primigenius Blum. var. trogontherii Pohl. ZufTardi Ibid., p. 167.
Cf Elephas primigenius mutation
. aslensis Deperet and Mayet, 1923, below, this list.

1914 Elephas afriranii.i Fran.s.seni iichoutcdcu Rev. Zool. Africaine, III, Fasc 2, p. 396. Type loc: Loxodonta africana
M'Paa near Bongo, northwest of Lake Leopold II. Type fig. Schouteden, : fransseni
ibid., PI. XI, figs. 1, 2. Living form.

1914 Elephas tnuximiis hirsutus Lydekker Abstract, Proc Zool. Soc. London, I, p. 20 (name only). Elephas indicus hirsu-
Type loc: Kuala Pila district of the Negri Sembilan province, Malay <ms [of doubtful val-
Peniasula. Type fig.: Lydekker, 1914, Proc. Zool. Soc London, I, text fig. 1, idity — see footnotes
p. 285; 1916, "Catalogue Ungulate Mammals in British Museum," text fig. on page 1332 of Chap-
25, p. 84. Formerly in the Gardens of the Zoological Society of London. ter XX above. Ed-—
itor.]

1914 Tetrabelodon willisloni Barbour Univ. Studies, Univ. Neb., XIV, No. 2, pp. 192-194. Type loc: Trilophodon {Geno-
Devil's (iulch, Niobrara River, Brown County, Nebraska. Type figs.: mastodon) willistoni
Barbour, ibid., Pis. v, vi, viii.

1914 Eubelodon 7norrilli Barhom Ibid., pp. 194-197. Type loc: Devil's Gulch, Niobrara River, Eubelodon morrilli
Brown County, Nebraska. Type figs. Barbour, ibid., Pis. ix-xii.
:

1914 Tetndyelodon liilli Rarbour Neb. Geol. Surv., IV, Pt. 14, p. 217. On this same page, in a footnote, Megabelodon lulli
the subgenus Megabelodon was suggested. Type loc: Exposures bordering
Snake River, Cheriy County, Nebraska. Type figs.: Barbour, ibid., Pis.
iii-vi. See also Barbour, 1917.1, p. 512.

1914 Mammut progenium Ua.y Iowa Geol.

Surv., Ann. Rept. for 1912, XXIIl, i)p. 368 373. Type Mastodon progenius
loc: gravel pit, Missouri N'alley, Harrison County, Iowa. Type fig.:
Cox
Hay, ibid., PI. xliv, figs. 1 and 2 (specimen first described and figured by
Calvin, 1911, Bull. Geol. Soc Amer., XXII, p. 213, Pis. xx, xxi, under the
name Mastodon a7nericanus).

1914 /{hdhdobiiniis mirifiriis Hay Ibid., p. 374. Stegomastodon ini-

1912 (those Mastodon mirificus Leidy, 18.58 (see above, this list), as t.lu' lijiciis
PoIiHk in
g(Uiotypc of Sicgoniaxlodtm (see Generic List above, p. 1379). In 1914 Hay made
Leidy's M. mirificus the type of his new genus Rhabdobunus, which, however, is invalid
because preoccupied by Pohlig's Stegomastodon (cf. Ilay, 1930, p. 633).

1915 Elephas hysiidrirns primitivus Soorgel Centraib. Min. Geol. Pal., No. 8, p. 250, No. 9, p. 283. (Not determined by the
present author]

1915 Dinotherium styriacum W\\hcT Mitt. Naturwiss. Ver. Steiermark, LI, pp. 113 117. Type loc: (Not determined by the
Oberdorf bei Weiz, Styria. Type fig.: Hilber, ibid., Taf. i, fig. 1. present author]
 NOMENCLATURE OF THE PROBOSCIDEA 1405

Reference in
Year Name Author Present Memoir

1915 Elephas hayi Barhour Amer. Jouni. Sci., (4), XL, pp. 129-134. Type loc: Croto, Saline Archidiskodon hayi
County, Nebraska. Type figs.: Hay, ibid., p. 130, fig. 1; p. 133, fig. 3; p.
134, fig. M.
1915 Elephas aurorx Matsumoto Scientific Gazette, Tokyo, III, No. 5, pp. 308-315 (Japanese only); Stegodnn nitrone
1918 Tohoku Imp. Univ., (2), Geol., Ill, No. 2,
1918, Sci. Kept. p. 52. Type loc.:
Mt. Tomuro, Kaga, Japan. Type fig.: Matsumoto, 1918, ibid., PI. xx, figs.
1-3.
Subsequently (1924, Journ. Geol. Soc. Tokyo, XXXI, No. 371, pp. 256, 257, 262)
made the genoty|ie of Parastegodon.

1915 .Ma.'itodon pirayuiensis Gez "Generalidades sobre paleontologia argentina: HI Mastodon platen- Cuvieroniiis pirnyuien-
sis —
Amegh. de Corrientes," 2' edicion, p. 35. sis

1915 Ma.'itodnn "ligoniferits" Cope and Matthew "Hitherto Unpublished Plates of Tertiary Mam- Tiilophndon ligoniferus
malia and Permian Vertebrata," Amer. Mus. Monograph Series No. 2. Type
loc: Black Hills, South Dakota. Type figs.: Cope and Matthew, op. rit.,
PI. cxxrv', figs. 1 and 2.

1916 Elephas maximus maximus Lydekker "Cat. Ungulate Mammals, British Museum," V, p. 82. Elephas indiais ben-
galensis

1916 Elephas anticjaus Recki Dietrich Arch. Biontol., IV, Heft I, p. 22. Type loc: Oldoway-Tufle, Palseoloxodon recki
Serengetisteppe, northern Tanganyika Territory, Africa. Lectotype fig.:
Dietrich, ibid., Taf. i, fig. 2; cotypes, Taf. i-viii (in part).

1916 Tetrabelodon osborni Barhour Amer. Journ. Sci., (4), XLI, pp. 522-529. Type loc: Near Trilophodon {Genomas-
Bristow, Boyd County, Nebraska. Type figs. : Barbour, /6/rf., p. 523, fig. 1; todon) osborni
p. 524, fig. 2; p. 526, fig. 3; p. 528, fig. 4.

1917 Mastodon (Riinolophodon) angtistidens Cuv. f. subtapiroidea Schlesinger Denk. Naturhist. Hof- Serridentinus subtapi-
mus., I, Geol.-Pal. Reihe I, pp. 30, 31, 35, 37. Type loc: Wies, near Eibis- roideus
wald (Styria), Austria. Cotj^pefigs. Schlesinger, ?6k/., Taf. iii, fig. 2; Taf.
:

IV, fig. 1 Taf. VII, fig. 3; Taf. viii, figs. 1 and 2, and p. 31, text fig. 3.
;

1917 Mantodon (Bunolophodon) longirostre Kaw^) forma sublatidens Schlesinger Ibid., pp. 101, 102. Stegolophodon .s)/6-

Tyjieloc: Near Teschen (Schlesien), Austria. Type fig. : Schlesinger, /6/f/., latidens
Taf. XVII, fig. 2.

1917 Mastodon (Bunolophodon) grandincisivuni Schlesinger Ibid., p. 119. Type loc: Maragha, Per.sia. Tetralophodon grandin-
Type fig.: Schlesinger, ibid., Taf. xxxiv, figs. 1 and 2. Paratypes: Taf. xv, cisivus
figs. 1 and 2, Mannersdorf near Angern, Austria.

1917 Goviphotherium gratumYiay Bull. Univ. Texas, No. LXXI, pp. 18-21. Type loc: Pittbridge, Cordillerion gratnm
Burleson County, Texas, on the Brazos River. Type figs.: Hay, ibid., PI. iii,
figs. 3 and 4, PI. iv, figs. 1 and 2.

1917 Gomphotherium elegans Hay Ibid., pp. 21, 22. Type loc: McPherson, Kan.sas. Type fig.: Hay, Tetralophodon elegans
1917, Proc U. S. Nat. Mus., LIII, No. 2198, PI. xxvi, figs. 1 and 2.

1918 Mastodon angustidens, Cuvier var. libyca Fourtau Ministry of Finance, Egypt, Surv. Dept., Trilophodon angusti-
pp. 84-89. Type loc: Moghara, northern Egypt. Type fig. Fourtau, op. : dens libycus
rit., text fig. 58.

1918 [Mastodon angustidens] mut. Pontileviensis Mayet (In Fourtau, op. rit., p. 88 name only.) — Trilophodon ponti-
Hab.: Chevilly, Pontlevoy, France. Cotypefigs.: Mayet, 1908, Ann. Univ. leviensis
Lyon, Nouv. Ser. I,— Sci., Med., Fasc. 24, Pis. vii, figs. 5, 6, xi, fig. 2 (as
Mastodon angustidens). See Vol. I, p. 283 and fig. 230 of the present Memoir.

1918 Mastodon Spe?iceri Fourtau Op. rit., pp. 89-91. Type loc: Moghara Desert, northern Egypt. Rhynehotherium{1)
Type figs.: Fourtau, op. cit., p. 89, fig. 60; p. 90, fig. 61. spenreri

1918 Rhynchotherium tlasralse Oshorn Bull. Geol. Soc. Amer., XXIX, pp. 134, 135. Type loc: Rhynehotherium
Tlascala, Valley of Mexico. Type fig. Vol. I, fig. 448, of present
: Memoir. tlo.'icala'

See Rhynehotherium browni O.sborn, 1936, below, this list.

1406 OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir

1919 Mtidofloii {.\[arnrnid) n/nerirajius Pcnn. fortna praetypicaScUesmgQV Mitt. (ieol. (les. Wien, XI, Pliumadoduii ameri-
1922 1). 142. Cotypp loc. Szabudka ( = Maria-Theresiopel), Batta Erd, Rakos-
: conns pmetypica
kcre.sztur, Szentlorincz, and Ajnaesko, Hungary. Original figures: Schlo-
singer, ibid., 1919, Taf. vi, figs. 2-4. Supplementary de.scri])tion and figures:
Schlesingcr, 1922, Gcol. Hungarica, Ed. Sep., II, Fase. 1, pi). 115, 116, 227-
230, Pis. xiv-xix.

1921 Trilophorh)) giga nteus Oshorn Amer. Mus. Novitates, No. 1, pp. 6-10. Type loc: Eastview, Trilophodon giganteus
near Dallas, (Gregory County, South Dakota. Type fig.: Osborn, ibid., p. 9,
fig. 4C.

1921 Tetralophodon bnrbouri Oshorn Ibid., pp. 9, 10. Type loc: Cambridge, Furnas County, ^[nn•iIlinba)^bf^ul•i.
Nebraska. Type fig.: Osborn, 1924, Amer. Mus. Novitates, No. 154, fig. 1.
See also figs. SOSF, 329D, and 337 of the present Memoir.

1921 Ma.'<liidon imtthem Osborn Amer. Mus. Novitates, No. 10, pp. 2-6. Typ(> loc: Snake Creek, Pliomadodon inatthewi
Sioux County, Nebraska. Type fig. Osborn, ibid., p. 3, fig. lA (Fig. 98A of
:

the present Memoir).

Osborn (1922. .')64, p. 4) lufurretl this sjjecies to Miomasludon malthewi; finally

(1926.706, p. 1) he mado it the type of his new genus Plionmxtodon.

1921 M<t.<<todon merr in m i Oahorn Ibid., pp. 4-6. Type loc: Thousand Creek, Humboldt County, Miomastodon merrinmi
Nevada. Type figs. : Osborn, /b/rf., p. 3, fig. IB, and p. 5, fig. 2.

1921 Madodon lapiioidex (uiuiicanus'&iMesingn (In Osborn, Amer. Mus. Novitates, No. 10, p. 2; Miomastodon lapirnides
1922 fig. 1, D and Dl, p. 3.) Schlosinger, 1922, Geol. Hungarica, II, Fasc. 1, pp. americanus
224-227, Taf. xiii, figs. 6 and 7, and Taf. xiv, figs. 1-4. Type loc. Tasnad, :

Usztato Komitat, Hungary.

1921 Trilophodon (Tetrabelodon) shepardi edensis Frick Bull. Dept. Geol. Univ. Calif., XII, No. 5, pp. Rhynchotherium
405-409. Type loc: Mt. Eden Hot Springs, San Bernardino County, shepardi edense
southern Cahfornia. Type figs.: Frick, ibid., p. 406, figs. 160-165.
Id the original description of Trilophodon (Tetrabelodon) shepardi edensis Frick, 1921,
the author described material which ultimately proved to belong to two different genera,
namely, Rhynchotherium, and IHbehdon { = C ordilkrioti) The molars (P'rick, 1921, figs. .

100 -165) are referable to Khyiicholherium shepardi edense (see Frick, 1926, Bull. Amer. Mus.
Nat. Hist., T,VI, Art. II, pp. 169-176); the premaxilte and tu.sks (Frick, 1921, PI. L, figs.
1,2) to Cordillerion edensis, 1936 (see below, this list).

1922 Dibelodon edensis Osborn (in part) Amer. Mus. Novitates, No. 49, p. 2.
Synonym of Cordillerion edensis, 1936 (.see below, this list).

1922 Miomastodon matthewi Osborn Ibid., p. 4.

,See note under Mastodon matthewi Osborn, 1921, above, this li.st.

1922 LoTodonta griqua U&ughion Trans. (!eol. Soc S. Africa, XXIV, jtp. 11 13. Type loc: Griqua- Melarchidiskodon
land West, Africa. Type fig.: Haughton, ibid., Pi. i, figs. 1, 2. griqua

1922 Pcdifomasiodon intermedius Matsumoto Amer. Mus. Novitates, No. 51, p. 2. Type loc: North I'alxotnastodon inler-
of Lake Qurun, Fayflm, Egypt. Type fig.: Matsumoto, ibid., text fig. 1. medius

1922 /'/uowiio os6orm Matsumoto Ibid., pp. 3, 4. Type loc: AUixandria Trail, Fayiim, Egypt. Phiomia osborni
Type fig.: Matsumoto, ibid., p. 4, fig. 3.

1922 M (idodon engelswiesensis IsX^hn "Die badischen Mastodonten," pp. x, 30. Type loc: Engels- Trilophodon en^els-
wies, Baden, ( iermany. Type fig. : Kliihn, op. rit.. p. 30, fig. 10. wiesensis

1922 Mn.stodon .fleinheimensi.-i Kliihn pp. x, xi, 35, 76. Type loc: Stcinheini, Baden, Gcr-
Op. rit., Trilophodon deinheim-
many. Fraas, 1870, .Jahresh. Vcr. naturk. Wiirttemb., XXN'l,
Typc fig.: en.sis

Hefte II, III, Taf. v, fig. 1 (as Mastodon arvernensis).

1922 .Mastodon gigantorostris Klahn Op. cil., pp. xi, 48, 50, 87, 131. Typc loc: Hcrmcrsheim, Hhcin- Tetralophodon giganto-
hessen, Germany. rostri.f
 NOMENCLATURE OF THE PROBOSCIDEA 1407

Reference in
Year Name Author Present Memoir
1922 Mastodon esselborne^isis Kliihn Op. ril., pp. xi, 50, 73. Typo loc: Esselborn and Westhofen, Trilophodon esselborn-
Rheinhessen, Germany. Type fig. Kliihn, op. crt., p. 92, fig. 24.
: ensis

1922 Mastodon wahlheimensis Kl&hn Op. cil., pp. xi, 76. Type loc: Wahlheim and Esselborn, Turicius wahlheimensis
Rheinhessen, Germany. Cotype figs.: Klahn, op. cit., p. 77, fig. 17; p. 94,
fig. 25.

1922 Mastodon minutoarvernensislsXahn Op. cit., pp. xii, 102. Type loc: Herbolzheim, Baden, Anancus minutoarvern-
Germany. ensis

1922 Mastodon gigantarvernensis Klahn Loc. cit. Type loc. : Herbolzheim, Baden, Germany. Anancus gigantarvern-
ensis

1922 Trilophodon hicksi Cook Proc. Colo. Mus. Nat. Hist., IV, No. 1, p. 5. Type loc: Near Wray, Amebelodoit {Trilopho-
Yuma County, Colorado. Type figs.: Cook, ibid., p. 9, figs. 2, 3; p. 10, fig. don) hicksi
1; p. 11, fig. 2.

1922 Trilophodon paladentatus Cook Ibid., p. 6. Type loc: Near Wray, Yuma C^ounty, Colorado. Amebelodon (Trilopho-
Type figs. : Cook, ibid., p. 9, fig. 1 ; p. 13, fig. 5; p. 14, fig. 6. don) paladentatus

1922 Elephas jeffersonii Oahorn Amer. Mus. Novitates, No. 41, pp. 11-16. Type loc: Near Jones- Parelephas jeffersonii
boro, Indiana. Type fig.: O.sborn, ibid., p. 11, fig. 10.

1922 Elephas roosevelti Hay Proc Biol. Soc. Washington, XXXV, pp. 100, 101 (description without Parelephas jeffersonii
figure). Type loc: A.shland, Cass County, Illinois. See figure 968, p. 1096,
above, this Memoir.

1922 Mastodon oligobunis var. antiquissima Freudenberg Geol. u. Pal. Abh., XIV (XVIII), Heft III, Cordillerion oligobunis
pp. 118-120. Type loc: Valley of Amajaque, Hidalgo, Mexico. Type fig. : antiqiiissimus
Freudenberg, ibid., Taf. ix, figs. 3, 3a.

1922 Mastodon oligobunis vnr. Felicis Freudenberg Ibid., pp. 120-123. Type loc: Puebla, Mexico. Cordillerion{1) oligo-
Type fig. : Freudenberg, ibid., Taf. x, figs. 1, la. bunis felicis

1922 Mastodon oligobunis v&r. intermedia Freudenberg Ibid., pp. 123-126. Type loc: Mexico, Cordillerion{1) oligo-
exact locality not recorded. Type fig.: Freudenberg, ibid., Taf. xii, figs. bunis intermedius
1,2.

1922 M. [Mastodon] oligobunis var. progressa Frcudenherg Ibid., p. 127. Type loc: Cannada [can- Cordillerion oligobunis
yon] de Aculcingo, Mexico. Type fig. Freudenberg, /b/rf., Taf. xiv, figs. 1, 2.
: progressus

1922 El. [Elephas] Columbi var. Felicis Freudenberg 147-152. Type loc: Tecamachalco,
Ibid., pp. Parelephas columbi
Puebla, Mexico. Type fig. : Freudenberg, ibid., Taf. xvi, fig. 4. felicis

1922 El. [Elephas] Columbi var. silvestris Freudenberg Ibid., pp. 152, 153. Type loc: Ejutla, Archidiskodon impera-
Oaxaca, Mexico. Type fig.: Freudenberg, ibid., p. 146, fig. 19. tor silvestri.'i

1922 El. [Elepha.'i] Columbi var. Falconeri Freudenberg /6td., pp. 153-160. Type loc: Tequixquiac Archidiskodon impera-
Valley of Mexico. Paratype fig.: Freudenberg, ibid., p. 154, fig. 21 (original- tor falconeri
ly figured by Villada, 1903, Ann. Mus. Nac Mexico, VII, Lam. viii, as El.
primigenius Blum.)

1922 El. [Elephas] Columbi var. imperator Leidy, Freudenberg Ibid., pp. 160-171. Typical example Archidiskodon impera-
from Spokam Bar, near Helena, Montana. Fig.: Freudenberg, i6td., p. 55, tor
fig. 22.

1923 E. [Elephas] meridionalis mutation cromerensis Dep^ret and Mayet Ann. Univ. Lyon, Nouv. Archidiskodon meridi-
M6d., Fasc 43, Deuxieme Partie, pp. 150, 152, 157. Type loc:
Ser., I,-Sci., onalis cromerensis
Kessingland, Suffolk, England. Type fig.: Dcperet and Mayet, ibid., PI. ix,
fig. 1.

1923 E. [Elephas] p7-imigenius mutation astensis Dep&ret and Mayet Ibid., pp. IS3, I8i. Type loc: Mammonteus primi-
San Paolo de Villafranca, northern Italy. Type fig.: Dep^ret and Mayet, genius a-ttensis
ibid., PI. XI, fig. 5 (original in Musfe du Palais Carignan, Turin). Paratype
fig.: Ibid., PL xi, fig. 6 (original in Mus. Geol., Turin); figured by Zuffardi,
1913, Palaeont. ital., XIX, Tav. vi [xii], fig. 2a, and Tav. v [xi] fig. 8a, as
Elephas primigenius Blum var. trogontherii Pohl.
 :

1408 OSBORN: THE PROBOSCIDEA

Reference in
Year Name Author Present Memoir
1923 Murilheriiim aiiccdnde Pctronievics Aim. Mag. Nat. Ilist., (9), XII, p. 57. Tyjx'loc.: FayOm, MariUienuin unceslrale
Egypt. Type fig. Petronievics, ibid., p. 56, fig. 1.
:

1923 Anancus brazosius Hay Pan-Amcr. Geol., XXXIX, pp. 112-114. Type loc. : Brazos River, near Trilophodon iJTetra-
San Felipe, Texa.s. Type fig.: Hay, ibid., PI. viii, figs. 1, 2. lophodon) brazosius

1923 Serridentinns .fimpliridens Osborn Amer. Mus. Novitates, No. 99, p. 2. Type lop.: Near Pierce, Trilophodon simplici-
Polk County, Florida. Type fig. : Figure 231 of the present Memoir. den.s

1923 Trilophodon pmgrea.'ms Oshorn Type loc: Driftwood Creek, Hitchcock County,
Ibid., p. 3. Serridentinus progres-
Nebraska. Type
Cope, 1889, Amer. Naturalist, XXIII, p. 202, PI. xi,
fig.: sus
first as Tetrabeiodon angustiden.'i proauus, then as Tetrabelodoii jiroavu.'s (sec
Figs. 360 and 361 of the present Memoir; also p. 403, text).

1923 Ti'lndDphndon premmpester Oahorn Loc. cit. Type loc: Harlan County, Nebraska. Type figs. Morrill in bnrbouri
Figures 308E, 329E, and 339 of the present Memoir.

1923 Rhyncholherium rectidens O.sborn Loc. cit. Type loc: li^ight miles west of Pawnee Buttes, Khynrhotherium
Weld County, Colorado. Type figs.: Figures 461 and 481 of the present rectidens
Memoir.
1923 Rhyncholherium falconeri Osborn Ibid., pp. 3, 4. Type loc: Mt. Blanco, Llano Estacado, Texas. Rhynchotherium
Type Figure 468 of present Memoir (originally figured as "Tetrabeiodon
fig. : falconeri
shepardii Leidy" by Coi)e, 1893, 4th Ann. Rept. Geol. Surv. Texas, PI. xv).

1923 Elephas ivashinglonii Osborn Ibid., p. 4.Type loc Pine Creek, Whitman County, Washington.
: Parelephas ica.<ih ing-
Type figs.: Figures 972, 975, 893, B, Bl, of the present Memoir. tonii

1924 Ilnrpaqonotherium cuiiddenxe Fischer de Waldheim (In Sherborn, "Index Animalium," 1924, Pt.
\', p. 1022.)
Probably same as Harpagnwtherium canadense Fischer, 1808, above, thi.s list.

1924 Serridentiyius rnongolietisi.'i Oshorn Amer. Mus. Novitates, No. 148, pp. 1-3. Type loc: Loh, Serridentinus mongo-
near camp, Mongolia. Type fig.: Osborn, ibid., p. 2, fig. 1. liensis

1924 Elephas antiquum rumanus S. StefSnescu Compt. Rend. Acad. Sci. (Paris), CLXXIX, p. 1418. Archidiskodon plani-
Type loc: Tvilucesti (Covurlui), Rumania. Type fig.: Athanasiu, 1912 frnns riinianiis
[1915], Annuar. Inst. (ieol. Romaniei, \T, PI. xvii, fig. 4 (as Elepha.s cfr.
meridionalis). Refigured by StefSnescu, 1927, "30 Notes sur I'Organisation
des Molaires et sur la Phylogenie des Elephants et des Ma.stodontes" (see
Fig. 857 of the present Memoir).

1924 Elephas ajdiquiis germanicus S. Stefanescu Loc. cit. Type loc: Tanganu (Ilfov), Rumania. Ilesperoloxodon a ntiqu-
Type figs.: Steffinescu, 1927, op. cit., Figs. A, B (see Fig. 1089 of the present ns germanicus
Memoir).
1924 Elephas namadiais naumaru)i Makiyama, —
Chikyu The Globe, I, p. 381 (in Japanese; fide PaliFoloj-odon namadi-
Matsumoto, original not available to present author) 1924, Mem. College ; cus niiiimnnni
Sci. Kyoto Imp. Univ., (B), I, No. 2, pp. 255-264 (in English). Type loc:
Sahamma, Totomi Province, Japan. Type figs.: Makiyama, Chikyd, I, PI.
viii; Mem. College Sci. Kyoto Imp. Univ., (B), I, Pis. xii-xv, and xvi, fig. 1.
The following is an excerpt from Dr. Makiyama '.s article on "Japonic Probosciflea,"
Mem. Kyoto Imp. Univ., Ser. B, Vol. XIV, No. 1, Art. 1, p. 40, giving c(;rtain of
Coll. Sci.
the results of his recent studies (May, 1938): "This is the most common fossil ('Icphant in
Japan. The name of subs)x;cies was given to the stenocoroninc form |which| came from
Sahanma near Haniamatu in Sizuoka-ken (Totomi) covering the Naumann's E. luimadicun
from Yokosuka as it be different from the type E. namadicus Falconer & Cautley, 184ti
which is euryeoronine. ,\fter that time, I have had a number of chances to make interesting
observation about the different forms of teeth, and now I have a thought that all the
Japonic forms are equally separated from the Indian E. namadicun. The names listed are
the synonyms:
E. namadicus namadi Pohlig, 1893 of Makiyama, 1924
Parelephns proiomamnmnteus Matsumoto, 192-1
Loxodonia {Pdlaeoloxodon) tuiinadica yabei Matsumoto, 1929
EU'phas iPalaeoloxodott) namadicus setoensis Makiyama, 1929
E. ituiicus buski Matsumoto, 1929
Euekphns Irogontherii oi Matsumoto, 1924, and Makiy;iinn, 1924
Palaeoloxodon yokohamanus Tokunaga, 1934)"
Also see note below under "1929 Elephas (Palxoloxodon) namadicus setoensis Maki-
yama."
 .

NOMENCLAIURE OF THE PROBOSCIDEA 1409

Reference in
Tear Name Author Present Memoir

1924 Elephas namadicus namadi Makiyama Chikyu, I, p. 381 (in Japanese); Mem. College Sci.
Kyoto Imp. Univ., I, No. 2, pp. 263, 264 (in English). Type loc.
(B), Dredged
:

off island of Shodo, Saniiki Province, Japan. Type figs.: Makiyama,

Chikyu, I, PI. viii; Mem. College Sci. Kyoto Imp. Univ., (B), I, PI. xvi, fig. 2.
See note above under "1924 Elephas namadicus naumanni," also note below under
"1929 Elephas (Palseoloxodon) namadicus setoensis Makiyama."

1924 Euelephas protomammonteus Matsumoto Journ. Geol. Soc. Tokyo, XXXI, p. 262 (in Japanese); Palieoloxodon proto-
1926, Sci. Kept. Tohoku Imp. Univ., (2), Geol., X, No. 2. pp. 43-50 (in mammonteus
English). Type loc: Nagahama, Town of Minato, Kimitsii District,
Province of Kazusa, Japan. Type figs.: Matsumoto, 1926, ibid., PI. xviii,
figs. 1, 2 (as Parelephas protomamnwnteus) PI. xix, figs. 1-3 (the paratype,
;

from same locality as type).

Sec note above, 1924, under Elephas namadicus naumanni Makiyama.

1924 Loxodonla {Palxoloxodon) tokunagai Mastumoto Ibid., p. 267 (in Japanese); 1929, Sci. Kept. Palxoloxodon tokuna-
Tohoku Imp. Univ., (2), Geol., XIII, No. 1, pp. 7-11 (in English). Type gai
loc: Soyama, Gokayama, Hira-mura Higashi-Tonami Dist., Province of
Etchu, Japan. Type figure not found by present axithor.
Referred to Archidiskodon tokunagai by Teilhard de Chardin and Trassaert in 1937
(Pal. Sin, (C), XIII, Fasc. 1, p. 44).

1924 Stegodon Matsumoto Ibid., pp. 333-335 (in Japanese). Type loc: Island
orietitaIi.'< fthodocn.si.s Stegodon orientalis
of Mitsugo (Mitsugo-Shima) and island of Shodo, Inland Sea, Japan. shodoensis
Type fig.: See Makiyama, 1938, Mem. Coll. Sci. Kyoto Imp. Univ., (B),
XIV, No. 1; p. 18, fig. 7 (holotype palate).

1924 Hcmimastodon annectens Matsumoto Ibid., pp. 401, 405 (in Japanese). Type loc: Banjobora, Serridenlinus annectens
Kaminogo-mura, Kani District, Province of Mino, Japan. Type figs.:
Matsumoto, 1926, Sci. Kept. Tohoku Imp. Univ., (2), Geol., X, No. 1, Pis.
I, figs. 1 and 2, ii, figs. 1-3.

1924 Trilophodon sendaicu.s MaUxinmU} /6if/., pp. 402,408 (in Japanese). Type loc: Kitayama, near Trilophodon sendaicus
Sendai, Province of Rikuzen, Japan. Type figs.: Matsumoto, 1926, Sci.
Rept. Tohoku Imp. Univ., (2), Geol., X, No. 1, Pis. iii and iv.

1924 Elephas africanus mogambiru.'i Frade Bull. Soc Portugaise Sci. nat., IX, Fasc. 3, pp. 131, 133. Loxodonta africana
Hab.: Maputo, Mozambique. Type fig.: Frade, ibid., text fig. 5. Female moQambica
formerly in the Jardin de Zoologique, Lisbon.

1924 Loxodonta africana Zukowskyi Strand (In Zukow.sky, Arch. Naturges., XC, Abth. A, Heft I, p. Loxodonta africana
68.) Hab.: Kaoko District, southwest Africa. Living form. zukoirskiji

1924 Panlephas jeffersonii Osborn Amer. Mus. Novitates, No. 152, pp. 1, 4, 7. Type loc:
progres.'ius Parelephas progressus
Zanesville,Mu.skingum County, Ohio. Type figs.: Osborn, 1922, Amer.
Mus. Novitates, No. 41, p. 13, fig. 11, and p. 14, fig. 12 (as paratypes of
Elephas jeffersonii)

1924 Mammonteus primigeniu.s compressus Osborn Ibid., pp. 5-7. Type loc: Rochester, Indiana. Mammonteus primi-
Type Osborn, 1922, Amer. Mus. Novitates, No. 41, p. 8, fig. 8 (as
figs.: genius compressus
Elephas primigenius) 1924, Amer. Mus. Novitates, No. 152, p. 6, fig. 2.
;

Paratype fig.: Osborn, 1922, Amer. Mus. Novitates, No. 41, p. 7, fig. 7
(from Eschscholtz Bay, Ala.ska).

1924 Stego7nastodon texanus Osborn Amer. Mus. Novitates, No. 154, p. 3. Type loc: Blanco for- Stegomastodon texanus
mation, Llano Estacado, Texas. Type figs.: Figures 639, 642, 644A, 645D,
of the present Memoir.

1924 StcgomasLodon aflunix Oabovn Loc. cil. Type loc: Abovit two miles east of Akron, Plymouth titegomastodon aftonix
County, Iowa. Type r.AP originally figured as Mammul mirijicuni by Calvin,
1909, Bull. Geol. Soc. Amer., XX, PI. xxvii; refigurcd as Rhabdobunus
mirificus by Hay, 1914, Iowa Geol. Surv., XXIII, PI. l. Type l.M', see
figure 650 of the present Memoir.
1410 OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir
1924 Stegomaslodon arizonsp flidlcy Amcr. Mus. Xovitatcs, No. 154, p. 4 from (lidley
(In Oshoni, — Slegomastodoii arizonse
1926 MS.); S. Geol. Surv., Profe.ss. Paper 140-B, p. 86.
(lidley, 1926, U. Type
loc: Curtis Flats, Cochise County, Arizona. Type figs.: Gidley, ibid., Pis.
xxxiii-xxxv, PI. XXXVIII, figs. 1-3, PI. XXXIX, figs. 1-3.

1924 Stegomastodon nebrascensis Oshorn Ibid., p. 5. Type loc: Snake Creek, Sioux County, we.«tern Serridenlt7ius (Ocalien-
Ncbraska. Type fig.: Osborn, ibid., p. 4, fig. 2 B. tinus?) nebrascensis

1925 Mammonteits primigenius ameriranus Osborn Proe. Amer. Phil. Soc, LXIV, No. 1, p. 33. Mammonleus primi-
Sec E. americanus DeKay, 1842, above, from near Rochester, New York. 9^" '
"-^ americanus

1925 Elephas scotti Barbour Nob. State Mus., Bull. 2, I, pp. 21-24. Type loc.: Five miles south of Archidiskodon inipera-
Staplehurst, Seward County, Nebraska. Type figs. Barbour, i6«rf., text figs. : lor scotti
7-10.
[On page 1025, of the pre.sent Memoir, appears the following statement by Pro-
fessor Osborn: "Awaiting furtlier evidence, O.sborn is inclined to regard the type of
'Elephas scolli' as representing a young individual of Archidiskodon imperalor." Neverthe-
l<'ss he treated this species under the heading Archidiskodon impfrator scotti.]

1925 Telrabelodon abeli Bsxrhom Neb. State Mus., Bull. 9, 1, pp. 91-94. Type loc: Devil's Gulch, Trilophodon abeli
Brown County, Nebraska. Type figs. : Barbour, ibid., pp. 92, 93, figs. 52-56.

1925 Elephas maibeni BsLrhouT Neb. State Mus., Bull. 10, I, pp. 95-118. Type loc: About sixteen Archidiskodon. impera-
miles north of Curtis, Lincoln County, Nebraska. Type figs.: Barbour, ibid., tor maibeni
text figs. 58-60, 63-70, 72, 74, 76-87.
In 1926 described by Barbour as Archidiskodon maibeni.

1926 Archidiskodon maibeni Barbour Neb. State Mus. Bull. 11, I.

See Elephas maibeni Barbour, 1925, preceding item.

1926 M amw ul francisi H&y Journ. Wash. Acad. Sci., XVI, No. 2, pp. 35-39. Type loc: Brazos ?Rhynchotherium
River, Pittbridge, Burleson County, Texas. Type figs.: Hay, ibid.,p.3&. francisi
fig. 3; p. 37, fig. 4.

1926 Mammul oregonense Hay /6/r/. ,])]>. 39, 40. Type loc: Rye Valley, Dixie Creek, Baker County, Mastodon oregonensis
Oregon. Type figs. : Hay, ibid., p. 36, figs. 1, 2.

1926 .M astodoK americaiius plicatus Oshorn Amcr. Mus. Novitates, No. 238, p. 1. Type loc: Walnut, Mastodon americanus
Bureau County, Illinois. Type fig.: O.'^born, ibid., p. 2, fig. 1. plicatus

1926 Zj/golophodnn pyrriinicii.s niirelianensis Osborn Ibid., j). 2. Hab.: Chevilly(?), Cussegrain, Zygolophodon pyrenai-
Avaray, France. No type figure available to the present author. cus aitrelianensis

1926 Tiiriciiis liiricensis si7>u)rrrn.tis Osborn Ibid., pp. 3, 4. Type loc: Near Simorre (Gers), Frances Turicius turicensis
Ty])e figs.: Lartet, 1859, Bull. Soc geol. France, (2), XVI, PI. xv, fig. 3 (as simoi-rensis
Mastodon tapiroides) Osborn, 1926, ibid, ]). 4, fig. 2.
;

1926 Serridenlinus browni Oshorn Ibid., p. i. Type loc: Near Chinji Bungalow, India. Type figs. : Serridentinus browni
Osborn, ibid., p. 5, fig. 3; p. 6, fig. 4.

1926 Serridentinus republicanus Oi^horn Ibid., yi. (i. Type loc: Republican River formation, north- OcaUentinus {Serriden-
westcrn Kansas. Type figs.: Osborn, ibid., p. 7, fig. 5; p. 8, fig. 6. linus) republicanus

1926 Serridcntinu.f obliquidens Oshorn Ibid., p. 9. Typ(\ loc: Phosphate beds near Charleston, Ocalientinus (Scrriden-
South Carolina. Type fig.: Oshorn, ibid., p. 9, fig. 7. tin.us) obliquidens

1926 Serridentinus anguirivalis Oshorn Ibid.,\). 10. Tyjx'loc: South-central Sioux County, western Scrridrnlinus aiiguiri-
Nebraska. Type fig.: Osborn, ibid., p. 10, fig. 8. valis

1926 Serridentinus brewslerensis Oshorn Ibid., p. 11. Type loc: Brewster, Polk County, Florida. Serridentinusbrewster-
Type fig.: Osborn, ibid., p. 11, fig. 9. ensis
 :

NOMENCLATURE OF THE PROBOSCIDEA 1411

Rsferenco in
Year Name Author Present Memoir
1926 Serridentinu^ guatemalensis Oi^hovn Ibid., p. 12. Type lor.: C'liiiiautla, Guatemala, Central Serridenlinus guatemal-
America. Type fig.: O.sborn, ibid., p. 12, fig. 10. ensis

1926 Rhijncholherium anginrivalis Oshorn Ibid., p. 13. Typeloc: Sioux County, western Nebraska, Rhynchoiherium angui-
Quarry 3 (Olcott Hill). Type fig.: Oshorn, ibid., p. 13, fig. 11. rivale

1926 Anancus falconeri Oshorn Ibid., pp. 13, 14. Type loc.: Red or Norwich Crag of Suffolk, Anancus falconeri
England. Type fig. Originally described and figured by Falconer as Masto-
:

don (Tetralophodon) arvernensis (1857, Quart. Journ. Geol. Soc. London,

XIII, PI. XII, figs. 3, 4; 1868, "Palaeont. Mem.," II, PI. iv, figs. 3, 4). Osborn,
ibid., p. 14, fig. 12.

1926 Parelephas protomamnwnteus (Matsumoto) typicus Matsumoto Sci. Rept. Tohoku Imp. Univ., Palxoloxodon proto-
(2), Geol., X, No. 2, pp. 43-48. jnamnionteus
Originally named Euelephas yrotomamtiwnteus by Matsumoto in 1924, p. 262 (publish-
ed See above, this list.
in Japanese).
[According to Shikama (1937, p. 164) thi.s species is synonymous with the species
antiquus. —
Colbert-Simpson, letter, July 12, 1939. Editor.] —
1926 Parelephas protomammonieus proximus, nnit. nov. Ma.tsun\oto Ibid., pp. 48-50. Type loc: Palseolojodon proto-
Isone,Kokubo, Onuki-mura, Kimitsu District, Pro\ince of Kazusa, Japan. mammonieus proxi-
Type fig.: Matsumoto, ibid., PI. xxiv, figs. 1, 2. m us
[If protomamnwnteus, iiccoiding to Shikama, is a synonym of aniiquiis (see preceding

note under Parelephas proiomamnwmteus lypicus Masumoto), then Shikama was justified
in emending the name of Matsumoto's second species to read Parelephas proximus proxi-
—
mus. Colbert-Simp.son, letter, July 12, 1939.— Editor.]

1926 Elephas eellsi Hay Journ. Wash. Acad. Sci., XVI, No. 6, pp. 154-159. Type loc. Port Williams, : Parelephas{1) eellsi
Clallam County, Washington. Type figs. Hay, ibid., p. 156, figs. 1, 2.
:

1926 Anancus orarius Hay Proc. U. S. Nat. Mus., LXVUI (1927), No. 2625, Art. XXIV, pp. 8-14. Cordillerion orarius
Type loc. West bank of Aransas River, San Patricio County, near town of
:

Sinton, Texas. Type figs.: Hay, ibid., PI. ii, fig. 1 PI. in, fig. 1; PI. rv, fig.
;

1; PI. v, figs. 1-3.

1926 Anancus defloccatus Hay Ibid., pj). 14-16, 18. Type loc: West bank of Aransas River, San Cordillerion defloccatus
Patricio County, near town of Sinton, Texas. Type figs.: Hay, ibid., PI. vii,
and PI. VIII, fig. 1.

1926 Anancus bensonensis Gidley U. S. Geol. Surv., Profess. Paper 140-B, pp. 85, 86. Type loc: Cordillerion bensonen-
Near Benson, Cochise County, west side of San Pedro \'alley, Arizona. Type sis
fig.: Gidley, ibid., PI. xxxii.

1926 Trilophodon (Serridentinu.s) pojoaquensis Frick Bull. Amer. Mus. Nat. Hist., LVI, Art. II, pp. Trilophodon pojoaquen-
125, 142-150, 161, 162-165. Type loc: About twenty-four miles north of sis
Santa Fe, Santa Fe County, near Pojuaque, New Mexico. Type figs.:
Frick, ibid., p. 148, fig. 26, and p. 144, fig. 22A. Amer. Mus. 21115(type skull).
The skull (F.: A. M. 2112.'>), originally considered by Fr'u:k as a jmratype of Triloph-
odon (Serridenlinus) pojoaquensis, proved on subsequent examination to be referable to his
new genus and species Ocalientinus ojocaliensis.

1926 ^Trilophodon {Serridenlinus) leidii Frick Ibid., pp. 137, 140, 141, 155, 169. Type loc: Alachua Ocalientinus (Serri-
clays, Mixson's bone bed, near Williston, Levy County, Florida. Type figs. denlinus) floridanus
Leidy and Lucas, 1896, Trans. Wagner Free Inst. Sci., IV, PI. iv, figs. 9-11; leidii
Frick, 1926, ibid., figs. 20A,B.

1926 'Uthynchotherium (Dibelodon) edensis Frick Ibid., pp. 169-176. Rhynchoiherium

See note under Trilophodon (Telrabelodon) shepardi edensis Frick, 1921, above, this list. shepardi edense

1927 E. [Elephas] primigenius Matsumotoi Dietrich Neues Jahrb. Min., I, .\bth. B, Referate, p. 314. [Not determined by the
[The specimen considered as the type of Elephas primigenius malsuinotoi by Dietrich, present author]
1927, is regarded by Shikama (1937, pp. 164, 16.i) as belonging to Parelephas proximus (sec
above, this list, under Parelephas protomammonteus proximus mut. nov. Matsumoto, 1926);
therefore the former of these names becomes a synonym of the latter. Colbert-Simpson, —
letter, July 12, 1939.— Editor.]
 :

1412 OSBORN: THE PROBOSCIDEA

Reference in
Year Name Author Present Memoir

1927 A inebebdon flick i Bavhour Neb. State Mils., Bull. 13, I, i)]).
131-134. Type loc: Freedom, AmebeUxlon fricki
FrontiiM- (
'ouiity, Nebra.'ika. Type fig.s. : Barbour, ibid., text figs. 89-91.

1927 Archiditikodon trannvaak-n.'iis ]:)art Supplement, Nature, No. 3032, pp. 41^8. Type loe.: Near Pala'olo.rodon trans-
Bloemhof, South Africa. Type figs.: Dart, ibid., p. 47, fig. 6 (right); p. 48, vaalensis
fig. 7 (left).

1927 Archidiskodon Sheppardi Dart Loc. cit.. Type loc.: Near Bloemhof, South Africa. Type figs. Palseoloxodon sheppardi
Dart, ibid., p. 47, fig. 6 (left) ; p. 48, fig. 7 (right).

1927 Elephas indinis Buski Matsumoto Sci. Rcpt. Tohoku Imp. Univ., (2), (ieoL, X, No. 3, pp. 57, IPalseoloxodon buski
58. Type loc: Ninohe District, Province of Mutsu, Japan. Type fig.:
Matsumoto, ibid., PI. xxvii, figs. 2, 3.

1927 Leith-Adnnma siwalikiensisi Matsumoto Japanese Journ. (ieol. and Geog., V, No. 4, Art. XII. Archidi.skodon plani-
Type loc: India. Type figs.: Falconer and Cautley, 1845, "Fauna Antiqua frons
Sivalensis," PI. xi, fig. 4; 1847, PI. xiv, fig. 8 (as Elephas planifrons).

1928 Elephas priniigeriitis Blumenbacli var. n. pachiiganalis Schroeder Jahrb. preuss. gcol. Landes- [Not determined by the
anstalt, XLVHI,
pp. 701, 702, 718. Type loc: Rixdorf, Germany. Type present author]
fig.: Schroeder, ibid., Taf. xxxvi, figs, la, lb.

1928 Trilophodon {Serridentinus?) inopinaius Borissiak and Beliaeva Bull. Acad. Sci. U. S. S. R., CI. Trilophodon inopinaius
Sci., Phys.-Math., pp. 241-252. Type loc: Jilancik beds of the Turgai
region, V. S. S. R. Typi^ fis^-: Borissiak and Beliaeva, ibid., Pis. i, ii.

1928 Airhidiiikodon .vib plan ifionsOsborn Nature, ( 'XXI, No. 3052, i)p. 672, 673. Type loc. Sydney- : Archidiskodon sub-
on-Vaal, Vaal River, South Africa. Type fig. (Osborn, ibid., p. 672, fig. 1. : planifrons

1928 Archidiskodon broomi Osborn Ibid., pp. 672, 673. Type loc: The Bend, Vaal River, near Archidiskodon broomi
Kimberley, South Africa. Type fig. : Osborn, ibid., p. 672, fig. 2.

1928 Elephas exilis Stock and Furlong Science, N. S.,LXVIII, No. 1754, p. 140. Type loc: Santa Archidiskodon exilis
Rosa Island, California. Type fig.: Stock, 1935, Sci. Monthly, XLI,
September, p. 210, fig. 6 (see also Figs. 920 and 921 of the present Memoir).

1928 Elephas haroldcooki Hay Proc Colo. Mus. Nat. Hist., VIII, No. 2, Pt. I, p. 33. Type loc: Archidiskodon harold-
Frederick, Oklahoma. Type figs.: Hay and Cook, 1930, ibid., IX, No. 2, cooki
PI. Ill, fig. 1, PI. V, fig. 1, Pis. XIII and xiv.

1928 Trilophodon phippsi Cook Proc. Colo. Mus. Nat. Hist., VIII, No. 4, pp. 37-43. Type loc: Near Megabelodon phippsi
.\ins\vorth. Brown County, Nebraska. Type figs.: Cook, ibid., Pis. i-iii.

(Ill !i footnote on imgo 328 of Volume I of tho present Memoir Trilophodon phippsi is

included among the s|jeeies referable to Megabelodon, the"Spoonbill Mastodonts." While on

])age 707 Professor Osborn mentions Megabelodon Itilli, M. cruziensis, and M. joraki, we
fail to find anywhere term Megabelodon phippsi. However, it was evidently
tlie use of tlie

his intention to assign phippsi to Megabrlodnn because of its tuskless mandible, the dis-
tinctive character of this genus. Therefore on page 738 it has been listed vmder Megabelo-
don with the three above-mentioned species. Editor.] —
1928 Serridenliinisfricki Peterson Mem. ( 'arnegie Mus., XI, No. 2, ])]i. 111-121. Type loc: Northern Trilophodon fricki
flanks of Douglas Mountain on Weller Ranch, Moffat County, Colorado.
l>pe figs.: Peter-son, ibid., Pis. xi-xiv, and text figs. 21,22; also restoration
by A. AvinotT (Pi. xv).

1928 lM.rndonlu afiicana (ingolensis Frade "Titulos e trabalhos cientificos (Curriculum vitae)," p. 15. Loxodonta africana
Type loc: Region of Cunene, .southern Angola, Africa. Living form. (uigolensis

1928 Mastodon andium Kraglievichii Berro "Un nuevo Mastodon en la R. (>. del Uruguay." Cordillerion andium
AVdc Cabrera, 1929, Kov. Mus. La Plata, XXXII, 119, 111; not seen by the kraglievichii
pp.
l)rescnt author.

1928 Mastodonte de Alangasi.

See '1931 Bunolophodon poslremus Spillmann," below, this list.
 :

NOMENCLATURE OF THE PROBOSCIDEA 1413

Reference in
Year Name Author Present Memoir
1928 Platybelodon Danovi Borissiak Ann. Soc. Paleont. Russie, VII, pp. 105-120. Type loc: Kuban Platybelodon danovi
region, North Caucasus, Rus.sia. Type fig.: Borissiak, (6id., Taf. viii, fig.s.
1-4.

1928 Tetrabelodon {Bunolophodori) Ayora Spillmann El Ecuador Comercial, Ano VI, No. 57. Type Cuvieronius ayorx
1931 loc: Quebrada [canyon] of Chalang, near Punin, southeast of Riobamba,
Province of Chimborazo, Ecuador. Type fig.: Spillmann, ibid., fig. 2 (see
also figures 548 and 549 of the present Memoir). Supplementary description
Spillmann, 1931, "Die Saugetiere Ecuadors im Wandel der Zeit," Erster
Tell, p. 67.

1929 Elephas (Palasoloxodon) nnmadicits seioe7isis Makiyama Chikyil The Clobe, XII, No. 5, pp. — [Not determined by the
364, 365 (in Japanese). Type loc: Seto, Japan. present author]
[Dr. Jiro Makiyama in a letter to the editor, dated Kyoto, Augu.st 26, 1937, stated that
the name Elephas (Pals-oloxodon) nanmdiciis setoensis Makiyama appears on page 365 of his
article of 1929 in Chikyil, without diagnosis or figure, but by "designation points, as an
example, to the specimen given in his paper 'Notes on Fossil Elephant from Sahamma,
Totonii' (Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B, Vol. I, No. 2, 1924, pp. 263, 264, PI.

XVI, fig, 2, as Elephas namadicus namadi)." This was an outline in English of the note
subsequently given by him in Japanese in the publication ChikyO^The Globe, XII, No. .5,
PI). 364, 36.'j, November, 1929, with the translated title "Scientific Names of Fossils and
the International Rules of Zoological Nomenclature," pp. 3.58-366. Doctor Makiyama
further stated that during the intervening years he had changed his opinion somewhat and
that he hoped to publish the results of his studies in the near future. This article has since
ap|)eared under the title "Japonic Proboseidea," Mus. Coll. Sci. Kyoto Imp. Univ., Ser.
B, Vol. XIV, No. 1, .\rt. 1, May, 193S.— Editor.]

1929 A tnebelodon grangeii Oshom Nat. Hist., XXIX, No. 1, pp. 12-16. Type loc: Tairum Nor Platybelodon grangeri
1931 Basin, Mongolia. Typo fig.: Osborn, ibid., p. 15 (plate). Supplementary
description: Osborn and Granger, 1931, Amer. Mus. Novitates, No. 470, pp.
6, 7, text figs. 1 and 3A, Al.
Transferred to genus Platybelodon, ow-ing to fundamental differences in the stracture
of the tusks, e. g., the presence of dentinal rod-cones in Platybelodon (characteristic of
grangeri), but which are absent in Amebelodon.

1929 Toryriobelodon loomisi B&rhour Neb. State Mus., Bull. 16, I, pp. 147-150. Type loc: Sand Torynobelodon loomisi
Canyon, two and a half miles southwest of Republican City, Harlan County,
Nebraska. Type figs. Barbour, i6/rf., text figs. 98-100.
:

1929 Trilophodon atigustidens gaillardi Osborn Amer. Mus. Novitates, No. 393, p. 1. Type loc: Trilophodon angusli-
Villefranche d'Astarac (Gers), France. Type fig.: Osborn, ibid., p. 1, fig. 1. dens gaillardi

1929 Serridentinus bifoliatus Osborn Ibid., p. 2. Type loc: Brewster, Polk County, Florida. Type Ocalientimis (Serri-
fig.: Osborn, ibid., p. 2, fig. 2. deniinus) bifolialus

1929 Serrideniinus hasnotensis Osborn Loc. cit. Type loc: Near Hasnot, India. Cotype figs.: Serrideniinus hasnoten-
Osborn, ibid., p. 3, fig. 3. sis

1929 Serrideniinus metachinjiensis Osborn Ibid., pp. 4, 5. Type loc: One mile northwest of Chinji Serrideniinus metachin-
Bungalow, India. Type fig.: Osborn, ibid., p. 3, fig. 4. jiensis

1929 Serrideniinus chinjiensis Osborn Ibid., p. 5. Type loc: One mile and a half west of Chinji Serrideniinus chinjien-
Bungalow, India. Type fig.: Osborn, ibid., p. 4, fig. 5. sis

1929 Serrideniinus prochinjiensis Osborn Ibid., p. 6. Type loc. : Two miles west of Chinji Bungalow, Serrideniinus prochin-
India. Type fig.: Osborn, ibid., p. 5, fig. 6. jiensis

1929 Serrideniinus florescens Osborn Ibid., p. 6. Type loc: Kholobolchi Nor region, Mongolia, five Ocalienlinus (Serri-
toeight miles north of camp. Type fig. Osborn, i6td., p. 7, fig. 7.
: deniinus) florescens

1929 Hhyncholherium paredensis Osborn Ibid., pp. 6-8. Type loc: Mt. Eden Hot Springs, San Rhyncholherium
Bernardino County, California. Type fig.: Osborn, ibid., p. 7, fig. 8. shepardi edense
Originally figured by Frick, 1926, Bull. Amer. Mus. Nat. Hist., LVI, Art. II,
figs. 2, 8, 9 as {l)Hhynrholherium (Dibelodon) eden.'iis.

In the present Memoir (Vol. I, p. 496, also fig. 474) R. paredensis is regarded as a
synonym of R. shepardi edense.
 :

1414 OSBORN: THE PROBOSCIDEA

Reference in
Year Name Author Present Memoir

1929 Rhtinrhotheriutn clunjiensis Osburu Ibid., p. 8. Type loc: Two mik-s west of Chinji Bungalow, Rhynchotheriiim
India. Type fig.: Osborn, t6irf., p. 8, fig. 9. chinjiense

1929 Si/ricoriolophus dhokpathanensis Oshorn 76?rf., pp. 10-12. Type loc: Three miles west of Dhok Synconolophus dhok-
Pathan, India. Type fig.: Osborn, ibid., p. 9, fig. 10. pathanensis

1929 Synco7iolophus ptychodus Osborn Ibid., p. 12. Type loc: Four miles west of Chinji Bungalow, Synconolophus ply-
India. Type fig.: Osborn, ibid., p. 11, fig. 12. chodiis

1929 Synconolophus propathmiensis Oshovn Ibid., p\^. 12, 13. Type loc Three : miles east of Dhok Syuronolophns pro-
Pathan, India. Type fig. : O.sborn, ibid., p. 13, fig. 13. pathanensis

1929 Stegolophodon nathotensis Ofihorn 76«/., pp. 13-15. Type loc: Near Nathol, India. Type fig.: Skgolophodon nalhoten-
Osborn, ibid., ]).14, fig. 14. sis

1929 Slegulnphoilon raulleyi progres.ms Ofihorn Ibid., p. 15. Type loc Twelve miles ea.st of Chinji : Slegolophodon vaulkyi
Bungalow, India. Type fig.: Osborn, i6irf., p. 14, fig. 15. progressus

1929 Slegodon insignisbinnanints Oshorn /6zW., pp. 15, 16. Typeloe.: Mingoon, opposite Mandalay, Stegodon insignis
Burma. Type fig.: Osborn, ibid., p. 16, fig. 16. birmanicus

1929 Stegodon orientalis grangeri Osborn Ibid., pp. 16, 17. Typeloe: Yenchingkou, near Wanh.Kien, Stegodon orienlnlis
Provinec of Sz(>chuan, China. Tyi)e fig.: Osborn, ibid., p. 16, fig. 16. grangeri

1929 Stegodon pin jorensis Osborn Ibid., p. l&. Type loc: Three miles north of Siswan, India. Stegodon pin jorensis
Type fig.: O.sborn, ibid., p. 17, fig. 17.

1929 A irh idiskodon .'ionoriensisOsborn Ibid., p. 18. Type loc: One mile ea.st of Arizpe, northern Arehidi.'ikodon sonori-
Sonora, Mexico, on the Sonora River. Type fig.: Osborn, ibid., p. 18, fig. 18. ensis

1929 Parelephasfioridanus Osborn Ibid., p. 20. Type loc: Manatee County, Florida, two miles Parelephas Jtoridanus
south of Bradenton. Type fig. Osborn, ibid., p. 19, fig. 19. :

1929 Poirleplids rolinnbi cayennensis Os\)oru Ibid., pp. 20, 21. Tyjie loc: Cayenne, French (iui- Pnrelephas eohimhi
ana, South America. Type fig. Osborn, /6(W., p. 21, fig. 20. : '-ni/ennensis

1929 Klephas ploli/rcpholiis O.shoni Ibid., pji. 21, 22. Type loc: Near Si.'^wan, bed of Amilec Creek, Pldlclephos pltilyreph-
Simla Hills, India. Type fig.: Osborn, ib/rf., j). 22, fig. 21. aliis

1929 Elephas platyeephalii.'i nngii.^tidens Oshorn Ibid., jip. 22, 23. Type loc: Three miles west of fli/pselephas hi/sudrieiis
Chandigarh, Siwalik Hills, India. Type fig. : Osborn, ibjrf., p. 23, fig. 22.

1929 Losodonla (Paia;olo.rodon) nanuidiea (Yabci) Matsumoto 8ci. Rept. Tohoku Imp. Univ., (2), l^tih'olo.rodon naniadi-
CJeol., XIII, No. 1, pp. 4, 5. Type loc: Inland Sea, Japan. Type figs. cusyabei
Matsumoto, ibid., PI. in, fig. 2, PI. iv.

1929 /.o.r. C/\//.) 7V)A-//;mf/fn _////(/«;, niut. nov., Matsumoto Ibid., p. 10. Typeloe: .Iai)an, precis(> Polpeolo.rodon (lArehi-
localily unknown. Tyix' fig.: Matsumoto, ibid., PI. vii, figs. 1, 2. diskodon) tokunagai
nxni. junior

1929 Xotiomastodon or»o///s Cabrera Rev. Mus. La Plata, XXXII, p. 91. Tyi)e loc: Monte Her- XoliouKislodDii tn-natus
moso, Provinc(> of Buenos Aires, Argentina. Tvpe figs.: Cabrera, ibid., p. 91,
fig. 2; p. 93, fig. 4.

1929 .{irhidiskodon vanalpheni Dart So. iVfr. .lourn. Sci., XX\'I, p. 704. Type loc: Sydney-on-Vaal, Arehidiskodon vanal-
South Africa. Type figs. Dart, /b(W., ]). 704, : figs. 8, 9. pheni

1929 Areh id i.'<kiid<>u millctli Da,ri Ibid., i)|).

706-708. Typ<' loc: S}nlney-on-\'aal, Soulli .Vfiica. .\ichidi.-ikiid(in inillclli

Type figs.: Dart, ibid., p. 706, figs. 10, 11.

1929 Arehidiskodon loxodonloides Dart Ibid., pp. 709 711. TyP'' 'o<"-- Sydney-on-Vaal, South Arehiili.skodiin lo.rodon-
Africa. Type fig. : Dart, (^jd., p. 709, tig. 13. toides
 NOMENCLATURE OF THE PROBOSCIDEA 1415

Reference in
Year Name Author Present Memoir
1929 Archidiskodori andrewsi Dart Ibid., pp. 711-713. Type loc: Gong-Gong, Vaal River, South Palxoloxodon{'?) an-
Africa. Type fig.: Dart, ihid., p. 711, fig. 14. dreivsi

1929 Archidiskodon hanekomi DsLi-t Ibid., pp. 713-715. Type loc.: Delport's Hope, Vaal River, Palseoloxodon hanekomi
South Mriea. Type figs.:Dart, ibid., p. 713, fig.s. 15, 16.

1929 Archidiskodon yorki Dart Ibid., pp. 715, 717. Type loc: Vanasswegenshoek — Bloemheuvel, Archidiskodon yorki
near Christiana, South Africa. Type fig.: Dart, ibid., p. 717, fig. 19.

1929 Pilgrimia yorki Dart Ibid., pp. 719, 720. Type loc.: Below Christiana, Vaal River, South Palseoloxodon yorki
Africa. Type figs.: Dart, ibid., p. 719, figs. 20, 21.

1929 Pilgrimia imlmani Dart Ibid., pp. 720-722. Type loc: Below Christiana, Vaal River, South Palseoloxodon wilmani
Africa. Type fig.:Dart, «6irf., p. 721, fig. 22.

1929 Pilgrimia kuhni Dart Ibid., pp. 723, 724. Type loc: Pniel Estate, South Africa. Type fig.: Palseoloxodon kuhni
Dart, ibid., p. 725, fig. 24.

1929 Loxodonta prima Dart Ibid., pp. 724-726. Type loc: Pilandsberg, Transvaal, South Africa. Loxodonla prima
Type figs.: Dart, ibid., p. 725, figs. 25, 26.

1929 Loxodonta africana \ar. obliqua Dart Ibid., pp. 72&-72H. Type loc: Valley of Steelport River, Loxodonta ajricanavar.
tributary of Oliphants River, northeast Transvaal, South Africa. Type figs.: obli(/ua
Dart, ibid., p. 726, figs. 27, 28.

1930 Gomphoiheriitm'? emmonsi Hay

"Second Bib. and Cat. of Fossil Vertebrata of North America," Ocalientinu.i cnimonsi
II, p. 636.loc. Halifax County, North Carolina. Type figs.
Type : Emmons, :

1858, "Rept. North-Carolina Geol. Surv.," p. 199, fig. 23; Emmons, 1860,
"Manual of Geology," p. 218, fig. 186.

1930 Gomphotherium serpentirivale Hay Op. cit., p. 639. Nomen nudum

1930 Am.ebelodon si nclairi Barhonr Neb. State Mus., Bull. 17, I, pp. 155-158. Type loc: Freedom, Amebelodon sinclairi
Frontier County, Nebraska. Type fig. Barbour, tbirf., fig. 101. :

1930 Pliomastodon sellardsi Simpson Bull. Amer. Mus. Nat. Hist., LIX, Art. Ill, pp. 203-206. Type Pliomastodon sellardsi
loc: Brewster, Florida. Type figs. Simpson, i6irf.. p. 204, fig. 30; : p. 205,
fig. 31.

1930 Gomphotherium priestleyi Hay and Cook Proc Colo. Mus. Nat. Hist., IX, pp. 31, 32. Type Stegomastodon priesi-
loc: Near Frederick, Tillman County, Oklahoma. Type fig.: Hay and leyi
Cook, ibid., PI. xii, figs. 1, 2.

1930 Prodinotherium hungaricum tlhik Geol. Hungarica, Ser. Palaeont., Fasc. 6, pp. 1-14. Type loc: Deinotheriumhungari-
Kotyhaza (Dep. Nograd), Hungary. Type figs.: fihik, ibid., PI. i, figs. 1-3; cum
Pis. ii-iv.

1930 Pliomastodon vexillarius Matthew Bull. Dept. Geol. Univ. Calif., XIX, No. 16, pp. 335-341. Pliomastodon vexil-
Type loc: Southeast of Coalinga, Fresno County, California. Type figs.: larius
Matthew, ibid., PI. xli-xliv, text figs. 1, 2.

1931 Bunolophodon postremus Spillmann "Die Saugetiere Ecuadors im Wandel der Zeit," ErstcrTeil, Cuvieronius postremus
pp. 73-107. Type loc: Quebrada [canyon] of Cachihuayco, near Alangasi,
Province of Pichincha, east of Quito, Ecuador. Type figs. Spillmann, 1929, :

Natur und Museum, LIX, Heft II, text figs. 3, 4, and as.sociated skeleton,
fig. 1.

Material fir.st described in "El Ecuador Comcrcial," 1928, Aiio VI, No. 57, p. 7-3, as
"Mastodonte de Alangasi."

1931 Teleobunomaslodon Rcvilliod, 1931, bolivianus Philippi, 1893. Revilliod, Mem. Soc Paleont. Cordillerion {'i)bolivi-
Suisse, LI, pp. 1-21. Type loc: Exact locality unknown, probably from the anus
vicinity of Calacoto, valley of the Rio Desaguadero, Bolivia, south of Corocoro,
or from Concordia. Type figs. Revilliod, ibid., Pis. i, ii, and fig. 3, p. 9.
:
 :

1416 OSBORN: THE PROBOSCIDEA

Reference in
Year Name Author Present Memoir

1931 Torynobclodon banuonhniinu Har\nmy Neb. Statf Mus., l^ull. 22, I, pp. 191-198. Type loc: Toripiohilodini bnniiim-
Siiakc River, Cherry Couiitv, soutluvest of N'aleiitiiie, Nebraska. Type figs.: bivwiii
Barbour, ibid., figs. 123, 124.

1931 Mastodon moodiei Barbour Neb. State Mus., Bull. 24, I, jip. 203-210. Type loc.: West Blue Madodnn moodiei
River, southwest of Milford, Seward County, Nebraska. Type figs.: Barbour,
ibid., figs. 130, 131, 1326.

1931 Parelephas protomammonteus (Matsumoto) matsumotoi Saheki Japanese Journ. Geol. and Geog., [Not determined by the
VIII, No. 3, pp. 125-129. Type loe.: Mishima, Kimitsu district, Chiija present author]
Prefecture, Province of Kazusa, Japan. Type fig.: Saheki, ibid., PI. xv, figs.
1-3.
[Parekphn.t protomammonlpus matsumoloi Saheki, 1931, is invalitl, since matsumoloi in
this case is a liomonym of malsumotin Dietrich, 1927 (sec above, this list, under E. [Elephas]

primigenimt Matsutiwtoi Dietrich, 1927). If the material described by Saheki is di.stinct, as

lie thought it was, it is material at present without a name. — Colbert-Simi)son, letter, July
12, 1939.— Editor.)

1931 Pal^oloxodon antiquus italicus Osborn Amer. Mus. Novitates, No. 460, pp. 1-24. Tyjje loc: Hesperoloxodon antiqu-
Pignataro Interamna, near Cassino, Italy. Type figs. O.sborn, /6/rf., figs. : us itnlicus
1, 4-16.

1931 I'ala-olo.rodon aidiquii.'t {aiidreir.'ii'!)Oshoru Ibid., \)\). 1, 19, 21, 23. Type loc: Upnor on tiic [Hcsperdlo.rodon anliqii-
banks of theMedway, Kent, England. Type fig.: Osborn, ibid., p. 21, fig. iis (fide Simpson, sec

15; also Andrews and Cooper, 1928, "On a Specimen of Elephas antiquus p. 1222 above)]
from Upnor," Pis. i-viii, text figs. 1-5, and figs. 1079-1082 and 1084 of the
present Memoir.

1932 Pilgrimia archidiskodontoides Haughton Trans. Roy. Soc So. Africa, XXI, Pt. I, i>p. 4-8. Type Paholit.rodnii (irchi-
loc: Sydney-on-Vaal Breakwater, bed of Vaal River, South Africa. Type diskodonloides
figs.: Haughton, ibid., Pis. i-iii.

1932 Pilgrimia subantiqua Haughton Ibid., pp. 8-10. Type loc: Delport's Hope, half a mile from Lo.rodoida siibanliqua
Vaal River, South Africa. Type fig. Haughton, ibid., PI. iv, figs. 1, 2.
:

1932 SerriderUinus gobiensis Osborn and Granger Amer. Mus. Novitates, No. 537, pj). 11-13. Type Serridenl.inii.'i gobiensis
loc About forty miles southeast of Iren Dabasu, Inner Mongolia. Type fig.
:

O.sborn and (iranger, ibid., p. 12, fig. 8.

1932 Archidiskodon meridionalis tiebrascensis Osborn Proc Colo. Mus. Nat. Hist., XI, No. 1, pp. 1-3. Arrhidiskodon meridi-
Type loc: Near Angus, Nuckolls County, Nebraska. Type figs.: Osborn, onalis nebni.sccnsis
ibid., figs. 1, 2.

1932 Trilophodon cooperi Osborn Amer. Mus. Novitates, No. 585, pp. 1-6. Type loc: Dera Bugti, Trilophndon cooperi
Baluchistan. Type fig.: O.sborn, ibid., p. 2, fig. 1; paratype, p. 3, fig. 2
(described by Forster Cooper, 1922, Proc. Zool. Soc. London, p. 610, as
referal)le to Bunolophodon angustidens).

1932 Telralophodon bumiajuensis van der Maiu'cl "Contribution to thi> Knowledge of the l<"ossil Mam- Trtralophodoii biimia-
malian Fauna of Java," pp. 2, 3, 108-121. Tyjjc loc: Humiaju, central jiten.sis

Java. Type figs.: van der Maarel, op. cii.. Pis. viii, ix, x, and text figs. 17-21.

1932 Stegodon bondolensis vnn der Ma,a,rv\ Op. n7., pp. 158-164. Type loc: Bondol near Kuwung, Slegodon bondoknsis
District Randublatimg, R(>gency Blora, Residency Rembang, Java. Type
figs. : van der Maarel, «/). ril., PI. xiv, figs. 1, 4, 5, and text figs. 24, 25.

1933 Blirkolhcrium blicki Frick Bull. Amer. Mus. Nat. Hist., LIX, Art. IX, pp. 509, 515, 527-531. Blickolhciiam blicki
Type loc: Near Tapasuma, Gracios, Honduras. Type figs.: Yr'wk, ibid.,
figs. 3, 4.

1933 Aybelodon hondurensis Frick Ibid., pp. 527, 528, 532. Type loc: Near Tapasuma, Gracios, Hon- Ai/belodon hondurensis
duras. Tyjje figs.: Frick, ibid., figs. 5, 13, 18.
 NOMENCLATURE OF THE PROBOSCIDEA 1417

Reference in
Year Name Author Present Memoir
1933 Serruleidiniis filholi Vr'ick /6/V/., pp. 509, 535. Typcloc: Gors, Franco. Serridentinii.^ filholi
The mandible (Mas. d'Hist. Nat. Paris .A.C 20.J8, 2062) of t hi.s species was refi rred to in
the article by Frick, 1926, Bull. .\mer. Mus. Nat. Hi.st., LVI, Art. II, pp. 177, 178, but the
name Serridentinus filholi was not as.signed until 1933.

1933 Trilophnclon rruziensis Frick 76;V/., pp. 505, 579. Typeloc: Santa Cruz, New Mexico. Type Megabelodon cruziensis
fig.s.: Frick, ibuL, hg^. 10, 12A, 17, 23A, 25.

1933 Ocalienlinus ojocaliensis Frick pp. 509, 576, 579.

/?)/>/., Typeloc: Ojo Caliente, New Mexico. Ocalientinus ojocalien-
Type figs.: Frick, ibid., fig.s. 7, 18. sis

1933 Trobelodon taoensis Frick Ibid., pp. 505, 580. Type loc. : Santa Cruz, New Mexico. Type figs. : Trobelodon taoensis
Frick, ibid., figs. 2, 13, 18.

1933 Tatabelodan riogranden.ns Frick Ibid., pp. 505, 581. Typo loc: Battleship Mountain, New Trilophodon {Tatabelo-
Mexico. Typo figs.: Frick, ibid., figs. 6, 13. don) riograndensis

1933 {'!)Amebelodon joraki Frick Ibid., pp. 505, 582. Type loc: Santa Cruz, New Mexico. Type Megabelodon joraki
fig.: Frick, ibid., fig. 18.
This type (a tuskless mandible) was first referred by the present author (Vol. I, p. 326,
this Memoir) to the genus Trilophodon, but finally in the Appendix to Volume I, pp. 707,
738, he definitely assigned it to Megabelodon.

1933 Serbelodon barbourensis Frick /fe«d., pp. 506, 592, 594. Typeloc: Christmas quarry, near Ain.s- Serbelodon barbourensis
worth, Nebraska. Type figs. Frick, /6/r/., figs. 14, 16, 27.
:

1933 Tatabelodon gregorii Frick Ibid., pp. 506, 597. Type loc: Vicinity of Ainsworth, Nebraska. Trilophodon {Tatabelo-
Type figs.: Frick, ibid., figs. 13, 27A. don) gregorii

1933 {?)Trilophodon barstonis Frick Ibid., pp. 506, 607. Typo loc: Mohave Desert, California. Serridentinus barstonis
Typo figs.: Frick, ibid., figs. 33, 36.

1933 Mastodon raki Frick Ibid., pp. 506, 630. Typo loc: Hot Springs, Now Mexico. Typo figs.: Mastodon rnki
Frick, ibid., figs. 25A, 29 A.

1933 Mastodon americanus alaskensis Frick Ibid., pp. 506, 631. Typo loc: Vicinity of Fairbanks, Mastodon americanus
Alaska. Typo fig. : Frick, f6;rf., fig. 29A. alaskensis

1933 Elephas primigenius alaskensis Osborn (In Frick, ibid., pp. 631, 632.) Type loc: Vicinity of Mammonteus primi-
Fairbanks, Alaska. Type figs. Osborn, this Memoir, Vol. II, figs. 1025, 1026.
: genius alaskensis

1933 Serbelodon burnhanii Oshorn Amor. Mus. Novitates, No. 639, pp. 1-5. Typo loc: Near Serbelodon bur nhami
Ricardo, San Bernardino County, California. Type figs.: Osborn, ibid., figs.
1,2.

1933 Stegodon trigonocephalus praecursor von Koonigswald Wetenschappelijke Mededoelingon, Dienst Stegodon trigonoceph-
Mijnbouw Nederl. -Indie, I Toil, No. 23, pp. 104, 105. Type loc: Bumiaju, alus praecursor
Java. Type fig. von Koonigswald, ibid., Taf. xxvii, fig. 2.
:

1933 Cryptomastodon martini yon Koenigswakl Ibid., pp. 111-119, Taf. xxviii, figs. 1-3, and text ?Sirenian
figs. 8, 9.

1934 D. [Dinotherium] Bozasi Arambourg Compt. Rend. Soc. g6ol. France, No. 6, pp. 86, 87. [Not determined by the
Type loc: Valley of the Omo, Abys.sinia. Type fig.: Arambourg, 1935, present author]
Bull. Soc. g^ol. France, (5), IV, PI. xviii (mandible). Supplementary
description: Arambourg, op. cii., pp. 305-310.

1934 Mastodon grangeri Yiarhonr Neb. State Mus., Bull. 35, I, pp. 287-290. Type loc: Pender, .^^Mastodon grangeri
Thurston County, Nebraska. Type fig.: Barboiu-, /6k/., p. 289, fig. 170.

1934 Palseoloxodon yokohamanus Tokunaga. Journ. Goog. (Tokyo), XLVI, No. 546, pp. 363-371 (in [Not determined by the
Japanese). Typeloc: Mouth of Tsurumi-gawa, Yokohama, Japan. Type present author]
fig.: Tokunaga, ibid., PI. viii, figs. 1, 2. r
 —

1418 OSBORN: THE PROBOSCIDEA

Reference in
Year Name Author Present Memoir
1934 Pnranlegodoii? kifniitoenm's Toknna^a Ibid., p|). 365-369 (in Japanoso). Typo loc: Kakio, (Not dctcmiiiicd by tho
Kanagawa Picl'ccturc, Japan. Type fig.: Tokiiiiaga, ibid., PI. ix, figs. 1-3. pro.iont aiitlioi']

1934 Archidixkodori proplnnifrons Osborn Amor. Mas. Novitates, No. 741, pp. 10-12. Type loc: Airhidixkodon prn-
(long-CJong, near tlic Vaal Rivor, South Africa. Typo fig.: Osborn, ibid., p. planifron.s
5, fig. 2.

1935 Palseoloxodon ])risru.^ var. bosei Proc. 22nd Indian Soi. Congross, Calcutta, ]>. 209.
Chakravarti [Not dotomiined by the
Typo loc: Parkalta near Jamniu, India. Typo fig.: Bo.so, 1929, Quarl. present autlior]
Journ. Gool. Min. and Motallurg. Soc India, II, No. 3, PI. v, fig.10, as Steg<i-
don bomhifrona. Supplementary description: Chakravarti, 1937, Quart.
Jouni. Cool. jMin. and Motallurg. Soc India, IX, No. 2, j)]). 39 42, PI. vi
(figvH'od as Palseoloxodon sp.).

1935 (Itmlhabelodon thorpei Barbour and Sternberg Neb. State Mus., Bull. 42, I, pp. 395-404. Typo Gnathabelodon thorpei
loc: Near Ogallah, Trego County, western Kansas. Typo figs. Barbour and :

Sternberg, ibid., figs. 187-191.

1935 Stegodon yiit:hensis Young Pal. Sinica, (C), IX, Fasc. 2, pp. 26-28. Type loc: Yushc, China. [Not determined by
Type fig.: Young, ibid., PI. v, figs. 1, la. the present author]

1935 Stegodon zdanskyi Hopwood (In Young, ibid., p. 28);Hopwood, Pal. Sinica, (C), IX, Fasc. 3, Slegodon zdanskyi
p. 75. Type loc: Unknown. Type fig. Hopwood, ?6k/.,
: PI. vii, fig. 5.

1935 Slegodon offidnalis Hopwood (In Young, ibid., Fasc. 2, pp. 27, 30.) Hopwood, ibid., Fa.sc. 3, p. 73. Slegodon officinalis
Typo loc: (?)Szechuan, China. Type fig. : Hopwood, ('fc/rf., PI. vii. fig. 3.

1935 T rilnphodon connexusUopvf ood Ibid., p. 14. Type loc: Kansu, Sining Fu, SW 20 li, Shui Trilophodon connexus
Ch'iian, P'u, SE 5 li, Tiao Kou, China. Tvpo fig.: Hopwood, ibid., PI. v,
figs. 1,2.

1935 Trilophodon wimani Hopwood Ibid., p. 19. Type loc: Pa P'an Shan, China. Type fig.: Serridenlinus wimani
Hopwood, ibid., PI. v, fig. 3.

1935 Trilophodon spectabilis Hopwood Ibid., p. 30. Type loc: Said to have come from Sian, China. Trilophodon spectabilis
Type fig.: Hopwood, ibid., PI. vi, fig. 2.

1935 Tetralophodon exoletus Hopwood Ibid., p. 35. Type loc: Shansi, China. Type fig. : Hopwood, Telralophodon exolelus
ibid., PI. VI, fig. 3.

1935 Pentalophodon sinensis Hopwood Ibid., p. 57. Type loc: Yii She Hsien, Shansi, China. Type A nanc as sinensis
fig.: Hopwood, ibid., PI. vii, fig. 2.

1935 Parastegodon mgiyaimii Tokuniiga Proc. Imp. Acad. Tokyo, XI, p. 434. Type loc: Iruhi, in [Stegodon{?) sugiyarrmi]
Saida village, Shikoku, Japan. Type fig. Tokunaga, : i6/rf., p. 433, text fig.

[Not determined by the present author, but regarded by Dr. E. H. Colbert as referable
to Slegodon rather than to Paraslegodon (which was considered by Professor Osborn as
possibly c(iual to Archidiskodon Pohlif? or to a progre.ssive Slegodon). Consequently this
species is described in the Slegodon chapter, pp. 899 and 900 above. Editor.] —
1936 Mastodon alavus Borissiak Travaux do I'Institut Paleozoologiciuo do 1' Academic des Sciences do [Not determined by the
rU.S.S.R., V, pp. 171-234. Tyj)o loc: Dschilantschik River, Turgai, Russia. present author]
Type figs.: Borissiak, ibid., Taf. i v, viii, and Taf. vi, vii (in part), also text
figs. 1-16. Pal. In.st. No. 2280, Leningrad.

1936 PlioTna.stodon nevadanii.^'Atock Publ. Carnegie Instn. Wash., No. 473, p. 37. Type loc: Thous- [Not determined by the
and Crook atxmt four miles northwest of the Hot Spring and on oast side
ba.sin, [jre.seut author]
of Railroad Ridge, Humboldt County, northwostorn Nevada. Ty])o fig.:
Stock, /6/f/., PI. (incomplete skull representing most of the palate, second and
I

third superior molars of each side, and a complete right tusk ('alif. Inst.
Toch. Coll. Vert. Pal. No. 1922).
 . .

NOMENCLATURE OF THE PROBOSCIDEA 1419

Reference in
Year Name Author Present Memoir
1936 raJ:rohi.ro(l(>ii (Kinioripiifn's T()\n\]\Ag,A No. 564, Fcliniary, pp.
.lourn. (loop;. (Tokyo), XLVHI, [Not dcHcrmiiiod by tlic
67-70 (in .l:ii)tine.se). Typcloc:
on the Sliichinolie-gavva, Toiijinbayaslii, iircsent author]
near entrance to town of Sluchinoho, Kamikita-gun, Aomori Prefecture,
Japan. Type fig.: Tokunaga, ibid., PI. i. Supplementaiy description:
Tokunaga and Takai, 1936, .Journ. Geol. Soc. Japan, XLIII, No. 511, April
20, pp. 254-258 (in English).

1936 Deinotherium hopivoodi Oshorn This Memoir, I, p. 117. Type loc: Olduvai, near southeast Deinolheriuin hopwoodi
shore of Lake Victoria, Tanganyika Territory, Africa. Type fig.: Osborn,
op. cit., p. 104, fig. 68a.
Ri'gartlcd by Doctor Hopwood a.s a synonym of IHnolheriitin lio'asi AramfjoiirK, 103^.
Sep abovf, thi.s li.st

1936 Triluphodon hasnotensis Oahorn Op. cit., p. 279. Type loc: Near Hasnot, India. Type fig.: Trihphodon hasnoten-
Osborn, op. cit., p. 454, fig. 417. .si'.s

1936 Tetralaphodon fricki (Miovn Op. cit., p. 375. Type loc: Near Clarendon, northern Texas. Tetndophodonfricki
Type figs.: Frick, 1933, Bull. Amer. Mus. Nat. Hist., LIX, Art. IX, fig. 12
(lower) and fig. 23B (upper).

1936 Rhyncholherium broumi Osborn Op. cit., p. 494. Type loc: San Jose de Pimas, Sonora, Mexico. Rhynchotherium browni
Type O.sboin, 1921, Amer. Mus. Novitates, No. 1, fig. 2C, as neotype of
fig. :

Rhynchotherium tlascalx. See also this Memoir, I, fig. 467.

On locating tlie type cast of Rhynchotherium tlascalae in tlw. Geneva Mu.seum, tlic neo-
type mandible was found to be quite different; con.sequently the present author made it

the type of a new species, Rhynchotherium browni.

1936 Cordillerion edensis Osborn This Memoir, I, p. 560. Type loc: Mt. Eden Hot Springs, San Cordillerion edensis
Bernardino County, California. Type fig.: Frick, 1921, Bull. Dept. Geol.
LTniv. Calif., XII, No. 5, PI. l (as Trilophodon shepardi edensis); Osborn,
1922, Amer. Mus. Novitates, No. 49, fig. 1, Al, A2 (as Dibelodon edensis).
See also this Memoir, I, figs. 522, 523.
Cf. notes under Trilophodon (Tetrahelodon) shepardi edensis Frick, 1921, and Dibelodon
edensis Osborn, 1922, above, this list.

1936 Anancus properimensis OshoTX). This Memoir, I, p. 647. Type loc: Near Chinji Bungalow, Anancus properimensis
India. Type figs. Osborn, op. cit., figs. 609, 613.
:

1936 P entalophodon falconeri Oshom Op. cit., p. 653. Type loc: Siwalik Hills, India. Type figs. : Pentalophodon falconeri
Falconer and Cautley, 1847, "Fauna Antiqua Sivalensis," Pis. xxxii, and
XXXIII, figs. 1, 2 (as Mastodon sivalensi.'s).

1936 Miomastodon depereti Oshoni Op. cit., p. 693. Type loc: Chevilly, France. Type fig.: Mayet, Miomastodon depereti
1908, Ann. Univ. Lyon, Nouv. Ser., I, Sci., M6d., Fasc. 24, PI. vii, fig. 3 (as
Mastodon angustidens)

1936 Mastodon pavlowi Osborn Op. ct7., p. 694. Type loc: Pestchana, Podolia, Russia. Type fig. : Mastodon pavlowi
Pavlow, 1894, Mem. Acad. Imp. Sci. St. Petersb., (8), I, No. 3, PI. i, figs. 1,
2, 3 (as Mastodon ohioticus).

1936 Mastodon acutidens Osborn Op. c/f., p. 696. Type loc. : Rochester, Indiana. Type figs. : Osborn, Mastodon acutidens
op. cit., figs. 131, A1-A4, 135, 656, and PI. i, L.

1936 Stegolophodon lydekkeri Osborn Op.Type loc: Vicinity of Bruni, northwest coast
cit., p. 700. Stegolophodon lydekkeri
of Borneo. Lydekker, 1885, Proc Zool. Soc. London, PI. xlviii
Type fig.:
(as Mastodon latidens); Lydekker, 1886, "Cat. Foss. Mamm. Brit. Mus.,"
fig. 19 (as M. latidens).

1936 Stegomastodon primitivus Osborn Op. cit., p. 72Q. Type loc: Northeast of Ainsworth, Nebraska. Stegomastodon primiti-
Typefigs. Osborn, op. «'<., figs. 674, 675.
: vus
1420 OSBORN: THE PROBOSCIDEA
Reference in
Year Name Author Present Memoir
19.SG J '(im.'ilegudon akashiensis Takai Proc. Imi). Acad. Tokyo, XII, No. 1, pp. 19 21. Tyi)o loc: [Not lietermincd by the
"Shore of the cliffy coast," west of Ni.'^liiyagi, ()kiil)()-nuira, Akashi-gun, present author]
Hyogo Prefecture, Japan. Type figs. Takai, ibid., p. 20, figs. 1, 2.
:

[Both Mr. Fuyuji Takai and Mr. Tokio Shikama .simultaneously studied the same fossil

material from the Akasi District, neither being aware of the deseription of the other. Mr.
Takai chose a molar as his tyjM', naming the spfU'ies Parastegudun akashiensis, while Mr.
Shikama extended his studies on this material, in<-luding two skulls with molars, a lower
jaw with molars, and some other isolated teeth from the same locality, choosing as his type
one of the skulls, and naming the species Paraslegodon nipponicus in his manuscript.
Meanwhile Mr. Takai had announced his species in a lecture delivered at the meeting of tlie
Palffontological Society of Japan, November 30, 1935. While this announcement did not
constitute priority of description, Mr. Shikama very generously withdrew his unpublished
manuscript name in favor of Mr. Takai's species name PnriiMegodon nknshiensis. (Cf.
footnote by H. Yabe, in Takai, 1936, *;(/., p. 19).— Editor.]

1937 Peritalophodon runeatus Teilhard de Chardin and Trassaert Pal. Sinica, (C), XIII, Fasc. 1, p. 11. [Not determined by the
Type loc: Southeastern Shansi (Yushe Basin), China. Type fig. Teilhard : [iresent author]
de Chardin and Trassaert, ibid., PI. in, fig. 4.

1937 Mastudun i/iYe;vnerfiu.s Teilhard de Chardin and Tras.saert Ibid., p. 22. Type loc: Southeastern [Not determined by the
Shansi (Yushe Basin), China. Type fig.: Teilhard de Chardin and Tras- present author]
.saert ibid., PI. iii, fig. 2 a-c.
,

1937 Siegiodo/i /;'ce«/i' Teilhard de Chardin and Trassaert Ibid., p. 27. Type loc: Southeastern [Not determined by the
Shansi (Yushe Basin), China. Type figs.: Teilhard de Chardin and Tra.s- present author]
saert, ibid., PI. viii, figs, la, lb, 2, and text fig. 3.

1937 Paraslegodon infrequens Shikama, Japanese Joiu-n. (!eol. Ceogr., XIV, pp. 127-131. Type loc: [Not determined by the
Near Akasi (precise locality luiknown). Type fig.: Shikama, ibid., PI. ix. pre.sent author|
Type: Anterior portion of left ramus with Ma in situ. Collection of Taki-
kawa Middle School in Kobe.

1937 Parelephas proximus uehataensis Shikama Ibid., p. 165. [Not determined by the
present author]
If the name Parelephas protomammonteus typicus is synonymous witli aniiquus, then
Shikama was justified in giving a new name, Parelephas proxtmus uehataensis, to inchidc
secondary specimens of P. protomammonteus typicus plus those of E. primigenius inalsu-
moloi.— Colbert-Simpson, letter, July 12, 1939. (See notes under Parelephas protomam-
monteus typicus Matsumoto, 1926, and E. [Eteplias] primigenius Matsumoloi Dietrich, 1927,
above, this list.)

1937 E. [Elrphn.^\ antiqmis mut. ruthenensis Astre Bull. Soc Hist. Nat. Toulou.se, LXXI, p. 30. [Not determined by the
Type loc: Salles-la-Source (Aveyron), France. Type fig.: Astre, ibid., PI. i pre.sent author]
(incomplete molar), figured as "Elephas aniiquus, de Salle.s-la-Source."

1938 Bunolophodon yokolii Makiyama Mem. Coll. Sci. Kyoto Imp. Univ. (B), XIV, No. 1, pp. 12-14. [Not determined by the
Type loc: Upper Banko Sandstone of the Mei.sen series, at Senkaibo in the pre.sent author]
Meisen di.strict, North Kankyo-do, Ja])an. Type figs.: Makiyama, ibid.,
text figs. ^i\ and 5b.

1938 Stegodon shodoensis akashiensis {Takai, 19'.i(i) Makiyama Ibid., pp. 21-27. Cotypeloc: Eiga- [Not determined by the
sima near Aka.'ii, and inid(>r the sea off Hayasi-zaki, Jai)an. Cotyjx' figs.: present author|
Makiyama, ibid., figs. 10 12.

1939 Palxoloxodon darli Cooke and Clark Trans. Roy. Soc So. Africa, XXVII, Pt. 3, pj). 296-302. [Not determined by tiie

Tyi)(> loc: Victoria Falls, northern Rhodesia. Tyi)e figs.: Cooke and Clark, present author]
ibid., Pis. XII and xiii.

1939 Archidiskodon pnrnmammonteus Matsumoto Zool. Mag. (Tokyo), LI, No. 10, p. 704 (in Japan- [Not determined by the
ese), )). 716 (lOnglish resume). Tyi)e loc Nagahama, Minato Town, Province
: present author]
of Kazusa, Japan. Type fig. Mat.simioto, /7»V/., fig. 3. Type: I'Yaginent of
:

molar.
 Chapter XXII

THE GEOLOGIC SUCCESSION OF THE PROBOSCIDEA

By Edwin H. Colbert

I. Introduction. V. Asia.

II. Africa. 1. Introduction.

1. Introduction. 2. The Miocene of Mongolia and Central Asia.

2. The Eocene and Oligocene of North Africa. 3. The Miocene of North China.
3. The Miocene of North Africa. 4. The Pliocene of Mongolia.
4. The Miocene of Central and East Africa. 5. The Pliocene of North China.
5. The Pleistocene of North Africa. 6. The Pleistocene of North China.
6. The Pleistocene of Central and East Africa. The Miocene to Pleistocene of Japan.
7. The Pleistocene of South Africa.
III. The Orient.
1. Introduction.
2. The Miocene of Baluchistan and Sind.
3. —
The Siwalik Series (Miocene Pleistocene) of North
India.
4. The Pleistocene of Central India.
5. The Pleistocene of Ceylon.
6. The Pleistocene of Burma.
The Pleistocene of South China.
The Pleistocene of Indo-China.
9. The Pleistocene of the East Indies (Java, Borneo,
Philippines).

IV. vr\.y

1.
1422 OSBORN: THE PROBOSCIDEA

scideans. Thus Ihad the opportunity to become familiar with Professor Osborn's views as to geologic ages and
correlations, and to learn, to some extent, the manner in which he planned to write and present this chapter on
the geologic succession of the fossil Proboscidea.

A detailed presentation of the geologic succession, such as Professor Osborn would have written, is not now
considered practicable. The original author is not here to write this chapter or to supervise its writing, and it is

not considered feasible to imitate his style

^EOIOQIC^ RELjiTlONSKlPS OF
or his method of presentation. Therefore
AFR-ICAN^ PROBOSCIDEA.
this chapter will be more or less in the
form of a running narrative, describing
the distribution, development, and corre-
lation of the various Proboscidea-bearing
horizons of the world. Charts and faunal
lists will be included only where it is

thought necessary to clarify the text.

In writing this chapter, an attempt

will be made to set forth the various view-
points —often very divergent —regarding
the geologic age and the relationships of
the horizons being considered. In all cases.
Professor Osborn's views will be stressed
(when known), and his opinions will be

compared with the opinions of other palae-

ontologists, including those of the present

author.
 GEOLOGIC SUCCESSION: AFRICA 1423

orders ofmammals, supporting the idea that these animals had a northern, Palaearctic origin, rather than an
Ethiopian origin as postulated by Professor Osborn.

North Africa, where the earliest known proboscideans are found, is at the present time in the Palaearctic
zoogeographic region. Consequently, upon the basis of the present-day distributionof mammals, the first probo-
scideans and the sediments in which they occur might logically be considered in connection with the discussion of
Eurasiatic horizons. It has been thought best, however, to include the whole of Africa in this one section, so that
the Eocene and Oligocene sediments of the Egyptian Fayum, containing the ancestral types of proboscideans, will

be taken up at this place.

2. THE EOCENE AND OLIGOCENE OF NORTH AFRICA

Fay^tm District, Egypt. —The Fayum District is a circular depression in the Libyan Desert, located to the
west of that portion of the Nile River between Kafr el Ayat and Feshn. Its central portion is occupied by a lake
known as Birket el Qurun —the remnants of a once much larger lake known as Moeris, which in the time of the
ancient Egyptian dynasties served as a reservoir that controlled the Nile irrigation system. A series of marine and
fluviatile deposits, ranging in age from the Middle Eocene through the Lower OUgocene, form the escarpments
around this depression ; these are the Fayum deposits in which several distinct and characteristic early Tertiary
faunas have been discovered.

The Fayum sediments, ranging from the lower to the upper beds, represent a succession of facies beginning
with marine sediments, passing through a series of mixed marine and fluviatile beds and finally developing at the
top into fluviatile deposits. As Andrews and Beadnell have shown, this gradual change in the character of the
Fayum deposits from the lower to the upper beds is to be attributed to an uplift of the Ethiopian region progress-
ing from the south to the north.

The succession of sediments in the Fayum District, and their correlation by different authors, is as follows.

Pliocene and Pleistocene and Recent

 :

1424 OSBORN: THE PROBOSCIDEA

strictly marine vertebrates —notably very primitive cetaceans among the mammals—while the sediments above
the Fluvio-marine series are virtually unfossiliferous so far as mammals are concerned.

In the original descriptions of the Fayum mammals by Andrews, the ages of the Qasr el Sagha and the
Fluvio-marine beds were established as Middle Eocene and Upper Eocene respectively. Subsequently several
authors, notably Stromer (1907), Osborn (1910) and Schlosser (1911) designated the Qasr el Sagha beds as of
Upper Eocene age, thereby shifting the Fluvio-marine beds up into the lowest part of the Oligocene period. These
differences of interpretation of the Fayum sediments may be attributed to divergent opinions as to the affinities

of the contained faunas.

Andrews'' views as to the age and affinities of the Fayum mammals were expressed in 1906 as follows:

—
The Mammals may bo divided into tliree sections: (1) the land-mammals which seem to be truly endemic to the Ethiopian
region; these occur both in the Upper and Middle Eocene beds, and include such genera as Mwrilherium, Pals-omadodon,
Arainoiiherium, Barj/therium, Megaluhyrax, Saghatheriiim, and perhaps Geniohyus: (2) forms of which close allies occur in
other regions in approximately contemporary deposits; these, so far as at present known, occur only in the Upper Eocene
beds, and include such genera as Ancodon, lihagatherimn, Hysfiiodon, Pterodon, Apterodon, and Sinopa: (3) the aquatic mam-
mals so far not found in the Upper Eocene beds, and comprising Eosrren, Zeughdon, and Prozeuglodon. It seems probable that
some of these last, like the genera included in section 1, are of endemic origin, having originated from land-mammals inhabiting
the region.

In 1907 Stromer- suggested that the fauna of the Fluvio-marine beds might properly be placed in the Oligo-
cene.
Nachdem nun jener ausgezcichnete Kcnner tertijirer Wirbelloser zu dem Schlusse kam (1906, S. 347), dass sie ganz oder
doch zum Toil dem Eartonien, also dem Obereociin entspreche, wijrde der unmittelbar auflagernden Fluviomarinstufe mit
ihren verkieselten Holzern und Rc])til- und Landsiiugetier-Resten (Andrews, 1906) unteroligoctines Alter zuzuschreiben sein.

In 1910, Osborn indicated the Fluvio-marine beds as of Lower Oligocene age, and this same interpretation
was followed by Schlosser in 1911, in his monograph on the "OUgozanen Landsaugetiere aus dem Fayum."

It may be well to review briefly at this place the evidence in favor of these differing interpretations as to the
age of the Qasr el Sagha and of the Fluvio-marine series.

Qasr el Sagha Series

The four genera of importance in the Qasr el Sagha beds are
Barytherium
Mceritherium
Eosiren
Zeuglodon

Of these the first two genera are autochthonous to northern Africa and therefore are of no value for correl-
ative purposes. The other two genera, Eosiren and Zeuglodon, are decidedly of Eocene age. It is difficult to say
whether these forms should be placed in the Middle or Upper Eocene, but comparisons with
in the related genera
in other parts of the world (particularly the wide-spread Zeuglodon) would seem to indicate that their affinities
might be with Upper Eocene forms.

Fluvio-marine Beds
In reviewing the Fluvio-marine fauna, it is well to remember Andrews' separation of the various nianunalian
genera into endemic African types and mammals common not only to Africa but to other parts of the world as
"Andrews, C. W., 1906. "A Descriptive Catalogue of the Tertiary Vertebrata of the FayOm, Egypt," pp. xii-xiii.
^Stromer, E., 1907. Abhandl. Senekenbergisehen Naturforsehenden Gesellschaft, XXIX, p. 144.
 GEOLOGIC SUCCESSION: AFRICA 1425

well. Schlosser, in 1911, lists nine orders of mammals from the Fluvio-marine deposits of the Fayum. Of these,
three — the Hyracoidea, Proboscidea, and Embrithopoda — were at that time probably indigenous to North
Africa, and hence should be excluded from any consideration of intercontinental correlations.

Of the remaining orders, the following genera are important. Their relationships, both zoologic and geologic,
are indicated.

Primates
Parapithecus Generally more advanced toward the anthropoid habitus than the Burmese genus,
Amphipithecus, which latter is of uppermost Eocene age. Therefore the evidence would
be in favor of an Oligocene age for the Fayum genus.
Propliopithecus Related to Pliopithecus. Propliopithecus is definitely so far advanced as an anthropoid
as to warrant its position in the Oligocene.

Insectivora
Metoldobotes Schlosser has compared this genus with the North American Oldobotes (Mixodectes) , of
Paleocene age. Insectivores are, however, notoriously persistent primitive forms in all

mammaHan faunas.

Creodonta
Ptolemaia Regarded by Schlosser as a specialized descendant of the Phosphorite genus, Cynohyasno-
don.
Metasinopa Closely related to "Sinopa" aethiopica of the Fajolm.
Sinopa An Eocene genus.
Apterodon Originally described from the Phosphorites of Quercy.
Pterodon An advanced type of hysenodont.
Hysenodon Typically an Oligocene genus.

RODENTIA
Phiomys Schlosser has compared these two genera with Theridomys of the Lower OHgocene, and
Metaphiomys Trechomys of the Upper Eocene of France.

Artiodactyla
Ancodon This is typically an Oligocene genus in Europe and America.
Brachyodus Schlosser referred to this genus the species that Andrews placed in Ancodon. Brachyodus
first appears in the OUgocene and continues into the Miocene.

Considering the possibilities offered by the above review of certain diagnostic genera, it would seem likely
that the fauna of the Fluvio-marine beds is of Lower OUgocene age. Of course this faima contains various Eocene
types, but these may very well be holdovers from an earlier period —a phenomenon that is very common in the
development of faunal assemblages. On the other hand, the presence in the fauna of numerous typically Oligocene
forms, particularly among the hysenodonts and the anthracotheres, would argue strongly for the OHgocene age of
the Fluvio-marine sediments. It is the presence of these newcomers that is important in determining the age of
a fauna.
1426 OSBORN: THE PROBOSCIDEA

Proboscideans:

Phiomia osborni Matsumoto

Phiomia xointoni (Andrews)
Phiomia minor (Andrews)
Phiomia serridens Andrews and Beadnell Fluvio-marine Beds
Palxomostodon parvus Andrews
Palxomostodon inlermedius Matsumoto Lower Oligocene
Palseomastodon beadnelli Andrews
Moeritherium trigodon Andrews
Mosritherium andrewsi Schlosser

Mceritherium lyonsi Andrews ( Qasr-el-Sagha Beds

Mcpritherium gracile Andrews <

Moeritherium ancestrale Petronievics I

Upper Eocene

3. THE MIOCENE OF NORTH AFRICA

IssERViLLE, North Africa. — In 1897 Charles Deperet^ published a short notice descriptive of a single
proboscidean tooth found near Isserville in North Africa. This tooth he regarded as representing a very small
race of Trilophodon angustidens, and he gave it a subspecific name, making its designation Mastodon angustidens
pygmxus. Comparing this specimen with the typical Miocene Trilophodon angustidens of Europe, Deperet came
to the conclusion that the African specimen represented a more primitive and an earlier race of the T. angustidens

stock. Consequently he considered its age as Cartennien, which in turn he regarded as repre.senting the beginning
of the Miocene.

Je terminerai en faisant remarquer I'interet qui s'attache a la constatation precise de la presence du genre Mastodonte en
Afrique al'^poque eartennienne, c'est-a-dire des le debut du Miocene, exactement comme en Europe, sans que, dans I'etat actuel
de nos connaissances, on puis.se dire de quelle region du globe proviennent ces premiers Proboscidiens, ni quelle forme animale
a pu leur donner naissance.

Subsequently Professor Osborn came to the conclusion that the tooth that Deperet had compared with
Trilophodon angustidens might be more properly placed in the genus Phiomia. Because of this apparent relation-
ship between the specimen under consideration and the advanced mastodonts of the Fayum, Osborn placed the
beds at Isserville as questionably of Upper Oligocene age, rather than as of Lower Miocene affinities.

Proboscidean: Phiomia pygmseus (Deperet).

MoGHARA Desert, Egypt. —The most abundant remains of Miocene proboscideans in Africa were discovered
in northern Egypt, in the desert between the Nile Delta and the boundary of Tripoli. These specimens were
described by Fourtau- in 1918 and 1920, in his publication "Contribution a I'Etude des Vertebras Miocenes de
I'Egypte."

The mammalian fauna of the Moghara deposits consists of certain genera of cetaceans, a brachypodine rhi-
noceros designated by Fourtau as" TeZeoceras," several species of anthracotheres referred to the genera Brachyodus
and Masritherium, two types of proboscideans, assigned by Professor Osborn to the genera Trilophodon and
Rhynchotherium, and two primates, Prohylobates and Dryopithecus (?). The complexion of this fauna is distinctly
Miocene. In addition to the mammals, the dei)osits at Moghara contain numerous and various crocodilians,
chelonians, and fish.

'Dcp6ret, C, 1897. Bull. Soc. grol. France, (3), XXV, pp. 520, 521.
^Fourtau, R., 1918. Ministry of Finance, Survey Dept., Cairo, p. 98.
 :

GEOLOGIC SUCCESSION: AFRICA 1427

It is quite evident that such a fauna represents a region in the vicinity of a shoreUne a strand crossed by-

rivers and bordered by the sea. This fact was recognized by Fourtau, who differentiated three categories of
mammals in the Moghara deposits.

1. Aquatic mammals. The cetaceans, of which two types are recognizable, one inhabiting the open
water and the other an estuarine form.
2. Semiaquatic mammals. The anthracotheres and the brachypodine rhinoceros. Living along the
rivers and in the marshes.
3. Land mammals. The proboscideans and the primates.

This fauna was compared by Fourtau with the Burdigalian, Lower Miocene, assemblages of Europe, par-
ticularly those of the Orleanais sands and the Eggenburg deposits (Fourtau, 1918, op. cit., p. 98)

"Quant a son age, les afRnites des vertebres nous amenent a synchroniser ce gisement avec ceux des sables
de rOrleanais, en France, et avec celui d'Eggenburg, en Autriche, et a en faire un depot datant du BurdigaUen."

This author differed from Haug, who placed the deposits under consideration in the Aquitanian —a division
that has been variously interpreted as either of Upper Oligocene or of Lower Miocene age.

The evidence of the mammalian fossils strongly supports Fourtau 's conclusion that the Moghara deposits
are of Lower Miocene age. The anthracotheres, Brachyodus and Masritherium, and the proboscideans, Trilopho-

don and Rhynchotherium, are particularly significant in fixing the age of the deposits as Lower Miocene.

Proboscideans:
Trilophodon angustidens libycus (Fourtau)
Rhynchotherium spencer i (Fourtau)

Osborn (Vol. I of this Memoir) designated Trilophodon angustidens libycus as of Lower (?) Miocene age, while
he placed Rhynchotherium spenceri in the Middle Miocene. However, Fourtau's description of the fauna from
Moghara would indicate that this is a single assemblage of mammals coming from one horizon. Consequently it

would seem likely that the age of both proboscideans is Lower Miocene, as incUcated by Fourtau.
Hopwood, in a personal communication to the present author, states that he has an unrecorded tooth of
Dinotherium from Moghara.

—
Smendou. Gervais (Zoologie et Paleontologie Francaises) described a single tooth which was found at
Smendou in the Province of Constantine in northern Africa, as a form closely related to if not identical with
Zygolophodon borsoni. This identification was revised by Deperet in 1897, who referred the specimen to Turicius
tapiroides, a change of considerable significance since the former species is of Upper PHocene age while the latter is

characteristic of the Miocene. Li speaking of the specimen, Deperet' made the following statement: "11 resulte
de cette description fort claire que la molaire du Smendou appartenait a un Mastodonte du groupe a molaires
tapiroides, et vraisemblablement au Mastodon turicensis Schinz ( = tapiroides Cuv.), espece repandue dans toute la
hauteur du Miocene europeen."

In discussing the beds in which this particular specimen was discovered, Deperet said that: "M. Ficheur dans
un important travail sur les terrains tertiaires de ce bassin {B. S. G. F., 3e ser., t. 22, p. 544), a ete amene a
considerer ces couches comme un equivalent lacustre du Miocene inferieur ou Cartennien."
Proboscidean: Turicius tapiroides (Cuvier).
'Deperet, C, 1897. Bull. Soc. gcol. France, (3), XXV, p. 518.
 .

1428 OSBORN: THE PROBOSCIDEA

Khenchela. —This deposit is probably correlative with certain other North African localities considered in
these pages, namely, those of Isserville and Smendou, which are of Lower Miocene age. A proboscidean tooth was
found at this locality, and Deperet' made the following statement concerning it: "Cette molaire, remarquable
par ses dimensions tres petites, ressemble, dit M. Gaudry, aux dents du Mastodon turicensis et provient sans doute
du terrain miocene."

Proboscidean : Turicius tapiroides (Cuvier).

Cherichera. — In Tunis, at Cherichera, are deposits that would seem to be somewhat later in age than the
beds exposed near Isserville and at Khenchela. Here was found a jaw referable to Trilophodon angiistidens,

typically a Middle Miocene species: "Enfin, dans le meme Memoire, M. Gaudry a decrit et figure . . . une belle

moitie de mandibule droite d'un Mastodonte d'un autre groupe, le Mastodon angustidens Cuv. aux molaires
pourvues de mamelons arrondis, non tapiroides; cette belle piece provient du Miocene moyen (probablement
Helvetien) du Cherichera, pres Kairouan (Tunisie)."^

Proboscidean : Trilophodon angustidens (Cuvier)

4. THE MIOCENE OF CENTRAL AND EAST AFRK^A

Several localities are known in Central and East Africa where Miocene proboscideans have been discovered.
These will be discussed briefly in the following pages.

Karungu, British East Africa. — In 1911 C. W. Andrews described a new species of Dinotherium that was
found on the east side of Lake Victoria.

In a subsequent paper (1914)^ on the fauna from Karungu, Andrews made the following remarks as to the
age of the deposits.

The general character of the fauna indicates that the age of the deposits is probably Lower Miocene (Biirdigalien) and
that it was contemporary with the faunas of the Sables de I'Orl^anais and of Moghara, and probably also with the recently-
discovered fauna of the Biigti Hills in British Baluchistan. In all these localities Anthracotheres of similar type apjicar as an
important constituent of the fauna; and, although at present the characteristic small form of Dinotherhim has not yet been
found at Moghara, ncxcrthcicss a primitive Tetrabelodon, closely similar to T. angudidens, wliich elsewhere accomi)anies the
Dinotherium, ha.s been found in that locality.

In discussing the Karungu fauna, Andrews notes particularly the presence of a peculiar hyracoid, probably
descended from the Eocene hyracoids of northern Africa, and a rodent seemingly a direct descendant of the
Fayuin genus, Phiomys. The mammalian fauna of Karungu is as follows.

Pseudaelurus africanus Andrews

Creodont(?)
Dinotherium hobleyi Andrews
Myohyrax oswaldi Andrews
Paraphiomys jngotti Andrews
Rhinoceros
Meiycops africanus Andrews
Tragulids

'Dcp<5rct, C, 1897. Bull. Soc. gC-ol. France, (3), XXV, pp. 518, .")19.
-Dcporet, C, 1897. Bull. Soc. gi'ol. Franco, XXV, .519.
(3), !>.

'Andrews, C. W., 1914. (^uart. Journ. Geol. Soc, London, LXX, p. 163.
 GEOLOGIC SUCCESSION: AFRICA 1429

Osborn (Volume I, p. Ill, of this Monograph) placed the Karungu deposits in the Lower Miocene. This
would seem to be as satisfactory a correlation as may be made at the present time.

Proboscidean: See fauna! list, above.

KoRU, Kenya Colony. —This deposit, the fossil fauna of which has been described by Hopwood, is very
important because of the presence in it of several primitive dryopithecoids. The Proboscidea are represented by
Dinotherium hobleyi, which leads Hopwood to think that these beds are closely related to and correlative with the
Karungu deposits —that is. Lower Miocene.

In 1933, Hopwood' described the fauna (exclusive of the higher primates) as follows: "The fauna consists of
two or three genera, which may prove to be lemuroids, Dinotherium hobleyi, three genera of Creodonts, an
amphicyonine carnivore, several Rodents, two Insectivores —one very close to Potamogale, — as well as small pigs
and ruminants."

Proboscidean : Dinotherium hobleyi Andrews.

Hopwood's detailed study of fossil vertebrates from Kenya is now in the course of preparation.

Lake Rudolph, East Africa. — Closely related to the fauna of Kenya is that recently reported by Arani-
bourg as coming from beds of Burdigahan age, along the western border of Lake Rudolph. This fauna was
discovered in tuffs interstratified between heavy basalts and other eruptive rocks of that vicinity. According to
Arambourg,- the constitution of the fauna is as follows:

Proboscidien.s Mastodon af. angustidens Cuv.

:

Perissodactyles Aceratherium? sp.

:

Hyracoide.s Pliohyrax nov. sp.

:

Artiodactyles : Brachyodus sp.

Listriodon nov. sp.

Snide indetennine, voisin de Palaeochcerus.
Dorcatherium nov. sp.
Antilope indeterminee.

Par sa composition, cette faune s'apparcnte ^troitement a cellos qui out ete roconnues deja en quclqucs points do I'Afrique
et attribuees an Miocene infericnr: celles de Moghara, dans lo d^soit Libyque, do Karungn pros dn Lac Victoria on Afriquo
orientalc ot du Namib, dans los Diamantemviiste do I'ancion Sud-Ouest africain allemand.

Proboscidean : See faunal list, above.

5. THE PLEISTOCENE OF NORTH AP^RICA

The Pleistocene is perhaps the most difficult period to summarize of any of the geological ages included in
this consideration of Proboscidea-bearing beds. Even in Europe and North America, where our knowledge of the

Pleistocene has been advanced to its greatest degree of perfection, the problems of correlation —particularly
where evidences of glaciation are missing — are extraordinarily difficult. Therefore it becomes evident that in the

more southerly portions and South American regions, correlations

of the world, particularly the African, Oriental,
within the Pleistocene must of necessity be at the present time very questionable, since they are based for the most
part on the development of river terraces or the expression of faunas, and in many cases there is no possibility of
connecting them with the comparable Eurasiatic and American stages. In this discussion of the Pleistocene of
Africa, the continent will be more or less arbitrarily divided into a North, a Middle, and a South section, and the

deposits in each of these sections will be in turn listed and discussed.

'Hopwood, A. T., 1933. Journ. Linnean Soc. Zool., London, XXXVIII, p. 437.

^Arambourg, C, 1933. C. R. Soc. gcol. Franco, No. 14, pp. 221 222. ,
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Algeria. — In a very succinct account of the Pleistocene of Algeria, Ronier (1928)' summarizes the work of
previous authors, notably Thomas, Pomel, Tournouer, Pallary, Joleaud, Boule, Stromer, Lamothe, Deperet, and
Solignac, by showing that in North Africa a division of the Pleistocene must of necessity be based on the sequence
of mammalian faunas, the sequence of cultures, and the development of shorelines and river terraces. As Romer
has remarked, the development of correlations by the use of the above methods is very apt to lead into a "vicious
circle," whereby each line of evidence depends upon and in turn supports every other line of evidence; conse-

quently the application of these several methods of correlative attack must needs be accomphshed with the great-
est of circumspection. When, however, such studies are carried out with a great deal of care, and when all modify-

ing factors have been accounted for, the following facts as to the Pleistocene sequence would seem to be evident.
Romer, following Deperet and Mayet, placed the base of the Pleistocene in North Africa above the Villa-
franchian and the St. Prestian and below the Lower Sicilian. In this he followed the general practice of European
palaeontologists. Recently (Matthew, 1929,^ Hopwood, 1935,* Colbert, 1937^) it has been argued that the Villa-
franchian is truly of Pleistocene age, for it contains certain immigrant types that may be taken as diagnostic of
the beginning of the Pleistocene. These are particularly the true elephants of the genus Archidiskodon and the
true horse of the genus Equus. If this correlation holds good, then the Pleistocene in North Africa would open
with the negative movement of the Villafranchian, followed by the establishment of the high 150 meter terraces,
and this in turn followed by the Lower Sicilian negative movement.
The Pleistocene succession of Algeria is characterized by some very interesting faunal developments, of
which brief notice may be taken here.
The close of the PUocene in the North African region was a period of warmth, with a rather humid climate.
The fauna was distinctly African in its character, and it contained such typically Pliocene genera as Machairodus,
Hipparion, a mastodont, a primitive hippopotamus, H. hipponensis, and a gigantic giraffid, Libytheriuni. Accord-
ing to Romer, Equus stenonis was present at this time —a slightly earlier occurrence of this horse than its more
characteristic Villafranchian appearance. It is to be noted in this connection that Equus stenonis may not belong
to the genus Equus, but rather may show relationships with the Upper Pliocene American equid, Plesippus.
The beginning of the Pleistocene, here regarded as the advent of the Villafranchian, was marked by the
continuation of the warm, moist climate of the Upper PUocene, and concomitantly by the continuation of many
characteristic Upper PUocene mammals. In this period, however, certain new and immigrant forms appeared;
these were the true elephants, particularly Archidiskodon planifrons, the true Equus, the modern type of Hip-
popotamus, and a varied array of bovids — antelope and cattle. These immigrant types that set the ViUafranchian
apart as the opening of a new era in mammalian history continued for the most part to the end of the Pleistocene
and on into recent times, while the holdover PUocene forms that give to the ViUafranchian its rather PUocene ap-
pearance died out sooner or later in the Pleistocene. Yet some of these characteristic PUocene mammals, par-
ticularly Hipparion, persisted well into the Pleistocene in Algeria —a teUing demonstration of the fact that
mammals are prone to continue beyond the period of their "typical" expression in geologic time. So it is that new
and immigrant types are much more reUable as diagnostic time markers in the study of faunal successions than are
the "characteristic" or "typical" forms, which may persist from one period to the next.
The development of the Pleistocene in Algeria was marked by a long period of warm, to hot, humid cUmates,
beginning in the Pliocene and lasting to the opening of Upper Pleistocene times. During a portion of the Upper
'Romer, A. S., 1928. Bull. Logan Miis., I, No. 2.
-Matthew, W. D., 1929. Bull. Aracr. Mus. Nat. Hist., LVI, pp. 437 -.")00.
'Hopwood, A. T., 1935. Proc. Geol. Assoc, XLVI, Pt. I, p. Hi.
"Colbert, E. H., 1937. "The Pleistocene Mammals of North America and Their Relations to Eurasian Forms," in Early Man as Depicted by Leading
.\uthorities at the International Symposium The Academy of Natural Sciences Philadelphia, March, 1937. 8vo, J. B. Lippincott Company, Philadelphia,
pp. 173-184.
1432 OSBORN: THE PROBOSCIDEA

Pleistocene, the Monastirian, there would seem to have been an interval of relatively cold, but humid weather,
which in turn was followed by a rise in temperature and a final desiccation of the entire North African region.

At the beginning of the long warm, humid period, the fauna of North Africa was quite similar to that of

Europe, due in large part to the northward extension of many But during the de-
Ethiopian types of mammals.
velopment of the Pleistocene there were difTerential changes in the relationship between the North African,
Eurasiatic, and Ethiopian faunas, due to the "de- Africanization" of the European fauna and the "Europeanization"
of the North African fauna. In other words, there was a succession of faunal movements with results somewhat
as follows.

1. Similarity between North African and European faunas, due to extension northwardly of African

types of mammals. Africanization of Europe.

2. ^^'ithdrawal of African forms from Europe, causing a dissimilarity between this region and North
Africa. De-Africanization of Europe.

3. European types into North Africa again establishing a similarity between the North
Influx of
African and European faunas, but causing the former to be different from the lower African fauna or faunas.
Europeanization of North Africa.

This history would seem to have been complicated in its end stages by the fluctuations in the desiccation of

North Africa. According to Romer, there was a period of aridity following the Mousterian, causing the more
tropical elements in the North African fauna to migrate southwardly. This period was followed by a humid
period during the Neolithic, at which time these warmth loving animals reestablished themselves in the North
African region, where they continued, but in gradually diminishing strength, until recent times. The final, slow
diminution of the North African fauna has been due largely to the post-Pleistocene desiccation of the region,
which has been quite marked during historic times.

Proboscideans: See table, above.

6. THE PLEISTOCENE OF CENTRAL AND EAST AFRICA

Fossiliferous manunal localities are known in Central Africa from Kenya (^olony, Uganda, Tanganyika
Territory, Nyasaland, and northern Rhodesia. The occurrences and geologic ages of these Central African

faunas have been excellently summarized by Hopwood (1929).

Hopwootl lias shown that the fossil mannnals of Central Africa are, for the most part, innnigrant types that

have migrated into this region from the north and from the east. Here, protected in something of an ecological
"backwash," there has been a tendency for primitive forms to persist to relatively late geologic dates, thereby
giving to the African faunas a somewhat less advanced aspect than is consistent with their real ages. In this
resj^ect, there is a close similarity between the fossil faunas of Africa and India, in which latter region there is

a tendency for the fossil mammals to be homotaxially related to assemblages of earlier ages in other portions of the

world. And it is an interesting fact that India, like Africa, was more or less separated from the rest of the world

during late Tertiary times — the one by the high Himalaya Mountains, the other by the developing Mediterranean
Sea and the expansive waste of northern Afri(;a.

Of the Pleistocene faunas most important and the most comjilete is the Oldoway
of Central Africa, by far the

or Olduvai fauna. This assemblage^ Hopwood considers to be no okler than Middle Pleistocene^ in age. The other
Pleistocene faunas of Central Africa are those of Lake Rudoli)h, Karungu, Homa Mountain, Kaiso, and Lake
Nyasa — all related to the Oldoway fauna and all essentially equivalent to it in age.
 r.EOLOGIC SUCCESSION: AFRICA 1433

In the following pages the Oldoway fauna will be considered in some detail, while the other Pleistocene faunas
named above, since they are essentially similar to the Oldoway faima, will l^e treated more briefly.

—
Olduvai (Oldoway). The Olduvai (or Oldoway) locality in Tanganyika Territory, along the southeastern
shore of Lake Victoria, was first explored for fossil vertebrates in 1913, by an expedition under the leadership
of Dr. H. Reck, and more recently in 1931, by an expedition led by Dr. L. S. B. Leakey, who had as one of his
coworkers Dr. A. T. Hopwood. The principal contributors to the problem of the age of the Olduvai fauna have
been Dr. W. O. Dietrich, working on the basis of the material collected by Dr. Reck and his associates, and Dr.
Hopwood, basing his studies on his own personal observations of the region and on material that he collected there.

The deposits at Olduvai are of volcanic origin, being composed of fine volcanic tuffs that seemingly were
accumulated in shallow lake basins. Lithologically the Olduvai beds are divisible into five horizons or layers,
which have been described by Hopwood' as follows:

Exposed in the cliffs of the gorge are four main beds, lying on a thick flow of lava. The first, or bottom bed is known as
Bed I. It consists of silvery grey tuffs, with yellow pumiceous layers and concretionary bands. It is the thickest of the four,
with very little earthy contamination, and its deposition, although discontintious, was very rapid. To the east it is practically
barren of fossils, but to the west it yielded crocodiles, antelopes, elephants, Dinotherium, the three-toed horse (Htpparion), and
zebras.

BedII is mainly buff-coloured. It is more massive and earthy than Bed I, with which it is entirely conformable. At the
time itwas deposited, the rainfall was of varying intensity, and extra wet sea.sons are represented by beds of pebbles. The
fossils from this bed are much the same as those from Bed I, but we did not find Dinotherium, and in 1913 Professor Reck re-
covered a human skeleton from here.
Bed the most useful for stratigraphical purpo.ses owing to its bright red colour. It is conformable to Bed II, but dies
III is
out to the west. This suggests that the volcanoes were dormant, and their reddened decomposition products were washed into
the lake from the east The bed itself is hard and close grained, with occasional layers of pebbles and gravel.
.

Conformable to the last is Bed IV, a grey or brown bed, with a reddish phase in its western extension. It reflects in its
structure and composition the gradual drying up of the lake. In a pebbly layer near the top we found an incomplete skeleton
of a hippopotamus with over a hundred quart zite hand-axes lying on or among the bones.

Next comes a marked unconformity, followed by a complex series of deposits of wind-blown dust known collectively
as Bed V.

Recently (1933) Dietrich^ has published his conclusions as to the age of the beds at Olduvai. The sequence of
deposits, according to Dietrich, range throughout the extent of the Pleistocene: Horizon 1 is of Lower Pleistocene

age. Horizons 2, 3, and 4 are of Middle Pleistocene age, and Horizon 5 may be assigned to the Upper Pleistocene.
Dietrich bases his conclusions on the supposed restriction of the Pliocene elements, Dinotherium and Stylohipparion
to the lowest horizon, on the general Middle Pleistocene character of the bulk of the fauna, foimd in Horizons
2 to 4 inclusive, associated with Chellean and Acheulean industries, and on the supposed discordance between the
topmost horizon and the bed beneath it, together with the presence of Mousterian and Ainignacian cultures in
this uppermost layer.
Horizont 5 Steppenkalk ('Loss'). Moustier? Aurignac. Recente Fauna.
Diskordanz
Horizont 4 Graue Tuffite mit Gerollagen. Elephas antiquus recki, Hippo-
potamus gorgops, Pelorovis oldowayensis, Fische, Muscheln.
Acheul-Industrie.
Horizont 3 Grellrote harte Tuffite mit eingelagerten Schottern ('Rote
Bank'), Fauna wie in 2. Jiingeres Chelles.
Horizont 2 Stumpffarbige, erdige Tuffite mit Gerollagen. Fauna wie in 1,
aber ohne Dinotherium. Chelles-Industrie.

Horizont 1 Graue Tuffite mit Praechelles-Industrie. Fauna mit Dino-

therium, Elephas antiquus recki, dreizehigen Pferden, Zebras,
Antilopen, Krokodilen, Schildkroten.

Lava.
'Hopwood, A. T., 1932. "The Olduvai Expedition, 1931." Nat. Hist. Mag., Ill, pp. 219, 220.
-Dietrich, W. 0., 1933. Centralblatt f. Min., etc., Jahrg. 1933, Abt. B, No. 5, p. 300.
1434 OSBORN: THE PROBOSCIDEA

Somewhat opposed to Dietrich's conclusions are Hopwood's final conclusions as to the age of the Olduvai
deposits. This author, as a result of his field work at Olduvai and his subsequent studies of the fauna, has decided
that the four lower successive beds represent a more or less continuous series, all of Middle Pleistocene age, for he

found that the lowest bed contained a large representation of Olduvai mammals, in all respects similar to the forms

from the beds above. This was in contrast to Dietrich's conclusion that Horizon 1 is relatively barren of fossils.

Therefore, in speaking of the Olduvai fauna, Hopwood' said that: "Die fauna von Oldoway muss daher als

einheitlich, und zwar als mitteldiluvial, betrachtet werden."

As to the uppermost bed. Horizon 5, Hopwood makes no comment in this latest contribution of his. Since it

is separated from the underlying sediments by a distinct disconformity and since its origin is entirely different

from that of the beds preceding it (this bed is composed of a complex series of wind-blown dust) the evidence
would seem to be strongly in favor of a later age for this uppermost bed.
Therefore, the sequence of deposits at Oldoway might be represented in the following manner.

Horizon 5 — Late Pleistocene

4
3 Middle Pleistocene
2
1

Lava

A complete list of the Olduvai fauna as known at the present time is presented below. An examination of
this fauna will show at once that it is decidedly African in its constitution, and for the most part the various
genera and species comprising the fauna are rather closely related to the modern African mammals of the same or
nearby regions. There are, however, some curious associations in the Olduvai fauna, and since these have con-
siderable bearing on the age of the assemblage, they will be commented on at this place.

First of all, it is to be noted that the Olduvai fauna contains a number of really primitive holdovers —mam-
mals that are not generally considered as being of Pleistocene age. These are, specifically, an Hipparion {Stylo-

hipparion albertensis) and a dinothere {Dinotherium hopwoodi"-) , mammals that are to be expected in the Pliocene
rather than in the Pleistocene. The Stylohipparion is found at Olduvai associated with true Equus of the zebra
type. A similar association of Hipparion and Equus has been noted in North Africa (above) so it would seem ,

that the Pleistocene persistence of Hipparion and its relatives is a phenomenon restricted to Africa.

As against this occurrence of primitive Tertiary types in the Olduvai beds there is the presence of certain very
advanced forms. Most notable of these is Hippopotamus gorgops, a species that must be considered as the most
specialized member of the Hippopotamidse. Its extremely elevated orbits and constricted muzzle, with widely
flaring canines and long face, mark this form as being structurally more advanced than the modern Hippopotamus
amphibius.

The bulk of the fauna is, as indicated above, closely related to the modern African fauna of the same general
locality. This is particularly apparent among the carnivores, the pigs, and the antelopes. It might be well to say
'Hopwood, A. T., 1937. "Die Fossilen Pforde von Oldoway." Wissenschaftlirho Ergobnisse der Oldoway-Expedition 1913, horaus. v. H. Reck, N. F.,

Heft 4, p. 135.

^Dinotherium bozasi (Qgborn's D. hopwoodi is a .synonym). Hopwood, A. T. — Personal communication.

 GEOLOGIC SUCCESSION: AFRICA 1435

that certain members of the Olduvai fauna are undoubtedly immigrants from the north and from the east, and
these forms have been used to advantage in correlating the assemblage with contemj^ioraneous Eurasiatic faunas.
Characteristic of these immigrant forms are 'Elephas' antiqun>< recki, closely related to Hesperoloxodon antiquus
of the European Pleistocene, and Sivatherium olduvaiensis, a link with Sivathernmi giganteum of the Upper Siwalik
beds of India.
The Olduvai Fauna
Canis africanus Pohle Hippopotamus gorgops Dietrich
Lupulella 7neso7nelas latii-ostris Pohle Sivatherium oldowayensis Hopwood
Prototocyon recti Pohle Giraffa sp. Dietrich
Hyxna brunnea Thunberg
aff. Pelorovis oldowayensis Reck
Desmarest
Felis aff. caffra Philantomba monticola (Thunberg)
Deinotherium hopwoodi Osborn (a synonym of Dinotherium Tragelaphus scriptus (Pallas)
bozasi Arambourg —
Hojiwood) Tragelaphus speckii stromeri Schwarz
Hesperoloxodon antic/uus recki (Dietrich) = Palaeoloxodon recki Tragelaphus strepsiceros (Pallas)
(Dietrich) —
Osborn (p. 1275 of the present memoir) Thaleroceros radiciformis Reck
Papio sp. Remane Taurotragus oryx pachyceros Schwarz
Simopiihecus leakeyi Hopwood Nesotragus moschatus von Duben
Stylohipparion albertensis (Hopwood) Gazella gazella praecursor Schwarz
Equus oldowayensis Hopwood Gazella granti Brooke
Rhinoceros simus germaiio-africanus Hilzheimer Phenacotragus recki Schwarz
Koiropotamiis majus Hopwood Beatragus hunteri Sclater
Koiropotamus sp. Dietrich Damaliscus angusticornis Schwarz
Notochoerus dietrichi Hopwood Pultiphagonides africanus Hopwood
Notochoerus sp. Dietrich Parmularius altidens Hopwood
Phacochoerus sp. Dietrich

It might be well at this place to present the following citation from a recent paper by Hopwood.'

Neither did big faimal changes take place in Africa until late in the Pleistocene. For example, the lower Pleistocene de-
posits of Kenya Colony and Uganda contain primitive elephants, mastodonts, Stegodon, Deinotherium, Sivatherium, Stylohip-
parion, Equus oldoicayensis, Hippopotamus gorgops, H. imaguncula, and large pigs of the Notochoerus-Metridiochoerus group.
In the Middle Pleistocene of Kenya Colony antl Tanganyika Territory the primitive elephants, Stegodon and Hippopotamus
imaguncula, have become extinct, but the other species continue. Deinotherium and a mastodont are both known from Bed II
at Olduvai; they have not yet been found in the higher beds (III, IV). The most striking newcomer is the elephant, E. antiqu-
us recki, which replaces the primitive forms of the lower Pleistocene. ...
The upper Pleistocene fauna, so far as we are at present acquainted with it, contains nothing but Recent species. All the
forms mentioned in the list of the lower and middle Pleistocene faunas appear to have become extinct.

Proboscideans: See list of Olduvai fauna, above.

Kaiso. —The Kaiso bone beds are located on the eastern shore of Lake Albert, Uganda. These deposits have
yielded a fairly extensive series of vertebrate remains, of both aquatic and terrestrial types, associated with
freshwater Mollusca. The vertebrates include such mammals as "Rhinoceros," two kinds of equids, various pigs,

Hippopotamus, and proboscideans, aquatic reptiles, namely turtles and crocodiles, and fish. The association of
these varied vertebrates with the Mollusca, coupled with the physical expression of the sediments, shows clearly
that the Kaiso deposits contain two facies —one freshwater, the other terrestrial, which intergrade laterally each

into the other.

In his description of the Kaiso deposits, Wayland^ made the following remarks:

Lithologically the bone beds are ironstones ranging in composition from extremely ferruginous sandstones to moderately
pure limonite in which oolitic structure is locally developed. They are not uniformly fo.ssiliferous, but very few, if indeed
. . .

any, are entirely without organic remains. ...

It was thought at one time that the different ferruginous horizons were characterized by different types of fo.ssils, but within
the limits of the investigation this view has proved erroneous, for the transition of lacustrine to terrestrial types of organisms, of

'Hopwood, A. T., 1940. Proc. Geol. Assoc, LI, Pt. 1, pp. 86, 87.
-Waylaiul, E. J., 1926, "The Geology and Paleontology of the Kaiso Bone-Beds." Uganda Protectorate, Geol. Surv. Dept., Occasional Paper, No. 2,
pp. 9-11.
1436 OSRORN: THE PROBOSCIDEA

whifh this faunistic ilirtViviicc consists, is found to be lateral i-atlicr than vcftifal. Indeed it appears probable that, could one
trace an ironstone patch coniiiletely to its limits, and study in detail the variation of its fossil contents, the purely lacustrine
organisms woukl. in Reneral, i)e found grouped toward the centre, the terrestrial forms round the periphery, and a graduated
mixtiu'c of aquatic and land-freciuentiiig species between. At any rate the evidence .so far gathered points that way. It would
appear then that the ironstone patches of any one horizon represent, singly or .severally, a luimber of pools that constituted the
remain.s of a desiccated Albertine lake of Plio-Pleistocene days.

Hopwood, who described the mammals from the Kaiso deposits, identified the forms given in the Ust below.

Mackairodus sp. flylochwrus euilus

Rhinoceros srotli Hippopotamus ainphibiiifs kaiserisis
Hipparion ulbericiisis IIippopoUitims inidgiuirula
Kquus zebra Klephas zidii (Lo.rodoiita zulu of this Memoir)
Chalicothere Elephas afl'. nieridionalis {Archidiskodon ineridionalis of
Sus Umnetes this Memoir)

Hopwood's original fonclusions were that this fauna is for the most part typically African in its complexion,
tliat it contains certain immigrant forms from the northeast, namely, Hipparion and Archidiskodon meridionalis,
that the small hippoiootamus, H. imaguncxda, is closely related to the Pleistocene Mediterranean form, //, viada-

gascariensis, and that the fauna is of Pliocene age. Subsequently (1929), he decided that the Kaiso fauna, like

the other related faunas of Central Africa, is certainly of Pleistocene age, a conclusion thoroughly justified by the
facts of the case. The presence of Machairodus, Hipparion, and a chalicothere in the Kaiso fauna is certainly due

to the persistence of these forms into the Pleistocene.

In a recent communication to the author Hopwood makes the following statement: "Additional material
seems to show that only one elephant occui's at Kaiso, namely 'Loxodonta griqiin' Haughton. An imdescribed
Stegodon has also been foinid there."

Proboscide.\ns: See faunal list, above.

Lake Nyasa. — The fossils from Lake Nyasa come from the Chiwondo Beds of Uraha Hill. The specimens
are extremely fragmentary and scanty, being broken proboscidean teeth identified by Hopwood as belonging to

a "Bunolophodont Mastodon," and several bones referred to Hippopotamus. The age is probably Pleistocene.

HoMA Mountain. — At Homa Mountain, near the eastern shore of the Victoria Nyanza, there have been
found various fossil remains of antelopes, Phacochoerus, Hippopotamus, and a baboon, described by Andrews as
Simopithecus oswaldi. In 1926 Hopwood described additional mammals from Homa Mountain, as follows:

Hippopotamus amphibius Linn.

Metridiochwrus andrewsi Hopwood
PhacochcETus xthiopicus Linn.
Bos sp.
Elephas antiquus recki Dietrich'

The complexion of this fauna is distinctly Pleistocene, and since the primate is closely related to the Papio
from Oldoway, described by Remane, while there is complete identity between the proboscideans of these two
regions, there is every reason to think that the Homa Mountain deposits are contemporaneous with those of
Oldoway. This would make the age Middle Pleistocene.

Proboscideans : Bee faunal list, above.

—
Lake Rudolph. Fo.ssils from this area, which may be designated as belonging to the Omo fauna, were first

described l^y Haug in 191 In recent years the Omo fauna has been made a subject of special
1 . investigation by
Arambourg,- who has shown that:
'"Kanjora Elephas antiquus rrcki. Thi.s locality includes Himia Mountain." Hopwood, A. T. — Pcnsonal coramunication.
-ArambourK, C, 193.'). Bull. Soc, rcoI. France, (.5), IV, pp. :«)(>, 307.
 .

GEOLOGIC SUCCESSION: AFRICA 1437

1 The Omo fauna closely resembles that of Olduvai

2. Like the Olduvai fauna it contains:
a. Many typically Afi'ican genera similar to the modern forms in the same ai-ea.
b. Certain persistent types, that would seem to be holdovers from Tertiary times.

Les gisements de Vertebres de I'Orao n'ont point livre d'industries humaines. Leur association faunique parait corre-
spondre en partie a celle que Leakey a observee daiisle ravin d'Oldoway (Tanganyika) iniin^diatement au-dessous des niveaux
a industries de facies chelleen; il parait done logique de les considerer comme appartenant a cette periode do transition qui
separe, on Europe, la fin du Pliocene de I'apparition des industries humaines, si tant est que Ton puisse admettre le synchronisme
de celles-ci entre des regions aussi eloign^es.

It is a foregone conclusion, therefore, that since the mammalian assemblage at Omo would seem to be virtually
identical with that of Olduvai, it is contemporaneous with the latter and consequently is of approximate Middle
Pleistocene age (compare faunal list on page 1435).

Proboscideans :

Dinolherium hozasi Araiiibourg {D. hopivoodi Osborn a synonym — Hopwood)

Palseoloxodon antiquus recki (Dietrich)

7. THE PLEISTOCENE OF SOUTH AFRICA

ZuLULAND. —The fauna described by Scott (1907)^ from Zululand has a decidedly advanced aspect. It is

doubtful whether this fauna is any older than the Middle Pleistocene and it might be even later than this in age.
Hopwood," by inference, would make this assemblage more or less contemporaneous with the Middle Pleistocene
faunas of Central Africa.

"The fossil mammals of South Africa are all of comparatively recent date. They have been described by
various authors, . . . and the general assemblage differs from those of the same date in Central Africa only in mat-
ters of detail."

In a recent letter to the author, Hopwood states that the Zululand fossils are, in his opinion, equivalent in
age to the Olduvai fauna. The fauna as described by Scott, is as follows.

Opsiceros simplicidens Scott

Loxodonta zulu Scott
Hippopotamus ponderosus Scott
Bubalus andersoni Scott

The fossils from Zululand were placed by Scott in the Upper PHocene, because of the seeming presence of
marine beds with Tertiary molluscs above the sediments in which the mammals were found. On the other hand,
Scott recognized the obvious Pleistocene character of the mammalian remains, admitting that they might be
more properly placed in the Pleistocene than in the Pliocene.
Concerning the position and mode of occurrence of these fossils Mr. Anderson writes as follows: 'The fossils were scattered
over a large flat outcrop of shales, which occurs below the level of ordinary low-water mark, and is only exposed under the ex-
ceptional circumstances of a strong south-easterly gale and a neap tide, when the large covering of sand is removed. Overlying
this bed are a series of shales with a few scattered bones and crustacean and fish remains. Above these a thin layer containing
Foraminifera, and then a foot or so containing marine Mollusca, which Mr. Etlieridge referred to the Tertiary period above ;

this a thick series (probably over 100 feet) of false bedded sands of various colours covered by the Recent sand dunes.'
Seeing that these mammals all belong to existing genera, and that, with the exception of the elephant, they differ but com-
paratively little from Recent Ethiopian species, it is obvious that the fossils cannot have any very great geological antiquity,
and probably they should be referred to the later Pliocene. So far, however, as the mammals themselves are concerned, they
might almost equally well be regarded as early Pleistocene.
'Scott, W. B., 1907. Third Report of the Geological Survey of Natal and Zululand, pp. 253, 2.54.

^Hopwood, A. T., 1929. Amer. Journ. Sci., (5), XVII, p. 103.

1438 OSBORN: THE PROBOSCIDEA

Due to the conditions under whicli these fossils were found, it is indeed doul^tful whether they were in pUxce
beneath marine Tertiary sediments.

PROBOscroEAN : See faunal list, above.

Vtlal River. —From the Vaal and Limpopo rivers (principally the former) of the Transvaal, South Africa,
a bewildering array of fossil elephants has been described during the course of the past fifteen years. The first

descriptions were made by Haughton more recent years by the rather prolific creation of
(1922), supplemented in
species by Dart (1927, 1929), with additional contributions by Osborn (1928). These fossils for the most part come
from various locahties and from different stratigraphic ])ositions along the Vaal River, and since there exists a con-
siderable amount of confusion as to their stratigraphic positions and geologic ages, the problem of their deposi-
tion and associations must be approached with a great deal of caution.

Essentially, the condition along the Vaal River, south of Johannesburg and in the vicinity of Bloemhof, is as
follows.

There are three terraces exposed along the river, a high terrace some 200 to 300 feet above the river bed,
a middle terrace at approximately 60 to 80 feet above the water, and a low terrace grading into river-bed gravels
contiguous to or within the present river. The high terrace is, so far as known at present, unfossiliferous. It is

from the middle terrace and the low terrace and river gravels that the several types of fossil elephant teeth have
been found. It might be added, too, that in the middle and low terraces and the river gravels several distinct
stone cultures have been discovered and have been used for the stratigraphic differentiation of the sediments.

Dart concluded that the Vaal River terraces represent successive phases by the early Vaal
of aggradation

River, the high terrace being of Pliocene age, the middle terrace very probably containing two levels, one of
Pliocene age, above which is a later Pleistocene level, and the low terrace being of late Pleistocene age, containing,

as suggested by van Riet Lowe, two distinct gravel layers.

In 1932 Haughton' raised the question as to the vaUdity of this sequence. He pointed out the very important
fact that "the earliest date that can be assigned to any undisturbed river-gravel is the date at which the lowest
beds in a continuous sequence were deposited ; and that comparisons between gravels at varying distances from
a river or at various places along a river valley should be based upon the altitude of the bottom-bed of each de-
posit with respect to the present river-level, and not upon that of the top of the gravel."

Haughton then went on to show that the middle terrace of the Vaal River is very close to the present river
channel, and that at each of the places where fossils or flint implements were discovered in this middle terrace, the
base of the terrace is only a few feet above the present river-bed level. With this consideration in mind, Haughton
presents the possibility that the middle terrace beds and the river gravels or low terrace beds are in reality part of
a single, continuous series of deposits. If such be the case, the conclusions of Dart would be greatly modified,
since there would be no time-gap between the fossils found in the middle terrace and those of the river gravels.
Essentially, the specimens from the middle terrace beds would therefore become much younger in their geologic

age than they were considered by Dart or by Osborn.

Which conclusion as to the sequence of the Vaal River deposits is valid, Dart's or Haughton's, is a question
that cannot be answered at the present time. As Haughton has pointed out, "Much detailed investigation is

necessary before the true sequence of events can be elucidated." It might be mentioned, however, that the
'Haughton, S. H., 1932. Trans. Roy. Soc. S. Afr., XXI, Pt. 1, p. 15.
 GEOLOGIC SUCCESSION: THE ORIENT 1439

evidence in favor of Haughton's views is very strong. In this connection Hopwood, in a personal communication
to the author, expresses his opinion that Haughton's work is at the present time the most satisfactory interpre-
tation of the Vaal River deposits.

Stbatigraphic Relationships of the Vaal River Proboscidea

High Terrace Level not specified
Lozodonta suhantiqua (Haughton) Palseoloxodon archidiskodontoides (Haughton)

Middle Terrace Lower Terrace (River Bed Gravels) —Pleistocene

Upper level (Lower Pleistocene — Dart; Pleistocene —Haugh- —
Archidiskodon broomi Osborn washed in from Middle
ton) Terrace?
Metarchidiskodon grif/ua (Haughton) Palseoloxodon transvaalemjs (Dart)
Archidiskodon yorki Dart Palseoloxodon ^heppardi (Dart)
—
Palseoloxodon hanekomi (Dart) more recent than P.
T /TDi-
— T~i i T>i
Lower level (Pliocene Dart; Pleistocene Haughton)
1 1
•
i
— TT 1, ^ \ transvaalensisi

Bunolophodont mastodont River Gravels— possibly related to the Lower Terrace gravels
PalxoloxMonif) andrexosi (Dart)-from a depth of 80 feet
Palxoloxodon yorki (Dart)
within terrace t> i„ j f-n
,,.,.,, ,
Archidiskodon vanalpneni T-.
Darti
,
i
Palxoloxodon wilmani (Dart)
i t^

,,.,.,, 11 /, r\ ,
d ;~ ? j /ti
Palxoloxodon kuhni (Dart)'
; i, *^
Archidiskodon ynilletti iJart
Archidiskodon loxodontoides Dart Recent
Archidiskodon subplanifrons Osborn —washed in from Loxodonta prima Dart
High Terrace? Loxodonta africana obligua Dart

III. THE ORIENT

1. INTRODUCTION
The Oriental region as here defined is that portion of Asia south of the Himalaya Mountains and the Tibetan
Plateau, namely, India and Burma and adjacent countries, South China, Indo-China, and the East Indian Islands.
Thus the Oriental region as used in this chapter coincides with the Oriental Region or Realm as used by the zooge-
ographers. In this case it is practicable to make the zoogeographical term and the term as used herein equal in
value —whereas in Africa, as we have seen above, the strict utiUzation of the zoogeographic division was not
convenient for our purposes.

By far the most important part of the Oriental region is northern India, where the tremendous thickness of the
Siwalik Series constitutes one of the most remarkable and productive sequences of fossil-bearing beds in the
world. Here, through a vertical extent of some twenty thousand feet of sediments, the development of the
Upper Tertiary and Quaternary mammalian faunas of India can be traced with the greatest of detail. And
throughout this long and virtually unbroken series of fossiHferous beds the remains of fossil proboscideans are of
major importance.

But for the earKest Proboscidea-bearing sediments of the Oriental region it is necessary to go outside of the
Siwalik area, to Baluchistan and Sind, where the early Miocene continental beds have yielded a considerable fauna.

2. THE MIOCENE OF BALUCHISTAN AND SIND

—
Baluchistan, Bugti Beds. Fossil vertebrates were found in Baluchistan as long ago as 1846 by Vicary, but
it was not until 1882, when Blanford visited the region, that serious attempts at collecting were made. Blanford
brought back a few anthracothere remains, which were subsequently described by Lydekker.
 :

1440 OSBORN: THE PROBOSCIDEA

But the mammalian fauna of Baluchistan remained virtually unknown until the years 1907-1910, when
Pilgrim and Forster Cooper independently conducted several expeditions to that region, particularly to the area
of the Bugti Hills, in the vicinity of Dera Bugti, Kumbhi, and Chur Lando, with the result that a large mamma-
lian fauna was discovered.

The mammal-bearing beds of the Bugti Hills have been described by Pilgrim' in part as follows

The ossiferous Upper Nari series [these beds were subsequently correlated by Pilgrim with the Gaj series, rather than with
the older Nari series] of the Bugti country attains a thickness of 1,000 feet, the rocks are subject to considerable lateral variation,
and it is often difficult to trace any particular bed for more than a short distance. As it happens, the same fossil species occur
at different horizons, so that it has been found impracticable to regard the series otherwise than as a whole. . . .

Fossil bones and teeth are met with at various horizons, from the base of the series up to within 100 feet of the top, on
either .side of the Zen range south of Dera Bugti and the Sihaf valley, particularly near Gandoi and Kumbhi which are situated
on the same meridian on exactly opposite sides of the range. Unfortunately, though well preserved, they are for the most part
. .

isolated fragments of small size.

^EOlS)GICAL REl^UlOAy^/PS OF OR/£NT:iZ. PROBOSC/DE^

CENTRAL AND SOUTH CHINA AND
B.AUJCHISTAN
AND SIND P1I.CWM LEwii NORTHERN INDIA SOUTHERN INDIA CEYLON BURMA INDO-CHINA JAVA PHILIPPINES
BORNEO
K.4RNUL EUph,as indicus SAMPO^'JVG
CAVES Elephas indicua
'mKrbXda /^GANDOA^G 5.triqonocephalus
k GODAVAR/ A10G0K '

Pala'>o(oxodon ,na- *-^

BOULDER Steaodon
w to 5 re. ^aneaa, Sfe^odon orlenfalis Sfe3odon tn^ono- Steoodon
CO^GIOMERATE pr'ientalis Palaeoloxodon
D^E T/S cephalu3| ndanens
OJ Polieolo-^ovion Palaeoloxodon namadlcus
0. [TAIV/). namadtcus naniadicus Stegodon tria. praecursor '-
'^

~UPP>BK Sfeqodon Elephas sp,

btegolophodon
selepn Tiusu'dr
=>?;
PINJOK ppatele phasplarucepKal us IRRAyVADD'Y elephan^olde5, KAZ/ GLAGAHX ^udekker.
Arch idiskodon
piamrrons ^te^olophodon insignis Tefralophodon bumtaiuenais I^ORNEO)
Ste^odon bombifroMS, ganesa, insidnis, pinjorensis Palaeolo^odon latidens Steqodon tnqonocephalus praecursor
TATROT ^^S?^9?^P^H^ stegodonfoides Stegodon ArcKid'iskodon planlfrorB
' Penralophodon ralconeri sivale elephanroides,
jnsi^gnis birman icus
TJJ D^OELAUG^
1
Sfe^odon sp.
nLjpselephas hi.i^sudricus _
TJJ SA/YDE.
5t.
Dinufherium indicum. an^ustidens oTTicrnalts,'
Tnlophodon hasno^ensls
^HOK Tetralophodon sinensis '

UJUJ falconer!, punjabiensis

>u Tv/TT/j^Khunchorherium chinuense
''O t^j-ijrvHjy
Sur'iconolopnus dhokpatnanensis, propahnanensis,
-a corru^atus, pfiycHodus, hasnott
Ste^olophodon latidens, cauMeui
-^
3teqodon bomDirrons
•5
Anancus perlVnensis
MACRf

.LU Dinotherium sindlense, indrcurn, pentapotamiae ,.

Tnlophodon anaystidens palaeindicus, macrognathus, cninjie

^ty,., ,, Tetralophodon jalconeri ...,,. ,, ..
c/r/Aiy/ 5errid^nt"inu5 hasnofensis, mefachiniiensia, broiuni, chinjiensis, prochinjiensis
.

Si^nconolophus ptuchodus
3£ Anancu^ properimensis
^o ^ L 2 tSteaotophodon cautleui prqaressus.i
. t.

. UJ
-c Ox £ ^r.
?.inotherium sindienae
^ ,
I
riiophodon pandionis, macroanatnus
r — u
4
BUGTI LDoubrful
occurrence in
the Pleistocene P.F U^-^tT SraJUjf

Fig. 1221

In 1912, as the result of more detailed and considered studies of the Bugti fauna, Pilgrim- came to the con-
clusion that it should be correlated with the Gaj series, of Aquitanian-Burdigalian age, rather than with the Ndri
beds.

In his discussion of the Bugti fauna, Pilgrim shows first that it is .separated by con.siderable differences from
the Lower Siwalik fauna of Sind and the Punjab. He therefore concludes that there is a great hiatus between
these faunas, a conclusion that is substantiated by the stratigraphic evidence. Continuing, with a comparison
between the Bugti fauna and like faunas in Europe and northern Africa, Pilgrim finally concludes that "it seems
improbable that the bone beds of the Bugti hills are younger than uppermost aquitanian."
'Pilgrim, G. E., 1908. Rec. Geol. Surv. India, XXXVII, Pt. 2, i)|). 141, 143.
^Pilgrim, G. E., 1912. Pal. Indira, N. S,, IV, Mpin. No. 2, pp. 2, .5.
 GEOLOGIC SUCCESSION: THE ORIENT 1441

Both Pilgrim and Forster Cooper, as the result of their independent studies on the fossils of the Bugti beds,
decided that these remains represent a unit fauna of essentially Lower Miocene age.

In preparing preliminary manuscript and notes for this chapter, Professor Osborn came to the conclusion
Dera Bugti and at Chur Lando might be assigned to two stratigraphic horizons.
that the fossiliferous beds at
This determination was based on the supposed presence of Baluchilherium, at Chur Lando and its absence at Dera
Bugti, denoting in his opinion the fact that the beds at the former locality were older than those at the latter place.

"Upper level: Dera Bugti bone beds; no remains of Paraceratherium or Baluchitherium. Lower level:
Chur-Lando bone beds; Baluchitherium or Paraceratherium life zone." (Osborn, H. F., Notes.)

In his most recent notes, Professor Osborn marked the "Chur-Lando bone bed" as of Lower Miocene age and
the "Dera Bugti bone bed" as of Lower to Middle Miocene age.'

Since Professor Osborn's supposition of two zones or levels in the Bugti beds was quite contrary to the publish-
ed opinions of both Pilgrim and Forster Cooper, it was thought advisable at the time to secure statements from
these authorities as to the probabilities of such a division. I take the liberty of quoting from their letters, written
in reply to the request.

Personally I am not convinced of the necessity for erecting a stage for the Chur Lando bone bed with Baluchitherium,,
separate from the rest of the Bugti stage of the Gaj, but in any case it seems impossible that any part of the Bugti stage can be
newer than the Burdigalian. This is proved by the fact that 4000 feet or more of Murrees north of the Salt Range intervenes
between the Kamlial stage (Helvetian) and the Fatehjang stage which contains Baluchitherium, Anthracotherium cf. bugtiense,
Brachyodus cf. africanus, Hemimeryx, Palaeochoerus pascoei and Brachypotheriutn fatehjangense. Thus even if the Kiunbhi and
Gandoi beds are the equivalent of a stage in the Murree a considerable way above the Fatehjang, they must still fall into the
Burdigalian.
If the Chur Lando bone bed is earlier than the oyster beds with which presumably the Kumbhi fauna is associated, there
is much more to be said for Professor Osborn's view, but this seems hardly likely, nor apparently does Cooper suggest anything
in support of this conjecture.
Baluchitherium may of coiuse have died out suddenly before the bulk of the Bugti beds was deposited, but it seems to me
as reasonable that the Chiu' Lando bone bed was a sort of cemetery for that particular species, and may have been contem-
poraneous with the Kumbhi beds.
If one is determined to invent stratigraphical possibilities, then I should say that the Chur Lando bone bed may be of any
age later than the Stampian, that an unconformity separates it from the Kumbiii and Khajuri beds, which through the as-
sociated oysters must be at any rate Burdigalian, probably Lower Burdigalian. At the same time I do not see why Baluchi-
therium bugtiense should not be Burdigalian just as much as Cadurcotherium indicum, since the latter genus elsewhere occurs in
the Oligocene. Cooper alone is in a position to estimate the stratigraphical value of the Chur Lando bone bed, and if he thinks
that both it and the Kumbhi beds are later than the oyster beds then both Professor Osborn's stages must lie in the Burdi-
galian. (Pilgrim, G. E., September, 1931. Personal communication to E. H. Colbert.)

A good number of my specimens were picked up on the ground washed out by rains so that their precise level must remain
in doubt. Personally I cannot feel that there are sufficient grounds for establishing two horizons, and it is pretty clear that Paracera-
therium bugtiense is not confined to the Churlando deposit as a number of large lower molars were found scattered in other parts.
This is sufficiently clear when you remember that Pilgrim was the first to describe the species and that the Churlando bone bed
was not discovered and opened up until I went to Baluchistan some time later. No one of Pilgrim's specimens therefore can be
considered as coming strictly from this bed. (Cooper, C. Forster, 1932. Personal communication to E. H. Colbert.)

Proboscideans:
Trilophodon pandionis Falconer
Trilophodon angustidens pabeindicus (Lydekker)
Trilophodon angustidens (Cuvier)
Trilophodon cooperi Osborn
Dinotherium indicum gajense Pilgrim
Hemimastodon crepusculi Pilgrim [Suina of Osborn j

In Volume I of this Monograi>h (page 275, caption to figure 221) the "Chur-Lando"liorizon i.s designated a.s being of Upper Oligocene age. Profe,s.sor
Osborn was inclined at times to regard this correlation as the correct one, thereby making the "Dera Bugti" beds of Lower Miocene age. In this discussion his
later views, as presented above, are followed.
1442 OSBORN: THE PROBOSCIDEA

SiND. —Anj' remarks as to the Bugti beds in Baluchistan apply equally well to the contemporaneous de-
posits of Sind. Pilgrim' discussed the Lower Miocene of Sind as follows.

I liavo elsewhere remarked upon the close similarity to the Bugti bono beds presented by certain unfossiliferous sands and
conglomerates at Bhagathoro in Ix)\ver Sind. These also rest upon Lower Nari limestones and were referred by Blanford to the
Upper Nari. Mr. Vredenburg, however, considers that a shell bed, which overlies them, is Upper Gaj. There seems, therefore,
little doubt that these, like the Bugti beds, represent a fresh water facies of the Gaj. Generally, these beds do not contain
. . .

more than mere fragments of vertebrates, but near Fatehjang, Anthmcotherium hugtiense, Brachyodus cf. africanus, Teleoceras
fatehjangeiise, a species very closely allied to Teleoceras blanfordi, and a species of Hemimeryx have been found. Across the
Indus, in the Kohat district, there occurs in similar beds a ribbed Unio, which may be the same as one of the curious ribbed
species found in the Gaj series of the Bugti hills.

3. THE SIWALIK SERIES (MIOCENE— PLEISTOCENE) OF NORTH INDIA

Punjab, Northwestern India. —In the northern Punjab district, along the Siwalik Hills and in the Salt
Range are found tremendously thick deposits of continental sediments, constituting the Siwalik Series, the most
important mammal-bearing beds in the Oriental region and Ukewise, one of the most important mammal-bearing
series in tlie world. Here are exposed more than twenty thousand feet of sediments, deposited as a more or less

unbroken sequence ranging from the Miocene through the Lower or Middle Pleistocene periods. Since the
Siwalik Series is a continuous sequence of sediments, it will be convenient to discuss the various divisions of the
series together.

The Siwalik Series has been divided in its larger aspects into three main divisions or groups, namely, the
Lower Siwalik, the Middle Siwalik, and the Upper Siwalik beds. Each of these three principal groups of the
Series has again been divided into formations or zones, as follows.

fBoulder Conglomerate (Tawi of Lewis)

Upper Siwalik jPmjor
j^Tatrot of Lewis)

Middle SiwaUk
Dhok Pathan
Nagri
Chinji
Lower Siwalik
Kamlial

The establishment of this detailed sequence of deposits is the result of studies that began with the pioneer
work of Hugh Falconer, a century ago, and which have continued up to the present time. Numerous authors have
contributed to the Siwalik problem, notably Falconer, Cautley, Lydekker, Pilgrim, Matthew, Colbert, Lewis, and
de Terra and Teilhard. Naturally opinions have differed a.s to the proper correlation of the several zones or
formations within the SiwaUk Series, but generally speaking it may be said that the lowest Siwalik horizons are of
Miocene age, while the highest ones are located within the Pleistocene. The intervening beds cover the period of
time between the Miocene and the Pleistocene.

Proboscideans appear in the Kamlial formation and they form a very important portion of each fauna through
all of the succeeding beds.

show the results of alluvial sediments accumulating in a region closely adjacent to

Lithologically the Siwaliks
a rapidly ui)lifting momitain ma.ss. The Lower Siwaliks are sands and clays, with occasional beds of heavier
material. These (h^jwsits, i)articularly those of the Chinji zone, are bright red in color and contain what Pilgrim
has called "pseudo-conglomerates"~bands of concretionary beds. Passing up into the Middle Siwaliks, the
'Pilgrim, G. E., 1912. Pal. Indica, N. S., IV, Mem. No. 2, p. 2.
 GEOLOGIC SUCCESSION: THE ORIENT 1443

sandstones become somewhat coarser, with clay beds less frequent. These Middle Siwalik beds are light gray in
color —not red — and they often contain large amounts of relatively unweathered minerals. In the Upper Siwaliks
there are sands and clays, often heavily cemented, and capping the series is a very heavy conglomerate made up of
extremely coarse gravels and large stones.

These sediments pass upward from the lower to the higher beds with but few apparent breaks. Recent work
has shown that there is a distinct angular unconformity between the Dhok Pathan horizon and the overlying
Tatrot zone, wliile the Upper Siwalik Pinjor zone is separated from the capping Boulder Conglomerate by another
unconformity.

Although fossils range throughout the extent of the Siwalik Series, three main faunas characterize the
sequence. These are the Lower, Middle, and Upper faunas, characteristic of the three groups to which these names
are appUed. The Lower Siwalik fauna is typically developed in the Chinji formation, the Middle Siwalik fauna
in the Dhok Pathan formation, and the Upper Siwahk fauna in the Pinjor formation. The fossils of the Kamlial
formation are genetically related to those of the Chinji, while the Nagri fossils are transitory between the Chinji
and Dhok Pathan faunas. The Tatrot fossils may be either transitional between the Dhok Pathan and the
Pinjor or identical with the latter fauna.

Falconer, who knew only the Upper Siwahk fauna, regarded it as of Miocene age. Lydekker, who described
a greater portion of the Middle Siwahk fauna, placed the tw^o faunas with which he was acquainted in the Phocene.
It remained for Dr. Pilgrim to discover the Lower Siwahk fauna, and to show the true relationsliips of these faunas
to each other and to point out their probable ages.

After many years of intensive study of the Siwahk mammals. Pilgrim came to the conclusion that the Lower
Siwaliks, Kamhal and Middle Miocene age (Helvetian and Tortonian, respectively), while the Nagri
Chinji, are of
horizon is to be placed in the Sarmatian or Upper Miocene. Furthermore, he argued that the Dhok Pathan
should most properly be placed in the Pontian, and the Tatrot and Pinjor in the middle and upper portions of the
Pliocene, respectively. The Boulder Conglomerate was placed by Pilgrim in the Lower Pleistocene. Pilgrim's
views,' which have been recently epitomized in a short paper on the correlation of the Siwaliks (1934), may be
shown to advantage as follows.

Lower Pleistocene Boulder Conglomerate

Upper Pliocene (Val d'Arno) Pinjor
Middle Pliocene (Montpellier) Tatrot
Lower Pliocene — Pontian Dhok Pathan
Upper Miocene — Sarmatian Nagri
-..,,, ^.. f
Tortonian Chinji
Middle Miocene— <^ ,, , . ., ,. ,

I Helvetian Kamlial

Pilgrim's views as to the correlation of the Siwahks were based on his comparison of the several Siwahk
faunas with what seemed to be their equivalents in the European sequence. The outstanding characteristics of
the Siwahk faunas, on this basis, would be the close resemblance of the Chinji fauna to the typical Tortonian fauna
ofLa Grive-St.-Alban, similarly the resemblance of the Dhok Pathan fauna to the wide-spread Pontian faunas of
Pikermi, Samos, and Maragha, and finally the close comparison between the Pinjor fauna and the so-called Upper
Phocene fauna of Val d'Arno.
'Pilgrim, G. E., 1934. Amer. Mus. Novitates, No. 704.
1444 OSBORN: THE PROBOSCIDEA

In 1929 Dr. Matthew,' having studied the Siwahk fossils inLondon and Calcutta, attempted to modify
Pilgrim's correlation by stepping the whole Siwalik Series up in the geologic column from the position assigned to

it by Pilgrim. In doing this Matthew was guided by two main considerations.

1. The appearance of new invading elements in a fauna is a safer guide to its correlation than the disappearance of old
elements or the average composition of the fauna as a whole. The appearance of these new elements must be interpreted in the
light of what is known of their origin and dispersal. When this is as directly recorded and fully documented as it is in the case
of Tertiary Equid» or Camelidse, the evidence appears not open to any effective challenge. But more often the appearance of
new elements in a fauna may be explained in several ways, the relative probability of which is not easy to test.
2. India and the Oriental region generally are today characterized by the survival of many primitive types of mammals
as well as by the absence, scarcity, or recent appearance of some of the most progressive and specialized mammals. It compares
in these respects with West Africa and tropical America. While it does not necessarily follow that this was true during the later
Tertiary, yet it should be so considered until evidence proves the contrary; and so far from proving the contrary I believe that
all of the evidence conforms with this assumption and much of it is difficult to explain in any other way.

Matthew argued that since Hipparion in India first appears in the Chinji beds, these deposits cannot be older
than the oldest Hipparion-hearing beds of North America, which latter mark the first appearance of Hipparion
as a direct descendant of the North American Merychippxis. In other words, Hipparion arose in North America
and subsequently migrated to the Old World — consequently its appearance in the eastern hemisphere must of
necessity be later than its first appearance in North America. And Matthew regarded the appearance of Hip-
parion as indicative of the favmal change that marked the transition from the Miocene into the Pliocene.

Likewise, Matthew jwstulated that Equus arising in North America at the beginning of the Pleistocene, did

not reach Eurasia until after its first appearance in the New World. Consequently, the Upper Siwalik beds con-
taining Equus would of necessity be of Pleistocene age. In this respect, it might be said that Matthew ])laced the

\'al d'Arno and Villafranchian faunas of Europe in the Lower Pleistocene because of the presence of Equus in

these horizons —a decision differing from the traditional European practice of assigning this stage to the Upper
Pliocene.

It might be .said that Matthew's correlation of the Siwaliks was not based on the evidence of the Equida*

alone. He showed that the Chinji giraffes are comparable to the Pontian giraffes, while the Dhok Pathan giraffes

are more advanced than any giraffes of tlie Pontian. And in the Upper Siwaliks, he cited the appearance of the

camel as additional evidence of the Pleistocene age of these beds.

Therefore, lie argued, the general resemblances of the Siwalik faunas show their homotaxial but not their
correlative identities. These are relict faunas, in which the influx of new types from the outside furnish the real
clews as to their age.

Pilgrim,- in 1931 , answered Matthew's argument as follows:

Matthew (1929) ha.s recently sought to replace my coirelation of lln' Dhok Pathan stage with the Pontian l)y another
which, to judge from his diagram on ]). 441, puts both it, as well as the earlier Nagri stage, later than Pikcrmi; assumes a gap,
which does not exist, between the Middle and Lower Siwalik, and makes the Chinji and Kamlial stages start in the Vindobonian
and end at an horizon which is the equivalent of Pikermi. His ai'gument is mainly i)ased on the first occurrence of Hipparion
in India at the top of the Chinji stage, but he considers that the remainder of the fauna, iticluding the Carnivora, support it.
Apparently, while admitting in part the occurrence in the Dhok Pathan of species allied to those of Pikermi, he regards these as
relics of an earlier age. If this is so, we have; the choice of alternatives:
. .
(1) either such forms nuist have migrated from the
Holarctic region in Pikermi times and lingered on in India to a much later epoch; or (2) such migration did not take place until
post-Pikermi times. If we adopt the first alternative we ought to find that the Nagri fauna and that of the uppermost Chinji,
which by hypothesis are the equivalent of Pikermi, contain Pikermi species or species at a similar stage of development; but
if the second, neither the Nagri nor Chinji fauna ought to contain any immediately ancestral typ(>s of the Pontian fauna of the

Holarctic region. Actually, however, neither is true of the Carnivora, nor it may be said of other mammalian orders. We do

'Matthew, W. D., 1929. Hull, Amcr. Mus. Nat. Hi.st., LVI, p]). 1 12, 443.
'Pilgrim, G. E., 1931. Catalogue of the Pontian Carnivora of Europe, Brit. Mus, (Nat. Hist.), |)|>. 151, 152,
 :

GEOLOGIC SUCCESSION: THE ORIENT 1445

not find Pikermi spocies at any earlier stage than the Dhok Pathan (the Perim Ishind beds I now regard as lieloiigiiig to the
Dhok Pathan stage); while the Xagri and, still more so, the Chinji stages contain species that are quite definitely ancestral to
(or at any rate more primitive than) those of the Dhok Pathan and the Pontian of Europe alike.
Matthew's argument from Hipparion is more difficult to answer. If flipparion originated in North America in Pontian
times it obviously could not have reached India before the Pontian. But his j)reniises lack absolute jiroof. First, although badly
preserved equid teeth do occur at the top of the Chinji, arc these «>rtainly Hipparion and what stage of development do they
represent? Secondly, did Hipparion originate in North America and not in C'entral Asia? Thirdly, if it did, is the age of the
Valentine, in which Hipparion first appears in America, demonstrably contemporaneous with and not earlier than, say, the
Sebastopol faima of the Black Sea region in Europe? The discovery of a fauna which contains the immediate ancestors of the
hitherto described 'Hipparion fauna' of China will settle the question definitely. Personally, I anticipate that such a fauna will
be comparable to some extent with that of the Chinji stage of India.

In 1935 the present author, as the result of protracted studies on the Siwahk collection in the American
Museum of Natural History, came to a conclusion essentially similar to Dr. Matthew's views as to the age of the

Siwaliks, in short, that the Chinji fauna is equivalent to the "Valentine" or transitional Miocene-Pliocene of
North America and is shghtly pre-Pontian, or Pontian, in age, that the Dhok Pathan fauna is definitely post-Pon-

tian and that the Pinjor fauna is definitely of Lower Pleistocene age. There were a few sUght differences between
my interpretations and those of Matthew, namely, the Lower Pliocene and the con-
raising of the Chinji into the
sideration of the Siwaliks as a continuous series rather than as separated by two major breaks, the view taken by
Matthew.

Consequently the correlation adopted by the present author is as follows

Lower Pleistocene /Boulder Conglomerate

\ Pinjor

Transitional Tat rot

Middle to Upi)er Pliocene |Dhok Pathan

^Nagri

Lower Pliocene Chinji

Upper Miocene Kamlial

Subsequently (1937) G. E. Lewis' published still another opinion as to the ages of the several Siwalik faunas.
On the basis of new evidence, especially his stratigraphic field studies, he adopted the following correlation of the
beds forming the Indian sequence.

Middle Pleistocene Tawi (new name for Boulder Conglomerate)

Lower Pleistocene Tatrot (including Tatrot and Pinjor)
Upper Pliocene Break
Middle Pliocene Dhok Pathan
Lower Pliocene Nagri
Upper Miocene Chinji
Middle Miocene Kamlial

This correlation differs from those of Matthew and of Colbert by placing the Lower Siwaliks even lower than
was granted by these two authors, but not so far down as they were placed by Pilgrim, and by extending the Upper
Siwaliks higher into the Pleistocene than had previously been done.

Lewis' views as to the age of the Chinji beds are based to a great extent on the supposed Upper Miocene
appearance of Hipparion in North America, in the Mint Canyon beds of the Pacific coast. He points out that
this Hipparion mohavense is very close to the Siwahk Hipparion, that it is probably close to the ancestor of the
Asiatic species, and that the Mint Canyon formation in which it appears is topped by marine beds carrying
Miocene invertebrates.
'Lewis, G. E., 1937. Amer. Journ. Sci., (5), XXXIII, p. 197.
1446 OSBORN: THE PROBOSCIDEA

As Lewis has remarked, it is probable that Hipparion mohavense is as close or closer to the Asiatic Hipparion
than are any other North American species of this genus, while Merychippus sumani is very likely ancestral to
tliese Hipparion tyjies. Consequently Merychippus sumani and Hipparion mohavense are the forms that should
be compared with the Asiatic Hipparion. The real crux of Lewis' argument is the question as to whether the
Mint Canyon formation is of Upper IVIiocene age. This is the interpretation given by Maxson, who described the
mammahan fauna from these beds. It should be pointed out here, however, that Stirton (1933)' has disputed the

Miocene age of the Mint Canyon, linking this formation with the Ricardo of Lower Pliocene age, rather than with
the Barstow of —
Upper Miocene age as was done by Maxson, and placing it above the Barstow but below the
Ricardo in the geologic column.

One important point in this discussion of the Mint Canyon is, of course, the age of the overlying marine beds.
It should be noted that the invertebrate fossils are not well preserved, and that Woodring's correlation of the de-
posits with the Cierbo formation was more or less tentative. Stirton places the Mint Canyon assemblage as

equivalent in age to the Cierbo and regards both of these formations as being about as close in their time relation-
ships to the Ricardo as they are to the Barstow.

Lewis regards the Nagri as the Pontian equivalent in the Siwalik Series, and places the Dhok Pathan in the

Middle Pliocene, as did Matthew and Colbert.

Finally Lewis shows a great stratigraphic break between the Dhok Pathan and the Upper SiwaUks, a fact of

which the present author was unaware when work was carried forward. The Tatrot, which the present author
his

considered as possibly transitional between the Upper Phocene and the Lower Pleistocene, is regarded by Lewis as
an equivalent of the Pinjor horizon. Lewis chooses the name Tatrot for this level, a choice that may not be general-
ly followed, due to the long-established use of the term "Pinjor." Likewise, Lewis' designation of the Boulder
Conglomerate as "TaAAi" may not be generally accepted.

In 1936 de Terra and Teilhard placed the Tatrot and Pinjor in the Lower Pleistocene and the Boulder Con-
glomerate in the Middle Pleistocene, as did Lewis. Since these two authors were concerned only with the Pleisto-
cene of India, their conclusions are not discussed here.

In several recent papers (1939, 1940)-', Pilgrim has defended his views as to the correlation of the
Siwahks, and has offered rebuttals to the arguments of Matthew, Colbert, de Terra and Teilhard, and Lewis.
While admitting as proven fact the North American origins of Hipparion and Equus, Pilgrim nevertheless main-
tains that the general characters of the Chinji and Dhok Pathan faunas prove their pre-Pontian and Pontian
relationships, respectively. This author holds that Matthew's and Coll)ert's idea that the Siwalik faunas are
homotaxially similar to but correlatively later than comjjarable faunas in Europe is untenable. Therefore, in
order to explain the presence of Hipparion in the ('hinji and to retain this formation at a pre-Pontian level.

Pilgrim suggests that the various Lower Pliocene formations of North America may be actually older than has
hitherto been admitted by American [lalaiontologists.

Pilgrim disagrees with Lewis' procedure of making the Dhok Pathan an equivalent of Roussillon and Montpel-
Her and placing these latter in the Plaisancian. As he shows by his arguments, these European faunas are of
Astian age, while the Plaisancian represents a general faunal gap between the Lower and the Ui)per Pliocene in

'Stirton, R. A., 1933. Amer. Journ. Sci., (5), XXVI, p. .570.

^Pilgrim, G. E., and A. T. Ilopwood, 1939. Rcc. Gcol. Surv. India, I-X.\III, Tt. 4. Pilgrim, 1939.1. Pal. Iiidica, N. S., XXVI; MilO.l, Gool. Mag.,
LXXVII.pp. 1 27.
 GEOLOGIC SUCCESSION: THE ORIENT 1447

Europe. Pilgrim further maintains that the Tatrot is a truly distinct horizon, not closely related to the Pinjor as
considered by de Terra and Teilhard, or identical with it as claimed by Lewis.

In the light of these recent discussions, Pilgrim presents his latest correlation of the Siwalik Series as follows:

Pleistocene fPost-Cromeriaii Boulder Conglomerate

\Villafranchian Pinjor

Pliocene / Astian Tatrot

\Pontian Dhok Pathan
[Sarmatian Nagri
Miocene ^Tortonian Chinji
[Tortonian Kamlial

In conclusion, it may be said that the Siwalik Series of India represent a period of deposition ranging from
the middle or upper part of the Miocene well into the Pleistocene. The lowest Siwalik horizon, the Kamlial
formation or zone, is definitely of pre-Pontian age. Above this is the Chinji formation, carrying Hipparion, which
may be correlated as either Pontian or as representing a period of time immediately preceding the Pontian,
according to the manner in which the origin and migrations of Hipparion from North America are interpreted.
Following the Chinji is the Nagri formation, transitional between the underlying Chinji and the overlying Dhok
Pathan. The Dhok Pathan is Pontian or post-Pontian, according to the manner in which the Siwalik faunas are
interpreted in relation to Eurasiatic and North American faunas. Between the Dhok Pathan and the succeeding
Upper Siwaliks is a break, representing an Upper Phocene interval during which no sediments were deposited.
The Upper Siwaliks are composed of two horizons of Lower Pleistocene age (Tatrot plus Pinjor) separated by an
erosional break from the uppermost Boulder Conglomerate (Tawi of Lewis), possibly of Middle Pleistocene age.

—
Perim Island. The fauna fromPerim Island was originally considered by Dr. Pilgrim to be equivalent to
the Lower Siwahk fauna. Subsequently, however, he changed his opinion and placed the Perim Island assemblage
in the Dhok Pathan stage. This latter procedure would seem to express correctly the true relationships of the
Perim Island beds.

4. THE PLEISTOCENE OF CENTRAL INDIA

Narbada-Godavari Alluvium. —Extensive deposits of Pleistocene age are exposed in Central India in the
Narbada and the Godavari valleys. These sediments are approximately contemporaneous with each other
(Pilgrim, 1905) and they represent, at least in part, the final stages of the Pleistocene in India.

The Narbada and the Godavari rivers have been known for many years, and from
alluvial deposits of the
time to time fossil mammals, definitely related to the Upper Siwahk fauna, have been found in them. Consequent-
ly various papers have been pubhshed in which these sediments and their contained fossils are discussed. One of

the latest contributions is that of de Terra and Teilhard (1936), in which the problem of the Upper Siwahk and
later Pleistocene deposits of India is considered.

These authors find that in northern India the Boulder Conglomerate stage is followed by a long erosion inter-
val, after which there comes the Potwar silt, a fine deposit, partly of fiuviatile, partly of eolian origin, of Upper
Pleistocene age. The Potwar silt is succeeded by another long erosion interval, while finally, at the top of the
section, are redeposited Potwar sediments and loesses.

Turning now to the Narbada valley, de Terra and Teilhard' make the following remarks as to the Narbada
alluvium.
'Terra, H. de, and P. Teilhard do Chardin, 1936. Proe. Amer. Philos. Soc, LXXVI, No. 6, pp. 820-822.
1448 OSBORN: THE PROBOSCIDEA

Pkoboscidea:
 GEOLOGIC SUCCESSION: THE ORIENT 1449

The formation consists hero [Narhada valloyl of two different horizons, each of which begins with a basal gravel overlain
by brown and pinkish or f)range coloured concretionary clays and silts. In the lower zone the conglomerate is coarser and more
cemented, the clay is more intensely coloured and also richer in concretions than in the upper zone.
Fossils occur chiefly near the disconformity which separates both zones. . .

The 'lower zone' of theNarbadda Pleistocene can be equated with the Upper Siwalik 'Boulder Conglomerate' on faunistic,
archffiological and lithological grounds. The association of advanced Elephas with Hippopotamus and large Bos suggests a stage
slightly younger than the older Upper Siwaliks. In harmony with this is the appearance of an early Paleolithic culture in the
basal gravel, clearly calling to one's mind the picture of heavy accumulation of river deposits during the glacio-pluvial stage of
late Siwalik times in the Punjab. It follows that on these grounds a further correlation between the 'Upper Zone' and the
'Pot war becomes rather plausible. Both are .separated by a long erosion interval from the underlying beds and both contain
silt'

implements of Soan type. The cotton .soil might then well represent the latest Pleistocene which possibly is homotaxial with the
redeposited Potwar .silt and the second loess in the Punjab.

According to these authors, the relationships of the Narbadda aUuvium might be represented in the following
manner.
1450 OSBORN: THE PROBOSCIDEA

5. THE PLEISTOCENE OF CEYLON

Ceylon. — During the years 1935 and 1936 Upper Siwalik fossils were discovered in Ceylon, and described by
Deraniyagala.' These fossils are extremely fragmentary but are sufficiently complete to show that they are
true Upper Siwalik forms. The faunal list is as follows.

Palseoloxodon
Rhinoceros
Hexaprotodon fsivalensis

6. THE PLEISTOCENE OF BURMA

Irrawaddy Valley (Irrawaddy Series). —The Cenozoic history of Burma may be summed up by saying
that there was continuous subsidence accompanied by continuous deposition. Consequently the sediments have
accumulated to a great thickness, with but minor breaks in the series. Of course there were variations in the rate

of subsidence and deposition, causing a considerable amount of interfingering of marine and continental deposits,
but on the whole since Middle Eocene times a gradual encroachment of the continental beds from the north to
the south took place, thereby causing the Burmese gulf to retreat southwardly.

A series of deposits represent the Tertiary in Burma, of which the Pondaung sandstone of Upper Eocene age
and the Pegu beds of Oligocene to Pliocene age are the continental, mammal-bearing facies. Correlative with the
upper portions of the Pegu beds are the lower phases of the Irrawaddy series. These Irrawaddy beds constitute
the upper part of the sedimentary section in Burma, ranging from the Lower Pliocene up through the lower phases
of the Pleistocene.

Fossils are fragmentary and scarce in the Pegu beds, and it is only in the Irrawaddy series that proboscideans
are found.

Two faunas have been identified in the Irrawaddy beds, a lower one of Pliocene age, probably correlative
with the Dhok Pathan fauna of the Siwaliks, and an upper one of Pleistocene age, correlative with the Pinjor
fauna of the Siwaliks. The two faunas are separated from each other by a vertical range of about 4500 feet of

sediments.

There is some question Lower Irrawaddy beds. Pilgrim, in

as to the actual presence of proboscideans in the
1910, listed the mastodont, Stegolophodon latidens, as belonging to the Lower Irrawaddy fauna, but this occurrence
was questioned by Stamp in 1922.- Subsequent work in Burma would seem to indicate that this animal is a member
of the Upper Irrawaddy fauna; certainly a number of discoveries made in recent years establish it as definitely
belonging in the Lower Pleistocene of Burma. Therefore, since its presence in the Lower Irrawaddy beds is very
doubtful, it will be considered here as limited to the Pleistocene in Burma.

The Upper Irrawaddy fauna is in all respects an eastward extension of the typical Upper Siwalik fauna of

India. In numerous cases there is a specific identity between elements constituting the Burmese faunas and
those elements in the Pinjor fauna of northwestern India. Where species are not identical in the two regions,
they are so closely related as to leave Uttle doubt as to their derivation from common ancestral types. Therefore
there is no reason to regard the Upper Irrawaddy fauna as other than of Lower Pleistocene age, strictly correlative
with the Pinjor fauna to the west.
'Deraniyagala, P. E. P., 1935. Journ. Roy. Asiatic Soc. (Ceylon), XXXIII, No. 88; 1936, Geol. Mag. (London), LXXIII, No. 805.
='Stamp, L. Dudley, 1922. Geol. Mag. (London), I-TX, No. XI, pp. 481-501.
 GEOLOGIC SUCCESSION: THE ORIENT 1451

That there was an extension Upper Siwalik fauna eastwardly throughout a considerable portion of the
of the

Oriental region, is shown by the presence of characteristic Siwahk types in various East Indian Islands, to the
south and east of Burma. This extension of the Upper Siwalik fauna has been designated by von Koenigswald' as
the "Siva-Malayan" fauna, having its origin in northern India, pushing into Burma and reaching Java and adja-
cent regions in Lower Pleistocene times.

Proboscideans:
Stegolophodon latidens (Clift)
Stegodon elephantoides (Clift)
Stegodon insignis birmanicus Osborn
Hypselephas hysudricus (Falconer and Cautley)

Shan Plateau (Mogok Fissures). —The Irrawaddy faunas of Burma are found in the Irrawaddy River
valley, in stream and flood-plain deposits. As contrasted with this, is the fauna found in the limestone caverns or
fissures of the Shan Plateau. This fauna is distinguished by Stegodon orientalis, Palseoloxodon namadicus, Ailuro-
poda baconi, and Hystrix. These are essentially the very forms that characterize the mammaUan assemblages
found in Hmestone caverns and fissures to the east of Burma, in Yunnan, Shansi, and Szechwan. There are other
mammals also in the Burma caverns that show the close relationships of these deposits to the cave deposits of
China, animals such as Rhinoceros, Sus, and various ruminants. But it is upon the basis of the first four forms
enumerated that the affinities of tliis cavern fauna in various regions may be established.

Of late years, particularly because of the work of Teilhard and de Terra, Pei, Bien, von Koenigswald, and
others, there is a tendency to regard the cave faunas of China as belonging to a Middle Pleistocene stage of de-
velopment. In Burma there is the Upper Irrawaddy fauna which seems to be definitely older than the mammals
from the Mogok caves, while in China there are such assemblages as those of Nihowan and Ma-Kai, which would
seem to precede the cave faunas. In short, the cave faunas represent a post-Villafranchian development through-
out the Orient.

So far as Burma is concerned, the Mogok fauna may be compared with the Narbada assemblage on the west,
as well as with the cave faunas to the east. But an analysis of such a comparison will show that although certain
Narbada elements are to be found in the Mogok fauna, the bulk of the assemblage is closely related to, if not
identical with the cave faunas of China.

Of course the most important correlations are with the Choukoutien fauna of North China, containing
Sinanthropus, and the Trinil fauna of Java, containing Pithecanthropus.

Von Koenigswald, recognizing the continuity over a broad area in the Orient of this Middle Pleistocene fauna,
designated it as the "Sino-Malayan" fauna. Seemingly having had its origin in the southern part of China, the
Sino-Malayan fauna spread to the south to the East Indian Islands, and to the west to Burma. Consequently the
two Pleistocene faunas of Burma represent invading assemblages, the Lower Pleistocene or Upper Irrawaddy
fauna coming in from the west, and the Middle Pleistocene or Mogok fauna coming in from the east.

Proboscideans:
Stegodon orientalis Owen
Palxoloxodon namadicus (Falconer and Cautley)

'Koenigswald, G. H. Ralph von, 1938-1939. Peking Nat. Hist. Bull., XIII, Pt. 4, pp. 293-298.
1452 OSBORN: THE PROBOSCIDEA

7. THE PLEISTOCENE OF SOUTH CHINA

SzECHWAN Province, China. —In southwestern China, in the province of Szechwan, are numerous pits and
fissures developed in the Umestone that forms the surface outcrops, and many of these pits contain rich deposits of

fossil mammals. They have been worked by the Chinese since ancient times for their "dragon bones," which
constitute one of the staples of the Chinese drug trade.

The fossils from the Szechwan pits were first described by Owen, who regarded them as of Pliocene age.
Subsequently various authors, notably Koken, Schlosser, and Matsumoto, described fossils from the Szechwan pits.

Finally, in 1923, Matthew and Granger' described a new and a very complete series of fossils from Szechwan
(collected by the latter author). At the time, these authors retained the older viewpoint of an Upper Pliocene age
for the material with which they were working. Subsequently, however, Matthew came to the conclusion that

the Szechwan fauna is more properly to be placed in the Lower Pleistocene, a view that was never published, but
clearly stated in manuscript notes.

Perhaps the most important mammals in the Szechwan fauna are Stegodon orientalis, a large stegodont with
very long grinding teeth, a gibbon, Bunopithenis, a langhur monkey, a large Hystrix, a dhole related to the modern
Indian dhole, a giant panda, a gigantic tapir, Megatapirus, a chalicothere, Nestor itherium sinense, a Rhinoceros, and
an extremely large gaur, BIbos geron.

The generally modern character of this fauna places it as definitely of Pleistocene age, for there are too many
advanced elements in it to allow its inclusion in the Pliocene. The association of Stegodon orientalis, Hystrix, and
the giant panda at once estabhshes a strong resemblance to the Mogok fauna of Burma, discussed on a preceding
page. Indeed, upon the basis of the entire faunal assemblage at Mogok and in the Szechwan fissures, there is
every reason to think that they are contemporaneous, representing essentially a single fauna stretching from
Burma through southern China. As has been pointed out in the discussion of the Mogok fauna, there is good
reason to think that these cave faunas are of Middle Pleistocene age since they are preceded both in Burma and
in China by faunas having a Villafranchian aspect.

I take the liberty to quote at this place from Matthew's unfinished manuscript on the Szechwan fauna:

This is a fairly typical fauna of southeastern Asia, phis a number of species now e.xtinet or hmited in their range. Marked
features are the presence of Stegodon, but no mastodons or mammoths,''-' the giant tapir, and a rhinoceros, l)ut no horses, a single
tooth of Chalicotherium. Among the Carnivora is a hyaena related to the spotted hyaenas, Cyon but no Catiin, a very large marten
as big as the American fisiier Maries pennanti, and a tiger, civet, particolored bear and true bear not very different from modern
survivors. The artiodactyls include muntjac and sambhur gazelle, serow, gaur and yak, also some other species which we
have not yet succeeded in identifying; they may be extinct species. The rodents are almost wholly a large bamboo rat allied to
the Chinese species but as big as the Malayan.
.\ltogether this fauna appears to be the fauna of South Chinese forests and mountain valleys, as we may suppose it to have
been before civilized or semicivilized man cut down the forests, cultivated the valleys, and brought about the extinction of the
larger and more specialized animals, driving the remainder of the fauna into the hills.
The absence of horses, of true dogs, of mammoths and of mastodons is in nuu'ked contrast to the Pliocene fauna of North
China, where all tiiose animals had already appeared. The latter was apparently a plains fauna.

Teilhard, Young and Szechwan and adjacent localities as

others,' in 1933, indicated the fissure dejoosits of

extending over a period of time in the Lower Pleistocene that embraced both the Sanmenian and the Choukoutien
dejjosits. In another part of this same paper, they indicated that the fi.ssure deposits of South China might be
limited to a period of time contemporaneous with the Choukoutien deposits. This is their latest opinion, and is

now generally held by authorities on the Pleistocene mammals of Asia.

'Matthew, W. D., and Walter Granger, 1923. Bull. Amer. Mus. Nat. Hist., XLVIII, Art. XVII, pp. 563-598.
'Pala-oloxodon nainadicus has recently been discovered in Szi^cliwan, and described by Yoiiiiji;.
'Black, Teilhard de Chardin, Voung and Pei, 1933. Mem. Geol. Surv. China, Ser. A, No. 11, Table 111, p. 158; Map III, opposite page 164.
 —

GEOLOGIC SUCCESSION: THE ORIENT 1453

The mammalian faunas from other caverns in southern China are so similar to that of the Yenchingkou pits
as to need no particular consideration at this place. It is quite evident that all of these occurrences represent

a single fauna that was rather widely spread during Middle Pleistocene times.

Mention should be made of the caves in Kwangsi, the fauna of which has been described by Teilhard, Young,
Pei and Chang, and the Hoshangtun Cave in Yunnan, described by Bien and Chia. Also there should be mention-
ed the isolated occurrence of a stegodont said to have been found near Shanghai and described by Owen as Stegodon
sinensis. This is probably of the same age as the other South Chinese Stegodonts.

Proboscideans:
Stegodon orienlalis Owen
Palxoloxodon namadicus (Falconer and Cautley)

8. THE PLEISTOCENE OF INDO-CHINA

French Indo-China. —There are cavern deposits in northern French Indo-China, seemingly quite similar
to those of southern China, described above. Since they contain the same fauna as the Chinese caverns, they
must be considered as correlative with the latter, representing a southern extension of the characteristic Middle
Pleistocene cave fauna of the Orient. Mansuy,'who described the mammalian fossils from Indo-China, identified
the stegodont material as belonging to Stegodon "clifti" and Stegodon insignis. A comparison of his excellent
plates with material on hand suggests the possibility that the Stegodon from Indo-China is of the species orientalis.
In fact, Mansuy regarded S. orientalis as probably synonymous with *S. insignis, and this view undoubtedly
influenced him in his identification of the material.

Proboscideans:
Stegodon orientalis Owen. {S. insignis according to Mansuy.)
Stegodon elephantoides (Clift). — (This species? probably orientalis.)

Palxoloxodon namadicus (Falconer and Cautley)

9. THE PLEISTOCENE OF THE EAST INDIES

Java. — Since the discovery of Pithecanthropus in 1892, Java has assumed a position of great importance to
the students of human prehistory and the evolution of Ufe. Consequently numerous contributions have appeared
during the course of the past forty years, discussing the phylogenetic relationships and the probable geologic age
of Pithecanthropus. It has been only recently that detailed geological investigations have been carried on in
Java, with a view to clarifying some of the hitherto obscure points as to the succession of upper Cenozoic sedi-
ments in Java.

For many years was supposed that the beds at Trinil, in which Pithecanthropus was discovered, might be
it

placed in the Upper Pliocene or the Lower Pleistocene, thereby allocating to Pithecanthropus the honored position
of being the oldest known hominoid. But recent studies by various authors have shown that the upper Cenozoic
stratigraphy of Java is much more complex than it was originally thought to be therefore our ideas as to the age —
of Pithecanthropus and the associated fauna have necessarily undergone considerable revision.

In 1931 van Es published a detailed geologic study entitled "The Age of Pithecanthropus," in which he
showed that there was a volcanic boulder breccia and beneath it a sand and conglomerate underlying the Trinil
beds. Both of these older horizons he regarded as of Pleistocene age.

'Mansuy, H., 1916. Mem. Surv. Geol. Iiidochine, V, Fuse. II, pp. 1-26.
1454 OSBORN: THE PROBOSCIDEA

In 1932 van der Maarel' described a new fauna from Bumiaju, in Central Java, and came to the following
conclusions as to the stratigraphic relationships of the Javanese strata:

"1. The Bumiaju fauna is older than the Trinil fauna.

2. The Trinil fauna is certainly of Pleistocene age, more particularly either Lower [or] Middle Pleisto-
cene, but not Upper Pleistocene.

3. Accordingly the Bumiaju fauna is of upper pliocene or lower pleistocene age."

In recent years von Koenigswald- has carried on a series of careful studies regarding the upper Cenozoic
mammal-bearing sediments in Java, and he has distinguished a number of successive horizons within the Pleisto-
cene. These and correlative horizons in India may be listed as follows:
 GEOLOGIC SUCCESSION: THE ORIENT 1455

The faunas of the several Javanese Pleistocene horizons are too extensive to list here, so only the probo-
scideans will be enumerated.

Proboscideans:

Sampoeng zone
Elephas indicus Linnseus

Ngandong zone
Stegodon trigonocephalus Martin
Palseoloxodon cf. namadicus (Falconer and Cautley)

Trinil zone
ICryptomastodon martini von Koenigswald [
= Sirenian? (Osborn)]
Stegodon trigonocephalus Martin
Palseoloxodon cf. namadicus (Falconer and Cautley)

Djetis zone
Stegodon trigonocephalus praecursor von Koenigswald
Elephas sp.

Kali Glagah zone

Tetralophodon bumiajuensis Maarel
Stegodon trigonocephalus praecursor von Koenigswald
Archidiskodon planifrons (Falconer and Cautley)

Tji Djoelang zone

Stegodon sp.

Borneo. — In 1885 Lydekker' described a tooth that he identified as Mastodon latidens, from the northwest
coast of Borneo.

The specimen forming the subject of the present notice was forwarded from Borneo to the Secretary of this Society by Mr.
A. H. Everett, C.M.Z.S., who stated that it was found during the early part of the present year by a Kadayan in the jungle in
the vicinity of Bruni, on the north-west coast of Borneo. Owing to the country being in a disturbed state Mr. Everett could
not visit the locality to make further inquiries but there seems no doubt that the history of the specimen is a true one.
;

Comparing this tooth with the many teeth of Mastodon latidens from Burma and India, Lydekker came to the
conclusion that the tooth from Borneo was specifically identical with the Burmese-Indian forms, and therefore he
decided that the deposits in Borneo from whence this tooth was recovered might be questionably of Pliocene age.

Subsequently (1936, Volume I of this Monograph, p. 700) Professor Osborn made the tooth from Borneo the
type of a new species, Stegolophodon lydekkeri.

Stegolophodon latidens from Burma, where the type was discovered, would seem to be restricted to the Upper
Irrawaddy beds of Pleistocene age, while in India the species ranges down into the Pliocene Dhok Pathan horizon.
It is an interesting fact that Lydekker emphasized the close mineralogical comparison between the Borneo tooth
and those teeth of M. latidens from Burma. With this fact in mind, and remembering that the Stegolophodon
latidens in Burma is found in the Pleistocene level, there would seem to be some reason for thinking that the

'Lydekker, R., 1885. Proc. Zool. Soc, London, 188.5, p. 777.

1456 OSBORN: THE PROBOSCIDEA

Borneo specimen might be of Pleistocene age. It should be pointed out, however, that Professor Osborn placed
this specimen in the Pliocene (but with a query) as did Lydokker.

Proboscidean: Stegolophodon lydekkeri Osborn.

Philippine Islands. — On the basis of published records it would seem that fossil proboscideans from the
Philippine Islands are represented by a single tooth from the island of Mindanao. This specimen was described
many years ago by E. Naumann, under the name of Stegodon mindanensis. Naumann compared it with Stegodon
trigonocephalus of Java, pointing out the fact that there were great similarities between the species. Therefore
it would seem possible that the beds containing these two forms in Java and Mindanao are geologically more or
less equivalent to each other. In tliis connection it may be pointed out that Stegodon trigonocephalus in Java is
found typically in the Ngandong beds, above the Trinil zone of probable Middle Pleistocene affinities. It is an
interesting fact, however, that stegodonts closely related to S. trigonocephalus are found in lower beds in Java,
ranging down to the basal Pleistocene. Therefore it would seem that the Mindanao deposits might be repre-
sentative either of Lower or of Middle Pleistocene times in the Phihppine region.

Proboscidean: Stegodon mindanensis Naumann.

IV. EUROPE
1. INTRODUCTION
It may be said that the Proboscidea entered Europe at the beginning of the Miocene —considering the
Burdigalian to be the opening stage of this period. From that time until the final stages of the Pleistocene, these
animals were prominent in the extinct mammalian faunas of the European region ; therefore, a consideration of the

Proboscidea-bearing beds of Europe is virtually a discussion of the continental stratigraphy of the Miocene,
Pliocene, and Pleistocene of that region.

A thorough discussion of the continental Tertiary and Quaternary sediments of Europe is indeed a large
order, too much to be included within the scope of this present work, and beyond the capabilities of one who does
not have an intimate first-hand knowledge of the complex relationships of these Old World mammal-bearing
horizons. Therefore, it is proposed to discuss very broadly the general succession of upper Tertiary and Pleisto-
cene continental deposits in Europe, giving particular attention to the more typical localities at which the several
stages are best developed. In doing this the localities at which proboscidean types were discovered will, of course,

be especially stressed.

In dealing with the Tertiary mammahan faunas of Europe one general consideration must be kept in mind,
namely, that this was primarily a mediterranean area —particularly during the earlier phases of the Tertiary
period —interspersed with numerous islands and peninsulas. In this respect, the European region is to be con-

trasted with America and Asia, which from the end of the Cretaceous were, generally speaking, broad continental
platforms, with the shallow sea borders lapping over the edges, but not in the main encroaching materially beyond
their present limitations. It was not until late Tertiary times that Europe underwent a general emergence so
that most of its present area was permanently lifted above sea level. Naturally, the broken-up character of the
land areas of Europe in earlier Tertiary times had a distinct influence on the development of the mammalian
faunas, and similarly the final emergence of this region as a continental mass also is reflected in the expression of
 GEOLOGIC SUCCESSION: EUROPE 1457

the assemblages of land animals. So it is that the earlier separated faunas are finally replaced by associations of
mammals that become more and
more cosmopolitan, with an eventual cuhnination in the appearance of the
almost world-wide Hipparion faunas and subsequent assemblages that are so characteristic of the last phases of
Tertiary times.

^EOl£>GICA.L KEIJLTIOI/SHIPS OF EUROPEA.N^ PKOBOSCIDEA.

FRANCE ITALV AND CENTRAL EUROPE EASTERN EUROPE ASIA MINOR
BELGIUM W MEDITERRANEAN GERMAN\' RU55IA GREECE
ENGLAND SWITZERLAND SPAIN ISLAND5 AUSTRIA POLAND E, MEDITERRANEAN
HUNGARY RUMANIA ISLANDS
Hesperoloxodon Mammonteus pnmiaenius Hesperoloxodon antic^u us Parelephas ujiisK Palaeoloxodon
anttquus hudrunfinus
Mammonteus primiaeniu; platurhunchus Loxodonta cornaliae
qerma icus
Hesperoloxodon
ci^priofes
Pal^o/oxodon
(Widelu disfvibutccj over Europe) Pajieoloxodon mnaidriensis. antiquus aermanicus creticus
falconcri, melifcnsis, lamarmorae
Hesperoloxodon antiquus tralicus
Parelephas irvrermedius Parelephas troqontherii Mastodon pavlotoi Parelephas
Mammonteua primiaenius armentacus
He s pe ro odon
I O.X aa ti q u u s rraasi
[Wideli^ distributed over Europe) if t< leith-adamsi

Parelephas Parelephas troqontherii

•5 *
troao
ontherii Mammonteus
nestii
primiaenius
rraasi
« *• leith-adamsi

¥'
GO//Z Archidiskodon
meridionalis
Anancus arvernensis Archidiskodon meridionalis
Parelephas frogontherioides
Archidiskodon
planifrons rumanus
cromerensis Mammonteus primiaenius astensis
Hesperoloxiodon antiquus ausonius
a (t nanus
';yz/x-
'^KAN'CHJAh
Anancus arvernensis Zuqolophodon borsoni Zuqolophodon borsoni laddach
Anancus a a brcvirostris
Anancus ^i^antarvernensis
ASTIAJi falconcri « minute arvernensis
(c « dissimiJis
tt u macroplus
Zuqolophodon borsoni buffonis
a (( vellavus
Turiclus viraatidens Dinotherium
PZA/S- (t ** vialeti Steaolophodon aiaantissimum
subiatidens
AHC/ATi

Dinotherium medium Dinotherium aiaanteum. Dinotherium proavus Trilophodon

Turret usluahlneimensis n podolicum (Choerolophodon)
Miomastodon tap iro ides
(4 uralense pentelicus
^, amencanus
rliomastodon amencanus Anancus arvern- Turicius atticus
r/A/f praett^pica ensis pro^ressor Tetralophodon
TIrilopnodon
IU A esselbornensis Anancus qrandincisivus
Tetralophodon lonairostris intermedi us
« ^iaantorostris
SARMA- Platubelodon danovi
T/Ay
Dinotherium intermedium Dinotherium. bavaricum
Dinotherium levius « secundarium
Zuaolophodon purenaicus Trilophodon steinheimensis
VINIDO- Turicius turiccnsis « enaelaujiesensis
BO/TfA^
<* « simorrensis « anaustidens minufus
Trilophodon anaustidens tt {£ austro-qermanicus
(t It ^aujaci Serridentinus sub^api^oideus
it it aaillardi

Dinotherium cuvieri |

Zt^qolophodon pijrenarcus aurelianensis

Turicius tapiroides '

BURDl-
Miomastodon depe re ti
GAUAX Trilophodon pontileviensis
a anaustidens cuvieri Dinotherium hunqaricur
Serridentinus Tiiholi I

DFjeyett RraJJeij

Fig. 1222

2. THE LOWER MIOCENE: BURDIGALIAN

It is not the purpose in this exposition to go into the question as to whether the Aquitanian is more properly
to be considered as the closing stage of the Ohgocene or the opening stage of the Miocene, for this horizon
precedes the appearance of proboscideans in Europe, and its consideration here would be fruitless. Suffice it to
say, therefore, that the Miocene may be considered for the purposes of convenience to have opened with the
Burdigalian.
1458 OSBORN: THE PROBOSCIDEA

The development of Burdigalian mammals in Europe is perhaps best exemplified by the fauna of the Sables
de rOrleanais of central France. This fauna, known since the time of Cuvier, has long been regarded as typifying
the BurtUgalian stage in the European region, but in recent years as knowledge of the stratigraphy and palaeon-
tology of these beds has become more exact, there have arisen the inevitable differences of opinion as to the
exact relationships of the several horizons and their contained mammalian faunas within the Sables de I'Orleanais
each to the other. The problem was stated by Mayet' in 1908 as follows.

Les sables de I'Orleanais ne sont pas une formation g^ologique ayant la remarquable unite qu'on lui a longtemps atri-
buee. A priori, il etait a supposer que cette masse enorme d'alluvions granitiques qui couvre encore actuellement une partie du
centre de la France, avait exige un temps fort long pour etre form^e, charri^e, deposee. Les renseignements stratigraphiques et
les donnees paleomammalogiques me paraissent confirmcr cette induction. On pent distinguer dans le Burdigalien de I'Orleanais,
dont I'expression 'Sables de I'Orleanais' est pour ainsi dire synonyme, plusieurs facics locaux, d'agc tres probablement different.

It is Mayet's contention, and in this he follows the lead of his illustrious countryman. Professor Deperet,
that the Burdigahan of central France is inaugurated by the Calcaire de Montabuzard, underlying the Sables de
I'Orleanais,and between these two horizons he would place the Sables de Chitenay. Thus, his Burdigalian
section would be, generally speaking, as follows:

Sables de la Sologne
Marnes de I'Orleanais
Sables de I'Orleanais
Marnes du Blesois
Sables de Chitenay
Calcaire de Montabuzard

Mayet has envisaged the Burdigalian in central France as beginning with the deposition of sediments in
a vast lake, the "lac de Beauce." The Calcaire de Montabuzard is a local facies of this deposition, and as such
contains a mammalian fauna which Mayet finds to be essentially more primitive than the typical Sables de
I'Orleanais fauna. The Sables de Chitenay are also regarded by Mayet as constituting a local facies of the basal
Burdigahan, deposited by a large Miocene river which flowed from the Central Plateau into the lac de Beauce at
about the time the deposition of the Calcaire was coming to an end. Either contemporaneous with or immediately
succeeding the sands of Chitenay are the Marnes du Blesois. Then come the Sables de I'Orleanais in the strict
sense of the word, with the large mammalian fauna typical of these sands. And finally at the top of the section
are unfossiliferous beds, the Marnes de I'Orleanais and the Sables de la Sologne.

A much different interpretation of the Burdigahan and associated faunas of central France was put forward by
Stehhn in 1908.'- A decade or so before this time, Deperet had called attention to the very important fact that the
fauna of the Sables de I'Orleanais contains two elements, one consisting of indigenous mammals, derived directly
from the Aquitanian manmials of the same region — the other being composed of immigrant forms coming in from
African, Asiatic, and American centers of origin. Stehlin elaborated on this thesis and attempted to show that the
relative abundance of inuTiigrant forms as compared with indigenous types is indicative to a certain extent of the
general advancement in age of the several faunas within which they are contained.

Following this line of thought, Stehlin came to the conclusion that the fauna of Chitenay is the most archaic
of the Burdigalian mammalian assemblages, because it contains tlie smallest percentage of immigrant forms.
Thus, he would designate the Chitenay fauna as lower Burdigalian, intermediate in position between the upper
Aquitanian faunas, such as Saint Gerand-le-Puy and the characteristic middle Burdigahan fauna of the true Sables
de I'Orleanais.
Mayet, L., 1908. Aim. Univ. Lyon, N. S., I, Fasc. 24, p. 313.
Stehlin, H. G., 1908. Bull. Soc. gcol. Franco, (4), VII, p. 545.
 :

GEOLOGIC SUCCESSION: EUROPE I459

Perhaps the greatest discrepancy between the views of StehHn and those of Deperet, Mayet and others, is
his assignment of the Calcaire de Montabuzard to a position not below the Sables de I'Orleanais, but rather to an

elevated place considerably above the Sables. Here again StehUn has based his views on the fact that there is
a very large immigrant element in the Montabuzard fauna, as compared with the indigenous mammals. Thus,
this authority would regard the Montabuzard assemblage as being perhaps equivalent to or slightly later than the
typical Sansan fauna. Stehlin admits, however, that

Si, un jour, on decouvre une faunule de Mammiferes dans quelque lambeau de calcaire de I'Orleanais
indubitable, c'est-
a-dire directement recouvert par des sables fluviatiles burdigaliens fossiliferes, la question de Montabuzard sera tranchee.

The locaUties at wliich Burdigalian faunules have been found in central France are numerous and are Usted
by Deperet, by Mayet, and by StehUn, to which authors the reader is referred. Mayet, particularly, has given
detailed discussions of the important locaUties at which Sables de I'Orleanais deposits containing mammaUan
remains have been discovered.

As to the fauna of the Sables de I'Orleanais, the foUowing forms might be accorded particular attention:

Pliopithecus— according to Stehlin present in the Faluns de Pontlevoy and not in the Sables de I'Orleanais,
strictly speaking. But Mayet placed this genus in the Sables, as weU as in the Faluns. Certainly an
advanced anthropoid, that might be of Middle rather than of Lower Miocene affinities.

Amphicyon —typically Miocene.

Pseudselurus— also typically Miocene.

Steneofiber—a characteristic Lower Miocene form.

Proboscideans — aU essentially primitive — Lower to Middle Miocene.
Brachyodus — a persistent type from the Oligocene.
Palseochoerus — another persisting Oligocene form.

Listriodon— characteristic of the Miocene. L. lockharti of the Sables is a very primitive member of the genus,
and might be considered as indicative of the beginning of the Miocene.
Palaeomeryx, Amphimoschus, Procervulus, Dicrocerus — all primitive Miocene cervuUnes.
Anchitherium — the appearance of
earliest genus in Europe. this

Macrotherium — quite characteristic of the Miocene, and may occur as early as the beginning of the period.
it

Dicer atherium — genus? tliis

Aceratherium, Teleoceras — again typical Miocene forms.

It might be well to mention here some important locaUties in addition to those discussed, namely, Chitenay
CheviUy, Eggenburg in Lower Austria, and Brlittelen in Switzerland, at wliich latter place a shoreUne facies has
been discovered, with an association of marine forms.

In a recent paper (1934) by Roman and

BurdigaUan fauna from La Romieu is described. This
Viret,' a
fauna is found to be contemporaneous with the upper BurdigaUan of the Sables de I'Orleanais, particularly as
characterizedby the faunas of Baigneaux and of CheviUy. In this connection, it might be said that these authors
make some interesting comments as to the characters of the upper BurdigaUan fauna in general, as compared with

'Roman, F., and J. Viret, 1934. Mem. Soc. gool. France, N. S., IX, No. 21.
 : :

1460 OSBORN: THE PROBOSCIDEA

the lower Burdigalian assemblage below, and the Helvetian fauna above. In short, the conclusions of Roman and
Viret as to the characters of the upper Burdigalian fauna or faunas of Central Europe are as follows:

The upper Burdigalian fauna differs from the older lower Burdigalian fauna in that:

There are more of Brachyodus onoideus, Amphitragulus, Palseomeryx garsonnini, and typical Brachypotherium
aurelianense.

The following forms appear for the first time

Listriodon lockharti, Hemicyon, near H. goriachensis, Pseudselurus quadridentatus, Lagomeryx of small size,
Dorcatherium, Dinotherium, as characterized by the small D. cuvieri, and a large Brachypotherium.

The upper Burdigalian differs from the younger Helvetian in that:

There have not as yet appeared such forms as Pliopithecus, Machairodus, Ursavus, Micromerijx, Potamo-
therium, and above all the antelopes, which latter appear suddenly in the Middle Miocene of Sansan.

Anchiiherium aurelianense is present but as a small mutation — that is, it has not attained the size of the
typical later Miocene Anchitherium. The same is true of Steneofiber. Palasochoerus is present but not Hyotherium.

The jjroboscideans of the BurdigaUan are as follows

Dinotherium cuvieri Kaup —

Type Chevilly
Turicius tapiroides (Cuvier) —
Type Calcaire de Montabuzard
Turicius turicensis (Schinz) (According to Mayet, type from vSarmatian)
Dinotherium bavaricum von Meyer (According to Mayet, type from Helvetian)
Miomastodon depereti Osborn Type — Chevilly
Trilophodon pontileviensis Mayet and Fourtau Type Pontlevoy —
Trilophodon angustidens cuvieri (Pomel) Type Gers —
Serridentinus filholi Frick Type Gers —
Zygolophodon pyrenaicus aurelianensis Osborn Type Chevilly —
Dinotherium hungaricum Ehik Type Kotyhaza, Hungary —

3. THE MIDDLE MIOCENE: HELVETIAN AND TORTONIAN = VINDOBONIAN

Helvetian. —The Middle Miocene of Europe, the Vindobonian, may be divided into two stages, the Helve-
tian and the Tortonian, each typified by distinctive mammalian faunas. The characteristic Helvetian fauna is

that of Sansan.

The Sansan fauna, discovered by Lartet, and studied by this same authority and by Milne-Edwards, and
later by Filhol, comes from a hill near the little village of Sansan in southern France. At this place there is a suc-
cession of freshwater sediments, begirming with a clay bed at the bottom, followed by a calcareous layer, which in

turn is capped by a series of marls containing freshwater molluscs and numerous fossil vertebrates —particularly
mammals and birds.

In contradistinction to the preceding Burdigalian fauna, as developed in the Sables de I'Orleanais, the Sansan
assemblage is definitively a more modernized fauna, with no important jjersisting Oligocene types, which were so
common in the Burdigahan faunas. Indeed, the Sansan fauna is characterized by the appearance of many mam-
 :

GEOLOGIC SUCCESSION: EUROPE 1461

malian forms indicative of the beginning of evolutionary developments that culminated in the modernized faunas
of uppermost Cenozoic and Recent times.

The origin of the Sansan deposits was discussed by Lartet, whose conclusions were substantiated by Filhol.

Lartet had the following to say about the genesis of these beds

Toutes COS circonstances m'avient porte a penser que I'accumulation successive des fossiles organiqucs qui coiistituc le
d^pot de Sansan se serait effectu^e au fond d'un marais, ou si Ton veut d'un petit lac ou auraient vecu les especes aquat-
iques dont on retrouve des ddbris; tandis que les eaux torrentielles auraient pu entrainer dans ce bas-fond, soit des osse-
ments dispers6s, soit les cadavres entiers des animaux terrestres etablis a demeure ou du moins se montrant passagferement sur
les terres environnantes, emergees pour un temps plus ou moins long.'

L'hypothese de Lartet parait bien justifiee. II y a cu ti Sansan un lac, dont la plus grande partie a disparu, a ete emportee
a I'epoqiie du creusement des vallecs. Dans ce lac venaient se d^verser diff^rents cours d'eau, susceptibles dc deborder a certains
moments et do couvrir probablement d'assez vastes espaces, d'ou ils emportaient les animaux y vivant au moment de leurs
crues subites, pour les deposer en un point oii existait un remous.-

Deperet has discussed at length the composition and relationships of the Sansan fauna, consequently the
fauna in its entirety will not be listed here. But the following important genera contained within the fauna may
be given some consideration.

Pliopithecus —here we see the continuation of a relatively advanced anthropoid, which already has appeared in the

Burdigahan fauna of the Sables de I'Orleanais and the Faluns de Pontlevoy.

Insectivores —several genera, such as Erinaceus, Sorex, Talpa, which are similar to the modern forms.
Rodents — the insectivores, genera leading to modern types. There should be mentioned particularly:
like

Steneofiber — a characteristic Lower to Middle Miocene castoroid. This, or a related type, found in the Lower is

Miocene of North America.

—
Amphicyon a typical Miocene canid, showing advanced characters.
—
Hemicyon one of the more advanced "bear-dogs," distinctly later than Lower Miocene, and found at various
Middle to Upper Miocene locahties in Europe, Asia, and North America.
Machairodus and Pseudselurus —Miocene Fehdse.
Other carnivores are distinctly leading to more modern types.
Anchitherium — the typical Miocene equid of the Eastern Hemisphere.
Macrotherium — similarly, the typical Old World Miocene chalicothere.
Aceratherium— continuation of a form that appeared in the Burdigalian.
first

Listriodon —typical Miocene suid.

Palxomeryx— a very typical element in the Miocene faunas of Europe.
Trilophodon.
Turicius.

Of the European deposits correlative with those of Sansan, perhaps the most important are those of Engels-

wies and Georgensgmlind in Germany, and Eibiswald and Goriach in Austria.

Klahn (1922) describes the Engelswies deposits as chalky, brackish water sediments, probably deposited in an
estuary or the mouth of a river that flowed out into the Middle Miocene mediterranean of Central Europe.

These beds contain a typical Middle to Upper Miocene fauna, characterized notably by Dinotherium, Barchypo-
therium, Aceratherium, Anchitherium, Macrotherium, and Hyotherium. Klahn would correlate this fauna with the
Steinheim assemblage, which would place it somewhat higher than it is regarded in the present work.
'Lartet, E., quoted by H. Filhol, 1891, pp. 5, 6.
^Filhol, H., 1891, p. 6.
 ;

1462 OSBORN: THE PROBOSCIDEA

The fauna from Goriach, described by various authors but particularly by Toula and by Hofmann, would
seem to be a typically Middle Miocene mammalian assemblage. The presence in this fauna of characteristic
Miocene genera, the lack of truly advanced Miocene forms, and the persistence of certain types from the Lower
Miocene seem to favor the correlation of Goriach as approximately of Sansan age. There might be noted here
such genera as Amphicijon, Dinocyon, Hemicyon, Trilophodon, Anchithermm, Steneofiber, Aceratherium, Hyo-
therium, Palseomeryx, Dicrocerus and other primitive deer, all of which relate this fauna very closely with the
Sansan assemblage. The genus Cebochoerus, described by Hofmann as from Goriach, is at best of doubtful refer-
ence, for the material is very fragmentary. One would hardly expect this form in the Miocene.

Others faunas of Helvetian age will compare with the Sansan fauna in about the same way as these, as con-
sidered above.

Proboscideans: See list of Vindobonian proboscideans (p. 1464 below).

ToRTONiAN. —Of the Tortonian deposits in which fossil mammals have been found, none is perhaps better
known than La Grive-Saint-Alban, Isere, in southern France. This locality has been known for many years,
and has been studied extensively by Deperet, who published several large papers dealing with the fauna and
the stratigraphic relationships of that deposit.

Deperet has shown that the mammal-bearing beds of La Grive-Saint-Alban are in reality fissure deposits,

formed during Middle Miocene times, in pits, fissures, and caves that were eroded out of Mesozoic limestones. In
similar pockets at Mont-Ceindre, near Lyons, Deperet discovered such a wealth of bat remains that he was led to
think that perhaps some of the deposits in this general complex or series were formed in Miocene bat caves.

Some fifty years ago this author' described the typical deposits at La Grive-Saint-Alban, as follows (Deperet,
1887, p. 60):

Ces fentes sont toutes ouvertes vers le haut, ct penetrent plu.s ou moins profond^ment dans I'epaisseur du terrain jurassique
la plupart cependant attcignont lo niveavi du sol de la carri^re. Le rcmpiissagp de ces fentes, s'est op^rc, il me somble, par le
haut, et sous I'influenee d'un simple ruissellement a la surface du calcaire; la faible vitesse des eaux phiviales sur ec plateau
horizontal, et la lent(>ur du jihenomeiie de remplissage me jiaraissent suffire a expliquer I'absence si remarquahle, dans les mat6-
riaux charries, de cailloux roules et meme de graviers, qui i)ourraient faire penser a un charriage op6r6 par un cours d'eau de
quel(|ue importance. I.es sculs elements etrangers a I'argile que j'ai observes sont des cristaux spathiques, qui paraissent
resulter de I'evaporation reiietee d'eaux riches cii acide carbonique. Quant a I'argile rouge, je pense qu'elle n'est ici que le simple
rcsidu de la lente dissolution du calcaire, oper^ed'mie maniere continue par les eaux pluviales, pendant la longue emersion du
continent mayencien, et accumulee dans les fentes par les eaux de ruissellement.

A few years later (1892) Deperet- had the following to say as to the age and relationships of the fauna from
La Grive-Saint-Alban.
Ainsi que je I'ai d^ja indiqu^ en 1887, la presence a la drive du Rhinoceio.s brachypvs et du Dinolherium, {\\\\ mantjuent k
Sansan, mais se retrouvent dans I'horizon un pen plus elev6 de Simnrre (Cters) annonce des affinites importaiites ent re la faune de
la drive et celle fie cette derniere localite, sur laquelle nos cormaissances sont malheureusement fort incompletes. (
'e dernier

l)arall61isme est d'autant plus probable que, ainsi que je I'ai d^ja indiqu^, les animaux de la drive, tels que le Pliopllherus, le
Marhairodiis Jourdani, la Lutra Lorteli, le Cricetndnn Rhodaiiiriim, Ic Frotrngocenif! Chanirei paraissent representer dans leur
en.semble vui degr^ d 'evolution legferement plus avanc^ que celui des especes representatives dans le gisement dc Sansan. . . .

Le gisement des sables de Steinheim (Wurtemberg) doit etre consid6r6 comme I'equivalent exact de celui de la (<riv(?. Kn
effet, sur les quarante-.sept especes de la (Jrive, dix-sejit .se trouvcnt aussi a Steinheim. . . .

Les autres gisements de Suisse, d'Allcmagne ou d'Autriclie ((ue Ton ])eut consid^rer (^onunc a pen jtres synchronitiucs de
celui dc la drive-Saint-Alban sont: les lignites dc (loriach (Styric); <lri>rqensqniund; (lUn.sbiirg et If-s sahlrx d Dinothrriiini de
Ba\ierc; KlUi, Kapfiioch Hies (Nordlingcn)
; \'cimcs {.luni liernois)
; mais la faune dc ces diver.ses localit(5s est Ix^aucoup plus
;

pauvre que celle des autres gi.semcnts prdcites.

'Dcporol, C, 1887. Arcli. Mils. hi.st. nat. Lyon, IV, p. fiO.

=Dci)6rct, C, 1892. Arch. Mus. hist. nat. Lyon, V, Pt. 2, i>p. 4, 5.
 GEOLOGIC SUCCESSION: EUROPE 1463

Subsequently, in 1899, Gaillard' published a paper supplementing the studies of Deperet, and describing new
or little known mammals from La Grive-Saint-Alban. Gaillard's conclusions as to the age of this deposit were
essentially the same as those of Deperet. Since the time that Deperet and Gaillard made their detailed studies of
the fauna of La Grive, the general trend has been to substantiate and strengthen the conclusions of these authors
as to the age of this mammalian assemblage.

That the Helvetian and Tortonian stages are very closely related is shown by a comparison of their respective
faunas. Thus, the Sansan and La Grive faunas may be compared, by genera, as follows.

Sansan La Grive-Saint-Alban Sansan La Grive-Saint-Alban

Pliopithecus Pliopithecus Viverra Viverra
Vespertilio Vespertilio Herpestes
Rhinolophus Vespenigo Progenetta
Erinaceus Erinaceus Machairodus Madiairodus
Galerix Galerix Pseudxlurus Pseiidxlurus
Lantharwiherium Aclurogale
Mrjgale Dimylus Trilophodon Trilophodon
Sorex Sorex Turicius Dinolheriuni
Talpa Talpa Anchitherium Anchitherium
Sciurus Sciurus Aceratherium Aceratherium
Cricetodon Cricetodon Macrotherium
Myoxus Myoxus Choeromorus Choeromorus
Meriones? Choerotherrum
Arvicola? Listriodon Listriodon
Steneofiher Hyotheriiim
Lagopsis Hysemoschus Hysemoschus
Prolagus Micromeryx M icromeryx
Hemicyon Hemicyon Palseomeryx Palxomeryx
Pseudocyon Morphelaphus Dicrocerus
Amphicyon Amphicyon Strogulognathiis
Trochitis Trochitis Palseocervus
Mustela Plesidis Cervus
Pidorius? Martes Antilope Prolragocerus
Proputorius Lutra

From this comparative list it may be seen that there is little definitive evidence that enables one logically to
separate the typical Helvetian faunas from those of Tortonian age. Indeed many of these faunas are considered

as of one or the other of these two stages, by different authors, and no two authorities will correlate the Middle
Miocene deposits of Central Europe in exactly the same way. Therefore there is much to be said in favor of
lumping the several Middle Miocene faunas within one time division, the Vindobonian.

Of the other so-called Tortonian faunas,some of the most important are those of Simorre and Villefranche
d'Astarac in France, of Steinheim, Mosskirch, and Oppeln in Germany.

The Steinheim mud, contains the following diagnostic genera:

fauna, preserved in an impure chalky ooze, or
Amphicyon, Viverra, Chalicomys, Aceratherium, Macrotherium, Anchitherium, Listriodon, Dicrocerus, and Palseo-
meryx. This assemblage is generally regarded as of Middle to Upper Miocene age, equal to the La Grive fauna
and perhaps younger than the typical Helvetian faunas. Klahn- speaks of it as follows:

In Bogloitung des unten zu beschroibenden Mast, .steinheimensis deutet die Fauna auf obermioziines Alter bin. Diese ist
dieiselbc wie die von Kgl. Neudorf bci Oppeln und Grive-St. Alban.

The fauna of Simorre is in all respects correlative with that of La Grive-Saint-Alban, and for this reason is
often considered as typical of the Tortonian in Europe. The remarks as to the age and relationships of the fauna
of La Grive apply equally well to that of Simorre.
'Gaillard, Claude, 1899. Arch. Mus. hist. nat. Lyon, VIL
^Klahn, H., 1922. "Die badischen Mastodontcn, etc.," p. 33.
 : ;

1464 OSBORN: THE PROBOSCIDEA

Certain other European faunas, representing the final stages of the Vindobonian or Tortonian, may be con-
sidered here. Of these, particular mention should be made of (Eningen, Saint Gaudens, Elgg, and Monte Bamboli
Kapffnach, Giinsburg, and San Isidro.

The ffiningen fauna, and flora preserved in fine-grained calcareous beds near Zurich, Switzerland, has long
been famous for the excellence and diversity of the fossils comprising it. It is especially characterized by numerous
plants, insects, reptilian and amphibian remains, as well as those mammals. Incidentally, it
of fossil was here that
the type of Andrias scheuzeri was discovered. De Lapparent' makes the following remarks concerning the
(Eningen fossils

On attribue au sommct du tortonien les couches d'ffiningen, sur les bords dvi lac de Constance, calcaires en minces pla-
quettcs, oxtraordinairoment riches en fossiles, insectes, poissons (Leuriscus), reptiles, etc. Mais c'est surtout par sa flore que le
gitc d'CEningen est celebre. Heer en a d^crit pres de 500 especes, parmi lesquclies bon nombrc de formes europecnnes sont melees
a des types asiatiques, af ricains, australiens ou meme americains. L'assise inferieure ou couche k insectes d'ffiningen est compos^e
d'environ 250 feuillets, ou Ton distingue jusqu'aux saisons succcssives: les fleurs de camphrier annon^ant le printemps, les
fruits d'orme et de peuplier I'et^, ceux de camphrier ct de Diospyros I'approche de I'automne. . .

Heer pense qu'il regnait a (Eningen un climat analogue a celui de Madere, du Japon meridional et de la G^orgie, suit une
moyenne annuelle de 18 a 19 degres.

The numerous faunas contemporaneous with or correlative with (Eningen and Saint Gaudens were listed by
Deperet in 1906.

Proboscideans : See list of Vindobonian proboscideans, below.

Vindobonian Proboscideans
Dinotherium levius Jourdan T. La Grive-Saint-Alban
Dinotherium intermedium Blainville Type locality? France
Dinotherium, bavaricum Meyer H. Gmiind
Dinotherium secundarium Kaup T. Simorre
Zygolophodon pyrenaicus (Lartet) T. Ile-en-Dodon
Zygolophodon borsoni affinis (Jourdan) Type locality?

Trilophodon angustidens (Cuvier) T. Simorre

Trilophodon angustidens minutus (Cuvier) Saxony
Trilophodon angustidens gaujaci (Lartet) Lombez
Trilophodon angustidens austro-germanicus Wegner Oppeln
Trilophodon angustidens gaillardi Osborn T. Villefranche d'Astarac
Trilophodon engelswiesensis Klahn H. Engelswies
Trilophodon steinheimensis Klahn T. Steinheim
Serridentinus s ubtapiroideus (Schlesinger) H. Wies
Turicius turicensis (Schinz) T. Elgg
Turicius turicensis simorrensis Osborn T. Simorre
T.—Tortonian
H. — Helvetian
4. THE UPPER MIOCENE: SARMATIAN
The close of the Miocene and the opening of the Pliocene in Eurasia was inaugurated by a new cycle of

sedimentation. This was, in brief, the beginning of the regression of Miocene seas, with a consequent inauguration
of extended continental sedimentation, characterized at first by littoral deposits, and then, in a progressive fashion.

'De Lapparent, A. de, 1906. Traitcdc Geologic, p. 1615.

 GEOLOGIC SUCCESSION: EUROPE 1465

by lake beds and river channels. The change from a predominantly mediterranean and island type of landscape,
typical of the European Miocene, to a truly continental type of landscape, so characteristic of the Pliocene, may
be said to have begun in the Upper Miocene, progressing with increasing rapidity through the Sarmatian and
finding its culmination in the Pontian.

The question of the correlation of Sarmatian deposits and faunas, which are characteristically of marine,
shallow-water facies, with the continental mammalian faunas of the same period is indeed difficult. This problem
is chiefly concerned with the query as to just what may be considered as truly Sarmatian or equivalent mam-
malian faunas.

Certain deposits, such as Saint Gaudens, (Eningen, Monte Bamboli and related beds, listed by Osborn in
1910 (p. 257) as of Sarmatian age, are more properly to be considered as representing the uppermost phases of the
Vindobonian or Tortonian in Europe, as shown in preceding pages of this chapter. This would seem to be the
general consensus of opinion of authorities on the matter and need not be discussed at any greater length here.

Yet the question of whether or not any mammalian faunas can be truly correlated with the marine Sarmatian
stage is an important one, and it has been recently considered by von Koenigswald and by Tobien.

Borissiak, in 1914 and 1915, described a mammaUan fauna that was found at Sebastopol, on the borders of
the Black Sea, in beds containing vmmistakable Sarmatian moUuscs. According to this author, the fossils were
found as a "breche ossifere" in a small calcareous lens, intercalated within the upper zone of the middle Sarmatian
of that region. Naturally, the occurrence of this fauna led the author, and numerous subsequent writers, to beheve
that the Sebastopol fauna was in truth a mammalian assemblage of Sarmatian age.

Yet it is a very curious fact that the mammaUan fauna of Sebastopol is typically Pontian in its characters.
It contains Achtaria, a giraffid quite similar to Palxotragus, Tragoceras, Aceratherium, and Hipparion; all in all an
assemblage that recalls Pikermi and Samos. Naturally, if this fauna is of true Sarmatian age, then it offers

undoubted proof of the pre-Pontian occurrence of Hipparion — a fact of prime importance. For, if as some
authors think, Hipparion appears in Eurasia before the beginning of the Pontian, the most important of the several
criteria definitive of the Pontian loses much of its diagnostic value.

The idea that the Sebastopol fauna is truly of Sarmatian age, or that Hipparion occurs in the Old World
prior to the advent of the Pontian, has recently been disputed by von Koenigswald. This author beUeves that
rather than a pre-Pontian appearance of what seems to be a typically Pontian fauna, the Sebastopol deposit
represents a cave, or possibly a fissure deposit, in which animals of a later age have been intruded into sediments
older than the period during which they lived. This explanation of the Sebastopol occurrence, if valid, goes
a longway towards clearing up what has always been a stumbHng block in the correlation of Eurasiatic mam-
malian horizons. Von Koenigswald' says that: "Das sarmatische Alter der Hipparion-FebXina von Sebastopol
kann bei der Art der Einlagerung und der Erhaltung der Reste nicht als unbedingt erwiesen angesehen werden."

Von Koenigswald goes on in the same contribution to dispute the idea that Hipparion is present as a pre-
Pontian form in any of the several localities where it has been reported as of an age preceding the characteristic
Pontian fauna.

As opposed to this view is the recent work of Tobien (1938), in which it is maintained that not only at
Sebastopol, but at various other localities, particularly in eastern Europe, there are beds of true Sarmatian age,

'Koenigswald, G. H. R. von, 1931. Sonderabdruck aus dem Centralblatt f. Min., etc., Abt. B, No. 1, p. 45.
'Tobien, H., 1938. Zeits. deutsch. Geol. Ges., XC, Heft 4.
 .

1466 OSBORN: THE PROBOSCIDEA

bearing Hipparion. The localities other than Sebastopol cited by Tobien are the trans-Caucasian region (Alekse-
ev, 1930), Linz (Giimbel, 1894), Bavarian Flinz (Freudenberg, 1928), Odessa (Andrussow, 1905), Constantinople
(Dietrich, 1933), Taraklia in Bessarabia (Khomenko, 1913, 1914), Thrace (von Arabu, 1916, 1919), Rumania
(Krejci-Graf, 1932). And in addition, Tobien describes some teeth and astragali of Hipparion from CEningen
(Ohningen)

So the problem stands at the present time. If these localities are truly of Sarmatian age, then they represent
what might be considered as the final stage of the continental Miocene in Europe, a stage during which Hipparion
and other Pontian types appear, foreshadowing the tremendous spread of the Hipparion faunas of the true Pontian.
If, on the other hand, these localities are of Pontian age (at least those in which Hipparion is present) then it may
be said that for practical purposes there are no continental mammalian faunas that may be proven as the exact
equivalents of the marine Sarmatian, and that the continental Miocene of Europe comes to a close with such
upper Vindobonian or Tortonian assemblages as those of Saint Gaudens and the like. At best the problem works
down to one of definitions, and as such is a difficult one to settle satisfactorily.

5. THE LOWER PLIOCENE: PONTIAN

The new faunal and sedimentation cycle that had its beginnings in the Sarmatian continued with increased
momentum into the following Tertiary subdivision in Europe, the Pontian. The mediterranean sea, so character-

istic of the Middle Miocene of Central Europe, became increasingly restricted, continental flood-plain and
lacustrine deposits replaced marine beds, and mammalian faunas became exceedingly wide-spread and character-
istic.

In this connection, there need only be repeated the well-known fact that the Pontian was the time during
which the characteristic Hipparion fauna spread throughout Eurasia. This fauna, which all in all was remark-
ably uniform throughout the extent of its range, stretched from Spain and western Europe to China, from the
shores of the Mediterranean as it was at that time into northern Europe. It was a wide-spread assemblage of

mammals living in a plains or steppe environment, and showing a great preponderance of grazing perissodactyls

and artiodactyls, accompanied by the carnivores that might be expected with an ecological and faunal association
such as this.

The presence of the Hipparion fauna in the Oriental and the Asiatic areas, and its significance with regard to

the correlation of the upper Tertiary beds of these regions, has been discussed in other sections of this chapter.
At the present time the discussion will be limited to the occurrence and significance of the Pontian Hipparion
fauna in Europe.

In a discussion of the continental Pontian faunas, the fact must be kept in mammalian mind that these
assemblages mark the beginning of a new period in the history of mammalian faunas. This was the time when
truly modernized types of advanced manunals made their first appearance, and particularly when inunigrant
forms suddenly became conspicuous in the Eurasiatic faunas, to characterize these faunas wherever they might
be developed.

Of course, as the name implies, the most characteristic of the immigrant genera is Hipparion. That this

genus is of undoubted North American origin can no longer be questioned by anyone who has made a careful
study of the succession of the Equida?. Consequently, as has already been pointed out in other sections of this
chapter, the appearance of Hipparion in the Old World must be subsequent to the time at which it arose in North
 GEOLOGIC SUCCESSION: EUROPE 1467

America —whatever that time may be. Certainly Hipparion did not become a genus distinct from its ancestor
Merychippiis, before the uppermost part of the Miocene in North America. And even though some evidence
would seem to indicate that Hipparion might have made its first appearance in North America during Upper
Miocene times, other evidence has been interpreted to indicate the fact that the genus did not appear in the New
World until the beginning of the Pliocene.

Turning now to a consideration of other Pontian types, we see numerous immigrant and specialized forms
appearing for the first time in Eurasia.

Among the carnivores it is interesting to note that true bears such as Indarctos and Ursavus —as distinguished
from the ancestral "bear-dogs," such as Hemicyon — first appear in the Pontian.

Likewise, the hyaenas, of Asiatic and Oriental origin, become well-established elements in the Tertiary faunas
in Pontian times. True enough, these animals first appear in Upper Miocene times, in the immediately pre-Pontian
deposits, but it is in the Pontian that the hyaenas become widely distributed throughout the European region.

A conspicuous element in the Pontian faunas of Europe, particularly along the eastern border of the Mediter-

ranean, is the aardvark, Oryderopus. The tubulidentates, though now of African distribution, did not necessarily

originate or evolve on that continent; indeed, all of the fossil aardvarks of pre- Pleistocene age come from the
Eurasiatic and Oriental regions, so that the group may well have originated somewhere in the north.

In a different category is the gigantic hyracoid, Pliohyrax — a conspicuous member of the Pikermi fauna. We
know quite definitely that the hyracoids originated in Africa in Eocene times, and so the presence of a member of

this group in the European Pontian implies an immigration into the fauna from a southerly source.

The most conspicuous elements of the Pontian fauna, with the exception of Hipparion, are the numerous
ruminants. Here is an advanced group of mammals, appearing for the most part as newly evolved elements in

the mammalian assemblage. Our evidence points rather strongly to the fact that the ruminants of advanced type
are relatively recent developments in the mammaUan world. That is, the common ancestor of the deer, of the
giraffids, and of the bovids is probably to be found in the Miocene.

In the European Pontian is a very primitive giraffe, Palseotragus, not greatly different in most of its characters
from the modern African okapi. With Palseotragus is Samotherium, which is nothing more nor less than an en-
larged type of Palxotragus, and Helladotherium, a gigantic girafiid belonging to a group that was to become
widely spread during Upper PUocene and Pleistocene times. It would seem probable that the origin of the giraffids

might have been Oriental —that they were immigrants into Europe and into Africa, and that the modern okapi
represents a persistent primitive form, pushed to the periphery of the range for this family by more specialized

types developing at or near the center of origin.

This was the age of the first flowering for the gazelles and the antelopes, and these animals are to be found in
great profusion throughout the Eurasiatic and Oriental Pontian. There is a host of forms (recently monographed
by Pilgrim) that give to the Hipparion fauna a very African appearance. It would seem, however, that these
various gazelles, sheep, antelopes, and pre-cattle were of northern origin and migrated in Pontian or post-Pontian
times to the southern continent.
 :

1468 OSBORN: THE PROBOSCIDEA

With these considerations in mind, what is the age of the Pontian? Among French scholars the European
Pontian has been generally accepted as of Upper Miocene age, while many Germans would place the Pontian at
the beginning of the Pliocene. Many years ago, as pointed out elsewhere in this chapter, Matthew suggested the
desirability of making the continental Pliocene throughout the world coincident with the appearance of Hip-
parion. This definition for the beginning of the Pliocene is, on the whole, as satisfactory a designation as has ever
been proposed. Consequently, the Pontian is here considered as of Lower Pliocene age, corresponding in Europe
to those stages in Asia, the Orient, and North America at which Hipparion first becomes a definitely identifiable

element in the mammalian faunas.

The outstanding Pontian localities of Europe are listed below taken from the
, list as presented by Pilgrim
in his "Pontian Carnivora of Europe." These localities, stretching from Spain on the west to the Black Sea region
and Asia Minor on the east, are characterized and correlated by the uniformity and the similarity of their faunas.

Spain Aragon, Catalonia, La Mancha

France Montredon, Orignac, Cerdagne, Mont L^beron, Cucuron, Croix Rousse
Germany Eppelsheim, Salmendingen, Melchingen, Trochtelfingen
Sicily Gravitelli
Greece Pikermi
Macedonia Salonica, Veles
Samos Samos
Hungary Baltavar, Polgardi, Csakvar, Baroth-kopecz
Black Sea (Sebastopol), Novo Elisavetovka, Taraklia Kischinev, Grebeniki, Tschobrotschi
Persia Maragha

Of these, perhaps the best known from their faunas, are: Mont Leberon, Cucuron, Eppelsheim, Pikermi,
Samos, and Maragha.

Of course, the presence of typical Pontian faunas characterized by Hipparion at the above localities affords

strong evidence as to their general unity and contemporaneity. Yet it is quite possible, as Pilgrim has pointed
out, that all of these faunas may not be exactly equivalent, each to the other, in age. That is, homotaxial factors
may play a certain part in the spread and delimitation of the Pontian complex in Europe—just as they most
probably did (as shown in another section of this paper) in India. Indeed, certain authorities would subdivide
the Pontian, particularly on the basis of the geology of the Black Sea area, into at least three zones. But in

spite of these considerations it is to be remembered that the Pontian faunas represent essentially a definite phase

in the evolution of mammaUan faunas, and as such are for all practical purposes to be coasidered together as a unit,
even though there may not be exact time identities between them —due to homotaxial lags dependent upon mi-
grations or other causes. This whole question has been no better stated than by Pilgrim ( 1931) ,• as follows

At the same time, there is no doubt that the 'Hipparion fauna' does occur at different levels, of which the lowest, including
perhaps the Sebastopol fauna described by Borissiak [but see remarks in the discussion preceding this on the Sarmatian problem]
(1915), has been referred to the Upper Sarmatian; the fauna of Taraklia (Khomenko, 1914) and Novo-Elisavetovka (Alexcjevv,
1916) are classed as Maeotic, or intermediate between the Sarmatian and the Pontian, while the remaining localities are truly
Pontian.
It is hardly conceivable that these three stages should not be represented among the so-called 'Pontian' deposits of the rest
of Europe. It may be that increased material, and intensive study of what is already in our museums, may one day render such
a correlation possible, but in my
opinion the state of our knowledge does not permit of it at present. . .

However this may be, there can be no doubt that we are dealing with a definite faunistic unit. Its homogeneous character
is specially striking when we compare it with the Vindobonian fauna of La-Grive-Saint-Alban which preceded it or that of Monte
Olivola, Rousillon and Montpellier, which followed it. The question, however, arises as to whether such a fauna, which includes
both Sarmatian and Pontian elements, can strictly be called Pontian. These names were definitely applied to certain marine
strata in the Black Sea region, and their application should not be either restricted or extended. Many writers, seeing this,
have written of this fauna as the 'Hipparion fauna', but since the genus Hipparion .seems to have a wider range even in Europe
and still more so in Asia and North America than the fauna we are considering, that term does not meet the need, and we seem
therefore bound to speak of the Sarmato-Pontian fauna.

'Pilgrim, G. E., 1931. Catalogiip of the Pontian Carnivora of Europe. Brit. Mus. (Nat. Hist.), pp. 147, 148.
 GEOLOGIC SUCCESSION: EUROPE 1469

PONTIAN PrOBOSCIDEA
Trilophodon (Choerolophodon) pentelicus (Gaudry and Lartet) Pikermi
Turicius atticus (Wagner) Pikermi
Tetralophodon grandincisivus (Schlesinger) Maragha
Tetralophodon longirostris (Kaup) Eppelsheim
Dinotherium medium Kaup Eppelsheim
Dinotherium giganteum Kaup Eppelsheim
Trilophodon esselbornensis (Kliihn) Esselborn
Turicius wahlheimensis (Kliihn) Wahlheim
Tetralophodon gigantorostris (Klahn) Bermersheim
Miomastodon tapiroides americanus (Schlesinger) Tasnad
Pliomastodon americanus praeiypica (Schlesinger) Batta-Erd
Dinotherium proavum Eichwald Podolia, Russia
Dinotherium podolicum Eichwald Podolia
Dinotherium uralense Eichwald Ural Mts.
Anancus arvernensis progressor Khomenko S. Bessarabia
Anancus intermedins (Eichwald) Volhynia
Platybelodon danovi Borissiak (Sarmatian) Kuban, N. Caucasus,
Chokrak Beds

6. THE MIDDLE PLIOCENE: PLAISANCIAN

Following the expansion of Pontian mammals in Europe, there would seem to be a gap in the succession of
mammalian faunas — perhaps the result of extensive marine inundations. This fact has been pointed out by
numerous authors, notably by Deperet, who was the first student to recognize and elucidate this distinct break
in the sequence of Pliocene mammalian faunas in Europe. Osborn (1910)^ emphasized the importance of Deperet's
conclusions, as quoted below, and more recently Pilgrim (1939)' pointed out the reality of a Middle Pliocene
hiatus in the European fossil mammal faunas.
As this is chiefly a marine phase, the terrestrial mammalian fauna is imperfectly known. The typical deposits are those of
the lignites of Casino (Tuscany) which arc correlated by Deperet with the lacustrine deposits of Autrey in the valley of the
Saone, France. On the east coast of England is a marine formation, the Coralline Crag of Suffolk, containing a mastodon and
a rhinoceros {Dicerorhinus).

Of course this gap is not complete, for there are scattered mammalian remains known from deposits of
Plaisancian age, but generally speaking the continental facies in this stage are of little importance. And it is

interesting to note that in many cases the fossil mammals of Plaisancian age are found in littoral deposits, some-
times associated with diagnostic marine invertebrates. All of these facts were made clear by Deperet, in liis

classic studies of some fifty years ago.

Deperet^* considered the Plaisancian and the succeeding Astian as closely related, and together they constituted
his "older Pliocene" which he distinguished from the "new Pliocene" or SiciHan. In the older Pliocene, according
to this author, were numerous holdovers from the Pontian fauna, while the Sicilian is marked by the influx of
a new and modernized fauna, quite distinct from those preceding it. According to the chronology adopted in this

'Osborn, H. F., 1910. "The Age of Mammals," pp. 311, 312.

^Pilgrim, G. E., and A. Tindell Hopwood, 1939. Rec. Geol. Surv. India, LXXIII, Pt. 4, pp. 445, 446.
^Deperet, C, 1893 (1894). Bull. See. geol. France, (3), XXI, pp. 529, 530.
 : ] —

1470 OSBORN: THE PROBOSCIDEA

present work, the Pontian, of course, represents the beginning of the Phocene, while the Plaisanoian and the
Astian, still showing mammalian types of Pontian origin, constitute the middle and upper phases of the Pliocene,
respectively. The Sicilian, with the influx of new and modernized types, such as Equus and Arcfiidiskodon,
represents the beginning of the Pleistocene (Dep6ret, 1894, op. cit., pp. 529, 530)

On
distingue nettement dans le facifes marin du Pliocene trois Stages: le Pliocene inferieur ou Plaisanc.ien, dont le type est
dans bleucs siibapennines; ensiiite le Pliocene moyen ou Aslien, fonde sur les sables jauncs tres fossilifferes du pays
les argiles
d'Asti ou ils surmontent avec evidence I'etage des argiles bleues.
Ces deux etages marins .... sont etroitement unis I'un a I'autre par leur distribution geographique et meme par leur faune
(qui denote une mer assez chaude), de sorte que Ton pourrait a la rigueur, avec MM.
de Rouville, de Stefani, Welscli, etc., les
consid^rer comme les deux phases .succe.ssives, la seconde plus littorale, d'un meme depot, sur un fond de mer en voie d'exhaus.se-
ment graduel.

Dep6ret then goes on to show that in Italy mammalian remains are rare in the marine facies of the Plaisancian,
and in the Astian as well, while to the north, especially in France, the Plaisancian deposits constitute a series of

blue clays with invertebrate remains, underlying or antecedent to the mammaliferous Astian sediments.

Pilgrim has recently restated Deperet's arguments, and has particularly emphasized the Plaisancian faunal
gap by demonstrating that the succeeding Astian fauna shows closer resemblances to the Sicilian fauna above it,

than to the older Pontian fauna.

Very little of the Roussillon and Montpellier fauna became extinct between that stage and that of the Val d'Arno, Perrier
and Seneze, as compared with the multitude of families and genera which appear for the last time in the Pontian. I can see no
grounds for regarding the two faunas as very diffiM-ent in age, while on the contrary a great zoological gap is suggested between
Roussillon and the Pontian. Two littoral or marine stages, the Plaisancian and the Astian, intervene between the Villafranchian
and the stage which corresponds to the mammaliferous lignites of Casino. The fauna of the Casino lignites is perhaps slightly
younger than that of the Pontian. Deperet, therefore, most reasonably considered that the Roussillon fauna corresponded with
the Astian, while the Plaisancian filled the faunal gap between the Pontian and Roussillon faunas. There is no manunal fauna
known in Europe which can be said to correspond with certainty to the Plaisancian. [Italics my own.

Three proboscideans have been referred very questionably to the Plaisancian. These are:

Turicius virgatidens (von Meyer) Fulda, Germany

Stegolophodon suhlatidens (Schlesinger) Teschen, Austria
Dinotherium gigantissimurn Stefanescu Gaiceana, Rumania

7. THE UPPER PLIOCENE: ASTIAN

The final Tertiary stage in Europe is that of the Astian, typified by the deposits of Villanova, Asti, in Italy,
and characterized by the very rich mammalian faunas of Roussillon and Montpellier in France. Deperet demon-
strated very ably, as already shown above, the fact that the Astian fauna or faunas are essentially modernized
much closer to the Pleistocene and Recent faunas of the Holarctic region than to the Pliocene faunas preceding this
stage. This point has been strongly emphasized by Pilgrim, in a recent paper, and it was more generally stated

by Osborn in 1910,' as follows:

Surveyingthis. Pliocene fauna as a whole we are struck by the great predominance of animals closely related to exist-
. .

ing forms. the living zoologist should imagine himself in France at this period, he would see only four animals which would
If
appear entirely novel and unknown, namely, the saber-tooth tiger, the mastodon, the hijiparion, and the hyaMiarctos; all the
rest of the fauna would seem to he a very strange (ommingliiig, or congre.s.s, of African, i'Ain)])eaii, and Asiatic manmials of the
present day. Not a single North American element would be observed in this assemblage, unless we except those elements of
more remote migration, such as the hares, the tai)irs, and possibly the hipparions and the foxes.

'Osborn, H. F., 1910. "The Age of Mamm.als," \>. 317.

 GEOLOGIC SUCCESSION: EUROPE 1471

One of the chief deposits of Astian age, perhaps the most important one in Europe, is that of Roussillon, fully
described by Deperet in his classic monograph of 1890.' According to this author, after the retreat of the Plaisan-

cian sea, the Roussillon basin was filled with a considerable thickness of argillaceous and siUcious sandstones, and
in addition calcareous muds, all having a fluviatile origin. It is in these latter beds that the characteristic Rous-
sillon fauna was deposited.

En revanche, ces limons sont le gisement d'une riche faune de vertebras terrestres et fluviatiles, dont les debris charries par
les courants, se retrouvent aujourd'hui dans quelques points privilegies, qui correspondent sans doute a des remous ou a des
parties plus stagnantes du cours des anciennes rivieres. Les pieces osseuses sont presque toujours isolees, souvent brisees et
meme un peu roulees il est assez rare de rencontrer plusieurs os en connexion. Cependant di verses parties d'un meme squelette
;

se retrouvent quelquefois dan.s un rayon peu etendu; meme le squelette de la Testudo Perpiniana a pu etre deterre tout entier
grace a la protection efficace de la boite osseuse, et a I'habitude qu'ont les Tortues de retracter leurs membres et leur tete dans
I'int^rieur de la carapace.
Les ossements se rencontrent dans les limons d'eau douce sur presque toute I'etendue du bassin du Roussillon les localites ;

les plus riches sont: Villemolaque, Trouillas, le mas Bclrich dans la vallee du R^art; Thuir, Millas, le Soler, le Serrat d'en Vac-
quer, la citadelle et les briqueteries de.s portes Canet et St-Martin, a Perpignan, dans la vallee de la Tet; les briqueteries de
Rivesaltes dans la vallee de I'Agly.

Pilgrim^ (1939) has presented a very helpful review of the Roussillon fauna, particularly with regard to its
relationships with European assemblages below and above it. Some of his remarks are as follows:

The character of the European fauna altered considerably between the Pontian and the stage of Roussillon. Numerous
families and genera have disappeared. . . .

There are very few surviving genera from the Pontian, and almost all of these differ specifically. . . .

On the other hand numerous species from Roussillon are identical or nearly so with Villafranchian forms.
The differences which the fauna of the Villafranchian displays from that of Roussillon are mainly due to what are invading
forms, which does not necessarily imply any great difference in age. . . .

When we observe that the comparable forms are either specifically the same or differ very little from one another, we are
forced to conclude that little development took place between the two levels. Very little of the Roussillon and Montpellier
fauna became e.xtinct between that .stage and that of the Val'Arno, Perrier and Seneze, as compared with the multitude of fami-
lies and genera which appear for the last time in the Pontian. I can see no grounds for regarding the two faunas as very differ-
ent in age, while on the contrary a great zoological gap is suggested between Rou.Sisillon and the Pontian.

The Astian fauna, as known from the assemblages of Roussillon and Montpellier, is large and varied and
shows a composition somewhat as follows:

Among the primates are Dolichopithecus, Semnopithecus, and Macacus, the first of which failed to survive into

Pleistocene times.

The carnivores display a considerable variety, including Vulpes, Ursus (both appearing for the first time in
Europe), Agriotherium, Lutra, Viverra, Hyaena, Megantereon, Epimachaerodus, and Felis.

A large group of rodents, including Hystrix, and one form, Ruscinomys, which became extinct at the end of
the Pliocene.

The proboscidean genera, Zygolophodon and Anancus.

Among the perissodactyls, Hipparion as found in the Astian represents a long-pensistent survivor from
Pontian times. Tliis genus, wliich generally became extinct at the end of the Pliocene, did persist in some regions
into the Lower Pleistocene. Also there is to be noted Tapirus and Dicer orhinus.

Among the artiodactyls Potamochcerus and Capreolus appear here for the first time. There are also various
bovids, such as Palseoryx and Gazella.

'Deperet, C, 1890. Mem. Soc. geol. France, III, p. 9.

^Pilgrim, G. E., and A. Tindell Hopwood, 1939, op. cit., pp. 443-445.
1472 OSBORN: THE PROBOSCIDEA

There is also Oryderopus, a large species as compared with the characteristic Pontian forms —although smaller
than the recent types.

Of particular significance is the absence of certain diagnostic genera from the Astian faunas, notably Equus,
Elephas (Archidiskodon), and bovines, such as Leptobos. It is the absence of these types that gives conclusive proof
of the distinction of the Astian fauna from the succeeding Villafranchian, where these animals as immigrant forms
appear for the first time in Europe. This difference was noted by Deperet, and has been emphasized by later
authors.

In addition to the characteristic Astian mammaUan faunas of France, and to a lesser extent of Italy, there
are limited occurrences ofmammals of this age in other parts of Europe, notably along the eastern coast of
England. Here is found the Red Crag deposit, a marine sediment containing occasional mammals. The mam-
mals, when found in the Red Crag, prove to be of the same types as those found at Montpellier and Roussillon.

The occurrences of the types of Astian proboscideans are given by Osborn as follows. In this connection it

might be mentioned that Pilgrim (1939) denies the presence of Zygolophodon borsoni at Roussillon, although

Deperet in his monograpliic study lists this species among the Roussillon fauna.

Astian Proboscideans
Zygolophodon borsoni (Hays) Villanova, Asti, Italy
Zygolophodon borsoni buffonis (Pomel) Auvergne, France
Zygolophodon borsoni zaddachi (Jentzsch) Thorn, Germany
Zygolophodon borsoni vellavus (Aymard) Velay, France
Zygolophodon borsoni vialetii (Aymard) Vialette, France
Anancus arvernensis (Croizet and Jobert) Perrier, France
Anancus arvernensis brevirostris (Gervais and de Serres) Montpellier, France
Anancus arvernensis dissimilis (Jourdan) Saone Basin, France
Anancus arvernensis macroplus Aymard Mt. Coupet, Puy-en-Velay, France
Anancus gigantarvernensis (Klahn) Herbolzheim, Germany
Anancus minutoarvernensis (Klahn) Herbolzheim, Germany
Anancus falconeri Osborn Suffolk, England

8. THE PLEISTOCENE
There was no abrupt change from the Pliocene to the Pleistocene in Europe, a condition which was generally
true for the other parts of the World. Consequently the boundary between the two epochs is difficult to define,

with the result that there are manifest differences of ojiinion as to what event or events in geologic history may be
properly regarded as significant of the opening of Pleistocene times in this region. In a general way, the Pleisto-
cene is the Great Ice Age, when portions of the northern hemisphere were partially covered by several successive
glaciations, and it might be supposed that the advance of the first continental glacier would serve to mark the
beginning of Pleistocene history. But some of the European authorities, notably Boule, would place the first
glaciation in the Upper Pliocene. Moreover a great portion of the Earth's surface was not affected by glacial
phenomena, so that other criteria must be used, no matter what may be the opinion as to the time of the first

glacial advance. The sequence of pluvial and dry periods, the development of marine and river terraces and other
geological phenomena have been widely studied and variously used as aids in Pleistocene chronology in the non-
 GEOLOGIC SUCCESSION: EUROPE 1473

glaciated regions of the Earth, yet even these evidences, useful as they are in estabhshing the succession of
diastrophic, climatic or sedimentary events within the Pleistocene period, have not been generally satisfactory in
an attack upon the problem of the beginning of the Pleistocene. Therefore the evidences of wide-spread changes in
the mammals are perhaps the most significant and the most useful of the criteria studied, in establishing the open-
ing of Pleistocene times. The mammals were rapidly evolving animals, able to migrate quickly over most of the
Earth's surface, so that changes in mammalian faunas necessitated by the adaptations required by the development
ofnew and unusual environmental conditions would almost instantly (from a geological point of view) be reflected

throughout wide areas. The problem has been excellently stated by Hopwood,' as follows:

Apparently the boundary between the Pliocene and Pleistocene is difficult to determine on purely geological evidence, so that
necessary to approach the problem in some other way, preferably by means of the fossil mammals since they furnish the raw
it is
material for the rest of the inquiry. There are many factors involved, but two of them are far more important than the others.
First of these is the proposition that the best boundaries in time are those expressed by a change in fauna, which means in
practice the incursion of new types, rather than the disappearance of old ones. The second proposition is, that the time necessary
for the distribution of new types of active quadrupeds, e.g., horse or bison, over a very wide area is negligible from the geologi-
cal point of view.

At the beginning of the so-called Sicilian or Villafrancliian time division in Europe and Asia, there was
a sudden appearance of mammals, part of which were the descendants of indigenous
new and modernized types of

ancestors, part of which were immigrants. Notable among these new types were the modern forms of horses
(Equus), of cattle (Bos), and of elephants {Archidiskodon, Elephas and related genera). As long ago as 1911, Haug
suggested that the appearance of these key types, the first an immigrant from the New World, the others of Old
World origin, should mark the beginning of the Pleistocene in Europe. In recent years this view has been reiterat-
ed, notablyby Matthew and by Hopwood. If accepted, then the typical SiciUan or Villafranchian faunas of the
European region are to be regarded as of Lower or Basal Pleistocene age, and it is the influx of these new types
that marks the beginnings of the period.

As opposed to this, many European authorities are inclined to regard the Villafranchian fauna, containing
Equus, Bos, and Elephas, as marking the summit of the Pliocene. In the present chapter the former interpreta-
tion will be favored.

Granting that the opening of the Pleistocene is marked by the appearance of numerous modernized mammals,
especially Equus, Bos, and Elephas (in the broad sense of the term), we may now consider the problem of dat-
ing the sequence of events within Pleistocene times. This has occupied the attention of many students over a
long period of years. In Europe the criteria most generally used for establishing dates within the Pleistocene
are those of the glacial succession, as delineated especially by Penck and by Geikie, and of the sequence of marine
and river terraces, as described by de Lamothe, Deperet, and others.

In northern Europe there are evidences of four major glaciations, with interglacial periods between them. In
the Mediterranean region there are the remains of four marine terraces, which may be traced in part around the
Atlantic coast and into the North Sea. In 1919, Deperet indicated the relationships between the European
glaciations and marine terraces in the following tabular form.

Northern Europe Alps Mediterranean

Fourth glaciation
 .

1474 OSBORN: THE PROBOSCIDEA

It must not be supposed that this compilation represents an exact correhition, since it is known tliat the
marine terraces may be correlated with the river terraces, while these latter underlie the moraines of the glaciations
to which they are most closely related. Therefore, it is probable that the terraces are more or less of interglacial

age, and immediately precede the glaciations with which they may be roughly correlated. The important fact

is that there is a fourfold division of the Pleistocene in Europe, as based either upon the development of marine
and river terraces, or upon the sequence of glaciations. And each glaciation may be considered as occupying the
latter portion of the general terrace stage, as shown below.

TV /Upper Monastirian IV Wiirm

\Lower Monastirian

TTT /Upper Tyrrhenian III Riss

\ Lower Tyrrhenian

TT /Upper Milazzian II Mindel

\ Lower Milazzian

-r /Upper Sicilian I Giinz

\ Lower Sicilian

Many students, following particularly the work of Penck, have envisaged a succession of "warm" and "cold"
mammalian faunas in Europe that might be correlated more or less closely with the successive advances and
retreats of the great ice sheets. According to Penck's scheme, the four glaciations were more or less equally

developed, but separated from each other by unequal intervals, of which the median one, the Mindel-Riss inter-
glacial, was by far the most protracted, and the last one, the Riss-Wurm, the shortest. He also distinguished the
Wlirm glaciation from the others by his supposition that it terminated in a series of minor fluctuations, rather
than by a .single and steady decline. This idea of alternating faunas controlled by glacial phenomena has been
expressed by Osborn' in the following words:

Tho principal contributors to the theory of northward and southward migrations and to the succession of faunas are
Nchring, Woldrich (1882), and more recently Penck. In considering the distribution and migration of the mammals throughout
the (ilacial Period, we must constantly keep in mind the differences of latitude. . .

Penck also observes that we cannot hope to trace a continuous evolution of forms during Pleistocene times, because we are
not dealing with a development of one successive series in one locality, but with the cyclical alternation of a number of different
faimas compelled to migrate through the alternations in the temperature and in the floras, the mammals disappearing and re-
turning at intervals too brief to allow of any marked evolutionary changes. Herein lies our difficulty when we attempt to distin-
guish between the tundra faunas of the late glaciations and the forest faunas of the late interglacial epochs, because the faunas
return not only with the same generic but the same specific types, as especially illustrated in the case of the mammoth (E.
primigenius) and the giant deer (Cervus inegaceros)

A considerably different picture has been presented recently (1933) by Stehlin,- who thinks that the glacial

periods were not only separated each from the other by interglacial periods of varying lengths, but also that they
were in themselves much more different from each other in intensity than was admitted by Penck. Thus Stehlin,

ba.sing his conclusions upon the development of the mammahan faunas, supposes that the glaciation at the begin-
ning of the Pleistocene was relatively much less intense than the two glaciations coming after the long Middle
Pleistocene interglacial period. (Incidentally, this author, like Boule, recognizes only three glaciations in the
Pleistocene.) According to Stehlin, the maximum of glacial activity was reached towards the end of the Wiirm
'Osborn, H. F., 1910, op. cit., pp. 388, 389.
-Dubois, A., and H. G. Stehlin, 1933. M(:m. Soc. Pul. Suisse, LIl-LIII, p. 272.
 GEOLOGIC SUCCESSION: EUROPE 1475

glacial period, a conception based upon the sequence of European Pleistocene faunas and which regards the
"warm" or temperate types of mammals as prevaiUng through a great part of the Pleistocene, to be followed in
Wiirm and then in post-Wtirm times by an invading arctic assemblage: "... la plus grand invasion des animaux
arctiques dans les latitudes moyennes et meridionales ne coincide avec le developpement maximal des glaciers
wurmiens, mais lui succede. .
."

This view is somewhat in accord with the recent tendency to discount to a large degree the reality of alternat-

ing "cold" and "warm" faunas, and to regard the development of the Pleistocene mammals as a rather continuous
process, with minor fluctuations due to the alternation of mild and severe temperature conditions.

Hopwood (1939, 1940), for instance, has the following to say on this subject:

At this stage it may be useful to discuss some of the points which govern the use of mammahan faunas as indices of the age
of Pleistocene deposits. It should be self-evident that the composition of a fauna will, in the main, depend on the climate as well
as on the environment; the literature is full of reference to 'cold' and 'warm' faunas, and also to faunas of 'forest,' 'parkland,'
or 'plains' type Moreover, one cannot be absolutely certain that any particular species indicates a particular type of climate.
. . .

. . . 'warm-climate' mammals are considered it is well to keep in mind the wide range of climate acceptable to such animals
When
as the leopard and tiger, and to remember that even a hippopotamus can survive an English winter with a minimum of extra
shelter and warmth.
of a long series of fossils, even an approximation to the true date is all but impossible in the ab.sence of
Except with the aid
such easily recognisable species as Equus robustus Pomel, Rhinoceros etruscus Falconer, Elephas meridionalis Nesti, or Rayigifer
larandus (Linn.)'"
Geographical changes of this magnitude [uplift of Alps, etc.] might be expected to have influenced the climate and the
fauna in some way or another, but this does not .seem to have happened. Admittedly, much has been written about alternating
'warm' and 'cold' faunas, particularly in Germany, where the teachings of Penck were most influential, but in fact there is very
little evidence to support the more extreme expositions of this view. .'-'
.

It has been the practice among many European palaeontologists to recognize three general mammaUan faunas
in the Pleistocene of Europe, corresponding roughly to the lower, middle, and upper phases of the period.
Although, as de Lapparent has stated, the various individual animals constituting these faunas extend beyond the
vague limits of each assemblage to mingle with each other, nevertheless associations of certain mammals are
characteristic of the early, middle, and late portions of Pleistocene times. Thus he would recognize an early warm
fauna, typified by Elephas antiquus, a middle cool and humid period with Elephas primigenius and Rhinoceros
tichorhinus, and finally a late cold fauna with the woolly mammoth and reindeer predominating. This was the
general succession recognized by Osborn in his "Age of Mammals" in 1910, and described in a somewhat similar
form recently (1935) by Boule and Piveteau.'^

While tliis conception of the succession of Pleistocene faunas in Europe may be true in a general way, it is

probably oversimplified. In the first place, it seems evident from recent work in various parts of the world that
many if not most of the Pleistocene mammals appeared at an early stage in the history of the period, to continue
with but little change through its extent. With the exception of certain rapidly developing types, such as the

hominids and the proboscideans, there was probably but little evolution of a super-specific nature occurring within
the relatively short duration of Pleistocene times. There were some extinctions, it is true, and these, as much as
anything else, serve to tUstinguish the characters of the faunas in successive phases of the Ice Age. As regards
this, it would appear that the evidence for extinctions at various times within the Pleistocene is certainly much
more convincing in Eurasia than it is in the Americas.

'Hopwood, A. T., 1939. Proc. Prchist. See, N. S., V, Ft. 1, pp. 13, 14.
^Hopwood, A. T., 1940. Proc. Geol. Assoc, LI, Pt. 1, p. 85.
'Boule, Marccllin, and Jean Piveteau, 1935. "Les Fossilcs."
 — —

1476 OSBORN: THE PROBOSCIDEA

Hopwood has outlined the relationships of the appearance and disappearance within the Pleistocene in
Europe as follows.

Glacial IV. — Extinction of Elephas primigenius, Rhinoceros tichorhinus, Hyaena spelxa, Ursus spelxus.
Interglacial — Extinction of Elephas antiquus, Rhinoceros megarhinus, Felis spelxa. leo

Glacial III.—
Interglacial — First appearance of Bos primigenius, Rangifer tarandus, Rhinoceros tichorhinus, Ursus
spelxus, Hyxna spelxa.

Glacial II. —Extinction of Hyxna arvernensis, Canis nescherensis, Equus robustus, Equus stenonis, Rhi-
noceros etruscus, Elephas meridionalis.

Interglacial

Glacial I.

These were some of the principal changes that took place during the development of the Pleistocene faunas in
Europe, and it is mainly by looking at them that we can see the progress of Pleistocene history reflected in the
assemblages of Ice Age mammals. Compared with changes such as these, the influx or egress of mammals
adapted to warm or cold climates was of relatively slight importance. As Hopwood states': "Climatic variations
may have favoured first one type and then the other, but neither type was completely expelled from the area."
The Lower Pleistocene proboscideans of Europe were Archidiskodon planijrons, Archidiskodon
typical Basal or
meridionalis, and Hesperoloxodon antiquus. Parelephas trogontherii and Elephas primigenius, perhaps not quite so
ancient in their first appearance as the above mentioned forms, nevertheless were present in the European Pleis-
tocene before the close of its lower phase. Seemingly there was an early extinction of the Archidiskodon group,
specifically ^.pZam/rons and A. meridionalis, probably in the interval marking the close of the first interglacial

and the opening of the second glacial periods. Hesperoloxodon antiquus probably became extinct in middle or
later Pleistocene times, perhaps near the end of the third interglacial period according to Hopwood. Finally,

during or after the fourth glaciation there was an extinction of Elephas primigenius.

Proboscideans:
Mammonteus primigenius (Blumenbach) Palxoloxodon lamarmorae (Major)
Mammonteus primigenius hydruntinus (Botti) Palxoloxodon Cypriotes (Bate)
Mammonteus primigenius fraasi (Dietrich) Palxoloxodon creticus (Bate)
Mammonteus^ primigenius leith-adamsi (Pohlig) Parelephas trogontherii (Pohlig)
Mammonteus primigenius astensis (Deperet and Parelephas trogontherii nestii (Pohlig)
Mayet) Parelephas trogonlherioides (Zuffardi)
Loxodonta cornaliae Aradas Parelephas intermedins (Jourdan)
Hesperoloxodon antiquus (Falconer and Cautley) Parelephas wusti (Pavlow)
Hesperoloxodon antiquus germanicus (Stefanescu) Parelephas armeniacus (Falconer)
Hesperoloxodon antiquus italicus Osborn Archidiskodon meridionalis (Nesti)
Hesperoloxodon antiquus platyrhynchus (Graells) Archidiskodon meridionalis cromerensis (Deperet
Hesperoloxodon antiquus ausonius (Major) and Mayet)
Hesperoloxodon antiquus nanus (Acconci) Archidiskodon planijrons (Falconer and Cautley)
Pnlxoloxodon mnaidriensis (Adams) Archidiskodon planijrons rumanus (Stefanescu)
Palxoloxodon falconeri (Busk) Mastodon pavlowi Osborn
Palxoloxodon melitensis (Falconer) Anancus arvernensis (Croizet and Jobert)
'Hopwood, A. T., 1940, op. cil., p. 86.
 :

GEOLOGIC SUCCESSION: ASIA 1477

V. ASIA
1. INTRODUCTION
Asia, as here used, comprises that portion of the continent north and west of the Himalaya Mountains and the
Tibetan Plateau. Thus there is included within it Siberia, Mongolia, North China, Japan, and Turkestan. This
separation of the more northerly and westerly sections of Asia from the Oriental Region is logical and natural,
being based in large part on the modern zoogeographic realms. That part of Asia delimited above constitutes at
the present time the eastern moiety of the Palsearctic Realm, and the evidence would seem to be indicative of the
fact that during the later stages of the Cenozoic, the Palsearctic portion of Asia was zoogeographically distinct from
the more southerly and easterly Oriental section, just as it is today.

2. THE MIOCENE OF MONGOLIA AND CENTRAL ASIA

Mongolia: Loh Formation.— The Loh formation is exposed to the south of Uskuk Mountain in the Tsagan
Nor basin of MongoUa, where it overhes the Upper Ohgocene Hsanda Gol beds. The only fossils identified to date
from the Loh formation are Serridentinus mongoliensis ^£02^GIC24J. REI^^fONSmPS OF
Osborn and Baluchitherium mongoliensis Osborn. (This ASIATfC PR. OB OS CIDEt^
MONGOLIA CENTRAL NORTH CHINA JAPAN
probably not Baluchitherium but rather a ASIA
latter form is
Elephss indicus bu»ki
(Palaeoloxodon?)
rhinoceros of the Coelodonta group.) From the evidence
of these fossils, and due to the fact that the Loh rests Mammonteus
primiqcnii
Parelephas
tro^ontherii
directly on the Upper Oligocene Hsanda Gol, it would Palaeoloxodon
narriadicus
naumanni
seem very probable that the Loh formation represents ^A laeoloxodon nam adieus
setoensis
tpkuna^ai namadicus uabei
a Lower Miocene or BurdigaUan phase in the sedimen- Palafeoloxodon jodon
Steqo
namadicLiS: onenfaiis,
sinensis
tary history of Mongolia. It might be pointed out Archfdiskodon
planifrons .

here that the fossils of the Loh beds might be placed

in later phases of the Miocene, particularly the Middle
(SHANSI) -

Parelephas
protomammonteus
Miocene, with as much justification as in the Lower Archidiokodon planifrons
Sfeggdon zdanskui, oricntali:
(=Paliioloxodon)I
Palieoloxodoh
"Lower ZugolopKodon borsoni
Miocene. Indeed, this series was designated as "Mastodon" intermedins
fokuna^ai •
Trilophodon.
sendaiciis
Middle Miocene" by Professor Osborn Volume
"

to in I
Sfeaodon
aur-orae
of this Monograph. KHUf/UK HIPPARION- ClAY^ bonibifrons
elephantoides
Serridentinus Serridentinus Luimani, Ii^dekkeri
(Ocalienfinus) Tnlophodon spectabilis
Floreacens Tetralop^iodon exoletus, sinensis
Pentalophodon sinensis, cuneatus
Berkey and Morris' described the Loh formation as 249olophodon borsom '

"Mastodon" intermcdius
Stegodon licentij zdanskt^i
follows
TVHG GUR.
PlafLjbelodpn TrilopKodon Serridentinus
connexus annectens
Directly overlying the Hsanda Gol clays, about five Semdenrinus
miles south of Uskuk Mountain, there is a group of olive qobiensis Xrilpphodon
palaeindicus
green clays less than one hundred feet thick. No clearly LOH JfLAUCIK Steaolophodon
defined physical break can be seen between the two for- Serridentinus Trtlophodon
latidens^
monqoliensis inopinafus
mations, but the upper olive clays yielded fossils which a Ta V u s 5teaodbn sp,2
Dr. W. D. Matthew correlates with the Lower Miocene of cf. anqu5^ldens

Europe. 1

J Doubtfuf occurrence in l"he P ioccne PrUyett 3ra.dlei^

2Poubtful occurrence in ttie Miocene

Proboscidean: Serridentinus mongoliensis Osborn. Fig. 1223

Tung Gur Formation. — The TungGur formation of Mongolia is typically exposed along a northeast to
southwest trending escarpment in the immediate vicinity of Gur Tung Khara Usu, Inner Mongolia. This escarp-

^Berkey. C. P.. and F. K. Morris, 1927. Natural History of Central Asia, II, p. 365.
1478 OSBORN: THE PROBOSCIDEA

ment is part of a broad table-land situated to the northeast of the Kalgan-Urga trail and near the border between
Inner and Outer Mongolia.

The Tung Gur formation was described by Spock in 1929" and again by the same author in 193U.' On the
basis of geologic evidence alone Spock suggested that this formation might be of Pliocene age, which was the
opinion first held by Professor Osborn as a result of his studies of Plahjhelodon, the most important and by far
the most spectacular fossil mammal from this horizon.
Soon after the Tung Gur fauna was discovered, however, P. Teilhard de Chardin suggested (verbally) that
the fauna might be of Miocene age —or more specifically of pre-Pontian affinities. Teilhard's conclusion was
based upon his observations of the fossils as they were collected in the field. This correlation of the Tung
Gur was subsequently by the detailed studies of the fauna by various authors including the present writer.
verified

Consequently, Professor Osborn came to regard the Tung Gur formation as of Upper Miocene age, thereby placing
Platybelodon as an earher stage of shovel-tusked mastodont than he had first considered it to be. (See Volume
I, pages 463-466, of this Monograph.)

The Tung Gur fauna is too large for detailed consideration or listing at this place. Some of the important
forms constituting the fauna, however, may be briefly considered.

As for the rodents, there is an Amblycastor closely related to Miocene species of the same genus occurring
in North America. Among the carnivores, the giant canid or ursid, Hemicyon, may be linked with certain
Miocene species referable to this genus, particularly as the Tung Gur form shows numerous relatively primitive,
dog-like characters. There is a hyaena of the Crocuta group, which might, on the other hand, perfectly well be of
Pliocene age. The same is true of Platybelodon, since it shows many specialized characters. Perhaps one of the
most significant of the Tung Gur animals is Anchitherium, definitely a Miocene type. The presence of this

forest horse and the complete absence of any trace of Hippnrion constitute strong evidence in favor of a Miocene

age for the Tung Gur formation. The rhinoceroses, too, would seem to be of Upper Miocene affinities. A pig,

Listriodon, on the whole shows affinities with Miocene species, although this is a genus that persisted from
the Miocene into the Pliocene in some localities. The deer, Stephanocemas and Lagomeryx, show definite

relationships with comparable Upper Miocene forms from Europe and Asia. Of the bovids, one species, Oioceras
noverca is, according to Pilgrim, a small form that might have been ancestral to certain Pontian species.

From this brief review it becomes evident that the Tung Gur fauna, although containing some possible
Pliocene elements, is, in its general aspects, of Upper Miocene affinities, a conclusion that is greatly strengthened by
the presence of Anchitherium and the absence of Hipparion in the assemblage, particularly since the absence of
the latter form would seem to be real and not due to accidents of collecting. It would seem that the Tung Gur
fauna, with a mixture of woodland and plains species, represents a borderland assemblage of mammals.

Proboscideans:
Platybelodon grangeri (Osborn)
Serridentinus gobiensis Osborn

—
Central Asia: Jilancik Beds, Turcjai Region. In a series of papers published between 1927 and the
present time Borissiak has described a mammalian fauna from the Turgai region of Central Asia, discovered in the
Jilancik (or Dschilantschik) beds of Miocene age. The Jilancik beds r(>st on a series of Oligocene deposits contain-
ing Indricotherium, and in turn are succeeded by a Pliocene horizon in which Hipparion is present. At the
'Spock, L. E., 1929. Amcr. Mus. Novitatcs, No. 391; ibid, 1930, No. 407.
 GEOLOGIC SUCCESSION: ASIA 1479

present time the Jilancik fauna (excavated during a long period of years, beginning in 1914) would seem to be
composed entirely of proboscideans and rhinoceroses, as follows.

TrilopJwdon inopinatus Borissiak and Beliaeva

Trilophodon cf. angustidens (Cuvier)
Trilophodon atavus (Borissiak)'
Brachypotherium aurelianense Nouel
Aceratherium depereti Borissiak

This fauna has a distinct IMiocene aspect, and due to the determination of elements within it identical with or
close to certain Lower Miocene species of Europe, Borissiak has referred the assemblage to the Burdigalian. With
regard to the age of the Jilancik beds, the following remarks might be appropriate at this place.

Trilophodon inopinatus
Although Borissiak thought that this species was close to Serridentinus mongoliensis of the Loh forma-
tion, Osborn placed it definitely in the genus Trilophodon. Borissiak, as the result of his comparisons, considered
the Loh formation to be virtually identical with the Jilancik beds in age. Osborn, on the other hand, compared
Trilophodon inopinatus with the primitive Trilophodon cooperi, from the Bugti beds of Baluchistan. It is inter-

esting to notice that both comparisons are with Lower Miocene horizons. This agrees with other evidence on the
age of the Jilancik beds.

Trilophodon angustidens

If this identification is valid, the Jilancik beds may be correlated with the Lower Miocene or BurcUgahan of
Europe.

Trilophodon atavus
This species is very close to T. angustidens. This again is a link with the Lower Miocene of Europe.

Brachypotherium aurelianense
According to Borissiak, this characteristic European Burdigahan form is present in the Jilancik sediments.
He has shown, however, that the Jilancik form displays certain advanced traits in the direction of the Middle
Miocene B. brachypus of Europe.

Aceratherium depereti
Placed by Borissiak between A. lemanense and A. tetradactylum in its evolutionary stage of development.
This species shows certain speciahzations, notably the very much elongated nasals, which mark an advance
beyond the typical Burdigalian forms.

Proboscideans: See faunal list, above.

3. THE MIOCENE OF NORTH CHINA

China. —The study of the fossil mammals of China dates from the middle of the last century, when Davidson
published a short notice of some palseontological material gathered together in Shanghai by W. Lockhart. Among
these first-known of the Chinese fossils was a tooth of an elephant. From that time until the present day a suc-

'This form has been recently described by Borissiak, under the name of Mastodon atavus. It is closely related to Trilophodon angustidens, as Borissiak
points out; consequently it is here referred to this latter genus.
1480 OSBORN: THE PROBOSCIDEA

cession of students, notably Owen, Gaudry, Koken, Schlosser, and in recent years the various authors describing
the collections made by the Geological Survey of China, the Pala-ontological Institute of Upsala University, and
the American Museum of Natural History, have added increasingly important and voluminous data concerning
the palseontological history of northern China and Mongolia.

The liistory of Upper Cenozoic mammals

North China would seem to begin with the Middle or Upper
in

Miocene and to continue almost uninterruptedly through the PHocene into the Pleistocene. Through this stretch
of geologic time two periods are marked by the expansion and the unusual abundance of fossil mammals. These are

the Lower PUocene or Pontian, in which the large, widely distributed "Hipparion fauna" occurs, and the Lower
and Middle Pleistocene, in which there are several faunas, notably the "Equus fauna" or Villafranchian assem-
blages of the flood-plain deposits and the later Choukoutien cave faunas.

As to the beds preceding the almost universal Hipparion clays, there is little to be said. At best, these

sediments are physically but sUghtly differentiated from the Pontian deposits, if they are at all distinguishable.

There have been described, from time to time, certain fossils that would seem to be of definitely Upper Miocene
rather than of Pontian affinities, and it is on the basis of these discoveries, as much as anything, that the presence
of pre-Pontian sediments in North China is inferred.

Such is the case of Trilophodon connexus, which Hopwood regards as a very primitive stage in the buno-
dont mastodon ts, closely comparable to the Miocene Trilophodon cooperi of Baluchistan. Trilophodon
cooperi is a Lower Miocene species; whether, on the basis of this, Trilophodon connexus should be regarded
as a very early Miocene form, or rather a structurally primitive species persisting into the Upper Miocene, is

a question open to some doubt. It is possible that this species may represent a Sarmatian or an equivalent age in
North China. On the other hand, there is no reason why the supposed pre-Pontian species in North China might
not be structurally primitive forms persisting into the Lower Pliocene.

In this connection it might be noted that Trilophodon wimani, considered by Hopwood as possibly of

Sarmatian age, was placed by Teilhard in the Pontian. Hopwood based his conclusions on the primitive structure
of Trilophodon wimani, and also on the fact that it was associated with Listriodon gigas, a pig showing affini-

ties, according to Miss Pearson, with certain Miocene forms of Europe. Yet Teilhard's conclusions as to the age of

Trilophodon wimani were based on his thorough knowledge of the stratigraphic relationships of the Cenozoic
deposits of China, and therefore they carry great weight.

Proboscideans:
Trilophodon connexus Hopwood
Trilophodon wimani Hopwood (Pontian, according to Teilhard)

Andersson,' in his discussion of the Cenozoic of North China, describes the Lu Tzu Kou beds, exposed in the

Pao Te Hsien area of Shansi. According to Andersson these beds, discovered by Zdansky, are of Upper Cenozoic
age, but they underlie the Hipparion clay. Yet although the Lu Tzu Kou beds are exposed beneath the Hip-
parion beds, there is a strong possibility that the age difference between the two series is not great.

As the Hipparion beds represent the trau.sition from Miocene to Pliocene, tlie Lu Tzvl Kou beds could eventually be sup-
posed to represent the Miocene, but there are some facts at hand which indicate that they are in age nearly related to the
Hipparion beds.

'Andersson, J. G., 1923. Mem. Geol. Surv. China, Scr. A, No. 3, p. 107.
 .

GEOLOGIC SUCCESSION: ASIA 1481

4. THE PLIOCENE OF MONGOLIA

Khunuk Formation. — The Khunuk formation was described by Granger' as follows:

In the Kholobolchi Nor region of Mongolia there are three areas of late deposits considered, in the field, as being of Pleis-
tocene age; the geologists were of the opinion that these three exposures were of the same age, and the name 'Khunuk' was
given to the formation. The three exposures mentioned in the Kholobolchi Nor region are respectively north, northeast, and
east of the lake; the type of Serridentinus florescens comes from the northern exposure of the 'Khunuk' and represents the only
specimen obtained in that particular locality.

As Osborn has shown, it is possible that the Khunuk formation is approximately correlative with the Hung
Kureh beds of the Tsagan Nor region, which latter deposits contain a small mammalian fauna of Pliocene re-
lationships. The single mastodont described from the Khunuk formation is an advanced type of serridentine,

which would favor the assigning of a Pliocene age to these sediments.

It is not possible to say at what stage in the Pliocene the Khunuk and Hung Kureh formations should be
placed, but it is very probable that they are in the main Pontian equivalents, representing a phase in the sedi-
mentary history of Mongolia immediately subsequent to the period during which the very fossiUferous Tung Gur
Upper Miocene sediments were deposited.

Proboscidean: Ocalientinus {Serridentinus) florescens (Osborn).

5. THE PLIOCENE OF NORTH CHINA

LOWER PLIOCENE
The Hipparion clays of North China form a mantle covering extensive areas of valley and plateau country,
dissected by streams. The localities at which fossils have been discovered in the Hipparion clays are so numerous
that they cannot be listed here, nor can comparisons between them be attempted. It might be well, however, to
quote Andersson's^ description of the Hipparion clays at Chi Chia Kou in Pao Te Hsien, one of the most richly

fossiUferous and best-known locaHties in this horizon.

The Chi Chia Kou region a plateau land dissected by an intricate system of ravines, in the bottom of which the sub-
is

stratum of the Hipparion clay almost everywhere visible. The basement rock is formed by the Carboniferous coal series in
is

nearly horizontal beds. The basal layer of the Hipparion series is a conglomerate bed, at most 4 meters thick with gray matrix.
Above this basal conglomerate rests the red Hipparion clay with a maximum thickness of 65 meters. In the clay there are inter-
bedded gravel beds of little persistence and occasionally also lenses of sand. In certain horizons there are also irregular lime con-
cretions in the clay. Round the fossil mammal bones are seen infiltrations of lime.
In the 65 meters of red, mostly entirely barren clay there is a well defined bone-carrying horizon 25 meters above the bottom
and 35 meters underneath the top of the deposit. The bone layer is mostly less than a meter thick and so nearly horizontal
that there is not more than 5 meters variation of altitude of the bone bed throughout the whole Chi Chia Kou area. In the
bone horizon there are not bones everywhere, but rather pockets or nests rich in bones separated by some meters of barren clay. .

According to Dr. Wiman and Dr. Zdansky the Hipparion fauna indicates steppe conditions, but the occurrence of Giraffinse
and Suidae points to the existence of groups of trees and of water pools spread over this steppe. As the clays are mostly barren
over wide areas and rich in mammals only in the three areas mentioned, it might be inferred that in the otherwise very dry
steppe there were locally oa.ses with trees, water pools and occasionally also sheet floods after the rains. It seems as if these
genial conditions prevailed only during a short, well defined period.

•Granger, Walter, in H. F. Osborn, 1936. Volume I of this Monograph, p. 397.

^Andersson, J. G., 1923, op. cit., pp. 107, 108, 110.

1482 OSBORN: THE PROBOSCIDEA

Proboscideans: (Pontian of China)

Serridenlinus mmani (Hopwood)

Serridentinus lydekkeri (Sclilosser)
Trilophodon spectabilis Hopwood
Tetralophodon exoletus Hopwood
Tetralophodon sinensis (Koken) (This horizon?)
Pentalophodon sinensis Hopwood = Anancus sinensis {fide Osborn)
Pentalophodon cuneatus Teilhard and Trassaert
Zijgolophudon borsoni (Hays) (This horizon?)
Mastodon americamis (Kerr) (This genus and species?)
"Mastodon" sp.

Mastodon interme.dius Teilhard and Trassaert

Stegodon licenti Teilhard and Trassaert
Stegodon zdanskyi Hopwood (This horizon?)

MIDDLE PLIOCENE
Only within recent years has the Middle Pliocene been recognized in North China. The distinction of this

stage has been difficult because it is hthologically closely related to the typical Pontian deposits, and particularly
because the fauna is not well known. In their important contribution entitled "The Pliocene Lacustrine Series in
Central Shansi," Licent and Trassaert' describe sediments overlying the Pontian beds but underneath the San-
menian or Lower Pleistocene.

These authors have shown that the lacustrine sediments of Central Shansi occupy an extensive, shallow
Permo-Triassic syncline. The beds representative of the Upper Cenozoic are 100 meters or more in thickness and
entirely of a freshwater type. They are characterized by the "extremely deceptive recurrency of the same
facies: rusty or reddish sands, green marls, etc.," thereby making the differentiation of the several horizons con-
tained witliin the series extraordinarily difficult. Nevertheless, Licent and Trassaert recognize three zones
within these lacustrine sediments. The lowest zone, designated as Zone 1, is of Pontian affinities. Above it is

Zone 2, of Middle Pliocene age, succeeded by Zone 3, representing the Sanmenian. In addition it is pointed out
that there is very likely a fourth zone, coming between zones 2 and 3, representative of a stage between the
Middle PUocene and the Sanmenian.

Zone 2. In the following zone 2, better studied in the Changtsun basin, the deposits become less coarse, and a typical
lacustrine condition is prevailing: green and bluish marls, containing many bird, turtle, fish-remains, freshwater shells {Lym-
nsea, Planorbis, thin-shelled Unionidx) and carbonised plant-remains. A small Hipparioii, the Chilotherium, a tapiroid
Mastodon and the Stegodon are still present (as in zone 1). But a remarkable type of strepsiceros Antilope (cf. Aiitilospira
T. & Y.) seems to appear for the first time, and also a special Castor (characteristic of the Ertemte fauna of Mongolia) Dipoides
majori Schl. A middle Pliocene age seems to be indicated.
Suggestion of a fourth sedimentary zone. The pala?ontological analysis of the fauna collected in the Yiinchu basin suggests
that, between zone 2 and zone 3 a fourth horizon might be eventually recognised in the Pliocene deposits. First, collected by
country people, from such localities as Malan, w.e have, embedded in a characteristic matrix of a dark red hue, canon-bones of
a big Hipparion, already associated with a Bison, but without any sure trace of Equus. Furthermore, amongst the best fos.sils
purchased in the area, are several teeth of a thick lamelled Elephant {Elephas cf. planifrons), a form never observed in Nihowan,
and yet too much advanced for being conveniently referred to the M
astodon-Stegodon fauna of zone 2.
would seem therefore that the lower part of zone 3 might have to be separated as a special
It unit, distinguishable from the
true Nihowan beds by the absence of Horse and the presence of archaic Elephant.
This point however is not yet supported by clear stratigraphical evidences.

'Licent, E., and M. Trassaert, 1935. Bull. Geol. Soc. China, XIV, No. 2, pp. 214, 216.
 :

GEOLOGIC SUCCESSION: ASIA 1483

UPPER PLIOCENE (?)

Teilhard and Trassaert,' in 1937, divided the Pliocene (as they considered the extent of its upper and lower
limits) into three "zones," namely: (1) A lower zone comprising the Pontian Hipparion beds, (2) a middle zone of
Pliocene age but subsequent to the Pontian, and (3) an upper zone comprising the Villafranchian, wliich these
authors placed in the Upper Pliocene.

In addition to the characteristic "Middle Pliocene," as recognized by Teilhard and Trassaert, and as described
by Licent and Trassaert (see above), there would seem to be an upper zone or subzone which is immediately
antecedent to the base of the ViUafranchian. Since the Villafranchian is considered as Lower Pleistocene (as will
be shown below) it would seem logical to suppose that this upper portion of "Zone 11" represents the uppermost
Pliocene in North China. Teilhard and Trassaert made the following comments concerning it

But, near C'hinglo (N. Shansi, cf. Teilhard and Young, 1931, p. 52, fig. 15), in an horizon representing clearly Zone II
(rather large Deer and Hipparion, Antilospira, no Equus .), Teilhard and Young have collected several years ago an isolated
.

lamella of D3 decidedly referable to a primitive Elephant, and not to a Stegodont. An upper horizon (planifrons subzone) might
therefore have to be recognised some day at the top of Zone II.

Proboscideans: (Middle and Upper Pliocene of China)

Zygolophodon borsoni (Hays)

Mastodon intermedius Teilhard and Trassaert
Stegodon zdanskyi Hopwood
Stegodon orientalis Owen (This zone?)
Archidiskodon planifrons (Falconer and Cautley) (Uppermost Pliocene)

6. THE PLEISTOCENE OF NORTH CHINA

The Pleistocene of China is of great importance, not only because it has yielded large and rich mammalian
faunas that are directly ancestral to the modern faunas of northeastern Asia, but also because it has produced
some of the most complete and the most significant remains of fossil man. For this reason, a rather complete and
detailed discussion of the Quaternary beds of China will be necessary.

The Villafranchian of China. —The Pleistocene succession of North China may be regarded as beginning
with the Villafranchian deposits of Nihowan, and also possibly of Yushe. This assignment of the Villafranchian of
China to the beginning of the Pleistocene is not entirely in accord with the views of Teilhard,- who recognizes the
fact that there is a growing sentiment among vertebrate palaeontologists to regard the appearance of Equus,
elephants, and cattle as truly indicative of the beginning of the Pleistocene throughout the world, but who feels

that in China it is most convenient to end the Tertiary sequence with the diastrophic movements that resulted in

the cutting of the gorges, subsequent to the deposition of the typical Villafranchian sediments. Consequently, he
would regard the accumulation of the Choukoutien deposits and the first appearance of man as indicative of the

beginning of the Pleistocene in North China.

In North China, the major period of gorges cutting (Fenho stage) first placed by Bailey Willis at the base of the Malan
loess, and later transferred by Barbour and myself to a pre-Nihowan stage (2), would find its true place in an intermediate
position, namely between the Sanmenian (Nihowan) and the Choukoutien stages of deposition. . . .

'Teilhard de Chardin, P., and M. Trassaert, 1937. Palaeontologia Sinica, Ser. C, XIII, Fasc. 1, p. 53.

^Teilhard de Chardin, P., 1937. Bull. Geol. Soc. China, XVII, No. 2, pp. 173-175.
1484 OSBORN: THE PROBOSCIDEA

In North China, again, (just as in France and also in North India), the first appearance of horse (more and more generally
accepted by the palaeontologists as indicating the base of the Pleistocene in the Old World) antecedes clearly the major diastro-
phisms which would be the best limit between the Tertiary and the Quaternary from a geological point of view.
Therefore, the advantage of keeping the Villafranchian (first 'Equus-beds'} in the Pliocene, instead of referring it to the
Pleistocene, should be re-considered. So far as North China is concerned, the most natural base of the Quaternary is given by
the appearance of Man on a modernized topography, after the Villafranchian.

YusHE. —According to Teilhard,' the basal portion of the Villafranchian of China is to be found in the Yushe
basin, and consequently he has distinguished this part of the Pleistocene sequence as a separate subzone within
the Villafranchian.

Strangely enough, no Archidiskodon has been recorded up to now from the typical Villafranchian formation of Nihowan.
In order to explain this difference (if it really does exist), we may suppose that the Nihowan beds represent only the top of the
Villafranchian {'riamadicus subzone?'), the base of the Villafranchian being on the contrary only present in Yushe, and
characterized perhaps there by the latest Archidiskodon {'tokunagai subzone'?). . .

In Yushe, as well as in Nihowan, the Stegodonts seem to have disappeared before the beginning of the Villafranchian.
Yet, some new finds made in S. Shansi suggest that the group (represented by Stegodon orientalis) has lasted as far up as the
Lower Pleistocene inclusively along the northern border of the Tsinling, as it did in Central and Southern China (Szechuan, etc.).

This definition of a basal zone of the Villafranchian would seem to be in accord with the evidence put forward
by Licent and Trassaert for a distinct zone between the Middle PUocene and the Villafranchian of Central Shansi-
Since the relationships of this distinct subzone or horizon in Yushe is with the overlying Villafranchian, rather
than with the underlying Pliocene sediments, it would seem best to place it as the first member of the Pleistocene

group of sediments in North China.

Proboscideans:
Palxoloxodon tokunagai (Matsumoto)
Palseoloxodon namadicus (Falconer and Cautley)

Nihowan (Sanmenian). —The true Villafranchian of North China is best exemplified by the sediments and
their contained fauna as exposed in Nihowan. Teilhard- has shown that originally only two distinct phases were

recognized in the Upper Cenozoic of North China, namely, the Pontian Hipparion beds and the Pleistocene loess.
Subsequently, as the result of more detailed work in the North China area, a series of post-Pontian, pre-Loess
deposits came to light, occupying the interval betv/een the levels originally known.

But then again the first impression of the geologists was that they had to deal with a single polymorphous complex. On
one hand, no internal disconformities were observed, at that time, in the newly recognised sedimentary block. And, on the
other hand, the common occurrence in all these 'post-Pontian and pre-Loessic' deposits of closely related types of Horse, Hy-
aena, Rhinoceros, Rodents and MoUusca suggested for all of them a same general age.
Thus was borne the conception of the 'Sanmenian,' a single stratigraphical and faunistical unit, rather vaguely referred
either to the Late Pliocene, or to the Lower Pleistocene.

As Teilhard has shown, there is a definite faunal and physiographic break between the Nihowan and Chou-
koutien phases, hitherto included in the Sanmenian. Consequently, he has felt it necessary to limit the term
Sanmenian to the Nihowan deposits and their included fauna, thereby making the Choukoutien formation
definitely post-Sanmenian in its relationships. In accordance with his views as to the Upper Pliocene age of the
Villafranchian, Teilhard has placed the Sanmenian as the latest part of the Pliocene, relating it to the lacustrine

Middle Pliocene sediments of North China, and separating it quite definitely from the overlying Choukoutien
phase. It is proposed here to regard the Sanmenian as a lower phase of the Pleistocene sequence in China (preced-
ed perhaps by the basal Villafranchian beds of Yushe) and related to the Choukoutien deposits. Of course, in

'Teilhard de Chardin, P., and M. Trassaert, 1937. Palaeontologia Sinica, Ser. C, XIII, Faso. 1, p. 53.

^Teilhard de Chardin, P., 1937. Bull. Gaol. Soc. China, XVII, No. 2, p. 170.
 GEOLOGIC SUCCESSION: ASIA 1485

following this line of procedure, the weight of the argument is being given to the fauna! evidence, as based on the
appearance of new and advanced Pleistocene genera, rather than to the physiographic evidence cited by Teilhard,
as based on diastrophism and the rejuvenation of the streams.

It all comes down to a choice between two lines of evidence which do not coincide, as shown by the following
diagram.
1486 OSBORN: THE PROBOSCIDEA

Various authors have described the several groups of mammals constituting the Choukoutien fauna, so that
the detailed information as to this mammalian assemblage is voluminous, to say the least. Recently Teilhard has
sunnnarized the Choukoutien manunalian fauna by showing that from the standpoint of its geological significance
the following classification may be formulated.

1. Forms that were present in the Sanmenian of Nihowan (Villafranchian), but which did not persist into
Choukoutien times.

2. Nihowan forms present in the Choukoutien deposits, but absent in the subsequent Malan Loess.

3. Loess forms present in Choukoutien, but absent in Nihowan.

4. Loess forms absent from Choukoutien.

5. Forms that were present in all three formations.

The distribution of the significant mammaUan types in the three formations enumerated above may be
shown graphically in the following manner.

Villafranchian Late Pleistocene

Sanmen-Nihowan Choukoutien Malan Loess
Mustela pachygnatha-^
Elasmotherium
Postschizotherium —
Hipparion
Eucladoceros
Rusa
Ochotonoides —
Siphneus tingi-
Machairodus — Machairodus
Hyxna cf. sinensis — Hyxna cf. sinensis
Equus sanmeniensis— Equus sanmeniensis
Rhinoceros cf. mercki- Rhinoceros mercki
Ovibovids Ovibovids
Nyctereutes -
Nyctereutes Nyctereutes
Meles Meles Meles
Camelus Camelus Camelus
Gazella Gazella Gazella
Spirocerus— Spirocerus Spirocerus
Bison— Bison Bison
Ovis Ovis Ovis
Cyon Cyon
Siphneus ci.fontanieri- Siphneus d.fontanieri
Euryceros Euryceros
Bubalus Bubalus
Ilysena spelsea

Equus hemioniis
Cervus elaphus
Bos primigenius
 GEOLOGIC SUCCESSION: ASIA 1487

From this it will be seen that the Choukoutien fauna is truly intermediate between the Villafranchian fauna
of Nihowan and the late Pleistocene fauna of the Malan Loess. As to the relationships of the Clioukoutien mam-

malian assemblage to the faunas either above or below it, Teilhard and Young' made the following statements.

Taking the facts more in detail, tlie Choukoutien types in many cases are distinctly different from their corresponding
Nihowan or Sjara-osso-gol form.«, being less primitive than the former, and less advanced to some extent than the latter. For
example: Hyaena cf. sinensis, Rhinoceros cf. tichorhinus, Spirocerus wongi, Bison palaeosinensis, Ovis shantungensis, from
Nihowan are more primitive than Hyaena sinensis, R. tichorhinus, Spirocerus peii, Bison sp., Ovis cf. ammon from Choukoutien.
On the contrary. Hyaena sinensis, Euryceros pachyosteus, Bubalus teilhardi from Choukoutien can be held as more archaic than
Hyaena spelaea, Euryceros ordosianus, Bubalus wansjocki from the Loess.
The conclusion is obvious. In full accordance with the stratigraphical facts which will be set forth subsequently below in
section 5, the Choukoutien fauna fits so exactly between the Late PUocene (Nihowan) and the Upper Pleistocene (Sjara-osso-
gol) ages that it is not easy to decide to which of them it stands more closely related. The Choukoutien formation has conse-
quently broadly to be considered as of Lower Pleistocene age.
In order to account for the change between the Nihowan and the Choukoutien faunas, a long period of time must be allowed
during which some climatic changes most probably took place. The fauna is still dominantly of a palaearctic type, but with a
clearer tendency to break into a special east Asiatic type. Some southern migration, possibly along the sea coast, may be sug-
gested by the appearance of the Bubalus.

Recently Teilhard has placed the Choukoutien deposits in the Middle Pleistocene, as a result of field work and
correlative studies made throughout southeastern Asia. By following this procedure, the Choukoutien deposits
are brought into line with the cave deposits of southern China and Burma, described in a preceding section of this
chapter. On the basis of present evidence, this would seem to be the most logical correlation for the Choukoutien
beds. There seems to be but little doubt that there was a widely spread Lower Pleistocene fauna in Asia and the
Upper Irrawaddy, Djetis, Nihowan) followed by an equally extensive Middle Pleistocene
Orient (Upper Siwalik,
complex, which upon the mainland is commonly preserved in caves (Mogok, Hoshangtun, Trinil, Kwangsi,
Szechwan, Choukoutien).

The Loess of North China. —At the top of the Cenozoic succession in North China is the Loess, a character-
isticand wide-spread deposit fornung a prominent and easily-recognized capping layer over a large area north of
the Yellow River. Fossils of an advanced type have been found in the Loess, and for this reason the deposit is
generally regarded as of Upper Pleistocene age. The following remarks were made by Andersson,^ in 1923, re-
garding the mammalian and avian fauna of the North China Loess.

The most common mammal remains in the loess are tusks and molars of an Elephas, which according to Dr. Zdansky may
possibly be Elephas namadicus, which was originally found in the Pleistocene alluvium of the Narbada valley of India, where it
occurs together with two species of Hippopotamus, and several other mammals. The reappearance of this Indian species in the
North China loess would be very surprising, especially in view of Richthofen's eolian theory. However, Dr. Zdansky's pro-
visional determination has to be tested by a much closer study of the specimens. At any rate, it is beyond doubt that the com-
mon loess elephant is not the Mammoth as has formerly been suggested.
Among the numerous isolated finds which have been made in loess-like material the following species can with fair safety
be assigned to the loess: Rhinoceros affinis simus, Ovis? sp.. Hyena sp., Ursus sp. A skull of a Castorid also was obtained from
undoubted loess, and this is another find which tends to weaken Richthofen's eolian theory. The same appUes to a recently
found Sus sp.
Only a single case, have we ever come across what deserves the name of a small bone accumulation in the loess. This
in
was in SE
Shansi, in Yuan Chii Hsien, and the locality has been studied by Dr. Zdansky who has communicated the section
fig. 39. In this place were found: Hyena, Equus, Cervus and a turtle. The deer, a forest animal, and the turtle, a water
animal, are further finds which hardly agree with Richthofen's eolian theory.
Lastly we have to mention a fossil which seems to conform better with the said theory, namely the egg shells of an Ostrich,
a bird described under the name Strulhiolithus. The modern ostrich is a steppe bird, and the mere mode of occurrence of these
. .

unbroken shells, often two or more together, seems to indicate that the nests were occasionally covered by a wind-drift deposit
which prevented the eggs from maturing and preserved the shells in an unbroken state.

Earlier authors attributed a great thickness to the loess deposits of North Cliina —in some cases as much as
1,500 feet. According to Andersson, however, the loess never exceeds a thickness of more than 50 or 60 meters.
'Teilhard de Chardin, P., and C. C. Young, 1933. (In Black, Teilhard, Young and Pei), Mem. Geol. Surv. China, Ser. A, No. 11, pp. 48, 49.
^Andersson, J. G., 1923, op. cit., p. 127.
 —

1488 OSBORN: THE PROBOSCIDEA

Andersson's revised figures for the thickness of the loess are based on the fact that he differentiated the older beds,
which the earlier students had confused with the loess and included in it.

The North China, named the Mnlan, has been divided by Andersson into two beds or horizons. The
loess of

lower deposit he has called the primary loess or the true loess. The upper bed is the secondary loess or the re-
deposited loess. As is indicated by the name, this secondary loess is merely material from the jirimary loess,

reworked in very late Pleistocene times.

The redeposited loess, because of the taxonomic relationships and the slight mineralization of its contained
fossils, has been defined by Andersson as a post-loessic deposit. Thus it may be considered as an horizon bridging
the time between the latest Pleistocene and the beginning of Recent conditions in North Cliina.

Lately Teilhard and Young have thrown new light on the problem of the post-loessic faunas of North China
in their study entitled, "On the Mammalian Remains from the Archaeological Site of Anyang." These authors
show that in the ancient city of Anyang, capital of the Shang dynasty during the period between 1400 B.C. and
1100 B.C., there were numerous mammals similar to the post-loessic fossils found in other parts of Cliina. In
their analysis of the Anyang fauna, Teilhard and Young list three faunal divisions, namely:

1. The wild indigenous animals, such as the raccoon-dog, bear, badger, tiger, bamboo-rat, hare, water-deer
and the sika.

2. The domesticated animals, including the pig, dog, sheep, goat, cattle, water-buffalo, and the macaque.

3. Imported mammals, the whale (probably bones collected along the seacoast), the elephant, tapir, and
a small bear.

The presence of the elephant, which has been identified as Elephas indicus, in North China is interesting,
to say the least. The evidence would seem to show, however, that this elephant was never wild here, but rather
was imported (probably in the form of tribute) by the ancient peoples of this old city.

As Teilhard and Young' have remarked, "Such beds [the redeposited loess and similar deposits] are still,

geologically, a 'no man's land' more attention should be paid in [the] future to the redeposited
; loess, to peat-bogs
and to subrecent cave deposits."

The difficulty in dealing with these redeposited loess and post-loessic beds is that they really do bridge the
period between the Pleistocene and the present day. Consequently such beds are defined only in a most provision-
al manner. As Teilhard and Young have shown, if Rhinoceros, Elephas namadicus or Elephas primigenius, Hysena,
and Euryceros are present, the deposits may be called Pleistocene with some degree of assurance. On the other
hand, there is no reason why some of these forms may not persist beyond the closing of the Pleistocene period
whenever that might have been. Therefore the problem resolves itself into a question of drawing an artificial

line across a series of continuous events which are so close to us that it is not possible for us to gain a true perspec-
tive as to their imi^ortance or their interrelationships.

Proboscideans:
Palseoloxodon namadicus (Falconer and Cautley) Loess
Elephas primigenius Blumenbach Upper Pleistocene, North China, Mongolia
Elephas indicus Linn.
'Teilhard, P., and C. C. Young, 1936. Palaeontologi.a Sinica, Ser. C, XII, Fa.sc.
Sub-Recent, Anyang
1, p. 57.
I
LIST OF THE FOSSIL LAND MAMMALS OF JAPAN AND KOREA
1490 OSBORN: THE PROBOSCIDEA

7. THE MIOCENE TO PLEISTOCENE OF JAPAN

Japan. — The fossil inanniials of Japan, thougli for the most part very poorly i)reserved, have been found at
numerous localities, showing that Cenozoic continental deposits, particularly those of the Upper Cenozoic, are
widely distributed throughout the islands. Most of the mammal-bearing localities are on the island of Honshu,
and a majority of these deposits are of Pleistocene age. In all of the Japanese deposits, whether of late Tertiary or
of Pleistocene age, the proboscideans are the dominant elements in the faunas.

The evidence of the fossil mammals would seem to indicate that Japan was connected with Asia during
Miocene, Pliocene, and early Pleistocene times. Certain elements of the Miocene fauna of Asia reached Japan,
and in subsequent times there was likewise a constant infiltration of new migrants from the mainland. This
influx from Asia into Japan reached its height seemingly during the early part of the Pleistocene, for at that time
numerous proboscideans as well as certain other forms found in Japan were specifically identical with mainland
types.

An interesting feature of the Japanese faunas is the dominance of the proboscideans, especially during Pleisto-

cene times. Whether this dominance of the Proboscidea is real or illusory is a question difficult to decide on the
basis of extant evidence. That is, there may be many proboscideans in the Japanese deposits because these fossils

are large and more apt to be preserved than are more delicate, smaller mammalian types. Or it may be that the
proboscideans actually effected a crossing from the mainland to the islands where other smaller, less adaptable
mannnals failed.

Recently Dr. Tokunaga' pubUshed a compendium of all the fossil land mammals found in Japan and Korea.
His list of genera and species, with their horizons and localities, is repeated here (p. 1489) in so far as it bears
on Upper Cenozoic forms.

VI. NORTH AMERICA

1. INTRODUCTION
It would seem that the Proboscidea first reached North America from the Old World in Upper Miocene times,
and from that time until the end of the Pleistocene, these animals played an important role in the development of

the mammalian faunas of the New World. Moreover, the evidence of recent discoveries shows that certain of the
mammoths and mastodons did not become extinct until after man had crossed from Asia into America and was
well established in his New World home.
In the following pages the relationships of the various Proboscidea-bearing formations of North America will
be discussed. Only those beds in which proboscideans are actually recorded will be included in the discussion,
and they will be grouped by their ages, according to whether they are of Miocene, Pliocene, or Pleistocene affinities.

In considering the Miocene and Pliocene beds, the several formations as named according to prevalent American
usage will be described and compared. To give the discussions a certain degree of order and comprehension, the
formations will be grouped according to their occurrence in the eastern portion of North America, in the Great

Mountain and Pacific Coast region respectively. The treatment of the Pleistocene must
Plains rogif)n, and in the
of necessity be somewhat different. Here the sequence of tlie glacial and intcrglacial periods in North America
in relation to the occurrence of Pleistocene proboscideans will be discussed.
'Tokunaga, Shigeyasu, 1933. .\raer. Mus. Novitates, No. 627, pp. 2-4.
 GEOLOGIC SUCCESSION: NORTH AMERICA 1491

2. THE UPPER MIOCENE: BARSTOVIAN

Pawnee Creek Formation, Colorado. — Dr. Matthew, in his first description of the Pawnee Creek beds of

northeastern Colorado, recognized two fossiliferous layers or zones within the formation, but at that time he
pointed out the fact that the separation between these zones, either stratigraphically or faunistically, was very
inconstant, so he was inclined to regard the zoning of the Pawnee Creek beds as of little importance. Later, in
1909, in 1918, and in 1924, he regarded the Pawnee Creek formation as a unit, containing a single fauna.

^EOlpqrCAL KEl^[0?/iHIP3 OF NORTf< 2^^JIIC24^^^P/1030SCIDEA^

PACIFIC COAST INTERMONTANE PLATEAUS GREAT PLAINS AND MOUNTAINS EASTERN U.S.

NEVADA OREGON MEXICO

•KA'p'<^-icc\ ARIZONA TEXAS COLORADO NEBRAS KA
CALIFORNIA CALIFORNIA IDAHO NEW MEXICO OKLAHOMA KANSA5
(WASHINGTON) Parelephas loashinatonii fMasfodon americanus, Parelephas colu bi, Archidiskodon imperafor, Mammontcus prim aeniusj i

Parciephas celsi Archidiskodon maibeni, scotti, fiaut.mertd

Parelephas
?RhqncKothcrium Archidiskodon
.

columbi tclicis naroldcooki Morrillta barbouri, ional;s_ nebpasc^nsis

Wideli^ disfribufed ArchidisKodon francisi Mastodon ^ranaeri. [INDIANA)
over N. America. sonoriensis, Stego rna stodon Mastodon mooaiei, Mastodon
Mastodon imperator. Mastodon priesMeui Steqo mastodon acutidens
oreaonensrs si Ivestn's, raki Cordillerion YlOWA) mirincus, Parelephas
Archidiskodon irnperator qratum Mastodon pro^enius Steaomastodon Jeftersonii
exilis ralconcri sAJsr ore^omasrodon artoniae primitivus
TEHA.MA HAGSKMA/^ Cordillerion PEDRO Horizon uncertain \
Horizon uncertain (S.CAROLrNA)
tropicus, Sfegomasrodon Cordiltenon orarius Trilophodon li^oniferus
Redepostted in
Stcgomastodon ^teg oma sfodon olicjobunis^, ari zonae C. defloccatus PLEISTOCENE
Tori^nobelodon loomisi
Ujui
cf. arizonae mirirtcus ii'^ntiquissimui
(( felicis,
Cordtilerion
bensoncnsts
RLA ATCO I

\
Rhi^nchotfieriurn^ Talconen
Amebeiodon frtcki Ocalientinus
Amcbelodon sinclairi oblit^uidens
« infermedius Serbelodon praccursor Gnathabelodon thorpei Ocalientinus
u proqressus Steoomastodon minficus
S. successor
emmon si
S. texanus
Horizon uncertain a/t: edejs^ THOUSAAfD CR. RATTLESNAKE Rhunchotherium HEMPHILL OPTIMA
kl
Rht^nchoHierium Rhi^ncKothcrium Miomastodon tiascalae,
sbepardi shepardi edcnse merriami broiuni
Cord llerion
I Pliomastodon
edensis nevadanus

B rCHEGO/JV
Pliomastod on
vexi llarius (FLORIDA)
ORINDA R/CARDO a CEDAR MT Serrtdenhrius
serridens
CLARENDOlsr
Serridentinus
WRAY UPPER SMAKE
Pliomastodon mattheiui
CR. ALACHUA
BONE VALLEY
Serbelodon Trilophodqrj Amebeiodon Rhunchotherium an^uirrvale Serridentinus
burnhami productus paladentatus
CHAJVAC dinoiherioidcs S. serridens Serridentinus an^uirivalis floridanus
S. serrjdens A. hicksi S. nebrascensts S. simplicidens
fGUATEMAL^Scrridcntinus ^uatemalensis Ci marronis VALENTINE S. breujsterensis
[HONDURAS) Blickotherium bllcki Serbelodon Trilophodon abeli Pliomastodon
Anbelodon honaurenais praecursor T ujillistoni sellardsi
Te trilophodon T ph.ppsi Ocalientinus
MONTANA! SANTA fricki \.

T
qregorii
^
giganteus
•"
qa n te u s 0-
floridanus
bifoliatus
CANYO/sT BAR^TOfV TRUCKEE MADISON FE REPURLiCAU KIVER,
Serridcntinus VALLEY Serridentinus productus SeAeloX^n
barbourensjs {-VALENTINE. PUOCENE)
barstonrs S. pojoaquensis
Eubclodon Blickotherium cuhtipodon
Trilophodon cruziensis morn In Tetralophodon campestcr
CUYAMA Ocattenftnus ojocaliensis Megabelodon
lulli
Serridentinus prpgressus
Trilophodon dinotherioidcs
Trobelodon taoensis
Torunobelodon Ocalientinus republicanu*
Trilophodon rio^randensis barnunbroujni
Amebeiodon joraki
N. COAU?/GA VIRGIN SKULL DEEP RIVER PAWUEE CR.
VALLEY SPRING Rhunchotherium Serridenrinus proavus
Rhunchotherium recfidens
Miomastodon brevidens Miimasfodon merriami
merriami (MARYLAND)
SASKATCHEWAN BKOWNS PARK Horizon uncertain
ivoon MT. Trilophodon iricki Trilophodon
obscurus
Itrlu'ttt BnuUtt/

Figure 1224

Professor Osborn, however, held to the view that the Pawnee Creek formation consists of two levels, which he
designated as "A" and "B," for the lower and upper divisions respectively. This view was expressed in 1918, in
the monograph on the Tertiary Equidse of North America, and will be found repeated in Volume I of the present

Memoir.

In 1924 Matthew clearly demonstrated the virtual equivalence of the Pawnee Creek beds with the
Lower Snake Creek beds of Nebraska. The correspondence of the two formations is indeed close, not only in the
constitution of their mammalian faunas, but also in the fact that they are located only a hundred miles or so apart.
Therefore there is every reason for regarding them as al)Out contemporaneous with each other.

The Pawnee Creek formation and Lower Snake Creek would seem to be representative of the beginning of
the Upper Miocene in North America. As Matthew has pointed out, certain elements in the faunas, particularly
1492 OSBORN: THE PROBOSCIDEA

the horses, show the admixture of typically Middle Miocene types with more progressive forms that foreshadow
to some extent Phocene species.

Proboscideans in the Pawnee Creek formation are extremely scanty, and it is interesting to note that they
are completely absent from the closely related Lower Snake Creek beds. Perhaps these beds were deposited soon
after the proboscideans first reached America, so that the group had not had an opportunity to become widely
spread.

Proboscideans:
Serridentinus proavus (Cope)
Rhynchotherium rectidens Osborn
Miomastodon merriami Osborn

—
Brown's Park Formation, Colorado. The Brown's Park formation, located in the extreme northwest
corner of Colorado, was fully described by Peterson in 1928, in his comprehensive memoir on the Brown's Park
Fauna. According to Peterson, this formation consists of "soft, almost chalk-white sands and sandstones cement-
ed with hme, sometimes intercalated with slightly argillaceous bands."

The lower part of the formation, consisting of a rather hard, resistant sandstone, was named the Weller
sandstone by Peterson, and it is at this level that the fossil mammals representative of the formation were found.

At its earUest, the Brown's Park formation is possibly correlative with the Sheep Creek of Nebraska, of
Middle Miocene age. The presence of a primitive procyonid, a chalicothere (chahcotheres extend upwardly into
the Sheep Creek in Nebraska), and an oreodont probably related to Ticholephis, would be evidence in favor of
this correlation for the assemblage. The presence of Trilophodon, however, may be an argument in favor of
a somewhat more advanced position for the fauna, that is, roughly equivalent to the Deep River, Pawnee Creek,
and Lower Snake Creek formations.

Proboscidean: Trilophodon fricki (Peterson).

—
Santa Fe Beds, New Mexico. For many years the Upper Tertiary sediments around Santa Fe, New
Mexico, have been known as the Santa Fe marls, and as a result of the early explorations and publications of Cope
these beds have come to be regarded as of Upper Miocene age. In 1909 Matthew listed the Santa Fe as being more
or less the equivalent of the "Fort Niobrara" and Little White River beds in Nebraska and Dakota, and of the
Clarendon in Texas. This would indicate a possible position in the Lower Pliocene, rather than in the Miocene.

During recent years the expeditions sponsored by Mr. Childs Frick have made extensive collections in the
Santa Fe beds, and much new material, far finer than any hitherto discovered, has been found. In an extensive
paper on Tertiary proboscideans of North America Frick has described and figured some extraordinarily complete
Santa Fe mastodonts, which arc hsted below.

Prick's work in New Mexico would seem to give evidence for the fact that the Santa F6 beds are not a unit
horizon, but rather cover a considerable period of time, beginning in the
Upper Miocene and extending throughout
an apjireciable portion of the Pliocene.
Th(> proboscideans discovered by Frick come from an intermediate horizon
within the Sante Fe, which, because of the presence of Hipparion, is very probably of Lower Pliocene age, being
more or less equivalent to the Republican River and Upper Snake Creek beds to the north. Above this zone
 :

GEOLOGIC SUCCESSION: NORTH AMERICA 1493

carrying Hipparion and proboscideans are beds that Frick regards as of Upper Pliocene age. Frick's' remarks as
to the age of the Santa Fe are as follows

The deposits of the Santa Fe basin of northeastern New Mexico, widely known in the literature as the Santa Fe marls, have
been currently interpreted as of Upper Miocene age. More recent investigation indicates that the accumulations of this
portion of the Rio Grande basin range from the Mid-Miocene to Pleistocene. The Pleistocene occurs in remnants of aeolian ori-
gin that here and there cap the irregular Pliocene-Miocene surface. While no mastodonts have so far been encountered in limit-
. .

ed exposures of probable Uppermost Pliocene facics, their remains are fairly common in the upper half of the earlier deposits.
A possible time equivalent of the Little White River of South Dakota, the Republican River of Kansas and the Upper Snake
Creek of Nebraska is indicated by the presence in certain locaHties of species of Hipparion and of advanced Pliohippus. . . .

The mastodont forms of the Ojo Caliente and the Santa Cruz sections of the Santa F6 basin are tentatively interpreted as of
the Hipparion zone.

More "The Lower Snake Creek, Pawnee Creek, Santa Fe and Barstow
recently (1937) Frick- states that:
ostensibly include a number of overlapping phases of the main Late Tertiary." According to this author's classifi-
cation of Tertiary sediments, the formations named above might occur in the interval between Middle Miocene
and about Middle Pliocene times.

Within the past year Charles S. Denny' has published a study of the Santa Fe formation in its type locality,

principally in the Espanola Valley, north of the city of Santa Fe. This is strictly a geologic and sedimentation
study, so that the question of the contained vertebrate fauna and its probable age is not considered. Denny con-
cludes that the Santa Fe formation is made up of several sedimentary horizons, deposited as broad alluvial fans
from rivers with low gradients arising in the relatively low ancestral Sangre de Cristo Mountains and associated
highlands. He thinks that during the time these sediments were being deposited the climate was semi-arid to
humid and that there were occasional showers of volcanic ash from nearby cones. There was a considerable
amount of vegetation on the floodplain and along the bordering highlands.

It would be most desirable to have a supplementary study of the Santa Fe formation, based largely upon the
contained vertebrate faunas, to determine as nearly as possible the zoologic and stratigraphic relationships of
these mammalian assemblages.

Proboscideans:
Serridentinus produdus (Cope) Trobelodon taoensis Frick
Trilophodon pojoaquensis Frick Trilophodon {Tatahelodon) riograndensis (Frick)
Megabelodon cruziensis (Frick) Trilophodon joraki (Frick)
Ocalientinus ojocaliensis Frick

Deep River Beds, Montana. — Some confusion exists as to the name of this formation, for the term was
originally applied by Scott to the sediments along the Smith River, which was once Deep Creek. As called

originally described, two faunas were recognized from this series of deposits: a lower one of Upper Oligocene age,
and an upper one of approximately Middle Miocene age. Consequently, Douglass in 1903* limited the name Deep
River to the upper fauna, and applied a new name, Fort Logan, to the lower mammalian assemblage.

According to Douglass, the Deep River beds, in the strict sense of the word, consist of marl, volcanic dust,
soft limestone, and fine sand, and they attain a thickness of approximately 400 feet.

'Frick, Childs, 1933. Bull. Amer. Mus. Nat. Hist., LIX, Art. IX, pp. 549, 571.

^Frick, Childs, 1937. Bull. Amer. Mus. Nat. Hist., LXIX, p. 7.

'Denny, Charles S., 1940. Bull. Geol. Soc. Amer., LI, No. 5, pp. 677-693.
^Douglass, Earl, 1903. Ann. Carnegie Mus., II, No. 2, p. 150.
1494 OSBORN: THE PROBOSCIDEA

The fauna is of Micklle or Upper Miocene affinities, and may be correlated with the faunas of the Pawnee,
Virgin Valley, and Mascall formations. A new, detailed study of this formation and its contained fauna is now
being made by H. A. Koerner, and when published will offer needed and valuable information on a long neglected
subject.

Proboscidean: Rhynchotherium brevidens (Cope).

Virgin Valley Beds, Nevada. —The Virgin Valley beds, which were thoroughly studied and described by
Merriam (1910, 1911)' are located in the northwestern corner of Nevada in a high, intermontane basin. These
sediments reach a thickness of about 1500 feet, and according to Merriam consist of three zones, the lower two of
which constitute the Lower Virgin Valley and are separated from the upper zone, designated as the Upi)er Virgin
Valley, by an unconformity. Most of the fossils come from a comparatively thin zone situated near the middle of
the section.

An extensive fauna is known from the Virgin Valley beds, which is definitely of Middle or Upper Miocene
relationships.

Tho closest relationship of the Virgin Valley fauna .seems to be with that of the Mascall Beds of Oregon and of the Pawnee
Creek Beds of Colorado. The Snake Creek Beds of Nebraska contain a larger percentage of the Virgin Valley species than
either the Mascall or Pawnee Creek, but there seems, nevertheless, good reason for considering the relationship with the other
faunas a.s closer.

Proboscidean :

Miomastodon merriami Osborn

"Mastodon {Tetmbelodon ?, sp.)"

Barstow Formation, Mohave Desert, California. — One of the characteristic Upper Miocene faunas of
North America is that of the Barstow formation, named and by Merriam. The Barstow beds are expos-
described
ed in southern California, in the southwestern .section of the Great Basin, and they form a part of the hot, dry
Mohave Desert. According to Merriam five members (as defined by Baker) constitute the Barstow formation.
These begin with a basal breccia at the bottom, and grade upward through a tuff-breccia, a fine ashy and shaly
tuff, a re.sistant breccia, to a fossiUferous tuff constituting tho topmost bed, and in which the vertebrate remains
are found.

The fauna of the Barstow formation is exten.sive and closely comparable to the Santa Fe "fauna," as originally
known. Thus the evidence would seem to be conclusively in favor of an uppermost Miocene age for the Barstow,
slightly later than the Virgin Valley, Mascall, Pawnee, and Lower Snake Creek beds, but earlier than the Ricardo,
which also is exposed in the Mohave Desert.

Merriam- has made the following remarks with regard to tho Barstow formation, and its fainia:

The fauna of the Barstow beds is as a whole that of an open country affording fairly abundant grass and herbage, and evi-
dently better watered than the Mohave Desert at the present day. The abundance of remains of grazing horses of the Mery-
chippus type, the pre.sence of mastodons, oreodonts, abundant merycodonts, a considerable variety of camels, and a peccary all
indicate that nutritious vegetation must have been more abundant than at present. The Mcri/rhippiis forms would i)robably not
have been present in such numbers unless gra.s.ses were well represented. . . .

The fauna of the liarstow beds represents a stage in the evolution of Tertiary mammalian faunas not jjreviously distinctly
recognized in the (Ireat F^asin Province. It seems clearly later than the Middle .\Iiocenc stage of the Mascall and Virgin \'allcy;
and is markedly older than Rattlesnake, Thousand Creek, and Ricardo, representing the next known stage following the
'Merriam, .I.e., 1911. Bull. Dopt. Geol., Univ. Calif., VI, No. 11, p. 206.
^Merriam, .1. C, 1919. Bull. Dept. Geol., Univ. Calif., XI, No. .3, pp. 450, 4ol, 453.
 GEOLOGIC SUCCESSION: NORTH AMERICA 1495

Middle Miocene in the Great Basin. The fauna of the Barstow has few if any species in common with that of the Ricardo, and

is of a distinctly older type. Its nearest relationships are with the fauna! assemblage of the Cedar Mountain region of south-
western Nevada, from which it possibly differs somewhat in stage. . . .

relationships of the Barstow fauna outside the Great Basin are with the Santa Fe beds of New Mexico. Several
The nearest
types which are among the most important forms of the Santa Fe beds are similar to species in the Barstow fauna. These in-
clude Aelurodon wheelerianus, Merychippus calamarius, Procamelus near gracili><, and Merycodus necatus.
As a considerable distance separates the Barstow geographically from the Santa Fe some difference in fauna is to be ex-
pected. It is also possible that the Santa Fe beds represent more than one horizon, or may include beds ranging into stages
older or younger than the Barstow.

Proboscideans:
Serridentinus barstonis (Frick)
Tetrabelodon ? sp.

3. THE LOWER PLIOCENE; CLARENDONIAN

Alachua and Bone Valley Formations, Florida. — Two formations represent the Pliocene in Florida;
these are the Alachua and Bone Valley beds exposed in the northwestern and central western portions of the State

respectively. Although different from each other in their physical expression and the relations with underlying
and overlying beds, these two formations are correlative in age and therefore may be considered together.
The Alachua formation consists of clays and phosphate rocks resting on the Eocene Ocala limestone and
overlain by soft Pleistocene or Recent sands. Due to the action of ground waters there has been a great deal of
solution and slumping, before, during, and after the deposition of the Alachua deposits, so that the relationships of

the Alachua clays and their contained mammals with underlying and overlying deposits and faunas have been
much confused. Simpson has shown that the Alachua formation contains a true, pure Pliocene fauna, not to be

confused with the Eocene and Miocene vertebrates that are found in older beds below it, or with Pleistocene and
Recent species from above.

The Bone Valley formation is made up of gravels, or more properly of "pebble phosphates" which grade up-
ward into true sands. The Bone Valley everywhere rests upon the Miocene Hawthorn formation, and according
to Simpson much of the material constituting it has been derived from the underlying Hawthorn sediments.
Simpson beUeves that the Bone Valley is of estuarine origin. Although there is some false association of verte-
brates from the older deposits in the Bone Valley beds, such association is readily interpreted, and does not lead
to the confusion of various faunal elements as is the case with the Alachua formation. Fossil remains from over-
lying, younger beds also are apt to be included within the Bone Valley deposits.

It might be said at this place that the Peace Creek beds, often quoted in the older Hterature as representing in

part the PHocene of Florida, are in reality of Pleistocene age, containing an admixture of derived Pliocene species.

Various expressions of opinion have been made and the Bone Valley formations.
as to the age of the Alachua
Osborn and Matthew in 1909 considered these formations as equivalent to the "Peraceras zone" of Upper Mio-
cene or Lower PHocene affinities; Osborn in 1910 placed them in the Pliocene; Sellards in 1916 made the two for-
mations correlatively equivalent to each other and placed them in the PUocene; Kellogg in 1924 (on the basis of
the marine mammals) regarded the Bone Valley as of pre-Pliocene age, while Hay in 1923 referred the two for-

mations to the Pleistocene.

Simpson's recent work of 1930' is the most thorough that has been done on these formations. As the result

of his studies, he has come to the following conclusions.

'Simpson, G. G., 1930. Bull. Amer. Mus. Nat. Hist., LIX, pp. 149-211.
 :

1496 OSBORN: THE PROBOSCIDEA

1. The Alachua and Bone Valley formations are approximately equivalent in age, since their vertebrate
faunas are essentially similar.

The land mammals found in these formations compare most closely with the Lower Pliocene faunas of
2.

western North America, more particularly with the faunas of the Upper Snake Creek and the Repubhcan River
beds.

3. The marine mammals of the Alachua and Bone Valley formations seemingly show definite pre-Pliocene
affinities (perhaps as old as the Helvetian, according to Kellogg) . However, this discrepancy between the evidence
of the land and marine forms may be due to

a. Derivation of the marine fossils from an older formation.

b. Survival of older marine types to a later period in Florida.

c. Tentative identifications, due to faulty material, that would result in the reference of the marine mammals
to genera of older geologic age than they represent.

Stirton (1936)' places the Alachua in the Middle Phocene, as about equivalent to the Hemphill.

Proboscideans:
Serridentinus floridanus (Leidy)
Trilophodon simplicidens (Osborn)
Serridentinus breivsterensis Osborn
Pliomastodon sellardsi Simpson
Ocalientinus floridanus leidii (Frick)
Ocalientinus hijoliatus (Osborn)

Valentine Beds, North Central Nebraska.— The name "Valentine" was given by Barbour and Cook- in

1917 to a series of beds in northern Nebraska which these authors considered as representing a stratigraphic unit,
containing a single fauna. Barbour and Cook, as a result of their study of the fossils from the Valentine beds,
decided that these sediments are of Lower Pliocene age, below the Upper Snake Creek beds, as defined by Matthew.

Subsequently the name "Valentine" came into general acceptance for the lowermost Pliocene, or transitional
Miocene to Pliocene phase of the Great Plains area — not only as a formation name, but also as a time term, due
particularly to its usage by Matthew, Osborn, Simpson, and other students discussing general problems of cor-
relation.

Stirton and McGrew, in 1935'', as a result of their extensive field work in the Valentine area, divided the
Valentine into three horizons, one of Upper Miocene age, one of Lower Pliocene age, and one of an age transitional
between the Lower and Middle Pliocene. These authors restricted the name "Valentine" to the uppermost of
these horizons, applying to the lowest horizon the name "Niobrara River" and to the middle one the name
"Burge."

In an effort to modify the change in usage proposed by Stirton and McGrew, Johnson, in 1936,^ suggested

that the name "Valentine" be applied to the lowest of the three horizons in question, that a new name, "Cap
Rock beds," be applied to the highest level, while the name "Burge" be retained for the middle horizon.
'Stirton, R. .\., 193(1. Amer. Journ. Sci., (5), XXXII, pp. Kil 20<i.
-Barbour, E. H., and Harold J. Cook, 1917. Neb. Geol. Siirv., Vll, Pt. 19, p. 173.
Stirton, R. A., and P. O. McGrew, lOSf). Amer. Journ. Sci., (o), XXIX, pp. 125-132.
^Johnson, F. W., 1936. Amer. Journ. Sci., (.5), XXXI, pp. 467-475.
 GEOLOGIC SUCCESSION: NORTH AMERICA 1497

Since then the relationships and nomenclature of the various horizons comprising the Upper Miocene and
Lower Pliocene of northern Nebraska have been extensively discussed by several authors. Suffice it to say that at

present several shades of opinion exist, so that the "Valentine Problem" has become rather complex and difficult
to follow. There is a recent tendency among those interested in this problem, to regard the name "Valentine"
as a rather inclusive term, designating the uppermost Miocene and the lowermost Pliocene in the north central

Nebraska region. Such a view has been expressed by Lugn,' in a paper reviewing and redefining the entire
Tertiary sequence in Nebraska. Lugn defines a large stratigraphic group of Pliocene age, which he calls the
Ogallala —using a name that had long been more or less abandoned. In the lower portion of this group he includes
the Valentine formation consisting of the original Valentine fauna, or Niobrara River assemblage at the base,
contained in loose, white sands, with the Burge channel member above it. Above the Valentine is the Ash
Hollow formation, the lower portion of which is formed by the hard cap rock, containing the "Valentine" fauna of
Stirton and McGrew. Incidentally, this cap rock layer with its included fauna was renamed the "Minnechaduza"
by Stirton,- in a paper that came out almost simultaneously with the above mentioned contribution by Lugn.

Several authors, notably Cook and Cook, Stirton and McGrew, Johnson, and McGrew, have given faunal
lists in which the mammalian assemblages from the three zones or levels of the Valentine and lower Ash Hollow
formations are defined. There is much disagreement among these authors as to the position of many genera and
species in the three horizons, and due to the lack, of knowledge as to the exact level at which most of the earlier

collections were made, some well-known forms cannot be definitely placed. The important point brought out by
the work of Stirton and McGrew is that in the lowest horizon Merychippus is the characteristic horse, while mem-
bers of the Hipparion group are absent. This is the definitive argument for placing this fauna in the Upper
Miocene. The Hipparion types, Nannippus particularly, appear first in the middle horizon, and this constitutes

the most telling argument for considering this zone as of basal Pliocene age. Finally, in the upper horizon more
advanced types of Hipparion appear, which would seem to connect this fauna with the progressive Lower
Pliocene faunas of other parts of North America.

Stirton and McGrew

1935
Upper Horizon
1498 OSBORN: THE PROBOSCIDEA

Proboscideans:
Trilophodon abeli (Barbour) Devil's Gulch
" "
Trilophodon wiUistoni (Barbour)
" "
TrilopJwdon pMp-psi Cook
" "
Trilophodon {Talahelodon) gregorii (Frick)
Trilophodon giganteus Osborn Oak Creek
Trilophodon (Genomastodon) osborni Barbour Bristow
Serbelodon barbourensis Frick Devil's Gulch
" "
Eubelodon morriUi Barbour
Megabelodon lulli Barbour Valentine
"
Torynobelodon barnumbrowni Barboiu-

Devil's Gulch, North Central Nebraska. — The Devil's Gulch beds were named by Barbour in 1914,'

for the Pliocene sediments as typically exposed in Devil's Gulch, north of Ainsworth, Nebraska. In his original
description of this fomiation, Barbour considered it as of Pliocene age.

As to the geological horizon, the faunal evidence suggests Pliocene equivalent to the Snake Creek beds of southern Sioux
County. Possibly some Pleistocene may be represented. Further study will be necessary to determine accurately the geologic
position of this newly explored fossil field. Faunal comparisons show this bed to be much earlier than that of Hay Springs, and
later than that of the famous Agate Springs Quarries. It will not be far wrong to call the Devil's Gulch deposits. Pliocene.

Barbour, and Barbour and Cook, in the early work on the Devil's Gulch beds, definitely considered these sedi-
ments as being rather well uj) in the Pliocene, certainly later than the Valentine and probably later than the Upper
Snake Creek.

In 1918 Osborn- listed two levels for the Devil's Gulch, a lower one more or less equivalent to the Valentine,
and an upper one of much more advanced, post-Snake Creek affinities.

Recent field work (as yet unpublished) in the Ainsworth regions would seem to indicate that the Devil's
Gulch beds probably are correlative with the Valentine formation to the west, and cover a fairly long period of

time, ranging from late Miocene well into the Pliocene. It is very possible that there are several faunal zones
here, corresponding to the zones of the Valentine area, as distinguished by Stirton and McGrew.
Proboscideans: See preceding list.

Oak Creek Formation, South Dakota. — The Oak Creek beds were named by Troxell, in 1916,'* who de-
scribed a new equine, Pliohippus lullianus, as coming from this horizon. Osborn, in 1918, placed the Oak Creek
in the Lower Phocene as an approximate equivalent of the Valentine beds, as then known.

In discussing Trilophodon giganteus, in Volume I of the present Monograph, Osborn'' says:

Whereas on the lower slopes of the Keyapaha there occur true Miocene beds, near the tops of the hills along the river
Keyapaha, the upi)er zone on the north side of the river (near Dallas) corresponds to the Pliocene quarry near Springview, and
to the beds near by in which was found the tyjie of Peraceraa troxelli. This whole formation seems to be as recent as the Oak
Creek beds in which was found the type of Pliohipputs lullianus; and all indicate an early Pliocene age.

The Oak Creek beds are undoubtedly the same as certain phases of the Devil's Gulch deposits to the south,
and of the Valentine beds to the southwest. The strata in which Trilophodon giganteus was found are i)robably

'Barbour, E. H., 1914. Neb. Geol. Surv., IV, Pt. XI, p. 183. (Reprint from the University Studies.)
^Osborn, H. F., 1918. Equidte of the Oligocene, Miocene, ami Pliocene of North America, Mem. Amer. Miis. N.at. Hist., N. S., II, Pt. I, p. 29.

^Troxell, E. L., 1916. Araer. Journ. Sci., (4), XLII, pp. ;j3r) 348.
••Osborn, H. F., 1936. Vol. I of this Monograph, p. 30.').
 GEOLOGIC SUCCESSION: NORTH AMERICA 1499

correlative with the middle horizon of the Valentine beds, the Burge of Stirton and McGrew, in which Megabelodon
lulli was discovered. Certainly, in view of the very close relationship that undoubtedly exists between the two
above species, the contemporaneity of the beds in which they were found seems to be well established.

Proboscideans: See preceding list.

Upper Snake Creek Beds, Northwestern Nebraska. — The Snake Creek beds were named by Matthew
and Cook in 1909,^ who at the time considered them as constituting a stratigraphic unit containing a single fauna.
This fauna was regarded as of Lower Pliocene age, but it was noted that there were Miocene species contained in
the assemblage and these were thought to be persistent primitive forms, contemporaneous with the Lower
Pliocene mammals. Merriam suggested that the seeming mixture of Miocene and Phocene mammals in a single
fauna might, in fact, be due to the stratigraphic mixing of two faunas. This same view was expressed by Matthew
in 1918, but it was not until 1924" that he was able to define the stratigraphic horizons in the Snake Creek area.

Matthew has shown quite clearly that there are three principal horizons and faunas in the Snake Creek
region. These are:

Upper Snake Creek beds Lower Pliocene Hipparion affine

Lower Snake Creek beds Upper Miocene Merychippus paniensis

Sheep Creek beds Middle Miocene Merychippus primus

In addition Matthew distinguished a fourth zone, which he called the Pliohippus leidyanus zone. This
horizon is later than the Upper Snake Creek in age, but since it has a very fragmentary fauna it has not received
a definite geographic name.

Proboscideans have been described only from the Upper Snake Creek beds, so this horizon in the Sioux
County series will be the only one considered at this place. According to Matthew:

The principal upper Sjiake Creek fauna is clearly distinct and of Pliocene age, comparable with that of the Republican River
beds. There are some forms in it suggesting a later stage, but they are rare and imperfectly known, and their evidence is not
weighty. The correspondence, however, is by no means so close as that between Pawnee Creek and lower Snake Creek. There
are few species in common. Teleoceras fossiger, so abundant in Republican River, is not positively recorded from the Snake
Creek. The Equidae correspond fairly well as to genera, but not as to species. ^Elurodon is the characteristic canid in both
faunas.

Subsequently, in 1930,^ Matthew suggested that the Upper Snake Creek fauna is nearly equivalent to the
Hemphill fauna : "The Upper Snake Creek fauna is also rather nearly equivalent [to the Hemphill] including , some
species more advanced, others somewhat less so."

Generally speaking, the Upper Snake Creek may be regarded as representative of the upper portion of the
Lower Pliocene in North America, probably approaching related Middle Pliocene deposits and faunas in its broad
relationships.

There has been some tendency among recent students to assign the Upper Snake Creek to the Middle or
Upper Pliocene (Lugn, 1939) or to spread it over a large extent of the Pliocene (Stirton, 1936), this latter correla-
tion being made upon the basis of an apparent mixing of faunas in this horizon.

'Matthew, W. D., and Harold J. Cook, 1909. Bull. Amer. Mus. Nat. Hist., XXVI, Art. XXVII, p. 363.

^Matthew, W. D., 1924. Bull. Amer. Mas. Nat. Hist., L, pp. 72, 73.

'Matthew, W. D., and R. A. Stirton, 1930. Univ. Calif. Publ., Bull. Dept. Geol. Sci., XIX, No. 17, p. 367.
1500 OSBORN: THE PROBOSCIDEA

Proboscideans:
PUomastodon matthemi Osborn
Rhynchotherium anguirivale Osborn
Serrideniinus anguirivalis Osborn
Serridentinus nehrascensis Osborn

Republican River Formation, Southern Nebraska and Northern Kansas. — Simpson,' in 1933, made the
following remarks about the Republican River beds.

Lower Pliocene of northwestern Kansas, and, by extension, the corresponding stage of the Pliocene generally. The beds
were extensively explored, especially for Marsh, and a large fauna is known, but there is no recent faunal or stratigraphic revision
aside from such syntheses as that of Osborn and Matthew (1909) or references in describing individual fossils or faunas of similar
age. It was generally placed in the 'Loup Fork,' 'Upper Loup Fork,' or 'Ogalalla,' but the more definite local name
has been
consistently applied by Matthew and is now generally accepted. Matthew (in Osborn and Matthew, 1909) tentatively con-
sidered it as .slightly later than the 'Nebraska' (Valentine) later reaffirming this more positively (1924) Mo.st recently, Matthew
, .

and Stirton (1930) have, in passing, suggested that the Republican River may be in part composite, with the beds on Sappa
Creek and elsewhere to the southwest later than those on Driftwood Creek and to the north. They continue to place the fauna,
as a whole, as younger than the Valentine, adding that it seems to be .slightly older than their Goodnight-Hemphill (and hence
equivalent to Clarendon).

Lugn,2 in his review of the Tertiary of Nebraska, makes the following remarks about the Republican River
beds:

Considering all available evidence, it seems certain that the 'Republican River' represents a composite of lithologic and
faunal horizons and .should be abandoned as a stratigraphic term. Furthermore, many of the fossil finds were obtained from the
unconsolidated sands and gravels in pits under the lower slopes of the valleys west and southwest of McCook, Nebraska. Some
fossils have been collected from higher beds, from the 'mortar beds' along the sides of the valleys. The sand
and gravels belong
to the upper part of the Valentine formation, and the 'mortar beds' levels contain the Krynitzkia fossil seed zone and the lower-
most part of the Biorbia fossil seed zone, all in the lower part of the Ash Hollow formation. Therefore, the fauna is much
mixed, and also the described and published sections do not include the lower (Valentine) beds in the exposures.

Proboscideans:
Blickoiherium euhypodon (Cope)
Tetralophodon campester (Cope)
Serridentinus progressus Osborn
Trilophodon dinotherioides Andrews
Ocalientinus republicanus (Osborn)

Wray Beds, Northeastern Colorado. —A considerable fauna from the Pliocene of northeastern Colorado,
near Wray, was described by Cook' in 1922. The fossils came from a relatively thin deposit of stream channel
sands and gravels, resting directly on the Cretaceous Pierre shales. Cook regarded the Wray beds, as he desig-
nated them, as being very closely related to the Upper Snake Creek of northwestern Nebraska.

This fauna from Yuma County, Colorado, which we may designate for local convenience, the 'Wray' beds, is a very close
equivalent of the upper phases of the Snake Creek beds, but shows certain faunal differences. This may be partly due to the
fact that we do not know both faunas completely and have found different representatives from each.
Some of the species. . .

from these beds are closely related to forms described by Merriam from California and Nevada, from beds slightly older or young-
er or of equivalent age.

Stirton (1936) regards the Wray as being somewhat later in age than it was originally designated by Cook,
placing it in the Middle PHocene as about an equivalent of the Hemphill.

'Simpson, G. G., 1933. Bull. Amer. Mus. Nat. Hist., LXVII, p. 107.

^Lugn, A. L., 1939. Bull. Geol. Soc. Amer., L, pp. 1272, 1273.

'Cook, H. J., 1922. Proc. Colorado Mus. Nat. Hist., IV, No. 2, p. 4.
 GEOLOGIC SUCCESSION: NORTH AMERICA 1501

Proboscideans:
Amebelodon paladentatus (Cook)
Amebelodon hicksi (Cook)

—
Santa Fe Beds, New Mexico. As has been shown above (p. 1492), the Santa Fe beds seemingly contain
more than one horizon and cover a time range including the Upper Miocene and the Lower Pliocene. Prick's
work of the past few years in the Santa Fe region has given evidence of a definite Pliocene Hipparion level within
this series of sediments.

Loup Fork and Ogalalla. — "Loup Fork" is an old term in the literature dealing with the continental
Tertiary of Nortli America, established through the early writings of Leidy and Cope. From the beginning, its

meaning has been obscure and uncertain, so that it has no value as a stratigraphic name —and at the present time
is for the most part ignored. Applied to Upper Tertiary beds in Nebraska of Miocene and Pliocene age, it was
frequently confused with the "Loup River" of Meek and Hayden —a term applied to beds, mainly of Pleistocene
age, along the Loup River
in central Nebraska. Fossils described by the earlier writers as of "Loup Fork" age
may be from almost any horizon between the Middle Miocene and the Upper Pliocene.
The name "Ogallala" was used by Darton in 1905' to designate the Upper Tertiary of Nebraska above the
"Arikaree," and as such it included the Upper Miocene, Lower and Middle Pliocene of the Great Plains region.
At the present time the Nebraska Geological Survey regards the Ogallala as a valid group name, containing
Pliocene formations in Nebraska.

Clarendon Beds, Northwestern Texas.— The Clarendon beds were named by Gidley- in 1903 for Upper
Tertiary deposits to the north of Clarendon, Texas, which he considered as being correlative with the "Loup Fork"
of the Great Plains. At the time these sediments were considered as coming within the Miocene.
The main body of the beds consists for the most part of cross-bedded sands and sandstones intermixing more or less and
cross-bedding with the clays. These channels all take a chrection nearly east and west, or approximately the same as that of the
streams draining the country at the present time. Some of them are traceable for long distances. It is in these peculiar beds of
sandy clays that all the fossils of this region occur.

According to Gidley, there are about 400 feet of thickness represented in the Clarendon horizon at its typical
outcrops, and the fossiliferous stratum lies at the top of the section. Gidley included the Goodnight beds, as
named by Cmnmins,^ within show why these Goodnight sediments
the Clarendon, advancing a long argument to
are not separable. Subsequent data show that the Goodnight beds are distinct from and later than the Clarendon.
The Clarendon fauna is now generally recognized as representing a distinct phase in the Phocene history of
North America, older than the Hemphill-Goodnight and about equivalent to the Lower PUocene of the Great
Plains area.

"The typical Clarendon species are nearly allied to those from Hemphill, although more primitive, and they
may well have been comparatively direct ancestral stages or mutations." Matthew, W. D., and R. A. Stirton,
(op. dt., 1930, p. 386).

Proboscideans:
Serridentinus productus (Cope)
Serridentinus serridens (Cope)
Serridentinus serridens cimarronis (Cope)
Serbelodon praecursor (Cope)
Tetralophodon fricki Osborn
'Darton, N. H., 1905. U. S. Geol. Surv. Profess. Paper No. 32, p. 178.
^Gidley, J. W., 1903. Bull. Amer. Mus. Nat. Hist., XIX, p. 633.
'Cummins, W. F., 1893. Texas Geol. Surv., Fourth Annual Report, Pt. I, pp. 201, 203.
1502 OSBORN: THE PROBOSCIDEA

RiCARDO Formation, Mohave Desert, California. —The Ricardo beds are located in the southwestern
corner of the Great Basin area, not far from the region where the tyjiical Barstow Miocene sediments are exposed.
Merriam, who studied l)oth tlie Ricardo and the Barstow faunas, early recognizeil the fact that the former as-
semblage is of later age than the latter because of an almost complete specific separation between them and conse-
quently he considered the relationships of the more advanced Ricardo fauna to be with certain Pliocene deposits
of the The Ricardo, typified by Hipparion and Pliohippus, is definitely of
Great Basin and the Plains regions.
Lower Pliocene age, and is probably older than the Thousand Creek or the Rattlesnake deposits of the Great Basin
region, since the Ricardo species, particularly among the Equidse, are slightly more primitive than the forms from
the Rattlesnake and Thousand Creek.

In its general age relationships the Ricardo, therefore, would seem to be more or less comparable with the
Valentine and Clarendon of the Great Plains region.
Proboscideans:
Trilophodon sp.
Serhelodon burnhami Osborn

4. THE MIDDLE PLIOCENE: HEMPHILLIAN

Thousand Creek Beds, Northwestern Nevada. —The Thousand Creek beds, named from Thousand
Creek in the northwestern corner of Nevada, consist of an alternation of tuffs, ashes, sands, gravels, and ancient

soil accumulations. From these beds a distinctive Pliocene fauna has been described by Merriam.

As Merriam has shown, the Thousand Creek fauna contains advanced Tertiary carnivores, rodents, horses,
camelids, and antilocaprids, yet in spite of the numerous progressive elements in the fauna, it must be regarded
as a purely PUocene assemblage. That is, none of the Thousand Creek mammals would warrant the inclusion of
the beds within the Pleistocene. It might be mentioned, however, that Pleistocene terraces containing mammalian
remains are found contiguous to the Thousand Creek exposures, and due to the difficulty of separating the sedi-
ments of the two epochs there is a possibiUty of mixing the faunal elements in making collections in this area.

The Thousand Creek beds are definitely much later than the Virgin Valley beds, which are exposed nearby,
and are more nearly comparable to the Rattlesnake, and especially to the Upper Snake Creek of Nebraska.
Merriam' made the following remarks with regard to the Thousand Creek beds:

In so far as correlation with the American mammaHaii faunas is concerned the Thousand Creek fauna would seem neces-
sarily to take a place later than that of the Snake Creek [Upper Snake Creek] and earlier than that of the Blanco.

In 1930 Matthew and Stirton suggested that the Thousand Creek fauna might be more or less equivalent
to the Hemphill fauna.

Proboscideans:
"Tetrabelodon ? sp."
Miomastodon merriami Osborn
Pliomastodon nevadanus Stock

Mount Eden Beds, Southern California.— The Eden beds were named by Frick" in 1921 to distinguish
a series of Lower Pliocene sediments exposed in the San Timoteo badlands in southern Cahfornia. In his original
'Merriam, J. C, 1911. Univ. Calif. PubL, Bull. Dept. Geol. Sci., VI, No. 11, p. 217.
''Frick, Childs, 1921. Univ. Calif. Publ., Bull. Dept. Geol. Sci., Xll, pp. 283-288.
 : .

GEOLOGIC SUCCESSION: NORTH AMERICA 1503

publication Frick placed the Eden beds in the middle portion of the Pliocene, as correlative with the Middle
Etchegoin, Rattlesnake, and Snake Creek (Upper), but subsequently (1933) he stated that the Eden beds are of
uppermost Pliocene age.

That the Eden beds are fairly well advanced in the Pliocene is made evident by certain progressive forms
present in the faunal assemblage. These are particularly, two sloths, Nothrotherium or Pronoihrotherium and
Megalomjx, a sabre-toothed cat referred to Smilodon, a bear, Hysenarctos, and the advanced peccary, Platy-
gonus. On the other hand there are many typical Lower Pliocene forms in the Eden fauna, such as Pliohippus
among the horses, Prosthennops, Pliauchenia and Procamelus, Merycodus, and Trilophodon. Frick has called
attention to the absence of Hipparion in the Eden fauna, which might argue for a relatively advanced age for the

assemblage. Likewise, rhinoceroses are absent in this faunal assemblage as they are in post-mid-Pliocene deposits
of North America, but the same is true of the Ricardo fauna, so that this hne of evidence must not be accorded
too much importance in the consideration of age relationships.

All in all, it would seem probable that the Eden fauna is a moderately advanced Phocene assemblage, con-
taining many progressive types of mammals that foreshadow the coming of the Pleistocene in North America,
but the absence of truly advanced horses such as those characterizing the Hagerman of Idaho, would seem to
preclude the assignment of the Eden to the uppermost Pliocene.

The Eden, as originally named by Frick, has been changed to "Mount Eden" by Eraser,' the name "Eden"
being preoccupied by a Paleozoic formation.

Proboscideans:
Rhynchotherium shepardi edense (Frick)
Cordillerion edensis Osborn

Etchegoin, Southern California. — Merriam recognized three horizons or zones in the Etchegoin area,
on the western border of the San Joaquin Valley, which he named as follows
Pliohippus proversus zone
Pliohippus coalingensis zone
IHipparion zone.

Generally speaking these three zones cover a period of time extending from the upper part of the Lower
Pliocene, comparable to the Rattlesnake or Upper Snake Creek, to the Upper Pliocene, comparable to the Blanco.
Fragmentary mastodont remains have been found in the upper portion of the Etchegoin

Proboscideans:
"Mastodon"
Pliomastodon vexillarius Matthew

5. THE UPPER PLIOCENE: BLANCAN

Blanco Formation, Northwestern Texas. —The Blanco beds have for many years been regarded as
representative of one of the later phases of the Pliocene in North America. These deposits were first studied and
described by Cummins and by Cope in 1893, the former author reporting on their stratigraphic relationships and
the latter describing the contained fauna. Subsequently (1903) Gidley redescribed the Blanco, limiting its hori-
zontal and vertical extent as compared with the original descriptions of Cummins and Cope.
'Fraser, D. M., 1931. Mining in Calif., XXVII, No. 4, pp. 511-514.
 —

1304 OSBORN: THE PROBOSCIDEA

According to Gidley, the Blanco beds consist of streana channel deposits —sands, clays, and diatomaceous
earth —cut into the older Miocene sediments, and exposed in a relatively narrow belt trending in a northwesterly
to southeasterly direction along either side of the Blanco River.

The Blanco fauna, though small, is well known due to the writings of Cope, Osborn, and Gidley, and therefore
has become well established as one of the most characteristic later Pliocene faunas of America. It has always
been recognized as of PUocene age, although the earlier authors did not specify its exact position within the Plio-
cene. In 1909, however, Matthew definitely placed it as of Middle Pliocene age — a correlation that has been
generally followed in subsequent publications.

Recent evidence favors the idea that the Blanco might be even later than Middle Pliocene in its age. Thus,
Gazin, who has described the uppermost PUocene Hagerman beds of Idaho, regards the fauna of these sediments
as showing many close relationships to the Blanco fauna. Illustrative of the close relationships between the
Blanco and the Hagerman, there might be mentioned the very advanced monodactyl horse Plesippus, which occurs
in both formations. Plesippus is certainly directly ancestral to Equus, and its first appearance cannot antedate
the Pliocene-Pleistocene transition by any great length of time.

Proboscideans:
Rhynchotherium falconeri Osborn
Serbelodon praecursor (Cope)
Stegomastodon mirificus (Leidy)
Stegomastodon successor (Cope)
Stegomastodon texanus Osborn

—
San Pedro Valley Beds, Arizona. On either side of the San Pedro valley, in southern Arizona, are
fossiliferous localities at which Upper Pliocene vertebrates have been found. Gidley, who has described the
mammals from the San Pedro deposits, has shown that the sediments, as interpreted by Bryan, consist of a con-
siderable thickness of sand, conglomerate, and clay valley fill, deformed to a certain extent by subsequent dias-
trophic movements.

On the west side of the valley fossils were found near the town of Benson, while to the east the fossiliferous
deposits are exposed on the Curtis Flats, near Curtis Ranch. The names Benson and Curtis Ranch have been
assigned accordingly to the faunas from the two areas.

Although the beds at the Benson and Curtis Ranch localities seem to be much the same stratigraphically, the

faunas from the two areas show decided differences. From the Benson horizon are Cordillerion bensonensis,
Neohipparion, Pliohippus, and Merycodus, while the mammalian assemblage from Curtis Ranch, containing as it
does such forms as Stegomastodon arizonx, Glyptotherium, and Plesippus, would seem to be definitely later in age
than the Benson fauna. Gidley' considered the Benson fauna to be fairly closely related to the Blanco fauna
perhaps somewhat earlier in age than the Texas assemblage, while he placed the Curtis Flats fauna in the Upper
Pliocene as definitely later than the Blanco fauna.

The bones occur for the most part in relatively small patches or layers of greenish tuffaceous clay, which, according to
Bryan, interfingcr on one side with arkosic gravel and conglomerate typical of deposition on alluvial slopes and on the other
with the lake beds. This position seems to confirm Bryan's view that these bone-bearing patches of gnjenish clay represent the
marginal and fresh-water springs that are characteristic of the borders of salt lakes in such basins. The localities thus probably
constituted the chief watering places for the animals of the region, and here, naturally, occur their fossil remains.

'Gidley, J. W., 1926. U. S. Geol. Surv., Profess. Paper 140-B, p. 84.

 GEOLOGIC SUCCESSION: NORTH AMERICA 1505

. although the general process of sedimentation in the San Pedro basin was continuous, the marginal springs of the two
. .

localities may have belonged to lakes of slightly different levels. Thus the time interval between the active existence of the
watering places on the west side of the salt lakes at the Benson locality and those on the east side of the lakes at Curtis Flats
may well have been long.
During this interval the faunas changed, but it does not necessarily follow that this change was greatly affected by evolution
in this locality, for, although considered long as measured in years, the interval was probably in a geologic sense relatively short,
and it may well be that the entire change was accomplished by migration. though many of the species of the Curtis Flats
. . .

locality are closely related to the species found in the Benson locality, the former are not descendant forms of the latter.

More recently, Dr. Gazin has expressed his opinion in conversations with the writer that the San Pedro beds
at Curtis Ranch are of Lower Pleistocene, rather than of Phocene age. He would regard the Benson deposits as
approximately equivalent in age to the Blanco of Texas, a view that is coming to be generally accepted.

Proboscideans:
Curtis Ranch Stegomastodon arizonx Gidley
Benson Cordillerion bensonensis (Gidley)

Hagerman Formation, Southwestern Idaho. —The name "Idaho" has been used as a vague term indica-
tive of a Upper Tertiary and Pleistocene age in southern Idaho. That the name is rather
series of deposits of

inclusive was recognized by Merriam in 1917, in his study of the Phocene mammalian faunas of western United
States. Certainly, the fauna, as he lists it, shows a mixture of Pliocene and Pleistocene types.

In recent years certain parts of the Idaho beds have been closely studied by Gazin,' who has shown that the
deposits from which his collections were made are definitely of very late Pliocene age. These beds have been
named by Gazin the Hagermaai, or Hagerman lake beds, and are representative of the uppermost Phocene of
North America. They are characterized by an advanced horse, Plesippus, which is certainly the direct ancestor
of the genus Equus. According to Gazin, the Hagerman beds are probably correlative with the Blanco of Texas.

Gazin states that the Hagerman lake beds, as he defines them, are probably somewhat older than other
exposures identified as the Idaho, particularly at Sinker Creek and certain other localities. Merriam mentions
that Stegomastodon mirificus, which he lists as from the Idaho formation, came from Sinker Creek. He then goes
on to say that this is a Pliocene type of proboscidean. It would seem to be adequately proven that the type of
Stegomastodon mirificus from Nebraska comes from beds definitely of Pleistocene age (see Lugn and Schultz,
1934). Therefore it is very possible that the portion of the Idaho beds from which the above species was obtained
is of Pleistocene age, and later than the Hagerman, as was surmised by Gazin.

Hagerman beds with the Blanco,

Gazin, in comparing the says that: "The Blanco shows a greater wealth of
Proboscidea but fewer members of the microfauna."

At the present time proboscideans have not been described from the Hagerman beds.

Proboscidean (Idaho): Stegomastodon mirificus (Leidy).

Tehama Formation, Sacramento Valley, California. —The Tehama formation, described by Russell and
VanderHoof ,- is a thick series of tuffaceous greenish gray to buff sandy clays, containing intercalated cross-bedded
sands and gravels. A small fauna was discovered in this formation, which would seem to be indicative of a late
Pliocene age, for it contains such advanced Tertiary genera as Hyasnognathus, Plesippus, lAntilocapra, Odocoileus,
Stegomastodon, and a megalonychid sloth.

'Gazin, C. L., 1936. Proc. U. S. Nat. Mus., LXXXIII, No. 2985, p. 287.
^VanderHoof, V. L., 1933. Amer. Journ. Sci., (5), XXV, p. 384.
1506 OSBOKN: THE PROBOSCIDEA

'riio horses are more ach'aiieed than those in the upper Etohegoiii (Plesippiis proversus) stage of evohition and there is an
influx of forms which appear to be the foreruimers of those reaching their maximum development in the lower Pleistocene.
From this faunal evidence, the Tehama formation may be regarded as being uppermost Pliocene in age.

Proboscidean: Stegomastodon cf. arizonx Gidley.

6. THE PLIOCENE OF MEXICO

The first proboscideans that reached Mexico would seem to have appeared in that region in Lower Pliocene
times. These early forms, referred by Freudenberg to two species, Trilophodon dinotherioides and Serridentinus
serridens, were seemingly contemporaneous with a fauna of general Lower or Middle Pliocene affinities.

Generally speaking, the proboscideans of Mexico range from the PUocene throughout the extent of the
Pleistocene. Whether their history, as well as that of the contemporaneous mammals, is a continuous story in
this region, or whether it is a record broken by considerable gaps (particularly in Upper Pliocene times) is a point
that cannot at the present time be definitely decided. Freudenberg' (1922) considered that there were, broadly
speaking, just two mammalian faunas in Mexico. He distinguished an earlier one of Lower Pliocene age,

which in fact may range into Middle Pliocene times, and a later one covering the extent of the Pleistocene. The
Pleistocene "fauna" Freudenberg thought might be divisible into three successive mammalian assemblages, but
the evidence for any such separation is admittedly uncertain.

Es bleibt also bei zwei Saugetier-Faunen in Mexiko, einer unterpliocanen imd einer dihivialen sensu lato. Die crstere ist
wohl einer einzigen kiirzeren Epoche zuzuweisen. Sie ist reich an Pferdeformen, die samthch auf der Hipparion-i^tuie stehen.
Dcmgegeniiber enthalten die Beckensedimente des Hochlandes nur die Gatung Equus, als Vertreter des Pferdestammes.

Lower and Middle Pliocene. —The Pliocene fauna described by Freudenberg is found in the states of

Vera Cruz and Hidalgo, in freshwater and lake deposits in the vicinity of Zacualtipan and Tehuichila. In these
sediments is a mastodont referred to Serridentinus serridens, an Hyxnardos, peccary, and Procamelus, as well as
Hipparion peninsulatum, Hipparion rectidens, and Protohippus castillai.

This mammalian assemblage is most certainly of Pliocene age, probably later than "upper Miocene or lower
Pliocene" as it by Freudenberg. The presence in it of Serridentinus serridens would point to-
was thought to be
wards an approximate correlation with the Lower Pliocene Clarendon of Texas. In this connection it is to be
noted that Freudenberg has also described material which he referred to Trilophodon dinotherioides, a Lower
Pliocene form the type of which is from the Republican River beds of Kansas.

Proboscideans:
Serridentinus serridens (Cope)
Trilophodon dinotherioides (Andrews)

Middle and Upper Pliocene. — It is very difficult to be certain as to the presence of late Pliocene deposits
in central Mexico. Two species of rhynchotheres, described by Osborn, may be indicative of sediments geo-
logically younger than those in by Freudenberg occurs. Osborn, in Volume I of this
which the fauna listed

Monograph, considered these forms as questionably of Upper Pliocene age. It is, of course, quite possible that
Rhynchotherium tlascalx and Rhynchotherium browni are of Upper Pliocene affinities; certainly they are later than
the lowest Pliocene of North America. On the other hand, there is no evidence against their contemporaneity
with the Vera Cruz-Hidalgo fauna, which is of late Lower or Middle Pliocene age.

Proboscideans:
Rhimcholherium tlascalae Osborn 1 _, . , , ... n*^- n r.i-
, , . , .
^ ,
These species may i

be from the Lower or Middle Pliocene

Rhynchothertum broivm Osborn I

'Freudenberg, W., 1922. Geol. und Pal. Abh., N. Folge, Band XIV, Heft III, p. lOt.
 GEOLOGIC SUCCESSION: NORTH AMERICA 1507

7. PROBOSCIDEANS FROM UNDETERMINED LEVELS IN THE MIOCENE AND PLIOCENE

OF NORTH AMERICA
A number of proboscideans from the Pliocene of North America have been described from horizons or levels
that are at the present time either unknown or not definitely determined. In some cases the exact age of the
types will never be known, for the species were described many years ago and precise data as to their levels or
localities are not to be had. On the other hand, certain forms are known to have been found at definite localities
and at well-determined geologic positions, but the uncertain knowledge as to the exact geological relation-
ships of the beds within which these types were found makes their stratigraphic position for the time being
somewhat uncertain. The species falling into these categories will be discussed below.

Trilophodon ligoniferus (Cope and Matthew)

This species is known from a type that was found in the Black Hills, according to Cope's original record. The
horizon and locality for this species must be regarded as uncertain, to say the least.

Tonjnohelodon loomisi Barbour

In the type description, Barbour states that this form was found in "Sand Canyon, just east of Indian Hill,

two and one-half miles southwest of Republican City, Harlan County, Nebraska, the formation being Late
Pliocene to Early Pleistocene."

Amebelodon fricki Barbour

Amebelodon fricki was discovered near Freedom, Frontier County, Nebraska, in the southwestern portion of
the state, some miles to the north of the Republican River. Barbour regarded its age as "late Pliocene or possibly
early Pleistocene," on the basis of his observations of the type locality.

Amebelodon sinclairi Barbour

The type of this specieswas found only a few hundred yards distant from the place where Amebelodon fricki
was discovered. Therefore it is very probable that it is contemporaneous with the above species in geologic age.

Gnathabelodon thorpei Barbour and Sternberg

According to Barbour,' "Professor Maxim K. Elias, of the University of Kansas, who has made special studies
of Trego County, when consulted, pronounced this bed Late Pliocene in all probability, although he had not seen
a section at this particular gravel pit, and based his opinion on neighboring exposures."

In a recent communication. Dr. C. Bertrand Schultz of the University of Nebraska tells me that the above
four species "all come from deposits which appear to be late Pliocene in age. Perhaps it would be better to say
'middle to late Pliocene.' They do come from the upper part of the Ogallala."

Rhynchotherium shepardi (Leidy)

The horizon of the type of this species cannot be positively determined on the basis of evidence now at hand.
The opinions of Stock and Buwalda with regard to this question are quoted on page 487 of Volume I of this Mono-
graph. It might be pointed out that a subspecies described by Frick, namely, Rhynchotherium shepardi edense,
comes from the Mount Eden formation of about Middle Pliocene age.
'Barbour, E. H., and George F. Sternberg, 1935. Bull. Neb. State Mus., I, Bull. 42, p. 396.
1508 OSBORN: THE PROBOSCIDEA

Cordillerion orarius (Hay)

Cordillerion defloccatus (Hay)
These species were described from teeth and jaws found on the west bank of the Aransas River, near Sin ton,
Texas. Hay, who described them, evidently regarded the age as Lower Pleistocene. In Volume I of this Mono-
graph Professor Osborn has assigned them provisionally to the Pliocene.

Stegomastodon priestleyi (Hay and Cook)

Although found in a deposit of Pleistocene gravels, the type of this species is possibly a derived fossil, of

Upper Pliocene age.

Trilophodon obscurus (Leidy)

The type of Trilophodon obscurus was found at Greensboro, Caroline County, Maryland, which would mean
that the probabilities are that it was found in the Choptank formation of Miocene age. The marine invertebrate
fauna would seem to indicate that the Choptank is of approximately Middle Miocene age. (See Volimie I of this
Monograph, page 285.)

Ocalientinus obliquidens (Osborn)

The phosphate beds, near Charleston, South Carolina, are definitely of Pleistocene age, as is shown by the
presence in them of numerous teeth referable to Parelephas columbi. It is notable, however, that the type tooth
of Ocalientinus obliquidens is structurally a Pliocene type of mastodont. Since redeposition is common in the

phosphate beds, and since remanie fossils of an earlier age are often found in younger sediments, the probabilities
are strongly in favor of the species under consideration being of Upper Pliocene age.

Ocalientinus emmonsi (Hay)

It is very likely that the age of this form is similar to that of Ocalientinus obliquidens, discussed above. That
is, it is an Upper Tertiary specimen redeposited in a Pleistocene sediment. Hay regarded this species as question-
ably of Pleistocene age.

8. NORTH AMERICAN TERTIARY HORIZONS CONTAINING FRAGMENTARY

PROBOSCIDEAN REMAINS
A number of important mammal-bearing horizons Upper Tertiary age have not been included in the fore-
of

going discussion, since they contain but fragmentary remains of proboscideans which have not been specifically,
and, for the most part, not even generically identified. Considering, however, the fact that there are indications of
proboscideans in these horizons —even though the evidence is largely incomplete — it may be well to discuss very

briefly these mammal-bearing sediments.

Wood Mountain Gravels, Southern Saskatchewan.— Described by Sternberg,' who records a very scanty
and fragmentary fauna, including an indeterminate mastodont. The age is either Middle or Upper Miocene.

Madison Valley Beds, Montana.— Originally described by Douglass, as the "Loup Fork of Madison Valley,
Montana." The fauna indicates a close relationship in age to the Barstow and to the lower portions of the
—
Santa Fe hence of Upper Miocene affinities. Proboscidean: a "mastodon."
'Sternberg, C. M., 1930. Trans. Roy. Soc. Canada, (3), XXIV, Sec. 4, pp. 29 and 30.
 GEOLOGIC SUCCESSION: NORTH AMERICA 1509

Skull Springs Beds, Southeastern Oregon. —Gazin, who described the fauna from this horizon, compared
it with the faunas of Virgin Valley, Mascall, Payette, Lower Snake Creek, and Pawnee. The evidence therefore
indicates an Upper Miocene age. A mastodont is known from this series.
Cedar Mountain Beds, Western Nevada. —Described by Merriam and compared with the Barstow and
Santa Fe. Stirton divided the Cedar Mountain into two horizons, one of Middle Miocene, the other of Lower
Pliocene age. "Tetrabelodon" is known from this series.

CuYAMA Formation, Southern California. — This horizon with its contained fauna has been carefully
studied by Gazin, who regards it as related in a general way to the Mint Canyon and to the Barstow beds, especial-
ly the latter. Gazin lists "mastodont sp." as from the Cuyama, now designated as the Quatal Canyon.

North Coalinga Beds, California. —One of a series of sediments north of Los Angeles. Considered by
Merriam, on the basis of a scanty fauna, to be related to the Mascall and the Virgin Valley; consequently of
Middle or Upper Miocene age. "Tetrabelodon" (?)sp. is recorded.

Mint Canyon Beds, California. —An important mammal-bearing horizon, described by Maxson in 1930.

The age of the Mint Canyon has been variously debated. Considered by Maxson as of Upper Miocene age
perhaps later than the Barstow. Stirton, on the other hand, regards it as of Lower Pliocene age, closely related to
the Ricardo. Perhaps this is a "boundary-line" horizon, representing the transition from the uppermost Miocene
into the lowermost Pliocene. Trilophodon sp. is known from this horizon.

Truckee Beds, Western Nevada. —The back portion of a third molar from these sediments was described
by Buwalda as Tetrabelodon (?), and on the basis of this specimen the beds were placed in the Upper Tertiary.
The tooth appears to be referable to the genus Miomastodon; the age is probably Upper Miocene.

—
Hemphill Beds, Texas. An important horizon, containing a fauna of about Middle Pliocene age. Matthew
and Stirton, who studied the Hemphill fauna, decided that it is somewhat later in age than the Clarendon but
earlier than the Blanco. It is equivalent to the Goodnight assemblage described many years ago by Cope. The

only known proboscidean is a fragment questionably referred to Rhynchotherium.

Optima Formation, Oklahoma. —Described by Hesse, in 1936.' This author regards it as closely related to
the Hemphill of Texas, both as to its fauna and as to its age. "A lower molar of a Proboscidean, the only indi-
cation of this family at Optima, was determined by the late W. D. Matthew as Miomastodon."

Rattlesnake Formation, Eastern Oregon. —Described in detail by Merriam, Stock, and Moody, in 1925.^
These authors regard it as later than the Ricardo, and rather closely related to the Thousand Creek. Therefore
of Middle Pliocene age. "Proboscidean remains" are recorded from the Rattlesnake.

Chanac Formation, Southern California. —Described by Merriam as closely related to the Ricardo and
to the Middle Etchegoin of the Pacific Coast area. Tetrabelodon (?) sp. is known from this horizon.

Orindan and Siestan Formations, Berkeley Hills; Pinole Tuff, San Pablo Bay, California. These —
formations, in the San Francisco area, are closely related to each other. The scanty fossils may be compared with
those of the Ricardo. Tetrabelodon (?) sp. has been recorded from the Orindan.

The formations discussed in the preceding pages of this section have been listed in detail by Wilmarth
(1938),'' to which the reader is referred.

'Hesse, C.J., 1936. Univ. Calif. Publ., Bull. Dept. Geol. Sei., XXIV, No. 3, p. 66.
-Merriam, John C, Chester Stock, and C. L. Moody, 1925. Contributions to Palaeontology, III, Carnegie Inst. Washington, No. 347, pp. 43-92.
'Wilmarth, M. Grace, 1938. U. S. Geol. Surv. Bull. 896, Pts. 1 and 2.
1510 OSBORN: THE PROBOSCIDEA

9. THE PLEISTOCENE OF NORTH AMERICA

As in Eurasia, the Pleistocene in North America may be regarded
commenced with the extension of
as having

the first continental glacier from the North, and with the appearance and spread of certain new and advanced
types of mamals in all parts of the continent. Those mammals particularly diagnostic of the beginning of the

Pleistocene in North America are the modern horse, Equus, the mammoth, Archidiskodon, and cattle. Bison. To
these three types there may be added the modern camelids, as exemplified in the New World by Cnmelops. Of the
foregoing eniunerated forms, the first and last were autochthonous and their first appearance marks the actual
beginning of Pleistocene times in the North American region. The other two forms, being immigrants from Asia,
may not have arrived in North America until after the opening of the Pleistocene, but if such is the case, the time

lag was not great.

An important clue to the changes that were taking place with the evolution of an Upjjer Pliocene fauna into

one of Pleistocene aspects is to be seen in the evidence of the horse, Plesippus, characteristic of the uppermost
Tertiary horizons of North America. This equid is in most of its characters much closer to the true Equus of the

Pleistocene than it is to any other Pliocene horses, and for this reason it may be regarded as immediately ancestral
to the horses of the Pleistocene. Thus we see in the appearance of Plesippus a near approach to the Pleistocene

similar and parallel to that signalized in the Old World by the appearance of primitive, ancestral types of Archi-
diskodon in horizons of uppermost Pliocene age.

In North America, as in Europe and Asia, the Pleistocene may be divided according to the advance and re-

treat of continental glaciers. In Europe there is a difference of opinion as to the presence of three or four glacial
advances during Pleistocene times, according to the manner in which the Pliocene-Pleistocene boundary is limited.
In America, on the other hand, the argument is concerned with the presence of four or five glacial advances. The
general trend of opinion is to recognize five glaciations in North America, separated from each other by four
interglacial stages. But some students of Pleistocene chronology in the New World would limit the glaciations

in North America to four, regarding one of the generally recognized glacial advances (lowan) as a minor substage
of the larger fourth glaciation.

The subdivision of the Pleistocene in North America has been arranged by Kay,' one of the outstanding au-

thorities on this subject, as follows:

Epoch
 GEOLOGIC SUCCESSION: NORTH AMERICA 1511

The question which concerns us at this place is, of course, the appearance, sequence, and extinction of mam-
mals within the Pleistocene of North America. This subject has been variously investigated by numerous
workers over a long period of years, with the result that two generally opposing views have been formulated. One
of these concepts as to the relationships of the Pleistocene mammals of North America, developed particularly
by Osborn, and elaborated in a much modified form by Hay, is that there was a succession of faunas within the
Quaternary of North America distinguished by the gradual and progressive extinction of many mammalian types,
and perhaps to a by the influx of other types, as a result of changing environmental and climatic
lesser extent

factors. The other theory, which has grown up in recent years among the later students of Pleistocene mammals,
and which has been particularly well by Romer, regards the Pleistocene mammals of North America as
set forth
comprising essentially a unit fauna, appearing for the most part at the beginning of the period and persisting
throughout its extent, to continue into Recent times. Such extinctions as have occurred, and it is largely by these
that the Pleistocene fauna of North America can be distinguished from that of Recent times, took place for the
most part at the "end" of Pleistocene times, which was at no very great date of geological antiquity.

Perhaps Cope was the first author to recognize a distinction of faunas within the Pleistocene, when he desig-
nated an Equus fauna characteristic of the western plains and a Megalonyx fauna typical of the eastern wood-
lands.

Osborn,' in 1910, recognized four faunal "zones" within the Pleistocene, basing them in part upon their sup-
posed sequence in time and in part upon their geographic facies. In designating these divisions he admitted that
the "lines of separation between these zones are by no means clearly defined at present, and will depend in the
future upon the more accurate definition of species." They may be listed as follows:

IV. Cervus zone. Prehistoric fauna of the forests of eastern and western North America. Mammals
entirely of modern types; "Pleistocene" forms extinct.

III. Ovibos-Rangifer zone. Plains and forest faunas. A "cold" fauna accompanying the last glaciation,
and dominated by muskoxen, reindeer, American mastodon, and woolly mammoth.

II. Megalonyx zone. Temperate faunas of middle and late Pleistocene times. Some primitive early
Pleistocene forms still survive, while certain of the "late" mammals, such as Rangifer,
have not as yet appeared.

I. Equus-Mylodon zone. Lower and Middle Pleistocene faunas of the Plains regions. Many types svu'-
viving from Pliocene times. Certain advanced forms, such as true Eurasiatic deer, bear,
mountain goats, sheep or bison have not as yet appeared.

"These zones are not sharply distinguishable chronologically at present; they partly overlap and are partly
successive."

Hay,-' who devoted a considerable portion of his life to the study of Pleistocene mammals in North America,
also recognized a sequence of faunas dependent upon gradual extinction throughout the Pleistocene. But his
theory as to mammalian succession within the was widely divergent from that held by Osborn.
Pleistocene period
According to Hay, there was an early Pleistocene fauna, usually designated by him as of Aftonian age, containing
many genera of mammals, such as Archidiskodon, Equus, Camelops, and the like, which became extinct at the end
'Osborn, H. F.. 1910. "The Age of Mammals," p. 438, and succeeding pages.
-Hay, O. P., 1925. ,Jouni. Wash. Acad. Sci., XV, No. 6, p. 128.
1512 OSBORN: THE PROBOSCIDEA

of "Lower" Pleistocene times. Thus he envisaged a sharp break between the Lower Pleistocene and succeeding
times, due to the wide-spread elimination of forms with the advent of the second glaciation. Subsequently, during
Middle and Upper Pleistocene times there were still more extinctions, so that only a few genera such as Mastodon,
Mammonteus, and Megalonyx persisted to the end of the Quaternary.

We know from actual discovery about the vertebrate life of the firsst glacial stage, the Nebraskan but it must have
little ;

been made up some species from South America, some from Asia, but principally of native species remaining over from the
of
Pliocene. . The first glacial stage.
. . almost annihilated the descendants of the Pliocene mammals, not inured to a severe
. . .

climate. The second glacial stage had nearly wiped out the South American contingent. The survivors were mostly of northern
Asiatic origin, hardened to an inclement environment. Elephas imperator had probably reached North America from southern
Asia and was a weakling. The faunal change that occurred during the first three stages appears to have been more profound
than that of the rest of the Pleistocene. Hence I believe that the division of the Pleistocene into Earlier and Later expresses
best the history of the North American vertebrate animals during the Pleistocene.

Later Pleistocene (Yarmouth Wisconsin) —

"Few edentates, few or no horses, no camels, no Elephas imperator, one or two mastodons, fewer large cats."
Earlier Pleistocene (Nebraskan — Kansan)
"Many edentates, many horses, many camels, Elephas imperator, several mastodons, many large cats."

Among the students who have been working in recent years upon the question of Pleistocene mammals in

North America, there is strong support for the idea that the characteristic Quaternary fauna appeared at the
beginning of that period, to persist throughout its extent, finally to be decimated by a wide-spread series of ex-

tinctions at a relatively recent date —seemingly after the entrance of man into the New World. According to the
proponents of this view, there was very little gradual and progressive extinction during the extent of the Pleisto-
cene.

This theory, although at first sight seemingly less logical than the theory of gradual extinctions occurring
during the Pleistocene, is nevertheless based upon very strong evidence —the result of recent careful stratigraphic
and palaeontologic work in the field. And with the passing of time, each successive year of field work strengthens
the theory of a continuous Pleistocene fauna. Hay's thesis depended upon his assumption that many of the ex-

tinct types of Pleistocene mammals in North America did not survive long after their period of greatest develop-

ment, namely, the Aftonian or first interglacial period. Yet most of the supposed Aftonian faunas, cited by Hay
in sujjport of this theory, cannot be definitely proven as of Aftonian age, while as a result of careful work carried
on in recent years, many of them can be definitely proven as later than Aftonian in age. Moreover, the work of

recent years in the southwestern United States in caverns, and in such valley deposits as that at Clovis, has proven
beyond doubt that practically all of the mammals supposed by Hay to be diagnostic of his Earlier Pleistocene
were still living at a relatively recent date, and were associated with early man in America.

In Romer's' extremely valuable paper on this subject, the conclusions are summarized as follows:

.\ hypothesis which implies that practically all the important fossil forms had existed until a comparatively Recent date and

then become extinct in a geologically short period of time had seemed equally improbable to the writer; and yet it is to such
a conclusion that a study of the evidence leads. A considerable proportion of the large Pleistocene forms now extinct in this
country appear to have existed until post-glacial or sub- Recent time in either the north-east or south-west .... The overwhelm-
ing trend of the evidence is that very little extinction appears to have taken place among mammals during the Pleistocene
proper, and that a vast amount of extinction, reducing the fauna to its present impoverished condition, has taken place in a
comparatively short period which presumably cannot have had its initiation more than, roughly, 20,000 years or so ago.

Of course the idea of a continuous Pleistocene fauna in North America cannot be expressed as an unqualified
fact. Undoubtedly there were some exceptions to the general rule that the Pleistocene mammals in this region

continued from the beginning until the end of the period. For instance, there is every reason to believe upon the
'Romer, A. S., 1933. .\rticle II in "The American Aborigines." Edited by Diamond Jenness. Univ. Toronto Press, pp. 7!), 76.
 GEOLOGIC SUCCESSION: NORTH AMERICA 1513

basis of available evidence, that certain immigrant forms, particularly some of the ground sloths and the capybara
from South America and the woolly mammoth and the bisons from Eurasia, did not reach this continent until
after the opening of Pleistocene times. And again certain forms, notably Stegomastodon, failed to persist into late
Pleistocene times. But except for these few types, all of the Pleistocene mammals in North America lived from
the beginning of the period until a few thousand years ago.

This picture of a generally continuous Pleistocene fauna in North America, while at variance with the earlier
ideas of successive "warm" and "cold" faunas in Europe, corresponds in a general way to the more recent hy-
pothesis of the European succession, by Stehlin, Hopwood, and other students and outlined on
as expressed
previous pages of this chapter. Even so, there would seem to be some real differences between Europe and North
America in the development of their Pleistocene mammals. Thus in Europe the persistence of Pleistocene genera
from the beginning until the end of the period does not seem to be so complete as is the case in North America.
In other words, there was more extinction during the progress of Quaternary times in Europe, and less of a wide-
spread suppression of types at the end or after the end of the last glaciation.

Of course it may be that this difference is illusory rather than real, due to the incompleteness of the knowledge
in both regions. On the other hand, recent work tends to point to the reality of the difference as outlined above.

It is possible, although in the present state of our knowledge the contributing factors are difficult to envisage,

that the difference between the extinction of large Pleistocene mammals in Eurasia and North America may
have been due in part to the evolution of man as a potent force in the history of later Quaternary times. In
Eurasia, man "grew up" with the Pleistocene faunas of that region, so that he was always more or less in a state
of ecological balance mammals around him. In North America, on the other hand, he came in as a late,
with the
aggressive element, and it may have been the entrance of this destructive animal, even though not at first im-
portant, that caused the final extinction of so many great mammals in the New World. Not that man was directly
responsible for the killing off of numerous and large herds of mammoths, horses, and camels, but rather by his

entrance he may have upset an ecological balance, he may have contributed

may have introduced epidemics, or he
in other ways which from this distance are quite obscure, to the downfall of numerous seemingly successful mam-

malian lines. This fact we do know, that almost all of the large Pleistocene mammals of North America were
still living when man entered this continent, and that they became extinct after his appearance in the New World.

Of course it is evident that with a continuous fauna inhabiting North America from the beginning until the
end of the Pleistocene, there arise great difficulties in any attempt to date events within the Pleistocene upon the
basis of mammalian evidence alone. Hopwood 's statement with regard to the Pleistocene mammals of Europe
that "Except with the aid of a long series of fossils, even an approximation to the true date is all but impossible...,"
holds to even a greater degree in North America. It may be possible at some future date, with numerous long
series of fossils at hand, to arrive at approximate dates within the Pleistocene upon the basis of mammalian as-
sociations, yet even this method seems in the state of our present knowledge to be incapable of yielding any very
positive results. For it is so subject to geographic and ecologic factors as to be almost abrogated without the
most complete kind of evidence correlated in all possible ways.

Nor is there much hope for dating within the Pleistocene upon the basis of mammalian evolution, because
generally speaking there was very little evolutionary progress made during the relatively short extent of this
period. Certainly generic evolution was virtually nonexistent. And while it is widely held that many early
Pleistocene species became extinct before the end of the period even this criterion is too vague and tenuous to be
 :

1514 OSBORN: THE PROBOSCIDEA

of much lielp in coi relation studies. Moreover, we know that in many eases the same species continued from the

beginninfi; to the end of the Pleistocene and even into Recent times. Again, it is all too true that much Pleistocene

material is not specifically identifiable, while many Pleistocene species are not valid.

As for the Pleistocene proboscideans of North America, it would seem to be quite certain that the mastodont
genus Stegomastodon lived only through the earlier portion of the period. The same is very probably true of the
genus Cordillerion in Mexico. Conversely, although the evidence is not definitive, it would seem that the woolly
mammoth, "Mammonteus," was not present in the lowermost phases of the Pleistocene. The other genera. Masto-

don, Archidiskodon, and Parelephas, seemingly lived pretty much throughout the extent of the Quaternary period,
while there is very good evidence based upon recent work to show that Mastodon, Parelephas, and Mammonteus
persisted until a relatively recent date and were contemporaneous with early Man in North America.

The stratigraphic position of the mastodont, Morrillia, is not certain. Although Osborn considered it as of

Middle Pleistocene age, this is more jjrobably a Pliocene form.

The range of i)roboscidean genera in the Pleistocene of North America may be indicated as follows

Pleistocene
)wer
Cordillerion
Stegomastodon
Mastodon
Archidiskodon
Parelephas
Mammonteus
 GEOLOGIC SUCCESSION: NORTH AMERICA 1515

10. THE PLEISTOCENE OF MEXICO

Freudenberg (1921/ 1922^) has recognized in a general way three successive divisions in the Pleistocene of
Mexico, as based on the occurrence of mammalian fossils. As he has pointed out, a precise subdivision of the
Quaternary here is not as yet possible, for there remains much to be learned about the succession and relationships
of the Pleistocene mammals
south of the Rio Grande. Naturally, any attempts to correlate the Pleistocene
mammals Mexico with the several glacial or interglacial periods of northern United States and Canada are
of
necessarily of the most provisional nature, particularly in view of the fact that such correlations in North America
are anything but certain.

According to Freudenberg, the truest knowledge as to the succession of Pleistocene faunas in Mexico is to be
had in Tequixquiac, where some fifty species of mammals have been discovered in Pleistocene sediments. The
oldest fauna in this region Freudenberg would correlate approximately with the first interglacial or Aftonian of
North America. This fauna is characterized by the presence of Equus giganteus, Hyaenognathus^ and Preptoceras,
and by the fact that the bones composing it are of a dark brown color. Whether the correlation proposed by
—
Freudenberg is valid is a debatable question for it may be that this author was unduly influenced by the work of
Hay, who regarded many Pleistocene types demonstrably ranging through the period as being restricted to the
Aftonian. The reader is referred to the discussion of the Pleistocene of North America, above, in which this
problem is treated at some length.

From the younger Pleistocene, according to Freudenberg, come Ardotherium simum, Smilodontopsis hysenoides
and many other typical Pleistocene genera. He considers Parelephas columbi as representative of the "jungdilu-
viale Steppenfauna" while he would place Archidiskodon imperator as somewhat older, in an interglacial age. He
correlates a deposit in which a tapir, Tapirus cf. tarijensis, was found with the lUinoian to the north. Finally, he
states that the Megalonyx-Mylodon fauna of Mexico is quite young, as it is in the United States, occurring in
river gravels in Mexico that are correlative with the cave deposits to the north.
Freudenberg lists Cordillerion oligobunis and the four subspecies described by him, as well as Cordillerion
tropicus, in his "Quartarfauna des Hochtals von Mexico."- Yet elsewhere in the same work (1922) he definitely
places two of the subspecies, C. oligobunis antiquissimus and C. oligobunis Jelicis, in the Pliocene. It may be that
Cordillerion in Mexico is of Pleistocene age, very possibly confined to the lower levels. On the other hand, one
cannot be sure but that some of the Cordillerion may come from Upper Pliocene deposits as it would seem evident —
that certain other genera, notably Hyaenognathus and possibly Rhynchotherium, do represent the Upper Pliocene in
Mexico.
On the basis of our present evidence, therefore, it must be recognized that most of the Pleistocene mammals of
Mexico, as listed by Freudenberg, may with good reason range from the lower to the upper reaches of the period.

Proboscideans:
Cordillerion tropicus (Cope)
Cordillerion oligobunis (Cope)
Cordillerion oligobunis antiquissimus (Freudenberg)
Cordillerionil) oligobunis Jelicis (Freudenberg)
>Some of these forms may be of Pliocene age.
Cordillerion{'?) oligobunis intermedius (Freudenberg)
Cordillerion oligobunis progressus (Freudenberg)
Parelephas columbi jelicis (Freudenberg)
Archidiskodon sonoriensis Osborn
Archidiskodon imperator silvestris (Freudenberg)
Archidiskodon imperator falconeri (Freudenberg)
'Freudenberg, W., 1921. Geologie von Mexiko, Svo, Borntraeger, Berlin, pp. 134-145.
-Freudenberg, W., 1922. Geol. u. Pal. Abh., N. F., XIV (XVIII), Heft III, pp. 104, 105, 171-176.
^It might be said in this connection that Hyxnogtiathus, listed by Freudenberg in the Pleistocene of Mexico, is probably of Pliocene age. This genus
ranges from the Lower to the Upper Pliocene in the United States.
 VII. CENTRAL AND SOUTH AMERICA
1. INTRODUCTION
Although proboscideans had migrated into Central America from the north by late Pliocene times, it would
seem that the geographic conditions prevailing in the isthmus may have been sufficiently severe to serve as a
barrier that prevented these great animals from reaching South ^eoipgicj4jL rei^tionShips of
soutk america^ proboscid£a
America in appreciable numbers until the beginning of Pleistocene

times. The world-wide depression of average temperatures, so

characteristic of the Pleistocene, though making itself felt on the

climate and the life of South America, did not cause in this region

the formation of extensive continental glaciers, as in the northern

hemisphere; in fact, glaciation in South America was confined to
the Andean uplift and to Patagonia, in the southernmost tip of

the continent. Over a major portion of the land, cool and dry
conditions, perhaps alternating with warm, moist periods, were
prevalent. Thus there were accumulated over a vast tract the
extensive loess deposits designated as the Pampean beds, in which

fossil mammals of Pleistocene age are abundant. Here and else-

where in South America, in beds of related age, the proboscideans

make their first appearance and follow the course of their charac-
Neotropical development. F'S"'^'^ 1225
teristic
 GEOLOGIC SUCCESSION: CENTRAL AND SOUTH AMERICA 1517

distance. The Pampean loess, as typically exposed, is probably for the most part of eolian origin, but fluviatile

and lacustrine facies are common, with their consequent variations in the composition of the sediments. Thus
there are bands or zones of coarser materials, concretions, 'loesskindl,' limy deposits and the like. The loess
itself, as is typical of this kind of sediment, is very fine-grained, compact, and rather well consolidated, forming
a resistant mantle, and where erosion has cut through it, vertical cliffs. In many localities, particularly in the
easternmost portions of the Pampean exposures, there are intercalated marine deposits, representing temporary
incursions of the sea due to slight depressions of the land surface.

The first Pampean beds were made some hundred years ago by d'Orbigny and by
serious studies of the
Darwin, and subsequently by Bravard and by Burmeister. However it remained for Ameghino, the great Argentine
palaeontologist, to attempt a classification or stratigraphic subdivision of these deposits.

Ameghino, in 1880,' suggested the separation of the Pampean into three successive horizons, as follows:

Terreno pampeano lacustre

Terreno pampeano superior
Terreno pampeano antiquo.

Subsequently locality names were given to these horieons, so that they became:

Lujanense = Pampeano lacustre = Upper Pampean

Bonaerense = Pampeano superior = Middle Pampean
Ensenadense = Pampeano inferior = Lower Pampean.

Naturally in the course of time Ameghino greatly extended his division of the Pampean beds, and subsequent
authors added to or modified the subdivision for the series that had been proposed. Consequently the views as to
the proper stratigraphic classification of the several units comprising the Pampean deposits became diverse, and
the system of classification for these subdivisions in many cases became complex. Suffice it to say at this point
that the three divisions given above constitute the basic separation of the Pampean deposits, and the more refined
or more complex systems of stratigraphic nomenclature for these sediments are based on the essentially tripartite
plan first proposed by Ameghino.

Ameghino, who was inclined to regard all of the South American formations and their contained mammalian
faunas as being much older than is justified by the facts, placed the entire Pampean sequence in the Pliocene.
Needless to say, all other authorities have differed from this interpretation of the age of the Pampean. Roverto, in
1914, regarded the Pampean as essentially of Pleistocene age, but he included within it the Puelchense stage or
division, which by most authors is assigned to the series antecedent to the Pampean (Uquiana or Araucana).
Later, in 1934, Kraglievich regarded the Middle and Upper Pampean as of true Pleistocene age, but he designated
the Lower Pampean, or Ensenadense, as "Upper Pliocene or Pliopleistocene." In 1937, Rusconi, like Kraglievich,
placed the Middle and Upper Pampean in the Pleistocene, but regarded the Lower Pampean as of Upper Pliocene
age, a view that seemingly is shared by Castellanos in his recent work on the Lower Pampean and Araucanean

sediments of the Valle de Los Reartes.

A comparison of the several Pampean faunas, particularly on the bases as outlined elsewhere in this chapter,
would seem to put a somewhat different interpretation on the age of the Pampean beds, than that outlined in the
preceding paragraph. Briefly, the mammals of the Pampean, as compared with those of other portions of the
'Ameghino, F., 1880. "La Antiquedad del Hombre en El Plata," Tome I, p. Ill; 1881, Tome II, p. 334.
1518 OSBORN: THE PROBOSCIDEA

world, give every evidence for us to suppose that the entire Pampean sequence is of Pleistocene age. Thus, the
Ensenadense, instead of being Upper Pliocene, as generally regarded by South American authorities, would seem
to be, on the basis of faunal evidence, of Lower Pleistocene age.

The Pleistocene age of the Pampean wasnumber of years ago by the late W. D. Mat-
effectively set forth a

thew, in his comprehensive essay "Climate and Evolution.'" Matthew showed most clearly that the Holarctic
genera in the Pampean of South America most certainly were immigrants from North America, and that their
appearance in the Neotropical region must of necessity have been made subsequent to their origin in the north.

The first appearance of true equines in South America is in the Pampean. The three be.st-knovvn genera are Equus,
Hippidion and Onohippidion. The first might be regarded as of Palearctic origin;''^' the second and third have no Old World
predecessors, but may be directly derived from the North American Pliohtppus. They are, however, much larger and more pro-
gressive than Pliohippiis, and in size, reduction of the lateral digits, etc., are equivalent to Equus. We can hardly doubt that
they came to South America from North America, nor can I see any practical alternative to believing that Equus arrived by the
same route. Now, the first appearance of Equus in North America is at the ba.se of the Pleistocene. In Argentina, it first
appears in the middle Pampean. The middle Pampean cannot therefore be older and is presumably younger than Lower
Pleistocene. Hippidion and Onohippidion are found {fide Roth) in somewhat older levels; but as they are much advanced
over anything in our Middle Pliocene (Blanco), it would seem that their first occurrence in the Pampean must be placed at the
top of the Pliocene or preferably in the I^ower Pleistocene. I conclude that the Pampean formation approximately represents the
Pleistocene epoch.
. . . the genus Arclotherium. of the true Pampean in South America, unknown in North America until the Pleistocene,
indicates, like Equus, that the Pampean is a Pleistocene formation.
The distribution of Smilodon in North and South America is in exact accord with that of Arciotherium. The relations of
the South American Proboscidea to those of North America correspond to tho.se of the Equidae. The Camelidse, Cervidse,
Canidae, etc., also support the Pleistocene age of the true Pampean.

The differentiation of distinct, successive faunas within the Pampean is not so clear-cut or definitive as might
be expected. For, as is modern genera appear at the base of
typical of the Pleistocene in other parts of the world,
the Pampean, and continue uninterruptedly throughout the sequence, from bottom to top. Therefore, the faunas
of the Ensenadense, Bonaerense, and Lujanense show successive appearances of the same genera, with only specific
changes to differentiate them. And specific differences are of little value in establishing time sequences within the
Pleistocene, as has been shown by Romer in his illuminating paper on the Pleistocene of North America (1933).

Of course there are many relatively primitive forms, characteristically South American, which have persisted
from older beds into the Pampean. But the presence of these conservative or holdover types in the Pampean does
not argue for the antiquity of the beds in which they are found, but rather constitutes once again an example of
the persistence of earlier types into a period later than that of their typical expression. Indeed, the very presence
in the Ensenadense of a large group of thoroughly modern genera is one of the strongest arguments for the Pleisto-
cene age of the base of the Pampean.

There would seem, however, to be certain differences that distinguish the Ensenadense from the overlying
beds. One of these is the presence of the genus Typotherium in the lowest of the Pampean stages and its absence
from the upper beds, for there seems to be valid proof that Typotherium persisted from the Tertiary into the
Ensenadense, at the end of which stage it became extinct. Conversely, the absence of Equus in the Ensenadense
and its presence in the Bonaerense and Lujanense also serve to distinguish this lower horizon from the upper ones.
But, for the most part, the Pampean mammals run through from the lowest to the highest phases of the for-
mation. This is true not only of the autochthonous genera, such as the edentates and 'certain of the noto-

ungulates, but also of the immigrant forms, such as Arctotherium, Canis, Smilodon, Hippidion, the artiodactyls,
and the proboscideans.
'Matthew, W. D., 1915. Ann. N. Y. Acad. Sci., XXIV, pp. 196, 198.
-Studies made subsequent to the writing of Matthew's paper have shown most conclusively that Wguu.s is of North American origin, having been derived
from the Upper Pliocene genus Plesippws, this latter a direct descendant of Pliohippus.
 GEOLOGIC SUCCESSION: CENTRAL AND SOUTH AMERICA 1519

At the present time there is no evidence definitely confirming the fact that any of the South American probo-
scidean genera are Hmited to certain stages of the Pampean. Thus, it would seem that Cuvieronius, Cordillerion,
and N otiomastodon range through the Pampean, although the last of these genera may be limited to the lower
phases of the formation. With regard to this question the reader is referred to page 595 of Volume I of the present
Monograph, where the geologic age of certain species of Cordillerion and Cuvieronius is discussed by Dr. G. G.
Simpson. (Dr. Simpson considers among other things the supposed Lower and Middle Pampean age of Cuvieroni-
us platensis and the Upper Pampean age of Cuvieronius superbus and shows that such restriction of these species
rests upon unsatisfactory evidence.)

Pampean Proboscideans:
N otiomastodon ornatus Cabrera
Cuvieronius superbus (Ameghino)
Cuvieronius platensis (Ameghino)
Cuvieronius maderianus (Ameghino)
Cuvieronius perayuiensis (Gez)
Cuvieronius bonaerensis (Moreno)
Cuvieronius rectus (Ameghino)

4. THE PLEISTOCENE OF THE ANDEAN VALLEYS

Tarija, Bolivia. —During the time that the diverse loesses of the Pampean formation were being deposited
on the broad plains of Argentina, sediments of somewhat different origins but of a similar age were being ac-
cumulated in the high mountain valleys of the western Cordilleras. These were the sandstones, clays, and gravels
of fluviatile origin, which were laid down by mountain streams
form discontinuous but locally extensive de-
to
posits in the numerous valleys at the bases of the ever-uplifting Andean mountains.

Typical of these mountain valley deposits are the sediments exposed in the valley of Tarija in southern
Bolivia, the fauna of which has been thoroughly monographed by Boule and Thevenin. These deposits consist
of sand and clays, which in many areas have been extensively eroded to form small areas of badlands.

The fauna from the valley of Tarija is comparable to the Pampean fauna, an indication of the virtual
closely
contemporaneity of the mammals in the two regions. Of particular interest in the Tarijan fauna are Cordillerion
andium, Macrauchenia, Toxodon, Equus, Hippidion, Onohippidium, Smilodon, and various ground sloths. One
difference is to be noted between the Pampean assemblage and that of Tarija, namely, the presence of Typotherium
in the former and its absence in the latter. As has been shown above, this genus presumably is found only in the
Lower Pampean beds of Argentina, having become extinct before the deposition of the Middle and Upper Pam-
pean deposits. Therefore, its absence from the Tarija fauna may mean
one of two things, either that these beds
are later than Lower Pampean, being generally correlative with the Middle and Upper Pampean of the plains, or
that the ecological conditions were such as to prevent the genus Typotherium from venturing into the elevated
areas of the mountains.

Enrico de Carles divided the Tarija beds into two horizons, a lower level containing among other things
Cordillerion, Hippidion, Macrauchenia, and Equus, and an upper level with Glyptodon, Mylodon, Megatherium,

and again Equus. But, as Boule and Thevenin have shown, this division will not hold —rather the fauna seems to
continue throughout the extent of the beds.
1520 OSBORN: THE PROBOSCIDEA

It has been the opinion of many authors that the Tarija deposits are in part at least of Upper Tertiary age.
Ameghino even went so far as to place these beds in the Lower Yet as Boule and Thevenin have most
Pliocene.

amply demonstrated, the Tarija deposits are fully correlative with the Pampean to the east, and as such are to be

regarded entirely as of Pleistocene age.

Proboscideans:
Notiomastodon argentinus (Ameghino)
Cordillerion tarijensis (Ameghino)

What has been said about the Tarija deposits applies, for the most part, to other mountain valley beds of the
Andes. That is, they are essentially of Pampean age, and therefore come within the Pleistocene.

Ulloma, Bolivia. —Philippi, in 1893, described a small mammalian fauna from UUoma, which in most
respects is similar to the Tarija fauna and likewise to the Pampean fauna. This assemblage from Ulloma is
characterized by Hippidion, Cordillerion, and various ground sloths.

Proboscidean: Ulloma, Bolivia

Cordillerion bolivianus (Philippi)

Proboscidean: Lake Tagua-Tagua, Chile

Cordillerion chilensis (Philippi)

Ecuador. —The localities and ages of Cuvier's types of Cordillerion andium and Cuvieronius humboldtii.

The type of Cordillerion andium, described by Cuvier, was found by von Humboldt near the volcano of

Imbaburra, at Quito, Ecuador. This specimen came from an elevation of some 7,600 feet —undoubtedly from
mountain valley beds contemporaneous in some degree with the sediments of Tarija, and with some portion of the
Pampean series of Argentina. Consequently the age of this species is Pleistocene, and it is very probable that
Cordillerion andium persisted into very late Pleistocene times. It is the typical notorostrine of the high cordilleran
valleys of South America.

Although Cuvieronius humboldtii, described by Cuvier, is the typical Pampean plains form of Argentina, the

type of this species was found in the Andean region. The locality is near Concepcion, either in Chile or in Ecuador.

Whatever may be the locality for the type, there would seem to be little doubt that it was found in one of the high
mountain valleys, in beds that were probably more or less equivalent to the Tarija deposits and to the Pampean
of the plains. Therefore the age of this form is Pleistocene.

Proboscideans:
Cordillerion andium (Cuvier)
Cuvieronius humboldtii (Cuvier)

In a notable study of the fossil mammals of Ecuador, Spillmann in 1931 presented a detailed description of
the geologic relationships of the various extinct mammals of that region. The facts are somewhat as follows.

During Pleistocene times an extensive tuff or volcanic ash deposit was built up throughout the mountain
valleys of the region. This deposit, which in places reached a considerable thickness, is of the nature of an eolian

accumulation and therefore forms a wide-spread mantle through the numerous mountain valleys. It would seem
to be indicative of deposition under dry conditions, not only by reason of the nature of the sediments themselves,

but also because the contained fauna shows adaptations to such a habitat.
 GEOLOGIC SUCCESSION: CENTRAL AND SOUTH AMERICA 1521

In the Quebrada of Chalang, a canyon tributary to the Rio Colorado valley and opposite Punin, Spillmann
found a fairly extensive fossiliferous deposit in the tuffaceous beds, known locally as the Cangahua. The fossils

were in the basal portion of the tufT at this locality, which may be rather high up in the tuflf as it is generally
developed. The fauna includes certain characteristic South American Pleistocene genera, such as Mylodon,
Megatherium, Protauchenia, Neohippus, Hippidion, and Cuvieronius; consequently it is to be considered as of
Pleistocene age, probably equivalent to some part of the Pampean assemblage. Such was the opinion of Spill-
mann, who described the material.

In Volume Monograph, page 567, will be found a map and a stratigraphic diagram, showing the
I of this

location and the geological occurrence of the Chalang fauna.

Proboscidean: Cuvieronius mjorae (Spillmann).

Near the town of Alangasi, in the Quebrada Cachihuayco, Spillmann discovered a rather complete skeleton of
Cuvieronius in a superficial layer, associated with man. This remarkable find is clearly indicative of the fact that
some of the South American proboscideans persisted until a very recent date —certainly to within a few thousands
of years ago.

The Alangasi specimen was found in a fine, bluish clay, which rests on the tufT or Cangahua. Evidently the
clay is a late, post-Pleistocene deposit, originally a soft mud in which the animal was trapped and finally perished.

As Spillmann has shown, the very nature of this deposit not only was responsible for the death of the proboscidean
but also for its exceptional preservation. These sediments form a locally developed terrace at the place where the
fossil was discovered.

The specimen shows certain marks that may be due to the inflicting of wounds by arrows or spears. With it
were pottery fragments and other clear signs of human workmanship. Thus the age probably is "post-Pleisto-
cene," contemporaneous with some of the early South American Indians.

For further details as to the occurrence of this specimen, the reader is referred to Volume I of this Monograph,
pages 571-574.

Proboscidean: Cuvieronius postremus (Spillmann).

5. THE PLEISTOCENE OF BRAZIL AND FRENCH GUIANA

Brazil. —One of the notable discoveries of a South American proboscidean is Cuvieronius brasiliensis, first

described by Lund almost a hundred years ago. This specimen was found in a limestone cave, in association with
sloths, carnivores, and other typical South American mammals. The age is difficult to determine, but probably
is late Pleistocene.

Proboscidean: Cuvieronius brasiliensis (Lund).

French Guiana. —A fragment of a tooth constitutes the only record, known at the present time, of the
extension of the elephantines into South America. This specimen was found in Cayenne, French Guiana, and is

listed by Osborn as "probably Upper Pleistocene." Nothing more can be said as to its age.

Proboscidean : Parelephas columbi cayennensis Osborn.

 uniform scale of one o.ic-hnndredth natural
FiK 1226 Models of Recent and extinct Mammoths and Mastodons, all ledufcd to a
Knight 1914, to show the external characters and pro-
Executed under the direction of Henry Fairfield Osborn by Charles R.
in
SIZR
World Series of Elephants and Mastodons" (1914.411).
portions. See Osborn. "Restoration of the

Imperial Elephant (Pleistocene) =ylrc/iid;sA-()(7oR impcratnr.

African Elephant (Recent) = Loxodonta africana.
Woolly Mammoth (Pleistocene) = MamDioiilms primigm-
"Pigmy" African Elephant (Recent) = Loxod/ynla africana
lUS.
cyclolis (Elephas cyclotin of Matschic).
American Mastodon (Pleistocene) =Ma)iUidon americanus.
Asiatic Elephant (Recent) = FAephnu indicun ceylaniciiK.

1522
 :

Chapter XXIII

AFFINITIES, MIGRATIONS, AND PHYLOGENY OF

THE PROBOSCIDEA: A SUMMARY

1. Introduction together with hst.s of: 3. Characters, affinities, and migrations of the Proboscidea:

^ J •,• McEritheres, Deinotheres, Longirostrines, Gnatha-

'

belodonts, Amehelodonts, Tetralophodonts, Notoros-

Families
trines, Rhynchorostrines, Brevirostrines, Humboldt-
Snhfnmilips
ma^, Serridentines, Platybelodonts, Notiomastodonts,
Cpnera
Palaiomastodonts, Mastodonts, Zygolophodonts,
Species"
*^ (with horizons)
Stpgolophodonts, Stegodonts, Mammontines, Loxo-
dontines, Elephautines.
2. ExpUmation of terms used throughout the text of the present 4. Skeletal material.
Memoir. 5. Heights of proboscideans, estimated and actual.

1. INTRODUCTION
During a period of fifteen years, namely, from 1920, when intensive work on the present Memoir was inaugu-
rated, to 1935, Professor Osborn collected notes, articles, and illustrations for his proposed chapter on the affinities,

migrations, and phylogeny of the Proboscidea, in which he intended to summarize the results of his own researches
as well as those of the palaeontologists of the world. It is a regrettable fact that this chapter was never written,
and in the following attempt to present his conclusions, it cannot be too strongly impressed upon the mind of
the reader that in no respect any deviation from the views expressed by him in his various publications,
is there
especially those in his final articles of 1934 and 1935, and in Volume I of the present Memoir. Quite naturally
certain of his earlier writings have become obsolete owing to the advance in knowledge of these interesting animals,
and for this reason modifications in some of the citations have had to be made, such alterations being indicated by
the use of dots (to show the omission of statements regarded as erroneous at the present time), or by square
brackets (embodying his final conclusions). For the sake of clarity and ease of presentation, however, there is in
general purposive avoidance of exact quotations; rather is this a paraphrased narrative of proboscidean evolution
since Eocene time, as revealed in the fossil remains unearthed over a period of more than three hundred years and
as interpreted by Professor Osborn in his writings and illustrations.

The accompanying jihylogenetic charts by Mrs. Margret Flinsch Buba, with migration maps by Miss D. F.
Levett Bradley, while the conception of Professor Osborn, were not wholly prepared under his direction but
(since the death of Professor Osborn in 1935) in consultation with the various curators of the American Museum
conversant with the subject, and in consistent agreement with the text and illustrations of the Memoir, thus
expressing his final views.

Perhaps no more fitting introduction to this chapter could be chosen than Professor Osborn's opening words
in his article in Natural History of January-February, 1925 (Osborn, 1925.637)

There are few joys in life comparable with that which the naturalist experiences when one of his predictions or prophecies
happens to be fulfilled. In 1900 I predicted that Africa would prove to be the cradle of the Proboscidea; in 1903 this prophecy
was verified by British explorers in Egypt. Naturally eager to visit the scene of this discovery at once, I refrained until my
British friends had fully described and published this and other discoveries and gained the world-wide reputation therefor to
which they were richly entitled. I then asked President Theodore Roosevelt for an introduction to Lord Cromer, at the time
Viceroy of Egypt, and through the generosity of President Jesup of the American Museum an expedition was fitted out, carrying
a,s credentials a thoroughly characteristic note from President Roosevelt to Lord Cromer [who.se] brief and simple diplomatic
. . .

message opened the doors of Egypt to the American Museum party. On our arrival at Shepheard's Hotel on the morning of
1523
1524 OSBORN: THE PROBOSCIDEA

January 23 a card was sent up announcing Captain H. G. Lyons, then director of the Geological Survey of Egypt, who
[1907],
thereupon assured —
me that all the resources of the Survey would be placed at our disposal, a camel caravan, a supply of the
absolutely essentisd fantasses for carrying water, and, best of all, the guidance of a most intelligent and delightful member of the
Survey staff, Mr. Hartley T. Ferrar. A personal caravan was also engaged. Thus, sixty camels strong, we wound our way past
the pyramids of the eastern side of the Nile, skirted the fertile basin of the Fayum, and struck southwest into the waterless
desert until we reached the region that represented the ancient cradle of the elephant family. We at once set to work with a
very superior force of Egyptian excavators from Kuft, under the direction of Mr. Walter Granger and Mr. George Olsen, two
of the best fossil hunters of America, who .stuck to their arduous post for nearly two months, until driven out by sandstorms
and excessive heat. With their skilled aid, we soon discovered the burial .sites of three of the early elephant dynasties; the
McERiTHERiuM, the abundant Phiomia, and finally the rare Pal^omastodon. The last-mentioned name is derived from the
uncorrupted Greek words iraXatos, /xao-ros, and 686vs, signifying 'the ancient nipple tooth.'
The Fayum Expedition .... aroused in the writer's mind the liveliest interest in these relatively small and primitive probo-
scideans, and a desire to compare them closely with the large proboscideans of France and South America, which were first
described in 1806 by the famous Cuvier, also the wish to compare them with the proboscideans described and figured by the
British explorers Falconer and Cautley in India between the years 1845 and 1847, and finally the hope to trace all these animals
from their ancestral homes in Africa and Eurasia through their migrations to America.
An insatiable Wanderlust has always possessed the souls of elephants as it has those of the tribes and races of man. Not
only to overcome the changes and chances of this mortal life, but also to gratify their intelligent curiosity ever to explore
afresh forests, pastures, fields, rivers, and streams, they have gone to the very ends of the earth and have far surpassed man
in adapting their clothing and teeth to all possible conditions of life. Thus the romances of elephant migration and conquest are
second only to the romances of human migration and conquest. Variety is the spice of elephant life, as it is of human life, and the
very longing for a change of scene and of diet has been the indirect cause of what in scientific parlance we term adaptive radia-
tion— the reaching out in every direction for every kind of food, every kind of habitat, in itself the cause of radiating or divergent
evolution and adaptation. It is to this predisposition to local, continental or insular, and world-wide wanderings that we at-
tribute the many branches and sub-branches which have been developed in this remarkable family.

The by Professor Osborn (see Vol. I, p. 21) "is the

distinguishing feature of the present Memoir, as stressed
separation of many direct phyletie Unes of descent from each other and the recognition of many mastodonts and
elephants which are very much alike in certain characters, but which are still more unlike in other characters and
cannot possibly be descended one from the other."

It will be noted by comparing Plates x and xi of Volume I that the author in 1935 departed somewhat
radically from his phylogenetic arrangement of 1934. The present treatment, therefore, will follow the author's

1935 chart. The reader is also referred to Chapter XXI of the present volume which gives a detailed history of

proboscidean nomenclature.

Five Superfamilies
Four main divisions (suborders or superfamilies) of the Proboscidea were adopted in 1921 (Osborn, 1921.515,
p. 2) as follows: "With the reservations, first, that we should not expect to find different orders of mammals
subdivided into branches of equal rank and, second, that the subdivisions of the Proboscidea are either of sub-
ordinal or of superfamily value, we may adopt as the four primary divisions:

I. MCERITHERIOIDEA typified by the Mceritherium in the Oligocene of North Africa

II. DINOTHERIOIDEA [Deinotherioidea] typified by the Miocene and Pliocene Dinotheres [Deino-
theres] of Eurasia

III. MA8T0D0NT0IDEA to include the Bunomastodontidae, new family, and the Mastodontidae
IV. ELEPHANTOIDEA to include the Elephantina?, Stegodontinae, and Mammontinse"
This fourfold classification was retained by the author until the publication of Volume I of the present

Memoir in which he withdrew the true Stegodon, subfamily Stegodontinae, from the Elephantoidea, creating for

this genus and subfamily a new superfamily, the

V. STEGODONTOIDEA (see Vol. I, pp. 22 and 25, also Vol. II, Chap. XIV, p. 807)

Thus the Proboscidea are now divided into five suborders or superfamilies.
 : :

SUMMARY 1525

Eight Families
The author's classification of 1933 included five families (see pp. 30 to 33 of Vol. I). While the family name
Bunomastodontidse Osborn ismost appropriate as characterizing the dentition of its included genera and species,
it is without a type genus. Of family names based on generic names, a choice between Trilophodontidae (as in
Simpson, 1931, p. 281) and Gomphotheriidae (as in Cabrera, 1929, pp. 74, 75) would depend upon the acceptance of
Gomphotherium Burmeister, 1837, as a valid genus, and its priority over Trilophodon Falconer and Cautley, 1846.
Doctor Simpson (in a personal communication to the editor) now considers Gomphotheriidae as preferable under
the International Rules and common usage. Professor Osborn 's judgment was that Trilophodon should be used
rather than Gomphotherium. If, therefore, he had followed the ordinary rule for family names, he would have
preferred Trilophodontidae.

A similar condition exists in connection with the family name Humboldtidse Osborn, i.e., it is without a type
genus. In 1934 (see PI. x, Vol. I), Professor Osborn withdrew Stegomastodon Pohlig, 1912 (which antedates both
Eubelodon Barbour, 1914, and Cuvieronius Osborn, 1923) from the Brevirostrinae and placed it in the Humboldtine
group. Dr. William Berryman Scott in his revision of "A History of the Land Mammals of the Western Hemi-
sphere," 1937, pp. 267, 281, 294, suggested the name Stegomastodontidae for the name Humboldtidae.

Professor Osborn did not accept the ordinary rule that a family or subfamily name must be based upon an
already named genus. In giving names to new families and subfamilies he followed the precedents in naming
suborders and orders from appropriate characters (e.g. Proboscidea) rather than from type genera. Consequently
his names are used in the present chapter, which is a summary of his work.

1. MCERITHERIID.E Andrews-Osborn (1906-1921)

2. CURTOGNATHID.E (Curtognati Kaup, 1833; Curtognathids Osborn, 1933)
3. MASTODONTIDiE Girard, 1852; Osborn, 1918, including the Palaeomastodontidae of Andrews, 1906
4. BUNOMASTODONTIDiE Osborn, 1921 (Gomphotheriidae Cabrera, 1929, in part; Trilophodontidse Simpson, 1931, p. 281)
5. ELEPHANTID^ Gray, 1821

In the Appendix to Volume I two new families are described (pp. 722 and 729 respectively)

6. HUMB0LDTIDJ5, to include Eubelodon, Cuvieronius, and Stegomastodon, as distinguished from Cordillerion of the
family Bunomastodontidse, subfamily Notorostrinse, in which subfamily it was thought at
one time (see p. 541) that these four genera should be included. Professor Scott in his new
edition of "A History of the Land Mammals of the Western Hemisphere," 1937, has sub-
stituted the name Stegomastodontidae for Humboldtidse, based on the genus Stegomastodon.

7. SERRIDENTIDiE, to include Serridentinus Osborn, typified by S. serridens Cope, Ocalientinus Frick, Serbelodon Frick,
and Trobelodon Frick, all of the subfamily Serridentinse; Platybelodon Borissiak and Tonj-
nobelodon Barbour of the subfamily Platybelodontinae, and N
otiomastodon Cabrera of the
subfamily Notiomastodontinae.

To these should be added the family recently (1935) described by Drs. Chung-Chien Young and A. Tindell
Hopwood
8. STEGODONTIDjE, regarded by these authors as comprising two groups, one with "compressed, tectiform ridges" (Stegorfon
Falconer and Cautley), the other with ridges more blunt and "composed of rounded
—
conules" {Stegolophodon Schlesinger) see Hopwood, 1935, p. 72. According to the views
of the present author, this family would embrace the true Stegodon only, the Stegolophodonts
having been removed by him to the Mastodontidse, subfamily Stegolophodontinse (see p.
700 of Vol. I for definition).

Therefore, in the final classification of the present Memoir, there are eight families.
 : ;

1526 OSBORN: THE PROBOSCIDEA

Twenty-one Subfamilies
In direct opposition to the morphological and horizontal system of classification of Cope, and despite the
highly \'alued opinion of such colleagues as Dr. William Diller Matthew and Dr. William King Gregory, who
differed with Professor Osborn as to the practicability of a phylogenetic system of classification, which phyla
were designated as subfamilies, he held to the last the firm conviction that a subfamily division could be "proper-
Thus, beginning in 1910 ("Age of Mammals," pp. 558, 559), he adopted
ly applied to vertical lines of succession."
three subfamilies of the Proboscidea, namely, the Dinotheriinse, Mastodontina?, and Elephantina?, the number
increasing until in 1933 (see Vol. I, pp. 30-33) he had included seventeen, and in 1935 (his final classification) as
many as twenty-one subfamilies, of more than half of which he was the author

^- McERiTHERiiN^ Wlnge-Osbom (1906-1923)

^^MirithM-iidffi

^" Deinotheriin^ Bonaparte-Winge-Osborn (1841, 1850-1906-1910)

^Curto^nathidffi
3. LongirostrinjE Osborn, 1918
Mastodontoidc'i ^" Gnathabelodontin^ Bailjour and Sternberg, 1935
'
Bunomastodontidffi 5. Amebelodontin^ Barbour 1929
[= Trilophodontid£e "• Ietralophodontin.e van der Maarel, 1932
Simpson, 1931]'' NoTOHOSTRiNiE Osborn 1921
l
8. Rhynchorostrin^ Osborn, 1918
9. Brevirostrin^ Osborn, 1918
Humboldtidffi
[
= Stegomastodontidse 10. HuMSOLDTiNiB Osborn, 1934, 1936
Scott]

11. SerridentinjE Osborn, 1921

Serridentidae 12. Platybelodontin^ Borissiak, 1928
13. N0TIOMASTODONTIN.E Osborn, 1936

Mastodontidffi
^^- Pal^omastodontin^ Osborn, 1936
= jMammutida> ^^' Mastodontin.e Brandt-Osborn (1869-1910)
J

Cabrera
'' in part I
^^' Zygolophodontin^ Osborn, 1923
17. STEGOLOPHODONTiNiE Osborn, 1936
Stegodontoidca
^g Stegodontin^ Osborn, 1918
Stegodontidse

Elephantoidea 1^. Mammontin^ Osborn, 1921

Elephantidffi 20. Loxodontin^ Osborn, 1918
21. Elephantine Osborn, 1910

Forty-four Genera
Out of the more than ninety names Chap. XXI, pp. 1371-1382) applied generically or subgenerically to
(cf.

the Proboscidea since the time of Linnaeus in 1735, Professor Osborn selected forty-four as valid. This will seem
a surprisingly large number in view of the fact that for many years only two, Elephas and Mastodon, were accepted
by the majority of scientists.

Elephas. —The name Elephas was first used by John Ray in his "Synopsis Methodica Animalium (Juadru-
pedum et Serpentini Generis" of 1693, p. 131; in 1735 Linnaeus in the first edition of the "Systema Naturae,"
p. 10, placed Elephas in the same division as the rhinoceroses; in 1754 in his "Animalia Rariora, Imprimis, et
Exotica," p. 11, he gave Elephas indicus as the type, for which he subsequently (10th edition of the "Systema
Naturae," 1758, p. 33) substituted Elephas maximus. This genus, together with other names appearing in the
same edition, was officially adopted by the members of the Fifth International Zoological Congress of 1901
hence the genus Elephas dates from 1758.

'[See previous page where Doctor Simpson states tliat he now (October, 1910) prefers Goinphotlicriidic Cabrera, 1929. — Editor.)
 SUMMARY 1527

Mastodon. —The name Mastodon in its original or French form was the "Mastodonte" of Cuvier, 1806.

Ten years later (1816) Oken first used the form Mastodon, based on the five classic species of Cuvier of 1806, thus
anticipating Cuvier by one year, who in 1817 in his "Le Regne Animal," p. 233, described Mastodon giganteum

and M. angusiidens. Inasmuch, however, as Cuvier was the author of Mastodonte, he has been given precedence
over Oken in the use of the scientific form Mastodon.

Genera of the Mceritherioidea, Deinotherioidea, and Mastodontoidea Regarded

AS Valid by the Present Author
1. Mceritkeriimi Andrews, 1901 17. Amebdodon Barbour, 1927
2. Deinotherium Kaup, 1829 18. Tetralophodon Falconer, 1847, 1857
3. Palseomastodon Andrews, 1901 19. Morrillia Osborn, 1924
4. Miomastodon Osborn, 1922 20. Cordillerion Osborn, 1926
5. Pliomastodon Osborn, 1926 21. Pentalophodon Falconer, 1857, 1865
6. Mastodon Cuvier, 1806, 1817 22. Anancus Aymard, 1855, 1859
7. Zygolophodon Vacek, 1877 23. Synconolophus Osborn, 1929
8. Turicius Osborn, 1926 24. Cuvieronius Osborn, 1923
9. Slegolophodon Schlesinger, 1917 25. Eubelodon Barbour, 1914
10. Rhynchothermm Falconer, 1856 (MS.), 1863, 1868 26. Stegomastodon Pohlig, 1912
11. BUckotherium Frick, 1933 27. Serridentinus Osborn, 1923
12. Aybelodon Frick, 1933 28. Ocalientinus Frick, 1933
13. Trilophodon Falconer, 1846, 1857 29. Serbelodon Frick, 1933
14. Megabelodon Barbour, 1914, 1917 30. Trobelodon Frick, 1933
15. Gnathabelodon Barbour and Sternberg, 1935 31. Platybelodon Borissiak, 1928
16. Phiomia Andrews and Beadnell, 1902 32. Torynobelodon Barbour, 1929
33. Notiomastodon Cabrera, 1929

Genera of the Stegodontoidea and Elephantoidea Regarded

AS Valid by the Present Author
34. Slegodon Falconer and Cautley, 1847, 1857 39. Loxodonta F. Cuvier, 1825-1827
35. Archidiskodon Pohlig, 1888 40. Palxoloxodon Matsumoto, 1924
36. M etarchidiskodon Osborn, 1934 41. Hesperoloxodon Osborn, 1931
37. Parelephas Osborn, 1924 42. Elephas Linnaus, 1735-1758
38. Mainmonteus Camper-Osborn, 1788-1924 43. Hypselephas Osborn, 1934, 1936
44. Plaielephas Osborn, 1934, 1936

Valid Species of Proboscidea

A complete list of the 552 species and subspecies of the Proboscidea described since 1754 will be found in
Chapter XXI of the present Memoir, pages 1382 to 1420. A list of the valid species of the Mceritherioidea (5),

Deinotherioidea (12), and Mastodontoidea (205), together with migration map (Fig. 1227) is repeated here
from Volume I, pages 735-743, followed by a similar list of the valid species of the Stegodontoidea (19) and
Elephantoidea (111), with migration map (Fig. 1228)—in all 352 valid species and subspecies as determined by the
present author, of which at least 22 are existing species, subspecies, or geographical varieties. For a detailed
treatment of the geologic succession of the Proboscidea, reference should be made to Chapter XXII above by
Dr. Edwin H. Colbert.
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 OSBORN'S FINAL (1935)

CLASSIFICATION OF THE MCERITHERIOIDEA, DEINOTHERIOIDEA, AND MASTODONTOIDEA

Superfamily: MCERITHERIOIDEA
Fam.: MCERITHERIIDiE. Subfam.: Mceritheriin^

Genus : Moeritherium

Primitive aquatic Proboscideans, Hippopotamus-like, or Sirenian-like bodies

McERiTHERiuM Andrews, 1901 Page

Mceiitherium andrewsi Schlosser, 1911, Fay(im, Egypt, Fluvio-marine formation Lower Oligocene 61, 74
Moeritherium trigodon Andrews, 1904, Fayijm, Egypt, Fluvio-marine formation Lower Oligocene 57, 74
Moeritherium lyonsi Andrews, 1901, FayOm, Egypt, Qasr-el-Sagha formation Upper Eocene 54, 72
Moeritherium gracile Andrews, 1902, Fayum, Egypt, Qasr-el-Sagha formation Upper Eocene 56, 73
Moeritherium ancestrale Petronievics, 1923, Fayum, Egypt, Qasr-el-Sagha (?) formation Upper Eocene 65, 76

Superfamily: DEINOTHERIOIDEA
Fam.: CURTOGNATHIDiE. Subfam.: Deinotheriin^

Genus : Deinotherium

Very large downturned inferior tusks, no superior tusks; [Hat cranium], elephantine bodies

Deinotherium Kaup, 1829. Partial list of species

Deinotherium hopwoodi sp. nov., Olduvai, southeast shore Lake Victoria, Africa Middle Pleistocene 117
Deinotherium gigantissimum Stefanescu, 1892, Gaiceana, Rumania Lower or Middle Pliocene 95
Deinotherium indicum Falconer, 1845, Perim Island, India Middle Pliocene 90, 105, 114
Deinotherium giganteum Kaup, 1829, Eppelsheim, Germany Lower Pliocene 80, 84, 86, 114
Deinotherium uralense Eichwald, 1835, Ural Mountains Lower Pliocene 85, 87, 114
Deinotherium podolicum Eichwald, 1835, Podolia, Russia Pliocene (?) 85, 87
Deinotherium intermedium De Blainville, 1845, France Upper Miocene (?) 85, 90, 115
Deinotherium bavaricum von Meyer, 1831, Gmiind, Bavaria Upper Miocene 84, 99, 107, 115
Deinotherium hungaricum Ehik, 1930, Kotyhaza, Hungary, Burdigalian (?) Lower Miocene (?) 115, 116
Deinotherium indicum gajense Pilgrim, 1912, Bugti Hills, Baluchistan, Bugti beds Lower Miocene 85, 105, 115
Deinotherium cuvieri Kaup, 1832, Chev-illy, France Lower Miocene 85, 87, 90, 107, 115
Deinotherium hobleyi Andrews, 1911, Karungu, British East Africa Lower Miocene 104, 115

Superfamily: MASTODONTOIDEA
Fam.: MASTODONTIDiE. Subfam. nov.: PALiEOMASTODONTiNvE

Genus : Palseomastodon

Superior tusks unknown; inferior tusks short, rounded. Progressive grinders, with six cones, proto- and metaconules closing the
median sulcus. Not ancestral to Mastodontina?

PaLjEomastodon Andrews, 1901

Palseomastodon beadnelli Andrews, 1901, Fayum, Egypt, Fluvio-marine formation Lower Oligocene 54, 147
Palseomastodon intermedius Matsumoto, 1922, Fayum, Egypt, Fluvio-marine formation Lower Oligocene 63, 146
Palseomastodon parvus Andrews, 1905, Fayiim, Egypt, Fluvio-marine formation Lower Oligocene 59, 146

1529
1530 OSBORN: THE PROBOSCIDEA

Fam.: MASTODONTlDiE. Subfam.: Mastodontin^

Genera: Miomastodon, Pliomastodoti, Mastodon

Superior tusks and inferior tusks (wlien present) rounded, enamel disappearing; typical mastodontine bodies
Species of Eurasia and North America in approximate theoretic descending order, apparently monophyl(>tic, directly ancestral to
typical Mastodon

Mastodon Cuvier, 1806, 1817. Superior tusks without enamel; molars with median sulcus and progressively sharpened crests Page
Mastodon acutidens sp. nov., Rochester, Indiana Pleistocene —
Postglacial (IV) 696
Mastodon atnen'canus alaskensis Frick, 1933, near Fairbanks, Alaska Pleistocene 176
Mastodon oregonensis Hay, 1926, Rye Valley, Baker County, Oregon Pleistocene(?) 173
Mastodon americanus plicatus Osborn, 1926, Walnut, Bureau County, Illinois —
Late Pleistocene Postglacial
(IV) or post-Wisconsin (?) 173
Mastodon americanus rugosidens Leidy, 1890, Beaufort County, South Carolina Pleistocene 171
Mastodon raki Frick, 1933, Hot Springs, New Mexico Pleistocene 175
Mastodon americanus rupertianus Richardson, 1854, Lake Winnipeg basin, Manitoba, Canada . . Pleistocene 137
Mastodon americanus Kerr, 1792, Big-Bone Lick, Boone County, Kentucky Late Pleistocene (IV Gla-
cial = Wisconsin time) 170
Mastodon grangeri Barliour, 1934, Pender, Thurston County, Nebraska Pleistocene (1st Interglacial,
Aftonian, or II Glacial) 175, 695
Mastodon moodiei Barbour, 1931, near Milford, Seward County, Nebraska Pleistocene (II Glacial) 174
Mastodon progenius Hay, 1914, Missouri Valley, Harrison County, Iowa Early Pleistocene 172
Mastodon panlowi sp. nov., Pestchana, Podolia, Russia Pleistocene 694

Pliomastodon Osborn, 1926. Superior tusks with enamel; molar crests intermediate in character (subhypsodont)
Pliomastodon vexillarius Matthew, 1930, near Coalinga, Fresno County, California Late Pliocene 161
Pliomastodon americanus praetijpica Schlesinger, 1919, 1922, Batta-fird, Hungary Middle Pliocene 159
Pliomastodon sellardsi Simp.son, 1930, Brewster, Florida, Bone Valley formation Lower Pliocene 160
Pliomastodon maUhewi Osborn, 1921-1926, Snake Creek B ["Upper Snake Creek"], Sioux
County, western Nebraska Lower Pliocene 157

Miomastodon O.sborn, 1922. Superior tusks with enamel; molars with rounded crests
Miomastodon tapiroides americanus Schlesinger, 1921, 1922, Tasnad, Hungary Lower Pliocene 156
Miomastodon merriami Osborn, 1921, Thousand Creek, Humboldt Covinty, Nevada, Virgin
Valley formation (type) Middle Miocene 154
Pawnee Creek horizon, Colorado (ref.) Middle Miocene 155
Miomastodon depereti sp. nov., Chevilly, Sables de I'Orl^anais, France Lower Miocene 693

Fam.: MASTODONTID^. Subfam.: Zygolophodontin^

Genera: Ztjgolophodon, Turiciua

Mastodontoid molar crowns, valleys open, conelets progressively added

Zygolophodon Vacek, 1877. Molars transversely crested, conelets 4-6, valleys open, median sulcus disappearing
Zygolophodon borsoni zaddachi Jentzsch, 1883, Thorn, west Prussia Upper Pliocene 209
Zygolophodon borsoni Hays, 1834, near Villanova, Asti, Piedmont, Italy (type); western
Eurasia (ref.) Middle to Upper Pliocene 207
Zygolophodon pyrenaicus Lartet MS. (in Falconer, 1857), Lartet, 1859, Ile-en-Dodon
(Haute-Garonne), France (type) Middle Miocene 206
Zygolophodon pyrenaicus aurelianensis Osborn, 1926, Sables de I'Orl^anais, Chevilly (?),
France Lower Miocene 207

TuRicius Osborn, 1926. Molars transversely crested, conelets 4-25, valleys oixni, median sulcus disappearing
Turicius wahlheimensis Klahn, 1922, Wahlheim and Esselborn, Rhcinliessen, Germany Pliocene 282
Turicius virgatidens von Meyer, 1867, Fulda, northeast of Frankfort, (iermany,
'Gelben Lehme' Middle (?) Pliocene 221
Turicius atticus Wagner, 1857, Pikermi, Greece Lower Pliocene 220
Turicius turicensis Schinz, 1824, Elgg, Canton Zurich, Switzerland, Tortonian Upper Miocene 219
Turicius turicensis simorrensis Osborn, 1926, near Simorrc (Gers), France Upper Middle Miocene 219
Turicius tapiroides Cuvier, 1806, 1821-1824; (in Desinarest, 1820-1822), Calcaire de
Montabusard, France Lower Miocene 21/
 '

SUMMARY 1531

Fam.: MASTODONTID^. Subfam. nov.: Stegolophodontin^

Genus: Stegolophodon
Stegodontoid rounded molar cones, valleys compressed; median sulcus persisting in anterior crests
Stegolophodon Schlesingcr, 1917. Molars transversely crested, conelets 4-5+. Page
Stegolophodon stegodontoides Pilgrim, 1913, Lehri, Punjab, India, Upper Siwaliks (?) Upper (?)Pliocene PI. iv, I, 701
Stegolophodon cautleyi Lydekker, 1886, Perim Island, India Middle Pliocene PI. iv, H, 701
Stegolophodon sublatidens Schlesinger, 1917, Teschen (Schlesien), Austria Middle (?) Pliocene PI. iv' A
Stegolophodon lydekkeri sp. nov., Bruni, northwest Borneo Pliocene (?) PI. iv, CJ, 700
Stegolophodon latidens Clift, 1828, near Yenangyaung, Irrawaddy River, Burma,
Irrawaddy Series (fluviatile) Lower Pliocene' PI. iv, D,E,F,701
Stegolopjiodon nathotensis Osborn, 1929, Nathot, Lower Middle Siwaliks, India, Lower
Chinji horizon Mio-Pliocene PI. iv, C
Stegolophodon cautleyi progressus Osborn, 1929, near Chinji Bungalow, summit of Lower
Chinji, 2,000 feet above base of Lower Siwaliks, India Mio-Pliocene PI. iv, B

Fam.: BUNOMASTODONTIDiE. Subfam.: Ryhnchorostrin^

Genera: Rhynchotherium, Blickotherium, Aybelodon
Typical beak-jawed Mastodonts. Mandible strongly deflected; enamel band on superior and inferior incisive tusks (excepting
Aybelodon), inferior tusks upturned; molars with variable central conules, enamel thick, few ridge-crests (4-5)
Rhynchorostrines of the western United States and Mexico
Rhynchotherium Falconer, 1856 (MS.), 1863, 1868
Rhynchotherium falconeri Osborn, 1923, Mt. Blanco, Staked Plains, Texas tapper Pliocene 494
Rhyjichotherium shepardi edense Frick, 1921, Mt. Eden Hot Springs, San Bernardino County,
California Upper Pliocene 496
Rhynchotherium hrowni .sp. nov., San Jose de Pimas, Sonora, Mexico Upper Pliocene 494
Rhynchotherium tlascalse Osborn, 1918, 1921, Tlascala, Valley of Mexico Upper(?) Pliocene 493
{?)Rhynchotheriumfrancisi Hay, 1926, Brazos River, Pittbridge, Burleson County, Texas. .Pliocene(?) . .
501
Rhynchotherium anguirivale Osborn, 1926, Snake Creek B, Sioux County, Nebraska Lower Pliocene 491
Rhynchotherium rectidens Osborn, 1923, near Pawnee Buttes, Weld County, Colorado,
Pawnee Creek horizon Upper Miocene 488
Rhynchotherium shepardi Leidy, 1871, Dry Creek, Stanislaus County, California Upper(?) Miocene 487
Rhynchotherium brevidens Cope, 1889, Smith River, Meagher County, Montana, Deep River
horizon Middle to Upper Miocene 485
Rhynchorostrines of Asia and Africa
Rhynchotherium chinjiense Osborn, 1929, near Chinji Bungalow, India, 600 feet above base
of Lower Siwahks Mio-Pliocene 502
Rhynchotherium spenceri Fburtau, 1918, Moghara Desert, northern Egypt Middle Miocene 485
Rhynchorostrines of Central America
Blickotherium Frick, 1933. Mandible progre.ssively slender; incisors with enamel
Blickotherium blicki Frick, 1933, near Tapasuma, Honduras Pliocene 508
Blickotherium euhypodon Cope, 1884, Driftwood Creek, Hitchcock County, Nebraska,
Repulilican River beds Lower Pliocene 489
Aybelodon Mandible attenuate; inferior incisors without enamel
Frick, 1933.
Aybelodon hondurensis Frick, 1933, near Tapasuma, Honduras Pliocene .509

Fam.: BUNOMASTODONTID^. Subfam.: Longirostrin^

Genera: Trilophodon, Megabelodon
Intermediate molars trilophodont (3 crests) third molars with 4-5+ crests and central conules. Superior tusks flattened, recurved,
;

with enamel band; inferior tusks rounded, triangular, flattened-horizontal, flattened-oblique, massive, sometimes absent. Mandible
hyperlongirostral
Trilophodon of Eurasia and Africa
Trilophodon Falconer, 1846, 1857
Trilophodon hasnotensis sp. nov., near Ha.snot, India, 1,000 feet below Bhandar bone bed
( = Dhok Pathan horizon) Middle Pliocene 279
Trilophodon esselbornensis Klahn, 1922, Esselborn, Westhofen, Rheinhessen, Germany Pliocene 281
Trilophodon {C hoerolophodon) pentelicus Oaudry and Lartet, 1856-1862, Pikermi, Greece
(type) Lower Pliocene 263
Samos Island (ref .) Lower Pliocene 264

^[See page 824 above where reasons are given for regarding this species as of Lower Pleistocene age. — Editor.]
1532 OSBORN: THE PROBOSCIDEA

Trilophudon sendaicus Matsumoto, 1924, Kitayama, near Sendai, Province of Rikuzen, Page
Japan Lower Pliocene 280
Trilophodon connexus Hopwood, 1935, Kansu, China Miocene 702
Trilophodon spectabilis Hopwood, 1935, locality and horizon unknown 702
Trilophodon inopinatus Boris.siak and Beliaeva, 1928, Jilanfiik beds, Turgai region.
Central Asia Miocene 278
Trilophodon angustidens var. libycus Fourtau, 1918, Moghara, northern Egypt, Africa Lower(?) Miocene 260
Trilophodon macrognalhus Pilgrim, 1913, near Chinji, India, Upper Chinji Mio-Pliocene 274
Trilophodon chinjiensis Pilgrim, 1913, Osborn, 1932, near Chinji Bungalow, Lower Chinji. . . Mio-Pliocene 272
Trilophodon angustidens gaillardi Osborn, 1929, Villefranche d'Astarac (Gers), France Upper(?) Miocene 259
Trilophodon engelswiesensis Klahn, 1922, Engelswies, Baden, Germany Miocene 281
Trilophodon sleinheimensis Klahn, 1922, Steinheim, Baden, Germany Middle to Upper Miocene 281
Trilophodon angustidens var. austro-germanicus Wegner, 1908, 1913, Oppeln, eastern
( iermany Middle Miocene 259
Trilophodon palxindicus Lydekker, 1884, near Kamlial, northern Punjab, India,
KamUal horizon Middle Miocene 268
Trilophodon pandionis Falconer, 1857, Larkana district, Sind, India, Lower Manchhar Middle Miocene 267
Trilophodon angustidens minutus Cuvier, 1806, 1824, Saxony Middle Miocene 252
Trilophodon angustidens Cuvier, 1806, 1817, Simorre, France Middle Miocene 252
Trilophodon angustidens cuvieri Pomel, 1848, Gers and Sables de I'Orl^anais, France Lower Miocene 250
Trilophodon angustidens gaujaci Lartet, 1851, Lombez, France Middle(?) Miocene 250
Trilophodon cooperi Osborn, 1932, near Dera Bugti, Baluchistan, Bugti horizon Lower Miocene 275
Trilophodon poniileviensis Mayet-Fourtau, 1918, Chevilly, Pontlevoy, Sables de I'Orl^anais,
France Lower Miocene 283

Trilophodon (Typical, angustidens), 'Prod-tuskers' ('Scaptobelodonts') of America

cf. T.

Mandible tusks rounded, rodlike or oval, horizontal or directed upwards

slender, nearly horizontal or slightly decurved;
Trilophodon pojoaquensis Frick, 1926, Santa F6 marls. New Mexico Mio-Pliocene 320
Trilophodon (Genomastodon) osborni Barbour, 1916, Bristow, Boyd County, Nebraska Lower Pliocene 298
Trilophodon abeli Barbour, 1925, Devil's Gulch, Brown Co., Nebraska Lower Pliocene 311
Trilophodon {Genomastodon) willistoni Barbour, 1914, Devil's Gulch, Niobrara River,
Brown County, Nebraska Lower Pliocene 293
Trilophodon fricki Peterson, 1928, Brown's Park formation, flanks of Douglas Mountain,
Moffat County, Colorado Middle to Upper Miocene 312

Trilophodon (Tatabelodon), 'Ancient-tuskers'

Mandible massive at symphysis, inferior incisors elongate, rounded or triangular, of massive cross-section
Trilophodon (Tatabelodon) riograndensis Frick, 1933, Santa F6 marls, Battleship Mountain,
New Mexico Mio-Phocene 324
Trilophodon (Tatabelodon) gregorii Frick, 1933, Ainsworth, Nebraska Lower Pliocene 324

Trilophodon, 'Oblique-tuskers'
Mandible more
ma.ssive, laterally compressed, or less strongly decurved tusks oblique or
; laterally flattened, directed downwards
Trilophodon dinotherioides Andrews, 1909, northwestern Kansas (?Republican River) Lower Pliocene 291
Trilophodon ligoniferus Cope and Matthew, 1915, Black Hills, South Dakota
(?Republican River level) Lower(?) Pliocene 298
Trilophodon giganteus Osborn, 1921, Eastview, near Dallas, Gregory County, South Dakota. .Lower Pliocene 304

Trilophodon of Eastern Coast of North America

Trilophodon (ITetralophodon) brazosius Hay, 1923, Brazos River, near San Felipe, Texas Pliocene(?) 374
Trilophodon simplicidens Osborn, 1923, Bone Valley formation, near Pierce, Polk County,
Florida Lower Pliocene 285
Trilophodon obscurus Leidy, 1869, Greensburgh, Caroline County, Maryland (?Choptank
formation) Miocene(?) 285

Megabelodon, 'Spoonbill Mastodonts'

Megabelodon Barbour, Mandible tuskless, round in section, extremely elongate, straight or slightly deflected, slender
1914, 1917.
Megabelodon lulli Barbour, 1914, 1917, Snake River, Cherry County, Nebraska Lower(?) Pliocene 294
Megabelodon joraki Frick, 1933, Santa F6 maris, Santa Cruz, New Mexico Mio-Pliocene 326
Megabelodon cruziensis Frick, 1933, Santa F6 maris, Santa Cruz, New Mexico Mio-Pliocene 323
Trilophodon phippsi^ Cook, 1928, Ainsworth, Brown County, Nebraska Lower Pliocene 315

'(This species is included with Megabelodon because of its tuskless mandible. It was evidently the intention of Professor Osborn to place Trilophodon

phippsi in the genus Megabelodon. See footnote on page 329 of Volume I.— Editor.)
 ;

SUMMARY 1533

Fam.: BUNOMASTODONTID^. Subfam.: Gnathabelodontin^

Genus : Gnathabelodon

Mandible tuskless, elongate, spreading broadly at edentulous rostrum, horizontal; no inferior tusks, superior tusks massive,
rounded, without enamel, upcurved, elongate. Molars of bunomastodont type, with central conules
Gnathabelodon Barbour and Sternberg, 1935 Page
Gnathabelodon thorpei Barbour and Sternberg, 1935, Ogallah, Trego County, western Kansas. . .Middle(?) Pliocene 713

Fam.: BUNOMASTODONTID^. Subfam.: Amebelodontin^

Genera: Phiomia, Amebelodon

Phiomia-Amebelodon, 'Shovel-tuskers' of northern Africa and North America

Mandible elongate, slightly upcurved; [lower] tusks elongate, broadened, appressed, alveolar portion concave. Conules large
Amebelodon Barbour, 1927. Inferior incisors with concentric dentinal laminae
Amebelodon frkki Barbour, 1927, Freedom, Frontier County, Nebraska Middle Pliocene 335
Amebelodon {T rilophodon) hicksi Cook, 1922, Wray, Yuma County, Colorado Middle Plioceiie(?) 307
Amebelodon (Trilophodon) paladenlaius Cook, 1922, Wray, Yuma County, Colorado Middle Pliocene 309
Amebelodon sinclairi Barbour, 1930, Freedom, Frontier County, Nebraska Middle Pliocene 337

Ancestral Amebelodontin.e of North Africa

Phiomia Andrews and Beadnell, 1902. [Inferior tusks with concentric dentinal laminae]
Phiomia pygm.se.us Dep6ret, 1897, Cartennien de Kabylie, near Isserville, Algeria Upper(?) Oligocene 246
Phiomia osborni Matsumoto, 1922, Fluvio-marine of the Fayum, Egypt Lower Oligocene 64, 244
Phiomia wintoni Andrews, 1905, Fluvio-marine of the Fayum, Egypt Lower Oligocene 60, 241
Phioinia wintorii (cf. serridens) Andrews ref., Quarry B, FayOm, Egypt Lower Oligocene 240, 241
Phiomia serridens Andrews and Beadnell, 1902, Fluvio-marine of the Fayum, Egypt Lower Oligocene 55, 239
Phiomia minor Andrews, 1904, Fluvio-marine of the Fayilm, Egypt Lower OHgocene 58, 239

Fam.: BUNOMASTODONTIDiE. Subfam.: Tetralophodontin^

Genera: Tetralophodon, Morrillia

Second molars tetralophodont. Mandibles and inferior tusks of medium length (medilongirostral). Inferior tusks flattened, oval
superior tusks enamelbanded
Tetralophodonts of Eurasia
(?)Subgenus Lydekkeria Osborn, 1924. Mio-Pliocene. Third molars four to five crested, brachyodont; single to rudimentary double
trefoils

Tetralophodon (Lydekkeria) falconeri Lydekker, 1877, Potwar district, Punjab, India

[Dhok Pathan horizon] Middle Pliocene 354
Tetralophodon (Lydekkeria) sinensis Koken, 1885, Yunnan, China Mio-Pliocene 355
Tetralophodon Falconer, 1847, 1857. Ridge-crests 5}2-6, brachyodont; double trefoils
Tetralophodon punjabiensis Lydekker, 1886, Punjab, Siwaliks, India [Dhok Pathan horizon]. Middle Pliocene 362
Tetralophodon grandincisivus Schlesinger, 1917, Maragha, Persia Lower(?) Pliocene 360
Tetralophodon bumiajuensis van der Maarel, 1932, Bumiaju, central Java Pliocene 365
Tetralophodon gigantorostris Klahn, 1922, Bermersheim, Rheinhessen, Germany Pliocene 282
Tetralophodon longirostris Kaup, 1832, Eppelsheim, Germany Lower Pliocene 357
Tetralophodon exoleius Hopwood, 1935, Shansi, China, Pontian Lower Pliocene 704

Tetralophodonts of North America

Morrillia Osborn, 1924. Ridge-crests 7^-8)3, hypsodont; double trefoils, thickly coated with cement
Morrillia barbouri Osborn, 1921, Cambridge, Furnas County, Nebraska Middle Pleistocene 377
Tetralophodon
Tetralophodon elegans Hay, 1917, McPherson, Kansas Middle(?) Pliocene 372
Tetralophodon campester Cope, 1878, Sappa Creek, Rawlins County, Kansas, Republican
River horizon Lower Pliocene 369
Tetralophodon fricki sp. nov., near Clarendon, northern Texas Lower to Middle Pliocene 375
1534 OSBORN: THE PROBOSCIDEA

Fam.: RUNOMASTODONTID^. Subfam.: Notorostrin^

Genus: Cordillerion
True Notorostrines typified by 'Madodon' andium. Mandible brevirostral. Su]:)erior tuslis si)iial or horizontal, with broad enamel
band. Inferior tusks wanting in adult stages; probably present in juvenile stages. INlolars persistently brachyodont with single trefoils,
paralleling Trilophodon; ridge-crests 4-4)^-5)2.

Cordillerion Osborn, 1926 Page

Cordillerion andium Cuvier, 1806, 1824, near volcano of Imbaburra, Quito, Ecuador Pleistocene 549
Cordillerion tarijensis Ameghino, 1902, Valley of Tarija, Bolivia Pleistocene 550
Cordillerion bolivianus Philippi, 1893, Ulloma, Bolivia Pleistocene 551

Cordillerion of Mexico
Cordillerion tropicus Cope, 1884, State of iSIichoacan, Valley of Mexico Pliocene or Pleistocene 553
Cordillerion oligobunis Cope, 1893, Tequixquiac, Valley of Mexico Pliocene or Pleistocene 554
Cordillerion oligobunis antiquissiynus Freudenberg, 1922, Valley of Amajaque, Hidalgo,
Mexico Pliocene 555
CordillerionC!) oligobunis intermedius Freudenberg, 1922, Mexico, exact locality not recorded. Pliocene or Pleistocene
. 557
Cordillerion oligobunis progressus Freudenberg, 1922, Canada de Aculcingo, Mexico Pliocene or Pleistocene 558
Cordillerion^!) oligobunis felicis Freudenberg, 1922, Puebla, Mexico Pliocene 556

Cordillerion of southwestern United States

Cordillerion edensis Osborn, 1922, Mt. Eden Hot Springs, San Bernardino County, California . . . Upper Pliocene 560
Cordillerion bensonensis Cidley, 1926, near Benson, Cochise County, Arizona, San Pedro
formation Upper Pliocene 565
Cordillerion gralum Hay, 1917, Brazos River, Pittbridge, Burleson County, Texas Pliocene 559
Cordillerion defloccalus Hay, 1926, Aransas River, San Patricio County, near Sinton, Texas. . .Pliocene(?) 564
Cordillerion orarius Hay, 1926, Aransas River, San Patricio County, near Sinton, Texas Pliocene(?) 562

Fam.: BUNOMASTODONTID.E. Subfam.: Brevirostrin.^

Genera: Pentalophodon, Anancus, Synconolophus
Mandible rapidly abbreviating,in correlation with loss of lower tusks; completely brevirostral in Upper Pliocene time. Grinding
teeth, originally short crowned, brachyodont, finally becoming subhypsodont with folded enamel, ptychodont; internal and external
cones progressively alternating and proverted central conules
;

Pentalophodon of India
Pentalophodon Falconer, 1857, 1865. Five-crested 'intermediate molars'
Pentalophodon sivalensis Cautley, 1836, vicinity of Nahun, southeast India, Upper Siwaliks(?). . . Upper Pliocene 650
Pentalophodon fakoneri sp. nov., Upper Siwaliks, Tatrot (?), 1,000 feet above Middle
Pliocene Dhok Pathan, India Upper Pliocene 653

Anancus of Eurasia
Anancus Aymard, 1855, 1859. Superior tusks straight, attaining great length. Four-crested intermediate molars
Anancus fakoneri Osborn, 1926, Red or Norwich Crag of Suffolk, England Uppermost Pliocene 636
Anancus arvernensis Croiz. & Job., 1828, Perrier, Auvergne, France, Villafi'anchian-Astian. . . .Upper Pliocene 632
Anancus arvernensis brevirostris Ciervais and de Serres, 1846, Montpellier, Ilerault, France.. . .Upper Pliocene 634
Anancus perimensis Falconer and Cautley, 1847, Perim Island, India = Dhok Pathan]
[ Middle Pliocene 643
Anancus minutoarvernensis Klahn, 1922, Herbolzheim, Baden, (lermany Pliocene 283
Anancus gigantarvernensis Kliihn, 1922, Herbolzheim, Baden, Germany Pliocene 283
Anancus intermedius Eichwald, 1831, Volhynia, Russia Pliocene 639
Anancus arvernensis progressor Khomenko, 1912, southern Bessarabia, Russia Pliocene 639
Anancus properimensis sp. nov., near Chinji Bungalow, Lower Chinji, 800 feet above
base of Lower Siwaliks Mio-Pliocene 647
Anancus sinensis Hopwood, 1935, Shansi, China PlioceneC?) 721

Synconolophus of India
Synconolophus Osborn, 1929. Warped, compacted conelets and conules, no trefoils
Synconolophus dhokpathanensis Osborn, 1929, Dhok Pathan, level of Hipparion punjabiense (?)
ref. quarry, 500 feet below top of Middle Siwaliks, India Middle Pliocene 661
Synconolophus hasnoti Pilgrim, 1913, near Bhimbar, India [=Dliok Pathan horizon] Middle Pliocene 659
Synconolophus corrugatus Pilgrim, 1913, Hasnot, India = Dh()k Pathan horizon]
[ Middle Pliocene 658
Synconolophus propathnnensis Osborn, 1929, near l^hok Pathan, India, Ilippitrion quarry level,
500 feet below top of Middle Siwaliks (Dhok Pathan horizon) Middle Pliocene 665
Synconolophus ptychodus Osborn, 1929, near Chinji Bungalow, India, Lower Chinji, 800 feet
above base of Lower Siwaliks Mio-Pliocone 657
 SUMMARY 1535

Fam. nov.: HUMBOLDTID.E. Subfam.: Humboldtin^

Genera: Cuvieronius, Eubelodon, Stegomastodon

Superior tusks without enamel. Molars with single, double, or quadruple trefoils; ridge-crests 4}^ to 7%; inferior ridge-crests pio-
verted superior ridge-crests transverse, centroverted, or retroverted. Central conules rudimentary or wanting.
;

All species below arranged in ascencUng order of the evolution of the grinding teeth. Geologic levels of South American species
uncertain.
North American Species
Eubelodon Barbour, Grinders broad, double trefoils
1914. Page
Eubelodon morrilU Barbour, 1914, Devil's Gulch B, Brown County, Nebraska Middle(?) Pliocene 602
South American Species
Cuvieronius Osborn, 1923. Grinders broad, double trefoils; single trefoils in primitive species
Cuvieronius humboldtii Cuvier, 1806 (in Desmarest, 1818), near Concepcion, Chile
[Ecuador?] Pleistocene 576
Cuvieronius platensis Ameghino, 1888, San Nicolas de los Arroyos, Province of Buenos Aires,
Argentina, Lower Pampeaii formation Pleistocene 579
Cuvieronius superbus Ameghino, 1888, Pergamino, Province of Buenos Aires, Argentina,
probably Superior Pampean Pleistocene 580
Cuvieronius bonaerensis Moreno, 1888, Arrecifes, Province of Buenos Aires, Argentina,
Lower Pampean Pleistocene 579
Cuvieronius rectus Ameghino, 1889, Ensenada, near La Plata, Argentina [Pleistocene] 580
Cuvieronius maderianus Ameghino, 1891, Puerto Madero in Buenos Aires, Argentina [Pleistocene] 581
Cuvieronius chilensis Philippi, 1893, near Lake Tagua-Tagua, Chile [Pleistocene] 581
Cuvieronius pirayuiensis Gez, 1915, Corrientes, Argentina [Pleistocene] 582
Cuvieronius brasiliensis Lund (in Lesson, 1842), valley of the Velhas River, Province of
Minas Gei'aes, Brazil, limestone cave [Pleistocene] 578
Cuvieronius postremus Spillmann, 1928-1931, Quebrada of Cachihuayco, near Alangasi,
Province of Pichincha, Ecuador Superior Pleistocene 585
Cuvieronius ayorse Spillmann, 1928-1931, Quebrada of Chalang, near Punin, Province of
Chimborazo, Ecuador Pleistocene 583

Stegomastodon of North America

Stegomastodon Grinders with quadruple to multiple
Pohlig, 1912. trefoils completely blocking the valleys; inferior ridge-crests
proverted; cones ptychodont, labyrinthodont, subhypsodont
Stegomastodon priestleyi Hay and Cook, 1930, near Frederick, Tillman County, Oklahoma. .[Pleistocene] . . 684
Stegomastodon primiiivus sp. nov., near Ainsworth, Nebraska Lower Pleistocene 726
Stegomastodon mirificus Leidy, 1858, "Loup Fork of Platte River," Nebraska [probably near
Seneca, Thomas County, Nebraska {fide Hay, 1924, p. 100) Hooker County? {fide Lugn
;

and Schultz,
1934, p. 372)] Upper Pliocene(?)i 669
Stegomastodon successor Cope, 1892, Blanco formation, Texas Upper Pliocene 671
Stegomastodon texanus Osborn, 1924, Llano Estacado, Texas, Blanco formation Upper Pliocene 673
Stegomastodo7i arizonx Gidley, 1924, 1926, Curtis Flats, Cochise County, Arizona,
San Pedro beds Uppermost Pliocene 678
Stegomastodon aftonise- Osborn, 1924, near Akron, Plymouth County, Iowa Lower Pleistocene^ 682
Incert^ sedis
Stegomastodon chapmani Hay, 1834, 1843, unrecorded locality in United States 669

Fam. nov.: SERRIDENTID^. Subfam.: Serridentin^

Genera: Serridentinus, Ocalientinus, Serbelodon, Trobelodon

Serrate-toothed mastodonts. Superior and inferior molars universally characterized by more or less perfectly developed and serrated
crests springingfrom the ectoconelets of the inferior molars, from the entoconelets of the superior molars. Generic divergence chiefly in
inferior incisors —
horizontal oval in Serridentinus, oval, greatly enlarged in Serbelodon, flattened into a pair of broad shovels in Platybelo-
don, abbreviated and rounded in Torynobelodon, and entirely wanting, with brevirostral mandible, in Notiomastodon.

Serridentines of Eurasia
Serridentinus Osborn, 1923
Serridentinus hasnotensis Osborn, 1929, near Hasnot, India, Upper Middle Siwaliks
(Dhok Pathan horizon) Middle Pliocene 452
Serridentinus metachinjiensis Osbom, 1929, near Chinji Bungalow, India, Lower Chinji,
800 feet above base of Lower Siwaliks (cf. S. productus) Mio-Pliocene 456

'[Lower to Middle Pleistocene (see footnote on page 669 of Volume I). — -Editor.)

'[Lower to Middle Pleistocene (see footnote on page 671 of Volume I). — Editor.)
1536 OSBORN: THE PROBOSCIDEA

Serridentinus prochinjiensis Osboni, 1929, near Chinji Bungalow, India, Lower Chinji, 600 Page
feet above base of Lower Siwaliks, small, primitive (of. .S. productus) Mio-Pliocene 457
Serridentinus chinjiensis Osborn, 1929, near Chinji Bungalow, India, Lower (Chinji, 700 feet
above base of Lower Siwaliks, sharp crested (cf. »S. serridens) Mio-Plioeene 456
Serridentinus browni Osborn, 1926, near Chinji Bungalow, India, Lower Chinji, 800 feet above
base of Lower Siwaliks, blunt crested (cf. S. productus) Mio-Pliocene 452
Serridentinus gobiensis Osborn and Granger, 1932, near Iren Dabasu, Inner Mongolia,
Tung Gur Upper Miocene 398
Serridentinus mongoliensis Osborn, 1924, Loh formation, Mongolia, blunt crested Lower to Middle Miocene 396

Serridentines of China and Japan

Serridentinus lydekkeri Schlosser, 1903 (1906), Red Clays, North China Lower Pliocene(?) 457
Serridentinus wimani Hopwood, 1935, Kansu, China Miocene(?) 732
Serridentinus annedens Matsumoto, 1924, Province of Mino, Japan, Hiramaki formation,
probably Burdigalian Lower Miocene 457

Serridentines of Europe
Serridentinus suhtapiroideus Schlesinger, 1917, Wies, near Eibiswald (Styria), Austria (Lower
Helvetian) Lower Middle Miocene 394
Serridentinus filholi Frick, 1926, 1933, Gers, France Lower Miocene 473

Serridentines of North and Central America

Sharp-crested Forms
Serridentinus serridens cimarronis Cope, 1893, east of Llano Estacado, Texas, Clarendon
formation, Procamelus-Hipparion zone Lower Pliocene 429
Serridentinus serridens Cope, 1884, Clarendon formation, Texas Lower Pliocene 423
Serridentinus anguirivalis Osborn, 1926, Snake Creek B, Sioux County, western Nebraska. . . .Lower Pliocene 425
Serridentinus brewsterensis Osborn, 1926, Brewster, Polk County, Florida, Bone Valley
formation Lower Pliocene 430

Blunt-crested Forms
Serridentinus guatemalensis Osborn, 1926, Chinautla, Guatemala Upper Pliocene 432
Serridentinus progressus Osborn, 1923, Driftwood Creek, Hitchcock County, Nebraska Pliocene(?) 401
Serridentinus (Ocalientinus?) nebrascensis Osborn, 1924, Snake Creek B, Sioux County,
Nebraska Lower Pliocene 473
Serridentinus productus Cope, 1875, Santa F6 marls. New Mexico (type) Upper Miocene 404
Clarendon formation, Texas (ref.) Lower Pliocene 404
Serridentinus proavus Cope, 1873, Pawnee Buttes, Pawnee Creek B, Weld County, Colorado. . Upper Miocene 403

Incert^ sedis
Serridentinus barstonis Frick, 1933, Mohave Desert, California, Barstow beds Mio-Pliocene 447

Ocalientinus of Mongolia
Ocalientinus Frick, 1933. mandible longirostral, deeply grooved above inferior tusks of 'prod-tusk' type
Trefoils florescent ; ;

Ocalientinus florescens Osborn, 1929, Kholobolchi Nor region, Khunuk formation, Mongolia.. .Pliocene 397

Ocalientinus of North America

Ocalientinus obliquidens Osborn, 1926, Charleston, South Carolina, phosphate beds Upper Pliocene, redeposited
in Lower Pleistocene 419
Ocalientinus bifoliatus Osborn, 1929, Brewster, Polk County, Florida, Alachua clays Lower Pliocene 415
Ocalientinus floridanus Leidy, 1886, near Williston, Levy County, Florida, Alachua clays Lower Pliocene 416
Ocalientinus floridanus leidii Frick, 1926, near Williston, Levy County, Florida,
Alachua clays Lower Pliocene 419
Ocalientinus republicanus Osborn, 1926, Republican River formation, northwestern Kansas. . .Lower Phocene 414
Ocalientinus ojocaliensis Frick, 1933, Ojo Calientc, New Mexico, Santa F6 marls (upper) Mio-Pliocene 435
Ocalientinus emmonsi Hay, 1930, marl beds of Halifax County, North Carolina Miocene 733

Serbelodon of North America

Serbelodon Frick, 1933, 'sub-shovel-tuskers.' Mandible massive, inferior tusks concavo-convex, increasingly massive, without dentinal
rod-cones
Serbelodon burnhami Osborn, 1933, near Ricardo, San Bernardino County, California Upper Pliocene 444
Serbelodon barbourensis Frick, 1933, near Ainsworth, Nebraska, Christmas quarry Lower Pliocene 443

Phyletic relationship uncertain

Serbelodoni?) precursor Cope, 1892, 1893, Mt. Blanco, Llano Estacado, Blanco formation,
Texas, sharp crested, trefoils simple, cement in valleys Upper Pliocene 431
 SUMMARY 1537

Trobelodon of North America

Trobelodon Frick, 1933, 'sub-shovel-tuskers.' [Inferior tusks moderately heavy, biconvex, pointed] Page
Trobelodon taoensis Frick, 1933, Santa Cruz, New Mexico, Santa Fe marls (rare) Mio-PIiocene 446

Fam. nov. : SERRIDENTID.E. Subfam.: Platybelodontin^

Genera: Platybelodon, Torynobelodon

Platybelodonts of Asia and North America

Platybelodon Borissiak, 1928, 1929, typical 'flat-tuskers.' Incisors solid with closely compacted rod-cones
Platybelodon danovi Borissiak, 1928, Kuban region, North Caucasus, Chokrak beds Upper Miocene 461
Platybelodon grangeri Osborn, 1929, Tairum Nor basin, MongoUa, Tung Gur horizon Upper Miocene 463

Torynobelodon Barbour, 1929, 'dredge-tuskers.' [Inferior tusks with dentinal rod-cones]

Torynobelodon loomisi Barbour, 1929, Sand Canyon, east of Indian Hill, vicinity of
Republican City, Harlan County, Nebraska Middle Pliocene 338
Torynobelodon barnumbrowni Barbour, 1931, Pliocene gravels on Snake River,
Cherry County, Nebraska Pliocene 470

Fam. nov.: SERRIDENTID.E. Subfam. nov.: Notiomastodontin^

Genus : Notiomastodon

Serridentines of South America

Notiomastodon Cabrera, 1929, [brevirostal mandible, tusks wanting).
Notiomastodon ornatus Cabrera, 1929, Monte Hermoso, Province of Buenos Aires,
Argentina Lower Pleistocene(?) 590, 731
Notiomastodon argentinvs Ameghino, 1888, Valley of Tarija, Argentina, exact locality
unrecorded Pliocene 550
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 OSBORN'S FINAL (1935)

CLASSIFICATION OF THE STEGODONTOIDEA AND ELEPHANTOIDEA

The and subfamihes of the Proboscidea first adopted by Osborn (1933) will be found on
classification of the superfamihcs, famihes,
pages 30 to 33 of Chapter II, Volume I, Memoir. Osborn's final (1935) classification of the Moeritherioidea, Deinotherioi-
of the present
dea, and Mastodontoidea immediately precedes this list, and, like the following list of the Stegodontoidea and Elephantoidea, embraces
the genera and species regarded by him as valid.

Superfamily: STEGODONTOIDEA
Fam.: STEGODONTID^ Subfam.: Stegodontin^
Genus : Stegodon

Phyhmi parallel to that of the true Archidiskodon and Elephas, not directly ancestral. Crania subelephantoid to extremely abbrevi-
ated (female?), to more elongated (male?), to triangular in form. Tusks attaining great dimensions. Molars generally more brachyodont
than those of the Elephantoidea; ridge-crests increasing from nine to fifteen and a half in the third lower molars; conelets multiplying
by binary fission to 20 +
In section, the valleys separating the adjacent ridges are found to be closed or V-shaped, whereas in the
.

elephantoid molars the valleys are open or U-shaped.

Stegodon Falconer and Cautley, 1847, 1857 Page

Stegodon insignis Falconer and Cautley, 1845, 1846, Siwalik Hills, India Lower Pleistocene to
Upper Pleistocene 836, 866, 874
Stegodon ganesa Falconer and Cautley, 1845, 1846, Siwalik Hills, India Lower Pleistocene to
Upper Pleistocene 836, 869, 874
Stegodon airdwana Martin, 1890, Alas-Tuwa, Trinil, Java, Kendeng-Schichten Middle(?) Pleistocene 836, 885
Stegodon ganesa javanicus = S. airdwana or S. trigonocephalus] Dubois, 1908, Trinil, Kendeng-
[

Schichten, Java Middle Pleistocene 836, 889

Stegodon orientialis shodoensis Matsumoto, 1924, island of Mitsugo (Mitsugo-shima) and island
of Shodo, Inland Sea, Japan Middle Pleistocene 836, 893
Stegodon (Archidiskodoni) mindanensis Naumann, 1890, Mindanao, Philippine Islands Lower(?) to Middle
Pleistocene 836, 892
Stegodon trigo7iocephalus Martin, 1887, probably vicinity of Surakarta, Java Lower(?) Pleistocene
(fide Matsumoto) 836, 890
Stegodon pinjorensis Osborn, 1929, near Siswan, India, upper levels of Pinjor horizon Lower Pleistocene 836, 883
Stegodon orientalis Owen, 1870, near Chungkingfoo, Szechuan, China Lower(?) Pleistocene 836, 884
Stegodon orientalis graiigeri Osborn, 1929, Yenchingkou, near Wanhsien, Province of Szechuan,
China Lower Pleistocene 836,875
Stegodon aurorse INIatsumoto, 1915, 1918, Mt. Tomuro, Kaga, Japan LTpper(?) Pliocene 836,892
Stegodon insignis birmanicus Osborn, 1929, Mingoon, I3urma, upper levels, Irrawaddy Series. . Upper Pliocene' 836,874
Stegodon bombifrons Falconer and Cautley, 1846, Siwalik Hills, India Middle Pliocene 836,863
Stegodon sinensis Owen, 1870, vicinity of Shanghai(?), China Lower(?) Pliocene 836,860
Stegodon elephantoides Clift, 1828, near Yenangyaung, Burma, Irrawaddy Series Lower Pliocene' 836,861
Stegodon elephantoides { = cliftii) Falconer and Cautley, 1846, near Yenangyaung, Burma, Ir-
rawaddy Series Lower Pliocene' 81 1, 814, 827
828, 831, 836

Not Determined by the Present Author

Stegodon bondolensis van der Maarel, 1932, Bondol, near Kuwung, Java 894
Stegodon trigonocephalus praecursor von Koenigswald, 1933, Bumiaju, Schichtcn of Kali Glagah, Java 896
Parastegodoni kwantoensis Tokunaga, 1934, Kakio, Kanagawa Prefecture, Japan 897
Stegodon yushensis Young, 1935, Ytishe, China, 'violet sands.' 897
Stegodon officinalis Hopwood, 1935, Szechuan (?), China. Horizon unknown 898
Stegodon zdanskyi Hopwood, 1935, China. Horizon unknown 899
Parastegodon sugiyamai Tokunaga, 1935, Iruhi, in Saida village, Shikoku, Japan 899
Parastegodon akashiensis Takai, 1936, Okubo-mura, Akashi-gun, Hyogo Prefecture, Japan 1420
Stegodon licenti Teilhard de Chardin and Trassaert, 1937, southeastern Shansi (Yiishe Basin), China 1420
Parastegodon infrequens Shikama, 1937, near Akasi, Japan 1420
Stegodon shodoensis akashiensis (Takai, 1936) Makiyama, 1938, Eigasima, near Akasi, and under the sea off Hayasi-zaki,
Japan 1420

'[See page 824 above where reasons are given for regarding these species as of Lower Pleistocene age. — Editor.]

1539
1540 OSBORN: THE PROBOSCIDEA

Superfamily: ELEPHANTOIDEA
Fam.: ELEPHANTID^ Subfam.: Mammontin^
Genera: Archidiskodon, Metarchidiskodon
Cranium brachycephalic, bathycephalic, hypsicephalic. Superior tusks large, incurved, crossing in old males. Primitive grinding
teeth, subloxodont, subhypsodont; ridge-plates extremely broad, enamel borders thickened, more or less crenulate, cement usually
thick.
Archidiskodonts of Eurasia
Archidiskodon Pohlig, 1885, 1888
Archidiskodon meridionalis cromerensis Dep^ret and Mayet, 1923, Kessingland, Suffolk, England Page
Cromerian or Norfolkian Lower Pleistocene 946, 980
Archidiskodon meridionalis Nesti, 1825, Val d'Arno supdrieure, northern Italy Upper Pliocene' to
Lower Pleistocene 946, 969
Archidiskodon planifrons Falconer and Cautley, 1845, Siwalik Hills, India. Pinjor horizon . . . Upper Pliocene and
Lower Pleistocene 946, 950
Archidiskodon planifrons rumatius Stefanescu, 1924, Tulucesti (Covurlui), Rumania LTpper Pliocene 968
Archidiskodonts of South Africa
Archidiskodon Pohhg, 1885, 1888
Archidiskodon broonii Osborn, 1928, The Bend, Vaal River, near Kimberley, South Africa. . . .Lower or Middle
Pleistocene 946, 989
Archidiskodon vanalpheni Dart, 1929, Sydney-on-Vaal, South Africa Lower(?) Pleistocene 990
Archidiskodon millelli Dart, 1929, Sydney-on-Vaal, South Africa Lower(?) Pleistocene 991
Archidiskodon loxodontoides Dart, 1929, Sydney-on-Vaal, South Africa Lower(?) Pleistocene 991
Archidiskodon yorki Dart, 1929, Vanasswegenshoek-Bloemheuvel, near Christiana, South
Africa Lower(?) Pleistocene 992
Archidiskodon subplanifrons Osborn, 1928, Sydney-on-Vaal, South Africa Upper(?) Pliocene
[Middle Pliocene] 946, 987
Archidiskodon proplanifrons Osborn, 1928, Gong-Gong, near Vaal River, South Africa ]Middle(?) Pliocene 986

Metarchidiskodon Osborn, 1934

Metarchidiskodon griqua Haughton, 1922, Griqualand West, South Africa Lower(?) Pleistocene 946, 994

Archidiskodonts of North America

Archidiskodon Pohlig, 1885, 1888
Archidiskodon imperator maibeni Barbour, 1925, near Curtis, Lincoln County, Nebraska Upper Pleistocene 946, 1027
Archidiskodon meridionalis nebrascensis Osborn, 1932, near Angus, Nuckolls County, Nebraska. Lower to Middle Pleistocene 1033 .

Archidiskodon imperalor Leidy, 1858, Platte River, Thomas County? (fide Hay, 1924), Hooker
County? (fide Lugn and Schultz, 1934), Nebra.ska Lower Pleistocene 946, 998
Archidiskodon exilis Stock and Furlong, 1928, Santa Rosa Island, California Pleistocene 946, 1031
Archidiskodon haroldcooki Hay, 1928, Frederick, Oklahoma Lower(?) Pleistocene 1029
Archidiskodon imperator scoiti Barbour, 1925, Staplehurst, Seward County, Nebraska Lower Pleistocene 946, 1025
Archidiskodon hayi Barbour, 1915, Crete, Saline County, Nebraska Lower Pleistocene 946, 1023
Archidiskodon imperator falconeri Freudenberg, 1922, Tequixquiac, Valley of Mexico Lower(?) Pleistocene 946, 1016
Archidiskodon imperator silrestris Freudenberg, 1922, Ejutla, State of Oaxaca, Mexico Lower(?) Pleistocene 946, 1015
Archidiskodon sonoriensis O.sborn, 1929, near Arizpe, northern Sonora, Mexico Lower Pleistocene 1033

Not Determined by the Present Author

Archidiskodon paramammonteus Matsumoto, 1939, Nagahama, Minato Town, Province of Kazusa, Japan 1420

Fam.: ELEPHANTID^ Subfam.: Mammontin.e

Genus : Parelephas
Cranium intermediate in form between Archidiskodon and Mammonlens; frontals concave; occipital crest elevated. Tusks with
remarkable incurvature. Molars with constantly increasing ridge formula in 3. M
Eurasiatic Species
Parelephas Osborn, 1924
Parelephas wiisti Pavlow, 1909, Tiraspol (gouv. Kherson), southern Russia Upper Pleistocene 1048, 1065
Parelephas intermedins Jourdan, 1861, near Lyons, Rhone Valley, France Pleistocene 1048, 1062
Parelephas armeniacus Falconer, 1857, near Khanoos, Province of Erzerum, Armenia Pleistocene 1048, 1060
Parelephas trogontherii Pohlig, 1885, 1888-1891, Siisscnborn, near Weimar, Germany Middle Pleistocene 1048, 1056
Parelephas{1) trogontherii nestii Pohlig, 1891, Forest Bed (Norfolk), Walton (Essex), and
Southwold (Suffolk), England Lower Pleistocene 1048, 1059
Parelephas trogontherioides Zuffardi, 1913, Piedmont, northern Italy Upper Pliocene' 1048, 1055

'[Possibly Lower Pleistocene (see footnote 1 on p. 1049). — Editor.]

 SUMMARY 1541

North American Species Pago

Parelephas prugressus Osborn, 1924, Zanesville, Muskingum County, Ohio Upper Pleistoceno 1048, 1097
Parelephas jejfersonii Osborn, 1922, Joiiesboro, Indiana Upper Pleistocene 1048, 1083
Parelephas roosevelti Hay [ = syn. Parelephas jejfersonii], Asliland, Cass County, Illinois Pleistocene 1048, 1095
Parelephas jacksoni Mather, 1838, Jackson County, Ohio Pleistocene 1048, 1068
Parelephas columbi cayennends Osborn, 1929, Cayenne, French Guiana, South America Upper(?) Pleistocene 1048, 1083
Parelephas floridanus Osborn, 1929, near Bradenton, Manatee County, Florida Upper(?) Pleistocene 1048, 1108
Parelephas colinnbi felicis Freudenberg, 1922, Tecamachalco, Puebla, Mexico Pleistocene 946, 1082
Parelephas columbi Falconer, 1857, 1863, 1868, Brunswick Canal, near Darien, Georgia Upper(?) Pleistocene 1048, 1071
Parelephas washingtonii Osborn, 1923, Pine Creek, Wliitman County, Washington Pleistocene 1048, 1101
Parelephasf eellsi Hay, 1926, Port Williams, Clallam County, Washington (?) 1104

Not Determined by the Present Author

Parelephas proximus uehataensis Shikama, 1937, Japan 1420

Fam.: ELEPHANTID^ Subfam.: Mammontin.e

Genus : Mammonteus
Cranium extremely acrocephalic, hypsicephalic, bathycephalic ; frontals concave; occipital crest greatly elevated ; occiput slight-
ly convex. Superior tusks greatly incurved, crossing in old age. Molar crowns broad, with extreme compression and very high ridge-
plate formula (M 3 io-al ^fi) » enamel borders more or less crimped or sinuous.

EuBASiATic Species
Mammonteum Camper, 1788; Mammonteus Osborn, 1924
Mammonteus primigenius Blumenbach, 1799, Europe and Asia Late or Upper Pleistocene 1138,1141
MammonieusC!) primigenius Leith-Adamsi PohHg, 1888, Thuringia, (iermany Upper-Lower Pleistocene 1137, 1150
Mammonteus primigenius hydruntinus Botti, 1891, Otranto, Italy Upper-Lower Pleistocene 1137, 1150
Mammonteus primigenius fraasi Dietrich, 1912, Steinheim, (iermany Upper-Lower Pleistocene 1137, 1152
Mammonteus primigenius astensis Dep^ret and Mayet, 1923, San Paolo de Villafranca, Italy. .Upper Pliocene' 1138, 1154

North American Species

Mammonteus primigenius compressjis Osborn, 1924, Rochester, Indiana Upper Pleistocene 1138, 1157
Mammonteus primigeniiis alaskensis sp. nov., vicinity of Fairbanks, Alaska Pleistocene 1159
Mammonteus primigenius americanus De Kay, 1842, near Rochester, New York Upper Pleistocene 1138, 1156

Fam.: ELEPHANTIDiE Subfam.: Loxodontin^

Genera: Loxodonta, Palxoloxodon, Hesperoloxodon
Cranium relatively primitive, platycephalic, brachycephalic (Loxodonta) somewhat more elevated or hypsicephalic (Palseoloxodon,
;

Hesperoloxodon); prominent parietofrontal crest {Palxoloxodon namadicus, P. melitensis, P. mnaidriensis). Premaxillary rostrum
broadened for insertion of widely divergent tusks; tusks relatively straight or slightly incurved. Molars moderately hypsodont, with
strong 'loxodont sinus' (Loxodonta), rudimentary or absent (Palseoloxodon, Hesperoloxodon).

Loxodonta F. Cuvier, 1825, 1827. Molar crowns low, broadly open 'loxodont sinus.'
Loxodonta prima Dart, 1929, Pilandsberg, Transvaal, South Africa [?Recent (fide Dart)] 1287
Loxodonta africana var. obliqua Dart, 1929, valley of Steelport River, northeastern Transvaal,
South Africa [?Recent (fide Dart)] 1287
Loxodonta subantiqua Haughton, 1932, Delport's Hope, near Vaal River, South Africa Level unknown Pleistocene — 1288
Loxodonta africana Blumenbach, 1797, probably from Cape region. South Africa Upper Pleistocene to Recent 1197
Loxodonta zulu Scott, 1907, Zululand, South Africa Pleistocene 1286
Loxodonta cornaliae Aradas, 1870, near monastery of Santa Chiara, Catania, Sicily [Quaternary or post-
Pliocene (fide Aradas)] 1204

Palseoloxodon (svn. Sivalikia) of India

Palxoloxodon Matsumoto, Cranium vnih
1924. broad rugose parietofrontal crest.
Palxoloxodon namadicus Falconer and Cautley, 1846, 1847, valley of the Nerbudda, India.. . .Upper Pleistocene 1211

'[Possibly Lower Pleistocene (see footnote 1 on p. 1049 above). — Editor.]

1542 OSBORN: THE PROBOSCIDEA

Pal^oloxodon (syn. Pilghimia) of the Mediterranean Islands Page

Palseoloxodon mnaidriensis Adams, 1870, Miiaidra Gap, island of Malta Pleistocene 1265
Palsenloxodon falconeri Busk, 1867, Zebbug Cave, island of Malta Pleistocene 1263
Palspoloxodun mclitenais Falconer, 1862, Zebbug Cave, island of Malta Pleistocene 1262
Palxoloiodon lamarmorae Forsyth Major, 1883, Quaternary sands of Morimentu near
Cionnesa, Sardinia Pleistocene 1266
Palseoloxodon Cypriotes Bate, 1903, Kcrynia Hills, island of Cyprus Pleistocene 1266
Palseoloxodon creticus Bate, 1907, near Cape Maleka, island of Crete Pleistocene 1267

Palseoloxodon of Africa
Palxoloxodon x/orki Dart, 1929, near Christiana, Vaal River, South Africa Middle(?) Pleistocene 1280
Palxoloiodon kuhni Dart, 1929, Pnicl Estate, South Africa Middle(?) Pleistocene 1281
Palseoloxodon irilmani Dart, 1929, below Christiana, Vaal River, South Africa Middle(?) Pleistocene 1280
Palseoloxodon{?) andreivsi Dart, 1929, Gong-Gong, Vaal River, South Africa Lower(?) Pleistocene 993, 1278
Palseoloxodon atlanljcits Pomel, 1879, near Mascara, Algeria, North Africa Pleistocene 1274
Palif'oloxodon jolensis Pomcl, 1895, Algerian scacoast. North Africa Pleistocene 1274
Palseoloxodon recki Dietrich, 1916, Scrcngetisteppe, Tanganyika Territory, East Africa Pleistocene 1275
Palseoloxodon Iransvaalensis Dart, 1927, near Bloomhof, Vaal River, South Africa Pleistocene 946, 993, 1284
Palseoloxodon .sheppardi Dart, 1927, near Bloemhof, Vaal River, South Africa Pleistocene 946, 993, 1285
Palseoloxodon arch idiskodontoides Haughton, 1932, Sydncy-on-Vaal Breakwater, bed of Vaal
River, South Africa Level unknown — Pleistocene 1282
Palseoloxodon hanekomi Dart, 1929, Delport's Hope, Vaal River, South Africa Level unkno^vn — Pleistocene 1279

Pal.eoloxodon of Japan
Palseoloxodon buski Matsumoto, 1927, Ninohe District, Province of Mutsu Post-Pleistocene to
Recent 1333
Palseoloxodon na)nadieus naumanni Makiyama, 1924, Sahamma, Totonii Province, Japan Middle(?) Pleistocene 1289, 1295
Palseoloxodon namadicus namadi Makiyama, 1924, dredged off island of Shodo, Sanuki
Province, Japan Middle(?) Pleistocene 1289, 1296
Palseoloxodon namadicus yabei Matsumoto, 1929, Inland Sea, Japan Middle Pleistocene
{fide Matsumoto) 1289, 1299
Palseoloxodon yokohamanus Tokunaga, 1934, Yokohama, Japan [Lower(?) Pleistocene
{fide Tokunaga)] 1301
Palseoloxodon prolomammonieus Matsumoto, 1924, 1926, Nagahama, Town of Minato,
Kimitsu District, Province of Kazusa, Japan Upper Pliocene(?),
Lower Pleistocene 1289, 1297
Palseoloxodon {1 Archidiskodon) tokunagai mut. junior Matsumoto, 1929, precise locality
unknown, Japan Upper PlioceneC?) or
Lower Pleistocene 1289, 1299
Palnoloxodon tokunagai Matsumoto, 1924, Soyama, Gokayama, Hiramura, Higashi-Tonami
District, Province of Etchu, Jai)an Upper Pliocene or
Lower Pleistocene 1289, 1298
(so recorded)
Pulnvloxodon prolomammonteus proximus Mat.sumoto, 1926, Kimitsu District, Province of
Kazusa, Japan Upper Pliocene(?) 1289, 1298

Palseoloxodon of Java
Palxoloxodon hysudrindicus Dubois, 1908, Kendeng-Schichten, .lava Middle(?) Pleistocene 1289, 1302

Not Determined by the Present Author

Parelephas prolomammonteus (Matsumoto) matsumotoi Saheki, 1931, Mishima, Kimitsu
District, Province of Kazusa, Japan [See Chap. XXI, \). 1416,
note inidcr matsumoloi\ 1300
Palaoloxodoii priscus var. ho.sei Chakravarti, 1935, Parkalta, near .lamniu, India 1418
Palifoloxodon aoniorienMs Tokunaga, 1936, Tenjiiibayashi, Aoniori Prefofture 1289, 1419
Palseoloxodon darti Cooke and Clark, 1939, Victoria Falls, northern Rhodesia, Africa 1420

Hesperoloxodon of Europe
Hesperoloxodon Osborn, 1931. Cranium domelike with flattened forehead. Premaxillaries extremely broad; tusks widely divergent,
slightly upcurved and incurvetl. ( Irinders hyjjsodoiit 'loxodont sinus' vestigial or alisent.
,

Hesperoloxodon antiquus italicus Osborn, 1931, Pignataro Interamna, near Cassino, Italy Upper Pleistocene 1245
Hesperoloxodon antiquus gcrmani.cus S. Stefanescu, 1924, Tanganu (Ilfov), Rumania U])p('r Phustocene 1233
Hesperoloxodon antiquus plnliirhnnehus Graclls, 1897, San Isidro del Campo, near Madrid,
Spain (?)L()\ver or Midtlle
Pleistocene 1231
Hesperoloxodon antiquus Falconer and Cautley, 1847, 1857, locality not recorded Lower(?) Pleistocene 1217
 SUMMARY 1543

Ilespcroloxodon untiquus nanus Acconci, 1880, near Monti Pisani, Cucigliana, Tuscany, Italy. Geologic age unknown 1230
Hespcroloxodon anliquus aitsonius Major, 1875, Verri, 1886, Dcperct and Mayet, 1923, San
Romano and San Paolo de Villafranca, Italy Upper Pliocene' 1232

Not Determined by the Present Author

E. [Elephas] antiqmis mut. rulhenensis Astre, 1937, Salles-la-Source (Avejrron,) France 1420

Fam.: ELEPHANTIDiE Subfam.: Elephantine

Genera: Elephas, Hypselephas, Platelephas

Elephas of India, Sumatra, and the Malay Peninsula

Cranium bathycephalic, cyrtocephalic, hypsicephalic, occipitofrontal dome more or less rounded; frontals gently concave.
Superior tusk.s relatively straight, incurved. Mandibular rostrum extremely abbreviate. Molars with finely plicated enamel; not
extremely hypsodont ridge-plate formula:
;
24
7-
M3 2 4-2

Elephas Linnaeus, 1735-1758

Elephas indicus Linnaeus, 1754, 1758,- Ceylon, India Upper Pleistocene and
Recent 1321, 1323
Elephas indicus ceylanicus de Blainville, 1845, Ceylon and mainland of India .Upper Pleistocene and
Recent 1327
Elephas indicus bengalensis de Blainville, 1845, chiefly Bengal, India Upper Pleistocene and
Recent 1327
Elephas indicus sumatranus Temminck, 1847, District of Palembang, Sumatra Upper Pleistocene and
Recent 1329
Elephas indicus hirsutus Lydekker, 1914, Negri Sembilan province, Malay Peninsula Upper Pleistocene and
Recent 1332
Hypselephas of India
Cranium hypsicephalic occiput elevated with broadly transverse frontal crest frontals deeply concave. Tusks relatively straight,
; ;

incurved, somewhat divergent at base. Molar crowns low; traceof median ioxodont sinus.' Ridge-plate formula: M3 ur^^T^^.

Hypselephas Osborn, 1934 {nornen nudum), 1936

Hypselephas hysudricus Falconer and Cautley, 1845, 1846, Siwalik Hills, near Siswan,
Chandigarh, Charnian, Kalka, as recorded by Barnum Brown Lower Pleistocene 1340

Platelephas of India
Cranium relatively elongate, dohchocephalic, platycephalic. Tusks unknown. Molars imperfectly known; ridge-plates relatively
low, directly transverse as in Elephas, no rudiment of a 'Ioxodont sinus.' Ridge-plate formula, as far as known 3 li^. : M
Platelephas Osborn, 1934 {nomen nudum), 1936
Platelephas platycephalus Osborn, 1929, near Siswan, Simla Hills, India Upper Pliocene or
Lower Pleistocene 1359

'[Possibly Lower Pleistocene, see footnote 1 on p. 1049 above. — Editor.)

-[Elephas maximum Linnajus, 1758, was adopted by the International Commission on Zoological Nomenclature at the meeting of the Fifth Inter-
national Congress of Zoology, held at Berlin in 1901. The present author, however, thought it preferable to retain Linnaeus' first designation, namely,
Elephas indicus, for the reasons given above on pages 1308-1311 of Chapter XX. — Editor.]
 Palaeomastodon intermedius Palaeomastodon intermedius
Amer. Mus. 13449 Paratype Amer. Mus. 14547 Type

Di upper m.3
external
mefacone
metaconid
metaconuleJ f

3rd crest

protoconid

hypocone
Natural size
hypoconid central conule
irrterna/ cin^u/um

Ani5898fc:Qst)

AM, 13431

An.l3437 AX^. 14547

as Kigs.
Fig. 1229. Four-coned ancestral grinders of the Proboscidea (e.g., ured with the six-coned PaUeomaslodon molius (sanu'
Marithtrium) corapar
93 and 94 of Volume I). All figures natural size.
thclu-xahun-
(Upper) Key to the hexabunodont upper and lower molar crowns of Palxoinmlodon intermedius. Observe the relatively broad proportions ;

odont crown.s of M'. namely, protocone, protocoinde, paracone, hypocone, raetacone, metaconule; the fundamental arrangement m two transverse cres *,
tnlophodont .rowii; both the
i.e.. protoloph, metaloph; the two intermediate cusps, i.e., protoconule, metaconule; the lower molar, M^, with rudimentary
any species Phiomia; a central conule rudiment in M.i; absence of median sulcus.
upper and lower molars wholly distinct in proportions from those of of

hexabunodont (D, E) teeth, M'-M.i, of the true Palxomastodon inlcrmedius, as compared with the
(Lower) Detailed studies of third superior and inferior

essentially tetrabunodont (A, B, C) Maritherium teeth.

.\, Maritherium lyonsi ref., right M''^ drawn from Mus. 15898.
cast Amer.
B, Marilhcrium trigodon ref., drawn from Amer. Mus. 13431.
right superior grinders
C, Maritherium andrewsi ref., left M2.3 drawn from Amer. Mus. 13437.
D, Palaomastodon intermedins paratype, third right superior molar, r.M', .4mer. Mus. 13449.
E, Palseomaslodon intermedius type, third left inferior molar, I.M3, Amer. Mus. 14547 (reversed in drawing).

1544
 2. EXPLANATION OF TERMS USED THROUGHOUT THE TEXT OF THE PRESENT MEMOIR
All proboscidean molars are derived from four-coned ancestral grinding teeth, such as are found in Moeritherium.
While the molar crowns in Palxommtodon are basically tetrabunodont, there being four principal cones in the
two well-developed crests of each tooth, the retention of intermediate conules in the upper molars makes these
teeth actually hexabunodont. In the last lower molar of this genus the development of a strong third crest leads
to a hexabunodont condition. The main external and internal bunoid cones never unite transversely into a com-
pleted ridge-crest in the Mastodontinae {Miomastodon, Pliomastodon, Mastodon), but remain separate or
bunolophodont, never becoming fully zygolophodont or j^oke crested, as in the Zygolophodontinae {Zygo-
lophodon, Turicius). In general the cones are arranged transversely to the long axis of the crowns, but in the
Brevirostrinse (e.g., Pentalophodon) there is an alternation of both the external and internal cones —a distinctive
character. There is also a slight alternation in the Humboldtine molar, Stegomastodon.

The numerical terminology of the superior (-loph) and inferior (-lophid) ridge-crests is as follows:

Pro-protoloph — id = One-half, anterior rudimentary ridge Pentaloph id— = Fifth ridge

Post-metaloph — id = One-half, posterior rudimentary ridge Hexaloph id — = Sixth ridge
Protoloph —id = First primary ridge = protocone and Heptaloph id— = Seventh ridge
paracone of Ungulata Octaloph id — = Eighth ridge
Metaloph — id = Second primary ridge = hypocone and Ennealoph id — = Ninth ridge
metacone of Ungulata Decaloph id — = Tenth ridge
Tritoloph — id = Third ridge Endecaloph id — = Eleventh ridge
Tetartoloph — id = Fourth ridge Dodecaloph id — = Twelfth ridge

Aristogenesis and Alloiometry. —A definition of aristogenesis (first known under the term "definite
variation," then as "rectigradation") will be found on page 1580 of the present chapter together with figure 1239.
See also page 1581 for figure and explanation of the term alloiometry.

—
Conules and Trefoils. In the Bunomastodontidae the very important coronal element or aristogene,
known as a conule, which successively appears between the proto- and metalophs, then between the meta- and
tritolophs, then between the trito- and tetartolophs, and so on, attaches itself to the external cones in the in-
ferior molars, forming on wear the outer trefoil or ectotrefoil, and to the internal cones of the superior molars,
forming the inner trefoil or entotrefoil. Molars of by the term bunomastodont.
this structure are characterized

Conules are highly characteristic of the Longirostrinse and Amebelodontinse and are more or less conspicuous in
the Tetralophodontinse and Notorostrinae as well as in the Palaeomastodontinae and in certain of the Brevirostrinse;
they are absent, however, in the molars of the Mceritheriinae (Moeritheres), the Deinotheriinae (Deinotheres), the
Mastodontinae, the Stegolophodontinse, the Stegodontinae (genus Htegodon), and the Elephantidae (Mammontinse,
Loxodontinse, and Elephantinae) ; vestigial or absent in the Zygolophodontinae and Humboldtinae and functional-
;

ly replaced in the Serridentidae (Serridentinae, Platybelodontinae, Notiomastodontinae) by lateral spurs which

gradually arise on the internal and external cones of the superior and inferior molars respectively, which, in turn,
subdivide into from two to six small conelets giving a serrated appearance to the spurs or crests, hence Serridentidae.
The absence in the Elephantoidea of conule development into trefoils, so characteristic of the mastodontoids,
and the early tendency to form evenly transverse, more or less mammillate crests, is one of the distinctive features
of this superfamily.

Conelets. —Another coronal element or aristogene is the conelet, which usually arises by binary fission,

rarely by ternary fission. For example, the primary or paired cones in the protoloph, referred to in the first

paragraph of this section, each splits into two making four (or 4+) conelets in the metaloph; each of these four
1545
 :

1546 OSBORN: THE PROBOSCIDEA

conelets tends to split into two, tending to form from five to eight conelets. But the binary fission is not so regular
as this. The newer anterior and posterior crests exhibit fewer conelets than the older mid-crests. To illustrate,

the maximum number of cones and conelets in each crest runs as follows
Molar cones and conelets: Primitive (Palxomastodon) 2-4-6-8-12-20+ progressive (Stegodon airdwana).

This conelet structure is also well illustrated in the Zygolophodon and Turicius molars (PI. in, pp. 134-135) in
which the conelets range from 4 to 25 in number. Professor Osborn in Chapter XIV of this volume, pages 809 to
812, has given a detailed explanation of this interesting transformation of the original crests or lophs by fission
into conelets.

AAa<iXyaTL SuZeu^ D o vbh he TrefoiLs

MecLtoTL ConzUes jSerrcite^ Spixrs

Fig. 1230. Molar diagrams showing typical crown pattern of Ms of each of the four families (geologic range, Oligocone to late Pleistocene) of the Masto-
dontoidea. All figures reduced to a uniform one-half scale.

Mastodontidak : A, Maslodon ameriainus ref. (Amer. Mus. 14294), from Fulton, Indiana. Ridge-crests 4K; distinct median sulcus. Plei.stocene.
See figure 134 B of the present Memoir.
Btjnomastodontid.e: B, Trilophodon pabnndicus type (Ind. Mus. A.426; cast Amer. Mus. 9905), from northern Punjab, India. Ridge-crests 4)^;

prominent central conules. Middle Miocene. See figure 211 of the pre.sent Memoir.
Ht'MH(ii.[)Tii).E: ref. (Buenos Aires Mus. 44; cast Amer. Mus. 277i)()).
C, CuvieroniuK supirbus Ri<lge-ereRts 4,!j4-, double trefoils; highly plicated.
Probably Upper Pleistocene. See figure 672 of the present Memoir.
SEnRiDKNTiD.E: D, SerriderdiTius produclus ref. (Amer Mus. 10582), from the Clarendon beds of Texas. Ridge-crests 4),i; serrated crests or spurs
springing from the ectoconelets. Lower Pliocene. See figure 369 C of the present Memoir.

Median Sulcus. — Also in dental structure the commissure of

median longitudinal sulcus (the fissure or

Hays) plays an important part (Fig. 1230, also PI. p. 134) it is distinctive of the molars of the Mastodontinae
i, ;

(e.g., Miomastodon, Pliomastodon, Mastodon), rapidly disappearing in the Zygolophodon tinjp (Zi/gnlnphndon

and Turicius), and persistent in the anterior ridge-crests only of the Stegolo])ii()(lontinir {Slegolupliodon).
This sulcus separates the external and internal cones both in the superior and inferior molars, further demonstrat-
ing that the ancestral proboscidean molar was tetrabunodont, as in Ma-ritherium, and not hexabunodont, as in
Palxomastodon ; in fact, the presence of proto- and metaconules blocking the median sulcus in the Palseomastodon
 SUMMARY 1547

molar forbids its direct ancestry to Mastodon in which the valleys between the crests are uninterrupted by either
conules or trefoils.

—
Ganometry. The ganometric method, from "ganos" signifying enamel, is, in brief, the measuring of the
combined enamel foldings of the grinding teeth, which, when drawn out of their closely plicated arrangement for
the finer comminution of the herbage on which these animals subsisted, steadily increase in length from Upper
Pliocene time when Eoa?ithropus lived to closing Pleistocene time when the late Cro-Magnon man lived.
In Professor Osborn's experimentation with the elephantine molars, he was ably assisted by Dr. Edwin H.
Colbert, who was measurements were approximate; that out of the thirty-six teeth
careful to state that the
studied, only two were unworn consequently that there were large estimated factors in most of the measurements
;

(cf. Osborn, with Colbert, 1931.858, p. 191). The accompanying figure (Fig. 1231) is inserted here to further dem-
onstrate this method as a possible contributory factor in "estimating the duration of the sub-divisions of the age of
man which have hitherto been dated chiefly by geologists calculating the length of the four glacial and three
interglacial epochs," and, according to Professor Osborn's opinion, in ascertaining "just how long it took to
produce a centimeter of enamel length" (op. cit., p. 188).
Cnomel length 1113 mm
thickness 5 mm
Fig. 1231. (Upper figure.s) Accelerated elephantine ridge-plates in No of folds lott
Cnomel Lenslh 6420 mm
thickness 55mm
Intensely accelerated evolution of the ridge-plates from the Quenije height 53 mm
Archidiskodon. Noof folds, 13
Ouerage height 234mm
Archidiskodon planifrotis of southern Eurasia into the Archidiskodon, imperator
of the United States and Mexico, all occurring from Upper Pliocene to Middle
Pleistocene time. Length of enamel foldings:

Archidiskodon imperator of Te.xas =8420 mm.

Archidiskodon planifrons of India =2204
Archidiskodon planifrons of India =1113

In these southern mammoths of Africa, Eurasia, and North America,

gigantic sizeis attained in a relatively short period of geologic time.

(Lower figure) Slegodon grangeri of the Pliocene of Szechuan, China, type

l.M^ with 11)2 ridge-crests (Amer. Mus. 18714):

Enamel length 740 mm.

Enamel thickness 7.5
Average height of ridge-crests 31 tnomel lenglh 2 204 mm

Ridge-crests in 100 mm. 3 3^ thickness 5 mm

Nq of folds 13
Queraae heisht 65mm
This figure illustrates one of the important differences between the
stegodontoid and elephantoid molars. As will be observed, the valleys
separating the adjacent molar ridges are closed or V-shaped at the bottom in Upper niiddie
the Stegodontoids, whereas in the Elephantoids they are U-shaped. Drawing
modified after Osborn, 1934.926, fig. 4.
Pliocene J leistocene

Dental Formula. —The canines are lacking in the Proboscidea, excepting in Mxritherium, in which the
third superior canine is pre.sent but greatly reduced nor are there any first deciduous premolars. The deciduous
;

premolars (or deciduous molars as they are frequently called) consist of Dp 2, Dp 3, Dp 4. In Moeritherium and
Palxomastodon P 2-4 are present; in Deinotherium, Phiomia, Serridentinus, Ocalientinus, and Platybelodon P 3-4
persist; in Trilophodon, Pliomastodon, Miomastodon, and possibly Blickotherium, P 4 only persists. In Mastodon
the fourth true superior and inferior premolars, P*, P4, form in the jaw but do not erupt;
in Elephas they are
suppressed entirely, and, with the exception of Archidiskodon (of the Mammontinse), there is also a complete
suppression of this premolar in the Mammoths. Incisors present in Moeritherium, but I 2 enlarged. Only I 2
persisting in other proboscideans as tusks (cf. pp. 1550-1552 below).

—
Ridge Formula. In certain of the Mastodontidse and in all of the Elephantidae the grinding teeth increase
the number of their ridges by adding crests especially by the addition of crests posteriorly, for example, post-
protolophid, post-metalophid, etc. The intermediate molars (Dp 4, 1, 2) vary in the number of their ridges M M
 :

1548 OSBORN: THE PROBOSCIDEA

from 3 to 5 in the Mastodontoidea (e.g., 3 in Trilophodon, 4 in Tetralophodon, and 5 in Pentalophodon). The

count is much liigher and apparently less uniform in the Stegodontoidea and Elephantoidea. In the Masto-
dontoidea the number of ridge-crests in the third molar (M 3) reaches 8'?; in the Tetralophodontina^ (genus
Morrillia); in the Stegodontoidea 15K {titegodon airdwana). In the Elephantoidea the number of ridge-plates

reaches fh {Mammonteus), f| (Parelephas). It is noteworthy that in Parelephas the ridge-plate count apparently
is higher in the superior than in the inferior molars —a marked distinction from the ridge formula of other

members of the Proboscidea, in which the inferior ridges exceed the superior ridges in number. If this character
should prove to be constant, it would aid greatly in the determination of molars of the genus Parelephas.

DoLiCHODONTY, Hypsodonty, AND Brachyodonty. —Other terms illustrating the great variability in the

molars of the Proboscidea are as follows

Dolichodoiity, referring to the long and narrow proportions of the molar crowns.
Hypsodonty, descriptive of the elevation or heightening of the crowns.
Brachyodonty, which indicates the broad, short, low-crowned character of the molar crowns.
Mas. La Plata
IM^ £x ternat 8-19
r.Mz External Paris Museum.

/ n ternat - Retrouers ion

Ref Cuvieronlus platen-sis Amegh

Univ. loiua Mus. 213

J.M5 Ex ten or
Cast Am.. Mas. 2573

Brit. Alus. 2 3 4 5
Internal -Proi/ersLon M.2857 internal - Centrover.5ion
Anancus perimensis Fate ana Caul. Stecfo mastodon afion.iae Osbcrn
Ref. "^P^

Brevirostrin*: Proversion OF Ridge-crests in Ananccs. FourConules Humboldtin/e: Retroversion and Ckntroversion of the Superior
ox Each Crest (1,2, 3. 4); Central Conules (CCC) between the Crests Ridge-crests in Cuvikroni0s and Stegomastodon
I'ig. 1232. Top: Anaiicus bremroslris ref. Third right inferior molar, Fii^. 1233. Top: Intermil retroversion in o + ridge-orested third left su-

r.Ms, .strong internal proversion of .5}2 ridge-crests. perior molar of Cuvieronius platensis.

Bottom: Anaiicuii pcriinensis ref. Third left superior molar, l.M'', Bottom: Internal centroversion in 7/2 ridge-crested third left superior
internal prover.sion of .)'2 ridge-crest.'i. molar of Stegomastodon afloni.f.

Chcerodonty and Ptychodonty. — Molar crowns covered with tubercles (choerodonty), and plication or
infolding of the enamel bordei-s with grooving of the sides of the molars (ptychodonty) are variable characters of
the proboscidean molar. The development of hypsodonty and chcerodonty among the mastodontoid (longirostrine
and brevirostrine) browsers is analogous to that in the hippopotami and the hypsodont suillines. The elephantoid
molars become hypsodont and polylophodont in the highest degree, in adaptation chiefly to grazing habits. The
acme of hypsodonty is represented in the Mastodontoidea by the Morrillia molar, and in the Elephantoidea by the
Mammonteus molar. Synconolophus, a brevirostral mastodont (see Fig. 623), and Stegomastodon, an humboldtine
mastodont (Fig. 645), present the most labyrinthine crown pattern in the entire range of proboscidean molars.
 :

SUMMARY 1549

Proversion, Retroversion, and Centro version. —The following descriptive terms are used in the present
Memoir in connection with the molars of certain of the Brevirostrines and Humbold tines (see Figs. 1232, 1233)
Ridge-crests extending obliquely from external to internal faces, toward anterior end of molars = Proversion
Ridge-crests extending jbliquely from external to internal faces, toward posterior end of molars = Retroversion
Ridge-crests extending obliquely from external to internal faces, toward center of molars = Centroversion
LoxoDONT Sinus. —A mesial expansion of the ridge-plates of the grinding teeth into a broad lozenge-shaped
sinus or cavity is especially characteristic of the genus Loxodonta (Fig. 1234) ; it is rudimentary or vestigial in
Palxoloxodon and Hesperoloxodon; there is a trace of it in the molars of primitive species of Archidiskodon, also in
Elephas hysudricus; in Metarchidiskodon there are prominent post-sinus folds instead of the median sinus expan-
sion of the primitive Archidiskodon.

Cement. —The Proboscidea, as a whole, are progressive, consequently cement is present more or less abund-
antly in the molars of all the Elephantoidea, even in some of the most primitive species, as, for example, Archi-
diskodon subplanifrons. In the Mastodontoidea, however, this is not the case. There are indications of cement in
the molars of such genera as Pentalophodon and Eubelo-
fy- «8- Fig. 89,
don, also it is present in the more progressive stages, e.g.,

J' J^S^\ Cordillerion, Morrillia, Stegomastodon, and Synconolo-

phus, especially the last-mentioned genus, the molars of
which are heavily cemented.

Cement Dentine Enamel

5 6

RM3 McQregor Mus. 5920

African Elephant. \ nat. size. Asiatic Elephant.

Type. Archidiskodon subplanifrons. Osbom. I928
' ' Drawn 6y Df.Uv&tt Bradltif

Fig. 1234. Crown view of the third ijil'erior molar of the right side of; Fig. I''ig. 1235. Section of third right inferior molar, r.M.3, of Archidiskodon
88, Loxodonta africana; Fig. 89, Elephas indicus, one-third natural .size. subplanifrons type (McGregor Mus. 3920), with si.K ridge-crests. From
Compare Owen, "Hi.story of British Fossil Mammals, and Birds," 1846, pp. Kimberley, South Africa. Drawn by Miss D. F. Levett Bradley, and repro-
230-232, figs. 88, 89. duced herewith slightly more than two-thirds natural size.

Cranium. —In a comparison of the crania of the Proboscidea it will be observed that there is great variability,
for example, the elongate (dolichocephalic), low cranium of the mastodonts as contrasted with that of the ele-
phants, which is vertically high (hypsicephalic), peaked (acrocephalic), deep (bathy cephalic), with great fore-and-
aft compression (cyrtocephalic). In one respect, however, there is uniformity, for the skull bones surrounding
the brain-case of but the most primitive proboscideans are strongly cancellate to afford broad attachments for
all

the heavy muscles necessary to support the weight of the tusks and of the trunk. As noted by Dr. C. W. Andrews
(see p. 102 of Vol. I) : "In the true Elephants and Mastodons the peculiar form of the skull is mainly due to the
enormous development of cellular bone in the occipital region." With age there is apparently a shortening and
1550 OSBORN: THE PROBOSCIDEA

deepening of the cranium in all species of elephants. In certain genera (e.g., Phiomia) the facial region lengthens
more rapidly than the cranial (that is, it is dolichopic), in contrast to certain other genera (e.g., Mceritherium) in

which the facial region shortens more rapidly than the cranial (that is, it is brachyopic).

Tusks. —The Proboscidea are distinguished by a greatly enlarged pair of cutting teeth or incisors, which
were originally opposed as feeding organs; this hypertrophy of P, I2, through feeding function, resulted in the
loss of both the first and third pairs of incisors; in the Moeritheres only does the first pair, P, Ii, persist, the

third superior incisor being greatly reduced, the inferior wanting. In the Deinotheres the upper incisors, I-, are
entirely aborted; the inferior pair (probably I.) is downturned and retroverted. In many of the Mastodontoidea
and in all of the Stegodontoidea and Elephantoidea the lower tusks (P) are greatly reduced or completely
suppressed.

Of the Mastodontoidea the most extraordinary tusks are those of: (1) The Amebelodonts, described as
'shovel-tusks' because of the high degree of specialization of the lower tusks for shoveling; these tusks with round-

ed tips are composed of concentric dentinal laminae. (2) The Platybelodonts or 'flat-tuskers,' with broad, flat,

square-tipped tusks, composed of dentinal rod-cones instead of dentinal laminae as in the Amebelodonts, and in
form like a coal shovel. (3) The Torynobelodonts or 'dredge-tuskers,' with broad, flat, chisei-Uke tusks, also

Chief Head and Dental Forms of four of the Superfamilies (I-IV) of the Proboscidea
Fig. 1236. (1) Ma?ritherioidea, one-twelfth natural size. (2) Deinotherioidea, (3) Mastodontoidea, and (4) Elephantoidea, all one-fiftieth natural size.

composed of dentinal rod-cones. (4) The Andean mastodonts {Cordillerion andium) with long spirally-

twisted tusks in which the enamel band follows the curved line of torsion. The Notorostrines, the Humboldtines,
and the Brevirostrines were without inferior tusks. In the true Mastodon {M. americanus) the presence or
absence of the inferior tusks varied with the individual.

The famous Stegodon ganesa upper tusks are striking not only because of their great length and symmetry,
but also because of the apparently impossible position in the jaw which left insufficient space for the pendant
trunk. This is a moot question, however, and is fully treated in the present volume, p. 857, caption to figure 733.

Mammontinae, the upper tusks attain gigantic size, e.g., the

Finally, in the Elephantoidea, especially in the
Archidiskodon imperator tusk from near Post, Texas, measuring 16-|- feet in length, also the example formerly in the
Geological Survey of Mexico with an estimated length of 13-(- feet. In certain of the Mammontines (i.e.,
Mammonteus primigenius) the tusks cross in old age producing a striking wheel-like appearance. The very early
loss of the lower tusks is one of the prime distinctions in the Elephantoidea and Stegodontoidea.
 SUMMARY 1551

Tusk Enamel. —A dental chararteristic of all primitive bunomastodonts is the presence of an enamel band
on the superior tusks as opposed to tlie elongate, flattened inferior tusks which generally lack the enamel band.

This betokens the primitive functional use of the upper and lower tusks in opposition, correlated with a grinding

Trilophodon
iLilli

T^hynchotheriuTn
tlascaldJe

All '/a^f. -natural size

Divergent Adaptive Radiation ok Crania and Incisive Tusks in Six Bunomastodont Subfamilies. Scale uniform
Fig. 1237. Comparative outline illustrations, side view, to same scale, of the completely restored skulls, jaws, and tusks, in descending order, of:

Trilophodon (Megabelodon) lullf Rhynchotherimn tlascahe, restored Stegomaslodon aTii,ons

Trilophodon (Genomaslodon) unllisloni, juvenile Observe persistent enamel band on inferior Anancus arvernensis
Phiomia minor tusks, also apposition of and superior
inferior Cuvieroniiis humboldtii, juvenile
Serridentinus prodiiclus tusks. In contrast with superior enamel band C'ordillcrion anttiti m
Tetralophodon grandincisivus only in Trilophodon, Phiomia, Srrridentinus, and
Tetralophodon campester Tetralophodon, and absence of enamel band on
inferior tusks.

(Osborn, 1934) Serridentinus belongs in the new family Serridentidic; Stegomaslodon and Cuvieionius in the new family Humboldtidffi

'The type mandible of Trilophodon lulli, originally restored by Barbour in 1914 with a pair of slender inferior incisive tusks (Fig. 244 of present Memoir),
is now found to be tuskless. The rostrum is expanded at the tips. See Barbour, 1934.2.
1552 OSBORN: THE PROBOSCIDEA

and triturating molar action rather than the chopping motion characteristic of the true mastodon ts. The important
functional distinction of the Mastodontoidea is that for a very long period of time the upper tusks abraded the
outer side of the lower tusks, which probably explains the retention of the superior enamel band (Mastodontinae
fide Schlesinger, and Notorostrinse, mde Cordillerion andium). The Rhynchorostrines are an exception, in that
the enamel band persists on the inferior tusks (see Fig. 1237) . In all the mastodonts related to the classic Mastodon
[Trilophodon] angustidens of Cuvier and the M. [Tetralophodon] longirostris of Kaup, as well as in certain of the
Serridentines, the superior incisors are extremely formidable as weapons, sharp, down- and out-turned, the
dentine or ivory being strengthened with a lateral enamel band. There seems to be no trace of an enamel band
on the Stegodontoid tusks, and those of the Elephantoidea are devoid of enamel, except at the tips in the young
stage.

Summary of Proportional Changes

Cyrtocephaly Fore-and-aft faciocranial abbreviation.
: Dolichocephaly: Lengthening of the cranium in proportion to breadth.
Cyrtodonty: Fore-and-aft molar-crown abbrcv^iation. Brachycephaly: Broadening of the occiput or of the zygomatic arches.
Hypsicephaly: Vertical heighteningof the cranium and jaws. Cyptocephaly Downward flexure of the facial to the basicranial a.xes.
:

Hypsodonty: Vertical heightening of the molar crowns. Orthoccphaly: Lackof inclination of the basifacial to the basicranial axis.
Brachyodonty: Vertical shallowness of the molar crowns. Dolichopy: Lengthening of the face.
Acrocephaly: Vertical heightening of the occipitofrontal apex. Brachyopy: Shortening of the face.
Bathycephaly Vertical deepening of the basicranium, molar alveoli, and jaws.
:

3. CHARACTERS, AFFINITIES, AND MIGRATIONS OF THE PROBOSCIDEA

MOERITHERIOIDEA
Superfamily: MCERITHERIOIDEA Osborn, 1921. Family: MffiRITHERI I D^ Andrews, 1906
Subfamily: Mceritheriin^ Winge-Osborn (1906-1923)
Genus: Moeritherium Andrews, 1901-1906
(Cf. Vol. I, Chap. Ill, and Pis. x, xi, and Figs. 23 and 24; also Vol. II, p. 1529, and Figs. 1220, 1227, and
PI. XIV)

The Mceritheres, named from Lake Moeris of the Greeks. Small amphibious quadrupeds of the
North African rivers and lakes. Cranium primitive, elongated (dolichocephalic), tubular brain cavity,
zygomatic arches slender. Face abbreviated (brachyopic) orbits very small, shallow, opening upward
;

and outward. Eyes very far forward, well raised toward top of face. Auditory meatus elevated, an
aquatic adaptation. Mandible and symphysis relatively short. Upper and lower lips opposing each other.
First superior and inferior incisors partly functional; second greatly enlarged, directly opposed, tusk-
like, curved, gliriform, partly sheathed in enamel; third superior incisors and superior canines greatly
reduced; third inferior incisors and inferior canines entirely wanting. Superior premolars, P''\ tri-
tubercular, tetartocones rudimentary or absent; last premolar not bilophodont. Molars tetrabunodont
(quadritubercular), bilophodont, with incipient trilophodon ty, quadrate or slightly elongate in pro-
portion; superior molars, M", strictly bilophodont, M" with rudimentary third crest, M^ with enlarged
third crest; inferior molars. Mi, bilophodont, M2, trilophodont, rudimentary, M3, trilophodont, trilopho-
—
donty not pronounced all three pairs of premolars and molars functioning at the same time. No trace of
trefoil pattern or of intermediate conules. Skeleton partly known vertebral structure indicating an ;

ambulatory and amphibious habit; probably pentadactyl. Palustral, amphibious, and semi-aquatic,
with ancestral genetic affinities to the Proboscidea in dentition, and analogies to the Sirenia and
Hyracoidea in the skull, but much closer to the Sirenia than to the Hyracoidea.
Dental formula: i;;^r| Ci P f^ M {ff
Ridge-crest formula: P 21 P
41^ 31It P2|^ 3^M M M
Horizon. — Upper Eocene (Qasr-el-Sagha formation) to Lower Oligocene (Fluvio-marine formation).
Compare p. 1529, also figures 23, 24, and 1227.
 /

DE-INOThERlUM HOPiVOODI. l-k£ i-icTOm-), Africa

PEINOTHERIUM CIG Ar\:TISSI MU M. Rumania

DLINOTHERIUM GIGA\'TE Uf'h OERyiAN^-

DEINOTHERIUM HAVAKICUM. bjii^aria

DEINOTHERIUM hung ARICUM, mumsary

<^^ MOERITHERIUM ANDREWS!, fayum
<^^ MOERtTHERtUM TRIGODON. F^yOr-i

^S^ MOERITHERIUM LYONS I. fayum DEINOTHERIUM HOBLEYI. east Africa

^^^-1^ mo'eritheriui^i gracile. f^yOm

^t-A mo~eritherium ancestrale. FAYOt^

30* 45* 60' 75' 9Q'E t05' 120" 135' ISO' 165' ISO" 165' J50' Ij5' 120" r05' 90'W 15' 60' PF Levrft BraJJei) I9M

Ongla, Ailgration and EvoLation of

oMoerittierium. ^Deinotheriam. Osborn 1955
PLATE XIV
Geologic range: Mxritherium, Upper Eocene to Lower Oligocene. Deinotherium, Lower Miocene to Middle Pleistocene.
 "

SUMMARY 1553

Habits and Affinities. — (Cf. Vol. I, p. 48): "The conclusions drawn from the front teeth, P- I2, from the
simple bunodont grinding teeth, from the very short face (brachyopy), from the long cranium (dolichocrany), and
from the extremely small bony eye sockets, are that Moeritherium was a confirmed and continual river-living
animal, feeding mainly under water and along the banks of rivers, more specialized for aquatic life than the
hippopotamus, as proven by its feeble pelvic bones, but far less specialized for aquatic life in its limb structure
than the Sirenia. This does not prove that Moeritherium is of the order Sirenia, as Osborn suggested in 1909 ; it is

certainly an independent member of the Proboscidea, as Andrews originally maintained and as Matsumoto has
stoutly contended. Its cranial analogies are with the sirenian skull ; its cranial and dental homologies are with the
Proboscidea. Its limbs and skeleton relate it to the primitive Proboscidea."

{Op. cii., "Recent studies by Gregory (1910-1920), Matsumoto (1923), and Simpson (1932) point
p. 39):

towards the existence in Upper Cretaceous and Lower Eocene time in Africa of a common ancestral form of
mammal which by adaptive radiation through ground-loving, shore-loving, and water-loving habits, may have
given rise to the Sirenians . . ., the Mceritheres, and the Proboscideans. Widely different and profoundly
divergent as the two great orders of Sirenians and Proboscideans are today, they still exhibit certain common
characters in their internal anatomy, certain common characters in their cranial and labial structure, as well as

one unique character in their grinding teeth, namely, trilophodonhj or the evolution of three crests on the superior
.

an d in f erior molars

Professor Osborn considered that the structure of the muzzle proves that Maeritherium had heavy and
fleshy lips capable of closing over the tusks when the mouth was shut; that it had no proboscis.

Migration. — So far as known at the present time Moeritherium was confined to the African continent. The
Moeritherium sp. from Baluchistan, described by Pilgrim in 1912 (see Vol. I, p. 78), is provisionally referred to
Trilophodon pandionis.

DEINOTHERIOIDEA
Superfamily: DEINOTHERIOIDEA Osborn, 1921. Family: Curtognathidaj Kaup-Osborn (1833-1936)
Subfamily: Dinotheriin^ Osborn, 1910
Genus: Deinotherium Kaup, 1829
(Cf. Vol. I, Chap. IV, also Fig. 56, and Pis. x, xi; Vol. II, p. 1529 and Figs. 1220-1222, 1227, also PI. xiv)

The Deinotheres, implying proboscideans of terrifying size, existed in Europe and Asia in Miocene and
Pliocene times, also in Africa in Middle Pleistocene time.

Cranium abbreviate; proportions brachycephalic ; summit flattened; occiput forwardly inclined.

Probably long proboscis capable of reaching higher branches of trees as well as the ground. Lower
jaw elongated (see Kittl's description on p. 99 of Vol. I, this Memoir, although on p. 112 it is describ-
ed as abbreviate) and sharply bent downwards. Superior tusks early aborted inferior tusks rounded, ;

directly downcurved, and bent backwards. Primitive Deinotheres present simple, bilophodont grinders,
similar to those of Moeritherium, and are progressively trilophodont the upper grinders attain a stage;

which parallels the molar pattern of the tapir, but show a tendency to the trilophodont structure char-
acteristic of the primitive mastodonts and elephants. As compared with other proboscideans, both
mastodon toid and elephantoid, the dentition of the Deinotheres is relatively non-progressive; the
fundamental pattern of the grinding teeth was established extremely early in geologic time, certainly dur-
ing the unknown Oligocene stages, because in the Lower Miocene specific stages it is fully established, es-
pecially the number of ridges and the character of the crests of the upper and lower true molars,
M1-M3. Grinding teeth sharply crested, persistently brachyodont, never hypsodont. Subdivision of
 1554 OSBORN: THE PROBOSCIDEA

summit of crests into fourteen to sixteen conelets {Deinotherium hopwoodi^), a character not previously
mentioned, so far as known, in connection with any other species of Deinotherium. Replacement of
Dp 3-4 by P 3-4; no premolar replacement of Dp 2. Dp 4 and 1 trilophodont; P 4, 2, and M
3 M M
bilophodont. All premolars and molars of both jaws functioning at same time. Pre- and i)ostcingulum
sometimes present. The ivory of the tusks "presents the fine concentric structure of those of the Hippo-
potamus, not the decussating curvilinear character which characterizes the ivory of the Eleohant .t and
u i
Mastodon" {fide Owen, 1868, III, p. 359).

% 4.

Fig. 1238.Juvenile jaw of Deinotherium giganteum, after Lartet, 1859,

PI. .XIII, fig. 4, one-third natural size,
from the Middle Miocene, Ile-en-Dodon
(Haute-Garonne), France. This jaw shows tlie replacement molars, as follows:
A and B = P3 and P4
D =M->
a-c = Dp>, Dps, Dpi, C = Ml
PI. XIII, fiij. 3, fragment of another jaw, one-third natural size, also
from Middle Miocene, Ile-en-Dodon (Haute-Garonne), France:
c = Dp, being replaced by B = P4: C = Mi [H. F. O.l

Limbs elongated, increasingly elephantoid, raising body well off the ground. Progressive increase
from Lower Miocene stage to Pliocene stage. Digits II, III, IV; vestigial D.I in the pes. Dorso-
in size
lumbar vertebrae abbreviate.
Dental formula: Di Itl Dc § I tl^l Dp =:| P ^:| M ;;|.
Ridge-crest formula: Dp 2* Dp 3f Dp 41 P 31 P 41 M If M 21 M 3*.
Horizon.— Lower Miocene to Middle Pleistocene. Compare Vol. I, p. 735, and figure 56; also Vol.
II, p. 1529 and figures 1220, 1222, 1227.

Habits and Affinities.— According to Weinsheimer (1883—see Vol. I, pp. 86-95), various authors during
the long period of excavation and description of Deinotherium remains compared them to fossils (mastodonts)
from the Ohio River, from Peru in South America, and from Asia, stating that they could not be distinguished
from the Siberian mammoth; others ascribed them to Cuvier's "Tapirs gigantesques," still others believed them
'[Regarded by Doctor Hopwood as a synonym of Dinulherium bozasi of Arambourg, 1934. — Editor.]
 SUMMARY 1555

to be related to the hippopotamus and to the rhinoceros. Many Buckland (1835) and
of the early authors, e.g.,

de Blainville (1837) were of the opinion that Deinotherium frequented freshwater lakes and rivers and was most
nearly related to the tapirs.
The skeleton of Deinotlxerium bavaricum found at Franzensbad in 1883 fails to support the original theory
that the Deinotheres were a fluviatile or water-loving animal which frequented streams and used its lower tusks
for the prehension of food from the banks, although during the warm seasons it doubtless bathed in shallow
waters, like the modern Indian elephant. On the contrary, one is it was a forest-living
inclined to conclude that
animal, subsisting upon leafage and tree boughs, to which its sharply crested grinding teeth were adapted like
those of the tapir and of the tree- or shrub-browsing types of rhinoceroses. The very powerful trunk was eminently
adapted to the collecting of tree boughs and leafage; the open supranarial space is enormous. The absence of
large superior tusks, which are seen in all the Mastodontidse and Elephantidse, explains the relatively flattened
form of the top of the cranium, and the forwardly inclined occiput. Deinotherium was neither mastodontoid nor
elephantoid in profile but relatively flattened and depressed, and the body height and length of limb approximated
that of the largest Proboscidea (see Figs. 63, 65 and 70).

It is not an unreasonable hypothesis of extinction that the brachyodont Deinotheres were unable to compete
with the incoming Stegodonts with their very numerous transverse crests and tendency to hypsodonty. At all

events, the climax of the series {Deinotherium gigantissimum) in southern Eurasia became extinct in Middle Pliocene
time, just prior to the appearance of numerous species of Stegodon. In Africa the Deinotheres survived into
Middle Pleistocene time {Deinotherium hopwoodi^).
Migration. — The evidence available seems to point to an African origin of Deinotherium, where specimens
have been found as far south as Tanganyika Territory, since there is no indication of ancestral forms of this
genus in any Eocene or Oligocene horizon of Eurasia at present known. From Africa these animals spread north-
ward through southern France, Germany, Hungary, Rumania, and as far east as Baluchistan and India; their
most northerly range, so far recorded, is the Ural Mountain region.

MASTODONTOIDEA
Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID^ Osborn, 1921
(Syn. TRILOPHODONTID^ Simpson, 1931)-
Subfamily: Longirostrin^ Osborn, 1918 (Syn. Trilophodontinse Scott, 1937)
Genera: Trilophodon Falconer, 1846, 1857; Megabelodon Barbour, 1914, 1917
Subgenera: Genomasiodon Barbour, 1914; Choerolophodon Schlesinger, 1917; Tatahelodon Frick, 1933
(Compare Vol. I, Chap. VIII, also Figs. 189, 277, 680, also Vol. II, p. 1531 and Figs. 1221, 1222, 1227, and PI. xv)
The Longirostrines, long-jawed bunomastodonts of Eurasia and North America, including Trilophodon
and Megabelodon. Probably flood-plain and low savanna habitat.
The history of the subfamily Longirostrinse is complex. While it was at first thought to embrace both the
hyperlongirostral {Trilophodon) and the medilongirostral {Tetralophodon) mastodonts (see Vol. I, p. 231), it is
now restricted to the former, the Tetralophodonts having been removed by van der Maarel in 1932 to his new
subfamily Tetralophodontinae, under which heading (pp. 343-379 of the present Memoir) they are treated. The
ancestral form of the Longirostrinse was believed to be Phiomia of the Oligocene of the Fayum of Egypt, owing
chiefly to the fundamental bunomastodont pattern of its molars, with 'central conules' conspicuous in Phiomia
'ISec footnote —
on opposite page. Editor.]
'[See page 1.525above, where Doctor Simpson states that he now prefers Gomphotheriida; Cabrera, 1929. —Editor.]
 :

1556 OSBORN: THE PROBOSCIDEA

osborni (see p. 715), but subsequently, after comparing the sectioned tusks of Phiomia, Trilophodon, and Amebelo-
don, Professor Osborn came to the conchision that the Phiomia tusk was more nearly like the tusk of Amebelodon
than that of Trilophodon, and as a consequence he placed Phiomia in the direct line of ancestry of Amebelodon
rather than of Trilophodon (see Amebelodontinae below, also PI. xv, opposite, indicated by interrupted line).
The following is a citation from page 249 of Volume I
It has taken many years to establish the fact that a single generic phylum arose from an unknown
ancestor in North Africa and gave rise to the series of long-jawed species in Europe, Asia, and North
America, separated from other phyla by constant increment in the following distinctive characters,
namely, the long, narrow teeth and the excessively long lower jaw. Typified by Trilophodon angustidens
of Simorre. Jaws progressively elongating to an extreme; conules arising in the center of the molar
valleys to form trefoils; Inferior incisors scaptobelodont, rodlike {Trilophodon angustidens, T. osborni,
. . .

T. abeli, T. fricki), more or less elongate or laterally compressed (T. dinotherioides, T. giganteus) some- ;

times tuskless (T. phippsi, T. gaillardi, T. lulli); subcylindrical ('prod-tuskers'); not spatulate or ex-
panded as in the 'shovel-tuskers' {Amebelodon, Platybelodon, Serbelodon).

The principal generic characters of Trilophodon may be summarized as follows:

Trilophodon. —Cranium low, elongate (dolichocephalic). Lower jaws hyperlongirostral. Limited

development of the proboscis, probably a prehensile upper lip, compensated for by great elongation of
the jaw. Superior tusks flattened, recurved, with enamel band; inferior tusks rounded, triangular,
fiattened-horizontal, flattened-oblique, massive, without enamel band; sometimes absent. Grinding
teeth persistently brachyodont, narrow; conules in center of valleys between crests, single mesotrefoils
in superior and inferior molars; rudiments of double trefoils in late geologic stages. Cones and conelets
regular, with simple, smooth enamel. Intermediate molars trilophodont. Third molars with four and
a half to five plus crests. Cement in T. dinotherioides. Third molars only functional in extreme old age.
Limbs mastodon toid feet tetradactyl to pentadactyl digits abbreviated. Low bodied pelvis broad.
; ;
;

Dental formula: Di H:^ C * I H^f Dp ^ P 5^ M ^ef.

Ridge-crest formula: Dp 2||| Dp 3f|| Dp 41 P 4; M If M2 3^^ M 3 ^^1^,
Horizon.— Lower Miocene to Middle Pliocene. Compare Vol. II, p. 1531, also figures 1222-1224, 1227.

Megabelodon. —The genus Megabelodon Barbour was regarded by the present author as a synonym of Trilopho-
don until the discovery in Nebraska in 1935 by Professor Barbour and his field parties of two rostra, one of which
was recovered at, or near, the type locality of Tetrabelodon [Megabelodon] lulli, which prove Megabelodon to be not
only widely distinct from Trilophodon, but a member of an independent generic line characterized by the entire
absence of the inferior tusks and by the broadening of the mandibular extremity into an obtuse, toothless digging
expansion which varies in different specdes. Megabelodon lulli was originally restored with a pair of slender
inferior tusks. Its possible relationship to Gnathabelodon has been suggested.

Horizon.— Of Mio-Pliocene and Lower Pliocene age Vol. also Figs. 1224, 1227).
(cf. II, p. 1.532,

Hemimastodon. — In the case of Hemimastodon Pilgrim, 1912, type Tetrabelodon crepusadi, the present author
was of the opinion that it was not a proboscidean ; the triangular form and simple bunodont structure of the type
molar appeared to him to relate it remotely to the Suina of the piglike Artiodactyla.

Choerolophodon (synonym of Trilophodon). —The subgenus Choerolophodon Schlesinger, 1917, type Mastodon
pentelicus Gaudry, based entirely on juvenile and jaws, was long under consideration by Professor Osborn,
skulls
who finally concluded that its affinities were more nearly with Trilophodon than with Tetralophodon. He was led
to this decision by the fortunate discovery Samos by Dr. Barnum Brown of a palate
in of a young individual with
molars Dp^ to M' in situ, the latter exhibiting two large conules in the median valleys.

Horizon.— Of Lower Pliocene age (cf. Vol. II, p. 1531, also Figs. 1222, 1227).
 Origin. Migration and Evolution of kJrilophodon, uMcgabelodon, ^Gnathabelodon,
oPhioinia, AAmebelodon, *Tetralophodon, xMorrillia. Osborn 1935
PLATE XV
Geologic range: Trilophodon, Lower Miocene to Middle Pliocene; Megabelodon, Mio-Pliocene and Lower Pliocene. Gnalhahdodon, Middle(?) Pliocene.
Phiomia, Lower Oligocene and Upper(?) Oligocene; Amebelodon, Middle Pliocene. Tetralophodon. Mio-Pliocene to Middle Pliocene; Morrillia, Middle
Pleistocene.
 SUMMARY 1557

Genomastodon and Tatabelodon. —As regards Genomastodon and Talabelodon, Professor Osborn was unable to
find clear generic distinctions to validate these terms. Of the latter he remarks on page 325 of Volume I that the
"superiorly grooved and upcurved longirostral symphyses of these types [Tatabelodon riograndensis and T.

gregorii of Frick] scarcely serve for generic distinction from Trilophodon to which they appear to be related both in

length of jaw and in the dimensions of the 4}^ ridge-crested third inferior grinding teeth." It will be noted,
however, that both Genomastodon and Tatabelodon are retained as subgenera or synonyms of Trilophodon (see
Vol. I, pp. 293, 298, and 324).

Migration. —The classic Trilophodon angustidens of the Middle Miocene of France is now traced to the
T. angustidens libycns of North Africa through the very primitive T. pontileviensis of the Lower Miocene of

France and the T. cooperi of the Lower Miocene of Baluchistan ; it passed through the Lower and Middle Miocene
stages of Europe and Asia into the extremely long-jawed American types of the Upper Miocene and the Pliocene
of Dakota, Nebraska, Colorado, and New Mexico, to which different generic names have been assigned, such as
Megabelodon and Genomastodon (sec migration map. Fig. 1227). Dr. Chester Stock lists Trilophodon sp. as occurring
in the Ricardo deposits of the Mohave Desert, California (see Stock, 1928.1, pp. 43-47). Recently Doctor
Hopwood has described members of this subfamily from China (see p. 702 of Vol. I of this Memoir). So far as
known, Trilophodon became rare, or migrated from Eurasia to America, at the close of Miocene or beginning of
Pliocene time both in Europe and Asia.

Superfamily. MASTODONTOIDEA Osborn, 1921. Family: liUNOMASTODONTID^ Osborn, 1921

(Syn. TRILOPHODONTID^ Simpson, 1931)'

Subfamily: Gnathabelodontin^ Barbour and Sternberg, 1935

Genus: Gnathabelodon Barbour and Sternberg, 1935

(Cf. Vol. I, Appendix, pp. 710-714, and Fig. 668; Vol. II, p. 1533, and Fig. 1227, also PI. xv)

Gnathabelodonts, typical "shovel-jawed longirostrines," of progressive bunomastodont type, with broad

tuskless mandibular symphysis and very broad molars. The present author states on page 711 of Volume I that

the "mandibular and dental characters widely separate Gnathabelodon from the shovel-tuskers, flat-tuskers, or any
of the remaining genera of the Bunomastodontidse and Serridentidae and are in full accord with Barbour's con-
ception of this as representing a distinct subfamily remotely related to the Amebelodontinae."

Gnathabelodon. —
Skull very large, the width across the occipital condyles being 294 mm. Lower
jaws longirostral, expanding broadly towards the extremities with an elongated and longitudinally groov-
—
ed symphysis a broad, birdlike, toothless bill analogous to that of the duck-billed Platypus in the
Monotreme order. Superior tusks massive, rounded, and outcurved, with upturned tips; no enamel.
Inferior tusks wanting, replaced by a broadly expanded, sharpened bony border, probably sheathed with
hardened epidermis. Superior and inferior grinding teeth tetralophoid to pentalophoid, ptychodont,
with double entotrefoils and rudimentary ectotrefoils in superior molars, double ectotrefoils on inferior
molars. In general, of progressive, bunomastodont type with central conules. Intermediate molars
trilophodont. In old age reduced to a single pair above and below. Cement in valleys of third superior
molars.
Dental formula incompletely known : I ^~ M^
Ridge-crest formula: M 2l^ M 37^^.

Horizon.— Middle(?) Pliocene. Compare Vol. II, p. 1533, and figures 1224, 1227.

Migration. —The genotype Gyiathabelodon thorpei is the only species of this genus thus far discovered, and
was found near Ogallah, Trego County, western Kansas.
'[See footnote 2 on page 1555 of this chapter. — Editor.]
1558 OSBORN: THE PROBOSCIDEA

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID^ Osborn, 1921

(Syn. TRILOPHODONTID.E Simpson, 1931)'

Subfamily: Amebelodontin^ Barbour, 1929

Genera: Ajnebelodon Barbour, 1927; Phiomia Andrews and Beadnell, 1902
(Cf. Vol. I, pp. 333-338, 690, 715-719, Pls.v-vii, x, xi. Fig. 277; Vol. II, p. 1533, Figs. 1220, 1224, 1227, PI. xv)

Amebelodonts, longirostral trilophodont mastodonts of North Africa and Central United States, differing
from the typical Longirostrinae in the high degree of specialization for the shoveling or digging functions of the

inferior tusks, which are reinforced by internal dentinal laminae. This tusk structure precludes the analogy of
Amebelodon to Platybelodon in which the inferior tusks consist of closely compacted rod-cones. The postsymphy-
seal portions of the jaw and grinding teeth undergo little progressive evolution except in size; it is the anterior
symphyseal and rostral region of the amebelodont shovel-tuskers that undergoes such an extraordinary special-
ization, while the grinding teeth remain substantially the same.
Amebelodon, typical 'shovel-tusker' of Nebraska, descendant of Phiomia of the Fayum. Cranium
and rostrum uniformly abbreviated and reduced in size. Mandibular rostrum relatively long and
slender. Lower jaws elongated (longirostral), nearly horizontal, moderately expanded towards the
extremities. Superior tusks reduced in size, downcurved, outcurved, with persistent outer enamel
band; inferior tusks without enamel band, composed of concentric dentinal laminie (1-9), instead
of dentinal rod-cones as in Plahjbelodon, greatly elongated, somewhat broadened, closely appressed in the
median line, tips rounded, alveolar portion plano-concave above. Mass of the Aynebclodon tusk more
than seventy times greater than that of the Phiomia tusk. Molars large, narrow, with double trefoils and
central conules; inferior molars with five ridge-crests; cement present. From external cingulum rise
many short, blunt cones.
Dental formula: Incompletely known.
Horizon.— Middle Phocene. Compare Vol. II, p. 1533, and figures 1224, 1227.

Phiomia, ancestral 'Fayum shovel-tusker' of North Africa. Of medilongirostral type; cranium low,
abbreviated; palate long and narrow; symphysis long; progressive dolichopy, also progressive doli-
chodonty and bunomastodonty, the latter character tending to place these animals in the direct line
of ancestry of Amebelodon fricki Superior tusks sharply pointed, downcurved, with enamel band on out-
.

er surface. Inferior tusks horizontal, spatulate, without enamel, composed of concentric dentinal
laminae (1-7); incisive alveoli elongate (P. osborni); T)i, with straight inner border, convex outer
border marked by crenulations or serrations, enamel confined to the tip. Molars typically bunodont,
relatively long and narrow (progressive narrowing of M3 prophetic of Amebelodon fricki) brachyodont, ,

rudimentary or progressive conules; generally trilophodont, last inferior molar in Phiomia osborni
subtetralophodont.
Dental formula: Di „:^ C § I H^ Dp fi^ P |^ MH
Ridge-crest formula: Dp 2\t Dp 3|* Dp 4f P 3; P 4/+ M If M 21 M 33^,
HoHizoN. — Lower Oligocene (Fluvio-marine formation) and Upper(?) Oligocene. Compare Vol. I,
figure 23; Vol. II, p. 1533, and figures 1220, 1224, 1227.

Migration. — Up to the year 1934 the progressive species of Phiomia from the Fayum of North Africa were
believed by the present author to be broadly ancestral to the Trilophodon angusiidens phylum of Europe, chiefly
because of the fundamental bunomastodont pattern of the Phiomia molars with 'central conules' conspicuous in
Phiomia osborni. Finally, however, he was of the opinion that Phiomia was the direct ancestor of Amebelodon,
as stated in Volume I, p. 715: "Not only in external form but in the internal macroscopic and microscopic
structure of the incisive tusks (.see PI. v, A-F3) Phiomia wintoni leads into Phiomia osborni and after an immense
interval of geologic time'-' spanning the Upper Oligocene, the entire Miocene, and Lower Pliocene periods, the
'(Sec footmitc 2 on i)agc of this chapter.
1."),').")
—
Editor.)
-["Tliirty million year evohition of the lower jaw and paired inferior tusks from the Phiomia mitior of the Oligocene of Nortli Africa into the Amebelodon
fricki of the Pliocene of Nebraska" (cf. Osborn, 1935.937, p. 408, fig. 3; also PI. v — — —
with modifications of Vol. I of the present Memoir). Editor.)
 SUMMARY 1559

Middle Pliocene Amehelodon fricki appears in Nebraska as the indubitable successor of Pliiomia osborni of the

Oligocene River Nile of Egypt." The abundance of Phiomia is attributable to its having a flood-plain habitat
similar to that of Triluphodon and other longirostrines, which are by far the most numerous proboscideans through-
out Ohgocene, Miocene, and Lower Pliocene times.

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID^ Osborn, 1921

(Syn. TRILOPHODONTID^ Simpson, 1931)^
Subfamily: Tetralophodontin^ van der Maarel, 1932
Genera: Teim/op/iodon Falconer, 1847, 1857; AfomMf a Osborn, 1924. Subgenus: Lydekkeria, O^hovn, \'d2'^

(Cf. Vol. I, pp. 342-379, 690, 703-705, Pis. x, xi, and Fig. 307; Vol. II, p. 1533, Figs. 1221, 1224, 1227, PL xv)
The Tetralophodonts, imperfectly known Medilongirostrines of Eurasia and North America, with four-
crested or tetralophodont intermediate molars, single trefoils in the Miocene stage {Lydekkeria) rudimentary ,

double trefoils in the Pliocene stage {Tetralophodon) ; brachyodont progres.sive to hypsodont, with cement and
complete doubling of trefoils in Pleistocene time (Morrillia).

Cranium brachycephalic, with rounded temporal region; moderate elongation of the rostrum.
Palate contracted, with posterior nares directly opposite pentaloph of M^. Lower jaw medilongirostral.
Superior tusks with enamel band. Inferior tusks flattened, oval, retrogressive, absent(?) in females
{T. campester), without enamel band (the exception being T. grandincisivus) Superior and inferior tusks .

straight (Fig. 320). Molars progre.ssively brachyodont to subhypsodont {Tetralophodon), to hypsodont

and covered with cement {Morrillia). Intermediate molars tetralophodont. Third molars long and nar-
row, with 5Kto6j3 ridge-crests (Tetralophodon), to 8% superior crests {Morrillia) third superior molars in ;

T. fricki exhibit forward inclination of the crests. Central conules variable, cones and conelets relatively
smooth and regular; progressive doubling of the trefoils. Third grinders only in use in old age.
Dental formula: 1 H^ Dp f^ M i^
Ridge-crest formula: Dp 2^ Dp 3? Dp 4^ M 1^ M 2|ii M 3f|fcf||

Horizon. — Mio-Pliocene Middle Pliocene {Tetralophodon), to Middle Pleistocene {Morrillia).

to
Compare Vol. I, figure 307; Vol. II, p. 1533, and figures 1221, 1224, 1227, also PI. xv.

It has been erroneously assumed by some authors that animals of the Trilophodon angustidens type gave
rise to animals of the Tetralophodon type by abbreviation of the jaw and by the addition of a fourth crest on the
intermediate molars. This assumption is disproven by three facts: (1) The true Trilophodon persisted after the
true Tetralophodon had appeared; (2) Tetralophodon is a Medilongirostrine, whereas Trilophodon is a Hyper-
longirostrine ; (3) a true ancestral species of Tetralophodon {Lydekkeria), e.g.. Mastodon [= Tetralophodon {Lydek-
keria)] falconeri Lydekker, is recorded in the Middle Pliocene, Dhok Pathan beds, of India. Con.sequently the
separation by Maarel in 1932 of the Tetralophodonts from the Trilophodonts under the subfamily name Tetralo-
phodontinse has been adopted in the present Memoir.

These medilongirostral mastodonts arose independently from an ancestral Oligocene stage which might be
comprised within the genus Phiomia. This stage was less specialized than Phiomia osborni, which leads directly
into Amehelodon fricki, as mentioned above under the Amebelodontinse (p. 1558). The distinctions between the
genus Trilophodon, the typical genus Tetralophodon, and the genus Morrillia are entirely in progressive characters
which developed from the Mio-Pliocene into Pleistocene times.

The subgenus Lydekkeria Osborn, 1924, genotype Mastodon falconeri Lydekker, 1877, of supposed Middle
Miocene age by Lydekker, Pflgrim, and Osborn, now proves to be of Middle Pliocene age, according to Hopwood
'[See footnote 2 ou page 1555 of this chapter. — Editor.)
1560 OSBORN: THE PROBOSCIDEA

(see Vol. I, p. and therefore a contemporary of 'Mastodon' [Tetralophodon] punjabiensis Lydekker, 1886.
353),
This being the case, Lydekkeria may become a synonym of Tetralophodon, and T. punjabiensis may become
a synon>Tn of T. falconeri. This leaves the Miocene ancestry of Tetralophodon uncertain, but the primitive Mio-
Pliocene stage {Lydekkeria) appears to be ancastral to the Lower Pliocene Tetralophodon.

Migration. — By referring to figure 1227 of the present Memoir, also to PI. xv, it will be noted that the
general trend of migration is from the west to the east. A fragmentary molar of a member of the Tetralophodon-
tinae ('il/.' longirostris?) is Smendou north of Constantine, northern Africa.
recorded from the lacustrine beds of
Professor Osborn in 1925 in his article in Natural History on "The Elephants and Mastodonts Arrive in Amer-
ica," pp. 12 and 13, states that the "Tetralophodonts are in all countries very rare, yet we can trace their long
migration through eastern Europe into India and China, until finally they arrive in Kansas and Nebraska."
Since that time Tetralophodon bumiajuensis of Java has been described by van der Maarel, and the present author,
in the first volume of this Memoir, described T. fricki from northern Texas. From the limited evidence now at
hand it would appear that Tetralophodon became rare in Eurasia early in Pliocene time but lingered until Middle
Pleistocene time on the ancient flood-plains of Nebraska.

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID^ Osborn, 1921

(Syn. TRILOPHODONTID^ Simpson, 1931)'
Subfamily: Notorostrin^ Osborn, 1921 (syn. Cordillerioninse Scott, 1937)
Genus: Cordillerion Osborn, 1926
(Cf. Vol. I, pp. 543-566, also Pis. x, xi, and Fig. 501; Vol. II, p. 1534, and Figs. 1224, 1225, 1227, PI. xvi)

Cordillerion. —
Cranium mastodontoid, of relatively low, simple character as in the primitive
Trilophodonts rostrum elongate. Jaws broad, deep, and abbreviated (brevirostral) symphysis is of medi-
; ;

um length. Superior incisive tusks spiral or horizontal, down- and outcurved, around which is wound
more or less spirally a broad enamel band. Inferior incisive tusks wanting, probably present in juvenile
stages. Grinding teeth brachyodont, never hypsodont, moderately elongate, with single trefoils, parallel
in this respect with Trilophodon, central conules more or less conspicuous, cement in progressive stages.
Intermediate molars with tendency to tetralophodonty (subtetralophoid) in 2. In final phyletic M
stages grinding tooth action concentrated on second and third superior and inferior molars. Adaptation
chiefly to a mountain habitat, grinders adapted principally to a browsing habit, tusks probably used in
the uprooting of plants for food.

Undoubtedly of Old World ancestry, although no member of the Notorostrinae has as yet been found
in the Eastern Hemisphere.
Dental formula: I '-'" Dp jf^ ^M
Ridge-crest formula: Dp 2] Dp 3J Dp 4} It M M 2'^!^ M 3 jTrfsw

Horizon — Pliocene and Pleistocene. Compare Vol. I, figures 501 and 680; Vol. II, p. 1534, and
figures 1224, 1225, 1227, also PI. xvi.

The Notorostrines. —The earliest member of the subfamily Notorostrinae is the South American Masto-
donte des Cordilieres of Cuvier described in 1806 by him Mastodon andium, thus
and later (1824) designated

superseding the name Mastodon cordillerarum of Desmarest, 1820-1822. This species was for a long time confused
with the Mastodonte humboldien, also of South America, described by Cuvier in the same publication of 1806.

In January of 1921 —a century later— Professor Osborn reviewed (Osborn, 1921.515, p. 10) these two classic
species, including also the Dibelodon [
= Cordillerion] tropicus and Mastodon = Stegomastodon]
[ successor of Cope,
designating them as the "Brevirostrines of South America," and in September of the same year he established
'(See footnote 2 on page 1555 of this chapter. — Editor.)
t
 RHYNCHOTHCRIUM FALCONEKI, re^^s

PENTALQPHODON FALCONERI,
NDIA

CORDILLERION
EDENSIS, CALIFORNIA {iRHYNCHOTHERIUM SHEPARDI
EDENSE, CALIFORNIA
ANANCUS AR\ ERNEN&IS, S\ NCONOLOPHUS DIIOkPATtlANENStS,
FRANCE. '^'^^ ^'^^ """^
PENTALOPHODON
SiyALENSIS, INDIM

RHYNCHOTHERIUM BROIVNTl mej(ico AYBELODON

HONDURENSIS, mohouR-M

ANANCUS PER/MENSIS.
INDIA

RHYNCHOTHERIUf^t TLASCALAE. S YNCONOL OPHUSCORR UCA TUS, india

MEXICO

BLICROTHERIUf^l BLICKI, Honduras

^
3 YNCONOL OPH US
PROPATHANENSIS. india

BLICKOTHERIUM
EUHYPODON.
NEBRASKA

6YNCONOL0PHUS
RHYNCHOTHERIUM CHINJIENSE, india PTYCHODUS, INDIA
ANANCUS PROPERIMENSIS. india

RHYNCHOTHERIUM SPENCERI, egypt

Oricim. Aligratlon and Evolution of nCordillerLon. mRhynchotheriarn. oBlickotherium,

A^AybcLodon, ^Anancus, *Pentalophodoa, xSl/izco no tophus. Osborn 1955
PLATE XVI
Geologic range: Cordillerion, Pliocene and Pleistocene. Rhynchotherium, Middle Miocene to Upper Pliocene; Blickotherium, Pliocene; Aybelodon,
Pliocene. ATiancus, Mio-Pliocene to uppermost Pliocene; Penialophodon, Upper Pliocene; Synconolophus, Mio-Pliocene to Middle Pliocene.
 SUMMARY 1561

his new subfamily, the Notorostrinse (1921.526, p. 330) to embrace not only the four species just mentioned but
six additional species, namely, Mastodon bolivianus and M. chilensis of Philippi, and M. platensis, M. rectus, M.
argentinus, and M. superhus of Ameghino, on the ground that they were profoundly separated from other buno-
mastodonts by the progressive abbreviation of the jaw, and that they were the only members of the great order
Proboscidea which, as far as known (1921), entered the South American continent. Professor Osborn's researches
had not then revealed the fact that other mastodonts, namely, Rhynchotherium Falconer and Notiomastodon
Cabrera, had also wandered into South America as well as one species of elephant belonging to the Pnrelephas
phylum, i.e., Parelephas columbi cayennensis Osborn, of French Guiana.

Perhaps a more important step was taken by Professor Osborn in 1923 (1923.601, p. 1) when he separated
still

the Mastodon humboldtii group from the M. andium group, making M. humboldtii the type of his genus Cuvieronius,
distinguished by double trefoils on the superior and inferior molars, and by simple, rounded, outwardly and up-
wardly curved superior tusks, lacking enamel band (see p. 540 of Vol. I), in contradistinction to M. andium with
single trefoils, more or less conspicuous central conules in the molars, and broad, spirally enameled superior tusks,
a peculiarity not observed in any other proboscidean, to which subsequently (1926.706, p. 15) he assigned the
generic name Cordillerion. This subfamily of short-jawed bunomastodonts, therefore, is monophyletic, including
the genus Cordillerion only.
Migration. —As mentioned above, the Notorostrinse probably originated in the Old World; fossil evidence
of this fact, however, is lacking, for the earliest occurrence, as far as known, of the genus Cordillerion is the Upper
Pliocene C. edensis Frick-Osborn, found at Mt. Eden Hot Springs, San Bernardino County, California; other
discoveries are at Benson, Arizona, and near Sinton, Texas, also of the Pliocene. It would seem, therefore, that

the trend of migration was southward, through Mexico, into South America, as far as Uruguay.

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID^ Osborn, 1921

(Syn. TRILOPHODONTID.^ Simpson, 1931)'

Subfamily: Rhynchorostrin^ Osborn, 1918, 1921 (syn. Rhynchotheriinse Cabrera, 1929)

Genera: Rhynchotherium Falconer, 1862, 1868; Blickotherium Frick, 1933; Aybelodon Fvick, 1933
(Cf. Vol. I, pp. 475-513, 690, also Pis. x, xi, and Figs. 451, 483; Vol. II, p. 1531, and Figs. 1220, 1221, 1224,
1227, also PL xvi)

The Rhynchorostrines, beak-jawed mastodonts, typified by Rhynchotherium Falconer, so named because

of the sharply deflected rostrum. The subfamily Rhynchorostrinae is the least known of any of the bunomasto-
donts. The genotype of Rhynchotherium is the cast of a lower jaw, the original of which was found at Tlascala,
Valley of Mexico. It was at first characterized by Falconer (in a letter to Lartet of Sept. 12, 1856) as resembling
"very much the figure in Alcide D'Orbigny's Voyage, described by Laurillard as M. Andium," but he stated that
the "Genoese palaeontologists had provisionally named it Rhynchotherium, from the enormous development of the
beak, approaching Dinotherium." This letter was not published until 1868 (see Pal. Mem., Vol. II, pp. 74, 75).

He further stated that he saw this cast in Genoa, an unfortunate substitution for Geneva (see Falconer, 1863, p.

56), an error to which Felix and Lenk called attention in 1891, but which was apparently unnoticed until Professor
Osborn's researches for the present Memoir led him to make inquiry of Dr. Hans Georg Stehlin, who kindly
located the cast in the Museum d'Histoire Naturelle of Geneva. Through the courtesy of the Director, Dr.

Pierre Revilliod, a duplicate cast was presented to the American Museum.

Professor Osborn was the first to use the specific name tlascalse in 1918 (Osborn, 1918.468, table opposite
p. 134); thus the genotype is Rhijnchotherium tlascalx. The discovery in 1911 by Dr. Barnum Brown near San
'[See footnote 2 on page 1555 of this chapter. — Editor.)
1562 OSBORN: THE PROBOSCIDEA

Jose de Pimas, Sonora, Mexico, of a fine specimen of a lower jaw referable to Rhynchotherium — made the
in fact,

neotype by Professor Osborn in 1921 (1921.515, pp. 5, 6), owing to the supposed loss of the cast —^served to confirm

the presence of this genus in Mexico. On recovery of the cast in Geneva, was found that the type and neotype
it

differed sufficiently to warrant the naming of a new species; consequently Professor Osborn assigned to the neo-
type the name Rhynchotherium browni (see Vol. I of the present Memoir, p. 494). During the years following
Falconer's description, specimens of Rhynchotherium were unearthed in California, Colorado, Montana, Nebraska,
and Texas.
The Rhynchorostrina^ include the fewest and the least known species of the bunomastodonts. Tracing the
phylum back into its Eurasiatic or Old World ancestry, we find a primitive lower jaw with downturned symphysis
and laterally compressed incisive avleoli (Fig. 455) described by Fourtau in 1918 as Mastodon spenceri from
Moghai-a, Egyi:)t, but provisionally referred by Professor Osborn to Rhynchotherium spenceri, which may confirm
the hyi)othesis of the African origin of the Rhynchorostrinse. Also doubtfully referred by Professor Osborn to
Rhynchotherium {R. chinjiense) is a left ramus, with Ma, M3, in situ, found by Dr. Barnum Brown in 1922 in the
Mio-Pliocene of India, near Chinji Bungalow. Dr. D. K. Chakravarti in 1935, p. 209, states that he sees no
evidence of the down turning of the symphysis, and that the "warping of the alveolus of the tusk appears to the

present author to be a case of distortion superinduced upon the specimen after its entombment"; he suggests
that "it should be referred to Trilophodon angustidens (Cuvier) var. chinjiensis (a name proposed by Pilgrim in
)."
1913 for a form discovered in the Salt Range. . .

Rhynchotherium. — No Face of medium length, maxilla partly down-

complete cranium known.
turned. Mandible of medium symphyseal portion strongly deflected. Superior and inferior
length;
tusks, I", lo, laterally compressed, bent downwards and outwards, with broad external enamel band;
inferior tusks slightly upcurved. Grinding teeth of bunomastodont type, relatively broad, simple,
brachyodont, single trefoils only, inferior 'central conules' variable, superior 'central conules' absent.
Molar enamel thick. Third superior grmders with three and a half to four plus ridge-crests; third
inferior grinders not exceeding four and a half to five ridge-crests functional grinders in adults reduced
;

toM 2-M 3, finally to 3. M

Intermediate grinders trilophodont.
Dental formula: I ^ffff Dp H M iff
Ridge-crest formula: Dp f^ Dp 31 Dp 41 M If M 2^ M 3=*^
Horizon. — Middle Miocene to Upper Pliocene. Compare Vol. II, p. 1531, and figures 1220, 1224,
1227, also PL xvi.

Up to the year 1929 the Rhynchorostrinse had been considered as monophyletic. The discovery, however,
near Tapasuma, Honduras, by Mr. John C. Blick of the Frick Expedition of 1929, of two long slender mandibles,
one adolescent, the other mature, amplified our knowledge of these rare and probably forest-living animals, with
the surprising result, as described by Mr. Childs Frick in 1933 in his article on "New Remains of Trilophodont-

Tetrabelodont Mastodon ts," that two new genera were added to this subfamily, namely, BUckoUierium, genotype
Blickotherium blicki, and Aijhelodon, genotype Aybelodon hondurensis. The distinctive feature of Aybelodon is the
enamel-less incisors, differing in this respect from other members of the Rhynchorostrinse, but the profile of the

type jaw reveals its rhynchotherine affinity and places it in a "distinct and unique line of succession."

Blickotherium. —
No complete cranium known; see, however, figure 462 A for palate of Mastodon
euhypodon Cope referred to Blickotherium by the present author. Mandible elongate, extremely slender,
less deflected than in Rhynchotherium. Symphysis laterally compressed, downwardly curved. Inferior
tusks upcurved, compressed laterally, broad external enamel band. Apparent presence of a replace-
ment P4 in the adolescent mandible.
Ridge-crest formula (Frick): Dp 4:^+ (possible replacement Pj) M 1? M 23^4
Ridge-crest formula (Osborn): M i^ M 2t M 33^4
Horizon. —Pliocene. Compare Vol. II, p. 1531, and figures 1224, 1227, also PI. xvi.
 ;

SUMMARY 1563

Axjhelodon. —
Cranium unknown. Mandibular rami of long, relatively slender type; vertical ramus,
condyle and coronoid process low. Symphysis strong, very elongate, laterally and vertically compressed,
upcurved, extremely narrow, transverse exceeding vertical diameter, as contrasted with RUck other ium
blicki. Inferior tusks large, broad, closely appressed, upcurved, without trace of enamel band. Superior
tusk (referred), possibly representing a distal section of upper left side, exhibits tendency toward inward
and forward curvature, the presumed outer surface with remnants of enamel band. First and second
inferior molars small; third inferior molars relatively broad and short; very thick enamel. Second
inferior molars probably trilophoid third inferior molars fully tetralophoid with rudimentary pentalophid.
;

Ridge-crest formula: M 2y^ M 3^^^

Horizon. —Pliocene. Compare Vol. II, p. 1531, and figures 1224, 1227, also PL xvi.

In summarizing the persistent primitive and progressive characters of the Rhynchorostrines, we may mention
the following:

Persistent Primitive Characters. — (1) Grinding teeth persistently brachyodont or short crowned; (2) no
evidence of hypsodonty or deposition of cement; (3) enamel unusually thick; (4) simple external trefoil spurs on
inferior molars and internal trefoil spurs on superior molars, no trace of doubling of the trefoil spurs; (5) inferior

'central conules' variable, superior 'central conules' absent.

Progressive Characters. — Superior tusks progressive, elongated, and laterally flattened, with broad persistent
enamel band. Inferior tusks progressive, compressed laterally, elongated, upturned, with persistent lateral
enamel band. The tusks of Aijbelodon hondurensis, however, are entirely without enamel.

Migration. —From the Miocene of ?Egypt, through the Mio-Pliocene of ?India, to the Middle to Upper
Miocene of Montana, Upper Miocene-Upper Pliocene of California, Nebraska, Texas, Colorado, Mexico, and
Honduras. See especially figure 483 of Volume I of the present Memoir, and the accompanying PI. xvi.

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: BUNOMASTODONTID^ O.sborn, 1921

(Syn. TRILOPHODONTID.E Simpson, 1931)'

Subfamily: Brevirostrin^ Osborn, 1918 (syn. Pentalophodontinse Scott, 1937)

Genera: Ajiancus Aymard, 1855, 1859; Pentalophodon Falconer, 1857, 1865; Synconolophus Osborn, 1929
(Cf. Vol. I, pp. 617-665, 690, Pis. x, xi, and Fig. 590; Vol. II, p. 1534, and figures 1221-1223, 1227, also PI. xvi)

The Brevirostrines. —We now come to one of the mastodont divisions that never reached America, as far
as is known at present, namely, the Brevirostrinse, a subfamily provisionally named by Professor Osborn in 1918
(1918.468, p. 136) and fully defined by him in 1921 (1921.515, p. 11). In Volume I, p. 627, of the present Memoir,
he summarizes the characters of this subfamily, as follows: "These short-jawed bunomastodonts are readily
distinguished from all others by four principal characters: First, the skull and jaws short, as in the elephants;

second, the grinding teeth finally reduced to one above and one below, as also in the elephants and in the Stegodonts
third,perhaps most distinctive and unique, the alternation of the main internal and external cones which are
placed diagonally instead of transversely to the long axis of the crowns, a character first observed by Cautley in
describing his species Mastodon sivalensis also by Falconer (1846, p. 50) in comparing the Mastodon sivalensis of
the SiwaUks with M. latidens, and later (1868, II, pp. 29-31) with the Mastodon arvernensis of the British coast
Upper Pliocene; the fourth distinctive character is the pUcation or folding {Sxjnconolophus) of the enamel borders
of the molar ridges, giving the hippopotamus- and pigUke character termed 'chcerodont' by Schlesinger, the
'[See footnote 2 on page 1555 of this chapter. —Editor.]
1564 OSBORN: THE PROBOSCIDEA

grinding action of the teeth being obviously as in pigs, and the hippopotami. The grinders, originally brachy-
odont, become subhypsodont; thus the single superior and inferior molar teeth become highly effective masti-
cating organs, labyrinthodont in pattern and serving effectually in the comminution of food."

It appears that the Brevirostrina? include three phyla, namely, Anancus Aymard, 1855, 1859, Pentalophodon
Falconer, 1857, 1865, and Synconolophus Osborn, 1929. Stegomastodon Pohlig, 1912, at first regarded as belonging
to the Brevirostrinse, was removed by Professor Osborn to his new family Humboldtidse, because of a "fundamental
resemblace of the grinding teeth to those of 'Mastodon' humboldtii."

Ananctis, 'straight-tusked brevirostrine' of southern Eurasia. Aymard based his genus on Anancus macro-plus,
1855, a generic and specific name applied to bones and teeth of an adult mastodont similar to the species described
and figured by Croizet and Jobert as Mastodon arvernensis from Auvergne, based on the milk dentition. Anancus
macroplus is therefore regarded as synonymous or identical with A. arvernensis, which latter becomes the genotype.
The term Anancus signifies without bend, probably in allusion to the straight tusks or upper incisors. This
phylum is purely Eurasiatic. A. arvernensis and A. falconeri may possibly be derived from A. perimensis or A.
properimensis of India.

^na«cws. —Cranium approaching true elephantine type, with brachycephaly, hypsicephaly, and fore-
and-aft abbreviation relatively primitive elongation of the rostrum in front of the orbits, an adaptation
;

to the presence of two grinding teeth in use at the same time, namely, M
2-M 3 frontal region relative-
;

ly narrow or laterally compressed; summit also narrow, since the temporal fossae are divided by a
relatively narrow frontoparietal plate as compared with the very broad frontoparietal plate in Pentalo-
—
phodon falconeri; occipital condyle very prominent and pedunculate a unique feature in the Probo-
scidea. Anterior portion of symphyseal region abbreviate. Superior tusks straight, elongate. No in-
inferior tusks. True molars brachyodont to subhypsodont; cones simple, smooth, compressed fore and
aft. Single external median conules in inferior molars; internal median conules in superior molars.
Deciduous molars (premolars) grooved, ptychoid, or plicated, externally. No trefoils. Crowns brachyo-
dont, cones erect {Anancus macroplus and related species) crowns subhypsodont, strongly inclined and
;

alternating {A. falconeri). Internal proversion of superior and inferior ridge-crests. Intermediate
molars tetralophodont.
Dental formula: I " ="Dp §;| M ;:|
Ridge-crest formula:
Horizon.— Mio-Pliocene
Dp 2| Dp 3 Dp^ 4 ^^ M It M 2ff^ M 3 §i^
to uppermost Pliocene. Compare Vol. II, p. 1534, and figures 1221, 1222,
1227, also PI. XVI.

Pentalophodon, 'five-crested straight-tusker' of the Siwaliks. Falconer in 1857 characterized this genus as
having the intermediate molars divided upon a quinary ridge formula, evidently with Mastodon sivalensis of
Cautley in mind at the time, for later (1865, p. 262) he clearly designated it as the genotype. His descriptions
and definitions are also based on specimens of superior and inferior molars figured in plates xxxvi, xxxvii of
the "Fauna Antiqua Sivalensis," as well as on a cranium figured in plate xxxiii, fig. 2, with altogether different
and much more progressive grinding teeth. A close comparison of the molars shows that this referred cranium
belongs to a much more progressive species, if not genus, than the type molar of 'Mastodon' sivalensis, which
Professor Osborn in Volume I, p. 653, of the present Memoir made the type of a new species, namely, Pentalo-
phodon falconeri. This Indian form {Pentalophodon sivalensis) has a line of southern Asiatic ancestry of its own,
branching from Anancus perimensis and distinct from the west European A. arvernensis.

—
Pentalophodon. Cranium lofty, hypsicephalic, brachyopic. Occipital condyles elevated. Sym-
physis abbreviate. Incisive tusks unknown. Intermediate molars pentalophodont. Molars subhypso-
dont; cones elevated, strongly alternating, inclined forwards in the lower molars; superior and inferior
ridge-crests extending obliquely forwards from the external to the internal faces, that is, internal
proversion; typically composed of three main irregular cones set widely apart {P. sivalensis), closely
 '

SUMMARY 1565

compacted (P. falconeri) crown progressively chcerodont with twenty-two aristogenes in M^ (P.
;

sivalensis), twenty-seven in M^ (P. Jalconeri); enamel smooth, not ptychoid; no ecto- and entotre-
foils; central conules present in two anterior valleys. Small amount of cement sometimes present.

Dental formula: X^„ Dp

I §;* M {^
Ridge-crest formula: Dp ^ ?M 1^^ M 2^^ M S^f^^^Hf
Horizon.— Upper Pliocene. Compare Vol. I, p. 648; Vol. II, p. 1534, Figs. 1221, 1223, 1227, and
Pi. XVI.

Synconolophus, 'syncone-crested brevirostrine' of the Siwaliks. Falconer's plates or descriptions do not seem
to show grinding teeth referable to this peculiar genus, but memoir of 1880 on "Siwalik and
Lydekker in his

Narbada Proboscidia" figures examples of teeth certainly referable to Synconolophus. The warping or dislocation
of the cones and crests in the superior and inferior molars is much more extreme than in Anancus or in Stegomasto-
don, also the multiplication of separate intermediate conelets, originally springing from the trefoils, is quite
distinct from the regular single or double trefoils of Stegomastodon. The whole surface of the crown becomes
crowded with a labyrinthine pattern of cones, conelets, and more or less separate intermediate conelets, with
warped or dislocated ridge-crests in apparent disorder, giving the appearance best expressed in the Greek generic
term Synconolophus. The genotype is Synconolophus dhokpathanensis, based on a giant cranium found by Dr.
Barnum Brown in 1922 in the Dhok Pathan horizon of India. The three-ridged second molar, M-, suggested
comparison with Trilophodon, but the broad four-ridged third molar, M^, differed from that of Trilophodon and
also from that of Tetralophodon punjahiensis which occurs in the same geologic horizon. It was not until perfect

superior molars were found (see Fig. 631 —paratype), also from the same horizon a pair of referred superior and
inferior molars (Fig. 632), that the structural and phyletic key to this very aberrant genus could be determined
(cf. Vol. I, pp. 654-665).
Synconolophus. —
Cranium platycephalic, dolichopic, brachycranial (Vol. I, p. 649) [dolichocranial?
—see p. 347], somewhat more elongate than in Anancus or Pentalophodon. Postnarial opening far back of
grinders, a generic distinction from Tetralophodon. Mandibular symphysis broad, trough shaped,
concave superiorly {S. propathanensis) indicating a large, fleshy lower lip. Superior tusks large, up-
,

turned, and out-turned, without enamel. Inferior tusks reduced or absent (*S. propathanensis). Milk
tusks with enamel. Grinders with ridge-crests strongly arched or dislocated valleys blocked with warp- ;

ed, compacted conelets and conules, giving on wear a strongly choerodont, ptychoid, crowded, compressed,
enamel-bordered appearance; internal pro version. No trefoils. Cement strongly developed in S. dhokpath-
anensis and S. propathanensis. Cones smooth to deeply grooved. Intermediate molars with trilophodont
ridge formula and small rudimentary half ridge-crests, representing pro-protoloph and tetartoloph.

Dental formula: Di ^^ I "P Dp H M '^,

Ridge-crest formula: Dp 4"-' M V^ M 2*^ M 3J|^

Horizon. — Mio-Pliocene to Middle Pliocene. Compare Vol. II, p. 1534, and figures 1221, 1227, also PI.
XVI.

Geographic Range. — (Vol. I, p. 629) : "Armed with very powerful upper tusks, the enamel bands of which
early disappear, propelled by massive limbs supported on a powerful frame, these animals appear to have been
adapted to browsing on the succulent vegetation of the warm-temperate zones, chiefly of the 40th and 30th
parallels, from France and the southeast coast of England, through northern Italy, then through India, but never
reaching North America, so far as we know at present. Analogous but unrelated are the species of Stegomastodon.'
A step further must be taken in the migration of these animals, owing to the determination by Dr. A. Tindell
Hopwood (1935.1, pp. 57-60) of molars from Shansi, China, as belonging to Pentalophodon sinensis. Professor
Osborn accepted Doctor Hopwood's reference of these molars to a member of the Brevirostrinae, but regarded
them as referable rather to Anancus than more progressive Pentalophodon which has a greater number
to the of
ridge-crests than are present in the Chinese molars (see Appendix to Vol. I, p. 722).
1566 OSBORN: THE PROBOSCIDEA

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: HUMBOLDTID^ Osborn, 1935, 1936

(Syn. STEG0MAST0D0NTIDJ5 Scott, 1937)

Subfamily: Humboldtin^ Osborn, 1934, 1936 (syn. Stegomastodontin» Scott, 1937)

Genera: Cuvieronius Osborn, 1923; Eubelodon Barbour, 1914; Stegomastodon Pohlig, 1912

(Compare Vol. I, Chap. XII, Chap. XIII, pp. 667-684, and Appendix, pp. 722-728, also Pis. x, xi, and figures

501, 590; Vol. II, p. 1535, and figures 1224, 1225, 1227, also PI. xvii)

The Humboldtines, 'hyperptychoid Huraboldtine mastodonts' of the southern United States and South
America. The decision of Professor Osborn in 1934 (Osborn, 1934.924, p. 183, fig. 2) to dismember his subfamily
Notorostrinae (which included the Mastodon [Cordillerion] andluni and M. [Cuvieronius] humholdlii groups, as
well as the Eubelodon morrilli of Barbour) was occasioned by his recognition of the differences in the crania and
dentition of the members of these groups. For example, the Cuvieronius skull converges toward that of Ele-
phas and of Mastodon americanus, while the Cordillerion skull retains the relatively low, simple character of
the primitive Trilophodonts ; furthermore, in Cuvieronius humholdlii the superior tusks are straight or upcurved
without enamel band, and the superior and inferior molars are broad with typically double trefoils and central
conules either inconspicuous or wanting; whereas in Cordillerion andium a broad enamel band winds more or
less spirally around the elongated superior tusks and the molars are narrow with single trefoils on the internal
side of the upper molars and on the external side of the lower molars, as in other bunomastodonts.

In the same article (1934, p. 183) Professor Osborn definitely placed Eubelodon morrilli in the direct ancestral
line of the Humboldtines or Humboldtina? (fig. 2, our PI. x), and accepted Dr. Cabrera's inclusion of Leidy's

Mastodon = Stegomastodon]
[ mirijlcus, 1858, as well as other Stegomastodonts, in the Humboldtinae, which sub-
family is defined on pages 575 and 615 of Volume I of the present Memoir.

Eubelodon. — From
when Doctor Barbour described Eubelodon morrilli, until 1929, the type specimen
1914,
was the single representative of this genus. The discovery of a superb cranium by the Frick Expedition of 1929,
at Horsethief Canyon, near Ainsworth, Nebraska, and referred by Mr. C'hilds Frick in 1933 to this species, shed
additional light on the dentition of Eubelodon, as both the second and third superior molars are present, the latter
beautifully displaying the coronal pattern of 4)1 ridge-crests with double trefoils, at once reminding one of Cuvier's
type of 'Mastodon' humholdlii.

Characters of Eubelodon, 'ancestral Humboldtine mastodont' of Nebraska. Cranial and facial

region primitive, subequal, less elevated than in Cuvieronius humholdlii. Occipital condyle low com-
pared with elevated condyle and occipital crest of Cuvieronius superbus. Rostrum and inferior mandible
elongate, somewhat downturned and pointed, with no trace of alveolus for tusks. Superior tusks
relatively .short, simple, rounded, out-turned, tips chisel shaped, slight downcurvature, conserving the
primitive downcurvature of the paired superior incisors of all primitive mastodonts, divergent, without
enamel band. No inferior tusks. Grinders broad with double trefoils, four ridge-crests and talon.
Superior molars with strong entotrefoils, incipient ectotrefoils indications of cement; broad internal
;

cinguliun outer side more convex, inner side more plane or concave. Cingulum less distinct in inferior
;

molars. Pelvis excessively broad; humerus robust; inferior in height and width to Warren Mastodon
{Mastodon americanus).
Ridge-crest formula: M 2; M 3*;i
— Middle(?) Pliocene.
Horizon. Compare Vol. II, p. 1535, and figures 1224, 1227, also PI. xvir.

Cuvieronius. — In giving the history of the genotype of Cuvieronius, namely, the Maslodonte humboldien
Cuvier, 1806 = Mastodon
[ humboldtii Cuvier in Desmarest, 1818, Mastodon humboldii Cuvier, 1824], Professor
 STEGOMASTODON
AFTONIAt.

CUyiEKONIUS PLATENSIS, Argentina CUl lERON/US POSTKEMUS. hcuadoh.

STEGOMASTODON
MIRIFICUS, ("c^'' '^

hJ£&RASKA

CUIIERONIUS A>ORAE, Ecuador

STEGOMASTODON
PRIMITIl/US,
NEBRASKA

STEGOMASTODON
ARIZONAE,
ARIZONA

STEGOMASTODON
TEXANUS.
TEXA3
STEGOMASTODON
SUCCESSOR.

EUBELODON MORRILLI, Nebraska

Origin. Migration and Evolution oF

•Eubelodon, ^Cuvieronias, *Stegomastodon. Osborn 1935
PLATE XVII
Geologic range: Eubelodon, Middle(?) Pliocene; Cuvieronius, Pleistocene, Upper Pleistocene (C. postrernus); Stegomaslodon, Upper Pliocene and Lower to
Middle Pleistocene.
 SUMMARY 1567

Osborn states in Volume I of the present Memoir, p. 575, as follows: "The weight of Cuvier's authority has been
so great that since 1806 the generic name Mastodon was applied by all authors to humboldtii, until Cope proposed
(1884.2, p. 2) the ill-fated generic name Dihelodon, signifying two-tusker, which he based upon the four-tusker
Mastodon shepardi of Leidy. Cope's practice of including Mastodon andiurn and M. humboldtii also within the
genus Dihelodon was followed by Lull and Osborn until Osborn discovered Cope's error and proposed the generic
name Cuvieronius (1923.601). Meanwhile all the European authors continued to use Cuvier's name Mastodon,
including Boule in his important Memoir of 1920."

Characters of Cuvieronius, 'Humboldtine mastodont' of the Pampean and Andean regions of South
America. Cranium elevated, facial region prominent, expanded, with air-cell chambers as in Mastodon
and Elephas. Rostrum of inferior mandible relatively abbreviate. Tusks relatively large, simple,
rounded, upturned and out-turned or straight, without enamel band; relatively short, robust, upturned
in primitive species (C. ayorx and C. postremus). No inferior tusks. Molars broad with typically double
trefoils; single trefoils in primitive species (C. aijorx, C. postremus). Third superior and inferior mo-
lars tetralophodont to pentalophodont. Intermediate molars with rudimentary tetartoloph. Superior
ridge-crests transverse or retroverted internally; inferior ridge-crests pro verted internally. Superior
and inferior crowns either plane or curved on external side. Central conules inconspicuous or absent.
Ridge-crest formula: M 1|| M 2|^ M 3^^^^
Horizon. —Pleistocene. Upper Pleistocene (C. postremus). Compare Vol. II, p. 1535, and figures
1225, 1227, also PI. xvii.

Stegomastodon. — This genus was thought by Professor Osborn to be the American representative of
first

his subfamily Brevirostrinse, but it has recently been determined that the migration of the true Brevirostrines
{Anancus, Pentalophodon, and Synconolophus) did not extend to the North Ajnerican continent. We are still

without knowledge of the Asiatic and possibly African forbears of Stegomastodon. The Stegomastodon mirificus
group now proves to be closely related in its molar-tooth structure to the Cuvieronius humboldtii group of the
subfamily Humboldtinae. This observation was first made by Dr. Angel Cabrera in 1929 and was confirmed by
Professor Osborn (see Vol. I, p. 612, this Memoir) after a further "close comparison of the fundamental grinding-
tooth pattern in Stegomastodon," which disclosed the fact that "both the superior and inferior ridge-crests of the
true Brevirostrines of Eurasia are proverted, whereas in the Humboldtines of America the inferior ridge-crests
are proverted and the superior ridge-crests are either retroverted or transverse (Cuvieronius) or centroverted
(Stegomastodon)."

Characters of Stegomastodon, Pohlig's 'roof-toothed Humboldtine mastodont' of the southwestern

United States. Cranium abbreviate; palate elongate, postnarial opening far back of grinders (S.
texanus). Mandibular rostrum progressively abbreviated. Superior tusks short, massive, upcurved,
outcurved, without enamel band. No inferior tusks. Tusks in S. primitivus first projected directly out
from the alveoli after which they were simply outcurved and then by inward rotation the tips were
finally incurved at the extremities. This peculiar tusk structure may remove it from the direct ancestry
of the other species of Stegomastodon and place it in a line of its own. Juvenile tusks straight. Molars
brachyodont to subhypsodont cones slightly alternating, deeply plicated, ptychoid, folded, giving
;

a labyrinthine pattern to the worn grinding surfaces double to quadruple to multiple trefoils in both
;

superior and inferior molars, completely blocking the valleys; enamel thick; no central conules;
external pericones block the spaces between the five main crests (S. texanus ref.). Superior ridge-crests
centroverted; inferior ridge-crests proverted internally as in Anancus, Pentalophodon, and Snyconolo-
phus. Cement in transverse valleys. Intermediate molars with trilophodont to tetralophodont ridge
formula, that is, 3K ridge-crests.

Ridge-crest formula: Dp 41 M M 2^ M Z \VT^XtZ

1|{|

Horizon. —Upper Pliocene and Lower to Middle Pleistocene. Compare Vol. II, p. 1535, and figures
1224, 1227, also PI. xvii.
1568 OSBORN: THE PROBOSCIDEA

The final step taken by Professor Osborn in 1935 was the creation of a new family, the HumboldtidiB (see
Osborn, 1935.937, fig. 2, for name only; 1936, Vol. I, p. 722, for definition), to embrace the Eubelodon-Cuvieronius-
Stegomastodon group, or Humboldtinse.

Migration. — (Vol. I, p. 611) : "Putting together the important resemblances between specimens of Eubelo-
don morrilli of Nebraska and the relatively little-known primitive remains of tusks and grinding teeth from the
Inferior Pampean of Argentina, we reach the very important conclusion that E. morrilli is a Middle Pliocene
ancestral stage in the subfamily Humboldtinse on its way from Eurasia through North America to South America.
In the Andean region there appear the primitive short-limbed forms Cuvieroniiis ayorae and C. postremus, and the
primitive typical C. humboldtii of Bolivia, (?)Ecuador, and Chile, quadrupeds of smaller size and possibly of
mountain habitat. Possibly a direct descendant of Eubelodon is the straight-tusked Cuvieronius rectus of the
Inferior Pampean of the Argentine. Finally there appears in the Pampean the giant Cuvieronius superbus and
the more specialized C. platen-sis as distinguished by Cabrera." On a subsequent page (p. 685) Professor Osborn
states that we may be certain that at some future time we may trace Eubelodon back to northern or central Asia
and ultimately perhaps to Africa.

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: SERRIDENTID^ Osborn, 1935, 1936

Subfamily: Serridentin^ Osborn. 1921

Genera: Serridentinus Osborn, 1923; Ocalientinus, Serbelodon, and Trobelodon Frick, 1933

(Cf. Vol. I, pp. 380-473, 729, also Pis. x, xi, and Figs. 344, 373; Vol. II, p. 1535, and figures 1221-1225, 1227,

also PI. xviii)

Perhaps in no other subfamily has there been such confusion in the determination of species as in the Ser-
ridentinse, especially in the differentiation of the Trilophodont and Serridentine unworn molars, in fact, for a

century these animals have been confused with Trilophodon. The salient features which clearly separate the
subfamily Serridentinse from the Longirostrinse (or Trilophodon ts) are: First, the central conules observed in all

species of Trilophodon are entirely absent in the Serridentinse; second, these central conules are functionally
replaced in the Serridentina? by crests or spurs which arise on the sides of the superior internal cones and on the
inferior external cones; and tlxird, these internal and external crests subdivide into two, three, four to six smaller
conelets, which produce a serrated aspect, hence the term 'serrate-toothed mastodonts' or Serridentinae.

By referring to Professor Osborn's first papers, it will be noted (Osborn, 1921.515, p. 8) that he designated as
the "T. [Trilophodon] serridens Phylum" four species, namely, Mastodon serridens Cope, Tetrabelodon serridens
cimarronis Cope, Dibelodon prxcursor Cope, and questionably Mastodon turicense Schinz. Again in his definition of

the new subfamily Serridentinse (Osborn, 1921.526, p. 330) he stated that the "Serridentinse apparently spring from
M. turicense Schinz, 1824, of the Middle Miocene of France and Switzerland —a rare animal, probably because a
forest dweller," and calls attention to the distinguishing spur or crest on the sides of the superior and inferior
grinders of the Serridentinae. Professor Osborn's final conclusions are given in Volume I, p. 382, of the present
Memoir: "One of the most interesting results of the prolonged research in preparation for this Memoir is the
discovery of two clearly distinguishable phyla of mastodonts {TuriciUrS and Serridentinus) which by all previous
investigators both in Europe and America had been confused with the phylum Trilophodon. In the present . .

chapter [Chap. X] it is shown that the grinding teeth, e.g., of Mastodon (B.) subtapiroidea of the Lower Miocene

lignites. . . widely differ from those of Trilophodon and strongly resemble those of Serridentinus. . . [see Figs. 350,

351, 352]; it is possible that Tnricius [genotype Mastodon turicense Schinz] and Serridentinus arise from a similar
stock not unUke Serridentinus subtapiroideus. The evolution of the true Turicius phylum, however, is entirely
 NOTIOMASTOPON OR\ATUS.
ARGENTINA

OCALIENTINUS FLORLSCENS. Mongolia

JEK5LL0D0N BURNHAMI. NOTIOMASTOPON

CALIFORNIA SERRIDENTINUS ARGENTINUS.
ARGENTINA
GUATEMALENSIS,
GUATEMjALA
OCALIENTINUS
OBLIOUIDENS, TORYNOBELODON
30UTH CAROLINA BARNUI^IBROiVNI, NESKASKA

SERRIDENTIhLUS ^NGUIRIi-ALIS TORYNOBELODON

NEBRA3KA
LOOMtSI.
NEBRASKA
SERBELODON
BARBOURENSIS. NESR^SKA

OCALIENTINUS FLORIDANUS. ,-lorida

SERRIDENTINUS SERRIDENS.^
SERRIDENTINUS PRODUCTUS.
E^^ MEXICO

SERRIDENTINUS BROWNI.
TROBELODON TAOENSIS. i/t%{ '^°'^
OCALIENTINUS
NEi^ MEXICO SERRIDENTINUS
PROCHINJIENSIS. OJOCALIENSIS.
INDIA
NEVi/ MEXICO PLATYBELOCTON G/TANGERI, MONGOLIA

SERRIDENTINUS GOBIENSIS,
INNER. MONGOLIA PLATYBELODON danovi,
north caucasus

SERRIDENTINUS
SUBTAPIROIDEUS.'SERR'PENTINUS MONGOLI E NSI S, MONGOL/A
AUSTRIA 'i:^ ^ ^
A^^
SflRRIDENTtNUS ANNECTE\S. u^p>^n

I?' 0"
i5' 30' ^5* 60' 75' 9Q'E 105' 120" 155' I5Q' \kb' '30' i65" iSQ^ ijs' 120° IPS' gij'w 15" 60° PFUvtft Bradlti) t9i6/
,

Origin, Micjration and Evolution of Alrobelodon. xScrbelodon, * SerrLdentiaus.

'Ocaiientinus. uPlatybclodon. iJorynobelodon, ^Notio mastodon. Osborn 1955
PLATE XVIII
Geologic range: Trohelodon, Mio-Pliocene; Serbelodon, Lower Pliocene and Upper Pliocene; 8fm(iere(ireus, Lower Miocene to Upper Pliocene; Ocalien-
linus, Miocene to Upper Pliocene (O. obliqiddens redeposited in Lower Pleistocene). Platybelodon. LTpper Miocene; Torymbelodon, Pliocene. Notiomastodon,
Pliocene and Lower Pleistocene (?).
 —

SUMMARY 1569

distinct, leading into sharply crested forms such as T. virgatidens, while the Serridentinus phylum is (A) increasing-
ly Serridentine, ending in Ocalientinus {Ser.) floridanus, or (B) more crested, ending in S. guatemalensis." While,
therefore, there is a strong resemblance in the Miocene stages between species of Serridentinus and species of
Turicius, there is very Kttle resemblance between final species in these two phyla which are very widely divergent.

The Serridentinse are polyphyletic, embracing four genera Serrideniimis Osborn, 1923, and the Ocalientinus,
Serbelodon, and Trobelodon of Frick, 1933. Professor Osborn 's researches led him in 1935 (Osborn, 1935.937, fig.

2 —see also Vol. I, p. 729, of the present Memoir) definitely to separate these 'serrate-toothed mastodonts' (the
Serridentinse), also the Platybelodontinse {Platyhelodon, Torynobelodon) and the Notiomastodontinse (Notiomasto-
don) from all other mastodonts, thereby constituting a very large and highly diversified new family, the Serri-
dentidse, which at the present time includes no less than forty species (Vol. I, p. 741), united by the common
character of the serrated crests arising from the ectoconelets in the lower molars and from the entoconelets in the
upper molars, also by the enamel bands on the superior tusks, but generically separated by the widely radiating
adaptations of the inferior tusks —horizontal oval in Serridentinus, oval, greatly enlarged in Serbelodon, flattened

into a pair of broad shovels in Platybelodon, abbreviated and rounded in Torynobelodon, and entirely wanting, with
brevirostral mandible, in Notiomastodon.

Habits. — Members of the Serridentidse, especially the true Serridentinus, rarely occur in the same fossil
beds with members of the Bunomastodontidse, hence it is inferred that they had different feeding habits. The
more or less perfectly serrated crests on the inner side of the upper molars and on the outer side of the lower
molars point to leaf-cutting and browsing habits, especially true of the typical Serridentines, which, it is thought,
may have frequented forests and were principally browsers, while the typical flat-tuskers (Platybelodon), the
cutting-tuskers (Serbelodon), and the sub-shovel tuskers (Trobelodon) frequented shallow lakes and river borders.

Characters of Serridentinus, typical 'serrate-toothed serridentine,' 'prod-tusker' or 'scaptobelodont,'

of Eurasia and North America. Cranium low, dolichocephalic; undoubtedly rounded above like the
skulls of Trilophodon and Tetralophodon the chief difference between the skull of Serridentinus and that
;

of Trilophodon is the abbreviation of the rostrum, maxillo-premaxillary above, and symphyseal portion
of the mandible below. The symphysis in species of American Serridentinus is decurved and contains
two large tusks of flattened oval section. Jaws moderately elongate (medilongirostral), relatively stout,
tending to decurvature. Superior tusks laterally compressed, downcurved and outcurved, retaining
broad lateral enamel band. Inferior tusks without enamel band, large, horizontally flattened, closely
appressed, and never greatly elongated as in Trilophodon. Intermediate molars trilophodont from the
beginning to the end of the phylum as in the Trilophodon phylum. Grinding teeth relatively broader than
in Trilophodon, primitively brachyodont progressive to subhypsodont prominent serrated trefoil spurs
;

or crests arise from the anterior and posterior faces of the entocones in the superior molars, from the
ectocones in the inferior molars; median conules absent in the vafleys, unlike Trilophodon in which they
are present. Three superior and inferior grinding teeth (M 1-M 3) in use at the same time; functional
retention and usage of upper and lower second and third molars, in contrast to the Longirostrinse
{Trilophodon) and Tetralophodontinae (Tetralophodon) in which the mastication is finally concentrated on
the third upper and lower molars. Internal cingulum more or less crenulate on superior molars, extend-
ing as far back as the antero-internal border of third crest of the third molars; rudimentary external
cingulum only in valley between first and second crests (S. productus).
Dental formula: I H^ Dp |^ P H M i^^
Ridge-crest formula: Dp 2^ Dp 3| Dp 41 P 3^ P 4f± M II M 21 M 3^*?^
Horizon. — Lower Miocene to Upper Pliocene. Compare Vol. II, p. 1535, and figures 1221-1224,
1227, also PI. XVIII.

Characters of Ocalientinus, 'multiserrate-crested serridentine,' 'prod-tusker' or 'scaptobelodont,' of

the United States and Mongolia, directly ancestral to 'Serridentinus' republicanus of Kansas and to 'S.'
floridanus of Florida. Cranium tall, narrow. Mandible elongate, symphysis extremely narrow with deep
 — :

1570 OSBORN: THE PROBOSCIDEA

superior longitudinal channel. Superior tusks as in Serridentinus, with latero-inferior enamel hand.
Inferior tusks without enamel, upcurved, relatively short. Molars with roundly blunted cones and
conelets, trefoil spurs dominant, florescent. Relative breadth of molars correlated with foreshortening
of cranium and heightening of occiput. Both superior and inferior ridge-crests sloping forward, slowly
increasing in height.

Ridge-crest formula: Dp 4§ P 3- P 4t M If M 21 M 3,,!^^

Horizon.— Miocene to Upper Pliocene (0. obliquidens redeposited in Lower Pleistocene). (Compare
Vol. II, p. 1536, and figures 1223, 1224, 1227, also PI. xviii.

Characters of Serbelodon, Frick's 'sub-shovel-tusked serridentine' of the western United States.

Cranium imperfectly known, probably more elongate than that of Trohelodon. Mandible extremely
abbreviated and broadened, the length equaling that of tierridentinuH productus. Symphysis short,
sym))hyseal rostrum extremely short and relatively broad. Superior tusks heavy, downwardly and
outwardly curved, with strong enamel band. Inferior tusks concavo-convex, relatively short, flattened,
heavy, spatulate; distinguished from Platybelodon by total absence of dentinal rod-cones, the interior
being composed of dentinal lamina^. Molars relatively long and narrow with serridentine pattern
superior molars with entoconelet spurs, inferior molars with ectoconelet spurs, but without serrated
borders.

Ridge-crest formula: M 21 M 31
Horizon. — Lower Pliocene, Upper Pliocene. Compare Vol. II, p. 1536, and figures 1224, 1227,
also PI. XVII I.

Characters of Trohelodon, Frick's 'brevidentate serridentine' of New Mexico, of 'sub-shovel-tusk'

type. Cranium broad or brachycephalic. Mandible relatively decurved. Symphysis relatively elongate,
narrow, slightly upcurved. Superior tusks with moderate enamel band inferior tusks moderately heavy,
;

biconvex, pointed, absence of dentinal rod-cones. Third superior molars short and broad, with blunt
cones; expanded entotrefoils as in Serridentinus; third inferior molars also short and broad, with ex-
—
panded ectotrefoils trefoil structure of primitive serridentine type.
Ridge-crest formula: M 21 M 31
— Mio-Pliocene.
Horizon. Compare Vol. II, p. 1537, and figures 1224, 1227, also PI. xviii.

Migration. — The followingis the geographic distribution of species of the Serridentinge, the trend of mi-
gration being eastward from southwestern Euroj^e to North America (see PI. xviii, also Figs. 373 and 1227)
France, Austria, India, Mongolia, China, Japan, Califoinia, Nebraska, Kansas, Colorado, New Mexico, Texas,
Guatemala, and eastward in the United States to North and South Carolina and Florida.

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: SIORRIDENTID^ Osborn, 1935, 1936

Subfamily: Platybelodontin^ Borissiak, 1928
Genera: Platybelodon Borissiak, 1928, 1929; Torynobelodon Barbour, 1929
(Cf. Vol. I, pp. 338, 339, 459-472, 729, 730, also Pis. x, xi, and Figs. 344, 373; Vol. II, p. 1537, and figures 1222,
1223, 1227, also PL xviii)

The subfamily Platybelodon tina? was founded by Borissiak in 1928 on his Platybelodon danovi of the Chokrak
beds, Kuban district. North Caucasus, the chief distinguishing character being the long and wide spoon-shaped
symphysis with wide, flat incisors. Prior to this description, and wholly unknown to Professor Borissiak, a speci-
men found in Nebraska had been described by Dr. E. H. Barbour as Amebelodon fricki in 1927 and made the type
of his family Amebelodon tidae (subsequently, 1929, modified to Amebelodontinae). Professor Osborn assumed
(1931) that the Amebelodon tidir Barbour, 1927, preoccupied the Platybelodon tina? Borissiak, 1928, especially
as both po.s.ses.sed large shovel tusks supposedly penetrated by dentinal tubules or rod-cones; also at the time he
was not aware of the highly progressive serridentine pattern of the Platybelodon molars, in widest contrast to the
trilophodont pattern of the Amebelodon grinders, or of the fact that the second superior and inferior molars
possessed four ridge-crests, unlike Trilophodon and members of the subfamily Serridentinae with three ridge-crests.
 :

SUMMARY 1571

As stated above under the subfamily Amebelodontinae (p. 1558), the structure of the incisive tusks of Amebclo-
don and of its ancestral form Phiomia is that of progressively concentric dentinal laminae, an observation made

in 1934, 1935, when the inferior tusks of Phiomia were sectioned and carefully compared with those of Amebelodon.
Likewise the inferior tusks of Platybelodon were sectioned, revealing compact interior dentinal rod-cones instead of
concentric dentinal laminae also the tusk of the American related genus Tonjnobelodon was found to be composed
;

of rod-cones, resulting in the reference of this genus to the Platybelodon tinse.

Quoting Dr. Edwin H. Colbert in Volume I, p. Memoir, "It would seem possible, then,
715, of the present
that the presence or absence of these rods distinguishes the two subfamilies Platybelodontinae and Amebelodon tinae
respectively ... It would seem likely, also, that in the Platybelodontinae the tusks are square across the ends,
whereas in the Amebelodon tinae the ends of the tusks are rounded. The origin of the rods constitutes a perplexing
. .

question. To me [Colbert] they appear to be derived from the lamellar ivory; perhaps they represent a secondary
breaking down of the concentric cones of dentine that form the tusk. Perhaps they are structures developed in
the pulp cavity independently of the lamellar ivory. . . A magnified cross-section of a dentinal rod of Torynobelodon
loomisi shows that it is built up of concentric layers of dentine—presumably unvascular orthodentine. An
examination of the tusk of T. loomisi shows that the rods occupy the central portion of the tusk, and that they are
surrounded by typical ivory. The rods are quite variable as to size, ranging from one to five millimeters in di-
ameter. Many of the rods branch. There are numerous nodules or egg-shaped excrescences on the sides of various
rods, and on some rods the external surface is granular. The rods are of varying lengths. The rods are 'bundled'
together, so to speak, in the tusk, but they are not compressed; consequently they are round and not hexagonal.
The interstices between the rods were seemingly empty ; if there was a filhng of dentine it has been destroyed
during the process of fossihzation. In many places the rods seem to grade into the lamellar ivory."
Characters of Platybelodon, Bori.ssiak's typical 'flat-tusker' of Mongolia. Cranium broad, low.
dolichopic. Mandible with long and wide spoon- or shovel-shaped symphysis extreme constriction and ;

narrowing of mandibular groove (according to the theory of Dr. William K. Gregory, 1934, the lower sur-
face of the tongue occupies this narrow groove which broadens out into the deeply hollowed suprarostral
concavity). Low and massive rami (type). Superior tusks small, rounded, relatively short and slightly
outturned and downturned inferior tusks wide and flat, with regularly and sharply defined borders, com-
;

posed of numerous and quite closely compacted dentinal rod-cones. Upper surface without enamel, with
irregular longitudinal grooves, lower surface protected by enamel [?dentine] longitudinally striated {fide
Borissiak). Anterior serration of the entocones in the superior molars and of the ectocones in the inferior
molars, as distinguished from the double or anterior and posterior serration of the corresponding ento-
and ectocones in the Serridentinus and Ocalientinus molars. Fourth superior and inferior deciduous
premolars and second molars with four ridge-crests instead of three ridge-crests as in Trilophodon,
Amebelodon, and members of the subfamily Serridentinae, resembling Tetralophodon in this respect.

Dental formula: Di |^ I H^ Dp |f| P |^ M 3-4 -^'-^

1-3
1-3

Ridge-crest formula (based on complete collection of P. grangeri in American Museum)

DplvDp2 „,X;::f"ot. Dp3,^Dp4iP3^:Hlg^P4tMlf^M2,.kM3.|^|^
Horizon.— Upper Miocene. Compare Vol. II, p. 1537, and figures 1223, 1227, also PI. xviii.

Characters of Torynobelodon, Barbour's 'dredge-tusker' of Nebraska. Cranial characters not yet

described by Doctor Barbour. Mandible thin, ladle shaped, with low ascending rami; abbreviated as
compared with Platybelodon grangeri symphysis relatively narrow with relatively broad bar between
;

rostrum and rami. Tusks short, broad, ladle or spoon shaped, upcurved, roughly and deeply corrugated,
composed of dentinal rod-cones.

Dental formula incompletely known : I „Tir:^ M ^r^

Ridge-crest formula incompletely known : M 23 M 3 ^^
Horizon. — Pliocene. Compare Vol. II, p. 1537, and figures 1224, 1227, also PI. xviii.
 :

1572 OSBORN: THE PROBOSCIDEA

Migration. — Up to the present time members of this subfamily have been found in the North Caucasus
and in the Gobi of MongoUa. A left lower incisor from Texas, in the Colorado Museum, may be referable to
Platybelodon also Torynobelodon, which appears to be a collateral
; branch, and somewhat less perfect mechanically,
of the Platybelodonts of Central Asia, comes from the Pliocene of Nebraska.

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: SERHIDENTID^ Osborn, 1935, 1936

Subfamily: NoTioMASTODONTiNyE Osborn, 1936
Genus: N otiomastodon Cabrera, 1929

(Compare Vol. I, pp. 541, 590-592, 691, 730, 731; also Pis. x-xii, and Fig. 501; Vol. II, p. 1537, and figures

1225, 1227, also PI. xviii)

The third subfamily of the Serridentidse is the Notiomastodontinse Osborn, based on the N otiomastodon
omatus of Cabrera, 1929. The reasons by the present author for his reference of this subfamily and
set forth

genus to the Serridentidse will be found on page 730 of Volume I of this Memoir, and are briefly as follows

The inferior molars have prominent serrated ridges arising from the ectoconelets as in Serridentines, and the
adjacent mesoconelets are reduced or wanting, in contrast to Cuvieronius and Cordillerion of South America, in
which the trefoU spurs arise from the mesoconelets; it is radically different from the fundamental molar pattern
of any of the Bunomastodontidse, the trefoil structure of which invariably rises from the mesoconelet or is in close
connection with it through the central conules. Also evidencing the Serridentine relationship are the laterally
compressed, downcurved superior tusks, with broad external enamel band, all trace of which band is absent in the
contemporary Cuvieronius of the Pampean region.

Characters of N
otiomastodon, Cabrera's 'short-jawed serridentine' of the Pampean. Mandible
abbreviated, typically stout, lacking inferior tusks; symphysis strongly abbreviated with a transversely
concave groove for the tongue; coronoid small and condyle elevated. Superior tusks down- and out-
turned, with broad enamel band as in Serridentines and Trilophodonts. Inferior molars bunolophodont,
with prominent serrated ridges arising from the ectoconelets as in the Serridentines; mesoconelets re-
duced or wanting. Double series of trefoils on wear (genotype).
Ridge-crest formula: M I3-+ M 2j+ M 87^1^)
Horizon. — Pliocene and Lower Pleistocene(?). Compare Vol. I, Pis. x and xi; Vol. II, p. 1537,
and figures 1225, 1227, also PI. xviii.

Migration. —The Notiomastodonts seem to have been confined to South America, as the genotype, Notio-
mastodon omatus from Buenos Aires, and a tusk from the valley of the Tarija River, described by Ameghino in
1888 as Mastodon argentinus, referred bj' the present author to N otiomastodon, are the only specimens recorded up
to the present time.

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: MASTODONTID^ Girard, 1852

(syn. in part MAMMUTID^ Cabrera, 1929)

Subfamily: Pal^omastodontin^e Osborn, 1936
Genus: Palaeomastodon Andrews, 1901
(Cf. Vol. I, pp. 138, 139, 143-149, 689, 691, 692, also Pis. i, 11, x, xi, and Figs. 24, 86, 123a; Vol. II, p. 1529,
and figure 1220, also PI. xix)
The theory that Moeritherium gave rise to Palxomaslodon, and that Palaeomastodon gave rise to Mastodon
angustidens, generally so interpreted from Dr. C. W. Andrews' Memoir of 1906, was not accepted by Profe.ssor

Osborn, who, in 1909, even went so far as to separate Moeritherium from direct relationship to the Proboscidea.
This opinion, however, he afterward (1921) modified, in fact, he definitely regarded Moeritherium as constituting
 MASTODON ICUTIDENS. .Jar. irvol^HA

t^
MASTODm AMgRICANUS, N. a^ierica

STEGOLOPHODON STEGOOONTOIDES. India

PLIOMASTODON VEXILLARIUS, California

STEGOLOPHODON LYDEKKERI, BOKNCO

TURICIUS URGATIDENS, gekmavy

STEGOLOrHODON LATIDENS.

M/OMASTOPON TAriROI DES AMERICANUS, HUnaAKf

STEGOLOPHODON CAUTLE1 1 PKOGRESSUS,
INDIA

PALAEOMAStODON BEAJWELLI, fayOm Ch J^

TURICIUS TURICENSIS. SniTZCRiAND

MIOMASTODON MERRIAMI, NEfAOA
PAL A£0M4STOP0N_JN TERMED/US.

f)

PALAEOMASTODON PARyUS. FAYUM

FA-yUM

m
MIOMASTOnO/^ DEPERETI, FRANCE TURICIUS TAPIK0I11£S, FRANCE

Origin. Migration and Evolution of •Paldeo mastodon, a. Mio mastodon, a Plio mastodon,
* Mastodon. iJliricLUs, xZi/^oLophodoa. ^Ste^olophodon. Osborn 1955

PLATE XIX
Geologic range: Palsomaslodon, Lower Oligocene. Mioinaslodon, Lower to Middle Miocene, and Lower Pliocene; Plioniastodon, Lower to late Pliocene;
Mastodon, early to late Pleistocene. Turicius, Lower Miocene to Middle(?) Pliocene; Zygolophodon, Lower Miocene to LTpper Pliocene. Stegolopkodon,
Mio-Pliocene to Upper(?) Pliocene (Lower Pleistocene?].
 SUMMARY 1573

one of the great primary stocks of the Proboscidea, namely, the Moeritherioidea. Doctor Andrews also (1909),
while still regarding Moeritherium as a proboscidean, reconsidered the possibility of its not being in the direct line
of ancestry of Palxomastodon; and Doctor Matsumoto (1924), while placing Moeritherium in a side line of
its own, concluded that Palxomastodon appeared nearly to correspond to a theoretical ancestry of the

Zygolophodon-Mastodon phylmn, thus suggesting that it was related to the subfamily which the present author
designated as the Mastodontinse. This theory was immediately approved by Professor Osborn, although it must
not be overlooked that in 1923 he had clearly separated the true Mastodon americanus from the Zygolophodon of
Vacek, at which time he described a new subfamily, the Zygolophodon tinse.

In 1934 Professor Osborn became convinced that Palaeomastodon should be removed from direct ancestry to
the true Mastodon, because he observed that the progressive third superior and inferior molars of P. intermedius
exhibited the presence of proto- and metaconules, thus blocking the median sulcus characteristic of all the Masto-
dontinse. On page 139 of Volume I of the present Memoir we find the following: "In the carefully drawn Plates
I, II, III, and IV, illustrating the evolution of the grinding teeth in Mastodon, Zygolophodon, Turicius, and Stegolo-
phodon, a median longitudinal sulcus separates the external and internal cones both in the superior and inferior
molars. This demonstrates that the ancestral proboscidean molar was tetrahunodont, as in Moeritherium, not
hexabunodont, as in Palxomastodon. . . By close comparison of all the figures of the upper and lower grinding teeth
of Palseomastodon, Miomastodon, PUomastodon, and Mastodon, it is observed: (1) That the molar crowns in
Palaeomastodon are mainly tetrahunodont, i.e., each protoloph (superior) and each protolophid (inferior) is com-
posed of a main external bunoid cone and a main internal bunoid cone; in the superior molars (Fig. 94D) where
the conules persist the main crown is hexabunodont. The presence of proto- and metaconules blocking the
median sulcus forbids the ancestry of Palseomastodon to Mastodon. (2) The vestigial intermediate protoconules
and metaconules are observed in the hexabunodont superior molars of Palaeomastodon intermedius, thus the
crested upper grinders are hexabunodont or six coned, whereas the lower grinders are subtrilophodont (Fig. 93,
M^-Mg, Fig. 94). This primitive condition of the cones connects Palaeomastodon with its undiscovered sexituber-
cular-quadritubercular ungulate ancestors ; the conules observed in the third superior molar, M^, of P. intermedius
(Figs. 93 and 89) are not seen in M^ of the same species (Fig. 92) ; the conules are vestigial or disappearing struct-
ures."

Accordingly Professor Osborn withdrew Palaeomastodon from the subfamily Mastodontinse and created a new
subfamily, Palseomastodontinse, for its reception (see Vol. I, Appendix, p. 691).

Palxomastodon, the 'ancient mastodon' of North Africa, Eocene-Oligocene. Skull imperfectly

known. Palate relatively short and broad. Mandible elongated anteroposteriorly (medium jawed)
mandibular symphysis relatively short as compared with Phiomia. Superior tusks unknown. Inferior
tusks rounded and relatively abbreviated. Molars brachyodont, of bunolophodont rather than of buno-
mastodont type, progressive in size and evolution from tetrahunodont cones {Palaeomastodon parvus)
to hexabunodont cones [P. intermedius, P. beadnelli); bilophodont superior molars, subtrilophodont in-
ferior molars, in other words, the tritoloph and tritolophid are in a formative stage, tritoloph rudimen-
tary even in M^. Intermediate proto- and metaconules progressively developing to perfect the transverse
crests, thus closing the median longitudinal sulcus characteristic of all the Mastodontinse. Molars exhibit-
ing central conules; trefoils rudimentary or absent. Basal cingula neither very strong nor very rough.

Dental formula: I^:^ C § Dp P | |^ M 1-3

Ridge-crest formula: P 2^ P 3\^ P 41 M 1^' M 2 "3^^

M 3f
Horizon. — Lower Oligocene (Fluvio-marine formation). Compare Vol. II, p. 1529, and figure
122U, also PI. XIX.

Habits and Habitat. — Probably forest loving with transversely secant molar crests for cutting herbage.
Up to the present time, however, specimens of Palxomastodon have been found in the Fayuni region only.
 :

1574 OSBORN: THE PROBOSCIDEA

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: MASTODONTID^ Girard, 1852

(S>Ti. in part MAMMUTID^ Cabrera, 1929)

Subfamily: MastodontiNvE Brandt, 1869— Osborn, 1910

Genera: Mastodon Cuvier, 1806-1817; Mzomastodon Osborn, 1922; Pliomastodon Oshorn, 192Q

(Cf. Vol. I, pp. 132, 138, 150-190, 690, 693-697; also Pis. i, x, xi, and Figs. 86, 123a, 123b; Vol. II, p. 1530,
and figures 1222, 1224, 1227, also PI. xix)
In Professor Osborn's article in Natural History of 1925 (Osborn, 1925.637), we find on page 10 the follow-
ing statement: "The true mastodons of our American forests appear to have arisen from the diminutive Palseo-
mastodon of the primordial river Nile. The reason these animals have left no trace of their 10,000-mile and
2,000,000-year journey from the Nile region to the forests bordering the Ohio and the Hudson rivers is that

fossilization of forest-living fauna has always been rare. The ancestral Palseomastodon of the Nile region is itself

very rare. . . not even fossilized teeth of this race were scattered in Europe to show the route."

During the ten-year interval from 1925 to 1935, Professor Osborn changed his views regarding Palxomastodon
being the ancestor of the true Mastodon, as expressed in the foregoing paragraphs (p. 1573) on the Palseomastodon-
tinse and also on pages 138 and 139 of Volume I of the present Memoir: "While we are certain that the true Pal-

xomastodon possesses a longer jaw but a broader and shorter skull than Lower Oligocene contemporary Phiomia,
its

while the superior grinding teeth are comparatively short and broad, and while in the crowns of the superior and
inferior grinding teeth the proto- and metaconules forbid the ancestral relationship of Palseomastodon to Plio-

mastodon, Miomastodon, and Mastodon, we must await further knowledge of the cranium and of the cutting teeth,
also of the incisors of Palxomastodon, before we can form a positive opinion on this very important and interesting
question." Again, in the Appendix to Volume I, p. 692, he voices uncertainty as to the ancestral forms of the
Mastodontinse which embraced the four genera, Palseomastodon, Miomastodon, Pliomastodon, Mastodon, until he
withdrew Palseomastodon, making it the type of a new subfamily Palseomastodontinse : "Despite the extreme
rarity of the remains of forest-loving and browsing mastodonts, exemplified by the rarity of Palseomastodon as

compared with Phiomia in the Fayiim deposits, several discoveries of isolated grinding teeth have been made in

the Tertiary deposits of Europe and North America which are now recognized as belonging to successors of still

unknown true Mastodontina? of Africa and ancestors of the true Mastodontina? of North America."

The contrasting characters, so far as known, of the three genera now constituting the Mastodontina>, are as
follows

Mastodon, 'typical mastodon' of Eurasiatic and North American forests. Cranium brachycephalic,
brachyopic. Mandible and symphysis abbreviated; progressive reduction of rostrum. Superior tusks
large, rounded, upturned, with slightly indicated annular ring growths; inferior tusks straight, cylindri-
cal, variable in old age; no enamel. Alarked sexual disparity in female tusks. Grinders relatively elon-
gate, bilobate, with strong median sulcus between inner and outer lobes; no conules; rudimentary tre-
foil spurs on superior iimer lobes, on inferior outer lobes; summits of lobes simple or crowned with
small conelets; extreme binary fission of summits of crests in Mastodon acutidens, in which as many as
twenty-one conelets may be counted; progressive plication of the surface enamel. Ridge-crests lopho-
dont, [)rogressively elevated, subhy[)sodont. Progressive development of third crest (tritoloph and
-lojjhid) in intermediate molars; progressive development of fifth crest rudiment (pentalophid) in
third inferior molars. Permanent |)rem()lars suppressed, except P*-P4, which form in the jaw but do not
erupt. Cingulum slightly stronger on inner side than on outer side.

Dental formula: I
ti:Dp l\ P (vestigial)
;;;;;:; M 1-:)

Ridge-crest fornuila: Dp 21 Dp 3* Dp 4i} M U M 25 M S"^

Horizon.— Early to late Pleistocene. Compare Vol. II, p. 1530, figures 1222, 1224, 1227, and PI. xix.
 SUMMARY 1575

PUomastodon, 'Pliocene mastodon' of Eurasia and North America, an ancestral stage of Mastodon.
Distinctive, low browed, flat crested; abbreviated postnarial region and broadly divergent premaxillary
region; approximation of condyles to superior molars (M'^), extreme backward recession of anterior
nares and shortening of lower jaw, probably also an unusual width of the occipital crest, suggesting an ex-
ceptionally large development of the proboscis (characters based on PUomastodon vexillarius Matthew,
—
1930 see Vol. I, pp. 161-163). Superior tusks suboval to rounded in section, upturned, enamel band
vestigial or wanting; a "narrow strip of thin enamel" within the alveolar base of the tusk in P. vexillarius.
Inferior tusks probably stout (as inferred from the alveoli —
about 50 mm. in diameter near the posterior
—
end in P. sellardsi Simpson, 1930). Molars intermediate in hypsodonty between Miomastodon and
Mastodon; a distinctive feature is the posterior narrowing of the crown of the third superior molars
(P. mattheioi). Ridge-crests with rudimentary trefoil spur extending from entocones of superior molars
and from ectocones of inferior molars; expanded ectotrefoils (P. matthewi). No median conules. Persis-
tent longitudinal sulcus also persistent four conelets.
;

Dental formula incompletely known : I ~|^ Dp | P | MI

Ridge-crest formula: P 4|| M 1| M 21 M 3y^
Horizon. —Lower to late Pliocene. Compare Vol. II, p. 1530, and figures 1222, 1224, 1227, also PI.
XIX.

Miomastodon, 'Miocene mastodon' of Eurasia and North America. Skull and skeleton unknown.
Mandible and symphysis moderately elongated; rostrum laterally compressed, abbreviate as compared
with Trilophodon or Serridentinus. Superior tusks downturned, broadly oval in vertical section;
broad enamel band on concave external surface (Miomastodon merriami), on convex external surface
(M. tapiroides americanus); inferior tusks rodlike, horizontal, vertically oval. Molars brachyodont to
subhypsodont; internal and external lobes or cones of each loph separated by median or longitudinal
sulcus; summit of each lobe or cone double or bifid, rounded; mesial expansion of rudimentary trefoils
in ento- and ectoconelets, upper and lower; central conules absent; retarded binary fission of cones and
conelets in pro to- and metalophs conelets never exceeding four in each loph or transverse ridge-crest.
;

Persistent median sulcus.

Dental formula incompletely known : I l',"^ Dp y P f M JtI

Ridge-crest formula: P 41^ M 1|: M 2f M 3^

—Lower to Middle
Horizon. Miocene, Lower Pliocene. Compare Vol. II, p. 1530, and figures
PL xix.
1222, 1224, 1227, also

Migration. — Vol.
(Cf.pp. I, 132, 133): "Springing from unknown ancestors of the Eocene-Oligocene of
North Lower Miocene Miomastodon [M. depereti] of western Eurasia
Africa, the true Mastodontinae appear in the
[France] and soon find their way eastward into North America, appearing in the Middle Miocene species of
Nevada, Miomastodon merriami, and in the Lower Pliocene species of western Nebraska, PUomastodon matthewi,
meanwhile leaving behind in Austria-Hungary their relatives, Miomastodon tapiroides americanus and PUomastodon
americanus praetypica, which give rise to the rare true Mastodon of southern Russia, referred to 'Mastodon ohioti-
cus' by Pavlow [and by the present author to Mastodon pavloun]. Rarely found in northern and western North
America, they multiply rapidly in the favorable forests of the middle and eastern United States in the typical
form Mastodon americanus." Fossil remains are found even as far northeast as Nova Scotia and New Brunswick,
as evidenced by a recent (1936) discovery at Hillsboro, N. B., and identified by Director William Macintosh of
the New Brunswick Museum as Mastodon americanus.
Superfamily: MASTODONTOIDEA Osborn, 1921. Family: MASTODONTID^ Girard, 1852
(Syn. in part MAMMUTID.E Cabrera, 1929)
Subfamily: Zygolophodontin^ Osborn, 1923
Genera: Zygolophodon Vacek, 1877; Turicius Osborn, 1926
(Cf. Vol. I, pp. 191-223, 282, 690, 697-699, also Pis. ii, iii, x, xi, and Figs. 137, 159; Vol. II, p. 1530, and figures
1222, 1227, also PI. xix)
When in 1923 Professor Osborn created his subfamily Zygolophodon tinae, he had not observed the pronounced
differences in the molar structure of species of Zygolophodon Vacek. 1877, which later (1926.706, p. 3) actuated him
1576 OSBORN: THE PROBOSCIDEA

to describe a new genus, Turicius. Especially noticeable were (1) the strong subdivision of the main cones of
certain species {Zygolophodon) into from four to six subequal conelets, whereas in other species {Turicius) the
cones subdivided into from five to nine conelets, and even into twenty-five (see PI. in, pp. 134, 135) ; also (2) the

presence of strong 'trefoil spurs' on the supero-internal and infero-external cones of the latter (Turicius) in contrast

to the absence (or vestiges) of these spurs in the former (Zygolophodon) . The species included within Turicius
show several resemblances to those included within the genus Serridentinus (e.g., the trefoil spurs just mentioned),
but in many other characters Turicius is clearly separable from Serridentinus and more closely related to Zygolo-
phodon.
The history of the Zygolophodontinse will be found in detail in Volume I, (chapter VII, covering the period
from 1770 to 1936, in which is set forth the grounds on which this subfamily is separable from the Mastodontinae,
to which it is most nearly related. In brief, the median longitudinal sulcus (Fig. 89) of Mastodon americanus
disappears in Zygolophodon and transversely perfected crests evolve ; in Mastodon the main cones feebly subdivide
at the summits into two conelets,' progressive to four (Miomastodon) whereas in Zygolophodon there are four to
,

six main conelets, and from five to nine and even twenty-five in certain specimens of Turicius. In other words,
from the principal characters of the type grinding teeth we observe (cf. Vol. I, p. 199) "that the universal generic
character both of Zygolophodon and of Turicius is the absence or reduction of the longitudinal sulcus, fissure, or
commisure, which Hays pointed out as the chief character distinguishing his type (Fig. 154) of M. [Zygolophodon]
borsoni from M. americanus. The absence or reduction of this sulcus distinguishes all the ascending species of
Zygolophodon and of Turicius, from early Miocene to late Pliocene or early Pleistocene time, from all the ascending
or ancestral stages of Mastodon in the corresponding period. We also observe that the fission of the cones into
'conelets' is rapidly progressive and distinct in Zygolophodon and in Turicius, whereas it is decidedly retarded in
Mastodon. [The final stage of evolution, however, in the Mastodon acutidens molar (Fig. 656) converges towards
the final evolution stage in the zygolophodont molar (Turicius virgatidens — Fig. 168).] In many other features . . .

the progressive dental and skeletal evolution of Zygolophodon parallels that of Mastodon (1) the abbreviation of
the jaws, (2) the suppression of the premolar dental succession, (3) the reduction of the lower incisive tusks, (4)
the progressive abbreviation of the cranium. . . . From the fact that the Zygolophodon molars are relatively broader
and shorter than those of Mastodon, it is probable that the cranium of Zygolophodon was more brachycephalic than
that of Mastodon."

The collateral relationship of the Zygolophodontinse to primitive species of Stegodonts (Stegolophodontinae,

Stegodontina?) of Asia is Volume I, p. 195, and
treated in in Chapter XIV of the present volume, p. 819. Such
relationship was first adumbrated by Schlosser as early as 1903, and was followed in 1917 by the suggestion of
Schlesinger that primitive European species may have given rise to primitive species of Stegodonts of Asia to
which he gave the new generic name Stegolophodon —both of which suggestions Professor Osborn was inclined to
favor. An excellent columnar presentation of the outstanding characters of the grinding teeth of the Mastodon,
Zygolophodon, Turicius, and Stegolophodon phyla is given on page 211 of Volume I, supplemented by Plates i-iv

(pp. 134-135) demonstrating the molar evolution. Reference is made in the last column to a description of the
collateral ancestor of Stegodon in Chapter XV; this should read Chapter XIV, as in revising the present volume
it was found desirable to rearrange the chapters somewhat (see pp. 819-822 above).

Professor Osborn made a further suggestion on page 202 of Volume I, to the effect that, owing to certain
strong resemblances (e.g., supero-internal and infero-external trefoil spurs) between the grinding teeth of Turicius
and Serridentinus, despite the entirely different lower tusks in the two genera, there might be an affinity of the

'[Sec, however, Mastodon acutidens Osborn (described on pj). 096, 697 of Vol. I of tlie present Memoir) in which as many as twcnty-onc oonolcts may be
—
counted on certain of the lophs. Editor.]
 :

SUMMARY 1577

genus Turicms to Serridentinus which a fuller knowledge would demonstrate. Among the distinguishing charac-
ters, in addition to the differences in the tusks, he mentions (p. 223) the development of a pure transverse crest
in the molars of Turicius, as against the development of corresponding conelets and crests in the American
Serridentinus in which the cones and conelets do not become connate and remain quite distinct at the summit,
also the progressive strengthening of the trefoil spurs in Serridentinus, which become more and more prominent
and conspicuous in progressive stages, surmounted with small conelets, whereas in progressive stages of Turicius,

Uke T. atticus and T. virgalidens, the trefoil spurs are reduced to fine ridges accented with numerous small conelets.
See, however, his final conclusions on pages 1568, 1569 above, namely, that the Turicius phylum is entirely

distinct.

A condensed summary of the contrasting characters in the two genera Zygolophodon and Turicius is as follows

Zygolophodon, typical 'yoke-crested mastodont' of Eurasiatic Miocene to Pleistocene. Cranium and

skeleton relatively unknown, probably similar to Mastodon americanus. Symphysis more abbreviated
than in Turicius. Superior tusks rounded, enamel band disappearing early, before Lower Miocene time.
Inferior tusks reducing early; no enamel (Z. pyrenaicus). Grinders permanently blunt, brachyodont
(Z. borsoni), not becoming subhypsodont. Ridge-crests or lophs directly transverse, not arched, each
loph divided into four to six distinct subequal conelets. Conules or tubercles in median valleys disappear-
ing in early stages (vestigial conules in anterior valley of Z. pyrenaicus). Trefoil spurs vestigial or absent.
Median longitudinal sulcus disappearing, primitive, vestigial. Fifth inferior crest, i.e., pentalophid,
slowly progressive. Intermediate molars trilophodont {fide Schlesinger). Suppression of the premolar
dental succession.

Dental formula: I ^l Dp 1:1 M -3

{:
1-3

Ridge-crest formula: Dp 21 Dp 3| Dp 41 M H M 2|i M 3|^

Horizon. —Lower Miocene to Upper Pliocene. Compare Vol. II, p. 1530, and figures 1222, 1223,
1227, also PI. XIX.

Turicius, sharp-crested 'Zurich mastodont' of the European Miocene and Pliocene. Cranium and
skeleton relatively unknown. Symphysis progressively pointed and reduced in length, horizontal not
decurved. Superior tusks oval in section, rodlike, with sharply defined enamel band except in Turicius
mrgatidens. Inferior tusks without enamel, straight, suboval in section, undergoing progressive reduction.
Molar ridge-crests sharply transverse and elevated, with conelets progressive from four to twenty-five.
Median longitudinal sulcus vestigial in early stages, completely disappearing in progressive stages. No
median conules. Progressively strong trefoil spurs on superior internal cones and on inferior external
cones. Conelets increasingly connate at summit, rising into sharp, subhypsodont transverse crests.
Postero-inferior molars with four well-developed crests, the tetartolophid slowly progressive, the pentalo-
phid rudimentary but progressive in the higher stages. Gradual repression of the premolar dental suc-
session, as in Mastodon americanus and Zygolophodon borsoni.

Dental formula: Di ^^ I tXl ^P B ^ B

Ridge-crest formula: Dp 2^^+ Dp 3f± Dp 41 M II M 2f± M 3^
Horizon.— Lower Miocene to Middle(?) Pliocene. Compare Vol. II, p. 1536, and figures 1220,
1222, 1227, also PI. xix.

Migration.— The types oi Zygolophodon and Turicius, as known up to the present time, occur in southern and
central Europe. As will be seen by referring to figure 159 and Plate xix, a referred specimen of Turicius
comes from North Africa, also, according to Matsumoto, one of Zygolophodon from Japan. Recently (cf. pp.
698 and 699 of the present Memoir) Doctor Hopwood has described a juvenile mandible regarded by him as

belonging to Mastodon americanus but according to Professor Osborn referable to Turicius, also a molar of Zygolo-
phodon borsoni, both from China. Zygolophodon borsoni is relatively abundant in Siberia. Again consulting
figure 159 and PI. xix, it would seem that the trend of migration was from North Africa, through southern and
central Europe eastward to Siberia, China, and Japan. The Zygolophodontinse apparently never reached
America.
 ;

1578 OSBORN: THE PROBOSCIDEA

Superfamily: MASTODONTOIDEA Osborn, 1921. Family: MASTODONTIDiE Girard, 1852

(Syn. in part MAMMUTID^. Cabrera, 1929)

Subfamily: Stegolophodontin.e Osborn, 1936

Genus: Stegolophodon Schlesinger, 1917

(Gf. Vol. I, p. 700, Pis. IV, XI and Fig. 661; Vol. II, Chap. XIV, p. 837, also p. 1531, and Figs. 691, 1221, 1223,
1228, PI. xix)

The subfamily Stegolophodontinse, type Stegolophodon Schlesinger, 1917, was defined by Professor Osborn
as "transitional in molar tooth structure between the Mastodon toidea and the Stegodontoidea" — the latter

a superfamily first by him to embrace both Stegolophodon and Stegodon (see Osborn, 1935.937, fig. 2). On
created
further study, however, the median sulcus characteristic of the Mastodontinse was found to persist in the anterior
ridge-crests of Stegolophodon, thus pointing toward a very remote relationship between the Mastodontoidea and
—
the Elephantoidea indeed, a comparison shows that the most primitive elephantoid molar known, that of
Archidiskodon proplanifrons of South Africa (subfamily Mammon tinae), is, in section, very like the molar teeth of
a Mdfitodon (subfamily Mastodontinse), that is, without trefoils or central conules blocking the valleys. Professor
Osborn, therefore, became convinced that Stegolophodon should be removed from the Elephantoidea and placed as
a terminal member of the Mastodontoidea, leaving the genus Stegodon as the sole generic representative of the
superfamily Stegodontoidea. The reader will find a detailed account of the phylogeny of the Stegolophodonts and
Stegodonts in Chapter XIV of the present volume, wherein they are treated together according to Professor

Osborn's original views, as unfortunately he had not reached this chapter in his final revision. We may summarize
the distinctive characters of Stegolophodon, as follows:

Stegolophodon, 'roof-crested i)ro-stegodont' parallel and collateral in evolution with the true
Stegodon of eastern and southern Eurasia. Cranium low, dolichocephalic, suggesting the primitive
type of skull and tusks seen in Trilophodon and Serridentinus. Mandibular symphysis not certainly
known, but probably short and tuskless. Superior tusks straight with broad enamel band. Inferior
tusks undiscovered. Lophs as in Mastodon and Zygolophodon, tendency to form from four to six trans-
versely arranged cones and conelets (conelets somewhat irregular) and to consolidate into ridge-crests;
molar pattern transitional between the Zygolophodon type and the Stegodon type; no median conules
and no trefoil spurs. Closure of enamel in base of transverse valleys, that is, V-shaped enamel thick ;

cement usually absent.

Dental formula: I "^'^'Dp |^ 1-3 M 1-3

Ridge-crest formula (based largely on Stegolophodon latidens) :

Dp 21 Dp 3: Dp 4^ M 1^^^^' M 2 fff M 3 -> +-

r>-6VS
+

Horizon. — Mio-Pliocene
to Upper (?) Pliocene [Lower Pleistocene?]. Compare Vol. II, p. 1531,
and figures 1221, 1223, 1228, also PI. xix.

Origin and Migration. — It was doubtfully suggested by Schlosser in 1903 (1903, p. 191) that west European
species, originally described as Mastodon M. pyrenaicus, etc., separated as Zygolophodon
turicensis [M. tapiroides,
by Vacek in 1877 and by Osborn in 1926 as Turicius {M. turicensis, M. tapiroides) and Zygolophodon (M. pyrenai-
cus)], may have given rise to primitive Asiatic species [such as Stegolophodon cautleyi]; also that M. turicensis

[= Turicius tapiroides] of the Lower Miocene of Europe may have given rise to the M. [Stegolophodon] latidens of
the Lower Pliocene [?Lower Pleistocene] of Asia from which in turn sprang off the true Stegodonts, such as
Stegodon insignis. Professor Osborn's final opinion regarding the relationships of the Stegolophodontinjp (see
pp. 197 and 819-822 above) would .seem to be in favor of Schlo.sser's suggestion of 1903, also that of Schlesinger
of 1917, that primitive species of Europe [Mastodon (Stegolophodon) sublatidens] may have given rise to primitive
 '

STEGODON AIRAWA/VA

STEGOOO^ TRIGONOCEPHAL U
.JA^A

STEGODON PINJORENS/S
^/SV^AN. INDtA

3TEGODON ORIENTALIS GRANGERI

^ZECHU^N, CHINA

STEGODON /NSIGN/S STEGOOO/V GANESA

SI WALIKS, IfVDIA SI\/i^>ALIKS. INDI^

STEGODON BOAIB/FRONS
SIU^AIIKS, INDIA
'"•' °" ''"
_ y 45'
y 75- yE lOf i;0- IK- [SO- I65' UO" IJO" r6S' fSo" 135' IJO" fOS" 9n"W 7f 60' tf~

30' -^5' 60' 75' 9Q'E 105" iZf fiS' 150' [65' 180' 165' 135" 120'
I50' 105' 90'W 75' 60" PFUvat BradJeijIW,

Origin. Migration and Evolution of

Ste^odon
PLATE XX
Geologic range: Stegodon, Lower Pliocene [?Lower Pleistocene, see p. 8'2-l above], Middle Pliocene to Upper Pleistocene. Since this map was prepared the
range of Slegodon has been extended into Africa, as far south as Kaiso, on Lake Albert.
 SUMMARY 1579

species of Asia to which Schlesinger has given the generic name Stegolophodon (cf. Vol. I, pp. 195, 197, and Pis.
ii-iv, also p. 700). The Stegolophodonts are not known to have reached America. Fossil remains have been
found in Austria, in Perim Island, the Punjab and the Siwaliks of India, also in Burma, Japan, and Borneo. Their
possible line of migration in indicated in figure 1228 and on the accompanying Plate xix.

STEGODONTOIDEA
Superfamily: STEGODONTOIDEA Osborn, 1935, 1936. Family: STEGODONTID^ Young-Hopwood, 1935
Subfamily: StegodontiNvE Osborn, 1918, 1921
Genus: Stegodon Falconer and Cautley, 1847, 1857
(Cf. Vol. I, PI. XI, and Chap. XIV of the present volume, pp. 807, 853, also Figs. 686-688, 691, 1221, 1223, 1228,
and PI. xx)
As fully stated in Chapter XIVabove, Professor Osborn separated the true Stegodonts from the Elephanto-
idea, giving them a new superfamily name, Stegodontoidea, which includes the genus Stegodon only of the sub-
family Stegodontinse, the genus Stegolophodon having been removed by him to the Mastodontoidea, subfamily
Stegolophodontinjae. The family Stegodontidse, first mentioned by Dr. C. C. Young in 1935 and fully defined by
Dr. A. Tindell Hopwood later in the same year, embraced both Stegodon and Stegolophodon. In the present
Memoir this family is logically included in the superfamily Stegodontoidea, although Professor Osborn never
indicated such reference other than in his chart of 1935 (Osborn, 1935.937, fig. 2), and even in this instance he
also included both genera, Stegodon and Stegolophodon. The separation of the Stegodontoids from the Elephan-
toids was based chiefly on the fact that in section the molars of the former showed the valleys separating the
adjacent ridges to be closed or V-shaped at the bottom and those of the latter to be open or U-shaped. Another
determining factor was the extremely short face of the Stegodontoids which, in his opinion, could not have given
rise to the longer face of the Elephantoids. With this classification in mind, the following characters have been
compiled pertaining solely to the Stegodontinse, genus Stegodon.

Stegodon, so named because of the resemblance of the toothed ridges of the grinding teeth to a series
of roof-gables; more primitive than the true elephants. Skull and tusks do not lead into either the
ElephantiniP or the Mammontinse types. Cranium brachycephalic, brachyopic, of mastodontoid
{Stegodon bombifrons), to extremely abbreviated, female? {S. insignis), more elongated, male? {S.
ganesa), more triangular {S. trigonocephalus) form rostrum elongated to support the tusks palate short,
;
;

depressed well below the occipital condyles (bathycephalic) narial openings elevated. Jaws greatly
;

abbreviated. Superior tusks horizontal or subhorizontal in direction, parallel, and slightly upcurved,
without trace of enamel band, attaining great dimensions {S. ganesa). Inferior tusks disappearing very
early. Grinding teeth short crowned; progressive heightening of ridge-crests (brachyodont to subhyp-
sodont), the distinctive feature being the rapid multiplication of transverse crests by the addition
of crests posteriorly; increasing curvature of the occlusal surface; enamel valleys closed or V-shaped
at the bottom instead of U-shaped as in the elephantoids, filling with cement. Particularly interesting
and significant is the transformation of the original cones by binary or rarely by ternary fission into
conelets (maximum 20 -|- in *S. airdwana). Ridge-crests intermediate between Stegolophodon and
Archidiskodon planifrons types. Probably browsers rather than grazers.
Dental formula: Di ""^ I "^^Dp fff M 1-3
1-3

Ridge-crest formula: Dp
2^f Dp 3H Dp 4^^ l?f| 2°;-^ M
3, M M
Horizon. — Lower Pliocene [?Lower Pleistocene, see p. 824 above], Middle Pliocene to Upper Pleisto-
cene. Compare this chapter, p. 1539, and figure 1228.

Origin and Migration. —Professor Osborn's views on the origin of the true Stegodonts are given on page 25
of Volume I of the present Memoir, as follows: "It has been assumed by practically all palaeontologists that the
Elephants were descended from the Stegodonts. . . . The Stegodonts were of independent origin and formed an
1580 OSBORN: THE PROBOSCIDEA

independent parallel branch terminating in the highly specialized Elephas [Stegodon] aurorse Mastumoto from the
Upper(?) Pliocene of Mt. Tomuro, Japan, now . . . separated by Osborn as the Stegodontoidea superfam. nov."
Also on page 853 of the present volume, where the generic characters of Stegodon are enumerated, occurs the
following, agreeing in substance with the above conclusions: "Phylum parallel to that of the true Archidiskodon

and Elephas, not directly ancestral, readily distinguished by cranial and dental characters." It will be noted on
page 819 above that Schlosser suggested the possible derivation of the true Stegodonts from certain Miocene
species of western Europe since referred by the present author to the Zygolophodontinae, that is to say, these may
have given rise to the Lower Pliocene [?Lower Pleistocene] M. [Stegolophodon] latidens of Asia, from which in
turn sprang off the true Stegodonts, such as Stegodon insignis. Professor Osborn regarded this phylogenetic
problem as of such importance that he stated on page 197 of Volume I that it would be treated more fully in

Volimie II, under the Stegodontinae [of the Stegodontoidea superfam. nov. ]. It is with regret that we are unable to

carry out his intention in this respect, as he left no record of his final views on the subject other than is given above.

The accompanying migration map (PI. xx), therefore, omits the hypothetical European point of departure
and indicates the Siwaliks of India as the center of dispersal, from which radiate three distinct routes, namely, to
Burma and Java, to central China, and to the northern part of China onward to Japan. Up to the present time

Stegodon has not been found in the Western Hemisphere. See Chapter XXII, p. 1436 above, where Dr. Edwin H.
Colbert cites from a letter of January 14, 1938, from Dr. A. Tindell Hopwood, in which the range of Stegodon is
extended into Africa as far south as Kaiso on the eastern shore of Lake Albert.

ELEPHANTOIDEA
It will be recalled that Professor Osborn long held the opinion that Africa was the center of adaptive radia-
tion—or the "homeland"— of the Proboscidea. With this thought ever in mind, he pursued his researches over
a period of nearly thirty-five years, and, happily, the results, so far as can be determined at the present time, have
confirmed his views, as will be seen from the following citation from his last paper on the subject, "The Ancestral
Tree of the Proboscidea. Discovery, Evolution, Migration and Extinction over a 50,000,000 Year Period,"
1935.937, p. 410:

In the Elephantoidea we have discovered a brilliant example of aristogenic growth and extensive migration in the evolution
of Archidiskodon proplanifrons, a very primitive stage found in the Vaal River gravels of South Africa, into the Archidiskodon
7naibeni of Nebraska. Archidiskodon proplanifrons has a molar crown pattern like that of a mastodont, with an enamel length
of 690 mm. Archidiskodon maibeni has eighteen tall ridge plates and an estimated enamel length of 9000 mm. This is the first
;

time that the evolution of the elephantoid molar from a theoretic mastodont prototype has ever been actually demonstrated . . .

Th(! 15,000 mil(! migration of Archidiskodon, from the Vaal River of South Africa through intermediate stages in North Africa,
France, Italy, Britain, India, then the long geographic break to the Niobrara River in Nebraska where Leidy discovered these
animals in 1858, is the most remarkable trek of any species of plant or animal thus far discovered in the life history of the earth.
The by-product is the fact that these stages give us an unprecedented opportunity of measuring precisely the actual I'ate of evo-
lution between Upper Pliocene and Middle Pleistocene time. The enamel length increasing from 690 mm. to an estimated
9000 mm. affords an evolution of approximately 15 mm. of enamel length per thousand years.

In this introduction to the subfamily members of the Elephantoidea it may not be amiss to give the present
author's definitions of alloiometrons and aristogenes, as well as an outline of his "elephant enamel or ganometric
method" of measuring Pleistocene time, particularly in view of the fact that reference has been made in the fore-

going quotation to aristogenic and ganometric usage in his researches. Compare page 1582, also figure 300 of
Vol. I, and figures 1231, 1239, 1240 of this chapter.

"Aristogenes are new adaptive units originating directly in the geneplasm and slowly evolving into important
functional service." Hence aristogenesis (first known under the term "definite variation," then as "rectigradation")
 o

SUMMARY 1581

"is a creative process from the geneplasm of entirely new germinal biomechanisms ; the process is continuous,
gradual, direct, definite in the direction of future adaptation."

In time contrast to alloiometrons (that is, changes of proportion or intensity which may be expressed in
measurements and indices, and which appear to be immediate and more or less temporal adaptive reactions to new
habits), aristogenes are secular, appearing very slowly in the course of long periods of geologic time. Lines of
ordinal, family, generic and specific descent may be distinguished by the potentiality of certain new geneplasmic
aristogenes. (Modified from Osborn, 1934.922, pp. 202, 210.)

For example, compare Osborn, op. cit., p. 221: "In

NEBRASKA the elephantoid division the low transverse ridge-crest is

UPPER PLEISTOCENE perfected in the upper Pliocene of the African ancestral

High'Crotumd giant Archidiskcdcn elephants (Archidiskodon) . The unique aristogenic

18 hffy rtdpe-plates. Enamel length = mm

A/lometric Adaptations of the Limbs Osborn- Gregory. I929

FEMTJK
NEBRASKA
Sfwrteniriif
LOWER PLEISTOCENE with Lengthening

increasing speed with

Medium-crowned ^rch idt'sko don
27.1% of total length increasiru^ weig/it
14 ridge-piates Enamel length ^ mm
AfrCMlDlSKODOrj MtwlDlONALIS NtbRASCENStS 367gmmj2'»t' 48.6% of total length
l«3 S HEBRASKA

FRANCE
~~~
LOWER PLEISTOCENE TIBIA
Length remaining relatively constant
Medium-croLuned elephant molars
31.7%
13~14 ndge-pUtes Enamel ien^th=2d24 mm

FRANCE 34. 3 '^o

LOWER PLEISTOCENE

Loui-croujned elephant molars

PES
Lengthen irig
15 ridge-plates Enamel length ° mm
ADCMIDISKODOM PLANlFfiONS 3745 mm.l£'3«' with Shortening
l')3l FRANCE CHAGMY
increasing speed
with
INDIA UPPER PLIOCENE •11.1% of total length increasing weight
Molars a^prlmitive elephant type 17.1%, of total length
TYPE
APCHIDISKOOON PLANIFRON& I9a0mm.6'6'6 10-12 rldqe-plates Enamel length - 1515mm.
J93) INDIA 3IWAUK5
ALLOMETRONS: Adaptit'e J/iecd and Height Prof>Qrtiuns
(le/t) Horse. Cursorial: Pes 4HZ of total lengtA.Femur 27 1 % of total length
SOUTH AFRICA. MIDDLE PLIOCENE
(/(ight) £lej>hant-gmviportal: 171X ~ - 4S-6X - -
TYPE(Ji
'
fS\\ Molars transitional from mastodon to elephant
AflCHlOlSKOOON 3USPLAN1FR0N5 ld00mnU4'7'e ' , ,
rtdge-piates.
. c
tnameiI I i-L
iength
£t:n
= ooumm.
Analogous ALLOMETRONS Speed and Hilght evolve, in Ajammals and Kyitiles
SOUTH AFRICA

Fig. 1240. (Left) Equus: Alloiometrons of speed; (Right) Elephas:

SOUTH AFRICA. MIDDLE PLIOCENE
Alloiometrons of weight. .4fter Osborn, 1938.1, fig. 6.
Mastodon - like molars
ARCHIDISKODON -rroplSn^ns i43omm J'R'e 5!i ndge-plates. Enamel length- 690mm.
Abbreviating femur of the horse, 27.1 %
of total length.
Elongating femur of the elephant, 48.6 %
of total length.
ARCHIDISKODON: THREE-MILLION-YEAR EVOLUTION AND MIGRATION Tibia of the horse relatively constant, 31.7 %
of total length.
SOUTH AFRICA TO AMERICA AND MEXICO H.F.o. 1935
Tibia of the elephant relatively constant, 34.3 of total length.%
Elongating pes of the horse, 41.1 "^J, of total length.
Fig. 1239. Geneplasmic evolution of the archaic-toothed mammoths Abbreviating pes of the elephant, 17.1 %
of total length.

during a three-million-year period so far as known to April, 1935. Similar cursorial alloiometrons evolve in all quadrupeds attaining speed,
Twenty-seven years of continuous exploration and research yielded all irrespective of phyletic relationship. Similar graviportal alloiometrons evolve
the ascending stages in the geneplasm of this archaic-toothed mammoth. in quadrupeds irrespective of mammalian (Proboscidea) or reptilian (Sauro-
After Osborn, 1938.1, fig. 11. poda) affinity.
 : :

1582 OSBORN: THE PROBOSCIDEA

potentiality of the transverse ridge-crests to convert pairs of cones by transverse binary fission into transverse
ridges is followed by the elevation of these ridge-crests into the elephantine ridge-plates, as first manifested in the
roof-toothed Stegodon of southern Eurasia. It is paralleled in the distinct Archidiskodon ridge-plated stock of the
upper Pliocene of South Africa. This dark continent gave rise to the world dominant elephantoid division of the
Proboscidea . . . Aristogenesis combined with alloiometric extension is carried to the biomechanical extreme in the
divergently adaptive grinders of the three mammoths [Archidiskodon, Parelephas, Mammonteus]. The contrasts
in the total length of the enamel foldings of the gigantic Archidiskodon (8,000 mm), of the gigantic Parelephas
(10,000 mm), of the relatively small Mammonteus (6,000 mm) are coordinated with the relative intensities of
their struggle for existence." To summarize, in Professor Osborn's own words (op. cit., p. 234)

In biomechanical evolution there are two distinct processes.The one, long known, consists in the alloiometric modification
of existing adaptations as in changes of proportion The other, discovered in course of researches on the phylo-
and of function.
geny of the horses, titanothcres and proboscideans, consists in the gradual geneplasmic origin of new and distinct adaptations;
it is to the latter originative and creative process that the term Aristogenesis is applied. Both processes become part of the
hereditary equipment of the organism.

We may now proceed to epitomize the three subfamilies embraced within the superfamily Elephantoidea,
family Elephantida", namely, the Mammontinae, Loxodontinse, and Elephantinae, in uniformity with the fifteen
subfamilies of the Mastodontoidea, and the single subfamily each of the Moeritherioidea, Deinotherioidea, and
Stegodontoidea.

Superfamily: ELEPHANTOIDEA Osborn, 1921. Family: ELEPHANTID^ Gray, 1821

Subfamily: Mammontin^ Osborn, 1921'

Genera: Archidiskodon FohWg, 1S85, ISS8; Metarchidiskodon Osborn, 1934:; Parelephas

Osborn, 1924; Mammonteus Camper-Osborn, 1788-1924'

(Cf. the present volume, Chapters XVI, XVII, and XVIII, also Pis. xxi and xxii, and Figs. 815, 822, 933,
1006, 1221-1225, 1228)

The name Mammontinae Osborn, 1921, was substituted for Euelephantinae Osborn, 1918, owing to the fact
that the genus Euelephas Falconer, 1857, was invalid (see Chap. XIX, p. 1175 above). Originally this subfamily,
aside from the mammoths, doubtfully included Elephas hysudricus of India, subsequently found to belong to the
Elephantinae, genus Hypselephas Osborn, because of profound cranial and dental differences. A definition of this

subfamily reads as follows on page 32 of Volume I

Subfamily Mammontinse (Mammoths) —Osborn, 1921.515, p. 1. Of close original affinity to the Elephantinae, including

(«) the southern mammoths Archidiskodon plnnifrona and A. meridionulis of southern Eurasia, A. impeiator of North America,
all with broad-plated teeth and few crests; al.so (6) the northern mammoths which apparently include Parelephas Irogontherii

of western Europe, P. columbi and P. jeffersonii of North America, and the widespread woolly mammoth {Mammonteus primi-
genius) of the northern steppes.

Great confusion had existed for years in the determination of species of Elephas columbi, E. imperator, and
E. primigenius. Superficially these three species are so similar that Cope in 1889 referred the remains of a fine
skull of Archidiskodon from Texas to Elephas primigenius columbi, and the present author also saw only resem-
blances to Elephas columbi in the fine Indiana skeleton (now the type of Parelephas jeffersonii.) , whereas Hay
referred the same skeleton to Elephas primigenius. Consequently Professor Osborn made a careful comparison of
'[Should Mammonteus prove to ho inv.'ilid, tliis would leave the subfamily Maninioiitiii.T without a type genus. — Editor.]
 ARCHIDISKODON IMPERATpR MAI&ENI

ARCHIDISKODON MERIDIONALIS. valj'arno, it^ly

ARCHIDISKODON EXILIS
ARCHIDISKODON s.anta rosa island
I M PER A TOR CALIFORNIA
TEX^S

ARCHIDISKODON MERIDIONALIS NEBRASCENSIS

NEBRASKA
ARCHIDISKODON MERIDIONA ^4
LIS DURFORT,

ARCHIDISKODON PLANIFRONS, chacn-t-, France

ARCHIDISKODON IMPER4T0R SCOTTI
1<Z NEBRASKA

ARCHIDISKODON PLANIFRONS. sii^aliks, India

ARCHIDISKODON SURPLAN I FRONS, vaal rii^er. south Africa

ARCHIDISKODON PROPLAN IFRONS, GONa-eoNC. south Africa

15' Q- 15' jtf .15' 60' 75' 90'E 105' 120' 135' 150' 165' IflO^ 165' I50^ ^.15' 120" 105° 90'W 15' 60* PFLevctt BraJ]enl9M/

Origin. Migration and Evolution of

o Archidiskodon, * Metarcfildiskodon
PLATE XXI
Geologic range: Archidiskodon, Mkklle Pliocene to Upper Pleistocene; ytetnrchidiskodon, Lo\ver(?) Pleistocene.
 — .

SUMMARY 1583

both type and referred specimens of these three species, which he found were united by distinctive cranial cliarac-
ters but separated by various types of dental characters, resulting in their reference by him to the following
genera (see his articles in Novitates, 1922.555, 1924.633, 1925.662):

Elephas columhi = Parelephas cohnnhi and P. jeffersonii

Elephas imperator = Archidiskodon imperator
Elephas primigenius = Mammonteus primigenius

In the present Memoir the Mammontinae embrace four genera, namely, Archidiskodon Pohlig, 1885, 1888, the
most primitive member, with the genotypic species, Elephas meridionalis Nesti, 1825, of the Val d'Arno, Italy,
and E. -planifrons Falconer and Cautley, 1846 [1845] of India; Metarchidiskodon Osborn, 1934, genotype Loxo-
donta griqua Haughton, 1922, of the Vaal River, South Africa; Parelephas Osborn, 1924, genotype Elephas
jeffersonii of Indiana, and Mammonteus^ Camper-Osborn, 1788-1924, the history and outstanding characters of
which are given in the following synopses.

Archidiskodon (referring to the archaic molar ridge-plates) of the southern and south temperate
zones; also known as the Southern Mammoth, in reference to its remote relationship to the Northern
Mammoth. Cranium foreshortened and broadened (brachycephalic), deepened (bathycephalic),
heightened (hypsicephalic) through hypsicephaly, orbits and occipital condyles approximated, i.e.,
;

brachycranial (.4. meridionalis), compressed fore and aft (cyrtocephalic), occipitofrontal apex vertically
heightened (acrocephalic) far closer to Mammonteus and Parelephas than to either Elephas or Loxodonta.
;

Nasals pointed. Forehead flattened {A. planijrons) concave {A. meridionalis, A. imperator)
, Mandible .

short; prolongation and beaklike depression of the symphysis (^4. planijrons); without beak, blunt,
—
obtuse (adult male A. meridionalis fide Deperet and Mayet, 1923, p. 156) relatively long and shallow ;

{A. hayi); symphysis prominent, ramus relatively slender (primitive stage). Superior tusks large
(max. 13+ to 16+ feet in length), incurved, crossing in old males; no inferior tusks. Molars short,
subhypsodont; subloxodont in primitive species; ridge-plates extremely broad, widely separated,
enamel borders thickened, more or less crenulate; cement usually very thick; A. planijrons without
cement outer coating. Ridge-plate formula slowly progressive from 3- {A. pro planijrons), to 3 ff^M M
{A. planijrons), to 3^1:1 ^ (^4.Mmeridionalis), to M3J8:M( A. imperator); succession of premolars, P 4,
P 4 (.4. planijrons)

Dental formula {A. planijrons) : I --— Dp ~ P |^ M 1^. This premolar succession does not occur,
so far as known, in any other species of the Elephantidae.

Ridge-plate formula (A. planijrons) : Dp 2^ Dp 3|^ P 3t P 4^^ Dp 4fi M 1^ M 2f M 3f^

—
Horizon. Middle Pliocene to Upper Pleistocene. Compare this chapter, p. 1540, and figures
1221, 1224, 1239, also PI. xxi.

Dr. Hans Pohlig, to the best of our knowledge, was the first to assign the name Archidiskodonten to elephant
molars with archetypal ridge-plates, specifying Elephas planijrons Falconer and Cautley, 1846 [1845], and E.
meridionalis Nesti, 1825, as types. This was in 1885. Subsequently (1888, p. 138) he supplemented his descrip-
tion, giving E. meridionalis only as the type, and on page 252 concluded with the introduction of three new
generic names Archidiskodon, Polydiskodon, and Loxo{-disko)don, of which Professor Osborn accepted Archidis-
kodon only as valid.

The first known species of this genus, therefore, was Elephas meridionalis Nesti. Leidy was the first to
describe (1858) an American species belonging to the genus Archidiskodon, namely, Elephas imperator from
Nebraska, regarded by the present author as a direct descendant of the E. meridionalis of Durfort, France —the
'[See Chapter XXI, pp. 1363-1367 for the history of this genus, which is of doubtful validity. Should Mammonteus prove to be invalid, this would
leave the subfamily Mammontinae without a type genus. Editor.] —
1584 OSBORN: THE PROBOSCIDEA

last Eurasian representative. A comparison of the Durfort skeleton with that of the recently discovered Archi-
diakodon meridionalis nebrascensis of Nuckolls County, Nebraska, led him to this conclusion as to lineage. In
1915 Dr. Erwin H. Barbour described Elephas hayi, with a jaw almost as primitive in structure as that of E.
planifrons, and a decade after appeared his description of the gigantic E. maiheni, both from Nebraska.

Through the courtesy of Director Wilman of the McGregor Museum at Kimberley, South Africa, two ele-

phant's teeth were forwarded to Professor Osborn in 1926-1927 for description. Simultaneously Professor
Raymond Dart described (Nature, 1927) two new species {Archtdiskodon transvaalensis and A. sheppardi),
which subsequently were found to belong to the genus Palaeoloxodon. In 1928 Professor Osborn contributed an
article to Nature in which he described the two molars from the McGregor Museum as Archtdiskodon subplanifrons
and A. broomi. Between 1929 and 1932 Profes.sor Dart described several species from South Africa, which were re-
viewed by Professor Osborn in American Museum Novitates of August, 1934; certain of these species were foimd
to be referable to Palaeoloxodon rather than to Archidiskodon. In this article he described the new genus Metarchi-
diskodon, genotype Loxodonta griqua Haughton, 1922, also the new species Archidiskodon proplanifrons from
Gong-Gong near the Vaal River, in his opinion the most primitive elephant tooth thus far discovered, even more
primitive than A. subplanifrons, and indubitably an ancestral Archidiskodon.

Migration. —The Archidiskodonts were widely distributed geographically, as will be seen from the ac-
companying Plate XXI, as well as from figures 815, 1228. Until the discovery of Archidiskodon proplanifrons
and A. subplanifrons of South Africa, the E. [Archidiskodon] planifrons of India was thought to be geologically
the earliest as well as the most primitive of all the members of this phylum. It is now believed that from the
region of the Vaal River, South Africa, the Archidiskodonts radiated into all the continents except Australia
(where up to the present time no proboscidean remains have been found) and South America. As stated above
on page 1580, the 15,000-mile journey from the Vaal River, through North Africa, France, Italy, England,
across to Rumania, onward to India, thence to North America (Saskatchewan, California, Montana, Wyoming,
Nebraska, Kansas, Oklahoma, Texas, Mexico, and Florida) is one of the most remarkable instances of the migra-
tion of any species of animal or plant ever recorded in the life history of the earth.

Metarchidiskodon, genotype Loxodonta griqua Haughton, 1922, of Griqualand West, South Africa.
A fragmentary molar with the following distinctive characters: Cement areas equal or exceed dentine
areas; pre-sinus folds absent or inconspicuous; very prominent post-sinus folds; very deep U-shaped
valleys extending to bottom of crown —
a very important point valleys filled to the summit with cement;
;

enamel ridge-plates very deej), closely compressed with very narrow dentinal areas between.
Horizon. — Lower(?) Pleistocene. Compare this chapter, page 1540, and figures 1220, 1228, also PI. xxi.

In reviewing the African material (see Osborn, 1934.925, p. 12) Professor Osborn observed that the frag-
mentary molar specimen described by Haughton in 1922 as Loxodonta griqua possessed characters (as enumerated
above) distinguishing it not only from Loxodonta, but from Archidiskodon as well, and accordingly he made it

the type of a new genus Metarchidiskodon. This group may also include certain relatively narrow grinders from
the Val d'Arno now in the British Museum, namely, Brit. Mus. M12641 and M12642.

Parelephas, in reference to the convergence or parallelism of the grinding teeth with those of the
true Elephas; of the intermediate and north temperate zones. Cranium relatively broad, elongate, and
rounded, intermediate in form between that of Archidiskodon and that of Mammonteus, namely, in
bathycejjhaly and acrocephaly. Frontals concave, occipital crest elevated, occiput more or less convex,
moderately compressed fore and aft (cyrtociephalic) moderately depressed molar-grinding area (bathy-
;
 ELEPHAS INDICUS
BENGALENSIS
BENGAiL, INDIA

PARELEPHAS TROGONTHERU PARELEPHAS COLUMBI

MOSBACH. GERMANY- GEORGIA

PARELEPHAS TROGONTHERH. taubach. German-,'

Origin,Migration and Evolution oF

o Parelephas, * Mammonteus, a Elephas

PLATE XXII
Geologic range: Parelephas, Upper(?) Pliocene [Lower(?) Pleistocene, see footnote 1 1049 above) to Upper Pleistocene. Mammonteus, Upper Pliocene
on p.
to late Pleistocene. Elephas, l^pper Pleistocene and Recent. phylum are Hypselephas of India (Lower Pleistocene)
Fossil representatives of the Elephantine
ami Platelephas also of India (Upiier Pliocene or Lower Pleistocene), restorations of which do not appear on the accompanying plate, nor elsewhere in this
Memoir, as they were never prejjared by Professor Osborn.
 SUMMARY 1585

cephalic), space between condyle and orbit broader than in Mammonteus primigenius anterior narial ;

opening broad and widely open divergent maxillo-premaxillary region, tusk sockets less elongate and
;

less parallel than in Mammonteus. Mandible robust, short and deep (bathy cephalic) ramus depressed ;

with rounded inferior border; rostrum prominent. Superior tusks long, with remarkable incurvature,
crossing in old age {Parelephas jeffersonii); shorter, more robust (P. floridanus). No inferior tusks.
Molars relatively narrow as compared with those ofArchidiskodon, with enamel of intermediate thickness,
more or less crimped or sinuous. Ridge-plates arcuate, converging toward the summit; consequently
the ridge-plate compression depends upon the level at which the count is taken the count ranges in —
P. jeffersonii molars from 7 in 100 mm. (base) to llK (summit) in the inferior molars, from 7 (base) to 10
(summit) in the superior molars. Relatively few ridge-plates in the Upper Pliocene to Middle Pleis-
tocene stages (M 3;*+), progressive Upper Pleistocene stages {P. progressiis) with multiple ridge-
plates (M 3 If). Thin cement (P. columbi). Adapted to continental plains or steppe environment,
grazing and browsing.

Dental formula: I ^^ Dp §f| M 1^ j^-3

Ridge-plate formulae:
P. columbi: Dp 2§ Dp 3'" Dp 4^^ M l^F M 2^{^ M 3,^^^
P. jeffersonii: Dp 3^ Dp 4|f M 1^ M 2^^^' M 3 H

Comparative Ridge-plate Formulae of M 3 in Parelephas

Parelephas progressus M3 If Parelephas columbi M3 1 9
X6 +
Parelephas jeffersonii M3H Parelephas intermedins M3 2 0-2 1

Parelephas icashingtonii M 3 ff Parelephas trogontherii M3 1

16 +
5 +

Parelephas floridanus M 3 —^2 Parelephas trogontherioides M3 ''^^.\\%

Horizon. —Upper(?) Pliocene [Lower(?) Pleistocene, see footnote 1 on p. 1049 above] to Upper
Pleistocene. Compare p. 1540, also figures 1222, 1224, 1225, and PI. xxii.

Professor Osborn introduces his chapter on Parelephas (Chap. XVII) with the following statement:

... it appears that Parelephas is A irhidiskodoti and Mammonteus in its cranial resemblances and great incurved
linked with
and ridge-plate formulae it is so nearly intermediate between these two genera as to have
incisive tusks, while in its grinding teeth
been mistaken for an actual connecting link. In the present chapter [XVII] it is shown to be an entirely distinct generic phylum
which during the more temperate interglacial periods occupied the same geographic range as that of the true woolly mammoth
. . .

{Mammonteus) during the glacial periods. [Compare Fig. 1241, below.]

While not sufficiently distinctive to preclude the subfamily imity of these three genera, there are certain

cranial differences, such as the intensification of characters like hypsicephaly, bathycephaly, and acrocephaly.
Their dental diversification, however, definitely separates them generically, for example, the broad, widely sep-
arated, thickly enamelled ridge-plates of Archidiskodon, as compared with the relatively narrow, more compressed
and less thickly enamelled ridge-plates of Parelephas, and the narrow, extremely compressed, thinly enamelled,
hypsodont ridge-plates of Mammonteus also the multiplication of the ridge-plates of M 3, which in Archidiskodon
;

increase from J"^ to -—, and in Mammonteus from J|^ to y^, whereas in Parelephas the increment is from
H^—^ to f|. It should also be stressed that, contrary to the normal ridge-plate structure, the superior count
generally exceeds that of the inferior in Parelephas.

The by which Parelephas crania may be distinguished from Mammonteus crania are
principal characters
summarized by Professor Osborn (Chap. XVII, p. 1051) as follows:
(1) In frontal aspect the crania of Parelephas are relatively broader, more spreading, and more brachycephalic than those of
Mammonteus, which are deeper and more bathycephalic; (2) in lateral aspect (a) the orbit is more widely separated from the
occipital condyle, (b) the occiput is much more convex, thus throwing the occipitoparietal apex farther forward, (c) the height
from the occipital apex to the superior molar crowns is less deep, i.e., less bathycephalic, (d) the apex formed at the summit of
the cranium is less acute, (e) the facial front is shorter and more deeply concave, (f) the maxillo-premaxillary sockets are less
1586 OSBOKN: THE PROBOSCIDEA
and the tusks emerge in a less vertical plane; (3) in frontal aspect (a) the premaxillary sockets arc more expanded at
vertical
tlie Mammontciis they arc more elongate and more closely compressed, (b) the transverse diameter of
extremities, whereas in
the frontals is relatively broader than in Mammonleus, (c) the anterior nares are proportionately broader transversely and less
deepened vertically; (4) in brief, the proportions of the cranium of Parelephas throughout are harmonious with those of the
grinding teeth, i.e., less compressed anteroposteriorly, le.ss bathycephalic and less hypsicephalic than those of Mammonleus.

This section would not be complete without some mention of Professor Osborn's clarification of the phyletic
position of the true "Elephas" columhi of Falconer, 1857, from Georgia, and his own "Elephas" [Parelephas]
jeffersonii from Indiana described in 1922. Falconer beheved E. columhi to be identical with Leidy's E. [Archi-
diskodon] imperator, and the present author for several years treated it under the genus Archidiskodon but as
a species quite distinct from imperator. Falconer was in error, as the two species are distinct, which will be seen
by the following comparison; moreover, according to the views of Professor Osborn, they were not geologically
contemporaneous, although the late Doctor Gidley [and many others] believed otherwise.

Elephas [=Parelephas] columhi Falconer, 1857-1868. Upper Pleistocene of southern United States and of
Mexico. Smaller animal, with narrower grinders (Fig. 887), thin cement outer coating;
maximum ridge-plate formula, M
3 ri^rh- Enamel ridge-plates arcuate, converging at
summit, giving the appearance in extremely worn grinding teeth of being as far apart in
mid-section as in .4. imperator.

Elephas [
= Archidiskodon] imperator Leidy, 1858. Lower [Early and Middle Pleistocene, .^d(^ Lugn and
Schultz, 1934, pp. 373-376] of southern United States and of Mexico. Larger animal in
size, with broader grinders (Fig. 887), very broad enamel plates, and heavy cement outer
coating; ridge-plate formula, M37|^. Enamel ridge-plates widely separated.

The present author was long misled by the widely separated ridge-plates seen in the types of both Elephas
columhi and E. imperator, but with the opportunity for further study through the acquisition of new and rich
materials from Florida and the phosphate beds of South Carolina, he perceived that the type of E. columhi
belonged within the phylum Parelephas.

Again, the type of the true Elephas [Parelephas] columhi of Georgia, Florida, and South Carolina must not be
confused with the Elephas [Parelephas] jeffersonii of the northern states, from which it is separable in its much
more primitive ridge formula. The species E. jeffersonii first described by Professor Osborn in 1922 (Osborn,
1922.555, i)p. 11-16) was based on a skeleton from Indiana referred to '^Elephas columhi" by Cope and by Osborn,
and to E. primiyenius by Hay. In his description he gave the distinctive characters of the grinding teeth of each
of the species E. imperator, E. columhi, and E. primigenius, remarking that these diagnoses left without a name the
animal which previously had been described in all the literature (excepting Soergel's Memoir of 1921) as Elephas
columhi. Unfortunately in his original description of E. jeffersonii Professor Osborn chose as paratypes a pair of
upper and lower molars of both sides from Zanesville, Ohio, described by Warren in 1855 as Elephas "primigenius,^^
which he finally made the type of a new subspecies, Parelephas jeffersonii progressus, because of the large number of
ridge-plates, namely, M Zll. In the meantime the molars of the aged type specimen from Indiana were cut out of
the jaw and sectioned, yielding a ridge-plate formula of M 3|f, still a much higher ridge-plate count than that
of the true "Elephas columhi," namely, M 3/^. After the establishment, therefore, of Elephas jeffersonii as
a species distinct from E. columhi and from E. primigenius, Professor Osborn (1924.633, p. 4) reached the con-
clusion that this species could be placed neither in the phylum of Archidiskodon nor in the phylum of Mammon-
teus ;
consequently he selected it as the type of a new genus, Parelephas.

In summation, Elephas columhi of Falconer, with a ridge-plate formula in M 3 of jVt ai^tl an average specific

ridge-plate frequency of 5-6}. in 100 mm., as compared with 7-9 (max. ll}.) in E. jeffersonii, 5-7 in E. imperator.
 —

SUMMARY 1587

and 8-13 in E. primigenius, is to be regarded as a distinct species of Parelephas, namely, P. columbi;^ E. columhi
(previously described as the Columbian Mammoth) is now to be known as Parelephas jeffersonii or the Jeffersonian
Mammoth. The skeleton from Brevard County, near Melbourne, Florida, now in the Amherst Museum, be-
longs to the true Parelephas columhi.

Migration. — A very careful comparison of all the known characters of the Elephas trogontherii phylum of

Europe and the Elephas jeffersonii of America, especially the cranial characters, established their close phyletic
relationship, justifying the linking of the European and American species in the new and distinct genus Parelephas.

The low ridge-plate formula of the true Parelephas columhi of Georgia and South Carolina suggested to Professor

Osborn the possibility of the early geologic entrance into America of relatively primitive species of Parelephas,

a theory which is supported by the primitive character of the lower jaw of P. xmshingtonii of Whitman County,
state of Washington, the molar ridge-plate formula of which agrees quite closely with that of P. intermedius of
southern France. Hence the evidence appears to indicate that the ancestors of P. columhi and of P. ivashingtonii
may have passed across Europe and Asia- and migrated far southward in North America during the 2d and even
possibly during the 1st Interglacial period, following the wave of migration into America of Archidiskodon. Profes-
sor Osborn was sanguine of the discovery (probably in northern Africa) of an ancestral stage more primitive than
the Parelephas trogontherioides of the Upper Pliocene^ of Italy. It will be noted from the accompanying Plate xxii
that members of this Parelephas phylum are recorded from France, Italy, England, Germany, southern Russia,
Asia Minor, Canada (Ontario), United States (Washington, Oregon, Montana, Colorado, Iowa, Kansas, Nebras-
ka, Indiana, Illinois, Ohio, Kentucky, South Carolina, Texas), Mexico, and French Guiana — the only representa-
tive of the Elephantidse that succeeded in reaching the South American continent, as far as we know at the
present time. Thus we have an elephant phylogeny and migration second only to that of Archidiskodon.

Mammonieus, the Northern or Woolly Mammoth, of the northerly and circumpolar zones. Cranium
related to that of Archidiskodon and of Parelephas, with fore-and-aft compression, resulting in bathyceph-
aly and hypsicephaly, which exceeds by far that of any other proboscidean or other mammalian skull
known. Frontals concave, occipital crest greatly elevated, occiput slightly convex. Mandible with
excessively short, deeply depressed ramus, terminating in a deflected and extremely narrow rostrum
in short, harmonic with the extreme hypsicephaly and bathycephaly of the cranium. Elevation of
coronoid and of mandibular condyle. Anterior nares small. Superior tusks of large proportions, greatly
incurved, crossing in old age; vertically placed tubular maxillo-premaxillary insertions of tusks, which
are relatively longer, narrower, and deeper than those of any species of Parelephas. No inferior tusks.
Molars with relatively numerous ridge-plates (polydiskodonty). Upper Pliocene stage {Mammonteus
primigenius astensis), M
S^'vao! typical Upper Pleistocene stage {M. primigenius), Sfl; final M
progressive stage {M. primigenius compressus), broad, M 3^
hypsodont, with enamel ridge-plates of
;

minimum thickness, more or less crimped or sinuous, in fact, 3 is the broadest and deepest probosci- M
dean molar known (compressus). Ridge-plates compressed in typical superior molars to 10-11-12 in
100 mm., in highly progressive superior molars 13 in 100 mm.; in progressive inferior molars 8-9-10 in
100 mm. As in Parelephas, the molars are arcuate at base, more compressed at summit, consequently
the ridge-plate count varies and as a rule should be taken at mid-section, both on the internal and ex-
ternal sides. Manus pentadactyl with five digital nails (fide Herz, 1902), manus and pes tetradactyl not
pentadactyl (fide Salensky,* 1904, p. 86) four digits (^rfe Pfizenmayer, 1926, p. 239) total phalanges in
; ;

'[Professor Osborn remarks (Osborn, 1930.837, p. 17): "It is not improbable that certain of the specimens found in Florida which hitherto have been re-
ferred to Parelephas columhi with its limited ridge formuhi, M 3f 2>, actually belong nearer to P. floridanus stage with its more progressive ridge formula,
M 3|ft."— Editor.)
[Panicphas trogoidltrrii is reported by Tokunaga (Amer. Mus. Novitates, 1933, No. 627, j). 2) as occurring at Honshu (Kazusa, Mikawa, Onii, Shinani)),
Inland Sea, Japan. See figure 1223 in Chapter XXII above, by Dr. Edwin H. Colbert. — Editor.)
'[Possibly Lower Pleistocene, see footnote 1 on p. 1049 above. — Editor.)
^[See Zalensky, Vladimir Vladimirovich, in Bibliography of Volume I of the present Memoir. — Editor.)
1588 OSBORN: THE PROBOSCIDEA

each foot reduced to nine in number, e.g., "

,riTi~iv v ^^Z'"' Dietrich (1912) records five digits in the
manus and pes of the large and more primitive Upper(?) Pleistocene M. primigenius fraasi. Tail
abbreviate, caudals 21.

Dental formula: I
""-"
Dp f:^ M ii|
Ridge-plate formula: Dp 2i Dp 3f Dp 4fg M
i;§ 2;^ M M
3=^ (typical M. primigenius)
Horizon. — Upper(?) Pliocene [?Lower Pleistocene] to late Pleistocene. Compare page 1541 of the
present volume.

The true mammoth is the only extinct proboscidean of which the characters of the soft parts and of the hairy
and woolly covering, as well as the nature of the food (with the exception of the Mastodon), are fully known. It
is probable that these northerly or woolly mammoths were the first mammalian fossils of northern Eurasia to be
discovered and recognized as extinct; the earliest descriptions are naturally lost in obscurity. The typical or
true mammoth was found in Siberia long before it was recognized in western Europe. Blumenbach had in mind
both Siberian and North German specimens in defining Elephas primigenius in 1799, also Cuvier had the Siberian
mammoth in mind in defining Elephas mammonteus (1796 MS., 1799). Out names applied to the
of a host of
northern mammoth between the years 1696 and 1888, the species Elephas primigenius of Blumenbach alone sur-
vives and is accepted in the scientific literature of the entire world. As to the genus, nearly a century and a half
of research since 1799 "demonstrates that the woolly mammoth belongs not to Elephas, but to a genus of its own,
distinct by all the canons of nomenclature from the true modern Elephas." In choosing, therefore, a new generic
name for E. primigenius. Professor Osborn had under consideration the Dictjclotherium of Geoffroy, 1837, the

Cheirolites of von Meyer, 1848, and the Synodontherium of Costa, 1850, also the Polydiskodon of Pohlig, 1888,
which he regarded as the first really appropriate name. He selected, however, the name Mammonteus (see
Osborn, 1924.633, p. 2), a term which he considered as most appropriate and as antedating the generic names
just mentioned, but unfortunately this term was used by Camper in 1788 in an adjectival sense (Mammonteum)
and applied to the remains of a Mastodon and not Mammoth. For a history of the nomenclature of this
to a
name and the substitution by Dr. A. Tindell Hopwood of the term Mammuthus Burnett, 1830, type Mammuthus
Borealis, see pages 1363 to 1367 of Chapter XXI above.

The characters differentiating Mammonteus from Archidiskodon and Parelephas are given on the immediately
preceding pages and are also observable by comparing the diagnoses accompanying these three genera, therefore
they need not be rei)eated here. Mammonteus, in a word, is the most acrocephalic, hy})sicephalic, bathycephalic,
and cyrtocephalic of theMammontines, with the greatest molar compression and an extremely high ridge-plate
formula; also the contour of the body is distinctive, especially in the sloping hind quarters and the sharp notch
behind the peaked skull, features brought out even in the drawings of Pateolithic age.

Migration. —The geographic distribution of the Northern or Woolly Mammoth has ])r()ven to be a most
difficult task, owing to the confusion that has existed in the separation of Elephas [Mammonteus] primigenius from
E. [Parelephas] trogontherii and P. jeffersonii in the previous records (see pp. 1041 , 1 133, 1 138, 1 140) ; in fact, all the

Pleistocene mammoths of western Europe and North America, including all the species of Parelephas as well as all

the primitive and geologically ancient species and subspecies of the true mammoth, have been attributed, or
referred, to the typical species Elephas primigenius. The ancestral relationship to Mammonteus of the Upper
Pliocene [?Lower Pleistocene] Mammonteus primigenius astensis of Italy is problematic, as it shares characters of
both Mammonteus and Parelephas; its ridge-plate fornnila, however, is higher (M 3jBr|f) than that of the
contemporary Parelephas species P. trogontherioides (M 3^^^)- "There can be no doubt," as stated by Pro-
fessor Osborn, "that during late Pleistocene (IV Glacial and Postglacial) time there existed all over northern
-S — — o

o
 —

1590 OSBORN: THE PROBOSCIDEA

Eurasia a single species of mammoth to which the name Mammonteus primigenim lias been assigned, since it

furnished the type to which the earliest as well as the original and later descriptions were applicable."

The accomjianying plate (PL xxii) also figures 1228 and 1241 are of necessity theoretical, but they have
been prepared with the utmost care, and only with a view to expressing Professor Osborn's latest determinations.

It will be observed that the woolly mammoth ranged from Lat. 45° (northern France) to 75° (New Siberian Is-

lands), according to Tolmachoff and Deperet and Mayet; in the Western Hemisphere from the 40th parallel

(New York State on the east) to the western coast, northward to the 70th parallel (Melville Island, fide Sternberg)
and Alaska.

Superfamily: ELEPHANTOIDEA Osborn, 1921. Family: ELEPHANTID^ Gray, 1821

Subfamily: Loxodontin^ Osborn, 1918

Genera: Loxodonta F. Cuvier, 1825, 1827; Palxoloxodon Matsumoto, 1924; Hesperoloxodon Osborn, 1931
Subgenera: Sivalikia Osborn, 1924; Pilgrimia Osborn, 1924, now regarded as synonyms of Palxoloxodon

(Cf. Vol. I, Pis. X, XI, also Chap. XIX of the present volume, Figs. 1220-1223, 1228, and PI. xxiii)

The subfamily Loxodontinse (embracing three genera Loxodonta, Palxoloxodon, and Hesperoloxodon) , like

the Elephantinte, is distinctive in that certain of its members survive to the present day, namely, the Loxodonta
africana of the African continent, the ancestry of which, in the opinion of the present author, is still in doubt, as
will be seen by the following quotation from page 1273 above: "L. africana cannot be descended from any known
typical species of Palxoloxodon but may have sprung from a more primitive ancestral form still to be discovered."
While the Loxodonta zulu of Scott, 1907, and the recently described species of Dart {Loxodonta prima, 1929, L.
subantiqiia, 1929, and L. africana ohliqua, 1932) may prove to be ancestral to the living African elephants. Profes-
sor Osborn was cautious in expressing his views on the subject (Osborn, 1934.925, pp. 7 and 6): "These occur
only on the more recent levels and are clearly related to the existing African elephant, distinguished by the above
[following] characters."

Loxodonta prima group

C'ruvMis relatively narrow, 74 mm.
prima) to 92 mm. (L. subaidiqiui). Enamel relatively thin, coarsely crimped;
(L.
conelctH numerous. Cement thin in middle, thick at edge. Ridge plates per 100 mm. =4 (L. africana obliqtia) to 5% (L.
subanliqua). Broad typical loxodont sinus expansion, double sinus foldings in contact. Total ridge plates 9 {L. prima) to 12-13
(L. Zulu).

In Doctor Dart's description, however, of Loxodonta prima in 1929 (pp. 724-726) he states: "The outstanding
character of this tooth (Figs. 25, 26 [Fig. 1149 of the present Memoir]) is its comparability with the living African
species. It provides the first indubitable fossil evidence of a very close approximation to the distinctive lozenge-
shaped lamellae of the living Loxodont grinding tooth. The absence of such a fossil type up to tiie present time

according to Osborn 'is a striking ('irc^umstance.' . . . There can be no doubt that in this narrow-(!rowned, loxodont-

simulating, few-plated, mountain-inhabiting specimen ... we have a long-sought ancestral type from which the
modern African elephant tooth might reasonably be derived by a progressive widening-out of the loxodont sinus,

for which reason I have named it Loxodonta prima. This fossil is of importance not only in demonstrating an
advancing, though still simple Loxodont tooth pattern in the Transvaal, but also in revealing Africa, and possibly
South Africa as the evolutionary home of the true Loxodonta. Its habitat in the elevated and relatively waterless
Pilandsljerg suggests the possible rea.sons for Loxodont persistence, namely, their b(H'oming inured to more arid
conditions and more active movement, as compared with their more ponderous Archidiskodont relatives."
 LOXODONTA LOXODONTA AFRICANA
AFRICANA PUMILIO OXYOTIS

PALAEOLOXODON MELITENSIS. PALAEOLOXODON

PALAEOLOXODON FALCONERI. NAMADICUS
PALAEOLOXODON MNAIDRIENSIS, /no/ -a

MA L T^
\ODON ANTIQUUS ITALIC US

HESPEROL OXODON
ANTIQUUS GERMANICUSy
<^£RM '^NY

HESPEROLOXODON ANTIQUUS
PL ATYRH YNCH US

HESPEROLOXODON ANTIQUUS of upnor. ENGL^r^D

Origin, Migration and Epoiation oF

^Loxodonta, -^PaLaeoloxodon, oHesperoloxodon
PLATE XXIII
Geologic range: Loxodonla, Pleistocene to Recent; Palxoloxodon, Upper Pliocene(?) or Lower Pleistocene to Upper Pleistocene; Hesperoloxodon,
Upper Pliocene (Lower(?) Pleistocene, see footnote 1 on p. 1049 above] to Upper Pleistocene.
 ;

SUMMARY 1591

Loxodonta, genotype probably the South African form Elephas ajricanus Bhimenbach, 1797; the
extreme South African form was distinguished by Cuvier in 1798 as Elephas capensis. Cranium relatively
low, brachycephalic, platycephalic, mesocephalic, bathycephalic, and cyrtocephalic (foreshortened)
cranial profile and section mucli more primitive, less hypsicephalic and bathycephalic than in the
Mammon tinae or Elephantine; i)arietofrontal cranial vertex low, rounded, platycephalic to subacro-
cephalic, of persistent primitive form; rostrum short and extremely broad; premaxillaries diverging to
—
the point where the tusks issue from the skull thus the bases of the tusks are very far apart instead of
being close together as in Mammonteus; nasals broad and rounded, narial openings hour-glass shaped;
anterior position of suture separating malar from maxillary portion of zygoma. Mandibular ramus elon-
gate; symphysis acute, relatively prominent and horizontal. Superior tusks elongate, widely divergent,
relatively straight, slightly upcurved and incurved, resulting in a lyre-shaped arrangement; marked
sexual disparity in tusks of females. No inferior tusks. Molars low crowned, relatively narrow and with
comparatively few ridge-plates, total ridge-plates Dp 4-M 3 1^ enamel borders thick and simple without
:

foldings or plications: cement thin; distinguished by broadly open 'loxodont sinus' (lozenge-shaped
median expansion) on wear. Ridge-plate formula constant and very conservative, since the 3 ridge M
formula of the living African elephant closely corresponds with that of the primitive Upper Pliocene
Archidiskodon planifrons.
Dental formula: I ^^ Dp |^ iff M
Ridge-plate formula (typical): Dp 2f Dp 3| Dp 4! 2,', 3 MUM M ,^
—
Horizon. Pleistocene to Recent. Compare page 1541, also figures 1220, 1222, and PL xxiii of the
present volume.

Comparison of Loxodonta with the genera Archidiskodon, Parelephas, Mammonteus, and Elephas
Taking, for example, the middle-aged skull of "Jumbo," a Sudanese subspecies, we observe that "in the fully
adult skull the dome is continuously rounded from the occipital condyles to the broad extremities of the nasals,
presenting the widest contrast to the profiles of Archidiskodon, of Parelephas, of Mammonteus, and of Elephas;
that while actually brachycephalic, the cranium of Loxodonta is also comparatively mesocephalic or elongate
(shown in the relatively long mandibular ramus, compared with the deeply depressed mandibular
[Fig. 1060]), as

ramus of Elephas indicus, or with the extremely bathycephalic and abbreviate mandibular ramus of Mammonteus
primigenius; that since both superior and inferior molars, M^, M3, are shorter and less hypsodont, the maxillary
and mandibular dental cavities are much less deep, thus accounting for the less hypsicephalic proportions of this
part of the skull; the superior aspect of the mandibular rami [Fig. 1060] also displays the relative prominence and
horizontal distinction of the rostrum, again presenting a very wide contrast to the deep, hypsicephalic rostrum of
the Mammontinse and of the Elephantinse" ; the superior or frontal aspect of the cranium (Fig. 1061A) shows the
short frontal bones, the massive orbital prominences, the broad narial openings, the widely separate maxillo-
premaxillary sockets for the enormous incisive tusks, similar to those in Palxoloxodon namadicus and Hesperoloxo-
don aiitiquus. It will be observed in a comparison of the palatal aspect of the Loxodonta cranium with that of the
Elephas indicus cranium (Fig. 800) that the former is relatively broader, more brachycephalic in all its dimensions
(cf. Chap. XIX, p. 1200 above).
Palxoloxodon. —The origin of the Palseoloxodonts is obscure. Professor Osborn suggests on page 14 of his
article in Novitates (Osborn, 1934.925) that the "problematic A. = Palxoloxodon] andrewsi" of Dart, 1929,
[

found at Gong-Gong, Vaal River, may be a "primitive or ancestral member of the Palxoloxodon group." In the
present Memoir (cf. p. 1207 above) he also suggests that this generic phylum (which includes Palxoloxodon and
its synonyms Sivalikia and Pilgrimia) may have sprung from the giant North African species Elephas atlanticus
Pomel, 1879. In any event, there can be no question of its clear generic distinction from Loxodonta. In speaking
of Palxoloxodon from the historic standpoint, he remarks (cf. p. 1207) tliat "It is not impossible that some
elephants of the Palxoloxodon type survived into early historic times, but it is an open question whether the
elephants described from Mesopotamia were of the ancient 'loxodontine' or of the modern 'elephantine' type.
1592 OSBORN: THE PROBOSCIDEA

probably the latter; drawings and inscriptions will probably be found some day which will determine these
relations." Pilgrim (1905) notes that Elephas [Palapoloxodon] nnmadiais is entirely absent from the Pliocene

Siwalik strata and that there is no ancestral type from which it might arise.

Palseoloxodon, genotype Elephas namadicus naumanni Makiyama, 1924. Cranium of genotype

unknown, but compare the Elephas [Palxoloxodon] namadicus of Falconer and Cautley, 1846, 1847,
from the Nerbudda, India, which is more hypsicephalic and bathycephalic, and relatively broader
and more platj'cephalic than that of Loxodonta. Parietofrontal cranial vertex somewhat more progres-
sive, acrocephalic, than in Loxodonta, with broad, smooth or rugose crest which in Palxoloxodon namad-
icus extends like a Phrygian cap down over the frontals almost to the nasals (seen also in the dwarfed
Elephas [Palxoloxodo?}] melitensis and E. [P.\ mTiaidriensis of Malta). Narial openings very broad and
shallow. Premaxillaries extremely broad, with widely divergent, relatively straight, slightly upcurved
and iiicur\-pd t>isks toward the extremities. No inferior tusks. Molars moderately hypsodont relative- ;

ly narrow in primitive stages; superior molars progressively narrow to broad, with numerous ridge-
plates composed of thin, plicated enamel foldings. Ridge-plates parallel, closely compressed. Loxodont
sinus rudimentary or absent. Dentine areas equal or exceed cement areas by relatively close compression
of the ridge-plates. Ridge-plate formula progre.ssive from Palxoloxodon atlanticus (M 3}|^yYi), to P.
melitensis (M 87—^), to P. namadicus (M 3y|), to P. namadicus naumanni (M S\j).
Dental formula: I ^^ Dp |^ iff M
Ridge-plate formula [fide Falconer): Dp 4ro^4 I^^ty^^ 2yy^ 3f|^M M M
—
Horizon. Upper Pliocene(?) or Lower Pleistocene to Upper Pleistocene. Compare page 1541
of the present volume, also figures 1220-1223, also PI. xxiii.

Sivalikia and Pilgrimia. — In 1924 both Matsumoto in Japan and the present author in America published
almost simultaneously the results of their researches on the Loxodontines, neither being aware of the activities
of the other. The genus Palxoloxodon of Matsumoto (described September 20, 1924) anticipated by three months
Professor O.sborn's description (December 20, 1924) of Sivalikia and Pilgrimia; hence technically it has priority
over the last two genera, which become synonyms of Palxoloxodon. In Chapter XIX of the jiresent Memoir
Professor Osborn has cxjilained the morphological reasons for regarding his Sivalikia and Pilgrimia as synonyms
of Palxoloxodon Matsumoto.

Dirarfed Species of the Mediterranean Islands. — Before taking up the genus Hesperoloxodon,
which Professor to
Osborn referred members of the 'Elephas antiqmis' group, the dwarfed species of the Mediterranean Islands
deserve .some mention. It is interesting to note, that besides the i^ygmy elephants of Malta, a specimen of nearly
normal size is also recorded from that island. Falconer and the older authorities related these insular species to
the Elephas antiquus of the European continent, but sub.sequent discovery has shown that they are more jirobably
derived from certain of the extinct ancestral African species described in the present Memoir as Palxoloxodon
(syn. Pilgrimia); moreover, the cranium and jaw of Elephas [Palxoloxodon] melitensis,^ with broadly overhanging
parietofrontal crest, more closely resembles the E. [Palxoloxodon]
namadicus of India than the E. [Hesperoloxodon]
antiquus of western Europe. A comparison of the type grinding teeth of the dwarfed Mediterranean sjjecies with
the type grinding to(>th of the extinct African species reveals a striking general resemblance in the narrow pro-
portions and in the rudiment or ab.sencc of the 'loxodont sinus,' characters which appear to relate these teeth to
the Palxoloxodon (syn. Pilgrimia) of Africa rather than to the typical Loxodonta africana or to the broad-toothed
Palxoloxodon namadicus of India. In Europe
some of the narrow-toothed varieties of Elephas [Hesperoloxodon]
antiquus may
be related to the dwarfed insular elephants. After the discovery and description of the cranium
from Pignataro Interamna, Italy, first referred by Professor Osborn to Palxoloxodon antiquus italicus and finally
made the type of his genus Hesperoloxodon, he expressed him.self as follows (see ('hap. XIX, p. 1252) "The entire :

cranial and dorsal hump silhouette (Fig. 1092) is quite different from that of the African elephant (Fig. 1093). . . The
cranial profile of Hesperoloxodon italicus is also entirely different from that of Palxoloxodon namadicus (Fig. 1070)

and of F. melitensis^^^ (Fig. 1 121), both of which are characterized by a very prominent transverse frontal ridge for the
'[Subsequently (see p. 1260 above) referred by Professor Osborn to Palxoloxodon mnaidriensis. — Editor.)
 SUMMARY 1593

attachment of the gigantic proboscis. ... In both cranial and dental characters H. italicus differs profoundly from
the African elephant, which is extremely conservative in its structure, in fact, much more conservative than any of
the known fossil Pleistocene elephants of Eurasia. . . . H. italicus affords additional and positive evidence that the
dwarfed elephants of the Mediterranean Islands {P.falconeri, P. melitensis, and P. mnaidriensis) were not derived
from the 'Elephas antiquus' phylum of Falconer, as hitherto universally believed, but sprang from some undis-
covered phylum of elephants of African origin, which gave rise on the one hand to the dwarfed elephants of the
Mediterranean Islands and on the other to the gigantic P. namadicus phylum of India and the Far East extending
to Japan."

Thus, according to the views last expressed by Professor Osborn, we are able to make the following summary:
Elephas namadicus of India (Nerbudda), varieties and subspecies- of
which are abundant in Japan =the true Palseoloxodon (syn.
Sivalikia of Osborn)

Elephas atlanticus, E. jolensis, E. recki, etc. of Africa = Palxoloxodon (syn. Pilgrimia

of Osborn)
Mediterranean Island species (pigmy and normal size), possibly derived
from the Elephas atlanticus group or the "Elephas
antiquus recki" = Palxoloxodon recki] of Dietrich.
[
= Palxoloxodon (syn. Pilgrimia
(Perhaps some of the narrow-toothed varieties of of Osborn)
"E. antiquus" of Europe may be related to these
dwarfed species.)

Hesperoloxodon. —Elephas antiquus Falconer and Cautley, 1847, described one year after the naming of E.
[Palaeoloxodon] namadicus Falconer and Cautley, 1846, is based on a molar, probably from England, although the
locality is not recorded. To the best of our knowledge no complete cranium of the true E. antiquus has as yet been
discovered, consequently the unearthing of a fine cranial specimen at Pignataro Interamna, Italy, in 1911-1912,
and acquired by the American Museum of Natural History in 1929, was of signal importance. Professor Osborn
immediately described the specimen, referring it first to Palxoloxodon antiquus italicus, but on comparing it more
closely with the cranium of P. namadicus, he observed at once that it was more hypsicephalic and bathycephalic
and that it lacked the prominent transverse frontal crest so characteristic of P. namadicus and P. mnaidriensis.
On these grounds he provisionally made it the type of a new genus, Hesperoloxodon. Again the discovery in 1926
and 1928 at Steinheim on the Murr, Germany, of two crania referable to Hesperoloxodon antiquus germanicus

convinced him more then ever of the distinction of the "Elephas antiquus" group from the Palxoloxodon group.
Taken together, these crania furnished the basis for determining the evolutionary history and relationships of the

classic Elephas antiquus, definitely assigned by Professor Osborn in the present Memoir to the genus Hespero-
loxodon.
Hesperoloxodon, signifying 'loxodont of the west,' of the European continent. Cranium domelike
with flattened forehead more hypsicephalic and bathycephalic than that of Palxoloxodon lacking the
; ;

prominent parietofrontal crest so distinctive of P. namadicus. Occiput relatively narrow and high
(broad and low in P. namadicus). Premaxillaries extremely broad, with widely divergent tusks, slightly
upcurved and incurved. Mandibular rostrum abruptly truncated symphysis wide. Molars hypsodont, ;

'loxodont sinus' vestigial or absent. Ridge formulae progressive from 3 Jevfrrr (typical), to 3'^~| M M
{germanicus), to M 3^
(italicus). Enamel plates relatively thick, more or less crimped or sinuous.
Cement present.
Dental formula: I "^ Dp ^^ i^ M
Ridge-plate formula (Hesperoloxodon antiquus): Dp 2| Dp 3| Dp 4;^ 1^^ 2 J| 3;^;i^ M M M
—
Horizon. Upper Pliocene' to Upper Pleistocene. Compare pages 1542, 1543 of the present
volume, also figure 1222, and PI. xxiii.
'[Possibly Lower Pleistocene (see footnote 1 on p. 10-19 above). — Editor.]
1594 OSBOKN: THE PROBOSCIDEA

In a comparison of the crania of Loxodonta africana, Palseoloxodon namndicus, and Hesperoloxodou antiquus,
we observe especially the following characters: Short and extremely broad rostrum; premaxillaries diverging to
the iK)int where the tusks issue from the skull— thus the bases of the tusks are very far apart instead of being close
together as in Mammonteus primigenius or relatively close as in E. indicus; whereas the premaxillary sockets are
relatively of the same length in all three species, the divergence of the sockets in P. tiamadicus is about the same
as in L. africana; the premaxillary sockets are relatively longer and diverge still more widely in H. antiquus; the
crania of both P. namadicus and H. antiquus are distinguished from the cranium of L. africana by the greater
development of the fronto-occipital crest which in P. namadicus engulfs the frontal bones so that there is a very
short space between the lower border of this crest and the extremities of the nasals and the narial openings; the
narial openings are extremely broad and shallow, they exhibit approximately the same hour-glass-shaped form
in L. africana, P. namadicus, and FI. antiquus; the skull of P. namadicus, moreover, is relatively broader and more
flattened or i)latycephalic than the skull of L. africana; this broadening and flattening of the summit of the cran-
ium reaches an extreme in the gigantic P. namadicus cranium (cf. Chap. XIX, p. 1209 above).

Distribution of Living Elephants: Loxodonta and Elephas

Lii^iii^ Elephants W:4m Fossil Elephants and Mastodonts

I'lo. 1212. WoHLDWiDE Distribution of the Pkoboscidea in Past and Present Time
This map on a larger scale constitutes the back end-paper of the present volume
.African Elephant: After Blanc, 1897; Bocage, 1890; Buckley, 1870; Chapman, 1808; Chubb, 1909-1919; Claridge, 1915; Cox, 1900; Heilprin, 1887;
Hippolyte, 1907; Johnston, 1907; May don, 1932; Roosevelt and Holler, 1914; Schultze, 1907; Sdater, 1900.
Indian Elephant: After Bartholomew, 1911; Blanford, 1888-1891; Blyth, 1872; Bishop, 1921; Champion, 1928; Hornaday, 1885; Hunter, 1808;
Jerdon, 1874; Laufer, 1925; Lydekker, 1900 1908; Sanderson, 1907; Sdater, 1899.

[Since Professor Osborn's intensive studies on the subject, recent observations by Pore Teilhard de Chardin, Dr. C. C. Young, Dr. Glover Allen (letter of
Dec. 24, 1938, to Dr. Pinkley), and Dr. CJeorge Pinkley have led to certain modifications of the map sliown in Volume 1, figure 0, namely, the area in Borneo,
as well as that in southwestern China, has been reduced somewhat. See Ch.aptor XX, p. 1;J02 above, for the views of Pi'to Teilliard and Doctor Young regard-
ing the e.\i.stence in historical times of the wild Indian elephant in China. —
Editor.)
 —

SUMMARY 1595

Migration of the Loxodontines

—
Loxodonta. The eighteen existing species and subspecies or varieties of Loxodonta africana (partly distin-
guished by size, partly by geographic distribution, and partly by the shape of the ears) are found from the Cape
northward to the southern border of the Sahara and the Sudan (see Fig. 1055). Fossil remains are principally of
African provenance, that is, from Zululand (L. zulu) and the Transvaal (L. prima, L. subantiqua, and L. obliqua),
although two or three locaUties in Europe have yielded types, e.g., Germany {E. prisons Goldfuss = L. africana)
and Italy (L. cornaliae), as shown on the accompanying plate (PI. xxiii).

Palseoloxodon. —The Palseoloxodonts, as we have just seen, are divided into two branches, (1) Palxoloxodon
(syn. Pilgrimia), embracing such species as atlanticus, jolensis, and recki from the northern and east-central regions
of Africa, and transvaalensis, sheppardi, andrewsi, hanekomi, mlmani, kuhni, and archidiskodontoides of
yorki,

South Africa, as well as the Mediterranean species melitensis, falconeri, mnaidriensis, lamarmorae, Cypriotes, and
creticus; and (2) Palseoloxodon (syn. Sivalikia), embracing the namadicus of Falconer and Cautley, type locaUty
the Nerbudda, India, the hysudrindicus of Java, and the various (possibly eight to ten) subspecies of Japan,
represented by the genotype of Palseoloxodon, namely, Elephas namadicus naumanni Makiyama. It would
appear, therefore, that the phylum Palxoloxodon originated in Africa, migrated northward into Europe, also
northeastward, through the Mediterranean Islands, leaving dwarfed descendants resembling either the 'E.
antiquus' group or the namadicus' group, finally arriving in southern and .southeastern Asia developing into
'£".

the 'Elephas namadicus' group, thence eastward to Java and northward to Japan, but as far as we know never
reaching America.

Hesperoloxodon. — Members of the 'Elephas antiquus' group are recorded from Italy {Hesperoloxodon antiquus
nanus, H. antiquus ausonius, H. antiquus from Spain {H. antiquus platyrhynchus) from England (the
italicus), ,

true H. antiquus especially), from Germany (represented by the H. antiquus germanicus of Steinheim), and from
Rumania (by H. antiquus germanicus ref. from Ilfov). Professor Osborn states in Chapter XIX, p. 1258 that some
of the narrow-toothed varieties of Elephas [Hesperoloxodon] antiquus in Europe may be related to the dwarfed
insular elephants; and again on page 1252 he calls attention to the "lofty cranial dome" of Hesperoloxodon antiquus

italicus which resembles that of the Elephas indicus of India and is in close agreement with the "Elephant trace
en rouge" in the cavern of Pindal, also with the anterior dorsal hump of the Spanish and Algerian elephants
(Fig. 1047).

Superfamily: ELEPHANTOIDEA Osborn, 1921. Family: ELEPHANTID^ Gray, 1821

Subfamily. ElephantinvE Osborn, 1910

Genera: Elephas Linn^us, 1735-1758; Hypselephas Osborn, 1934, 1936; Platelephas Osborn, 1934, 1936

(Cf. Chap. XX of present volume, also p. 1543, and figures 1173, 1221, 1228, and PI. xxii)

The history of the elephant in art, industry, and science is fascinatingly told by Dr. George F. Kunz in his
book on "Ivory and the Elephant." The early life phases, however, are so beclouded by superstition and myth

that we are without positive knowledge of these interesting animals until the time of Homer. Strangely enough,
also, the fossil ancestry of the Hving Asiatic elephant is still unknown, despite the untiring efforts of palaeontolo-

gists and zoologists everywhere to solve this problem. Consequently the subfamily phylogeny of most of the
elephantoids is better that of the Elephantina?, or, as a matter of fact, of the Loxodontinse, for, as
known than
noted in the preceding section, there is still uncertainty as to the ancestral forms of the living African elephant.
Professor Osborn was of the opinion that "the conclusion is inevitable, that in some as yet unexplored region of
1596 OSBORN: THE PROBOSCIDEA

Asia the direct ancestors of E. indicus were slowly evolving, while in some unexplored area, probably west of
central Africa, the very conservative ancestors of L. africana were also slowly evolving." As our knowledge stands
at present, the Indian elephant suddenly appears fully formed during the Age of Man.

The nomenclatural problems in connection with the genotype of Elephas are no less complex. After long
consideration Professor Osborn adopted Elephas indicus rather than Elephas maximus in the present Memoir,
giving his reasons therefor in Chapter XX, pages 1308-1311 above.

Falconer, in his discussion of the unity or plurality of species among the existing Asiatic elephants, namely,
types from Ceylon, Nepal, Bengal, and Sumatra, decided in favor of the unity of the species Elephas indicus
(Falconer, 1868, Vol. II, p. 269): "the evidence in every aspect appears to fail in showing that the Elephant of
Ceylon and Sumatra is of a species distinct from the Continental Indian form . . . The result of this range of

observation, combined with long osteological study, has been to establish the conviction in my mind that there is

but a single species of Asiatic Elephant at present known, modified, doubtless, according to his more northern
or southern habitat, but not to an extent exceeding that of a slight geographical variety." He admits that the
Ceylon elephants are occasionally imported into Bengal but does not admit, as claimed by Schlegel, that this is an
explanation of the variation in the vertebral and rib formulae of the Indian elephants. The present author ex-
pressed his belief (seep. 1315, Chap. XX above) that the "wide variations in cranial and vertebral characters as
well as in dental and dermal characters and in the shape of the external ear support the subdivision of Elephas
indicus into at least four out of the large number (12) of geographic varieties or subspecific forms," namely:

Elephas indicus ceylanicus (cf. Mukna var.)

Elephas indicus bengalensis (cf. Dauntela var.)
Elephas sumatranus
Elephas maximus hirsutus { = E. indicus hirsutus)

Professor Osborn left the question of the continental and insular races and subspecies for future investigation.
He states on page 1329 that the "profound cranial differences which divide the Ceylon and Bengal elephants
from each other as well as from the Sumatran elejjhant in all probability will be found to differentiate the Sumatran
and other still undiscovered extinct types"; that by "comparison with the evolution of other Pleistocene un-
gulates it appears probable that a very long period of time separated these continental and insular subspecies and
species from each other, a period of time equivalent perhaps to nearly half of Pleistocene time or 500,000 years,
during which through isolation and segregation the subspecific and specific characters were thoroughly founded.
Here again monographic research is essential before we can reach a final conclusion."

As to the extinct forms, it is remarkable that no fossil Pliocene ancestors of the recent Indian elephant have
as yet been discovered. Elephas hysudricus found "below the conglomerates" in the upper Siwaliks of India,
according to Dr. Barnum Brown, shows few resemblances in the cranium to the E. indicus (Dauntela var.) and
no very marked resemblances in the grinding teeth; it appears to be unique, and in 1936 (see Vol. I, p. 12) it was
made one of the types of the genus Hypselephas Osborn.

The third genus included within the subfamily Elephantina? is the Platelephas of Osborn, genotype Elephas
platycephalus, a very ancient and primitive animal. The inclusion by Professor Osborn of these three genera in
the subfamily Elephantinae was provisional, because Elephas hysudricus Falconer and Cautley and E. platycephalus
Osborn are at [)resent known chiefly by cranial characters, our knowledge of the dentition of platycephalus being
confined to the third superior molar only; they appear, however, to represent generic or subgeneric phyla distinct
from the true Elephas.
 — : — : ;

SUMMARY 1597

Characters of these three genera, as far as they are known

Elephas Hypselephas Platelephas

Modernized elephant of India. Highly Primitive elephant of India with elevated Elephant of India with flattened cranium,
progressive and distinctive in cranial cranium primitive
structure

Typified by Elephas indicus Typified by Elephas hysudricus Typified by Elephas platycephalus

Cranium bathycephalic, cyrtocephaUc, Cranium hypsicephalic, brachycephalic Cranium relatively elongate, dolichoce-
hypsicephalic, occipitofrontal dome condyles well raised above grinding phalic, platycephalic, occipital con-
more or less rounded, not acute, with surface of molars, occiput elevated dyles not greatly elevated above level of
expanding diploe, frontals gently con- with broadly transverse rugose frontal grinding surface of molars; deeply in-
cave crest, relatively flat, sloping backward, dented supra-occipital border, without
frontals deeply concave rugosity

Premaxillaries relatively narrow, sub- Premaxillaries relatively narrow or later- Premaxillaries greatly elongated in front
vertical. Orbits large, relatively ele- ally compressed. Orbits large, depress- of superior third molars, somewhat
vated, i.e., directly opposite occipital ed, near maxillary rostrum, unlike divergent. Ro.strum somewhat broad.
condyle Loxodonta or Elephas Orbits large, elevated, near frontal
profile, set widely apart. Posterior
nares deeply indented

Superior tusks relatively straight, in- Superior tusks relatively straight, in- Tusks imperfectly known
curved, upcurved with bases of tusks curved, somewhat divergent at base.
relatively close together. No inferior No inferior tusks
tu.sks

Mandible with extremely abbreviated Mandible with elongate, prominent sym-

symphysis. Ramus deeply depressed physis; ramus shallow

Molars of intermediate breadth, hyp.so- Molars low crowned, long, narrow; ridge- Molars imperfectly known relatively low,
:

dont (less extreme than in Parelephas, plates with pUcated enamel, convexo- ridge-plates directly transverse, as in
Archidiskodon, or Mammonteus); ab- concave, reversed above and below; Elephas. No rudiment of 'loxodont
sence of 'loxodont sinus' {E. indicus); widely separated, with cement filled sinus.' Ridge-plates broad, depressed,
moderately compressed enamel ridges valleys (juvenile); trace of a median limited in number: M3*^^*
of intermediate thickness, extremely 'loxodont sinus.' Ridge-plate formula:
crimped or sinuous in E. indicus. M —
l»± 3
>-l±
"17-18-19
Ma.ximum number of plicated ridge-
plates: M 3„-A|^

Dental formula: I ^^^ Dp |ti M^

Ridge-plate Formulae

Elephas {E. indicus, fide Falconer): Dp 2t Dp 3^ Dp 4; J M Iff M 2;^ M S^^

Hypselephas (E. hysudricus) :

(Falconer, 1868): Dp 3^ 4±^ M l||^ M 2^-^^ M Srfrn

Dp
(Osborn, 1930): Dp 3^, Dp 4f| M Ij^ M 2^^^^, M 3^

Platelephas (as far as known) : M 3^^

HORIZON. Elephas, Upper Pleistocene and Recent. Hypselephas, Lower Pleistocene. Platelephas, Upper Pliocene or Lower
Compare page 1543 of this volume, also figures 1221, 1228, and PI. xxii.
Pleistocene.
 1598 OSBORN: THE PROBOSCIDEA

CoMP.UtlSON OF HyPSELEPHAS AND PlATELEPHAS WITH ElEPHAS

With the exception of the decidedly low position of the orbits, the front view of the Hypselephas hysudricus
cranium bears a closer resemblance to the broad-narial variety Elephns indicus bengalensLs than to the narrow-
narial variety E. indicus ceylanicus; also the low, narrow grinding teeth of the type of H. hysudricus are
entirely different in proportion from those of E. indicus, they exhibit some resemblance or analogy to the extinct
Palseoloxodon of Africa both in their long, narrow proportions and in their 'loxodont sinus.' The juvenile cranium
of H. hysudricus resembles somewhat closely the juvenile cranium of E. indicus at the time when the fourth de-
ciduous molars are in use. From cranial and juvenile characters alone one would be inclined to regard H. hysudri-
cus as an early offshoot in Lower Pleistocene times of the main stem which gave rise to E. indicus of recent times,
hut the less rugose orbits in a similarl}- depressed position as that seen in crania in the British and Amherst
Museums referable to H. hysudricus, that is, lying directly upon the premaxillary sockets of the tusks instead of
being raised above the tusk sockets, demonstrate afresh that H. hysudricus is not to be regarded as ancestral to
the collective Elephas indicus type but that it belongs in a totally distinct phylum or line of descent. In short,
//. hysudricus had a modernized cranium with a relatively primitive condition of tlie molars.

The Platelephas platycephcdus cranium is of very primitive elephantine affinity —low and flattened —widely
different from the elevated Hypselephas hysudricus or the greatly elevated Elephas indicus crania. Both in the
cranium and the grinding teeth it is also profoundly distinct from those of Archidiskodon planifrons. Therefore,
Platelephas platycephalus, in the opinion of the present author, was not ancestral to either H. hysudricus or E.
indicus, nor was it related to Archidiskodon; it would seem to belong to a primitive stage in the evolution of the
ElephantiniP. The specific name has reference to the highly characteristic and primitive lowering of the fronto-
occipital profile and the placing of the occipital condyle only slightly above the horizontal level of the maxillary

border of the superior grinders.

# ^€
(Sir
«:«m:<m
'liliii;i[

/«
"nil
 SUMMARY 1599

In an article by Mr. George G. Goodwin in the Journal of Mammalogy, November, 1925, the date of the
landing in the United States (at New York) of the first Asiatic elephant is given as April 13, 1796, on the ship
"America" of Salem, Massachusetts.

Fig. 1244. Refeneil Jilephas mdicus showing contrast in proportions be-

tween adult and young. Mother, six montlis after capture, and her baby,
one month old. Photograph reproduced through the courtesy of Underwood
and Underwood of New York.
 :

1600 OSBORN: THE PROBOSCIDEA

4. SKELETAL MATERIAL
Inasmuch as the axial and appendicular skeleton of the Proboscidea has been touched upon but briefly in the

foregoing pages of this chapter, it may be helpful to summarize the skeletal material mentioned in the present
Memoir, noting certain important observations made by Professor Osborn.
MffiRITHERIOIDEA
Mieritherium Skeleton partly known (Fig. 43, and for restorations see Figs. 19, 41, 42, 51, also Pis. x and xiv). Verte-
bral structure indicates an ambulatory and amphibious habit. Limbs relatively short and stout.
Vertebrae estimated: Cervicals 7, dorsals 19-20, lumbars 4, sacrals 4. Length of humerus 240-260 mm.
(distally of jirimitive locomotor type). Femur straight with ligamentum teres pit, greater and lesser
trochanter, absence of inner trochanter. Proportions of ulna and radius not fully known. Pelvis,
scapulae, and girdles primitive, with analogies to the sirenian type, not expanded as in the proboscidean
graviportal type. Probably pentadactyl.

DEINOTHERIOIDEA
Deinotherium Skeleton almost completely known in the Franzensbad and Rumanian specimens (pp. 99, 100, also 97
and 98 respectively; for restorations see Figs. 65, 66, 67, 70, 72, 73, and Pis. x and xiv). Limbs
elongate, increasingly elephantoid, raising body well off the ground. Humerus 815 mm., ulnaSOO mm.,
—
femur 1055 mm. {D. bavaricum Franzensbad skeleton). D. gigantissimum ref. of Rumania, including
two two scapulae (one measuring 1150 mm. in diameter), also vertebrae, four limbs, eighteen ribs,
iliae,

as well as tibia and pes, was probably double the size of D. bavaricum. Feet reduced to three short
functional digits in manus and pes, with vestigial D.I in the pes. Dorso-lumbar vertebrae and trunk
abbreviate.
MASTODONTOIDEA
Trilophodoit Referring to the mounted skeleton (Fig. 199) in the Paris Museum, Galerie de Paleontologie, there is
some doubt as to the reference of this specimen to Trilophodon only there is a possibility that some of
;

the parts may be referable to Senidentinus. It is certainly an animal of larger size than Serridentinus
productus in the American Museum, according to Dr. W. D. Matthew. There is, however, a fairly
complete skeleton of Trilophodon giganteus in the American Museum (Figs. 257 and 372), the important
measurements of which are as follows
Humerus,

Phiomia

Amebelodon

Megabelodon

Tetralophodon
 :

SUMMARY 1601

Morrillia Skeleton unknown. For restoration see PI. xv.

Cordillerion The restorations of Cordillerion are also based on the characters of the skull, especially the superior tusks,
and on the lower jaw. For restorations sec figures 75, 484, 498, 500, 502, 503, and Pis. x and xvi.

lihynchotheriuin The only skeletal parts known of the genus Rhynchotherium are those of the adolescent neotype of Rhyn-
Blickotherium chotherium shepardi edense, consisting of the humerus (660 mm.), femur (860 mm.), radius (500 mm.),
Aybelodon tibia (490 mm.); also metacarpals and metatarsals 1-5. The restorations of Rhynchotherium, Blick-
otherium, and Ayhelodon are based on these fossil remains as well as on the lower jaws; no complete
cranium is known. For restorations see figures 75, 447, and Pis. x and xvi.

Two skeletons of Anaticus arvernensis from Italy are in the Turin Museum and Royal University of
Pentalophodon Bologna respectively. The Turin .specimen (Fig. 584) is "deficient only in the cranial portion of the
Synconolophus head, right hind-leg, part of the scapula and pelvis, and some of the bones of the carpus and tarsus.
The upper and lower jaws, with the tusks entire to their tips, are preserved." Falconer says in regard
to the Bologna specimen (Fig. 585) that it is nearly as perfect [as the Turin specimen].
Skeleton of Pentalophodon and of Synconolophus unknown.
For restorations see figures 75, 582, 583, 594, 595, 621, and Pis. x and xvi.

Cuvieronius Skeletal remains of Cuvieronius superbus and C. platensis furnished the basis of the restorations in the
present Memoir. The outlines of the crania and limbs in figure 566 rest entirely on the largest of these
specimens in the La Plata Museum
C. superbus C. platensis
1030 mm.
795
750
780

Other parts include the atlas, axis, and first and third dorsals. There is also a fine skeleton of Cuvieroni-
us postremus from Ecuador in the Museos de la Uni^'ersidad Central del Ecuador (Fig. 533) : Femur
(890 mm.), humerus (740 mm. ), tibia (505 mm. ), radius (630 mm.). For restorations see figures
75, 485, 498, 502, and Pis. x and xvii.

Stegomaslodon In the San Pedro valley of Arizona was found a portion of a young adult male skeleton of Stegomaslodon
(S. arizonse) now in the U. S. National Museum, consisting of "the base portion of the skull with tusks
and cheek teeth in place; the lower jaws; both fore limbs (one with nearly complete foot); parts of
both hind limbs; the pelvis; and several of the more characteristic vertebrae and ribs." For skeleton
(see Fig. 634), for restoration (Fig. 635). Humerus (850 mm.), femur (1010 mm.), scapula (765 mm.),
width of pelvis (1475 mm.). For other restorations see figures 75, 498, 642, 648, and Pis. x and xvii.

Eubelodon The following are measurements of the type pelvis and of referred specimens of Eubelodon morrilli in the
Nebraska State Museum Pelvis (type), extreme width 1424 mm. referred femur (940 mm.), humerus
: ;

(760 mm.), radius (610 mm.), fibula (545 mm.), ulna (685 mm.), scapula (760 mm.). See figures 573
(skeleton) and 577 (restoration) also figures 75, 498, and Pis. x and xvii (restorations).
;

Serridentinus The reproductions of Serridentinus produclus (Figs. 371, 372) give only an approximate idea of the com-
plete skeleton. The original skull and jaws are mounted with the skeleton; also thirteen vertebrae,
most of the ribs, pelvis and sacrum, the right hindlimb bones with the exception of the foot and some
of the tarsals are original bone. The feet are all restored, excepting the left hind pes.
 :

1602 OSBORN: THE PROBOSCIDEA

Torynobdodvn Skeleton unknown, liestoration based on mandible in the Nebraska State Museum (see Y'lfx,. 423 as
—
Plaiybelodon baniumbrou'ni finally belie\ed by Professor Osborn to be referable to Torynubclodon).
See also PI. x, where it appeared as Platybelodon barrmmbrowni, and PI. xviii as Torynnbelodun
barnumbroivni.

Palseoniastodon See figure 96 for limbs and vertebne oi Palxomadodon bcadnelli, namely, axis vertebra, seapula, humerus,
femur, and tibia. Important because of certain primitive, Mastodon-like characters. The restoration
(Fig. 97) wii.s based on these remains. See also figures 41, 41a, 110, and Pis. x and xix.

Maslodoti The largest fidly adult male skeleton of the true Mastodon (M. aniericanus) is the mount in the Ohio
State University (Fig. 114), with a skeletal height of 2920 mm. or 9 ft. 7 in., and a height in the flesh of
3105 mm. or 10 ft. 2% in. This, together ^vith the complete M. americanus skeleton (Fig. 124) in the
American Museum, known as the '"Warren ]\Iastodon," with a skeletal height of 2780 mm., or 9 ft. 1^
in., and a height in the flesh of 2956 mm., or 9 ft. 9 in., furnished the measurements on which the restora-
tions in the present Memoir were based. See also for skeleton, figures 82, 83, and 125; for restorations,
figures 109, 110, 136, and 158, also Pis. xandxix. The following are the principal measurements of the
"Warren Mastodon"
Humerus

Pliomastodon

Miomastodon

Zygolophodoii
Turicius

Slegolophodoii

STEGODONTOIDEA
Slegodon

ELEPHANTOIDEA
Archidiskodon
 SUMMARY 1603

Parelephas Well represented both in Europe and America by skeletal remains. P. intermedim in the Lyons Mu-
seum (Fig. 944) is a complete skeleton and the beautiful frontispiece to Chapter XVII (/-". trogonthcrii)
is partly based on this skeleton, but chiefly on the superb P. jeffersonii in the American Museum
(Figs. 931, 966, and 988), the follo\\ing parts of which, however, are mi.Ksing and restored in plaster:
Radius and ulna of both sides, right tibia and fibula, and both fore- and hindfeet. The length of the
femur is 1250-1255 mm., of the humerus 1085-1120 mm. A record humerus is that of P. trogovtherii of
Germany which measures 1480 mm. A fine skeleton of P. columhi from Florida is in the Amherst
Museum; it lacks four dorsals, parts of the limb bones, ilium, manus and pes; otherwise largely
original (Fig. 955). Finally, the P. floridanus skeletal remains are among the most important; this
material, with mea.?urements, is enumerated in Chapter XVII, pp. 1111-1114. For other restorations
see figures 930, 936, and PI. xxii.

Mammonteus There an abundance of skeletal material of the woolly mammoth, namely, from the Kolyma-Beresowka
is
River, Siberia, Steinheim on the Miut, AViirttemberg, and Borna, Germany, also Lierre, Province
d'Anvers, Belgium (p. 1130); the Moravian skeleton from Briinn, Czechoslovakia (Fig. 1007), and the
fine skeleton of M. primigenius fraasi at Stuttgart (Fig. 1018) finally, and perhaps the most important
;

skeleton discovered (as it included a piece of the hide with hair in place) is the Lena River specimen,
known as the "Adams skeleton," which is mounted in the Zoological Museum of the Academy of Sci-
ences, Leningrad (Fig. 1014). In 1929 a splendid lot of cranial and skeletal material was unearthed in the
vicinity of Fairbanks, Ala.ska. This material is listed on page 1160 above. Length of the humeri
980-750 mm., of the femora 1130-965 mm., of the tibiae 680-440 mm., of the radii 610-560 mm. For
restorations see figures 990, 994, 1000-1002, 1013, 1034, and PI. xxii.
Loxodonia As one of the living genera of the Proboscidea there is endless skeletal material. In the American Museum
is the skeleton of "Jumbo" (Loxodonia africana oxyotis) and the Akeley mount of two geographical
varieties of Loxodonia africana, namely, albertensis and peeli. A table at the close of this section gives
the maximum height and other measurements of the African elephant, so that it is not necessary to
enumerate the extensive materials, excepting to mention the length of the humerus (1078 mm. see —
Fig. 912) and of the femur (1246 mm. —
see p. 1230) of "Jumbo," and to call attention to the comparative
figure (Fig. 1084) of the vertebral columns of L. africana, Elephas indicus, Mammonteus primigenius,
Parelephas jeffersonii, and Hesperoloxodon antiquus. According to Falconer (cf. p. 931 above) the
African elephant \'aries in the number of dorsal vertebrae from 20 to 21.
Palseoloxodon The extinct branches of the Loxodontinae are not well represented by skeletal material. Doctor Dietrich
gives the length of the humerus of Palseoloxodon recki of East Africa as 1235 mm., of the femur as 1470
mm. Professor Osborn suggested that the dwarfed elephants of the Mediterranean Islands might be
descended from such forms as P. recki, P. allanticus, etc.
Dwarfed Palaeoloxodonts of the Mediterranean Islands. These animals vary in size; observe particularly
figures 1118 and 1119, in which the large stage (Palseoloxodon mnaidrieyisis) has a humeral length of 573
mm.; the intermediate stage (P. melitensis) has a humeral length of 478 mm., and the small stage
(P. falconeri) has a lumieral length of 270 mm. The skeleton of P. falconeri is fragmentaiy, and P.
lamarmorae from Sardinia is represented by carpal and tarsal bones only. The heights of these insular
species vary from 2 ft. 6 in. to 6 to 7 feet.
The Palseoloxodon phylum culminates in the P. namadicus of the Nerbudda of India, the cranium of which
has been used as a ba.sis for the restorations in the present Memoir (Fig. 1068). Pilgrim, 1905, ])]). 211,
212, gives the measurements of certain skeletal material which he recovered from the Godavari beds and
regarded as referable to this species, but it is somewhat fragmentary; it consists of portions of a femur
and of the pelvic bones; the cranium and tusks are very fine. For other restorations, .see figure 1127,
and PI. XXIII.

HesTperoloxodon The most complete skeleton of Elephas [Hesperoloxodon] antiquus is in the British Museum, although it
lacks the cranium (Fig. 1080). It is important to note that .since the discovery of this Upnor specimen,
there have been several crania described, among them that of Palseoloxodon [Hesperoloxodon] antiquus
italicus from Italy (Fig.s. 1105-1108) and two of Hesperoloxodon antiquus germanicus from Steinheim
(Fig. 1114), affording the much-needed knowledge regarding the skull and tusks of this ancient elephant
of Europe. The Upnor and Pignataro specimens furnished the necessary data for the restoration (Fig.
1074). The skeleton of the Upnor //. antiquus is shown in figure 1079; as noted in the caption, the
backbone lacks two \'ertebrae, the radius of the left fore leg is restored in plaster, the right radius is
complete. Fortunately the humerus, although not quite complete, was present and measures 1290 mm.
Of the Pignataro Interamna specimen (//. ant. italicus) the right scapula is preserved. A femur of the
large Taubach skeleton of H. ant. germanicus measures 1600e mm., and another from Rome of H. ant.
italicus measures 1500e mm., according to Pohlig (cf. p. 1251 above). See figures 1068, 1083, 1092, and
PI. XXIII for restorations.
Elephas Like Loxodonta the skeleton of the living Indian elephant (Elephas indicus) is well known. The average
length of the humerus varies from 816 mm. to 895 mm. in three individuals (Fig. 1194). According
to P^alconer (cf. p. 931 above) E. indicus, including the continental and insular varieties, varied in the
number of dorsal vertebree from 19 to 20.
Hypselephas The cranium, dentition, and tusks only of Elephas [Hypselephas] hysudricus are known.
Platclcphas Up to the present time the skeleton of Platelephas is unknown. It is represented by the type cranium and
imperfectly known grinding teeth. The tusks also are unknown.
1604 OSBORN: THE PROBOSCIDEA

5. HEIGHTS OF PROBOSCIDEANS, ESTIMATED AND ACTUAL

In a recent monograph (1936) on "The Physiology by Dr. Francis G. Benedict, Director
of the Elephant," of

the Nutrition Laboratorj^, Carnegie Institution of Washington, much accurate and important information is

given regarding the heights and Aveights of the elephant, especially of the Indian elephant. It is stated on page
290 that the weight of a "large" elephant is over 8,000 pounds or 3630 kilograms. The weight of "Jap" (the
female Indian elephant used in the physiological experiments on which much of Doctor Benedict's conclusions are
based) is given as 8095 lbs. or 3672 kg. (see p. 104). The weight of "Khartum" (a gigantic male African elephant
formerly living in the New York Zoological Park), on which an autopsy was made by Dr. W. Reid Blair and Dr.
C. V. Noback, was about 10,040+ lbs. or 4710 kg. (see p. 106).

The following is a list of the heights (in millimeters and feet, estimated and actual) of tlie Proboscidea, as
recorded in the present Memoir.
SUMMARY 1605
1606 OSBORN: THE PROBOSCIDEA

CONCLUSION
The proboscideans are geologically long lived —55,()()0,0()() to Go,()00,()()0 years (Osborn, 1933.901, p. 159),
originating (Osborn, 1934.924, p. 177) "exclusively in Africa, probably inUpper Cretaceous or Lower Eocene
time." According to Professor Osborn (1934.922, p. 211) they "rank next to man in biological importance, and
far surpass the mechanically inferior man in demonstration of all the main principles of biomechanical aristo-
genesis and alloiometry."
 o

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ADJACENT TEBRITORY.
) 10 go 30 40 50 60 70 60 9 100 MM fS copic struc-
number of
nent dental
on of them,
ults desired

al intention

phs than as
had not yet
Aemselves,
only to the
inadequate
le shapes of

s that have
subject the
'graphs has
Anderson's
e benefit of

Anderson,
e polarized

nining the
)ut, by the
so clearly
lich can be
in subject,

d either its

distinctive
may be of
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es radically

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'ing animal

Ervvin J. Raisz A.M.

)y the American Museum.

 MAP OF
~V NORTHWESTERN
AND
INDIA
^ V ADJACENT TERRITORY.

Il

^"^^^^^ OF NATURAL HISTORY

CoSh*!^
irom the
192/
Survey of India I loooooo sheets Nrs, 38,39 43.4A. 52. 53.

and the adjacent Salt Range and the important localities where vertebrate fossils have been recovered by the American Museum.
Plate XXV. Map of Northwestern India showing the Siwalik Hils
 APPENDIX TO VOLUME II

PROBOSCIDEAN DENTAL HISTOLOGY

By George Gaylord Simpson

It was Professor Osborn's intention to include in this Memoir an appendix devoted

to the microscopic struc-
ture of proboscidean teeth. For purpose he had Mr. A. E. Anderson prepare and photograph a number of
this
thin sections of such teeth and write notes on some aspects of their stud3^ It was intended that an eminent dental
histologist, the late Dr. J. Leon Williams, should also study these sections and prepare an interpretation of them.
Unfortunately no part of this plan was carried far enough for it to be possible now to achieve the results desired
without a vast amount of further research impossible to us at this time and going far beyond the editorial intention
of publishing the work done under Professor Osborn's direction without substantial change.

Mr. Anderson's notes were intended and are useful rather as a guide for the study of his photographs than as
Dr. WiUiams prepared a few additional photographs of the same sections, but had not yet
results of such study.
supplied an accompanying manuscript when he died suddenly on February 23, 1932. The sections themselves,
and the photographs of them, pertain to only three genera, Elephas, Phiomia, and Trilo-phodon, and only to the
tusks of these except for two sections of a Trilo-phodon molar. Even within this limited sphere, they are inadequate
to reveal all the important details. For instance, there are no tangential sections and without them the shapes of
the enamel rods cannot be determined.

Despite this very fragmentary character of the data left, it includes photographs and observations that have
not been made obsolete by any pubhcation in the meantime and that do give at least a glimpse at a subject the
complete omission of which would be a fault in this Memoir. A selection of the most important photographs has

therefore been made them have been prepared in part from Mr. Anderson's
for publication here, legends for

notes and in part from independent observation, and this introductory comment written, all without the benefit of
Professor Osborn's direction but making available material that was in his hands.

With a by Dr. Williams) these photographs were made by Mr. Anderson.

single exception (PI. xxviii, Fig. 2,

A few were made with ordinary Ught, but most by means of a petrographic microscope, some in plane polarized
light, and some between crossed nicols. The use of crossed nicols is not only valuable for determining the
optical properties and mineralogy of the tissues, a matter of secondary interest here, but also to bring out, by the
phenomena of birefringence, details of tissue structure and orientation that are not visible, or not so clearly
visible, in ordinary or in plane light. This
a technique of considerable importance, the value of which can be
is

seen, for instance, by comparing figures 3 and 4 of Plate xxvii. Although the two are identical in subject,
each reveals important structural features not visible in the other.

The histology of a tooth is an important part of its structure. It is impossible really to understand either its

gross structure or its functional characters without knowing something of its microstructure and of the distinctive
features of the tissues composing it. There is also good reason to believe that histological characters may be of
value in the identification of fossils and in the determination of animal affinities. Dental tissues individually and
in combination have complex characters, fairly constant within limited taxonomic groups and sometimes radically
distinct in different groups. Some of these distinctions seem to be adaptive or habitus characters, but others are

probably deep-seated heritage characters and they may prove to be exceptionally reliable helps in tracing animal
descent.
1607
I(i08 OSBORN: THE PROBOSCIDEA

The present material gives examples of proboscidean dental tissues that must be more or less typical of that
group. This purely descriptive aspect is sufficiently expressed by the photographs and their legends. These data
also show that for the three genera in question the microstructure even of a simple transverse section of a tusk is

determinative. Elephas, Pkiomia, and Trilophodon differ in dental histology and on this basis alone any one of
these genera can at once be distinguished from the other two. This fact should stimulate the preparation of the
many what extent thin sections can be useful in the diagnosis and
additional sections necessary to determine to
recognition of taxonomic groups of Proboscidea. The histology of these three genera also shows that they have
many characters in common, some of which do not appear to be shared by more distantly related animals. More-
over Phiomia and Trilophodon resemble each other more than either resembles Elephas. These facts warrant and
urge further study to see how far such resemblances can be used to determine relationships.

Although inadequate to establish them definitely, these imperfect data also hint at evolutionary trends of
considerable interest functionally and in other ways. This is seen, for instance, in the fine structure of the tusk

enamel. In Phiomia (e. g., PI. xxviii, Fig. 2), the rods have a pronounced spiral arrangement which makes for

toughness and resistance to injury, suggestive of the very tough enamel of rodent incisors. In Trilophodon (e.g.,

PI. XXIX, Fig. 3), this arrangement persists, but in appreciably weaker form. In Elephas (PI. xxvi, Fig. 3), it has
practically disappeared and the enamel is weak in histologic structure. Thus these three genera, at least, show
progressive simplification and weakening of the enamel in microscopic characters as it becomes more reduced
macroscopically and less important functionally. The peculiar enamel stringers in Trilophodon and the relation-
ships of cement in Trilophodon and Elephas also suggest potentially fruitful lines of investigation. Another will be
found in the evolution of the "engine-turning" effect of the dentine and another in the probable changes in enamel
structure in the molars, with changes from the primitive condition to the heavy cones of some later mastodonts
or the lamella^ of the elephants. Another of the more interesting of the many })oints merely glimpsed in these

sections is the possible relationship between the dentine cylinders of some mastodonts, such as Plalybelodon,
and the peculiar dentine modification seen in the most juvenile part of a tusk of Elephas (PI. xxvi. Fig. 1).
 CATALOGUE NUMBERS OF SPECIMENS MENTIONED IN THE PROBOSCIDEA MEMOIR, VOLUME II

Algiers: School of Sciences Los Angeles: Museum of History, Science and Art
Amherst: Amherst College Museum Lyons: Museum of Natural Sciences
Amsterdam: Zoological Gardens

Marseilles: Museum of Natural History

Berlin: Geological-Palseontological Institute of the University
Mexico: Engineering School
Museum for Natural History
Geological Institute
Bologna: University Museum
Bonn: Museum Moscow: University
Brno: Moravian Government Museum Munich: State Zoological Museum
Brussels: Museum
Bucharest Laboratory of Geology of the University
:

Laboratory of Palaeontology of the University

New York: American Museum of Natural History
Zoological Society
Buenos Aires: Argentinian Museum of Natural Sciences
Norwich: Norwich Castle Museum and Art Gallery

Calcutta: Asiatic Society of Bengal

Indian Museum Oran: Museum
Canterbury: Museum Oxford: Museum
Capetown: South African Museum
Chicago: Field Museum of Natural History
College: University of Alaska (formerly Alaska Agricultural College and Paris: Museum of Natural History

School of Mines) Pasadena: California Institute of Technology

Columbus: Ohio State University Museum Philadelphia: Academy of Natural Sciences

Cromer: Savin Collection' American Philosophical Society

Wagner Free Institute of Science

Florence: Natural History Museum Private Collections: Walter W. Holmes, Florida

Fiikuoka: Kyushu Imperial University Kato. Kokubo, Japan
Percy Madeira, Philadelphia
Natsume, Minato, Japan
Geneva: Museum of Natural History
Due Wood Norton, England
d'Orleans,
Gottingen: Zoological Institute of the University
C. V. A. Peel, Oxford, England
Gotha: Museum
Powell-Cotton, Birchington, England
Lord Rothschild, Tring, England^
Halle: Museum

Iowa City: University of Iowa Quito: Museum of the Central University of Ecuador
Ipswich: Natural History Museum
Rome: University Museum
Jena: Museum Rotterdam: Zoological Gardens

Kimberley: McGregor Museum Saint John, New Brunswick: Museum

Kobe: Takikawa Middle School Sendai: Tohoku Imperial University
Kyoto: Imperial University Sendai (probably): Second High School
Stockholm: Museum of King Adolph Frederick
La Plata: Museum Stuttgart: Wurttemburg Natural History Collection
Leiden: Dubois Collection
Natural History Museum
Tallahassee: Florida Geological Survey
Royal Museum of Geology and Mineralogy
Tervueren: Museum of the Belgian Congo
Leipzig: Palaeontological Collection of the University Tokyo: Geological Institute of the Imperial University
State Ethnographical Museum (Grassi Museum) Museum of Education at Ochanomizu
Leningrad: Museum of the Institute of Mines, Academy of Sciences of the Uyeno Museum
U.S.S.R.
Turin: Museum
Palseozoologic Institute of the Academy of Sciences
Zoological Museum of the Academy of Sciences
Lincoln: Nebraska State Museum Vienna: Natural History Museum
Lisbon : Zoological Gardens
London: British Museum (Natural History)
Imperial Institute Washington: United States National Museum
Zoological Park
India House
Museum of the Geological Society Weimar: Natural History Museum
Royal College of Surgeons Rebling Collection
Zoological Society Schwabe Collection

'[The entire collection of vertebrate fossils made by Alfred C. Savin was purchased by the British Museum in 1897. —Editor.)
^[Probably .at the British Museum. — Editor.)
1609
1610 OSBORN: THE PROBOSCIDEA

Algiers: School of Sciences

CAT. NO. PAGE
Palspoloxodon jolensis type 1275

Amherst: Amherst College Museum

25-1 Parelephu, rolnmbi ref 1051, 1052, 1079, 1080, 1081, 1106, 1111, 1112, 1603
Ilypselephnx hysudricus ref 1351, 1353

Amsterdam: Zoological Gardens

Elephas indicus sumatramis ref 1314

Berlin: Geological-Pal.«ontological Institute of the University

XI] 1 711 Palxoloxodon recki ref 1275
'X\'I I 1384 Palxoloxodon recti lectotype 1275, 1276

Berlin : Museum of Natural History

Loxodonta africana cyclolis type 1 193
Loxodonla africana knochenha\ieri type 1193

Bologna: University Museum

A rianciin nrrcrnensin ref 1601

Bonn: Museum
Ma))in)onlciis (?) primigenius leilh-adamsi type 1150

Brno: Moravian Covernment Museum

Mainmonleus primigenius ref 1 128, 1139

Brussels: Museum
Maniniontciis primigenius ref 1130

Bucharest: Laboratory of Geology of the University

Ilesperoloxodon anliquus germanicus type 1235
Archidiskodon planifrons rumanus type 968, 969

Bucharest: Laboratory of Paleontology of the University

Archidiskodon planifrons rumanus ref 969

Buenos Aires: Argentinian Museum of Nati'ral Sciences

44 Cuvieronius superbus ref 1546

Calcutta: Asiatic Society of Bengal

'^Pnhidldxodon nnmadicus ref 1211

Calcutta: Indian Museum

.\ 48 Tetralophcidnn pnnjabiensis cotype . ., 841
A 86 Stegolophodnn skgodontoides type 834, 847
.\ 355 Slegolopliudon cautleyi ref 842
.\ 426 Trilophodon pala^indicus tyy>e 1546
.\ 437 Stegolophodon cautleyi cotype 841, 842

Canterbury: Museum
Uesperoloxndnn anliquus ref 1215
 LIST OF CATALOGUE NUMBERS 1611

Capetown: South African Mitseitm

CAT. NO. PAGE
Loxodonta africana toxotis type 1193

Chicago : Field Museum of Natural History

201616 Hesperoloxodon antiquus germanicus ref 1236, 1237

College: University of Alaska

Mammon te us priniigenius ahiskensis cotype 1159, 1160, 1161

Columbus: Ohio State University Museum

Mastodon americamis ref 1602, 1605

Cromer: Savin Collection'

197 Archidiskodon meridionalis cromei-ensis rof 963, 980, 981, 1155
1240 Porelephas (?) irogontherii sp.? 981, 1155
Hesperoloxodon anliquus ausonius (?) ref 981

Denver: Colorado Museum of Natural History

472 Parelephas washingtonii ref 1103, 1104
1057 Archidiskodon haroldcooki type 1029
1359 Arrhidiskodon meridionalis nebrascensis type 1033, 1035

Florence : Natural History Museum

Hesperoloxodon antiquus ausonius type 1215, 1232, 1233
Hesperoloxodon antiquus ausonius ref 1215
Archidiskodon meridionalis lectotype 938, 970, 972, 975
Archidiskodon meridionalis cotype 971, 977
Archidiskodon meridionalis ref 975, 1 251

Fukuoka: Kyushu Imperial University

Stegodon orienlalis shodoensis ref 893

Geneva: Museum of Natural History

Rhynehotberium tlascalx genotype (cast) 1377, 1419, 1561

Gottingen: Zoological Institute of the University

Mammonteus priniigenius lectotype 1122, 1123, 1141

GoTHA : Museum
Hesperoloxodon antiquus ref 1119, 1122, 1181, 1236, 1365

Halle: Museum
Hesperoloxodon antiquus germanicus neotype 1235

Iowa City: University of Iowa

167 Parelephas jeffersonii or P. washingtonii ref 1088
213 Stegomastodon aftonise type 1548

Ipswich : Natural History Museum

Archidiskodon ? planifrons ref 963, 981, 1155
Parelephas (?) trogontherii sp? 981, 1155
Mammonteus primigenius? astensis ref 981, 1155

Jena: Museum
Hesperoloxodon antiquus germanicus neotype 1234, 1235

'[The entire collection of vertebrate fossils made by Alfred C. Savin was purchased by the British Museum in 1897. —Editor.)
1612 OSBORN: THE PROBOSCIDEA

Kimbeuley: McGregor Museum

''*°*=
CAT. NO. ,

Pals-oloxndon andrewsi typo 99-^> 1278

43o
2930 Palifoloj-ixlon hanekomi type 993, 1279

3682 Archidiskodon broomi type ^^^t 9^9

Metairhidiskodon griqtta genotype 994, 995
3686
3920 Archidiskodon subplanifrons type 987, 988, 1549

4073 Archidiskodon yorki type 992, 993

Palxoloxodon yorki type 1280
4074
4075 Palseoloiodon wilmani type 1280, 1281

4077 Loiodontn prima type 1287

4078 Loxodonki africana var. obliqria type 1287, 1288

4085 Archidiskodon milletti type 991

4086 Archidiskodon vanalphcni type 990
4087 Archidiskodon loiodonloides type 992
Palscoloxodcm kuhni type 1281
4144
4286 Loxodonta snbaniiqiia type 1288
4334 Archidiskodon proplaiiifrons type 986, 987
Palspoloxodon archidiskodontoides cotypes 1282

Kobe: Takikawa Middle School

Parastegodon infrequens Shikama, type' 1-120

Kyoto: Imperial University

Palspoloxodon namadicus naumanni genotype 1294
PaLroloxodon {Archidiskodon'!) tokiinagai mut. junior type 1300
Stegodon orienlalis shodoensis ref. (cast) 893
Mammontetis primigenius ref 907

La Plata: Museum
8-19 Cuvieronius platensis ref 1^48

Leiden: Dubois Collection

B 85 Palseoloxodon hysudrindicus cotype 1302, 1303

W 122 E 2 r Pala'oloxodon hysudrindicus cotype 1302

li 122 E 3 PaUeoloxodon hysudrindicus cotype 1302

Leiden: Natural History Museum

Elephas indicus sumatrnHus cotypes 1330, 1331, 1393

Leiden: Royal Museum of Geology and Mineralogy

Skgodon trigonocephalus type 890, 891

Leipzig: Pal.«ontological Collection of the University

4402 Archidiskodon imperator siluestris type 1015
Parelephns cohnnbi felicis type 1082

Leipzig: State Ethnographical .Museum (Grassi Museum)

Mammonleus primigenius ref 1130, 1228

Leningrad: Museum of the Institute of Mines, Academy of Sciences of the U.S.S.R.

"Elephas" affinis Eichwald 1393

Leningrad: Pal.eozoologic Institute of the Academy of Sciences

2280 MaJtlodon alacus Borissiak, type' 1418

'[Not determined hy the present author. — Exlitor.j

 LIST OF CATALOGUE NUMBERS 1613

Leningrad: Zoological Museum of the Academy of Sciences

cat. no. page
Marnmonteus primigenius ref 1130
Mammonteus primigenius ref 1130, 1148, 1387, 1603
Marnmonteus primigenius ref 1388
"Elephas" affinis Brandt 1388

Lincoln: Nebraska State Museum

1-4-15 Parelephas jeffersonii ref 1012, 1019, 1051, 1052, 1091, 1092, 1093, 1106, 1111
1-4-26 Archidiskodon imperator maibeni ref 1012, 1027
1-11-8-17E Archidiskodon imperator ref 1011, 1012
2-7-17B Archidiskodon imperator ref 1009, 1012
2}2-3-8-19 Archidiskodon imperator ref 1012
4-12-13 Archidiskodon imperator ref 1013, 1018
5-9-22 Archidiskodon imperator maibeni type 947, 1009, 1012, 1017, 1020, 1021, 1022, 1027, 1028, 1080, 1111
5-11-20 Archidiskodon imperator ref 1012, 1013
8-7-08 Archidiskodon imperator ref 1012
11-3-13 Archidiskodon imperator ref 1009, 1012
13-24-10-14 Archidiskodon imperator ref 1013, 1017, 1018
16-6-16 Archidiskodon imperator ref 1009, 1012
18-2-22 Archidiskodon imperator scotti type 1012, 1026, 1027
19-9-17 Archidiskodon imperator ref 1012
23-6-14 Archidiskodon hayi type 1006, 1012, 1023, 1024
29-25-11-18 Archidiskodon imperator ref 1012

Lisbon: Zoological Gardens

Loxodonta africana mo^ambica type 1193, 1409
Loxodonta africana angolensis type 1 193

London: British Museum (Natural History)

E 595 Archidiskodon planifrons ref 966
E 622 A rchidiskodoii planifrons ref 966
M 2004 Archidiskodon meridionalis ref 980
M 2006 Hesperoloxodon antiquus lectotype 1182, 1217, 1218, 1219
M 2009 ^Archidiskodon planifrons ref 963
M 2010 Archidiskodon planifrons cotype 950, 951
M 2498 Stegolophodon lydekkeri type (cast) 851, 1305
M 2705 Stegolophodon cautleyi lectotype 821, 841
M 2817 Stegolophodon cautleyi cotype 821, 840, 841, 842
M 2851 Stegolophodon cautleyi ref. (cast) 842
M 2857 Anancus perimensis ref 1548
M 2882 Anancns perimensis paratype 1392
M 2884 Stegolophodon cautleyi cotype 841
2887 Stegolophodon cautleyi cotype (east) 841, 842
M 2978 Stegodon bombifrons lectotype 830, 864, 865
M 2979 Stegodon bombifrons cotype 830, 858, 865, 891
M 3008 Stegodon ganesa ref 858, 871, 882
M 3015 Stegodon insignis lectotype 867, 881
M 3039 Stegodon insignis cotype 867
M 3060 Archidiskodon planifrons ref 938, 953, 960, 1352
M 3068 Archidiskodon planifrons lectotype 950, 951, 952, 953, 967
M 3070 Archidiskodon planifrons ref 953, 959, 960
M 3092 Palseoloxodon namadicus type 1211
M 3109 Hypselephas hysudricus ref 1341, 1350, 1351
M 3110 Hypselephas hysudricus ref 1352
M 3114 Hypselephas hysudricus ref 1354, 1355
1614 OSBORN: THE PROBOSCIDEA

London: British Museum (continued)

CAT. NO. PAGE
M 3127 Hyp.selephas hysudricus type 1341
M 3146 Hypselephas hysudricus paratype 1341, 1342
M 3428 Tetralophodon pimjabiensis cotype (cast) 841
7388 Stegodon elephanlaides ( = cliftii) cotype (cast) 863
7393 Stegodon elephanlmdes { = cUf(ii) lectotype (cast) 862
7436 Hesperoloxodon antiquus ref 989
M 8588 Pabfoloxodon Cypriotes cotype 1266
M 8591 Palitoloiodon Cypriotes cotype 1266
M 9378 Palxoloxodon creticus cotype 1267
M 9.381 Palxoloxodon creticus cotype 1267
M 10520 Stegodon elephantoides { = cliftii) cotype 825, 826, 831, 862, 863
M 10941 "Mammuthus" primigenius ref 1366
:M 12639 Loxodonla zulu ref 1287
.M 12641 Metarchidiskodon griqua ref 985, 994, 995, 1584
M 12642 Metarchidiskodon griqua ref 985, 994, 995, 1584
M 14102 "Mammuthus" primigenius ref 1366
14759 Stegodon elephantoides ( = cliftii) ref 863
16229 Hesperoloxodon antiquus ref 1220, 1234
17420 Mastodon americanus ref 844
1S489 Stegodon ganesa lectotype 871, 874
21680 lElephas antiquus ref 1223
27872 "Elephas" meridionalis ref 1226
27915 Parelephas (?) trogontherii nestii cotype 1059
29007 Palxoloxodon namadicus ref 1062, 1212
322.50 Parelephas armeniacus type 1060, 1061
32251 Parelephas armeniacus type 1060
32252 Parelephas armeniacus type 1060
32253 Parelephas armeniacus type 1060
32254 Parelephas armeniacus type 1060
32256 Parelephas armeniacus type 1060
33218 Parelephas columbi ref 1073, 1075
33327 Parelephas (?) trogontherii nestii cotype 10.59, 1060
33334 Archidiskodon meridionalis cromerensis ref 980
36695 Archidiskodon planifrons ref 959, 960
39370 Hesperoloxodon antiquus ref 1182, 1219
39463 Parelephas{l) trogontherii nestii cotype '
1059
.39464 Hesperoloxodon antiquus ref 1234
40769 Parelephas columbi type 1071, 1072, 1075
41925 Stegodon sinensis type 860
41926-7 Stegodon orientalis type 884, 885
44140 ?yl rchidiskodon planifrons ref 964
44304 Palaoloxodon mnaidriensis type 1264, 1265
44306 Palxoloxodon mnaidriensis paratype 1264, 1265
44312 Palxoloxodon melitensis type 1262
44895 "^Archidiskodon planifrons ref 963
14.2.16.1 Elephas indicus hirsutus type 1332
Hesperoloxodon antiquus ref 1223, 1224, 1227, 1249
Loxodonta africana ravendishi type 1 193
Loxodonta africana albertensis type 1193

London: Imperial Institute

Loxodonla africana selousi type 1 193

London: India House

Elephas indicus ref 1324
 \

LIST OF CATALOGUE NUMBERS 1615

London: Museum of the Geological Society

CAT. NO. PAGE
Stegodon elephantoides ( = cliftii) lectotype 862

London: Royal College of Surgeons

741a Archidiskodon imperator falconeri lectotype 1016
2656 Elephas indicus ceylanicus ref 1312

London: Zoological Society

Stegolophodon lydekkeri type 851

Los Angeles: Museum of History, Science and Art

3800-1
Archidiskodon imperator vei 939, 976, 977, 1007, 1009, 1053
3801-1/
Lyons : Museum of Natural Sciences
Parelephas intermedins ref 1063, 1064, 1395, 1603

Marseilles: Museum of Natural History

8449 Parelephas columbi cayennensis type 1083

Mexico: Engineering School

1 Archidiskodon imperator ref 1013
2 Parelephas columbi ref 1081
3 Parelephas columbi ref 1081

Mexico: Geological Institute

207 Archidiskodon imperator ref 1013
210 Archidiskodon imperator ref 1013
v-211 Archidiskodon imperator (?) ref 1013, 1014
212 Archidiskodon imperator ref 947, 1009, 1013, 1014
213 Parelephas columbi ref 1081
214 Parelephas columbi ref 1081
Parelephas columbi ref 1081

Moscow: University'
Parelephas vmsti cotypes 1065, 1066
"Ekphas hysudricus" ref 1065

Munich: State Zoological Museum

Elephas indicus sumatranus ref 1314, 1330, 1331

New York: American Museum of Natural History

44 (Dept. Mamm.) Elephas indicus ref 917, 918
1747 Parelephas columbi type (ca,st) 1000, 1071, 1072, 1074, 1075
2011 Palseoloxodon melitensis ref 1263
2568 Archidiskodon imperator type (ca.st) 998, 999, 1000, 1005
2573 Stegomaslodon aftoniie type (cast) '
1548
3283 (Dept. Manim.) Loxodonta africaaa oxyotis ref 931, 1095, 1131, 1176, 1193, 1194, 1199, 1200, 1227, 1228, 1249, 1250
3819 (Dept. Mamm.) Elephas indicus var. bengalensis ref 918, 1092
6835 Stegodon airdwana ref. (cast) 813, 887
8681 (Cope Coll.) Parelephas washingtonii ref 939, 1051, 1053, 1084, 1088, 1090, 1091, 1100, 1101, 1102, 1103, 1111, 1144
8681 A Parelephas washingtonii type 1006, 1100, 1101, 1102, 1103
9905 Trilophodon palmndicus type (cast) 1546
9950 Parelephas jeffersonii genotype 931, 939, 1006, 1041, 1051, 1053, 1084, 1085, 1086, 1087, 1088,
1089, 1090, 1091, 1094, 1095, 1103, 1108, 1109, 1110, 1111, 1112, 1113, 1114, 1131, 1144, 1227, 1228
10374 Serridentinus lydekkeri type (cast) 885
10378 Stegodon bombifrons cotype (cast) 830, 858, 865, 891
10381 (Warren Coll.) Palxoloxodon namadicus type (cast) 1211
10382 (Warren Coll.) Stegodon elephantoides ( = cliftii) cotype (cast) 826, 831, 862
10457 (Warren Coll.) Parelephas progressus type 1084, 1085, 1097, 1098, 1099
10582 Serridentinus productus ref 1546
1616 OSBORN: THE PROBOSCIDEA

New York: Ameuic.'VN Museum (continued)

CAT. NO. PAGE
10598 ArchicUskodon imperator ref 990, 1005, 1006, 1007, 1010, 1017, 1018, 1022, 1030, 1080
10655 He.speroloxodon antiquus germanicus ref 1253, 1255
10656 Mammonteiig priinigeniiis ref 1143
11871 Archkliskodon imperaior neotype 999, 1000, 1001, 1005, 1549
13225 Parelephas jeffersonii ideotype 1006, 1070, 1084, 1085, 1087, 1088, 1089, 1096
13431 Maritherium trigodon ref 1544
13437 Motriiherium andrewsi ref 1544
13449 Paheomastodon intermedius paratype 1544
13707 (Cohen Coll.) Parelephas columbi neotype 1013, 1072, 1073, 1074, 1075, 1076
13708a-z (Cohen Coll.) Parelephas colionbi ref 1075, 1076, 1105
13708aa-kk(Cohen Coll.) Parelephas columhi ref 1076
13709 (Cohen Coll.) Parelephas columbi ref 1075, 1076
13749 Mammonteus primigenius rompressus paratype 1127, 1135, 1142, 1143, 1157, 1159
13752 Mammaideus primigenius ref 1 142, 1 143
14294 Mastodon americanus ref 1546
14343 Mammonteus primigenius ref 1 143
14371 Mammonteus primigenius ref 1 142, 1 143
14475 (Cope Coll.) Archidiskodon imperator ref 922, 924, 1004, 1005, 1039, 1084
14476 Archidiskodon imperator ref 938, 989, 990, 1001, 1005, 1007, 1008, 1009, 1017, 1028
14547 Palsponiaslodon intermedius type 1 544
14558 Archidiskodon imperator ref 989, 990, 1001, 1002, 1005, 1006, 1007, 1089
14559 Mommonteiis primigenius compressus type 922, 931, 1095, 1131, 1142, 1145, 1157, 1158
14610 Archidiskodon imperaior scotti type (cast) 1026, 1027
15878 Hypselephas hysudricus ref 1343, 1348
15898 Mferitherium hjonsi ref 1544
17355 Archidiskodon imperator ref 1011, 1012
18536 Stegodon orientalis grangeri ref 808
18629 Stegodon orientalis grangeri ref 880
18630 Stegodon orientalis grangeri ref 880, 891
18630 a Stegodon orientalis grangeri ref 876, 878
18632 Stegodon orientalis grangeri ref 891
18636 Stegodon orientalis grangeri ref 876, 878, 880
18638 Stegodon orientalis grangeri ref 880
18640 Stegodon orientalis grangeri ref 880
18642 Stegodon orientalis grangeri ref 876, 879
18702 Stegodon orientalis grangeri ref 880
18705 Stegodon orientalis grangeri ref 813, 876, 878, 880
18708 Stegodon orientalis grangeri ref 880
18711 Stegodon orientalis grangeri ref 876, 878, 880
18714 Stegodon orientalis grangeri type 875, 876, 877, 879, 880, 881, 1547
19416 (Brown Coll.) Hypselephas hysudricus ref 1346, 1347, 1348
19421 Trilophodon chinjiensis holotype 1404
19446 Stegolophodon cautkyi progressus type 835, 848, 849, 850, PI. XIII
19455 Stegolophodon nalhotensis type 847, 848, PI. XIII
19772 Stegodon pinjorensis type 835, 881, 882, 883, 891, 938, 960, 1360
19773 Stegodon ganesa ref 869, 882
19778 Archidiskodon planifrons ref 954, 955, 957
19783 (Brown Coll.) Hypselephas hysudricus ref 1347, 1348
19786 Hypselephas hysudricus ref 1346, 1347, 1348
19798 Archidiskodon planifrons ref 954, 955, 967
19799 (Brown Coll.) Hypselephas hysudricus ref 1346, 1347, 1348
19804 Stegodon insignis rtsf 869
19809 (Brown Coll.) Hypselephas hysudricus ref 1346, 1347, 1348
19809A (Brown Coll.)
19818 (Brown Coll.)
Hypselephas hysudricus ref
Plalelephiis plnlycephalus type
1347, 1348, 1351
1347, 1348, 1352, 1359, 1360, 1361
I
 LIST OF CATALOGUE NUMBERS 1617

New York: American Museum (continued)

CAT. NO. PAGE
19819 Archidiskodon planijrons ref 954, 955, 958, 1549
19821 ArchidiskodoH planifrons ref 954, 955, 956, 967, 1549
19830 (Brown Coll.) Hypsekphas hysudricus ref 1345, 1347, 1348
19858 Stegodon insignis ref 869, 870
19859 Stegodon insignis ref 869, 871
19861 Archidiskodon planifrons ref 954, 955, 958
19862 Archidiskodon planifrons ref 954, 955, 957
1986,3 (Brown Coll.) Hypsekphas hysudricus ref 1344, 1346, 1347, 1348, 1353
19864 Archidiskodon planifrmis ref 954, 955, 957, 963
19866 (Brown Coll.) Hypselephas hysudricus ref 1347, 1348, 1349, 1354, 1355, 1356, 1357
19867 (Brown Coll.) Hypselephas husudricus ref 1342, 1346, 1347, 1348
19869 Stegodon insignis ref 868, 869, 877
19869 a (Brown Coll.) Hypselephas hysudricus ref 1342, 1344, 1347, 1348
19870 Archidiskodo7i planifrons ref 954, 955
19871 Archidiskodon planifrons ref 954, 955
19873 Archidiskodon planifrons ref 954, 955, 957
19879 Archidiskodon planifrons ref 954, 955, 959
19880 Archidiskodon planifrons ref 954, 955
19881 Archidiskodon planifrons ref 954, 955, 956
19882 Arckidiskodon planifrons ref 954, 955
19915 (Brown Coll.) Hypselephas hysudricus ref 1347, 1348, 1354, 1357
19916 Archidiskodon planifrons ref 954, 955
19917 Archidiskodon plamfrons ref 954, 955
19950 Archidiskodon planifrons ref 954, 955, 956
19951 Archidiskodon planifrons ref 954, 955, 958, 959, 967
19952 Archidiskodon. planifrons ref 954, 955, 956
19955 Archidiskodon planifrons ref 954, 955, 956
19956 (Brown Coll.) Hypselephas hysudricus ref 1347, 1348
19961 Archidiskodon planifrons ref 954, 955
19962 (Brown Coll.) Hypselephas hysudricus ref 1346, 1347, 1348
19964 Stegodon insignis ref 869, 870
19965 Archidiskodon planifrons ref 954, 955, 957, 986, 987, 988, 989
19967 Archidiskodon planifrons ref 955
19968 Archidiskodon planifrons ref 954, 955, 958
20002 Stegodon insignis birmanicus type 874, 875, 877
20044 Stegodon bombifrons ref 866
20069 Archidiskodon imperaior ref. (cast) 1012
21115 (Frick Coll.) Trilophodon pojoaguensis type 1411
21125 (Frick Coll.) Ocalientinus ojocaliensis ref 1411
21872 Stegodon zdanskyi type (cast) 899
21878 Stegodon officinalis type (cast) 898
21879 Stegodon officinalis ref. (cast) 898
21889 Loxodonta africana peeli ref 1198, 1248, 1249
21891 Archidiskodon meridionalis ref. (cast) 1034, 1037
21892 Parelephas jeffersonii ideotype 1084, 1087, 1088, 1089
21894 Archidiskodon meridionalis ref. (cast) 1034, 1037
21895 Archidiskodon meridionalis ref. (cast) 1034, 1037
21907 Archidiskodon broomi type (cast) 989
21924 Archidiskodon subplanifrons type (cast) 987, 988
21933 Parelephas cohtmbi cayennensis type (cast) 1083
21978 Stegolophodon latidens cotype (cast) 826, 843, 844, PI. XIII
22481 Archidiskodon imperator ref 947, 1007, 1008
22616 Stegolophodon latidens ref. (cast) 845
22634 Hesperoloxodon anliquus italicus type 1210, 1216, 1234, 1238, 1241, 1242, 1243, 1244, 1245, 1246,
1247, 1248, 1249, 1251, 1252, 1255
1618 OSBORN: THE PllOBOSCIDEA

New York: American Museum (continued)

CAT. NO. PAGE
2263G Stegodon airdwana ref 881
22637 Archidiskodon sonoriensis type 1033
22722 Archidiskodon milletti type (cast) 991
22723 Archidiskodon vaiialpheni type (cast) 990
22724 Archidiskodon loiodontoides type (cast) 992
22725 Palxoloxodon kuhni type (cast) 1281
22726 Palxoloxodon icilmani type (cast) 1280, 1281
22727 Palseoloxodon yorki type (cast) 1280
25500 (Frick Coll.) Archidiskodon imperator ref 1012
25500 A (Frick Coll.) Archidiskodon imperator ref 1012
25501 (Frick Coll.) Archidiskodon imperator ref 1012
25501 A (Frick Coll.) Archidiskodon imperator ref 1012
25501 B (Frick Coll.) A rchidiskodon imperator ref 1012
25505 A-D (Frick Coll.) Archidiskodon imperator ref 1012
25506 (Frick Coll.) Archidiskodon imperator ref 1012
26820 Parelephas floridanus type 1051, 1105, 1106, 1107, 1108, 1109, 1110, 1111, 1113, 1114
26821 Parelephasfloridanus paratype 1105, 1106, 1107, 1108, 1109, 1110, 1111, 1113, 1114
26822 Parelephas floridanus paratype 1107, 1108, 1109
26823 Parelephas floridanus ref 1107, 1109
26824 Parelephas floridanus ref 1 107, 1109
26825 Parelephas floridanus ref 1107, 1109
26826 Parelephas floridanus ref 1 107, 1109
26833 a-f J^arelephas floridanus ref 1 107, 1 1 14
26965 Stegolophodon cautleyi cotype (cast) 821, 840
26966 Stegolophodon cautleyi lectotype (cast) 821, 841
26968 Palseoloxodon andrewsi type (cast) 993, 1278
26969 Archidiskodon proplanifrons type (cast) 986, 987
26980 Mammonteus primigenins lectotype (cast) 1122, 1123
26981 Mammontens primigenius lectotype (cast) 1122, 1123
26987 Loxodonta prima type (cast) 1287
26988 Loxodonta africana var. obliqua type (cast) 1287, 1288
26989 (Frick Coll.) Mammonteus primigenius alaskensis cotype 1159, 1160, 1161
26990 (Frick Coll.) Mammonteus primigenius alaskensis cotype 1159, 1160, 1161
26991 (Frick Coll.) Mammonteus primigenius alaskensis cotype 1159, 1160, 1161
27010 (Frick Coll.) Mammonteus primigenius alaskensis ref 1159
27769 Palseoloxodon transvaalensis type (cast) 1284
27796 Cuvi(ronius superbus ref. (cast) 1546
32727 (Dept. JMamm.) Loxodonta africana peeli ref 1190
32732 (Dopt. Mamni. Loxodonta africana peeli ref 1190
32734 (l)opt. Mamiii. Loxodonta africana albertensis ref 1190
35591 (Dopt. Mamm.) Loxodonta africana pumilio type 1193, 1259
39081 (Dei)t. Mamm. Elephas indicus ref 1337, 1605
39082 (Dopt. Mamm. Elephas indicus ref 1227, 1228, 1337
39083 (Dept. Mamm.) Loxodonta africana ref 1006
51939 (Dept. Mamm. Loxodonta africana ref 1251, 1252, 1335
54085 (Dept. Mamm. Loxodonta africana albertensis ref 1190, 1202, 1239
54261 (Dept. Mamm.) Elephas indicus ref 1006, 1251, 1252, 1335, 1360
54452 (Dept. Mamm.) Elephas indicus ref 1311
54453 (Dept. Mamm.) Elephas indicus ref 1249, 131 1, 1337

New York: Zoological Society

Loxodonta africana oxyotis ref 1201, 1239

Norwich: Norwich Castle Museum and Art (Jallery

1570 Archidiskodon mcridionalis ref 973
1703 Hespei'oloxodon antiquus ref 1397
 LIST OF CATALOGUE NUMBERS 1619

Oran: Museum
CAT. NO. PAGE
Palseolojrodon allaiiticiis cotype 1274
Pabeoloxodon allanticus ref 1260

Oxford : Museum
Archidiskodon meridionalis ref 973

Palermo: Museum
Coll. I Palxoloxodon mnaidriensis ref 1260
Coll. VI Palxoloxodon mnaidriensis ref 1260

Paris: Museum of Natural History

AC 20.58 Serridenlinus filholi type 1417
AC 2062 Serridentinus filholi type 1417
A 8014 Elephas indicus reylanicus ref 1317
A 8016 Elephas indicus hengalensis type 1312, 1313, 1316, 1317
Archidiskodon meridionalis ref 977, 978, 979, 1034, 1093, 1094, 1095
Turicius tapiroides ref 819
Anaiwus hrevirostris ref 1548

Pasadena: California Institute of Technology

14 (Coll. Vert. Pal.) Archidiskodon exilis type 1030
1922 (Coll. Vert. Pal.) Pliomastodon nevadanus Stock, type' 1418

Philadelphia: Academy of Natural Sciences

Mastodon americanus rugosidens type 1399
Archidiskodon imperator ref 1002

Philadelphia: American Philosophical Society

Mastodon americanus ref 1388

Philadelphia: Wagner Free Institute of Science

Parelephas columbi ref 1076

Private Collection: Walter W. Holmes, Florida

Parelephas columbi ref 1076

Private Collection: Mr. Kato, Kokubo, Japan

Pabeoloxodon protomammonteus proximus type 1298

Private Collection: Percy Madeira, Philadelphia

Loxodonia africana peeli ref 1202

Private Collection: Mr. Natsume, Minato, Japan

Palxoloxodon protomammonteus type 1297

Private Collection: Due d'Orleans, Wood Norton, England

Loxodonta africana orleansi type 1193

Private Collection: C. V. A. Peel, Oxford, England

Loxodonta africana peeli type 1193

Private Collection: Mr. Powell-Cotton, Birchington, England

Loxodonta africana cottoni type 1193

Private Collection: Lord Rothschild, Tring, England-

Loxodonta africana peeli cotype [paratype] 1193

'[Not determined by the present author. —Editor.]

^[Probably at the British Museum. — Editor.)
1620 OSBORN: THE PROBOSCIDEA

Quito: Museum of the Central University of Ecuador

cat. no. page
Cuvieronius poslremus ref 1601

Rome: University Museum

Hesperoloiodon anliquus italicus ref 1236, 1247

Rotterdam: Zoological Gardens

Ekphas indicus sumatranus ref 1329, 1331

Saint John, New Brunswick: Museum

tPalxoloxodon hysudrindicus ref 1334

Sendai: Tohoku Imperial University

IPalxoloxodon buski type 1333
Palxoloxodon namadicus yabei ref 1299

Sendai (probably) : Second High School

PaUeoloxodon namadicus yabei type 1299

Stockholm : Museum of King Adolph Frederick

Ekphas indicus type 1309, 1310

Stuttgart: Wurttemburg Natural History Collection

12837 Mnmvwnteus primigenius fraasi type 1130, 1152, 11.53, 1604
15344 ] I esperoloxodon antiquus germanicus ref 1254, 1255, 1256
1.5930 H esperoloxodon antiquus germanicus ref 1254, 1255, 1256
16274 fI esperoloxodon antiquus germanicus ref 1236
16515 H esperoloxodon antiquus germanicus ref 1255

Tallahassee: Florida Geological Survey

\-4529 Parekphas floridanus ref 1115

Tervueren: Museum of the Belgian Congo

Loxodonla africana fransseni type 1193

Tokyo: Geological Institute of the Imperial University

iilcgodon aurorse type 893
Stegodon orientalis shodoensis ref 893

Tokyo: Museum of Education at Ochanomizu

Palasoloxodon tokunagai type 1298

ToYKo: Uyeno Museum

2194 Stegodon orientalis shodoensis type 893
2208 (See Museum of Education, Ochanomizu, Tokyo)

Turin: Museum
15 Mrimmonteus primigenius astensis paratype 1154, 1407
Mammonteus primigenius astensis type 1154, 1407
A nancus arvernensis ref 1601
I'arekphas trogonthcrioides lectof ype 1054
Parekphas trogontherioidcs cotypc 1054

Vienna: Natural History Museum

Sicgolophodon sublatidens type 846
 LIST OF CATALOGUE NUMBERS 1621

Washington: United States National Museum

CAT. NO. PAGE
185 Archkliskodon imperator type 998, 999, 1001, 1002, 1005
287 Parelephas jeffersonii or P. washingtonii ref 1088
1614 Parelephas columhi ref 1077
2195 Parelephas jeffersonii ref 1086, 1087, 1088, 1089, 1095, 1096, 1097
2216 Archidiskodon imperator ref 1002
2217 Archidiskodon imperator ref 1002
4162 Marnmonteus primigenius ref 1145
4761 Parelephas jeffersonii ref 1097
6052 Parekphas jeffersonii or P. icashingtonii ref 1088
6662 Archidiskodon imperator ref 100.3
6663 Archidiskodon imperator ref 1003
6666 Marnmonteus primigenius ref 1 145, 1146
6668 Parelephas jeffersonii or P. umshingtonii ref 1088
6669 Parelephas jeffersonii or P. washingtonii ref 1088
8579 Mammovfeus primigenius ref 1091, 1145, 1146
8580 Marnmonteus primigenius ref 922, 939, 1051, 1091, 1143, 1144
10261 Parelephas jeffersonii ref 922, 1052, 1084, 1087, 1088, 1090
11620 Archidiskodon imperator ref 996, 1005, 1078
11805 Archidiskodon imperator ref 996, 1005
11806 Parelephas floridanus ref 996, 1077, 1078, 1079, 1108
11808 Parelephas floridanus ref 996, 1077, 1078, 1079, 1108
11810 Parelephas floridanus ref 996, 1077, 1078, 1079, 1 108
11814 Archidiskodon imperator ref 996, 1005

Washington: Zoological Park

Elephas indicus sumalranus ref 1314

Weimar: Natural History Museum

10 Parelephas trogontherii ref 1056
17 Parelephas trogontherii ref 1056
32 Parelephas trogontherii ref 1056
46 Parelephas trogontherii ref 1056
Hesperoloxodon antiquus germanicus neotype 1234, 1235

Weimar: Reeling Collection

69 Parelephas trogontherii ref 1056
80 Parelephas trogontherii ref 1056
87 Parelephas trogontherii ref 1056
89 Parelephas trogontherii ref 1056

Weimar: Schwabe Collection

Parelephas trogontherii type 1057
 SUPPLEMENTARY BIBLIOGRAPHY
See this Memoir, Volume I, page 761

Adams, Michael Arambourg, Camille

1807.1 Relation d'un Voyage a la Mek Glaciale et Decouverte 1933.1 Decouverte d'un gisement de Mammiferls burdiqaliens
DES RESTE8 d'un Mammouth. Jimm. du Noid, St. Petersburg, No. d.\ns le Bassin du Lac Rodolphe (Afrique Orientale). Compt.
XXXIII, Aoflt. Rend. Soc. geol. France, No. 14, November 20, pp. 221, 222.
Origina! not seen by the present author. Translated by Sir Joseph Banks (see Mastodon af. anijustidens.
entry under Michael Adams, this Memoir, Vol. I, p. 762).
1934.1 Le Dinothehium des gisements de l'Omo. Compt. Rend.
Soc. g-^ol. France, No. 6, March 19, pp. 86, 87.
Agassiz, Jean Louis Rodolphe
New species: D. Bozasi, p. 87. See .\ranibourg, lOS.^J.!, for type figure.
1850.1 On the Fossil Remains of an Eleph.\nt fodnd in Vermont.
Proc. Amer. Assoc. Adv. Sci., 2d Mpeting, pp. 100, 101. 1936.1 Le Dinothbrium des gisements de l'Omo (Abyssinie). Bull.
Tliis reference appears in the Bibliografthy, \'(il, I, p. 7fi2, but without specific Soc. g^ol. France, (5), IV, February, pp. 305-310, PI. xviii.
determination by tlie present author, uf the fossil elephant remains found
on the "slope of Mt. Holly, Vermont." Subsequent research has confirmed Type figure of Di'ivthfrinin Boznsi, PI. xvill.
the identification of Warren (185.5, PI. xxviii) as Ehpfias [Mammonteuxl
jiritriiof^'nus.
Astre, Gaston
1937.1 SuH UN Elephas .\ntiuuus a Formule Laminaire Elevee.
Alekseev [Alexejew], A.
Bull. Soc. Hist. Nat. Toulou.se, LXXI, Fasc. 1-2, .luiie 30, pp. 26-32,
1930.1 Die obersarmatische Sabgetierfauna von Eldar. I. PI. I.

Achtiaria Borissiakii n. sp. Trav. Musee Geol., Acad. Sci. New nmtation: Elephas aritifjuu,-^ ruthenensis.

U. R. S. S., VII, pp. 167-204, Pis. i-v, text figs. 1-6.

Barbour, Erwin Hinckley, and Harold James Cook

Allen, Glover Morrill

1917.1 Notes on the Skull of Metoreodon. Neb. Geol. Suiv., VII,
Pt. 18, April 15, pp. 165-172, text figs. 1-8.
1936.1 Zoological Results of the George Vanderbilt African
Expedition of 1934. Part II, -The Forest Elephant of Africa. 1917.2 Skull of Aeldrodon platyrhinus, sp. nov. Neb. Geol. Surv.,
Proc. Acad. Nat. Sci. Phila., LXXXVIU, pp. 1.5-44, Pis. i-iv, text VII, Pt. 19, April 15, pp. 173-180, te.xt figs. 1-11.
figs. 1, 2, folding map. The name "Valentine" given to a series of beds in northern Nebraska.
Forest and bush elephants, especially the smaller forest elephant, the so-called
"pygmy elephant," of the Cameroons. Barbour, Erwin Hinckley, and Charles Bertrand Schultz
1937.1 An Early Pleistocene Fauna from Nebr.aska. Amer. Mus.
Ameghino, Florentine Novitates, No. 942, September 10, pp. 1-10, text figs. 1-4.
1880-1881 La Antiquedad del Hombre en el Plata. 8vo, Paris and Hay .Springs fauna, pp. 3. 4, 6.

Buenos Aires: I, 1880, pp. i-xiv, 1-640, Pis. i-xvi; II, 1881, pp.
1-557, Pis. xvii-xxv, folding table. Beliaeva, £.
Separation of the Pampean into three successive horizons: Terreno pampeano 1936.1 EiN Fund von Elephas in T.\d.schikistan. Trudy Paleo-
lacustre, Terreno pampeano superior, and Terreno pampeano antique.
zool. Inst., Akad. Nauk SSSR, V, pp. 103-109, 1 pi., 2 text figs.,

1 map.
Andersson, J. Gunnar
1923.1 Essays on the Cenozoic of Northern China. Mem. Geol. Benedict, Francis G.
Surv. China, (A), No. 3, March, pp. 1-152 (also 16 pp. in Chinese), 1936.1 The Physiology of the Eleph.ant. Publ. Carnegie In.stn.
Pis. i-ix, text figs. 1-42, maps i-iii, 2 folding tables. Wash., No. 474, pp. vii-f-302. Pis. i-viil, text figs. 1-13, tables 1-43.
Describes the Lu Tzu Kou beds of .Shansi, discovered bj' Zdansky.

Berkey, Charles P., and Frederick K. Morris

Andrews, Charles William
1927.1 Geology of Mongolia. N.atural History of Central Asia,
1914.1 On the Lower Miocene Vertebrates from British East II. 4to, American Museum of Natural History, New York, pp.
Africa, collected by Dr. Felix Oswald. Quart. .lourn. Geol. xxxi-l-475, 44 pis., 161 text figs.
Soc. London, LXX, pp. 163-186, Pis. xxvii-xxix, text figs. 1-3.

Black, Davidson, Pierre Teilhard de Chardin, Chung-Chien Young,

Andrussow [Andrusov], Nikolai Ivanovich and W. C. Pei
1905.1 Maeotische Stufe. Verh. russisch.-kaiserl. Min. Ges. St. 1933.1 Fossil M.\n in China. Mem. Geol. Surv. China, (k), No. 11,
Petersb., (2), XLIII, pp. 289-449, Taf. v, vi. May, pp. i-x, 1-166 (also 5 pp. in Chinese), text figs. 1-82, maps i-vi.

Arabu, N. von Blainville, Henri Marie Ducrotay de

1916.1 Etudes sur les formations terti.ures du bassin de la mer 1817.1 Dents. Nouv. Diet. d'Hist. Nat., nouvelle edition, IX, pp.
DE M.\RMAR,\: classification ET PARALLELISML DES DERNIERES 252-352, synoptic table. Paris.
The woril MnKf'ntoiitum u.sed for "Genre Mastodonte" on page 27fi.
couches NEOGENES de LA REGION ET DES REGIONS VOISINES. Compt.
Rend. Acad. Sci., CLXII, pp. 332-334. Paris.
Blumenbach, Johann Friedrich
1916.2 Existence de la faune a Hippahion dans le Sarmatien du 1803.3 Das Mammut-Ohioticum, nun wirklich in London. Maga-
BASSIN DE LA MER DE MaRM-\RA ET SES CONSEQUENCES POUR LA zin den neuesten Zustand der Natuikunde, herausgegeben
f. .1. H.
CLASSIFICATION DU NeOGBNE DANS l'EuROPE SUD-ORIENTALE. Voigt, V, .lanuary, pp. 1-6. Weimar.
Compt. Rend. Acad. Sci., CLXII, pp. 424-426. Paris.
1913.1 Ueber die FOS.SILEN Gebeine von Elephanten und Mam-
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DU BASSIN DE LA Mer DE Marmara. Bull. Soc. geol. France, (4), Pfl.\nzen-Reste. besonders aus den Hannoverschen L.\NDEN.
XVII, for the year 1917, pp. 390-405, text fig. 1. Annalen der Physik, XLV, pp. 425-436. Leipzig.

1623
1624 OSBORN: THE PROBOSCIDEA

Borissiak, Alexei Alexievich Clark, J. Desmond, and H. B. S. Cooke

1914.1 Mammikeres FossiLEs DE Sebastopol. I. Mom. Comitc 1939.1 See Cooke, H. B. S., and Desmond
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Potersburg.
Colbert, Edwin Harris
1916.1 Mammikere.s Fo.ssilkw de Seb.\stofol. II. Mom. Comiti': 1937.1 The Pleistocene Mammals of North America and Their
Gool., (N. S.), I.ivr. 187. pp. 1 47, Pis. i-iii, toxt figs. 1-3. St. Rel.ations to Eurasian Forms. In "Early Man as Dei)ictpd by

Ppfersbuig. Leading Authorities at the International Symposium The Academy

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1936.1 MA.'iTODON .AT.WUS N. SP., DER PRIMITIVSTE VeRTRETEB DER cott Company, Philadelphia, pp. 173-184.

Gruppe M. angustidens. Trudy Paleozool. Inst., Akad. Nauk

SSSR, V, pp. 171 234, S pis., 16 text figs. 1938.1 Fo.ssil Mammals from Burma in the American Museum of
Natural History. Bull. Amer. Mus. Nat. Hist., LXXIV, Art. VI,
October 14, pp. 2.55-436, text figs. 1-64.
Bose, B. K.
1929.1 Notice of Mammalian Remains krom the Siwaliks of Cook, Harold James, and Erwin Hinckley Barbour
.Iammf. Quart. ,Iourn. Geol., Mining and Mctallurg. Soc. India, II,
1917.1 and 1917.2 See Barbour, Erwin Hinckley, and Harold James
No. 3, September, pp. 121-125, PI. v.
Cook.
Type of Falseoloxodon pritcmi vat. hosei Cliakraviirti, 193.'), figured (PI. v. fig.
10) without name.

Cook, Harold James, and Margaret C. Cook

Boule, Marcellin, and Jean Piveteau 1933.1 Faunal Lists of the Tertiary Vertebr.\ta of Nebraska and
1936.1 l.ES FossiLEs. Klementm de Paleontoloqie. 8vo, Masson Adjacent Are.as. Neb. Geol. Surv., Paper No. 5, 58 pp.
et Cic, Paris, pp. vii+899, frontispiece and 1330 text figs.

Succession of Pleistocene faunas in Europe.

Cook, Margaret C, and Harold James Cook
1933.1 See Cook, Harold James, and Margaret C. Cook.
Brayley, Kdward William
1833.1 [FossiLE Wirbelthier-Reste iM Arktischen Kreise.I Neues
Jahrb. Min., pp. 370-372. (Unsigned review.)
Cooke, H. B. S., and J. Desmond Clark
Elejitifi" primurtiiittis, p. 371?. 1939.1 New Fossil Elephant Remains from the Victoria Falls,
Northern Rhodesia, and a Preliminary Note on the Geology
and Archaeology of the Deposit. Trans. Roy. Soc. So. Afr.,
Caruana, A. A. XXVII, Pt. 3, December, pp. 287-319, Pis. xii, xiii, text figs. 1-1 1,

1870.1 FlHTHER DISCOVERY OF THE FoSSIL ELEPHANTS OF MaLTA. 1 table.

(^uart. .lonrn. Geol. .Sor. London, XXVI, pp. 434-436. New spei'iep: Palafohxodcn (tarti, pp. 29t)-302.

Cummins, W. F.
Castellanos, Alfredo
1893.1 Notes on the Geology of Northwest Texas. 4th Ann.
1936.1 Los sedimentos del Pampe.ano inferior y del Arauoano en
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el Valle de Los Reartks (Sierra de C6rdoba). Publ. Fac.
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Cien. Mat., Fis.-Quim. y Nat., Univ. Nac. Litoral, (Ser. Tecnico-
Cientifica), No. 6, pp. 1-1 10, text figs. 1-22. Rosario, Argentina.
Darton, N. H.
1906.1 Preliminary Report on the Geology and Underground
Chakravarti, D. K. Water Resources of the Central Great Plains. U. S. Geol.
1936.1 On the occurrence of a new Loxodo.ntine form of Eleph.^nt Surv. Profess. Paper No. 32, 433 pp.. Pis. I-Lxxil, text figs. 1-18.
Tfie name "Ogallala" used to designate the tapper Tertiary of Nebraska above
IN the Indian Cainozoic: Pal.eoloxodon priscus var. bosei. the ".\rikaree."
Proc. 22nd Indian Sci. Congress, Calcutta, (Fourth Circuit), p. 209.

1936.2 On the generic reference of a doubtful Rhynchorostrine Dehm, Richard

Bunomastodontid from Chinji in the Salt R.\nge. Proc. 22nd 1937.1 EiN Waldelefanten-Vorkommen in zwischeneiszeitlichen
Indian Sci. Congress, Calcutta, (Fourth Circuit), p. 209. .\blagerungen am Alpenrand. Bayerische Vorge.schichtsbl.'itter
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1937.1 K NEW Stage in the Evolution of Steoodons. Steoodon
ELEPHANToiDES (Clift). Quart. .Journ. Geol., Mining and Metal-
hirg. Soc. India, !.\, No. 2, June, pp. 33-37. Delafosse, Wilfrid
1936.1 Deuxi^me Contribution X l'etude de la Faune guArERNAiRE
1937.2 On a Primitive Loxodontine Form of Elephant from the DU dep.\btment de la Moselle.- II les Elephants. Bull. Soc.
Siwaliks of Jammu. Quart. Journ. Geol., Mining and Metalling. d'Hist. Nat. Moselle, 34"= Cahier, pp. 167-212, Pis. i-xii, 7 text fig.s.

.Soc. India, IX, No. 2, June, pp. 39-42, PI. vi. Hemains of E. itierulintinlis. E. njitititius, E. ifrimit/cniiin have been found in
Moselle.
Supplementary (leHerlption of I'nlxuloiodon />rinr.un var. bonri.

Denny, Charles S.
Chang, H. T.
1940.1 Santa Fe Formation in the EspaSola V.\lley, New Mexico.
1926.1 On the Question of the Existence of Elephants and Bull. Geol. Soc. Amer., LI, No. 5, May 1, pp. 677-693, Pis. i-iv, text
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Mnslotherium ohiolicum, p. 252. LI, pp. 111-132, Taf. 1-4.
1626 OSBORN: THE PROBOSCIDEA

Hopwood, Arthur Tindell 1914.1 La faune meotique du village Taraklia on dlstrict de

1929.1 A Rkvikw of the Fossil Mammals of Clntral Africa. Amcr. Bendery. Fissipedia, Rodentia, Rhinocerinae, Equinab, Suidae,
.lourn. Sci., (o), XVII, pp. 101-118. Proboscidea. Trudy Besstirahskoe ohshcliestvo estcstvoispytatelei,
V, pp. 1-55, Pis. i-iv. Kisliinef.
1932.1 The Oluuvai Expf.uitiox, 1931. Nut. ?Iist. Mag., Ill, pp.
214-22,j, text 1-11.
fig.-i.
Kirk, John
1933.1 Miocene Primates from Kenya. .louin. Linn. See. London, 1864.1 List of M.^mmalia met with in Zambesia, East Tropical
(Zoology), XXXVIII, No. 260, Novomhoi- 7, pp. 437-464, PI. vi. Africa. Proc. Zool. Soc. London, pp. 649-660.

1935.2 Fossil Elephants and M.\n. Proc. Geol. Assoc, XLVI, Pt.
Koenigswald, G. H. Ralph von
I . March 28, pp. 46-60. London.
Table summarizing "the of the more important of the larger
cli.stribution 1931.1 Die Bedeutung der Equiden fOr die Altersstellung des
flpeeies of maminuU found associated with human remains in Europe" kheinhessischen Dinotheriensandes. Centialb. Min. Geol. Pal.,
(p. 60K Of the elephants. E. jtlanifruns. E. aatiquus, and E. primuj^nius
are mentioned. Abt. B, pp. 42-48, 1 text fig. Stuttgart.

1939.1 Das Pleistocan Javas. Qu:irtar (Berlin), II, pp. 2S-53, Taf.
1937.1 The Identity of Elephas TROaoNTiiKRii Pohlio. Bull. Gool.
ix-xi. Abb. 1-6.
In.'^t. Univ. Up.sala, XXVII, pp. 19-24.
Mounted skeleton of SItyiMhn tnui>fioct]tfniliifi in the Geological Museum of
Bandoeng.
1937.2 Wiss. Ergebn. Oldoway-
Die FossiLEN Pferde VON Oldoway.
Expeditioii, 1913, horausgcgcbcn von H. Rt-ck, Hoft 4, pp. 111-136, 1939.2 The Rel.\tionship between the Fossil M.\mmalian Faunae
PI. VII. OP Java and China, with Special Reference to Early Man.
Peking Nat. Hist. Bull., XIII, Pt. 4, June, pp. 293-298, 1 text fig.
1940.1 Fossil Mamm.\ls and Pleistocene Correlation. Proc. Geol.
.\ssoc., LI, Pt. 1, March 29, pp. 79-88. London.
Kraglievich, Lucas
1934.1 La antiquedad pliocena de l.^s faunas de Monte Heemoso
Hopwood, Arthur Tindell, and J. Reid Moir
Y Chapadmalal, deducidas de su comparaci6n con l.\s que le
1939.1 Sec Moir, ,J. Reid, uiul Aitliur Timlcll Hopwood,
precedieron y suCEDiERON. 8vo, "El Siglo Ilustrado," Montevideo,
136 pp., fontispiece and 1 text fig.
Hopwood, Arthur Tindell, and Guy Ellcock Pilgrim Designated the Lower Pampean, or Ensenadense. as "llpper Pliocene or
Pliopleistocene."
1939.1 .~^i'c Pilfjrim, Guy EUouck, ;tml .'\rtliur Tindell Hopwood.
Krejci-Graf, Karl
Hutchinson, G. E., and Hellmut de Terra 1932.1 Parallellsierung des strDOSTEUROpXiscHEN Pliozans. Geol.
1936.1 Sec; Terra, Hellmut de, and G. E. Hutchinson. Rundschau, XXIII, pp. 300-339, text fig. 1.
Beds of true Sarmatian age, bearing Hipparion.

Issel, A.
Lapparent, A. de
1879.1 Descrizione di due denti d'Elefante raccolti nella
LiQURiA occidentals. .\iin. Mus. Civ. Storia Nat. Geneva, XIV, 1906.1 Traite de Geologie. Fifth edition, 8vo, Mas.'^on et Cie.,

1-4. Paris, III, pp. 1289-2015, text figs. 584-883.

pp. 153-168, te.\t figs.
Remarks concerning the Qiningen fossils.

Johnson, F. Walker Frank

Leverett,
1936.1 The Sx.vrus of the Name "Valentine" in Tertiary Geology
1902.1 Glacial Form.\tions and Drainage Features op the Erie
AND Paleontology. Amcr. .lourn. Sci., (.5), XXXI, June, pp.
AND Ohio Basins. Monogr. U. S. Geol. Surv., XLI, pp. 1-802, Pis.
467-475, text figs. 1,2.
i-xxvi, text figs. 1-8.

Johnston, H. H. (Henry Hamilton] Lewis, G. Edward

No date The Openi.vg I p of .Africa. 16mo, Henry Holt and Com- 1937.1 A New Siwalik Correlation. .\mer. Journ. Sci., (5), XXXIII,
pany, New York, Williams and Norgate, London, 255 pp., 3 maps. March, pp. 191-204, text figs. 1, 2.

Tawi, new name for Boulder Conglomerate.

1906.1 A
History of the Colonization of .\frica by Alien Races.
8vo, Cambridge University Press, pp. xiii+349, 8 pis. (maps).
Licent, Emile, and M. Trassaert
1936.1 The Pliocene Lacustrine Series in Central Shansi. Bull.
Kay, George F.
Geol. Soc. China, XIV, pp. 211-219, text fig. 1, 2 maps.
1931.1 Classification and Duration of the Pleistocene Period.
Bull. Geol. Soc. Amer., XLI I, March 31, pp. 425-466, text figs. 1-9.
Lugn, Alvin L.
1936.1 The Plelstocene Geology op Nebraska. Neb. Geol. Surv,,
Kellogg, Remington (2), Bull. 10, pp. 1-223, Pis. I, II, text figs. 1-38, tables A-D.
1924.1 Tertiary Pelagic M.\mmals of Eastern North America.
Bull. Geol. Soc. Amer., XXXV, December .30, pp. 75.5-766.
1939.1 Classification of the Tertiary System in Nebraska. Bull.

.\ge of the Bone Vallej' formation.

Geol. Soc. Amer., L, No. 8, pp. 1245 1275, I pi.

Use of the long abandoned name Ogallalu for a large slratigraphic trroup r)f

Pliocene age.

Khomenko, J.

1913.1 La faune meotique du village Taraklia du district de MacCurdy, George Grant

BbNDERY. I. LeS ANCETRES des CeRVINAE. II. GiRAFFINAE ET 1924.1 Human Origins. A Manual of Prehlstory. Svo, D. Apple-
Cavicor.nia. .'Vnnuiiire Min. Ru.ssie, XV, Livr. 4-6, pp.
Geol. ton and Company, New York, I, pp. xxxviii 4-440, frontispiece, text
107-143, Pis. vi-ix. Novo-Alexandria. figs. 1-254; 11, i)p, xvi+516, frontispiece, text figs. 255 -410.
 BIBLIOGRAPHY 1627

Makiyama, Jiro Moir, J. Reid, and Arthur Tindell Hopwood

1929.1 Chikyu-Thc Globi', XII, No. o, pp. 364, 365 (in Jiipancso). Not 1939.1 Excavations .vr Brundon, Suffolk (1935-37). Part I.
available to tlie present author. Str.\tigraphy and Archaeology (Moir). Part II. Fossil Mam-
New subspecies: Elcpluis {Paixoloxod'jii) niimadicas aetoensis. mals (Hopwood). Proc. Prehist. Soc, (N. S.), V, No. 1, pp. 1-32,
Pis. I, II, text figs. 1-18.
1938.1 Japonic Proboscidea. Mem. College Sci. Kyoto Imp. Univ.,
(B), XIV, No. 1, Art. 1, May, pp. 1-59, text figs. 1-31.
New species; Bunolophodon yolcotii,\i\i. 12-14, figs, oa, 5b. Stegodon shodoensis Moody, C. L., John Campbell Merriam, and Chester Stock
akashiensis (Takai, 1936) = Parastegodon akashiensis (p. 21). Stegodon
insignis sugiyaniai (Tokunaga, 1936) =Parastegon sugiyamai (p. 27). 1926.1 Sec Merriam, John Campbell, Chester Stock, and C. L. Moody.

Mansuy, H. Morris, Frederick K., and Charles P. Berkey

1916.1 SuR QuELQUES Mammiferes Fossiles Recemment Decouverts 1927.1 See Berkey, Charles P., and Frederick K. Morris.
EN Indochine. Mem. Serv. Geol. Indochine, V, Fasc. II, pp. 1-26,
Pis. i-vii, 1 text fig. Osborn, Henry Fairfield
Stegodon insignis, S. Cliftii, Etephas sp. ? aff. E. naniadicus, E. naniadicus, and
E. indicus. 1933.901 Biological Inductions from the Evolution op the
Proboscidea. Proc. Nat. Acad. Sci., XIX, No. 1, January, pp.
1.59-163.
Mather, Cotton
1717.1 An Extract of Several Letters from Cotton Mather, D. D. 1936.940 The Record Proboscidean Tusk. Nat Hist., XXXV,
to John Woodward, M. D. .\nd Richard Waller, Esq.; S. R. April, p. 357. (Unsigned article.)
Record tusk of Archidiskodon imperalor from Texas.
Secr. Phil. Trans., XXIX, No. 339, for April, May, and June,
1714, pp. 62-71. 1936.961 L'AoE Geologique de l'Homme de Piltdown (Eoanthro-
pus) et de l'Homme de Trinil (Pithecanthropus). Melanges de
Prehistoire et d'Anthropologie offerts au Professeur Henri Begouen
Matsumoto, Hikoshichiro
a I'occasion de son 70" Anniversaire (20 Novembre, 1863-1933) par
1939.1 On Some Fossil Elephants from Province of Kazusa, from
ses Eleves, ses Collegues et ses Amis, pp. 23-36, text figs. 1-6.
Province of Shimotsuke, and from other Localities. Dobu- These sheets were received September 28, 1936, with a letter from Comte
tugaku Zassi (Zool. Mag., Tokyo), LI, No. 10, October, pp. 701-717, B^gouen in which he says; " L'impression fut meme un instant arret^e
. .

et elle vient de reconimencer. Je vousenvoie, sous pH separ6. en juatificatif

te.xt figs. 1-8. les premit^res bonnes feuilles. -"
. No other sheets have been received.
.

New species; Archidiskudon iMtramarnmonieus, pp. 704, and 710 (English).

1938.962 Eighteen Principles <.if Adapt.\tion in Alloiometrons
.\ND Aristogenes. Palaeobiologica, VI, pp. 273-302, text figs. 1-12.
Matthew, William Diller
1924.1 Third Contribution to the Snake Creek Fauna. Bull.
Paterson, T. T., Hellmut de Terra, and Pierre Teilhard de Chardin
Anier. Mus. Nat. Hi.st., L, Art. II, July 3, pp. 59-210, text figs. 1-63.
1936.1 See Terra, Hellmut de, Pierre Teilhard de Chardin, and T. T.
Paterson.
Matthew, William Diller, and Ruben Arthur Stirton
1930.1 Equidae from the Pliocene of Texas. Bull. Dept. Geol. Peale, Rembrandt
Univ. Calif., XIX, No. 17, November 29, pp. 349-396, Pis. xlv-lviii. 1802.1 Account of the Skeleton of the Mammoth, a Non-Descript
Carnivorous Animal of immense Size found in America. Sm.
Maxson, John H. 8vo, London, 46 pp.

1930.1 A Tertiary Mammali.an Fauna from the Mint Canyon 1803.1 An Historical Disquisition on the Mammoth, or, Gre.at
FoRM.\TioN OF Southern California. Publ. Carnegie Instn. Wash., American Incognitum, .\n Extinct, Immense, Carnivorous
No. 404, August, pp. 77-112, text figs. 1-lS. Animal, whose Fossil Remains have been found in North
America. 12mo, E. Lawrence, London, pp. viii+Ol, 1 pi.

Merriam, John Campbell

1919.1 Tertiary Mammalian Faunas of the Mohave Desert.
Pei, W. C, Davidson Black, Pierre Teilhard de Chardin, and Chung-
Chien Young
Bull. Dept. Geol. Univ. Calif., XI, No. 5, pp. 437a-e, 438-585, text
figs. 1-253. 1933.1 Sec Black, Davidson, Pierre Teilhard de Chardin, Chung-Chicn
Fauna of the Barstow formation. Young, and W. C. Pei.

Merriam, John Campbell, Chester Stock, and C. L. Moody Pfizenmayer, F. W.

1926.1 The Pliocene Rattlesnake Formation and Fauna of 1937.1 Mammut-Funde in Sibirien. Natur u. Volk, LXVIl,
Eastern Oregon, with Notes on the Geology of the Rattle- pp. 279-288, text figs. 6-14.
Sangajurach-Mammuts.
snake and Mascall Deposits. Publ. Carnegie Instn. Wash., No.
347, October 8, pp. 43-92, text figs. 1-45.
Piette, Edouard
1907.1 L'Art pend.ant l'Age du Renne. 4to, Masson et Cie., Paris,
Meyer, Hermann von pp. iv-f-12, 100 pis., 128 text figs.
1841.1 FossiLE Knochen von Wiesbaden. Neues Jahrb. Min., pp.
4.58-461.
Pilgrim, Guy EUcock
New species; Dinolheriiiin minntnm, p. 4.'>0.

1931.1 Catalogue ok the Pontian Carnivora of Europe in the

DiiPARTMENT OF Geology. 4to, British Mu.seuni (Natmal History),
Middendorf, Alexander Theodor von London, pp. vi-f-174, 2 pis., 30 text figs.
1860.1 Ubersicht DER N.-VTUR NoRD- UND Ost-Sibiriens. Reise in den
Aussersten Norden und Osten Sibiriens wahrend der Jahrc 1843 und 1934.1 Correlation of Ossiferous Sections in the Upper Cenozoic
1844. . ., IV, Th. I, Lief. 2, pp. 201-332. 4to, St. Petersburg. of India. Amer. Mus. Novitates, No. 704, March 15, pp. 1-5.
1628 OSBORN: THE PROBOSCIDEA

Pilgrim, Guy Ellcock — Continued 1930.1 Alounos comentario.s sobre .mastodontes chilenos. Revista
1939.1 The Fossil Bovidae of India. Mem. Geol. Surv. India, Universitaiia (Univ. Cat61ica dc Chile), No. 8, Ano XV, pp. 886-893.
Palaoont. Indica, N. S., XXVI, Mom. 1, pp. iii+356,8pls., 35 text figs. Santiago.
Remarks on the questionablp typo lo(-;ility of Mofltidon [CuvieroniuA It it iitholdtii.
1940.1 The Application of the Europe.\n Time Scale to the
Upper Tertiary of North America. Geol. Mag., LXXVTI, pp.
1-27. Schroeder, Henry
1928.1 t'sER Elephas .\ntiquu8 und trogontherii aus DEM Diluvium
Pilgrim, Guy Ellcock, and Arthur Tindell Hopwood DER Mittelmark. Jahib. Preuss. Geol. Landesanstalt, Berlin,
XLVIII, for the year 1927, pp. 699-723, Pis. xxxiv-.xxxvi, text fig. 1.
1939.1 .\FiE THK EgriDAE Reliable koh the Correlation of the
New subspecies: Elepltas prijnifjeniu.-^ Blumenb. var. pachygnnalis, p, 718.
ii.

SiWALiKS with the Coenozoic Stages of North America (Pilgrim).

Appendix on the Correlation of Certain Tehtiarv Deposits
of India and Europe (Hopwood). Rec. Geol. Surv. India, LXXIII, Schultz, Charles Bertrand, and Erwin Hinckley Barbour
Pt. 4, Decemb(M-, 1938, pp. 437-482. 1937.1 See Barbour, Erwin Hinckley, and Charles Bertrand Schultz.

Piveteau, Jean, and Marcellin Boule

Scott, William Berryman
1935.1 .See Boule, MarculMii, and Jean Piveteau.
1937.1 A History of L.\nd Mammals in the Western Hemisphere.
Revised edition, 8vo, Macmillan Company, New York, pp. xiv+786,
Pohlig, Hans frontispiece, 420 text figs.
1887.3 [Casts of Elephant Teeth, E. antiquus typds .\nd E. .\ntiqu- New family: Stegoraastodontidie. New subfamilies: Trilophodontin«,
Pentalophodontinte, Cordillerionin:e, Stegomastodnntin:e.
us var. minor] Verb, natur. Vereins preuss. Rhein., Jahrg. XLIV,
p. 115. Bonn.
New subspecies: E. \Elcpfias] antiquus mr. minor. Serebr3rakov, A. H.
1938.1 Elephas mammonteus Cuvier versus E. Primigenius Blumen-
Pontier, G.
bach. Bull. Acad. Sci. U. S. S. R., Classe des Sci. Math, et Nat.,
1930.1 ."V PROPOS i)'.\nomalies dentaires observees CHEZ LES Probo- Serie Biol., pp. 1063-1068.
srioiKNs. .'VnM. Soc. gAjl. du Nord, LV, pp. 2-10, Pis. I, ii. He regards the name Elephas primigenius Blumcnbach (1799) as an indis-
putable nonien nudum, hence must be rejected, and maintains that E.
mammonteus Cuvier (1799) must be restored, recognizing as the tyjie the
skull described bv .Messerschmidt in 172-1, or .Adams' mammoth (Tilesius,
Roman, F., and J. Viret 1813).
1934.1 La Faune db Mammiferes du Burdicalien de la Romieu
(Gers). Mem. Soc. geol. France, (N. S. ),IX, Fasc. 2-3, (Mem. No.
Shikama, Tokio
21), pp. 1-67, Pis. i-xix, text figs. 1-25.
1936.1 Note on Parastegodon .\kashiensis Takai from the Akasi
District. Proc. Imp. Acad., Tokyo, XII, No. 1, January, pp.
Romer, Alfred Sherwood 22-24, text figs. 1-4.
1933.1 Pleistocene Vertebrates and their Bearing on the
Problem of Hum,\n .\ntiquity in North America. In "The 1937.1 Parastegodon infrequens sp. nov. from the Akasi District.

.\merican Aborigines, Their Origin and Antiquity," edited by Japanese .lourn. Geol. and Geog., XIV, Nos. 3-4. October, pp. 127-
Diamond Jenness. 8vo, University of Toronto Press, pp. 47-83. 131, PI. IX.

1937.2 Nomenclative Notes on Parelephas Protomammonteus

Roverto, Cayetano (Matsumoto). Japanese Journ. Geo!, and Geog., XIV, Nos. 3-4,
1914.1 l.os Estr.vtos Araucanos y Sus Fosiles. An. Mus. Nac. October, pp. 163-166.
Hist. Nat. Buenos Aires, XXV, pp. 1-249, Pis. i-xxxi, text figs. 1-92. New subspecies: Parelephas proximus uelialaensis.

Rusconi, Carlos Shuler, Ellis W.

1937.1 CONTRIBUCION AL CONOCIMIENTO DE LA OEOLOOIA DE LA CIUDAD 1934.1 Collecting Fossil Elephants .\t Dallas, Texas. Bull.
DE Buenos Aires y sus alrbdbdohes y referencia de su fauna. Texas Archaeol. and Palaeont. ,Soc., VI, September, pp. 7.5-79, PI. xn.
Actas Acad. Nac. Cicn. C6rdoba, X, Nos. 3-t, December 20, pp.
177-384, Pis. i-xix, text figs. 1 ,58. Buenos Aires.
Soergel, Wolfgang
Russell, R. Dana, and V. L. VanderHoof 1921.3 Die Ursachen der diluvialbn Auf.schotterung und Erosion.
8vo, Borntraeger, Berlin, pp. iv+74, 1 text fig.
1931.1 Vertebrate Fauna from a New Phocbnh Formation in
.\

Northern California. Bull. Dept. Geol. Univ. Calif., XX, No. 2,

February .5, pp. 11-21, text figs. 1-7. 1926.1 Die S.xuqbtibrfauna des altdiluvialen Tonlagers von
Tehama formation named and described. Contains Slegotnaatodon cf. ariionse. JocKGRiM IN DER Pfalz. Zeits. Deiitsch. Geol. Ges., LXXVII,
Heft 3, November 20, pp. 405-438, Taf. xvii, 2 tabellen.
Saheki, Shiro
1931.1 On Parelephas protomammonteus (Matsumoto) Recently Spock, Leslie Erskine
Found in the Province of Kazusa. Japanese .lourn. Geol. and 1930.1 New Mesozoic and Cenozoic Form.\tu)Ns encountered by
Gcog., VIII, No. 3, Fi-bruary, pp. 12.t 129, PI. xv, 1 (cxt fig.
THE Central Asiatic Expeditions in 1928. Amer. Mus. Novitates,
New Hubspeeies; Parelephas prulomavunonteiis (Matsumotn) maisumoloi. n.
127.
No. 407, March 18, pp. 1-8, text figs. 1-6.

Schneider, Carlos Oliver Stamp, L. Dudley

1929.1 La Di.stribuci6n GEooK.iFiCA de los Mastodontes en Chile. 1922.1 An Outline of the Tertiary Geology of Burma. Geol, Mag.,
Actcs Soc. Scientifique Chili, XXXV'I, pp. 73-83, 1 map. Santiago. LIX, No. 11, November, pp. 481-501, text figs. 1-6.
 BIBLIOGRAPHY 1629

Stehlin, Hans Georg Teilhard de Chardin, Pierre, Davidson Black, Chung-Chien Young,
1908.1 Notices paleomammalogiques sur quelqdes depots Mio- and W. C. Pei
cfcNES DES BaSSINS DE LA LoiHE ET DE l'AlLIER. BuU. SoC. gool. 1933.1 See Black, Davidison, Pierre Teilhard dc Chardin, Chung-Chien
France, (4), VII, for the year 1907, pp. 525-550, text figs. 1-3. Young, and W. C. Pei.

Stehlin, Hans Georg, and Auguste Dubois Teilhard de Chardin, Pierre, and Hellmut de Terra
1932.1 and 1933.1 See Dubois, Auguste, and Hans Georg Stehlin. 1936.1 See Terra, Hellmut de, and Pierre Teilhard de Chaidin.

Sternberg, C. M. Teilhard de Chardin, Pierre, Hellmut de Terra, and T. T. Paterson

1930.2 Miocene Gravels in Southern Saskatchewan. Trans. Roy. 1936.1 See Terra, Hellmut de, Pierre Teilhard de Chardin, and T. T.
Soc. Canada, (3), XXIV, Sec. 4, pp. 29, 30. Paterson.
The scanty and fragmentary fauna includes an indeterminate Mastodont.

Teilhard de Chardin, Pierre, and M. Trassaert

Stlrton, Ruben Arthur
1937.1 The Pkoboscide.vns of South-Easteun Shansi. (Yushe
1933.1 A Critical Review of the Mint Canyon Mammalian Fauna
basin). Pal, Sinica, (C), XIII, Fasc. 1, March, pp. 1-84 (also 4 pp.
and its Correlative Significance. Amer. Journ. Sci., (5), XXVI,
in Cliinese), Pis, i-xiii, text figs. 1-6.
December, pp. 569-576.
New species: Pentalopliodon cuneatits, p. 11; .yfnstodon intermedium, p. 22;
Stefjodun licenti. p. 27.
1936.1 Succession of North American Continental Pliocene
Mammalian Faunas. Amer. Journ. Sci., (5), XXXII, pp. 161-206.
Teilhard de Chardin, Pierre, and Chung-Chien Young
1939.1 Methods and Procedure in THE Valentine Question. Amer. 1936.1 On the Mammalian Remains from the -\hch.eolooical Site
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Stlrton, Ruben Arthur, and William Diller Matthew
1930.1 See Matthew, William Diller, and Ruben .\rthur Stirton. Terra, Hellmut de
1936.1 Late Cenozoic History in India. Nature, CXXXVII, No.
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1936.1 D.\ta on Post-Glacial Clim.atic Changes in North-West
Trihphodon sp., consisting of fragmentary skull, incomplete mandibular India. Current Science, V, July, pp. 5-10. Bangalore City.
ramus, humerus, femur, and a calcaneum, found at three distmct localities
in the Ricardo deposits.

1936.1 A Pliomastodon Skull from the Thousand Creek Beds, Terra, Hellmut de, and Pierre Teilhard de Chardin
Northwestern Nevada. Publ. Carnegie Instn. Wash., No. 473, 1936.1 Observations on the Upper Siwalik Form.ation and L.ateu
July 10, pp. 35-39, PI. i.
Pleistocene Deposits in India. Proc. .^mer. Phil. Soc, LXXVI,
New species: Pliomastodon nfvaiianus, p. 37.
pp. 791-822, text figs. 1-14.

Stock, Chester, John Campbell Merriam, and C. L. Moody Terra, Hellmut de, Pierre Teilhard de Chardin, and T. T. Paterson
1926.1 See Merriam, John Campbell, Chester Stock, and C. L. Moody.
1936.1 Joint Geological and Prehistoric Studies of the Late
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1907.1 Geologische Beobachtungen im Fajum und am iwteren
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Hyoqo Prefecture, Japan. Proc. Imp. Acad., Tokyo, XII, No. 1, Tobien, H.
January, pp. 19-21, text figs. 1, 2.
Uber Hipparion-Restb aus der obermioz.anen SOsswasser-
1938.1
New species: Parastegodon akaahiensis, p. 20.
molasse Sudwestdeutschl.\nds. Zeits. Deutsch. Geol. Ges., XC,
1936.2 Fossil Elephants from Tiba Prefecture, Jap,\n. Japanese Heft 4, May 5, pp, 177-192, T.af, vi.
Journ. Geol. and Geog., XIII, Nos. 3-4, October, pp. 197-203, PI.
XXIV, text fig. 1.
Tokunaga, Shigeyasu
The author states that from "this district, the following species have thus far
been reported: 1933.2 A List of the Fossil Land Mammals of Japan and Korea
Steyodon orientalis Owen.
Parelephas protonuivimonteus typicus Matsumoto. with Descriptions of New Eocene Forms from Korea. .\mer.
P. protoniammonteits proximus Matsumoto.
P. protomammontcus matsitmotoi Saheki.
Mus. Novitates, No. 627, May 27, pp. 1-7, text figs. 1, 2.
Paiaeoloxodon iioumaiiiu (Makiyama)."
1934.1 Fossil Eleph.\nt teeth found at Yokohama and Kakio,
K.ANAGAWA Prefecture. Journ. Geog., XLVI, No. 546, July, pp.
Takai, Fuyuji, and Shigeyasu Tokunaga
363-371, Pis. VIII, IX, 5 text figs. Tokyo.
1936.1 .'-'(e Tdkunaga, Shigeyasu, and Fuyuji Takai. New species: Paiaeoloxodon yokoharnanus, p. .363, PI. Vll; Parastegodon?
kwantoensis, p. 365. PI. ix. Text in .Japanese.

Teilhard de Chardin, Pierre 1936.1 A New Fossil Elephant found in Shikoku, Japan. Proc,
1937.1 The Post-Villafranchian Interval in North China, Bull. Imp. Acad., Tokyo, XI, No. 10, December, pp. 4.32 434, text fig. 1.
Geol. Soc, China, XVII, No, 2, June, pp, 169-176, text fig. 1. New species: Parastegodon sugiynmai.
1630 OSBORN: THE PROBOSCIDEA

Tokunaga, Shigeyasu Conliiuitd Viret, J., and F. Roman

1936.1 Geology of the District of Shichinohe and Fossil Ele- 1934.1 See Roman, F., and J. Viret.

phant FOUND there. Jouin. Gpog., XI.VHI, No. 064, February,

pp. 67-70, PI. I. Tokyo.
Original description fin Japanope) of Palneohixotiun aomorienais, p. 70.
Ward, Rowland
1922.1 Rowland Ward's Records of Big Game with Their Distri-
Tokunaga, Shigeyasu, and Fuyuji Takai bution, Characteristics, Dimensions, Weights, and Horn and
1936.1 U.N A Fo.ssiL Elephant, Palaeoloxodon aomokiensis, from Tusk Measurements. Eighth edition edited by J. G. DoUnian
Shichinohe, Kamik.\t.\-gux, Aomohi Prefecture, Japan. Journ. and J. B. Burlace, 8vo, Rowland Ward, London, pp. xiii +527,
Geol. Soc. Japan, XLIII, No. 511, April 20, pp. 254-258, Pis. xiii illustrated.

(in)i XIV (iv).

Type tlescription (in Eneli?!)) uf Palaeoloxodon nomorirnais. 1928.1 Rowland Ward's Records of Big Game with Their Distri-
bution, Ch.aracterlstics, Dimensions, Weights, and Horn and
Trassaert, M., and Emile Licent Tusk Measurements. Ninth edition edited by J. G. Dollinan and
1936.1 .So I.iccnt, Eiiulr, and M. Trassaert. J. B. Burlace, 8vo, Rowland Ward, London, pp. xiii-1-523, illustrated.

Trassaert, M., and Pierre Teilhard de Chardin

1937.1 .S'c Tcilliard dc Chardin, Pierre, and M. Tra.ssaert. Weber, Max
1896.1 (18971 Vorstudien iiber das Hirngewicht der Saugethiere.
Troxell, Edward L. Festschrift z. Siebenzigsten Geburtstage von Carl Gegenbaur, III,

1916.1 .\n Early Pliocene One-Toed Horse, Pliohippus lullianus, pp. 103-123.

sp. NOV. .\mer. Journ. Sci., (4), XLII, October, pp. 335-348, text figs.

17.
Oak C'rcfk foriimtion naniofi antl described, pp. 34.5-34.S.
Wilmarth, M. Grace
1938.1 Lexicon of Geologic Names of the United St.\tes (includ-
VanderHoof, V. L. ing Al.\ska). Bull. U. S. Geol. Surv. 896: Part 1, A-L, pp. 1-1244;
1933.1 Additions to the Fauna of the Tehama Upper Pliocene of Part 2, M-Z, pp. 1245-2396.
Northern California. .\mrr. .lourn. Sci., (5), XXV, pp. 382-384.

VanderHoof, V. L., and R. Dana Russell Young, Chung-Chien, and Pierre Teilhard de Chardin
1931.1 See Rus.sell, R. Dana, and V. L. VanderHoof. 1936.1 Sei' Teilhard dc Chardin, Pierre, and Chung-Chien Young.

VanEs, L. J. C.
1931.1 The Age of Pithecanthropus. 8vo, Martinus Nijhoff, The Young, Chung-Chien, Davidson Black, Pierre Teilhard de Chardin,
Hague, pp. xii-|-142, 4 pis., 11 maps. and W. C. Pei
1933.1 See Black, Davidson, Pierre Teilhard de Chardin, Chung-Chien
van Riet Lowe, C. Young, and W. C. Pei.
1929.1 Firther Notes on the Archaeology of Sheppard Island.
So. Afr. Journ. Sci., XXVI, pp. 66.5-683, text figs. 1-5.
Kxeellent tabli- ofassociated fauna during the Plcistoeene, ineluding anione tile Zittel, Karl Alfred von
Proboseidea Mastodon {Buiialuplojdon) sp. {=Trilvpliodon], Archidishodon
Mitfiptanijrons, and .1. I — Fol^oloxodoit] transvaaknsii< and .4. 1 = Polxoloxo- 1926.1 Text-book of Palaeontology. Ill, Mammalia. 8vo, Mac-
don] ahrppnrdi of the early Pleistocene, to A. brooini of the late Pleistocene.
Stellenhoseh industry of the Stone Age. millan and Company, London, pp. viii+316, 374 text figs.
PLATES XXVI— XXX
PLATE XXVI
 PLATE XXVI
Fig. 1. Eli-phas intticus. Complete transverse section near tip of unerupted tusk, probably of an immature female. Ordinary light. Seven times
natural size.
The dark central area is dentine. In it, to the left, is a triangular area of tubular or cylindrical structure, suggestive of the development in Flalybelodon and
its allies, but absent in the later dentine of Elephas. The "engine-turning" effect is not visible in this juvenile dentine. An enamel band encircles about two-
thirds of the circumference of the dentine, being absent only to the right and upper Much of it has been lost in making the section to
right in the photograph.
the left, where it is thickest, but elsewhere it is clearly visible as a relatively transparent and homogeneous tissue. The whole tusk is here encased in a thick
layer of cement, irregular in structure but in general with relatively dense, laminated intern,al and external zones and a thicker, non-laminated intermediate
zone, with large canals rather like the Haversian canals of bone.

Fig. 2. EUphas indicus. Area of thin section within the circle on figure 1. Plane light. Nineteen times natural size.

The dark, lower region is cement. Between these, to the left, is the
dentine and the irregular, spotty mass forming the greater part of the photograph is

enamel band, ending naturally near the middle of the photograph. It is sharply defined, the dentine border undulating and the cement border rugose. In the
cement, the part to the right clearly shows the three layers, inner and outer dense and vaguely laminated, middle spongy, all with innumerable lacuna;.

Fig. 3. Elephas indicus. Same as figure 2, photographed between crossed nicols.

The enamel is characterized by high birefringence and relative simplicity and homogeneity of rod arrangement, with a narrow zone of deflection, dark as
pli(>togra[)lic(l, parallel to the dentine border. The cement also shows birefringence but is highly irregular with little consistent orientation c.\cept in the
iiinor (lower) layer.
Osborn: The PnoBOfeciDEA, II Plate XXVI
PLATE XXVII
 PLATE XXVII
Fig. 1. Elephasindictts. Area within the circle on PI. xxvi, Fig. 2. Plane light. Three hundred thirty-seven times natural size.

In this innermost area of the cement, just above the enamel, lacuna? are very nuraerou.s and have many branching canaliculi. There are also more sparse,
small tubules, one of which appears in the upper right in this figure.

Fig. 2. Elephas indicus. Transverse section of about half of a small, mature tusk about 14 inches long. Ordinary light. Four times natural size.

The section taken about 3 inches from the tip of the tusk. Several inches had been worn off, including all the enamel and most of the cement. At this
is

point, there is only a thin band of cement, seen in the figure as a more translucent coating above the dense dentine. The dentine as a whole shows three distinct,
superimposed types of structure: (a) radiating tubes, too small to be clearly seen in this figure, directed away from the center of the tusk, (b) well-marked,
fine, concentric lamellse, (c) curving, criss-cross markings of coarser character, visible as dark lines in the thinner parts of this section, that give the "engine-

turning" effect. This last effect appears to be the result of regularly recurring undulations in the courses of the dentine tubes and in the orientation of the
surrounding calcified mass. (The prominent white lines are scratches on the section.)

Fig. 3. Elephas indicus. Part of the section within the circle on figure 2. Plane light. Forty times natural size.

This shows the whole thickness of the cement on this part of the tusk. In ordinary light the cement here appears vaguely laminated, with fairly numerous
lacuna; but no trabeculse or canals. The boundary against the dentine is smooth and very sharp.

Fig. 4. Elephas indicus. Same as figure 3, with crossed nicols. The effect of strong birefringence in the cement is to bring out a wavy, interwoven,
fabric-like structure in this tissue.

Fig. 5. Elephas indicus. A small area near the outer edge of the cement band of the section shown in figures 2-4. Plane light. Three hundred
thirty-seven times natural size.

This shows a fabric-like texture similar to that seen more plainly and on a larger scale in figure 4. The lacume have very few and very short canaliculi
and most of them appear to be empty (or filled with colorless fluid) in this section of a fresh tooth. Contrast with this figure 1, which shows the abundant
canaliculi and opaque appearance typical of younger cement. The inner part of the cement in the present section, near the dentine, is about intermediate
between the two conditions.

Fig. 6. Phiomia winloni. Transverse thin section about one-half inch from tip of small, worn tusk. Plane light. Thirty-eight times natural size.

The lower part of the section is dentine and above this isthe whole thickness of the enamel, cut at right angles to its surface. The clear bands in the
dentine are caused by infiltration of the embedding medium into the cut ends of tubes and these zones, in fact, have tubes like those seen so clearly in the lower
part of the picture. For details of enamel and of enamel-dentine junction see PI. xxviii. Fig. 2. There is no indication of cement on this tusk.

J
Osborn: Thk Proboscidea, II Plate XXVII

" 'i-n f^ • -'-

/. .
.
'^-4

if*
PLATE XXVIII
 PLATE XXVIII
Fig. 1. I'Iduinia wiiduiti. Same as PI. .x.wii, Fig. 6, with crossed nicols.

The dentine is not birefringent. The enamel is strongly birefringent and variations in extinction angle show the varying orientations of the rods through
the greater part of the thi( kness of the enamel, but near the outer surface (up in the photograph) the illumination is even and shows the rods there to be es-
sentially parallel.

Fig. 2. Phiomin winloni. .\rea within the circle on PI. xxvii, Fig. fi. Ordinary light, .\bout three hundred times natural size. Photograph liy Dr. J.

T/Bon Williams.

The small bottom area in the photograph shows part of the granular layer of the dentine. Above this is the sharp enamel boundary, and then the full
thickness of the enamel. Natural staining and variable refringence clearly brings out the pattern of the enamel rods, which lie in long, open spirals through
most of the thickness of the enamel and become approximately straight and perpendicular to the surface only in the outermost zone, which is on the order of
.02 mm. in thickness. The enamel as a whole is here about ..5 mm. in thickness.
Osborn: The Proboscidea, II Plate XXVIII
PLATE XXIX
 PLATE XXIX
Fig. 1. Trilophodon obscurun. Transverse section of part of tusk. Ordinarj' light. Four times natural size.

The bulk of the section is in the which the radial and concentric structures are shown, but not the "engine-turning" effect, which was, never-
dentine in

theless, present, .\bove this the whole width of the enamel band is shown. There is no cement on the specimen as preserved, but this probably occurred
when the tusk was fresh.

Fig. 2. Trilophodon obsmrus. Area within circle on figure 1. Crossed nicols. Forty times natural size.

In this region the enamel rods are only very gently and obscurely spiral and tend rather to be parallel to each other in simple curves from the dentine to
the outer surface. Toward the edge of the enamel band, to the left, the rods are strongly curved and the enamel overhangs a natural pocket. .'Vn isolated

stringer of enamel occurred beyond this and its edge is barely visible at the left margin of the photograph. The enamel is also marked throughout by fine
striations, not very prominent in the photograph, parallel to the outer surface even when this is strongly curved, as at tlie left of the photograph, and hence
approximately at right angles to the enamel rods throughout. The dentine is very feebly birefringent and shows uneven lamination parallel to the surface.

Fig. 3. Trilophodon ohscurus. Same tusk as figures 1 and 2, thin section across enamel cut in a plane vertical to the surface and parallel to the longi-
tudinal axis of the tooth. Crossed nicols. Forty times natural size.

.\way from the edge of the enamel band, the rods show a slight spiral arrangement much like that of Phiomia (PI. .xxviii) in kind but far less in degree.
somewhat more prominent in this photograph.
Striations as seen in figure 2 are

Fig. 4. Trilophodon {Megnbdodon) sp. Transverse thin section of tusk near edge of enamel band. Plane light. Eleven and a half times natural size.

The area covered by the lower part of the photograph was occupied by dentine, fragments of which can be seen although most of it h.as broken away.
Above this is the enamel and cement covering. To the left is the edge of the main band of enamel. Beyond this to the right were five long, isolated string-
ers of enamel. Transverse sections of two of these are seen in the middle and to the right in the photograph and another is cut by the right margin. Between
the main enamel band and the stringer in the middle of the ph(jtograph is a mass of cement. From thinner films around the stringers and main band and from
the character of the enamel surface it appears that the enamel was all embedded in a coating of cement and was wholly exposed only at the wearing edge. The
cement also extends in a thicker coating around the dentine beyond the last enamel stringer, outside the area of the photograph.

I'"ig. ."). Trilophodon (M egabelodon) sp. A different part of the same thin section as figure 4. Crossed nicols. Nineteen times natural size.

This photograph includes the last two enamel stringers, to the right of the tliree visible in figure 4. The strong birefringence clearly brings out the
peculiar, fan-like divergence of the enamel rods in these stringers, which is equally developed in the other .stringers and at the edge of the main enamel
band. The stringens are separated and in part surrounded by a mass of cement so feebly birefringent that it can hardly be seen in the photograph.
Osborn: The Proboscidea, II Plate XXIX

rMiWi^JTil

VrTi^PTl^iSiE '* 7//««K

PLATE XXX
 PLATE XXX
Fig. 1. Trilophodon (Megabelodon) sp. Vertical thin section of broken molar tubercle. Ordinary liglit. Four times natural size.

The worn apex of the tubercle is and enamel to the right. The enamel here reaches a thickness of about 5 mm. .4side from
at the to[), dentine to the left,
the mineral staining, largely casual but in part emphasizing natural structure, the most striking thing is the presence and orientation of the numerous enamel
striatioDs or laminations.

Fig. 2. Trilophodon (Megabelodon) &\t. Part of section in circle on figure 1. Crossed nicols. Twenty times natural size.

To the left may be seen worn into a pit above. The margin of the enamel is also a wear surface. Within the enamel the striations
a small part of the dentine,
are emphasized by mineral stain as well as by birefringence and are very prominent, especially in the more internal part of the enamel where they are more
wavy than externally. The enamel rods are less clearly distinguishable. In a thin inner zone, very obscure in the photograph (but see Fig. 4), they appear
to be spiral. In the bulk of the section they are simple, nearly parallel, arranged in a sweeping curve becoming more horizontal as they pass from the inner
(left) to outer (right) side of the enamel coating.

Fig. 3. Trilophodon (Megabelodon) sp. Transverse thin section of a molar cusp, from the same tooth as figures 1 and 2 but a different cusp. Ordinary
light. Four times natural size.

In the center is the circular core of dentine, about 4 mm. in left) the enamel, 5-7 mm. in
diameter, and surrounding this (except where broken away, lower
thickness. The dentine shows no pronounced structure. The enamel has an inner zone, about 1 mm.
with vague concentricity but no strong
in thickness,
striations, and an outer zone of 4-6 mm. with pronounced concentric striations, which are homologous with those cut in a different plane in figures 1 and 2.

Fig. 4. Trilophodon (Megabelodon) sp. Part of section in inner circle on figure 3. Plane light. Forty times natural size.

To the left is the dentine with sharply visible tubules and terminating in a granular layer below the enamel. The light band down tlie middle of the
photograph is the thin inner zone of enamel. Mineral staining shows the rods to be oriented in an irregular, highly complex, spiral pattern.

Fig. o. Trilophodon (Megabelodon) sp. Part of section in outer circle on figure 3. Plane light. Twenty times natural size.

This shows the outer part of the enamel, with its well-defined concentric striations and with the rods, not very clear in the photograph, generally radial
and much simpler in orientation than in the innermost enamel.
Osboun: The Proboscidea, II
Plate XXX

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INDEX
Page references in small roman figures refer to Volume I only

Abel, Othenio, 4, 13, 95, 100-102, 761 Ahiska, 137, 176, 177, 736, 7.53, 1088, 1091, 1099, 1127, 1134, 11.35, 1145, 1156,
abcli (see Trilophodon abeli) 1157, 1159, 1169, 1201
Aberdare Mountains, 1193 Alaska Agricultural College and Scliool of Mines, 1 161 1609, 161 ,

Abich, Otto Wilhelra Hermann von, 3, 761 Alaska CoUege-Frick American Museum Expedition, 11.59-1161
Absolon, Karel, 4, 13, 761, 1139, 1168 Alaska- Yukon (see Yukon)
Abtsdorf, 81 alaskensis (see Mammonteus primigenius alaskcnsis, and Mastodon americanus
Abyssinia, 117, 1176, 1200, 1417 alaskensis)
Academy of Natural Sciences, Philadelphia, 15, 209, 745, 1609, 1619 Alas-Tuwa, 885, 886
Academy of Sciences (Zoological Museum, Leningrad), 1128, 1148 Albany, New York, 136, 1602
Acconci, Luigi, 3, 761, 1187, 1230 Albert Nyanza, 995, 1193
Aceratherium. 117. 272, 1429, 1459, 1461, 1479 albertensis (see Loxodonta africana albertensis)
Acheulean, 1169, 1430, 1433 Alces latifrons, 971. 1155
acrocephaly, 919, 1552 Alcoi, Spain, 114
Aculcingo, Cannada de, Mexico, 537, 558, 740 Alekseev [Alexejew], A., 1466, 1623
acutidens (see Mastodon aculidens)