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UNIVERSITY OF

ILLINOIS LIBRARY
AT URBANA-CHAMPAIGN
NATURAL HIST. SURVEY
FIELD
Botany
NEW SERIES NO. 20
UBMW

PTERIDOPHYTA OF PERU
Parti
1. Ophioglossaceae-12. Cyatheaceae
Rolla M. Tryon
Robert G. Stolzc

January 31. 1989


Publication 1397

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY


nation diana

'

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.


FIELDIANA
Botany
NEW SERIES, NO. 20

PTERIDOPHYTA OF PERU
Parti
1. Ophioglossaceae-12. Cyatheaceae

Rolla M. Tryon
Gray Herbarium
Harvard University
22 Divinity Avenue
Cambridge, Massachusetts 02138

Robert G. Stolze
Collection Manager, Fern Herbarium
Department of Botany
Field Museum of Natural History
Roosevelt Road at Lake Shore Drive
Chicago, Illinois 60605-2496

Accepted February 19, 1988


Published January 31, 1989
Publication 1397

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY


1989 Field Museum of Natural History
Library of Congress Catalog Card Number: 88-80741
ISSN 00 15-0746
PRINTED IN THE UNITED STATES OF AMERICA
Table of Contents List of Illustrations

INTRODUCTION 1 1 .
Botrychium virginianum 7
ACKNOWLEDGMENTS 2 2. Ophioglossum: O. palmatum; O. crotal-
KEY TO FAMILIES OF PTERIDOPHYTA IN ophoroides; O. reticulatum 10
PERU 3 3. Marattia laevis 14
1 . OPHIOGLOSSACEAE 5 4. Danaea: D. moritziana; D. nodosa .... 16

Botrychium 6 5. Osmunda: O. regalis var. spectabilis;

Ophioglossum 8 O. cinnamomea 22
2. MARATTIACEAE 13 6. Anemia: A. pastinacaria; A. phylli-
Marattia 13 tidis 26
Danaea 15 7. Lygodium: L. venustum; L. volubile ... 31
3. OSMUNDACEAE 20 8. Schizaea elegans 35
Osmunda 21 9. Gleichenia bifida 40
4. SCHIZAEACEAE 23 10. Dicranopteris: D. pectinata; D. flex-
Anemia 24 uosa 48
Lygodium 30 11. Hymenophyllum: H. fucoides var. fu-
Schizaea 33 coides\ H. polyanthos; H. crispum 51
5. GLEICHENIACEAE 37 12. Trichomanes: T. radicans; T. pinnatum;
Gleichenia 37 T. hymenoides 77
Dicranopteris 46 13. Loxsomopsis pearcei 100
6. HYMENOPHYLLACEAE 49 14. Plagiogyria semicordata 102
Hymenophyllum 50 15. Culcita coniifolia 104
Trichomanes 76 16. Dicksonia sellowiana 106
7. LOXOMATACEAE 98 17. Lophosoria quadripinnata var. quadri-
Loxsomopsis 99 pinnata 108
8. PLAGIOGYRIACEAE 99 18. Metaxya rostrata 110
Plagiogyria 101 19. Sphaeropteris: S. elongata; S. quindiu-
9. DlCKSONIACEAE 101 ensis 113
Culcita 103 20. Alsophila engelii 117
Dicksonia 105 21. Nephelea cuspidata 119
10. LOPHOSORIACEAE 107 22. Trichipteris pubescens 121
Lophosoria 107 23. Cyathea caracasana var. boliviensis ... 134
11. METAXYACEAE 109 24. Cnemidaria speciosa 137
Metaxya 109
1 2. CYATHEACEAE Ill
Sphaeropteris 112
Alsophila 116
Nephelea 118
Trichipteris 1 20

Cyathea 129
Cnemidaria 1 36

MAP OF PERU 140


INDEX TO NAMES . .141

111
PTERIDOPHYTA OF PERU
Parti
1. Ophioglossaceae-12. Cyatheaceae

Introduction

The pteridophytes form a large and conspicuous their knowledge of particular genera, but several
element of the Peruvian flora, including 96 genera treatments will be contributed by specialists cur-
and about 1 ,000 species. Peru, which encompasses rently involved in monographic studies. The re-
one of the world's richest biotas, occupies a central sults will be published in five parts, divided as
position in the Andes, and accordingly a knowl- equally as practicable, beginning with the present
edge of its flora is basic to understanding the plant volume on the Ophioglossaceae through Cyathea-
life of the Andes as a whole. The Andes form a ceae.

largely tropical mountain chain with an essentially Studies are based primarily on the collections
north and south orientation, which is of special Museum, Harvard University Herbaria,
at Field

significance to its biogeography and the underlying and United States National Herbarium (Smith-
processes of speciation. sonian Institution), but specimens at many other
A portion of the Ferns of Peru was published United States and European institutions have been
by Rolla M. Tryon in 1964 (Contr. Gray Herb., examined. The extensive collections made under
encouragement and active sup-
vol. 194), with the the current Flora of Peru project, a joint under-
port of Theodore K. Just of Field Museum of Nat- taking of Field Museum and Missouri Botanical
ural History. A commitment was made at that Garden, are fully utilized. A close collaboration
time to complete the project after undertaking a has also been established with Universidad Na-
major work on the genera of pteridophytes in cional de Trujillo, Trujillo, Peru, and Museo de
America. The present work is a cooperative proj- Historia Natural "Javier Prado" de Universidad
ect of Harvard University Herbaria and Field Mu- Nacional Mayor de San Marcos, Lima, Peru, in-
seum of Natural History to present an account of cluding loans of their specimens. The type of each
all of the Pteridophyta of Peru. This is particularly name has been determined when possible, and an
important in relation to the Flora of Peru, initiated effort has been made to see the holotype, or at
at Field Museum over 50 years ago, and thus far least type photographs, or authentic material.

encompassing some 8,500 published pages. It is Original drawings illustrate the diagnostic features
the only comprehensive treatment of Pacific coast- of each genus and, where possible, some of the
al, Andean, and Amazonian plants, and is closer number of plates published
species. In addition, a
to completion than any flora of the larger South Botany, Ferns and fern allies of Gua-
in Fieldiana:
American countries. temala, have been used for species occurring in
The taxonomy is based not only on Peruvian Peru. Voucher specimens cited in the legends are
materials, but includes an assessment of the An- from Peru unless otherwise indicated.
dean species of a genus and, where appropriate, Prior to 1 944 the Department of Pasco was a
of all tropical American species. Most of the work part of Junin, and until recently Ucayali was a
will be accomplished by the authors, according to part of Loreto. An attempt has been made to ac-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.


count for these changes. This is not difficult when shipment of specimens from the two Peruvian her-
labels cite towns and provinces; however, on older baria. The original drawings were contributed by

specimens the labels often contain only sketchy Field Museum scientific illustrators, Zorica Da-
data,making it impossible to determine in which bich and Clara L. Richardson, and by volunteer
part of Loreto or Junin a plant was collected. The illustrators, Julia A. Liesse, Rosemarie Seitz, and
map of Peru at the end of the text shows the de- Lisa M. Thorns. To these fine artists we wish to
partments and indicates the sequence of the col- express our sincere appreciation, not only for their
lection citations. painstaking work, but also for their patience and
In the present work, Part I, the treatments of understanding. Special thanks are extended to sev-
the Ophioglossaceae through the Plagiogyriaceae eral persons who have been particularly helpful to
have been prepared by Robert G. Stolze, and those by providing special aid with the ex-
us, either
of the Dicksoniaceae through the Cyatheaceae, by amination of specimens at their institutions, or
Rolla M. Tryon. However, each author has re- through stimulating discussions or correspon-
viewed and edited the manuscript of the other, to dence concerning problems with various genera:
the extent that the treatments are a joint effort. Dr. David Lellinger (US), Dr. Alan R. Smith (UC),
New names published here are indicated by Dr. Henk van der Werff (MO), Dr. W. H. Wagner
boldface in the Index to Names. Collections have (MICH), and Dr. Paulo Windisch (Dept. Botanico,
been cited from each department of Peru from Rio Preto, Brazil). We also appreciate comments
which material has been seen, and further collec- on the manuscript by several reviewers.
tions are sometimes cited to include other her- We extend our thanks to the officers of the fol-
baria. In general, all collections seen are cited for lowing institutions for granting loans of their ma-
rare species, and a selection is cited for common terial or allowing us to examine specimens in their

species. Type collections, mentioned in the syn- herbaria: Botanischer Garten und Botanisches
onymy, are not repeated in the specimen citations, Museum, Berlin-Dahlem, Berlin (B); British Mu-
although they are included in the Peruvian range seum (Natural History), London (BM); Jardin Bo-
and ecology. tanique National de Belgique, Meise (BR); Royal
The nomenclature of the genera and species is Botanic Garden, Edinburgh (E); Field Museum of
not intended to be complete. Synonyms are listed Natural History, Chicago (F); Harvard University,
when they are considered useful and when the type Cambridge, Mass. most Gray Herbarium (GH),
of the name of a species or infraspecific taxon is some Arnold Arboretum (A); Institut fur Allge-
from Peru. meine Botanik und Botanischer Garten der Uni-
Appended to some of the generic treatments are versitat, Hamburg (HBG); Herbarium Truxil-
portions of text labeled Comments. Herein are in- lense,Universidad Nacional de Trujillo, Trujillo,
cluded species to be expected in Peru, names based Peru (HUT); Royal Botanic Gardens, Kew, En-
on Peruvian material but of uncertain application, gland (K); Rijksherbarium, Leiden, The Nether-
excluded species (erroneous reports that may have lands (L); Botanische Staatssammlung, Munich
special significance), and cultivated species that (M); University of Michigan, Ann Arbor (MICH);
are possibly adventive. Missouri Botanical Garden, St. Louis (MO); New
Abbreviations of periodical publications gen- York Botanical Garden, New York (NY); Museum
system of Botanico-Periodicum-
erally follow the National d'Histoire Naturelle, Paris (P); Univer-
Huntianum (1968), and abbreviations of authors' sityof California, Berkeley (UC); United States
names and of books generally follow TL-2 Taxo- National Herbarium, Smithsonian Institution,
nomic Literature by Stafleu and Cowan (1976 et Washington, D.C. (US); Museo de Historia Nat-
seq.). ural "Javier Prado" de Universidad Nacional

Mayor de San Marcos, Lima, Peru (USM); and


Naturhistorisches Museum, Vienna (W).
This project has been supported in part by grant
Acknowledgments #BSR-85 16358 from the National Science Foun-
dation, Systematic Biology Program. The work
The authors are indebted to Dr. Michael Dillon would not have been possible without this assis-
(F), Dr. Abundio Sagastegui (HUT), and Blanca tance; however, any opinions and conclusions ex-
Leon (USM) for their invaluable assistance in pre- pressed are those of the authors and do not nec-
paring loans and arranging for the packing and essarily reflect the views of the Foundation.

FIELDIANA: BOTANY
Key to Families of PTERIDOPHYTA in Peru

The following key is restricted to the families that occur in South America and to the genera that are
known from Peru or are likely to occur there. Several families are keyed out more than once in order
to simplify the headings and to provide for more accurate identification.

a. Two or more sporangia joined in a synangium b


b. Synangium single in the axil of a bifid, bractlike leaf; aerial stem slender, green, dichotomously
forked 24. Psilotaceae
b. Two elongate synangia at the apex of a spike, or many synangia on the abaxial surface of a leaf;
stem subterranean c
c. Two elongate synangia at the apex of a spike; stem small 1 .
Ophioglossaceae
c. Many synangia on the abaxial surface of a leaf; stem stout to massive 2. Marattiaceae

a. Sporangia separate d
d. Sporangia borne on the inner side of a peltate sporangiophore in an apical strobilus; leaves much

reduced, forming a sheath at the apex of each elongate, ridged interode Goint) of the stem
25. Equisetaceae
d. Sporangia single in or near the axil of a leaf, or on 1 lobe of a 2-lobed leaf, or several to many

sporangia borne on a leaf: at the margin, on the abaxial surface, on a specialized portion, or on
a specialized leaf e
e. A singlesporangium borne in or near the axil of a leaf f
f. Plants homosporous; leaves lacking a ligule 26. Lycopodiaceae
f. Plants heterosporous, with megasporangia and microsporangia; leaves with a ligule g
g. Leaves less than 1 cm long, borne along an elongate stem, fertile leaves in an apical
strobilus 27. Selaginellaceae
g. Leaves 2 cm long or usually longer, clustered at the apex of a compact to slightly elongate
stem; all leaves usually fertile 28. Isoetaceae
e. Several to many sporangia borne on a leaf, or a single sporangium borne on 1 lobe of a 2-
lobed leaf h
h. Plants heterosporous; the leaf bearing megasporangia and/or microsporangia, these enclosed
in small, specialized structures i

i. Plant with stem rooted in wet soil or underwater; leaves filiform, or with 4 leaflets at
the apex of the petiole 22. Marsileaceae
i. Plant floating on water; the floating leaves entire, oblong to suborbicular, or unequally
2-lobed with 1 lobe submerged 23. Salviniaceae
h. Plants homosporous, the isomorphic sporangia exposed at the margin or on the abaxial
surface of a leaf, or on a specialized portion of a leaf, or on a specialized leaf, sometimes
enclosed prior to maturity within an indusium j

j. Sporangia sessile or subsessile, or with a stalk of 4 or more rows of cells, annulus absent,
or if present then apical, lateral, or oblique, not interrupted by the stalk k
k. Annulus lacking, or sporangia with a poorly differentiated annulus 1

1. Sporangia lacking an annulus, borne on a specialized fertile branch of a leaf; spores


lacking chlorophyll 1 Ophioglossaceae .

1. Sporangia with a poorly differentiated lateral annulus, borne on fertile pinnae;


spores with chlorophyll (green) 3. Osmundaceae

k. Sporangia with a well-differentiated apical to oblique annulus m


m. Sporangia on the abaxial surface of a fertile portion of a leaf, distantly attached
in a cluster or single on a vein, or in wholly fertile panicles 4. Schizaeaceae . .

m. Sporangia contiguous on the receptacle of marginal or abaxial sori n


n. Sporangia in marginal sori o
o. Leaf very thin, 1-few cells thick, translucent, lacking stomata; stem pro-
tostelic 6. Hymenophyllaceae

o. Leaf thickened, with stomata; stem siphonostelic or dictyostelic p

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.


p. Stem long-creeping, with scattered, short, stiff trichomes; receptacle
elongate 7. Loxomataceae

p. Stem massive, short-creeping to arborescent, with a dense mass of very


long, soft trichomes; receptacle short or globose ... 9. Dicksoniaceae
n. Sporangia in abaxial sori q
q. Stem and leaves lacking evident indument, a mucilaginous secretion (flaky
when dry) sometimes present; leaves strongly dimorphic
8. Plagiogyriaceae
q. Stem and usually the leaves with evident trichomes and/or scales; leaves
monomorphic to somewhat dimorphic r
r. Stem slender, long-creeping, subterranean, freely branched; leaf usually
partly pseudodichotomously branched with axillary arrested or dormant
buds 5. Gleicheniaceae

r. Stem stout to massive, more or less epigeous or arborescent, sparingly


if at all branched; leaves wholly pinnately branched s

s. Stem and petiole bearing scales, trichomes may be present or absent


12. Cyatheaceae
s. Stem and petiole bearing only trichomes t

t. Lamina 2-pinnate-pinnatifid to 3-pinnate-pinnatisect


10. Lophosoriaceae
t. Lamina -pinnate
1 11. Metaxyaceae

j. Sporangia with a 1- to 3 -rowed stalk; the annulus vertical or nearly so, interrupted by
the stalk u
u. Petiole articulate at or very near the stem 21. Polypodiaceae
u. Petiole continuous with the stem, or articulate well above the base of the petiole v . . .

v. Stem scales clathrate w


w. Spores monolete x
x. Lamina indusium lacking
entire; 14. Vittariaceae
x. Lamina pinnatifid or more complex, or entire and the sori indusiate y . .

y. Spores with chlorophyll (green), or lamina more or less dichotomous


(furcate) 21. Polypodiaceae
y. Spores lacking chlorophyll (not green) and lamina entire or pinnately
branched z
z. Sori abaxial; indusia, when present, circular to reniform, with a some-
times narrow sinus 17. Dryopteridaceae
z. Sori abaxial, indusiate, the indusia elongate, or sori nearly marginal
18. Aspleniaceae
w. Spores trilete aa
aa. Sori very long, near and parallel to the margin, or following the anasto-
mosing veins, or in scattered groups 14. Vittariaceae
aa. Sori roundish or united in a short line close to the costa
21. Polypodiaceae
v. Stem scales not clathrate, or stem with trichomes bb
bb. Spores trilete, or monolete and the sori marginal cc
cc. Indusia absent dd
dd. Stem with trichomes only 13. Pteridaceae
dd. Stem with scales ee
ee. Sori elongate along the veins, or sporangia acrostichoid
13. Pteridaceae
ee. Sori roundish or united in a short line close to the costa
21. Polypodiaceae
cc. Indusia present ff

ff. Stem with trichomes, or with scales and an abaxial indusium present
15. Dennstaedtiaceae
.

FIELDIANA: BOTANY
ff. Stem with scales and an abaxial indusium absent gg
gg. Spores trilete 13. Pteridaceae
gg. Spores monolete 15. Dennstaedtiaceae
bb. Spores monolete and the sori or sporangia abaxial hh
hh. Sori elongate, adjacent to and parallel to the segment axis
20. Blechnaceae
hh. Sori roundish, or elongate, and most or all of them neither adjacent to
nor parallel to the segment axis, or sporangia acrostichoid ii

ii. Pinna stalks, if present, continuous with the rachis jj

jj. Spores with chlorophyll (green); lamina or main veins more or less
dichotomous (furcate) 21. Polypodiaceae

jj .
Spores lacking chlorophyll (not green) and lamina pinnately veined
or branched kk
kk. Petiole, at least basally, with 2 vascular bundles 11

11. Lamina with unicellular, acicular, or variously branched


trichomes, or if glabrate then indusia absent
16. Thelypteridaceae
11. Lamina lacking trichomes, or with multicellular ones, or
with minute, unicellular, blunt trichomes; indusia present
17. Dryopteridaceae
kk. Petiole, at least basally, with 3 or more vascular bundles . .

17. Dryopteridaceae
ii. Pinnae articulate to the rachis mm
mm. Sporangia acrostichoid, or in indusiate sori and the lamina 2-
pinnate, or 1 -pinnate and the pinnae with a large, basal, bas-
iscopic auricle 17. Dryopteridaceae
mm. Sporangia in indusiate sori, lamina 1 -pinnate, the pinnae cor-
date, or the basal, basiscopic side less developed
. 19. Davalliaceae

Family 1: OPHIOGLOSSACEAE and contains three genera and 50 to 60 species.


ilies

Botrychium and Ophioglossum occur in Peru and


Ophioglossaceae C. Agardh, Aphor. bot. 113.1 822. are virtually cosmopolitan, with representatives in
TYPE: Ophioglossum L. subarctic as well as tropical regions. The Ophiog-
lossaceae are distinctive in the fleshy, mycorrhizal
Stem erect or prostrate, fleshy, lacking indu- roots lacking root hairs, the erect or folded (rather
ment, or pubescent-scaly at apex. Leaves erect or than circinate) vernation, and the massive spo-
folded, not or scarcely circinate in vernation, en- rangia with an extremely high spore capacity. Some
pinnate or subpalmate or dichotomous,
tire to of the species tend to have very wide, and often
commonly partially dimorphic, the sterile leaf with disjunct, distributions.
an expanded lamina, the fertile one with sporangia
borne on a special branch (or branches) arising
from the base of or below the sterile lamina. Veins References
free or anastomosing. Sporangia sessile or subses-
sile, separate or laterally joined in a synangium, CLAUSEN, R. T. 1938. A monograph of the
lacking an annulus. Spores trilete, lacking chlo- Ophioglossaceae. Mem. Torrey Bot. Club, 19(2):
rophyll, from 1,500-15,000 in each sporangium. 1-177.
UNDERWOOD, L. M., AND R. C. BENEDICT. 1909.
This is the most primitive of all living fern fam- Ophioglossaceae, in N. Amer. fl., 16: 3-13.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.


Key to Genera of Ophioglossaceae

a. Sterile lamina ternately decompound; veins free; sporangia borne in a panicle, separate from each
other and not immersed in the fertile segment I. Botrychium

a. Sterile lamina entire or palmately or digitately lobed; veins anastomosing; sporangia joined laterally
in a synangial spike II. Ophioglossum

I. Botrychium America. Within the four subgenera of Botrychium


only four species of Sceptridium have been found
Botrychium Sw., J. Hot. (Schrader) 1800(2): 8, 110. in the Neotropics, and but a single species of sub-
1802.TYPE: Botrychium lunaria (L.) Sw. genus Osmundopteris (B. virginianum). The pau-
(Osmunda lunaria L.). Figure 1. city of species in the Neotropics may be partly a
natural phenomenon, but the problem may be also
Plants terrestrial, very rarely epiphytic. Stem attributed to poor and sparse collections, as ferns
erect, short, lackingindument. Leaves solitary or of this genus often go undetected. Increased col-
few, 2-70 cm long, glabrous to pubescent, bud of lecting efforts in North America have produced
the next leaf wholly or partly enclosed by the ex- more species, and similar results may be attained
panded stipular base of the current leaf. Sterile through greater efforts in the tropics. Collectors
lamina sessile to long-stalked, pinnatifid to de- should be aware that when a single plant is found
compound. Veins free. Fertile branch paniculate, in the field there are likely to be many more of
racemose or spicate, arising at or below the base that and other species nearby.
of the lamina. Sporangia protruding from the sur-
face of the fertile branch, not immersed in the References
tissue.

BUTTERS, F. K. 1917.
Botrychium virginianum
The genus contains about 25 species, with dis- and its American Rhodora, 19: 207.
varieties.
tribution in temperate to tropical regions in both MILDE, J. 1869. Botrychiorum monographia.
hemispheres, but is poorly represented in tropical Verh. Zool. Bot. Ges. Wien, 19: 55-190.

Key to Species of Botrychium

a. Fertile leaf with the sterile lamina long-stalked; leaf bud completely enclosed by the sheath of the
current leaf 1 . B. schafTneri
a. Fertile leaf with the sterile lamina sessile or nearly so; leaf bud partly exposed within the sheath of
the current leaf 2. B. virginianum

1. Botrychium schafTneri Underw., Bull. Torrey shaped sinuses, the margins entire to crenate, very
Bot. Club 30: 51. 1903. TYPE: Mexico, San rarely serrate.
Luis Potosi, Schaffner 9 (holotype, NY; iso-
type, K). Terrestrial, in high scrub forests and open mead-
ows, 2350-3600 m, Amazonas, Cuzco, Madre de
Dios(?).
Mexico; Honduras south to Argentina.
Fertile plants 12-30 cm tall (sometimes to 50 This is often difficult to distinguish from B. un-
cm outside Peru), the common stalk to 4 cm long, derwoodianum (Guatemala to Venezuela), which
the leaf bud completely enclosed by the sheath of is usually a much larger fern with stalks of sterile
the current leaf. Sterile lamina of fertile leaf long- laminae up to 28 cm long. However, Mexican
stalked (3-8 cm), fleshy, ternate, to 3-pinnate-pin- plants of B. schqffneri seem to rival the latter in
natifid,3-10 cm long, 6-10 cm broad. Ultimate size, and the more reliable diagnostic features ap-
segments 0.3-0.7 cm long, mostly obtuse to sub- pear to occur in the ultimate segments. Those of
acute, separated by usually narrow to broadly U- B. underwoodianum are 0.7-1 .4 cm long with mar-

FIELDIANA: BOTANY
FIG.1. Botrychium virginianum: a, lamina with fertile branch; b, stem and part of petiole; c, part of fertile pinna,
(a from Seller 865, El Salvador, GH; b-c from Lyonnet 896, Mexico, GH.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.


gins typically serrate, whereas in B. schaffneri they geography. Proctor (Ferns of Jamaica, 1985) has
are usually less than 0.7 cm long and entire to observed that "... the almost complete intergra-
crenate (only rarely serrate). The taxonomy of the dation of the alleged distinguishing characters ren-
species complex requires detailed analysis. ders even varietal recognition difficult to uphold."
We concur with this judgment.
Amazonas: Chachapoyas, along Rio Ventilla, 1-2 km
W of Molinopampa. Wurdack 1528 (us). Middle eastern Cajamarca: Prov. Celendin, canyon of Rio Maranon
Calla Calla slopes, km 41 1-416 of Leimebamba-Balsas above Balsas, Hutchison & Wright 5358 (uc). Amazonas:
road, Wurdack 1319 (us). Cuzco: Urubamba, Machu Prov. Chachapoyas, Cerros Calla Calla, 5 km above Lei-
Picchu, slopes of Altillero, Peyton & Peyton 504 (MO). mebamba, Hutchison & Wright 4885 (uc). Huanuco:
Dept. Unknown: "Pinasniocj, Panticalla Pass," Cook & Mito, in canyon, Bryan 387 (F). Muna, in dry woods,
Gilbert 1845 (us) (this may be on Cerro de Pantiacolla, Bryan 537 (F). Pasco: Prov. Oxapampa, 5 km SE of
Dept. Madre de Dios, near the Cuzco border). Oxapampa, D. Smith 2897 (MO). Junin: Huacapistana,
dense forest, Killip & Smith 24308 (us). Cuzco: Prov.
Calca, Vilcabamba, Vargas 3907 (us).

2. Botrychium virginianum (L.) Sw., J. Bot.


(Schrader) 1800(2): 111. 1802. Figure 1.
II. Ophioglossum
Osmunda virginiana L., Sp. pi. 1064. 1753. TYPE:
"America," Kalm (LINN 1244.3). Ophioglossum L., Sp. pi. 1062. 1753. TYPE:
Osmunda cicutaria Savigny in Lam., Encycl. 4: 650. Ophioglossum vulgatum L. Figure 2.
1797. TYPE: based on plants collected by Plu-
mier in Hispaniola, Traite foug. Amer. 136, t.
159. 1705. Cheiroglossa Presl, Suppl. tent, pterid. 56. 1846. TYPE:
Cheiroglossa palmata (L.) Presl
= Ophioglossum
Botrychium cicutarium (Savigny) Sw., Syn. fil. 171.
1806. palmatum L.

Botrychium virginianum var. mexicanum Grev. &


Hooker, Bot. Misc. 3: 223. 1833 (as B. virginicum Plants terrestrial or epiphytic. Stem erect, glo-
beta mexicanum). SYNTYPES: Mexico, Vera- bose or short-prostrate, small, lacking indument,
cruz, Chamisso s.n.; "Rigla," Veitch (both E? or
or filiform-scaly at apex. Leaves solitary or several,
K?).
1-75 cm long (or to more than 1.5 m in the Old
Fertile plants ca. 25-70 cm the common World O. pendulum), glabrous, the eroded stipular
tall,
base of the expanded leaf not enclosing the bud of
stalk 1 9-40 cm long, the leaf bud partly exposed
within the sheath of the current
the next leaf. Sterile lamina sessile or short-stalked,
leaf. Sterile lamina
entire,or palmately or digitately lobed. Veins
of fertile leaf sessile or short-stalked (to 5 mm),
membranous, subternate, to 3-pinnate-pinnatifid, anastomosing, often with free, included veinlets.
Fertile branch or branches borne on or below the
to 20 cm long and 25 cm broad. Ultimate segments
base of the sterile lamina. Sporangia joined lat-
0.5-0.9 cm by narrow, acute sinuses,
long, joined
to obovate, truncate, rounded or subacute, erally in a synangial spike.
elliptic
their apices variously dentate or lacerate.
Ophioglossum is a genus of 25-30 species scat-
tered in temperate to tropic regions around the
Forests and wooded canyons, 1800-2930 m,
world. Plants are often small and inconspicuous
Cajamarca, Amazonas, Huanuco, Pasco, Junin,
Cuzco.
and are commonly concealed in grassy turf or her-
baceous growth; consequently, current collection
Essentially cosmopolitan. In the Neotropics from
records probably do not represent true distribution
Mexico south to Bolivia and Brazil, and in the
Greater Antilles. patterns. Although six species are recognized here
from Peru, several others might be expected in the
Botrychium virginianum has been treated by
future.
earlier authors as several species or asone species
with several varieties or subspecies. Variants have
been distinguished primarily by relative length of
the fertile branch or degree of dissection of the Reference
sterile lamina. However, closer examination of

more recent and numerous collections throughout WAGNER, W. H., ET AL. 1984. Ophioglossum el-
the range of the species indicates that these are lipticum in Louisiana and the taxonomy of O.
variable characters and not wholly correlated with nudicaule. Castanea, 49: 99-1 10.

FIELDIANA: BOTANY
Key to Species of Ophioglossum

a. Sterile lamina deeply and irregularly palmately lobed; fertile spikes several; plants epiphytic
1 . O. palmatum
a. Sterile lamina entire; fertile spike solitary; plants terrestrial b
b. Stem globose 5. O. crotalophoroides

b. Stem cylindrical, sometimes swollen, but never globose c


c. Sterile lamina stalked, the common stalk and stalks of fertile and sterile segments hypogeous

and scarious 4. O. scariosum

c. Sterile lamina sessile or nearly so, the common stalk and stalks of sterile and fertile segments

mostly epigeous, not or scarcely scarious d


d. Primary areoles each enclosing several smaller secondary areoles and/or some free veinlets;
base of sterile lamina cordate, truncate or abruptly cuneate 2. O. reticulatum

d. Primary areoles not enclosing secondary ones or free veinlets; base of sterile lamina narrowly
cuneate to attenuate e
e. Sterile lamina elliptic or oblong, 1.5-3 times as long as broad, cuneate at base; plants of

higher elevations (over 2500 m) 3. O. nudicaule var. tenerum

e. Sterilelamina linear-lanceolate to narrow-oblanceolate, (4-)4.5-8 times as long as broad,


long-attenuate at base; plants of lower elevations (under 700 m)
6. O. lusitanicum ssp. coriaceum

1 .
Ophioglossum palmatum Sp. pi. 1063. 1 753.
1 ... in deep ravine, Allard21580 (us). Pasco: Prov. Oxapam-

TYPE: based on illustration of plant from pa, 1800 m, van der \Verff8352 (MO). Cuzco: Prov. La
Convention, Tunquimayo, Idma, Vargas 10681 (MICH,
Haiti, Plumier, Traite foug. Amer., /. 163.
MO).
1705. Figure 2a.

Cheiroglossa palmata (L.) Presl, Suppl. tent, pterid.


57. 1846.
2. Ophioglossum reticulatum L., Sp. pi. 1063.
1753. LECTOTYPE (designated by Proctor, Flora
Plants epiphytic. Stem stout, elongate, abun-
Lesser Antilles 2: 43. 1977): Plumier, Traite foug.
dantly provided at apex with tawny to orange scales on specimen from Haiti. Fig-
Amer., /. 164, based
which are hairlike for most of their length. Leaf ure 2c-d.
(15-)20-75 cm
long, with common stalk (5-) 10-
40 cm long. Sterile lamina deeply and irregularly
Ophioglossum petiolatum Hooker, Exot. fl. 1: 56. 1823.
palmately lobed, obdeltoid, narrowly cuneate at TYPE: based on plants from West Indies sent to
base, lacking a distinct midvein or pale median Liverpool Botanic Garden and cultivated there
band of tissue. Veins distinct, theprimary areoles (not located).
Ophioglossum peruvianum Presl, Suppl. tent, pterid.
large, often enclosing smaller secondary areoles
52. 1845. TYPE: Peru, Poeppig(noi located).
and occasionally a few free veinlets. Fertile seg-
ments (l-)3-l 2, borne on or below the base of the
Plants terrestrial. Stem cylindrical, lacking in-
lamina, the fertile spikes 0.5-6 cm long, on stalks
dument. Leaf cm long, common stalk
(5-) 10-40
0.2-2 cm long.
mostly epigeous, 3-20 cm long. Sterile lamina
membranaceous, 2-12 cm long, 1.5-5 cm broad,
Suberect or pendent from trunks or branches of
sessileor subsessile, ovate to nearly circular, apex
trees, in cloud forests, elfin forests, and shaded
obtuse to acute, base cordate to truncate or (oc-
ravines, 1000-2450 m, Amazonas, Huanuco, Pas-
casionally) abruptly cuneate. Veins distinct to in-
co, Cuzco.
distinct, primary areoles enclosing a few smaller
Southern Florida; West Indies; southern Mexico
secondary areoles which are scarcely differentiated
to Peru and Brazil; scattered in Old World tropics.
from the primary ones, free included veinlets sev-
eral to numerous. Fertile segment solitary, borne
Amazonas: Prov. Bagua, E of La Peca, Barbour 2512,
at baseof sterile lamina, on mature leaves some-
2860, 2940 (MO). Prov. Chachapoyas, along Rio Ventilla
1-2 km W
of Molinopampa, Wurdack 1478 (F, GH, NY, what to greatly exceeding the lamina, fertile spike
uc, us). Huanuco: East of Tingo Maria (as San Martin) 1.4-4 cm long, on a stalk 3.5-14.5 cm long.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.


3 mm

5mm

3cm

FIG. 2. Ophioglossum palmatum: a, habit. Ophioglossum crotalophoroides: b, habit. Ophioglossum reticulatum:


c, venation of sterile leaf; d, fertile spike, (a from Allard 21580, us; b from Vargas 3978, us; c-d from Tryon & Tryon
5419, F.)

10 FIELDIANA: BOTANY
From high, rocky meadows to the open forests and sedges, 2800-4000 m, Cajamarca, Lima, Cuz-
of middle elevations, and the grassy soil of coastal co.
lomas, 400-3800 m, Amazonas south to Cuzco. Colombia; Peru; Brazil.
Widely distributed in tropical America; Old In spite of Clausen's work (1938), considerable
World. study is still needed to understand the variation
Ophioglossum reticulatum is variable in the within O. nudicaule. The Peruvian plants here
shape and base of the sterile lamina and in the identified cannot be clearly placed within any of

frequency of secondary areoles and free, included the varieties delineated by Clausen; but, aside from
veinlets. These characters intergrade freely with the fact that their fertile segments do not rise too
those of the often recognized O. petiolatum, so that conspicuously beyond the sterile lamina, they seem
it is impractical to separate the two as distinct taxa. to fit best under var. tenerum. Diagnostic features
Thus circumscribed, O. reticulatum is perhaps the of O. nudicaule are so few and variable that it is

most ubiquitous of any species in the genus. often difficult to discern which are innate differ-
ences and which are merely induced by rigorous
Amazonas: Prov. Bongara, Shilla. Young & Eisenberg habitat or attributed to degree of maturity of the
460 (MO). La Libertad: Prov. Pataz, Pumatambo, Puerta individual plant. that this and other
It is likely
del Monte. Lope: & Sagdstegui 3438 (GH). San Martin:
Monte Campana, near Tarapoto, Spruce 4709 (p, us).
varieties acceptedby Clausen are not supportable,
as evidenced by the studies of Wagner et al. ( 1 984),
Lima: Lomas de Quilmana, Coronado 27 (us). Prov.
Chancay, Lomas de Lachay, Coronado & Velarde 15 (uc, which suggest that they "all should be subsumed
us); Tryon & Tryon 5419 (GH). Pasco: Prov. Oxapampa, under the single binomial, O. nudicaule."
Chontabamba, D. Smith & Brack 3056 (F, MO). Ayac-
ucho: Aina, between Huanta and Rio Apurimac, Killip
& Smith 22809 (GH, NY, us). Apurimac: Prov. Abancay, Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui
et al. 6412 (GH, HUT). Lima: Prov. Huarochiri, Dist. de
Cachora, Huillcayoc, Vargas 9096 (MICH, uc). Cuzco:
Mariatana, Cueva Mortero, Cerrateet al. 4812 (GH, USM).
Prov. Urubamba, Machu Picchu, Leon 452 (GH, USM);
Cuzco: Prov. Acomayo, Quenco Grande, near Acomayo,
Tryon & Tryon 5400 (GH, us), 5419 (GH). Madre de Dios:
Prov. Manu, Parque Nacional Manu, Foster et al. 11486
Vargas 9752 (F, GH, K, ucp.p., us). Prov. Espinar, Yauri,
Vargas 5619 (MICH). Prov. Urubamba, Vargas 14122
(F).
(GH).

3. Ophioglossum nudicaule L. f. var. tenerum 4. Ophioglossum scariosum Clausen, Mem. Tor-


(Mett.) Clausen, Mem. Torrey Bot. Club 19(2): rey Bot. Club 19(2): 153. 1938. TYPE: Peru,
151. 1938. Junin, vicinity of Oroya, Kalenborn 125 (ho-
lotype, us!; isotypes, GH!, MO!, NY).
Ophioglossum ypanemense Mart., Icon. pi. crypt. 39,
30, /. 73. 1 834. TYPE: based on specimens from
1
Plants terrestrial. Stem broadly elongate, 4-6
"Ypanema," Brazil.
Ophioglossum tenerum Mett. in Prantl, Ber. Deutsch.
mm thick, lacking indument. Leaf 3-6 cm long,
Bot. Ges. 1: 352. 1883. TYPE: United States,
with common stalk and bases of fertile and sterile
Georgia, collector undesignated (holotype, B). stalks hypogeous and scarious, common stalk 0.3-
0.8 cm long. Sterile lamina 1-1.5 cm broad, ob-
Plants terrestrial. Stem cylindrical to somewhat long-ovate to suborbicular, apex apiculate, at base
swollen, lacking indument. Leaf 2.5-7 cm long, truncate, reduced abruptly to an attenuate stalk
common stalk mostly epigeous, 0-4 cm long. Ster- 0.8-1.2 cm long. Veins indistinct to nearly ob-
ilelamina chartaceous to somewhat carnose, 0.8- scure, primary areoles usually enclosing some sec-
2.5 cm long, 0.3-1 .0 cm broad, 1.5-3 times as long ondary ones as well as free veinlets. Fertile seg-
as broad, sessile, elliptical to subovate, obtuse or ment solitary, borne at the base of the stalk of the
subacute, base cuneate. Veins indistinct to ob- sterile lamina, on mature leaves somewhat to

scure, areoles small, not enclosing secondary ar- greatly exceeding the lamina, fertile spike 0.6-1.2
eoles or (except rarely) free veinlets. Fertile seg- cm long, on a stalk 1.3-4 cm long.
ment solitary, borne at base of sterile lamina, on
mature leaves somewhat exceeding the lamina, In open, grassy areas, 2840-3850 m, Junin, Cuz-
fertile spike 0.5-1 .2 cm long, on a stalk 0.8-2. 5(-4) co, Puno.
cm long. Peru and Bolivia.
Leaves of this rare fern grow with proximal por-
In high rocky meadows and slopes, with grasses tions of leaves embedded in the soil. This is evi-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 11


dent even in dried material, as the scarious com- Ophioglossum coriaceum A. Cunn., Companion Bot.
mon stalk and proximal portions of the stalks of Mag. 2: 361. 1836. TYPE: based on specimens
from Matauri, New Zealand (not located).
fertile and sterile segments contrast sharply with
the dark color of the rest of the leaf.
Plants terrestrial. Stem cylindrical to somewhat
swollen, lacking indument. Leaf 4-8(-9) cm long,
Cuzco: Prov. Urubamba, Vargas 9165 (MICH). Puno:
Lake Titicaca, Bishop 1994 (us).
common stalk mostly epigeous, 0.5-2.5 cm long.
Sterile lamina chartaceous to somewhat carnose,
1.54 cm long, 0.3-0.6 cm broad (ca. 4.5-8 times
as long as broad), sessile, linear-lanceolate to nar-
5. Ophioglossum crotalophoroides Walter, Flora
Caroliniana 256. 1788. TYPE: Carolina, row-oblanceolate, apex acute or subacute, base
Walter (holotype, BM?). Figure 2b. narrowly cuneate to (typically) long-attenuate.
Veins indistinct to obscure, areoles commonly long
Plants terrestrial. Stem globose, 4-12 mm in and narrow, not enclosing secondary areoles or
free veinlets. Fertile segment solitary, borne at base
diameter, lacking indument. Leaf 3-15 cm long,
with common stalk 0.5-4 cm long. Sterile lamina of sterile lamina, on mature leaves somewhat to
ovate to deltoid, acute or rarely obtuse, cordate or conspicuously exceeding the lamina, fertile spike
truncate and abruptly attenuate at base, often lon- 0.4-1.7 cm long, on a stalk 1-7 cm long.

gitudinally folded. Veins distinct or indistinct, pri-


In soil pockets or on thin soil over rocks, in Peru
mary areoles not or rarely enclosing secondary ones
or free veinlets. Fertile segment solitary, borne at possibly confined to the coastal lomas in the vi-
the base of the sterile lamina; in mature leaves the cinity of Lima, 400-650 m.
fertile segments somewhat to greatly exceeding the Peru; Bolivia; Chile; New Caledonia; Australia;
lamina, fertile spike 0.5-1.5 cm long, on a stalk New Zealand.
to 10 cm long. In Peru, this is typically a very delicate plant,
with long, narrow, sterile leaves and a fragile, fil-

On moist, open, grassy slopes, 600-3900 m, iform, fertile stalk, and has thus far been found

Huanuco, Lima, Junin, Apurimac, Cuzco. only at low elevations. However, we have exam-
Southern United States; Mexico to Honduras; ined some Bolivian plants of this subspecies, from
Colombia and Venezuela south to Argentina and elevations to 3000 m, which are more variable:
Chile. sometimes more robust, with stouter fertile stalks,
and with sterile laminae broader and less attenuate
Among the diminutive species of Ophioglos-
to base and apex. Subspecies lusitanicum, from
sum, a few plants are found with stems which are
somewhat swollen. However, only in O. crotalo- Europe, Asia, and Africa, has obtuse sterile lam-
is the stem so conspicuously globose. inae with larger and fewer areoles. Subspecies cal-
phoroides
With this should probably be included O. opacum ifornicum (Prantl) Clausen (California & Mexico)
has laminae larger and broader, with less attenuate
Carmichael, from Tristan da Cunha. It is said to
differ from the former merely in the somewhat bases than in ssp. coriaceum and, according to
shorter fertile segment and slightly smaller globose Clausen (1938), perhaps does not merit distinc-
stem. tion.As with the O. nudicaule complex (q.v.), far
more study is needed to clarify the taxonomy, at
Huanuco: Prov. Huanuco, Huanuco-Tingo Maria road, both inter- and intraspecific levels.
from Bandera Blanca to Carpish Tunnel, Luteyn & Lu-
teyn 5464 (NY). Lima: Prov. Lima, Lomas de Lurin, Lima: Prov. Chancay, Lomas de Lachay, Coronado &
Ferreyra 9566 p.p. (USM). Junin: Huaytapata. near Ha- Velarde 16 (GH, uc, us); Tryon & Tryon 5414 (GH p.p.,
cienda Conocancha, Tiller 14 (USM). Apurimac: Prov. us).Lomas de Quilmana, Coronado 27 (uc, us p.p.).
Abancay, Cachora, Vargas 9054 (MICH p.p., MO). Cuzco: Lomas de Lurin. Ferreyra 9566 (GH, USM p.p.). Lomas
Prov. Calca, Vilcabamba, Vargas 3978 (us). Prov. Can- de Atocongo, Grant 7509 (GH).
as, slopes of San Andres de Checca, Vargas 11010 (F,

uc).

Comments
6. Ophioglossum lusitanicum L. ssp. coriaceum (A.
Ophioglossum ellipticum Hooker & Grev., Icon,
Cunn.) Clausen, Mem. Torrey Bot. Club 19(2):
fil. 1, t. 40A. 1831. TYPE: illustration, based
161. 1938.

12 FIELDIANA: BOTANY
on specimens collected in Demerara, "British to pinnately compound or rarely simple or pal-
Guiana," Parker. mate, monomorphic to somewhat dimorphic. Pet-
iole with an expanded, stipular base, there swollen
This perhapsmay be expected in Peru, as its and sometimes subarticulate, frequently also with
range thus
isfar reported from Guatemala to Costa several swollen and darkened nodes throughout
Rica, and the Guianas southward to Brazil and its Pinnae commonly borne in opposite
length.
(possibly) Bolivia. Although usually a smaller plant pairs from swollen, darkened nodes on the rachis.
than Ophioglossum reticulatum, the aspect of the Veins free or (in Christensenia) reticulate. Spo-
sterile lamina is similar in that the primary areoles rangia borne on the abaxial surface of leaves, with
contain a number of secondary areoles and usually thickened walls, lacking an annulus, separate and
some However, the secondary ar-
free veinlets. contiguous or fused into indurated synangia and
eoles are much more numerous in O. ellipticum, opening by terminal pores. Spores ellipsoidal and
and the walls are thinner and far less distinct than monolete or trilete, rugose or echinate, very nu-
those of the primary ones. Additionally, laminae merous (1,500-1,700) in each sporangium.
in O. ellipticum have a somewhat distinct midvein
and a pale, median band of tissue which persists Of the seven genera (about 1 50 species) in Mar-
to within a short distance of the apex. It is most attiaceae, two are represented in Peru: Danaea,
closely related to O. nudicaule, in fact, the plants confined to the Neotropics, and Marattia, pan-
heretofore recognized as O. ellipticum may be tropical. Next to Ophioglossaceae, it is considered
merely O. nudicaule with larger and more fully to be the most primitive of the living fern families
developed leaves (see Wagner et al., 1984). and can be easily recognized by the character of
the sporangia being fused into thick, elongate, dou-
ble-rowed synangia.

Family 2: MARATTIACEAE
References
Marattiaceae Bercht. & J. S. Presl, Prir. rostlin 1 :

272. 1820. TYPE: Marattia Sw. DEVRIESE, W. H.,ANDP. HARTING. 1853. Mon-
ographic des Marattiacees, Leiden, Nether-
Danaeaceae Agardh, Aphor. hot. 117. 1822. TYPE: lands.
Danaea Sm. TRYON, R. M., AND A. F. TRYON. 1982. Mar-
attiaceae, pp. 39-50, in Ferns and allied plants,
Stem fleshy, stout to massive, creeping or erect, Springer- Verlag, New York.
sometimes scaly. Leaves moderate in size to huge UNDERWOOD, L. M. 1909. Marattiaceae, in N.
(3 m or more), circinate in vernation, 1 -pinnate Amer. fl. 16: 15-23.

Key to Genera of Marattiaceae

a. Leaves 1-3-pinnate or more, monomorphic; synangia scattered, borne near the ends of veins
I. Marattia
a. Leaves 1 -pinnate, somewhat dimorphic (fertile ones commonly longer and narrower and with smaller
pinnae); synangia nearly covering the abaxial surface of fertile pinnae II. Danaea

I. Marattia sentially glabrous, but in some species sparsely to


moderately scaly or with a few trichomes on the
Marattia Sw., Prodr. 128. 1788. TYPE: M. alata axes abaxially. Penultimate divisions with axes
Sw. Figure 3. commonly broadly alate, the wings narrowing
abruptly at the base of each segment. Veins free.
Plants terrestrial. Leaves coarse, to ca. 3(-4) m Sporangia in two rows, opening by longitudinal
long, 2-4-pinnate, often ternate in architecture, slits, fused into bivalvate, ovoid synangia, which

monomorphic. Lamina with expanded tissue es- are borne near the ends of veins on a receptacle,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 13


FIG. 3. Marattia laevis: a, portion of lamina; b, segment base, with synangia. (a from Macbride 4817, F, b from
Haught 6145, Colombia, F.)

14 FIELDIANA: BOTANY
sessile or (in M. laevis) on short, thickened stalks. some Brazilian specimens of M. laevis with "soros
Spores ellipsoid, monolete. ... stipitati," upon which character Hooker (fol-
lowing Smith's unpublished description) partially
This is a pantropical genus of about 40 species, based the new species M. Kaulfussii (1839). Ob-
commonly occurring in dense, wet forests. It is viously Hooker had seen the type of M. laevis, and
readily separated from other Neotropical genera supposed it lacked the stalked synangia; therefore,
by the large, decompound, subternate leaves with he surmised that the Brazilian collections of Kaul-
stipular bases, and the distinctive ovoid synangia. fuss represented a new species. Finally in 1936
A single species is known in Peru. Alston (J. Bot., p. 174) combined the two names,
but failed to specify reasons for doing so. In turn-

ing to the British Museum for clarification of the


1. Marat tia laevis Sm., Plantarum Icones Inedi- matter, this reply was received from Miss Jose-
tae ... 2: /. 47. 1790. TYPE: as "Polypodium phine M. Camus: "I have examined the (type)
pedicellata, in pedunculat(o?), He
argento specimen in the Linnaean Herbarium. It does in-
Dominique (Dominican Republic, not Do- deed have synangia with very elongate receptacles.
minica), Thiery" (holotype, LINN). Figure 3. The original label bears the name Polypodium ped-
icellata in Linnaeus fil.'s hand. The 'awns' are pres-
Marattia alata sensu Raddi, PI. bras. nov. gen. 1 74, : ent on the upper surface of the pinnule axes... ."
t. 83-84. 1825.
Thus, there seems to be no question that the fern
Marattia Kaulfussii Hooker, Gen. fil., t. 26 (as to note).
considered to be M. Kaulfussii for nearly 1 50 years
1839. TYPE: Brazil, 1816, Cunningham s.n.

(holotype, BM!; photo, F!).


is M. laevis.

Eupodium Kaulfussii (Hooker) Hooker, Sec. cent, ferns,


/. 95. 1860. Amazonas: Prov. Bagua, Cordillera Colon SE of La
Marattia alata Sw. var. laevis (Sm.) Farw., Amer. Midi. Peca, Barbour 3900 (MO). Huanuco: Churubamba, Trail
Naturalist 12: 308. 1931. Puente Durand to Exito, Mt. Santo Toribio, Mexia 8250
(F, GH, MICH, NY, uc, us). Pasco: Cushi, Macbride 4817

Leaves 24 m long, commonly 3-pinnate-pin- (F, GH, us). Junin: Schunke Hacienda above San Ramon,

Killip & Smith 24540 (NY, us). Pichis Trail, Enenas,


natifid, at least as to base of lamina. Axes (at least
Killip & Smith 25738 (GH, NY, us). Chanchamayo Valley,
minor ones) sparsely to moderately provided on C. Schunke 671 (F). Cuzco: La Convention, above Knox's
abaxial side with filiform to lanceolate scales, these Cascade, Dudley 10619 (GH, MO).
orange to light brown, flaccid, 0.5-3 long, mm
often caducous. Costae of penultimate segments
bearing on the adaxial side 1 -several well-spaced, II. Danaea
often appressed, awns, these 0.2-1 long (or mm
on older leaves sometimes withering or broken Danaea Sm., Mem. Acad. Roy. Sci. (Turin) 5: 420.
away near their bases). Synangia 8-16(-18)-lo- 1793, nom. consent., not Allioni, 1785. TYPE:
cular, mature ones obviously stipitate, the stalks Danaea nodosa (L.) Sm. (Acrostichum nodos-
stout, to 0.7 mm
long. Spores echinate. um L.). Figure 4.

Occurring in dense forests, principally along the Plants terrestrial, with stems creeping to decum-
Cordilleras Centraland Oriental, in Amazonas, bent or erect. Leaves to 2 m
long, 1 -pinnate (or
Huanuco, Pasco, Junin, and Cuzco, 1400-2400 m. simple in 2 species outside Peru), monomorphic
Costa Rica; Cuba; Hispaniola; Colombia; Ven- to somewhat dimorphic (fertile ones commonly
ezuela; Brazil; Ecuador; Peru; Bolivia. longer and narrower and with smaller pinnae). Pet-
In various herbaria, M.
been confused
laevis has iole (in most species) nodose, scaly, bearing sti-
with M. alata, a species from the Greater Antilles. pules at the base. Rachis nodose, nonalate to con-
Although leaf outline is similar in each, M. laevis spicuously alate at least distally, glabrous or more
is easily distinguished by its stalked synangia and often with appressed scales. Lamina with a distinct
awned penultimate axes, these characters are apical segment, or this replaced by a proliferous
unique in the genus, at least in New World species. bud, provided with scales abaxially, those of the
There has been some confusion as to the identity laminar tissue minute and scattered. Sterile pinnae
of M. laevis, beginning with Smith's original de- opposite or subopposite, straight to subfalcate, ses-
scription, which mentioned nothing about the dis- sile or short-stalked, entire and often undulate, or
tinctive stalked synangia and adaxial awns. Kaul- serrate near the apex. Veins free. Fertile pinnae
fuss (Enum. fil., p. 32, 1824) mentioned seeing similar to the sterile ones, but commonly reduced

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 15


FIG. 4. Danaea moritziana: a, habit. Danaea nodosa: b, apex of sterile lamina; c, portion of fertile pinna with
synangia. (a from Tryon & Tryon 5278, F, b-c from Wurdack 1934, F.)

16 FIELDIANA: BOTANY
in size, and with vegetative tissue nearly lacking. America may be synonymous with some in the
Sporangia nearly covering the abaxial surface, co- West Indies. Although about 35 species have been
alescing into much elongated, indurated synangia recognized, 20-25 is a more realistic total to expect
and opening by terminal pores. Spores monolete, once a greatly needed revision has been under-
the surface prominently echinate. taken. Six species from Peru are recognized in the
following treatment.
Danaea
confined to the Neotropics. With its
is

nodose leaf axes and the distinctive synangia near-


ly covering the abaxial surface of simple, usually References
opposite, pinnae, it should not be confused with
other Peruvian genera. However, distinguishing PROCTOR, G. R. 1977. Danaea, pp. 45-49, in
species is another matter. To date there has been Howard, R. A., ed., Flora of the Lesser Antilles:
no comprehensive study of the genus, and many Leeward and Windward Islands, Vol. 2, Pteri-
of the characters used by Underwood to separate dophyta, Arnold Arboretum, Harvard Univer-
taxa are highly suspect: e.g., length of fertile pin- sity, Jamaica Plain, Mass.

nae, number of veins per centimeter, veins paired UNDERWOOD, L. M. 1 902. A review of the genus
in origin versus distinctly forked. Species previ- Danaea. Bull. Torrey Hot. Club, 29: 669-679.
ously thought of as distinct in Central or South

Key to Species of Danaea

a. Larger pinnae (5-)6-40 cm long; larger fertile pinnae 4-25 cm long


sterile b
b. Sterile pinnae 2-6 pairs; fertile pinnae 3-5 pairs 2. D. elliptica

b. Sterile pinnae (7-)8-16 pairs; fertile pinnae 7-25 pairs c


c. Petiole of mature plants lacking nodes; apices of sterile pinnae entire (very rarely serrulate);

larger sterile pinnae (2.5-)3-6.5 cm broad 1. D. nodosa

c. Petiole of mature plants with 1-3 nodes; apices of sterile pinnae serrate; larger sterile pinnae

1 .2-2.8 cm broad d
d. Sterilepinnae broadly oblanceolate, broadest above the middle, abruptly acuminate at the
apex; veins predominantly simple, occasionally paired in origin or forked
3. D. oblanceolata

d. Sterile pinnae lanceolate to oblong-lanceolate, broadest near or below the middle, tapering

gradually to an acuminate or attenuate apex; veins predominantly forked, only occasionally


simple or paired at the base 4. D. moritziana
a. Larger pinnae 1.5-3.5(-4) cm long; larger fertile pinnae 1.1-1.7 cm long
sterile e
e. Sterile pinnae strongly inequilateral at base, narrow and rounded basiscopically, much broader
and abruptly cuneate to truncate acroscopically; veins predominantly forked, some of them simple;
petiole nodes 1-3; terminal segment of sterile lamina equaling or longer than the pinnae, lacking
a proliferous bud 5. D. humilis

e. Sterilepinnae subequilateral at the truncate base; veins simple, rarely forked; petiole nodes lacking;
terminal segment of sterile lamina much shorter than larger pinnae, or replaced by a proliferous
bud 6. D. trichomanoides

1 . Danaea nodosa (L.) Sm., Mem. Acad. Roy. Sci. Danaea grandifolia Underw., N. Amer. fl. 16: 18. 1909.

(Turin) 5: 420. 1793 Figure 4b-c TYPE: Colombia, Valparaiso, Santa Marta, //.
H. Smith 992 (holotype, NY; isotypes, BM, F!, GH!,
MO!, us!).
Acrostichum nodosum L., Sp. pi. 1070. 1753. LEC-
TOTYPE (designated by Proctor, Flora Lesser
Antilles 2: 48. 1977): Plumier, Traite foug. Amer., Sterile leaves to 2 m long and 60 cm broad,
/. 108, based on specimen from Martinique or
imparipinnate; petiole lacking nodes, sparsely sca-

Dan^ealongifolia Desv., Ges. Naturf. Freunde Berlin


* or abundantly so at base, the scales mostly ap-

Mag. Neuesten Entdeck. Gesammten Naturk. 5: pressed, dark brown, peltate, nearly circular and
307. 1811. TYPE: "in antillis" (p). ca. 1 mm in diameter or to 3 mm
long, margins

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 17


often fimbriate; rachis not or scarcely alate, sparse- Sterile leaves to 1 m
long and 30 cm broad,
ly scaly; pinnae 7-16 pairs, subsessile to short- imparipinnate; petiole with 1-5 nodes, sparsely
stalked, elliptic to oblanceolate, larger ones 20-40 scaly, the scales appressed, dark brown, amor-
cm long and (2.5-)3-6.5 cm broad, base cuneate, phous, peltate, often fimbriate, commonly less than
apex long-acuminate, margins entire (very rarely 2 mm
long; rachis nonalate to narrowly so distally
serrulate along the acuminate tip), abaxial surface (or alate throughout in juvenile leaves), sparsely
and veins sparsely provided with minute (ca. 0. 1 scaly; pinnae 2-6 pairs, subsessile to short-stalked,
mm) scales, these appressed, nearly circular, dark more or less elliptic, 8-18 cm long, (2.5-)3 4.5 cm
or reddish brown, commonly erose to stellate; veins broad, base cuneate, apex acute to acuminate,
mostly paired or 1 -forked near the base. Fertile margins entire and plane to somewhat undulate,
pinnae 7-14 pairs, 12-25 cm long, 1.5-2.5 cm abaxial surface and veins very sparsely provided
broad, subsessile to short-stalked, the apex acute with minute (ca. 0. 1 mm) scales, these appressed,
to acuminate. nearly circular, dark or reddish brown, commonly
erose to stellate; veins mostly paired or 1 -forked
dense forests, 100-2100 m, Ama-
Terrestrial, in near the base. Fertile pinnae 3-5 pairs, 5-14 cm
zonas and Loreto to Cuzco and Madre de Dios. long, (l-)1.7-2.8(-3) cm broad, short-stalked, the
Southern Mexico to Panama; West Indies; Co- apex acute to acuminate.
lombia; Venezuela; Surinam; Ecuador; Peru; Bra-
zil. In dense forests, principally along the Cordillera
Danaea grandifolia is said to differ from D. no- Oriental, Amazonas to Junin, 125-1600 m.
dosa in the broader sterile pinnae and more broad- Southern Mexico to Panama; West Indies; the
ly cuneate bases of fertile pinnae. The latter char- Guianas to Colombia and south to Brazil and Peru.
acter is not consistent, and the former is hardly There is no justification in recognizing Rosen-
sufficient reason for separation even at infraspe- stock's var. crispula as a separate entity; suppos-
cific level. There is also some question as to the edly it differed from the typical in having one less

merit of separating D. nodosa and D. elliptica. pair of pinnae, these broadest at or above the mid-
While the characters of nodose or non-nodose pet- dle, with margins more crisped. All fall within the
iolesand pinnae number usually serve to distin- natural variation of this species.
guish the two, even these have not been reliable
consistently in specimens seen from Peru. Further Amazonas: Prov. de Bagua, valley of Rio Maranon
study is needed. near Cascadas de Mayasi, Wurdack 1934 (GH, us). San
Martin: On ridge in jungle E of Tingo Maria, Allard
21381 (GH, us). Loreto: Fierro Cano a km 4 del Centre
Ama/onas: E of Huampami, near Pedro Alberto, Ber- al. 1439 (MO).
Forestal Jenaro Herrera, Spichiger et
lin 1524 (MO). San Martin: Prov. Rioja, km 390, Pedro
Huanuco: Churubamba, Hacienda Mercedes, trail
Dist.
Ruiz-Moyobamba road. D. Smith & Vasquez 4726 (F, to Balsa-playa, Mexia 817 Oa (uc). Pasco: Prov. Oxa-
MO). Loreto: Balsapuerto (lower Rio Huallaga basin), Kil-
pampa, Palcazu Valley, Iscozacin, Foster 9502 (MO).
lip & Smith 28410 (GH, us). Huanuco: Prov. Pachitea, Junin: Pichis Trail, Yapas, Killip & Smith 25507 (GH,
Dist. Honoria, en frente a Tournavista, al este del Rio
NY, US).
Pachitea, J. Schunke 1800 (F, GH). Pasco: Puerto Ber-
mudez (as Junin), Killip & Smith 2651 1 (NY, us). Junin:
Prov. Satipo, near Satipo, van der Werffet al. 8644 (MO).
Cuzco: Prov. Quispicanchi, Punkiri Rio Arriba, Vargas 3. Danaea oblanceolata Stolze, Amer. Fern J. 77:
16121 (GH). Madre de Dios: Prov. Tambopata, SSW of 33. 1 987. TYPE: Peru, Dept. Pasco (as Junin),
Puerto Maldonado, Barbour 5178 (F, GH, MO).
Cahuapanas, on Rio Pichis, Killip & Smith
26777 (holotype, us!; frag., F!).

2. Danaea elliptica Sm. in Rees, Cycl. 1 1 Danaea :


40-50 cm long, 13-18 cm broad,
Sterile leaves
No. 2. 1808. LECTOTYPE (designated by
apical segment (on mature leaves) replaced by a
Proctor, Flora Lesser Antilles 2: 48. 1977):
proliferous bud; petiole with 1-2 nodes, moder-
Sloane Herb. 1: 85, based on plant from Mt.
ately to abundantly scaly; rachis narrowly alate;
Diablo, Jamaica (BM).
pinnae 10-12 pairs, mostly short-stalked, oblong
to (more commonly) broadly oblanceolate, larger
Danaea elliptica var. crispula Rosenst., Repert. Spec. ones 7-11 cm long and 2-2.8 cm broad, inequi-
Nov. Regni Veg. 7: 310. 1909. TYPE: Peru, San
lateral at base, narrow and rounded to cordate
Martin, "In montibus secus flumen Mayo, prope
Tarapoto," Spruce 4770 (holotype, K; isotypes, basiscopically, broader and cuneate acroscopical-
GH!, L, P!, uc!. us!; photo, F of L!). ly, terminating abruptly in an acuminate and ser-

18 FIELDIANA: BOTANY
rate apex, abaxial surface amply provided with Colombia; Venezuela; Peru.
minute, dark brown scales; veins commonly sim- Except for Danaea nodosa, this is the most widely
ple, but sometimes paired at origin or forked. Fer- distributed representative of Danaea in Peru.
tile pinnae 12-14 pairs, larger ones 7-8 cm long Moreover it is likely that the Central American D.
and 0.8-1 cm broad, short-stalked, the apex ob- cuspidata Liebm. also belongs here, along with one
tuse. or two West Indian species. A number of taxa with
pinnae under 2.5 cm broad were separated by Un-
0-500 m, thus far
Terrestrial in dense forests, derwood (1902), merely on the degree of forking
known only from Peru: Pasco and Ucayali. and spacing of veins, an inconsistent character cor-
This is perhaps most closely related to D. alata related rarely or not at all by other features, hence
Sm., of the West Indies and Venezuela, especially a comprehensive revision of the genus should prove
in that both species have predominantly simple many of these to be synonymous. Peruvian spec-
veins. However, in D. oblanceolata pinnae are imens of D. moritziana apparently are more vari-
fewer and relatively shorter and broader, and most able than in other areas, sometimes having smaller
are broadest well above the middle, where the and fewer pinnae as in Venezuela, sometimes with
margins then bend abruptly to a short-acuminate more and larger pinnae as in "D. cuspidata" of
apex. In D. alata, as in all members of the D. Costa Rica. A few very large specimens from Tin-
moritziana complex, pinnae are broadest at or near go Maria have pinnae with rather glossy under-
the middle, from whence they taper gradually to surfaces and extremely attenuate apices. Very
a moderately acuminate or attenuate apex. closely related to this is D. alata Sm. of the Lesser
Antilles, Trinidad, and Venezuela. Although D.
Pasco: Oxapampa, Palcazu Valley, Iscozacin, ./?. Foster alata tends to have more numerous pinnae than
9466 (MO), 10049 (F). Ucayali: Vicinity of Aguaytia, Croat D. moritziana, the only consistent difference be-
20938 (MO).
tween the two is that of simple versus forked veins.
To further add to the confusion, some specimens
of D. moritziana in Peru have been incorrectly
determined as D. stenophylla Kunze, which is
4. Danaea moritziana Presl, Abh. Konigl. Bohm
Ges. Wiss. 5(4): 35. 1845. TYPE: Venezuela, merely a synonym for D. alata.
Underwood (1902) described both of the latter
Colonia Tovar, Moritz 257 (holotype, PR?; is-
as having unforked veins and distinguished them
otype, L; frag., us!; photo, us). Figure 4a.
principally by the degree of crowding of veins, and
that D. stenophylla was a "small" plant. Later
Sterile leaves to 1.2 m long and 35 cm broad
(1909) he reversed himself by describing D. steno-
(somewhat broader than fertile ones), imparipin-
phylla as a "rather tall plant" and as having mostly
nate, or sometimes with the apical segment re-
forked veins. Obviously he had not seen the type
placed by a proliferous bud; petiole with 1-3 nodes,
collection, for the isotype (2 sheets) at Kew con-
sparsely scaly; rachis scarcely alate, or slightly so
tains leaves to nearly a meter long, with veins
in younger plants; pinnae 8-16 pairs, subsessile to
obviously simple (only rarely forked), differing
short-stalked, elliptic to oblong-lanceolate, larger
from D. alata in no way that we can perceive. We
ones (5-)6-20 cm long and 1 .2-2.2 cm broad, ine-
cannot be certain which West Indian plants with
quilateral at base, narrow and rounded to cordate
forked veins were seen by Underwood, and later
basiscopically, broader and cuneate to subtruncate
by Proctor (1977), but perhaps they are D. ja-
acroscopically, gradually tapering to an acuminate
maicensis.
or long-attenuate apex, margin subentire but con-
spicuously serrate at apex, veins (and often the
tissuebetween the veins) amply provided with mi- Cajamarca: Tambillo, Jelski 1073 (P). Amazonas: Bag-
nute scales; veins predominantly forked at or near ua, ca. 25 km E of La Peca, Barbour 2969 (MO). San
the base, occasionally simple, or paired at origin. Martin: Monte Campana, Tarapoto, Spruce 4711 (P).
Loreto: Rio Corrientes, Cachuela, McDaniel 11198 (F,
Fertile pinnae 8-15 pairs, larger ones 4-11 cm GH, MO). Huanuco: Tingo Maria, Tryon & Tryon 5278
long, 0.6-0.8 cm broad, subsessile or short-stalked, (F, GH, uc, us). Pasco: Prov. Oxapampa, San Alberto,
the apex acute to apiculate. Cordillera de Yanachaga, van der Werffet al. 8434 (MO).
Junin: Chanchamayo Valley, C. Schunke 156 (F, us).
Cuzco: Prov. La Convention, ca. 10 km from Hacienda
Terrestrial, in dense forests, often along streams
Luisiana, Dudley 10412 (GH). Madre de Dios: Prov. Manu,
or in ravines, 300-2300 m, Cajamarca to Loreto, Cerro de Pantiacolla, Rio Palotoa 10-15 km NNWof
south to Cuzco and Madre de Dios. Shintuya, Foster 10750 (F).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 19


5. Danaea humilis Moore, Index fil. 286. 1861. rounded to subacute, margin slightly crispate; veins
LECTOTYPE (designated by Lellinger in Proc. predominantly simple, rarely forked. Fertile lam-
Biol. Soc. Wash. 89: 710. 1987): Peru, Dept. ina with rachis scarcely alate; pinnae 1.4-1.7 cm
San Martin, Tarapoto, Spruce 4769 (lecto- long, 0.3-0.4 cm broad, obviously stalked, apex
type, K; isolectotypes, BM!, GH!, L!, P!, us!). rounded.

Sterile leaves 20-34 cm long, 3.5-7 cm broad, Thus far known


only from the type collection.
larger than the fertile ones, imparipinnate, the ter- When Moore published the description of the
minal segment equaling or longer than the largest species, taken from a Spruce manuscript, he noted
pinnae, with or without a proliferous bud; petiole that pinnae were "unequal at the base." Perhaps
with 1-3 nodes, amply to abundantly scaly; lamina this conclusion was drawn from observing the cris-
slightly shorter than the petiole, the tissue between pate margins of some of the pinnae which were
the veins abundantly provided with minute scales; partially folded on drying. However, study of the
rachis conspicuously alate, the margins of wings two type specimens reveals that the bases of most
sometimes revolute, but not crispate; pinnae 14 pinnae are subequally truncate and approximately
24 pairs, oblong-elliptic, subfalcate, 2.2-3.5(-4) cm as broad on either side of the costa. This is in
long, 0.6-1(-1.4) cm broad, strongly inequilateral marked contrast with most species of Danaea,
at base, narrow and rounded basiscopically, much whose bases are commonly broader and less
broader and abruptly cuneate to truncate acros- abruptly tapered on the acroscopic side.
copically, apex acute to subacuminate and usually Danaea trichomanoides is comparable to D.
serrulate,margin subentire, essentially plane; veins wendlandii Reichb. f., from Costa Rica, for the
predominantly forked above the base, yet many two are similar in size, and have nodeless petioles
of them simple. Fertile lamina scarcely alate along and crispate pinnae, with the presence (usually) of
rachis; pinnae 12-25 pairs, 1.1-1.4 cm long, 0.2- a terminal bud. However, the latter has pinnae
0.3 cm broad, subsessile to short-stalked, mostly with inequilateral bases,veins predominantly
acute to subapiculate. forking, and rachis wing plane or revolute (not
crispate).
In Peru known only from the type collection:
San Martin.
Rare, terrestrial, in rain forests, 1000-1400 m,
Venezuela; Colombia; Ecuador, Peru; Bolivia.
This and Danaea trichomanoides are among the Family 3: OSMUNDACEAE
smallest species of the genus, with even the larger
sterile pinnae commonly less than 3 cm long. Oth- Osmundaceae Bercht. & J. S. Presl, Prir. rostlin 1:

er than size,however, they share few other similar 272. 1820. TYPE: Osmunda L.

features, as evidenced by the characters listed in


the key. Stem massive, woody, decumbent to erect, lack-

ing indument, with hard, fibrous roots. Leaves cir-


cinate in vernation, densely caespitose, pinnate to
6. Danaea trichomanoides Moore, Index fil. 288. pinnately compound, monomorphic to (in Os-
1861. TYPE: Peru, "in monte Guayrapurima, munda) partly or fully dimorphic. Petiole with an
prope Tarapoto" (San Martin), Aug. 1856, expanded, stipular base, not articulate to the stem.
Spruce 4710 (holotype, K!; frag., NY!; photos, Veins free. Sporangia not in definite sori, borne
F, GH; isotypes, BM!, P!). abaxially along the segments or (in Osmunda)
completely replacing the vegetative tissue of the
Sterile leaves 10-12 cm long, 34 cm broad, lamina or of only some of the pinnae, exindusiate,
shorter than the fertile ones, with a terminal seg- short-stalked, with walls 1 cell thick, the annulus
ment much shorter than the larger pinnae, or re- lacking or of only a few thickened cells near the
placed by a proliferous bud; petiole lacking nodes, distal end. Spores uniform, green, 1 20-5 1 2 in each

abundantly scaly; lamina 2-4 times longer than sporangium.


the petiole, the tissue between veins essentially
lacking scales; rachis broadly crispate-alate; pin- The three genera of the Osmundaceae are rep-
nae 11-15 pairs, oblong, 1.5-2.2 cm long, 0.5-0.9 resented by 1 5-20 species. Osmunda is nearly cos-
cm broad, subequilateral at the truncate base, apex mopolitan, while Leptopteris and Todea are con-

20 FIELDIANA: BOTANY
fined to the Old World. This is a very old fern I. Osmunda
family; only Ophioglossaceae and Marattiaceae are
considered to be more primitive. Like the other Osmunda L., Sp. pi. 1063. 1 753. TYPE: O. regalis
leptosporangiate members of the Filicales, Os- L. Figure 5.
mundaceae have sporangial walls only one cell

thick, yet the family is similar to the Eusporan- Plants terrestrial. Leaves coarse, 1-2 or long- m
giatae in that the annulus is lacking or is composed er, 1-2-pinnate, completely or partially dimor-
of only a few thickened cells. phic, the fertile ones (or portions of them) lacking
green leaf tissue. Sterile lamina commonly gla-
brous at maturity, but the axes often sparsely to
References moderately provided with trichomes. Veins free,
at least 1 -forked. Sporangia relatively
large, de-
BENEDICT, R. C. 1909. Osmundaceae, in N. veloping simultaneously, commonly borne in
Amer. fl., 16: 27-28. clusters on the segments of the fertile leaves. Spores

TRYON, R. M., AND A. F. TRYON. 1982. Os- tetrahedral-globose, trilete, rugose or crested with
mundaceae, pp. 50-57, in Ferns and allied plants, slender echinate processes.
Springer- Verlag, New York.
This is a genus of about 1 species which gen-
erally grow in swampy areas in temperate and

tropical regions of both hemispheres. Two species


occur in Peru.

Key to Species of Osmunda

a. Sterile lamina 1-pinnate-pinnatisect; ultimate segments entire, the veins commonly 1 -forked; fertile
lamina nonfoliaceous throughout 1 O. cinnamomea .

a. Sterile lamina 2-pinnate; ultimate segments serrulate, the veins commonly 2-forked; fertile lamina
nonfoliaceous and fertile only in the distal portion 2. O. regalis var. spectabilis

1. Osmunda cinnamomea L., Sp. pi. 1066. 1753. trichomes abundantly interspersed among the
TYPE: United States, Maryland, Kalm (ho- crowded sporangia.
lotype, LINN 1 244. 1 2; photo, F). Figure 5c-d.
In wet places, usually in sphagnum swamps,

Leaves completely dimorphic, vegetative tissue 1700-2800 m, Cajamarca, Amazonas, and Pasco.
Eastern and central United States and Canada;
essentially lacking in fertile ones. Petiole glabrous
or moderately provided with tortuous, filiform, West Indies; southern Mexico to Honduras; Costa
Rica; Colombia; Venezuela; Brazil; Peru; Para-
pluricellular, reddish brown trichomes, sheathed
at the base. Sterile lamina 1-pinnate-pinnatifid, guay; southeast Asia.
tissue glabrous, but with filiform trichomes as on Some earlier authors recognized this in the Neo-
the petiole appearing especially at bases of pinnae, tropics as var. imbricala (Kunze) Milde, distin-

rachis narrowly alate adaxially; pinnae to 10 cm guishing it from North American plants on the
basis of darker-colored tomentum and thicker lam-
long, spreading to strongly ascending, subopposite,
articulate with age at the rachis, deeply incised to
ina. We find little justification for the separation.

form broad, subfalcate, obtuse or subacute seg-


ments, the segment margins subentire and bearing Cajamarca: Jaen, Quebrada de Pajonal, above Taba-
conas, 19km ESE Huancabamba, Fosberg 27795 (MICH,
short, filiform trichomes; veins 1 -forked. Fertile
NY). Amazonas: Chachapoyas, Jalca zone, 3-6 km of W
lamina nearly 2-pinnate, the pinnae more strongly Molinopampa, Wurdack 1404 (us). Pasco: (as Junin)
ascending than the sterile ones, with dark, filiform Pichis trail, Enenas, Killip & Smith 25708 (NY, us).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 21


FIG. 5. Osmunda regalis var. spectabilis: a, habit; b, sterile ultimate segment. Osmunda cinnamomea: c, habit; d,
sterile ultimate segment. (From Stolze, Ferns and fern allies of Guatemala, 1976.)

22 FIELDIANA: BOTANY
2. Osmunda regalis L. var. spectabilis (Willd.) A. length of their stalks. The characters are too in-
Gray, Manual ed. 2: 600. 1856. Figure 5a-b. significant and inconsistent to merit such a dis-
tinction.
Osmunda spectabilis Willd., Sp. pi. 5: 98. 1810. TYPE:
United States, Pennsylvania, Muhlenberg (holo- Amazonas: Mendoza, on rocky wall of ravine, Woyl-
type, B, Herb. Willd. 19504; photo, GH). kowski 8253 (GH, MO, us). Prov. Bongara, W
& S margins
Osmunda palustris Schrader, Gott. Gel. Anz. 866. of Laguna Pomacocha, Wurdack 896 (us). Pasco: Prov.
1824. TYPE: not located.
Oxapampa, Canyon de Huancabamba, Leon 612 (GH).
Osmunda regalis ssp. palustris (Schrader) Love & Love, Cuzco: Empalizada, Biies 1705 (us).
Taxon 26: 324. 1977.

Leaves partly dimorphic, fertile ones with fertile

and sterile portions combined on the same lamina, Family 4: SCHIZAEACEAE


the fertile pinnae commonly distal and essentially
lacking vegetative tissue. Petiole glabrous at ma- Schizaeaceae Kaulf., Wesen farrenkr. 119. 1827.
turity, sheathed at the base. Sterile lamina 2-pin- TYPE: Schizaea Sm.
nate, chartaceous to subcoriaceous, tissue gla-
brous, but a few tortuous, pluricellular, reddish Stem erect to decumbent, usually small and
brown trichomes often clustered at the base of sometimes branched, or long-creeping, slender and
pinnules, the rachis nonalate; pinnae to 20 cm freely branched, provided with trichomes or (in
long, strongly ascending, subopposite, short- Mohria of Africa) with scales. Leaves circinate in
stalked, articulate with age at the rachis, costae vernation, entire or filiform, or dichotomous, or
very narrowly alate (at least distally) on adaxial pinnate, glabrous to pubescent or (in Mohria) with
side; pinnules distant, subopposite, short-stalked, scales, partially or wholly dimorphic. Sporangia
articulate with age at the costa, oblong to narrow- borne abaxially on slightly to strongly modified
elliptic, obtuse to subacute, margins serrulate. Veins portions of the leaf, separate, or crowded on each
mostly 2-forked. Fertile pinnae 2(-3)-pinnate, more side of a vein, or in loose clusters on wholly fertile
strongly ascending than the sterile ones, the crowd- panicles, sessile, or with a short,
many-rowed stalk,
ed sporangia almost completely replacing the leaf and an apical annulus. Spores tetrahedral-globose
tissue. and trilete, or ellipsoidal and monolete, lacking
chlorophyll.
In damp places near lakes or streams, occasion-
ally among rocks, 1300-2200 m, Amazonas, Pas- This is an old family, with the four extant genera
co, Cuzco. so highly diverse in habit and form that some
Eastern and central United States and Canada; authors prefer to segregate them into distinct fam-
West Indies; southern Mexico; Guatemala; Hon- ilies. Mohria is confined to Africa and Madagas-
duras; Costa Rica; Colombia; Venezuela; Brazil; car; Anemia is inAmerica, Africa, and southern
Ecuador, Peru; Paraguay; Uruguay. India; the other two genera are pantropic. All ex-
Variety regalis, which occurs in the Old World, cept Mohria occur throughout tropical America
is said to differ in its broader leaves, narrower and are all well represented in Peru.
panicles, and more numerous blackish to casta-
neous trichomes persistent along the rachis. Al-
though specimens we have seen from both the References
New and Old World do not seem to differ strongly
or consistently enough to warrant separation, we LELLINGER, D. B. 1969. Schizaeaceae, in The
maintain the traditional varietal distinction for botany of the Guayana Highland Part VIII.
purposes of this treatment. Earlier workers pro- Mem. New York Bot. Gard., 18: 1-11.
posed yet another variant, ssp. palustris, for South MAXON, W. R. 1909. Schizaeaceae, in N. Amer.
American plants, based on size of segments and fl., 16: 31-52.

Key to Genera of Schizaeaceae

a. Leaves erect or suberect; pinnae lacking arrested buds in branch axils; sporangia not covered by a
laminar flange b

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 23


b. Leaf pinnate or pinnately decompound, rarely pinnatifid I. Anemia
b. Leaf filiform or with dichotomous veins or branches III. Schizaea
a. Leaves vinelike, spreading and scandent; pinnae short-stalked, with an arrested bud in the axil of
the first, pseudodichotomous branch; sporangia each covered by a laminar flange . . II. Lygodium

I. Anemia The genus contains about 80 species, which oc-


cur in tropical to subtropical regions of both hemi-
Anemia Sw., Syn. fil. 6, 155. 1806, nom. conserv. spheres, predominantly in the Neotropics. Species
(sometimes as "Aneimia"). TYPE: Anemia limits are not always clearly drawn, as diagnostic

phyllitidis (L.) Sw. (Osmunda phyllitidis L.). features are often quite variable, such as: relative
Figure 6. length of fertile pinnae and sterile portion of lam-
ina; color of rhizome indument; degree of dissec-
Coptophyllum Gardn., London J. Bot. 1: 133. 1842. tion of pinnae. Color of trichomes is particularly
TYPE: Coptophyllum buniifolium Gardn. = Ane- difficult to assess, inasmuch as each observer sees
mia buniifolia (Gardn.) Moore.
colors in different ways, and definitions of color
Anemirhiza J. Sm., Seemann, Bot. Voy. Herald 243.
1854. TYPE: Anemirhiza adiantifolia (L.) J. Sm.
terms are usually difficult to interpret. Rhizome
(Osmunda adiantifolia L.)
= Anemia adiantifolia indument in Anemia generally grades from orange
(L.) Sw. to reddish orange to red to brownish red; in many
specimens the distinctions are clear, but in others
Plants terrestrial. Stem decumbent to creeping, they are somewhat inconstant. Hybridization is
rarely erect, bearing short to long trichomes and not uncommon in the genus and must be taken
fibrous roots. Leaves ca. 1-75 cm long, erect, par- into consideration when attempting identification
tially (as in Peruvian species) dimorphic, with a of collections. The following key will be more ef-
pair of fertile pinnae at the base of or below the fective if attention is given to combinations of
sterile lamina, or with similar leaves longer and characters, whenever possible.
more erect than the wholly sterile leaves, or wholly
dimorphic, the whole leaf either sterile or fertile.
Sterile lamina 1-2-pinnate (rarely pinnatifid); veins Reference
free, or rarely anastomosing without included free
veinlets. Sporangia borne on fertile segments which MICKEL, J. T. 1962. A monographic study of the
are reduced to mere axes or have narrow borders fern genus Anemia, subgenus Coptophyllum.
of laminar tissue along the axes. Spores tetrahe- Iowa State Coll. J. Sci., 36: 349-482.
dral-globose, trilete.

Key to Species of Anemia

a. Fertile pinnae borne 3 mm


or more below the base of the sterile lamina; ultimate fertile segments
with narrow (but distinct) borders of laminar tissue along the axes b
b. Sterile pinnae crenate to pinnatisect, not cut entirely to the costa (except rarely as to proximal
lobes of basal pinnae) 2. A. villosa
b. Sterile pinnae 1-2-pinnate c
c. Apex of stem ascending or erect, covered with petiole bases; fertile pinnae commonly shorter
than the sterile portion of the lamina d
d. Petiole slender (less than 1 mm
thick), dark brown; fertile pinnae suberect, with stalk 0.3-
0.8 cm long 1 A. clinata .

d. Petiole stout (1-2 mm thick), commonly yellow; fertile pinnae erect, with stalk 1.5-5 cm
long 3. A. flexuosa
c. Apex of stem horizontal, exposed beyond the most recent petiole bases; fertile pinnae commonly
longer than sterile portion of the lamina e
e. Lamina hirsute to glabrate, petiole commonly glabrate and atropurpureous (rarely light
brown to yellow) 4. A. ferruginea

24 FIELDIANA: BOTANY
Lamina and petiole conspicuously lanate, petiole yellow
e. 5. A. myriophylla

Fertilepinnae borne at base of sterile lamina, or less than 1 mm


below it; ultimate fertile segments
with laminar tissue lacking or essentially so f
f. Veins free; stalk of fertile pinnae usually 2-4 times the length of the panicle; sterile pinnae less
than 4 cm long g
g. 8-25 cm long; lamina gradually reduced to a pinnatind apex (or
Petiole (at least the fertile)
rarely terminating in a subconform lanceolate segment); apex of sterile pinnae acute or subacute
h
h. Sterile pinnae mostly deeply incised, the segments commonly narrow and cuneate
6. A. hirsute

h. Sterile pinnae crenulate or denticulate, or rarely with a few deep lobes on several proximal

pinnae 7. A. pastinacaria

g. Petiole less than 6 cm long; lamina abruptly terminating in a broadly obovate or obdeltoid
apical segment; apex of sterile pinnae broadly rounded 8. A. oblongifolia

f. Veins anastomosing; stalk of fertile pinnae usually equaling or shorter than the panicle; sterile
pinnae (larger ones) 5-14 cm long 9. A. phyllitidis

1. Anemia din at a Mickel, Amer. Fern J. 56: 58. 2. Anemia villosa Willd., Sp. pi. ed. 4, 5: 92. 1 8 0.
1

1966. TYPE: Peru, Junin, along Rio Perene, TYPE: "America meridionale," Humboldt &
Killip & Smith 21594 (holotype, us!; isotypes, Bonpland (holotype, B, Herb. Willd. 19496;
F!, NY!). photos, GH, us). Venezuela, Prov. Cumana,
"Between Catuaro and Cariaco": fide HBK.,
Stem with apex ascending and thickly covered Nov. gen. sp. fol. ed. 1: 26. 1815.
with petiole bases, densely provided with orange
trichomes. Leaves 10-25 cm long, 4-9 cm broad, Stem with apex ascending and thickly covered
pilose to hirsute, the suberect fertile pinnae borne with petiole bases, or the apex horizontal and ex-
8-16 mm below the sterile portion of the lamina; posed beyond the most recent petioles, densely
petiole dark brown, less than mm thick. Sterile
1 provided with orange to brownish red trichomes.
lamina 1 -pinnate-pinnatisect to 2-pinnate, with 5- Leaves 10-55 cm long, villous to hirsute, the erect
1 1 pairs of pinnae; pinnae dissected nearly or quite fertilepinnae borne 3-18 mm
below the base of
to the costa, the ultimate segments joined at the the sterile portion of the lamina; petiole yellow to
base or discrete, but broadly adnate, not stalked; light brown, 1-2 mm thick. Sterile lamina 1 -pin-
veins free. Fertile pinnae suberect, usually shorter nate to pinnate-pinnatisect, with 7-19 pairs of pin-
than the sterile portion of the lamina, short-pe- nae; pinnae with rounded lobes, not cut entirely
tiolulate (stalk 0.3-0.8 cm long), the ultimate fer- to the costa (except rarely as to proximal lobes of
tile segments with narrow bands of laminar tissue basal pinnae); veins free. Fertile pinnae erect,
on each side of the axes. slightly shorter than to (more typically) somewhat
longer than the sterile portion of the lamina, their
In thickets and deep forests, 600-1 300 m, Junin. stalks 1-5 cm long, the ultimate fertile segments

Panama; Colombia and Venezuela to Bolivia with narrow bands of laminar tissue on each side
and western Brazil. of the axes.
One of the distinguishing features of clinata A
is the suberect or slightly spreading position of the
On rocky, open slopes, 900-2000 m, Cajamarca,
fertile pinnae, which are rigidly erect in most species
Amazonas, San Martin, Ayacucho, Cuzco.
of Anemia. It is also one of the smaller and more
Surinam to Colombia, south to Peru; east coast
delicate species, with slender, wiry petioles less
of Brazil.
than 1 mm
thick.
This appears to differ only quantitatively from
Anemia flexuosa, the latter with sterile pinnae more
Junin: Near La Merced, E of Quimiri Bridge, Killip &
Smith 23591 (NY, us). Colonia Perene, Killip & Smith strongly dissected and, typically, with fertile pin-
25036 (F, NY, us). La Merced, Kunkel 564 (OH). Chan- nae shorter than the sterile portion of the lamina.
chamayo Valley, C. Schunke 78 (F, us). Fertile pinnae of A. villosa are commonly longer

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 25


FIG. 6. Anemia paslinacaria: a, habit; b, apex of fertile pinna. Anemia phyllitidis: c, sterile pinna, (a-b from Irwin
et al. 24260, Brazil, p, c from Stork & Morton 9432, F.)

26 FIELDIANA: BOTANY
than the lamina, but occasionally equal or exceed gas 19836, Prov. La Convencion, alt. 1650 (GH). m
it. Evidently these are the reasons why Prantl con- Leaves are much thinner in texture than typical
sidered it merely a variety of A. flexuosa. See Mickel A. flexuosa, with some anastomosing veins, and
(1962) for more detailed discussion. fertile pinnules are long-petiolulate.

Cajamarca: Prov. Jaen, San Patricio, Chontali, Chi- Amazonas: Rodriguez de Mendoza, Soukup 5023 (us).
moy65 (USM). Amazonas: Prov. Chachapoyas, Cano Santa Huanuco: Vilcabamba, on Rio Chinchao, Macbride4985
Lucia, Wurdack 599 (F, GH, NY, uc, us). San Martin: (F, NY, us), Stork & Morton 9876 (F, uc, us). Junin:

Tarapoto, in monte Campana, Spruce 4708 (GH, NY, p, Huacapistana, Killip & Smith 24152, 24324 (F, NY, us).
us). Ayacucho: Prov. La Mar, Aina, Ldpez & Soukup Ayacucho: Ccarrapa, between Huanta and R;o Apuri-
15.088 (GH). Cuzco: Prov. La Convencion, Urusaiwa, mac, Killip & Smith 22319 (F, GH, NY, us). Cuzco: Prov.
Vargas 22349 (GH). La Convencion, Valley of the Sambray, Mexia 8038 (F,
GH, MO, uc).

3. Anemia flexuosa (Savigny) Sw., Syn. fil. 156.


1806.
4. Anemia ferruginea HBK., Nov. gen. sp. 1: 32.
1815, var. ferruginea. TYPE: Venezuela, Prov.
Osmunda flexuosa Savigny in Lam., Encycl. 4: 652.
1797. TYPE: based on undesignated specimen in
Cumana, "prope Guardia de San Augustin,"
Jussieu herbarium (not located).
Humboldt & Bonpland (holotype, P?).
Anemia flexuosa var. setosa Prantl, Unters. Morph.
Gefasskrypt. 2: 95. 1881. SYNTYPES: Brazil, Stem with apex horizontal, and exposed beyond
Lagoa Santa, Warming (not located): Peru, Cu- the most recent petioles, densely provided with
chero, Poeppig (not located). ISOSYNTYPE:
red or brownish red trichomes. Leaves 10-55 cm
Poeppig (L; photos, GH, us).
long, the erect fertile pinnae borne 3-7 mm
below
the base of the sterile portion of the lamina; petiole
Stem with apex ascending and thickly covered
with petiole bases, densely provided with orange commonly atropurpureous (rarely yellowish), gla-

or reddish orange trichomes. Leaves 18-65 cm brate, 1-2 mm lamina 2-pinnate,


thick. Sterile
hirsute to subglabrous, with 8-12 pairs of pinnae;
long, hirsute or glabrate, the erect fertile pinnae
borne 5-25 mm
below the base of the sterile por- pinnae dissected to the costa, the pinnules mostly
discrete, adnate at base, acute or subacute at apex,
tion of the lamina; petiole yellow to light brown,
1-2 mmthick. Sterile lamina 2-pinnate (some-
sometimes those of proximal pinnae shallowly
times 2-pinnate-pinnatifid), with 6-12 pairs of pinnatifid; veins free. Fertile pinnae erect, com-
pinnae; pinnae (many of them) divided to the costa
monly longer than the sterile portion of the lamina,
into broadly adnate, obtuse pinnules; veins free.
their stalks 1 .5-6 cm long, the ultimate fertile seg-

shorter than the


ments with narrow bands of laminar tissue on each
Fertile pinnae erect, commonly
side of the axes.
sterile portion of the lamina (rarely equal to it or
slightly longer), their stalks 1.5-5 cm long, the
In sunny places, open woods and clearings, on
ultimate fertile segments with narrow bands of

laminar tissue on each side of the axes. rocky slopes and clay banks, 850-2200 m, San
Martin, Huancavelica, Cuzco.
and open woods or on moist rocky
In thickets Honduras; Guyana to Colombia, south along
the Andes to Bolivia and Brazil.
slopes or clay banks, 900-3000 m, Amazonas,
Huanuco, Junin, Ayacucho, Cuzco.
The variety ahenobarba (Christ) Mickel, from
Surinam to Colombia, south to Bolivia. Brazil, differs in its more deeply dissected pinnules

Peruvian specimens of Anemia flexuosa with acute to acuminate tips. Both are difficult to
Many
are to be found in various herbaria; especially distinguish from Anemia tomentosa (Savigny) Sw.,
abundant are collections from Huanuco, Junin, especially var. anthriscifolia (Schrader) Mickel,
and Cuzco. This is a highly variable species, par-
from northern Argentina, Bolivia, Paraguay, and
southeastern Brazil. See Mickel (1962) for further
ticularly in the dissection of sterile laminae, some-
times 2-pinnate-pinnatifid, typically 2-pinnate, comments.
occasionally approaching the 1 -pinnate-pinnatifid San Martin: Tarapoto, Spruce 4044 (GH, NY, p, us).
condition of A. villosa (q.v.) and thus is often con- Huancavelica: Prov. Tayacaja, Dist. Huachocolpa, near

fused with it. As in some


other species of Anemia, Quintabamba, Tovar 4157, 4693 (GH). Cuzco: Prov.
Convencion, Valley of the Sambray, Mexia 8038 (NY,
was a hybrid
this tends to hybridize; in fact there
us). Prov. Convencion, Hacienda Sahuayaco, Vargas
found in Cuzco: A. flexuosa x A. phyllitidis, Var- 1656 (GH, MO).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 27


5. Anemia myriophylla Christ, Bull. Herb. Bois- pinnae to 2.5 cm long and 0.8 cm broad, com-
sier 2(7): 793. 1907. TYPE: Bolivia, Padcaya, monly inequilateral at base, truncate acroscopi-
Fiebrig 2541 (isotype, p). cally and cuneate basiscopically, obliquely incised
into linear or narrowly cuneate segments, obtuse
Stem with apex horizontal and exposed beyond to subacute at apex; veins free. Fertile pinnae to
the most recent petioles, densely provided with 18 cm long, the stalk (1.5-)2-3 times the length of
reddish to orange trichomes. Leaves 20-50 cm the panicle, the ultimate fertile segments essen-
long, the erect fertile pinnae borne 3-8 below mm tially lacking laminar tissue.
the base of the sterile portion of the lamina; petiole
yellow, conspicuously lanate, 1-2 mm
thick. Ster- on slopes or mesas, 1 200-
In dry, grassy areas,
ile lamina 2- to nearly 3-pinnate, conspicuously 2 1 50 m, Huanuco, Cuzco.
lanate, with 7-12 pairs of pinnae; pinnae with pin- Mexico to Panama; Greater Antilles; Trinidad
nules cut deeply or quite to the costule, the tips of & Tobago; Venezuela and Colombia south to Bo-
ultimate segments commonly broadly rounded; livia and Brazil.
veins free. Fertile pinnae erect,
commonly longer This is scarcely distinct from Anemia pastina-
than the portion of the lamina, their stalks
sterile caria, essentially differing only by the characters
1.5-5 cm long, the ultimate fertile segments with noted in the key. More detailed studies are needed
narrow bands of marginal tissue on each side of to clarify relationships of the several variable taxa
the axes. of the A. hirsuta group which, furthermore, appear
to hybridize readily with a number of other species
On open slopes and rocky banks, 2000-2200 m, of Anemia throughout the entire range. For ex-
Cajamarca, Amazonas. ample, there is a specimen of A. pastinacaria at
Peru; Bolivia; Argentina. Paris (San Martin, Tarapoto, Spruce 4134) which
Anemia myriophylla has a more highly divided also contains a very large, highly dissected leaf
leaf than any Anemia in Peru. This and the densely with abortive spores possibly a hybrid between
lanate lamina and petiole make it rather easy to A. hirsuta and A. phyllitidis.
recognize.
Huanuco: Muna, Bryan 427 (F, GH). In grass steppe,
Cajamarca: Between San Marcos and Cajabamba, Woytkowski 219 (uc). Cuzco: Quillabamba, Coronado
Correll & Smith P911 (GH, us). Prov. Celendin, Sagds- 124 (uc). Dept. Unknown: Mat hews 3299 (us).
legui et al. 8457 (F, HUT, MO, uc). Amazonas: of W
Chachapoyas on road to Gaelic. Hutchison & Bennett
4519 (F, GH, MO, NY, uc, us). Anemia
7. pastinacaria Prantl, Unters. Morph.
Gefasskrypt. 2: 110. 1881. LECTOTYPE
(designated by Lellinger in Proc. Biol. Soc.
6. Anemia hirsuta (L.) Sw., Syn. fil. 155. 1806. Wash. 98: 367. 1 985): Venezuela, "in convalli
del Tigre," Moritz 26 (B; probable isolecto-
Osmunda hirsuta L., Sp. pi. 1064. 1753. LECTO- type, GH!). Figure 6a-b.
TYPE (designated by Lellinger in Proc. Biol. Soc.
Wash. 98: 387. 1985): Plumier, Traite foug. Amer.,
/. 762, based on a Plumier collection from His-
This differs from Anemia hirsuta essentially only
paniola. in the following characters: Leaves to 45 cm long
Anemia hirsuta var. humboldtiana Hieron., Bot. Jahrb. and 7 cm broad. Sterile lamina gradually tapering
Syst. 34: 566. 1905. LECTOTYPE (designated by to a pinnatifid apex, but occasionally with a dis-
Lellinger in Proc. Biol. Soc. Wash. 98: 366. 1985):
Venezuela, border Edo. Sucre- Monagas, Hum-
tinct, nonconform apical segment; pinnae to 3.5(-5)
boldt 459 (B, Herb. Willd. 19495-2; photos, GH, cm long and 1 .5(-2) cm broad, margins denticulate
us). or crenate, or rarely with a few deep lobes on sev-
eral proximal pinnae.
Stem
short-creeping, densely provided with or-
ange trichomes. Leaves to 35 cm long and 5 cm On dry, rocky slopes or in forest clearings, 750-
broad, glabrate or somewhat villous on axes and 2000 m, San Martin, Loreto, Huanuco, Junin, Ay-
lamina, the erect, fertile pinnae borne at the base acucho, Cuzco.
of the sterile portion of the lamina; petiole (at least Mexico to Panama; Cuba; Jamaica; Trinidad;
on fertile leaves) 8-1 8 cm long. Sterile lamina pin- Surinam to Colombia, south to Bolivia and Brazil.
nate-pinnatifid to nearly 2-pinnate, with 7-1 2 pairs This and Anemia hirsuta differ only in their rel-
of pinnae, gradually tapering to a pinnatifid apex; ative size and depth of dissection of pinnae, and

28 FIELDIANA: BOTANY
a few specimens in Peru tend to be intermediate these characters are too variable and inconsistent
even in these characters. See A. hirsuta for further to be of real value. We
tentatively maintain it here
discussion. as a synonym, although we have not seen the type
to fully substantiate this.
San Martin: Tarapoto, Spruce 4 134 (p in part). Loreto:
Yanayacu, Biies 1037 (us). Huanuco: Yanano, Macbride Cuzco: Echarate, Camino de Sahuayacu, Bites 845 (us).
3816 (F, us). Junin: Prov. Huancayo, Pariahuanca, Tovar Ccochayoc, Biies 1712, 1719 (us). Prov. Convencion,
7948a (USM). Ayacucho: Aina, between Huanta and Rio Valle de Lucumayo-Pistipata, Vargas 4176 (MO, uc, us).
Apurimac, Killip & Smith 22614 (F, NY, us). Cuzco: Prov.
Convencion, Hacienda Sahuayaco, Vargas 1655 (GH),
1657 (GH, MO).
9. Anemia phyllitidis (L.) Sw., Syn. fil. 155. 1806.
Figure 6c.

8. Anemia oblongifolia (Cav.) Sw., Syn. fil. 156.


Osmunda 1064. 1753. TYPE:
phyllitidis L., Sp. pi.
1806. based on illustration of Hispaniola plant in Plu-
mier, Traite foug. Amer., /. 156. 1705.
Osmunda /. 592, / 2.
Anemia haenkei Presl, Reliq. haenk. 1: 74. 1825.
oblongifolia Cav., Icon. 6: 69,
TYPE: Panama, Nee TYPE: Peru, "in vallibus Cordillerarum Peru-
1801. (holotype, MA), ver-
ified by C. Christensen [Dansk. Bot. Ark. 9(3): viae," Haenke (holotype, PRC?).
31. 1937].
Osmunda humilis Cav., Icon. 6: 69, /. 592, f. 3. 1801. Stem decumbent (rarely short-creeping), rather
TYPE: Panama, Taboga Island,Nee(nol located);
densely clothed with orange or brownish tri-
earlier presumed at MA, but not seen by C. Chr.
chomes. Leaves to 80 cm long and 20 cm broad,
[Dansk. Bot. Ark. 9(3): 31. 1937].
Anemia humilis (Cav.) Sw., Syn. fil. 156. 1806. the erect fertile pinnae borne at the base of the
Anemia oblongifolia var. humilis (Cav.) Hooker & sterile portion of the lamina or less than 1 mm
Baker, Syn. fil. 431. 1868. below it;petiole (at least on fertile leaves) ca. 10-
35 cm long. Sterile lamina 1 -pinnate, with 3-7
Stem decumbent,rather densely provided with
pairs of pinnae and a conform apical one; pinnae
orange trichomes. Leaves to 30 cm long and 3.5 (larger ones) 5-14 cm long and 1.3-3 cm broad,
cm broad, sparsely to densely villous on axes and subequilateral, ovate to lanceolate, acute to acu-
lamina, the erect pinnae borne at the base
fertile
minate, the margins crenulate to dentate (or rarely
of the sterile portion of the lamina; petiole com- the basal pair lobed); veins copiously anastomos-
monly less than 6 cm long. Sterile lamina 1 -pin- ing. Fertile pinnae to 40 cm long, the stalk equaling
nate, with 2-9 pairs of pinnae, abruptly terminat- or (commonly) shorter than the panicle; the ulti-

ing in an obovate or obdeltoid apical segment which mate fertile segments essentially lacking laminar
is often as broad as long; pinnae to 1.8 cm long tissue.
and 0.7 cm
broad, strongly inequilateral at base,
truncate or rounded acroscopically and cuneate
Moist, often rocky slopes and ravine banks, in
basiscopically, broadly rounded at apex, the mar-
and at edges of forests, 700-1700 m, Amazonas
gins crenulate or subentire; veins free. Fertile pin- and Cuzco.
to Loreto, south to Huancavelica
nae to 1 8 cm long, the stalk 24
times the length
Mexico to Panama; Greater Antilles; Trinidad;
of the panicle, the ultimate fertile segments essen-
Venezuela and Colombia to Argentina and Uru-
tially lacking laminar tissue.
guay.
This is one of the most common species of Ane-
In forests, on rocky slopes, 1000-1 100 m, Cuz-
mia in Peru and, with its large, subentire pinnae
co.
and reticulate venation, is the easiest to recognize.
Mexico to Panama; Venezuela to Brazil and Bo-
livia.
Amazonas: Prov. Bagua, S of Bagua Grande on Rio
Anemiaoblongifolia is rather distinctive in its
Utcubamba, Hutchison 1479 (uc, us). San Martin: Ze-
coarse pinnae which are broadly rounded at their
pelacio, near Moyobamba, Klug 3440 (F, MO, NY, uc,
us). Loreto: Yanayacu, Biies 2005 (us). Huanuco:
tips and often nearly as broad as long. Sterile leaves Prov.
are commonly very short-petiolate (often subses- Huanuco, Puente Durand, Stork & Morton 9432 (F, GH,
MO, uc, us). Pasco: Oxapampa, Soukup 2361 (F, GH).
sile), and typically are so densely caespitose that
Junin: Yucapata, Woytkowski 6642 (GH, MO, us). Huan-
they give the appearance of a rosette. Some authors cavelica: Prov. Tayacaja, SE of Tintay, Tovar 46 11 (GH).
have separated A. humilis on the strength of its Ayacucho: Aina, between Huanta and Rio Apurimac,
villous sterile pinnae with crenulate margins; but Killip & Smith 22719 (NY, us). Cuzco: Prov.
La Con-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 29


vencion, upper valley of Rio Sanbray, Mexia 8056a, (F, 1-10 m long, vinelike, indeterminate, spreading
GH, uc, us). and scandent, partially dimorphic, the fertile por-
tions somewhat contracted with marginal fertile
Comments lobes, pinnate, glabrous to somewhat pubescent.
Pinnae short-stalked, pseudodichotomously
Anemia hispida Kunze, Linnaea 9: 20. 1834. branched with an arrested bud in the axil, each
TYPE: "In fruticetis ad Chibangata fl. Pe- primary pinna-branch radiately lobed or branched
ruv.." 1829, Poeppig diar. 1165 (LZ? de- or pinnate. Veins reticulate or (in Peruvian species)
stroyed, B?, w?). free. Sporangia borne separately on marginal lobes

of a pinna segment or on a wholly fertile segment,


Apparently known thus far only from the type each covered by a laminar outgrowth (flange).
collection. Spores tetrahedral-globose, trilete.
Kunze's plant was described as having: creeping
rhizome; pinnate, hispid leaves; fertile pinnae twice Lygodium is a natural group, easily distin-
as long as the sterile lamina; pinnae remote, pat- guished within the family by its long leaves and
ent, sessile, dimidiate, oblong, obtuse, with mar- vinelike and scrambling habit, often climbing to
gins crenate or subincised. He likened it to A nemia considerable heights by twining around the small-
1

repens Raddi (= A. hirsuta ?), but having pinnae er branches of shrubs and trees. It is widely dis-
less incised.Assuming it belongs to subg. Anemia tributed, primarily pantropic, with several extra-
(unsubstantiated by the description), it could be tropical species both in the New and Old World.
A. pastinacaria, but we cannot be certain without There are 30-35 species, but the number could be
examining the type. somewhat reduced by further study, as some ap-
pear to intergrade rather freely. Three species are
recognized in Peru.
II. Lygodium

Lygodium Sw., J. Bot. (Schrader) 1800(2): 106.


1 802, nom. consent TYPE: Lygodium scan- References
dens (L.) Sw. (Ophioglossum scandens L.).

Figure 7. DUEK, J. J. 1978. FeddesRepert., 89:411^23.


TRYON, R. M., AND A. F. TRYON. 1982. Lygo-
Plants terrestrial. Stem slender, short- to long- dium, pp. 69-76, in Ferns and allied plants,
creeping, provided with short trichomes. Leaves Springer- Verlag, New York.

Key to Species of Lygodium

a. Primary pinna-branch pinnate, the pinnules stalked, except sometimes near the apex of the primary
branch b
b. Pinnules (at least fertile ones) diminishing in size toward the apex of the primary branch, mostly

expanded at base into lobes, the lobes often discrete or even stalked 1 L. venustum .

b. Pinnules subequal, not expanded at base, subentire, very rarely some basal ones lobed at the base

2. L. volubile
a. Primary pinna-branch radiate, the ultimate segments deeply lobed, joined at the base
. 3. L. radiatum

1. Lygodium venustum Sw., J. Bot. (Schrader) lular trichomes. Primary pinna-branches pinnate
1801(2): 303. 1803. TYPE: Jakob Breyne, to (rarely) 2-pinnate, sessile to short-stalked,
Cent. I, t. 96. 1678, not "Brazil, Breynius" broadly to narrowly deltoid. Fertile pinnules 9-
(designated by Proctor, Flora Lesser Antilles 2 1 alternate and commonly widely spaced, all but
,

2: 51. 1977), a specimen which probably does the distal ones stalked, stalk often nodose at its
not exist. Figure 7a. juncture with pinnule base, sparsely to abundantly
pilose on axes, veins and (occasionally) the leaf
Pinna-stalks 2-10 mm moderately to
long, tissue, larger ones 2.5-5 cm long, gradually di-
densely pubescent with tawny to orange pluricel- minishing in size toward the apex of the primary

30 FIELDIANA: BOTANY
FIG. 7. Lygodium venustum: a, primary axis and pair of pinnae. Lygodium volubile: b, pinna; c, base of pinnule,
(a from Irwin et al. 21469, Brazil, F,b from Jeanpert s.n., Brazil, F, c from L. B. Smith 1352, Brazil, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 31


branch, most of them expanded at base into lobes, cm long, proximal and distal ones subequal,
the lobes rounded to cordate or subpalmate. Ster- broadly cuneate to truncate at base, only rarely
ile pinnules similar to the fertile, but often larger lobed at base. Sterile pinnules similar to the fertile,
and not much reduced toward the apex of the pri- but often somewhat larger. Veins free, several
mary branch. Veins free, 1 -several times-forked, times-forked, prominulous.
prominulous.
In wet, often inundated, lowland forests and open
Along roadsides and stream banks, in fields or fields, scandent on shrubs and low trees, often
along
in thickets and open forests, usually twining over stream and river banks, sea level to 250 m, Ama-
low shrubs, 100-1000 m, San Martin, Loreto, zonas, San Martin, Loreto, Madre de Dios.
Huanuco, Junin, Ucayali, Cuzco. Southern Mexico; Guatemala; Belize; Panama;
Tropical America. Greater Antilles; Trinidad; Venezuela to the
This species is widely distributed throughout the Guianas, south to Argentina and Brazil.
Neotropics. A number of other species have been This and Lygodium micans have been separated
confused with it or should be included within it. on the basis of several, variable, quantitative char-
For detailed discussions see Smith, Flora of Chia- acters, although these may appear more distinct

pas: Pteridophytes, Calif. Acad. Sci. p. 145. 1981, in certain regions outside South America: relative
and Stolze, Ferns & Fern Allies of Guatemala: Part size and shape of ultimate segments and their bas-
I, Fieldiana, Bot. 39: 38. 1976. The closely related es; degree of pubescence on abaxial surface; angle
Lygodium volubile readily can be distinguishedby of veins. None of these are consistently correlated
the fewer, larger, and subequal pinnules on each in the many Peruvian specimens examined.
pinna-branch; the distal ones are nearly or
i.e.,

quite as large as the proximal ones. Pinnules (at Amazonas: Quebrada Huampami, Kayap 1066 (MO).
least fertile ones) of L. venustum gradually dimin- San Martin: Prov. San Martin, Laguna Sauce, Ramirez
ish in size and shape so that subapical ones are
& Sotero 092-85 (F, GH, HUT). Loreto: Rio Tacsha Cur-
aray, Croat 20394 (F, GH, MO, NY). Gamitanacocha, Rio
often one-third to one-half the size of those near
Mazan, J. Schunke 234 (F, GH, NY, uc, us). Madre de
the pinna base. The Breyne plate was cited by Dios: El Pilar, Lopez & Soukup 4601 (GH).
Swartz and is clearly this species.

San Martin: Dist. San Martin, 2 km NW


of Tarapoto, 3. Lygodium radiatum Prantl, Unters. Morph.
Belshaw 3353 (F, GH, MO, NY, uc, us). Juan Jui, Alto Rio Gefasskrypt. 2: 66. 1881. TYPE: based on L.
Huallaga. Klug 4218 (F, GH, MO, NY, uc, us). Loreto: digitatum D. C. Eaton.
Puerto Arturo, lower Rio Huallaga below Yurimaguas,
Killip & Smith 27847 (NY, us). Huanuco: Prov. Leoncio Lygodium digitatum D. C. Eaton, Mem. Amer. Acad.
Prado, Huachipa, Plowman 5957 (F, GH). Junin: La Arts n.s. 2, 8: 217. 1860 (not Presl, 1825). LEC-
Merced, Killip & Smith 23379 (F, NY, us). Ucayali: Pu- TOTYPE (inferentially designated by Duek,
callpa (as Loreto), Soukup 3049 (F). Cuzco: Santa Ana, Feddes Repert. 89: 417. 1978): Panama, Gatun,
Cook & Gilbert 1614 (us). Hayes 25 (YU; isolectotype, us!).

Pinna-stalks reduced to short (0.5 mm) projec-


volubile Bot.
tions along the primary axis (or sometimes ob-
2. Lygodium Sw., J. (Schrader)
1801(2): 304. 1803. TYPE: Jamaica, Swartz solete).Primary pinna-branches long-stalked (1.5-
4 cm), the pinnae radiate (subpalmate), divided
(holotype, S-PA; photo, us). Figure 7b-c.
nearly to the base into (2)3-7 ultimate segments.
Fertile ultimate segments linear-lanceolate to nar-
Lygodium micansj. W. Sturm in Mart., Fl. bras. 1(2):
178. 1859. TYPE: "British Guiana," Schom- rowly oblanceolate, 5-15 cm long and 1.5-2 cm
burgk 399 (holotype, B). broad, the margins serrate, glabrous, or sparsely
pubescent along the midrib. Sterile segments sim-
Pinna-stalks 0.5-3 mm long, essentially gla- ilar to the fertile, but often somewhat longer (to
brous. Primary pinna-branches pinnate or (very 25 cm). Veins free, mostly 1- or 2-forked, prom-
rarely) 2-pinnate at base, the pinnae stalked (1.5- inulous.
5 cm), broadly oblong. Fertile pinnules 4-10, al-
ternateand widely spaced, conspicuously stalked, Rare, in wet forests, often along stream and river
nodose and often articulate at juncture with the banks, scandent on shrubs and small trees, 135-
pinnule base, glabrous to sparsely pilose on axes 850 m, Amazonas, Loreto, Pasco, Ucayali.
and veins (especially at bases of sporangia), 6-14 Panama; Colombia; Ecuador; Peru.

32 FIELDIANA: BOTANY
Amazonas: Al lado de Huampami, Kayap 1217 (MO, dichotoma (L.) Sm. (Acrostichum dichoto-
Santa Rosa, lower Rio Huallaga below Yu-
us). Loreto: mum L.). Figure 8.
rimaguas, Killip & Smith 28937 (F, GH, NY, us). Puerto
Melendez, below Pongo de Manseriche, Tessmann 4735
(NY). Pasco: Prov. Oxapampa, west side of Cordillera de Lophidium Rich., Actes Soc. Hist. Nat. Paris 1: 114.
San Matias, D. Smith 2011 (F, MO). Ucayali: Prov. Co- 1 792. TYPE: Lophidium latifolium Rich. = Schi-
ronel Portillo, km 89 Carretera Federico Basadre, C. zaea elegans (Vahl.) Sw.
Vdsquez 1 (USM). Actinostachys Wall., Numerical list of plants in East
Indies Company Museum, 1829, description
1.

from R. Br., Prod. 162. 1810. TYPE: Actinosta-


chys digitata (L.) Wall. (Acrostichum digitatum
Comments L.)
= Schizaea digitata (L.) Sw.

Lygodium oligostachyum (Willd.) Desv., Mem. Soc. Plants terrestrial. Stem erect or ascending, rath-
Linn. Paris 6: 205. 1827. er densely provided with septate trichomes and
with slender, fibrous roots. Leaves ca. 5-50 cm
ed. 4, 5:
Hydroglossum oligostachyum Willd., Sp. pi. long, caespitose, long-petiolate, glabrous or with
81. 1810. TYPE:
Plumier, Traite foug. Amer., t.
scattered, small trichomes, slightly to partially or
92. 1705 (based on specimens from Haiti, near
Lake Miragoan). wholly dimorphic, fertile ones with elongate fertile
segments borne pinnately or subdigitately at the
This is similar in general aspect to Lygodium apex of laminar axes which are simple or dichot-
venustum, but differs in the conspicuously 2-pin- omously branched. Lamina filiform, grasslike and

nate pinnae and the strongly flexuous (zigzag) and scarcely or not at all foliaceous, or fusiform to
flabelliform (in general outline) and foliaceous.
more delicate primary and secondary pinna rach-
Veins free. Sporangia crowded in or more rows 1
ises.Although L. oligostachyum has been consid-
ered by most authors to be confined to Hispaniola,
on the scarcely foliaceous segments (sporangio-
Duek (1978) assigned to it a broad distribution phores). Spores ellipsoidal, monolete.

(West Indies; parts of Central and South America)


and cited two specimens at Prague from Peru: Schizaea sometimes has been divided into sev-

Poeppigand Cuming, neither of which he had ex-


eral genera or subgenera on the basis of the more
amined. Under L. oligostachyum he also placed expanded, foliaceous lamina (Lophidium) or the
L. mexicanum Presl and two varieties of L. po- subdigitate (Actinostachys) versus pinnate (Schi-
zaea) fertile branches, as well as some differences
lymorphyum (Cav.) HBK. (illeg.); however, plants
usually identified under either name have usually
in the gametophytes. Nevertheless, the groups are
turned out to be L. venustum. We
have not seen quite closely allied and are here treated as a single

the Poeppig and Cuming collections in question, genus. About 30 species occur in tropical or ex-

but they are probably L. venustum and not L. oli- tratropical regions of both hemispheres, five of
which are found in Peru.
gostachyum as suggested. All specimens of the lat-
ter which we have examined in various herbaria
have been collected in Hispaniola, so it is unlikely
to be found in Peru.
Reference

TRYON, R. M., AND A. F. TRYON. 1982. Schi-

III. Schizaea zaea, pp. 76-83, in Ferns and allied plants,


Springer- Verlag, New York.

Schizaea Sm., Mem. Acad. Roy. Sci. (Turin) 5:

419. 1793, nom. conserv. TYPE: Schizaea

Key to Species of Schizaea

a. Leaves with lamina obviously foliaceous, flabelliform in general outline, the divisions linear-oblong
toobovate 1 S. elegans .

a. Leaves not or scarcely foliaceous, but filiform and grasslike or, if flabelliform in outline, then the
divisions filiform b
b. Sporangiophore pinnatifid, its segments bearing sporangia in a single row on each side of the vein
. .c

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 33


c. Leaves 0-2 times dichotomous, monomorphic; axis of the sporangiophore straight to slightly
curved at maturity d
d. Leaves 7-10 cm long, unbranched; sporangiophores with 5-7 pairs of ultimate segments
2. S. pusilla
d. Leaves (12-) 15-50 cm long, simple to once or twice acutely and dichotomously branched;
sporangiophores with (10-) 12-25 pairs of ultimate segments 3. S. incurvata

c. Leaves 3-6 times dichotomous, dimorphic; axis of the sporangiophore strongly curved at
maturity 4. S. poeppigiana
b. Sporangiophore subdigitate, its segments bearing sporangia in 2 or more (sometimes indistinct)
rows on each side of the vein 5. S. pennula

1. Schizaea elegans (Vahl) Sw., J. Bot. (Schrader) 2. Schizaea pusilla Pursh, Fl. Amer. sept. 2: 657.
1800(2): 103. 1 80 1 Figure
. 8. 1814. TYPE: United States, New Jersey, Bur-
lington Co., Quaker Bridge, Pursh? (not lo-
Acrostichum elegans Vahl, Symb. hot. 2: 104, t. 50. cated).
1791. TYPE: Trinidad, von Rohr (c).
Schizaea flabellum Mart., Icon. pi. crypt. 115, /. 55.
1834. TYPE: Brazil, Prov. Rio Negro, Martins Leaves unbranched, 7-10 cm long, linear, grass-
(holotype, BR; isotype, B; photo, us of B). with expanded lamina essentially lacking, pet-
like,
Lophidium elegans (Vahl) Presl, Suppl. tent, pterid.
77. 1845.
iole 0.40.5 mm broad, rather flattened, or sub-
Lophidium flabellum (Mart.) Presl, Suppl. tent, pterid. terete at base, sterile and fertile leaves erect and
77. 1845. ones apically
straight or slightly flexuous, fertile
bearing a pinnatifid, conduplicate, scarcely folia-
Leaves 20-80 cm long, petiole 15-65 cm long, ceous sporangiophore. Sporangiophore 0.5-0.7 cm
1-2.5 mm in diameter, subterete, obtusely angled long, with 5-7 pairs of linear segments,
abundantly
abaxially. Lamina subcoriaceous, essentially gla- provided on the margins and among the sporangia
brous, flabelliform (in general outline), 1 -several with deep orange to reddish brown, flexuous tri-
times-dichotomously forked or cleft, the divisions chomes. Sporangia in a single row of 5-7 on each
oblong to obovate, lateral margins entire, distal side of the vein.
ones strongly and sharply lacerate. Veins dichot-
omously forked within the ultimate divisions.
Sporangiophores borne along the distal margins
Thus far known only from Pasco, "plant colo-
of the lamina, pinnately branched at the tips of nizing sandy, wet landslide; shrubland on white
the marginal lacerations. Sporangia in a single row sandstone, 2700-2800 m."
on each side of the vein. United States (New Jersey, New York); Canada
(Newfoundland, Nova Scotia).
humus on forest floor, or in sandy or rocky
In Heretofore, Schizaea pusilla has been known to
soilon wooded slopes and ridges, 130-2200 m, occur only in a few scattered locations in north-
eastern North America. The recent collection in
Amazonas, San Martin, Loreto, Huanuco, Pasco,
Junin, Cuzco.
Pasco duplicates the northern plants in every fea-
Southern Mexico to Panama; Trinidad to Co- ture (including spores) except that the sterile leaves
are straight to slightly flexuous instead of con-
lombia, and south to Bolivia and Brazil.
Several varieties have been recognized, based spicuously curling. The great disjunction in dis-
tribution is at first startling until one understands
upon number and shape of laminar divisions,
characters which do not appear to merit distinc- the problems of encountering this very inconspic-

tion. uous fern. This and the other diminutive species


of Schizaea (like those of Ophioglossaceae and
Amazonas: Mendoza, Woytkowski8211 (GH, MO). San
Martin: Zepelacio, near Moyobamba, Klug 3415 (F, GH, Hymenophyllaceae) are very difficult to detect.
MO, NY, us). Loreto: Balsapuerto, Klug 2885 (F, GH, MO, Hence true distribution patterns may never come
NY, us). Huanuco: Prov. Huanuco, Tingo Maria, Tryon to light. It is most closely allied to S. incurvata,
& Tryon 5292 (BM, GH, NY, u, uc, us, USM). Pasco: Prov. under which see further discussion. Less closely
Oxapampa, Cordillera San Matias, Leon 324 (F, GH).
related is the Old World S.fistulosa Labill. and its
Junin: La Merced, Hacienda Schunke, Macbride 5601
(F, us). Cuzco: Valle de Urubamba, Machu Picchu, Her- variety australis (Gaud.) Fosb. from Chile and Ar-
rera 3299 (us). gentina, which differ in the lack of pubescence on

34 FIELDIANA: BOTANY
FIG. 8. Schizaea elegans: a, habit; b, sporangiophore; c, sporangiophore segment, abaxial side; d, sporangiophore
segment, adaxial side. (From Stolze, Ferns and fern allies of Guatemala, 1976.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 35


the sporangiophore and in the erose-lacerate mar- 4. Schizaea poeppigiana J.W. Sturm in Mart., Fl.
gins of the fertile ultimate segments. bras. 1(2): 181. 1859. SYNTYPES: Peru, "ad
Ventanilla de Cassapillo, 1829," Poeppig (w?);
Pasco: Oxapampa, Cordillera de Yanachaga, Cerro "British Guiana, in montibus Canuku," Rich.
Pajonal, Foster 9065 (F, GH, MO, us). Schomburgk 1189(ei).

Lophidium poeppigianum (J. W. Sturm) Underw., N.


3. Schizaea incurvata Schkuhr, 24 Kl. Linn. Pfl.- Amer. fl. 16: 38. 1909.
Syst. 1: 138, t. 137. 1809. TYPE: "British
Guiana, habitat in India Occidentali circa Es- Leaves dichotomous, 12-40 cm long, dimor-
sequebum," Gartner (not located). phic; petiole 10-30 cm long, ca. 1 mm
broad, con-
vex abaxially, flattened to concave adaxially,
Leaves simple to or 2-forked at an acute angle
1 -
sparsely to amply pilose. Sterile lamina circular
(branches subparallel), mature ones ( 1 2-) 1 5-50 cm in outline (often appearing conduplicate and fla-

long, linear, grasslike, with an expanded lamina belliform after pressing), 5-8 times dichotomously
essentially lacking, petiole 0.6-0.8 in diam- mm forked, the divisions linear, scarcely or slightly
convex abaxially, shallowly sulcate adaxially,
eter, foliaceous. Fertile lamina on somewhat longer pet-
somewhat flattened, or terete at base, sterile and ioles than the sterile, 3 or 4 times dichotomous,
fertile leaves straight or slightly flexuous, fertile the divisions linear, nonfoliaceous, bearing spo-
ones apically bearing a pinnatifid, conduplicate, rangiophores at their apices. Sporangiophores 1.5-
scarcely foliaceous sporangiophore. Sporangio- 2 cm long, with 12-25 pairs of linear segments,
phore 1.2-3 cm long, with (10-) 12-2 5 pairs of amply provided on the segment margins and among
linear segments, abundantly provided on the mar- the sporangia with pale to light brown, flexuous
gins and among the sporangia with pale to light trichomes. Sporangia in a single row of 1 5-25 on
brown, flexuous trichomes. Sporangia in a single each side of the vein.
row of (8-) 10- 18 on each side of the vein.
Thus far known in Peru only from one of the
Thus far known from a single collection (dept. syn types.
unknown) in Peru; elsewhere in open woods or Greater Antilles; Mexico (Chiapas); Costa Rica
clearings and savannas, in sand or sandy or rocky to Venezuela and Guyana; Peru; Bolivia.
soil, sea level to 400 m.
Surinam to Venezuela and northern Brazil;
Peru.
Of the many specimens seen by us in various 5. Schizaea pennula Sw., Syn. fil. 150, 379. 1806.
herbaria, only one was (possibly) collected in Peru: TYPE: "America meridionalis ... habitat in
Poeppig (NY). The label is worded simply "In Pe- America calidiore" (holotype, S-PA).
ruvia, diar. Kunze, 1834." However, Kunze (Lin-
naea9: 19. 1834) lists only one species of Schizaea Actinostachys pennula (Sw.) Hooker, Gen. fil., /. Ilia.
from Peru (41. 5". dichotomd), which is S. poep- 1842.

pigiana. Since there is no "Diar." number on


the label of the New York sheet, the data are Leaves ca. 1 2-50 cm long, unbranched, linear,
suspect. That the collecting site is correctly des- grasslike, with expanded lamina essentially lack-
ignated is also problematical, for the known range ing; petiole 1-1.8 mm broad, triquetrous and the
of the species is somewhat remote from Peru. faces sulcate, often flattened apically and subterete
However, as noted in the discussion of 5. pusilla at base, sterile and fertile leaves erectand straight
(q.v.),the grasslike appearance of several species or slightly flexuous, fertile ones apically bearing a
of Schizaea masks them well in any savanna-type subdigitate, scarcely foliaceous sporangiophore.
vegetation, so that more specimens may yet be Sporangiophore 1-4 cm long, with 6-14 linear,
discovered throughout the Neotropics after dili- strongly ascending segments, these essentially with
gent search, especially in Peruvian Amazonia in entire and glabrous margins, and with abundant,
sandy areas. Schizaea pusilla is similar to S. in- flexuous, reddish brown trichomes interspersed
curvata, both in general habit and pubescence of among the sporangia. Sporangia in 2 or more
the sporangiophore, but the two are easily sepa- (sometimes indistinct) crowded rows on each side
rated by the characters noted in the key. of the vein.

36 FIELDIANA: BOTANY
In sand and sandy soil of open forests and clear- pinnatisect (rarely pinnate) pectinate. Sporangia
ing, sea level to 260 m, San Martin, Loreto. borne in exindusiate sori on the abaxial surface of
Costa Rica; Puerto Rico; Guadeloupe; Trinidad; ultimate segments, with a short many-rowed stalk
Surinam to Colombia, south to Bolivia and Brazil. of cells and a central to oblique or nearly apical
annulus not interrupted by the stalk. Spores
San Martin: Chazuta, Rio Huallaga, Klug 3986 (F, GH, monolete or trilete, lacking chlorophyll, 1 20 to ca.
MO, NY, uc, us). Loreto: San Juan, vicinity of Iquitos, 800 in each sporangium.
Asplund 14414 (F, p, NY, us). Dist. Iquitos, Maynas, Que-
brada Shushuna, Rimachi 3995 (F, NY), 4870 (NY).
The pseudodichotomous branching of leaves and
the usually pectinate penultimate segments make
this one of the most distinctive of fern families.
These ferns frequently are found in dry, open areas
Family 5: GLEICHENIACEAE and species with larger leaves sometimes spread
over low shrubs to form dense thickets. Approx-
Gleicheniaceae (R. Br.) Presl, Reliq. haenk. 1 70. :

imately 1 20 species have been recognized by var-


1825, as Order GleicheniaeR. Br., Prod. 160. The
ious authors in as many as eight genera. fol-
1811, as Tribe of Filices. TYPE: Gleichenia
lowing treatment is based generally on the
Sm.
classification of Holttum (1957). Two genera are

recognized in Peru.
Terrestrial. Stem long-creeping, bearing tri-

chomes and/or scales. Leaves 20 cm long, erect,


to over 5 m
long and scandent or trailing, often References
forming dense thickets, circinate in vernation,
monomorphic, once or several times pseudodi- 1957. Florae Malesianae Prae-
HOLTTUM, R. E.
chotomously branched (rarely simple or with sev- cursores XVI. On the taxonomic subdivision of
eral 2-pinnate pinnae), indeterminate, with a per-
the Gleicheniaceae... .
Reinwardtia, 4: 257-280.
manently arrested bud at the fork of its branches, Amer.
MAXON, W. R. 1909. Gleicheniaceae, in N.
or the lamina partially pinnately branched, the
fl., 16: 53-63.
rachis (and sometimes pinna-rachis) with a pe-
NAKAI, T. 1950. A new classification of Glei-
riodically dormant bud between the last developed cheniales. Bull. Natl. Sci. Mus., 29: 1-71.
branches, a pair of accessory foliaceous segments M. 1907. American ferns VIII.
UNDERWOOD, L.
sometimes present at the base of otherwise usually
A preliminary review of the North American
naked axes, in addition to stipule-like segments
Gleicheniaceae. Bull. Torrey Bot. Club, 34: 243-
borne within the forks. Lamina bearing scales and/
262.
or trichomes. Veins free. Penultimate segments

Key to Genera of Gleicheniaceae

a. Axils and axillary buds with scales; veins simple to 1 -forked; sporangia 2-4(-5) per sorus
I. Gleichenia
a. Axils and axillary buds with trichomes; veins 2-4-forked; sporangia ca. 8-15 per sorus
II. Dicranopteris

I. Gleichenia 29: 47. 1950. Gleichenia section Diplopterygium


Diels, Nat. Pflanz. 1(4): 353. 1900. TYPE: Glei-
chenia glauca (Houtt.) Hooker (Polypodium
Gleichenia Sm., Mem. Acad. Roy. Sci. (Turin) 5:
glaucum Houtt.).
419. 1793. TYPE: G. polypodioides (L.) Sm.
(Onoclea polypodioides L.). Figure 9. Stem provided with setose or short-ciliate scales,
and sometimes stellate trichomes. Leaves once or
Sticherus Presl, Tent, pterid. 51. 1836. TYPE: S. lae-
several times pseudodichotomously branched (but
vigatus (Willd.) Presl (Mertensia laevigata Willd.)
= Gleichenia truncata (Willd.) Sprengel. simple in G. simplex, or with several 2-pinnate
Diplopterygium (Diels) Nakai, Bull. Natl. Sci. Mus. pinnae in G. bancroftii), with the penultimate seg-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 37


ments in diverging, subequal, usually pectinate 100 species. The three subgenera, Gleichenia, Di-
pairs. Axils of forks bearing a dense tuft of setose plopterygium, and Mertensia, are treated as genera
to ciliate (very rarely entire) scales, these often by some authors,
flanked by reduced, stipule-like appendages. Lam-
ina, at least abaxially, provided with setose to cil-
iate scales or stellate trichomes. Veins simple to Reference
1 -forked. Sori commonly paraphysate. Sporangia
2-4(-5) per sorus. CHING, R. C. 1940. On the genus Gleichenia
Smith. Sunyatsenia, 5(4): 269-288.
Gleichenia is a pantropical genus of more than

Key to Species of Gleichenia

a. Leaves simple (or very rarely with a single fork), deeply pinnatisect to pinnate; lamina linear
2. G. simplex

a. Leaves pseudodichotomously forked, or with 1-3 pairs of 2-pinnate pinnae; lamina broad, never
linear b
b. Pinnae 2-pinnate, not forked; axillary scales with margins entire 1 . G. bancroftii
b. Pinnae once to several times pseudodichotomous; axillary scales conspicuously ciliate or setose
(or entire in G. nitiduld) c
c. Ultimate segments moderately to densely tomentose on midrib and/or veins abaxially, the
whitish to orange tomentum often obscuring the segment tissue (sometimes thinning in age)
d
d. Larger penultimate segments 1.4-2.2 cm broad; larger ultimate segments 1.2 cm long or
less; pinnae usually 3-4 times-forked; branches strongly ascending, usually tightly crowded
and subparallel with adjacent ones 3. G. pennigera

d. Larger penultimate segments 2.5-8 cm broad; larger ultimate segments 1.5-4.5 cm long;
pinnae 1 - or 2-forked; branches spreading to moderately ascending, not crowded e
e. Sori inframedial, crowding or touching the segment midrib; ultimate segments (larger

ones) 1.8-3 mm broad beyond the dilated base, usually strongly revolute 4. G. bifida
e. Sori medial, rarely crowding the midrib; ultimate segments (larger ones) (3-)3.5 4.5 mm
broad, plane to moderately revolute 5. G. tomentosa

c. Ultimate segments naked to scaly abaxially, never tomentose (if trichomes present, these
scattered, short, or rigid) f
f. Penultimate segments less than 2 cm broad; ultimate segments 2-3.5(-4) times longer than
broad g
g. Scales ample abundant on costae and veins abaxially
to 6. G. revoluta
g. Scales essentially lacking on lamina h
h. Axillary scales broad, ciliate; primary axis conspicuously tuberculate to muricate . .

7. G. tuberculata

h. Axillary scales narrow and rigid, entire or (rarely) sparingly dentate or short-setose;
primary axis smooth 8. G. nitidula
f. Penultimate segments (at least larger ones) 2.2-8 cm broad; ultimate segments (4-)5-12
times longer than broad i

i. Midribs of ultimate segments naked 9. G. lechleri

i. Midribs of ultimate segments conspicuously scaly, at least abaxially j

j. Ultimate segments (most of them) remote, separated by a space once or twice their
width; sori medial to supramedial 10. G. remota

j. Ultimate segments approximate, contiguous at their bases; sori commonly inframedial


(or some of them medial) k
k. Costa scales commonly less than 1 mm
long; midrib scales deltoid, the scale body
5 or more cells wide; axillary scales 0.5-1.5 mm
long, with whitish or pale orange,
lax cilia 11. G. peruviana

38 FIELDIANA: BOTANY
k. Costa scales 1.5-3 mm long; midrib scales filiform, the scale body 1-3 cells wide;
axillary scales 2-3 mm long, with dark, rigid setae 1

1. Penultimate segments 4-9 cm broad; margins of ultimate segments plane to


slightly revolute; veins not or slightly raised 12. G. longipinnata
1. Penultimate segments 2.2-3.2 cm broad; margins of ultimate segments mod-
erately to strongly revolute; veins strongly raised abaxially . . 1 3. G. rubiginosa

1. Gleichenia bum-mini Hooker, Sp. til. 1: 5. 1844. long and less than 3 cm broad. Scales of the petiole
TYPE: Jamaica, Bancroft (holotype, K). essentially lacking, but abundant on the rachis
abaxially, these lanceolate, attenuate, orange to
Mertensia bancroftii (Hooker) Kunze, Linnaea 18: 307. reddish brown, the margins long-ciliate. Ultimate
1844.
segments contiguous at the very base or (especially
Dicranopteris bancroftii (Hooker) Underw., Bull. Tor-
proximal ones) discrete and almost their own width
rey Bot. Club 34: 252. 1907.
apart, strongly revolute, on the abaxial side slightly
Hicriopteris bancroftii (Hooker) Ching, Sunyatsenia 5:
278. 1940. to conspicuously pruinose and scaly, scales along
Diplopterygium bancroftii (Hooker) A. R. Smith, Amer. the midrib similar to those of the rachis, but to-
Fern J. 70: 26. 1980. ward the segment apex grading to pluricellular,
stellate trichomes. Sori medial to inframedial.
Leaves with 1-3 pairs of 2-pinnate pinnae, these
to 1.5 m
long and 40 cm broad, the axes naked or Road cuts, banks and open rocky slopes,
with scattered, linear, or filiform scales. Axillary to Cajamarca, south
(1900-)2700-3800 m, Piura
scales lanceolate to ovate, long-acuminate to at- to Huancavelica and Cuzco.
tenuate, yellowish to light brown, with entire mar- Colombia to Bolivia.
gins. Penultimate segments (pinnules) very nu- There has been some confusion as to publication
merous, crowded, at nearly right angles to the costa, of the name Mertensia simplex. In the Diet. Sci.
pectinate, cut nearly or quite to the costa. Ultimate Nat. 1 827 there is no text, but the plate with anal-
segments strongly revolute, midribs abaxially often ysis constitutes a valid publication. A description
provided with scattered, filiform scales. Sori in- was given at approximately the same time by Des-
framedial. vaux in Mem. Soc. Linn. Paris 6: 200. 1827.

In open forests, commonly on slopes or ravine Piura: Prov. Huancabamba, road to Canchaque,
banks, 2400-2800 m, infrequent in Peru: Huan- Hutchison 1617 GH, NY, uc). Lambayeque: Dist. In-
(F,

uco, Pasco, Cuzco. cahuasi, Laguna Tembladera, Sagdsteguiet al. 12774 (F,
Mexico to Panama; West HUT, MO). Cajamarca: Prov. San Miguel, El Tingo, Sa-
Indies; Colombia;
gdstegui et al. 9517 (F, MO, NY). Amazonas: Prov. Chach-
Venezuela; Ecuador; Peru; Bolivia.
apoyas, banks above swamp on summit of Cerros de
Calla-Calla, Wurdack 1212 (F, GH, NY, us). La Libertad:
Huanuco: Huanuco, within 5 km of Carpish, Tryon & Prov. Pataz, Huaylillas, Puerta del Monte, Lopez & Sa-
Tryon 5316 (F, OH, uc, us). Pasco: Border Prov. Oxa- gdstegui 3457 (GH). San Martin: Dist. Huallaga, Valley
pampa and Pasco, van der Werffet al. 8604 (MO). Cuzco: of Rio Apisoncho, Hamilton & Holligan 540 (us). An-
La Convencion, Valle de San Miguel, Biies 2126 (us). cash: Prov. Huari, Huascaran National Park, Quebrada
Pachachaca, D. Smith et al. 12564 (F, MO). Huanuco:
Yanano, stony clay bank, Macbride 4925 (F, GH, us).
2. Gleichenia simplex (Desv.) Hooker, Icon, pi., Huancavelica: Prov. Tayacaja, Chuspi-Tocas, between
/. 92. 1837.
Colcabamba and Paucarbamba, Tovar 2066 (GH, USM).
Cuzco: La Convencion, Biies 2078 (GH, us).

Mertensia simplex Desv., Dictionaire sciences natu-


relles 62(2), /. 91. 1827; Mem. Soc. Linn. Paris
6: 200. 1 827. TYPE: Peru, collector unknown (ho- 3. Gleichenia pennigera (Mart.) Moore, Index fil.

lotype, photo, us).


P; 381. 1862.
Dicranopteris simplex (Desv.) Maxon, Contr. U.S. Natl.
Herb. 24: 50. 1922.
Sticherus simplex (Desv.) Ching, Sunyatsenia 5: 285.
Mertensia pennigera Mart., Icon. pi. crypt. 130, t. 59,
1940. f. 1. 1834. TYPE: Brazil, "Prov. Minarum, in
Serra de S. Geraldo," Martius (M; photo, us).
Dicranopteris pennigera (Mart.) Maxon, Contr. U.S.
Leaves simple (very rarely with a single fork), Natl. Herb. 24: 48. 1922.
deeply pinnatisect to pinnate, linear, 2050 cm Sticherus penniger (Mart.) Copel., Gen. fil. 27. 1947.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 39


FIG. 9. Gleichenia bifida: a, habit; b, leaf axil with bud; c, ultimate segment, (a from Schunke V. 5976, F, b from
Tryon & Tryon 6605, Brazil, F, c from Luteyn & Luteyn 6735, Ecuador, F.)

40 FIELDIANA: BOTANY
Leaves pseudodichotomously forked. Pinnae Sticherus bifidus (Willd.) Ching, Sunyatsenia 5: 282.
1940.
(2-)3 4-forked, the branches (at least in Peruvian
Sticherus mathewsii (Hooker) Nakai, Bull. Nat. Sci.
specimens) strongly ascending, often tightly Mus. 29: 22. 1950.
crowded and subparallel with adjacent ones. Ax-
illary scales lanceolate, orange, the margins co- Leaves pseudodichotomously forked. Pinnae 1-
piously beset with long, tortuous pale or whitish or 2-forked, the branches spreading to moderately
cilia. Larger penultimate segments 1.4-2.2 cm
ascending. Axillary scales lanceolate to ovate, at-
broad, the costae abaxially provided with a mix-
tenuate, light brown to pale orange, the margins
ture of orange or whitish long-ciliate scales and a
amply ciliate. Larger penultimate segments (2.5-)3-
scattered whitish tomentum, adaxially naked or 7 cm broad, the costa on the abaxial side amply
sparsely to amply white-tomentose. Ultimate seg- to abundantly provided with orange, long-ciliate
ments slightly to strongly revolute, larger ones to
scales, on the adaxial naked or sometimes with
1.2 cm long, densely covered abaxially with whit-
sparse whitish tomentum. Ultimate segments
ish tomentum (often thinning in age). Sori infra-
moderately to (typically) strongly revolute, larger
medial, mostly crowding or touching the midrib. mm mm
ones 1 5-35 long and 1 .8-3 broad (beyond
the dilated base), densely covered abaxially with
Open places in forests, on slopes or stream banks,
orange to whitish tomentum (sometimes thinning
1000-2100 m, Loreto?, Huanuco, Pasco, Junin,
in age), the midribs also sparsely provided with
Cuzco.
tawny to whitish, filiform, ciliate scales. Sori in-
Colombia; Venezuela; Peru; Bolivia; Brazil.
framedial, mostly crowding or touching the mid-
Special note should be made of the branching rib.
pattern throughout the range of this species.
Branches may be very strongly ascending, thus
and clearings, often on banks
In forests, thickets
quite crowded and even subparallel with each oth- and 150-1800 m (very rarely to 2200 m)
slopes,
er; or they may diverge at broader angles, as seen
Piura to Loreto, south to Madre de Dios and Puno.
in the type. Thus far, all specimens examined from
Mexico to Panama; West Indies; Colombia and
Peru exhibit the crowded pattern, which might
Venezuela south to Bolivia and Paraguay.
tempt one to consider separation at subspecific
level. However, this difference appears uncorre- Piura: Prov. Huancabamba, slopes of Cerro La Viuda,
lated with other characters,and specimens, at least Sagdsteguiet al. 8 209 p.p. (NY). Antazonas: Prov. Bagua,
outside Peru, seem to grade freely from one pattern along roadside, 37 km NE of Chiriaco, Barbour 4486
to the other. (MO). San Martin: Zepelacio, near Moyobamba, Klug
3458 (F, GH, MO, NY, us). Loreto: Pumayacu, between

" Balsapuerto and Moyobamba, Klug 3242 (F, GH, MO, NY,
Loreto?: Altura de Salarayacu" (Sarayacu?), Biies 847
us). Huanuco: Dist. Churubamba, trail Cotirarda to
(us). Huanuco: Between Huanuco and Pampayacu, Ka-
Mercedes, Mexia 8191 (F, GH, MICH, MO, NY, uc, us).
nehira 153 (GH, us). Pasco: Prov. Oxapampa, Gran Pa-
Pasco: Prov. Oxapampa; Palcazii, Rio Alto Iscozacin,
jonal, D. Smith 5073 (F, MO). Junin: Concepcion, Her- Foster & d'Achille 10098 (F). Junin: Huacapistana, be-
rera 142 (us). Cuzco: Valle de Lares, Hda. Huy-huy, Biies
tween Tarma &
San Ramon, Ferreyra 286 (GH, USM).
1826 (us). Prov. Convention, Lucumayo, Vargas 4221
Ucayali: Vicinity of Aguaytia, on steep slopes along Rio
(us).
Aguaytia, Croat 21003 (F, MO, uc). Ayacucho: Ca. 25
km SW from Hacienda Luisiana and Rio Apurimac, ca.
25 km from Tambo, Dudley 11887 (GH). Cuzco: Prov.
4. Gleichenia bifida (Willd.) Sprengel, Syst. veg. Paucartambo, near Asuncion, West 7122 (MICH, uc).
Madre de Dios: Prov. Manii, Rio Inambari, Chavez 1057
4: 27. 1827. Figure 9.
(MO). Puno: San Gaban to Ollachea, Dillon et al. 1243
(F, MO, NY).
Mertensia bifida Willd., Kongl. Vetensk. Acad. Nya
Handl. 25: 168. 1804. TYPE: Venezuela, Czn-
cas,Bredemeyer(ho\olype,B,Herb. Willd. 19468;
photo, GH). 5. Gleichenia tomentosa (Sw.) Sprengel, Syst. veg.
Dicranopteris bifida (Willd.) Maxon, N. Amer. fl. 16:
4: 27. 1827.
60. 1909.
Gleichenia mathewsii Hooker, Sp. fil. 1 9. : 1 844. TYPE:
Peru, Mathews 1092, in part (holotype, K; photo, Mertensia tomentosa Sw., Kongl. Vetensk. Acad. Nya.
us). Handl. 25: 1 77, /. 5J. 4. 1 804. TYPE: Peru, Herb.
Mertensia mathewsii (Hooker) Fee, Mem. Foug. 1 1: Cavanilles (holotype, s; isotype, p?; photos, us of
122. 1866. S&P).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 41


Mertensia velata Kunze, Linnaea 9: 15. 1834. TYPE: midrib; also the plants seem to occur at lower
Peru, Huanuco, Pampayacu, July 1829, Poeppig elevations in Peru: 150-1800 m
(rarely to 2200
1117) (holotype, B?; isotype, M!; probable
(diar.
& m) as compared with 1 500-2700 in G. tomen-m
isotype, P!; frag., us!; photos, us of M p).
Gleichenia velata (Kunze) Mett., Fil. hort. bot. Lips. tosa. Color of scales on the buds and major axes
113. 1856. affords another, but less consistent, diagnostic fea-
Gleichenia buchtienii Christ & Rosenst., Repert. Spec. ture; in both species these are pale to deep orange,
Nov. Regni Veg. 5: 229. 1908. TYPE: Bolivia, but in G. tomentosa they are often liberally inter-
"Yungas australis, Sirupaya prope Yanacachi,"
Buchtien 496 (holotype, P!; isotypes, F!, P!, s).
mixed with castaneous ones. A few specimens have
Dicranopteris velata (Kunze) Maxon, Contr. U.S. Natl. been seen which are intermediate, thus emphasiz-
Herb. 24: 50. 1922. ing the close relationship of the two species. For
Sticherus buchtienii (Christ & Rosenst.) Copel., Gen. example, the only collection of G. tomentosa (cited
fil. 28. 1947.

Sticherus velatus (Kunze) Copel., Gen. fil. 28. 1947. below) from Cajamarca has segments strongly re-
volute and 3 mm
broad or less, but sori are medial
and scales frequently are castaneous at base.
Leaves pseudodichotomously forked. Pinnae 1
-

or 2-forked, the branches spreading to moderately Cajamarca: Prov. Cutervo, La Achira (San Andres-
ascending. Axillary scales lanceolate to ovate, at- Socota), growing over bushes,Lopez & Sagdstegui 5464
tenuate, deep to pale orange, or castaneous at base, (GH, HUT). Amazonas: Prov. Bagua, Cordillera Colan SE
of La Peca, Barbour 3980 (MO). San Martin: "In monte
the margins amply long-ciliate. Larger penultimate
Campana prope Tarapoto," Spruce 4707 (p, us). Huan-
segments 4-7 cm broad, the costa abundantly pro- uco: Playapampa, steep banks, Macbride 4509 (F, us).
vided abaxially with orange, long-ciliate scales, and Pasco: Cushi, trail to Tambo de Vaca, Bryan 687 (F).
adaxially with orange to whitish filiform scales Cuzco: La Convention, ca. 12 km NE from Hacienda
which often grade into a pale tomentum (this Luisiana and Rio Apurimac, Dudley 10545 (GH, us).

sometimes caducous). Ultimate segments plane to

moderately revolute, larger ones 2045 long mm


and (3-)3.5-4.5 mm
broad above the dilated base, 6. Gleichenia revoluta HBK., Nov. gen. sp. 1: 29.
densely covered abaxially with orange to whitish 1815. TYPE: Ecuador, Saraguru, Humboldt
tomentum (this sometimes thinning in age), the
& Bonpland (holotype, p; isotype, B!; frag., us
midribs provided also with orange, ciliate scales. of B; photos, us of B & p).

Sori mostly medial, rarely, if ever, crowding the


midrib. Mertensia revoluta (HBK.) Desv., Mem. Soc. Linn.
Paris 6: 200. 1827.
Mertensia pruinosa Mart., Icon. pi. crypt. 109. 1834.
In cloud or elfin forests, usually in open sites on TYPE: Brazil, Minas Gerais, Freyreiss (probable
holotype, M!; isotype, s; photos, F &
us of M).
ridges, slopes or ravine banks, 1 500-2700 m, Ca-
Gleichenia pruinosa (Mart.) Mett., Ann. Mus. Bot.
jamarca, Amazonas, San Martin, Huanuco, Pasco, Lugduno-Batavum 1: 49. 1863.
Cuzco. Gleichenia affinis Kuhn, Linnaea 36: 167. 1869. TYPE:
Colombia; Venezuela; Ecuador; Peru; Bolivia. Peru, Dept. Puno, San Gaban, Lechler 2265 (ho-
Much confusion has attended circumscription lotype, B!; photo, us).

of the densely tomentose species of Gleichenia, Dicranopteris affinis (Kuhn) Maxon, Contr. U.S. Natl.
Herb. 24: 47. 1922.
which include G. bifida as well as those combined Maxon, Contr. U.S.
Dicranopteris pruinosa (Mart.)
here under G. tomentosa. We have been unable to Natl. Herb. 24: 49. 1922.
obtain the type specimens of the latter, but type Sticherus pruinosus (Mart.) Ching, Sunyatsenia 5: 284.
1940.
photos (us) indicate clearly enough that ultimate
Sticherus revolutus (HBK.) Ching, Sunyatsenia 5: 285.
segments are too broad and soral lines appear too 1940.
farfrom midribs to be G. bifida. The photos, along Sticherus affinis (Kuhn) Nakai, Bull. Natl. Sci. Mus.
with Swartz's original description ( 1 804) of Mer- 29: 13. 1950.
tensia tomentosa and his subsequent Latin de-
scription (Syn. fil. 164, p. 392, 1806) make it ev- Leaves pseudodichotomously forked. Pinnae 2-
ident that G. tomentosa and G. velata are the same. 4-forked. Axillary scales lanceolate to ovate, lax,
All this confusion within the species complex is thin-textured, orange to reddish brown, the mar-
-
warranted, for G. bifida and G. tomentosa do not gins amply ciliate. Larger penultimate segments I

As seen in
differ strongly. the key, the former has 1.8(-2) cm broad, the costa on the abaxial side
narrower, more strongly revolute ultimate seg- amply to copiously provided with orange to red-
ments, and soral lines usually are closer to the dish brown, long-ciliate scales, on the adaxial side

42 FIELDIANA: BOTANY
the costa naked or filiform-scaly. Ultimate seg- Leaves pseudodichotomously forked, petiole and
ments moderately to strongly revolute, larger ones rachis conspicuously tuberculate to muricate. Pin-
2-3.5(-4) times longer than broad, the midrib and nae 2-3-forked. Axillary scales closely imbricate,
veins scaly as on the costa, those of the veins often ovate to lanceolate, castaneous to reddish brown,
filiform or grading to tortuous, stellate trichomes, the margins amply ciliate. Larger penultimate seg-
the veins not or scarcely raised. Sori medial to ments 1 .3-1.5 cm broad, pruinose abaxially, scales
supramedial. essentially lacking. Ultimate segments moderately
revolute, larger ones ca. 3 times longer than broad.
In elfin forests, clearings, and along roadsides, Sori supramedial.
1770-3400 m, Piura to Amazonas, south to Cuzco
and Puno. Thus far represented only by the type collection,
Costa Rica & Panama; Colombia; Venezuela; from Puno. This species appears to differ from
Ecuador; Peru; Brazil; Bolivia. Gleichenia revoluta only in the lack of scales on
This species is locally abundant in middle to the laminar surface and in the tuberculate to mur-
upper elevations and is sometimes confused with icate primary axis. However, the specimens were
Gleichenia bifida, probably because of the often taken from a plant apparently quite advanced in
copious indument borne on the abaxial surfaces. age, for few sporangia remain on the segments,
These scales, especially on the veins, often become and it is possible that any scales once borne on
filiform and even grade into stellate trichomes; the axes may have fallen away as well. Density of
however, the surfaces are never truly tomentose, indument varies within some species of Gleichen-
as in G. bifida. Also, penultimate segments of the ia, depending upon maturity. Furthermore, tu-
latter are muchbroader, with linear ultimate seg- berculate primary axes are not uncommon in the
ments, and the sori crowd the midrib, whereas in genus, although never so conspicuously so as in
G. revoluta sori are medial to supramedial, most G. tuberculata. The species is maintained as dis-
of them quite remote from the midrib. Further- tinct here, but with reservations.
more, G. bifida grows at lower elevations. It is
likely that G. revoluta occurs also in Costa Rica
and Panama [as G. costaricensis (Underw.) C. Chr.]
and in Bolivia [as G. boliviensis(Maxon & Morton) 8. Gleichenia nitidula Rosenst., Repert. Spec. Nov.
Lell.]. Probably G. subandina Sodiro and G. hy-
Regni Veg. 10: 275. 1912. TYPE: Costa Rica,
poleuca Sodiro of Ecuador are also synonymous San Carlos, Brade & Brade 503 (holotype, not
with G. revoluta.
located; isotypes, NY, s).

Huancabamba, above Huancabamba, road


Piura: Prov. Sticherus nitidulus (Rosenst.) Copel., Gen. fil. 28. 1 947.
to Canchaque, Hutchinson 1618 (F, OH, uc, us). Caja-
marca: Prov. Cutervo, alrededores de Cutervo, Ldpez &
Leaves pseudodichotomously forked, petiole and
Sagdstegui 5314 (GH, MO). Amazonas: Prov. Chacha-
poyas, slopes of Puma-urcu SE of Chachapoyas, Wur- rachis not or scarcely tuberculate. Pinnae 1-3-
dack 552 (F, GH, NY). La Libertad: Prov. Pataz, Puerta forked. Axillary scales not or slightly imbricate,
del Monte, Paso La Sabana, Lopez & Sagdstegui 3463
linear or linear-lanceolate, with subcaudate tips,
(GH). Pasco: Prov. Oxapampa, San Alberto, Cordillera
de Yanachaga, van der Werffet al. 8477 (MO). Huanuco: deep reddish brown, the margins entire or (rarely)

Playapampa, in sphagnum montana, Macbride 4508 (F, sparingly dentate or short-setose. Larger penulti-
GH, NY, us). Cuzco: Cordillera Vilcabamba, 23 km NE mate segments 1.4-1.8 cm broad, not or scarcely
from Hacienda Luisiana, Dudley 11133 (GH, us). Puno:
pruinose abaxially, scales lacking. Ultimate seg-
Prov. Sandia, near Limbani, Metcalf 30539 (MO, us;
ments slightly revolute, larger ones ca. 3 times
atypical in narrower segments and darker, narrower
scales). longer than broad.

Scandent on wet, shady banks, in forests, 1 600-


7. Gleichenia tuberculata Kuhn, Linnaea 36: 166. 1900 m, San Martin, Pasco.
1869. TYPE: Peru, Dept. Puno, Tatanara, Costa Rica; Panama; Ecuador, Peru.
Lechler2572 (holotype, B!; isotype, B!; photos,
The species is notable for its disjunct distribu-
F & MICH of B). tion, and has rarely been collected in South Amer-
ica. Distinctive characteristics are the lack of scales
Sticherus tuberculatus (Kuhn) Nakai, Bull. Natl. Sci. on the narrow penultimate segments, and the
Mus. 29: 30. 1950. sparse, rigid axillary scales with entire margins.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.


Apparently represented thus far in Peru by the follow- 10. Gleichenia remota (Kaulf.) Sprengel, Syst. veg.
ing two collections: San Martin: "Prope Tarapoto, in 4: 27. 1927.
monte Guayrapurima," Spruce 4018 (K). Pasco: (as Jun-
in) Pichis Trail, Enenas, dense forest, Killip & Smith
Mertensia remota Kaulf., Enum. fil. 39. 1824. TYPE:
25777 (F).
Brazil, Ilha de Sta. Catarina, Chamisso (holotype,
LZ destroyed; isotype, LE?).
Dicranopteris remota (Kaulf.) Maxon, Contr. U.S. Natl.
Herb. 24: 50. 1922.
9. Gleichenia lechleri Kuhn, Linnaea 36: 167.
1869. TYPE: Peru, "Tabina," Lechler 2040
(holotype, B!; isotypes, K, L; frag., us!; photos,
F, GH, us of B). Leaves pseudodichotomously forked, with fo-
liaceous appendages usually borne within each axil
Gleichenia yungensis Rosenst., Repert. Spec. Nov.
(especially proximally). Pinnae 1 or 2-forked. Ax-
-
Regni Veg. 5: 228. 1908. TYPE: Bolivia, Yungas,
illary scales lanceolate to ovate, attenuate, rigid,
Unduavi, 3300 m, Buchtien 902 (holotype, s; is-
otypes, uc!, us!).
1-1.5 mm long, often somewhat convex, casta-
Dicranopteris yungensis (Rosenst.) Maxon, Contr. U.S. neous to reddish brown, lustrous, not or slightly
Natl. Herb. 24: 50. 1922.
imbricate, their margins short-setose, or rarely en-
Sticherus yungensis (Rosenst.) Copel., Gen. fil. 28.
tire or with a few short cilia at base. Penultimate
1947.
Sticherus lechleri (Kuhn) Nakai, Bull. Natl. Sci. Mus. segments (larger ones) 4-9 cm broad, costae abax-
29: 21. 1950. ially provided with scattered castaneous or dark
reddish brown scales less than 1 mm
long, these
ovate to lanceolate, with setose to short-ciliate
Leaves pseudodichotomously forked, with fo-
margins, adaxially virtually naked (or occasionally
liaceous, often pinnatifid,appendages borne with-
with scattered patches of arachnoid scurf). Ulti-
in each axil (especially proximally). Pinnae 1-3-
mate segments remote, most of them separated by
forked. Axillary scales ovate to linear-lanceolate,
a space once or twice their width, slightly to strong-
orange to reddish brown, attenuate, the margins
not or slightly pruinose abaxially, larg-
ly revolute,
short-ciliate. Penultimate segments (larger ones)
2.5-6 cm broad, costae sparsely to amply scaly
er ones 20-50 mmlong and 1.7-2.5 mm
broad,
the midribs amply to copiously scaly on abaxial
abaxially, the scales orange, linear or linear-lan-
somewhat side, the scales deltoid, orange to tawny, with long,
ceolate, long-attenuate, the margins
on the margins, or toward segment tips
pale cilia
short-ciliate. Ultimate segments plane or slightly
sometimes becoming substellate. Veins often
revolute, commonly pruinose abaxially,larger ones
raised. Sori medial to supramedial.
12-28 mm long and 2.5-4.5 mm broad, the mid-
ribs lacking scales. Sori commonly supramedial.

At edges of forests, roadsides, and on rocky


In mountain forests, commonly on exposed banks, 350-2000 m, Amazonas, San Martin,
ridges, (1 1
00-) 1 800-3000 m, Huanuco, Cuzco. Huanuco.
Trinidad; Colombia; Ecuador; Peru; Bolivia. Costa Rica; Panama; Cuba; Trinidad; Colombia
This species is characterized by lack of midrib
to the Guianas; Ecuador; Peru; Bolivia; Brazil
scales, commonly pruinose lamina, plane or slight- (Amazonas, Ilha Sta. Catarina).
ly revolute ultimate segments, and the suprame- This and Gleichenia longipinnata are superfi-
dial sori. With it should be included Dicranopteris
cially similar in the great size of their penultimate
brittonii Maxon of Trinidad, and probably also G.
segments (sometimes to 50 cm long and 9 cm
leucocarpa Sod. of Ecuador. Although we have not
broad) and the extremely long and narrow ultimate
seen the type of the latter, a type fragment and
segments. However, besides the characters noted
photo (us) reveal no significant differences be- in the key, G. remota can also be sharply distin-
tween the two species.
guished by the laminar scales. Those within the
axils are relatively short (1-1.5 mm) and broad,
while the costal scales are even shorter (less than
Huanuco: Carpish, Coronado 78 (us). Cuzco: Machu and broader. In G. longipinnata, both the
Picchu, Valle del Urubamba, Herrera 3286 (us). Cerro
1 mm)
de Cusilluyoc, forest along Rio Pilahuata, Pennell 13940 axillary scales and costal scales are linear to fili-
(F, us). Prov. Paucartambo, "Pillawata," Yanamayo- form, the former 2-3 mm
long and the latter 1.5-
Tambomayo, Vargas 16702, 16705 (GH). 2 mm long.

44 FIELDIANA: BOTANY
Ama/onas: Prov. Bagua, roadside, 37 km NE of Chi- us
lotype, K; isotypes, BM, K; frag., us!; photo,
Barbour 4485 (MO), 4486 (USM). San Martin: Tar-
riaco,
ofK).
apoto-Yurimaguas Hwy., km 39, McDaniel 14209 (GH,
MO). Tarapoto, Carretera Tarapoto-Yurimaguas, Ri-
machi 5163, 5251 (MO, NY). Huanuco: Monzon, con- Mertensia longipinnata (Hooker) Klotzsch, Linnaea
fluencia con Huallaga, cerca de Tingo Maria, Ferreyra 18: 537. 1844.
10047 (GH). Dicranopteris longipinnata (Hooker) Maxon, Contr.
U.S. Nail. Herb. 24:48. 1922.
Sticherus longipinnatus (Hooker) Nakai, Bull. Nat. Sci.
Mus. 29: 21. 1950.
11. Gleichenia peruviana (Maxon) 1 ell.. Amer.
Fern J. 74: 57. 1984.
Leaves pseudodichotomously forked, occasion-
ally bearing foliaceous appendages within the ax-
Dicranopteris peruviana Maxon, Amer. Fern J. 33: ils. Pinnae commonly 1 -forked. Axillary scales
133. 1943. TYPE: Peru, Huanuco, Playapampa,
ca. 2700 m, Macbride 4510 (holotype, F!; isotype,
linear to filiform, 2-3 mm
long, lustrous reddish
us!). brown to deep orange, not or rarely tightly im-
bricate, the margins with dark, rigid, ascending
Leaves pseudodichotomously forked, with fo- setae. Penultimate segments (larger ones) 4-9 cm

liaceous appendages often borne within some of broad, costae on abaxial side amply to abundantly
the more proximal axils. Pinnae 1-3-forked. Ax- scaly, the scales 1.5-2 mm
long, lustrous reddish
illary scales ovate or lance-ovate, attenuate, 0.5- brown, setose and filiform or substellate, on adax-
1.5 mm long, castaneous to reddish brown, closely ialside often appressed filiform-scaly. Ultimate

imbricate, their margins amply ciliate, the cilia lax, segments approximate, contiguous at their bases
short and whitish to pale orange. Penultimate seg- (very rarely some proximal ones discrete and
ments (larger ones) 2.5^4(-5) cm broad, costae on somewhat apart), plane to slightly revolute, not or
abaxial side amply to copiously scaly, the scales ones 25-45
slightly pruinose, larger mm
long and
lustrous castaneous to reddish brown, ovate to lan- 2.5-3.5 mmbroad, abaxially bearing filiform or
ceolate, acuminate or attenuate, commonly less substellate scales on midrib and veins and often
than 1 mm long, their margins short-ciliate, costae some pale, delicate trichomes at the very margin.
on adaxial side sparsely to amply provided with Veins not or slightly prominulous. Sori mostly
tortuous trichomes or filiform scales. Ultimate seg- raised.
ments crowded, contiguous at their bases, slightly
to moderately revolute, not or slightly pruinose, on slopes and ridges, 300-1600 m
In forests,
larger ones 2-24 mm
long and 1 .5-2.5
1 broad mm (reported on Dudley 10448 label as also seen at
(beyond the dilated base), abaxially the midribs 3300 m, but probably mistaken for G. peruviana),
and (usually) veins minutely scaly, ciliate- or se- Amazonas, Loreto, Pasco, Huanuco, Cuzco.
tose-scaly, the scales often becoming filiform or Surinam; Peru; Brazil.
grading into stellate, whitish trichomes. Veins often Despite obvious differences, this and Gleichenia
raised. Sori mostly inframedial. rubiginosa share a distinctive type of laminar scale
not found in other species in Peru. The scales along
In thickets, open forests and ravines, often on the costae of penultimate divisions, abaxially, are
exposed rocky ridges, 2000-3400 m, La Libertad, typically long and narrow, the marginal cell walls
Huanuco, Pasco, Cuzco. deep reddish brown and bearing rigid dark setae.
Apparently confined to Peru, Cordilleras Cen- Along midrib and veins, these grade into filiform
tral and Oriental. scales often only two to three cells wide (the setae
then much longer than the width of the scale body)
La Libertad: Prov. Pataz, Pampa de Huayno-huincho, and thence into delicate, pluricellular trichomes.
Huaylillas, Ldpez & Sagdstegui 3517 (GH, HUT). Huan- The Costa Rican G. mellifera Christ also bears this
uco: Wet, dense jungle, 12 mi S of Panao, Macbride & kind of laminar scale, and although we have not
Featherstone 2217 Pasco: Prov. Oxapampa, Dist.
(F).
seen the type, it probably belongs here.
Oxapampa, Rio San Alberto, Foster et al. 10306 (F);
Leon 641 (F, GH); D. Smith & Pretel 7594 (F, MO). Cuzco:
Prov. La Convention, on open exposed ridge, Dudley Amazonas: Prov. Bagua, valley of Rio Maranon above
10710 (GH, MO). Cascadas de Mayasi, Wurdack 2049 (F, NY, uc, us).
Huanuco: SW slope of Rio Llullapichis watershed, Dud-
ley 13193 (GH). Ucayali: Prov. Coronel Portillo, Padre
12. Gleichenia longipinnata Hooker, Sp. til. 1: 9. Abad, Chacra de Cesar Vela, J. Schunke 5465, (F, GH,
1844. TYPE: Surinam, Hostmann 238 (ho- MO, NY, us). Pasco: Prov. Oxapampa, Valle de Palcazii,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 45


Foster 4504 (F). Cuzco: Prov. Convention, Cordillera Huancabamba, Dist. Sondor, subiendo
Piura: Prov.
Vilcabamba, Dudley 10448 (GH). alCerro La Viuda, Sagdstegui et al. 8209 p.p. (NY).
Huanuco: Prov. Huanuco, Carpish, between Huanuco
and Tingo Maria, Ferreyra 10009 (GH, USM). Huanuco-
Tingo Maria road, S of Chinchayo, Gentry et al. 19322
(F,MO). Pasco: Prov. Oxapampa, road between Oxapam-
13. Gleichenia rubiginosa Mett., Ann. Sci. Nat.
pa and Paucartambo, D. Smith 1592 (F, MO). Ucayali:
Bot. 5, 2: 267. 1864. TYPE: Colombia, Puente Prov. Coronel Portillo, Cordillera Azul, km 43 Tingo-
Nacional, 1650 m, Lindig 71 (holotype, B!; Maria-Pucallpa Road, Young & Sullivan 737 (F, MO).
P; frag., us!).
Cuzco: Valle San Miguel, La Convention, Bites 2061
isotype,
(us). Puno: Prov. Carabaya, San Gaban Carretera, Var-
gas 18877 (GH).
Gleichenia rubiginosa f. virescens Hieron., Bot. Jahrb.
Syst. 34: 561. 1905. SYNTYPE: Peru, Matthews
1092 (us).
Dicranopteris rubiginosa (Mett.) Maxon, Contr. U.S. II. Dicranopteris
Natl. Herb. 24: 50. 1922.
Sticherus rubiginosus (Mett.) Nakai, Bull. Nat. Sci.
Mus. 29: 28. 1950. Dicranopteris Bernh., Neues J. Bot. 1(2): 38.
("1806") 1805, nom. nov. for Mertensia Willd.
(not Roth), and with same type. Figure 10.
Leaves pseudodichotomously forked, rarely
bearing foliaceous appendages in the axils. Pinnae Mertensia Willd., Kongl. Vetensk. Acad. Nya Handl.
1- or 2-forked. Axillary scales linear or linear-
25:165.1 804, illeg. (not Roth 1 790). TYPE: Mer-
tensia dichotoma (Murray) Willd. (Polypodium
lanceolate, attenuate, 2-3 mm long, sublustrous, dichotomum Murray), Dicranopteris dichotoma
reddish brown, not or loosely imbricate, the mar- (Murray) Bernh. = Dicranopteris linearis (Burm.
gins with dark, rigid, ascending setae. Penultimate f.) Underw.
segments (larger ones) 2.2-3.2 cm broad, costae
on abaxial side copiously scaly, the scales 2-3 mm Stem provided with pluricellular trichomes,
long, deep orange, sublustrous, lanceolate or lin- scales lacking. Leaves 14 (rarely to 20) m long,
ear-lanceolate and attenuate, with setose margins, pseudodichotomously branched, with 1 -several
on adaxial side naked or occasionally sparsely ar- pairs of opposite pinnae which again typically
achnoid-scaly. Ultimate segments approximate, branch 1 or more times in opposite pairs, or with
contiguous at their bases, moderately to strongly several pinnae which branch unequally. Axils of
revolute, pruinose or not, larger ones 12-18 mm forks each bearing a dense tuft of trichomes and
long, (2-)2.5-3 mm broad, abaxially bearing fili- often a pair of reduced, stipule-like appendages.
form and setose to substellate scales on midrib Lamina glabrous, or pubescent with simple or stel-
and veins, and sometimes some delicate trichomes late trichomes, scales lacking. Veins 2 4-forked.
at the very margin. Veins raised abaxially. Sori Sori lacking paraphyses. Sporangia ca. 8-15 per
inframedial or occasionally medial. sorus.

In wet forests and thickets, 1 100-2850 m, Piura, This, like Gleichenia, is an essentially pantrop-
Huanuco, Pasco, Ucayali, Cuzco, Puno. ical genus. The two are readily distinguished by
Colombia; Venezuela; Ecuador, Peru. the character of their indument: that of Gleichenia
Although easily distinguished from G. longipin- being scales and trichomes, while scales are com-
nata apparently closely related to the
(q.v.), this is pletely lacking in Dicranopteris. Three species of
latter by virtue of the distinctive laminar scales. the latter are recognized in Peru.

Key to Species of Dicranopteris

a. Leaves with 1 -several


opposite pairs of stalked pinnae, which again branch 1 or more times; ultimate
segments contiguous at base; lamina abaxially glabrous or moderately pubescent with orange, stellate
trichomes (rarely tomentose) b
b. Pinna-forks branching in subequal pairs; accessory, mostly reduced, leafy segments usually borne
in pairs at the base of each fork (these in addition to stipule-like segments borne within the forks);

spores trilete 1 D. flexuosa .

b. Pinna-forks obviously branching unequally; accessory leafy segments lacking at the base of each

46 FIELDIANA: BOTANY
fork (not to be confused with stipule-like segments often borne within the forks); spores monolete
3. D. pectinate
a. Leaves 1 -forked, each branch consisting of a single, sessile, pinnate (to distally pinnatisect) pinna;
ultimate segments mostly well-spaced, not contiguous; lamina abaxially with dense, reddish brown
tomentum 2. D. nervosa .

1. Dicranopteris flexuosa (Schrader) Underw., Rio Chinchao, Macbride 5010 (F, NY, us). Ucayali: Prov.
Bull. Torrey Bot. Club 34: 254. 1907. Figure Coronel Portillo, Obeteni, E ridge of basin, Chrostowski
66-13 (uc). Cuzco: Prov. Convention, Choquallo, Var-
lOb-c.
gas 8153 (MO, uc). Puno: Prov. Sandia, Asalaya, Vargas
14835 (OH).
Mertensia flexuosa Schrader, Gott. Gel. Anz. 863.
1 824. TYPE: Brazil, Maximilian Prinz Neuwied

s.n. (holotype, LZ? destroyed; isotype, M?).


Mertensia rigida Kunze, Linnaea 9: 16. 1834. TYPE: 2. Dicranopteris nervosa (Kaulf.) Maxon, Contr.
Peru, "Chibangata," Poeppig 1153 (holotype, LZ U.S. Natl. Herb. 24: 49. 1922.
destroyed; isotype, p).
Gleichenia flexuosa (Schrader) Men., Ann. Mus. Bot.
Mertensia nervosa Kaulf., Enum. fil. 37. 1824. TYPE:
Lugduno-Batavum 1: 50. 1863.
Gleichenia rigida (Kunze) Bommer & Brazil, Sta. Catarina Is., Chamisso (holotype, not
Christ, Bull.
Soc. Roy. Bot. Belgique 35: 174. 1896, not G. located, LZ? destroyed; isotype, LE?, s?).
Sm. Gleichenia nervosa (Kaulf.) Sprengel, Syst. veg. 4: 25.
rigida J.
1827.
Leaves with 1-several opposite pairs of stalked
pinnae, these repeatedly pseudodichotomous, with Leaves 1 -forked, each branch consisting of a sin-
all subsequent branches forking in subequal pairs pinnate (to distally pinnatisect) pinna.
gle, sessile,

(i.e., each of the stalks subequal in length and di- Axils bearing a dense tuft of long, lax, reddish
verging at about the same angle), and a pair of brown trichomes and usually a pair of reduced
reduced, accessory, pectinate segments commonly stipule-like appendages.Pinnae (larger ones) 1 5-
produced at the base of each (but rarely the ulti- 40 long, 5-12 cm broad, the axes and laminar
cm
mate) fork. Axils bearing a tuft of stout, rigid, surfaces abaxially covered with a dense reddish
castaneous trichomes and a pair of reduced sti- brown tomentum (this sometimes gray to whitish
pule-like segments. Pinnae and their branches long- in age). Ultimate segments well-spaced, not con-
stalked, the axes and laminar surfaces glabrous, tiguous at base, to 60 mm
long and 3.5 mm
broad,
the forked, penultimate segments sessile, to 30 cm coriaceous, glaucous abaxially, the margins strong-
long and 6 cm broad. Ultimate segments contig- ly revolute. Spores trilete.

uous at base, to 30 mm
long and 4 broad,mm
coriaceous, glaucous abaxially, the margins strong- Thus far represented in Peru by a single collection:
Puno: Region of Rio Inambari, trail from Aricoma Pass
ly revolute. Spores trilete.
to Santo Domingo, 1800 m, McCarroll 125 (MICH).

On ridges and open slopes, at edges of forests,


750-2750 m, San Martin, Huanuco, Ucayali, Cuz- Venezuela?; Peru; Bolivia; Brazil.
This is linked to Dicranopteris schomburgkiana
co, Puno.
West Mexico to Panama, S to (Sturm) Morton and D. seminuda (KJotzsch) Max-
Indies; southern
and Paraguay (some isolated col-
on by the tomentose abaxial surfaces. The three
Brazil, Bolivia,
are usually separated by size of segments, color of
lections in the United States from coastal Ala-
tomentum, and other quantitative features; how-
bama).
ever, D. nervosa appears to be further distin-
This species is very closely related to Dicran-
Underw. of the Paleotrop- guished by the delicate, somewhat flexuous axil-
opteris linearis (Burm.)
ics. Suzanne Roth, who is working on a revision lary trichomes and the single pair of sessile pinnae.

of the Neotropical Gleicheniaceae, has suggested


The related species have either stout, rigid axillary
trichomes, or stalked, branching pinnae, or both.
(in litt.) that the two are probably conspecific. If
Whether these characters are sufficient to consti-
so, D. linearis has priority.
tute specific or subspecific distinction is open to

San Martin: NW
of San Martin, Rioja, Rio Negro, question. Further study of the species complex is
Soukup 5156 (GH). Huanuco: Vilcabamba, hacienda on needed.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 47


FIG. 10. Dicranopteris pectinata: a, primary axis and one pinna. Dicranopteris flexuosa: b, lamina, habit; c, axil
of primary fork, (a from Belshaw 3435, F, b-c from Britton 7186, Cuba, F.)

48 FIELDIANA: BOTANY
3. Dicranopteris pectinate (Willd). Underw., Bull. Tingo Maria to Pucallpa, Ridoutt s.n. (GH). Cuzco: Prov.
Torrey Hot. Club 34: 260. 1907. Figure lOa. Paucartambo, near Asuncion, West 7121 (MICH p.p., MO,
vcp.p.).

MertensiapectinataWMd., Kongl. Vetensk. Acad. Nya.


Handl. 25: 168. 1804. TYPE: Venezuela, Cara-
cas, Bredemeyer(hololype,B,Herb. Willd. 19465; Comments
photo, GH).
Gleichenia pectinata (Willd.) Presl, Reliq. haenk. 1 :

Dicranopteris flexuosa f. monstrosa (Hieron.) Na-


71. 1825.
kai, Bull. Natl. Sci. Mus. 29: 60. 1950.

Leaves with 1-several opposite pairs of stalked


Gleichenia flexuosa f. monstrosa Hieron. Hedwigia
pinnae, these each bearing several to many un- 48: 289. 1909. TYPE: Peru, Amazonas, Quebra-
equal branches which are again unequally branched da de Santa Lucia near Chachapoyas, Stiibel 1070
or which terminate in a pair of penultimate seg- (holotype, not located).
ments (i.e., each of the stalks unequal in length
and diverging at much different angles), lacking This supposed form of Dicranopteris flexuosa
accessory leafy segments at the base of the forks. was described from a plant which was rusty-to-
Axils bearing a tuft of orange to reddish brown, mentose on the juvenile segments of more distal
curved to flexuous trichomes and often a pair of pinnae. We have not seen evidence of tomentum
reduced stipule-like segments (at least within the on D. flexuosa, either on mature or juvenile plants,
proximal fork). Pinnae and their branches long- but it is possible this could represent a form of the
stalked, the axes commonly glabrous, the midribs species, or perhaps a hybrid. It is more likely that
and veins glabrous or, more often, sparsely to am- itbelongs to the complex of tomentose species
ply provided on the abaxial side with orange, stel- which are discussed above in the treatment of D.
late trichomes, the forked, penultimate segments nervosa, particularly D. seminuda of Venezuela,
sessile, to 23 cm long and 5 cm broad. Ultimate which has the general aspect of D. flexuosa. Another
segments contiguous at base, to 25 mm long and likely possibility is thatit may be a densely pu-

5 mm broad, chartaceous, glaucous abaxially, the bescent variant of D. pectinata (q.v.), although Hi-
margins scarcely to moderately revolute. Spores eronymus certainly should have mentioned the
monolete. distinctive branching pattern of the latter if this
were the case. Pending examination of the type, it

Forest clearings and thickets, along roadsides is fruitless to conjecture further.


and on open banks, 100-2000 m, Amazonas to
Loreto, south to Cuzco and Puno.
Degree of pubescence on abaxial surfaces varies
greatly in this species. In Central America and the
West Indies the lamina is commonly glabrous, or
with a few orange, stellate trichomes scattered along
Family 6: HYMENOPHYLLACEAE
the veins; but in South America, especially in Peru,
some pubescence is to be expectedvarying from Hymenophyllaceae Link, Handbuch 3: 36. 1833.
sparse to copious and on a few specimens we TYPE: Hymenophyllum Sm.
have examined it is so dense as to nearly obscure
the sori. It was perhaps one of these tomentose
Stem erect to decumbent, often slender and long-
plants on which Hieronymus based his Gleichenia short tri-
creeping, usually bearing scattered,
(Dicranopteris) flexuosa f. monstrosa (see discus- chomes. Leaves commonly circinate in vernation,
sion below under Comments). entire or pinnatifid to decompound, typically
monomorphic, or dimorphic in Trichom-
rarely
Amazonas: Prov. Bagua, along roadside from Chiriaco
anes, glabrous or pubescent, lamina very thin (often
to Puente Venezuela, Barbour 4488 (MO). San Martin:
1 cell thick), lacking stomates. Veins free, or in a
Prov. Lamas, Dist. Lamas, Belshaw 3435 (F, GH, MICH,
MO, NY, uc, us). Loreto: Mishuyacu, near Iquitos, Klug few species reticulate. Sporangia borne in margin-
850 (F, NY, us). Huanuco: Prov. Huanuco, Dist. Chu- al sori on a short to elongate receptacle, enclosed
rubamba, Pampa Hermosa, Mexia 8147 (F, GH, MICH, within a bivalvate or tubular indusium, with a
MO, NY, uc, us). Pasco: Prov. Oxapampa, road between
short stalk and an oblique annulus not interrupted
Puente Paucartambo and Oxapampa, D. Smith 1472 (F,
MO). Junin: La Merced, thickets, Killip & Smith 23804 by the stalk. Spores trilete, tetrahedral-globose to
(F, GH, NY, us). Ucayali: Prov. Coronel Portillo, road spheroidal, commonly with chlorophyll.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 49


The Hymenophyllaceae are a natural and dis- References
tinctive family, with representatives found in trop-
ical towet temperate regions nearly throughout COPELAND, E. B. 1938. Genera Hymenophylla-
the world.Most of the nearly 600 species have the cearum. Philipp. J. Sci., 67: 1-1 10.
lamina only one cell thick, which has earned them DIEM, J., AND J. LICHTENSTEIN. 1959. Las Hy-
the common name "filmy fern." The
filmy leaves menofilaceas del area Argentina-Chilena del sud.
and the marginal sori shaped like a cup or pouch Darwiniana, 11: 61 1-760.
make them readily recognizable as a family, but LELLINGER, D. B. 1984. Hymenophyllaceae, in
through the years there have been great differences Botany of the Guayana Highland Part X. Mem.
of opinion as to its subdivision. Various authors New York Bot. Gard., 38: 9-38.
have split it into as many as 42 genera, whereas MORTON, C. V. 1 968. The genera, subgenera and
but two are recognized here. sections of the Hymenophyllaceae. Contr. U.S.
The term marginate is used in this family to Natl. Herb., 38: 153-214.
describe an axis of the leaf that has a very narrow PRANTL, K. 1875. Die Hymenophyllaceen. Un-
band of green tissue on each side. Frequently this ters. Morph. Gefasskrypt., 1: 1-73.
band is scarcely raised beyond the surface of the PRESL, K. B. 1843. Hymenophyllaceae, pp. 1-
axis and is hardly discernible except for the color 70 (from Abh. Konigl. Bohm. Ges. Wiss., 5).
contrast with the axis. It is a rudimentary wing STOLZE, R. G. 1976. Hymenophyllaceae, Ferns
and represents the ultimate reduction of the con- and fern allies of Guatemala. Fieldiana, Bot.,
ditions elsewhere described in the text as "broadly 39: 51-90.
alate" or "narrowly alate." TRYON, R. M., AND A. F. TRYON. 1982. Hy-
The publications of Presl on the family have menophyllaceae, pp. 97-1 24, in Ferns and allied
been cited according to the first publication, with plants, Springer- Verlag, New York.
the date and page number of the separately issued
publication. Later publication with different pa-
gination in the Abh. Konigl. Bohm Ges. Wiss. is
not mentioned.

Key to Genera of Hymenophyllaceae

a. Indusium bivalvate, the valves '/2 or more the length of the indusium (rarely less); receptacle not or
and slightly exserted; venation anadromous; ultimate segments sometimes serrate, lacking false
rarely
veins I. Hymenophyllum
a. Indusium tubular and (and then the valves not more than '/a the
entire, or bilabiate, rarely bivalvate
length of the indusium; receptacle commonly exserted at maturity, often strongly so; venation anad-
romous or catadromous; ultimate segments not serrate, with false veins sometimes present
. . II. Trichomanes

I. Hymenophyllum
Hymenophyllum subg. Sphaerocionium (Presl) C. Chr.,
Index fil. Suppl. 3: 5. 1934.
Hymenophyilum Sm., Mem. Acad. Roy. Sci. (Tur- Mecodium(Cope\.)CopeL, Philipp. J. Sci. 67: 17. 1938.
in) 5: 418. 1793. LECTOTYPE (designated Hymenophyllum subg. Mecodium Copel., Philipp. J.
by Presl, Hymenophyllaceae 31. 1843): Hy- Sci. 64: 93. 1937. TYPE: Hymenophyllum po-
lyanthos (Sw.) Sw.
menophyllum tunbridgense (L.) Sm. Figure 11.

Plants epiphytic, occasionally epipetric or ter-


Leptocionium Presl, Hymenophyllaceae 26. 1843.
TYPE: Leptocionium dicranotrichum Presl = Hy- Stem filamentous, long-creeping, usually
restrial.

menophyllum dicranotrichum (Presl) Sadeb. bearing scattered trichomes and small, delicate
Hymenophyllum subg. Leptocionium (Presl) Christ, roots. Leaves monomorphic, petiolate, commonly
Farnkr. Erde 20. 1897.
1-20(-50) cm long or, in H. speciosum, to nearly
Sphaerocionium Presl, Hymenophyllaceae 33. 1843.
TYPE: Sphaerocionium hirsutum (L.) Presl (Tri- 2 m long. Lamina simple and entire (outside Peru)
chomanes hirsutum L.) = Hymenophyllum hir- or pinnatifid to decompound, glabrous or sparsely
sutum (L.) Sw. to densely pubescent. Ultimate segments entire to

50 FIELDIANA: BOTANY
FIG. 1
Hymenophyllum fucoides \ar.fucoides: a, habit; b, pinna with son. Hymenophyllum polyanthos: c, pinna
1.

with Hymenophyllum crispum: d, pinna with sori. (a from Seller 295, El Salvador, F, b from Plowman 6074, F,
sori.
c from Schunke 462, F, d from Wurdack 1510, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 51


serrate, bearing usually a single vein. Veins free section Ciliata) from those in which trichomes are
(reticulate in a few Old World species), anadro- borne also on the segment surface between the veins
mous, false veins absent. Sori terminal on the veins. and margins (his subsection Lanata). In a few
1 ml usiiim bivalvate, the valves Vz or more the length species trichomes between the veins may be
of the indusium (rarely less), the sporangia borne sparse present only in areas toward the base of
on an elongate, sometimes subglobose, receptacle. segments. Care must be exercised that these are
Receptacle not or rarely exserted beyond the mouth not overlooked. Conversely, there are several
of the indusium. species with stellate trichomes so dense on veins
and margins, and with stalks so long, that they
Nearly 300 species of Hymenophyllum are found may at first appear to spring from the intervening
throughout tropical regions of the world, although tissue.
a few occur in wet subtropical and temperate areas. Furthermore, certain groups of species in subg.
While most are epiphytes, several species have Hymenophyllum and Leptocionium are distin-
been reported from wet clay or rocky banks in guished by the presence of lamellae: accessory green
dense forests. A number of genera and/or subgen- wings or crests of tissue that emerge from the veins
era have been segregated by various authors, but or costae, but not on the same plane as the lamina.
in the following treatment Peruvian species are Typically these lamellae are made conspicuous by
aligned into three well-defined subgenera: Hy- abundance, but in H. loba-
their size, shape, or

menophyllum, Mecodium, and Leptocionium. A toalatum they are small and scattered, and in H.
good revision of the latter was done by Morton plumosum they are not at all foliose, but instead
( 1 947) under the name Sphaerocionium, but this are low and inconspicuous, and usually obscured
and others still require careful study, and some of by the dense, matted tomentum covering the en-
the species concepts are only provisionally treated tirelamina. In H. mirificum they could be mis-
below. Although there is no attempt here at a for- taken for trichomes, since many of them, although
mal subgeneric classification, the species are so four to six cells wide at the base, abruptly become
designated in the key and their descriptions follow uniseriate and grade to a filiform tip.
in natural order.
To facilitate use of the key to species, some ex-
planation of the diagnostic characters may be help- Reference
ful here, especially in subg. Leptocionium. Caution
is recommended when using the couplets that sep- MORTON, C. V. 1947. The American species of
arate those species having trichomes borne only Hymenophyllum sect. Sphaerocionium. Contr.
on veins and margins of segments (Morton's sub- U.S. Natl. Herb., 29: 139-201.

Key to Species of Hymenophyllum

a. Ultimate segments conspicuously serrate, glabrous, or rarely the serrations tipped by a short, pluri-
trichome [subg. Hymenophyllum]
cellular b
b. Costae and veins lacking lamellae c
c. Pinnae subequilateral at least beyond the base; sori (most of them) borne in the same plane as
the lamina, only occasionally arcuate; petiole usually more than 0.3 mm
in diameter
1 H. fucoides .

c. Pinnae dimidiate secondary segments all borne on acroscopic side); sori (most of them)
(i.e.,

strongly arcuate, thus arranged nearly perpendicular to the plane of the lamina; petiole less
than 0.3 mmin diameter 2. H. peltatum
b. Costae or veins (at least adaxially) bearing few to many lamellae not in the plane of the lamina,
these several to many cells wide at base, becoming uniseriate toward apex d
d. Petiole nonalate; rachis flexuous, nonalate, or slightly so to marginate distally; lamellae scattered
and inconspicuous 3. H. mirificum
d. Petiole broadly alate, often nearly to base; rachis straight, broadly alate throughout; lamellae
frequent and conspicuous 4. H. lamellatum
a. Ultimate segments entire, sometimes undulate to crispate, but never serrate, or rarely minutely
denticulate and then the teeth tipped by unicellular or stellate trichomes e

52 FIELDIANA: BOTANY
e. Lamina glabrous; ultimate segments with margins entire (although often crispate) [subg. Mecodium]
f
f. Leaves 1.5-4(-5) cm long; pinnae 3-6 pairs; rachis strongly flexuous to fractiflex
5. H. apiculatum
f. Leaves (mature ones) 6-30 cm long; pinnae 8-30 pairs; rachis straight or slightly (rarely strongly)
flexuous g
g. Mature indusia ovoid, or rhomboid, most of them somewhat to markedly longer
elliptical,
than broad, apex obtuse to acute, base narrowly to broadly cuneate (occasionally rounded)
and slightly to deeply immersed in the segment tissue; soriferous segment tips not or slightly
constricted, as broad as (or slightly narrower than) mature indusia h
h. Mature indusia broad- or narrow-ovoid, broadest toward the rounded or broadly cuneate
base, slightly immersed in the segment tissue; receptacle filiform to narrow-cylindrical,
short, not or rarely exserted i

i. Petiole marginate or alate near apex or throughout (wings sometimes partly deciduous);
rachis alate throughout; leaves commonly less than 15 cm long; growing at 100-
2000(-2400) m 6. H. polyanthos

i. Petiole neither alate nor marginate; rachis commonly nonalate at base, at least on one
side; leaves (mature ones) commonly 14-30 cm long; growing at 2100-3150 m . . . .

7. H. mathewsii

h. Mature indusia rhomboid, or conical, broadest at or beyond the middle, the base
elliptic,

narrow-cuneate, deeply (often halfway) immersed in the segment tissue; receptacle fili-
form, long, often exserted 8. H. trichomanoides

g. Mature indusia subglobose, circular in outline to broader than long, the apex rounded, the
base rounded or subtruncate, not or scarcely immersed in the segment tissue; soriferous
segment tips somewhat to greatly constricted, narrower than the mature indusia (usually so
much so that indusia appear pedicellate) j

j. Sporangia (5-)6-20 per sorus; petiole marginate or alate, at least distally (wing sometimes
deciduous); rachis alate throughout k
k. Indusium apex erose; 1 -several elongated pinnae intermixed among normal ones . .

9. H. ferax
k. Indusium apex entire; elongated pinnae rare or lacking among normal ones
10. H. myriocarpum
j. Sporangia l-4(-5) per sorus; petiole neither marginate nor alate; rachis commonly non-
alate toward base (or discontinuously alate on either side due to decurrent pinna bases)
1 1 . H. u lid u a
I 1 11 ii i

e. Lamina pubescent (at least on margins or veins), often densely so; ultimate segments entire or
rarely minutely denticulate [subg. Leptocionium} 1

1. Trichomes lacking on tissue between the veins m


m. Petiole slender, 0.2-0.3 in diameter mm n
n. Rachis nonalate at least in the proximal portion, or weakly and irregularly alate due to
the decurrent pinna bases; petiole nonalate, or weakly alate on one side due to the
decurrent pinna base o
o. Pinnae (at leastproximal ones) short-stalked, abundantly to densely pubescent on
veins and margins p
p. Costae regularly and conspicuously alate throughout on both sides; pinnae reg-
ularly diminishing in length toward lamina apex, larger ones with 4-9 pairs of
segments 12. H. molle

p. Costae partially nonalate, or discontinuously alate due to the decurrent bases of


segments; pinnae often irregular in length, some of them greatly elongate, normal
ones with 2-3(-4) pairs of segments 13. H. trichophyllum

o. Pinnae sessile or adnate, sparsely to moderately pubescent on veins and margins .

Q
q. Marginal trichomes mostly forked at base or 2-forked (a few simple or stellate),
petiole trichomes simple or forked; leaves determinate 14. H. elegans

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 53


q. Marginal and petiole trichomes mostly stellate; leaves indeterminate
1 5. H. adiantoides
n. Rachis regularly and conspicuously alate throughout; petiole alate on both sides, at least
at its apex r

r. Segments essentially plane; trichomes of segment margins mostly stellate or 2-forked


16. H. hirsutum
Segments undulate-crispate; trichomes of segment margins simple or forked from
r.

base 1 7. H. crispum
m. Petiole (of mature leaves) stout, 0.4-1 .5 in diameter mm s
s. Rachis conspicuously alate throughout; petiole usually alate just below lamina; trichomes
lacking on adaxial side of lamina t

t. Ultimate segments and rachis wing slightly to strongly undulate; stellate trichomes
lacking on petiole 1 8. H. valvatum

t. Ultimate segments and rachis wing not or scarcely undulate; stellate trichomes fre-
quent on petiole among the simple and forked ones 19. H. microcarpum
s. Rachis nonalate, at least in the proximal portion; petiole nonalate; trichomes present
on axes and veins both abaxially and adaxially 20. H. ruizianum
1. Trichomes (few to many) borne on the segment surface between the veins as well as on the
veins and margins u
u. Veins lacking lamellae v
v. Petiole slender, 0.2-0.3 mm
in diameter (or sometimes to 0.4 in H. amabile) mm w . . .

w. Pinnae or primary segments simple, entire; leaves to 7 cm long ...21. H. simplex


w. Pinnae or primary segments lobed to pinnatisect; leaves over 12 cm long (except
sometimes less in H. fragile) x
x. Lamina moderately to abundantly pubescent, but veins and tissue always clearly

visible y
y. Rachis broadly alate throughout; leaves determinate; petiole trichomes simple
to forked and often stellate 22. H. fragile
Rachis nonalate (sometimes alate distally); leaves indeterminate; petiole tri-
y.
chomes simple or forked, not or rarely stellate 23. H. elegantulum
x. Lamina densely tomentose, the veins and tissue often obscured

24. H. amabile
v. Petiole (of mature leaves) stout, 0.4-1.0 mm
in diameter z
z. Lamina densely tomentose, the veins and tissue mostly obscured by the tomentum;

rachis nonalate throughout aa


aa. Trichomes on rachis subsessile or short-stalked, very tightly appressed; larger

pinnae (2.5-)3-8 cm long, with 6-14 pairs of segments 25. H. speciosum


aa. Trichomes on rachis short- to long-stalked, spreading; larger pinnae 1-2.3 cm
long, with 4-7 pairs of segments 26. H. karstenianum
z. Lamina moderately to abundantly but surfaces not obscured by the trichomes;
pilose,
rachis commonly only occasionally nonalate
alate, at least distally, bb
bb. Petiole (8-)10-15 cm long, 0.7-1.2 mm
in diameter; lamina ovate to ovate-

lanceolate, 8-15 cm broad, not or slightly reduced at base .... 27. H. lindenii
bb. Petiole 2-7 cm long, 0.4-0.6(-0.7) mm in diameter; lamina linear, 3-8 cm broad,
and gradually reduced toward base
often strongly 28. H. plumieri
u. Veins bearing conspicuous lamellae (or, in H. plumosum, the outgrowths low and incon-
spicuous, these and the veins obscured by dense tomentum) cc
cc. Rachis nonalate nearly or wholly throughout dd
dd. Lamellae minute and inconspicuous, these and the veins obscured by dense to-
mentum 29. H. plumosum
dd. Lamellae broad and conspicuous, especially abaxially (if surface densely tomentose,
most lamellae still usually visible among the trichomes) ee
ee. Trichomes frequent, but not forming a dense cover; lamellae conspicuous abax-

ially, sometimes less so adaxially 30. H. multialatum

54 FIELDIANA: BOTANY
ee. Trichomes densely covering (sometimes obscuring) tissue and veins; lamellae
sometimes lacking adaxially 31. H. tomentosum
cc. Rachis alate nearly or wholly throughout ff

ff. Petiole rather stout, 0.4-0.7 mm in diameter; rachis strongly and regularly alate, at
least above the base; leaves 9-60 cm long; pinnae (larger ones) with 6-24 segments
gg
gg. Pinnae mostly narrow-triangular, narrowly acute to attenuate, larger ones 20-
45 mm long, with (6-)7-l 2 pairs of segments hh
hh. Lamellae on abaxial side mostly consisting of scattered flanges, about as
broad as long, lacking adaxially 32. H. lobatoalatum
hh. Lamellae on abaxial side abundant, conspicuous, elongate, often nearly the
length of the vein, sometimes less abundant and conspicuous adaxially . .

H. pyramidatum 33.

gg. Pinnae oblong to broadly triangular, obtuse or subacute, larger ones 6-20 mm
long, with 3-6(-8) pairs of segments 34. H. verecundum
ff. Petiole slender, 0. 1 0-0. 1 5 mm in diameter; rachis weakly and irregularly alate due
to the decurrent pinna bases; leaves less than 7 cm long; pinnae with 2-4 segments
35. H. tarapotense

1. Hymenophyllum fucoides (Sw.) Sw., J. Hot. (BM) of Hymenophyllum cristatum Hooker Grev. &
(Schrader) 1800(2): 99. 1802. This sheet contains two specimens of//, fucoides,
while pinned to it is an exact copy of/. 148, Icon,

Leaves 4-25(-30) cm long, 1 .5-8 cm broad. Pet- 1829, the illustration of//, cristatum, complete
fil.

iole 0.3-0.8 mm in diameter, nonalate or margin- with globose receptacle and conspicuous lamellae
ate to alate distally, glabrous to sparsely or mod- on the veins, which is an Ecuadorean species of a
erately pubescent with catenate trichomes. Lamina different section, Buesia (Morton) Morton. This
broadly or narrowly oblong, not or rather abruptly led to another error, for the subsequent illustration
reduced at base, 3-4-pinnatisect. Rachis broadly H. fucoides in Hooker's "Century of Ferns" (1854)
to narrowly alate throughout, or nonalate just at was evidently produced from this mixed sheet,
the base, sparsely to moderately provided with thus misleading later investigators about some of
flexuous, catenate trichomes. Pinnae 3-20(-25) the principal diagnostic features (for a full discus-
pairs, adnate, orproximal 1-2 pairs short-stalked, sion see Stolze, Amer. Fern J. 77: 137-140. 1987).
subequilateral except at base, there truncate ac- A second source of confusion has been the high-
roscopically, cuneate basiscopically, with 2-8 pairs ly variable nature of the species, which has
of pinnatisect to 2-pinnatisect segments. Sori 1-8 prompted the recognition of a number of related
per pinna, not or scarcely immersed in the segment taxa at various levels over the years. Morpholog-
tissue, most of them borne in the same plane as ical features are rather constant in the West Indian
the lamina, not or only occasionally arcuate. In- specimens, where leaves are quite small and com-
dusia narrowly to broadly oblong or elliptic, the pact; but variability increases in Central America,
apex obtuse to subacute and entire to laciniate, where some leaves are larger and more elongate,
receptacle narrow-cylindrical or fusiform to ovoid, and apices of indusia are deeply dentate. Even
not exserted or occasionally slightly so. greater differences are found in South American
plants, especially in Peru, where variability is at
The species occurs in the West Indies; southern its peak. It is likely that part of this may be due to

Mexico to Panama; Venezuela and Colombia to the effects of ecological niches and altitudinal levels,
Brazil and Bolivia. but this kind of analysis must wait for a much-
This species has been misunderstood, both in needed revision of this part of the genus.
itsnomenclature and taxonomy, since it was first For purposes of this treatment //. fucoides has
recognized by Swartz in 1788. The source of the been segregated into four varieties, a key to which
greatest confusion is a supposed type collection is provided here.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 55


Key to Varieties

a. Rachis marginate to narrow-alate, or if broadly alate then alae essentially plane, ultimate segments
plane b
b. Apex of indusium entire, erose or denticulate c
c. Rachis narrow-alate, or sometimes nonalate at base; petiole nonalate, or occasionally marginate
near lamina base; basal pinnae often short-stalked; sori rarely more than 3 to a pinna ......
la. var. fucoides
c. Rachis broadly alate throughout, each wing broader than the rachis; petiole usually alate distally,
at least near lamina base; basal and other pinnae adnate; sori often 5-8 to a pinna
Ib. var. calodictyon
b. Apex of indusium spinulose-dentate to laciniate var. pedicellatum
Ic.

a. Rachis broadly alate its entire length, alae and/or ultimate segments often undulate and their margins
discontinuously conduplicate Id. var. chachapoyense

la. Hymenophyllum fucoides var. fucoides. Fig- Amazonas: Trail east of La Peca in Serrania de Bagua,
ure lla-b. Gentry et al. 23034 (F, MO, us). San Martin: Venceremos,
near Amazonas border, Gentry et al. 45402 (MO). Huan-
uco: Cani, NE of Mito, Bryan 206, 384 (F); Macbride
Trichomanes fucoides Sw., Prodr. 136. 1788. TYPE: 3397 (B, F, us). Cuzco: Cerro Chuyapi, Bues ASS (GH,
Jamaica, Swartz (holotype, s; isotypes, B, BM; us), A56 (us). Puno: Sandia, Weberbauer 713 (B).
phOtOS, F & GH Of B, US of BM).
Leptocionium fucoides (Sw.) Presl, Hymenophyllaceae
27. 1843.
Ib. Hymenophyllum fucoides var. calodictyon
Meringium fucoides (Sw.) Copel., Philipp. J. Sci. 67:
45. 1938. (Bosch) Stolze, comb. & stat. nov.

Petiole (0.3-)0.4 0.8 mm in diameter, nonalate, Hymenophyllum calodictyon Bosch, Ned. Kruidk.
Arch. 5(3): 172. 1863. TYPE: Peru, Ruiz 84 (ho-
occasionally marginate near apex. Rachis narrow-
lotype, B!).
alate or marginate, at least distally, alae plane.
Pinnae adnate, or 1-2 proximal pairs short-stalked,
ultimate segments plane. Sori l-3(-4) pairs to a
Petiole 0.5-0.8 mm in diameter, conspicuously
alate to marginate distally. Rachis broadly alate
pinna, apex of indusia entire to denticulate, rarely
throughout, each of the alae plane and much wider
dentate.
than the rachis. Pinnae all adnate, with ultimate
segments plane. Sori ( l-)2-8 pairs to a pinna, apex
On
trunks and branches of trees in wet forest,
of indusia entire.
rarely on wet rock walls, 1 700-3300 m, Amazonas,
San Martin, Huanuco to Puno.
On trees in wet forests, Cajamarca and Ama-
West Indies; southern Mexico to Panama; Ven-
zonas south to Cuzco, 2100-3300 m.
ezuela and Colombia to Brazil and Bolivia.
Ecuador; Peru.
The most diminutive of these ferns scarcely dif-
In addition to the characters mentioned in the
fer from H. tunbridgense (L.) J. Sm., the type of
key, var. calodictyon commonly may be distin-
the genus, which is said to differ in its smaller
guished from the other varieties by the broader
leaves and in its strongly arched sori. The latter
segments, membranaceous tissue, and the lighter,
species supposedly occurs in the West Indies and
yellow-green color.
in Europe and Africa, but it may be only another

component in this complex. Some authors place


Cajamarca: Prov. Cutervo, Dist. San Andres, Agua
it, with H. peltatum and a few others, in sect. Diaz et al. 5340 (USM). Amazonas: Prov. Bagua,
Fria, A.
Hymenophyllum, distinguished principally by the Cordillera Colan SE of La Peca, Barbour 3586 (F, MO,
strongly arched sori, which are borne nearly per- USM). Prov. Bongara, near Pomacocha, Wurdack 865 (F,
GH, uc, us). Huanuco: Prov. Huanuco, Carpish, Plow-
pendicular to the plane of the lamina. In Old World
man 6074 (F, GH, us). Pasco: Prov. Oxapampa, E of Abra
specimens especially, this character is inconstant
Cantarizu, Skog et al. 5108 (us). Cuzco: Prov. Paucar-
and is but one of the problems which needs closer
tambo, Dist. Marcachea, near Achirani, Vargas 11142
scrutiny when the group is revised. (F, uc, us).

56 FIELDIANA: BOTANY
Ic. Hymenophyllum fucoides var. pedicellatum In undulate to crispate rachis wings and ulti-
its

(Klotzsch) Hieron., Bot. Jahrb. Syst. 34: 435. mate segments,var. chachapoyense resembles Hy-
1904. menophyllum tortuosum Hooker & Grev. of Chile;
however, alae and segments of the latter are much
Hymenophyllum pedicellatum Klotzsch, Linnaea 20: more strongly crispate and are setose-serrate as
439. 1847. TYPE: Venezuela, Merida, Moritz 346 well.
(holotype, B; isotypes, BR, us in part!).
Leptocionium pedicellatum (Klotzsch) Fourn., Bull.
Soc. Bot. France 19: 249. 1872.
Amazonas: Prov. Chachapoyas, Cerros de Calla Calla,
26 km above Leimebamba on road to Balsas, Hutchison
& Wright 6987 in part (F, GH, uc). Huanuco: Tambo de
Petiole 0.3-0.5 mm in diameter, nonalate. Vaca, Macbride 4460 (F, us).
Rachis broadly to narrowly alate, alae plane. Pin-
nae adnate, or basal pair short-stalked, ultimate
segments plane. Sori 1-3 pairs to a pinna, apex of
indusia laciniate to spinulose-dentate.
2. Hymenophyllum peltatum (Poir.) Desv., Mem.
Soc. Linn. Paris 6: 333. 1827.

On trees in wet forests, 500-2400 m, Amazonas.


Trichomanes peltatum Poir. in Lam., Encycl. 8: 76.
Venezuela; Colombia; Ecuador; Peru. TYPE:
1808. Mauritius, "Ile-de-France, Bory de
This may be synonymous with H. peruvianum, Saint-Vincent (V.s. in Herb, du Petit-Thouars)"
which see under Comments. It is likely that //. (not located).
ectocarpon Fee, of Central America and the Lesser
Antilles, should be included here also.
Leaves 2.5-8 cm long, 1-2 cm broad. Petiole
0.15-0.3 mm in diameter, nonalate, glabrous.
Amazonas: Prov. Bagua, Cordillera Colan, SE of La
Lamina narrowly oblong to lanceolate, slightly re-
Peca, Barbour 3598 (F, MO, USM), 3976 (F, MO). Prov.
duced at base, 2-3-pinnatisect. Rachis nonalate,
Bagua, Montenegro-Chiriaco, Sagdstegui 5931 (GH).
occasionally marginate distally, essentially gla-
brous. Pinnae 4-10 pairs, adnate, or 1-2 proximal
pairs short-stalked, dimidiate (i.e., the 1-4 sec-
Id. Hymenophyllum fucoides var. chachapoyense ondary segments all bome on acroscopic side). Sori
Stolze, var. nov. 1 or 2 per pinna, not immersed in the segment
tissue,most of them strongly arcuate, thus ar-
Differt a var. fucoide characteribus sequentibus: alae ranged nearly perpendicular to the plane of the
rhachidis latae et saepe undulatae; segmenta ultima saepe lamina. Indusia subspheroid to ovoid, the apex
undulata.
obtuse, entire, receptacle narrow-cylindrical, not
or rarely exserted.
Petiole 0.5-0.9 mm in diameter, commonly alate
Rachis broadly alate
distally, at least at the apex.
throughout, the alae often undulate. Pinnae ad- On rocky slopes and cliffs, 2800-4700 m, Junin,
nate, the ultimate segments often undulate and Cuzco, Madre de Dios, Puno.
their margins discontinuously conduplicate. Sori Peru; Bolivia; Argentina; Chile; Europe; Africa.
commonly 3-10 to a pinna, apex of indusia entire. Diem and Lichtenstein (1959) reported that H.
peltatum is highly variable in southern South
TYPE Peru, Dept. Amazonas, Prov. Chacha- America. They recognized six varieties in Argen-
poyas, Cerros de Calla Calla, 18 km above Lei- tina and/or Chile and also consider that it is also
mebamba on road to Balsas, 3100 m, Hutchison present in one form or another in areas of the Old
& Wright 5659 (holotype, GH!; isotype, uc!). World. Apparently only var. peltatum occurs in
On trunks and branches of trees, in wet forests Peru. It is a much more delicate fern than H. fu-
and moist, wooded ravines, 2900-3700 m, Ama- coides and can be easily distinguished from the
zonas, Huanuco. latter by the strongly dimidiate pinnae. Most sori
Colombia (Dept. del Valle); Ecuador (Prov. Car- are rather conspicuously arched out of the plane
chi & Napo); Peru. of the lamina, a character which Morton (1968)
This is a robust variety of higher elevations, used to separate sect. Hymenophyllum from others
similar to var. calodictyon in its stout petioles and in subg. Hymenophyllum. This feature is not con-
larger leaves (to 25 cm long, occasionally longer). stant in some other species, notably H. tunbridg-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 57


ense (L.) Sm., and its value is questionable as a and apices of indusia entire to denticulate. Rach-

diagnostic character above the level of species. ises in H. cristatum are obviously alate through-
out, lamellae are numerous and conspicuous, and
Junin: Prov. Huancayo, Dist. Huancayo, Huaytapal- indusia are deeply dentate to laciniate at apex. For
lana. Sounders 1 167 (GH). Cuzco: "Ccarcco" (Ccorca?), further discussion about the confused identity of
Biies 1395 (us). ?Madre de Dios: Pinasniocj, "Panticalla
H. cristatum, see H. fucoides.
Pass" (Pantiacolla?), Cook & Gilbert 1868, 1870 (us).
Puno: Prov. Carabaya, Ayapata, Vargas 10705 (GH).
Cuzco: Prov. La Convencion, Vilcabamba, Biies 1592,
1593, 2104 (us). Prov. Paucartambo, Acanacu, Vargas
29 (GH); West 7144 (uc, us). Prov. Urubamba, Puyu-
3. Hymenophyllum mirificum Morton, Bot. Gaz. pata-Yuncapata, Vargas 2922 (MO, us).
(Crawfordsville) 93: 338. 1932. TYPE: Peru,
Cuzco, Prov. La Convencion, Vilcabamba,
Biies 1600 (holotype, us!; photo, F).
4. Hymenophyllum lamella turn Stolze, sp. nov.

Buesia mirifica (Morton) Copel., Philipp. J. Sci. 67:


48. 1938.
Folia 7-1 5 cm longa, 2.5-4.5(-6) cm lata. Petiolus 1 .5-
5 cmlongus, 0.35-0.7 mm
diametro, conspicue alatus.
Lamina 2-pinnatisecta vel 3-pinnatisecta, conspicue la-
Leaves 16-38 cm long, (5-) 7-1 4 cm broad. Pet- mellata. Rhachis recta, late alata, lamellis conspicuis in-
iole 4.5-7 cm long, (0.4 )0. 5-0. 8 mm in diameter, structa. Pinnae 9-13-jugatae, confertae vel imbricatae,

nonalate or, rarely, very narrowly so or marginate adnatae, l-2.5(-3) cm longae, costis et venis lamellatis
praeditae. Segmenta saepe ultima undulata. marginibus
at apex, sparsely provided with brownish, flex- serratis.Indusia sphaeroidea vel ovoidea, brevipedicel-
uous, catenate trichomes. Lamina broadly or nar- lata, integris, receptaculis subglobosis vel
marginibus
rowly oblong, not or abruptly reduced at base, ovoideis.

deeply 3-4-pinnatisect, sparsely and inconspic-


uously lamellate adaxially. Rachis strongly flex- Leaves 7-15 cm long, 2.5-4.5(-6) cm broad.
uous, nonalate, or weakly alate toward the apex, Petiole 1.5-5 cm long, 0.37-0.7 mm in diameter,
sparsely provided with flexuous, catenate tri- conspicuously alate well toward, or quite to, base,
chomes and (adaxially) also with widely scattered sparsely provided with brownish, flexuous, caten-
lamellae, these usually 46 cells wide at base, ate trichomes. Lamina oblong-lanceolate, not or
abruptly becoming uniseriate and often grading to scarcely reduced at base, 2-3-pinnatisect, con-
a sharp, filiform tip. Pinnae 1 0-20 pairs, proximal spicuously lamellate adaxially. Rachis straight,
ones subdistant and short-stalked, distal ones ap- broadly alate throughout, sparsely to amply pu-
proximate and adnate, costae flexuous and nar- bescent with flexuous, catenate trichomes and also
rowly alate, with 6-10 pairs of 2-pinnatisect sec- provided with conspicuous scalelike lamellae, these
ondary segments, ultimate segments long and several to many cells wide at base, long-acuminate
narrow, glabrous, plane, the margins serrate, la- and often grading to a sharp, filiform tip. Pinnae
mellae inconspicuous and often widely scattered 9-13 pairs, crowded to imbricate, adnate at base,
along veins adaxially. Sori several to many, es- l-2.5(-3) cm long, adaxially lamellate on costae
pecially on distal pinnae, not or scarcely immersed and veins as on the rachis, with 3-6 pairs of 2-
in thesegment tissue, lying essentially in the plane pinnatisect or (distally) pinnatisect segments, ul-
of the lamina. Indusia subspherical to ovoid, the timate segments glabrous, usually slightly to
margins subentire, receptacle subglobose, not strongly undulate, the margins serrate, often con-
exserted. duplicate. Sori few, commonly 1 per pinna on the
basal acroscopic segment, not immersed in the
Endemic. In damp ravines and on trees, in wet segment tissue, lying essentially in the plane of the
2800-3650 m, Cuzco.
forests, lamina. Indusia subspherical to ovoid, the margins
Thisclosely related to Hymenophyllum cris-
is entire, receptacle subglobose or ovoid, not or
tatum Hooker & Grev. of Ecuador, which has the slightly exserted.
same general aspect and habit. Both species have
elongate, pendent leaves with flexuous rachises, TYPE Peru, Huanuco, Prov. Huanuco, trail
subspherical to ovoid indusia, subglobose recep- from S entrance of Carpish tunnel to crest of ridge,
tacles, and lamellae on axes and veins. But H. Luteyn & Luteyn 5474 (holotype, us!; isotype, NY!).
mirificum has larger pinnae, rachis wings essen- Endemic. In wet forests, on tree trunks and
tially lacking, lamellae sparse and inconspicuous, branches, 2750-3000 m, Huanuco, Cuzco, Puno.

58 FIELDIANA: BOTANY
Some of the specimens cited below were earlier fewer pinnae, but the character of its sori is dif-
suspected of being new species. On the sheets of ferent. The narrowly cuneate base of the indusium
Bites A 17 & A20 (us) are Morton's penciled re- is deeply immersed in the segment tip, whereas in

marks, "sect. Buesia, n. sp.?"; and Lechler 2568 H. polyanthos the indusium base is broadly cu-
(B) is annotated by Mettenius, "//. latipes Mett." neate to obtuse and only slightly immersed in the
Apparently Mettenius intended to publish, but a tissue.

search of the literature has revealed nothing under


this name. In its small size, compact lamina, and San Martin: Prov. Mariscal Caceres, Dist. Tocache
conspicuously alate axes, this is the most distinc- Nuevo, Cerro Sinsin, Plowman & Schunke 11478 (F).
Loreto: Mishuyacu, near Iquitos, Klug 1518 (F, NY, us).
tive species in sect. Buesia.
Huanuco: Environs of Jingo Maria, Aguilar 304 (uc).
Tingo Maria (as San Martin), Allard 20741, 21485 (GH,
Cuzco: Alturas de Sicre, Biies 1562 (us). Valley of Rio us). Pasco: Puerto Bermudez (as Junin), Killip & Smith
Urubamba, Biies A- 17, A-20 (us). Prov. La Convention, 26415 (F, us). Cuzco: Valle de Lares, alturas del Rio
at Klaus' Folly, above Camp 5, Dudley 1 WOO (GH). Puno: Lachac, Biies 1813a (us).
Tatanara, Lechler 2568 (B).

6. Hymenophyllum polyanthos (Sw.) Sw., J. Bot.


5. Hymenophyllum apiculatum Kuhn, Linnaea 35:
(Schrader) 1800(2): 102. 1802. Figure lie.
391. 1868. TYPE: Venezuela, Edo. Aragua,
Colonia Tovar, Fendler 32 (holotype, B; iso-
Trichomanes polyanthos S\v., Prodr. 137. 1788. TYPE:
type, us). Jamaica, Swartz (holotype, s; isotypes, B, BM;
photOS, US Of S, F & GH Of B).
Hymenophyllum dendritis Rosenst., Repert. Spec. Nov. Mecodium polyanthos (Sw.) Copel., Philipp. J. Sci. 67:
Regni Veg. 6: 308. 1909. TYPE: Bolivia, San Car- 19. 1938.
los near Mapiri, Buchtien 1093 (holotype, B?; is- Mecodium mexiae Copel., Univ. Calif. Publ. Bot. 19:
otype, us!). 294, 48. 1941. TYPE: Peru, Huanuco, near
/.

Mecodium dendritis (Rosenst.) Copel., Philipp. J. Sci. confluence of Rio Huallaga, Mexia 8282 (holo-
67: 26. 1938. type, uc!; isotypes, B!, F!, MO!).
Mecodium apiculatum (Kuhn) Vareschi, Flora Ven- Hymenophyllum mexiae (Copel.) Morton, Contr. U.S.
ezuela, Caracas 1: 198. 1969. Natl. Herb. 38: 173. 1968.

Leaves 1.5-4(-5) cm long, 0.8-1.8 cm broad. Leaves (6-)8-15 cm long, 2-5(-6) cm broad.
Petiole 0.3-1 .5 cm long, 0. 1-0.25 mm in diameter, Petiole 0.5-5 cm long, 0.3-0.6 mm in diameter,

marginate to narrow-alate, glabrous or with a few alate near theapex or throughout, glabrous. Lam-
scattered trichomes, especially at very base. Lam- ina ovate or elliptic, scarcely reduced at base (or
ina ovate or deltoid-ovate, not or scarcely reduced proximal 1-2 pairs of pinnae slightly reduced), 3-
at base, 2-3-pinnatifid, glabrous. Rachis glabrous, 4-pinnatisect, glabrous. Rachis glabrous, straight
strongly flexuous to fractiflex, alate throughout. or slightly flexuous, alate throughout. Pinnae 8-
Pinnae 3-6 pairs, adnate, larger ones 0.5-1.2 cm 14 pairs, essentially adnate, but not strongly over-
long, bearing 14 pairs of secondary segments. Ul- lapping the rachis, larger ones 1 -4 cm long, bearing
timate segments entire, plane, or occasionally the 2-many secondary segments. Ultimate segments
margins involute or conduplicate. Sori 1-4 to a entire, plane or occasionally a few with condupli-
pinna, usually confined to apical portion of lam- cate segment margins, larger ones commonly more
ina. Indusia rhomboid or ovoid, apex entire and than 1 mm
broad, the free portion relatively short,
obtuse to subacute, with broadly cuneate base, usually about twice as long as broad. Sori 2-many
about halfway immersed in the segment tissue, per pinna. Indusia relatively small, usually about
receptacle filiform, not exserted, bearing 6-12 spo- as broad as the segment apex, ovoid, elliptic, or
rangia. lanceolate, broadest toward the base, apex obtuse
or subacute, the broadly cuneate base slightly im-
on branches and trunks of trees
In dense forests mersed in the segment tissue, margins entire, re-
and on fallen logs, 100-1 500(-2000) m, San Mar- ceptacle filiform to narrowly cylindrical or fusi-
tin, Loreto, Huanuco, Pasco, Cuzco. form, not exserted, bearing 6-15 sporangia.
Venezuela; Colombia; Peru; Bolivia.
This would appear to be simply a reduced form In dense forests, on fallen logs, and on trunks
of Hymenophyllum polyanthos; however, it not of trees and tree ferns, 1 00-2000(-2400) m, Ama-
only has smaller leaves, more delicate petioles, and zonas, Loreto, San Martin south to Puno.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 59


Pantropical. Amazonas: Prov. Bagua, 1 2 mi E of La Peca, Harbour
This one of the most widely distributed ferns
is
2466 (uc in part). San Martin: Ravine E of Tingo Maria,
Allard 21446 (GH, us). Prov. Mariscal Caceres, Tocache
of tropical regions around the world and with its
Nuevo, J. Schunke 5693 (F). Loreto: Sierra del Pongo,
many close relatives represents one of the more Mexia 6291a (GH, uc). Huanuco: Prov. Huanuco, Cerro
taxonomically difficult species complexes. Com- Cucharas, Woytkowski 1152(cH). Prov. Huamalies, Rio
ponents of the group each have only a few distin- Monzon, 20 km W
of Tingo Maria, Tryon & Tryon 5299
(GH). Pasco: Prov. Oxapampa, Cordillera San Matias,
guishing characters and these are often inconstant.
Leon 317 (F, GH, USM). Pichis Trail, Enenas (as Junin),
Some of the features seem to vary more or less in
Killip & Smith 25651 (F, NY, us). Junin: La Merced, E
different geographic areas, so that species concepts of Quimiri Bridge, Killip & Smith 24022 (F, NY, us).
in each of these regions vary proportionately. The Madre de Dies: Prov. Manu, Cerro de Pantiacolla, Rio
Palotoa, Foster et al. 10683 (F). Cuzco: Prov. Paucartam-
group has long been misunderstood; consequently,
bo, Kosnipata, Vargas 17860 (GH). Puno: San Gaban,
in most herbaria, a number of specimens may be
Lechler 3327 (B).
filed incorrectly under Hymenophyllum polyan-
thos.
For purposes of this treatment, H. polyanthos
includes H. mexiae, which differs only in its small- 7. Hymenophyllum mathewsii Bosch, Ned.
er leaves. Leaves of H. apiculatum are also very Kruidk. Arch. 5: 162. 1863. SYNTYPES:
small, but this species (q.v). is characterized by "Peruvia," Mathews (B!; isosyntype, p; prob-
other, quantitative, features, and so should be con- able isosyntype, B!; photos, F, GH, & us of p).
sidered distinct. Hymenophyllum mathewsii may Ecuador, Quito, Cuming (B).
be merely a more luxuriant, higher elevation form
of H. polyanthos, with larger leaves and sori, and Leaves (6-) 1 4-30 cm long, 1.5-4 cm broad. Pet-
the axes less distinctly alate. Another species of iole 2.5-6(-8) cm long, 0.35-0.5 mm in diameter,
the subgenus occurring in Peru is H. myriocarpum, nonalate, glabrous. Lamina linear-lanceolate to
which differs principally in shape and character of oblanceolate, rather abruptly reduced or strongly
the indusia: in most specimens sori appear to be and gradually reduced at base, 2-3-pinnatisect,

pedicellate, for the apices of ultimate segments are glabrous. Rachis glabrous, alate throughout except
strongly constricted; furthermore, mature indusia usually nonalate at base, or there narrow-alate on
are subglobose, or often noticeably broader than only one side. Pinnae 14-18 pairs, adnate and often
long. This contrasts with the essentially ovoid in- overlapping the rachis, or basal ones short-stalked,
dusia of//, polyanthos et al., commonly somewhat larger ones (1.5-)2-3.5 cm long, bearing 4-8 pairs
longer than broad and slightly to strongly im- of secondary segments. Ultimate segments entire,
mersed in the segment tips, which are not or plane or occasionally undulate, or a few with mar-
scarcely constricted. While this character is prob- gins conduplicate. Sori 4-many per pinna. Indusia
ably the most reliable one in subg. Mecodium in relatively large, usually slightly broader than the
Peru, unfortunately even it is not 1 00% constant, segment apex, ovoid, apex obtuse to subacute, the
for on occasional specimens indusia may vary broadly cuneate base slightly immersed in the seg-
somewhat from one shape to another. Conse- ment tissue, margins entire, receptacle short and
quently, most species in the subgenus must be sep- filiform, bearing 6-12 sporangia.
arated by suites of characters. Ultimately, taxo-
nomic problems will be solved only by thorough Cloud forests, commonly on trunks and branch-
study of species groups throughout their entire es of trees, 2 1 00-3 1 50 m, Huanuco, Pasco, Cuzco.
range. Ecuador; Peru.
Two other characters are often helpful in sep- Hutchison 1751 (cited below) has delicate, di-
arating H. polyanthos (at least in Peru) from H. minutive leaves only 6-8 cm long; this depauper-
myriocarpum particularly useful where sterile ate condition probably results from the unusual
specimens are involved. Pinnae of H. myriocar- habitat, "in cave." This species is very closely re-
pum commonly have basal segments strongly Hymenophyllum polyanthos (q.v.) and may
lated to

overlapping the rachis. In H. polyanthos pinnae not merit distinction at the species level. Normal
are adnate and basal segments often crowd or touch specimens of//, mathewsii tend to have larger sori,
the rachis, but rarely do some of them overlap. in relation to the segment apex, and longer, pen-
Also, H. polyanthos occurs at lower elevations in dent leaves, with no trace of tissue along the petiole
Peru, 100-2000(-2400) m; whereas H. myriocar- and, commonly, none between the proximal two
pum is found at ( 1 700-)2000-4200 m. pinnae. Leaves of H. polyanthos are smaller and

60 FIELDIANA: BOTANY
more compact, erect or arching, with rachis dis- guished by the narrower ultimate segments. Larger
tinctly alate. The petiole is marginate or alate ones are commonly 0.6-0.9 mm
broad and linear.
throughout, or at least distally. Often the wing is Other species of subg. Mecodium have relatively
deciduous upon drying, but traces can usually be broader ultimate segments, most of them over 1
found by close scrutiny. mm broad, or if not, then only about twice as long
as broad. This is a rather reliable way to identify
Huanuco: Prov. Huanuco, Carpish, Plowman 6074A sterile specimens.
(F).Pasco: Prov. Oxapampa, San Alberto, Yanachaga, Hymenophyllum trichomanoides differs sharply
van der Werffel al. 8437 (MO). Cuzco: Huadquina, Bues
from all other species of subg. Mecodium in Peru
1346 (us). Prov. La Convention, Cochapata, Valle de
San Miguel, Bues 2176 (us). Prov. Urubamba, summit by the narrowly cuneate sori which are deeply im-
of Huayna Picchu, Hutchison 1751 (atypical) (F, uc). mersed in the segment tissue. Indusia are broadest
at or beyond the middle and, at least in Peru, are

long and narrow: either narrowly elliptic with sub-


acute apices, or conical, with rounded apices. All
8. Hymenophyllum trichomanoides Bosch, Ned. specimens seen from Peru clearly match the Spruce
Kruidk. Arch. 5(3): 158. 1863. SYNTYPES: syntype. Some specimens examined from Colom-
Peru, San Martin, Tarapoto, Spruce 4696 bia and Venezuela have shorter and broader in-

Colombia, dusia and much shorter petioles, and it is possible


(isosyntypes, GH!, NY!, P!, us!).
Moritz (BM?), Schomburgk(BM?). Ecuador, Pi- that these represent a variant. However, this has

Cuming (not not been substantiated by comparison with the


chincha, Quito, located). "Brit-
ish Guiana," Schomburgk (BM?). other syntypes from northern South America,
which we have not seen. With H. trichomanoides
Mecodium trichomanoides (Bosch) Pic.-Ser., Webbia probably should be included H. trianae Hieron.
28: 469. 1973. of Colombia. A syntype of the latter, Lehmann
7410 (us), matches Spruce 4696, except that re-
Leaves (9-)l 1-26 cm long, 4-6(-8) cm broad. ceptacles are not exserted.
Petiole 5-12 cm long, 0.4-0.8 cm in diameter,
marginate to alate distally (rarely nonalate), with Huanuco: Cerros del Sira, SW
slope of Rio Llullapichis
a few scattered trichomes. Lamina broadly or nar- watershed, Dudley 13375 (GH). Pasco: Prov. Oxapampa,
Palcazu Valley, Cabeza de Mono, D. Smith 3756 (F, MO).
rowly ovate (1 or 2 pairs of proximal pinnae com- Junin: Schunke Hacienda, above San Ramon, Killip &
monly somewhat reduced), 2-3-pinnatisect, gla- Smith 24844 (F, GH, NY, us). Cuzco: Sahuayacu, Bues
brous. Rachis glabrous, slightly flexuous, narrowly 804, 817 (us).
to broadly alate throughout, the wing margin
sometimes conduplicate. Pinnae 7-14 pairs, ad-
nate and overlapping the rachis, or 1-2 proximal
pairs short-stalked, larger ones 2.2-4 cm long, 9. Hymenophyllum ferax Bosch, Ned. Kruidk.
bearing 4-8 pairs of secondary segments. Ultimate Arch 4: 392. 1 859. TYPE: Venezuela, Merida,
segments entire, plane, or some of them with in- Funck & Schlim 1578 (holotype, L?; isotype,
volute margins, larger ones commonly less than 1 p; photo, us of P).
mm broad, the free portion linear, usually 2.5-4
times as long as broad. Sori 6-12 per pinna. In- Mecodium ferax (Bosch) Copel., Philipp. J. Sci. 67:
dusia relatively small, usually about as broad as 25. 1938.
the segment apex, rhomboid, elliptical or conical,
broadest at or beyond the middle, apex acute to Leaves 16-38 cm long, 4-8 cm broad. Petiole
obtuse, the sharply cuneate base deeply (often half- 2-9 cm long, 0.4-0.9 cm in diameter, alate toward
way) immersed in the segment tissue, margins en- apex (wings sometimes deciduous), essentially gla-
tire,receptacle long, filiform, often exserted, bear- brous. Lamina ovate to narrow-elliptic, scarcely
ing 6-10 sporangia. to somewhat reduced at base (1-several proximal
pairs of pinnae reduced), 3-pinnatisect, glabrous.
In dense, wet forests, on trunks of trees, 300- Rachis glabrous, straight or slightly flexuous, alate
3000 m, San Martin to Cuzco. throughout, rachis wings and those of other axes
Surinam to Colombia, south to Bolivia. sometimes conduplicate. Pinnae 1 2-24 pairs, ad-
Besides the characters used in the key, Hymen- nate, and basal segments overlapping the rachis,
ophyllum trichomanoides can often be distin- larger ones 2.5-5 cm long, bearing 7-10 pairs of

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 61


secondary segments, some elongate pinnae inter- weg 1530 (holotype, K!; isotypes, BM, p; photo,
mixed with normal ones. Ultimate segments en- F of P).
tire, plane to slightly undulate. Sori 8-many per

pinna. Indusia relatively large, subrotund, circular Leaves 7-28 cm long, 2.5-5(-6) cm broad. Pet-
in outline or broader than long, apex erose, base iole 1.5-8(-10) cm long, 0.3-0.7 mm in diameter,
rounded, not immersed in the greatly constricted marginate to alate throughout or at least distally
segment apex, receptacle short, filiform, bearing (wings sometimes deciduous), essentially gla-
7-12 sporangia. brous. Lamina lanceolate to ovate, elliptic, or ob-
lanceolate, scarcely to strongly reduced at base, 2-
On trees in wet forests, 2400-3300 m, Cuzco. 3-pinnatisect, glabrous. Rachis glabrous, straight
Venezuela; Colombia to Peru. or slightly flexuous, alate throughout, rachis wings
This is most closely related to Hymenophyllum and those of other axes often conduplicate. Pinnae
myriocarpum var. endiviifolium, from which it dif- 8-30 adnate and basal segments strongly
pairs,
fers in the eroseindusia tips and the presence of overlapping the rachis, larger ones 1-4 cm long,
one to several unusually elongate pinnae which bearing 5-10 pairs of secondary segments. Ulti-
are scattered among the normal ones the latter mate segments entire, plane to undulate, often with
a condition similar to that of H. undulatum. A conduplicate margins. Sori 5-many per pinna. In-
remarkably huge specimen, Lechler 2420 (B; not dusia subglobose or broader than long, much
other 2420 at B, F, K: H. elegans), was collected broader than (and not at all immersed in) the
inPeru at an undesignated site. This is apparently strongly constricted segment apex, rounded at base,
a monstrous form of H. ferax. It matches this rounded and entire at apex, receptacle short to
species in every way, including the erose indusia, nearly obsolete, filiform, bearing 8-20 sporangia.
but the leaf is 85 cm long and the petiole nearly
2 mm
in diameter. It is by far the stoutest specimen This, like H. polyanthos (q.v.), is a widespread
of Hymenophyllaceae we have seen and belies the and greatly misunderstood species which is sorely
appellation "filmy fern." in need of detailed study. It seems to consist of

several, not highly distinctive components which


Cuzco: Vilcabamba, Bues 1601 (us). Prov. Paucartam- are recognized as varieties. With var. myriocar-
bo, Pilahuata, Vargas 4912 (us). Dept. Unknown: Lech- pum might possibly be included H. axillare Sw.
ler 2420 (B in part). of the West Indies and H. andinum Bosch of Ec-
uador. Of all the names, H. axillare has priority.
The species occurs in Mexico and Central Amer-
10. Hymenophyllum myriocarpum Hooker, Sp. fil. icaand in South America from Venezuela and
1: 106, /. 37 d. 1844. TYPE: Colombia, Hart- Colombia to Bolivia.

Key to Varieties

a. Lamina elliptic to oblanceolate, broadest at or beyond the middle; several or many proximal pinnae
somewhat to greatly reduced lOa. var. myriocarpum
a. Lamina ovate to lanceolate, broadest near the base; 1 or 2 proximal pinnae (if any) slightly reduced
b
b. Larger leaves 7-1 5 cm long; pinnae commonly patent and strongly imbricate

lOb. var. nigrescens


b. Larger leaves ( 1 2-) 1 5-28 cm long; pinnae (most of them) ascending and contiguous or only slightly
imbricate . lOc. var. endiviifolium

1 Oa. Hymenophyllum myriocarpum Hooker, var. Leaves (9-) 12-28 cm long. Lamina elliptic, ob-
myriocarpum. long or oblanceolate, broadest at or beyond the
middle, several proximal pinnae somewhat to
greatly reduced. Rachis and ultimate segments
Mecodium myriocarpum (Hooker) Copel., Philipp. J. plane, some of their margins occasional condu-
Sci. 67: 25. 1938. plicate. Sori 10-20 per pinna.

62 FIELDIANA: BOTANY
In wet forests, usually pendent from tree trunks Skoget al. 5157 (us). Cuzco: Valley of Urubamba, Sues
and branches, or on wet, mossy rocks, rarely on A18,A19(us).
the forest floor, (1 700-)2000-3200 m, Cajamarca,
Amazonas, Huanuco to Cuzco.
Mexico to Costa Rica; Venezuela; Colombia to lOc. Hymenophyllum myriocarpum var. endivi-
Bolivia. ifolium (Desv.) Stolze, stat. & comb. nov.

Cajamarca: Prov. Hualgayoc, Hacienda Taulis, Hymenophyllum endiviifolium Desv., Mem. Soc. Linn.
Hutchison & Bismarck 6415 (F, GH [as Lambayeque], Paris 6: 334. 1827. TYPE: "Peruvia" (holotype,
MO, uc, us). Amazonas: Prov. Bongara, WSW
of Po- p; photos, GH, uc, us).
macocha, Wurdack 866 (GH, us). Huanuco: Mito, Bryan Hymenophyllum multiflorum Rosenst., Meded. Rijks-
206a (F). Pasco: Enenas (as Junin), Pichis Trail, Killip & Herb. 19: 5. 1913. TYPE: Bolivia,Comarapa,
Smith 25700 (F, NY, us). Cuzco: Prov. Urubamba, along Herzog 1951 (holotype, B; isotype, us!).
Rio Urubamba near town of Machu Picchu, Tryon & Mecodium multiflorum (Rosenst.) Copel., Philipp. J.

Tryon 5410 (F, GH, us). Sci. 67: 25. 1938.


Mecodium endiviifolium (Desv.) Pic. -Sen, Webbia 28:
469. 1973.

lOb. Hymenophyllum myriocarpum var. nigres- Leaves (12-) 15-28 cm long. Lamina ovate or
cens (Liebm.) Stolze, stat. & comb. nov. ovate-lanceolate. Rachis and ultimate segments
slightly to strongly undulate, their margins often

Hymenophyllum nigrescens Liebm., Kongel. Danske conduplicate. Sori 1 6-many per pinna.
Vidensk. Selsk. Skr., Naturvidensk. Afd. 5,1: 292.
1849. LECTOTYPE (designated by A. R. Smith, On trees and wet banks in deep forest, 2400-
Flora of Chiapas 2: 133. 1981): Mexico, Puebla,
4200 m, Amazonas, Huanuco, Huancavelica,
Chinantla, Liebmann 537 (lectotype, c; isolec-
Cuzco.
totypes, BM, K, P).

Sphaerocionium nigricans KJotzsch, Linnaea 18: 536. Colombia; Ecuador; Peru; Bolivia.
1 844, not Hymenophyllum nigricans Colla, 1836.
SYNTYPES: Peru, Dombey 87 (syntype, u!; iso-
Amazonas: Prov. Chachapoyas, Calla Calla slopes near
syntype, B); Venezuela, Aragua, Colonia Tovar, km 4 5-4 1 8 of Leimebamba-Balsas road, Wurdack 1750
1

Moritz 268 (syntype, B; isosyntype, in part, K!),


(us, USM). Huanuco: Prov. Huanuco, Mirador, road Aco-
268b (syntype, B; isosyntype, BM).
mayo to Chanchao, Mexia 7761 (F, GH, K, MO, uc, us).
Hymenophyllum nigricans (KJotzsch) Kunze, Bot. Zeit.
Huancavelica: Prov. Tayacaja, Chuspi-Tocas, between
(Berlin) 244. 1847. Colcabamba and Paucarbamba, Tovar 2050 (GH, USM).
Hymenophyllum nigricans (KJotzsch) Copel., Phillip. Cuzco: Prov. Paucartambo, Dist. Marcachea, near
J. Sci. 67: 25. 1938.
Achirani, Vargas 11143 (F, K. in part, uc).

Leaves 7-1 5 cm long. Lamina ovate. Rachis and


ultimate segments plane, their margins frequently
conduplicate. Sori 3-15 per pinna.
11. Hymenophyllum undulatum (Sw.) Sw., J. Bot.
(Schrader) 1800(2): 101. 1802.
Erect or arching from tree trunks or branches,
rarely from wet rocks, in thickets and wet forests,
Leaves 6-35(-40) cm long, 0.7-2.5 cm broad.
1750-3400 m, Amazonas to Cuzco. Petiole 0.5-3(-5) cm long, 0.15-0.3 mm in di-

Southern Mexico to Costa Rica; Venezuela; Co- ameter, nonalate, essentially glabrous. Lamina lin-
lombia to Bolivia. ear to narrow-elliptic, often strongly and gradually
reduced at base, 2-3-pinnatisect, glabrous. Rachis
Sterile specimens can easily be confused with
H. polyanthos, but the latter occurs at lower ele- glabrous, marginate to crispate-alate, commonly
nonalate toward base, or discontinuously alate on
vations, and laminae are less compact. See dis-
cussion of H. polyanthos for further comments. alternate sides due to the long-decurrent pinna
bases. Pinnae 1
6-many pairs, adnate, commonly
Amazonas: Prov. Chachapoyas, Cerros de Calla Calla less than 2 cm few of them greatly
long, but often a
above Leimebamba, Hutchison & Wright 6987 (in part: elongate and lamina-like, normal ones bearing 3-
F, GH, uc). San Martin: Prov. Rioja, Pedro Ruiz-Moy- 8(-10) pairs of secondary segments. Ultimate seg-
obamba Road, D. Smith 4376 (F, MO). Pasco: Prov. Ox- ments entire, plane, or the margins conduplicate
apampa, Abra los Mellizos, 4-8 km from Enenas, Skog
in drying, to undulate or markedly crispate. Sori
et al. 5033 (vs). Junin: Huacapistana, Killip & Smith
24164 (us). Ucayali: Plantation Azul, Ridoutt (USM). Prov. 3-many per pinna. Indusia nearly circular, but on
Coronel Portillo (as Loreto), NE of pass at La Divisoria, mature son most of them broader than long, not

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 63


or scarcely immersed in the segment tissue, com- from tree branches sometimes reach 40 cm in

monly truncate at base, the margins entire and length. Several species have been described based
plane to crispate, receptacle very short, filiform, on whether the tissue is plane or undulate or
bearing 1^4(-5) sporangia. strongly crispate. However, comparison of indi-
viduals throughout the entire range seems to in-
This species varies greatly as to length and shape dicate that few of these characters are very con-
of lamina and margins of rachis wings and ulti- sistent. Only the conspicuously crispate variants
mate segments. In rocky habitats leaves usually seem to merit recognition, as indicated in the fol-
tend to be less than 10 cm long and narrowly el- lowing key.
liptic, whereas the flaccid, linear leaves pendent

Key to Varieties

a. Ultimate segments and rachis wings plane to undulate, margins often discontinuously conduplicate
in drying; rachis nonalate in the proximal '/3 to /2 or nearly throughout
l
1 la. var. undulatum
a. Ultimate segments and rachis wings markedly crispate, often so strongly as to appear dentate; rachis
commonly alate except near lamina base, or wings sometimes lacking in the proximal 'A or lk of the
rachis . ..lib. var. fendlerianum

1 1 a. H ymenoph y Hum undulatum var. undulatum. cartambo, slopes of Pilahuata, Vargas 4909 (MO, us).
Puno: Sandia, Weberbauer 1293 (B).

TrichomanesundulatumSw.,Prodr. 137. 1788. TYPE:


Jamaica, Swartz (holotype, s; isotypes, B, Herb.
Willd. 20238, BM!, P; photos, F & us of s).
Hymenophyllum reniforme Hooker, Sp. fil. 1: 110, /. 1 1 b. Hymenophyllum undulatum var. fendlerian-
38c. 1844. TYPE: Peru, Mathews 1783 (holo- um W. Sturm) comb. & stat. nov.
(J. Stolze,
type, K.!; isotypes, B!, F!, K!, P!; photos, F & us of
P).
Hymenophyllum fendlerianum J. W. Sturm in Mart.,
Hymenophyllum rimbachii Sodiro, Anal. Univ. Quito
13: 47 (Crypt, vase. quit. 33). 1893. TYPE: Ec-
Fl. bras. 1(2): LECTOTYPE (desig-
291. 1859.

uador, Prov. Azuay, Rimbach (holotype, not lo-


nated by Lellinger, Mem. New York Dot. Gard.
38: 12. 1984): Venezuela, Edo. Aragua, Colonia
cated; isotype, us!).
Mecodium undulatum (Sw.) Copel., Philipp. J. Sci. 67: Tovar, Fendler 35 (us; isolectotype, BR; photo, us
26. 1938. of BR).
Hymenophyllum contortum Bosch, Ned. Kruidk. Arch.
5(3): 170. 1863. TYPE: Costa Rica, Aguacate,
In wet forests on rock cliffs or pendent from tree
Hoffman (B).
trunks and branches, 1500-4100 m, Huanuco to Hymenophyllum polycarpum Kuhn, Linnaea 35: 391.
Puno. 1868. TYPE: "Peruvia" (holotype, B!; photo, F).
Greater Antilles; Guadeloupe; southern Mexico Mecodium fendlerianum (J. W. Sturm) Copel., Phil-
ipp. J. Sci. 67: 26. 1938.
to Panama; Surinam to Colombia, south to Bo-
Mecodium contortum (Bosch) Copel., Philipp. J. Sci.
livia; southeast Brazil. 67: 26. 1938.
Hymenophyllum reniforme and H. rimbachii
were based on depauperate specimens in which on wet, rocky cliffs, or pendent from
In forests
ultimate segments are mostly plane but with their tree trunksand branches, 1250-2900 m, Ama-
margins conduplicate. Thus the segments appear zonas, Huanuco, Junin, Puno.
to be much narrower than typical H. undulatum.
Hispaniola; southern Mexico; Guatemala; Cos-
Some of this folding of the margins appears to be ta Rica; Surinam to Colombia, south to Bolivia;
a product of drying and some of it merely part of southeastern Brazil.
the natural variation of the species.
Amazonas: Prov. Bagua, Cordillera Colan SE of La
Huanuco: Tambo
de Vaca, Bryan 624 (F, us). Pasco: Peca, Barbour 3539, 3738 p.p. (MO). Huanuco: Cushi,
Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, trail to Tambo de Vaca, Bryan 684 (F, us). Playapampa,
van der Werffet al. 8482 (MO, uc). Junin: Prov. Tarma, Macbride 4498 (F, us). Junin: Schunke Hacienda, above
Yanango Mountains, Weberbauer 2135 (B). Cuzco: Pau- San Ramon, Killip & Smith 24843 (NY, us). Puno: Prov.

64 FIELDIANA: BOTANY
Carabaya, road from San Gaban to Macusani, Maas et 13. Hymenophyilum trichophyllum HBK., Nov.
al. 6109 (MO). Dept. Unknown: Without location, Dom-
gen. sp. 1: 27. 1815.
bey (P).

Leaves indeterminate, to 25 cm long and 3 cm


broad. Petiole 0.5-4 cm long, 0.2-0.25 in di- mm
ameter, nonalate, sparsely pubescent with simple
to stellate trichomes, or glabrate. Lamina 2-pin-
12. Hymenophyllum molle Morton, Contr. U.S.
nate-pinnatisect, abundantly pubescent, the rachis
Natl. Herb. 29: 149. 1947.TYPE: Peru, Cuz-
commonly nonalate throughout. Pinnae short-
co, Vilcabamba, Achiyayoc Inca ruins, Biies
stalked (at least proximal ones), cut deeply or quite
2102 (holotype, us!; isotype, GH!).
to costa, costae partially nonalate or discontin-

uously alate by the decurrent bases of segments,


Sphaerocionium molle (Morton) Pic.-Ser., Webbia 28:
normal pinnae with 2-3(-4) pairs of segments (but
471. 1973.
often a few of them greatly elongate and equaling
the lamina in size and shape), the segments (or
Leaves indeterminate, to 35 cm long and 3.5 cm
broad. Petiole 3-6 cm long, 0.2-0.3 mm
in di- pinnules) simple or bifid, plane to undulate or cris-
pate, veins and margins abundantly to densely pu-
ameter, nonalate, sparsely pubescent with simple
bescent with stellate trichomes, but trichomes
to (rarely) forked trichomes, or glabrate. Lamina
lacking between veins and margins. Indusia about
2-pinnatisect or nearly 2-pinnate, abundantly pu-
as broad as long and nearly as broad as the segment
bescent, the rachis commonly nonalate through-
out. Pinnae short-stalked (at least proximal ones), tips.

cut deeply or nearly to costa, costae regularly and


The species occurs in Guatemala; Costa Rica;
conspicuously alate on both sides, the 4-9 pairs
of segments simple or bifid, entire, plane, veins Panama; Guyana to Venezuela and south to Bo-
livia.
and margins abundantly or densely pubescent with
stellate trichomes, but trichomes lacking between
The leaves of this delicate fern sometimes in-

veins and margins. Indusia slightly broader than


tertwine to form mats on the trunks of trees. A
singular feature the irregular character of the
is
long, often broader than the segment tips.
pinnae: often, but not always, one to several pin-
nae become greatly elongate and once again pin-
Endemic. In forests, 2200-3600 m, Cuzco.
nate-pinnatisect, each of these sometimes equaling
the length of the leaf on which it is borne. Of the
Cuzco: Huadquina, Biies 708 (us). Prov. La Conven-
cion, Biies 2072, 2115 (us). Prov. Paucartambo, Vargas several varieties of Hymenophyllum trichophyl-
3636 (GH). lum, two are recognized in Peru.

Key to Varieties

a. Ultimate segments plane to slightly undulate 13a. var. trichophyllum


a. Ultimate segments conspicuously undulate to crispate 13b. var. buesii

13a. Hymenophyllum trichophyllum var. tricho- Elsewhere, in wet forests, on rocks, clay banks
phyllum. TYPE: Venezuela, Cumana, be- or trees, 2000-3800 m; range the same as the
tween Cocollar and Guardia de San Augustin, species.
Humboldt (holotype, P). The name H. procerum was based on H. pul-
chellum sensu Mett., for which there was no de-
Hymenophyllum procerum Bosch, Ned. Kruidk. Arch. scription. The specimen probably intended as the
4: 409. 1859, nom. nud.
type is the only collection of var. trichophyllum
Sphaerocionium trichophyllum (HBK.) Copel., Phil- thus far known from Peru. It closely matches other
ipp. J. Sci. 67: 32. 1938.
specimens of var. trichophyllum and is cited here.

In Peru, thus far known only from one collec-


tion: on shaded rocks, Puno. Puno: San Gaban, Lechler 2250 (B, BR, F, P).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 65


13b. Hymenophyllum trichophyllum var. buesii only by lack of trichomes on the veins, and by
Morton, Contr. U.S. Natl. Herb. 29: 1 52. 1 947. having marginal trichomes mostly three-forked at
TYPE: Peru, Cuzco, Prov. La Convention, the base. However, in H. elegans, trichomes are
Abra Mirador, Bues 2076 (holotype, us!; is- only occasional to rare or lacking on the veins, and
otype, GH!). marginal trichomes are highly variable, often in-
cluding many which are three-forked at base. The
Thus far known only from the type collection, isotype of H. tenerrimum has occasional tri-
3150m. chomes on the veins, and a number of marginal
Although Morton distinguished his var. buesii ones that are simple or one-forked. In fact, both
from var. trichophyllum solely on the obviously taxa may be only a variant of//, lineare (Sw.) Sw.,
undulate segments, it also appears that the former of Central America and the West Indies, which
has less densely pubescent laminae. However, un- also lacks trichomes on veins, but which appears
tilmore specimens are collected it remains to be to have more pinna segments and simpler and less
seen if either of the differences are sufficiently sig- variable marginal trichomes. Further study is

nificant to merit even varietal separation. needed.

San Martin: Tarapoto, "in monte Campana," Spruce


14. Hymenophyllum elegans Sprengel, Syst. veg. 4700, 4701 (K). Puno: San Gaban, Lechler (B). Dept.
Unknown: Lechler (NY, us), Lechler 2420 (B, F, K; another
4: 133. 1827. TYPE: Brazil, Sellow (holotype,
2420 at B is H. ferax).
LZ destroyed; isotype, B!).

Hymenophyllum tenerrimum Bosch, Ned. Kruidk.


Arch. 5: 185. 1863. TYPE: Peru, San Martin,
15. Hymenophyllum adiantoides Bosch, Ned.
5: 184. 1863. TYPE: Peru, San
Kruidk. Arch.
Tarapoto, "in monte Campana," Spruce 4702
(holotype, K!; isotype, BM!). Martin, Tarapoto, Spruce (holotype, L; prob-
Sphaerocionium tenerrimum (Bosch) Pic.-Ser., Web- able isotypes, B!, GH!, K!, L!, NY, us; photos,
bia28:472. 1973.
GH & us of L).
Sphaerocionium elegans (Sprengel) Copel., Philipp. J.
Sci. 67: 32. 1938.
Sphaerocionium adiantoides (Bosch) Pic.-Ser., Web-
bia28:470. 1973.
Leaves determinate, to 20 cm long and 2 cm
broad. Petiole 0.5-2 cm long, 0.15-0.2 in di- mm Leaves indeterminate, to 1 2 cm long and 2 cm
ameter, nonalate, sparsely pubescent with simple
or forked trichomes. Lamina
broad. Petiole 1-2 cm long, 0.2-0.25 mm
in di-
pinnate-pinnatisect,
ameter, nonalate, sparsely pubescent with (mostly)
sparsely to moderately pubescent, the rachis non-
stellate trichomes. Lamina pinnate-pinnatisect,
alate, or narrowly and irregularly alate distally or
sparsely to moderately pubescent, the rachis non-
throughout due to the decurrent bases of pinnae.
alate near the base, narrowly alate distally or ir-
Pinnae sessile or adnate, pinnatifid or pinnatisect,
regularly alate due to the decurrent bases of pin-
the l-2(-3) pairs of segments, simple or (basal
nae. Pinnae adnate, pinnatifid, or pinnatisect, the
ones) bifid, entire, plane to slightly undulate, mar-
3-4 pairs of segments simple or bifid, entire, slight-
gins sparsely to moderately pubescent, with tri-
ly undulate, veins and margins sparsely to mod-
chomes highly variable: simple to 2-3-forked at
erately pubescent with (mostly) stellate trichomes,
base, or the branches again forked; trichomes rare
but trichomes lacking between veins and margins.
on veins and lacking between veins and margins.
Indusia as long as or slightly longer than broad,
Indusia slightly broader than long, about as broad
about as broad as the segment tips.
as the segment tips.

On shaded rocks, San Martin, Puno, and from Apparently known only from the type collec-
tion. With this probably should be included Hy-
two other collections, localities unspecified, 3000
m. menophyllum sprucei Baker, under which see
Comments.
Elsewhere, in wet forests, on rocks, clay banks
or tree trunks, 800-2400 m; Guatemala; Costa
Rica; Panama; West Indies; Venezuela; Colombia;
Peru; Bolivia; Brazil. 16. Hymenophyllum hirsutum (L.) Sw., J. Bot.
Hymenophyllum tenerrimum was said to differ (Schrader) 1800(2): 99. 1802.

66 FIELDIANA: BOTANY
Trichomanes hirsutum L., Sp. pi. 2: 1098. 1753. TYPE: W side of Cordillera de San Matias, D. Smith 2054 (MO).
based on t. 50b of Plumier, Traite foug. Amer., Cuzco: Prov. La Convencion, Valle San Miguel, San Ja-
apparently representing a plant from the West cinto, Bues 2139 (us). Puno: San Gaban, Lechler 2296
Indies. (B).
Trichomanes ciliatum Sw., Prodr. 136. 1788. TYPE:
Jamaica, Swartz (holotype, s; isotype, B; photo,
us of s).
Hymenophyllum ciliatum (Sw.) Sw., J. Bot. (Schrader) 1 7. Hymenophyllum crispum HBK., Nov. gen. sp.
1800(2): 100. 1802. 1: 26. 1815. TYPE: Venezuela, Silla de Ca-

Sphaerocionium ciliatum (Sw.) Presl, Hymenophyl- racas, Humboldt & Bonpland (holotype, p;
laceae 34. 1843.
photos, GH & us of P). Figure lid.
Sphaerocionium hirsutum (Sw.) Presl, Hymenophyl-
Iaceae34. 1843.
Sphaerocionium crispum (HBK.) Klotzsch, Linnaea
1: 537. 1844.
Leaves determinate, to 12 cm long and 4 cm
broad. Petiole 0.3-2.3 cm long, 0.2-0.3 mm in
Leaves determinate, to 20 cm long and 4 cm
diameter, strongly alate (often nearly to base),
broad. Petiole 0.5-5 cm long, 0.1-0.3 in di- mm
sparsely provided with stellate trichomes. Lamina
ameter, alate near apex, sparsely provided with
2-pinnatisect, sparsely to moderately pubescent,
delicate, simple or 1 -forked trichomes. Lamina
the rachis regularly and conspicuously alate
pinnate to nearly 3-pinnate, sparsely pubescent,
throughout. Pinnae 6-15 pairs, ovate or oblong- the rachis regularly and conspicuously alate
ovate, deeply incised, the 2-5 pairs of segments
throughout, the wing strongly undulate or undu-
plane, simple to bifid, the veins and margins pu-
late-crispate. Pinnae 8-20 pairs, simple (distally)
bescent with stellate or (on the margins) 2-forked
or bifid to nearly 2-pinnate, the 1-8 pairs of seg-
trichomes, but trichomes lacking between veins
ments undulate or crispate, the veins and margins
and margins. Indusia as long as or (usually) longer
sparsely pubescent, marginal trichomes simple or
than broad, somewhat broader than the segment
1 -forked, but trichomes
lacking between veins and
tips.
margins. Indusia as long as or (usually) longer than
broad, Oiten broader than the segment tips.
on branches and trunks of
In dense, wet forests,
trees,500-2000 m, Loreto, Amazonas to Puno. The species occurs in Jamaica, Mexico, Gua-
Mexico to Panama; West Indies; Guianas to temala, Costa Rica, Venezuela, Colombia, Peru,
Venezuela, south to Bolivia and Brazil. Bolivia, and Brazil.
Hymenophyllum crispum might be confused with
Amazonas: Prov. Bagua, E of La Peca, Barbour 2753 H. valvatum, another species in the subgenus with
(MO). San Martin: Tarapoto, Monte Campana, Spruce undulate segments. However, the latter is larger
4698 (K). Loreto: Balsapuerto, lower Rio Huallaga basin,
and coarser with a much thicker petiole, and the
Killip & Smith 28539 (F, GH, NY, us). Huanuco: Prov.
Huanuco, Dist. Churubamba, crest of Santo Toribio, rachis wing and segments are not as strongly un-
Mexia 8145 (B, F p.p., GH, us). Pasco: Prov. Oxapampa, dulate. Two varieties are recognized here.

Key to Varieties
a. Pinnae simple secondary segments 1-2 pairs, simple to bifid
(distally) to trifid or, if pinnatisect, the
1 7a. var. crispum
a. Pinnae (larger ones) 2-pinnatisect, the secondary segments 4-8 pairs, pinnatisect
1 7b. var. bipinnatisectum

1 7a. Hymenophyllum crispum var. crispum. Fig- Rio Lachac, Sues 1813 (GH, us). Prov. Urubamba, Ma-
ure lid. chu Picchu, lit is et al. 1061 (GH, uc, us). Puno: Sandia,
Weberbauer 797 (B).

In forests, on trees or wet banks or cliffs, 2 1 50-


3000 m, Amazonas, Cuzco, Puno.
Range the same as the species.
1 7h. Hymenophyllum crispum var. bipinnatisec-
Amazonas: Prov. Chachapoyas, along Rio Ventilla, W tum Morton, Contr. U.S. Natl. Herb. 29: 161.
of Molinopampa, Wurdack 1510 (F, GH, uc, us). Cuzco: 1947. TYPE: Peru, Cuzco, Alturas de Sieve,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 67


Bties 1580 (holotype, us!; isotype, GH!; photo, imens determined to be H. microcarpum, the ul-

us). timate segments are only slightly undulate. A per-


fect intermediate is the isotype of H. platylobum,

Endemic. which is placed here in synonymy with H. val-

vatum. The petiole trichomes of this specimen


Thus far known only from the type and one other match those of//, microcarpum, but the segments
collection from Peru: Cuzco: Alturas de Sieve, Bues 1528 and rachis wings are scarcely or only slightly un-
(us). dulate. Other features considered by Morton and
subsequent workers seem not to be significant, at
least in specimens from Peru. Furthermore, both

18. Hymenophyllum valvatum Hooker & Grev., species may be only robust forms of H. hirsutum,
Icon. fil. 2, /. 279. 1831. TYPE: Ecuador, for- which seems to differ chiefly in its smaller lamina
ests of Esmeraldas, on trunks of trees, Jame- and more delicate petiole. The entire complex is
son (holotype, K; possible isotype, K!). in need of detailed study.

Hymenophyllum crispatulum Bosch, Ned. Kruidk. Huanuco: Rio Llullapichis watershed, Cerros del Sira,
Arch. 4: 412. 1859. TYPE: Peru, Tatanara (Puno), Dudley 13339 (GH). Junin: Chanchamayo Valley, C.
Lechler 2530 (holotype, L?; isotypes, B!, P!; pho- Schunke 1491, 1559 (F, us). Puno: "St. Gavan" (San
tos, GH & us of P). Gaban), Lechler (K). Prov. Sandia, Chunchusmayo, We-
Hymenophyllum platylobum Bosch, Ned. Kruidk. berbauer 1237 (B).
Arch. 5:189.1863. TYPE: Peru, Dept. Puno, San
Gaban, Lechler 2489 (apparently cited in error
originally as 2498) (holotype, P; isotype, F!; photo,
us! of P).
Sphaerocionium valvatum (Hooker & Grev.) Copel., 19. Hymenophyllum microcarpum Desv., Mem.
Philipp. J. Sci. 67: 31. 1938. Soc. Linn. Paris 6: 333. 1827. TYPE: His-
paniola (holotype, P; photo, uc).
Leaves determinate, to 20 cm longhand 6 cm
broad. Petiole of mature leaves (2-)3-lOcm long, Hymenophyllum beyrichianum Kunze, Linnaea 9: 108.
0.4-1.0 mm in diameter, commonly alate just be- 1834. TYPE: Peru, Huanuco, Pampayacu, Poep-

low lamina base, pubescent with simple or forked pig (holotype, LZ destroyed).
Mecodium microcarpum (Desv.) Copel., Philipp. J.
trichomes or glabrate. Lamina pinnate-pinnatisect
Sci. 67: 25. 1938.
to nearly 2-pinnate-pinnatifid, sparsely to mod- Sphaerocionium microcarpum (Desv.) Copel., Phil-
erately pubescent, the rachis alate throughout, the ipp. J. Sci. 67: 34. 1938.

wing somewhat undulate. Pinnae 8-1 5 pairs, ovate-


to elliptic-lanceolate, incised deeply to the costa Leaves determinate, to 30 cm long and 10 cm
into bifid or pinnatifid pinnules, ultimate segments broad. Petiole of mature leaves (3-)5-15 cm long,
slightly to strongly undulate, the veins (abaxially) 0.6-1.0 mmin diameter, broadly alate just below
and margins sparsely to amply pubescent, but tri- lamina base, pubescent with simple, forked, and
chomes lacking on tissue between veins and mar- stellate trichomes or glabrate. Lamina pinnate-

gins. Indusia commonly longer than broad, and pinnatisect to nearly 2-pinnate-pinnatifid, sparse-
slightly longer than the segment tips. ly to moderately pubescent, the rachis conspicu-
ously alate throughout. Pinnae 10-16 pairs, ovate-
On wet
tree trunks in forests, 900-1800 m, to elliptic-lanceolate, incised deeply or nearly to
Huanuco, Junin, Puno. the costa into bifid or pinnatifid pinnules, the ul-
Lesser Antilles; Guyana to Colombia, south to timate segments entire or deeply bifid, the veins
Bolivia. (abaxially)and margins sparsely to moderately pu-
It is with some reluctance that this
is maintained bescent, but trichomes lacking on tissue between
as distinct from Hymenophyllum microcarpum, the veins and margins. Indusia commonly longer
from which it seems to differ only in the undulate than broad, and slightly broader than the segment
ultimate segments and (sometimes) rachis wings, tips.
and the character of the petiole trichomes. Even
these features are variable, as only in a few spec- On tree trunks and clay banks in wet forests,
imens examined are the segments and rachis wings 1500-2700 m, Amazonas, Huanuco to Cuzco.
markedly undulate to crispate most are only Greater Antilles; Guatemala to Panama; Guy-
moderately so. Conversely, in a number of spec- ana to Colombia, south to Bolivia and Brazil.

68 FIELDIANA: BOTANY
With this probably might be included H. val- 21. Hymenophyllum simplex Morton, Contr. U.S.
vatum, under which see for further discussion. Natl. Herb. 29. 171. 1947. TYPE: Peru,
Huanuco, Tambillo, Jelski 897 (us!).
Amazonas: Prov. Bagua, E of La Peca, Barbour 2547
(MO). Huanuco: Rio Llullapichis watershed, ascent of Sphaerocionium simplex (Morton) Pic.-Ser., Webbia
Cerros del Sira, Dudley 13290B (MO). Pasco: Prov. Ox- 28: 472. 1973.
apampa, Palmazu, van der Werff et al. 8410 (MO, uc).
Junin: Chanchamayo Valley, C. Schunke 463 (f, us).
Cuzco: Prov. La Convention, Bites 2018, 2035 (us), 2034
Leaves determinate, fragile and small, 5-7 cm
(GH). Prov. La Convention, Alturas de Pintobamba, long. Petiole slender, 0.1-0.15 mm
diameter, in

Vargas 3294 (MO, us). nonalate, glabrate or sparsely pubescent with sim-
ple to stellate trichomes. Lamina simply pinnate
or pinnatisect, moderately pubescent, the rachis
20. Hymenophyllum ruizianum (Klotzsch) Kunze, narrowly alate distally, nonalate at base or in the
Hot. Zeit. (Berlin) 1847: 199. 1847. proximal Pinnae few, ascending, simple, lin-
half.

ear, entire, the tissueand veins amply provided


Sphaerocionium ruizianum Klotzsch, Linnaea 18: 535. with orange, stellate trichomes. Veins lacking ac-
1 844. TYPE: Peru, Ruiz 85 (holotype, B!; isotype,
cessory wings. Indusia about as broad as long, usu-
MA?, P; photos, GH & us of P).
ally narrower than the pinna tips.

Leaves determinate, to 45 cm long and 12 cm


Endemic. Besides the type (Huanuco) thus far
broad. Petiole of mature leaves (7-) 1 0-20 cm long,
known only from two other collections: Cajamarca
0.7-1.5 mmin diameter, nonalate, densely pu-
and San Martin. Habitat and elevation not re-
bescent with mostly simple or forked trichomes.
corded.
Lamina 2-3-pinnatifid, sparsely to amply pubes-
This delicate little fern is distinctive among the
cent on both sides of the lamina, the rachis alate
species of subg. Leptocionium in its few, ascend-
distally. Pinnae commonly patent or their tips
ing, linear, entire pinnae. Although we now know
drooping, 10-18 pairs, ovate-lanceolate, incised it only from two Peruvian collections, its small,
nearly to the costa into pinnatifid or 2-pinnatifid
inconspicuous leaves make it difficult to detect. It
pinnules, the ultimate segments entire or bifid, the is hoped that more diligent search will uncover
veins and margins sparsely to moderately pubes-
additional specimens throughout Peru and, per-
cent, but trichomes lacking on tissue between veins
haps, in other areas of South America as well.
and margins. Indusia varying from broader than
long to longer than broad, commonly slightly
Cajamarca: Cutervo, Jelski 914 (P, us). San Martin:
broader than the segment tips.
Tarapoto, Spruce (K), collection is without number, but
with notation "conf. 4048"; #4048 is type number of H.
In wet forests, on tree trunks and banks of rav- tarapotense.

ines, sphagnum on forest floor, 1000-3300


and in

m, Amazonas, Huanuco, Pasco, Cuzco, Puno.


Colombia; Ecuador; Peru; Bolivia.
A singular feature of this species is the tendency 22. Hymenophyllum fragile (Hedwig) Morton,
Contr. U.S. Natl. Herb. 29: 172. 1947.
for the long, flexible pinnae to droop gracefully at
their tips, a character which is usually preserved
Trichomanes fragile Hedwig, Fil. gen. sp., t. 18. 1802.
in driedspecimens. Perhaps its nearest relative is TYPE: based on illustration from plant collected
H. trapezoidale Liebm., from Mexico and north- in "America meridionalis."
ern South America, which is a much smaller fern Sphaerocionium fragile (Hedwig) Pic.-Ser., Webbia 28:
with pinnae which are rather strong-
less dissected 471. 1973.

ly ascending (never drooping) and with more del-


icate, merely sparsely pubescent, petioles. Leaves determinate, (3-)6-18 cm long, 1.5-2.5
cm broad. Petiole ca. 0.2 mm in diameter, sparsely
Amazonas: Prov. Chachapoyas, above Leimebamba, to moderately pubescent with simple, forked, or
Hutchison & Wright 5553 (F, GH, MO, uc, us). Huanuco: often stellate trichomes. Lamina pinnate-pinna-
Playapampa, Macbride 4516 (F, us). Pasco: Prov. Oxa- moderately pubescent, the rachis broadly
tisect,
pampa, San Alberto, Cordillera de Yanachaga, van der and consistently alate throughout. Pinnae (3-)5-
Werffet al. 8423 (MO), 8439, 8491 (MO, uc). Cuzco: Prov.
La Convention, Dudley 10790 (F, GH, MO). Puno: San 1 2 pairs, deeply 1 -forked (distally) to pinnatisect,
Gaban, Lechler 2260 (B). the 1-2 pairs of segments entire, the tissue and

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 69


veins moderately provided with orange, short- the laminae have the rachis narrowly winged well
stalked, stellate trichomes. Veins lacking accessory into the proximal half, and these specimens might

wings. Indusia about as broad as long, usually nar- be confused with H. fragile. However, the alate
rower than the ultimate segments, the receptacle condition in these atypical leaves is irregular and
not exserted. mostly due to the decurrent bases of pinnae,
whereas in H. fragile the wings are very conspic-
In dense forests, on tree trunks or mossy banks, uous and of nearly consistent width throughout.
800-2700 m, Pasco, Junin, Ucayali, Cuzco.
Mexico to Panama; Greater Antilles; Venezuela Ama/onas: Prov. Bagua, Cordillera Colan NE of La
and Colombia to Bolivia and Brazil. Peca, Barbour 3415 (F, MO). Cuzco: Michihuanuncca,
Biies 720 (us). Puno: Prov. Carabaya, Ayapata, Vargas
This may be confused with Hymenophyllum ele-
1071 7 (GH).
gantulum, under which see for further discussion.

Pasco: Oxapampa, Soukup 2650 (GH). Junin: East of


Quimiri Bridge, near La Merced, Killip & Smith 23834
24. Hymenophyllum amabile Morton, J. Wash.
(NY, us). Ucayali (as Loreto): Prov. Coronel Portillo, NE
Acad. Sci. 22: 63. 1932. TYPE: Peru, Cuzco,
of pass at La Divisoria, Skog et al. 5156 (us). Cuzco: Prov. La Convencion, Huadquina, Bites 715
Prov. La Convencion, Pintobamba, Vargas 3262 (MO,
(holotype, us!; isotype, GH!; photo, F of us).
us).

Leaves indeterminate, to 40 cm long and 4 cm


broad. Petiole 0.2-0.3(-0.4) cm in diameter, non-
23. Hymenophyllum elegantulum Bosch, Ned. alate, sparsely pubescent with mostly stellate tri-
Kruidk. Arch. 4: 408. 1859. LECTOTYPE chomes. Lamina pinnate-pinnatisect, densely to-
(designated as type by Morton, Contr. U.S. mentose, the rachis nonalate throughout, or rarely
Natl. Herb. 29: 170. 1947): based on t. 33a
slightly alate due to decurrent bases of pinnae.
(Hooker, Sp. fil. 1 1 844), which in turn, ac-
.
Pinnae numerous, oblong or ovate-oblong, obtuse,
cording to Morton, was based on Ecuador, often incised nearly to the costa, the 4-8 pairs of
Pillzhum, Jameson 366 (K.?; probable isolec-
segments entire to forked, ultimate divisions en-
totype, NY!). the tissue and veins often obscured by a dense
tire,
tomentum of orange, stalked, stellate or bistellate
Hymenophyllum pulchellum sensu Hooker, Sp. fil. 1,
trichomes. Veins lacking accessory wings. Indusia
/. 33a. 1844, not Schlecht. & Cham. 1830.
broader than long, about as broad as the segment
Sphaerocionium elegantulum (Bosch) Copel., Philipp.
J. Sci. 67: 32. 1938. tips, the receptacle very short, not exserted.

Leaves indeterminate, 12-35 cm long, 2-4 cm Pendent from tree trunks and branches in wet

broad. Petiole ca. 0.2 mm


in diameter, glabrate or forests,2600-4200 m, Huanuco, Cuzco, Puno.
sparsely pubescent with simple or forked tri- Ecuador and Peru.
chomes. Lamina pinnate-pinnatisect, amply pu-
bescent, the rachis flexuous, nonalate, or some- Huanuco: Tambo de Vaca, Bryan 628 (F, us). Cuzco:
Prov. La Convencion, Vilcabamba, Biies 2108, 2109,
times alate in the distal portion. Pinnae subdistant
2110 (us). Prov. La Convencion, Loma Grande, Biies
to remote, ascending, 8-24 pairs, deeply pinnati-
2148 (F, GH, uc, us). Urubamba, Machu Picchu, Peyton
sect, the 4-6 pairs of segments 1-2-forked or bifid, & Peyton 1 104 (MO). Puno: Sandia, Weberbauer 796 (B).

the tissue and veins moderately provided with or-


ange, sessile to short-stalked, stellate trichomes.
Veins lacking accessory wings. Indusia about as 25. Hymenophyllum speciosum Bosch, Ned.
broad as long, and usually as broad as the ultimate Kruidk. Arch. 5: 181. 1863. LECTOTYPE
segments. (designated as type by Morton, Contr. U.S.
On tree trunks, in forests and wet clearings, 3000- Natl. Herb. 29: 175. 1947): Peru, Puno, San
3600 m, Amazonas, Cuzco, Puno. Gaban, on tree trunks, Lechler 2246 (L?; iso-

lectotypes, B!, F!, p; probable isolectotype, NY!;


Mexico to Panama; Greater Antilles; Venezuela
photos, GH, us of P).
and Colombia south to Peru and Bolivia.
Some leaves of Hymenophyllum elegantulum
Hymenophyllum spectabile Kuhn, Linnaea 35: 392.
vary somewhat in the character of the rachis wing, 1868. TYPE: Bolivia, Yungas, D'Orbigny 175
as in the case of Biies 720, cited below. Some of (holotype, B?; isotype, p; photo, us of P).

70 FIELDIANA: BOTANY
Sphaerocionium spectabile (Kuhn) Copel., Philipp. J. to long-stalked stellate trichomes, these mostly
Sci. 67:31. 1938.
spreading, not tightly appressed. Pinnae numer-
ous, larger ones 1-2.3 cm long, lanceolate or lin-
Leaves indeterminate, mature ones 30-180 cm
ear-lanceolate, acute to subattenuate, shallowly or
long, 4-9 cm broad. Petiole (of mature leaves) 0.5-
deeply incised, the 4-7 pairs of segments entire to
0.7 mm in diameter, nonalate, glabrate or sparsely bifid, the tissue and veins mostly obscured by a
pubescent with simple to stellate trichomes. Lam- dense tomentum or orange to tawny, short-stalked,
ina pinnate-pinnatifid or pinnate-pinnatisect,
stellate trichomes. Veins lacking accessory wings.
densely tomentose, the rachis nonalate through- Indusia commonly as broad as long, and as broad
out, commonly flexuous, with a tightly appressed as the segment tips, the receptacle not exserted.
covering of subsessile or short-stalked, stellate tri-
chomes. Pinnae numerous, larger ones (2.5-)3-8
Pendent from
trees or clay banks in wet forests,
cm long, lanceolate or linear-lanceolate, often at- 700-1900 m
throughout the range, but thus far
tenuate, incised deeply or nearly to the costa, the
apparently known in Peru only from one collection
6-14 pairs of segments shallowly to deeply bifid, cited below.
the ultimate divisions entire, the tissue and veins
Venezuela; Colombia; Peru.
mostly obscured by a dense tomentum of orange This and Hymenophyllum speciosum are very
(aging whitish), short-stalked, stellate trichomes.
closely related. See discussion of the latter for ad-
Veins lacking accessory wings. Indusia about as
ditional comments.
broad as long and often broader than the segment
tips, the receptacle not exserted.
San Martin: Near Tarapoto, in Monte Campana,
Spruce 4694 (BR, us).
Pendent from trees or clay banks in wet forests,
1800-3350 m, Amazonas, Pasco, Cuzco, Puno.
Peru and Bolivia.
This might better be treated as a variety ofHy- 27. Hymenophyllum lindenii Hooker, Sp. I'll. 1:

menophyllum karstenianum, as the two differ only 94, t. 34c. 1844. TYPE: Venezuela, Caracas,
quantitatively, e.g.,appressed versus spreading to- Linden 173 (holotype, K!; isotype, P; photos,
mentum, F of K, us of P).
strongly versus slightly flexuous rachis,
relative size and number of segments of pinnae.
Sphaerocionium lindenii (Hooker) Vareschi, Flora
Venezuela 1: 217. 1969.
Amazonas: Prov. Chachapoyas, Hutchison & Wright
5569 (F,GH, MO, uc, us); Sagastegui 7453 (F, HUT). Pasco:
Prov. Oxapampa, Cordillera San Gutardo, Leon et al. Leaves indeterminate, to 35 cm long and 8-15
534(usM). Cuzco: Vallede Lares, Biies 1808, 1817, 1821, cm broad. Petiole (8-) 10-1 5 cm long, 0.7-1 .2 mm
1931 (us). Valle de San Miguel, La Convention, Biies
in diameter, commonly nonalate, pubescent with
2178 (GH, MO, us).
simple to stellate trichomes. Lamina ovate or ovate-
lanceolate, not or slightly reduced at base, nearly
2-pinnate-pinnatisect, moderately pilose, the rachis
26. Hymenophyllum karstenianum J. W. Sturm, abundantly stellate-pubescent, scarcely alate, or
Bot. Zeit. (Berlin) 17: 298. 1859. LECTO- alate distally (or sometimes to base). Pinnae patent

TYPE to ascending, 14-20 pairs, 4-9 cm long, acuminate


(designated as type by Morton, Contr.
U.S. Natl. Herb. 29: 176. 1947): Venezuela, to subattenuate, incised nearly to the costa into 8-

Merida, Moritz 381 p.p. 14 deeply pinnatifid pinnules, the ultimate seg-
(BR).
ments entire to deeply bifid, rarely undulate or

Sphaerocionium karstenianum (J. W. Sturm) Pic.-Ser.,


between the veins sparsely provid-
crispate, tissue
Webbia28:471. 1973. ed with scattered, sessile or stalked, stellate tri-
chomes. Veins lacking accessory wings. Indusia
Leaves indeterminate, mature ones 15-50 cm broader than long, and about as broad as the seg-
long, 2-3.5(-4) cm broad. Petiole 0.4-0.5 mm in ment tips, receptacle not exserted.
diameter, nonalate, glabrate or sparsely pubescent
with simple to stellate trichomes. Lamina pinnate- Pendent from tree trunks in wet forests, 2300-
pinnatifid to -pinnatisect, densely tomentose, the 2450 m, Amazonas, Puno.
rachis nonalate throughout, straight or slightly Venezuela; Colombia; Ecuador(?); Peru.
flexuous, abundantly to densely covered with short- Hymenophyllum lindenii is one of the most ro-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 71


bust species in the subgenus, the long, stout petiole Pendent from tree trunks and clay banks in wet
often measuring a full millimeter in diameter. The forests, 600-2200 m, San Martin to Puno.
presence of a rachis wing, usually a good diagnostic Venezuela and Colombia to Peru and Bolivia.
feature inmost species of subg. Leptocionium, is It is not possible to separate this satisfactorily
not consistent in this or in the closely related H. from Hymenophyllum interruptum, which was said
plumieri. The rachis in H. lindenii is occasionally to differ in its groups of fertile pinnae interrupted
alate to the base of the lamina; more commonly by a few sterile ones, by its rachis not alate toward
the wing is present in the distal half but sometimes the base, and by variations in lamina base and in
nearly lacking throughout. The two Lechler
it is marginal trichomes. In a number of specimens
specimens cited below are most atypical, being examined during this study (identified as H. in-
broadly alate even well down the petiole, and with terruptum) fertile pinnae are continuous, and rach-
the wings very strongly crispate. The latter feature ises are fully to partially alate or (occasionally)

apparently was what prompted Mettenius's orig- nonalate throughout. The entire species complex,
inal annotation on this sheet as "//. valvatum." which also involves H. dependens Morton and H.
Lechler 2489 is a mixed collection, as the other superbum Morton from northern South America,
four specimens of 2489 at Berlin are isotypes of needs closer examination. All may be conspecif-
H. platylobum (= H. valvatum). ic.

San Martin: In monte Guayrapurima, prope Tara-


Amazonas: Prov. Bagua, Cordillera Colan, Barbour poto, Spruce 4025 (K, P). Huanuco: Rio Llullapichis wa-
3554 (MO, USM), 3602, 3751 (MO). Puno: near San Gaban, tershed, ascent of Cerros del Sira, Dudley 13014 (GH,
Lechler 2489 (u). Tatanara, Lechler 2563 (B, K). us), 13204, 13377 (GH). Junin: Colonia del Perene, Killip
& Smith 24955 (F, GH, NY, us). Cuzco: Prov. Paucartam-
bo, Pilahuata, Vargas 4914 (MO, us), 76756, 76767 (GH).
Madre de Dios: Prov. Manu, Shintuya, Chavez 861 (MO).
& Puno: Prov. Carabaya, Vargas 17562 (GH).
28. Hymenophyllum plumieri Hooker Grev.,
Icon. fil. 2, /. 123. 1829. TYPE: Ecuador,
western side of Pichincha, Jameson (holo-
type, K.!; photo, F). 29. Hymenophyllum plumosum Kaulf., Enum. fil.
267. 1824. TYPE: Brazil, Chamisso (holo-
Hymenophyllum interruptum Kunze, Linnaea 9: 107. type, LZ destroyed; isotype, LE?).
1834. TYPE: Peru, Huanuco, Pampayacu, Poep-
pig 1104 (holotype, w?; isotype, HBG).
Sphaerocionium plumosum (Kaulf.) Copel., Philipp.
Sphaerocionium plumieri (Hooker & Grev.) Presl, Hy-
J. Sci. 67: 30. 1938.
menophyllaceae 34. 1 843.
Sphaerocionium interruptum (Kunze) Presl, Hymen-
ophyllaceae 34. 1843. Leaves indeterminate, to 75 cm long and 6 cm
broad. Petiole 0.3-0.7 mm
in diameter, nonalate,
Leaves indeterminate, to 50 cm long and 3-8 pubescent with simple to stellate trichomes or gla-
cm broad. Petiole 2-7 cm long, 0.4-0.6(-0.7) mm brate. Lamina
pinnate-pinnatifid to rarely nearly
in diameter, nonalate (or sometimes alate just at 2-pinnate, densely appressed-tomentose, the rach-
the apex), pubescent with mostly forked to stellate is nonalate throughout. Pinnae numerous, lanceo-

trichomes. Lamina linear, usually strongly and late, pinnatifid to pinnatisect, or rarely incised to

gradually reduced at base, often interruptedly fer- the costa, the ultimate segments entire (some prox-
tile (e.g., groups of fertile pinnae separated by some imal ones bifid), the tissue and veins completely
sterile ones), pinnate-pinnatifid to -pinnatisect, obscured by a tomentum of sessile to short-stalked,
moderately pilose, the rachis alate throughout, or stellate trichomes. Veins (especially abaxially) with

occasionally alate only distally or not at all. Pinnae low, inconspicuous lamellae. Indusia about as
patent to ascending, 1 5-many pairs, 1 .5-6 cm long, broad as long, usually slightly broader than the
acute to subattenuate, the 5-12 pairs of segments segment tips.
simple and entire to deeply bifid, the tissue be-
tween the veins sparsely to moderately provided Epiphyte on trunks or branches in dense, wet
with sessile or stalked, stellate trichomes. Veins pendent from wet, clay
forests, or occasionally

lacking accessory wings. Indusia about as broad banks, 550-3100 m, Huanuco, Junin, Cuzco,
as long or usually broader, as broad as the segment Madre de Dios, Puno.
tips. Costa Rica to Colombia; Peru; Bolivia; Brazil.

72 FIELDIANA: BOTANY
The lamellae along the veins in this species are irregularly interrupted, conspicuous lamellae. In-
scarcely "wings" in the sense of being broad and dusia about as broad as long, equaling or slightly
thin flanges of tissue. Instead, they are commonly broader than the segment tips.
long, low outgrowths borne along the abaxial or
(sometimes) adaxial sides of the veins contiguous Epiphyte in dense, wet forests, 2000-3200 m,
to, but not in the same plane with, the segment Amazonas, Huanuco, Pasco, Cuzco.
tissue. They are often difficult to discern, since the Colombia; Ecuador; Peru.
veins and laminar surface are so densely tomen- Shape of lamina and pinnae can be variable and
tose. The best way to observe them is on the older irregular in this species. Typically the leaves are

(proximal) pinnae wheresome of the indument linear, with numerous pinnae 3 or 4 cm long, yet
at times, scattered along the rachis, there may be
may have away. For further discussion see
fallen
also Hymenophyllum tomentosum. a few greatly elongated, attenuate pinnae, occa-
1 5 cm in
Easily confused with Hymenophyllum plumo- sionally reaching length. Although Hy-
sum are H. karstenianum and H. speciosum, which menophyllum multialatum has been thus far re-
have no lamellae on the veins, but are similar to ported from four departments in Peru, the great
the former in most other features, including the majority are represented by Biies's collections in
dense matting of tomentum obscuring the laminar Cuzco. For further discussion see H. tomentosum.
surface. Extreme care and patience must be ex-
Amazonas: Prov. Bagua, Cordillera Colan, Barbour
ercised to determine whether the low outgrowths
3740 (F, MO). Huanuco: Prov. Huanuco, 5 km from Car-
are present or lacking. In fact, the real importance pish, Tryon & Tryon 5318-B (GH). Pasco: Prov. Oxa-
of the presence or absence of lamellae bears more pampa, San Alberto, Cordillera de Yanachaga, van der
During this study of
careful analysis in the future. Werffet al. 8495 (MO). Cuzco: Valley of Rio Urubamba,
Biies A- 16 (F, GH, us). Montana de Calca, Biies 1916 (GH,
Peruvian species of Hymenophyllum, it has been
us), 1919, 1923, 1932 (us). Prov. La Convencion, Cor-
noted that size and frequency of these outgrowths, dillera Vilcabamba, Dudley 1 1 137 (GH, us).
on one or both surfaces, may be more variable
than earlier assumed, and thus may be of much
less taxonomic significance. 31. Hymenophyllum tomentosum Kunze, Lin-
naea 9: 107. 1834. TYPE: Peru, Huanuco,
Huanuco: Vilcabamba, Hacienda on Rio Chinchao, Pampayacu, 1829, Poeppig Diar. 1134 (ho-
Macbride 5146 (F, us). Junin: Schunke Hacienda above lotype, w?; isotype, BM?; frag., B!; probable
San Ramon, Killip & Smith 24840 (F, GH, us). Cuzco:
isotype, MO!; photo, us of MO).
Prov. Paucartambo, above Tambomayo, West 7108 (MO,
uc). Madre de Dies: Prov. Mania, Shintuya, Chavez 860
(MO). Puno: San Gaban, Lechler 2264 (B). Sphaerocionium tomentosum (Kunze) Presl, Hymen-
ophyllaceae 34. 1843.
Hymenophyllum fusagasugense J. W. Sturm, Bot. Zeit.
(Berlin) 1859: 297. 1859 (as "fusugasugense").
30. Hymenophyllum mult ialat urn Morton, Contr. TYPE: Colombia, Cundinamarca, "Fusugasuga"
U.S. Natl. Herb. 29: 185. 1947. TYPE: (Fusagazuga) Karsten (not located).
Peru,
Hymenophyllum fusagasugense var. aberrans Mor-
Cuzco, Prov. La Convencion, Alturas de Sieve,
ton, Contr. U.S. Natl. Herb. 29: 187. 1947 (as
Biies 1575 (holotype, us!; frag., GH!). "fusugasugense"). TYPE: Venezuela, Merida,
Tabay, Gehriger 584 (holotype, us!; isotypes, F!,
Sphaerocionium multialatum (Morton) Pic.-Ser., GH!).
Webbia28: 471. 1973.
Leaves indeterminate, to 50 cm long and 2.5 cm
Leaves indeterminate, to 50 cm long, 3-8(-15) broad. Petiole 0.5-0.6 mm
in diameter, nonalate,
cm broad. Petiole 0.4-0.7 mm in diameter, non- sparsely pubescent with mostly simple trichomes.
alate, sparsely hirsute with mostly simple tri- Lamina pinnate-pinnatifid, densely hirsute, the
chomes. Lamina pinnate-pinnatisect, hirsute, the rachis nonalate throughout. Pinnae numerous, ob-
rachis nonalate throughout. Pinnae numerous, long or narrow-deltoid, pinnatifid (sometimes
narrow-oblong or -deltoid, pinnatisect, sometimes deeply so), the ultimate segments simple or (oc-

irregularly and greatly elongate, the ultimate seg- casionally) bifid, the tissue and veins densely cov-
ments commonly bifid, the tissue and veins mod- ered with (and often obscured by) sessile to short-
erately provided with sessile to short-stalked ses- stalked trichomes. Veins abaxially bearing irreg-
sile trichomes. Veins bearing (especially abaxially) ularly interrupted, conspicuous winglike lamellae,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 73


these wings much smaller or sometimes lacking vided with light or dark brown, short-stalked, stel-

adaxially. Indusia about as broad as long, equaling late trichomes. Veins abaxially bearing small,
or slightly broader than the segment tips. scattered lamellae, these about as broad as long,
but lacking adaxially. Indusia slightly longer than
On
steep banks and tree trunks, 2750-3100 m, broad, usually as broad as the segment tips, re-
Amazonas, Huanuco. ceptacle not exserted.
Colombia; Venezuela; Peru.
This is distinguished from Hymenophyllum from trunks and branch-
In wet forests, pendent
multialatum and H. plumosum only by the char- es of trees,600-1300 m, Loreto and Huanuco.
acters used in the key. All three appear to be alike, Colombia; Ecuador; Peru.
except for the size and frequency of the lamellae There is little, beyond the key characters, to
along the veins and by the relative density of pu- distinguish this from Hymenophyllum pyrami-
bescence. Of these, H. plumosum (q.v.) is the most datum. Leaves of the latter are perhaps more ro-
distinct, in that the lamellae are merely low and bust, with relatively narrower pinnae and much
inconspicuous outgrowths and the trichomes are denser pubescence. Both also share nearly the same
so densely matted as to completely obscure the range, and more study may prove them to be con-
veins and tissue. In H. tomentosum the indument specific. Also very closely related is H. verecun-
isalso densely matted, but the lamellae are so large dum, which is more easily separated by its much
and conspicuous that many of them emerge through lighter coloredtrichomes and shorter, relatively
the tomentum. Morton (1947) chose to separate broader and more crowded pinnae. But even these
H. fusagasugense and var. aberrans on the pres- characters sometimes tend to be intermediate.
ence or absence of these processes adaxially. How-
ever, the size or lack of lamellae on the adaxial Loreto: Pumayucu, between Balsapuerto and Moy-
side varies considerably in the species, and the obamba, Klug 3248 (B, F, GH, MO, us). Huanuco: Prov.
character is difficult to observe due to the dense
Huanuco, Dist. Churubamba, Mt. Santo Toribio, Mexia
8258 (B, F, GH, uc, us).
surface tomentum. To prove that the lamellae are
completely absent on a given specimen would re-
quire scraping away all the tomentum of each leaf.
33. Hymenophyllum pyramidatum Desv., Mem.
A thorough study of the species complex through- Soc. Linn. Paris 6: 332. 1827. TYPE: "habitat
out its range is needed.
inAmerica" (possibly a Dombey collection
from Peru) (holotype, P; photos, GH, us).
Amazonas: Prov. Chachapoyas, Cerros de Calla Calla,
19 km above Leimebamba, Hutchison & Wright 5570
(uc). Prov. Chachapoyas, near km 422, Balsas-Leime- Sphaerocionium pyramidatum (Desv.) Copel., Phil-
bamba road, Wurdack 1249 (F, GH, uc, us). ipp. J. Sci. 67: 30. 1938.

Leaves indeterminate, to 65 cm long and 6 cm


32. Hymenophyllum lobatoalatum Klotzsch, Lin- broad. Petiole 0.5-0.7 mm
in diameter, often alate

naea 20: 438. 1847. TYPE: Peru, Panatahua, 1-2 cmnear the apex, pubescent with simple to
Ruiz 83 (holotype, stellate trichomes. Lamina pinnate-pinnatifid,
B!; isotype, MA?).
densely pubescent with simple to stellate tri-

Sphaerocionium lobatoalatum (Klotzsch) Pic.-Ser., chomes, the rachis alate throughout. Pinnae nu-
Webbia28:471. 1973. merous, larger ones 20-45 mm
long, 6-1 1 mm
broad at base, most of them narrow-triangular,
Leaves indeterminate, to 60 cm long and 8 cm the apex narrowly acute to attenuate, deeply lobed
broad. Petiole 0.4-0.7 mm
in diameter, alate, if to pinnatifid, the 6-12 pairs of ultimate segments
at all, only near base of lamina, pubescent with simple to bifid, the tissue and veins usually co-
simple to stellate trichomes or glabrate. Lamina piously provided with orange to dark brown (rare-
pinnate-pinnatifid, pubescent with stellate tri- ly tawny), subsessile to short-stalked, stellate
chomes, the rachis strongly alate throughout. Pin- trichomes. Veins abaxially with numerous, con-
nae numerous, larger ones 20-45 long, 7-1 1 mm spicuous, elongate, winglike lamellae (often nearly
mm broad at base, most of them narrow-trian- the length of the vein), these sometimes less con-
gular, the apex narrowly acute to subattenuate, spicuous and abundant on the adaxial surface. In-
deeply pinnatind, the 7-12 pairs of ultimate seg- dusia slightly longer than broad, usually as broad
ments bifid, the tissue and veins moderately pro- as the segment tips, receptacle not exserted.

74 FIELDIANA: BOTANY
In dense, wet forests, pendent from stumps, tree (us). Madre de Dios: Prov. Manii, Shintuya, Chavez 833
and branches, and on wet banks, 600-1 800 p.p. (MO). Puno: Prov. Sandia, E of Oconeque, Hodge
trunks,
6090 (F, GH, us).
m, along Cordillera Central from San Martin to
Cuzco.
Colombia; Peru; Bolivia. 35. Hymenophyllum tarapotense Stolze, sp. nov.
This belongs to a close-knit complex of species
involving Hymenophyllum verecundum and H. Folia usque ad 6 cm longa et 1.6 cm lata. Petiolus 1-
lobatoalatum. See discussion of the latter for fur- 2 cm loneus. 0.1-0.15 mm
diametro. Lamina pinnato-
ther comments. pinnatifida. Rhachis versus basin non alata. Pinnae 6-
12-jugatae, bifidae vel pinnatifidae, usque ad 1 cm lon-
San Martin: Tingo Maria, Allard 20818 (us). Huan- gae, pinnae basales admodum reductae, marginibus, venis
et interveniis trichomatibus stellatis instructis. Venae
uco: Pampayacu. Kanehira 123 (F, GH, K, us). Pasco:
lamellis cristiformibus numerosis praeditae.
Prov. Oxapampa, 4-8 km from Enenas, Skoget al. 5077
(us). Junin: Chanchamayo Valley, C. Schunke 466, 926
(F.us), 7055 (F). Cuzco: Prov. Paucartambo, between Leaves indeterminate, to 6 cm long and 1.6 cm
Mistiana and Keros, Vargas 7380 p.p. (MO). broad. Petiole 1-2 cm long, 0.1-0.15 in di- mm
ameter, nonalate, sparsely provided with simple
trichomes. Lamina pinnate-pinnatifid, pubescent
34. Hymenophyllum verecundum Morton, Contr. with stellate trichomes, the rachis weakly alate dis-
U.S. Natl. Herb. 29: 183. 1947. TYPE: Peru, nonalate toward the base. Pinnae 6-12 pairs,
tally,
Huanuco, Churubamba, Mexia 8143-A (ho- bifid, or distal ones pinnatifid with 3-4 segments,
lotype, us!; isotypes, GH!, MO!, uc!). larger ones 1 cm long, basal ones often reduced to
mere auricles, the margins, veins, and intervening
Leaves indeterminate, to 50 cm long and 4 cm tissuemoderately to amply provided with tawny
broad. Petiole 0.4-0.6 mm in diameter, alate, if trichomes. Veins bearing numerous, conspicuous
at all, just below the lamina, glabrate or slightly accessory foliar crests not in the plane of the lam-
pubescent with simple to stellate trichomes. Lam- ina, these less frequent on the adaxial surface. In-
ina pinnate-pinnatifid, densely pubescent with dusia about as long as broad, as broad as the seg-
stellate trichomes, the rachis alate throughout, or ment tips, receptacle minute, not exserted.
sometimes nonalate at base. Pinnae numerous,
larger ones6-20 mm
long, 3-7 mm
broad at base, TYPE Peru, Tarapoto (Dept. San Martin) in
oblong to very broadly triangular, the apex obtuse monte Guayrapurima, Aug. 1856, Spruce 4048
to subacute, pinnatifid, the 3-6(-8) pairs of ulti- Another Spruce specimen
(holotype, K!; photo, F).
mate segments simple to retuse or occasionally at K, with notation "conf. 4048," is H. simplex.
bifid, the tissue and veins copiously provided with Known only from the type.
whitish to tawny, subsessile or short-stalked, stel- This tiny fern is unquestionably the smallest of
late trichomes. Veins abaxially with numerous the species in subg. Leptocionium which bear ac-
conspicuous, elongate, winglike lamellae, these less cessory tissue on their veins. These foliar processes
conspicuous and abundant on the adaxial surface. are not long, low wings as seen in Hymenophyllum
Indusia about as long as broad and usually as broad of
pyramidatum, but are separate, crestlike flanges
as the segment tips, receptacle not exserted. tissue similar to those frequently seen in more
delicate specimens of//, verecundum. Superficially
from tree trunks,
In dense, wet forests, pendent //.tarapotense closely resembles H. fragile, in its
branches, and wet banks, (5 50-) 1500-2 700 m, small size, delicate petiole, and reduced, often bi-
Amazonas, Huanuco, Junin, Cuzco, Madre de fid, pinnae. It may seem strange that this distinc-
Dios, Puno. tive species has not been described before, nor
Colombia; Peru; Bolivia. collected again in over 1 00 years. However, many
This is closely related to Hymenophyllum pyr- of these diminutive members of the genus are very
amidatum and H. lobatoalatum. See discussion of easily overlooked by collectors.
the latter for further comments.

Amazonas: Prov. Bagua. Cordillera Colan, Barbour Comments


3740 (MO in part). Huanuco: Rio Llullapichis watershed,
ascent of Cerros del Sira, Dudley 13370 (GH, us). Junin:
Pichis Trail. Porvenir, Killip & Smith 25948 (F, GH, us). Hymenophyllum peruvianum Hooker & Grev.,
Cuzco: Hacienda Pintobamba, Chaupimayo, Biies 1943 Icon. fil. 2, /. 208. 1831. TYPE: Ecuador (as

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 75


Peru), Prov. Esmeraldas, Jameson (holotype, In the original description the type was cited as
K). having come from Peru, but Esmeraldas is an Ecu-
adorean province in which Jameson did much of
The original drawing seems to illustrate most of his collecting. Yet Hymenophyllum trifidum still
the features characteristic of Hymenophyllum fu- might be expected in Peru, as it could be merely
coides var. pedicellatum, except that the apex of a form of H. elegans. From the description and
the indusium is deeply dentate rather than lacin- plate it appears to be very similar to the latter, but
iate. It is likely that the two taxa are synonymous, until the type is located the question must remain
but we have not seen the type to verify this. The unanswered.
problem is but one of many in the H. fucoides
complex begging solution through a complete re-
vision of the group. II. Trichomanes

Trichomanes L., Sp. pi. 1097. 1753. TYPE: Tri-


Hymenophyllum poeppigianum Presl, Hymeno-
TYPE: chomanes scandens L. Figure 12.
phyllaceae 54. 1843. Peru, Huanuco,
Pampayacu, Poeppig (holotype, PR; probable
isotype, P; photos, GH & us of P). Didymoglossum Desv., Mem. Soc. Linn. Paris 6: 330.
1827. TYPE: Didymoglossum muscoides (Sw.)
Desv. (Trichomanes muscoides Sw.) = Trichom-
Some
authors have placed this with Hymeno- anes hymenoides Hedwig.
phyllum polyanthos. The type has not been seen, Trichomanes subg. Didymoglossum (Desv.) C. Chr.,
but judging from the protologue and photos of a Index fil. xiv. 1906.
Trichomanes subg. Achomanes Presl, Hymenophyl-
probable isotype (P) the species is more likely to
laceae 15. 1843. TYPE: Trichomanes crispum L.
be H. myriocarpum. The protologue refers to: the Trichomanes subg. Pachychaetum Presl, Hymeno-
indusia "ovate-subrotund"; the rachis alate phyllaceae 16. 1843. TYPE: Trichomanes rigi-
throughout; the petiole alate toward the apex; the dum Sw.
lamina linear-lanceolate with the base acute. The Homoeotes Presl, Gefassbiindel Farm, 23. 1847.
TYPE: Homoeotes heterophylla Presl = Trichom-
description of the indusia could conceivably fit anes humboldtii Lell.
either H. polyanthos or H. myriocarpum, but that
of the lamina (as well as the photo of the possible
Plants epiphytic, occasionally epipetric or ter-
isotype) more closely match H. myricarpum or H. restrial. Stem stout and erect or decumbent to long-
undulatum. However, the petiole and (usually) the
creeping and filiform, bearing scattered to abun-
rachis base of H. undulatum are nonalate.
dant trichomes and delicate to large, fibrous roots
or (especially in subg. Didymoglossum) lacking
Hymenophyllum sprucei Baker in Hooker Bak- & roots. Leaves monomorphic or (less commonly)

er, Syn. fil. 65. 1 867. TYPE: Peru, San Martin, dimorphic, 0.540 cm or longer. Lamina entire to
Tarapoto, Spruce (not located). nearly 5 -pinnate, subsessile to petiolate, with pet-
iole commonly shorter than the lamina, glabrous
Morton (p. 155, 1947) considered that this is or with trichomes borne mostly on costae and
the same as Hymenophyllum adiantoides, and there margins. Veins free or, in Trichomanes diversi-
is nothing in the description to argue otherwise. It frons, reticulate near the lamina margin, pinnate
is possible that Baker was unaware of the publi- (anadromous or catadromous) or sometimes fla-
cation of the latter species, and even that he based bellate, false veins also present in some species,
his new species on the same specimen cited earlier these mostly parallel, but in a few species perpen-
by van den Bosch. Specimens (B, GH, K, L, NY, us) dicular to the true veins. Sori terminal on the veins.
thought to be probable isotypes of H. adiantoides Indusium mostly tubular, the mouth bilabiate or
are those of Spruce from Tarapoto. They bear the entire and often
flaring, or very rarely nearly bi-
notation "conf. 4700" and are conceivably the ones valved, the sporangia borne on an elongate recep-
on which Baker based his "//. sprucei." tacle. Receptacle commonly exserted beyond the
mouth of the indusium, often greatly so.

Hymenophyllum trifidum Hooker & Grev., Icon, There are about 300 species in the genus, oc-
fil. 2, 1. 196. 1 83 1 TYPE: Ecuador (as "Peru"),
.
curring in tropical to subtropical regions of both
Esmeraldas, Jameson (E?). hemispheres. As in Hymenophyllum, most are

76 FIELDIANA: BOTANY
FIG. 12. Trichomanes radicans: a, habit; b, pinna with sori. Trichomanes pinnatum: c, portion of pinna with false
veins and sori. Trichomanes hymenoides: d, apical portion of leaf, (a from Acosta-Solis 13782, Ecuador, F, b from
Malhias & Taylor 5075, F, c from Barbour 5194, F, d from Wurdack 1513, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 77


found as epiphytes in dark, wet forests, but a num- to the plane of the lamina. One of these, T. mar-
ber of species may be found on wet, rocky cliffs tiusii, occurs in Peru.
or occasionally on clay banks. Also as in Hymen- The group of Trichomanes crispum L. in subg.
ophyllum, Trichomanes has been divided by pre- Achomanes is the most taxonomically difficult in
vious authors into a number of genera and/or sub- the genus, and a few of these species occurring in
genera. Only one genus is recognized here, with Peru may eventually be found to merit only infra-
the key and species order reflecting as nearly as specific status. Some of the difficulties are very
possible four of the subgenera. The monotypic subg. likely the result of yet unsolved questions involv-
Cardiomanes (Presl) Christ occurs in New Zea- ing hybridization.
land.
"False veins" may be found on laminae in two
of the subgenera. These are long sclerenchymatous References
strands which can be observed as faint to rather
distinct lines, parallel or perpendicular to the true BOER, J. G. W. 1962. The new world species of
veins. They may be connected to the true veins Trichomanes sect. Didymoglossum and Micro-
but are much more commonly free from them. gonium. Acta Hot. Neerl., 11: 277-330.
Their position and frequency are important char- WINDISCH, P. G. 1977. Synopsis of the genus
acters in recognizing some of the species. As in Trichomanes, subgenus Achomanes. Ph.D. the-
Hymenophyllum, veins of several species of Tri- sis, Department of Biology, Harvard Univer-
chomanes bear crestlike lamellae, perpendicular sity, Cambridge, Mass., 198 pages.

Key to Species of Trichomanes

a. Venation anadromous, never flabellate; false veins lacking b


b. Stem long-creeping; leaves well-spaced to remote [subg. Trichomanes] c
c. Stems relatively stout, 0.7-2.2 mm in diameter; mature leaves 15-50 cm long d
d. Pinnae 2-3-pinnatisect; stem amply to densely provided with castaneous trichomes; ultimate
segments narrow, commonly linear e
e. Indusium narrow-cylindrical or -elliptic, often subfusiform, the apex not or scarcely

expanded; petiole (1 )4 15 cm long; elevation 0-500 m 1. T. radicans


e. Indusium apex expanded into a wide-flaring mouth; petiole 0.3-3 cm long;
conical,
elevation (400-)700-3200 m 2. T. collariatum

d. Pinnae pinnatifid or pinnatisect (sometimes 2-pinnatifid as to basal segments); stem with


trichomes sparse to widely scattered; ultimate segments relatively short and broad, not
linear 3. T. rupestre
c. Stems filiform, 0.2-0.5 mm in diameter, leaves 3-15(-18) cm long f
f. Rachis alate throughout; pinnae commonly adnate, the costae alate to base (rarely nonalate
just at base) g
g. Tube of indusium broadly conical, 1-1.5 times as long as broad, mouth not or slightly

flaring; tissue of most ultimate segments with elongate, narrow folds parallel to the veins
4. T. pyxidiferum

g. Tube of indusium narrowly cylindrical; 24 times as long as broad,


mouth abruptly and
widely flaring; tissue of
segments essentially plane (margin sometimes undulate)
5. T. diaphanum

f. Rachis nonalate nearly to the apex; pinnae stalked, the costae not alate at base h
h. Costae alate, except not below the basal pinnule; ultimate segments 0.5-0.8 broad; mm
sori 1-5 per pinna; indusia commonly alate, the wings several cells wide
6. T. angustatum
h. Costae nonalate except near the apex; ultimate segments 0.1-0.4 mm broad; sori rarely
more than 1 per pinna; indusia not or scarcely alate, wings (if any) 1 cell wide
7. T. capillaceum
b. Stem erect, or occasionally short-creeping; leaves caespitose to (occasionally) approximate [subg.
Pachychaetum] i

78 FIELDIANA: BOTANY
i. Tissue of segments opaque, the cells small and occluded, not evident when observed with
magnification of 8-12 x; leaves (mature ones) 15-90 cm long; petiole (0.6-)0.7-3.5 mm in
diameter j

j. Rachis obtusely quadrangular, alate (often broadly so) at least in the distal half; pinnae
adnate, the costae alate; lamina usually more than 1 cell thick, at least away from margins
8. T. elegans

j. Rachis or marginate to slightly alate near apex; pinnae (at least proximal
terete, nonalate,

ones) short-stalked, costae not alate at base; lamina essentially 1 cell thick throughout . . .

9. T. rigidum
i. Tissue of segments translucent, the cells large and clear, quite evident with magnification of
8-12 x; leaves commonly less than 15 cm long; petiole 0.3-0.6 mm
in diameter
10. T. cellulosum
a. Venation at least partly sometimes flabellate); false veins present or lacking
catadromous (or k . . .

k. False veins present, these submarginal or borne between and parallel with the true veins [subg.
Didymoglossum] 1

1. Leaves with simple to stellate, commonly dark, trichomes along the margins; indusia distinctly
bilabiate and the lips commonly dark-margined m
m. Venation flabellate, a percurrent costa lacking 11. T. punctatum ssp. sphenoides
m. Venation pinnate, or if subflabellate, the costa essentially percurrent n
n. Mature leaves entire, or distally lobed; stellate trichomes borne all along the margin
1 2. T. angustifrons
n. Mature leaves 1-2-pinnatifid; stellate trichomes confined to segment sinuses, with simple
or bifid trichomes borne on outer margin o
o. Son protruding from the leaf tissue, or slightly immersed at base; false veinlets

extending toward the lamina margin, but rarely running parallel to it p


p. False veinlets sparse, scattered; lips of indusium mostly broader than long; cells
of lamina mostly isodiametric 13. T. hymenoides

p. False veinlets commonly abundant; lips of indusium mostly as long as or longer


than broad; cells of lamina mostly elongate 14. T. reptans
o. Sori partly to deeply immersed in the leaf tissue, or at least broadly alate; false
veinlets (many of them) parallel to and very near the margin 15. T. krausii
1. Leaves with margins glabrous or provided with circular scalelike processes (these sometimes
deciduous in age); indusia not or scarcely bilabiate, the mouth not dark-margined q
q. Lamina lacking paired, circular scalelike processes, but bearing a peripheral, submarginal
false vein; venation essentially pinnate 16. T. kapplerianum

q. Lamina bearing numerous paired, circular, scalelike processes all along the margin (these
sometimes deciduous in age), but lacking a submarginal false vein; venation flabellate . . .

1 7. T. membranaceum
k. False veins lacking or, in T. pinnatum, perpendicular to the true veins (some short, veinlike lines
in sect. Lacostea); [subg. Achomanes] r

r. Lamina trichomes stellate or forked from the base; stem filiform, 0.3-0.4 mm in diameter
18. T. polypodioides
r. Lamina trichomes simple or lacking; stem relatively stout and wiry, 0.5-1.2 mm in diameter
s

s. Stem long-creeping up tree trunks, leaves subdistant to remote; lamina appressed to tree
trunks by means of rhizoids on axes t

t. Lamina linear, subentire to lobed, 1-1.5 cm broad; indusia mouth rather widely flaring
1 9. T. tanaicum
t. Lamina oblong, pinnatisect or nearly pinnate, mature ones ( 1 .8-)2-9 cm broad; indusium
mouth not or slightly flaring u
u. Segments of sterile laminae entire or shallowly crenate; veins simple, not forked;
indusium mouth not expanded; rare in Peru 20. T. tuerckheimii
u. Segments of sterile laminae deeply crenate or crenate-serrate; veins (some or many)
1 -forked; indusium mouth slightly flaring; locally common 21. T. ankersii

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 79


s. Stem erect to decumbent and the leaves crowded to caespitose, or if in some species long-
creeping and the leaves subdistant, then the lamina not appressed to tree trunks by rhizoids
on leaf axes v
v. Leaves dimorphic w
w. Stem decumbent; leaves crowded to caespitose; fertile leaves simple and
erect or

essentially entire, or without laminar tissue and the sori completely free x
x. Fertile leaves with lamina entire to pinnatisect; sori fully immersed in tissue y . .

y. Sterile lamina with 4-8 veins issuing from the rachis between adjacent costae;
mouths of indusia neither flaring nor recurved, strongly indented between the
vein branches 23. T. diversifrons

y. Sterile lamina with 2(-3) veins issuing from the rachis between adjacent costae;
mouths of indusia slightly flaring and recurved, but not indented between the
vein branches 24. T. trollii
x. Fertile leaves without laminar tissue; sori completely free 25. T. botryoides
. . .

w. Stem wiry, long- or short-creeping; leaves subdistant to remote; fertile leaves with
lamina foliose and deeply pinnatisect to nearly pinnate 28. T. humboldtii
v. Leaves monomorphic z
z. Lamina 3-4-pinnatisect; petiole conspicuously alate throughout, the alae broader than
the petiole quite or nearly to base 22. T. bicorne
z. Lamina not more than 1 -pinnate-pinnatisect; petiole nonalate in the proximal half
or throughout, or very narrowly alate in the proximal portion aa
aa. Lamina abruptly terminating in a conform apical segment or the rachis prolonged,

flagellate, and proliferous at tip; indusia fully exserted from segment margin, often
stalked bb
bb. False veins copious and perpendicular to the true
veins; marginal vein strong,
continuous; pinna margins sharply serrate to spinulose .26. T. pinnatum .

bb. False veins rare or lacking; marginal vein lacking or faint and discontinuous,
pinna margins commonly obtusely serrate 27. T. hostmannianum
aa. Lamina gradually reduced to a pinnatifid apex, the rachis not prolonged and
flagellate; indusia at least partially immersed in the laminar tissue, wholly or
partially alate on each side cc
cc. Pinnae regularly and deeply lobed to pinnatisect dd
dd. Pinnae lobed to pinnatifid; lamina l-2(-2.5) cm broad, not at all obscured
by the moderate pubescence 29. T. crinitum
dd. Pinnae pinnatisect (rarely the segments again lobed); lamina (2.5-)3-12
cm broad, partially to completely obscured by the dense covering of
trichomes 31. T. lucens
cc. Pinnae entire to serrate or crenate (or rarely a few with some irregularly
scattered lobes) ee
ee. Veins bearing crestlike lamellae on abaxial side; pinnae in proximal por-
tion of lamina deflexed 30. T. martiusii
ee. Veins lacking lamellae; pinnae patent or a few basal ones deflexed ff . .

ff. Petiole conspicuously alate halfway or more to the stem; stem erect
to decumbent, leaves contiguous to caespitose .... 32. T. pellucens
ff. Petiole not alate, or scarcely so near apex due to decurrent lamina
base; stem short-creeping or decumbent, the leaves subdistant to con-
tiguous (rarely caespitose) gg
gg. Rachis trichomes on abaxial side predominantly dark brown, stout,
rigid, most of them terete toward their base 33. T. plumosum
. .

gg. Rachis trichomes tawny to orange, delicate, tortuous, the cells

flattened hh
hh. Lamina mature ones commonly
linear or linear-lanceolate,
5-12 times as long as broad; mature leaves (10-) 15-50 cm

80 FIELDIANA: BOTANY
long; rachis abaxially provided with trichomes predominantly
unicellular beyond the basal one, occasionally with 2-3 cells
34. T. cristatum
hh. Lamina oblong- to ovate-lanceolate, 3-4 times as long as
broad; mature leaves 5-15 cm long; rachis abaxially provided
with trichomes of 2-8 cells beyond basal one, as well as with
many unicellular ones 35. T. vandenboschii

1. Trichomanes radicans Sw., J. Hot. (Schrader) rupestre form a close-knit species complex which
1800(2): 97. 1802. TYPE: Jamaica, Swartz is need of closer study. The three are only pro-
in

(holotype, s; photo, us). Figure 12a-b. visionally maintained here at the species level. In
Central America T. radicans and T. collariatum

Trichomanes kunzeanum Hooker, Sp. fil. 1: 127. 1844.


are more distinct, e.g., leaves of the latter are con-
SYNTYPES: Peru, Huanuco, Pampayacu, Poep- sistently reduced to a very short and broadly alate
pig 1132 (K!; photo, us); Peru, Junin, Pangoa, petiole; whereas in T. radicans leaves are not or
Mathews 1088 (K.!; photos. A, us); Venezuela,
scarcely reduced at base, and petioles are much
Merida, Linden 176 (K.!; isosyntypes, BM, n).
Trichomanes brachyblastos Kuhn, Linnaea 35: 388. longer and not or very narrowly alate or marginate.
1868. TYPE: Peru, San Martin, Tarapoto, Monte These characters are also somewhat diagnostic in
Guayrapurima, Spruce 4703 (holotype, B?; iso- South America, but are much more variable, and
types, BM!, BR!, GH!, K!, w!). are therefore not thoroughly reliable. In Peru, often
Vandenboschia radicans (Sw.) Copel., Philipp. J. Sci.
the two species can be positively separated only
67: 54. 1938.
Trichomanes radicans var. kunzeanum (Hooker) Duek by the soral features.
& Lell., Amer. Fern J. 68: 120. 1978.
San Martin: Prov. Mariscal Caceres, Dist. Tocache
Nuevo, J. Schunke 4361 (F, us). Huanuco: Near Tingo
Stem long-creeping, relatively stout, 1-2.2 mm Maria, confluence of Monzon and Huallaga Rivers, Stork
in diameter, amply to densely covered with cas- & Norton 9501 (F, GH, MO, uc, us). Pasco: Paujil near
taneous trichomes. Leaves remote, 20-50 cm long, Puerto Bermudez, Leon 288 (GH, USM). Junin: Chan-
axes often provided with scattered septate tri- chamayo Valley, C. Schunke 459 (F, us). Ucayali: Prov.
Coronel Portillo, Sinchono, between Tingo Maria and
chomes abaxially. Petiole (l-)4-15 cm long, 0.8-
Pucallpa, Aguilar 883 (GH). Ayacucho: Prov. La Mar,
2 mm in diameter, nonalate to marginate or alate between Tambo San Miguel, Ayna and Hacienda Lu-
(often the wings deciduous). Lamina 2-3-pinna- isiana, Dudley 11904 (GH). Cuzco: Prov. Paucartambo,
ovate or deltoid-ovate and scarcely reduced
tisect, bosques de Keros, Vargas 7382 (uc). Puno: Prov. Car-
at base, or occasionally elliptic and somewhat abaya, Hacienda Palmera, Vargas 16137 (GH).

strongly reduced toward base. Rachis narrowly to


broadly alate. Pinnae mostly 2-pinnatisect, slight-
ly to strongly adnate to the rachis. Ultimate seg-
2. Trichomanes collariatum Bosch, Ned. Kruidk.
ments commonly narrow, often with simple
linear,
Arch 4: 368. 1859. TYPE: Mexico, Tabasco,
or 1 -forked veins, false veinlets lacking. Venation
Linden (holotype, L?; probable isotype, K;
anadromous. Indusia very slender, cylindrical or
photo, us of K).
subconical, often subfusiform, not
elliptic, rarely
or scarcely alate, the wings of tissue (if any) 1-
Stem
long-creeping, relatively stout, 1-1.7 mm
2(-3) cells wide, apex truncate, not expanded, re-
in diameter, amply to densely covered with cas-
ceptacle short- or long-exserted.
taneous trichomes. Leaves remote, 1 5-38(-45) cm
long, axes often provided with scattered, septate
Hemiepiphytic on tree trunks in dense, wet for- trichomes abaxially. Petiole 0.3-3 cm long, 0.7-
ests, (400-)700-3200 m, San Martin to Puno. 1.5 mm in diameter, broadly alate nearly or quite
Mexico to Panama; West Indies; Guianas to Co- throughout. Lamina 2-3-pinnatisect, oblong or
lombia and southward to Brazil and Paraguay; Old oblong-lanceolate, often abruptly reduced at base.
World. Rachis commonly broad-alate throughout. Pinnae
Trichomanes radicans, T. collariatum, and T. mostly 2-pinnatisect, slightly to strongly adnate to

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 81


rachis. Ultimate segments commonly narrow, often shape of indusia. The indusium in the latter is
linear, with simple or forked veins, false veinlets conical, with a conspicuous and widely flaring
lacking. Venation anadromous. Indusia conical, mouth; in T. rupestre it is narrower, with mouth
gradually widening from base to a conspicuously only slightly flaring, and in T. radicans the indu-
flaring mouth, alate, the wings of tissue on each sium is even more slender, often narrowing slight-
side 4-6 cells wide, receptacle long-exserted. ly, with the apex truncate, not or scarcely spread-

ing. See T. radicans for further discussion.


Hemiepiphyte on tree trunks in wet forests, sea
level to 500 m, San Martin, Loreto, Madre de Huanuco: Prov. Huanuco, Dist. Churubamba, Cotir-
Dios. arda, Mexia 8221 (F, GH, MO, uc, us). Tingo Maria, Selva
Real, Morrow 11132 (GH). Ucayali: Prov. Coronel Por-
Southern Mexico to Panama; British Guiana to
tillo (as Loreto), Aguaytia, Ridoutt 13082 (USM).
Colombia, south to Peru; Brazil.
This is not greatly distinct from T. radicans,
under which see further discussion.
4. Trichomanes pyxidiferum L., Sp.pl. 1098. 1753.
TYPE: based on Plumier, Traite foug. Amer.,
San Martin: Prov. Mariscal Caceres, Dist. Campan-
/. 50E, 1705, illustrating a plant from Haiti.
margin of Rio Huallaga, J. Schunke 4120 (F, GH,
illa, left

us). Loreto: Soledad, on Rio Ilaya, Killip & Smith 29739


(us). Madre de Dios: Prov. Tambopata, Lago Tres Chim- Vandenboschia pyxidifera (L.) Copel., Philipp. J. Sci.
badas, Barbour 5669 (MO). Parque Nacional del Manu, 67: 53. 1938.
Cocha Cashu Biological Station, M. S. Foster P-84-4
(uc), P-84-67 (MO); R. Foster 6868 (F). mm in di-
Stem long-creeping, filiform, 0.4-0.5
ameter (excluding indument), densely covered with
blackish trichomes. Leaves well-spaced to remote,
3-12 cm long, axes sometimes provided abaxially
3. Trichomanes rupestre (Raddi) Bosch, Ned.
with scattered, minute trichomes. Petiole 1-3 cm
Kruidk. Arch. 4: 370. 1859.
long, 0.3-0.6 mm in diameter, flattened, or terete
PI. Bras. nov. gen.
at narrowly to broadly alate distally or
base,
Hymenophyllum rupestre Raddi,
1 :
67, /. 825. TYPE: Brazil, Raddi (holotype,
80. 1 throughout, the wings plane to somewhat crispate.
FI?; isotype, K.; photos, A us of K). & Lamina 2-3-pinnatifid. Rachis alate throughout,
the wings commonly undulate to crispate. Pinnae
Stem
long-creeping, stout, 0.7-1.2 in di- mm 4-10 pairs, strongly adnate to the rachis (the costae
ameter, provided with a few scattered to sparse alate). Ultimate segments linear, margins plane to
castaneous trichomes. Leaves remote, mature ones undulate, tissue commonly with elongate, narrow
15-30 cm long, axes sometimes provided with folds parallel to the veins, false veinlets lacking.
scattered, septate trichomes on abaxial side. Pet- Venation anadromous. Sori 1 -several per pinna.

iole 0.2-2 cm long, 0.6-0.8 mm in diameter, par- Indusia broadly conical, the tube 1-1.5 times as
tially alate. Lamina commonly 1-pinnate-pinna- long as broad, not or scarcely bilabiate, the mouth
oblong or elliptic-oblong. Rachis broadly alate
tifid, not or slightly flaring, receptacle commonly long-
throughout. Pinnae pinnatifid or pinnatisect, or exserted.
rarely 2-pinnatifid as to basal segments, strongly
adnate to rachis. Ultimate segments relatively short On trees in wet forests, 900-1 200 m, Amazonas,
and broad, not linear, with simple or forked veins, San Martin.
false veinlets lacking. Venation anadromous. In- Pantropical.
dusia conical, gradually widening from base to a A distinctive feature of Trichomanes pyxidifer-
slightly flaring mouth, narrowly alate on each side, um is the rather sharp, longitudinal folding of seg-
receptacle short- to long-exserted. ment tissue parallel to the veins. Other related
species have plane segments, or some with the
Creeping on tree trunks or wet banks in dense, margins undulate, but none exhibit this unique
wet forests, 1 100-1600 m, Huanuco and Ucayali. character. Perhaps it was this which prompted
Venezuela and Colombia; Peru; Bolivia; Brazil Morton's incorrect observation (1968) of false veins
(also reported from Costa Rica by Alan Smith, in. in the species, since the sharper folds of tissue

litt). sometimes cause dark shadow lines in the seg-


This is not as common as Trichomanes radicans ments. However, in careful study of specimens
or T. collariatum and is midway between them in throughout the range, no leaves were seen with the

82 FIELDIANA: BOTANY
sclerenchymatous strands which produce these axes and indusia, crowding of pinnae and dissec-
"false veins." The species thus far is represented tion of lamina. Each of these may appear to be
in Peru by only two collections, even though its important in individual specimens, but when many
full range extends throughout tropical regions of collections are examined throughout the range of
the world. distribution, it becomes evident that they are quite
variable and uncorrelated.
Amazonas: Prov. Chachapoyas, .la/an (Ingenio-
Chachapoyas), Ldpe: el al. 4264 (GH, HUT). San Martin: Amazonas: Prov. Bagua, Cordillera Colan SE of La
Tarapoto, Spruce 4761 (BR, OH, K, L, us). Peca, Harbour 3 904 (MO). I .ore to: Sierra del Pongo, Mex-
ia 6288B (GH, uc, us). Pasco: Prov. Oxapampa, Dist.
Oxapampa, road Oxapampa to Villa Rica, B. Ledn 660
(USM La Merced, E of Quimiri Bridge,
in part). Junin:
5. Trichomanes diaphanum HBK., Nov. gen. sp. Killip & Smith 24006 (F), 24023 (GH, us). Cuzco: Prov.
1:25 (fol. 1 6). 1 8 1 6. TYPE: Venezuela, Hum- La Convencion, "El Dorado," Vargas 3518 (GH). Puno:
boldt & Bonpland (holotype, p; isotype, B!).
San Gaban, Lechler (L).

Trichomanes leptophyllum Bosch, Ned. Kruidk. Arch.


4: 363. 1859 (not A. Cunn., 1836), nom. illeg.
TYPE: based on T. pyxidiferum Hooker & Grev. 6. Trichomanes angustatum Carm., Trans. Linn.
Trichomanes hymenophylloides Bosch, Ned. Kruidk. Soc. London 12: 513. 1818. TYPE: Tristan
Arch. 5: 209. 1863, nom. nov. for T. leptophyllum da Cunha, Carmichael (holotype, K!; isotype,
Bosch and type based on that name.
BM!).
Vandenboschia diaphana (HBK.) Copel., Philipp. J.
Sci. 67: 53. 1938.
Vandenboschia hymenophylloides (Bosch) Copel., Trichomanes tenerum Sprengel, Syst. veg. 4: 1 29. 1 827.
Philipp. J. Sci. 67: 53/1938. TYPE: Brazil, collector not stated (holotype, LZ
destroyed).
Vandenboschia tenera (Sprengel) Copel., Philipp.
Stem long-creeping, filiform, 0.2-0.5 mm in di- Sci. 67: 53. 1941.
J.

ameter (excluding indument), sparsely to abun- Vandenboschia angustata (Carm.) Copel., Philipp. J.

dantly provided with castaneous trichomes. Leaves Sci. 73: 466. 1941.

well-spaced to remote, 5-15(-18) cm long, axes


glabrous, or the rachis sometimes abaxially pro- Stem long-creeping, filiform, 0.2-0.3 mm in di-
vided with scattered, minute trichomes. Petiole ameter, amply provided with castaneous tri-

0.5-6 cm long, 0.3-0.8 mm in diameter, flattened, chomes. Leaves well-spaced to remote, 4-14(-18)
or terete at base, nonalate, or narrowly to broadly cm long, glabrous. Petiole 0.5-5 cm long, 0.2-0.4
alate for most of its length, the wings plane or mm in diameter, terete, nonalate. Lamina 2-pin-

occasionally undulate. Lamina essentially 3-pin- nate-pinnatifid to 4-pinnate, linear-lanceolate to


natifid.Rachis alate throughout, the wings narrow ovate-lanceolate, abruptly or gradually reduced at
to broad, plane to undulate or occasionally cris- base. Rachis nonalate, or scarcely alate near the
pate. Pinnae 4-12 pairs, adnate to the rachis, the apex. Pinnae 6-many pairs, short-stalked, the cos-
costae alate (rarely nonalate at base). Ultimate seg- tae alate except below the basal pinnule. Ultimate
ments linear, 0.6-0.9 mm
broad, plane, not folded, segments linear, 0.5-0.8 mm
broad, each bearing
Venation anadromous. Sori
false veinlets lacking. a single vein, false veinlets lacking. Venation anad-
1-10 per pinna. Indusia salverform, the alate tube romous. Sori 1-5 per pinna. Indusia narrowly fun-
narrowly cylindrical, 2-4 times as long as broad, nelform to salverform, the tube 2-4 times as long
abruptly expanding into a broad-flaring mouth, as broad, the mouth flaring, margins commonly
receptacle commonly long-exserted. alate, with wings several cells wide, receptacle long-
exserted.
On moist banks and (rarely) wet rocks, in
trees,
deep forests 700-2900 m, Amazonas and Loreto In deep forests, on trees, wet cliffs and banks,
to Puno. and on decaying logs, 600-2800 m, Cajamarca and
Southern Mexico to Panama; West Indies; Trin- San Martin to Ayacucho and Cuzco.
idad; the Guianas to Colombia, south to Peru and Southern Mexico to Honduras; Greater Antilles;
Brazil. Venezuela and Colombia to NW
Argentina and
Previous authors have separated Trichomanes Brazil.

hymenophylloides on characters such as number This is very closely related to Trichomanes cap-
of sori per pinna, width of tissue wings flanking illaceum, under which see further discussion.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 83


Cajamarca: Colasay, Woytkowski 7002 (us). San Mar- partments which both species are known to oc-
in
tin: Monte Campana, near Tarapoto, Spruce 4705 (BR, based only on collections from Peru,
cur. Therefore,
K, NY). Pasco: Prov. Oxapampa, San Alberto, Cordillera
one would be tempted to combine the two species,
de Yanachaga, van der Werffet al. 8428 (MO, uc). Junin:
but a thorough study of the characters of both taxa
Carpapata, above Huacapistana, Killip & Smith 24492
(F,uc, us). Ayacucho: Ccarrapa, between Huanta and throughout their range is necessary before further
Rio Apurimac, Killip & Smith 22406 (F, GH, us). Cuzco: classification is attempted.
Valley of Urubamba, Machu-Picchu, base of Huayna
Picchu, Iltis el al. 1064 (GH, MO, uc, us).
Pasco: Pichis Trail, San Nicolas (as Junin), Killip &
Smith 25990 (F, GH, us). Cuzco: Prov. Paucartambo,

Cosnipata Valley, Rio Tono, Wachter et al. 199 (F).


7. Trichomanescapillaceum L., Sp. pi. 1099. 1753.
LECTOTYPE (designated by Proctor, Ferns
of Jamaica, British Museum, London, p. 109. Trichomanes elegans Rich., Actes Soc. Hist.
1985): Plumier, Traite foug. Amer., t. 99D, Nat. Paris 1: 114. 1792. TYPE: French
based on a Plumier collection from Haiti. Guiana, Le Blond (P) (not T. elegans Rudge,
1805).
Vandenboschia capillacea (L.) Copel., Philipp. J. Sci.
67: 53. 1938.
Trichomanes prieurii Kunze, Analecta Pteridogr. 48.
1837. SYNTYPES: French Guiana, Leprieur (LZ
This differs from Trichomanes angustatum only destroyed; isosyntype, F!); Brazil, Amazonas, Rio
in the characters used in the key. Japura, Martius (M); British Guiana, Rudge (K).
Trichomanes opacum Bosch, Ned. Kruidk. Arch. 5(2):
1 75. 86 1 TYPE: Peru, Puno, San Gaban, Lech-
1 .

In deep forests, on tree trunks, 800-1100 m, ler 2175 (holotype, L?; isotype, P!; photo, F, frag.,
Pasco and Cuzco. L!).

Southern Mexico to Panama; Greater Antilles; Davalliopsis elegans (Rich.) Copel., Philipp. J. Sci. 67:
82. 1938.
Venezuela and Colombia to Peru.
With this might possibly be included Trichom-
anes angustatum, but the two are separated here Terrestrial, rarely epiphytic or epipetric. Stem
provisionally, pending much needed monographic Leaves caespitose to
erect or rarely short-creeping.
revision. Typically, T. capillaceum has very little occasionally approximate, 20-90 cm long, rachis,
expanded tissue along the axes, and even the ul- costae, and often costules amply to abundantly
timate segments are rarely more than 0.3 mm provided with castaneous, pluricellular trichomes
broad. Indusia are not or scarcely alate, and son on abaxial side. Petiole 10-40 cm long, 1-3.5 mm
are solitary on each fertile pinna. Conversely, be- in diameter, usually alate distally. Lamina 2-3-

yond the first pinnule in T. angustatum, the costae pinnate-pinnatifid, deltoid, opaque, usually more
and costules are obviously alate throughout, and than 1 cell thick, at least away from the margins,
ultimate segments are often nearly 1 mm broad. the cells small and occluded, not evident even when
The tissue also forms wings on each side of the magnified at 10-15x. Rachis obtusely quadran-
indusia, and sori are frequently four or five per gular, alate (often broadly so) at least in distal half.
pinna. Pinnae adnate, the costae alate to base. Ultimate
Even some of the characters listed in the key are segments laciniate, with linear lobes, each of these
subject to variation in certain geographic regions, bearing a single vein, false veins lacking. Venation
and intermediates are found frequently. Never- anadromous. Sori usually bent down away from
theless there are areas in which only one or the the plane of the lamina. Indusia subconical to
other occur. Only Trichomanes angustatum oc- somewhat urceolate, not bilabiate, the mouth
curs in Bolivia and southern Brazil, whereas in truncate or slightly (rarely strongly) flaring, recep-
Central America it has not been found south of tacle short- to long-exserted.
Guatemala, a region where T. capillaceum is rath-
er common. In Peru, the latter is rare, and even In dense, wet forests or wet, wooded ravines, on
the few specimens located are not typically skeletal ground, or rarely on tree trunks or wet rocks, sea
in their appearance, whereas Trichomanes angus- level to 1300 m, Amazonas and Loreto south to
tatum is and the specimens
rather widespread Madre de Dios and Puno.
rather typical. One specimen from Cuzco, Vargas Lesser Antilles and Trinidad; Nicaragua to Pan-
8653 (MO), is exactly intermediate between the ama; Guianas to Colombia and south to Bolivia
two; significantly, Cuzco is one of the two de- and Brazil.

84 FIELDIANA: BOTANY
With its large,blue green leaves and laciniate essentially in the plane of the lamina. Indusia sub-
pinnae, this is one of the most beautiful species in conical, not bilabiate, the mouth truncate, not or
the family. Its most distinctive features are the scarcely flaring, receptacle commonly long-exsert-
usually strongly arching sori (bent downward out ed.
of the plane of the lamina) and the relatively thick
tissue. Although the lamina in Hymenophyllaceae On slopes and ridges of dense forests and in

is one cell thick, the tissue in Tri- wooded ravines, occasionally on wet rocks or cliffs,
typically only
chomanes elegans is often several cells thick, es- rarely on bases of tree trunks, 350-2500 m, Ca-
pecially away from the segment margins. The jamarca and San Martin, south to Cuzco and Madre
de Dios.
broadly alate rachis traditionally has been used to
Southern Mexico to Panama; West Indies; South
separate this from T. rigidum (with which it often
grows), and although usually it is a good diagnostic
America, to Bolivia and Brazil; probably Old World
feature, occasional specimens of T. elegans can be tropics.

found throughout the range in which rachis alae This is sometimes confused with Trichomanes
are very narrow or lacking in the proximal portion. elegans, with which it often grows. For compari-
son, see discussion under the latter species.
Consequently, supporting key characters should
be employed for accurate identification.
Cajamarca: Tambillo, Jelski 891 (us). San Martin:
Tarapoto, in monte Guayrapurima, Spruce 4047 (BR, K).
Amazonas: Prov. Bagua, left bank of Rio Maranon Huanuco: East of Tingo Maria (as San Martin), Allard
opposite Quebrada Mirana, Wurdack 2031 (F, GH, us). 22356 (GH, us). Pasco: Oxapampa, Cordillera San Ma-
San Martin: Prov. Mariscal Caceres, Tocache Nuevo, J. tias, Leon 326a (USM). Junin: La Merced, Hacienda
Schunke4855 (F, GH, us), 6955 (F, MO in part, us). Loreto: Schunke, Macbride 5633 (F, us). Ucayali: Vicinity of
Sierra del Pongo, Mexia 1 195 (F, GH, MO, us). Huanuco:
Aguaytia, along Rio Aguaytia (as Loreto), Croat 20927
Prov. Pachitea, Dist. Honoria, /. Schunke 2259 (F, GH).
(MO). Cuzco: Prov. Urubamba, base of Huayna Picchu,
Pasco: Prov. Oxapampa, Valle del Palcazu, Iscozacin, Iltis et al. 1062 (GH). Madre de Dios: Prov. Manu, Cerro
Le6n 692 (F, GH). Junin: Puerto Bermudez, Killip & Smith de Pantiacolla, Foster 10903 (F).
26537 (GH, us). Ucayali: Prov. Coronel Portillo (as Lor-
eto), Padre Abad, J. Schunke 5393 (F, us). Cuzco: Bajada
de Tocate, Biies 1748 (us). Madre de Dios: Prov. Tam-
10. Trichomanes cellulosum Klotzsch, Linnaea 18:
bopata, Rio Tambopata Nature Reserve, Barbour 5165
(MO). Puno: Prov. Carabaya, San Gaban, Vargas 18864 531. 1844. TYPE: "British Guiana," (Guy-
(GH). ana). Kanuku Mountains, Rich. Schomburgk
1186 (holotype, B!; photo, F; isotypes, B! 2
sheets, BM!).
9. Trichomanes rigidum Swartz, Prodr. 137. 1788.
TYPE: Jamaica, Swartz (holotype, s; isotype, Epiphytic, terrestrial, or sometimes epipetric.
B, Herb. Willd. 20202- 7; photos, us of s, GH Stem erect to short-creeping. Leaves caespitose to
& us of B). (occasionally) approximate, 6-15 cm long, gla-
brous. Petiole 1-7 cm long, 0.3-0.6 mm
in di-
Selenodesmium rigidum (Sw.) Copel., Philipp. J. Sci. ameter, narrowly alate at least distally. Lamina 4-
67:81. 1938.
5-pinnate, lanceolate to ovate, translucent, the cells
largeand clear and quite evident when magnified
Terrestrial, rarely epiphytic or epipetric. Stem at 8-12x. Rachis terete, alate throughout. Pinna
erect or occasionally short-creeping. Leaves caes- divisions skeleton-like, the costae, costules, and
sometimes approximate, 1 5-35 cm long,
pitose to ultimate segments with bands of tissue only 1-
lamina glabrous, the axes glabrous, or rarely the several rows wide. Ultimate segments linear, 0.3-
rachis or petiole base with a few, scattered tri-
0.7 mm broad, each bearing a single vein, false
chomes. Petiole 5-16 cm long, 0.6-1.6 mm in di- veins lacking. Venation anadromous. Indusia sub-
ameter, nonalate. Lamina 3-4-pinnate-pinnatifid, conical to somewhat urceolate, not bilabiate, the
deltoid, opaque, 1 cell thick, but cells small and mouth truncate or slightly flaring, receptacle short-
occluded, not evident even when magnified at 1 0- to long-exserted.
1 5 x Rachis terete, nonalate, or marginate to
.

slightly alatetoward apex. Pinnae (at least prox- In wet forests, on trees or on the forest floor,
imal ones) short-stalked and the costae not alate sometimes on wet rocks, 300-1 300 m, Amazonas,
at base. Ultimate segments with linear lobes, each Loreto, Huanuco.
of these bearing a single vein, false veins lacking. Surinam to Colombia; Peru; Brazil.
Venation anadromous. Sori (most of them) borne With this perhaps should be included Trichom-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 85


anes sprucei Baker of northern South America, 1 2. Trichomanes angustifrons (Fee) Boer, Fl. Neth.
which is probably only a more robust form, with Antill. I (Pterid.): 17. 1962.
broader segments and broader bands of tissue along
the axes. Didymoglossum angustifrons Fee, Mem. foug. 11: 113,
t. 28, f. 5. 1866. TYPE:
Guadeloupe, I'Herminier
Amazonas: Prov. Bagua. roadside from Chiriaco to (holotype, P!).

Puente Venezuela, Barbour 4455 (MO, USM). Loreto: Pu-


mayacu, between Balsapuerto and Moyobamba, Klug Stem long-creeping, filiform. Leaves approxi-
3192 (F, GH, MO, us). Tierra Doble on Rio Nanay, LI. mate or well-spaced, 0.5-1.5 cm long, to 0.6 cm
Williams 1057 (F, us). Huanuco: Rio Llullapichis wa-
broad, subsessile or short-petiolate. Lamina nar-
tershed, ascent of Cerros del Sira, Dudley 13032 (GH),
Wolfe 12352 (GH, us).
rowly or broadly oblong, entire, or distally lobed,
costa percurrent, glabrous except for the dark, stel-
late trichomes borne along the margin. Venation

1 1 . Trichomanes punctatum Poiret ssp. sphen- catadromous, pinnate, with false veinlets sparsely
oides (Kunze) Boer, Acta Bot. Neerl. 1 1: 301. to amply distributed between the true veins. Sori

1962. solitary at lamina apex, or few and terminating


each distal lobe, somewhat to deeply immersed in
Trichomanes sphenoides Kunze, Farrnkrauter 216, /. the tissue. Indusia narrow-conical, bilabiate, the
88, f. 2. \ 840. TYPE: Peru, Cuchero (Dept. Huan- lips mostly wide-flaring and narrowly dark-mar-
uco), Poeppig (holotype, w!; isotype, MO!). gined, receptacle commonly exserted.
Didymoglossum sphenoides (Kunze) Presl, Hymeno-
phyllaceae 23. 1843.
Thus far represented in Peru by a single speci-
men from Huanuco, on ridge in jungle, 850 m;
Stem long-creeping, filiform. Leaves approxi-
elsewhere apparently always epiphytic.
mate or widely spaced, to 1.5 cm long and 1 cm
Costa Rica and Cocos Island; West Indies; Trin-
broad, subsessile or short-petiolate. Lamina cir-
idad and Tobago; the Guianas to Brazil; Peru;
cular to obovate or spathulate, margin entire to
Paraguay (reported from Chiapas, Mexico by Alan
irregularly lobed, a costa lacking or, in some fertile
Smith, in litt.).
laminae, distinct only in the proximal portion, gla-
Collections of this species are rather sparse, ex-
brous except for the dark, stellate, marginal tri-
cept from Brazil and the West Indies; yet it is
chomes. Venation flabellate, the veins crowded,
surely a much more common plant than the spec-
with false veinlets parallel to them. Sori several,
imens would indicate. Since it is such a tiny and
or sometimes solitary, partially immersed in the
inconspicuous fern, it certainly has been over-
lamina tissue, commonly borne between lobes, thus
looked and probably is not accurately represented
rarely extending beyond the outline circumscribed
in herbaria.
by the lobe apices. Indusia narrow-cylindrical, bil-
abiate, the lips mostly wide-flaring and narrowly
Huanuco (as San Martin): E of Tingo Maria, Allard
dark-margined, receptacle not or scarcely exsert- 21383a (us).
ed.
In dense, wet forests and wooded ravines, on
tree trunks, 100-1 100 m, Loreto to Pasco. 1 3. Trichomanes hymenoides Hedwig, Fil. gen. sp.,
Guatemala; Costa Rica; Panama; Greater An- /. 3,f. 3. 1799. LECTOTYPE
(designated by
tilles; Venezuela; Colombia to Bolivia. Boer, Acta Bot. Neerl. 11: 306. 1962): Hed-
Boer (1962) recognized four subspecies of Tri-
wig, t. 3, f. 3, probably based on a Swartz
chomanes punctatum, and only this one occurs in
specimen from Jamaica. Figure 12d.
Peru. Others differ from ssp. sphenoides in having
more distinctly lobed laminae and/or larger and Trichomanes muscoides Sw., J. Bot. (Schrader) 1 800(2):
more broadly dark-margined indusium lips. 95. 1802. TYPE: Jamaica, Swartz (holotype, S-PA).
Didymoglossum hymenoides (Hedwig) Desv., Mem.
Soc. Linn. Paris 6: 330. 1827.
Loreto: Maynas. Iquitos,Ushpa-Cana across from Rio Didymoglossum muscoides (Sw.) Desv. Mem. Soc.
Itaya, McDaniel 11381 (GH, MO). Huanuco: Prov. Huan- Linn. Paris 6: 330. 1827.
uco, near confluence of Rio Cayumba with Huallaga,
Mexia 8275 (F, MO, us). Pasco: Prov. Oxapampa, Valle
de Palcazu, Cacazu, Leon 681 (F). Puerto Bermudez (as Stem long-creeping, densely covered with dark
Junin), Killip & Smith 26592 (us). trichomes that extend onto the petioles. Leaves

86 FIELDIANA: BOTANY
well-spaced, (0.5-)l-3.5 cm long, to 1.5 cm broad, nuses and simple to bifid on outer margins, lam-
subsessile or short-petiolate. Lamina obovate to inar cells mostly elongate. Venation catadromous,
elliptic, 1-pinnatifid or rarely 2-pinnatifid, the cos- pinnate, false veinlets commonly abundant, ex-
ta percurrent (but juvenile laminae often subren- tending toward lamina margin but not (or very
iform and merely lobed and the costa indistinct rarely) parallel to it. Sori 1 or a few, borne near
distally),glabrous except for the dark, marginal lamina apex, conspicuously protruding from the
trichomes which are stellate in sinuses and simple leaf tissue, or only slightly immersed at base, not
to bifid on outer margin, laminar cells commonly or scarcely alate. Indusia narrow-funnelform, bil-
isodiametric. Venation catadromous, commonly abiate, the lips dark-margined, and sometimes
pinnate (occasionally subflabellate in juvenile flaring, most of them as long as or longer than
leaves), false veinlets sparse, scattered between true broad, receptacle short- to long-exserted.
veins, extending toward lamina margin but not
parallel to it. Sori 1-several borne near lamina On wet rocks in forests, 1900-2750 m, Ama-
apex, conspicuously exserted, or only slightly im- zonas, Huanuco.
mersed at base, not or very narrowly alate. Indusia Southern Mexico to Panama; Greater Antilles;
narrow-funnelform, bilabiate, but the lips quite Venezuela; Colombia; Ecuador; Peru; Brazil; Ar-
short, commonly broader than long, and usually gentina.
(not always) dark-margined, receptacle long- Although found thus far in Peru only on wet
exserted. rocks, Trichomanes reptans may be expected also
on trees, since it frequently has been reported as
In forests on trees, or rarely on rocky or sandy an epiphyte throughout much of its range.
soil, 700-2400 m, Amazonas, Loreto.
Southern Mexico to Panama; West Indies; Trin- Amazonas: Prov. Bagua, Cordillera Colan SE of La
idad; Venezuela and Colombia, south to Argentina Peca, Harbour3903 (MO, USM). Huanuco: Mito, Bryan
392 (F).
and Uruguay.
Juvenile leaves are often nearly reniform and
shallowly lobed, with veins subflabellate and the
costae sometimes indistinct toward the apex. Hence
15. Trichomanes krausii Hooker & Grev., Icon,
fil. 2, /. 149. 1831. TYPE: Dominica, Kraus
they can be confused with Trichomanes puncta-
tum. However, fertile leaves are pinnatifid, with (holotype, E).

stellate trichomes borne only in the sinuses, and


son strongly protrude from the leaf tissue. In T. Didymoglossum krausii (Hooker & Grev.) Presl, Hy-
menophyllaceae 23. 1843.
punctatum ssp. sphenoides sori are partially im-
mersed, and marginal trichomes are consistently
Stem long-creeping, densely covered with dark
stellate.
trichomes which extend onto the petioles. Leaves
well-spaced, 2-8 cm long, 1-3 cm broad, subsessile
Amazonas: Prov. Chachapoyas, Rio Ventilla of W to short-petiolate. Lamina narrow-ovate to -ellip-
Molinopampa, Wurdack 1513 (F, OH, us). Loreto: Sierra
del Pongo, Mexia 6289B (GH, us).
tic or obovate, 1 -pinnatisect to 2-pinnatifid, the
rachis distinct throughout and often abaxially dark-
pubescent at least near the base, the lamina pro-
14. Trichomanes reptans Sw., Prodr. 136. 1788. vided with simple to bifid trichomes, but with
TYPE: Jamaica, Swartz (holotype, S-PA). stellate ones in the segment sinuses. Venation ca-

tadromous, pinnate, false veinlets seldom numer-


Didymoglossum reptans (Sw.) Presl, Hymenophylla- ous, many of them parallel to and very near the
ceae 23. 1843. lamina margin. Sori 1 or a few, terminating the
segments, partly to deeply immersed in the seg-
Stem long-creeping, densely covered with dark ment tissue, the protruding portion narrowly or
trichomes that extend onto the petioles. Leaves broadly alate. Indusia salverform, bilabiate, the
well-spaced, 2-9 cm long, 14 cm broad, subsessile lips commonly dark-margined and flaring, recep-
to short-petiolate. Lamina lanceolate to ovate or tacle short- to long-exserted.

elliptic, 1-pinnatifid or rarely 2-pinnatifid, the


rachis distinct throughout, glabrous except for the Found near 800 m, San Martin, Huanuco. There
dark, marginal trichomes which are stellate in si- are no other label data on the only two collections

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 87


found thus far in Peru, but Trichomanes krausii Detailed study of many specimens throughout
occurs in forests on trees or wet rocks elsewhere the Neotropics indicates that Trichomanes ek-
throughout its range. manii is not distinct. The latter was said by Boer
United States (Florida); Mexico to Panama; West (1962) to differ in the uninterrupted submarginal
Indies; Trinidad; the Guianas to Colombia, south false vein, bordered on the outside by cubical cells,
to Paraguay and Argentina. whereas the submarginal vein of T. kapplerianum
Besides the characters used in the key, Trichom- was said to be sometimes interrupted and bor-
anes krausii often can be distinguished from closely dered by tangentially lengthened cells. These fea-
related species by the dark trichomes that fre- tures prove to be minor ones that are quite vari-
quently extend up the petiole onto the abaxial side able. In careful study of many specimens, including
of the lower rachis. Rachises of the other species types, one can nearly always find some interrup-
are essentially glabrous. tions in submarginal false veins; furthermore, shape
of the border cells is seldom constant. Many leaves
San Martin: Tarapoto, Spruce 3991 (BR, w). Huanuco: on the type specimen of T. kapplerianum, for ex-
Fundo Chela, Sinchono, Rio Chino, Aguilar 912 (USM). ample, have nearly as many cubical cells as tan-
gentially lengthened ones, and some cells are even
radially lengthened (supposedly a distinguishing
feature of T. hookeri Presl, of the Greater Antilles).
Additionally, most of the type specimens of T.
16. Trichomanes kapplerianum J. W. Sturm, Fl.
ekmanii have few leaves with somewhat
at least a
bras. 1(2): 276. 1859. TYPE: Surinam, near
interrupted submarginal veins. There is probably
Station Victoria, Kappler 1760 (holotype, w!).
no justification for separating either species from
T. hookeri, which Boer distinguished on similar
Hemiphlebium kapplerianum (J. W. Sturm) Prantl,
characters.However, the type of the latter has not
Hymenophyllaceae 46. 1875.
Trichomanes ekmanii Boer, Acta Bot. Neerl. 11: 319. been examined and T. kapplerianum is provision-
1962. TYPE: Dominican Republic, La Cumbre, ally maintained here as distinct.
Cordillera Central, Ekman H- 14342 (holotype,
u; isotypes, F!, G, GH!, MO!, s, uc, us!). San Martin: Near Tarapoto, Spruce 4762 (BR, w). Lor-
eto: San Antonio, on Rio Ataya, Killip & Smith 29522
Stem long-creeping, filiform, sparsely to abun- (F, GH). Prov. Maynas, on the Amazon, 50 miles down-
river from Iquitos, Moran 3661 (F). Huanuco: E of Tingo
dantly provided with brown trichomes which ex-
Maria (as San Martin), Allard 20912 (us). Puno: Near
tend onto the petioles. Leaves well-spaced, to 3
San "Gavan" (Gaban), Lechler 2297 (BR).
cm long and 1.5 cm broad, subsessile or short -
petiolate. Lamina obovate or oblong (or in juve-
nile ones often base rounded, cordate or
circular),
17. Trichomanes membranaceum L., Sp. pi. 1097.
cuneate, entire or distally lobed, with a distinct
1753. TYPE: "America," without specific lo-
costa at least in the proximal portion, glabrous, or
cality (holotype, LINN 1253.1).
sparsely pubescent basally along the costa. Vena-
tion catadromous, essentially pinnate (but some-
Lecanium membranaceum (L.) Presl, Hymenophyl-
times subflabellate veins ample to
distally), false laceae 12. 1843.
abundant, parallel to the true veins and with a Didymoglossum membranaceum (L.) Vareschi, Flora
submarginal, continuous or sometimes interrupt- Venezuela 1: 222. 1969.
ed false vein around the perimeter of the lamina.
Sori several to ample near lamina apex or on the Stem long-creeping, slender, densely covered
lobes, fully immersed in the tissue. Indusia fun- with dark brown trichomes which extend onto the
nelform, not bilabiate, the mouth expanded, but short petioles. Leaves well-spaced, 2-6 cm long
not dark-margined, receptacle short- or long- and often nearly as broad, subsessile. Lamina near-
exserted. ly circular and entire, or spathulate and incised
into regular lobes, lacking a distinct midrib, gla-
In dense forests and wooded ravines, on tree brous, but bearing along the margin numerous,
trunks (also on rocks, outside Peru), sea level to paired, circular, scalelike processes (these some-
1000 m, San Martin, Loreto, Huanuco, Puno. times deciduous in age). Venation flabellate, the
Guatemala to Panama; West Indies; the Guian- veins repeatedly dichotomous, false veinlets abun-
as to Venezuela; Peru; Brazil; Bolivia. dant and parallel with the true veins, but a con-

88 FIELDIANA: BOTANY
tinuous submarginal false vein lacking. Sori sev- sparsely scattered along the stem, unlike the dense,
eral to many on vein tips toward the lamina apex, blackish indument found on stems of T. reptans
partially to fully immersed in the tissue. Indusia and its relatives.

narrow-funnelform, not or scarcely bilabiate, the


mouth neither flaring nor dark-margined, recep- Mexia 6228 (GH, MO, uc,
Loreto: Sierra del Pongo,
tacle short- or long-exserted. us). Huanuco: E of Tingo Maria (as San Martin), Allard
21391 (GH, us). Pasco: Pichis Trail, Yapas (as Junin),
Killip & Smith 25543 (F, GH), 25555 (F). Junin: Schunke
In deep forests or wooded ravines, on tree trunks Hacienda above San Ramon, C. Schunke A240 (GH, us).
or wet rocks or cliffs, sea level to 850 m, Ama- Cuzco: Altura del Rio Tocate, Bues 1744 (us). Madre de
zonas, Loreto, Huanuco. Dios: Prov. Manu, Cerro de Pantiacolla, Foster el al.

Southern Mexico to Panama; West Indies; the 10713 (F).

Guianas to Colombia, south to Bolivia.

Amazonas: Prov. Bagua, Rio Maranon opposite Que-


brada Mirana, Wurdack 2040 (F, us). Loreto: Dist. Tigre, 19. Trichomanes tanaicum J. W. Sturm in Mart.,
Rio Tigre near Tigre, McDaniel & Rimachi 18531 (GH). Fl. bras. 1(2): 260. 1859. TYPE: Brazil, Para,
Pongo de Manseriche, Mexia 6198 (F, OH, MO, us). Rio Acara, Spruce 410 (holotype,
Huanuco: E of Tingo Maria (as San Martin), Allard 21549 B; possible
(GH, us). isotypes ["Spruce 44"], K.!, us; photo, A of K).

Lacostea tanaica (J. W. Sturm) Prantl, Unters. Morph.


18. Trichomanes polypodioides 1098. Gefasskrypt. 1: 50. 1875.
L., Sp. pi.
Trichomanes ankersii Hooker & Grev. var. tanaicum
1753. NEOTYPE (designated by Proctor, (Sturm) Sadebeck in Engler & Prantl, Nat. Pflan-
Flora Lesser Antilles 92. 1977): Montserrat, zenfam. 1(4): 105. 1899.
Proctor 19068 (A).

Stem long-creeping, stout and wiry, 0.5-0.8 mm


Trichomanes poeppigii Presl, Hymenophyllaceae 4 1 .
in diameter, amply provided with brown rhizoids
1843. TYPE: "Habitat in Peruvia," Poeppig(ho- which extend onto the rachis and adhere to tree
lotype, w?).
trunks. Leaves subdistant to remote, mature ones
3-15 cm long, 0.7-1.5 cm broad, subsessile or
Stem long-creeping, filiform, sparsely provided
scarcely petiolate. Lamina linear, subentire or lobed
with scattered brown trichomes. Leaves well-
(at base occasionally incised nearly to the rachis),
spaced, 4-12(-18) cm long, 1-3 cm broad, sub-
the rachis abaxially often provided with rhizoids,
sessile or short-petiolate. Lamina linear- or ob-
otherwise glabrous. Venation catadromous, veins
long-lanceolate, deeply lobed to pinnatifid, the axes,
free,1 -several times-forked in the lobes, false veins
veins and margin provided with dark brown tri-
lacking. Sori1 or a few, terminating the lobes, not
chomes that are stellate or forked from the base.
or scarcely immersed in the segment tissue. In-
Venation catadromous, lacking false veins. Sori
dusia salverform, not or scarcely bilabiate, the
1-few at or near segment apex, fully immersed in
mouth rather widely flaring, not dark-margined,
the segment tissue. Indusia salverform, neither bil-
receptacle scarcely to long-exserted.
abiate nor dark-margined, the mouth flaring, re-
ceptacle long-exserted. In deep forests,commonly hemiepiphytic on
tree trunks, 100-130 m, Loreto, Ucayali.
In deep forests, on tree trunks, very rarely on
Venezuela; Colombia; Brazil; Amazonian Peru.
wet clay banks, 350-2300 m, Loreto and Huanuco
The lamina in Trichomanes tanaicum is com-
to Cuzco and Madre de Dios.
monly lobed less than one-third to the rachis, but
Southern Mexico to Panama; West Indies; Trin-
Klug 92) some of the leaves are deeply
rarely (as in
idad; South America, to Brazil and Uruguay.
pinnatifid. Nevertheless, the narrow laminae with
Smaller specimens could be mistaken for Tri-
their linear outline distinguish them from other
chomanes reptans or other related species of subg.
species in section Lacostea.
Didymoglossum, which are similar in their deli-

cate, long-creeping stems, small, pinnatifid leaves,


Loreto: Mishuyacu, near Iquitos, Klug 92 (F, us). Prov.
marginal stellatetrichomes, and general aspect. Maynas, Nanay, Santa Maria de Nanay, J. Schunke
Dist.
However T. polypodioides lacks the false veinlets V. 2449 (F, GH). Ucayali: Prov. Coronel Portillo (as Lor-

common in that subgenus, and trichomes are eto), Rio Mazan, J. Schunke 300 (F, GH, uc, us, USM).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 89


20. Trichomanes tuerckheimii Christ, Hedwigia (1.8-)2-6 cm broad, subsessile. Lamina oblong,
44: 361. 1905. LECTOTYPE (designated here): pinnatisect, or pinnate to a broadly
winged rachis,
Guatemala, Alia Verapaz, Cubilquitz, von and (sometimes) veins abaxially
the rachis, costae,
Tuerckheim 8348 (lectotype, P!; isolectotypes, provided with simple, reddish brown trichomes
K., P!). LECTOPARATYPE: Peru, Loreto, near that adhere to tree trunks, free portion of larger
Leticia, Ule 6228 (K!, P!). segments 1.5-2.8(-3) cm long and 0.5-0.8 cm
broad, deeply crenate or crenate-serrate. Venation
Stem long-creeping, relatively stout and wiry, at least partly catadromous, veins free, commonly
0.5-1.2 mm in diameter, sparsely to abundantly simple, but some- to many-forked, false veins
provided with brown rhizoids that extend onto the lacking, but often some very short, darkened lines
leaf axes and adhere to tree trunks. Leaves sub- scattered between the veins. Sori few to several on
distant to remote, larger ones (9-)12-20(-30) cm the pinna lobes, not or scarcely immersed in the

long, 2-9 cm broad, subsessile. Lamina broadly segment tissue. Indusia tubular, neither bilabiate
oblong, pinnatisect, the rachis, costae and (some- nor dark-margined, mouth scarcely to slightly ex-
times) veins and margins abaxially provided with panded, receptacle commonly long-exserted.
simple, reddish brown trichomes that adhere to
tree trunks, free portion of larger segments 2.8-
4.5 cm long and 0.8-1.2 cm
broad, entire to shal- In forests, commonly hemiepiphytic, tightly ap-

lowly crenate. Venation catadromous, veins free, pressed to tree trunks, sea level to 1 000 m, Loreto,
simple, false veins lacking, but often some very Huanuco, Pasco, Cuzco.
short,darkened lines scattered between the veins. The Guianas to Colombia, south to Brazil and
Sori few to several on the pinna-lobes, not or Bolivia.

scarcely immersed in the segment tissue. Indusia This, Trichomanes tuerckheimii, and T. tanai-

tubular, neither bilabiate nor dark-margined, cum, all members of sect.


Lacostea, are commonly
mouth not expanded, receptacle commonly long- hemiepiphytic. Their leaves usually have the dis-
exserted. tinctive habit of being tightly appressed to tree
trunks. They cling to the bark by means of rust-
Thus far known
in Peru only from the lecto- colored, prehensile trichomes that are abundant
paratype, cited above, on tree trunks; elsewhere along the stems, rachises, and in T. ankersii and
hemiepiphytic, in forests on tree trunks, sea level T. tuerckheimii, on midribs and veins. Some col-
to 500 m. lectors' labels describe them as "plastered to the
Southern Mexico to Panama; Surinam to Co- bark of trees." Another character peculiar to the
lombia; Amazonian Peru. three species (though very rare in T. tanaicum) is
Included on the sheet with the syntype at Paris the presence of very short, dark lines seen scattered
is a fragment of a 2-pinnatifid leaf with nearly between and parallel with the veins. They some-
laciniate lobes, which might have been mounted what resemble the false veins of sect. Didymo-
here by mistake. It resembles the more highly di- glossum, except that they are not lengthy scler-
vided specimens which have been described as enchymatous strands. Rather, they merely appear
Trichomanes pedicellatum and T. subsessile. Re- to be a series of (usually) two or three constricted
lationships with these are discussed under T. an- and occluded cells that are mostly found toward
kersii.(Also described there are the dark lines in the segment margin.
the lamina which might be mistaken for the false Very closely related to Trichomanes ankersii are
veins of sect. Didymoglossum.) T. pedicellatum Desv. and T. subsessile Splitg., of
northern South America and the West Indies. These
appear todiffer only in their more highly divided
21. Trichomanes ankersii Hooker & Grev., Icon, lamina, which is 2- to 3-pinnatifid. Further
sterile

fil. 2, t. 201. 1831. TYPE: Guyana, Demerara, study may prove them all to be conspecific.
Ankers (holotype, K; photo, us).

Stem long-creeping, relatively stout and wiry, Loreto: Mishuyacu, near Iquitos, Klug 149, 1135 (F,
0.5-1.2 mm in diameter, sparsely to
abundantly us). Sierra delPongo, Mexia 6289 (GH, MO, uc, us).
Huanuco: 5 km NE of Tingo Maria, Stork & Horton
provided with brown rhizoids that extend onto the 9518 (F, GH, uc, us). Pasco: Prov. Oxapampa, Valle del
leaf axes and adhere to tree trunks. Leaves sub-
Palcazu, Leon 706 (F, GH). Cuzco: Prov. Quispicanchi,
distant to remote, larger ones 5-18 cm long, near Inambari, Vargas 16460 (GH).

90 FIELDIANA: BOTANY
22. Trichomanes bicorne Hooker, Icon, pi., t. 892. Trichomanes elegans Rudge, PI. Guian. 24, /. 35. 1805,
1854. LECTOTYPE (designated here): Brazil, nom. illeg. (not L. C. Rich., 1792, or Poiret, 1808).
Hvmenostachys diversifrons Bory, Diet, class, hist. nat.
Amazonas, Barra do Rio Negro, Spruce 1 178 8: 462. 1825. TYPE: "Guiane," Poiteau (holo-
(lectotype, K!; isolectotype, us!). LECTO- type, P?; isotype, L; photos, GH & us of L).
PARATYPE: Brazil, Amazonas, Sao Gabriel, Feea diversifrons (Bory) Copel., Phillip. J. Sci. 67: 74.
1938.
Spruce 2334 (K!, us).

Ptilophyllum bicorne (Hooker) Prantl, Unters. Morph. Stem stout, erect. Leaves crowded to caespitose,
Gefasskrypt. 1: 48. 1875. 10-35 cm long, strongly dimorphic, sterile ones
elliptic or lanceolate, deeply pinnatisect, short-pet-
Stem stout, decumbent or erect, abundantly iolate, fertile ones commonly longer, linear, sub-
provided with lustrous, castaneous, pluricellular lamina (2-)3-8 cm
entire, long-petiolate. Sterile
trichomes. Leaves monomorphic, crowded or broad, deeply divided into subfalcate segments 4-
caespitose, 3-10 cm long, 3-4-pinnatisect, the axes 7 mm
broad, the rachis abaxially provided with
(and sometimes veins) sparsely provided on the dark, simple trichomes and often flagellate and
abaxial side with simple, castaneous, pluricellular proliferous at the tip. Venation catadromous, a few
trichomes. Petiole 0.5-4 cm long, conspicuously to many veins anastomosing toward the segment
alate throughout, the alae broader than the petiole margin, 4-8 of them issuing from the rachis be-
quite or nearly to base. Lamina ovate, the axes tween adjacent costae, false veins lacking. Fertile
broadly alate, alae and segments often undulate. lamina simple, essentially entire, with sori ar-
Venation catadromous, veins free, false veins lack- ranged in a nearly continuous line along each mar-
ing. Sori immersed in the segment tissue. Indusia gin, fully immersed in tissue in the forks of veins.
short-tubular, borne in the forks of veins, each of Indusia subconical, not bilabiate, the mouth nei-
which extend well beyond the indusial mouth, the ther flaring nor recurved, strongly indented be-
mouth not dark-margined, receptacle long-exsert- tween the vein tips, receptacle short- to long-
ed. exserted at maturity.

In dense forests, epiphytic, or on wet humus or


Terrestrial in wet forests, in wooded ravines and
decaying logs, sea level to 1 50 m, Loreto.
on ridges and banks, 1 50-1 300 m, Amazonas and
A lectoparatype (Spruce 2334) is mounted on
Loreto to Junin, Madre de Dios.
the same sheet with the lectotype at Kew. How-
Southern Mexico to Panama; the Guianas to
ever, there are other specimens bearing this Spruce
Colombia, south to Brazil and Bolivia.
number that represent the type collection of Tri-
This is often confused with Trichomanes trollii,
chomanes spruceanum Hooker, a closely related
under which see further discussion.
species with dimorphic leaves which occurs in
northern South America. One sheet (BM) is the
isotype of T. spruceanum. Another, presumably Amazonas: Prov. Bagua, Quebrada Tambillo, valley
of Rio Maranon above Cascadas de Mayasi, Wurdack
the holotype, is at Kew.
2057 (GH, us). San Martin: Prov. Mariscal Caceres, Dist.
The species is aptly named for its sori, which Tocache Nuevo, Santa Rosa de Mishollo, J. Schunke V.
are borne at segment apices in the forks of veins. 6815 (F, MO, uc). Loreto: Prov. Maynas, Brillo Nuevo
The bordered on the outside with narrow
veinlets, and vicinity, Plowman et al. 6829 (GH). Huanuco: Tingo
wings of tissue, each extend well beyond the mouths Maria, Tryon & Tryon 5277 (F, GH, us). Pasco: Prov.
of indusia and are often curved, thus appearing Oxapampa, Cordillera San Matias, Ledn 320 (F, USM).
Junin: Cahuapanas, on Rio Pichis, Killip & Smith 26762
like two "horns." GH, us). Madre de Dios: Prov. Manu, Cerro de Pan-
(F,
tiacolla NNW of Shintuya, Foster et al. 10948 (F).

Loreto: Vicinity of Lago Llanchama near Rio Nanay,


Croat 18705 (F, MO). Mishuyacu, near Iquitos, Killip &
Smith 29988 (F, us). Maynas, Dist. Iquitos, Cacerio Mis- 24. Trichomanes trollii Bergdolt, Flora 127: 256,
hana, Rimachi 1256 (GH, MO).
264, t. 3. 1933. TYPE:
Bolivia, Dept. La Paz,
San Carlos (Mapiri), Troll 2531 (holotype, B;
frag., us!).
23. Trichomanes diversifrons (Bory) Sadebeck in

Engler & Prantl, Nat. Pflanzenfam. 1(4): 108. Feea trollii (Bergdolt) Vareschi, Flora Venezuela 1 :

1899. 247. 1969.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 91


Stem decumbent or erect. Leaves crowded to nate, elliptic-lanceolate, short-petiolate, fertile ones
caespitose, 10-25 cm long, strongly dimorphic, commonly longer and long-petiolate, sori subdis-
sterile ones elliptic, pectinate, short-petiolate, fer- tichous on the nonfoliaceous axis. Sterile lamina
tile ones commonly longer, linear, subentire, long- 1.2-3 cm broad, the pinnae oblong-lanceolate,
petiolate. Sterile lamina 1.5-4.5 cm broad, cut subentire to serrate, 34 mm
broad, the petiole
nearly to the rachis into subfalcate segments 1.5- and rachis with scattered, brown, pluricellular tri-
3( 4) mmbroad, the rachis abaxially provided chomes, the rachis often flagellate and proliferous
with simple, dark brown trichomes and often fla- at the tip. Venation catadromous, the veins free,

gellate and proliferous at the tip. Venation catad- commonly 1 -forked, false veins lacking. Fertile

romous, a few to many veins anastomosing toward leaves bearing numerous, stalked or subsessile, as-
the segment margin, 2(-3) of them issuing from cending sori on each side of the primary axis. In-
the rachis between adjacent costae, false veins dusia urceolate, scarcely or not bilabiate, the mouth
lacking. Fertile lamina simple, essentially entire, not flaring, receptacle strongly exserted at matu-
with sori arranged in a nearly continuous line along rity.
each margin, fully immersed in tissue in the forks
of veins. Indusia subconical, not bilabiate, the Thus far known in Peru from a single collection,
mouth very slightly flaring and recurved, but not roadside along a sandy, rocky stream bank, 350-
indented between the vein tips, receptacle strongly 700 m, Amazonas.
exserted at maturity. Elsewhere terrestrial or epipetric, commonly near
banks of streams or rivers, 250-800 m; Panama;
Terrestrial in dense, wet forests, often in wet the Guianas and Colombia.
ravines or along stream banks, 350-850 m, Ama- The small size, strongly dimorphic leaves, and
zonas to Pasco. nonfoliaceous fertile leaves easily distinguish this
Surinam to Colombia; Peru; Bolivia. from others in the genus.
This is closely related to the more robust Tri-
chomanes diversifrons. Besides the characters in Amazonas: Prov. Bagua, ca. 40-43 km NE of Chiriaco,
the key the latter can usually be recognized by its Barbour 4524 (MO, USM).
broader lamina (to 8 cm) and segments (4-7 mm),
these being separated from the rachis by a con-
spicuous wing ca. 2 mm
broad on each side. The 26. Trichomanes pinnatum Hedwig, Fil. gen. sp.,
lamina of T. trollii is rarely more than 4 cm broad 1799. LECTOTYPE
/. 4,f. 1. (designated by
and are cut nearly to the rachis into segments which Flora Lesser Antilles. 89.
Proctor, 1977):
are commonly 3 cm broad or less. Additionally,
Hedwig, /. 4,f. 1, based on specimen allegedly
the fertile lamina of T. diversifrons has dentate
from Jamaica. Figure 12c.
margins, due to the apices of the indusia which
are deeply indented between the vein forks. In-
Trichomanes pennatum Kaulf., Enum. fil. 264. 1824.
dusial mouths of T. trollii are typically flush with TYPE: French Guiana, without collector (holo-
the vein tips, so that the leaf margins are essentially type, LZ destroyed).
entire throughout. Neurophyllum pinnatum (Hedwig) Presl, Hymeno-
phyllaceae 19. 1843.

Amazonas: Prov. Bagua. along roadside from Chiriaco


to Puente Venezuela, Barbour 4461 (MO in part). San Stem short-creeping to decumbent, provided
Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo,
with dark brown or blackish pluricellular tri-
Palo Blanco. Plowman & Schunke 7431 (F). Huanuco:
Ascent of Cerros del Sira. Wolfe 12263A (GH, MO, us). chomes. Leaves essentially monomorphic (but fer-
Pasco: Prov. Oxapampa. W
side of Cordillera de San tile ones often somewhat larger, with longer pet-
Matias, D. Smith 2001 (MO). ioles),approximate or subcaespitose, to 70 cm long,
1 -pinnate, the axes sparsely to amply provided on

the abaxial side with castaneous trichomes. Petiole


25. Trichomanes botryoides Kaulf, Enum. fil. 263. about as long as the lamina, nonalate (sometimes
1824. TYPE: French Guiana, Poiteau (holo- marginate toward the lamina). Lamina ovate or
type?, P). subdeltoid, terminating in a conform apical seg-
ment, or the rachis prolonged, flagellate and pro-
Stem erect. Leaves crowded to caespitose, 4-12 liferous at the tip, the tissue thin and translucent,
cm long, strongly dimorphic, sterile ones 1 -pin- glabrous. Venation catadromous, the veins free,

92 FIELDIANA: BOTANY
except connected at the tips by a continuous mar- proliferous at the tip, the tissue relatively thick,
ginal vein, false veins copious and perpendicular commonly obscure, glabrous, pinna margins ser-
to true veins. Sori arranged in a nearly uninter- rate, the serrations obtuse to subacute. Venation
rupted line along pinna margins at the tips of veins, catadromous, the veins free, a lateral marginal vein
not immersed in the tissue, often stalked. Indusia faint and discontinuous or (more commonly) lack-

narrowly tubular, not or scarcely bilabiate nor dark ing, false veins lacking or a few very rare and faint
margined, mouth not flaring, receptacle long- ones perpendicular to true veins. Sori arranged in
exserted. a nearly uninterrupted line along pinna margins
at tips of veins, not immersed in the tissue, fre-

deep forests or wooded ravines,


Terrestrial, in quently stalked. Indusia conical to tubular, neither
inhumus or clay banks of streams, 150-1000 m, bilabiate nor dark-margined, mouth not flaring,

Amazonas and Loreto to Junin and Madre de Dios. receptacle exserted.


West Indies and Trinidad; Mexico to Panama
and south to Brazil and Bolivia.
Terrestrial in dark, wet forests and wooded rav-
Further differences between this and Trichom-
ines, often in frequently inundated areas along
anes hostmannianum are reviewed below in dis-
streams, sea level to 450 m, Amazonas, San Mar-
cussion of that species.
tin, Loreto.
Surinam to Colombia; Amazonian Brazil and
Amazonas: Prov. Bagua, valley of Rio Maranon near Peru.
Cascadas de Mayasi, Wurdack 1939 (GH, us). San Mar-
This and the closely related Trichomanes pin-
tin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Gran-

ja Santa Ines, J. Schunke V. 3654 (F, GH). Loreto: 1 7 km


natum are sometimes confused in herbaria. Be-
SW of Iquitos on road to Puerto Almendra, Croat 18475 sides the differences noted in the key they also
(F, MO, uc). Huanuco: Prov. Pachitea, Dist. Honoria, differ in size and in leaf texture. Leaves of T. host-
Bosque Nacional de Iparia, J. Schunke V. 1333 (F, GH, mannianum do not exceed 30 cm, pinnae are rare-
us). Pasco: Prov. Oxapampa, Palcazu Valley, Cabeza de
Mono, D. Smith 3741 (F, MO). Junin: E of Quimiri Bridge, ly more than 6 cm long and 1 cm broad, and tissue
near La Merced, Killip & Smith 23924 (F, us). Ucayali: of mature leaves is commonly dull green and ob-
Along trail to Arboretum of Bosque von Humboldt Ex- scure. Leaves of T. pinnatum often are 70 cm long,
perimental Station, km 86 of Pucallpa-Tingo Maria road, with pinnae to 1 5 cm long and 2 cm broad, and
D. Smith 1226 (F, MO). Madre de Dios: Prov. Tambo-
tissue is typically thin and translucent, so that the
pata, Vargas 18625, 18711 (GH).
false veins are easily visible with little magnifi-
cation.
Contrary to previous reports, false veins do oc-
27. Trichomanes hostmannianum (Klotzsch) cur in T. hostmannianum. They are infrequent and
Kunze, Bot. Zeit. (Berlin) 5: 352. 1847. scattered and so faint as to be overlooked, or un-
detectable, in the thick leaf tissue. However, in
Neurophyllum hostmannianum Klotzsch, Linnaea 1 8: immature leaves or in the rare thin-textured ones,
532. 1844. TYPE: Surinam, Hostmann 75 (ho- high magnification sometimes reveals a few or sev-
lotype, B; isotypes, K, NY, u, us!; photos, GH & us eralof them, perpendicular to the true veins as in
ofK).
Odontomanes hostmannianum (Klotzsch) Presl, Epi-
T. pinnatum. As this character is so difficult to

mel. hot. 21. 1849. observe, rather ineffective in delineating T.


it is

Ptilophyllum hostmannianum (Klotzsch) Prantl, Un- hostmannianum from species in other sections of
ters. Morph. Gefasskrypt. 1: 49. 1875. the subgenus; yet it serves to further establish its

Neurophyllum (Presl) Moore, along


position in sect.
Stem decumbent to erect, provided with dark with T. pinnatum. Morton ( 1 968) placed it outside
brown pluricellular trichomes. Leaves essentially the section because of the supposed lack of false

monomorphic (but fertile ones often larger, with veinlets.

longer petioles), caespitose, to 30 cm long, 1 -pin-


nate, the axes sparsely provided on the abaxial
side with brown trichomes. Petiole commonly as Amazonas: Nazareth, Osgood 25 (F, us). San Martin:
Prov. Mariscal Caceres, Dist. Tocache Nuevo, near
long as or longer than the lamina, not alate, or
Granja San Ysabel, J. Schunke V. 10313 (F, MO). Loreto:
narrowly so toward the lamina. Lamina ovate or Rio Tacsha Curaray, Croat 20403 (F, MO, uc). Dist. Iqui-
subdeltoid, terminating in a subconform apical tos, trail through Versailles, Mexia 6500 (GH, MO, uc,
segment, or the rachis prolonged, flagellate and us).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 93


28. Trichomanes humboldtii Lell., Mem. New McDanielet al. 22121 (F). Mishuyacu, near Iquitos, Klug
1246 (F, us).
York Bot. Gard. 38: 35. 1984, nom. nov. for
Trichomanes heterophyllum Willd. and with
the same type.
29. Trichomanes crinitum Sw., Prodr. 136. 1788.
Trichomanes heterophyllum Willd., Sp. pi. ed. 4, 5:
TYPE: Jamaica, Swartz (holotype, SET; iso-
503. 1810, nom. illeg., not T. heterophyllum (Sm.) types, BM!, s; photos, us of BM & s).
Poir. 1808. TYPE: Brazil, Amazonas, Javita,
Humboldt (holotype, B, Herb. Willd. 20210; pho- 16.
Ragatelus crinitus (Sw.) Presl, Hymenophyllaceae
tos, F, GH). 1843.
Homoeotes heterophylla Presl, Gefassbiindel Farm 24.
1847, nom. nov., based on Trichomanes hetero-
phyllum Willd. and with the same type.
Stem Leaves monomorphic, caespitose,
erect.

Feea humboldtii Bosch, Ned. Kruidk. Arch. 4: 347. 5-25 cm cm broad, moderately to
long, l-2(-2.5)
1859, nom. nov., based on Trichomanes hetero- amply pubescent, the trichomes delicate, un-
phyllum Willd. and with the same type. branched, unicellular (or rarely pluricellular) be-
Feea heterophylla Copel., Philipp. J. Sci. 67: 73. 1938,
nom. nov., based on Trichomanes heterophyllum yond a short, enlarged, basal cell. Petiole nonalate,
Willd. and with the same type. or narrowly so toward the lamina. Lamina linear,
cut nearly or quite to the rachis into deeply lobed
to pinnatifid, patent, pinnae, gradually reduced to
long- to short-creeping. Leaves deep-
Stem wiry, a pinnatifid apex, lamellae lacking on veins. Ve-
ly pinnatisect to nearly pinnate, subdistant or re- nation catadromous, the veins free, false veins
mote, to 22 cm long, dimorphic, sterile ones short- on the apical
lacking. Sori terminating the veins
petiolate, to 3 cm broad, fertile ones long-petio-
part of the pinnae. Indusia tubular, deeply im-
late. far exceeding the sterile ones and 1 cm broad mersed in the segment tissue or, if partially im-
or less, the petiole essentially nonalate, sparsely or mouth
mersed, broadly alate on each side, the
amply provided with castaneous, pluricellular tri- bilabiate and somewhat flaring, the lips not dark-
chomes. Sterile lamina oblong or oblong-elliptic,
margined, receptacle exserted.
commonly equaling or longer than the petiole, the
apex pinnatifid. Venation catadromous, veins free
Apparently known thus far from two collections
and dichotomously branched, false veins lacking. on tree trunks, ca.
in Peru, in damp elfin forest,
Fertile lamina linear, much shorter than the pet-
1850 m, Huanuco. Elsewhere epiphytic or epi-
iole, with a few sori terminal on each pinna and
700-2300 m.
petric,
fully immersed in the tissue. Indusia subconical West Indies; Costa Rica; Venezuela; Colombia;
or broadly tubular, not bilabiate, the mouth not
Ecuador; Peru.
(or scarcely) flaring nor dark-margined, the recep-
Trichomes in many species of subg. Achomanes
tacle long-exserted.
are pluricellular, with two to several greatly elon-
gate cells, these often springing from a very short,
Terrestrial, in open to dense forests, commonly widened basal cell. In species most closely related
in wet, sandy places, sea level to 1400 m, San to Trichomanes crinitum, these are often mixed
Martin, Loreto. with a number of unicellular (above the basal cell)
Venezuela; Colombia; Amazonian Peru and trichomes. However, T. crinitum differs from its
Brazil.
Peruvian relatives in having all, or nearly all, of
There has been some confusion with nomen- its laminar trichomes unicellular above the basal
clature of this species. Lellinger's (1984) citing of
cell.
Trichomanes humboldtii "Bosch" (1858) was in
error, since therewas no such name; instead, it SW
Huanuco: slope of Rio Llullapichis watershed, on
was "Feea humboldtif that Bosch had published. the ascent of Cerros del Sira, Dudley 13525 (GH), 13545
However, Lellinger's use of 'T. humboldtii"'' is le- (GH, us).

gitimate, as a nom. nov. (Art. 72); therefore this


name is correct for the species, with Lellinger as
the author.
30. Trichomanes martiusii Presl, Hymenophyl-
San Martin: Rioja, Soukup 5055 (us). Rio Negro,
laceae 36. 1843. TYPE: a renaming of T. pi-

Woytkowski 6211 (GH, MO, us). Loreto: Prov. Maynas, losum (sensu Martius in part, Icon. pi. crypt.
Rio Nanay, Mishana, Foster & Foster 4103 (GH); 104, t. 68 right) presumably based on "Brazil,

94 FIELDIANA: BOTANY
Arara-coara" (Araracuara, on Rio Caqueta, Iquitos, Klug 196, 51 1 (F, us). Prov. Maynas, 10 km S
of Iquitos, Tryon & Tryon 5179 (GH, us). Timbuchi on
now Colombia), Martius (holotype, M; iso-
Rio Nanay, LI. Williams 957 (F, us).
types, BR, L; photo, us of L).

Trichomanes plumula Presl, Hymenophyllaceae 15,


36. 1843. TYPE: based on Martius in part. Icon.
31. Trichomanes lucens Sw., Prodr. 136. 1788.
pi. crypt. 104, /. 68 left, specimens probably as
TYPE: Jamaica, Swartz (holotype, SET; iso-
for T. martiusii. type, BM; frag., B; photos, us of SBT & BM), not
Ptilophyllum martiusii (Presl) Prantl, Unters. Morph. T. lucens Hooker & Grev., 1827.
Gefasskrypt. 1: 48. 1875.

Trichomanes lambertianum Hooker, Sp. fil. 1: 139, /.


Stem erect to decumbent. Leaves monomor- 4 IB. 1846. TYPE: Peru, Huanuco, Pillao, Ruiz
phic, crowded to caespitose, 0-40 cm long, (2-)3-
1 & Pavon (holotype, K; isotype, B; photos, GH &
8 cm broad, densely (on axes) to amply pubescent, us of K).
the trichomes unbranched, unicellular beyond the Ptilophyllum lambertianum (Hooker) Prantl, Unters.
Morph. Gefasskrypt. 1: 48. 1875.
short basal cell, or pluricellular on petiole and
rachis. Petiole nonalate. Lamina narrowly elliptic
Stem erect to decumbent. Leaves monomor-
or lanceolate, gradually reduced to a pinnatifid
phic, crowded to caespitose, 10-60 cm long,
apex, cut nearly or quite to the rachis into linear
(2.5-)3-12 cm broad, amply to densely pubescent
pinnae, the proximal ones commonly deflexed,
(indument often obscuring the laminar surface),
subentire to crenulate or serrate, or rarely with a
the trichomes unbranched, unicellular and pluri-
few, scattered lobes, with numerous, crestlike la-
mellae borne on the veins abaxially. Venation ca-
cellular beyond the scarcely evident basal cell. Pet-
iole nonalate. Lamina linear to lanceolate, pin-
tadromous, the veins free, false veins lacking. Sori
nate-pinnatifid (occasionally 2-pinnate-pinnatifid
terminating the veins on the apical part of seg-
toward base), gradually reduced to a pinnatifid
ments. Indusia tubular or narrow-conical, deeply
mouth apex, pinnae narrow-deltoid, mostly ascending,
immersed in the segment tissue, the trun-
lacking lamellae. Venation catadromous, the veins
cate, not dark-margined, sometimes apparently
free, false veins lacking. Sori commonly terminal
bilabiate due to narrow wings of tissue extending
on ultimate segments near apex of pinnae. Indusia
beyond the mouth on two sides, receptacle com-
tubular, deeply immersed in the segment tissue,
monly exserted.
the mouth scarcely or slightly flaring, not dark-

On tree trunks in forests, or on clay banks along margined, sometimes apparently bilabiate due to
the narrow wings of tissue extending beyond the
roads and streams, sea level to 150 m, thus far
mouth on two sides, receptacle commonly exsert-
found only in Loreto, but locally common there.
ed.
to Colombia; Peru; Amazonian Brazil.
Surinam
The shape of the indusium is somewhat vari-
able. The mouth is commonly truncate and not at and wooded ravines, pendent from
In forests
all wings of tissue along
bilabiate, but frequently the trees or in sphagnum on sandy
wet rock crevices, or
the sides extend beyond the mouth, giving it a banks, 2150-3500 m, Cajamarca, Amazonas,
"horned" appearance, much like Trichomanes bi- Huanuco, Pasco, Cuzco.
corne. This a distinctive species in Peru, because
is Jamaica; Costa Rica; Venezuela and Colombia,
of the crestlike lamellae borne on many of the south to Bolivia; Brazil.
veins on the abaxial side, perpendicular to the With its l-(2-)pinnate-pinnatifid leaves and the
plane of the lamina. These processes are similar copious, long trichomes that often obscure the
to those occurring in several species of Hymeno- lamina surface, this should not be mistaken for
phyllum. On some of the more densely hirsute any other species of subg. Achomanes.
specimens of T. martiusii, the lamellae are par-
tially obscured by the indument and thus they Cajamarca: Prov. Jaen, Paso de Huascarai, 15 km SE
have been mistaken for other species in subg. of Huancabamba, Fosberg 27842 (us). Amazonas: Prov.

Achomanes, particularly T. pilosum Raddi, of Bo- Chachapoyas, Cerros de Calla Calla above Leimebamba,
Hutchison & Wright 5568 (F, GH, MO, uc, us). San Mar-
livia and southern South America. tin: Prov. Mariscal Caceres, Rio Abiseo National Park,

Young & Ledn 4471 (F). Huanuco: Playapampa, Mac-


Loreto: Prov. Maynas, Rio Nanay, behind Mishana, bride 4499 (F, GH, us). Pasco: Prov. Oxapampa, Cordi-
Gentry & Revilla 20704 (MO, uc, USM). Mishayucu, near llera Yanachaga, Cerro Pajonal, Foster 9047 (F, MO).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 95


Cuzco: Prov. La Convention, Cordillera Vilcabamba, Trichomanes elatum Desv., Mem Soc. Linn. Paris 6(3):
Dudley 10789 (F, GH, MO). 327. 1827 (not Forst., 1786, or Bosch, 1861).
TYPE: "habitat in America calidiori," without
collector (holotype, P; photo, GH).

32. Trichomanes pellucens Kunze, Linnaea 9: 104.


1 834. TYPE: Peru, Tocache, upper Rio Hual-
Stem short-creeping to decumbent. Leaves
monomorphic, subdistant to contiguous, (rarely
laga, Poeppig (holotype, LZ destroyed; iso-
caespitose), mature ones 1 5-50 cm long, 3-8 cm
types, BR!, w; possible isotype, L!; photos, GH,
& us of BR). The epithet broad, moderately to amply pubescent, the tri-
HB T. pellucidum was
chomes unbranched, those of the veins and mar-
used by Presl (Hymenophyllaceae 15. 1843),
but later he indicated this was an error for T. gins unicellular beyond a very short basal cell,
those of the rachis (especially abaxially) predom-
pellucens Kunze (Epimel. bot. 16. 1851).
inantly dark brown, 1-5-celled above basal cell,
stout, rigid and spreading, most of them terete
Ptilophyllum pellucens (Kunze) Prantl, Unters. Morph.
Gefasskrypt. 48. 1875.
toward their base. Petiole essentially nonalate, am-
ply provided with dark brown trichomes. Lamina
Stem erect to decumbent. Leaves monomor- lanceolate, often broadly so, gradually reduced to
a pinnatifid apex, cut nearly to the rachis into
phic, close, contiguous or caespitose, 8-30 cm long,
3-7 cm broad, sparsely to moderately pubescent, linear to narrow-deltoid or -oblong segments, these

the trichomes unbranched, light to dark brown, patent (or the basal ones deflexed), lacking lamel-
those of the veins and margins unicellular beyond lae, the margins subentire to crenate or serrate,

a very short basal cell, those of primary axes 1-3- often undulate. Venation catadromous, the veins
celled beyond the basal cell. Petiole rather con- free, false ones lacking. Sori borne near and at

spicuously alate halfway or more to the stem. Lam- segment apices, at tips of veins, sometimes flanked
ina narrowly or broadly lanceolate, gradually re- by vein forks. Indusia tubular or narrow-conical,
duced to a pinnatifid apex, cut nearly or quite to deeply to fully immersed in the segment tissue,
the rachis into adnate, linear or linear-oblong, pat- the mouth neither bilabiate nor dark-margined,
ent pinnae (or basal ones deflexed), their margins not or scarcely flaring, receptacle exserted.

subentire to crenate, lacking lamellae. Venation


catadromous, the veins free, false veins lacking. In dense, wet forests, occasionally epiphytic, but
Sori borne at tips of veins in the forks, near and commonly on wet sandy or mossy soil, 550-2100
at segment apices. Indusia tubular, deeply im- m, San Martin to Puno.
mersed in the segment mouth not bil-
tissue, the Ecuador; Peru; Bolivia; Brazil.
abiate, flaring, or dark-margined, receptacle com- This and Trichomanes cristatum are easily con-
monly exserted. fused. Some collections from Peru are interme-
diate between the two, and there is evidence of
Epiphytic in forests, or terrestrial, most com- hybridization involving these and other closely re-
monly in sandy soil in ravines or on stream banks, lated species. The indument on the abaxial side of
1 50-800 m, Amazonas, San Martin, Cuzco, Madre the rachis seems to be the best character for dis-
de Dios. tinction, as indicated in the key. In T. cristatum
Venezuela; Colombia; Ecuador; Peru; Brazil. most of these trichomes are tawny to orange, rath-
er long, delicate and tortuous, with their usually
Amazonas: Prov. Bagua, NE of Chiriaco, Barbour 4461 four to six cells greatly flattened. Those of T. plu-
(MO 4462 (USM), 4575, 4518 (MO). San Martin:
in part),
mosum are predominantly dark brown (some
Prov. Mariscal Caceres, Dist. Tocache Nuevo, Palo
blackish), shorter, stout and rigid, and terete at
Blanco, Plowman & J. Schunke V. 7434 (F); J. Schunke
V. 5722 (F, MO). Cuzco: Prov. Paucartambo, Cosnipata leasttoward the base. Careful examination of spec-
Valley, Rio Tono, Wachter el al. 134 (F). imens is necessary to view these trichomes, as those
on the adaxial side of the rachis differ little, being
primarily light brown, even tawny, on both species.
33. Trichomanes plumosum Kunze, Linnaea 9: Curiously (and unfortunately) most of the speci-
104. 1834. TYPE: Peru, "in sylvis montanis mens examined for this study had been mounted
ad Pampayaco" (Pampayacu, Dept. Huanu- with abaxial side down, so that it was very difficult
co), Poeppig diar. 1107 (holotype, B; isotypes, to determine the character of trichomes on the
BR!, K; photo, BM of BR). other side. Besides differences in indument, T. plu-

96 FIELDIANA: BOTANY
mosum can usually be distinguished from T. cris- to slight projections of tissue along the flanking
tatum Peru) by lamina shape: that of
(at least in vein forks, receptacle exserted.
the latter is commonly linear, rarely more than 4
cm broad, whereas the lamina of T. plumosum is Terrestrial, or occasionally on tree trunks, com-
typically broadly lanceolate, usually more than 4 monly in wet forests on sandy soil or in open
cm broad. sphagnum bogs, sea level to 2000 m, San Martin
and Loreto to Puno.
San Martin: Tarapoto, Spruce 4766 (BR, w). Huanuco: Venezuela and Colombia, south to Brazil and
SW slope of Rio Llullapichis watershed, ascent of Cerros Argentina.
del Sira, Dudley 13051 (GH), 1 3128 (GH, us), 1 3512 (GH). This species very easily confused with Tri-
Pasco: Prov. Oxapampa, 1 9 km W of Oxapampa, D.
is

Smith 2209 (MO). Junin: La Merced, Macbride 5634 (F, chomanes plumosum; characters are discussed
GH). Cuzco: La Convencion, Biies 2028, 2140 (us). Madre above in the treatment of the latter. Trichomanes
de Dies: Prov. Manu, Shintuya, Chavez 804 (MO). Puno: cristatum and T. plumosum share nearly the same
Prov. Carabaya, San Gaban, Vargas 18875 (GH). Dept.
distribution and are often found growing together.
Unknown: "ad saxa humida prope Sangari," Lechler 2548
In Peru, at least, there are frequent intermediates,
(BR, K, w 3 sheets); although this is the type number of
T. undulatum (= T. vandenboschii), these specimens rep- which indicates a strong probability of hybridiza-
resent part of a mixed collection). tion. These problems are not confined to species
of Peru but also occur in other taxa of the group
of T. crispum L. throughout the Neotropics, as
pointed out by Windisch (1988). Obviously there
34. Trichomanes cristatum Kaulf., Enum. fil. 265. is a great need for careful analysis of plants in situ

1824. TYPE: Brazil, collector undesignated as well as further caryological study before ac-

(holotype, LZ destroyed). curate relationships are understood.

Trichomanes sellowianum Presl, Hymenophyllaceae San Martin: Rio Negro, Woytkowski 6210 (GH, MO,
15, 37. 1843. TYPE: "Brasilia," Sellow 197 (ho- uc, us). Loreto: Prov. Maynas, Dist. Iquitos, Rio Nanay,
lotype, PR?; possible isotypes, 2 unnumbered Sel- McDaniel & Rimachi 18924 (F, MO). Huanuco: Tingo
low sheets from "Brasilia," B!, K!). Maria (as San Martin), Allard 21585 (GH, us), 27559
(us). Pasco: Prov. Oxapampa, Palcazu, van der Werff
el at. 8376 (MO, uc). Junin: E of Quimiri Bridge near La
Stem short-creeping to decumbent, rarely erect.
Merced, Killip & Smith 23952 (F, us). Cuzco: Prov. La
Leaves monomorphic, subdistant to contiguous, Convencion, Valle de Santa Ana, Herrera 3008 (us).
mature ones 15-50 cm long, 2-4(-5) cm broad, Puno: Prov. Sandia, Vargas 1 1832 (GH).
moderately to rather densely pubescent, the tri-

chomes unbranched, those of the veins and mar-


gins unicellular beyond a very short basal cell, 35. Trichomanes vandenboschii Windisch, Bra-
those of the rachis abaxially tawny to orange, del- dea 5(4): 57. 1988, nom. nov. for T. undula-
icate, most of them long and tortuous and with tum Bosch.
the cells flattened, predominantly unicellular be-
yond the basal one (occasionally with 2-3 cells). Trichomanes undulatum Bosch, Ned. Kruidk. Arch.
Petiole essentially nonalate, amply provided with 5(2): 147. 1861, nom. itleg. (not Swartz, 1788).
LECTOTYPE (designated by Windisch, Bradea
long, orange to tawny trichomes, or these dark
5(4): 57. 1988): "Peruvia, prope Sangari" (Dept.
brown toward the stem. Lamina linear to linear-
Puno, Prov. Carabaya, Dist. Ayapata) Lechler
lanceolate, commonly 5-12 times as long as broad, 2548 (L; isolectotypes, BR!, P; photos, HB oft, GH
gradually reduced to a pinnatifid apex, cut nearly of BR, GH & us of P). Specimens of Lechler 2548,
to the rachis into linear or narrowly oblong seg- a mixed collection at BR, K, and w (3 sheets), are
not type material, but are instead T. plumosum.
ments, these patent (or basal ones deflexed), lack- LECTOPARATYPES: "Peruvia, Tatanara"
ing lamellae, the margins subentire to crenulate, (Dept. Puno, Prov. Carabaya, Dist. Ayapata),
often undulate. Venation catadromous, the veins Lechler 2503. 2571 (L?, P?).
free, false veins lacking. Sori borne near and at
segment apices, at tips of veins, often flanked by Stem short-creeping. Leaves monomorphic, ap-
the vein forks. Indusia tubular or narrow-conical, proximate to subdistant, 5-15 cm long, 1.8-3 cm
deeply to fully immersed in the segment tissue, broad, moderately to densely pubescent, the tri-
the mouth not dark-margined, not or slightly flar- chomes unbranched, those of the veins and mar-
ing, not bilabiate but sometimes appearing so due gins unicellular beyond a very short basal cell,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 97


those of the rachis tawny to orange, delicate, many chomes (to 4 mm), reflexed pinnae in the proximal
of them long and tortuous and with the cells flat- portion of the lamina, and the petiole very nar-
tened, frequently with 2-8 cells beyond basal one, rowly alate halfway to the stem. Windisch (in litt.)
as well as with many unicellular ones. Petiole es- stated that he had seen one specimen (AAU) from
sentially nonalate, sparsely provided with short, Peru (Loreto, Prov. Maynas, Plowman et al. 6633).
light to dark brown trichomes. Lamina oblong- to Although this specimen has not been examined,
ovate-lanceolate, 3-4 times as long as broad, grad- duplicates (F, GH) are scarcely distinct from T. cris-
ually reduced to a pinnatifid apex, cut nearly or tatum, as rachis trichomes are less than 2 long mm
quite to the rachis into narrowly oblong segments, and proximal pinnae are patent or just a few of
these patent (or basal ones deflexed), lacking la- them reflexed. Only a few petioles are obscurely
mellae, the margins entire to crenulate, often un- alate to marginate. The species is found in Trin-
dulate. Venation catadromous, the veins free, false idad, the Guianas, Venezuela, Colombia, and Bra-
veins lacking. Sori commonly borne at segment and future collections of this species complex
zil,

apices, at tips of veins, often flanked by the vein from Peru need to be examined carefully to de-
forks. Indusia urceolate, fully immersed in the seg- termine if T. accedens truly occurs in Peru.
ment tissue, the mouth somewhat flaring, not dark-
Trichomanes delicatum Bosch, Ned. Kruidk. Arch.
margined, not or scarcely bilabiate, receptacle
exserted at maturity. 5(2): 145. 1861. TYPE: Ecuador, Quito, Cum-
ing 21 (holotype, B; frag., L).

In deep, wet forests, on tree trunks or in wet,


This is closely related to T. crinitum in that the
boggy places, rarely in crevices of wet rocks, 750-
1 800 m, Cuzco. trichomes are unicellular above the short basal
cell. It differs from that species in the alate petiole,
Venezuela; Colombia; Ecuador; Peru; Bolivia;
northern Brazil. the more shallowly lobed pinnae, and the glabrous

This is closely related to Trichomanes cristatum, margins of the indusia. It is to be expected in Peru,
since it has been found in Colombia, Ecuador, and
from which it is distinguished primarily by its
Bolivia.
smaller size and relatively broader lamina. Besides
the key characters, it can usually be separated from Trichomanes haenkeanum Presl, Hymenophylla-
both T. cristatum and T. plumosum by its some- ceae 15, 36, 65. 1843. TYPE: Peru, mountains
what flaring indusium mouth. That of the other of Huanuco, Haenke (holotype, PR?).
two species is scarcely or not at all expanded. The
Trichomanes crispum var. haenkeanum C. Chr., In-
confused identities of species within this complex
dex fil. 641. 1905.
is illustrated by the fact that the number of the
In the protologue, Presl indicated a close rela-
type collection is shared with specimens of T. plu-
mosum, as noted above in the citation of types. tionship with Trichomanes crispum. Characters
used in the description certainly align T. haen-
Cuzco: Maranura, Beatriz, Bites 895 (us). Prov. La keanum with this group of species but are insuf-
Convencion, Sahuayaco, Vargas 6288 (GH). Prov. La ficient to place it precisely. Until the type is located
Convencion, Choquello, Vargas 8184 (uc). Prov. Pau- and compared with other species in the complex,
cartambo, Atalaya, Valle Kosnipata, Vargas 23155 (OH).
nothing more can be determined.

Comments FamilyT: LOXOMATACEAE


Trichomanes accedens Presl, Epimel. bot. 14: 1851. Loxomataceae Presl, Gefassbiindel Farm 31.1 847,
SYNTYPES: Guyana, Rich. Schomburgk 27 1 as Loxsomaceae. TYPE: Loxoma Cunn. (often
(B; photos, GH, us); Hooker &
Grev., Icon, altered to Loxsoma).
fil., t.12, based on a Guilding collection from
St. Vincent (specimen not seen by Presl). Stem long-creeping, slender to rather stout,
branched, densely pubescent with stiff trichomes
This belongs to the Trichomanes crispum com- enlarged at base. Leaves to 5 m
long, pinnate,
plex and is allied to T. cristatum. It differs from commonly pubescent abaxially, circinate in ver-
the latter primarily in its predominantly erect stem nation. Petiole lacking stipules. Sporangia borne
with contiguous leaves, much longer rachis tri- in marginal sori on an elongate receptacle, within

98 FIELDIANA: BOTANY
a more or less urceolate indusium, short-stalked glaucous abaxially. Petiole dark brown to atro-
(ca. 6 rows of cells), with an oblique annulus not purpureous, sublustrous. Lamina subdeltoid,
interrupted by the stalk. sparsely to amply provided on surface, veins and
costules with light to dark brown, tortuous, septate
This small family contains two genera: Loxoma trichomes, glabrous adaxially. Pinnae commonly
of New Zealand, and the Neotropical Loxsomop- well-spaced, subdeltoid, somewhat or slightly re-
sis. duced at base, with pinnules strongly ascending,
segments and venation catadromous. Veins pin-
nately branched, prominulous. Indusia urceolate
I. Loxsomopsis to (especially at maturity) narrow-cyathiform.

Loxsomopsis Christ, Bull. Herb. Boissier II. 4: 399. Scandent or trailing, or arching to pendent from
1904. TYPE: Loxsomopsis costaricensis clay banks, in on exposed ridges,
elfin forests, often

Christ. Figure 13. 2200-3400 m, Huanuco, Pasco, Cuzco.


Ecuador and Peru.
Terrestrial. Stem bearing scattered fibrous roots Range of the species perhaps should be extended
and abundant, rigid, lustrous, dark trichomes which to include Costa Rica and Bolivia. Specimens de-
are enlarged at the base. Leaves monomorphic, scribed as Loxsomopsis costaricensis Christ differ
well-spaced on the stem, 2-pinnate-pinnatifid to at all, from L. pearcei. The type (Werckle
little, if

nearly 3-pinnate. Lamina subcoriaceous, glabrous & Brune 279, Costa Rica) has not been seen, but
adaxially, glabrous to pubescent abaxially. Veins the original description and illustrations suggest
free. Sorus marginal, paraphysate, the indusium the two species are synonymous. Loxsomopsis no-
narrow-cyathiform to urceolate, the rim entire, the tabilis Slosson (type from Bolivia, R. S. Williams

elongate receptacle exserted. Spores trilete, glo- 1303) is supposedly distinguished by its glaucous
bose-tetrahedral, the surface coarsely tuberculate. abaxial surface and by the larger pinnae being
greatly reduced at base. Two sheets of isotype (us)
The genus
is probably represented by a single seem to illustrate this clearly, and yet another is-

species. However, two very closely related species otype (P) is only slightly glaucous and pinnae are
have been described (from Costa Rica and Bolivia, not very strongly reduced at base. Lack of consis-
respectively), based on supposed differences in size tency in all these features suggests that there is but
of leaf, color of petiole, and shape of pinnae and one species of Loxsomopsis.
indusia. Two others have been separated on the
basis of other variable and doubtfully significant Huanuco: SW
slope of Rio Llullapichis watershed, as-
characters. Observations of a number of cent of Cerros del Sira, Dudley 13420 (GH). Playapampa,
speci-
Macbride4521 (F, GH. us). Pasco: Prov. Oxapampa, Cord.
mens throughout the range suggest that such char-
San Gutardo, Ledn 532 (USM). Cuzco: Valle San Miguel,
acters vary with age of the plant and maturity of
La Convention, Bues 2119 (us). Prov. La Convention,
sori, and they demonstrate no significant corre- Cordillera de Vilcabamba, Dudley 10713 (GH, MO).
lation.

1 Loxsomopsis pearcei (Baker) Maxon, Proc. Biol.


.

Soc. Wash. 46: 105. 1933. Figure 13. Family 8: PLAGIOGYRIACEAE

Dicksonia pearcei Baker, Ann. Bot. (London) 5: 197.


Plagiogyriaceae Bower, Ann. Bot. (London) 40:
1891. TYPE: Ecuador, "Eastern Andes, 8000- 484. 1926. TYPE: Plagiogyria (Kunze) Mett.
9000 ft.," Pearce 251 (holotype, K!; photo, us).
Loxsomopsis lehmannii Hieron.. Bot. Jahrb. Syst. 34: Stem stout, erect to decumbent, indurated, bear-
435. 1904. TYPE: Ecuador, prope Chinguinda, fibrous roots, essentially lacking indu-
ing stiff,
Cordillera Oriental de Sigsig, 800-2500 m, Leh-
1

ment. Leaves pinnate, circinate in vernation, di-


mann 5061 (holotype, LZ destroyed; isotype, us!).
Dennstaedtia pearcei (Baker) C. Chr., Index fil. 218. morphic. Petiole expanded basally into indurated
1905. stipules, commonly with a double row of aero-
phores at base (these rarely discernible in dried
Leaves 0.4-5 m long, 2-pinnate-pinnatifid to plants). Sporangia exindusiate, not paraphysate,
nearly 3-pinnate, sometimes inconspicuously commonly covering the abaxial surface of fertile

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 99


FIG. 1 3. Loxsomopsis pearcei: a, habit; b, portion of fertile pinna. (From Prieto P-259, Ecuador, F.)

100 FIELDIANA: BOTANY


pinnae, long-stalked (4-6 rows of cells), with an LELLINGER, D. B. 1971. The American species
oblique annulus not interrupted by the stalk. Spores of Plagiogyria sect. Carinatae. Amer. Fern J.,
trilete, tetrahedral-globose, the surface irregularly 61: 110-118.
tuberculate.

The family consists of a single genus essentially 1 .


Plagiogyria semicordata (Presl) Christ, Farnkr.
confined to wet, montane areas in Asia, Austra- Erde. 176. 1897. Figure 14.
lasia, and tropical America. These ferns resemble
some species ofBlechnum because the sterile leaves Lomaridium semicordatum Presl, Epimel. hot. 155.
arise obliquelyfrom the thick rhizome in a circular 1849. TYPE: "In sylvis Columbia," without col-
lector (holotype, PRC!).
pattern, from the center of which spring several Linnaea 36: 149. 1869.
Plagiogyria costaricensis Kuhn,
dimorphic, fertile leaves. Stems in Plagiogyria, TYPE: Costa Rica, Volcan Barba, Wendland 1066
however, lack indument whereas those of Blech- (holotype, c?; drawing, B!).
num are scaly, an easily observed character which Plagiogyria latifolia Copel., Philipp. J. Sci. 38: 411.
1929. TYPE: Peru, Huanuco, Cani, 7 mi NE of
immediately distinguishes the former from simi-
Mito, ca. 2600 m, Macbride 3432 (holotype, us!;
lar-appearing Blechnum species.
isotypes, F!, OH!).
Plagiogyria denticulata Copel., Philipp. J. Sci. 38: 4 1 2.
TYPE: Bolivia, Dept. Santa Cruz, alt. 2600 m,
I. Plagiogyria Herzog 1954 (holotype, us!).

Petiole and rachis stramineous, the former brown


Plagiogyria (Kunze) Mett., Abh. Senckenberg Na-
turf. Ges. 2: 265. 1858. TYPE: Lomaria eu-
at base, 8-30 cm long. Sterile lamina to 60 cm
phlebia Kunze
= Plagiogyria euphlebia long and 18 cm broad,
-pinnate to deeply pin-
1

natisect, lanceolate-elliptic, tapering to a pinna-


(Kunze) Mett. Figure 14.
tifid apex, gradually reduced to base, where pinnae
Terrestrial. Leaves to 2 m
long, pinnatisect to
are 2
as long as central ones; pinnae contiguous
'/2- /j

at base with adjacent ones, or widely spaced, 3-


1 -pinnate, dimorphic (fertile ones erect, common-
ly with longer petioles and narrower, constricted
10 cm long, 0.5-1.0 cm broad, margins serrulate
to biserrate, apex acute to short-attenuate; veins
pinnae). Sterile pinnae subentire to serrulate or
biserrate. Veins free, simple, paired at or near the commonly 1-2-forked, or a few simple or paired
lamina slightly longer and nar-
at the costa. Fertile
base, or forked. Fertile pinnae subentire to erose,
the margins at first slightly to strongly reflexed to rower than the sterile; pinnae very widely spaced,

protect developing sporangia, later spreading, or commonly 3-4 mm


broad, margins strongly re-
sometimes so strongly retroflexed that both edges troflexed at maturity.

touch on the adaxial side.


At edges of forests, ravines, on shaded clay and
The most recent studies on Plagiogyria have rocks banks, 1800-3600 m, Cajamarca, Amazon-
as, San Martin, Huanuco.
recognized about 50 species in the genus; however,
these have been based on a number of mostly Cuba; Jamaica; Mexico; Guatemala; Costa Rica
to Peru and Bolivia.
overlapping or quantitative characters, e.g., degree
of serration on pinna margins, relative length of
basal pinnae, degree of branching of veins, none Cajamarca: Hualgayoc, Soukup & Carmona Fa.5008
(us). Amazonas: Prov. Chachapoyas, Summit of Puma-
of which seem to be consistent or significant.
Urcu SE of Chachapoyas, Wurdack 1159(F, GH, NY, uc,
Therefore, we believe the actual number of species us). San Martin: Prov. Mariscal Caceres, Puerta del
in Plagiogyria is more likely to be nearer 5. Based
1 Monte, Young 1980 (USM). Huanuco: Yanano, Macbride
on study of type collections and on Peruvian spec- 3830 (F, GH, us).

imens attributed to various species, we have con-


cluded that there is only one somewhat variable
species in Peru.

References Family 9: DICKSONIACEAE


COPELAND, E. B. 1929. The fern genus Plagio- Dicksoniaceae Bower, Origin land fl. 591. 1908,
gyria. Philipp. J. Sci., 38: 377-417. as Dicksonieae. TYPE: Dicksonia L'Her.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 101


5cm

FIG. 14. Plagiogyria semicordata: a-b, habit; c, apex of fertile leaf; d, apex of fertile pinna, (a from Cuatrecasas
5465, Colombia, F, b from Wurdack 1159, F, c-d from Little 9184, Colombia, F.)

102 FIELDIANA: BOTANY


Stem stout to massive, usually not branched, The Dicksoniaceae are a family of five genera
prostrate to decumbent to erect and then often and 3540 species. Two genera are in Peru.
arborescent, indurated, densely covered with usu- The family, with the possible exception of Cys-
ally long trichomes. Leaves usually large, ca. 1-3 todium of Malaysia, is natural and distinctive. It
m long, circinate in vernation, monomorphic to has been united with the Cyatheaceae, but that
dimorphic (the fertile nearly lacking green tissue family differs in having scales on the stem and an
and more complex than the sterile), pinnate, gla- abaxial sorus and the two are of uncertain affinity.
brous to pubescent. Petiole lacking stipules, not
articulate to the stem. Veins free. Sori marginal,

usually paraphysate with slender trichomes, en- Reference


closed within a usually firm and partly green adax-
ial indusium and a thinner, light brown abaxial TRYON, R. M., AND A. F. TRYON. 1982. Dick-
indusium, these separate or basally or fully joined. soniaceae, pp. 138-155, in Ferns and allied
Sporangia with a 4-6-rowed stalk and a complete, plants, Springer- Verlag, New York.
oblique annulus.

Key to Genera of Dicksoniaceae

a. Lamina 4-5-pinnate-pinnatifid, broadest at the base, its tertiary axes grooved on the adaxial side
I. Culcita

a. Lamina 2-3-pinnate-pinnatifid (in America), reduced at the base, its tertiary axes ridged (not grooved)
on the adaxial side . II. Dicksonia

I. Culcita Dicksonia coniifolia Hooker, Sp. fil. 1: 70, t. 24A.


1 844. TYPE: Venezuela, (Dist. Federal), Caracas,

Linden 538 (holotype, K; isotype, BR; photo, GH).


Culcita Presl, Tent, pterid. 135. 1836. TYPE: Cul-
cita macrocarpa Presl. Figure 15.
Stem
prostrate to 1 m
tall, the apex with few

Terrestrial. Stem prostrate, decumbent or rarely


leaves.Leaves to 3 m
long, the petiole with a dense

m tall, stout. Leaves monomorphic, covering of trichomes at the base, often longer than
erect and to 3
the lamina. Lamina deltoid, to 4- or 5-pinnate-
to ca. 3 m long. Lamina 4-5-pinnate or slightly
pinnatifid at the base, the major segments stalked,
more complex, the tertiary axes grooved adaxially,
thinly pubescent, or glabrous with age, sterile ul-
veins free. Sori marginal, the adaxial indusium
timate segments with bluntly acute lobes. Sorus
joined basally to the thinner abaxial indusium, or
single on each ultimate lobe.
the two separate. Spores tetrahedral-globose, tri-
In wet elfin forests at ca. 2400-3100 m, Ama-
lete, nearly smooth, rugulose, or tuberculate.
zonas south to Cuzco.
Southern Mexico, Central America, Greater
Culcita coniifolia of tropical America and C.
Antilles, Venezuela and Colombia south to Peru;
macrocarpa of the Canary Islands, Azores, and
Mt. Itatiaia, Brazil.
Spain form subg. Culcita. The other five species, and
Culcita coniifolia is a distinctive species
of Malaysia to Samoa and Australia, comprise the
should be mistaken for no others. Although of
subgenus (or, better, the genus) Calochlaena Max-
wide distribution, it has been rarely collected in
on.
Peru.

Ama/onas: Prov. Chachapoyas, 18 km above Lei-


1 Culcita coniifolia (Hooker) Maxon, Annual Rep. road to Balsas, Cerros de Calla Calla,
.
mebamba on
Smithsonian 1911: 488. 1 9 1 2; J. Wash. Acad. Hutchison & Wright 5687 (F, GH). Along Rio Ventilla,
Sci. 12:456. 1922. Figure 15. 1-2 km W
of Molinopampa, Wurdack 1468 (GH, USM).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 103


FIG. 1 5. Culcita coniifolia: a, stem and part of petiole; b, pinna; c, rachis and base of pinna, adaxial side; d, fertile
ultimate segment, (a from Macbride 4519, r, b-d from Lent 732, Costa Rica, F.)

104 FIELDIANA: BOTANY


Huanuco: Playapampa, Macbride 4519 (F). Pasco: Prov. 4-pinnate, reduced at the base, tertiary axes adax-
Oxapampa, San Alberto, Cordillera de Yanachaga, van ially ridged (not grooved), veins free. Sori mar-
der Werffet al. 8436 (MO). Cuzco: El Dorado, Rio Uru-
ginal, the adaxial indusium joined basally to the
bamba valley, Aug. 12, 1941, Bites (GH). Prov. La Con-
vention, Dudley 10712 (GH).
thinner abaxial indusium. Spores tetrahedral-glo-
bose, trilete, granulate or reticulate.

II. Dicksonia Dicksonia is a genus of about 20 species, from


Malaysia to Samoa, St. Helena, tropical America,
Dicksonia L'Her., Sert. angl. 30. 1788. TYPE: and the Juan Fernandez Islands.
Dicksonia arborescens L'Her. Figure 16.

Terrestrial. Stem erect, to 10 m tall, or decum- Reference


bent at the base, usually massive. Leaves mono-
morphic to dimorphic (the fertile nearly lacking STOLZE, R. G. 1 976. Dicksonia, in Ferns and fern
and more complex than the sterile), allies of Guatemala. Part I. Fieldiana, Bot., 39:
green tissue
to ca. 3.5 m long. Lamina 2-pinnate-pinnatifid to 99-102.

Key to Species of Dicksonia

a. Tertiary segments of the larger fertile pinnae each with few to several sori 1 D. sellowiana.

a. Tertiary (ultimate) segments of the larger fertile pinnae each with a single sorus ... 2. D. stuebelii

1. Dicksonia sellowiana Hooker, Sp. til. 1: 67. In wet woods or cloud forests, at 1 550-2400 m,
1844. TYPE: Brazil, Sellow (lectotype desig- Cajamarca and Amazonas, south to Cuzco.
nated here, K; isolectotype, HBG; photos, GH, Southern Mexico, Central America, Venezuela,
us of HBG; lectoparatype, Brazil, Miers, K). and Colombia south to Bolivia, Paraguay, Uru-
Figure 16. guay and southeastern Brazil.
Dicksonia sellowiana is a widespread and vari-
Balantium karstenianum Klotzsch, Linnaea 20: 444. able species. Several segregates are sometimes rec-
1847. TYPE: "Columbia," Karsten, no. 9 (Coll.
ognized but these evidently represent minor vari-
II) (holotype, B?; isotype, HBG; photo, GH of HBG).
ations. The other American species are D. stuebelii
Dicksonia gigantea Karsten, Fl. columb. 2: 1 77, 1. 193.
1 869. TYPE: Colombia, (Cundinamarca), Andes
of Peru and D. berteriana (Colla) C. Chr. of the
of Bogota, Guadeloupe, Karsten (not located) Juan Fernandez Islands.
(Karsten, HBG; photo, GH, may be authentic).
Dicksonia karsteniana (Klotzsch) Moore, Index fil.
Cajamarca: Prov. Cutervo, La Pucarilla, Ldpez & Sa-
190(1860), 313(1861). gdstegui 5457 (GH, HUT). Amazonas: Prov. Bagua, ca. 20
Dicksonia spruceana Kuhn, Linnaea 36: 153. 1869. km E of La Peca, Harbour 2870 (MO). Huanuco: Cerros
TYPE: Peru, (San Martin), Tarapoto, Spruce 4728 del Sira, Rio Llullapichis watershed, Dudley 13254 (F,
(holotype, B; isotypes, A!, BM, GH!; photos, GH, GH, MO), 75259, 13373 (GH). Pasco: Oxapampa (as Jun-
MO, uc of BM). in), Soukup 2334 (F, GH). Prov. Oxapampa, Canyon de
Huancabamba, Ledn 614 (F). Cuzco: Prov. La Conven-
Stem to 10 m tall, enclosed, at least basally in tion, Cordillera Vilcabamba, Dudley 10439, 10603,

dense fibrous roots, with persistent leaf bases, to 11308(GH).


ca. 25 cm in diameter, bearing many leaves in a
crown. Leaves to 3 m
long, the petiole very short, 2. Dicksonia stuebelii Hieron., Hedwigia 45: 228,
densely covered with long trichomes. Lamina 2- /. 72, / /. 1906. TYPE: Peru, (Amazonas),
3-pinnate-pinnatifid, the secondary and tertiary Tambo Ventilla, Pascomayo to Moyobamba,
segments nearly sessile, glabrous to thinly pubes- Stiibel 1076 (holotype, B).

cent abaxially, sterile lobes bluntly acute to acute,


the margin flat or curved. Sori few to several on Similar to Dicksonia sellowiana, but the lamina
the tertiary segments of the larger fertile pinnae. is 2-pinnate-pinnatifid to 2-pinnate-pinnatisect,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 105


FIG. 16. Dicksonia sellowiana: a, 2 pinnae; b, fertile ultimate segments, abaxial side; c, secondary axis, adaxial
side,with ultimate segments, (a from Reiss 61, Brazil, F, b from Killip & Smith 18095, Colombia, F, c from Holm-
Nielsen el al. 5645, Ecuador, F.)

106 FIELDIANA: BOTANY


the larger fertile pinnae have the tertiary (ultimate) I. Lophosoria
segments bearing a single sorus, and the sterile
margins are recurved. Lophosoria Presl, Gefassbiindel Farm 36. 1847.
TYPE: Lophosoria pruinata (Sw.) Presl = Lo-
In the Jalca zone and in wet sandy soil, ca. 2200- phosoria quadripinnata (Gmelin) C. Chr. Fig-
3400 m, Amazonas. ure 17.
Endemic in northern Peru.
This is evidently a rare and local species. It is Terrestrial. Leaves rarely 0.25 m , usually 2-3
unusual in growing within the range of the related m, sometimes to 5 m long. Lamina 2-pinnate-
Dicksonia sellowiana. pinnatifid to 3-pinnate-pinnatisect, segments often
glaucous abaxially and slightly to densely pubes-
Amazonas: Cerro de Fraijaco (Huaui-Huni), NE of cent. Sori round, borne on the veins, the receptacle
Tambo de Ventilla. Pennell 15855 (GH). Prov. Chach-
hardly elevated. Spores trilete, tetrahedral-glo-
apoyas, 3-6 km W
of Molinopampa, Wurdack 1410
bose, with a large equatorial flange, coarsely tu-
(GH). Cerro Yama-uma, above Taulia, 12-15 km SE of
Molinopampa, Wurdack 1679 (F, GH, uc, us, USM). berculate to rugose.

Amonotypic genus, ranging from Mexico and


the Greater Antilles, south to Bolivia and Brazil,

LOPHOSORIACEAE also in southern Argentina and Chile, and the Juan


Family 10:
Fernandez Islands.

Lophosoriaceae Pic.-Ser., Webbia 24: 700. 1970. Lophosoria quadripinnata is a widely distrib-
TYPE: Lophosoria Presl.
uted and distinctive species of the cloud forest
zone. One high altitude variety that is local in

Stem sometimes branched, massive, decumbent Ecuador and Peru may be recognized.
to erect and arborescent, indurated, densely cov-
ered by long trichomes. Leaves usually large, ca.
1-5 m
long, circinate in vernation, monomorphic,
1. Lophosoria quadripinnata (Gmelin) C. Chr.,
more or Skottsb. Nat. hist. Juan Fernand. 2: 16. 1920.
pinnate, less pubescent. Petiole lacking
stipules, not articulate to the stem. Veins free. Sori
exindusiate, on the abaxial surface of the segments, Stem apex with few leaves. Petiole about as long
as the lamina or somewhat shorter, with 3 con-
paraphysate with slender trichomes. Sporangia
with a short, 6-rowed stalk and a complete, oblique voluted vascular bundles. Lamina very narrowly
annulus. lanceolate to usually broadly ovate, to 3-pinnate-
pinnatisect, with sessile or short-stalked segments,
The family Lophosoriaceae contains a single,
very sparsely to densely pubescent and often glau-
American genus. cous abaxially, ultimate segments obtuse to sub-
acute. Sori single on a fertile vein.

Reference Range of the genus.

TRYON, R. M., AND A. F. TRYON. 1982. Lopho-


soriaceae, pp. 156-161, in Ferns and allied
plants, Springer- Verlag, New York.

Key to Varieties

a. Leaf ca. 2-4 rn long, pinnae spaced, patent, the central ones ca. 0.5-1.0 m long
la. var. quadripinnata
a. Leaf ca. 0.25-1 m long, pinnae imbricate, ascending, the central ones ca. 0.05-0.20 m long
. Ib. var. contracta

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 107


FIG. 1 7. Lophosoria quadripinnata var. quadripinnata: a, stem and portion of petiole; b, pinna; c, base of fertile
segment, (a from Wurdack 1752, F, b-c from Stork & Morton 10348, F.)

108 FIELDIANA: BOTANY


1 a. Lophosoria quadripinnata var. quadripinnata. Amazonas: Prov. Chachapoyas, Pomacocha, Lopez el
al.4392 (GH). Prov. Chachapoyas, Cerros de Calla Calla
Figure 17.
slopes, Wurdack 1752 (GH, us, USM). Prov. Chachapoyas,
between Leimebamba and Calla-Calla, Smith & I'dsquez
Polypodium glaucum Sw., Prodr. 1 34. 1 788, not Houtt.
4978 (GH). La Libertad: Prov. Bolivar, cerca Nevado de
1 783. TYPE: Jamaica, Swam (holotype, B!, Herb.
Cajamarquilla, Ferreyra 1349 (GH, USM).
Willd. 19723; photo, GH, designated as type by
Hieronymus, it may be the holotype or an iso-
type).
Polypodium quadripinnatum Gmelin, Syst. nat. 2(2):
1314. 1791, nom. nov. for Polypodium glaucum
Sw. (not Houtt.) and with the same type.
Polypodium pruinatum Sw., J. Bot. (Schrader) 1800(2):
29. 1802, nom. nov. for Polypodium glaucum Sw. Family 11: METAXYACEAE
(not Houtt.) and with the same type.
Alsophila pruinata (Sw.) Kunze, Linnaea 9: 99. 1834.
Farm Metaxyaceae Pic.-Ser., Webbia 24: 701. 1970.
Lophosoria pruinata (Sw.) Presl, Gefassbiindel
37. 1847.
TYPE: Metaxya Presl.
Alsophila quadripinnata (Gmelin) C. Chr., Index fil.

44. 1905. Stem rather stout, sometimes branched, pros-


trate to nearly erect, indurated, with dense, long
on brushy slopes,
In wet forests, forest borders,
trichomes, especially toward the apex. Leaves usu-
in cloud forests and elfin forests, 700-3100 m, ally large, to 2 m long, circinate in vernation,
Cajamarca and Amazonas, south to Puno. monomorphic, pinnate, glabrate. Petiole lacking
Range of the genus. stipules, not articulate to the stem. Veins free. Sori
Throughout most of mountainous tropical exindusiate, on the abaxial surface of the pinnae,
America, this variety is a typical element of the
paraphysate with slender trichomes. Sporangia
cloud forest zone. with a 4-rowed stalk and a complete, slightly
oblique annulus.
Cajamarca: Prov. Celendin, Agua Colorado, Sanchez
206 (GH). Amazonas: Cerros de Calla Calla, above Lei-
mebamba, Hutchison & Wright 5686 (F, GH, uc, us). The family Metaxyaceae contains a single,
Cerro Puma Urco, Chachapoyas, Soukup 4086 (F, us). American genus.
San Martin: Monte Campana, Tarapoto, Spruce 4248
(GH, us). Huanuco: Panao, Bryan 407 (F, GH). Playapam-
pa, Macbride 4860 (F, us). Pasco: Quillasu, Soukup 3290
Reference
(F,GH, uc, us). Yapas, Pichis Trail (as Junin), Killip &
Smith 25456 (F, GH, us). Junin: Huacapistana, Ferreyra
3654 (USM). Ucayali: La Divisoria, Aguilar 849 (GH). TRYON, R. M., AND A. F. TRYON. 1982. Metax-
Huancavelica: Prov. Tayacaja, Surcubamba, Stork &
yaceae, pp. 162-165, in Ferns and allied plants,
Horton 10348 (F, GH, uc, us). Cuzco: Prov. La Conven-
Springer- Verlag, New York.
tion, Dudley 10883 (F, GH). Pilahuata, Cerro de Cusil-
luyoc, Pennell 13938 (GH, us). Puno: Prov. Sandia, San-
dia to Chunchusmayo, Weberbauer 1333 (USM).
I. Metaxya

1 b. Lophosoria quadripinnata var. contracta (Hi- Metaxya Presl, Tent, pterid. 59. 1836. TYPE: Me-
eron.) R. & A. Tryon, Rhodora 84: 1 26. 1 982. taxya rostrata (HBK.) Presl (Aspidium rostra-
turn HBK.). Figure 18.
Alsophila contracta Hieron., Hedwigia 45: 236, /. 75,
/ 8. 1906. SYNTYPES: Peru, (Amazonas), near Terrestrial or rarely on tree bases. Leaves to ca.
Inez and Calle-calle, Stitbel 1067 (B!; frag., GH!);
(Amazonas), near Challuayacu and Tambo Cen- 2 m long. Petiole usually with short trichomes at

tamala, Stubel 1066 (B!). the very base. Lamina 1 -pinnate, with simple pin-
nae, veins free. Sori roundish to elongate, the re-
Wet, shrubby areas, 2800-3500 m, in Amazon- ceptacle nearly flat. Spores globose, trilete, gran-
as and La Libertad. ulate.
Ecuador and Peru.
The var. contracta is evidently a high-altitude A monotypic genus ranging from southern Mex-
ecotype. The following collection is intermediate icoand Central America; Guadeloupe; Trinidad;
between the two varieties: Peru, Amazonas, Puma- and French Guiana west to Colombia and south
Urcu, SE of Chachapoyas, Wurdack 791 (us). to Bolivia; Amazonian Brazil.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 109


FIG. 18. Metaxya rostrata: a, habit; b, portion of fertile pinna, abaxial side. (From Wurdack 1872, F.)

110 FIELDIANA: BOTANY


1. Metaxya rostrata (HBK.) Presl, Tent, pterid. rarely entire, glabrous, pubescent and (or) scaly
60. 1836. Figure 18. abaxially. Petiole lacking stipules, not articulate to
the stem. Veins free or rarely reticulate. Sori ex-
Polypodium rostratum Willd., Sp. pi. ed. 4, 5: 193. indusiate or indusiate, on the abaxial surface of
1810, not Burm. 768. TYPE: Venezuela, Javita,
1
the segments, usually paraphysate with slender tri-
Humboldt 966 (holotype, B!, Herb. Willd. 19691;
chomes or rarely with enlarged ones. Sporangia
photo, GH).
with a 4-rowed stalk and a complete, oblique an-
Aspidium rostratum HBK., Nov. gen. sp. 1: 12. 1815,
nom. nov. for Polypodium rostratum Willd. (not nulus.
Burm.) and with the same type.
Alsophila blechnoides Hooker, Sp. fil. 1: 35. 1844, The Cyatheaceae are a family of six genera and
nom. superfl. for Alsophila rostrata (HBK.) Mart.
a few hundred species. All of the genera occur in
and with the same type.
Peru.

Stem prostrate, the apex with few leaves. Petiole


This treatment has been adapted from the lit-

erature cited under the family and the genera. There


about as long as the lamina, with 1 convoluted
are at least 1 9 species, especially of Ecuador but
vascular bundle. Lamina narrowly lanceolate to
also of Bolivia, that may well also occur in Peru,
ovate, 1 -pinnate, with a conform terminal pinna,
but these are too numerous to mention.
the pinnae stalked, glabrous or nearly so abaxially,
Characters of all parts of the leaf are useful and
entire to finely serrate, the serrate apex acuminate
often necessary for the accurate identification of
to caudate. Sori 1-3 on each fertile vein.
the genera and species of Cyatheaceae. Of special

Woods and dense forests, usually in sandy, moist, importance are characters of the petiole scales.
well-drained soil, 100-800 m, Amazonas to Cuzco However, the scales may be abraded on the petiole
of a mature leaf and their characters are best seen
and Madre de Dios.
on the croziers. The height and diameter of the
Range of the genus.
arborescent stem vary in a species depending upon
Metaxya is unusual in having the 1-pinnate-
the age of the plant and the growing conditions.
pinnatifid leaves of juvenile plants more complex
than the 1 -pinnate leaves of adult plants.
Accordingly, these characters as well as other
quantitative ones are usually not included in the
Amazonas: Prov. Bagua, Rio Maranon, above Cas- species descriptions.
cadas de Mayasi, Wurdack 1872 (us, USM). San Martin: Several special terms that are used in the Cy-
Prov. Mariscal Caceres, Isla de Pucunuchu, J. Schunke atheaceae are the following.
4783 (F, GH). Prov. Mariscal Caceres, Rio Sion, J. Schunke PETIOLE SPINES Corticinate: The spine is an ex-
3524 GH, us). Loreto: Iquitos, Killip & Smith 27489
(F,
tension of the cortex of the petiole; each bears, at
(GH, us). Huanuco: Prov. Pachitea, Bosque Nacional de
least when young, a scale at its apex which is rather
Iparia, J. Schunke 1620, 1833 (F, GH, us). Pasco: Prov.
Oxapampa, Cordillera San Matias, Leon 332 (USM). blunt when the scale falls off. Squaminate: The
Ucayali: vicinity of Aguaytia, Croat 20966 (us). Prov. spine is not an extension of the petiole; it breaks,
Coronel Portillo, Bosque Nacional Humboldt, Vdsquez or can be broken, from the petiole at its base. It
3910 (F, MO). Cuzco: Prov. Quispicanchi, Vargas 16462
has a sharp apex that does not bear a scale. These
(GH). Prov. Paucartambo, Vargas 1 1245 (GH). Madre de
Dios: Prov. Tambopata, Tambopata Nature Reserve, spines have evidently evolved from petiole scales.
Barbour5191 (F). PETIOLE SCALES Conform: All cells, except those
that may be borne on the edge, are similar in ori-

entation, shape, and usually in size and color.


Marginate: With a narrow to broad margin of cells
Family 12: CYATHEACEAE different from those in the central portion of the
scale in orientation, size, and usually in shape and
Cyatheaceae Kaulf., Wesen Farrenkr. 119. 1827. color. Body: All of the scale except for cilia, teeth,
TYPE: Cyathea Smith. or setae borne on the edge. Margins: The cellularly
differentiated areason each side of the central por-
Stem usually massive, erect, arborescent, un- tion of marginate scales. Concolorous: All of the
branched, very rarely slender and scandent, to scale of one color or nearly so. Bicolorous: The
small, decumbent and short-creeping, indurated, margins definitely of lighter color than the central
bearing scales and sometimes spines. Leaves usu- portion. Concordantly bicolorous: The lighter col-
ally large, ca. 1-4 m
long, circinate in vernation, or confined to the differentiated margins and the
monomorphic to rarely dimorphic, pinnate or very darker color to the central portion of elongate cells.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. Ill


Discordantly bicolorous: The lighter color of the the sporangia except for a rather small opening at
margins extends into part of the central portion of the apex. Sphaeropteroid: Completely enclosing
elongate cells. the sporangia and with an apical umbo.
PETIOLE SCURF The scales are
Large scales:
small, but definitely larger than most of the scurf
which is composed of minute scales and tri- References
chomes. TRYON, R. 1970. The classification of the Cy-
INDUSIA Hemitelioid: Attached at the base and atheaceae. Contr. Gray Herb., 200: 2-53.
often at the sides of the receptacle, but not com- . 1986. Cyatheaceae, in Harling, G., and
pletely surrounding Meniscoid: Completely sur-
it. Anderson, eds., Flora of Ecuador, 27: 17-56.
L.

rounding the receptacle, nearly flat or with slightly TRYON, R. M., AND A. F. TRYON. 1982. Cy-
upturned edges. Cyathiform: Cup-shaped. Urceo- atheaceae, pp. 166-212, in Ferns and allied
late: Urn-shaped. Subsphaeropteroid: Enclosing plants, Springer- Verlag, New York.

Key to Genera of Cyatheaceae

a. Petiole scales conform I. Sphaeropteris


a. Petiole scales marginate b
b. Petiole scales with a dark, opaque apical seta c
c. Petiole lacking spines, or with corticinate spines that bear, at least when young, a scale at the
apex, other petiole scales almost appressed to the surface and attached at a usually pseudopeltate
base; croziers lacking spines II. Alsophila

c. Petiole with squaminate spines, many of these large, black, with a sharp apex; petiole scales
patent, attached at a usually narrowed base; croziers with large, black, sharp spines
III. Nephelea
b. Petiole scales lacking a differentiated apical seta, the apex rounded to filamentous d
d. Indusium absent IV. Trichipteris
d. Indusium present e
e. Spores lacking large pores, sometimes with variously distributed small pits or pores; veins
free, the basal ones of adjacent segments extending to the margin above the sinus, rarely a
few costal areolae present V. Cyathea
e. Spores with three large equatorial pores, often also with smaller pits or pores; veins anas-
tomosing to form costal areolae, or if free then the basal veins of adjacent segments connivent
to the sinus . VI. Cnemidaria .

I. Sphaeropteris borne at the fork of a vein, exindusiate or with a


hemitelioid or sphaeropteroid indusium. Spores
Sphaeropteris Bernh., J. Bot. (Schrader) 1800(2): tetrahedral-globose, trilete, echinate, porate, ver-
122. 1802. TYPE: Sphaeropteris medullaris ruca te or nearly smooth.
(Forster) Bernh. (Polypodium medullare Fors-
ter). Figure 19. Sphaeropteris occurs in tropical America, India
and southeastern Asia to New Zealand and Pit-
Terrestrial. Stem stout, erect and arborescent or cairn Island. Among the 120 species, there are 23
rarely short-decumbent, lacking spines. Leaves in America and seven in Peru.

monomorphic to slightly dimorphic, with scales,


especially on the croziers and the base of the pet-
iole, that are conform and with or without a dark References
apical seta. Petiole lacking spines or with corticin-
ate spines. Lamina 1 -pinnate to 3-pinnate-pin- TRYON, R. 1971. The American tree ferns allied
or rarely partially anastomosing
natifid, veins free to Sphaeropteris horrida. Rhodora, 73: 1-19.
without included free veinlets. Sori round, often WINDISCH, P. G. 1977. Synopsis of the Genus

112 FIELDIANA: BOTANY


FIG. 19. Sphaeropteris elongata: a, portion of pinna. Sphaeropteris quindiuensis: b, portion of fertile segment; c,

portion of petiole scale, (a from Soukup 1651, F, b from Soukup 2336, F, c from Plowman 6062, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 113


Sphaeropteris (Cyatheaceae) with a revision of . 1978. Sphaeropteris (Cyatheaceae), the
the neotropical exindusiate species. Bot. Jahrb. systematics of the group of Sphaeropteris hir-
Syst, 98: 176-198. suta. Mem. New York Bot. Gard., 29: 2-22.

Key to Species of Sphaeropteris

a. Petiole scales without a dark apical seta and without dark marginal setae, rarely the margins ciliate

toward the apex [subg. Sclephropteris Windisch] b


b. Veins free c
c. Sori exindusiate 1 . S. aterrima
c. Sori indusiate, the indusium sometimes inconspicuous d
d. Sori with a small, hemitelioid (flabellate) indusium e
e. Costa and costules with whitish, fimbriate or highly dissected scales abaxially

2. S. rufescens

Costa and costules lacking scales or with scales various, but not whitish, fimbriate, or
e.

highly dissected 3. S. macrosora

d. Sori with a sphaeropteroid indusium 5. S. atahuallpa

b. Veins regularly anastomosing, especially along the costa, sori with a hemitelioid indusium ....
4. S. bradei
Petiole scales with a dark apical seta and dark marginal setae [subg. Sphaeropteris] f
f. Sori exindusiate 6. S. elongata
f. Sori with a sphaeropteroid indusium 7. S. quindiuensis

1 .
Sphaeropteris aterrima (Hooker) Tryon, Contr. 2. Sphaeropteris rufescens (Kuhn) Windisch,
Gray Herb. 200:20. 1970. Bradea 1: 372. 1973.

Ahophila aterrima Hooker, Syn. fil. 38. 1866. TYPE: Hemitelia rufescens Kuhn, Linnaea 36: 159. 1869.
Peru, (San Martin), Tarapoto, Spruce 4713 (ho- TYPE: Peru, (San Martin), Monte Guairapurima,
lotype, K!; isotypes, P!, us!). Tarapoto, Spruce 4727 (holotype, B!; isotype, K.;
Cyathea aterrima (Hooker) Domin, Pterid. 262. 1 929. photo, GH of K).
Cyathea rufescens (Kuhn) Domin, Pterid. 264. 1 929.

Petiole without spines or sparingly short-acu-


leate, with very narrow scales intergrading to tri- Petiole muricate, pubescent, scales brown with
chomes, these rather dense, especially toward the lighter margins to wholly light colored, the edges
base of the petiole, straw-colored to light brown, Lamina
toothed. 2-pinnate-pinnatifid, with tri-
lacking a dark apical seta and dark marginal setae, chomes, bullate scales,and long whitish, fimbriate
sometimes ciliate. Lamina 1-2-pinnate-pinnatifid, or highly dissected scales abaxially, veins free. Sori
pinnules long-pubescent and sometimes with a few usually at the fork of a vein, paraphyses longer
flattish scales abaxially, veins free. Sori with pa-
than the sporangia, indusium hemitelioid (flabel-
raphyses about as long as the sporangia, exindu- late).
siate.

This species has been collected only in San Mar-


Known only from the type collection from San
Martin.
tin at mid-elevations.
Endemic to Peru.
Colombia and Peru.
The whitish fimbriate scales beneath and the
Sphaeropteris aterrima is notable for its petiole
small indusium make this a distinctive species.
indument which consists of scales intergrading to
trichomes.

San Martin: Monte Morro de Moyobamba, Stiibel 3. Sphaeropteris macrosora (Baker) Windisch,
/77J(B, NY). Bradea 1: 372. 1973.

114 FIELDIANA: BOTANY


Alsophila macrosora Baker, Timehri 5: 211. 1886. San Martin: Prov. Lamas. Caserio Bonilla, km. 75 of
TYPE: Venezuela, (Bolivar), Ml. Roraima, 1m Tarapoto-Yurimaguas road, Knapp & Mallet 7141 (MO).
Thurn 87 (holotype, K!; isotype, us!). Loreto: Mishuyacu, near Iquitos, Klug 1546 (F, NY, us).
Cyathea macrosora (Baker) Domin, Pterid. 263. 1929. Estacion Biologica Callicebus. Rio Nanay, 2 horas rio
urn ha de Iquitos, Vasque: el al. 649 (GH). Near mouth
of Rio Napo, Croat 20194 (F, uc).
Petiole glabrous to sparingly pubescent, nearly
smooth to muricate with the scales whitish to
brownish and with lighter margins, these some-
times broad, or most of the scales wholly whitish,
5. Sphaeropteris atahuallpa Tryon, Rhodora 74:
442. 1972. TYPE: Peru, Amazonas, Prov.
lacking a dark apical seta and dark marginal setae,
sometimes ciliolate, especially Lamina 2- Chachapoyas, Cerros de Calla Calla, above
apically.
Balsas on road to Leimebamba, Hutchison &
pinnate-pinnatifid, the pinnules glabrous, pubes-
cent and (or) slightly scaly abaxially, the scales Wright 6922 (holotype, GH!; isotypes, MO, uc).
various, but not whitish, fimbriate, or highly dis-
Petiole sparingly very short-aculeate, with large,
sected. Sori with paraphyses longer than the spo-
broad scales to 6 cm long, these dense well above
rangia, indusium hemitelioid (broadly flabellate to
the base of the petiole, brownish white, lacking a
reduced).
dark apical seta and dark marginal setae, sparsely

At 400-500 m, Pasco.
and finely ciliate, especially toward the apex. Lam-
ina 2-pinnate-pinnatifid to 2-pinnate-pinnatisect,
Guyana and Brazil, west to Colombia and Peru.
pinnules with prominent whitish, flattish scales
Sphaeropteris macrosora represented in Peru
is
abaxially, veins free. Sori with paraphyses about
by var. reginae Windisch which occurs throughout
as long as the sporangia, indusia sphaeropteroid.
the range of the species. Variety macrosora is in
Guiana and Venezuela and var. vaupensis
British
This species has been collected in Peru only at
Windisch is in Colombia and Venezuela.
ca. 3000-3300 m
in the Department of Ama-
zonas.
Pasco: Prov. Oxapampa, Palcazu, Foster & d'Achille
10168 Ecuador and Peru.
(F).
Sphaeropteris atahuallpa is a distinctive and at-
tractive species. The
croziers are completely en-
veloped by large, whitish scales and these form a
4. Sphaeropteris bradei Windisch, Bradea 1: 372. dense covering to the petiole. Some of the scales
1973. TYPE: Colombia, Vaupes, Cerro Mitu, cm
are up to 6 long and several cells thick at their
Schultes, Raffauf& Soejarto 2422 9 (holotype, base.
GH!).
Amazonas: Summit of Puma-Urcu, SE of Chachapoy-
Petiole sparingly aculeate, with rather broad as, Wurdack 1153 (GH, us). Prov. Chachapoyas, Calla-
scales, these usually dense, especially toward the Calla, Smith & Vasquez 5012 (GH).

base of the petiole, shining, dark brown, lacking a


dark apical seta and dark marginal setae, usually
6. Sphaeropteris elongata (Hooker) Tryon, Contr.
finely short-ciliate. Lamina
2-pinnate-pinnatifid,
Gray Herb. 200: 20. 1970. Figure 19a.
pinnules with scattered, short, often crispate tri-
chomes and sometimes a few bullate scales abax-
Alsophila elongata Hooker, Sp. fit. 1 : 43. 1 844. TYPE:
veins regularly anastomosing, especially along
ially,
Colombia, Hartweg 1528 (holotype, K!; isotype,
the costa. Sori with paraphyses shorter than the
LD), not Cyathea elongata Karsten.
sporangia, indusium hemitelioid, usually 2-lobed. Alsophila poeppigii Hooker, Sp. fil. 43. 844. TYPE:
1 : 1

Peru, 1829, Poeppig (holotype, K!), a synonym of


Sphaeropteris elongata by Windisch, Bot. Jahrb.
Primary forests, 130-200 m, San Martin and Syst. 98: 189. 1977.
Loreto. Cyathea poeppigii (Hooker) Domin, Pterid. 263. 1929.

Colombia, Ecuador, and Peru.


The regularly anastomosing veins of Sphaer- Petiole aculeate, with rather narrow scales, these
opteris bradei are unique among the American Cy- usually sparingly persistent, whitish to light brown
atheaceae, except for species of the genus Cnem- or with a darker base, with a dark apical seta and
idaria. dark marginal setae. Lamina 2-pinnate-pinnatifid

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 115


to 2-pinnate-pinnatisect, pinnules sparingly to def- On steep mountain slopes, in rain forests and
initelypubescent and sometimes with a few flattish cloud forests, 1700-2500 m, Amazonas south to
scales abaxially, veins free. Sori with persistent Cuzco.
paraphyses mostly longer than the sporangia, ex- Colombia south to Bolivia.
indusiate. This species is unique among those of Peru in
having some of the scales at the top of the crozier
In primary forests, rain forests,and on open with the marginal teeth retrorse at the scale apex
hillsides, at 450-2000 m, Amazonas and Loreto, but changing to antrorse below.
south to Puno.
Costa Rica south to Bolivia; also in southeastern Amazonas: Prov. Bagua, Cordillera Colin, SE of La
Brazil. Peca, Barbour 3749 (MO). Huanuco: Prov. Leoncio Pra-
do, Cordillera Azul, Gentry et al. 29573 (GH). Playapam-
Sphaeropteris elongata is the most widely dis-
pa, Macbride 4856 (F). Pasco: Oxapampa (as Junin), Sou-
tributed species of the genus in America and con-
kup 2336 (F, GH). Prov. Oxapampa, road Oxapampa-
sequently is a variable species. Alsophila poeppigii Paucartambo, Smith & Pretel 1643 (F, MO). Junin: Prov.
was described on the basis of a plant with unusu- Chanchamayo, ca. 26 km S of San Ramon, Smith &
Palacios 2649 (F, MO). Cuzco: Prov. La Convention, Cor-
ally coriaceous leaves. Specimens without petiole
dillera Vilcabamba, Dudley 11313 (GH).
scales may be similar to those of Trichipteris con-
jugata, but that species has sessile rather than
stalked pinnae.
II. Alsophila

Amazonas: Prov. Bagua, Chiriaco to Puente Venezue-


la.Barbour 4385 (MO, USM). San Martin: Prope Tara- Alsophila R. Br., Prodr. 158. 1810. TYPE: Also-
poto. Spruce 4722 (GH). Loreto: Cerca a Pongo, entre phila australis R. Br. Figure 20.
Yurimaguas y Tarapoto, Ferreyra 17474 (GH, USM).
Huanuco: Huamalies, Valle de Monzon, Weberbauer Stem and arborescent or
Terrestrial. stout, erect
3471 (USM). Chinchao, Ferreyra 16950 (GH, USM). Junin:
La Merced, Soukup 1061 (F). Cerca a San Ramon, Cer- very rarely short and decumbent or climbing, lack-
raie 2870 (GH, USM). 10 km SW of San Ramon, Tryon ing spines or these rarely present. Leaves mono-
& Tryon 5445 (F, GH). Lcayali: 10 km NE of Aguaytia, morphic or sometimes dimorphic, with scales, es-
Gentry el al. 41422 (F, MO, USM). Ayacucho: Prov. La peciallyon the crozier and petiole base, that are
Mar, Hacienda Santa Rosa, Dudley 11697 (GH). Ayna,
marginate and with a usually dark apical seta. Pet-
between Huanta and Rio Apurimac, Killip & Smith 22716
iole lacking spines or with corticinate spines. Lam-
(NY, us). Puno: Prov. Carabaya, Puente Inambari, Var-
gas 18431 (GH). ina entire to 4-pinnate, veins free. Sori round, often
at the fork of a vein, exindusiate or with a hem-
itelioid to sphaeropteroid indusium. Spores tet-

rahedral-globose, trilete, prominently ridged.


7. Sphaeropteris quindiuensis (Karsten) Tryon,
Alsophila is a pantropical genus of ca. 230 species
Contr. Gray Herb. 200: 20. 1970. Figure 19b-
with 3 of them in America, mostly in the Greater
1
c.
Antilles, and a single one in Peru.
Conant (1983), the first 13
In the treatment of
Cyathea Quindiuensis Karsten, Linnaea 28: 454. 1 857.
TYPE: Colombia, (Tolima), between Rios Mag- species are Alsophila and the others are treated
dalena and Cauca, Karsten (not located). (Toli- here in the next genus, Nephelea.
ma), Paramo Quindio, Karsten (Herb. Mett. B! is
authentic).

Petiole without spines, with long, narrow scales, Reference


these dense, usually well above the base of the
petiole, whitish to shining brown, with a dark api- CONANT, D. S. 1983. A revision of the genus
cal seta and dark marginal setae. Lamina 2-pin- Alsophila (Cyatheaceae) in the Americas. J. Ar-
nate-pinnatifid to 2-pinnate-pinnatisect, pinnules nold Arbor., 64: 333-382.
with a few to many bullate scales, especially on
the costules of fertile segments, and often some
trichomes abaxially. Sori with paraphyses about 1 .
Alsophila engelii Tryon, Contr. Gray Herb. 200:
as long as the sporangia, indusium sphaeropteroid. 29. 1970, nom. nov. for Cyathea elongata

116 FIELDIANA: BOTANY


4 cm

3 mm

FIG. 20. Alsophila engelii: a, pinna; b, ultimate segment and son; c, apex of petiole scale. (From Dudley 11944,
OH.)

Karsten, not Alsophila elongata Hooker. Fig- Petiole smooth to tuberculate, brown, bearing
ure 20. mostly blackish to light brown scales with 1 apical
seta. Lamina 2-pinnate-pinnatifid, gradually ta-
pering to the apex, rachis light brown to brown,
with usua "y numerous sma11 dissected scales, pin-
>
Cyathea elongata Karsten, Fl. Columb. 2: 1 59. 1 869.
TYPE: Venezuela, (Merida), Merida, Engel 138
nae sessile or nearly so. Indusia deeply cyathiform
(holotype, B!). to sphaeropteroid, glabrous.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 117


Montane rain forests, 1 700-1 900 m, Amazonas, Terrestrial. Stem stout, erect and arborescent,
San Martin, and Ayacucho. sometimes branched, with spines. Leaves mono-
Venezuela and Colombia, south to Peru. morphic, with on the croziers and
scales, especially
petiole base, that are marginate and with a dark
Amazonas: Prov. Bongara, Shilla, Young & Eisenberg apical seta. Petiole with squaminate spines. Lam-
442 (uc). San Martin: Prov. Rioja, Pedro Ruiz-Moy- ina 1-3-pinnate-pinnatifid, veins free. Sori round,
obamba, D. Smith 4454 (GH). Ayacucho: Prov. La Mar,
usually at the fork of a vein, exindusiate or with
along Inca trail between Huanhuachayo and Punccu,
Cordillera Central, Dudley 11944 (GH).
a hemitelioid to sphaeropteroid indusium. Spores
tetrahedral-globose, trilete, with prominent short
to long ridges.

Comments
Nephelea is a tropical American genus of 17
species, with three in Peru. It ranges from southern
Mexico and the West Indies, south to northern
Alsophila paucifolia Baker of Colombia and Ec-
uador may be found in Peru. It differs from A. Argentina and southern Brazil, centering in the
Greater Antilles where nine species occur.
engelii in having a 1-pinnate-pinnatifid lamina. Al-
The squaminate spines on the petiole are dis-
sophila capensis (L. f.) John Sm. ssp. polypodioides
(Sw.) Conant of southeastern Brazil may also occur
tinctive of Nephelea. They are blackish, have a
in Peru. It is characterized by the highly dissected very sharp tip, and at least on old leaves they may
be broken off at their base.
aphlebiae at the base of the petiole.

Reference
III. Nephelea

GASTONY, G. J. 1973. A revision of the fern


Nephelea Tryon, Contr. Gray Herb. 200: 37.1 970.
TYPE: Nephelea polystichoides (Christ) Tryon genus Nephelea. Contr. Gray Herb., 203: 81-
= (Cyathea polystichoides Christ). Figure 21. 148.

Key to Species of Nephelea

a. Veins with stellate trichomes or squamules abaxially b


b. Indusium glabrous; pinna-rachises not green-alate between the more distal, sessile pinnules; petiole
scales on the abaxial side lacking lateral setae 1 N. erinacea
.

b. Indusium with whitish trichomes; pinna-rachises usually (sometimes narrowly) green-alate be-

tween the more distal, sessile pinnules; petiole scales on the abaxial side with dark lateral setae
2. N. cuspidata

a. Veins lacking indument abaxially, or present and not stellate; pinna-rachises green-alate between the
more distal, sessile pinnules; petiole scales lacking lateral setae, indusium with trichomes
.3. N. incana

1 .
Nephelea erinacea (Karsten) Tryon, Contr. Gray natifid, usually abruptly reduced to a pinna-like
Herb. 200: 40. 1970. apex, pinna-rachises not green-alate between the
more distal sessile pinnules, veins with minute,
Cyathea erinacea Karsten, Linnaea 28: 453. 1857. whitish, stellate squamules abaxially. Indusia cy-
TYPE: Venezuela, (Merida), Merida, 2000 m, athiform to urceolate or rarely subsphaeropteroid,
Karsten (holotype, not located; isotype, B!).
glabrous to densely scaly.
Alsophila erinacea (Karsten) Conant, J. Arnold Arbor.
64: 371. 1983.

In forests at ca. 1200-2000 m, Amazonas to


Scales on the abaxial side of the petiole with a Huanuco and Pasco.
dark apical seta and without lateral setae. Lamina Costa Rica south to Venezuela, Colombia, and
usually coriaceous, predominantly 2-pinnate-pin- Bolivia.

118 FIELDIANA: BOTANY


FIG. 2 1 Nephelea cuspidata: a. basal portion of pinna; b, expanding crozier, c, ultimate segments and sori; d,
.

apex of petiole scale, (a from Cuatrecasas 14843, Colombia, F, b, d from Davis 1109, Bolivia, F, c from Rimachi
481, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 119


Nephelea erinacea var. purpurascens (Sodiro) Prov. La Convention, Rio Apurimac, above Boca del
Tigre rapids, Davis et al. 1304 (F, GH). Madre de Dios:
Gastony has the indusium densely and persistently
Prov. Tambopata, SSW of Puerto Maldonado, Barbour
than glabrous to sparsely scaly as in
scaly, rather 5297 (F, MO). Puno: Prov. Carabaya, San Gaban, Puente
var. erinacea. Itis presently known only from a
Arica, Vargas 1891 9 (GH).
few volcanoes in Ecuador.

Amazonas: Prov. Bongara, 5 km N of N end of lake 3. Nephelea incana (Karsten) Gastony, Contr.
Pomacocha, on road to Rioja, Hutchison & Wright 6803 Gray Herb. 203: 137. 1973.
(GH, uc, USM). Rio Cenepa, 10 km E of Huampami,
Berlin 219 (F, MO). Huanuco: Macbride 4843 (F, us).
Cyathea incana Karsten, Fl. Columb. 1: 75, t. 37.
Pasco: Prov. Oxapampa, trail to summit of Cordillera
1860. TYPE: Colombia, (Cundinamarca), Andes
Yanachaga, D. Smith et al 7817, 7851 (F, MO). Prov. of Bogota, 2500 m, Lindig(nol located).
Oxapampa, Rio San Alberto, Smith & Pretel 7942 (USM). Alsophila incana (Karsten) Conant, J. Arnold Arbor.
64: 371. 1983.

2. Nephelea cuspidate (Kunze) Tryon, Contr. Gray Scales on the abaxial side of the petiole with a
Herb. 200: 40. 1970. Figure 21. dark apical seta and without lateral setae, or rarely
with an additional apical seta and lateral setae.
Cyathea cuspidata Kunze, Linnaea 9: 101. 1834. Lamina rigidly papyraceous, 2-pinnate-pinnatifid,
TYPE: Peru, (Loreto), Prov. Maynas, Feb. 1831,
abruptly reduced to a pinna-like apex, pinna-rach-
Poeppig diar. 2286 (holotype, LZ destroyed; iso-
ises green-alate between the more distal sessile
types, B!, P; photo, GH of p).
Alsophila cuspidata (Kunze) Conant, J. Arnold Arbor. pinnules, veins glabrous abaxially, or with occa-
64: 371. 1983. sional trichomes. Indusia meniscoid to subsphaer-
opteroid, pubescent with rather soft and often cris-
Scales on the abaxial side of the petiole with a pate trichomes or trichomoid processes.
dark apical seta and usually with numerous, small-
er apical and lateral setae. Lamina usually papyr- In primary forests at ca. 800-2400 m, Caja-
aceous, predominantly 2-pinnate-pinnatifid, re- marca and Amazonas south to Junin and Ucay-
duced to a pinna-like apex, pinna-rachises green- ali.

alate between the more distal sessile pinnules, veins Colombia south to northwestern Argentina.
with trichomes or stellate squamules abax-
stellate

ially. Indusia rarely cyathiform, to urceolate to Cajamarca: Prov. Hualgayoc, Soukup 3810 (F). Prov.
sphaeropteroid, glabrous or with stellate indu- Cutervo, San Andres, Lopez et al. 6684 (GH, HUT). Ama-
ment. zonas: Laguna Pomacocha, NW
of Jumbilla, Soukup
2560a (GH). Pasco: Prov. Oxapampa, Oxapampa, Leon
502 (USM). Junin: Esposto 670 (USM). Prov. Tarma, ca.
In low-elevation rain forests and less often in 3 km SE of San Ramon, Iltis & Iltis 249 (GH, USM).
cloud forests, usually at 1 50-900 m, rarely to 2200 Ucayali: Prov. Coronel Portillo, Cordillera Azul, Young
m. Amazonas and Loreto south to Puno. & Sullivan 668 (F, MO).

Nicaragua to Panama, northern South America,


south to Bolivia and Paraguay.
IV. Trichipteris
Nephelea cuspidata is the most widely distrib-
uted species of the genus. Among Peruvian species
Trichipteris Presl, Delic. prag. 1: 172. 1822. TYPE:
it is distinctive in having the scales of the petiole, = Trichipteris cor-
Trichipteris excelsa Presl
especially on the abaxial side, with several dark covadensis (Raddi) Copel. Figure 22.
apical setae and dark marginal setae.

Stem stout, erect and arborescent or


Terrestrial.
Amazonas: Rio Cenepa, ca. 5 km E of Chavez Val-
rarely decumbent, lacking spines. Leaves mono-
divia, Berlin 2066 (F, MO, uc). Loreto: Near mouth of
Rio Santiago, above Pongo de Manseriche, Mexia 6197 morphic, with scales especially on the croziers and
(F, GH, MO, NY, uc, us). Alto Rio Itaya, LI. Williams petiole base that are marginate and lack a dark,
3288 (F, GH, NY, us). Huanuco: Tingo Maria (as San apical seta. Petiole lacking spines or with corticin-
Martin), Allard 20618 (GH). Junin: E of Quimiri Bridge, ate spines. Lamina 1 -pinnate to 3-pinnate-pin-
near La Merced, Killip & Smith 23986 (F, NY, us). Ucay-
natifid, veins free. Sori round, often at the fork of
ali:Prov. Coronel Portillo, Cordillera Azul, Young &
Sullivan 705 (F, MO). Ayacucho: Estrella, between Huanta a vein, exindusiate. Spores tetrahedral-globose,
and Rio Apurimac, Killip & Smith 22654 (GH). Cuzco: trilete, porate, verrucate, or finely echinate.

120 FIELDIANA: BOTANY


FIG. 22. Trichipteris pubescens: a, apical portion of lamina; b, fertile ultimate segments; c, portion of segment,
showing a sorus and a receptacle; d, portion of petiole scale, (a from Buchtien 5304, Bolivia, F, b-c from Killip &
Smith 24871, F, d from Ollgaard et al. 9106, Ecuador, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 121


An American genus of 55 species, from Mexico References
and the West Indies south to northern Argentina
and southern Brazil. Fifteen species are in Peru, HARRINGTON, D. S. 1978. A revision of the genus
many formerly treated in the genus Alsophila be- Trichipteris. Contr. Gray Herb., 208: 3-91.
cause of the exindusiate sori. RIBA, R. 1969 (1967). Revision monografica del
The name is sometimes incorrectly spelled as complejo Alsophila Swartziana Mart. (Cyathea-
Trichopteris. The International Code of Botanical ceae). Ann. Inst. Biol. Univ. Nac. Auton. Mex-
Nomenclature, Art. 73.8 confines the use of an ico, ser. bot., 38(1): 61-100.
incorrect compounding form to epithets, thus ex-
cluding generic names. The original spelling Tri-
chipteris should be maintained.

Key to Species of Trichipteris

a. Lamina 1 -pinnate-pinnatifid b
b. Pinnae sessile to short-stalked c
c. Petiole 15-30 cm long; lamina not or not much reduced at the base 10. T. pubescens
c. Petiole 3-6 cm long; lamina much reduced at the base 11. T. phegopteroides
b. Pinnae, especially below the lamina apex, very long-stalked 4. T. latevagans
a. Lamina 2-pinnate or more complex d
d. Lamina mostly 3-pinnate-pinnatifid; pinnules abundantly short-pubescent adaxially, or petiole
scales with a narrow, dark center stripe and broad white margins e
e. Leaf ca. 1.5 m
long, pinnules with many small, bullate scales abaxially, abundantly short-
pubescent adaxially; petiole scales nearly concolorous, brown with very narrow, lighter margins
7. T. flava

e. Leaf ca. 4 m
long or more, pinnules with a few flattish scales abaxially, glabrate or with a few
long trichomes adaxially; petiole scales with a narrow dark center stripe and broad whitish
margins 9. T. serpens

d. Lamina 2-pinnate-shallowly lobed to 2-pinnate-pinnatisect; if rarely partly 3-pinnate-pinnatifid


then sparingly long-pubescent adaxially and the petiole scales with narrow, lighter margins . . . f
f. Lamina abruptly reduced at the apex g
g. Abaxial and lateral sides of the pinna-rachises lacking trichomes to (usually) sparingly

pubescent with usually brown trichomes to 1 long mm h


h. Fertile and sterile veins simple; or if rarely forked (T. procerd) then the ultimate lobes
entire and the paraphyses much shorter than the sporangia i

i. Petiole aculeate, pinnules usually sessile, basal veins end above a sinus, paraphyses
much shorter than the sporangia 1 . T. procera
i. Petiole muricate to tuberculate, pinnules stalked, basal veins connivent to a sinus,
paraphyses longer than the sporangia 13. T. lechleri
h. Fertile and sterile veins forked; ultimate lobes usually crenulate; paraphyses longer than
the sporangia 5. T. nigra

g. Abaxial and
lateral sides of the pinna-rachises usually densely pubescent with usually whitish
trichomes mostly 1.5-2 long mm 6. T. pilosissima
f. Lamina gradually reduced at the apex j

j. Pinna-rachises with few to several sharp spines 12. T. microdonta

j. Pinna-rachises lacking spines k


k. Pinnae and pinnules mostly long-stalked; ultimate segments subobtuse to acuminate or
apiculate 3. T. kalbreyeri
k. Pinnae mostly sessile to short-stalked; pinnules sessile to short-stalked or rarely long-
stalked and then the ultimate segments obtuse 1

1. Pinnules with prominent, large, flattish scales abaxially; paramo and subparamo (elfin
forest) 8. T. frigida

122 FIELDIANA: BOTANY


Pinnules glabrate or pubescent abaxially or somewhat scaly with bullate or small
montane forest and rain forest
flattish scales; m
m. Paraphyses shorter than the sporangia; petiole scales brown to dark brown, nearly
concolorous; central and basal pinnae short- to long-stalked .... 2. T. nigripes
m. Paraphyses longer than the sporangia n
n. Petiole scales light brown to dark brown, with lighter margins; tertiary (ulti-

mate) segments apically entire to coarsely toothed; central and basal pinnae
short- to long-stalked (rarely sessile) 6. T. pilosissima

n. Petiole scales dark brown to atropurpureous, usually with broad lighter mar-
gins, or whitish to light brown and concolorous; tertiary (usually ultimate)
segments apically crenulate or denticulate; central and basal pinnae sessile
o
o. Petiole with long trichomes, soon glabrous, the caducous trichomes leaving
a smooth petiole surface 14. T. conjugate
o. Petiole with long trichomes, these more or less persistent, when deciduous
leaving a hard base and a scabrous petiole surface 15. T. tryonorum

1 .
Trichipteris procera (Willd.) Tryon, Contr. Gray Petiole lacking trichomes, the scales dark brown
Herb. 200: 46. 1970. or with whitish margins. Lamina 2-pinnate-pin-
natifid, abruptly reduced at the apex, central and
basal pinnae short- to long-stalked, their pinnules
Polypodium procerum Willd., Sp. pi. ed. 4, 5: 206. sessile to short-stalked, abaxial and lateral sides
1810. TYPE: Brazil, Hoffmannsegg (holotype, B!,
of the pinna-rachises sparingly short-aculeate or
Herb. Willd. 19717).
Polypodium pungens Willd., Sp. pi. ed. 4, 5: 206. 1810, lacking spines, glabrate or with scattered to rather
asynonym of Trichipteris procera (Willd.) Tryon numerous brownish trichomes. Pinnules obtuse to
by Harrington, Contr. Gray Herb. 208: 23. 1978. acuminate, glabrate or with some small scales and
Alsophila procera (Willd.) Desv., Mem. Soc. Linn. Paris
(or) trichomes abaxially, ultimate segments obtuse
6: 319. 1827.
to subacute, fertile veins simple or rarely forked.
Alsophila dombeyi Desv., Mem. Soc. Linn. Paris 6:
320. 1827. TYPE: Peru, (Huanuco), Cochero, Paraphyses much shorter than the sporangia.
Dombey (holotype, Herb. Desv. P!; isotype, P!).
Alsophila infesta Kunze, Linnaea 9: 98. 1834. TYPE: In original rain forests or cloud forests, often in
Peru, (San Martin), Tocache, Rio Huallaga, Poep-
secondary forests or in other partly disturbed vege-
pig (holotype, LZ destroyed; isotype, B!).
Alsophila armigera Kunze, Linnaea 9: 98. 1 834. TYPE: tation, 100-1500 m, Amazonas and San Martin
Peru, (Huanuco), Ventanilla de Cassapi, Jul. 1829, south to Puno.
Poeppig (holotype, LZ destroyed; isotypes, MO!, P!; West Indies; South America south to northern
photos, GH, us of P). Peru and Bolivia.
Brazil,
Alsophila pycnocarpa Kunze, Linnaea 9: 97. 1834.
TYPE: Peru, (Huanuco), Pampayacu, Jul. 1829, Trichipteris dombeyi, although recognized by

Poeppig (holotype, LZ destroyed; isotype, Poeppig Barrington, is very close to T. procera and is here
201, B!). included within it. Trichipteris procera has an ex-
Alsophila peruviana Klotzsch, Linnaea 20: 441. 1847. tensive distribution and is quite variable as the
TYPE: Peru, (Junin), Tarma, Ruiz Herb. 66 (ho-
many synonyms described from Peru imply.
lotype, B!).

Alsophila pterorachis Baker, Syn. fil., ed. 2, 456. 1874.


TYPE: Peru, (San Martin), Tarapoto, Spruce 47 17 Amazonas: Prov. Bagua. Rio Maranon, above Cas-
(holotype, K.!). cadas de Mayasi, Wurdack 1825 (GH). San Martin: Cam-
Alsophila floribunda Baker, Syn. fil., ed. 2, 458. 1874. pana, near Tarapoto, Spruce 4323bis (GH). Prov. Rioja,
TYPE: Peru, (San Martin), Cerro Campana, Rioja, Ferreyra 18455 (USM). Loreto: Gamitanacocha,
Spruce 4715 (holotype, K; isotype, P!). Rio Mazan, J. Schunke 269 (F, GH, NY, uc). Iquitos,
Cyathea pungens (Willd.) Domin, Pterid. 263. 1929. Mexia 6497 (F, GH, MO, uc, us). Huanuco: Tingo Maria,
Cyathea willdenowiana Domin, Acta Bot. Bohemica Tryon & Tryon 5256 (F, GH). Prov. Huanuco, near con-
9: 171. 1930, nom. nov. for Polypodium procerum fluence of Rio Cayumba with Huallaga, Mexia 8293 (F,
Willd., not Cyathea procera Brause. GH, MO, NY, uc, us). Pasco: Prov. Oxapampa, Enenas,
Trichipteris infesta (Kunze) Tryon, Contr. Gray Herb. Soukup 6706 (GH). Prov. Oxapampa, Rio Palcazu, Ledn
200: 45. 1970. 702 (F). Junin: Above San Ramon, Killip & Smith 24541
Trichipteris dombeyi (Desv.) Barr., Contr. Gray Herb. (F, GH, NY, us). Ucayali: Prov.
Coronel Portillo, Dist.
208: 27. 1978. Padre Abad (as Loreto), J. Schunke 9852 (MO). Cuzco:

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 123


Prov. Paucartambo, Kosnipata-Santa Ines, Vargas 11318 Alsophila podophylla Baker, J. Bot. 19: 202. 1881.
(GH). Madre de Dios: Pantiacolla, Gentry el al. 2735 1 TYPE: Colombia, Antioquia, Kalbreyer 1375
(uc). Puno: Prov. Carabaya, Hacienda Palmares, Vargas (holotype, K; isotype, B!).
16145 (GH). Cyathea kalbreyeri (Baker) Domin, Pterid. 262. 1929.

Petiole lacking trichomes, the scales atropur-


2. Trichipteris nigripes (C. Chr.) Barr., Rhodora pureous or with lighter margins. Lamina 2-pin-
78: 4. 1976. nate-pinnatifid, rather gradually reduced at the

Alsophila nigripes C. Chr., Index fil. 45. 1905, nom. apex, pinnae and most of the pinnules long-stalked,
nov. for Alsophila melanopus Hooker (not Hassk.) pinna-rachises lacking spines, the abaxial and lat-
and with the same type. eral sides glabrous or with some short, brownish
Alsophila melanopus Hooker, Syn. fil. 37. 1866 (not trichomes. Pinnules acute to acuminate, glabrate
Hassk., 1855). TYPE: Ecuador, Chimborazo,
Spruce 5742 (holotype, K!; isotype, P!). abaxially, ultimate segments subobtuse, acute,
Cyathea nigripes (C. Chr.) Domin, Pterid. 263. 1929. acuminate or subapiculate, fertile veins simple or
forked. Paraphyses shorter than the sporangia.
Petiole lacking trichomes, the scales brown to
dark brown, nearly concolorous. Lamina 2-pin- In forests, 1200-1600 m, San Martin.

nate-pinnatifid to 2-pinnate-pinnatisect, gradually Colombia south to Bolivia.


reduced at the apex, central and basal pinnae short-
to long-stalked, their pinnules sessile to nearly San Martin: Zepelacio, near Moyobamba, Klug 3256
short-stalked, pinna-rachises lacking spines, the (F, GH). Monte Campana, near Tarapoto, Spruce 4330
(GH, NY).
abaxial and lateral sides glabrous or with a few to
sometimes many light brown to brown, rarely
whitish trichomes. Pinnules obtuse to attenuate, 4. Trichipteris latevagans (Baker) Tryon, Contr.
glabrous or with short to rather long, mostly scat- Gray Herb. 200: 45. 1970.
tered trichomes abaxially, ultimate segments ob-
tuse to acute, fertile veins simple or usually forked. Alsophila latevagans Baker, J. Bot. 19: 203. 1881.
Paraphyses shorter than the sporangia. TYPE: Colombia, Antioquia, Kalbreyer 1327
(holotype, K).
Cyathea latevagans (Baker) Domin, Pterid. 262. 1929.
In primary forests and somewhat disturbed for-
ests, wet ravines, 200-800 m, Amazonas, San Petiole lacking trichomes, the scales at the base
Martin and Loreto south to Madre de Dios. brown, nearly concolorous. Lamina 1-pinnate-
Costa Rica south to Peru.
pinnatifid, gradually reduced at the apex, pinnae
Recent collections have shown that Trichipteris very long-stalked, especially the central and lower
nigripes var. brunnescens of Colombia and Ec- ones, their apex acute. Pinnules glabrous or some-
uador, with well-developed petiole spines, pubes- what pubescent on the margins, obtuse to sub-
cent pinna-rachises, and sessile pinnules, is only acute, fertile veins forked. Paraphyses shorter than
one extreme within a spectrum of variation. to longer than the sporangia.

Amazonas: Prov. Bagua, Dist. Senepa, Tillett 672-115 In pajonal vegetation, 2200 m, San Martin.
(GH). San Martin: Prov. Mariscal Caceres, Puente Palo Colombia and Peru.
Blanco, Rio Tocache, Plowman et al. 1 1354 (GH). Loreto:
Above Pongo de Manseriche, Mexia6191 (F, GH). Huan-
The very long-stalked pinnatifid pinnae and the
uco: Hills E of Tingo Maria, Croat 21153 (MO). Ucayali: atropurpureous petiole and rachis are distinctive
Km 86 on Pucallpa-Tingo Maria road, Smith et al. 1 179 features of this species.
(GH). Cuzco: Prov. Paucartambo, Cosnipata valley,
Wachler et al. 221 (F, GH). Madre de Dios: Prov. Tam-
San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba, D.
bopata. SSW of Puerto Maldonado, Barbour 4934 (MO). Smith 4799 (GH).

3. Trichipteris kalbreyeri (Baker) Tryon, Contr. 5. Trichipteris nigra (Mart.) Tryon, Contr. Gray
Gray Herb. 200: 45. 1970. Herb. 200:46. 1970.

Alsophila kalbreyeri Baker, Summary new ferns 9. Alsophila nigra Mart.. Icon. pi. crypt. 71.1 834. TYPE:
1892, nom. nov. for Alsophila podophylla Baker Brazil, Prov. Rio Negro, Rio Japura, Martius (ho-
(not Hooker) and with the same type. lotype, not located: isotypes, B!, K).

124 FIELDIANA: BOTANY


Alsophila lasiosora Kuhn, Linnaea 36: 157. 1869. 6. Trichipteris pilosissima (Baker) Barr., Contr.
TYPE: Peru, Spruce 4349 (holotype, B; isotypes, Gray Herb. 208: 76. 1978.
GH!, P!, us!).
Alsophila tarapotensis Rosenst., Repert. Spec. Nov.
Alsophila pilosissima Baker, Syn. fil., ed. 2, 457. 1874.
Regni Veg. 7: 291. 1909. TYPE: Peru, (San Mar-
TYPE: Peru, (San Martin), Mt. Campana, Spruce
tin), Tarapoto, Spruce 4349 (holotype, P!; iso-
4322 (holotype, K!; isotype, BR; photos, GH of K,
types, GH!, us!).
GH, us of BR).
Cyathea lasiosora (Kuhn) Domin, Pterid. 262. 1929.
Cyathea primaeva Domin, Pterid. 263. 1929, nom. Cyathea pilosissima (Baker) Domin, Pterid. 262. 1929.
nov. for Alsophila nigra Mart., not Cyathea nigra
Fourn. sometimes with abundant short to mod-
Petiole
Alsophila killipiiMaxon, Amer. Fern J. 32: 58. 1942. brown to
erately long trichomes, the scales light
TYPE: Peru, Loreto, between Yurimaguas and
dark brown with lighter, sometimes nearly whit-
Balsapuerto, Killip & Smith 28133 (holotype, us!;
isotype, F!; numerous paratypes cited). ish, margins. Lamina 2-pinnate-pinnatifid to 2-

Trichipteris lasiosora (Kuhn) Tryon, Contr. Gray Herb. pinnate-pinnatisect, gradually to abruptly reduced
200: 45. 1970. at the apex, central and basal pinnae short- to long-

stalked, rarely sessile, their pinnules, especially to-

Petiole lacking trichomes, the scales brown or ward the base, sessile to usually short-stalked, pin-
dark brown with lighter margins. Lamina 2-pin- na-rachises lacking spines, the abaxial and lateral
sides with abundant long, mostly whitish tri-
nate-pinnatifid to 2-pinnate-pinnatisect, abruptly
reduced at the apex, pinnae short- to long-stalked, chomes. Pinnules obtuse to attenuate, with long,
pinna-rachises lacking spines, the abaxial and lat- mostly whitish trichomes abaxially, ultimate seg-
somewhat pubescent, some-
eral sides glabrate to ments obtuse to subacute, the apex entire to
times with rather numerous, long, mostly brown coarsely toothed, fertile veins simple or usually
to light brown trichomes. Pinnules acuminate to forked. Paraphyses longer than the sporangia.

attenuate, with usually scattered, short to mod-


and secondary forests, rarely in sea-
In primary
erately long trichomes and some small scales abax-
ultimate segments obtuse to acute, fertile sonally inundated forests, 50-800 m, Tumbes,
ially,
veins forked. Paraphyses longer than the sporan- Amazonas, and Loreto, south to Cuzco and Madre
de Dios.
gia.
Panama south to Peru.
This species is close to Trichipteris nigra, and
In wet primary forests, 100-1 100 m, Amazonas their relationship requires further study. At the
south to Madre de Dios. Dudley 13532, from
present time it is maintained as a distinct species,
Huanuco, a juvenile plant, probably of Trichip-
is
although there seems to be intergradation with T.
teris nigra; it was collected in elfin forest at 1850
nigra.
m.
Venezuela and northern Brazil, Colombia south Tumbes: Cerro Alegria, Canchaya 5169 (GH). Ama-
to Bolivia. zonas: Prov. Bagua, Quebrada Chuivi, Wurdack 1927
The few collections with long trichomes on the (GH). San Martin: Prov. Mariscal Caceres, Tocache,
abaxial and lateral sides of the pinna-rachises and Plowman & Kennedy 5790 (F, GH). Loreto: Prov. May-

nas, Mishana, Ldpez et al. 8677 (HUT). Prov. Maynas,


the abaxial side of the segments suggest that this
Rio Ampiyacu, Plowman et al. 6601 (F, GH, us). Huan-
species may not be wholly distinct from Trichip- uco: E of Tingo Maria, Croat 21153 (MO, uc). Pasco:
teris pilosissima. Prov. Oxapampa, near confluence of Rio Palcazu and
Rio Iscozacin, Smith & Franzen 1921 (F, MO). Junin:
Near La Merced, Killip & Smith 23922 (F). Ucayali:
Amazonas: Prov. Bagua, Chiriaco to Puente Venezue- Vicinity of Aguaytia (as Loreto), Croat 20952 (F, MO,
la, Barbour 4465 (MO). San Martin: Prov. Mariscal Cac- uc). Cuzco: Prov. Paucartambo, Cosnipata valley,
eres, Tocache Nuevo, J. Schunke V. 4781 (F, MO, us),
Wachterel al. 156 (F, GH). Madre de Dios: Prov. Manu,
5644 (GH). Loreto: Sierra del Pongo, Mexia 6270 (GH, Cerro de Pantiacolla, Foster et al. 10701 (F, GH).

uc). Prov. Maynas, Mishana, Rio Nanay, Gentry et al.


21109 (F, MO). Huanuco: Tingo Maria (as San Martin),
Allard 20607 (GH, us). Pasco: Prov. Oxapampa, Valle
7. Trichipteris flava Tryon, sp. nov.
del Palcazu, Ledn 703 (GH). Prov. Oxapampa, Rio Is-
cozacin, Gentry et al. 41911 (USM). Junin: Santa Rosa,
Pichis Trail, Killip & Smith 26169 (GH, NY). Ucayali: La Petiolus squamis brunneis vel atrobrunneis. Lamina
Divisoria (as Huanuco), Gentry et al. 18874 (USM). Madre ca. 1 m
longa tripinnata vel triptnnato-pinnatifida. Pin-
de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. nulae acutae supra breviter pubescentes subter squa-
10703 (F, GH). mulis multis pallide brunneis. Soris deest.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 125


Petiole lacking trichomes, the scales brown, con- Trichipteris frigida typically grows at higher al-
colorous or with very narrow lighter margins. titudes than other species of the genus.
Lamina 3-pinnate-pinnatifid nearly throughout,
gradually reduced at the apex, pinnae moderately Amazonas: Prov. Chachapoyas, Molinopampa-Dios-
an pass, Wurdack 1654 (F, GH, us). Huanuco: Cerros al
long-stalked, pinna-rachises lacking spines, the
sudoestede Monzon, Weberbauer 3389(\jstA). Huanuco-
abaxial and lateral sides with scattered, short to
Tingo Maria road, near Carpish, Gentry & Smith 44863
long trichomes. Pinnules short-stalked, acute, with (F, MO). Pasco: Prov. Oxapampa, Santa Barbara, D. Smith
many small, light brown, bullatc scales abaxially, 8172 (F, MO). San Gotardo, van der Werff et al. 8562
ultimate segments mostly acute, many falcate. Sori (MO, uc). Junin: Huacapistana, Weberbauer2272(?, USM).
absent.

HOLOTYPE Peru, Dept. Huanuco, above Chin- 9. Trichipteris serpens Tryon, sp. nov.
chao, Tingo Maria to Chinchao, 1 Aug. 1965, D.
Soejarto 1432 (GH 2 sheets). Petiolus deest. Pedum circinatum sine trichomata,

Only known from the type collection from an squamis centralibus fuscatis angustis marginatis late niv-
eis. Lamina ca. 3-4 m longa tripinnato-pinnatifida pinnis
exposed hillside at 2500 m in Huanuco.
pinnulisque longe petiolulatis. Pinnulae acuminatae sub-
Endemic to Peru. ter squamis paucis brunneis complanatis. Venae fertiles
The stem is reported on the label as slender and furcatae. Paraphyses longiores quam sporangia.
ca. 75 cm tall, the leaves ca. 1.5 m long, and the
cut stem exuding yellow sap. Although the collec- Petiole lacking trichomes, the scales with a dark,
tion is sterile, it very probably represents a species narrow central stripe and broad whitish margins
of Trichipteris. The lamina complexity and the (characters from croziers). Lamina 3-pinnate-pin-
abundant short-pubescence on the upper surface natifid nearly throughout, apex absent, pinnae long-
of the segments amply distinguish the species. to very long-stalked, pinna-rachises lacking spines,
the abaxial and lateral sides somewhat appressed
pubescent and scaly. Pinnules short- to long-
8. Trichipteris frigida (Karsten) Tryon, Contr. stalked, acuminate, glabrate or with a few large,
Gray Herb. 200:45. 1970. flattish, brown scales abaxially, fertile veins forked.

Paraphyses longer than the sporangia.


Alsophila frigida Karsten, Fl. Columb. 1 61, t. 30.
:

1859. TYPE: Colombia, (Cundinamarca), Andes TYPE Peru, Dept. Cuzco, Prov. La Conven-
of Bogota, 2600 m, Karsten (holotype, not locat-
tion, Cordillera Vilcabamba, 10 July 1968, T. R.
ed; isotype, B!).

Cyathea frigida (Karsten) Domin, Pterid. 262. 1929. Dudley 10949 (holotype, NA 2 sheets; isotype, GH
2 sheets).
Petiole lacking trichomes, the scales light to dark Known only from the type collection in elfin

brown usually with rather narrow lighter margins. forest at 2900 m in Cuzco.
Lamina 2-pinnate-pinnatifid, rarely partly 3-pin-
Endemic to Peru.
nate-pinnatifid, gradually reduced at the apex, pin-
The leaves of this new species are unusual: they
nae sessile to long-stalked, pinna-rachises lacking are 4-6 m long, or more, and trail or are scandent
spines, the abaxial and lateral sides with large flat- over low vegetation. The scales of the croziers,
tish scales or theirhard bases, with scattered brown which will in part become petiole scales, are quite
to light brown trichomes or often with dense, ca- different from those of the related Trichipteris fri-

ducous, slender, matted, whitish trichomes or gida.

strongly dissected scales. Pinnules sessile to short-


stalked, obtuse to rarely attenuate, with promi-
nent, large flattish scales abaxially, ultimate seg- 10. Trichipteris pubescens (Baker) Tryon, Contr.
ments obtuse to subacute, fertile veins forked. Pa- Gray Herb. 200: 46. 1970. Figure 22.

raphyses shorter than the sporangia or of the same


length. Alsophila pubescens Baker, Syn. fil. 449. 1868. TYPE:
Peru, (San Martin), Mt. Guayrapurima, Tara-
poto, Spruce 4712 (holotype, K; isotypes, OH!, NY!,
Growing in moist scrub forests, 2500-3500 m,
P!).
Amazonas south to Junin. Cyathea bipinnatifida (Baker) Domin, Pterid. 262.
Venezuela and Colombia south to Peru. 1929.

126 FIELDIANA: BOTANY


Cyathea pubens Domin, Pterid. 263. 1929, nom. nov. Growing at 350-1750 m, in San Martin and
for Alsophila pubescens Baker, not Cyathea pu-
(probably) Puno.
bescens Kuhn.
Endemic to Peru.
This species has a small leafless than 1 m long
Petiole lacking spines, the scales light brown
and a very short petiole ca. 3-6 cm long; the basal
with lighter margins. Lamina 1-pinnate-pinnati-
pinnae are much reduced.
fid, gradually reduced at the apex, pinnae sessile
to short-stalked, with abundant, short to moder-
San Martin: Mt. Guayrapurima, near Tarapoto, Spruce
ately long trichomes abaxially or rarely these few, 4028 (GH, NY). Prov. Riqja, Pedro Ruiz-Moyobamba,
ultimate segments obtuse, fertile veins simple or D. Smith 4452 (GH). Puno(?): Lechler (F).
forked. Paraphyses as long as the sporangia.

In montane rain forests, cloud forests, or ceja 12. Trichipteris microdonta (Desv.) Tryon, Contr.
de la montana, 850-2200 m, Amazonas south to Gray Herb. 200: 46. 1970.
Puno.
Guyana, Venezuela, and Colombia south to Bo- Polypodium microdontum Desv., Ges. Naturf. Freunde
livia. Berlin Mag. Neuesten Entdeck. Gesammten Na-
turk. 5: 319. 1811. TYPE: "America australis"
This species and the next are the only ones in
(holotype, Herb. Desv. P!; photos, GH, us).
Peru with a 1-pinnate-pinnatifid lamina. Trichip-
Alsophila microdonta (Desv.) Desv., Mem. Soc. Linn.
teris pubescens is widely distributed and, although Paris 6: 319. 1827.
variable, characterized by the features
is men- Cyathea microdonta (Desv.) Domin, Pterid. 263. 1929.

tioned in the key.


Petiole lacking trichomes, the scales light brown
to dark brown, concolorous. Lamina 2-pinnate-
Amazonas: Prov. Bagua. La Peca, Barbour 2764 (MO,
pinnatifid to 2-pinnate-pinnatisect, gradually re-
uc), 3987 (F, MO). San Martin: Prov. Rioja, Pedro Ruiz-
duced at the apex central and basal pinnae short-
Moyobamba, D. Smith 4798 (GH). Huanuco: Cerros del
Sira, Rio Llullapichis watershed, Dudley 13371, 13413, to long-stalked, their pinnules sessile to short-
13538 (GH). Pasco: Prov. Oxapampa, Palcazu, D. Smith stalked, pinna-rachises with few to several sharp
8553 (F, MO). Junin: Above San Ramon, Killip & Smith and
spines, their abaxial lateral sides glabrate or
24871 (F, us). Ucayali: Cerca a La Divisoria (as Loreto),
with usually numerous short to moderately long
Ferreyra 1071 (GH, USM). Ayacucho: Prov. La Mar, be-
tween Huanhuachayo and Punccu, Cordillera Central, trichomes. Pinnules acute to attenuate, with few
Dudley 11941 (GH). Cuzco: Prov. La Convention, Rio to usually many, short to usually long trichomes
Mapitunuari, Dudley 10159B (GH). Puno: San Gaban, abaxially, ultimate segments obtuse to acute, fer-
Lechler 2190 (K). Prov. Sandia, San Juan del Oro, Fer-
tile veins forked. Paraphyses longer than the spo-
reyra 16678, 16701 (GH, USM), 16704 (GH).
rangia or the same length.

In primary forests or disturbed forests, 100-750

11. Trichipteris m, San Martin, Loreto, and Cuzco.


phegopteroides (Hooker) Tryon,
Contr. Gray Herb. 200: 46. 1970.
Mexico and the Greater Antilles, south to Peru
and Brazil.
The sharp spines on the pinna-rachises are a
Alsophila phegopteroides Hooker, Syn. fil. 32. 1865.
TYPE: Peru, (San Martin), Tarapoto, Spruce 4020 distinctive feature of Trichipteris microdonta.
(holotype, K; isotypes, P!, us!). Rarely, they may be few and rather short, as in
Cyathea phegopteroides (Hooker) Domin, Pterid. 263. Luna 895.
1929.

San Martin: Near Tarapoto, Spruce 4726 (NY). Loreto:


Mishuyacu, near Iquitos, Killip & Smith 29876 (F, GH,
Petiole pubescent, also bearing dark brown scales
NY). Quistococha, 12 km from Iquitos, Luna 895 (F).
with lighter brown margins. Lamina 1-pinnate- Rio Itaya. Iquitos, Tryon & Tryon 5172 (F, GH). Cuzco:
pinnatifid, gradually reduced at the apex, pinnae Prov. Quispicanchi. Quince Mil, Vargas 15340 (GH).
sessile, rachis scaly and densely pubescent with
long trichomes. Pinnule-segments obtuse, pubes-
cent adaxially and abaxially, fertile veins forked. 13. Trichipteris lechleri (Mett.) Tryon, Contr.
Paraphyses shorter than the sporangia. Gray Herb. 200:45. 1970.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 127


Alsophila lechleri Men., Fil. lechl. 2: 28. 1859. TYPE: abundant long trichomes abaxially, ultimate seg-
Peru, (Puno), Tatanara, Lechler 2532 (holotype, ments obtuse to acute, the apex denticulate, fertile
Herb. Mett. B!; photo, GH).
veins forked. Paraphyses longer than the sporan-
Alsophila w/e; Christ, Hedwigia 44: 367. 1905. TYPE:
Peru, (Amazonas), Cerro Ponasa, Ule 6901 (ho- gia.

lotype, P?; isotypes, B!, L; photos, GH of B, us of


L). In montane forests, 700-2000 m, Huanuco south
Cyathea subtropica Domin, Pterid. 263. 1 929, nom. to Cuzco.
nov. for Alsophila lechleri Mett., not Cyathea lech-
leri Mett.
Venezuela and Colombia south to Bolivia.
Cyathea ulei (Christ) Domin, Acta Hot. Bohemica 9: Trichipteris conjugata and the next species, T.
168. 1930. tryonorum, are members of a group of some 1 3
species with dark denticulate margins on the pet-
Petiole lacking trichomes, the scales dark brown iole scales. The dark denticulate margins, how-
to atropurpureous with rather broad, lighter mar- ever, are often absent in the two Peruvian species.
gins. Lamina 2-pinnate-shallowly lobed to 2-pin-
nate-pinnatifid, abruptly reduced at the apex, cen- Huanuco: Prov. Leoncio Prado, Young & Sullivan 867
tral and basal pinnae long-stalked, their pinnules, (F, MO). Prov. Huanuco, Yuracyaca, Ridoutt in 1 943 (GH,
us, USM). Junin: Above San Ramon, Killip & Smith 24643
especially toward the base, long-stalked, pinna-
(F, GH, us). Ucayali: Prov. Coronel Portillo, La Divisoria,
rachises lacking spines, abaxial and lateral sides
Aguilar 848 (F, GH). Ayacucho: Prov. La Mar, between
glabrate. Pinnules acuminate to attenuate, glabrate Huanhuachayo and Punccu, Dudley 1 1943 (F, GH). Cuz-
abaxially, lobes or ultimate segments more or less co: Prov. Urubamba, near town of Machu Picchu, Rio

obtuse, fertile veins simple. Paraphyses longer than Urubamba, Tryon & Tryon 541 1 (GH, USM). Machu Pic-
chu, Iltis et al. 1024 (GH, uc).
the sporangia.

Wet forests and cloud forests, 500-1300 m,


Amazonas, Huanuco, Junin, and Puno. 15. Trichipteris tryonorum (Riba) Tryon, Contr.
Venezuela and Colombia south to Bolivia. Gray Herb. 200:46. 1970.
Trichipteris lechleri usually has the pinnules
rather shallowly pinnatifid or lobed and the basal Alsophila tryonorum Riba, Rhodora 69: 66. 1967.

veins on an ultimate segment are connivent to a TYPE: Colombia, Cundinamarca, Cuesta "Fu-
sugasuga," (Fusagasuga), Cuatrecasas 8036 (ho-
sinus.
lotype, us!).

Huanuco: Cerros del Sira, Rio Llullapichis watershed, Petiole with long trichomes, these persistent or
Dudley 13047, 13214, 13220 (GH). Junin: Prov. Satipo,
S of Chequitavo, D. Smith 5130 (F).
deciduous and then leaving a hard base and a sca-
brous petiole surface, the scales dark brown to
atropurpureous, usually with broad, lighter mar-
gins, or whitish to light brown and concolorous,
14. Trichipteris conjugata (Hooker) Tryon, Contr.
the margins slightly or not dark denticulate. Lam-
Gray Herb. 200: 45. 1970.
ina 2-pinnate-pinnatifid to 2-pinnate-pinnatisect,

fil. 37. 1 866. TYPE:


gradually reduced at the apex, pinnae sessile, pin-
Alsophila conjugata Hooker, Syn.
Ecuador, Chimborazo, Spruce 4745 as published, na-rachises lacking spines, the abaxial and lateral
5745 is correct (holotype, K; frag., us!). sides with usually abundant long, whitish to
brownish trichomes. Pinnules sessile, acute to usu-
Petiole with long trichomes, soon glabrous, the ally acuminate or attenuate, with usually abundant
caducous trichomes leaving a smooth surface, the long trichomes and often some whitish bullate
scales dark brown to atropurpureous with usually scales abaxially, ultimate segments acute to ob-
rather broad, light brown to whitish margins, or tuse, the apex crenulate to coarsely denticulate,
light brown to whitish and concolorous, the mar- fertile veins forked. Paraphyses longer than the
gins slightly or not dark-denticulate apically. Lam- sporangia.
ina 2-pinnate-pinnatifid to 2-pinnate-pinnatisect,
gradually reduced at the apex, pinnae sessile, pin- Growing at 200-2400 m, Amazonas and Pasco.
na-rachises lacking spines, the abaxial and lateral Venezuela, Colombia, south to northern Peru.
sides with usually abundant, long, whitish to This species is especially distinguished from Tri-
brownish trichomes. Pinnules sessile, acuminate chipteris conjugata by the persistent trichomes on
to mostly attenuate, with rather sparse to usually the petiole, or their hard bases that provide a sca-

128 FIELDIANA: BOTANY


brous surface when they fall off. Trichipteris con- morphic, with scales, especially on the croziers and
jugata has caducous trichomes on the petiole that the petiole base, that are marginate and lack a dark
leave a smooth surface. apical seta. Petiole lacking spines or with corticin-
ate spines. Lamina 1-4-pinnate, veins free or rare-
Amazonas: Prov. Bagua, Cordillera de Colan, SE of ly anastomosing without included veinlets. Sori
La Peca, Barbour 3750 (MO). Pasco: Prov. Oxapampa,
round, often at the fork of a vein, with a hemi-
Rio Yamaquizu valley, D. Smith 4207 (OH).
telioid to sphaeropteroid indusium. Spores tetra-

hedral-globose, trilete, nearly smooth, porate, ver-


rucate or minutely echinate.
V. Cyathea

Cyathea is a tropical American genus of ca. 40


Cyathea Sm., Mem.
Acad. Roy. Sci. Turin 5: 416.
species, with 14 of them in Peru. The genus is
1 793. TYPE: Cyathea arborea (L.) Sm.
(Poly- sometimes treated in a broad sense to include nearly
podium arboreum L.). Figure 23. allof the family Cyatheaceae. Here it is restricted
to a natural American group.
Hemitelia R. Br., Prodr. 158. TYPE:
1810. Hemitelia
multiflora (Sm.) Sprengel
= Cyathea multiflora
Sm.
Reference

Stem stout, erect and arborescent,


Terrestrial.
slender and scandent or short, lacking TRYON, R. 1976. A revision of the genus Cy-
rarely very
athea. Contr. Gray Herb., 206: 19-98.
spines. Leaves monomorphic to somewhat di-

Key to Species of Cyathea

a. Indusia hemitelioid b
b. Sori costal 3. C. vilhelmii
b. Sori medial to submarginal c
c. Basal veins straight or nearly so, ending above a sinus d
d. Ultimate segments with simple fertile veins and usually entire (rarely coarsely dentate), or
with forked fertile veins and coarsely dentate 1 C. multiflora .

d. Ultimate segments with forked fertile veins, entire or finely dentate to strongly crenate . .

2. C. andina

c. Basal veins curved, connivent to a sinus, sometimes forming costal areolae ... 4. C. petiolata

a. Indusia sphaeropteroid, sometimes evanescent and then the apical umbo or a basal remnant is usually
present e
e. Petiole scales either whitish to light brownish and concolorous or nearly so, or discordantly
bicolorous with whitish to brownish margins; petiole scurf whitish to light brown, or rarely brown
at the base of the petiole f
f. Lamina 2-pinnate-pinnatifid or 3-pinnate only at the base of the pinnae g
g. Large scales of the petiole scurf flattish h
h. Pinnae and pinnules mostly long-stalked 5. C. divergens
h. Pinnae and pinnules sessile or nearly so 7. C. ruiziana
g. Large scales of the petiole scurf mostly, or many of them, crested 6. C. pallescens
f. Lamina 3-4-pinnate nearly throughout i

i. Tertiary segments entire or basally 2-lobed; bullate scales on the abaxial side of the pinnule-
rachis dark brown 8. C. microphylla

i. Tertiary segments with up to usually 5 or more segments or lobes; bullate scales on the
brown
abaxial side of the pinnule-rachis light 9. C. multisegmenta

e. Petiole scales eitherbrown and nearly concolorous, or (sometimes narrowly) concordantly bico-
lorous with light brown to brown margins; petiole scurf brown or absent j

j. Lamina 2-pinnate-pinnatifid or 3-pinnate only at the base of central and basal pinnae; larger
ultimate segments usually ca. 1 mm long k
k. Petiole scales light brown to brown, concolorous to narrowly bicolorous; pinnules usually

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 129


with few or no scales abaxially, abundantly to moderately long- pubescent
10. C. delgadii
k. Petiole scales with a reddish brown to usually dark reddish brown or atropurpureous body,
or with dark areas or streaks, bicolorous with sometimes narrow margins 1

1. Petiole aculeate to abundantly, rarely sparsely, muricate, with abundant, sometimes


caducous scurf 11. C. caracasana
1. Petiole nearly or quite smooth, or with some scattered tubercles, scurf absent or sparse
m
m. Pinnules sessile to short-stalked 12. C. lechleri
m. Pinnules long-stalked 13. C. ebenina
Lamina 3-pinnate nearly throughout; larger ultimate segments ca. 5 mm long
14. C. dudleyi

1. Cyathea multiflora Sm., Mem. Acad. Roy. Sci. Hemitelia andina Karsten, Linnaea 28: 452. 1856.
TYPE: Colombia, "Santa Martha," 2500 m, Kar-
(Turin) 5: 416. 1793. TYPE: "Amer. merid.,"
sten (not located; authentic specimen, Karsten,
R. Shakespeare (holotype, Herb. Banks BM!;
Herb. Men. B!; photo, GH!).
photos, GH, NY, us).

Petiole slightly muricate to usually sparingly


Hemitelia multiflora (Sm.) Sprengel, Syst. veg. 4: 1 26.
aculeate, the scales light brown, or partly whitish,
1827.
to dark brown and concolorous, or usually nar-

Petiole aculeate or less often sparingly aculeate,


rowly concordantly or discordantly bicolorous,
scurf dense, often caducous, the large scales flat-
the scales light brown to dark brown, nearly con-
tish. Lamina 2-pinnate-pinnatifid to 2-pinnate-
colorous to usually, sometimes narrowly, concor-
pinnatisect, pinnules nearly sessile to rather long-
dantly bicolorous, scurf dense, often caducous, the
stalked, glabrate, slightly scaly or pubescent abax-
large scales flattish. Lamina
2-pinnate-pinnatifid
ially, fertile veins forked, rarely simple. Sori me-
to 2-pinnate-pinnatisect, pinnules sessile to short-
dial to submarginal, indusia hemitelioid.
stalked, glabrate to usually slightly to prominently
long-pubescent abaxially and usually with a few
scales, ultimate segments entire or coarsely den- In dense forests and forest borders 20-1300 m,
veins simple or forked. Sori medial to
tate, fertile Loreto south to Puno.
nearly submarginal, indusia hemitelioid. Greater Antilles, French Guiana west to Colom-
bia and south to Bolivia.
100-800 m, Amazonas south
In dense forests, Cyathea andina is close to C. multiflora, and
to Cuzco and Madre de Dios. the two species have a similar range in the Andes.
Belize south to Bolivia; northern Brazil. The indusium of C. andina is usually larger than
Although close to the next species, Cyathea mul- in C. multiflora and it commonly splits at maturity
tiflora is evidently distinct. In addition to the char- into two segments.
acters mentioned in the key, the small indusium
is entire.
Loreto: Yanamona, Rio Amazonas above mouth of
Rio Napo, Gentry et al. 36580 (MO). Pasco: Prov. Ox-
Amazonas: Laguna Pomacocha, NW
of Jumbilla, Sou- apampa, Gran Pajonal, N of Chequitavo, D. Smith 5088
kup 5260B (GH). Loreto: Gamitanacocha, Rio Mazan, J. (GH). Junin: E of Quimiri Bridge, near La Merced, Killip
Schunke 1 17 (GH, MO, NY, uc, USM). San Antonio, Rio & Smith 23995 (GH, NY, us). Prov. Tarma, Esposto 659
Itaya, Killip & Smith 29379 (F, NY, us). Junin: E of (USM). Ucayali: La Divisoria, Ferreyra 1694 (us). Prov.
Quimiri Bridge, near La Merced, Killip & Smith 23979 Coronel Portillo, Obeteni, Chrostowski 66-14 (uc). Madre
(F, NY, us). Cuzco: Cerca de Atalaya, Vargas 16274 (GH). de Dios: Prov. Tambopata, Rio Tambopata, Barbour
Entre Quince Mil y San Lorenzo, Vargas 11754 (GH). 4775 (F, MO). Puno: Prov. Carabaya, Vargas 17536 (GH).
Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster
et al. 10721 (F).

3. Cyathea vilhelmii Domin, Pterid. 264. 1929,


2. Cyathea andina (Karsten) Domin, Pterid. 263. nom. nov. for Hemitelia lechleri Mett., not
1929. Cyathea lechleri Mett.

130 FIELDIANA: BOTANY


Hemitelia lechleri Mett., Fil. lechl. 2: 28. 1859. LEC- (Diar. 1163) (holotype, LZ destroyed; authentic
TOTYPE (designated by Tryon, Contr. Gray specimen, Poeppig 2 19 (Diar. 1152), B!, P!; photo.
Herb. 206: 41. 1976): Peru, (Puno), Tatanara, GH of P).
Lechler2654 (holotype. Herb. Mett. B!; frag., GH!; Cyathea equestris Kunze, Linnaea 9: 100. 1834. TYPE:
lectoparatype, Peru, Tatanara, Lechler2650, Herb. Peru, (Huanuco), Cerro de Cristobal, Pampayacu,
Mett. B!). 1829, Poeppig (holotype. LZ destroyed; frag., K).

Petiole lacking, scales and scurf unknown. Lam- Petiole short-aculeate to aculeate, the scales very
ina 3-pinnate-pinnatifid, pinnules long-stalked, light brown to dark brown, predominantly dis-
glabrate abaxially, ultimate segments subentire to cordantly bicolorous with lighter margins, scurf
slightly toothed, fertile veins forked. Sori costal, dense, usually persistent, the large scales Hattish.
indusia hemitelioid, small, nearly concealed by the Lamina 2-pinnate-pinnatifid to 2-pinnate-pinna-

sporangia. tisect, pinnules short- to long-stalked, glabrous,


slightly pubescent or with a few scales abaxially,
Only known from the two Lechler collections ultimate segments subentire, slightly denticulate
from Puno. or shallowly crenate, fertile veins forked. Sori cos-
Endemic to Peru. tal or nearly so, or rarely medial, indusia sphaer-

opteroid.

4. Cyathea petiolata (Hooker) Tryon, Contr. Gray Montane forests and cloud forests, 200-2000 1

Herb. 206: 42. 1976. m, San Martin south to Cuzco.


The var. divergens occurs from Costa Rica and
Hemitelia petiolata Hooker, Sp. fil. 1: 31, t. 26. 1844. Panama; Guyana west to Colombia and
TYPE: Isthmus of Panama, Dr. Sinclair (hole- south to Peru.
type, K). Two varieties are recognized by Tryon (1976).
Cyathea panamensis Domin, Pterid. 264. 1929, nom.
Only var. divergens occurs in South America; the
super/1. Intended as a nom. nov. for Hemitelia
petiolata Hooker, not John Sm., 1841, but the other, var. tuerckheimii, is confined to Mexico and
latter is invalid. Guatemala. In Ecuador and Peru, var. divergens

usually has very long pinnule stalks.


Petiole nearly smooth, the scales brown to dark
brown, uniformly so or in patches, discordantly San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba,
bicolorous, with whitish to light brown margins, Smith & Vdsquez 4637 (GH). Huanuco: Pampayacu, Ka-
nehira 120 (GH). Cushi, Macbride 4819(r). Junin: Prov.
scurf rather sparse and caducous, large scales flat-
Satipo, Gran Pajonal, S of Chequitavo, D. Smith 5123
tish when present. Lamina 2-pinnate-pinnatifid, (GH). Cuzco: Sahuayaco, Chaupiorcco, Biies 840 (us).
pinnules long-stalked, glabrate abaxially, ultimate
segments toothed toward the apex, fertile veins
simple or forked. Sori submarginal, indusia hem-
itelioid. 6. Cyathea pallescens (Sodiro) Domin, Pterid. 263.
1929.
Montane forests and cloud forests, to 1750 m,
Huanuco and Cuzco. Alsophila pallescens Sodiro, Recens. crypt, vase. Quit.
20. 1883. TYPE: Ecuador, Bosques de Nanegal,
Panama south to Peru.
Sodiro (holotype, not located; authentic speci-
Unusual features of this species are the basal
men, Sodiro, P!; photo, GH).
veins that are connivent to a sinus where they
sometimes form costal areolae.
Petiole muricate to aculeate, the scales whitish
and concolorous to brown or rarely dark brown
Huanuco: Tingo Maria (as San Martin), Allard 22360
and discordantly bicolorous, with lighter margins,
(GH) is probably a juvenile plant. Cuzco: Prov. La Con-
scurf dense, mostly persistent, the large scales
vention, Cordillera de Vilcabamba, Dudley 10616 (GH).
crested or rarely absent. Lamina 2-pinnate-pin-
natifid to 2-pinnate-pinnatisect, pinnules sessile or

5. Cyathea divergens var. divergens nearly so, glabrate to pubescent and (or) scaly
abaxially, ultimate segments subentire to crenate
or rarely lobed, fertile veins simple or forked. Sori
Cyathea divergens Kunze, Linnaea 9: 1 00. 1 834. TYPE:
Peru, (Huanuco), Pampayacu, Jul. 1 829, Poeppig costal to nearly medial, indusia sphaeropteroid.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 131


On steep slopes of ceja de la montana, in cloud mate segments entire, fertile veins simple. Sori
forests and in grasslands, less often in secondary subcostal, indusia sphaeropteroid.
growth, 1 500-3300 m, Amazonas, Pasco, Ucayali,
and Cuzco. In thickets and forests, 2100-2800 m, Cuzco
Colombia south to Bolivia. and Puno.
Although this is a variable species, it is char- Endemic to Peru.
acterized by the crested large scales of the petiole The type collection, Lechler 2160, is clearly a
scurf. These are expanded into two or more planes mixture. It is cited by Mettenius under Cyathea
and usually dissected at the apex. The indusium microphylla and also under Cyathea schanschin
is rarely subsphaeropteroid or less developed. (= C. delgadii). There is a specimen at Paris of the
collection correctly identified as the latter species.
Amazonas: Prov. Bagua, Cordillera Colan, SE of La Cyathea microphylla is a rare species with small
Peca, Barbour 3590 (F, MO), 3722 (MO). Pasco: Prov. leaves to ca. 75 cm long.
Oxapampa, Rio San Alberto, Foster et al. 10285 (F, GH).
Ucayali: Prov. Coronel Portillo, Cordillera Azul, Young
Cuzco: Prov. La Convention, Dudley 10943 (GH). Puno:
& Sullivan 762 (F, MO). Cuzco: Prov. La Convention,
Valle Grande, Sandia, Vargas 11858 (GH).
antes de San Luis, Vargas 22780 (GH). Prov. La Con-
vention, Dudley 10950B, 11030 (GH), 11149 (GH, us).

9. Cyathea multisegmenta Try on, Contr. Gray


Herb. 206: 65. 1976. TYPE: Peru, Cuzco, Prov.
7. Cyathea ruiziana Klotzsch, Linnaea 20: 439. La Convention, Dudley 11326 (holotype, NA!;
1847. TYPE: "In peruviae andium nemori-
isotypes, GH!, us!).
bus," Ruiz 72 (holotype, B!; isotype, us!).
Petiole nearly smooth, the scales whitish to light
Petiole aculeate, the scales with a dark center,
brown, broad, concolorous, scurf dense, persis-
discordantly bicolorous with whitish margins, scurf tent, the large scales flattish. Lamina 3-pinnate-
dense, rather persistent, the large scales flattish. pinnatifid to 4-pinnate, pinnules sessile, pubes-
Lamina 2-pinnate-pinnatifid to 2-pinnate-pinna-
cent, and scaly abaxially, ultimate segments entire
tisect,pinnules sessile or nearly so, rather scaly to lobed, fertile veins simple or forked. Sori sub-
abaxially, ultimate segments subentire, fertile veins costal, indusia sphaeropteroid.
forked. Sori medial, indusia sphaeropteroid.

In dense cloud forest, 1 800 m, Cuzco.


Inmontane forests, ca. 2000 m, Huanuco. Endemic to Peru.
Endemic to Peru. Known only from the type collection. The leaves
Cyathea ruiziana has the large scales of the pet- are noted as 4 m or more long.
iole scurf unusually well-developed and 3-4 mm
long: they are whitish and mostly concolorous.
10. Cyathea delgadii Sternb., Vers. Fl. Vorwelt 1:
Huanuco: Huacachi, near Muna, Macbride 4135 (F, B. 1820. TYPE: Gancho
47, /. Brazil, Goias,
us). The Ruiz collection also bears the name of Pana-
General Delgado, via ad Caldas Novas, Pohl
tahuas, formerly a province west of Huanuco.
(holotype, PRC; frag., GH!).

Cyathea schanschin Mart., Icon. pi. crypt. 77, /. 54.


1834. TYPE: Brazil, Sao Paulo, Martius (holo-
8. Cyathea microphylla Melt., Fil. lechl. 1:
23, /.
type, M?; isotype, BM!; photo, GH).
3,f. 1-6. 1856. TYPE:
Peru, (Puno), St. Ga-
Cyathea oligocarpa Kunze, Linnaea 9: 101. 1834.
ban (Rio San Gaban), Lechler 2160 (LZ de- TYPE: Peru, (Huanuco), Pampayacu, Jul. 1829,
stroyed; authentic specimen, Lechler 2569, Poeppig Diar. 1101 (holotype, LZ destroyed; is-
Herb. Mett. duplicate, F! (frag.), P!; otypes, Poeppig 218, B!, MO!, P!).
B!; photo,
GH of P). Cyathea pilosa Baker, Syn. fil., ed. 2, 19. 1874. TYPE:
Peru, (San Martin), Tarapoto, Spruce 4729 (ho-
lotype, K; isotypes, BM!, GH!, P!; photos, GH, uc,
Petiole smooth, the scales whitish to light brown, us of K, GH of P).
broad, concolorous, scurf rather dense, small, more
or less persistent. Lamina 3-pinnate to 3-pinnate- Petiole slightly muricate to aculeate, the scales
pinnatifid, pinnules sessile, scaly abaxially, ulti- light brown to brown, nearly concolorous or nar-

132 FIELDIANA: BOTANY


rowly concordantly bicolorous, scurf sometimes 1 1. Cyathea caracasana (KJotzsch) Domin, Pter-
dense, usually caducous, large scales flattish. Lam- id. 262. 1929.
ina 2-pinnate-pinnatifid to 3-pinnate at the base
of the central and basal pinnae, pinnules sessile to Alsophila caracasana KJotzsch, Linnaea 1 8: 54 1 . 1 844.
short-stalked, abundantly to moderately long-pu- TYPE: Venezuela, (Dist. Federal), Caracas, Mor-
itz 117 (holotype, not located; isotypes, GH!,
bescent abaxially, sometimes with few long tri- P!).

chomes or few to several concolorous scales, ul-


Petiole muricate to aculeate, the scales predom-
timate segments entire to partly lobed, fertile veins
forked or rarely simple. Sori costal or nearly so, inantly dark reddish brown to atropurpureous,

indusia sphaeropteroid. concordantly, sometimes narrowly, bicolorous with

In montane forests, primary or open forests, or lighter margins, scurf dense, persistent or ca-
ducous, the large scales flattish. Lamina 2-pinnate-
cloud forests, 850-2880 m, Amazonas, San Mar-
pinnatifid to 2-pinnate-pinnatisect, pinnules ses-
tin, Huanuco, Pasco, and Cuzco.
sile to short-stalked or rarely long-stalked, gla-
Costa Rica and Panama; northern South Amer-
ica south to Bolivia and southeastern Brazil. brous, pubescent and (or) scaly abaxially, some-
times densely so, ultimate segments subentire to
Cyathea delgadii is characterized by the long
trichomes abaxially, which are usually abundant crenate, fertile veins forked, rarely simple. Sori
costal to medial, indusia sphaeropteroid.
and the rather few, if any, scales. It is one of the
most widely distributed species of Cyathea.
The species occurs in the Greater Antilles; Costa
Rica; and Venezuela and Colombia south to Bo-
livia.
Amazonas: Prov. Bagua, Cordillera Colan, SE of La
Peca, Barbour 3612 (F, MO, USM). San Martin: Rioja, Cyathea caracasana is the most variable species
Soukup 5223 (GH). Huanuco: Tingo Maria, Tryon & Tryon among the American Cyatheaceae. Five varieties
5219 (F, GH, us). Cerros del Sira, Rio Llullapichis wa- were recognized by Tryon (1976) in order to fa-
tershed, Dudley 13070 (GH). Pasco: Prov. Oxapampa,
cilitate the study of this complex. In addition to
Yanachaga National Park, Leon 1014 (GH). Cuzco: Prov.
La Convention, Dudley 10058, 10268 (GH). Prov. Uru- the two varieties in Peru, var. maxonii is endemic
bamba, camino a Huinayhuayna, Chavez 3442 (MO). to Costa Rica, var. caracasana is in the Greater
Antilles and northern South America, and var.
chimborazensis is in Venezuela, Colombia, and
Ecuador.

Key to Varieties

a. Pinnules usually tapering to the apex from beyond the middle, abaxially usually with abundant scales
and trichomes 1 la. var. boliviensis
a. Pinnules, especially toward the base of the central and basal pinnae, tapering to the apex from the
base, indument abaxially sparse or absent lib. var. meridensis

1 la. Cyathea caracasana var. boliviensis (Ro- Amazonas: Prov. Chachapoyas, entre Ingenio y Po-
macocha, Ldpez et al. 4312 (GH, HUT). Prov. Chacha-
senst.)Tryon, Contr. Gray Herb. 206: 77.
poyas, Molinopampa-Diosan pass, Wurdack 1653 (F,
1976. Figure 23.
GH, us). Huanuco: Within 5 km of Carpish, Tryon &
Tryon 5326 (F, GH, NY, uc, us). Pasco: Prov. Oxapampa,
Cyathea mexicana var. boliviensis Rosenst., Repert. Rio San Alberto valley. Smith & Pretel 8003 (F, MO).
Spec. Nov. Regni Veg. 25: 56. 1928. TYPE: Bo-
Oxapampa (as Junin), Soukup 2335 (F, GH). Ucayali:
livia, Hacienda Simaco, above Tipuani, Buchtien
Prov. Coronel Portillo, La Divisoria, (as Loreto), Fer-
5140 (holotype, not located; isotypes, F!, GH!, NY!,
reyra 1074 (GH, us), 7696 (us, USM). Huancavelica: Prov.
us!).
Tayacaja, entre Huachocolpa y Tintay, Tovar 4201 (GH).
Cuzco: Prov. Paucartambo, near Santa Isabel, Pilahuata
In wooded ravines, mountain forests, and es-
to Patria, Plowman & Davis 4988 (GH, USM). Prov. Uru-
pecially in cloud forests, 300-2800 m, Amazonas
1
bamba. Puente Ruinas, Machu Picchu, Iltis et al. 1025
south to Puno. (GH, uc, us). Puno: Cerca a San Juan del Oro, Valle del
Venezuela and Colombia south to Bolivia. Alto Tambopata, Ferreyra 16684 (GH, USM).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 133


X ^^ri ^^ri^T^fc^li
^^ '*^z^r
~<^s *'/':

FIG. 23. Cyathea caracasana \ar. boliviensis: a, rachis and base of pinna; b, sori (one opened); c, portion of petiole
scale. (From Wurdack 1653, F.)

134 FIELDIANA: BOTANY


lib. Cyathea caracasana var. meridensis (Kar- Petiole smooth or with a few scattered tubercles,
sten) Tryon, Contr. Gray Herb. 206: 79. 1 976. the scales reddish brown to atropurpureous, con-
cordantly bicolorous with lighter margins, scurf
Cyathea meridensis Karsten, Fl. Columb. 2: 161, t. absent. Lamina 2-pinnate-pinnatind, pinnules
184. 1869. TYPE: Venezuela, (Merida), Merida, long-stalked, glabrous to slightly pubescent abax-
2000 m, Karsten (holotype, not located; Karsten's
ially, ultimate segments subentire to shallowly
illustration is probably taken from the type col-
lection).
crenate, fertile veins simple or forked. Sori more
or less medial, indusia sphaeropteroid.

In forests, 1800-2300 m, Amazonas, south to


Junin. Growing at 2100 m, Amazonas.
Venezuela and Colombia, south to Peru.
Venezuela, Colombia, Ecuador, and Peru.
This species is unique in the genus in the char-
acters of the basal segments of the pinnules being
Amazonas: Prov. Bongara, Pomacocha a Yambras-
bamba, Ferreyra 15243 (GH). Serrania de Bagua, E of La decurrent onto the pinnule-stalk which is of a
Peca, Gentry et al. 22897 (MO). San Martin: Prov. Rioja, strongly contrasting lighter color than the dark
Pedro Ruiz-Moyobamba, D. Smith 4401 (GH). Huanuco:
pinna-rachis.
Prov. Leoncio Prado, Sullivan & Young 1178 (USM). Pas-
co: Prov. Oxapampa, Rio San Alberto valley, Smith & Amazonas: Prov. Bongara, N end of lake Pomacocha,
Pretel 7999 (F, MO). Junin: Prov. Chanchamayo, Chilpez, Hutchison & Wright 6814 (GH, NY, us).
S of San Ramon, Smith & Palacios 2641 (F, MO).

14. Cyathea dudleyi Tryon, Contr. Gray Herb.


12. Cyathea lechleri Mett., Fil. lechl. 2: 32. 1859. 206: 85. 1976. TYPE: Peru, Cuzco, Prov. La
TYPE: Peru, (Puno), St. Gaban (Rio San Ga- Convencion, Cordillera Vilcabamba, Dudley
ban) Lechler (holotype, LZ destroyed; authen- 10867B (holotype, GH!; paratypes, same lo-
tic specimen, Lechler 2309, Herb. Mett. B!). cality, Dudley 10738, 10867 GH!).

Cyathea castanea Baker, Syn. fil. ed. 2, 451. 1874. Petiole nearly smooth, the scales brown to dark
TYPE: Peru, (San Martin), Tarapoto, Spruce 47 23 brown, nearly concolorous, scurf absent. Lamina
(holotype, K; isotypes, GH!, MO!, P!, us!; photos,
3-pinnate nearly throughout, pinnules sessile to
GH, US Of P).
short-stalked, somewhat scaly and pubescent
abaxially, ultimate segments entire, fertile veins
Petiole nearly smooth, the scales reddish brown
simple. Sori subcostal, indusia sphaeropteroid.
to atropurpureous, concordantly bicolorous with

lighter margins, scurf essentially absent. Lamina


Known only from five collections, in dwarf for-
2-pinnate-pinnatind to 2-pinnate-pinnatisect, pin-
ests or wet, dense cloud forests, 2100-2700 m,
nules sessile to short-stalked, glabrate abaxially,
Pasco and Cuzco.
ultimate segments subentire to slightly crenate,
Endemic to Peru.
fertile veins forked. Sori medial, indusia sphaer-
Cyathea dudleyi has a more complex lamina
opteroid.
than the related C. lechleri and the concolorous
petiole scales distinguish it from the somewhat
In wet forests and cloud forests, ca. 600-1500
smaller C. microphylla with bicolorous scales.
m, San Martin, Huanuco, Pasco, and Puno.
Venezuela, Peru, and Bolivia.
Pasco: San Gotardo, border of Prov. Oxapampa and
Pasco, van der Werff el al. 8589 (MO, uc). Prov. Oxa-
San Martin: 36 km NE of Tarapoto, Rio Cainarache, pampa, 20 km W
of Oxapampa. D. Smith 5375 (GH).
Gentry et al. 37928 (GH, MO). Huanuco: Cerros del Sira,
Rio Llullapichis watershed, Dudley 1 3007, 13213, 13262
(GH). Pasco: Prov. Oxapampa, W
side of Cordillera de
San Matias, D. Smith 2046 (F, MO).
Comments

13. Cyathea ebenina Karsten, Linnaea 28: 461.


1856. TYPE: Venezuela (Aragua), between Two collections from Dept. Amazonas, Prov.
Caracas and Puerto Cabello, Karsten (holo- Bagua, Cordillera Colan, Barbour 3748 and 3920
type, not located; isotype, Caracas, Karsten, (MO), may be Cyathea fulva (Mart. & Gal.) Fee.
B!). The materials are not wholly adequate for certain

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 135


identification and the species is not treated under ina 1
-pinnate to 1 -pinnate-pinnatisect, veins free
Cyathea. It isknown in Ecuador and may well and the basal ones connivent to a sinus, or usually
occur in northern Peru. regularly anastomosing without included free
Specimens of Cyathea fulva will key out near C. veinlets, especially along the costa. Sori round,
delgadii, from which it differs in having few or no borne on the veins, rarely at a fork, with a hem-
scales or trichomes on the segments abaxially, or itelioid or meniscoid indusium. Spores tetrahe-
with some trichomes and also some scales. Cy- dral-globose, trilete, nearly smooth, with three large
athea delgadii has few or no scales on the abaxial equatorial pores and often smaller ones.
side of the segments, and the abaxial side is mod-
erately to usually abundantly long-pubescent. In A
tropical American genus of 25 species, with
addition, Cyathea fulva has some to many of the fiveof them in Peru.
large scales of the petiole scurf crested, while in Species of Cnemidaria were formerly placed in
C. delgadii they are flattish. Hemitelia because of the small indusia. The genus
is characterized by a lack of trichomes on the adax-
ial side of the costae and costules and by the spores

VI. Cnemidaria with three large equatorial pores. All of the Pe-
ruvian species have regularly anastomosing veins,

Cnemidaria Presl, Tent, pterid. 56. 1836. TYPE: except for C. uleana, which sometimes has free
veins.
Cnemidaria speciosa Presl. Figure 24.

Terrestrial. Stem rather small to stout, ascend- Reference


ing to erect, usually hardly arborescent, rarely to
3 mtall. Leaves monomorphic, with scales, es- STOLZE, R. 1974. A taxonomic revision of the

pecially on the croziers and petiole base, that are genus Cnemidaria (Cyatheaceae). Fieldiana,
Bot., 37: 1-98.
marginate and lack a dark apical seta. Petiole
smooth, muricate, or with corticinate spines. Lam-

Key to Species of Cnemidaria

a. Pinnae (excluding the basal pair and reduced apical ones) deeply pinnatifid or pinnatisect; the segment
sinuses extending %
or more to the costa b
b. Lamina gradually reduced to a nonconform pinnatifid apex; rachis not alate; petiole scales bi-

colorous, dark brown with broad to narrow whitish margins c


c. Basal basiscopic veins commonly arising from the costule or from its junction with the costa;
rachis and petiole muricate to spiny; scales of costae and costules, if present, deep brown . . .

l.C. horrida
c. Basal basiscopic veins, especially toward the basal and apical portions of pinnae, commonly
arisingfrom the costa; rachis smooth, petiole smooth or tuberculate; scales of costae and costules
whitish 2. C. uleana
b. Lamina abruptly reduced to aconform or subconform apical segment similar to the lateral pinnae;
rachis conspicuously alate, especially distally; petiole scales predominantly whitish
3. C. alatissima

a. Pinnae (excluding the basal pair and reduced apical ones) subentire to shallowly pinnatifid; the segment
sinuses, if present, extending less than halfway to the costa d
d. Pinnae shallowly and obtusely lobed to shallowly pinnatifid; indusia more or less semicircular,
attached on the costular side of the receptacle 4. C. speciosa
d. Pinnae subentire to broadly and coarsely serrate; indusia circular, completely surrounding the

receptacle 5. C. nervosa

136 FIELDIANA: BOTANY


FIG. 24. Cnemidaria speciosa: a. part of rachis, with pinnae; b, pinna midrib, venation pattern, and sori; c, petiole
scale, (a from Tryon & Tryon 5260, F, b-c from Buchtien 5224, Bolivia, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 137


1. Cnemidaria horrida (L.) Presl, Tent, pterid. 57. ish scales abaxially, basal basiscopic veins, espe-
1836. cially toward the basal and apical portions of the
pinnae, commonly arising from the costa. Soral
Polypodium horridum L., Sp. pi. 1092. 1753. TYPE: lines medial to supramedial between the costule
Plumier descript. pi. Amer., /. 4. 1693 (Traite and segment margin, indusia semicircular, sub-
foug. Amer., /. 8. 1705). entire to several-lobed.
Hemitelia horrida (L.) R. Br., Prodr. 158. 1810.

In cloud forests, 1700-2200 m, Cuzco.


Petiole muricate, or more often with stout spines
Southern Peru and southeastern Brazil.
to 5 mm long, the scales, usually only at the base
Cnemidaria uleana is unique in the genus in that
of the petiole, bicolorous, dark brown with narrow
anastomosing of the basal veins may be present
whitish margins. Lamina gradually reduced to a
or absent. All other Peruvian species have costal
nonconform pinnatind apex, rachis not alate, mur-
areolae. Specimens of var. uleana with anasto-
icate or often spiny. Pinnae incised % or more to
the costa, the costa and costules rarely with dull
mosing veins can be separated from Cnemidaria
horrida by the characters in the key.
brown scales abaxially, basal basiscopic veins usu-
The species ranges from Colombia to Peru, and
ally arising from the costule. Soral lines supra-
in southeastern Brazil. The collections from Peru
medial to submarginal between the costule and
are var. uleana (also in Brazil), with a gradually
segment margin, indusia more or less semicircular,
reduced pinnatifid lamina apex and usually with
subentire to erose.
costal areolae. The var. abitaguensis (Domin) Stolze
occurs in Colombia and Ecuador, with a conform
In forest, on stream banks, and mountain slopes,
or subconform apical segment and free veins.
300-1 500 m, Amazonas and Loreto, south to Cuz-
co.
Cuzco: Prov. La Convention, Cordillera Vilcabamba,
Greater Antilles; Costa Rica; Venezuela and Co-
Dudley 10459 (GH, NA), 706/7 (GH), 77J72 (GH, NA).
lombia south to Peru.
Cnemidaria horrida is the most widely distrib-
uted species of the genus and has the greatest range
of altitude. It is also the largest species with trunks Cnemidaria alatissima
3. Stolze, Fieldiana, Bot.
sometimes to 3 m tall and 7 cm in diameter. 37: 55. 1974. TYPE: Peru, Huanuco, Rio
Llullapichis watershed, Dudley 13282 (holo-
Amazonas: Prov. Bagua, Rio Maranon, above Cas-
type, GH! 2 sheets; photo, F; isotype, NA!).
cadas de Mayasi, Wurdack 1991 (us). San Martin: Near
Tarapoto, Spruce 3943 (GH, us). San Roque, LI. Wil-
liams 7156 (F, NY, us). Loreto: Pumayacu, between Bal- Petiole with minute spines, thickly covered near
sapuerto and Moyobamba, Klug 3182 (F, GH, MO, NY, the base by large, ovate, whitish scales. Lamina
us). Pasco: Prov. Oxapampa, Rio Palcazu, Smith & Sal- abruptly reduced to a subconform apical segment,
ick 8377 (MO). Cuzco: Prov. Paucartambo, Villa Carmen,
rachis with a green wing 1-2 mm
wide on each
Kosnipata-Pilcopata, Vargas 77269(GH).
side, smooth. Pinnae incised %-% to the costa, the
costa and costules lacking scales abaxially, basal

basiscopic veins usually arising from the costule.


2. Cnemidaria uleana (Samp.) Tryon var.
Soral lines about medial between the costule and
uleana.
segment margin, indusia often rudimentary, usu-
Cnemidaria uleana (Samp.) Tryon, Contr. Gray Herb. than semicircular, subentire.
ally less
200: 52. 1970.
Hemitelia uleana Samp., Bol. Mus. Nac. Rio de Ja-
neiro 1:65.1 923. TYPE: Brazil, (Rio de Janeiro), In dense cloud forests, ca. 1 500 m, Huanuco.
Perto de Nova Friburgo, Alta da Serra, Vie (ho- Endemic to Peru.
lotype, R!; photos, F, GH). This species is characterized by the abundant
white petiole scales and the broad green alate rach-
Petiolesmooth to slightly muricate, the scales is, the wings extending down onto the petiole. It
bicolorous, dark brown with broad whitish mar- is known only from the type collection and one
gins. Lamina gradually reduced to a nonconform
juvenile plant.
pinnatind apex, rachis not alate, essentially smooth.
Pinnae incised %-% to the costa, the costa and Huanuco: Rio Llullapichis watershed, Dudley 13281
costules usually with broad, pale yellowish to whit- (GH, us), a juvenile plant.

138 FIELDIANA: BOTANY


4. Cnemidaria speciosa Presl, Tent, pterid. t. 1, f. Pantiacolla, Foster10907 (F). Junin: Near La Merced,
Smith 23889 (NY, us). Chanchamayo valley, C
&
16-17. 1836. TYPE: Peru, (Huanuco), Pam- Killip
Schunke 52 (F). Ucayali: Prov. Coronel Portillo, Padre
payacu, Jul. 1829, Poeppig 221 (Diar. 1144) Abad, J. Schunke V. 5477 (F, GH, us). Aguaytia, Hua-
(holotype, PR or PRC; frag., NY!, us!; isotypes, palla 2470 (USM). Cuzco: Prov. Quispicanchi, entre In-
B!, BR, P!). Figure 24. ambari y Quince Mil, Vargas 16502 (GH). Madre de
Dies: Prov. Manu, Vargas 17738 (GH). Puno: Prov. Car-

Hemitelia subincisa Kunze, Bot. Zeit. (Berlin) 2: 296. abaya, San Gaban, Vargas 18863 (GH).

1844, nom. nov. for Cnemidaria speciosa Presl,


not Hemitelia speciosa (Willd.) Kaulf.
5. Cnemidaria nervosa (Maxon) Tryon, Contr.
Petiole smooth or rarely tuberculate at the base, Gray Herb. 200: 52. 1970.
the scales lanceolate to ovate, bicolorous, whitish
with a brown central stripe. Lamina abruptly re- Hemitelia nervosa Maxon, J. Wash. Acad. Sci. 34:

duced to a conform or subconform apical segment, 309. 1944. TYPE: Peru, Loreto, mouth of Rio
Santiago, Mexia 6291 (holotype, us! 3 sheets; is-
rachis not alate, essentially smooth. Pinnae very
otypes, F!, GH!, NY!, uc!).
deeply crenate to lobed less than '/2 to the costa,
the costa and costules rarely with dull, brown scales
Petiole smooth to muricate, the scales, mostly
abaxially, basal basiscopic veins mostly arising
near the base, lanceolate or linear-lanceolate, bi-
from the costule or its base. Soral lines supra-
colorous, brown with narrow whitish margins.
medial between the costule and segment margin,
Lamina abruptly reduced to a conform or subcon-
indusia more or less semicircular, rather large.
form apical segment, rachis not alate, smooth. Pin-
nae subentire to broadly serrate, the costa and pri-
In forests and at forest borders, along stream
mary veins lacking scales abaxially, basal basiscopic
banks, and on mountain slopes, 1 1 5-1900 m, San vein arising from a primary vein. Soral lines be-
Martin and Loreto south to Puno. tween the primary veins, indusia commonly cir-
Peru and Bolivia.
cular, completely surrounding the receptacle, en-
Three collections from Loreto, Varadero de Ma- tire to lobed.
zan, (Rio Amazonas to Rio Napo), Croat 19500
(MO, uc), 79574 (F, MO) and 20797 (MO, uc) have Rain forests, 300450 m, Amazonas and Lor-
a few broad whitish scales on the costae abaxially, eto.
rather than few small brown ones or none. These Ecuador and Peru.
may represent a distinct variety of the species. The circular indusium is a character in
the genus
shared only by Cnemidaria cocleana of Panama.
San Martin: Prov. Mariscal Caceres, Palo Blanco, To- Cnemidaria nervosa is a rare fern, represented by
cache Nuevo, J. Schunke V. 5682 (F, GH, NY). Loreto:
Prov. Maynas, Varadero, Vdsquez 721 (MO). Huanuco:
one collection from Ecuador and two from Peru.
Prov. Huanuco, Tingo Maria, Tryon & Tryon 5260 (F,
GH, u, us). Pasco: Yapas, Pichis Trail, Killip & Smith Amazonas: Prov. Bagua. valley of the Rio Maranon,
25563 (GH, NY, us). Prov. Oxapampa, Palcazu, Cerro de Wurdack 2059 (NY, us).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 139


Colombia

1. Tumbes
2. Piura
3. Lambayeque
4. Cajamarca
5. Amazonas
6. La Libertad
7. San Martin
8. Loreto
9. Ancash
10. Hulnuco
11. Lima
12. Pasco Bolivia
13. Junlh
14. Ucayali
15. lea
16. Huancavelica
17. Ayacucho
18. ApurTmac
19. Cuzco
20. Madre de Dios
21. Arequipa
22. Puno
23. Moquegua
24. Tacna Chile

DEPARTMENTS OF PERU

140 FIELDIANA: BOTANY


Index to Names

Accepted names are in roman type, synonyms are in italics, and new names are in boldface. A page
number is provided for the principal place, or the only place, where the name occurs.

Acrostichum Anemia Culcita 103


dichotomum 33 clinata 25 subg. Calochlaena 103
digit at um 33 ferruginea 27 coniifolia 103
elegans 34 var. ahenobarba 27 macrocarpa 103
nodosum 15 var. ferruginea 27 Cyathea 129
Actinostachys 33 flexuosa 27 andina 130
digit at a 33 x phyllitidis 27 arborea 129
pennula 36 var. setosa 27 aterrima 1 14
Alsophila 1 16 haenkei 29 bipinnatifida 126
armigera 123 hirsuta 28 caracasana 133
aterrima 1 14 var. humboldtiana 28 var. boliviensis 133
a usual is 1 16 hispida 30 var. caracasana 1 33

blechnoides 1 1 1 humilis 29 var. chimborazensis 133


capon sis 1 18 myriophylla 28 var. maxonii 133
ssp. polypodioides 1 1 8 oblongifolia 29 var. meridensis 135
caracasana 133 var. humilis 29 castanea 135
conjugata 128 pastinacaria 28 cuspidata 120
contracta 109 phyllitidis 29 delgadii 132
cuspidata 120 repens 30 divergens 131
dombeyi 123 tomentosa 27 var. divergens 1 3 1
elongata 1 1 5 villosa 25 var. tuerckheimii 131
engelii 1 1 6 var. anthriscifolia 27 dudleyi 135
erinacea 1 1 8 Anemirhiza 24 ebenina 135
floribunda 123 adiantifolia 24 elongata 1 1 7
frigida 126 Aspidium equestris 131
incana 120 rostratum 1 1 1 erinacea 1 1 8
infesta 123 frigida 126
kalbreyeri 124 fulva 135
killipii 125 incana 120
Balantium
lasiosora 125 kalbreyeri 124
karstenianum 105
latevagans 124 lasiosora 125
Blechnum 101
fecAter/ 128 latevagans 124
macrosora \ 1 5 Botrychium 6 lechleri 135
subg. Osmundopteris 6
melanopus 124 macrosora 1 1 5
subg. Sceptridium 6
microdonta 127 meridensis 135
cicutarium 8
mgra 124 mexicana 133
lunaria 6
nigripes 124 var. boliviensis 1 33
schaffneri 6
pallescens 131 microdonta 127
underwoodianum 6
paucifolia 1 18 microphylla 132
virginianum 8
peruviana 123 multiflora 130
var. mexicanum 8
phegopteroides 127 multisegmenta 132
virginicum 8
pilosissima 125 nigra 125
fotfa mexicanum 8
podophylla 124 /foes/a
nigripes 124
poeppigii 1 1 5 oligocarpa 132
mirifica 58
procera 123 oreites 1 1 5
pruinata 109 pallescens 131
pterorachis 123 panamensis 131
pubescens 126 Cheiroglossa 8 petiolata 131
pycnocarpa 123 palmata 9 phegopteroides 127
quadripinnata 109 Christensenia 13 /7/7o.sa 132
rostrata 1 1 1 Cnemidaria 136 pilosissima 125
sprucei 1 1 5 alatissima 138 poeppigii \ 1 5
swartziana 122 cocleana 139 polystichoides 1 1 8
tarapotensis 125 horrida 138 primaeva 125
tryonorum 128 nervosa 139 procera 123
/ 128 speciosa 139 pubens 127
Aneimia 24 uleana 138 pubescens 127
Anemia 24 var. abitaguensis 1 38 pungens 123
subg. Coptophyllum 24 var. uleana 138 quindiuensis 1 1 6
adiantifolia 24 Coptophyllum 24 rufescens 1 14
buniifolia 24 buniifolium 24 rui /i ana 132

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 141


Hemitelia
Didymoglossum nervosa 139
Cyathea krausii 87
schanschin 132 petiolata 131
membranaceum 88 4
sprucei 1 1 5 rufescens 1 1
muscoides 86
subtropica 128 speciosa 139
reptans 87
ulei 128 subincisa 139
vilhelmii 130
sphenoides 86
uleana 138
willdenowiana 123 Diplopterygium 37
Hicriopteris
bancroftii 39
Cyatheaceae 1 1 1
bancroftii 39
Cystodium 103 Homoeotes 76
Eupodium 15 heterophylla 94
kaulfussii 15 Hydroglossum 33
Danaea 15 oligostachyum 33
cuspidata 19 Hymenophyllaceae 49
elliptica 18 Hymenophyllum 50
var. crispula 18 diversifrons 9 1 sect. Buesia 59
grandifolia 17 heterophylla 94 sect. Hymenophyllum 56
humilis 20 humboldtii 94 sect. Sphaerocionium 52
jamaicensis 19 91 subg. Hymenophyllum 52
longifolia 17 subg. Leptocionium
50
moritziana 19 subg. Mecodium 50
nodosa 17 Gleichenia 37 50
subg. Sphaerocionium
oblanceolata 18 Diplopterygium 37
sect. adiantoides 66
stenophylla 19 o$n/s 42 amabile 70
trichomanoides 20 bancroftii 39 andinum 62
wendlandii 20 bifida 41 apiculatum 59
Danaeaceae 13 boliviensis 43 axillare 62
42
Davalliopsis buchtienii beyrichianum 68
elegans 84 coslaricensis 43 calodictyon 56
Dennstaedtia flexuosa 47 ciliatum 67
pearcei 99 f. monstrosa 49 contortum 64
Dicksonia 105 glauca 37 crispatulum 68
arborescens 105 hypoleuca 43 crispum 67
berteriana 105 lechleri 44 var. bipinnatisectum 67
coniifolia 103 leucocarpa 44 var.crispum 67
gigantea 105 longipinnata 45 cristatum 55
karsteniana 105 mafAwsn 41 dendritis 59
pearcei 99 mellifera 45 dependens 72
sellowiana 105 nervosa 47 dicranotrichum 50
spruceana 105 nitidula 43 ectocarpon 57
stuebelii 105 pectinata 49 elegans 66
Dicksoniaceae 101 pennigera 39 elegantulum 70
Dicranopteris 46 peruviana 45 endiviifolium 63
affinis 42 polypodioides 37 fendlerianum 64
bancroftii 39 pruinosa 42 ferax 61
bifida 41 remota 44 fragile 69
44
brittonii revoluta 42 fucoides 55
dichotoma 46 r//rffl 47 var. calodictyon 56
flexuosa 47 rubiginosa 46 var. chachapoyense 57
monstrosa 49
f. f. virescens 46 var. fucoides 56
linearis 47 simplex 39 var. pedicellatum 57
longipinnata 45 subandina 43 fusagasugense 73
nervosa 47 tomentosa 4 1 var. aberrans 73
pectinala 49 truncata 37 fusugasugense 73
pennigera 39 tuberculata 43 hirsutum 66
peruviana 45 ve/a/a 42 interruptum 72
pruinosa 42 yungensis 44 karstenianum 71
remold 44 Gleicheniaceae 37 1.n H' la in in 58
1 I

mbiginosa 46 /ar/pes 59
schomburgkiana 47 lindenii 7 1

seminuda 47 Hemiphlebium
lineare 66
kapplerianum 88
simplex 39 lobatoalatum 74
Hemitelia 129
velata 42 mathewsii 60
andina 130
yungensis 44 mexiae 59
horrida 138
Didymoglossum 76 microcarpum 68
fccAferi 131
angustifrons 86 mirificum 58
multiflora 130
hymenoides 86

FIELDIANA: BOTANY
142
Hymenophyllum Lomaridium Mertensia
molle 65 semicordatum 101 pectinata 49
multialatum 73 Lophidium 33 pennigera 39
multiflorum 63 elegans 34 pruinosa 42
myriocarpum 62 flabellum 34 remota 44
var. endiviifolium 63 latifolium 33 revoluta 42
var. myriocarpum 62 poeppigianum 36
var. nigrescens 63 Lophosoria 107 simplex 37
nigrescens 63 pruinata 109 tomentosa 4 1
nigricans 63 quadripinnata 107
nigricans 63 var. contracta 109 Metaxya 109
pedicellatum 57 var. quadripinnata 109 rostrata 1 1 1

peltatum 57 Lophosoriaceae 107 Metaxyaceae 109


peruvianum 75 Loxoma 98 Mohria 23
platylobum 68 Loxomataceae 98
plumieri 72 Loxsoma 98
plumosum 72 Loxsomopsis 99 Nephelea 118
poeppigianum 76 costaricensis 99 cuspidata 120
polyanthos 59 lehmannii 99 erinacea 1 1 8
polycarpum 64 notabilis 99 var. erinacea 1 1 8
procerum 65 pearcei 99 var.purpurascens 120
pulchellum Hooker 70 Lygodium 30 incana 120
pulchellum Mett. 65 digitatum 32 polystichoides 1 1 8
pyramidatum 74 mexicanum 33 Neuromanes
reniforme 64 micans 32 pinnatum 92
rimbachii 64 oligostachyum 33 Neurophyllum
ruizianum 69 polymorphyum 33 hostmannianum 93
rupestre 82 radiatum 32 pinnatum 92
simplex 69 scandens 30
speciosum 70 venustum 30
spectabile 70 volubile 32
sprucei 76 Odontomanes
superbum 72 hostmannianum 93
tarapotense 75 Onoclea
tenerrimum 66 Marattia 13 polypodioides 37
tomentosum 73 alata 15 Ophioglossaceae 5
tortuosum 57 var. laevis 1 5 Ophioglossum 8
trapezoidale 69 alata 15 coriaceum 12
trianae 61 kaulfussii 15 crotalophoroides 12
trichomanoides 61 laevis 15 ellipticum 12
trichophyllum 65 Marattiaceae 13 lusitanicum 12
var. buesii 66 Mecodium 50 ssp. californicum 12
var. trichophyllum 65 apiculatum 59 ssp. coriaceum 12
trifidum 76 contortum 64 ssp.lusitanicum 12
tunbridgense 50 dendritis 59 nudicaule 1 1
undulatum 63 endiviifolium 63 var. tenerum 1 1

var. fendlerianum64 fendlerianum 64 opacum 12


var. undulatum 64 ferax 6 1 palmatum 9
valvatum 68 mexiae 59 pendulum 8
verecundum 75 microcarpum 68 peruvianum 9
Hymenostachys multiflorum 63 petiolatum 9
diversifrons 9 1 myriocarpum 62 reticulatum 9
polyanthos 59 scandens 30
trichomanoides 6 1 scariosum 1 1
undulatum 64 tenerum 1 1

Lacostea Meringium vulgatum 8


tanaica 89 fucoides 56 ypanemense 1 1

Lecanium Mertensia 46 Osmunda 21


membranaceum 88 bancroftii 39 adiantifolia 24
Leptocionium 50 cicutaria 8
dicranotrichum 50 dichotoma 46 cinnamomea 21
fucoides 56 flexuosa 47 var. imbricata 21
pedicellatum 57 laevigata 37 flexuosa 27
Leptopteris 20 longipinnata 45 hirsuta 28
Lomaria mathewsii 4 1 humilis 29
euphlebia 101 nervosa 47 lunaria 6

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 143


Osmunda Sphaerocionium Trichipteris

oblongifolia 29 interruptum 72 lechleri 127


palustris 23 karstenianum 71 microdonta 127
phyllitidis 29 lindenii 1 1
nigra 124
regalis 23 lobatoalatum 74 nigripes 124
23
ssp. palustris microcarpum 68 var. brunnescens 124
var. regalis 23 phegopteroides 127
var. spectabilis 23 multialatum 73 pilosissima 125
spectabilis 23 nigricans 63 procera 123
virginiana 8 plumieri 72 pubescens 126
Osmundaceae 20 plumosum 72 serpens 126
pyramidatum 74 tryonorum 128
ruizianum 69 Trichomanes 76
Plagiogyria 101
sect.Carinatae 101 simplex 69 sect. Didymoglossum 90
costaricensis 101 spectabile 1 1 sect. Lacostea 89
denticulata 101 tenerrimum 66 sect. Microgonium 78
tomentosum 73 sect. Neurophyllum 93
euphlebia 101
latifolia 101
trichophyllum 65 subg. Achomanes 76
semicordata 101 valvatum 68 subg. Cardiomanes 78
Plagiogyriaceae 99 Sphaeropteris 1 1 2 subg. Didymoglossum 76
subg. Sclephropteris 1 14 subg. Pachychaetum 76
Polypodium
arboreum 129 subg. Sphaeropteris 1 14 subg. Trichomanes 78
dicholomum 46 atahuallpa 1 1 5 accedens 98
aterrima 1 14 angustatum 83
glaucum 109
horridum 138 bradei 115 angustifrons 86
medullare 1 1 2 elongata 1 1 5 ankersii 90
microdontum 127 hirsuta 1 14 var. tanaicum 89
horrida 112 bicorne 9 1
pedicellata 15
macrosora 1 1 4 botryoides 92
procerum 123
var. macrosora 1 1 4 brachyblastos 8 1
pruinatum 109
pungens 123 var. reginae 1 1 5 capillaceum 84
var. vaupensis 1 5 cellulosum 85
quadripinnatum 109
1

rostratum 1 1 1
medullaris 1 1 2 ciliatum 67

Ptilophyllum quindiuensis 1 16 collariatum 81


bicorne 9 1 rufescens 1 14 crinitum 94
hostmannianum 93 Sticherus 37 crispum 76
lambertianum 95 affinis42 var. haenkeanum 98
martiusii 95 bifidus 4 1 cristatum 97
pellucens 96
buchtienii 42 delicatum 98
laevigatus 37 diaphanum 83
lechleri 44 diversifrons 9 1

Ragatelus longipinnatus 45 ekmanii 88


crinitus 94 mathewsii 4 1 elatum 96
nitidulus 43 elegans 84
Schizaea 33 penniger 39 elegans 9 1

dichotoma 33 pruinosus 42 fragile69


digitata 33 revolutus 42 fucoides 56
elegans 34 rubiginosus 46 haenkeanum 98
tistulosa 34 simplex 39 heterophyllum 94
var. australis 34 tuberculatus 43 hirsutum 67
flabellum 34 velatus 42 hookeri 88
incurvata 36 yungensis 44 hostmannianum 93
pennula 36 humboldtii 94
poeppigiana 36 hymenoides 86
pusilla 34 Todea 20 hymenophylloides 83
Schizaeaceae 23 Trichipteris 120 kapplerianum 88
Selenodesmium conjugata 128 krausii 87

rigidum 85 corcovadensis 120 kunzeanum 81


Sphaerocionium 50 dombeyi 123 lambertianum 95
adiantoides 66 excelsa 120 leptophyllum 83
ciliatum 67 flava 125 lucens 95
crispum 67 frigida 126 martiusii 94
elegans 66 infest a 123 membranaceum 88
elegantulum 70 kalbreyeri 124 muscoides 76
fragile 69 lasiosora 125 opacum 84
hirsutum 67 latevagans 124 pedicellatum 90

144 FIELDIANA: BOTANY


Trichomanes Trichomanes Trichomanes
pellucens 96 pyxidiferum 82 tuerckheimii 90
pellucidum 96 radicans 8 1 undulatum 97
peltatum 57 var. kunzeanum 8 1 vandenboschii 97
pennatum 92 reptans 87 Trichopteris 122
pilosum 95 rigidum 85
pilosum 94 rupestre 82
pinnatum 92 scandens 76 Vandenboschia
plumosum 96 sellowianum 97 angustata 83
plumula 95 sphenoides 86 capillacea 84
poeppigii 89 spruceanum 91 diaphana 83
polyanthos 59 sprucei 86 hymenophylloides 83
polypodioides 89 subsessile 90 pyxidifera 82
84
prieurii tanaicum 89 radicans 8 1
punctatum 86 tenerum 83 tenera 83
ssp. sphenoides 86 trollii 9 1

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 145


281 5/93
Other Fieldiana: Botany Titles Available

Publication 1246, SI 0.50

Publication 13*

Publication 1349.
UNIVERSITY OF ILLINOIS URBANA
580 5FBN.S COOS
FIELDIANA BOTANY CHGO
20 1988

30112017564094
UNIVERSITY OF ILLINOIS-URBANA

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