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HARVARD UNIVERSITY
Library of the
Museum of
Comparative Zoology
B R i
V I R A
niseuiiii of Comparative Zoology
HARVARD UNIVERSITY
Numbers 464-487
1981-1986
MC2
LIBRARY
'
NOV 1 9 ir
CAMBRIDGE, MASSACHUSETTS 02138 U.S.A.

1986
BREVIORA
Museum of Comparative Zoology
CONTENTS
Numbers 464-487
1981
No. 464. The Origin of the Crocodiloid Tarsi and the Inter-
relationships of Thecodontian Archosaurs. By Donald
Brinkman. 23 pp. December 30.
No. 465. The Structure and Relationships of the Dromasaurs

(Reptilia: Therapsida) By Donald Brinkman. 34 pp.
December 30.
1982
No. 466. Systematics of the Mexicana Species Group of the

Colubrid Genus Lampropeltis, with an Hypothesis
Mimicry. By William R. Garstka. 35 pp. June 30.
No. 467. Three New Species of the Anolis Punctatus Complex

from Amazonian and Inter-Andean Colombia, with
Comments on the Eastern Members of the Punctatus
Species Group. By Ernest E. Williams. 38 pp. June
30.
No. 468. A New Forest Skink from Ponape. By A. Ross Kiester.

10 pp. June 30.
No. 469. Catalog of the Primary Types of Bombyliidae (Diptera)

in the Entomological Collections of the Museum of
Comparative Zoology, with Designations of Lecto-

types. By Neal L. Evenhuis. 23 pp. June 30.
No. 470. Systematic Implications of Innervation Patterns in Tele-

ost Myotomes. By Quentin Bone and R. Dana Ono.
23 pp. June 30.
No. 471. Arthur Loveridge —A Life in Retrospect. By Ernest E.
Williams. 12 pp. June 30.
No. 472. Fishes of the Suborder Labroidei (Pisces: Perciformes):

Phylogeny, Ecology, and Evolutionary Significance.
By Leslie S. Kaufman and Karel F. Liem. 19 pp. June
30.
1983
No. 473. The Interrelationships of Pelycosaurs. By Donald

Brinkman and David E. Eberth. 35 pp. April 29.
No. 474. Ruthiromia Elcobriensis, A New Pelycosaur from El

Cobre Canyon, New Mexico. By David A. Eberth
and Donald Brinkman. 26 pp. April 29.
No. 475. New or Problematic Anolis from Colombia. L Anolis

Calimae, New Species,from the Cloud Forest of
Western Colombia. By Stephen Ayala, Dennis Har-
ris, and Ernest E. Williams. 1
pp. April 29.
1
No. 476. Townsend's Unmapped North Atlantic Right Whales

(Eubalaena Glacialis). By William E. Schevill and
Karen E. Moore. April 29.
1984
No. 477. New or Problematic Anolis from Colombia. IL Anolis

Propinquus, Another New Species from the Cloud
Forest of Western Colombia. By Ernest E. Williams.
7 pp. September 7.
No. 478. New or Problematic Anolis from Colombia. IIL Two

New Semiaquatic Anoles from Antioquia and Choco,
Colombia. By Ernest E. Williams. 22 pp. September 7.
No. 479. Agonistic and Courtship Displays of Male Anolis Sagrei.

By Michelle P. Scott. 22 pp. September 7.
1985
No. 480. Three New Lizards of the Genus ^^©/^(Scincidae) from

Southern New Guinea. By Waher C. Brown and Fred
Parker. 12 pp. June 21.
No. 481. A New Anolis of the Lionotus Group from Northwest-

ern Ecuador and Southwestern Colombia (Suaria:
Iguanidae) By Kenneth Miyata. June 21.
No. 482. New or Problematic Anolis from Colombia. IV. Anolis

Antioquiae, New Species of the Anolis Eulaemus
Subgroup from Western Colombia. By Ernest E.
WilUams. 9 pp. June 21.
No. 483. Notes on Pristidactylus (Sqamata: Iguanidae). By

Richard Etheridge and Ernest E. Williams. 18 pp.
June 21.
No. 484. Male Aggressive Behavior in a Pair of Sympatric Sibling

Species. By Jonathan B. Losos. 30 pp. June 21.
1986
No. 485. The Anatomy and Relationships of Stereophallodon

and Baldwinonus {KepiiMa., Pelycosauria). By Donald
Brinkman and David A. Eberth. 36 pp. August 30.
No. 486. Thelodus "Macintoshi" Stetson 1928, The Largest

Known Thelodont (Agnatha: Thelodonti). By Susan
Turner. 20 pp. August 30.
No. 487. The Identity of Larval Parasudis (Teleostei, Chloroph-

thalmidae), with Notes on the Relationships of Auli-
poform Fishes. By Karsten E. Hartel and Melanie L.
J. Stiassny. 24 pp. August 30.
BREVIORA
Index of Authors
Numbers 464-487
1981-1986
Ayala, Stephen 475
Bone, Quentin 470
Brinkman, Donald 464, 465, 473, 474, 485
Brown, Walter C 480
Eberth, David E 473, 474, 485
Etheridge, Richard 483
Evenhuis, Neal L 469
Garstka, William R 466
Hartel, Karsten E 487
Harris, Dennis 475
Kaufman, Leslie S 472
Losos, Jonathon B 484
Liem, Karel F 472
MiYATA, Kenneth 481
Moore, Karen E 476

Ono, R. Dana 470
Parker, Fred 480
Scott, Michele P 479
Shevill, William E 476
Stiassny, Melanie L. J 487
Turner, Susan 486
Williams, Ernest E 467, 471, 475, 477, 478, 482, 483

. COM?. ^°^
B R E.XI O R A
lis ISSN 0006 9698
Cambridge, Mass. 30 December 1981 Number 464
THE ORIGIN OF THE CROCODILOID TARSI

AND THE INTERRELATIONSHIPS OF
THECODONTIAN ARCHOSAURS
Donald Brinkman'
Abstract. The tarsus of the proterosuchian Chasmatosaurus represents the

primitive archosaur tarsus. This kind of tarsus is also present in rhynchosaurids,
triiophosaurids, prolacertids, and Protorosaums, and suggests

that these reptiles are
members of a single radiation. Two distinct kinds of crocodiloid tarsi are present in
thecodonts, a crocodile-normal tarsus and a crocodile-reversed tarsus. The crocodile-
reversed tarsus could have originated from the crocodile-normal tarsus, but the
reverse relationship is not plausible. Gracilisuchus. the only "ornithosuchid" with a
crocodile-normal tarsus, shows features of the skull that are not consistent with its
placement in the Ornithosuchidae. Euparkeria. on the basis of both cranial
and
but could not be
postcranial characters, is a plausible ornithosuchid ancestor
ancestral to a pseudosuchian with a crocodile-normal tarsus. The tarsus of
Erythosuchus neither contradicts nor supports a relationship between Erythrosuchus

and rauisuchids.
INTRODUCTION
In recent years, been recognized that a number of
it has
structurally distinct kinds of tarsi are present in archosaurs,
and an
understanding of the evolution of this structural complex is
of the
necessary for an understanding of the interrelationships
group. In the tarsus of crocodiles and typical pseudosuchians, the
ankle joint passes between the astragalus and calcaneum, the
astragalus being locked to the tibia and the calcaneum integrated
with the In dinosaurs,
pes. the ankle joint passes distal to the
astragalus and calcaneum. Krebs (1963, 1973) argued that this
Museum of Comparative Zoology, Harvard University, Cambridge, Massachsuetts

02138.
2 BREVIORA No. 464
difference precludes derivation of dinosaurs from any known

pseudosuchian. Recently, two additional kinds of tarsi have been
recognized in thecodonts. Proterosuchian thecodonts of the family
Proterosuchidae have a distinct tarsus that is in many ways
primitive (Cruickshank, 1972; Carroll, 1976). Bonaparte (1971)
recognized that two distinct kinds of crocodilelike tarsi are present
in pseudosuchians, one like that of crocodiles in which the
astragalus has a peg that fits in a socket on the calcaneum, and one
seen in advanced ornithosuchids in which the calcaneum has a
process that fits in a socket on the astragalus. Chatterjee (1978)
termed these the crocodile-normal and crocodile-reversed tarsus
respectively. Also, two pseudoschians with mesotarsal ankle joints
have been described (Romer, 1971). One of these, Lagerpeton, has a
fully developed mesotarsal joint. The second, Lagosuchus, retains a
posteriorly directed calcaneal tuber and a complex articulation
between the astragalus and calcaneum (Bonaparte, 1975a).
The evolution of these tarsal patterns was recently discussed by
Cruickshank (1979). Cruickshank showed that the proterosuchian
tarsus is an excellent structural ancestor to the crocodile-normal
tarsus and argued that the two kinds of crocodile tarsi can be used
to separate pseudosuchians into two groups. Based on this,
Cruickshank suggested that Gracilisuchus, which has a crocodile-
normal tarsus, be removed from the Ornithosuchidae, all other
members of which have a crocodile-reversed tarsus. However, the
origin of the crocodile-reversed tarsus remains unknown. If the
crocodile-normal tarsus was ancestral to the crocodile-reversed
tarsus, then a crocodile-normal tarsus could have been present in
primitive ornithosuchids, and the presence of a crocodile-normal
tarsus would not bar Gracilisuchus from the Ornithosuchidae.
Thus, in order to use the structure of the tarsus as a basis for
interpreting the interrelationships of archosaurs, it is necessary to
obtain a more precise understanding of both the origin of the
crocodile-reversed tarsus and the phylogenetic position of Gracili-
suchus.
THE CROCODILE-NORMAL TARSUS

In extant crocodiles, five elements are present in the tarsus: the
astragalus, the calcaneum, and the second to fourth distal tarsals.

I98I THE ORIGIN OF THE CROCODILOID TARSI
The astragalus (Fig. IC-D) supports the tibia and contacts the fibula
by articular surfaces that almost completely cover the proximal
surface of the bone. Anteriorly, the astragalus has a strongly covex
surface that articulates with the proximal end of the first two
metatarsals and the medial surface of the second and third distal
tarsals. Above this, the anterior face of the astragalus is formed by a
concave area covered by finished bone. Laterally, a distinctive
articular surface for the calcaneum is present. This is divisible into
two separate areas. The ventral area has the shape of a portion of a
fib art surf
wheel-sfiaped
art surf
wheel -shaped
arl surf
bond-shaped
\ art surf
med flange
5 mm
Figure 1. The right astragalus and calcaneum of Caiman sclerops. A) calcaneum

in proximal view; B) calcaneum in medial view; C) astragalus in dorsal view; D)
astragalus in medial view.
Ahhreviaiiom: band-shaped art surf, band-shaped articular surface; cai tub,
calcaneal tuber; cone-shaped cone-shaped articular surface; fib art surf,

art surf,
fibular articular surface; med flange, medial flange; met art surf, metatarsal
articular surface; tib art surf, tibial articular surface; wheel-shaped art surf,
wheel-shaped articular surface; ast art surf, astragalar articular surface.

4 BREVIORA No. 464
cone, its apex forming the tip of the laterally directed peg. Dorsal to
this, a notch edge of the astragalus leads to a band-
in the lateral
shaped articular surfce. These two areas meet along a ridge that
terminates on the tip of the lateral peg.
The calcaneum (Fig. A-B) has three characteristic areas: a-dorsal
1
area that has the form of a portion of a wheel, a medially directed

flange that underlies the astragalus, and a posteriorly directed tuber.
The medial half of the wheel-shaped articular surface fits in the
notch on the lateral edge of the astragalus and is overlapped by the
astragalus. The lateral half supports the fibula. The fibular and
astragalar surfaces are differentiated by a slight change in the
curvature of the articular surface. The medially directed flange
articulates behind the cone-shaped articular surface of the astra-
galus. The calcaneal tuber extends across the full width of the bone.
The distalend of the tuber is expanded and has a vertical groove in
which lie tendons of the long pedal flexors. Anterodistally, the
calcaneum as a flat articular surface that abuts the fourth distal
tarsal.
A crocodile-normal tarsus is present in the Rauisuchidae (Krebs,

1965, 1973; Sill, 1974), the Aetosauridae (Sawin, 1947; Walker,
1961; Bonaparte, 1971; and Gracilisuchus Bonaparte,
Sill, 1974),
1975b). A crocodile-normal calcaneum from the uppermost Lower
or lowermost Middle Triassic was figured by Young (1964, Fig. 60)
and attributed to Wangisuchus.
The astragalus in these pseudosuchians, where known, differs
from that of crocodiles in having more extensive development of
finished bone on its anterior face and in the proportions of the
articular surfaces, the metatarsal articular surface being narrower
mediolaterally in most genera, as in Gracilisuchus (Fig. 4B). The

proportions of the calcaneum also show some variation, the
calcaneal tuber of aetosaurs being considerably broader than in
crocodiles and other pseudosuchians (Sawin, 1947). Despite this
variation, the structure of the joint between the astragalus and
calcaneum is like that of crocodiles.
THE TARSUS OF CHASMATOSAURUS

The tarsus of the early proterosuchid Chasmatosaurus^ (Fig. 2) is
primitive in the presence of a separate astragalus and centrale, and
1981 THE ORIGIN OF 1 HE CROCODILOID TARSI
foramen between the astragalus and calcaneum.

the retention of a
However, comparison with an eosuchian tarsus, such as that of a
tangasaurid (Fig. 3), demonstrates that a number of derived features
are present. The articular surface between the astragalus and
calcaneum in eosuchians is flat and forms a straight line when the
tarsus is seen in dorsal view. In Chasmatosaurus, the portion of the
articular surface proximal to the perforating foramen is inclined
relative to the distal portion, with the astragalus overlying the
calcaneum. The
surface between the astragalus and
articular
calcaneum complex concave-convex
is a joint. The portion of the
articular surface distal to the perforating foramen is a ball and
socket joint, with the socket on the calcaneum. Proximal to the
perforating foramen, a concave-convex joint is also present, but the
concavity is on the astragalus. The proximal edge of the calcaneum
is not preserved in Chasmatosaurus vanhoepeni. but in the
cm
cen
Figure 2. The tarsus of Chasmatosaurus vanhoepeni. Left calcaneum, astragalus,

and centrale in A) ventral. B) dorsal, and C) proximal views. Drawing based on cast
of NM C30I6, Nasionale Museum, Bloemfontein.
Abbreviations: ast, astragalus; cal, calcaneum; cen, centrale.
Following Charig and Sues ( 976), Proterosuchus is considered a nomen dubium,

'
1
and the remaining proterosuchid material from South Africa is referred to the genus
Chasmatosaurus.
BREVIORA No. 464
Figure 3. The left tarsus of Hovasaurus. a tangasaurid eosuchian, in dorsal view.

Drawn from cast of MNHN 1925-5-61, National Museum of Natural History, Paris.
Abbreviations: ast, astragalus; cal. calcaneum; cen, centrale.

1981 THE ORIGIN OF THE CROCODILOID TARSI 7
calcaneum of Chasmatosaurus yuani illustrated by Young (1936,
Fig. 9D) the convexity of the calcaneal surface is seen to continue

laterally to form a dorsoventrally convex fibular articular surface,
as it does in the early rhynchosaur Noteosuchus (Fig. 6A).
The calcaneum has been modified from the primitive platelike
condition by the development of a laterally extending tuber. The
distal edge is thin, and a thickened medial buttress extends
transversely across bone to the expanded cartilage-covered

the
lateral edge.Judging from Young's illustration of the calcaneum of
Chasmatosaurus yuani. the proximal edge was covered by unfin-
ished bone, as it is in the early rhynchosaur Noteosuchus (Fig. 6A).
The centrale has shifted its position so that it is now located
laterally, rather than distally, to the astragalus. It may aid in
support of the tibia.
Cruickshank (1972), based on the similarity of the tarsus of

Chasmatosaurus and early rhynchosaurs and the presence in both of
a downturned premaxilla, suggested that Chasmatosaurus was a
carnivorous rhynchocephalian. If the proterosuchian tarsus was
restricted to these animals, it would suggest that Chasmatosaurus is
phylogenetically a rhynchosaur and thus would bring into question

the validity of using the proterosuchian tarsus as the primitive
archosaur pattern. However, a number of additional groups of
diapsid reptiles have a tarsus that, in so far as comparison is
possible, is like that of Chasmatosaurus. The tarsus of Prolacerta
(Gow, 1975, Fig. 33) shows all the advanced features seen in
Chasmatosaurus and early rhynchosaurs. As would be expected, the
aquatic members of the Prolacertiformes show a decrease in the
ossification of the tarsus. Despite this, mature specimens of
Tanystropheus (Wild, Fig. 75) and Macrocnemus (Peyer,
1973;
1937; PI. 56, Fig. 2) show
the diagnostic features of a laterally
directed dorsoventrally compressed tuber on the calcaneum, and a
complex concave-convex articulation between the astragalus and
calcaneum. A proterosuchian tarsus is also seen in Protorosaurus
from the Permian of Europe (von Meyer, 1856, PI. 9), and
Trilophosaurus from the Triassic of Texas (Fig. 4).
At first glance, this assemblage of reptiles seems to be an
unnatural one, bringing together animals with markedly different
adaptations and skull configurations. However, a more detailed
consideration shows that this assemblage is not as artificial as first
BREVIORA No. 464
Figure 4. The right astragalus and calcaneum of Trilophosaiirus. Calcaneum in
A) ventral, B) dorsal, C) proximal, and D) distal views; astragalus in E) ventral, F),

medial, and G) dorsal views. Calcaneum: specimen TMM
31025-258; astragalus:
specimen TMM 31025-259, Texas Memorial Museum.
appears. Prolacerta and Protorosaurus have long been recognized

as being closely related (Camp, 1945; Watson, 1958). Romer (1956)
denied the presence of such a relationship, choosing to place
Protorosaurus in the Euryapsida because of the presumed presence
of a solid cheek. However, as noted by Chatterjee (1980), much
uncertainty about the structure of the skull oi Protorosaurus exists.
Details of the skull and postcranial skeleton that are known for
certain are similar to Prolacerta, and Chatterjee unites these genera
in the suborder Prolacertiformes. Gow (1975) concluded on the
basis of evidence from the skull of Prolacerta that the Prolacerti-
formes are related to archosaurs.
The remaining groups in this assemblage, rhynchosaurids and
trilophosaurids, have highly specialized skulls that could be derived
from any primitive diapsid, so the skull neither supports nor negates
a relationship between these groups and the archosaur-protoro-
saurid-prolacertid group. Postcranially, both trilophosaurids and

rhynchosaurids are less specialized, and a similarity in the structure
of their postcranial skeletons and the postcranial skeleton of some
members of the archosaur-protorosaurid-prolacertid group has
been noted (Gregory, 1945; Carroll, 1976), although it is not certain
whether these represent derived features or primitive features
widespread in diapsid reptiles.
Thus, there seems to be no need to hypothesize a multiple origin
of the proterosuchian tarsus. If rhynchosaurs, prolacertiformes,
prolacertids, trilophosaurids, and archosaurs
are a natural group,
the proterosuchian tarsus could have originated only once. One
implication of this is that the proterosuchian tarsus is the primitive

archosaur tarsus and is the ultimate structural ancestor of the
various kinds of tarsi seen in advanced archosaurs. In some cases,
an intermediate structural complex may have been present, but, as
shown by Cruickshank (1979), the proterosuchian tarsus was
probably the direct structural antecedent of the crocodile-normal
tarsus. It is useful to identify the structural changes that would have
occurred during this transition before considering the structure and
origin of the crocodile-reversed tarsus.
ORIGIN OF THE CROCODILE-NORMAL TARSUS

As recognized by Cruickshank (1979), the astragalus-centrale unit
of the proterosuchian tarsus is directly comparable to the astragalus
of the crocodile-normal tarsus (Fig. 5). The area distal to the
perforating foramen is homologous to the cone-shaped articular

surface of the crocodile-normal tarsus, but differs in being smaller
and less strongly curved. The area proximal to the perforating
foramen is homologous to the dorsal half of the notch on the lateral
edge of the crocodile-normal astragalus, the main difference being

that in the proterosuchian tarsus this surface is separated from the
distal surfaceby finished bone.
The calcaneum of the crocodile-normal and proterosuchian tarsi
(Fig. 6) differs in the orientation of the calcaneal tuber; in the
proterosuchian tarsus, this is directed laterally, while in the

crocodile-normal tarsus, this is directed more posteriorly. If the
calcaneal tuber of the proterosuchian tarsus were oriented so that it
extended posteriorly, the articular surface for the fibula and
astragalus would be oriented along the long axis of the bone rather
10 BREVIORA No. 464
1 cm
1 cm
Figure 5. The left astragalus of the proterosuchian and crocodile-normal tarsus

in A) astragalus and centrale of Noteosuchus: B) astragalus of
anterior view.
Gracilisuchus. Not drawn to scale. Noteosuchus based on cast of Albany Museum
3591. Grasilisuchus based on cast of specimen in collection of Paleontologia
Vertebrados de la Fundacion M. Lillo.
n
cal tub
3
n
3
fib art surf

ast art surf
ast art surf
art surf
Figure 6. The left calcaneum of the proterosuchian and crocodile-normal tarsus.
A) calcaneum of Noteosuchus: B) calcaneum of GraciHsuchus.
Abbreviations: ast art surf, astragalar articular surface; cal tub, calcaneal
tuber; fib art surf, fibular articular surface.
1981 THE ORIGIN OF THE CROCODILOID TARSI 1 1
than across it. A simple enlargement of the articular surface,

together with the extension of the proximal portion of the astragalar
articular surface onto the medial edge of the perforating foramen,
would form the wheel-shaped articular surface of the crocodile-
normal calcaneum. An enlargement of the ventral half of the
articular surface for the astragalus would form the medial flange of
the crocodile-normal calcaneum.
Thus structurally, the proterosuchian tarsus is an excellent
ancestor of the crocodile-normal tarsus. The mechanical changes
involved in this transition were probably minor, since, as noted by
Thulborn (1980), the joint between the astragalus and calcaneum
was probably movable in the proterosuchian tarsus.
THE CROCODILE-REVERSED TARSUS

The crocodile-reversed tarsus is best known in the ornithosuchid
Riojasuchus (Bonaparte, 1971, 1975b). Dorsally, the calcaneum
(Fig. 7B) has an articular surface that is convex both mediolaterally
and proximodistally. This surface supports the fibula along its
lateral edge and the astragalus along its medial edge. These are
exactly the relationships of the wheel-shaped articular surface of the

crocodile-normal tarsus (Fig. 8A-B). The major difference is that
the medial edge of this area is hypertrophied in Riojasuchus to form
a medially directed process. This process is functionally equivalent
to the ventral flange of the crocodile-normal calcaneum in that it
underlies the astragalus (Fig. 8C-D). Consequently, it is not
surprising that the ventral flange or an homologous area is not

present in the crocodile-reversed calcaneum. The absence of the
ventral flange is associated with a reduction in the width of the
calcaneal tuber; in the crocodile-reversed tarsus, the calcaneal tuber
does not extend the full width of the bone. In addition, the tuber of
the crocodile-reversed tarsus is distinctive in that its distal end
curves medially and is without a groove for the tendon of the

gastrocnemial muscles.
The differences in structure of the astragalus of the crocodile-
normal and crocodile-reversed tarsus correspond to the differences
in structure of the calcaneum: the hypertrophy of the medial edge of
the wheel-shaped articular surface is associated with the elongation
of the overlying portion of the astragalus, and the loss of the ventral
flange is associated with the loss of the cone-shaped articular
surface.
12 BREVIORA No. 464
tib art surf.
fib art cal tub

surf
met art
Figure 7. The left astragalus and calcaneum of the crocodile-reversed tarsus. A)

Astragalus and B) calcaneum of Riojasuchus. Drawn from cast of PVL 3827,
Paleontologia Vertebrados de la Fundacion M. Lillo.
Abbreviations: ast art surf, astragalar articular surface; cal tub, calcaneal
tuber; fib art surf, fibular articular surface; met art surf, metatarsal articular
surface; tib art surf, tibial articular surface.
From this comparison, it can be seen that the crocodile-normal

tarsus is a plausible ancestor of the crocodile-reversed tarsus. The
major changes involved in such a transition would be the medial
elongation of the wheel-shaped articular surface of the calcaneum
and the loss of the ventral flange. Derivation of the crocodile-
reversed tarsus directly from the primitive archosaur tarsus is also
possible. However, derivation of the crocodile-normal tarsus from
the crocodile-reversed tarsus can be discounted as being improbable
since it would involve the redevelopment of the ventral flange, a
structure that is present in the crocodile-normal tarsus and the
primitive archosaur tarsus but absent in the crocodile-reversed

tarsus.
Given this relationship of the crocodile-normal and crocodile-
reversed tarsi, the structure of the tarsus cannot be used to exclude
Gracilisuchus from the Ornithosuchidae. Rather, the systematic
position of Gracilisuchus has implications for the evolution of the
tarsus. If Gracilisuchus is a true ornithosuchid, the crocodile-
normal tarsus must have given rise to the crocodile-reversed tarsus,
and the structure of the tarsus has little real phylogenetic
significance. If however, Gracilisuchus is not an ornithosuchid, the
crocodile-reversed tarsus may have originated independently from
Figure 8. The left calcaneum of the crocodile-normal and crocodile-reversed

tarsi. A) crocodile-normal calcaneum of Gracilisuchus
in proximal view; B)
crocodile-reversed calcaneum of Riojasuchus proximal view; C) section through

in
an articulated crocodile-normal astragalus and calcaneum; D) section through an

articulated crocodile-reversed astragalus and calcaneum. Not drawn to scale.
the proterosuchian tarsus, and the presence of the crocodile-

reversed tarsus can be used as the defining feature of some
taxonomic group. Thus it is necessary to reconsider the relation-
ships of Gracilisuchus.
THE RELATIONSHIPS OF GRACILISUCHUS

The skull of Gracilisuchus was reconstructed by Romer(1972) on
the basis of MCZ 41 17, a complete, three dimensional skull. One of
14 BREVIORA No. 464
the unusual features seen in this skull is a small lower

temporal
opening. However, other material, particularly MCZ 4116 and
MCZ 41 18, show that MCZ 41 17 has been slightly crushed and the
quadratojugal and squamosal have been displaced. In MCZ 4116,
the preorbital bar is slender and the antorbital opening is larger than
in MCZ 41 17 (Fig. 9A). In MCZ 41 18, the postorbital bar is tall and
slender, the quadratojugal and squamosal are separated from the

postorbital, and the lower temporal opening is large and rectangular
(Fig 9B). Based on these skulls, the arrangement of the temporal
region and the height of the face in the reconstruction of the skull of
Gracilisuchus is modified (Fig. 93).
Gracilisuchus differs from the advanced ornithosuchids as
defined by Bonaparte (1975b) (Fig. 10) in the following features:
1) The antorbital fenestra is rectangular in Gracilisuchus and is
triangular in the advanced ornithosuchids.
2) The ventral border of the orbit is round in Gracilisuchus, and a
distinct antorbital ramus of the jugal
not present. In the
is
advanced ornithosuchids, a distinct preorbital ramus of the

jugal is present; this is close to the postorbital ramus at its base,
so the ventral margin of the orbit is pointed.
3) The quadratojugal of Gracilisuchus is a tall, slender element
that extends nearly the full height of the lower temporal
opening. In the advanced ornithosuchids, the quadratojugal is
broader and is limted to the ventral half of the lower temporal
opening.
4) The upper tooth row is complete, and all the teeth of the lower
jaw fit inside the upper teeth row in Gracilisuchus. In advanced
ornithosuchids, a gap is present between the anterior tooth of
the maxilla and the posterior tooth of the premaxilla, with the
anterior one or two dentary teeth passing lateral to the maxilla
in thisgap.
5) In Gracilisuchus, the lower temporal fenestra is rectangular,
and no anterior inflection of the quadratojugal and squamosal
is present. In the advanced ornithosuchids, a large anterior
inflection of the quadratojugal and squamosal results in the
presence of an L-shaped lower temporal fenestra.
6) In Gracilisuchus, the squamosal has a peculiar, posteriorly
concave flange on its dorsal end. No such flange is present in
the advanced ornithosuchids.
Figure 9. The skull of Gracilisuchus. A) specimen drawing of MCZ 4116; B)

specimen drawing of MCZ 4118; C) reconstruction of skull.
16 BREVIORA No. 464
7) In Gracilisiichus. the posterior end of the dentary extends

dorsal to the mandibular fenestra. In the advanced ornitho-
suchids, the posterior end of the dentary is forked, with one
branch extending dorsal and one branch extending ventral to
the lateral mandibular fenestra.
8) In Gracilisuchus, the splenial forms the ventral margin of the
jaw along the posterior half of the dentary. In the advanced
ornithosuchids, the splenial is restricted to the inner surface of
the jaw.
9) In Gracilisuchus, the cervical vertebrae are not keeled. In the
advanced ornithosuchids, the cervical vertebrae, where known
(Riojasuchus and Ornithosuchus), are keeled.
10) In Gracilisuchus. there are two pairs of dermal scutes per
vertebra in the cervical region of the vertebral column. In the
advanced ornithosuchids, where known (Ornithosuchus and
Riojasuchus), there is one pair of scutes per vertebra.
This list of differences between Gracilisuchus and the advanced
ornithosuchids shows that the advanced ornithosuchids are more
similar to each other than to Gracilisuchus. But these features by
themselves do not demonstrate that Gracilisuchus is not an
ornithosuchid; Gracilisuchus occurs much earlier in time than the
advanced ornithosuchids, so the differences may represent succes-
sive grades of evolution in a single radiation. In order to be used
phylogenetically, it is necessary to determine which of the character-

states represent derived features. Euparkeria (Fig. lOD), the oldest
known pseudosuchian, had traditionally been considered to be the
structural ancestor of later pseudosuchians and thus can be used as
the outgroup in determining which character-states are primitive or
advanced.
Features that are shared by Gracilisuchus and Euparkeria, and
thus can be considered primitive, are the presence of a complete
upper tooth row with all the dentary teeth fitting inside the upper
tooth row, the shape of the lower temporal fenestra, and the shape
of the antorbital fenestra (features 1, 4, and 5 in the above list). In all
other features, Euparkeria is like the advanced ornithosuchids and
unlike Gracilisuchus: the jugal has a well-developed antorbital
process, and the base of this is near the postorbital process; the
squamosal without the peculiar posteriorly concave flange seen in
is
Gracilisuchus; the posterior end of the dentary is forked, with a

branch above and a branch below the lateral mandibular fenestra;
Figure 10. The skulls of the advanced ornithosuchids and Euparkeria. A)

I'enaticosuchus; B) Ornithusuclms: C) Riojasuchus; D) Euparkeria. From Bona-
parte, 1975.
the splenial is restricted to the internal surface of the lower jaw; the
cervical vertebrae are keeled; and one pair of dermal scutes is
present per vertebral segment. For these features, the character-state

present in the advanced ornithosuchids must be considered primi-
tive, and the Gracilisuchus condition derived.
Thus, if an ornithosuchid, Gracilisuchus isderived from the
ornithosuchid pattern in a way different from the advanced
ornithosuchids. Alternatively, Gracilisuchus may be a member of a
radiation distinct from that of ornithosuchids. This latter possibility
is suggested by the presence of some of the derived features of
Gracilisuchus in Sphenosuchus, Pseudohesperosuchus. and Lewi-
suchus, pseudosuchians that are thought to be unrelated to
ornithosuchids (Romer, 1972). These features include the tall.
18 BREVIORA No. 464
slender quadratojugal, the posteriorflange on the squamosal

(known in Pseudohesperosuchus and Sphenosuchus), and the
presence of unkeeled cervical vertebrae (known in Lewisuchus).
THE TARSUS OF EVPARKERIA

The similarity between ornithosuchids and Euparkeria raises the
possibility that Euparkeria is closely related to the Ornithosuchidae,

rather than being a generalized primitive pseudosuchian. Such a
relationship is supported by the structure of the tarsus. The

astragalusand calcaneum of Euparkeria were illustrated by
Bonaparte (1975) and Cruickshank (1979). As noted by these
authors, these elements are directly comparable to those of the
crocodile-reversed tarsus (Fig. 11). The dorsal portion of the
articular surface of the calcaneum is divisible into two areas, a
narrow area along the edge of the bone that would have
lateral
articulated with the fibula, and a medial area that would have
articulated with the astragalus. The astragalar area is in the shape of
a portion of a cone with its apex directed medially. No area
comparable to the ventral flange of the crocodile-normal tarsus is
present. The calcaneal tuber

directed posteriorly and is narrow.
is
Thus the calcaneum of Euparkeria is similar to the crocodile-

reversed tarsus in all the features by which the crocodile-normal and
crocodile-reversed tarsi differ. Of special importance is the absence
of an area homologous to the ventral flange of the astragalus of the
crocodile-normal tarsus. As was argued above, the absence of this
area prevents derivation of a crocodile-normal tarsus from a
crocodile-reversed tarsus without the redevelopment of a lost
structure. On this basis, Euparkeria can be considered to be more
closely related to the ornithosuchids than to any pseudosuchian
with a crocodile-normal tarsus.
THE TARSUS OF ERYTH ROSUCHUS

An additional problem to be considered is the relationship of
Erythrosuchus. (1974) showed that a striking similarity exists
Sill
between the skull of Erythrosuchus and rauisuchids, so that a

relationship between these two groups is probable. Cruickshank
(1978, 1979), however, felt was
that the tarsus of Erythrosuchus
most similar to that of Euparkeria, and placed Erythrosuchus with
Euparkeria and the ornithosuchids in his Ornithosuchoidea.
198i THE ORIGIN OF THE CROCODILOID TARSI 19
cd tub
tib art
surf B
fib art surf
Figure 1 1. The
right astragalus and calcaneum of Euparkeria. A) astragalus; B)
calcaneum; and C) astragalus and calcaneum in articulation. A and B from
Bonaparte (1975), C based on drawings of South African Museum specimen SAM
6049 made by J. Bonaparte.
Abbreviations: ast art surf, astragalaj articular surface; cal tub, calcaneal
tuber; fib art surf, fibular articular surface; tib art surf, tibial articular surface.
A general similarity between the tarsus of Erythrosuchus and

Euparkeria is present, but this can be attributed to the poor
ossification of the bones. More fundamental is the similarity
between the tarsus of Erythrosuchus and Chasmaiosaurus. A
comparison of the calcaneum of these two animals (Fig. 12) shows
that the calcaneal tuber extends laterally, has thin proximal and
distal edges, and has a thickened medial buttress that extends across
the tuber to an expanded cartilage-covered lateral edge. The poor
ossification of the
astragalus the
of Erythrosuchus obscured
structure of the articulation between the astragalus and calcaneum.
20 BREVIORA No. 464
Figure 12. The left calcaneum of Erylhrosuchus in A) ventral, and B) distal

views; and Chasniatosaurus in C) ventral, and D) distal views. Erythrosuihus
drawing based on cast of Bernard Price Institute F. 2069 M 405, Chasniatosaurus
drawing based on cast of NM C 3016, Nasionale Museum. Bloemfontein.
although Cruickshank (1978) identifies a perforating foramen, a

primitive feature that is also present in Chasniatosaurus.
Thus the tarsus of Erythrosuchus is best considered a poorly
ossified proterosuchian tarsus. A reduction of ossification is
commonly seen in
aquatic and probably reflects an
animals,
adaptation for an aquatic habitat, rather than the development of a
mechanically distinct structural complex. A fully terrestrial erythro-
suchid would, therefore, be expected to have a tarsus like that of
Chasmatosaurus. Since this kind of tarsus is the structural ancestor
of the crocodile-normal tarsus, the structure of the erythrosuchid
tarsus neither supports nor negates a relationship between erythro-
suchids and rauisuchids. If a relationship between erythrosuchids
and rauisuchids accepted on the basis of similarities in the skull, it

is
is necessary to assume that the crocodile-normal tarsus originated
after the origin of this radiation of thecodonts, and therefore the

tarsus cannot be used as the defining feature of the group.
SUMMARY
A summary of the relationships suggested above may be made in
the form of a phylogenetic diagram (Fig. 13). According to this
phylogeny. many of the higher taxa of diapsid reptiles are artificial

assemblages, and major modifications of reptilian classification are
necessary to accurately reflect reptilian relationships, it would,
however, be useful to test these relationships through examination
of other structural complexes before making the necessary changes.
Stogonolepdidae
Ornithosuchidae Rauisuchidae / Grocilisuchus
Erythrosuchidae
Lacertilia
Sauropterygia
Trilophosaundae
Figure 13. The interrelationships of thecodonts.
ACKNOWLEDGMENTS
I would like to thank Dr. R. L. Carroll, at whose suggestion this
project was undertaken. For reading the manuscript and making
many useful suggestions, thank Hans Dieter Sues, Museum of
I
Comparative Zoology, Harvard University; Michael Parrish, Uni-

versity of Chicago; Dr. Sankar Chatterjee, Texas Tech University;
and Robert Long and Dr. S. Welles, University of California,
22 BREVIORA No. 464
Berkeley. I also thank Lillian Maloney, who proofread the

manuscript. Figure 9 was drawn by Linda Krause. Publication costs
of this study were covered in part by a grant from the Wetmore
Colles Fund.
LITERATURE CITED
Bonaparte, J. F. 1971. Los tetrapodos del sector superior de la formation Los
Colorados, La Rioja, Argentina. Opera Lilloana, 22: 1-183.
1975a. Nuevos materials de Lagosuchus talampayensis Romer (Theco-
dontia-Pseudosuchia) y su significade en el origen de los Saurischia. Acta Geol.
Lilloana, 13: 5-87.
1975b. The family Ornithosuchidae (Archosauria: Thecodontia). Col-
loques int. Cent. Natn. Rech. Scient. No. 218, pp. 485-506.
Camp, C. L. 1945. Prolacena and the protorosaurian reptiles. Am. J. Sci., 243:
17-32, 84-101.
Carroll, R. L. 1976. Noteosuchus. the oldest known rhynchosaur. Ann. S. Afr.
Mus., 72: 37-57.

Charig, a. J., AND H. -D. Sues. 1976. Proterosuchia, pp. 1 1-39. In A. J. Charig,
B. Krebs, H. -D. Sues,and F. Westphal, Handbuch der Palaoherpetologie, 13,
Stuttgart and Portland, Gustav Fischer, 137 pp.
Chatterjee, S. 1978. A primitive parasuchid (phytosaur) reptile from the Upper-
Triassic Maleri Formation of India. Paleontology, 21: 83-127.
1980. Malerisaurus, a new eosuchian reptile from the late Triassic of
India. Phil. Trans. Roy. Soc. Lond. (B). 291: 163 200.
Cruickshank, a. R. I. 1972. The proterosuchian thecodonts, 89-119. In K. A.

Joysey, and T. S. Kemp (eds.). Studies in Vertebrate Evolution: Essays Pre-
sented to Dr. F. R. Parrington F. R. S. Edinburgh, Oliver and Boyd, 284 pp.
1978. The pes of Erylhrosuchus africanus Broom, J. Linnean Soc.
(Zool.), 62: 161-178.
1979. The ankle joint in some early archosaurs. S. Afr. J. Sci., 75:
168-178.
Gow, C. E. 1975. The morphology and relationships of Youngina capensis
Broom and Prolacerta broomi Parrington. Paleontol. Afr., 18: 89-131.
Gregory, J. T. 1945. Osteology and relationships of rr/Vo/^/ioâwrui. U. of Texas

Publ. No. 4401, pp. 273-359.
Krebs, B. 1963. Bau und Function des Tarsus eines Pseudosuchiers aus der Trias
des Monte San Giorgio (Kanton Tessin, Schweiz.). Palaontol. Z., 37: 88 95.
1965. Tictinosuchus ferox nov. gen., nov. sp. Ein neuer Pseudosuchier
aus der Trias des Monte San Giogio. Schweiz. Palaontol. Abh., 81: 140. 1
1973. Der Tarsus von Rauisuchus (Pseudosuchia, Mittel Trias). Mitt.

Bayer. Staatsamml. Palaontol. Hist. Geol., 13: 95-101.
Peyer, B. 1937. Die Triasfauna der Tessiner Kalkalpen, XII Macrocnemus
bassanii Nopsca. Schweiz. Palaontol. Abh., 59: 1-140.
ROMER, A. S. 1956. The Osteology of the Reptiles. Chicago, University of Chica-
go Press. 772 pp.

1971. The Chaiiares (Argentina) Triassic reptile Fauna X. Two new but
incompletely known long-limbed pseudosuchians. Mus. Comp. Zool., Breviora
No. 378. pp. 1-10.
1972. The Chanares (Argentina Triassic Reptile Fauna, XIII. An early
ornithosuchid pseudosuchian. Gracilisuchus slipaniaicorum. gen. et sp. nov.
Mus. Comp. Zool.. Breviora No. 389, pp. 1-24.
S.AWiN, H. J. 1947. The pseudosuchian reptile Typothorax meadei. J. Paleontol.,
21: 201-238.
Sill. W. D. 1974. The anatomy of Saurosuchus galilei and the relationships of the
rauisuchid thecodonts. Bull. Mus. Comp. Zool.. 146: 317-362.
Thulborn, R. a. The ankle joints of archosaurs. Alcheringa, 4: 241-261.
1980.
VON Meyer. H. Zur Fauna der Vorwelt. Saurier aus dem Kupferschiefer
1856.
der Zechstein-Formation. Frankfut-am-main.
Walker. A. D. 1961. Triassic reptiles from the Elgin area; Stagonolepis,
Dasygnathus and their allies. Phil. Trans. Roy. Soc. Lond. (B), 244: 103-204.
1970. A revision of the Jurassic reptile Hallopus victor (Marsh), with
remarks on the classification of crocodiles. Phil. Trans. Roy. Soc. Lond. (B),
257: 323-372.
Watson, D. M. S, 1958. On Millerosaurus and the early history of the sauropsid
reptiles. Roy. Soc. Lond. (B). 240: 355-400.

Phil. Trans.
Wild, R. 1973. Die Triasfauna der Tessiner Kalkalpen XXHL Tanystropheus

longobardicus (Bassani) (Neue Ergebnisse). Schweiz. Palaontol. Abh.. 95:
,
1-162.
Young, C. C. 1936. On a new Chasmaiosaurus from Sinckiang. Bull. Geol. Soc.
China. 15: 291 320.

1964. The pseudosuchians in China. Palaeontol. Sinica. New Ser. C. 19:
1-205.
J*'^.
MU3. COMP. ZOOL
LfHRARY
MAR 1 8 1985
B R E te±?lO R A
Miiseiiiii ol^
Cu}ai])arative Zoology
us ISSN 0006 9698
Cambridge, Mass. 30 December 1981 Number 465
THE STRUCTURE AND RELATIONSHIPS

OF THE DROMASAURS (REPTILIA: THERAPSIDA)
Donald Brinkman'
Abstract. The dromasaurs are primitive anomodonts that are advanced over the
venjukoviamorph grade of evolution in the reduction of the postorbital and
zygomatic branches of the jugal, the reduction of the septomaxilla, and the
enlargement of the area on the dentary for the insertion of the external mandibular
muscle. Derived features that distinguish dromasaurs from dicynodonts are the lack
of canines, the presence of a tall, slender postorbital. and the rodlike lower temporal
bar. The three known dromasaurs form a structural series showing changes in the
proportions of the face, a loss of premaxillary teeth, changes in the curvature of the
humerus, and a loss of the ectepicondylar foramen.
INTRODUCTION
Frequently, the earliest representatives of higher taxa include
groups that are at a primitive grade of evolution but are, in some
features, highly specialized. Within the Therapsida, one such group
is the Dromasauria. The dromasaur to be recognized, Gale-
first
chirus, was initially considered to be more primitive than any other

therapsid known at that time (Broom, 1907). Later, two additional
genera, Galepus and Galeops, were recognized, with Galeops being

considered sufficiently distinct to be placed in its own family, the
Galeopsidae (Broom, 1910, 1912). The better understanding of the
dromasaur skull provided by these genera showed that they were, in
many respects, derived from the primitive therapsid condition, and
'Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts

02138.
2 BREVIORA No. 465
Broom (1914: 12) suggested that they were "an aberrant group of
small primitive therapsids."
While there has been little question of the reality of the group, its
relationships have been a matter of much uncertainty. Romer and
Price (1940) noted that many of the features that separated droma-
saurs from other therapsids were primitive for pelycosaurs of the
suborder Edaphosauria, implying that a close relationship existed
between the two groups. Later, in a review of therapsid classifica-
tion, Romer and Watson (1956) suggested that a relationship
between dromasaurs and dicynodonts was likely, and placed the
dromasauria within the Suborder Anomodontia as an infraorder
equal in rank to the Dicynodontia. The possibility that both of these
relationships are correct was raised by Olson (1962), who suggested
that the therapsids had a multiple origin, with the anomodonts
originating from within the Edaphosauria.

Uncertainty about the relationships of the dromasaurs is a result
of an incomplete understanding of the anatomy of the genera
comprising the group. Although much of the skeleton is preserved,
the original material consists of natural casts, so that Broom was
able to determine little more than the outlines of the bones.
Recognizing these problems, it was decided to reexamine the

specimens studied by Broom using the methods developed for the
study of specimens preserved as natural casts by Baird (1951, 1955)
and Carroll (1976), in which latex peels of the specimens provide a
positive image of the bones. Because it is the most completely
preserved of the dromasaurs, Galeops will be described first.
G A LEO PS
and Methods: Galeops is represented by a single
Materials
species,Galeops whaitsi, and by a single specimen, 5536, AMNH
that comes from the Tapinocephalus zone of South Africa (Kitch-
ing, 1977). It consists of the impressions of the skull and the front
half of the postcranial skeleton in a fine-grained sandstone. Both
part and counterpart are present. Latex peels of the specimen

provided a positive image of the skeleton, and in some cases, it was
possible to cast the two halves of the individual elements and join
them together to give three-dimensional replicas of the original
bone.
1981 THF DROMASAURS 3
Skull Roof: The skull of Galeops (Figs. 1-5) is small for a

therapsid, its length from snout to quadrate being 55 mm. The
orbits and e.xternal narial openings are large and face laterally. The
face and postorbital regions are short. Sclerotic rings are present.
The ventral margin of the postorbital region is deeply emarginated,
reducing the zygomatic bar to a slender rod. The posterior edge of

the upper temporal opening is folded backwards, providing a large
area for the temporal muscles. No such outfolding of the squamosal
is present ventral to the zygomatic bar. The occiput slopes forward
and grades into the skull roof. A
large pineal opening is present at
the level of the frontal-parietal suture ( Fig. 4). The teeth are reduced
in size and are restricted to the maxilla —
presumably a horny beak
would have been present anteriorly.
Much is formed by the maxilla. In
of the lateral surface of the face
bone is triangular with its apex directed dorsally. It
lateral view, this
is sculptured by numerous small
pits and grooves, with these being
most strongly developed anteriorly. The relationship of the maxilla
to the surrounding bones of the face is similar to that of dicyno-
donts: it and prefrontal poste-
articulates with the jugal, lacrimal,
riorly;the nasal dorsally; and the premaxilla midway below the

external narial opening. The contacts of the maxilla with the
prefrontal and nasal are strongly interdigitating sutures. The lacri-
mal appears to overlay the maxilla, with the suture line being
straight in external view. The details of the remaining contacts are
uncertain.
Ventrally, the maxilla has a well-developed palatal flange that
borders the internal narial opening (Fig. 5). Four sockets are present
in the and five in the right maxilla. These increase in size
left
posteriorly from the first to third, with the fourth being about equal
in size to the second in the case of the right maxilla. A distinct
canine not present.
is
A small portion of the lateral surface of the premaxilla is present

ventral to the external narial opening. This has a roughened surface
like that of the maxilla. Ventrally, the
paired premaxillae have large
palatal flanges that form a dicynodontlike secondary palate ( Fig. 5).
The ventral surfaces of these flanges are roughened by numerous
posteriorly projecting spicules of bone. A distinct suture separates
the two premaxillae from each other.
BREVIORA No. 465
1981 THE DROMASAURS 5
A small septomaxilla spans the premaxilla-maxilla suture (Fig.

1). It has no exposure on the lateral surface of the face. The
septomaxilla foramen is a small opening located between the
septomaxilla and maxilla.
The nasal has a large exposure on the lateral surface of the face
and forms the dorsal half of the posterior margin of the external
narial opening. The anterior three quarters of the bone has a
roughened pitted texture like that of the maxilla. Posterior to this,

the lateral surface of the nasal is smooth.
The
lacrimal, in lateral view, appears to be a small subcircular
bone between the maxilla, jugal, and prefrontal. The orbital
fitting
edge of the bone has an internally directed flange, although this is
incompletely preserved (Fig. 2).
The jugal is reduced to a slender rodlike bone forming the ventral
margin of the orbit and articulating with the postfrontal and
squamosal posteriorly and with the maxilla and lacrimal anteriorly.
No contribution to the formation of the postorbital or zygomatic
bar is made by the jugal. Anteriorly, a well-defined groove is present
on the lateral surface of the jugal. This may have been associated
with the horny beak.
The prefrontal is a triangular bone forming the anterodorsal edge
of the orbit. It contacts the lacrimal ventrally, the maxilla and nasal
anteriorly, and the frontal dorsally. All of these sutures are squa-
mous interdigitating contacts. The prefrontal is restricted to the
lateral surface of the face, the transition from the lateral to dorsal
surface occurring at the area of the prefrontal-frontal contact. The
lateral surface of the prefrontal is smooth.
The frontals are seen in ventral view. Each frontal contacts the
nasal and prefrontal anteriorly, the postfrontal and parietal poste-
riorly, and its fellow at the midline. No preparietal bone is present.
Figure 1.
Galeops whaitsi. Drawing based on latex peel taken from AMNH-
5536.
Abbreviations: an, angular; ar, articular; C, clavicle; Co, coracoid; d, dentary; f,

frontal;H, humerus; i, intermedium; IC, interclavicle; j, jugal; 1, lacrymal; m,
maxilla; n, nasal; p, parietal; pa, prearticular; PCo, procoracoid; pf, postfrontal; pm,
premaxilla; po, postorbital; prf, prefrontal; pro, prootic; pt, pterygoid; q, quadrate;
qj, quadratojugal; r, radiale; R, radius; S, scapula; sa, surangular; sm, septomaxilla;
sq, squamosal; u, ulnare; U, ulna.
BREVIORA No. 465
SO the frontals form the anterior border of the pineal foramen.

Laterally, they contribute to the margin of the orbit. Ventrally,
strong ridges form the border of the nasal tracts.
The postfrontal is a small, triangular bone wedged between the
frontal and postorbital. It has a smooth concave lateral surface.
The postorbital is a narrow bone separating the orbit from

tall,
the temporal opening. supported by the jugal and squamosal

It is
ventrally and articulates with the parietal and postfrontal dorsally.

It does not extend posteriorly along the parietal to meet the
squamosal as is the case in dicynodonts and primitive therapsids
generally.
The squamosal is a complex bone that is, in its general structure,
like that of dicynodonts. As in dicynodonts. the main body of the
bone forms the lateral portion of the occipital plate. The dorsal
portion of this area folded posteriorly so that the posterior rim of
is
the temporal fenestra overhangs the occiput. No such outfolding is

seen ventral to the zygomatic bar. A ventrally directed ramus
supports the quadrate and quadratojugal and forms the lateral
portion of the occipital plate. Laterally, an anteriorly directed
zygomatic ramus extends to the jugal. This is a narrow rodlike bone,
rather than being flat and beamlike as in dicynodonts. The zygoma-
tic ramus of the squamosal
overlaps the jugal, extending well
anterior to the posterior edge of the orbit.
The parietals are seen in ventral view ( Fig. 4). They have a sutural
contact with the frontals and postorbitals anteriorly and with the
postparietal posteriorly. Large tabulars are probably present as
well, but sutures cannot be clearly identified. Ventrally, each
parietal has a crescent-shaped ridge that served as the area of
attachment of the epipterygoid and braincase to the skull roof. The
Figure 2. Galeops whaitsi. Drawing based on latex peel taken from counterpart
slab of AMNH 5536. Skull and lower jaw bones are shown from medial surfaces.
Abbreviations: aa, atlas arch; ac, atlas centrum; an, angular; ar. articular; ax, axis;
C, clavicle; Co, coracoid; d, dentary; f, frontal; H, humerus; ic, mtercentrum; IC,

interclavicle; j, jugal; 1. lacrymal; m, maxilla; n, nasal; p. parietal; PCo, procoracoid;
pf. postfrontal; pm. premaxilla; prf, prefrontal; pro. prootic; q, quadrate; R, radius;
S, scapula; sp, splenial; sq. squamosal; si pit. stapedial pit; U, ulna.
BREVIORA No. 465
Figure 3. Galeops whaitsi. A) reconstruction of skull and lower jaw; B)

reconstruction of lower jaw, medial view; C) reconstruction of articular, dorsoposte-
rior view.
Abbreviations: see Figure 2.
parietals extend lateral to this ridge as a ventrally facing shelf that

would have provided an area for muscle attachment. This shelf
forms a portion of the lateral edge of the temporal fenestra.
Anteriorly, a cup-shaped depression is present just anterior and
lateral to the area of attachment of the braincase.
The quadratojugal is a small splintlike bone resting on the
posterolateral corner of the quadrate ventrally and fitting in a
groove on the anterior face of the squamosal dorsally. An oval
quadratojugal foramen lies between the quadrate and quadrato-
jugal.
The quadrate is a large bone with a dorsally directed process that
fits in a groove in the squamosal, an anteriorly directed flange that
1981 THE DROMASAURS
meets the pterygoid, and a ventral expansion that forms the

articular surface for the lower jaw. The articular surface is divided
into two condyles separated by a deep groove. The lateral condyle is
located slightly posterior to the inner condyle. A pit is present on the
internal surface of the quadrate that would have received the stapes,
although the stapes itself is not preserved.
Braincase: The braincase of Galeops is represented by the internal
surface of the dorsal half of the occiput (Fig. 4). the lateral surface
of the prootic (Fig. 1), and the ventral surface of the basisphenoid
(Fig. 5). No sutures can be seen in the occiput. A small posttemporal
fenestra is present, surrounded by the squamosal laterally and the
occipital plate medially. A groove on the lateral surface of the
prootic leads to the temporal fenestra. Two large paired internal
carotid foramina are present in the basisphenoid just medial and
Figure 4. Galeops whaitsi. Internal view of posterior surface of skull drawn

from latex peel of AMNH 5536.
Abbreviations: oc, occiput; p, parietal; par f, parietal foramen; pp. postparietal;

pro, prootic; pt, pterygoid; ptf, posttemporal fenestra; q, quadrate; qj. quadratojugal;
sq, squamosal; t, tabular.
10 BREVIORA No. 465
posterior to the basipterygoid articulation. Grooves in the basi-

sphenoid lead to these foramina from the posterior edge of the bone.
Posterior to the basipterygoid articulation, the lateral edge of the
basisphenoid extends ventrally as a large semicircular flange that
forms the ventral edge of the fenestra ovalis. This is located well
above the level of the stapedial pit on the quadrate, so the stapes
would have sloped ventrally, rather than being nearly horizontal as
in dicynodonts.
Palate: The
palate of Galeops (Fig. 5) is primitive in its general
structure; a complement of palatal bones is present, the
full
transverse flange of the pterygoid is present, and the interptery-

goidal vacuity is small. An advanced feature is the enlargement of
the palatal flanges of the prema.xillae to form a secondary palate.
The vomers, as in dicynodonts, form vertical plates, although in
Galeops they are separate ventrally. The palatines have the relation-
ships seen in the primitive anomodont Otsheria: each palatine
contacts the maxilla and ectopterygoid laterally, the vomers and
opposite palatine medially, and the pterygoid posteriorly. The
palatines are arched to roof the posterior portion of the internal
narial openings. A sharp ridge borders this vaulted area laterally,
suggesting that a soft palate was present. The ectopterygoid is a

small rectangular bone extending from the transverse flange of the
pterygoid to the maxilla. The bone has been displaced slightly, so it
isimpossible to determine if a lateral palatal foramen was present as
in dicynodonts (Cluver, 1975). The pterygoids are partially sepa-
rated by a small interpterygoid vacuity similar in size and propor-
tions to that of Otsheria. The two pterygoids contact one another
anterior to this vacuity but do not meet posteriorly, as is the case in
dicynodonts, including Eodicynodon, the earliest known dicyno-
dont (Barry, 1974). The basipterygoid articulation is unfused. The
lateral edge of the transverse flange of the pterygoid is expanded
and covered by unfinished bone.
Lower Jaw: The lower jaw of Galeops (Figs. 1-3) is short and
deep with a vertical unfused symphysis and a straight ventral edge.
A fenestra is present on the lateral surface of the jaw between the
dentary, surangular, and angular. A distinct coronoid process is not
present, although the posterior edge of the dentary extends above
Figure 5. The palate of Galeops whaitsi. A) as preserved in AMNH 5536; B)

reconstruction.
Abbreviations: bsp. basisphenoid: ect. ectopterygoid; ic, foramen for internal

carotid artery; m, maxilla; pal, palatine; pm, premaxilla; pt. pterygoid; ptf. transverse
flange of pterygoid; v, vomer.
the level of the postdentary bones. A small ventrally directed

retroarticular process is
present.
The dentary forms the anterior half of the lower jaw. E.\cept for a
small posterior area, the lateral surface of the dentary is covered by
numerous small pits and grooves. Posteriorly, the surface is smooth
and is depressed below the anterior portion of the bone. This
arrangement, which is similar to that of dicynodonts, suggests that
the muscles extended onto the posterior portion of the dentary and
that a horny beak was present anteriorly. In contrast to most
dicynodonts, no longitudinal groove is present on the upper edge of
the dentary lateral to the tooth row.
12 BREVIORA No. 465
Internally, only the upper half of the dentary is exposed, the

ventral half being covered by the splenial. The anterior end of the
dentary is without teeth; posterior to this, eight sockets are present
for teeth along the lateral margin of the bone. The anterior sockets
are slightly larger than the posterior sockets, indicating that tooth
size increased anteriorly. There is no indication that a distinct
canine was present. All of the sockets are vertical. The tooth row of
the dentary is longer than the tooth row of the maxilla, suggesting
that some anteroposterior movement of the jaw occurred during
closing.
The splenial is restricted to the internal surface of the lower jaw.
Posteriorly, the splenial is wedged between the angular and prear-
ticular. It is incompletely preserved anteriorly, although grooves in
the dentary show that it extended to the symphysis.

The angular forms most of the lateral surface of the posterior half
of the jaw. most prominent feature is the reflected lamina, which
Its
separates from the angular at the posterior edge of the lateral

mandibular fenestra and extends posteriorly and ventrally as a thin
sheath of bone.
The prearticular is a broad crescent-shaped bone that extends
from the articular to the dentary. It forms the ventral edge of the
lower jaw posteriorly, but its lateral exposure is reduced anteriorly
so that at the posterior edge of the reflected lamina it is restricted to

the internal surface of the jaw. The prearticular most probably
formed the margin of the adductor fossa, but since the dorsal
portion of the bone is not preserved, the full extent of the fossa is
unknown.
The surangular extends from the dentary to the articular, forming
the dorsal margin of the postdentary region of the jaw. It is
extensively overlapped by the angular so that, in lateral view, it is
restricted to the dorsal margin of the jaw, but in medial view, it
extends halfway down the internal surface of the jaw.

The articular is supported by the prearticular, angular, and
surangular. It forms both the articular surface for the quadrate and
the retroarticular process. The articular surface faces strongly
posteriorly. It is differentiated into two grooves separated by a

rounded ridge. The groovelateral is
elongate anteroposteriorly,
strongly concave mediolaterally and slightly so anteroposteriorly. A
small medially directed shelf forms the medial groove.
The internal surface of the postdentary region is partially covered
by the transverse flange of the pterygoid, so the presence of a

coronoid is uncertain.
Vertebrae: The fourteen vertebrae of Galeops are present

first
although many of these are incompletely preserved. Cox

(Figs. 6. 7),
(1959) differentiated the cervical from the trunk vertebrae of the
dicynodont Kingoria on the basis of the size of the parapophysis

and diapophysis and the thickness of the associated ribs, the cervical
vertebrae having more poorly developed parapophyses and more
slender ribs. In Galeops, a well-developed parapophysis is first seen
in the seventh vertebra. No parapophysis can be seen on the
corresponding area of the sixth vertebra, although a small one may

•
have been present ventrally. Thus, the first six vertebrae can be
considered to be cervicals.
The
atlas-axis complex is well-preserved (Fig. 6). It is primitive in
itsgeneral construction: the atlas centrum and axis are separate, and
judging from the articular surfaces, separate atlas and axis inter-
centra and proatlases would have been present. The atlas arches
pro at
art surf
intc intc
art surf
art
surf
Figure 6. The atlas-axis complex of Galeops whaitsi. A) lateral view; B) anterior

view of atlas arch and centrum. Drawing based on latex peels from AMNH
5536.
Abbreviations: aa, atlas arch; aa art surf, articular surface for atlas arch; ac, atlas
centrum; ax, axis; intc, intercentrum; intc art surf, articular surface for intercentrum;
prezyg, prezygopophysis; pro at art surf, articular surface for proatlas.
14 BREVIORA No. 465
meet above the neural canal. Transverse processes are present on

these elements sloping posteriorly and ventrally. The atlas centrum
does not extend to the ventral edge of the vertebral column. It has a
trefoil-shaped articular surface anteriorly. The neural spine of the
axis is not preserved. The prezygapophysis is convex and faces
dorsolaterally. A well-developed transverse process is present on the
axis extending posteriorly and ventrally from the base of the neural
arch.
Both the cervical and dorsal vertebrae are deeply amphicoelous.
A sharp ventral keel is present on the centra. The neural arches are
not although the sutural attachment is
fused to the centrum,
intimate. The base of is located anteriorly on the
the neural arch
centrum. A large intercentrum is present between the sixth and
seventh vertebrae, and presumably intercentra would have been
present between the more anterior vertebrae. A small intercentrum
is located between the seventh and eighth vertebrae. None is present
posterior to this.
A number of structural changes can be seen posteriorly along the
vertebral column:
1) The transverse process moves
dorsally. The transverse process
of the axis located at the dorsal margin of the centrum. On the
is
fourth vertebra, it is located more dorsally on the lateral surface of

the neural arch. The position of
the process of the following three
vertebrae is similar. A
further dorsal migration is seen between the
seventh and ninth vertebrae, with the transverse process reaching
the level of the zygapophysis.
2) The inclination of the transverse process changes from ventral
to dorsal. On the axis, the transverse process slopes ventrally and
posteriorly. On the fourth vertebra, the process is nearly horizontal

and directed laterally. On the eighth vertebra, a distinct dorsal
is
inclination is seen. This is further accentuated on the more posterior

vertebrae.
3) The parapophyses move dorsally. A distinct parapophysis is
first seen on the seventh vertebra, where it is located well down on

the anterior edge of the centrum. The intercentrum and posterior
edge of the sixth vertebra also contribute to its formation. The

parapophysis of the eighth vertebra is located slightly higher, and
the articular surface does not extend onto the seventh vertebra. The
parapophysis of the eleventh vertebra is located about midway

5=^PPf^^
ic PP
Figure 7. The vertebral column posterior to the axis of Galeops whaitsi in A)

dorsal and B) lateral views. Based on latex peels taken from AMNH
5536.
Abbreviations: ic, intercentrum; pp, parapophysis; tp, transverse process.
between the dorsal and ventral edges of the centrum. On the

fourteenth vertebra, the parapophysis is located on the dorsal
margin of the centrum close to the transverse process, although the

capitular and tubercular articular surfaces of the rib head remain
distinct. There is no indication that the diapophysis migrates
ventrally from the transverse process, as in dicynodonts (Cox,
1959).
4) The zygapophyses move closer to the midline. On the sixth
vertebra, the vertebra on which the zygapophyses are clearly
first
preserved, the zygapophyses are located at the edge of the centrum

and are inclined about 45° to the horizontal. The seventh to
eleventh vertebrae show a medial movement of the zygapophyses
and an increase in the angle that they make with the horizontal from
about 45° to about 60°. The medial movement of the postzygapo-
physes results in a coalescence of the processes supporting the
articular surfaces, resulting in a single posteriorly directed process.
Ribs: Of the cervical ribs of Galeops, only those associated with

the fifth and sixth vertebrae are preserved (Figs. 1, 2). Although
both of these are incomplete, they are distinctly less robust than the
more posterior ribs.
The first nine thoracic ribs are at least partially preserved. All of
these are double-headed. The tuberculum terminates the shaft of the
rib,and the capitular articular surface is formed by a process that
extends ventrally at an angle of about 35° to the shaft. The thoracic
ribs are only slightly curved. In the right rib associated with the
16 BREVIORA No. 465
twelfth vertebra, the shaft of the rib extends nearly straight from the
tubercular surface for about a quarter of the length of the rib. Thus
Galeops probably had a tall laterally compressed body.
Pectoral Girdle: The is formed by the

pectoral girdle of Galeops
paired clavicles, coracoids, procoracoids, and scapulae and by a
single median interclavicle (Figs. 1, 2). There is no evidence that
either a cleithrum or an ossified sternum was present.
The scapula has a tall, slender blade and an expanded platelike
base. A prominent tubercle for the scapular head of the triceps is
present on the posterior edge of the bone just proximal to the

glenoid. A distinct acromion process is not present, nor is a scapular
spine developed as is the case in dicynodonts (Boonstra, 1966).
The coracoid and procoracoid are subequal in size. The glenoid is
formed by the scapula and coracoid. The primitive screw-shaped
structure of the glenoid has been lost; the glenoid oi Galeops is short
and faces posterolaterally. The coracoid extends posterior to the
glenoid. The coracoid foramen is located within the procoracoid.
The clavicle has a triangular base and a rodlike stem. The

interclavicle is a long paddle-shaped element. Its head is not
preserved. Distinct grooves for a sternum are not present in the

stem.
Humerus: The humerus of Galeops is a slender bone with

moderately expanded proximal and distal ends in a distinct shaft
(Fig. 8). The proximal and distal ends are distinctly, but not
strongly, twisted on one another; when viewed proximally, the angle
between the long axes of the two ends is about 40°. The proximal
end of the humerus curves strongly dorsally; when seen in antero-
dorsal view (Fig. 8B), the dorsal third of the humerus makes an
angle of about 120° with the distal third of the humerus. The
entepicondyle is well developed, and an entepicondylar foramen is
present. The ectepicondyle is poorly developed, and no ectepicon-
dylar foramen is present. A groove passing proximodistally along
the lateral edge of the bone presumably held the nerves and vessels
that usually pass through the ectepicondylar foramen. The ulnar
articular surface is distinct and well developed. It faces primarily
ventrally, although it extends onto the distal end of the humerus.

The radial condyle, although distinct, is not a strongly rounded
capitulum like that seen in gorgonopsians and dicynodonts (Boon-

stra, 1965, 1966).
18 BREVIORA No. 465
Radius: The radius of Galeops is a slender bone about 60 percent

of the length of the humerus. The proximal articular surface is oval
in outline and is slightly concave. The distal articular surface is also
oval.
The ulna of Galeops is slightly longer than the radius (Figs.

Ulna:
1, The olecranon process is prominent, extending well past the
2).
humeral articular surface. The humeral articular surface is a well-
defined notch. Lateral to this, a distinct articular surface for the
radius is present.
Carpus: Only the dorsal surface of the proximal carpal bones of

Galeops is seen (Fig. 1). The radiale is short and has a small area of
finished bone on its dorsal surface. The intermedium and ulnare are
elongate. They have aof contact with one another

large area
proximal to the
perforating foramen. A portion of the lateral
centrale is present. This supports the intermedium and forms the
distolateral corner of the perforating foramen.
GALE PUS
Materials and Methods: Galepus
is represented by a single
species, Galepus jouberti, a and by
single specimen, 5541, AMNH
that comes from the Cistecephalus zone of the Beaufort series of
South Africa. Much of the skull and skeleton is represented by
impressions in a coarse sandstone from which latex peels were

made. Unfortunately, the counterpart block was not preserved, so it
was not possible to cast individual elements in three dimensions.
Skull: The skull of Galepus is represented by a cast of the internal

surface of the roofing bones and of the palatal flanges of the
premaxillae (Figs. 9, 12). It is similar to that of Galeops in its
general structure: the orbits and external narial openings are large;
the face and postorbital region of the skull are short; a large oval
pineal opening is present; the ventral margin of the postorbital

region of the skull is deeply emarginated; the occiput slopes
forwards; the orbit bounded posteriorly by a tall narrow postor-
is
bital; a secondary palate formed by the palatal flanges of the

premaxillae is present; and the internal surface of the parietal is
marked by a crescentic ridge and a cup-shaped depression antero-
Figure 9. Galepus jouberti. Drawing based on latex peel taken from AMNH
5541. Skull and lower jaw bones viewed from medial surface.
Abbreviations: C, clavicle; cal, calcaneum; cen. centrale; Co, coracoid; ect for.
ectepicondylar foramen; F, femur; H, humerus; IC, interclavicle; IL, ilium; IS.

ischium; mc metacarpal; mt
1. first 5, fifth metatarsal; PCo, procoracoid; PU, pubis;
r, radiale; R, radius; U, ulna.
20 BREVIORA No. 465
Unfortunately the sutures referred to by Broom

lateral to this ridge.
(1914) have been obliterated, so the exact relationships of the

individual skull elements are uncertain.
The left lower jaw is visible in internal view. It is more slender
than the jaw of Galeops, and its outline differs: the ventral margin of
the jaw, rather than being straight as in Galeops, is slightly concave
in profile,and the dorsal margin of the postdentary region extends
nearly straight from the dentary to the articular, rather than being
gently convex as in Galeops. A splenial is not preserved, but grooves
in the dentary indicate that it extended to the symphysis. The
sutures between the postdentary bones cannot be identified.
Six small peglike teeth are preserved in the dentary and an empty
aveolus is present anterior to these. A small edentulous region is
present anterior to the tooth row. Unlike Galeops where the sockets,
and presumably the teeth they contained, are vertical, the teeth of
Galepus slope anteriorly.
Postcranial Skeleton: A series of eighteen dorsal vertebrae of
Galepus is These are deeply amphicoelous and show a

present.
sutural attachment between the neural arch and centrum. The
structure of the neural arch is seen clearly only in the fifth vertebra
from the anterior end of the preserved series. It is like the neural
arch of the corresponding vertebra in Galeops: the zygapophyses are
strongly tilted and are near the midline, the postzygapophyses are
coalesced so the articular surfaces appear to be formed by a single
posteriorly directed process, and the transverse processes are large
dorsolaterally directed structures located at the level of the zyga-
pophyses.
The caudal region is represented by two disconnected series of
vertebrae, an anterior and a posterior series
series of nine vertebrae
of eight vertebrae.
They are separated by a space that would have
been occupied by six vertebrae and are separated from the sacral
region by a space that would have been occupied by about eight
vertebrae. Thus the tail would have included at least 31 vertebrae.
These decrease both height and length posteriorly. The zyga-
in
pophyses slope less steeply than in the dorsal series. A prominent

transverse process is located at the base of the neural arch on the
first vertebra in the series. No transverse process is seen on any of
the more posterior vertebrae. Haemal spines and arches are present
between all the vertebrae in the proximal series and between the first
two vertebrae in the posterior series.

Eighteen ribs are at least partially preserved on the right and six
on the left side. In all the ribs, the curvature is greatest near the
head, with the distal half of the rib being nearly straight and at an
angle of 30° to the proximal portion. The heads of the first three
ribs on the left side and the last two ribs on the right side are visible.
The anterior ribs are double-headed. The posterior ribs are single-
headed.
The pectoral girdle is seen in internal view. A large interclavicle is
present with a paddle-shaped stem and a diamond-shaped head. No
ossified sternum is present. The clavicle has a long narrow riblike
stem and a triangular head. The coracoids, procoracoids, and
scapulae are too poorly preserved for their structure to be fully
determined.
The humerus of Galepus is like that of Galeops in its proportions
and general features. In both, the entepicondyle is well developed,
the ectepicondyle is reduced, the proximal and distal ends are
distinctly twisted on one another, the ulnar surface faces strongly

ventral but also extends onto the distal surface of the bone, and the
pectoral crest is restricted to the proximal third of the bone. The

humerus of Galepus differs from that of Galeops in having an
ectepicondylar foramen and in the lack of an upturning of the
proximal end, the dorsal edge being nearly straight in anterodorsal
view.
Only the medial surface of the proximal end of the ulna is
preserved. As in Galeops an olecranon process is present, and the

medial surface of the proximal end of the ulna is concave. The right
radius is seen in lateral aspect. It is like the radius of Galeops in its
proportions and in having a slight S-shaped curve.
The hand overlies the foot, although much of the hand is missing,
exposing the metatarsals and phalanges of the foot.
Only three elements of the carpus are present, these being the
radiale, lateral centrale and first distal carpal. The radiale is short
and without finished bone. The first distal carpal is subcircular and
has a large area of finished bone on its ventral surface.
Of the metacarpus, only the first metacarpal is present. This is a
short phalangelike element. A large tubercle is present on the lateral
edge of the bone near its proximal end. The toes are represented by
three long, clawlike terminal phalanges.
22 BREVIORA No. 465
The pelvis is seen in internal view. It is primitive in having a large
pubis and ischium and in having well-developed anterior and

posterior extensions of the iliac blade. An advanced feature is the
enlargement of the obturator foramen.

The dorsal surface of the proximal end of the left femur is visible.
The proximal end of the femur is tilted dorsally about 35° from the
shaft of the bone.
The tarsus is represented by an incompletely preserved right
calcaneum, centrale, and the first four distal tarsals, all visible in
ventral view. The calcaneum is a tall, platelike element with a notch
for the perforating foramen on its medial edge. The centrale is a
cubical cartilage-covered element. The first three distal tarsals are
subequal in size. The fourth is larger, although it is less than twice
the size of the third in its linear dimensions.
All five metatarsals are present, visible in ventral view. They
increase in size from the first to the fourth with the fifth being about
equal to the third in length. The phalangeal formula of the pes is

2,3,?,?, 3. The terminal phalanges, where known, are elongate,
clawlike elements.
GALECHIRUS
and Methods: Galechirus is represented by a single
Materials
species, Galechirus scholtzi. and by two specimens: 1068, SAM
which shows the front half of the skull and the lower jaw in lateral
view and much of the postcranial skeleton (Fig. 10); and AMNH
5516 (now housed in the South African Museum), which shows a
pelvic and pectoral limb and fragmentary ribs and vertebrae (Fig.
11). Both of these specimens come from the Cistecephalus zone of
the Beaufort series of South Africa (Kitching, 1977). They are

preserved as natural casts. Molds taken from these specimens
provided a positive image that served as the basis for the descrip-
tions and drawings.
Skull: The skull of Galechirus (Fig. 10) is like that of Galeops in
having large orbits and external narial openings, a short temporal

region, and a deep emargination of the cheek posterior to the orbit.
In contrast to the condition in Galeops. Galechirus retains premax-
illary teeth and has a more elongate face.

Figure 10. Galechirus scholizi. Drawing based on latex peel taken from SAM
1068.
Ahhri'viations: a, astragalus; ac, atlas centrum; ax, axis; ax ic. axis intercentrum;
C, clavicle; cal, calcaneum; cer rib, cervicle rib; dt 1, first distal tarsal; ect for,
ectepicondylar foramen; F, femur; FIB, fibula; H, humerus, i, intermedium; IS.
ischium;], jugal; m cen, medial centrale; pi. pisiform; pm, premaxilla; prf, prefrontal;
PU, pubis; T, tibia; u, ulnare; U. ulna.
24 BREVIORA No. 465
The arrangement of the bones of the face in Galechirus is similar

to of Galeops. The premaxilla-maxilla contact is located
that
midway below the external narial opening. A separate bone,

presumably the septomaxilla, is present internal to the premaxilla-
maxilla contact. The maxilla meets the nasals midway along the
posterior edge of the external narial opening. The maxilla-nasal

suture slopes dorsoposteriorly from this point. Unfortunately, the
sutures between the maxilla and more posterior bones are obscure,
so the relationship of these bones is uncertain. Six small peglike
teeth are present in the maxilla, and an empty socket is present
posterior to these. The anterior three teeth are subequal in size and
are slightly larger than the posterior three teeth.
The premaxillae are separate. Each premaxilla has a well-
developed dorsal ramus that extends between the nasals and a
ventral ramus that meets the maxilla midway below the external
narial opening. The right premaxilla, which is seen in internal view,
contains two large, procumbent, chisel-shaped teeth. Three teeth are
present in the preserved portion of the left premaxilla. The posterior
two are approximately equal to the maxillary teeth in size and are
vertically oriented.The more anterior tooth is slightly larger and

slopes anteriorly. The missing portion of the premaxilla probably
held two large procumbent teeth similar to those seen in the right
premaxilla.
The prefrontal, which has been displaced slightly, is a triangular
element forming the anterodorsal margin of the orbit. Its lateral
surface is smooth and curves onto the dorsal surface of the skull.
The suborbital portion of the jugal is a narrow rodlike bone.
Posteriorly, a groove present on its lateral surface, probably for
is
an overlying zygomatic branch of the squamosal.
Lower Jaw: The lower jaw of Galechirus is long and slender. Its
ventral edge curves upwards, and the symphysis slopes forward. The
dentary forms the anterior two-thirds of the jaw. Its lateral surface is
covered by numerous small and grooves anteriorly. Posteriorly,
pits
the surface is smooth and
recessed below the more anterior
is
portion of the dentary. Presumably the adductor muscles inserted

on this area of the dentary. Only one tooth is preserved in the
dentary. This is a small peglike element located well anteriorly.
There does not appear to be room anterior to this for large
procumbent teeth similar to those seen in the premaxilla. nor is

there any evidence that such teeth were present.
The dorsal edge of the postdentary region of the jaw curves evenly
from the posterior edge of the dentary to the articular. A fenestra is
present between the dentary and the postdentary bones. The
reflected lamina of the angular separates from the angular at the
posterior edge of the fenestra. A small ventrally directed retroarticu-

larprocess present beneath the jaw joint. The sutures between the
is
individual bones of the postdentary region of the jaw are not visible.
Postcranial Skeleton: The vertebral column of Galechirus is
partially preserved in SAM 1068. Anteriorly, four cervical vertebrae

are present between the posterior end of the jaw and the pectoral
girdle. The first vertebra in this series is the atlas, the centrum of
which is seen in lateral view. As in Galeops, it does not reach the
ventral surface of the vertebral column. A large axis intercentrum is
present as a distinct element between the atlas centrum and the axis.
The following vertebrae are short and broad, their length being
about equal to their width across the posterior end of the centrum.
The axis has a posteroventrally directed transverse process located
well down on the neural arch. The centrum does not have a sharp
ventral keel, although a low ridge is present. The following two
vertebrae are seen in ventral view. A distinct ventral keel is present,
and intercentra are present between each of these vertebrae.
A series of three dorsal vertebrae, probably the tenth to twelfth,
liesbetween the shoulder girdle and the hand. These are seen in
ventral view. The keel has been reduced and intercentra are absent.
Distinct articular surfaces for the heads of the ribs are not visible.
Faint traces of eight presacral vertebrae are present between the
wrist and the knee. Little of their structure can be determined.
The caudal vertebrae are represented by two series: an anterior
series of twelve vertebrae and a more posterior series of three
vertebrae. A space equal in length to four centra is present between
the anterior series and the pelvis, and the two sections of the tail are
separated by a space equal in length to about ten centra. Thus, the
tail would have had in excess of 29 vertebrae.
The caudal vertebrae decrease both in height and length poste-

riorly. Haemal arches are present between each of the first ten
vertebrae, and transverse processes are present on each of the
26 BREVIORA No. 465
The more posterior vertebrae have

vertebrae in the anterior series.
neither transverse processes nor haemal arches.
Three ribs are preserved in SAM
1068. These are associated with
the three vertebrae seen between the shoulder girdle and the hand.
The more anterior of these is clearly double-headed. The separation
of the two articular surfaces is less distinct posteriorly.
Small, needle-like gastralia are present posteriorly, but these are
disturbed so their natural arrangement is unknown. This is the only
evidence of gastralia in therapsids (Romer, 1956).
Figure II. Galechirus scholtzi. Drawing based on cast of AMNH 5516 (now
housed in the South African Museum).
Abbreviations: a,astragalus; cal, calcaneum; cen, centrale; ect for. ectepicondyiar

foramen; ent for, entepicondylar foramen; F, femur; FIB, fibula; IL, ilium; IS,
ischium; 1 cen, lateral centrale; PU, pubis; r, radiale; R, radius; S, scapula; T, tibia; u,
ulnare; U, ulna.
The pectoral girdle is seen in ventral view in SAM 1068 and in
lateral view in AMNH 5516. The scapulae have tall, slender blades
and expanded platelike bases. A distinct rugosity on the posterior
edge of the scapular blade just proximal to the glenoid marks the
site of the origin of the scapular head of the triceps. An acromion
process is not present.

The coracoid and procoracoid are both large, with the coracoid
extending well posterior to the glenoid. The procoracoid foramen is
located within the procoracoid just proximal to the glenoid. The
interclavicle has a diamond-shaped head and a broad paddle-shaped
stem. The head of the interclavicle is recessed ventrally to receive the
medial end of the clavicle.
The left humerus is seen in ventral view in both specimens. The
proportions of this element are similar to the proportions of the
humerus of Galeops: it is a slender bone with a well-defined shaft
and with little expansion of the proximal and distal ends. The
pectoral crest is restricted to the proximal third of the bone. In
contrast to the condition in Galeops, both an ectepicondylar and
ectepicondylar foramen are present.
The radius is a slender bone, slightly shorter than the humerus. It
has a slight S-shaped curvature. The ulna is almost identical to that

of Galeops in its proportions and in the development of the articular
surfaces, although the olecranon process is developed. This
less well
difference may be a result of differences in the ontogenetic stage of

development of the specimens.
A complete carpus is present in SAM 1068 and most of the carpus
is present in AMNH
5516. All the elements primitively present in
the synapsid carpus are present, although these are all shortened
proximodistally so the length of the carpus is less than that of the

metacarpus. A distinctive feature is the expansion of the ulnare to
form a flat platelike bone.
The metacarpals increase in length from the first to the fourth,
with the fifthbeing about equal to the second in length. The
phalangeal formula is 2.3,3,3,3. The terminal phalanges are elon-
gate, laterally compressed, clawlike elements.

The pelvis is seen in ventral view in SAM 1068 and in lateral view
in AMNH 5516. The pubis and ischium are large, platelike ele-
ments. The pubis is damaged, so the size of the obturator foramen is
uncertain. The ilium is not completely known, although the pre-
28 BREVIORA No. 465
served portion shows that it had a tall, slender blade that extends
posterior to the acetabulum.

The femur is seen in postaxial view in both specimens. It is equal
in length to the humerus. The head of the femur is incompletely
preserved, although there is no indication of a dorsal tilting of the
proximal end such as is seen in Galepus. Also, no downturning of
the distal end of the femur is seen.
The tibia and fibula are elongate, slender bones approximately
equal in length to the femur and substantially longer than the radius
and ulna. Both bones are slightly curved with the concave edges of
the bones facing one another. The tibia is more robust than the
fibula, the diameter of the tibia at its narrowest point being about
twice the minimum diameter of the fibula.
A
nearly complete tarsus is seen in dorsal view in AMNH 5516,
and an astragalus is seen in dorsal view in SAM 1068. The
calcaneum is a large platelike bone. L-shaped. A
The astragalus is
perforating foramen passes between the astragalus and calcaneum.

Distal to the perforating foramen, the articular surface on the
astragalus for the calcaneum is strongly convex and extends onto

the dorsal surface of the bone. Proximal to the perforating foramen,
the articular surface on the astragalus for the calcaneum is concave.
The tibial articular surface is restricted to the medial edge of the
astragalus. The centrale is a small, cartilage-covered bone located
between the astragalus and first four distal tarsals. The distal tarsal
row consists of four elements, the fifth distal tarsal having been lost.
The first three are subequal in size, the fourth is about three times
the size of the third in its linear dimensions.

The metatarsals increase in length from the first to the fourth,
with the fifth being about equal to the third in length. The
phalangeal formula is 2,3,3,3,3. The terminal phalanges are elon-
gate, laterally compressed clawlike elements.
DISCUSSION
In considering the general evolution of therapsids, Hopson(1969)
recognized three basic groups: the Dinocephalia, characterized by

an interlocking of the incisors and the presence of a jaw joint that
permits no anteroposterior movement of the jaw; the Theriodontia,
characterized by the development of a movable quadrate permitting
some anteroposterior movement of the lower jaw; and the Anomo-
dontia, characterized by the ventral emargination of the cheek

posterior to the orbit and a jaw joint that permits considerable
anteroposterior movement of the lower jaw on the fixed quadrate.
The emargination of the cheek in the dromasaurs immediately
indicates that they belong within the Anomodontia. This placement
is supported by features of the skull of dromasaurs that are similar
to other anomodonts, including: 1) the formation of a secondary

palate by palatal flanges of the premaxillae and 2) the presence of a
lateral mandibular fenestra.
The most primitive anomodonts known are the venjukovia-
morphs Otsheria and Venjukovia from the Middle Permian of
Russia. These show the diagnostic feature of an emarginated cheek,
but, relative to dicynodonts, retain a primitivejaw joint, a primitive
palate, and primitive features jn the temporal region (Watson, 1948;
Olson, 1962; Barghusen, 1976). In many respects, the dromasaurs
are at the same grade of evolution as are the venjukoviamorphs.
This is demonstrated by the following features, which are shared by
dromasaurs and the venjukoviamorphs and which are different
from the more derived condition seen in dicynodonts:
1) No outfolding of the squamosal is present ventral to the

zygomatic raiûs of the squamosal. In dicynodonts, suth an
outfolding is present.
2) The palate is more primitive in showing a small interpterygoid

vacuity, transverse flanges on the pterygoids, an unfused
basipterygoid joint, and a smaller secondary palate. In dicyno-
donts, the interpterygoid vacuity has enlarged so that it
extends forward between the posterior ends of the vomers, the
transverse flanges of the pterygoids have been lost, and the
pterygoids are sutured to one another posterior to the inter-
pterygoid vacuity (Cluver, 1970).
3) The articular, as in Venjukovia (Watson, 1948), retains a
central ridge with two concave depressions on either side of it.
In dicynodonts, the central ridge is convex in profile, and a
concave area is present anterior to this (Crompton and
Hotton, 1967).
In addition, a number of features seen in the dromasaurs, but not

known at present in the venjukoviamorphs, are more primitive than
the condition in dicynodonts. These include:
30 BREVIORA No. 465
1) A separate atlas centrum and axis intercentrum are present in

the dromasaurs. In dicynodonts, these fuse to the axis (Cox,
1959). However, the centrum and axis may fuse during
atlas
ontogenetic development dromasaurs.

in the
2) No scapular spine or acromion process is present on the

scapulocoracoid. In dicynodonts, both these features are
present (Boonstra, 1966).
3) The shape of the interclavicle and the absence of a sternum in
the dromasaurs is primitive. In dicynodonts, the interclavicle
short and a large sternum is present (Boonstra, 1966).
is
4) The iliac blade in the dromasaurs is primitive in having a well-
developed posterior process. In dicynodonts, the anterior

ramus of the ilium is more strongly developed than the
posterior ramus (Boonstra, 1966).
Despite the primitive structure of the dromasaurs, features are

present that are advanced over the condition seen in the venjukovia-
morphs and are similar to the condition seen in dicynodonts. One of
these is the structure of the jugal. In Otsheria, the jugal is a triradiate
bone retaining well-developed zygomatic and postorbital branches.
In Venjukovia, the postorbital branch has been reduced although it
is still present (Barghusen, 1976). In both dromasaurs and dicyno-
donts, the postorbital and zygomatic branches of the jugal have

been reduced or lost. A second advanced feature is the presence of a
smooth area on the posterior end of the dentary that suggests that
the adductor muscles extended onto the lateral surface of the
dentary. In Venjukovia, an external mandibular muscle was present,

although it had only a small area of insertion on the lateral surface
of the dentary (Barghusen, 1976). A third advanced feature is the
structure of the septomaxilla. In Otsheria and Venjukovia, as in
primitive therapsids generally, the septomaxilla is a large element

with an exposure on the lateral surface of the face. In dicynodonts
and dromasaurs, the septomaxilla is reduced; in the dromasaurs, it
is restricted to the floor of the external narial opening.

This combination of primitive and derived features demonstrates
that the dromasaurs are members of a grade of evolution interme-
diate between the venjukoviamorphs and the dicynodonts, but it
does not demonstrate that it is a monophyletic group. For this, it is
necessary to show that features are present in the three genera that
are advanced over the primitive anomodont condition but are not
shared with dicynodonts. The most obvious such feature is the

dentition,which consists of reduced pegUke teeth. This is a clear
difference from both the primitive anomodont condition and the
dicynodont condition in which distinct canines are present. An
additional derived feature that separates dromasaurs from dicyno-

donts is the presence of a tall, narrow postorbital and a slender,
rodlike lower temporal bar. In dicynodonts, the postorbital bar is
low and the lower temporal bar is a flat, beamlike element.
Also the absence of a contact between the postorbital and
squamosal is a derived feature different from the condition in both
dicynodonts and venjukoviamorphs, where the postorbital extends
along the upper margin of the temporal opening to reach the
squamosal. However, the only dromasaur in which the posterior
extent of the postorbital is known is Galeops, so the possibility that
this is a derived feature present only in that genus cannot be
discounted.
Thus the present evidence, although not conclusive, suggests that
thedromasaurs are a natural group. Within the group, the three
genera can be placed in a single structural sequence with Galechirus
being most primitive and Galeops being the most derived
the
member of the sequence. The features that document this series are:
1) The Tooth Row: In Galechirus. premaxillary teeth are pres-

ent; in Galepus, premaxillary teeth are absent but the eden-
tulous region is short; in Galeops, a large edentulous region is
present.
2) The Proportions of the Face: In Galechirus. the face is
relatively long; Galepus and Galeops show a progressive
reduction in the length of the face (Fig. 12).
3) The Lower Jaw: is slender and
In Galechirus, the lower jaw
the ventral border is concave; in
Galepus and Galeops. a
progressive decrease in the length of the jaw is seen and the
ventral margin becomes straight (Fig. 12).
4) Humerus: In Galechirus and Galepus, the dorsal margin of the
humerus is straight when seen in anterodorsal view. In
Galeops, the proximal end of the humerus curves strongly
dorsally.
5) Ectepicondylar Foramen: In Galechirus. a well-developed
ectepicondylar foramen is present; in Galepus. this foramen is
reduced; in Galeops, an ectepicondylar foramen is absent.
32 BREVIORA No. 465
Figure 12. Reconstructions of the skulls of the dromasaurs. A) Galechirus

sholtzi; B) Galepus jouberti; C) Galeops whaitsi. Not drawn to scale.
Since the most derived member of this sequence is the earhest,

these three genera cannot represent successive stages in a single
evolving lineage. Rather, they must be regarded as members of
distinct grades of evolution in a single monopholetic radiation.
ACKNOWLEDGMENTS
I would like to thank Dr. R. L. Carroll of McGill University at
whose suggestion this project was undertaken. Dr. M. A. Cluver

provided sihcone rubber casts of the specimens housed in the South
African Museum. Drs. M. A.,Cluver, A. W. Crompton, and R. L.
Carroll read the manuscript and made many useful suggestions that
led to its improvement. Drs. J. A. Hopson and H. R. Barghusen
freelyprovided important information on the structure of the

venjukoviamorphs. I would also like to thank LilHan Mahoney for
proofreading the manuscript.
LITERATURE CITED
Bairp, D. 1951. Latex molds in paleontology. Compass of Sigma Gamma
Epsilon, 28: 339-345.
1955. Latex micro-molding and latex-plaster molding mixture. Science,
122: 202.
Barghusen, H. R. 1976. Notes on the adductor jaw musculature of Venjukovia.
a primitive anomodont therapsid from the Permian of theUSSR. Ann. S. Afr.
Mus., 69: 249-260.
Barry, T. H. 1974. A new dicynodont ancestor from the Upper Ecca (Lower
Middle Permian) of South Africa. Ann. S. Afr. Mus., 64: 117-136.
Boonstra, L. D. 1965. The limbs and girdles of the Gorgonopsia of the Tapino-
cephalus zone. Ann. S. Afr. Mus., 48: 237-249.
1966. The girdles and limbs of the Dicynodontia of the Tapinocephalus
zone. Ann. S. Afr. Mus., 50: 1-11.
Broom, R. 1907. On some new fossil reptiles from the Karroo beds of Victoria
West, South Africa. Trans. S. Afr. Phil. Soc, 18: 31-42.
1910. A
comparison of the Permian reptiles of North America with those
of South Africa. Bull. Am. Mus. Nat. Hist., 28: 197-234.
1912. On some new fossil reptiles from the Permian and Triassic beds
of South Africa. Proc. Zool. Soc, 1912: 859-876.
1914. On the origin of mammals. Phil. Trans. Roy. Soc. Lond. B, 206:
1-48.
34 BREVIORA No. 465
Carroll, R. L. 1976. Noteosuchus, the oldest known rhynochosaur. Ann. S. Afr.
Mus., 72: 37-51.

Cluver, M. a. 1970. The palate and mandible in some specimens of Dicynodon
testudirostris Broom & Haughtbn (Reptilia, Therapsida). Ann. S. Afr. Mus.,
56: 133-153.
1975. A new dicynodont reptile from the Tapinocephalus zone (Karoo
system, Beaufort series) of South Africa, with evidence of the jaw adductor
musculature. Ann. S. Afr. Mus., 67: 7-23.
Cox, C. B. On
the anatomy of a new dicynodont genus with evidence on
1959. the
position of the
tympanum. Proc. Zool. Soc. Lond., 132: 231-367.
Crompton, a. W., and N. Hotton III. 1967. Functional morphology of the
masticatory apparatus of two dicynodonts (Reptilia, Therapsida). Postilla, 109:

1-51.
HoPSON, J. A. 1969. The origin and adaptive radiation of mammal-like reptiles
and nontherian mammals. Ann. New York Acad. Sci., 167: 199-216.
KiTCHiNG, J. W. 1977. The distribution of the Karroo vertebrate fauna. Memoir
1. Bernard Price Institute for Paleontological research. Johannesburg, Univer-
sity of Witwatersrand, 131 pp.
Olson, E. C. 1962. Late Permian terrestrial vertebrates, USA and USSR. Trans.
Amer. Phil. Soc. N.S., 25: 1-225.
RoMER, A. S. 1956. The Osteology of the Reptiles. Chicago, University of
Chicago Press, 772 pp.
RoMER, A. S., AND L. 1. Price. 1940. Review of the Pelycosauria. Geol. Soc.
Amer. Spec. Pap. No. 28, 538 pp.
RoMER, A. S., AND D. M. S. Watson. 1956. A classification of therapsid reptiles.
Bull. Mus. Comp. Zool., 114: 37-89.
Watson, D. M. S. 1948. Dicvnodon and its allies. Proc. Zool. Soc, 118: 823-877.
B R E V^-0 R A
Mniseum of Comparative Zoology
us ISSN 0006 9698
Cambridge, Mass. 30JrNEl982 NrMBER466
SYSTEMATICS OF THE MEX/CANA

SPECIES GROUP OF THE
COLUBRID GENUS LAMPROPELTIS,
WITH AN HYPOTHESIS
MIMICRY
William R. Garstka'
Abstract. Variation within the/»('.v/(m?flgroupofkingsnakes, genus LaiiipiDpcl-

tis (Fit/inger). isdiscussed. Three species are recognized: L. niexicana (Ciarman). L.
alierna (Brown), and
L. //////\£'/;; (Blanchard). which is added to the species group. L.
leonis (Gunther) placed in the synonymy of L. me.xicana. QuaHtati\e and quantita-

is
tive characters of external and internal morphology are used to diagnose the ta.xa.
Outgroup comparisons with other Laiupropeltis and other colubrids indicate that L.
niexicana primiti\e relati\e to L. alierna. E\olution within the group and an
is
hypothesis of mimicry to explain pattern polymorphisms are discussed.

L. alierna as interpreted here is restricted to areas north of the Anticline of Arteaga,
Mexico and occurs throughout the Bolson de Mapimi and north to the foot of the
Guadalupe Mountains. /.. me.xicana is a montane and plateau form occurring on and
to the south of the Anticline of Arteaga. L. ruihveni has been found in nttrthwestern
Michoacan and adjacent Queretaro in the transverse \olcanic region. .S\mpatr\ is not
recorded for the taxa.
INTRODUCTION
The kingsnakes with Lamprope/tis me.xicana (Carman) are
allied
relatively small (less than one meter) little-known snakes which

inhabit the Chihuahuan Desert and adjacent areas. They occur from
'Museum of Comparative Zoology, Harvard Uni\ersity, Cambridge. Massachusetts

02138. Present address: Department of Biological Sciences. Universits of Alabama in
Huntsville. Huntsville, Alabama 35899.

2 BREVIORA No. 466
northern Michoacan, Mexico, north to perhaps southern New Mex-

ico. The taxa have been considered closely related and have been
group (Smith, 1942, 1944; Webb. 1961)ora

called variously a species
species complex (Gehlbach and Baker, 1962; Gehlbach and McCoy,

1965; Gehlbach, 1967; Tanzer, 1970). The nominal taxa are L. mexi-
cana (Garman), L. thayeri Loveridge, L. greeri Webb, L. leonis
Gunther, L. alterna (Brown), and L. hlairi Flury. These species have
been considered as a unit because of the following shared characteris-
tics: a light-edged red-centered black blotch, mottled speckling, dis-
tinct head, and long (Gehlbach and Baker, 1962). 1 also include in
tail
this group L. n///7\'6'A7/

(Blanchard) because it clearly fits these crite-
ria, and is sympatric with L. trianguluni.
The findings reported here suggest that the niexicana group con-
sists of three species-level taxa which are distinctive in several charac-
ter suites: L. alterna (including L. alterna and L. blairi), L. mexicana
(including L. mexicana, L. thayeri, L. greeri, and L. leonis), and L.
ruthveni (removed from the synonymy of L.t. arcifera).
The mexicana group is related closely to L. triangulum on the basis
of shared hemipenial characters (Smith, 1942) and may also show
close relationship to L. pyromelana and L. zonata when pertinent
characters are more closely examined in those species. The mexicana
group has been considered both as ancestral to the remaining species
of Lampropelt is (Webb, 1961) and as the most derived species (Blan-
chard, 1921; Smith, 1942; 1944; Tanner, 1953). Data will be presented
demonstrating that L. alterna is the most derived species in the group,
and that both it and L. ruthveni are closely related to L. mexicana.
Although the distribution of all three species appears to be presently
patchy and perhaps relictual, this is not sufficient justification for
considering them ancestral.
Four specimens of Lampropeltis, seemingly related to L. mexi-
cana. were collected by the author in Coahuila and Nuevo Leon,
Mexico, during 1975 and 1976. It was not possible to refer three of the
four specimens to any presently recognized taxon when compared to
the key in Webb (1961) and the descriptions in Gehlbach (1967).
Comparison with reference material did not clarify the situation. Of

the four specimens previously known from the vicinity of Galeana,
Nuevo Leon (Gehlbach and McCoy, 1965; Gehlbach, 1967), one was
identified as L. mexicana thayeri, one as L.m. alterna, and two as
L.m. thayeri X L.m. mexicana intergrades. The only additional
specimen from Nuevo Leon was the holotype of the obviously closely
1982 LAMPROrriTIS \1E\ICA\A SF'FCirS GFÔliP 3
related L. leonis. It was clear then that a revision ot the goup and
reidentification of the reference material was necessary.
One of the above specimens collected by the author (from San
Lazaro, Coahuila) has a series of 24 laterally truncated body rings;
thus it should have been classed as L.ni. ihayeri (Webb, 1961).
However, in comparison with the type specimen of ihuyeri and a
living specimen quite like the type, this Coahuila specimen was
clearly different. The head was much more triangular, as has been
noted for altertia (Brown, 1901; Gehlbach, 1967); the number of
ventral scaleswas nearly 209f greater than reported for ihayeri, and
the iris was silver-grey, like specimens from Texas and unlike the
gold-brown iris of animals collected in the mountains of Nuevo Leon.

Living specimens collected in the mountains of Durango, Mexico,
were similar in these three characters with those from Nuevo Leon,
and were likewise similar to specimens from Queretaro, except in
ventral scale number.
The specimens recently collected in Queretaro, Mexico, posed
another problem, as they closely resembled the single pre\ ious spec-
imen identified as L. ruthveni (Blanchard, 1921), which had been
synonymized (Williams, 1978) with L.t. anifera. These specimens
share many of the same features of L. me.xicana. and indeed are

distinguishable only on ventral scale number from some L. me.xicana
from Nuevo Leon. They are clearly not L. triaugulum nelsoni nor L.t.
anifera, with which they are sympatric; both subspecies of L. triangu-
lum have been collected (Williams, 1978) in the valleys that surround
the montane area where the ruthveni were collected. Therefore an
initialhypothesis of three valid taxa, aherna, me.xicana, and ruth-
veni, was considered, and characters were analyzed within that
framework.
The most striking feature of kingsnakes is their often brilliant color
and pattern. Color and pattern have been relied upon exclusively in
diagnoses of the species of Lampropehis (Blanchard, 1920, 1921;
Gehlbach, 1967; Smith, 1942; Webb, 1961) and the subspecies of L.
triangulum (Williams, 1978). The extreme variability of color and
pattern among individuals in the me.xicana group and the similarity
of the patterns o{ ruthveni and some me.xicana with some subspecies
o{ triangulum makes using only this character untenable.
Color and pattern variability is also well documented in Texas
aherna (Gehlbach and Baker, 1962; Gehlbach and McCoy, 1965;
Gehlbach, 1967; Tanzer, 1970; Miller, 1979). There is a gradation of
4 BREVIORA No. 466
patterns from extremes of more than 1 7 narrow rings with alternating

dots (a/ierna) to a low number (9- 7) of major bands with no alter-
1
nating reduced markings (hiairi). Tanzer (1970) reported a brood

from an alterna female which contained both hiairi and alterna
offspring. Melanism is also known in both alterna (M iller, 979) and 1
mexicana.
The morphological characters examined in this study are: 1) lepi-
and pattern, includingcolor of the iris, 3) osteology of
dosis, 2) color
the skull and vertebrae, and 4) hemipenes. The variation in these
characters isanalyzed in mexicana, ruthveni, and alterna, and each
taxon is compared with its respective sympatric triangulum popula-
tion. The three taxa are redefined, and a hypothesis of relationship
and of selection pressures leading to evolution within the group is
presented.
MATERIALS AND METHODS

Specimens
Both preserved and living specimens including all age classes were
examined. Embryo and hatchling specimens were examined and used
in ontogenetic comparisons with adults. Living snakes were sexed by
the method of Laszlo (1975) using probes manufactured by J & M

Specialty Parts. All living specimens in the author's collection will be
deposited in the Museum of Comparative Zoology. Wild caught
specimens only were used in the character analysis as captive bred
animals cannot be considered as statistically independent samples.
External Morphology
Scale counts recorded included ventrals (method of Dowling,

and dorsal
1951), subcaudals, infralabials, supralabials, temporals,
scale rows at mid-body (V method of Peters, 1964). The location of
the umbilical scar on hatchling specimens was noted. Dissected and
everted hemipenes of each taxon were examined. Statistical analysis
of data used methods from Sokal and Rohlf ( 1969). All sample means
indicated in the text are followed by ± one standard error of the
mean.
X-Rays
Radiographs of preserved specimens were taken on an F.E. Faxi-

tron X-ray machine using Kodak Industrcx R film. Measurements of
1982 I.AMPROPI.ITIS \U:\ICA\\ SIM (11 S (IROl I' 5
five thoracic vertebrae posterior to #20 were taken from each speci-
men radiographed. Coiling of preserved specimens prevented exami-
nation of the same five vertebrae on every animal. Measurements and
terminology of the vertebrae are from Johnson 1955a. 1955b) and (
Auffenberg(1963). Comparisons were made between replicate radio-

graphs of the same vertebrae, and between measurements from a
prepared skeleton and radiographs of the same vertebrae to be confi-
dent of accuracy and precision. Differences were less than 39f- Dor-
soventral radiographs of heads and measurements of the length and
width of the skull at various levels were also taken. Individual osteo-
logical elements of the skull were not examined. Comparisons were
again made of three sets of duplicate radiographs, and the differences
were less than 2%.
Field Work
During 1972, 1973, and 1975 trips were made to the Chihuahuan
Desert areas of Texas, principally to Brewster. Presidio, and Val
Verde counties. During 1974, 1975, 1976, 1979. and 1980 trips were
made Mexico. Collection of living specimens
to various localities in
provided information on the ecology and extent of distribution of the

group.
ANALYSIS OF CHARACTERS
Lepidosis
There is a marked difference in the numbers of ventral scales of the

three taxa (see Table 1 and Fig. 1). A one-way ANOVA shows that
there is significant heterogeneity of mean ventral scale number
among taxa (F = 85.3 at 2/83 df. 0.005). In a Student-New man-
p<
Keuls of differences between ranked pairs of means, all differ-
test
ences are significant at the \% level. In order to be assured that

distinctpopulations were sampled, the distributions were tested for
normality. The individual taxon distributions are not different from
normal by the Kolmogorov-Smirnov cumulative test (D {alterna] =
0.09. critical value = 0.29. a = 0.01; D [mexicana] = 0.07. critical
value = 0.30, a = 0.01; Dlruthveni] = 0.13, critical value = 0.30, a =
0.01 ). The total distribution was tested for normality and was found
to be significantly different from normal (D = 0.27. critical value =
0.18).
BREVIORA No. 466
1982 LAMPROPEITIS MEMCASA SPrClFS CROIP 7
The variation among the ventral scale means oialterna, niexicana.

and ruthveni appears as a north to south cline. This variation, how-
ever, is not clinal within a taxon, and the differences among taxa are
stable in captive-bred animals. Clinal variation in ventral scale
number has been documented for Coluber constrictor (Auffenberg,
1955) and Lampropeltis triangiilum (Williams, 1978). In both taxa

the change is gradual, with counts high in the south, low in the north.
The effects of differing temperatures of incubation on scale counts
(Fox et a/.. 1961) could explain that situation, although no experi-
mental evidence was given for either species.
The geographic distribution of ventral scale numbers within each
taxon appears random (Table 2). The southernmost specimens of
alterna from just north of Saltillo, Coahuila (USNM 10819) and 1
from near San Antonio, Durango (TCWC 36892), have 223 and 218
ventrals, respectively. The northernmost mexicana specimens, three
from the Rio Mimbres area of Durango (UCM 21061 and 2 Garstka
collection) and one from the Anticline of Arteaga near Los Lirios,
Coahuila (ITESM uncataloged), have 203, 202, 204, and 190, respec-
tively. The Los Lirios mexicana has the lowest ventral number of any
niexicana examined.
The hypothesis that the north to south cline in ventral scale counts
may be a result of differing incubation temperatures was examined by

incubating two clutches oi mexicana eggs and one clutch oi alterna
eggs in the same container at the same time {alterna: 3 August 23
October; mexicana: 14 July 25 September; 28 June- 18 September
1977). The same means of ventral counts are alterna = 218.5 ± 2.50
(H=2)\ mexicana = 204.6 ± 1.45 (N=10), and the difference is signifi-
cant (t = 8.47, p < 0.001). In an additional sample of captive-bred
alterna (218.8 ± 0.38, N=40) and captive-bred mexicana (201.6 ±
7.87, N=7), the difference is also significant (t = 8.28, p < 0.00 The
1 1 ).
conditions and dates of incubation for the latter sample are not
known.
Juvenile specimens usually possess an umbilical scar. The number
of ventral scales anterior to the scar is 70.9 ± .48 for 1 1 1 mexicana
1
and 190.0 ± 1.69 for \2 alterna (1-= 12.98, p< 0.001 ). The length of
the scar is the same in both taxa; the mean lengths in the samples are
4.5 ± 0.25 scales for mexicana and 5.0 ± 0.37 scales for alterna (t
=
1.25, 0.3 > p > 0.2). No sex differences were noted. The single
juvenile rw//7V'£'/7/(KU 155528) had no visible scar.
BREVIORA No. 466
3
a.u
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53
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1982 LAMPR()Pi:i.riS Mi:\KASA SIM C II S (.ROl
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en
10 BREVIORA No. 466
The pattern of ventral scale counts in these taxa is clearly differ-

ent from that of each sympatric triangulum variety (Williams, 1978).
L. triangulum celaenops and L.t. annulata are sympatric with
alterna, L.t. nelsoni and L.t. arcifera with ruthveni, and there is
apparently no triangulum sympatric with the montane me.xicana.

The ventral scale counts of Williams' (1978) and Quinn's (in prepa-
ration) samples of celaenops, annulata, arcifera, nelsoni, and a new
subspecies can serve to distinguish each species of the mexicana
group from its sympatric L. triangulum subspecies (Table .) I
There is no clear separation of the taxa in other scale characters.

The uppermost preoculars in most mexicana and ruthveni extend
onto the top of the head to or nearly to the frontal, whereas in
alterna they rarely do so. There is much variability and overlap
between taxa in the number and arrangement of the temporal scales.
There is also no differences among taxa in numbers of labial scales,
subcaudals, or dorsal scale rows, and all specimens examined pos-
sess a loreal (Table 3).
Color Pattern
The variable head and snout markings of this group can be

grouped into four categories: I) asymmetric spot or blotch markings
without red, II) snout evenly black to the middle or posterior of the
parietals, with or without a red infusion. 111) a red-centered, three-
lobed cloverlike marking with the anterior lobe across the frontal or
usually the prefrontals, and IV) a similar three-lobed marking,
without red, that can have the anterior lobe either split to form a
double Y, detached, or enlarged and subrectangular (Fig. 2). None
of these categories is exclusive to any taxon, but most ruthveni are
category 11, and most alterna are category 1. The head markings
connect to the nuchal blotch only in some mexicana. The nuchal
blotch oi tnexicana and alterna is usually light-centered and can be
split longitudinally. Postocular stripes are usually present in all taxa
(absent in UCM 21061, a mexicana), but may be reduced or obs-
cured, as occurs in category 11.
All living specimens examined o{ alterna have a distinctive silver-
grey iris; mexicana and ruthveni have a golden-brown iris. Other
Lampropeltis (getulus, triangulimi, pyromelana, and zonata) exam-

ined also have a golden-brown iris.
1982 LAMPROPI.I TIS \n:\ICA\A SPFCIFS GROCP 11
12 BREVIORA No. 466
0\
oo
1982 LAMPROPKI/riS MKXICASA SPFCHS (iKOUP 13
The number of primary, usually red-centered, body markings var-

ies considerably and is not by itself indicative of any taxon (Table
1). The hlairi form of alienia has the fewest primarv markings
(9-17).
The 47 offspring produced in ten broods by one male and two
female mexicana collected from a single site near San Juanito,
Nuevo Leon, are variously ringed, blotched, or spotted. All possible
variants of the dorsal head pattern occur in these offspring, and
ground color ranges from lemon-yellow through various shades of
brown, orange, and grey. The blotches vary in number from fewer
than 20 to more than 40, and in shape from spots to rings extending
around the body. The number of body bands falls within the ranges
of all the forms except hlairi. In some, alternating reduced markings
occur in one or several rows. One specimen has red only in the
nuchal blotch. Thus, neither number, level of truncation (Webb,
1961) or alternating reduction (Blanchard. 1920. 1921: Smith. 1942)
of body bands are diagnostic characters. The local \ariabilii\
characteristic of mexicana and alterna is not present in any
triariguluni subspecies, nor are alternating reduced markings
recorded for any L. triangu/um.
Ventral pigmentation ranges from nearly entirely black to nearly
entirely background color in alterna and mexicana. The ruthveni
examined most commonly have rings which extend around the
body; this also occurs with the anterior rings in some mexicana. The
venter oi alterna and mexicana can also be checkered in black and
background color as in L.i. iriangulum and Elaphe guttata (see
figure in Smith, 1942, pp. 202 203), and some ruthveni have a
checkered red pattern ventrally.
A ventrolateral expansion of the first (anteriormost) tail blotch
with red extending onto (usually across) the subcaudals is character-

istic of mexicana. This character is not evident in any of the alterna
examined. The underside of the tail of one specimen of mexicana
(BMNH 1946-1-4-10) has red infused anteriorly in a longitudinal

black line. The tails of the ruthveni examined have a ringed condi-
tion similar to mexicana; however, the first tail ring is not usually
widened ventrolaterally. The anterior tail rings are not different
from body rings in any of the L. triangulum subspecies.
The L. triangulum subspecies sympatric with each taxon can be
distinguished by a combination of color patterns and scale charac-
ters from the sympatric member of the mexicana group (Table 1).
14 BREVIORA No. 466
The snout color and symmetric body annuli of ruthveni make these
animals easily distinguishable from the sympatric nelsoni. Similarly,
the high number of annuli of celaenops and annulata are different
from the usually few annuli of the ringed or saddled blairi form of
alterna. In the other cases, some combination of color pattern with
ventral scale number serves to diagnose members of the mexicana
group.
Skull
Data from dorsoventral radiographs of similarly sized, adult

alterna (N = 7) and mexicana (N = 12) were compared to quantify
the observation of a more triangular head shape in alterna. The
premaxilla to occipital condyle length, the width of the maxilla at
L. alterna
L. mexicana I I
CO
O
I
o
<
LU
DC
LU
GQ
.72 .76 .80
PRE- -^ POST-ORBITAL WIDTH

Figure 3. The ratio of pre-orbital and post-orbital width of L. alterna and L.
mexicana adults.
1982 L. A. \f PROPt:LT/S MEX/CA\ A SPECIES GROVP 15
the prefrontals, the least width of the frontals, the maximum widths
across the post-orbitals, across the supratemporals, and across the
quadrates were measured. The ratio of preorbital to postorbital
width was used to quantify triangularity of the skull, as that should
be least influenced by kinesis. The mean ratio of the alterna sampled
is 0.66 ± 0.007 and of the mexicana
sampled is 0.77 ± 0.010. They
are significantly different (t = 7.343, p<
0.001) (Fig. 3). The single
ruthveni adult examined for this character (USNM
46558) has a
ratio of 0.74.
Additionally, ten juveniles each of alterna and mexicana were

compared for this character. The mean ratio of the sample oi mexi-
cana juveniles is 0.74 ± 0.007, with one specimen outside the range
of variation of the adults, in the direction of alterna. In the sample
of juvenile alterna, five are outside the variational range of adult
alterna in the direction of mexicana,and the mean is 0.69 ± 0.008.
Thus the two taxa are more similar as juveniles, and, importantly,
are more like mexicana than alterna as juveniles. This indicates that
there is differential growth to the adult stage in alterna.
Adults of other species of Lampropeltis were also examined in
outgroup comparison for this character. L. triangulum amaura and
L.t. polyzona both had a preorbital to postorbital ratio of 0.77, a
single L. pyromelana had a ratio of 0.73, and a single L. calligaster

had a ratio of 0.73. Thus,
appears that the lesser triangularity of
it
the skull in the mexicana condition is not only the common condi-
tion for juveniles, but also for other Lampropeltis.
Vertebrae
Eight or more individual thoracic vertebrae (Johnson, 1955a,

1955b;
=
first precaudal series, Auffenberg, 1963; = lumbar series.
Bullock and Tanner, 1966) prepared from each of eight adult speci-
mens oi alterna and four adult mexicana were examined. X-rays in
lateraland dorsoventral aspect of an additional three adult and five
juvenile alterna and six adult and nine juvenile mexicana were also
examined. No ruthveni vertebrae were available. None of the com-
monly used measures (Johnson, 1955a, 1955b) or ratios (Auffen-
berg, 1963) showed any significant difference between adults of the
two taxa. The amount of individual variation in qualitative appear-
ance of the vertebrae is substantial, however, even in the small
sample examined; because of the lack of descriptions of the range of

16 BREVIORA No. 466
o
D.
'J
r-
N
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O 5
r--
r-
o
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1982 ^/1 A/ /'/?0/'£/,r/5 A/ /:".V/C/<;V/1 SPECIES GROUP 17
individual variation of colubrid vertebrae, and the importance of

vertebral characters in the systematics of fossil forms, an analysis of
the vertebrae follows (Fig. 4).
The neural spine may be blunt or pointed, and in all individuals of
mexicana examined it overhangs both anteriorly and posteriorly. In
lateral X-ray the neural spine of a single alierna, USNM 110819,
appears to not overhang anteriorly. The zygosphene is crenate to

convex from above and usually has a broad shallow notch. In ante-
rior aspect the dorsum of the neural arch is flattened or sHghtly
arched and is thinned centrally; the sides are obtusely angled, form-
ing a dorsolateral articulating surface. The dorsal articulating surfa-
ces of the anterior zygapophyses are ovoid to reniform and their
can be blunted or pointed, directed later-
lateral accessory processes
upward, and if upward, may be curved. The processes may be

ally or
rounded to dorsoventrally compressed, and if compressed, may be
tilted to be parallel to the centrum. The cotyle may be round to
ovoid and if ovoid, may have accessory flanges or fillets,
usually
ventrally, that make it appear rectangular. The parapophyses may
or may not project ventral to the cotyle and
may be projected ante-
riorly. The paradiapophysialarticulating surfaces are in the form of
a figure-8 with the upper portion larger and directed anteriorly, and
the lower surface directed laterally to posteriorly. There is a small
foramen on the parapophyses lateral to the cotyle on each side
approximately at its midline. The haemal keel is usually ridgelike,

though it may be flattened or absent. The subcentrum ridges are
straight to slightly concave. There are two pairs of subcentrum
foramina, one pair on either side of the haemal keel and the other
more lateral and dorsal, just ventral to the lateral flange of the body.
The condyle is oblique. In posterior view, the neural arch is rounded
dorsally and flattened ventrally with a central ventral ridge. There
ate also lateral minor ridges extending the length of the canal on
either side approximately one third of the distance superior to the
base. The ventral exterior articulating surfaces of the posterior
zygapophyses are ovoid to reniform with the long axis perpendicu-

lar to the centrum. The medial articulations that match the dorsal
zygosphene corners, the zygantra, appear as enlarged triangular

holes lateral to the neural arch. There are small projections poste-
rior and lateral to the posterior zygapophyses just dorsal to the
ventral articulating surfaces.
18 BREVIORA No. 466
Juvenile vertebrae present quite a different appearance from

those of adults, which is most noticeable in the proportionally larger
neural canal. The ratio of length of the centrum to width of the
neural canal in a sample of three adult mexicana is 2.1 ± 0.10, of
three adult alterna is 2.3 ±
and of eight juvenile mexicana is 1.2
0.08,
± 0.03. The neural spine
also proportionally higher. The ratio of
is
length to height of the neural spine of a sample of three mexicana

adults is 2.4 ±0.34 and of three juvenile mexicana is 1.5 ± 0.09. The
juvenile alterna sampled had vertebrae insufficiently ossified to
obtain the measurements for comparison. Since nervous tissues
have grown most in relation to other tissues at birth, and since
ossification is incomplete then, these differences are to be expected.
The thoracic vertebrae of L. triangulum differ from alterna and
mexicana in the form of the zygosphene which is arched rather than
flattened (Brattstrom, 1955; Auffenberg, 1963). All of the tricolored
kingsnakes examined (L. triangulum, L. pyromelana, L. zonata, L.

alterna,and L. mexicana) share the single foramina lateral to the
cotyleand the two pairs of subcentrum foramina. L. getulus and L.
calligaster,however, have two pair of foramina adjacent to the
haemal keel and a pair of foramina on each side of the cotyle.
Hemipenes
Previous workers (Smith, 1942; Gehlbach and Baker, 1962) have
reported on the hemipenes of alterna and mexicana, and allied the
taxa with L. triangulum on the basis of overall similarity of gross
structure. The hemipenes of three adult mexicana, five adult alterna,
and two subadult ruthveni have been examined. They are uniformly
longitudinally flounced proximally, then abruptly spinulate, quickly
grading into slightly spinulate fringed calyces. The sulcus is single
and terminates laterally on the very slightly bilobed organ. This
morphology is similar to that of L. triangulum.
The spination on the hemipenes is clearly different among the
mexicana group, however (Fig. 5). The spines o{ mexicana are very
slightly recurved and are about 0.4 mm long. They appear diamond-
shaped in cross section and are tightly packed on the organ. The
spines of alterna and ruthveni are also recurved but are longer, to
approximately 0.8 mm in alterna and 0.6 mm in ruthveni, and the
supporting tissue makes them appear more ovoid or flattened in
cross section. The spines also appear to be fewer in number in these
1982 LAMPROPELTIS MEXICAN A SPECIES GROUP
L alterna
L. mexicana
L. ruthveni
mexicana, and L. ruthveni. Lateral

Figure 5. Hemipenial spines of L. alterna. L.
(L) and distal ( R) aspects.
taxa than in mexicana. The spines of both ruthveni examined are
more strongly recurved than those of both mexicana and alterna.
Ecology
The distribution of the entire mexicana group has been character-

ized as xeric to subhumid (Gehlbach and Baker, 1962) and roughly
corresponds to the limits of the Chihuahuan Desert as described by
Morafka(1977)(Fig.6).
20 BREVIORA No. 466
L. alterna is found within the Chihuahuan Desert, usually in

association with typically desert floral elements such as Prosopis
and La/TPfl (Jameson and Flury,I949; Mecham and Milstead, 1949;
Worthington, 1974). All known localities are within or north of the
Bolson de Mapimi. Specimens collected in the Chisos Mountains,
Brewster County, Texas (Murray, 1939) and near Saltillo, Coahuila,
Mexico (Smith, 1941) were found in rock crevices, though the spe-
cies appears to be primarily nocturnal (Miller, 1979).
L. mexicana seems to be limited, except for CM 59980, to for-
ested mesic areas peripheral to the desert, but with xerophytic vege-
tation such as Echinocactus and Ariocarpus. At mexicana localities
in Nuevo Leon, San Luis Potosi, and Durango, Mexico, the vegeta-
tion may be termed "mesic to arid oak-pine forest" (J. Henrickson,
personal communication), with the following dominant plants in
common at the three localities: Finns cembroides, Quercus sp..
Juniper us sp., Buddleja cordaia, and Bouvardia ternifolia. The

author has collected specimens in rock crevices, under stones in an
arroyo, and active on the road at night.
L. ruthveni is presently known from the northeastern portion of
the trans-volcanic region in a habitat very similar to mexicana. In
the vicinity of Amealco, Queretaro, Mexico, ruthveni are locally
abundant in areas with scattered patches of Quercus-dom\n?iied
woodland along arroyos and on hilltops. Bouvardia is common here
as well. All recent collections have been of animals either in crevices
or diurnally active.
Both mexicana and ruthveni appear to be more mesic species than
aUerna, occurring in more temperate forested areas. However, CM
59980 was collected in a desert area with stunted Larrea, cacti, and
short grasses (D. Morafka, personal communication). The elevation
of this rocky area is over 2,000 m, and while it is surrounded by a
it is also closely adjacent to mesic upland areas. Many
sandy plain,
similar hilly areas in the vicinity have been deforested in recent times
(T. Wendt, personal communication) and the area may have been
woodland as recently as 100 years ago. L. aherna may also occupy
upland (Worthington, 1974) or riparian habitats within the desert;

however, it appears to be primarily a desert species. The higher
ventral scale numbers oi aherna may indicate adaptation to a desert
environment, as Klauber (1941) found a correlation of higher ven-
tral counts in desert habitats and lower counts in more humid coas-
tal habitats in 1 2 of 1 3 taxa tested.

1982 i.A\inR()ri:i.Tis MEXicASA sim cii s (iKoui' 21
DISCUSSION
Phylogeny
Underwood (1967) considered Lampropeltis as derived from

Elaphe on the basis of similarity of a long intrapulmonary bron-
chus, a series of skull characters, presence of similar scale tubercles,
similar arrangement of pits on the head, and similar paired pits on
trunk scales. The characters that define Lampropeltis as a group
distinctfrom Elaphe are the entire anal scale and the unkeeled
dorsal scales. Minton (1976) and Minton and Salanitro (1972) dis-
cussed the phylogeny of colubrids, based on the immunoelectropho-
retic analysis of serum proteins and found that both E. guttata and
L. getidiis have the same degree of difference from E. obsoleta.
Karyological evidence also demonstrates this similarity to Elaphe. A

= 36 (16 macrochromosomes and 20 micro-
karyotype of 2N
chromosomes) has been reported for L. getulus, calligaster, and
most Elaphe
allerna, as well as for (E. suhocularis is an exception)
(Bury, et al. 1970; Baker, et al. 1972). This close relationship
between Lampropeltis and Elaphe is also substantiated by vertebral
morphology. The form of the neural spine and the arrangement of
the vertebral foramina are similar in L. getulus, L. calligaster, and
Elaphe.
All of the tricolored kingsnakes (alterna, mexicana, pyromelana.
ruthveni, triangulum, zonata) share grossly similar derived hemipe-
nial (Smith, 1942) and vertebral character states that are unlike
those of the remaining species in the genus, L. calligaster and L.
getulus, which are in turn similar to each other and to Elaphe (Cope,
1898).
Captive-breeding experiments (Miller, 1979; Tryon and Garstka,
in preparation) have produced alterna
X mexicana, pyromelana X
alterna, and pyrotnelana X zonata hybrids. None of these species
will mate with any triangulum subspecies tested in captivity. This
can be taken as tentative evidence of their relationships. Therefore,
it
appears that all the tricolored kingsnakes share a common ances-
tor,and that the mexicana group forms a logical unit within that
lineage, perhaps most closely related to pyromelana and zonata.

Within the mexicana group, alterna is the most derived species, as
shown by its iris color and head shape. These characters suggest that
alterna is the most derived species within the genus as well (Fig. 6).
22 BREVIORA No. 466
mexicana alterna
/
g
ruthveni
zonata
pyromelana
triangulum
getulus
calligaster
ELAPHE LAMPROPELTIS
/
b
Figure 6. Hypothesis of relationship within the genus Lampropeltis. Derived

character states uniting the taxa are: a) paired pits on trunk scales, long intrapulmo-
nary bronchus; b) entire anal scale, unkeeled dorsal scales; c) single paired foramina
lateral to hemal keel and cotyle, hemipenes flounced proximally and spinulate dis-
tally, pattern in three colors including red; d) behavioral traits in male aggression and
reproduction; e) light-edged, red-centered blotch; alternating reduced markings
possible; and g) grey iris, "triangular" skull.
1982 LAMPROPEl.T/S \tEX/CA.\'A SPECIES GROVP 23
The data presented here suggest that the three taxa hypothesized.
L. niexicana, L. altertia, and L. ruthveni, are distinct morphological
species. However, the taxa are allopatric. as are the two apparently
most closely related species, L. pyrome/ana and L. zonata. There-
fore the allocation of species status must still be regarded as an
hypothesis until appropriate species borders or contact zones are

established.
The species borders of niexicana and alterna, that is, the Anticline
of Arteaga and the uplands of central Durango at the periphery of
the desert, should be further collected. The presence o{ alterna at the
base (USNM II08I9) and niexicana (ITESM uncataloged) at the
top of the Anticline of Arteaga indicate a high probability of a
contact zone. Southern San Luis Potosi, Guanajuato, and northern
Michoacan including the Sierra de Santa Rosa (Duges, 1897) and
the escarpment of the basin of the Rio Lerma should be explored as
well for a species boundary or area of intergradation as this is
between the localities oi niexicana and ruthveni. Finally, the moun-
tainous area of northern Durango north of Otinapa and west of
Santiago Papasquiaro and the upper reaches of the Rios San Juan
and Verde need investigation. There are no kingsnakes known from
that large area between the localities of niexicana and pyronielana
(Taylor, 1940).
In addition, small sample sizes of some of the Mexican Plateau
subspecies oi triangiduni make it difficult to determine the extent of

sympatry, especially with ruthveni, and make comparisons difficult.
More collections from that area, and revision of the genus or at least
of the tricolored group, are necessary before the relationships
among the various taxa can be better understood.
Variation ami Selection of Color and Pattern
Many explanations of the significance of color and pattern in

snakes have been proposed (see review by Jackson et ai. 1976).
Aposematism, to warn potential predators, has been proposed as an
explanation of bright colors on venomous snakes (Goodman and
Goodman, 1976; Gehlbach, 1972; Smith, 1969). Mimicry of coral
snakes by nonvenomous forms has been discussed extensively
(Smith, 1969; Gehlbach, 1972; Greene and Pyburn, 1973; Echter-
nacht, 1973; Hecht and Marien, 1956) as an explanation for the
brightly colored rings on many snakes. Crypsis on broken back-
24 BREVIORA No. 466
108 106 104 102 100
Figure Distribution of the species in the mexicana group. State boundaries are
7.
shown as a dotted Une, national borders as a broken hne, and the extent of the
Chihuahuan Desert (after Morafka, 1977) as a solid line. The Anticline of Arteaga is
the band of mountains extending into the desert across southern Coahuila at about
latitude 25° N. North of that is the Mapimian portion of the desert, and south of the
Anticline is the Saladan portion of the desert (Morafka, 1977). Symbols denote only
localities and not numbers of specimens.
1982 LAMPROPKI.TIS Mi:\IC WA SPFCIFS (iROl'P 25
grounds (Pough, 1976; Brattstrom, 1955), or lack of crypsis on a

unicolored background (Camin and Erhlich, 1958) have also been
cited to explain the presence or absence of a ringed or blotched
pattern.
Gehlbach (1972) reported experimental evidence that self-mimicry,
that is, the coral snake behavior of flipping the tail over, thwarted
predation by coatimundis {Nasua) and peccaries {Tayassu). This

effect was enhanced if the rubber models were painted aposemati-
cally. Evidence of the effectiveness of coral snake mimicry has been
provided by experiments offering striped and ringed stick models to

naive captive-raised motmots {Eumomota superciliosa) (Smith,
1975). The birds responded to red and yellow ringed models by
avoidance and, in some cases, alarm notes.
L. mexicana and L. alterna, even though they may be brightly

colored, do not look very much like coral snakes. Gans (1961) dis-
cussed an hypothesis of mimicry in Dasypeltis, a small nonveno-
mous snake that eats eggs of birds large enough to eat it. He
demonstrated a correspondence of pattern between Dasypeltis and
the local possible viperid models over its entire range. The pattern of
the particular similar-sized viper in its range was mimicked. There-
fore, the distribution of venomous snakes of the same general size
and habits as Lampropeltis, and occurring within the range of the
mexicana group, was examined. Crotaline models were considered,
as the fine dark speckling on the scales lends an illusion of texture to
the smooth-scaled Lampropeltis.
Agkistrodon mokeson pictigaster occurs in Texas west through

the Big Bend region, but west of Val Verde County occurs only in
the vicinity of permanent water sources. Crotalus lepidus is found
throughout the range of alterna and is present in the same micro-

habitat. Crotalus lepidus, with its alternating broken pattern, can
easily be mistaken for alterna. A similar pattern is also seen on
Trimorphodon from west Texas. Micrurus fulvius is found in Texas

west to Val Verde County.
The variation in Texas alterna could be a function of multiple
models, the blairi form of alterna being a mimic of Agkistrodon.
The constancy of pattern in alterna outside of the range of
relative
Agkistrodon could be due to having a single available model in

those areas, C. lepidus. In recent years Val Verde County, Texas,
has had increased rainfall, and the hlairi form, first collected in 1948
26 BREVIORA No. 466
Figure 8. Hypothesis of mimicry in the mexicana group, a) Crolalus lepidus (L)

with sympatric L. alterna (R); b) Agkistrodon mokeson (L) with sympatric hlairi
morph of L. alterna (R); and c) Crotalus iriseriatus (L) with sympatric L. mexicana
(R).
1982 LAMPROPELT/S MEXJCANA SPECIES GROUP 27
(Axtell, 1951), is today quite commonly found. Agkistrodon are
common along the Devil's and Pecos river drainages as well, in

similar habitat with blairi; Micrurus fulvius is found there as well.
The sporadic appearance of blairi-\'\ke animals in Presidio and
Brewster counties in the Big Bend region is correlated with the
distribution of Agkistrodon. Both A.m. pictigaster and M. fulvius
are very conspicuous, brightly colored animals, and though C. lepi-
dus is also sympatric in Val Verde County, its dull appearance may
make it a less efficient model. The increased red in the blairi morph
may be a result of stimulus generalization by predators.
In San Luis Potosi Crotalus triseriatus aquilis shares the range
and habitat of mexicana. The pattern match of these animals is
quite remarkable. In life, C.t. aquilis is gray-brown with black speck-
ling and a mid-dorsal row of 28 to 40 (mean
= 34.5, Klauber, 1952)
almost rectangular red-brown irregular blotches. Ventrolateral black
blotches are also present. The W.W. Brown 1923 collection at
Alvarez for the MCZcontained 10 mexicana and 20 C.t. aquilis.
The color variability oi mexicana in the Nuevo Leon population
sampled could reflect the local variability of the model, Crotalus
lepidus, or a multiple model system. In that area C. lepidus varies
from ochre to blue in background color, with deep reddish-orange
on the ventral side of the tail, which is also apparent in C.t. aquilis.
Some mexicana from Nuevo Leon are clearly ringed, like Micrurus.
Micrurus fulvius is present on the Atlantic versant of the Sierra
Madre. The mobility of predatory birds, as well as the relative rarity
of the crotaline model, could affect the occurrence of mimicry in
mexicana.
There are two situations which the mimicry hypothesis is less
in
certain. The population of mexicana in Durango, Mexico, is rather
uniform in appearance and not clearly like any other snake found
there. Possible models which share the same canyons with mexicana
are C. lepidus, C. pricei. and C. willardi. The mexicana are most like
male C lepidus in that both are yellow-green to gray-green in back-
ground color with contrasting narrow dark crossbands. The mexi-
cana, however, have many more bands and the bands are usually
red-centered. The second situation concerns ruthveni. On the same
hills in southwestern Queretaro where ruthveni is locally abundant,
C. triseriatus is also abundant, yet ruthveni is a brightly ringed

animal. Micrurus fitzingeri is sympatric with ruthveni over much of
28 BREVIORA No. 466
itsrange, but appears not to be present at those localities. M.fitzin-

geri is ringed similarly to ruthveni, with up to 28 triads of more or
less equal annuli (Smith and Taylor, 1966). Again, the conspicuous-
ness of the red color may contribute to its stimulus value.
An made (Levene, 1953) that the areas
ahernate prediction can be
showing polymorphism some way spatially patchy. The habi-
are in
tat oi mexicana in Nuevo Leon could be considered patchy in that
there are, in close proximity, areas of pine forest, chaparral, open

desert, and natural and man-made meadows. There is, however, no
correlation of observed morph to microhabitat. There is some spa-
tial heterogeneity in alterna habitat also, with alterna found in both
riparian and desert situations. However, this does not correlate with
the distribution of the Z)/fl/>/ and fl//£'/-/7flmorphs.
Parallel selection processes for crypsis in both the "model" and
"mimic" could also explain the similarity of pattern in each situa-
tion. The blairi case, however, with the change of color pattern of
alterna to the more abundant and conspicuous model Agkistrodon,
even in the presence of C lepidus, argues strongly for something
otherlhan parallel selection.
DESCRIPTIONS AND DIAGNOSES OFTHETAXA

Lampropeltis alterna (Brown)
Ophibolus alternus Brown 1901
Lampropeltis alterna, Stejnegerand Barbour 1917
Lampropeltis blairi ¥\my 1950
Lampropeltis mexicana blairi, Gehlbachand Baker 1962
Lampropeltis mexicana alterna, Gehlbachand Baker 1962
Type specimen {\\o\oiy^Q): Acad. Nat. Sci. Phil. 14977

Type locality: Davis Mountains, Jeff Davis County, Texas
Description and Diagnosis: A moderately sized (to about m) 1
snake with a very distinct head and overall mottled grey color. The
that may
pattern is a series of white-edged black blotches or saddles
be red-centered. Alternating reduced markings may be present
between major markings. The iris of the relatively large eye is silver-
grey in color. The number of ventral scales is 210-232. The proxi-
mate spines on the hemipenes are ovoid in cross section and
approximately 0.7 mm long.
1982 LAMPROPELTIS MHX/CAXA SPECIES GROUP 29
Distribution: Specimens have been collected over the entire

Mapimian portion of the Chihuahuan Desert (Morafka, 1977).
These localities range from latitude 32° N (Worthington, 1974)
south to latitude 25° N(Tanzer, 1970).
Lampropeltis mexicana (Garman)

Ophibolus triangulus var. mexicanus Garman 884 1
Oreophis boulengeri Duges 1897

Coronella mexicana, Gunther 1900
Coronella leonis Gunther 1900
Lampropeltis mexicana, Blanchard 1921
Lampropeltis leonis, Blanchard 1921
Lampropeltis leonis, Loveridge 1924

Lampropeltis thayeri Loveridge 1924
Lampropeltis greeri Wehh 1961
Lampropeltis mexicana greeri, Gehlbachand Baker 1962
Lampropeltis mexicana thayeri, Gehlbachand Baker 1962
Lampropeltis mexicana mexicana, Gehlbachand Baker 1962
Type specimen (syntypes): MCZ 4652, 4653

Type locality: near Ciudad San Luis Potosi, Mexico
Description and Diagnosis: A moderately sized (to about m) 1
snake with a slightly distinct head and overall mottled grey to yellow
or brown color. The pattern is a seriesof white-edged black
blotches, saddles or rings that may be red-centered. Alternating
reduced markings may be present, usually ventrolaterally, between
major markings. The anterior tail blotch is enlarged ventrolaterally
and red extends onto or usually across the subcaudals. The iris of
the relatively large eye is yellow-brown in color. The number of
ventral scales falls between 1 90 and 2 1 2. The proximate spines of the
hemipenes are rhomboidal in cross section and are approximately

0.4 mm long.
Distribution: Specimens have been collected mainly from the
mountains surrounding the Saladan portion of the Chihuahuan
Desert (Morafka, 1977). The localities have ranged from about lati-
tude 25° N in the Sierra Madre Oriental (P. Bartlett, personal com-
munication) south to 21° (Duges 1897). A single desert locality
N
(D. Morafka, personal communication) may be a result of recent
man-altered changes in the habitat.
30 BREVIORA No. 466
Lanipropeltis rutliveni Blanchard

Lampropeltis ruthveni Blanchard 1 92 1
Lainpropeltis trianguluni arcifera, Williams 1978 (in part)
Type spedmen (ho\olype): USNM 46558

Type locality: Poirenaro= Patzcuaro?, Michoacan, Mexico
Description and Diagnosis: A moderately sized (to about 0.8 m)
snake with a slightly distinct black head and a uniformly ringed
pattern. The black rings are light-edged and red-centered and
extend entirely around the body. The small amount of background
color is a mottled tan to lime green. The iris is yellow-brown in
color. The number of ventral scales falls betwen 82 and 196.
1
L. ruthveni can be distinguished from the central Mexican milk

snakes L.t. nelsoni and L.t. arcifera on several features of external
morphology. head of L. ruthveni is distinct from the neck,

First, the
as in L. inexicana. Second, the ringed pattern of L. ruthveni is
uniform; the red is not much broader nor is the background color
much narrower than the black as in L.t. nelsoni. Third, there is no
tendency in any of the L. ruthveni examined for the black to extend
along the dorsal midline through the red, as is the case in both L.t.
nelsoni and L.t. arcifera (Smith, 1942; Williams, 1978). Fourth, the
black rings of L. ruthveni are bordered with a lighter color, usually a
pale lime green; this is similar to L. mexicana and is distinct from L.
trianguluni. Finally, the ventral scales of the L. ruthveni examined
were never greater than 196 (mean = 188), while the range of ven-
trals of L. t. nelsoni is 203-23 and of L. t. arcifera
1 is 197-217.
Distribution: The type locality is cited (Blanchard, 1921) as Patz-
cuaro, Michoacan, but this may be incorrect. The locality with the
specimen is Potrenaro, Michoacan. The base camp of the collector,
W.E. Nelson, in August 1892 was Patzcuaro (F. McCuUough, per-
sonal communication). Recent collections have been in the vicinity
of La Piedad, Queretaro (C. Lieb, J. Dixon, E. Wagner, personal
communication and personal observation), Morelia (UMSNH) and
Contepec, Michoacan (D. Armstrong and J. Campbell, personal
communication) and Tapalpa, Jalisco. All of the recent collections
have been in rocky, wooded uplands. The range of this species may
extend entirely across the Mexican Plateau.
1982 LAMPROt'l.l.TIS MEXICASA SPECIES (i Rmi> 31
SPECIMENS EXAMINED
Lampropeltis altenia. MEXICO: Coahuila: Cuatrocienegas
(FMNH 47090); Cruz Verde Mt.c. Saltillo(USNM 1108 19); Puente
de la Muralla c. Monclova (Garstka coll., spec). Durango: 26 mi. I
N San Juan del Rio (TCWC 36892). UNITED STATES. TEXAS;

Brewster Co.: hills N (MCZ 157763). Jeff Davis Co.:
of Study Butte
17.7 mi. N Fort Davis (TCWC
26181). Presidio Co.: 25 mi. W
Lajitas (UTA 7875). Val Verde Co.: W
of Comstock (UTA 2633); 8
mi. W of Comstock (UTA 2941); 8.5 mi. N of Comstock (UTA
8690); 32 mi. NW
of Comstock (TCWC 33759); 2 mi. of Com- W
stock (TCWC 30515); N of Comstock (MCZ 157764, 157765;
Garstka coll., spec); 15 mi. NE Del Rio (TCWC 26179); 1.5 mi. E
1
of Langtry (UTA 6680); 7.5 mi. E of Langtry (UTA 6681); N of

Langtry (UTA N of Langtry (UTA 8671); Langtry Loop
7188); 6 mi.
Road (MCZ 156175); mi. E of Langtry (Garstka coll., spec); 11
1 1
mi. N of Loma Alta (UTA 8668); c 10 mi. S of Loma Alta (UTA

7874); 0.5 mi. E of Pecos River on US 90
(UTA 8095); Roadside rest
c. (UTA 8568); Pecos River overlook (TCWC 26180).
Pecos River
NO SPECIFIC LOCALITY: (UTA 7969, 8179); (Wagner coll.. no.
19). CAPTIVE-HATCHED: Gravid female collected 8.7 mi. N of
Comstock (TCWC 33761 33763). CAPTIVE-BRED: Male parent
collected Brewster Co., Christmas Mts.; Female parent collected
Val Verde Co., c Langtry (MCZ 156271 156273); Both parents
collected Val Verde Co., c Langtry (MCZ 157724 157728; 156173,
156174; Garstka coll., 10 spec); Both parents collected no specific
locality (Texas) (MCZ
156259-156270, 157755-157758, 157760-
157762, 158326-158335; Garstka coll., 1
spec, UTA 8126 8129,
7873; Wagner coll., 1 , 5, 7, 1 9, 2 1 -25, 34, 44).
Lampropeltis me.xieana. MEXICO: Durango: 42 mi. S of Cd.

Dgo. (UCM 21061); 23 mi. S of Cd. Dgo. (Wagner coll., Gl. G2);
Canyon of the Rio Tunal (Garstka coll., 2 spec); Highway 40 at the
Rio Chico (LACM 107230, 107231);Barbara (Hous- Rancho Sta.
ton Zoo, spec);1no specific locality (San Diego Zoo, 9 spec).
Nuevo Leon: c Galeana, Linares-San Roberto highway (ITESM
2507, 2508); Galeana (TCWC 56823); 5 mi. SE of Galeana (TU
16483); c Los Lirios (ITESM uncataloged); Ojo de Agua c Galeana
(FMNH 30819-30821); c La Angostura (Garstka coll., 3 spec); no
specific locality (BMNH 146-1 4 10). San Luis Potosi: AKare/
32 BREVIORA No. 466
(MCZ 19022 19025, 24976 24979; AMNH 77602; USNM 120823);

c.Armadillo de los Infantes (Wagner coll., spec.); c. Rioverde (KU
1
85010); c. Cd. San Luis Potosi (MCZ 4652, 4653); 52 mi. WNW
of
Cd. San Luis Potosi (CM 59980). Tamaulipas: Miquihuana (MCZ
19551). CAPTIVE-BRED: Both parents collected Mexico, Du-
rango, no specific locality (MCZ 157754; Garstka coll., spec;1
Wagner G3, G5). Both parents collected Mexico, Nuevo Leon,

coll.,
c. La Angostura (MCZ 56274- 56277, 57766; also Garstka coll.)
1 1 1
Lampropeltis ruthveni. MEXICO: Michoacdn: Club Campestre

at Morelia(UMSNH uncataloged); Potrenaro= Patzcuaro?(USNM
46558). Queretaro: Canyon of the Rio Galindo c. Amealco (MCZ

161010-161012; Mexican govt, coll., Agencia Forestal y de la Fauna
Z-06586, Z-06587; Garstka coll., 5 spec; Wagner coll., 9 spec; 5
spec, released at site of capture). Jalisco: Mts. W
Zacoalco (KU
155528); Tapalpa (SDMNH 46093); 6.5 mi. E Tapalpa (LACM
37307).
ACKNOWLEDGMENTS
I would most like to thank E. E. Williams for his patience and
presence throughout this study. T. Fritts was instrumental in the
early stages of the study, as were many discussions with D. F. Retes,
E. Wagner, and others. G. C. Mayer criticized several drafts. D.
Crews, J. Dixon, M. Scott, and two reviewers also commented on
the manuscript. Access to specimens was provided by: J. Alvarado,

Universidad Michoacana de San Nicolas Hidalgo, Morelia, Michoa-
can, Mexico (UMSNH); P. Bartlett, Instituto Tecnologico y Escu-
ela Superior de Monterrey, Monterrey, Nuevo Leon, Mexico
(ITESM); R. Bezy & J. Wright, Los Angeles County Museum

(LACM); J. Dixon, Texas Cooperative Wildlife Collection (TCWC);
W. Duellman, University of Kansas (KU); H. Dundee, Tulane Uni-
versity (TU); T. Fritts, San Diego Museum of Natural History
(SDMNH); A. G. C. Grandison, British Museum (Natural History)
(BMNH); W. R. Heyer, United States National Museum (USNM);
D. Hoffmeister, University, of Illinois Museum of Natural History
(UIMNH); P. Maslin, University of Colorado Museum (UCM); C.
J. McCoy, Carnegie Museum (CM); W. Pyburn, University of
Texas Arlington (UTA); T. Van Devender, University of Arizona

at
(UA); and R. Zweifel, American Museum of Natural History

(AMNH). Access to specimens was also provided by numerous zoo
1982 LAMPROPELTIS MEXICANA SJ>F.CIF.S (ikOlP 33
and private Bacon (Zool. Soc. San Diego),

collections, especially J.
J.Murphy (Zool. Soc. Dallas), H. Quinn, and B. Tryon (Zool. Soc.
Houston), E. Wagner (Zool. Soc. Seattle) and D. F. Retes. Botani-
cal material was identified by J. Henrickson, T. Wendt, and M.
Donaghue.
in Mexico was issued by Lie. M. L. Cossio-
Permission to collect
Gabucio, L Ibarrola-Bejar, and Lie. M. Gonzalez-Escamilla, all
Lie.
of the Agencia Forestal y de la Fauna.
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* r^M
'ARVARD
UMIVERSiTY
B R E V I O R A
us ISSN 0006-9698
Cambridge, Mass. 30June1982 Number 467
THREE NEW SPECIES OF THE ANOLIS PVNCTATVS

COMPLEX FROM AMAZONIAN AND INTER-ANDEAN
COLOMBIA, WITH COMMENTS ON THE EASTERN
MEMBERS OF THE PVNCTATUS SPECIES GROUP
Ernest E. Williams'
Abstract. New species of the Anolis punctaius group are described from central
and eastern Colombia: A. vaupesianiis from Comisaria of Vaupes, A. hiiilae from the
Departments of Huila and Tolima, A. santamariae from the south slope of the Sierra
Nevada de Santa Marta. The characters useful in discriminating the eastern members
of the Anolis punciaius species group are discussed and evaluated, and the species
themselves are diagnosed.
INTRODUCTION
Important new collections from Colombia have yielded Anolis
Those dealt with here are allied to Anolis
that require description.
punctatus of Amazonia and the Atlantic Forest of Brazil, and, like

the other recently described members of the punctatus group
(Williams, 1965, 1974), they occur areas peripheral to the range of
in
that central and widely dispersed species. The three new taxa thus
fill in blank areas in the known range of the complex. However, like
many others of the punctatus species group, excepting only the

central widespread and best-known species punctatus, these are
species almost without distributions or ecologies. There are so few
localities from which they are known, so little is known about their
biology, that they are question marks rather than known compo-
nents of fauna. They testify primarily to our basic ignorance of
South American lizards.
'Museum of Comparative Zoology. Harvard University, Cambridge, Massachusetts

02138.
2 BREVIORA No. 467
I first describe the Amazonian taxa. Since the three best
specimens are from the Comisaria of Vaupes, 1 call it:
Anolis vaupesianus, new species

(Figs. 1-5 and Map)
Type: MCZ 156309 (formerly UTA 3626). adult male.
Type locality: Timbo, a Cubeo village ca. N 70 on the Rio
1 W
Vaupes, E of Mitii, Vaupes, Colombia, William F. Pyburn coll., 24
July 1972.
Paratypes: COLOMBIA: Vaupes. UTA 6850, 154592 MCZ
(formerly UTA 5051)' same locality as type. J. K. Salser, Jr. coll.;
ICN 4542, Cano Ti near Pamopeta, W. W. Lamar coll.; UTA 10283,
Wacara, Lois Lores and Marilyn Cathcart coll. Amazonas. MCZ
79655, lower Rio Apaporis, L Cabrera coll.; MCZ 112098, Tio
Miriti-Parana, La Providencia, F. Medem coll.
Description (paratype variation in parentheses). Head: Head
scales moderately sized, weakly (sharply) keeled toward tip of snout,
smooth posteriorly. Ten (8-1 1) scales across snout between second
canthals. Frontal depression distinct, scales within it smaller than
those anterior to it (not or not conspicuously so). Eleven (9-10)
scales border rostral posteriorly. Anterior nasal scale divided (not
divided), and inferior nasal scale in contact with rostral. Eight
it
(6 between supranasals. Rostral area swollen, protuberant,

7) scales
rostral extending well beyond mental on male (not swollen or
protuberant, rostral not extending much beyond mental in female).

Supraorbital semicircles separated by one minute scale (or by one
or two larger scales or narrowly in contact), separated from the
supraocular disk by one row of granules. Supraocular disk ill-
defined (better-defined), grading into surrounding granular scales.
One elongate supraciliary followed by minute granules. Canthus
second largest. Six (5-7) loreal
distinct, five (5 6) canthal scales, the
rows counting down from the second canthal, the ventralmost
largest.
Temporals and supratemporals granular, the latter grading into
around the interparietal. An ill-defined and tapering
larger scales
double intertemporal row of somewhat enlarged scales. Inter-
'This is a specimen referred to as A. puiulaius by Greene (1977).

1982 THREE NEW COLOMBIAN A SOUS
Figure 1. Anolis vaupesianus. n. sp. Type $. MCZ 156309. Dorsal view ot head.
Figure 2. Anolis vaupesianus. n. sp. Type Q, MCZ 156309. lateral view ot head.
BREVIORA No. 467
Figure 3. Anolis vaupesianus. n. sp. Type $, MCZ 156309. Ventral view of head.
1982 THREE NEW COLOMBIAN ANOLIS
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BREVIORA No. 467
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1982 THREE NEW COLOMBIAN /I, VO/./5 7
parietal much larger than ear (larger), separated from the supra-
orbital semicircles by two (3) rather large scales. Scales surrounding
interparietal largest anterolaterally, posteriorly grading through an

area of minute scales into the dorsals that are as large as the
temporals.
Suboculars weakly keeled, in contact with supralabials. Seven to
eight (7 9) supralabials to center of eye.
Mental semidivided, each half distinctly wider than long, in
contact with four scales between the keeled sublabials. Three (5)
sublabials in contact with infralabials. Gulars small, smooth,
increasing in size laterally.
Trunk: Dorsals granular, subequal, smaller than the weakly

keeled ventrals which are subimbricate (juxtaposed), in transverse
rows.
Dewlap: Moderate in male, not extending posteriorly much
beyond axilla. Lateral scales elongate, larger than ventrals, in close-
packed rows. Edge scales shorter, crowded, subimbricate. Vestigial
in female.
Limbs: Largest scales on forelimb unicarinate, on digits weakly

multicarinate. On hind limbs scales in front of thigh and lower leg
weakly unicarinate, on back of thigh and lower leg granular.
Supradigital scales weakly multicarinate. Twenty-five (22 26) la-
mellae under phalanges ii and iii of fourth toe.
Tail: Weakly compressed. No dorsal crest. No verticils. Scales
above weakly carinate. A double middorsal row. Scales behind vent
at most very weakly carinate, becoming distinctly so posteriorly.
Postanal scales not well defined in male, absent in female.
Size: Male mm
snout to vent; largest male paratype: 78
type: 82
mm snout to vent. Female paratype: 75 mm
snout to vent.
Field notes and color in life: F. Pyburn presents the following
notes on the male holotype: "24 July 1972 I collected a dark green
anole on the base of a small tree in village (Timbo) about one hour
before dark. Dorsum of head, body, legs dull uniform green with
scattered blue flecks; venter grey; dewlap skin black; scales of
dewlap grey. Lizard turns dull brown when handled. Has fleshy keel
on nape and along back."
For MCZ 154592, a male, the notes below are abridged from
those of J. K. Salser, Jr.: "Anolis sp.; Timbo (Vaupes), Colombia;
March 20, 1974: very hot noon-time; caught in an aged avocado tree
8 BREVIORA No. 467
- scaled tree and jumped from its top in attempt to escape. Blue . .
dots dorsolaterally placed in stripes from vertebra down, angling

down to lower sides; changes from dark green to green-brown
mottled very quickly; eyes golden, eyelid border yellow..."
For the female paratype, MCZ 112098 (Fig. 5), from the Rio
Miriti-Parana, Federico Medem has provided the following infor-
mation: "Caught 8:30 a.m. on underbrush, one meter high, in open
country. Gular sack small, greenish. Iris golden. Intense green on
. .
body with lateral small blue spots. Middorsally 16 squarish sand-

colored spots. Dark transverse bars on tail. Venter light gray with
dark vermiculations."
For MCZ 79655, a male, Medem reports the color as: "Dorsally
-
green with lateral blue spots; dewlap 'morado' violet, mulberry-
colored."
Comparisons: Very few specimens of A. punctatus are recorded

from Colombia. The nearest to any of the type series of A.
vaupesianus are two recently collected specimens in the Paris
Museum (MHNP-1978-3086-87) from Hameau Kuira, Rio Agara
Parana, a tributary of the Rio Putumayo, in Amazonas Territory.
Both are males with strongly keeled ventrals, dorsum an unpat-
terned dark purplish and dewlap with light skin with minute
dispersed melanophores and purplish scales.
The A. vaupesianus most nearly adjacent to these is MCZ 79655
from La Providencia, Rio Miriti-Parana, a tributary of the Rio
Caqueta, also in Amazonas. It is unfortunately a female. MHNP-
1978-3086 has therefore been compared with the topotypic paratype
UTA 6850, which is of the same size and sex. The dewlap in the
paratype contrasts strongly with the Amazonas punctatus in its

intensely black skin and white scales, and the ventral scales in
vaupesianus are smaller and decidedly more weakly keeled. No
other scale differences are evident, but the color and pattern of the
Mitu paratype as preserved are strikingly different from anything
observed in punctatus: it is brown, strongly blotched with darker,
the dorsal blotches tending to form transverse series across the back.
Anteriorly, a dark V, the apex backward, formed of somewhat

irregular blotches, reaches the middle of the orbit on each side. A
less well defined V in medial to this, reaching to the tops of the
orbits.
This very distinctive color and pattern are not, however, seen in
the remainder of the type series of vaupesianus which, as preserved.
1982 THREE NEW COLOMBIAN /I AY;/./.S 9
are not strikingly dissimilar to A. pimctatiis. At best the distinctive
pattern of UTA6850 represents an element of the pattern repertoire

of A. vaupesianus not present in A. punctatus. It clearly will not
serve to distinguish most specimens.
Habitat: The few notes on vaupesianus available give little
indication of its ecology, except that it is associated with trees. The
was caught in a village probably indicates that it is
fact that the type
an edge animal, not characteristic of the dark forest.
The second species was first discovered in the Departamento del
Huila. It has therefore been called:
Anolis huilae, new species

(Figs. 6-9 and Map)
Type: ICN 3725, an adult male, collected by Helen Chin, October,

1976.
Type locality: Herberto Herrera's coffee plantation, Palestina,

Huila, Colombia.
Paratypes: COLOMBIA: Huila. KU 169830-31, Parque Ar-
queologico San Agustin, 3 km SW San Agustin, 1750 m; ICN 3726,
MCZ 159121-22: Palestina; MCZ 156305: Quebrada La Cascajosa,
Parque Nacional Natural La Cueva de los Guacharos (northern
boundary); INDERENA R-0297, -0562, -0665. MCZ 156306,
156316: Parque Nacional Natural La Cueva de los Guacharos; ICN
4461-62: 62 km (by road) NW
San Isidro de Isnos, 1940 m. Tolima.
ICN 3732: Amaine; ICN 3727-31. 3733, INDERENA (numbers not
now available, formerly UVMP 4400. 4404), MCZ 159112-117:
Cajamarca; Cali (Univ. Valle, Depto. Biol.) UVC 38, 191, ICN
3735-36, MCZ 159119-120: Juntas; ICN 3734, MCZ 159118:
Llanitas, 10 km N Ibague.
[Referred specimen: Meta: ICN 4541 ca. 3-4 km NNE of
Manzanares.]
Description (paratype variation in parentheses). Head: Most head
scalessmooth, slightly concave, a few convex, obtusely keeled. Ten
(8-11) scales across snout between second canthals. A few narrow
scales in the frontal depression much smaller than surrounding
scales. Eight (7-9) scales border rostral posteriorly. Anterior and
inferior nasal scales in contact with rostral. Seven (6-7) scales
dorsally between nasals. Snout not protuberant in male.
Supraorbital semicircles broadly in contact with each othei and
with the supraocular disks. Supraocular disks with few (several)
10 BREVIORA No. 467
Figure 6. Anolis huilae. n. sp. Paratype 5, MCZ 159015. Dorsal view of head.
Figure 7. Anolis huilae. n. sp. Paratype 5, MCZ 159015. Lateral view of head.
1982 THREE NEW COLOMBIAN ANOLIS 11
Figure 8. Anolis huilae, n. sp. Paratype $, MCZ 159015. Ventral view of head.
large smooth scales. One elongate supraciliary on each side followed

by granules. Canthus blunt, ca. 8 canthal scales, the second and
third largest. Five (4-5) loreal rows, counting down from the second
canthal, subequal.
Temporals slightly smaller than the supratemporals, both granu-
lar. No
differentiated intertemporal line of enlarged scales. The
scales surrounding the interparietal abruptly larger, those anterior
and lateral largest (not so in female). Interparietal equal to (smaller
12 BREVIORA No. 467
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1982 THREE NEW COLOMBIAN /l,VO/./5 13
or larger than) ear, separated from the supraorbital semicircles by

2-3 (0-2) scales. Large scales posterior to interparietal followed by
minute granules on nape, distinctly smaller than the granular
dorsals.
Suboculars keeled, in contact with supralabials, continued
posterior to the eye by two distinct (less distinct) rows of smaller
scales, anteriorly meeting canthals. Seven (5 7) supralabials to
center of eye (supralabial five is much the longest in one paratype,
an obvious anomaly; in the type, the supralabials are nearly
subequal).
Mental semidivided, in contact with three ( 1
4) scales between the
very large, smooth sublabials. Three sublabials ( -4) in contact with
1
infralabials. Central throat scales small, swollen, oval, becoming
abruptly longer laterally alongside the sublabials.

Trunk: a few middorsal (2-4) rows indistinctly enlarged, weakly
or distinctly keeled, grading into granular flank scales. Ventrals
much larger than largest dorsals, smooth, imbricate or subimbricate
(juxtaposed in female), not in distinct transverse rows (in transverse
rows).
Dewlap: Large in male, extending to midbelly (absent in female).
Scales at edge larger than ventrals, smooth. Lateral scales smooth,
in double rows separated by naked skin.
Limbs: Limb scales weakly uni- or multicarinate. Supradigital
scales multicarinate. Twenty-one (22) lamellae under phalanges ii
and iii of fourth toe.
Tail: compressed, without crest. Enlarged postanals
Slightly
present (absent A double row of keeled scale middorsally
in female).
on tail. Scales behind vent smooth.
Size: The male type is 75 mm
in snout-vent length. The largest
male reaches 82 mm.
Color in life: Helen Chin reports topotypic animals as 'parrot
green' in the trees. Her color transparencies show tan and reticulate
animals after capture.
Color notes are available for two MCZ specimens. MCZ 156405.
a male: "Dorsum
bright green with yellow spots (seven large spots
along the spine). Tail banded green and black. A white dot inside a
black ring on side of neck behind the ear. Eyelid yellow, iris grey.
Chin grass-green. Dewlap yellow with six green stripes, these
bordered by numerous black dots. Rest of underparts greyish."
MCZ 156306, a female: "Dorsum with wide tan stripe from back
14 BREVIORA No. 467
of head to halfway down tail, at which point it becomes speckled

with black dots. Rest of back grey-green, darker next to tan stripe,
and lighter with dark patches on the sides. Legs banded black and
light green. Head light green with dark around eyes. Tip of tail
bronzy. Venter yellow with few small black spots."
Both these specimens are near-topotypes. Palestina is a settlement
at thebeginning of the 22 km trail leading to the Parque Nacional
Natural La Cueva de los Guacharos.
The colors in life of the Kansas specimens from Parque
Arqueologico de San Agustin are described rather differently
(although not in the same detail): "At night dorsum pale green with
-
brown reticulations. Dewlap cream with grey-bordered white

streaks. Iris creamy-grey. Tongue orange."
Notes are at hand also for two females collected by Pedro Ruiz
and J. D. Lynch near San Isidro de Isnos (notes by Lynch):
ICN 4461: "Sleeping on leaf in forest near stream. Iris yellowish.
Ground brown barring; greenish cream spots
color dull green with
on flanks; tail barredbrown and medium brown with cream edges;
venter dirty cream with brown spots. Throat the same but with
greenish cast."
ICN 4462: "Sleeping on Ortega, 3 m above stream at edge of corn
field. grey brown. Tongue pale orange yellow."
Iris
Dennis M. Harris has provided notes on the series from Tolima:

"Male: Dorsum pale leaf-green, spotted most vividly on the sides
by 1-2 mm diameter circular dots of greenish-yellow. Head slightly
brownish. Eyelid yellow. Nape with a lateral spot of blue-green to
black, sometimes surrounded by a yellow area, sometimes with a
barred or not with bands equal in width to the
light center. Tail
green. Venter white to grayish. Dewlap skin pale yellow, scales
cream, in rows that are edged with black spotting.
Male may change to a deep brown with vivid yellow spots. It does
so when handled and possibly as a response to another male. The
one brown male seen in the field was never a green one, and had a
vertically flattened body and extended dewlap. (The cream scales of
the dewlap may become drab greenish in the brown-phase male.)
Female: Banded or striped, light or dark brown. The stripe is
Crayola brown with dark brown flecks. There is no color change
except for minor tone changes."
Color notes have been made available by W. W. Lamar for the
single animal from Manzanares, Meta: "Dorsum dark gray with
1982 THREE NEW COLOMBIAN /4,VOL/5 15
numerous gray-green spots. Mid-dorsum brighter than rest of

light
the back. More green towardthe center of the flanks. Tops of
forelegs gray-green with lighter yellow-green elbows and feet. Hind
legs with 3 light bands above knee. Three broken bands below knee.
Tail pale green with 11-12 dusty bands. A black spot with an
aquamarine center on the side of the neck. Yellow eye ring and
snout aqua-blue-green, darker between the eyes. Venter pale yellow
with dark gray spots on the sides, brighter yellow near vent, broad
portion of tail, and inguinal area. A patch of aqua-blue on tail 10-25
mm from vent, rest of tail dark gray. Dewlap an overall aqua green-
blue striped with pale green and dotted with black. Spaces between
stripes are pale yellow. Nostril yellow. Tongue yellow."
Comparison of these notes for the Meta specimen described
shortly after death with the detailed account from Huila and Tolima
shows various discrepancies. Color differences in Anolis, however,
may be described differently by different individuals or by the same
person at different times. This, plus the capacity for color change,
suggest caution in attributing taxonomic significance to color
differences of the sort seen here. (See also comment below.) There
are no scale differences.
Color as preserved: Most male .4. huilae are strikingly distinctive
because of the bold lateral spotting, plus the lines of dark spots on
the dewlap. Some specimens display a more muted spotting. This is
especially true of the individual from Manzanares, Meta. The latter

seems because the spotting on the dewlap is
at first distinguishable
not in lines but quite random. Some specimens from Tolima,
however, have the spotting on the dewlap irregular also. The Meta
animal does not, on present evidence, fully match the type series;
since italso geographically peripheral (on the east side of the
is
Andes, rather than in the Magdalena Valley) I have not made it a

paratype. With only one specimen and no structural differences it is
not permissible to describe it taxonomically.

Habitat: Dennis Harris reports: "found on relatively smooth bark
tree trunks usually, between and 7 meters. Males almost
1
exclusively there, females sometimes on bushes nearby, m high. 1
Usually only single male/ tree trunk (25 cm+ diameter). Shrubs
seem to be only peripheral environments. Not found on very
smooth-bark guayaba (guava) trees. Not found on trunks with
heavy moss growth. Common right in town of Cajamarca and along
edges of road. Males more frequently seen and more of them
16 BREVIORA No. 467
caught. Only time male and female seen together on tree trunk they
were copulating. Usually seen head down, out {\'\ke frenatus) on
trunk. At Anaime Prionodactylus argulus found at same site. At
Llanitas, 10 km from
Ibague, Anolis antonii and A. huilae; at
Juntas, much higher, Phenacosaurus and A. huilae.'"'
MCZ156316 was found below a cave-hole of Rupicola rupicola
among defecated seeds. Apparently a food item, it was in relatively
good condition and quite recognizable.

Comparisons: A. huilae, although obviously a member of the
punctatus species group, is not obviously closely related to
any
other presently described member. The color pattern of males is
highly distinctive.
A third species comes from the southern slopes of the isolated
Santa Marta range. It is to be called:
Anolis santamartae, new species

(Figs. 10-13 and Map)
Type: CAS 113922, adult male.

Type San Sebastian de Rabago, Sierra Nevada de Santa
locality:
Marta, Cesar, Colombia (10°4" N, 73° 16" W), Borys Malkin coll.,
1-10 April, 1968.
Paratypes: same data as type: CAS 113924, 156311. MCZ
Description (paratype variation in parentheses). Head: Moderate-
lyelongate. Head scales rather large, smooth, flat except for smaller
bluntly keeled or swollen scales at tip of snout. Six (4 5) scales
across snout between second canthals. Frontal depression distinct,
the scales within it larger (some smaller) than those anterior to it.
Anterior and inferior nasal scales in contact with rostral. Four (5-6)
scales dorsallybetween supranasals. Rostral area protuberant in
male, overhanging lower lip (not so in female).
Supraorbital semicircles broadly in contact with each other and
with the supraocular disks. Supraocular disks well defined, com-
prised of 8 to 10 smooth scales. Remainder of supraocular area with
large granules. Canthus distinct, four or five canthal scales, the
second and third (or first and second) largest. Five (4-5) loreal rows,
the lowermost largest. Temporal and supratemporal scales granular,
larger at the angle of the mouth an at the angle between temporal

and supratemporal regions. Scales surrounding the interparietal
rather abruptly larger, largest laterally, grading into dorsals
1982 THREE NEW COLOMBIAN ANOI.IS 17
Figure 10. Anolis saniamartae. Type $, CAS 113922. Dorsal view of head.
Figure II. Anolis saniamartae. Type S- CAS 113922. Lateral view of head.
BREVIORA No. 467
Figure 12. Anolis santamartae. Type $, CAS 113922. Ventral view of head.
posteriorly. Interparietal much larger than ear, broadly in contact

with the supraorbital semicircles.
Suboculars weakly keeled, in contact with supralabials, anteriorly
separated from the canthal ridge by two (1-2) scales, posteriorly
grading into the temporals. Seven supralabials to the center of the
eye.
Mental semidivided (mental region abnormal in MCZ
15631 1), in
contact with two large scales between the well-developed sublabials.
1982 THREE NEW COLOMBIAN ANOLIS 19
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20 BREVIORA No. 467
Figure 14. Distribution of the species described herein, t : Anolis santamartae:

O: Anolis huilae: and : Anolis vaupesiamis.
1982 THREE NEW COLOMBIAN /I /VO/./.V 21
Four (3) sublabials in contact with infralabials. Gulars rather large,

swollen, increasing in size laterally and anteriorly.
Trunk: Ca. four to six middorsal rows flat, weakly keeled,
subimbricate, much larger than flank granules but grading into
them. Ventrals still larger, smooth or obtusely keeled. Chest scales
distinctly keeled.
Dewlap: Large in male, only indicated in female. Scales in close
set rows, elongate and narrow, except at edge where they are flat
and smooth.
Limbs: Scales on limbs uni- to multicarinate, those on digits
multicarinate. Eighteen (19 21) lamellae under phalanges ii and iii
of fourth toe.
Tail: Compressed. Scales weakly to strongly carinate. A double
middorsal row. No tail crest. No verticils. Scales behind vent

carinate. Post-anals large (broken up on one side type, absent in
female.)
Size: Male type: 50 mm, snout to vent. Male paratype: 55 mm.
Female paratype: 53 mm.
Color as preserved. Female paratype: Gray-brown. A light line
from suboculars to above ear. A narrow dark interorbital bar. A
series of ovoid or rhomboidal dark blotches paravertebrally.
Irregular oblique dark streaking on flanks. Limbs irregularly

banded. Tail very weakly banded. Throat and belly smudged with
darker.
Male type: Gray. A light line from suboculars to above ear.
Another light line from lower edge of ear along lateral edge of
dewlap onto underside of upper arm. Weak traces of blotching
dorsally. Irregular oblique dark streaks on flanks. Limbs and tail
with very weak and obscure markings. Underneath smudged with
brown. Dewlap skin white, scales very light brown.
Male paratype: A dark interorbital bar. A light stripe from
suboculars to above ear, another from below ear to shoulder.
Middorsum and dorsum of tail obscurely marked with light and
dark. Flanks with oblique dark streaks. Limbs smudged above.
Belly with irregular dark markings. Throat with vaguer markings.
Habitat: San Sebastian is at the border between the Departments
of Cesar and Magdalena, and the specimens may have been
collected in either Department. The elevation is at, or above, 2,000
m. There are no ecological notes.
22 BREVIORA No. 467
DISCUSSION
Comparison oi santamartae and the other two new species of the
punctatus complex with each other and with the other already-
described members of the complex could be made in tabular form,
as in Williams (1974). Here, however, I
prefer to discuss and
elevuate the characters on which I
recognize the several species. I
begin with the color differences.
Color in Life
in life of
For this purpose the few detailed descriptions of color
Anolis punctatus must be quoted.
Hoogmoed (1973) has a description for /I. punctatus 'punctatus'^
from Surinam: "Color above with sky-blue dots,

in life grass-green
eyelids purplish with a yellow rim. Chin bluish,

throat greenish.
Belly white to purple-green. In bright sunlight the hzards

are purple
with sky-blue dots. Dewlap in males yellow ochre with rows of white
scales; in females the scales on the dewlap are yellow. Iris orange-
brown."
A parallel description is provided by Dixon and Soini (1975) for
A. punctatus 'boulengeri' from the Iquitos area in Peru: "The dorsal
coloration of adults is leaf-green with six to seven very narrow white
cross-bands confined to the upper body. The lower sides are spotted
with minute white dots, and the limbs are spotted with minute dark
brown Occasionally specimens are deep purple dorsally,
spots.
especially after being in captivity for a few hours.
The white cross-
bands and white spots are very distinct when the latter color
prevails. Male dewlaps are orange to reddish-orange with the

enlarged lateral scale rows light yellow."
P. E. Vanzolini has provided me with a translation of the
unpublished field description of Helmut Sick of an A. punctatus

'punctatus' from the Atlantic Forest in Brazil (Rio Itaunas, Espirito
Santo): "General body color alive: grayish sepia, dewlap chrome
yellow. Freshly killed: moss green, eyelid yellowish green, dewlap
chrome yellow with stripes composed of whitish green spot, chin
whitish-green. Underparts greenish-white. Digits and
tail with
'Western specimens oi punctatus with keeled ventrals are usually called boulengeri
O'Shaughnessy whether as subspecies or species. See Williams and Vanzolini (1980)
for evidence that 'boulengeri' is a morph and not a valid taxon.
1982 THREE NEW COLOMBIAN /lA'OZ.AS- 23
vestiges of dark brown crossbands. Posterior two-thirds of tail
grayish sepia. Flanks with lines of white spots...." Vanzolini

himself describes dewlap colors for two additional specimens:
(Santo Amaro dos Brotas, Sergipe): 'pure yellow'; (Lago Mapori,
Rio Japura, Amazonas): 'rusty orange.'
No color descriptions from living specimens are available (or A.
santamartae, A. caquetae, A. deltae, or A. nigropunctatus. The
color descriptions of A. vaupesianus and huilae have been given
above.
There are serious problems in the comparisons of color and
pattern in the of this complex. The few good reports are
members
for the local populations of widely scattered regions or for
individual specimens. This would be a difficulty in any taxonomic
group. In Anolis an additional problem exists: color repertoire may
be extensive and tends to be especially so in the green species. In the
latter, the difference between light and dark phases may be extreme,
and there may be in life intermediate patterns or colors that are only
briefly or occasionally seen. (This may have been true, for example,
of the Meta specimen referred to A. huilae above.) Preserved
specimens compound the dilemma already posed. Pattern or lack of
pattern may be due entirely to the preservative, or to its strength.
Specimens done in strong formalin tend to a muddy uniformity;
many of the older specimens in collections are of this type. It is
fortunate that none of the specimens considered in this paper are of

so inferior a quality. Nevertheless, they have been preserved by
different individuals with different preferences in technique. The
preserved specimens are therefore only minimally comparable.

But even the available descriptions of color in life are imperfectly
comparable. They were not made to be checked against one
another, and they therefore do not stress quite the same details.
They were never checked against the same color standards.
Yet it is quite clear that enough information exists to assure us
that differences exist. While no possibility exists of entirely avoiding
error, some plausible conclusions may be drawn.
Color in anoles is best discussed by breaking it into components.
It is a fact about Anolis in general that dewlaps are least modified by
color change and most readily interpretable in preserved specimens.

This fact quite accords with the view (e.g.. Rand and Williams,
1970; Williams and Rand, 1977) that the dewlap is an intra- and
interspecific signal which should have high constancy in order
24 BREVIORA No. 467
always to convey the same message. (A caveat: this rule may have
less validity in mainland Anolis than it appears to have in the species
of the islands.)
In the present case we have the following evidence of dewlap
color:
punctatus pimctatus (Surinam)

— yellow ochre with white scales (Hoogmoed). The other reports
from the range of subspecies (Sergipe, Espirito Santo) stress a
yellow color also.
punctatus 'boulengeri'
— orange to reddish-orange (Dixon and Soini). Clearly a deeper
shade than in punctatus punctatus, but not sharply different.
Preserved specimens of both taxa show only light skin and
scales.
vaupesianus
— 'black' (Pyburn); 'mulberry red' (Medem) presumably on the
authority of the collector: 'greenish' in the female; Medem, in

this case himself the collector. There is an obvious discrepancy
here, but we are evidently dealing with a dark dewlap, and the
dewlap skin in all preserved specimens is darkly pigmented.
Fromthese notes it is clear that vaupesianus is sharply distinct in
dewlap color from the animals that in size and morphology it most
resembles, western A. punctatus, and from A. huilae. The dewlap of
the latter, with dark spots placed in lines or randomly, is quite
unlike any of the others. Insufficient information exists for any
really useful discussion of dewlap color in the other taxa.
As regards body color, I am very uncertain what can be inferred.
From the descriptions of Hoogmoed for Surinam and of Dixon and
Soini for Peru, it would appear plausible that a purple phase exists
inwestern A. punctatus which does not occur in A. vaupesianus, but
Vanzolini has not noticed such a purple phase in the punctatus of
Amazonia and eastern ^xdizW punctatus that he has caught. Absence
of purple may therefore not be fully diagnostic for vaupesianus.
Other aspects of the color of vaupesianus involve tints and special
aspects of its color repertoire which again are of uncertain value.
Size, body, and dewlap color seem safer guides to identification. For
A. huilae we do not know enough about color repertoire to evaluate
the colors in life that we do know. The bold flank spotting seen in
huilae may be useful, like the nape spot, but neither are obvious in
females and are not mentioned in some of the descriptions of colors
1982 THREE NEW COLOMBIAN /I,Vr;/./5 25
in See above for discussion of the specimen from Meta. For

life.
nigropunctatus, caquetae, santamariae. and deltae, discussions of

preserved body color are of very dubious value. Are the spots of
preserved nigropunctatus surely of diagnostic value? There is at
present no way to tell. The light line in santamartae from suboculars
to ear seems distinctive.
It is less evident in the
type than in the other
darker specimens. It may or may not be readily visible in life. For
these last four species body color cannot be disregarded but
determination of specimens must depend upon size, scale characters,
and locality. Let us look at these features in sequence.
Size
A. vaupesianus, A. huilae, and A. nigropunctatus are all within

the size range oi A. p. punctatus (70-80+ mm
snout to vent length).
A. dissimilis, A. caquetae, A. santamartae, and A. deltae are, on
present evidence, smaller (50-60 mm) (Table 1). (1 include in these
tables and discussions all described members oi\.\\t punctatus group
east of the Pacific slopes of the Andes.)
Table I. Size oi puiutatiis group anoles from east of the Andes and from the
inter-Andean valleys.
Maximum male snout to \ent length
(in mm.)
punctatus 85
iransversalis 83
vaupesianus 82
huilae 82
jacare lli
nigropunctatus 72
deltae 58
caquetae 57
dissimilis 56
santamartae 55
Scale characters
The more significant of these may be examined one by one:

Ventrals: In Anolis it is probable that smooth ventrals are
26 BREVIORA No. 467
primitive, keeled ventrals the derived condition. (The argument

derives from the apparently more frequent correlation of smooth
than keeled ventrals with other characters considered primitive; this
is not a strong argument.) Most pimctatus group species have
smooth ventrals, but a few have strong keels on the venter. Table 2
again lists the conditions of the several species east of the Andes or
in the inter-Andean valleys.
Table 2. Ventral scales in punctatus group anoles from east of the Andes and
from the inter-Andean valleys.
punctatus smooth or weakly to strongly keeled
vaupesianus weakly keeled

transversalis smooth
huilae smooth
jacare smooth
nigropunctatus smooth
deltae smooth
caquetae weakly keeled

dissimilis smooth
santamartae smooth or weakly keeled
Nostril-rostral relationship: This character is somewhat variable

within species, but within a narrow range. The punctatus group
shows only a fraction of the possible range, and each species shows
only a segment of this restricted range. The relevant conditions and
the species in which these are seen are listed below:
1) Circunmasal scale (=nasal) directly in contact with rostral:
nigropunctatus, jacare.
2) Circunmasal scale separated from rostral by a small round
undifferentiated scale: nigropunctatus, dissimilis.
3) A differentiated anterior nasal scale present and in contact with
rostral (the anterior nasal is characteristically triangular and
overlaps posteriorly part of the circunmasal): deltae, huilae.

4) Anterior plus an inferior nasal in contact with rostral: puncta-
tus, houlengeri, transversalis, vaupesianus, caquetae, huilae,
santamartae.
It is noteworthy that, while a species may exhibit two of these
four categories, these are always adjacent ones and no species is
1982 THREE NEW COLOMBIAN /I ;VO/./.9 27
known to range the full gamut. This character will assist in
discriminating species.
Snout protuberant or not in males: This character, well known in
A. punctatus, is clearly present in A. vaupesianus, less obvious in
nigropunctatus, and absent in the other taxa. It again assists in
identifying species.
Scales across the snout between second canthals: The range in the
punctatus group moderate, most species ranging between 6 and 9.
is
transversalis including, however, values as low as 4, and caquetae,
nigropunctatus and vaupesianus reaching 10. Very low counts here

may point to transversalis. Counts otherwise are not diagnostic.
Scales between the semicircles: the minimum number between the
semicircles is 0, usually implying broad contact. In the species set as
a whole the modal conditions are and 1; a count of 2 is relatively

infrequent. This character cannot be relied on to distinguish species.
Loreals: The maximum number of loreals (counted down from
the second canthals) may be as low as 3 (transversalis) or as high as
7 (vaupesianus). More frequently the count ranges from 4 to 6.
Again, a very low number of loreals may point to transversalis, but

in general loreals are not diagnostic.
Interparietal: A very large interparietal in contact with the
semicircles is a distinguishing feature of the unique type of caquetae
and of the type series of santamartae and of dissimilis and deltae.
An interparietal smaller than the small ear is characteristic of
nigropunctatus (clearly so only in the type). For the other members

of the punctatus group the interparietal is not diagnostic.
Number of supra labia Is to the center of the eye: The exceptional
number of 11 is distinctive for A. dissimilis. A range of 6 to 9 is
characteristic for all the other species and hence does not discrimi-
nate among them.
Lamellae under phalanges ii and Hi offourth toe: Number of toe
lamellae in Anolis correlates with two factors — perch and size. In
the known punctatus group anoles tend to high perches—

cases,
usually in the crown and should have high lamellar counts for
their size. Correspondingly, the better-known, larger species, known
—
to correspond to expectations in perch punctatus, vaupesianus
—
have counts in the mid- to upper 20s. A. nigropunctatus, A. jacare,
and A. huilae have relatively low counts for punctatus group
animals of their size (20-24).
28 BREVIORA No. 467
Tail crests: Two species

— dissimilis and deltae — have tail crests
and hence are unique in the group. Others have a double row
without a crest.
Species Diagnoses
What combinations of characters diagnose the punctatus group

species I have described herein and the others previously described?
(See also Tables 3 and 4.)
1) Vaupesianus is distinctive in its black dewlap, strongly
protuberant snout in males, weakly keeled ventrals, and
moderate size. (Distribution: Vaupes and Amazonas.)
2) Huilae is spotted or streaked dewlap, and
distinctive in its
(usually) in the vivid mottling of its flanks and in its moderate

size (ventrals smooth). (Distribution: the upper Magdelena
valley in Huila and Tolima, perhaps at high elevations on the

eastern slope of the Andes in Meta [Manzanares].)
3) Santamartae is distinctive in the large interparietal broadly in
contact with the semicircles, plus smooth or weakly keeled

ventrals, small size,and a light line from supralabials to ear and
from ear to upper arm. (Distribution: south slope of Santa
Marta mountains.)
4) Caquetae is diagnosed by its large interparietal broadly in
contact with the semicircles, plus keeled ventrals, small size,
and no light line from suboculars to ear. (Distribution: known
only from the Upper Apaporis in Caqueta, Colombia.)
5) Nigropunctatus stands out by its small interparietal, smaller
than the small ear, an irregularly and weakly punctate pattern
on the flanks (in males on middorsum also), snout protuberant
in males, and by moderate size (ventrals smooth). (Distribu-
tion: from Norte de Santander in Colombia to Tachira and
Trujillo in Venezuela.)
6) Deltae has the large interparietal broadly in contact with the
semicircles and small size of caquetae but has smooth ventrals,
a tail crest and a short, blunt head. (Distribution: Delta of the
Orinoco in Venezuela.)
7) Dissimilis has a very large interparietal in contact with the
semicircles, small size, smooth ventrals and tail crest like those
of deltae, but a strikingly elongate head. (Distribution: Madre
de Dios in Peru.)
1982 THREE NEW COLOMBIAN /lA'OZ./S' 29
8) Jacare is distinctive in moderate size, male without a

protu-
berant snout, with an immaculate yellow dewlap and a strongly
reticulate body pattern with often an oblique light streak from
the throat to above the shoulder; /ema/e with unspotted body
and unmarked throat (ventrals smooth). (Distribution: the
Sierra Madre de Merida in Venezuela.)
9) Transversalis is \\k.& jacare in moderate size and the lack of a

protuberant snout in males, but the male has a yellow dewlap
with streaksand spots, and a body pattern of oblique light lines
enclosing rows of black spots; and females with bold transverse
bands on the body and the throat vividly cross-marked with
black {ventrals smooth). (Distribution: western Amazonia.)
10) Punctatus has moderate size and a strongly protuberant snout
in males and a yellow or orange dewlap. {Ventrals smooth or
keeled). (Distribution: Amazonia, the Guianas, and the Atlan-
tic Forest of Brazil.)
The Geography of Difference
The species o{ iht punctatus group that have been discussed here
are all those thus far reported that are east of the Andes or inter-
Andean. (For convenience, santamartae is counted in this group; it
lies inan eastward-looking valley in the southern part of the isolated
Sierra Nevada de Santa Marta.) This assemblage, as it turns out, is,
if not a natural group, clearly as natural as a group which also
includes taxa west of the Andes, currently referred to the punctatus

species group. None of the western species are demonstrably close to
the species here considered.'
'I no longer consider A. nigropunclatus to be especially close to A. nigrolineatus
(Williams, 1965). A. nigropunclatus. as will be suggested below, is probably closer to

A. jacare. and A. nigrolineatus may be a strict synonym of A. festae Peracca
(syntypes examined). Both nominal species have the same highly peculiar dewlap
with an elongate dark spot. There are no scale differences. There is variability in head
shape. The sole known differences between southern Ecuadorian specimens
(Machala, El Oro) known from recent large collections by Fitch, Echelle and Echelle.
and northern populations (Pichincha) known from equally large collections made by
Kenneth Miyata are smaller size in the northern populations (Miyata, personal
communication) and a blue iris in the north (Miyata) as compared with the yellow
iris reported by Fitch et al. (1976) for southern animals. No recently collected
— —
animals live or preserved show the narrow dark middorsal line believed diag-
nostic for A. nigrolineatus.
30 BREVIORA No. 467
I I
I I
crj
1982 THREE NEW COLOMBIAN ANOUS 31
^
a
2 + 2
to
^
a
so
o
^-^
3
II +
— 7
i^
Z
.6c
(S3
H
32 BREVIORA No. 467
>3
3
>
u
C/3
—
1982 THREE NEW COLOMBIAN ^/VO/,/5 33
The eastern and inter-Andean assemblage is, itself, clearly

morphologically heterogeneous, showing several levels of differen-
tiation. Yet it is a striking fact that only one of the forms here 1 1
called species is widely sympatric with any of the others: A.

transversalis occurs within the range of A. punctatus and A.
vaupesianus. It is abundantly distinct not only from the two taxa

with which it sympatric, but from
is allother punctatus group
species. It differs not only in the striking color characters
emphasized in the section immediately above, but it stands at one
extreme in scale characters also (low counts of loreals and scales
across the snout).
None of the other nine described forms is known to co-occur.
Two of them, however, are in their own way sharply distinct. A.
dissimilis, as its name implies, seemed very distinct when it was first
made known, but the tail crest, otherwise unique within the
punctatus group, has since been found in widely allopatric deltae
(which differs, however, inter alia, in its much shorter head).
The remaining taxa are allopatric; they do, however, divide into
low-land and high-land forms, and this, while it is not prima facie
evidence against conspecificity, makes it a little less likely.
The low-land species are punctatus, vaupesianus, and caquetae.
Here it is clear that as regards the first two I have relied on color in
life. The however,
difference, sharp, and, is if there are any
we can specify where they must
intermediate populations, be: in
Amazonas between the Rios Putumayo and Caqueta.

It is, of course, crucial to the final allocation of all allopatric taxa
that are phenetically similar that, when possible, we endeavor to

discover whether there is a contact zone and to determine what
happens have expressly chosen not to call distinctive
there. I
populations subspecies merely because they are related. Unless

variation is clinal and the relevant characters not congruent in —
which case taxonomic description is not warranted the alternative —
to the hypothesis of allospecies, which I am using here, is subspecies,
a highly restrictive
hypothesis according to which distinctive
populations with congruent characters meet in well-defined zones of
intergradation. I agree with the long-expressed opinion of Wilson
and Brown ( 953) that these restrictive conditions are rarely met. A.
1
punctatus and A. vaupesianus are, indeed, distinctive populations

but closest relatives that are either allospecies, the ends of a cline, or
supspecies. There is not present evidence of clinal change or of
34 BREVIORA No. 467
intergradation. My decision therefore is to tentatively regard the

two populations as allospecies.
The case of A. caquetae is different. It is a single specimen
peculiar in some aspects of color and squamation
—
in combination
outside the known limits of other forms. As a single specimen, it is
not clearly representative of any distinctive population. None of its
characters is individually unusual within the broad variation of the
punctatus group. The large interparietal in contact with supra-

orbital semicircles is not at all usual in iht punctatus species group,
but see A. santamartae. The color as preserved is distinctive but
suffers the difficulty of preserved color in Anolis, discussed at some
length above. Clearly, acceptance oi A. caquetae as a valid species
involves risk of error. But if it were to be synonymized, what would
itbe synonymized with? A. punctatus or A. vaupesianusl Or would
both sink into a variable species to be called A. punctatusl The large
interparietal in contact with semicircles is rare in A. punctatus (one
known case in 157 specimens), unknown in A. vaupesianus (but the
sample is too small for a secure statement). It is closer to A.
vaupesianus, but it would seem erroneous on present evidence to
synonymize A. vaupesianus with caquetae. The case here is one of
inadequate evidence, and, above all, of inadequate search for
evidence. The fauna of Rios Vaupes, Caqueta, and Putumayo and
their tributaries is very poorly known.
The remaining taxa with which we are concerned—yafare,
—
nigropunctatus, huilae, and santamartae are all animals of high
elevations. Of these, only jacare is an old name, and huilae and
santamartae have been described herein.
Santamartae is the most distinctive of these. In the tendency to
keeling of the ventrals and the light line from the labials toward the
ear it is unlike the others. It seems also to be smaller.
The other three, it must be confessed, are primarily distinctive in
coloration. A. huilae seems to stand out in this series. The bold light
spotting and nape spot in males, as well as the contact of
differentiated anterior and inferior nasals with the rostral, contrast
with the dark spotting of nigropunctatus and the vermiculation of
jacare and with the primitive circumnasal scale characteristic of
both the latter.
Betweenyocare and nigropunctatus there are no conspicuous scale
differences other than the smaller interparietal of nigropunctatus
(but this is sharply evident only for the male type oi nigropunctatus,
1982 THREE NEW COLOMBIAN /I A'OL/5' 35
the only known male). They differ, however, in color; the sparse
dark spots oinigropunctatus are quite unlike the bold vermiculation
of male jacare. I am reinforced in my confidence of this color
character by examination of a series of specimens o{ jacare from
1 1
three localities in the state of Merida made available to me by James

R. Dixon. The males seem clearly distinctive; the females are not as
readily assigned. Between these two, however, and even between
nigropunctatus and huilae, there might be a contact zone. The
appropriate intermediate regions remain uncollected.
I have not here gone beyond a minimal analysis of the phenetic
similarities of the 10 species of the easiern punctatus species group. I
do not attempt any history or even any dendrogram of relationship.

It seems to me totally premature to do so.
Of the 10 species only three —punctatus, transversalis, and

—
jacare have been known for more than 15 years. Of these, only
punctatus is widespread and relatively common. A. transversalis is
widespread but rather scarce in collections, although the collections
of Pekka Soini (Soini and Dixon) indicate that it must be
moderately common, at least in the Iquitos region. A. jacare is
known only from the Merida region in Venezuela where, from the
experience of Carlos Rivero Blanco and Williams (Williams et al,
1970, and the recent collections by James Dixon), it is commoner
than the infrequent sightings of it and the meager collections
indicate.
Anoles of the punctatus group are animals usually of the canopy
and appear rather infrequently on the trunks or near the ground {A.
huilae fide Dennis Harris may be an exception since it is, according
to his report, more of a trunk-ground adapted species like A.
frenatus [cf. Scott et al., 1976 for the latter]). Most of them are,
therefore, not conspicuous Uzards, whatever their real abundance.
Other species may be genuinely rare, rare enough that the
question of intraspecies and interspecies, even intergeneric compe-
tition, becomes moot. It is difficult, however, to demonstrate
that
failure to observe is equivalent to real rarity. Many species of Anolis

also are highly cryptic and behaviorally highly adapted to crypsis,
almost never visible unless they move. Some, readily capturable
while sleeping and therefore known to be at least locally common,
are rarely seen in the daytime, and their diurnal activity and ecology
are therefore almost unknown.
36 BREVIORA No. 467
However, quite apart from any characteristics of the animals that

may make acquisition of knowledge about them difficult, a primary
and evident difficulty is failure to explore and to collect. Anolis
huilae not only a species just now described, but only very recently
is
collected. Dennis Harris is the collector of all the Tolima specimens

— 23 obtained in one 10-day trip which did not specialize in Anolis
but sought lizards of all kinds. Such a collection in a brief span
implies that prior sampling of this area, which has not been totally
neglected
—
it is the
type locality oi Anolis tolimensis and A. ihague
-
has nevertheless been quite inadequate.

If this is true of areas from which there is moderately good
representation in old collections, what may still be coming from

areas that are still altogether or nearly untouched?
The Department of Nariiio is one such area. Collections by even
Ayala's group have been only in the low-lands near Tumaco; there is
no reason to believe that even near Tumaco there is not much more
to find.
An undescribed species is known from the west side of the Sierra
Nevada de Santa Marta, and one on the south side has been de-
scribed herein. What occurs on the east side? Are the new species the
only ones on the west and south sides? Is the fauna of the Choco
completely known, of Cauca, Antioquia, Bolivar, Santander, and
Norte de Santander? One can, in fact, call the roster of the
Departments of Colombia and it will not be possible to say even of
the relatively well collected areas that the herpetofauna is com-
pletely known. We
in fact do not know enough to estimate the frac-
tion that unknown. (However, to shirk the frustrating effort of
is
dealing with incomplete and inadequate evidence would be to

acquiesce in the perpetuation of ignorance. Our obligation is, while
not avoiding these thorny thickets, to explore carefully and to leave
the paths that we pursue plainly marked.)
What has been done in this paper is I do not
to attempt — and
vouch for more than an —
answer the first and most
attempt to
elementary (but most fundamental) of biological questions: who?
-
i.e.. what is the cast of characters? After we have answered this first
question we may begin to ask the question where? On the way we

may begin to answer other questions: what does it do? with whom
does it interact? Much the picture is much clearer, we
later, when
can try to answer historical questions: how did this system come
about. On the evidence before us the picture is too incomplete, the
1982 THREE NEW COLOMBIAN /I A'O/./S 37
unfilled places in the evidence too blatant and glaring, to begin an
analysis of species group history or even, in any serious sense, of

inter-group relationship. We
do not yet know enough to ask the
right questions.
ACKNOWLEDGMENTS
I am indebted to William F. Pyburn for permission to describe
the new species collected by him, and to Alan E. Leviton (California
Academy of Sciences [CAS]), Pedro Ruiz (Instituto de Ciencias
Naturales, Bogota [ICN]), William E. Duellman (University of
Kansas Museum of Natural History [KU]), James Dixon (Texas A
& M University), and Jean Lescure (Museum National de Histoire
Naturelle, Paris [MNHP]) for permission to examine material in
their charge. Crucial to the study was newly collected material made
available by Stephen Ayala, Dennis Harris, and William Lamar.
Stephen Ayala also prepared the distribution map. Laszlo Meszoly
in preparing the figures has done me an indispensible service.
LITERATURE CITED
Dixon, J. R., and P. Soini. 1975. The reptiles of the Upper Amazon Basin.
Iquitos, Region, Peru. I. Lizards and amphisbaenians. Contrib. Biol. Geol.
Milwaukee Public Mus. No. 4, pp. 1-58.
FiTCFi, H. S., A. F. EcHELLE. AND A, A. ECHELLE. 1976. Field observations on
rare or little known mainland anoles. Univ. Kansas Mus. Nat. Hist. Sci. Bull..
51: 91-128.
Greene, H. W. 1977. Lizards of the genus Uracentron (Iguanidae) in east-central

Colombia. Herpetologica, 33: 256-260.
HooGMOED. M, S. 1973. Notes on the herpetofauna of Surinam. IV. The lizards
and amphisbaenians of Surinam. The Hague, W. Junk. 410 pp.
Rand, A. S., and E. E. Williams. 1970. An estimation of redundancy and
information content of anole dewlaps. Amer. Nat.. 104: 99-103.
Scott, N. J. Jr., D. E. Wilson, C. Jones, and R. M. Andrews. 1976. The
choice of perch dimensions by lizards of the genus Anolis (Reptilia, Lacertilia,
Iguanidae). J. Herp.. 10: 75-84.
Williams, E. E. 1965. South American Anolis (Sauria. Iguanidae): Two new
species oi i\\Q punctaiiis group. Breviora Mus. Comp. Zool. No. 233, pp. 15. 1
1974. South American Anolis: Three new species related to Anolis

nigrolineatus and A. Jissimilis. Breviora Mus. Comp. Zool. No. 422. pp. 1 15.
Williams. E. E., and A. S. Rand. 1977. Species recognition, dewlap function.

and faunal size. Amer. Zool., 17: 261-270.
38 BREVIORA No. 467
Williams, E. E., O. A. Reig, P. Kiblisky and C. Rivero-Blanco. 1970. Anoli.s

of Venezuela. Breviora Mus.
jacare Boulenger, a solitary anole from the Andes
Comp. Zool. No. 353, pp. 1-15.
Williams, E. E.. and P. E. Vanzolini. 1980. Notes and biogeographic com-
ments on anoles from Brasil. Pap. Avuls. Zool. S. Paulo. 34: 99-108.
Wilson, E. O.. and W. L. Brown, Jr. 1953. The subspecies concept and its
taxonomic application. Syst. Zool., 2: 97-111.
MAR 1 8 1985
lARVARD
'
I'M'VERdlTY
B R E V I R A
useiim of Coiiiparaitive Zoology
IS ISSN ()()()6 9(i98
Cambridge, Mass. 30June1982 Number 468
A NEW FOREST SKINK FROM PONAPE
A. Ross Kiester'
Abstract. Emoia ponapea. new species, is described from Ponape in the Caroline
Islands where it
distinguished from all other Emoia bv the
inhabits deep forest. It is
presence of 13 premaxillary teeth and a palate intermediate between the alpha and
beta conditions.
INTRODUCTION
A survey of the scincid genus Emoia, undertaken after a collecting
trip to Ponape in the Caroline Islands and other localities in
Micronesia, showed that three specimens taken on Ponape repre-
sent a distinct species of Emoia. In fact, as discussed below, this
form possesses some characters which make its generic allocation
somewhat problematic. However, an analysis of the boundaries of
the genus Emoia is beyond the scope of this paper, and so the new
taxon is here described as:
Emoia ponapea, new species

(Figs. 15)
Holotype: MCZ 121041, forest 1/4 mile above Agricultural

Station, Kolonia, Ponape Island, Eastern Caroline Islands. Col-
lected by A. R. Kiester, 28 July 68.
Paratypes: Eastern Caroline Islands, Ponape Island: 121042- MCZ
43, same data as for holotype (The skull of 121042 has been
removed and prepared. This individual was a sexually mature
'Smithsonian Tropical Research Institute, APO Miami 34002.

BREVIORA No. 468
Head scalation of the holotype (MCZ 121041) ot Emuia ponapea.

Figure 1.
Dorsal view.
1982 A NEW FOREST SKINK
.5
5
I
Q.
C
O
3
SO
4 BREVIORA No. 468
Figure 3. Skull of paratype (MCZ 121042) of Emoia ponapea. Ventral view.

1982 A NEW FOREST SKINK
Figure 4. Skull of paratype (MCZ 121042) of Emoia ponapea. Dorsal view.

BREVIORA No. 468
Figure 5. Skull of paratype (MCZ I2I042) of Emoia ponapea. Lateral view.

1982 A NEW FOREST SKINK 7
USNM 138985-86, Nanpil River; USNM 138991, Dolo-

female.);
nier; USNM 139002-05, N end of ridge parallel to Tavensorola
River; USNM 139006-07, Dolen Eirike; CAS 152222 (formerly
USNM 138987), Nanpil River. All USNM and CAS specimens
collected between 13 September 55 and 5 December 55 by J. T.
Marshall, Jr. See acknowledgments for abbreviations.
Diagnosis: A member of the genus Enioia, as it iscurrently and
broadly construed (Greer, 1974), differing from all other members
of the genus by the possession of 13 premaxillary teeth. It is
distinguishable externally by the combination of the following
characters: interparietal present; subdigital lamellae of the 4th toe
38-46; midbody scale rows 30-32; middorsal scale rows 48-54
counting from the nuchals to the anterior insertion of the hind limb
and 52-60 counting to the point directly above the vent; body form
small (snout-vent length less than 50 mm) and distinctly slender;
head narrow and relatively pointed; coloration without metallic
blues or greens.
DESCRIPTION OF HOLOTYPE
The range of paratypical variation is given in parentheses after the
description of the holotype character.
General Appearance: Body small with a snout-vent length of 46
mm (21 mm to 49 mm for the paratypes), form distinctly slender and
gracile. Tail long, times the snout-vent length. Limbs well
over 1.5
developed, overlapping easily when adpressed to the body; hindlimb

length 23 mm, forelimb length 13 mm. Five digits on each foot.
Head and Head Scalation (Figs. 1-2): Head narrow and some-
what depressed. Rostral twice .as long as deep; supranasals
separated,much wider anteriorly than posteriorly; prefrontals
moderately to narrowly separated by frontonasal and frontal;
supraoculars 4; frontoparietals fused; interparietal single, distinct,
and one pair of parietals in contact behind interparietals; two
large;
pairs of temporals corresponding to the primary and secondary
temporals of Taylor (1935) with only the secondary temporals
placed toward the top of the head; supraciliaries 9-9; loreals two;
single pair of nuchals; 6th supralabial subocular; nasal divided
below nostril; eyelid window undivided; postmental contacts first
two infralabials on each side.

8 BREVIORA No. 468
Ear: Tympanum sunk below level of epidermis. A single larger

anterior lobule with two smaller ones below (a single smaller one on
one side of one of the paratypes). Vertical (longest) diameter of ear
opening 1.1 mm.
Body Scalation: Scales smooth. Midbody scale rows 30 (30-32).
Middorsal scale rows 48 (48-54) counting from the first scale
posterior to the nuchals to the anterior insertion of the hind leg and
53 (52-60) counting posterior to a point directly above the vent.
Subdigital lamellae on the 4th toe of the hind foot rounded with 42
on both sides (38-46). Preanals not enlarged.
Coloration: Dorsum with a central tan stripe with irregular edges
extending from the snout to the end of the tail. A small number of
black spots on the stripe on the head (some of the paratypes show
black spotting on the stripe on the back as well). A poorly defined
laterodorsal stripe on either side of this beginning behind the eye
and extending one third of the way down the tail. These stripes are
formed by two irregular rows of black dashes on the tan color. Sides
reddish brown becoming spotted over cream color toward the
venter. Limbs reticulated brownish red over cream above. Chin,
throat, venter, and undersides of limbs and tail all immaculate
cream.
DESCRIPTION OF SKULL
Skull MCZ 121042, paratype (Figs. 3-5): General appearance
very delicate and somewhat narrow (as compared, say, to a similarly
sized E. cyanura). Premaxillary teeth 13. Secondary palate moder-
meet extensively along the midline
ately well developed. Palatines
and send pointed projections posteriorly between the anterior
portions of the pterygoids. One of the projections is folded slightly
under the other. Pterygoids intermediate in condition between the
alpha and beta configurations of Greer and Parker (1968) and Greer
(1974), with the anterior portion showing distinct expansion toward
the midline without achieving a strongly recurved process.
Lower Jaw: The Meckelian canal is as the second of the two
conditions described by Greer (1974) for the Leiolopisma group
of lygosomine genera: canal closed with no suture evident.
1982 A NEW FOREST SKINK 9
DISTRIBUTION AND ECOLOGICAL OBSERVATIONS

At present this species is known only from the island of Ponape in
the Caroline Islands. Ponape is also known as Ascension Island in
some of the older literature.
All three MCZ specimens were collected in the late morning on
the floor of a mature forest in an area where sun flecks occurred. No
other emoias were seen in this deep forest habitat. E. cyanura, E.
caeruleocauda, and E. hoettgeri were found only in more open areas
towards the coast. Ecological notes by J. T. Marshall, Jr. ac-
companying the other specimens indicate that they were also col-
lected on the floor of the forest.This species bears a similar
ecological relationship to its congeners as E. parkeri does to the
other emoias in the Fiji Islands (Brown et al., 1980).
Two of the USNM
specimens are hatchlings. Marshall notes that
the eggs were collected on 27 October 55 in a rotted palm stump in
the rain forest and that they hatched on 6 November 55. A clutch
size of two is typical for most species in this genus (Greer, 1968).
DISCUSSION
The current state of confusion in the genus Emoia prevents any
accurate assessment of the relationships of Emoia ponapea. A
thorough revision of the genus would be necessary to understand
the relationships of any but the most closely related of its species.
Lacking this, a comparison with the smaller emoias found in

Micronesia may be useful. E. cyanura and E. caeruleocauda, both
also found on Ponape, have true beta palates. This character is
considered derived in the genus (Greer, 1974), as is the very high
subdigital lamellae count of cyanura itself. Thus, since E. ponapea

has a palate intermediate between the alpha and beta conditions, it
is unlikely that it is derived from any of the members oiiht cyanura
group directly. Within Micronesia a possible candidate for a related

species could be E. mivarli. However, there are notable differences
from this species as well. E mivarti has the interparietal fused with
the frontoparietal and is a much stouter animal. None of these
species seems closely related to E. ponapea; a search for its relatives
will have to extend outside of Micronesia.
10 BREVIORA No. 468
The most unusual character of E. ponapea is the possession of 13

premaxillary teeth. All other members of the genus Emoia have 1 1,
as do all of the genera regarded as related to Emoia (the members of
Group II of Greer, 1974). The only other leiolopismid genus with 13
premaxillaries is Carlia, which is clearly unrelated on other grounds

(Greer, For the genus Emoia this
1974). character state must be
derived. This character and the intermediate condition of the palate
argue that E. ponapea is rather different from the rest of the genus.
Just how different it really is will have to be determined by future
work.
ACKNOWLEDGMENTS
I wish to thank W. C. Brown, Allen Greer, the late T Preston
Webster, and Ernest E. Williams for much help and discussion on
matters related to this paper. The illustrations are by Lazlo Meszoly.
This work was supported by the Evolutionary Biology Committee
of Harvard University, The Society of Sigma Xi, and Ernest E.
Williams. For permission to examine specimens in their care, I
thank A. G. C. Grandison, British Museum (Natural History); H.
Marx, Field Museum of Natural History; A. Leviton, California
Academy of Sciences (CAS); Ernest E. Williams, Museum of
Comparative Zoology, Harvard University (MCZ); R. Stebbins,
Museum of Vertebrate Zoology, University of California at Berk-
eley; and the late J. Peters and W. R. Heyer, United States National
Museum (USNM).
LITERATURE CITED
Brown, W. C, J. C. Pernetta, and D. Watling. 1980. A new lizard of the
genus Emoia (Scincidae) from the Fiji Islands. Proc. Biol. Soc. Wash., 93:
350-356.
Greer, A. E. 1968. Clutch size in the genus Emoia. Copeia, 1968: 417-418.
1974. The generic relationships of the scincid lizard genus Leiolopisma
and its relatives. Austr. Jour. Zool. Suppl. Ser. No. 31, pp. 1-67.
Greer, A. E., and F. Parker. 1968. Geomyersia glabra, a new genus and species
of scincid lizard from Bougainville, Solomon Islands, with comments on the
relationships of some lygosomine genera. Breviora Mus. Comp, Zool. No. 302,
pp. 1-17.
Taylor, E. H. 1935. A taxonomic study of the cosmopolitan scincoid lizards of
the genus Eumeces, with an account of the distribution and relationships of
its species. Kans. Univ. Sci. Bull., 23: 1-643.
Webster, T. P. 1969. Aspects of the morphological and ecological variation in
the cyanura group of the lizard genus Emoia (Sauria: Scincidae) in the Solomon
Islands. Honors Thesis, Harvard University.
MUS. COMP. ZOOL
LfBRARY
B R E
Museum
IXO
of Comparative Zoology
R A
us ISSN 0006-9698
Cambridge, Mass. 30 June 1982 Nimbi k 469
CATALOG OF THE PRIMARY TYPES OF

BOMBYLIIDAE (DIPTERA)
INTHE ENTOMOLOGICAL COLLECTIONS OF
THE MUSEUM OF COMPARATIVE ZOOLOGY,
WITH DESIGNATIONS OF LECTOTYPES
Neal L. Evenhuis'
Abstract. One hundred and nine holotypes. lectotypes. and s\nt\pes of Bomb\-
liidae deposited in the entomological collections of the Museum of Comparati\e
Zoology are listed. Lectotypes are designated for 27 taxa based on material in MCZ.
INTRODUCTION
This catalog lists the primary types of 109 ta.xa of Bombyliidae
located in the entomological collections of the Museum of Compara-
tive Zoology. Harvard University. Cambridge (MCZ). Fifty-sexen
of these ta.xa are represented by holotypes, 7 by syntype series, and
1
35 by lectotypes. All but one of the types listed here are deposited in
the general collection of the MCZ. The unique male holot\pe of
Epihaies harrisi Osten Sacken is deposited in the Thaddeus W.
Harris collection, which is maintained separately from the general
collection. Twenty-seven new lectotype designations are recorded
herein including two by Jack C. Hall, University of California. Riv-
erside. Of the remaining 25 new lectotypes, three are designated in
Hall and Evenhuis (1981). The format used here follows that of
Radovsky ei al. ( 1976) with modifications. Information presented in
brackets [ ]
is data additional to or corrected from data recorded in
the original description. The present combination of each taxon
'Department of Entomolog\. Bishop Museum. P.O. Box 19000-A. Honolulu.

Hawaii 96819.
2 BREVIORA No. 469
it differs from the

listed, if original combination, is also included
after the remarks section under each taxon concerned.
The majority of species recorded here were described by Loew in
his "Centuries," Osten Sacken in his "Western Diptera," and C.W.
Johnson various publications. Most of the types or type series in
in
the collections carry a red label with a type register number placed
there by Nathan Banks when he curated the collection. Recent addi-
tions to the type collection carry a similar label with a type number
placed there by various curatorial technicians since Banks. Banks'

type labeling has posed problems because he sometimes failed to
label all of the syntypes in a syntype series or labeled more than the
original description of that particular taxon was based upon (this is

especially true for many of Loew's species). In compiling this
catalog, all original descriptions were checked to make sure the
correct number of syntypes were located and labeled. Discrepancies
in thenumber of specimens found versus the number of specimens
upon which that species was based are noted in the remarks section
under each taxon concerned. In some cases, syntypes of certain taxa
were found to be located in other museums. These museums include
the United States National Museum of Natural History, Washing-
ton, D.C. (USNM), the American Museum of Natural History, New
York (AMNH), the Snow Collection at the University of Kansas,
Lawrence (SNOW) and the Zoologisches Museum of the Museum
fiir Naturkunde der Humboldt-Universitat, Berlin (ZMHU). This
information is supplied in the remarks section under each taxon

concerned.
Some syntypes of the bombyliid taxa described by Osten Sacken
in "Biologia Centrali-Americana" are deposited in the MCZ. Lecto-
types for most of these species were designated by Painter and Paint-
er (1962) from material in the British Museum (Natural History),
where in most cases the majority of the syntypes of these particular
taxa are housed. Specimens of these species represented in the MCZ
carry labels signifying that the lectotype for that species is in the
BMNH.
SPECIES-GROUP INVENTORY
adusta Loew, Anthrax SYNTYPES 2 $ 12672
1869 Berliner Entomol. Z. 13: 26 pin
CUBA: [no further data] Gundlach
1981 BOMBYLIIDAE TYPES IN MCZ 3
The specimens in the MCZ are left here as syntypes pending revision ot the genus.
Present name: Villa adusia (Loew)
agassizi Loew, Exoprosopa HOLOTYPE $ 12635

UNITED STATES: California: [no further data] Agassiz
albicapillus Loew, Bombylius LECTOTYPE 9 12686

1872 Berliner Entomol. Z. 16: 78-79 pin
UNITED STATES: [California]: San Francisco [no date]
H. Edwards
Lectotype designated by Evenhuis in Hall and Evenhuis (1980: 6).
albidipennis Loew, Geron HOLOTYPE (S 12713

aldrichi Johnson, Phthiria SYNTYPES 2 (5, 1 $ 7555

1903 Psyche 10: 184 pin
UNITED STATES: Idaho: Caldwell 24TV.1901 C.W. John-
son Collection
These specimens are left as syntypes pending revision of the genus.
Present name: Phthiria (Poecilognathus) aldrichi Johnson
alpha Osten Sacken, Anthrax LECTOTYPE? 12660

1877 Bull. U.S. Geol. Geogr. Surv. pin
Terr. 3: 239-40
UNITED STATES: Wyoming: Cheyenne 21.VIIL O. Sacken
Four (three male,one female) of the fivt specimens on which Osten Sacken based
his description of alpha have been located at the MCZ. The fifth syntype is deposited
in AMNH. The best preserved of the MCZ material, the female, is here designated
lectotype.
Present name: Poecilanthrax alpha (Osten Sacken)
amabilis Osten Sacken, Ploas LECTOTYPE (5 12703

Terr. 3: 261-62
UNITED STATES: California: Yosemite 5.VI. O. Sacken
Osten Sacken based his description of amabilis on two female and one male. The
male is here designated lectotype.
Present name: Conophorus amabilis (Osten Sacken)
4 BREVIORA No. 469
arenicola Johnson, Anthrax lateralis LECTOTYPE S 7548

190S Psyche 15: 15 pin
UNITED STATES: New Jersey: Clementon I6.V. 1917
C.W. Johnson Collection
Lectotype male and allolectotype female designated herein by J.C. Hall.
Present name: Villa lateralis arenicola (Johnson)
arizonicus Banks, Systropus SYNTYPES 2 (5 13484

1909 Entomol News 20: 18 pin
UNITED STATES: Arizona: Palmerlee [no date] N. Banks
Collection
These specimens are left as syntypes pending revision of the genus.
atratula Loew, Picas HOLOTYPE $ 12698

UNITED STATES: California: [no further data] H. Edwards
Present name: Conophorus atratulus (Loew)
atriceps Loew, Bombylius LECTOTYPE $ 12688

UNITED STATES: Florida: [no further data] O. Sacken
aurifer Osten Sacken, Bombylius LECTOTYPE $ 12691

Terr. 3: 249-50
UNITED STATES: California: Webber Lake 26. VII.
O. Sacken
banksi Johnson, Dipalta HOLOTYPE S 13485

1921 Occas. Pap. Boston Soc. Nat. pin
Hist. 5: 12-13
UNITED STATES: Virginia: Great Falls 12.IX. Collection
N. Banks
barbatus Osten Sacken, Anastoechus LECTOTYPE (5 12719

Terr. 3: 252-53
UNITED STATES: Wyoming: Cheyenne 21. VIII. O. Sacken
Lectotype designated by Hall in Hall and Evenhuis (1981: 153-54).
bifurca Loew, Exoprosopa HOLOTYPE 9 12636

Loew's description of bifurca was apparently based on a single specimen because
there is no range of measurements recorded. There are two female specimens in the
MCZ labeled as types. The one in the best condition is considered to be the holotype. It
fitsLoew's description and measurements and carries a label "bifurca m." in Loew's
handwriting. The other specimen is denuded, smaller than the holotype,
and carries a
label "bifurca" in Banks' handwriting. The latter specimen probably does not belong to
the original type series.
bigradata Loew, Anthrax HOLOTYPE [$] 12655

Original description incorrectly cites sex of type as "(5".
borealis Johnson, Phthiria HOLOTYPE $ 27054

\9\Q Psyche \1: 229 pin
UNITED STATES: Maine: Fort Kent 7.VIII. 1910
Present name: Phthiria (Poecilognathus) borealis Johnson
brevicornis Loew, Sparnopolius SYNTYPES 2 $ 12723

UNITED STATES: Texas: [no further data] Beifrage
Though Loew gives no range of measurements, two specimens are labeled as types
in the MCZ. They are left here as syntypes until their type status can be clarified.
cachinnans Osten Sacken, Bombylius LECTOTYPE S 12693

Terr. 3: 250
UNITED STATES: California: Sonoma County 2.IV.-9.V.
R. Osten Sacken
Male and female on same Lectotype designated by Evenhuis //; Hall and
pin.
Evenhuis(1980: 29).
calvus Loew, Geron SYNTYPES 2 $ 12711

UNITED STATES: New York [no further data]
Loew's description does not include a range of measurements though two speci-
mens are labeled as types in the MCZ. Until this discrepancy can be clarified, they are
left as syntypes.
6 BREVIORA No. 469
candidulus Loew, Systoechus LECTOTYPE (5 12716

UNITED STATES: Wisconsin: [no further data] O. Sacken
Loew's description cites a range of measurements though only one male was
located in the MCZ. The male is here designated leclolype.
canuta Hall, Thevenemyia HOLOTYPE ^ 31339

1969 Univ. Calif. Publ. Entomol. pin
56: 28-29 Fig. 8
UNITED STATES: Arizona: Reef lO.X. C.W. Johnson
Collection
capito Osten Sacken, Pantarbes LECTOTYPE (5 12720

Terr. 3: 256
UNITED STATES: CaUfornia: Sonoma County 2.IV.-9.V.
R. Osten Sacken
Lectotype male designated herein by J.C. Hall.
cervinus Loew, Aphoebantus HOLOTYPE $ 12684

UNITED STATES: Texas: [no further data] Belfrage
ceyx Loew, Anthrax LECTOTYPE^ 12664

UNITED STATES: Virginia [no further data]
Loew's description cites a range of measurements though only one male was found
in the MCZ. It is here designated lectotype.
Present name: Poecilanthrax demogorgon (Walker)
contigua Loew, Argyromoeba HOLOTYPE $ 12679

UNITED STATES: Virginia: [no further data] Loew
Present name: Anthrax argyropygus argyropygus
Wiedemann
coquilletti Johnson, Phthiria SYNTYPES 2 (5, 4 9 7554

1902 Can. Entomol. 34: 240-41 pin
UNITED STATES: New Jersey: 1
(5, 4 $, Riverton 4.VII.1891;
1 (5, Jamesburg 4.VII.1891 C.W. Johnson Collection
Three syntypes are in USNM; one syntype is in SNOW; these specimens are left as
syntypes pending generic revision.

Present name: Phthiria (Poecilognathus) coquilletti Johnson
cubana Loew, Exoprosopa SYNTYPES (5, 1 1

$ 12641
Two syntypes are also located in ZMHU. These specimens are left as syntypes
pending generic revision.
curta Loew, Anthrax HOLOTYPE (5 12654

Present name: Hemipenthes curta (Loew)
cyanoceps Johnson, Phthiria HOLOTYPE S 27053

\903> Psyche W. \Â pin
UNITED STATES: Massachusetts: Cohasset 8TX.
Present name: Phthiria (Poecilognathus) cyanoceps Johnson
decora Loew, Exoprosopa HOLOTYPE 9 12630

IS69 Berliner Entomol. Z. 13: 13 pin
UNITED STATES: Wisconsin: [no further data] Kennicot
delila Loew, Argyromoeba HOLOTYPE? 12674

Present name: Anthrax delila (Loew)
diagonalis Loew, Anthrax HOLOTYPE? 12656

Present name: Paravilla diagonalis (Loew)
dodrans Osten Sacken, Exoprosopa SYNTYPES 2 (5 12634

Terr. 3: 234-35
UNITED STATES: [Colorado]: Colorado Springs [no date]
O. Sacken
These specimens are left here as syntypes pending generic revision.
8 BREVIORA No. 469
dorcadion Osten Sacken, Exoprosopa LECTOTYPE $ 12631

1877 Bull. U.S. Geo I. Geogr. Surv. pin
Terr. 3:231-33
UNITED STATES: Maine [no further data]

Osten Sacken's original description of dorcadion is based on syntypes from
"Summit Station, Central Pacific Railroad, California (July 17); Webber Lake,
Sierra Nevada, California (July 26); Shasta district, California (H. Edwards);
Washington Territory (the same); Georgetown, Colorado (August 12); Twin Lake
Creek, Colorado (W.L. Carpenter); White Mountains, New Hampshire (H.K. Mor-
rison); Maine
— 2 specimens from Denver, Colorado (Uhler)." The MCZ has all but
the New Hampshire and Shasta district specimens. The Summit Station specimen
also carries the label "Sierra Nevada." All the specimens are females except for the
one male from Denver, Colorado. The female specimen from Maine is in the best
condition and is here designated leciotype. Three syntypes are also deposited in
AMNH.
doris Osten Sacken, Exoprosopa SYNTYPE 9 12632

ISn Bull. U.S. Geo Geogr. Surv.
I.
pin
Terr. 3: 235-36
UNITED STATES: Nevada: Humboldt Station 29. VII.
O. Sacken
One syntype (headless) is deposited in AMNH. These specimens are left as syn-
types pending generic revision.
edwardsii Loew, Allocotus HOLOTYPE [S] 12704

Original description incorrectly states sex as "9".
Present name: Paracosmus edwardsii (Loew)
egerminans Loew, Phthiria LECTOTYPE [9] 12708

Original description incorrectly cites sex of type as "(5". Loew's description cites a
range of measurements though only one female was found in the MCZ. It is here
designated leciolype.
eremita Osten Sacken, Exoprosopa HOLOTYPE^ 12637

Terr. 3: 236
UNITED STATES: California: Shasta District [15.VII.]
H. Edwards
euplanes Loew, Argyromoeba HOLOTYPE $ 12681

Present name: Anthrax euplanes (Loew)
fenestrata Osten Sacken, Ploas LECTOTYPE $ 12702

\S11 Bull. U.S. Geol. Geogr. Surv. pin
Terr. 3: 260-61
UNITED STATES: California: San Raphael 12.IV. O. Sacken
Osten Sacken based his description oi fenestrata on two males and one female. Of
these syntypes, the best preserved (the female) is here designated leciotype.
Present name: Conophorus fenestratus (Osten Sacken)
flaviceps Loew, Anthrax LECTOTYPE? 12663

MEXICO: Tamaulipas [no further data]
Of the two female syntypes of this species in the MCZ, the best preserved is here
designated lectotype.
Present name: Poecilanthrax flaviceps (Loew)
fraudulentus Johnson, Bombylius HOLOTYPE S 7552

1907 P^vf/ze 14: 99-100 pin
UNITED STATES: Massachusetts: Provincetown 27. VI. 1904
fuliginosa Loew, Anthrax HOLOTYPE (5 12661

UNITED STATES: California [no further data]
Specimen rubbed.
Present name: Poecilanthrax fuliginosus (Loew)
fur Osten Sacken, Argyromoeba LECTOTYPE c5 12678

Terr. 3: 244-45
UNITED STATES: Texas: Dallas [no date] Boll
Osten Sacken based his description of fur on one male and two female syntypes.
The best preserved of these (the male) is here designated lectotype.
Present name: Anthrax limatulus fur (Osten Sacken)
gazophylax Loew, Exoprosopa HOLOTYPE? 12629

10 BREVIORA No. 469
1869 Berliner Entomol Z. 13: 12-13 pin

Head missing.
Present name: Ligyra gazophylax (Loew)
gibbus Loew, Lordotus HOLOTYPE $ 12724

MEXICO: [Tamaulipas]: Matamoros [no further data]
The label data cites the locality as "Matamoras."
haemorrhoicus Loew, Bombylius HOLOTYPE S 12696

CUBA: [no further data] [Gundlach]
Original description cites collector as "Riehl," label data cites "Gundlach."
Present name: Heterostylum haemorrhoicum (Loew)
harrisi Osten Sacken, Epibates HOLOTYPE S 26400

Terr. 3: 273
[No locality label] H. Gray
Hall (1969: 38) proposed a lectotype male (MCZ Type No. 31340) and allolecto-
type female for harrisi under the assumption that the original type specimen was
lost.' The type male is, in fact, present in the T.W. Harris Collection [in MCZ] as
described by Osten Sacken.
Present name: Thevettemyia harrisi (Osten Sacken)
humilis Osten Sacken, Phthiria HOLOTYPE 5 12710

Terr. 3: 264
UNITED STATES: CaHfornia: Sonoma County 4. VII.
O. Sacken
incanus Johnson, Bombylius HOLOTYPE S 7553

1907 Psyche 14: 97-98 pin
UNITED STATES: Massachusetts: Provincetown 2. VI. 1904
Present name: Bombylius (Zephyrectes) incanus Johnson
'Though neotypes should be proposed for lost or destroyed types. Hall mistakingly
proposed a lectotype and allolectotype which, therefore, have no validity no-
menclatorially.
johnsoni Painter, Geron HOLOTYPE S 27266

1932 Trans. Am. Entomol. Soc. pin
58: 155
UNITED STATES: Massachusetts: Horseneck Beach 30.
VII. 1913 C.W. Johnson Collection
lancifer Osten Sacken, Bombylius LECTOTYPE ^ 12692

Terr. 3:251
UNITED STATES: California: [no further data] Sacken
Lectotype male designated by Evenhuis in Hall and Evenhuis (1980: 57).
lepidotoides Johnson, Chrysanthrax HOLOTYPE S 7550

\9\9 Psyche 16: \2 pin
UNITED STATES: New Jersey: lona 16. VI. 1902
limbata Loew, Picas LECTOTYPE? 12700

UNITED STATES: New Mexico [no further data]
Specimen badly rubbed; right wing glued to point below specimen; head, left wing,
leftfore- and hind- and right mid- and hindlegs beyond coxae missing. Loew's
description cites a range of measurements though only one female was found in the
MCZ. It is here designated lectotype.
Present name: Conophorus limbatus (Loew)
luctifer Osten Sacken, Epibates HOLOTYPE S 12728

\%11 Bull. U.S. Geol. Geogr. Surv. pin
Terr. 3: 271-72
[CANADA: British Columbia]: Vancouver Island [no date]
G.R. Crotch
Present name: Thevenemyia luctifera (Osten Sacken)
macer Loew, Systropus LECTOTYPE 9 12726

UNITED STATES: Pennsylvania [no further data]
Of the three syntypes oi macer (1 male, 2 females) in the MCZ, the best preserved
of these (a female) is here designated lectotype.
macropterus Loew, Geron HOLOTYPE S 12712

12 BREVIORA No. 469
1869 Berliner Entomol. Z. pin

13: 172-73
UNITED STATES: New York: Geneseo [no further data]
Head and thorax damaged; abdomen beyond third segment missing.
Present name: Geron (Empidigeron) calvus Loew
magnus Osten Sacken, Epibates HOLOTYPE $ 12731

Terr. 3: 272-73
[CANADA: British Columbia]: Vancouver Island [no date]
G.R. Crotch
Present name: Thevenemyia magna (Osten Sacken)
marginatus Osten Sacken, Epibates HOLOTYPE $ 12730

Terr. 3: 272
UNITED STATES: California: San Francisco [no date]
H. Edwards
Present name: Thevenemyia marginata (Osten Sacken)
metopium Osten Sacken, Bombylius HOLOTYPE (5 12690

Terr. 3: 249
UNITED STATES: California: [San Francisco] 19.IV
O. Sacken
modestus Loew, Leptochilus SYNTYPES 1

(5, 1
9 12685
UNITED STATES: Texas: [no further data] Belfrage
The specimens are left here as syntypes pending revision of the genus.
Present name: Epacmus modestus (Loew)
molitor Loew, Anthrax HOLOTYPE 9 12652

Head glued to thorax.
Present name: Villa molitor (Loew)
mucorea Loew, Anthrax HOLOTYPE? 12651


UNITED STATES: Nebraska: [no further data] [F. V. Hayden]
Collector recorded in Osten Sacken ( 1903: 95).
Present name: Villa mucorea (Loew)
muricatus Osten Sacken, Epibates HOLOTYPE (5 12729

Terr. 3: 272
UNITED STATES: California: Sierra Nevada [no date]
H. Edwards
Present name: Thevenemyia muricata. (Osten Sacken)
itius Osten Sacken, Triodites LECTOTYPE ^ 12683

Terr. 3: 246-47
UNITED STATES: Utah: Sah Lake City l.VIII. O. Sacken
Of the three syntypes of mus (1 male, 2 females) located in the MCZ, the best
preserved of these (the male) is here selected as lectotype.
Present name: Aphoebantus mus mus (Osten Sacken)
nigricauda Loew, Anthrax HOLOTYPE $ 12650

UNITED STATES: Massachusetts: [no further data] Scudder
Head missing.
Present name: Villa fulviana nigricauda (Loew)
nigripennis Loew, Ploas LECTOTYPE? 12697

Loew's description cites a range of measurements though only one female was
found in the MCZ. It is here designated lectotype.
Present name: Conophorus nigripennis (Loew)
notata Loew, Phthiria HOLOTYPE? 12706

Present name: Phthiria (Poecilognathus) loewi Painter
nubifera Loew, Exoprosopa LECTOTYPE S 12639

14 BREVIORA No. 469
CUBA: [no further data] [Poey]

Original description cites collector as "Gundlach," label data cites "Poey." Loew's
description cites a range of measurements though only one male was found in the
MCZ. The male is here designated lectotype.
obesula Loew, Ploas SYNTYPES 3 (5 12699

Loew's original description oi obesula was based on an unstated number of males
and females. Only three males were found in the MCZ. They are left here as syntypes
until the genus is revised.
Present name: Conophorus obesulus (Loew)
obsoleta Loew, Argyromoeba HOLOTYPE [9] 12675

UNITED STATES: Missouri [no further data]
Original description incorrectly states sex of type as "(5".
Present name: Anthrax limatulus limatulus Say
occidentalis Johnson, Spogostylum SYNTYPES 2 9 7547

1913 Bull. Am. Mus. Nat. Hist. pin
32:56
UNITED STATES: 1 $, Colorado: Denver 5.VIII.1897; 1 9,
Washington: Seattle [no further data] C.W. Johnson

Collection
These specimens are left here as syntypes pending generic revision.
Present name: Anthrax analis Say
oreas Osten Sacken, Systoechus LECTOTYPE (5 12718

Terr. 3: 254
UNITED STATES: California: Webber Lake 22. VII.
O. Sacken
Lectotype designated by Hall in Hall and Evenhuis (1981: 147).
palliata Loew, Anthrax HOLOTYPE? 12657

UNITED STATES: Illinois: [no further data] Osten Sacken
Present name: Paravilla palliata (Loew)
parva Loew, Exoprosopa LECTOTYPE (5 12640

Though Loew's description cites a range of measurements, only one male was
located in the MCZ. It is here
Present name: Neodiplocampta (Neodiplocampta) parva (Loew)
parvicornis Loew, Anthrax HOLOTYPE (5 12658

UNITED STATES: Illinois: [Chicago] [no date] Osten Sacken
Present name: Rhynchanthrax parvicornis (Loew)
pauper Loew, Argyromoeba HOLOTYPE 9 12677

UNITED STATES: IlUnois [no further data]
Present name: Anthrax pauper (Loew)
pertusa Loew, Anthrax HOLOTYPE^ 12659

UNITED STATES: New Mexico: [lOTV.] [no further data]
Present name: Thyridanthrax pertusa (Loew)
planus Osten Sacken, Lordotus HOLOTYPE (5 12725

1877 Bull. U.S. Geol. Geogr. pin
Surv. Terr. 3: 258-59
UNITED STATES: California: Marin County [no date]
O. Sacken
proboscidea Loew, Anthrax HOLOTYPE (5 12653

MEXICO: Sonora: [no further data] Schott
Present name: Lepidanthrax proboscideus (Loew)
pulchellus Loew, Bombylius LECTOTYPE (5 12687

UNITED STATES: Illinois [no further data]
Lectotype designated by Evenhuis in Hall and Evenhuis ( 1980:79).
quinquenotata Johnson, Phthiria HOLOTYPE $ 7556

16 BREVIORA No. 469
1903 Psyche 10: IS5 pin

UNITED STATES: Colorado: Grand Junction 25.V.1900
Present name: Oligodranes quinquenotata (Johnson)
ravus Loew, Bombylius HOLOTYPE $ 12694

MEXICO [no further data]
Present name: Bombylius (Zephyrectes) ravus Loew
rufula Osten Sacken, Picas LECTOTYPE (5 12701

Terr. 3: 261
UNITED STATES: California: San Geronimo 19TV.
O. Sacken
Present name: Conophorus rufulus (Osten Sacken)
sagata Loew, Anthrax LECTOTYPE (5 12671

MEXICO: [Tamaulipas]: Matamoros [no further data]
Head missing; label data cites locality as "Matamoras". Loew's description cites a
range of measurements, though only one male was found in the MCZ. It is desig-
nated here as lectotype.
Present name: Hemipenthes sagata (Loew)
scolopax Osten Sacken, Phthiria SYNTYPES [2^, 1$] 12789

Terr. 3: 263-64
UNITED STATES: Colorado: Manitou 17-18.VIII. O. Sacken
The original description of scolopax was based on one male and three females.
Two male and one female syntypes are located in the MCZ. These specimens are left
here as syntypes pending generic revision.
Present name: Phthiria (Poecilognathus) scolopax Osten Sacken
scrobiculata Loew, Anthrax LECTOTYPE S 12648

Of the two male syntypes of scrobiculata in the MCZ, the best of these is here
Present name: Villa scrobiculata (Loew)
seminigra Loew, Hemipenthes HOLOTYPE [(5] 12673

CANADA: Saskatchewan: [no further data] Kennicot
Head missing; original description incorrectly cites sex of type as "9"
Present name: Hemipenthes seminigra Loew
semirufus Loew, Bombylius LECTOTYPE (5 12695

HAITI: [Geremie]: [no date] P.R. Uhler
The location of Geremie is cited from the label data. It was omitted from the original
description. Of the two syntypes oi semirufus (one male, one female) in the MCZ, the
best preserved (the male) is here designated lectotype.
Present name: Heterostylum semirufum (Loew)
serpentina Osten Sacken, Dipalta LECTOTYPE? 12646

Terr. 3: 237
UNITED STATES: California: Mt. Shasta District [no date]
O. Sacken
Osten Sacken based his description of serpentina on one male and two female
syntypes. The best preserved of these (a female) is here designated lectotype.
shawii Johnson, Anthrax HOLOTYPEcJ 7549

\90S Psyche 15: 14-15 pin
UNITED STATES: New Hampshire: Hampton 27.VIII. 1906
S.A. Snow
The holotype also carries a type label reading "holotype no. 599".'
Present name: Villa shawii (Johnson)
sima Osten Sacken, Exoprosopa LECTOTYPE [(5] 12628

Terr. 3:231-33
UNITED STATES: Nevada: Humboldt Station: Central Paci-
fic Railroad [no further data]
Osten Sacken's original description of sima incorrectly cites the sex of the three
syntypes as female. They are, in fact, all males. The best preserved is here designated
lectotype.
This number probably pertains to Johnson's personal collection prior to MCZ

accession (Woodley, personal communication).
18 BREVIORA No. 469
slossonae Johnson, Spogostylum HOLOTYPE S 7546

1913 Bull. Am. Mus. Nat. Hist. pin
32:55
UNITED STATES: Kentucky: Cumberland Gap VII. 1876 G.
Dimmock
Present name: Anthrax aterrimus (Bigot)
sordida Loew, Exoprosopa SYNTYPES [2$] 12642

MEXICO: 1$, Tamaulipas: [no further data]; 1$, [Tamauli-
pas]: Matamoros [no further data]
Loew's original description of sordida cites an unspecified number of males and
females. Only two males were found in the MCZ; label data cites locality of the latter
syntype as "Matamoras." These specimens are left here as syntypes pending generic
revision.
stellans Loew, Argyromoeba HOLOTYPE [$] 12676

UNITED STATES: Oregon [no further data]
Right wing broken; original description incorrectly cites sex of type as "5".
Present name: Anthrax stellans (Loew)
stenozona Loew, Anthrax LECTOTYPE $ 12649

A lectotype is here designated for the one female in MCZ
as Loew's description
cites a range of measurements, implying a series of specimens.
Present name: Villa stenozona (Loew)
subauratus Loew, Geron SYNTYPES 2 (5, 1 $ 12714

UNITED STATES: Pennsylvania: [no further data] Osten
Sacken
The specimens are left here as syntypes until the genus is revised.
subvarius Johnson, Bombylius LECTOTYPE? 7551

1907 Psyche 14: 98-99 pin
UNITED STATES: Pennsylvania: Lehigh Gap 26. VI. 1901
H.L. Viereck
Lectotype female designated herein.
Present name: Bombylius atriceps Loew
sulphurea Loew, Phthiria HOLOTYPE $ 12705

UNITED STATES: New Jersey: [no further data] Glover
Present name: Phthiria (Poecilognathus) sulphurea Loew
titubans Osten Sacken, Exoprosopa HOLOTYPE [c5] 12633

Terr. 3: 233-34
UNITED STATES: [Colorado]: Denver [no date] O. Sacken
Original description incorrectly states sex of type as "9".
trabalis Loew, Exoprosopa HOLOTYPE? 12638

MEXICO: [Veracruz]: Jalapa [no further data]
Present name: Exoprosopa anthracoidea Jaennicke
validus Loew, Bombylius LECTOTYPE S 12689

1863 Berliner Entomol. Z. 7: 300 -
pin
UNITED STATES: Virginia: [no further data] Osten Sacken
virgata Osten Sacken, Toxophora SYNTYPES 1 (5, 1

$ 12727
Terr. 3: 266-67
UNITED STATES: Georgia [no further data]
Osten Sacken based his description of virgata on two males and two females from
Georgia and Texas. A male and a female from Georgia are in the MCZ. The Texas
specimens have not been located. These specimens are left here as syntypes pending
generic revision.
vitripennis Loew, Geron SYNTYPES 2$ 12713

UNITED STATES: "Middle States" [no further data]
No locahty labels on type specimens. Until the genus is revised they are left as
syntypes.
vulgaris Loew, Systoechus LECTOTYPE^ 12717

UNITED STATES: Nebraska: [no further data] [F. V. Hayden]
Lectotype designated by Hall in Hall and Evenhuis (1981: 151); original descrip-
tion incorrectly states collector as "Osten Sacken." Osten Sacken (1903: 95) gives the
correct collector.
20 BREVIORA No. 469
webberi Johnson, Villa HOLOTYPE S 12680

1919 Psyche 26: 11 pin
UNITED STATES: Massachusetts: Lunenberg 3. VI. 1914 [no
collector]
Present name: Hemipenthes webberi (Johnson)
willistoni Osten Sacken, Pantarbes HOLOTYPE (5 12722

1887 Biologia Centrali- Americana pin
1: 153
UNITED STATES: Arizona: [no further data] O. Sacken
xanthomeros Marston, Anthrax HOLOTYPE $ 32555

1970 Smithson. Contrib. Zool. pin
43: 52-53 PI. 3g
BRITISH HONDURAS: Benque Viejo [no date] Father
Stanton
GENUS-GROUP INVENTORY
(indexed by present combination)
Anastoechus: barbatus
Anthrax: bigradata, contigua, delila, euplanes, fur, obsoleta, occi-
dentalis, pauper, slossonae, stellans, xanthomeros
Aphoebantus: cervinus, mus mus
Bombylius (Bombylius): albicapillus, atriceps, aurifer, cachinnans,
fraudulentus, lancifer, metopium, pulchellus, subvarius, validus
Bombylius (Zephyrectes): incanus, ravus
Chrysanthrax: lepidotoides
Conophorus: amabilis, atratulus, fenestratus, limbatus, nigripennis,
obesulus, rufulus
Dipalta: banksi, serpentina
Epacmus: modestus
Exoprosopa: agassizi, bifurca, cubana, decora, dodrans, dorcadion,
doris, eremita, nubifera, sima, sordida, titubens, trabalis
Geron: albidipennis, calvus, johnsoni, macropterus, subauratus,
vitripennis
Hemipenthes: curta, sagata, seminigra, webberi
Heterostylum: haemorrhoicum, semirufum
Lepidanthrax: proboscideus
Ligyra: gazophylax
Lordotus: gibbus, planus
Neodiplocampta (Neodiplocampta): parva
Oligodranes: quinquenotata
Pantarbes: capito, willistoni
Paravilla: diagonalis, palliata
Paracosmus: edwardsii
Phthiria (Phthiria): egerminans, humilis
Phthiria (Poecilognathus): aldrichi, borealis, coquilletti, cyanoceps,
loewi, scolopax, sulphurea
Poecilanthrax: alpha, ceyx, flaviceps, fuUginosus
Rhynchanthrax: parvicornis
Sparnopolius: brevicornis
Systoechus: candidulus, oreas, vulgaris
Systropus: arizonicus, macer
Thevenemyia: canuta, harrisi, luctifera, magna, marginata
Thyridanthrax: pertusa
Toxophora: virgata
Villa: adjusta, lateralis arenicola, molitor, mucorea, fulviana nigri-
cauda, scrobiculata, shawii, stenozona
ACKNOWLEDGMENTS
I wish to thank Mr. Jack C. Hall for reviewing the manuscript of
this paper and Drs. Pedro Wygodzinsky of the American Museum
of Natural History and H. Schumann of the Museum fiir Natur-
kunde der Humboldt-Universitat, Berlin for their help in locating
Osten Sacken and Loew types, respectively, in their museums. I also
my sincere gratitude to Mr. Norman E. Woodley for
wish to express
his assistance during my visit to the MCZ and for his comments and
suggestions concerning the manuscript of this study. I thank Ms.
Margaret K. Thayer for her help in obtaining for me the initial loans
of some of the type material in the MCZ.
Travel for this study was supported by National Science Founda-
tion Grant DEB 79-10192 to the University of California, Riverside.
LITERATURE CITED
Banks, N. 1909. A new species of Syslropus (Bombyliidae). Entomol. News, 20: 18.
22 BREVIORA No. 469
Cole, F.R. 1917. Notes on Osten Sacken's group "Poecilanthrax," with descriptions
of new species. J. N.Y. Entomol. Soc, 25: 67-80.
Hall, J.C. 1969.A review of the subfamily Cylleniinae with a world revision of the
genus Thevenemyia Bigot (Eclimus auct.) (Diptera: Bombyliidae). Univ. Calif.
Publ. Entomol., 56: 1-85.
Hall, J.C, and N.L. Evenhuis. 1980. Part 13, Number 1. Family Bombyliidae, pp.
1-96. In Griffiths, G.C.D. (ed.), Flies of the Nearctic Region. Vol. V. Stuttgart, E.
Schweizerbart'sche.
1981. Part 13, Number 2. Family Bombyliidae, pp. 97-184. In Griffiths,
G.C.D. (ed.). Flies of the Nearctic Region. Vol. V. Stuttgart, E. Schwei-
zerbart'sche.
Johnson, C.W. 1902. New North American Diptera. Can. Entomol., 34: 240-42.
. 1903. Descriptions of three new Diptera of the genus Phthiria. Psyche, 10:
184-85.
. 1907. A review of the species of the genus Bombylius of the eastern United
States. Psyche, 14:95-100.
. 1908. Notes on New England Bombyliidae, with a description of a new species
o{ Anthrax. Psyche, 15: 14-15.
. 1910. Some additions of the dipteran fauna of New England. Psyche, 17:
228-35.
1913. Insects of Florida. I.
Diptera. Bull. Amer. Mus. Nat. Hist., 32: 37-90.
. 1919. New species of the genus Villa (Anthrax). Psyche, 26: 1 1-13.
. 1921. New species of Diptera. Occas. Pap. Boston Soc. Nat. Hist., 38: 57-99.
LoEW, H. 1863a. Diptera Americae septentrionalis indigena. Centuria tertia. Berliner

Entomol. Z., 7: 1-55.
. 1863b. Diptera Americae septentrionalis indjgena. Centuria quarta. Berliner
Entomol. Z., 7: 275-326.
. 1869a. Diptera Americae septentrionalis indigena. Centura octava. Berliner
Entomol. Z., 13: 1-52.
. 1869b. Diptera Americae septentrionalis indigena. Centuria nona. Berliner
Entomol. Z., 13: 129-186.
. 1872. Diptera Americae septentrionalis indigena. Centuria decima. Berliner
Entomol. Z., 16:49-115.
Marston, N. 1970. Revision of New World species of /I /7//7/-a.v (Diptera: Bombyliidae)
other than the Anthrax albofasciatus group. Smithson. Contrib. Zool., 43:
1-148.
Osten Sacken, C.R. 1877a. Art. XIII. Western Diptera: Descriptions of new genera
and species of Diptera from the region west of the Mississippi and especially
from California. Bull. U.S. Geol. Geogr. Surv. Terr., 3: 189-354.
1887b. Diptera. pp. 129-216. In Godman, F.D. and O. Salvin (eds.), Biologia
Centrali-Americana. Zoologia-Insecta-Diptera. Vol. I. London, Taylor & Fran-
cis, 378 pp.
. 1903.Record of my life work in entomology. Cambridge, Mass., University

Press, John Wilson & Sons, 204 pp.
Painter, R.H. 1932. A monographic study of the genus Geron Meigen as it occurs in
the United States (Diptera: BombyUidae). Trans. Amer. Entomol. Soc, 58:
139-67.
Painter, R.H., and E.M. Painter. 1962. Notes on and redescriptions of types of
North American Bombyliidae (Diptera) in European Museums. J. Kans.
Entomol. Soc, 35: 1-164.
Radovsky, F.J., G.A. Samuelson, and W.A. Steffan. 1976. Catalog of entomological
types in the Bishop Museum. Introduction. Pac. Insects, 17: 1-5.

LfDRARY
MAR 1 8 1985
B R E V-'-F^ R A
IS ISSN ()()()6 9(i!)8
Camhridge, Mass. 30JunkI982 NrviBHR470
SYSTEMATIC IMPLICATIONS OF
INNERVATION PATTERNS
IN TELEOST MYOTOMES
QuENTiN Bone' and R. Dana Onc)^
Abstract. The peripheral innervation patterns of the red and white myotomal
muscles from over 230 species representing more than 125 families of teleosts uere
studied. A distributed, multiple innervation pattern ot telcost red superficial
myotomal muscles was found without exception in all groups examined. 1 here is
innervation patterns of the white m\otomal muscles, however. A

variabilitN' in the
terminally innervated pattern seems to be present in the basal groups of teleosts.

while a trend toward distributed mnervation occurs in the Neoteleostei. Stomii-
formes possess a rather different distributed pattern which \ve suggest is the early,
transitional stage from terminal to distributed innervation patterns in teleosts. There
appears to be a distinct functional difference in the distributed and terminal

innervation patterns. The innervation of the uhite mvotomal libers should be
considered a taxonomicallv useful character in elucidating familial relaiionships.
INTRODUCTION
In all fishes, two main muscle fiber types in the
there are
myotomes, usually naked eye when the fish is
readily visible to the
sectioned transversely. Small-diameter, well-vascularized red mus-
cle fibers normally form a thin superficial layer covering the much
more numerous, larger-diameter. poorl\-vascularized white muscle

fibers that make up the major portion of the myotome. In some
fishes, but not in all, other myotomal fiber types are present. These
minor myotomal components will not concern us here. In all fishes.
'The Laboratory. Citadel Hill. Plymouth PI 1 2PB. Devon. England.

-Museum of Comparative Zoology. Harvard I'niversitv. Cambridge. Massachusctt--
02138.
BREVIORA No. 470
i2 S E
1982 MYOTOME INNERVATION
OSTARIOPHYSI
O
s
o
o
o
BREVIORA No. 470
Figure 3. Distributed innervation pattern of motor endplates in white myotomal

muscle fibers of Morune lahrax as revealed by acetylcholinesterase (AChE) studies.
(Mag. X90).
'^. "V
Figure 4. Terminal innervation pattern of motor endplates in white myotomal

muscle fibers of the clupeomorph. Sprattus sprattus. as revealed by AChE studies.
(Mag. y 90).
1982 MYOrOME INNERVATION
Figure 5. Silver impregnated axons of terminal innervation pattern m the white
myotomal muscle fibers of the alepocephalid, Xenock'rnucluhvs copci. Wmkelmann

and Schmitt technique. (Mag. "190).
Ifc
Figure 6.
^ X
Silver impregnated axons of the distributed inner\ation pattern m the

white myotomal muscle fibers of the gymnotid, Eiênniannia vircsccns. VVinkclmann
and Schmitt technique. (Mag. X228).
6 BREVIORA No. 470
Chondrostei (Acipenseridae, Polydontidae), hagfish, Elasmobran-

chiomorpha, and Latimeria chalunmae have shown that terminal
innervation universal in non-teleost groups (Bone, 1964 and
is
unpublished; Ono, unpublished; Sakharov and Kashapova, 1979).

In addition, terminal innervation is found in the Dipnoi, in the
myotomal musculature of adult urodeles, and in urodele and anuran

larvae (Best and Bone, 1973; Bone, unpublished; Ono, unpub-
lished).
distribution of the terminal innervation pattern of white
The wide
muscle fibers led to a preliminary attempt (Bone, 1970) to link the
occurrence of the terminal pattern in teleosts with the systematic
position of the families in which it was found. This attempt

was
unsuccessful since relatively few families were examined. In this
study, we report the results of a more detailed survey of the
innervation pattern of white muscle fibers in different teleost groups
that suggests that this character may be of interest to systematists.

The innervation pattern of white muscle fibers was studied either
by supravital staining with

methylene blue in
Ringerteleost
solution, or by silver staining of 10% unbuffered formalin and

Bouin-fixed material using the methods of Palmgren (1948) and
Winkelmann and Schmitt (1957). During the last several years, we
have collected marine fish material from the Caribbean, the Indian
Ocean, the North Atlantic Ocean, and the Bay of Biscay. The
freshwater fish material was collected by us from North American
and European waters. In addition, we have been much assisted by
other workers who have provided specially fixed or museum
material from the waters of South America, Africa, Japan, New
Zealand, and several deep-sea localities. We owe a particular debt of
gratitude to Dr. P. H. Greenwood (BMNH) and to Mr. J. B.
Badcock and Mr. N. Merrit (lOS) who have generously provided
material from their collections.
TELEOST GROUPS AND THEIR

INNVERVATION PATTERNS
The status of the different groups of teleosts is still subject to
review, and different authors have adopted different systems of
classification to indicate their views of the relationships of the
1982 MYOTOME INNERVATION 7
various groups. In this paper we have adopted the classification

scheme of Rosen (1973) and Fink and Weitzman{ 1982). A complete
checklist showing the teleosts examined is given in Table 1.
1) Osteoglossomorpha
Nine genera from this group were examined. With the exception
of the two Hiodon species, all possess the distributed pattern of
white muscle innervation.
2) Elopomorpha
We
have examined 12 genera from this group representing the
Elopiformes, Anguilliformes, and Notacanthiformes. All possess
the terminal innervation pattern. Both the adults and leptocephali
of Elops sp. and Albula sp. have the terminal pattern of white
muscle innervation.
3) Clupeomorpha
Twenty-one species of clupeids belonging to 16 genera were
studied. All possess the terminal innervation pattern with the sole
exception of Denticeps, in which the white myotomal muscle has a

distributed innervation.
4) Ostariophysi
Wehave examined 32 genera of ostariophysans representing the
Gonorynchiformes, Characiformes, Cypriniformes, and Siluri-
formes. Some ostariophysans possess the terminal pattern (some
siluroids and all gonorynchids), but the majority have the distrib-
uted pattern of innervation.
5) Protacanthopterygii
There is considerable uncertainty about the limits of this group,
which has been reduced from its original size (Fink and Weitzman,
1982). The 16 species representing 14 genera listed in Table
are 1
provisionally placed in this group.
6) Stomiiformes
Fifteen genera representing the two Infraorders Gonostomata
and Photichthya (Weitzman, 1974) were studied. Two innervation
patterns are found in these two lineages of Stomiiformes.
While the
Gonostomata possess the usual distributed innervation pattern
found in other teleosts, members within the Photichthya have the
white zone of the myotome innervated in a different way. Terminal
endplates are seen on many fibers which are probably innervated
at
8 BREVIORA No. 470
both ends. In addition, some axons pass from the myosepta into the
midregions of the myotome to form a sparsely distributed innerva-
tion pattern. Thus, the Photichthya possess a third type of
innervation pattern (Fig. IC), distinct from either the terminal or
distributed patterns, apparently combining elements of both.
7) Eurypterygii
examined in the Aulopi-
All 125 species representing 99 genera
formes, Myctophiformes, Paracanthopterygii, and Acanthopterygii
possess the distributed innervation pattern (see Table 1 for list of
eurypterygians examined).
DISCUSSION
All non-teleost fishes possess terminally innervated white muscle
fibers (Ono, unpublished). It is reasonable to suppose therefore that
this pattern is the primitive condition and that the distributed
pattern of innervation is derived. The universal occurrence of the
distributed pattern in the Eurypterygii supports this view.
Two questions arise. First, is there a functional advantage in the
distributed pattern that has led to its universal appearance in
"higher" teleosts? Secondly, can white muscle innervation prove

useful in dealing with the controversial questions of the limits and
interrelationships of different teleost groups?
Consideration of the habits of fishes possessing the two types of
innervation pattern, for example, Hiodon and Heterotis or A /hula
and Chanos, shows no obvious correlation between habitat and
innervation pattern, nor any obvious differences in locomotor
ability. But studies of fishes swimming under controlled conditions
have revealed a notable difference in locomotor ability between
fishes where the white muscle is innervated terminally, and where it
is innervated in the distributed manner. Few fishes have yet been
studied in this
way (where activity in different zones of the myotome
is monitored by electromyography as the fish swims at different
speeds), but results are consistent, and are probably generally
applicable. In all fishes studied, the red muscle fiber zone of the
myotome is active during slow cruise
swimming that can be
maintained indefinitely. In fishes where the white muscle fibers are
terminally innervated, these are only active during bursts of rapid
swimming, and are rapidly exhausted (Bone, 1966; Bone ei al..
1982 MYOTOME INNERVAIION 9
1978). Where the white muscle fibers are innervated in the

distributed manner, they operate quite differently, for they are
active not only during bursts of rapid swimming, but also during
cruise swimming at intermediate speeds (Hudson, 1973; Johnston ei
ai, 1977; Bone et ai. 1978).
Electromyographic records from the white muscle zone during

cruise swimming in such fishes are different from those obtained
during rapid swimming, and it

appears that where the fibers receive
distributed innervation, they can operate in two ways propagating
action potentials only during rapid swimming (Bone et ai, 1978).
Extracellular recordings cannot provide other than suggestive
evidence on this point, and definite proof of the hypothesis that
contraction of the same fast muscle fiber in such fishes can either
follow local potentials or propagated action potentials awaits
intracellular investigation.
However, the results of these electromyographic studies clearly

show that fishes with the distributed pattern of innervation of the
white muscles are able to recruit these fibers to give a wider range of
sustainable cruising speeds than can be obtained when the white
fibers are terminally innervated and are used only during burst
swimming. Evidently, this ability to swim for long periods over a
wide speed range could confer a significant advantage to fishes in
some particular habitats. For example, it may be advantageous to
fishes living in streams where flow varies and the fishes are required
to keep stationary. This ability to swim over
a wide speed range
would be insignificant to benthic fishes, however, at least in the
adult stage due to the relatively stable flow regime of the water
column.
We conclude that the distributed pattern of innervation was
derived in teleostean evolution when adults or larvae were pelagic,
and has been retained in those groups where the adults today are
relatively sedentary, so that the habits of the adults today are not
correlated with the innervation pattern.
The first attempt to use the innervation pattern as a systematic
character made the simple assumption that this derivation from the
ancestral terminal pattern occurred only once during teleost
phylogeny (Bone. 1970). The present more complete survey

(unfortunately still lacking data on several important species)
demonstrates that this assumption can no longer by justified. Figure
10 BREVIORA No. 470
2 shows a current view of the relationships among teleost groups
(Fink and Weitzman, 1982), and indicates points where transition

from terminal to distributed innervation is assumed to have taken
place. In this view of the relationships of the groups, the change
presumably took place independently on at least eight occasions.
In general, our survey has shown that the more primitive groups
possess the terminal innervation pattern, as expected. Thus in the

Osteoglossomorpha, the Hiodontoidei is a phylogenetic relict in the
sense that the two living Hiodon species form a lineage that has
retained a large number of primitive features (Greenwood, 1970);
only these members of the group possess the terminal pattern.
Both the Elopomorpha and Clupeomorpha (with the sole
exception of Denticeps) also possess the terminal pattern. The
freshwater Denticeps retains many primitive characters, (Green-
wood, 1968), but is unique among clupeomorphs in having achieved
the distributed innervation pattern. Since a wider sustainable speed
range might be a distinct advantage in the fluviatile freshwater
environment, we examined other African freshwater clupeids, but
all five genera studied possessed terminally innervated white muscle
fibers.
In the Ostariophysi, all Gonorynchiformes and some Siluriformes
have the terminal innervation pattern, and if the scheme of
ostariophysan relationships proposed by Fink and Fink (1981) is
accepted, this implies that convergent origin of the distributed
pattern has occurred in all lines except that leading to the
Gonorynchiformes (Fig. 2). Certainly, although the phylogenetic
position of Siluriform families is uncertain, the innervation pattern
is terminal in the most primitive, the relict Diplomystidae.
Perhaps the most interesting group with respect to muscle

innervation pattern is the Stomiiformes. This group has been less
thoroughly studied than the Ostariophysi, and despite recent work
by Fink and Weitzman (1982), and an earlier study by Weitzman
(1974), relationships within the group are still uncertain. It is
notable that the two major sister groups forming the Stomiiformes,
the Gonostomata and Photichthya (Weitzman, 1974), have different
innervation patterns. In the Gonostomata, innervation is dis-
tributed, but in the Photichthya, innervation of the white fibers is
mixed. Relatively few axons course through the white portion of the
myotome, and in addition there are axons terminating on the fiber
1982 MYOTOME INNERVATION I I
ends as in the terminal pattern. Perhaps in this group of teleosts. we

witness the distributed innervation pattern at an early stage in its
development. It would be interesting to see whether the mi.xed
pattern of adult photichthyans develops from an initially terminal
pattern in the larval stage.
Few studies involving the nervous system in fishes have been used
in a comparative fashion to solve problems of classification. On the
whole, our survey suggests that the innervation of white myotomal
fibers,despite evident convergence in its origin, should be con-
sidered a systematically useful character, and may prove particu-
larly helpful in elucidating interrelationships within the Siluriformes
and the Stomiiformes, respectively.
ACKNOWLEDGMENTS
We express our gratitude to Drs. P. H. Greenwood, William L.
Fink, Karel F. Liem, Ian A. Johnston, as well as to Mr. Julian B.
Badcock and Ms. Sara V. Fink for providing helpful comments and
specimens. R. D. O. was partially supported with funds from a
Raney Award (American Society of Ichthyologists and Herpeto-
logists). National Science Foundation Grant BNS-79I5308, and
National Institutes of Health Predoctoral Fellowship GMO-7117
during the tenure of this project.
LITERATURE CITED
Barets, a. Contribution a I'etude des systemes moteurs lent-et rapid du
1961.
muscle lateral des teleosteens. Arch. Anat. Morphol. E.xp.. 50 (suppl.): 91 187.
Best, A. C. G., and Q. Bone. 1973. The terminal neuromuscular junctions of
lower chordates. Z. Zellforsch., 143: 495-504.
Bone, Q. 1964. Patterns of muscular innervation in the lower chordates. Int. Rev.
Neurobiol., 6: 99 147.
1966. On the function of the two types of m\otomal muscle fibre in
elasmobranch fish. J. Mar. Biol. Ass. U.K.. 46: 321 349.

1970. Muscular innervation and fish classification, pp. 369-377. In A.
de Haro
(ed.), Simposio Internacional de Zoofilogenia, Fac. Ciencias,
1 Llniv.
Salamanca, 492 pp.

Bone, Q., J. Kicenuik, and D. R. Jone.s. 1978. On the role of different fibre
types in fish myotomes at intermediate swimming speeds. Fi^h. Bull..
76: 691 699.

Fink, S. V., and W. L. Fink. 1981. Interrelationships of the ostariophysan
fishes (Teleostei). Zool. J. Linn. Soc, 72(4): 297 353.
BREVIORA No. 470
Fink. W. 1... \Nn S. H. Wi it/man. 1982. Relationships of the Stomiiform fishes

(Teleostei) with a description of Diplophos. Bull. Mus. Comp. Zool., 150: 31-93.
Grhfnwood. p. H. I96S. The osteology and relationships of the Denticipitidae,
a family of clupeomorph Mus. Nat. Hist. (Zool.).. 16: 213-273.
fishes. Bull. Br.
1970. On the genus, Lvn'pieia. and its relationships with the family
Hiodontidae (Pisces, Osteoglossomorpha). Bull. Br. Mus, Nat. Hist. (Zool.).,
19: 257 285.
Hi'D.soN, R. C. L. 1973. On the function of the white muscles in teleosls at
intermediate swimming speeds. J. Exp., Biol., 58: 509 522.
.loHNSioN, 1. A., W. Davidson, and G. Goldspink. 1977. Energy metabolism
of carp swimming muscles. J. Comp. Physiol., 114: 203 216.
Palmgren. a. 1948. A rapidmethod for selective silver staining of ner\e fibres
and nerve endings in mounted paraffin sections. Acta Zool., 29: 378-392.
Rosen, D. E. 1973. Interrelationships of higher euteleostean fishes, pp. 397-513.

In P. H. Greenwood. R. S. Miles, and C. Patterson (eds.), London, Academic
Press, 536 pp.
Sakharov, D. a., and L. A. Kashapova. 1979. The primitive pattern of the
vertebrate body muscle innervation: Ultrastructural e\idence for two synaptic
transmitters. Comp. Biochem. Physiol., 62(A): 771 776.
Weitzman, S. Osteolog\' and evolutionary relationships of the

H. 1974.
Sternoptychidae with a new classification of stomiatoid families. Bull. Am. Nat.

Hist., 153(3): 329-478.
Winkelmann, R. K., and R. W. Schmitt. 1957. A simple silver method for
nerve axoplasm. Proc. Staff Meeting Mayo Clinic. 32: 217 222.
1982 MVOIOML IWI l<\ \l 1()\ 13
Table 1. Patterns of Innervation m the White M\otomal Muscle fibers ot
Teleosts.
White Myolomal
Muscle Kibcrs
List of Teleosts Examined for Terminal Distributed
Innervation Pattern Pattern Pattern
Osteoglossomorpha
Notopteridae
+
Soiopierus chiiala
+
Xencmystus nig,ri
Osteoglossidae
Heterotis niloticus +
+
Osieoglossum hicirrosum
Pantodontidae
Paniodon hiicholzi
Mormyridae
Mormyrus sp.
Mormyrops engystoma
Gnathonennis petersii
Hiodontidae
Hiodun tergisus +
Hiodon alosoides +
Elopomorpha
Elopiformes
Elopidae
El ops sp. +
Albulidae
Alhula +
sp.
Anguilliformes
Anguillidae
"•
Anguilla sp.
Muraenidae
+
Gyninolhorax sp.
Congridae
+
Conner sp. (2)
^
Paraconger sp.
Muraenesocidae
\luraencso.\ sp. +
Ophichthidae
•
Ophichlhus sp.
14 BREVIORA No. 470
Table 1. continued
White Myotomal
Muscle Fibers

Eurypharyngidae
Eurypharynx pelecanoides
Notacanthiformes
Halosauridae
Halosaurus sp. +
Halosauropsis sp. +
Notacanthidae
Polyacanthonotus sp.
Clupeomorpha
Clupeidae
Clupea sp. +
Harengula sp. (2) +
Sardine I la sp. +
Pellonula atzeliusi +
Opisthoplerus sp. +
Limnothrissa miodon +
Thrissodes sp. +
Euplatygaster sp. +
Cynothrissa memo +
Poeciloihrissa congicae +
Stolothrissa tanganicae +
Microthrissa sp. +
Opisthopterus sp. +
Alosa pseudoharengus +
Alosa aestivalis +
Alosa sp. +
Sprattus sprattus +
Engraulidae
Anchoa mit chilli +
Anchoa sp. +
Chirocentidae
Chirocentrus dorab +9
Denticipitidae
Demiceps clupeoides
EUTELEOSTEI
Ostariophysi
Gonorynchiformes
Table continued
White Myotomal
Muscle Fibers

Chanidae
Chanos chanos
Kneriidae
Kneria mitiei
Phractolaemidae
Phraeiolaemus ansurgei
Gonorynchidae
Gonorynchus gonorynchus
Characiformes
Characidae
Crenuchus spilurus
Hyphessobrycon flammeus
Hyphessobrycon serpae X
Hyphessobrycon collistus
Hyphessobrycon pulchrspinnis
Astyanax mexicanus
Lebiasinidae
Nannostomus nannostomus
Gasteropelecidae
Gasteropelecus sp.
Hemiodontidae
Hemiodus sp.
Cypriniformes
Cyprinidae
Nolropis hudsonius +
Cyprinus +
sp.
Gyrinocheilidae
Gyrinocheilus aymonieri
Catostomidae
Catostomus catosiomus
Cobitidae
Noemacheihis sp.
Siluriformes
Siluroidei
16 BREVIORA No. 470
Table 1. continued
White Myotomal
Muscle Fibers
Bagridae
Parauchenoglanis macrostoma
Siluridae
Kryptopierus hicirrhis
Malapteruridae
Malapterurus eleciricus
Pangasiidae
Pangasius sutchi
Chacidae
Chaca chaca
Mochokidae
Synodontis sp.
Aspredinidae
Bunocephalus sp.
Callichthyidae
Corydoras sp.
Hoplosternum sp.
Diane ma sp.
Loricariidae
Ancistrus sp.
Ictaluridae
Icialurus sp.
Dipiomystidae
Diplomysies sp.
Ariidae
Arius sp.
Doradidae
Doras sp.
Pimelodidae
Soruhim limas
Pimelodella sp.
Gymnotoidei
Rhamphichthyidae
Eigenmannia virescens
Table continued
White Myotomai
Muscle Fibers

Protacanthopterygii
Salmoniformes
Esocidae
Esox niger
Esox americanus
Umbridae
Umhra liml
Dallia pectoralis
Salmonidae
Salmo iriitta
Salmo sp.
Retropinnidae
Reiropinna sp.
Galaxiidae
Galaxias sp.
Osmeridae
Osmerus mordax
Plecoglossidae
Plecoglossus altivelis
Argentinidae
Argentina sp.
Opisthoproctidae
Opisthoprocius sp.
Alepocephalidae
Alepocephalus sp.
Xenodermichthys copei
Baihylaco nigricans
Searsiidae
Searsia sp.
Stomiiformes
Gonostomata
Gonostomatidae
Cvclothone ohscura
Gonosioma e/ongaium
Maurolicus sp.
BREVIORA No. 470
Table I. continued
White Myotomal
Muscle Fibers

Sternoptychidae
Argyropelecus sp. (2)
Sternoptyx sp.
Photichthya
Chauliodontidae
Chauliodus sp.
Stomiatidae
Stomias sp.
Macrostomias longibarbatus
Astronesthidae
Astronesthes sp.
Melanostomiatidae
Melanostomias sp. +
Eustomias sp. +
Echiostoma barbalum +
Malacosteidae
Malacosteus sp. +
Photostomias sp. +
Idiacanthidae
Idiacanthus sp.
EURYPTERYGII
Aulopiformes
Aulopodidae
Aulopus sp.
Synodontidae
Sy nodus sp. (2)
Giganturidae
Gigantura sp.
Bathypteroidae
Bathypierois sp. (2)
Myctophiformes
Myctophidae
Myctophum sp.
Diaphus sp.
Table I. continued
White Myotomal
Muscle Fibers

Paralepididae
Paralepis sp.
Omosudidae
Omusmiis sp.
Evermannellidae
Coccorella sp.
Scopelarchidae
Scopelarchus sp.
Paracanthopterygii
Gadiformes
Moridae
Antimora sp.
Macrouridae
Nematonurus sp.
Lophiiformes
Lophiidae
Lophius sp.
Antennariidae
Antennarius hispidus
Antennarius scaber
Acanthopterygii
Atheriniformes
Exocoetidae
Parexocoetus sp.
Cypselurus sp.
Hemiramphus sp. (2)
Belonidae
Be lone sp.
Platybelone sp.
Tylosurus sp.
Cyprinodontidae
Belonesox belizanus
Scomberesocidae
Scomberesox sp.
20 BREVIORA No. 470
Table I. continued
White Myotomal
Muscle Fibers

Atherinidae
Meniciia metiidia
Atherinomorus sp.
Beryciformes
Polymixiidae
Poly mixta lowei
Polymixia japonica
Holocentridae
Holocentrus sp. (3)
Cetomimidae
Cetomimus sp.
Melamphaeidae
Melamphaes sp.
Dactylopteriformes
Dactylopteridae
Dactylopterus sp.
Zeiformes
Zeidae
Zeus faher
Sygnathiformes
Aulostomidae
Aiilostomus sp.
Fistulariidae
Fistularia sp, (2)
Sygnathidae
Nerophis sp. +
Hippocampus sp. +
Scorpaeniformes
Triglidae
Trivia sp.
Cottidae
Cotlus cognathus
Perciformes
Centrarchidae
Pomoxis nigromaculatus
Lepomis gibbosus
1982 MYOrOME INNERVAI ION 21
Table 1. continued
White Myotomal
Muscle Fibers
Priacanthidae
Priacanthus sp.
Carangidae
+
Caranx sp. (3)
+
Oligoplites sp.
+
Serial a sp.
Lutjanidae
+
Lutjanus sp. (2)
+
Ocyurus sp.
Plector\nchidae
Gaterin sp.
Pentapodidae
Monolaxis sp.
Serranidae
+
Epinephelus sp. (3)
+
Petrumetapun sp.
+
Serranus sp.
+
Cephalopholis sp.
Pomadasyidae
+
Haemulon sp. (2)
+
Anisoi remits sp.
Percichthyidae
Morone Uihra.x
Sphyraenidae
Sphyraena sp.
Grammistidae
Rypticiis sp.
Scaridae
Sparisuma sp. (3)
Percidae
Etheostvma olmstedi
Stromateidae
StromaleicI sp.
Istiophoridae
Makaira sp.
22 BREVIORA No. 470
Table I. continued
White Myotomal
Muscle Fibers

Coryphaenidae
Coryphaena sp.
Sciaenidae
+
Cynoscion regalis
+
Equetus sp.
Cichlidae
Julidochromis sp.
Belontiidae
Trichogaster trichopterus
Channidae
Channa micropeltes
Mullidae
MuUus surmuletus +
Mulloidichthys +
sp.
Kyphosidae
Kyphosus sp.
Chaetodontidae
Chaetodon sp. (6)
Pomacanthus sp.
Pomacanthodes sp.
Pomacentridae
Abudefduf sp.
Cepolidae
Cepola rubescens
Centropomidae
Cenlropomus sp.
Acanthuridae
Adanthurus +
sp. (4)
Zanclus +
sp.
Scombridae
Scomber +
sp. (2)
Thunnus +
sp.
Euthynnus +
sp.
Gerreidae
Gerres +
sp.
+
Eager res sp.
Table I. continued
White Myotomal
Muscle Fibers

Siganidae
Siganus sp.
Labridae
Halichoeres sp.
Lepidaplois sp.
Anampses sp.
Lachnolaimus maximus
Gobiidae
Periophlhalmus koelreuteri
Ephippidae
Chaetodipterus sp.
Tetraodontiformes
Balistidae
Batistes sp. +
+
Xanthichthys sp.
+
Melichthys sp.
Cantherhines sp. +
+
Balistapus sp.
Ostraciotidae
Acanthostracion +
sp. (2)
+
Lactophrys sp.
Diodontidae
Diodon +
sp. (2)
+
Chilomycterus sp.
Tetradontidae
Sphoeroides sp.
In all the fishes listed above, the red myotomal muscle fibers had the distributed
pattern of innervation.
MU3. COMP. Zowu
LfRRARY
i'r
B R ¥«i:„0 R A
1#'"^^
E UNIVERSITY
iiseiim of CoiiiparatiYe Zoology
us ISSN 0006-9698
Cambridge, Mass. 30 June 1982 NrMBhR47l
ARTHUR LOVERIDGE-A LIFE IN RETROSPECT
Ernest E. Williams'
On February 16. 1980, at the age of almost eighty-nine, Arthur

Loveridge, former Curator of Reptiles and Amphibians at the
Museum of Comparative Zoology, died on the island of St. Helena
in the South Atlantic after a short illness.
In 1924 A. Lawrence Lowell, then President of Harvard, wrote to
the Immigration Department in Boston in these terms:
"This is to inform you that Arthur Loveridge, Esq.. formerly of
the Manchester University Museum. National Museum of Wales,
and latterly Director of the British East African Museum in
Nairobi, a gentleman standing high in his chosen field, is due on the
steamer Laconia. arriving in Boston on or about May 1st.
"On March 14th of this year Mr. Loveridge was appointed by the
Faculty of the Museum of Comparative Zoology in Harvard
University to the position of Associate in Zoology in the Har\ard
University Museum, where he will exercise his profession during the
coming years Harvard University.
as an officer of
"Since I informed that the British quota is full, am anxious
am I
that you should know in advance that Mr. Loveridge is a teacher,

scientist and author of high professional standing, and that he
comes here already appointed to a University position in Harvard.
"Any kindness you may show him in expediting his entr\ will be
very greatly appreciated by me."

It is obvious that President Lowell's plea was effective. It is
known that Glover Allen, then Curator of Mammals, met Lo\eridge
'Museum of Comparative Zoology, Harvard University, Cambridge. Massachusetts

02138.
2 BREVIORA No. 471
at the boat and drove him Cambridge, where he was to spend

to
thirty-three years (till 1957). At first he was Thomas Barbour's
assistant, reorganizing the Museum's herpetological collection and

then, when Barbour was appointed Director of the Museum,
continuing to supervise with surpassing care the expansion of one of
the world's great collections of reptiles and amphibians. From 1931
he had the volunteer assistance of Benjamin Shreve.
Thomas Barbour brought Loveridge to the New World, but
Loveridge's heart remained in the continent of Africa where he had
spent almost ten years. There he had made the reputation that
brought the Harvard appointment. There he had met his wife, and
there he had indulged to the full his passion for collecting and for
general natural history that had been his since childhood indulged
—
it despite (or by means of) service with the East African Mounted
Rifles, the Nairobi Museum, and the Game Department in
Tanganyika (now Tanzania).

Loveridge's association with Harvard was no boh from the blue;
it was a tie that had gradually strengthened. In the Museum Report
for 1919-1920, there is a mention of a first gift from Loveridge. His

name recurs in 1921 1922, and then, in 1923 1924, there is
Barbour's comment: "This year has been eventful in that during its
course the Arthur Loveridge African collection was received and
Mr. Loveridge arrived to assist in a general overhauling of the study
series."
Harvard got a bargain in Loveridge. Harvard bought Loveridge's
collection, but with it came a Curator — one the collection des-
perately needed. The other side of Loveridge's passion for collecting

was a passion for order and for tidiness: what he brought back or
had brought back to him had to be as perfectly classified, ticketed,
and put away as human power could manage.
For a while he surely had the best of his two worlds. On the one
hand, he was in charge of a major but crowded, ill-labeled, ill-
organized gathering of collections. He was able to transform it into
a model of collections as he felt they should be taking frogs,
—
snakes, and lizards out of tanks and putting them in fine glass-
stoppered bottles with labels written in hard pencil in his own neat
hand and carefully arranging them within trays, each with neatly
typed labels. His collection, when he finished, was a thing of beauty
(and fiercely kept so).
1982 LOVERIDGE'S LIFE IN RETROSPECT 3
For the other part, in the first years he was repeatedly able to go
back to Africa and, doing what he most wanted to do, simultane-
ously enrich the Harvard collections and provide for himself the
study material he needed. Clearly this had been part of the
understanding that went with the Harvard appointment. He did
general collecting, not only herpetological collecting. He had always
done so, and museum workers in that day were always, whatever
their specialty, general collectors. He did sometimes collect large
mammals, but this, I am told, was not to his preference. He did get
to Africa and to parts of it he had not seen before. In the years
between 1924 and 1940 he was away from Cambridge four times
(1925-1926, 1928-1929, 1933 1934, and 1938-1939). On each
occasion he was away a full year. In terms of his additions to the
Harvard collections, this was his prime time.
The first years were active years in many ways. These were years
of affluence for the Museum. Barbour's money immensely aug-
mented collections that Louis Agassiz had been at feverish pains to
acquire. Although Loveridge's African expeditions were undoub-
tedly the greatest source of additions to the herpetological
collections during these years, Barbour did not cease, so long as he
was able, to encourage and directly finance every sort of acquisition
from any part of the world. This flood of material was certainly
Loveridge's joy.
Loveridge was something of a public figure in the first years. He
routinely gave lectures, wrote articles for "Fauna," "Frontiers," and
"Natural History" and in 1928 gave a series of twenty lectures for
the Boston Society of Natural History on Boston's WBET entitled
"Tales from Tanganyika." He made "Who's Who" in 1938.
There is much to indicate that the world changed for Loveridge
after the 30s. The Depression had come; if its impact was not
immediate, it was fundamental. The concomitant diminution of
Barbour's fortune meant that the flood of specimens began to come
to an end. (Loveridge once showed me how plainly this change was
demonstrated on our species cards.) It was later in this period of
diminished affluence that Loveridge refused to take more than two
of a series, offered by Vanzolini, of a species not represented in the
MCZ collections; "Bottles," he is reported to have said, "are
precious." For some time the momentum of previous activity
continued. By 1942, the number of species and subspecies in the
4 BREVIORA No. 471
collection surpassed 6,000. The Department had to be enlarged, and

a new room (the old Aquarium) was taken over for snakes.
But already in the previous year (1941), Barbour's report as
Director had begun to take a mournful tone: "Increased taxes are
going to make it difficult or impossible for the Museum to expect to
receive the private assistance which it has received in the past."
In 1942 Shreve left for the army. In 1942-1943 only 400 specimens
were catalogued; 140 of these were exchanges. Loveridge's own
collecting suffered also. There was to be only one more African trip.
Decreased curating and collecting did not impair Loveridge's
productivity, however. His previous work had been primarily
reports of collections and faunal studies. He now began revisions,
and in a popular vein began the series of books that gave him wider
fame. "Many Happy Days I've Squandered" (1944) was the first.
"Tomorrow's a Holiday" succeeded it in 1947, then "I Drank the
Zambesi" (1953), and "Forest Safari" (1956). All included accounts,
highly entertaining, of his African experiences.
The end of the Second World War had brought some bonuses.
Shreve came back. W. H. Stickel, Sergeant Beck, Captain Jarvo, an
Australian, Gunner Tovell, and others sent to Loveridge material
from the Pacific area collected during their service. Loveridge had
written a little book, "Reptiles of the Pacific World" (100,000 copies
were printed for the Armed Forces, and it has recently been
reprinted), and these collections were its rewards. Loveridge
dutifully reported on the collections and on similar material ob-
tained by the National Museum.
(It is curious that the most massive of all the acquisitions since
Loveridge were the result of these Lov-
activities peripheral to
eridge's major Correspondence with Fred Parker in 1960

interest.
was initiated by a request for Loveridge's Australian and New
Guinean papers and has resulted in the MCZ's now huge Solomons
and New Guinea collections.)
Loveridge remained in charge of the collections for almost ten
years more. He madethe last African expedition of his Harvard
career, that to Nyassaland and Tete in 1948-1949. Thereafter, the
entire period was devoted to his intended summary of East African
herpetology, most of it to the series of revisions that he had begun

earlier, in 1940, some snake genera, and that culminated with
with
the East African Check List published in the year of his retirement,
1957. One paper on "The Cryptodira of Africa" was in collaboration
with the man who was to succeed him — myself— published again in
1957.
Loveridge left Cambridge in 1957, immediately after his retire-
ment, for the island of St. Helena in the South Atlantic. Although
he did visit England, and I once saw him in the British Museum, and
although we maintained a correspondence and he even published on
material he sent to the Museum, he never returned to the United
States. It is not known why — perhaps because the collection was no
longer his in the special sense that it had been for thirty-three years.
The Boston Globe of July 21, 1957 headed its four column
account of Loveridge's retirement with the statement: "Retiring
Curator to Avoid Work Temptation." If that was genuinely
Loveridge's intention, it did not turn out that way. He did make his
retirement home at Varney's on St. Helena, but it often seemed that
he was only a little less active in these final years than he had been in
the MCZ.
He kept up an intense interest in both African herpetology and
the Museum and in collecting: his letters of 1958 are full of
impatience to get the tubes to collect St. Helena spiders. His

correspondence, in fact, began on the boat to the island, and he was
soon to start numbering his letters. There were already 2,472 in early
1965, and they were nearing 7,000 at the time of his death. (He was a
punctilious writer, always answering a letter, but always insisting
also that his letters be answered before he would write again.)
He travelled also, not infrequently to England, and at least once
collected again in Africa —
a small collection —
Chamaeleo. Mabuya,
and frogs from Mau Narok at 9,000 ft. in Kenya, donated to the
MCZ. He received specimens from correspondents
also and
published on some of them, reporting Hemidactylus mercatorius as
new to Ascension Island and describing new amphisbaenids
collected by lonides in Tanganyika. His most interesting paper from
his "exile" on St. Helena may be unique in herpetology: his own
report, published at his own expense, on "The status of new

vertebrates described or collected by Loveridge."
His wife died suddenly on St. Helena in 1972. His son Brian
joined him on the island four years later. In another four years
Loveridge himself was dead.
What of the man behind the Curator? He was, of course, a very
special individual in his own right, but he was also one of a breed
that is now extinct because the times have made its life style no
longer viable.
6 BREVIORA No. 471
Born in Penarth, Glamorgan. Wales, 28 May 1891. Loveridge was

thirty-three when he came to the Harvard Museum. He had already
been Curator in Nairobi and served in museums in Wales and
England. As he reports in "Many Happy Days Tve Squandered." he
had decided to become a Museum Curator at the age often. He tells
in the preface of that book of "the acquiescence of a kindly father."
However, the Harvard Archives contains his application in 1914 for
the newly created post of Curator at the Nairobi Museum. This
reveals that he had to "serve time" for two years as apprentice in the
family business of ship furnishing, and that only then was he

allowed to take a year's course in Zoology and Botany in the
University College of South Wales on the way to appointments first
at the Manchester University Museum and then in the Temporary
Museum in Cardiff.
It was while he was inthe latter post, and. in addition to his
regular duties, making a card index of the whole British Fauna

(about, he reports, 23,000 cards), that he serendipitously received
knowledge of an open position in Africa. Although he already had a
private collection of "nearly 250 jars of preserved reptiles and over
300 glass topped drawers containing birds' eggs, insects and other
specimens," he was always avid for more. When he heard about a
civil engineer from British East Africa due home on leave who "had
in his youth shown a fondness for snakes," he tried to inveigle the

man stamps and
into collecting for him. Utilizing a joint interest in
bribing him with duplicates of these, Loveridge extorted a promise

to pickle lizards and snakes. An inquiry six months later produced
an apologetic reply which included the news that the East Africa
and Uganda Natural History Society needed a curator for a new
museum that would have government support. "Why don't you
apply for the post and then you can collect your own bally snakes"
was the advice. Loveridge applied at once, was accepted, and
arrived in Nairobi in mid-1914.
The first World War very soon cast its shadow over Africa; it did
not interrupt Loveridge's career as a naturalist. Although he joined
the local forces shortly after his arrival and after six months'
training was on active duty, it is often difficult, from his account of

the next four years, to be conscious that a war was on. The
occasional moment of danger was memorable for him because of
the capture of a rare animal. His story of the capture of his first
Boulengerula houlenfêri is characteristic: "This rare Caecilian was

1982 LO\ ERIDGE'S LIFE IN RETROSI'ECl 7
obtained under rather unusual circumstanees during the Hast

African campaign. We were busily engaged in 'digging in' under an
unpleasant shellfire. when it was unearthed b\ one ot m\ fellow
troopers inMounted Riiles. He hurnourouslv called out that one
the
of my snakes had escaped and that, if did not come over and take 1
charge of it at once, he would run his bayonet through it. Needless

to say, w hen saw what it was, 1 very gladly took charge of it." The
i
same total devotion to natural history made him as he himself

recounts — badger
first his sergeant and then the general in the midst
of his staff for permission to obtain bottles of pickled snakes from

an abandoned German house.
He got the snakes; this was no isolated incident. One of his
periodical summaries of his activities preserved in the Harvard
Archives mentions: "Travelling through German East Africa from
north to south provided exceptional opportunities for collecting . . .
All necessary preservatives and pickling jars were 'found' in
captured German towns." (The 'found' is in quotation marks in liis
own typescript.)
We have here the image of a man wholly de\ oted to an avocation
that he made his vocation and who found his life "one long holidav.
Gratefully I confess to being one of the favored few whose waking
thoughts in the
morning consist of the pleasant planning of the dav's
work." It was he who also wrote: "Probably only a zoologist can
look at an uncaught cobra and feel the jov- a child feels on Christmas
morning."
It is this spirit of Loveridge that is well-caught in the cartoon that
I have chosen to illustrate this memorial of a life, it is a man would
I
have liked to have known. It is not. 1 think, the man knew.

1
came into Loveridge's ambience late, in 1947. after the Second

I
World War, and while was working on mv' thesis. Visiting the
I
Museum. was able to re-identify one or two turtles and so gained

I
his confidence. This began a cordial relationship. Eventually, after 1
came to Harvard, I was able to call him "Arthur." (The first level o\'
intimacy was "Loveridge" rather than "Mr. Loveridge.")

The man knew was stiffish. Some called him "Sir Arthur." Ihis
I
was probaby mere Englishness plus a firm insistence on standards

that he did not allow to be relaxed, not for himself, not foranvone.
Romer called him the "Demon Curator," and this was the aspect
that most of us knew, who knew him late in his career.
He could be kind and very helpful. Manv of his correspondents.
8 BREVIORA No. 471
the visitors to the Department, the young questioner, and even some
of the anatomists in search of specimens for study knew his kindness
and assistance well. The more demanding might get short shrift, and
for poseurs — —
and frauds so he regarded Ivan Sanderson he had no
kindness at all. His review of a book of Sanderson's, and of its
gentler reviewers, is classic vitriol.
His tidiness extended to classification.He preferred clarity, was
unhappy with complication, was impatient of subtlety. He wanted
problems solved cleanly, once and for all. Therefore he was very
much a lumper. He was so much a lumper that many of our species
cards record the species name with an interval between the genus
—
and species name an interval for the eventual insertion of the
species name of which Loveridge was sure the taxon in question
could only be a subspecies. (He was very insistent also that
subspecies be readily recognizable from museum material. He
reportedly gave K. P. Schmidt the chance to sort out unlabeiled
MCZ specimens into subspecies that K. P. was describing: K. P.
flunked. It is known that he gave a similar test to Vanzolini on the
subspecies oi Amphisbaena fuUginosa that Vanzolini passed hand-
somely. Vanzolini is now not certain that one of his subspecies is
valid.)
This fervor for lumping and tidiness sometimes caused disagree-
ments. I was invited into collaboration with him on the
Cryptodira
of Africa. As he told me, this was partly because I knew turtles, but
also because I could read German and translate type descriptions.
That the collaboration succeeded is evidenced by a thickish volume,
but there were moments of discord. My discussions were too
theoretical and too verbose, and my taxonomy too splitting.
Loveridge told me that he had lost a year of his scheduled program

because of me, and a well-known footnote (softened at the advice of
his wife) testifies to our taxonomic disagreement. (It was character-
istic of him that this did not impair a good relationship.)
I, and others of my time, knew Loveridge only in the Curator-
facet of his life. We knew him after his last field trip. That part of his
life had ended.
But more, I think, was gone by then than just the opportunity for
year-long field trips. The world had changed. The British Empire
was diminished if not extinct. Africa had changed. Harvard had
changed. His chosen profession as naturalist-curator was no longer
highly regarded at Harvard, or elsewhere. Africa was not the same
1982 LOVERIDGE'S LIFE IN RETROSPECT
Cartoon of Arthur Loveridge from the newspaper East Africa.

10 BREVIORA No. 471
land in which Salimu, his favorite No. 1

Boy, had chosen to be
naturahst-servant to a naturalist-master.
It is notable that he did not choose to go back to Africa upon his
retirement. There is a story that his wife chose isolated St. Helena
because they had once stopped there on their way to Africa in one of
thetwo months of the year when the climate is pleasant. But surely,
even if that were true, the choice involved more than that. Africa
was no longer the Africa he had loved.

The man of the later years was not. at least on the surface, the
man one would e.xpect to write a book with the title "Many Happy
Days Tve Squandered." The man seemed sterner and more prim,
—
more New England as though he had acquired the characteristics
of the New England breed for whom the verb "to squander" borders
on obscenity.
Loveridge's aspect as Demon Curator provoked as much
astonishment as appreciation. Romer, the new Director after
Barbour, regarded Loveridge with a respect not unmixed with
amusement. Their psychologies were nearly antithetical, and
Romer's appreciation of Loveridge's value was very incomplete. It is
true that Loveridge could not have been quite happy with Barbour's
exuberant and insouciant carelessness, but at least Barbour and
Loveridge were of one mind about collecting and collections.
Romer marked the first of the transitions to another museum style.
The element of fanaticism in Loveridge's neatness quite naturally
evoked legends. There is a tale that there was in the Department a
drawer labelled "string too short to use." Neatness and routine were
at times extreme. Shreve's work counter had to be cleared at 4:30
when he left. Books had to be put back. Loveridge told both Carl
Cans and Vanzolini that gaps on the book shelves were to him like
teeth that had been knocked out. Even the chairs had to be in
correct positions under the counter. was reprimanded one
I
Monday, when, working over a weekend, I left all three chairs

improperly aligned.
Loveridge's fanatic passion for his collection astonished his
colleagues; clearly he did not fit too well in the new world at
Harvard. But fanatic attention to detail is a good thing in a curator,
and certainly the Museum was well served by Loveridge's devotion.
The organization of the herpetology collection was his and his
alone; the task that confronted him when he first arrived must have
been fabulous, and the order he achieved remains an achievement as
great as his African contributions. When 1 took over the collection,

all was in perfect shape. There were few curatorial tasks to do: only
the one collection from the Riu-Kiu Islands that had not yet been
wholly identified and put away.
Loveridge served in a University Museum, but he was in no sense
an academic. He belonged to another generation and another life
style
—he was pre-eminently a collector-naturalist. It is interesting to
put him in context in the succession of herpetological curators at the
MCZ. Agassiz must be counted here, but he was clearly unique a —
European emigre, professor, builder and acquirer of collections,
intellectual parent to whole generations of natural historians in the
United States, he is not at
comparable to anyone else.
all
Of Garman. we know too little, not much beyond

the next in line,
Barbour's unsympathetic remarks and some plaintive autobio-
graphical notes of his own. Before the MCZ, he apparent!}' had an
irregular career; at the MCZ, fide Barbour, he remained isolated
and apart. Clearly in his day he was useful and respected. He began
the MCZ's West Indian interest which Barbour so much cultivated
after him.
Barbour had. in contrast, the full academic panoply, a doctoral
degree, and, at least late in life, professorial status. For all that, he
was throughout his life the Wealthy Amateur, never quite willing to
go very deep, never quite serious enough to be professional. He
could be pontifical and very disparaging of others, but many of the
criticisms could have been turned against him. He had notable
proteges
— —
G. K. Noble and E. R. Dunn and was. within American
herpetology. for a while something of a father figure with equal
colleagues but no admitted superior.
Loveridge was very different. -British always and a Briton of the
Empire, he was a man who. without inherited wealth, had chosen,
—
very stubbornly, a poorly remunerative career the career of Bates
and Wallace, the naturalist-collector. He had chosen also a
continent. His eyes and his interests turned eastward toward Atrica;
he was hardly part of American herpetology. For all that in
Cambridge he was physically close to his American colleagues, it
was hardly different from what it might have been had he been
across the sea. If ever he was further west than New York. ha\e no 1
record or report of it.

In a sense that Barbour was not, Loveridge was a professional.
Without interest in theory or in biology beyond field and museum
12 BREVIORA No. 471
natural history, he was professional in what he did

totally
—
completely dedicated to competence in that rather narrow area. He
never formally taught, and his own formal instruction was limited.
In herpetology. like hrs'predecessors (and like myself), he was self-
taught. (The new curator at MCZ

is the first to have had formal
specific instruction in herpetology.) What he chose to be. he was/?a/'

excellence —
Curator-Collector-Naturalist.
With Loveridge's departure
— and his literal departure from
Cambridge for St. Helena came only a few weeks after his formal
retirement as Curator — the Department settled into quite another
style. I was thecurator to be fully a product of Academia -not
first
only the holder of the conventional degrees, but one who needed
them for a living. The world has changed post-Loveridge; the pure
naturalist-collector is. when he e.xists at all. an anachronism.
Arthur and Mary Loveridge's one son. Brian, was schooled at

Harvard, and had early gone to England for his career. Apart most
of their lives. Brian joined his father on St. Helena and was building
a home there near Varney's when his father died. Brian has said of
his father that his work was his life. That is an affirmation that is
also a tribute, and the tribute that Arthur Loveridge would have
most wanted.
In one of the entryways to the MCZ there is a plaque on the wall
honoring Alexander Agassiz and with the Latin motto, "Omnia

quae hie vides monumentum." The Herpetology Department might
very reasonably display a similar motto in Loveridge's honor. The
collection's order and style have his imprint. have added somewhat
I
to that collection, but he provided the solid base.

His taxonomic work is now history; his revisions are now revised.
It would disappoint him bitterly that this is true, but his passion for
simplicity is now judged to have gone too far. He was concerned in

his "Status" —
paper to learn certainly with a twinge of heart which
—
of his species had been synonymized. He would view with dismay, if
not distaste, the sibling species that are now commonplace. He
would not understand the concerns and disputes of modern
taxonomists, nor care to. In this sense time has passed him by, but
his own collections and the collections he so diligently curated are
his enduring monument.
'MAR 1 8 iQfls
HARVARD
B R E V I'U'R A
Miiseiim of Comparative Zoology
us ISSN ()()()() 9G9S
Cambridge, Mass. 30 June 1982 Ni:mhir472
FISHES OF THE SUBORDER LABROIDEI

(PISCES: PERCIFORMES):
PHYLOGENY, ECOLOGY, AND
EVOLUTIONARY SIGNIFICANCE
Leslie S. Kaufman' and Karel F. Liem-
Abstract. We the
Pomacentridae, Cichlidae, Embiotocidae,
postulate that
Labridae, Odacidae. and Scaridae

comprise a monophyletic assemblage: the
Labroidei. Four groups within the Labroidei can be defined as monophyletic
assemblages on the basis of shared derived characters: the Pomacentridae, Cichlidae,
Embiotocidae, and Labridae (which includes the Scaridae and Odacidae). The
Pomacentridae considered the primitive sister group of all other Labroidei; the
is
Cichlidae is a sister group of embiotocids and labrids, and the Embiotocidae is a

sistergroup of the Labridae. Labroids are characterized by (I) united or fused fifth
ceratobranchials resulting in the formation of one functional unit: (2) a true
diarthrosis between upper pharyngeal jaws and the basicranium without an
inter\ening part of the transversus dorsalis anterior muscle: and (3) the presence of
an undivided sphincter oesophagi muscle forming a continuous sheet. It is proposed
that ( the ecological and functional versatility of the trophic apparatus is correlated
I
)
with a characteristic structural design, and that (2) this design has contributed to the
dominant position of labroids in diurnal communities of tropical marine and lentic
fresh waters.
INTRODUCTION
Liem and Greenwood (1981) have recently reviewed the compara-
tive functional morphology of the pharyngeal jaw mechanism in
acanthopterygian fishes. On the basis of functional considerations
they have proposed that the Cichlidae, Embiotocidae, Labridae,
Odacidae, and Scaridae comprise a monophyletic assemblage.
'-Museum of Comparative Zoology, Harvard University. Cambridge, Massa-

chusetts 02138.
2 BREVIORA No. 472
Pharyngognathy, as expressed in the Cichlidae, has been correlated

with several functional and ecological attributes which distinguish
cichlids from most other Acanthopterygii. First, cichlids can
mechanically process a broader range of food types (Liem, 1974).
Second, they have greater feeding versatility (Liem and Osse, 1975;
Liem, 1980) and hence a broadened fundamental niche {sensu
Hutchinson, 1958, 1965). Finally, cichlids exhibit an extremely high
speciesdiversity, perhaps because extinction rates in changing
environments are decreased (Liem, 1982). To test this hypothesis,
evolutionary patterns in the Cichlidae must be compared with those
of related fishes both more and less specialized with respect to
pharyngeal jaw morphology. Such an analysis requires a thorough
knowledge of phylogenetic relationships (Lauder, in preparation).
In this paper we offer a revised hypothesis of the phylogenetic

relationships of pharyngognath acanthopterygians first proposed by
Liem and Greenwood (1981). New morphological evidence offered
here and by Stiassny (1981, personal communication) requires the
inclusion of the Pomacentridae in the monophyletic assemblage
Pharyngognathi (sensu Liem and Greenwood, 1981), and a re-
arrangement of the Embiotocidae as the sister group of the Labridae
instead of the Cichlidae. The resulting scheme of classification is as
follows:
Suborder Labroidei
Family Pomacentridae
Family Cichlidae
Family Embiotocidae
Family Labridae (including Labridae, Odacidae, Scaridae)
The proposed phylogenetic relationships provide a basis for
assessing patterns of change in the feeding apparatus, historical
consequences of new feeding mechanisms, and the evolution of
coral reef fish communities.

Morphological studies were conducted with the aid of a Wild-M5
dissecting microscope and camera lucida. Clearing and staining
followed the techniques of Taylor (1967). Scanning electron
M
microscopy was conducted on an A R- 000, and x-ray cineradiog-
1
raphy of feeding labrids and cichlids was carried out using the
Siemens Cineradiographic Unit at 150 frames sec '. The following
material was examined:
1982 I ABROII) l>HV1.0GENY 3
Pomacentridae: Ahucli'fchif launis MCZ 42755. Aniplupiitifi \(inihtirti\ MCZ

14852, A. perculii MCZ 33399. Dascvllus iiimaculaia MCZ 14837. /). ulhisclla MCZ
51671. EupomcHcnlnis planifrons MCZ 44745. accipiiUcnsiis MCZ 43961.
/:'.
Panuncnirus littoialis MCZ 5794. Chromis alrilohaius MCZ 44640.

Cichlidae:
"
Haplm hraDiis" /cmlscusMCZ 49517.
Embiotocidae: Rlumnhilus vacca MCZ 57708. Panwlicliilivs vacca MCZ 54333.
CvmaiofiiisU'r ai;i;rciâui MCZ 57707. Plninerocion Jurcatu.s MCZ 54334, Emhioioca
javksoni MCZ 54332.
Labridae: TauioKolahnis adspcrsus uncat.. Taiitoâ oniiis iincat.; Halichoeres
hiviiiamm. Scariis croicensls. Spari.snnui viridc. all MCZ acq. 1981-002-6.
Caribbean reef fishes were studied in Salt River Canyon and

Tague Bay. Croix, U.S. Virgin Islands, and also studied at the
St.
Discovery Bay Marine Laboratory, Jamaica, W.I. Ecological

classifications of reef fishes were based in part on observations made
from the NULS-I Hydrolab during mission 81-8.
ANALYSIS OF CHARACTERS
Definition of the Lahroidei
We postulate that the Pomacentridae. Cichlidae. Embiotocidae.

Labridae. Odacidae. and Scaridae comprise a monophyletic lineage,
the Labroidei. A cladogram defining this group and expressing
relationships among its major clades has been derived on the basis
of three investigations: Liem and Greenwood (1981). Stiassny's
analysis of the phylogenetic relationships of the Cichlidae (in which
extensive out-group comparisons are described, Stiassny, 1981 and
personal communication), and this study (Fig. 1).
All Labroidei share the following three derived characters: (1)
junction or fusion of the two fifth ceratobranchial bones into a

single unit, (2) diarthrosis between the upper pharyngeal jaws and
the basicranium (Fig. 2 A
F: APU; Stiassny, personal communica-
tion), and (3) the presence of the sphincter oesophagi muscle as a

continuous sheet, with no dorsal subdivision (Fig. 2; Stiassny
personal communication). Within Perciformes, fused or joined
lower pharyngeal jaws also occur among the Anabantidae (all),
Kyphosidae (Girella tricuspidata), and Sciaenidae (Pogonias chro-
mis and Aplodinotus gnmniens). The pharyngeal jaw morphology
and biting mechanisms of these fishes differ appreciably, however,
from those of the Labroidei (Liem and Greenwood. I98I). Other
acanthopterygians show some form of articulation between the
upper pharyngeal jaws and the basicranium (e.g.. Sparidae,
BREVIORA No. 472
Figure 1. Cladogram illustrating interrelationships of the major labroid clades.

Bars represent shared derived characters: (1) United or fused fifth ceratobranchials;
(2) True diarthrosis between upper pharyngeal jaws and basicranium; (3) Undivided
tissue joining lower jaw
sphincter oesophagi muscle; (4) Strong sheet of connective
with a ligament which inserts on the ceratohyal bone; (5) Nipple-like bony process
on ventral surface of lower pharyngeal jaw; (6) Pharyngo-cleithral articulation of
characteristic form; (7) Obliquus posterior dominant muscle to lower pharyngeal
jaw; levator externus 4 and posterior vertically aligned on fourth
obliquus
epibranchial, separated by oblique aponeurosis or tendon; (8)
Transverus dorsalis
muscle subdivided into four parts; (9) Premaxillae and maxillae functionally
decoupled; (10) Cartilagenous cap on anterior border of epibranchial 2; (11)
Micro-
branchiospinae of characteristic form present on outer faces of second, third, and

fourth gill arches; (12) A: and Av. portions
of adductor mandibulae complex lacking
major structural association; insertion of large ventral division of A: onto angulo-

articular; (13) Head of epibranchial 4 distinctly expanded; (14) Intra-uterine
development of young with strongly modified vascularized median fins; (15) Mus-
cular sheet joining Ai and Am portions of adductor mandibulae; (16) Levator
posterior dominant muscle to the lower pharyngeal jaw, forming a force couple with
the pharyngocleithralis muscle; (17) Toothplates of fourth pharyngobranchials
absent (either lost or fused with pharyngobranchial 3), first pharyngobranchials
absent or reduced; (18) Fourth epibranchials highly modified, articulating with
upper pharyngeal jaws; (19) True pharyngo-cleithral articulation functioning as
sliding and hinge joint; (20) Levator externus 4 is a continuous muscle joining
for insertion of
prootic region to muscular process on lower jaw; (21) Predisposition
levator posterior muscle on lower pharyngeal jaw; (22) Loss of second pharyngo-
branchial toothplates; (23) First three branchial adductor muscles cover antero-
dorsal faces of the epibranchials; (24) Ligament connecting postmaxillary process of
maxilla with anterior border of palatine and ectopterygoid; (25) Tooth rows
arranged radially across the lower pharyngeal jaw, teeth located directly over the
symphysis between left and right fifth ceratobranchials. LPJ toothplate composed of
an anterior, small-toothed field and a posterior, large-toothed pavement replaced by
addition along the rear margin of the LPJ.
1982 LABROID PHYLOGENY 5
Gerreidae, Pogonias, Aplodinotus), but only in the Labroidei is

there a true diarthrosis. In other perciforms a portion of the
transversus dorsalis muscle or its aponeurosis passes between the
apophyses of the upper pharyngeal jaws and basicranium (Stiassny,
1981).
Synapomorphies Characterizing the Pomacentridae

The damselfishes can be defined on the basis of four characters.
(1) Stiassny (1981: 286) observed that "A strong sheet of connective
tissue originates from the dorsal border of the bony ridge on the
medial face of the lower jaw [dentary] and merges with a cylindrical
ligament that passes posteriorly and inserts onto the ceratohyal
bone." In other acanthopterygians she examined this ligament was
wanting. The remaining three characters concern the structure of
the lower pharyngealjaw (LPJ). (2) The LPJ's of all pomacentrids
we have examined bear on their ventral surfaces a pair of small
nipplelike processes, which serve as the insertion sites for the

pharyngohyoideus muscle. These processes are absent in all other
acanthopterygians examined. (3) In primitive acanthopterygians
there is no contact between the fifth ceratobranchial and the
cleithrum. In most pomacentrids, however, the muscular processes
of the LPJ abut upon the cleithrum and slide along it by means of
articular facets. Two such facets may be present (e.g., Pomacentrus
littoralis, MCZ5794): a dorsal facet lying parellel to the dorso-
ventral plane, and a ventral facet which is curved slightly outwards
from this plane and may provide the LPJ with some lateral freedom
of movement. Labrids, in contrast, have a true pharyngo-cleithral
joint.The degree of pharyngo-cleithral articulation varies con-
siderably among pomacentrids. Even when the two bones are
closely related, the nature of the articulation differs from that seen
in the Labridae (Liem and Greenwood, 1981). This difference is also
reflected in the unique and complex shape of the muscular processes
of the LPJ pomacentrids, a feature related to their peculiar
in
musculature (Figs. 2, 3). In some pomacentrids (e.g., Microspatho-
don) pharyngo-cleithral articulation appears to have been lost as
part of a general reduction of the pharyngeal apparatus. (4) In
pomacentrids, as in all more primitive perciforms, the fourth levator
externus muscle (Fig. 2 A-F: LE4) and levator posterior (LP) insert
BREVIORA No. 472
RD TOP
Figure 2. Dorsal aspect of the branchial musculature viewed from posterior to

elucidate the muscles surrounding the esophagus and posterior branchial arches.
A) Pomacentrus litioralis; B) Abudefduf taurus; C) Taulogolabrus adspersus;
D) Amphiprion xanthurus; E) "Haplochromis" leuciscus; F) Embiotoca jacksoni.
Abbreviations: AD, adductor branchialis; APU, apophysis of upper pharyngeal

jaw (third pharyngobranchial); CBs, fifth ceratobranchial (lower pharyngeal jaw,
LPJ); EB, epibranchial; ES, esophagus; LE, levator externus muscle; LI, levator
internus muscle; LP, levator posterior muscle; OD, obliquus dorsalis muscle; OP,
obliquus posterior muscle; PB, pharyngobranchial; RD, retractor dorsalis muscle;

SE, sphincter oesophagi muscle; TDS, transversus dorsalis anterior muscle; TDP,
transversus dorsalis posterior muscle.
1982 LABROID PHYLOGENY
<
8 BREVIORA No. 472
on the dorsal aspect of the fourth epibranchials (Liem, 1974).

However, the pomacentrid LE4 (Fig. 2A,B,D) splits near its
insertion site on the fourth epibranchial (EB4). The larger head
inserts near the insertion site of LP, while a smaller medial head
joins an oblique aponeurosis that runs anteromedially along the
border of EB4. The obliquus posterior (OP) is the major muscle
operating the posterior region of the LPJ. The OP
muscle runs
between the broad dorsal flange of each muscular process and the
flange along the EB4 just below the insertion site of LE4 (Fig. 2A, B,
D). Here the LE4 and OP muscles are separated by the oblique
aponeurosis. Thus the sites of origin and insertion for OP in
pomacentrids are the same as the other acanthopterygians, but the
extremely close relationship between LE4 and OP on the fourth
epibranchial is a derived character. The aponeurosis is present in all
pomacentrids examined, and it clearly separates the fibers of the
LE4 from those of the OP. Nevertheless, the insertions for the two
muscles are extremely close together, a condition which seems to
foreshadow the muscular sling of cichlids and other Labroidei.
There is a broad flange on EB4 to receive LE4, and the posterior
flanges on EB4 and the fifth ceratobranchial (CB?) are vertically
ahgned. Between them, and possibly contributing to the mechanical
linkage between the two, is the stout dorsal portion of CB4. Thus,
while the muscles of the LPJ in pomacentrids still originate and
inserton the same bones as in non-labroid acanthopterygians, the
geometry of these insertions has been modified in a characteristic
fashion. The resulting condition may represent a primitive counter-
part to the cichlid muscular sling (Liem, 1974).
Synapomorphies Characterizing the CichHdae

Six characters clearly distinguish the Cichlidae as a monophyletic
assemblage: (1) The transversus dorsalis muscle is subdivided into
four parts (Liem and Greenwood, 1981; Fig. 2E:TDA). (2) The
premaxillae and maxillae of all cichlids are functionally decoupled;
i.e.,premaxillary protrusion can be regulated independently from
motion of the maxillae by means of multiple mechanical pathways
(Liem, 1978, 1979). Four additional characters have been described
by Stiassny ( 98 ): (3) There is an extensive cartilaginous cap on the
1 1
anterior border of EB2 (Cichla ocellaris being the exception). (4)

Microbranchiospinae of characteristic form are present on the outer
faces of the second, third, and fourth gill arches. (5) The A: and Au
portions of the adductor mandibulae complex have lost a major
structural association, and there is an insertion of a large ventral
division of A: onto the angulo-articular. (6) The head of EB4 is
distinctly expanded.
Synapomorphies Characterizing the Embiotocidae

Two characters are considered synapomorphies for the Embio-
tocidae. (1) All embiotocids are fully viviparous fishes, which deliver
large, well-developed young. The males have a small intromittent

organ derived from anal fin rays. The young are usually closely
packed in the ovarian sacs which function in the fashion of a uterus.

Monophyly of the embiotocids can be established on the basis of the
highly specialized mode of "intra-uterine" development with its
associated structural, physiological, and behavioral features, invol-
ving spatulate extensions of the webs of median and caudal fins,
each with a rich blood supply (Webb and Brett, 1972). (2) Stiassny
(1981) has found a small sheet of parallel muscle fibers arising from
the anteromedial region of the Ai part of the adductor mandibulae
muscle and inserting upon the dorsal aponeurosis of A2+3 part of this
muscle.
Synapomorphies Characterizing the Labridae

The families Labridae, Odacidae, and Scaridae have been
recognized as close relatives within the Labroidei (Greenwood et al,
1966). Here the three groups are recognized as a single family, the
Labridae, since the monophyletic nature of this assemblage is
strongly indicated by both morphological and functional characters.
(1) The levator posterior muscle (Fig. 2C:LP) is the dominant
muscle of the LPJ, forming a force couple with the pharyngo-
cleithralis externus muscle (Liem and Greenwood, 1981). (2) The
toothplates of the fourth pharyngobranchials are absent (either lost

or fused with pharyngobranchial 3; Stiassny, 1981), while the first
pharyngobranchials are absent or reduced. (3) The fourth epi-
branchials are highly modified and of characteristic form, articulat-
ing with the upper pharyngeal jaws (Yamaoka, 1978, 1980; Gobalet,
1978). (4) There is some form of physical contact between the LPJ
and the cleithrum in all members of the clade (Liem and
Greenwood, 1981).
10 BREVIORA No. 472
The posterior face of the muscular process on each side of the

LPJ bears an articular facet, which fits against a small fossa on the
cleithrum. The mechanics of this joint are quite complex. Cine-
radiography of the LPJ in Tautogolabrus adspersus during mastica-
tionshows that there is a biphasic pattern involving both a hingelike
and a sliding movement. The dynamics are reminiscent of those in
the human temporo-mandibular joint. At rest and during the bite
the LPJ is in close contact with the cleithrum and exhibits the
pattern of a hinge-joint. During protraction the LPJ slides down
and forward, using the anterior face of the cleithrum as a track. The
morphology of the pharyngocleithral joints of odacids and scarids is
basically similar to that of labrids, though the support system is
stronger, and it is unknown whether the LPJ disarticulates during
protraction. The condition in odacids and scarids overlaps broadly
with that of the Labridae. Cryptotomus roseus is a scarid with a
labridlike LPJ; both it and Nicholsina denticulata have many
wrasselike features (see Gobalet, 1980 for comparative discussion of
Nicholsina). Pseudodax moUucanus is a labrid with a scaridlike LPJ
(Bleeker, 1862).
Interrelationships Among the Labroidei
The from those of previous

relationships proposed here differ
investigators (Greenwood al, 1966; Nelson, 1967; Liem and
et
Greenwood, 1981) in three important ways. The Pomacentridae is

considered the primitive sister group of other Labroidei. The
Embiotocidae is postulated to be the sister group of the Labridae,
and not the Cichlidae. Finally, the Labridae, Scaridae, and
Odacidae are united in a single family, Labridae, to reflect striking
similarities in their morphological as well as functional specialized
features (Fig. 1).
Chief evidence for the proposed primitive sister-group relation-

ship of the Pomacentridae to other Labroidei is found in the
osteology and myology of the LPJ. In the Cichlidae, Embiotocidae,
and Labridae, the LE4 and (in general) LP muscles insert on the
LPJ. The LE4 and LP muscles of Pomacentridae insert on the
fourth epibranchials, which represents the primitive condition in the
Perciformes. Aerts and Verraes (1982) have demonstrated that the
LE4 of the cichlid Haplochromis elegans (and presumably, other
cichids as well) is actually a composite of LE4 plus a large medial
head of the obliquus posterior muscle, the two fusing during
development. This condition is never fully developed in Poma-

centridae (Fig. 2A, B, D: OP, LE4). Obliquus posterior (OP) is the
dominant muscle to the LPJ while LEj remains a separate muscle,
though it is large, and is aligned with the OP. The result is that the
two muslces together bridge the gap between prootic and LPJ,
meeting across the fourth epibranchial, where they are separated by
an aponeurosis (Fig. 2A, B, D: OP, LEj).
Four derived characters establish sister-group relationship be-
tween the Embiotocidae and Labridae. (1) These groups have lost
the second pharyngobranchial toothplates (Nelson, 1967; Stiassny
1981). (2) The first three branchial adductor muscles cover the
anterodorsal faces of the epibranchials (Stiassny, 1980 and Fig. 2C,
F: ADi+:). (3) A ligament connects the postmaxillary process of the
maxilla with the anterior border of the palatine and ectopterygoid
(Stiassny, 1980). (4) Tooth rows are arranged radially across the
LPJ, with teeth located directly over the symphysis between left and
right fifth ceratobranchials. The LPJ toothplate is composed of two
fields: an anterior durophages), and a posterior
field (often lost in
field bearing the large cardinal teeth used in crushing or grinding.

New cardinal teeth are added only to the rear margin of the LPJ,
forming a conveyor-like pavement with greatest wear toward the
front (Fig. 3; see also Embiotocidae: Damalichthys vacca; Labridae:
Pseudodax mollucanus; all scarids).
The phylogeny proposed in Figure assumes that similarities in
1
pharyngeal dentition between the Labridae and Embiotocidae on

the one hand, and Pomacentridae on the other, are homoplasies. In
cichlids the LPJ toothplate is divided into left and right regions,
with the largest teeth arranged in two main rows parallel to the
symphysis, and there are no teeth located directly over the
•
symphysis. This is clearly the primitive condition, since (a) the LPJ
of labroids is derived from two separate fifth ceratobranchial bones,
and (b) this condition is displayed in those non-labroid perciforms
possessing fused or joined fifth ceratobranchials. In pomacentrids,

as well as in embiotocids and labrids, the teeth cross the plate in a
radial series and there are teeth located
directly over the symphysis.
As a sole for pomacentrids, embiotocids, and labrids
synapomorphy
this character seems too tentative. It appears independently in the
Beloniformes, another group with fused lower pharyngeal jaws,

which is clearly unrelated to the Labroidei (Collette, 1966).
Pharyngocleithral joints appear in both the Pomacentridae and the
12 BREVIORA No. 472
Labridae, but are clearly dissimilar in form. Thus in terms of the

specialized osteological and myological characters discussed above,
the pharyngeal jaws of the Cichlidae, Embiotocidae, and Labridae
resemble each other much more closely than any one of these groups
resembles the Pomacentridae.
Independent investigations (Stiassny, 1981; Liem and Green-
wood, 1981, and this study) have resulted in very similar hypotheses
of labroid phylogenetic relationships. Stiassny (1981, personal
communication) has based her studies primarily on soft-tissue
characters, while we have concentrated on functional and osteo-
logical characters. The relationships postulated in Figure 1 deviate
drastically from previous schemes. This is mainly because important
new osteological, dental, and myological evidence has emerged.
DISCUSSION
Ecology of the Labroidei
The new phylogenetic scheme of the Labroidei has important
implications ourfor
perception of the ecology and functional
morphology of this group. The gradal nature of former classifica-
tions obscures relationships and thereby masks evolutionary se-
quences, ecological diversity, and changes in functional patterns. To

illustrate this point we will briefly discuss some of the implications
of the new phylogeny for ecological concepts as they pertain to the
Labroidei.
Labroidei as defined here unites at least 1,470 species (some 5 to
10 percent of living fishes) that are extremely diverse ecologically.
Nevertheless, the majority of labroids occur within one general type
of environment: warm, slow-moving water with abundant habitat
structure. Tropicalmarine reefs are densely populated by pomacen-
trids and These are joined by embiotocids on temperate
labrids.
Pacific reefs. All four clades figure prominently in aquatic
macrophyte forests such as grass beds, kelp beds, algal reefs, or

heavily vegetated pond and stream edges. Relatively few labroids
are abundant in pelagic, soft-bottom, or strictly lotic assemblages.
Modes of life exhibited by marine labroids differ characteris-
tically from those of sympatrically occurring non-labroid and
functionally intermediate forms. We illustrate these patterns with
data from a coral reef fish assemblage observed in Salt River
Canyon, St. Croix, U.S. Virgin Islands (Table 1). The 137 species
Table 1. Relationship between functional morphology of the pharyngeal jaws

and potential anti-predator mechanisms for 137 species of coral reef fishes observed
during visual censuses in Salt River Canyon, St. Croix. U.S. Virgin islands
(Kaufman and Ebersole, in preparation).
Potential
14 BREVIORA No. 472
The data in Table 1

suggest that there are characteristic modes of
Hfe for coral reef labroids. Most of them both feed and refuge on the
reef. They are not heavily armed. They are, for the most part, either
defend a general area as
territorial or schooling. Territorial labroids
a multipurpose territory rather than occupying one specific hole or
cavity as do many inquiline gobies, blennies, or jawfishes (Opis-
thognathidae). The residents exclude potential competitors for
food, hiding places, and mates, as well as species that threaten the
integrity of the territory (Low, 1971; Thresher, 1976; Kaufman,
1977, 1979; Potts, 1977; WiUiams, 1979, 1980; Ebersole, 1977;
Lobel, 1980). Even the schooling and planktivorous labroids are
strongly reef-associated, utilizing benthic cover as the ultimate
means of escape from predators. In summary, members of the
Labroidei generally rely on the reef for refuge, exploiting those
foods which are, or can be made available without travelling over
long distances. There is an exception to this pattern. Adults of the
larger species (such as the huge Caribbean parrotfishes Scarus
guacamaia and 5". coelestinus) sometimes forage or migrate to and
from the reef as individuals. For these fishes, large size alone may be
a sufficient deterrent against predators.
Other reef-dwelling fishes exhibit modes of life that contrast with
those of the labroids. The non-labroid category in Table 1, including
such fishes as berycoids, apogonids, serranids, and lutjanids,
consists of both diurnal .and nocturnal predators which stalk small
soft-bodied prey. The intermediate pharyngeal jaw category is
dominated by two groups: armed fishes that feed on the reef by day
and rest on the reef at night, and unarmed fishes that feed off-reef at
night and shelter on the reef during the day. The heavily armed
plectognaths, chaetodontids, and acanthurids comprising the bulk
of the first group together exploit almost as broad a range of foods
as the labroids. Individually, however, they exhibit functional
limitations related to gape, mouth position, jaw mobility and
pharyngeal jaw mechanics that should seriously limit feeding

versatility in comparison to that of similar-sized pomacentrids or
labrids. This is reflected in what is known of their diet (e.g., Randall,
1967; Hobson, 1974; Reese, 1975). Relatively few of these species

enhance their own local food supply by defending feeding territories
(possibly Acanthurus sohal, Vine, 1974; Chaetodon trifascialis
[formerly Megaprotodon strigangulus], Reese, 1975). The nocturnal
off-reef predators (e.g., Pomadasyidae) prey chiefly on small

benthic invertebrates.
We postulate that feeding versatility was a chief factor in shaping
the characteristic modes of life exhibited by marine labroids; i.e.,
schooling or territorial behavior with a strong reliance on the reef

for both food and shelter. Trophic mobility can be one key to
survival when spatial mobility is limited by a high risk of predation.
The antithesis of this strategy, rarely exhibited by labroids, is to
reduce the risk of predation by adopting some active defensive
mechanism.
Labroids play a disproportionate role in determining the dis-
tribution and abundance of benthic organisms in tropical marine
hard-bottom communities (Randall, 1961, 1974; Ogden and Lobel,
1978; Brock 1979). In part this is due to broad-spectrum feeding
capabilities (hard-shelled invertebrates, coral rock, coral, algae).
Many labroids locally manipulate the substratum and its occupants
to suite their ownneeds (Brawley and Adey, 1977; Kaufman, 1977,
1979; Wellington, 1981). This constitutes a patchy disturbance to
sessile invertebrates (Kaufman, 1977; Council, 1978) and could be a
principal factor regulating food abundance for other reef organisms.
Labroid Phylogeny and the Evolution of

Acanthopterygian Feeding Mechanisms
Whenthe cichlid pharyngeal jaw mechanism was first described,
itappeared to represent an abrupt breakthrough in the acantho-
pterygian feeding mechanism, radically different from anything
known in the cichlids' presumed ancestors (Liem, 1974). Subsequent
radiation seemed to involve little modification of the basic feeding
mechanism (Fryer and lies, 1972; Greenwood, 1974). The new
hypothesis on the genealogical relationships of the Labroidei
presented here requires that these views be revised considerably.
One erroneous hypothesis was that the pharyngeal jaw complex is
unique to cichlids; present evidence rejects such a hypothesis (Liem
and Greenwood, 1981). Many features present in cichlids are
present in Labroidei. was thought to be a large
Second, there
morphological and functional gap between cichlids and their
primitive (ancestral) counterparts; there is not. The apparent gap

was an artifact of insufficient data, now bridged by primitive
labroids and certain non-labroid perciforms. Aerts and Verraes
(1982) have shown that during the ontogeny of a cichlid {Astatoti-
16 BREVIORA No. 472
LE4 splits into a lateral and a medial head. The

lapia elegans) the
medial head of LE4 unites with the medial head of the OP, thus
establishing a functionally as well as a structurally uninterrrupted
muscle between the prootic and the muscular process of the LPJ,
i.e., a compound LE4. The characteristic arrangement of LE4 and
OP in pomacentrids (e.g., Pomacentrus littoralis, Fig. 2) resembles
those early ontogenetic stages in cichlids before the compound
muscle is formed. The Pomacentridae is clearly primitive, however,
in terms of the origins and insertions of LE4 and OP. Thus the
Pomacentridae is intermediate between the more derived labroids
(cichlids, embiotocids, and labrids) and other perciforms.
Other perciforms which approach the labroid condition in one or
more respects are morphologically (and perhaps phylogenetically)
intermediate between labroids and primitive perciforms. The
anabantoids, Kyphosidae, and Sciaenidae have fused or joined
pharyngeal jaws in some members. The fifth ceratobranchials of the
Gerreidae and Pomadasyidae (especially Anisotremus surinamen-
have no true bony junction, but are in some species very tightly
sis)
bound together by strong ligaments. Certain Sciaenidae (e.g.,
Pogonias chromis, Aplodinotus grunniens) and Gerreidae have an
articulation between the upper pharyngeal jaws and the basicranium
although it is not as well developed as in labroids. The shell-cracker

centrarchid Lepomis microlophus and the molluscivorous Carangi-
dae {Trachinotus spp.) have broad, hypertrophied lower pharyngeal
jaw elements which meet closely at the midline (Kaufman and Ono,
in preparation). Nearly all of these morphologically intermediate
forms feed habitually on hard-shelled benthic invertebrates in
addition to a wide variety of other organisms, both hard and soft.
Selection favoring a broader, more inclusive diet could have been a
major factor in the early evolution of labroids. This hypothesis can
not be tested without first developing a better picture of perciform
phylogeny. However, the presence of so many intermediate forms
(one of which may represent the primitive sister group of the
Labroidei) and the intermediate characteristics displayed in Poma-
centridae, indicate that advanced acanthopterygian pharyngeal jaws
are the result of a series of morphological changes. There was no
single "adaptive breakthrough" {sensu Simpson, 1944, 1953; Liem,
1974).
Insummary, this new model of labroid relationships will permit

us toexamine the nature of evolutionary change in a structurally
complex mechanical system. Judging from the great ecological
diversity of labroids, it seems that-their specialized pharyngeal jaw
apparatus has greater structural potential and functional flexibility
than that of its more primitive counterparts. The more precise
labroid phylogenetic scheme will allow us to determine if there is a
general relationship between design versatility and historical pat-

terns of morphological change.
ACKNOWLEDGMENTS
We owe much to the useful comments made by William and Sara
Fink, Humphry Greenwood, and Dana Ono. Melanie Stiassny
contributed generously from her own observations on labroid
systematics. Karsten Hartel aided in locating study material. Ed
Seling and Al Coleman assisted in electron microscopy and
photography. William B. Hamner, Jr. made important contribu-
tions to studies of the pharyngeal jaw in labrids, and helped in
collecting specimens; William McFarland contributed many embio-

tocid specimens. We thank the staffs of the West Indies Marine
Laboratory and the Discovery Bay Marine Laboratory for assist-

ance in specimen collection and field observations. This work was
supported by National Science Foundation Grant DEB-79-00955 to
K. Liem; National Oceanographic and Atmospheric Administra-
tion Grant NULS-1 Mission 81-8 to L. Kaufman and J. Ebersole;
and National Science Foundation Grant OCE 79-12674 to J. B.C.
Jackson. This paper contribution No. 261 from the Discovery Bay
is
Marine Laboratory, Discovery Bay, Jamaica, W.I.
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uruRARY
MAR 1 8 1985
B R -^
R A
11 seiim of Comparative Zoology
L'S ISSN 0006 9698
Cambridge, Mass. 29 April 1983 Number 473
THE INTERRELATIONSHIPS OF PELYCOSAURS

Donald Brinkman' and David A. Eberth'
Abstract. An analysis of 17 characters forms the basis of a hypothesis of

relationships of seven pelycosaur genera. These genera include representatives from
most ol the higher taxa recognized b\ Romer and Price 1940). in agreement with the
(
phylogen\ proposed by Romer and Price, Edaphosaunts is considered to be more

closely related to DinieiroJon than is Ophlacodun. In contrast to the phylogeny
proposed by Romjr and Price, Opluacotlon is considered more closely related to
Dinienoiion than are I'aranop.s and Aerosauius. Three character-states that are
interpreted as being derived are shared by Casea, Varanops. and Aeru.sauru.s.
suggesting that these three genera are members of a clade distinct from the clade
mcluding Ophiacocloti. Edaphosauius, and Dinieiiudun.
INTRODUCTION
Pelycosaurs occupy a central position in amniote evolution. As a
paraphyletic taxon within the clade including mammals, pelyco-
saurs have played an important role in considerations of the origin
ofmammals. Also, pelycosaurs include some of the most primitive
known reptiles, and are an important element in consideration of
the early evolution of amniotes. Thus, an understanding of pelyco-
saur interrelationships has implications for many broader problems
of reptile diversification,
Pelycosaurs were the subject of a detailed monographic study by
Romer and Price ( 1940), and as a result are one of the best under-
stood groups of Paleozoic reptiles. Romer and Price used evolu-
tionary systematics in their study of pelycosaurs (Fig. 1

A). In line

02138. Present Address: Tyrrell Museum of Paleontology, Bo,\ 7500, Drumheller,
Alberta, Canada, TOJ OYO.
-Department of Geology, University of Toronto, Toronto, Ontario, Canada, M5T
IR7.
BREVIORA No. 473
To ThERAPSIOS Caseidae
SPHENACOOONTiNfte E OAPHOSAUR DACI
LUPEOSAURIOAE
Sphenacooontidae
VARANOPSIOAE NiTOSAURIDAE
SPHENACODONTiA
EOTHYRIDIDAE
^\ /
EDAPHOSAURIA
O.PHI ACOOOHTIDAE
OPHIACODONTIA
A
Oiapstda EothyndHjae Caseidae EdaphouundBfl Ophiicodoniida^ VaianopsidBr Sphenacodontidae Therapsida

Captorhinomorpha
Figure I. Hypotheses of pelycosaur interrelationships. A) Phylogeny of pely-

cosaurs presented by Romer and Price (1940); B) Cladogram showing the inter-
relationships ol peiycosaurian lamiiies presented by Reis/ (1980). A is from Romer

and Price, 1940; B is from Reis/, 1980.
with this approach, taxa were established on the basis of phenetic

similarity. This is reflected in the diagnoses, which consist of gene-
ralized descriptions that encompass the anatomy of most of the
members of the group. The taxonomic rank assigned to a group was
based on both the diversity within the assemblage and the morpho-
logical distance between that and other groups. Ecological interpre-
tations also played a significant role in their study of pelycosaur
evolution, and a scenario uniting the morphological and ecological
interpretations was developed.
1983 PELYCOSAUR INTERRELATIONSHIPS 3
Some aspects of the phyiogeny proposed by Romer and Price

were brought into question by Langston (1965). who proposed a
close relationship between Oedaleops and Euihyris, and argued that
these genera were close to the ancestry of caseids. This implied that
caseids were not closely related to the edaphosaurids, as was sug-
gested by Romer and Price.
Recently, Reisz (1980) reviewed pelycosaur interrelationships

using a cladistic analysis of the characters available to Romer and
Price. Reisz concluded that many of the families recognized by
Romer and Price were monophyletic. but the interrelationships of
the families that Reisz proposed (Fig. IB) differed from those sug-
gested by Romer and Price. Also, the relationships that Reisz rec-
ognized did not conform to the scenario of pelycosaur evolution
proposed by Romer and Price.
During the study of a new pelycosaur from El Cobre Canyon,
New Mexico, it became clear that much additional morphological
information could be brought to bear on the problem of pelycosaur
interrelationships, and a review of the group was undertaken. The
provide a testable hypothesis of
results of this review are intended to
pelycosaur interrelationships, and to serve as a framework in which

detailed taxonomic and morphological revisions of individual gen-
era and families can be interpreted.
The central problem in a cladistic analysis is establishing the

polarity of character-states. Out-group comparison has been consid-
ered the most powerful tool for this (Watrous and Wheeler, 1981).
The interrelationships of the groups used in determining the polarity
of character-states within pelycosaurs are shown in Figure 2A.
These relationships are based on Carroll (1970), except for the posi-
tion of Diadectes, which, following Heaton (1980), is placed with
Liinnoscelis and Tseajaia in the Diadectomorpha. Also, following
Kemp ( 1980), pelycosaurs are considered to be the sister-group of all

other reptiles.
In using out-group comparison to interpret polarities, two princi-
ples are used. One is the principle of parsimony: the hypothesis of
polarities that requires the least number of evolutionary events to
account for the distribution of character-states will be accepted.
BREVIORA No. 473
PRIMITIVE OUT-GROUPS IN-GROUP SISTER-GROUP

1
1 I
Anthrocosoria Diodectomorpho Pelycosouno oil other reptiles
primitive sister- primitive sister-
out-groups Pelycosourio group out-groups Pelycosouno group
primitive Sister- primitive sister-
out-groups Pelycosouno group out-groups Pelycosouna group
primitive sister- primitive sister-
out-groups Pelycosouna group out-groups Pelycosouno group

This principle is the basis for the interpretation of polarities for the
character-states whose distribution is shown in Figure 2B E.
The second principle is the co-occurrence of primitixe character-
states. According to this principle, primitive character-states tend to
occur together. Using this principle, the character-state that occurs

in the outgroup that, on the basis of other evidence, is thought to be
more primitive is accepted as the primitive character-state. This is

the basis for the interpretation of polarities for the character-states
whose distribution is shown in Figure 2F and G. It shoud be noted
the assumption that reversals are less
that implicit in this principle is
likel\ to occur than are independent acquisitions. The use of this
principle, therefore, limits the extent to which the results presented

here can be used to test this assumption.
Characters of uncertain polarity will not be used to support rela-
tionships. However, such characters may provide corroboration of
relationships proposed on the basis of derived character-states, since
the distribution of uncertain character-states will be either consis-
tent with the cladogram or will require that the occurrence of parallel
evolution be hypothesized.
One of the problems in the study of the evolution of groups
represented by fossil material is the incorporation of data both trom
well-known animals and from animals represented by incomplete
specimens. This is especially true in the study of the evolution of
where material is rare and, in many cases, animals are
early reptiles,
represented by single fragmentary specimens. Often animals repre-
sented by such material have been assigned to a taxon erected on the
basis of well-known animals, and discussions of the evolution of the
group have focused on the interrelationships of the higher taxa.
Figure 2. Ihe interpretation ot poiantieh ol character-states by outgroup compari-

son. A) Cladogram showing the interrelationships ol the taxa used as out-groups
lor mterpreting the polarity ol character-stales within pelycosaurs; Diadectomorpha

includes /J/«(/<'<7f.v, LininoMi'll.',. and B G) Interpretation
T.sci/ala; ol polarities ol
character-states. In B E, this is based on parsmion\, in Hand G. this is based on the
principle ol co-occurrence ol primiti\e character-states. represent the deri\ed

in-group character-states; D represent the primitive in-group character-states; E repre-
sent character-states not present in the in-group.
6 BREVIORA No. 473
In order that information from both fragmentary and well-known

animals can be incorporated into the analysis of pelycosaur inter-
relationships, this study will be separated into two steps. The first
will be to construct a cladogram showing the interrelationships of
the most completely known genera using all available morphological
features. This will provide aframework in which data from the less
well known genera can be interpreted. The results of this part of the
study are presented here. The second part will be the inclusion of the
less well known animals in the analysis. Through this, the clad-
ogram will be tested and a more

detailed understanding of pelyco-
saur interrelationships will be obtained.
By using genera as the basis for considering pelycosaur interrela-
tionships, all morphological diversity known to be present within
higher taxa will be incorporated in a single analysis of diversifica-
tion of pelycosaurs. Also, assumptions about pelycosaur evolution
are minimized. Some currently recognized higher taxa are almost
certainly monophyletic and could be treated as units in a discussion

of pelycosaur interrelationships. Others, such as the Ophiacodon-
tidae, as currently defined may be either polyphyletic or paraphyletic.
By treating each genus as an independent entity, this potential

sou-rce of error is avoided.
The pelycosaurs considered hereare Casea. Varanops. Aero-
sawus, Ophiacudon, Edaphusaurus, Dimeirudon, and Sphenacodun.
The restriction of the study to these genera was necessitated by the
accessibility of material for direct observation and the fragmentary
nature of available specimens of other genera. Only characters that
were actually observed were utilized in the analysis, and many of the
reconstructions presented here are modified from previously pub-
lished reconstructions on the basis of observations made during the
course of this work. This required that a large amount of material be
examined, and supplying a complete list of specimens studied is not
possible. To facilitate future work, the specimens that showed par-
ticular structures most clearly are listed in Table 1, and the speci-
mens that are the basis for the modifications in the reconstructions
are listed in the figure legends. In all the features considered here,
SphenacodomLnd Dimelrodon ave indistinguishable (Eberth, 1981).
Thus, illustrations from only one of these genera are included here.
1983 PELYCOSAUR INTERRELATIONSHIF'S
Table I.
8 BREVIORA No. 473
ventral cheek margin. They have the distribution shown in Figure

2C, with the derived character-state present in pelycosaurs being the
presence of a concave cheek margin. The condition seen in Hylo-
iionius is a separate, derived character-state.
Figure 3. The skulls in right lateral view of A) Diaciectes: B) Casea: C) Vara-

nups; D) Ophiacodon: E) Eciaphusaurus: and F) Sphenacodon. Drawing of Dia-
decles based on 1739,MCZ MCZ
1736, and MCZ
2086; Casea based on UC 698 and
UC656; Varanops based on MCZ 1926 and UR 2423; Ophiacodon based on UC671;
Edaphosaurus based on MCZ 1762. MCZ 1764. and USNM 299844; Sphenacodon
from Eberth(1981).
/I /'/7A'('\'/a/(V;/7.v.- f, frontal; j, jugal; la, lacrymal; max, maxilla; na, nasal; pf, post-
frontal; pmx, premaxilla; po. postorbital; prf, prefrontal; pt, pterygoid; q. quad-
rate; qj, quadratojugal; sq, squamosal; st, supratemporal.
Premaxilla
In Dimetrodon, Sphenacodon (Fig. 3F), Edaphosaurus (Fig. 3E),

and Ophiacodon (Fig. 3D), the anterior margin of the premaxilla
slopes posteriorly from the anterior termination of the tooth row
giving a convex outline to the snout.
In Casea and Varanops (Fig. 3B-C), the anterior margin of the
premaxilla first extends anteriorly from the anterior termination of

the tooth row, giving a forwardly projecting rostrum. Langston and
Reisz (1981) argue that the forward sloping premaxilla seen in
Aerosaurus specimen UCMP 40096 is a result of displacement of
the element, and they reconstruct the skull with a posteriorly sloping
premaxilla. However, apart from the slope of the premaxilla, there

is no direct evidence that the premaxilla has been displaced, and no
evidence that the structure of this part of the skull was different than
in Varanops.
In Diadectes, the dorsal process of the premaxilla is vertical (Fig.
3A). Limnoscelis Varanops. Casea, and Aerosaurus in having

is like
a forwardly projecting rostrum (Romer, 1946). Anthracosaurs, with

an elongate, flattened face, are not directly comparable to either
condition seen in pelycosaurs. The structure of the premaxilla in
Hylonomus and Paleothyris is not known for certain. In Romeria,
the dorsal ramus of the premaxilla is nearly vertical (Heaton, 1979),
a condition not specifically similar to either of the character-states
present in pelycosaurs. Thus, these character-states have the distri-

bution shown in Figure 2E, with the presence of a forwardly sloping
dorsal ramus of the premaxilla being the primitive condition.
Front ah
In Dimetrodon and Sphenacodon (Fig. 4E), the frontal has a

laterally directed lappet that is greater in mediolateral width than is
the posterior end of the frontal. Edaphosaurus was figured as hav-
ing a broad flange extending laterally along the entire anterior half
of the frontal by Romer and Price (1940). However, isolated front-
als show that the lateral flange illustrated by Romer and Price is
formed in part by the prefrontal, and that a lateral lappet like that of
Dimetrodon and Sphenacodon is present (Fig. 4D).

In Ophiacodon (Fig. 4C), the frontal is without a strongly devel-
oped lateral lappet. A small projection extends between the base of

10 BREVIORA No. 473
1983 PELYCOSAUR INTERRELATIONSHIPS II
the prefrontal and postfrontal, but the width of this is much less
than the width of the posterior end of the frontal.
In Casea (Fig. 4A), Varanops (Fig. 4B), and Aerosaurus (Lang-
ston and Reisz, 1981), frontal lappets are absent.
Frontal lappets are absent in Limnoscelis (Romer, 1946). Dia-
dectes (Lewis and Vaughn. 1965). anthracosaurs (Panchen. 1970),
and Paleothyris (Carroll. 1969). Thus, these character-states have
the distribution shown in Figure 2B, with the absence of a frontal
lappet being the primitive character-state.
Maxilla
In Dimetrodon, Spenacodon (Fig. 3F), Edaphosaurus (Fig. 3E),

and Ophiacodon (Fig. 3D), the maxilla does not extend posterior to'
the orbit and does not meet the quadratojugal. In Varanops (Fig.
3C) and Aerosaurus (Langston and Reisz. 1981), the maxilla
extends posterior to the orbit and meets the quadratojugal. exclud-
ing the jugal from the ventral margin of the skull. In cross section,
the jugal slopes laterally so that a ridge is present at the contact of
the jugal and maxilla. In Casea the maxilla meets the quadratojugal,
excluding the jugal from the ventral margin of the skull (Fig. 3B). A
ridge is present along the contact of the maxilla and jugal as in
Aerosaurus and Varanops.
Thus two structural patterns are present: the Dimetrodon pattern
in which the jugal enters the ventral margin of the skull and no
ridge
ispresent at the contact of the jugal and maxilla, and the Varanops
pattern in which the jugal is excluded from the ventral margin of the
skull by a contact between the maxilla and quadratojugal and a
ridge is present along the contact of the maxilla and jugal. Diadecies
(Fig. 3A). Limnoscelis (Romer. 1946). Paleothyris (Carroll. 1969).
Figure 4. The skulls in dorsal view of A) Casea; B) Varanops: C) Ophiacodon:

D) Edaphosaurus: and E) Sphenacocion. Reconstruction of Casea based on \JC 656
and UC 698; I'aranops based on MCZ 1926 and UR 2423: Ophiacodon based on
MCZ 1366; Edaphosaurus based on MCZ 1762 and USNM 299844; and Sphena-
codon from Eberth 1981 { ),
/I /)/7rf\7a/;V)mv f, frontal; j, jugal; la, lacrymal; na, nasal; op, opisthotic; p, pari-
etal; pf, postfrontal; prf, prefrontal; so, supraoccipital; sq, squamosal; tab, tabu-
lar.
12 BREVIORA No. 473
and anthracosaurs (Panchen. 1970) are like DInietroc/on in these

features. Thus, these character-states have the distribution shown in
Figure 2B. with the absence of a contact between the maxilla and
quadratojugal and the absence of a ridge along the contact of the
maxilla and jugal being the primitive character-state.
Quadratojuiâl
and Sphenacudon (Fig. 3F), the quadratojugal is

In Dinu'troc/on
a small bone sitting on the posterolateral corner of the quadrate and
is without an anterior zygomatic process.
EUaphosaurus was reconstructed with a large quadratojugal

(Romer and Price, 1940), but specimen MCZ 1762 shows that a
small quadratojugal like that of Dimetrudun was present (Fig. 3E).
In Ophiacudon {V\g. 3D), Varanops (F\g. 3C), Aerosaurus {Lang-
ston and Reisz, 1981), and Casea (Fig. 3B), the quadratojugal
extends forward from the posterior corner of the skull forming the
ventral border of at least the posterior half of the cheek.
In Diadecies (Fig. 3 A), Lininoscelis (Romer, 1946), anthraco-
saurs (Panchen, 1970), and Paleoihyris (Carroll, 1969), the quadra-
tojugal is a large element extending well anteriorly. Thus, these
character-states have the distributionshown in Figure 2B, with the
presence of a large quadratojugal being the primitive condition.
Quadrate Ramus of Pterygoid

In Dinietrodon (F\g. 5F), Sphenacodon, and Edaphosaurus (Fig.
5E), the quadrate ramus of the pterygoid is a vertical sheet with a
rounded ventral edge. In Ophiacodon (Fig. 5D), Varanops (Fig.
5C), and Casea (Fig. 58), a shelf extends medially from the ventral
edge of the quadrate ramus, flooring the lateral most part of the
tympanic cavity. The quadrate ramus of the pterygoid in Diadectes
has a similar shelf (Fig. 5 A), as does Limnoscelis (Romer, 1946). In
anthracosaurs, a tympanic shelf is not present, but the quadrate
ramus of the pterygoid is little differentiated from the more anterior
part of the bone (Panchen, 1970), and is not comparable to the
quadrate flange of pelycosaurs in structural details. Paleothyris is
like Dinietrodon and Edaphosaurus in not having a medially
directed tympanic tlange. Thus, these character-states have the dis-

tribution shown in Figure 2G, with the presence of a medially
Figure 5. Ventral view of the posterior half of the skulls of A) Diadectes;

B) Casea; C) I'aranops; D) Ophiacodon: E) Edaphosaurus; F) Dimetrodon. Re-
construction of Diadectes based on MCZ1739 and MCZ
2042; Casea based on UC
698; Varanops based onP 12841; Ophiacodon based on MCZ 1366 and UC 1638;
Edaphosaurus based on MCZ 1762; and Dimetrodon based on MCZ 1365, MCZ
2168. and MCZ 5950.
Abbreviations: bo, basioccipital; bsp. pasisphenoid; ec, ectopterygoid; ept, epi-

pterygoid;j, jugal; op, opisthotic; pt. pterygoid; q, quadrate; sq, squamosal.
14 BREVIORA No. 473
directed tympanic cavity being the primitive

shelf tlooring the
character-state and the absence of this flange being a derived
condition.
Supraoccipiial
In Dimetrodon, Sphenacodon (Fig. 6D), Edaphosaurus (Fig.

6C), and /IfTOiai/rwi (Langston and Reisz, 1981), the supraoccipital
has an area dorsal to the posttemporal fenestra forming the dorsal
margin of the posttemporal fenestra, termed the lateral process by
Romer and Price (1940). In Ophiacodon (Fig. 6B), the supraoccipi-
tal is without an ossified lateral process. Romer and Price inter-
preted the absence of a lateral process as a result of the tendency

for
poor ossification in that genus, and reconstructed cartilaginous lat-
eral processes. Reisz (1980) accepted this hypothesis and showed a

lateral process of the supraoccipital bordering the posttemporal
fenestra in his reconstruction of the occiput of Ophiacodon. While
cartilaginous lateral processes of the supraoccipital may have been

present, these would not have been visible in posterior view since
the
tabular of Ophiacodon, as illustrated by Romer and Price (1940),
contacts the opisthotic and has a finished ventral surface that would
have formed the dorsal edge of the posttemporal fenstra. In Casea
(Fig. 6A), the arrangement of the supraoccipital and tabular is like
that of Ophiacodon: the supraoccipital is not exposed above the
posttemporal fenestra when seen in occipital view, and the tabular
contacts the paroccipital process, excluding the supraoccipital from
the margin of the posttemporal fenestra.
In Diadectes,' the elements of the occiput are fused. In Limnosce-
Hs, the relationships of the tabular are like those of the Ophiacodon
(Romer, 1946).Paleothyris is without lateral processes on the
supraoccipital, and the dorsal margin of the posttemporal fenestra is
formed by .the tabular (Carroll, 1969), as in Ophiacodon. However,
in contrast to that genus, Paleothyris has a large posttemporal
fenestra that is bounded medially by the supraoccipital, and thus is
not distinctly like either condition seen pelycosaurs. Thus, these

in
character-states have the distribution shown in Figure 2E, with the

exclusion of the supraoccipital from the border of the posttemporal
fenestra by the tabular being the primitive character-state, and the
presence of a lateral process of the supraoccipital forming the dorsal

border of the posttemporal fenestra being a derived character-state.
Bas ip terygo id Pro cesses
The basipterygoid processes of Dimeiiodon (Fig. 7G). Sphenu-

cuJon. and Edaphosaurus (Fig. 7F) extend anteriorly, ventrally,
and laterally from the lateral surface of the basisphenoid. Each
articular surface is divided into two areas, a flat
anteroventrally
facing area and a dorsally facing area at nearly 90 degrees to this.
Figure 6. Occipital view of the skulls of A) Casea:^) Ophiacodon; C) Edapho-

saurus; and D) Sphenacodon. Drawing of Casea based on UC 698 and \}C 656;
Ophiacodon based on MCZ 1366, MCZ 1426. and MCZ 1 121; Edaphosaurus based
on MCZ 1762; and Sphenacodon from Eberth (1981).
Abbreviations: bo. basioccipitai; eo, exoccipital; op, opisthotic; par. parietal;

pp. postparietal; q. quadrate; qj, quadratojugal; sq. squamosal; tab. tabular.
16 BREVIORA No. 473
In OphiacocJon, the basipterygoid processes extend ventrally (Fig.

7E). The long axis of the articular surfaces are anteroposteriorly
oriented and the surface curves smoothly from its ventral to its
lateral aspect.
In Casca (Fig. 7C). the basipterygoid processes extend laterally
from the side of the cultriform process and have strongly curved
articular surfaces. Aerusaurus (Langslon -dnd Reisz, 1981) is similar
to Casea, but differs in that the articular surfaces are more elongate
mediolaterally.
Thus, two distinct characters can be recognized in the structure of
the basipterygoid process. One of these is the curvature of the artic-
ular surfaces: the curved articular surfaces such as are seen in Aero-
saurus being one character-state and two flat articular surfaces set at
Figure 7. The basisphenoid in ventral view of A) Archeria, B) Diadectes,

C) Casea. D) I'aranops. E) Ophiacodon, F) Edaphosaurus. and G) Dimeirodon.
Drawing of Archeria based on MCZ 8736; Diadectes based on MCZ 3277; Casea
based on UC 698; I'aranops based on UR 2423; Ophiacodon based on UC 1638 and
MCZ 4820; Edaphosaurus based on MCZ 1762; and Dimeirodon based on MCZ
1697.
Ahhrevialions: h^s. basisphenoid shelf; bs w, basisphenoid wings; bst tub, basi-

pterygoid tubercula.
nearly 90 degrees to each other such as in Dimetrudon being a

second character-state. In Diadecies (Fig. 7B), Archeria (Fig. 7 A),
and Eucapturhinus (Heaton, 1979), the basipterygoid processes are
curved. Thus, for the character of the curvature of the articular
surfaces, the character-states have the distribution shown in Figure
2B, with the presence of curved articular surfaces being the primitive
character-state and the flat articular surfaces of Dimetrodun, Sphe-
nacudon, and Edaphosaurus being a derived character-state.
The second character is the orientation of the basipterygoid pro-
cesses: the laterally oriented basipterygoid processes such as is seen
in Aerosaurus being one character-state, anteroventrally oriented
is seen in Dimetrodun being a second character-

processes such as
state,and ventrally oriented basipterygoid processes with antero-
posteriorly oriented articular surfaces such as is seen in Ophiacodon
being a third character-state. Inall the outgroups, the basipterygoid
processes extend ventrolaterally from the side of the cultriform pro-

cess as in Dimetrudon. Thus, for the character of the orientation of
the basipterygoid processes, the character-states have the distribu-
tion shown in Figure 2B, with the presence of ventrolaterally
oriented basipterygoid processes being the primitive character-state.
The laterally orientated basipterygoid process seen in Casea and
Aerosaurus is one derived character-state, and the ventrally oriented
basipterygoid process with anteroposteriorly oriented articular sur-
faces ofOphiacudun is a second derived character-state.
Shelf Between Basisphenuid Wings
In Dimetrudun (Fig. 7G), Sphenacudun, and Edaphusaurus, the
basisphenoid wings extend posteriolaterally from the base of the

basipterygoid tubercula with a smooth trough between them. In
Ophiacudun. the arrangement of the basisphenoid wings is similar,
but a shelf (bs.s.. Fig. 7E) extends between the base of the wings,
roofing over the anterior end of the trough. This is also the case in
Varanups {Fig. 7D)and Aerusaurus {Langston and Reisz, 1981). In
Casea (Fig. 7C), a basisphenoid shelf is present, although its poste-
rior edge, rather than being straight, is V-shaped with the lateral
ends of the V raised.
In Limnuscehs (Romer, 1946), a shelf is present roofing the ante-
rior portion of the trough between the basisphenoid wings. In Dia-
dectes, this shelf is hypertrophied and extends to the posterior ends
18 BREVIORA No. 473
of the basisphenoid wings so that, in ventral view, the shelf appears

to occupy the position of the wings (Fig. 7B). The anthracosaur
Anheria is like Limnoscelis in having a shelf covering the anterior
portion of the trough between the basisphenoid wings (Fig. 7A).
Paleoihyris is without such a shelf.
Thus, these character-states have the distribution shown in Figure
2F, with the presence of a shelf between the bases of the basis-
phenoid wings being the primitive character-state, and the absence
of the shelf being a derived condition.
Stapes
The stapes of Dimetrodon (Fig. 8E) consists of a triangular plate

oriented anteromedially with a posteromedially oriented footplate
(fp.. Fig. 8E) directed about 90 degrees from the medial edge of the
triangular plate. The wide end of the triangle is the dorsal process
(dp.. Fig. 8E), and the narrow end is the quadrate process (qp.. Fig.
8E). The dorsal process articulates in a socket on the ventral surface
of the paroccipital process.
The stapes of Edaphosaurus (Fig. 8C) differs from that of Dime-
trodon only in proportions, the dorsal process being relatively wider
Figures. The left stapes in posterior view of A) Casea:^) Varanops:C) Ophia-

codon: D) Edaphosaurus: E) Dimetrodon. Drawing of Casea based on UC 698;
Varanops based on UR 2423: Ophiacodon based on MCZ 1366; Edaphosaurus based
on MCZ 1762; and Dimetrodon based on MCZ 1347.
Abbreviations: d\), dorsal process; fp, footplate; qp, quadrate process.

1983 PEl YCOSAIR INTERRELATIONSHIPS 19
and the quadrate process shorter. The dorsal process articulates in a

socket on the paroccipital process as in Dimetrodon.
In Ophiacodon (Fig. 8C). the stapes is more rodlike than Dime-
trodon. The dorsal process of the stapes is narrower, although it still
articulates in a socket on the paroccipital process, and the footplate

is oval, rather than circular in end view as in Dimetrodon and
Edaphosaurus.
In Casea (Fig. 8A), the stapes consists of a footplate and a single
triangular flange of bone extending laterally from this. One corner

of the flange approaches the paroccipital process and presumably
represents the base of the unossified dorsal process. The other
corner is directed towards the stapedial pit of the quadrate. A dis-
tinct socket in the paroccipital process for the dorsal process of Ihe
stapes not present.

is
stapes of Varanops (Fig. 8B) differs from that of Casea in the

The
presence of a V-shaped notch in its lateral edge. The edges of this
notch are damaged, and therefore the notch may be articicial. In
other regards, the stapes is directly comparable to that of Casea. As
in Casea, no socket is present on the paroccipital process for the
dorsal process of the stapes.
The stapes o{ Aerosaurus not known, but the paroccipital pro-
is
cess shows no articular surface for the dorsal process of the stapes
(Langston and Reisz, 1981).

The stapes of Paleothyris (Carroll, 1969) appears most similar to
the rodlike laterally directed stapes in captorhinids (Heaton, 1979),
in which the dorsal process does not articulate in a socket on the
paroccipital process. The stapes of Diadectes, as described by Olson

(1966) is not readily comparable to that of any other tetrapod. A
stapes is not known in Limnoscelis or anthracosaurs. In the early
temnospondyle Greererpeton, the stapes is similar to that of Vara-
nops and Casea in the presence of a triangular lateral process that
does not have an ossified dorsal head articulating in a socket on the
paroccipital process (Carroll, 1980). Since Greererpeton and pelyco-
saurs are both without an otic notch, probably primitively so (Lom-
bard and Bolt, 1979; Carroll, 1980), the similarity of the stapes in
these animals can be used as evidence that the stapes of Casea and
Varanops is primitive in not having a socket on the paroccipital

process that receives the dorsal process of the stapes.
20 BREVIORA No. 473
Prearticular
In Dimetrodon (Fig. 9C), Sphenacodon, and Edaphosaurus. the

prearticular underlies the pterygoideus process of the articular, a
distinctive twisting of the bone being present as it passes under the
process.
in (Fig. 9A), the prearticular underlies the medial
Ophiacodon
portion of the articular but does not show the twisting seen in
Dimetrodon. This is also the case in Varanops (Fig. 10) and Aero-
saurus (Langston and Reisz, 1981). The condition in Casea is
unknown.
ptpr
Figure 9. The right lower jaw in medial view of A) Ophiacodon: B) Edapho-

saurus; C) Dimetrodon. From Romer and Price, 1940.
Abbreviations: ang, angular; ar, articular; prear, prearticular; pt pr, pterygoi-

deus process; sur ang, surangular. Arrows indicate the position of sections shown in
Figure 1 1.
Figure 10. The lower jaw of Varanops in A) ventral view; B) medial view of
articular region; and C) section through articular and pterygoideus process. Draw-
ings based on MCZ 1926.
Abbreviations: pt pr, pterygoideus process; ret ar pr, retroarticular process.
Didaectes is without the twisting of the prearticular, as are cap-

torhinids (Heaton, 1979), and anthracosaurs (Panchen, 1970). Thus,
these character-states have the distribution shown in Figure 2B,
with the absence of a twisted prearticular being the primitive
character-state.
Angular
In Dinietrodon (Fig. HE) and Sphenacodon the angular is a
vertically oriented plate that supports the prearticular and surangu-

lar by its dorsal edge (Fig. HE), and the articular by its medial
surface. The posterior edge of the angular is notched, forming the
reflected lamina of the angular.
The angular of Edaphusawus (Fig. ID) and Ophiacodon (Fig.
1
IIC) is like that of Dinietrodon in being a vertically oriented plate.

They differ in that they are without a posterior notch.
22 BREVIORA No. 473
ang \Vy/ang
ang )/ang
fang
Figure 1 1 . Sections through the postdentary bones of the lower jaw at the posi-
tion just posterior to the dentary (marked by an arrow for the jaws shown in Figure
9). A) Varanops: B) Casea: C) Ophiacodon; D) Edaphosaurus; and E) Dime-
trodon. Drawing of Varanops based on UR 2423; Casea based on UC 698; Ophia-
codon, Dimeirodon and Edaphosaurus from Romer and Price, 1940.
Abbreviation: ang, angular.
In Casea (Fig. A), Varanops (Fig.

1 1 B), and Aerosaurus {Lang-
1 1
ston and Reisz, 1981), the angular is a troughlike bone. In Vara-

nops, a keel is present on its ventral edge, but in no position along
the angular does the bone form an extended vertical plate.
The angular in Limnoscelis and Diadectes is like that of Casea
and Varanops in being troughlike. In Paleothyris, a keel is present
in the region of the angular, but the bone does not form an extended
vertical plate. Thus, these character-states have the distribution

shown in Figure 2B, with the absence of an extended vertical plate
being the primitive character-state.
Pterygoideus Process of Articular
In Dimetrodon (Fig. 9C), Sphenacodon, and Edaphosaurus (Fig.
9C), a pterygoideus process is present as a distinct process on the

articular. In primitive species of Dimetrodon, and in Sphenacodon
and Edaphosaurus, this is located medial to the glenoid.
In Ophiacodon (Fig. 9A) a distinct pterygoideus process is not
present on the articular. A flange on the prearticular just anterior to

the articular may be functionally equivalent to the pterygoideus
process of Dimetrodon.
In Varanops (Fig. and Aerosaurus (Langston and Reisz,
10)
1981), a pterygoideus process is not present on the articular.
although a well-developed flange on the prearticular just anterior to

the articular is present. The condition in Casea is unknown.
In Diadecies, the articular does not have a distinct pterygoideus
process.The articular extends medially from the inner surface of the

jaw (Romer, 1956; Fig. 107C), but the medially projecting portion is
located beneath the medial condyle of the gleniod and is not a
distinct process. No flange is present on the prearticular anterior to
the articular. This is also the case in anthracosaurs (Panchen, 1970).
The is not known in Paleuthyris. In
internal surface of the articular
captorhinids, a pterygoideus process is absent or poorly developed,

although a medial flange formed by the prearticular just anterior to
the articularis present (Heaton, 1979). Thus, these character-states
have the distribution shown in Figure 2D, with the absence of a'
pterygoideus process being primitive for tetrapods, the presence of a
flange on the prearticular just anterior to the articular being primi-
tive for reptiles, and the presence of a pterygoideus process formed
by the articular being a derived feature within pelycosaurs.
Neural A rch
In Dimetrodon (Fig. 12F) and Sphenacodon, the neural arch has
a pit in its lateral surface above the level of the transverse process. In
Edaphosaurus boanerges (Fig. 12E), the neural arch is without pits,
although in an undescribed species of Edaphosaurus from near
Garnett, Kansas, Sphenacodon-Wkt pits are present along the length

of the column (Reisz et al. 1982). Varanops (Fig. i2C) and Aero-
saurus (Langston and Reisz, 1981) are like Dimetrodon in having a
pit in the lateral surface of the neural arch. Ophiacodon (Fig. 12D)
and Casea (Fig. 12B) are without a pit in the lateral surface of the
neural arch.
The neural arches of Limnoscelis, Diadectes, anthracosaurs, and
Paleothyris are without pits.Thus, these character-states have the
distribution shown in Figure 2B, with the absence of pits in the
neural arch being the primitive character-state.
Transverse Processes
In Dimetrodon (Fig. 12F) and Sphenacodon, the longest trans-

verse processes are in the area of the posterior cervicals and anterior
dorsals. They extend laterally a distance about equal to the width of
the centrum.
24 BREVIORA No. 473
Figure 12. Anterior dorsal vertebrae in lateral and anterior view, and a cross
section of an anterior dorsal centrum of A) Lahidosaurus: B) Casea; C)
Varanops:
D) OphiacoJon: E) Edaphosaurus; and F) Dimeirodon. Drawing of Lahidosaurus
based on MCZ 8724; Casea based on P 12841; Varanops based on MCZ 1926;
Ophiacodon based on MCA 5912; Edaphosaurus based on MCZ 1359; and Dime-
trodon based on MCZ 5216.
Ahhreviaiions: pit, pit in the lateral surface of the neural arch; vent flange, ventral
flange supporting the transverse process.

In Edaphosaurus, the longest transverse processes are shorter than

the width of the centrum and the transverse processes are braced by
a vertical flange of bone ventrally.
Ophiacodon (Fig. 12D) has short transverse processes directed
more ventrally than in Edaphosaurus and Dimetrodon. The ventral
flange forms a complete web of bone connecting the transverse
proces to the anterior edge of the centrum.
Varanops (Fig. 12C), Aerosaurus (Langston and Reisz, 1981),
and Casea (Fig. 12B) have long laterally directed transverse pro-
cesses. Limnoscelis, Diadectes, anthracosaurs, and Paleothyris all
have short transverse processes, although only in captorhinids (Fig.
12 A) are these supported by a web of bone that extends to the
anterior edge of the centrum. Thus, for the character of the trans-
verse processes, the character-states have the distribution shown in
Figure 2D, with the presence of short transverse processes con-

nected to the anterior edge of the centrum by a web of bone being
the primitive character-state for reptiles, and the long transverse
processes being a derived character-state present within pelycosaurs.
Centrum
In Dimetrodon (Fig. 12F) and Sphenacodon, the vertebrae in the

cervical and anterior to mid-dorsal regions of the vertebral column
have well-developed keels.
In Edaphosaurus (Fig. 12E), keels are present only on the cervical
vertebrae. The dorsal vertebrae have broadly rounded ventral
surfaces.
In Ophiacodon (Fig. 12D), the cervical vertebrae are keeled. The
mid-dorsal vertebrae are wedge-shaped in cross section but have a
pair of ridges ventrally, rather than a single keel. The posterior
dorsal and lumbar vertebrae are circular in end view.
In Varanops (Fig. 12C), the cervical and mid-dorsal vertebrae are
keeled.
In Casea the most anterior cervical vertebrae are unknown. All
the more posterior vertebrae are without keels (Fig. 12B). In cross
section, the centra have flatter ventral surfaces than in Edapho-
saurus.
The centra of Diadectes and Limnoscelis are not specifically sim-
ilar to any pelycosaur. In cross section, they are generally wedge-
shaped, but are without a distinct keel. The vertebrae of Proiero-

26 BREVIORA No. 473
2°ridge
Figure 13. The humerus in distal ventral view of A) Casea: B) Aerosaurus:

C) Varanops;D) Ophiacodon; E) Edaphosaurus; and F) Dimetrodon. Drav/'ing of
Casea from WiUiston (1911), drawing of Aerosaurus based on UC 464; Varanops
based on UR 695; Ophiacodon based on MCZ I486; Edaphosaurus based on MCZ
3417; Dimeirodun based on MCZ 1304.
Abbreviations: delt. deltopectoral crest; ect, ectepicondyle; ect f, ectepicondyl-
car foramen; ent f, entepicondylar foramen; pect fossa, pectoralis fossa; sup, supi-
nator process; 2° ridge, secondary pectoralis ridge.
gyrinus an anthracosaur with a protoreptilian vertebral pattern

(Holmes and Carroll, 1977), are without keels. This is also the case
in captorhinids (Fig. I2A). Thus, the absence of a distinct keel can
be considered a primitive feature.
Dehopectoral Crest
Sphenacodon and Dimetrodon (Fig. I3F), the dehopectoral

In
bulbous distal end and a sharp knife-edge base. In Ophia-
crest has a
codon and Edaphosaurus. the distal end of the dehopectoral crest is
bulbous, but the proximal end is a broad ridge (Fig. 13 D-E).
In Varanops (Fig. 13C), Aerosaiirus (Fig. 13B). and Casea (Fig.
13 A), a secondary ridge is present extending from the distal end of
the dehopectoral crest to amore medial position on the proximal
end of the humerus. In Casea and Aerosaurus, this is a sharp crest
that results in the presence of a distinct fossa at the base of the
dehopectoral crest. In Varanops this is a low, rounded ridge.
In captorhinids (Holmes, 1977), the crest is absent. The tuberosity

is located on the lateral edge of the proximal end of the humerus. In
Diadectes, the dehopectoral crest is like that of Ophiacodon,
although it forms a more obtuse angle with the proximal end of the
humerus. Thus, these character-states have the distribution shown
in Figure 2E, with the presence of a dehopectoral crest formed by a
single broad ridge extending from the tuberosity to the proximal

end of the bone being a primitive reptilian condition, and the pres-
ence of a secondary ridge extending from the dehopectoral tuber-
osity to the proximal end of the bone being a derived condition.
Results
The distribution of the character-states for which polarity can be
interpreted, and the cladogram that requires the fewest number of

reverseals or convergent evolutionary events to explain this distribu-
tion, is shown in Figure 14.
CHARACTER ANALYSIS: II. FEATURES FOR WHICH

POLARITY CANNOT BE INTERPRETED
As argued in the materials and methods section, characters for
which polarity of the character-states cannot be interpreted are of
use in providing a test of the caldogram presented on the basis of
other characters. Three such characters are considered here.
28 BREVIORA No. 473
Casea Varanops Aerosaurus Ophiacodon Edaphosaurus Dimetrodon Sphenocodon
Figure 14. Hypothesis of relationships of selected genera of pelycosaurs based on

the characters described in the te.xt for which polarity can be interpreted. D indicate
primitive character-states, and ED indicate derived character-states. For description

of characters and character-states see Table 2.
Table 2. The characters that form the basis for the cladogram shown in Figure
14. The numbers refer to the characters shown in Figure 14.
1) humerus, deltopectoral crest:
primitive character-state: the presence of a single ridge leading from the distal end of
the deltopectoral crest to the proximal end of the humerus;
derived character-state: the presence of a secondary ridge leading from the distal end
of the deltopectoral crest to a more medial position on the proximal end of the
humerus with a fossa at the base of the crest.
2) maxilla:
primitive character-state: no contact between the maxilla and the quadratojugal;

derived character-state: the maxilla contacts the quadratojugal with the bones raised
to form a ridge along the contact of the maxilla and jugal.
3) basipterygoid process:
primitive character-state: basipterygoid process directed anteroventrally;

derived character-state #1 (indicated by in Figure 14): basipterygoid process
directed laterally and are mediolaterally elongate:

derived character-state #2 (indicated by ID) in Figure 14): basipterygoid processes
directed ventrally and articular surface oriented anteroposteriorly.
4) cheek margin:
primitive character-state: cheek margin convex;
derived character-state: cheek margin concave.
5) premaxilla:
primitive character-state: anterior margin of the premaxilla extends anteriorly from
the anterior termination of the tooth row;
derived character-state: anterior margin of the premaxilla slopes posteriorly from the
anterior termination of the tooth row.
6) stapes:
primitive character-state: dorsal process of stapes not articulating in socket on the

paroccipital process;
derived character-state: dorsal process of stapes articulating in a socket on the paroc-
cipital process.
7) angular:
primitive character-state: angular without an extended ventral plate in the region

anterior to the articular;
derived character-state: angular with an extended ventral plate in the region of the
articular.
8) basipterygoid articular surfaces:

primitive character-state: articular surfaces curved;
derived character-state: articular surfaces differentiated into two tlal areas at right
angles to each other.
9) shelf between basisphenoid wings:

primitive character-slate: shelf between basisphenoid wings present;
derived character-state: no shelf between basisphenoid wings.
30 BREVIORA No. 473
Table 2. Continued
10) frontal:
primitive character-state: frontal without a lateral lappet;

derived character-state: frontal with a lateral lappet.
1 1
) quadratojugal:
primitive character-state: quadratojugal extends anteriorly forming
ventral margin ot
skull along posterior half of cheek;

derived character-state: quadratojugal restricted to ventro-lateral corner of skull.
12) prearticular;
primitive character-state: prearticular not twisted;
derived character-state: prearticular twisted.
13) pterygoideus process:
primitive character-state: pterygoideus process not formed by articular alone;

derived character-state: pterygoideus process formed entirely by articular.
14) quadrate ramus of pterygoid:
primitive character-state: quadrate ramus of pterygoid with a medially directed tym-

panic flange along its ventral edge;
derived character-state; quadrate ramus of pterigoid with rounded ventral edge.
15) supraoccipital:
primitive character-state: supraoccipital without a lateral process, dorsal border

of
posttemporal fenestra formed by tabular;

derived character-state: supraoccipital with a lateral process that forms the dorsal
border of the posttemporal fenestra.
16) neural arch:
primitive character-state: neural arch without pits;

derived character-state: neural arch with pits on its lateral surface at the level of the
zygapophyses.
17) transverse processes:
primitive character-state: transverse processes supported by a web of bone extending

to the anterior edge of the centrum;
derived character-state: transverse processes extend laterally, ventral flange not
extending to the anterior edge of the centrum.
Postparietal
In Dimetrodon, Sphenacodon (Fig. 6D), and Edaphosaurus (Fig.

6C), the postparietal is a single median element that broadly overlies
the supraoccipital.
The Ophiacodon, a postparietal was not observed, but the
supraoccipital shows that the postparietal did not greatly overlap
that bone.
In Casea (Fig. 6A), the postparietal

is a paired element.
The postparietal of Varanops and Aerosaurus is unknown.
Limnoscelis is like Dimeirodon and Edaphosaurus in having a

single postparietal (Romer.
1946). In anthracosaurs, as in primitive
tetrapods generally, the postparietal is a paired element (Panchen,
1970).
Two explanations for this distribution of character-states are
equally possible: that Liinnuscelis apomorphic in having a single
is
postparietal and a paired postparietal is primitive for reptiles, or

that a single postparietal is primitive for reptiles and Casea is
derived in having a paired postparietal. In view of this, the presence
of single or paired postparietals must be considered character-states
for which polarity cannot be determined.
Paroccipiial processes
The paroccipital processes in Dimetrudun and Sphenacodon are

rodlike structures sloping ventrally and posteriorly. In cross section,
they are triangular with their height less than twice their width. In
Edaphosaurus. the processes are like those in Dimetrodon, but they
and extend more directly laterally (Fig. 6C). In Ophia-
are shorter
codon, the process short and laterally directed (Fig. 6B). It is
is
rodlike, although end is unossfied so that, as preserved, it

its distal
does not reach the cheek. In Varanops (Fig. 4B) and Aerosaurus
(Langston and Reisz, 1981), the paroccipital process is platelike: in
lateral view its height is more than three times its width. In Casea,
the platelike nature of the paroccipital process is exaggerated (Fig.
6A).
Thus, two distinct character-states can be recognized: one present
in Varanops and Casea in which the paroccipital process is platelike,
and one seen in Ophiacodon, Edaphosaurus. Dimetrodon. and
Sphenacodon which the paroccipital process is rodlike. The
in
paroccipital process of Limnoscehs. as described by Romer 1946), (
is not directly comparable to either pelycosaur condition. The
paroccipital process of Paleothyris is unossified (Carroll, 1969).

Thus, there is no basis for interpreting the polarity of these
character-states.
Supinator Process
Dimetrodon and Sphenacodon, the supinator process

In is elon-
gate proximodistally, and its distal end curves distally (Fig. 13F).
32 BREVIORA No. 473
In Edaphosaurus (Fig. I3E), an ectepicondylar foramen is pres-

ent. Romer and Price ( 1940) interpreted this as a result of the devel-
opment of a bony connection between the ectepicondyle and the
distalend of the supinator process. Without this connection, the
supinator process would have the morphology of that of Dime-
trodon.
In Ophiacodon(Fig. 13D), the supinator process is a narrow
triangular process that projects laterally.
Casea(¥\g. 13A)and Varanops(¥'\g. 13C) are like D//??e//'of/on in
the development of their supinator process. The morphology of the
process in an adult humerus of Aerosaurus is not known.
The structure of the supinator process is variable in early tetra-
pods. In anthracosaurs, this is represented by a crest running the full
length of the humerus (Panchen, 1970). In Limnoscelis and Dia-
dectes, it is a narrow laterally directed process located distal to the
radial condyle (Romer. 1956). In its shape, it is like that of Ophia-
codon, but in Ophiacodon. as in other pelycosaurs and in contrast
to Diadectes and Limnoscehs, the supinator process is located dis-
tally and is separated from the radial condyle by a distinct groove.

The supinator process of Paleothyris is located near the distal end of
the humerus and is separated from the radial condyle by a groove
(Carroll, 1969). Its shape does not compare directly with either
pelycosaur condition. This distribution of character-states does not
allow polarity of the character in pelycosaurs to be interpreted.
Resuhs
The characters for which polarity cannot be interpreted can be
separated two groups: those whose distribution does not

into
require hypothesizing the occurrence of parallel evolution, and

those whose distribution requires a more complicated hypothesis of
evolution of the character. In the first group are the structure of the
paroccipital process and the presence of paired postparietals. The
structure of the paroccipital process is in accordance with the sepa-
ration of Ophiacodon, Edaphosaurus, Dimetrodon, and Sphena-
codon from Aerosaurus, Varanops, and Casea, and thus is in
agreement with the separation of pelycosaurs into these two groups.
In the second group is the structure of the supinator
process. If
the supinator process of Ophiacodon is autapomorphic, these
character-states are consistent with the cladogram; if primitive, then
the proximodistally elongate supinator process must have evolved
independently in the clade including Casea and Varanops and the

clade including Edaphosaurus, Dimetrodon, and Sphenacodon.
Thus, this character is not necessarily in conflict with the clado-
gram, but requires a more restrictive hypothesis of polarities to

account for the distribution of the character-states.
DISCUSSION
These from those of Reisz ( 980) in three features: )
results differ 1 1
Edaphosaurus and Dimetrodon are interpreted as members of a

clade more derived than Ophiacodon; 2) Varanops and Aerosaurus
are interpreted as being more primitive than the clade including
Ophiacodon, Edaphosaurus, Dimetrodon, and Sphenacodon; and
3) Casea, Varanops, and Aerosaurus are interpreted as being
members of a single clade distinct from the clade including Ophia-
codon, Edaphosaurus, and Dimetrodon.
The more primitive position of Ophiacodon relative to Edapho-
saurus and Dimetrodon is the same as that in the phylogeny pro-
posed by Romer and Price (1940). The alternate interpretation of

relationships of these genera proposed by Reisz (1980) was based
primarily on cranial features, including the structure of the frontal,
supratemporal, and the proportions of the skull. The structure of
the frontal of Edaphosaurus was incorrectly interpreted by Romer

and Price ( 940). As discussed above, the frontal of Edaphosaurus is
1
directly comparable to that of Dimetrodon, and the frontals in these

animals share features that can be interpreted as being derived with
respect to Ophiacodon. Thus, the frontal supports a relationship
between Edaphosaurus and Dimetrodon, rather than between Ophi-
acodon and Dimetrodon. The structure of the supratemporal is
poorly known in most genera. The element that Romer and Price
identified as thesupratemporal in Casea is better interpreted as the
proximal end of the squamosal. This is also the case in Edaphosau-
rus. Thus, this character cannot be used at present in interpreting
pelycosaur interrelationships. The proportions of the skull of Eda-

phosaurus are not known with sufficient certainty to be used in
interpreting relationships.
The position of Varanops and Aerosaurus is the most striking
difference from both the cladogram of Reisz (1980) and the phyl-
ogeny of Romer and Price (1940). This difference is based on the
34 BREVIORA No. 473
interpretation that Ophiacodon and Dimetrodon share derived

cranial features not present in Varanops and Aerosaurus, and that
Casea, Varanops, and Aerosaurus share derived cranial features not
present in Ophiacodon. Necessarily, we interpret the derived verte-
bral features shared in Dimetrodon and Varanops, the features that
Romer and Price used to unite these genera, as convergent.
The suggestion that Casea, Varanops, and Aerosaurus are mem-
bers of a single clade is the least well supported of the relationships
proposed above. This reflects the problems inherent in determining

relationships of primitive members of clades. Greater certainty
about the primitive character-state of various features in pelyco-
saurs will come from a better understanding of the interrelation-
ships of early reptiles and the use of an out-group that is more

closely related to pelycosaurs than are diadectomorphs in the
analysis of polarities. This will allow better separation of characters
that are primitive for pelycosaurs from those that are derived for the
clade including Casea and Varanops.
ACKNOWLEDGMENTS
The authors are greatly indebted to the scientific and curatorial
staff of the Field Museum of Natural History and the American
Museum of Natural History for permission to study specimens in
their collections. Grants from the K. P. Schmitt Fund of the Field
Museum enabled the authors to visit the Field Museum. Versions of
thismanuscript were read by J. Hopson, H. Barghusen, R. Reisz,
M. Heaton, F. Jenkins, Jr., H. Sues, and G. Meyer. Their critical
comments substantially improved the manuscript.
Publication costs of this study were covered in part by a grant
from the Wetmore CoUes Fund.
LITERATURE CITED
Carroll, R. L. 1964. The earliest reptiles. J. Linn. Soc. (Zool.), 45: 61-83.
1969. A Middle Pennsylvanian captorhinomorph and the interrelation-

ships of primitive reptiles. J. Paleontol., 43: 151-170.
1970. The ancestry of reptiles. Phil. Trans. Roy. Soc. Lond. B, 257:
267-308.
1980. The hyomandibular as a supporting element in the skull of
primitive tetrapods pp. 293-317. In A. L. Panchen (ed.). The Terrestrial

Environment and the Origin of Land Vertebrates, London and New York,
Academic Press.
Eberth, D. 1981. The skuU of Sphenacodonferocior and comparisons within the
subfamily Sphenacodontinae. MSc thesis. Department of Paleontology, Berke-
ley, University of California.
Heaton, M. H. 1979. Cranial anatomy of primitive captorhinid reptiles from the
Late Pennsylvanian and Early Permian, Oklahoma and Texas. Oklahoma
Geol. Surv. Bull. No. 127, 84 pp.
1980. The Cotylosauria: A reconsideration of a group of archaic tetra-
pods, pp. 479-551. /n A. L. Panchen (ed.). The Terrestrial Environment and the
Origin of Land Vertebrates, London and New York, Academic Press.
Holmes, R. 1977. The osteology and musculature of the pectoral limb of small
captorhinids. J. Morphoi., 152: 101-140.
Holmes, R., and R. L. Carroll. 1977. A temnospondyl amphibian from the
Mississippian of Scotland. Bull. Mus. Comp. Zool., 147: 489-511.
Kemp, T. S. 1980. Origin of mammal-like reptiles. Nature, 283: 378-380.
Langston, W. 1965. Oedaleops camp/ (Reptilia: Pelycosauria). A new genus and
species from the Lower Permian of New Mexico and the family Eothyrididae.
Bull. Texas Mem. Mus., 9: 1^6.
Langston, W., and R. Reisz. 1981. Aerosaurus wellesi. new species, a varanop-
seid mammal-like reptile (Synapsida: Pelycosauria) from the Lower Permian of
New Mexico. J. Vertebr. Paleontol., 1: 73-96.
Lewis, G. E., and P. P. Vaughan.
Early Permian vertebrates from the Cutler
1965.
Formation of the Placerville area, Colorado. Contrib. Paleontol. Geol. Surv.
Prof. Pap. No. 503-C, pp. 1^6.
Lombard, R. E., and J. R. Bolt. 1979. Evolution of the tetrapod ear: an analysis
and reinterpretation. Biol. J. Linn. Soc, 11: 19-76.
Olson, E. C. 1966. —
The middle ear morphological types in amphibians and rep-
tiles. Am. Zool., 6: 399^19.
Panchen, A. L. 1970. Batrachosauria, part A. Anthracosauria. In O. Kuhn (ed.),

Handbuch der Palaoherpetologie, Stuttgart, Germany and Portland, USA,
Gustav Fischer Verlag, 84 pp.
Reisz, R. 1980. The Pelycosauria: A review of phylogenetic relationships, pp.
553-592. In A. L. Panchen (ed.). The Terrestrial Environment and the Origin of
Land Vertebrates, London and New York, Academic Press.
Reisz, R.,M. J. Heaton, and B. R. Pynn. 1982. Vertebrate fauna of the Late
Pennsylvanian Rock Lake Shale near Garnett, Kansas: Pelycosauria. J. Paleon-
tol., 56: 741-750.
Romer, a. S. 1946. The primitive reptile Limnoscelis restudied. Am. J. Sci., 244:
151-188.
1956. The Osteology of the Reptiles. Chicago and London, University of
Romer, A. S., and L. I. Price. 1940. Review of the Pelycosauria. Geol. Soc. Am.
spec. Pap. No. 28, 538 pp.
Watrous, L. E., and Q. D. Wheeler. 1981. The out-group comparison method
of character analysis. Syst. Zool., 30: 1-1 1.
Williston, S. W. 1911. American Permian Vertebrates. Chicago, University of

—S. COMP. ZOOL-
MAf? J 8 1985
B K -i
^ ro ff A
Mseum of Comparative Zoology
us ISSN 0006-9698
RVTHIROMIA ELCOBRIENSIS, A NEW PELYCOSAUR

FROM EL COBRE CANYON, NEW MEXICO
David A. Eberth' and Donald Brinkman^
Abstract. Riithiioniia elcuhrien.si.s. a new genus and species of pelycosaur from

thePermo-Carbonilerous redbeds of El Cobre Canyon. New Mexico, is based upon
an articulated portion ot the vertebral column, partial appendicular skeleton, and
associated vertebral and cranial fragments. Ruthirumia is tentatively included within
the Varanopseidae on the basis of postcranial characters, it is distinguished from all
other varanopseids by its larger size, pinched-in lumbar centra, and massively
developed delto-pectoral crest.
INTRODUCTION
The Permo-Carboniferous, Cutler formation redbeds of El Cobre
Canyon, New Mexico, have yielded a unique assemblage of fossil
tetrapods. El Cobre Canyon, near Abiquiu, New Mexico, is a large
box canyon that has been sporadically prospected by collectors and
paleontologists since the late 1870's. The fossil fauna and flora,
known from both the canyon walls and floor, was most recently
reviewed by Fracasso (1980).
In 1965 A. Lewis and S. Olsen collected the larger part of the
postcranial skeleton and disarticulated cranial fragments of a
'Department of Geologs. Universit\ of I oronto. Toronto, Ontario, Canada, M5T

IR7.
-Museum of Comparati\c Zoolog\, Harvard University, Cambridge, Massachusetts
02138. Present Address: lyrrellMuseum of Paleontology. Bo.\ 7500. Drumheller.
Alberta. Canada. lOJ OYO.
2 BREVIORA No. 474
pelycosaur from the west wall of the canyon. The specimen, referred
to Ophiacodon navajovicus, remained largely unprepared in the
collections at the Museum of Comparative Zoology at Harvard
University until the spring of 1980 when the authors undertook a

study of the specimen (MCZ 3150). Preparation and study of the
material has shown that while it is clearly a pelycosaur, it is
unassignable to Ophiacodon. It is here described as a new genus and
species, a primitive member of the family Varanopseidae.
SYSTEMATIC PALEONTOLOGY
Class: Reptilia
Order: Pelycosauria
Family: Varanopseidae
Genus: Ruthirumia gen. nov.
Type species: Ruthirumia elcuhriensis gen. et sp. nov.
Etymology: Named in honor of Ruth Romer.

Diagnosis: A large varanopseid distinguished from all other
members of the family in the following features: linear measure-
ments approximately 30 percent greater than those of Varanodon.
previously the largest known varanopseid; the first eight centra
directly anterior to the pelvis deeply pinched-in laterally, just below
the level of the notochordal pit, giving the centra an hour-glass
appearance when seen in cross section; each of the eight centra

displays a broad rounded ventral surface; the delto-pectoral crest of
the humerus is massively developed and continues distally as a sharp
ridge below the area of pectoralis muscle insertion; the neck of the
astragulus represents 35 percent of the total height of that element.
Ruthiromia elcohriensis differs from Varanops brevirostris in
having a keeled angular in the region of the articular; a large medial
process of the articular positioned farther forward than in the latter

genus; and a low bladed ilium. Ruthiromia elcohriensis differs from
Aerosaurus greenleeorum in not exhibiting the excavated anterior
lip in its anterior dorsal vertebrae. Ruthiromia elcohriensis differs

from Aerosaurus wellesii in retaining a short paroccipital process
and two sacral ribs.
1983 A NEW PELYCOSAUR 3
Ruthiroinia elcohriensis gen. et sp. nov.

(Figs. 2-8.9-5)
Etymology: Named in reference to El Cobre Canyon, the type
locality.
Diagnosis: Same as for the genus.
Holoiype (MCZ 3150): An articulated, partial \ertebral column.

peKis and right hind limb found in close association with disarticu-
lated elements. All material is considered to represent a single indi-
vidual. Material includes: an occipital fragment including supra-
occipital and opisthotic; a left quadrate fragment in articulation

with a partial quadratojugal and squamosal; the posterior portion
of the left mandible, the atlas centrum and axis intercentrum; a
cervical rib; four disarticulated dorsal vertebrae; one disarticulated
neural arch; one partial neural spine; articulated posterior axial
elements including the last seven presacral vertebrae, two sacral
vertebrae and five caudal vertebrae; right and left partial scapulaco-
racoids; a complete right pelvic girdle; a partial left pelvic girdle
including pubis, ischium and a partial ilium; a right humerus; a
right femur; a left proximal femur; a right tibia, fibula, tarsus, and
pes.
Type Locality and Age: Cutler Formation. Lower Permian. Col-
lectedfrom the fluvial redbeds in the west wall of El Cobre Can\on.
Specific collecting site unknown.
METHODS
Although comparisons of Ruthiromia elcobriensis with other
pelycosaur taxa have been performed in line with phylogenetic sys-
tematic principles, a few words on polarity determinations within
the pelycosaurs are in order. Following Brinkman and Eberth
(1983), we accept a very primitive position for the varanopseids
within the paraphyletic taxon Pelycosauria (Fig. 16). Thus any pel\-
cosaur taxon may appear closely related to the varanopseids.
caseids. and eothyridids simply by exhibiting primitive characters.
To guard against the use of symplesiomorphy in establishing the
affinities of Ruthiromia elcobriensis, we have evaluated character
state polarities within the context of what we consider to be a gener-
ally accepted view of amphibian reptile relationships. In this paper

4 BREVIORA No. 474
\\c adopt the classification ot Carroll (1969) moditied b\ Heaton

(1980) (Fig. I). It is b\ ictereiice to the Diadectomorpha (including
Tscajaia. Lininosie/i.s. and Ditulccics) as the closest sister ta.xon of
reptiles and the rclati\e distributit)n oi character states in reptiles

that v\e determine character polarit\ in primitixe pelycosaurs and
specilicalU Rmhlroniia clcohiicusis.
DESCRIPTION AND COMPARISON

Skull: A small fragment of the left side of the occiput is preserved
(Fig. 2A,B). The anteroposteriorK flattened paroccipital process
tall
extends directl\' laterall\ beneath the posttemporal fenestra and

terminates in a con\e.\, thin edge as in \
aranops and Aerosaurus
wcllcsii. No trace ol a suture between the opisthotic and supraoccip-
ital is posterior \iev\. unfinished lateral edges of the
\isible. In
supraoccipital suggest that the absence of lateral processes is a result

of incomplete preservation. he anterior surface of the occipital
1
fragment displays a small gap between the anterior surface of the

supraoccipital and an anteromedial continuation of the opisthotic.
No notch for the trigeminal is visible however, and it appears that
only the corner of the braincase in this region is perserved.
As pelycosaurs and primitive reptiles generallv the quadrate is
in .
quite tall (Fig. 3A,B.C). Both cond\les are oriented with their long
a.xes anteromediallv. the medial condyle being the more elongate. A
saddle-shaped concavity occurs between the two condyles. Frag-

mentarv bone lateral to the posterior edge of the element probably
represents the squamosal and a portion of the quadratojugal. Distal
extremities are not preserved. No stapedial recess is discernible on
the medial surface of the quadrate.
Ihe posterior portion of the left mandible (Fig. 4 A,B,C), includes
and possibly prearticu-
the articular, part of the surangular, angular
lar and measures cm
long. Ihe condylar recesses of the articular
7.7
are in the horizontal plane. The articular has a large triangular,
mediall\ projecting process whose medial apex is in line with or
slightlv ahead of the posterior margin of the Meckelian trough and
well ahead of the condylar recesses. This is closely comparable to
the condition seen in Aerosaurus wellesii. In Varanops the medial
apex of the process is located just slightlv anterior to the cond\lar
recesses. In the area of the articular, the angular is keeled and com-
pares quite well with A. wellesii and I aranoJon. I

aranops differs in
1983 A NEW PELYCOSAUR
PROTOROTHYRIDIDA£
ANTHRACOSAURIA DIADECTOMORPHA SYNAPSIDA ALL OTHEr'rEPTILES
Figure I. Cladogram depicting relationships of taxa closely involved in the

amphibian, reptilian transition. Derived from Carroll (1969) and modified by Heaton
(1980).
Figure 2. R. elcobriensis, MCZ 3 1 50. A. Posterior view of left opisthotic/ supraoc-

cipital fragment. B. Anterior view of the same. Scale equals I cm.
Abbreviations: op, opisthotic; st, supratemporal.
having an angular that is gently rounded in this portion of the jaw.

The posterior limit of a medial fenestra is preserved at the anterior
most portion of the mandibular fragment and appears to be

bounded by the angular and the prearticular.
Axial Skeleton: The vertebral column is known from an articu-
lated series of 14 vertebrae and five disarticulated vertebrae. All
regions are represented, but only the atlas is present from the cervi-
cal region. The centra are clearly larger than their counterparts in
other varanopseid taxa (Table ) and appear to be relatively shorter
1
than those of Varanops and A. greenleeorum (Table 2).

BREVIORA No. 474
Figure 3. R. elcohriensis, MCZ

3150. A. Anterior view of lett quadrate, squamo-
sal,quadratojugal fragment. B. Medial view of the same. C. Ventral view of the same.
Scale equals 1 cm.
Abbreviations: q, quadrate; sq, squamosal.

1983 A NEW PELYCOSAl R
c.r.
Figure 4. R. elcohriensis, MCZ3150. A. Dorsal view of left mandibular fragment.

B. Medial view of the same. C. Lateral view of the same. Scale equals I cm.
Abbreviations: a, angular; ar, articular; cr. condylar recesses of articular; mp,

medial process of the articular; mt, Meckelian trough; sa, surangular.
BREVIORA No. 474
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12 BREVIORA No. 474
Figure 5. R. eUohriemis. MCZ 3150. A. Anterior same and right cervical rib.
Scale equals 1 cm.
Abbreviations: ai, a.xis intercentrum.
A small block of matri.x contains the atlas centrum and the axis
intercentrum as well as a spatulate cer\ical rib Fig. 5A,B). The atlas
(
centrum is fully ossified and is exposed \entrally. The axis intercen-

trum is a large crescentic element fused to the posterior surface of
the atlas centrum. Although the atlas centrum is visible ventrally. it
is the axis intercentrum that more prominent, contributing to
is
more than 50 percent The condition in

of the ventral surface.
Varanops is like that in Ruthironua. The condition in aranudun, \
A. i^ri'enleeoruni, and A. wellesii is unknown.

The head of the cer\ ical rib exhibits separate capitular and tuber-
cular processes. The shaft is expanded anteroposteriorly as a spatu-
late convex blade.
A disarticulated anterior dorsal vertebra includes the centrum
and and displays an anterolateralK' projecting
partial neural arch
diapophysis that extends 6.8 mm from the unexpanded neural arch
(Fig. 6). The ventral surface of the centrum is sharply keeled. This is
the typical morphology of all varanopseids with the exception o{ A.
greenleeuruin. There, the anterior
lip of the centrum
is deeply
recessed to receive the intercentrum and the ventral surfaces are

never as strikingly keeled. The floor of the neural canal is not ossi-
fied,resulting in a deep excavation which extends nearly to the
mid-height of the centrum as in Varanops and A. greenleeoruni. The
articulating surface of the zygapophysis is slightly inturned and
forms an angle of 20 degrees to the horizontal.
The three remaining disarticulated dorsal vertebrae (Fig. 7) can
be arranged relative to one another but position in the column is
Figure 6. R. ekobriensis, MCZ 3150. Left lateral view of anterior dorsal vertebra
and a cross section through the mid-length. Scale equals 1 cm.
uncertain. All three display a small pit on the lateral surface of the
centrum wall at mid-height. The more anterior of the three is keeled,
although not as sharpl\ as the anterior dorsal vertebra just de-

scribed. The ventral surface of the centrum displays three closely set
ridges extending anteroposteriorly. in P'a/a/76)/;i comparable ridges
occur on the 22nd presacral vertebra (Fig. 9).
The second vertebra retains a lateralis projecting diapophysis that
extends 7 mm
from the neural arch. The ventral surface is gently
rounded, however, and shows no trace of a keel or ridges.
The third vertebra differs from the preceding two in that the
centrum is more broadly rounded.
ventral surface of the
Neural arch excavations are present on the third vertebra and
extend anteroventrally and open onto the surface of the neural arch
between the diapophyses and the anterior zygapophyses. The dia-
pophysis projects posterolaterally and measures 5 mm from the
surface of the neural arch. The
articulating surface of the zyga-
pophysis appears nearh horizontal. The orientation of the diapo-
physis and the smoothly rounded \entral edge of the centrum
suggest that this vertebra was a lumbar.
Although the last seven presacral vertebrae are in articulation
(Fig. 8). onh' the first four of the series are preserved well enough to
merit discussion. As
the preceding vertebrae, each displays a
in
small pit in the lateral wall of the centrum at mid-height. The lateral
14 BREVIORA No. 474
Figure 7. /?. elcohriensis, MCZ 3150. Left lateral views and cross sections of three
dorsal centra arranged with the anterior most to the left. Scale equals 1 cm.
walls of the centra are strongly concave and the ventral surfaces are
broadly rounded giving the centra an hour-glass shape in cross sec-
tion. This condition is termed pinched-in. No other varanopseid
displays this condition in the posterior dorsal centra. Varanops dis-

plays broadly rounded ventral surfaces on its last four presacral
vertebrae (Fig. 9), but there is no indication of pinching-in or pitting
of the lateral walls, of the centra. A posterior dorsal vertebra of an
immature A. wellesii (UCMP 20096?) displays a pit in the lateral
wall of the centrum, but lacks the pinched-in condition.
The neural arches of the last three presacral vertebrae are deeply
excavated at the level of the articular surfaces of the zygapophyses.
All neural spines in this articulated series are imperfectly pre-
served and show only the primitive condition of being laterally
compressed. The centra of all of the vertebrae posterior to the third
vertebra of the series are either inaccessible or poorly preserved.
A disarticulated neural arch with a complete neural spine is
poorly preserved but shows that the spine is relatively lower in

height than those of Varanops and A. wellesii (Table 2).
Figure 8. R. elcubriensis. MCZ 3150. Right lateral view of articulated lumbar,

sacral, proximal caudal vertebrae and pelvis. Scale equals 1 cm.
Ahhreviaiions: cv, caudal vertebrae; l\, lumbar vertebrae.
Figure 9. \aranops hievirosiris. MCZ 1926. Ventral view and cross sections ol
last eight presacral \ertebrae. Anteriormost to the right. Scale equals 1cm.
The neural arches of the sacral vertebrae are deeply excavated

above the articular surfaces of the zygapophyses. Two poorly pre-
served sacra! ribs are present. Aerosaurus wellesii has three sacral
ribsand appears to be unique in this feature within the \aranopseids
(Langston and Reisz. 1981).

16 BREVIORA No. 474
trie cor
Figure 10. R. ehohnensis. MCZ 3150. Lateral view ol right scapulocoracoid

fragment. Scale equals cm.
I
Ahhrevluiiun.s: g, glenoid; trie cor, process tor the coracoid head ot the M. triceps.
The first caudal vertebra is distorted and thus appears wider than
the more posterior caudals. The neural spines of the caudal verte-
brae are smaller than those of the sacral or presacral vertebrae and
appear almost triangular in lateral aspect. All of the five articulated
caudals display excavated neural arches.
Appendicular Skeleton: Fragmentary right and left scapulocora-
coids are preserved. Both display at least a portion of the glenoid
and a well-developed process for the origin of the coracoid head of
the M. triceps (Fig. 10). In primitive fashion the glenoid is not raised
above the surface of the coracoid and continues back onto the tri-
ceps process. Similar conditions exist elsewhere in all known
varanopseids.
The right humerus is well preserved (Fig. 1) and has proportions
1
conformable with those present in Varanops (Table 2). Varanodon

and A. wellesii have a proportionately narrower proximal ends. The
proximal end of the humerus displays a stout M. latissimus tubercle.
This tubercle is poorly ossified in Varanops and A. wellesii but is
well ossified in A. greenleeorum and most other pelycosaurs. The
proximal surface in the region of the deltopectoral crest exhibits two

well-developed parallel ridges with a depression between them. Such
a condition exists in A. greenleeorum and elsewhere in pelycosaurs
only in caseids (Brinkman and Eberth, 1983). A feature of the

humerus not seen in other varanopseid taxa is that of a massively
developed delto-pectoral crest that continues distally as a sharp
ridge. This distal extension is so prominent that the entire region
appears almost square in anterior aspect. Romer and Price (1940)
distinguished A. greenleeorutn partly on the basis of a well-
d.p.cr.
lat tub
Figure 11./?. eliohriensis. MCZ 3150. A. Proximal dorsal view ol left humerus.
B. Proximal ventral view of the same. C. Proximal view of the same. D. Distal view
of the same. Scale equals cm.
I
Ahhreviaiions: dpcr. delto-pectoral crest; lat tub. latissimus tubercle.

18 BREVIORA No. 474
developed deltopectoral crest. Direct comparison indicates that A.

greenleeorum lacks the distal ridge. A strong ectepicondyle occurs
distally and is well separated from the partially preserved supinator
process. Much of the entepicondyle is missing, and the position of
the entepicondylar foramen is uncertain. Distoventrally, a large
radial condyle is preserved.
The is well preserved and has been completely pre-
right pelvis
pared laterally and ventrally. The medial surface of the ilium has
also been prepared. The pelvis is quite primitive in general structure,
having a posteriorly projecting iliac blade and a solid puboischiadic

plate. The ilium (Fig. 12A,B) is triangular in lateral aspect. It
becomes wider and longer ventrally where it forms the dorsal por-
tion of the acetabulum. The elongate, triangular posterior process is
low as in A. wellesii and Varanops but appears longer exhibiting
lower base/length and neck/ length proportions than in those two
species (Table 2). A small anterior projection of the ilium is poorly
preserved. In medial aspect (Fig. 12B) a ridge bisects the length of
the posterior process and appears conformable with that present in
allother varanopseids. The pubis is primitively short and ends ante-
riorly in a slightly thickened knob (Fig. 12D). In lateral aspect, the
dorsal margin of the pubis is thickened and appears concave. A
poorly preserved bump on the lateral surface, directly anterior to
the acetabulum, may be a pubic tubercle. A pubic tubercle is
unknown other varanopseid taxa, but is found in the ophiaco-
in
dontids Clepsydrops and Varanosaurus as well as caseids and eda-

phosaurids. The widespread occurrence of a pubic tubercle in
pelycosaurs as well as many cotylosaurs suggests that it is primitive.
The ischium is complete and also displays a slightly thickened
dorsal margin in lateral aspect (Fig. 12C). The ischiadic plate is thin
and meets its counterpart of the left side in the mid line. The ischium
appears conformable with the known morphology of that same ele-
ment in other varanopseids and ophiacodontids.
The right femur is complete (Fig. 13) although a small portion of
the proximal end is broken off and remains in articulation with the
acetabulum, obscuring the morphology of the latter. The proximal
end of the left femur is also preserved. Measurements and propor-
tions are listed in Tables 1 and 2 respectively. No significant dif-
ferences in femoral proportions between Ruthiromia and other
varanopseid taxa are apparent. In anterior aspect, the proximal end
Figure 12. R. elcohriensis. MCZ 3150. A. Lateral view of right ilium. B. Medial
view ol the same. C. Ventrolateral view ot right ischium. D. Ventrolateral view of
right pubis. Scale equals I cm.
Ahhrt'viaiuin: pt, pubic tubercle.
Figure 13. R. elcohriensis. MCZ 3150. A. Dorsal view of right femur. B. Ventral
view ol the same. C. Posterior view of the same. D. Anterior view of the same. Scale
equals 1 cm.
Ahhreviaiions: ac, adductor crest; 4th t, 4th trochanter; it, internal trochanter.
20 BREVIORA No. 474
of the right femur displays a large, tall internal trochanter well set
offfrom the thickened proximal surface of the femur. Varanops (the
only varanopseid where this condition can be compared) does not
exhibit such a well-developed internal trochanter. The intertrochan-
teric fossa is extensive, occupying about a third of the length of the
femur as in most non sphenacodontid pelycosaurs. A large knob

marks the point of attachment of the M. pubo ischio femoralis
internus. The distinct, although small, fourth trochanter is present
just distal to the intertrochanteric fossa. A prominent, well-ossified
adductor crest extends across the ventral surface of the element to
the fibulad edge. The crest is relatively larger than in any other
varanopseids and appears to be a primitive feature. In cross section
the shaft appears tear drop-shaped, in contrast to Varanops where a
cross section appears oval. In dorsal view, the two well-developed
ossified tibial condyles are well separated by a deep intercondylar
fossa. Their dorsal surfaces are essentially flat, and their articular
surfaces are oriented ventro laterally.
In dorsal view (Fig. 14) the right tibia and fibula have straight
external and concave internal edges. The fibula is slightly shorter
than the tibia (Table 1). The lateral femoral cotyle of the tibia is not
strongly compressed. Sharp ridges are present on the lateral and
medial edges of the distal half of the bone. The proximal end of the
fibula is damaged, so the extent of the dorsal tubercle is unknown.
The distal end is wide mediolaterally and compressed dorsoven-
trally. Similar overall conditions are seen to exist in all vara-
nopseids.
Eight elements are present in the tarsus: the astragulus, calca-
neum, a centrale and five distal tarsals (Fig. 15). The astragulus
(Fig.I5A,B) is L-shaped and supports both the tibia and fibula. The
neck of the astragulus is elongate and contributes to 35 percent of
the height of the element; it thus appears more like that of a sphena-
codontid than all other known varanopseids. It is unclear at present
whether this is a primitive or derived feature. In contrast to sphena-
codontids however, no tubercle is present on the ventral surface of
the astragulus medial to the arterial groove. The calcaneum (Fig.
I5A,B) is primitive in being wide and having a convex lateral edge
when seen in dorsal view. In contrast to the varanopseid genera A.

wellesii and Varanops, the element is much taller than it is wide.
The single centrale (Fig. 15C) is a small element with a limited
area of finished bone on its dorsal surface. The largest of the distal
t-"^"^
Figure 14. R. elcohriensis. MCZ

3150. A. Dorsal view of right tibia and fibula.
B. Ventral \iew of the same. Scale equals cm.
!
tarsals is the fourth. In dorsal view (Fig. 15C) it is essentially pen-
tagonal incontrast to the rectangular fourth distal tarsal of A.

wellesii and Varanops. The fourth distal tarsal also shares an equal
contact with the astragulus. The first and third distal tarsals are
subequal in size and the second is slightly smaller than these. This is
also the case in Varanops and A. wellesii. The metatarsals increase
in length from first to fourth (Table 1). The first metatarsal is sup-
ported entirely by the first distal tarsal. In dorsal view, the second
and third articulate with the distal tarsal of the same number. In
ventral view (Fig. 15D) each of these metatarsals is supported by
both the distal tarsal of the same number and the laterally adjacent
distal tarsal. The fourth metatarsal is longer than the combined
length of the firstmetatarsal and digit. This condition is seen devel-
oped to a greater degree in Varanops and to a lesser degree in A.
wellesii and occurs elsewhere within the pelycosaurs only in Varano-
saurus and Ophiacocion mirus. The first toe has a clawlike terminal
phalanx. The remaining toes are incompletely preserved or not
present.
In overall appendicular proportions (Table 2), Ruthiromia elco-
briensis appears more primitive than any other varanopseid in hav-
ing a humerus and femur of subequal lengths and a lower tibia/

femur ratio.
22 BREVIORA No. 474
Figure 15. R. ekuhriensis. MCZ 3150. A. Dorsal view of right astragalus/ calca-
neum complex. B. Ventral view of the same. C. Dorsal view of right pes. D. Ventral
view of the same. Scale equals 1 cm.
Abbreviation: c, centrale.
DISCUSSION
Ruthiromia elcobriemis considered a pelycosaur on the basis of
is
the combined, derived characters (within reptiles) of excavated neu-

ral arches, keeled mid-dorsal vertebrae, a strongly developed pro-
cess for the coracoid head of the triceps and a keeled angular in the
region of the articular. These constitute a suite of characters which

is found only within the pelycosaurian families Varanopseidae and
Sphenacodontidae.
The Varanopseidae is currently defined by cranial characters
(Langston and Reisz, 1981), yet shows a unique combination of two
derived, postcranial characters: excavated neural arches and the
presence of a shallow fossa on the proximal surface of the deltopec-

toral crest(Brinkman and Eberth, 1983). Ruthironiia eUuhriensis \s
tentatively assigned to the Varanopseidae on the basis of displaying
thesesame characters. Following the cladistic classification of
Brinkman and Eberth (1983, Fig. 16) for the better known pelyco-
saur genera, the derived character of excavated neural arches is
thought to have developed either two (Fig. 16A) or three (Fig. 16B)
times. All varanopseid and sphenacodontid taxa, where postcranial
material is available, show excavated neural
arches. Within the eda-
phosaurids an undescribed specimen (E2) from the Pennsylvanian

of Kansas shows this condition (R. Reisz, personal communica-
tion). Within this context and assuming parsimony, four equally
correct positions denoting propinquity of descent of Ruthiromia
elcobriensis may be depicted in a cladogram (Fig. 16C).
Position 1
places Ruthiromia elcobriensis as a member of the
varanopseids in the clade including Varanopsj Aerosaurus and
Casea. This clade is united by the derived morphology of the prox-
imal surface of the humerus in the region of the delto-pectoral crest.
As described above, Ruthiromia elcobriensis shares this same
morphology.
Relationships indicated by positions 2, 3, and 4 each necessitate
reversals of those characters at nodes (c), (c,d,e) and (c,d) respec-
tively. The clade including Ophiacodon, Edaphosaurus, and Dime-

trodon defined by the shared presence of a rodlike paroccipital
is
process with a socket on the ventral surface for contact with a

distinct dorsal process of the stapes, a concave ventral margin of the
cheek and a single postparietal (node c). R. elcobriensis has a nar-
row paroccipital process that shows no evidence of a socket along its
ventral edge for contact with a dorsal stapedial process. A relation-
of this
ship indicated by position 2 therefore involves the reversal
character. Other regions are not preserved in the holotype. Sim-
ilarly, Ruthiromia elcobriensis does not show

the derived characters
of the twisted prearticular and ventromedial projecting pterygoideus
process which partially define the clade consisting of Edaphosaurus
and Dimetrodon (node d). In this case, a total of three reversals
(reversal at both c and d) isnecessary to explain the relationship
indicated by position 4. Finally, the relationship indicated by posi-
tion 3 necessitates the reversals listed above as well as reversals of
the autapomorphic characters that define the Edaphosauridae and

24 BREVIORA No. 474
EDAPHOSAURIDAE
OTHER REPTILES VARANOPSEIDÊ OPHIACODONTIDAE EDAPHOSAURUS E2 SPHENACODONTIDAE
EDAPHOSAURIDAE
OTHER REPTILES VARANOPSEIDAE OPHIACODONTIDAE EDAPHOSAURUS E2 SPHENACODONTIDAE
EDAPHOSAURIDAE
OTHER REPTILES VARANOPSEIDAE OPHIACODONTIDAE EDAPHOSAURUS SPHENACODONTIDAE
higurc 16. Cladograms depicting relationships of pelycosaur families. Adopted

and modified from Brinkman and Eberth (1983). A&B. Two equally parsimonious
hypotheses for the appearance and distribution of excavated neural arches in pelyco-
saurs. Solid bars represent those points where excavated neural arches are exhibited.
Dashed bar represents point of character loss. C. Four equally correct positions for
Ruihironiia on the basis of the derived character of excavated neural arches.
are not seen in Ruthiromia elcobriensis (elongate neural spines with

lateral tubercles, low position of the jaw articulation relative to the
tooth row). Clearly, the most parsimonious course to take is to
tentatively place Ruthiromia elcobriensis among the Varanopseidae

as a primitive member
of that family. The tentative nature of this
placement both the lack of diagnostic cranial material in the
reflects
type and the employment of a combination of derived postcranial

characters, none of which can serve alone as an autapomorphic
character of the family Varanopseidae.
Ruthiromia appears to be more closely allied with A. wellesii in
sharing the unique character of the forwardly placed medial process

of the articular and the keeled angular.
Ruthiromia shares only primitive characters with Varanops and
appears similar retaining a laterally projecting paroccipital pro-
in
cess and having only two sacral vertebrae.

in
The morphology of the lumbar vertebrae and more specifically

the position in the column where keeled vertebrae first appear is
more The seven articulated presacral vertebrae of
difficult to assess.
Ruthiromia show no signs of possessing keels and all retain a gently
rounded ventral surface. The inclusion in the holotype of one disar-
ticulated lumbar (posterior dorsal) with a similarly unkeeled ventral
surface places the minimum location for the transitional vertebra at
a position nine vertebrae in front of the pelvis. In addition, a signifi-
cant difference in diapophysis orientation between the transitional
vertebra and the isolated lumbar (lateral vs. postero lateral respec-
tively) warrants placement of the transitional vertebra even further

forward in the column. Varanops develops a fully keeled vertebra
six vertebrae ahead of the pelvis (Fig. 9). The fifth is transitional,
displaying three parallel ridges along the ventral surface. The only
fully prepared vertebrae known from A. wellesii that can be placed
in the column with certainty are those nine and ten positions ahead
of the pelvis. These show keeled ventral surfaces. No idea concern-

ing the initial position of keeled vertebrae can be determined in
either A. greenleeorum or Varanodon, but A. greenleeorum does
not appear to develop strong keels of the sort seen in the one known
anterior dorsal of Ruthiromia anywhere in its column. Ruthiromia
similarly does not share with A. greenleeorum the condition of the

deeply recessed anteroventral lips of the anterior dorsals. Keeled
lumbar vertebrae are considered an advanced character state (Romer
26 BREVIORA No. 474
and Ruihiromia appears more primitive than either

Price, 1940).
Varanops or A. wellesii in displaying a greater number of unkeeled
vertebrae in the presacral region of the vertebral column.

In summary, Ruihiromia elcobriensis represents a varanopseid
which very primitive postcranially, with some unique vertebral
is
specializations. Cranially, it shows some of the same advanced spe-

cializations seen in A. wellesii and perhaps Varanodon.
ACKNOWLEDGMENTS
The authors wish to express their gratitude and appreciation to
Robert Reisz for his many valuable comments and criticisms as well
as the technical assistance that he provided during the preparation
of this manuscript. Although his help must be regarded as a crucial
factor in the production of this paper, the authors assume full
responsibility for any errors that might be present.

Travel and research expenses were covered, in part, by a Schmitt
Fund research grant provided by the Field Museum of Natural
History, Chicago.
LITERATURE CITED
Brinkman, D., and D. Eberth. 1983. The interrelationships of Pelycosaurs.
Breviora Mus. Comp. Zool., No. 473.
Carroll, R. L. 1969. The origin of reptiles, pp. 1-44. In A.d'A. Bellairs el al.
Biology of the Reptilia. New York, Academic Press, 373 pp.

(eds.),
Fracasso, M. a. 1980. Age of the Permo-Carboniferous Cutler Formation ver-

tebrate fauna from ElCobre Canyon, New Mexico. J. Paleontol., 44: 156-163.
Heaton, M. J. The Cotylosauria: A reconsideration of a group of archaic
1980.
tetrapods, pp. 497-551. In. A. L. Panchen (ed.). The Terrestrial Environment

and the Origin of Land Vertebrates. Systematics Assoc. Spec. Vol. No. 15,
London, Academic Press, 633 pp.
Langston, W. Jr., and R. R. Reisz. 1981. Aerosaurus wellesii, new species, a
varanopseid mammal-like reptile (Synapsida: Pelycosauria) from the lower
Permian of New Mexico. J. Vertebr. Paleontol., 1(1): 73-96.
Olson, E. C. 1965. New Permian vertebrates from the Chickasha Formation in
Oklahoma. Oklahoma Geol. Surv. Circ. No. 70, 70 pp.
Romer, a. S., and L. I. Price. 1940. Review of the Pelycosauria. Geol. See.
Amer. Spec. Pap. No. 28, 538 pp.
^^5. COMP. ZOOL
MAR 1 8 1985
^VARO
B R I O R A
11 seiini of Comjâralivo Zoology
IS ISSN ()()()6 9698
NEW OR PROBLEMATIC AISOLIS FROM COLOMBIA.

l.ANOUS CAL/MAE, NEW SPECIES, FROM THE
CLOLD FOREST OF WESTERN COLOMBIA
Stephen Avala,' Dennis Harris,- and Ernest E. Williams-*
Abstract. A noli.', calimae, new species, is described from the cloud forest of
western Colombia in Departmento Valledel Cauca. Referable to the /I /7(;//.v /;;//;< /t//i/.v
species group, it is distinctive in its coloration, in the absence of an interparietal scale,
and in the apparent trend to reduction of the elongate anterior supraciliar> scale usual
in Ano/i.s.
INTRODUCTION
In a recent paper (Williams, 1982), the description of three new
specieswas made the occasion of a summary of the eastern members
odhe punctatus species group. Several new species must be described
before a similar summary will be possible for the western punclalus
group. The first is here described, a small species and initially recog-
nized only from a single specimen collected in cloud forest near Lake
Calima. Even with one specimen, its striking color pattern and dis-
tinctive habitus made it obvious that it is a new species. Subsequently,
three additional specimens have been obtained, one from the original
locality, two from Television Tower Mountain near Cali. These four
specimens have been divided between the Museum of Comparative
'120 Vista View Place, Petaluma, California 94952.

-Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109.
'Museum of Comparative Zoology. Harvard University, Cambridge. Massachusetts
02138.
2 imi-N lORA No. 475
Zoology and the InstitutodeCiencias Naturales(ICN, Bogota). Still

more recently a Museo La Salle (MLS, Bogota) specimen from a
third locality has been recognized as belonging to this species. At the
suggestion of Fernando Castro, we call it after the place of its first

discovery:
A noils calinwe, new species
(Figs. 1-6)
Type: MCZ 158392, adult male.

Type localliy: San Antonio, Television Tower Mountain, Depto.
Valle del Cauca, Colombia, (3° 28'N 76° 40' W) 1,800 m elevation,
Dennis Harris, Humberto and Fanny Carvajal, coll., 23 January
1980.
Paratypes (aW from Valle): MCZ 158393, adult female, same data
as type; ICN
3678. adult female, approx. km below Lake Calima
1
dam, (3° 50'N 76° 32'W) Dennis Harris, coll., 18 January 1980; ICN
3679, adult male,same place as ICN 3678, William Duellman and
Fernando Castro, coll., 17 March 1979. MLS 122: Mares. 3 km N of
San Antonio (3° 30'N 76° 40'W).
Diagnosis. A
small green cloud forest species of the punetatus
group distinguished by its short body, legs and tail, consistent
absence of the interparietal scale, a blunt and weak canthal ridge with
poorly differentiated canthal scales, only a short supraciliary scale
followed by granules or granules only, small, low number of loreal
scales (4-5), smooth ventralsand very short stubby toes with 15 to 17
lamellae under phalanges ii and iii of fourth toe. Dewlap present in
both sexes, small in female and with larger scales.
Description. Head. Rather short. Head scales small, flat, very
slightly wrinkled. Seven to ten scales across the snout between the
second canthals. Frontal depression shallow, the scales within it as
large or larger than some of those anterior to it. Four to seven scales
border rostral posteriorly. Circumnasal scales of each side separated
from rostral by one elliptical scale which lies above the suture
between rostral and flrst supralabial. Six scales between supranasals
dorsally. Snout elongate, slightly protuberant, extending slightly
beyond mental.
Supraorbital semicircles separated in both males by two rows of
large scales, as large as the scales of the semicircles, in the females in
contact or separated by one row of small scales or granules. Supraor-
1983 ANOLIS CAI'LIMAE
Figure 1. Anulis calinwe. new species, in life. Female paratype above, male type
below.
bital scales in contact with or barely separated from the supraocular
disk of each side, which consists of six to ten enlarged, slightly

wrinkled scales, remainder of the supraocular area granular or sub-
granular. One or two short supraciliaries on each side followed by
granules, or only granules present. Canthus blunt, canthals small,
poorly differentiated, the first or first and second largest. Loreal rows
four or five, the lowermost slightly larger.
Temporals granular. An indistinct intertemporal double line of
slightly enlarged scales. Supratemporals granular laterally, becoming
larger and flattened toward the interparietal area, which is bounded
by weak ridges. No differentiated interparietal, but a zone of enlarged
scales, largest anteriorly and laterally, posteriorly grading slowly into
the dorsal granules. Ear opening small, elliptical, but larger than any
scale in the interparietal area. Occiput with small blunt median knob,
obvious in males, not evident in females.
Suboculars in contact with supralabials. Seven supralabials to the
center of the eye.
BREVIORA No. 475
Figure 2. Anulis calimae, new species. Male type, MCZ 158392. Dorsal view of
head.
Mental semidivided, each part slightly or distinctly wider than

deep. Four granules are in contact with the mental between the large
first sublabials. Two to five sublabials in contact with the infralabials.
Gular scales subgranular, very little enlarged laterally where they
border the sublabials.
Trunk. Dorsal scales granular, convex, subequal. Ventrals much
larger,smooth, juxtaposed or subimbricate, rounded or slightly
pointed in males, broader, squarish in females, in tranverse rows.
Lateral chest scales not keeled. Males may develop a low nuchal crest
when aroused.
1983 ANOLIS CAULIMAE
Figure 3. Anolis calimae. new species. Male type, MCZ 158392, Lateral view of
head.
Dewlap. Present in both sexes. Large in male, reaching anterior

abdomen, scales small, smaller than ventrals, weak, in rows, one scale
wide, separated by naked skin, edge scales larger, somewhat crowded,
imbricate, smooth; small to moderate in females, scales larger, as
large or larger than ventrals, in close-packed rows, smaller and much
more numerous along the edge.
Limbs and digits. Upper arm scales granular. Lower arm scales
granular to larger and unicarinate. Leg scales granular posteriorly,
larger and uni- to multicarinate anteriorly. All supradigital scales
multicarinate. Toes relatively short and stubby. Digital dilations
moderately wide. Fifteen to 18 scales under phalanges ii and iii of

fourth toe.
Tail. Compressed, short, ca. 1.5 X body length, possibly prehen-
sile. tail crest. Two weakly keeled middorsal rows, ventral rows
No
larger and more strongly keeled, verticils not evident. Scales poste-
rior to vent smooth. Large postanals in males, none in females. Tail
base prominently swollen to accommodate hemipenes in adult males.
Size. Anolis calimae is a small but somewhat robust species. Sizes
of the four recent specimens are: 59 (type), 58, 55, and 58 mm
snout-vent length respectively.
Color. This is a green anole with a considerable capacity for rapid
pattern and color change. It may be almost uniform green or yellow-

green with little or no pattern, or it can have three prominent broad
BREVIORA No. 475
Figure 4. Anolis caliniae. new species. Male type, MCZ 158392. Ventral view of
head.
black bands across the back and sides (apparently more prominent in
cream colored spots in the dark bands.
the males), with pale yellow or
At another state of excitement, the sides and back are mostly green-
grey with scattered small black spots on the sides and neck and a few
dark brown crossbars on the vertebral line. Females especially may
show alternating sets of narrow, light yellow and dark brown-black
spots along the midline. There is no prominent dark band across the
head between the eyes. The large male dewlep is unpigmented: pale
yellow-green with salmon pink near anterior edge and white or pale
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8 BREVIORA No. 475
yellow scales. The small female dewlap is pigmented: blue with yellow
or white scales. The tail is banded: broad dark bands in the male and
narrow bands in the female. The lining of the throat is pale. The iris of
ICN 3678 (a female) was golden yellow above and below a zone of
orange; that of the holotype male, MCZ 158392, was orange.
Preserved specimens are greenish-grey with few to many small but
prominent dark (and light in the male) spots on the back and sides,
elongate dark and light spots along the dorsal midline and a darker
brown, unpatterned head. The belly is pale, with small grey spots
under the chin. The dewlap has rows of white scales over white skin
(males) or pigmented skin (females). The peritoneal kjning is heavily
pigmented.
Habitat and reproduction. The two recent collection sites are
about 50 km apart in the same cloud forest region (tropical premon-
tane wet forest) in the western Colombian cordillera. All four speci-
mens were collectedat night while they were sleeping in exposed sites
40 to 150 cm above the ground, the type and first paratype on a fern
leaf and a low shrub, in a cool forested region at about 1,800 m
elevation. The two Lake Calima paratypes were on exposed twigs, in
a somewhat warmer, more densely vegetated area also subject to
frequent rains and cool fogs at 1,300 m elevation. Other anoline
species known to occur in the same sites or in the same general area
arey4. ventrimaculatus, A. eulaemus, A.fraseri, A. antonii, Phenaco-
saiirus heterodermus and an undescribed punctatus group anole
known at present from a single specimen.
The specimens were in reproductive condition at the time of cap-
ture: the males with enlarged testes (6. x 4.0 mm approx.), and the
1
females with a single oviducal egg on one side and an enlarged,

yolking follicle on the other.
Etymology. The name calimae refers to the site where William
Duellman and Fernando Castro collected the first specimen. Lake
Calima is in turn named after the Calima Indians who inhabited the
region centuries ago.
Comparisons. A. ra///;m£' is a very distinctive anole. No previously
described member punctatus group is known to lack an inter-
o{\\\q
parietal scale. This condition is unusual in any group of Anolis, but it

is curious that it is known as a moderately common variation in two
species of the aequatorialis species group (A. ventri/naculatus and /4.
gemmosu.s) which, like calimae, are inhabitants of Andean cloud

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10 BREVIORA No. 475
forest.It is possible, since there are only four specimens of calimae
thus far known, that absence of an interparietal will be found to be

inconstant in this species too.
The wholly granular supraciliary margin found in one specimen of
calimae is unique, not only for the puncaiatus group, but for the
genus Anolis. The alternative and commoner condition (in
three of
the four calimae) of one elongate but short supraciliary scale is

known elsewhere (e.g., in the ligrinus species group); the extreme
shortness of the scale in the three specimens may, however, be indica-
tive of a strong trend toward de-differentiation of this scale, which
then culminates in the completely granular margin.

The presence of dewlap in both sexes, and the fact that it is
a
smaller and differently pigmented in the female, may be a primitive
feature of calimae and occurs erratically in a number of Anolis
species groups. In the pwutatus group the condition is known in
chocorum and in transversalis, and in these species, as in calimae, is

associated with a more or less marked difference in body color and
pattern between the sexes.
size of the scales in the
Unique to calimae is the larger dewlap
female. The
reverse or equal-sized scales is typical for Anolis species
in which the female retains a dewlap.
Unusual also and requiring confirmation by additional material is

the apparent difference between males and females of calimae in the
size and number of scales between the supraorbital semicircles.
From two other previously described punctatus group species
the
of western Colombia A. calimae is as distinctive in color as in scales.
Both these species are also Pacific lowland forms: Anolis chocorum
(also a larger species, reaching 79 mm snout-vent length) has the
dorsum uniform green or with oblique rows of dark green blotches on
the flanks, never with prominent black bands enclosing yellow or
cream spots. The male dewlap is orange laterally, green basally; or the
female dewlap isgreen with pale yellow or gray at the base. The rows
of scales in the dewlaps are three to four scales wide. Anolis chloris
(about the same A. calimae, ca. 55
size as mm
snout-vent length ) is
grass-green with
in color, the potentiality of turning a dark olive
green which then may have diagonal darker bars, but again there are
never the black bands of calimae. The dewlap color in males is yellow
or white, never green, the rows of scales in the dewlap are one scale
wide and there is no dewlap in the female. In contrast to A. calimae,
both A. chocorum and A. chloris turn blue or purple in preservative
1983 ANOLIS CAULIMAE 11
rather than the greyish green of A. calimae. The occiput knob is not
seen in A. chocorum or A. chloris.
There has been no evident close relative for A. calimae. However,
an undescribed species from the same region, known only from a
juvenile male, seems closer than any other, although still sharply
distinct.
ACKNOWLEDGMENTS
We are grateful to the collectors of the species here described,
William E. Duellman, Humberto and Fanny Carvajal, and Fernando
Castro. Ayala's work in Colombia was sponsored by grants from the
U.S. Public Health Service (NlAlDAl-A21511),Tulane University
(NIAID A1-I0050), and COLCIENCIAS, the Colombia National
Science Institute. The drawings are by Laszlo Meszoly.
LITERATURE CITED
Williams. E.E. 1982. Three new species of the Anulis puintaius complex irom
Amazonian and mter-Andean Colombia, with comments on the eastern members
o{ \.\\e punctatus group. Breviora Mus. Comp. Zool. No. 467. pp. 1-38.
MUa COMP, ZOOL
LfRr?ARY
MAR 1 8 1985
ARVARD
B K E V-'f^'t) R A
Meseiim of Comparative Zoology
us ISSN 0006 9698
TOWNSEND'S UNMAPPED NORTH ATLANTIC

RIGHT WHALES (EUBALAENA GLACIALIS)
William E. Schevill' and Karen E. Moore'
Abstract. Townsend's detailed maps of worldwide whale distribution from 19th

century American whaling logbooks (1935, Zoologica, N.Y., 19: 1-50) omitted his
North Atlantic records for Euhalaena. We have read 12 of his 15 sources and have
mapped the right whales therein recorded.
INTRODUCTION
Very little is known about the distribution of right whales (Euba-
laena glacialis) in the North Atlantic Ocean. Among others, J. A.
Allen (1908), Collett (1909), G. M. Allen (1916), Thompson (1928),
Slijper, van Utrecht, and Naaktgeboren (1974), Reeves, Mead, and
Katona (1978), Schevill, Moore, and Watkins (1981), Kraus and
Prescott ( 98 ), Reeves and Brownell ( 982), and Watkins and Sche-
1 1 1
vill (in press) have discussed sightings and catches in particular areas.
Townsend (1935), in his study of "the distribution of certain whales as

shown by logbook records of American whaleships," examined log-
books from 1,665 voyages, worldwide, mostly of the 19th century.
Out of the 8,415 Eubalaena taken, he found only 35 in the North
Atlantic, recorded on 5 voyages, and did not bother to plot these on
1
his charts by latitude, longitude, and month as he did the rest of his
totalof 53,877 whales ofsix species (see his pp. 10 and 18). Our note is
a response to this omission, and like Townsend's paper is based only
'Museum of Comparative Zoology. Cambridge, MA 02 138 and Woods HoleOceano-

graphic Institution,Woods Hole, MA
02543.
-Woods Hole Oceanographic Institution. Woods Hole. MA 02543.
2 BREVIORA No. 476
on Yankee whaling logbooks. Thus it does not deal with the historic
Eubalaena fishery of the eastern North Atlantic, well-established in
the Bay of Biscay by the 12th century, and continuing intermittently
along the western coast of Europe from the Mediterranean to the
Barents Sea (where this whale got the name of nordkaper) and
Iceland; this fishery pretty well stopped in the 1920's, and now
Eubalaena is very rarely seen in those waters.
Because so little is known, the location and season of even Town-
send's few North Atlantic right whales are important. We have con-
sulted 1 2 of his 15 sources (Table 1
), but failed to find the other three
logbooks, which he credits with one North Atlantic rightwhale each
(sloop Greyhound, 1753; ship Governor Troup, 1868-70; bark Sea
Ranger, 1879-84). The Providence Public Library has a partial
journal by William H. Tilton, a boatsteerer/ third mate of Sea
Ranger during this voyage, but it does not tell where whales were
encountered. The 2 logbooks that we read are listed by Townsend as
1
recording the other 32 Eubalaena glacialis.

Our results do not entirely agree with Townsend's. For one thing,
as he explains on page 6, his counts of whales include "not only the
1
whales turned into oil ['saved', as the whalers said], but also those
killed. but subsequently lost." In our logbook reading we have
. .
tried to separate these categories, and since we are interested more in

whale distribution than in oil production, we have also counted
whales sighted (and identified) but not struck. The logs do not always
specify the numbers sighted if greater than or 2. When they said 1
merely "right whales," we counted it as 2+; the sum of 5+ and 4+ is

given conservatively as 9+, though the actual count might have been
appreciably higher. We assume that these old-time whalers were
competent recognizing the species of interest
at probably much
—
more reliably than most seafarers of today; sometimes sightings were
logged simply as "whales," and these we did not count. We cannot
explain our discrepancies with Townsend, and can only say that we
have reported what we found in our reading. We failed to find
mention of the single Eubalaena each that he recorded for Jireh
Swift and Emma Jane, and of the 3 he ascribed to Endeavor.
1983 NORTH ATLANTIC RIGHT WHALES
Ji
BREVIORA No. 476
FINDINGS
The 12 voyages summarized (Table \) for Eubalaenaglacialis in the
North Atlantic were made by 1
whaleships over a 45-year span ( 853
1 1
to 1898). The highest and lowest latitudes reached by these are well
within the known range of this species. No Euhalaena were recorded
in the months of January, April, September, October, and December.
The records from November through March are all south of N. lat.
32°, and those from May through August are between N. lat. 46° and
62°, but there is no detailed evidence for migration patterns. We find
the whales in high latitudes in summer and lower in winter, but there
are no helpful hints of routes to and fro. The wint«r records are right
along the shore on both sides of the Atlantic (Cintra Bay, N. lat.
23° in western Africa, and N. lat. 31° at the Georgia-Florida border
in North America); on each coast the ship sometimes anchored and
whaled from the boats.

Six voyages (Ansel GiBBS in 1868, Daniel Webster and A. J. Ross
in 1877, A. R. Tucker in 1880, Palmetto in 1886, and Canton in
1897) reported whales in June, July, and August to the eastward of

Cape Farewell, Greenland, between N. lat. 59° and 62°, W. long. 32°
and 38°, at the southern entrance to Denmark Strait. They logged 12
whales saved, 7 struck but lost, and at least 16 more sighted; Town-
send recorded 16 whales, but without specifying the region (3 of these
voyages whaled also in other areas). These 35 whales were encoun-
tered in 26 ship-weeks of hunting in five years, which implies a
consistently good hunting ground. This ground is not referred to by

name in these logs. We found no hint that bowheads were seen or
taken, although they might have been expected in these waters. (For a
recent report, see Jonsgard, 1982, Fig. 1.) As pointed out by J. A.
Allen (1908: 281-288 and especially 286), since the 17th century
whalers have distinguished bowhead {Balaena mysticetus) from
nordkaper, or right whale in American usage {Eubalaena glacialis),
both on morphology and oil yield, as well as on behavior. While the
Ross was on these Denmark Strait grounds near N. lat. 60° and W.
long. 35° the whales were specified as "Wright Whales" and seven or
,
more were sighted, of which four were saved. After leaving these
grounds, the Ross went through the Canaries, and on 24 November
"Saw A Wright Whale" in "Senter Bay" (Cintra Bay) on the African
coast. Therefore we conclude that only Eubalaena glacialis were
found on these voyages.
70° N
^
\3^ VII-68
mj VI.VIII-77
^V VI.VII-80
VII.VIII-86
CINTRA BAY-
XI- 57
E.m-58
XI-77
80° W 60° 40° 20° 0°
Figure Distribution of North Atlantic right whales ( Eubalaena glacialis) recorded

1 .
in American whaling logbooks between 1857 and 1897. The dots show where whales
were encountered at sea, and the coastal encounters are indicated by arrows. Months
are indicated by roman numerals.
Although these whaleships departed and returned to port (usually

New Bedford) in all seasons, there is no mention of right whales seen
near the New England or Canadian coasts. Before the mid 1700's
right whales were caught in Massachusetts waters
from small shore-
based boats, mostly in the winter and spring (Macy, 1835). From
1955 to the present, right whales have been observed locally in all
seasons (Schevill, Moore, and Watkins, 1981; Watkins and Schevill,

in press). It seems highly unlikely that these 19th century whalers
would have ignored such valuable animals wherever seen, and so it
may be that Eubalaena were then very scarce in these coastal waters,
although they had been abundant there from the 16th well into the
18th centuries.
6 BREVIORA No. 476
Against the background of scanty and often imprecise data on the

seasonal and geographic distribution of Eubalaena glacialis, even
these few additional points are significant. We have mapped about 60
whales logged on 9 voyages by 8 ships during the 40 years from 1857
to 1897. half of these were in the Denmark Strait area in
More than
June, July, and August. More than 3 of the remainder were at Cintra
1
Bay in November, February, and March.

A question about the North Atlantic Eubalaena has been whether
there was more than one stock in the population. Did the relative
abundance that we report for Denmark Strait a hundred years ago
hint that right whales from both sides of the North Atlantic sum-
mered together, indicating one general population for this ocean? Or
might there have been two stocks, with Cintra Bay perhaps being in
the winter range of the nordkapers of the Barents Sea? This is the sort
of problem for which radio tagging and tracking are needed. There
are still at least several hundred Eubalaena in the North Atlantic.
ACKNOWLEDGMENTS
We would like to acknowledge help from
Stuart C. Sherman, who
told us where Richard Kugler and Virginia
to find the logbooks,
Adams of the Old Dartmouth Historical Society, Paul Cyr of the
New Bedford Public Library, and Lance Bauer of the Providence
Public Library; Colleen Hurter helped with bibliothecal matters.
William A. Watkins and Richard H. Backus helpfully commented on
the manuscript. Support for this study was from the Oceanic Biology
Program of the Office of Naval Research, Contract No. NOOO 14-82-

C0019 NR083-004. This is Contribution No. 5241 from the Woods
Hole Oceanographic Institution.
LITERATURE CITED
Allen, G. M. 1916. The whalebone whales of New England. Mem. Boston Soc. Nat.
Hist. 8(2): 107-322, plates 8-16.
Allen, J. A. 1908. The North Atlantic right whale and its near allies. Bull. Am.
Mus. Nat. Hist., 24: 277-329, plates 19-24.
CoLLETT, R. 1909. A few notes on the whale Balaena glacialis and its capture in recent
years in the North Atlantic by Norwegian whalers. Proc. Zool. Soc. Lond., 1909:
91-98, plates 25-27.
JonsgArd, a. 1982. Bowhead (fla/acAia rnysticetus) surveys in the Arctic Northeast
Atlantic waters in 1980. Rep. Int. Whal. Commn., 32: 355-356.
Krais, S. D.. and J. I98L Distribution, abundance and notes on

H. PRtscori.
Bay of Fundy, summer and fall 1980. Final Report to
the large cetaceans of the
the U.S. Department of Commerce. NOAA, National Marine Fisheries Service,
87 pp.
Macv, O. 1835. The History of Nantucket. Milliard, Gray, and Co.. Boston, 300
pp. Reprinted by Research Reprints Inc.. York, 1970.New
Reeves. R. R. and R. L. Brow nell, Jr. 1982. Baleen whales. Euhalaena ^lai iaiis
and allies, pp. 415-444. //; J. Chapman, G. Feldhamer (eds.). Wild Mammals of
North America. The Johns Hopkins University Press.
Reeves, R. R., J. G. Mead, and S. Katona. 1978. The right whale. Eubalaena
glacialis, in the western North Atlantic. Rep. Int. Whaling Comm., 28: 303-312.
SciiEviLL, W. E.. K. E. MooRh, AND W. A. Watkins. 1981. Right whale. Euba-
laena glacialis. sightings in Cape Cod waters. Ret. No. 81-50, Woods HoleOcean-
ographic Institution, 16 pp.
Slijper, E. J., W. L. van Utrecht, and C. Naaktgeboren. 1964. Remarks on the
distribution and migration of whales, based on observations from Netherlands
ships. Bijdragen tot de
Dierkunde, Aflevering 34: 3-93.
Thompson, D'Arcv W. On whales landed at the Scottish whaling stations
1928.
during the years 1908 1914and 1920-1927. Fishery Board Scotl., Sci. Invest. 3,
39 pp.
Townsend, C. H. 1935. The distribution of certain whales as shown by logbook
records of American whaleships. Zoologica, 19 ( 1): 1-50, plates 1^.
Watkins, W. A., AND W.E.ScHEViLL. In press. Observations of right whales (£w/>a/aeA7fl
glacialis) in Cape Cod waters. Fishery Bull.
POSTSCRIPT
While this paper was in press, we were enabled by the courtesy of
Professor Howard E. Winn of the University of Rhode Island to
consult a collection of about half the worksheets used in preparing
the maps of Townsend's 1935 compilation. These had been pre-
served at the New York Zoological Society; for each logbook they
tally the date, location, and species of each whale taken. The log-
books are of course the primary sources, but these tallies tell us
something of the compilation, and account for occasional discrep-
ancies. Thus we have learned that the particular Emma Jane log-
book read by us is incomplete; it ends on January 1880. The tally 1
shows that the voyage continued into August 1881. and that this
schooner did take a right whale on 5 February 880 off Brunswick,
1 1
Georgia, near where Golden City took her 2 whales of 1876 and
1882, also in the latter part of February. The tally for Endeavor
1854-1856 confirms our failure to find in her logbook any right
whale captures during either North Atlantic passage, and shows that
8 "^'^''BREVIORA No. 476
the 3 whales pubHshed as frorn that ocean were actually taken in the
Sea of Okhotsk. The GoverS/or Troup tally for 1868-1870 tells us
that the whale listed for the North Atlantic was taken in the South
Atlantic, 20 miles east of Tristan da Cunha in about 37° 15'S. We
still have not seen her actual logbook. The Canton tally for
1897-1898 counts of the 2 whales that we list as struck and lost in
1
July; the other 2 cited as North Atlantic turn out to have been taken
in the South Atlantic in November. The sheets for Jireh Swift are
missing, as are those for Sea Ranger and the sloop Greyhound; we
hope that someone will find our three missing logbooks.
MAR 18m
B R E V I -R A
useum of Comparative Zoology
us ISSN 0006-9698
Cambridge, Mass. 7 September 1984 Number 477
NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA.

II. ANOLIS PROPINQUUS, ANOTHER NEW SPECIES
FROM THE CLOUD FOREST OF WESTERN COLOMBIA.

Ernest E. Williams'
Abstract. A. propinquus. new species, from the cloud forest of Departamento

Valle in western Colombia resembles sympatric A. calimae Ayala, Harris, and
Williams in the absence of an interparietal scale, but differs by its uniform coloration,
distinctly keeled head scales, elongate supraciliary scales, and blue rather than yellow
dewlap.
Another western punctatus group species has languished un-

recognized in the Museum of Natural History, the University of
Kansas (KU), since its collection in 1974. Known only from a male
near hatchling, is unremarkable as
it regards color, but it is
structurally nearly as distinctive an animal as the recently described
A. calimae with which it occurs.
Because it is another member of an anole fauna that is just
beginning to be known, it receives the Latin name that means
'neighbor':
Anolis propinquus, new species

(Figs. 1-3)
Type: KU 169833, male juvenile.

Type locality: Rio Calima, 1.5 km Lago Calima, Valle, W
Colombia, W. E. Duellman, coll., 15 September 1974.
Diagnosis. Another green cloud forest species of the punctatus
group somewhat similar to A. calimae and, like that species, lacks a

02138.
BREVIORA No. 477
Figure 1. Anolis propinquus, new species. Holotype, KU 169833. Dorsal view of

head.
1984 ANOUS PROP/NQUUS. NEW SPECIES
Figure 2. Anolis propinquus. new species. Holotype, KU 169833. Lateral view of

head.
parietal eye, but distinguished among other features by keeled head

scales, an elongate supraciliary scale which occupies half the
supraciliary margin, a higher number of loreal rows (7) and a higher
lamellar count under phalanges ii and iii of the fourth toe (25), and
its blue rather than yellow dewlap.
Description. Head. Blunt, short. Head scales rather small, most
scales unicarinate, the keels irregular in direction. Twelve scales
across snout between second canthals. Frontal depression very
shallow, the scales within it larger than those anterior to it. Six
scales border rostral Anterior nasal scale weakly
posteriorly.
differentiated, narrow, separated from the rostral-first supralabial
suture by one scale. Eight scales between supranasals dorsally.
Supraorbital semicircles separated by three rows of scales, the
lateral ones almost as large as the scales of the semicircles and
keeled, the middle row minute, granular. Supraocular disk mod-

erately developed, containing ca. 21 keeled scales, those next to the
semicircles largest, grading laterally into smaller but still keeled
scales,two rows of subgranular scales separating the disk on each
side from the supraciliaries. Anterolateral corner of supraocular
area with subgranular scales, posterolateral corner with minute
granular scales. One elongate supraciliary extending half the
supraciliary distance and followed by granules. Canthus blunt, of
ca. seven scales, the first, second and third larger. Seven loreal rows
below second canthal.
BREVIORA No. 477
Figure 3. Anolis propinquus, new species. Holotype, KU 169833. Ventral view of

head.
Temporal scales granular, a triangle of subgranular intertemporal

scales. Supratemporal scales granular, minute, becoming abruptly
larger toward the interparietal area. Ear small, but larger than any
presumptive interparietal. Enlarged scales of interparietal area

largest laterally, all abruptly larger than the minute nape scales
which are distinctly smaller than the dorsals. No parietal eye.
Suboculars weakly keeled, separated from the supralabials by one
row of scales. Seven to eight supralabials to the center of the eye.
Mental divided, each half a little wider than long. Two small
scales posteriorly in the notch between the mentals. Two much
larger rectangular scales lateral to them, between the trapezoidal
first sublabials. Three sublabials in contact with the infralabials on
each side.
Throat scales swollen, rather elongate, minute posteriorly,
becoming larger anteriorly and laterally.
1984 ANOLIS PROPINQUUS. Ê\^ SPECIES 5
Trunk. Dorsal scales granular, convex, subequal. Ventrals not

much larger, swollen, smooth, juxtaposed or subimbricate, in
transverse rows. An umbilical scar still detectable.
Dewlap. Retracted in the unique type, not readily visible. Large,

scales crowded, swollen, imbricate, raised into series of rows or
ridges, a little smaller than the ventrals.

Limbs and Digits. Anterior arm and leg scales unicarinate
except multicarinate at the knee. Supradigital scales multicarinate.
Ca. 25 lamellae under phalanges ii and iii of fourth toe.
Tail. Weakly compressed. No dorsal crest. One row of keeled
scales middorsally, not differentiated from those lateral to it. No
evident verticils.
Enlarged postanals present (5).
Size. The snout-vent length of the unique type is 41 mm.
Color. As preserved, the specimen shows no pattern: it is dark
above, light below and the dewlap skin appears to be dark. W. E.
Duellman provides notes of color in life: "Dorsum green. Venter
pale bluish green. Dewlap pale blue. Iris dark brown. Mouth bluish
black, tongue pink."
Habitat. The only notes on habitat or ecology are Duellman's:
"Sleeping on herb ca. 0.5 m above ground at night."
Comparisons. The most pertinent comparison of A. propinquus
is with sympatric and just-described A. calimae (Ayala et al., 1983)
(Table 1). The unique type is a near hatchling, but the dewlap is
already well indicated, extending posterior to the axilla and the high
number of lamellae under the fourth toe (25) indicates a species of at
least moderate size.
It again astonishing to find the interparietal reduced (i.e., a

is
parietal eye lacking). In contrast to A. calimae, the supraciliary

scales are unusually well developed and the head scales very
distinctly keeled. The blue dewlap is also distinctive.
ACKNOWLEDGMENTS
I am grateful to William E. Duellman, Curator at the Museum of
Natural History, University of Kansas for the opportunity to
examine and describe this peculiar animal. The drawings are by
Laszlo Meszoly,
BREVIORA No. 477
Table 1. Comparison of Anolis calimae and A. propinquus.

1984 ANOLIS PROPINQUUS. NEW SPECIES
Table 1. Comparison of Anolis calimae and A. propinquus.

calimae propmquus
scales posterior
:)^
'^^S. COMP. 200L
MAR381985
B R E V iWn A
useum of Comparsitive Zoology
us ISSN 0006-9698
Cambridge, Mass. 7 September 1984 Number 478
NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA.

III. TWO NEW SEMIAQUATIC ANOLES
FROM ANTIOQUIA AND CH0C6, COLOMBIA.

Ernest E. Williams'
Abstract. Two new semiaquatic anoles from Colombia, partly sympatric, and
sometimes syntopic, prove to belong to distinct lineages despite convergence in habits
— —
and habitat. The larger of the two A. maculigula, new species belongs to the alpha
section of the genus Anolis and ihe eulaemus subgroup of the A. aequatorialis species
group. It is confined, so far as known, to the Departamento Antioquia and

apparently to larger streams. The smaller species, A. rivalis. new species, is a beta
anole of the lionotus species group and tends to prefer smaller streams and extends
beyond the known range of A. maculigula in Antioquia and in Choco.
In 1968 Norman J. Scott, collecting on a tributary of the Rio

Arquia western Antioquia, Colombia, obtained a large anole of
in
aquatic habits which he thought resembled A. aquaticus. It proves,

however, to be a new species belonging to the alpha section of the
genus. Syntopic with this species was a smaller form which Scott
recognized as a member of the lionotus species group (beta section).
This also proves to be new. These specimens were deposited in the
collections of the Los Angeles County Museum (LACM).
Since these first collections, additional material of both species
has been obtained in another area of western Antioquia some 70 km
farther north by Juan Manuel Renjifo and Vladimir Corredor
(material in the Instituto de Ciencias Naturales, Bogota [ICN]).
Specimens of the larger species have been collected at Urrao,
Parque Las Orquideas, ca. 50 to 60 km east of the type locality, by

02138.
BREVIORA No. 478
Horatio Echeverri for Marco Serna (collection of Colegio San Jose,

Medellin (CSJ). Specimens of the smaller species have been
collected in theChoc6 independently by Philip Silverstone (MCZ)
and Charles Myers (ICN). A further Choc6 specimen of the smaller
species has been found in the collections of the San Diego Natural
History Society (SDSNH) collected by C. B. Perkins.
The
larger new anole may be named, with reference to its heavily
spotted throat:
Anolis maculigula, new species

(Figs. 1-3)
Holotype: LACM 42150, adult male.

Type locality: Quebrada San Lorenzo, tributary of the Rio
Arquia near the small town of Belen (6° 15'N 76° 39'W), about 10 to
15 km upstream from the junction of the Rio Arquia with the Rio
Atrato, western Antioquia, Colombia, N. J. Scott coll., 17 April
1968.
Paratypes: Antioquia: LACM 42142, 42144-49, 42151-54, same

data as the type; ICN 5917,
Camp Pegadorcito (Ingeominas), (about
6° 42'N 76° 27'W,) on the height of the Cordillera Occidental
between Frontino on the south and Dabeiba on the north, Rio
Figure 1. Anolis maculigula, new species. Type, LACM 42150. Dorsal aspect of
head.
1984 TWO NEW SEMIAQUATIC ANGLES
o
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2
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o
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BREVIORA No. 478
Figure 3. A. maculigula, new species. Type, LACM 42150. Throat pattern.
Amparracl6, 805 m elevation, Juan M. Renjifo and Vladimir

Corredor coll., 16 September 1981; ICN 5918, at the first stream
beyond Pegadorcito, J. M. Renjifo and V. Corredor coll., 14
September 1981; CSJ 308-309, Urrao, Parque Las Orquideas,
Horatio Echeverri coll., 21 May 1981; CSJ 431, 445, 447, Urrao
(Rio Calles), Marco A. Serna and Horatio Echeverri coll., 10-12
July 1983.
1984 TWO NEW SEMIAQUATIC ANGLES 5
Diagnosis. An alpha anole of moderate size related to A.

eulaemus Boulenger and, like the latter, having small head scales
and narrow digital dilations, but differing in having fewer scales
between the supraorbital semicircles and between the semicircles
and the interparietal, differing also in color, habits and habitat.
Description. Head. Head scales small, pustulose, swollen,
keeled. Twelve to 19 scales between the second canthals. Eight to 1 1
scales bordering rostral posteriorly. Anterior nasal separated from
rostral by one or two scales.
Two to three scales between supraorbital semicircles. A few
supraocular scales distinctly enlarged, no well-defined supraocular
disk. One to two elongate supraciliaries on each side followed by
granules. Canthus distinct, seven canthal scales, the second longest.

Eight to 11 loreal rows, scales small, subequal.
Temporal and supratemporal scales granular. A very indistinct

double row of enlarged granules at margin between temporal and
supratemporal areas. Scales around interparietal enlarged, inter-
parietal larger than or about equal to ear, separated from
supraorbital semicircles by one to three scales on each side or in
contact.
Suboculars separated from supralabials by one row of scales,
posteriorly grading into temporal granules, anteriorly diminishing
gradually in size in front of eye. Seven to ten supralabials to center
of eye.
Mentals wider than deep, in straight line contact with eight to ten
scales between infralabials, grading in size from the ihfralabials
toward the center. Sublabials not clearly differentiated. Central
gular scales small, swollen, even conical, grading laterally into much
larger scales.
Trunk. Two rows slightly enlarged, tricar-
to three middorsal
inate, imbricate, grading into small, keeled but juxtaposed flank
scales, showing small granules between them. Ventral scales equal to
or smaller than the middorsals, smooth, flat or swollen, sub-
imbricate or juxtaposed, sometimes showing small granules.
Dewlap. Large in male, extending onto first third of belly. Edge
scales ca. equal to ventrals. Lateral scales in rows, several scales
across, widely separated by naked skin. Barely indicated in female.
Limbs and Digits. Largest arm and leg scales larger than
ventrals and uni- to multicarinate. Supradigital scales multicarinate.
Sixteen to 22 lamellae under phalanges ii and iii of fourth toe.
6 BREVIORA No. 478
Tail. Distinctly compressed to very strongly compressed, most

strongly in the adult male, a low crest of unicarinate scales, the crest
scales at least 2X those of the lateral rows. The other caudal scales
uni- to multicarinate. No enlarged postanals in male.
Size. The male type measures 98 mm snout to vent. A topotypic
male measures 72 mm, while the largest topotypic female is 73 mm
snout-vent length. The male from Urrao (CSJ 308) is 107 mm
snout
to vent; the unregenerated tail is 215 mm
long. The female (CSJ 309)
from the same locality has 75 mm
snout-vent length.
Color in Life. Normal Scott has provided notes from life for
topotypic specimens. "Adult male: Dorsum with an obscure pattern
of five dark brown blocks separated by lighter gray brown areas and
broken by, on each side, a dorsolateral stripe of the same color.
Dorsal crest paler and green. Tail and legs banded with dark gray
brown and light gray green. On flanks a mixture of olive gray and
orange flecks on a green ground. Side of head a mixture of orange
and gray flecks, lips mottled and with blue flecks. Soles of feet olive
brown. Throat with a series of orange and blue stripes. Chin mottled
with orange, blue and gray, the blue predominant. Iris as in female
but white ring tinged with green. Dewlap color complex: base with
orange stripes on a blue gray ground, anterior third pale bluish rose,
posterior portion white becoming pale blue toward belly. Juveniles
colored much as females but green brighter on head, neck and sides.
Adult female: Dorsum dark gray brown with gray black bar, lighter
areas on neck, head and legs greenish gray. Sides mottled gray, olive
green and black with a few light gray punctations. Side of head
mostly greenish. Venter clear white, throat mottled green and
white, underside of head patterned with white and gray, the chin
with green and gray. Iris dark brown enclosing a narrow white
ring."
Juan Renjifo in a note to Stephen Ayala reports the color in life
of theAmparrado maculigula (translated): "Color green (lichenate)

with dark blotches on the back, the first above the shoulders. A
blotch at the shoulder dark with a white spot in the center."
Color in Preservative. Color now differs substantially from that
reported for the fresh specimens. All blues, pinks and greens are
gone. There are only light and dark grays but the complex pattern is
retained. The dark crossbars of the middorsum contain light spots
and the lower flanks tend The throat is boldly to
to be vermiculate.
weakly vermiculate, less strongly vermiculate in males than in

females, in which the heavily marked throat contrasts strikingly

with the light and unmarked belly. The skin of the throat fan is gray.
Habitat and Habits. The type locality, Quebrada San Lorenzo,
is described as a stream about 10 m
wide with occasional waterfalls,
with a moderately steep gradient and huge boulders. A. maculigula
was most common where there were 3 to 4 m diameter moss-covered
boulders at the head of pools, when first seen most were on the
upstream vertical face of the boulders spray zone and tried to
in the
escape under the overhanging edge of the boulders. In Quebrada

San Lorenzo the smaller (lionotus group) species also occurred but
was less common. (See further below.) No comparable details are
available for the habitat of ICN 5917 and 5918, but the latter
specimen is reported as on rocks in a stream.
Comparisons. A. maculigula is clearly an alpha anole; a caudal
vertebra dissected from a broken tail shows no trace of transverse
processes. It is referred to the aequatorialis species group on the
basis of very small head scales, large size and narrow digital
dilations. Since its toe pads are "raised," i.e., overlap distally the
proximal scales under the first phalanx, it is further referred to the
eulaemus subgroup in contrast to species with pads "not raised,"
i.e.,not overlapping but continuous with the scales of the first
phalanx. (The latter are the aequatorialis subgroup.)

Within the eulaemus subgroup, in which there are a confusing
number of undescribed populations, A. maculigula is distinctive in
its pustulose head scales, its small ventrals, usually smaller than the
small dorsals, and in the relatively large size of the interparietal

which is almost always larger than the ear, in the very compressed
tail and in a distinctive coloration. It may well be unique among the
known eulaemoid forms in its semiaquatic habits also. Certainly no

other member of the group shows so compressed a tail or
maculigula's tendency to a lined pattern.
The second and smaller species may very appropriately receive
the Latin name that means "user of the same stream":
Anolis rivalis
(Figs. 4-7, 12, 14)
Holotype: LACM 42124, adult male.

Type locality: Helen, Rio Arquia, Antioquia, Colombia (6°
I5'N, 76° 39'W), Norman J. Scott coll., 17 April 1968.

8 BREVIORA No. 478
Paratypes: Antid(\\x\2i: LACM 42128-133, 42135, 42137-139.

MCZ 115720-722 from the type locality, Norman J. Scott coll.;
LACM 42125-127, 42141, 45002-07, 51540, Finca Los Llanos, Rio
Arquia, Philip Silverstone coll., 1968; ICN 5912, Camp Pegadorcito
(Ingeominas), 45 minutes by helicopter from Medellin, on the height
of the Cordillera Occidental between the towns of Frontino to the
south and Dabeiba to the north, Rio Amparrad6, 805 m elevation,
(about 6° 42'N, 76° 27'W,) Juan M. Renjifo and Vladimir Corredor
coll., 13 September 1981; ICN 5913, Filo Amparrad6, the same area
and collectors, 16 September 1981; ICN 5914, Camp Chontaduro,
same area and collectors, 9-12 September 1981; ICN 5915, same
area and collectors, 14 September 1981. Choco: LACM 72766,
72772, MCZ
100353, Alto de Buey, P. Silverstone coll., 1968; ICN
4053, Quebrada Mutate, 200 m, northern base of Alto de Buey, C.
W. Myers, John Daly and Michael G. A. Hill coll., 18-24 October
1978; SDSNH31163, "Port Utria," = Puerto Utria (6° 02'N, 76°
23'W), C. B. Perkins coll., 25 February 1938.
Diagnosis. A member of the lionotus group of beta anoles,
differing from the remainder of the group in the combination of
small but not minute head scales (13 to 18 across snout between
second canthals), one to two rows of scales between the supraorbital
semicircles, interparietal in contact with the semicircles or separated
by no more than two rows of scales,and by a zone of moderately
large, indistinctly keeled, flat scales on the middorsum in 11 to 18
rows.
Description. Head. Scales small, uni- to multicarinate. Thirteen
to 18 scales across snout between second canthals. Seven to nine
scales border rostral posteriorly. Anterior nasal scale separated
from by one scale. Nine scales between supranasals. Scales in
rostral
posterior portion of frontal depression smaller than those anteriorly

placed.
Supraorbital semicircles separated medially by one to three
scales, in contact laterally with the largest scales of the supraocular
disks which consist of a variable number of wrinkled or keeled
scales, a few of which may be much larger than the others. One to
three elongate supraciliaries continued posteriorly by a series of
smaller scales. Canthus distinct, canthals 9. second largest. Six to 10
loreal rows,lowermost largest.
Temporals small and flat, not granular. A distinct double
supratemporal row, dorsad of which nearly granular scales grade
1984 TWO NEW SEMIAQUATIC ANGLES
Figure 4. A. rivalis, new species. Type, LAGM 42124. Dorsal aspect of head.
Figure 5. A. rivalis. new species. Type, LACM 42124. Lateral aspect of head.
into the flat scales surrounding the large interparietal, usually larger
than ear and in contact with the semicircles or separated by one to
two scales. Scales posterior to interparietal not different in size from
middorsals but grading into smaller nape scales which then grade
posteriorly into the dorsals. Suboculars weakly keeled, narrowly in
contact with supralabials or separated by one scale row, grading
10 BREVIORA No. 478
Figure 6. A. rivalis, new species. Type, LACM 42124. Ventral view of chin.
anteriorly into loreals, posteriorly more sharply distinct from the

temporals. Six to nine supralabials to the center of the eye.
Mental divided, wider than long, in contact with six or seven
scales between infralabials. Gular scales smallest medially, quadrate,
swollen.
Trunk. Middorsal scales flat, hexagonal, wrinkled, in ca. 1 1 to
17 rows, grading quite gradually into subgranular flank scales,
larger in the center of the body than on the nape, becoming slightly
smaller again in the sacral region and on the base of the tail.
Ventrals smaller than dorsals, keeled, imbricate.
Dewlap. Moderate, extending onto first third of belly, lateral
scales weak, in densely packed rows, edge scales keeled, slightly
larger than ventrals.
Limbs and Digits. Scales on limbs strongly unicarinate except
at knees and elbows where they are multicarinate. Supradigital
scales multicarinate. Twelve to 18 scales under phalanges ii and iii of
fourth toe.
1984 TWO NEW SEMIAQUATIC ANGLES u
£ .5
c
BREVIORA No. 478
12
Figure 8. A. oxylophus. KU 34262. Frontal and middorsal scales.
•I r^ *»1 »«>4 wv*iAAr\re>a\ cnCt\/^K.

Figure 10. A. macrolepis, MCZ 133000. Frontal and middorsal scales.
c:r....-o II A r,^^îl^r>,,c If II M'X')AQ Frnntnl nnH miHrlnr«al scales

14 BREVIORA No. 478
Figure 12. A. rivalis, LACM 42129. Frontal and middorsal scales.
Tail. Compressed without dorsal crest. Verticils indistinct.
Enlarged postanals absent.

Color as Preserved. Brown with darker flanks. Middorsum
banded. Two narrow light lines on
one above shoulder, one
sides,
starting at axilla. Spotting on lower flanks. Limbs banded. Belly
and throat white or very weakly spotted. Dewlap white. Nape white
to above level of ear where a darker margin sets it off against the
dorsal light brown.
Size. The type male is 62 mm in snout-vent length. MCZ 1 15722
and LACM 42138, topotypic males, are each 64 mm
snout to vent.
Color in Life. Scott provides color notes on the topotypic
specimens: "Adult male: Dorsum dark brown with thin green brown
vertical bands on flanks. Several stripes from axilla to groin, the
lower ones heavily suffused with red. Light areas on sides dusty
rose. Venter yellowish white with heavy red suffusion on sides of
abdomen, throat and underside of head. Iris dark chestnut enclosing

a narrow yellow ring. Dewlap solid orange. Adult female: Dorsum
Figure 13. A. rivalis from Quebrada Mutati, 200 m, northern base of Alto del
Buey, Choc6, Colombia. Photo by C. W. Myers.
as in male but with only a faint hint of red suffusion. Light areas of
head white becoming green on snout. Venter yellowish white.
Dewlap area with pale orange spot, throat anteriorly white. Iris as in
male."
Notes by C. W. Myers on the specimen obtained from Quebrada
Mutate record the following colors: "Brown, changeable to
brownish green, with lateral line and lower side of neck dirty white.
Throat gray, dewlap light orange, venter greenish gray. Iris brown.
Tongue light gray."
Juan Manuel Renjifo has supplied color notes on ICN 5912 from
the Rio Amparrad6 region: Greenish brown with bars of lighter
greenish brown forming chevrons on the middle of the back. Labials

and flanks rosy salmon. Venter yellowish cream. Gular sac white.
One entry in Scott's field notes cites an unspecified rivalis from
Belen as having an orange dewlap. There would appear to be a
range of dewlap colors in this species.
Habitat and Habits. At Quebrada San Lorenzo, Scott reports
A. rivalis both to be less common and to occur on small rocks near
the bottom of pools rather than on the huge moss covered boulders
preferred by A. maculigula. They escaped by running from rock to

rock over the water surface. Some were taken at night sleeping on
16 BREVIORA No. 478
vegetation overhanging the stream. He specifically mentions (letter

of February 28, 1979): "Quebrada San Lorenzo was the only place
where I
got both aquatic species. Nearby Quebrada Barrero had less
water and seemed to have only the "lionotus" [= rivalis] type."
The single specimen of A. rivalis obtained by the Myers group in
1978 was again found in a quebrada, but the notes do not cite any
larger aquatic species.
Renjifo in his field notes does not mention close or co-occurrence
of the two species and confirms the semi-aquatic habits of rivalis in
the Amparrado region. He cites ICN 5912 as (translated) "in forest
on trunk above the stream, meter above ground level;" ICN 5914
'/2
"in stream on a stone;" ICN 5915 "sleeping on rocks in the stream
alongside the current;" and ICN 5918 "in stream.... on rocks."

Comparisons. A. rivalis is a member of the beta series and of the
lionotus species group.
The latter is a series of taxa (the other referred species: oxylophus
Cope, 1875, lionotus Cope, \%()\, poecilopus Cope, 1862, macro-
lepis Boulenger 1911) that extend from Nicaragua to Ecuador,
replacing one another with minimal or no overlap, so far as known.
A sixth taxon belonging to this group occurs in northwestern
Ecuador and southwestern Colombia. It will be described by
Kenneth Miyata. All are characterized ecologically by "aquatic"
habit and riparian habitat and (usually) by a zone of enlarged dorsal
scales, more or less similar in size over an area of 14 to 20 rows, then
grading laterally into the flank scales. They have also small keeled
ventrals and a more or less well developed flank stripe.
The species of this group ring thechanges on just a few
morphological characters: the size of the head scales, the number of
scale rows between the supraorbital semicircles, the number of
scales between the interparietal and the semicircles, the size of
supratemporal and nape scales, and the size and also the keeling or
lack of keeling of the middorsals. Each of these characters varies
independently, and adjacent species tend to be sharply distinct in
one or more features.
No revision of the lionotus species group exists nor has South
American macrolepis been recognized as a member of it until
recently (Williams, 1976).
Boulenger (191 describing mac ro/ep/^ did, indeed, associate it
1) in
with poecilopus but also suggested a relationship to notopholis

{humilis species group), to which it is only superficially similar.
Figure 14. Map: Localities for semiaquatic anoles in Colombia and adjacent
Panama.
BREVIORA No. 478
18
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20 BREVIORA No. 478
Taylor (1956) in his discussion of the lizards of Costa Rica called

attention todiscrepancies between the type description of
the
lionotus Cope, 1861 (with a type locality in Panama) and that of
oxylophus Cope, 1875 (without exact locality but presumably from
Costa Rica), which was supposedly a synonym. However, lacking
Panamanian specimens, he followed then and current usage in
applying the name lionotus to Costa Rican animals. My own
examination of most of the material available in American
museums shows significant differences between eastern Panamanian
and western Panamanian and Costa Rican-Nicaraguan specimens.
Campbell (1973) noted the character of true lionotus that has most
impressed me: dorsal scales about twice the size of the ventrals. In
oxylophus (as in poecilopus) the dorsals and ventrals are about
equal but xnpoecilopus the dorsals are strongly keeled, in oxylophus
smooth or weakly keeled. Table reports the characters dis-
1
tinguishing members of this group.

The new species is distinguished from all other members of the
lionotus group by a combination of characters: small but not minute
head scales, few scales between the supraorbital semicircles, small to
moderate supratemporal scales, nape scales distinctly smaller than
the enlarged dorsals which are large, larger than the ventrals,
wrinkled or weakly keeled and weakly imbricate. From poecilopus
it is distinguished by its much larger middorsals, fewer rows between
the supraorbital semicircles and its larger circum-interparietal

scales. From macrolepis it is distinguished by its somewhat larger
middorsals, the nape scales distinctly smaller than the large
middorsals (rather than subequal) and by smaller head scales.
Discussion. The sympatric occurrence of two semiaquatic
anoles is not unique: within the lionotus group there is the example
of the local sympatry oi poecilopus and lionotus in the vicinity of

the Panama Canal (Campbell, 1973). In this case the overlap zone is
suspected to be narrow, but knowledge of the distributions is not

adequate to demonstrate this. In the case of poecilopus and
lionotus, the two overlapping species are very different within the
group in dorsal and head scale
size. (In general, in the lionotus
group, although the species are primarily allopatric, the differences

summarized in Table 1 are greater in adjacent than in distant taxa.)
The two sympatric semiaquatic species here described are quite
different in their affinities, belonging to different sections of the
genus Anolis. They are adequately different in scale characters but

not quite so strikingly as the most different species pairs within the
lionotus group. The similarities in habits, in the pronounced flank
stripes, and in the strongly compressed tails, are indeed more
impressive than their differences.
But these similarities are parallels only and are seen in other
semiaquatic anoles not at all closely related, as Schwartz has
demonstrated taxonomically quite isolated semi-
in describing a
aquatic anole from Hispaniola (Schwartz, 1978). The semiaquatic

anoles, except for the allo-parapatric series that constitutes the
lionotus group, are not at all a lineage, but only an ecomorph in the
sense of Williams (1972, 1983).
ACKNOWLEDGMENTS
I am indebted to the curators who have provided me the
opportunity to describe the new species: John Wright and Robert
Bezy of the Los Angeles County Museum, Pedro Ruiz of the
Instituto de Ciencias Naturales, Bogota, Colombia and Gregory
Pregill of theSan Diego Natural History Society. Norman Scott
and Charles Myers have allowed me to quote from their field notes,
and Pedro Ruiz has transmitted the field localities and observations
of Juan Renjifo and Vladimir Corredor. Stephen Ayala has
commented on the manuscript and provided the map which
compares the distributions of Colombian aquatic anoles. Laszlo
Meszoly has done the illustrations. The photograph of A. rivalis was
provided by Charles Myers.
LITERATURE CITED
BouLENGER, G. A. 191 1. Description of new reptiles from the Andes of South
America, preserved in the British Museum. Ann. Mag. Nat. Hist. ser. 8, 8:
19-25.
Campbell, H. W. 1973. Ecological observations on Anolis lionotus and A.
poecilopus (Reptilia, Sauria) in Panama. Amer. Mus. Novitates, No. 2516, pp.
1-29.
Cope, E. D. 1861. Notes and descriptions of anoles. Proc. Acad. Nat. Sci.
Phila., 1861: 209 215.

1862. Contributions to Neotropical saurology. Proc. Acad. Nat. Sci.
Phila., 1862: 159-188.

1875, Gn the Batrachia and Reptilia of Costa Rica. J. Acad. Nat. Sci.,
8: 93-157.
Schwartz, A. 1978. A new species of aquatic anole (Sauria, Iguanidae) from
Hispaniola. Ann. Carnegie Mus., 47: 261-279.
22 BREVIORA No. 478
Taylor, E. 1956. A review of the lizards of Costa Rica. Univ. Kansas Sci.
Bull., 38: 3-222.
Williams, E. E. 1972. The origin of faunas. Evolution of lizard congeners in a
complex island fauna: a trial analysis. Evolutionary Biology, 6: 47-89.
1976. South American anoles: the species groups. Pap. Avuls. Zool. S.
Paulo, 29: 259-268.
1983. Ecomorphs, faunas, island size and diverse end points in island
radiations of Anolis pp. 326-370, 481-484. In Huey, R., E. Pianka, and T.
Schoener (eds.). Lizard Ecology - Studies of a Model Organism, Harvard
University Press, Cambridge, Mass.
,- ^^ .-sw^-/--.'
r' /
B R E V I ff^H A
useum of Comparative Zoology
us ISSN 0006-9698
Cambridge. Mass 7 Sfptember 1984 Nimber 479
AGONISTIC AND COURTSHIP DISPLAYS OF

MALE ANOLIS SAGREI
Michelle P. Scott'
Abstract. Male Anolis sagrei perform a dewlap fanning display and four different
bobbing displays in agonistic and courtship encounters. Only one of the bobbing
displays has a species-specific, stereotyped pattern. The total number of bobs and the
number of bobs in each unit of this display and the use of the dewlap showed
considerable variability both within and between individuals.The frequency
distribution of the type of display used in aggressive encounters showed more inter-
than intra-individual variability. Dewlap fanning displays were used at a relatively
higher frequency before crests were raised, but the dewlap was used with the bobbing
display at a relatively higher frequency by males after the erection of crests.
Submissive animals displayed less frequently overall and were more apt to use a
dewlap fanning display, but they were less apt to use the dewlap with a bobbing
display than a dominant male. Male A. sagrei were less apt to use the species-specific
stereotyped pattern in courtship than in agressive encounters. The "jiggle" bob
display immediately preceeded half of the attempted matings. A dewlap fanning with
erect posture was associated with the termination of a copulation.
INTRODUCTION
The brightly colored dewlaps and stereotyped bobbing displays of
males of the iguanid lizard genus Anolis may communicate
information regarding species, sex, reproductive state, dominance
and territorial status, intentions, and level of arousal. Dewlap color
alone sometimes does not appear to be an important element in
species recognition and female choice for the solitary anole A.
carolinensis (Greenberg and Noble, 1944; Crews, 1975a). However,
in a complex Anolis fauna, color and dewlap proportions may

02138.
2 BRFVIORA No. 479
provide redundant information for species recognition (Williams

and Rand, 1977). Jenssen's work (1970a) with Anolis nebulosus
indicates that the male's species-specific bobbing display does
indeed play a significant role in sexual selection and female choice.
In examining anoline displays, recent researchers have focused on
the degree of stereotypy (the amount and possible significance of
variability within and among individuals) and the complexity of
display repertoires of different species. Displays may vary either in
temporal patterns or by the presence or absence of various
components within the display. Species differ considerably in the
amount of stereotypy of their displays, and the source of variability
may be intra-individual or between individuals (Jenssen 1978).
Anolis carolinensis (Crews, 1975b), A. aeneus (Stamps and Barlow,
1973), and A. nebulosus (Jenssen, 1971) have very stereotyped
displays. Each individual performs a consistent display, but there is
variability between individuals. Different populations of A. nebu-
losus have been examined and still further variability between
populations was found. Other species, however, like A. limifrons
(Jenssen and Hover, 1976) show greater intra-individual variability
than inter-individual differences. At the extreme, almost all (97%) of
the variability of A. townsendi displays is attributable to intra-
individual variation (Jenssen and Rothblum, 1977). Anolis opal-
inus, on the other hand, was found to be almost completely lacking
stereotypy at any level, individual or population, in that there was

no single discrete display pattern. However, 75% of the analyzed
displays fit a theme which explained the highly variable bob number
and display cadence (Jenssen, 1979).
The complexity of the display repertoire of a species is another
important feature. Researchers (e.g.. Carpenter and Grubitz, 1961)
first described only two basic types of displays, a courtship pattern
of rapid head bobs and a species-unique "signature" pattern. The

lattertype was originally labeled an "assertion" display that, with
modifiers, blends into a "challenge" display (Carpenter, 1967;
Jenssen, 1970b, 1977; Crews, 1975b). Typically, if other display
types were described, they consisted of the signature display plus
modifiers (e.g., four-legged push-up posture preceding dewlap
fanning), or they may be characterized by a different coordination

of the dewlap extensions during the head bob sequence. Recently,
researchers have found greater complexity in the display repertoire
of some anoles (Hover and Jenssen, 1976; Jenssen and Rothblum,
1984 DISPLAYS OF ANOUS SAGREI 3
1977). Anolis aeneus, A. limifrons, and A. townsendi, for example,

have several clearly distinct display types which differ not just by the
modifiers or the context in which they appear, but also by different
bob patterns.
This paper described the display repertoire of male Anolis sagrei
in agonistic and in courtship situations. A description and analysis
of the species-specific stereotyped display and its variable elements
are given. An attempt is made to analyze these displays and their
modifiers in terms of the social context in which they appear and the
effect they have on conspecifics. The possible role of the species-
specific signals used in sexual selection is discussed.

Anolis sagrei has a wide distribution, occurring on Cuba,
Jamaica, the Bahamas, LittleCayman, Cayman Brae, Swan Island,
and coastal areas of Mexico and Belize, and has recently been
introduced into Florida (Williams, 1976). It occupies a relatively
open habitat.During the breeding season, males ond females are
strongly territorial. A male territory has been described by Evans
(1936) as at least 50 m- and containing up to three females. In
cultivated areas, it is typical for males to occupy a hedge row or
fence at approximately 6 m intervals (Evans, 1936; Scott, personal
observation).
Twenty males (55-62 mm snout to vent length), 12 females and 8
juveniles were collected near the Universtiy of South Florida,

Tampa, in May 1979 when the mature males and females were
sexually active.
The were held in experimental cages of two designs. In
lizards
Design 1, one large male, a juvenile male and two to three females
inhabited an aquarium 75 cm long X 32 cm deep X 45 cm high. Two
of these aquaria so arranged were placed end to end with a
removable opaque divider between them. Each contained a live
plant and a prominent perching branch. The far side of each
aquarium was covered by bark which was also favored as a perching
site. Thus, when the opaque divider was removed, the two territorial
males could see each other but were physically separated by a

5 pairs of 10 different males
double pane of glass. In this design,
were tested against each other three to five times. In Design 2, a wire
cage 47 cm long X 51 cm deep X 39 cm high was divided in half

4 BREVIORA No. 479
longitudinally by a removable opaque partition. A single large male

was housed in each which also contained a plant, a perch, and
half,
a bark wall. In these cages, the animals were in physical as well as
visual contact with each other when the partition was removed. In
this design, three pairs of six different males were effectively tested
against each other only once since subsequent tests would have been
declared "no contest" at the outset by the participants. These wire
cages were kept in Sherer-Gillet environmental chambers with
transparent doors. All animals experienced a constant environ-
mental regime of 14L:10D photoperiod and a corresponding
temperature cycle of 32:23° C. All animals received food (meal-
worms and crickets) and water ad lib. Observation of the aquaria
animals in Design was made from behind a cardboard blind to
1
ensure that the displays were directed at conspecifics and not the
observer. The animals in the lighted environmental chambers of
Design 2 were observed in a darkened room, making the observer
relatively inconspicuous. The cages were arranged in each case so
that no animal could see a conspecific except those in its
experimental setup. When the partition was in place males were

only occasionally observed displaying at the juveniles or females.
Males were housed individually before they were placed in the
experimental cages.
All males were allowed at least one week to acclimate to a new
cage before testing was begun. Agonistic tests were at least 15

minutes long and continued 5 minutes after the situation appeared
stable (that is, when there was no change in positions and display
types used by the two animals). Courtship tests were terminated

when the animals finished copulating.
Male agonistic behavior was studied in both types of cage designs
with the opaque dividers removed. Males in the aquaria of Design 1
were repeatedly tested against each other whereas those in the wire
cages of Design 2, which allowed physical contact, only met once
because, once beaten, the subordinate animal did not display but
retreated. Courtship behavior was observed only in the cages of
Design 2; the partitions were left intact and a female was introduced
into a wire cage which housed a single male.
Parameters of male behavior were recorded with a 10 channel
EsterlineAngus operation recorder Model A. Displays were taped
with a Sony video camera AVC 3210 with a 75 mm fl2.5 lens and
analyzed with a Sony videorecorder AV 3650 with which the tapes
1984 DISPLAYS OF /I A'0Z./5' 5/1 G^£/ 5
could be slowed to 8% actual speed. The head and dewlap

movements were transcribed onto a strip of acetate by running the
known speed, using
acetate in front of the video screen at a constant
the gears of the event recorder and following the action with a felt
tipped pen placed on the rostrum of the videotaped lizard. The time
framework could be superimposed on the tracing making correc-
tions for the video recorder speed and the gear ratio of the event
recorder. Only animals from the wire cages of Design 2 were
videotaped because the T.V. equipment was less conspicuous to the

subjects.
A total of 620 agonistic displays by 12 different males were
observed in 18.8 hours. Two hundred
twenty-nine displays of eight
different courting males were observed in 12.4 hours. Thirty-one
displays of three males in two entire agonistic events and three

different males in entire courtship tests were taped and analyzed.
Recording data on the video equipment and event recorder
provided different benefits. When
using the event recorder, the
social context of the display and the response of the recipient could
easily be included. In this way the meaning of the display could be

defined by the response it provoked. With the video equipment it
was possible to record data on the modifiers of the displays in fine
enough detail to compare and discuss variability within and between
individuals. The cadence of bobs could also be recorded to
discriminate the type of bobbing display and to relate it to the social
context.
For purposes of discriminating dominant and submissive be-
havior, the animals of all the tests in both cage designs were divided
into two groups on the basis of their behavior at the end of the test.
An animal was judged to have been acting dominant throughout the
test if he was still facing his opponent at the end with an alert
posture. Those anoles that ran from their opponent to the far side of
the cage and had a posture low to the substrate were classed as
submissive for the whole test. Since these classifications were
somewhat subjective, borderline cases in which either animal
showed a mixture of dominant and submissive behavior at the end
of the test were not used in the following analysis. Within each
group there was little variance in the frequency of each type of
display performed (Table 1); therefore, for statistical purposes, data
from all dominant males were pooled, as were those from all
subordinate males.
BREVIORA No. 479
Table 1
Frequency ± variance of displays by type per minute of animals judged
.
at the end of the test to be dominant or subordinate.
Dominant animals Subordinant animals

(N = 34) (N = 11) Difference
Dewlap only 0.07 ± 0.01 0.07 ± 0.01 n.s.
Bob only 0.43 ± 0.06 0.21 ± 0.04 t

= 2.70, p < 0.01
Bob + dewlap 0.16 ±0.02 0.01 ± 0.00 t
= 3.49, p < 0.001
Any encounter escalated (if it were going to) in a predictable

fashion. Modifiers were added to displays and other behaviors
appeared in the same order each time, as the states of arousal of the
participants increased and became more tense. To
their bodies
facilitate analysis and
of behaviorsresponses in agonistic en-
counters, confrontations were divided into three stages which were
differentiated by changes of body posture. In the first stage, a male
had not yet erected either a nucal or dorsal crest or laterally
compressed his body and was said to be in state A. In the second
stage, a male had both crests (state B) and in the third, most aroused
stage, he had laterally compressed his body as well (state C).
Statistical tests usea were t-test of proportions, arcsin trans-
formed, to test the difference between the means of two samples and
a one-way analysis of variance to examine the inter- and intra-
individual variance in display stereotypy. Means are expressed ± the
standard error.
RESULTS AND DISCUSSION

Displays and Modifiers
A display is a sequence of behaviors, usually performed in its
entirety, which communicates information about the sender. In the

following context, it refers to the change in amplitude over time of a
lizard's head and/or dewlap and encompasses any consistantly

associated stereotyped movements or postures. A modifier is a
posture (static modifier) or movement (dynamic modifier) which is
not always present with a particular stereotyped display but may be
added (Jenssen, 1978). Anolis sagrei sometimes performed these
modifying behaviors, particularly postures, separately from stereo-
typed head bob displays.
A brief description of the displays investigated follows. The
frequency distributions of the display types, degree of stereotypy.

1984 DISPLAYS OF /I A'OL/5S/lG/?£/ 7
and information conveyed are all discussed later in the context in

which the display was used.
Dewlap Fanning. The dewlap, which in Florida A. sagrei is
bright orange with a yellow margin, is extended by the hyoid

apparatus. The pattern and context of this action is extremely
variable. It can occur as a single extension or in a fast or slow series
of as many as 15 separate extensions. The series have no discernible
pattern, and the display is used in all observed contexts and may be
directed at a humanobserver as well as a male or female conspecific.
Bobbing. The head can be bobbed from the neck, from mid-
body, or as a four-legged push-up. Bobs from the neck or mid-body
can occur singly or in a series of a single posture, or the display may
begin as bobs from the neck and finish as bobs from the mid-body.
These postures may demonstrate increasing states of arousal in that
order. Bobs with a four-legged push-up posture are not performed
singly. These four-legged push-up bobbing displays are typically
8-10 bobs but may contain 18 to 20 and last as long as 30 seconds.
There are three different sorts of bobbing displays.
(A) Anolis sagrei can perform a species-specific stereotyped
display which has a distinctive cadence to a series of bobs which
may be called its "signature" display (Stamps and Barlow, 1973). It
consists of four units: 2 quick initial bobs of increasing amplitude, a
long bob, a short bob, followed by a series of 3 to 12 even-tempo

bobs dampening in amplitude (Fig. 1). There may be variation in
bob number of any unit of the pattern. If the dewlap is extended
during the display, italways flashes on the third (long) bob and
variably during the dampening series of bobs at the end. This
signature display occurs in an assertion context (i.e., non-directed or
low conflict situation), in both agonistic and courtship tests, and at
every state of arousal.

However, sometimes this bobbing display does not consist of this
characteristic pattern. It may, however, have some similar elements
to the signature pattern such as a long bob accompanied by a
dewlap extension followed by a short bob. These displays also may

or may not be accompanied by dewlap extensions and without them
as a marker, the elements similar to the signature pattern often
could not be identified. For example, in the display illustrated by
Figure the dewlap extension at 5 seconds marks what might be
2,
the long bob followed by the short bob characteristic of the

signature pattern but it is unrecognizable by the pattern of bobs
alone. These displays also may occur in any context.
8 BREVIORA No. 479
T3
_jj\nAnnnrv\-
— a
e
«:
5 y-Y^-^—y \r~"
0)
5 10 15
time (sec)
Figure 1. The signature display of Anolis sagrei.
-vÂY^/^l^'V"'~^^^'Â^vvvvvv-~^
a xaanpxiimi jutagaMiJWPaagtnKiii^^
'^gaKr
'
fT
1;
5 10 15 20 25
time (sec)
Figure 2. A typical example of a display that did not fit the signature pattern.
•a
m
JZ .:,jyv\vv
Hmnwaiiim'iiiiiiniii «B»V*aft:
5 f \.r 1S
time (sec)
Figure 3. The quick rhythmic bobbing display.
(B) The second bobbing display is characterized by a quick

(about 1-2/sec) rhythmic bobbing with no inter-bob pause and
without any dewlap extension (Fig. 3). The amplitude of these bobs
was less than that of the bobs of the signature display, and usually it
was performed with the head and body close to the substrate. In
these tests, 49 of 53 times it was performed by an animal in a
submissive situation. However, it was occasionally (4 times)
1984 DISPLAYS OF /4AÔZ./5 5/4 G^£/ 9
performed by a highly aggressive dominant male in a context when

the opponent had a submissive posture.
(C) The third type of bobbing display is jiggling action of the
head. The amplitude of these bobs was very much less than that of
other bobbing displays and the rate was much faster (8- 10/ sec). In
these tests, all 34 times this display was used was in courtship
situations and was performed by the male usually as he approached
the female to take a neck grip prior to mating. It may be performed
as a separate display or tacked on to the end of another.
Modifiers. Many different behaviors

may be used to modify a
display. Some wereconsistently used in a particular context and
state of arousal and often caused a consistent response in the
recipient and therefore these modifiers may impart specific informa-

tion on the intent and motivation level of the animal displaying. See
Table 2 for a list of modifiers frequently used, their social context,
and state of arousal when used.
Male-Male Encounters
The encounters between two males were staged such that they
would both be expected to act territorially. A total of 12 different
males were used in 26 tests. In Design 1, the two aquaria which each
housed resident males were sufficiently large that the males were
often 150 cm apart when they first saw each other, so that in 45% of
the 25 tests they escalated aggression (see below) and were only
prevented from combat by the glass partitions. When these same
two males confronted each other in future tests, both continued to
act aggressively. The situation in the wire cages of Design 2 was
different. The animals were less than 50 cm apart when they first
saw each other, and there was no physical partition. In every
instance the rank order of the two was quickly apparent when one
male fled without a contest; there was no escalation of aggression.
The physical proximity seemed to be a deterrent to bluffing by the
less aggressive male and did not allow for a slow escalation of
aggressive behavior and evaluation of each other.

The sequence of events in an escalating fight is typical of that
described for other anoles (Crews, 1975b). Behaviors and modifiers
to displays are added in a sequence that is predictable. In a aquaria
of Design 1, males would display towards each other with first a
nuchal crest and then a dorsal crest being erected. The whole body
10 BREVIORA No. 479
Table 2. Modifiers used male A. sagrei and their context. The steps
in displays of
in escalation of an encounter from display to conclusion (combat or neck grip)

first
were consistently ordered (see text). The state of arousal refers to the stage of
escalation.
Modifier
1984 DISPLAYS OF /I A'OZ./^ 5/4 G/?£/ 11
Table 3. The probability of an animal in an agonistic confrontation taking a

more aggressive posture. A, B, and C are the states of arousal, and A — B is the
first state going to the second. An animal in state A has a
probability of a male in the
body posture without nucal or dorsal crests or lateral compression of the body. An
animal in state B has both crests and one in state C has crests and lateral
compression.
ff tests A-B A-C B-C C-B C-A B-A
Dominant
animals 34 0.91 0.09 0.74
Submissive
animals 11 0.45 0.27 0.09 0.18
Twenty-four displays by males clearly acting territorially were

taped and analyzed; all had some recognizable components of the
signature display. There was however a great deal of variability.
Five of seven displays by one male, and four of five by another, were
of the signature pattern. The remaining displays for these two
animals were varied by having either one or three quick initial bobs
instead of two. The third male whose displays were taped had only
three of 12 showing the signature pattern. The number of initial
bobs (2, 3, or 4), slow bobs (1 or 2), and quick bobs (0, 1, or 2)
varied so that for some displays the signature pattern was barely
recognizable. The mean number of bobs in these filmed displays was
11.1 ± 0.4. The mean time elapsed was 17.4 ± 0.8 seconds. The
coefficients of variation for the number of bobs in these variable
units of the signature display in these agonistic encounters are: 24%
for the initial bobs, 32% for the slow bob, 40% for the quick bob,
and 23% for the total number of bobs in the display. Twenty of these
24 taped displays were accompanied by dewlapping, whereas only
23% of the bobbing displays included dewlap extensions in the total
sample of all displays observed. Twenty-two of the taped bobbing
displays were two-legged push-ups with crests erected; in 12 displays
the body was compressed as well, as the state of arousal
laterally
during the confrontation increased. Each of the three animals gave
one or two displays that began as four-legged push-ups, usually
raising the tail on the up stroke of the bob (Fig. 4), and dampened to
two-legged push-ups. Tail lifts were used as a modifier of two-legged
push-ups in conjunction with nuchal and dorsal crests three times
and once with lateral compression as well. One display which
included crests and lateral compression also was accompanied by a
tongue protrusion and a lip smack.
12 BREVIORA No. 479
Figure 4. Aggressive display with a 4-legged push-up, nucal and dorsal crests,
lateral compression of the body, and a tail lift.
Information on the context of each display type was gained by

pooling the data from all the staged encounters in both cage designs
and examining the frequency distributions of display types (dewlap
flashes, bobbing displays, and bobbing displays with dewlap
extensions). Four bobs and dewlap flashes in a continuous sequence
were arbitrarily chosen as the minimum to constitute a display. The
mean proportion of dewlap fanning displays of all displays was 0. 14
±0.02. There was little intra-individual variation in the proportion
of use of this display form from one test to another. In a one-way
analysis of variance, 87% of the variation in the frequency of use of
this display type was between individuals and 13% between tests of a
single individual (F
= 7.28, p < 0.01). This tendency was enhanced
by the fact that each male in the cages of Design 1 was always
matched against the same opponent and most animals responded in
a characteristic fashion in each test. The frequency with which a
bobbing display was accompanied by dewlap extension was
similarly consistent. Eighty-three percent of the variation was
between animals (F = 4.77, p < 0.01). The mean proportion of
bobbing displays with dewlap modifier was 0.23 ± 0.03 (Fig. 5a).
A male was judged to be either submissive or dominant on a test-
by-test basis, but he probably performed as both a dominant and
submissive individual during the test. Most animals that "lost"
ultimately responded with varying intensity to the challenge of
still
the more one. In addition, the relative ranks of

aggressive
submission and dominance were not necessarily consistent even
1984 DISPLAYS OF ANOLIS SAGREI 13
14 BREVIORA No. 479
when the same opponents were repeatedly matched in the separated

aquaria.
Submissive animals displayed (either bobbing or fanning) less
than half as often (0.29/minute, N=ll) as dominant ones (0.63/
minute, N=34) (Table 1). Dewlap fanning formed a significantly
higher proportion of the total displays by submissive animals than
by dominant animals (x
= 0.34 ± 0.08 vs. x = 0.08 ± 0.01; t
= 2.29,
p < 0.05) (Fig. 5b, c). A significantly lower proportion of the bob-
bing displays of submissive animals was accompanied by dewlap
extensions, x = 0.03 ± 0.01, than of dominant animals, x
= 0.30 ±
0.03 (t
= 3.36, p < 0.01). Therefore, submissive animals use their
dewlaps for fanning displays but not to modify bobbing displays. A
significantly higher proportion of the bobbing displays of sub-
missive animals were the rapid bob (Fig. 3), x = 0.25 ± 0.09 for
submissive animals versus x = 0.01 ± 0.01 for dominant animals (t =
2.99, p < 0.01). Dominant animals also took progressively more
aggressive postures and reached a higher arousal state more
frequently than submissive ones did (Table 3).
The pacing of agonistic encounters varies with each fight, even
when the combatants are known to each other. Of the pairs that
always escalated to a full confrontation, usually the same individual
was faster to be aroused. The mean latency time of aggressive
animals for developing nuchal and dorsal crests was 4.6 ± 0.4
minutes (N=38) and for lateral compression was 7.4 ± 0.5 minutes
(N=26). There was a great deal of intra-individual variation in the
kind of displays, if any, that were performed before both crests were
erected and the body compressed. But the frequency distribution of
display types performed in this low arousal state was different for
animals that would be judged dominant or submissive at the end of
the test, i.e., there were behavioral differences in the two groups
from the beginning of the test (Fig. 6).
A display by one male A. sagrei often prompted a display by the
other in these tests. Forty percent of all displays seen occurred
within 10 seconds after a display or single bob or dewlap flash by the
opponent, but there was no particular tendency to use the same sort
in answer as the one just seen. For example, a bobbing display was
followed by dewlap fanning display or a bobbing display. Forty to
50% of the fanning displays and the bobbing displays with and
without dewlap modifier of both dominant and submissive animals
was in response to the opponents display. There was a slightly
1984 DISPLAYS OF A NOUS SAGREI 15
dewlap bob + bob

only dewlap only
a. dominant
males
TO
submissive
16 BREVIORA No. 479
Table 4. Proportion and number of each type of display for submissive and
dominant animals which were in response to a display by the opponent rather than
spontaneously performed.
1984 DISPLAYS OF /I A'0Z./5 5/1 G/?£/ 17
Of the seven video taped bobbing displays in courtship sequences

of three males, only one was the signature pattern. The pattern of
the rest could not be characterized. All four displays in the two
courtships that ended in neck grips were four-legged push-ups with
the tail lifted as the head bobbed up and the dewlap pulsed. This tail
lift was a very common (28%) modifier in all courtship bobbing
displays.
The overall frequency of all types of displays in courtship
sequences by 8 differentmales in 33 tests was 0.35/ minute which is
about half that of a dominant male in an agonistic situation
(0.66/ minute). As was the case of the agonistic tests, the data can be
combined because variance of display frequency between per-
formances in different tests was relatively low (0.05). The frequency
of bobbing displays with dewlap flashes were similar in premating
and postmating displays (0.32 vs. 0.21) which was similar to that in
agonistic encounters (0.23). There was both inter- (57%) and intra-
(43%) individual variation in the use of the
dewlap (F 1.3, p >
=
0.20). The proportion of displays which were dewlap fanning
displays were similar in courtship (0.15) and agonistic situations
(0.14), but a higherproportion of the displays after mating (0.50)
than before (0.10) were dewlap fanning (Fig. 7). Six of the seven
complete matings, and none of the short matings, were followed by
a dewlap fanning display, most of which had a distinctive erect
posture (Fig. 8). But bobbing displays and further attempts to
regain a neck grip followed one of the incomplete matings. Thus,
bobbing displays may be predominantly a premating signal, and in
this context fanning displays may be a signal of a successful
copulation.
CONCLUSIONS
The male Anolis sagrei of this study show variability in both
pattern and contextual use of displays. Anolis sagrei has only one
species specific stereotypic pattern which may be used in every
context. This signature display when used by males in the agonistic
encounters was quite variable both in the total number of bobs and
the number of bobs in each unit. In addition, the accompaniment of
the dewlap extension was also variable; although, if it was used, it
was always associated with one particular unit of the display, the
BREVIORA No. 479
18
dewlap bob + bob

only dewlap only
a. total
1984 DISPLAYS OF ANOLIS SAGREI 19
Figure 8. Typical post-copulatory display of dewlap fanning and erect posture.
long bob. This display is so variable that sometimes the signature

pattern cannot be recognized. Video taped agonistic and courtship
encounters showed that the species-specific signature pattern was
used in almost all displays directed toward other territorial males,
but was hardly or not at all recognizable in courtship displays.
The only displays which are consistent in their form and use are
not species-specific. The jiggle bob, although not always performed
in courtships of unreceptive females, was never seen in any other
context. The display of rapid rhythmic bobs, shown by submissive
males, was not performed by courting males and only rarely by
and then modified by a very aggressive posture.
territorial males,
The dewlap fanning display was seen in all contexts but was used
consistently after copulation with a distinct erect posture and may
be associated with successful copulation. It has not been described
as such for other species, but probably occurs in A. carolinensis
(Crews, personal communication) and may be used by other anoles.

The color of the dewlap is also a species specific signal. But it may
not be a symbol to aid species recognition as much as a means of
emphasizing the bobbing display with the bright flash of color
(Greenberg and Nobel, 1944; Williams and Rand, 1977). Dominant
males in agonistic situations use the dewlap in bobbing displays
significantly more than submissive males, but the dewlap is used for
fanning displays at a significantly higher frequency by submissive
males. Therefore, dewlap flashes alone may be less aggressive
signals than when they accompany bobbing displays.
If the dewlap per se were being used as a species-specific signal,
one might hypothesize an increase in its use in courtship. However,

20 BREVIORA No. 479
fanning displays comprise a small proportion of premating displays,

which is lower than its average use in other contexts, and the dewlap
is used bobbing displays at only slightly higher frequencies during
in
courtship prior to mating than at other times. Therefore the two

signals, the dewlap and the species-specific signature pattern, which
one might expect to play a role in species recognition and female
choice, are not particularly associated with courship, at least not
when the animals are at relatively close range as they were in these
tests.
Anolis sagrei originates from Cuba, an island with a very complex

Anolis fauna (22-28 species, Williams, 1969). The stereotyped
display pattern is more variable than one might expect with so many
congeneric species needing reproductive isolation (Ruibal, 1967).
Information to enable species recognition may be redundantly
coded by many physical and behavioral characteristics (Williams
and Rand, 1977). This seems to be the case with A. opalinus on
Jamaica, which has no discrete display pattern even though it is
sympatric with its two closest relatives, A. garmani and A. grahami.
All three species have similar dewlap color and display structure but
differ greatly in body size, shape, and color (Jenssen, 1979).
The signature display and increased use of the dewlap with the
display are associated with territorial status in agonistic encounters.
But neither shows an increase in use as the encounter escalates. The
state of arousal and the willingness to escalate may be communi-
cated by the modifiers of the displays, especially body posture and
shape, and probably by the timing of the events. The more
aggressive male of a pair was usually the first to erect his crests,
engorge his throat, and laterally compress his body. Small shifts in
posture or position in tense moments are probably also used as

aggressive signals. Characterization of the behavioral repertoire of
A. sagrei has been elusive (Ruibal, 1967) because of this subtlety.
ACKNOWLEDGMENTS
I would like to thank E. E. Williams, D. Crews, W. Gartska, T. A.
Jenssen, and A. S. Rand for their critical comments on earlier drafts
of this paper; J. F. A. Traniello for his technical help and
encouragement; B. Scott for the art work; and J. S. Godley for
organizing the lizard hunt. This study was supported in part by
NIMH Research Scientists Development Award 00135 to D. Crews.
1984 DISPLAYS OF /I A'OZ./5S/lG/?£/ 21
LITERATURE CITED
Carpenter, C. C. 1967. Aggression and social structure in iguanid lizards, pp.
87-105. In W. Milstead (ed.) lizard Ecology: a Symposium, Columbia Univ.

Missouri Press.
Carpenter, C. C, and G. Gri^bitz. 1961. Time-motion study of a lizard.
Ecology 42: 199-200.

Crews, D. P. 1975a. Effects of different components of male courtship behavior
on environmentally induced ovarian recrudescence and mating preferences in
the lizard, Anolis carolinensis. Anim. Behav., 23: 349-356.
1975b. Inter- and intraindividual variation in display patterns in the
lizard. Anolis carolinensis. Herpetologica. 31: 37-47.

1977. The annotated anole: studies on the control of lizard reproduction.
Am. Sci.. 65: 428-436.
Evans, L. T. 1936. A study of a social hierarchy in the lizard, Anolis carolinensis.

J. Genet. Psychol., 48: 88-111.
1938. Courtship behavior and sexual selection of Anolis. J. Comp.
Psychol., 26: 475-497.
Greenberg, B., and G. K. Noble. 1944. Social behavior of the American
chameleon (Anolis carolinensis Voight). Physiol. Zool., 17: 392-439.
Greenberg, N. 1977. A neurethological study of the display behavior in the
lizard, Anolis carolinensis (Reptiiia, Lacertilia, Iguanidae). Am. Zool., 17:
191-201.
Hover, E., and T. Jenssen. 1976. Descriptive analysis and social correlates of
agonistic displays of Anolis limifrons (Sauria: Iguanidae). Behav., 58: 173-191.
Jenssen, T. A. 1970a. Female response to filmed displays of Anolis nebulosus
(Sauria: Iguanidae). Anim. Behav., 18: 640-647.
1970b. The ethoecology of Anolis nebulosus (Sauria: Iguanidae). J.
Herp., 4: 1-38.
1971. Display analysis of Anolis nebulosus (Sauria: Iguanidae). Copeia,
1971: 197-209.
1977. Evolution of anoline lizard display behavior. Amer. Zool., 17:
203-215
1978. Display diversity in anoline lizards and problems of interpretation,
pp. 269-285. In N. Greenberg and P. D. MacLean (eds.) Behavior and
Neurology of Lizards, NIMH.

1979. Display behavior of male Anolis opalinus (Sauria, Iguanidae): a
case of weak display stereotypy. Anim. Behav., 27: 173-184.
Jenssen, T. A., and E. Hover. 1976, Display analysis of the signature display of
Anolis limifrons (Sauria: Iguanidae). Behav., 57: 227-240.
Jenssen, T. A., and L. M. Rothblum. 1977. Display repertoire analysis oi Anolis
lownsendi (Sauria: Iguanidae) from Cocos Island. Copeia, 1977: 103-109.
Noble, G. K., and H. T. Bradley. 1933. The mating behavior of lizards: its
bearing on the theory of sexual selection. Ann. N. Y. Acad. Sci., 35: 25-100.
Rand, A. S., and E. E. Williams. 1970. An estimate of redundancy and
information content of anole dewlaps. Am. Natur., 104: 99 103.
22 BREVIORA No. 479
RuiBAL, R. 1967. Evolution and behavior in West Indian anoles. pp. 16 140. In
1
W. W. Milstead Lizard Ecology: a Symposium, Columbia Univ. Missouri

(ed.).
Press.
Stamps, J., and G. Barlow. 1973. Variation and stereotypy in the displays of
Anolis aeneus (Sauria: Iguanidae). Behav.. 48: 67-94.
Williams, E. E. 1969. The ecology of colonization as seen in the zoogeography of
anoline lizards on small islands. Quart. Rev. Biol., 44: 345-389.
1976. West Indian Anoles: a taxonomic and evolutionary summary. I.
Introduction and species list. Breviora Mus. Comp. Zool. No. 440, pp. 1-21.
Williams, E. E., and A. S. Rand. 1977. Species recognition, dewlap function
and faunal size. Am. Zool., 17: 265-274.
' 1^ • •
liBRARY
JUN 1 7 1985
B R T"V I O R A
Miiiseiiim of Comparative Zoology
us ISSN 0006-9698
Cambridge, Mass. 21 June 1985 Nimber 480
THREE NEW LIZARDS OF THE

GE^VS EMO/A (SCINCIDAE)
FROM SOUTHERN NEW GUINEA
Walter C. Brown' and Fred Parker-
AiiSTRACT. Three new species of skinks in the genus Enioia arc described from
provinces south ol the central mountain range. New Guinea.
INTRODUCTION
The genus Emoia is widespread through much of the Indo-
AustraHan Archipelago and the islands of the Pacific, and ranges
into the Philippines in the north and northeastern Australia and
New Hebrides in the south. The greatest diversity of species occurs
in New Guinea and surrounding smaller islands. The three new
species described in this paper are from south of the central moun-
tain range in New Guinea.

One species is based on a series of specimens collected primarily
by the junior author in the Fly River and Bamu River drainages.
Western Province, Papua New Guinea. The second ranges from
Western Province westward through southern Irian Jaya. Both spe-
cies are in the E. haudini-suhmetallica group as defined by Brown,
1953. The third species which belongs to the E. physicae group
(Brown, 1953) has a wide range in several provinces of southern

Papua New Guinea. The last two, in addition to the series collected
in earlier collections.
by the junior author, are also represented
of Sciences, San Francisco.

'Department of Herpetology. California Academy
Menlo College. Menlo Park. California 94025.
California 94118. and.
-717 Ross River Road. Kirwan. Queensland, Australia.
2 BRFVIORA No. 480
These had been misidentified as juveniles of E. p. physicae, E. p.

tropldolepis, or E. haudini.
Emoia aurulenia new species

(Fig. I)
Holotype: MCZ 142327, adult male, collected at Migalsimbip in
the upper Fly River drainage at an elevation of about 1,200 m.

Western Province, Papua New Guinea, Fred Parker Coll., 16 August
1972.
Paratypes: The following localities in Western Province, Papua

New Guinea. Migalsimbip: SAM 11637; AM 40778; MCZ
142322-26, 142328-30, 152268-71, 152273-75. CAS 154186,
AMNH 111718; UPNG 6477-78, 6480, 6483; Derongo: MCZ
131895-900; AMNH 103961; Menemsore: MCZ 131901; Emeti:
MCZ 152265-67, 152291, AMNH 1 11715; Tingkem: MCZ 131894.
Diagnosis. This species differs from the other New Guine?n spe-
cies of the Emoia haudini section in the following combination of
characters: (1) small size, 37+ to 49.5 mm
snout-vent length at
maturity; (2) pattern of gold-orange markings on the sides of the
neck and the lateral surfaces of the body in life (these fade to dirty,
silvery-white in preservative); (3) low number of midbody scale

rows; 26 to 32 (mean 28+), rarely greater than 30; (4) number of
subdigital lamellae, 39 to 48 beneath the fourth toe; and (5) number
of paravertebral scale rows between the parietals and base of tail 45
to 52.
Description. A small Emoia. snout-vent length of 41.3 to 49.5 mm

for 16 males and 37.4 to 47.5 mm
for eight females; habitus moder-
moderately tapered, bluntly rounded, its length

ately slender; snout
35 to of head length and 52 to 60% of head breadth; head
40%
breadth 64 to 69% of head length and 15 to 18% of snout-vent
length; eye moderately large, its diameter 7 to 85%) of snout length
1
and 40 to 45% of head breadth; ear diameter about one third to one
half of eye diameter; rostral broader than high, forming long, nearly
straight suture with frontonasal; supranasals widely separated, nar-

rowly triangular, in contact with anterior loreal; prefrontals moder-
ately to widely separated; frontal longer than broad, rounded

posteriorly, shorter than fused fronto-interparietal shield, in contact
with first and second supraoculars; four large supraoculars;
parietals large, in contact posteriorly; one pair of nuchals; anterior
1985 NEW SPECIES OF EMOIA
Figure I. An adult A.', auiulenla.
loreal higher and shorter than posterior, in contact with second, or

firstand second supralabials; six to seven upper labials, fifth (very
rarely sixth) enlarged and beneath eye; six to seven lower labials;
dorsal scales smooth, vertebral rows not distinctly enlarged; 26 to 32
(usually 28 to 30) midbody scale rows; 45 to 52 (mean 48.3) paraver-
tebral rows between parietals and base of tail, ventrals about same
size as dorsals;preanals somewhat enlarged; limbs well developed,
length of extended hind limb 90 to 10% of axilla-groin distance and
1
47 to 54% of snout-vent length; 39 to 48 (mean 42.6) smooth,

rounded lamellae beneath fourth toe; 10 to 14 lamellae under first
toe; rank of adpressed toes from longest to shortest four, three, two
to five, one; tail slender and much longer than body.
Measurements (in mm) of Holotype. Snout-vent length 49.0;
axilla-groin distance 24.8; hind limb length 24.1; head length 1.9; 1
head breadth 8.1; snout length 4.35; eye diameter 3.4; ear diameter
1.5; tail length 76.4.
Color in Preservative. The basic ground color on the dorsum
ranges from light olive greenish-brown to brown marked by lighter
and darker blotches, usually involving all dorsal scale rows (or occa-
sionally not including the paravertebral rows). The light and dark
4 BREVIORA No. 480
blotches may occupy alternate scales or involve two or more scales.

The upper lateral surfaces are dark brown, bordered dorsally by a
row of scattered, single or clustered, pale scales. There are also
scattered pale scales between and on the limbs. The neck is marked
by pale and dark blotches, and the posterior labials by dark bars.
The venter is dirty white to grayish, lightest on the chin and in the
limb regions.
in life, the top of the head is bronze with a lighter line from above
the eye to the nuchals or for a few specimens extending further
posterior onto the neck. The dorsum is gray or brown with alternate
black scales or paravertebral rows of black spots in a checkerboard
pattern. The upper flanks are black or mottled black and brown.
The lower flanks and usually the side of the neck are marked by
golden yellow spots. The venter is yellowish to cream.
Comparison. Enwia aurulenta is related to Papuan species of the
E. hauiilni-suhmeial/ica group, but is readily distinguished in life by
such features of the color pattern as the checker-board pattern on

the dorsum andthe prominent golden yellow spots on the sides. It
also characterized by a lower number of midbody scale rows than
is
other known species of this section. Also, the number of subdigital

lamellae is greater than for most other species of this group, only
overlapping slightly with E. s. popei and two other undescribed
species (Table I).
Etymology. The name aurulenta refers to the golden yellow spots
on the sides.
Note on Reproduction. No data are available for this species.
Note on Habitat. This is a diurnal species which is active on the
rain forest floor beneath the unbroken canopy.
Range. Etnoia aurulenta is known only from the Fly and Bamu
River drainages in the Western Province, Papua New Guinea.
Another undescribed species represented by samples from several
populations south of the central mountain range between Western
Province in Papua New Guinea and the Jamur Lake area in western
IrianJaya can not be identified with any previously described spe-
cies ofEmoia. Four specimens from various localities in Western
Province, Papua New Guinea were collected by Fred Parker in 1969
and 1971. A series of specimens in the Leiden Museum collected
during a 1955 survey and the 1959 expedition along the Digul
River and several of its tributaries in southeast Irian Jaya and sev-
eral collected earlier at more westerly locations (identified as E.
1985 NEW SPECIES OF KMOIA
haudini) also belong to this species. These were collected in part by

the British Ornithological Expedition in the Mimika River area in
1913 and in part by the Royal Netherlands Geographical Society
Expedition in the Jamur Lake area in 1959.
Emoia aenea new species

(Fig. 2)
Lygosonia hauc/ini, (part) Boulenger, 1914, Trans. Zool. Soc. Lon-

don. 20: 259.
Holotype: MCZ 131949, adult male, collected at Menemsore,
Western Province, Papua New Guinea, Fred Parker coll., 30 March
1969.
Paraiypes: The following localities in Western Province, Papua

New Guinea, Matkomrae: MCZ 144393, Emeti: MCZ 144386,
Kiunga: MCZ 13 1948; the following localities in Irian Jaya, Mimika
River area:BMNH 1913.1 81-82, 1913. 10.31. I64F;
1. 1. Gariau,
Jamur Lake area: RMHN 21278; lower DigulTanah Merah: River,
RMHN 21180-85, 21273 74, 21276-77; 21279; Tanah Tinggih:
RMHN 21275, 21280-82; Kouh: RMHN 21186 89; CAS 156680;
Mariang: RMHN 21 190-94.
Figure 2. An adult K. aciwa.

BREVIORA No. 480
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1985 NEW SPECIES OF AA/OM 7
Diagnosis. This species can be differentiated from other species of

the E. submeiallica complex on the basis of the following combina-
tion of characters: (1)31 to 36 midbody scale rows; (2) 49 to 54
paravertebral scale rows between the parietals and the base of the
tail; (3) 39 to 46 rounded lamellae under the fourth toe; (4) sixth
(rarely fifth) upper labial enlarged and beneath the eye; (5) snout-
vent length at maturity 53.5 to 71.0 mm; (6) such features of the
color pattern as dorsum medium brown (in life bronzy anteriorly),
sometimes with darker blotches in longitudinal series mostly lateral
to the paravertebral scale rows; upper lateral surfaces (two or three
rows of scales) darker brown to blackish, lower lateral surfaces
grayish slate; venter whitish to ivory, occasionally more grayish on

head and abdominal regions, pale spot on neck absent.
Description. An Emoia of intermediate size, snout-vent length
53.5 to 71.8 mm for 1 1 males and 59.6 to 69.5 mm for three females;
habitus moderately robust, limbs well developed; snout moderately
tapered, bluntly rounded, its length 54 to 62% of head breadth and
34 to 39% of head length; head breadth 58 to 69% (rarely less than
60%) of head length and 14 to 17% of snout-vent length; eye rela-
tively large, its diameter 67 to 90% of snout length and 36 to 50% of
head breadth; ear diameter one quarter to nearly one half of eye
diameter; rostral broader than high, forming long, nearly straight
suture with frontonasal; supranasals narrow, elongate, in contact
with anterior loreal; prefrontals moderately to widely separated;
frontal longer than wide, about same length as fused fronto-
interparietal shield, broadly rounded posteriorly, in contact with
first and second supraoculars; four large supraoculars; seven or
eight supraciliaries; parietals large, in broad contact posteriorly; one
pair of nuchals; anterior loreal shorter and higher than posterior, in
contact with first and second, second, or second and third upper
labials; usually six to eight upper labials, sixth (occasionally fifth)
enlarged and beneath eye; seven or eight lower labials; dorsal scales
smooth, paravertebral scales not or scarcely enlarged; midbody
scale rows 31 to 36; paravertebral scale rows between parietals and
base of 49 to 54; preanals not or slightly enlarged; limbs well
tail
developed, length of extended hind limb 91 to \20% of axilla-groin

distance and 45 to 54% of snout-vent length; 39 to 46 rounded
lamellae under fourth toe; 10 to 14 lamellae under first toe; rank of
adpressed toes from longest to shortest four, three, two, five, one;
tail longer than body.
8 BREVIORA No. 480
Measurement (in mm) of Holotype. Snout-vent length 71.0;
axilla-groin distance 34.7; hind limb length 31.7; head length 17.8;
1.5; tail length 104±.
Color in Preservative. The dorsal ground color ranges from tan to
a vaguely grayish-brown or a light brown. It may be nearly uniform
or be marked by a longitudinal row of dark brown blotches or
sometimes dark brown, narrow, marginal lines on the scale rows
lying lateral to the paravertebral rows. The top of the head, except
sometimes the margins of the scales is relatively uniform. The upper
lateral surface is marked by a narrow, darker brown band varying
from one or two to three or four scale rows in breadth, narrowing
on the neck and head and scarcely evident on the snout. This band is
sometimes bordered by scattered, pale scales along its dorsal margin
and there are numerous pale scales on the lower lateral surfaces, but
the dark band itself it usually unmarked. The upper part of the
upper labials are dusky, and there are some dark marks on some of
the lower labials. The venter is whitish ivory, unmarked, and the
undersurface of the tail has scattered, small, blackish or grayish
spots.
in life the head and neck are bronzy, the belly white. The Kiunga
specimen has a patch of pink just anterior to the vent, the Menem-
sore specimen a patch of orange. The dorsal scales are smooth and
iridescent.
Etymology. The name aenea refers to the bronzy coloration on

the top of the head and anterior body.
Comparisons. This species is close to E. suhmetallica ohscura and
E. suhmetallica popei in scale counts. In color E. aenea is more
s. ohscura than to E. s. popei, lacking the small, whitish
similar to E.
blotch on the side of the neck which is typical of the latter. E. aenea
also appears to be slightly larger in size and the number of subdigital
lamellae may be slightly greater than for either of the two subspecies
o( suhmetallica (Jab\Q 1), although it is closest to E. s. popei m this
character. Also the sixth upper labial is usually the one which is
enlarged and beneath the eye for E. s. popei and E. aenea, whereas
it is the fifth for E. s. ohscura.
Note on Reproduction. RMHN 21281, a gravid female measuring

62.2 mm snout-vent length, has two eggs in the oviducts.
1985 NEW SPECIES OF A/WO/zl 9
Habitat Note. Parker's specimens were found in primary forest in

areas of high rainfall. All localities are at low elevation between sea
level and 100 m.
Range. Known from Western Province, Papua New Guinea, and
Mimika River, Digul River and Jamur Lake areas in southern Irian
Jaya.
Emoia physicina new specices

(Fig. 3)
Holotype. MCZ 152287, adult male, collected near Emeti,

Western Province, Papua New Guinea at an altitude of about 100
m, Fred Parker coll., October 1971.
Paratypes. Same locality as holotype: MCZ
142567-72, 152283-86,
152288. 152290, AMNH111713-14; other localities in Western
Province, Menemsore: MCZ 131945 46, 152280; Kiunga: MCZ
152281-82; Derongo: AMNH
111712, MCZ
131938-39, 131942;
Matkomrae: MCZ
131943 44; localities in other provinces of
Papua New Guinea, Chimbu Province, Karimui: 97308-09, MCZ
99193-96, 109579-82, 109584-96, CAS 1 17731, 1 17750, 1 18769-70,
118807; Soliabeda: CAS 117733 34. MCZ 109509, 109589-96,
109598-99; Dege: MCZ 90750; Bomai: MCZ 90478-501; Eastern
Highlands Province; AMNH
98570a-b; Gulf Province, Pio River:
AMNH 102233, CAS MCZ 109601-03; Koni: CAS
118871-72;
117726-27; MCZ 109514, 109529, 109532; Uraru: CAS 117753-54,
MCZ 109572-75, 109577- 78; Oroi: MCZ 109833; Weiana: MCZ
109521-23; localities in Irian Jaya, Assike on the Digul River:
RMHN 5081a-b.
Diagnosis. This species can be distinguished by the following
combination of characters: (1) relatively weak keels on the dorsal
scales, especially on the posterior half of the body; (2) small size,
39.0 to 50.7 mm snout-vent length; (3) relatively low midbody scale
count 30 to 38; (4) fifth upper labial enlarged and beneath the eye;
(5) number of vertebral scale rows between the parietals and
base of
tail 45 to 54, and (6) of fourth toe lamellae 34 to 43.
number
Description. A
relatively small Emoia, snout-vent length 39.0 to
50.7 mm
for twelve mature males and 43.3 to 50.0 for fourteen mm
mature females; habitus moderately slender with well-developed
limbs; snout moderately tapered, broadly rounded at tip, its length
10 BREVIORA No. 480
hijiuic 3. All adult /.. phvsiiiiia.
51 to 60% of head breadth and 35 to 39% of head length; head

breadth 60 to 72% of head length and 15 to 18% of snout-vent
length; eye relatively large, its diameter 72 to 85% of snout length
and 39 to 46% of head breadth; ear moderate, its diameter about
two fifths to three fifths of eye diameter, usually with two lobules
anteriorly; rostral broader than high, forming long, convex or trun-
cate suture with frontonasal; prefrontals moderately to widely
separated; frontal slightly longer than broad, about as long as
fronto-interparietal shield, in contact with two anterior supraocu-
lars;four large supraoculars; frontoparietals fused with interparietal
into one shield; parietals in contact; one pair of nuchals; anterior
loreal shorter and slightly higher than posterior loreal, in contact
with first or first and second upperseven or eight upper
labials;
labials, fifth (very rarely sixth) enlarged and beneath eye; six or
seven lower labials; dorsal scales, at least posteriorly, with three
weak to moderate keels; 30 to 38 scale rows at midbody; 45 to 54
(rarely greater than 53) paravertebral rows between parietals and
base of tail; limbs well developed, length of extended hind limb 9 to 1
105% of axilla-groin distance and 44 to 54% of snout-vent length; 34

to 43 rounded lamellae beneath fourth toe (only one specimen has
1985 NEW SPECIES OF /lA/O/zI 11
more than 40); 8 to 10 beneath first toe; rank of adpressed toes from
longest to shortest four, three, five, two, one; tail much longer than
body.
Measurements (in mm) of Holoiype. Snout-vent length 48.1;
axilla-groin distance 21.9; hind limb length 22.6; head length 12.9;
1.6; tail length 74.8.
Color in Preservative. The dorsum (six to eight scale rows) is
medium brown, nearly uniform or with rows of darker brown spots

marginal to the paravertebral rows. The lateral surfaces, at least five
to six upper scale rows, are usually much darker brown,
nearly
uniform or with scattered single or small groups of pale or whitish
scales and often one or two light scales on neck. The upper labials
are dusky, at least on the dorsal half, and the lower labials may or
may not have dusky blotches. The venter is grayish slate, more dusky
tan or ivory in the limb region.
For living specimens, the dorsum is grayish-brown to brown,
usually with two rows of paravertebral dark spots. Lateral surfaces
are blackish with scattered light spots on the lower flanks. Some
specimens have the upper and lower surfaces of the snout suffused
with red.
Etymology. The name physicina refers to the fact that the species
has been confused with juveniles of other keel-scaled species such as
E. physicae.
Comparisons. Emoia physicina can be distinguished from all
other known species of the E. physicae section except E. callistica
(1) on the basis of its sm.all size, (2) the very weak keels which
separate it from all but E. kuekenthali, (3) in having the fifth instead
of sixth upper labial enlarged and beneath the eye. It differs from E.
callistica in lower number of midbody scale rows and subdigital
lamellae, in the distinct nuchals, and in the very weak rather than
strong keels on the dorsal scales.
Note on Reproduction. Gravid females CAS 110360, 49.3 mm
snout-vent length and CAS 18770, 50.0 1 mm snout-vent length have
two eggs in the oviducts. One hatchling (MCZ 90750) measures 24
mm snout-vent length.
Habitat Note. This species occupies the rain forest floor in areas
shaded by the canopy. Specimens do bask in sunflecked areas, how-
ever. It is strictly diurnal.
12 BREVIORA No. 480
-^|SS.\^
Range. E. physicina is-Jcnown from the following provinces in

southern Papua New Guinea: Eastern Highlands, Gulf, Chimbu,
and Western.
'^^* ACKNOWLEDGMENTS
We wish and Ernest E. Williams, Museum
to thank Pere Alberch
of Comparative Zoology (MCZ), Allen E. Greer, Australian
Museum (AM), Richard G. Zweifel, American Museum of Natural
History (AMNH), John Pernetta, University of Papua New Guinea
(UPNG), R. C. Drews and A. E. Leviton, California Academy of
Sciences, and Terry D. Schawner, South Australian Museum
(SAM) for the loan of material used in this study. The senior author
also thanks the trustees of the Australian Museum and the Science
and Industry Endowment Fund, Commonwealth Scientific and
Industrial Research Organization of Australia, for their financial
assistance while studying in the museums of Australia. Robert
Drewes and Allen Greer have read the manuscript and been most
helpful with their suggestions.
LITERATURE CITED
Brown, W. C. 1953. Results of the Archbold Expeditions No. 69. A review of
New Guinea li/ards allied to Hnioia hauc/ini and Enioia physicae. Amer. Mu.s.
Novitatcs. No. 1627, pp. I 25.

B R E'"V I O R A
useiiiii^^S^^mparative Zoology
SSN 0006-9698
Cambridge, Mass. 21 June 1985 Number 481
A NEW ANOLIS OF THE LIONOTUS GROUP FROM

NORTHWESTERN ECUADOR AND SOUTHWESTERN
COLOMBIA (SAURIA: IGUANIDAE)
Kenneth Miyata'
Abstract. Anulis lynchi. new species,

is described from several lowland rain
Ecuador and southwestern Colombia. It is allied to

forest localitites in northwestern
the semiaquatic anoles of the lionoius species group, and appears to be most closely
related to Anolis puecilopus Cope of Panama and northwestern Colombia. The
lionotus species group and the known distribution of its South American representa-
tives is discussed.
INTRODUCTION
Much of the complexity of the South American Anolis fauna is
contained within the narrow confines of the wet Pacific coastal
region of Colombia and Ecuador (see Table 2 in Williams, 1976).
Although these anole communities may be richer than similar
Amazonian communities (up to perhaps twelve sympatric species
vs. six in Amazonia), a major feature of the region seems to be more
restricted ranges of the individual species and narrower microhabi-

tat preferences. Localities close together geographically may have
strikingly different faunas, and within a single fauna there may be
distinct forest and forest edge components.
Until recently there were few representative series of most South
American anoles. Moderate to large series of some species from the
Andes and their western and eastern flanks have been collected
'Deceased; formerly Museum of Comparative Zoology, Harvard University. Cam-

bridge, Massachusetts 02138.
2 BREVIORA No. 481
during the past decade by workers interested primarily in frogs, who

find sleeping anoles at night. These have sometimes been the ones
most poorly represented in museum collections; in some cases it
seems as if a different anole fauna can be found at night. Field
parties from the University of Kansas led by Dr. John D. Lynch
have night-collected a series of anoles from the vicinity of Santo
Domingo de los Colorados, Provincia del Pichincha, Ecuador.
Some of these prove to represent a new species, which here name: 1
Anolis lynchi, new species

(Figs. 1-4)
Holotype: MCZ 124406, an adult male from Santo Domingo de

los Colorados, 600 m elevation, Provincia del Pichincha, Ecuador,
R. W. Henderson coll.. 31 July 1968.
Paratypes: ECUADOR: Provincia del Pichincha: KU 178953,
Santo Domingo de los Colorados, 580 m, elevation J. D. Lynch
coll., 12 June 1977; KU 178954-178958, 2 km E, km S Santo I
Domingo de los Colorados, 600 m elevation, T. Berger and J. D.

Lynch coll., 13 June 1977, and KU 178959 from the same locality, J.
D. Lynch coll., July 1977; MCZ 157156, 2 km E,
1 1 km N Santo 1
Domingo de los Colorados, 620 m elevation, J. D. Lynch coll., 31

July 1977; Provincia de Esmeraldas: USNM 21 1222, 1-2 km El W
Placer, 390-410 m elevation, J. A. Peters coll., December 1958.
1
Refered specimens: COLOMBIA: Departmento del Cauca:

AMNH 107864-70, 109598-602, Quebrada Guangui, 0.5 km above
Rio Patia, upper Saija drainage, 100-200 m elevation.
Diagnosis. Anolis lynchi can be distinguished from all other
known species of South American Anolis by the high number of
scales across the snout ( 8-29 at level of second canthals), the small,
1
subequal scales in the supraocular area, distinct pale flank stripes,

and the greatly enlarged bilobed hemipenes of adult males.
'
Description. Head. Head scales small, flat to slightly granular.

(Counts for holotype are in parentheses.) Eighteen to 29 (28) scales
across snout at level of second canthals. Seven to 10 (8) scales
border rostral posteriorly. Circumnasal scale separated from rostral
by one to two (2) scales. Nine to 12(10) scales between supranasals.
Supraorbital semicircles distinct, raised, separated medially by
three to six (4) scales. Supraocular scales small, subequal, slightly
1985 ANOLIS LYNCHI
i
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BREVIORA No. 481
Figure 2. Anoll.s lynchl. new species. Lateral view of head of holotype (MCZ
124406).
keeled to granular, to 19 (18) across widest point. Supraciliaries

1 1
three to five (4). Canthus distinct in adults, less so in juveniles; five

to six (5) canthals, the second the largest. Loreal rows seven to ten
(10), subequal. Nasal area slightly swollen in adults.

Temporals and intertemporals granular. Interparietal slightly
smaller than ear opening, separated from supraorbital semicircles
by four to six (4) scales. Interparietal much smaller than ear opening
in some specimens. Scales anterior to interparietal enlarged or sub-
equal (enlarged) with respect to temporals; scales posterior to inter-

parietal slightly flattened or granular (flattened), larger than, but
grading into, dorsal scales. No enlarged supratemporal rows. Dis-
tinct thin fold extending from supraorbital semicircles laterally to
mid-orbit, where it extends
posteriorly above ear opening and fades
out in axillary region.
Suboculars slightly keeled, separated from supralabials by one

row of scales. Seven to 10(10) supralabials to center of orbit.
Mental groove extending approximately three quarters across,
each part much wider than deep. Six to eight (7) scales in contact
with mental posteriorly. Eight to 11 (II) infralabials. Chin scales
granular, except for some which are slightly keeled.
1985 A NOI.IS I. YNCHI
Figure 3. Anolis lynchi. new species. Dorsal view ot head of holotype (MCZ
124406).
BREVIORA No. 481
Figure 4. Anolis lynchi. new species. Ventral view of head of holotype (MCZ
124406).
Dewlap. Dewlap of adult males medium-size, extending from

under the midpoint of the orbit posteriorly to a point just under the
axilla. Scales form 13 to 16 longitudinal rows when dewlap
extended, but arranged irregularly and not parallel. Dewlap scales
narrowly compressed laterally and elongate, especially at the edge.
Dewlap of adult females small.
Trunk. Middorsal scales flattened or slightly keeled (flat); two
medial rows slightly larger than adjoining rows. Lateral scales gran-
ular, grading into larger, subimbricate ventral scales. Ventral scales
1985 ANOUS l.YNCHI 7
slightly tomoderately keeled; keeling more prominent midventrally

than anteriorly or posteriorly.
Limbs and Digits. Scales on limbs smooth to lightly keeled;
generally smoother on upper arm and thigh than on lower arm or
calf. Scales on forearm subequal; those on hind limbs are larger
anteriorly. Keeling on limb scales unicarinate. Supradigital scales

multicarinate, usually with two longitudinal keels. Fifteen to 18(17)
lamellae under phalanges ii and iii of fourth tow. Distal phalanx
raised slightly above adjacent phalanx.
Tail. Base of tail greatly swollen in adult males, slightly thicker
than trunk tapering abruptly about one quarter
in pelvic region,
SVL behind the vent. Tail slightly compressed laterally, height

about 20 percent more than width. Caudal scales unicarinate, more
strongly keeled ventrally than laterally or dorsally. Males with a
distinctly keeled middorsal scale row beginning where the swollen
tailbase ends. Postanals not enlarged.
Heniipenes. The hemipenes of adult males are greatly enlarged,
accounting for the extreme swelling at the base of the tail. The
everted hemipenes are bifurcate, the split extending back about one
third of the distance from the distal end to the base. The distal end
has small calyces that become larger proximally. The shaft has large
folds, becoming smoother on the base. The sulcus spermaticus has
several longitudinal folds.
Measurements. Adult males 54.2 to 61.8 mm snout-vent length
(SVL), adult females 49.5 to 59.3 SVL. mm
Color as Preserved. The dorsum ranges from cinnamon-brown
to deep fuscous. Males show indistinct lighter brown longitudinal
stripes dorsolaterally; females have a distinct gray middorsal stripe.
Some males show broken and indistinct series of small, elongate
a
dark brown marks middorsally. Both sexes show traces of indistinct
pale and dark brown lateral stripes that are very thin just above a
prominent muddy gray flank stripe. The gray is heavily flecked with
brown, but the flank stripes are very distinct in most specimens.
Below this flank stripe is a dark brown stripe heavily flecked with
gray. The gray flank stripe extends to the orbit, and the sides of the
head are gray with brown flecks and spots. The venter is an almost
immaculate light gray to cream, with some gray flecking laterally.
The undersides of the limbs and tail are lightly mottled with brown.
Color in Life. The following description of Ecuadorian speci-
8 BREVIORA No. 481
mens from Santo Domingo de los Colorados is compiled from the

field notes and Kodachrome transparencies of living animals pro-
vided by John D. Lynch.

The dorsum is brown to reddish brown, with an obscure series of
dark brown middorsal blotches in males and a pale gray middorsal
stripe in females. The flank stripe is dull cream and is bordered
above by a thin, yellowish cream stripe. The flanks are mottled
brown, with an olive wash in some specimens. The sides of the head
are off-white with dark brown marks under the orbits. The chin,
venter, and underside of the limbs are cream; the underside of the
tail of females and the underside of the
tail of males posterior to the
swollen tail base is yellowish.The dewlap is orange with white

scales. The iris is bright coppery brown. The tip of the tongue is
gray.
Color notes of the Colombian specimens were provided by
Charles W. Myers. They were similar to the Ecuadorian specimens,
except Myers noted grayish chins, a dull gray reticulum under the
limbs, and the dewlaps were dull orange (Myers' italics). Myers also
noted that the throat lining was unpigmented.
Etymology. The species is named after John D. Lynch, whose
indefatigable excursions into the Ecuadorian night are responsible
for many of the known specimens.
Variation. The Colombian specimens referred to Anolis lynchi
differ somewhat from the Ecuadorian sample. They have fewer
scales across the snout at the second canthals (18-24 vs. 25-29) and
fewer scales across the supraocular region (11-15 vs. 15-18). Both
counts reflect slightly larger head scales in the northern population.
There also appears to be a slight difference in color of living ani-
mals. The Ecuadorian specimens had immaculate cream venters
while those of the Colombian specimens were cream with a dull
gray reticulum under the limbs and a grayish throat. The dewlaps of
adult males from both areas were basically orange, but those of the
northern population were characterized as a dull orange while those
of the southern populations were a fairly bright orange.
Despite the differences seen between the Ecuadorian and Colom-
bian populations, I
regard them as conspecific. The differences may
well be clinal, and there is no evidence for sharp discontinuities in
character states that might reflect breaks in gene flow. However,

since there is a possibility that the Colombian populations may be
distinct, 1 have not designated these specimens as paratypes.
1985 ASOI.IS l.YSCHI 9
Comparisons. A no/Is lynchi most closely resembles A. poecl/o-

pus of Panama and northwestern Colombia. The two are closely
related (see discussion below), but can be distinguished by the char-
acters summarized in Table I.
The reduced supraocular scales of A. lynchi are paralleled in two
other groups of South American anoles. The latifrons group
includes several giant anoles with adult SVLs of 125 mm and more.
The aequatorialis group is a poorly known complex of medium to
larger size anoles (adult SVLs generally over 80 mm). Species in
both groups lack transverse processes on the posterior caudal verte-

brae and are alpha anoles in the terminology of Etheridge (1960).
Since Anolis lynchi has these transverse processes, it falls into the
beta section of the genus. Anolis lynchi can be easily distinguished
from any known member of the latifrons or aequatorialis groups by
its smaller adult size (less than 65 mm
SVL vs. 80 mm
or more),
much smaller supraocular scales (11-19 across), smaller scales
across the snout (at least 18 at level of second canthals), and the
greatly enlarged, bifurcate hemipenes of adult males.
Anolis granuliceps, a small beta anole of uncertain relationship
from western Colombia and northwestern Ecuador, shows a similar
reduction in the size of the supraocular scales, but the central scales
in the supraocular area are slightly enlarged and keeled, while those
of lynchi are subequal and usually granular. The hemipenes of
granuliceps are small and not bifurcate, and the small male dewlap
is golden orange.
Natural History.All of the Ecuadorian specimens of A. lynchi
for which there are field notes were collected at night as they slept
on low vegetation adjacent to small streams in closed-canopy rain
forest. They were sleeping head-up on leaves and small ferns along-
side, or overhanging, the creek beds, usually within m of the 1
ground. The Colombian specimens were collected in a forested

ravine on low vegetation near a stream during the day. It appears
that A. lynchi is associated with streamside lowland rain forest
habitats, although admittedly collectors may have concentrated on
streamside habitats, or the lizards may simply have been easier to
see in these areas. Charles W. Myers, who collected the Colombian
sample, believes that it is not an aquatic anole in the same sense as
A. poecilopus. At his collecting site this niche was apparently occu-
pied by A. niacrolepis, another member of the lionotus group.
However, the association between A. Ivnchi and streamside forest
10 BREVIORA No. 481
habitats seems real, at least in the southern part of its distribution.

Despite extensive collecting in the vicinity of Santo Domingo de los
Colorados, the only available specimens were found in the imme-
diate vicinity of forest streams. Other anoles found within 200 m of
one such creek (2 km E, 1 km N Santo Domingo de los Colorados)
include A. chloris, fraseri, gracilipes, peraccae, princeps, and an
undescribed species related to vitiigerus.
Discussion. Williams (1976) recognized two South American
anoles, A. poecilopus Cope and A. macro/epis Boulenger, in his
iionotus species group. He has subsequently described a third, A.
riva/isWilliams, 1984. The group was characterized in Williams'
1976 key by the presence of a zone of enlarged flat or slightly keeled
middorsal scales. In A. niacrolepis and A. riva/is the enlarged mid-
dorsals are considerably larger than the lateral scales and are
arranged in from 11 to 17 longitudinal series. In A. poecilopus the
enlarged middorsals are only slightly larger than the laterals, into
which they grade, in A. lynchi the zone of enlarged middorsals is
scarcely evident. There are only two slightly larger and flatter para-
vertebral scale rows, and the difference in size and shape is slight.
The lack of an obvious zone of enlarged middorsal scales, and the
extreme reduction in the size of scales on the supraocular discs, at
first obscured the relationships of A. lynchi and suggested an asso-
ciation with the aequatorialis group (sensu Williams, 1976). How-

ever, the presence of transverse processes on the posterior caudal
vertebrae madethis impossible.
The
large bilobed hemipenes of A. lynchi provide the clearest
morphological evidence linking A. lynchi with the Iionotus species
group. Many anoles show a tendency for the distal end of the
hemipenis to be bilobed, but few show a strongly forked hemlpenis
with the bifurcation extending as much as a third the length of the
everted organ. This condition characterizes several semiaquatic
anoles in Central America and northwestern South America allied
to A. Iionotus Cope. In addition to a semiaquatic streamside habi-
tat and bilobed hemipenes, these lizards share a similar body pat-
The group consists of six currently
tern with distinct lateral stripes.
recognized species: A. Iionotus of central Panama, A. oxylophus of

Costa Rica, Nicaragua and western Panama, A. poecilopus of east-
ern Panama and northwestern Colombia, A. rivalis and A.
niacrolepis of western Colombia, and A. lynchi. Each is distinct in
1985 ANOI.IS l.YNCHI II
external morphology, and occupies largely exclusive ranges. Anulis

lynchi and A. niacrolepis have both been collected at Quebrada
Guangui Cauca, Colombia and A. poecilopus and A. lionutus
in
occur insympatry in central Panama (Campbell, 1973), but the

zones of overlap are probably narrow.
Anolis lynchi is the southernmost representative of the lionutus
group. The distribution of the South American species of the group
is shown in Figure 3. The range o{ A. poecilopus, which on the basis
of external similarity seems to be the closest relative oi A. lynchi, is

separated from that of A. lynchi by a substantial gap of approxi-
mately 1,000 km. The intervening area is inhabitated by A.
macrolepis and A. rivalis, which are not morphologically close to
either species.
ACKNOWLEDGMENTS
John D. Lynch collected most of the specimens of the new species
and provided information about the localities, field notes, and
Kodachrome transparencies. Charles W. Myers of the American
Museum of Natural History (AMNH) loaned Colombian speci-
mens and field notes. Ernest E. Williams shared his wealth of
knowledge on South American Anolis and commented on the

manuscript. Stephen Ayala commented on the manuscript and
answered questions regarding Colombian specimens. Other speci-
mens were loaned by William E. Duellman, Museum of Natural
History of the University of Kansas (KU), W. Ronald Heyer and
Ronald I. Crombie, National Museum of Natural History (USNM),
and Arnold G. Kluge. University of Michigan Museum of Zoology
(UMMZ). My own unsuccessful efforts to collect Anolis lynchi in
Ecuador were partially funded by Earthwatch and the Center for

Field Research of Belmont, Massachusetts and by the Museu de
Zoologia of the Universidade de Sao Paulo. thank Jeannie

I
Sellmer and Laszlo Meszoly for the illustrations.

12 BREVIOFtA No. 481
Figure 5. Distribution of the Anolis lionotus species group in South America

and lower Central America. Triangles: A. lynchi. n. sp. Solid squares: A. macrolepis.
Open squares: A. rivalis. Circles: A. poecilopus. Localities for A. macrulepis and A.

rivalis were provided by Stephen Ayala and Ernest E. Williams.
1985 ANOI.IS l.YNCHI 13
LITERATURE CITED
Cami'HM.i., H. W. 1973. Ecological observations on Anoli.s lionotus and Anolis
poecilopus (Reptilia, Sauria) in Panama. Amer. Mus. Nat. Hist. Novitatcs.
2516: I 29.
EiiiiKiDCi. R. E. I960. The relationships of the anoles (Reptilia: Sauria: Iguani-

dae). An interpretation based on skeletal morphology. Ann Arbor, University
Microfilms, xiv + 236 pp.
Wii.i.i.AMs, E. E. 1976. South American anoles: the species groups. Pap. Avuls.
Zool.. S. Paulo. 29: 259 268.
1984. New or problematic Aiuili.s from Colombia. 111. Two new semia-
quatic anoles from Antioquia and Choco. Colombia. Breviora Mus. Comp.
Zool.. No. 478, pp. I 22.
'^'"S.
COMP. zooc
B R EXI R a
iiseum '^
'd'f*
GoHiparative Zoology
us ISSN 0006-9698
NEW OR PROBLEMATIC ANOLIS FROM COLOMBIA. IV.

ANOL/S ANTIOQUIAE, NEW SPECIES OF THE
ANOLIS EULAEMVS SUBGROUP FROM
WESTERN COLOMBIA.
Ernest E. Williams'
Abstract. A new species of the Anolis eulaemus subgroup of the A. aequalorialis

species group from the western part of the Departmento Antioquia, Colombia is
closer to A. fitchi Williams and Duellman of southern Colombia and northern
Ecuador than to neighboring A. ventrimaculatus Boulenger and A. eulaemus Bou-
lenger of the Departments of Valle and Risavalda.
Three adult female Anolis from the Cordillera Occidental of

Antioquia, Colombia represent a distinctive new species of the
eulaemus subgroup of the Anolis aequatorialis species group. Other
members of this subgroup include A. eulaemus Boulenger, A. ven-
trimaculatus Boulenger, A. gemmosus O'Shaughnessy, A. maculig-
ula Williams and A. fitchi Williams and Duellman, all ranging
farther to the south in Colombia and Ecuador. Several novelties in
the genus Anolis have recently been discovered in Antioquia, so it
seems appropriate to emphasize the interest of the new material by
naming this lizard:

02138.
BREVIORA No. 482
U
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CL
B
u
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1985 ANOl IS ANTIOQUIAE
Kiîurc 2. Aii(>li\ (in/ioc/iilac. 1 ACM 72763. I.iitcral \icw ol head.
Anulis antiuquiae, new species

(Figs. 1-4)
Type: INDERENA 0277. adult female.

Type locality: Along a road paralleling Quebrada Chaparral,
Rio San Juan drainage, 10 km E of Andes (town), western Antio-
quia, Colombia. 2200 2300 m. Philip A. Silverstone, Carlos Arturo
Escobar and Luis Carlos Acevedo Cuartas coll., July 1971.
Paratypes. Antioquia: LACM 12763. adult female, same local-
ity as type. CSJ 310, adult female, Urrao.
Description. Head. Head scales minute, anteriorly weakly uni-
to multicarinate, posteriorly pustulose. Sixteen to 19 scales across
snout between second canthals. Six to seven scales border rostral
posteriorly. Anterior nasal scale separated from rostral by one scale
or in contact.
Supraorbital semicircles separated medially by four to five scales.

The medial supraocular scales enlarged, wrinkled or keeled, but
supraocular disk not or very weakly defined. Three short supercil-

iaries on each side, followed by granules. Canthus projecting later-
ally, sharp-edged, slightly overhanging loreal region. Second and

third or third and fourth or fourth and fifth scales longest. Ten to
twelve loreal rows.
BREVIORA No. 482
Figure 3. Anulis aniioquiae, LACM 72763. Dorsal view of head.

1985 ANOI.IS ASriOQUIAE
Figure 4 Aiuilis ciniuiqitiac. I. ACM 72763. Ventral view of head.

6 BRFVIORA No. 482
Temporals and supratemporals granular. An intertemporal dou-

ble row of enlarged scales hardly indicated. No interparietal. Scales
in the interparietal area granular posteriorly, grading into parietal
area, bounded by oblique lateral ridges which converge on but do
not meet a blunt occipital knob, larger scales alongside the
supraorbital semicircles. Ear moderate but less than half eye
aperture.
Suboculars weakly keeled, narrowly and irregularly in contact
with the supralabials. Six to seven supralabials to below the center
of the eye.
Mental completely or incompletely divided, each half as wide as
deep, indented medially by six to seven small swollen smooth scales
between the large smooth sublabials, only the first of which on each
side iscontact with the infralabials (on one side of INDERENA
in
0277 the second sublabial also is in contact with the infralabials).
Central gular scales smooth or keeled, imbricate, projecting, sub-

equal, becoming rather abruptly larger, laterally adjacent to the
infralabials.
Trunk. A central zone of small keeled juxtaposed scales grading
into smaller projecting and bluntly or sharply pointed flank scales
that are juxtaposed or separated by minute granules. Ventrals
larger, smooth, juxtaposed or subimbricate, not in transverse rows.

Dewlap. Moderate (female), extending posteriorly to the level
of the axilla. Scales close packed, more or less projecting, smaller
than ventrals.
Limbs and digits. Scales on limbs uni- to multicarinate, the
largest (on anterior thigh) as large or larger than ventrals, on digits
multicarinate. Digital dilations narrow, 21 to 22 lamellae under
phalanges ii and iii of fourth toe.

Compressed, no dorsal crest. Tail scales small, larger
Tail.
below, keeled except those under the base of tail smooth. Two
all
middorsal rows enlarged.

Size. The snout-vent length of the INDERENA type is 72 mm,
of CSJ 310 77 mm, and of LACM 72763 74 mm. The tail is broken
in all three specimens.
Co/or (aspreserved). Light purplish with brown markings
expanding at intervals on the dorsum (Type and LACM 72763) or
dark purplish with a dorsal zone brown with a black line on each
side (CSJ 310). Throat and belly very weakly spotted on a light grey
1985 A NOUS ANTIOQUIA E
Table I .
Comparison of Anuli.s antioquiae and A. fin hi.
aniioquiae fitchi
head
8 BREVIORA No. 482
ground (Type) or light purplish laterally smudged medially (CSJ

310). The limbs are very weakly banded, the tail darkish above,
lighter below. Dewlap white, blotched and spotted with intense
black.
Color in life. P. A. Silverstone has provided color notes for the
Type and LACM 72763: "Dorsum green with black median stripe
with paired light green spots. Venter brown with white spots. Dew-
lap red orange with black blotches and yellow border."
Ecology. Silverstone reports the two specimens collected by his
party "on ferns at the edge of forest next to pasture."
Comparisons. Anolis antioquiae is closest to recently described
Anolis fitchi Williams and Duellman. In both the female has a
blotched dewlap. In both there is a noticeable indication of a small
round parietal spine or convexity (Fig. 2) just anterior to the
enlarged median name scales. This small prominence is not evident
in other species of the group. Of the other recognized species of the
—
eulaemus subgroup, two A. ventrimaculatus and A. gemmosus —
entirely lack a dewlap in females, in the one female referred to A.
eulaemus the dewlap is much reduced, apparently with uniformly
dark skin. All three examined specimens of A. antioquiae lack an
interparietal; this scale is always present in A. fitchi and in A.
eulaemus, but sometimes absent in A. getnmosus and often %o\n A.
ventritnaculatus.
A. antioquiae
is unique one respect: the very sharp canthal
in
ridge projecting above the although an apparently

loreal region,
trivial character, has not been seen by me in any other species of the
group or indeed in any Anolis. Within the group the three subequal
short superciliaries are unusual. The median angular indentation of
the mental by postmental scales contrasts with the straight trans-
verse border seen at the mental gular contact in A. fitchi. The
conical flank scales often surrounded by granules are again very
different from other members of the group.
Although apparently allopatric to somewhat to

this species is
—
the north of — A.
eulaemus and A. ventrimaculatus, and although
only females are known (and the female reported as that of eulae-
mus is questionably so referred), there is no question of conspecific-
ity or even of very close relationship. Even with A. fitchi, very

distant in Napo in Ecuador and Putamayo in Colombia, despite
more resemblances, there is no demonstration of close affinity
except for position with the same subgroup of a species group.
1985 ANOI.IS ANTIOQVIAE 9
ACKNOWLEDGMENTS
Iam grateful to Jorge Hernandez of INDERENA, John Wright
and Robert Bezy of the Los Angeles County Museum (LACM).
and Marco A. Serna of the Colegio San Jose, Medellin. Colombia
(CSJ) for the loan of specimens. Laszlo Meszoly made the
drawings.
LITERATURE CITED
Wii I lAMs. E. E.. ANO W. E. Di II 1 MAN, 1984. Anolis finhi. a new species of the
Anolis acquatoiialis group trom Ecuador and Colombia, pp. 257 266. //; Seigel
R. A. CI at. (cds.). Vertebrate Ecology and Systematics A Tribute to Henry
Filch. Lawrence. Kansas.
B R E'S3 R a
iiseiiiiii of Comparative Zoology
us ISSN 0006-9698
NOTES ON PRISTIDACTYLUS (SQUAMATA: IGUANIDAE)

Richard Etheridge' and Ernest E. Williams^
Abstract. Pristidactylus is diagnosed as a genus of austral South American

anoloid iguanid lizards allied to Enyalius and possibly paraphyletic with respect to
Leiosaurus, Aperopristis, and Diplolaemus. West of the Andes Pristidactylus torqua-
tus occurs in forested coastal areas and the inland cordilleras of central Chile; P.
valeriae (new comb.) and P. alvaroi {new comb.) are restricted to disjunct patches of
relict forests to the north in Santiago Province. East of the Andes four species occur
Argentina. P. casuhatiensis lives in high, wet, rocky meadows atop the Sierra de
in la
Ventana in southern Buenos Aires Province, and P. achalensis occurs in a similar

habitat atop the Sierra de Cordoba in Cordoba Province. P. scapulatus is known
from arid, rocky habitats at disjunct localities in the cordilleras and precordilleras
from western Catamarca Province southward to Mendoza Province, and on isolated
basaltic mesetas at lower altitudes from Mendoza southward to Chubut Province. In
cordilleran and precordilleran populations adult females virtually lose the bold
the latter are not
juvenile pattern whereas those from the basaltics do not. Although
recognized nomenclatorially, the name P. araucanus is available for them. P.fascia-
tus, type species of the genus, lives in a narrow belt of sandy steppe from San Juan
Province southward to Rio Negro and extreme southern Buenos Aires Provinces. It
has frequently been incorrectly referred to as P. araucanus. Leiosaurus hardensis is a
synonym of P.fasciatus. A key to the species of Pristidactylus is provided.
INTRODUCTION
Pristidactylus a genus of austral South American iguanid
is
and Argentina, and an

lizards with a disjunct distribution in Chile
exceedingly complex taxonomic history. Fitzinger (1843: 58) de-

scribed Pristidactylus as a subgenus of Leiosaurus Dumeril and
Bibron, 1837, and designated Leiosaurus fasciatus Dumeril and
California 92182.
'Department of Biology, San Diego State University, San Diego,
^Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts
02138.
2 BRFVIORA No. 483
Bibron, 1837 as its type species. Boulenger (1885: 127) employed

Pristidactylus, but only for the species fasciatus. In this century, up
until 1976, fasciatus and all of the other species now referred to
Pristidactylus were variously placed in Leiosaurus, Urostrophus
Dumeril and Bibron, 1837 and Cupriguanus Gallardo, 1964. In 1976
Etheridge {in PauU, Williams, and Hall, 1976: 10) pointed out that if
Barrio (1969) was correct in placing Cupriguanus araucanus Gal-
lardo, 1964 in the synonomy of Leiosaurus fasciatus Dumeril and
Bibron, 1837 and if the latter were congeneric with Cupriguanus
achalensis Gallardo, 1964, type species of Cupriguanus, then Cupri-
guanus is a junior synonym of Pristidactylus Fitzinger, 1843. In this
work seven species are referred to Pristidactylus: achalensis Gal-
lardo, 1964; alvaroi Donoso-Barros, 1975; casuhatiensis Gallardo,
196^; fasciatus Dumeril and Bibron, 1837; scapulatus Burmeister,
1861; torquatus Philippi, 1861; and valeriae Donoso-Barros, 1966.
Pristidactylus in this modern sense has never had a proper diag-
nosis,nor have the species within it been adequately distinguished
from one another. We
plan a future, more formal review of Pristi-
dactylus and related taxa; we here propose only to provide a diag-
nosis and discussion of the genus as now understood, enough
description of the named formsto permit their recognition, and
lastly a key which arranged to display the relationships of the

is
species as we now understand them. These notes are based on exam-

inations of 221 specimens of Pristidactylus in European, North and
South American museums (see Acknowledgments), and field work
in Argentina by Etheridge in late January, February, and March
during the austral summer of 1983.
Pristidactylus Fitzinger 1843
1843 Leiosaurus (Pristidactylus) Fitzinger, Syst. Rept., Wien, 1: 16.— Type species
(by indication): Leiosaurus fasciatus Dumeril and Bibron 1837.
1845 Leiosaurus (Pentadactyius) Gray, Cat. Spec. Liz. Brit. Mus., London, 224. —
Type species (by monotypy and original designation) Leiosaurus fasciatus
D'Orbigny in Dumeril and Bibron 1837.
1885 Pristidactylus
—
Boulenger, Cat. Liz. Brit. Mus., London, 2: 127.
1964 Cupriguanus Gallardo, Neotropica, B. Aires, 10(33): 127.— Type species (sub-
sequent designation by Gallardo, 1967: Cupriguanus achalensis Gallardo
1964.)
1976 Pristidactylus Etheridge in PauU, Williams, and Hall, Breviora, Mus. Comp.
Zool., No. 441, p. 10.
1985 PRISTIDACTYl.US 3
Diagnosis. Pristidactylus is a member of the "anoloid" group of
iguanid genera, specified by the acquisition of endolymphatic sacs

that extend back between the parietal and occipital bones into the
dorsal neck musculature (Etheridge and de Queiroz, in prepara-
tion). Within anoloids Polychrus is considered the sister group
of
the remaining genera, the latter having lost femoral pores and
acquired a spinulate Oberhauchen with flat cell borders. Within this

lattergroup Pristidactylus together with Diplolaemus, Leiosaurus,
Aperopristis and Enyalius retain four pairs of sternal ribs and have
acquired a unique structure of the distal two or three subdigital
scales:each has a median longitudinal groove. Akhough grooved
distal subdigitals areobscure or lacking in forms with distinctly
multicarinate subdigital scales {P. fasciatus, A. catamarcensis, A.
paronae, E. leechii, and E. brasiliensis), their presence is considered
a derived characteristic of the group, their absence the resuk of loss.
This subgroup containing Pristidactylus forms the sister taxon to a
group formed by the remaining anoloid genera: Urostrophus, Ani-
solepis, Aptycholaemus, Chanmeleolis, Phenacosaurus,
Chamaeli-
norops, and Anolis, all of which have three rather than four pairs of
sternal ribs.
Relationships the genera that form the subgroup contain-

among
ing Pristidactylus remain unresolved, and a preliminary analysis
indicates that Pristidactylus itself may be paraphyletic. Neverthe-
less, Pristidactylus can be diagnosed within this group.
It differs
from Aperopristis, Diplolaemus, and Leiosaurus in having a black

bar within the antehumeral fold and marked sexual dichromatism;
itfurther differs from Aperopristis and Leiosaurus in having fully
autotomic caudal vertebrae and a different color pattern, and from
Diplolaemus in possessing pterygoid teeth. Pristidactylus differs
from Enyalius in having the caudal scales smooth proximally and
keeled distally (keeled throughout in Enyalius); slightly convex
rather than conical mesoptychials; wide, smooth, scarcely imbricate,
lamellarlike supradigitals rather than sharply unicarinate, rhomboi-
dal and imbricate supradigitals; the prefrontal-frontonasal part of
the skull roof rounded rather than flat; a more robust body (some-
what compressed in Enyalius); shorter limbs (adpressed hind limb
reaches to between the antehumeral fold and orbit in Pristidactylus,
4 BREVIORA No. 483
to the orbit or beyond in Enyalius)\ and a shorter tail (tail/ snout-

vent 0.73-1.60 in Pristidactylus, 1.60-3.30 in Enyalius).
Juveniles of Pristidactylus and Diplolaemus have a light grey or
yellowish background with bold, dark crossbands over the head,
body, tail, and limbs, with a characteristic horseshoe-shaped mark-
ing over the back of the head, and the crossbands of the back
usually constricted laterally. In both sexes with increasing size the
dark markings typically break up or acquire irregular borders.
Large adult male Pristidactylus become nearly unicolor except for a
conspicuous, black antehumeral bar, and have at most only a faint
trace of the juvenile pattern. The juvenile pattern, though altered,
remains bold in adult females,
except for the northern populations
of P. scapulatus. The juvenile markings show various patterns of
ontogenetic change in Diplolaemus, but never disappear entirely,
and are identical in large individuals of both sexes. The distinctive
butterfly or fleur-de-lis markings of Leiosaurus and Aperoptistis are

present at hatching and change little in either sex. Thus, marked
sexual dichromatism is a characteristic of Pristidactylus, shared
with Enyalius, but distinguishing it from Aperopristis, Diplolaemus
and Leiosaurus.
Boulenger in 1885 transferred Leiosaurus torquatus, and in 1889
Leiosaurus scapulatus, to Urostrophus Dumeril and Bibron, 1837;
type species U. vautieri Dumeril and Bibron, 1837. Gallardo (1964)
recognized that a number of differences exist between Urostrophus
vautieriand Leiosaurus scapulatus. He proposed the genus Cupri-
guanus to include scapulatus and two additional species from
Argentina described in the same work, araucanus from Laguna

Blanca, Neuquen Province, and achalensis from the Pampa de
Achala in Cordoba Province. He further suggested that Leiosaurus
torquatus probably should be transferred to Cupriguanus. Gallardo
pointed out that Cupriguanus differs from Leiosaurus (in which he
included Diplolaemus and Aperopristis) in having compressed and
pectinate infradigitals at the bases of the first, second, and third toes
of the hind foot. The pectinate scales on the inner border of the first,
second, and third toes, conspicuous in P. fasciatus and emphasized
by Gallardo (1964) as characteristic of Cupriguanus, are not uni-
formly developed in all species of Pristidactylus, and are least prom-
inent in P. casuhatiensis. Also, these scales are conspicuously
swollen, though not pectinate, in Diplolaemus, so that this feature is
1985 PRISTIDACTYl.VS 5
not as clear-cut as Gallardo thought. Gallardo also pointed out that

Pristidactylus differsfrom Urostrophus in having a shorter tail,
more and more scales between the supralabials and
labial scales
suboculars, no projecting scales around the eye, the trunk and tail
not compressed, and the fourth toe shorter. Gallardo's (1979) use of
the binomial Pristidactylus vautieri for Urostrophus vautieri was
unjustified and probably unintended. As indicated earlier, Urostro-

phus is a member of the sister taxon of the group containing Pristi-
dactylus, distinguished from the latter by the loss of a pair of sternal
ribs and the absence of grooved distal subdigital scales. Thus,
although not all of Gallardo's characters for Cupriguanus (= Pristi-

dactylus) are useful, Pristidactylus can be distinguished from Uro-
strophus as well as from other anoloid genera as indicated in the
diagnosis.
Cei (1973a) compared Cupriguanus with Leiosaurus, Aperopris-
tis, and Diplolaemus. His concept of Cupriguanus
was based upon
the Argentinian species, achalensis, scapulatus, casuhatiensis, and
"fasciatus" (—araucanus). Leiosaurus bardensis (—Pristidactylus

fasciatus) was included with belU and catamarcensis in
the genus
Leiosaurus, and Aperopristis was considered monotypic with the

bibro-
species paronae. Diplolaemus included the species darwinii,
nii, and leopardinus. He pointed out correctly that Diplolaemus
differs from the others in lacking pterygoid teeth. The subocular was
said to be elongate in Diplolaemus and Cupriguanus, represented by
an arc of small, subequal scales in Leiosaurus and Aperopristis;
however, we find this condition variable in both Diplolaemus and
Pristidactylus. The subdigital scales were said to be smooth in
Diplolaemus and Cupriguanus, unicarinate or multicarinate in Lei-

osaurus, and multicarinate in Aperopristis. The subdigital scales
are smooth in Diplolaemus and multicarinate in Aperopristis; how-
ever, in Pristidactylus they are smooth, indistinctly unicarinate or
multicarinate. Cei's description of the subdigital scales of A. paro-
nae and L. catamarcensis as having three to five distinct keels is
correct, but his statement that the subdigital scales of belli have one
to three keels is not entirely in accord with our observations. In 64
Rio Negro,
specimens of L. belli from Chubut, Mendoza, Neuquen,
and Santa Cruz Provinces we find the subdigital scales either
and
entirely smooth, as stated in the type description (Dumeril
Bibron, 1837), or with one, rarely two, indistinct keels, as mentioned
6 BREVIORA No. 483
by Koslowsky (1898). The distal caudal scales were said to be

smooth in Diplolaemus, but we find they are keeled in D. darwinii,
as first noticed by Stejneger (1909).
A precis of the recognized species of Pristidactylus follows:
Pristidactylus fasciatus (Dumeril and Bibron 1837)
1837 Leiosaurus fasciatus Dumeril and Bibron, Erpet. Gen., Paris, 4: 244. Type
—
locality: not specified,specimen shipped from Buenos Aires. Restricted type
locality (D'Orbigny, 1847): Carmen de Patagones, Buenos Aires Province,
Argentina.
1843 Leiosaurus ( Pristidactylus) fasciatus
— Fitzinger, Syst. Rept., Wien, 58. 1:
1845 Leiosaurus (Ptenodactylus) fasciatus — Gray, Cat. Spec. Liz. Coll. Brit. Mus.,
London, 224.
1885 Pristidactylus fasciatus
— Boulenger, Cat. Liz. Brit. Mus., London, 2: 127.
1896 Pristidactylus fasciatus

— Koslowsky, Revta Mus. La Plata, 7: 447.
1933 Leiosaurus fasciatus — Burt and Burt, Trans. Acad. Sci. St. Louis, 27(1): 30.
1968 Leiosaurus hardensis Gallardo, Neotropica, Buenos Aires, 14(43): 5, unnumb.
Fig.
— Type locality: Puelen, Cochico, Altos de Chochico, La Pampa Province,
Argentina.
1973 —
Cupriguanus bardensis Cei and Roig, Deserta, Mendoza, 4: 71.
—
1978a Pristidactylus bardensis Cei and Castro, Publ. Occ. Inst. Biol. Anim. Univ.
Nac. Cuyo, Ser. Sci., 5: 3.
Diagnosis. Pristidactylus fasciatus is unique within the genus in

having multicarinate subdigital scales and the tail less than 50% of
the total length.
Taxonomy. The type description oi fasciatus was based upon a

juvenile specimen (42 mm snout-vent) collected on the wall of the
fort at Carmen de Patagones in extreme southern Buenos Aires
Province, just north of the mouth of the Rio Negro. Dumeril and
Bibron (1837: 244) attribute the name to D'Orbigny, who collected
the animal and for a time observed it alive. in an
D'Orbigny (1847),
expanded description with notes on the live animal, described it as a
"charming species with all of the body a lively yellow, the top of the
head brown, two large, transverse bands on the back of the head, six
on the back all interrupted on the flanks. The upper part of the tail
is likewise ringed with black,
just as are the Umbs."
Dumeril and Bibron's (1837) description oi fasciatus is for the
most part applicable to juvenile individuals of all species now
referred to Pristidactylus, except that the subdigital scales are said
to be multicarinate, and the internal border of the foot has a dentic-
ulation, both variable in the genus, and the tail is shorter than in
Other forms. That the tail was mutilated was noted in the type
description, but the figure provided by D'Orbigny (1847) shows an
animal with a complete tail, and when measured on the illustration
the tail appears to be between 0.8 and 0.9 times the snout-vent
length.
Boulenger (1885) listed Pristidactylus fasciatus on the authority
of Dumeril and Bibron (1837), but inexplicably stated that the tail is
nearly twice as long as the head and body. Koslowsky (1896)
reported specimens under that name from Rio Negro and Neuquen,
the latter having subdigital keels completely lacking, and later Kos-
lowsky (1898), under the name Leiosaurus fasciatus, reported a
juvenile from Neuquen with subdigital scales tricarinate at the
extremities but otherwise smooth. In the latter the tail/ snout-vent
ratio was 1.18. It seems Koslowsky's specimens from
likely that
Neuquen form
are referable to the described from Laguna Blanca by
Gallardo (1964) as Cupriguanus araucanus rather than to P. fascia-
tus. In the former, the tail is longer than the snout-vent length and
the subdigital scales are smooth except for the grooved distal sub-
digitals. Koslowsky's Rio Negro specimen was not described. Kos-

lowsky's (1898) use of the binomial Leiosaurus fasciatus was fol-
lowed by Burt and Burt (1933) and Lieberman (1939).
Barrio (1969) compared the types of Pristidactylus fasciatus and
Cupriguanus araucanus Gallardo, 1964. He concluded that they
were synonyms because in both the juvenile pattern was one of
distinctive dark crossbands on the head, body, limbs, and tail; there
were pectinate scales on the infradigitals of toes 1, 2, and 3; and
measurements taken from D'Orbigny's (1847) figure yield a snout-
vent/ tail ratio of 0.89, only slightly larger than that of C. araucanus.
However, the distinctive juvenile markings are characteristic and
very similar in all species of Pristidactylus and Diplolaemus, most
other species of Pristidactylus have pectinate subdigitals on the foot,
and according to Gallardo's ( 964) figures and our own calculations
1
the snout-vent/tail length ratio in specimens from Laguna Blanca,

the type locality of araucanus, is greater than 1 .0. Barrio noted that
Gallardo had specifically rejected the synonomy of araucanus with
fasciatus because of the shorter tail in the latter, and that Gallardo
must have obtained his measurements from the illustration of
D'Orbigny, Although Barrio directly compared the holotypes of

both species he apparently failed to note the presence of distinctly
multicarinate subdigitals '\n
fasciatus, as contrasted with the smooth
8 BREVIORA No. 483
subdigitals of araucanus. The synonomy o{ araucanus w'lih fasciatus

was accepted by Cei (1973a, 1973b, 1975, 1979), Cei and Castro
(1978a, 1978b), and Cei and Roig (1973).
Gallardo (1968) described Leiosaurus bardensis from Cochico, La
Pampa Province and reported additional specimens from Ishigua-
lasto, San Juan Province, some distance north of the type localities
oi fasciatus and araucanus. He compared the species in detail with
Diplolaemus darwinii and D. bibronii, but not with any of the
species now referred to Pristidactylus. The species was recognized as
Cupriguanus bardensis by Cei and Roig (1973), and as Pristidacty-
lus bardensis by Cei and Castro (1978a, 1978b). Its distinctive fea-
tures, i.e., multicarinate subdigital scales, pectinate subdigitals on

the foot, and a tail shorter than head and body, are just those
characteristics which distinguish P. fasciatus from other species of
Pristidactylus. We have examined the types oi bardensis diXidi fascia-

tus and a dozen other specimens from Rio Negro, Mendoza, and
San Juan Provinces, and we find they agree in all essential details.
Distribution and Habitat. Pristidactylus fasciatus has been taken
at widely scattered localities between Ishigualasto (+30°10'S,
57°50'W) at about 1,700 m altitude in the north and the type locality
near sea level in the south. All records are within the Southern
Chaco Province, or Monte
(Cei, 1979). Cei and Castro (1978a)
stated that in theBermejo Basin of San Juan Province it occurs in
areas of sand dunes. In Mendoza Province the species lives on flat,
sandy steppe, between 1,000 and 1,750 m, and Gallardo
at altitudes
(1968) stated that at Chochico in La Pampa Province (type locality

of bardensis) it lives in loose sand, in small burrows of Microcavia
sp. constructed in mounds of sand that accumulate around the bases
of Prosopis alpataco. Other species of Argentinian Pristidactylus
appear always to be associated with rocky areas, and at the same
latitudes occur at higher altitudes. South of La Pampa Province
there are no specific locality records except for the type locality, but
Koslowsky (1898) reported the species from Rio Negro Province,

and we have seen a specimen in the Zoologiske Museum, Universi-
tetes K0benhaven also from "Rio Negro."
Pristidactylus scapulatus (Burmeister)
1861 Leiosaurus scapulatus Burmeister, Raise La Plata-Staat., Halle, 2: 523. Type

de Uspallata, near Uspallata, about 5,000 ft. altitude, Men-
locality: Sierra
doza Province, Argentina.
1861 Leiosaurus multipunctalus Burmeister, Reise La Plata-Staat., Halle, 2: 524.

Type locality: Sierra de Uspallata, near Paramillo, about 8,000 ft. altitude,
Mendoza Province, Argentina.
1889 Urostrophus scapulatus Bou\engeT, Proc. Zool. Soc. London, 144; PI. 15,
Fig. 2.
1941 Leiosaurus scapulalus—MxiWer, Zeitschr. f. Naturwiss, 94: 184.
1964 Cupriguanus araucanus—GaWardo, Neotropica, B. Aires, 10(37): 129; Fig. 2, 3
— Type locality:
Laguna Blanca, Neuquen Province, Argentina.
1964 Cupriguanus scapulalusGaWardo, Neotropica, B. Aires, 10(37): 128.
1976 Prislidactylus scapulalus Eiheridge, in Paull, Williams, and Hall, Breviora,
Mus. Comp. Zool., No. 441, p. 10.
Diagnosis. Pristidactylus scapulatus differs from P.fasciatus and

P. casuhatiensis in having perfectly smooth subdigital scales, from
P. achalensis in having the crowns of the posterior marginal teeth
slightly tapered with small anterior and posterior cusps, and from
the Chilean forms {torquatus, alvaroi, and valeriae) in having one or
two scales interposed between the nasal and lateral postrostral
scales.
Taxonomy. The descriptions of Leiosaurus scapulatus and L.

multipunctatus Burmeister, 1861 were accompanied by the descrip-
tion of a third species, Leiosaurus marmoratus, from several locali-
tieswest of Catamarca. Boulenger (1889) stated that scapulatus
represented the adult male, multipunctatus the adult female, and
marmoratus the juveniles of a single species, which he referred to as
Urostrophus scapulatus. MCiller (1928) pointed out that the types of
marmoratus belonged to the genus Phrynosaura Werner in Burger,
1907 (subsequently transferred to Ctenoblepharis by Cei [1974] and
then to Liolaemus [Cei, 1980]). Miiller (1941) later confirmed Bou-
lenger's synonomy o{ multipunctatus with scapulatus after compar-
ing the types at the Museum in Halle.
As presently recognized, Pristidactylus scapulatus is a species of

the high Cordilleras and precordilleras of western Argentina in the
provinces of Catamarca, San Juan and Mendoza (Cei, 1979). How-

ever,we now believe that specimens from the basaltic tablelands of
southern Mendoza, western Neuquen and southern Chubut pro-
araucanus and later to P.fasciatus (see
vinces, previously referred to P.
above discussion under fascial us), may be conspecific with P. scapu-
latus. In characters of scalation and proportions, and in the color
pattern exhibited by adult males, specimens from the cordilleras and

precordilleras cannot be distinguished from those from the basaltic
tablelands. In these populations as in all other forms of Pristidacty-
lus, the bold juvenile pattern becomes progressively more obscured
10
'
BREVIORA No. 483
in males,becoming faint or fading entirely, except for the black

antihumeral bars, in large adults.
Adult females from the basaltic tablelands do, however, differ
from those of the Cordilleras and precordilleras in the extent to
which they retain the juvenile pattern. Females 98 mm snout-vent
length and larger, from northwestern San Juan Province, and from
the Sierra de Uspallata and the vicinity of Paso de Choique in
western Mendoza Province usually have numerous scattered dark
brown spots on both upper and lower surfaces, but retain only a
faint indication of the juvenile crossbands (see Boulenger, 1889, Fig.
2). Those from the basaltic meseta around Volcan Payiin in south-
western Mendoza Province, from the vicinity of Laguna Blanca in
western Neuquen Province, and from Meseta Canquel in southern
Chubut Province, also usually have scattered dark spots, but retain
with little fragmentation or loss of contrast the bold pattern of
juveniles. Especially prominent is the dark, horseshoe-shaped mark
across the back of the head. Retention of a bold pattern in adult
females is other species of Pristidactylus, thus
characteristic of all
the loss of pattern in adult females in the Cordilleras and precordille-

ras is unique within the genus.
A question that we have not resolved to our own satisfaction is
how to treat this apparent step-clinal variation in the

ontogenetic
development of the female color pattern taxonomically. Our six
populations are widely separated from one another, except for Paso
del Choique and Payiin, where the step cline is abrupt. The pattern
of small, dark spots on a washed-out greyish background character-
istic of the Cordilleran females, like that of the males, matches very
well the mostly granitic rocks of their habitat, whereas the bold
female pattern is more cryptic on the dark brown, eroded substrate
of the basaltic mesetas. We may be dealing with relict populations
that are now no
longer and the situation appears com-
in contact,
parable to a chain of islands extending from north to south, so that

any taxonomic decision must perforce be arbitrary.
Serological comparisons of specimens from Uspallata, Choique,
Payiin, and Batra (near Payiin) show a high degree (>84%) of homo-
geneity (Cei and Castro, 1975). Future karyological, electrophoretic
or immunological studies may shed light on the problem. For now,
however, we follow a conservative nomenclature, formally recogniz-
ing only scapulatus, although in the key below we utilize adult
female color pattern and locality to separate the cordilleran and

precordilleran populations from those of the basaltics. If the latter
are eventually recognized taxonomically the name P. araucanus is
available (see discussion under P. fasciatus).

Distribution and The most northerly record is an
Habitat.
Catamarca Province
unspecified locality in the high cordilleras of
(Koslowsky, 1895). In northwestern San Juan Province at an alti-
tude of 3,800 m in the Reserva Provincial de San Guillermo
(29°18'S, 69°15'W) they are closely associated with granitic out-
crops and live in burrows under stones or at the bases of small
bushes, and appear to be active in the early morning and late after-
noon. They occur in similar habitats but at lower altitudes in the
Sierra de Uspallata (32°30-4rS, 69°00-06'W; 2,000-3,000 m) and
the vicinity of Paso de Choique (36°25-27'S, 69°25-45'W; 1,800-
2,000 m) in western Mendoza Province. Fewer than 50 km eastward
of Paso de Choique they occur on the basaltic meseta dominated by
Volcan Payun (36°26'S, 69°16-25'W; 1,800 2,000 m), where they
are also closely associated with stony areas and are active in the
morning and late afternoon. Likewise they occur in similar habitats

but at lower altitudes to the south, in the vicinity of Laguna Blanca
(39°03-45'S, 70°23-37'W; 1,200 m) in western Neuquen Province,
and the Meseta Canquel (44° 10-33'S, 68° 16-20'W; 900 m) in south-
ern Chubut Province. A single specimen is known from Puerto
Madryn on the coast in northeastern Chubut.
Pristidactylus achalensis (Gallardo)
1964 Cupriguanus achalensis Gallardo, Neotropica, Buenos Aires, 10(33): 132; Fig.
4. —
Type locality: Postade Pampa de Achala, Cordoba Province, Argentina.
1976 Pristidactylus achalensis
— Etheridge in Paul!, Williams, and Hall, Breviora,
Mus. Comp. Zool., No. 441, p. 10.
Diagnosis. Pristidactylus achalensis is unique within the genus in

having the crowns of the posterior marginal teeth rather strongly
compressed linguo-labially, flared in an anterior-posterior direction,
and deeply cusped, and in having an adult male color pattern of
uniform bright green with an immaculate yellowish belly.
and Habitat. The species
Distribution islimited to a small region
in western Cordoba Province, from the Pampa de San Luis
(3I°20'S, 64°46'W) in the north, southward at altitudes of 2,000 to
12 BREVIORA No. 483
2,500 m to Cerro Champaqui (31°59'S, 64°56'W). It is abundant on

the Achala, where the habitat is a high pampa with many
Pampa de
low rock outcrops that emerge from a wet meadow covered with low
herbaceous vegetation. Numerous small sinkholes full of rain water
dot the landscape. The bright green adult males are conspicuous
throughout the day. Unlike other members of the genus this species
isomnivorous, feeding on a variety of insects, including butterflies,
and flowers (di Tada et al., 1977a and 1977b). Its diet is reflected in
the species' unique tooth form. Gallardo (1967) reported that 130
individuals were obtained by two collectors in a period of three
hours on Cerro Champaqui in February.
Pristidactylus casuhatiensis (Gallardo)
1968 Cupriguanus casuhatiensis Gallardo, Neotropica, B. Aires, 14(43): 2; unnumb.

fig.
—
Type locality: Sierra de la Ventana, Buenos Aires Province, Argentina.
1976 Pristidactylus casuhatiensis
—
Etheridge in Paull, Williams, and Hall, Brevi-
ora, Mus. Comp. Zool., Cambridge, No. 441, P. 10.
Diagnosis. Pristidactylus casuhatiensis is unique within the genus

in having one or two indistinct keels on most of the subdigital scales,
the posterior marginal tooth crowns swollen, and in having an adult
male color pattern consisting of a green background with a fine,
dark reticulum, especially prominent on the sides; their venter is
immaculate yellow. The adult females have a brown background
color with obscure transverse bars.
Distribution and Habitat. The species is known only from Cerro
Tres Picos (38°27'S, 62° 12'W) and Cerro de la Ventana (38°08'S,
61°47'W) in the Sierra de la Ventana, at altitudes of about 1,000 m
in southern Buenos Aires Province. The habitat is one of a high
pampa with rocks, herbaceous vegetation and small sinkholes
flat
full of rainwater. P. casuhatiensis feeds on beetles, spiders, and
(Plagiodontes patagonicus) (Gallardo, 1970), the

terrestrial snails
latterperhaps correlated with its distinctive tooth form. According
to Gallardo (1970), the populations from Cerro Tres Picos and
Cerro de la Ventana exhibit slight differences in their characteristics.
Pristidactylus torquatus (Philippi)
1861 Leiosaurus torquatus Philippi in Philippi and Landeck, Arch. Naturgesch.,

Berlin, 27(1): 295. Type locality: in the neighborhood of Concepcion, Chile.
1985 PRISTIDACTYLUS 13
1861 Leiosaurus valdivianus Philippi in Philippi and Landeck, Arch. Naturgesch.,

Type locality: in the Province of Valdivia, Chile.
Berlin, 27(1): 298.
1885 Urostrophus /or^wa/wi— Boulenger, Cat. Liz. Brit. Mus., London, 2: 124.
1970 —
Cupriguanus torquatus Donoso-Barros, Bol. Mus. Nac. Hist. Nat. Chile,
Santiago, 49(24): 86.
1979 Pristidactylus torquatus
— Montecinos Espinoza and Formas, Herp. Review,
10(4): 121.
Diagnosis. Pristidactylus torquatus differs from the Argentinian

species in having the nasal scale in broad contact with the lateral
postrostral, and from the Chilean species P. alvaroi and valeriae in
having more than 140 scales around midbody, and generally smaller
and more numerous scales overall. Some individuals have an
inconspicuous and incomplete row of slightly enlarged middorsal
scales, but they are not continuous on the lumbar region as is
alvaroi.
Distribution and Habitat. Pristidactylus torquatus occurs in for-
ested areas of the coastal and inland Cordilleras of central Chile from
Curico Province (35° lO'S) southward to southern Llanquihue Prov-
ince (41°50'S),and on Isla de Chiloe (Donoso-Barros, 1966; For-
mas, 1979; Montecinos Espinosa and Formas 1979). The species is
arboreal, but also forages in dense brush (Donoso-Barros, 1966).
Pristidactylus alvaroi (Donoso-Barros) new comb.
1975 Cupriguanus alvaroi Donoso-Barros, Bol. Soc. Biol. Concepcion, 47(1974):

221; Figs. 3-5. —
Type locality: Cerro E! Roble, Santiago Province, Chile.
Diagnosis. Pristidactylus alvaroi differs from other members of

the genus in having a markedly compressed tail, a middorsal row of
enlarged scales continuous on the back, at least on the lumbosacral

region, and a different color pattern.
Distribution and Habitat. The species is known only from the
type where it occurs in relict clusters of Nothofagus

locality in Chile,
forest at the base of Cerro El Roble, Santiago Province (32°58'S,
71°0rW). It is said to be abundant (Donoso-Barros, 1975).
Pristidactylus valeriae (Donoso-Barros) new comb.

-
1966 Urostrophus valeriae Donoso-Barros, Rept. Chile, Santiago, 369: PI. 83.
Type locality: Alhue, Chile.

1975 Cupriguanus valeriae
— Donoso-Barros, Bol. Soc. Biol. Concepcion, 47(1974):
223.
14 BREVIORA No. 483
Diagnosis. Pristidactylus valeriae differs from Argentinian spe-

cies and from torquatus in having fewer, larger scales all over, with
fewer than 140 scales around midbody. It differs from alvaroi in
lacking a continuous row of enlarged middorsal scales on the back,
and in color pattern, and in not having a strongly compressed tail.
Distribution and Habitat. The species is known in Chile from
Nothofagus forests of the
Cordillera de la Costa near Alhue
(34°09'S, 71°24'W), and from Cerro El Roble (32°58'S, 71°01'W),
Santiago Province (Donoso-Barros, 1966). According to Donoso-
Barros (1966) the species is not as arboreal as P. torquatus, and is
often found in dense thickets and relict formations of Chusquea
quila.
Key to the Species of Pristidactylus
la. Scales larger and less numerous, i.e., nasal scale in broad contact with lateral
postrostral; nasal in narrow contact with anterior supralabial, or narrowly

separated from it by contact of lateral postrostral and anteromost loreolabial;
largest supraoculars equal to or larger than scales of the supraorbital semicir-
cles at narrowest width of frontal region; 6 to 8 scales in a horizontal line across
widest part of supraorbital region between superciharies and supraorbital semi-
circle; 8 to 16 scales bordering supralabials above Chilean species 2
lb. Scales smaller and more numerous, i.e., one or two scales interposed between
nasal and lateral postrostral; nasal scale well separated from anterior suprala-
bial by one or two scales. Largest supraoculars smaller than scales of the
supraorbital semicircles at narrowest width of frontal region; 8 to 10 scales in a

horizontal line across widest portion of supraorbital region between superciha-
ries and supraorbital semicircles; 15 to 23 scales bordering supralabials above. . .
Argentinian species 4
2a. Scales smaller and more numerous, i.e., scales around midbody more than 140;
scales bordering supralabials above more than 13; 13 to 22 scales across tem-
poral region in a straight line from postoculars to anterior margin of ear

torquatus
2b. Scales larger and less numerous, i.e., scales around midbody fewer than 140;
scales bordering above supralabials fewer than 13; 10 to 14 scales across tem-
poral region in a straight line between postocular and anterior margin of ear. .3 .
3a. Tail more strongly compressed; a continuous row of enlarged middorsal scales
on the lumbar region; dorsum greyish, venter yellow, iris red; tail conspicuously
banded in adults alvaroi
3b. Tail slightly compressed; a middorsal scale row absent on the lumbar region;
dorsum bluish, venter reddish, iris blue; tail in adult not conspicuously banded;
(antehumeral black mark absent ma\Q fide Donoso-Barros 1966).
in valeriae . .
4a. Subdigital scales distinctly multicarinate; tail less than 48% total length
fasciatus
1985 PRISTIDACTYLiS 15
4b. Subdigital scales smooth or with one or two weak keels; tail more than 48%
total length 5
5a. Subdigital scales with one or two weak keels; crowns of posterior marginal
teeth somewhat swollen, the anterior and posterior cusps absent or only faintly
indicated; adult male dorsal pattern a fine, dark reticulum, especially evident on
the sides casuhatiensis
5b. Subdigital scales perfectly smooth; crowns of posterior marginal teeth some-
what compressed, anterior and posterior cusps moderate or small but distinctly
present; adult male dorsum nearly uniform, a fine reticulation absent 6
6a. Crowns of posterior marginal teeth flared, more strongly compressed, with
larger anterior and posterior cusps; adult male uniform blue-green, green or
yellow-green above achalensis
6b. Crowns of posterior marginal teeth tapered, slightly compressed, with small
anterior and posterior cusps; adult male grey or yellowish-grey above, with or
without small, scattered dark spots scapulatus 7
7a. Adult female color pattern obscure, typically broken into scattered dark spots,
with dorsal cross bars faintly or not all indicated; no distinct horseshoe-shaped
mark on the back of the head Precordilleras and Cordilleras
7b. Adult female color pattern consisting of bold dark and light markings, a con-
spicuous dark horseshoe-shaped mark across the back of the head; throat and
belly usually with scattered dark spots
Payiin, Laguna Blanca, and Canquel
ACKNOWLEDGMENTS
We are grateful to the following individuals for permission to
examine specimens in their care: A. G. Kluge, The University of
Michigan Museum of Zoology; H. W. Greene, Museum of Verte-
brate Zoology, Berkeley; R. Sage, private collection; G. Peters and
R. Giinther, Zoologisches Museum der Humboldt-Universitat, Ber-
lin; J. Cranwell, Museo Argentine de Ciencias Naturales, Buenos
Aires; P. Alberch, Museum of Comparative Zoology, Harvard Uni-
versity; S. W. Braestrup, Zoologiske Museum, Universitetes K0ben-
haven; T. Cekalovic, Institute de Zoologia, Universidad de Concep-
cion; K. Klemmer, Natur-Museum Senckenberg, Frankfurt-am-
Main; W. Ladiges and H. Koepcke, Zoologisches Museum,
Hamburg; J. D. Williams, Museo de La Plata; W. E. Duellman,
Museum of Natural History, University of Kansas; A. G. C. Gran-

dison and E. N. Arnold, British Museum (Natural History); L. Cas-
tro, Institute de Biologia, Universidad Nacional de Cuyo, Mendoza;
R. Braun, Institute Argentine de Investigacienes de las Zonas Ari-
das, Mendoza; F. Achaval, Universidad de Uruguay; W. Hellmich
and U. Gruber, Zoologisches Sammlung des Bayerischen Staates,
16 BREVIORA No. 483
Munchen; J. Guibe, Museum National d'Histoire Naturelle, Paris;

C. McCoy, Carnegie Museum of Natural History; A. Leviton, Cali-
fornia Academy of Sciences; H. La Gilia, Museo Historia Natural
de San Rafael; R. Laurent, Fundacion Miguel Lillo, Tucuman; P.
E. Vanzolini, Museu de Zoologia da Universidade de Sao Paulo; U.
Parenti, Civico de Storia Naturale, Torino; J. Eiselt and F.
Museo
Tiedeman, Naturhistorisches Museum, Wien; and G. Zug, National
Museum of Natural History, Washington.
Funds for travel in the United States and field work in Argentina
were provided by a grant from the National Geographic Society
Committee on Research and Exploration. For providing laboratory
facilities and access to field vehicles we wish to thank Eng. Rolando
Braun, Director of the Instituto Argentino de Investigaciones de las
Zonas Aridas, Mendoza, and Dr. Oswald H. Sala, Director of the

Centro Nacional Patagonico in Puerto Madryn. We also thank Don
Eleodoro A. Sanches, Jefe Division de la Fauna of San Juan Prov-
ince for permission to work in the Reserva Provincial de San
Guillermo.
Dr. Jose M. Cei for
We especially wish to express our gratitude to
his tireless efforts to make our field work
Argentina a success, in
and to Fernando Videla and Jorge D. Williams for their assistance

in the field.
LITERATURE CITED
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. 1889. On some specimens of lizards in the Zoological Museum of Halle
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. 1978a. Datos preliminares sobre las componentes de la herpetofauna de
la Provincia de San Juan. Pubis Occ. Inst. Biol. Anim., Univ. Nac. de Cuyo,
Mendoza, Ser. Cien., 5: 14.
. 1978b. Atlas de los vertebrados inferiores de
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Cei, J. M., and G. V. Roig. 1973. Fauna y ecosistemas del Oeste arido argentino.
I.
Reptiles de la Provincia de Mendoza. Deserta, Mendoza, 4: 69 91.
Di Tada, I., R. Martori, B. Kutner, and A. Ocana. 1977. Reproductive cycle
and feeding behavior of Cupriguanus achalensis (Sauria:Iguanidae). Herp. Rev.,
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Di Tada, I., R. Martori, B. Kutner, A. Ocana, and M. Beatriz.
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tofauna endemica de de Achala (Cordoba, Argentina) I Reun. Ibero-
la Pampa
amer. Zool. Vert. La Rabida, 1977: 493 512.
Donoso-Barros, R. 1966. Reptiles de Chile., Santiago, xclvi + 458 pp.
. 1970. Catologo herpetologico chileno. Bol. Mus. Nac. Hust. Nat. Santi-
ago, 31: 49 124.
. 1975. Nuevos reptiles y anfibios de Chile. Bol. Soc. Biol. Concepcion,
48(1974): 217-229.
D'Orbigny, A. 1 847. Voyage dans I'Amerique meridionale . . .
pendant les annees
1862. . . 1833. Paris, 5(1): 1-12; Pis. 115.
DumCril, a. M. C, and G. Bibron. 1837. Erpetologie generale ou Histoire
Naturalle complete des reptiles. Paris, 4: 1-572.
FiTZiNGER, L. 1843. Systema Reptilium. Fasciculus primus, Amblyglossae. Wien,

vi + 106 pp.
FoRMAS, J. R. 1979. La herpetofauna de los bosques temperados de Sud Amer-
ica, pp. 341 369. In W. E. Duellman (ed.). The South American herpetofauna:
its origin, evolution and dispersal. Mus. Nat. Hist. Univ. Kansas, Monog., 7:
1-485.
Gallardo, J. M. 1964. Los generos "Urostrophus" D. et B. y "Cupriguanus"
gen. nov. (Sauria, Iguanidae) y sus especies. Neotropica, Buenos Aires, 10(33):
125 136.
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1967. A proposito del genero Cupriguanus Gallardo (Sauria, Iguani-

dae). Neotropica, Buenos Aires, 13(41): 56.
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Aires, 12: 67-68.
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pp. 299-307. In W. E. Duellman (ed.). The South American herpetofauna: its
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London, xxviii + 289 pp.
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Argentina) recogidos durante los meses de Febrero a Mayo de 1895. Rvsta Mus.
La Plata, 6: 357-370.
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Rvsta Mus. La Plata, 7: 447-457.
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argentinos. Rvsta Mus. La Plata, 8: 161-200.
LiEBERMANN, J. 1939. Catalogo sistematico y zoogeografico de los lacertillios
argentinos. Physis, 16(48): 61-82.

MoNTECiNOS EsPiNOSA, R. M., AND J. R. FoRMAS. 1979. Distribution of the
Chilean lizard Pristiciactylus torquatus. Herp. Rev., 10(4): 121-122.
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. 1941. Ober die in der Sammlungdes Zoologischen Institutes der Univer-
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Paull, D., E. E. Williams, and W. P. Hall. 1976. Lizard karyotypes from the
Galapagos Islands: chromosomes in phylogeny and evolution. Breviora, Mus.
Comp. Zool., No. 441, pp. 1-31.
Philippi, R. a., and L. Landeck. 1861. Neue Wirbelthiere von Chile. Arch.
Naturgesch., Berlin, 27(1): 289-301.

Stejneger, L. 1909. Batrachians and Reptiles, Part U, pp. 211-224. In W. B.
Scott (ed.), 1905 1911; Reports of the Princeton University Expeditions to
Patagonia, 1896-1899, Vol. Ill, 1, Zoology, Princeton, xiii + 374 pp.
B R Ej;SSI O R A
us ISSN 0006-9698
MALE AGGRESSIVE BEHAVIOR IN A PAIR

OF SYMPATRIC SIBLING SPECIES
Jonathan B. Losos'
Abstract. encounters were staged between adult male Anolis mar-

Intra.specific
lanoi and A. cyhoie^ order to describe their aggressive behavior. The major
in
component of stationary behavior is the dewlap display, accompanied by various

static and dynamic modifiers. Advancing behaviors include moving toward an oppo-
nent, threatening attack, and actual biting. The agressive behavior of /I. nwnanoi is
progressive and hierarchically structured. By contrast, the behavior of A. cyhotes is

less elaborate and ritualized than that of its sibling and lacks progressive structure.
INTRODUCTION
Defined simply as "pairs or even larger groups of related species
which are so similar that they are considered as belonging to one
species until a more satisfactory analysis clears up this mistake,"
(Mayr, 1942) sibling species have been discovered at an increasing
pace as the level of systematic analysis has become finer, passing
from examinations of morphological characters to investigations of
molecular properties.
The different patterns of distribution of sibling species raise inter-
esting ecological questions. On one hand, sibling species may be

completely allopatric, which raises questions more about the evolu-
tionary forces operating to produce such species than about the
'Museum of Comparative Zoology. Harvard University, Cambridge, Massachusetts

02138. Present Address: Museum of Vertebrate Zoology and Department of
Zoology. University of California, Berkeley, California 94720.
2 BREVIORA No. 484
ecological relationships between them. On the other hand, parapat-

ric and sympatric sibling species distributions bear directly on eco-
logical theory. In theory, two species occupying the same niche

cannot long coexist unless differences sufficient to lessen competi-
tive pressures evolve. The parapatric ranges of many sibling species
pairs or groups attest to the difficulty such species have in coexisting
(e.g., the members of the Rana pipiens complex in North America

[Pace, 1974] and the Anolis chlorocyanus [Williams, 1965] and A.
r/r6>/Y//7 in Hispaniola). One would expect
[Schwartz, 1974] groups
that sympatric sibling specieswould exhibit differences in mor-
phology, physiology, or behavior that would permit coexistence.
Indeed, resource partitioning of some kind is observed in almost all
known cases of sympatric sibling species (Mayr, 1942; Schoener,
1977).
In this regard, the curious case of the Hispaniolan sibling species
Anolis cyhotes and A. marcanoi warrants attention. Anolis cybotes,
the larger of the two, is found throughout the island, while A.
marcanoi is restricted to a small area in the southwest portion of the
Dominican Republic (Williams, 1975). The two are classic sibling
species, consistently separable morphologically only by the color of
the dewlap of males and the throat of females: Anolis marcanoi has
a red dewlap or throat, while that of A. cybotes is usually either
white or yellow, depending on locality (Williams, 1975). Electro-
phoretic studies by Webster (1975), however, clearly reveal that the
two are distinct species. Table provides a review of differences
1
between the species.

Ecologically, the species are just as similar. Unlike the cases of
no apparent differences in physi-
sibling species already discussed,
ology or microhabitat preference have been discovered. They are
patchily sympatric throughout the range of A. marcanoi and have
been noted on adjacent fenceposts in some areas (Hertz, 1980),
while in other locations only one species is present. Hertz suggests
that A. marcanoi may be better adapted for hotter, more open
microhabitats, but no consistent differences have been noted in the
field(Williams, 1975; Hertz, 1980; personal observation^
Atissue is the manner in which these species interact. Are the
differences between the species in dewlap color sufficient to allow
species recognition, or are there also behavioral differences of
importance? If species recognition is highly accurate, do the similar-
1985 /lA'O/./SSIBIINCi SPECIES BEHAVIOR
Table I. Differences between Anolis marianoi and Anolis cvhotes.*
A. cvhotes A. nwrcanoi
range widespread throughout Peravia Province, south-

Hispaniola west Dominican Republic
dewlap color normally white to yellow, red
except some populations on

the extreme end of the
southwestern peninsula in
Haiti, which are red
scale characters middorsal and midsacral middorsal and midsacral

scales sometimes enlarged scales rarely enlarged
maximum snout- 81 mm 65 mm
vent length
heat tolerance:
Experimental 34.3° C 35.7° C

Voluntary
Maximum
Critical Thermal 38.4° C 40.5° C
Maximum
After Williams. 1975 and Hertz, 1980.
itiesbetween the species lead to resource competition and interspe-

cific aggression? The goal of this study is to provide a thorough
description of intraspecific male aggressive behavior in the two spe-
cies. In themselves, reports on the intraspecific male aggressive
behavior of both species are important because detailed characteri-

zations of the components and progression of male aggressive
behavior in Anolis are not common (the only reports of comparable
detail are Greenberg and Noble, 1944, and Greenberg, 1977, on A.
carolinensis and Jenssen 1979a and 1979b on A. opalinus). Furth-
ermore, in order to analyze interspecific interactions, it is first neces-
sary to understand the dynamics of intraspecific behavior. This
knowledge then may be used as a control with which interspecific
aggressive behavior can be compared.

In January 1983, adult male A. marcanoi 'were collected past the
first Recodo road, approximately 6 to 10 km north of
ford on the El
4 BRFVIORA No. 484
Bani, Peravia Province, Dominican Republic. Adult A. cybotes

were collected at that locality and in Santo Domingo, Dominican
Republic. The lizards were housed in terraria in the Biological
Laboratories, Harvard University, and provided with perches, small
potted plants, and a soil substrate, and offered several crickets one
to two timesa week and water three to four times a week. Room
temperature was maintained at approximately 26 to 30°C. Individu-

als were separated from other males in the same terrarium by card-
board partitions.
Encounters between two conspecific males were staged between 23
March and 16 December 1983. Lizards were placed in the parti-
tioned halves of either a 4! X 22 X 20 cm (the first 14 A. mananoi
encounters) or a 74 X 32 X 29 glass terrarium (the last 12 A. mana-
noi and all 19/1. cybotes encounters) covered with a mesh lid and
provided with perches in each half of the terrarium. Lizards were
left undisturbed in the terrarium for a period ranging from 7 hours 1
to four days, but usually lasting two days. Water was only provided
when an encounter had to be postponed, forcing the period to
exceed two days. Lizards were allowed to acclimate in order to gain
familiarity and enhance territorial defense.
Approximately 15 to 30 minutes prior to an encounter, a 250 W
infrared lightwas switched on, warming the terrarium to 25 to 32°C
and markedly increasing the activity level of the lizards. Tempera-
ture in the room in which the encounters were staged was approxi-
mately 20° C.
Encounters were initiated by lifting the partition. All lights in the
room were extinguished, except two 60 W
desk lamps pointed
directly into the terrarium which the display arena and made it
lit
difficult for the lizards to see observers in the room. Encounters

were videotaped on a Panasonic NV-8050 time lapse recorder with a
MTl-65 video camera and a Canon 16-100 mm
TV zoom lens and
then analyzed at normal and slow-motion speeds on a Panasonic
WV-5350 video monitor. Encounters normally lasted 45 minutes,
but were terminated when one lizard tried repeatedly to escape from
the other, and were extended when the lizards were still intensely
interacting at the end of the alloted time.
Twenty-one A. marcanoi and ten A. cybotes were used in these
encounters. Lizards were paired randomly, with the constraints that
they had not been housed in the same terrarium, that they had no
1985 /1AY;//.SSIB1 lN(i SPFCIES BEHAVIOR 5
prior experience with each other (two exceptions), and that there
was no more than a 5 mm
difference in snout-vent-length.
RESULTS
The aggressive behavior of the Hzards can be divided into two
categories: stationary and advancing behavior. In stationary behav-
ior, the Hzard remains in one area and primarily moves its head,
bobbing with or without extension of the dewlap. Several modifiers

of this display, either morphological changes or movements of body
parts, are correlated with aggressive intensity, particularly in A.
marcanoi. The level of intensity oi A. marcanoi could be judged by
the correlated appearance of behavioral and morphological re-
sponses. As
displays progressed, lizards generally displayed a
sequence of actions and display modifiers, adopted a more threaten-
ing posture with body and head raised and apparent size maxim-
ized,and became more active. By contrast, in A. cyhotes there was
littlecorrelation between the appearance of particular aspects of
aggressive behavior. Thus, level of intensity was more difficult to
gauge.
In advancing behavior, the lizard moves toward its opponent. In
the early, less intense stages, the lizard makes various intention
movements, while later stages involve lunging and biting.
The male aggressive behavior of the two species differed in two
major respects. First, though elements of both stationary and
advancing behavior were displayed by A. cybotes, many of the
components of these behaviors were absent or much less elaborate
than those exhibited by A. marcanoi. Second, the ordered appear-
ance of progressively more aggressive behaviors, so integral a part
of male-male interactions of A. marcanoi, was absent in the behav-
ior of A. cybotes. In most cases, particularly among stationary be-
haviors, any particular behavior did not necessarily follow any
other particular behavior, nor did particular behaviors connote dif-
ferent levels of aggression.
Stationary Behavior
The primary stationary display of the lizards was the dewlap
display, composed mainly of head-bobbing and dewlap extension.

6 BRFVIORA No. 484
Dewlap Display
Anolis mananoi
There were generally three levels of intensity of the display exhi-
bited by A. mananoi. Intensity is best correlated with the degree to
which the lizard elevated its body above the substrate.
The low level dewlap display, often the first display the lizard
would make in an encounter, involved little body elevation. The
body occasionally remained completely in contact with the sub-
strate, but usually the anterior portion of the body was slightly lifted
by extending the front limbs, holding them out to the side in the
typical reptilian stance (Fig. la). The was that the forequar-
result
ters were raised several mm off the substrate. The snout was also
angled upward at 30 to 45 degrees to the ground. The head usually
was kept in the vertical plane of the body. During the process of
raising the body and snout, the dewlap usually was extended com-
pletely with several rapid head bobs. On a number of occasions, the
forebody distinctly raised first, and then the dewlap extended, but
the two usually occurred simultaneously.
This display occurred primarily at the start of an encounter,
shortly after the partition had been raised, though sometimes it was
omitted altogether. Occasionally, when an encounter had proven
indecisive and the lizards had calmed, they would revert to this level.
The intermediate dewlap display involved an increase in body
elevation. Generally, it took one of two forms: either the forelimbs
were completely extended with the snout pointed at a very high
angle (75 to 90 degrees), giving the whole body a very steeply sloped
configuration, or all four legs pushed up, with the hind legs out to
the side, elevating the whole body off the substrate (Fig. lb). The
tail was then usually also lifted, either held rigid and completely
elevated, or slightly arched with the posterior half dropping to the

substrate. The tail was also on occasion held higher than the body or
was even curled back above it. The
dewlap was extended completely
in this display; the snout's higher angle and the higher elevation of
the forequarters (at least a little higher in the second variation than
in the low level dewlap display) allowed the dewlap to be lifted
completely off the substrate.

In the highest level of intensity display, clearly distinct from the
first two, the entire body was elevated high off the substrate (Fig.
Ic). In ihe full elevation display, the forelimbs were held under the
1985 A \()l IS S\li\ INCi SPFCIFS HFHAVIOR
(b)
Figure I. Male Anolis mananoi performing (a) low-level, (b) intermediate, and
elevation displays. Display modifiers shown here, such as crest erection and
(c) full
head spot darkening, were noted in all three display levels, through they were more
common in the more aggressive levels.
8 BREVIORA No. 484
body and almost completely straightened, while the hindlimbs were

pushing up greatly either sprawled out to the side or also under the
body. The tail was usually rigidly held straight back or even held
higher than the body. The snout was very high, and the dewlap was
well off the substrate.
Several particular aspects of the display varied independently of
these levels, though in some cases relationships were noted. The
most variable, and perhaps most significant, component of the dis-
play involved the dynamics of dewlap extension. In general, the
degree of extension varied as a function both of time elapsed and
intensity of the encounter. The dewlap was usually completely
extended and maintained or pulsed (retracted and reextended) to a
barely perceptible extent at the beginning of encounters and when a
high level of intensity was observed. As encpunters progressed into
series of alternating exchanged displays, dewlap extension tended to
decrease both extent and duration. Often, the dewlap would only
in
be pulsed out briefly at the end of a series of bobs; the amount of

dewlap extended varied similarly, sometimes only revealing a slight
bit of the dewlap, and sometimes not at all. If intensity increased,
for example as a result of an advance by one of the lizards, dewlap
extension would increase in subsequent bobbing.
A no/is
cy boles
Anolis cvhotes'only has one level of dewlap display, comparable
to the low level dewlap display of A. marcanoi. A lizard displayed
with posterior half in contact with the substrate, while its anterior
its
was raised a variableamount by extension of the forelegs (Fig. 2),

either out to the side or directly under the body. The snout also was
angled upward to a variable degree. On rare occasions, usually when

itwas on the ground, a lizard displayed with all four legs out to the
side, pushing its body up off the ground.
The dewlap was usually extended, at least in the initial bout of
displaying, with concurrent head-bobbing. The amplitude, number,
and even presence of bobs varied. On occasion, usually later in an
encounter, the dewlap was extended and retracted without any head
movement at all. As with A. marcanoi, sometimes the dewlap was
extended and then retracted, while other times it was maintained at
full extension.
1985 /1AY;//.S SIBI l\(i SPECIES BEHAVIOR
Figure 2. Typical display posture of male Anolis cyhoies. The forequarters are
elevated, but the hindquarters are firmly planted on the substrate. The nuchal crest is
occasionally erected more fully, and the dorsal crest is sometimes apparent. The
dewlap often is extended more fully.
Head Bobbing
Anolis nmrcanoi
Though the bobs of the head usually were part of the dewlap
extension process, similar to the fanbob of A. aeneus (Stamps and
Barlow, 1973), in later stages of the encounter, especially when the
lizards were alternating displays, the bobs were increasingly empha-
sized and independent of dewlap extension. A series of 2 to 10 bobs,
either of the normal amplitude or greatly exaggerated with several
times that amplitude, were performed without dewlap extension, or
as a distinct precursor to extension. At other times, the dewlap was
pulsed out with exaggerated jerking bobs, similar to the jerkbob of

A. aeneus. The lizards sometimes bobbed several times at full dew-
lap extension, with a slight pulse of thedewlap during each bob and
between. This usually occurred when the display
slight retractions in
intensity was high. In several cases, a subordinate lizard raised and
lowered its head extremely slowly with large amplitude and no dew-
lap extension. Thiswas usually repeated several times.
Jenssen (1983) found that A. cybotes performed only one stereo-
typed head bobbing pattern, in contrast to the greater repertoires of
other Anolis (Jenssen, 1977, 1978). A detailed investigation was not
conducted, but from an analysis of the diplays of several lizards, it
appears that A. marcanoi has at least two distinct display types.
However, neither the extent of variation nor the degree of stereo-
typy of the head-bobbing patterns were determined. It is possible
10 HRFVIORA No. 484
also that A. mananoi has several other head-bobbing patterns that

were not disco\ered. No relationship between level of dewlap dis-
play and head-bobbing pattern was found.

in the first pattern, equivalent to the signature display (Stamps
and Barlow, 1973; Jenssen, 1978), the head was bobbed once or
twice with great amplitude, often with a pause in between bobs, and
then rapidly bobbed another 7 to 17 times with the snout remaining
elevated (Fig. 3). The dewlap began to expand between bobs six to
nine, at which time the bobbing would begin to slow. Several times
this display was prefaced with a lowering and raising of the head,
akin to. though slower than, the "head-dipping" of A. cybotes
(Jenssen. 1983). Jenssen (personal communication) pointed out the
close similarity between this display pattern and the sole one
reported for A. cybotes. Only the number of initial large-amplitude

bobs— one or two in a /I. mananoi, three in A. cybotes —
distinguishes the two, attesting to the close relationship of the two
species. This pattern was observed most often early in encounters,
though it was also seen sometimes at high levels of intensity. This
description must be considered tentative, however, because it is
based on only four displays of three lizards.

A second display, probably functioning as a challenge display
(Jenssen, 1978), was often seen late in encounters, especially when
lizards were fairly close to each other and alternating displays. It
was characterized by a large amplitude dip of the head, a pause, and
then three or four rapid low amplitude bobs with the head not quite
F-igurc y. Sample display-action pattern graph ot the first head-bobbing pattern

ot Aii()li\ manunoi. The line represents approximate elevation through time. The
shaded region represents dewlap extension through time. This pattern was sometimes
preceded by a dip ol the head.
1985 /4/VO//.S SIBI lN(i SF'ECIES BFHAViOR il
dropping back to its Sometimes, there was a

original level (Fig. 4).
pause after the bob. with the head often not returning to its original
level. Usually the dewlap was brought out only at the end of the
bobbing sequence, if at all. This description is based on an analysis

of nine displays by four lizards. It did seem to be fairly stereotyped,
however, and was observed in the displays of many of the other
lizards.
Intensity was reflected in the length and magnitude of dewlap

display and head-bobbing. More intense encounters tended to have
longer displays with greater number of bobs and pulses and greater
dewlap extension. The pace of the displays was also faster in more
intense displays. If these intense encounters did not immediately
progress into more direct aggressive action, however, displays

tended to become shorter and less animated, often devolving into
the alternating bouts of medium speed bobbing mentioned above.
Anolis cybotes
Because Jenssen has already extensively examined the stereo-
typed head-bobbing patterns of A. cybotes, they were not investi-
gated here. Two distinct methods of bobbing were noted in A.
cybotes. In one, by far the more common in A. marcanoi, the whole
head was bobbed, passively moving the dewlap along with it. but
without greatly changing its amplitude. This method of bobbing
produces the head-bobbing display action patterns so extensively
studied in Anolis.
Figure 4. Sample of the second head-bobbing pattern of Anolis marcanoi.

Though not indicated here, the dewlap was occasionally extended at the end of the
display.
12 RRFVIORA No. 484
—
By contrast, dewlap bobbing rarely exhibited by A. marcanoi,
but —
commonly performed by A. cyhotes primarily involved mov-
ing the dewlap up-and-down by raising and lowering the posterior
portion of the hyoid, causing a large amplitude change in the dew-
lap, but moving the head only slightly. This bobbing type was usu-
ally preceded by the first type and appeared when the lizard was
displaying intensely. Only the first type was involved in dewlap
extension.
The quick inverted head bob or "head dip" noted in wild A.
cyhotes by Jenssen was observed on a number of occasions. His
observation that head dips were often performed independently of
dewlap displays was confirmed, though no long series of head dips,
which Jenssen also reported, were noted in these experiments.
Static Modifiers
Several —
morphological responses Jenssen's static modifiers
(Jenssen and —
Hover, 1976; Jenssen, 1977, 1978) were also indica-
tive of level of intensity, particularly in A. marcanoi.
A no/is marcanoi
Both the nuchal and dorsal crests were often erected during dis-
plays (Fig. lb. and Ic). Often, they were erected, retracted, and
erected again several times in an encounter. The nuchal crest always
appeared before and disappeared after or simultaneously with the

The crests often at first were erected fully, but then
dorsal crest.
retracted to only a fraction of their full size as the encounter
continued.
The presence of fully erected crests indicated a high intensity
state,but their absence did not imply the converse. When the dis-
tance separating the lizards was not great, actions more aggressive
than the /w// elevation display were generally accompanied by full
crest erection, as were most /t/// elevation and some intermediate
elevation displays. Crests were noted less frequently when the
lizards were displaying from across the large terrarium. Lizards that
appeared clearly subordinate — indicated by display level, activity,
posture, and, retrospectively, by the outcome of the encounter (see

below) — were much slower to raise their crests and maintained them
generally for a shorter period.

1985 .4.\Y;//.S SIHI INCi SF'FCIFS BFHAVIOR 13
Similarly, the appearance of a dark spot on the side of a lizard's

head correlated broadly with intensity. Between the eye and the
tympanic opening, a black circular spot slightly larger than the
tympanic opening would appear when a lizard was aroused (Fig.
Ic). A similar well-defined dark patch with a similar function has
been reported for A. carolinensis (Greenberg and Noble. 1944;
Greenberg, 1977), but in that species the patch is rectangular and
directly postorbital without intervening undarkened scales. The
appearance of the spot in both species seems clearly a function of
arousal; lizards handled during transfer to and from the experimen-
tal arena almost invariably exhibited it and often tried to bite. Many
other Anolis display an irregular darkening in that region when

aroused (G. C. Mayer, personal communication; personal observa-
tion). Dominant lizards were much more likely to exhibit the spot
than subordinate ones, which rarely displayed it except at the high-
est levels of intensity, and only then when putting up resistance
instead of fleeing. As with the nuchal and dorsal crests, black spots
were usually apparent, especially on dominant lizards, at high levels
of intensity, and sometimes at lower levels, ranging from low level
dewlap displays io full elevation displays. Darkening of the spot
—
almost invariably was preceded by erection though not necessarily
full erection —
of the crests. As with crest erection, the spot appeared
much more frequently when the distance separating the lizards was
not great.
Body orientation and appearance also were important compo-
nents of the display. Lizards, especially dominant ones, increased
their apparent body size several ways. The apparent size of the head
was increased by erecting the nuchal crest, engorging the head, and
lowering the hyoid apparatus of the throat. Such enlargement is a
common aggressive response in iguanids (Greenberg and Noble,
1944; Carpenter. 1967). Presumably, dewlap extension has the same
effect of making the lizard appear larger. When retracted, the dew-
lap remained apparent, ranging in size from a slight rim to a fairly

large crescent, making the head region appear larger. The body was
also made apparently larger by expansion of the dorsal crest and
lateral compresion of the body. Subordinate lizards, as well as dom-
inant lizards displaying after winning an encounter (indicated by
quiescence or flight of their opponents) rarely attempted to increase
their apparent size.
14 BRFVIORA No. 484
Lizards also attempted to assume a position that would expose

the greatest part of their bodies to the other lizard, increasing their
apparent size. When both lizards were displaying, the most common
position was a parallel alignment, each lizard exposing its broadside
to the other. As with many other Anolis (e.g.. Carpenter, 1965,
1967), when not aligned parallel, one or both lizards (usually the one
with less apparent broadside exposure to the other) would turn its
head, so that the dewlap when extended would be parallel to the
other's head and thus appear as large as possible. Maximizing
apparent size was most obvious when one lizard was on a perch
while the other was nearby on the ground. In these instances, dis-
playing in an upright position on the branch would not present as
largean image as possible to the lizard on the ground. Invariably,
the lizardon the branch would display on the side of the branch.
The closer the lizard on the ground was to the branch, the greater
would be the angle from the vertical of the lizard on the branch.
When the lizard on the ground was directly underneath the branch
of the other lizard, the latter would arrange itself horizontally, at a
90° angle from the upright. By contrast, a lizard displaying to
another lizard also on a perch would always display directly upright
on the perch.
Anolis cy botes
As with A. marcanoi\ several morphological responses were noted
as part of the male-male interactive behavior of A. cyhoies. In the
latter species, however, lack of a particular static modifier — or of all
static modifiers — did not necessarily correlate with low levels of

intensity.
Like A. marcanoi, A. cybotes possesses both a nuchal and a
dorsal crest. The crests appear to be smaller in A. cybotes and are
fully erected much less frequently. Often, the dorsal crest was visible
only as a low ridge along the lizard's back. Crest erection appears to
serve the same function in both species, increasing apparent size and
indicating heightened level of intensity. In the eight encounters in
which dominance could be determined (with A. cybotes, dominant
lizards could only be determined post facto; dominance could not be
determined by the presence of hierarchically higher displays and
modifiers as it could with A. marcanoi), five of the dominant lizards
displayed crest erection. In two of the three other instances, no other
1985 ,-l,\r;//.S SIBi INCi SI'F.CIES Bt HAVIOR 15
preliminaries to a direct attack occurred. Only two of the eight

subordinate lizards displayed crest erection, both in encounters in
which the dominant lizard also displayed crest erection.
Anolis cyhotes does not have a well-defined circular head spot
oi A. mananoi. It can, however, change skin hue (as can A.
like that
marcanoi): often, dominant lizards became lighter. An ill-defined
darkening of the skin in the postorbital region of the head, where

the dark spot in A. marcanoi is located, was noted on several occa-
sions, usually when the lizard was aroused by another male or was
handled. Only in one lizard, however, was anything approaching a
circular spot apparent.
As with A. marcanoi. A. cyhotes attempted to increase its appar-
ent size in various ways. This was observed more often in dominant
lizards, though in some cases both lizards attempted to increase
their size. Apparent size was increased in several ways, all also
exhibited by A. marcanoi. Most common, other than dewlap exten-
sion, was enlargement of the head. Lizards also increased their
apparent size by lateral compression and throat enlargement.
Dynamic Modifiers
Several display modifiers involved moving body parts and thus
constituted dynamic modifiers (Hover and Jenssen, 1976; Jenssen
1977, 1978). Instead of bobbing with the head, on several occasions
lizards raised and lowered the body by pushing up with the legs,
primarily the forelegs. Push-ups have been widely reported among
/lA7o//5(e.g., Greenberg, 1977; Jenssen, 1979a), and iguanids in gen-
eral (Carpenter, 1967), but no correlation with intensity was disco-
vered in A. marcanoi or A. cyhotes.
Anolis marcanoi
The full elevation display was the most active level of dewlap
display. The rear of the body was moved independently in several
ways. Most common were push-ups with the rear legs, usually in
series of two to six causing the posterior to oscillate greatly, similar
to "rearing" observed in A. opalinus (Jenssen, 1979a). The precur-
sor of these pelvic push-ups could be seen in several intermediate
dewlap displays in which the rear legs were raise and lowered
slightly as the dewlap was thrust out during head-bobbing. In the
full elevation display, the lizards alsosometimes jumped backwards
16 BREVIORA No. 484
several times in rapid succession, forcing the posterior of the body
up-and-down in a manner similar to that caused by hind-leg push-

ups. Similar behavior in the same context has been noted in male A.
lineatopus {Rand, 1967). The full elevation display was usually only
performed in response to a dewlap display or other action by the
other lizard in an encounter. Posterior movements were never per-

formed without such provocation.
Head orientation was also related to intensity level. Throughout
the dewlap displays, and especially at low intensity levels, lizards
changed their head orientation frequently and usually rather slowly.

The snout was often pointed at the other lizard, but this action in
itself did not seem highly significant. On the other hand, a lizard
rapidly turning its snout toward the other, either at the end of a
display by the former or during the display of the latter, appeared to
indicate a high level of intensity. Generally, pointing occurred dur-
ing intermediate or full elevation displays or more intense behavior

and was more often performed by the dominant lizard in the en-
counter. When a dominant lizard rapidly pointed its snout at a
displaying subordinate lizard, the latter often ceased its display,
retracted its dewlap, and dropped to a less elevated posture.
Anolis cy botes
There was no indication in A. cybotes of anything resembling the
pelvic push-ups of A. marcanoi. The posterior was always station-
ary and usually firmly on the substrate, though occasionally the
entire body was lifted off the substrate.
Pointing with the snout at an opponent was commonly seen in A.

cybotes, though neither as frequently nor in as defined a manner as
in A. marcanoi. Several times, a lizard rapidly turned, its snout and
even its whole body directed at its opponent, briefly paused, and
then leapt at or toward it. The derivation of pointing in A. marcanoi
may thus be revealed; perhaps pointing in A. marcanoi has become
ritualized, detached from its original threat of imminent attack.
Pointing was never observed by a lizard which would subsequently

lose an encounter.
A common action more frequently displayed by A. cybotes than
by A. marcanoi was tilting or cocking the head downward toward
an opponent. This often occurred when the lizard was on a branch,
looking down upon an opponent on the ground and often was
1985 /4AY;/.AV SIBLING SPECIES BEHAVIOR 17
associated with some movement or action by the other lizard.
Though cocking of the head may simply allow better vision of the
other lizard's actions, it
probably has a communicative function,
perhaps as an intention movement, indicating awareness of the
other lizard's actions and readiness to respond. Anolis also com-
monly performs this behavior prior to attacking potential prey items
(G. C. Mayer, personal communication). Dominant lizards more
commonly performed this behavior.
Advancing Behavior
Past the stationary display, the steps of increasing intensity of
aggression become more stereotypical, involving movement toward
an opponent and threat of or actual attack, though often inter-
rupted by more bouts of unilateral, alternating, or simultaneous
dewlap displays. In most cases, the progression to increasing levels
of intensity was only stopped by fleeing or quiescence of the subor-
dinate lizard, particularly in A. niarcanoi, though several times both
lizards ceased behaving and assumed less intense postures without
either clearly "winning" the encounter.
Anolis marcanoi
Figure 5 presents the flow of advancing behaviors of A marcanoi. .
Rapid pointing of the snout toward the other lizard was the low-
est level of advancing behavior as well as a display modifier, though
it was usually tied to the interspersed dewlap displays.
still
Taking one or several steps toward the other lizard is the first
overt threat of attack. At close proximity, stepping toward the other
lizard threatened imminent attack and often caused the other lizard
to cease its display and/ or assume a more subordinate position,
retreat one or several steps, or seek escape from the encounter.
Also at closer proximity, intent, or at least threat, to attack was
indicated by the assumption of a "poised" position. The body was
held off the ground with the legs out to the side and appearing ready
to spring. The head, enlarged by crest and throat expansion, with
the dewlap not usually greatly extended, was tilted somewhat for-
ward and toward the other lizard. Rarely was this position assumed
without a subsequent attack. At this time, the lizards were usually in
the faceoff position (Carpenter, 1962, 1978), their bodies parallel
with heads pointed in opposite directions. When displaying on a
18 BR[ VIORA No. 484
1985 /1Af;//.SSlBIIN(i SPECIES BFHAVIOK 19
perch, this poised position often took the form of a crouch, clearly
threatening a jump toward the other lizard. Echelle et al. ( 1971 ), in a
study of aggressive behavior in nine Costa Rican Anolis. noted the
assumption of a similar position in males that subsequently attacked
their opponents.
Usually following the poised position was an advance toward the

rear of the other lizard, culminating in an open-mouthed lunge
toward the other lizard's hindquarters. Biting was never observed at
this stage.Often the tongue was bunched at the front of the open
mouth. Though tongue-bunching is characteristic of aggressive
interactions in many Anolis species (e.g., Greenberg, 1977), it does
not appear to have consistent importance in A. marcanoi. Attacking
individuals always lunged with mouth open and often kept the
mouth open while displaying after lunging. In addition, two lizards
opened their mouths in threat displays without immediately lunging
afterwards.
The attacked lizardresponded either by jumping back and flee-
ing, or pivoting, usually with a jump backwards, away from the
attacking lizard, with its head turned over the shoulder toward the
attacking lizard. The two lizards, now in semi-circular positions
with heads at opposite ends, displayed at each other, bodies com-
ground and snouts angled up, with mouths,
pletely elevated off the
wide open. Occasionally, the attacking lizard con-
at least initially,
tinued moving toward the other's rear so that both circled around,
maintaining the same relative position. In several instances, the
attacked lizard, after displaying, lunged for the other's rear, though
in no instance was it successful in driving the former off. In all but
two cases, after a sequence of displays, lunges, and circling motions,

the attacked lizard fled and attempted to escape the terrarium by
running to a distant wall and clawing wildly at it.
In two instances, the two lizards locked jaws for less than two
seconds, shaking each other violently, until they released, jumping
backwards, with the attacked lizard fleeing. In one of these in-
stances, the attacking lizard chased the other lizard several steps
before ignoring it.
In all instances in which the losing lizard fled, the winning lizard
continued displaying for up to several minutes. Slowly, the lizard's
displays and postures descended the stages of intensity, though they
would temporarily reascend if the other lizard came nearby in its
20 BRFVIORA No. 484
attempts to escape the terrarium. Ultimately, the winning lizard

would assume a non-aggresive posture, stretched out on a perch or
the ground with its head on the substrate or barely elevated and its
rear legs stretched back behind it. The overall effect was to increase
itslength while decreasing its apparent height. This is the normal

resting position of the lizards when undisturbed.
Anolis cyhutes
Advancing behavior was progressively ordered to an extent, but
not completely (Fig. 6). In the most structured sequences, a lizard
performed some display behaviors and then moved towards its
opponent, either jumping off a branch or running half or more of
the distance separating the lizards. This was usually done at a rapid
pace. The advancing lizard then adopted a stalking, "poised" posi-

tion, somewhat like that exhibited by A. mananoi, but with all four
seemingly prepared to spring, with the long axis
legs out to the side,
of the body pointed directly at the other lizard. This was not always
seen, and sometimes a lizard hardly paused between moving toward
and actually attacking the other lizard.
The act of biting an opponent was very different in the two spe-
cies. Encounters involving A. mananoi only resulted in biting when
both lizards continued to behave intensely. The sequence of faceoff

position-lunging-biting was seen only once in the
A. cyhotes
encounters. Instead, most incidents of biting involved very little
ritualized behavior. One lizard leapt or ran at the other and bit it
behind the head, attempting to pin it to the substrate. Attempts to

escape by the attacked lizard often resulted in that lizard carrying
the other around on its back. The attacked lizard often was not
displaying vigorously and in several cases was trying to escape from

the attacking lizard. Anolis mananoi rarely exhibited chasing
behavior, but this was common in A. cyhotes. In the latter species,
on occasion, lizards were chased allover the aquarium until they
could be cornered and bitten.
Most surprising were two instances in which a lizard, with no
preliminaries, ran directly and rapidly at the other lizard,
which also
had exhibited no aggressive behavior, and bit it behind the head, in
both cases, the attacking lizard persistently bit and chased the other
lizard until the encounter was ended. This was reminiscent of the
"vicious biting attack" termed "abnormal" by Greenberg and Noble
1985 •^ vc;//.s siHi i\(; si>fcifs bfhavior 21
o a
o o
in
a>
1/5
n
22 BRl viOKA No. 484
in A.Carolinensis (1944). This behavior was observed by these

investigators in 18 or 800 males, as well as in some females and
surgically-altered males.
Submissive Behavior
A no/is mananoi
As indicated previously, subordinate individuals usually did not
posture as intensely as dominant lizards. Dewlap displays were gen-
erally less elevated, shorter in duration, less frequent, and often
without crests or dark spots. Subordinate lizards also did not as
often attempt to enlarge their apparent head or body size.
More submissive behavior generally took the form of staying very
low to the ground, with the head elevated little and the snout angled
up only slightly, if at all. Carried to the extreme, completely submis-
completely flat on the ground, appearing as incon-
sive lizards laid
spicuous as possible.
At higher levels of intensity, particularly in response to advancing
behavior, submissiveness was usually indicated by fleeing the en-
counter and attempting to escape the aquarium.
Anolis cy botes
One
of the most surprising aspects of A. cybotes" behavior is the
lack of submissive behavior in the lizards that were judged, by the
eventual outcome of an encounter, to be subordinate. Unlike A.
mananoi. these lizards did not behave less vigorously or aggres-
sively, nor did they necessarily display fewer behaviors connoting
higher levels of intensity than the dominant lizard in the encounter.

In several cases, the subordinate lizard initiated aggression by mov-
ing toward the other lizard. This was never seen in the A. mananoi
encounters.
More generally, lizards did not adopt a submissive posture as A.
mananoi did. They sometimes moved away or displayed less, but
they did not flatten themselves or otherwise appear inconspicuous.
Often, a lizard appeared to be trying to escape the aquarium, though
neither persistently nor frantically enough to warrant ending an
encounter. When this proved futile, the lizard turned back and
behaved aggressively again toward the other lizard. Such behavior
was rarely exhibited by A. mananoi.
1985 /I Vr;/ AS SIBIINCi SPECIES BEHAVIOR 23
Index of Aggression
In order to quantify levels of intensity in interactions, an index of

aggression, similar to that developed for Puerto Rican anoles by
Ortiz and Jenssen (1982) was constructed. The index assigns progres-
sively higher point values for actions indicating greater intensity
(Table 2). This index, then, can be used to compare quantitatively
the level of aggressive intensity of A. marcanoi in inter- and intra-
specific encounters (Losos. in press). Points for any behavior are

scored only once per encounter even though many behaviors are
repeated periodically. The score of a lizard in an encounter is the
sum of points awarded for behaviors which it exhibited.
The behavior of A. marcanoi was progressive, with the less
aggressive behaviors preceding the more intense ones. Because this
was .not the case for A. cyboies, no index of aggression could be
developed for that species. Occasionally, the actions of A marcanoi .
during the dewlap display did not necessarily follow a set hierarchy
of increasing intensity either. The most notable exception occurred
when a lizard, usually at a distance from its opponent, sometimes
moved toward the other before pointing its snout rapidly and
directly at it.
Table 2. Index of Aggression used to score encounters between adult male Anolis
marcanoi.
Points Action or Display Modifier
1 low level and intermediate dewlap display

2 full elevation dewlap display
2 crests erected
3 head spot apparent

3 lifting the pelvic region and or moving backwards during
dewlap display
3 pointing snout rapidly at opponent at conclusion of own
display or during opponent's display
4 moving toward opponent (including jumping onto or off of
perch)
6 adopting poised position
7 lunging toward opponent
8 locking jaws
8 opponent tlees
24 BRFVIORA No. 484
Several notes on the construction of this index are necessary. Only

two levels of dewlap display are scored although three were recog-
nized. The intermediate dewlap display stage was considered only
slightly more intense than the low level dewlap display, and conse-
quently the two were grouped together. Though the nuchal crest is
almost always erected before and retracted after the dorsal crest,
they are not considered independent behaviors denoting different
levels of aggressive intensity. The scores of the lizards agree well
with the qualitative evaluations of the 26 encounters. The encoun-
ters can be split into four categories based on their outcome: I.
neither lizard supplants the other; 2. displaying and moving toward,
one flees; 3. lunging, one flees; and 4. jaw-locking, one flees.
The index well represents the level of intensity of aggression in an

encounter (Table 3). Even within particular categories, encounters
that qualitatively appeared to be more intense scored higher on the

index of aggression. The one discrepancy involves several of the
higher scoring encounters in the first category which scored higher
than several of the lower scores in the second category. In the
former encounters, the dominant lizard was fairly aggressive, adopt-
ing the poised position, but then subsiding without forcing the other
lizard to flee. In the latter encounters, on the other hand, the subor-
dinate lizard fled immediately after the dominant one jumped down
from a perch toward These encounters probably would have
it.
scored appreciably higher if the subordinate lizard had not so

promptly fled; the points awarded for forcing the other lizard to flee
were not sufficient to compensate for those lost from the abrupt
termination of the encounter. Notwithstanding this one shortcom-
ing, the index appears to satisfactorily evaluate levels of intensity in
aggressive encounters.
A second confirmation, with four exceptions, of the index's accu-
racy comes from comparison of the scores of the lizards judged
dominant by their behavior, level of activity, and posture with the
scores of those judged subordinate. In no case did the subordinate
lizard achieve a higher score than the dominant lizard, while in four
instances both lizards received the same score.
DISCUSSION
The male aggressive behavior of A . tnarcanoi appears very similar
to that of most other Anolis studied, both overall and in particular
1985 /1,\Y;//.SS1B1ING SPECIES BEHAVIOR 25
Tabic 3. Results of encounters between male Anolis nwnanoi.

26 BRI vioKA No. 484
the ordering of some behaviors and display modifiers depended on

distance separating the lizards. At a given distance, behaviors and
modifiers generally did not appear prematurely, though many ear-
lier behaviors and modifiers recurred at higher levels of intensity. A
comparison of the index of aggression derived for A. marcanoi vj'ith

that derived by Ortiz and Jenssen for Puerto Rican Anolis shows
great similarity, though the latter is more detailed than the index
presented here.
Particular behaviors also resemble those reported for related spe-
cies. Stamps and Barlow (1973) reported several stages in the dew-
lap display of A. aeneus: fan, fanbob, bob, and jerkbob. Not only
wereall four stages observed in A. marcanoi, but they appeared,
very approximately, m
equal proportions to those of A. aeneus.
Crest erection, assumption of the faceoff position, lunging, and jaw-
locking have been observed in a number of species. Maximizing
apparent size by body configuration and orientation is also preval-
ent among Anolis (Jenssen, 1977) and iguanids in general (Carpen-
ter, 1967).
However, there are some differences in the behavior of /I. marca-

noi a?, snout has been reported for only one
well. Pointing with the
other species, A. carolinensis (and then only at very close range
[Cooper. 1977]). Anolis carolinensis is also the only species reported
to exhibit a well-defined postorbital dark spot. Behaviors noted
occasionally in A. marcanoi, such as open mouth threats and mouth-

smacking, are of greater importance in other species such as A.
opalinus (Jenssen, 1979a) and A. aeneus (Stamps and Barlow,
1973). while actions never observed in A. marcanoi, such as tongue
protrusion and foreleg lifting, have been observed in many other
species (e.g..Rand, 1967; Jenssen, 1979a; Jenssen and Rothblum,
1977; Ortiz and Jenssen, 1982). It appears that as well as having its
own stereotyped display-action patterns, each species oi Anolis has
its own behavioral
repertoire, sharing some behaviors with other
species while possessing a few unique behaviors of its own.
With regard to A. cyhotes, two related questions merit address-
ing. Why is the aggressive behavior of A. cybotes so much simpler
than that of /I. marcanoi, and, indeed, of all Anolis examined? And
why there such little order in the appearance of aggressive behav-

is
iors,unlike the highly ordered and progresive pathways of increas-

ing aggression in A. marcanoi and other Anolisl
1985 ,1 \Y;//.S SIMI l\(. Sl'iClFS BFHAVIOK 27
Before these questions can be answered, several points of caution

must be raised. First, it is possible that the experimental arena was
not large enough to elicit the full range and ordering of natural
aggressive behavior. Much larger experimental arenas have been
used by other investigators (e.g., Jenssen, 1970, 1975; Jenssen and
Hover, 1976; Stamps and Barlow, 1973), which was not possible in
is true, however, it is curious that the same problem
this case. If this
did not occur in A. nwrcanoi, which is so similar to A. cyhotes.

Second, the lizards had been in captivity for more than ten months
when the last of these experiments were performed. Though most
appeared to be in reasonably good health, the long stretch of captive

care may have taken its toll on their behavior. The lizards did,
however, react quickly and seemingly normal ways during the
in
encounters; there was just no pattern from lizard-to-lizard or

encounter-to-encounter. Third, the encounters involving A. cyhotes
were staged during the winter, a refractory period for males of the
species in which their aggressive behavior is diminished in the wild
(Jenssen, personal communication). Several A. nwrcanoi tested dur-
ing the winter (which presumably is the refractory period for this
species as well), after the same length of time in captivity, exhibited
typical behavior.
Jenssen (1983) argues that the single stereotyped head-bobbing
display pattern of A. cvhotes is a primitive trait with respect to all
Anolis. The rest of A. cyhotes" aggressive behavior — dewlap dis-
plays, display modifiers, and advancing behavior
— are also simpler
and less ritualized than that of its sibling. A. nwrcanoi. That this
simplicity could not be derived from the more complex behavior of

A. nwrcanoi, however, is not clear.
Even these behaviors are retained primitive ones, the ultimate
if
question, why they have not been expanded and elaborated as in

other species,still needs an answer. Jenssen (1983, personal com-
munication) suggests that the determinants of social spacing in A.

—
cyhotes a large lizard with large territories, few neighbors, low
turnover, and low density — would mitigate against many territorial
challenges. Lizards would know their neighbors and rarely chal-

lenge them. Intrusion would thus be a matter to be seriously dealt
with because interlopers are likely to be lizards without a territory
seeking to displace a resident, and because, due to low turnover,
available territories would be hard to come b\. Consequently, an
28 BRivioRA No. 484
intrusion would rapidly result in a fight and elaborate communica-

tory rituals would be unnecessary. This hypothesis can explain the
simplified aggressive behavior of A. cyhotes compared to that oi A.
distichus (with which it was contrasted by Jenssen), but why such
radical differences with its sibling species, A. marcanoi, and other
truck-ground ecomorphs {sensu Williams, 1972), such as A. crista-

te/lus, A. cooki, A. gundlachi (Ortiz and Jenssen, 1982), and A.
lineatopus (Rand, 1967)? All that can be said is that nothing is

known about the spacing and turnover oi A. marcanoi populations,
so direct comparisons with its sibling species cannot be drawn.
One importantpoint can be made about the lack of distinct sub-

missive behavior in A. cybotes. In A. marcanoi, submissive behavior
appeared to have a pacifying effect on the dominant lizard. Only
rarely and to a limited extent did a dominant lizard continue to act
aggressively toward a clearly submissive opponent. By contrast,
submissive behavior elicited no such complacency in A. cybotes.
Most incidents of biting occurred upon lizards that were not dis-
playing high levels of aggression. Consequently, submissive behav-

ior did not seem to gain any advantage to the lizard displaying it.
Rather, perhaps the best defense if escape is impossible is to put up
an aggressive front, threatening retaliation in case of attack.
Whether these findings and speculations relate to natural behav-
ior needs to be verified in the field. Studies are needed on the social
structure and spacing of populations of both A. marcanoi and A.
cybotes in nature. Futhermore, it is important to find out whether
A. cybotes does, indeed, need a much greater space between indi-
viduals than A. marcanoi for naturally progressive behavior to
appear.
Other than expanding the general pool of knowledge of Anolis
behavior, the data here reported hopefully will be of value in inves-
tigating the ecological and behavioral interactions of these sibling
species. Knowledge of the intraspecific behavior of males of both
species will serve as a control with which to compare the results of
experiments investigating interspecific male aggressive behavior and
the mechanisms of species recognition.
ACKNOWLEDGMENTS
am deeply indebted to Sibel Akyol, Pere Alberch, Ernest E.
1
Williams, and, particularly, Greg Mayer for their continuous

1985 /1\Y;//.SS1BI l\(i SPECIES BEHAVIOR 29
encouragement, advice, and support. I am also deeply grateful to

Sixto and Ivon Inchaustegui for their assistance when 1 was in the
Dominican Republic. Thanks also to Emily Gale, Carlos Garcia,
Harry Greene, Paul Hertz, Thomas Jenssen, James Knight, Jose

Rosado, the members of the Herpetology Department, Museum of
Comparative Zoology, and the staff of the Biological Laboratories,
Harvard University, for their help, and to Crimson Camera for the
loan of a video camera. Laszlo Meszoly and Gene Christman kindly
drew the illustrations.
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1978. Ritualistic social behaviors in lizards, pp. 253-267. In N.

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and P. D. Behavior and Neurology of Lizards: An Inter-
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CooPHR. W. E. 1977. Information analysis of agonistic behavioral sequences in
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leon (Anolis carolinensis). Physiol. Zool. 17: 392-439.
Grhhnbkrg, N. 1977. A neuroethological study of display behavior in the lizard
Anolis carolinensis. Amer. Zool.. 17: 191-201.
Hhrtz. P.E. 1980. Comparative physiological ecology of the sibling species Ano-
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lis J. Herp.. 14: 92-95.
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Herp.. 4: 1-38.
1 975. Display repertoire of a male Phenacosaurus heterodermus (Sauria:
Iguanidae). Herpetologica. 31: 48-55.
1977. Evolution of anoline lizard display behavior. Amer. Zool., 17:
203-215.
1978. Display diversity of anoline lizards and problems of interpretation,
pp. 269 285. In N. Greenberg and P. D. MacLean (eds.). Behavior and Neurol-
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ogy of l-i/ards: An Interdisciplinary Colloquium. Rockville, Md., NIMH, vi +

.^52 pp.
1979a. Display modifiers o\ Anoli.s oijaliniis. Herpetologica. 35: 21 30.

1979b. Display behavior of male Anolis opalinus (Sauria, Iguanidae): A
case ol weak display stereotypy. Anim. Behav.. 27 173 184.
Display behavior of two Haitian li/ards. Anolis cyhotesanA Anolis
19S3.
clisiiclnis. 552 569. /// A. G. J. Rhodin and K. Miyata (eds.). Advances in
Herpclolof;} and Evolutionary Bi(>lof;y: Essays in Honor of Ernest E. lîllianis.

Cambridge. Mus. Comp. Zool.. x + 725 pp.
JiNssi \. T. A.. AND E. I.. Hovi K. 1976. Display analysis of the signature display
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Ji:n.ssi:n. T. A., ami 1,. Rotiiiu i'M. 1977. Display repertoire analysis of Anolis
t ownsendi (Sau&d. Iguanidae). Copeia, 1977: 103 109.
MA^ R. E. 1942. Svstentatics and the Orifîn oj S/)ecies. New York. Columbia
University Press, x + 334 pp.
Okm/. p. R., AND T. A. Jknssi N. 1982. Interspecific aggression bewteen li/ard
competitors. Anolis cook i and Anolis cristatellus. Z. Tierpsychol., 60: 227 238.
I'ac 1 A. E.
. 1974. Systematic and biological studies of the leopard frogs (Rana
pipiens complex) of the United States. Miscellaneous Publications of the
Museum of Zoology, Univ. of Michigan. 148: 140. I
Rand. A. Ecology and social organization in the iguanid li/ard Anolis

S. 1967.
lineaiopus. Proc. U.S. Nat. Mus,. 122: 79. I
S(II()i:ni:r. T. W. 1977. Competition and the niche, pp. 35 136. //; C. Cans and
D. W. Tinkle (eds.) Biology of the Reptilia. Vol. 7. London, Academic Press, viii
+ 720 pp.
S( iiv\ ARIZ, A. 1974. An analysis of variation in the Hispaniolan giant anole
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Stami's. .1. A.. AND G. W. Bari ()w . 1973. Variation and stereotypy in the displays
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r n
B R E V I R A
iiseiim of Comparative Zoology
us ISSN 0006-9698
Cambridge, Mass. 29 AUGUST 1986 Number 485
THE ANATOMY AND RELATIONSHIPS OF

STEREOPHALLODON AND BALDWINONUS
(REPTILIA, PELYCOSAURIA)
Donald Brinkman' and David A. Eberth^
Abstract. As representatives of pelycosaurs from the earliest terrestrial deposits

of the Texas and New Mexico Permo-Pennsylvanian sequence, Stereophallodon and
Baldwinonus are important in our understanding of pelycosaur evolution. However,
their variable placement in the phylogenetic schemes of Romer and Price (1940),
Langston (1965), and Reisz (1980) has underscored the fact that their morphology is
poorly known and that their relationships are openly questioned. Reanalysis of
previously described specimens and heretofore undescribed specimens has allowed
morphology of Stereophallodon and Baldwinonus and
for a reevaluation of both the
the phylogenetic relationships of these taxa with other pelycosaurs.
Stereophallodon is interpreted as the sister-taxon of Ophiacodon on the basis of

five derived features: the shape of the cultriform process, the ventrally directed
basipterygoid process, the anteroposteriorly oriented basipterygoid articular sur-
faces, the presence of a ridge that is triangular in cross section extending dorsally from
the caniniform-tooth buttress, and the presence of a notch on the postero-medial
edge of the quadrate. Stereophallodon is primitive with respect to Ophiacodon in
retaining the following characters: dorsal vertebrae that have flattened ventral sur-
faces and subparallel sides, a femur with an adductor crest running along the mid-
ventral surface of the bone, and a long neck on the astragalus. Apomorphic features
of Stereophallodon are the presence of two greatly enlarged caniniform teeth sup-
ported by a caniniform-tooth buttress and the triangular cross section of the prox-
imal end of the fibula.
'Tyrrell Museum of Palaeontology, Box 7500, Drumheller, Alberta, Canada,

TOJ OYO
^Tyrrell Museum of Palaeontology, Box 7500, Drumheller, Alberta, Canada,
TOJ OYO
'
SEP 1 7
HARV
2 BREVIORA No. 485
Baldwinonus is represented by the weathered remains of a single individual from

New Mexico. It is considered related to Stereophallodon on the basis of shared
derived features of the maxilla.
INTRODUCTION
In a previous paper (Brinkman and Eberth, 1983), the interrela-
tionships of a number of genera of pelycosaurs were considered. The

genera included representatives of most of the families recognized
by Romer and Price (1940), and thus the analysis served as a test of
their long accepted phylogeny. The relationships of the Ophiaco-
dontidae, Edaphosauridae, and Sphenacodontidae that they pro-
posed were supported by this analysis, but the relationships of the
Varanopseidae and the Caseidae were not. Rather than considering
Varanops and Aerosaurus as primitive sphenacodonts and Casea as
a primitive edaphosaurian, it was concluded that these genera are
members of a single clade that is primitive in a number of features
with respect to the clade that includes Dimetrodon, Edaphosaurus,
and Ophiacodon.
Brinkman and Eberth restricted their analysis to better preserved
taxa in order to establish ". .a framework in which detailed taxo-
.
nomic and morphological revisions of individual genera and fami-

lies can be interpreted." (Brinkman and Eberth, 1983: 3). Since that
review, two problematic taxa, Stereophallodon and Baldwinonus,

have been studied by the authors and their relationships with other
pelycosaurs have been reevaluated. In this paper, we describe the
morphology of these two taxa and present our phylogenetic
conclusions.
Stereophallodon was first described by Romer (1937: 90) and was
characterized by the presence of "ophiacodont" vertebrae and two
greatly enlarged caniniform teeth. In a later description (Romer
and
Price, 1940), it was stated that Stereophallodon differed from
Ophiacodon in the presence of two large caniniform teeth, the
greater development of the prefrontal pocket, abrupt decrease
in
length of the postcaniniform teeth, and in showing a tendency

towards a greater ventral flattening of the centra. In the same paper,
Romer and Price (1940) named a new pelycosaur, Baldwinonus
trux, and described it as having ophiacodont-like quadrate,
vertebral and tooth morphology, but sharing with Stereophallodon
the specialized characters of enlarged caniniform teeth, close-set
1985 STEREOPHALLODON MiD BALDWINONUS 3
zygapophyses and flattened ventral centra. Noting similar carnivo-

rous specializations in the diminutive Eothyris, considered by
Romer and Price to be an especially primitive ophiacodont, and
Stereorhachis, a pelycosaur from Autun, France, with Ophiacodon-
like postcrania,Romer and Price erected the taxon Eothyrididae for
the inclusion of Eothyris, Baldwinonus, Stereophallodon, and
Stereorhachis and characterized the family as follows:
Ophiacodont pelycosaurs, primitive in most known

regards but paralleling the higher sphenacodonts in the
development of much enlarged canines and showing a
tendency towards elongation of the vertebral column
(Romer and Price, 1940: 246-247).
The occurrence of Stereophallodon in the earliest terrestrial
deposits of the Texas Permo-Pennsylvanian sequence (Pueblo

Formation) led Romer and Price to suggest that the eothyridids
represented an early radiation of carnivorous pelycosaurs that,
ecologically, were supplanted by sphenacodonts and were, in
themselves, a phyletic dead end. Contrary to this opinion, Watson
(1954, 1957) suspected a close phylogenetic relationship between
Eothyris and the caseids, Casea and Cotylorhynchus. According to
Romer and Price (1940) the caseids were derived from an obscure
group, the Nitosauridae, the taxa of which appear to be based on

fragmentary remains of questionable association (Reisz, 1980).
Watson's insight gained more firm support following Vaughn's 1958
description of Colobomycter pholeter, a small pelycosaur from the
Fort Sill fissure fills of Oklahoma that displays strong affinities to
both Eothyris (short face, anteroposteriorly thickened postorbital
bar, large supratemporal, large parietal foramen, large naris and
similar maxillary dentition) and caseids in general (rugose skull,
short face, large naris, large supratemporal, large parietal foramen
and the presence of an infraorbital foramen). Langston (1965)
described a new pelycosaur from the Cutler Formation of north-
central New Mexico {Oedaleops) that likewise showed a number of
features shared in common by the eothyridids Eothyris and
Colobomycter and the caseids. His reevaluation of the eothyridid
relationships within the Pelycosauria led him to conclude (as had

Vaughn) that the Eothyrididae (including Stereophallodon, Stereo-
4 BREVIORA No. 485
rhachis, Baldwinonus, Eothyris, Oedaleops, Colobomycter, as well

as three genera no longer accepted as members of that
family, the
very poorly known Delorhynchus, Bayloria and Tetraceratops)
were antecedent to the caseids and that, in contradiction to Romer
and Price, the nitosaurids represented a phyletic dead end group of
ophiacodonts convergent with caseids only a few characters of
in
proportion. Most recently, Reisz (1980), in a cladistic analysis of

pelycosaur families, concluded that the relationship between eothy-
ridids {Eothyris and Oedaleops) and caseids was well founded,
citing the shared derived characters of an overhanging rostrum, an

elongate narial opening, and a maxillary contribution to the ventral
border of the orbit.
While the
£'o//2>'r/5, Oedaleops, Co/oZjomj^c/er/ Caseidae relation-
ship appears now to be well established, the position of Stereophal-

lodon, Stereorhachis, and Baldwinonus are less certain. Langston
(1965) chose to retain these taxa within the Eothyrididae although
he cited the enlarged caniniform teeth as the only reason for doing
so and remained uncertain of the validity of the grouping. Reisz
went one step further and removed Stereorhachis and Stereophal-
lodon from the Eothyrididae and placed them in the Ophiacodon-
tidae, (1980: 566, 568). He questioned the reality of Baldwinonus,
identifying the maxilla and neural spines of the type specimen as a
sphenacodont and the vertebrae as pertaining to an ophiacodont. In
our previous (Brinkman and Eberth, 1983) we have
paper,
questioned phylogeny and have proposed a different
Reisz's
arrangement. For this reason and the fact that the relationships of
Baldwinonus, Stereophallodon, and Stereorhachis, and the validity
of Baldwinonus remain uncertain, and because a number of new
specimens of Stereophallodon have become available, we have
undertaken a review of the specimens of these taxa available to us
for examination. Stereorhachis, known from the
Stephanian near
Autun, France, was not available for study and must unfortunately
remain excluded from the following analysis. In considering the
position of Baldwinonus and Stereophallodon the phylogenetic
conclusions of our earlier study were tested and our understanding
of the morphological diversity present in pelycosaurs has been
increased.
1985 STEREOPHALLODON AND BALDWINONUS

The methods used in interpreting polarities of the character-states
are those outlined by Brinkman and Eberth (1983: 3). In that
analysis, the polarity of character-states was interpreted using
outgroup comparison. All other reptiles were considered the closest
sister-group to the pelycosaurs; diadectomorphs were considered
the closest sister-group to the pelycosaurs and all other reptiles; and
anthracosaur amphibians were considered the sister-group of the
clade composed of the foregoing taxa. This framework is used as a
basis for interpreting polarities in this study as well.
The following specimens were examined during the course of this
work:
AMNH 4780: (type and only known specimen of Baldwinonus

trux) fragmentary remains of a single individual
including maxilla, quadrate, twelve partial verte-
brae, and partial ribs;
MCZ 1535: (type of Stereophallodon ciscoensis) fragmentary
remains of a single individual including the pre-
maxilla, maxilla, quadrate, articular, left half of
occiput, and five centra;

MCZ 1944: fragmentary remains of a single large individual
identified as Stereophallodon on the basis of the
maxilla and articular;
MCZ 6618: fragmentary remains of a single large individual,
identified as Stereophallodon on the basis of the
quadrate and articular;

AMNH 4768: associated vertebrae, sacral ribs and premaxilla of
a single large individual, identified as Stereophal-
lodon on the basis of the premaxilla and the
similarity of the vertebrae with those preserved in
MCZ 6618.
A number of isolated elements present in the collections from
Prideaux Pocket, a bonebed in the Pueblo Formation, can be
identified as belonging to Stereophallodon. MCZ
These include:
6358, three premaxillae; MCZ 6354, five maxillary fragments; MCZ
6352, two prefrontals; MCZ 6353, one postfrontal; MCZ 6348, five
6 BREVIORA No. 485
basioccipitals; MCZ 6349, three basi-parasphenoids; MCZ 6350,

four quadrates; MCZ 6359, eight dentary fragments; MCZ 6352,
three articulars; MCZ 6371, one axis centrum; MCZ 6357, partial
centrum of a cervical vertebra; MCZ 6355, twenty mid-dorsal
centra; MCZ 6356, twenty-two centra from various positions along
the vertebral column; MCZ 6768, three ilia; MCZ 7083, part of the
distalend of a femur; MCZ 6766, proximal end of femur; MCZ
6765, two proximal ends of fibulae; and MCZ 6767, four proximal
ends of tibiae.
The following acronyms are used in this paper; AMNH,
American Museum of Natural History; MCZ, Museum of Com-
parative Zoology; UCMP, University of California, Berkeley,
Museum of Paleontology.
DESCRIPTION
Stereophallodon
An almost complete premaxilla is preserved in AMNH

4768 (Fig.
lA-B). This has a posteriorly sloping dorsal ramus and a tooth
bearing base. A foramen is present on the posterior surface of the
premaxilla at the base of the dorsal ramus. Sockets for five teeth are
present in the premaxilla. The diameter of the anterior two sockets
is about twice that of the more posterior sockets. The premaxilla of
Stereophallodon is like that of Ophiacodon (Fig. ID) and different

from that of the sphenacodontines (Fig. IE) in the structure of the
dorsal ramus. In Stereophallodon and Ophiacodon the posterior
edge of the dorsal ramus meets the tooth-bearing base of the bone
just above the maxillary suture. In the sphenacodontines, the
posterior edge of the dorsal ramus meets the base of the premaxilla
anterior to the maxillary suture.
The maxilla (Fig. IC) is represented by a number of specimens
which show a caniniform-tooth buttress and adjacent areas. The
sutural surface for the palatine extends forward onto the buttress to
a point above the exposed posterior edge of the second caniniform
tooth. The maxilla is broken at the dorsal apex of the buttress. The
horizontal cross section at this point shows that a rodlike brace
extended dorsad from the buttress along the inner surface of the
maxilla as in Ophiacodon. In the sphenacodontines no such ridge is
present; the caniniform-tooth buttress is a broad, gently contoured
wedge of bone that merges smoothly and evenly with the medial
1985 STEREOPHALLODON AND BALDWINONUS
Figure 1. The right premaxilla of Stereophallodon in A) lateral and B) medial

views; C) the right maxilla of Stereophallodon in medial view; D) the right
premaxilla of Ophiacodon in lateral view, and E) the right premaxilla of
Sphenacodon in lateral view. Arrows on the premaxillae indicate the postero^-entral
edge of the dorsal ramus of the premaxilla. Stereophallodon premaxilla drawing

based on AMNH 4768, reconstructed portion based on the articular surface for the
premaxilla preserved on the maxilla in MCZ 1944; shaded portion of the maxilla
drawing based on MCZ 1535, with the outline drawn on the basis of MCZ 1944;
Ophiacodon drawing based on UCMP 40035; and Sphenacodon drawing based on
UCMP 83047.
Abbreviations: BUT, caniniform-tooth buttress; BUT, DOR, EX, dorsal extension
of the caniniform-tooth buttress; F, foramen; M S S, maxillary sutural surface; N S S,
nasal sutural surface; PAL S S, palatine sutural surface; VS S, vomer sutural surface.
surface of the maxilla. Moreover, a precaniniform-tooth step is not

present in Stereophallodon. Rather, the ventral edge of the maxilla
slopes gently dorsally and bears well-developed precaniniform teeth.
Two large caniniform teeth are present on the maxilla, each
having a basal diameter of about three times that of any of the more
posterior teeth. At least three teeth are present anterior to the
caniniform teeth. The postcaniniform teeth, as described by Romer
and Price (1940), are extremly short, measuring about one sixth the
length of the caniniform teeth, and are more broadly exposed on the
internal surface of the maxilla than on the external surface.
Fragments of the dermal skull elements present in the type

specimen include the prefrontal, postfrontal, and frontal. Isolate!
8 BREVIORA No. 485
prefrontals are present in the Prideaux Pocket locality. These

elements show that the frontal of Stereophallodon
is like that of
Ophiacodon in having a small lateral lappet. The prefrontal has a

lateral pocket anterior to the orbit, although this does not seem
significantly larger than a similar pocket seen in the prefrontal of
larger specimens of Ophiacodon.
Pterygoids are preserved in MCZ
1944 (Fig. 2 A). The right and
leftpterygoids are preserved together in a single block with the
transverse flange of the right pterygoid visible in dorsal view and the
quadrate ramus of the left pterygoid visible in medial view. The tip
of the transverse flange of the right pterygoid is preserved separately

(Fig. 2C). As in Ophiacodon (Fig. 2B), a tympanic flange extends
TRANS
I cm
Figure 2. The right pterygoid of A) Stereophallodon and B) Ophiacodon in

medial view and C) lateral view of the transverse flange of the pterygoid of Stereo-
phallodon. Drawing of Stereophallodon a composite based on right and left
MCZ 1944. Ophiacodon drawn from MCZ specimen.

pterygoid preserved in
Abbreviations: BSPT, basipterygoid articular surface; TRANS, transverse flange

of the pterygoid; TYMP FLANGE, tympanic flange; ?Q, ?Quadrate.
1985 STEREOPHALLODON AJ<D BALDWINONUS
medially from the ventral edge of the quadrate ramus of the

pterygoid.The transverse flange of the pterygoid is like that of
Ophiacodon in that its lateral edge is thinner relative to its height
than that of Dimetrodon. This difference is consistent in both large
and small specimens of Dimetrodon and Ophiacodon and thus is
independent of size. As in Ophiacodon, a single row of many small
teeth is present on the ventral edge of the transverse flange. Five
teeth are preserved, and more were probably present on the missing
portion of the transverse process (dashed area. Fig. 2A).

Quadrates are present in MCZ
1535 (Fig. 3A-C) and MCZ 6618.
Four isolated quadrates are present in the material from Prideaux
Pocket. The articular surface is differentiated into two condyles
with a saddle-shaped surface extending between them. The lateral
condyle has articular surface extending from its medial to its lateral
surface. articular surface developed only on
The medial condyle has
its and ventral surfaces. A stapedial pit is present on the
lateral
medial edge of the bone just above the medial condyle. This is a
shallow depression containing a round tuberosity. In posterior view
the dorsal edge of the quadrate is notched. This notch is similar in
position to the shelf on the posteromedial surface of the quadrate as

in Ophiacodon and is considered a less well-developed stage of the
same character. The shape of the articular surface in the two genera
is different in that the lateral condyle is broader mediolaterally in
Stereophallodon than in Ophiacodon.
The left half of the occiput
is preserved in 1535 (Fig. 4). MCZ
Although not visible in this view, the fused supraoccipital and
NOTCH
Figure 3. The right quadrate of Stereophallodon in A) posterior, B) medial, and

C) articular views. Drawing based on MCZ 1535.
Abbreviations: LAT COND, lateral condyle; MED COND, medial condyle;

NOTCH, notch in dorsal edge of quadrate; S, stapedial pit.
10 BREVIORA No. 485
TAB
PRO
Figure 4. Left half of the occiput of Stereophallodon, posterior view. Drawing

based on MCZ 1535.
Abbreviations: F M, dorsal edge of foramen magnum; PP, postparietal; PRO,

prootic; TAB, tabular.
prootic end laterally in an oval area of unfinished bone as in

Ophiacodon. Romer and Price (1940) identified a piece of bone on
the lateral corner of the supraoccipital as a displaced postparietal.
However, a separate element is present medial to this in a position
normally occupied by the postparietal in all other pelycosaurs and is
thus more reasonably interpreted as that element. The more lateral
bone is therefore interpreted as a fragment of the tabular.
A nearly complete basi-parasphenoid is present in MCZ 1944
(Fig. 5), and, based on
specimen, a number of fragmentary
this
basi-parasphenoids can be identified among the Prideaux Pocket

material. As in Ophiacodon, the basipterygoid processes are
directed ventrally and the long axis of their articular surfaces are
oriented anteroposteriorly. The cultriform process is like that of
Ophiacodon in being nearly as wide as it is high. This is unlike the
condition in Dimetrodon and Edaphosaurus where the cultriform
process is more bladelike in cross section. The basi-parasphenoid
wings are broadly separated. A
small shelf covers the anterior
portion of the groove between the wings.
Five isolated basioccipitals from Prideaux Pocket can be referred
to Stereophallodon (Fig. 6) on the basis of their large size. No other
pelycosaur from that locality is as large. These are like the

basioccipitals of Ophiacodon in that the occipital condyles are
1985 STEREOPHALLODON AND BALDWINONUS II
CUL PR
Figure 5. Basi-parasphenoid of Stereophallodon in A) dorsal and B) ventral

views. Drawing based on MCZ 1944.
Abbreviations: BS S. basisphenoid shelf; BST TUB, basipterygoid tubercula;

BS W, basi-parasphenoid wings; CUL PR, cultriform process; D S, dorsum sella;
P F, pituitary fossa.
VAG F.
OPS
Figure 6. Basioccipital of Stereophallodon in A) posterior, B) right lateral, C)
dorsal, and D) ventral view. Drawing based on MCZ 6348.
Abbreviations: BS S, basi-parasphenoid sutural surface; OP S, opisthotic sutural

surface; VAG F, medial edge of vagus foramen; XII, foramen for twelfth cranial
nerve.
flattened inferiorly. Also, in both genera, the ventral surface is

marked by a series of ridges and grooves between the basioccipital
tubercula. In Dimetrodon and Edaphosaurus, no ridges are
present on the ventral surface of the basioccipital between the
basioccipital tubercula.
12 BREVIORA No. 485
Articulars are present in MCZ 1535 (Fig. 7), MCZ 1944, and
MCZ 6618. Three articulars are known from the Prideaux Pocket
locality. The most complete of these is MCZ 6618, in which the
angular and prearticular remain in sutural contact with the
articular. The shape of the cotyles correspond to the condyles of the
quadrate: the lateral cotyle is a cup-shaped depression, and the
medial cotyle faces medially. No retroarticular process is
present on
the articular. Instead, a ridge extends from the ventral edge of the
angular to the posterior tip of the articular. A distinct process for
MED PROC
COT
PTM
PROC
Figure 7. Articular of Stereophallodon in A) dorsal, and B) medial views, and the
posterior end of the jaw of C) Ophiacodon and D) Dimetrodon. Stereophallodon

drawing based on MCZ1535, Ophiacodon and Dimetrodon from Romer and Price
(1940).
Abbreviations: A, articular; ANG, angular; C, coronoid; LAT COT, lateral

cotyle; MED COT, medial cotyle; PR A, prearticular; PROC, medial process on
angular; PTM, pterygoid process of articular; RP, retroarticular process; SANG,
surangular.
1985 STEREOPHALLODON AND BALDW/NONUS 13
muscle attachment ispresent on the angular just anterior to the

articular. As in Ophiacodon and Varanops, this is a medially
directed process that is triangular in dorsal view. In Dimetrodon
and Edaphosaurus the process is formed by the articular and it
extends ventromediad. The articular has a ridge on its ventral
surface that marks the limit of contact with the angular. As in
Ophiacodon, Edaphosaurus, and Dimetrodon, the angular extends
below the adductor fossa as a ventrally extended platelike element.
The anterior end of the dentary is preserved in specimen MCZ
1944, and a number of dentary fragments are known from Prideaux
Pocket. In contrast to the slender dentary of Ophiacodon, the
dentary of Stereophallodon is robust and deep. The anterior three
teeth are enlarged, although they are smaller than the caniniform
teeth of the maxilla. Behind these, the dentary narrows medio-
laterally, presumably to allow the upper caniniform teeth to pass

lateral to the dentary.
Twenty associated vertebrae are present in AMNH

4768. These
include two sacral, four caudal, four lumbar, and ten dorsal
vertebrae. Some of these are preserved in articulation. An atlas
centrum and axis are present in MCZ 1944. The axis and four centra
from the cervical or anterior dorsal region are present in MCZ 1535,
and a number of isolated centra are known from Prideaux Pocket.
An atlas intercentrum from this locality is referred to Stereophal-
lodon on the basis of its large size. Thus most morphological areas
along the vertebral column are represented, although the total
number number of vertebrae present
of presacral vertebrae and the
in each areaunknown.
is
The atlas centrum is directly comparable to the fused atlas cen-

trum and axis intercentrum of an adult Ophiacodon in outline,
position of notochordal pits, and width of the ventral surface. Thus,
although no sutures are visible, it is assumed that the axis intercen-
trum is fused to the atlas centrum in this specimen.
The axis is keeled. Anteriorly, a well-developed lip is present to
receive the intercentrum (Fig. 8 A). The transverse processes are
short and slop ventrolaterally.
The four more posterior cervical and anterior dorsal centra
preserved in MCZ
1535 show a transition from a keeled ventral
surface on the cervical centra to a flattened ventral surface
possessing a pair of widely separated ventral ridges on the anterior
14 BREVIORA No. 485
Figure 8. A J) vertebrae of
Stereophallodon. A) axis, B) anterior cervical verte-
bra; C) mid-cervical vertebra; D) anterior dorsal vertebra; E) dorsal vertebra; F)
lumbar vertebra; G) sacral vertebra in lateral and ventral views. H) dorsal vertebra in
proximal view; I) lumbar vertebra; J) and sacral vertebra in distal views. K-N) cross
sections of dorsal vertebrae of K) Stereophallodon; L) Ophiacodon; M) Edaphosau-
rus;and N) Dimetrodon. Drawings A-D based on MCZ 1535; E-I based on AMNH
4768; and K-N from Romer and Price (1940).
Abbreviations: CAP ART SURF, capitular articular surface; IC, intercentrum;

KEEL, mid-ventral keel; LIP, anterior lip on axis; POST ZYG, postzygapophysis;
PRE ZYG, prezygapophysis; RIB, lumbar rib in place; SAC RIB, sacral rib; TRANS
PR, transverse process.
dorsal centra. The most broadly rounded

anterior of the cervicals is
in cross section; its ventral surface possesses a midline keel and two,
more lateral, longitudinal ridges (Fig. 8B). The intercentrum is
preserved in place. Without it, a well-developed lip would be
present. A large articular surface for the head of the rib is present on
each side of the centrum. This extends onto the intercentrum. The
1985 STEREOPHALLODON AND BALDW/NONUS 15
second vertebra in this series (Fig. 8C) is approximately wedge-

shaped in cross section, but has a pair of closely placed ridges
ventrally rather than a single mid-ventral keel. The capitular
articular surfaces are less well developed than in the more anterior
vertebra. A lip to receive the intercentrum is not present. The
following two vertebrae, the best of which is illustrated in Figure 8D,
are more rectangular in cross section. The pair of ridges on the
ventral surface of the centrum are widely separated (Fig. 8D). The
articular surface for the capitular head of the rib is very small in
comparison to the more anterior vertebrae.

Ten dorsal vertebrae are present in AMNH
4768. These are
subrectangular end view and in cross section (Fig. 8H). The
in
ventral surfaces of each centrum is flattened and does not possess a

keel (Fig. 8E). The ventral half of the lateral surfaces of the centra
are subparallel to one another dorsoventrally. small articular A
surface for the capitular head of the rib is located near the dorsal
edge of each centrum. The transverse processes are short and are
separated from the capitular articular surfaces by a small notch. The
zygapophyses are set close to the midline and are strongly sloped.
The neural spines are short, their height being about one and a half
times the height of the centrum. The spines are expanded
anteroposteriorly at their tips.
Four lumbar vertebrae are present in AMNH 4768 (Fig. 8F).
Thesediffer from the dorsal vertebrae in being wider and having a
trefoil-shaped cross section. In end view (Fig. 81) the centrum is
round, rather than sub-rectangular as in the dorsal vertebrae. The
articular surfaces for the ribs on the lumbars are larger than those
on the dorsal vertebrae, and in some specimens ribs remain in place,
fused to the centrum. The zygapophyses are more nearly horizontal
than they are in the dorsal vertebrae. The neural spines are similar
to those of the dorsal vertebrae in their proportions and structure.
The sacral vertebrae can be identified by the exaggeration in the
size of the rib articular surfaces. The ventral half of the centrum is
compressed mediolaterally, forming a rounded ventral ridge that

extends between the ends of the centrum (Fig. 8G). The zyga-
pophyses are similar to those of the dorsal vertebrae in being large,
sub-horizontally oriented surfaces. The neural spine is narrower

anteroposteriorly than in the more anterior vertebrae, and slopes
posteriorly. The centrum is oval in end view (Fig. 8J).
16 BREVIORA No. 485
The caudal vertebrae are round in cross section. The articular

surfaces for the ribs are large, the capitular articular surfaces being
slightly smaller than the tubercular surfaces. Both the proximal and
distal edges of the centrum are deeply bevelled for the intercentra.
The anterior cervical vertebrae of Stereophallodon are like those
of Ophiacodon, Edaphosaurus, and the sphenacodontines: the cen-
and the anterior end is strongly bevelled for the inter-
tra are keeled
centrum. The dorsal vertebrae of Stereophallodon bear little
resemblance to those of Ophiacodon, Edaphosaurus, or the sphena-
codontines. In Ophiacodon, the mid-dorsal vertebrae are wedge-
shaped in cross section (Fig. 8L) and the posterior dorsal vertebrae
are circular in end view and in cross section. Vertebrae that are
subrectangular in cross section like those of the dorsal vertebrae of
Stereophallodon (Fig. 8K) are not present. The dorsal vertebrae of
Edaphosaurus differ from those of Stereophallodon in that in cross
section the centra are rounded rather than flattened ventrally (Fig.
8M). The sphenacodontines have dorsal vertebrae that are strongly
keeled (Fig. 8N). The lumbar vertebrae of Stereophallodon are
unlike those Ophiacodon, Edaphosaurus, or the sphenacodontines
in having a centrum that is trefoil-shaped in cross section.
The proximal end of the ulna is present in MCZ 6618 (Fig. 9). The
element broad, medio-laterally flattened, and its posterior edge is
is
strongly convex. The trochlear notch is shallow and has a distinct

articular surface for the proximal end of the radius. The ulna of
Stereophallodon is like that of Ophiacodon in being broad at the

level of the trochlear notch. In Edaphosaurus and Dimetrodon the
ulna is narrower in this area.
The acetabular region of
the pelvis is present in 6618 (Fig. MCZ
10). Three partial are ilia
present in the collection from the Prideaux
Pocket. The ilium lacks an anterior process and has only a short
posterior process. As in Ophiacodon and Ruthiromia, a shelf
extends medially above the area of articulation with the sacral rib.
This shelf is located at the dorsal edge of the iliac blade.
The distal end of the femur is present in MCZ
6618 (Fig. 1 1). As
in primitive reptiles generally, the dorsal surface of the distal end of
the femur has a well-developed intercondylar groove. The finished
bone covering this groove extends around the distal end of the
femur towards the fibula. The shaft of the femur is circular in cross
section and the adductor crest is a low ridge extending along i*s
1985 STEREOPHALLODON AND BALDWINONUS 17
Figure 9. Proximal end of ulna of A) Stereophallodon and ulna of B) Ophiaco-

don, and C) Dimetrodon. Drawing of Stereophallodon based on MCZ
6618. Draw-
ings of Ophiacodon and Dimetrodon from Romer and Price (1940).
Figure 10. Partial pelvis of Stereophallodon in lateral and medial views. Drawing
based on MCZ 6618.
Abbreviations: IL, ilium; PUB, pubis; SAC, articular surface for sacral rib;
SUPRA AC BUT, supra-acetabular buttress.
mid-ventral surface. This is different from the condition in Ophiaco-

don, where the ridge is a sharp crest located along the fibulad edge
18 BREVIORA No. 485
-ADD CR
1 cm
Figure 1 1 Right femur of A) Stereophallodon, B) Ophiacodon and C) Dime-

.
trodon. Stereophallodon drawn from MCZ

6618. Ophiacodon and Dimetrodon
from Romer and Price (1940).
Abbreviations: ADD CR, adductor crest.
of the bone. In Ruthiromia, Casea, and Edaphosaurus, the adductor

crest is located in a mid-ventral position, but this is a sharp crest
rather than a low rugosity, and gives the femur a tear-drop-shaped

cross section.
The proximal end of the left tibia and the shaft and the distal end
of the right tibia are preserved in MCZ
6618. Three proximal ends
of tibiae are present in the material from Prideaux Pocket. This
element (Fig.12) is much more fully ossified than any tibia known
in Ophiacodon, so detailed comparison is not possible with that
genus. The element differs from the tibiae of Dimetrodon and
Edaphosaurus in that the outline of the lateral condyle of the
articular surface is circular, rather than oval in articular view. The
shaft has well-developed ridges on both the medial and lateral
surfaces. Well-developed ridges are present on the lateral edges of
all other pelycosaur tibiae examined. A
slight ridge is present on the
internal edge of the tibia in Ophiacodon, although it is not as
strongly developed. Ruthiromia is similar to Stereophallodon in
having a well-developed ridge on the internal surface of the tibia.
The proximal and distal ends of a fibula are present in MCZ 6618.
Two proximal ends of fibulae are present in the material from
MED CON
CNEM
1 cm
Figure 12. Left tibia of Stereophallodon in articular and anterior views. Anterior
view is a composite drawing based on right and left elements present in MCZ6618.
Abbreviations: CNEM, cnemial tuberosity; LAT CON, lateral condyle, MED

CON, medial condyle.
Prideaux Pocket (Fig. 13). The proximal end of the fibula is
triangular in proximal view, with the base of the triangle forming

the articular surface for the femur. A muscle scar is present on the
lateral tip of the triangle. The proximal end of the fibula is unlike
20 BREVIORA No. 485
TUB
FEM
Figure 13. A-C) the proximal end of the right fibula of Stereophallodon, in A)
lateral view, B) medial view, and C) proximal view; D) left fibula of Stereophallodon,
actual length unknown; E) Ophiacodon, and F) Dimetrodon. Stereophallodon draw-
ing based on MCZ 6618, Ophiacodon and Dimetrodon from Romer and Price
(1940).
Abbreviations: FEM, articular surface for femur; TAR, articular surface for
tarsus; TUB, tuber of uncertain function.
those of other pelycosaurs in its triangular shape. In Ophiacodon,
Dimetrodon, and Edaphosaurus the proximal end of the fibula is

rectangular in end view and has a proximo-distally oriented
tuberosity on its lateral edge. The distal end of the fibula of Stereo-
phallodon is wide and dorso-ventrally compressed. The astragalar
and calcaneal articular surfaces are set at a low angle to each other
and are differentiated by a slight narrowing of the distal end of the
bone. Ophiacodon, Edaphosaurus, and Ruthiromia are like Stereo-
phallodon in having broad distal ends of the fibulae. In sphenaco-
dontines, the distal end of the fibula is narrower mediolaterally.
A nearly complete left astragalus and the distal half of the right
astragalus are present in MCZ
6618 (Fig. 14). As in primitive
reptiles generally, this element is L-shaped with a tibial articular
surface located on the lateral branch of the L, and the dorsal branch
contributes to the support of the fibula. A perforating notch is
present on the lateral face of the astragalus. The proportions of the
astragalus are like those of the element in Edaphosaurus in that the
neck is elongate, constituting nearly half of the proximodistal length
of the bone. This is unlike the condition in Ophiacodon, where the
neck is less than 20% of the length of the element.
TIB
PERF
FOR
Figure 14. The astragalus of Stereophallodon in A) dorsal and B) ventral views,

and the astragalus of C) Ophiacodon and D) Dimetrodon. Stereophallodon drawing
based on MCZ 6618. The shaded area is the preserved portion of the right astragalus;
the dotted outline is based on the preserved portion of the left astragalus. Ophiaco-
don and Dimetrodon from Romer and Price (1940).
Abbreviations: PER FOR, medial edge of perforating foramen, TIB, articular

surface for tibia.
22 BREVIORA No. 485
Baldwinonus
A fragmentary right maxilla is preserved and displays 25 teeth or
alveoli including two that are greatly enlarged (Fig. 15). In lateral
view the ventral edge of the maxilla is nearly straight posterior to
the caniniform teeth and shows a steep incline anterior to them. No
edentulous stepis present as in sphenacodontines. The bases of five
precaniniform teeth are present. These increase in size posteriorly.

All were apparently erupted at the time of death. The base of one
caniniform tooth is present and is suboval in cross section with a
shallow dorsoventral groove along its anterior and posterior
surfaces.The base of this tooth is nine times greater in cross-
sectional area than those of the adjacent teeth as originally observed
by Romer and Price, 1940. Eighteen postcaniniform teeth or alveoli

are present.Those teeth that are fully preserved show compression
and recurvature only at the tips. Slight uncrenulated cutting edges
are present along these tips.
In medial view, the alveolar ridge is swollen above the caniniform
teeth forming a caniniform-tooth buttress. This swelling accommo-
dates the sockets for the caniniform teeth. A horizontal cross section
of the dorsal-most extent of the buttress triangular in outline and
is
thus indicates that the buttress extended dorsad against the medial
surface of the maxillary wall as a vertical rod or ridge. Such a
BUT
PM s s
Figure 15. A) the right maxilla of Baldwinonus in internal view. Arrows mark the
position of B) cross-section through the maxilla. Drawing of AMNH 4780.

Abbreviations: BUT, caniniform buttress; MX PAL V S, maxillary/ palatine ves-
sel scar; PAL S S, palatine sutural surface; PM S S, premaxilla sutural surface.
1985 STEREOPHALLODON A^D BALDWINONUS 23
Structureis seen in Ophiacodon but not in the sphenacodontines
where the caniniform-tooth buttress extends dorsally as a broad

smoothly contoured wedge. The presence of a rodlike extension is
also indicated by the down turning of the maxillary/ palatine vessel
scar as passes forward along the medial surface of the buttress.
it An
identical condition exists in Ophiacodon where the maxillary
palatine vessel is downturned presumably to avoid contact

with the
edges of the rodlike extension. In sphenacodontines, where the

buttress swelling smoothly extends into adjacent surfaces over a
broad area at its dorsal terminus, the vessel scar extends forward
with no downturning.
Posterior to the buttress the aveolar ridge contracts, forming, in
cross section, a V-shaped structure (Fig. 15B). In sphenacodontines,
the alveolar ridge remains rectangular in cross section.
The medial half of a left quadrate is preserved (Fig. 16). The
articular surface is a saddle-shaped joint. The articular surface of
the medial condyle developed only on its lateral-facing surface.
is
The lateral not fully preserved. The stapedial pit contains

condyle is
a strong tuberosity, presumably marking the site of contact with the
stapes. A notch is present on the dorsal edge of the bone.

In all these
features, the quadrate of Baldwinonus is like that of Stereophallo-
don.
Thirteen partial vertebrae are present. Each can be placed in one
of three groups defined on the basis of the morphology of the
centrum and transverse process, and the relative position of these
NOTCH
Figure 16. The left quadrate of Baldwinonus in posterior, medial, and articular
views. Drawing based on AMNH 4780.
Abbreviations: LAT CON, lateral condyle; MED CON, medial condyle;

NOTCH, notch in dorsal edge of quadrate; S, stapedial pit.
24 BREVIORA No. 485
groups can be established on the basis of comparison with

Stereophallodon.
The first group comprises four vertebrae; each possesses a sharp,
ventral keel, a wedge-shaped cross section, and a bevelled antero-
ventral edge. This group is represented by the specimen illustrated in
Figure 17 A. When preserved, the transverse process slopes postero-
laterally and is about equal in length to thewidth of the centrum. It

has a large oval articular surface for the rib. The neural spine,
preserved on an unillustrated specimen measures about twice

the
height of the centrum. The slope of the transverse process indicates

that these vertebrae are located in the cervical region of the vertebral
column.
The second group comprises three vertebrae, two of which are
illustrated. Figure 17B and C. These vertebrae are longer than those
of the first group and possess a centrum whose ventral surface
displays two, close-set parallel ridges. A
transverse process, pre-
served on one of these, is a short laterally directed structure with a
ventral buttress extending to the anterior end of the centrum. These
vertebrae correspond to the anterior dorsal vertebrae of Stereo-
phallodon.
1cm
Figure 17. Presacral vertebrae of Baldwinonus. A) anterior cervical vertebra, B)

mid-cervical vertebra, C) posterior-cervical vertebra, and D) dorsal vertebra. Draw-
ings based on AMNH4780.
Abbreviations: CAP ART SURF, capitular articular surface.

1985 STEREOPHALLODON M^D BALDIV/NONUS 25
Six vertebrae constitute the third group and are best exemplified
by the illustrated specimen, Figure 17D. These vertebrae possess
smooth, flat ventral surfaces. The lateral surfaces of the ventral half
of the centra are approximately parallel dorsoventrally, and thus the
centra are subrectangular in cross section. These vertebrae cor-
respond to the dorsal vertebrae of Stereophallodon.
DISCUSSION
Ophiacodon and Stereophallodon
In the review of pelycosaurs presented by Brinkman and Eberth,
(1983) two major clades were recognized. The first includes Casea,
Ruthiromia, Varanops, and Aerosaurus. This clade is referred to
below as the Casea- Varanops clade. It was defined on the basis of
three derived character-states: 1) the presence of a secondary ridge
leading from the distal end of the deltopectoral crest to a more
medial position on the proximal end of the humerus; 2) the presence
of a contact between the maxilla and quadratojugal with these
bones raised to form a ridge along the contact and; 3) the presence
of basipterygoid processes directed laterally and articular surfaces
elongate mediolaterally. The second includes Ophiacodon, Edapho-
saurus, Dimetrodon, and Sphenacodon. This clade is referred to
below as the Ophiacodon- Dimetrodon clade. It is defined on the
basis of four shared derived characters: 1) a concave ventral cheek
margin; 2) a posteriorly sloping anterior margin of the premaxilla;

3) a dorsal process of the stapes that articulates with the ventral
surface of the paroccipital process of the opisthotic and; 4) an
extended ventral plate of the angular in the region of the articular.
The Stereophallodon material at hand includes only one of the
skeletal regions used in defining the Casea- Varanops clade (the
basipterygoid processes of the basi-parasphenoid). In Stereophallo-
don the processes extend ventrad and possess smooth antero-
posteriorly elongate articulating surfaces. Such a morphology is
absent in members of the Casea- Varanops clade and had been
considered a unique derived character of Ophiacodon (Brinkman
and Eberth, 1983).
The present Stereophallodon material includes two of the four
Ophiacodon- Dimetrodon clade
skeletal regions used in defining the
(the premaxilla and the angular). The premaxilla is like those of

members of the Ophiacodon- Dimetrodon clade in that it does not
26 BREVIORA No. 485
project anteriad as is the case primitively in pelycosaurs. The

angular shares with those of members of the Ophiacodon-Dimetro-
don clade the presence of an extended ventral plate in the region of
the articular. Thus these character-states indicate that Stereophallo-
don is a member of the Ophiacodon- Dimetrodon clade.
Within the Ophiacodon- Dimetrodon clade, Stereophallodon is
most parsimoniously placed as the closest sister-taxon of Ophiaco-
don (Fig. 18) and forms with Ophiacodon a monophyletic taxon
defined by five characters. Two of these, the ventral orientation of
the basipterygoid processes and the antero-posteriorly oriented
articular surfaces have been discussed above and are present
elsewhere in pelycosaurs only in Ophiacodon.
Three additional features not discussed by Brinkman and Eberth
(1983) can be interpreted as derived and shared by only Ophiacodon
and Stereophallodon. In interpreting the polarity of these charac-
ters, themethodology used in Brinkman and Eberth (1983) was
employed. The outgroups used are, 1) members of the Casea-
Varanops clade, which is the closest sister-taxon of the Ophiacodon-
Dimetrodon clade and, 2) the taxa that were used as outgroups in
interpreting the polarities of character-states within pelycosaurs.
The polarity of character-states was interpreted according to the
distributions shown in Brinkman and Eberth (1983: Fig. 2).
Casea Varanops Aerosaurus Ophiacodon Edaphosaurus Dimetrodon Sphenocodon
Figure 18. Cladogram showing the interrelationships of seven genera of pelyco-

saurs. From Brinkman and Eberth (1983).
1985 STEREOPHALLODON AJ<D BALDWINONUS 27
One of these features is the presence of a ridge that is triangular in

cross section extending dorsally from the caniniform-tooth buttress.
This is not present in Edaphosaurus or the sphenacodontines. The
condition is not known in the members of the Casea-Varanops
clade, but in other reptiles used as outgroups, including Paleothyris,

Captorhinus, and Diadectes, no ridge is present. Thus the absence
of such a ridge is interpreted as primitive for pelycosaurs, and its
presence is interpreted as a derived feature. Since it is only known in
Stereophallodon and Ophiacodon, it
supports the hypothesis that
these genera are members of the same clade.
The second feature is the presence of a notch on the postero-
medial edge of the quadrate. Although differing in detail, the
similarity in position of this and the shelf on the medial surface of
the quadrate of Ophiacodon indicates that they are different
variants on a single character-state. No other pelycosaur shows a
notch or shelf in this position. None is present in captorhinids or
diadectomorphs. The absence of such a notch, therefore, is
interpreted as primitive for pelycosaurs, and the condition in Stereo-
phallodon and Ophiacodon is interpreted as a derived character-
state shared by those two genera.
The third character-state is the presence of a cultriform process
that is wide in cross section. In Dimetrodon, Edaphosaurus,
Varanops, Casea, and Diadectes, the cultriform process is bladelike

(thatis, much higher than it is wide) in cross section. In Stereophal-
lodon and Ophiacodon, the cultriform process is wider than it is

high and has a broad U-shaped cross section. The presence of the
bladelike cultriform process in both the members of the Casea-
Varanops clade and in Diadectes indicates that it is primitive for
pelycosaurs. The wide cultriform process is therefore interpreted as
a derived character state that is shared by Stereophallodon and
Ophiacodon.
An alternative hypothesis of relationship that would place Stereo-
phallodon as a member of the Dimetrodon-Sphenacodon clade is

weakly supported by two characters: the presence of an enlarged
caniniform-teeth buttress and two greatly enlarged caniniform
teeth. Although the Dimetrodon-Sphenacodon clade was not
discussed by Brinkman and Eberth (1983), a number of shared
derived characters have been cited by Reisz (1980) and Reisz (in
press) as defining the clade. These are the presence of 1) an
28 BREVIORA No. 485
edentulous step anterior to the caniniform teeth; 2) enlarged

caniniform teeth; 3) a swollen caniniform tooth buttress; 4) a
reflected lamina of the angular; 5) a tall septomaxilla; 6) keeled
dorsal vertebrae; 7) a ventrolaterally oriented paroccipital process;
8) a short lacrimal; 9) a tall skull and; 10) a strongly convex
maxillary margin. The absence in Stereophallodon of an edentulous
step anterior to the caniniform teeth, a reflected lamina, keeled
dorsal vertebrae, and ventrolaterally oriented paroccipital processes
indicates that Stereophallodon cannot be a member of this clade.
Furthermore the absence in Stereophallodon of characters that
define the Edaphosaurus-Dimetrodon-Sphenacodon clade (basi-
pterygoid articular surfaces at right angles; absence of a shelf
between the basipterygoid tubera, a well-developed lateral lappet of
the frontal, a downturned prearticular, a pterygoideus process
formed by the articular, a quadrate ramus of the pterygoid without
a tympanic flange, a well-developed lateral process of the supra-
occipital, lateral excavations on the neural arches, transverse

processes of vertebrae lacking a ventral web of bone) indicates that
Stereophallodon is not the sister taxon of the Dimetrodon-
Sphenacodon Thus Stereophallodon is interpreted as the
clade.
Ophiacodon that has paralleled members of

closest sister-taxon of
the Dimetrodon-Sphenacodon clade in the development of enlarged
caniniform teeth supported by a large caniniform-tooth buttress.
This hypothesis of relationship is in agreement with the recom-
mendation of Reisz (1980) that Stereophallodon be separated from
the Eothyrididae and placed in the Ophiacodontidae.
In some characters, Stereophallodon and Ophiacodon are differ-
ent. Outgroup comparison was used to determine whether the
character-states that occur in Stereophallodon are primitive for the
Ophiacodon- Stereophallodon clade or apomorphic for Stereophal-
lodon.The outgroups used in this analysis are the Edaphosaurus-
Dimetrodon clade (the sister-taxon of the Ophiacodon- Stereophal-
lodon clade), the Casea-Varanops clade (the sister-taxon of the
Ophiacodon- Dimetodon clade), other reptiles (the sister-taxon to
the pelycosaurs) and diadectomorphs (the sister-taxon to all of
these).The polarity of character-states was interpreted according to
the distributions shown in Brinkman and Eberth (1983: Fig. 2).
One of the differences between Stereophallodon and Ophiacodon
is in the structure of the centra. In Stereophallodon the dorsal
vertebrae are rectangular in end view, and have flat ventral surfaces
and subparallel sides. In Ophiacodon vertebrae of the cervical and
dorsal region are wedge-shaped in cross section and vertebrae of the
lumbar region are circular in end view and in cross section. The
dorsal vertebrae of Casea are like those of Stereophallodon. Those
of Ruthiromia differ in detail, but are similar to Stereophallodon in
that the posterior dorsal vertebrae tend to have a flat ventral
surface. Also, the trefoil shaped cross section seen in the lumbar
vertebrae of Stereophallodon can be considered a structural ante-
cedent to the "pinched-in" condition of the lumbar vertebrae of
Ruthiromia (see Eberth and Brinkman, 1983). Since Casea and
Ruthiromia are primitive members of the Casea- Varanops clade,
and Stereophallodon is primitive within the Ophiacodon- Dimetro-
don clade, the morphology of the centra of the dorsal vertebrae seen
inStereophallodon is interpreted as primitive with respect to Ophia-
codon. Inherent in this conclusion is the interpretation that this
centrum morphology is primitive for pelycosaurs.
A second difference between Stereophallodon and Ophiacodon is
seen in the structure of the femur. The femur of Ophiacodon
possesses a sharp adductor crest that runs along its postero-ventral
edge. Stereophallodon differs from Ophiacodon and is like other
pelycosaurs in that the adductor crest is located in a mid-ventral
position. The wide distribution of this latter character-state indicates
that it is a primitive feature in pelycosaurs. Thus in this feature,
Stereophallodon is more
primitive than Ophiacodon.
A third difference between Stereophallodon and Ophiacodon is
seen in the structure of the astragalus. In Stereophallodon the
astragalus has a long neck, in Ophiacodon it is short. The astragalus

of Ophiacodon has been thought to be primitive, but establishing
polarities has been (Eberth and Brinkman, 1983). A long
difficult
neck is
present Ruthiromia
in (the primitive member of the
Varanops clade), Edaphosaurus (the primitive member of the
Edaphosaurus-Sphenacodon-Dimetrodon clade), and Stereophallo-
don. Thus the Stereophallodon character-state is interpreted as the
primitive condition for pelycosaurs.
A
fourth difference between Stereophallodon and Ophiacodon is
seen in the structure of the quadrate. The quadrate of Stereophallo-

don is distinctive in the shape of the articular surface. Edaphosaurus,
Ophiacodon, the sphenacodontids, Casea, and the varanopseids
30 BREVIORA No. 485
each have a different quadrate articular morphology. Thus it is not

possible to determine which is the primitive pattern using outgroup
comparison.
A fifth difference between Stereophallodon and Ophiacodon is
seen in the structure of the fibula. The fibula of Stereophallodon is
distinctive in the triangular cross-sectional shape of its proximal

end. In Ophiacodon, the fibula has a rectangular proximal end. This
is also the case in Edaphosaurus, sphenacodontines, and the
varanopseids. Thus the character-state seen in Ophiacodon is
interpreted as the primitive character-state and that seen in Stereo-
phallodon is considered apomorphic for that genus.
from Ophiacodon in the presence

Finally, Stereophallodon differs
of a caniniform-tooth buttress and the large size of the caniniform
teeth. As discussed above, these are interpreted as derived features
acquired in parallel with the derived members of the Dimetrodon-

Sphenacodon clade.
In summary, Stereophallodon is interpreted as the closest sister-
taxon of Ophiacodon. position in the Ophiacodon- Edaphosaurus-

Its
Dimetrodon clade is supported by seven shared derived character-

states. Two of these are shared with all other members of the clade;
five are derived features shared only with Ophiacodon. The two
features shared with all other members of the clade are the presence
of a platelike extension of the angular beneath the articular and the
posterior slope of the dorsal ramus of the premaxilla. The features
that unite Stereophallodon with Ophiacodon are the shape of the
cultriform process of the basi-parasphenoid, the ventrally directed
basipterygoid process, the anteroposteriorly oriented basipterygoid
articular surfaces, the presence of a ridge that is triangular in cross
section extending dorsally from the caniniform-tooth buttress, and
the presence of a notch on the postero-medial edge of the quadrate.
Stereophallodon shares five features with Ophiacodon that are
primitive relative to Edaphosaurus and the sphenacodontines.
These are the presence of a triangular process located medial and
anterior to the articular, the presence of a tympanic flange on the
ventral edge of the quadrate ramus of the pterygoid, the absence of
an anterior process on the ilium, the presence of a shelf covering the
anterior end of the groove between the basisphenoid wings, and the
absence of a retroarticular process. Stereophallodon is primitive
with respect to Ophiacodon in retaining the following characters:
dorsal vertebrae that have flattened ventral surfaces and subparallel

sides, a femur with an adductor crest running along the mid-ventral
surface of the bone, and a long neck on the astragalus. Stereophallo-
don is derived relative to Ophiacodon in the
presence of two greatly
enlarged caniniform teeth supported by a caniniform-tooth buttress,
and a fibula that has a triangular cross section at its proximal end.
Baldwinonus
Before considering the systematic position of Baldwinonus it is
necessary to consider whether or not more than one individual is
present in the type specimen. This was questioned by Reisz (1980),

who identified the centra as ophiacodont elements, and the maxilla
and neural spines as sphenacodont. Thus Reisz concluded that at
two individuals were present.
least
In identifying the maxilla as that of a sphenacodont, rather than a
—
sphenacodontine the only group of sphenacodonts known from
—
New Mexico Reisz recognized that it differs significantly from the
maxilla of other betterknown sphenacodontines. Also, in identify-
ing the vertebrae as ophiacodont vertebrae Reisz recognized that
they were not from Ophiacodon, the only ophiacodontid present in
New Mexico. In the features in which the vertebrae and maxilla of
Baldwinonus are different from the sphenacodontines and Ophiaco-
don, they are like Stereophallodon. The quadrate of Baldwinonus,
not considered by Reisz, is also like Stereophallodon and unlike the
quadrate of Ophiacodon or the sphenacodontines. While studying
the type specimen, it was possible to piece together many of the
vertebral fragments. Some of the resulting vertebrae are in articula-
tion, and in one case a neural spine, identified as a sphenacodont
neural spine by Reisz, could be attached to a centrum. Thus, it is
concluded that the type specimen of Baldwinonus is most probably
a single individual that is morphologically similar to Stereophal-
lodon.
In most features in which both Baldwinonus and Stereophallodon
are known, Baldwinonus is simply a smaller version of Stereophallo-
don. A difference in the structure of the maxilla is present: in Stereo-
phallodon, the palatine sutural surface extends onto the caniniform-

tooth buttress; in Baldwinonus, the palatine sutural surface ends
just posterior to the caniniform-tooth buttress. On the basis of this
difference, Baldwinonus is considered a valid genus.
32 BREVIORA No. 485
Two derived character-states that unite Stereophallodon and

Ophiacodon are also present in Baldwinonus; therefore, the three
genera can be combined in a monophyletic group. These are the

presence of the rodUke dorsal extension of the caniniform-tooth
buttress on the internal surface of the maxilla and the presence of a
notch in the postero-medial surface of the quadrate.
That Baldwinonus and Stereophallodon may be more closely
related to each other than either
is to Ophiacodon is indicated by the
shared derived character-state of greatly enlarged caniniform teeth

supported by a buttress on the internal surface of the maxilla. Other
character-states of Stereophallodon that were interpreted as being
derived, specifically the structure of the proximal end of the fibula,
are not known in Baldwinonus.
Character-states shared by Baldwinonus and Stereophallodon
that can be considered primitive for pelycosaurs in general are seen
in the structure of the vertebrae. The vertebrae of Baldwinonus are
like those of Stereophallodon in that the dorsal vertebrae have flat
ventral surfaces and subparallel sides. Thus the discussion of the

polarity of the vertebral features of Stereophallodon given above
also applies to Baldwinonus.
The structure of the maxilla of Baldwinonus is more fully known
than that of Stereophallodon. Like the maxilla fragments of Stereo-
phallodon, only superficially similar to the sphenacodontines.
it is
The include the absence of the following derived

differences
sphenacodontine characters: a maxillary step, a reduced tooth

count, compressed teeth with fully developed cutting edges, an
evenly tapering caniniform-tooth buttress, and a rectangular post-
buttress alveolar ridge.
In summary, from the characters discussed here, it is concluded
that Baldwinonus closely related to Stereophallodon. These genera
is
are interpreted as primitive members of a clade that also includes
Ophiacodon, and therefore replace Ophiacodon as the most

primitive known member of the Ophiacodon- Dimetrodon clade
(Fig. 19). Character-states shared
by Stereophallodon, Baldwinonus,
and primitive members of the Casea-Varanops clade can be
considered primitive for pelycosaurs. One such feature is the
structure of the centra of the dorsal vertebrae. This is of significance
in providing evidence that pelycosaurs, as traditionally defined, had
a monophyletic origin.
Casea Ruthiromia Aerosaurus Voronops Optiiocodon Stereophollodon Baldwinonus Edophosourus Sptienocodon Dimetrodon
Figure 19. Cladogram showing the interrelationships of Stereophallodon, Bald-

winonus and the pelycosaurian genera considered by Brinkman and Eberth (1983)
and Eberth and Brinkman (1983).
ACKNOWLEDGMENTS
The authors are greatly indebted to the scientific and curatorial
staff of theAmerican Museum of Natural History for permission
to study specimens in their care. The manuscript was read by Phil
Currie. His critical comments were greatly appreciated. The
specimen drawings, except for Figures IC-E, 5, and 7A-B, were

made by Linda Krause of the Tyrrell Museum of Palaeontology.
Figures IC and 7A-B were made by Donna Sloan of the Tyrrell
Museum. The manuscript was typed by Judy McMillan. The
comments of two anonymous reviewers also greatly improved the
manuscript. The responsibility for any errors in methodology or
conclusion rest solely with the authors.
LITERATURE CITED
Brinkman, D. B., and D. A. Eberth. 1983. The interrelationships of pelycosaurs.
Breviora No. 473, 35 pp.
Eberth, D. A., and D. B. Brinkman. 1983. Ruthiromia elcobriensis, a new
pelycosaur from El Cobre Canyon, New Mexico. Breviora, No. 474, 26. pp.
Langston, W., Jr. 1965. Oedaleops campi (Reptilia: Pelycosauria). A new genus
and species from the Lower Permian of New Mexico and the family
Eothyrididae. Bull. Texas Mem. Mus., 9: 1-46.
34 BREVIORA No. 485
Reisz, R. R. 1980. The Pelycosauria: a review of phylogenetic relationships, pp.

553-592. In A. L. Panchen (ed.). The Terrestrial Environment and the Origin of
Land Vertebrates, London and New York, Academic Press.
.
Pelycosauria. Handbuch der Palaeoherpetologie. Fischer, Stuttgart.
RoMER, A. S. 1937. New genera and species of pelycosaurian reptiles. Proc. New
Engl. Zool. Club, 16: 89-96.
RoMER, A. S., AND L. L Price. 1940. Review of the Pelycosauria. Geol. Soc. Am.
Spec. Pap. No. 28, 538 pp.
Vaughn, P. P. 1958. On a new pelycosaur from the Lower Permian of
Oklahoma, and the origin of the family Caseidae. J. Paleont., 32: 981-991.
Watson, D. M. S. 1954. On Bolosaurus and the origin and classification of
reptiles. Bull. Mus. Comp. Zool., Ill: 299-249.

. 1957. On Millerosaurus and the early history of the sauropsid reptiles.
Phil. Trans. Roy. Soc. Lond., B, 240: 325-400.
B R E V I R A
us ISSN 0006 9698
Cambridge, Mass. 29 August 1986 Number 486
THELODUS MACINTOSH! STETSON 1928,

THE LARGEST KNOWN THELODONT
(AGNATHA: THELODONTI)
Susan Turner'
Abstract. Material attributed to the thelodont Thelodus macintoshi Stetson,

from the Silurian of New Brunswick, Canada, is shown to be heterogeneous. It
comprises not only the type thelodont, T. parvidens Agassiz 1839, but also a species
of Loganella similar to L. ludlowiensis or L. martinssoni (Gross 1967) and an
acanthodian resembling Gomphonchus. Scales of T. bicostatus Hoppe and T. trilo-
batus Hoppe are found in association with those of T. parvidens for the first time.
Measurements of Stetson's type material confirm that T. parvidens is the largest
known thelodont, with a total length of about one meter. The vertebrate fauna
including these thelodonts could be as old as late Llandovery (within the Long Reach
Formation) or as young as early Pridoii (in the Jones Creek Formation). The New
Brunswick thelodonts are similar in preservation to those from the Lower Silurian
fish beds of Scotland, but their phylogenetic affinities are closer to those from the
Silurian of England and the Baltic region.
INTRODUCTION
In 1928 Harold Stetson described articulated thelodont material,
preserved in calcareous concretions, from a site in the Silurian rocks

near Nerepis, Kings Co., New Brunswick, eastern Canada. This was
the first definite thelodont material found in North America. The
Cunningham's Brook (Cunningham's Creek on the map of
site, at
MacKenzie 1964), has also yielded the heterostracan Cyathaspis

acadica (Matthew), (see Denison 1964), spines of an acanthodian
related to C//m6r//u5 (Denison 1956, p. 385), and a possible anaspid.
" '"
'Queensland Museum, Fortitude Valley, Queensland 4006, Australia
:0
2 BREVIORA No. 486
Ctenopleuron nerepisense Matthew 1907, which remains indetermi-

nate (e.g.. Woodward 1920). Associated invertebrates include cera-
tiocarids, conodonts and a xiphosuran (Denison 1956; Blieck 1982).

The nodules containing this thelodont material were collected
from siltstones by Stetson and W. E. Schevill of the Museum of
Comparative Zoology during a Harvard expedition in 1927, follow-
ing up the lead of William Macintosh of the Natural History
Society of New Brunswick. The type material is housed at the
Museum of Comparative Zoology, Harvard University.
Subsequently, during the 1950s and 1960s, Robert Denison (then
of the Field Museum of Natural History, Chicago) collected more
material from two separate horizons. Denison referred all the thelo-
donts he collected to Thelodus macintoshi Stetson. In 1956 he stated
that the specimens came from the Long Reach Formation, follow-
ing the lead of Matthew (1888) who stated that the fish beds were
in the Mascarene Group, Division 2 which was subsequently called
the Long Reach Formation (e.g., MacKenzie 1951). In 1964 Deni-
son corrected his statement by referring the Cyathaspis acadica
material to the Jones Creek Formation, on the basis of MacKenzie 's
1951 field appraisal. This interpretation was followed by Blieck
(1982).
Evidence, however, from the 1964 maps of the region by Mac-
Kenzie suggests that all the fish beds are within the older Long Reach
Formation. The position of the two formations has been confused
because in the 1950s, and on the 1964 Saint John and Hampstead
maps, MacKenzie placed the Jones Creek Formation below the

Long Reach Formation. In fact, the reverse order is the case (Berry
and Boucot 1970; McCutcheon 1981). One recent source places the
fish beds definitely within the Long Reach Formation (Smith 1966).
In the notes of the 1964 Hampstead map there is even a reference to
a new locality in the Long Reach Formation for "a primitive fish" in
a brook, northeast of Armstrong Corner. This specimen was not
identified and I have no further information about it. It may be that
the cyathaspids did not come from the same horizon as the thelo-
donts and other elements of the fauna.
In the early 1970s, Denison sent me, on request, a sample of loose
scales from one of the nodules from the New Brunswick site. I
1986 THELODUS MACINTOSH/ 3
attempted to make thin sections, but the histological structure of the

scales was disrupted by post-mortem algal or fungal attack, or
recrystallization (see PI. 1). In general shape, however, the scales
seemed identical to those of T. parvidens. I also examined a speci-
men of T. macintoshi in the British Museum (Natural History),
collected from New Brunswick by W. Graham-Smith in 1937. On
the basis of these investigations, I suggested that T. macintoshi was
virtually identical to T. parvidens, at least in its scale morphology

(Turner 1973, 1976). [I also stated that the Long Reach Formation
was the stratotype, following Denison 1956; see also Blieck 1982].
From a perusal of Stetson's figures in 1968 I decided to write a
paper on the closure of lapetus, the early Palaeozoic ocean (Turner

1970). For if T. parvidens and T. macintoshi were the same species,
then New Brunswick and the Anglo- Welsh region must have been
closely aligned in the late Silurian.

In his original work Stetson (1928, Figs. 1-3) described and fig-
ured scales he thought looked like typical T. parvidens scales but
which bore long thin extensions on the posterolateral rims of the
crowns (Figs. 2, 3). This character prompted him to distinguish T.
macintoshi as a separate species. In spring 1983 I examined both
Stetson's and Denison's collections. Specimens PF are housed at the
Field Museum of Natural History, Chicago (FMNH), MCZ at the
Museum of Comparative Zoology, and BMNH at the British
Museum of Natural History.
MATERIALS EXAMINED
MCZ 2035 HOLOTYPE Thelodus macintoshi Stetson 1928 is
T. parvidens Agassiz with T. trilobatus scale. Area at
least 30 + X 250 mm. Holotype includes thin
sections.
MCZ 2037 Paratype T. macintoshi is T. parvidens.
MCZ 2036 Paratype T. macintoshi is Loganella cf. L. ludlo-
wiensis (Gross 1967).
MCZ 13007 in part to 13015, includes the specimens from which
some thin sections were cut, are referred to T. parvid-
ens Agassiz.
MCZ 13007 in part; one nodule has Gomphonchus type scales.
MCZ 13014 includes T. bicostatus {Hoppe 1931) scales.

MCZ 13012 is at least 240 mm long.
4 BREVIORA No. 486
PF 1805 T. macintoshi is T. parvidens, area 300 X 170 mm

(Denison coll.).
PF 1 809 T. macintoshi is T. parvidens horizon A, area at least
300 X 170 mm (Denison coll.).

PF 1804 T. macintoshi is T. parvidens, horizon B (Denison
coll).
PF 1808 loose thelodont scales in coprolite includes T. par-
videns, T. bicostatus and T. trilobatus, horizon B.

PF is a coprolite containing Gomphonchus type scales.
is a coprolite with thelodont scales, T. parvidens,
horizon B.
jumbled T. parvidens scales in coprolite, (Denison
coll. 1961).
T. macintoshi -
a good articulated T. parvidens,
incomplete 150 X 90 mm in area (Denison coll.

1961).
PF 3477 and PF 1801 are acanthodians and PF 1698 is an acantho-

dian ?spine. At least three uncatalogued specimens, including 126-52
and 1969 Denison coll. FMNH, are coprolites from horizon B con-
taining well-preserved T. parvidens scales.
BMNH P. 52444. T. macintoshi (Graham-Smith coll.) is T. parvid-

ens with T. trilobatus'} scales.
1986 THELODUS MACINTOSH I
BREVIORA No. 486
Figure I. Thelodus macintoshi Stetson 1928= Thelodus parvidens Agassiz 1838.

Holotype, MCZ
2035. Close dotting: small T. parvidens scales in part, towards the
sides of the heart-shaped nodule (probably the region of the pectoral fins).
Larger
scales in center; good articulated patches shown. Asterisk mark: one scale of T.
thlohatus. Parallel bars: cracks in the nodule with slickensides. Clear rectangles:
specimen labels. Arrow points to anterior.
kESULTS
Material. Examination of Stetson's specimens and his thin sec-
tions (to which the 1928 plates do not do justice), and of Denison's
material, reveals that the material attributed to T. macintoshi is
heterogeneous: it
comprises at least two genera of thelodonts, and
one specimen is actually an acanthodian. T. macintoshi {s.s.) is
almost certainly synonymous with T. parvidens. I propose that T.
macintoshi Stetson should be formally considered a junior synonym
of T. parvidens, but I shall refer to T. macintoshi in an informal
sense to distinguish the New Brunswick material from other
material.
1986 THELODUS MACINTOSH!
Figure 2. Isolated scales from nodule PF 9603 from horizon B of Denison,

Cunningham's Brook, Nerepis, New Brunswick. A) T. parvidens. lateral view show-
ing neck riblets, PF 9603. 1 ;
B) T. parvidens. ventral view, PF 9603.2; C) T. bicos-
tatus, dorsal view of broken crown, PF 9603.3;
D) T. trilobatus, ventral and dorsal
views, PF 9603.4; E) T. trilobatus, lateral and dorsal views, PF 9603.5; F) T.
trilobatus. dorsal view of smooth crown, PF 9603.6; G) T. trilobatus, lateral and
ventral views, PF 9603.7.
8 BREVIORA No. 486
.50 mm
Figure 3. Scales on paratype of T. macintoshi. MCZ 2036. A) Loganella sp.,

two scales seen in cross section; B) Loganella sp., body scale in crown view showing
unbroken posterolateral spinelets; C-D) Loganella sp., body scales in crown view
with small anterior basal process and posterolateral neck spinelets.
The scales of the T. macintoshi specimens are large, up to 1.5 mm

long, and an advanced stage of growth with well-developed bases
in
(see Fig. 2 A, PI. lA-C); they are undoubtedly scales of a mature

animal. They differ from European examples of T. parvidens
(including the type specimen) in the clear expression of the numer-
ous riblets on the neck region (PI. 2). However, as most European
scales are waterworn to some degree it seems possible that the full
extent of neck ribbing in the type species has never been clearly seen
in the isolated scales. Gross did show one scale with 12 riblets on
one side ofthe neck (1967, PI. 1, Fig. 3 A) but their number is usually
much smaller. The slight differences in rib ornament on the neck of
scales may reflect local variation within a single population of The-
lodus parvidens.
Furthermore, some of the specimens, including coprolitic masses
and thin sections, show that scales of T. bicostatus type are part of
the T. parvidens squamation (see Fig. 2). Despite the fact that the
scales of T. bicostatus are very distinctive, both Gross (1967) and

Turner (1973, 1976) thought they might be special scales of T. par-
videns, while retaining the specific name bicostatus. One scale on
the type specimen (MCZ 2035) appears to belong to T. trilobatus
(see Fig. 1). A patch of small striated scales among the larger rhom-
boid T. parvidens scales on specimen BMNH
P52444 could also
belong to T. trilobatus. Scales of this species are also seen in copro-
litic masses (Fig. 2D-G). Thin sections of scales from the nodules
1986 THELODUS MACINTOSH I
Plate 2. Scales of Thelodus parvidens from coprolitic nodule from Cunningham's

Brook, Nerepis, New Brunswick. A) Detail of neck in lateral view showing riblets
on lower neck, PF 9603.8, approx. X60; B) Scale in lateral view, PF 9603.8,
approx. X45; C) Scale in posterobasal view showing pulp cavity, PF 9603.10,
approx. X45.
confirm the presence of T. parvidens in association with T. bicosta-

tus and T. trilobatus (see PI. 1). Thus the T. macintoshi material
confirms the synonymy of T. parvidens, T. trilobatus and T. bicos-
tatus postulated initially by Gross (1967).
The most interesting discovery in this re-examination of Stetson's
type material is the presence of at least one specimen of a loganiid.
The paratype of T. macintoshi (MCZ 2036) is not a thelodontidid,
but a specimen of Loganella sp. cf L. ludlowiensis (Gross 1967) or

perhaps cf L. martinssoni (Gross 1967). [The genus name Logania
formerly attributed to this and other loganiid species is preoccupied
—
{Logania Distant Lepidoptera; Whitley 1976). Here I propose to
use the replacement name Loganella.] The figures in Stetson's paper
depicting scales with rim spines actually represent sagittal sections
through Loganella scales (see Fig. 3). The scales in the nodule are
sometimes broken through, exposing the slit-like pulp canal and
10 BREVIORA No. 486
navicular shape of typical loganiid body scales (cf Gross 1967 e.g..
Figs. 11 I-K).
At two specimens from Nerepis, one in each of the collec-
least
tions at MCZ
and the Field Museum, are not thelodonts but
acanthodians. The scales are those of an ischnacanthid cf
Gomphonchus.
not really difficult to see why Stetson did not recognise the
It is
different thelodonts in his fauna. Before 1928 only a few cross-

sections of Thelodus scales, except those of T. parvidens, had been
illustrated. Knowledge of the histological structure of scales did not
become refined until the comprehensive study by Gross (1967), in
which he separated the genus Logania from Thelodus. The MCZ
houses the thin sections Stetson used; those cut from 2036 MCZ
show the teardrop-shaped outlines of Loganella scales cut in horiz-
ontal section, with a slit-like pulp cavity. Nine slides of nodule
pieces containing scales of T. macintoshi are housed at the Field
Museum.
In the main, these scales are well-preserved and the sections show
clearly that the scales belong to T. parvidens: some sections of the

scales are mere "ghosts," the internal structure of the dentine having
been recrystallised. One slide, PF slide 4187/52-102b, contains
cross-sections of two scales of T. bicostatus (Hoppe 1931) (see PI.
IE). Many parvidens, including the bicostatus and
of the scales of T.
trilobatus forms, exhibit clear incremental lines in the dentine; there

are from nine to in large scales and two to three in small scales
1 1
(see PI. I). The incremental lines in thelodont scales are a measure of
scale growth (Gross 1967). Whether the lines represent annual or

seasonal increments never be ascertained; if the animals were
may
living in warm
near-equatorial waters, as has been suggested for
Siluro-Devonian vertebrates on other occasions (e.g., Halstead and
Turner 1973), then it seems possible that the lines represent seasonal
fluctuations in the availability of calcium and/ or phosphate. Some

scales are also penetrated by what look like fungal hyphae, similar
to those described by Goujet and Locquin (1977) and Karatajute-
Talimaa (1978) (see PI. A, F). 1
Size of T. parvidens. Several of the nodules (e.g., those contain-

ing the holotype, see Fig. 1) contain articulated squamation which
indicates that the thelodonts were very large. Stetson was correct in
estimating that T. macintoshi ^diS longer than Turinia pagei {Powrie

1986 THELODUS MACINTOSH! 11
1870), the largest known thelodont at that time. He considered that

some eight to ten inches would need to be added to the length of the
holotype Turinia, making an estimate of 22 inches (around 550 mm)

for T. macintoshi. Study of the type specimen shows that scales
towards the center are very large, around 1.5 mm
square, and
closely packed in neat diagonal rows, whereas towards the antero-
lateral margins the scales are progressively smaller, around 0.5 mm
square and along the lateral margins the very small scales are much
disrupted (Fig. 1). I suggest that, by comparison with the type spec-
imen of Turinia pagei (see Turner 1982), the type specimen exhibits
part of the cephalothorax extending to the region of the proximal
tips of the pectoral fins. The specimen measures about 300 long mm
by 250 mm
wide. Given that the cephalothorax was probably about
one quarter to one third of total body length, this New Brunswick
thelodont might have had a length between 900 and 1200 mm. Other
nodules in the collections examined show areas of squamation at
specimen. With a length around one
least as large as that in the type
meter, these specimens of T. parvidens are the largest thelodonts

found to date. This more than confirms Stetson's prediction about
the size of the animals.
Age of the Nerepis fauna. Matthew (1888) considered the fish-
bearing beds to be of Niagaran age because of the associated fauna,

which includes Ceratiocaris pusillis and the xiphosuran Bunodella
horrida (Denison 1956). Bailey and Mclnnes (1888) reported that
Matthew regarded the beds to be about the same age as those con-
taining Palaeaspis in the United States, that is, the Medina and
Clinton Groups, and equivalent to Divisions 2 and 3 of the Anticosti
Group and Groups B and B' of Arisaig. Westoll (1958a) placed the
beds within the Wenlock, equivalent to the Lockport of the United
States.
Both the Long Reach and Jones Creek Formation beds lie within
the Mascarene Group, referred to as the Upper Silurian by the
Canadian Geological Survey (MacKenzie 1964). When MacKenzie's
maps were published in 1964 it was still not certain whether the
Jones Creek Formation lay above, or below, the Long Reach For-
mation. The latter has been dated as Upper Llandovery/ Lower
Wenlock (C6), because of its Costistricklandia- Eocoelia community
(Berry and Boucot 1970). MacKenzie (1951) had placed the Long
Reach Formation above the Jones Creek Formation, but Berry and
12 BREVIORA No. 486
Boucot, following a statement by MacKenzie, considered that it

must rest below and that the Jones Creek Formation was of Lud-
low/ Pridoli age (see also Blieck 1982). McCutcheon (1981) has reit-
erated this interpretation, which would imply a large time gap
between the two formations. I am not able to ascertain from the
literature if this hiatus is discernible in the field. On their correlation
chart Berry and Boucot (1970) show the Long Reach Formation
extending from ?Upper Llandovery to Ludlow, but based on the
brachiopod fauna, they prefer a Late Llandovery/ Early Wenlock
age. The Jones Creek Formation is possibly as old as Upper Ludlow
(Ludfordian), for, as Berry and Boucot point out, the beds contain a
Salopina community which is thought to signify Ludlow shallow
water conditions in the Welsh Borderland (Lawson 1975).
Berry and Boucot compared the Jones Creek Formation with the
Pembroke Formation of Maine. Devonian fish material has been
discovered in the Eastport Formation of this region by Denison
(Field Museum collections) but is not described yet. As the strati-
graphy and structure of the Nerepis area seem complex, perhaps a
more thorough search for fish remains in equivalent Siluro-
Devonian beds nearby in Canada and in the United States would
help clarify the succession.
The association of cyathaspids, acanthodians and T. parvidens
(including thlobatus and bicostatus varieties) and a Loganella sp. cf.

L. ludlowiensis in New Brunswick could be as old as early Wenlock
and no younger than early Downton. A similar assemblage of these
thelodont species and acanthodian scales has been found in sedi-
ments as old as Lower Wenlock in the Welsh Borderland (Turner
1973) and also in the Upper Llandovery of Norway (Turner 1984),
and it is also found throughout the Upper Ludlow and early Down-
ton of Europe (Turner 1973; Karatajute-Talimaa 1978; Marss
1982b). Marss (1982b) suggests that the pre-late Ludlow loganiid
in the Welsh Borderland is in fact L. martinssoni. The possibility
that the New Brunswick loganiid may belong to this latter species
cannot be ruled out. Cyathaspidids are known from supposed
Upper Llandovery and Wenlock sequences in the Canadian Arctic
but details of these early forms are not yet published (Denison 1964;
Thorsteinsson 1967; Dineley and Loeffler 1976). In their review of
cyathaspids Dineley and Loeffler (1976) compared Cyathaspis acad-
ica with C. banksi, known from the Ludlow and early Downton of
the Welsh Borderland.It seems likely, however, following the rea-
soning of Elliott (1978) and of Dineley and Loeffler, that cyathas-

pids occur earlier in Canada than in Europe. More work on the
Canadian cyathaspids might help clarify their relationships and

biostratigraphic significance.
DISCUSSION
Implications. As 1 predicted (Turner 1970), T. macintoshi is
very closely related if not identical to the type thelodont T. parvid-
ens. Also the type material contains T. bicostatus and T. trilobatus.
These three scale forms, T. parvidens, T. bicostatus and T. triloba-
tus, are elements of the Baltic-Anglo-East Canadian Silurian fauna

(the T. parvidens assemblage of Turner 1973). This assemblage may
represent a facies fauna indicative of the onset of "red bed"
conditions.
If the thelodonts do come from the Long Reach Formation and
the age of the fish beds is confirmed as early Wenlock, or even late
Llandovery, this will add weight to the predicted occurrence of T.
parvidens throughout the southern Laurasian region in the early

Silurian. The appearance of T. parvidens before the late Wenlock is
disputed by Karatajute-Talimaa (1978), who has not found it in any

of the older localities which she has investigated. However, the fact
that T. parvidens did appear in the late Llandovery is confirmed by
itsdiscovery in the Norwegian succession. If this is the oldest occur-
rence then there must have been the possibility of dispersal of T.
parvidens between the west Baltic, the Welsh Borderland and New
Brunswick. This would imply a shallow water connection between
at least two of these three regions in mid-Silurian times.
Theparvidens assemblage could also represent a dine, with the
T.
T. macintoshi form to the "west" of the range in New Brunswick,
and forms such as T. costatus (Pander 1856), T. sculp tills Gross

1967, and T. admirabilis Marss 1982 to the "east" in the southern
Baltic — even, perhaps, including T. marginatus Karatajute-Talimaa
1978. T. trilobatus, T. bicostatus, and T. pugniformis Gross 1967
would be included in T. parvidens as varieties because all possess a
similar histological structure. These thelodonts would be placed
within Blieck's Ichthyofacies IIA (Blieck 1982). Blieck (1982, Fig. 5)
envisaged the New Brunswick site as located within the Appalachian
channel with a possible link to the Welsh Borderland and elsewhere
14 BREVIORA No. 486
in western Europe via a narrow shallow water passage. New Bruns-

wick was almost certainly in connection with the Anglo-Welsh
cuvette, Norway and probably elsewhere in the Baltic hinterland at
some point during late Llandovery to early Downton times (see also
Turner and Tarling 1982).
Scales of T. parvidens have also been found in the Ludlow Moy-
dart Formation of Nova Scotia ((/)rvig in Boucot et al. 1974) and
those of ""Thelodus sp." in the Upper Silurian (Lower Devonian?)
Oriskany Sandstone of Nictaux Falls, Nova Scotia (Eastman 1907,
Gardiner 1966). Thus there are indications that the T. parvidens
fauna was quite widespread in eastern Canada by late Silurian
times.
Environment. Denison (1956); Robertson (1957); and White
(1958) considered that the fauna at Nerepis indicated a marginal
marine or non-marine environment. From the state of preservation
itwould seem that the animals were living in or near a quiet back-
water, possibly a supratidal pool or a lagoon. The style of preserva-
tion in the New Brunswick fish beds is virtually identical to that in
the Lower Silurian fish beds of southern Scotland (see e.g., Ritchie
1968). At Lesmahagow and Hagshaw thelodonts, anaspids, ceratio-
carids, eurypterids and rare xiphosurans are found, often contained
in nodules (considered to be coprolitic) within black shales. Some
near-complete thelodonts are preserved in large nodules, and
numerous scales occur in the center of nodules, probably the
remains of consumed thelodonts. At Cunningham's Brook these
coprolitic masses seem to be confined to one horizon (Denison's
horizon B). There or no evidence of transport seen on any of
is little
the specimens; thelodont scales are not noticeably waterworn and a
range of scale size is seen in the disarticulated scale masses in

nodules as well as in articulated squamation (see Figs. 1, 2).
Denison ( 964) discussed the state of preservation of the cyathas-
1
pids and found that one specimen, also found in a coprolite, was a
juvenile on the evidence of its thin plates and scales. This also sug-
gests that the fish were living and dying in a low-energy
environment.
The large thelodonts in the Nerepis fauna suggests that
size of the
this was an environment for these creatures which until now
ideal
have invariably been thought of as small insignificant agnathans.
1986 THELODUS MACINTOSHI 15
The thelodonts in the Lower

Silurian of Scotland, belonging to
Loganella scotica Traquair 1898, which seem to have lived in a

similar environment, also exhibit a range of size from a few centime-
ters (juveniles) to quite large individuals (up to an estimated 400
mm). If components and style of preservation
similarity of faunal
are taken into account then the Nerepis site and Scottish lower fish
beds could be of equivalent age, which would be late Llandovery to
early Wenlock (e.g., Rolfe 1973).
ACKNOWLEDGMENTS
I would thank Mrs. Ruth Romer, Dr. and Mrs. R. H.
like to
Denison, Dr. W. and M. C. Schaff for their help and

E. Schevill
hospitality in Cambridge, Mass. Dr. Denison made helpful com-

ments upon the manuscript. Dr. and Mrs. John Bolt, J. Clay
Bruner, and Dr. and Mrs. W. Turnbull gave me their help and
hospitality while I was in Chicago. The opportunity to examine
the
Field Museum material was provided by the award of a grant from
the Thomas Dee Visiting Scientist Fellowship. Dr. Godfrey S. Now-
lan helped me to obtain information on the geology of New Bruns-
wick. Dr. Roger Miles allowed me to examine and cast the BMNH
specimen then under his care. Part of this work was done under the
auspices of an Honorary Research Fellowship of the Queensland
Museum, for which I thank Dr. Alan Bartholomai and the Board of
Trustees.
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ford, Radnor, Montgomery, Caermarthen, Brecon, Pembroke, Monmouth,
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Bailey, L. W., and W. McInnes. Report on explorations and surveys in
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Berry, W. B. N., and A. J. Boucot. 1970. Correlation of the North American
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1964. The Cyathaspididae, a family of Silurian and Devonian jawless
vertebrates. Fieldiana, Geology, 13: 31 1-473.
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and associated Siluro-Devonian formations. Northwest Territories, Canada.
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Eastman, C. R. 1907. Devonic fishes of the New York formations. Memoir of the
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Elliott, D. K. 1978. New Pteraspididae from the Canadian Arctic, p. 27. PADS
Abstracts. International Symposium on the Devonian System 1978. The
Palaeontological Assoc, Univ. Bristol: 61 pp.
Gardiner, B. G. 1966. Catalogue of Canadian fossil fishes. Roy. Ont. Mus. Life
Sciences, Contrib. No. 68: 1-154.
Goujet, D., and M. V. LocQUiN. 1979. Decouverte de spores fongiques dans les
ecailles de poissons d'agnathes paleozoiques: Mycobystrovia lepidophaga

et
gen. et sp. nov. 104e Congres National des Societes Savantes, Bordeaux,
sciences. Fascicule 1: 87-99.
Gross, W. Uber Thelodontier-Schuppen. Palaeontographica, 127A: 1-64.
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Halstead, L. B. and S. Turner. 1973. Silurian and Devonian ostracoderms, pp.
67-79. In A. Hallam (ed.). Atlas of Palaeobiogeography. Amsterdam: Elsevier,
531 pp.
Hoppe, K. H. 1931. Die Coelolepiden und Acanthodier des Obersilurs der Insel
Oesel. Palaeontographica 76: 35-94.
Karatajute-Talimaa, V. N. 1978. Silurian and Devonian Thelodonts of the
USSR and Spitsbergen. Vilnius: Mosklas, 1-334 [in Russian].
Lawson, J. D. 1975. Ludlow Benthonic Assemblages. Palaeontology 18 (3):
509-525.
Mackenzie, G. S. 1951. Geol. Surv, Can. Pap. 1951-15 (out of print, not seen).
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1113A, 1 inch: 1 mile, 1:63360.
1964. Geology, Hampstead, New Brunswick. Geol. Surv. Can. Map
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Marss, T. 1982a. Thelodus admirabilis n. sp. (Agnatha) from the Upper Silurian
of the East Baltic. Eesti NSV Teaduste Akadeemia Toimetised 31, K. Geol. no.
31: 112-116.
1982b. Vertebrate zones in the East Baltic Silurian, pp. 97-105. In D.
Kaljo and E. Klaaman (eds.), Ecostratigraphy of the East Baltic Silurian.
Tallinn: Academy of Sciences Estonian SSR, Inst. Geol.
Matthew, G. F. 1888. On some remarkable organisms of the Silurian and
Devonian rocks in Southern New Brunswick. Proc. Trans. Roy. Soc. Can.,
Ottawa, Section 6(1)6, Section 4: 49-62.
1907. A new Genus and a New Species of Silurian Fish. Trans. Roy. Soc.
Can. (3), 1, section 4: 7 9.
McCuTCHEON, S. R. 1981. Revised stratigraphy of the Long Reach area, south-

ern New Brunswick: evidence for major, northwestward-directed Acadian
thrusting. Can. J. Earth Sciences, 18 (3): 646-656.
Pander, C. H. 1856. Monographic der fossilen Fischedes silurischen Systems der
russisch-baltischen Gouvernements. St. Petersburg: x -I- 1-91.
PowRiE, J. 870. 1On the earliest known vestiges of vertebrate life; being a descrip-
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Ritchie, A. 1968. New evidence on Jamoytius kerwoodi White, an important
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21-39.
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156-87.
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Glasgow Districts. Glasgow Geol. Soc. Glasgow: 181 pp.
Smith, J. C. 1966. Geology of Southwestern New Brunswick, pp. 1-12. In W. H.
Poole (ed.). Geology of parts of Atlantic provinces. Guidebook, Geol. Assoc.
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Stetson, H. C. 1928. A New American Thelodus. Amer. J. Science, 16: 221-31.
Thorsteinsson, R. 1967. Preliminary note on Silurian and Devonian ostraco-
derms from Cornwallis and Somerset Islands, Canadian Arctic Archipelago, pp.
45-47. In Evolution des Vertebres. Problems actuel de Paleontologie, 163,
Paris: C. N. R. S. Masson et Cie.
Traquir, R. H. 1898. Report on Fossil Fishes. Summary of Progress for 1897,

Geol. Surv. G. B., 72-76.
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Nature, London, 227, No. 5253: 90.
1970b. Fish help to trace continental movements. Spectrum, Bull. Cent.
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Catalogus I: Animalia, Pars 122: 14 pp. 1
1982. A new articulated thelodont (Agnatha) from the Early Devonian of

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B R E V I R A
us ISSN 0006 9698
Cambridge, Mass. 29 August 1986 Number 487
THE IDENTIFICATION OF l.XR\ AlL PARASUDIS

(TELEOSTEl, CHLOROPHTHALMIDAE); WITH
NOTES ON THE ANATOMY AND RELATIONSHIPS
OF AULOPIFORM FISHES
Karsten E. Hartel and Melanie L. J. Stiassny'
Abstract. Larvae and transforming individuals of Parasudis are identified and

described for the first time. The phylogenetic integrity of the family Chlorophthalmi-
dae is investigated and restricted to the sistergenera, Parasudis and Chlorophthal-

mus. Twoderived morphological characters support the monophyly of an extended
Ipnopidae which now includes the genus Bathysauropsis. The "rostral cartilages" in
aulopiform fishes are reviewed and the homology of these and associated structures
throughout the lower Neoteleostei is questioned.
INTRODUCTION
Parasudis truculenta (Goode and Bean, 1895) is a relatively com-
mon offshore fish found from off northern New England to equato-
Adult specimens have been taken only in bottom trawls
rial Brazil.
at depths of ca. 180 to 480 m, but dietary studies suggest that Para-
sudis moves off the bottom to feed in midwater (Mead 1966). The
species hermaphroditic, and ripe specimens have been collected
is
from February to May (Mead 1960). Neither the eggs nor the larvae
of this species have yet been described (Okiyama 1984).
The purpose of this paper is to describe larval and transforming
Parasudis, and to comment on aspects of anatomy and relation-
ships. There is considerable confusion in current literature regarding

02138. Order of authorship is alphabetical. ''*;
SEP 1<
2 BREVIORA No. 487
the systematics and classification of Rosen's (1973) Aulopiformes

(seeJohnson 1982; Okiyama 1984; Rosen 1985; Stiassny 1986). The
resolution of this problem is beyond the scope of this paper and for
the purpose of this study we adhere to Rosen's classification of the
Order (1973:509).

Specimens are from the Woods Hole Oceanographic Institution
(WHOI) collections housed at the Museum of Comparative Zool-
ogy, Harvard University (MCZ); the Atlantic Research Centre, New

Brunswick (ARC); the DANA collections of the Zoological
Museum, University of Copenhagen (ZMUC); the National Mu-
seum of Natural History, Washington, D.C. (USNM); the British
Museum of Natural History (BMNH) and the University of Miami
(UMML). The specimens used in this study are listed in appendix
under material examined.
Counts and measurements follow Hubbs and Lagler (1964). For
osteological examination, selected specimens were cleared and
stained following the procedure of Dingerkus and Uhler (1977);
these specimens are indicated "c.s." under material examined.
Anatomical drawings were made with the aid of a Zeiss SV-8 stereo-
microscope with a camera lucida attachment. Osteological and syn-
desmological nomenclature follows Stiassny (1986).
DIAGNOSIS
Parasudis larvae are characterized by the distinctive bulbous
shape of the snout, the pattern of snout, opercular and caudal pig-
mentation, as well as by a particular basihyal form and dentition.
The larvae are further distinguished from those of other chloro-
phthalmoid taxa by having 38 to 39 vertebrae and/ or myomeres
(Okiyama 1984:208).
DESCRIPTION
Larvae
(Figs. lA, BandC)
Preflexion or flexion larvae are not present among the material
examined, and the following description is based on 85 postflexion
1986 LARVAL PARASUDIS RELATIONSHIPS 3
specimens ranging from 10.6 to 80 mm

standard length (SL). Spec-
imens of 15.4 and 30.6 mm
SL are illustrated in Figs. lA-C. The
body is elongate, oval to round in cross section at maximum depth;
maximum depth 6.4 to 9.4 times in SL; maximum body width sub-
equal or equal to maximum body depth (7.8 to 1 1.7 in SL). Head
length (HL) 2.6 to 3.9 times in SL, probably negatively allometric.
Bulbous snout rounded in anterodorsal profile, 2.2 to 3.0 times in
HL. Eyes large, 3.5 to 5.0 times in HL. The eyes often appear
stalked or partially stalked (Fig. IB). The presence (or absence) of
stalked eyes may be a result of damage incurred during collection;
however, stalked and non-stalked specimens do appear in the same
samples. The eyes of both larval and transforming individuals lack
the "keyhole" shaped aphakic space characteristic of adult
chlorophthalmids.
All larvae examined appear to have a full complement of fin
elements, although, the fragile fin rays are often damaged. The
origin of the dorsal fin (10 rays) is slightly posterior to the base of
the pectoral fin (17 rays). The narrow-based adipose fin is posi-
tioned above the second or third anal ray (8-9 anal rays). The pecto-
rals are at mid-body level with their bases almost under the thin
opercular flap. Pelvic fin (9 rays) origin is below the anterior half of
the dorsal fin. The caudal fin is forked even in the smallest speci-
mens. Procurrent caudal rays are present and increase in number
with growth. The anus is located well caudad of the pelvic fin base
but closer to the insertion of the pelvic than to the insertion of the
anal fin.
The dentition remains relatively constant throughout the larval
period. Both the premaxillae and the dentaries bear single rows of
evenly spaced caniniform teeth (Fig. 2). Anteriorly, these larval
teeth are recurved, but are markedly retrose in the posterior aspect
of both upper and lower jaws. Two to three small recurved
caniniform teeth are implanted along the anteriolateral aspect of the
dentary (Fig. 2).
In most specimens a few small caniniform teeth are inserted along
the body of the palatine cartilages, and in the ventral hypobranchial
apparatus a markedly elongate basihyal cartilage bears small
recurved fang-like teeth on its anterior margin (Fig. 3A).
BREVIORA No. 487
Figure 1. Parasudis truculenta. (A) Lateral view of larva 15.4 mm SL (MCZ

62400). (B) Dorsal view of head of same specimen. (C) Lateral view of larva 30.6mm
SL (MCZ 62399). (D) Lateral view of transforming juvenile 85 mm SL (MCZ 62401).
Note that the dotted lines on the body in (A) and (C) represent myosepta while the
solid lines in (D) represent the pigmented edges of scale pockets. Drawn by S.
Landry.
1986 LARVAL PARASUDIS RELATIONSHIPS
PMX
Figure 2. Isolated buccal jaws of larval Parasudis (MCZ 62397, 38 mm SL).

Abbreviations: Dent, dentary; Mx, maxilla; Pmx, premaxilla.
Pigmentation
Body color in preserved specimens is a uniform opaque white

although very infrequently a light tan or brown wash is present. Live
coloration is unknown, but based on observations of live pretrans-
formation Chlorophthalmus (Hartel personal observation), living
Parasudis larvae are presumed to be almost translucent in seawater.
Opercular pigmentation covers an area almost as large as that of
the eye and at times may be quite dark. Opercular pigmentation is
present in the smallest specimen examined.

Initially, asmall group of melanophores is situated on the caudal
peduncle above the midline. With growth (>30 SL) the pig- mm
mented area becomes roughly triangular in outline with the pos-
terior lower corner extending below the midline (Fig. IC). A melan-
ophore distinct from the triangular patch and situated in the upper
corner of the caudal base is found in specimens greater than 22 mm
BREVIORA No. 487
Figure 3. Parasudis. (A) Ventral gill arch elements and hyoid apparatus of
larva 30 mm SL (MCZ 62398). (B) First epibranchial and associated structures of
adult specimen (MCZ 40561). Chlorophthalmus. (C) Ventral gill arch elements of
larva 22 mmSL (MCZ 62403). (D) First epibranchial and associated structures of
adult specimen (MCZ 40509).
SL, and a second is usually developed at the lower corner of the

caudal base by 30 mm SL.
Pigmentation in the snout region first appears as a single melano-
phore above the anterior end of the maxilla at 14 mm
SL. Addi-
tional melanophores areadded posteriorly in a line just above the
maxilla and by 50 mm SL, the row reaches a point almost under the
anterior margin of the orbit.
body pigmentation begins as one or two mid-body
Lateral
melanophores at 22 mm SL. The melanophores increase laterally in
two groups. Four are present under the dorsal, and three are under
the adipose fin by 24 mm SL. At 40 mm SL the melanophores,
which are always found directly over a myomere, form an almost
continuous band. There are one, two or sometimes three melano-
phores per myomere.
Deep pigment is present in the dorsal midline as a small dark spot
just posterior to the dorsal fin. Ventrally a similar small midline spot
isfound at mid-body. An additional line of deep pigment is present
along the midventral caudal peduncle. All of these deeper markings
become less noticeable with growth as the muscle tissues become
denser. Internal pigment is often visible through the bodywall below
the base of the pectoral fin (Fig. 1 A). This darkened area is densely
pigmented mesentery supporting the viscera and areas of visceral

peritoneum wrapping partially around the developing anterior
viscera.The pigmented mesentary extends caudad almost to the
level of the anal fin origin (Fig. 4).
Transforming specimens
(Fig. ID)
This description is based upon two specimens, 75.5 mm SL (MCZ

57922) and 85 mm SL (MCZ 62401), illustrated in Fig. ID. The
larger specimen is only 5 mm larger than the largest available larval
specimen (ARC 6879), yet marked morphological changes are evi-
dent.
General body proportions remain constant, but the form of
the head is altered by the loss (via reabsorption?) of the character-
istic bulbous larval snout. The smaller individual retains a few of the
larval retrose jaw and has a strong basihyal dentition. In the
teeth
larger transforming fish the

jaws are edentulate, apparently having
lost the larval teeth and not yet having developed the juvenile/ adult
BREVIORA No. 487
P.m
Figure 4. Internal pigmentation of larval Parasudis (based on two specimens;

MCZ 63055, 40 mm SL and MCZ 63056, 31 mm SL). Drawn by S. Landry.
Abbreviations: B.c, body cavity; P.m, pigmented mesentery; V.p, visceral

peritoneum.
dentition. A few small palatine teeth are present, and small teeth are
embedded in the dermis overlying the basihyal element. As the
smaller of the two transforming specimens is badly damaged and
contorted, the remaining description is based solely upon the 85 mm

SL specimen.
The diagnostic larval pigmentation pattern is faint but the rem-
nants of snout, opercular, lateral and caudal pigmentation are still
discernible. Patches of scales are present and, as is indicated in Fig.
ID, these have a distinctive pattern of implantation. Each scale
is located on a continuous
pocket flap of skin which is strongly
pigmented along its distal margin. The resultant "herringbone"
pattern is clearly visible in adult specimens (Mead 1966: Fig. 47)
and where scales have been lost, corresponds to approximately two
pigment lines per scale.

Transformation apparently occurs rapidly. We have seen a fully
transformed juvenile (MCZ 41840) the same length as the larger
transforming larva. The fully transformed fish has the definitive
adult dentition (see Mead 1966), is scaled, and has lost almost all
traces of larval pigmentation. During transformation the snout
becomes increasingly flattened and "shrunken," lending the fish its
characteristic "duckbilled" appearance. Apparently, a loss (reab-
sorption?) of the somewhat gelatinous larval body tissues results in a

more attenuated body form in the recently transformed juvenile.

Halliday (1968: Fig. 1) illustrates a 99.0 mm SL transformed juve-
nile that shows initial body deepening.
The internal pigmentation of the anterior viscera is present in
transforming specimens. Apparently during transformation, pig-
ment migrates from the visceral peritoneum onto the parietal peri-
toneum, and the pigmentation of the caudad extension of the
mesentery greatly reduced. In fully transformed juveniles the pa-
is
rietal peritoneum is strongly pigmented throughout the abdominal

cavity but little pigment is associated with the visceral
peritoneum.
We have examined a number of transforming Chlorophthalmus
specimens from both the North Atlantic Ocean and the Coral Sea.
In each of these specimens, tranformation occurs at a considerably
smaller size (ca. 35 to 40 mm
SL) than that observed in our Para-
sudis specimens.
DISTRIBUTION
The western Atlantic specimens we have examined come from
areas off the Brazilian coast to areas east of the southern tip of Nova
Scotia (Fig. 5). Included in our study is one eastern Atlantic
specimen (MCZ 62402). Preliminary morphometric analysis of this
larva and its eastern Atlantic locality indicate that it may represent
Parasudis fraser-brunneri (Poll 1953). However, due to a lack of
additional material for detailed study and the close morphological
resemblance of P. fraser-brunneri to P. truculenta (see Mead
1966:184), we are unable to confirm the specific identity of the
eastern Atlantic specimen.
All of the larval Parasudis were taken in midwater trawls. Speci-
mens from the WHOIopening and closing nets (MOC 10 and 20)
indicate that the larvae are most commonly taken above 150 m (five
collections: 70 to m; to 100 m; 70 to 140m; to 203 m; 70 to 150
m). Information from non-closing nets indicate similar distribu-
tions. However, other collections taken with non-closing nets indi-
cate that the larvaemay approach the surface rather closely, since
Parasudis larvae are found in collections taken above 50 m. Except
for a few larvae from the relatively shallow Straits of Florida, all
specimens were taken in trawls over ocean bottoms from 1937 to

4934 m deep.
10 BREVIORA No. 487
4U
Figure 6. Parasudis. (A) Posterior region of the neurocranium, vertebral

column and predorsal bones of adult specimen (MCZ 40561). (B) Larva 30 mm SL
(MCZ 62398). Chlorophthalmus. (C) Posterior region of the neurocranium, vertebral
column and predorsal bones of adult specimen (MCZ 41444). (D) Larva 22 mm SL
(MCZ 62403). Arrows indicate predorsal bones.
12 BREVIORA No. 487
Johnson (1982) discussed predorsal (=supraneural) number among

aulopiforms and noted a trend towards predorsal reduction
throughout the assemblage. He cited notosudids (=scopelosaurids)
and Omosudis as other taxa with a single predorsal. Sulak (1977)
stated that ipnopine genera bear one of two predorsals.
The anal position and lack of anal pigmentation during trans-
formation of Parasudis contrasts with that of Chlorophthalmus
where the anus is situated closer to the pelvic base (see Mead 1966),
and is highly pigmented during transformation. Other pigment
characters of the anterior visceral cavity separate Parasudis and
Chlorophthalmus. In Parasudis the larval pigmentation is confined
to the visceral peritoneum and associated mesentery, while in Chlo-
rophthalmus it is a discrete patch closely associated with the parietal

peritoneum which lines the body cavity. In addition, larval Parasu-
dis differs from larval Chlorophthalmus by the presence in the
former of a caudad extension of pigmented mesentry (Fig. 4). Deep
visceral pigmentation in other aulopiforms is discussed by Johnson
(1974).
COMPARISON WITH OTHER AULOPIFORM TAXA

To assess the phylogenetic significance of certain features de-
scribed in Parasudis we compare them here with those found in
Chlorophthalmus and other aulopiform taxa. Clearly, the following
anatomical comparisons are far from exhaustive, and we have
selected only those features which appear to be phylogenetically
informative at the present level of analysis.
Rosen (1971) found basihyal teeth of a similar, although larger,
form and distribution in larval Chlorophthalmus (see also our Fig.
3C), as well as in the ipnopid larvae he described (see Okiyama 1981
for other ipnopid larvae). Sulak (1977) described comparable basi-
hyal dentition in larval Bathypterois, and Bathytyphlops. Rosen

(1971) noted that a similar basihyal dentition is unknown in any
adult fish (although the pattern is approximated in Glossanodon).
Sulak (1977:80) stated that the ". .condition appears to be unique
.
to the juvenile stages of basal myctophiform fishes." However, until

the full limit of the distribution of this type of larval dentition are
known, particularly in larval aulopids and neoscopelids, we can
draw little phylogenetic inference from its occurrence in Parasudis
larvae.
Basihyal morphology easily distinguishes Chlorophthalmus from

Parasudis, which in Chlorophthalmus is short and strongly spatu-
late (Figs. 3A, C), and
Parasudis is elongate. These differences,
in
although less
markedly than in larvae, are present in the ossified
adult elements and help to distinguish the taxa (compare Figs. 7B,
C). Basihyal morphologies similar to those of larval Chlorophthal-
mus are found in the larvae of Bathypterois (Sulak 1977; personal
observation) and other ipnopids (Rosen 1971; Okiyama 1981), as

well as in adult Aulopus, Bathysaurus and Scopelosaurus (Bertelsen
et al, 1976; personal observation). Thus, based upon its limited
distribution, we interpret the broad and elongate basihyal of larval
Parasudis as an autapomorphic feature.
Interestingly, adult ipnopids share a derived condition of the basi-
hyal element, which is relatively tiny [or entirely absent in some
bathypteroids (Mead 1966)] and obliquely aligned (Fig. 7A). A sim-
ilar condition of the basihyal present in adult Bathysauropsis,
is
where unfortunately the larval" condition is unknown (Okiyama

1984).2
Certain stomiatoids have a similar basihyal arrangement, how-
ever, we agree with Weitzman (1974) that the presence of a moder-
ately sized and horizontally aligned basihyal is primitive for the
Stomiiformes as a whole. Among alepisauroids (sensu Rosen 1973),
a small (toothed) basihyal is found in certain scopelarchid taxa
(Johnson 1974) and in Bathysaurus, but a small vertically aligned
basihyal appears to be restricted in its distribution among aulopi-

forms to adult Bathysauropsis and the Ipnopidae.^ The common
occurrence of this basihyal morphology suggests an immediate
common ancestry between them.
Corroboration of the phyletic integrity of a grouping comprising
Bathysauropsis and the ipnopids is found in the degree of develop-
ment of the branchiostegal membranes and gular fold. Mead
(1966:115) described the region in bathypteroids in the following
^We have been able to examine only two specimens of Bathysauropsis gracilis but
have kindly been given access to notes and radiographs made by K. Sulak after his
examination of specimens of B. gigas and B. mayanus.
^Following Johnson (1982) and Okiyama (1984) the Ipnopidae is taken here to
include the Bathypteroidae of Mead (1966) and is equivalent to the Bathypteroidae of
Marshall and Staiger (1975) and the sub-family Ipnopinae of Sulak (1977) and
Nelson (1984).
14 BREVIORA No. 487
Figure 7. Basihyal and associated structures in adult (A) Bathypterois (MCZ

57624). (B) Chlorophthalmus (MCZ 41444). (C) Parasudis (MCZ 40561).
Abbreviations: Bbl-2, basibranchials 1-2; Bhy, basihyal; Cbl, ceratobranchi-

al
terms which apply equally to Bathysauropsis and the ipnopids:

"Branchiostegal membranes broad and separate from one another
and from the isthmus, overlapping anteriorly, and crossed near the
tip of the jaw by a thick but narrow gular fold ..." This morphology
appears to be unique among aulopiforms and although a poorly
developed gular fold present in some stomiiform fishes (e.g.,

is
Gonostoma and Polymetme) it is not as elaborate as that found in

Bathysauropsis and the ipnopids.
Chlorophthalmus shares with Parasudis the pigmented scale
pocket skin flap character. In Parasudis the dorsal and ventral
scale rows converge rostrad, "herringbone" fashion, upon the lateral
line (Mead 1966; Fig. 47), while in Chlorophthalmus, dorsal and
ventral scale rows diverge from the midline resulting in a slightly
"zig-zag" pattern (Mead 1966; Fig. 46). This highly distinctive feature
of squamation is unique to Parasudis and Chlorophthalmus, and
is here interpreted as a synapomorphy uniting the two genera.
In the eyes of adult Chlorophthalmus (Theisen 1965; Fig. 3), as

well as in adult Parasudis, a "keyhole" shaped aphakic space is
present. Although well-developed aphakic spaces are

not uncom-
mon to the eyes of many benthic marine fishes (Munk 1966; Mar-
shall 1971), a "keyhole" shaped space is restricted to aduks of these
two genera. A similar space is absent in the eyes of Bathysauropsis

and it is here interpreted as a synapomorphy of Parasudis and
Chlorophthalmus.
RELATIONSHIPS
In reviewing the literature pertaining to Parasudis and its sup-
posed relatives, it is clear that many problems remain (Johnson

1982; Rosen 1985; Stiassny 1986). However, at the intrafamilial
level, aside from the suggestion that Parasudis may share a close
phylogenetic relationship with paralepidid alepisauroids (Mead
1966), most authors who have considered the chlorophthalmids
(e.g., Gosline et al. 1966; Marshall and Staiger 1975; Sulak 1977;
Nelson 1984; Stiassny 1986) imply that Chlorophthalmus and Para-
sudis are closely related, and our own investigation confirms a sister-
group relationship between these two taxa. The genus Bathysaurop-
sis is poorly known anatomically, and consequently, its phylogenetic
placement is less clear. Mead (1966) suggested that further investiga-
tion of this genus may indicate a need for familial reallocation, a
finding that is corroborated by this study. We propose that the

notion of an "intermediate" or "transitional" position of Bathysau-
ropsis,forming a "link" between Parasudis and Chlorophthalmus
on the one hand and the ipnopids on the other, and thus serving as
". .the primary basis of the present incorporation of the ipnopine
.
genera into the Chlorophthalmidae" (Sulak 1977:64) be replaced by

a phylogenetic scheme in which Bathysauropsis is removed from the
Chlorophthalmidae and incorporated within the Ipnopidae (Fig. 8).
This relocation of Bathysauropsis is based on our observations of
the presence of a small, obliquely aligned basihyal and a well-
developed gular fold (Fig. 8: characters one and two). The position
of Aulopus in relation to these taxa is here unresolved; larval
aulopid material was unavailable for this study. Elsewhere Stiassny
(1986) argued that Chlorophthalmus, Parasudis and Aulopus (but
not Bathysauropsis and the other ipnopid genera) share with cte-
nosquamate fishes an advanced type of palatine/ maxillary associ-
ation and morphology (see also Rosen 1985 for a consideration of
aulopid relationships). For this reason Aulopus is included in our

cladogram, albeit in an unresolved position (Fig. 8).
16 BREVIORA No. 487
"AULOPOIDEI" CTENOSQUAMATA
.^^
Figure 8. Cladogram of eurpterygian relationships, incorporating the resuhs of

the present study. Characters are the presence of; 1) A small obliquely aligned basi-
hyal bone; 2) A thick well-developed gular fold; 3) A "keyhole" shaped aphakic space;
4) A peculiar scale pocket morphology and pigmentation; A) See Rosen (1985) for
synapomorphies uniting the Aulopoidei with the Ctenosquamata; B) See Stiassny
(1986) for synapomorphies uniting Aulopus, Chlorophihalmus and Parasudis with
the Ctenosquamata; C) See Lauder and Liem (1983) for synapomorphies uniting the
Ctenosquamata. Note: Following Rosen (1973), "other aulopoids" are the Bathy-
sauridae and Notosudidae. "Other Ipnopidae" are Bathypterois, Bathytyphlops,
Balhymicrops and Ipnops.
Significance of the Rostral Cartilage
Given this admittedly tentative scheme of relationships for the

chlorophthalmids, it is extremely interesting to note the highly var-
ied condition of the so-called "rostral cartilage" in these and in other
aulopiform taxa. The "rostral cartilage" of both larval and adult

Parasudis is a single median structure which is bound by a well-
developed ethmo-rostroid ligament to both the ethmoid and pre-
maxillae (Figs. 9 A, B). The attachment of the cartilage to the ethmoid
is strong in the larval fish and becomes weaker with growth. In
Parasudis, the rostral cartilage stains normally with Alcian Blue and
istherefore presumably fully chondrified hyaline cartilage. In con-
trast, the "cartilage" in Chlorophthalmus (Figs. 9C, D; Rosen 1985;
Fig. 40C) is paired, and the "cartilages" are associated with the
symphyseal processes of the premaxillae. An ethmo-rostroid liga-
ment is lacking, but the whole region is invested with loose connec-
tive tissue fibers. These rostral structures do not stain normally with
Alcian Blue; staining is weak and diffuse. Similarly, in Aulopus
(Rosen 1985; Fig. 41C; Stiassny 1986; Fig. 6) the single median
structure does not stain as hyaline cartilage and is highly fibrous.
Theisen (1965) illustrated paired rostral structures
in Ipnops similar
to those of Chlorophthalmus. Rosen (1985) also illustrated paired
"rostral cartilages" of an apparently remarkably similar type and
arrangement in the stomiiform, Maurolicus. Other stomiiform taxa

examined in this study either lack a "rostral cartilage" entirely (e.g.,
Photichthys), or bear a single median structure ligamentously

bound to the ethmoid region (e.g., Gonostoma and Diplophos). We
have been unable to locate "rostral cartilages" in either Bathypterois
or Bathysauropsis. In both of these genera the premaxillae are
bound to the ethmoid region by a well-developed ethmo-rostroid
ligament. We interpret the similarities between the rostral morphol-
ogies of Chlorophthalmus and Ipnops (and Maurolicus) to be
homoplasious. To assess them otherwise, that is, to argue for the
phyletic alignment of Chlorophthalmus with Ipnops (and Mauroli-
cus) on the basis of this one "rostral cartilage" character would
require losses and reversals in a series of other characters (see Fink
and Weitzman 1982; Marshall and Staiger 1975).
Bertelsen et al. (1976) described and illustrated the rostral mor-
phology of certain notosudids where a large (often ossified) "rostral
cartilage" is bound to the ethmoid and the premaxillae by a well-
developed ethmo-rostroid ligament similar to that of the stomiiform,
Gonostoma (Stiassny 1986), or to that illustrated here for Parasudis.
Rostral morphologies are also confusing among other aulopiform
lineages; for example, among synodontids, Synodus (Fig. lOA)
bears a large medial and fully chondrified "rostral cartilage"
strongly bound to the ethmoid. Saurida (Fig. lOB), on the other
hand, lacks the structure as do both Harpadon and Bathysaurus.
The paralepidid Sudis bears small paired hyaline cartilages, which
in adults are associated with the symphyseal processes of the pre-
18 BREVIORA No. 487
PMX
''R.CART"
Figure 9. Parasudis. (A)Ethmovomer and upper jaws of adult specimen in

dorsal view (MCZ 40561). (B)Ethmovomer of larva 30 mm SL (MCZ 62398) in
dorsal view. Chlorophthalmus. (C) Ethmovomer and upper jaws of adult specimen
(MCZ 40564) in dorsal view. (D) Ethmovomer of larva 22 mm SL (MCZ 62403) in
dorsal view.
maxillae. In larvae these structures appear more strongly bound to

the ethmoid. Finally, in Alepisaurus a condition much like that
described in Salmo (Fink and Weitzman 1982; Rosen 1985) per-
tains. Paired hyaline cartilages are strongly bound to the inner faces
of the premaxillary symphyseal processes.
Even from this cursory review it is clear the homologies of rostral
morphologies and other associated structures throughout the lower
Neoteleostei are questionable. By the level of the Acanthomorpha,
the situation has stabilized such that there appears little doubt of the
homology of the single median chondrified rostral cartilage strongly
bound to the premaxillary ascending processes by a well-developed
maxillo-rostroid ligament (Stiassny 1986). Until the distribution
and homologies of the non-acanthomorph rostral structures are bet-
ter understood, perhaps the term rostral cartilage should be re-
stricted to that structure in the Acanthomorpha. When referring to
the various rostral structures in non-acanthomorph neoteleosts
before their homologies are resolved, the term "rostral cartilage" or
"rostral structure" should be employed within quotation marks.
ACKNOWLEDGMENTS
We would like to thank the following individuals and institutions
for the loan of material and /or helpful information and comments:
PMX
PMX
R.CART
ME.P
Figure 10. Ethmovomer and upper jaw of adult specimens of (A) Synodus
(MCZ 47490). (B) Saurida (MCZ 561 11).
Abbreviations: Fr, frontal; Me, mesethmoid; Me.p, mesethmoid process; Mx,

maxilla; Pal, palatine; Pmx, premaxilla; "R.cart", "rostral cartilage". Ligaments: II,
ethmo-rostroid; IV, median palato-maxillary; XI, ethmo-maxillary; XII, palato-

premaxillary.
20 BREVIORA No. 487
(ZMUC), R. H. Gibbs and R. P. Vari (USNM), R. K.

E. Bertelsen
Johnson (FMNH), D. F. Markle (formerly ARC), K. J. Sulak
(ARC), J. Webb (Boston University), and A. Wheeler (BMNH).
For their critical review of the manuscript we are grateful to J. E.
Craddock (WHOI), G. D. Johnson (USNM), and K. F. Liem
(MCZ). The late Sally Richardson offered invaluable comments on
this paper and we dedicate our first work on larval fishes in her
memory. We
are especially grateful to Sally Landry for her skillful
preparation of Figures 1 and 4.

This research was carried out during the tenure of NSF BSR
84-07449 (MLJS). NSF DEB 77-23726 (K. F. Liem) provided sup-
port for the transfer, sorting and curation of the WHOI specimens
used in the study.
LITERATURE CITED
Bertelsen, E., G. Krefft and N. B. Marshall. 1976. The fishes of the family
Notosudidae. Dana Rept., 86: 3 14. 1
DiNGERKUS, G. AND L. D. Uhler. 1977. Enzyme clearing of Alcian Blue stained

whole small vertebrates for demonstration of cartilage. Stain Tech., 52(4):
229-232.
Fink, W. L. and S. H. Weitzman. 1982. Relationships of the stomiiform fishes
(Teleostei), with a description of Diplophos. Bull. Mus. Comp. Zool., 150(2):
31-93.
Gosline, W. a., N. B. Marshall and G. W. Mead. 1966. Order Iniomi. In
Fishes of the Western North Atlantic. Mem. Sears Fdn. mar. Res., 1: 1-18.
Halliday, R. G. 1968. Occurences of Parasudis trunculentus (Goode and Bean)
1895 (Iniomi: Chlorophthalmidae) off La Have Bank, Nova Scotia. J. Fish. Res.
Bd. Canada, 25(2): 421-422.
HuBBS, C. L. AND K. F. Lagler. 1964. Fishes of the Great Lakes Region. Ann
Arbor: Univ. Michigan Press, 213 pp.
Johnson, R. K. 1974. A revision of the alepisauroid family Scopelarchidae (Pis-
ces:Myctophiformes). Fieldiana (Zool.) 66: 1-249.

1982. A revision of the alepisauroid families Evermannellidae and Scope-
larchidae: Systematics, morphology, interrelationships and zoogeography. Field-
iana (Zool.) N. S. 12: 1-252.
Lauder, G. V. and K. F. Liem. 1983. The evolution and relationships of the
actinopterygian fishes. Bull. Mus. Comp. Zool., 150: 95-197.
Marshall, N. B. 1971. Explorations in the lives of fishes. Cambridge: Harvard
Univ. Press, 204 pp.
Marshall, N. B. and J. C. Staiger. 1975. Aspects of the structure, relationships

and biology of the deep-sea fish Ipnops murrayi (Family Bathypteroidae). Bull.
Mar. Sci., 25(1): 101-111.
Mead, G. W. I960. Hermaphroditism in archibenthic and pelagic fishes of the

order Iniomi. Deep-Sea Research, 6: 234-235.
1966. Family Chlorophthalmidae. In Fishes of the Western North Atlan-
tic, Mem. Sears Fdn. mar. Res., 1: 162-189.
MuNK, O. 1966. Ocular anatomy of some deep-sea teleosts. Dana Rept., 70:1-62.
Nelson, J. S. 1984. Fishes of the World. 2nd Ed. New York: Wiley and Sons,
ix + 523 pp.
Okiyama, M. 1981. A larval Ipnops and its possible metamorphosis. Jap. J. Ich-
thyol., 28(3): 247-253.

1984. Myctophiformes: Development. In H. G. Moser, (ed.-in-chieO,
Ontogeny and Systematics of Fishes. Spec. Pubis. Am. Soc. Ichthyol. Herptol.,
No.l: 206-218.
Rosen, D. E. 1971. The Nacristiidae, a Ctenothrissiform family based on juvenile
and larval scopelomorph fishes. Am. Mus. Novit., 2452: 1-22.
1973. Interrelationships of higher euteleostean fishes. In Greenwood, P.
H., R. S. Miles, and C. Patterson (eds.). Interrelationships of Fishes. Zool. J.
Linn. Soc. 53 Suppl. 1: 397-513.
1985. An essay on euteleostean classification. Am. Mus. Novit., 22877:
1-45.
Stiassny, M. L. J. 1986. The limits and relationships of the acanthomorph tele-
osts. J. Zool., Lond., (B)l: 1-50.

SuLAK, K. J. 1977. The systematics and biology of Bathypterois (Pisces, Chlo-
rophthalmidae) with a revised classification of benthic myctophiform fishes.
Galathea Rept., 14:49-108.
Theisen, B. 1965. On the cranial morphology of Ipnops murrayi Gunther, 1878
with special reference to the relations between the eyes and the skull. Galathea
Rept., 8:7-17.
Weitzman, S. H. 1974. Osteology and evolutionary relationships of the Sternop-
tychidae, with a new classification of stomiatoid families. Bull. Am. Mus. Nat.
Hist., 153(3): 329-478.
APPENDIX
MA TERM L EX A MINED
Parasudis material
The single figure given in parenthesis after the catalogue number
is SL; if more than one specimen is registered the number of speci-
mens is given and followed by their size range.
P. truculent a
Larvae: ARC uncat. (42.8mm), 42°27'N 58°27'W, 0-300 m, 4

Apr. 1979, USSR Belogorsk, 35-4. ARC 6729 (37.3 mm), 40°
sta.
22'N 62°38'W, 0-200 m, 20 Mar. 1979. ARC 6732 (28.0 mm, c.s.), 40°
22 BREVIORA No. 487
Sl'N 6r34'W, 0-50 m, 21 Mar. 1979. ARC 6845 (41.2 mm), 39°44'N
56°59'W, 0-200 m, Apr. 1979. ARC 6879 (80.0 mm), 42°27'N 58°
1
27'W, 0-300 m, 4 Apr. 1979. MCZ

58531 (28.5 mm), JEC 7755, 1° 1
12'N 53°49'W, 60 m, 27 Mar. 1977. MCZ 62397 (38.0 mm, c.s.), JEC
7715, 0°13'N 35°44'W, 90-lOOm, 15 Mar. 1977. MCZ 62398 (30.0
mm, C.S.), JEC 7750, 10°48'N 52° 17'W, c.80 m, 26 Mar. 1977. MCZ
62399 (30.6 mm), JEC 7753, 10°48'N 52°17'W, c.l40 m, 27 Mar.
1977. MCZ 62400 (15.4 mm), MOC
10-118, 3, 34° 12'N 75°00'W,
70-0 m, 14 Aug. 1978. MCZ 63055 (40.0 mm), JEC 7737, 38°55'N 72°
25'W, 0-825 m, 3-4 Sept. 1976. MCZ 63056 (31.0 mm), JEC 7738,
38°5rN 72°27'W,0-750m, 4 Sept. 1976. Plus an additional 32 uncat-
alogued MCZ specimens (11.7-49.1 mm) from the WHOI collec-
tion. USNM 258610 (20.0 mm), 31°50'N 63°56'W, 40 m, 23 Feb.
1972.
(ZMUC) DANA 1166V (11, 14.6-24.1 mm), 100 meters of wire
out (mw) andDANA 1166VI (2, 14.3-21.2 mm), 50 mw, both 10°
16'N 40°4rW, 11 Nov. 1921. DANA 1168IV (41.4 mm), 300 mw,
and DANA 168VI (22, 13.8-23.9), 50 mw, both 9°30'N 42°4rW, 12
1
Nov. 1921. DANA 190VIII (13.1 mm), 17°58'N 64°45'W, 100 mw,
1
13 Dec. 1921. DANA 1941 (10.6 mm), 17°58'N 64°4rW, 320 mw,
1
12 Dec. 1921. DANA 1202IV (12.5 mm), 100 mw, and DANA 1202
(1 1.3mm), 50 mw, both 9°40'N 79°56'W, 10 Jan. 1922.

Transforming larvae: MCZ 57922 (75.5 mm), 1°36'N 62°46'W, 1
530m, 19 Apr. 1960. MCZ 62401 (85.0 mm) MOC

20-19, 0, 39° 13.5'N
71° 17.6'W, 0-1027m, 15 June 1982.
Adults: MCZ 39968 (9, 130-190 mm), 07° lO'N 53°07'W, 360 m,
6 Nov. 1957. MCZ 40561 (22, 90-175 mm), 07°34'N 54°50'W,
360 m, 6 Nov. 1957. MCZ 40561 (3, 90-102.2 mm
c.s.), 07°34'N
54°50'W, 6 Nov. 1957. MCZ 41840 (85.0 mm), 18° 16'N 67° I7'W,
450 m, 6 Oct. 1959. UMML 15608 (120 mm, c.s.) no data.
P. fraser-brunneri
Larvae: MCZ 62402 (31.5 mm), RHB 2053, 16°32'N 19°35'W,

50-56 m, 13 Nov. 1970.
Adults: USNM 245646 (3, 84-138), 04°26'N 08°29'W, 200 m.,
31 Oct., 1963, BBC 843. MCZ 63153 (3, 97-105, one specimen
C.S.), same data as USNM 245646.
Comparative material
STOMIIFORMES
Gonostomatidae — Diplophos taenia: MCZ 52562, MCZ uncat. c.s.;
Gonostoma elongatum: MCZ 62404; Gonostoma sp.: MCZ
uncat. c.s.
Sternoptychidae
— Maurolicus muelleri: MCZ 62598, MCZ uncat.
c.s.
Photichthyidae
— Photichthys argenteus: MCZ 56953; Polymetme
corytheola: MCZ 56968, MCZ uncat. c.s.
AULOPIFORMES
Aulopidae
— Aulopus nanae: MCZ 40516; A. japonicus: MCZ
45169 c.s.
—
Chlorophthalmidae Chlorophthalmus agazzisi: MCZ 40539, MCZ
40509 c.s., MCZ 41444 c.s.;C bicornis: BMNH 1939.5.24:457;
C. brasiliensis: MCZ 51365, MCZ 40564 c.s.; C. chalybeius:
MCZ 62155, MCZ C nigripirmis: BMNH
40564 c.s.;
1887.12.7:207; Chlorophthalmus sp.: MCZ 62403 c.s., MCZ

62591, MCZ 62592, MCZ 62593, MCZ 62597.
—
Ipnopidae Ipnops murrayi: MCZ 41133; Bathypterois phenax:
MCZ 57624; B. quadrifilis: MCZ 35598; B. viridensis: MCZ
40567 c.s.; Bathysauropsis gracilis: BMNH 1887.12.7:209-210.
Notosudidae — Scopelosaurus argenteus: MCZ 62405 c.s., MCZ
62406 C.S.; S. harryi: MCZ 40512.
—
Scopelarchidae Scopelarchus analis: MCZ 62599 c.s.
—
Bathysauridae Bathysaurus agazzisi: MCZ 55305 c.s., MCZ 62409
c.s.; fi. mo///5.MCZ41140
Synodontidae
— Synodus synodus: MCZ 47378, MCZ 47490 c.s.; 5".
jaculum: MCZ 46972 c.s.; Saurida brasiliensis: MCZ 62408

C.S.; Sa. tumbil: MCZ 59273; Sa. undosquamis: MCZ 56111
c.s.
Harpadontidae
— Harpadon MCZ uncat. sp.: c.s.
—
Alepisauridae Alepisaurus brevirostris: MCZ 60345, MCZ 43153
c.s.
Evermanellidae — Evermanella MCZ uncat. sp.: c.s.
Paralepididae
— Paralepis elongata: MCZ 43140; P. speciosa: MCZ
60332 c.s.; Lestrolepis intermedia: MCZ 62407 c.s.; Sudis
atrox: MCZ 60336 c.s.
-(^
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r-jgg
Harvard MCZ Libra
3 2044 066 302 829
Date Due

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111

CAMBRIDGE, MASSACHUSETTS 02138 U.S.A.

Museum of Comparative Zoology

No. 465. The Structure and Relationships of the Dromasaurs

No. 466. Systematics of the Mexicana Species Group of the

No. 467. Three New Species of the Anolis Punctatus Complex

No. 468. A New Forest Skink from Ponape. By A. Ross Kiester.

No. 469. Catalog of the Primary Types of Bombyliidae (Diptera)

Comparative Zoology, with Designations of Lecto-

No. 470. Systematic Implications of Innervation Patterns in Tele-

No. 472. Fishes of the Suborder Labroidei (Pisces: Perciformes):

No. 473. The Interrelationships of Pelycosaurs. By Donald

No. 474. Ruthiromia Elcobriensis, A New Pelycosaur from El

No. 475. New or Problematic Anolis from Colombia. L Anolis

No. 476. Townsend's Unmapped North Atlantic Right Whales

No. 477. New or Problematic Anolis from Colombia. IL Anolis

No. 478. New or Problematic Anolis from Colombia. IIL Two

No. 479. Agonistic and Courtship Displays of Male Anolis Sagrei.

No. 480. Three New Lizards of the Genus ^^©/^(Scincidae) from

No. 481. A New Anolis of the Lionotus Group from Northwest-

No. 482. New or Problematic Anolis from Colombia. IV. Anolis

No. 483. Notes on Pristidactylus (Sqamata: Iguanidae). By

No. 484. Male Aggressive Behavior in a Pair of Sympatric Sibling

No. 485. The Anatomy and Relationships of Stereophallodon

No. 486. Thelodus "Macintoshi" Stetson 1928, The Largest

No. 487. The Identity of Larval Parasudis (Teleostei, Chloroph-

Museum of Comparative Zoology

Ayala, Stephen 475

Bone, Quentin 470

Brinkman, Donald 464, 465, 473, 474, 485

Brown, Walter C 480

Eberth, David E 473, 474, 485

Etheridge, Richard 483

Evenhuis, Neal L 469

Garstka, William R 466

Hartel, Karsten E 487

Harris, Dennis 475

Kaufman, Leslie S 472

Losos, Jonathon B 484

Liem, Karel F 472

MiYATA, Kenneth 481

Moore, Karen E 476

Parker, Fred 480

Scott, Michele P 479

Shevill, William E 476

Stiassny, Melanie L. J 487

Turner, Susan 486

Williams, Ernest E 467, 471, 475, 477, 478, 482, 483

Cambridge, Mass. 30 December 1981 Number 464

THE ORIGIN OF THE CROCODILOID TARSI

Abstract. The tarsus of the proterosuchian Chasmatosaurus represents the

triiophosaurids, prolacertids, and Protorosaums, and suggests

Erythosuchus neither contradicts nor supports a relationship between Erythrosuchus

Museum of Comparative Zoology, Harvard University, Cambridge, Massachsuetts

difference precludes derivation of dinosaurs from any known

THE CROCODILE-NORMAL TARSUS

astragalus, the calcaneum, and the second to fourth distal tarsals.

fib art surf

Figure 1. The right astragalus and calcaneum of Caiman sclerops. A) calcaneum