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 .

ACKNOWLEDGMENTS 2 2.

KEY TO FAMILIES OF PTERIDOPHYTA IN

PERU 3 3.

1 . OPHIOGLOSSACEAE 5 4.

Botrychium 6 5.

Ophioglossum 8

2. MARATTIACEAE 13 6.

Marattia 13

Danaea 15 7.

3. OSMUNDACEAE 20 8.

Osmunda 21 9.

4. SCHIZAEACEAE 23 10.

Anemia 24

Lygodium 30 11.

Schizaea 33

5. GLEICHENIACEAE 37 12.

Gleichenia 37

Dicranopteris 46 13.

6. HYMENOPHYLLACEAE 49 14.

Hymenophyllum 50 15.

Trichomanes 76 16.

7. LOXOMATACEAE 98 17.

Loxsomopsis 99

8. PLAGIOGYRIACEAE 99 18.

Plagiogyria 101 19.

9. DlCKSONIACEAE 101

Culcita 103 20.

Dicksonia 105 21.

10. LOPHOSORIACEAE 107 22.

Lophosoria 107 23.

11. METAXYACEAE 109 24.

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

Botrychium virginianum 7

Ophioglossum: O. palmatum; O. crotal-

ophoroides; O. reticulatum 10

Marattia laevis 14

Danaea: D. moritziana; D. nodosa .... 16

Osmunda: O. regalis var. spectabilis;

O. cinnamomea 22

Anemia: A. pastinacaria; A. phylli-

tidis 26

Lygodium: L. venustum; L. volubile ... 31

Schizaea elegans 35

Gleichenia bifida 40

Dicranopteris: D. pectinata; D. flex-

uosa 48

Hymenophyllum: H. fucoides var. fu-

coides\ H. polyanthos; H. crispum 51

Trichomanes: T. radicans; T. pinnatum;

T. hymenoides 77

Loxsomopsis pearcei 100

Plagiogyria semicordata 102

Culcita coniifolia 104

Dicksonia sellowiana 106

Lophosoria quadripinnata var. quadri-

pinnata 108

Metaxya rostrata 110

Sphaeropteris: S. elongata; S. quindiu-

ensis 113

Alsophila engelii 117

Nephelea cuspidata 119

Trichipteris pubescens 121

Cyathea caracasana var. boliviensis ... 1 34

Cnemidaria speciosa 137

111

PTERIDOPHYTA OF PERU

Parti

1. Ophioglossaceae-12. Cyatheaceae

Introduction

The pteridophytes form a large and conspicuous

element of the Peruvian flora, including 96 genera

and about 1 ,000 species. Peru, which encompasses

one of the world's richest biotas, occupies a central

position in the Andes, and accordingly a knowl-

edge of its flora is basic to understanding the plant

life of the Andes as a whole. The Andes form a

largely tropical mountain chain with an essentially

north and south orientation, which is of special

significance to its biogeography and the underlying

processes of speciation.

A portion of the Ferns of Peru was published

by Rolla M. Tryon in 1964 (Contr. Gray Herb.,

vol. 194), with the encouragement and active sup-

port of Theodore K. Just of Field Museum of Nat-

ural History. A commitment was made at that

time to complete the project after undertaking a

major work on the genera of pteridophytes in

America. The present work is a cooperative proj-

ect of Harvard University Herbaria and Field Mu-

seum of Natural History to present an account of

all of the Pteridophyta of Peru. This is particularly

important in relation to the Flora of Peru, initiated

at Field Museum over 50 years ago, and thus far

encompassing some 8,500 published pages. It is

the only comprehensive treatment of Pacific coast-

al, Andean, and Amazonian plants, and is closer

to completion than any flora of the larger South

American countries.

The taxonomy is based not only on Peruvian

materials, but includes an assessment of the An-

dean species of a genus and, where appropriate,

of all tropical American species. Most of the work

will be accomplished by the authors, according to

their knowledge of particular genera, but several

treatments will be contributed by specialists cur-

rently involved in monographic studies. The re-

sults will be published in five parts, divided as

equally as practicable, beginning with the present

volume on the Ophioglossaceae through Cyathea-

ceae.

Studies are based primarily on the collections

at Field Museum, Harvard University Herbaria,

and United States National Herbarium (Smith-

sonian Institution), but specimens at many other

United States and European institutions have been

examined. The extensive collections made under

the current Flora of Peru project, a joint under-

taking of Field Museum and Missouri Botanical

Garden, are fully utilized. A close collaboration

has also been established with Universidad Na-

cional de Trujillo, Trujillo, Peru, and Museo de

Historia Natural "Javier Prado" de Universidad

Nacional Mayor de San Marcos, Lima, Peru, in-

cluding loans of their specimens. The type of each

name has been determined when possible, and an

effort has been made to see the holotype, or at

least type photographs, or authentic material.

Original drawings illustrate the diagnostic features

of each genus and, where possible, some of the

species. In addition, a number of plates published

in Fieldiana: Botany, Ferns and fern allies of Gua-

temala, have been used for species occurring in

Peru. Voucher specimens cited in the legends are

from Peru unless otherwise indicated.

Prior to 1 944 the Department of Pasco was a

part of Junin, and until recently Ucayali was a

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

labels cite towns and provinces; however, on older

specimens the labels often contain only sketchy

data, making it impossible to determine in which

part of Loreto or Junin a plant was collected. The

map of Peru at the end of the text shows the de-

partments and indicates the sequence of the col-

lection citations.

In the present work, Part I, the treatments of

the Ophioglossaceae through the Plagiogyriaceae

have been prepared by Robert G. Stolze, and those

of the Dicksoniaceae through the Cyatheaceae, by

Rolla M. Tryon. However, each author has re-

viewed and edited the manuscript of the other, to

the extent that the treatments are a joint effort.

New names published here are indicated by

boldface in the Index to Names. Collections have

been cited from each department of Peru from

which material has been seen, and further collec-

tions are sometimes cited to include other her-

baria. In general, all collections seen are cited for

rare species, and a selection is cited for common

species. Type collections, mentioned in the syn-

onymy, are not repeated in the specimen citations,

although they are included in the Peruvian range

and ecology.

The nomenclature of the genera and species is

not intended to be complete. Synonyms are listed

when they are considered useful and when the type

of the name of a species or infraspecific taxon is

from Peru.

Appended to some of the generic treatments are

portions of text labeled Comments. Herein are in-

cluded species to be expected in Peru, names based

on Peruvian material but of uncertain application,

excluded species (erroneous reports that may have

special significance), and cultivated species that

are possibly adventive.

Abbreviations of periodical publications gen-

erally follow the system of Botanico-Periodicum-

Huntianum (1968), and abbreviations of authors'

names and of books generally follow TL-2 Taxo-

nomic Literature by Stafleu and Cowan (1976 et

seq.).

Acknowledgments

The authors are indebted to Dr. Michael Dillon

(F), Dr. Abundio Sagastegui (HUT), and Blanca

Leon (USM) for their invaluable assistance in pre-

paring loans and arranging for the packing and

shipment of specimens from the two Peruvian her-

baria. The original drawings were contributed by

Field Museum scientific illustrators, Zorica Da-

bich and Clara L. Richardson, and by volunteer

illustrators, Julia A. Liesse, Rosemarie Seitz, and

Lisa M. Thorns. To these fine artists we wish to

express our sincere appreciation, not only for their

painstaking work, but also for their patience and

understanding. Special thanks are extended to sev-

eral persons who have been particularly helpful to

us, either by providing special aid with the ex-

amination of specimens at their institutions, or

through stimulating discussions or correspon-

dence concerning problems with various genera:

Dr. David Lellinger (US), Dr. Alan R. Smith (UC),

Dr. Henk van der Werff (MO), Dr. W. H. Wagner

(MICH), and Dr. Paulo Windisch (Dept. Botanico,

Rio Preto, Brazil). We also appreciate comments

on the manuscript by several reviewers.

We extend our thanks to the officers of the fol-

lowing institutions for granting loans of their ma-

terial or allowing us to examine specimens in their

herbaria: Botanischer Garten und Botanisches

Museum, Berlin-Dahlem, Berlin (B); British Mu-

seum (Natural History), London (BM); Jardin Bo-

tanique National de Belgique, Meise (BR); Royal

Botanic Garden, Edinburgh (E); Field Museum of

Natural History, Chicago (F); Harvard University,

Cambridge, Mass. — most Gray Herbarium (GH),

some Arnold Arboretum (A); Institut fur Allge-

meine Botanik und Botanischer Garten der Uni-

versitat, Hamburg (HBG); Herbarium Truxil-

lense, Universidad Nacional de Trujillo, Trujillo,

Peru (HUT); Royal Botanic Gardens, Kew, En-

gland (K); Rijksherbarium, Leiden, The Nether-

lands (L); Botanische Staatssammlung, Munich

(M); University of Michigan, Ann Arbor (MICH);

Missouri Botanical Garden, St. Louis (MO); New

York Botanical Garden, New York (NY); Museum

National d'Histoire Naturelle, Paris (P); Univer-

sity of California, Berkeley (UC); United States

National Herbarium, Smithsonian Institution,

Washington, D.C. (US); Museo de Historia Nat-

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

#BSR-85 16358 from the National Science Foun-

dation, Systematic Biology Program. The work

would not have been possible without this assis-

tance; however, any opinions and conclusions ex-

pressed are those of the authors and do not nec-

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 single sporangium 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 1 -pinnate 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 entire; indusium lacking 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

Ophioglossaceae C. Agardh, Aphor. bot. 113.1 822.

TYPE: Ophioglossum L.

Stem erect or prostrate, fleshy, lacking indu-

ment, or pubescent-scaly at apex. Leaves erect or

folded, not or scarcely circinate in vernation, en-

tire to pinnate or subpalmate or dichotomous,

commonly partially dimorphic, the sterile leaf with

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

free or anastomosing. Sporangia sessile or subses-

sile, separate or laterally joined in a synangium,

lacking an annulus. Spores trilete, lacking chlo-

rophyll, from 1,500-15,000 in each sporangium.

This is the most primitive of all living fern fam-

ilies and contains three genera and 50 to 60 species.

Botrychium and Ophioglossum occur in Peru and

are virtually cosmopolitan, with representatives in

subarctic as well as tropical regions. The Ophiog-

lossaceae are distinctive in the fleshy, mycorrhizal

roots lacking root hairs, the erect or folded (rather

than circinate) vernation, and the massive spo-

rangia with an extremely high spore capacity. Some

of the species tend to have very wide, and often

disjunct, distributions.

References

CLAUSEN, R. T. 1938. A monograph of the

Ophioglossaceae. Mem. Torrey Bot. Club, 19(2):

1-177.

UNDERWOOD, L. M., AND R. C. BENEDICT. 1909.

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

Botrychium Sw., J. Hot. (Schrader) 1800(2): 8, 110.

1802. TYPE: Botrychium lunaria (L.) Sw.

(Osmunda lunaria L.). Figure 1.

Plants terrestrial, very rarely epiphytic. Stem

erect, short, lacking indument. Leaves solitary or

few, 2-70 cm long, glabrous to pubescent, bud of

the next leaf wholly or partly enclosed by the ex-

panded stipular base of the current leaf. Sterile

lamina sessile to long-stalked, pinnatifid to de-

compound. Veins free. Fertile branch paniculate,

racemose or spicate, arising at or below the base

of the lamina. Sporangia protruding from the sur-

face of the fertile branch, not immersed in the

tissue.

The genus contains about 25 species, with dis-

tribution in temperate to tropical regions in both

hemispheres, but is poorly represented in tropical

America. Within the four subgenera of Botrychium

only four species of Sceptridium have been found

in the Neotropics, and but a single species of sub-

genus Osmundopteris (B. virginianum). The pau-

city of species in the Neotropics may be partly a

natural phenomenon, but the problem may be also

attributed to poor and sparse collections, as ferns

of this genus often go undetected. Increased col-

lecting efforts in North America have produced

more species, and similar results may be attained

through greater efforts in the tropics. Collectors

should be aware that when a single plant is found

in the field there are likely to be many more of

that and other species nearby.

References

BUTTERS, F. K. 1917. Botrychium virginianum

and its American varieties. Rhodora, 19: 207.

MILDE, J. 1869. Botrychiorum monographia.

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

Bot. Club 30: 51. 1903. TYPE: Mexico, San

Luis Potosi, Schaffner 9 (holotype, NY; iso-

type, K).

Fertile plants 12-30 cm tall (sometimes to 50

cm outside Peru), the common stalk to 4 cm long,

the leaf bud completely enclosed by the sheath of

the current leaf. Sterile lamina of fertile leaf long-

stalked (3-8 cm), fleshy, ternate, to 3-pinnate-pin-

natifid, 3-10 cm long, 6-10 cm broad. Ultimate

segments 0.3-0.7 cm long, mostly obtuse to sub-

acute, separated by usually narrow to broadly U-

shaped sinuses, the margins entire to crenate, very

rarely serrate.

Terrestrial, in high scrub forests and open mead-

ows, 2350-3600 m, Amazonas, Cuzco, Madre de

Dios(?).

Mexico; Honduras south to Argentina.

This is often difficult to distinguish from B. un-

derwoodianum (Guatemala to Venezuela), which

is usually a much larger fern with stalks of sterile

laminae up to 28 cm long. However, Mexican

plants of B. schqffneri seem to rival the latter in

size, and the more reliable diagnostic features ap-

pear to occur in the ultimate segments. Those of

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

are usually less than 0.7 cm long and entire to

crenate (only rarely serrate). The taxonomy of the

species complex requires detailed analysis.

Amazonas: Chachapoyas, along Rio Ventilla, 1-2 km

W of Molinopampa. Wurdack 1528 (us). Middle eastern

Calla Calla slopes, km 41 1-416 of Leimebamba-Balsas

road, Wurdack 1319 (us). Cuzco: Urubamba, Machu

Picchu, slopes of Altillero, Peyton & Peyton 504 (MO).

Dept. Unknown: "Pinasniocj, Panticalla Pass," Cook &

Gilbert 1845 (us) (this may be on Cerro de Pantiacolla,

Dept. Madre de Dios, near the Cuzco border).

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

(Schrader) 1800(2): 111. 1802. Figure 1.

Osmunda virginiana L., Sp. pi. 1064. 1753. TYPE:

"America," Kalm (LINN 1244.3).

Osmunda cicutaria Savigny in Lam., Encycl. 4: 650.

1797. TYPE: based on plants collected by Plu-

mier in Hispaniola, Traite foug. Amer. 136, t.

159. 1705.

Botrychium cicutarium (Savigny) Sw., Syn. fil. 171.

1806.

Botrychium virginianum var. mexicanum Grev. &

Hooker, Bot. Misc. 3: 223. 1833 (as B. virginicum

beta mexicanum). SYNTYPES: Mexico, Vera-

cruz, Chamisso s.n.; "Rigla," Veitch (both E? or

K?).

Fertile plants ca. 25-70 cm tall, the common

stalk 1 9-40 cm long, the leaf bud partly exposed

within the sheath of the current leaf. Sterile lamina

of fertile leaf sessile or short-stalked (to 5 mm),

membranous, subternate, to 3-pinnate-pinnatifid,

to 20 cm long and 25 cm broad. Ultimate segments

0.5-0.9 cm long, joined by narrow, acute sinuses,

elliptic to obovate, truncate, rounded or subacute,

their apices variously dentate or lacerate.

Forests and wooded canyons, 1800-2930 m,

Cajamarca, Amazonas, Huanuco, Pasco, Junin,

Cuzco.

Essentially cosmopolitan. In the Neotropics from

Mexico south to Bolivia and Brazil, and in the

Greater Antilles.

Botrychium virginianum has been treated by

earlier authors as several species or as one species

with several varieties or subspecies. Variants have

been distinguished primarily by relative length of

the fertile branch or degree of dissection of the

sterile lamina. However, closer examination of

more recent and numerous collections throughout

the range of the species indicates that these are

variable characters and not wholly correlated with

geography. Proctor (Ferns of Jamaica, 1985) has

observed that "... the almost complete intergra-

dation of the alleged distinguishing characters ren-

ders even varietal recognition difficult to uphold."

We concur with this judgment.

Cajamarca: Prov. Celendin, canyon of Rio Maranon

above Balsas, Hutchison & Wright 5358 (uc). Amazonas:

Prov. Chachapoyas, Cerros Calla Calla, 5 km above Lei-

mebamba, Hutchison & Wright 4885 (uc). Huanuco:

Mito, in canyon, Bryan 387 (F). Muna, in dry woods,

Bryan 537 (F). Pasco: Prov. Oxapampa, 5 km SE of

Oxapampa, D. Smith 2897 (MO). Junin: Huacapistana,

dense forest, Killip & Smith 24308 (us). Cuzco: Prov.

Calca, Vilcabamba, Vargas 3907 (us).

II. Ophioglossum

Ophioglossum L., Sp. pi. 1062. 1753. TYPE:

Ophioglossum vulgatum L. Figure 2.

Cheiroglossa Presl, Suppl. tent, pterid. 56. 1846. TYPE:

Cheiroglossa palmata (L.) Presl = Ophioglossum

palmatum L.

Plants terrestrial or epiphytic. Stem erect, glo-

bose or short-prostrate, small, lacking indument,

or filiform-scaly at apex. Leaves solitary or several,

1-75 cm long (or to more than 1.5 m in the Old

World O. pendulum), glabrous, the eroded stipular

base of the expanded leaf not enclosing the bud of

the next leaf. Sterile lamina sessile or short-stalked,

entire, or palmately or digitately lobed. Veins

anastomosing, often with free, included veinlets.

Fertile branch or branches borne on or below the

base of the sterile lamina. Sporangia joined lat-

erally in a synangial spike.

Ophioglossum is a genus of 25-30 species scat-

tered in temperate to tropic regions around the

world. Plants are often small and inconspicuous

and are commonly concealed in grassy turf or her-

baceous growth; consequently, current collection

records probably do not represent true distribution

patterns. Although six species are recognized here

from Peru, several others might be expected in the

future.

Reference

WAGNER, W. H., ET AL. 1984. Ophioglossum el-

lipticum in Louisiana and the taxonomy of O.

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. Sterile lamina 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 1 ... Sp. pi. 1063. 1 753.

TYPE: based on illustration of plant from

Haiti, Plumier, Traite foug. Amer., /. 163.

1705. Figure 2a.

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

57. 1846.

Plants epiphytic. Stem stout, elongate, abun-

dantly provided at apex with tawny to orange scales

which are hairlike for most of their length. Leaf

(15-)20-75 cm long, with common stalk (5-) 10-

40 cm long. Sterile lamina deeply and irregularly

palmately lobed, obdeltoid, narrowly cuneate at

base, lacking a distinct midvein or pale median

band of tissue. Veins distinct, the primary areoles

large, often enclosing smaller secondary areoles

and occasionally a few free veinlets. Fertile seg-

ments (l-)3-l 2, borne on or below the base of the

lamina, the fertile spikes 0.5-6 cm long, on stalks

0.2-2 cm long.

Suberect or pendent from trunks or branches of

trees, in cloud forests, elfin forests, and shaded

ravines, 1000-2450 m, Amazonas, Huanuco, Pas-

co, Cuzco.

Southern Florida; West Indies; southern Mexico

to Peru and Brazil; scattered in Old World tropics.

Amazonas: Prov. Bagua, E of La Peca, Barbour 2512,

2860, 2940 (MO). Prov. Chachapoyas, along Rio Ventilla

1-2 km W of Molinopampa, Wurdack 1478 (F, GH, NY,

uc, us). Huanuco: East of Tingo Maria (as San Martin)

in deep ravine, Allard21580 (us). Pasco: Prov. Oxapam-

pa, 1800 m, van der \Verff8352 (MO). Cuzco: Prov. La

Convention, Tunquimayo, Idma, Vargas 10681 (MICH,

MO).

2. Ophioglossum reticulatum L., Sp. pi. 1063.

1753. LECTOTYPE (designated by Proctor, Flora

Lesser Antilles 2: 43. 1977): Plumier, Traite foug.

Amer., /. 164, based on specimen from Haiti. Fig-

ure 2c-d.

Ophioglossum petiolatum Hooker, Exot. fl. 1: 56. 1823.

TYPE: based on plants from West Indies sent to

Liverpool Botanic Garden and cultivated there

(not located).

Ophioglossum peruvianum Presl, Suppl. tent, pterid.

52. 1845. TYPE: Peru, Poeppig(noi located).

Plants terrestrial. Stem cylindrical, lacking in-

dument. Leaf (5-) 10-40 cm long, common stalk

mostly epigeous, 3-20 cm long. Sterile lamina

membranaceous, 2-12 cm long, 1.5-5 cm broad,

sessile or subsessile, ovate to nearly circular, apex

obtuse to acute, base cordate to truncate or (oc-

casionally) abruptly cuneate. Veins distinct to in-

distinct, primary areoles enclosing a few smaller

secondary areoles which are scarcely differentiated

from the primary ones, free included veinlets sev-

eral to numerous. Fertile segment solitary, borne

at base of sterile lamina, on mature leaves some-

what to greatly exceeding the lamina, fertile spike

1.4-4 cm long, on a stalk 3.5-14.5 cm long.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

3 mm

3cm

5mm

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

of middle elevations, and the grassy soil of coastal

lomas, 400-3800 m, Amazonas south to Cuzco.

Widely distributed in tropical America; Old

World.

Ophioglossum reticulatum is variable in the

shape and base of the sterile lamina and in the

frequency of secondary areoles and free, included

veinlets. These characters intergrade freely with

those of the often recognized O. petiolatum, so that

it is impractical to separate the two as distinct taxa.

Thus circumscribed, O. reticulatum is perhaps the

most ubiquitous of any species in the genus.

Amazonas: Prov. Bongara, Shilla. Young & Eisenberg

460 (MO). La Libertad: Prov. Pataz, Pumatambo, Puerta

del Monte. Lope: & Sagdstegui 3438 (GH). San Martin:

Monte Campana, near Tarapoto, Spruce 4709 (p, us).

Lima: Lomas de Quilmana, Coronado 27 (us). Prov.

Chancay, Lomas de Lachay, Coronado & Velarde 15 (uc,

us); Tryon & Tryon 5419 (GH). Pasco: Prov. Oxapampa,

Chontabamba, D. Smith & Brack 3056 (F, MO). Ayac-

ucho: Aina, between Huanta and Rio Apurimac, Killip

& Smith 22809 (GH, NY, us). Apurimac: Prov. Abancay,

Cachora, Huillcayoc, Vargas 9096 (MICH, uc). Cuzco:

Prov. Urubamba, Machu Picchu, Leon 452 (GH, USM);

Tryon & Tryon 5400 (GH, us), 5419 (GH). Madre de Dios:

Prov. Manu, Parque Nacional Manu, Foster et al. 11486

(F).

and sedges, 2800-4000 m, Cajamarca, Lima, Cuz-

co.

Colombia; Peru; Brazil.

In spite of Clausen's work (1938), considerable

study is still needed to understand the variation

within O. nudicaule. The Peruvian plants here

identified cannot be clearly placed within any of

the varieties delineated by Clausen; but, aside from

the fact that their fertile segments do not rise too

conspicuously beyond the sterile lamina, they seem

to fit best under var. tenerum. Diagnostic features

of O. nudicaule are so few and variable that it is

often difficult to discern which are innate differ-

ences and which are merely induced by rigorous

habitat or attributed to degree of maturity of the

individual plant. It is likely that this and other

varieties accepted by Clausen are not supportable,

as evidenced by the studies of Wagner et al. ( 1 984),

which suggest that they "all should be subsumed

under the single binomial, O. nudicaule."

Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui

et al. 6412 (GH, HUT). Lima: Prov. Huarochiri, Dist. de

Mariatana, Cueva Mortero, Cerrateet al. 4812 (GH, USM).

Cuzco: Prov. Acomayo, Quenco Grande, near Acomayo,

Vargas 9752 (F, GH, K, ucp.p., us). Prov. Espinar, Yauri,

Vargas 5619 (MICH). Prov. Urubamba, Vargas 14122

(GH).

3. Ophioglossum nudicaule L. f. var. tenerum

(Mett.) Clausen, Mem. Torrey Bot. Club 19(2):

151. 1938.

Ophioglossum ypanemense Mart., Icon. pi. crypt. 39,

1 30, /. 73. 1 834. TYPE: based on specimens from

"Ypanema," Brazil.

Ophioglossum tenerum Mett. in Prantl, Ber. Deutsch.

Bot. Ges. 1: 352. 1883. TYPE: United States,

Georgia, collector undesignated (holotype, B).

Plants terrestrial. Stem cylindrical to somewhat

swollen, lacking indument. Leaf 2.5-7 cm long,

common stalk mostly epigeous, 0-4 cm long. Ster-

ile lamina chartaceous to somewhat carnose, 0.8-

2.5 cm long, 0.3-1 .0 cm broad, 1.5-3 times as long

as broad, sessile, elliptical to subovate, obtuse or

subacute, base cuneate. Veins indistinct to ob-

scure, areoles small, not enclosing secondary ar-

eoles or (except rarely) free veinlets. Fertile seg-

ment solitary, borne at base of sterile lamina, on

mature leaves somewhat exceeding the lamina,

fertile spike 0.5-1 .2 cm long, on a stalk 0.8-2. 5(-4)

cm long.

In high rocky meadows and slopes, with grasses

4. Ophioglossum scariosum Clausen, Mem. Tor-

rey Bot. Club 19(2): 153. 1938. TYPE: Peru,

Junin, vicinity of Oroya, Kalenborn 125 (ho-

lotype, us!; isotypes, GH!, MO!, NY).

Plants terrestrial. Stem broadly elongate, 4-6

mm thick, lacking indument. Leaf 3-6 cm long,

with common stalk and bases of fertile and sterile

stalks hypogeous and scarious, common stalk 0.3-

0.8 cm long. Sterile lamina 1-1.5 cm broad, ob-

long-ovate to suborbicular, apex apiculate, at base

truncate, reduced abruptly to an attenuate stalk

0.8-1.2 cm long. Veins indistinct to nearly ob-

scure, primary areoles usually enclosing some sec-

ondary ones as well as free veinlets. Fertile seg-

ment solitary, borne at the base of the stalk of the

sterile lamina, on mature leaves somewhat to

greatly exceeding the lamina, fertile spike 0.6-1.2

cm long, on a stalk 1.3-4 cm long.

In open, grassy areas, 2840-3850 m, Junin, Cuz-

co, Puno.

Peru and Bolivia.

Leaves of this rare fern grow with proximal por-

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-

mon stalk and proximal portions of the stalks of

fertile and sterile segments contrast sharply with

the dark color of the rest of the leaf.

Cuzco: Prov. Urubamba, Vargas 9165 (MICH). Puno:

Lake Titicaca, Bishop 1994 (us).

5. Ophioglossum crotalophoroides Walter, Flora

Caroliniana 256. 1788. TYPE: Carolina,

Walter (holotype, BM?). Figure 2b.

Plants terrestrial. Stem globose, 4-12 mm in

diameter, lacking indument. Leaf 3-15 cm long,

with common stalk 0.5-4 cm long. Sterile lamina

ovate to deltoid, acute or rarely obtuse, cordate or

truncate and abruptly attenuate at base, often lon-

gitudinally folded. Veins distinct or indistinct, pri-

mary areoles not or rarely enclosing secondary ones

or free veinlets. Fertile segment solitary, borne at

the base of the sterile lamina; in mature leaves the

fertile segments somewhat to greatly exceeding the

lamina, fertile spike 0.5-1.5 cm long, on a stalk

to 10 cm long.

On moist, open, grassy slopes, 600-3900 m,

Huanuco, Lima, Junin, Apurimac, Cuzco.

Southern United States; Mexico to Honduras;

Colombia and Venezuela south to Argentina and

Chile.

Among the diminutive species of Ophioglos-

sum, a few plants are found with stems which are

somewhat swollen. However, only in O. crotalo-

phoroides is the stem so conspicuously globose.

With this should probably be included O. opacum

Carmichael, from Tristan da Cunha. It is said to

differ from the former merely in the somewhat

shorter fertile segment and slightly smaller globose

stem.

Huanuco: Prov. Huanuco, Huanuco-Tingo Maria road,

from Bandera Blanca to Carpish Tunnel, Luteyn & Lu-

teyn 5464 (NY). Lima: Prov. Lima, Lomas de Lurin,

Ferreyra 9566 p.p. (USM). Junin: Huaytapata. near Ha-

cienda Conocancha, Tiller 14 (USM). Apurimac: Prov.

Abancay, Cachora, Vargas 9054 (MICH p.p., MO). Cuzco:

Prov. Calca, Vilcabamba, Vargas 3978 (us). Prov. Can-

as, slopes of San Andres de Checca, Vargas 11010 (F,

uc).

6. Ophioglossum lusitanicum L. ssp. coriaceum (A.

Cunn.) Clausen, Mem. Torrey Bot. Club 19(2):

161. 1938.

Ophioglossum coriaceum A. Cunn., Companion Bot.

Mag. 2: 361. 1836. TYPE: based on specimens

from Matauri, New Zealand (not located).

Plants terrestrial. Stem cylindrical to somewhat

swollen, lacking indument. Leaf 4-8(-9) cm long,

common stalk mostly epigeous, 0.5-2.5 cm long.

Sterile lamina chartaceous to somewhat carnose,

1.5—4 cm long, 0.3-0.6 cm broad (ca. 4.5-8 times

as long as broad), sessile, linear-lanceolate to nar-

row-oblanceolate, apex acute or subacute, base

narrowly cuneate to (typically) long-attenuate.

Veins indistinct to obscure, areoles commonly long

and narrow, not enclosing secondary areoles or

free veinlets. Fertile segment solitary, borne at base

of sterile lamina, on mature leaves somewhat to

conspicuously exceeding the lamina, fertile spike

0.4-1.7 cm long, on a stalk 1-7 cm long.

In soil pockets or on thin soil over rocks, in Peru

possibly confined to the coastal lomas in the vi-

cinity of Lima, 400-650 m.

Peru; Bolivia; Chile; New Caledonia; Australia;

New Zealand.

In Peru, this is typically a very delicate plant,

with long, narrow, sterile leaves and a fragile, fil-

iform, fertile stalk, and has thus far been found

only at low elevations. However, we have exam-

ined some Bolivian plants of this subspecies, from

elevations to 3000 m, which are more variable:

sometimes more robust, with stouter fertile stalks,

and with sterile laminae broader and less attenuate

to base and apex. Subspecies lusitanicum, from

Europe, Asia, and Africa, has obtuse sterile lam-

inae with larger and fewer areoles. Subspecies cal-

ifornicum (Prantl) Clausen (California & Mexico)

has laminae larger and broader, with less attenuate

bases than in ssp. coriaceum and, according to

Clausen (1938), perhaps does not merit distinc-

tion. As with the O. nudicaule complex (q.v.), far

more study is needed to clarify the taxonomy, at

both inter- and intraspecific levels.

Lima: Prov. Chancay, Lomas de Lachay, Coronado &

Velarde 16 (GH, uc, us); Tryon & Tryon 5414 (GH p.p.,

us). Lomas de Quilmana, Coronado 27 (uc, us p.p.).

Lomas de Lurin. Ferreyra 9566 (GH, USM p.p.). Lomas

de Atocongo, Grant 7509 (GH).

Comments

Ophioglossum ellipticum Hooker & Grev., Icon,

fil. 1, t. 40A. 1831. TYPE: illustration, based

12

FIELDIANA: BOTANY

on specimens collected in Demerara, "British

Guiana," Parker.

This perhaps may be expected in Peru, as its

range is thus far reported from Guatemala to Costa

Rica, and the Guianas southward to Brazil and

(possibly) Bolivia. Although usually a smaller plant

than Ophioglossum reticulatum, the aspect of the

sterile lamina is similar in that the primary areoles

contain a number of secondary areoles and usually

some free veinlets. However, the secondary ar-

eoles are much more numerous in O. ellipticum,

and the walls are thinner and far less distinct than

those of the primary ones. Additionally, laminae

in O. ellipticum have a somewhat distinct midvein

and a pale, median band of tissue which persists

to within a short distance of the apex. It is most

closely related to O. nudicaule, in fact, the plants

heretofore recognized as O. ellipticum may be

merely O. nudicaule with larger and more fully

developed leaves (see Wagner et al., 1984).

Family 2: MARATTIACEAE

Marattiaceae Bercht. & J. S. Presl, Prir. rostlin 1 :

272. 1820. TYPE: Marattia Sw.

Danaeaceae Agardh, Aphor. hot. 117. 1822. TYPE:

Danaea Sm.

Stem fleshy, stout to massive, creeping or erect,

sometimes scaly. Leaves moderate in size to huge

(3 m or more), circinate in vernation, 1 -pinnate

to pinnately compound or rarely simple or pal-

mate, monomorphic to somewhat dimorphic. Pet-

iole with an expanded, stipular base, there swollen

and sometimes subarticulate, frequently also with

several swollen and darkened nodes throughout

its length. Pinnae commonly borne in opposite

pairs from swollen, darkened nodes on the rachis.

Veins free or (in Christensenia) reticulate. Spo-

rangia borne on the abaxial surface of leaves, with

thickened walls, lacking an annulus, separate and

contiguous or fused into indurated synangia and

opening by terminal pores. Spores ellipsoidal and

monolete or trilete, rugose or echinate, very nu-

merous (1,500-1,700) in each sporangium.

Of the seven genera (about 1 50 species) in Mar-

attiaceae, two are represented in Peru: Danaea,

confined to the Neotropics, and Marattia, pan-

tropical. Next to Ophioglossaceae, it is considered

to be the most primitive of the living fern families

and can be easily recognized by the character of

the sporangia being fused into thick, elongate, dou-

ble-rowed synangia.

References

DEVRIESE, W. H.,ANDP. HARTING. 1853. Mon-

ographic des Marattiacees, Leiden, Nether-

lands.

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

attiaceae, pp. 39-50, in Ferns and allied plants,

Springer- Verlag, New York.

UNDERWOOD, L. M. 1909. Marattiaceae, in N.

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

Marattia Sw., Prodr. 128. 1788. TYPE: M. alata

Sw. Figure 3.

Plants terrestrial. Leaves coarse, to ca. 3(-4) m

long, 2-4-pinnate, often ternate in architecture,

monomorphic. Lamina with expanded tissue es-

sentially glabrous, but in some species sparsely to

moderately scaly or with a few trichomes on the

axes abaxially. Penultimate divisions with axes

commonly broadly alate, the wings narrowing

abruptly at the base of each segment. Veins free.

Sporangia in two rows, opening by longitudinal

slits, fused into bivalvate, ovoid synangia, which

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.

Spores ellipsoid, monolete.

This is a pantropical genus of about 40 species,

commonly occurring in dense, wet forests. It is

readily separated from other Neotropical genera

by the large, decompound, subternate leaves with

stipular bases, and the distinctive ovoid synangia.

A single species is known in Peru.

1. Marat tia laevis Sm., Plantarum Icones Inedi-

tae ... 2: /. 47. 1790. TYPE: as "Polypodium

pedicellata, in argento pedunculat(o?), He

Dominique (Dominican Republic, not Do-

minica), Thiery" (holotype, LINN). Figure 3.

Marattia alata sensu Raddi, PI. bras. nov. gen. 1 : 74,

t. 83-84. 1825.

Marattia Kaulfussii Hooker, Gen. fil., t. 26 (as to note).

1839. TYPE: Brazil, 1816, Cunningham s.n.

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

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

/. 95. 1860.

Marattia alata Sw. var. laevis (Sm.) Farw., Amer. Midi.

Naturalist 12: 308. 1931.

Leaves 2—4 m long, commonly 3-pinnate-pin-

natifid, at least as to base of lamina. Axes (at least

minor ones) sparsely to moderately provided on

abaxial side with filiform to lanceolate scales, these

orange to light brown, flaccid, 0.5-3 mm long,

often caducous. Costae of penultimate segments

bearing on the adaxial side 1 -several well-spaced,

often appressed, awns, these 0.2-1 mm long (or

on older leaves sometimes withering or broken

away near their bases). Synangia 8-16(-18)-lo-

cular, mature ones obviously stipitate, the stalks

stout, to 0.7 mm long. Spores echinate.

Occurring in dense forests, principally along the

Cordilleras Central and Oriental, in Amazonas,

Huanuco, Pasco, Junin, and Cuzco, 1400-2400 m.

Costa Rica; Cuba; Hispaniola; Colombia; Ven-

ezuela; Brazil; Ecuador; Peru; Bolivia.

In various herbaria, M. laevis has been confused

with M. alata, a species from the Greater Antilles.

Although leaf outline is similar in each, M. laevis

is easily distinguished by its stalked synangia and

awned penultimate axes, these characters are

unique in the genus, at least in New World species.

There has been some confusion as to the identity

of M. laevis, beginning with Smith's original de-

scription, which mentioned nothing about the dis-

tinctive stalked synangia and adaxial awns. Kaul-

fuss (Enum. fil., p. 32, 1824) mentioned seeing

some Brazilian specimens of M. laevis with "soros

... stipitati," upon which character Hooker (fol-

lowing Smith's unpublished description) partially

based the new species M. Kaulfussii (1839). Ob-

viously Hooker had seen the type of M. laevis, and

supposed it lacked the stalked synangia; therefore,

he surmised that the Brazilian collections of Kaul-

fuss represented a new species. Finally in 1936

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

matter, this reply was received from Miss Jose-

phine M. Camus: "I have examined the (type)

specimen in the Linnaean Herbarium. It does in-

deed have synangia with very elongate receptacles.

The original label bears the name Polypodium ped-

icellata in Linnaeus fil.'s hand. The 'awns' are pres-

ent on the upper surface of the pinnule axes... ."

Thus, there seems to be no question that the fern

considered to be M. Kaulfussii for nearly 1 50 years

is M. laevis.

Amazonas: Prov. Bagua, Cordillera Colon SE of La

Peca, Barbour 3900 (MO). Huanuco: Churubamba, Trail

Puente Durand to Exito, Mt. Santo Toribio, Mexia 8250

(F, GH, MICH, NY, uc, us). Pasco: Cushi, Macbride 4817

(F, GH, us). Junin: Schunke Hacienda above San Ramon,

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

Killip & Smith 25738 (GH, NY, us). Chanchamayo Valley,

C. Schunke 671 (F). Cuzco: La Convention, above Knox's

Cascade, Dudley 10619 (GH, MO).

II. Danaea

Danaea Sm., Mem. Acad. Roy. Sci. (Turin) 5: 420.

1793, nom. consent., not Allioni, 1785. TYPE:

Danaea nodosa (L.) Sm. (Acrostichum nodos-

um L.). Figure 4.

Plants terrestrial, with stems creeping to decum-

bent or erect. Leaves to 2 m long, 1 -pinnate (or

simple in 2 species outside Peru), monomorphic

to somewhat dimorphic (fertile ones commonly

longer and narrower and with smaller pinnae). Pet-

iole (in most species) nodose, scaly, bearing sti-

pules at the base. Rachis nodose, nonalate to con-

spicuously alate at least distally, glabrous or more

often with appressed scales. Lamina with a distinct

apical segment, or this replaced by a proliferous

bud, provided with scales abaxially, those of the

laminar tissue minute and scattered. Sterile pinnae

opposite or subopposite, straight to subfalcate, ses-

sile or short-stalked, entire and often undulate, or

serrate near the apex. Veins free. Fertile pinnae

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.

Sporangia nearly covering the abaxial surface, co-

alescing into much elongated, indurated synangia

and opening by terminal pores. Spores monolete,

the surface prominently echinate.

Danaea is confined to the Neotropics. With its

nodose leaf axes and the distinctive synangia near-

ly covering the abaxial surface of simple, usually

opposite, pinnae, it should not be confused with

other Peruvian genera. However, distinguishing

species is another matter. To date there has been

no comprehensive study of the genus, and many

of the characters used by Underwood to separate

taxa are highly suspect: e.g., length of fertile pin-

nae, number of veins per centimeter, veins paired

in origin versus distinctly forked. Species previ-

ously thought of as distinct in Central or South

America may be synonymous with some in the

West Indies. Although about 35 species have been

recognized, 20-25 is a more realistic total to expect

once a greatly needed revision has been under-

taken. Six species from Peru are recognized in the

following treatment.

References

PROCTOR, G. R. 1977. Danaea, pp. 45-49, in

Howard, R. A., ed., Flora of the Lesser Antilles:

Leeward and Windward Islands, Vol. 2, Pteri-

dophyta, Arnold Arboretum, Harvard Univer-

sity, Jamaica Plain, Mass.

UNDERWOOD, L. M. 1 902. A review of the genus

Danaea. Bull. Torrey Hot. Club, 29: 669-679.

Key to Species of Danaea

a. Larger sterile pinnae (5-)6-40 cm long; larger fertile pinnae 4-25 cm long 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. Sterile pinnae 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 sterile pinnae 1.5-3.5(-4) cm long; larger fertile pinnae 1.1-1.7 cm long 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. Sterile pinnae 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-

ly scaly; pinnae 7-16 pairs, subsessile to short-

stalked, elliptic to oblanceolate, larger ones 20-40

cm long and (2.5-)3-6.5 cm broad, base cuneate,

apex long-acuminate, margins entire (very rarely

serrulate along the acuminate tip), abaxial surface

and veins sparsely provided with minute (ca. 0. 1

mm) scales, these appressed, nearly circular, dark

or reddish brown, commonly erose to stellate; veins

mostly paired or 1 -forked near the base. Fertile

pinnae 7-14 pairs, 12-25 cm long, 1.5-2.5 cm

broad, subsessile to short-stalked, the apex acute

to acuminate.

Terrestrial, in dense forests, 100-2100 m, Ama-

zonas and Loreto to Cuzco and Madre de Dios.

Southern Mexico to Panama; West Indies; Co-

lombia; Venezuela; Surinam; Ecuador; Peru; Bra-

zil.

Danaea grandifolia is said to differ from D. no-

dosa in the broader sterile pinnae and more broad-

ly cuneate bases of fertile pinnae. The latter char-

acter is not consistent, and the former is hardly

sufficient reason for separation even at infraspe-

cific level. There is also some question as to the

merit of separating D. nodosa and D. elliptica.

While the characters of nodose or non-nodose pet-

ioles and pinnae number usually serve to distin-

guish the two, even these have not been reliable

consistently in specimens seen from Peru. Further

study is needed.

Ama/onas: E of Huampami, near Pedro Alberto, Ber-

lin 1524 (MO). San Martin: Prov. Rioja, km 390, Pedro

Ruiz-Moyobamba road. D. Smith & Vasquez 4726 (F,

MO). Loreto: Balsapuerto (lower Rio Huallaga basin), Kil-

lip & Smith 28410 (GH, us). Huanuco: Prov. Pachitea,

Dist. Honoria, en frente a Tournavista, al este del Rio

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

16121 (GH). Madre de Dios: Prov. Tambopata, SSW of

Puerto Maldonado, Barbour 5178 (F, GH, MO).

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

No. 2. 1808. LECTOTYPE (designated by

Proctor, Flora Lesser Antilles 2: 48. 1977):

Sloane Herb. 1: 85, based on plant from Mt.

Diablo, Jamaica (BM).

Danaea elliptica var. crispula Rosenst., Repert. Spec.

Nov. Regni Veg. 7: 310. 1909. TYPE: Peru, San

Martin, "In montibus secus flumen Mayo, prope

Tarapoto," Spruce 4770 (holotype, K; isotypes,

GH!, L, P!, uc!. us!; photo, F of L!).

Sterile leaves to 1 m long and 30 cm broad,

imparipinnate; petiole with 1-5 nodes, sparsely

scaly, the scales appressed, dark brown, amor-

phous, peltate, often fimbriate, commonly less than

2 mm long; rachis nonalate to narrowly so distally

(or alate throughout in juvenile leaves), sparsely

scaly; pinnae 2-6 pairs, subsessile to short-stalked,

more or less elliptic, 8-18 cm long, (2.5-)3— 4.5 cm

broad, base cuneate, apex acute to acuminate,

margins entire and plane to somewhat undulate,

abaxial surface and veins very sparsely provided

with minute (ca. 0. 1 mm) scales, these appressed,

nearly circular, dark or reddish brown, commonly

erose to stellate; veins mostly paired or 1 -forked

near the base. Fertile pinnae 3-5 pairs, 5-14 cm

long, (l-)1.7-2.8(-3) cm broad, short-stalked, the

apex acute to acuminate.

In dense forests, principally along the Cordillera

Oriental, Amazonas to Junin, 125-1600 m.

Southern Mexico to Panama; West Indies; the

Guianas to Colombia and south to Brazil and Peru.

There is no justification in recognizing Rosen-

stock's var. crispula as a separate entity; suppos-

edly it differed from the typical in having one less

pair of pinnae, these broadest at or above the mid-

dle, with margins more crisped. All fall within the

natural variation of this species.

Amazonas: Prov. de Bagua, valley of Rio Maranon

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

Forestal Jenaro Herrera, Spichiger et al. 1439 (MO).

Huanuco: Dist. Churubamba, Hacienda Mercedes, trail

to Balsa-playa, Mexia 817 Oa (uc). Pasco: Prov. Oxa-

pampa, Palcazu Valley, Iscozacin, Foster 9502 (MO).

Junin: Pichis Trail, Yapas, Killip & Smith 25507 (GH,

NY, US).

3. Danaea oblanceolata Stolze, Amer. Fern J. 77:

33. 1 987. TYPE: Peru, Dept. Pasco (as Junin),

Cahuapanas, on Rio Pichis, Killip & Smith

26777 (holotype, us!; frag., F!).

Sterile leaves 40-50 cm long, 13-18 cm broad,

apical segment (on mature leaves) replaced by a

proliferous bud; petiole with 1-2 nodes, moder-

ately to abundantly scaly; rachis narrowly alate;

pinnae 10-12 pairs, mostly short-stalked, oblong

to (more commonly) broadly oblanceolate, larger

ones 7-11 cm long and 2-2.8 cm broad, inequi-

lateral at base, narrow and rounded to cordate

basiscopically, broader and cuneate acroscopical-

ly, terminating abruptly in an acuminate and ser-

18

FIELDIANA: BOTANY

rate apex, abaxial surface amply provided with

minute, dark brown scales; veins commonly sim-

ple, but sometimes paired at origin or forked. Fer-

tile pinnae 12-14 pairs, larger ones 7-8 cm long

and 0.8-1 cm broad, short-stalked, the apex ob-

tuse.

Terrestrial in dense forests, 0-500 m, thus far

known only from Peru: Pasco and Ucayali.

This is perhaps most closely related to D. alata

Sm., of the West Indies and Venezuela, especially

in that both species have predominantly simple

veins. However, in D. oblanceolata pinnae are

fewer and relatively shorter and broader, and most

are broadest well above the middle, where the

margins then bend abruptly to a short-acuminate

apex. In D. alata, as in all members of the D.

moritziana complex, pinnae are broadest at or near

the middle, from whence they taper gradually to

a moderately acuminate or attenuate apex.

Pasco: Oxapampa, Palcazu Valley, Iscozacin, ./?. Foster

9466 (MO), 10049 (F). Ucayali: Vicinity of Aguaytia, Croat

20938 (MO).

4. Danaea moritziana Presl, Abh. Konigl. Bohm

Ges. Wiss. 5(4): 35. 1845. TYPE: Venezuela,

Colonia Tovar, Moritz 257 (holotype, PR?; is-

otype, L; frag., us!; photo, us). Figure 4a.

Sterile leaves to 1.2 m long and 35 cm broad

(somewhat broader than fertile ones), imparipin-

nate, or sometimes with the apical segment re-

placed by a proliferous bud; petiole with 1-3 nodes,

sparsely scaly; rachis scarcely alate, or slightly so

in younger plants; pinnae 8-16 pairs, subsessile to

short-stalked, elliptic to oblong-lanceolate, larger

ones (5-)6-20 cm long and 1 .2-2.2 cm broad, ine-

quilateral at base, narrow and rounded to cordate

basiscopically, broader and cuneate to subtruncate

acroscopically, gradually tapering to an acuminate

or long-attenuate apex, margin subentire but con-

spicuously serrate at apex, veins (and often the

tissue between the veins) amply provided with mi-

nute scales; veins predominantly forked at or near

the base, occasionally simple, or paired at origin.

Fertile pinnae 8-15 pairs, larger ones 4-11 cm

long, 0.6-0.8 cm broad, subsessile or short-stalked,

the apex acute to apiculate.

Terrestrial, in dense forests, often along streams

or in ravines, 300-2300 m, Cajamarca to Loreto,

south to Cuzco and Madre de Dios.

Colombia; Venezuela; Peru.

Except for Danaea nodosa, this is the most widely

distributed representative of Danaea in Peru.

Moreover it is likely that the Central American D.

cuspidata Liebm. also belongs here, along with one

or two West Indian species. A number of taxa with

pinnae under 2.5 cm broad were separated by Un-

derwood (1902), merely on the degree of forking

and spacing of veins, an inconsistent character cor-

related rarely or not at all by other features, hence

a comprehensive revision of the genus should prove

many of these to be synonymous. Peruvian spec-

imens of D. moritziana apparently are more vari-

able than in other areas, sometimes having smaller

and fewer pinnae as in Venezuela, sometimes with

more and larger pinnae as in "D. cuspidata" of

Costa Rica. A few very large specimens from Tin-

go Maria have pinnae with rather glossy under-

surfaces and extremely attenuate apices. Very

closely related to this is D. alata Sm. of the Lesser

Antilles, Trinidad, and Venezuela. Although D.

alata tends to have more numerous pinnae than

D. moritziana, the only consistent difference be-

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

merely a synonym for D. alata.

Underwood (1902) described both of the latter

as having unforked veins and distinguished them

principally by the degree of crowding of veins, and

that D. stenophylla was a "small" plant. Later

(1909) he reversed himself by describing D. steno-

phylla as a "rather tall plant" and as having mostly

forked veins. Obviously he had not seen the type

collection, for the isotype (2 sheets) at Kew con-

tains leaves to nearly a meter long, with veins

obviously simple (only rarely forked), differing

from D. alata in no way that we can perceive. We

cannot be certain which West Indian plants with

forked veins were seen by Underwood, and later

by Proctor (1977), but perhaps they are D. ja-

maicensis.

Cajamarca: Tambillo, Jelski 1073 (P). Amazonas: Bag-

ua, ca. 25 km E of La Peca, Barbour 2969 (MO). San

Martin: Monte Campana, Tarapoto, Spruce 4711 (P).

Loreto: Rio Corrientes, Cachuela, McDaniel 11198 (F,

GH, MO). Huanuco: Tingo Maria, Tryon & Tryon 5278

(F, GH, uc, us). Pasco: Prov. Oxapampa, San Alberto,

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

Luisiana, Dudley 10412 (GH). Madre de Dios: Prov. Manu,

Cerro de Pantiacolla, Rio Palotoa 10-15 km NNW of

Shintuya, Foster 10750 (F).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

19

5. Danaea humilis Moore, Index fil. 286. 1861.

LECTOTYPE (designated by Lellinger in Proc.

Biol. Soc. Wash. 89: 710. 1987): Peru, Dept.

San Martin, Tarapoto, Spruce 4769 (lecto-

type, K; isolectotypes, BM!, GH!, L!, P!, us!).

Sterile leaves 20-34 cm long, 3.5-7 cm broad,

larger than the fertile ones, imparipinnate, the ter-

minal segment equaling or longer than the largest

pinnae, with or without a proliferous bud; petiole

with 1-3 nodes, amply to abundantly scaly; lamina

slightly shorter than the petiole, the tissue between

the veins abundantly provided with minute scales;

rachis conspicuously alate, the margins of wings

sometimes revolute, but not crispate; pinnae 14—

24 pairs, oblong-elliptic, subfalcate, 2.2-3.5(-4) cm

long, 0.6-1(-1.4) cm broad, strongly inequilateral

at base, narrow and rounded basiscopically, much

broader and abruptly cuneate to truncate acros-

copically, apex acute to subacuminate and usually

serrulate, margin subentire, essentially plane; veins

predominantly forked above the base, yet many

of them simple. Fertile lamina scarcely alate along

rachis; pinnae 12-25 pairs, 1.1-1.4 cm long, 0.2-

0.3 cm broad, subsessile to short-stalked, mostly

acute to subapiculate.

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

smallest species of the genus, with even the larger

sterile pinnae commonly less than 3 cm long. Oth-

er than size, however, they share few other similar

features, as evidenced by the characters listed in

the key.

6. Danaea trichomanoides Moore, Index fil. 288.

1861. TYPE: Peru, "in monte Guayrapurima,

prope Tarapoto" (San Martin), Aug. 1856,

Spruce 4710 (holotype, K!; frag., NY!; photos,

F, GH; isotypes, BM!, P!).

Sterile leaves 10-12 cm long, 3—4 cm broad,

shorter than the fertile ones, with a terminal seg-

ment much shorter than the larger pinnae, or re-

placed by a proliferous bud; petiole lacking nodes,

abundantly scaly; lamina 2-4 times longer than

the petiole, the tissue between veins essentially

lacking scales; rachis broadly crispate-alate; pin-

nae 1 1-15 pairs, oblong, 1.5-2.2 cm long, 0.5-0.9

cm broad, subequilateral at the truncate base, apex

rounded to subacute, margin slightly crispate; veins

predominantly simple, rarely forked. Fertile lam-

ina with rachis scarcely alate; pinnae 1.4-1.7 cm

long, 0.3-0.4 cm broad, obviously stalked, apex

rounded.

Thus far known only from the type collection.

When Moore published the description of the

species, taken from a Spruce manuscript, he noted

that pinnae were "unequal at the base." Perhaps

this conclusion was drawn from observing the cris-

pate margins of some of the pinnae which were

partially folded on drying. However, study of the

two type specimens reveals that the bases of most

pinnae are subequally truncate and approximately

as broad on either side of the costa. This is in

marked contrast with most species of Danaea,

whose bases are commonly broader and less

abruptly tapered on the acroscopic side.

Danaea trichomanoides is comparable to D.

wendlandii Reichb. f., from Costa Rica, for the

two are similar in size, and have nodeless petioles

and crispate pinnae, with the presence (usually) of

a terminal bud. However, the latter has pinnae

with inequilateral bases, veins predominantly

forking, and rachis wing plane or revolute (not

crispate).

Family 3: OSMUNDACEAE

Osmundaceae Bercht. & J. S. Presl, Prir. rostlin 1:

272. 1820. TYPE: Osmunda L.

Stem massive, woody, decumbent to erect, lack-

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

cinate in vernation, densely caespitose, pinnate to

pinnately compound, monomorphic to (in Os-

munda) partly or fully dimorphic. Petiole with an

expanded, stipular base, not articulate to the stem.

Veins free. Sporangia not in definite sori, borne

abaxially along the segments or (in Osmunda)

completely replacing the vegetative tissue of the

lamina or of only some of the pinnae, exindusiate,

short-stalked, with walls 1 cell thick, the annulus

lacking or of only a few thickened cells near the

distal end. Spores uniform, green, 1 20-5 1 2 in each

sporangium.

The three genera of the Osmundaceae are rep-

resented by 1 5-20 species. Osmunda is nearly cos-

mopolitan, while Leptopteris and Todea are con-

20

FIELDIANA: BOTANY

fined to the Old World. This is a very old fern

family; only Ophioglossaceae and Marattiaceae are

considered to be more primitive. Like the other

leptosporangiate members of the Filicales, Os-

mundaceae have sporangial walls only one cell

thick, yet the family is similar to the Eusporan-

giatae in that the annulus is lacking or is composed

of only a few thickened cells.

References

BENEDICT, R. C. 1909. Osmundaceae, in N.

Amer. fl., 16: 27-28.

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

mundaceae, pp. 50-57, in Ferns and allied plants,

Springer- Verlag, New York.

I. Osmunda

Osmunda L., Sp. pi. 1063. 1 753. TYPE: O. regalis

L. Figure 5.

Plants terrestrial. Leaves coarse, 1-2 m or long-

er, 1-2-pinnate, completely or partially dimor-

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

moderately provided with trichomes. Veins free,

at least 1 -forked. Sporangia relatively large, de-

veloping simultaneously, commonly borne in

clusters on the segments of the fertile leaves. Spores

tetrahedral-globose, trilete, rugose or crested with

slender echinate processes.

This is a genus of about 1 0 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.

TYPE: United States, Maryland, Kalm (ho-

lotype, LINN 1 244. 1 2; photo, F). Figure 5c-d.

Leaves completely dimorphic, vegetative tissue

essentially lacking in fertile ones. Petiole glabrous

or moderately provided with tortuous, filiform,

pluricellular, reddish brown trichomes, sheathed

at the base. Sterile lamina 1-pinnate-pinnatifid,

tissue glabrous, but with filiform trichomes as on

the petiole appearing especially at bases of pinnae,

rachis narrowly alate adaxially; pinnae to 10 cm

long, spreading to strongly ascending, subopposite,

articulate with age at the rachis, deeply incised to

form broad, subfalcate, obtuse or subacute seg-

ments, the segment margins subentire and bearing

short, filiform trichomes; veins 1 -forked. Fertile

lamina nearly 2-pinnate, the pinnae more strongly

ascending than the sterile ones, with dark, filiform

trichomes abundantly interspersed among the

crowded sporangia.

In wet places, usually in sphagnum swamps,

1 700-2800 m, Cajamarca, Amazonas, and Pasco.

Eastern and central United States and Canada;

West Indies; southern Mexico to Honduras; Costa

Rica; Colombia; Venezuela; Brazil; Peru; Para-

guay; southeast Asia.

Some earlier authors recognized this in the Neo-

tropics as var. imbricala (Kunze) Milde, distin-

guishing it from North American plants on the

basis of darker-colored tomentum and thicker lam-

ina. We find little justification for the separation.

Cajamarca: Jaen, Quebrada de Pajonal, above Taba-

conas, 19 km ESE Huancabamba, Fosberg 27795 (MICH,

NY). Amazonas: Chachapoyas, Jalca zone, 3-6 km W of

Molinopampa, Wurdack 1404 (us). Pasco: (as Junin)

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.

Gray, Manual ed. 2: 600. 1856. Figure 5a-b.

Osmunda spectabilis Willd., Sp. pi. 5: 98. 1810. TYPE:

United States, Pennsylvania, Muhlenberg (holo-

type, B, Herb. Willd. 19504; photo, GH).

Osmunda palustris Schrader, Gott. Gel. Anz. 866.

1824. TYPE: not located.

Osmunda regalis ssp. palustris (Schrader) Love & Love,

Taxon 26: 324. 1977.

length of their stalks. The characters are too in-

significant and inconsistent to merit such a dis-

tinction.

Amazonas: Mendoza, on rocky wall of ravine, Woyl-

kowski 8253 (GH, MO, us). Prov. Bongara, W & S margins

of Laguna Pomacocha, Wurdack 896 (us). Pasco: Prov.

Oxapampa, Canyon de Huancabamba, Leon 612 (GH).

Cuzco: Empalizada, Biies 1705 (us).

Leaves partly dimorphic, fertile ones with fertile

and sterile portions combined on the same lamina,

the fertile pinnae commonly distal and essentially

lacking vegetative tissue. Petiole glabrous at ma-

turity, sheathed at the base. Sterile lamina 2-pin-

nate, chartaceous to subcoriaceous, tissue gla-

brous, but a few tortuous, pluricellular, reddish

brown trichomes often clustered at the base of

pinnules, the rachis nonalate; pinnae to 20 cm

long, strongly ascending, subopposite, short-

stalked, articulate with age at the rachis, costae

very narrowly alate (at least distally) on adaxial

side; pinnules distant, subopposite, short-stalked,

articulate with age at the costa, oblong to narrow-

elliptic, obtuse to subacute, margins serrulate. Veins

mostly 2-forked. Fertile pinnae 2(-3)-pinnate, more

strongly ascending than the sterile ones, the crowd-

ed sporangia almost completely replacing the leaf

tissue.

In damp places near lakes or streams, occasion-

ally among rocks, 1300-2200 m, Amazonas, Pas-

co, Cuzco.

Eastern and central United States and Canada;

West Indies; southern Mexico; Guatemala; Hon-

duras; Costa Rica; Colombia; Venezuela; Brazil;

Ecuador, Peru; Paraguay; Uruguay.

Variety regalis, which occurs in the Old World,

is said to differ in its broader leaves, narrower

panicles, and more numerous blackish to casta-

neous trichomes persistent along the rachis. Al-

though specimens we have seen from both the

New and Old World do not seem to differ strongly

or consistently enough to warrant separation, we

maintain the traditional varietal distinction for

purposes of this treatment. Earlier workers pro-

posed yet another variant, ssp. palustris, for South

American plants, based on size of segments and

Family 4: SCHIZAEACEAE

Schizaeaceae Kaulf., Wesen farrenkr. 119. 1827.

TYPE: Schizaea Sm.

Stem erect to decumbent, usually small and

sometimes branched, or long-creeping, slender and

freely branched, provided with trichomes or (in

Mohria of Africa) with scales. Leaves circinate in

vernation, entire or filiform, or dichotomous, or

pinnate, glabrous to pubescent or (in Mohria) with

scales, partially or wholly dimorphic. Sporangia

borne abaxially on slightly to strongly modified

portions of the leaf, separate, or crowded on each

side of a vein, or in loose clusters on wholly fertile

panicles, sessile, or with a short, many-rowed stalk,

and an apical annulus. Spores tetrahedral-globose

and trilete, or ellipsoidal and monolete, lacking

chlorophyll.

This is an old family, with the four extant genera

so highly diverse in habit and form that some

authors prefer to segregate them into distinct fam-

ilies. Mohria is confined to Africa and Madagas-

car; Anemia is in America, Africa, and southern

India; the other two genera are pantropic. All ex-

cept Mohria occur throughout tropical America

and are all well represented in Peru.

References

LELLINGER, D. B. 1969. Schizaeaceae, in The

botany of the Guayana Highland— Part VIII.

Mem. New York Bot. Gard., 18: 1-11.

MAXON, W. R. 1909. Schizaeaceae, in N. Amer.

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

Anemia Sw., Syn. fil. 6, 155. 1806, nom. conserv.

(sometimes as "Aneimia"). TYPE: Anemia

phyllitidis (L.) Sw. (Osmunda phyllitidis L.).

Figure 6.

Coptophyllum Gardn., London J. Bot. 1: 133. 1842.

TYPE: Coptophyllum buniifolium Gardn. = Ane-

mia buniifolia (Gardn.) Moore.

Anemirhiza J. Sm., Seemann, Bot. Voy. Herald 243.

1854. TYPE: Anemirhiza adiantifolia (L.) J. Sm.

(Osmunda adiantifolia L.) = Anemia adiantifolia

(L.) Sw.

Plants terrestrial. Stem decumbent to creeping,

rarely erect, bearing short to long trichomes and

fibrous roots. Leaves ca. 1-75 cm long, erect, par-

tially (as in Peruvian species) dimorphic, with a

pair of fertile pinnae at the base of or below the

sterile lamina, or with similar leaves longer and

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

free, or rarely anastomosing without included free

veinlets. Sporangia borne on fertile segments which

are reduced to mere axes or have narrow borders

of laminar tissue along the axes. Spores tetrahe-

dral-globose, trilete.

The genus contains about 80 species, which oc-

cur in tropical to subtropical regions of both hemi-

spheres, predominantly in the Neotropics. Species

limits are not always clearly drawn, as diagnostic

features are often quite variable, such as: relative

length of fertile pinnae and sterile portion of lam-

ina; color of rhizome indument; degree of dissec-

tion of pinnae. Color of trichomes is particularly

difficult to assess, inasmuch as each observer sees

colors in different ways, and definitions of color

terms are usually difficult to interpret. Rhizome

indument in Anemia generally grades from orange

to reddish orange to red to brownish red; in many

specimens the distinctions are clear, but in others

they are somewhat inconstant. Hybridization is

not uncommon in the genus and must be taken

into consideration when attempting identification

of collections. The following key will be more ef-

fective if attention is given to combinations of

characters, whenever possible.

Reference

MICKEL, J. T. 1962. A monographic study of the

fern genus Anemia, subgenus Coptophyllum.

Iowa State Coll. J. Sci., 36: 349-482.

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

e. Lamina and petiole conspicuously lanate, petiole yellow 5. A. myriophylla

Fertile pinnae 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. Petiole (at least the fertile) 8-25 cm long; lamina gradually reduced to a pinnatind apex (or

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.

1966. TYPE: Peru, Junin, along Rio Perene,

Killip & Smith 21594 (holotype, us!; isotypes,

F!, NY!).

Stem with apex ascending and thickly covered

with petiole bases, densely provided with orange

trichomes. Leaves 10-25 cm long, 4-9 cm broad,

pilose to hirsute, the suberect fertile pinnae borne

8-16 mm below the sterile portion of the lamina;

petiole dark brown, less than 1 mm thick. Sterile

lamina 1 -pinnate-pinnatisect to 2-pinnate, with 5-

1 1 pairs of pinnae; pinnae dissected nearly or quite

to the costa, the ultimate segments joined at the

base or discrete, but broadly adnate, not stalked;

veins free. Fertile pinnae suberect, usually shorter

than the sterile portion of the lamina, short-pe-

tiolulate (stalk 0.3-0.8 cm long), the ultimate fer-

tile segments with narrow bands of laminar tissue

on each side of the axes.

In thickets and deep forests, 600-1 300 m, Junin.

Panama; Colombia and Venezuela to Bolivia

and western Brazil.

One of the distinguishing features of A clinata

is the suberect or slightly spreading position of the

fertile pinnae, which are rigidly erect in most species

of Anemia. It is also one of the smaller and more

delicate species, with slender, wiry petioles less

than 1 mm thick.

Junin: Near La Merced, E of Quimiri Bridge, Killip &

Smith 23591 (NY, us). Colonia Perene, Killip & Smith

25036 (F, NY, us). La Merced, Kunkel 564 (OH). Chan-

chamayo Valley, C. Schunke 78 (F, us).

2. Anemia villosa Willd., Sp. pi. ed. 4, 5: 92. 1 8 1 0.

TYPE: "America meridionale," Humboldt &

Bonpland (holotype, B, Herb. Willd. 19496;

photos, GH, us). Venezuela, Prov. Cumana,

"Between Catuaro and Cariaco": fide HBK.,

Nov. gen. sp. fol. ed. 1: 26. 1815.

Stem with apex ascending and thickly covered

with petiole bases, or the apex horizontal and ex-

posed beyond the most recent petioles, densely

provided with orange to brownish red trichomes.

Leaves 10-55 cm long, villous to hirsute, the erect

fertile pinnae borne 3-18 mm below the base of

the sterile portion of the lamina; petiole yellow to

light brown, 1-2 mm thick. Sterile lamina 1 -pin-

nate to pinnate-pinnatisect, with 7-19 pairs of pin-

nae; pinnae with rounded lobes, not cut entirely

to the costa (except rarely as to proximal lobes of

basal pinnae); veins free. Fertile pinnae erect,

slightly shorter than to (more typically) somewhat

longer than the sterile portion of the lamina, their

stalks 1-5 cm long, the ultimate fertile segments

with narrow bands of laminar tissue on each side

of the axes.

On rocky, open slopes, 900-2000 m, Cajamarca,

Amazonas, San Martin, Ayacucho, Cuzco.

Surinam to Colombia, south to Peru; east coast

of Brazil.

This appears to differ only quantitatively from

Anemia flexuosa, the latter with sterile pinnae more

strongly dissected and, typically, with fertile pin-

nae shorter than the sterile portion of the lamina.

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

it. Evidently these are the reasons why Prantl con-

sidered it merely a variety of A. flexuosa. See Mickel

(1962) for more detailed discussion.

Cajamarca: Prov. Jaen, San Patricio, Chontali, Chi-

moy65 (USM). Amazonas: Prov. Chachapoyas, Cano Santa

Lucia, Wurdack 599 (F, GH, NY, uc, us). San Martin:

Tarapoto, in monte Campana, Spruce 4708 (GH, NY, p,

us). Ayacucho: Prov. La Mar, Aina, Ldpez & Soukup

15.088 (GH). Cuzco: Prov. La Convencion, Urusaiwa,

Vargas 22349 (GH).

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

1806.

Osmunda flexuosa Savigny in Lam., Encycl. 4: 652.

1797. TYPE: based on undesignated specimen in

Jussieu herbarium (not located).

Anemia flexuosa var. setosa Prantl, Unters. Morph.

Gefasskrypt. 2: 95. 1881. SYNTYPES: Brazil,

Lagoa Santa, Warming (not located): Peru, Cu-

chero, Poeppig (not located). ISOSYNTYPE:

Poeppig (L; photos, GH, us).

Stem with apex ascending and thickly covered

with petiole bases, densely provided with orange

or reddish orange trichomes. Leaves 18-65 cm

long, hirsute or glabrate, the erect fertile pinnae

borne 5-25 mm below the base of the sterile por-

tion of the lamina; petiole yellow to light brown,

1-2 mm thick. Sterile lamina 2-pinnate (some-

times 2-pinnate-pinnatifid), with 6-12 pairs of

pinnae; pinnae (many of them) divided to the costa

into broadly adnate, obtuse pinnules; veins free.

Fertile pinnae erect, commonly shorter than the

sterile portion of the lamina (rarely equal to it or

slightly longer), their stalks 1.5-5 cm long, the

ultimate fertile segments with narrow bands of

laminar tissue on each side of the axes.

In thickets and open woods or on moist rocky

slopes or clay banks, 900-3000 m, Amazonas,

Huanuco, Junin, Ayacucho, Cuzco.

Surinam to Colombia, south to Bolivia.

Many Peruvian specimens of Anemia flexuosa

are to be found in various herbaria; especially

abundant are collections from Huanuco, Junin,

and Cuzco. This is a highly variable species, par-

ticularly in the dissection of sterile laminae, some-

times 2-pinnate-pinnatifid, typically 2-pinnate,

occasionally approaching the 1 -pinnate-pinnatifid

condition of A. villosa (q.v.) and thus is often con-

fused with it. As in some other species of Anemia,

this tends to hybridize; in fact there was a hybrid

found in Cuzco: A. flexuosa x A. phyllitidis, Var-

gas 19836, Prov. La Convencion, alt. 1650 m (GH).

Leaves are much thinner in texture than typical

A. flexuosa, with some anastomosing veins, and

fertile pinnules are long-petiolulate.

Amazonas: Rodriguez de Mendoza, Soukup 5023 (us).

Huanuco: Vilcabamba, on Rio Chinchao, Macbride4985

(F, NY, us), Stork & Morton 9876 (F, uc, us). Junin:

Huacapistana, Killip & Smith 24152, 24324 (F, NY, us).

Ayacucho: Ccarrapa, between Huanta and R;o Apuri-

mac, Killip & Smith 22319 (F, GH, NY, us). Cuzco: Prov.

La Convencion, Valley of the Sambray, Mexia 8038 (F,

GH, MO, uc).

4. Anemia ferruginea HBK., Nov. gen. sp. 1: 32.

1815, var. ferruginea. TYPE: Venezuela, Prov.

Cumana, "prope Guardia de San Augustin,"

Humboldt & Bonpland (holotype, P?).

Stem with apex horizontal, and exposed beyond

the most recent petioles, densely provided with

red or brownish red trichomes. Leaves 10-55 cm

long, the erect fertile pinnae borne 3-7 mm below

the base of the sterile portion of the lamina; petiole

commonly atropurpureous (rarely yellowish), gla-

brate, 1-2 mm thick. Sterile lamina 2-pinnate,

hirsute to subglabrous, with 8-12 pairs of pinnae;

pinnae dissected to the costa, the pinnules mostly

discrete, adnate at base, acute or subacute at apex,

sometimes those of proximal pinnae shallowly

pinnatifid; veins free. Fertile pinnae erect, com-

monly longer than the sterile portion of the lamina,

their stalks 1 .5-6 cm long, the ultimate fertile seg-

ments with narrow bands of laminar tissue on each

side of the axes.

In sunny places, open woods and clearings, on

rocky slopes and clay banks, 850-2200 m, San

Martin, Huancavelica, Cuzco.

Honduras; Guyana to Colombia, south along

the Andes to Bolivia and Brazil.

The variety ahenobarba (Christ) Mickel, from

Brazil, differs in its more deeply dissected pinnules

with acute to acuminate tips. Both are difficult to

distinguish from Anemia tomentosa (Savigny) Sw.,

especially var. anthriscifolia (Schrader) Mickel,

from northern Argentina, Bolivia, Paraguay, and

southeastern Brazil. See Mickel (1962) for further

comments.

San Martin: Tarapoto, Spruce 4044 (GH, NY, p, us).

Huancavelica: Prov. Tayacaja, Dist. Huachocolpa, near

Quintabamba, Tovar 4157, 4693 (GH). Cuzco: Prov.

Convencion, Valley of the Sambray, Mexia 8038 (NY,

us). Prov. Convencion, Hacienda Sahuayaco, Vargas

1656 (GH, MO).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

27

5. Anemia myriophylla Christ, Bull. Herb. Bois-

sier 2(7): 793. 1907. TYPE: Bolivia, Padcaya,

Fiebrig 2541 (isotype, p).

Stem with apex horizontal and exposed beyond

the most recent petioles, densely provided with

reddish to orange trichomes. Leaves 20-50 cm

long, the erect fertile pinnae borne 3-8 mm below

the base of the sterile portion of the lamina; petiole

yellow, conspicuously lanate, 1-2 mm thick. Ster-

ile lamina 2- to nearly 3-pinnate, conspicuously

lanate, with 7-12 pairs of pinnae; pinnae with pin-

nules cut deeply or quite to the costule, the tips of

ultimate segments commonly broadly rounded;

veins free. Fertile pinnae erect, commonly longer

than the sterile portion of the lamina, their stalks

1.5-5 cm long, the ultimate fertile segments with

narrow bands of marginal tissue on each side of

the axes.

On open slopes and rocky banks, 2000-2200 m,

Cajamarca, Amazonas.

Peru; Bolivia; Argentina.

Anemia myriophylla has a more highly divided

leaf than any Anemia in Peru. This and the densely

lanate lamina and petiole make it rather easy to

recognize.

Cajamarca: Between San Marcos and Cajabamba,

Correll & Smith P911 (GH, us). Prov. Celendin, Sagds-

legui et al. 8457 (F, HUT, MO, uc). Amazonas: W of

Chachapoyas on road to Gaelic. Hutchison & Bennett

4519 (F, GH, MO, NY, uc, us).

6. Anemia hirsuta (L.) Sw., Syn. fil. 155. 1806.

Osmunda hirsuta L., Sp. pi. 1064. 1753. LECTO-

TYPE (designated by Lellinger in Proc. Biol. Soc.

Wash. 98: 387. 1985): Plumier, Traite foug. Amer.,

/. 762, based on a Plumier collection from His-

paniola.

Anemia hirsuta var. humboldtiana Hieron., Bot. Jahrb.

Syst. 34: 566. 1905. LECTOTYPE (designated by

Lellinger in Proc. Biol. Soc. Wash. 98: 366. 1985):

Venezuela, border Edo. Sucre- Monagas, Hum-

boldt 459 (B, Herb. Willd. 19495-2; photos, GH,

us).

Stem short-creeping, densely provided with or-

ange trichomes. Leaves to 35 cm long and 5 cm

broad, glabrate or somewhat villous on axes and

lamina, the erect, fertile pinnae borne at the base

of the sterile portion of the lamina; petiole (at least

on fertile leaves) 8-1 8 cm long. Sterile lamina pin-

nate-pinnatifid to nearly 2-pinnate, with 7-1 2 pairs

of pinnae, gradually tapering to a pinnatifid apex;

pinnae to 2.5 cm long and 0.8 cm broad, com-

monly inequilateral at base, truncate acroscopi-

cally and cuneate basiscopically, obliquely incised

into linear or narrowly cuneate segments, obtuse

to subacute at apex; veins free. Fertile pinnae to

18 cm long, the stalk (1.5-)2-3 times the length of

the panicle, the ultimate fertile segments essen-

tially lacking laminar tissue.

In dry, grassy areas, on slopes or mesas, 1 200-

2 1 50 m, Huanuco, Cuzco.

Mexico to Panama; Greater Antilles; Trinidad

& Tobago; Venezuela and Colombia south to Bo-

livia and Brazil.

This is scarcely distinct from Anemia pastina-

caria, essentially differing only by the characters

noted in the key. More detailed studies are needed

to clarify relationships of the several variable taxa

of the A. hirsuta group which, furthermore, appear

to hybridize readily with a number of other species

of Anemia throughout the entire range. For ex-

ample, there is a specimen of A. pastinacaria at

Paris (San Martin, Tarapoto, Spruce 4134) which

also contains a very large, highly dissected leaf

with abortive spores— possibly a hybrid between

A. hirsuta and A. phyllitidis.

Huanuco: Muna, Bryan 427 (F, GH). In grass steppe,

Woytkowski 219 (uc). Cuzco: Quillabamba, Coronado

124 (uc). Dept. Unknown: Mat hews 3299 (us).

7. Anemia pastinacaria Prantl, Unters. Morph.

Gefasskrypt. 2: 110. 1881. LECTOTYPE

(designated by Lellinger in Proc. Biol. Soc.

Wash. 98: 367. 1 985): Venezuela, "in convalli

del Tigre," Moritz 26 (B; probable isolecto-

type, GH!). Figure 6a-b.

This differs from Anemia hirsuta essentially only

in the following characters: Leaves to 45 cm long

and 7 cm broad. Sterile lamina gradually tapering

to a pinnatifid apex, but occasionally with a dis-

tinct, nonconform apical segment; pinnae to 3.5(-5)

cm long and 1 .5(-2) cm broad, margins denticulate

or crenate, or rarely with a few deep lobes on sev-

eral proximal pinnae.

On dry, rocky slopes or in forest clearings, 750-

2000 m, San Martin, Loreto, Huanuco, Junin, Ay-

acucho, Cuzco.

Mexico to Panama; Cuba; Jamaica; Trinidad;

Surinam to Colombia, south to Bolivia and Brazil.

This and Anemia hirsuta differ only in their rel-

ative size and depth of dissection of pinnae, and

28

FIELDIANA: BOTANY

a few specimens in Peru tend to be intermediate

even in these characters. See A. hirsuta for further

discussion.

San Martin: Tarapoto, Spruce 4 134 (p in part). Loreto:

Yanayacu, Biies 1037 (us). Huanuco: Yanano, Macbride

3816 (F, us). Junin: Prov. Huancayo, Pariahuanca, Tovar

7948a (USM). Ayacucho: Aina, between Huanta and Rio

Apurimac, Killip & Smith 22614 (F, NY, us). Cuzco: Prov.

Convencion, Hacienda Sahuayaco, Vargas 1655 (GH),

1657 (GH, MO).

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

1806.

Osmunda oblongifolia Cav., Icon. 6: 69, /. 592, / 2.

1801. TYPE: Panama, Nee (holotype, MA), ver-

ified by C. Christensen [Dansk. Bot. Ark. 9(3):

31. 1937].

Osmunda humilis Cav., Icon. 6: 69, /. 592, f. 3. 1801.

TYPE: Panama, Taboga Island, Nee(nol located);

earlier presumed at MA, but not seen by C. Chr.

[Dansk. Bot. Ark. 9(3): 31. 1937].

Anemia humilis (Cav.) Sw., Syn. fil. 156. 1806.

Anemia oblongifolia var. humilis (Cav.) Hooker &

Baker, Syn. fil. 431. 1868.

Stem decumbent, rather densely provided with

orange trichomes. Leaves to 30 cm long and 3.5

cm broad, sparsely to densely villous on axes and

lamina, the erect fertile pinnae borne at the base

of the sterile portion of the lamina; petiole com-

monly less than 6 cm long. Sterile lamina 1 -pin-

nate, with 2-9 pairs of pinnae, abruptly terminat-

ing in an obovate or obdeltoid apical segment which

is often as broad as long; pinnae to 1.8 cm long

and 0.7 cm broad, strongly inequilateral at base,

truncate or rounded acroscopically and cuneate

basiscopically, broadly rounded at apex, the mar-

gins crenulate or subentire; veins free. Fertile pin-

nae to 1 8 cm long, the stalk 2—4 times the length

of the panicle, the ultimate fertile segments essen-

tially lacking laminar tissue.

In forests, on rocky slopes, 1000-1 100 m, Cuz-

co.

Mexico to Panama; Venezuela to Brazil and Bo-

livia.

Anemia oblongifolia is rather distinctive in its

coarse pinnae which are broadly rounded at their

tips and often nearly as broad as long. Sterile leaves

are commonly very short-petiolate (often subses-

sile), and typically are so densely caespitose that

they give the appearance of a rosette. Some authors

have separated A. humilis on the strength of its

villous sterile pinnae with crenulate margins; but

these characters are too variable and inconsistent

to be of real value. We tentatively maintain it here

as a synonym, although we have not seen the type

to fully substantiate this.

Cuzco: Echarate, Camino de Sahuayacu, Bites 845 (us).

Ccochayoc, Biies 1712, 1719 (us). Prov. Convencion,

Valle de Lucumayo-Pistipata, Vargas 4176 (MO, uc, us).

9. Anemia phyllitidis (L.) Sw., Syn. fil. 155. 1806.

Figure 6c.

Osmunda phyllitidis L., Sp. pi. 1064. 1753. TYPE:

based on illustration of Hispaniola plant in Plu-

mier, Traite foug. Amer., /. 156. 1705.

Anemia haenkei Presl, Reliq. haenk. 1: 74. 1825.

TYPE: Peru, "in vallibus Cordillerarum Peru-

viae," Haenke (holotype, PRC?).

Stem decumbent (rarely short-creeping), rather

densely clothed with orange or brownish tri-

chomes. Leaves to 80 cm long and 20 cm broad,

the erect fertile pinnae borne at the base of the

sterile portion of the lamina or less than 1 mm

below it; petiole (at least on fertile leaves) ca. 10-

35 cm long. Sterile lamina 1 -pinnate, with 3-7

pairs of pinnae and a conform apical one; pinnae

(larger ones) 5-14 cm long and 1.3-3 cm broad,

subequilateral, ovate to lanceolate, acute to acu-

minate, the margins crenulate to dentate (or rarely

the basal pair lobed); veins copiously anastomos-

ing. Fertile pinnae to 40 cm long, the stalk equaling

or (commonly) shorter than the panicle; the ulti-

mate fertile segments essentially lacking laminar

tissue.

Moist, often rocky slopes and ravine banks, in

and at edges of forests, 700-1700 m, Amazonas

to Loreto, south to Huancavelica and Cuzco.

Mexico to Panama; Greater Antilles; Trinidad;

Venezuela and Colombia to Argentina and Uru-

guay.

This is one of the most common species of Ane-

mia in Peru and, with its large, subentire pinnae

and reticulate venation, is the easiest to recognize.

Amazonas: Prov. Bagua, S of Bagua Grande on Rio

Utcubamba, Hutchison 1479 (uc, us). San Martin: Ze-

pelacio, near Moyobamba, Klug 3440 (F, MO, NY, uc,

us). Loreto: Yanayacu, Biies 2005 (us). Huanuco: Prov.

Huanuco, Puente Durand, Stork & Morton 9432 (F, GH,

MO, uc, us). Pasco: Oxapampa, Soukup 2361 (F, GH).

Junin: Yucapata, Woytkowski 6642 (GH, MO, us). Huan-

cavelica: Prov. Tayacaja, SE of Tintay, Tovar 46 11 (GH).

Ayacucho: Aina, between Huanta and Rio Apurimac,

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,

GH, uc, us).

Comments

Anemia hispida Kunze, Linnaea 9: 20. 1834.

TYPE: "In fruticetis ad Chibangata fl. Pe-

ruv.." 1829, Poeppig diar. 1165 (LZ? de-

stroyed, B?, w?).

Apparently known thus far only from the type

collection.

Kunze's plant was described as having: creeping

rhizome; pinnate, hispid leaves; fertile pinnae twice

as long as the sterile lamina; pinnae remote, pat-

ent, sessile, dimidiate, oblong, obtuse, with mar-

gins crenate or subincised. He likened it to A nemia

repens Raddi (= A. hirsuta1?), but having pinnae

less incised. Assuming it belongs to subg. Anemia

(unsubstantiated by the description), it could be

A. pastinacaria, but we cannot be certain without

examining the type.

II. Lygodium

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

1 802, nom. consent TYPE: Lygodium scan-

dens (L.) Sw. (Ophioglossum scandens L.).

Figure 7.

Plants terrestrial. Stem slender, short- to long-

creeping, provided with short trichomes. Leaves

1-10 m long, vinelike, indeterminate, spreading

and scandent, partially dimorphic, the fertile por-

tions somewhat contracted with marginal fertile

lobes, pinnate, glabrous to somewhat pubescent.

Pinnae short-stalked, pseudodichotomously

branched with an arrested bud in the axil, each

primary pinna-branch radiately lobed or branched

or pinnate. Veins reticulate or (in Peruvian species)

free. Sporangia borne separately on marginal lobes

of a pinna segment or on a wholly fertile segment,

each covered by a laminar outgrowth (flange).

Spores tetrahedral-globose, trilete.

Lygodium is a natural group, easily distin-

guished within the family by its long leaves and

vinelike and scrambling habit, often climbing to

considerable heights by twining around the small-

er branches of shrubs and trees. It is widely dis-

tributed, primarily pantropic, with several extra-

tropical species both in the New and Old World.

There are 30-35 species, but the number could be

somewhat reduced by further study, as some ap-

pear to intergrade rather freely. Three species are

recognized in Peru.

References

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

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

dium, pp. 69-76, in Ferns and allied plants,

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)

1801(2): 303. 1803. TYPE: Jakob Breyne,

Cent. I, t. 96. 1678, not "Brazil, Breynius"

(designated by Proctor, Flora Lesser Antilles

2: 51. 1977), a specimen which probably does

not exist. Figure 7a.

Pinna-stalks 2-10 mm long, moderately to

densely pubescent with tawny to orange pluricel-

lular trichomes. Primary pinna-branches pinnate

to (rarely) 2-pinnate, sessile to short-stalked,

broadly to narrowly deltoid. Fertile pinnules 9-

2 1 , alternate and commonly widely spaced, all but

the distal ones stalked, stalk often nodose at its

juncture with pinnule base, sparsely to abundantly

pilose on axes, veins and (occasionally) the leaf

tissue, larger ones 2.5-5 cm long, gradually di-

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,

the lobes rounded to cordate or subpalmate. Ster-

ile pinnules similar to the fertile, but often larger

and not much reduced toward the apex of the pri-

mary branch. Veins free, 1 -several times-forked,

prominulous.

Along roadsides and stream banks, in fields or

in thickets and open forests, usually twining over

low shrubs, 100-1000 m, San Martin, Loreto,

Huanuco, Junin, Ucayali, Cuzco.

Tropical America.

This species is widely distributed throughout the

Neotropics. A number of other species have been

confused with it or should be included within it.

For detailed discussions see Smith, Flora of Chia-

pas: Pteridophytes, Calif. Acad. Sci. p. 145. 1981,

and Stolze, Ferns & Fern Allies of Guatemala: Part

I, Fieldiana, Bot. 39: 38. 1976. The closely related

Lygodium volubile readily can be distinguished by

the fewer, larger, and subequal pinnules on each

pinna-branch; i.e., the distal ones are nearly or

quite as large as the proximal ones. Pinnules (at

least fertile ones) of L. venustum gradually dimin-

ish in size and shape so that subapical ones are

often one-third to one-half the size of those near

the pinna base. The Breyne plate was cited by

Swartz and is clearly this species.

San Martin: Dist. San Martin, 2 km NW of Tarapoto,

Belshaw 3353 (F, GH, MO, NY, uc, us). Juan Jui, Alto Rio

Huallaga. Klug 4218 (F, GH, MO, NY, uc, us). Loreto:

Puerto Arturo, lower Rio Huallaga below Yurimaguas,

Killip & Smith 27847 (NY, us). Huanuco: Prov. Leoncio

Prado, Huachipa, Plowman 5957 (F, GH). Junin: La

Merced, Killip & Smith 23379 (F, NY, us). Ucayali: Pu-

callpa (as Loreto), Soukup 3049 (F). Cuzco: Santa Ana,

Cook & Gilbert 1614 (us).

2. Lygodium volubile Sw., J. Bot. (Schrader)

1801(2): 304. 1803. TYPE: Jamaica, Swartz

(holotype, S-PA; photo, us). Figure 7b-c.

Lygodium micansj. W. Sturm in Mart., Fl. bras. 1(2):

178. 1859. TYPE: "British Guiana," Schom-

burgk 399 (holotype, B).

Pinna-stalks 0.5-3 mm long, essentially gla-

brous. Primary pinna-branches pinnate or (very

rarely) 2-pinnate at base, the pinnae stalked (1.5-

5 cm), broadly oblong. Fertile pinnules 4-10, al-

ternate and widely spaced, conspicuously stalked,

nodose and often articulate at juncture with the

pinnule base, glabrous to sparsely pilose on axes

and veins (especially at bases of sporangia), 6-14

cm long, proximal and distal ones subequal,

broadly cuneate to truncate at base, only rarely

lobed at base. Sterile pinnules similar to the fertile,

but often somewhat larger. Veins free, several

times-forked, prominulous.

In wet, often inundated, lowland forests and open

fields, scandent on shrubs and low trees, often along

stream and river banks, sea level to 250 m, Ama-

zonas, San Martin, Loreto, Madre de Dios.

Southern Mexico; Guatemala; Belize; Panama;

Greater Antilles; Trinidad; Venezuela to the

Guianas, south to Argentina and Brazil.

This and Lygodium micans have been separated

on the basis of several, variable, quantitative char-

acters, although these may appear more distinct

in certain regions outside South America: relative

size and shape of ultimate segments and their bas-

es; degree of pubescence on abaxial surface; angle

of veins. None of these are consistently correlated

in the many Peruvian specimens examined.

Amazonas: Quebrada Huampami, Kayap 1066 (MO).

San Martin: Prov. San Martin, Laguna Sauce, Ramirez

& Sotero 092-85 (F, GH, HUT). Loreto: Rio Tacsha Cur-

aray, Croat 20394 (F, GH, MO, NY). Gamitanacocha, Rio

Mazan, J. Schunke 234 (F, GH, NY, uc, us). Madre de

Dios: El Pilar, Lopez & Soukup 4601 (GH).

3. Lygodium radiatum Prantl, Unters. Morph.

Gefasskrypt. 2: 66. 1881. TYPE: based on L.

digitatum D. C. Eaton.

Lygodium digitatum D. C. Eaton, Mem. Amer. Acad.

Arts n.s. 2, 8: 217. 1860 (not Presl, 1825). LEC-

TOTYPE (inferentially designated by Duek,

Feddes Repert. 89: 417. 1978): Panama, Gatun,

Hayes 25 (YU; isolectotype, us!).

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

tions along the primary axis (or sometimes ob-

solete). Primary pinna-branches long-stalked (1.5-

4 cm), the pinnae radiate (subpalmate), divided

nearly to the base into (2)3-7 ultimate segments.

Fertile ultimate segments linear-lanceolate to nar-

rowly oblanceolate, 5-15 cm long and 1.5-2 cm

broad, the margins serrate, glabrous, or sparsely

pubescent along the midrib. Sterile segments sim-

ilar to the fertile, but often somewhat longer (to

25 cm). Veins free, mostly 1- or 2-forked, prom-

inulous.

Rare, in wet forests, often along stream and river

banks, scandent on shrubs and small trees, 135-

850 m, Amazonas, Loreto, Pasco, Ucayali.

Panama; Colombia; Ecuador; Peru.

32

FIELDIANA: BOTANY

Amazonas: Al lado de Huampami, Kayap 1217 (MO,

us). Loreto: Santa Rosa, lower Rio Huallaga below Yu-

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

San Matias, D. Smith 2011 (F, MO). Ucayali: Prov. Co-

ronel Portillo, km 89 Carretera Federico Basadre, C.

Vdsquez 1 (USM).

Comments

Lygodium oligostachyum (Willd.) Desv., Mem. Soc.

Linn. Paris 6: 205. 1827.

Hydroglossum oligostachyum Willd., Sp. pi. ed. 4, 5:

81. 1810. TYPE: Plumier, Traite foug. Amer., t.

92. 1705 (based on specimens from Haiti, near

Lake Miragoan).

This is similar in general aspect to Lygodium

venustum, but differs in the conspicuously 2-pin-

nate pinnae and the strongly flexuous (zigzag) and

more delicate primary and secondary pinna rach-

ises. Although L. oligostachyum has been consid-

ered by most authors to be confined to Hispaniola,

Duek (1978) assigned to it a broad distribution

(West Indies; parts of Central and South America)

and cited two specimens at Prague from Peru:

Poeppigand Cuming, neither of which he had ex-

amined. Under L. oligostachyum he also placed

L. mexicanum Presl and two varieties of L. po-

lymorphyum (Cav.) HBK. (illeg.); however, plants

usually identified under either name have usually

turned out to be L. venustum. We have not seen

the Poeppig and Cuming collections in question,

but they are probably L. venustum and not L. oli-

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.

dichotoma (L.) Sm. (Acrostichum dichoto-

mum L.). Figure 8.

Lophidium Rich., Actes Soc. Hist. Nat. Paris 1: 114.

1 792. TYPE: Lophidium latifolium Rich. = Schi-

zaea elegans (Vahl.) Sw.

Actinostachys Wall., Numerical list of plants in East

Indies Company Museum, 1. 1829, description

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

chys digitata (L.) Wall. (Acrostichum digitatum

L.) = Schizaea digitata (L.) Sw.

Plants terrestrial. Stem erect or ascending, rath-

er densely provided with septate trichomes and

with slender, fibrous roots. Leaves ca. 5-50 cm

long, caespitose, long-petiolate, glabrous or with

scattered, small trichomes, slightly to partially or

wholly dimorphic, fertile ones with elongate fertile

segments borne pinnately or subdigitately at the

apex of laminar axes which are simple or dichot-

omously branched. Lamina filiform, grasslike and

scarcely or not at all foliaceous, or fusiform to

flabelliform (in general outline) and foliaceous.

Veins free. Sporangia crowded in 1 or more rows

on the scarcely foliaceous segments (sporangio-

phores). Spores ellipsoidal, monolete.

Schizaea sometimes has been divided into sev-

eral genera or subgenera on the basis of the more

expanded, foliaceous lamina (Lophidium) or the

subdigitate (Actinostachys) versus pinnate (Schi-

zaea) fertile branches, as well as some differences

in the gametophytes. Nevertheless, the groups are

quite closely allied and are here treated as a single

genus. About 30 species occur in tropical or ex-

tratropical regions of both hemispheres, five of

which are found in Peru.

Reference

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

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

Springer- Verlag, New York.

III. Schizaea

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

to obovate 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)

1800(2): 103. 1 80 1 . Figure 8.

Acrostichum elegans Vahl, Symb. hot. 2: 104, t. 50.

1791. TYPE: Trinidad, von Rohr (c).

Schizaea flabellum Mart., Icon. pi. crypt. 115, /. 55.

1834. TYPE: Brazil, Prov. Rio Negro, Martins

(holotype, BR; isotype, B; photo, us of B).

Lophidium elegans (Vahl) Presl, Suppl. tent, pterid.

77. 1845.

Lophidium flabellum (Mart.) Presl, Suppl. tent, pterid.

77. 1845.

Leaves 20-80 cm long, petiole 15-65 cm long,

1-2.5 mm in diameter, subterete, obtusely angled

abaxially. Lamina subcoriaceous, essentially gla-

brous, flabelliform (in general outline), 1 -several

times-dichotomously forked or cleft, the divisions

oblong to obovate, lateral margins entire, distal

ones strongly and sharply lacerate. Veins dichot-

omously forked within the ultimate divisions.

Sporangiophores borne along the distal margins

of the lamina, pinnately branched at the tips of

the marginal lacerations. Sporangia in a single row

on each side of the vein.

In humus on forest floor, or in sandy or rocky

soil on wooded slopes and ridges, 130-2200 m,

Amazonas, San Martin, Loreto, Huanuco, Pasco,

Junin, Cuzco.

Southern Mexico to Panama; Trinidad to Co-

lombia, and south to Bolivia and Brazil.

Several varieties have been recognized, based

upon number and shape of laminar divisions,

characters which do not appear to merit distinc-

tion.

Amazonas: Mendoza, Woytkowski8211 (GH, MO). San

Martin: Zepelacio, near Moyobamba, Klug 3415 (F, GH,

MO, NY, us). Loreto: Balsapuerto, Klug 2885 (F, GH, MO,

NY, us). Huanuco: Prov. Huanuco, Tingo Maria, Tryon

& Tryon 5292 (BM, GH, NY, u, uc, us, USM). Pasco: Prov.

Oxapampa, Cordillera San Matias, Leon 324 (F, GH).

Junin: La Merced, Hacienda Schunke, Macbride 5601

(F, us). Cuzco: Valle de Urubamba, Machu Picchu, Her-

rera 3299 (us).

2. Schizaea pusilla Pursh, Fl. Amer. sept. 2: 657.

1814. TYPE: United States, New Jersey, Bur-

lington Co., Quaker Bridge, Pursh? (not lo-

cated).

Leaves unbranched, 7-10 cm long, linear, grass-

like, with expanded lamina essentially lacking, pet-

iole 0.4—0.5 mm broad, rather flattened, or sub-

terete at base, sterile and fertile leaves erect and

straight or slightly flexuous, fertile ones apically

bearing a pinnatifid, conduplicate, scarcely folia-

ceous sporangiophore. Sporangiophore 0.5-0.7 cm

long, with 5-7 pairs of linear segments, abundantly

provided on the margins and among the sporangia

with deep orange to reddish brown, flexuous tri-

chomes. Sporangia in a single row of 5-7 on each

side of the vein.

Thus far known only from Pasco, "plant colo-

nizing sandy, wet landslide; shrubland on white

sandstone, 2700-2800 m."

United States (New Jersey, New York); Canada

(Newfoundland, Nova Scotia).

Heretofore, Schizaea pusilla has been known to

occur only in a few scattered locations in north-

eastern North America. The recent collection in

Pasco duplicates the northern plants in every fea-

ture (including spores) except that the sterile leaves

are straight to slightly flexuous instead of con-

spicuously curling. The great disjunction in dis-

tribution is at first startling until one understands

the problems of encountering this very inconspic-

uous fern. This and the other diminutive species

of Schizaea (like those of Ophioglossaceae and

Hymenophyllaceae) are very difficult to detect.

Hence true distribution patterns may never come

to light. It is most closely allied to S. incurvata,

under which see further discussion. Less closely

related is the Old World S.fistulosa Labill. and its

variety australis (Gaud.) Fosb. from Chile and Ar-

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-

gins of the fertile ultimate segments.

Pasco: Oxapampa, Cordillera de Yanachaga, Cerro

Pajonal, Foster 9065 (F, GH, MO, us).

3. Schizaea incurvata Schkuhr, 24 Kl. Linn. Pfl.-

Syst. 1: 138, t. 137. 1809. TYPE: "British

Guiana, habitat in India Occidentali circa Es-

sequebum," Gartner (not located).

Leaves simple to 1 - or 2-forked at an acute angle

(branches subparallel), mature ones ( 1 2-) 1 5-50 cm

long, linear, grasslike, with an expanded lamina

essentially lacking, petiole 0.6-0.8 mm in diam-

eter, convex abaxially, shallowly sulcate adaxially,

somewhat flattened, or terete at base, sterile and

fertile leaves straight or slightly flexuous, fertile

ones apically bearing a pinnatifid, conduplicate,

scarcely foliaceous sporangiophore. Sporangio-

phore 1.2-3 cm long, with (10-) 12-2 5 pairs of

linear segments, abundantly provided on the mar-

gins and among the sporangia with pale to light

brown, flexuous trichomes. Sporangia in a single

row of (8-) 10- 18 on each side of the vein.

Thus far known from a single collection (dept.

unknown) in Peru; elsewhere in open woods or

clearings and savannas, in sand or sandy or rocky

soil, sea level to 400 m.

Surinam to Venezuela and northern Brazil;

Peru.

Of the many specimens seen by us in various

herbaria, only one was (possibly) collected in Peru:

Poeppig (NY). The label is worded simply "In Pe-

ruvia, diar. Kunze, 1834." However, Kunze (Lin-

naea9: 19. 1834) lists only one species of Schizaea

from Peru (41. 5". dichotomd), which is S. poep-

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

the label of the New York sheet, the data are

suspect. That the collecting site is correctly des-

ignated is also problematical, for the known range

of the species is somewhat remote from Peru.

However, as noted in the discussion of 5. pusilla

(q.v.), the grasslike appearance of several species

of Schizaea masks them well in any savanna-type

vegetation, so that more specimens may yet be

discovered throughout the Neotropics after dili-

gent search, especially in Peruvian Amazonia in

sandy areas. Schizaea pusilla is similar to S. in-

curvata, both in general habit and pubescence of

the sporangiophore, but the two are easily sepa-

rated by the characters noted in the key.

4. Schizaea poeppigiana J. W. Sturm in Mart., Fl.

bras. 1(2): 181. 1859. SYNTYPES: Peru, "ad

Ventanilla de Cassapillo, 1829," Poeppig (w?);

"British Guiana, in montibus Canuku," Rich.

Schomburgk 1189(ei).

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

Amer. fl. 16: 38. 1909.

Leaves dichotomous, 12-40 cm long, dimor-

phic; petiole 10-30 cm long, ca. 1 mm broad, con-

vex abaxially, flattened to concave adaxially,

sparsely to amply pilose. Sterile lamina circular

in outline (often appearing conduplicate and fla-

belliform after pressing), 5-8 times dichotomously

forked, the divisions linear, scarcely or slightly

foliaceous. Fertile lamina on somewhat longer pet-

ioles than the sterile, 3 or 4 times dichotomous,

the divisions linear, nonfoliaceous, bearing spo-

rangiophores at their apices. Sporangiophores 1.5-

2 cm long, with 12-25 pairs of linear segments,

amply provided on the segment margins and among

the sporangia with pale to light brown, flexuous

trichomes. Sporangia in a single row of 1 5-25 on

each side of the vein.

Thus far known in Peru only from one of the

syn types.

Greater Antilles; Mexico (Chiapas); Costa Rica

to Venezuela and Guyana; Peru; Bolivia.

5. Schizaea pennula Sw., Syn. fil. 150, 379. 1806.

TYPE: "America meridionalis ... habitat in

America calidiore" (holotype, S-PA).

Actinostachys pennula (Sw.) Hooker, Gen. fil., /. Ilia.

1842.

Leaves ca. 1 2-50 cm long, unbranched, linear,

grasslike, with expanded lamina essentially lack-

ing; petiole 1-1.8 mm broad, triquetrous and the

faces sulcate, often flattened apically and subterete

at base, sterile and fertile leaves erect and straight

or slightly flexuous, fertile ones apically bearing a

subdigitate, scarcely foliaceous sporangiophore.

Sporangiophore 1-4 cm long, with 6-14 linear,

strongly ascending segments, these essentially with

entire and glabrous margins, and with abundant,

flexuous, reddish brown trichomes interspersed

among the sporangia. Sporangia in 2 or more

(sometimes indistinct) crowded rows on each side

of the vein.

36

FIELDIANA: BOTANY

In sand and sandy soil of open forests and clear-

ing, sea level to 260 m, San Martin, Loreto.

Costa Rica; Puerto Rico; Guadeloupe; Trinidad;

Surinam to Colombia, south to Bolivia and Brazil.

San Martin: Chazuta, Rio Huallaga, Klug 3986 (F, GH,

MO, NY, uc, us). Loreto: San Juan, vicinity of Iquitos,

Asplund 14414 (F, p, NY, us). Dist. Iquitos, Maynas, Que-

brada Shushuna, Rimachi 3995 (F, NY), 4870 (NY).

Family 5: GLEICHENIACEAE

Gleicheniaceae (R. Br.) Presl, Reliq. haenk. 1 : 70.

1825, as Order GleicheniaeR. Br., Prod. 160.

1811, as Tribe of Filices. TYPE: Gleichenia

Sm.

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

forming dense thickets, circinate in vernation,

monomorphic, once or several times pseudodi-

chotomously branched (rarely simple or with sev-

eral 2-pinnate pinnae), indeterminate, with a per-

manently arrested bud at the fork of its branches,

or the lamina partially pinnately branched, the

rachis (and sometimes pinna-rachis) with a pe-

riodically dormant bud between the last developed

branches, a pair of accessory foliaceous segments

sometimes present at the base of otherwise usually

naked axes, in addition to stipule-like segments

borne within the forks. Lamina bearing scales and/

or trichomes. Veins free. Penultimate segments

pinnatisect (rarely pinnate) pectinate. Sporangia

borne in exindusiate sori on the abaxial surface of

ultimate segments, with a short many-rowed stalk

of cells and a central to oblique or nearly apical

annulus not interrupted by the stalk. Spores

monolete or trilete, lacking chlorophyll, 1 20 to ca.

800 in each sporangium.

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

and species with larger leaves sometimes spread

over low shrubs to form dense thickets. Approx-

imately 1 20 species have been recognized by var-

ious authors in as many as eight genera. The fol-

lowing treatment is based generally on the

classification of Holttum (1957). Two genera are

recognized in Peru.

References

HOLTTUM, R. E. 1957. Florae Malesianae Prae-

cursores XVI. On the taxonomic subdivision of

the Gleicheniaceae... . Reinwardtia, 4: 257-280.

MAXON, W. R. 1909. Gleicheniaceae, in N. Amer.

fl., 16: 53-63.

NAKAI, T. 1950. A new classification of Glei-

cheniales. Bull. Natl. Sci. Mus., 29: 1-71.

UNDERWOOD, L. M. 1907. American ferns VIII.

A preliminary review of the North American

Gleicheniaceae. Bull. Torrey Bot. Club, 34: 243-

262.

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

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

419. 1793. TYPE: G. polypodioides (L.) Sm.

(Onoclea polypodioides L.). Figure 9.

Sticherus Presl, Tent, pterid. 51. 1836. TYPE: S. lae-

vigatus (Willd.) Presl (Mertensia laevigata Willd.)

= Gleichenia truncata (Willd.) Sprengel.

Diplopterygium (Diels) Nakai, Bull. Natl. Sci. Mus.

29: 47. 1950. Gleichenia section Diplopterygium

Diels, Nat. Pflanz. 1(4): 353. 1900. TYPE: Glei-

chenia glauca (Houtt.) Hooker (Polypodium

glaucum Houtt.).

Stem provided with setose or short-ciliate scales,

and sometimes stellate trichomes. Leaves once or

several times pseudodichotomously branched (but

simple in G. simplex, or with several 2-pinnate

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 to abundant on costae and veins abaxially 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.

TYPE: Jamaica, Bancroft (holotype, K).

Mertensia bancroftii (Hooker) Kunze, Linnaea 18: 307.

1844.

Dicranopteris bancroftii (Hooker) Underw., Bull. Tor-

rey Bot. Club 34: 252. 1907.

Hicriopteris bancroftii (Hooker) Ching, Sunyatsenia 5:

278. 1940.

Diplopterygium bancroftii (Hooker) A. R. Smith, Amer.

Fern J. 70: 26. 1980.

Leaves with 1-3 pairs of 2-pinnate pinnae, these

to 1.5 m long and 40 cm broad, the axes naked or

with scattered, linear, or filiform scales. Axillary

scales lanceolate to ovate, long-acuminate to at-

tenuate, yellowish to light brown, with entire mar-

gins. Penultimate segments (pinnules) very nu-

merous, crowded, at nearly right angles to the costa,

pectinate, cut nearly or quite to the costa. Ultimate

segments strongly revolute, midribs abaxially often

provided with scattered, filiform scales. Sori in-

framedial.

In open forests, commonly on slopes or ravine

banks, 2400-2800 m, infrequent in Peru: Huan-

uco, Pasco, Cuzco.

Mexico to Panama; West Indies; Colombia;

Venezuela; Ecuador; Peru; Bolivia.

Huanuco: Huanuco, within 5 km of Carpish, Tryon &

Tryon 5316 (F, OH, uc, us). Pasco: Border Prov. Oxa-

pampa and Pasco, van der Werffet al. 8604 (MO). Cuzco:

La Convencion, Valle de San Miguel, Biies 2126 (us).

2. Gleichenia simplex (Desv.) Hooker, Icon, pi.,

/. 92. 1837.

Mertensia simplex Desv., Dictionaire sciences natu-

relles 62(2), /. 91. 1827; Mem. Soc. Linn. Paris

6: 200. 1 827. TYPE: Peru, collector unknown (ho-

lotype, P; photo, us).

Dicranopteris simplex (Desv.) Maxon, Contr. U.S. Natl.

Herb. 24: 50. 1922.

Sticherus simplex (Desv.) Ching, Sunyatsenia 5: 285.

1940.

Leaves simple (very rarely with a single fork),

deeply pinnatisect to pinnate, linear, 20—50 cm

long and less than 3 cm broad. Scales of the petiole

essentially lacking, but abundant on the rachis

abaxially, these lanceolate, attenuate, orange to

reddish brown, the margins long-ciliate. Ultimate

segments contiguous at the very base or (especially

proximal ones) discrete and almost their own width

apart, strongly revolute, on the abaxial side slightly

to conspicuously pruinose and scaly, scales along

the midrib similar to those of the rachis, but to-

ward the segment apex grading to pluricellular,

stellate trichomes. Sori medial to inframedial.

Road cuts, banks and open rocky slopes,

(1900-)2700-3800 m, Piura to Cajamarca, south

to Huancavelica and Cuzco.

Colombia to Bolivia.

There has been some confusion as to publication

of the name Mertensia simplex. In the Diet. Sci.

Nat. 1 827 there is no text, but the plate with anal-

ysis constitutes a valid publication. A description

was given at approximately the same time by Des-

vaux in Mem. Soc. Linn. Paris 6: 200. 1827.

Piura: Prov. Huancabamba, road to Canchaque,

Hutchison 1617 (F, GH, NY, uc). Lambayeque: Dist. In-

cahuasi, Laguna Tembladera, Sagdsteguiet al. 12774 (F,

HUT, MO). Cajamarca: Prov. San Miguel, El Tingo, Sa-

gdstegui et al. 9517 (F, MO, NY). Amazonas: Prov. Chach-

apoyas, banks above swamp on summit of Cerros de

Calla-Calla, Wurdack 1212 (F, GH, NY, us). La Libertad:

Prov. Pataz, Huaylillas, Puerta del Monte, Lopez & Sa-

gdstegui 3457 (GH). San Martin: Dist. Huallaga, Valley

of Rio Apisoncho, Hamilton & Holligan 540 (us). An-

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).

Huancavelica: Prov. Tayacaja, Chuspi-Tocas, between

Colcabamba and Paucarbamba, Tovar 2066 (GH, USM).

Cuzco: La Convencion, Biies 2078 (GH, us).

3. Gleichenia pennigera (Mart.) Moore, Index fil.

381. 1862.

Mertensia pennigera Mart., Icon. pi. crypt. 130, t. 59,

f. 1. 1834. TYPE: Brazil, "Prov. Minarum, in

Serra de S. Geraldo," Martius (M; photo, us).

Dicranopteris pennigera (Mart.) Maxon, Contr. U.S.

Natl. Herb. 24: 48. 1922.

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

(2-)3— 4-forked, the branches (at least in Peruvian

specimens) strongly ascending, often tightly

crowded and subparallel with adjacent ones. Ax-

illary scales lanceolate, orange, the margins co-

piously beset with long, tortuous pale or whitish

cilia. Larger penultimate segments 1.4-2.2 cm

broad, the costae abaxially provided with a mix-

ture of orange or whitish long-ciliate scales and a

scattered whitish tomentum, adaxially naked or

sparsely to amply white-tomentose. Ultimate seg-

ments slightly to strongly revolute, larger ones to

1.2 cm long, densely covered abaxially with whit-

ish tomentum (often thinning in age). Sori infra-

medial, mostly crowding or touching the midrib.

Open places in forests, on slopes or stream banks,

1000-2100 m, Loreto?, Huanuco, Pasco, Junin,

Cuzco.

Colombia; Venezuela; Peru; Bolivia; Brazil.

Special note should be made of the branching

pattern throughout the range of this species.

Branches may be very strongly ascending, thus

quite crowded and even subparallel with each oth-

er; or they may diverge at broader angles, as seen

in the type. Thus far, all specimens examined from

Peru exhibit the crowded pattern, which might

tempt one to consider separation at subspecific

level. However, this difference appears uncorre-

lated with other characters, and specimens, at least

outside Peru, seem to grade freely from one pattern

to the other.

Loreto?: " Altura de Salarayacu" (Sarayacu?), Biies 847

(us). Huanuco: Between Huanuco and Pampayacu, Ka-

nehira 153 (GH, us). Pasco: Prov. Oxapampa, Gran Pa-

jonal, D. Smith 5073 (F, MO). Junin: Concepcion, Her-

rera 142 (us). Cuzco: Valle de Lares, Hda. Huy-huy, Biies

1826 (us). Prov. Convention, Lucumayo, Vargas 4221

(us).

4. Gleichenia bifida (Willd.) Sprengel, Syst. veg.

4: 27. 1827. Figure 9.

Mertensia bifida Willd., Kongl. Vetensk. Acad. Nya

Handl. 25: 168. 1804. TYPE: Venezuela, Czn-

cas,Bredemeyer(ho\olype,B,Herb. Willd. 19468;

photo, GH).

Dicranopteris bifida (Willd.) Maxon, N. Amer. fl. 16:

60. 1909.

Gleichenia mathewsii Hooker, Sp. fil. 1 : 9. 1 844. TYPE:

Peru, Mathews 1092, in part (holotype, K; photo,

us).

Mertensia mathewsii (Hooker) Fee, Mem. Foug. 1 1:

122. 1866.

Sticherus bifidus (Willd.) Ching, Sunyatsenia 5: 282.

1940.

Sticherus mathewsii (Hooker) Nakai, Bull. Nat. Sci.

Mus. 29: 22. 1950.

Leaves pseudodichotomously forked. Pinnae 1-

or 2-forked, the branches spreading to moderately

ascending. Axillary scales lanceolate to ovate, at-

tenuate, light brown to pale orange, the margins

amply ciliate. Larger penultimate segments (2.5-)3-

7 cm broad, the costa on the abaxial side amply

to abundantly provided with orange, long-ciliate

scales, on the adaxial naked or sometimes with

sparse whitish tomentum. Ultimate segments

moderately to (typically) strongly revolute, larger

ones 1 5-35 mm long and 1 .8-3 mm broad (beyond

the dilated base), densely covered abaxially with

orange to whitish tomentum (sometimes thinning

in age), the midribs also sparsely provided with

tawny to whitish, filiform, ciliate scales. Sori in-

framedial, mostly crowding or touching the mid-

rib.

In forests, thickets and clearings, often on banks

and slopes, 150-1800 m (very rarely to 2200 m)

Piura to Loreto, south to Madre de Dios and Puno.

Mexico to Panama; West Indies; Colombia and

Venezuela south to Bolivia and Paraguay.

Piura: Prov. Huancabamba, slopes of Cerro La Viuda,

Sagdsteguiet al. 8 209 p.p. (NY). Antazonas: Prov. Bagua,

along roadside, 37 km NE of Chiriaco, Barbour 4486

(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,

us). Huanuco: Dist. Churubamba, trail Cotirarda to

Mercedes, Mexia 8191 (F, GH, MICH, MO, NY, uc, us).

Pasco: Prov. Oxapampa; Palcazii, Rio Alto Iscozacin,

Foster & d'Achille 10098 (F). Junin: Huacapistana, be-

tween Tarma & San Ramon, Ferreyra 286 (GH, USM).

Ucayali: Vicinity of Aguaytia, on steep slopes along Rio

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.

Paucartambo, near Asuncion, West 7122 (MICH, uc).

Madre de Dios: Prov. Manii, Rio Inambari, Chavez 1057

(MO). Puno: San Gaban to Ollachea, Dillon et al. 1243

(F, MO, NY).

5. Gleichenia tomentosa (Sw.) Sprengel, Syst. veg.

4: 27. 1827.

Mertensia tomentosa Sw., Kongl. Vetensk. Acad. Nya.

Handl. 25: 1 77, /. 5J. 4. 1 804. TYPE: Peru, Herb.

Cavanilles (holotype, s; isotype, p?; photos, us of

S&P).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

41

Mertensia velata Kunze, Linnaea 9: 15. 1834. TYPE:

Peru, Huanuco, Pampayacu, July 1829, Poeppig

(diar. 1117) (holotype, B?; isotype, M!; probable

isotype, P!; frag., us!; photos, us of M & p).

Gleichenia velata (Kunze) Mett., Fil. hort. bot. Lips.

113. 1856.

Gleichenia buchtienii Christ & Rosenst., Repert. Spec.

Nov. Regni Veg. 5: 229. 1908. TYPE: Bolivia,

"Yungas australis, Sirupaya prope Yanacachi,"

Buchtien 496 (holotype, P!; isotypes, F!, P!, s).

Dicranopteris velata (Kunze) Maxon, Contr. U.S. Natl.

Herb. 24: 50. 1922.

Sticherus buchtienii (Christ & Rosenst.) Copel., Gen.

fil. 28. 1947.

Sticherus velatus (Kunze) Copel., Gen. fil. 28. 1947.

Leaves pseudodichotomously forked. Pinnae 1 -

or 2-forked, the branches spreading to moderately

ascending. Axillary scales lanceolate to ovate, at-

tenuate, deep to pale orange, or castaneous at base,

the margins amply long-ciliate. Larger penultimate

segments 4-7 cm broad, the costa abundantly pro-

vided abaxially with orange, long-ciliate scales, and

adaxially with orange to whitish filiform scales

which often grade into a pale tomentum (this

sometimes caducous). Ultimate segments plane to

moderately revolute, larger ones 20—45 mm long

and (3-)3.5-4.5 mm broad above the dilated base,

densely covered abaxially with orange to whitish

tomentum (this sometimes thinning in age), the

midribs provided also with orange, ciliate scales.

Sori mostly medial, rarely, if ever, crowding the

midrib.

In cloud or elfin forests, usually in open sites on

ridges, slopes or ravine banks, 1 500-2700 m, Ca-

jamarca, Amazonas, San Martin, Huanuco, Pasco,

Cuzco.

Colombia; Venezuela; Ecuador; Peru; Bolivia.

Much confusion has attended circumscription

of the densely tomentose species of Gleichenia,

which include G. bifida as well as those combined

here under G. tomentosa. We have been unable to

obtain the type specimens of the latter, but type

photos (us) indicate clearly enough that ultimate

segments are too broad and soral lines appear too

far from midribs to be G. bifida. The photos, along

with Swartz's original description ( 1 804) of Mer-

tensia tomentosa and his subsequent Latin de-

scription (Syn. fil. 164, p. 392, 1806) make it ev-

ident that G. tomentosa and G. velata are the same.

All this confusion within the species complex is

warranted, for G. bifida and G. tomentosa do not

differ strongly. As seen in the key, the former has

narrower, more strongly revolute ultimate seg-

ments, and soral lines usually are closer to the

midrib; also the plants seem to occur at lower

elevations in Peru: 150-1800 m (rarely to 2200

m) as compared with 1 500-2700 m in G. tomen-

tosa. Color of scales on the buds and major axes

affords another, but less consistent, diagnostic fea-

ture; in both species these are pale to deep orange,

but in G. tomentosa they are often liberally inter-

mixed with castaneous ones. A few specimens have

been seen which are intermediate, thus emphasiz-

ing the close relationship of the two species. For

example, the only collection of G. tomentosa (cited

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.

Cajamarca: Prov. Cutervo, La Achira (San Andres-

Socota), growing over bushes, Lopez & Sagdstegui 5464

(GH, HUT). Amazonas: Prov. Bagua, Cordillera Colan SE

of La Peca, Barbour 3980 (MO). San Martin: "In monte

Campana prope Tarapoto," Spruce 4707 (p, us). Huan-

uco: Playapampa, steep banks, Macbride 4509 (F, us).

Pasco: Cushi, trail to Tambo de Vaca, Bryan 687 (F).

Cuzco: La Convention, ca. 12 km NE from Hacienda

Luisiana and Rio Apurimac, Dudley 10545 (GH, us).

6. Gleichenia revoluta HBK., Nov. gen. sp. 1: 29.

1815. TYPE: Ecuador, Saraguru, Humboldt

& Bonpland (holotype, p; isotype, B!; frag., us

of B; photos, us of B & p).

Mertensia revoluta (HBK.) Desv., Mem. Soc. Linn.

Paris 6: 200. 1827.

Mertensia pruinosa Mart., Icon. pi. crypt. 109. 1834.

TYPE: Brazil, Minas Gerais, Freyreiss (probable

holotype, M!; isotype, s; photos, F & us of M).

Gleichenia pruinosa (Mart.) Mett., Ann. Mus. Bot.

Lugduno-Batavum 1: 49. 1863.

Gleichenia affinis Kuhn, Linnaea 36: 167. 1869. TYPE:

Peru, Dept. Puno, San Gaban, Lechler 2265 (ho-

lotype, B!; photo, us).

Dicranopteris affinis (Kuhn) Maxon, Contr. U.S. Natl.

Herb. 24: 47. 1922.

Dicranopteris pruinosa (Mart.) Maxon, Contr. U.S.

Natl. Herb. 24: 49. 1922.

Sticherus pruinosus (Mart.) Ching, Sunyatsenia 5: 284.

1940.

Sticherus revolutus (HBK.) Ching, Sunyatsenia 5: 285.

1940.

Sticherus affinis (Kuhn) Nakai, Bull. Natl. Sci. Mus.

29: 13. 1950.

Leaves pseudodichotomously forked. Pinnae 2-

4-forked. Axillary scales lanceolate to ovate, ±lax,

thin-textured, orange to reddish brown, the mar-

gins amply ciliate. Larger penultimate segments I -

1.8(-2) cm broad, the costa on the abaxial side

amply to copiously provided with orange to red-

dish brown, long-ciliate scales, on the adaxial side

42

FIELDIANA: BOTANY

the costa naked or filiform-scaly. Ultimate seg-

ments moderately to strongly revolute, larger ones

2-3.5(-4) times longer than broad, the midrib and

veins scaly as on the costa, those of the veins often

filiform or grading to tortuous, stellate trichomes,

the veins not or scarcely raised. Sori medial to

supramedial.

In elfin forests, clearings, and along roadsides,

1 770-3400 m, Piura to Amazonas, south to Cuzco

and Puno.

Costa Rica & Panama; Colombia; Venezuela;

Ecuador; Peru; Brazil; Bolivia.

This species is locally abundant in middle to

upper elevations and is sometimes confused with

Gleichenia bifida, probably because of the often

copious indument borne on the abaxial surfaces.

These scales, especially on the veins, often become

filiform and even grade into stellate trichomes;

however, the surfaces are never truly tomentose,

as in G. bifida. Also, penultimate segments of the

latter are much broader, with linear ultimate seg-

ments, and the sori crowd the midrib, whereas in

G. revoluta sori are medial to supramedial, most

of them quite remote from the midrib. Further-

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)

Lell.]. Probably G. subandina Sodiro and G. hy-

poleuca Sodiro of Ecuador are also synonymous

with G. revoluta.

Piura: Prov. Huancabamba, above Huancabamba, road

to Canchaque, Hutchinson 1618 (F, OH, uc, us). Caja-

marca: Prov. Cutervo, alrededores de Cutervo, Ldpez &

Sagdstegui 5314 (GH, MO). Amazonas: Prov. Chacha-

poyas, slopes of Puma-urcu SE of Chachapoyas, Wur-

dack 552 (F, GH, NY). La Libertad: Prov. Pataz, Puerta

del Monte, Paso La Sabana, Lopez & Sagdstegui 3463

(GH). Pasco: Prov. Oxapampa, San Alberto, Cordillera

de Yanachaga, van der Werffet al. 8477 (MO). Huanuco:

Playapampa, in sphagnum montana, Macbride 4508 (F,

GH, NY, us). Cuzco: Cordillera Vilcabamba, 23 km NE

from Hacienda Luisiana, Dudley 11133 (GH, us). Puno:

Prov. Sandia, near Limbani, Metcalf 30539 (MO, us;

atypical in narrower segments and darker, narrower

scales).

7. Gleichenia tuberculata Kuhn, Linnaea 36: 166.

1869. TYPE: Peru, Dept. Puno, Tatanara,

Lechler2572 (holotype, B!; isotype, B!; photos,

F & MICH of B).

Sticherus tuberculatus (Kuhn) Nakai, Bull. Natl. Sci.

Mus. 29: 30. 1950.

Leaves pseudodichotomously forked, petiole and

rachis conspicuously tuberculate to muricate. Pin-

nae 2-3-forked. Axillary scales closely imbricate,

ovate to lanceolate, castaneous to reddish brown,

the margins amply ciliate. Larger penultimate seg-

ments 1 .3-1.5 cm broad, pruinose abaxially, scales

essentially lacking. Ultimate segments moderately

revolute, larger ones ca. 3 times longer than broad.

Sori supramedial.

Thus far represented only by the type collection,

from Puno. This species appears to differ from

Gleichenia revoluta only in the lack of scales on

the laminar surface and in the tuberculate to mur-

icate primary axis. However, the specimens were

taken from a plant apparently quite advanced in

age, for few sporangia remain on the segments,

and it is possible that any scales once borne on

the axes may have fallen away as well. Density of

indument varies within some species of Gleichen-

ia, depending upon maturity. Furthermore, tu-

berculate primary axes are not uncommon in the

genus, although never so conspicuously so as in

G. tuberculata. The species is maintained as dis-

tinct here, but with reservations.

8. Gleichenia nitidula Rosenst., Repert. Spec. Nov.

Regni Veg. 10: 275. 1912. TYPE: Costa Rica,

San Carlos, Brade & Brade 503 (holotype, not

located; isotypes, NY, s).

Sticherus nitidulus (Rosenst.) Copel., Gen. fil. 28. 1 947.

Leaves pseudodichotomously forked, petiole and

rachis not or scarcely tuberculate. Pinnae 1-3-

forked. Axillary scales not or slightly imbricate,

linear or linear-lanceolate, with subcaudate tips,

deep reddish brown, the margins entire or (rarely)

sparingly dentate or short-setose. Larger penulti-

mate segments 1.4-1.8 cm broad, not or scarcely

pruinose abaxially, scales lacking. Ultimate seg-

ments slightly revolute, larger ones ca. 3 times

longer than broad.

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

1900 m, San Martin, Pasco.

Costa Rica; Panama; Ecuador, Peru.

The species is notable for its disjunct distribu-

tion, and has rarely been collected in South Amer-

ica. Distinctive characteristics are the lack of scales

on the narrow penultimate segments, and the

sparse, rigid axillary scales with entire margins.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

Apparently represented thus far in Peru by the follow-

ing two collections: San Martin: "Prope Tarapoto, in

monte Guayrapurima," Spruce 4018 (K). Pasco: (as Jun-

in) Pichis Trail, Enenas, dense forest, Killip & Smith

25777 (F).

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).

Gleichenia yungensis Rosenst., Repert. Spec. Nov.

Regni Veg. 5: 228. 1908. TYPE: Bolivia, Yungas,

Unduavi, 3300 m, Buchtien 902 (holotype, s; is-

otypes, uc!, us!).

Dicranopteris yungensis (Rosenst.) Maxon, Contr. U.S.

Natl. Herb. 24: 50. 1922.

Sticherus yungensis (Rosenst.) Copel., Gen. fil. 28.

1947.

Sticherus lechleri (Kuhn) Nakai, Bull. Natl. Sci. Mus.

29: 21. 1950.

Leaves pseudodichotomously forked, with fo-

liaceous, often pinnatifid, appendages borne with-

in each axil (especially proximally). Pinnae 1-3-

forked. Axillary scales ovate to linear-lanceolate,

orange to reddish brown, attenuate, the margins

short-ciliate. Penultimate segments (larger ones)

2.5-6 cm broad, costae sparsely to amply scaly

abaxially, the scales orange, linear or linear-lan-

ceolate, long-attenuate, the margins somewhat

short-ciliate. Ultimate segments plane or slightly

revolute, commonly pruinose abaxially, larger ones

12-28 mm long and 2.5-4.5 mm broad, the mid-

ribs lacking scales. Sori commonly supramedial.

In mountain forests, commonly on exposed

ridges, (1 1 00-) 1 800-3000 m, Huanuco, Cuzco.

Trinidad; Colombia; Ecuador; Peru; Bolivia.

This species is characterized by lack of midrib

scales, commonly pruinose lamina, plane or slight-

ly revolute ultimate segments, and the suprame-

dial sori. With it should be included Dicranopteris

brittonii Maxon of Trinidad, and probably also G.

leucocarpa Sod. of Ecuador. Although we have not

seen the type of the latter, a type fragment and

photo (us) reveal no significant differences be-

tween the two species.

Huanuco: Carpish, Coronado 78 (us). Cuzco: Machu

Picchu, Valle del Urubamba, Herrera 3286 (us). Cerro

de Cusilluyoc, forest along Rio Pilahuata, Pennell 13940

(F, us). Prov. Paucartambo, "Pillawata," Yanamayo-

Tambomayo, Vargas 16702, 16705 (GH).

10. Gleichenia remota (Kaulf.) Sprengel, Syst. veg.

4: 27. 1927.

Mertensia remota Kaulf., Enum. fil. 39. 1824. TYPE:

Brazil, Ilha de Sta. Catarina, Chamisso (holotype,

LZ destroyed; isotype, LE?).

Dicranopteris remota (Kaulf.) Maxon, Contr. U.S. Natl.

Herb. 24: 50. 1922.

Leaves pseudodichotomously forked, with fo-

liaceous appendages usually borne within each axil

(especially proximally). Pinnae 1 - or 2-forked. Ax-

illary scales lanceolate to ovate, attenuate, rigid,

1-1.5 mm long, often somewhat convex, casta-

neous to reddish brown, lustrous, not or slightly

imbricate, their margins short-setose, or rarely en-

tire or with a few short cilia at base. Penultimate

segments (larger ones) 4-9 cm broad, costae abax-

ially provided with scattered castaneous or dark

reddish brown scales less than 1 mm long, these

ovate to lanceolate, with setose to short-ciliate

margins, adaxially virtually naked (or occasionally

with scattered patches of arachnoid scurf). Ulti-

mate segments remote, most of them separated by

a space once or twice their width, slightly to strong-

ly revolute, not or slightly pruinose abaxially, larg-

er ones 20-50 mm long and 1.7-2.5 mm broad,

the midribs amply to copiously scaly on abaxial

side, the scales deltoid, orange to tawny, with long,

pale cilia on the margins, or toward segment tips

sometimes becoming substellate. Veins often

raised. Sori medial to supramedial.

At edges of forests, roadsides, and on rocky

banks, 350-2000 m, Amazonas, San Martin,

Huanuco.

Costa Rica; Panama; Cuba; Trinidad; Colombia

to the Guianas; Ecuador; Peru; Bolivia; Brazil

(Amazonas, Ilha Sta. Catarina).

This and Gleichenia longipinnata are superfi-

cially similar in the great size of their penultimate

segments (sometimes to 50 cm long and 9 cm

broad) and the extremely long and narrow ultimate

segments. However, besides the characters noted

in the key, G. remota can also be sharply distin-

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

1 mm) and broader. In G. longipinnata, both the

axillary scales and costal scales are linear to fili-

form, the former 2-3 mm long and the latter 1.5-

2 mm long.

44

FIELDIANA: BOTANY

Ama/onas: Prov. Bagua, roadside, 37 km NE of Chi-

riaco, Barbour 4485 (MO), 4486 (USM). San Martin: Tar-

apoto-Yurimaguas Hwy., km 39, McDaniel 14209 (GH,

MO). Tarapoto, Carretera Tarapoto-Yurimaguas, Ri-

machi 5163, 5251 (MO, NY). Huanuco: Monzon, con-

fluencia con Huallaga, cerca de Tingo Maria, Ferreyra

10047 (GH).

11. Gleichenia peruviana (Maxon) 1 ell.. Amer.

Fern J. 74: 57. 1984.

Dicranopteris peruviana Maxon, Amer. Fern J. 33:

133. 1943. TYPE: Peru, Huanuco, Playapampa,

ca. 2700 m, Macbride 4510 (holotype, F!; isotype,

us!).

Leaves pseudodichotomously forked, with fo-

liaceous appendages often borne within some of

the more proximal axils. Pinnae 1-3-forked. Ax-

illary scales ovate or lance-ovate, attenuate, 0.5-

1.5 mm long, castaneous to reddish brown, closely

imbricate, their margins amply ciliate, the cilia lax,

short and whitish to pale orange. Penultimate seg-

ments (larger ones) 2.5^4(-5) cm broad, costae on

abaxial side amply to copiously scaly, the scales

lustrous castaneous to reddish brown, ovate to lan-

ceolate, acuminate or attenuate, commonly less

than 1 mm long, their margins short-ciliate, costae

on adaxial side sparsely to amply provided with

tortuous trichomes or filiform scales. Ultimate seg-

ments crowded, contiguous at their bases, slightly

to moderately revolute, not or slightly pruinose,

larger ones 1 2-24 mm long and 1 .5-2.5 mm broad

(beyond the dilated base), abaxially the midribs

and (usually) veins minutely scaly, ciliate- or se-

tose-scaly, the scales often becoming filiform or

grading into stellate, whitish trichomes. Veins often

raised. Sori mostly inframedial.

In thickets, open forests and ravines, often on

exposed rocky ridges, 2000-3400 m, La Libertad,

Huanuco, Pasco, Cuzco.

Apparently confined to Peru, Cordilleras Cen-

tral and Oriental.

La Libertad: Prov. Pataz, Pampa de Huayno-huincho,

Huaylillas, Ldpez & Sagdstegui 3517 (GH, HUT). Huan-

uco: Wet, dense jungle, 12 mi S of Panao, Macbride &

Featherstone 2217 (F). Pasco: Prov. Oxapampa, Dist.

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

10710 (GH, MO).

12. Gleichenia longipinnata Hooker, Sp. til. 1: 9.

1844. TYPE: Surinam, Hostmann 238 (ho-

lotype, K; isotypes, BM, K; frag., us!; photo, us

ofK).

Mertensia longipinnata (Hooker) Klotzsch, Linnaea

18: 537. 1844.

Dicranopteris longipinnata (Hooker) Maxon, Contr.

U.S. Nail. Herb. 24:48. 1922.

Sticherus longipinnatus (Hooker) Nakai, Bull. Nat. Sci.

Mus. 29: 21. 1950.

Leaves pseudodichotomously forked, occasion-

ally bearing foliaceous appendages within the ax-

ils. Pinnae commonly 1 -forked. Axillary scales

linear to filiform, 2-3 mm long, lustrous reddish

brown to deep orange, not or rarely tightly im-

bricate, the margins with dark, rigid, ascending

setae. Penultimate segments (larger ones) 4-9 cm

broad, costae on abaxial side amply to abundantly

scaly, the scales 1.5-2 mm long, lustrous reddish

brown, setose and filiform or substellate, on adax-

ial side often appressed filiform-scaly. Ultimate

segments approximate, contiguous at their bases

(very rarely some proximal ones discrete and

somewhat apart), plane to slightly revolute, not or

slightly pruinose, larger ones 25-45 mm long and

2.5-3.5 mm broad, abaxially bearing filiform or

substellate scales on midrib and veins and often

some pale, delicate trichomes at the very margin.

Veins not or slightly prominulous. Sori mostly

raised.

In forests, on slopes and ridges, 300-1600 m

(reported on Dudley 10448 label as also seen at

3300 m, but probably mistaken for G. peruviana),

Amazonas, Loreto, Pasco, Huanuco, Cuzco.

Surinam; Peru; Brazil.

Despite obvious differences, this and Gleichenia

rubiginosa share a distinctive type of laminar scale

not found in other species in Peru. The scales along

the costae of penultimate divisions, abaxially, are

typically long and narrow, the marginal cell walls

deep reddish brown and bearing rigid dark setae.

Along midrib and veins, these grade into filiform

scales often only two to three cells wide (the setae

then much longer than the width of the scale body)

and thence into delicate, pluricellular trichomes.

The Costa Rican G. mellifera Christ also bears this

kind of laminar scale, and although we have not

seen the type, it probably belongs here.

Amazonas: Prov. Bagua, valley of Rio Maranon above

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

Abad, Chacra de Cesar Vela, J. Schunke 5465, (F, GH,

MO, NY, us). Pasco: Prov. Oxapampa, Valle de Palcazii,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

45

Foster 4504 (F). Cuzco: Prov. Convention, Cordillera

Vilcabamba, Dudley 10448 (GH).

13. Gleichenia rubiginosa Mett., Ann. Sci. Nat.

Bot. 5, 2: 267. 1864. TYPE: Colombia, Puente

Nacional, 1650 m, Lindig 71 (holotype, B!;

isotype, P; frag., us!).

Gleichenia rubiginosa f. virescens Hieron., Bot. Jahrb.

Syst. 34: 561. 1905. SYNTYPE: Peru, Matthews

1092 (us).

Dicranopteris rubiginosa (Mett.) Maxon, Contr. U.S.

Natl. Herb. 24: 50. 1922.

Sticherus rubiginosus (Mett.) Nakai, Bull. Nat. Sci.

Mus. 29: 28. 1950.

Leaves pseudodichotomously forked, rarely

bearing foliaceous appendages in the axils. Pinnae

1- or 2-forked. Axillary scales linear or linear-

lanceolate, attenuate, 2-3 mm long, sublustrous,

reddish brown, not or loosely imbricate, the mar-

gins with dark, rigid, ascending setae. Penultimate

segments (larger ones) 2.2-3.2 cm broad, costae

on abaxial side copiously scaly, the scales 2-3 mm

long, deep orange, sublustrous, lanceolate or lin-

ear-lanceolate and attenuate, with setose margins,

on adaxial side naked or occasionally sparsely ar-

achnoid-scaly. Ultimate segments approximate,

contiguous at their bases, moderately to strongly

revolute, pruinose or not, larger ones 12-18 mm

long, (2-)2.5-3 mm broad, abaxially bearing fili-

form and setose to substellate scales on midrib

and veins, and sometimes some delicate trichomes

at the very margin. Veins raised abaxially. Sori

inframedial or occasionally medial.

In wet forests and thickets, 1 100-2850 m, Piura,

Huanuco, Pasco, Ucayali, Cuzco, Puno.

Colombia; Venezuela; Ecuador, Peru.

Although easily distinguished from G. longipin-

nata (q.v.), this is apparently closely related to the

latter by virtue of the distinctive laminar scales.

Piura: Prov. Huancabamba, Dist. Sondor, subiendo

al Cerro 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-

pa and Paucartambo, D. Smith 1592 (F, MO). Ucayali:

Prov. Coronel Portillo, Cordillera Azul, km 43 Tingo-

Maria-Pucallpa Road, Young & Sullivan 737 (F, MO).

Cuzco: Valle San Miguel, La Convention, Bites 2061

(us). Puno: Prov. Carabaya, San Gaban Carretera, Var-

gas 18877 (GH).

II. Dicranopteris

Dicranopteris Bernh., Neues J. Bot. 1(2): 38.

("1806") 1805, nom. nov. for Mertensia Willd.

(not Roth), and with same type. Figure 10.

Mertensia Willd., Kongl. Vetensk. Acad. Nya Handl.

25:165.1 804, illeg. (not Roth 1 790). TYPE: Mer-

tensia dichotoma (Murray) Willd. (Polypodium

dichotomum Murray), Dicranopteris dichotoma

(Murray) Bernh. = Dicranopteris linearis (Burm.

f.) Underw.

Stem provided with pluricellular trichomes,

scales lacking. Leaves 1—4 (rarely to 20) m long,

pseudodichotomously branched, with 1 -several

pairs of opposite pinnae which again typically

branch 1 or more times in opposite pairs, or with

several pinnae which branch unequally. Axils of

forks each bearing a dense tuft of trichomes and

often a pair of reduced, stipule-like appendages.

Lamina glabrous, or pubescent with simple or stel-

late trichomes, scales lacking. Veins 2— 4-forked.

Sori lacking paraphyses. Sporangia ca. 8-15 per

sorus.

This, like Gleichenia, is an essentially pantrop-

ical genus. The two are readily distinguished by

the character of their indument: that of Gleichenia

being scales and trichomes, while scales are com-

pletely lacking in Dicranopteris. Three species of

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.,

Bull. Torrey Bot. Club 34: 254. 1907. Figure

lOb-c.

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:

Peru, "Chibangata," Poeppig 1153 (holotype, LZ

destroyed; isotype, p).

Gleichenia flexuosa (Schrader) Men., Ann. Mus. Bot.

Lugduno-Batavum 1: 50. 1863.

Gleichenia rigida (Kunze) Bommer & Christ, Bull.

Soc. Roy. Bot. Belgique 35: 174. 1896, not G.

rigida J. Sm.

Leaves with 1-several opposite pairs of stalked

pinnae, these repeatedly pseudodichotomous, with

all subsequent branches forking in subequal pairs

(i.e., each of the stalks subequal in length and di-

verging at about the same angle), and a pair of

reduced, accessory, pectinate segments commonly

produced at the base of each (but rarely the ulti-

mate) fork. Axils bearing a tuft of stout, rigid,

castaneous trichomes and a pair of reduced sti-

pule-like segments. Pinnae and their branches long-

stalked, the axes and laminar surfaces glabrous,

the forked, penultimate segments sessile, to 30 cm

long and 6 cm broad. Ultimate segments contig-

uous at base, to 30 mm long and 4 mm broad,

coriaceous, glaucous abaxially, the margins strong-

ly revolute. Spores trilete.

On ridges and open slopes, at edges of forests,

750-2750 m, San Martin, Huanuco, Ucayali, Cuz-

co, Puno.

West Indies; southern Mexico to Panama, S to

Brazil, Bolivia, and Paraguay (some isolated col-

lections in the United States from coastal Ala-

bama).

This species is very closely related to Dicran-

opteris linearis (Burm.) Underw. of the Paleotrop-

ics. Suzanne Roth, who is working on a revision

of the Neotropical Gleicheniaceae, has suggested

(in litt.) that the two are probably conspecific. If

so, D. linearis has priority.

San Martin: NW of San Martin, Rioja, Rio Negro,

Soukup 5156 (GH). Huanuco: Vilcabamba, hacienda on

Rio Chinchao, Macbride 5010 (F, NY, us). Ucayali: Prov.

Coronel Portillo, Obeteni, E ridge of basin, Chrostowski

66-13 (uc). Cuzco: Prov. Convention, Choquallo, Var-

gas 8153 (MO, uc). Puno: Prov. Sandia, Asalaya, Vargas

14835 (OH).

2. Dicranopteris nervosa (Kaulf.) Maxon, Contr.

U.S. Natl. Herb. 24: 49. 1922.

Mertensia nervosa Kaulf., Enum. fil. 37. 1824. TYPE:

Brazil, Sta. Catarina Is., Chamisso (holotype, not

located, LZ? destroyed; isotype, LE?, s?).

Gleichenia nervosa (Kaulf.) Sprengel, Syst. veg. 4: 25.

1827.

Leaves 1 -forked, each branch consisting of a sin-

gle, sessile, pinnate (to distally pinnatisect) pinna.

Axils bearing a dense tuft of long, lax, reddish

brown trichomes and usually a pair of reduced

stipule-like appendages. Pinnae (larger ones) 1 5-

40 cm long, 5-12 cm broad, the axes and laminar

surfaces abaxially covered with a dense reddish

brown tomentum (this sometimes gray to whitish

in age). Ultimate segments well-spaced, not con-

tiguous at base, to 60 mm long and 3.5 mm broad,

coriaceous, glaucous abaxially, the margins strong-

ly revolute. Spores trilete.

Thus far represented in Peru by a single collection:

Puno: Region of Rio Inambari, trail from Aricoma Pass

to Santo Domingo, 1800 m, McCarroll 125 (MICH).

Venezuela?; Peru; Bolivia; Brazil.

This is linked to Dicranopteris schomburgkiana

(Sturm) Morton and D. seminuda (KJotzsch) Max-

on by the tomentose abaxial surfaces. The three

are usually separated by size of segments, color of

tomentum, and other quantitative features; how-

ever, D. nervosa appears to be further distin-

guished by the delicate, somewhat flexuous axil-

lary trichomes and the single pair of sessile pinnae.

The related species have either stout, rigid axillary

trichomes, or stalked, branching pinnae, or both.

Whether these characters are sufficient to consti-

tute specific or subspecific distinction is open to

question. Further study of the species complex is

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.

Torrey Hot. Club 34: 260. 1907. Figure lOa.

MertensiapectinataWMd., Kongl. Vetensk. Acad. Nya.

Handl. 25: 168. 1804. TYPE: Venezuela, Cara-

cas, Bredemeyer(hololype,B,Herb. Willd. 19465;

photo, GH).

Gleichenia pectinata (Willd.) Presl, Reliq. haenk. 1 :

71. 1825.

Leaves with 1-several opposite pairs of stalked

pinnae, these each bearing several to many un-

equal branches which are again unequally branched

or which terminate in a pair of penultimate seg-

ments (i.e., each of the stalks unequal in length

and diverging at much different angles), lacking

accessory leafy segments at the base of the forks.

Axils bearing a tuft of orange to reddish brown,

curved to flexuous trichomes and often a pair of

reduced stipule-like segments (at least within the

proximal fork). Pinnae and their branches long-

stalked, the axes commonly glabrous, the midribs

and veins glabrous or, more often, sparsely to am-

ply provided on the abaxial side with orange, stel-

late trichomes, the forked, penultimate segments

sessile, to 23 cm long and 5 cm broad. Ultimate

segments contiguous at base, to 25 mm long and

5 mm broad, chartaceous, glaucous abaxially, the

margins scarcely to moderately revolute. Spores

monolete.

Forest clearings and thickets, along roadsides

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

the veins; but in South America, especially in Peru,

some pubescence is to be expected— varying from

sparse to copious— and on a few specimens we

have examined it is so dense as to nearly obscure

the sori. It was perhaps one of these tomentose

plants on which Hieronymus based his Gleichenia

(Dicranopteris) flexuosa f. monstrosa (see discus-

sion below under Comments).

Amazonas: Prov. Bagua, along roadside from Chiriaco

to Puente Venezuela, Barbour 4488 (MO). San Martin:

Prov. Lamas, Dist. Lamas, Belshaw 3435 (F, GH, MICH,

MO, NY, uc, us). Loreto: Mishuyacu, near Iquitos, Klug

850 (F, NY, us). Huanuco: Prov. Huanuco, Dist. Chu-

rubamba, Pampa Hermosa, Mexia 8147 (F, GH, MICH,

MO, NY, uc, us). Pasco: Prov. Oxapampa, road between

Puente Paucartambo and Oxapampa, D. Smith 1472 (F,

MO). Junin: La Merced, thickets, Killip & Smith 23804

(F, GH, NY, us). Ucayali: Prov. Coronel Portillo, road

Tingo Maria to Pucallpa, Ridoutt s.n. (GH). Cuzco: Prov.

Paucartambo, near Asuncion, West 7121 (MICH p.p., MO,

vcp.p.).

Comments

Dicranopteris flexuosa f. monstrosa (Hieron.) Na-

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

Gleichenia flexuosa f. monstrosa Hieron. Hedwigia

48: 289. 1909. TYPE: Peru, Amazonas, Quebra-

da de Santa Lucia near Chachapoyas, Stiibel 1070

(holotype, not located).

This supposed form of Dicranopteris flexuosa

was described from a plant which was rusty-to-

mentose on the juvenile segments of more distal

pinnae. We have not seen evidence of tomentum

on D. flexuosa, either on mature or juvenile plants,

but it is possible this could represent a form of the

species, or perhaps a hybrid. It is more likely that

it belongs to the complex of tomentose species

which are discussed above in the treatment of D.

nervosa, particularly D. seminuda of Venezuela,

which has the general aspect of D. flexuosa. Another

likely possibility is that it may be a densely pu-

bescent variant of D. pectinata (q.v.), although Hi-

eronymus certainly should have mentioned the

distinctive branching pattern of the latter if this

were the case. Pending examination of the type, it

is fruitless to conjecture further.

Family 6: HYMENOPHYLLACEAE

Hymenophyllaceae Link, Handbuch 3: 36. 1833.

TYPE: Hymenophyllum Sm.

Stem erect to decumbent, often slender and long-

creeping, usually bearing scattered, short tri-

chomes. Leaves commonly circinate in vernation,

entire or pinnatifid to decompound, typically

monomorphic, or rarely dimorphic in Trichom-

anes, glabrous or pubescent, lamina very thin (often

1 cell thick), lacking stomates. Veins free, or in a

few species reticulate. Sporangia borne in margin-

al sori on a short to elongate receptacle, enclosed

within a bivalvate or tubular indusium, with a

short stalk and an oblique annulus not interrupted

by the stalk. Spores trilete, tetrahedral-globose to

spheroidal, commonly with chlorophyll.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

49

The Hymenophyllaceae are a natural and dis-

tinctive family, with representatives found in trop-

ical to wet temperate regions nearly throughout

the world. Most of the nearly 600 species have the

lamina only one cell thick, which has earned them

the common name "filmy fern." The filmy leaves

and the marginal sori shaped like a cup or pouch

make them readily recognizable as a family, but

through the years there have been great differences

of opinion as to its subdivision. Various authors

have split it into as many as 42 genera, whereas

but two are recognized here.

The term marginate is used in this family to

describe an axis of the leaf that has a very narrow

band of green tissue on each side. Frequently this

band is scarcely raised beyond the surface of the

axis and is hardly discernible except for the color

contrast with the axis. It is a rudimentary wing

and represents the ultimate reduction of the con-

ditions elsewhere described in the text as "broadly

alate" or "narrowly alate."

The publications of Presl on the family have

been cited according to the first publication, with

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.

References

COPELAND, E. B. 1938. Genera Hymenophylla-

cearum. Philipp. J. Sci., 67: 1-1 10.

DIEM, J., AND J. LICHTENSTEIN. 1959. Las Hy-

menofilaceas del area Argentina-Chilena del sud.

Darwiniana, 11: 61 1-760.

LELLINGER, D. B. 1984. Hymenophyllaceae, in

Botany of the Guayana Highland— Part X. Mem.

New York Bot. Gard., 38: 9-38.

MORTON, C. V. 1 968. The genera, subgenera and

sections of the Hymenophyllaceae. Contr. U.S.

Natl. Herb., 38: 153-214.

PRANTL, K. 1875. Die Hymenophyllaceen. Un-

ters. Morph. Gefasskrypt., 1: 1-73.

PRESL, K. B. 1843. Hymenophyllaceae, pp. 1-

70 (from Abh. Konigl. Bohm. Ges. Wiss., 5).

STOLZE, R. G. 1976. Hymenophyllaceae, Ferns

and fern allies of Guatemala. Fieldiana, Bot.,

39: 51-90.

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

menophyllaceae, pp. 97-1 24, in Ferns and allied

plants, Springer- Verlag, New York.

Key to Genera of Hymenophyllaceae

a. Indusium bivalvate, the valves '/2 or more the length of the indusium (rarely less); receptacle not or

rarely and slightly exserted; venation anadromous; ultimate segments sometimes serrate, lacking false

veins I. Hymenophyllum

a. Indusium tubular and entire, or bilabiate, rarely bivalvate (and then the valves not more than '/a the

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

Hymenophyilum Sm., Mem. Acad. Roy. Sci. (Tur-

in) 5: 418. 1793. LECTOTYPE (designated

by Presl, Hymenophyllaceae 31. 1843): Hy-

menophyllum tunbridgense (L.) Sm. Figure 11.

Leptocionium Presl, Hymenophyllaceae 26. 1843.

TYPE: Leptocionium dicranotrichum Presl = Hy-

menophyllum dicranotrichum (Presl) Sadeb.

Hymenophyllum subg. Leptocionium (Presl) Christ,

Farnkr. Erde 20. 1897.

Sphaerocionium Presl, Hymenophyllaceae 33. 1843.

TYPE: Sphaerocionium hirsutum (L.) Presl (Tri-

chomanes hirsutum L.) = Hymenophyllum hir-

sutum (L.) Sw.

Hymenophyllum subg. Sphaerocionium (Presl) C. Chr.,

Index fil. Suppl. 3: 5. 1934.

Mecodium(Cope\.)CopeL, Philipp. J. Sci. 67: 17. 1938.

Hymenophyllum subg. Mecodium Copel., Philipp. J.

Sci. 64: 93. 1937. TYPE: Hymenophyllum po-

lyanthos (Sw.) Sw.

Plants epiphytic, occasionally epipetric or ter-

restrial. Stem filamentous, long-creeping, usually

bearing scattered trichomes and small, delicate

roots. Leaves monomorphic, petiolate, commonly

1-20(-50) cm long or, in H. speciosum, to nearly

2 m long. Lamina simple and entire (outside Peru)

or pinnatifid to decompound, glabrous or sparsely

to densely pubescent. Ultimate segments entire to

50

FIELDIANA: BOTANY

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

with sori. Hymenophyllum crispum: d, pinna with sori. (a from Seller 295, El Salvador, F, b from Plowman 6074, F,

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

(reticulate in a few Old World species), anadro-

mous, false veins absent. Sori terminal on the veins.

1 ml usiiim bivalvate, the valves Vz or more the length

of the indusium (rarely less), the sporangia borne

on an elongate, sometimes subglobose, receptacle.

Receptacle not or rarely exserted beyond the mouth

of the indusium.

Nearly 300 species of Hymenophyllum are found

throughout tropical regions of the world, although

a few occur in wet subtropical and temperate areas.

While most are epiphytes, several species have

been reported from wet clay or rocky banks in

dense forests. A number of genera and/or subgen-

era have been segregated by various authors, but

in the following treatment Peruvian species are

aligned into three well-defined subgenera: Hy-

menophyllum, Mecodium, and Leptocionium. A

good revision of the latter was done by Morton

( 1 947) under the name Sphaerocionium, but this

and others still require careful study, and some of

the species concepts are only provisionally treated

below. Although there is no attempt here at a for-

mal subgeneric classification, the species are so

designated in the key and their descriptions follow

in natural order.

To facilitate use of the key to species, some ex-

planation of the diagnostic characters may be help-

ful here, especially in subg. Leptocionium. Caution

is recommended when using the couplets that sep-

arate those species having trichomes borne only

on veins and margins of segments (Morton's sub-

section Ciliata) from those in which trichomes are

borne also on the segment surface between the veins

and margins (his subsection Lanata). In a few

species trichomes between the veins may be

sparse— present only in areas toward the base of

segments. Care must be exercised that these are

not overlooked. Conversely, there are several

species with stellate trichomes so dense on veins

and margins, and with stalks so long, that they

may at first appear to spring from the intervening

tissue.

Furthermore, certain groups of species in subg.

Hymenophyllum and Leptocionium are distin-

guished by the presence of lamellae: accessory green

wings or crests of tissue that emerge from the veins

or costae, but not on the same plane as the lamina.

Typically these lamellae are made conspicuous by

their size, shape, or abundance, but in H. loba-

toalatum they are small and scattered, and in H.

plumosum they are not at all foliose, but instead

are low and inconspicuous, and usually obscured

by the dense, matted tomentum covering the en-

tire lamina. In H. mirificum they could be mis-

taken for trichomes, since many of them, although

four to six cells wide at the base, abruptly become

uniseriate and grade to a filiform tip.

Reference

MORTON, C. V. 1947. The American species of

Hymenophyllum sect. Sphaerocionium. Contr.

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-

cellular trichome [subg. Hymenophyllum] 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 (i.e., secondary segments all borne on acroscopic side); sori (most of them)

strongly arcuate, thus arranged nearly perpendicular to the plane of the lamina; petiole less

than 0.3 mm in 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, elliptical, or rhomboid, most of them somewhat to markedly longer

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, elliptic, or conical, broadest at or beyond the middle, the base

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 I a 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 mm in diameter 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 least proximal 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

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

base 1 7. H. crispum

m. Petiole (of mature leaves) stout, 0.4-1 .5 mm in diameter 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 mm in H. amabile) . . . 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

y. Rachis nonalate (sometimes alate distally); leaves indeterminate; petiole tri-

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 pilose, but surfaces not obscured by the trichomes;

rachis commonly alate, at least distally, only occasionally nonalate 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,

often strongly and gradually reduced toward base 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 . .

33. H. pyramidatum

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.

(Schrader) 1800(2): 99. 1802.

Leaves 4-25(-30) cm long, 1 .5-8 cm broad. Pet-

iole 0.3-0.8 mm in diameter, nonalate or margin-

ate to alate distally, glabrous to sparsely or mod-

erately pubescent with catenate trichomes. Lamina

broadly or narrowly oblong, not or rather abruptly

reduced at base, 3-4-pinnatisect. Rachis broadly

to narrowly alate throughout, or nonalate just at

the base, sparsely to moderately provided with

flexuous, catenate trichomes. Pinnae 3-20(-25)

pairs, adnate, or proximal 1-2 pairs short-stalked,

subequilateral except at base, there truncate ac-

roscopically, cuneate basiscopically, with 2-8 pairs

of pinnatisect to 2-pinnatisect segments. Sori 1-8

per pinna, not or scarcely immersed in the segment

tissue, most of them borne in the same plane as

the lamina, not or only occasionally arcuate. In-

dusia narrowly to broadly oblong or elliptic, the

apex obtuse to subacute and entire to laciniate,

receptacle narrow-cylindrical or fusiform to ovoid,

not exserted or occasionally slightly so.

The species occurs in the West Indies; southern

Mexico to Panama; Venezuela and Colombia to

Brazil and Bolivia.

This species has been misunderstood, both in

its nomenclature and taxonomy, since it was first

recognized by Swartz in 1788. The source of the

greatest confusion is a supposed type collection

(BM) of Hymenophyllum cristatum Hooker & Grev.

This sheet contains two specimens of//, fucoides,

while pinned to it is an exact copy of/. 148, Icon,

fil. 1829, the illustration of//, cristatum, complete

with globose receptacle and conspicuous lamellae

on the veins, which is an Ecuadorean species of a

different section, Buesia (Morton) Morton. This

led to another error, for the subsequent illustration

H. fucoides in Hooker's "Century of Ferns" (1854)

was evidently produced from this mixed sheet,

thus misleading later investigators about some of

the principal diagnostic features (for a full discus-

sion see Stolze, Amer. Fern J. 77: 137-140. 1987).

A second source of confusion has been the high-

ly variable nature of the species, which has

prompted the recognition of a number of related

taxa at various levels over the years. Morpholog-

ical features are rather constant in the West Indian

specimens, where leaves are quite small and com-

pact; but variability increases in Central America,

where some leaves are larger and more elongate,

and apices of indusia are deeply dentate. Even

greater differences are found in South American

plants, especially in Peru, where variability is at

its peak. It is likely that part of this may be due to

the effects of ecological niches and altitudinal levels,

but this kind of analysis must wait for a much-

needed revision of this part of the genus.

For purposes of this treatment //. fucoides has

been segregated into four varieties, a key to which

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 Ic. var. pedicellatum

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-

ure lla-b.

Trichomanes fucoides Sw., Prodr. 136. 1788. TYPE:

Jamaica, Swartz (holotype, s; isotypes, B, BM;

phOtOS, F & GH Of B, US of BM).

Leptocionium fucoides (Sw.) Presl, Hymenophyllaceae

27. 1843.

Meringium fucoides (Sw.) Copel., Philipp. J. Sci. 67:

45. 1938.

Petiole (0.3-)0.4— 0.8 mm in diameter, nonalate,

occasionally marginate near apex. Rachis narrow-

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

pinna, apex of indusia entire to denticulate, rarely

dentate.

On trunks and branches of trees in wet forest,

rarely on wet rock walls, 1 700-3300 m, Amazonas,

San Martin, Huanuco to Puno.

West Indies; southern Mexico to Panama; Ven-

ezuela and Colombia to Brazil and Bolivia.

The most diminutive of these ferns scarcely dif-

fer from H. tunbridgense (L.) J. Sm., the type of

the genus, which is said to differ in its smaller

leaves and in its strongly arched sori. The latter

species supposedly occurs in the West Indies and

in Europe and Africa, but it may be only another

component in this complex. Some authors place

it, with H. peltatum and a few others, in sect.

Hymenophyllum, distinguished principally by the

strongly arched sori, which are borne nearly per-

pendicular to the plane of the lamina. In Old World

specimens especially, this character is inconstant

and is but one of the problems which needs closer

scrutiny when the group is revised.

Amazonas: Trail east of La Peca in Serrania de Bagua,

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

3397 (B, F, us). Cuzco: Cerro Chuyapi, Bues ASS (GH,

us), A56 (us). Puno: Sandia, Weberbauer 713 (B).

Ib. Hymenophyllum fucoides var. calodictyon

(Bosch) Stolze, comb. & stat. nov.

Hymenophyllum calodictyon Bosch, Ned. Kruidk.

Arch. 5(3): 172. 1863. TYPE: Peru, Ruiz 84 (ho-

lotype, B!).

Petiole 0.5-0.8 mm in diameter, conspicuously

alate to marginate distally. Rachis broadly alate

throughout, each of the alae plane and much wider

than the rachis. Pinnae all adnate, with ultimate

segments plane. Sori ( l-)2-8 pairs to a pinna, apex

of indusia entire.

On trees in wet forests, Cajamarca and Ama-

zonas south to Cuzco, 2100-3300 m.

Ecuador; Peru.

In addition to the characters mentioned in the

key, var. calodictyon commonly may be distin-

guished from the other varieties by the broader

segments, membranaceous tissue, and the lighter,

yellow-green color.

Cajamarca: Prov. Cutervo, Dist. San Andres, Agua

Fria, A. Diaz et al. 5340 (USM). Amazonas: Prov. Bagua,

Cordillera Colan SE of La Peca, Barbour 3586 (F, MO,

USM). Prov. Bongara, near Pomacocha, Wurdack 865 (F,

GH, uc, us). Huanuco: Prov. Huanuco, Carpish, Plow-

man 6074 (F, GH, us). Pasco: Prov. Oxapampa, E of Abra

Cantarizu, Skog et al. 5108 (us). Cuzco: Prov. Paucar-

tambo, Dist. Marcachea, near Achirani, Vargas 11142

(F, uc, us).

56

FIELDIANA: BOTANY

Ic. Hymenophyllum fucoides var. pedicellatum

(Klotzsch) Hieron., Bot. Jahrb. Syst. 34: 435.

1904.

Hymenophyllum pedicellatum Klotzsch, Linnaea 20:

439. 1847. TYPE: Venezuela, Merida, Moritz 346

(holotype, B; isotypes, BR, us in part!).

Leptocionium pedicellatum (Klotzsch) Fourn., Bull.

Soc. Bot. France 19: 249. 1872.

Petiole 0.3-0.5 mm in diameter, nonalate.

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.

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

Venezuela; Colombia; Ecuador; Peru.

This may be synonymous with H. peruvianum,

which see under Comments. It is likely that //.

ectocarpon Fee, of Central America and the Lesser

Antilles, should be included here also.

Amazonas: Prov. Bagua, Cordillera Colan, SE of La

Peca, Barbour 3598 (F, MO, USM), 3976 (F, MO). Prov.

Bagua, Montenegro-Chiriaco, Sagdstegui 5931 (GH).

Id. Hymenophyllum fucoides var. chachapoyense

Stolze, var. nov.

Differt a var. fucoide characteribus sequentibus: alae

rhachidis latae et saepe undulatae; segmenta ultima saepe

undulata.

Petiole 0.5-0.9 mm in diameter, commonly alate

distally, at least at the apex. Rachis broadly alate

throughout, the alae often undulate. Pinnae ad-

nate, the ultimate segments often undulate and

their margins discontinuously conduplicate. Sori

commonly 3-10 to a pinna, apex of indusia entire.

TYPE— Peru, Dept. Amazonas, Prov. Chacha-

poyas, Cerros de Calla Calla, 18 km above Lei-

mebamba on road to Balsas, 3100 m, Hutchison

& Wright 5659 (holotype, GH!; isotype, uc!).

On trunks and branches of trees, in wet forests

and moist, wooded ravines, 2900-3700 m, Ama-

zonas, Huanuco.

Colombia (Dept. del Valle); Ecuador (Prov. Car-

chi & Napo); Peru.

This is a robust variety of higher elevations,

similar to var. calodictyon in its stout petioles and

larger leaves (to 25 cm long, occasionally longer).

In its undulate to crispate rachis wings and ulti-

mate segments, var. chachapoyense resembles Hy-

menophyllum tortuosum Hooker & Grev. of Chile;

however, alae and segments of the latter are much

more strongly crispate and are setose-serrate as

well.

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

Vaca, Macbride 4460 (F, us).

2. Hymenophyllum peltatum (Poir.) Desv., Mem.

Soc. Linn. Paris 6: 333. 1827.

Trichomanes peltatum Poir. in Lam., Encycl. 8: 76.

1808. TYPE: Mauritius, "Ile-de-France, Bory de

Saint-Vincent (V.s. in Herb, du Petit-Thouars)"

(not located).

Leaves 2.5-8 cm long, 1-2 cm broad. Petiole

0.15-0.3 mm in diameter, nonalate, glabrous.

Lamina narrowly oblong to lanceolate, slightly re-

duced at base, 2-3-pinnatisect. Rachis nonalate,

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-

ondary segments all bome on acroscopic side). Sori

1 or 2 per pinna, not immersed in the segment

tissue, most of them strongly arcuate, thus ar-

ranged nearly perpendicular to the plane of the

lamina. Indusia subspheroid to ovoid, the apex

obtuse, entire, receptacle narrow-cylindrical, not

or rarely exserted.

On rocky slopes and cliffs, 2800-4700 m, Junin,

Cuzco, Madre de Dios, Puno.

Peru; Bolivia; Argentina; Chile; Europe; Africa.

Diem and Lichtenstein (1959) reported that H.

peltatum is highly variable in southern South

America. They recognized six varieties in Argen-

tina and/or Chile and also consider that it is also

present in one form or another in areas of the Old

World. Apparently only var. peltatum occurs in

Peru. It is a much more delicate fern than H. fu-

coides and can be easily distinguished from the

latter by the strongly dimidiate pinnae. Most sori

are rather conspicuously arched out of the plane

of the lamina, a character which Morton (1968)

used to separate sect. Hymenophyllum from others

in subg. Hymenophyllum. This feature is not con-

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

diagnostic character above the level of species.

Junin: Prov. Huancayo, Dist. Huancayo, Huaytapal-

lana. Sounders 1 167 (GH). Cuzco: "Ccarcco" (Ccorca?),

Biies 1395 (us). ?Madre de Dios: Pinasniocj, "Panticalla

Pass" (Pantiacolla?), Cook & Gilbert 1868, 1870 (us).

Puno: Prov. Carabaya, Ayapata, Vargas 10705 (GH).

3. Hymenophyllum mirificum Morton, Bot. Gaz.

(Crawfordsville) 93: 338. 1932. TYPE: Peru,

Cuzco, Prov. La Convencion, Vilcabamba,

Biies 1600 (holotype, us!; photo, F).

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

48. 1938.

Leaves 16-38 cm long, (5-) 7-1 4 cm broad. Pet-

iole 4.5-7 cm long, (0.4— )0. 5-0. 8 mm in diameter,

nonalate or, rarely, very narrowly so or marginate

at apex, sparsely provided with brownish, flex-

uous, catenate trichomes. Lamina broadly or nar-

rowly oblong, not or abruptly reduced at base,

deeply 3-4-pinnatisect, sparsely and inconspic-

uously lamellate adaxially. Rachis strongly flex-

uous, nonalate, or weakly alate toward the apex,

sparsely provided with flexuous, catenate tri-

chomes and (adaxially) also with widely scattered

lamellae, these usually 4—6 cells wide at base,

abruptly becoming uniseriate and often grading to

a sharp, filiform tip. Pinnae 1 0-20 pairs, proximal

ones subdistant and short-stalked, distal ones ap-

proximate and adnate, costae flexuous and nar-

rowly alate, with 6-10 pairs of 2-pinnatisect sec-

ondary segments, ultimate segments long and

narrow, glabrous, plane, the margins serrate, la-

mellae inconspicuous and often widely scattered

along veins adaxially. Sori several to many, es-

pecially on distal pinnae, not or scarcely immersed

in the segment tissue, lying essentially in the plane

of the lamina. Indusia subspherical to ovoid, the

margins subentire, receptacle subglobose, not

exserted.

Endemic. In damp ravines and on trees, in wet

forests, 2800-3650 m, Cuzco.

This is closely related to Hymenophyllum cris-

tatum Hooker & Grev. of Ecuador, which has the

same general aspect and habit. Both species have

elongate, pendent leaves with flexuous rachises,

subspherical to ovoid indusia, subglobose recep-

tacles, and lamellae on axes and veins. But H.

mirificum has larger pinnae, rachis wings essen-

tially lacking, lamellae sparse and inconspicuous,

and apices of indusia entire to denticulate. Rach-

ises in H. cristatum are obviously alate through-

out, lamellae are numerous and conspicuous, and

indusia are deeply dentate to laciniate at apex. For

further discussion about the confused identity of

H. cristatum, see H. fucoides.

Cuzco: Prov. La Convencion, Vilcabamba, Biies 1592,

1593, 2104 (us). Prov. Paucartambo, Acanacu, Vargas

29 (GH); West 7144 (uc, us). Prov. Urubamba, Puyu-

pata-Yuncapata, Vargas 2922 (MO, us).

4. Hymenophyllum lamella turn Stolze, sp. nov.

Folia 7-1 5 cm longa, 2.5-4.5(-6) cm lata. Petiolus 1 .5-

5 cm longus, 0.35-0.7 mm diametro, conspicue alatus.

Lamina 2-pinnatisecta vel 3-pinnatisecta, conspicue la-

mellata. Rhachis recta, late alata, lamellis conspicuis in-

structa. Pinnae 9-13-jugatae, confertae vel imbricatae,

adnatae, l-2.5(-3) cm longae, costis et venis lamellatis

praeditae. Segmenta saepe ultima undulata. marginibus

serratis. Indusia sphaeroidea vel ovoidea, brevipedicel-

lata, marginibus integris, receptaculis subglobosis vel

ovoideis.

Leaves 7-15 cm long, 2.5-4.5(-6) cm broad.

Petiole 1.5-5 cm long, 0.37-0.7 mm in diameter,

conspicuously alate well toward, or quite to, base,

sparsely provided with brownish, flexuous, caten-

ate trichomes. Lamina oblong-lanceolate, not or

scarcely reduced at base, 2-3-pinnatisect, con-

spicuously lamellate adaxially. Rachis straight,

broadly alate throughout, sparsely to amply pu-

bescent with flexuous, catenate trichomes and also

provided with conspicuous scalelike lamellae, these

several to many cells wide at base, long-acuminate

and often grading to a sharp, filiform tip. Pinnae

9-13 pairs, crowded to imbricate, adnate at base,

l-2.5(-3) cm long, adaxially lamellate on costae

and veins as on the rachis, with 3-6 pairs of 2-

pinnatisect or (distally) pinnatisect segments, ul-

timate segments glabrous, usually slightly to

strongly undulate, the margins serrate, often con-

duplicate. Sori few, commonly 1 per pinna on the

basal acroscopic segment, not immersed in the

segment tissue, lying essentially in the plane of the

lamina. Indusia subspherical to ovoid, the margins

entire, receptacle subglobose or ovoid, not or

slightly exserted.

TYPE— Peru, Huanuco, Prov. Huanuco, trail

from S entrance of Carpish tunnel to crest of ridge,

Luteyn & Luteyn 5474 (holotype, us!; isotype, NY!).

Endemic. In wet forests, on tree trunks and

branches, 2750-3000 m, Huanuco, Cuzco, Puno.

58

FIELDIANA: BOTANY

Some of the specimens cited below were earlier

suspected of being new species. On the sheets of

Bites A 17 & A20 (us) are Morton's penciled re-

marks, "sect. Buesia, n. sp.?"; and Lechler 2568

(B) is annotated by Mettenius, "//. latipes Mett."

Apparently Mettenius intended to publish, but a

search of the literature has revealed nothing under

this name. In its small size, compact lamina, and

conspicuously alate axes, this is the most distinc-

tive species in sect. Buesia.

Cuzco: Alturas de Sicre, Biies 1562 (us). Valley of Rio

Urubamba, Biies A- 17, A-20 (us). Prov. La Convention,

at Klaus' Folly, above Camp 5, Dudley 1 WOO (GH). Puno:

Tatanara, Lechler 2568 (B).

fewer pinnae, but the character of its sori is dif-

ferent. The narrowly cuneate base of the indusium

is deeply immersed in the segment tip, whereas in

H. polyanthos the indusium base is broadly cu-

neate to obtuse and only slightly immersed in the

tissue.

San Martin: Prov. Mariscal Caceres, Dist. Tocache

Nuevo, Cerro Sinsin, Plowman & Schunke 11478 (F).

Loreto: Mishuyacu, near Iquitos, Klug 1518 (F, NY, us).

Huanuco: Environs of Jingo Maria, Aguilar 304 (uc).

Tingo Maria (as San Martin), Allard 20741, 21485 (GH,

us). Pasco: Puerto Bermudez (as Junin), Killip & Smith

26415 (F, us). Cuzco: Valle de Lares, alturas del Rio

Lachac, Biies 1813a (us).

5. Hymenophyllum apiculatum Kuhn, Linnaea 35:

391. 1868. TYPE: Venezuela, Edo. Aragua,

Colonia Tovar, Fendler 32 (holotype, B; iso-

type, us).

Hymenophyllum dendritis Rosenst., Repert. Spec. Nov.

Regni Veg. 6: 308. 1909. TYPE: Bolivia, San Car-

los near Mapiri, Buchtien 1093 (holotype, B?; is-

otype, us!).

Mecodium dendritis (Rosenst.) Copel., Philipp. J. Sci.

67: 26. 1938.

Mecodium apiculatum (Kuhn) Vareschi, Flora Ven-

ezuela, Caracas 1: 198. 1969.

Leaves 1.5-4(-5) cm long, 0.8-1.8 cm broad.

Petiole 0.3-1 .5 cm long, 0. 1-0.25 mm in diameter,

marginate to narrow-alate, glabrous or with a few

scattered trichomes, especially at very base. Lam-

ina ovate or deltoid-ovate, not or scarcely reduced

at base, 2-3-pinnatifid, glabrous. Rachis glabrous,

strongly flexuous to fractiflex, alate throughout.

Pinnae 3-6 pairs, adnate, larger ones 0.5-1.2 cm

long, bearing 1—4 pairs of secondary segments. Ul-

timate segments entire, plane, or occasionally the

margins involute or conduplicate. Sori 1-4 to a

pinna, usually confined to apical portion of lam-

ina. Indusia rhomboid or ovoid, apex entire and

obtuse to subacute, with broadly cuneate base,

about halfway immersed in the segment tissue,

receptacle filiform, not exserted, bearing 6-12 spo-

rangia.

In dense forests on branches and trunks of trees

and on fallen logs, 100-1 500(-2000) m, San Mar-

tin, Loreto, Huanuco, Pasco, Cuzco.

Venezuela; Colombia; Peru; Bolivia.

This would appear to be simply a reduced form

of Hymenophyllum polyanthos; however, it not

only has smaller leaves, more delicate petioles, and

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

(Schrader) 1800(2): 102. 1802. Figure lie.

Trichomanes polyanthos S\v., Prodr. 137. 1788. TYPE:

Jamaica, Swartz (holotype, s; isotypes, B, BM;

photOS, US Of S, F & GH Of B).

Mecodium polyanthos (Sw.) Copel., Philipp. J. Sci. 67:

19. 1938.

Mecodium mexiae Copel., Univ. Calif. Publ. Bot. 19:

294, /. 48. 1941. TYPE: Peru, Huanuco, near

confluence of Rio Huallaga, Mexia 8282 (holo-

type, uc!; isotypes, B!, F!, MO!).

Hymenophyllum mexiae (Copel.) Morton, Contr. U.S.

Natl. Herb. 38: 173. 1968.

Leaves (6-)8-15 cm long, 2-5(-6) cm broad.

Petiole 0.5-5 cm long, 0.3-0.6 mm in diameter,

alate near the apex or throughout, glabrous. Lam-

ina ovate or elliptic, scarcely reduced at base (or

proximal 1-2 pairs of pinnae slightly reduced), 3-

4-pinnatisect, glabrous. Rachis glabrous, straight

or slightly flexuous, alate throughout. Pinnae 8-

14 pairs, essentially adnate, but not strongly over-

lapping the rachis, larger ones 1 -4 cm long, bearing

2-many secondary segments. Ultimate segments

entire, plane or occasionally a few with condupli-

cate segment margins, larger ones commonly more

than 1 mm broad, the free portion relatively short,

usually about twice as long as broad. Sori 2-many

per pinna. Indusia relatively small, usually about

as broad as the segment apex, ovoid, elliptic, or

lanceolate, broadest toward the base, apex obtuse

or subacute, the broadly cuneate base slightly im-

mersed in the segment tissue, margins entire, re-

ceptacle filiform to narrowly cylindrical or fusi-

form, not exserted, bearing 6-15 sporangia.

In dense forests, on fallen logs, and on trunks

of trees and tree ferns, 1 00-2000(-2400) m, Ama-

zonas, Loreto, San Martin south to Puno.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

59

Pantropical.

This is one of the most widely distributed ferns

of tropical regions around the world and with its

many close relatives represents one of the more

taxonomically difficult species complexes. Com-

ponents of the group each have only a few distin-

guishing characters and these are often inconstant.

Some of the features seem to vary more or less in

different geographic areas, so that species concepts

in each of these regions vary proportionately. The

group has long been misunderstood; consequently,

in most herbaria, a number of specimens may be

filed incorrectly under Hymenophyllum polyan-

thos.

For purposes of this treatment, H. polyanthos

includes H. mexiae, which differs only in its small-

er leaves. Leaves of H. apiculatum are also very

small, but this species (q.v). is characterized by

other, quantitative, features, and so should be con-

sidered distinct. Hymenophyllum mathewsii may

be merely a more luxuriant, higher elevation form

of H. polyanthos, with larger leaves and sori, and

the axes less distinctly alate. Another species of

the subgenus occurring in Peru is H. myriocarpum,

which differs principally in shape and character of

the indusia: in most specimens sori appear to be

pedicellate, for the apices of ultimate segments are

strongly constricted; furthermore, mature indusia

are subglobose, or often noticeably broader than

long. This contrasts with the essentially ovoid in-

dusia of//, polyanthos et al., commonly somewhat

longer than broad and slightly to strongly im-

mersed in the segment tips, which are not or

scarcely constricted. While this character is prob-

ably the most reliable one in subg. Mecodium in

Peru, unfortunately even it is not 1 00% constant,

for on occasional specimens indusia may vary

somewhat from one shape to another. Conse-

quently, most species in the subgenus must be sep-

arated by suites of characters. Ultimately, taxo-

nomic problems will be solved only by thorough

study of species groups throughout their entire

range.

Two other characters are often helpful in sep-

arating H. polyanthos (at least in Peru) from H.

myriocarpum— particularly useful where sterile

specimens are involved. Pinnae of H. myriocar-

pum commonly have basal segments strongly

overlapping the rachis. In H. polyanthos pinnae

are adnate and basal segments often crowd or touch

the rachis, but rarely do some of them overlap.

Also, H. polyanthos occurs at lower elevations in

Peru, 100-2000(-2400) m; whereas H. myriocar-

pum is found at ( 1 700-)2000-4200 m.

Amazonas: Prov. Bagua, 1 2 mi E of La Peca, Harbour

2466 (uc in part). San Martin: Ravine E of Tingo Maria,

Allard 21446 (GH, us). Prov. Mariscal Caceres, Tocache

Nuevo, J. Schunke 5693 (F). Loreto: Sierra del Pongo,

Mexia 6291a (GH, uc). Huanuco: Prov. Huanuco, Cerro

Cucharas, Woytkowski 1152(cH). Prov. Huamalies, Rio

Monzon, 20 km W of Tingo Maria, Tryon & Tryon 5299

(GH). Pasco: Prov. Oxapampa, Cordillera San Matias,

Leon 317 (F, GH, USM). Pichis Trail, Enenas (as Junin),

Killip & Smith 25651 (F, NY, us). Junin: La Merced, E

of Quimiri Bridge, Killip & Smith 24022 (F, NY, us).

Madre de Dies: Prov. Manu, Cerro de Pantiacolla, Rio

Palotoa, Foster et al. 10683 (F). Cuzco: Prov. Paucartam-

bo, Kosnipata, Vargas 17860 (GH). Puno: San Gaban,

Lechler 3327 (B).

7. Hymenophyllum mathewsii Bosch, Ned.

Kruidk. Arch. 5: 162. 1863. SYNTYPES:

"Peruvia," Mathews (B!; isosyntype, p; prob-

able isosyntype, B!; photos, F, GH, & us of p).

Ecuador, Quito, Cuming (B).

Leaves (6-) 1 4-30 cm long, 1.5-4 cm broad. Pet-

iole 2.5-6(-8) cm long, 0.35-0.5 mm in diameter,

nonalate, glabrous. Lamina linear-lanceolate to

oblanceolate, rather abruptly reduced or strongly

and gradually reduced at base, 2-3-pinnatisect,

glabrous. Rachis glabrous, alate throughout except

usually nonalate at base, or there narrow-alate on

only one side. Pinnae 14-18 pairs, adnate and often

overlapping the rachis, or basal ones short-stalked,

larger ones (1.5-)2-3.5 cm long, bearing 4-8 pairs

of secondary segments. Ultimate segments entire,

plane or occasionally undulate, or a few with mar-

gins conduplicate. Sori 4-many per pinna. Indusia

relatively large, usually slightly broader than the

segment apex, ovoid, apex obtuse to subacute, the

broadly cuneate base slightly immersed in the seg-

ment tissue, margins entire, receptacle short and

filiform, bearing 6-12 sporangia.

Cloud forests, commonly on trunks and branch-

es of trees, 2 1 00-3 1 50 m, Huanuco, Pasco, Cuzco.

Ecuador; Peru.

Hutchison 1751 (cited below) has delicate, di-

minutive leaves only 6-8 cm long; this depauper-

ate condition probably results from the unusual

habitat, "in cave." This species is very closely re-

lated to Hymenophyllum polyanthos (q.v.) and may

not merit distinction at the species level. Normal

specimens of//, mathewsii tend to have larger sori,

in relation to the segment apex, and longer, pen-

dent leaves, with no trace of tissue along the petiole

and, commonly, none between the proximal two

pinnae. Leaves of H. polyanthos are smaller and

60

FIELDIANA: BOTANY

more compact, erect or arching, with rachis dis-

tinctly alate. The petiole is marginate or alate

throughout, or at least distally. Often the wing is

deciduous upon drying, but traces can usually be

found by close scrutiny.

Huanuco: Prov. Huanuco, Carpish, Plowman 6074A

(F). Pasco: Prov. Oxapampa, San Alberto, Yanachaga,

van der Werffel al. 8437 (MO). Cuzco: Huadquina, Bues

1346 (us). Prov. La Convention, Cochapata, Valle de

San Miguel, Bues 2176 (us). Prov. Urubamba, summit

of Huayna Picchu, Hutchison 1751 (atypical) (F, uc).

8. Hymenophyllum trichomanoides Bosch, Ned.

Kruidk. Arch. 5(3): 158. 1863. SYNTYPES:

Peru, San Martin, Tarapoto, Spruce 4696

(isosyntypes, GH!, NY!, P!, us!). Colombia,

Moritz (BM?), Schomburgk(BM?). Ecuador, Pi-

chincha, Quito, Cuming (not located). "Brit-

ish Guiana," Schomburgk (BM?).

Mecodium trichomanoides (Bosch) Pic.-Ser., Webbia

28: 469. 1973.

Leaves (9-)l 1-26 cm long, 4-6(-8) cm broad.

Petiole 5-12 cm long, 0.4-0.8 cm in diameter,

marginate to alate distally (rarely nonalate), with

a few scattered trichomes. Lamina broadly or nar-

rowly ovate (1 or 2 pairs of proximal pinnae com-

monly somewhat reduced), 2-3-pinnatisect, gla-

brous. Rachis glabrous, slightly flexuous, narrowly

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,

bearing 4-8 pairs of secondary segments. Ultimate

segments entire, plane, or some of them with in-

volute margins, larger ones commonly less than 1

mm broad, the free portion linear, usually 2.5-4

times as long as broad. Sori 6-12 per pinna. In-

dusia relatively small, usually about as broad as

the segment apex, rhomboid, elliptical or conical,

broadest at or beyond the middle, apex acute to

obtuse, the sharply cuneate base deeply (often half-

way) immersed in the segment tissue, margins en-

tire, receptacle long, filiform, often exserted, bear-

ing 6-10 sporangia.

In dense, wet forests, on trunks of trees, 300-

3000 m, San Martin to Cuzco.

Surinam to Colombia, south to Bolivia.

Besides the characters used in the key, Hymen-

ophyllum trichomanoides can often be distin-

guished by the narrower ultimate segments. Larger

ones are commonly 0.6-0.9 mm broad and linear.

Other species of subg. Mecodium have relatively

broader ultimate segments, most of them over 1

mm broad, or if not, then only about twice as long

as broad. This is a rather reliable way to identify

sterile specimens.

Hymenophyllum trichomanoides differs sharply

from all other species of subg. Mecodium in Peru

by the narrowly cuneate sori which are deeply im-

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

specimens seen from Peru clearly match the Spruce

syntype. Some specimens examined from Colom-

bia and Venezuela have shorter and broader in-

dusia and much shorter petioles, and it is possible

that these represent a variant. However, this has

not been substantiated by comparison with the

other syntypes from northern South America,

which we have not seen. With H. trichomanoides

probably should be included H. trianae Hieron.

of Colombia. A syntype of the latter, Lehmann

7410 (us), matches Spruce 4696, except that re-

ceptacles are not exserted.

Huanuco: Cerros del Sira, SW slope of Rio Llullapichis

watershed, Dudley 13375 (GH). Pasco: Prov. Oxapampa,

Palcazu Valley, Cabeza de Mono, D. Smith 3756 (F, MO).

Junin: Schunke Hacienda, above San Ramon, Killip &

Smith 24844 (F, GH, NY, us). Cuzco: Sahuayacu, Bues

804, 817 (us).

9. Hymenophyllum ferax Bosch, Ned. Kruidk.

Arch 4: 392. 1 859. TYPE: Venezuela, Merida,

Funck & Schlim 1578 (holotype, L?; isotype,

p; photo, us of P).

Mecodium ferax (Bosch) Copel., Philipp. J. Sci. 67:

25. 1938.

Leaves 16-38 cm long, 4-8 cm broad. Petiole

2-9 cm long, 0.4-0.9 cm in diameter, alate toward

apex (wings sometimes deciduous), essentially gla-

brous. Lamina ovate to narrow-elliptic, scarcely

to somewhat reduced at base (1-several proximal

pairs of pinnae reduced), 3-pinnatisect, glabrous.

Rachis glabrous, straight or slightly flexuous, alate

throughout, rachis wings and those of other axes

sometimes conduplicate. Pinnae 1 2-24 pairs, ad-

nate, and basal segments overlapping the rachis,

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-

mixed with normal ones. Ultimate segments en-

tire, plane to slightly undulate. Sori 8-many per

pinna. Indusia relatively large, subrotund, circular

in outline or broader than long, apex erose, base

rounded, not immersed in the greatly constricted

segment apex, receptacle short, filiform, bearing

7-12 sporangia.

On trees in wet forests, 2400-3300 m, Cuzco.

Venezuela; Colombia to Peru.

This is most closely related to Hymenophyllum

myriocarpum var. endiviifolium, from which it dif-

fers in the erose indusia tips and the presence of

one to several unusually elongate pinnae which

are scattered among the normal ones— the latter

a condition similar to that of H. undulatum. A

remarkably huge specimen, Lechler 2420 (B; not

other 2420 at B, F, K: H. elegans), was collected

in Peru at an undesignated site. This is apparently

a monstrous form of H. ferax. It matches this

species in every way, including the erose indusia,

but the leaf is 85 cm long and the petiole nearly

2 mm in diameter. It is by far the stoutest specimen

of Hymenophyllaceae we have seen and belies the

appellation "filmy fern."

Cuzco: Vilcabamba, Bues 1601 (us). Prov. Paucartam-

bo, Pilahuata, Vargas 4912 (us). Dept. Unknown: Lech-

ler 2420 (B in part).

10. Hymenophyllum myriocarpum Hooker, Sp. fil.

1: 106, /. 37 d. 1844. TYPE: Colombia, Hart-

weg 1530 (holotype, K!; isotypes, BM, p; photo,

F of P).

Leaves 7-28 cm long, 2.5-5(-6) cm broad. Pet-

iole 1.5-8(-10) cm long, 0.3-0.7 mm in diameter,

marginate to alate throughout or at least distally

(wings sometimes deciduous), essentially gla-

brous. Lamina lanceolate to ovate, elliptic, or ob-

lanceolate, scarcely to strongly reduced at base, 2-

3-pinnatisect, glabrous. Rachis glabrous, straight

or slightly flexuous, alate throughout, rachis wings

and those of other axes often conduplicate. Pinnae

8-30 pairs, adnate and basal segments strongly

overlapping the rachis, larger ones 1-4 cm long,

bearing 5-10 pairs of secondary segments. Ulti-

mate segments entire, plane to undulate, often with

conduplicate margins. Sori 5-many per pinna. In-

dusia subglobose or broader than long, much

broader than (and not at all immersed in) the

strongly constricted segment apex, rounded at base,

rounded and entire at apex, receptacle short to

nearly obsolete, filiform, bearing 8-20 sporangia.

This, like H. polyanthos (q.v.), is a widespread

and greatly misunderstood species which is sorely

in need of detailed study. It seems to consist of

several, not highly distinctive components which

are recognized as varieties. With var. myriocar-

pum might possibly be included H. axillare Sw.

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-

ica and in South America from Venezuela and

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.

myriocarpum.

Mecodium myriocarpum (Hooker) Copel., Philipp. J.

Sci. 67: 25. 1938.

Leaves (9-) 12-28 cm long. Lamina elliptic, ob-

long or oblanceolate, broadest at or beyond the

middle, several proximal pinnae somewhat to

greatly reduced. Rachis and ultimate segments

plane, some of their margins occasional condu-

plicate. Sori 10-20 per pinna.

62

FIELDIANA: BOTANY

In wet forests, usually pendent from tree trunks

and branches, or on wet, mossy rocks, rarely on

the forest floor, (1 700-)2000-3200 m, Cajamarca,

Amazonas, Huanuco to Cuzco.

Mexico to Costa Rica; Venezuela; Colombia to

Bolivia.

Cajamarca: Prov. Hualgayoc, Hacienda Taulis,

Hutchison & Bismarck 6415 (F, GH [as Lambayeque],

MO, uc, us). Amazonas: Prov. Bongara, WSW of Po-

macocha, Wurdack 866 (GH, us). Huanuco: Mito, Bryan

206a (F). Pasco: Enenas (as Junin), Pichis Trail, Killip &

Smith 25700 (F, NY, us). Cuzco: Prov. Urubamba, along

Rio Urubamba near town of Machu Picchu, Tryon &

Tryon 5410 (F, GH, us).

lOb. Hymenophyllum myriocarpum var. nigres-

cens (Liebm.) Stolze, stat. & comb. nov.

Hymenophyllum nigrescens Liebm., Kongel. Danske

Vidensk. Selsk. Skr., Naturvidensk. Afd. 5,1: 292.

1849. LECTOTYPE (designated by A. R. Smith,

Flora of Chiapas 2: 133. 1981): Mexico, Puebla,

Chinantla, Liebmann 537 (lectotype, c; isolec-

totypes, BM, K, P).

Sphaerocionium nigricans KJotzsch, Linnaea 18: 536.

1 844, not Hymenophyllum nigricans Colla, 1836.

SYNTYPES: Peru, Dombey 87 (syntype, u!; iso-

syntype, B); Venezuela, Aragua, Colonia Tovar,

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

268b (syntype, B; isosyntype, BM).

Hymenophyllum nigricans (KJotzsch) Kunze, Bot. Zeit.

(Berlin) 244. 1847.

Hymenophyllum nigricans (KJotzsch) Copel., Phillip.

J. Sci. 67: 25. 1938.

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

ultimate segments plane, their margins frequently

conduplicate. Sori 3-15 per pinna.

Erect or arching from tree trunks or branches,

rarely from wet rocks, in thickets and wet forests,

1750-3400 m, Amazonas to Cuzco.

Southern Mexico to Costa Rica; Venezuela; Co-

lombia to Bolivia.

Sterile specimens can easily be confused with

H. polyanthos, but the latter occurs at lower ele-

vations, and laminae are less compact. See dis-

cussion of H. polyanthos for further comments.

Amazonas: Prov. Chachapoyas, Cerros de Calla Calla

above Leimebamba, Hutchison & Wright 6987 (in part:

F, GH, uc). San Martin: Prov. Rioja, Pedro Ruiz-Moy-

obamba Road, D. Smith 4376 (F, MO). Pasco: Prov. Ox-

apampa, Abra los Mellizos, 4-8 km from Enenas, Skog

et al. 5033 (vs). Junin: Huacapistana, Killip & Smith

24164 (us). Ucayali: Plantation Azul, Ridoutt (USM). Prov.

Coronel Portillo (as Loreto), NE of pass at La Divisoria,

Skoget al. 5157 (us). Cuzco: Valley of Urubamba, Sues

A18,A19(us).

lOc. Hymenophyllum myriocarpum var. endivi-

ifolium (Desv.) Stolze, stat. & comb. nov.

Hymenophyllum endiviifolium Desv., Mem. Soc. Linn.

Paris 6: 334. 1827. TYPE: "Peruvia" (holotype,

p; photos, GH, uc, us).

Hymenophyllum multiflorum Rosenst., Meded. Rijks-

Herb. 19: 5. 1913. TYPE: Bolivia, Comarapa,

Herzog 1951 (holotype, B; isotype, us!).

Mecodium multiflorum (Rosenst.) Copel., Philipp. J.

Sci. 67: 25. 1938.

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

469. 1973.

Leaves (12-) 15-28 cm long. Lamina ovate or

ovate-lanceolate. Rachis and ultimate segments

slightly to strongly undulate, their margins often

conduplicate. Sori 1 6-many per pinna.

On trees and wet banks in deep forest, 2400-

4200 m, Amazonas, Huanuco, Huancavelica,

Cuzco.

Colombia; Ecuador; Peru; Bolivia.

Amazonas: Prov. Chachapoyas, Calla Calla slopes near

km 4 1 5-4 1 8 of Leimebamba-Balsas road, Wurdack 1750

(us, USM). Huanuco: Prov. Huanuco, Mirador, road Aco-

mayo to Chanchao, Mexia 7761 (F, GH, K, MO, uc, us).

Huancavelica: Prov. Tayacaja, Chuspi-Tocas, between

Colcabamba and Paucarbamba, Tovar 2050 (GH, USM).

Cuzco: Prov. Paucartambo, Dist. Marcachea, near

Achirani, Vargas 11143 (F, K. in part, uc).

11. Hymenophyllum undulatum (Sw.) Sw., J. Bot.

(Schrader) 1800(2): 101. 1802.

Leaves 6-35(-40) cm long, 0.7-2.5 cm broad.

Petiole 0.5-3(-5) cm long, 0.15-0.3 mm in di-

ameter, nonalate, essentially glabrous. Lamina lin-

ear to narrow-elliptic, often strongly and gradually

reduced at base, 2-3-pinnatisect, glabrous. Rachis

glabrous, marginate to crispate-alate, commonly

nonalate toward base, or discontinuously alate on

alternate sides due to the long-decurrent pinna

bases. Pinnae 1 6-many pairs, adnate, commonly

less than 2 cm long, but often a few of them greatly

elongate and lamina-like, normal ones bearing 3-

8(-10) pairs of secondary segments. Ultimate seg-

ments entire, plane, or the margins conduplicate

in drying, to undulate or markedly crispate. Sori

3-many per pinna. Indusia nearly circular, but on

mature son most of them broader than long, not

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

63

or scarcely immersed in the segment tissue, com-

monly truncate at base, the margins entire and

plane to crispate, receptacle very short, filiform,

bearing 1^4(-5) sporangia.

This species varies greatly as to length and shape

of lamina and margins of rachis wings and ulti-

mate segments. In rocky habitats leaves usually

tend to be less than 10 cm long and narrowly el-

liptic, whereas the flaccid, linear leaves pendent

from tree branches sometimes reach 40 cm in

length. Several species have been described based

on whether the tissue is plane or undulate or

strongly crispate. However, comparison of indi-

viduals throughout the entire range seems to in-

dicate that few of these characters are very con-

sistent. Only the conspicuously crispate variants

seem to merit recognition, as indicated in the fol-

lowing key.

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 l/2 or nearly throughout 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.

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, /.

38c. 1844. TYPE: Peru, Mathews 1783 (holo-

type, K.!; isotypes, B!, F!, K!, P!; photos, F & us of

P).

Hymenophyllum rimbachii Sodiro, Anal. Univ. Quito

13: 47 (Crypt, vase. quit. 33). 1893. TYPE: Ec-

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

cated; isotype, us!).

Mecodium undulatum (Sw.) Copel., Philipp. J. Sci. 67:

26. 1938.

In wet forests on rock cliffs or pendent from tree

trunks and branches, 1500-4100 m, Huanuco to

Puno.

Greater Antilles; Guadeloupe; southern Mexico

to Panama; Surinam to Colombia, south to Bo-

livia; southeast Brazil.

Hymenophyllum reniforme and H. rimbachii

were based on depauperate specimens in which

ultimate segments are mostly plane but with their

margins conduplicate. Thus the segments appear

to be much narrower than typical H. undulatum.

Some of this folding of the margins appears to be

a product of drying and some of it merely part of

the natural variation of the species.

Huanuco: Tambo de Vaca, Bryan 624 (F, us). Pasco:

Prov. Oxapampa, San Alberto, Cordillera de Yanachaga,

van der Werffet al. 8482 (MO, uc). Junin: Prov. Tarma,

Yanango Mountains, Weberbauer 2135 (B). Cuzco: Pau-

cartambo, slopes of Pilahuata, Vargas 4909 (MO, us).

Puno: Sandia, Weberbauer 1293 (B).

1 1 b. Hymenophyllum undulatum var. fendlerian-

um (J. W. Sturm) Stolze, comb. & stat. nov.

Hymenophyllum fendlerianum J. W. Sturm in Mart.,

Fl. bras. 1(2): 291. 1859. LECTOTYPE (desig-

nated by Lellinger, Mem. New York Dot. Gard.

38: 12. 1984): Venezuela, Edo. Aragua, Colonia

Tovar, Fendler 35 (us; isolectotype, BR; photo, us

of BR).

Hymenophyllum contortum Bosch, Ned. Kruidk. Arch.

5(3): 170. 1863. TYPE: Costa Rica, Aguacate,

Hoffman (B).

Hymenophyllum polycarpum Kuhn, Linnaea 35: 391.

1868. TYPE: "Peruvia" (holotype, B!; photo, F).

Mecodium fendlerianum (J. W. Sturm) Copel., Phil-

ipp. J. Sci. 67: 26. 1938.

Mecodium contortum (Bosch) Copel., Philipp. J. Sci.

67: 26. 1938.

In forests on wet, rocky cliffs, or pendent from

tree trunks and branches, 1250-2900 m, Ama-

zonas, Huanuco, Junin, Puno.

Hispaniola; southern Mexico; Guatemala; Cos-

ta Rica; Surinam to Colombia, south to Bolivia;

southeastern Brazil.

Amazonas: Prov. Bagua, Cordillera Colan SE of La

Peca, Barbour 3539, 3738 p.p. (MO). Huanuco: Cushi,

trail to Tambo de Vaca, Bryan 684 (F, us). Playapampa,

Macbride 4498 (F, us). Junin: Schunke Hacienda, above

San Ramon, Killip & Smith 24843 (NY, us). Puno: Prov.

64

FIELDIANA: BOTANY

Carabaya, road from San Gaban to Macusani, Maas et

al. 6109 (MO). Dept. Unknown: Without location, Dom-

bey (P).

12. Hymenophyllum molle Morton, Contr. U.S.

Natl. Herb. 29: 149. 1947. TYPE: Peru, Cuz-

co, Vilcabamba, Achiyayoc Inca ruins, Biies

2102 (holotype, us!; isotype, GH!).

Sphaerocionium molle (Morton) Pic.-Ser., Webbia 28:

471. 1973.

Leaves indeterminate, to 35 cm long and 3.5 cm

broad. Petiole 3-6 cm long, 0.2-0.3 mm in di-

ameter, nonalate, sparsely pubescent with simple

to (rarely) forked trichomes, or glabrate. Lamina

2-pinnatisect or nearly 2-pinnate, abundantly pu-

bescent, the rachis commonly nonalate through-

out. Pinnae short-stalked (at least proximal ones),

cut deeply or nearly to costa, costae regularly and

conspicuously alate on both sides, the 4-9 pairs

of segments simple or bifid, entire, plane, veins

and margins abundantly or densely pubescent with

stellate trichomes, but trichomes lacking between

veins and margins. Indusia slightly broader than

long, often broader than the segment tips.

Endemic. In forests, 2200-3600 m, Cuzco.

Cuzco: Huadquina, Biies 708 (us). Prov. La Conven-

cion, Biies 2072, 2115 (us). Prov. Paucartambo, Vargas

3636 (GH).

13. Hymenophyilum trichophyllum HBK., Nov.

gen. sp. 1: 27. 1815.

Leaves indeterminate, to 25 cm long and 3 cm

broad. Petiole 0.5-4 cm long, 0.2-0.25 mm in di-

ameter, nonalate, sparsely pubescent with simple

to stellate trichomes, or glabrate. Lamina 2-pin-

nate-pinnatisect, abundantly pubescent, the rachis

commonly nonalate throughout. Pinnae short-

stalked (at least proximal ones), cut deeply or quite

to costa, costae partially nonalate or discontin-

uously alate by the decurrent bases of segments,

normal pinnae with 2-3(-4) pairs of segments (but

often a few of them greatly elongate and equaling

the lamina in size and shape), the segments (or

pinnules) simple or bifid, plane to undulate or cris-

pate, veins and margins abundantly to densely pu-

bescent with stellate trichomes, but trichomes

lacking between veins and margins. Indusia about

as broad as long and nearly as broad as the segment

tips.

The species occurs in Guatemala; Costa Rica;

Panama; Guyana to Venezuela and south to Bo-

livia.

The leaves of this delicate fern sometimes in-

tertwine to form mats on the trunks of trees. A

singular feature is the irregular character of the

pinnae: often, but not always, one to several pin-

nae become greatly elongate and once again pin-

nate-pinnatisect, each of these sometimes equaling

the length of the leaf on which it is borne. Of the

several varieties of Hymenophyllum trichophyl-

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-

phyllum. TYPE: Venezuela, Cumana, be-

tween Cocollar and Guardia de San Augustin,

Humboldt (holotype, P).

Hymenophyllum procerum Bosch, Ned. Kruidk. Arch.

4: 409. 1859, nom. nud.

Sphaerocionium trichophyllum (HBK.) Copel., Phil-

ipp. J. Sci. 67: 32. 1938.

In Peru, thus far known only from one collec-

tion: on shaded rocks, Puno.

Elsewhere, in wet forests, on rocks, clay banks

or trees, 2000-3800 m; range the same as the

species.

The name H. procerum was based on H. pul-

chellum sensu Mett., for which there was no de-

scription. The specimen probably intended as the

type is the only collection of var. trichophyllum

thus far known from Peru. It closely matches other

specimens of var. trichophyllum and is cited here.

Puno: San Gaban, Lechler 2250 (B, BR, F, P).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

65

13b. Hymenophyllum trichophyllum var. buesii

Morton, Contr. U.S. Natl. Herb. 29: 1 52. 1 947.

TYPE: Peru, Cuzco, Prov. La Convention,

Abra Mirador, Bues 2076 (holotype, us!; is-

otype, GH!).

Thus far known only from the type collection,

3150m.

Although Morton distinguished his var. buesii

from var. trichophyllum solely on the obviously

undulate segments, it also appears that the former

has less densely pubescent laminae. However, un-

til more specimens are collected it remains to be

seen if either of the differences are sufficiently sig-

nificant to merit even varietal separation.

14. Hymenophyllum elegans Sprengel, Syst. veg.

4: 133. 1827. TYPE: Brazil, Sellow (holotype,

LZ destroyed; isotype, B!).

Hymenophyllum tenerrimum Bosch, Ned. Kruidk.

Arch. 5: 185. 1863. TYPE: Peru, San Martin,

Tarapoto, "in monte Campana," Spruce 4702

(holotype, K!; isotype, BM!).

Sphaerocionium tenerrimum (Bosch) Pic.-Ser., Web-

bia28:472. 1973.

Sphaerocionium elegans (Sprengel) Copel., Philipp. J.

Sci. 67: 32. 1938.

Leaves determinate, to 20 cm long and 2 cm

broad. Petiole 0.5-2 cm long, 0.15-0.2 mm in di-

ameter, nonalate, sparsely pubescent with simple

or forked trichomes. Lamina pinnate-pinnatisect,

sparsely to moderately pubescent, the rachis non-

alate, or narrowly and irregularly alate distally or

throughout due to the decurrent bases of pinnae.

Pinnae sessile or adnate, pinnatifid or pinnatisect,

the l-2(-3) pairs of segments, simple or (basal

ones) bifid, entire, plane to slightly undulate, mar-

gins sparsely to moderately pubescent, with tri-

chomes highly variable: simple to 2-3-forked at

base, or the branches again forked; trichomes rare

on veins and lacking between veins and margins.

Indusia slightly broader than long, about as broad

as the segment tips.

On shaded rocks, San Martin, Puno, and from

two other collections, localities unspecified, 3000

m.

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.

Hymenophyllum tenerrimum was said to differ

only by lack of trichomes on the veins, and by

having marginal trichomes mostly three-forked at

the base. However, in H. elegans, trichomes are

only occasional to rare or lacking on the veins, and

marginal trichomes are highly variable, often in-

cluding many which are three-forked at base. The

isotype of H. tenerrimum has occasional tri-

chomes on the veins, and a number of marginal

ones that are simple or one-forked. In fact, both

taxa may be only a variant of//, lineare (Sw.) Sw.,

of Central America and the West Indies, which

also lacks trichomes on veins, but which appears

to have more pinna segments and simpler and less

variable marginal trichomes. Further study is

needed.

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

4700, 4701 (K). Puno: San Gaban, Lechler (B). Dept.

Unknown: Lechler (NY, us), Lechler 2420 (B, F, K; another

2420 at B is H. ferax).

15. Hymenophyllum adiantoides Bosch, Ned.

Kruidk. Arch. 5: 184. 1863. TYPE: Peru, San

Martin, Tarapoto, Spruce (holotype, L; prob-

able isotypes, B!, GH!, K!, L!, NY, us; photos,

GH & us of L).

Sphaerocionium adiantoides (Bosch) Pic.-Ser., Web-

bia28:470. 1973.

Leaves indeterminate, to 1 2 cm long and 2 cm

broad. Petiole 1-2 cm long, 0.2-0.25 mm in di-

ameter, nonalate, sparsely pubescent with (mostly)

stellate trichomes. Lamina pinnate-pinnatisect,

sparsely to moderately pubescent, the rachis non-

alate near the base, narrowly alate distally or ir-

regularly alate due to the decurrent bases of pin-

nae. Pinnae adnate, pinnatifid, or pinnatisect, the

3-4 pairs of segments simple or bifid, entire, slight-

ly undulate, veins and margins sparsely to mod-

erately pubescent with (mostly) stellate trichomes,

but trichomes lacking between veins and margins.

Indusia as long as or slightly longer than broad,

about as broad as the segment tips.

Apparently known only from the type collec-

tion. With this probably should be included Hy-

menophyllum sprucei Baker, under which see

Comments.

16. Hymenophyllum hirsutum (L.) Sw., J. Bot.

(Schrader) 1800(2): 99. 1802.

66

FIELDIANA: BOTANY

Trichomanes hirsutum L., Sp. pi. 2: 1098. 1753. TYPE:

based on t. 50b of Plumier, Traite foug. Amer.,

apparently representing a plant from the West

Indies.

Trichomanes ciliatum Sw., Prodr. 136. 1788. TYPE:

Jamaica, Swartz (holotype, s; isotype, B; photo,

us of s).

Hymenophyllum ciliatum (Sw.) Sw., J. Bot. (Schrader)

1800(2): 100. 1802.

Sphaerocionium ciliatum (Sw.) Presl, Hymenophyl-

laceae 34. 1843.

Sphaerocionium hirsutum (Sw.) Presl, Hymenophyl-

Iaceae34. 1843.

Leaves determinate, to 12 cm long and 4 cm

broad. Petiole 0.3-2.3 cm long, 0.2-0.3 mm in

diameter, strongly alate (often nearly to base),

sparsely provided with stellate trichomes. Lamina

2-pinnatisect, sparsely to moderately pubescent,

the rachis regularly and conspicuously alate

throughout. Pinnae 6-15 pairs, ovate or oblong-

ovate, deeply incised, the 2-5 pairs of segments

plane, simple to bifid, the veins and margins pu-

bescent with stellate or (on the margins) 2-forked

trichomes, but trichomes lacking between veins

and margins. Indusia as long as or (usually) longer

than broad, somewhat broader than the segment

tips.

In dense, wet forests, on branches and trunks of

trees, 500-2000 m, Loreto, Amazonas to Puno.

Mexico to Panama; West Indies; Guianas to

Venezuela, south to Bolivia and Brazil.

Amazonas: Prov. Bagua, E of La Peca, Barbour 2753

(MO). San Martin: Tarapoto, Monte Campana, Spruce

4698 (K). Loreto: Balsapuerto, lower Rio Huallaga basin,

Killip & Smith 28539 (F, GH, NY, us). Huanuco: Prov.

Huanuco, Dist. Churubamba, crest of Santo Toribio,

Mexia 8145 (B, F p.p., GH, us). Pasco: Prov. Oxapampa,

W side of Cordillera de San Matias, D. Smith 2054 (MO).

Cuzco: Prov. La Convencion, Valle San Miguel, San Ja-

cinto, Bues 2139 (us). Puno: San Gaban, Lechler 2296

(B).

1 7. Hymenophyllum crispum HBK., Nov. gen. sp.

1: 26. 1815. TYPE: Venezuela, Silla de Ca-

racas, Humboldt & Bonpland (holotype, p;

photos, GH & us of P). Figure lid.

Sphaerocionium crispum (HBK.) Klotzsch, Linnaea

1: 537. 1844.

Leaves determinate, to 20 cm long and 4 cm

broad. Petiole 0.5-5 cm long, 0.1-0.3 mm in di-

ameter, alate near apex, sparsely provided with

delicate, simple or 1 -forked trichomes. Lamina

pinnate to nearly 3-pinnate, sparsely pubescent,

the rachis regularly and conspicuously alate

throughout, the wing strongly undulate or undu-

late-crispate. Pinnae 8-20 pairs, simple (distally)

or bifid to nearly 2-pinnate, the 1-8 pairs of seg-

ments undulate or crispate, the veins and margins

sparsely pubescent, marginal trichomes simple or

1 -forked, but trichomes lacking between veins and

margins. Indusia as long as or (usually) longer than

broad, Oiten broader than the segment tips.

The species occurs in Jamaica, Mexico, Gua-

temala, Costa Rica, Venezuela, Colombia, Peru,

Bolivia, and Brazil.

Hymenophyllum crispum might be confused with

H. valvatum, another species in the subgenus with

undulate segments. However, the latter is larger

and coarser with a much thicker petiole, and the

rachis wing and segments are not as strongly un-

dulate. Two varieties are recognized here.

Key to Varieties

a. Pinnae simple (distally) to trifid or, if pinnatisect, the secondary segments 1-2 pairs, simple to bifid

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-

ure lid.

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

3000 m, Amazonas, Cuzco, Puno.

Range the same as the species.

Amazonas: Prov. Chachapoyas, along Rio Ventilla, W

of Molinopampa, Wurdack 1510 (F, GH, uc, us). Cuzco:

Rio Lachac, Sues 1813 (GH, us). Prov. Urubamba, Ma-

chu Picchu, lit is et al. 1061 (GH, uc, us). Puno: Sandia,

Weberbauer 797 (B).

1 7h. Hymenophyllum crispum var. bipinnatisec-

tum Morton, Contr. U.S. Natl. Herb. 29: 161.

1947. TYPE: Peru, Cuzco, Alturas de Sieve,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

67

Bties 1580 (holotype, us!; isotype, GH!; photo,

us).

Endemic.

Thus far known only from the type and one other

collection from Peru: Cuzco: Alturas de Sieve, Bues 1528

(us).

18. Hymenophyllum valvatum Hooker & Grev.,

Icon. fil. 2, /. 279. 1831. TYPE: Ecuador, for-

ests of Esmeraldas, on trunks of trees, Jame-

son (holotype, K; possible isotype, K!).

Hymenophyllum crispatulum Bosch, Ned. Kruidk.

Arch. 4: 412. 1859. TYPE: Peru, Tatanara (Puno),

Lechler 2530 (holotype, L?; isotypes, B!, P!; pho-

tos, GH & us of P).

Hymenophyllum platylobum Bosch, Ned. Kruidk.

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.,

Philipp. J. Sci. 67: 31. 1938.

Leaves determinate, to 20 cm longhand 6 cm

broad. Petiole of mature leaves (2-)3-lOcm long,

0.4-1.0 mm in diameter, commonly alate just be-

low lamina base, pubescent with simple or forked

trichomes or glabrate. Lamina pinnate-pinnatisect

to nearly 2-pinnate-pinnatifid, sparsely to mod-

erately pubescent, the rachis alate throughout, the

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

to elliptic-lanceolate, incised deeply to the costa

into bifid or pinnatifid pinnules, ultimate segments

slightly to strongly undulate, the veins (abaxially)

and margins sparsely to amply pubescent, but tri-

chomes lacking on tissue between veins and mar-

gins. Indusia commonly longer than broad, and

slightly longer than the segment tips.

On tree trunks in wet forests, 900-1800 m,

Huanuco, Junin, Puno.

Lesser Antilles; Guyana to Colombia, south to

Bolivia.

It is with some reluctance that this is maintained

as distinct from Hymenophyllum microcarpum,

from which it seems to differ only in the undulate

ultimate segments and (sometimes) rachis wings,

and the character of the petiole trichomes. Even

these features are variable, as only in a few spec-

imens examined are the segments and rachis wings

markedly undulate to crispate— most are only

moderately so. Conversely, in a number of spec-

imens determined to be H. microcarpum, the ul-

timate segments are only slightly undulate. A per-

fect intermediate is the isotype of H. platylobum,

which is placed here in synonymy with H. val-

vatum. The petiole trichomes of this specimen

match those of//, microcarpum, but the segments

and rachis wings are scarcely or only slightly un-

dulate. Other features considered by Morton and

subsequent workers seem not to be significant, at

least in specimens from Peru. Furthermore, both

species may be only robust forms of H. hirsutum,

which seems to differ chiefly in its smaller lamina

and more delicate petiole. The entire complex is

in need of detailed study.

Huanuco: Rio Llullapichis watershed, Cerros del Sira,

Dudley 13339 (GH). Junin: Chanchamayo Valley, C.

Schunke 1491, 1559 (F, us). Puno: "St. Gavan" (San

Gaban), Lechler (K). Prov. Sandia, Chunchusmayo, We-

berbauer 1237 (B).

19. Hymenophyllum microcarpum Desv., Mem.

Soc. Linn. Paris 6: 333. 1827. TYPE: His-

paniola (holotype, P; photo, uc).

Hymenophyllum beyrichianum Kunze, Linnaea 9: 108.

1834. TYPE: Peru, Huanuco, Pampayacu, Poep-

pig (holotype, LZ destroyed).

Mecodium microcarpum (Desv.) Copel., Philipp. J.

Sci. 67: 25. 1938.

Sphaerocionium microcarpum (Desv.) Copel., Phil-

ipp. J. Sci. 67: 34. 1938.

Leaves determinate, to 30 cm long and 10 cm

broad. Petiole of mature leaves (3-)5-15 cm long,

0.6-1.0 mm in diameter, broadly alate just below

lamina base, pubescent with simple, forked, and

stellate trichomes or glabrate. Lamina pinnate-

pinnatisect to nearly 2-pinnate-pinnatifid, sparse-

ly to moderately pubescent, the rachis conspicu-

ously alate throughout. Pinnae 10-16 pairs, ovate-

to elliptic-lanceolate, incised deeply or nearly to

the costa into bifid or pinnatifid pinnules, the ul-

timate segments entire or deeply bifid, the veins

(abaxially) and margins sparsely to moderately pu-

bescent, but trichomes lacking on tissue between

the veins and margins. Indusia commonly longer

than broad, and slightly broader than the segment

tips.

On tree trunks and clay banks in wet forests,

1 500-2700 m, Amazonas, Huanuco to Cuzco.

Greater Antilles; Guatemala to Panama; Guy-

ana to Colombia, south to Bolivia and Brazil.

68

FIELDIANA: BOTANY

With this probably might be included H. val-

vatum, under which see for further discussion.

Amazonas: Prov. Bagua, E of La Peca, Barbour 2547

(MO). Huanuco: Rio Llullapichis watershed, ascent of

Cerros del Sira, Dudley 13290B (MO). Pasco: Prov. Ox-

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

(GH). Prov. La Convention, Alturas de Pintobamba,

Vargas 3294 (MO, us).

20. Hymenophyllum ruizianum (Klotzsch) Kunze,

Hot. Zeit. (Berlin) 1847: 199. 1847.

Sphaerocionium ruizianum Klotzsch, Linnaea 18: 535.

1 844. TYPE: Peru, Ruiz 85 (holotype, B!; isotype,

MA?, P; photos, GH & us of P).

Leaves determinate, to 45 cm long and 12 cm

broad. Petiole of mature leaves (7-) 1 0-20 cm long,

0.7-1.5 mm in diameter, nonalate, densely pu-

bescent with mostly simple or forked trichomes.

Lamina 2-3-pinnatifid, sparsely to amply pubes-

cent on both sides of the lamina, the rachis alate

distally. Pinnae commonly patent or their tips

drooping, 10-18 pairs, ovate-lanceolate, incised

nearly to the costa into pinnatifid or 2-pinnatifid

pinnules, the ultimate segments entire or bifid, the

veins and margins sparsely to moderately pubes-

cent, but trichomes lacking on tissue between veins

and margins. Indusia varying from broader than

long to longer than broad, commonly slightly

broader than the segment tips.

In wet forests, on tree trunks and banks of rav-

ines, and in sphagnum on forest floor, 1000-3300

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

Colombia; Ecuador; Peru; Bolivia.

A singular feature of this species is the tendency

for the long, flexible pinnae to droop gracefully at

their tips, a character which is usually preserved

in dried specimens. Perhaps its nearest relative is

H. trapezoidale Liebm., from Mexico and north-

ern South America, which is a much smaller fern

with less dissected pinnae which are rather strong-

ly ascending (never drooping) and with more del-

icate, merely sparsely pubescent, petioles.

Amazonas: Prov. Chachapoyas, above Leimebamba,

Hutchison & Wright 5553 (F, GH, MO, uc, us). Huanuco:

Playapampa, Macbride 4516 (F, us). Pasco: Prov. Oxa-

pampa, San Alberto, Cordillera de Yanachaga, van der

Werff et al. 8423 (MO), 8439, 8491 (MO, uc). Cuzco: Prov.

La Convention, Dudley 10790 (F, GH, MO). Puno: San

Gaban, Lechler 2260 (B).

21. Hymenophyllum simplex Morton, Contr. U.S.

Natl. Herb. 29. 171. 1947. TYPE: Peru,

Huanuco, Tambillo, Jelski 897 (us!).

Sphaerocionium simplex (Morton) Pic.-Ser., Webbia

28: 472. 1973.

Leaves determinate, fragile and small, 5-7 cm

long. Petiole slender, 0.1-0.15 mm in diameter,

nonalate, glabrate or sparsely pubescent with sim-

ple to stellate trichomes. Lamina simply pinnate

or pinnatisect, moderately pubescent, the rachis

narrowly alate distally, nonalate at base or in the

proximal half. Pinnae few, ascending, simple, lin-

ear, entire, the tissue and veins amply provided

with orange, stellate trichomes. Veins lacking ac-

cessory wings. Indusia about as broad as long, usu-

ally narrower than the pinna tips.

Endemic. Besides the type (Huanuco) thus far

known only from two other collections: Cajamarca

and San Martin. Habitat and elevation not re-

corded.

This delicate little fern is distinctive among the

species of subg. Leptocionium in its few, ascend-

ing, linear, entire pinnae. Although we now know

it only from two Peruvian collections, its small,

inconspicuous leaves make it difficult to detect. It

is hoped that more diligent search will uncover

additional specimens throughout Peru and, per-

haps, in other areas of South America as well.

Cajamarca: Cutervo, Jelski 914 (P, us). San Martin:

Tarapoto, Spruce (K), collection is without number, but

with notation "conf. 4048"; #4048 is type number of H.

tarapotense.

22. Hymenophyllum fragile (Hedwig) Morton,

Contr. U.S. Natl. Herb. 29: 172. 1947.

Trichomanes fragile Hedwig, Fil. gen. sp., t. 18. 1802.

TYPE: based on illustration from plant collected

in "America meridionalis."

Sphaerocionium fragile (Hedwig) Pic.-Ser., Webbia 28:

471. 1973.

Leaves determinate, (3-)6-18 cm long, 1.5-2.5

cm broad. Petiole ca. 0.2 mm in diameter, sparsely

to moderately pubescent with simple, forked, or

often stellate trichomes. Lamina pinnate-pinna-

tisect, moderately pubescent, the rachis broadly

and consistently alate throughout. Pinnae (3-)5-

1 2 pairs, deeply 1 -forked (distally) to pinnatisect,

the 1-2 pairs of segments entire, the tissue and

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

69

veins moderately provided with orange, short-

stalked, stellate trichomes. Veins lacking accessory

wings. Indusia about as broad as long, usually nar-

rower than the ultimate segments, the receptacle

not exserted.

In dense forests, on tree trunks or mossy banks,

800-2700 m, Pasco, Junin, Ucayali, Cuzco.

Mexico to Panama; Greater Antilles; Venezuela

and Colombia to Bolivia and Brazil.

This may be confused with Hymenophyllum ele-

gantulum, under which see for further discussion.

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

Quimiri Bridge, near La Merced, Killip & Smith 23834

(NY, us). Ucayali (as Loreto): Prov. Coronel Portillo, NE

of pass at La Divisoria, Skog et al. 5156 (us). Cuzco:

Prov. La Convencion, Pintobamba, Vargas 3262 (MO,

us).

23. Hymenophyllum elegantulum Bosch, Ned.

Kruidk. Arch. 4: 408. 1859. LECTOTYPE

(designated as type by Morton, Contr. U.S.

Natl. Herb. 29: 170. 1947): based on t. 33a

(Hooker, Sp. fil. 1 . 1 844), which in turn, ac-

cording to Morton, was based on Ecuador,

Pillzhum, Jameson 366 (K.?; probable isolec-

totype, NY!).

Hymenophyllum pulchellum sensu Hooker, Sp. fil. 1,

/. 33a. 1844, not Schlecht. & Cham. 1830.

Sphaerocionium elegantulum (Bosch) Copel., Philipp.

J. Sci. 67: 32. 1938.

Leaves indeterminate, 12-35 cm long, 2-4 cm

broad. Petiole ca. 0.2 mm in diameter, glabrate or

sparsely pubescent with simple or forked tri-

chomes. Lamina pinnate-pinnatisect, amply pu-

bescent, the rachis flexuous, nonalate, or some-

times alate in the distal portion. Pinnae subdistant

to remote, ascending, 8-24 pairs, deeply pinnati-

sect, the 4-6 pairs of segments 1-2-forked or bifid,

the tissue and veins moderately provided with or-

ange, sessile to short-stalked, stellate trichomes.

Veins lacking accessory wings. Indusia about as

broad as long, and usually as broad as the ultimate

segments.

On tree trunks, in forests and wet clearings, 3000-

3600 m, Amazonas, Cuzco, Puno.

Mexico to Panama; Greater Antilles; Venezuela

and Colombia south to Peru and Bolivia.

Some leaves of Hymenophyllum elegantulum

vary somewhat in the character of the rachis wing,

as in the case of Biies 720, cited below. Some of

the laminae have the rachis narrowly winged well

into the proximal half, and these specimens might

be confused with H. fragile. However, the alate

condition in these atypical leaves is irregular and

mostly due to the decurrent bases of pinnae,

whereas in H. fragile the wings are very conspic-

uous and of nearly consistent width throughout.

Ama/onas: Prov. Bagua, Cordillera Colan NE of La

Peca, Barbour 3415 (F, MO). Cuzco: Michihuanuncca,

Biies 720 (us). Puno: Prov. Carabaya, Ayapata, Vargas

1071 7 (GH).

24. Hymenophyllum amabile Morton, J. Wash.

Acad. Sci. 22: 63. 1932. TYPE: Peru, Cuzco,

Prov. La Convencion, Huadquina, Bites 715

(holotype, us!; isotype, GH!; photo, F of us).

Leaves indeterminate, to 40 cm long and 4 cm

broad. Petiole 0.2-0.3(-0.4) cm in diameter, non-

alate, sparsely pubescent with mostly stellate tri-

chomes. Lamina pinnate-pinnatisect, densely to-

mentose, the rachis nonalate throughout, or rarely

slightly alate due to decurrent bases of pinnae.

Pinnae numerous, oblong or ovate-oblong, obtuse,

often incised nearly to the costa, the 4-8 pairs of

segments entire to forked, ultimate divisions en-

tire, the tissue and veins often obscured by a dense

tomentum of orange, stalked, stellate or bistellate

trichomes. Veins lacking accessory wings. Indusia

broader than long, about as broad as the segment

tips, the receptacle very short, not exserted.

Pendent from tree trunks and branches in wet

forests, 2600-4200 m, Huanuco, Cuzco, Puno.

Ecuador and Peru.

Huanuco: Tambo de Vaca, Bryan 628 (F, us). Cuzco:

Prov. La Convencion, Vilcabamba, Biies 2108, 2109,

2110 (us). Prov. La Convencion, Loma Grande, Biies

2148 (F, GH, uc, us). Urubamba, Machu Picchu, Peyton

& Peyton 1 104 (MO). Puno: Sandia, Weberbauer 796 (B).

25. Hymenophyllum speciosum Bosch, Ned.

Kruidk. Arch. 5: 181. 1863. LECTOTYPE

(designated as type by Morton, Contr. U.S.

Natl. Herb. 29: 175. 1947): Peru, Puno, San

Gaban, on tree trunks, Lechler 2246 (L?; iso-

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

photos, GH, us of P).

Hymenophyllum spectabile Kuhn, Linnaea 35: 392.

1868. TYPE: Bolivia, Yungas, D'Orbigny 175

(holotype, B?; isotype, p; photo, us of P).

70

FIELDIANA: BOTANY

Sphaerocionium spectabile (Kuhn) Copel., Philipp. J.

Sci. 67:31. 1938.

Leaves indeterminate, mature ones 30-180 cm

long, 4-9 cm broad. Petiole (of mature leaves) 0.5-

0.7 mm in diameter, nonalate, glabrate or sparsely

pubescent with simple to stellate trichomes. Lam-

ina pinnate-pinnatifid or pinnate-pinnatisect,

densely tomentose, the rachis nonalate through-

out, commonly flexuous, with a tightly appressed

covering of subsessile or short-stalked, stellate tri-

chomes. Pinnae numerous, larger ones (2.5-)3-8

cm long, lanceolate or linear-lanceolate, often at-

tenuate, incised deeply or nearly to the costa, the

6-14 pairs of segments shallowly to deeply bifid,

the ultimate divisions entire, the tissue and veins

mostly obscured by a dense tomentum of orange

(aging whitish), short-stalked, stellate trichomes.

Veins lacking accessory wings. Indusia about as

broad as long and often broader than the segment

tips, the receptacle not exserted.

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-

menophyllum karstenianum, as the two differ only

quantitatively, e.g., appressed versus spreading to-

mentum, strongly versus slightly flexuous rachis,

relative size and number of segments of pinnae.

Amazonas: Prov. Chachapoyas, Hutchison & Wright

5569 (F,GH, MO, uc, us); Sagastegui 7453 (F, HUT). Pasco:

Prov. Oxapampa, Cordillera San Gutardo, Leon et al.

534(usM). Cuzco: Vallede Lares, Biies 1808, 1817, 1821,

1931 (us). Valle de San Miguel, La Convention, Biies

2178 (GH, MO, us).

26. Hymenophyllum karstenianum J. W. Sturm,

Bot. Zeit. (Berlin) 17: 298. 1859. LECTO-

TYPE (designated as type by Morton, Contr.

U.S. Natl. Herb. 29: 176. 1947): Venezuela,

Merida, Moritz 381 p.p. (BR).

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

Webbia28:471. 1973.

Leaves indeterminate, mature ones 15-50 cm

long, 2-3.5(-4) cm broad. Petiole 0.4-0.5 mm in

diameter, nonalate, glabrate or sparsely pubescent

with simple to stellate trichomes. Lamina pinnate-

pinnatifid to -pinnatisect, densely tomentose, the

rachis nonalate throughout, straight or slightly

flexuous, abundantly to densely covered with short-

to long-stalked stellate trichomes, these mostly

spreading, not tightly appressed. Pinnae numer-

ous, larger ones 1-2.3 cm long, lanceolate or lin-

ear-lanceolate, acute to subattenuate, shallowly or

deeply incised, the 4-7 pairs of segments entire to

bifid, the tissue and veins mostly obscured by a

dense tomentum or orange to tawny, short-stalked,

stellate trichomes. Veins lacking accessory wings.

Indusia commonly as broad as long, and as broad

as the segment tips, the receptacle not exserted.

Pendent from trees or clay banks in wet forests,

700-1900 m throughout the range, but thus far

apparently known in Peru only from one collection

cited below.

Venezuela; Colombia; Peru.

This and Hymenophyllum speciosum are very

closely related. See discussion of the latter for ad-

ditional comments.

San Martin: Near Tarapoto, in Monte Campana,

Spruce 4694 (BR, us).

27. Hymenophyllum lindenii Hooker, Sp. I'll. 1:

94, t. 34c. 1844. TYPE: Venezuela, Caracas,

Linden 173 (holotype, K!; isotype, P; photos,

F of K, us of P).

Sphaerocionium lindenii (Hooker) Vareschi, Flora

Venezuela 1: 217. 1969.

Leaves indeterminate, to 35 cm long and 8-15

cm broad. Petiole (8-) 10-1 5 cm long, 0.7-1 .2 mm

in diameter, commonly nonalate, pubescent with

simple to stellate trichomes. Lamina ovate or ovate-

lanceolate, not or slightly reduced at base, nearly

2-pinnate-pinnatisect, moderately pilose, the rachis

abundantly stellate-pubescent, scarcely alate, or

alate distally (or sometimes to base). Pinnae patent

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

to subattenuate, incised nearly to the costa into 8-

14 deeply pinnatifid pinnules, the ultimate seg-

ments entire to deeply bifid, rarely undulate or

crispate, tissue between the veins sparsely provid-

ed with scattered, sessile or stalked, stellate tri-

chomes. Veins lacking accessory wings. Indusia

broader than long, and about as broad as the seg-

ment tips, receptacle not exserted.

Pendent from tree trunks in wet forests, 2300-

2450 m, Amazonas, Puno.

Venezuela; Colombia; Ecuador(?); Peru.

Hymenophyllum lindenii is one of the most ro-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

71

bust species in the subgenus, the long, stout petiole

often measuring a full millimeter in diameter. The

presence of a rachis wing, usually a good diagnostic

feature in most species of subg. Leptocionium, is

not consistent in this or in the closely related H.

plumieri. The rachis in H. lindenii is occasionally

alate to the base of the lamina; more commonly

the wing is present in the distal half but sometimes

it is nearly lacking throughout. The two Lechler

specimens cited below are most atypical, being

broadly alate even well down the petiole, and with

the wings very strongly crispate. The latter feature

apparently was what prompted Mettenius's orig-

inal annotation on this sheet as "//. valvatum."

Lechler 2489 is a mixed collection, as the other

four specimens of 2489 at Berlin are isotypes of

H. platylobum (= H. valvatum).

Amazonas: Prov. Bagua, Cordillera Colan, Barbour

3554 (MO, USM), 3602, 3751 (MO). Puno: near San Gaban,

Lechler 2489 (u). Tatanara, Lechler 2563 (B, K).

28. Hymenophyllum plumieri Hooker & Grev.,

Icon. fil. 2, /. 123. 1829. TYPE: Ecuador,

western side of Pichincha, Jameson (holo-

type, K.!; photo, F).

Hymenophyllum interruptum Kunze, Linnaea 9: 107.

1834. TYPE: Peru, Huanuco, Pampayacu, Poep-

pig 1104 (holotype, w?; isotype, HBG).

Sphaerocionium plumieri (Hooker & Grev.) Presl, Hy-

menophyllaceae 34. 1 843.

Sphaerocionium interruptum (Kunze) Presl, Hymen-

ophyllaceae 34. 1843.

Leaves indeterminate, to 50 cm long and 3-8

cm broad. Petiole 2-7 cm long, 0.4-0.6(-0.7) mm

in diameter, nonalate (or sometimes alate just at

the apex), pubescent with mostly forked to stellate

trichomes. Lamina linear, usually strongly and

gradually reduced at base, often interruptedly fer-

tile (e.g., groups of fertile pinnae separated by some

sterile ones), pinnate-pinnatifid to -pinnatisect,

moderately pilose, the rachis alate throughout, or

occasionally alate only distally or not at all. Pinnae

patent to ascending, 1 5-many pairs, 1 .5-6 cm long,

acute to subattenuate, the 5-12 pairs of segments

simple and entire to deeply bifid, the tissue be-

tween the veins sparsely to moderately provided

with sessile or stalked, stellate trichomes. Veins

lacking accessory wings. Indusia about as broad

as long or usually broader, as broad as the segment

tips.

Pendent from tree trunks and clay banks in wet

forests, 600-2200 m, San Martin to Puno.

Venezuela and Colombia to Peru and Bolivia.

It is not possible to separate this satisfactorily

from Hymenophyllum interruptum, which was said

to differ in its groups of fertile pinnae interrupted

by a few sterile ones, by its rachis not alate toward

the base, and by variations in lamina base and in

marginal trichomes. In a number of specimens

examined during this study (identified as H. in-

terruptum) fertile pinnae are continuous, and rach-

ises are fully to partially alate or (occasionally)

nonalate throughout. The entire species complex,

which also involves H. dependens Morton and H.

superbum Morton from northern South America,

needs closer examination. All may be conspecif-

ic.

San Martin: In monte Guayrapurima, prope Tara-

poto, Spruce 4025 (K, P). Huanuco: Rio Llullapichis wa-

tershed, ascent of Cerros del Sira, Dudley 13014 (GH,

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).

29. Hymenophyllum plumosum Kaulf., Enum. fil.

267. 1824. TYPE: Brazil, Chamisso (holo-

type, LZ destroyed; isotype, LE?).

Sphaerocionium plumosum (Kaulf.) Copel., Philipp.

J. Sci. 67: 30. 1938.

Leaves indeterminate, to 75 cm long and 6 cm

broad. Petiole 0.3-0.7 mm in diameter, nonalate,

pubescent with simple to stellate trichomes or gla-

brate. Lamina pinnate-pinnatifid to rarely nearly

2-pinnate, densely appressed-tomentose, the rach-

is nonalate throughout. Pinnae numerous, lanceo-

late, pinnatifid to pinnatisect, or rarely incised to

the costa, the ultimate segments entire (some prox-

imal ones bifid), the tissue and veins completely

obscured by a tomentum of sessile to short-stalked,

stellate trichomes. Veins (especially abaxially) with

low, inconspicuous lamellae. Indusia about as

broad as long, usually slightly broader than the

segment tips.

Epiphyte on trunks or branches in dense, wet

forests, or occasionally pendent from wet, clay

banks, 550-3100 m, Huanuco, Junin, Cuzco,

Madre de Dios, Puno.

Costa Rica to Colombia; Peru; Bolivia; Brazil.

72

FIELDIANA: BOTANY

The lamellae along the veins in this species are

scarcely "wings" in the sense of being broad and

thin flanges of tissue. Instead, they are commonly

long, low outgrowths borne along the abaxial or

(sometimes) adaxial sides of the veins contiguous

to, but not in the same plane with, the segment

tissue. They are often difficult to discern, since the

veins and laminar surface are so densely tomen-

tose. The best way to observe them is on the older

(proximal) pinnae where some of the indument

may have fallen away. For further discussion see

also Hymenophyllum tomentosum.

Easily confused with Hymenophyllum plumo-

sum are H. karstenianum and H. speciosum, which

have no lamellae on the veins, but are similar to

the former in most other features, including the

dense matting of tomentum obscuring the laminar

surface. Extreme care and patience must be ex-

ercised to determine whether the low outgrowths

are present or lacking. In fact, the real importance

of the presence or absence of lamellae bears more

careful analysis in the future. During this study of

Peruvian species of Hymenophyllum, it has been

noted that size and frequency of these outgrowths,

on one or both surfaces, may be more variable

than earlier assumed, and thus may be of much

less taxonomic significance.

Huanuco: Vilcabamba, Hacienda on Rio Chinchao,

Macbride 5146 (F, us). Junin: Schunke Hacienda above

San Ramon, Killip & Smith 24840 (F, GH, us). Cuzco:

Prov. Paucartambo, above Tambomayo, West 7108 (MO,

uc). Madre de Dies: Prov. Mania, Shintuya, Chavez 860

(MO). Puno: San Gaban, Lechler 2264 (B).

30. Hymenophyllum mult ialat urn Morton, Contr.

U.S. Natl. Herb. 29: 185. 1947. TYPE: Peru,

Cuzco, Prov. La Convencion, Alturas de Sieve,

Biies 1575 (holotype, us!; frag., GH!).

Sphaerocionium multialatum (Morton) Pic.-Ser.,

Webbia28: 471. 1973.

Leaves indeterminate, to 50 cm long, 3-8(-15)

cm broad. Petiole 0.4-0.7 mm in diameter, non-

alate, sparsely hirsute with mostly simple tri-

chomes. Lamina pinnate-pinnatisect, hirsute, the

rachis nonalate throughout. Pinnae numerous,

narrow-oblong or -deltoid, pinnatisect, sometimes

irregularly and greatly elongate, the ultimate seg-

ments commonly bifid, the tissue and veins mod-

erately provided with sessile to short-stalked ses-

sile trichomes. Veins bearing (especially abaxially)

irregularly interrupted, conspicuous lamellae. In-

dusia about as broad as long, equaling or slightly

broader than the segment tips.

Epiphyte in dense, wet forests, 2000-3200 m,

Amazonas, Huanuco, Pasco, Cuzco.

Colombia; Ecuador; Peru.

Shape of lamina and pinnae can be variable and

irregular in this species. Typically the leaves are

linear, with numerous pinnae 3 or 4 cm long, yet

at times, scattered along the rachis, there may be

a few greatly elongated, attenuate pinnae, occa-

sionally reaching 1 5 cm in length. Although Hy-

menophyllum multialatum has been thus far re-

ported from four departments in Peru, the great

majority are represented by Biies's collections in

Cuzco. For further discussion see H. tomentosum.

Amazonas: Prov. Bagua, Cordillera Colan, Barbour

3740 (F, MO). Huanuco: Prov. Huanuco, 5 km from Car-

pish, Tryon & Tryon 5318-B (GH). Pasco: Prov. Oxa-

pampa, San Alberto, Cordillera de Yanachaga, van der

Werffet al. 8495 (MO). Cuzco: Valley of Rio Urubamba,

Biies A- 16 (F, GH, us). Montana de Calca, Biies 1916 (GH,

us), 1919, 1923, 1932 (us). Prov. La Convencion, Cor-

dillera Vilcabamba, Dudley 1 1 137 (GH, us).

31. Hymenophyllum tomentosum Kunze, Lin-

naea 9: 107. 1834. TYPE: Peru, Huanuco,

Pampayacu, 1829, Poeppig Diar. 1134 (ho-

lotype, w?; isotype, BM?; frag., B!; probable

isotype, MO!; photo, us of MO).

Sphaerocionium tomentosum (Kunze) Presl, Hymen-

ophyllaceae 34. 1843.

Hymenophyllum fusagasugense J. W. Sturm, Bot. Zeit.

(Berlin) 1859: 297. 1859 (as "fusugasugense").

TYPE: Colombia, Cundinamarca, "Fusugasuga"

(Fusagazuga) Karsten (not located).

Hymenophyllum fusagasugense var. aberrans Mor-

ton, Contr. U.S. Natl. Herb. 29: 187. 1947 (as

"fusugasugense"). TYPE: Venezuela, Merida,

Tabay, Gehriger 584 (holotype, us!; isotypes, F!,

GH!).

Leaves indeterminate, to 50 cm long and 2.5 cm

broad. Petiole 0.5-0.6 mm in diameter, nonalate,

sparsely pubescent with mostly simple trichomes.

Lamina pinnate-pinnatifid, densely hirsute, the

rachis nonalate throughout. Pinnae numerous, ob-

long or narrow-deltoid, pinnatifid (sometimes

deeply so), the ultimate segments simple or (oc-

casionally) bifid, the tissue and veins densely cov-

ered with (and often obscured by) sessile to short-

stalked trichomes. Veins abaxially bearing irreg-

ularly interrupted, conspicuous winglike lamellae,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

73

these wings much smaller or sometimes lacking

adaxially. Indusia about as broad as long, equaling

or slightly broader than the segment tips.

On steep banks and tree trunks, 2750-3100 m,

Amazonas, Huanuco.

Colombia; Venezuela; Peru.

This is distinguished from Hymenophyllum

multialatum and H. plumosum only by the char-

acters used in the key. All three appear to be alike,

except for the size and frequency of the lamellae

along the veins and by the relative density of pu-

bescence. Of these, H. plumosum (q.v.) is the most

distinct, in that the lamellae are merely low and

inconspicuous outgrowths and the trichomes are

so densely matted as to completely obscure the

veins and tissue. In H. tomentosum the indument

is also densely matted, but the lamellae are so large

and conspicuous that many of them emerge through

the tomentum. Morton (1947) chose to separate

H. fusagasugense and var. aberrans on the pres-

ence or absence of these processes adaxially. How-

ever, the size or lack of lamellae on the adaxial

side varies considerably in the species, and the

character is difficult to observe due to the dense

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.

A thorough study of the species complex through-

out its range is needed.

Amazonas: Prov. Chachapoyas, Cerros de Calla Calla,

19 km above Leimebamba, Hutchison & Wright 5570

(uc). Prov. Chachapoyas, near km 422, Balsas-Leime-

bamba road, Wurdack 1249 (F, GH, uc, us).

32. Hymenophyllum lobatoalatum Klotzsch, Lin-

naea 20: 438. 1847. TYPE: Peru, Panatahua,

Ruiz 83 (holotype, B!; isotype, MA?).

Sphaerocionium lobatoalatum (Klotzsch) Pic.-Ser.,

Webbia28:471. 1973.

Leaves indeterminate, to 60 cm long and 8 cm

broad. Petiole 0.4-0.7 mm in diameter, alate, if

at all, only near base of lamina, pubescent with

simple to stellate trichomes or glabrate. Lamina

pinnate-pinnatifid, pubescent with stellate tri-

chomes, the rachis strongly alate throughout. Pin-

nae numerous, larger ones 20-45 mm long, 7-1 1

mm broad at base, most of them narrow-trian-

gular, the apex narrowly acute to subattenuate,

deeply pinnatind, the 7-12 pairs of ultimate seg-

ments bifid, the tissue and veins moderately pro-

vided with light or dark brown, short-stalked, stel-

late trichomes. Veins abaxially bearing small,

scattered lamellae, these about as broad as long,

but lacking adaxially. Indusia slightly longer than

broad, usually as broad as the segment tips, re-

ceptacle not exserted.

In wet forests, pendent from trunks and branch-

es of trees, 600-1300 m, Loreto and Huanuco.

Colombia; Ecuador; Peru.

There is little, beyond the key characters, to

distinguish this from Hymenophyllum pyrami-

datum. Leaves of the latter are perhaps more ro-

bust, with relatively narrower pinnae and much

denser pubescence. Both also share nearly the same

range, and more study may prove them to be con-

specific. Also very closely related is H. verecun-

dum, which is more easily separated by its much

lighter colored trichomes and shorter, relatively

broader and more crowded pinnae. But even these

characters sometimes tend to be intermediate.

Loreto: Pumayucu, between Balsapuerto and Moy-

obamba, Klug 3248 (B, F, GH, MO, us). Huanuco: Prov.

Huanuco, Dist. Churubamba, Mt. Santo Toribio, Mexia

8258 (B, F, GH, uc, us).

33. Hymenophyllum pyramidatum Desv., Mem.

Soc. Linn. Paris 6: 332. 1827. TYPE: "habitat

in America" (possibly a Dombey collection

from Peru) (holotype, P; photos, GH, us).

Sphaerocionium pyramidatum (Desv.) Copel., Phil-

ipp. J. Sci. 67: 30. 1938.

Leaves indeterminate, to 65 cm long and 6 cm

broad. Petiole 0.5-0.7 mm in diameter, often alate

1-2 cm near the apex, pubescent with simple to

stellate trichomes. Lamina pinnate-pinnatifid,

densely pubescent with simple to stellate tri-

chomes, the rachis alate throughout. Pinnae nu-

merous, larger ones 20-45 mm long, 6-1 1 mm

broad at base, most of them narrow-triangular,

the apex narrowly acute to attenuate, deeply lobed

to pinnatifid, the 6-12 pairs of ultimate segments

simple to bifid, the tissue and veins usually co-

piously provided with orange to dark brown (rare-

ly tawny), subsessile to short-stalked, stellate

trichomes. Veins abaxially with numerous, con-

spicuous, elongate, winglike lamellae (often nearly

the length of the vein), these sometimes less con-

spicuous and abundant on the adaxial surface. In-

dusia slightly longer than broad, usually as broad

as the segment tips, receptacle not exserted.

74

FIELDIANA: BOTANY

In dense, wet forests, pendent from stumps, tree

trunks, and branches, and on wet banks, 600-1 800

m, along Cordillera Central from San Martin to

Cuzco.

Colombia; Peru; Bolivia.

This belongs to a close-knit complex of species

involving Hymenophyllum verecundum and H.

lobatoalatum. See discussion of the latter for fur-

ther comments.

San Martin: Tingo Maria, Allard 20818 (us). Huan-

uco: Pampayacu. Kanehira 123 (F, GH, K, us). Pasco:

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

Mistiana and Keros, Vargas 7380 p.p. (MO).

34. Hymenophyllum verecundum Morton, Contr.

U.S. Natl. Herb. 29: 183. 1947. TYPE: Peru,

Huanuco, Churubamba, Mexia 8143-A (ho-

lotype, us!; isotypes, GH!, MO!, uc!).

Leaves indeterminate, to 50 cm long and 4 cm

broad. Petiole 0.4-0.6 mm in diameter, alate, if

at all, just below the lamina, glabrate or slightly

pubescent with simple to stellate trichomes. Lam-

ina pinnate-pinnatifid, densely pubescent with

stellate trichomes, the rachis alate throughout, or

sometimes nonalate at base. Pinnae numerous,

larger ones 6-20 mm long, 3-7 mm broad at base,

oblong to very broadly triangular, the apex obtuse

to subacute, pinnatifid, the 3-6(-8) pairs of ulti-

mate segments simple to retuse or occasionally

bifid, the tissue and veins copiously provided with

whitish to tawny, subsessile or short-stalked, stel-

late trichomes. Veins abaxially with numerous

conspicuous, elongate, winglike lamellae, these less

conspicuous and abundant on the adaxial surface.

Indusia about as long as broad and usually as broad

as the segment tips, receptacle not exserted.

In dense, wet forests, pendent from tree trunks,

branches, and wet banks, (5 50-) 1500-2 700 m,

Amazonas, Huanuco, Junin, Cuzco, Madre de

Dios, Puno.

Colombia; Peru; Bolivia.

This is closely related to Hymenophyllum pyr-

amidatum and H. lobatoalatum. See discussion of

the latter for further comments.

(us). Madre de Dios: Prov. Manii, Shintuya, Chavez 833

p.p. (MO). Puno: Prov. Sandia, E of Oconeque, Hodge

6090 (F, GH, us).

35. Hymenophyllum tarapotense Stolze, sp. nov.

Folia usque ad 6 cm longa et 1.6 cm lata. Petiolus 1-

2 cm loneus. 0.1-0.15 mm diametro. Lamina pinnato-

pinnatifida. Rhachis versus basin non alata. Pinnae 6-

12-jugatae, bifidae vel pinnatifidae, usque ad 1 cm lon-

gae, pinnae basales admodum reductae, marginibus, venis

et interveniis trichomatibus stellatis instructis. Venae

lamellis cristiformibus numerosis praeditae.

Leaves indeterminate, to 6 cm long and 1.6 cm

broad. Petiole 1-2 cm long, 0.1-0.15 mm in di-

ameter, nonalate, sparsely provided with simple

trichomes. Lamina pinnate-pinnatifid, pubescent

with stellate trichomes, the rachis weakly alate dis-

tally, nonalate toward the base. Pinnae 6-12 pairs,

bifid, or distal ones pinnatifid with 3-4 segments,

larger ones 1 cm long, basal ones often reduced to

mere auricles, the margins, veins, and intervening

tissue moderately to amply provided with tawny

trichomes. Veins bearing numerous, conspicuous

accessory foliar crests not in the plane of the lam-

ina, these less frequent on the adaxial surface. In-

dusia about as long as broad, as broad as the seg-

ment tips, receptacle minute, not exserted.

TYPE— Peru, Tarapoto (Dept. San Martin) in

monte Guayrapurima, Aug. 1856, Spruce 4048

(holotype, K!; photo, F). Another Spruce specimen

at K, with notation "conf. 4048," is H. simplex.

Known only from the type.

This tiny fern is unquestionably the smallest of

the species in subg. Leptocionium which bear ac-

cessory tissue on their veins. These foliar processes

are not long, low wings as seen in Hymenophyllum

pyramidatum, but are separate, crestlike flanges of

tissue similar to those frequently seen in more

delicate specimens of//, verecundum. Superficially

//. tarapotense closely resembles H. fragile, in its

small size, delicate petiole, and reduced, often bi-

fid, pinnae. It may seem strange that this distinc-

tive species has not been described before, nor

collected again in over 1 00 years. However, many

of these diminutive members of the genus are very

easily overlooked by collectors.

Amazonas: Prov. Bagua. Cordillera Colan, Barbour

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).

Cuzco: Hacienda Pintobamba, Chaupimayo, Biies 1943

Comments

Hymenophyllum peruvianum Hooker & Grev.,

Icon. fil. 2, /. 208. 1831. TYPE: Ecuador (as

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

75

Peru), Prov. Esmeraldas, Jameson (holotype,

K).

The original drawing seems to illustrate most of

the features characteristic of Hymenophyllum fu-

coides var. pedicellatum, except that the apex of

the indusium is deeply dentate rather than lacin-

iate. It is likely that the two taxa are synonymous,

but we have not seen the type to verify this. The

problem is but one of many in the H. fucoides

complex begging solution through a complete re-

vision of the group.

Hymenophyllum poeppigianum Presl, Hymeno-

phyllaceae 54. 1843. TYPE: Peru, Huanuco,

Pampayacu, Poeppig (holotype, PR; probable

isotype, P; photos, GH & us of P).

Some authors have placed this with Hymeno-

phyllum polyanthos. The type has not been seen,

but judging from the protologue and photos of a

probable isotype (P) the species is more likely to

be H. myriocarpum. The protologue refers to: the

indusia "ovate-subrotund"; the rachis alate

throughout; the petiole alate toward the apex; the

lamina linear-lanceolate with the base acute. The

description of the indusia could conceivably fit

either H. polyanthos or H. myriocarpum, but that

of the lamina (as well as the photo of the possible

isotype) more closely match H. myricarpum or H.

undulatum. However, the petiole and (usually) the

rachis base of H. undulatum are nonalate.

Hymenophyllum sprucei Baker in Hooker & Bak-

er, Syn. fil. 65. 1 867. TYPE: Peru, San Martin,

Tarapoto, Spruce (not located).

Morton (p. 155, 1947) considered that this is

the same as Hymenophyllum adiantoides, and there

is nothing in the description to argue otherwise. It

is possible that Baker was unaware of the publi-

cation of the latter species, and even that he based

his new species on the same specimen cited earlier

by van den Bosch. Specimens (B, GH, K, L, NY, us)

thought to be probable isotypes of H. adiantoides

are those of Spruce from Tarapoto. They bear the

notation "conf. 4700" and are conceivably the ones

on which Baker based his "//. sprucei."

Hymenophyllum trifidum Hooker & Grev., Icon,

fil. 2, 1. 196. 1 83 1 . TYPE: Ecuador (as "Peru"),

Esmeraldas, Jameson (E?).

In the original description the type was cited as

having come from Peru, but Esmeraldas is an Ecu-

adorean province in which Jameson did much of

his collecting. Yet Hymenophyllum trifidum still

might be expected in Peru, as it could be merely

a form of H. elegans. From the description and

plate it appears to be very similar to the latter, but

until the type is located the question must remain

unanswered.

II. Trichomanes

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

chomanes scandens L. Figure 12.

Didymoglossum Desv., Mem. Soc. Linn. Paris 6: 330.

1827. TYPE: Didymoglossum muscoides (Sw.)

Desv. (Trichomanes muscoides Sw.) = Trichom-

anes hymenoides Hedwig.

Trichomanes subg. Didymoglossum (Desv.) C. Chr.,

Index fil. xiv. 1906.

Trichomanes subg. Achomanes Presl, Hymenophyl-

laceae 15. 1843. TYPE: Trichomanes crispum L.

Trichomanes subg. Pachychaetum Presl, Hymeno-

phyllaceae 16. 1843. TYPE: Trichomanes rigi-

dum Sw.

Homoeotes Presl, Gefassbiindel Farm, 23. 1847.

TYPE: Homoeotes heterophylla Presl = Trichom-

anes humboldtii Lell.

Plants epiphytic, occasionally epipetric or ter-

restrial. Stem stout and erect or decumbent to long-

creeping and filiform, bearing scattered to abun-

dant trichomes and delicate to large, fibrous roots

or (especially in subg. Didymoglossum) lacking

roots. Leaves monomorphic or (less commonly)

dimorphic, 0.5—40 cm or longer. Lamina entire to

nearly 5 -pinnate, subsessile to petiolate, with pet-

iole commonly shorter than the lamina, glabrous

or with trichomes borne mostly on costae and

margins. Veins free or, in Trichomanes diversi-

frons, reticulate near the lamina margin, pinnate

(anadromous or catadromous) or sometimes fla-

bellate, false veins also present in some species,

these mostly parallel, but in a few species perpen-

dicular to the true veins. Sori terminal on the veins.

Indusium mostly tubular, the mouth bilabiate or

entire and often flaring, or very rarely nearly bi-

valved, the sporangia borne on an elongate recep-

tacle. Receptacle commonly exserted beyond the

mouth of the indusium, often greatly so.

There are about 300 species in the genus, oc-

curring in tropical to subtropical regions of both

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-

ber of species may be found on wet, rocky cliffs

or occasionally on clay banks. Also as in Hymen-

ophyllum, Trichomanes has been divided by pre-

vious authors into a number of genera and/or sub-

genera. Only one genus is recognized here, with

the key and species order reflecting as nearly as

possible four of the subgenera. The monotypic subg.

Cardiomanes (Presl) Christ occurs in New Zea-

land.

"False veins" may be found on laminae in two

of the subgenera. These are long sclerenchymatous

strands which can be observed as faint to rather

distinct lines, parallel or perpendicular to the true

veins. They may be connected to the true veins

but are much more commonly free from them.

Their position and frequency are important char-

acters in recognizing some of the species. As in

Hymenophyllum, veins of several species of Tri-

chomanes bear crestlike lamellae, perpendicular

to the plane of the lamina. One of these, T. mar-

tiusii, occurs in Peru.

The group of Trichomanes crispum L. in subg.

Achomanes is the most taxonomically difficult in

the genus, and a few of these species occurring in

Peru may eventually be found to merit only infra-

specific status. Some of the difficulties are very

likely the result of yet unsolved questions involv-

ing hybridization.

References

BOER, J. G. W. 1962. The new world species of

Trichomanes sect. Didymoglossum and Micro-

gonium. Acta Hot. Neerl., 11: 277-330.

WINDISCH, P. G. 1977. Synopsis of the genus

Trichomanes, subgenus Achomanes. Ph.D. the-

sis, Department of Biology, Harvard Univer-

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 conical, apex expanded into a wide-flaring mouth; petiole 0.3-3 cm long;

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; 2—4 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 mm broad;

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

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 catadromous (or sometimes flabellate); false veins present or lacking . . . 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 erect or decumbent; leaves crowded to caespitose; fertile leaves simple and

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 linear or linear-lanceolate, mature ones commonly

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)

1800(2): 97. 1802. TYPE: Jamaica, Swartz

(holotype, s; photo, us). Figure 12a-b.

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

SYNTYPES: Peru, Huanuco, Pampayacu, Poep-

pig 1132 (K!; photo, us); Peru, Junin, Pangoa,

Mathews 1088 (K.!; photos. A, us); Venezuela,

Merida, Linden 176 (K.!; isosyntypes, BM, n).

Trichomanes brachyblastos Kuhn, Linnaea 35: 388.

1868. TYPE: Peru, San Martin, Tarapoto, Monte

Guayrapurima, Spruce 4703 (holotype, B?; iso-

types, BM!, BR!, GH!, K!, w!).

Vandenboschia radicans (Sw.) Copel., Philipp. J. Sci.

67: 54. 1938.

Trichomanes radicans var. kunzeanum (Hooker) Duek

& Lell., Amer. Fern J. 68: 120. 1978.

Stem long-creeping, relatively stout, 1-2.2 mm

in diameter, amply to densely covered with cas-

taneous trichomes. Leaves remote, 20-50 cm long,

axes often provided with scattered septate tri-

chomes abaxially. Petiole (l-)4-15 cm long, 0.8-

2 mm in diameter, nonalate to marginate or alate

(often the wings deciduous). Lamina 2-3-pinna-

tisect, ovate or deltoid-ovate and scarcely reduced

at base, or occasionally elliptic and somewhat

strongly reduced toward base. Rachis narrowly to

broadly alate. Pinnae mostly 2-pinnatisect, slight-

ly to strongly adnate to the rachis. Ultimate seg-

ments commonly narrow, often linear, with simple

or 1 -forked veins, false veinlets lacking. Venation

anadromous. Indusia very slender, cylindrical or

elliptic, rarely subconical, often subfusiform, not

or scarcely alate, the wings of tissue (if any) 1-

2(-3) cells wide, apex truncate, not expanded, re-

ceptacle short- or long-exserted.

Hemiepiphytic on tree trunks in dense, wet for-

ests, (400-)700-3200 m, San Martin to Puno.

Mexico to Panama; West Indies; Guianas to Co-

lombia and southward to Brazil and Paraguay; Old

World.

Trichomanes radicans, T. collariatum, and T.

rupestre form a close-knit species complex which

is in need of closer study. The three are only pro-

visionally maintained here at the species level. In

Central America T. radicans and T. collariatum

are more distinct, e.g., leaves of the latter are con-

sistently reduced to a very short and broadly alate

petiole; whereas in T. radicans leaves are not or

scarcely reduced at base, and petioles are much

longer and not or very narrowly alate or marginate.

These characters are also somewhat diagnostic in

South America, but are much more variable, and

are therefore not thoroughly reliable. In Peru, often

the two species can be positively separated only

by the soral features.

San Martin: Prov. Mariscal Caceres, Dist. Tocache

Nuevo, J. Schunke 4361 (F, us). Huanuco: Near Tingo

Maria, confluence of Monzon and Huallaga Rivers, Stork

& Norton 9501 (F, GH, MO, uc, us). Pasco: Paujil near

Puerto Bermudez, Leon 288 (GH, USM). Junin: Chan-

chamayo Valley, C. Schunke 459 (F, us). Ucayali: Prov.

Coronel Portillo, Sinchono, between Tingo Maria and

Pucallpa, Aguilar 883 (GH). Ayacucho: Prov. La Mar,

between Tambo San Miguel, Ayna and Hacienda Lu-

isiana, Dudley 11904 (GH). Cuzco: Prov. Paucartambo,

bosques de Keros, Vargas 7382 (uc). Puno: Prov. Car-

abaya, Hacienda Palmera, Vargas 16137 (GH).

2. Trichomanes collariatum Bosch, Ned. Kruidk.

Arch 4: 368. 1859. TYPE: Mexico, Tabasco,

Linden (holotype, L?; probable isotype, K;

photo, us of K).

Stem long-creeping, relatively stout, 1-1.7 mm

in diameter, amply to densely covered with cas-

taneous trichomes. Leaves remote, 1 5-38(-45) cm

long, axes often provided with scattered, septate

trichomes abaxially. Petiole 0.3-3 cm long, 0.7-

1.5 mm in diameter, broadly alate nearly or quite

throughout. Lamina 2-3-pinnatisect, oblong or

oblong-lanceolate, often abruptly reduced at base.

Rachis commonly broad-alate throughout. Pinnae

mostly 2-pinnatisect, slightly to strongly adnate to

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

81

rachis. Ultimate segments commonly narrow, often

linear, with simple or forked veins, false veinlets

lacking. Venation anadromous. Indusia conical,

gradually widening from base to a conspicuously

flaring mouth, alate, the wings of tissue on each

side 4-6 cells wide, receptacle long-exserted.

Hemiepiphyte on tree trunks in wet forests, sea

level to 500 m, San Martin, Loreto, Madre de

Dios.

Southern Mexico to Panama; British Guiana to

Colombia, south to Peru; Brazil.

This is not greatly distinct from T. radicans,

under which see further discussion.

San Martin: Prov. Mariscal Caceres, Dist. Campan-

illa, left margin of Rio Huallaga, J. Schunke 4120 (F, GH,

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

(us). Madre de Dios: Prov. Tambopata, Lago Tres Chim-

badas, Barbour 5669 (MO). Parque Nacional del Manu,

Cocha Cashu Biological Station, M. S. Foster P-84-4

(uc), P-84-67 (MO); R. Foster 6868 (F).

3. Trichomanes rupestre (Raddi) Bosch, Ned.

Kruidk. Arch. 4: 370. 1859.

Hymenophyllum rupestre Raddi, PI. Bras. nov. gen.

1 : 67, /. 80. 1 825. TYPE: Brazil, Raddi (holotype,

FI?; isotype, K.; photos, A & us of K).

Stem long-creeping, stout, 0.7-1.2 mm in di-

ameter, provided with a few scattered to sparse

castaneous trichomes. Leaves remote, mature ones

15-30 cm long, axes sometimes provided with

scattered, septate trichomes on abaxial side. Pet-

iole 0.2-2 cm long, 0.6-0.8 mm in diameter, par-

tially alate. Lamina commonly 1-pinnate-pinna-

tifid, oblong or elliptic-oblong. Rachis broadly alate

throughout. Pinnae pinnatifid or pinnatisect, or

rarely 2-pinnatifid as to basal segments, strongly

adnate to rachis. Ultimate segments relatively short

and broad, not linear, with simple or forked veins,

false veinlets lacking. Venation anadromous. In-

dusia conical, gradually widening from base to a

slightly flaring mouth, narrowly alate on each side,

receptacle short- to long-exserted.

Creeping on tree trunks or wet banks in dense,

wet forests, 1 100-1600 m, Huanuco and Ucayali.

Venezuela and Colombia; Peru; Bolivia; Brazil

(also reported from Costa Rica by Alan Smith, in.

litt).

This is not as common as Trichomanes radicans

or T. collariatum and is midway between them in

shape of indusia. The indusium in the latter is

conical, with a conspicuous and widely flaring

mouth; in T. rupestre it is narrower, with mouth

only slightly flaring, and in T. radicans the indu-

sium is even more slender, often narrowing slight-

ly, with the apex truncate, not or scarcely spread-

ing. See T. radicans for further discussion.

Huanuco: Prov. Huanuco, Dist. Churubamba, Cotir-

arda, Mexia 8221 (F, GH, MO, uc, us). Tingo Maria, Selva

Real, Morrow 11132 (GH). Ucayali: Prov. Coronel Por-

tillo (as Loreto), Aguaytia, Ridoutt 13082 (USM).

4. Trichomanes pyxidiferum L., Sp.pl. 1098. 1753.

TYPE: based on Plumier, Traite foug. Amer.,

/. 50E, 1705, illustrating a plant from Haiti.

Vandenboschia pyxidifera (L.) Copel., Philipp. J. Sci.

67: 53. 1938.

Stem long-creeping, filiform, 0.4-0.5 mm in di-

ameter (excluding indument), densely covered with

blackish trichomes. Leaves well-spaced to remote,

3-12 cm long, axes sometimes provided abaxially

with scattered, minute trichomes. Petiole 1-3 cm

long, 0.3-0.6 mm in diameter, flattened, or terete

at base, narrowly to broadly alate distally or

throughout, the wings plane to somewhat crispate.

Lamina 2-3-pinnatifid. Rachis alate throughout,

the wings commonly undulate to crispate. Pinnae

4-10 pairs, strongly adnate to the rachis (the costae

alate). Ultimate segments linear, margins plane to

undulate, tissue commonly with elongate, narrow

folds parallel to the veins, false veinlets lacking.

Venation anadromous. Sori 1 -several per pinna.

Indusia broadly conical, the tube 1-1.5 times as

long as broad, not or scarcely bilabiate, the mouth

not or slightly flaring, receptacle commonly long-

exserted.

On trees in wet forests, 900-1 200 m, Amazonas,

San Martin.

Pantropical.

A distinctive feature of Trichomanes pyxidifer-

um is the rather sharp, longitudinal folding of seg-

ment tissue parallel to the veins. Other related

species have plane segments, or some with the

margins undulate, but none exhibit this unique

character. Perhaps it was this which prompted

Morton's incorrect observation (1968) of false veins

in the species, since the sharper folds of tissue

sometimes cause dark shadow lines in the seg-

ments. However, in careful study of specimens

throughout the range, no leaves were seen with the

82

FIELDIANA: BOTANY

sclerenchymatous strands which produce these

"false veins." The species thus far is represented

in Peru by only two collections, even though its

full range extends throughout tropical regions of

the world.

Amazonas: Prov. Chachapoyas, .la/an (Ingenio-

Chachapoyas), Ldpe: el al. 4264 (GH, HUT). San Martin:

Tarapoto, Spruce 4761 (BR, OH, K, L, us).

5. Trichomanes diaphanum HBK., Nov. gen. sp.

1:25 (fol. 1 6). 1 8 1 6. TYPE: Venezuela, Hum-

boldt & Bonpland (holotype, p; isotype, B!).

Trichomanes leptophyllum Bosch, Ned. Kruidk. Arch.

4: 363. 1859 (not A. Cunn., 1836), nom. illeg.

TYPE: based on T. pyxidiferum Hooker & Grev.

Trichomanes hymenophylloides Bosch, Ned. Kruidk.

Arch. 5: 209. 1863, nom. nov. for T. leptophyllum

Bosch and type based on that name.

Vandenboschia diaphana (HBK.) Copel., Philipp. J.

Sci. 67: 53. 1938.

Vandenboschia hymenophylloides (Bosch) Copel.,

Philipp. J. Sci. 67: 53/1938.

Stem long-creeping, filiform, 0.2-0.5 mm in di-

ameter (excluding indument), sparsely to abun-

dantly provided with castaneous trichomes. Leaves

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

glabrous, or the rachis sometimes abaxially pro-

vided with scattered, minute trichomes. Petiole

0.5-6 cm long, 0.3-0.8 mm in diameter, flattened,

or terete at base, nonalate, or narrowly to broadly

alate for most of its length, the wings plane or

occasionally undulate. Lamina essentially 3-pin-

natifid. Rachis alate throughout, the wings narrow

to broad, plane to undulate or occasionally cris-

pate. Pinnae 4-12 pairs, adnate to the rachis, the

costae alate (rarely nonalate at base). Ultimate seg-

ments linear, 0.6-0.9 mm broad, plane, not folded,

false veinlets lacking. Venation anadromous. Sori

1-10 per pinna. Indusia salverform, the alate tube

narrowly cylindrical, 2-4 times as long as broad,

abruptly expanding into a broad-flaring mouth,

receptacle commonly long-exserted.

On trees, moist banks and (rarely) wet rocks, in

deep forests 700-2900 m, Amazonas and Loreto

to Puno.

Southern Mexico to Panama; West Indies; Trin-

idad; the Guianas to Colombia, south to Peru and

Brazil.

Previous authors have separated Trichomanes

hymenophylloides on characters such as number

of sori per pinna, width of tissue wings flanking

axes and indusia, crowding of pinnae and dissec-

tion of lamina. Each of these may appear to be

important in individual specimens, but when many

collections are examined throughout the range of

distribution, it becomes evident that they are quite

variable and uncorrelated.

Amazonas: Prov. Bagua, Cordillera Colan SE of La

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 in part). Junin: La Merced, E of Quimiri Bridge,

Killip & Smith 24006 (F), 24023 (GH, us). Cuzco: Prov.

La Convencion, "El Dorado," Vargas 3518 (GH). Puno:

San Gaban, Lechler (L).

6. Trichomanes angustatum Carm., Trans. Linn.

Soc. London 12: 513. 1818. TYPE: Tristan

da Cunha, Carmichael (holotype, K!; isotype,

BM!).

Trichomanes tenerum Sprengel, Syst. veg. 4: 1 29. 1 827.

TYPE: Brazil, collector not stated (holotype, LZ

destroyed).

Vandenboschia tenera (Sprengel) Copel., Philipp. J.

Sci. 67: 53. 1941.

Vandenboschia angustata (Carm.) Copel., Philipp. J.

Sci. 73: 466. 1941.

Stem long-creeping, filiform, 0.2-0.3 mm in di-

ameter, amply provided with castaneous tri-

chomes. Leaves well-spaced to remote, 4-14(-18)

cm long, glabrous. Petiole 0.5-5 cm long, 0.2-0.4

mm in diameter, terete, nonalate. Lamina 2-pin-

nate-pinnatifid to 4-pinnate, linear-lanceolate to

ovate-lanceolate, abruptly or gradually reduced at

base. Rachis nonalate, or scarcely alate near the

apex. Pinnae 6-many pairs, short-stalked, the cos-

tae alate except below the basal pinnule. Ultimate

segments linear, 0.5-0.8 mm broad, each bearing

a single vein, false veinlets lacking. Venation anad-

romous. Sori 1-5 per pinna. Indusia narrowly fun-

nelform to salverform, the tube 2-4 times as long

as broad, the mouth flaring, margins commonly

alate, with wings several cells wide, receptacle long-

exserted.

In deep forests, on trees, wet cliffs and banks,

and on decaying logs, 600-2800 m, Cajamarca and

San Martin to Ayacucho and Cuzco.

Southern Mexico to Honduras; Greater Antilles;

Venezuela and Colombia to NW Argentina and

Brazil.

This is very closely related to Trichomanes cap-

illaceum, under which see further discussion.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

83

Cajamarca: Colasay, Woytkowski 7002 (us). San Mar-

tin: Monte Campana, near Tarapoto, Spruce 4705 (BR,

K, NY). Pasco: Prov. Oxapampa, San Alberto, Cordillera

de Yanachaga, van der Werffet al. 8428 (MO, uc). Junin:

Carpapata, above Huacapistana, Killip & Smith 24492

(F, uc, us). Ayacucho: Ccarrapa, between Huanta and

Rio Apurimac, Killip & Smith 22406 (F, GH, us). Cuzco:

Valley of Urubamba, Machu-Picchu, base of Huayna

Picchu, Iltis el al. 1064 (GH, MO, uc, us).

7. Trichomanescapillaceum L., Sp. pi. 1099. 1753.

LECTOTYPE (designated by Proctor, Ferns

of Jamaica, British Museum, London, p. 109.

1985): Plumier, Traite foug. Amer., t. 99D,

based on a Plumier collection from Haiti.

Vandenboschia capillacea (L.) Copel., Philipp. J. Sci.

67: 53. 1938.

This differs from Trichomanes angustatum only

in the characters used in the key.

In deep forests, on tree trunks, 800-1100 m,

Pasco and Cuzco.

Southern Mexico to Panama; Greater Antilles;

Venezuela and Colombia to Peru.

With this might possibly be included Trichom-

anes angustatum, but the two are separated here

provisionally, pending much needed monographic

revision. Typically, T. capillaceum has very little

expanded tissue along the axes, and even the ul-

timate segments are rarely more than 0.3 mm

broad. Indusia are not or scarcely alate, and son

are solitary on each fertile pinna. Conversely, be-

yond the first pinnule in T. angustatum, the costae

and costules are obviously alate throughout, and

ultimate segments are often nearly 1 mm broad.

The tissue also forms wings on each side of the

indusia, and sori are frequently four or five per

pinna.

Even some of the characters listed in the key are

subject to variation in certain geographic regions,

and intermediates are found frequently. Never-

theless there are areas in which only one or the

other occur. Only Trichomanes angustatum oc-

curs in Bolivia and southern Brazil, whereas in

Central America it has not been found south of

Guatemala, a region where T. capillaceum is rath-

er common. In Peru, the latter is rare, and even

the few specimens located are not typically skeletal

in their appearance, whereas Trichomanes angus-

tatum is rather widespread and the specimens

rather typical. One specimen from Cuzco, Vargas

8653 (MO), is exactly intermediate between the

two; significantly, Cuzco is one of the two de-

partments in which both species are known to oc-

cur. Therefore, based only on collections from Peru,

one would be tempted to combine the two species,

but a thorough study of the characters of both taxa

throughout their range is necessary before further

classification is attempted.

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).

Trichomanes elegans Rich., Actes Soc. Hist.

Nat. Paris 1: 114. 1792. TYPE: French

Guiana, Le Blond (P) (not T. elegans Rudge,

1805).

Trichomanes prieurii Kunze, Analecta Pteridogr. 48.

1837. SYNTYPES: French Guiana, Leprieur (LZ

destroyed; isosyntype, F!); Brazil, Amazonas, Rio

Japura, Martius (M); British Guiana, Rudge (K).

Trichomanes opacum Bosch, Ned. Kruidk. Arch. 5(2):

1 75. 1 86 1 . TYPE: Peru, Puno, San Gaban, Lech-

ler 2175 (holotype, L?; isotype, P!; photo, F, frag.,

L!).

Davalliopsis elegans (Rich.) Copel., Philipp. J. Sci. 67:

82. 1938.

Terrestrial, rarely epiphytic or epipetric. Stem

erect or rarely short-creeping. Leaves caespitose to

occasionally approximate, 20-90 cm long, rachis,

costae, and often costules amply to abundantly

provided with castaneous, pluricellular trichomes

on abaxial side. Petiole 10-40 cm long, 1-3.5 mm

in diameter, usually alate distally. Lamina 2-3-

pinnate-pinnatifid, deltoid, opaque, usually more

than 1 cell thick, at least away from the margins,

the cells small and occluded, not evident even when

magnified at 10-15x. Rachis obtusely quadran-

gular, alate (often broadly so) at least in distal half.

Pinnae adnate, the costae alate to base. Ultimate

segments laciniate, with linear lobes, each of these

bearing a single vein, false veins lacking. Venation

anadromous. Sori usually bent down away from

the plane of the lamina. Indusia subconical to

somewhat urceolate, not bilabiate, the mouth

truncate or slightly (rarely strongly) flaring, recep-

tacle short- to long-exserted.

In dense, wet forests or wet, wooded ravines, on

ground, or rarely on tree trunks or wet rocks, sea

level to 1300 m, Amazonas and Loreto south to

Madre de Dios and Puno.

Lesser Antilles and Trinidad; Nicaragua to Pan-

ama; Guianas to Colombia and south to Bolivia

and Brazil.

84

FIELDIANA: BOTANY

With its large, blue green leaves and laciniate

pinnae, this is one of the most beautiful species in

the family. Its most distinctive features are the

usually strongly arching sori (bent downward out

of the plane of the lamina) and the relatively thick

tissue. Although the lamina in Hymenophyllaceae

is typically only one cell thick, the tissue in Tri-

chomanes elegans is often several cells thick, es-

pecially away from the segment margins. The

broadly alate rachis traditionally has been used to

separate this from T. rigidum (with which it often

grows), and although usually it is a good diagnostic

feature, occasional specimens of T. elegans can be

found throughout the range in which rachis alae

are very narrow or lacking in the proximal portion.

Consequently, supporting key characters should

be employed for accurate identification.

Amazonas: Prov. Bagua, left bank of Rio Maranon

opposite Quebrada Mirana, Wurdack 2031 (F, GH, us).

San Martin: Prov. Mariscal Caceres, Tocache Nuevo, J.

Schunke4855 (F, GH, us), 6955 (F, MO in part, us). Loreto:

Sierra del Pongo, Mexia 1 195 (F, GH, MO, us). Huanuco:

Prov. Pachitea, Dist. Honoria, /. Schunke 2259 (F, GH).

Pasco: Prov. Oxapampa, Valle del Palcazu, Iscozacin,

Le6n 692 (F, GH). Junin: Puerto Bermudez, Killip & Smith

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-

bopata, Rio Tambopata Nature Reserve, Barbour 5165

(MO). Puno: Prov. Carabaya, San Gaban, Vargas 18864

(GH).

9. Trichomanes rigidum Swartz, Prodr. 137. 1788.

TYPE: Jamaica, Swartz (holotype, s; isotype,

B, Herb. Willd. 20202- 7; photos, us of s, GH

& us of B).

Selenodesmium rigidum (Sw.) Copel., Philipp. J. Sci.

67:81. 1938.

Terrestrial, rarely epiphytic or epipetric. Stem

erect or occasionally short-creeping. Leaves caes-

pitose to sometimes approximate, 1 5-35 cm long,

lamina glabrous, the axes glabrous, or rarely the

rachis or petiole base with a few, scattered tri-

chomes. Petiole 5-16 cm long, 0.6-1.6 mm in di-

ameter, nonalate. Lamina 3-4-pinnate-pinnatifid,

deltoid, opaque, 1 cell thick, but cells small and

occluded, not evident even when magnified at 1 0-

1 5 x . Rachis terete, nonalate, or marginate to

slightly alate toward apex. Pinnae (at least prox-

imal ones) short-stalked and the costae not alate

at base. Ultimate segments with linear lobes, each

of these bearing a single vein, false veins lacking.

Venation anadromous. Sori (most of them) borne

essentially in the plane of the lamina. Indusia sub-

conical, not bilabiate, the mouth truncate, not or

scarcely flaring, receptacle commonly long-exsert-

ed.

On slopes and ridges of dense forests and in

wooded ravines, occasionally on wet rocks or cliffs,

rarely on bases of tree trunks, 350-2500 m, Ca-

jamarca and San Martin, south to Cuzco and Madre

de Dios.

Southern Mexico to Panama; West Indies; South

America, to Bolivia and Brazil; probably Old World

tropics.

This is sometimes confused with Trichomanes

elegans, with which it often grows. For compari-

son, see discussion under the latter species.

Cajamarca: Tambillo, Jelski 891 (us). San Martin:

Tarapoto, in monte Guayrapurima, Spruce 4047 (BR, K).

Huanuco: East of Tingo Maria (as San Martin), Allard

22356 (GH, us). Pasco: Oxapampa, Cordillera San Ma-

tias, Leon 326a (USM). Junin: La Merced, Hacienda

Schunke, Macbride 5633 (F, us). Ucayali: Vicinity of

Aguaytia, along Rio Aguaytia (as Loreto), Croat 20927

(MO). Cuzco: Prov. Urubamba, base of Huayna Picchu,

Iltis et al. 1062 (GH). Madre de Dios: Prov. Manu, Cerro

de Pantiacolla, Foster 10903 (F).

10. Trichomanes cellulosum Klotzsch, Linnaea 18:

531. 1844. TYPE: "British Guiana," (Guy-

ana). Kanuku Mountains, Rich. Schomburgk

1186 (holotype, B!; photo, F; isotypes, B! 2

sheets, BM!).

Epiphytic, terrestrial, or sometimes epipetric.

Stem erect to short-creeping. Leaves caespitose to

(occasionally) approximate, 6-15 cm long, gla-

brous. Petiole 1-7 cm long, 0.3-0.6 mm in di-

ameter, narrowly alate at least distally. Lamina 4-

5-pinnate, lanceolate to ovate, translucent, the cells

large and clear and quite evident when magnified

at 8-12x. Rachis terete, alate throughout. Pinna

divisions skeleton-like, the costae, costules, and

ultimate segments with bands of tissue only 1-

several rows wide. Ultimate segments linear, 0.3-

0.7 mm broad, each bearing a single vein, false

veins lacking. Venation anadromous. Indusia sub-

conical to somewhat urceolate, not bilabiate, the

mouth truncate or slightly flaring, receptacle short-

to long-exserted.

In wet forests, on trees or on the forest floor,

sometimes on wet rocks, 300-1 300 m, Amazonas,

Loreto, Huanuco.

Surinam to Colombia; Peru; Brazil.

With this perhaps should be included Trichom-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

85

anes sprucei Baker of northern South America,

which is probably only a more robust form, with

broader segments and broader bands of tissue along

the axes.

Amazonas: Prov. Bagua. roadside from Chiriaco to

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

mayacu, between Balsapuerto and Moyobamba, Klug

3192 (F, GH, MO, us). Tierra Doble on Rio Nanay, LI.

Williams 1057 (F, us). Huanuco: Rio Llullapichis wa-

tershed, ascent of Cerros del Sira, Dudley 13032 (GH),

Wolfe 12352 (GH, us).

1 1 . Trichomanes punctatum Poiret ssp. sphen-

oides (Kunze) Boer, Acta Bot. Neerl. 1 1: 301.

1962.

Trichomanes sphenoides Kunze, Farrnkrauter 216, /.

88, f. 2. \ 840. TYPE: Peru, Cuchero (Dept. Huan-

uco), Poeppig (holotype, w!; isotype, MO!).

Didymoglossum sphenoides (Kunze) Presl, Hymeno-

phyllaceae 23. 1843.

Stem long-creeping, filiform. Leaves approxi-

mate or widely spaced, to 1.5 cm long and 1 cm

broad, subsessile or short-petiolate. Lamina cir-

cular to obovate or spathulate, margin entire to

irregularly lobed, a costa lacking or, in some fertile

laminae, distinct only in the proximal portion, gla-

brous except for the dark, stellate, marginal tri-

chomes. Venation flabellate, the veins crowded,

with false veinlets parallel to them. Sori several,

or sometimes solitary, partially immersed in the

lamina tissue, commonly borne between lobes, thus

rarely extending beyond the outline circumscribed

by the lobe apices. Indusia narrow-cylindrical, bil-

abiate, the lips mostly wide-flaring and narrowly

dark-margined, receptacle not or scarcely exsert-

ed.

In dense, wet forests and wooded ravines, on

tree trunks, 100-1 100 m, Loreto to Pasco.

Guatemala; Costa Rica; Panama; Greater An-

tilles; Venezuela; Colombia to Bolivia.

Boer (1962) recognized four subspecies of Tri-

chomanes punctatum, and only this one occurs in

Peru. Others differ from ssp. sphenoides in having

more distinctly lobed laminae and/or larger and

more broadly dark-margined indusium lips.

Loreto: Maynas. Iquitos, Ushpa-Cana across from Rio

Itaya, McDaniel 11381 (GH, MO). Huanuco: Prov. Huan-

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

Junin), Killip & Smith 26592 (us).

1 2. Trichomanes angustifrons (Fee) Boer, Fl. Neth.

Antill. I (Pterid.): 17. 1962.

Didymoglossum angustifrons Fee, Mem. foug. 11: 113,

t. 28, f. 5. 1866. TYPE: Guadeloupe, I'Herminier

(holotype, P!).

Stem long-creeping, filiform. Leaves approxi-

mate or well-spaced, 0.5-1.5 cm long, to 0.6 cm

broad, subsessile or short-petiolate. Lamina nar-

rowly or broadly oblong, entire, or distally lobed,

costa percurrent, glabrous except for the dark, stel-

late trichomes borne along the margin. Venation

catadromous, pinnate, with false veinlets sparsely

to amply distributed between the true veins. Sori

solitary at lamina apex, or few and terminating

each distal lobe, somewhat to deeply immersed in

the tissue. Indusia narrow-conical, bilabiate, the

lips mostly wide-flaring and narrowly dark-mar-

gined, receptacle commonly exserted.

Thus far represented in Peru by a single speci-

men from Huanuco, on ridge in jungle, 850 m;

elsewhere apparently always epiphytic.

Costa Rica and Cocos Island; West Indies; Trin-

idad and Tobago; the Guianas to Brazil; Peru;

Paraguay (reported from Chiapas, Mexico by Alan

Smith, in litt.).

Collections of this species are rather sparse, ex-

cept from Brazil and the West Indies; yet it is

surely a much more common plant than the spec-

imens would indicate. Since it is such a tiny and

inconspicuous fern, it certainly has been over-

looked and probably is not accurately represented

in herbaria.

Huanuco (as San Martin): E of Tingo Maria, Allard

21383a (us).

1 3. Trichomanes hymenoides Hedwig, Fil. gen. sp.,

/. 3,f. 3. 1799. LECTOTYPE (designated by

Boer, Acta Bot. Neerl. 11: 306. 1962): Hed-

wig, t. 3, f. 3, probably based on a Swartz

specimen from Jamaica. Figure 12d.

Trichomanes muscoides Sw., J. Bot. (Schrader) 1 800(2):

95. 1802. TYPE: Jamaica, Swartz (holotype, S-PA).

Didymoglossum hymenoides (Hedwig) Desv., Mem.

Soc. Linn. Paris 6: 330. 1827.

Didymoglossum muscoides (Sw.) Desv. Mem. Soc.

Linn. Paris 6: 330. 1827.

Stem long-creeping, densely covered with dark

trichomes that extend onto the petioles. Leaves

86

FIELDIANA: BOTANY

well-spaced, (0.5-)l-3.5 cm long, to 1.5 cm broad,

subsessile or short-petiolate. Lamina obovate to

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

ta percurrent (but juvenile laminae often subren-

iform and merely lobed and the costa indistinct

distally), glabrous except for the dark, marginal

trichomes which are stellate in sinuses and simple

to bifid on outer margin, laminar cells commonly

isodiametric. Venation catadromous, commonly

pinnate (occasionally subflabellate in juvenile

leaves), false veinlets sparse, scattered between true

veins, extending toward lamina margin but not

parallel to it. Sori 1-several borne near lamina

apex, conspicuously exserted, or only slightly im-

mersed at base, not or very narrowly alate. Indusia

narrow-funnelform, bilabiate, but the lips quite

short, commonly broader than long, and usually

(not always) dark-margined, receptacle long-

exserted.

In forests on trees, or rarely on rocky or sandy

soil, 700-2400 m, Amazonas, Loreto.

Southern Mexico to Panama; West Indies; Trin-

idad; Venezuela and Colombia, south to Argentina

and Uruguay.

Juvenile leaves are often nearly reniform and

shallowly lobed, with veins subflabellate and the

costae sometimes indistinct toward the apex. Hence

they can be confused with Trichomanes puncta-

tum. However, fertile leaves are pinnatifid, with

stellate trichomes borne only in the sinuses, and

son strongly protrude from the leaf tissue. In T.

punctatum ssp. sphenoides sori are partially im-

mersed, and marginal trichomes are consistently

stellate.

Amazonas: Prov. Chachapoyas, Rio Ventilla W of

Molinopampa, Wurdack 1513 (F, OH, us). Loreto: Sierra

del Pongo, Mexia 6289B (GH, us).

14. Trichomanes reptans Sw., Prodr. 136. 1788.

TYPE: Jamaica, Swartz (holotype, S-PA).

Didymoglossum reptans (Sw.) Presl, Hymenophylla-

ceae 23. 1843.

Stem long-creeping, densely covered with dark

trichomes that extend onto the petioles. Leaves

well-spaced, 2-9 cm long, 1—4 cm broad, subsessile

to short-petiolate. Lamina lanceolate to ovate or

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

rachis distinct throughout, glabrous except for the

dark, marginal trichomes which are stellate in si-

nuses and simple to bifid on outer margins, lam-

inar cells mostly elongate. Venation catadromous,

pinnate, false veinlets commonly abundant, ex-

tending toward lamina margin but not (or very

rarely) parallel to it. Sori 1 or a few, borne near

lamina apex, conspicuously protruding from the

leaf tissue, or only slightly immersed at base, not

or scarcely alate. Indusia narrow-funnelform, bil-

abiate, the lips dark-margined, and sometimes

flaring, most of them as long as or longer than

broad, receptacle short- to long-exserted.

On wet rocks in forests, 1900-2750 m, Ama-

zonas, Huanuco.

Southern Mexico to Panama; Greater Antilles;

Venezuela; Colombia; Ecuador; Peru; Brazil; Ar-

gentina.

Although found thus far in Peru only on wet

rocks, Trichomanes reptans may be expected also

on trees, since it frequently has been reported as

an epiphyte throughout much of its range.

Amazonas: Prov. Bagua, Cordillera Colan SE of La

Peca, Harbour 3903 (MO, USM). Huanuco: Mito, Bryan

392 (F).

15. Trichomanes krausii Hooker & Grev., Icon,

fil. 2, /. 149. 1831. TYPE: Dominica, Kraus

(holotype, E).

Didymoglossum krausii (Hooker & Grev.) Presl, Hy-

menophyllaceae 23. 1843.

Stem long-creeping, densely covered with dark

trichomes which extend onto the petioles. Leaves

well-spaced, 2-8 cm long, 1-3 cm broad, subsessile

to short-petiolate. Lamina narrow-ovate to -ellip-

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-

vided with simple to bifid trichomes, but with

stellate ones in the segment sinuses. Venation ca-

tadromous, pinnate, false veinlets seldom numer-

ous, many of them parallel to and very near the

lamina margin. Sori 1 or a few, terminating the

segments, partly to deeply immersed in the seg-

ment tissue, the protruding portion narrowly or

broadly alate. Indusia salverform, bilabiate, the

lips commonly dark-margined and flaring, recep-

tacle short- to long-exserted.

Found near 800 m, San Martin, Huanuco. There

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

occurs in forests on trees or wet rocks elsewhere

throughout its range.

United States (Florida); Mexico to Panama; West

Indies; Trinidad; the Guianas to Colombia, south

to Paraguay and Argentina.

Besides the characters used in the key, Trichom-

anes krausii often can be distinguished from closely

related species by the dark trichomes that fre-

quently extend up the petiole onto the abaxial side

of the lower rachis. Rachises of the other species

are essentially glabrous.

San Martin: Tarapoto, Spruce 3991 (BR, w). Huanuco:

Fundo Chela, Sinchono, Rio Chino, Aguilar 912 (USM).

16. Trichomanes kapplerianum J. W. Sturm, Fl.

bras. 1(2): 276. 1859. TYPE: Surinam, near

Station Victoria, Kappler 1760 (holotype, w!).

Hemiphlebium kapplerianum (J. W. Sturm) Prantl,

Hymenophyllaceae 46. 1875.

Trichomanes ekmanii Boer, Acta Bot. Neerl. 11: 319.

1962. TYPE: Dominican Republic, La Cumbre,

Cordillera Central, Ekman H- 14342 (holotype,

u; isotypes, F!, G, GH!, MO!, s, uc, us!).

Stem long-creeping, filiform, sparsely to abun-

dantly provided with brown trichomes which ex-

tend onto the petioles. Leaves well-spaced, to 3

cm long and 1.5 cm broad, subsessile or short -

petiolate. Lamina obovate or oblong (or in juve-

nile ones often circular), base rounded, cordate or

cuneate, entire or distally lobed, with a distinct

costa at least in the proximal portion, glabrous, or

sparsely pubescent basally along the costa. Vena-

tion catadromous, essentially pinnate (but some-

times subflabellate distally), false veins ample to

abundant, parallel to the true veins and with a

submarginal, continuous or sometimes interrupt-

ed false vein around the perimeter of the lamina.

Sori several to ample near lamina apex or on the

lobes, fully immersed in the tissue. Indusia fun-

nelform, not bilabiate, the mouth expanded, but

not dark-margined, receptacle short- or long-

exserted.

In dense forests and wooded ravines, on tree

trunks (also on rocks, outside Peru), sea level to

1000 m, San Martin, Loreto, Huanuco, Puno.

Guatemala to Panama; West Indies; the Guian-

as to Venezuela; Peru; Brazil; Bolivia.

Detailed study of many specimens throughout

the Neotropics indicates that Trichomanes ek-

manii is not distinct. The latter was said by Boer

(1962) to differ in the uninterrupted submarginal

false vein, bordered on the outside by cubical cells,

whereas the submarginal vein of T. kapplerianum

was said to be sometimes interrupted and bor-

dered by tangentially lengthened cells. These fea-

tures prove to be minor ones that are quite vari-

able. In careful study of many specimens, including

types, one can nearly always find some interrup-

tions in submarginal false veins; furthermore, shape

of the border cells is seldom constant. Many leaves

on the type specimen of T. kapplerianum, for ex-

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.

ekmanii have at least a few leaves with somewhat

interrupted submarginal veins. There is probably

no justification for separating either species from

T. hookeri, which Boer distinguished on similar

characters. However, the type of the latter has not

been examined and T. kapplerianum is provision-

ally maintained here as distinct.

San Martin: Near Tarapoto, Spruce 4762 (BR, w). Lor-

eto: San Antonio, on Rio Ataya, Killip & Smith 29522

(F, GH). Prov. Maynas, on the Amazon, 50 miles down-

river from Iquitos, Moran 3661 (F). Huanuco: E of Tingo

Maria (as San Martin), Allard 20912 (us). Puno: Near

San "Gavan" (Gaban), Lechler 2297 (BR).

17. Trichomanes membranaceum L., Sp. pi. 1097.

1753. TYPE: "America," without specific lo-

cality (holotype, LINN 1253.1).

Lecanium membranaceum (L.) Presl, Hymenophyl-

laceae 12. 1843.

Didymoglossum membranaceum (L.) Vareschi, Flora

Venezuela 1: 222. 1969.

Stem long-creeping, slender, densely covered

with dark brown trichomes which extend onto the

short petioles. Leaves well-spaced, 2-6 cm long

and often nearly as broad, subsessile. Lamina near-

ly circular and entire, or spathulate and incised

into regular lobes, lacking a distinct midrib, gla-

brous, but bearing along the margin numerous,

paired, circular, scalelike processes (these some-

times deciduous in age). Venation flabellate, the

veins repeatedly dichotomous, false veinlets abun-

dant and parallel with the true veins, but a con-

88

FIELDIANA: BOTANY

tinuous submarginal false vein lacking. Sori sev-

eral to many on vein tips toward the lamina apex,

partially to fully immersed in the tissue. Indusia

narrow-funnelform, not or scarcely bilabiate, the

mouth neither flaring nor dark-margined, recep-

tacle short- or long-exserted.

In deep forests or wooded ravines, on tree trunks

or wet rocks or cliffs, sea level to 850 m, Ama-

zonas, Loreto, Huanuco.

Southern Mexico to Panama; West Indies; the

Guianas to Colombia, south to Bolivia.

Amazonas: Prov. Bagua, Rio Maranon opposite Que-

brada Mirana, Wurdack 2040 (F, us). Loreto: Dist. Tigre,

Rio Tigre near Tigre, McDaniel & Rimachi 18531 (GH).

Pongo de Manseriche, Mexia 6198 (F, OH, MO, us).

Huanuco: E of Tingo Maria (as San Martin), Allard 21549

(GH, us).

18. Trichomanes polypodioides L., Sp. pi. 1098.

1753. NEOTYPE (designated by Proctor,

Flora Lesser Antilles 92. 1977): Montserrat,

Proctor 19068 (A).

Trichomanes poeppigii Presl, Hymenophyllaceae 4 1 .

1843. TYPE: "Habitat in Peruvia," Poeppig(ho-

lotype, w?).

Stem long-creeping, filiform, sparsely provided

with scattered brown trichomes. Leaves well-

spaced, 4-12(-18) cm long, 1-3 cm broad, sub-

sessile or short-petiolate. Lamina linear- or ob-

long-lanceolate, deeply lobed to pinnatifid, the axes,

veins and margin provided with dark brown tri-

chomes that are stellate or forked from the base.

Venation catadromous, lacking false veins. Sori

1-few at or near segment apex, fully immersed in

the segment tissue. Indusia salverform, neither bil-

abiate nor dark-margined, the mouth flaring, re-

ceptacle long-exserted.

In deep forests, on tree trunks, very rarely on

wet clay banks, 350-2300 m, Loreto and Huanuco

to Cuzco and Madre de Dios.

Southern Mexico to Panama; West Indies; Trin-

idad; South America, to Brazil and Uruguay.

Smaller specimens could be mistaken for Tri-

chomanes reptans or other related species of subg.

Didymoglossum, which are similar in their deli-

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

marginal stellate trichomes, and general aspect.

However T. polypodioides lacks the false veinlets

common in that subgenus, and trichomes are

sparsely scattered along the stem, unlike the dense,

blackish indument found on stems of T. reptans

and its relatives.

Loreto: Sierra del Pongo, Mexia 6228 (GH, MO, uc,

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

Hacienda above San Ramon, C. Schunke A240 (GH, us).

Cuzco: Altura del Rio Tocate, Bues 1744 (us). Madre de

Dios: Prov. Manu, Cerro de Pantiacolla, Foster el al.

10713 (F).

19. Trichomanes tanaicum J. W. Sturm in Mart.,

Fl. bras. 1(2): 260. 1859. TYPE: Brazil, Para,

Rio Acara, Spruce 410 (holotype, B; possible

isotypes ["Spruce 44"], K.!, us; photo, A of K).

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

Gefasskrypt. 1: 50. 1875.

Trichomanes ankersii Hooker & Grev. var. tanaicum

(Sturm) Sadebeck in Engler & Prantl, Nat. Pflan-

zenfam. 1(4): 105. 1899.

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

in diameter, amply provided with brown rhizoids

which extend onto the rachis and adhere to tree

trunks. Leaves subdistant to remote, mature ones

3-15 cm long, 0.7-1.5 cm broad, subsessile or

scarcely petiolate. Lamina linear, subentire or lobed

(at base occasionally incised nearly to the rachis),

the rachis abaxially often provided with rhizoids,

otherwise glabrous. Venation catadromous, veins

free, 1 -several times-forked in the lobes, false veins

lacking. Sori 1 or a few, terminating the lobes, not

or scarcely immersed in the segment tissue. In-

dusia salverform, not or scarcely bilabiate, the

mouth rather widely flaring, not dark-margined,

receptacle scarcely to long-exserted.

In deep forests, commonly hemiepiphytic on

tree trunks, 100-130 m, Loreto, Ucayali.

Venezuela; Colombia; Brazil; Amazonian Peru.

The lamina in Trichomanes tanaicum is com-

monly lobed less than one-third to the rachis, but

rarely (as in Klug 92) some of the leaves are deeply

pinnatifid. Nevertheless, the narrow laminae with

their linear outline distinguish them from other

species in section Lacostea.

Loreto: Mishuyacu, near Iquitos, Klug 92 (F, us). Prov.

Maynas, Dist. Nanay, Santa Maria de Nanay, J. Schunke

V. 2449 (F, GH). Ucayali: Prov. Coronel Portillo (as Lor-

eto), Rio Mazan, J. Schunke 300 (F, GH, uc, us, USM).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

89

20. Trichomanes tuerckheimii Christ, Hedwigia

44: 361. 1905. LECTOTYPE (designated here):

Guatemala, Alia Verapaz, Cubilquitz, von

Tuerckheim 8348 (lectotype, P!; isolectotypes,

K., P!). LECTOPARATYPE: Peru, Loreto, near

Leticia, Ule 6228 (K!, P!).

Stem long-creeping, relatively stout and wiry,

0.5-1.2 mm in diameter, sparsely to abundantly

provided with brown rhizoids that extend onto the

leaf axes and adhere to tree trunks. Leaves sub-

distant to remote, larger ones (9-)12-20(-30) cm

long, 2-9 cm broad, subsessile. Lamina broadly

oblong, pinnatisect, the rachis, costae and (some-

times) veins and margins abaxially provided with

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-

lowly crenate. Venation catadromous, veins free,

simple, false veins lacking, but often some very

short, darkened lines scattered between the veins.

Sori few to several on the pinna-lobes, not or

scarcely immersed in the segment tissue. Indusia

tubular, neither bilabiate nor dark-margined,

mouth not expanded, receptacle commonly long-

exserted.

Thus far known in Peru only from the lecto-

paratype, cited above, on tree trunks; elsewhere

hemiepiphytic, in forests on tree trunks, sea level

to 500 m.

Southern Mexico to Panama; Surinam to Co-

lombia; Amazonian Peru.

Included on the sheet with the syntype at Paris

is a fragment of a 2-pinnatifid leaf with nearly

laciniate lobes, which might have been mounted

here by mistake. It resembles the more highly di-

vided specimens which have been described as

Trichomanes pedicellatum and T. subsessile. Re-

lationships with these are discussed under T. an-

kersii. (Also described there are the dark lines in

the lamina which might be mistaken for the false

veins of sect. Didymoglossum.)

21. Trichomanes ankersii Hooker & Grev., Icon,

fil. 2, t. 201. 1831. TYPE: Guyana, Demerara,

Ankers (holotype, K; photo, us).

Stem long-creeping, relatively stout and wiry,

0.5-1.2 mm in diameter, sparsely to abundantly

provided with brown rhizoids that extend onto the

leaf axes and adhere to tree trunks. Leaves sub-

distant to remote, larger ones 5-18 cm long,

(1.8-)2-6 cm broad, subsessile. Lamina oblong,

pinnatisect, or pinnate to a broadly winged rachis,

the rachis, costae, and (sometimes) veins abaxially

provided with simple, reddish brown trichomes

that adhere to tree trunks, free portion of larger

segments 1.5-2.8(-3) cm long and 0.5-0.8 cm

broad, deeply crenate or crenate-serrate. Venation

at least partly catadromous, veins free, commonly

simple, but some- to many-forked, false veins

lacking, but often some very short, darkened lines

scattered between the veins. Sori few to several on

the pinna lobes, not or scarcely immersed in the

segment tissue. Indusia tubular, neither bilabiate

nor dark-margined, mouth scarcely to slightly ex-

panded, receptacle commonly long-exserted.

In forests, commonly hemiepiphytic, tightly ap-

pressed to tree trunks, sea level to 1 000 m, Loreto,

Huanuco, Pasco, Cuzco.

The Guianas to Colombia, south to Brazil and

Bolivia.

This, Trichomanes tuerckheimii, and T. tanai-

cum, all members of sect. Lacostea, are commonly

hemiepiphytic. Their leaves usually have the dis-

tinctive habit of being tightly appressed to tree

trunks. They cling to the bark by means of rust-

colored, prehensile trichomes that are abundant

along the stems, rachises, and in T. ankersii and

T. tuerckheimii, on midribs and veins. Some col-

lectors' labels describe them as "plastered to the

bark of trees." Another character peculiar to the

three species (though very rare in T. tanaicum) is

the presence of very short, dark lines seen scattered

between and parallel with the veins. They some-

what resemble the false veins of sect. Didymo-

glossum, except that they are not lengthy scler-

enchymatous strands. Rather, they merely appear

to be a series of (usually) two or three constricted

and occluded cells that are mostly found toward

the segment margin.

Very closely related to Trichomanes ankersii are

T. pedicellatum Desv. and T. subsessile Splitg., of

northern South America and the West Indies. These

appear to differ only in their more highly divided

sterile lamina, which is 2- to 3-pinnatifid. Further

study may prove them all to be conspecific.

Loreto: Mishuyacu, near Iquitos, Klug 149, 1135 (F,

us). Sierra del Pongo, Mexia 6289 (GH, MO, uc, us).

Huanuco: 5 km NE of Tingo Maria, Stork & Horton

9518 (F, GH, uc, us). Pasco: Prov. Oxapampa, Valle del

Palcazu, Leon 706 (F, GH). Cuzco: Prov. Quispicanchi,

near Inambari, Vargas 16460 (GH).

90

FIELDIANA: BOTANY

22. Trichomanes bicorne Hooker, Icon, pi., t. 892.

1854. LECTOTYPE (designated here): Brazil,

Amazonas, Barra do Rio Negro, Spruce 1 178

(lectotype, K!; isolectotype, us!). LECTO-

PARATYPE: Brazil, Amazonas, Sao Gabriel,

Spruce 2334 (K!, us).

Ptilophyllum bicorne (Hooker) Prantl, Unters. Morph.

Gefasskrypt. 1: 48. 1875.

Stem stout, decumbent or erect, abundantly

provided with lustrous, castaneous, pluricellular

trichomes. Leaves monomorphic, crowded or

caespitose, 3-10 cm long, 3-4-pinnatisect, the axes

(and sometimes veins) sparsely provided on the

abaxial side with simple, castaneous, pluricellular

trichomes. Petiole 0.5-4 cm long, conspicuously

alate throughout, the alae broader than the petiole

quite or nearly to base. Lamina ovate, the axes

broadly alate, alae and segments often undulate.

Venation catadromous, veins free, false veins lack-

ing. Sori immersed in the segment tissue. Indusia

short-tubular, borne in the forks of veins, each of

which extend well beyond the indusial mouth, the

mouth not dark-margined, receptacle long-exsert-

ed.

In dense forests, epiphytic, or on wet humus or

decaying logs, sea level to 1 50 m, Loreto.

A lectoparatype (Spruce 2334) is mounted on

the same sheet with the lectotype at Kew. How-

ever, there are other specimens bearing this Spruce

number that represent the type collection of Tri-

chomanes spruceanum Hooker, a closely related

species with dimorphic leaves which occurs in

northern South America. One sheet (BM) is the

isotype of T. spruceanum. Another, presumably

the holotype, is at Kew.

The species is aptly named for its sori, which

are borne at segment apices in the forks of veins.

The veinlets, bordered on the outside with narrow

wings of tissue, each extend well beyond the mouths

of indusia and are often curved, thus appearing

like two "horns."

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-

hana, Rimachi 1256 (GH, MO).

23. Trichomanes diversifrons (Bory) Sadebeck in

Engler & Prantl, Nat. Pflanzenfam. 1(4): 108.

1899.

Trichomanes elegans Rudge, PI. Guian. 24, /. 35. 1805,

nom. illeg. (not L. C. Rich., 1792, or Poiret, 1808).

Hvmenostachys diversifrons Bory, Diet, class, hist. nat.

8: 462. 1825. TYPE: "Guiane," Poiteau (holo-

type, P?; isotype, L; photos, GH & us of L).

Feea diversifrons (Bory) Copel., Phillip. J. Sci. 67: 74.

1938.

Stem stout, erect. Leaves crowded to caespitose,

10-35 cm long, strongly dimorphic, sterile ones

elliptic or lanceolate, deeply pinnatisect, short-pet-

iolate, fertile ones commonly longer, linear, sub-

entire, long-petiolate. Sterile lamina (2-)3-8 cm

broad, deeply divided into subfalcate segments 4-

7 mm broad, the rachis abaxially provided with

dark, simple trichomes and often flagellate and

proliferous at the tip. Venation catadromous, a few

to many veins anastomosing toward the segment

margin, 4-8 of them issuing from the rachis be-

tween adjacent costae, false veins lacking. Fertile

lamina simple, essentially entire, with sori ar-

ranged in a nearly continuous line along each mar-

gin, fully immersed in tissue in the forks of veins.

Indusia subconical, not bilabiate, the mouth nei-

ther flaring nor recurved, strongly indented be-

tween the vein tips, receptacle short- to long-

exserted at maturity.

Terrestrial in wet forests, in wooded ravines and

on ridges and banks, 1 50-1 300 m, Amazonas and

Loreto to Junin, Madre de Dios.

Southern Mexico to Panama; the Guianas to

Colombia, south to Brazil and Bolivia.

This is often confused with Trichomanes trollii,

under which see further discussion.

Amazonas: Prov. Bagua, Quebrada Tambillo, valley

of Rio Maranon above Cascadas de Mayasi, Wurdack

2057 (GH, us). San Martin: Prov. Mariscal Caceres, Dist.

Tocache Nuevo, Santa Rosa de Mishollo, J. Schunke V.

6815 (F, MO, uc). Loreto: Prov. Maynas, Brillo Nuevo

and vicinity, Plowman et al. 6829 (GH). Huanuco: Tingo

Maria, Tryon & Tryon 5277 (F, GH, us). Pasco: Prov.

Oxapampa, Cordillera San Matias, Ledn 320 (F, USM).

Junin: Cahuapanas, on Rio Pichis, Killip & Smith 26762

(F, GH, us). Madre de Dios: Prov. Manu, Cerro de Pan-

tiacolla NNW of Shintuya, Foster et al. 10948 (F).

24. Trichomanes trollii Bergdolt, Flora 127: 256,

264, t. 3. 1933. TYPE: Bolivia, Dept. La Paz,

San Carlos (Mapiri), Troll 2531 (holotype, B;

frag., us!).

Feea trollii (Bergdolt) Vareschi, Flora Venezuela 1 :

247. 1969.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

91

Stem decumbent or erect. Leaves crowded to

caespitose, 10-25 cm long, strongly dimorphic,

sterile ones elliptic, pectinate, short-petiolate, fer-

tile ones commonly longer, linear, subentire, long-

petiolate. Sterile lamina 1.5-4.5 cm broad, cut

nearly to the rachis into subfalcate segments 1.5-

3(— 4) mm broad, the rachis abaxially provided

with simple, dark brown trichomes and often fla-

gellate and proliferous at the tip. Venation catad-

romous, a few to many veins anastomosing toward

the segment margin, 2(-3) of them issuing from

the rachis between adjacent costae, false veins

lacking. Fertile lamina simple, essentially entire,

with sori arranged in a nearly continuous line along

each margin, fully immersed in tissue in the forks

of veins. Indusia subconical, not bilabiate, the

mouth very slightly flaring and recurved, but not

indented between the vein tips, receptacle strongly

exserted at maturity.

Terrestrial in dense, wet forests, often in wet

ravines or along stream banks, 350-850 m, Ama-

zonas to Pasco.

Surinam to Colombia; Peru; Bolivia.

This is closely related to the more robust Tri-

chomanes diversifrons. Besides the characters in

the key the latter can usually be recognized by its

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

lamina of T. trollii is rarely more than 4 cm broad

and are cut nearly to the rachis into segments which

are commonly 3 cm broad or less. Additionally,

the fertile lamina of T. diversifrons has dentate

margins, due to the apices of the indusia which

are deeply indented between the vein forks. In-

dusial mouths of T. trollii are typically flush with

the vein tips, so that the leaf margins are essentially

entire throughout.

Amazonas: Prov. Bagua. along roadside from Chiriaco

to Puente Venezuela, Barbour 4461 (MO in part). San

Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo,

Palo Blanco. Plowman & Schunke 7431 (F). Huanuco:

Ascent of Cerros del Sira. Wolfe 12263 A (GH, MO, us).

Pasco: Prov. Oxapampa. W side of Cordillera de San

Matias, D. Smith 2001 (MO).

25. Trichomanes botryoides Kaulf, Enum. fil. 263.

1824. TYPE: French Guiana, Poiteau (holo-

type?, P).

Stem erect. Leaves crowded to caespitose, 4-12

cm long, strongly dimorphic, sterile ones 1 -pin-

nate, elliptic-lanceolate, short-petiolate, fertile ones

commonly longer and long-petiolate, sori subdis-

tichous on the nonfoliaceous axis. Sterile lamina

1.2-3 cm broad, the pinnae oblong-lanceolate,

subentire to serrate, 3—4 mm broad, the petiole

and rachis with scattered, brown, pluricellular tri-

chomes, the rachis often flagellate and proliferous

at the tip. Venation catadromous, the veins free,

commonly 1 -forked, false veins lacking. Fertile

leaves bearing numerous, stalked or subsessile, as-

cending sori on each side of the primary axis. In-

dusia urceolate, scarcely or not bilabiate, the mouth

not flaring, receptacle strongly exserted at matu-

rity.

Thus far known in Peru from a single collection,

roadside along a sandy, rocky stream bank, 350-

700 m, Amazonas.

Elsewhere terrestrial or epipetric, commonly near

banks of streams or rivers, 250-800 m; Panama;

the Guianas and Colombia.

The small size, strongly dimorphic leaves, and

nonfoliaceous fertile leaves easily distinguish this

from others in the genus.

Amazonas: Prov. Bagua, ca. 40-43 km NE of Chiriaco,

Barbour 4524 (MO, USM).

26. Trichomanes pinnatum Hedwig, Fil. gen. sp.,

/. 4,f. 1. 1799. LECTOTYPE (designated by

Proctor, Flora Lesser Antilles. 89. 1977):

Hedwig, /. 4,f. 1, based on specimen allegedly

from Jamaica. Figure 12c.

Trichomanes pennatum Kaulf., Enum. fil. 264. 1824.

TYPE: French Guiana, without collector (holo-

type, LZ destroyed).

Neurophyllum pinnatum (Hedwig) Presl, Hymeno-

phyllaceae 19. 1843.

Stem short-creeping to decumbent, provided

with dark brown or blackish pluricellular tri-

chomes. Leaves essentially monomorphic (but fer-

tile ones often somewhat larger, with longer pet-

ioles), approximate or subcaespitose, to 70 cm long,

1 -pinnate, the axes sparsely to amply provided on

the abaxial side with castaneous trichomes. Petiole

about as long as the lamina, nonalate (sometimes

marginate toward the lamina). Lamina ovate or

subdeltoid, terminating in a conform apical seg-

ment, or the rachis prolonged, flagellate and pro-

liferous at the tip, the tissue thin and translucent,

glabrous. Venation catadromous, the veins free,

92

FIELDIANA: BOTANY

except connected at the tips by a continuous mar-

ginal vein, false veins copious and perpendicular

to true veins. Sori arranged in a nearly uninter-

rupted line along pinna margins at the tips of veins,

not immersed in the tissue, often stalked. Indusia

narrowly tubular, not or scarcely bilabiate nor dark

margined, mouth not flaring, receptacle long-

exserted.

Terrestrial, in deep forests or wooded ravines,

in humus or clay banks of streams, 150-1000 m,

Amazonas and Loreto to Junin and Madre de Dios.

West Indies and Trinidad; Mexico to Panama

and south to Brazil and Bolivia.

Further differences between this and Trichom-

anes hostmannianum are reviewed below in dis-

cussion of that species.

Amazonas: Prov. Bagua, valley of Rio Maranon near

Cascadas de Mayasi, Wurdack 1939 (GH, us). San Mar-

tin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Gran-

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

SW of Iquitos on road to Puerto Almendra, Croat 18475

(F, MO, uc). Huanuco: Prov. Pachitea, Dist. Honoria,

Bosque Nacional de Iparia, J. Schunke V. 1333 (F, GH,

us). Pasco: Prov. Oxapampa, Palcazu Valley, Cabeza de

Mono, D. Smith 3741 (F, MO). Junin: E of Quimiri Bridge,

near La Merced, Killip & Smith 23924 (F, us). Ucayali:

Along trail to Arboretum of Bosque von Humboldt Ex-

perimental Station, km 86 of Pucallpa-Tingo Maria road,

D. Smith 1226 (F, MO). Madre de Dios: Prov. Tambo-

pata, Vargas 18625, 18711 (GH).

27. Trichomanes hostmannianum (Klotzsch)

Kunze, Bot. Zeit. (Berlin) 5: 352. 1847.

Neurophyllum hostmannianum Klotzsch, Linnaea 1 8:

532. 1844. TYPE: Surinam, Hostmann 75 (ho-

lotype, B; isotypes, K, NY, u, us!; photos, GH & us

ofK).

Odontomanes hostmannianum (Klotzsch) Presl, Epi-

mel. hot. 21. 1849.

Ptilophyllum hostmannianum (Klotzsch) Prantl, Un-

ters. Morph. Gefasskrypt. 1: 49. 1875.

Stem decumbent to erect, provided with dark

brown pluricellular trichomes. Leaves essentially

monomorphic (but fertile ones often larger, with

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

nate, the axes sparsely provided on the abaxial

side with brown trichomes. Petiole commonly as

long as or longer than the lamina, not alate, or

narrowly so toward the lamina. Lamina ovate or

subdeltoid, terminating in a subconform apical

segment, or the rachis prolonged, flagellate and

proliferous at the tip, the tissue relatively thick,

commonly obscure, glabrous, pinna margins ser-

rate, the serrations obtuse to subacute. Venation

catadromous, the veins free, a lateral marginal vein

faint and discontinuous or (more commonly) lack-

ing, false veins lacking or a few very rare and faint

ones perpendicular to true veins. Sori arranged in

a nearly uninterrupted line along pinna margins

at tips of veins, not immersed in the tissue, fre-

quently stalked. Indusia conical to tubular, neither

bilabiate nor dark-margined, mouth not flaring,

receptacle exserted.

Terrestrial in dark, wet forests and wooded rav-

ines, often in frequently inundated areas along

streams, sea level to 450 m, Amazonas, San Mar-

tin, Loreto.

Surinam to Colombia; Amazonian Brazil and

Peru.

This and the closely related Trichomanes pin-

natum are sometimes confused in herbaria. Be-

sides the differences noted in the key they also

differ in size and in leaf texture. Leaves of T. host-

mannianum do not exceed 30 cm, pinnae are rare-

ly more than 6 cm long and 1 cm broad, and tissue

of mature leaves is commonly dull green and ob-

scure. Leaves of T. pinnatum often are 70 cm long,

with pinnae to 1 5 cm long and 2 cm broad, and

tissue is typically thin and translucent, so that the

false veins are easily visible with little magnifi-

cation.

Contrary to previous reports, false veins do oc-

cur in T. hostmannianum. They are infrequent and

scattered and so faint as to be overlooked, or un-

detectable, in the thick leaf tissue. However, in

immature leaves or in the rare thin-textured ones,

high magnification sometimes reveals a few or sev-

eral of them, perpendicular to the true veins as in

T. pinnatum. As this character is so difficult to

observe, it is rather ineffective in delineating T.

hostmannianum from species in other sections of

the subgenus; yet it serves to further establish its

position in sect. Neurophyllum (Presl) Moore, along

with T. pinnatum. Morton ( 1 968) placed it outside

the section because of the supposed lack of false

veinlets.

Amazonas: Nazareth, Osgood 25 (F, us). San Martin:

Prov. Mariscal Caceres, Dist. Tocache Nuevo, near

Granja San Ysabel, J. Schunke V. 10313 (F, MO). Loreto:

Rio Tacsha Curaray, Croat 20403 (F, MO, uc). Dist. Iqui-

tos, trail through Versailles, Mexia 6500 (GH, MO, uc,

us).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. I.

93

28. Trichomanes humboldtii Lell., Mem. New

York Bot. Gard. 38: 35. 1984, nom. nov. for

Trichomanes heterophyllum Willd. and with

the same type.

Trichomanes heterophyllum Willd., Sp. pi. ed. 4, 5:

503. 1810, nom. illeg., not T. heterophyllum (Sm.)

Poir. 1808. TYPE: Brazil, Amazonas, Javita,

Humboldt (holotype, B, Herb. Willd. 20210; pho-

tos, F, GH).

Homoeotes heterophylla Presl, Gefassbiindel Farm 24.

1847, nom. nov., based on Trichomanes hetero-

phyllum Willd. and with the same type.

Feea humboldtii Bosch, Ned. Kruidk. Arch. 4: 347.

1859, nom. nov., based on Trichomanes hetero-

phyllum Willd. and with the same type.

Feea heterophylla Copel., Philipp. J. Sci. 67: 73. 1938,

nom. nov., based on Trichomanes heterophyllum

Willd. and with the same type.

Stem wiry, long- to short-creeping. Leaves deep-

ly pinnatisect to nearly pinnate, subdistant or re-

mote, to 22 cm long, dimorphic, sterile ones short-

petiolate, to 3 cm broad, fertile ones long-petio-

late. far exceeding the sterile ones and 1 cm broad

or less, the petiole essentially nonalate, sparsely or

amply provided with castaneous, pluricellular tri-

chomes. Sterile lamina oblong or oblong-elliptic,

commonly equaling or longer than the petiole, the

apex pinnatifid. Venation catadromous, veins free

and dichotomously branched, false veins lacking.

Fertile lamina linear, much shorter than the pet-

iole, with a few sori terminal on each pinna and

fully immersed in the tissue. Indusia subconical

or broadly tubular, not bilabiate, the mouth not

(or scarcely) flaring nor dark-margined, the recep-

tacle long-exserted.

Terrestrial, in open to dense forests, commonly

in wet, sandy places, sea level to 1400 m, San

Martin, Loreto.

Venezuela; Colombia; Amazonian Peru and

Brazil.

There has been some confusion with nomen-

clature of this species. Lellinger's (1984) citing of

Trichomanes humboldtii "Bosch" (1858) was in

error, since there was no such name; instead, it

was "Feea humboldtif that Bosch had published.

However, Lellinger's use of 'T. humboldtii"'' is le-

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

name is correct for the species, with Lellinger as

the author.

San Martin: Rioja, Soukup 5055 (us). Rio Negro,

Woytkowski 6211 (GH, MO, us). Loreto: Prov. Maynas,

Rio Nanay, Mishana, Foster & Foster 4103 (GH);

McDanielet al. 22121 (F). Mishuyacu, near Iquitos, Klug

1246 (F, us).

29. Trichomanes crinitum Sw., Prodr. 136. 1788.

TYPE: Jamaica, Swartz (holotype, SET; iso-

types, BM!, s; photos, us of BM & s).

Ragatelus crinitus (Sw.) Presl, Hymenophyllaceae 16.

1843.

Stem erect. Leaves monomorphic, caespitose,

5-25 cm long, l-2(-2.5) cm broad, moderately to

amply pubescent, the trichomes delicate, un-

branched, unicellular (or rarely pluricellular) be-

yond a short, enlarged, basal cell. Petiole nonalate,

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

a pinnatifid apex, lamellae lacking on veins. Ve-

nation catadromous, the veins free, false veins

lacking. Sori terminating the veins on the apical

part of the pinnae. Indusia tubular, deeply im-

mersed in the segment tissue or, if partially im-

mersed, broadly alate on each side, the mouth

bilabiate and somewhat flaring, the lips not dark-

margined, receptacle exserted.

Apparently known thus far from two collections

in Peru, in damp elfin forest, on tree trunks, ca.

1850 m, Huanuco. Elsewhere epiphytic or epi-

petric, 700-2300 m.

West Indies; Costa Rica; Venezuela; Colombia;

Ecuador; Peru.

Trichomes in many species of subg. Achomanes

are pluricellular, with two to several greatly elon-

gate cells, these often springing from a very short,

widened basal cell. In species most closely related

to Trichomanes crinitum, these are often mixed

with a number of unicellular (above the basal cell)

trichomes. However, T. crinitum differs from its

Peruvian relatives in having all, or nearly all, of

its laminar trichomes unicellular above the basal

cell.

Huanuco: SW slope of Rio Llullapichis watershed, on

the ascent of Cerros del Sira, Dudley 13525 (GH), 13545

(GH, us).

30. Trichomanes martiusii Presl, Hymenophyl-

laceae 36. 1843. TYPE: a renaming of T. pi-

losum (sensu Martius in part, Icon. pi. crypt.

104, t. 68 right) presumably based on "Brazil,

94

FIELDIANA: BOTANY

Arara-coara" (Araracuara, on Rio Caqueta,

now Colombia), Martius (holotype, M; iso-

types, BR, L; photo, us of L).

Trichomanes plumula Presl, Hymenophyllaceae 15,

36. 1843. TYPE: based on Martius in part. Icon.

pi. crypt. 104, /. 68 left, specimens probably as

for T. martiusii.

Ptilophyllum martiusii (Presl) Prantl, Unters. Morph.

Gefasskrypt. 1: 48. 1875.

Stem erect to decumbent. Leaves monomor-

phic, crowded to caespitose, 1 0-40 cm long, (2-)3-

8 cm broad, densely (on axes) to amply pubescent,

the trichomes unbranched, unicellular beyond the

short basal cell, or pluricellular on petiole and

rachis. Petiole nonalate. Lamina narrowly elliptic

or lanceolate, gradually reduced to a pinnatifid

apex, cut nearly or quite to the rachis into linear

pinnae, the proximal ones commonly deflexed,

subentire to crenulate or serrate, or rarely with a

few, scattered lobes, with numerous, crestlike la-

mellae borne on the veins abaxially. Venation ca-

tadromous, the veins free, false veins lacking. Sori

terminating the veins on the apical part of seg-

ments. Indusia tubular or narrow-conical, deeply

immersed in the segment tissue, the mouth trun-

cate, not dark-margined, sometimes apparently

bilabiate due to narrow wings of tissue extending

beyond the mouth on two sides, receptacle com-

monly exserted.

On tree trunks in forests, or on clay banks along

roads and streams, sea level to 150 m, thus far

found only in Loreto, but locally common there.

Surinam to Colombia; Peru; Amazonian Brazil.

The shape of the indusium is somewhat vari-

able. The mouth is commonly truncate and not at

all bilabiate, but frequently the wings of tissue along

the sides extend beyond the mouth, giving it a

"horned" appearance, much like Trichomanes bi-

corne. This is a distinctive species in Peru, because

of the crestlike lamellae borne on many of the

veins on the abaxial side, perpendicular to the

plane of the lamina. These processes are similar

to those occurring in several species of Hymeno-

phyllum. On some of the more densely hirsute

specimens of T. martiusii, the lamellae are par-

tially obscured by the indument and thus they

have been mistaken for other species in subg.

Achomanes, particularly T. pilosum Raddi, of Bo-

livia and southern South America.

Loreto: Prov. Maynas, Rio Nanay, behind Mishana,

Gentry & Revilla 20704 (MO, uc, USM). Mishayucu, near

Iquitos, Klug 196, 51 1 (F, us). Prov. Maynas, 10 km S

of Iquitos, Tryon & Tryon 5179 (GH, us). Timbuchi on

Rio Nanay, LI. Williams 957 (F, us).

31. Trichomanes lucens Sw., Prodr. 136. 1788.

TYPE: Jamaica, Swartz (holotype, SET; iso-

type, BM; frag., B; photos, us of SBT & BM), not

T. lucens Hooker & Grev., 1827.

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

4 IB. 1846. TYPE: Peru, Huanuco, Pillao, Ruiz

& Pavon (holotype, K; isotype, B; photos, GH &

us of K).

Ptilophyllum lambertianum (Hooker) Prantl, Unters.

Morph. Gefasskrypt. 1: 48. 1875.

Stem erect to decumbent. Leaves monomor-

phic, crowded to caespitose, 10-60 cm long,

(2.5-)3-12 cm broad, amply to densely pubescent

(indument often obscuring the laminar surface),

the trichomes unbranched, unicellular and pluri-

cellular beyond the scarcely evident basal cell. Pet-

iole nonalate. Lamina linear to lanceolate, pin-

nate-pinnatifid (occasionally 2-pinnate-pinnatifid

toward base), gradually reduced to a pinnatifid

apex, pinnae narrow-deltoid, mostly ascending,

lacking lamellae. Venation catadromous, the veins

free, false veins lacking. Sori commonly terminal

on ultimate segments near apex of pinnae. Indusia

tubular, deeply immersed in the segment tissue,

the mouth scarcely or slightly flaring, not dark-

margined, sometimes apparently bilabiate due to

the narrow wings of tissue extending beyond the

mouth on two sides, receptacle commonly exsert-

ed.