BIOLOGV

Botany

PTERIDOPHYTA OF

PART V

18. Aspleniaceae-21. Polypodiacea<

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FIELDIANA

Botany

NEW SERIES, NO. 32

PTERIDOPHYTA OF PERU

PARTY

18. Aspleniaceae-21. Polypodiaceae

Rolla M. Tryon

Department of Biology

University of South Florida

Tampa, Florida 33620-5150

With the collaboration of:

Blanca Leon

Universidad Nacional Mayor de San Marcos

Lima, Peru

Robert G. Stolze

Associate Curator

Department of Botany

Field Museum of Natural History

Roosevelt Road at Lake Shore Drive

Chicago, Illinois 60605-2496

Accepted January 6, 1993

Published June 30, 1993

Publication 1447

BIOLOGY LIBRARY

101 BURRILL HALL

SEP 2 7 I993

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 1993 Field Museum of Natural History

Library of Congress Catalog Card Number: 93-71887

ISSN 00 15-0746

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

List of Illustrations

INTRODUCTION 1

ACKNOWLEDGMENTS 1

1 8. ASPLENIACEAE 2

Asplenium 2

1 9. DAVALLIACEAE 49

Nephrolepis 49

20. BLECHNACEAE 54

Blechnum 56

Salpichlaena 68

2 1 . POLYPODIACEAE 70

Grammitis 72

Pecluma 116

Polypodium 125

Pleopeltis 140

Dicranoglossum 145

Microgramma 148

Campyloneurum 158

Niphidium 173

Solanopteris 1 79

Platycerium 181

MAP OF PERU 1 84

INDEX TO NAMES . .185

1. Asplenium: A. pseudoangustum; A.

hallii; A. theciferum 3

2. Nephrolepis: N. pectinata; N. cordifo-

lia; N. multiflora; N. rivularis 50

3. Blechnum: B. asplenioides; B.fraxi-

neum; B. binervatum ssp. fragile 55

4. Salpichlaena: S. volubilis 69

5. Grammitis: G. myosuroides; G. limba-

ta\ G. david-smithii; G. myriophylla ... 73

6. Pecluma: P. hygrometrica; P. filicula;

P. eurybasis var. pilosa; P. curvans 117

7. Polypodium: P. decumanum; P. au-

reum; P. loriceum; P. triseriale 126

8. Pleopeltis: P. macrocarpa var. macro-

carpa; P. macrocarpa var. laciniata; P.

percussa 141

9. Dicranoglossum: D. subnudum; D. des-

vauxii; D. polypodioides 147

10. Microgramma: M. thurnii; M. reptans\

M. squamulosa 1 50

11. Campyloneurum: C. fasciale; C. phylli-

tidis; C. angustipaleatum\ C. ophiocau-

lon; C. amphostenon 159

12. Niphidium: N. crass ifolium; N. mac-

bridei 175

13. Solanopteris: S. bifrons; S. bismarckii . . 178

14. Platycerium: P. andinum 182

in

Back cover: Nephrolepis pectinata

PTERIDOPHYTA OF PERU

PartV

18. Aspleniaceae-21. Polypodiaceae

Introduction

This fifth part of the "Pteridophyta of Peru"

contains the fern families Aspleniaceae, Blech-

naceae, Davalliaceae, and Polypodiaceae. Al-

though it was stated in the introduction to Part I

that this pteridophyte flora would be contained in

five parts, it has been deemed more practical to

add a sixth part (now in progress), which will in-

clude the "water ferns," "fern allies," a brief bio-

geography of Peru, and a comprehensive index to

the entire work. The general style, typography, form

of citations, and so forth follow the previously

published parts. These matters are adequately dealt

with in Part I (Fieldiana: Botany, n.s., No. 20, 1989),

and it is not necessary to repeat them here.

Campyloneurum of the Polypodiaceae has been

contributed by Blanca Leon, and the other genera

are a joint effort of Rolla M. Tryon and Robert

G. Stolze, each critically reviewing the treatments

prepared by the other.

Type collections from Peru are mentioned in

the nomenclature but are not repeated in the spec-

imen citations. They are, however, included in the

Peru range and ecology. The nomenclature of the

genera and species is not intended to be complete.

It includes all names based on Peru material and

other names that are considered useful to mention.

Abbreviations of periodicals generally follow the

system of Botanico-Periodicum-Huntianum

(1968), while those of books and authors generally

follow the system of Taxonomic Literature (TL-

2, 1 976 et seq.). A notable exception is Karel Dom-

in's "The pteridophyta of the island of Dominica"

(1929), which is cited throughout as "Pterid.

Dominica," in the interest of brevity. According

to TL-2, the full citation is Rozpr. Krai. Ceske

Spolecn. Nauk, Tr. Mat. Prir. N.R. 2. The abbre-

viations for herbaria follow Index Herbariorum.

Acknowledgments

Blanca Leon is nearing completion of a mono-

graphic study of Campyloneurum (Polypodi-

aceae), a genus in which there are very few diag-

nostic characters to separate the species. She has

seen nearly all of the type collections during sev-

eral years of research and so has gained a thorough

knowledge of a taxonomically difficult group of

ferns. The authors are extremely fortunate to have

her contribution to this flora. We would like to

extend special thanks to Dr. Abundio Sagastegui

(HUT) and Blanca Leon (USM) for their invaluable

assistance in preparing loans and arranging for their

packing and shipment from those two important

Peruvian herbaria. The illustrations were contrib-

uted by Field Museum scientific illustrator Zorica

Dabich, who created the original drawings and

adapted the rest from those previously used in the

Fieldiana: Botany publication, "The Ferns and

Fern Allies of Guatemala." We are grateful to her,

for her art has provided a valuable complement

to the descriptions. We are indebted to Dr. Alan

R. Smith and Dr. Robbin C. Moran for their crit-

ical suggestions on the treatment of Grammitis

(Polypodiaceae), and to Dr. David B. Lellinger for

stimulating discussions and correspondence per-

taining to a wide variety of taxonomic problems.

To Dr. Brigitte Zimmer, of the Botanical Museum,

Berlin-Dahlem, we extend special thanks for valu-

FIELDIANA: BOTANY, N.S., NO. 32, JUNE 30, 1993, PP. 1-190

able comments on typification and nomenclature,

particularly in regard to Pecluma and Microgram-

ma. We also appreciate the valuable comments

presented by reviewers of the manuscript.

We are also grateful to the officers of the follow-

ing institutions for granting loans of their material

or allowing us to examine specimens in their her-

baria: Botanischer Garten und Botanisches Mu-

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

(Natural History), London (BM); Jardin Botanique

National de Belgique, Meise (BR); Field Museum

of Natural History, Chicago (F); Harvard Univer-

sity, Cambridge, Mass. — most Gray Herbarium

(GH), some Arnold Arboretum (A); Herbarium

Truxillense, Universidad Nacional de Trujillo,

Trujillo, Peru (HUT); Royal Botanic Gardens, Kew,

England (K); Rijksherbarium, Leiden, The Neth-

erlands (L); Missouri Botanical Garden, St. Louis

(MO); New York Botanical Garden, New York (NY);

Museum National d'Histoire Naturelle, Paris (P);

National Museum, Prague, Czechoslovakia (PR);

Botanical Institute, Charles University, Prague,

Czechoslovakia (PRC); Herbario del Institute de

Ciencias Naturales, Universidad Central, Quito,

Ecuador (Q); Rijksmuseum, Stockholm, Sweden

(s); University of California, Berkeley (uc); United

States National Herbarium, Smithsonian Institu-

tion, Washington, D.C. (us); Herbario San Mar-

cos, 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.

Family 18. ASPLENIACEAE

Aspleniaceae Newman, Hist. Brit, ferns, 6: 1840.

TYPE: Asplenium L.

Terrestrial, epiphytic, or epipetric. Stem com-

monly erect, sometimes slender and long-creeping,

bearing scales (and rarely also trichomes), the scales

obviously clathrate, or occasionally only incon-

spicuously so. Leaves circinate in vernation, pin-

nate to decompound, or in several species entire,

essentially monomorphic, of small to moderate

size, or rarely to 1 m long, petiole continuous with

the stem. Lamina essentially glabrous, or several

species with minute trichomes or reduced scales,

especially along the rachis, the rachis sometimes

proliferous at the pinna axils or apex. Veins free,

or in a few species casually to freely anastomosing.

Sori linear to elliptic, borne along a vein, covered

by a short to very long and narrow indusium, or

rarely with a nearly marginal pouch formed by the

indusium and opposed leaf tissue, paraphyses

lacking. Sporangia usually long-stalked, the cells

of the stalk uniseriate below its apex, the annulus

interrupted by the stalk. Spores monolete, bilat-

eral, lacking chlorophyll, usually with sharp folds

in the perispore.

The Aspleniaceae, as recognized here, are a fam-

ily of about 700 species, included in seven genera,

of which five are found in the New World: As-

plenium, Camptosorus, Holodictyum, Pleurosorus,

and Schaffneria. Only Asplenium occurs in Peru.

The family is typically characterized by small to

medium-sized leaves with clathrate stem scales

and nonarticulate petioles, veins commonly free

or, when anastomosing, the areoles without free

included veinlets, and elongate sori with a narrow

indusium borne along a vein.

References

MORTON, C. V., AND D. B. LELLINGER. 1986. The

Polypodiaceae subfamily Asplenioideae in Ven-

ezuela. Mem. New York Bot. Gard., 15: 1-49.

STOLZE, R. G. 1986. Polypodiaceae— Asplenioi-

deae. In Harling, G., and B. Sparre, eds., Flora

of Ecuador 23: 1-83. Stockholm.

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

iaceae, pp. 627-654. In Ferns and Allied Plants.

Springer- Verlag, New York.

I. Asplenium

Asplenium L., Sp. pi. 1079. 1753; Gen. pi., ed. 5,

485. 1754. TYPE: Asplenium marinum L.

Figure 1.

FIG. 1 . Asplenium pseudoangustum: a, habit. Asplenium hallii: b, habit; c, stem scale; d, base of pinna. Asplenium

theciferum: e, apex of pinna, (a from Schunke V. 6951, F; b, c, d from Acosta Solis 13761, Ecuador, F; e from Holm-

Nielsen et al. 4110, Ecudaor, F.)

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

Phyllitis Hill, Brit. herb. 525. 1757. TYPE: Phyllitis

scolopendrium Newman = Asplenium scolopen-

drium L.

Loxoscaphe Moore, Hooker's J. Bot. Kew Gard. Misc.

5: 227. 1853. TYPE: Loxoscaphe concinna

(Schrader) Moore (Davallia concinna Schrader) =

Asplenium concinnum (Schrader) Kuhn = As-

plenium theciferum (HBK.) Mett.

Leaves pinnate to 4-pinnate, in a few species

entire, 3-60(-100) cm long. Lamina terminating

in a conform apical segment, or more commonly

tapering to a pinnatifid apex, or occasionally with

a flagellate, proliferous tip. Pinnae, when present,

several to many, subequilateral to (more com-

monly) inequilateral at base. Veins simple or

forked, free, or reticulate in A. purdieanum. Sori

elliptic to (more commonly) linear, borne along

the acroscopic side of veins or, in several species,

a few of them also borne along the basiscopic side

(thus back-to-back, i.e., diplazioid). Indusia nar-

row, attached along the vein and opening toward

the midrib, commonly persistent, rarely obsolete,

or in A. theciferum cuplike or pouchlike, borne

singly near segment tips, with sporangia deeply

immersed. Sporangia on slender stalks composed

of a single row of cells, annular cells commonly

20-25.

There are more than 600 species in this nearly

cosmopolitan genus, a majority of these occurring

in wet forests of tropical regions. Unfortunately,

although the numerous species are rather easily

identified to genus, they are not readily (or at least

obviously) classified into infrageneric groups. As

in Diplazium, many can be separated on the basis

of pinna dissection, and yet very closely related

species (and even a species itself) can be 1-3-pin-

nate. One exception to this is the group of species

often placed in the genus Loxoscaphe. Most, in-

cluding Asplenium theciferum, are quite distinc-

tive in their indusia, which are generally pouch-

or cup-shaped and occur near the segment margin.

By means of this character, the latter species can-

not be confused with any other in Peru. But there

are a few Old World species in the group in which

the sori intergrade to a more typical Asplenium

sorus. For this reason Loxoscaphe is included with

Asplenium for purposes of this treatment.

A few species with back-to-back (diplazioid) sori

are frequently determined as Diplazium. With these

species, comparison of stem scales, and of spo-

rangia stalks and annular cells, is necessary for

positive identification to genus. (Also see Smith,

1976.)

References

MUTUI, K., N. MURIKAMI, AND K. IWATSUKI.

1989. Chromosomes and systematics of As-

plenium sect. Hymenasplenium (Aspleniaceae).

Amer. J. Bot., 76: 1689-1697.

SMITH, A. R. 1976. Diplazium delitescens and

the Neotropical species of Asplenium sect. Hy-

menasplenium. Amer. Fern J., 66: 1 16-120.

Key to Species of Asplenium

a. Lamina simple, margins entire or serrate; petiole obsolete or VJ0 to V20 the length of the lamina . b

b. Larger adult leaves 4.2-15 cm broad, 20-70(-100) cm long; veins usually spreading at 50-75°

from the midrib 1 . A. serratum

b. Larger mature leaves 0.5-3(-4) cm broad, 8-30(-40) cm long; veins spreading at 30-45° from the

midrib c

c. Midrib with scattered, minute, clathrate scales, at least abaxially; stem scales blackish, linear,

attenuate, obscurely clathrate; veins commonly evident (if not prominent) ... 2. A. angustum

c. Midrib lacking scales; stem scales brown, lanceolate to ovate, acute or acuminate; veins obscure

3. A. pseudoangustum

a. Lamina lobed or pinnate to decompound or (in a few species) simple and unlobed, and then petiole

'/4-'/2 the length of the lamina d

d. Indusium cup- or pocket-shaped, borne along the acroscopic side of ultimate segments near their

tips 62. A. theciferum

d. Indusium linear to narrow-elliptic, attached along the vein and opening toward the midrib . . e

e. Lamina simple, the margins entire to serrate or crenulate f

f. Lamina base extremely long-decurrent; midrib abaxially with scattered, minute, blackish

scales; lamina chartaceous, lacking glandular trichomes 4. A. stuebelianum

f. Lamina base short-decurrent; midrib lacking scales; lamina thin- to firm-herbaceous, with

scattered, minute (0.1-0.2 mm), appressed, often gland-tipped trichomes .. 5. A. pearcei

FIELDIANA: BOTANY

e. Lamina lobed to 1 -pinnate or decompound g

g. Lamina shallowly lobed to pinnatifid (or some juvenile leaves with a pair of lobed basal

pinnae); leaves diminutive, 2-6 cm long, 0.2-0.8 mm broad 6. A. escaleroense

g. Lamina regularly 1 -pinnate or decompound (in a few species juvenile leaves sometimes

simple and entire); leaves small to large h

h. Basal pair of pinnae much more strongly produced basiscopically than acroscopically;

lamina with few to many whitish, septate trichomes, especially on rachis, veins, and

margins 7. A. pumilum

h. Basal pair of pinnae subequilateral at base or more strongly produced acroscopically;

lamina lacking whitish, septate trichomes i

i. Petiole and base of rachis highly lustrous and dark brown, atropurpureous or blackish

j

j. Pinnae entire to sharply and deeply incised, 0.6-1.5 cm long k

k. Pinnae deeply incised 8. A. formosum

k. Pinnae entire to crenate, serrate, or shallowly lobed 1

1. Adaxial raised ribs of rachis conspicuously hirsute, the rigid, spreading, cas-

taneous trichomes to 0.3 mm long; pinnae subequilateral ... 9. A. extensum

1. Adaxial ribs or wings of rachis entire, denticulate, or pustulate; pinnae inequi-

lateral at base, cuneate basiscopically, truncate and often subauriculate ac-

roscopically m

m. Vein tips not dilated, inconspicuous, or obscured in leaf tissue; adaxial

ribs of rachis each with a pronounced, foliaceous green wing throughout

10. A. resiliens

m. Vein tips (most of them) somewhat to strongly dilated and conspicuous

on adaxial side of pinnae; adaxial ribs commonly lacking foliaceous green

wings 11. A. monanthes

j. Pinnae 1-3 -pinnate, or if subentire to incised, then larger ones (1.8-)2-12 cm long

n

n. Lamina gradually reduced at base; petiole 1-6 cm long o

o. Lamina 2-3-pinnate 12. A. rutaceum

o. Lamina 1 -pinnate-pinnatisect 13. A. hallii

n. Lamina truncate or nearly so p

p. Rachis not prolonged or proliferous, the lamina apex pinnatifid; stem short-

creeping (often inconspicuously so in A. laetum); pinnae subentire to shal-

lowly pinnatifid q

q. Pinnae 15-24 pairs, 2.5-4.5 cm long, most of them strongly inequilateral

for nearly half their length 31. A. laetum

q. Pinnae 7-1 1 pairs, larger ones 6-1 2 cm long, essentially equilateral except

at the base 32. A. melanopus

p. Rachis prolonged, often proliferous; stem erect or decumbent; pinnae sub-

entire to 3-pinnate r

r. Stem scales 0.6-1 mm broad, ovate or broadly lanceolate, acute, dark or

grayish brown and often with lighter brown margins . . 14. A. macrurum

r. Stem scales 0.1-0.3 mm broad, linear or narrowly lanceolate, attenuate,

gray-brown to blackish, concolorous 15. A. radicans

i. Petiole and base of rachis not or slightly lustrous, green, reddish or grayish brown

(rarely darker, but then not highly lustrous) s

s. Pinnae mostly trifoliolate or bifoliolate, occasionally with l(-2) additional pairs of

pinnules t

t. Leaves 1.5-5 cm long, remote, borne singly or in pairs from long-creeping,

stoloniform roots; pinnae 4-8 pairs; stem scales few, 0.5 mm long

1 6. A. repens

t. Leaves (4-)6-30 cm long, fasciculate from an erect stem, pinnae numerous; stem

scales many, 1-2 mm long 17. A. triphyllum

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

s. Pinnae simple or regularly pinnate to decompound, not trifoliolate or bifoliolate

u

u. Lamina 2-pinnate or more, at least at pinna bases; basal segments (pinnules) of

proximal pinnae obviously stalked v

v. Lamina conspicuously reduced at base, proximal pinnae often rudimentary

w

w. Fertile ultimate segments mostly less than 2 mm long, conspicuously

broader than adjacent sterile ones; stem scales bicolorous, blackish with

narrow but conspicuously brown margins 18. A. divaricatum

w. Fertile and sterile ultimate segments 2-4 mm long, subequal; stem scales

concolorous, or occasionally faintly brown-margined x

x. Lamina linear, 2-pinnate; pinnules 3-6 pairs on a pinna; plants epipetric

(rarely terrestrial), growing above 2900 m 19. A. haenkeanum

x. Lamina elliptic or elliptic-ovate, 2-pinnate-pinnatisect or 3-pinnate;

pinnules of larger pinnae 7-14 pairs; plants terrestrial (rarely epipetric

or epiphytic), at elevations from 250 to 2700 m y

y. Pinnae commonly patent, margins parallel for much of their length;

ultimate segments mostly narrow-elliptic, acute .20. A. cristatum

y. Pinnae commonly ascending, broadest near the base, from there

gradually to abruptly tapering to apex; ultimate segments mostly

obovate, obtuse to subacute 21. A. myriophyllum

v. Lamina not or scarcely reduced at base z

z. Leaves 1.5-8(-10) cm long, borne directly on stoloniform (commonly

long-creeping) roots; scales of petiole base 0.4-0.6 mm long a2

a2. Ultimate segments flabelliform, subdigitately incised at apex, veins

commonly 1 -forked in each segment; petiole to 0.5 cm long

16. A. repens

a2. Ultimate segments linear or narrowly oblanceolate, apex acute, veins

simple in each segment; petiole 1.5—4 cm long . . 22. A. delicatulum

z. Leaves (10-) 12- 100 cm long, borne on a stout, short-creeping to erect,

stem; scales of stem and petiole base 1.5-20 mm long b2

b2. Rachis obviously alate throughout; most pinnae oblong, margins par-

allel for much of their length, spreading at right angles, sessile, their

basal pinnules strongly overlapping the rachis . . . 20. A. cristatum

b2. Rachis nonalate, or narrowly so distally; pinnae ovate or deltate,

tapering from base to apex, ascending, short-stalked, basal pinnules

not or infrequently overlapping the rachis c2

c2. Lamina 1 5-20 cm broad; pinnules (2-)3-9 cm long; stem scales

10-20 mm long, 3-6 mm broad; indusia thin, commonly scarious

23. A. squamosum

c2. Lamina 2-1 2(-l 5) cm broad; pinnules 0.5-2 cm long; stem scales

2—4 mm long, 0.3-0.8 mm broad; indusia firm or thick and fleshy,

whitish to gray, or light brown d2

d2. Stem scales linear, attenuate, dark brown; lustrous; ultimate

segments obovate to subflabellate, broadly and obtusely den-

tate near apex; veins subdichotomously branched

56. A. cuneatum

d2. Stem scales lanceolate to ovate, dull orange to gray-brown;

ultimate segments linear-lanceolate or elliptic, acute or sub-

acute; veins 1 -forked to pinnately branched

24. A. cuspidatum

u. Lamina 1 -pinnate to 1 -pinnate-pinnatisect (in a few species nearly 2-pinnate at

bases of proximal pinnae); basal secondary segments (if any) adnate, not freely

stalked . . e2

FIELDIANA: BOTANY

e2. Lamina terminating in a distinct, conform or subconform apical segment

(this occasionally with a basal lobe or auricle) f2

f2. Veins anastomosing, especially toward pinna margin; petiole copiously

scaly 61. A. purdieanum

f2. Veins free; petiole naked or sparsely scaly (or sometimes in A. serra

conspicuously scaly) g2

g2. Leaves approximate to remote on a short- to long-creeping

stem h2

h2. Stem abundantly scaly; lamina firm-herbaceous to (more com-

monly) chartaceous, the petiole and rachis scaly (scales at times

caducous); pinnae subequilateral at base 57. A. serra

h2. Stem essentially lacking scales; lamina membranaceous or thin-

herbaceous, the petiole and rachis naked; pinnae conspicuously

inequilateral at base 25. A. repandulum

g2. Leaves crowded to caespitose on an erect to decumbent (rarely very

short-creeping) stem i2

i2. Pinnae conspicuously inequilateral at base, there acroscopically

truncate to cordate or auriculate, often overlapping the rachis

J2

j2. Larger pinnae 2.5-4 cm broad; pinnae bases (at least prox-

imal ones) mostly truncate to broadly rounded basiscopi-

cally 26. A. vomeriforme

j2. Larger pinnae 1-2 cm broad; pinna bases mostly narrowly

cuneate basiscopically 53. A. sal id folium

i2. Pinnae subequilateral at base, there broadly or narrowly cuneate

to attenuate k2

k2. Stem scales filiform, 0. 1-0.3(-0.4) mm broad

27. A. juglandifolium

k2. Stem scales lanceolate to ovate, 0.5-2 mm broad ... 12

12. Lamina with 1 (very rarely 2) pair of pinnae, these

subopposite and narrowly adnate; stem scales 3-4 mm

long 5. A. pearcei

12. Lamina with (l-)2-8 pairs of pinnae, these alternate,

sessile or short-stalked, not or rarely adnate; stem scales

mostly 5-1 5 mm long m2

m2. Stem scales 1-2.5 mm long, 0.5 mm broad, black-

ish, often slightly vaulted; apical lamina segment

1-1.7 mm broad or occasionally obso-

lete 28. A. tuerckheimii

m2. Stem scales 5-15 mm long, mostly 1-2 mm broad,

orange to brown, flat; apical segment 2.2-5.0 cm

broad n2

n2. Pinna margins entire; lamina glabrous

abaxially; plants epiphytic

29. A. tricholepis

n2. Pinna margins crenate-serrate; lamina mod-

erately to amply provided abaxially with

minute, appressed, dark brown trichomes;

plants terrestrial 30. A. davisii

e2. Lamina tapering to a pinnatifid, caudate, or flagelliform apex or, if with a

distinct terminal segment, this obviously differing in shape from lateral

segments o2

o2. Petiole, rachis and tissue sparsely to copiously scaly, the scales dark

and filiform, with prolonged, unicostate apices p2

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

p2. Pinnae serrate to deeply and irregularly erose-lacerate, not cune-

ately lobed 58. A. dimidiatum

p2. Pinnae deeply lobed, the 3-6 segments cuneiform, their apices often

lacerate 59. A. praemorsum

o2. Petiole, rachis and tissue with scales lacking or scattered and incon-

spicuous q2

q2. Stem short-creeping (sometimes not obviously so in A. laetum);

leaves closely spaced or crowded along the stem r2

r2. Pinnae 14-24 pairs, larger ones 2.5-5 cm long; sori not or

rarely diplazioid s2

s2. Stem scales orange; veins and sori subparallel, or diverging

at 5-10°; pinnae deeply pinnatisect 60. A. lividum

s2. Stem scales dark brown or blackish; veins and sori di-

verging at 30-40°; pinnae bicrenate or biserrate

31. A. laetum

r2. Pinnae 6-10(-12) pairs, larger ones 6-14 cm long; sori often

diplazioid t2

t2. Pinnae (at least proximal ones) lobed to pinnatifid on both

margins, the tip of each lobe with a few rounded teeth .

32. A. melanopus

t2. Pinnae serrate to deeply crenate acroscopically, serrate to

subentire basiscopically 33. A. delitescens

q2. Stem erect, not creeping; leaves fasciculate to densely caespi-

tose u2

u2. Pinnae incised nearly to the costa into 7-16 pairs of entire to

crenate segments 34. A. achilleifolium

u2. Pinnae subentire to crenate, or with a nearly free basal acro-

scopic auricle, but not regularly pinnatisect v2

v2. Pinnae lacking a definite midrib, the veins few and com-

monly subflabellately dichotomous w2

w2. Veins conspicuous, blackish, simple or 2-3-forked;

pinnae 2-3 mm long, with short, blackish, stalks . .

35. A. projectum

w2. Veins obscure, or if distinct only slightly darker than

the laminar tissue, commonly subflabellate, 2-3 times

dichotomous; pinnae 4-14 mm long, not black-

stalked x2

x2. Larger pinnae 1 0- 1 4 mm long; plants terrestrial,

growing below 1000 m 36. A. lorentzii

x2. Larger pinnae 4-8(-10) mm long; plants epi-

petric (rarely terrestrial), growing above 2700 m

y 2

y2. Petiole 0.2-0.4 mm thick, this and rachis

flexuous and greenish or stramineous; pin-

nae broadly and deeply lobed and cuspidate

37. A. gilliesii

y2. Petiole 0.5-1 mm thick, this and rachis rigid

and commonly reddish or grayish brown;

pinna margins entire to deeply dentate,

rarely lobed 38. A. peruvianum

v2. Pinnae, at least in the proximal half, with a definite, central

(or basiscopic in subdimidiate pinnae) axis, this either a

thick, raised, costa, or straight midvein bearing several to

many pinnately arranged veins z2

FIELDIANA: BOTANY

z2. Stem scales lacking; pinna midrib with 2(-3) acro-

scopic, and usually 1 basiscopic, veins

39. A. quitense

2.2. Stem scales ample and conspicuous; pinnae, larger

ones, with 3-many pairs of veins (sometimes less in

A. pulchellum) a3

a3. Pinnae subdimidiate, i.e., conspicuously inequi-

lateral for half their length or more; veins 1-3

(-4) on basiscopic side of the pinna, and the basi-

scopic margin entire b3

b3. Lamina apex acuminate to subcaudate; pin-

na apex acute; acroscopic margin deeply and

acutely cleft 40. A. pulchellum

b3. Lamina apex acute; pinna apex obtuse; ac-

roscopic margin crenate-serrate

41. A. discrepans

a3. Pinnae subequilateral, or inequilateral only to-

ward the base; veins 5-many on basiscopic side

(sometimes less in A. tabinense but then the basi-

scopic margin narrowly and deeply crenate) c3

c3. Veins all simple, or proximal 1-3 veins forked

on acroscopic side of pinnae d3

d3. Lamina gradually reduced to rudimen-

tary basal pinnae; petiole very short, '/5-

Vio the length of the lamina e3

e3. Rachis broadly alate, each wing of-

ten as wide as the rachis; acroscopic

pinna margin serrulate to deeply

serrate; elevation 400-1 100 m ...

42. A. pteropus

e3. Rachis narrow-alate; acroscopic

pinna margin deeply and narrowly

crenate to shallowly lobed, and of-

ten with a discrete basal auricle; el-

evation 1 500-3600 m f 3

f3. Basal acroscopic pinna auricle

cut nearly to costa; sori com-

monly 8-12 on larger pinnae .

43. A. sessilifolium

f3. Basal acroscopic pinna auricle

cut (if at all) only halfway to cos-

ta; sori commonly 4-7 on larger

pinnae 44. A. tabinense

d3. Lamina scarcely reduced at base, or sev-

eral pinnae reduced and the basal ones

half the size of largest ones; petiole '/»-

'/2 the length of the lamina g3

g3. Lamina apex with a subcaudate,

proliferous tip ... 45. A. herzogii

g3. Lamina apex pinnatifid, not prolif-

erous h3

h3. Pinna margins subentire to

crenulate; pinnae 1.2-2.2(-2.5)

cm long; stem scales 1.5-3 mm

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

long, blackish, attenuate

46. A. bangii

h3. Pinna margins deeply serrate to

serrately lobed; pinnae (larger

ones) mostly 2.5-9 cm long;

stem scales 4-10 mm long, gray

brown or, if dark brown to

blackish (in A. harpeodes) then

with a long, tortuous, unicos-

tate tip i3

i3. Stem scales deep brown to

blackish, linear, with long,

tortuous, unicostate tips;

pinnae (24-)30-50 pairs;

rachis atropurpureous . . .

47. A. harpeodes

i3. Stem scales gray-brown,

linear- to ovate-lanceolate,

acute to attenuate; pinnae

(12-) 15-24 pairs; rachis

gray- to red-brown

48. A. raddianum

c3. Veins all forked on acroscopic side of pinnae,

except 1-3 apical ones simple j3

j3. Lamina abruptly ending in a proliferous

tip; rachis (and usually petiole) broadly

alate, the wing 1-2 mm broad

49. A. alatum

j3. Lamina tip not proliferous; rachis and

petiole narrowly or vestigially alate, or

occasionally the rachis wing to 1 mm

broad k3

k3. Indusia thin and mostly scarious,

often hyaline until mature .... 13

13. Lamina chartaceous, abruptly

terminating in a broad-based,

nonconform, apical segment;

petiole 12-25 cm long; pinnae

4-12 pairs . 50. A. absciss urn

13. Lamina firm-herbaceous to

membranaceous, gradually re-

duced to a pinnatifid, usually

acuminate, apex; petiole 4-9 cm

long; pinnae 1 5—40 pairs

51. A. drepanoph yllum

k3. Indusia (at least mature ones) firm

and fleshy, opaque m3

m3. Lamina very gradually taper-

ing to a pinnatifid, often sub-

caudate apex; stem scales ecil-

iate 52. A. auritum

m3. Lamina abruptly reduced at

apex, the apical section ser-

rate-ligulate, broad-based and

nonconform, or indefinite;

10

FIELDIANA: BOTANY

stem scales with orange cil-

ia n3

n3. Veins mostly 2-forked;

lamina ending in a broad-

based, nonconform, some-

times hastate, terminal

segment

.... 53. A. salicifolium

n3. Veins simple to 1 -forked;

lamina ending in a nar-

row, serrate-ligulate, or

subcaudatc apex .... o3

o3. Stem scales 6-9 mm

long; pinna apex

commonly attenuate

... 54. A. hastatum

o3. Stem scales 2^*(-5)

mm long; pinna apex

acute or subacute . .

55. A. auriculatum

1. Asplenium serratum L., Sp. pi. 1079. 1753.

LECTOTYPE (designated by Proctor, in

Howard, Flora Lesser Antilles 2: 313. 1977):

Plumier, Descr. pi. Amer. /. 39. 1693.

Plants epiphytic, occasionally terrestrial. Stem

stout, provided with brown-tomentose roots and

usually iridescent, gray-brown (rarely blackish)

scales, these obviously clathrate, linear or linear-

lanceolate, attenuate, (5-)7-18 mm long. Leaves

densely caespitose, sessile or subsessile, simple and

unlobed, larger mature ones 20-70(-100) cm long

4.2-15 cm broad. Petiole 0-4 cm long, '/i0-'/2o the

length of the lamina. Lamina chartaceous, acute

to acuminate or abruptly caudate at the apex, grad-

ually reduced to a long-attenuate base, with entire

to crenulate or bluntly serrate margins, glabrous

and eglandular, but midrib with minute, scattered,

blackish, clathrate scales, at least abaxially. Veins

simple to 1 -forked, spreading at 50-75° from the

midrib. Sori densely crowded, greatly elongated.

Indusia elongated, thin, often scarious.

In deep, wet forests, on tree trunks and branches,

occasionally in alluvial soil on ravine or stream

banks, 100-800(-1200) m, Amazonas and Loreto

to Cuzco and Madre de Dios.

United States (southern Florida); West Indies;

southern Mexico to Argentina and Paraguay.

This and similar species such as the Old World

A. nidus L., have been given the common name

"Bird's Nest Ferns," due to their growth habit.

The large, densely caespitose leaves of older plants

often form a tight circle, the center of which then

becomes a receptacle for a mixture of organic mat-

ter that falls, or is blown in, from above. With this

perhaps should be included A. angustum and A.

stuebelianum, which are probably only variants.

The two taxa differ from A. serratum merely by

subtle characters of leaf shape, vein angle, and

stem scales, and a number of specimens examined

during this study are clearly intermediates. Stem

scales of A. serratum are typically 8-15 mm long,

gray-brown, and conspicuously clathrate, but oc-

casionally they are smaller, darker, and more nar-

rowly clathrate like those of A. stuebelianum.

Some specimens of A. pearcei have simple leaves

and are often determined as A. serratum. How-

ever, the former have long petioles like those of

A. stuebelianum as well as several other differing

characters, which are discussed under that species.

Amazonas: Prov. Bagua, NE of Chiriaco, Harbour 45 13

(F, MO, USM). San Martin: Chazuta, Rio Huallaga, Klug

4140 (F, OH). Loreto: Gamitanacocha, Rio Mazan, J.

Schunke 276 (F, GH, uc). Huanuco: Prov. Huanuco, near

confluence of Rio Cayumba with Huallaga, Mexia 8276

(F, OH, MO, uc, us). Pasco: Prov. Oxapampa, Palcazu

valley, Iscozacin, Foster 7973 (F, MO). Junin: Chancha-

mayo, La Merced, Soukup 101 1 (F). Ucayali (as Lorelo):

Prov. Coronel Portillo, between Divisoria and Boque-

ron, Ferreyra 4273 (GH, USM). Cuzco: Prov. La Conven-

tion, Rio Apurimac, below Puerto Capiro, Davis et al.

1300 (F, GH). Madre de Dios: Prov. Manu, Manu Na-

tional Park, Cocha Cashu Station, Foster et al. 6629 (F),

69/6 (F).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

11

2. Asplenium angustum Sw., Kongl. Vetensk.

Acad. Handl. 38: 66, t. 4,f. 1. 1817. TYPE:

"Habitat in Brasilia," Freyreis (holotype, s;

photos, F, us).

Plants epiphytic. Stem erect, provided with

brown-tomentose roots and blackish scales, these

linear, 4-5(-6) mm long including the attenuate,

unicostate tip, obscurely clathrate. Leaves caes-

pitose, sessile or subsessile, simple, 8-30(-40) cm

long, 0.5-3(-4) cm broad. Petiole 0-2 cm long,

'/io-'/2o the length of the lamina. Lamina firm-her-

baceous to chartaceous, long-attenuate to apex and

base, lacking proliferous buds, margins entire, gla-

brous and eglandular, but midrib provided with

scattered, minute, dark, clathrate scales, at least

abaxially. Veins simple to 1 -forked, spreading at

30-45° from the midrib. Sori approximate to sub-

distant, linear to elliptic. Indusia linear to elliptic,

whitish to pale greenish, relatively thick, not trans-

lucent.

In wet forests, on tree trunks, 100-1 100 m, Lo-

reto, Pasco, Madre de Dios.

The Guianas to Colombia; Peru; Brazil.

This is but provisionally maintained as distinct

from A. serratum. Whereas the extreme forms of

each appear to be discrete, a number of specimens

toward the edge of the range tend to be interme-

diate. Typical A. angustum, especially in Brazil,

has leaves less than 2.5 cm broad and very grad-

ually tapering to apex and base, the veins borne

at an angle of less than 45° from the midrib, the

indusia relatively broad and thick-textured, and

whitish and obscure, not crowded, and the stem

scales blackish and obscurely clathrate. This con-

trasts greatly with typical A. serratum, with lamina

8-12 cm broad and abruptly acute or acuminate

at apex, the veins at angles of 50-60°, the indusia

very long and narrow, scarious and densely crowd-

ed, and the stem scales brownish, iridescent, and

obviously clathrate. However, a number of small-

er specimens of A. serratum exhibit nearly the size

and vein angle of A. angustum and have blackish

and less conspicuously clathrate stem scales. Sev-

eral of the specimens from Peru cited below are

approximately intermediate but have features

much closer to A. angustum than to A. serratum

and therefore are so identified here. Monographic

study may prove the former to be merely a variant

of the latter.

Loreto: Prov. Maynas, Dist. Iquitos, Rio Momon, be-

low Balcon, Hickock 606 (GH). Lower Rio Momoncillo,

lower Rio Momon, near Iquitos, Jones & Davidson 9515

(MO, us). Prov. Maynas, Mishana, van der Werff el al.

10181 (MO, uc). Pasco (as Junin): Pichis Trail, San Ni-

colas, Killip & Smith 30698 (us). Madre de Dios: Tam-

bopata Nature Reserve, SE bank of Rio Tambopata,

Barbour 5185 (MO). Parque Nacional del Manu, Cocha

Cashu Biological Station, M. Foster P-84-52 (uc).

3. Asplenium pseudoangustum Stolze, Amer. Fern

J. 74: 49. 1984. TYPE: Peru, Huanuco, Tingo

Maria, Tryon & Tryon 5257 (holotype, GH!;

isotypes, F!, us!, USM!). Figure la.

Plants epiphytic. Stem small, erect to decum-

bent, provided with dark or light brown, clathrate

scales, these lanceolate or ovate, acute to acumi-

nate, 1.5-3 mm long. Leaves caespitose, sessile or

subsessile, simple, 8-22 cm long, 1.3-2.5(-3) cm

broad. Petiole essentially lacking. Lamina firm-

herbaceous, subcarnose, elliptic, apex acute

(sometimes somewhat attenuate), long-attenuate

at base, lacking proliferous buds, margins entire,

surfaces and midrib glabrous and lacking scales.

Veins 1 (-2)-forked, obscure, spreading at 30^45°

from the midrib. Sori well spaced, linear. Indusia

linear, pale green or yellow-green, not translucent,

becoming dark brown and thicker at maturity.

On tree trunks in wet forests, 350-1400 m, San

Martin to Cuzco and Madre de Dios.

Ecuador; Peru.

Although resembling A. angustum in size and

shape, this is quite a distinct species, as evidenced

by the key characters. In addition, A. pseudoan-

gustum has somewhat succulent leaves that often

commonly dry blackish green. Leaves of A. an-

gustum dry gray- or yellow-green.

San Martin: Prov. Mariscal Caceres, between Aucaya-

cu and Uchiza, Ferreyra 1 7038 (GH, USM). Prov. Mariscal

Caceres, Tocache Nuevo, /. Schunke V. 567 '1 (F, us).

Huanuco (as San Martin): Tingo Maria, Allard 21596a

(us), 22384 (us). Pasco: Paujil, near Puerto Bermudez,

Leon et al. 292 (GH, USM). Prov. Oxapampa, Valle de

Palcazu, near Iscozacin, Leon 715 (F, GH). Cuzco: Prov.

La Convention, Tupitani, Bites 5450 (us). Madre de

Dios: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa,

Foster et al. 10827 (F).

4. Asplenium stuebelianum Hieron., Hedwigia 47:

222. 1908. TYPE: Colombia, Prov. Cundi-

namarca, near Villavicencio and Oca, Stiibel

659 (holotype, presumably B).

Plants epiphytic, occasionally terrestrial. Stem

stout, erect or decumbent, provided with brown-

tomentose roots and gray-brown to blackish scales,

12

FIELDIANA: BOTANY

these obscurely to narrowly clathrate, linear or lin-

ear-lanceolate, attenuate, 2-4(-6) mm long. Leaves

densely caespitose, usually conspicuously petio-

late, simple and unlobed, larger mature ones 25-

50 cm long, 3.5-9 cm broad, proliferous buds oc-

casionally borne at the apex. Petiole (3-)5-18 cm

long, usually '/4-'/2 the length of the lamina. Lamina

chartaceous, acute to acuminate or abruptly cau-

date at the apex, at the base abruptly reduced and

then long-attenuate, margins entire to crenulate or

bluntly serrate, glabrous and eglandular, but mid-

rib with minute, scattered, blackish, clathrate

scales, at least abaxially. Veins simple to 1 -forked,

spreading at 60-75° from the midrib. Sori densely

crowded, greatly elongated. Indusia elongated, thin,

often scarious.

In deep, wet forests, on tree trunks and branches,

occasionally in alluvial soil or ravine or stream

banks, 100-1400 m, Amazonas and Loreto south

to Junin, Madre de Dios.

Venezuela; Colombia; Peru; Bolivia; Brazil.

Asplenium stuebelianum is perhaps merely a va-

riety of A. serratum, from which it differs princi-

pally in the petiolate or subpetiolate lamina. In

the latter the lamina tapers very gradually and

regularly to the stem, with petioles (if any) only a

few centimeters long. The lamina in A. stuebe-

lianum is reduced suddenly and markedly and then

is attenuate to a narrowly winged or naked petiole,

which is typically 6-12 cm long. For further com-

parison see treatment of A. serratum. The Wur-

dack collection, cited below from Amazonas, is

intermediate between the two taxa, in characters

of both lamina and stem scales.

Amazonas: Prov. Bagua, Rio Maranon above Casca-

das de Mayasi, Wurdack 1971 (F, GH, uc, us). San Mar-

tin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Puer-

to Pizana, Schunke V. 6947 (F). Loreto: Mishiyacu, near

Iquitos, Klug 1537 (F, us). Huanuco: Prov. Pachitea,

Dist. Puerto Inca, Bosque Nacional de Iparia, Schunke

V. 2968 (F, GH, us). Junin: Puente Perene, Coronado 254

(GH). Schunke Hacienda above San Ramon, Killip &

Smith 24705 (F). Madre de Dios: Prov. Manu, Atalaya,

Foster & Wachter 7461 (F). Prov. Manu, Cerro de Pan-

tiacolla, Rio Palotoa, Foster et al. 10694 (F).

5. Asplenium pearcei Baker, Syn. fil. ed. 2: 483.

1874. TYPE: Peru (Pasco), banks of Rio

Huancabamba, Pozuzo, Pearce 528 (holo-

type, K.!; photo, F).

Asplenium amazonicum Christ, Hedwigia 45: 191.

1906. LECTOTYPE (designated here): Brazil, Rio

Jurua, Vie 5525 (P!; isolectotype, K!; photo, F of

p). Vie 5525 (L!) is A. stuebelianum (mixed col-

lection).

Asplenium haplophyllum Domin, Pterid. Dominica

170, t. 29, f. 1 (Vie 5525, K.!). 1929.

Plants low-epiphytic, rarely terrestrial. Stem

stout, erect or decumbent, provided with clathrate,

gray-brown scales, these lanceolate, acuminate, 2-

4 mm long and mostly 1 mm broad. Leaves fas-

ciculate, 24-40 cm long, mostly long-petiolate,

simple to trifoliolate or (very rarely) with 2 pairs

of pinnae. Petiole commonly 5-18 cm long, dull

reddish or greenish brown, sparsely clathrate-scaly

at base. Lamina thin- to firm-herbaceous, in pin-

nate leaves the apical segment conform but larger

than lateral ones, sparsely to moderately provided

with minute (0.1-0.2 mm), appressed, pluricel-

lular trichomes, these often gland-tipped and vis-

ible only under high magnification, rachis (in pin-

nate specimens) and proximal part of petiole alate.

Pinnae (when present) opposite or subopposite,

narrowly and subequally cuneate at base, acute or

acuminate at apex, narrowly adnate to the rachis,

margins entire to crenulate. Veins 1-2-forked, in-

distinct. Indusia linear, green or yellow-green, en-

tire.

In wet forests, low on tree trunks, on fallen logs,

or rarely in wet, rocky soil, 1 00-500 m, San Martin

and Loreto south to Puno.

Venezuela; the Guianas; Ecuador; Peru; Brazil.

Two strange occurrences have led to confusion

attending the names of synonyms of A. pearcei.

First, Ule 5525, one of the syntypes of A. ama-

zonicum, represents a mixed collection, for the

specimen at Leiden is A. stuebelianum, a distinct,

albeit superficially similar, species. Second, Do-

min, obviously unaware of the identity of A. ama-

zonicum, later created A. haplophyllum, based on

Ule 5525 at Kew (here designated isolectotype of

the former name).

Specimens of^4. pearcei with simple leaves have

been confused with A. serratum, A. angustum, and

A. pseudoangustum. It is distinguished from these

three by the long petiole and thinner lamina and

from the first two by lack of small, black scales

along the costa abaxially.

San Martin: Prov. San Martin, road from Tarapoto

to Yurimaguas, Plowman 6001 (GH, USM). Prov. Maris-

cal Caceres, Tocache Nuevo, Schunke V. 7069 (F, MO,

USM). Loreto: Gamitanacocha, Rio Ma/an. J. Schunke

305 (F, GH, uc, us). Pasco (as Junin): Cahuapanas, on

Rio Pichis, Killip & Smith 26736 (us). Ucayali (as Lo-

reto): Rio Aguaytia above mouth of Yurac-Yacu, Croat

20905 (MO). Madre de Dios: Prov. Tambopata, Albergue

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

13

Cuzco Amazonico, Leon 890 (F, USM). Puno: Prov. Cara-

baya, Hacienda Palmera, Inambari, Vargas 16142 (GH).

6. Asplenium escaleroense Christ, Hedwigia 44:

366. 1905. TYPE: Peru (Loreto), "Cairo de

Escalero," Ule 6886 (holotype, P; isotype, B?).

Plants epiphytic, occasionally epipetric. Stems

minute, small and erect, often arising at subdistant

intervals from long-creeping, stoloniform roots,

scales lacking, or a few dark brown, clathrate ones

at the petiole base. Leaves single, or several and

fasciculate, 2-6(-9) cm long, 0.2-0.6(-1.0) cm

broad. Petiole thin, 0.4-2 cm long, green or green-

ish brown. Lamina thin-herbaceous, linear or lin-

ear-lanceolate (or juvenile ones subflabellate to

obovate), glabrous, shallowly lobed to pinnatifid,

or rarely with a pair of lobed basal pinnae. Seg-

ments or lobes 2-6 pairs, strongly ascending. Veins

simple in each segment, often indistinct. Sori one

per segment. Indusia to 3 mm long and 0.5 mm

broad, yellowish or greenish.

In rain forests, on trunks or branches of trees,

or occasionally on wet rocks or cliffs near water-

falls, 130-1200 m; in Peru thus far known from

the type and two other collections.

Colombia; Ecuador; Peru; Bolivia.

Scattered collections have been made of this

species from Colombia to Bolivia, but the leaves

are tiny and inconspicuous and therefore easily

overlooked, so it is likely to be far more common

within its reported range. It should be searched

for throughout Peru in wet forests, on trees, and

on rocks around waterfalls.

San Martin: Tarapoto-Yurimaguas Hwy., km 14-17,

McDaniel 13802 (GH). Loreto: Prov. Maynas, Yana-

mono Explorama Tourist Camp, van der Werff et al.

9923 (MO, uc).

7. Asplenium pumilum Sw., Prodr. 129. 1788.

TYPE: Swartz, Jamaica (holotype, s).

Plants terrestrial or epipetric. Stem small, erect,

provided at apex with filiform, blackish scales to

3 mm long. Leaves 1 -pinnate or -pinnatisect, sub-

caespitose, mature ones 4-20 cm long, 1.5-6 cm

broad. Petiole thin, 1-1 Ocm long, green to lustrous

and castaneous or atropurpureous at base abaxi-

ally. Lamina firm-herbaceous to somewhat fleshy,

not reduced at base, terminating in a nonconform,

pinnatifid apex, provided, especially on rachis,

veins, and margins, with few to many whitish,

septate trichomes. Pinnae 1-4 pairs, sessile and

adnate distally, the proximal ones often short-

stalked, subentire to crenate or lobed, basal pair

the largest, more strongly produced basiscopically

than acroscopically. Veins 1 - or 2-forked, or pin-

nately branched in basal lobes. Sori linear or el-

liptic. Indusia thin, yellow to whitish.

Rare, in wet forests, on rocks, or on the forest

floor, 400-1800 m, Tumbes, Huanuco, Cuzco.

United States (Florida); West Indies; Mexico to

Panama; Venezuela and Colombia to Brazil and

Argentina; Africa.

This is one of a few species in the genus having

basal pinnae more strongly produced on the basi-

scopic side. This and the whitish, septate tri-

chomes of the lamina easily separate it from other

species in Peru. Although it is rarely collected, it

is probably more common than thus far reported,

as it is but another of the very inconspicuous spe-

cies in Asplenium.

Tumbes: Between Caucho and Cotrina, Coronado 221

(GH, uc). Between Tumbes and Caucho, Coronado 225

(GH, uc). Huanuco: Yanano, Macbride 3815 (F). Cuzco:

Prov. La Convencion, Potrero, Bties A40 (us). Prov. La

Convencion, Playa de Balsa, Mexia 8053b (uc).

8. Asplenium formosum Willd., in L. Sp. pi. ed.

4, 5: 329. 1810. TYPE: Venezuela, Caracas,

Bredemeyer(ho\oiype,B,Herb. Willd. 1990S;

photos, F, GH).

Plants terrestrial, occasionally epipetric or epi-

phytic. Stem small, erect, provided with rigid, lin-

ear scales, these 1-3 mm long, obscurely clathrate,

black with narrow, brown margins. Leaves 1 -pin-

nate, caespitose, to 30 cm long and 2.5 cm broad.

Petiole rather stout, 1.5-4 cm long, lustrous, black

or atropurpureous, glabrous. Lamina firm-herba-

ceous to chartaceous, tapering to a pinnatifid apex,

gradually narrowed at base to mere auricles, gla-

brous, lacking proliferous buds, rachis adaxially

with narrow perpendicular wings. Pinnae numer-

ous, 0.6-1.5 cm long, patent, sessile, narrow and

subacute, subdimidiate, basiscopically cuneate at

base and the margin entire to biserrate, acroscop-

ically truncate and the margin biserrate to deeply

and sharply incised. Veins simple or 1 -forked, in-

distinct. Sori 1-3 to a pinna, elliptic to ovate, often

confluent at maturity. Indusia pale green to whit-

ish.

In wet forests, usually in loose soil of banks,

occasionally on tree trunks or in crevices of wet

14

FIELDIANA: BOTANY

cliffs, 200-1 200 m, Loreto, Junin, Ayacucho, Cuz-

co.

West Indies; Mexico to Panama; the Guianas

to Colombia, south to Brazil and Argentina.

Loreto: Banks of Rio Santiago above Pongo de Man-

seriche, Mexia 6318 (F, GH, MO, uc, us). Junin: Rio

Pinedo N of La Merced, Killip & Smith 23566 (F). Puente

Quimiri, Leon 469 (USM). La Merced, Macbride 5373

(F). Ayacucho: Estrella, between Huanta and Rio Apu-

rimac, Killip & Smith 23077 (us). Cuzco: Prov. La Con-

vencion. Potrero, W of Quillabamba, Tryon & Tryon

5382 (GH). Prov. La Convention, Rosalina, Vargas 12288

(GH).

9. Asplenium extensum Fee, Mem. foug. 7: 51, /.

13. 1857. TYPE: Colombia, Ocana, Schlim

629 (holotype, P; isotypes, L!, RB; frag., us! of

L; photos, F, GH & us of L).

Plants terrestrial or epipetric. Stem short, erect

or decumbent, provided with linear-lanceolate

scales, these 1-2 mm long, obscurely clathrate,

black, with narrow brown margins. Leaves 1 -pin-

nate, subcaespitose, to 40 cm long and 2 cm broad.

Petiole 3-5 cm long, terete, lustrous, atropurpu-

reous to blackish, essentially glabrous. Lamina

chartaceous to subcoriaceous, linear, strongly and

gradually tapering to apex, not or rarely reduced

at base, occasionally with proliferous buds at apex

or in pinna axils. Rachis adaxially provided with

strong parallel, winglike ribs, these conspicuously

hirsute with rigid, spreading, castaneous trichomes

to 0.3 mm long. Pinnae numerous, to 1 cm long,

patent, approximate to remote, sessile, oblong,

subequilateral, margins entire or broadly and shal-

lowly crenate. Veins obscure, but their tips com-

monly enlarged and conspicuous near the pinna

margin. Sori 2-7 to a pinna, sometimes confluent

at maturity. Indusia large, persistent, pale green

to whitish.

In forests, in soil, or in crevices of wet, rocky,

canyon walls, 1 800-3950 m, Cajamarca and Ama-

zonas to Junin.

Colombia; Peru.

Cajamarca: Prov. Celendin, canyon of Rio Maranon

above Balsas, Hutchison & Wright 5294 (F, GH, uc, us).

Amazonas: Prov. Chachapoyas, Cerros Calla Calla above

Leimebamba, Hutchison & Bennett 4546 (F, GH, uc, us).

San Martin: Prov. Mariscal Caceres, Rio Abiseo Na-

tional Park, Young 4224 (HUT). Huanuco: Yanano, Mac-

bride 3831 (F, GH, us). Tambo de Vaca, Macbride 4380

(F, us). Junin: Prov. Chanchamayo, Mina Pichita, van

der Werffet al. 8660 (MO, uc).

10. Asplenium resiliens Kunze, Linnaea 18: 331.

"1844" (1845), nom. nov. for Asplenium par-

vulum Mart. & Gal. and with the same type.

Asplenium parvulum Mart. & Gal., Nouv. Mem. Acad.

Roy. Sci. Bruxelles 15: 60. 1842, nom. illeg. (not

Hooker, 1840). TYPE: Mexico, Oaxaca, Galeotti

6462 (holotype, BR, photos, BM, F, us; isotype, BR,

photos, P, us).

Plants epipetric, rarely terrestrial. Stem short,

erect, provided at apex with linear to filamentous,

blackish scales, these 3-4 mm long, often with

narrow brown margins, obscurely clathrate. Leaves

1 -pinnate, densely caespitose, to 35 cm long and

2.5 cm broad. Petiole 1-5 cm long, lustrous, atro-

purpureous to blackish, essentially glabrous. Lam-

ina subcoriaceous, linear, gradually reduced to base

and diminished abruptly to a pinnatifid apex, lack-

ing proliferous buds. Rachis adaxially with parallel

ribs, each of these with a narrow but pronounced,

foliaceous, green wing nearly throughout. Pinnae

numerous, large ones 0.5-1.2 cm long, patent to

slightly deflexed, subsessile or short-stalked, ellip-

tic or broadly oblong, inequilateral at base, cu-

neate basiscopically, truncate and often subauricu-

late acroscopically, margins entire to broadly

crenulate. Veins obscure or indistinct, the tips not

dilated. Sori several pairs to a pinna, mostly

crowded at the pinna margins, usually confluent

at maturity. Indusia broad, pale yellow to whitish,

persistent, but often obscured at maturity by the

numerous sporangia.

In forests, on ledges and in crevices of rocky

cliffs, or on boulders, rarely in rocky soil, 1200-

3500 m, Cajamarca and Amazonas south to Cuz-

co.

Southern United States; Jamaica; Hispaniola;

Mexico; Guatemala; Colombia and Venezuela

south to Argentina and Brazil.

Cajamarca: About 40 km from Cajamarca on road to

Chilete, Correll & Smith P844 (GH, us), P844A (GH).

Amazonas: Prov. Chachapoyas, Jazan (Ingenio-Chacha-

poyas), Lopez et al. 4262 (GH, HUT). La Libertad: Prov.

Sanchez Carrion, between Sausagocha and Cajabamba,

Smith & Vasquez 3382 (MO, uc). Ancash: Prov. Carhuas,

Cordillera Blanca, Valley of Rio Marcara, Hutchison &

Wright 4362 (F, GH, uc, us). Huanuco: Huacachi, esta-

cion near Muna, Macbride 3871 (F, GH, us). Junin: South

of Huancayo, between Viques and Ingahuasi, Killip &

Smith 22157 (F, us). Apurimac: 5 km above Huamcara-

ma, West 3802 (uc). Cuzco: Road from "Ccasapata"

(Casapata) to Apurimac, Sues 1388 (us).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

15

11. Asplenium monanthes L., Mant. pi. 130. 1767.

TYPE: South Africa, Cape of Good Hope (ho-

lotype, LINN 1250.17; photo, us).

Plants epipetric, occasionally terrestrial, rarely

epiphytic. Stem short, erect, provided at apex with

filiform to narrow-deltate scales, these 1-5 mm

long, gray-brown to blackish, scarcely to obviously

clathrate. Leaves 1 -pinnate, caespitose, to 60 cm

long and 3 cm broad. Petiole 1-15 cm long, lus-

trous, dark brown, atropurpureous or blackish, es-

sentially glabrous. Lamina firm-membranaceous

to chartaceous, linear, tapering to a pinnatifid apex

and strongly reduced at base. Rachis sparsely pro-

vided with dark, fibrillose scales, adaxially with 2

parallel raised ribs, not proliferous at apex but, in

var. wagneri, often with scale-covered gemmae in

a distal pinna axil. Pinnae (larger ones) to 1.5 cm

long, sessile to minutely stalked, patent, blunt and

quadrangular to subacute and falcate, inequilateral

at base, often subauriculate acroscopically. Veins

often indistinct, but their tips conspicuous and

somewhat to strongly dilated adaxially. Sori sol-

itary near the basiscopic pinna margin, or borne

in several pairs, often confluent at maturity. In-

dusia broad, pale green to whitish, persistent, but

often obscured at maturity by spreading sporangia.

The Southern United States; Jamaica; Hispan-

iola; Mexico south to Panama; Colombia and Ven-

ezuela south to Argentina and Chile; Hawaii; Af-

rica.

Authors have recognized Asplenium monanthes

and two closely related species separated by a few,

mostly quantitative, features. Some of these char-

acters are relatively constant in certain parts of the

range, but variable in other areas, and interme-

diate specimens are not uncommon. (See discus-

sion of var. monanthes for detailed comparison of

characters.) In this treatment the three taxa are

recognized as geographical or elevational variants,

a key to which is provided here:

a. Ribs on adaxial side of rachis conspicuously pustulate or pustulate-dentate; stem scales l-2(-3) mm

long; a proliferous bud sometimes borne in a distal pinna axil lie. var. wagneri

a. Ribs on adaxial side of rachis entire to remotely and minutely denticulate; stem scales mostly 4-5

mm long; proliferous buds lacking b

b. Stem scales commonly blackish, or with a narrow brown margin, not or obscurely clathrate; pinnae

commonly 2-4 times as long as broad 1 la. var. monanthes

b. Stem scales commonly gray-brown, clathrate; pinnae l-1.5(-2) times as long as broad

1 1 b. var. castaneum

1 la. Asplenium monanthes var. monanthes.

Asplenium monanthemum Murray, in L., Syst. veg.

ed. 14: 933. 1784. nom. superfl. for Asplenium

monanthes and with the same type.

Stem scales commonly 4-5 mm long, blackish,

or with narrow brown margins, linear to filiform,

not or obscurely clathrate, lumina narrow and their

walls thick. Leaves to 60 cm long and 3 cm broad.

Petiole atropurpureous (sometimes dark brown or

blackish). Rachis with adaxial ribs entire to mi-

nutely denticulate, lacking proliferous buds. Pin-

nae (larger ones) mostly 2—4 times as long as broad,

subentire to crenate or crenate-serrate.

In forests and thickets, on and among rocks, in

crevices of rock cliffs, and occasionally in rocky

soil, (900-) 1300-4350 m, Lambayeque to Ama-

zonas, south to Ayacucho and Madre de Dios.

Southern United States; Jamaica; Hispaniola;

Mexico to Panama; Colombia and Venezuela south

to Argentina and Chile; Hawaii; Africa.

The range of var. monanthes is very broad; it

occurs at middle to very high elevations in Peru.

Variety castaneum does not occur in the Old World,

the West Indies, or middle Central America, is

(with a few exceptions) confined to Andean regions

in South America, and is found mostly over 3000

m in Peru. Variety wagneri thus far has been found

only in Colombia and Ecuador, and in Peru com-

monly between 3000 and 3800 m. The last has the

smallest leaves of the three varieties and is easiest

to recognize because of the pustulate rachis ribs,

the much smaller stem scales, and the occasional

rachis buds. Darker clathrate scales, darker peti-

oles, and relatively narrower pinnae usually dis-

tinguish var. monanthes from var. castaneum, but

all these vary frequently enough so that compar-

ison of a suite of characters is usually necessary

for positive identification. Some characters seem

to be more constant in certain geographic areas

than in others; consequently, the above key is es-

pecially designed to separate the varieties in Peru.

Lambayeque: Prov. Ferrenafe, Quiros 2521 (F). Ca-

16

FIELDIANA: BOTANY

jamarca: Prov. Celendin, Pumarrume, Mostacero el al.

1011 (F, GH, HUT, uc). Amazonas: Prov. Chachapoyas,

Cerros Calla Calla, above Balsas, Hutchison & Wright

5824 (F, GH, uc, us, USM). La Libertad: Prov. Santiago

de Chuco, 26 km from Santiago, D. Smith 2334 (F, MO).

San Martin: Dist. Huallaga. Valley of Rio Aposoncho,

Hamilton & Holligan 500 (us), 914 (us). Ancash: Prov.

Corongo, Nueva Victoria, Mostacero el al. 2009 (F, HUT).

Huanuco: Muna, Macbride 3939 (f, GH, us). Lima: Prov.

Yauyos, Laguna Huacracocha, Cerrate 1255 (USM). Pas-

co: Huariaca, Bryan 169 (F). Junin: Prov. Tarma, 5 km

SW of Huacapistana, Tryon & Tryon 5425 (F, GH, us).

Ayacucho: Prov. Huamanga, Totorabamba, Weberbauer

5469 (GH). Apurimac: Prov. Abanbcay, forests of Am-

pay, Vargas 1068 (F, GH). Cuzco: Prov. Urubamba, near

town of Machu Picchu, Tryon & Tryon 5405 (F, GH, us).

Mad re de Dios: Pinasniocj, Panticalla [Pantiacolla?] Pass,

Cook & Gilbert 1798 (us).

lib. Asplenium monanthes var. cast am* urn

(Schlecht. & Cham.) Stolze, Flora of Ecuador

23:45. 1986.

Asplenium castaneum Schlecht. & Cham., Linnaea 5:

611. 1830. TYPE: Mexico, Veracruz, Mt. Ori-

zaba, Schiede (holotype, HAL; frag., NY!).

Stem scales commonly 4-5 mm long and gray-

brown, linear to narrow-deltate, usually conspic-

uously clathrate, the lumina broad and clear and

the walls thin. Leaves to 35 cm long and 2 cm

broad. Petiole dark brown, sometimes castaneous,

rarely atropurpureous. Rachis with adaxial ribs

subentire, lacking proliferous buds. Pinnae (larger

ones) l-1.5(-2) times as long as broad, subentire

to crenate.

In forests, on rocks and in crevices of rock cliffs,

and sometimes in rocky soil, (600-)2900-4700 m,

Cajamarca and Amazonas south to Huancavelica

and Puno.

Southern Mexico; Guatemala; Costa Rica; Pan-

ama; Venezuela and Colombia south to Bolivia.

Authors often distinguish this from var. monan-

thes by the petiole color. Petioles of var. casta-

neum usually are dark brown and those of var.

monanthes atropurpureous, but, at least in Peru,

petioles on the same stem can be of both colors.

Furthermore, Stolze (1981) pointed out that in

Guatemala the short pinna stalks were usually green

in "A. monanthes," but often partly brown in "A.

castaneum." This character is useless in Peru, as

pinna stalks are nearly always concolorous in both

varieties.

Cajamarca: Prov. Contumaza, Jalca El Chuno, Sa-

gdstegui el al. 9368 (F, MO). Amazonas: Prov. Chacha-

poyas, Cerro Campanario, Wurdack 1558 (GH, uc, us).

La Libertad: Prov. Santiago de Chuco, near Santiago,

Sagdstegui el al. 11708 (HUT). San Martin: Prov. Maris-

cal Caceres, Rio Abiseo National Park, Young & Leon

4576 (USM). Ancash: Prov. Yungay, near Laguna Llan-

ganuco, Mostacero el al. 1401 (F, HUT, MO). Lima: Prov.

Huarochiri, Dist. San Mateo, Rio Blanco, Sounders 404

(F, uc). Pasco: Between La Quinua and Tambo, Soukup

5627 (us). Junin: Mt. La Juntay, near Huancayo, Killip

& Smith 22058 (GH, us). Huancavelica: Prov. Castro-

virreina, Choclococha, Tovar 2842 (USM). Cuzco: Prov.

Espinar, Imantata, Vargas 10509 (GH). Puno: Granja

Salcedo, Canon Viscachani, Mexia 4263 (uc).

1 Ic. Asplenium monanthes var. wagneri (Kuhn)

Stolze, Flora of Ecuador 23: 46. 1986.

Asplenium wagneri Kuhn, Linnaea 36: 96. 1 869. TYPE:

Ecuador, Pichincha, near "Turtillas" between

Quito and Esmeraldas, Wagner (holotype, M; iso-

type, BM!; photos, BM & uc of M, GH of BM).

Asplenium vargasii Abbiatti, Darwiniana 14: 61, /. 1.

1966. TYPE: Cuzco, "Sajasaihuaman" (Saxihua-

man), Vargas 362 (holotype, LIL).

Stem scales l-2(-3) mm long, blackish, linear

or narrow-deltate, not or obscurely clathrate.

Leaves to 22 cm long and 1.3 cm broad. Petiole

castaneous or atropurpureous. Rachis with adaxial

ribs conspicuously pustulate or pustulate-dentate,

sometimes with a proliferous bud in one of the

distal pinna axils. Pinnae 1-1.5 times as long as

broad, subentire to crenate or crenate-serrate.

In forests, on rocks or in crevices or rock cliffs,

or in rocky soil, (2700-)3000-3800(-4350) m, An-

cash, Junin, Cuzco.

Venezuela; Colombia; Ecuador; Peru.

In Ecuador, pinnae are sometimes deeply cre-

nate to shallowly lobed, but in specimens from

Peru they are commonly merely crenate-serrate.

The type of A. vargasii has not been examined,

but the diagnostic features of var. wagneri (deeply

dentate rachis ribs and short stem scales) are clear-

ly indicated in the former's description and illus-

tration.

Ancash: Prov. Bolognesi, Cochacutan, at the foot of

"El Carnicero," Cerrate 2294 (GH). Junin: Prov. Huan-

cayo, Dist. Huancayo, foot of Nevada Huaytapallana.

Sounders 1165 (GH). Cuzco: Prov. Cuzco, Hills of Saxi-

huaman, Herrera 1349 (us), 2370 (F), 2380 (F). Prov.

Cuzco, Saxihuaman. Tryon & Tryon 5348, in part (GH,

us).

12. Asplenium rutaceum (Willd.) Men.. Abh.

Senckenberg Naturf. Ges. 3: 173. 1858.

Aspidium rutaceum Willd., Sp. pi. ed. 4, 5: 266. 1810.

TYPE: Plumier, Traite foug. Amer. /. 57. 1805.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

17

Asplenium perkinsii Jenman, Gard. Chron. 3, 19: 8.

1896. TYPE: Guyana (as British Guiana), Kaie-

teur Falls, Perkins (holotype, NY!).

Asplenium conquisitum Christ, Bull. Herb. Boissier 2,

7: 270. 1907. LECTOTYPE (designated by Max-

on, Contr. U.S. Natl. Herb. 10: 488. 1908): Ja-

maica, Maxon 1558 (P; isolectotype, us!).

Plants epiphytic or terrestrial. Stem erect, pro-

vided with linear or lanceolate scales, these to 3

mm long, clathrate, castaneous or gray-brown.

Leaves 2-3-pinnate, rigid, fasciculate, to 40 cm

long and 10 cm broad. Petiole 0-2 cm long, lus-

trous, castaneous to atropurpureous, glabrous.

Lamina thin-herbaceous, gradually reduced to a

naked and flagelliform apex, this often with a pro-

liferous and radicant tip, strongly reduced at base,

surface apparently glabrous but sparsely provided

abaxially with minute, appressed trichomes. Ra-

chis lustrous, dark brown or atropurpureous, nar-

rowly alate adaxially. Pinnae 1 5-32 pairs, patent

to slightly ascending, or proximal ones deflexed,

sessile, the basal pinnules commonly overlapping

the rachis. Pinnules 6-10 pairs, inequilateral at

base, with several pairs of spatulate or obovate

segments, their apices obtuse to acute (occasion-

ally apiculate). Veins simple in most ultimate seg-

ments. Sori solitary on the vein of each segment.

Indusia thin, subentire, yellowish or greenish.

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

banks of ravines, 800-2600 m, Cajamarca and

Amazonas south to Cuzco.

Greater Antilles; southern Mexico to Panama;

the Guianas to Colombia and south to Bolivia.

This is often confused with other 2-3-pinnate

species of Asplenium in Peru, such as A. myrio-

phyllumandA. radicansvur. uniseriale. However,

only A. rutaceum has the combination of such

diagnostic features as: petiole dark, lustrous, and

nearly obsolete; lamina strongly reduced at base

and apex, with a naked, flagelliform, and usually

radicant tip.

Lellinger (Proc. Biol. Soc. Wash. 98: 372. 1985)

separated A. maxonii Lell. from A. rutaceum in

South America and southern Central America on

a number of subtle characters and cited one para-

type from Peru: Woytkowski 34518 (us). How-

ever, upon reexamination, he has advised (pers.

comm.) that the latter is actually A. rutaceum, thus

A. maxonii has not yet been found in Peru. The

latter is presumed to differ in having adnate ulti-

mate segments and the basal acroscopic lobe of

pinnules usually bifid. Ultimate segments of A.

rutaceum are short-stalked, and basal lobes are

never bifid. Other features cited in the original

description of A. maxonii vary too greatly in Pe-

ruvian specimens to be of significant value.

Cajamarca: Prov. Jaen, above Tabaconas, Quebrada

de Pajonal, Fosberg 27803 (us). Amazonas: Prov. Bagua,

E of La Peca, Barbour 2821 (F, MO, us). San Martin:

Prov. Rioja, Venceremos, D. Smith 4432 (F, MO, uc).

Huanuco: Prov. Leoncio Prado, between Tingo Maria

and Pucallpa, Sullivan & Young 1 159 (MO). Pasco: Prov.

Oxapampa, trail to summit of Cordillera Yanachaga, D.

Smith et al. 7815 (F, MO, USM). Junin: Prov. Tarma, Rio

Palca Valley, Hodge 6299 (GH). Ucayali (as Huanuco):

Divisoria, Woytkowski 34518 (uc, us). Cuzco: Prov.

Paucartambo, Cosnipata Valley, Rio Tono, Wachter et

al. 187 (F).

1 3. Asplenium hallii Hooker, Sp. fil. 3: 202. 1 860.

TYPE: Ecuador, forest of Esmeraldas, Hall

(holotype, K!, frag., B; photo, us of B). Figures

Ib-d.

Asplenium spruceanum Hieron., Hedwigia 60: 260.

1918. TYPE: Brazil, San Gabriel, Spruce 2357

(not 2375 as in protologue) (holotype, B!, frag,

us!; isotype, GH!).

Plants epiphytic. Stem rather stout, erect or de-

cumbent, provided with linear-lanceolate scales,

these 2-3 mm long, clathrate, dark brown to black-

ish. Leaves 1 -pinnate-pinnatifid or -pinnatisect,

erect, fasciculate, 18-40 cm long, (1.5-)2-9 cm

broad. Petiole 1-6 cm long, lustrous, castaneous

to atropurpureous, glabrous. Lamina thin- to firm-

herbaceous, glabrous, gradually reduced to apex

and base, the tip attenuate, usually flagelliform

and/or proliferous, rachis dark and lustrous. Pin-

nae 20-30 pairs, larger ones (1.5-)2—4 cm long,

patent, or a few proximal ones deflexed, sessile,

pinnatifid, or incised nearly to the costa at base,

the basal acroscopic segment commonly overlap-

ping the rachis. Veins simple in most segments,

forked to pinnate in basal ones. Sori solitary on

each vein. Indusia thin, yellowish or greenish.

In deep forests, on stumps or tree trunks and

branches, 100-200 m, San Martin and Loreto.

Colombia; Ecuador; Peru; Amazonian Brazil.

Some confusion has attended the nomenclature

and descriptions of this species and its synonym.

Hieronymus separated his A. spruceanum from A.

hallii on the basis of several quantitative charac-

ters and cited as its type a paratype of the latter

(incidentally citing the number incorrectly as 2375).

During the present study, comparison of the type

material showed there is no reason to recognize

18

FIELDIANA: BOTANY

two species. Furthermore, in the protologue, fol-

lowing the description of A. hallii, Hooker made

a curious statement: "My specimens from Col.

Hall have the pinnae again pinnated, especially in

the lower half . . . ." The Hall specimen at Kew

exhibits nothing of the kind and, in fact, is iden-

tical to Spruce 2357, except the pinnae are less

deeply incised and the segments a little broader.

Mexia 6184 (F, GH, MO, uc, us) from Loreto has

dull, gray-green rachis and petiole, stem scales light

brown and ovate, and roots somewhat golden-to-

mentose (all characters similar to those of A. au-

riculatum) but otherwise matches A. hallii. This

may be a hybrid involving the two species, for

many sporangia are barren or with some spores

malformed.

San Martin: Lamas, Santa Rosa de Davidcillo, near

road to Tioyacu, Knapp & Mallet 7195 (F, MO, uc). Lo-

reto: Mishuyacu, near Iquitos, Killip & Smith 29975 (F,

GH, us). Prov. Maynas, Mishara, Lopez et al. 8650 (HUT).

Prov. Requena, Jenaro Herrera, N of Requena, Tovar

6968 (USM). La Victoria on the Amazon River, LI. Wil-

liams 2989 (F, us).

14. Asplenium macrurum Mickel & Stolze, in

Stolze, Flora of Ecuador 23: 41. 1986, nom.

nov.

Asplenium longicaudatum Mickel & Stolze, Amer. Fem

J. 74: 115. 1984, nom. illeg., not A. longicauda-

tum Bonaparte. 1917. TYPE: Ecuador, Prov. Mo-

rona-Santiago, Cordillera Cutucu, Camp E-1283

(holotype, NY!; isotype, s).

Plants terrestrial or low-epiphytic. Stem stout,

decumbent to erect, provided with ovate to lan-

ceolate, acute scales, these 2-4 mm long, 0.6-1

mm broad, dark brown or gray-brown, or often

bicolorous with narrow, lighter brown margins,

subclathrate, the lumina isodiametric or slightly

elongated. Leaves 1 -pinnate, fasciculate, to 1 mm

long and 18 cm broad. Petiole 10-30 cm long,

lustrous or sublustrous, dark brown to atropur-

pureous, essentially glabrous. Lamina thin- to firm-

herbaceous, glabrous, lanceolate or deltate-lanceo-

late, not or scarcely reduced at base, gradually

reduced to a long, flagelliform apex. Rachis gla-

brous, not alate, dark reddish brown, the flagel-

liform tip often proliferous but fragile and usually

broken off in dried specimens. Pinnae 10-17 pairs,

to 12 cm long and 1 .8 cm broad, sessile or short-

stalked, equilateral except at base, there broadly

cuneate to truncate acroscopically, narrow-cune-

ate basiscopically, the apex narrowly acute to at-

tenuate, margins subentire to broadly crenulate.

Veins l(-2)-forked. Sori 5-8 mm long, linear or

narrow-oblong, slightly arcuate. Indusia yellow-

ish, firm, subentire.

On the floor of deep, wet forests or at the base

of tree trunks, 1000-1700 m, Amazonas, Ucayali.

Venezuela; Ecuador; Peru.

Heretofore known only by the type collection

from Ecuador, several collections of the species

now have been made in Peru. It is similar in ap-

pearance to A. radicans var. cirrhatum but differs

significantly in stem scales and in size and shape

of pinnae. The Peruvian, especially Schunke, col-

lections closely match the type, but pinnae of the

two Harbour specimens are not attenuate as on the

type; they are merely narrow-acute.

Amazonas: Prov. Bagua, 12 km E of La Peca, Barbour

2404 (MO, uc), 2507 (uc). Ucayali (as Loreto): Prov.

Coronel Portillo, Dist. Padre Abad, Rio Chino, J. Schunke

V. 9197 (MO, uc).

15. Asplenium radicans L., Syst. nat. ed. 10, 2:

1323. 1759. LECTOTYPE (designated by

Morton & Lellinger, Mem. New York Bot.

Gard. 1 5: 38. 1966): Jamaica, Browne 92 (LINN

1250.16, right-hand specimen; photo, us).

Plants terrestrial, rarely epiphytic or epipetric.

Stem stout, erect, provided at apex with linear or

lanceolate, attenuate, clathrate scales, these 2-5

mm long, gray-brown to blackish. Leaves 1 -pin-

nate to 3-pinnate-pinnatisect, fasciculate, to 1 m

long and 25 cm broad. Petiole 10-30 cm long,

lustrous, dark brown to atropurpureous, essen-

tially glabrous. Lamina thin- to firm-herbaceous,

lanceolate to deltate-ovate, not or scarcely reduced

at base, gradually tapering to a radicant apex. Ra-

chis essentially glabrous, bisulcate adaxially, not

or conspicuously green-alate, lustrous, terminat-

ing in a naked and flagelliform apex. Pinnae nu-

merous, extremely variable: simple, entire to cre-

nate-serrate and commonly 2-5 cm long by 1 cm

broad in var. cirrhatum, to 3-pinnate and 12-15

cm long by 4-5 cm broad in var. uniseriale; sessile

or subsessile, patent to slightly ascending. Veins

commonly 1-2-forked, or simple in ultimate seg-

ments of decompound varieties. Sori elongated,

or moderately so in decompound varieties. In-

dusia thin, yellowish or greenish, oblong to elliptic,

the margins subentire.

The species occurs in the West Indies, and from

Mexico to Bolivia and Brazil.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

19

Few fern species are as highly variable in lamina

architecture as Asplenium radicans. A combina-

tion of features distinguish it from other species

in the genus: lustrous, atropurpureous petiole and

rachis; flagelliform, proliferous leaf tips; scarcely

reduced lamina base. However, the lamina may

be simply pinnate, or 3-pinnate, or varying in an

infinite number of degrees between each extreme.

This has prompted some authors to recognize four

or more species, each based solely on very subtle

differences. Obviously, questions posed by this and

similar aggregate species cannot be satisfactorily

resolved by comparison of herbarium specimens,

within the limitations of a floristic treatment. Field

and greenhouse observations coupled with cyto-

logical studies are needed. A good beginning was

made by Walker (Trans. Roy. Soc. Edinburgh 66:

169-237. 1966; and 69: 109-235. 1969), who re-

vealed that some crossing is involved within and

also out of the species complex.

Tentatively, four varieties are recognized, which

may or may not represent geographic or ecological

taxa.

Key to Varieties

a. Lamina 1 -pinnate or 1 -pinnate-pinnatisect. Secondary segments (if any) adnate to a broadly alate

costa b

b. Pinnae lobed to deeply pinnatisect (but not dissected entirely to the costa) . . 1 5a. var. radicans

b. Pinnae entire to deeply crenate-serrate, not lobed, not or rarely auriculate 1 5b. var. cirrhatum

a. Lamina 2- to 3-pinnate, at least the larger secondary segments definitely stalked c

c. Pinnae 1 -pinnate (at least in proximal half), the pinnules coarsely dentate to deeply pinnati-

sect 1 5c. var. partitum

c. Pinnae 1 -pinnate-pinnatisect to 2-pinnate, the pinnules (at least proximal one) with 1 -several free

segments 1 5d. var. uniseriale

1 5a. Asplenium radicans var. radicans.

Asplenium rhizophyllum L., Sp. pi. 1540. 1763, based

on the same protologue as A. radicans. Not A.

rhizophyllum L., 1753.

(For full list of synonyms of A. radicans and varieties,

see Morton and Lellinger, 1966.)

Pinnae pinnatifid to deeply pinnatisect, basal

segments often free, but not fully stalked, obtuse

to subacute, subentire to dentate distally.

Rare, on slopes or ravine banks in wet forests,

600-2300 m, Amazonas, San Martin, Huanuco.

West Indies; southern Mexico south to Brazil

and Bolivia.

Degree of lamina dissection merges gradually

into that of var. partitum, the subtle distinction

here being that the latter has clearly stalked sec-

ondary segments with coarsely dentate apices.

Amazonas: Prov. Bongara, Shillac, D. Smith & Vas-

quez 4839 (MO, uc). San Martin: Mt. Guayrapurima,

near Tarapoto, Spruce 4021 (BR, GH, us). Huanuco (as

San Martin): East of Tingo Maria, Allard 21562 (GH).

15b. Asplenium radicans var. cirrhatum (Willd.)

Rosenst., Hedwigia 46: 102. 1906.

Asplenium cirrhatum Willd., Sp. pi. ed. 4, 5: 321.

1810. TYPE: Guadeloupe, L. C. Richard (holo-

type, B, Herb. Willd. 19894, frag., NY!; isotype, p;

photos, F, GH, us of B).

Pinnae entire to deeply crenate-serrate, acro-

scopically truncate and sometimes subauriculate,

basiscopically narrow-cuneate to excavate.

In forests, on sandy or rocky soil, or on old logs

or bases of tree trunks, 300-1700 m, Amazonas

and Loreto to Cuzco and Puno.

Guadeloupe; Greater Antilles; southern Mexi-

co; Guatemala; Nicaragua to Brazil and Bolivia.

Amazonas: Prov. Bagua, valley of Rio Maranon above

Cascadas de Mayasi, Wurdack 1871 (GH, us). San Mar-

tin: Prov. Mariscal Caceres, Dist. Uchiza, SE of Nuevo

Progreso, Schunke V. 3159 (F, GH). Loreto: Valseca-Ru-

dolpho, along Rio Corrientes, McDaniel & Marcos 11061

(GH, MO). Huanuco: Prov. Leoncio Prado, Castilo Alto,

Plowman 5849 (F, GH, USM). Pasco: Prov. Oxapampa,

Palcazu Valley, between Iscozacin and Villa America,

D. Smith 3871 (MO, uc). Junin: Schunke Hacienda, above

San Ramon, Killip & Smith 24577 (F, GH). Cuzco: Prov.

Paucartambo, Cosnipata Valley, Rio Tono, Wachter et

al. 201 (F, USM). Madre de Dios: Tambopata Nature

Reserve, between the lodge and Rio la Torre, Funk et

al. 8377 (uc, us). Puno: Prov. Carabaya, Palmera, Vargas

16150 (GH).

1 5c. Asplenium radicans var. partitum (Klotzsch)

Hieron., Bot. Jahrb. Syst. 34: 464. 1904.

20

FIELDIANA: BOTANY

Aspleniumflabellulatum var. dentatum Klotzsch, Lin-

naea 9: 71. 1834. SYNTYPES: Columbia [Ven-

ezuela], Moritz 44 (B?, P!; photos, F & us of P).

Venezuela, prope Caracas (B?).

Aspleniumflabellulatum Kunze var. partitum Klotzsch,

Linnaea 20: 357. 1847. LECTOTYPE (designat-

ed by Morton & Lellinger, Mem. New York Bot.

Card. 15: 37. 1966): Venezuela, Caracas, Otto

651 (holotype, B; photo, us).

Asplenium partitum (Klotzsch) C. Chr., Index fil. 1 25.

1905.

Asplenium radicans var. dentatum (Klotzsch) Bonap.,

Notes Pteridologiques 7: 349. 1918.

Pinnae 1 -pinnate, with at least several pairs of

free pinnules, these coarsely dentate and often

deeply incised.

On slopes and ravine banks of wet forests, 650-

2400 m, Amazonas, Huanuco, Pasco, Junin, Cuz-

co.

West Indies; Mexico south to Peru and Bolivia.

Stolze (1986) had not seen type material but

cited A. flabellulatum as a synonym of var. par-

titum, based on the original description. However,

having now examined the lectotype (with fully tri-

pinnate lamina), it can be seen that the former

name is more properly placed under var. unise-

riale. This is typical of the confusion surrounding

the many names and variants of A. radicans, all

of which have been distinguished merely on mi-

nute degrees of variation in lamina dissection.

Amazonas: Prov. Bongara, Shillac, Smith & Vdsquez

4893 (MO). Huanuco: La Divisoria, 25 km NE of Tingo

Maria, Moran 3700 (MO). Pasco: Prov. Oxapampa, Dist.

Oxapampa, Rio San Alberto, Leon 633 (F, GH). Junin:

Chanchamayo Valley, C. Schunke 109 (F), 110 (F), 1452

(F). Above San Ramon, C. Schunke A2 11 (uc). Prov.

Tarma, Agua Dulce, Woytkowski 3 5429 (MO, uc). Cuzco:

Torontoy, Urubamba Valley, Cook & Gilbert 1097 (us).

Prov. La Convention, Rio Apurimac, Davis et al. 1299

(F, GH).

15d. Asplenium radicans var. uniseriale (Raddi)

Gomez, Brenesia 8: 53. 1975.

Asplenium uniseriale Raddi, Opusc. Sci. 3: 29 1 . 1819.

TYPE: Brazil, Raddi (holotype, n).

Asplenium flabellulatum Kunze, Linnaea 9: 71. 1834.

TYPE: Peru, Pampayacu (Huanuco), Poeppig

(holotype, LZ, destroyed; lectotype, B!, frag., us!,

photo, us).

Pinnae 2-3-pinnate, with basal pinnules strong-

ly overlapping the rachis, most pinnules with 1 to

several free segments, these obovate to cuneiform,

or once again pinnate.

Terrestrial in and at edges of forests, 100-2700

m, Cajamarca to Loreto, south to Huancavelica

and Cuzco.

West Indies; southern Mexico to Brazil and Bo-

livia.

With this probably should be included A. bal-

liviani Rosenst. of Bolivia, which Rosenstock dis-

tinguished from A. uniseriale as having slightly less

divided laminae (2-pinnate-pinnatifid). Asple-

nium radicans var. uniseriale (including A. flabel-

lulatum) has laminae 3-pinnate or more. Detailed

analysis of the species complex is needed. Studies

by Stolze in Guatemala and Ecuador, and now in

Peru, have revealed no other morphological char-

acters or ecological data to corroborate differences

in degree of lamina dissection. Monographic study

possibly may substantiate the premise that A. rad-

icans is merely a species with highly varied leaf

architecture, whether this conclusion is "conve-

nient" to classification or not.

Cajamarca: Prov. Cutervo, Grutas de San Andres, Lla-

tas Quiroz & Suarez 2735 (F, uc). Amazonas: Prov. Ba-

gua, Cordillera Colon SE of La Peca, Barbour 4003 (MO).

San Martin: Zepelacio, near Moyobamba, Klug 3489 (F,

GH, MO), 3490 (B). Loreto: Mishuyacu, near Iquitos. Klug

208 (F), 392 (F). Huanuco: Prov. Huanuco, Carpish, As-

plund 12835 (s). Pasco: Prov. Oxapampa, Dist. Oxa-

pampa, Rio San Alberto, Leon 651 (F, GH, USM). Junin:

Chanchamayo Valley, C. Schunke 830 (F), 1399 (F).

Huancavelica: Prov. Tayacaja, Marcavalle, Tovar 4755

(GH). Cuzco: Rio Tacate, Bues 1745 (us).

16. Asplenium repens Hooker, Sec. cent, ferns, /.

31. 1860. TYPE: Ecuador (Napo), growing in

woods in Archedona, Jameson 787 (holotype,

K.!). (Jameson 786 cited in the protologue.)

Plants epiphytic, rarely terrestrial. Stems mi-

nute, borne on long-creeping, stoloniform roots.

Leaves remote, borne singly, or occasionally in

pairs, 2-pinnate (at least proximal ones), 1 .5-5 cm

long. Petiole to 0.5 cm long, yellow- or gray-green,

at the base bearing a few, inconspicuous scales,

these castaneous, clathrate, ovate, about 0.5 mm

long. Lamina membranaceous, oblong or deltate-

oblong, reduced to a pinnatifid apex, not or scarce-

ly reduced at base. Rachis glabrous, lacking pro-

liferous buds. Pinnae 4-8 pairs, approximate or

subdistant, stalked, essentially pinnate, typically

with only a single pinnule borne on the acroscopic

side of the costa (thus nearly bifoliolate). ultimate

segments about 3 mm long, cuneate, flabelliform,

subdigitately incised at apex. Veins commonly

1 -forked in each ultimate segment. Sori solitary

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

21

on each ultimate segment. Indusia thin, broad,

yellowish or greenish brown, entire.

Rarely collected in Peru, on trunks or branches

of trees, rarely on wet ground or mossy rocks,

1700-1800 m, Huanuco, Junin, Cuzco.

Colombia; Ecuador; Peru; Bolivia.

Jameson 786 was cited as the type by Hooker,

and he also referred to it in Species Filicum (3:

194. 1860). However, the specimen in the holo-

type folder is 787. A diligent search by Peter Ed-

wards (K), as well as a canvass of other herbaria,

failed to yield a collection numbered 786. Thus,

it appears likely Hooker erred in citing the type

number.

Although shape of ultimate segments and lam-

ina of A. repens are quite different from those of

A. delicatulum, the two species are very closely

related. Both have small, delicate and glabrous

leaves borne on stoloniform roots, with a few

clathrate petiole scales about 0.5 mm long. That

neither species is well represented in herbaria is

probably because they are so small and incon-

spicuous.

Huanuco: Prov. La Divisoria, 25 km NE of Tingo

Maria, Moran & Fernandez 3704 (MO, uc, USM). Junin:

La Merced, Chanchamayo, Soukup 1009 (F). Cuzco:

Tablahuas, Lucumayo Valley, Biies 1034 (us).

17. Asplenium triphyllum Presl, Reliq. haenk. 1:

45. 1825. TYPE: "Hab. in Cord. Peru" (but

possibly Ecuador), Haenke (holotype, PRC!;

photo, F).

Asplenium tenue Presl, Reliq. haenk. 1 : 44, t. 6, f. 5.

1825. TYPE: Ecuador (as Peru), monte Chim-

borazo, Haenke (holotype, PR!; isotype, PRC!;

probable isotype, NY!; photos, F & GH of PR, GH

of NY).

Asplenium ternatum Presl, Reliq. haenk. 1: 45. 1830.

TYPE: in vallibus cordillerarum Peruviae, Haenke

(PR!).

Asplenium rhomboideum Brack., U.S. expl. exped. 1 6:

156, t. 21, f 2. 1854. TYPE: (Huanuco) Peru,

Andes of Banos, Brackenridge 24 (holotype, us!).

Plants epipetric, rarely terrestrial. Stem small to

stout, provided with linear or linear-lanceolate

scales, these obscurely clathrate, 1-2 mm long,

blackish. Leaves fasciculate to densely caespitose,

(4-)6-30 cm long, 0.5-1.8 cm broad. Petiole 1-10

cm long, dull, or rarely sublustrous, gray-brown

to castaneous, sparsely provided with minute,

glandular trichomes. Lamina thin-herbaceous to

chartaceous, linear, gradually tapering to a pin-

natifid apex, abruptly to gradually reduced at base.

Rachis rigid to flexuous, sparsely to moderately

provided with filiform scales and pluricellular tri-

chomes, not or inconspicuously alate, often bear-

ing a proliferous bud in a pinna axil. Pinnae nu-

merous, subsessile to short-stalked, subdistant to

densely crowded, most of them trifoliolate or bi-

foliolate, or occasionally with another pair of lat-

eral pinnules, ultimate segments obovate or ob-

lanceolate, entire to bifid. Veins simple (rarely

1 -forked) in each ultimate segment. Sori solitary

on the vein of each segment. Indusia thin, pale

yellowish or translucent, subentire.

In crevices and ledges of rock walls, at cave

mouths, among rocks and boulders, rarely in wet,

mossy, soil, (2800-)3 100-4800 m, from Caja-

marca south to Arequipa and from Huanuco south

to Puno.

Colombia to Bolivia and Argentina.

The taxonomy of this species and its near rel-

atives has been very confused; consequently, a

rather lengthy discussion is in order here. The type

collection of A. tenue cited above contains leaves

merely 2—4 cm long and 0.6-0.8 mm broad, and

Stolze (1986) thought they closely resembled those

of A. fragile (= A. peruvianum). Although pinnae

are not conspicuously trifoliolate, close exami-

nation reveals that many of them are deeply cleft;

therefore, they resemble many of the smaller, bi-

foliolate pinnae of A. triphyllum. The stem scales

of the holotype of A. tenue are scarcely or very

narrowly clathrate, like those of A. triphyllum, and

not obviously clathrate like those of A. peruvia-

num. Thus, it seems that A. tenue is nothing more

than a depauperate form of A. triphyllum.

The types of both A. tenue and A. triphyllum

were collected by Haenke, and on the labels are

marked "Peru." However, boundaries of the two

countries were often confused, and the protologue

of the former cited that the plants were actually

collected in Monte Chimborazo, which is in Ec-

uador. Presl based his descriptions of both species

on the Haenke collections in hand, and it is quite

conceivable that Haenke found both at the same

collecting site. Despite the fact that these plants

evidently represent extreme forms of the same spe-

cies, Presl chose to describe them as different.

Asplenium trilobatum C. Chr. of Bolivia is very

closely related and also may be merely a depau-

perate form.

Other variability in leaves of Asplenium tri-

phyllum similarly prompts their recognition as dif-

ferent species. At one extreme, the stem is rather

22

FIELDIANA: BOTANY

small, the rachis slender and flexuous, and the

pinnae thin-herbaceous and subdistant; in other

leaves (as in the type of A. ternatum), the stem is

stout, the rachis stout and rigid, and the pinnae

chartaceous and crowded (even imbricate). When

proliferous buds are present, they tend to be found

only on the thin and flexuous leaves. Nevertheless,

it is doubtful that two distinct taxa are represented

here, as too many intermediate conditions occur,

and the extreme forms may simply indicate leaf

development responding to different habitats. Field

studies are needed to corroborate this.

Besides being closely related to A. peruvianum,

A. triphyllum also is part of a variable species com-

plex involving A. myriophyllum, A. haenkeanum,

and perhaps some others; in fact, several speci-

mens in Peru are intermediate between A. tri-

phyllum and A. haenkeanum. See treatment of the

latter for further discussion.

Cajamarca: Prov. Cajamarca, Cerro "Cumbe Mayo,"

Sanchez V. 39 (GH, HUT, us). La Libertad: Prov. Santiago

de Chuco, Laguna La Victoria, Sagdstegui el al. 6189

(GH, HUT). Ancash: Prov. Huari, Huascaran National

Park, D. Smith el al. 12411 (F, MO). Huanuco: Chasqui,

Macbride & Featherstone 1757 (F, GH, us). Lima: Prov.

Huarochiri, above Chumpicocha, Cerrate 2005 (F, GH,

USM). Junin: Road from Tarma to La Oroya, Correll &

Smith P786 (GH, us), P787 (GH, MO). Huancavelica: Prov.

Tayacaja, Tocas, between Colcabamba and Paucarbam-

ba, Tovar 2103 (GH, USM). Ayacucho: Between Quinua

and road to Tambo, Soukup 6185 (us). Cuzco: Prov.

Cuzco, near Saxihuaman, Tryon & Tryon 5361 (F, GH,

us, USM). Arequipa: Pichu Pichu, Stafford 676. Puno:

Near Puno, Soukup 302 (F, uc).

18. Asplenium divaricatum Kunze, Linnaea 9:71.

1834 (not Wallich, 1829, nom. nud.). TYPE:

Peru, "Cerro San Christobal (sic), prope Cas-

sapi," Poeppig (diar. 7757), 1829 (holotype,

LZ, destroyed; isotypes, BM!, P!, w!; photos, F,

GH, uc & us of BM; F of w).

Plants epipetric, rarely terrestrial. Stem small to

stout, erect, provided with linear or linear-lanceo-

late scales, these obscurely clathrate, 1-2 mm long,

bicolorous, blackish with narrow but conspicuous

brown bands of marginal cells. Leaves caespitose,

2-pinnate-pinnatisect to 3-pinnate, 5-30 cm long,

1-7 cm broad. Petiole 0-6 cm long, dull, reddish

to grayish brown. Lamina thin- to firm-herba-

ceous, linear to elliptic, gradually reduced to apex

and base. Rachis glabrous or with a few filiform

scales, narrow-alate, lacking proliferous buds. Pin-

nae numerous, sessile, crowded to imbricate, basal

pinnules overlapping the rachis. Pinnules of larger

pinnae 6-10 pairs, bifoliolate or with several pairs

of ultimate segments, segments commonly less than

2 mm long, obtuse, fertile ones nearly twice as

broad as adjacent sterile ones. Veins simple in each

ultimate segment. Sori solitary on the vein of each

segment, at maturity usually projecting beyond

segment margin. Indusia thin, translucent, elliptic-

oblong, margins entire.

Commonly in crevices of dry rocky walls or in

soil among rocks, 700-2700 m, generally along the

Cordillera Central from Amazonas south to Cuz-

co.

Endemic to Peru.

Superficially, this resembles A. rutaceum, but

the latter differs, among other characters, in its

darker, lustrous petiole and radiant leaf apex. A

distinctive feature of most leaves of A. divaricatum

is the relative size and shape of fertile and sterile

segments, which is especially evident on pinnules

that are not again pinnate but merely bifoliolate.

The fertile segment is usually obovate, while the

opposing sterile segment is often about half its

width and nearly linear.

Mickel and Beitel (Mem. New York Hot. Gard.

46: 55-56. 1988) included A. divaricatum in their

Pteridophyte Flora of Oaxaca and cited its distri-

bution from Mexico to Peru. They maintained that

this name applies to specimens from Central and

South America that traditionally have been in-

cluded under A. myriophyllum and that true A.

myriophyllum occurs only in the West Indies.

However, the Oaxaca fern they described and il-

lustrated as A. divaricatum and a supposed type

fragment (NY) are substantially different from ac-

tual type material (BM, P, w) of that species and

from specimens of true A. divaricatum in Peru

(cited below). References to neither the bicolorous

stem scales nor the size and differences in fertile

and sterile segments are seen in Mickel's descrip-

tion and illustration.

There is a fragmentary specimen at New York

labeled simply "Asplenium rhizophyllum Kze.,

Peru, Poeppig," which is identified and filed as a

type collection of A. divaricatum. Labels of all oth-

er type specimens (cited above) contain not only

the identification as A. divaricatum but also the

full data of type locality. Asplenium rhizophyllum

is a synonym of A. myriophyllum, and indeed the

"type collection" at New York matches the type

of the latter well. Perhaps it is this specimen that

prompted the incorrect application of the name of

A. divaricatum to the Mexican plants. This is also

typical of the understandable confusion that has

long attended the circumscription of species in this

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

23

highly variable complex. See discussion of A.

haenkeanum for further details.

Amazonas: Prov. Chachapoyas, Cerros Calla Calla,

above Leimebamba, Hutchison & Bennett 4545 (F, GH,

uc, us). Hacienda Limon, W of Balsas, Osgood & An-

derson 57 (F), 63 (F, us). San Martin: Trip from Pacas-

mayo to Moyobamba, Stiibel 1023 (B, us). Huanuco:

Muna, Bryan 432 (F, us). Huacachi, estacion near Muna,

Macbride 3870 (F, GH, us, w). Junin: Prov. Tarma, near

San Felix Bridge, Ferreyra 14955 (GH, USM). Prov. Tar-

ma, SW of Huacapistana, Tryon & Tryon 5433 (GH).

Cuzco: Masias, high above Apurimac River, Sues 1383

(us).

1 9. Asplenium haenkeanum (Presl) Hieron., Hed-

wigia47: 233. 1908.

Athyrium haenkeanum Presl, Tent, pterid. 98. 1836.

TYPE: Peru, near city of Huanuco, Haenke 52

(holotype, PRC!).

Asplenium cladolepton Fee, Mem. foug. 7: 55. 1857.

TYPE: Colombia, Prov. Ocana, Schlim 324 (ho-

lotype, P; isotype, L; frag., us!: photos, F & us of

L).

Asplenium cladolepton var. angustifolium Hieron., loc.

cit. SYNTYPES: Peru, (Loreto) prope Cuelapa

(Loreto), Stiibel 1019b (B); supra Tambo Mayo,

Amazonas, Stiibel 1043 (B).

Asplenium cladolepton var. minor Hieron. loc. cit.

TYPE: Peru (Loreto) prope Cuelapa, Stiibel 101 9a

(B).

Plants epipetric (rarely terrestrial). Stem small

to rather stout, erect or suberect, provided with

linear or deltate-lanceolate scales, these obscurely

clathrate, 1.5-3 mm long, blackish, or occasionally

faintly brown-margined. Leaves fasciculate,

2-pinnate, 12-38 cm long, 2.2-5(-7) cm broad.

Petiole 3-1 2 cm long, dull reddish or grayish brown.

Lamina thin-herbaceous or membranaceous, lin-

ear, gradually reduced to apex and base. Rachis

glabrous or with a few filiform scales, narrow-alate,

lacking proliferous buds. Pinnae 20-28 pairs, ap-

proximate to subdistant, mostly patent, or prox-

imal ones deflexed, sessile, basal pinnules over-

lapping the rachis. Pinnules of larger pinnae 3-6

pairs, shallowly to deeply bifid, fertile and sterile

segments subequal, 2-3 mm long. Veins simple in

each ultimate segment. Sori solitary on veins. In-

dusia thin, yellowish or greenish, margins entire.

In wet places, on or among rocks, or in crevices

of rock walls, 2900-3800 m, Amazonas, La Liber-

tad, Loreto, Huanuco, Pasco, Apurimac, Cuzco.

Colombia; Venezuela; Ecuador; Peru.

This is at the morphological center of a species

complex involving A cristatum, A. myriophyllum,

A. triphyllum, and, in a broader sense, A. divari-

catum. These species are alike in many significant

characters, although they have been separated tra-

ditionally by some variable, quantitative, features:

lamina dissection and shape, degree of reduction

of lamina base, degree of pinna angle, and number

of pinnules.

Typical A. triphyllum has very long and narrow

leaves with bi- or trifoliolate pinnae, in the axils

of which are frequently borne proliferous buds,

but not infrequently there are pinnae with an ad-

ditional one or two lateral segments, whose con-

dition thus approaches that of less robust forms

of A. haenkeanum (3-6 pairs of pinnules). How-

ever, no proliferous buds yet have been found in

the latter species. Conversely, more robust leaves

of A. haenkeanum (narrow-elliptic, with up to 6

pairs of pinnules) resemble narrower forms of A.

myriophyllum (7-10 pairs of pinnules). The latter

is characteristically terrestrial, growing at 1400-

2700 m in Peru, and A. haenkeanum is epipetric,

at 2900 m or higher. Asplenium cristatum and A.

myriophyllum have been distinguished by slight

differences in scale color, pinna shape, and degree

of reduction in the lamina base, which is seen

under/!, cristatum.

Some specimens of A. myriophyllum with un-

usually small ultimate segments might be confused

with A. divaricatum, but the latter commonly can

be distinguished by its conspicuously bicolorous

stem scales and the difference in size and shape of

the fertile and sterile segments. Obviously the en-

tire complex in the Neotropics is badly in need of

detailed study, and it is with strong reservations

that its various components are separated in this

flora.

La Libertad: Prov. Bolivar, Bolivar, Lopez & Sagds-

tegui 3274 (GH, HUT). Huanuco: Chasqui, Macbride &

Featherstone 1756 (F, us, in part). Pasco: Huariaca, Bry-

an 168 (F). Road to Cerro de Pasco, Gentry et al. 37486

(F, MO). Apurimac: Prov. Andahuaylas, Huancarama,

Vargas 8827 (uc). Cuzco: Ollantaytambo, Cook & Gilbert

395 (us). Prov. Urubamba, Chincheros, King et al. 141

(F, a variant).

20. Asplenium cristatum Lam., Encycl. 2: 310.

1786. TYPE: Locality and collector undesig-

nated (holotype, P, Herb. Jussieu 1276; photo,

us).

Asplenium cicutarium Sw., Prodr. 130. 1788. TYPE:

Jamaica, Swartz (holotype, s!; isotype, SET; pho-

tos, F & us of s).

24

FIELDIANA: BOTANY

Plants terrestrial, rarely epiphytic or epipetric.

Stem stout, erect, provided with linear or deltate-

lanceolate scales, these 2-3 mm long, subclathrate,

gray-brown or blackish, occasionally faintly brown-

margined. Leaves fasciculate, 2-pinnate-pinnati-

sect to 3-pinnate, to 60 cm long and 1 5 cm broad.

Petiole 5-25 cm long, dull gray or reddish brown

to blackish. Lamina thin-herbaceous or membra-

naceous, elliptic or oblong-lanceolate, gradually

reduced to a pinnatifid apex, scarcely reduced at

base, but occasionally with 1 or 2 proximal pairs

of pinnae strongly reduced. Rachis glabrous or with

a few, scattered, filiform scales, alate, lacking pro-

liferous buds. Pinnae 1 5-20 pairs, approximate to

crowded or imbricate, most of them patent, ob-

long, the margins parallel for much of their length,

sessile, basal pinnules overlapping the rachis. Pin-

nules of larger pinnae 9-16 pairs, deeply pinnati-

sect or pinnate, ultimate segments 2.5—4 mm long,

narrow-elliptic, acute. Sori 3-8 on a pinnule. In-

dusia thin, grayish or yellowish, margins subentire

or slightly erose.

On wet forest floor, in wooded ravines, rarely

on or among rocks or on bases of tree trunks, 250-

2400 m, Cajamarca and Amazonas south to Aya-

cucho and Madre de Dios.

United States (Florida); West Indies; Mexico to

the Guianas, south to Bolivia and Brazil.

With this perhaps should be included A. myri-

ophyllum, which shares nearly the same range and

habitat and has been distinguished by subtle dif-

ferences such as shape of lamina base, pinnae, and

ultimate segments. See A. haenkeanum for further

discussion.

Cajamarca: Prov. Hualgayoc, Monte Seco, Soukup

381 1 (F, us). Amazonas: Prov. Bagua, Cerro Tapur near

Rio Utcubamba, Hutchison 1480 (F, uc, us). San Martin:

Near Tarapoto, Spruce 3975 (BR). Huanuco: Prov. Hua-

nuco. Tingo Maria, Asplund 13201 (s). Mima, Bryan 424

(F). Pasco: Oxapampa, Soukup 2355 (F, GH). Junin:

Chanchamayo Valley, C. Schunke 7 (F, us), 65 (F, us).

Ayacucho: Aina [Ayna], between Huanta and Rio Apu-

rimac, Killip & Smith 22838 (us). Cuzco: Prov. La Con-

vention, Rio Apurimac below Puerto Capiro, Davis et

al. 1293 (F, GH). Madre de Dios: Maldonado, Lopez et

al. 4588 (GH).

21. Asplenium myriophyllum (Sw.) Presl, Reliq.

haenk. 1:48. 1825.

Caenopteris myriophylla Sw., J. Bot. (Schrader)

1800(2): 50. 1801. TYPE: Jamaica, Swam (ho-

lotype, s!; photo, F).

Plants terrestrial, rarely epipetric or epiphytic.

Stem small to rather stout, erect or suberect, pro-

vided with linear or deltate-lanceolate scales, these

obscurely clathrate, 3-4 mm long, cast an ecus to

gray-brown or blackish, occasionally faintly brown-

margined. Leaves fasciculate, 2-pinnate-pinnati-

sect or 3-pinnate, 12-35(-40) cm long, 3-7 cm

broad. Petiole 3-15 cm long, dull reddish or gray-

ish brown. Lamina thin-herbaceous or membra-

naceous, ovate to elliptic, gradually reduced to a

pinnatifid apex, abruptly reduced at base, with 2-

4(-6) proximal pairs of pinnae much shorter than

central ones. Rachis glabrous or with a few, scat-

tered, filiform scales, alate, lacking proliferous buds.

Pinnae 15-28 pairs, approximate to subdistant,

mostly ascending, broadest near base, tapering

gradually or abruptly to apex, sessile, basal pin-

nules overlapping rachis. Pinnules of larger pinnae

(6-) 7-10 pairs, with one or more free ultimate

segments, these 2-3 mm long, mostly obovate and

obtuse to subacute. Veins simple in each ultimate

segment. Sori solitary on the veins. Indusia thin,

yellowish or greenish, margins entire.

Infrequent and widely scattered in Peru, on moist

forest floor, rarely on rocks, 1400-2700 m, Ca-

jamarca, Amazonas, Huanuco, Pasco, Junin.

United States (Florida); Greater Antilles; Mex-

ico to Peru and Bolivia.

This is the most highly dissected of several spe-

cies in a confusing complex. See further discussion

under A. haenkeanum and A. divaricatum.

Cajamarca: Prov. Santa Cruz, Bosque de Monteseco,

Sagdstegui et al. 12387 (F, HUT, MO, uc). Colasay, Woyt-

kowski 7014 (GH, MO, us). Amazonas: Prov. Bagua, Cor-

dillera Colan SE of La Peca, Barbour 3896 (F, MO). Hua-

nuco: Mima, Woytkowski 5215 (GH, MO). Prov. Leoncio

Prado, km 478 on Lima-Tingo road, Young & Sullivan

850 (F, MO). Pasco: Prov. Oxapampa, 2 km from Oxa-

pampa, Leon 496 (USM). Junin: Huancayo, Soukup (F).

22. Asplenium delicatulum Presl, Reliq. haenk. 1:

47, /. 7,f. 5. 1825. TYPE: Ecuador, "in Cor-

dilleris Quitensibus," Haenke (holotype, PRC!;

isotype, HBG; photos, F, GH, us of HBO).

Plants epiphytic or epipetric. Stems very small,

crowded to subdistant, borne on (usually) long-

creeping, stoloniform roots. Leaves borne singly

or in small fascicles, 2-3-pinnate, 4— 7(-9) cm long.

Petiole 1.5—4 cm long, yellow- or gray-green, with

a few inconspicuous scales at the base, these dark

brown, clathrate, ovate, 0.4-0.6 mm long. Lamina

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

25

membranaceous, deltate-ovate, reduced to a pin-

natifid apex, not or scarcely reduced at base. Ra-

chis glabrous, lacking proliferous buds. Pinnae 5-

8 pairs, approximate or subdistant, stalked, with

2-3 pinnules, these simple or again pinnate, ulti-

mate segments 2—4 mm long, linear or narrow-

oblanceolate, acute. Veins simple in each ultimate

segment. Sori solitary on the veins. Indusia thin-

textured, narrow-elliptic, yellowish or grayish

green, entire.

In wet forests and wooded ravines, on tree trunks,

branches, and fallen logs, and on wet, moss-cov-

ered rocks, 500-1500 m, San Martin, Huanuco,

Pasco, Ucayali.

Colombia; Venezuela; Ecuador; Peru; Chile.

Although more than a dozen collections have

been found in herbaria thus far from Peru (most

of these in Huanuco), only a few specimens have

been seen from Colombia and Venezuela, and ap-

parently the species is represented to date in Ec-

uador only by the type. As with the closely related

A. repens, the paucity of existing collections is like-

ly due to the inconspicuous nature of the plant

rather than to its actual rarity. See A. repens for

further discussion.

San Martin: Prov. Mariscal Caceres, Cerro Verde, Leon

112 (USM). Near Tarapoto, Spruce 4035 (BR, GH, us).

Huanuco: Prov. Huanuco, near confluence of Rio Ca-

yumba with Huallaga, Mexia 8290 (F, GH, uc, us). Prov.

Leoncio Prado, Dist. Rupa Rupa, 5 km S of Tingo Maria,

Schunke V. 3252 (F, GH, us). Prov. Huanuco, Tingo Ma-

ria, Tryon & Tryon 5238 (F, GH, us, USM). Pasco: Prov.

Oxapampa, Valle del Palcazu, Rio Cacazu, Leon 682 (F).

Ucayali: Prov. Coronel Portillo, Sinchono, near La Divi-

soria, Aguilar 834 (GH).

23. Asplenium squamosum L., Sp. pi. 1082. 1753.

TYPE: Pet. fil. \\2,t.5,f. 2, illustration based

on a plant from Hispaniola.

Asplenium jamesonii Hooker, Sp. fil. 3: 184. 1860.

TYPE: Ecuador, Guayaquil, Jameson (holotype,

K; isotype, BM; frag., NY!; photo, us of BM).

Plants terrestrial. Stem stout, creeping or de-

cumbent, provided with broad, flaccid, light brown

scales, these 10-20 mm long, 3-6 mm broad, finely

clathrate, often remotely fimbriate. Leaves sub-

distant, 2-pinnate or 2-pinnate-pinnatifid, 40-100

cm long, 1 5-50 cm broad. Petiole to 40 cm long,

dull, light or grayish brown. Lamina firm-herba-

ceous to chartaceous, subdeltate, gradually re-

duced to a pinnatifid apex, not reduced at base,

most axes and veins sparsely to moderately pro-

vided with minute, brown, appressed, 2-3-celled,

articulate trichomes. Rachis light or gray-brown,

glabrous, with a few, scattered, broad scales, non-

alate, narrowly alate near apex, lacking proliferous

buds. Pinnae 8-16 pairs, approximate, ascending,

clearly stalked, deltate or oblong-deltate. Pinnules

2-9 cm long, ovate-lanceolate to subtrapeziform,

acute or subacute, cuneate at base, crenate-dentate

or deeply lobed. Veins several-forked, borne at

acute angles. Sori borne in a single line along each

side of costule. Indusia thin, often scarious, mar-

gins entire to erose.

In forests, thickets, or wooded ravines, in wet

or rocky soil, 2100-3600(-3900) m, San Martin,

Huanuco, Ayacucho, Apurimac, Cuzco.

Hispaniola; Costa Rica; Colombia and Vene-

zuela to Bolivia and Brazil.

This species is distinguished by its huge leaves

and the very large, nearly trapeziform, ultimate

segments. With it probably should be included A.

tucumanense Hieron. of Argentina.

San Martin: Prov. Mariscal Caceres, Rio Abiseo, Young

4168 (USM). Huanuco: Mito, Bryan 379 (F). Muna, Bryan

627 (F). Tambo de Vaca, Macbride 4464 (F, us). Aya-

cucho: Prov. La Mar, between El Tambo and Ayna,

Plowman & Davis 4681 (GH, USM). Apurimac: Prov.

Abancay, Sanchez 244 (USM). Cuzco: Prov. Paucartam-

bo, Pillahuata, Aldave 5024 (HUT). Calca, Vargas 15645

(GH).

24. Asplenium cuspidatum Lam., Encycl. 2: 310.

1786. TYPE: Peru, Jussieu (holotype, P, Herb.

Jussieu, photo, us; isotype, P, Herb. Lamarck;

photo, us).

Plants epiphytic or terrestrial, rarely epipetric.

Stem stout, erect, provided with lanceolate to ovate,

orange to gray-brown scales, these 2—4 mm long,

0.5-0.8 mm broad, acute, narrowly clathrate.

Leaves caespitose, 2-pinnate (at least as to basal

pinnae) to nearly 4-pinnate, 10-70 cm long, 2-

1 2(-l 5) cm broad. Petiole dull or sublustrous, green

or gray to reddish brown or blackish. Lamina firm-

herbaceous to chartaceous, lanceolate to ovate or

subdeltate, tapered to a pinnatifid or serrate, often

caudate, apex, not reduced at base, commonly gla-

brous. Rachis glabrous, but with a few, minute,

filiform scales, nonalate, or narrowly so distally,

lacking proliferous buds. Pinnae 6 to many pairs,

approximate to subdistant, most of them ascend-

ing, short-stalked, ovate to linear-lanceolate. Pin-

nules 0.5-2 cm long, lanceolate to ovate, acute or

subacute, subentire to nearly 3 -pinnate, at least

26

FIELDIANA: BOTANY

the basal ones of proximal pinnae obviously

stalked. Veins forked to pinnately branched. Sori

elliptic. Indusia relatively thick and fleshy, narrow,

grayish or whitish, subentire.

This and A. auritum are components of a highly

varied species complex. Pending monographic

study they are separated arbitrarily here as a mat-

ter of convenience (see A. auritum for detailed

discussion). In Central America and northern South

America, A. cuspidatum is very common, but south

of Ecuador it is encountered much less frequently.

Two varieties are provisionally recognized in Peru,

and var. tripinnatum is included in the following

key, as it is to be expected here.

The species occurs in the West Indies, from

Mexico to Panama, and from Colombia and Ven-

ezuela south to Bolivia and Brazil.

a. Proximal pinnae 1 -pinnate (at least near their base), the pinnules deeply serrate or a few with basal

acroscopic lobes 24a. var. cuspidatum

a. Proximal pinnae 1-pinnate-pinnatifid to nearly 3-pinnate, most of the pinnules deeply pinnatifid or

more b

b. Proximal pinnae 1-pinnate-pinnatifid to nearly 2-pinnate, the pinnules deeply pinnatifid, some

tertiary segments incised to the costule, but never freely stalked 24b. var. tripinnatum

b. Proximal pinnae 2-pinnate to nearly 3-pinnate, tertiary segments commonly stalked

24c. var. foeniculaceum

24a. Asplenium cuspidatum var. cuspidatum.

In forests, on wet slopes, rarely on mossy rocks

or on tree trunks, 1 500-3 100 m, Cajamarca, Ama-

zonas, La Libertad, San Martin, Cuzco.

Mexico to Panama; Colombia and Venezuela to

Bolivia and Brazil.

Plants with less dissected pinnae merge with,

and are often identified as, A. auritum.

Cajamarca: Prov. Santa Cruz, Bosque de Montesco,

Sagdstegui et al. 12382 (F, HUT, MO, uc). Amazonas:

Prov. Chachapoyas, Cerros Calla Calla, 45 km above

Balsas, Hutchison & Wright 5807 (GH, uc). La Libertad:

Prov. Bolivar, between Unamen and Bolivar, Lopez &

Sagdstegui 3331 (GH). San Martin: Prov. Mariscal Ca-

ceres, Rio Abiseo National Park, Young & Leon 4945

(HUT), 4994 (HUT). Cuzco: Prov. Urubamba, Machu Pic-

chu, above Rio Mandor, Peyton & Peyton 1286 (MO).

Prov. La Convention, Huayopata, 10 km SW of Inca-

tambo, Peyton & King 1428 (MO).

24b. Asplenium cuspidatum var. tripinnatum

(Fourn.) Morton & Lell., Mem. New York

Bot. Gard. 15: 29. 1966.

Asplenium fragransSv/., Prodr. 180. 1788. TYPE: Ja-

maica, Swartz (holotype, s; isotype, UPS; photo,

us of s).

Asplenium auritum Sw. var. tripinnatum Fourn., Mex-

ic. pi. 1: 107. 1872. TYPE: Based onA.fragrans

Sw.

Not yet reported from Peru, but to be expected.

Greater Antilles, except Puerto Rico; Mexico to

Panama; Colombia and Venezuela to Bolivia and

Brazil.

24c. Asplenium cuspidatum var. foeniculaceum

(HBK.) Morton & Lell., Mem. New York Bot.

Gard. 15: 29. 1966.

Asplenium foeniculaceum HBK., Nov. gen. sp. 1:15.

1815. TYPE: "Novae Andalusiae," Humboldt &

Bonpland (holotype, p; isotype, B, Herb. Willd.

19860; photos, us of p, F & GH of B).

Asplenium abrotanoides Presl, Reliq. haenk. 1 : 47, /.

8,f. 2. 1825. TYPE: Peru, Huanuco, Haenke(ho-

lotype, PRC!; frag., NY!).

Asplenium cuspidatum var. abrotanoides ( Presl) Mor-

ton & Lell., Mem. New York Bot. Gard. 15: 29.

1966.

In wet forests and wooded canyons, in humus,

on mossy logs, sometimes in rocky soil or on tree

trunks, 2000-3950 m, Piura, Huanuco, Huanca-

velica, Apurimac, Cuzco, Puno.

Mexico to Panama; Colombia and Venezuela to

Bolivia and Brazil.

Of the two varieties reported from Peru, var.

cuspidatum usually is found (with few exceptions)

in the northwest and var. foeniculaceum in the

central or (mostly) southern portions. With the

absence, thus far, of the intermediate var. tripin-

natum, the two others are easily separated, both

geographically and morphologically.

Piura: Prov. Ayabaca. near Ayabaca, Sagdstegui &

Cabanidas 8700 (MO). Huanuco: Tambo de Vaca. Bryan

647 (F, GH). Macbride 4459 (F, us). Huancavelica: Prov.

Tayacaja, Ampurco, between Salcabamba and Surcu-

bamba, Tovar 3767 (GH). Apurimac: Prov. Abancay, for-

ests of Ampay, Vargas 1066 (GH, us). Cuzco: Prov. Pau-

cartambo. Dist. Marcachea, near Achirani. I'argas 1 1 140

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

27

(F, GH, uc, us). Puno: Prov. Carabaya, between Ayapata

and Kahualluyoc, Vargas 10770 (GH).

25. Asplenium repandulum Kunze, Linnaea 9: 65.

1834. TYPE: Peru, Huanuco, "Pampayaco"

[Pampayacu], Poeppig'm July 1 829 (holotype,

LZ, destroyed; frag., NY!).

Plants epiphytic, at least in Peru. Stem short-

to long-creeping, scales lacking or rare and small

(less than 1 mm long). Leaves 1 -pinnate, subdis-

tant to remote, 20-40 cm long, 6-12 cm broad.

Petiole 3-1 5 cm long, fleshy, dull reddish or green-

ish brown, naked or with a few small scales at

base. Lamina thin-herbaceous or membranaceous,

oblong-lanceolate to ovate, not or only slightly

reduced at base, narrowing abruptly to a subcon-

form apical segment, this often with a broad, basal

lobe. Rachis scarcely to broadly alate, lacking pro-

liferous buds. Pinnae 10-20 pairs, approximate to

subdistant, sessile or short-stalked, apex acute to

attenuate (rarely obtuse), strongly inequilateral at

base, narrowly cuneate or excavate basiscopically,

truncate and often auriculate acroscopically, mar-

gins crenate to serrate. Veins 1-2-forked, ascend-

ing from the costa at 30-40° angles. Sori mostly

medial. Indusia linear, opaque, greenish or brown,

subentire.

In forests, on tree trunks, 1250-2000 m, Ama-

zonas to Junin, Madre de Dios.

Southern Mexico to Panama; Venezuela and

Colombia to Brazil and Bolivia.

With this perhaps should be included A. obtu-

sifolium L. of northern South America and the

West Indies, which supposedly differs from A.,

repandulum in having fewer pinnae with obtuse

apices and a basal lobe incised nearly to the costa.

Although A. repandulum is often epipetric or ter-

restrial in Mexico and Central America, most col-

lectors report it as epiphytic in Peru and Ecuador.

Amazonas: East of La Peca in Serrania de Bagua, Gen-

try et al. 23073 (MO, uc). Huanuco: Fundo Chela, Rio

"Chin" (Chinchao?), Aguilar 911 (USM). Prov. Leoncio

Prado, Dist. Hermilio Vladizan, La Divisoria, Plowman

& Schunke 7363 (F, GH) Pasco: Prov. Oxapampa, Gran

Pajonal, D. Smith 5168 (F, MO). Junin: Prov. Tarma,

Chanchamayo Valley, above La Merced, Hutchison 1 196

(F, GH, uc, us). Chanchamayo Valley, C. Schunke 155

(F, us). Madre de Dios: Cocha Cashu, between Panagua

and Tayakome, Foster et al. 3449 (F).

26. Asplenium vomeriforme Hooker, Sp. fil. 3: 109,

/. 762. 1860. TYPE: Peru, Mathews in 1835

(K!). (Mathews 1851 was cited by Hooker in

the protologue.)

Asplenium mathewsii Moore. Index fil. 145. 1857.

Nom. nud.

Plants epiphytic or terrestrial. Stem decumbent,

or rarely very short-creeping, provided with gold-

en-tomentose roots and gray-brown scales, these

narrowly clathrate, linear-lanceolate, attenuate, 4—

7 mm long, often ciliate toward base. Leaves

crowded, long-petiolate, usually pendent,

1 -pinnate, to 90 cm long and 25 cm broad. Petiole

1 5-40 cm long, dull, dark brown to yellow-brown,

glabrous. Lamina carnose or chartaceous, abruptly

terminating in a subconform apical segment, or

this often with a basal lobe or auricle, not or scarce-

ly reduced at base, rachis nonalate. Pinnae 6-14

pairs, larger ones 9-13 cm long. 2.5-4 cm broad,

patent, short-stalked, most of them narrowly to

broadly deltate, the margins entire or inconspic-

uously crenulate, conspicuously inequilateral at

base, acroscopically truncate to cordate or auric-

ulate, basiscopically rounded to broadly cuneate,

but proximal ones mostly truncate. Veins once or

twice dichotomously forked, indistinct or obscure.

Indusia firm, entire, inrolled at maturity.

In primary and secondary forests, pendent from

tree trunks and branches, or on the forest floor,

1200-1800 m, Pasco and Junin.

Endemic.

This is very closely related to A. salicifolium and

may be merely a very robust, local variant. Prin-

cipal differences are size, shape and margins of

pinnae.

Pasco: Prov. Oxapampa, 2 km from Oxapampa, Leon

et al. 500 (F, uc, us, USM). Prov. Oxapampa, in coffee

plantation near Villa Rica, van der Werffet al. 8278 (F,

MO, uc). Junin: Chanchamayo Valley, C. Schunke 695

(F), 696 (F, us). Prov. Satipo, Gran Pajonal, south of

Chequitavo, D. Smith 5151 (F, MO, uc).

27. Asplenium juglandifolium Lam., Encycl. 2:

307. 1786. TYPE: "Antilles" (probably Puer-

to Rico or Hispaniola), collector undesignated

(holotype, P, Herb. Jussieu 128 3 A & B; pho-

tos, F & us of 1283 A).

Asplenium integerrimum Sprengel, Nova Acta Phys.-

Med. Acad. Caes. Leop.-Carol. Nat. Cur. 10: 231.

1821. TYPE: Puerto Rico, Bertero (holotype, B;

isotypes, L, P; photos, us of B, F, & us of P).

Asplenium falcinellum Maxon, Contr. U.S. Natl. Herb.

28

FIELDIANA: BOTANY

13: 14. 1909. TYPE: Guatemala, Cubilguitz,

Tuerckheim 1910 (holotype, us!).

Plants epiphytic (rarely terrestrial outside Peru).

Stem decumbent (rarely very short-creeping out-

side Peru), provided with medium or dark brown

scales, these narrowly clathrate, mostly filiform,

3-10 mm long, 0. 1-0.3(-0.4) mm broad. Leaves

crowded, long-petiolate, 1 -pinnate (juvenile ones

rarely simple), to 80 cm long and 25 cm broad.

Petiole 10-30 cm long, dull reddish or greenish

brown, with a few linear scales at very base. Lam-

ina firm-herbaceous to chartaceous, abruptly ter-

minating in a conform apical segment, not reduced

at base, scarcely or moderately provided abaxially

with minute (0.1-0.2 mm), appressed, trichomes,

these often gland-tipped, rachis very narrowly alate

(if at all) near the apex. Pinnae (0-)2-12 pairs,

larger ones 7-20 cm long, 1.2-2.6(-4) cm broad,

ascending, short-stalked, apices commonly atten-

uate, subequilateral at the cunate base, margins

entire. Veins 1-2-forked, indistinct. Indusia linear,

firm and rather thick-textured, entire.

On tree trunks in wet forests, 100-1500 m,

Amazonas and Loreto to Cuzco.

Southern Mexico to Panama; Greater Antilles;

Trinidad; Colombia to Surinam, south to Bolivia

and Brazil.

This species is quite variable in number, size,

and shape of pinnae and in length of stem scales,

and these differences have encouraged earlier sep-

aration into several species and varieties. At one

extreme is A. integerrimum of the Greater Antilles,

with 2-5 pairs of pinnae well over 2 cm broad,

their bases quite inequilateral, and truncate to

broadly rounded. Asplenium falcinellum of Cen-

tral America typically has 6-12 pairs of pinnae

1.2-1.8 cm broad, their bases more narrowly and

more equally cuneate. Pinnae in the type specimen

of A. juglandifolium more closely resemble those

of the former species. Length of stem scales fre-

quently corresponds with pinna characters in that

those of A. falcinellum tend to be less than 5 mm

long and those of A. juglandifolium and A. inte-

gerrimum 6-10 mm long. However, throughout

South America, especially in Ecuador and Peru, a

number of specimens with numerous and narrow-

er pinnae have much longer scales, and in other

specimens the converse is true, which renders the

separation of these taxa untenable when examined

throughout the entire range of distribution. Also

included under A. juglandifolium should be A.

kapplerianum Kunze of Surinam and probably A.

neogranatense Fee of Colombia.

Amazonas: Prov. Bagua, Dist. Cenepa, Tillett 672-67

(GH). San Martin: Zepelacio, near Moyobamba, Klug

3430 (F, GH, MO, us). Prov. Mariscal Caceres, Tocache

Nuevo, Schunke V. 6948 (F, GH, MO). Loreto: Rio Ma-

mon near Rio Nanay, Croat 19950 (GH, MO, uc). Rio

Mazan, Salinas, J. Schunke 375 (F, GH, uc, us). Huanuco:

Above Rio Huallaga at Tingo Maria, Croat 21033 (MO,

us). Pasco (as Junin): Puerto Bermudez, Killip & Smith

26679 (us); Prov. Oxapampa, Cordillera San Matias,

Leon 322 (F, USM). Cuzco: Prov. Paucartambo, Callanga,

Woytkowski 392 (GH).

28. Asplenium tuerckheimii Maxon, Contr. U.S.

Natl. Herb. 13: 15. 1909. TYPE: Guatemala,

Baja Verapaz, Panzal, Tuerckheim 11-1677

(us!).

Plants terrestrial. Stem erect or decumbent, pro-

vided at apex with blackish brown, clathrate scales,

these ovate, 1-2.5 mm long, often somewhat

vaulted. Leaves crowded or fasciculate, to 50 cm

long and 1 5 cm broad, long-petiolate, 1 -pinnate.

Petiole to 25 cm long, dull grayish to reddish brown,

glabrous. Lamina firm-herbaceous, abruptly ter-

minating in a conform apical segment (this seg-

ment sometimes obsolete), not reduced at base,

glabrous except for scattered, minute (0. 1-0.2 mm),

appressed, sometimes glandular trichomes, rachis

narrowly alate. Pinnae typically 3-6 pairs, to 10

cm long and 1 .8 cm broad, alternate, at least prox-

imal ones stalked, not or rarely adnate, narrowly

and subequally cuneate at base, acute to attenuate

at apex, margins broadly crenate-serrate. Veins 1-

2-forked, not distinct. Indusia thin, yellowish

brown, entire.

On the floor of deep forests, 600-1 500 m, Junin

and Cuzco.

Southern Mexico; Guatemala; Peru.

The few collections made in Peru constitute a

wide disjunction in range for this species, here-

tofore known only from southern Mexico and

Guatemala. It is sometimes confused with A. ab-

scissum, or with specimens of A. juglandifolium

with narrow pinnae. The latter species has pinna

bases somewhat to strongly inequilateral, and long,

filiform stem scales, whereas pinnae in A. tuerck-

heimii are equilaterally and narrowly cuneate, and

stem scales are minute, ovate, and often somewhat

vaulted. Asplenium abscissum is much more close-

ly related, for it has identical stem scales. The latter

is usually distinguished by its pinnatifid or non-

conform apical segments and by strongly inequi-

lateral pinna bases.

Junin: Chanchamayo Valley, C. Schunke 88 (F, us),

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

29

1023 (F). Cuzco: Prov. La Convencion, Rio Apurimac,

below Puerto Capiro, Davis et al. 1311 (F, GH, uc).

29. Asplenium tricholepis Rosenst., Repert. Spec.

Nov. Regni Veg. 12: 468. 1913. TYPE: Bo-

livia, Yungas, Polo-Polo, near Coroico, Buch-

tien 3330 (holotype, s!; isotype, us!).

Asplenium rusbyanum Domin, Pterid. Dominica 171,

/. 29, f. 2. 1929. TYPE: Bolivia, Yungas, Rusby

383 (holotype, presumably K; iso types, F!, GH!,

us!).

Plants epiphytic. Stem stout, erect or decum-

bent, provided with orange to reddish brown,

clathrate, scales, these lanceolate to ovate, flat, 5-

15 mm long, 1-2 mm broad, margins (especially

at base) often bearing long, tortuous cilia. Leaves

few, crowded, to 60 cm long and 20 cm broad,

long-petiolate, 1 -pinnate. Petiole to 30 cm long,

dull, gray-green to purplish brown, glabrous. Lam-

ina firm-herbaceous to subcarnose, abruptly ter-

minating in a conform apical segment, not reduced

at base, glabrous, rachis not or scarcely alate. Pin-

nae (l-)2-5 pairs, to 25 cm long and 4 cm broad,

alternate, sessile or subsessile, not or rarely adnate,

narrowly and subequilaterally cuneate at base, api-

ces obtuse or acute, occasionally attenuate, mar-

gins entire. Veins l-2(-3)-forked, distinct or in-

distinct. Indusia thick, inrolled at maturity, entire.

On tree trunks in dense, wet forests, 800-2 1 00

m, Amazonas to Cuzco.

Ecuador; Peru; Bolivia.

The long-cilate stem scales may be helpful in

distinguishing this species, but they do occur, per-

haps less frequently, in A. davisii, A. juglandifoli-

um, and A. salicifolium and its relatives. However

the cilia often go unobserved, either because they

are readily broken off or are actually lacking.

Asplenium tricholepis has been confused with A.

oligophyllum Kaulf. (syn. A. escragnollei Fee) of

Venezuela and Brazil, which is very similar; how-

ever, the latter has crenulate pinna margins, and

blackish, filiform scales along the costae abaxially.

Amazonas: Prov. Bagua, E of La Peca, Barbour 2522

(MO), 2576A (MO). Huanuco: Prov. Huanuco, Tulumayo,

near Tingo Maria, Ferreyra 2151 (GH). Pasco (as Junin):

Pichis Trail, Yapas, Killip & Smith 25560A (us). Junin:

Schunke Hacienda above San Ramon, Killip & Smith

24837 (F, us). Cuzco: Prov. La Convencion, NE of Ha-

cienda Luisiana, Dudley 10398 (GH). Prov. La Conven-

cion, Tinkuri, Vargas 5453 (uc).

30. Asplenium davisii Stolze, sp. nov.

Plantae terrestres. Caulis erecta vel decumbens, pa-

leis brunneolis, clathratis, ovato-lanceolatis, 5-

10 mm longis, 1.5 mm latis. Folium 50-100 cm

longum, 16-28 cm latum. Petiolus 25-60 cm lon-

gus, obscurus, flavidus vel purpurascens. Lamina

1-pinnata, segmento apicali conformi, in pagina

abaxiali trichomatibus minutis, adpressis, brun-

neis, glandulosibus. Pinnae 3-7 jugae, 14-24 cm

longae, 3-4 cm latae, lanceolatae, attenuatae, ad

basin cuneatae et subequilaterale, marginibus

crenato-serratis.

Plants terrestrial. Stem stout, erect or decum-

bent, provided with brown or gray-brown, clath-

rate scales, these ovate-lanceolate, flat, 5-10 mm

long, and about 1.5 mm broad, the margins some-

times ciliate. Leaves few, crowded, 50-100 cm long,

16-28 cm broad, long-petiolate, 1 -pinnate. Petiole

25-60 cm long, dull, yellowish to purplish brown,

glabrous. Lamina chartaceous, abruptly terminat-

ing in a conform apical segment, not reduced at

base, amply to moderately provided abaxially with

minute (0. 1-0.2 mm) appressed, dark brown, plu-

ricellular, often gland-tipped trichomes, rachis

nonalate. Pinnae (2-)3-7 pairs, larger ones 14-24

cm long and 3-4 cm broad, alternate, stalked, lan-

ceolate, attenuate, broadly to narrowly cuneate and

subequilateral at base, margins crenulate to cre-

nate-serrate. Veins 1-2-forked, usually rather dis-

tinct. Indusia linear, 1.5-2.5 cm long, greenish or

yellowish, thin, often inrolled at maturity.

TYPE— Peru, Cuzco, Prov. La Convencion, Rio

Apurimac, above Boca del Tigre rapids, Davis et

al. 1292 (holotype, F!; isotypes, GH!, uc!).

Endemic. Terrestrial in wet forests, 100-1600

m, Amazonas and Loreto south to Cuzco and Ma-

dre de Dios.

This species has been most frequently identified

as A. oligophyllum Kaulf. (syn. A. escragnollei Fee)

of northern South America and Brazil, which dif-

fers most significantly in the blackish, filiform,

clathrate scales along the costa. Presence of these

scales indicates a relationship with the simple-

leaved A. serratum and A. angustum. Asplenium

oligophyllum is further distinguished from A. da-

visii by the absence of minute, appressed tri-

chomes on the abaxial laminar surface.

Asplenium davisii is probably more closely re-

lated to A. canelense Rosenst., a rare species from

Colombia and Ecuador (type: Ecuador, Spruce,

Herb. Bonaparte no. 10010, P!). That species dif-

fers by its much more numerous (12-14 pairs)

pinnae and the absence of glandular trichomes.

Another similar species from Ecuador is A. virens

Presl (syn. A.flavidum Sodiro), which differs from

30

FIELDIANA: BOTANY

A. davisii by its smaller (less than 40 cm) leaves,

its much smaller (3-5 mm) stem scales, more deeply

serrate, merely obtuse to acute, pinnae, and the

absence of laminar trichomes.

Amazonas: Prov. Bagua, Cerro Tapur on Rio Utcu-

bamba, Hutchison 1485 (F, uc, us). Loreto: Puerto Ar-

turo, lower Rio Huallaga below Yurimaguas, Killip &

Smith 27773 (us). Huanuco: Prov. Leoncio Prado, Dist.

Hermilio Valdizan, La Divisoria Plowman & Schunke

7377 (F). Junin: Chanchamayo Valley, C. Schunke 5 (F,

us), 75 (F, us), 452 (F, us). Cuzco: Prov. Paucartambo,

Kosnipata-Pilcopata, Vargas 11290 (GH). Madre de Dios:

Prov. Mann. Atalaya, Foster & Wachter 7462 (F, MO).

3 1 . Asplenium laetum Sw., Syn. fil. 79: 271.1 806.

TYPE: "Habitat in India occidentale," Swartz

(holotype, s).

Asplenium virens Desv., Mem. Soc. Linn. Paris 6: 273.

1827, not Presl, 1825. TYPE: "Habitat in Amer-

ica," collector undesignated (holotype, p, Herb.

Desv.; photo, us).

with petioles quite crowded, so on some specimens

it may appear to be decumbent. Besides the dif-

ferences cited in the key, it differs from A. deli-

tescens and A. melanopus in another subtle char-

acter. The typical lamina in A. laetum becomes

very gradually pinnatifid to a nearly caudate apex;

i.e., each pair of distal segments becomes slightly

shorter than the adjacent pair, the lamina finally

terminating in a subcaudate tip. In the other two

species, the distal segments are rather abruptly re-

duced, the lamina commonly terminating in a

nonconform apical segment.

Amazonas: Bagua, 1 2 km S of La Peca, Barbour 2500

(F). San Martin: Juan Jui, Alto Rio Huallaga, Klug 4253

(F, GH, uc, us). Huanuco: Prov. Leoncio Prado, Dist.

Rupa Rupa, 5 km S of Tingo Maria, J. Schunke V. 3261

(F, GH). Prov. Huanuco, Tingo Maria, Tryon & Tryon

5228 (F, GH, us, USM). Pasco: Prov. Oxapampa, Rio El

Tunqui, D. Smith et al. 1710 (F, MO). Junin: La Merced,

Chanchamayo, C. Schunke 20 (A). Cuzco: Prov. La Con-

vention, below Rosalina, Vargas 12293 (GH).

Plants terrestrial. Stem short-creeping (some-

times not obviously so), provided with linear scales,

these clathrate, 1-3 mm long, dark brown or black-

ish. Leaves 1 -pinnate, crowded, to 40 cm long, 4-

8 cm broad. Petiole 6-18 cm long, dull grayish or

greenish to lustrous and reddish brown or atro-

purpureous, not alate, sparsely to amply scaly.

Lamina thin-herbaceous, oblong-lanceolate, grad-

ually diminishing to a pinnatifid, often subcau-

date, apex, not or scarcely reduced at base. Rachis

essentially glabrous, scarcely alate distally, lacking

proliferous buds. Pinnae 1 5-24 pairs, larger ones

2.5-4.5 cm long, subsessile or short-stalked, most

of them strongly inequilateral for nearly half their

length, truncate acroscopically, narrow-cuneate or

excavate basiscopically, obtuse to acute at apex,

margins bicrenate or biserrate acroscopically, sub-

entire basiscopically. Veins 1-2-forked. Sori

spreading at 30—40° from costa, not or rarely di-

plazioid. Indusia thin, pale green or yellowish, lin-

ear, subentire.

On floor of forests, on rocky slopes, or on mossy

rocks or in soil pockets of rocky cliffs, 200-900

(-1600) m, Amazonas south to Cuzco.

Southern Mexico to Panama; West Indies; Co-

lombia to Surinam, south to Argentina and Par-

aguay.

This is one of a few species of Asplenium with

the petiole varying from light to dark in color and

from dull to lustrous, the variation observable on

the same plant. The stem is very short-creeping,

32. Asplenium melanopus Sodiro, Crypt, vase.

Quit. 189. 1893. TYPE: Ecuador, Chimbo-

razo, Pallatanga Valley near Chimbo Bridge,

Sodiro (holotype, QCA?).

Diplazium melanopus (Sodiro) Hieron., Bot. Jahrb.

Syst. 34: 456. 1904.

Plants terrestrial or epipetric. Stem short-creep-

ing, provided with linear to lanceolate scales, these

clathrate, 0.5-1.5 mm long, dark brown. Leaves

1 -pinnate, approximate to crowded, 28-64 cm long,

10-1 8 cm broad. Petiole 1 2-32 cm long, dark red-

dish brown to purplish, dull, or sometimes lus-

trous, nonalate, sparsely scaly as on the stem. Lam-

ina firm-herbaceous to membranaceous, deltate or

ovate-deltate, rather abruptly reduced to a pin-

natifid or nonconform apical segment, not or

scarcely reduced at base. Rachis essentially gla-

brous, not or scarcely alate, lacking proliferous

buds. Pinnae 7-1 1 pairs, larger ones 6-12 cm long

and 1.4-4 cm broad, short-stalked, equilateral ex-

cept at base, truncate acroscopically, abruptly cu-

neate or excavate basiscopically, acute to attenuate

at apex, both margins lobed to pinnatifid, the tip

of each lobe or segment with a few rounded teeth.

Veins distinct, several times forked. Sori borne on

the vein branches, often back-to-back (diplazioid).

Indusia thin, yellow to reddish brown, linear, sub-

entire.

In forests, in soil or among rocks, sometimes on

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

31

rocky outcrops, 250-1700 m, Amazonas, Loreto,

Junin, Madre de Dios.

Colombia; Ecuador; Peru.

This and A. delitescens have been included ear-

lier in Diplaziwn because of the frequent occur-

rence of back-to-back sori. However, each has the

principal diagnostic characters ofAsplenium: ob-

viously clathrate scales, single-rowed cells in spo-

rangia stalks, annular cells of sporangia 20 or more,

spores with folded perispore. The two species dif-

fer from each other chiefly in the degree of pinna

lobing and perhaps should be separated merely at

an infraspecific level.

Asplenium purpurascens Mett. of Ecuador, and

A. melanopus may also be synonymous; however,

the type of the former has not been examined dur-

ing this study. The name of A. purpurascens would

have priority.

Amazonas: Prov. Bagua, E of La Peca, Barbour 2500

(F, a depauperate specimen). Loreto: Right bank of Rio

Santiago, above Pongo de Manseriche (Amazonas), Mexia

6223 (GH, uc, us). Junin: Colonia Perene, Killip & Smith

24924 (F, us). Madre de Dios: Prov. Tambopata, Al-

bergue Cuzco Amazonico, Leon 866 (USM).

33. Asplenium delitescens (Maxon) Gomez, Bre-

nesia 8: 52. 1976.

Diplazium delitescens Maxon, contr. U.S. Natl. Herb.

10: 497. 1908. TYPE: Cuba, Oriente, vicinity of

San Luis, Pollard & Palmer 348 (holotype, us!).

Plants terrestrial, occasionally epipetric or epi-

phytic. Stem short-creeping, provided with linear

or lanceolate, attenuate scales, these clathrate, 1-

2 mm long, dark brown to blackish. Leaves

1 -pinnate, approximate to crowded, 30-54 cm long,

1 2-24 cm broad. Petiole 1 4-25 cm long, dark red-

dish brown to yellowish brown or greenish, dull

to sublustrous, nonalate, sparsely scaly. Lamina

thin- to firm-herbaceous, deltate or oblong-del-

tate, nonalate, sparsely scaly. Lamina thin- to firm-

herbaceous, deltate or oblong-deltate, gradually or

abruptly reduced to a nonconform, sometimes at-

tenuate, apical segment, not or scarcely reduced

at base. Rachis glabrous, or with a few filiform

scales, marginate or inconspicuously alate distally,

lacking proliferous buds. Pinnae 6-10 pairs, larger

ones 6-14 cm long and 1.5-2 cm broad, sessile,

or proximal ones short-stalked, equilateral except

at base, there truncate (and sometimes auriculate)

acroscopically, narrowly cuneate to excavate basi-

scopically, acute to attenuate at apex, margins ser-

rate to deeply crenulate acroscopically, serrate or

subentire basiscopically. Veins distinct or indis-

tinct, 1-2-forked. Sori (some of them) diplazioid.

Indusia yellow to light brown, linear, subentire.

In forests, on tree trunks, on clay banks, or on

the forest floor, occasionally on rocky cliffs, 135-

750 m, Loreto, Huanuco, Madre de Dios.

Southern Mexico to Panama; Cuba; Colombia;

Ecuador; Peru.

This is probably only a less-dissected form of

A. melanopus, under which see further discussion.

Loreto: Yurimaguas, lower Rio Huallaga, Killip & Smith

29078 (us). Huanuco: Tingo Maria, Moran 3676 (MO,

uc, USM). Prov. Pachitea, Dist. Puerto Inca, Bosque Na-

tional de Iparia, /. Schunke V. 2967 (F, GH, us). Madre

de Dios: Prov. Manu, Parque Nacional Manu, Pakitsa

Station, Foster & Baldeon 12806 (F).

34. Asplenium achilleifolium (Mart. & Gal.)

Liebm., Kongel. Danske Vidensk. Selsk. Skr.,

Naturv. Afd. V, 1: 249 (seors. 97). 1849, not

(Lam.)C. Chr. 1905.

Caenopteris achilleifolia Mart. & Gal., Nouv. Mem.

Acad. Roy. Sci. Bruxelles 15: 63. 1842. SYN-

TYPES: Mexico, Veracruz, vicinity of Mirador,

Galeotti 6279 (BR), 6295 (BR).

Athyrium achilleifolium (Mart. & Gal.) Fee (Mem.

foug. 5) Gen. fil.: 186. 1852.

Plants terrestrial. Stem erect or decumbent, am-

ply provided with obscurely clathrate, ovate or

lanceolate scales, these to 5 mm long and 3 mm

broad, obtuse or acute, light to dark brown. Leaves

1 -pinnate-pinnatisect or nearly 2-pinnate, caes-

pitose, 0.3-1.0 m long, 9-30 cm broad. Petiole

somewhat shorter than the lamina, dull greenish

or grayish brown, sparsely scaly, basal scales broad,

the rest filiform. Lamina firm-herbaceous, ovate

or lanceolate, tapering to an attenuate, pinnatifid

apex, not or slightly reduced at base. Rachis broad-

ly alate, lacking proliferous buds, sparsely filiform-

scaly. Pinnae numerous, 5-15 cm long, crowded

to subdistant, subsessile, truncate and narrowly

decurrent at base, cut nearly to costa into 7-16

pairs of segments, these subentire to crenate or

serrate. Veins 1 -several-forked, diverging from the

midrib at a wide angle. Indusia thin, elliptic, yel-

lowish to whitish.

Thus far known in Peru from one collection, in

wet forest, 2000 m, Amazonas.

Southern Mexico to Nicaragua; Ecuador?; Peru.

The collection cited below apparently is a de-

pauperate form of A. achilleifolium. It matches

32

FIELDIANA: BOTANY

typical plants in Central America except that the

leaves and pinnae are significantly shorter. Stolze

( 1 986, p. 76) cited a single collection from Ecuador

as another possible variant of this species. In any

case, this represents a strikingly disjunct distri-

bution.

Amazonas: Prov. Bagua, 20 km E of La Peca, Harbour

2759 (F, MO, uc).

35. Asplenium projectum Kunze, Linnaea 9: 68.

1835. TYPE: Peru, "Pampayaco, in rupibus

calidioribus" (Dept. Huanuco, Pampayacu),

Poeppig (diar. 1156), Jul. 1829 (holotype, B!;

isotypes, B!, p; photos, GH & uc of P).

Plants epipetric. Stem very small, erect, provid-

ed with linear-lanceolate scales, these 0.5-1 mm

long, clathrate, blackish, sublustrous. Leaves fas-

ciculate, to 7 cm long and 0.7 cm broad. Petiole

1-3 mm long, dull greenish or grayish brown, rare-

ly blackish, marginate or narrowly alate. Lamina

thin-herbaceous, linear, gradually tapering to a

pinnatifid or flagelliform apex, somewhat reduced

at base. Rachis dark brown or atropurpureous,

moderately provided with minute, glandular tri-

chomes, narrow-alate, often bearing a proliferous

bud at apex or in a pinna axil. Pinnae 12-18 pairs,

2-3 mm long, subopposite, with short, blackish

stalks, rotund to obovate, or shallowly and broadly

bi- or trilobate, costa lacking. Veins distinct, black,

simple or 2-3-forked in each pinna. Sori 1-2 on

a pinna. Indusia thin, yellow-green, oblong-ellip-

tic.

In rock crevices, and among rocks, about 2500

m, Amazonas, Huanuco.

Endemic.

This rare species is one of the smallest of the

genus in Peru and is closely related to A. peru-

vianum, A. gilliesii, and A. triphyllum. The tiny,

roundish pinnae and the black color of the stalks

that continues onto the veins set it apart from its

affiliates. The Lopez collection is the first for over

a century, but this is not surprising, as the fern is

so tiny and inconspicuous.

The Poeppig collection cited below is incorrectly

labeled as a type at Vienna.

Amazonas: Prov. Chachapoyas, Pomacocha (Leime-

hamha-Balsas). Lopez et al. 4398 (GH). Huanuco: "Cas-

sapi" (Dist. Chinchao, Casapi), Poeppig, Oct. 1 829 (BM,

w).

36. Asplenium lorentzii Hieron., Hot. Jahrb. Syst.

22: 375. 1896. LECTOTYPE (designated

here): Argentina, Prov. Salta, Yacone, at foot

of Nevada del Castillo, Hieronymus & Lo-

rentz 304 (B!, photo, F, isolectotype, B!). SYN-

TYPE: Argentina, Siambon, Sierra de Tucu-

man, Hieronymus & Lorentz 896 (B!).

Plants terrestrial. Stem erect or decumbent, pro-

vided with linear, attenuate scales, these 1-3 mm

long, clathrate, gray-brown to blackish. Leaves fas-

ciculate, 1 -pinnate, 16-30 cm long, 1.7-2.7 cm

broad. Petiole 4-10 cm long, 0.8-1 m thick, red-

dish- or greenish brown, not or narrowly alate,

glabrous, or with glandular trichomes. Lamina

membranceous, linear, gradually tapering to a pin-

natifid apex, somewhat reduced at base, glabrous,

or with scattered, minute, glandular trichomes.

Rachis straight or slightly flexuous, glabrous or

with glandular trichomes, narrow-alate, prolifer-

ous buds rare or lacking. Pinnae numerous, 10—

14 mm long, subopposite or alternate, subsessile,

subdimidiate, most of them rhombic, deeply den-

tate, patent, or proximal ones deflexed, lacking a

distinct midrib. Veins distinct, slightly darker than

the laminar tissue, 1-2 on basiscopic side of pinna,

4-6 on acroscopic side, 1-2-forked, the basal ac-

roscopic ones subflabellately dichotomous. Sori 4-

6 on larger pinnae. Indusia thin, yellowish or

greenish, oblong-elliptic.

Thus far known in Peru by the lone collection

cited below, on the ground, 700 m, Arequipa.

Peru; Bolivia; Argentina.

The sole collection from Peru differs slightly from

the type in that pinnae are patent and moderately

provided with minute, glandular trichomes on the

abaxial surface. In the lectotypes and paratypes

from Argentina laminae are glabrous, and pinnae

are frequently deflexed. If these differences are

substantiated by subsequent collections in Peru, it

may warrant separation at an infraspecific level.

This species is a low-elevation plant related to

A. peruvianum and A. gilliesii. It is a more robust

fern with stouter petiole, longer leaves, larger pin-

nae, and more numerous sori.

Arequipa: Lomas of Atequipa, Coronado 34 (GH, uc,

us).

37. Asplenium gilliesii Hooker, Exot. fl. 3: 208

(overleaf). 1827. TYPE: Argentina, Cerro

Grande Uspallata, Gillies, Jan. 31, 1825 (ho-

lotype, K!; frag., GH!; ISOTYPE, GH!).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

33

Asplenium gillesianum Hooker, Icon. fil. 1 : t. 73. 1 829,

nom. nov. (an illegitimate renaming of A. gilliesii,

and with the same type).

Plants epipetric, rarely terrestrial. Stem small,

erect, provided with linear, attenuate scales, these

1.5-3 mm long, narrow-clathrate, blackish, sub-

lustrous. Leaves fasciculate, 1 -pinnate, 8-20 cm

long, (0.8-)1-1.5 cm broad. Petiole 2-8 cm long,

0.2-0.4 mm thick, stramineous or greenish, mar-

ginate or very narrowly alate. Lamina thin-her-

baceous, linear, gradually tapering to a pinnatifid

apex, somewhat reduced at base, provided abax-

ially with minute, bacilliform glands. Rachis thin

and flexuous, greenish, sparsely provided with

minute, glandular trichomes, scarcely or narrowly

alate, proliferous buds rare or lacking. Pinnae 7-

20 pairs, 4-8 mm long, subopposite or alternate,

sessile or with short, greenish stalks, rhombic or

subflabellate, lacking a costa, veins ending in acute,

broad and deep, cuspidate lobes. Veins commonly

distinct, slightly darker than the lamina tissue,

subflabellately dichotomous. Sori 2-3 on a pinna.

Indusia thin, yellowish or grayish green, oblong-

elliptic.

In rock crevices, or among rocks in dry mead-

ows, 2750-4000 m, Ancash and Huanuco south

to Ayacucho and Puno.

Peru; Bolivia; Argentina.

Very closely related are A. peruvianum and A.

projectum, all three high-elevation ferns with lin-

ear leaves and blackish, linear scales. See A. pe-

ruvianum for further discussion.

Ancash: Prov. Bolognesi, E of Huasta, Cerrata 2173,

2473 (GH, USM). Huanuco: Yanahuanca, Macbride &

Featherstone 999 (F). Lima: Prov. Yauyos, Capia, below

Tupe, Cerrate 1069 (GH, USM). Ayacucho: Prov. Pari-

nacocha, Incuyo, Hutchison 1270 (F, NY, uc). Cuzco:

Prov. Urubamba, road to Ollanta, Vargas 23280 (F, GH).

Puno: Above Lake Titicaca, Mexia 7782 (F, GH, MO, uc,

us).

38. Asplenium peruvianum Desv., Mem. Soc.

Linn. Paris 6: 271. 1827. TYPE: Peru, col-

lector undesignated, perhaps Dombey (holo-

type, P!, Herb. Desv.; photos, GH, us).

Asplenium fragilePres\,Tenl.pterid. 108. 1836. TYPE:

"Montanis Peruviae," Haenke (holotype, PR!;

frag., NY!; isotype, PRO!; photos, F, GH & us of PR,

F of PRC).

Plants epipetric, rarely terrestrial. Stem erect,

provided with lanceolate or linear-lanceolate scales,

these 1-1.5 mm long, clathrate, blackish, sublus-

trous. Leaves densely caespitose, 1 -pinnate, 6-30

cm long, 0.6-1.5(-2) cm broad. Petiole 1-10 cm

long, commonly 0.5-1 mm thick, dull grayish to

reddish brown, scarcely alate, provided with a few

filiform scales and minute, clavate trichomes, or

glabrescent. Lamina firm-herbaceous or charta-

ceous, linear, gradually tapering to a pinnatifid

apex, somewhat to strongly reduced at base. Ra-

chis typically straight, rigid, and reddish brown,

occasionally thin, flexuous and greenish, with in-

dument as on the petiole, narrow-alate to margin-

ate, sometimes with a proliferous bud in a prox-

imal pinna axil. Pinnae 1 2-30 pairs, 4-8(-10) mm

long, subopposite, sessile or subsessile, rhombic,

a distinct costa lacking (or rudimentary at pinna

base), margins entire or shallowly to deeply den-

tate, rarely lobed. Veins distinct to obscure, subfla-

bellately dichotomous. Sori 4-6 on a pinna. In-

dusia yellowish or greenish, narrow-elliptic.

In open forests or thickets, in rock crevices, on

and among rocks, or in rocky soil, 2750-4500 m,

Cajamarca and La Libertad south to Ayacucho

and Puno.

Venezuela and Colombia south to Chile.

The type specimen of Asplenium peruvianum

consists merely of a single leaf, minus the stem,

so the valuable character of stem scales is lacking.

This matches the holotype of A. fragile, which also

lacks stem scales, and the isotype (PRC), which has

both stem and scales. The holotype label of A.

peruvianum cites no data as to collector, number,

or specific locality. There is another specimen at

Paris, collected by Dombey at Tarma (Junin) also

originally determined to be A. peruvianum. The

label of a Morton photo (us) states that ". . . it

may be the original plant from which the fragment

which is the type was taken." This specimen is

complete with a dozen densely caespitose leaves

and small, blackish, clathrate, stem scales and is

identical to the isotype of A. fragile in every way.

It is unfortunate that this familiar name for a com-

mon South American species of Asplenium must

be subsumed under A. peruvianum.

This is part of a species complex that includes

A. gilliesii and A. projectum as well as A. tricho-

manes-dentatum L. of Central America and the

West Indies. They are all small ferns with linear,

1 -pinnate leaves, blackish, linear stem scales, and

laminae with scattered filiform scales and glan-

dular trichomes. Asplenium gilliesii and A. peru-

vianum may be simply variants (witnessed by the

intermediates cited separately below), and A.

34

FIELDIANA: BOTANY

trichomanes-dentatum is a low-elevation form with

smaller sterile leaves in a rosette pattern. (The

name A. trichomanes-dentatum was later changed

by Linnaeus to A. dentatum, and some authors

still disagree on the application of the rules toward

determination of the valid name. For further dis-

cussion of the nomenclature problem, see Stolze

[Fieldiana, Bot., n.s., 6: 92. 1981].) Monographic

revision is needed.

These species have close affinities with A. tri-

phyllum and A. cladolepton, as well as with high-

altitude plants with narrow leaves. The last two

differ principally in their more dissected leaves. In

addition, A. triphyllum has scattered, minute,

glandular trichomes on the petiole.

Cajamarca: Prov. Cajamarca, El Gavilan, road Caja-

marca-Chilete, Ferreyra 3256 (USM). La Libertad: Prov.

Otuzco, Cerro facing Rio Bado. Aguado 6741 (GH). An-

cash: Prov. Recuay, Pativilea-Huaraz Hwy., D. Smith

8306 (MO). Huanuco: Mito, Bryan 207 (F). Lima: Rio

Blanco. Killip & Smith 21707 (F, GH, us). Junin: NE of

Huancayo, between Acopalca and Huari, Duncan et al.

2746 (MO, uc, us). Ayacucho: Between Huanta and Ha-

cienda Pargora, Killip & Smith 23316 (us). Cuzco: Prov.

Cuzco, near Saxihuaman, Tryon & Tryon 5350 (F, GH).

Puno: Valley 10 mi NE of Macusani, Webster 7 (K).

Intermediate between A. gilliesii and A. peru-

vianum:

Cajamarca: Near Hacienda Porcon, N of Cajamarca,

Correll& Smith P875 (GH, us). Lima: Chicla, Ball (GH).

Cuzco: Prov. Anta, below Chinchaypujio, Vargas 17186

(GH).

39. Asplenium quitense Hooker, Sec. cent, ferns,

/. 20. 1860. TYPE: Ecuador, Andes of Quito,

Archidona, Jameson 707 (holotype, K.!; prob-

able isotype, K.!).

Plants epiphytic. Stems minute, compact, bear-

ing leaves singly or in small clusters, several to

many of these commonly connected by long,

creeping, stoloniform roots, scales lacking. Leaves

1-6 to a stem, 1 -pinnate, 3-1 1 cm long, 0.8-1.8

cm broad. Petiole 0.5-3.5 cm long, marginate or

very narrow-alate to base, greenish to yellowish,

glabrous but sometimes with a few dark brown,

subclathrate scales at base, these 0. 1-0.4 mm long

and nearly as broad. Lamina thin- to firm-her-

baceous, glabrous, lanceolate or oblong-lanceo-

late, rather abruptly reduced to a pinnatifid apex

or with a nonconform terminal segment, not or

scarcely reduced at base. Rachis marginate or nar-

row-alate, lacking proliferous buds. Pinnae 5-10

pairs, to 10 mm long and 7 mm broad, stalked,

irregularly obovate or rhombic, strongly inequi-

lateral, the base acroscopically truncate and the

margin with 1-3 shallow or deep, obtuse lobes,

basiscopically narrow-cuneate and the margin en-

tire or with a single lobe toward the apex. Veins

simple or 1 -forked, mostly obscure, commonly 2

(3) on acroscopic side of midrib and only 1 basi-

scopically. Sori usually 1-3 per pinna. Indusia yel-

low to brownish, linear or narrow-oblong.

In dark, wet forests, on trunks or branches or

trees, or on fallen logs, 700-1200 m, Pasco and

Cuzco.

Colombia; Ecuador; Peru.

As in A. escaleroense, the stem of this species

lacks scales and often produces long, stoloniform

roots, which at intervals in turn produce other

stems. This root system may extend for more than

40 cm along a tree trunk or branch. However, the

former species has merely lobed or pinnatifid

leaves, whereas A. quitense has distinctly stalked

and lobed pinnae.

Pasco: Prov. Oxapampa, Palcazu Valley, Rio Cacazu,

Foster & Smith 9427 (F, MO). Prov. Oxapampa, Rio Ca-

cazu, 56 km from Villa Rica, D. Smith & Foster 2663

(F, MO). Cuzco: Altura de Tintininquiato, Bues 1 752 (us).

Prov. Paucartambo, Cosnipata Hacienda Maria. Vargas

10206 (uc). Prov. Paucartambo. Cosnipata Valley, Rio

Tono, Wachter et al. 189 (F).

40. Asplenium pulchellum Raddi, Opusc. Sci. 3:

291. 1819. TYPE: Brazil, Mandiocca, Raddi

(holotype, FI; isotype, BR; photos, BM, F & us

of BR).

Asplenium poloenseRosensl., Repert. Spec. Nov. Reg-

ni Veg. 12: 469. 1913. TYPE: Bolivia, North

Yungas, Polo-Polo, near Coroico, Buchtien 3345

(holotype, s!; isotypes, B!, us!).

Plants terrestrial or epipetric, very rarely epi-

phytic. Stem erect, provided with linear or lan-

ceolate, often attenuate, scales, these 1-3 mm long,

narrow-clathrate, blackish, sublustrous. Leaves

fasciculate, 1 -pinnate, 6-16 cm long, 1.3-3 cm

broad. Petiole 1-2.5 cm long, reddish brown to

greenish, narrow-alate to marginate, provided with

a few, scattered, dark, filiform scales. Lamina

membranceous, lanceolate or linear-lanceolate,

gradually reduced to a pinnatifid, long-acuminate

or subcaudate apex, somewhat reduced at base.

Rachis reddish or greenish brown, with a few, dark,

filiform scales, narrow-alate, proliferous buds

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

35

lacking. Pinnae 7-15 pairs, 8-14 mm long, short-

stalked, somewhat to strongly ascending, subdi-

midiate, conspicuously inequilateral for half their

length or more, acroscopic margin deeply and

acutely cleft, the basiscopic essentially entire, the

apex acute. Veins usually distinct, 3-5 on the ac-

roscopic side of pinnae, 1-3 on the basiscopic, all

simple except forked at the base acroscopically,

all borne along the principal vein running near and

parallel to the basiscopic margin. Sori commonly

3-7 on a pinna. Indusia thin, yellowish or greenish,

oblong-elliptic.

In deep, wet forests, on the ground or on or

among rocks, very rarely at the base of tree trunks,

400-1800 m, San Martin, Huanuco, Junin, Madre

de Dios.

Ecuador; Peru; Bolivia; Brazil.

This has been confused with A. otites (see Com-

ments below), but the latter is a more robust plant

with a stouter, much longer petiole; also pinnae

are larger and are equilateral except toward the

base, and have more veins and serrations. Asple-

nium otites has not been found in Peru, but it is

apparently somewhat common in northern South

America; A. pulchellum does not occur in northern

South America.

The Schunke collection from San Martin (cited

below) is atypical in that pinnae are not strongly

subdimidiate, but it matches A. pulchellum in ev-

ery other way.

San Martin: Prov. Mariscal Caceres, Dist. Tocache

Nuevo, Fundo La Campina, J. Schunke V. 3428 (F, GH,

us). Huanuco: Prov. Huanuco, Dist. Churubamba, Hda.

San Carlos, Mexia 825 la (uc, us), 8256a (us). Junin:

Chanchamayo Valley, C. Schunke 177, 180, 221, 773,

940, 1364 (all F). Madre de Dios: Prov. Mania, Manu

Park, Cocha Cashu uplands, Nunez 5775 (MO).

4 1 . Asplenium discrepans Rosenst., Repert. Spec.

Nov. Regni Veg. 12: 469. 1913. TYPE: Bo-

livia, North Yungas, Polo-Polo, near Coroico,

Buchtien 3341 (holotype, B?; isotypes, GH!,

us!).

Plants epipetric or terrestrial. Stem erect, pro-

vided with linear, attenuate scales, these 2-3 mm

long, narrow-clathrate, blackish, sublustrous.

Leaves fasciculate, 1 -pinnate, 6-16(-20) cm long,

1-3 cm broad. Petiole 1-3.5 cm long, reddish

brown to greenish, narrow-alate, provided with

scattered, dark, filiform scales. Lamina thin-her-

baceous, lanceolate, gradually tapering to an acute,

pinnatifid apex, somewhat reduced at base. Rachis

reddish brown or greenish, provided with scat-

tered, dark, filiform scales, narrow-alate, prolif-

erous buds lacking. Pinnae 10-20 pairs, 6-15 mm

long, sessile or with short, greenish stalks, patent

or slightly ascending, subdimidiate, conspicuously

inequilateral for half their length or more, acro-

scopic margin crenate-serrate, the basiscopic es-

sentially entire, the apex obtuse. Veins 4-6 and

mostly forked on acroscopic side of pinna, l-3(-

4) and unbranched basiscopically, all borne along

a principal rib running near and parallel to the

basiscopic margin. Sori commonly 4-8 on a pinna.

Indusia thin, yellowish or greenish, oblong-ellip-

tic.

In forests, on rocky hillsides and ravine banks,

or on and among rocks, 700-1400 m, San Martin,

Junin, Cuzco.

Peru; Bolivia; Paraguay.

This is frequently identified as A. pulchellum.

Besides the characters of the key, it also differs

from the latter in its usually patent or slightly as-

cending pinnae. Pinnae of A. pulchellum typically

are borne on the rachis at a very acute angle, al-

though occasionally they may be slightly ascend-

ing.

San Martin: Tarapoto, Spruce 3966 (BR, GH, us). Ju-

nin: San Francisco de Satipo, along Rio Satipo, Solomon

3233 (MO). Yaupi, Woytkowski 6501 (MO, us), 6504 (MO,

us). Cuzco: Santa Ana, Cook & Gilbert 1535 (us), 6501

(MO, us), 6504 (MO, us). Santa Rosa, Urubamba Valley,

Cook & Gilbert 1726 (us). Prov. La Convencion, Potrero,

Tryon & Tryon 5381 (GH).

42. Asplenium pteropus Kaulf, Enum. fil. 170.

1824. TYPE: Brazil, Chamisso (holotype, LE).

Plants epiphytic, sometimes terrestrial. Stem

erect or decumbent, provided with narrowly clath-

rate, linear or lanceolate, acute to attenuate scales,

these 3-5 mm long, gray-brown or castaneous.

Leaves 1 -pinnate, fasciculate, 1 5^4 5 (-60) cm long,

2.5-6 cm broad. Petiole 1-5 cm long, dull grayish

to reddish brown, with a few, scattered, linear to

filiform scales, broadly alate distally or through-

out. Lamina glabrous, thin- to firm-membrana-

ceous, narrow-elliptic, gradually and conspicu-

ously reduced at both ends, terminating in a

pinnatifid, occasionally proliferous, apex. Rachis

dull reddish or greenish brown, broadly alate, each

wing often as broad as the rachis, naked, or with

scattered filiform scales. Pinnae 1.5-3 cm long,

numerous, approximate to subdistant, sessile, acute

to obtuse, conspicuously inequilateral at base, ba-

36

FIELDIANA: BOTANY

siscopically cuneate to excavate, acroscopically

truncate and often subauriculate, the margins

deeply serrate to serrulate. Veins distinct, 5-10

pairs, simple, except forked at or near the pinna

base acroscopically. Sori commonly 7-1 2 on larger

pinnae. Indusia yellowish to hyaline, narrowly el-

liptic.

Rare in Peru, in deep forests, on tree trunks or

on the forest floor, 400-1950 m, San Martin,

Huanuco, Junin, Ayacucho.

Southern Mexico to Panama; West Indies; Co-

lombia to Guyana, south to Bolivia and Brazil.

The taxonomy of this species has been confused,

treated by various authors in a strict to a broad

sense, and it is in need of much closer scrutiny.

Broadly delimited, it includes rather thin-textured

plants with short or medium-length petioles, ta-

pered gradually to a pinnatifid apex and with a

somewhat or strongly reduced base, the rachis and

petiole broadly alate, pinnae conspicuously in-

equilateral at base, with veins mostly simple. In a

strict sense, the species would be confined to those

plants with petioles only a few centimeters in length

and with laminae strongly and gradually reduced

to auriclelike basal pinnae. Plants previously de-

termined as A. pteropus Hieron., with somewhat

longer petioles and with five or six proximal pin-

nae somewhat, but not greatly, reduced are prob-

ably other species: in Central America perhaps A.

barbaense Hieron., in South America perhaps A.

raddianum or (in Ecuador) A. rosenstockianum

Brade. The first two have narrow rachis wings, the

last strongly alate wings as in true A. pteropus. For

purposes of this flora, A. pteropus is delimited in

a strict sense and, as such, is not very common in

Peru.

San Martin: Prov. Mariscal Caceres, Dist. Tocache

Nuevo, Quebrada de Saule Chico, J. Schunke V. 4358

(F, us). Huanuco (as San Martin): Tingo Maria, across

the Huallaga on trail to "Monson" (Monzon), Allard

21864 (us). Junin: Prov. Chanchamayo, Rio Rondayacu,

45 km from San Ramon, D. Smith et al. 2614 (F, GH).

La Merced, Chanchamayo, Soukup 1067, in part (F).

Ayacucho: Rio Apurimac Valley, near Kimpitiriki, Killip

& Smith 22918 (F, us).

43. Asplenium sessilifolium Desv., Ges. Naturf.

Freunde Berlin Mag. Neuesten Entdeck. 5:

322. 1811. TYPE: "Habitat in America ae-

quinoctale," collector undesignated (holo-

type, P, Herb. Jussieu 1265; isotype, B; photos,

BM, GH, us of P).

Plants terrestrial, at least in Peru. Stem stout,

erect or decumbent, provided with clathrate, lin-

ear or lanceolate, attenuate scales, these 2-5 mm

long, gray-brown to blackish, often with narrow,

lighter brown margins. Leaves 1 -pinnate, caespi-

tose, 20-40 cm long, 2-5 cm broad. Petiole 3-16

cm long, gray-brown to atropurpureous or black-

ish, dull or slightly lustrous, naked or with a few

lanceolate scales, not or scarcely alate. Lamina gla-

brous, thin- to firm-herbaceous, linear or narrow-

elliptic, gradually and strongly reduced at both

ends, terminating in a pinnatifid (rarely prolifer-

ous) apex. Rachis castaneous to atropurpureous

(occasionally grayish brown), narrow-alate to mar-

ginate, naked or with a few dark, filiform scales.

Pinnae 1-3 cm long, numerous, approximate to

crowded, sessile, conspicuously inequilateral at

base, basiscopically cuneate to excavate, acroscop-

ically truncate and incised nearly to costa to form

a nearly free basal auricle (occasionally opposed

by a smaller basiscopic auricle), the rest of the

pinna serrate, crenate, or shallowly lobed. Veins

distinct, 4-7 pairs, simple, or subflabellate in the

auricles. Sori 8-12 on larger pinnae. Indusia thin,

linear or elliptic, yellowish or hyaline.

On rocky slopes and wet ravine banks of cloud

forests, 2500-3600 m, Cajamarca, Junin, Ayacu-

cho, Apurimac, Cuzco.

Southern Mexico to Costa Rica; Greater Antil-

les; Ecuador; Peru; Bolivia.

Pinnae of Asplenium triphyllum are typically bi-

or trifoliolate, but in occasional plants they may

vary in having extra lobes, crenations, or serra-

tions, and such specimens sometimes have been

identified as A. sessilifolium. However, the two

differ significantly in indument. In the latter, the

blackish stem scales are 2-5 mm long and often

have narrow, brown margins, and petiole and ra-

chis are naked or with a few, scattered, lanceolate

or filiform scales. In A. triphyllum, the blackish

stem scales are usually less than 2 mm long and

lack lighter colored margins, and axes are sparsely

to moderately provided with filiform scales and

glandular trichomes.

Cajamarca: Prov. Celendin, canyon of Rio Maranon

above Balsas, Hutchison & Wright 5295 (uc). Junin:

Huari, Kunkel, 1960 (B). Ayacucho: Prov. La Mar, above

Jano, Plowman & Davis 4677 (GH). Between La Quinua

and Tambo, Soukup 5625 (us). Apurimac: Forests of

Ampay, Vargas 1069 (GH, MO). Cuzco: Prov. La Con-

vention, Dist. Vilcabamba, between Rumichurco and

Alcobamba, Davis et al. 1235 (F, GH). Prov. La Conven-

tion, Choquequiran, Vargas 12999 (F).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

37

44. Asplenium tabinense Hieron. Hedwigia 60:

224. 1918. TYPE: Peru, near Tabina (Puno),

Lechler 2105 (holotype, B!; photo, F).

Plants terrestrial, at least in Peru. Stem erect or

decumbent, provided with narrowly clathrate, lan-

ceolate or linear-lanceolate, attenuate scales, these

2.5-4 mm long, blackish, with hairlike tip and

scattered marginal cilia (the latter processes usu-

ally broken away in dried plants). Leaves 1 -pin-

nate, fasciculate, 15-33 cm long, 2-4 cm broad.

Petiole 1-5 cm long, atropurpureous, dull or slightly

lustrous, with scattered, dark, filiform scales, nar-

row-alate or marginate. Lamina glabrous, mem-

branaceous, narrow-elliptic, gradually and con-

spicuously reduced at both ends, terminating in a

pinnatifid, usually attenuate, apex. Rachis atro-

purpureous, narrow-alate, naked or with scattered,

dark, filiform scales. Pinnae 1.2-2 cm long, nu-

merous, approximate to crowded, subsessile, con-

spicuously inequilateral at base, basiscopically cu-

neate to excavate, acroscopically truncate and often

auriculate, the auricle sometimes incised nearly

halfway to midrib, the rest of the pinna narrowly

and deeply crenate. Veins (3-)4-5 and simple on

the basiscopic side of larger pinnae, 5-7 on the

acroscopic side, simple, except subflabellate in the

basal auricle and forked in the adjacent lobe. Sori

commonly 4-7 on larger pinnae. Indusia thin, nar-

row-elliptic, yellowish or hyaline.

On densely wooded slopes or in thickets, 1500-

3100 m, Huancavelica, Ayacucho, Cuzco, Puno.

Peru; Bolivia.

All specimens seen from Bolivia are epiphytes,

whereas plants in Peru grow on the forest floor.

Besides the characters cited in the key, this also

differs from A. sessilifolium in the more delicate

lamina, usually firm-herbaceous in the latter, and

membranaceous in A. tabinense. The rachis in A.

sessilifolium varies from atropurpureous to cas-

taneous (rarely lighter colored), but it is always

atropurpureous in A. tabinense, at least on mature

leaves. This perhaps might be considered merely

a geographic variant of the widespread A. sessili-

folium.

Huancavelica: Prov. Tayacaja, Ampurco, between Sal-

cabamba and Surcubamba, Tovar 3768 (GH). Ayacucho:

Ccarrapa, between Huanta and Rio Apurimac, Killip &

Smith 22385 (us). Cuzco: Quillabamba, Santa Teresa,

0.5 km W of La Playa, Peyton & Peyton 1236 (MO). Prov.

La Convencion, Huayopata, Pistipata drainage, Peyton

& King 1429 (MO). Prov. Calca, near Mantio, Vargas

15594 (GH).

45. Asplenium herzogii Rosenst., Meded. Rijks.

Herb. 19: 12. 1913. TYPE: Bolivia, Incacor-

ral, Herzog 2211 (holotype, B?; isotype, us!;

photo, F of us).

Plants terrestrial. Stem erect, provided with ob-

scurely clathrate, lanceolate scales, these 2-3 mm

long, gray-brown, acuminate. Leaves 1 -pinnate,

fasciculate, 25-50 cm long, 3-5.5 cm broad. Pet-

iole 6-12 cm long, gray-brown, narrowly to ves-

tigially alate. Lamina linear, gradually reduced to

a subcaudate, proliferous apex, slightly reduced at

base, with a few proximal pairs of pinnae about

half the length of the longest ones. Rachis dull red-

or gray-brown, narrow-alate. Pinnae 2-2.8 cm long,

1 6-25 pairs, apex obtuse or subacute, inequilateral

at base, basiscopically cuneate, acroscopically

truncate and subauriculate, the margins crenate-

serrate. Veins 5-8 pairs, simple, except forked near

the pinna base acroscopically. Sori 9-15 on larger

pinnae. Indusia thin, hyaline or yellow-brown,

narrow-elliptic, entire.

Thus far known in Peru from one collection, in

thicket, about 2750 m, Huanuco.

Peru; Bolivia.

This may be confused with A. alatum because

of the subcaudate, proliferous rachis but is easily

separated from the latter by the narrow rachis wings

and by the strongly inequilateral pinna bases. In

A. alatum the wings on the rachis (and usually the

petiole) are very broad (1-2 mm), and pinnae are

mostly subequilateral. Asplenium herzogii is most

closely related to A. raddianum, from which it can

be distinguished by the proliferous rachis, the

smaller (2-3 mm vs. 4-6 mm) stem scales and the

linear lamina.

Huanuco: Chaglla, Macbride 3639 (F).

46. Asplenium bangii Hieron., Hedwigia 60: 245.

1918. TYPE: Bolivia, Bang 2419 (holotype,

B; isotypes, F!, NY!, us!).

Plants terrestrial, rarely epiphytic or epipetric.

Stem erect, provided with obscurely clathrate, lin-

ear or linear-lanceolate, attenuate, scales, these 1.5-

3 mm long, blackish, sublustrous, their margins

subentire. Leaves 1 -pinnate, caespitose, 30-65 cm

long, 3.5-5 cm broad. Petiole (5-)6-15 cm long,

dull purplish to reddish brown, narrowly to ves-

tigially alate. Lamina linear or linear-lanceolate,

gradually reduced to a pinnatifid apex scarcely

reduced at base or several proximal pairs about

38

FIELDIANA: BOTANY

half the length of the longest ones. Rachis dull

purplish or reddish brown, narrow-alate (rarely the

wing rather broad), lacking proliferous buds. Pin-

nae (larger ones) 1.2-2.2(-2.5) cm long, (30-)35-

45 pairs, usually obtuse at apex, conspicuously

inequilateral at base, basiscopically cuneate to ex-

cavate, acroscopically truncate and often subauric-

ulate, the margins subentire to crenulate. Veins 8-

12 on acroscopic side of costa, 5-8 basiscopically,

simple, except forked at or near the pinna base

acroscopically. Sori 12-18 on larger pinnae. In-

dusia thin, yellowish or hyaline, margins entire.

In damp forests, on wooded slopes and canyon

walls, commonly on the forest floor, rarely on cliffs

or tree trunks, 650-2150 m, Huanuco, Pasco, Ju-

nin.

Peru; Bolivia; Brazil.

With this perhaps should be included A. claus-

senii Hieron. of Brazil, northern South America,

and the Greater Antilles, of which Hieronymus

recognized three forms, based on minor differ-

ences in length, margin and apex of pinnae. Mor-

ton and Lellinger (1966, p. 16) designated Fendler

138, Venezuela (B; photo, us), the lectotype for

this name, but unfortunately this specimen was

unvailable for examination during our study. In

the Field Museum herbarium, there is a sheet of

Fendler 138, containing laminae of three different

species. The center one is possibly A. claussenii

and is very similar to A. bangii, but it lacks petiole

and stem for definitive comparison. Given the ap-

parent variability of A. claussenii, the entire com-

plex (including A. drepanophyllum and A. pul-

chellum) should be subjected to closer scrutiny.

Therefore, A. claussenii is excluded from Peru for

purposes of this treatment.

Huanuco: Mima, Bryan 433 (F, GH). Mima, Macbride

4010 (F, GH, us). Pasco: Prov. Oxapampa, Valle del Pal-

cazu, Rio Cacazu, Leon 678 (F, USM). Junin: La Merced,

Macbride 5372 (F, GH, us). Chanchamayo Valley, C.

Schunke 1371 (F in part, us). Prov. Tarma, Huacapis-

tana, Tryon & Tryon 5453 (F, GH, uc, us, USM).

47. Asplenium harpeodes Kunze, Linnaea 18: 329.

1844. LECTOTYPE (designated by Morton

& Lellinger, Mem. New York Bot. Gard. 15:

1 5. 1966): Mexico, Leibold26 (B!; photos, us).

Plants terrestrial or epiphytic. Stem erect or de-

cumbent, provided with clathrate, linear scales,

these 4-10 mm long, deep brown or blackish, lus-

trous, terminating in a long, tortuous, unicostate

tip. Leaves 1 -pinnate, caespitose, 30-120 cm long

4-12 cm broad. Petiole 5-10 cm long, dull yel-

lowish to reddish brown or purple-gray, wings

lacking or vestigial. Lamina linear or linear-lan-

ceolate, gradually reduced to a pinnatifid apex,

scarcely reduced at base, or several proximal pairs

about half the length of longest ones. Rachis atro-

purpureous, narrow-alate, lacking proliferous buds.

Pinnae (larger ones) 2.5-9 cm long, (24-)30-50

pairs, long-attenuate, inequilateral at base, basi-

scopically excavate, acroscopically truncate, often

subauriculate (but the basal lobe never incised more

than halfway to the costa), the margins deeply,

serrately lobed. Veins 1 2-20 pairs, mostly simple,

except subdichotomously forked in the acroscopic

auricle. Sori numerous. Indusia thin, yellowish,

brownish or hyaline, margins subentire.

Common, on the earth in wet forests and thick-

ets, in humus, on wet banks, or rocky soil or on

mossy rocks, often on tree trunks, 1 500-3600 m,

Cajamarca to San Martin, south to Huancavelica

and Puno.

Southern Mexico to Panama; Greater Antilles;

Colombia and Venezuela south to Bolivia and

Brazil.

In addition to the characters cited in the key,

this usually can be recognized by the narrowly and

deeply lobed pinnae that very gradually taper to

long-attenuate tips. However, several depauperate

specimens have been seen that have merely strong-

ly serrate margins and acute tips. In these, the

purple rachis and the stem scales with long, tor-

tuous tips are diagnostic. This is one of the most

common species of the genus in the Neotropics.

Cajamarca: Colosay, Woytkowski 7010 (GH, MO, us).

Amazonas: Prov. Chachapoyas, W of Molinopampa,

Wurdack 1471 (GH, us, USM). San Martin: Prov. Maris-

cal Caceres, Rio Abiseo National Park, Young 4 171 (USM),

Young & Leon 501 1 (USM). Huanuco: Cani, 7 mi NE of

Mito, Macbride 3390 (F, GH, us). Pasco: Prov. Oxapam-

pa, 2-4 km N of Mallampampa, D. Smith & Canne 5865

(MO). Junin: Chanchamayo Valley, above La Merced,

Hutchison 1197 (F, NY, uc, us). Huancavelica: Prov. Tay-

acaja. Chuspi, near Tocas, Tovar 2063 (USM). Ayacucho:

Prov. La Mar, between El Tambo and Ayna, Plowman

& Davis 4675 (GH), 4680 (GH). Apurimac: Forests of

Ampay, Vargas 1065 (GH)Cuzco: Prov. Urubamba, Cuz-

co-Quillabamba Road, Skog & Skog 5173 (us). Puno:

Prov. Carabaya, between Ayapata and K'ahualluyoc,

Vargas 10763 (GH).

48. Asplenium raddianum Gaud., in Freycinet,

Voy. Uranie, Bot., 316. 1828. Norn. nov. for

A. brasiliense Raddi and based on the same

type.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

39

Asplenium brasiliense Raddi, PI. bras. nov. gen. 1 : 36,

t. 51,f. 1. 1825, not Swartz, 1817. TYPE: Brazil,

Prov. Rio de Janeiro, Raddi (holotype, n; isotype,

BR; photo, BM of BR).

Plants terrestrial, occasionally epiphytic. Stem

erect, but often slender and elongate, provided with

obscurely clathrate, ovate- to linear-lanceolate

scales, these 4-6 mm long, gray-brown, acute to

attenuate. Leaves 1 -pinnate, fasciculate, 25-50 cm

long, 4-7 cm broad. Petiole 6-10 cm long, reddish

or grayish brown, wings narrow or vestigial. Lam-

ina lanceolate or elliptic-lanceolate, gradually re-

duced to a pinnatifid apex, scarcely reduced at

base, or several proximal pairs about half the length

of the longest ones. Rachis dull reddish or grayish

brown, narrow-alate, lacking proliferous buds.

Pinnae (larger ones) 2.5-3.5 cm long, (12-) 15-24

pairs, apex acute to obtuse, inequilateral at base,

basiscopically excavate, acroscopically truncate,

often subauriculate (but the basal lobe never in-

cised more than halfway to the costa), the margins

deeply serrate or serrately lobed. Veins usually 8-

1 1 on acroscopic side of costa, 6-9 basiscopically,

simple, except forked at or near the pinna base

acroscopically. Sori 10-16 on larger pinnae, fre-

quently diplazioid in the acroscopic auricle. In-

dusia yellowish to light brown, margins subentire.

On wet slopes or in litter on the forest floor,

occasionally on trunks of trees or tree ferns, 1000-

2800 m, Cajamarca and Amazonas south to Cuzco

and Puno.

General distribution uncertain; probably Ven-

ezuela and Colombia; Brazil; Peru; possibly Bo-

livia.

There is some question as to proper application

of this name for the plants so circumscribed here.

The type was not seen, but the specimens cited

below closely match the description and most her-

barium specimens identified as A. raddianum. As-

plenium barbaense Hieron., from Costa Rica, may

belong here, as the two species differ only quan-

titatively and even share the distinctive feature of

having relatively frequent diplazioid sori in the

basal acroscopic pinna auricle. Otherwise, A. bar-

baense essentially differs only in having narrower,

more delicate laminae, proximal pinnae with short

stalks, broader petiole and rachis wings. Perhaps

plants from Costa Rica and northern South Amer-

ica represent an infraspecific variant of those from

Peru, Bolivia, and Brazil. Further study is re-

quired.

Cajamarca: Colasay, Woytkowski 701 1 (MO, us). Ama-

zonas: Prov. Bongara, Shillac, N of Pedro Ruiz, D. Smith

& Vdsquez 4885 (MO, uc). San Martin: San Roque, Li

Williams 7189 (us). Ancash: Prov. Huaraz, Dist. Hua-

raz, behind Hotel Monterrey, Sounders 1282, atypical

(F, GH). Pasco (as Junin): Pichis Trail, "Yapas" [Yapaz],

Killip & Smith 25520 (us). Junin: Schunke Hacienda

above San Ramon, C. Schunke A208 (GH, us). Chan-

chamayo Valley, C. Schunke 108 (us). Cuzco: Urubam-

ba, Machu Picchu, Peyton & Peyton 372 (MO). Puno:

Prov. Carabaya, San Gaban, Vargas 18913, atypical (GH).

49. Asplenium alatum Willd., Sp. pi. ed. 4, 5: 319.

1810. TYPE: Venezuela, Nova Andalusia,

near Caripe, Humboldt & Bonpland (holo-

type, B, Herb. Willd. 19889; isotype, p; pho-

tos, F & GH of B).

Plants terrestrial, rarely epipetric. Stem erect,

provided with obscurely clathrate, ovate or ovate-

lanceolate scales, these 2-3 mm long, gray-brown

acute. Leaves 1 -pinnate, crowded to subfascicu-

late, 20-50 cm long, 7-11 cm broad. Petiole 6-15

cm long, dull greenish or grayish brown, conspic-

uously alate, often to base. Lamina broadly lan-

ceolate to oblong, rather abruptly terminating in

a short-ligulate, proliferous tip, not or scarcely re-

duced at base. Rachis dull reddish or greenish

brown, broadly alate, the wings each 1-2 mm

broad. Pinnae to 7 cm long and 1.5 cm broad, 12-

20 pairs, obtuse to acute, broadly cuneate to trun-

cate or scarcely auriculate at base, subequilateral,

the margins broadly serrate or biserrate. Veins dis-

tinct, mostly 1 -forked. Sori 9-12 pairs on larger

pinnae. Indusia thin, linear, yellowish or greenish,

margins entire.

On the floor of wet forests or banks of wooded

ravines, rarely on wet, mossy rocks, (700-) 1 200-

2400 m, along the Cordilleras Central and Ori-

ental, Cajamarca and Amazonas south to Cuzco.

Southern Mexico to Panama; Jamaica; Hispan-

iola; Colombia and Venezuela south to Bolivia

and Brazil.

The very broad rachis wing and ligulate, prolif-

erous rachis tip distinguish this from most other

species of Asplenium in Peru.

Cajamarca: Prov. Cutervo, Grutas de San Andres, Lla-

tas Quiroz & Suarez 2734 (F, uc). Amazonas: Prov. Bon-

gara, Quebrada Chachuaico, near Shilla, Young et al.

398 (MO, uc). San Martin: Between Moyobamba and

Chachapoyas, Croat 58166 (F, MO). Prov. Mariscal Ca-

ceres, Rio Abiseo National Park, Young 4221 (HUT, USM).

Huanuco: Mima, Macbride 4032 (F, us). Pasco: Pozuzo,

Hacienda Ballisteros, Bryan 692 (F, us). Junin: Chan-

chamayo Valley, C. Schunke 57 (F, us), 727 (F, us). Cuz-

co: Prov. Paucartambo, Kosnipata Valley, Plowman &

Davis 4936 (GH).

40

FIELDIANA: BOTANY

50. Asplenium abscissum Willd., Sp. pi. ed. 4, 5:

321. 1810. LECTOTYPE (designated by

Morton & Lellinger, Mem. New York. Hot.

Gard. 15: 26. 1966): Venezuela, Caracas, Bre-

demeyer (*, Herb. Willd. 19893-1, 19893-2;

photos, F & GH).

Plants terrestrial, occasionally epipetric, rarely

epiphytic. Stem erect or decumbent, provided with

obscurely clathrate, linear or lanceolate scales, these

1-2.5 mm long, blackish or with very narrow brown

margins. Leaves 1 -pinnate, fasciculate, 30-50 cm

long, 5-15 cm broad. Petiole 15-25 cm long, yel-

lowish to reddish brown, not or scarcely alate.

Lamina chartaceous, ovate to oblong, terminating

abruptly in a broad-based, nonconform apical seg-

ment, not reduced at base. Rachis dull, yellowish

or reddish brown, narrow-alate, lacking prolifer-

ous buds. Pinnae (larger ones) 1.5-12 cm long,

0.7-2 cm broad, 4-12 pairs, apex obtuse to atten-

uate, scarcely to strongly inequilateral at base, the

margins subentire to crenate-serrate. Veins few to

many, 1-2-forked. Sori linear or narrow-elliptic.

Indusia thin, yellow to brownish, scarious, often

hyaline until mature, margins subentire.

The species occurs in the United States (Flori-

da); southern Mexico to Peru; West Indies; Co-

lombia and Venezuela south to Bolivia and Uru-

guay.

Throughout most of its broad range the features

of this species are rather constant. However, in

Bolivia and southern Peru a number of specimens

have been found with elongated and more equi-

lateral pinnae. Also, throughout much of Peru and

occasionally growing alongside typical A. abscis-

sum, there occur plants with short, broad, and

obtuse pinnae, and a few of these, including a syn-

type of A. ruizianum, have been reported from

Venezuela. Since these differences are merely

quantitative, three taxa are recognized here as geo-

graphical variants of A. abscissum.

Another species, A. congestum C. Chr. of Ec-

uador, is very closely related but should be con-

sidered distinct, as its lamina is considerably

smaller and membranaceous and tapers gradually

to a narrow, pinnatifid apex. This contrasts with

the chartaceous lamina of A. abscissum, which

abruptly ends in a broad-based, nonconform apex.

Key to Varieties

a. Pinnae subequilateral at base, broadly cuneate acroscopically, narrowly so basiscopically, narrowly

acute to attenuate at apex, commonly 6-10 times as long as broad; veins of larger pinnae 12-20 pairs

50b. var. subaequilaterale

a. Pinnae conspicuously inequilateral at base, truncate acroscopically, cuneate to excavate basiscopically,

obtuse to narrowly acute at apex, commonly 2-4.5 times as long as broad; veins 4-10(-12) pairs

b

b. Pinnae 3-4.5(-6) times as long as broad, subacute to narrowly acute at apex

50a. var. abscissum

b. Pinnae 1.8-2.3 times as long as broad, broadly rounded at apex 50. var. ruizianum

50a. Asplenium abscissum var. abscissum.

In wet forests or wooded canyons, on slopes or

on the forest floor, occasionally on rocks or rocky

cliffs, rarely on bases of tree trunks or fallen logs,

300-2200 m, Cajamarca and Amazonas south to

Ayacucho and Madre de Dios.

Range the same as the species.

Cajamarca: Prov. Santa Cruz, 3.2 km ENE of Mon-

teseco, Santisteban & Guevara 23 (F, GH, HUT, uc). Ama-

zonas: Prov. Bagua, Cerro Tapur, near Rio Utcubamba,

Hutchison 1473 in part (F, uc). San Martin: Juan Jui,

Alto Rio Huallaga, Klug4312 (F, GH, MO, uc, us). Huanu-

co: Cliffs W of Rio Huallaga, above bridge, Croat 57967 'a

(MO in part, USM). Pasco: Cueva Grande, near Pozuzo,

Macbride 4791 (F, us). Junin: Hacienda Perene, Coro-

nado 247 (GH, uc). Ayacucho: Ayna, between Huanta

and Rio Apurimac, Killip & Smith 22835 (F, us). Cuzco:

Calca, Vargas 15595 (GH). Madre de Dios: Parque Na-

cional del Manu, Cocha Cashu Biological Station, M.

Foster P-84-50 (uc).

50b. Asplenium abscissum var. subaequilaterale

Rosenst., Repert. Spec. Nov. Regni Veg. 12:

470. 1913. TYPE: Bolivia, southern Yungas,

Polo-Polo, near Coroico, Buchtien 3349 (ho-

lotype, presumably B).

On the ground in dark, wet forests, 800-1600

m, Junin and Ayacucho.

Peru; Bolivia.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

41

This variety is very similar to A. tuerckheimii

in many aspects, but it is primarily distinguished

from the latter by the nonconform apical segment

and the 7-12 pairs of pinnae. The lamina of A.

tuerckheimii terminates in a conform apical seg-

ment and bears only 3-6 pairs of pinnae.

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

Smith 23900 (us). Chanchamayo Valley, C. Schunke 74

(F, us), 722 (us), 935 (F). Ayacucho: Ayna, between Huanta

and Rio Apurimac, Killip & Smith 22782 (F, us).

50c. Asplenium abscissum var. ruizianum

(Klotzsch) Stolze, comb, and stat. nov. As-

plenium ruizianum Klotzsch, Linnaea 20: 354.

1847. LECTOTYPE (designated by Morton

& Lellinger, Mem. New York Bot. Card. 15:

27. 1966): Andes of Peru, Ruiz (B!; fragment,

NY!; photos, F & us).

In wet forests and wooded canyons, in the earth

or on or among wet rocks, very rarely on tree

trunks, 800-2700 m, Tumbes to Cuzco.

Venezuela; Peru.

Tumbes: Prov. Zarumilla, Dist. Matapalo, Bosque Na-

cional de Tumbes, Schunke V. 2408 (F). Cajamarca: Co-

lasay, Woytkowski 7012 (MO, us). Amazonas: Prov. Ba-

gua, 5 km above La Peca, Croat 58353. Prov. Bagua,

Rio Utcubamba, Cerro Tapur, Hutchison 1473 (F in part,

uc in part, us). Huanuco: Muna, Bryan 434 (F, GH, us).

Junin: Chanchamayo Valley, C. Schunke 74 (F, us). Cuz-

co: Prov. La Convencion, Quebrada Vilcabamba, Vargas

23397 (USM).

51. Asplenium drepanophyllum Kunze, Linnaea

9: 66. 1834, not Baker, 1874. TYPE: Peru,

Pampayaco (Dept. Huanuco, Pampayacu),

Poeppig July, 1829 (holotype, w; frag., BM!;

photOS, BM Of W, F & GH of BM).

Plants terrestrial, rarely epipetric. Stem erect or

decumbent, provided with narrowly clathrate, lin-

ear-lanceolate, attenuate, scales, these 2-3 mm

long, dark brown to blackish. Leaves 1 -pinnate,

fasciculate, 15-40 cm long, 3-4.5(-5) cm broad.

Petiole 4-9(-12) cm long, yellowish to reddish

brown, not or scarcely alate. Lamina membrana-

ceous to firm-herbaceous, linear-lanceolate, grad-

ually reduced to a narrow, pinnatifid, usually acu-

minate apex, scarcely reduced at base or several

proximal pairs about half the length of the longest

ones. Rachis dull reddish brown, narrow-alate,

lacking proliferous buds. Pinnae (larger ones) 1 .2-

3 cm long, 0.4-0.8 cm broad, 15—40 pairs, apex

obtuse to attenuate, conspicuously inequilateral at

base, basiscopically cuneate to excavate, acroscop-

ically truncate and often subauriculate, the mar-

gins crenulate to deeply biserrate. Veins 6-12 on

acroscopic side of costa, (4-)5-10 basiscopically,

1-2-forked, except simple toward pinna base bas-

iscopically. Sori 6-12 on larger pinnae, occasion-

ally diplazioid. Indusia thin, yellowish or brown-

ish, scarious, often hyaline until mature, margins

entire.

In dense, wet forests, on the forest floor, rarely

on mossy rocks, 100-1600 m, San Martin and

Loreto south to Ayacucho and Madre de Dios.

Peru; Bolivia; Paraguay.

Pinna apices vary greatly from obtuse to acu-

minate, and extremes might be presumed different

species, yet these differ from each other in no other

way, and there are an infinite number of inter-

mediate conditions. Leaves with obtuse pinnae are

easily confused with A. bangii (under which see

further discussion), but veins of the latter are all

simple except at the acroscopic pinna base. As-

plenium drepanophyllum is also similar to the

enigmatic A. otites, discussed under Comments at

the end of the genus treatment.

San Martin: Along Rio Huallaga, NE of Shapaja on

road to Chazuta, Knapp & Mallet 6915 (F, MO). Loreto:

Puerto Arturo, lower Huallaga below Yurimaguas, Killip

& Smith 27794 (us). Junin: E of Quimiri Bridge near La

Merced, Killip & Smith 23909 (F, us). Between La Mer-

ced and Satipo, Leon 188 (F, USM). Ayacucho: Estrella,

between Huanta and Rio Apurimac, Killip & Smith 23078

(us). Cuzco: Prov. La Convencion, Rio Apurimac, be-

tween Puerto Capiro and Boca de Tigre rapids, Davis el

al. 1297 (F, GH). Prov. La Convencion, edges of Rio

Mapitunuari, Dudley 10128 (GH, MO). Madre de Dios:

Prov. Manu, Cocha Cashu uplands, Nunez 5766, in part

(MO), 6092.4 (MO).

52. Asplenium auritum Sw., J. Bot. (Schrader)

1800(2): 52. 1801. TYPE: Jamaica, Swartz

(s). (For detailed list of synonyms, see Morton

&Lell., 1966, pp. 17-20.)

Plants epiphytic or terrestrial, occasionally epi-

petric. Stem erect, provided with clathrate, lan-

ceolate to ovate scales, these 2—4 mm long, acute

or acuminate, dark or grayish brown, eciliate.

Leaves 1 -pinnate or 1 -pinnate-pinnatisect, rarely

nearly 2-pinnate at base of proximal pinnae, fas-

ciculate, 10-50 cm long. Petiole 5-20 cm long, dull

green, gray, reddish brown, or blackish, not or

scarcely alate. Lamina chartaceous to subcoria-

ceous, deltate to oblong-lanceolate, very gradually

tapering to a pinnatifid to serrate, often subcaudate

42

FIELDIANA: BOTANY

apex, not reduced at base. Rachis green, gray, or

reddish brown, scarcely alate, or sometimes broadly

so distally, lacking proliferous buds. Pinnae 1-9

cm long, usually numerous, approximate to sub-

distant, varying from obtuse and subentire and

scarcely auriculate at base to deeply pinnatifid,

subcaudate at apex, and with a nearly free basal

acroscopic auricle. Veins indistinct or obscure,

forked, or pinnately branched in larger segments.

Son elliptic, frequently confluent at maturity. In-

dusia firm and fleshy, opaque, commonly pale yel-

low or whitish, subentire.

In forests and thickets, on tree trunks and

branches, on the forest floor, or occasionally on

rocky soil or cliffs, 150-2900 m, Cajamarca to

Loreto, south to Ayacucho and Puno.

Southern United States (Florida); West Indies;

southern Mexico to Bolivia and Argentina.

This and A. cuspidatum are part of a highly vari-

able and widely distributed complex that is char-

acterized within the genus by the fleshy, light green

lamina that contrasts with the rich brown sori (the

latter often confluent at maturity) and by the thick,

fleshy, opaque indusia. Because of the variable

dissection of the lamina ( 1 -pinnate to 3-pinnate-

pinnatifid) and the obtuse to subcaudate pinnae,

authors have separated the species complex into

as many as a dozen taxa. However, individual

specimens essentially differ only in pinna shape

and degree of lamina dissection; thus, for purposes

of this treatment, only A. auritum and A. cuspi-

datum (including three varieties) are recognized

here. These are separated merely as a matter of

convenience, by purely quantitative characters, as

seen in the following key, and there are a number

of intermediates even in these two components of

the group.

a. Pinnae simple and entire to pinnatifid (rarely nearly pinnate at base of proximal pinnae; basal segments

(if any) adnate, never truly stalked A. auritum

a. Pinnae (at least proximal ones) 1 -pinnate to nearly 3 -pinnate; basal segments (pinnules) of proximal

pinnae obviously free and stalked A. cuspidatum

Cajamarca: San Ignacio, San Juan de Piedra Bola, Diaz

2028 (F, MO). Amazonas: Kayamas Creek flowing into

Rio Cenepa, Berlin 463 (MO, us, USM). San Martin: Tara-

poto, Spruce 3956 (BR), 4677 (us). Loreto: Rio Mazan,

J. Schunke 299 (F, GH, uc, us, USM). Huanuco: Prov.

Huanuco, Dist. Churubamba, Hacienda Mercedes, Mex-

ia 8179 (F, GH, uc). Pasco: Prov. Oxapampa, Dist. Ox-

apampa, Rio Alberto, Leon 628 (USM). Junin: Prov. Tar-

ma, Utcuyacu, Woytkowski 35363 (MO, uc). Ucayali:

Prov. Coronel Portillo, Bosque von Humboldt, Young

& Salazar 1026 (F, MO). Ayacucho: Ccarrapa, between

Huanta and Rio Apurimac, Killip & Smith 22369 (us),

22477 (us). Cuzco: Prov. Paucartambo, road from Pilla-

wata to Patria, Plowman & Davis 4941 (F, GH, USM).

Madre de Dios: Prov. Manu. Rio Manu, Cocha Cashu

Station, Foster 9703 (F, MO, USM). Puno: Prov. Carabaya,

San Gaban, Vargas 18922 (GH).

53. Asplenium salicifolium L., Sp. pi. 1080. 1753.

LECTOTYPE (designated by Morton & Lei-

linger, Mem. New York Bot. Gard. 15: 23.

1966): Plumier, Traite foug. Amer. 18, t. 60.

1 705, based on a plant from the West Indies.

Plants terrestrial (often epiphytic outside Peru),

the roots densely golden-tomentose. Stem erect,

provided with clathrate, lanceolate to ovate-lan-

ceolate scales, these 2-6 mm long, acute to atten-

uate, gray-brown, usually with long, orange cilia.

Leaves 1 -pinnate, subfasciculate, 30-80 cm long,

9-20 cm broad. Petiole 8-30 cm long, dull yel-

lowish to reddish brown, not alate, sparsely scaly

and often golden-tomentose at base. Lamina thin-

to firm-herbaceous, lanceolate to deltate-ovate,

abruptly reduced at apex to a broad-based, non-

conform or subconform segment, not or scarcely

reduced at base. Rachis dull yellowish or reddish

brown, narrow-alate, lacking proliferous buds.

Pinnae 1 2-20 pairs, larger ones 4-10 cm long, 1 .2-

2.4 cm broad, subdistant to remote, narrowly acute

to attenuate, inequilateral at base, commonly cor-

date and auriculate, the auricles often overlapping

the rachis acroscopically, narrow-cuneate basi-

scopically, the margins obtusely crenulate to cre-

nate or bicrenate. Veins distinct to obscure,

2-forked, at least in the proximal half of pinnae.

Sori linear or narrow-elliptic, inframedial. Indusia

firm and fleshy, opaque, yellowish to light brown.

Dry woods or cloud forests, on the forest floor

(but often on tree trunks and branches outside

Peru), 2150-2700 m, Amazonas and Huanuco.

Southern Mexico; Nicaragua; Costa Rica; West

Indies; Colombia to the Guianas, south to Peru

and Brazil.

This, A. auriculatum, and A. hastatum have been

combined by some authors. Their differences are

merely quantitative, but they are maintained sep-

arately here pending revision of the group. Morton

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

43

and Lellinger (1966) and Stolze (1986) partially

based their separation of the species on width of

rachis wing, texture of lamina, or acroscopic pinna

bases overlapping the rachis. None of these char-

acters are consistent or significant, whereas length

of stem scales, forking of veins, and shape of pinna

and lamina apex are far more reliable, over the

broad range of distribution.

Amazonas: Bongara, 4 km N of Pomacochas, Knapp

et al. 7523 (F, MO). Prov. Chachapoyas, slopes of Puma-

urcu SE of Chachapoyas, Wurdack 557 (F, GH, uc, us,

USM). Huanuco: Muna, Bryan 528 (F). Cushi, trail to

Tambo de Vaca, Bryan 618 (F). Huacachi, estacion near

Muna, Macbride 4107 (F, us).

54. Asplenium hastatum Kunze, Linnaea 23: 235,

305. 1850. LECTOTYPE (designated by

Morton & Lellinger, Mem. New York Bot.

Gard. 15: 21. 1966): Hort. Lips., originally

from Venezuela (B!, photos F, us; isolectotype,

Plants epiphytic or terrestrial, the roots densely

golden-tomentose. Stem erect, provided with

clathrate, linear-lanceolate, attenuate scales, these

commonly 6-9 mm long, attenuate, gray-brown,

usually with long, orange cilia. Leaves 1 -pinnate,

30-80 cm long, 8-16 cm broad. Petiole 12-30 cm

long, dull greenish to reddish brown, not or slightly

alate, sparsely scaly and often golden-tomentose

at base. Lamina firm-herbaceous to chartaceous,

lance-ovate to oblong, abruptly reduced to a nar-

row, serrate-ligulate or subcaudate apex, not or

scarcely reduced at base. Rachis dull reddish or

greenish brown, alate, lacking proliferous buds.

Pinnae 12-25 pairs, larger ones 6-9 cm long, 1-2

cm broad, subdistant to remote, commonly atten-

uate, inequilateral at base, truncate or subauricu-

late acroscopically, cuneate basiscopically, the

margins obtusely but prominently serrate or bi-

serrate. Veins indistinct or obscure, simple to once-

forked. Sori linear or narrow-elliptic, medial to

inframedial. Indusia firm to subcartilaginous, yel-

lowish to brown.

Thus far known in Peru from one collection, on

trees in mountains, San Martin.

Colombia; Venezuela; Ecuador; Peru.

The only collection seen from Peru perfectly

matches the type specimens. The species seems to

be intermediate between A. auriculatum and A.

salicifolium, and all three could be synonymous.

San Martin: Cerros de Campana, Spruce 4676 (us).

55. Asplenium auriculatum Sw., Kongl. Vetensk.

Acad. Handl. 1817: 64. 1817, not (Thunb.)

Kuhn, 1868. TYPE: Brazil, Minas Gerais,

Freyreis (holotype, presumably s; isotype, BM;

photos, F & us of BM).

Plants epiphytic, sometimes terrestrial, rarely

epipetric, the roots densely golden-tomentose. Stem

erect, provided with clathrate, ovate to lanceolate

scales, these 2-4(-5) mm long, acute, orange to

light brown, usually with sparse, long, orange cilia.

Leaves 1 -pinnate, 12-30(-50) cm long, 3.5-9 cm

broad. Petiole 4-1 5(-l 8) cm long, dull green, gray,

or reddish brown, rarely atropurpureous, non-

alate, sparsely scaly and often golden-tomentose

at base. Lamina thin- to firm-herbaceous, lanceo-

late to oblong, abruptly reduced to a narrow, ser-

rate-ligulate or subcaudate apex, not or scarcely

reduced at base. Rachis dull reddish to greenish

brown, marginate or narrow-alate, lacking prolif-

erous buds. Pinnae 9-15 pairs, larger ones 3-6 cm

long, 1-1.8 cm broad, approximate to subdistant,

acute or subacute, inequilateral at base, truncate

to subcordate and often overlapping the rachis ac-

roscopically, cuneate to excavate basiscopically,

the margins serrate or crenate- serrate. Veins sim-

ple or 1 -forked, indistinct to obscure. Sori infra-

medial, linear or elliptic. Indusia firm, fleshy,

opaque, yellow, green, or light brown.

In deep forests, commonly on tree trunks or

branches, sometimes on wooded slopes or the for-

est floor, rarely on mossy rocks, 500-2700 m, Ca-

jamarca and Amazonas to Cuzco and Madre de

Dios.

Mexico to Panama; West Indies; Colombia to

Trinidad, south to Bolivia and Brazil.

This is closely related to, and possibly synon-

ymous with, A. salicifolium and A. hastatum. See

treatment of the former for further discussion.

Cajamarca: Prov. Santa Cruz, ENE of Monteseco,

Santisteban & Guevara 22 (F, GH, HUT, uc). Amazonas:

Prov. Bagua, E of La Peca, Barbour 2492 (F, MO, uc).

San Martin: Prov. Mariscal Caceres, Dist. Tocache Nue-

vo, 4 km from Puerto Pizana, Schunke V. 4866 (F, us).

Huanuco: Tingo Maria, Stork & Hort on 9469 (F, GH, uc,

us). Pasco: Prov. Oxapampa, Dist. Oxapampa, Rio San

Alberto, Leon 629 (F). Junin: Chanchamayo Valley, C.

Schunke 178 (F), 1369(p). Cuzco: Quillabamba-Potrero,

Coronado 1 19(GH, uc, us). Madre de Dios: Prov. Manu,

Atalaya, near Hacienda Amazonia, Foster & Wachter

7414 (F, MO).

56. Asplenium cuneatum Lam., Encycl. 2: 309.

1786. TYPE: Locality and collector undesig-

44

FIELDIANA: BOTANY

nated (holotype, P, Herb. Jussieu 1 725; photo,

us).

Plants epiphytic, rarely terrestrial, outside Peru

sometimes epipetric. Stem stout, repent, provided

with linear, attenuate, dark brown scales, these 2-

4 mm long, 0.3 mm broad, lustrous, broadly and

conspicuously clathrate. Leaves fasciculate, 2-3-

pinnate, 1 5-35(— 45) cm long, 4-1 2(-l 5) cm broad.

Petiole 7-20 cm long, dull or sublustrous, reddish

brown. Lamina chartaceous, subdeltate- to ob-

long-lanceolate, reduced to a pinnatifid apex, not

or scarcely reduced at base, glabrous. Rachis dark

reddish brown, glabrous, but this and the costae

sparsely to moderately provided with dark, tor-

tuous, filiform scales, essentially nonalate, lacking

proliferous buds. Pinnae 12-25 pairs, approxi-

mate or subdistant, somewhat ascending, short-

stalked, broad- or narrow-deltate. Pinnules 0.6-2

cm long, irregularly ovate to obovate, obtuse, lobed

or with 1-2 pairs of segments, these obovate to

subflabellate, broadly and obtusely dentate toward

the apex. Veins indistinct, dichotomously

branched, commonly forked. Sori 3-5 on the seg-

ments, borne at a very acute angle to the costule.

Indusia light brown or yellowish, firm.

In dense forests, on trunks and branches of trees,

rarely on forest floor, 100-900 m, San Martin and

Loreto to Pasco, Puno.

West Indies; Guatemala to Costa Rica; Colom-

bia to the Guianas; Ecuador; Peru; Brazil; Africa;

Polynesia.

San Martin: Prov. Mariscal Caceres, Dist. Uchiza, E

of Nuevo Progreso, Schunke V. 3195 (F, GH, us). Dist.

Tocache Nuevo, Puerto Pizana, Schunke V. 6946 (F, MO,

uc). Loreto: Bank of Rio Santiago, above Pongo de Man-

seriche, Mexia 6222 (F, GH, MO, uc, us). Huanuco: Prov.

Huanuco, Las Palmas, Asplund 12635 (s). Pasco (as Ju-

nin): Puerto Bermudez, Killip & Smith 26518 (GH, us).

Puno: Prov. Carabaya, Palmera, Vargas 16161 (GH).

57. Asplenium serra Langsd. & Fisch., Icon. fil.

16, t. 19. 1810. TYPE: Brazil, Santa Catarina,

Langsdorff (holotype, LE; isotype, B, Herb,

mild. 19880; photos, GH & us of B).

Asplenium serra var. remotum Kunze, Linnaea 9: 64.

1834. TYPE: Peru, Pampayacu, Poeppigin 1829

(holotype, LZ, destroyed).

Plants terrestrial, occasionally epiphytic or epi-

petric. Stem short- to long-creeping, abundantly

provided with lanceolate, attenuate, brown or gray-

brown, iridescent, scales, these 3-5 mm long, con-

spicuously clathrate. Leaves 1 -pinnate, approxi-

mate to remote, to 1 m long and 30 cm broad.

Petiole to 50 cm long, dull or slightly lustrous,

dark brown to atropurpureous, sparsely (rarely co-

piously) scaly. Lamina chartaceous (occasionally

firm-herbaceous), oblong-lanceolate, terminating

in a distinct, conform apical segment (this rarely

with a basal lobe), not or scarcely reduced at base.

Rachis nonalate, lacking proliferous buds. Pinnae

8-16 pairs, well spaced, spreading to ascending,

stalked, long-attenuate, cuneate at base (often more

broadly so acroscopically), margins serrate or bi-

serrate (at least at apex). Veins 2-3-forked, as-

cending at very acute (mostly 10-20°) angles, often

subparallel to the costa. Sori inframedial, not ex-

tending into the marginal V? of the pinna. Indusia

yellowish or brown, long and narrow, often ob-

scured by the sporangia at maturity.

In forests, thickets, and edges of clearings, in

Peru commonly on the forest floor, but occasion-

ally low on tree trunks or among rocks, 100-3650

m, Cajamarca to Loreto, south to Ayacucho and

Puno.

West Indies; Mexico to Panama; Colombia and

Venezuela south to Argentina and Paraguay.

This species is commonly found as an epiphyte

from Mexico to Ecuador, yet in Peru and the West

Indies it is most often terrestrial. Also, the petiole

and rachis commonly bear only a few, scattered,

filiform scales, but several collections in Peru have

rachises copiously beset with linear, often atten-

uate, clathrate scales, to 5 mm long: C. Schunke

10, 137, and 655 (F) and Killip & Smith 24615 (F,

GH, us). Curiously, all plants were collected be-

tween 1927 and 1929 in Junin near Schunke's Ha-

cienda, at about 1500 m. It is likely they come

from the same site. Other than density of scales,

they are typical A. serra.

Cajamarca: Prov. Cutervo, "Petroglifos de Llipa,"

Mostacero et al. 1747 (F, GH, HUT, MO, uc). Amazonas:

Prov. Chachapoyas, Calla-Calla slopes, near km 411-

4 1 6 of Leimebamba-Balsas road, Wurdack 1343 (F, GH).

San Martin: Zepelacio, near Moyobamba, Klug 3671 (F,

GH, MO, us). Loreto: Mishuyacu, near Iquitos, Klug 378

(F, us), 7509 (F, us). Huanuco: Prov. Huanuco, Mirador,

road Acomayo to Chinchao, Mexia 7758 (GH, MO, uc,

us). Pasco (as Junin): Pichis Trail, Enenas, Killip & Smith

25649 (F, us). Junin: Schunke Hacienda, above San Ra-

mon, Killip & Smith 24615 (F, GH, us). Ayacucho: Prov.

La Mar, between Tambo San Miguel, Ayna, and Haci-

enda Luisiana, Dudley 11888 (GH). Cuzco: Prov. Pau-

cartambo, Pillawata, Vargas 16714 (GH). Puno: Churu-

mayo, Soukup 867 (F).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

45

58. AspleniumdimidiatumSw.,Prodr. 129. 1788.

TYPE: Jamaica, Swartz (holotype, s; isotypes,

B, Herb. Willd. 19905, UPS, Herb. Thunb.

24798; photos, us of s, GH & us of B).

Plants terrestrial, rarely epiphytic or epipetric.

Stem erect or short-decumbent, abundantly pro-

vided with narrowly clathrate, linear or filiform

scales, these 5-8 mm long, with a tortuous, uni-

costate tip, dark brown to blackish, often with a

fine, lighter brown margin. Leaves 1 -pinnate, sub-

caespitose, 1 8-50 cm long, 6-12 cm broad. Petiole

to 20 cm long, dull, dark brown to blackish, abun-

dantly clathrate-scaly at base, usually less so dis-

tally. Lamina chartaceous or subcoriaceous, grad-

ually reduced to a pinnatifid apex, or to a

nonconform apical segment, not or scarcely re-

duced at base. Rachis nonalate, lacking proliferous

buds, moderately to amply provided with narrow,

attenuate, blackish, clathrate scales. Pinnae 6-12

pairs, well-spaced, sessile or subsessile, lanceolate,

subdimidiate, lacking a costa, apex attenuate, in-

equilateral at base, cuneate acroscopically, excised

basiscopically, margins serrate to deeply and ir-

regularly erose-lacerate, abaxial surface and veins

sparsely to moderately filiform-scaly. Veins re-

peatedly subdichotomous, subparallel or diverg-

ing from each other at 5-10° angles. Sori linear,

crowded, at maturity usually covering pinnae

nearly to margin. Indusia narrow, firm, light or

yellow-brown, entire.

On forest floor or in moist ravines, rarely on

rocks or bases of tree trunks, 650-2000 m, Ama-

zonas, San Martin, Junin, Cuzco.

This has been confused with A. erosum L. of the

Greater Antilles; however, pinnae of the latter are

distinctly stalked and costate, and margins are more

regularly and evenly serrate.

Amazonas: Prov. Bagua, canyon of Rio Utcubamba

on Chachapoyas road, Hutchison and Wright 3793 (GH,

uc). San Martin: Near Tarapoto, Spruce 4753 (BR, GH,

us). Junin: Prov. Tarma, Chanchamayo, Esposto (USM).

Chanchamayo Valley, Puente Paucartambo to La Mer-

ced, Gentry et al. 39823 (F, MO). La Merced, Soukup

3407 (GH, us). Cuzco: Sahuayacu, Biies 843 (us). Prov.

La Convention, Rio Apurimac, 20 min. float below

Puerto Capiro, Davis et al. 1291 (F, uc).

59. Asplenium praemorsum Sw., Prodr. 130. 1788.

TYPE: Jamaica, Swartz (holotype, s; isotype,

UPS, Herb. Thunb. 24834; photo, us of s).

Asplenium nigricans Kunze, Linnaea 9: 69. 1834.

TYPE: Peru, Huanuco, Sierra de Huanuco, "ad

vicum Ambo," Poeppig, in Apr. 1830 (isotype,

w; frag., BM; photos, BM, F & us of w).

Plants epipetric, epiphytic, or sometimes ter-

restrial. Stem erect or decumbent, abundantly pro-

vided with narrowly clathrate, linear to filiform

scales, these 3-7 mm long, usually with a tortuous,

unicostate tip, dark brown to blackish, sometimes

with a fine, lighter brown margin. Leaves 1-pin-

nate-pinnatisect (in America), fasciculate, 12-60

cm long, 3-14 cm broad. Petiole somewhat shorter

than the lamina, dull to slightly lustrous, dark

brown to blackish, abundantly clathrate-scaly at

base, usually less so distally. Lamina subcoria-

ceous, tapering to a pinnatifid apex, not or scarcely

reduced at base. Rachis nonalate, lacking prolif-

erous buds, moderately to copiously scaly as on

the petiole. Pinnae 7-18 pairs, approximate or

subdistant, sessile, or proximal ones very short-

stalked, deeply lobed (or sometimes fully pinnate

at base), the segments cuneiform to obovate, their

apices often lacerate, abaxial surface and veins fi-

liform scaly. Veins indistinct, subparallel or di-

verging from each other at a 5-10° angle. Sori

linear, crowded. Indusia narrow, thin, light or yel-

low-brown, entire.

Common, in forests, usually on rocks and in

crevices or rocky cliffs, often on trunks and

branches of trees, or less frequently on the forest

floor, 600-3700 m, Cajamarca to San Martin, south

to Arequipa and Puno.

West Indies; Mexico to Panama; Venezuela and

Colombia to Brazil and Bolivia; Argentina; Ga-

lapagos Islands.

This variable species has been recognized for a

long time under a number of names, from such

diverse areas as Jamaica, Arabia, and South Af-

rica. Recently, Braithwaite (J. Linn. Soc., Bot. 93:

349. 1986) proposed that all, including A. prae-

morsum, should be included under the oldest name,

A. aethiopicum (Burm.) Bech. African species have

leaves no less than 2-pinnate, and two of the sub-

species recognized by Braithwaite are 2-pinnate-

pinnatifid. In the New World, and particularly in

Peru, leaves are often only 1 -pinnate-pinnatisect,

with the division of pinnae sometimes confined

to one or two basal lobes; so perhaps this taxon

should be recognized as an American variety. An

exhaustive study of the American representatives

of this species complex, such as Brathwaite's in

Africa, is needed before the problem is resolved.

For purposes of this treatment, however, A. prae-

morsum is tentatively maintained as a distinct spe-

cies.

46

FIELDIANA: BOTANY

Cajamarca: Proc. Contumaza, near Guzmango, Sa-

gdstegui 7703 (F, GH, MO). Amazonas: Prov. Chacha-

poyas, slopes of Cano Santa Lucia, Wurdack 732 (F, GH,

uc). La Libertad: Prov. Trujillo, Cerro Campana, Sa-

gdstegui 6193 (GH, MO). San Martin: Dist. Huallaga,

Valley of Rio Apisoncho, Hamilton & Holligan 940 (us).

Ancash: Callejon de Huallas, 8 km SSE of Yungay, Tryon

& Tryon 6548 (F, GH, us). Huanuco: Muna, Macbride

3931 (F, us). Lima: Prov. Canta, Muller & Lopez 12905

(GH). Junin: Satipo, Pichanaki, rodal del Proyecto Peru-

ano-Aleman, Leon 236 (F, uc). Ayacucho: Prov. La Mar,

Ayna, Ldpez & Soukup 15089 (GH). Apurimac: Prov.

Andahuaylas, W of Pincos, Stork & Norton 10714 (F,

uc, us). Cuzco: Machu Picchu, Leon 461 (F, USM). Are-

quipa: Lomas de Arequipa, Coronado 32 (GH, uc, us).

Puno: Prov. Carabaya, below Ollachea, Vargas 6906 (uc,

us), 75997 (GH).

60. Asplenium lividum Kuhn, Linnaea 36: 100.

1 869. TYPE: Venezuela, Aragua, Colonia To-

var, Fendler 156 (holotype, B; isotypes, F!, MO,

us; photo, us of B).

Plants terrestrial. Stem short-creeping, provided

with ovate, narrowly clathrate scales, these 0.5-1

mm long, orange, acute to acuminate. Leaves

1-pinnate-pinnatisect, approximate, 25—45 cm

long, 5-8 cm broad. Petiole somewhat shorter than

the lamina, dull, reddish or grayish brown, non-

alate, naked or with a few appressed scales like

those on the stem. Lamina chartaceous, tapering

to a pinnatifid apex, not or scarcely reduced at

base. Rachis nonalate, lacking proliferous buds,

this and the pinnae abaxially with scattered, ap-

pressed, acuminate scales to 1 mm long. Pinnae

14-20 pairs, larger ones 3-5 cm long, subdistant,

short-stalked, pinnatisect (to nearly pinnate at

base), the 3-6 pairs of segments linear to cunei-

form, their apices often lacerate. Veins indistinct,

subparallel or diverging from each other at 5-10°

angles. Sori linear, crowded. Indusia narrow, thin,

light or yellowish brown, entire.

Thus far known in Peru from one collection, in

damp, shaded woods, about 2100 m, Huanuco.

Venezuela; Peru; Africa.

This has been confused with A. praemorsum, as

the two species are nearly identical in lamina ar-

chitecture, texture, veins, and sori; yet they differ

significantly in habit and indument. The stem of

A. lividum is short-creeping, although somtimes

not obviously so (in the manner of A. laetum), and

bears only a few approximate leaves. Minute, or-

ange, mostly ovate-acuminate, scales are clustered

at the stem apex and also are sparsely scattered

along the axes and veins abaxially. Laminar scales

are only microscopically evident. In marked con-

trast are the erect (rarely decumbent) stem and

densely fasciculate leaves of A. praemorsum, yet

the latter differs even more conspicuously in the

long, dark, filiform, usually copious scales that

spread from the rachis, these quite evident to the

naked eye.

Although dozens of specimens of A. praemor-

sum were examined during the course of this study,

only the two sheets of A. lividum cited here have

been found. However, Andean collections in var-

ious herbaria should be searched for the latter. Any

specimen of A. praemorsum is suspect on which

dark, spreading lamina scales are not macroscop-

ically evident.

Huanuco: Muna, Bryan 417 (F, GH).

61. Asplenium purdieanum Hooker, Icon. PI. 10:

t. 938. 1 854. TYPE: Colombia, Sierra de Santa

Marta, La Fundacion (erroneously reported

by Hooker as Jamaica). Purdie (holotype, K;

isotype, BM; frag., NY).

Plants epipetric or terrestrial. Stem stout, erect,

provided with lanceolate, attenuate, gray-brown,

iridescent scales, these 3-6 mm long, conspicu-

ously clathrate. Leaves 1 -pinnate, caespitose, to

50 cm long. Petiole to 25 cm long, reddish brown

to blackish, dull or slightly lustrous, densely scaly

as on the stem. Lamina thin- to firm-herbaceous,

with 1-2 pairs of pinnae and a conform apical

segment, or in juvenile leaves simple and cordate

to sagittate. Rachis not or scarcely alate, moder-

ately scaly. Pinnae sessile or short-stalked, elliptic

or broadly lanceolate, acute to short-attenuate, cu-

neate at base, margins entire. Veins obscure or

indistinct, anastomosing beyond the middle of the

pinna to form elongate areoles, these lacking in-

cluded veinlets. Sori in a single row on either side

of the costa, spreading at acute angles. Indusia long

and narrow, thin, often hyaline, shrivelling at ma-

turity, the margin entire.

Rare throughout the range, thus far known in

Peru from one collection in San Martin.

Elsewhere, in forests, on rocky soil or in crevices

of limestone, 100-700 m, Honduras; Colombia;

Venezuela; Peru.

This is easily distinguished from other species

in the genus by the few pinnae and conform apical

segment, the densely scaly petiole, and the anas-

tomosing veins.

San Martin: Near Tarapoto, Spruce 4754 (BR, K).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

47

62. Asplenium theciferum (HBK.) Mett., Ann. Sci.

Nat. Bot. 2: 227. 1864. Figure le.

Davallia thecifera HBK., Nov. Gen. Sp. 1: 23. 1815.

TYPE: Venezuela, in monte Saraquen, Hwnboldt

& Bonpland (holotype, P).

Loxoscaphe thecifera (HBK.) Moore, Index fil. 302.

1861.

Plants epiphytic, occasionally epipetric. Stem

erect or decumbent, provided with ovate to lan-

ceolate, bicolorous scales, these 2-3 mm long, gray-

brown to blackish, with narrow lighter brown mar-

gins. Leaves 2-pinnate to 2-pinnate-pinnatifid, fas-

ciculate, 7-25 cm long. Petiole as long as or slightly

longer than the lamina, stout and fleshy, dull

greenish to yellowish, moderately to sparsely pro-

vided with dark, conspicuously clathrate scales

similar to those of the stem. Lamina subcoria-

ceous, deltate- to elliptic-ovate, tapering to a pin-

natifid apex. Rachis greenish, not or slightly alate,

inconspicuously scaly. Pinnae 5-8 pairs, stalked,

divided into linear to clavate segments or pin-

nules, these sometimes incised deeply into tertiary

segments. Veins completely obscured within the

thickened tissue. Indusia cuplike or pocketlike,

borne singly along the acroscopic side of ultimate

segments, near their tips, with sporangia deeply

immersed.

Known thus far in Peru from a single specimen

from Cuzco, 900 m (cited below).

Hispaniola; southern Mexico; Costa Rica; Pan-

ama; Colombia; Venezuela; Ecuador; Peru.

Cuzco: Mercedes, Bties 1736 (us).

Comments

Asplenium denudatum Kuhn, Linnaea 36: 93. 1 869.

TYPE: "Andes Peruvianae" (collector undes-

ignated, B?).

This was originally described by Mettenius, but

not published until Kuhn's emendation of it in

1869. Judging from Kuhn's detailed description,

it is probably A. monanthes. Stem scales are said

to be concolorous, reddish brown, and clathrate,

the lamina 6 in. long, and petioles lustrous and

castaneous, so it is likely to be var. castaneum.

Positive identification awaits study of the type.

Asplenium discolor Kunze, Linnaea 9: 65. 1835.

Kunze identified a collection by Poeppig as

this species ("Peru, on the upper Huallaga,

near Mission Tocache, Poeppig, July, 1830").

A search of several herbaria (B, BM, K, P, w)

proved fruitless, and the original description is too

incomplete to assign specimens to this name. The

species was said to be a foot long, with petiole 4

in. long, rachis subalate, purplish and puberulent,

and pinnae remote and obtusely serrate. These

characters are common to many species of As-

plenium.

Asplenium otites Link, Fil. spec. 2: 60. 1 833. TYPE:

Cultivated, presumably from a South Amer-

ican plant (Link, in Fil. spec. 91. 1841, cited

"Brasilia"), Hort. Mus. Bot. Berol., Cat. No.

053514 (holotype, B!; photos, F, us).

It is difficult to place this name, because the type

is similar to several other South American species,

yet the specimen is too incomplete for definite

comparison. It consists only of a single lamina

with 6 cm of petiole, it lacks a stem, and the lamina

is broken off near the tip. It has been compared

to A. pulchellum, but its pinnae are inequilateral

only at the base and bear 6-8 pairs of veins, which

are mostly forked acroscopically. Pinnae of the

latter are inequilateral throughout the proximal

half, and veins are fewer and unbranched above

the pinna base. It is probably more closely related

to A. drepanophyllum, which does have more nu-

merous and forked veins, but the latter has twice

the number of pinnae and the lamina is more at-

tenuate. The question is not likely to be resolved

with certainty.

Asplenium trapezoides Sw., Syn. fil. 76. 1806.

TYPE: Peru (Junin), Tarma, Lagasca (holo-

type, s; isotypes, B, Herb. Willd. 19869, P;

photos, F & GH of B).

The locality cited is undoubtedly incorrect. It

certainly should be Chile, because the specimen is

clearly the Chilean species A. trilobum Cav., and

the Willdenow Herbarium sheet was so identified

by Hieronymus. This rare fern has not been re-

ported outside Chile for nearly 200 years, since

the original citation of the type of A. trapezoides.

The fasciculate leaves are subtrapezoidal, deeply

crenate or with a pair of basal lobes. They lack a

distinct midrib, and veins are flabellately branched

from the lamina base. Admittedly, this is a small

and inconspicuous fern, but its leaf shape is cu-

rious and distinctive, and if it is indeed in Peru it

certainly should have been rediscovered by now.

48

FIELDIANA: BOTANY

Asplenium weberbaueri Hieron.. Hcdwigia 60: 2 1 1 .

1918. TYPE: Peru (Puno), Tambo Isilluma,

on the way from Sandia to Chanchasmayo,

Weberbauer 1198(B?).

This plant was described as having simple, lin-

ear leaves to 16 cm long and 1.5 cm broad that

taper gradually at both ends, serrate or dentate

margins, short (4 cm) petioles, veins spreading at

20-25° angles, and elongate-deltate stem scales. In

characters of leaf size and shape, it is similar to

Asplenium angustwn, but in lamina margin and

stem scales it seems nearer to A. serratum. In

Hieronymus's discussion, he refers to hook-shaped

spines (stiff scales?) along the midrib. Both the

above-named species have dark scales along the

midrib, but these cannot be termed "hook-shaped."

A canvass of European herbaria failed to locate

the type, so the name cannot be precisely placed

at this time.

Family 19. DAVALLIACEAE

Davalliaceae Frank, Syn. Pflanzensk. (Leunis), ed.

2, 3: 1474. 1877. TYPE: Davallia Sm.

NephrolepidaceaePic.-Ser., Webbia 29: 8. 1974. TYPE:

Nephrolepis Schott.

Stem erect, decumbent, or long-creeping, mod-

erately stout to slender, more or less indurated,

bearing scales. Leaves ca. 1 5 cm to 3 m long, entire

to usually pinnate, circinate in vernation, petiole

without stipules. Sori roundish to somewhat elon-

gate, rarely linear, abaxial, near or back of the

margin, or on an intramarginal commissure, usu-

ally not paraphysate, usually indusiate. Sporan-

gium with a 2-rowed stalk below its apex, with a

usually long stalk, the vertical annulus interrupted

by the stalk.

The Davalliaceae are a family of about 1 0 or

fewer genera of the Old World, with Nephrolepis

also in America. Most species are epiphytes with

a dorsiventral stele and leaves that are articulate

near the base of the petiole.

Nephrolepis, the only genus in America, is in-

cluded in the family with some uncertainty. The

tuberculate spores and staurocytic stomata suggest

a closer alliance with the davallioid ferns than with

other groups.

I. Nephrolepis

Nephrolepis Schott, Gen. fil. no. 3. 1834. TYPE:

Nephrolepis exaltata (L.) Schott (Polypodium

exaltatum L.). Figure 2.

Stem erect or decumbent, bearing scales, freely

stoloniferous. Leaves monomorphic or slightly di-

morphic with the fertile pinnae narrower than the

sterile. Lamina 1 -pinnate, the pinnae articulate,

entire or nearly so, glabrous, pubescent, and (or)

scaly, the scales usually dissected, with hydathodes

adaxially that usually have a thin, whitish, circular

secretion, veins free. Sori roundish to lunate, borne

on the vein ends, not paraphysate, covered by an

orbicular to lunate indusium. Spores ellipsoidal,

monolete, the surface irregularly tuberculate to ru-

gose.

Nephrolepis is a tropical and subtropical genus

of perhaps 20 species, with five native species in

America and four in Peru.

The species of Nephrolepis are not well under-

stood and, accordingly, current accounts of the

genus in tropical America and other tropical regions

often differ in the application of names and in the

primary characters employed for identification.

The following treatment is based primarily on Pe-

ruvian materials, and differences with other mod-

ern treatments do not necessarily imply alterations

in the taxonomy of American species in other

regions.

Several species and their cultivated variations

are commonly maintained in gardens and may

escape to more or less native habitats, either

through spore dispersal or by the growth of dis-

carded cultivated plants. Some of the many cul-

tivated variations (cultivars) have rarely been col-

lected in Peru. These usually have pinnatifid to

several times pinnate pinnae, forked pinnae or ra-

chises, or nearly orbicular pinnae. They are not

included in the treatment of the native and ad-

ventive species.

The key to species emphasizes the primary char-

acters of the species, while uncommon variations

are mostly mentioned in the descriptions.

References

NAUMAN, C. E. 1981. The genus Nephrolepis in

Florida. Amer. Fern J., 71: 35-40.

SMITH, A. R. 1981. Nephrolepis, pp. 151-154.

In Flora of Chiapas, part 2, Pteridophytes.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

49

50

FIELDIANA: BOTANY

California Academy of Sciences, San Fran-

cisco.

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

phrolepis, pp. 656-662. In Ferns and Allied

Plants. Springer- Verlag, New York.

Key to Species of Nephrolepis

\ . Indusia orbicular to orbicular-reniform, with a narrow to rather open sinus; mature sporangia pro-

jecting all around the indusium b

b. Pinnae short- or very short-stalked; costa definitely pubescent adaxially, or if glabrous then the

base of the pinnae equilateral or nearly so c

c. Base of the pinnae equilateral or nearly so, sometimes biauriculate, or slightly and broadly

auriculate only on the acroscopic side; costa adaxially with long, often flexuous, trichomes and

usually some dissected scales; stem scales light brown to brown, more or less spreading

1 . N. biserrata

c. Base of the pinnae, or most of them, inequilateral, the acroscopic side narrowly auriculate;

costa adaxially with short to rarely moderately long, stiff trichomes; stem scales dark brown

to blackish, appressed 2. N. multiflora

b. Pinnae sessile; costa glabrous or nearly so adaxially; base of the pinnae inequilateral, the acroscopic

side broadly and acutely to subacutely auriculate 3. N. rivularis

a. Indusia reniform, with a broad sinus, to lunate; mature sporangia projecting from the free portion

of the indusium d

d. Fertile pinnae with the basal sorus on the acroscopic side and the basal sorus on the basiscopic

side ca. equally distant from, or close to, the base of the pinna, or many of them so; the distal

pinnae, but not necessarily the apical ones, rounded, usually broadly so, at the base on the basiscopic

side 4. N. cordifolia

d. Fertile pinnae with the basal sorus on the acroscopic side much closer to the base of the pinna

than the basal sorus on the basiscopic side, or most of them so; especially the distal pinnae cuneate

at the base on the basiscopic side to sometimes slightly rounded 5. N. pectinata

1 . Nephrolepis biserrata (Sw.) Schott, Gen. fil. no.

3. 1834.

Aspidium biserratum Sw., J. Bot. (Schrader) 1800(2):

32. 1802. TYPE: Mauritius, Grondahl (Grondal)

(holotype, Herb. Sw., s!).

Tectaria fraxinea Cav., Descr. pi. 250. 1802. TYPE:

Peru. Obragillo, Nee, MA, seen by C. Chr. (Dansk

Bot. Ark. 9(3): 15. 1937 and referred here).

Stem scales light brown to brown, more or less

spreading. Pinnae short-stalked, pubescent to rarely

glabrous abaxially. the costa pubescent adaxially

with long, often flexuous, trichomes and usually

some dissected scales, rarely the trichomes short

or the costa glabrous, the base equilateral or nearly

so, sometimes biauriculate, or slightly and broadly

auriculate only on the acroscopic side, broadly

rounded to broadly cuneate on the basiscopic side.

Lamina of fertile leaves ca. 10-30 cm broad. Basal

sori on the acroscopic and basiscopic sides equally

close to the base of the pinna. Indusium orbicular,

with a narrow sinus, the mature sporangia pro-

jecting all around the indusium.

Epiphytic or usually terrestrial, along roadsides,

in secondary growth, along forest borders, and in

tall forests, 1 00-900 m, Cajamarca and Amazonas

to Ayacucho.

Tropical America; Old World.

The lamina of this species is usually consider-

ably broader than that of other Peruvian species.

Even in rather small leaves it is rarely less than

10 cm broad, and in large leaves, which may be

up to ca. 3 m long, it is commonly 30 cm broad

or more. Small plants may have leaves similar to

those of N. multiflora but that species has the pin-

nae inequilateral at the base, dark, appressed stem

FIG. 2. Nephrolepis pectinata: a, habit; b, fertile pinna. Nephrolepis cordifolia: c, fertile pinna. Nephrolepis mul-

tiflora: d, pinna base. Nephrolepis rivularis: e, pinna base. (Adapted from Stolze, Ferns and Fern Allies of Guatemala,

1981.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

51

scales, and usually has the pinnae abaxially fibril-

lose-scaly, especially at the base.

Cajamarca: Prov. Jaen, Quebrada Tabaconas, Ferrey-

ra & Sanchez 19646 (USM). Amazonas: Prov. Bagua, Chi-

riaco-Puente Venezuela, Barbour 4331 (MO). La Poza,

Rio Santiago, Berlin 3701 (MO). San Martin: NW of San

Martin, Soukup 5218 (GH). Prov. San Martin, Vaca Pozo,

Woytkowski 35136 (MO). Prov. San Martin, Ekin, Woyl-

kowski 35230 (MO, uc). Loreto: Prov. Maynas, Rio Itaya,

Tryon & Tryon 5176 (BM, F, GH, us, USM). Prov. Maynas,

3 km S of Iquitos, Tryon & Tryon 5163 (BM, F, GH, u,

us, USM). Huanuco: Prov. Pachitea, Bosque Nacional de

Iparia, J. Schunke V. 1735 (F, GH, us). Ayacucho: Kim-

pitiriki, Rio Apurimac, Killip & Smith 22856 (F, us).

2. Nephrolepis multiflora (Roxb.) Morton, Contr.

U.S. Natl. Herb. 38: 309. 1974. Figure 2d.

Davallia multiflora Roxb., Calcutta J. Nat. Hist. 4:

515, t. 31 (left), 1844. LECTOTYPE: by Morton,

he. cit., India, Roxburgh (BR; frag, and photo, us).

Stem scales dark brown to blackish, appressed.

Pinnae short- or very short-stalked, fibrillose-scaly

abaxially, especially toward the base and (or) pu-

bescent, the costa adaxially with usually short and

stiff, to moderately long, trichomes, rarely with

some dissected scales or long trichomes, the base

inequilateral, narrow auriculate on the acroscopic

side, broadly rounded to slightly auriculate on the

basiscopic side. Lamina of fertile leaves ca. 8-20

cm broad. Basal sori on the acroscopic and basi-

scopic sides equally close to the base of the pinna.

Indusium orbicular to orbicular- reniform, with a

narrow to rather open sinus, the mature sporangia

projecting all around the indusium.

Terrestrial, in primary forests and also disturbed

areas, 120-2100 m, Cajamarca and Loreto.

Paleotropics; sporadically adventive in tropical

America.

This species is probably more frequent in Peru

than the few collections indicate. It is distin-

guished by the abaxially fibrillose-scaly pinnae and

the stem scales that are dark and appressed. It has

perhaps been confused with another paleo tropical

species, Nephrolepis hirsutula (Forster) Presl, which

has the adaxial side of the costa moderately to

densely invested with dissected scales, rather than

pubescent as in N. multiflora. The species N. hir-

sutula is not known to be adventive in Peru.

Cajamarca: Provicutervo, San Andres, Quiroz 2840

(F). Loreto: Prov. Maynas, Puerto Almendras, Vasquez

& Jaramillo 1415 (MO). Prov. Maynas, Quistococha,

Vasquez & Jaramillo 4079 (F). Iquitos, Mitller & Miiller

2508 (GH). Quistococha, Gentry & Ayala 15531 (us).

3. Neprolepis rivularis (Vahl) Krug, Bot. Jahrb.

Syst. 24: 122. 1897.

Polypodium rivulare Vahl, Eclog. amer. 3: 51. 1807.

TYPE: West Indies, Montserrat, Ryan (holotype,

c).

Stem scales light to dark brown, appressed to

spreading. Pinnae sessile, glabrous to slightly pu-

bescent abaxially, and (or) with a few dissected

scales, rarely minutely pubescent, the costa gla-

brous or nearly so adaxially, the base inequilateral,

broadly acutely or subacutely auriculate on the

acroscopic side, cuneate to rounded on the basi-

scopic side. Lamina of fertile leaves ca. 8-15 cm

broad. Basal sori on the acroscopic and basiscopic

sides ca. equally close to the base of the pinna, or

closer on one side than the other. Indusium or-

bicular, with a narrow sinus, the mature sporangia

projecting all around the indusium.

Epiphytic, or usually terrestrial, in disturbed

vegetation, secondary forests, or primary forests,

100-900 m, San Martin to Puno.

Tropical America.

This is a distinctive species, especially in its

character of sessile pinnae. Contrary to some au-

thors, it appears that the combination Nephrolepis

rivularis was validly published by Krug, although

the basionym was not cited.

San Martin: Soritor, Woytkowski 6246 (GH, us). Lo-

reto: Gamitanacocha, Rio Mazan, Schunke 279 (GH, uc).

Prov. Maynas, Rio Nanay, McDaniel 13685 (GH), Hick-

ok 662 (GH). Quista Cocha (Quistococha), near Iquitos,

McDaniel 1 1739 (GH). Pasco: Prov. Oxapampa, Palcazu

valley, Foster 9503 (MO). Cuzco: Prov. Paucartambo,

Kosnipata-Pilcopata, Vargas 14750 (GH). Madre de Dios:

Prov. Tambopata, Tambopata Nature Reserve, Barbour

4783 (MO), 5124 (F, MO). Puno: San Gavan (Gaban),

Lechler 2515 (GH, K).

4. Nephrolepis cordifolia (L.) Presl, Tent.

Pterid. 79. 1836. Figure 2c.

Polypodium cordifolium L. Sp. pi. 2: 1089. 1 753. TYPE:

Petiver, Pteri-graphia amer., t. l,f. 11 (not seen

in Petiver, Hort. Sice, in Herb. Sloane, BM).

Aspidium pendulum Raddi, Opusc. Sci 3: 289. 1819.

TYPE: Brazil, Raddi (holotype, FI; isotype, K).

Nephrolepis pendula (Raddi) J. Sm., J. Bot. (Hooker)

4: 197. 1841.

52

FIELDIANA: BOTANY

Stem scales light to dark brown to blackish, more

or less appressed. Pinnae short- to very short-

stalked, essentially glabrous, the costa essentially

glabrous adaxially, the base inequilateral or some-

what inequilateral, slightly to definitely auriculate

on the acroscopic side, rounded to broadly round-

ed or sometimes slightly auriculate on the basi-

scopic side. Lamina of fertile leaves ca. 3-10 cm

broad. Basal sorus on the acroscopic and basi-

scopic sides ca. equally distant from, or close to,

the base of the pinna, or most of them so. Indusium

reniform, with a broad sinus, to lunate, mature

sporangia projecting from the free portion of the

indusium.

Epiphytic, rupestral, or terrestrial, rocky places,

rarely on cliffs, cloud forests, forest borders, pri-

mary forests, shrubby hillsides, or on fallen trunks

and branches, 125-2800 m, Cajamarca and Ama-

zonas south to Puno.

Tropical America; Old World.

The Petiver illustration is not adequate for the

purposes of accurate identification. It is said to be

copied from Plumier, Traite foug. amer., t. 71,

from Hispaniola, which is more clearly, and usu-

ally identified as, Nephrolepis cordifolia.

Some epiphytic plants have very long, pendent

leaves, up to 3 m and perhaps longer. However,

in comparison to the long leaves of Nephrolepis

biserrata, the lamina of those of N. cordifolia is

narrow, about 8-10 cm broad. Sometimes there

is a single basal basiscopic sorus closer to the base

of the pinna than the basal basiscopic sorus.

At times, Nephrolepis pendula is adopted for this

species because its application is certain. At other

times, N. pendula is considered to be a separate

species from N. cordifolia. At least in Peru the two

do not seem to be separable since there are inter-

mediate specimens.

This and the following species Nephrolepis pec-

tinata are the most widely distributed and com-

mon species of the genus in Peru. A discussion of

the differences between them is provided under N.

pectinata.

Cajamarca: Prov. Santa Cruz, Bosque de Monteseco,

Sagdstegui et al. 12402 (F, MO). Prov. Cutervo, Cutervo-

Socota, Lopez et al. 5316 (GH). Amazonas: Prov. Bon-

gara, Laguna Pomacocha, Wurdack 883 (GH, us). Prov.

Chachapoyas, entre Chachapoyas y Caclic, Lopez et al.

4350 (GH, MO). San Martin: Zepelacio, near Moyobam-

ba, Klug 3736 (F, GH, us). Prov. Mariscal Caceres, Dis-

trito Tocache Nuevo, J. Schunke V. 4337 (F, GH, us).

Loreto: Above Pongo de Manserische, Mexia 6163 (F,

GH, uc, us). Huanuco: Prov. Pachitea, Bosque Nacional

de Iparia, J. Schunke V. 2940 (F, GH, us). Hacienda Paty,

abajo de Carpish, Ferreyra 9353 (GH, USM). Pasco: Prov.

Oxapampa, Oxapampa-Cerro de Pasco, D. Smith 4083

(GH). Prov. Oxapampa, N of Chequitavo, D. Smith 5108

(GH). Junin: Huacapistana, Tryon & Tryon 5452 (BM, F,

GH, u, us, USM). Ucayali: Prov. Coronel Portillo, Bo-

queron del Padre Abad, Ridoutl (GH, USM). Ayacucho:

Prov. La Mar, eastern massif of the Cordillera Central,

Dudley 11724 (GH), 11880 (GH, us). Cuzco: Prov. La

Convention, Rio Apurimac, Davis et al. 1328 (F, GH).

Prov. Urubamba, Macchu Picchu station, Sounders 1230

(F, GH). Puno: Prov. Sandia, Asalaya, Vargas 1483 (F).

5. Nephrolepis pectinata (Willd.) Schott, Gen. fil.

no. 3. 1834.

Aspidium pectinatum Willd., Sp. pi. 5: 223. 1810.

TYPE:"exAiton"(holotype,//er6. Willd. 19753.

B!; photo, GH). Figures 2a-b.

Stem scales brown to blackish, more or less ap-

pressed. Pinnae sessile to usually short-stalked,

essentially glabrous abaxially, or rarely minutely

pubescent, the costa essentially glabrous adaxially,

to rarely partly minutely pubescent, the base in-

equilateral, auriculate on the acroscopic side, es-

pecially the distal pinnae cuneate to slightly round-

ed on the basiscopic side. Lamina of fertile leaves

ca. 2-4 cm broad. Basal sorus on the acroscopic

side much closer to the base of the pinna than the

basal sorus on the basiscopic side. Indusium reni-

form, with a broad sinus, to lunate, mature spo-

rangia projecting from the free portion of the in-

dusium.

Epiphytic, rupestral, or terrestrial, rocky places

in woods, in primary forests, cloud forests, on

stream banks, forest borders, in moist and rough

pastures, on fallen trunks and branches, 200-2900

m, Cajamarca and Amazonas to Puno.

Tropical America.

Nephrolepis pectinata is distinguished by hav-

ing, especially in the distal portion of the lamina,

the pinnae with a cuneate (evenly tapering) base

on the basiscopic side and having the basal sorus

on the acroscopic side much closer to the base of

the pinna than the basal sorus on the basiscopic

side. Sometimes N. pectinata has the base of the

pinna on the basiscopic side slightly rounded, and

N. cordifolia may have a single basal acroscopic

sorus closer to the base of the pinna than the basal

basiscopic sorus.

Nephrolepis occidentalisKun7.e(L\nnaea 18: 343.

1844) is usually recognized as a species, with a

poorly developed stem and thin, long-triangular,

crenate to shallowly lobed pinnae with the veins

clearly evident abaxially. However, at least in South

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

53

America, there is intergradation between speci-

mens with the characters of N. occidentalis and

those of N. pectinata and it is not possible to rec-

ognize the former as a distinct species in that con-

tinent. Plants referred to N. occidentalis usually

grow in especially damp and shaded situations,

and they may represent an ecological variation

(shade form) or an ecotype of N. pectinata.

Cajamarca: Prov. Santa Cruz, Monteseco, Sagdstegui

et al. 12402 (HUT). Amazonas: Prov. Bagua, Chiriaco to

Puente Venezuela, Barbour 4401 (MO). San Martin: Prov.

Lamas, Lamas, Belshaw 3441 (F, GH). Loreto: Above

Pongo de Manserische, Mexia 6219 (F, GH, uc, us). Pu-

mayacu, between Balsapuerto and Moyobamba, Klug

3246 (F, GH, us). Huanuco: Muna, Bryan 415 (F, GH).

Prov. Huanuco, above Cayumba, Rio Huallaga, Mexia

8324 (F, GH, uc, us). Prov. Huanuco, Hacienda Exito,

Mexia 8135 (F, GH, uc, us). Pasco: Prov. Oxapampa,

between Oxapampa and Paucartambo, Smith & Pretel

1485 (F, GH). Junin: Prov. Satipo, S of Chequitavo, D.

Smith 5154 (GH). Above San Ramon, Killip & Smith

24613 (F, GH, us). Ucayali: Prov. Coronel Portillo, La

Divisoria (as Loreto), Dillon 2630 (F, MO). Prov. Coronel

Portillo, Boqueron del Padre Abad, Ferreyra 8109 (GH),

8129 (GH). Cuzco: Prov. Quispicanchi, San Pedro, Var-

gas 9748 (GH). Prov. La Convention, Dudley 10432 (GH).

Madre de Dios: Manu, Atalaya, Foster & Wackier 7415

(MO). Puno: Prov. Carabaya, Hacienda Palmera, Vargas

16123 (GH).

Comments

Nephrolepis hirsutula (Forst.) Presl, Tent. Pterid.

79. 1836.

Polypodium hirsutulum Forst. Prod. 81. 1786. TYPE:

South Sea Islands, probably Tahiti, Forster, pho-

to, us of BM (see Morton, Contr. U.S. Natl. Herb.

38: 345. 1974).

This species is native in the Malaysian and Pa-

cific regions and is sparingly adventive in the

American tropics. It may be collected in Peru. It

is similar to N. biserrata but the adaxial surface

of the costa bears dissected scales and lacks tri-

chomes; also, a single, prominent, basal auricle is

usually on the acroscopic side of the pinnae.

Family 20. BLECHNACEAE

Blechnaceae (Presl) Copel., Gen. fil. 155. 1947.

Blechneae Presl, Epim. bot. 103. 1851, as Blechna-

ceae. TYPE: Blechnum L.

Stem erect and slender to very thick, to decum-

bent and small to stout, or long-creeping, or scan-

dent, hardly sclerotic to strongly indurated, bear-

ing scales. Leaves ca. 10 cm to 1 5 m long, circinate

in vernation, monomorphic to dimorphic, the

lamina entire to usually pinnatisect or 1-2-pin-

nate, glabrous, minutely glandular, pubescent, or

scaly abaxially. Petiole lacking stipules, not artic-

ulate to the stem. Veins wholly free or partly anas-

tomosing. Sori abaxial, short to very long, borne

on a vein (or sometimes the sporangia also borne

on the leaf-tissue), usually covered by an indusium

that opens on the side of the axis at the segment

or exindusiate, not paraphysate. Sporangia with a

2-3-rowed stalk and a vertical or nearly vertical

annulus, which is usually interrupted by the stalk.

The Blechnaceae are a family of nine genera and

about 175 species. There are two genera and 20

species in Peru. The spores are monolete, and the

young leaves are usually tinged with red.

Key to Genera of Blechnaceae

a. Leaf to ca. 2 m long, determinate; lamina entire to 1 -pinnate I. Blechnum

a. Leaf climbing to 15 m or more, indeterminate; lamina 2-pinnate II. Salpichlaena

FIG. 3. Blechnum asplenioides: a, habit; b, fertile pinnae. Blechnum fraxineum: c, habit. Blechnum binervatum

ssp. fragile: d, habit; e, portion of rachis with pinna base; f, stem scales. (Adapted from Stolze, Ferns and Fern Allies

of Guatemala, 1981.)

54

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

55

I. Blechnum

Blechnum L., Sp. pi. 1077. 1753; Gen. pi. ed. 5,

485, 1754. TYPE: Blechnum occidental L. (as B.

orientale). Figure 3.

Lomaria Willd., Ges. Naturf. Freunde Berlin Mag.

Neuesten Entdeck. Gesammten Naturk. 3: 160.

1809. TYPE: Lomaria nuda (Labill.) Willd. =

Blechnum nudum (Labill.) Luers.

Terrestrial, rupestral, or sometimes epiphytic,

or scandent-epiphytic. Stem decumbent, or long-

creeping, or scandent, or erect and small to mas-

sive, often producing stolons. Leaves monomor-

phic to dimorphic and the fertile usually longer

than the sterile and with narrower segments. Lam-

ina entire or usually pinnatisect to 1 -pinnate. Veins

free or rarely partly anastomosing and without in-

cluded veinlets. Sori short to long, on a vascular

commissure close to the costa, not paraphysate,

covered by an indusium of similar extent that opens

toward the costa. Spores ellipsoidal, monolete,

nearly smooth, slightly papillate, rugose, reticu-

late, or echinate.

Blechnum is a nearly worldwide genus of per-

haps 150 species, 19 of them in Peru. Although

infrageneric classifications are sometimes pre-

sented (Smith, 1985), there is uncertainty con-

cerning the relations of several species, and only

two subgenera are mentioned here: subgenus

Blechnum (species 1-7), with expanded green tis-

sue beyond the attachment of the indusium and

with monomorphic leaves, and subgenus Lomaria

(Willd.) C. Chr. (Index fil. 1 50, 40 1 . 1 905) (species

8-19), with little or no green tissue beyond the

attachment of the indusium and with usually di-

morphic leaves.

Species of Blechnum are often poorly defined,

due in part to genetic and phenotypic variation

and also to hybridization. Walker (1985) reports

two hybrids in subgenus Blechnum and suggests

that these probably represent but a few of those

that occur in nature. For the purposes of identi-

fication of the hybrid (Blechnum x caudatum (3 x )

B. fraxineum (2 x ) x occidental (4 x ) is placed

under B. occidentale, and the hybrid B. asplenioi-

des x fraxineum (4x) is treated under B. con-

fluens. Much more cytological data are needed be-

fore the taxonomy of the Andean Blechnum

occidentale complex (species 1-5) is adequately

known. Walker ( 1 985) and Mickel and Beitel (1988)

discuss the characters of hybrids in this complex.

Various characters, sometimes employed as

species distinctions, have been found to be too

variable in Peru to merit their use— for example,

the presence or absence of aerophores on the lam-

ina (at the base of the pinnae abaxially) or the

shape of the aerophores, and pubescence and glan-

dularity of the lamina.

In this treatment, a pinna is considered to be a

stalked or sessile segment or an adnate one that is

longer than broad. A lobe is considered to be a

reduced segment that is adnate and broader than

long. While this treatment is centered on the spe-

cies of Peru, it includes an assessment of all South

American Blechnum materials. However, species

tend to be more variable in the Andes than to the

north, and we do not wish to imply that other

species of Mexico, Central America, and the West

Indies are not probably recognized.

References

BROADHURST, J. 1912. The genus Struthiopteris

and its representatives in North America. I, II.

Bull. Torrey Bot. Club, 39: 257-278, 357-385.

LOOSER, G. 1947. Los Blechnum (Filices) de

Chile. Rev. Universit. (Univ. Catol. Chile), 32(2):

7-106.

MICKEL, J. T., AND J. M. BEITEL. 1988. Pteri-

dophyta flora of Oaxaca, Mexico. Mem. New

York Bot. Card., 46.

MURILLO, M. T. 1968. Blechnum subgenus

Blechnum en sur America. Nova Hedwigia, 16:

229-366.

SMITH, A. R. 1981. Flora of Chiapas, 2, pp. 1-

370. In Breedlove, ed., Pteridophytes. Califor-

nia Academy of Sciences, San Francisco.

. 1985. Pteridophytes of Venezuela, an

Annotated List. University of California, Berke-

ley (not formally published).

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

num, pp. 669-680. In Ferns and Allied Plants.

Springer- Verlag, New York.

WALKER, T. G. 1985. The cytological and tax-

onomic implications. In A. C. Jermy and T. G.

Walker, eds., Cytotaxonomic Studies of the Ferns

of Trinidad. Bull. Brit. Mus. (Nat. Hist.) Bot.,

13: 133-276.

56

FIELDIANA: BOTANY

Key to Species of Blechnum

a. Fertile segments with expanded green tissue beyond the attachment of the indusium; leaves mono-

morphic, the sterile segments with essentially flat margins b

b. Lamina entire, rarely with 1 or 2 small pinnae or lobes at the base 5. B. lanceola

b. Lamina pi n na t i lid to 1 -pinnate c

c. Lamina 1 -pinnate; pinnae with the short stalk articulate to the rachis 6. B. serrulatum

c. Lamina often pinnatifid, pinnae of a 1 -pinnate lamina with the stalk continuous with the rachis

d

d. Lamina with the apex abruptly reduced to an apical segment that is longer than the longest

pinna 3. B. fraxineum

d. Lamina with the apex gradually reduced, the apex pinnatifid or with an entire tip that is

usually shorter than the longest pinna e

e. Lamina usually not or only slightly reduced at the base; basal pinnae truncate or semi-

cordate to cordate, not fully adnate to the rachis 1 . B. occidentale

e. Lamina reduced at the base; basal pinnae fully adnate to the rachis although an auricle

may be partly free f

f. Scales at the base of the petiole usually few, brown to blackish and usually nearly

concolorous; the longest pinna ca. 0.5-4 cm long g

g. Lamina hardly reduced at the base, or more or less abruptly reduced to basal pinnae

that are as long as broad or longer than broad 2. B. confluens

g. Lamina gradually reduced at the base to lobes that are broader than long

4. B. asplenioides

f. Scales at the base of the petiole numerous, wholly blackish or with a blackish center

and brown margins; the longest pinna (7-)15-20(-25) cm long . . 7. B. brasiliense

a. Fertile segments contracted, with little or no green tissue extending beyond the attachment of the

indusium; leaves dimorphic or if monomorphic then the sterile segments also contracted and with

involute margins h

h. Scales at the base of the petiole absent, few, or numerous, lanceolate to usually lanceolate-ovate;

stem decument, erect or climbing, rarely arborescent i

i. Leaves monomorphic; pinnae of the sterile lamina strongly involute, the margins usually reach-

ing or nearly reaching the costa on the abaxial side 15. B. loxense

i. Leaves dimorphic; pinnae of the sterile lamina flat or somewhat involute and the margins

distant from the costa on the abaxial side j

j. Pinnae of the sterile lamina narrowly or broadly adnate, or most of them so k

k. Adult plants with a scandent stem, hemiepiphytic or epiphytic, its scales, especially the

older ones, mostly bicolorous with a blackish center and lighter margins

13. B. binervatum

k. Adult plants terrestrial, the stem scales concolorous 1

1. Sterile lamina abruptly reduced at the base, sometimes to a wing or to lobes, 1 2-30

cm broad, the petiole with some very small, inconspicuous lobes 12. B. divergens

1. Sterile lamina gradually reduced at the base, ca. 1.5-10 cm broad m

m. Sterile lamina reduced at the base to small pinnae that are 2-4 mm long

9. B. andinum

m. Sterile lamina reduced at the base to definite lobes n

n. Pinnae of the sterile lamina mostly curved, ascending 8. B. lehmannii

n. Pinnae of the sterile lamina straight, strictly patent or nearly so

10. B. penna-marina

j. Pinnae of the sterile lamina sessile or short-stalked, or most of them so o

o. Sterile lamina with often biauriculate pinnae, with a much prolonged, viviparous apex

of reduced pinnae 16. B. sprucei

o. Sterile lamina with cuneate, broadly rounded to cordate, not auriculate pinnae, the apex

gradually to abruptly reduced, not viviparous p

p. Apex of the sterile lamina gradually reduced to an entire tip; many or most (rarely

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 57

only some) of the scales of the costa and rachis abaxially denticulate to ciliate, more

or less spreading, light brown to dark brown and then sometimes bicolorous

14. B. stipitellatum

p. Apex of the sterile lamina abruptly reduced to a large, entire, terminal segment; scales

of the costae and rachis abaxially entire, rarely denticulate or ciliate, mostly appressed,

whitish to light brown 11. B. cordatum

h. Scales at the base of the petiole numerous, long, acicular beyond the usually broader base, curved;

stem usually arborescent q

q. Sterile lamina with few or no fully adnate pinnae distally, with the base of the lamina abruptly

and usually hardly reduced r

r. Sterile lamina with the pinnae mostly patent or slightly ascending; pinnae with the apex

acute to long-acuminate, the margins flat, or at least the tip flat or nearly so; stem mostly

bearing few leaves 17. B. schomburgkii

r. Sterile lamina with the pinnae ascending to strongly ascending; pinnae with the apex obtuse,

the margins involute to strongly involute, the tip involute; stem bearing a dense crown of

numerous leaves 18. B. aural um

q. Sterile lamina with many fully adnate pinnae distally, with the lamina base gradually reduced

.19. B. obtusifolium

1. Blechnum occidentale L., Sp. pi. 1077, as "ori-

entale." 1753. TYPE: LINN 1247.1, photo A

is this species.

Blechnum caudatum Cav., Descr. pi. 262. 1802. TYPE

(designated by C. Chr., Dansk Bot. Ark. 9(3): 20.

1937): Ecuador, San Antonio, Nee (holotype, MA).

Blechnum glandulosum Link, Enum. hort. berol. alt.

2: 462. 1822. TYPE: Cultivated, "Hab in Brasi-

lia" (holotype, Herb. Link, B; photo, GH).

Blechnum pectinatum Hooker, Hooker's Icon. PI. 1 :

t. 95. 1837, not Presl, 1825. TYPE: Peru, Casapi,

Mathews (holotype, K).

Blechnum x caudatum Cav. (pro sp.) (T. G. Walker,

1985).

Stem small, erect to usually decumbent and

short-creeping. Leaves monomorphic. Petiole with

basal scales rather few, short, mostly lanceolate to

ovate-lanceolate, stem scales wholly light brown

to dark brown or often with lighter margins. Lam-

ina pinnatisect to 1 -pinnate at the base, the apex

gradually reduced, pinnatifid or with an entire tip

shorter than the longest lateral pinna, not or slight-

ly reduced at the base (rarely more), the pinnae

essentially flat, the basal ones truncate, semicor-

date to cordate. Fertile pinnae with green tissues

extending well beyond the attachment of the in-

dusium.

Terrestrial and rupestral, lowland rain forests,

montane forests, cloud forests, along streams, on

banks, in thickets, among and on rocks, usually in

moist and shaded sites, 400-3800 m, usually at

700-2400 m, Tumbes south to Arequipa and Puno.

Tropical America.

This is a common and variable species, part of

the variation undoubtedly due to hybridization.

Walker (1985) noted the hybrid B. fraxineum x

occidentale, which is called Blechnum x caudatum

Cav. While the morphology of the hybrid may

agree with the holotype, the latter cannot be cy-

tologically known and may represent a homoploid

variation of B. occidentale. Walker also repeated

his report of Blechnum asplenioides (as B. unila-

terale) x occidentale. Mexia8216, Dept. Huanuco

(MO), is clearly B. fraxineum x occidentale; an-

other sheet of the same collection (GH) is B. oc-

cidentale.

Blechnum occidentale has a rachis that is gla-

brous or nearly so, and in Peru it grows from 400

to 3850 m. It is much more common than Blech-

num glandulosum (B. occidentale var. pubirhachis

Rosenst.), which has a definitely glandular-pubes-

cent rachis. The two variations are altitudinally

distinct in Central America (Stolze, Fieldiana, Bot.,

n.s., 6: 110-111. 1981) but not in Peru where there

is considerable overlap in the altitude. The glan-

dular-pubescent variation occurs from 650 to 2700

m. Accordingly, the two are treated as one taxon

in Peru.

Tumbes: Between Caucho and Cotrina, Coronado 219

(GH). Prov. Zarumilla, near Campo Verde, Simpson &

Schunke 457 (F, us, USM). Piura: Prov. Huancabamba,

Sagdstegui & Cabanillas 8543 (MO, us). Lambayeque:

Prov. Lambayeque, km 45 from Olmos, Plowman et al.

14282 (F, GH). Cajamarca: Prov. Contumaza, El Molino,

Sagdstegui et al. 10895 (F). Amazonas: Prov. Bagua, entre

Aramango y Montenegro, Lopez et al. 4217 (GH, us).

Just E of Chachapoyas, Wurdack 730 (F). La Libertad:

Prov. Otuzco, Dist. Huaranchal, Sagdstegui 95 (HUT).

San Martin: Prope Tarapoto, Spruce 3950 (GH). San

Roque, LI. Williams 7029, 7269, 7523 (F). Huanuco:

58

FIELDIANA: BOTANY

Prov. Huanuco, Dist. Churubamba, Mexia 8137 (F, GH,

MO). Prov. Pachitea, Panao, Ferreyra 1791 (GH, USM).

Prov. Huanuco, Puente Durand, Stork & Morton 9425

(F, MO, us). Lima: Lomas de Atocongo, Coronado 291

(F, GH). Chosica, Macbride & Featherstone 489 (F, us).

Prov. Huarochiri, Leon 177 (MO, USM). Pasco: 8 km N

of Huancabamba, Skog et al. 5074 (us). Junin: Cerca a

San Ramon, Cerrate 962 (GH). La Merced, Killip & Smith

23760 (F, us). Huancavelica: Prov. Tayacaja, SE of Tin-

tay, Tovar 4704 (GH). Ayacucho: Estrella, between Huan-

ta and Rio Apurimac, Killip & Smith 22659 (F, GH, us).

Cuzco: Prov. Urubamba, Machu Picchu, Mexia 8084 (F,

GH, MO). Prov. La Convention, Potrero, Tryon & Tryon

5395 (F, GH, us). Arequipa: Lomas de Atiquipa, Coro-

nado 30 (GH, us). Puno: Prov. Sandia, Sandia, Vargas

14813 (F).

2. Blechnum confluens Schlect. & Cham., Linnaea

5: 613. 1830. TYPE: Mexico, Hacienda de

Laguna, Schiede 779 (holotype, B, photo, GH;

isotype, Schiede & Deppe 779, B, photo, GH).

Blechnum triangulare Link. Hort. berol. 2: 76. 1833.

TYPE: Cultivated (holotype, B, Herb. Link, photo

GH; isotype,?, "e Mexico, Blechnum triangulare

Spr." Sprengel script., B, photo, GH). Although

Link published the name first, he says he received

it from Sprengel.

Stem small, erect to decumbent and short-creep-

ing. Leaves monomorphic. Petiole with basal scales

rather few, short, mostly lanceolate to ovate-lan-

ceolate, stem scales wholly light brown to dark

brown or often with lighter margins. Lamina pin-

natisect, the apex gradually reduced, pinnatifid,

reduced at the base to pinnae that are longer than

broad or as long as broad, the pinnae essentially

flat, the longest ca. 1—4 cm long. Fertile pinnae

with green tissue extending well beyond the at-

tachment of the indusium.

Terrestrial or rupestral, in forests, on wet banks,

among rocks, or in rock crevices, 670—2300 m,

Amazonas south to Junin.

Tropical America.

This species is more or less intermediate be-

tween B. occidentale and B. asplenioides; some

(all?) specimens might represent hybrids between

those species. The basal lobes in B. asplenioides

are fully adnate and broader than long, while in

B. confluens they are longer than broad or about

as long as broad and only partly adnate to the

rachis.

Amazonas: E of Chachapoyas, Wurdack 730 (GH). San

Martin: Monte Guayrapurima, prope Tarapoto, Spruce

4022 (GH, us). Prov. Lamas, Dist. Lamas, Belshaw 3424

(F, GH, us). Lamas, LI. Williams 6341 (F). Huanuco:

Prov. Leoncio Prado, Dist. Rupa Rupa, J. Schunke V.

5081 (GH). Between Tingo Maria and Monzon, Croat

57909 (F). La Divisoria, Gentry et al. 18864 (F). Pasco:

Prov. Oxapampa, Soukup 6810 (GH). Junin: Prov. Tar-

ma, Utcuyacu, Woytkowski 35371 (GH). Near La Mer-

ced, Killip & Smith 23994 (F).

3. Blechnum fraxineum Willd., Sp. pi. ed. 4, 5:

413. 1810. TYPE: Venezuela, Caracas, Bre-

demeyer (holotype, B, Herb. Willd. 20044,

photo, GH). Not Lomaria fraxinea Willd., op.

cit. 1810. Figure 3c.

Blechnum longifolium Willd., Sp. pi. ed. 4, 5: 413.

1810, not Cav. 1802. TYPE: Venezuela, Caripe,

Humboldt 464 (holotype, B, Herb. Willd. 20045,

photo, GH).

Blechnum graciletiaulf., Enum. fil. 158. 1824. TYPE:

"Brasilia, Otto comm." (not seen), ex char.

Stem small, erect to decumbent and short-creep-

ing. Leaves monomorphic. Petiole with basal scales

usually rather few, rather short, mostly lanceolate

to ovate-lanceolate, stem scales brown to dark

brown, sometimes with narrow lighter margins.

Lamina pinnatisect to 1 -pinnate, the apex abruptly

reduced to an apical pinna that is longer than the

longest lateral pinna, not or slightly reduced at the

base, the pinnae essentially flat. Fertile pinnae with

green tissue extending well beyond the attachment

of the indusium.

Terrestrial or rupestral, on clay banks, along

streams, and among rocks, 450-2000 m, Ama-

zonas, south to Cuzco.

Tropical America.

Walker (1985) reported a diploid and a tetra-

ploid from Trinidad. The tetraploid, which is more

dimorphic than the diploid and with additional

(6-14) pinnae, is evidently more common in Peru

than the diploid, with 2-4 pinnae. The identifi-

cation of Peru material is uncertain pending cy-

tological studies that have not been made. Blech-

num gracile is a name for the tetraploid and B.

meridense Kl. for the diploid. The type of B. fraxi-

neum is considered to be a triploid hybrid of B.

meridense and B. occidentale. The essential char-

acter of the species and its hybrids is the long

terminal segment.

Amazonas: Prov. Bagua, valley of Rio Maranon, above

Cascadas de Mayasi, Wurdack 1822 (GH. us. USM). San

Martin: San Roque, LI. Williams 7519 (F). Tarapoto,

Spruce 4026 (us). Junin: La Merced. Macbride 5500 (F.

us). Chanchamayo valley. C. Schunke 3 1 (F). / 32 (F. us).

1352 (F). Above San Ramon. Killip & Smith 24731 (us).

Ucayali: La Divisoria. Tryon & Tryon 5271 (GH. us).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

59

Cuzco: Prov. Paucartambo, Pillawata, Vargas 16729 (GH).

QUILLABAMBA, CORONADO 127 (US).

4. Blechnum asplenioides Sw., Kongl. Vetensk.

Acad. Handl. 1817: 72, t. 3, f. 3. TYPE: Bra-

zil, Villa Rica, Freyreis (holotype, s, Herb.

Swartz, photo, GH, us). Figures 3a-b.

Asplenium blechnoides Sw., Syn. fil. 76. 1806. TYPE:

Peru, evidently Nee ex Lagasca (holotype, s; pho-

to, GH, us).

Blechnum unilaterale Sw., Ges. Naturf. Freunde Ber-

lin Mag. Neuesten Entdeck. Gesammten Naturk.

4: 79. 1810, also Willd., Sp. pi. ed. 4, 5: 407.

1810. Both authors cite Asplenium blechnoides

Sw. in synonymy; hence the name is superfluous

and has the same type as Asplenium blechnoides.

Blechnum polypodioides Raddi, Opusc. Sci. 3: 294.

1819, not (Sw.) Kuhn, 1 868. TYPE (from Raddi,

PI. bras. nov. gen. 53, t. 60, f. 2. 1825): Brazil,

Corcovado, Raddi (holotype, PI?).

Blechnum blechnoides (Sw.) C. Chr., Index fil. 151.

1906, not Keyserl., 1873.

Stem small, erect to decumbent and short-creep-

ing. Leaves monomorphic. Petiole with basal scales

rather few, mostly short, lanceolate to narrowly

lanceolate, stem scales light brown to dark brown

or with a dark brown center. Lamina pinnatifid to

pinnatisect, the apex gradually reduced, pinnatifid,

the base gradually reduced to lobes that are broad-

er than long, pinnae essentially flat, the longest ca.

0.5-2.5 cm long. Fertile segments with green tissue

extending well beyond the attachment of the in-

dusium.

Terrestrial or rupestral, wet forests, secondary

forests, shaded stream banks, clay soil of road

banks, or on rocks, 300-1 800 m, Amazonas south

to Cuzco.

Tropical America.

This species is distinguished by the small basal

lobes of the lamina. It is usually a small plant,

although sometimes the leaf reaches a length of

ca. 50 cm.

Amazonas: Prov. Bagua, Rio Maranon, above Cas-

cadas de Mayasi, Wurdack 2056 (F, GH, us). Prov. Bagua,

NE of Chiriaco, Barbour 4509 (USM). San Martin: Ze-

pelacio, near Moyobamba, Klug 3358 (F, GH, us). Prov.

Lamas, Lamas, Ferreyra 4691 (GH, USM). San Roque, LI.

Williams 7144 (F). Loreto: Pampayacu, Klug 3237 (F,

GH, us). Huanuco: Villcabamba, Macbride 4991 (F, GH,

us). Tingo Maria, Tryon & Tryon 5259 (F, GH, us). Pasco:

Dos de Mayo, Pichis Trail, Killip & Smith 25861 (F, GH).

Junin: La Merced, Soukup 2374 (GH). Near La Merced,

Killip & Smith 23959 (F, us). Chanchamayo valley, C.

Schunke2, 26, 182, 777, 1374(p). Ucayali: La Divisoria,

Allard 22174 (GH). La Divisoria, Gentry et al. 18832 (F,

USM). Cuzco: Prov. Quispicanchis, entre Quinze Mil y

San Lorenzo, Vargas 11761 (GH). Prov. La Convention,

valley of Rio Vilcanota, Mexia 8052 (F, GH, us). Prov.

Paucartambo, Kosnipata valley, Plowman & Davis 5114

(GH). Madre de Dios: 39 km SW of Puerto Maldmado,

Smith et al. 1 1 28 (us).

5. Blechnum lanceola Sw., Kongl. Vetensk. Acad.

Handl. 1 8 1 7: 7 1 , t. 3, f. 2. TYPE: Brazil, Villa

Rica, Freyreis (holotype, s, Herb. Swartz,

photo, GH, us).

Stem small, erect to decumbent and very short-

creeping. Leaves monomorphic. Petiole with basal

scales few to nearly lacking, short, mostly lanceo-

late to ovate-lanceolate, stem scales light to dark

brown, or with lighter narrow margins. Lamina

entire or rarely with 1 or 2 small lobes at the base,

mostly very narrowly elliptical, essentially flat.

Fertile lamina with green tissue extending well be-

yond the attachment of the indusium.

A single collection from San Martin, Peru.

Peru south to Argentina and east to Brazil.

This species is represented by small plants 4—

1 5 cm, or occasionally to 30 cm, tall. The entire

lamina is distinctive, although rarely there may be

one or two lobes at the base. It is evidently rare

in the Andes and more common in southeastern

Brazil.

San Martin: Monte Campana, prope Tarapoto, Spruce

4672 (GH, us).

6. Blechnum serrulatum Rich., Actes. Soc. Hist.

Nat. Paris 1: 114. 1792. TYPE: "Cayenne"

(French Guiana), Le Blond (holotype, p, Herb.

Richard, photo, GH, us).

Stem widely creeping (in deep sandy soil), its

ascending branches bearing the leaves. Leaves

monomorphic. Petiole with few or no scales at the

base, stem scales wholly dark brown or with lighter

margins or small and light brown only. Lamina

1 -pinnate, with an entire apical segment, not or

hardly reduced at the base, pinnae essentially flat,

articulate to the rachis, more or less serrulate, the

largest ca. 5-1 5 cm long. Fertile pinnae with green

tissue extending well beyond the attachment of the

indusium.

Sandy soil in secondary vegetation, 1 80 m, Lo-

reto.

Tropical America, south to Peru and north-

eastern Argentina.

60

FIELDIANA: BOTANY

The species has monomorphic leaves and differs

from all species in South America in having ar-

ticulate pinnae. In sandy deep soil, the main stem

is deeply subterranean and has ascending branches

that bear the leaves.

Loreto: Jenaro Herrera, Mejia & Van der Well 221

(USM).

7. Blechnum brasiliense Desv., Ges. Naturf.

Freunde Berlin Mag. Neuesten Entdeck. Ge-

sammten Naturk. 5: 330. 1811. TYPE: Brazil,

Dombey (holotype, P, Herb. Jussieu 1390,

photo, GH, us).

Blechnum nigrosquamatum Gilbert, Bull. Torrey Bot.

Club 24: 258. 1897. TYPE: Bolivia, Colapampa,

Bang 2314 (holotype, not located; isotype, GH!).

Stem subarborescent to arborescent, stout, to 2

m tall. Leaves monomorphic. Petiole with scales

at the base many, mostly 10-15 mm long, acicular,

blackish or with a blackish center, stem scales dark

brown to blackish or with a dark center and lighter

(to brown) margins. Lamina pinnatifid to pinnati-

sect, the apex gradually reduced, pinnatifid, or with

a short entire tip, gradually reduced, pinnatifid, or

with a short entire tip, gradually reduced at the

base to small pinnae or lobes, pinnae essentially

flat, the largest ca. (7-)15-20(-25) cm long. Fertile

pinnae with green tissue extending well beyond the

attachment of the indusium.

Terrestrial, mountain forests, 1200-1600 m, San

Martin.

Guatemala; tropical South America.

The petiole is usually short to nearly absent. The

long, narrow scales at the base of the petiole com-

bined with the monomorphic leaves are distinc-

tive features of this species.

San Martin: Zepelacio, near Moyobamba, Klug 3433

(F, GH, us). Murillo (1968) also cites: Monte Campana,

Tarapoto, Spruce 4673 (BR).

8. Blechnum lehmannii Hieron., Bot. Jahrb. Syst.

34: 473. 1904. TYPE: Colombia, Rio Tim-

biqui, Lehmann 8928 (holotype, B; isotype,

us!).

Struthiopteris maxonii Broadh., Bull. Torrey Bot. Club

39: 268. 1912. TYPE: Panama, Chiriqui, Cerro

de la Horqueta, Maxon 5415 (holotype, us; iso-

type, GH!, us!).

Blechnum maxonii (Broadh.) C. Chr., Index fil. suppl.

16. 1913.

Stem more or less erect, rather slender. Leaves

dimorphic. Petiole with basal scales few to rather

numerous, brownish, mostly large, concolorous,

lanceolate-ovate or broader, stem scales acumi-

nate, brown to dark brown, to blackish (some),

concolorous. Sterile lamina pinnatisect, the apex

gradually reduced or with a short to long entire

tip, the base gradually reduced to lobes, pinnae

curved-ascending, fully adnate to the rachis, the

margins flat to somewhat involute, distant from

the costa. Fertile pinnae with green tissue hardly

extending beyond the attachment of the indusium.

Terrestrial, cloud forests, 1750 m, Cuzco.

Southern Mexico; Central America; Venezuela

and Colombia, south to Peru.

The stem may be erect and to 40 cm tall; the

sterile lamina is 3-10 cm, usually 4-6 cm broad.

This is closely related to, perhaps conspecific

with, B. I'herminieri (Kunze) Mett. of the West

Indies. That species has the petiole dark, and the

lamina is abruptly reduced basally in contrast to

B. lehmannii, which has a light-colored petiole and

a lamina that is gradually reduced at the base.

Lopez & Sagdstegui 5297 (GH, HUT), from Prov.

Chota, Cajamarca, seems to be this species, but

the petioles are mostly dark, rather than straw-

colored as in typical B. lehmannii. The collection

may represent an intermediate between the two

taxa.

Cuzco: Prov. La Convencion, Cordillera Vilcabamba,

Knox's Cascade, Dudley 10463 (GH). Dudley 1 0489 from

the same place (GH) is a sheet of juvenile plants, most

probably of this species.

9. Blechnum andinum (Baker) C. Chr. Ind. fil. 1 50.

1905.

Lomaria andina Baker, Syn. fil. ed. 2: 482. 1883.

TYPE: Andes, "Unduari" (Unduavi), Bolivia,

10,000 ft, Pearce (holotype, K; photo, GH, us).

Blechnum subtile Rosenst., Repert. Spec. Nov. Regni

Veg. 11: 54. 1912. TYPE: Unduavi, Bolivia.

Buchlien 2779 (holotype, not located; isotype, us!).

Stem small, erect, stoloniferous, the stolons very

slender and threadlike. Leaves small, dimorphic.

The fertile and sterile about the same length. Pet-

iole lacking scales or with a few, brownish, con-

colorous ones, stem scales more or less small,

brownish, sometimes with lighter margins. Sterile

lamina 1 -pinnate, the apex and base gradually re-

duced, the base reduced to small pinnae, the pin-

nae obtuse, narrowly to broadly adnate. Fertile

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

61

pinnae with green tissue hardly extending beyond

the attachment of the indusium.

Wet forests, on trees and in a cave, 3050 m,

Cuzco.

Peru; Bolivia; and southern Brazil.

This is a very small species, the leaves usually

6-12 cm long.

Cuzco: Michihianuca, Bues 973 (us). Huadquina, Bues

980 (us).

10. Blechnum penna-marina (Poiret) Kuhn, Fil.

Afr. 92. 1868.

Polypodium penna-marina Poiret in Lam., Encycl. 5:

520. 1804. TYPE: Straits of Magellan, Commer-

son (holotype, Herb. Desfontaine, p; photo, GH).

Stem small, more or less erect, freely stolonif-

erous and sometimes forming large clumps of

plants. Leaves small, dimorphic, the sterile shorter

and spreading, the fertile longer and erect. Petiole

with a few brownish, concolorous scales, stem

scales brown, more or less reddish brown, to light

brown or dark brown, concolorous, more or less

acute to acuminate. Sterile lamina mostly 5-10

(-15) cm long, narrow, pinnatisect, gradually re-

duced to the apex, at the base reduced to small

lobes. Fertile pinnae with green tissue hardly ex-

tending beyond the attachment of the indusium.

Terrestrial, 3390 m, Cuzco.

Peru and southeastern Brazil; to southernmost

Chile and Argentina; circumantarctic islands;

southeastern Australia; New Zealand. Mettenius

(Fil. Lech. 2: 15. 1859.) reported this species for

Peru as Blechnum alpinum var. elongatum (Puno,

Agapata, Lechler). I have not seen this specimen,

and it is possible that it is some other species. In

any event, the following is a recent collection.

Cuzco: Prov. Paucartambo, Cerro Macho Cruz, Manu

National Park, B. Leon et al. 2757 (cuz, USM).

11. Blechnum cordatum (Desv.) Hieron., Hed-

wigia47: 239. 1908.

Lomaria cordata Desv., Ges. Naturf. Freunde Berlin

Mag. Neuesten Entdeck. Gesammten Naturk. 5:

330. 1811. TYPE: omitted in the publication. Ho-

lotype evidently "Perou-Concepcion," Dombey,

p, Herb. Jussieu 1368; photo, GH, us.

Lomaria chilensisKau\f.,Enum.fi\. 154. 1824. TYPE:

Chile, Chamisso (holotype, LE?)

Lomaria ornifolia Presl, Reliq. haenk. 51.1 825. TYPE:

"Hab. in Peruviae?" Placed as a synonym of

Blechnum cordatum by Hieron., Hedwigia 47: 239.

1908.

Lomaria serrulosa Desv., Mem. Soc. Linn. Paris 6:

290. 1827. TYPE: "Peruvia" (holotype, P; photo,

GH, us).

Lomaria arborescens Klotzsch, Linnaea 20: 347. 1844.

SYNTYPES (designated by Hieron., Hedwigia 47:

240. 1908): Venezuela, Merida, Moritz 299 (B)

and Venezuela, Karsten Coll. II, 25 (B). Isosyn-

type, Moritz 299, BM; photo, GH; frag., us!.

Blechnum chilense (Kaulf.) Men., Fil. lechl. 1: 14.

1856.

Blechnum ornifolium (Presl) Ettingsh., Denkschr.

Kaiserl. Akad. Wiss. Wien. 23: 61. 1864.

Blechnum arborescens (Klotzsch) Hieron., Hedwigia

47: 239. 1908.

Blechnum peruvianum Hieron., Hedwigia 47: 245, /.

10. f. 19. 1908. TYPE: Peru, Rio Huallaga, Mo-

yobamba, Stiibel 1 105 (holotype, B; photo, us).

Stem erect and subarborescent to decumbent

and short-creeping, moderately to very stout.

Leaves dimorphic. Petiole with basal scales small

to rather large, few to many, concolorous and

mostly brown, linear to lanceolate to lanceolate-

ovate or broader, stem scales broad, thin, concol-

orous, light brown or reddish brown. Sterile lam-

ina 1 -pinnate, the apex abruptly reduced to a ter-

minal segment, the base not or only slightly

reduced, pinnae sessile to definitely stalked, strict-

ly patent to somewhat ascending, cuneate to

broadly rounded to cordate, the margins flat or

slightly involute, distant from the costa. Fertile

pinnae with green tissue extending hardly beyond

the attachment of the indusium.

Terrestrial, wet montane forests, cloud forests,

thickets, along streams, on clay road banks, and

on old landslides, 700-3300 m, Piura south to

Cuzco.

Venezuela and Colombia, south to Peru; east-

ward to southeastern Brazil; Chile; Juan Fernan-

dez Islands, West Falkland Islands; entire range

not certain.

This is a variable species, primarily of the Andes

from Venezuela and Colombia, south to Chile.

The leaves are often large, to 1 m or more long,

the apex is abruptly reduced to a large terminal

segment, and the scales of the costae and rachis

are mostly rather appressed and concolorous,

whitish to light brown.

The species is variable in many characters— for

example, aerophores are absent, or present on some

pinnae and then low to conical, the rachis may be

smooth or strongly roughened and it varies from

straw-colored to dark reddish. There seems to be

62

FIELDIANA: BOTANY

no basis, at least in Peru, for the recognition of

more than one taxon. A. R. Smith (1985) also

treated Blechnum cordatum as a wide-ranging spe-

cies with many synonyms. As in the case of other

variable species, related ones recognized to the

north of South America may be distinctive. A

careful revision of this complex is much needed.

The photograph of the type of Lomaria cordata

clearly shows that the label reads "Perou-Concep-

cion," rather than "Peru, Concepcion," as indi-

cated by Morton on the covering label. Dombey

never collected at or near a Concepcion in Peru,

but he did travel from Lima to Concepcion, Chile,

where he evidently collected the type specimen.

Piura: Prov. Ayabaca, Sagdstegui & Cabanillas 8702

(F, HUT). Lambayeque: 4 km NW of Incahuasi, Dillon &

Skill man 4155 (F, GH, USM). Arriba de Incahuasi, Sa-

gdstegui el al. 12893 (F, GH). Amazonas: Mountain E of

Balsas, Osgood & Anderson 72 (F). San Martin: Prov.

San Martin, Dist. Tarapoto, Hickok 636 (GH). Prov. La-

mas, Cerro Blanco, J. Schunke V. 6259 (F). Prov. Maris-

cal Caceres, Rio Abiseo National Park, Young 1768 (USM).

Loreto: Pumayacu, King 3243 (GH, us). Anacash: Above

Yungay, Correll & Smith P964(GH). Prov. Huaylas, Lopez

et al. 8378 (F, HUT). Prov. Yungay, Laguna Llanganuco,

Mostacero et al. 1402 (F, GH). Huanuco: Carpish, Tryon

& Tryon 5317 (GH). Prov. Huanuco, Dist. Churubamba,

Mexia 8174 (F, GH, us). Mito, Macbride & Feat herst one

1522 (F, us). Lima: Prov. Canta, 4 km from Huaman-

tanga, Sounders 1254 (GH). Pasco: Quillasu, Soukup 3289

(or 3284) (F, GH). Prov. Oxapampa, road Oxapampa-

Paucartambo, Smith & Pretel 1631 (F). Prov. Oxapampa,

"La Esperanza," Leon 613 (USM). Junin: Villa Amoretti,

Kunkel 655 (GH). Chanchamayo valley, C. Schunke 162

(F). La Merced, Soukup 1087 (F). Huancavelica: Prov.

Tayacaja, cerca a Tocas, Tovar 2044 (GH, USM). Aya-

cucho: Prov. La Mar, 20-25 km from Tambo, Dudley

1 1942 (GH). Cuzco: Prov. La Convencion, San Luis, Var-

gas 19858 (GH). Prov. La Convencion, Cordillera Vil-

cabamba, Dudley 10329 (GH), 10584 (GH), 70609 (GH).

5 km N of Aguas Calientes, Solomon 3169 (F, USM).

12. Blechnum divergens (Kunze) Mett., Ann. Sci.

Nat. Hot. 5, 2: 225. 1864.

Lomaria divergens Kunze, Linnaea 9: 57. 1 834. TYPE:

Peru, Dept. Huanuco, Pampayaco, Poeppig Diar.

1141 (not located).

Lomaria pteropus Kunze, Farnkr. 1: 97. 1842, nom.

nov. for Acrostichum heterophyllum Raddi, PI.

bras. nov. gen. 5, /. 17. 1825. TYPE: Brazil, Man-

diocca, Raddi (holotype, FI?), not Lomaria het-

erophylla Desv., 1811, nor Blechnum hetero-

phyllum (Desv.) Schlect., Adumbr. pi. 37. 1827.

Blechnum pteropus (Kunze) Mett., Fil. hort. bot. Lips.

61. 1856.

Stem erect to rather long-creeping, moderately

stout. Leaves dimorphic. Petiole with basal scales

few, rather large, concolorous, lanceolate to lan-

ceolate-ovate, stem scales broad, dark brown to

blackish, concolorous. Sterile lamina pinnatisect,

the apex rather abruptly reduced to a short or long,

entire tip, the base abruptly reduced, sometimes

to a wing or to lobes that may be distant and

inconspicuous, pinnae patent to somewhat as-

cending, fully adnate or rarely partly adnate, the

margins flat, distant from the costa. Fertile pinnae

with green tissue extending hardly beyond the at-

tachment of the indusium.

Terrestrial or rarely epiphytic, cloud forests,

dense, wet forests, 1500-2400 m, Cajamarca,

Huanuco, Junin, and Cuzco.

Tropical America.

In South America, specimens of Blechnum di-

vergens and B. binervatum ssp. acutum are not

always readily distinguished, due to variation in

both taxa. Blechnum divergens is a terrestrial spe-

cies, rarely epiphytic in very wet cloud forest, and

the stem has broad, concolorous scales. The sterile

lamina is broad, 12-30 cm, and is abruptly re-

duced at the base to a petiole that is either alate

or has lobes, often small and inconspicuous, that

are much broader than long. The basal or proximal

pinnae are usually not or not much decurrent and

hardly if at all narrowed toward the adnate base.

Blechnum binervatum ssp. acutum often has a

broad sterile lamina that is abruptly reduced at

the base, and the basal or proximal pinnae are

usually strongly decurrent or narrowed to the ex-

panded and adnate base.

Cajamarca: Prov. Cutervo, Parque Nacional de Cu-

tervo, Diaz & Osores 259911 (MO). Huanuco: SW slope

of Rio Llullapichis watershed, ascent of Cerros del Sira,

Dudley 13407B (GH). Junin: Chanchamayo valley, C.

Schunke 11, 159, 160, 1382 (F). Prov. Chanchamayo,

Rio Rondayacu, Smith & Bokor 2104 (F). Cuzco: Prov.

La Convencion, Quellouno to Chirumbia, Vargas 11396

(GH). Prov. Paucartambo, between Pillawata and Patria,

Plowman & Davis 4935 (GH, us).

1 3. Blechnum binervatum (Poiret) Morton & Lell.,

Amer. Fern J. 57: 67. 1967.

Polypodium binervatum Poiret in Lam., Encycl. 5:521.

1 804. TYPE: collector and locality not known but

probably in the Lesser Antilles (holotype, p, Herb.

Jussieu 1365; photo, GH).

Stem scandent, somewhat slender to rather stout.

Leaves dimorphic. Petiole with basal scales few or

numerous, short to moderately long, rather lan-

ceolate, mostly bicolorous (in Peru) with a black-

ish center and lighter margins, or concolorous (ssp.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

63

binervatum). Sterile lamina pinnatisect or

1 -pinnate proximally, the apex gradually or

abruptly reduced to an entire tip, the base some-

what to strongly reduced to lobes, pinnae patent

to ascending, mostly fully adnate, the margins flat

or nearly so, distant from the costa. Fertile pinnae

with green tissue extending hardly or only slightly

beyond the attachment of the indusium.

This is a variable species, with three subspecies

that formerly were usually recognized as species.

The ssp. binervatum, with concolorous stem scales,

is confined to the Lesser Antilles and Puerto Rico.

This is the only scandent-epiphytic species in

Peru. The stem may be rooted in the ground and

climbing on tree trunks, especially on Cyathea-

ceae, or it may (probably) be wholly epiphytic;

very rarely the plant is rupestral, or juvenile plants

may be terrestrial.

Southern Mexico and Central America; Antilles;

Venezuela to Colombia; south to Bolivia; south-

eastern Brazil.

Key to Subspecies of Blechnum binervatum

The following key provides the typical charac-

ters of the subspecies; these intergrade, and not all

specimens have the characteristic complement of

characters. For example, Peyton & King 1420,

Dept. Cuzco, at MO is nearly ssp. fragile, while at

GH it is more like ssp. acutum.

a. Sterile lamina 3-10 cm broad, gradually reduced at the base; pinnae usually close and the sinus

narrow; stem scales toothed, often freely so, especially near the apex of the stem 1 3a. ssp. fragile

a. Sterile lamina 10-25 cm broad, more or less abruptly reduced at the base; pinnae often spaced and

the sinus broad; stem scales with few and small teeth or none 1 3b. ssp. acutum

13a. Blechnum binervatum ssp. fragile (Liebm.)

Tryon & Stolze, comb. nov. Figures 3d-f.

Lomaria fragile Liebm., Kongel. Dankse Vid. Selsk.

Skr. Naturvidensk. Afd. 5, 1: 232. 1849. TYPE:

Mexico, Barranca de Huitamalco, Liebmann (Fl.

Mex. 773) (lectotype, selected by A. R. Smith,

1981): C folio, frag., us!; isolectotype, Barranca

de Huitamalco, L, photo, GH).

Blechnum fragile (Liebm.) Morton & Lell., Amer. Fern

J. 57: 68. 1967.

OsmundapolypodioidesSv/., Prod. 127. 1788. TYPE:

Jamaica, Swartz (holotype, s, photo, us; isotype,

BM, photo, GH).

Blechnum polypodioides(S'w.) Kuhn, Fil. afr. 92. 1 868,

not Raddi, 1819.

Other synonyms, mostly not currently used, are

in the reference for Blechnum fragile.

In dense rain forests and cloud forests, 1550-

3300 m, Cajamarca south to Cuzco.

Southern Mexico and Central America; Greater

Antilles; Venezuela and Colombia, south to Bo-

livia; southeastern Brazil.

Cajamarca: Prov. Cutervo, Parque Nacional de Cu-

tervo, Diaz & Osores 2596 (MO). Amazonas: Upper slopes

of Puma-urea, ESE of Chachapoyas, Wurdack 695 (GH).

Prov. Bongara, Shillac, Smith & Vasquez 4898 (MO, USM).

San Martin: Rio Abiseo National Park, Young & Watson

3350 (USM). Huanuco: SW slope of Rio Llullapichis wa-

tershed, ascent of Cerros del Sira, Dudley 13284 (GH).

Muna, Macbride 4292 (F, GH). Pasco: Oxapampa, Sou-

kup 2337 (F, GH). Rio San Alberto, Foster et al. 10316

(USM). Junin: Pichita Caluga, Walden 37, 74 (GH). Cuzco:

Prov. La Convencion, Huayopata, Peyton & King 1420

(MO).

13b. Blechnum binervatum ssp. acutum (Desv.)

Tryon & Stolze, comb. nov.

Lomaria angustifolia HBK., Nov. gen. sp. 1 : 18. 1816.

TYPE: Ecuador, Villa de Ibarra, Humboldt &

Bonpland (B or p?, not seen).

Lomaria acuta Desv., Mem. Soc. Linn. Paris 6: 290.

1827. LECTOTYPE: Dombey (designated by

Weatherby, Contr. Gray Herb. 114: 26. 1936):

"Habitat in Peruvia" (holotype, P; photo, GH).

Lomaria cuspidata Kunze, Linnaea 9: 59. 1 834. TYPE:

Peru, Pampayaco, Poeppig, Diar. 1 120 (isotype,

B?, not seen).

Lomaria meridensis Klotzsch, Linnaea 20: 345. 1847.

TYPE: Venezuela, Merida, Moritz 297 (holotype,

B, not seen).

Lomaria ensiformis Liebm., Kongel. Danske Vidensk.

Selsk. Skr. Naturvidensk. Afd. 5, 1: 234. 1849.

TYPE (selected by A. R. Smith, 1981): Mexico,

Oaxaca, Distr. Chinantla, Teotalcingo, Liebmann

PL Mex. 2342 (lectotype, c, folio, sheet with both

fertile and sterile leaves, frag., us!).

Blechnum meridense (Klotzsch) Mett., Fil. hort. lips.

61. 1856, based on Lomaria meridensis, not

Blechnum meridense Klotzsch, 1847.

Blechnum acutum (Desv.) Mett., Ann. Sci. Nat. Bot.

Ser. 5, 2: 225. 1864.

Blechnum angustifolium (HBK.) Hieron., Bot. Jahrb.

Syst. 34: 472. 1904, not Willd., 1810.

Blechnum ensiforme (Liebm.) C. Chr., Index fil. 153.

1905.

Blechnum kunthianum C. Chr., Index fil. suppl. 16:

1913, nom. nov. for Lomaria angustifolia HBK.,

not Blechnum angustifolium Willd., 1810.

64

FIELDIANA: BOTANY

Southern Mexico and Central America; Vene-

zuela and Colombia, south to Bolivia; southeast-

ern Brazil.

In humid forests, in rain forests, and in wet

cloud forests, 1000-2700 m, Amazonas south to

Cuzco.

In South America this subspecies is especially

variable in the characters of the sterile lamina.

Usually the distal pinnae are fully adnate and

broadest at the base, but neither excurrent nor

decurrent, and the pinnae are close with a narrow

sinus between them. Proximal pinnae may be sim-

ilar or they may be somewhat spaced and excur-

rently and decurrently broadened at the base. These

characters, however, vary to the extreme (Lomaria

acuta Desv.), with few and widely spaced pinnae

that are narrowed toward the base where they are

fully adnate.

Amazonas: Prov. Bagua, 12 km E of La Peca, Barbour

2507 (F, MO). San Martin: Prov. Rioja, Pedro Ruiz-Mo-

yobamba, D. Smith 4451 (F). Prov. Rioja, Pedro Ruiz-

Moyobamba, Smith & Vasquez 4709 (MO, USM). Huanu-

co: SW slope of Rio Llullapichis watershed, ascent of

Cerros del Sira, Dudley 13253 (GH). Pasco: Prov. Oxa-

pampa, Oxapampa, Soukup 2339 (GH). Prov. Oxapam-

pa, Rio El Tunqui, Smith & Alban 5524 (F). Prov. Oxa-

pampa, SW of Oxapampa, Foster 7624 (USM). Junin:

Prov. Chanchamayo, Rio Rondayacu, Smith et al. 2616

(F, USM). Ucayali: Prov. Leoncio Prado, La Divisoria,

Plowman & Schunke 7418 (F). Cuzco: Prov. La Con-

vention, 3 km from Incatambo, Peyton & Peyton 851

(GH, MO). Quillabamba, Santa Teresa, Peyton & Peyton

1273 (GH, MO); Prov. Paucartambo, Parque Nacional

Manu, Leon 2194 (F).

14. Blechnum stipitellatum (Sodiro) C. Chr., In-

dex fil. 160. 1905.

Lomaria stipitellata Sodiro, Recens. crypt, vase. Quit.

29. 1883. SYNTYPES: Ecuador, Corazon y Ata-

cazo, 2500-3400 m. Sodiro (syntype or isosyn-

type, Corazon, Sodiro, p; photo, GH).

Blechnum rubicundum Hieron., Hedwigia 47. 242, /.

4,f. 16. 1908. TYPE: Colombia, Cumbal to La

Ceja de Mayasquer, Stiibel 347 (holotype, B; pho-

to, us).

Stem moderately to rather stout, erect to de-

cumbent and very short-creeping. Leaves dimor-

phic. Petiole with basal scales mostly large, mostly

concolorous, sometimes bicolorous, brown, few to

many, linear to lanceolate to lanceolate-ovate, stem

scales large, brown to dark brown or reddish brown

often with lighter (broad) margins, acute more or

less thin, sometimes concolorous. Sterile lamina

1 -pinnate, the apex gradually reduced to an entire

tip, the base slightly or not reduced, pinnae usually

patent to slightly ascending, sessile to short-stalked,

more or less truncate, the margins flat to somewhat

involute, distant from the costa. Fertile pinnae

with green tissue extending only slightly beyond

the attachment of the indusium.

Terrestrial, in wet forests and among rocks,

2400-3700 m, Amazonas, south to Puno.

Colombia, south to Bolivia.

This seems to be a distinctive species, but it is

little known and its variation is uncertain.

Many or most of the scales on the costa and

rachis abaxially are more or less spreading, light

brown to dark brown and then sometimes bicolor-

ous. The scales, or at least some of them, are den-

ticulate to ciliate on the margins. The lamina has

many, usually 20—40 pairs of close, patent pinnae.

At least when dried, the pinnae are often promi-

nently striate adaxially— the tissue between the

veins being depressed.

This species seems close to some variations of

Blechnum cordatum, but the expanded sterile pin-

nae suggest it may be a shade form of Blechnum

loxense.

Amazonas: Prov. Bagua, Cordillera Colan, Barbour

3467 (MO). Huanuco: Between Huanacoand Pampayacu,

Kanehira 144 (GH, USM). Huanuco-Tingo Maria road,

Gentry et al. 33851 (F). Pasco: Prov. Oxapampa, Cor-

dillera San Gutardo, Leon et al. 517 (USM). Junin: Prov.

Satipo, Dist. Pampa Hermosa, Sounders 1069 (GH). Cuz-

co: Prov. Urubamba, above Palca, Peyton & Peyton 767

(GH, MO). Puno: Agapata, Lechler 2025, in part (GH).

15. Blechnum loxense (HBK.) Salomon, No-

mencl. Gefaskrypt. 117. 1883.

Lomaria loxensis HBK., Nov. gen. sp. 1: 18. 1816.

TYPE: Ecuador, "Andibus Peruviae inter Gon-

zanama et Loxa," Humboldt & Bonpland (holo-

type, P; photo, GH).

Lomaria squamulosa Desv., Mem. Soc. Linn. Paris

6: 290. 1827. "Habitat in Peruvia," Dombey (F,

photo, us).

Lomaria stenophylla Klotzsch, Linnaea 20: 346. 1847.

TYPE: of alpha. Peru, Dombey (B; photo, us).

This and also beta are both B. loxense according

to Hieronymus, Hedwigia 47: 241. 1908. Not

Blechnum stenophyllum Presl, 1851.

Blechnum stenophyllum (Klotzsch) Men., Fil. hort.

bot. lips. 64. 1856. Not Blechnum stenophvllum

Presl.

Blechnum squamulosum (Desv.) Men., Ann. Sci. Nat.

Bot. Ser. 5,2: 225. 1864.

Stem stout to massive, erect to somewhat de-

cumbent and very short-creeping. Leaves mono-

morphic. Petiole with basal scales usually numer-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

65

ous, large, brown and concolorous, stem scales

mostly broad, light brown to brown to more or

less reddish brown with or without a darker center.

Sterile lamina 1 -pinnate, the apex abruptly re-

duced to a terminal segment, the base hardly or

not reduced, the pinnae ascending, usually cor-

date, the margins strongly involute, usually reach-

ing or nearly reaching the costa on the abaxial side,

the tip usually strongly involute. Fertile pinnae

with green tissue extending only slightly beyond

the attachment of the indusium.

In open, often wet, sites, usually in the puna,

grasslands, steep roadside banks, among rocks or

in the crevices of rocks, 1750-4100 m, Lamba-

yeque south to Puno.

Costa Rica and Panama; Venezuela and Colom-

bia, south to Bolivia.

The stem is stout but only rarely (to ca. 2 m tall)

arborescent. It is usually soft, mucilaginous and

green inside, especially at the apex. Except for B.

auratum var. columbiense, this species is distinc-

tive among those of Peru in having very narrow

sterile pinnae as well as fertile ones.

Lambayeque: Dist. Incahuasi, Sagdstegui et al. 12781

(F, GH, HUT). Cajamarca: Prov. Hualgayoc, Carmona.

Soukup 501 1 (us). Amazonas: Paso de Calla-Calla, San-

chez 87 (GH). San Martin: Prov. Mariscal Caceres, Rio

Abiseo National Park, Young & Watson 3499 (USM),

4400 (F, USM). La Libertad: Prov. Bolivar, Laguna de

los Ichus, Lopez & Sagdstegui 3237 (GH). Prov. Santiago

de Chuco, Dist. Mollepata, Sounders 875 (GH). Prov.

Santiago de Chuco, Shoreyo-Trujillo, Smith 2344 (F).

Ancash: NW slope of Nevada de Huascaran, Correll &

Smith P958 (GH). Prov. Pallasca, Consuzo, Lopez 2394

(GH). Huascaran National Park, D. Smith 10613, Smith

et al. 10380 (F), 77627 (F, HUT), 12395, 12457 (F). Huan-

uco: Mito, Bryan 203 (F), Macbride & Featherstone 1790

(F, us), 2779 (F, us). Junin: Huancayo, Kunkel 425 (GH).

Huancavelica: Prov. Tayacaja, entre Colcabamba y Pau-

cartambo, Tovar 2005 (GH). Ayacucho: Between Huanta

and Rio Apurimac, Killip & Smith 23258 (F, us). Apu-

rimac: Prov. Andahuaylas, Dist. Chincheros, Saunders

736 (GH). Cuzco: Prov. La Convencion, Dist. Vilcabam-

ba, Davis et al. 1217 (F, GH). Puno: Agapata, Lechler

2025, in part (GH). Near Puno, Soukup 218 (F).

16. Blechnum sprucei C. Chr., Index fil. 160. 1905,

nom. nov. for Lomaria caudata Baker, not

Blechnum caudatum Cav., 1802.

Lomaria caudata Baker, in Hooker & Baker, Syn. fil.

1 79. 1 967. TYPE: Andes of Ecuador, Spruce 5329

(holotype, K, photo, Svensk Bot. Tidskr. 420.

1954).

Stem rather stout, nearly erect to decumbent

and very short-creeping. Leaves dimorphic. Pet-

iole with basal scales few to many, rather large,

concolorous, lanceolate to lanceolate-ovate or

broader, stem scales light to dark brown often with

lighter margins. Sterile lamina 1 -pinnate, the apex

gradually reduced, much prolonged with small

pinnae and with 1 or more proliferous buds (al-

ways?), the base gradually reduced, pinnae strictly

patent, sessile, truncate or often somewhat biau-

riculate, the margins flat, distant from the costa.

Fertile pinnae with green tissue extending hardly

or only slightly beyond the attachment of the in-

dusium.

Undoubtedly in Peru.

Costa Rica; Ecuador; Peru south to northwest

Argentina and east to southeast Brazil.

The much prolonged apex of the sterile lamina

makes this a very distinctive species.

17. Blechnum schomburgkii (Klotzsch) C. Chr.,

Index fil. 157. 1905.

Lomaria schomburgkii Klotzsch, Linnaea 20: 346.

1847. TYPE: "British Guiana," Schomburgk 1162

(B).

Stem stout, arborescent to 3 m tall, to decum-

bent. Leaves dimorphic. Petiole with basal scales

very numerous, long (to 3.5 cm or more), acicular,

the base usually broader, curved, mostly bicolor-

ous with a blackish or dark center and lighter,

brown margins, stem scales dark brown or with

lighter margins or light brown or brown. Sterile

lamina 1 -pinnate or pinnatisect distally, the apex

abruptly reduced to a large, entire segment, the

base hardly to somewhat reduced, the pinnae pa-

tent to somewhat ascending, sessile to short-stalked

or partly adnate distally, cuneate, truncate, broad-

ly rounded to cordate, the margins flat to involute,

distant from the costa. Fertile pinnae with green

tissue extending hardly or only slightly beyond the

attachment of the indusium.

Usually terrestrial, rarely rupestral, cloud forests

and elfin forests, or in open sites, seepage slopes,

grassy hillsides, or at the edge of rocks, 1 450-3400

m, Cajamarca south to Cuzco.

Costa Rica; Venezuela (Roraima) west to Co-

lombia, south to Bolivia; southeastern Brazil.

The leaves are often few and the pinnae are acute

to long-acuminate, and the tip is flat or nearly so.

Cajamarca: Namora, Correll & Smith P897 (GH).

Amazonas: Prov. Chachapoyas, entre Leimebamba y

Balsas, Lopez et al. 4446 (GH). Prov. Bongara, 3 km S

66

FIELDIANA: BOTANY

of Pomacocha. Wurdack 998 (F. GH). La Libertad: Prov.

Bolivar, Chomparen, Lopez & Sagdstegui 3207 (GH).

Huanuco: SW slope of Rio Llullapichis watershed, ascent

of Cerros del Sira. Dudlev 13417 A (GH). 13467 (GH),

13469 (GH). Mito. Macbride 3408 (F). Cuzco: Prov. La

Convencion. Cordillera Vilcabamba, Dudlev 10975 (GH),

11118(GH), 11135(GH, MO).

18. Blechnum auratum (Fee) Tryon & Stolze,

comb. nov.

Stem arborescent, often to ca. 1 m tall, some-

times multicipetal, stout. Leaves dimorphic or

monomorphic. Petiole with basal scales usually

many, to 3.5 cm long, curved, acicular beyond the

usually broader base, bicolorous with a blackish

center and lighter margins, or concolorous, brown,

stem scales long, narrow, dark brown to blackish

or center so but narrow margins lighter. Sterile

lamina 1 -pinnate or pinnatisect distally, the apex

abruptly reduced to a large, entire terminal seg-

ment, the base not or hardly reduced, to gradually

reduced to rather small pinnae, pinnae usually

strongly ascending, a few distal ones partly to fully

adnate, others sessile to short-stalked, cuneate to

broadly rounded to cordate. Fertile pinnae with

green tissue extending hardly or only slightly be-

yond the attachment of the indusium.

Costa Rica; Venezuela and Colombia; south to

Bolivia.

The stem is arborescent with a dense crown of

stiffly straight leaves, the pinnae are strongly as-

cending to rarely only slightly so, and their apex

is more or less obtuse, with the tip involute or

rarely only slightly involute.

Key to Subspecies of Blechnum auratum

a. Leaves dimorphic, the sterile pinnae broader than the fertile ones; margins of the sterile pinnae

involute but not to the costa abaxially ssp. auratum

a. Leaves monomorphic, the fertile and sterile pinnae of essentially the same width; margins of the

sterile pinnae very strongly involute, reaching to or nearly to the costa abaxially . . . ssp. columbiense

1 8a. Blechnum auratum ssp. auratum.

Lomaria aurata Fee. Mem. foug. 8: 71. 1857. TYPE:

Colombia. Ocana. Schlim 394 (holotype, p?).

Blechnum buchtienii Rosenst., Repert. Spec. Nov.

Regni Veg. 5: 231. 1908. TYPE: Bolivia, Un-

duavi, Buchtien 878 (holotype, not located; iso-

type, us).

In open sites, wet hillsides, grasslands, rocky

ground, and boggy places, 1900-3400 m. Piura

south to Puno.

Range of the species.

Piura: Prov. Huancabamba, above Huancabamba.

Hutchison 1620 (GH). Amazonas: Prov. Bongara, Dist.

Yambrasbamba, Tillet 673-340 (GH, USM). Balsas to Lei-

mebamba, Barro Negro, E. E. Smith (habitat photo, GH).

San Martin: Rio Abiseo National Park, Young 1981

(HUT, USM). Prov. Chachapoyas. south side of Molino-

pampa-Disan pass, Wurdack 1614 (USM). Pasco: Prov.

Oxapampa, Santa Barbara, D. Smith 8102 (F). Junin:

Prov. Satipo, Dist. Pampa Hermosa, Sounders 1030 (GH),

7066 (GH). Cuzco: Prov. La Convencion, Cordillera Vil-

cabamba, Dudley 11031 (GH). Prov. Urubamba. near

Sayacmarco. Peyton & Peyton 290 (GH). Paucartambo.

Soukup 375 (F). Tres Cruces, Gentry el al. 23440 (F).

Puno: Prov. Sandia, Vargas 14856 (GH). Prov. Sandia.

near Limbani. Metcalf 30519 (GH).

18b. Blechnum auratum ssp. columbiense (Hi-

eron.) Tryon & Stolze, comb. nov.

Blechnum columbiense Hieron., Hedwigia 47: 244, /.

5,f. 17. 1908. TYPE: Colombia, Huila. Paramo

de Moras, Stubel 1268 (holotype, B).

Blechnum columbiense var. bogotense Hieron.. Hed-

wigia loc. cit. 244. 1908. SYNTYPES: Colombia.

Bogota. Karsten (B), Lindig 27 (B): isosyntype,

Lindig 27 (BM; photo. GH).

Elfin forests and jalca, 2700-3300 m, Piura and

Amazonas. Venezuela and Colombia, south to

Peru.

D. Smith 7720 (USM), Dept. Pasco, is this variety

or near it.

Piura: Prov. Huancabamba, Hutchison 1619 (F, GH).

Amazonas: 20 km downstream from Balsas. Madison

1138 (GH). La Jalca. Chachapoyas. LI. Williams 7570

(F).

19. Blechnum obtusifolium Ettingsh.. Denkschr.

Kaiserl. Akad. Wiss. Wien. 23: 59, /. 8. f. 4.

1864. Also Ettingsh., Farnkr. Jetzw. 122. 1865.

Lomaria obtusifolia Presl, Tent. Pterid. 143. 1836.

nom. nud. Since Ettingshausen uses Presl's name,

the specimen cited by Presl. "Herb. bras. reg. be-

rol. 101" may be considered to be the type (not

located).

/

Stem more or less erect, massive, becoming ar-

borescent. Leaves dimorphic. Petiole with basal

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

67

scales very numerous, long (to 3.5 cm or more),

acicular beyond a usually broader base, usually

bicolorous with a blackish center and lighter, brown

margins, or mostly concolorous, brown, stem scales

very long and narrow, often curved, attenuate,

brownish often with dark brown to blackish cen-

ter. Sterile lamina pinnatisect distally, 1 -pinnate

proximally, the apex abruptly reduced to an entire,

usually large terminal segment, the base gradually

reduced to small pinnae or lobes, pinnae patent

to ascending, sessile with a cuneate to rounded

base proximally or broadly sessile or partly adnate

there, margins flat to somewhat involute, distant

from the costa. Fertile pinnae with green tissue

extending hardly or only slightly beyond the at-

tachment of the indusium.

The status and nomenclature of this species are

uncertain, the group of species related to Blech-

num magellanicum being in need of revision. The

description of Ettingshausen is not wholly satis-

factory; additional characters are mentioned by

Christ, Bull. Herb. Boisier 2, 4: 1091. 1904, and

also by Hieronymus, Hedwigia 47: 243. 1908.

Two collections, wet forest and forest border,

550 and 220 m, Cuzco, and Madre de Dios.

Peru and northwestern Argentina; southeastern

Brazil; range uncertain.

Cuzco: Prov. Paucartambo, Pillawata, Vargas 16698

(GH). Madre de Dios: Prov. Manu, Shintuya, Chavez 834

(fertile), 835 (sterile) (MO).

Comments

One collection of what may be a new species of

subgenus Blechnum has not been identified: Ama-

zonas, Prov. Bagua, Cordillera Colan, Barbour

3589 (MO). The specimen is evidently the central

portion of a large fertile leaf. The long and narrow

fertile pinnae are short-stalked and bear costal sori

with expanded green tissue well beyond the at-

tachment of the indusium.

The following names have not been associated

with a Peruvian species of Blechnum for the rea-

sons stated.

Blechnum ciliatum Presl, Reliq. Haenk. 1: 50.

1825. "Hab. in Peruvia et in Cordilleris Chilen-

sibus." Under Parablechnum ciliatum (Presl) Presl,

Epim. bot. 109. 1851, only Chile is cited. Placed

by Murillo (1968) as a synonym of Blechnum au-

riculatum Cav.

Blechnum cognatum Presl, Epim. bot. 107. 1851.

"Habitat in vallibus Cordillerae Peruviae

(Haenke)." Murillo (1968) places this name, ex

char, as a synonym of Blechnum occidentale L.

Blechnum delicatum Maxon & Morton, Bull.

Torrey Bot. Club 66: 41. 1939. TYPE: Robore,

Bolivia, Cardenas 3006 (holotype, us!) This seems

to be an unusual variation of Blechnum asple-

nioides.

Blechnum lechleri Mett., Fil. Lechl. 2: 17. 1859.

"Peruvia, Tatanara; Brasilia (Bongard)." I have

not examined authentic material of this species.

It may be Blechnum cordatum.

Lomaria linariaefolia Presl, Reliq. Haenk. 1 : op.

cit. 52. 1825. "Hab. in Peruviae?"

Blechnum malacothrix Maxon & Morton, Bull.

Torrey Bot. Club 66: 40. 1939. TYPE: Okara, Cor-

dillera Real, Bolivia, Tate 951 (holotype, NY; iso-

type, us!). This may be a valid species, or it may

be Blechnum confluens.

Blechnum trilobum Presl, loc. cit., t. 9,f.2.\ 825.

"Hab. in Peruvia." The figure is Blechnum auri-

culatum Cav. of Chila NE to southern Brazil.

II. Salpichlaena

Salpichlaena Hooker, Gen. fil., /. 93. 1842, as Sal-

pichloena in text, corrected in index. TYPE:

Salpichlaena volubilis (Kaulf.) Hooker (Blech-

num volubile Kaulf.). Figure 4.

Terrestrial. Stem long-creeping to short-creep-

ing and moderately stout. Leaves dimorphic or

partly dimorphic, the fertile pinnae with narrower,

sometimes very narrow segments, scrambling or

climbing to 15 m or more long, indeterminate.

Lamina 2-pinnate, the pinnae imparipinnate. Veins

free, connected by a marginal strand. Sori on a

long vascular commissure close to the costa, not

paraphysate, covered by an indusium of similar

extent, arching completely over the sporangia, and

opening toward the costa. Spores ellipsoidal,

monolete, with spherical deposit over the papil-

late-rugose surface.

FIG. 4. Salpichlaena volubilis: a, sterile pinna; b, fertile pinnule; c, section of pinnule with sori. (From Plowman

2550, F.)

68

FIELDIANA: BOTANY

5cm

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

69

Salpichlaena is a widely distributed tropical

American genus of one species. The leaves of ju-

venile plants are entire or 1 -pinnate, and deter-

minate, while those of adult plants are 2-pinnate,

indeterminate, and scrambling or high-climbing

in trees.

1. Salpichlaena volubilis (Kaulf.) Hooker, Gen.

fil., t. 93. 1842. Figure 4.

Blechnum volubile Kaulf., Enum. fil. 159. 1824. TYPE:

perhaps Brazil, Chamisso (holotype, LZ, de-

stroyed: no isotype located).

Blechnum scandens Bory, in Duperrey, Voy. Coquille

Bot. 272, t. 36. 1 829. TYPE: Brazil, St. Catharina,

D'Urville (holotype, P; isotype, GH!).

Lomaria volubilis Hooker, Sp. fil. 3: 39. 1860. TYPE:

Brazil, Barra do Rio Negro, Spruce 1263 (holo-

type, K; isotype, ?, GH!).

Blechnum volubile var. lomarioideum Baker in Mar-

tius, Fl. Brasil. 1(2): 428. 1870.

Spicanta hookeriana O. Kuntze, Revis. gen. pi. 821.

1891. Nom. nov. for Lomaria volubilis Hook, and

with the same type.

Salpichlaena hookeriana (O. Kuntze) Alston, Bull.

Misc. Inform 1432: 312.

Salpichlaena lomarioidea (Baker) A. R. Sm., Ann.

Missouri Bot. Gard. 77: 250. 1990.

Stem scales small, rigid, dark brown to blackish.

Pinnae usually subopposite. Segments acute, nar-

rowly acuminate to caudate, entire or often serrate

at the tip, sessile to rather long-stalked, cuneate to

broadly rounded at the base, the margins cartilagi-

nous, the costa prominently abaxially, the sterile

ones ca. 1.5 mm to 3 cm broad and 10-30 cm

long. Sori costal, very long.

Terrestrial in rain forests, montane forests, in-

undated thickets and forests near rivers, rarely in

cloud forests, 100-1900 m, mostly below 700 m,

Amazonas and especially Loreto, south to Cuzco.

Central America; Lesser Antilles; south to Bo-

livia and southeastern Brazil.

The chromosome number of n = 40 is unique

in the family. This is a very distinctive genus and

species, with a long scrambling or climbing leaf.

The habit is the same as that ofLygodium species.

Rarely the fertile and sterile segments are of es-

sentially the same breadth. It is probably more

widely distributed than the collections indicated

since the climbing leaves may be partly concealed

among tree branches, or the plant may not be rec-

ognized as a fern.

In some specimens, the fertile segments are very

narrow, with little or no green tissue extending

beyond the attachment of the indusium. These

have been named Salpichlaena hookeriana— for

example, Plowman 2550 and McDaniel 11758,

both Dept. Loreto (GH), are this narrow variant.

Some specimens have narrow segments and only

narrow green tissue beyond the indusium. Buds

may be present in the axils of the pinnae (less often

the axils of the pinnules), and these occur both in

specimens with narrow and broad fertile segments.

The narrow variant seems restricted to low ele-

vations, while the specimens with broader seg-

ments are also from higher elevations. Further

study is required before another taxon is recog-

nized with confidence.

Amazonas: Rio Maranon, above Cascadas de Mayasi,

Wurdack 1893 (F, GH). Prov. Bagua, 1 2 km E of La Peca,

Barbour 2402 (F, MO). E of La Peca, Gentry et al. 22905

(F, USM). San Martin: Zepelacio near Moyobamba, Klug

3514 (F, GH, MO). Prov. Mariscal Caceres, Dist. Tocache

Nuevo, J. Schunke V. 5541 (F, MO), 7079 (F, USM). Lo-

reto: Sierra del Pongo, Mexia 6276 (F, GH, MO). Pumaya-

cu, Klug 3 197 (F, GH, MO). Shiriara, Rio Nanay, Plowman

2550 (F, GH). Versailles, Iquitos, McDaniel 1 1758 (F, GH,

MO). Huanuco: Tingo Maria, Allard 21386 (GH). Pasco:

Prov. Oxapampa, Rio Pichis, 1 hr below Puerto Ber-

mudez, Gentry et al. 42123 (F). Prov. Oxapampa, N of

Chequitavo, D. Smith 5118 (MO). Junin: La Merced,

Soukup 1 106 (F). Near La Merced, Killip & Smith 23923

(F). Above San Ramon, Killip & Smith 25565 (F). Cuzco:

Prov. La Convencion, Cordillera Vilcabamba, Dudley

10451 (GH, MO, USM).

Family 21. POLYPODIACEAE

POLYPODIACEAE Bercht. & J. S. Presl, Priroz-

en. Rostlin 1: 272. 1820. TYPE: Polypo-

dium L.

Stem erect to short-creeping, or very long-creep-

ing, small to rather stout, or slender to filiform,

with scales. Leaves ca. 3 cm to 2 m long, circinate

in vernation, monomorphic or dimorphic, entire,

pinnatifid, or variously forked, or pinnate, gla-

brous, pubescent or scaly abaxially. Petiole lacking

stipules, often articulate to the stem. Veins free to

anastomosing. Sori abaxial, round to elongate,

borne on a vein, at the tip of a vein, at the junction

of veins, on a vascular commissure, or on a special

series of fertile veins, exindusiate, paraphysate or

not. Sporangia with a 1 - or 2-rowed stalk basally,

the vertical annulus interrupted by the stalk.

The Polypodiaceae are a large family of perhaps

40 genera and 1 000 species, most of these being

70

FIELDIANA: BOTANY

epiphytic or epipetric. The classification of this

group is difficult, and authors often disagree on

the taxa to be recognized and their rank. Three

tribes are adopted here, each of these sometimes

being treated as a family.

The tribe Polypodieae usually has the petiole

sulcate and articulate to the stem, the petiole and

lamina have short trichomes or none, the sporan-

gium has two rows of cells at its base, and the

spores are ellipsoidal, monolete and only rarely

green. There are nine genera in Peru.

The tribe Loxogrammeae is intermediate be-

tween the two other tribes, having some characters

of one and some of the other. It does not occur in

Peru.

The tribe Grammitideae usually has the petiole

terete and continuous (not articulate) with the stem,

the petiole and lamina often with long trichomes,

the sporangium has a single row of cells at its base,

and the spores are usually green and usually tet-

rahedral-globose, trilete. It is represented in Peru

by the single genus, Grammitis.

References

COPELAND, E. B. 1947. Polypodiaceae, pp. 174-

222. In Genera Filicum. Chronica Botanica,

Waltham, Mass.

TRYON, R. M., AND TRYON. 1982. Polypodi-

aceae, pp. 684-758. In Ferns and Allied Plants.

Springer- Verlag, New York.

Key to Genera of Polypodiaceae

a. Petiole and usually the lamina with long, multicellular, acuminate trichomes; sporangium stalk

uniseriate at base; spores usually with chlorophyll (green) and tetrahedral-globose . . I. Grammitis

a. Petiole and lamina without trichomes, or these present and usually unicellular, short; sporangium

stalk bi- or triseriate; spores lacking chlorophyll (with rare exception), ellipsoidal, monolete . . . . b

b. Lamina dichotomously or subdichotomously forked or veined c

c. Lamina with scattered scales V. Dicranoglossum

c. Lamina with stalked, stellate trichomes X. Platycerium

b. Lamina entire, lobed, or pinnatifid to 3-pinnate d

d. Stem bearing tubers; spores prominently echinate; epiphytic IX. Solanopteris

d. Stem without tubers; spores nearly smooth, verrucate, rugose, tuberculate, papillate or with

folds; terrestrial, rupestral, or epiphytic e

e. Lamina pinnatifid, pinnatisect, or to 3-pinnate f

f. Stem scales peltately attached III. Polypodium

f. Stem scales basally attached II. Pecluma

e. Lamina entire g

g. Stem more or less flattened, its scales not clathrate, attached near the center

VI. Microgramma

g. Stem more or less terete, its scales usually clathrate h

h. Sori in 2 rows between strong lateral veins, or lateral veins obscure or absent ... i

i. Immature sori with peltate paraphyses; most sori served by 2 or more veins; lamina

surface scaly (at least sparsely so) IV. Pleopeltis

i. Paraphyses absent or minute; most sori borne back of a single vein tip; scales absent

on the lamina or only a few on the costa VII. Campyloneurum

h. Sori in 1 row between strongly developed lateral veins j

j. Stem very long-creeping HI. Polypodium

j. Stem more or less erect, decumbent or short-creeping VIII. Niphidium

Marginariopsis C. Chr. (Dansk. Bot. Ark. 6(3):

42. 1 920) is an epiphyte that occurs from Ecuador

north to Costa Rica and it may grow in Peru. Its

stem is rather flattened and bears finely clathrate

scales, the leaves are dimorphic, the fertile much

narrower than the sterile, the lamina is entire, with

small scales, and the sporangia are borne on anas-

tomosing veins.

Many American species of Polypodium have

been placed in the genus Goniophlebium. That

name, however, applies to an Old World genus

and is typified by Polypodium subauriculatum

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

71

Blume (Goniophlebium subauriculatum (Blume)

Presl).

I. (.ram in it is

Grammitis Sw., J. Bot. (Schrad.) 1800(2): 17. 1801.

TYPE: Grammitis marginella (Sw.) Sw.

(Polypodium marginellum Sw.). Figure 5.

Xiphopteris Kaulf., Berl. Jahrb. Pharm. 21: 35. 1820.

LECTOTYPE (designated by John Sm., Hist. fil.

179. 1875: Xiphopteris serrulata (Sw.) Kaulf.

Grammitis section Xiphopteris (Kaulf.) Presl, Tent.

Pterid. 208. 1836.

Cochlidium Kaulf., Berl. Jahrb. Pharm. 21: 36. 1820.

TYPE: Grammitis graminoides (Sw.) Sw. (Acros-

tichum graminoides Sw., Cochlidium grami-

noides (Sw.) Kaulf.) (section Pleurogramme).

Ctenopteris Kunze, Bot. Zeit. (Berlin) 4: 425. 1846.

TYPE: Ctenopteris venulosa (Blume) Kunze

(Polypodium venulosum Blume = Grammitis ven-

ulosa (Blume) R. & A. Tryon (section Cryptoso-

rus).

Cryptosorus Fee, Mem. foug. 5 (Gen. fil.): 231. 1852:

TYPE: Cryptosorus blumei Fee, nom. super/], for

Polypodium obliquatum Blume = Grammitis ob-

liquata (Blume) Hassk. Polypodium section Cryp-

tosorus (Fee) Fourn., Ann. Sci. Nat. Bot., ser. 5,

18: 282. 1873. Grammitis section Cryptosorus

(Fee) R. & A. Tryon, Rhodora 84: 128. 1982.

Glyphotaenium (John sm.) John Sm., Hist. fil. 187.

1875. Ctenopteris subgenus Glyphotaenium John

Sm., Bot. Voy. Herald (Seemann) 227. 1854.

TYPE: Ctenopteris crispata John Sm. (Glypho-

taenium crispatum (John Sm.) John Sm.) =

Grammitis crispata (John Sm.) Morton. Gram-

mitis section Glyphotaenium (John Sm.) R. & A.

Tryon, Rhodora 84: 128. 1982.

Enterosora Baker, Timehri 5: 218. 1886. TYPE: En-

terosora campbellii Baker = Grammitis camp-

bellii (Baker) Proctor (section Glyphotaenium).

Micropolypodium Hayata, Bot. Mag. Tokyo 42: 302,

341. 1928. TYPE: Micropolypodium pseudotri-

chomanoides (Hayata) Hayata.

Ceradenia Bishop, Amer. Fern J. 78: 2. 1988. TYPE:

Polypodium curvatum Sw. (Ceradenia curvata)

(Sw.) Bishop) = Grammitis curvata (Sw.) Ching.

Zygophlebia Bishop, Amer. Fern J. 79: 107. 1989.

TYPE: Polypodium sectifrons Mett. (Zygophlebia

sectifrons (Mett.) Bishop) = Grammitis sectifrons

(Mett.) Seymour.

Lellingeria Smith & Moran, Amer. Fern J. 81: 76.

1991. TYPE: Polypodium apiculatum Klotzsch

(Lellingeria apiculata (Klotzsch) Smith & Mor-

an).

Plants commonly epiphytic, occasionally epi-

petric or terrestrial. Stem small, mostly short-

creeping, sometimes erect, commonly scaly (scales

clathrate or not), rarely naked, bearing slender,

fibrous roots. Leaves monomorphic or rarely the

apical fertile portion modified, 2-60 cm (rarely to

1 m) long, rarely articulate to the stem, scales lack-

ing, or present only at petiole base, petiole long to

obsolete. Lamina simple and entire to pinnatisect

(very rarely to 3-pinnate), glabrous or variously

pubescent (rarely densely so), the trichomes 1-

few-celled. Veins commonly free, sometimes anas-

tomosing and with or without free, included vein-

lets. Sori round to elongate, sometimes confluent,

borne on the vein or vein branch, sometimes deep-

ly immersed in the laminar tissue, or sporangia in

lines on each side of the costa, paraphysate or not,

exindusiate. Spores trilete, tetrahedral-globose,

greenish (chlorophyllous).

Grammitis is a pantropical and austral genus of

more than 400 species, with perhaps half of these

in America. From the time of Copeland's (1947)

"Genera Filicum" to the present it has been di-

vided by various authors into as many as 20 genera

or subgenera, and some of these divisions may be

warranted. Alan R. Smith and Robbin C. Moran

are currently attempting a revision of the Amer-

ican species that will recognize nine genera, the

concepts partially based on the recent work of

Bishop (1977, 1978, 1988, 1989a). The authors

have kindly provided provisional manuscripts of

these genera, which have been very helpful in con-

structing a key to the species in Peru. Nevertheless,

pending the results of these new studies, Gram-

mitis is recognized in the broadest sense for pur-

poses of the present treatment.

The key to species in Peru is based in part on

the generic divisions ultimately to be proposed by

Smith and Moran. For example, species 1-3 are

Grammitis s.s., essentially defined in the Neo-

tropics by the dark, sclerenchymatous border of

the lamina, and species 7-9 share the characters

of Enterosora (= Glyphotaenium): lamina subcar-

FIG. 5. Grammitis myosuroides: a, habit. Grammitis limbata: b, habit; c, leaf section, with sori. Grammitis david-

smithii: d, habit; e, pinna, adaxial side; f, stem scale. Grammitis myriophylla: g, section of rachis with fertile pinna

base, (a from Camp E-5131, Ecuador, F; b from Steyermark & Wurdack 1 178, Venezuela, F; c from D. Smith 2038,

F; d from Beck 1424 'A, Bolivia, F; e, f from Smith & Canne 5837, holotype, F; g from Leon 1005, F.)

72

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

73

nose and entire to lobed, with sori immersed in

the tissue, hydathodes lacking.

The character of hydathodes is effective in sep-

arating some major species groups, in addition to

Glyphotaenium. In many species, hydathodes are

easily visible to the naked eye— usually seen as

expanded vein tips differing in color from the ad-

jacent tissue. In other species, closer scrutiny is

necessary; i.e., hydathodes may be hidden by dense

indument, or they may be located at the very seg-

ment margin, the adaxial portion of which is

strongly revolute, or they may be less conspicuous

to the naked eye, yet quite evident with slight mag-

nification (10-12x).

Besides the group of Enterosora, two other an-

hydathodous groups are to be recognized by Smith

and Moran, these separated by a presumably good

and consistent character: sori with conspicuous,

pale, waxlike paraphyses (Ceradenid) vs. paraph-

yses lacking, or dark and not waxlike (Zygophle-

bid). Unfortunately, the presence and character of

the paraphyses often cannot be ascertained in ma-

ture sori nor in heat-dried specimens (Bishop, 1988,

p. 2). Species of Zygophlebia are further charac-

terized by strongly to casually anastomosing veins

(free in Ceradenid), but in two of three species in

Peru (G. dudleyi, G. werffii) the lamina texture is

so thick that veins can only be examined by the

lengthy process of clearing. Thus, the two most

important distinguishing features of these two

groups are in most cases functionally ineffective

insofar as rapid field or herbarium identification

is concerned.

Species 29-66, with hydathodes, are much more

difficult to separate into discrete groups; however,

the characters used in the key that best seem to

indicate species relations are those of stem scales,

laminar trichomes, and sporangia (capsular setae

present or lacking).

Several species complexes in this treatment are

still very poorly understood, and relationships of

their components are sorely in need of study. The

groups most in need of attention are those of G.

cultrata/heteromorpha, G. semihirsuta, G. monil-

iformis, and G. flabelliformis/pilosissima, each of

these probably containing four to eight taxa. The

species delineations herein seem to serve rather

well insofar as Peruvian collections are concerned,

but many of the taxa vary or intergrade consid-

erably in Mexico and Central America, the West

Indies, and other parts of South America. Most of

the problems are pointed out in the respective

species discussions that follow.

References

BISHOP, L. E. 1977. The American species of

Grammitis section Grammitis. Amer. Fern J.,

67: 101-106.

. 1978. Revision of the genus Cochlidium

(Grammitidaceae). Amer. Fern J., 68: 76-94.

-. 1988. Ceradenia, a new genus of Gram-

mitidaceae. Amer. Fern J., 78: 1-5.

. 1989a. New species of Ceradenia. Amer.

Fern J., 79: 14-25.

. 1989b. Zygophlebia, a new genus of

Grammitidaceae. Amer. Fern J., 79: 103-1 18.

BISHOP, L. E., AND A. R. SMITH. 1992. Revision

of the fern genus Enterosora (Grammitidaceae)

in the New World. Syst. Bot., 17: 345-362.

CHRISTENSEN, C. 1929. Taxonomic fern studies.

I. Revision of the polypodioid genera with lon-

gitudinal coenosori (Cochlidiinae and "Dry-

moglossinae"; with a discussion of their phy-

logeny. Dansk. Bot. Ark., 6(3): 1-93.

COPELAND, E. B. 1952a. Grammitis. Phillip. J.

Sci., 80:93-271.

. 1952b. The American species of Xiph-

opteris. Amer. Fern J., 42: 41-52, 93-1 10.

. 1956. Ctenopteris in America. Philipp.

J. Sci., 84:381-473.

MAXON, W. R. 1914. Notes upon Polypodium

duale and its allies. Contr. U.S. Natl. Herb., 17:

398-406.

. 1916. Polypodium trichomanoides and

its American allies. Contr. U.S. Natl. Herb., 17:

542-557.

MORTON, C. V. 1967. The genus Grammitis in

Ecuador. Contr. U.S. Natl. Herb., 38: 85-123.

SMITH, A. R., R. C. MORAN, AND L. E. BISHOP.

1 99 1 . Lellingeria, a new genus of Grammiti-

daceae. Amer. Fern J., 81: 76-88.

Key to Species of Grammitis

a. Lamina simple, entire, with a black or dark brown, sclerenchymatous border b

b. Lamina less than 3 mm broad; stem scales 2-6 cells across at base 3. G. paramicola

b. Lamina 3-8 mm broad; stem scales 8-12 cells across at base c

c. Hydathodes present; sclerenchymatous border commonly less than 0. 1 mm broad; lamina firm-

herbaceous to chartaceous . . 2. G. limbata

74

FIELDIANA: BOTANY

c. Hydathodes lacking; border commonly 0.2 mm broad; lamina subcoriaceous

1 . G. bryophila

a. Lamina lobed to decompound or, if simple, then lacking a dark, sclerenchymatous border d

d. Lamina (at least fertile portions) simple and subentire to repand or serrate-dentate, 1-2 mm broad,

the son confluent and commonly covering the costa at maturity e

e. Leaves monomorphic, fertile and sterile portions entire throughout, ca. 1 mm broad

4. G. pumila

e. Leaves partly dimorphic, fertile distal portion subentire, sterile leaf or portion serrate-dentate

to pinnatisect, to 4 mm broad f

f. Lamina trichomes castaneous, scattered but persistent at least on costa and veins; sporangia

borne at base of veins, confluent as 2 subcostal lines until merging into a broad single line

at full maturity; sterile portion of lamina pennatisect 5. G. myosuroides

f. Lamina trichomes lacking (or pale and caducous on juvenile plants); sporangia borne on

costa and veins, thus confluent as a single, costal line; sterile portion of lamina serrate-

dentate or with short, acute lobes 6. G. serrulata

d. Lamina pinnatisect to decompound throughout, or if simple to shallowly lobed then 4-20 mm

broad and sori discrete, in 1 or more lines on each side of, and subdistant from, the costa . . . g

g. Hydathodes lacking, or minute and discernible only with high (more than 20 x ) magnification

h

h. Lamina simple, subentire or lobed to pinnatifid (not cut nearly to the costa) i

i. Leaves 1-3.5 cm broad; stem scales yellow or yellowish brown; sori in (l-)2-5 series

between costa and margin j

j. Leaves 2-3.5 cm broad, cut Vj-'/z to the costa; petiole lustrous black or atropurpureous;

sori in 3-5 series between costa and margin 7. G. trifurcata

j. Leaves 1-1.5 cm broad, subentire to very shallowly lobed; petiole dull yellowish or

reddish brown; sori in (l-)2 series between costa and margin .... 8. G. trichosora

i. Leaves 0.4-0.8 cm broad; stem scales deep orange to castaneous; sori in a single series

between costa and margin k

k. Stem scales deep orange, entire or remotely ciliolate; lamina base attenuate; petiole

trichomes castaneous 9. G. parietina

k. Stem scales castaneous, conspicuously setose; lamina base cuneate; petiole trichomes

orange 10. G. phalacron

h. Lamina pinnatisect (cut nearly to the costa) to 3-pinnate 1

1. Lamina 2-3-pinnate; ultimate segments 0.7-3 mm long m

m. Stem scales clathrate, margins entire or with some short glands near apex; petiole and

lamina puberulent, but lacking long, spreading, castaneous trichomes

24. G. in i rahi I is

m. Stem scales nonclathrate, margins abundantly setose; petiole and lamina with spread-

ing castaneous trichomes to 1.5 mm long (and puberulent as well)

28. G. bipinnata

1. Lamina 1 -pinnate or deeply pinnatisect; ultimate segments 5 mm long or (usually) much

longer n

n. Veins regularly merging to form usually quite evident areoles, at least when held to

light 25. G. mathewsii

n. Veins free, or, if casually anastomosing, then completely obscure, even when held to

light o

o. Lamina not or scarcely reduced at base, or basal pair of segments more than half

as long as median ones; petiole about half as long as or longer than the lamina .

P

p. Pinnae, at least proximal ones, deeply serrate, the others crenulate

11. G. longipinnata

p. Pinnae subentire q

q. Stem scales rather flaccid, pale orange or yellow-brown, entire, lacking mar-

ginal setae; plants epiphytic 26. G. dudleyi

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 75

q. Stem scales rigid, deep orange to castaneous, with conspicuous marginal setae;

plants terrestrial r

r. Rachis and costae (and usually segments) amply provided on both sides

with long, spreading trichomes, these 0.5-2 mm long

12. G. meridensis

r. Rachis (and usually costae) densely short-puberulent adaxially, the tri-

chomes 0. 1-0.3(-0.4) mm long s

s. Setae of stem scales dark, the same color as the scale body; stem long-

creeping, the leaves mostly wide spaced 13. G. bishopii

s. Setae of stem scales whitish, in strong contrast with the dark scale body;

stem short-creeping to ascending, leaves approximate or crowded . . .

27. G. werffii

o. Lamina somewhat to strongly reduced at base, several to many proximal segments

much shorter than medial ones or reduced to mere auricles; petiole obsolete or

much shorter than the lamina t

t. Petiole lacking conspicuous, spreading trichomes u

u. Petiole essentially obsolete; sclerenchyma of the rachis and costae obscured

by the thick lamina tissue v

v. Lamina segments 1-2 times as long as broad; lamina moderately to densely

white-farinose, the farina mostly disposed in stellate patterns

14. G. discolor

v. Lamina segments (larger ones) 3-7 times as long as broad; lamina mod-

erately farinose to naked, the farina deposits solitary or in amorphous

clusters 15. G. curvata ssp. pearcei

u. Petiole (1-) 2- 10 cm long; sclerenchyma of the rachis (and sometimes costae)

exposed, at least proximally w

w. Lamina pinnatisect, segments closely spaced, joined by at least a narrow

wing of tissue, gradually narrowing from longest medial pinnae to nearly

auricular ones at base x

x. Stem scales 3-5 mm long, setulose; segments ascending at a 30-50°

angle; sori medial 16. G. terrestris

x. Stem scales ca. 1 mm long, subentire; segments ascending at a 60-80°

angle; sori supramedial 17. G. herrerae

w. Lamina 1 -pinnate, pinnae adnate, but discrete and mostly subdistant,

abruptly reduced at base, with only ca. 3-6 pinnae shorter than medial

ones y

y. Pinnae 1-3.5 cm long, 2.5-5 mm broad, ascending at a 60-80° angle,

base rounded to truncate acroscopically; sori generally confined within

pinna margins 19. G. farinosa

y. Pinnae to 10 cm long, ca. 1.5 mm broad, ascending at a 20-35° angle,

attenuate at base; sori at maturity extending well beyond pinna margins

18. G. dendrodoxa

t. Petiole (and often the rachis and lamina) amply provided with orange to cas-

taneous, spreading, unicellular trichomes, these 0.5-3 mm long z

z. Lamina 0.6-1 cm broad; pinnae circular, obovate or broadly oblong, obtuse

at the apex, 1-2 times as long as broad 20. G. congesta

z. Lamina 2-10 cm broad; pinnae linear to deltate, acute to attenuate at the

apex, several to many times longer than broad a2

a2. Pinna base rounded nearly to the costa acroscopically, short-decurrent

basiscopically; sori not extending beyond the pinna margin

21. G. praeclara

a2. Pinna base fully adnate and nearly perpendicular to the rachis acroscopi-

cally, long-decurrent basiscopically; sori mostly extending beyond the

pinna margin at maturity b2

76 FIELDIANA: BOTANY

b2. Lamina thin- to firm-herbaceous; rachis evident and usually lus-

trous, at least abaxially; petiole 0.2-0.4(-0.5) mm in diameter . . .

22. G. capillaris

b2. Lamina subcarnose; sclerenchyma of the rachis usually obscured by

laminar tissue, or rarely evident and then not lustrous; petiole

(0.4-)0.6-0.9 mm in diameter 23. G. pilipes

g. Hydathodes evident (or in a few species hidden by dense trichomes) c2

c2. Trichomes on petiole and/or lamina 0.1-0.3 mm long or lacking and the stem scales

conspicuously setose d2

d2. Lamina subtruncate at base, or a few proximal pairs of pinnae about half as long as

medial ones 30. G. apiculata

d2. Lamina attenuate at base, 2-6 proximal pairs of pinnae rudimentary and auriculiform

e2

e2. Rachis thin and conspicuously flexuous, sparsely puberulent adaxially; pinna base

inequilateral, rounded or truncate acroscopically, decurrent basiscopically; pinnae

ascending at a 20-45° angle 31. G. pseudocapillaris

e2. Rachis stout and rigid, not or slightly flexuous, densely puberulent adaxially; pinna

base subequilateral, either subtruncate or dilated; pinnae patent or slightly as-

cending (often strongly ascending in G. subsessilis) f 2

f2. Mature leaves 0.8-1.5 cm broad; petiole 0.3-0.4 mm in diameter; pinnae

oblong, 3—4 times longer than broad 32. G. tunguraguae

f2. Mature leaves 2-7 cm broad; petiole (0.4-)0.5-1.2 mm in diameter; pinnae

linear to deltate, 6-12 times longer than broad g2

g2. Pinnae often strongly ascending, linear from a strongly dilated base; stem

scales 2-4 mm long, obviously clathrate, the proximal cells in 5-10 series

across the scale and the lumina clear 33. G. subsessilis

g2. Pinnae patent to slightly ascending, deltate or oblong-deltate; stem scales

(4— )5-7 mm long, narrowly clathrate, the proximal cells in (12-) 15-20

series across the scale and the lumina mostly occluded . . 34. G. major

c2. Trichomes on petiole and/or lamina (0.4— )0. 5-2 mm long or, if shorter or lacking, then

stem scales entire h2

h2. Trichomes on petiole and/or lamina 0.1-0.3 mm long or lacking (occasionally a few,

scattered, longer ones present, especially on the petiole in #40) i2

i2. Pinnae deeply lobed to pinnatifid, most of them ascending at a 30—45° angle ....

35. G. gracilis

i2. Pinnae with margins entire to crenulate, patent or slightly ascending j2

j2. Pinnae conspicuously inequilateral at base (i.e., broadly rounded to subauricu-

late acroscopically, strongly decurrent basiscopically); lamina thin-herbaceous

36. G. phlegmaria

j2. Pinnae subequilateral, equally dilated at base, or basiscopically somewhat more

spreading; lamina chartaceous to coriaceous k2

k2. Pinnae or segments ascending at a 60-70° angle, linear-deltate, 6-8 times

as long as broad 37. G. youngii

k2. Pinnae or segments patent, semicircular to oblong or deltate, l-3(— 4) times

as long as broad 12

12. Laminar tissue amply provided with appressed, whitish, viscid tri-

chomes, these often obscuring the hydathodes 29. G. jamesonioides

12. Laminar tissue essentially glabrous, the hydathodes conspicuous ....

m2

m2. Stem scales l-2(-2.5) mm long, not iridescent, cell walls blackish;

segments plane 38. G. melanosticta

m2. Stem scales 3-8 mm long, iridescent, cell walls brown or reddish

brown; segment margins (at least fertile ones) revolute n2

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

n2. Leaves well spaced on the stem; segments 2—4 times as long

as broad; plants not aromatic 39. G. erecta

n2. Leaves commonly approximate to subfasciculate; segments

1-1.5 times as long as broad; plants (especially dried) with a

resinous scent 40. G. moniliformis

h2. Trichomes on petiole and/or lamina (0.4-)0.5-2 mm long o2

o2. Leaves 1 -pinnate and 1-2-forked 50. G. heteromorpha

o2. Leaves pinnatisect to 2(-3)-pinnate, not (or very rarely) forked p2

p2. Pinnae crenate or lobed (at least near the base) to l(-2)-pinnate q2

q2. Pinnae crenate to deeply lobed; stem scales black or dark brown

64. G. athyrioides

q2. Pinnae deeply pinnatisect to 1 (-2)-pinnate; stem scales orange or light

brown r2

r2. Ultimate segments linear; margins of stem scales entire or with a

few short glands; sporangia not setose 65. G. myriophylla

r2. Ultimate segments oblong to nearly circular; margins of stem scales

long-setose; sporangia long-setose s2

s2. Hydathodes commonly encrusted with white deposits; pale tri-

chomes of rachis (and segments) mixed with castaneous, stel-

lately branched ones 66. G. immixta

s2. Hydathodes lacking white encrustations; pa> trichomes of rachis

mixed with minute (0.1-0.3 mm), septate, gland-tipped ones .

67. G. variabilis

p2. Pinnae (or segments) entire t2

t2. Sporangia with long setae; axes with most of the trichomes sericeous and

hyaline to pale orange; stem scales setose u2

u2. Lamina deeply pinnatisect, not or scarcely reduced at base or only

1-2 basal pairs greatly reduced; petiole long, conspicuous

48. G. asplenifolia

u2. Lamina fully 1 -pinnate, strongly and gradually reduced at base; pet-

iole short or obsolete v2

v2. Rachis with stout, castaneous, sessile-stellate trichomes mixed

among the simple, sericeous ones w2

w2. Pinnae 0.4-1 cm long, oblong-elliptic, obtuse, most of them

narrowed at base; petiole 0.2-0.4 mm in diameter; sporan-

gia each with 1 seta 49. G. dependens

w2. Pinnae 1 . 1-4.5 cm long, deltate, acute or subacute, broadly

adnate at base; petiole 0.4-1 mm in diameter; sporangia

with 1-3 setae 5 Ib. G. lanigera var. stella

v2. Rachis lacking stout, castaneous, sessile-stellate trichomes (al-

though some thin, pale, stellate ones may be present) x2

x2. Pinnae 3-6 mm long; pinna base very constricted and in-

equilateral, strongly rounded acroscopically, decurrent basi-

scopically 52. G. subflabelliformis

x2. Pinnae 10—45 mm long; pinna base broadly adnate and

subequilateral, slightly rounded or truncate to somewhat

dilated y2

y2. Pinnae 60-80 pairs, linear, 0.2-0.3 mm broad, 12-20

pairs along 5 cm of rachis; rachis trichomes all simple

53. G. laxa

y2. Pinnae 20-50 pairs, oblong to deltate, (0.3-)0.4-1.2

mm broad, 4-10 pairs along 5 cm of rachis; rachis with

few to many sessile-stellate trichomes mixed among the

abundant simple ones z2

z2. Most pinnae slightly deflexed; petiole 0.2-0.3 mm

78 FIELDIANA: BOTANY

in diameter; lamina sparsely to moderately pilose;

sessile-stellate trichomes sparse on rachis and ab-

axial surface 54. G. cultrata

z2. Most pinnae patent to slightly ascending; petiole

0.4-1 mm in diameter; lamina densely pilose; sim-

ple and sessile-stellate trichomes everywhere abun-

dant 5 la. G. lanigera var. lanigera

t2. Sporangia lacking long setae; axes with stout, rigid, trichomes (except

sericeous in G. taxifolid), stem scales setose or not a3

a3. Stem scales with few to many rigid, marginal setae, these often as

long as the width of the scale body b3

b3. Lamina of mature leaves to 1 cm broad; veins solitary in each

segment and simple or with one acroscopic branch c3

c3. Petiole 0.5-1 .5 cm long; lamina chartaceous; segments often

somewhat gibbous at base acroscopically

55. G. blepharolepis

c3. Petiole obsolete or nearly so; lamina thin- to firm-herba-

ceous; segments not gibbous 56. G. truncicola

b3. Lamina of mature leaves ( 1 . 5-)2-8(- 1 2) cm broad; veins 3-many

pairs in a pinna or segment d3

d3. Stem scales gray-brown or black, clathrate or not; sori me-

dial to inframedial e3

e3. Stem scales gray-brown to blackish, essentially clath-

rate; lamina trichomes mostly 0.4-0.5 mm long; veins

6-14 pairs per segment; hydathodes lacking white de-

posits 57. G. david-smithii

e3. Stem scales lustrous black, nonclathrate; lamina tri-

chomes 1-2 mm long; veins 3-4 pairs per segment;

hydathodes mostly encrusted with white deposits . . .

58. G. pichinchensis

d3. Stem scales orange to castaneous, not or scarcely clathrate;

sori supramedial to submarginal f3

f3. Pinna surface amply and minutely setulose abaxially;

lamina not reduced at base, or with 1-3 pairs of pinnae

somewhat shortened; rachis trichomes 0.4-0.7 mm long

abaxially, shorter adaxially 59. G. alsopteris

f 3. Pinna surface glabrous abaxially; lamina conspicuously

reduced at base, 3-6 pairs of pinnae much shortened;

rachis trichomes mostly 1-2 mm long (sometimes part-

ly broken away on older leaves) g3

g3. Pinnae broad- or oblong-deltate, obtuse to sub-

acute, 4-6 times as long as broad; veins 6-13 pairs

per pinna 60. G. leucosticta

g3. Pinnae linear or linear-deltate, acute, 7-10 times

as long as broad; veins 1 6-24 pairs on larger pin-

nae 61. G. semihirsuta

a3. Stem scales with margins entire (scale apex setose in G. taxifolid) or

minutely glandular or ciliolate h3

h3. Stem scales less than 1 mm long, few and inconspicuous; stems

small (usually less than 1 mm), on long-creeping, stoloniform

roots 41. G. anfractuosa

h3. Stem scales (l-)2-8 mm long, conspicuous and usually in large

masses at least at stem apex; stems 1-5 mm thick, roots not

stoloniform i3

i3. Stem scales conspicuously clathrate, gray-brown, narrow-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V. 79

delta te or -lanceolate, 8-15 cells across at scale base and the

lumina large, clear and elongate j3

j3. Pinnae linear and many of them deflexed, conspicuously

inequilateral at base, strongly dilated acroscopically,

subtruncate basiscopically; lamina (2-)3-7 cm broad;

trichomes on rachis adaxially 0.3-0.5 mm long and per-

sistent 42. G. firma

j3. Pinnae oblong to narrow- or oblong-deltate, not de-

flexed, subequilaterally truncate or slightly dilated at base;

lamina 0.4-2.5 cm broad; rachis trichomes 0.6-1 mm

long (sometimes deciduous) k3

k3. Pinnae ascending, acute; lamina not or abruptly re-

duced at base, proximal 0-3 pairs of pinnae some-

what shortened 43. G. pilosissima

k3. Pinnae patent, obtuse or subacute; lamina strongly

and gradually reduced at base, with 3-many pairs

of pinnae much shortened 13

13. Pinnae mostly oblong, 2.5-6 times as long as

broad; petiole amply provided with spreading,

dark, unicellular trichomes, but minute, tortu-

ous, septate ones rare or lacking; dried speci-

mens strongly aromatic

44. G. xiphopteroides

13. Pinnae deltate or oblong-deltate, 1.5-2 times as

long as broad; petiole moderately provided with

minute, septate trichomes, but spreading, uni-

cellular ones sparse or lacking; specimens not

or weakly aromatic . . . 45. G. flabelliformis

i3. Stem scales not or obscurely clathrate, yellow to lustrous

orange or reddish brown, or if gray-brown then ovate or

broadly lanceolate and minutely clathrate, with 1 8-30 cells

across at scale base and most lumina very small and nearly

isodiametric m3

m3. Stem scales flaccid and dull yellow in mass, with scat-

tered, minute, marginal glands; sori solitary at segment

base; petiole 0.5-2.5 cm long . . . 62. G. blepharidea

m3. Stem scales stout and bright orange to red-brown or

thin and gray-brown, eglandular; sori 2-12 pairs on a

pinna; petiole 3-10 cm long n3

n3. Stem scales orange to red-brown, stout, rigid and

lustrous, nonclathrate, terminating in a pale, apical

(often deciduous) seta; rachis trichomes pale and

sericeous 63. G. taxifolia

n3. Stem scales light or gray-brown, thin, minutely

clathrate, lacking an apical seta; rachis trichomes

rigid and castaneous or lacking o3

o3. Pinnae narrowly deltate or oblong-deltate, 4-

8 times as long as broad, margins essentially

plane, apex acute or subacute

46. G. pseudonutans

o3. Pinnae oblong or broadly oblong-deltate, 1.5-

2 times as long as broad, margins mostly rev-

olute, apex obtuse (occasionally subacute) . .

. 47. G. andicola

80 FIELDIANA: BOTANY

1 . Grammitis bryophila (Maxon) Seymour, Phy-

tologia31: 172. 1975.

Polypodium bryophilum Maxon, Amer. Fern J. 16: 7.

1926. TYPE: Costa Rica, Pcia. San Jose, vicinity

of La Palma, Maxon & Harvey 7980 (holotype,

us!; photo, F).

Plants commonly epiphytic, rarely (and in Peru)

terrestrial. Stem erect to ascending, provided with

bright orange, lustrous, nonclathrate scales, these

1 .5-3 mm long, to 0.5 mm broad, 8-12 cells across

at the base. Leaves simple, entire, 4-25 cm long,

3-5 mm broad. Petiole dark brown to castaneous,

0-3 mm long, 0.2-0.8 mm in diameter. Lamina

linear to narrow-elliptic or -oblanceolate, erect,

subcoriaceous, attenuate at base, the apex acute

to attenuate, provided with a lustrous, black or

castaneous, sclerenchymatous border about 0.2

mm broad, essentially glabrous, but with scat-

tered, minute, subclavate, caducous trichomes, es-

pecially on costa and margin. Veins indistinct, di-

verging from the costa at 30-^tO(-50)° angles, sterile

ones simple, fertile ones forked, hydathodes lack-

ing. Sori elongate, often slightly immersed, usually

confluent at maturity.

In dwarf forest, in sphagnum, 2700 m, San Mar-

tin and Pasco.

Costa Rica; Panama; Venezuela; Colombia;

Peru; Bolivia.

The two specimens cited below from San Martin

are mounted on the same sheet at Kew, each with

the small, blue, handwritten label characteristic of

Spruce, and both are numbered 4643. It is as-

sumed the number is correct for the Spruce spec-

imen, since other specimens of his, collected near

this date, bear numbers in this sequence. Appar-

ently mounters inadvertently copied this number

onto the Nelson label; this specimen may have

come from Ecuador (see Comments).

San Martin: Mt. Picote near Moyobamba, Nelson in

Dec. 1856 (K). Mt. Campana, near Tarapoto, Spruce

4643 in Aug. 1 856 (K). Pasco: Border between Provinces

Pasco and Oxapampa, San "Cotardo" (Gutardo), van

der Werffet al. 8584 (MO, uc).

2. Grammitis limbata Fee (Mem. foug. 5) Gen.

fil.: 233. 1852. TYPE: Guadeloupe, Perrotet,

in 1824 (holotype, P!). Figures 5b-c.

Grammitis nigrolimbata Hooker, Sp. fil. 4: 164. 1862.

in synon.

Polypodium nigrolimbatum Jenman, Bull. Bot. Dept..

'n.s. 4: 69. 1897.

Polypodium limbatum (Fee) Maxon, Bull. Torrey Bot.

Club 42: 222. 1915.

Grammitis nigrolimbata (Jenman) Lell., Proc. Biol.

Soc. Wash. 89:715. 1977.

Plants terrestrial. Stem erect to ascending, pro-

vided with bright orange or golden orange, lus-

trous, rigid, nonclathrate scales, these 2-3 mm long,

to 0.5 mm broad, 8-12 cells across at the base.

Leaves simple, entire to weakly sinuate-undulate,

5-16 cm long, 3-8 mm broad. Petiole light brown,

1-5 mm long, 0.2-0.4 mm in diameter. Lamina

linear to narrow-elliptic or -oblanceolate, erect,

firm-herbaceous to chartaceous, attenuate at base,

the apex acute or subacute, rarely apiculate, pro-

vided with a lustrous black or castaneous, scler-

enchymatous border usually less than 0.1 mm

broad, essentially glabrous but with scattered,

minute, caducous trichomes, especially on costa

and margin, these simple and subclavate, rarely

branched. Veins distinct or indistinct, diverging

from the costa at 30—40° angles, sterile ones sim-

ple, fertile ones commonly with a short spur bear-

ing the sorus, hydathodes evident adaxially. Sori

superficial, rounded to, usually, elongate, some-

times confluent.

In cloud forests, growing on trees, 750-910 m,

Pasco and Cuzco.

West Indies; Venezuela; Peru; Bolivia; Brazil.

Pasco: Prov. Oxapampa, west divide of Cordillera de

San Matias between Iscozacin and summit, D. Smith

2038 (F, MO). Cuzco: Prov. Paucartambo, Rio Tono Road,

Foster et al. 10619(f).

3. Grammitis paramicola Bishop, Amer. Fern J.

67: 105. 1977. TYPE: Colombia, Depto. Hui-

la, 25 km southeast of Gigante, Little 8663

(holotype, us!; photo, F).

Plants terrestrial. Stem erect to ascending, pro-

vided with deep orange, lustrous, rigid, nonclath-

rate scales, these 1-2 mm long, ca. 0.2-0.3 mm

broad, 2-6 cells across at the base. Leaves simple,

entire to slightly sinuate, 2-7 cm long, 1.5-3 mm

broad. Petiole light brown, 1-5 mm long, 0.2-0.4

mm in diameter. Lamina linear, or slightly broad-

ened distally, rigid, subcoriaceous, attenuate at

base, the apex obtuse to subacute, rarely cuspidate,

provided with a black or dark brown, lustrous,

sclerenchymatous border 0.1-0.2 mm broad, es-

sentially glabrous, but with scattered, caducous,

minute, subclavate trichomes especially on costa

and margin. Veins obscure, diverging from the

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

81

costa at a very acute angle, sterile ones simple,

fertile ones furcate, hydathodes lacking. Sori

roundish to elongate, confluent at maturity and

sometimes slightly immersed in the tissue.

In open paramos or elfin forests, in sphagnum

or on mossy hummocks, 2750-3500 m, San Mar-

tin, Huanuco, Cuzco.

Costa Rica; Colombia; Peru.

This is one of the three species from Peru in the

genus with dark, sclerenchymatous borders

(Grammitis s.s., fide Smith and Moran). In addi-

tion to the key characters, it differs also from G.

bryophila and G. limbata in its smaller, more rig-

idly erect leaves and by the more acute vein angle.

In G. paramicola veins diverge from the costa at

about 10-15°, whereas the angle is commonly SO-

SO0 in the other two species.

San Martin: Prov. Mariscal Caceres, Chochos, NW

corner of Rio Abiseo National Park, Young & Leon 4760

(USM). Huanuco: Playapampa, Macbride 4520 (F, us).

Cuzco: Prov. La Convention, Cordillera Vilcabamba,

Dudley 11061 (GH). Huadquina, Biies 1271 (us).

4. Grammitis pumila (C. Chr.) Stolze, comb. nov.

Cochlidium pumilum C. Chr., Dansk. Bot. Ark. 6(3):

19, t. 3,f.3. 1929. TYPE: Ecuador, Chuquiribam-

ba, Massee, in 1868 (holotype, K).

Plants epiphytic or terrestrial. Stem erect or as-

cending, provided with bright orange, lanceolate,

attenuate, nonclathrate scales, these 2-3 mm long.

Leaves simple, entire to distally repand, 4-15 cm

long, ca. 1 mm broad. Petiole brown to blackish,

2-5 mm long, 0.2-0.5 mm in diameter. Lamina

linear, subcoriaceous, attenuate at base, fertile only

in the distal portion, this often conduplicate, gla-

brous. Veins obscure, reduced to short spurs,

hydathodes indistinct to obscure. Sori elongate,

arranged in interrupted or continuous lines (coe-

nosori).

In elfin forests or somewhat open places, on or

at bases of trees, 2100-2600 m, Cajamarca and

Huanuco.

Venezuela; Colombia to Bolivia; Brazil.

Cajamarca: Prov. Cutervo, La Pucarilla, Socota to San

Andres, Lopez et al. 6717 (F, GH). Huanuco: SW slope

of Rio Llullapichis watershed, ascent of Cerros del Sira,

Dudley 134 54 A (GH), 13490 (GH).

5. Grammitis myosuroides (Sw.) Sw., J. Bot.

(Schrader) 1800(2): 18. 1801. Figure 5a.

Polypodium myosuroides Sw., Prodr. 131.1 788. TYPE:

Jamaica, Swartz (holotype, s; photo, us).

Xiphopteris myosuroides (Sw.) Kaulf., Enum. fil. 85.

1824.

Xiphopteris jamesonii Hooker, Sec. cent, ferns, /. 14.

1860. TYPE: Ecuador, Andes of Quito, Jameson

(holotype, K).

Polypodium serrulatum var. strictissimum Hooker, Sp.

fil. 4: 175. 1862, based on Xiphopteris jamesonii

Hooker.

Polypodium jamesonii (Hooker) Jenman, Bull. Bot.

Dept. 4: 112. 1897, nom. illeg., not (Fee) Salo-

mon, 1883.

Polypodium strictissimum (Hooker) Hieron., Bot.

Jahrb. Syst. 34: 501. 1904.

Grammitis jamesonii (Hooker) Morton, Contr. U.S.

Natl. Herb. 38: 96. 1967.

Lellingeria myosuroides (Sw.) Smith & Moran, Amer.

Fern J. 81: 85. 1991.

Plants epiphytic. Stem small, erect, provided

with grayish brown, clathrate, narrow-deltate, at-

tenuate scales, these ca. 1 mm long. Leaves partly

dimorphic, 3-10 cm long, 3-4 mm broad. Petiole

dark brown to castaneous, 1-4 mm long, 0.2-0.4

mm in diameter. Lamina linear, chartaceous, at-

tenuate at base, fertile only in the subentire distal

portion, sterile portion deeply pinnatisect with 1 5-

20 pairs of segments, these deltate and acute to

oblong and obtuse, provided with scattered, mi-

nute, persistent, castaneous, setiform trichomes to

0.5 mm long. Veins simple and solitary in each

sterile segment, black and extending at broad an-

gles beyond the sporangia in the fertile portion,

hydathodes indistinct or lacking. Sporangia borne

at the base of veins, initially in discrete sori, but

eventually confluent as 2 subcostal lines, merging

into a broad, single, costal line at maturity.

Known thus far in Peru by a single collection

from Cajamarca, edge of the road, 2700 m.

Mexico (Chiapas); Costa Rica; Jamaica; Cuba;

Puerto Rico; Venezuela; Colombia to Peru.

Besides the key characters this differs also from

G. serrulata in the blackish veins of the fertile

portion of the lamina, which spread at a very broad

angle from the costa and contrast strongly with the

laminar tissue. Fertile veins of G. serrulata spread

at an acute angle and contrast little with the lam-

inar tissue. The sole collection of G. myosuroides

in Peru was found growing together with speci-

mens of the latter and G. pumila.

Cajamarca: Prov. Cutervo, La Pucarilla, between San

Andres and Socota, Lopez et al. 6716 (GH, HUT).

6. Grammitis serrulata (Sw.) Sw., J. Bot. (Schrad-

er) 1800(2): 18. 1801.

82

FIELDIANA: BOTANY

AcrostichumserrulatumSw.,Prodr. 128. 1788. TYPE:

Jamaica, Swart z (holotype, s; frag., us; isotypes,

B, Herb. Willd. 19589-1 &-3; photos, GH of B).

Xiphopteris serrulata (Sw.) Kaulf., Wes. Farrnkr. 87.

1827.

Polypodium serrulatum (Sw.) Mett., Fil. hort. hot. Lips.

30. 1856, not. Sw. 1802.

Polypodium duale Maxon, Contr. U.S. Natl. Herb. 1 6:

61. 1912, nom. nov. for Acrostichum serrulatum

Sw. and with the same type.

Cochlidium serrulatum (Sw.) Bishop, Amer. Fern J.

68: 80. 1978.

Plants epiphytic, occasionally epipetric. Stem

ascending, provided with brown, narrow-deltate,

nonclathrate scales, these 2-3 mm long. Leaves

partly dimorphic, 3-8 cm long, to 2.5 mm broad.

Petiole very short or obsolete. Lamina linear, firm-

herbaceous, attenuate at base, fertile only in the

subentire, sometimes conduplicate, distal portion,

sterile leaf or portion serrate-dentate or with short,

acute lobes, glabrous, or with some pale and ca-

ducous trichomes on juvenile plants. Veins indis-

tinct to obscure, simple and solitary on each sterile

segment, in fertile portion ascending at an acute

angle and not contrasting strongly in color with

the laminar tissue, hydathodes vestigial or lacking.

Sporangia borne on costa and veins, becoming

confluent in a single costal line (coenosorus).

Polypodium trifurcatumL., Sp.pl. 1084. 1753. TYPE:

Plumier t. 138, Traite foug. Amer., illustrating a

Plumier specimen from Martinique.

Glyphotaenium trifurcatum (L.) Lellinger, Amer. Fem

J. 75: 31. 1985.

Enterosora trifurcata (L.) Bishop, Syst. Bot. 17: 353.

1992.

Plants epiphytic. Stem short-creeping to as-

cending, provided at the apex with dense clusters

of closely imbricate, yellow or yellow-brown scales,

these narrow-deltate, not or scarcely clathrate, 2-

4 mm long, remotely ciliolate. Leaves monomor-

phic, 10-27 cm long, 2-3.5 cm broad. Petiole 3-

8 cm long, commonly '/j-'/z the length of the lam-

ina, lustrous, blackish or atropurpureous, amply

to densely provided with spreading, castaneous,

unicellular trichomes, these 1-1.5 mm long. Lam-

ina subcarnose, lanceolate or elliptic-lanceolate,

obtuse or subacute at apex, cuneate at base, cut

'/3-'/2 to the costa, the lobes obtuse (rarely sub-

acute), the margins sparsely provided with tri-

chomes like those of the petiole, or glabrate. Veins

obscure, free or rarely anastomosing, pinnately

branched in each lobe, hydathodes lacking. Sori

round to elongate, borne at the tips of acroscopic

vein branches, in 3-5 irregular series on each side

of the costa, somewhat immersed in the tissue.

In wet forests or clearings, on trees or fallen logs,

325-1 800(-2700) m, Cajamarca, Loreto, and

Huanuco south to Cuzco.

Southern Mexico to Panama; West Indies; Co-

lombia to the Guianas, south to Bolivia and Brazil;

wet tropical Africa.

This is one of the most widespread species of

Grammitis in the Neotropics and one of the easiest

to distinguish. For further comparisons, see dis-

cussion under G. myosuroides.

Cajamarca: Prov. Cutervo, La Pucarilla, between San

Andres and Socota, Ldpez et al. 6715 (GH). San Martin:

Prov. Mariscal Caceres, Dist. Tocache Nuevo, Schunke

y. 7807 (MO). Loreto: Sierra del Pongo, Mexia 6287 (GH,

MO, us). Huanuco: Prov. Huanuco, Dist. Churubamba,

Pampa Hermosa, Mexia 8258a (F, GH). Pasco: Prov.

Oxapampa, west side of Cordillera de San Matias, be-

tween Iscozacin and summit, D. Smith 2041 (F, MO).

Junin: East of Quimiri Bridge near La Merced, Killip &

Smith 23886 (F, us). Cuzco: Valle de Cosnipata, between

Yaramayo and Santa Isabel, Scolnik 863 (uc, us).

Uncommon, in dense forests, usually arched or

pendent from tree trunks, 700-1900 m, Huanuco,

Pasco, Madre de Dios.

West Indies; Costa Rica and Panama; the

Guianas to Colombia; Ecuador to Bolivia.

Specimens of G. trifurcata from the West Indies,

especially Jamaica, commonly bear two different

kinds of indument: intermixed with the long,

spreading, unicellular, trichomes can be found

some scattered, minute, tortuous ones with 3—4

cells, the apical cell often glandular. No such tri-

chomes have been observed in the few specimens

found thus far in Peru. The character merits fur-

ther examination in future monographic study.

Huanuco: SW slope of Rio Llullapichis watershed, as-

cent of Cerros del Sira, Dudley 13084b (GH). Pasco (as

Junin): Pichis Trail, Dos de Mayo, Killip & Smith 25803

(F, us). Pichis Trail, Porvenir, Killip & Smith 25894 (NY,

us). Madre de Dios: Prov. Manu, Cerro de Pantiacolla,

Rio Palotoa, Foster et al. 10847 (F, USM).

7. Grammitis trifurcata (L.) Copel., Gen. fil. 21 1.

1947.

8. Grammitis trichosora (Hooker) Ching, Bull.

Fan. Mem. Instil. Biol., Bot. 10: 241. 1941.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

83

Polypodium trichosorum Hooker, Sec. cent, ferns, /.

12. 1860. TYPE: Ecuador, Archidona, Andes of

Quito, Jameson 349 (holotype, K.!; photos, F, us).

Enterosora trichosora (Hooker) Bishop, Syst. Bot. 1 7:

357. 1992.

Plants epiphytic. Stem short-creeping or repent,

provided at the apex with dense clusters of closely

imbricate, pale yellow scales, these narrow-deltate,

nonclathrate, 1-2 mm long, entire to remotely

denticulate or ciliolate. Leaves monomorphic, 8-

18 cm long, 1-1.5 cm broad. Petiole 1-5 cm long,

0.4-0.8 mm in diameter, yellowish to reddish

brown, densely covered with spreading, casta-

neous, unicellular trichomes 2 mm long. Lamina

firm-herbaceous, elliptic to oblanceolate, apex

subacute, base cuneate, margins subentire to re-

mand or broadly and shallowly lobed, abundantly

provided with persistent trichomes like those of

the petiole. Veins obscure, pinnately branched, free

to irregularly anastomosing especially away from

the costa, hydathodes lacking. Sori round or slight-

ly elongate, in 1-2 irregular series on each side of

the costa, not or scarcely immersed in the tissue.

Thus far known in Peru by a single collection

from San Martin, epiphyte in montane rain forest,

2100 m.

Ecuador and Peru.

Very similar is Polypodium ecostatum Sodiro

(Central America; Antilles; northern South Amer-

ica). However, the latter is a much longer fern of

lower elevations (under 1 000 m), with sori rather

deeply immersed in the subcarnose tissue. Also,

the veins frequently anastomose along the costa,

whereas areoles of G. trichosora are quite irregular,

occurring infrequently along the costa, and are

much more common about midway between costa

and margin.

San Martin: Prov. Rioja, Venceremos, Pedro Ruiz-

Moyobamba Road, D. Smith & Vdsquez 4727 (MO, USM).

9. Gram mil is parietina (Klotzsch) Fee (Mem. foug.

5) Gen. fil. 233. 1852.

Polypodium parietinum Klotzsch, Linnaea 20: 373.

1847. TYPE: Venezuela (as "Columbia"), Colo-

nia Tovar, Moritz 253 (holotype, B!; isotype, BM;

photo, F of B).

Polypodium sprucei Hooker, Sec. cent, ferns, t. 10.

1 860. TYPE: Peru, San Martin, Tarapoto, Spruce

4746 (holotpye, K!; isotype, us!).

Grammitis sprucei (Hooker) John Sm., Hist. fil. 181.

1875.

Enterosora parietina (Klotzsch) Bishop, Syst. Bot. 17:

357. 1992.

Plants epiphytic. Stem small, ascending, pro-

vided at apex with a few inconspicuous, deep or-

ange scales, these entire or ciliolate, obscurely

clathrate, 0.5-1 mm long. Leaves monomorphic,

4-9 cm long, 0.4-0.8 cm broad. Petiole to 0.7 cm

long or obsolete, 0.4-0.6 mm in diameter, brown

to blackish, amply provided with spreading, cas-

taneous, unicellular trichomes about 1 mm long.

Lamina subcarnose to chartaceous, oblong-ellip-

tic, apex obtuse or subacute, base attenuate, mar-

gins subentire to sinuate or crenate, amply pro-

vided with trichomes like those of the petiole. Veins

obscure, free, commonly 1 -forked, hydathodes

lacking. Sori roundish, in a single series on each

side of the costa, superficial or slightly immersed

in the tissue.

In cloud forests, on tree trunks, 1900-2400 m,

Amazonas, Pasco, San Martin and Cuzco.

Southern Mexico; Guatemala; Costa Rica and

Panama; Jamaica; Hispaniola; Venezuela; Colom-

bia to Peru.

The species thus far is represented in Peru by

only three widely scattered collections. Because of

its small size and inconspicuous leaves it has likely

been overlooked, and a more diligent search should

reveal it to be in intervening localities.

Amazonas: Prov. Bagua, 12-20 km E of La Peca, Bar-

hour 2723 (F). Pasco: Prov. Oxapampa, San Alberto,

Cordillera de Yanachaga, van der Werffel al. 8496 (MO).

Cuzco: Prov. La Convencion, Cordillera Vilcabamba,

Dudley 11 302 (GH).

10. Grammitis phalacron Stolze, sp. nov.

Caulis breve gracilisque, paleis 1-1.5 mm longis,

castaneis, setosis, angusti-deltatis, non clathratis.

Folia 7-12 cm longa, 0.6-0.8 cm lata. Petiolus

1-2 cm longus, trichomatibus circa 2mm longis,

aurantiacis, unicellularibus. Lamina anguste el-

liptica, subintegra vel repanda, glabrata vel tri-

chomatibus 0.2-0.4 cm longis, pallescentibus,

tortuosis, pluricellularibus. Hydathodi nulli. Sori

rotundi, discreti, praphysibus glandulis.

Plants probably epiphytic. Stem short, slender,

provided with castaneous, narrow-deltate, setose,

nonclathrate scales, these 1-1.5 mm long. Leaves

monomorphic, 7-12 cm long, 0.6-0.8 cm broad.

Petiole 1-2 cm long, 0.2-0.4 mm in diameter, dark

brown to blackish, amply provided with spread-

ing, orange, unicellular trichomes, these to 2 mm

84

FIELDIANA: BOTANY

long. Lamina firm-herbaceous to chartaceous, nar-

row-elliptic, apex obtuse to subacute, cuneate at

base, margins subentire or repand to very broadly

and shallowly lobed, abaxially provided with scat-

tered, pale, tortuous, pluricellular trichomes, these

0.2-0.4 mm long, caducous. Veins indistinct, 1-

2-forked to pinnate, hydathodes lacking. Sori

round, borne on basal vein branches and forming

a single series on each side of the costa, paraphyses

glandular.

TYPE— Peru, Cuzco, Valley of Rio Urubamba,

Bues A29 (holotype, us!).

Known only from the type.

In general aspect, this appears to be interme-

diate between G. parietina and Polypodium

(Grammitis) ecostatum Sodiro (Ecuador). How-

ever, the completely different indument, the glan-

dular paraphyses and the thinner textured lamina

(vs. laminae with spongy parenchyma) argue

against such affinity. The specimen was annotated

by Bishop and Smith as Ceradenia phalacron, with

intent to publish at a later date. Their kind per-

mission for publication as Grammitis in this flora

is greatly appreciated.

1 1. Grammitis longipinnata (Copel.) Loll.. Amer.

Fern J. 74: 58. 1984.

Ctenopteris longipinnata Copel., Philipp. J. Sci. 84:

459. 1956. TYPE: Peru, Cuzco, Cabeceradel Ko-

ribeni, Bues 1952 (holotype, us!).

Ceradenia longipinnata (Copel.) Bishop, Amer. Fern

J. 78: 5. 1988.

Plants probably epiphytic. Stem unknown.

Leaves to 30 cm long and 10 cm broad. Petiole

8-10 cm long, 1 mm in diameter, blackish, sub-

lustrous, glabrescent. Lamina chartaceous to sub-

coriaceous, subdeltate, not or scarcely reduced at

base, deeply pinnatisect to fully 1 -pinnate toward

the base, rachis amply provided with spreading,

castaneous trichomes to 1 mm long; pinnae falcate

and somewhat ascending, linear-deltate, acute to

attenuate, margins with scattered, castaneous tri-

chomes and deeply serrate, but distal ones merely

crenulate or entire. Veins obscure, free, 1 -forked,

hydathodes lacking. Sori round to slightly elon-

gate, in a single supramedial to submarginal series

on each side of the costa.

Endemic. Thus far known from the type and

two other collections from Cuzco, 2750-3250 m.

The Dudley specimen was pendulous from trees

in a dark, wet, cloud forest.

Nothing is known of the stem characteristics of

this species, since leaves of the only known spec-

imens are broken off at the petiole base. No habit

was mentioned on labels of the type and the other

Biles collection.

Cuzco: Valle de Lares, Bues 1793 (us). Prov. La Con-

vention, 25 km from Hacienda Luisiana and the Apu-

rimac River, Dudley 1 1 106 (us).

12. Grammitis meridensis (Klotzsch) Seymour,

Phytologia 31: 179. 1975.

Polypodium meridense Klotzsch, Linnaea 20: 380.

1847. TYPE: Venezuela, Edo. Merida, Merida,

Moritz 335 (holotype, B!; photo, F).

Ctenopteris meridensis (Klotzsch) Copel., Philipp. J.

Sci. 84: 464. 1956.

Ceradenia meridensis (Klotzsch) Bishop, Amer. Fern

J. 78: 5. 1988.

Plants terrestrial, at least in Peru. Stem short-

creeping to ascending, abundantly provided with

linear, rigid, lustrous, deep orange to castaneous

scales, these (2-)3-6 mm long, nonclathrate, am-

ply setulose, the setae whitish. Leaves monomor-

phic, articulate and approximate on the stem, 20-

40(-50) cm long, 3-7 cm broad. Petiole 8-30 cm

long (usually as long as the lamina), 0.4-1 mm in

diameter, lustrous, castaneous to atropurpureous

or blackish, sparsely to amply pilose, or glabres-

cent at maturity, the trichomes castaneous.

spreading, unicellular, 0.4-1 mm long. Lamina

firm-herbaceous to chartaceous. oblong or oblong-

deltate, truncate at base, deeply pinnatisect, or

rarely 1 -pinnate at base, rachis moderately pro-

vided on both sides with spreading, castaneous,

trichomes to 1 mm long. Segments patent or slightly

ascending, linear-deltate, acute or subacute, tri-

chomes like those of the rachis. sparse along the

margins and sometimes on the abaxial surface,

especially among the sori. Veins completely ob-

scure, commonly once-forked, hydathodes lack-

ing. Sori superficial, round, supramedial to sub-

marginal, immature sporangia intermixed with

pale, waxy paraphyses.

In dense, wet forests or wooded rav ines. on clay

banks or in humus, 1500-3600 m. Huanuco to

Cuzco.

Costa Rica; Panama; Venezuela and Colombia

south to Brazil and Bolivia.

This species is especially distinguished by the

TRYON & STOLZE. PTERIDOPHYTA OF PERU. V.

85

long (to 1 mm), spreading, dark, unicellular tri-

chomes on both sides of the rachis and by the

truncate lamina base with basal pinnae longer or

nearly as long as medial ones. Very similar, and

probably conspecific, are Polypodium spixianum

Mett. of Brazil and P. abitaguae Hooker of Ec-

uador. Both apparently differ from G. meridensis

in the lighter orange color and greater length (to 9

mm) of stem scales, and sori of P. abitaguae are

medial rather than supramedial or submarginal.

Some specimens have been determined as P. spixi-

anum on the basis of slightly thinner lamina tex-

ture and nearly obtuse segment apices, but both

characters are too variable to be significant. This

condition seems to pertain to juvenile and/or de-

pauperate plants, while thicker texture and longer,

acute segments are common to fully mature leaves.

Huanuco: Cani, pueblo 7 mi NE of Mito, Macbride

3491 (F, us). Vilcabamba, Macbride 5145 (F in part, us).

Pasco: Pichis Trail, Porvenir (as Junin), Killip & Smith

25939 (F, NY, us). Prov. Oxapampa, Dist. Chontabamba,

near the summit of San Gutardo, Leon et al. 524 (USM,

juvenile specimens), 526 (F, USM). Junin: Chanchamayo

Valley, C. Schunke 853 (F), 1380 (F). Cuzco: Prov. La

Convencion, Valle San Miguel, Biles 2133 (us). Prov.

Paucartambo, near El Mirador, Manu National Park,

Leon 2265 (F, USM). Prov. Paucartambo, Cordillera de

Tres Cruces, Vargas 12191 (GH).

13. Grammitis bishopii Stolze, sp. nov.

Plantae terrestres. Caulis repens, 1.5-3 mm crassus,

paleisabundantibus, linearibus, rigentibus, niten-

tibus, castaneis, 2-4 mm longis, 0.2-0.4 mm latis,

setulosis, setulis concoloribus. Folia 18-34 cm

longa, 4-10 cm lata, subdistantia, ad phyllopodia

bulbosa affixa. Petiolus 10-24 cm longus, nitidus,

castaneus vel atropurpureus, glaber. Lamina

1-pinnata, deltata, basi non reducta. Rhachis et

costa trichomatibus minutis, castaneis, unicel-

lularibus, abundantibus praeditae. Venae ob-

scurae, 1-2-furcatae, raro anastomosantes. Hy-

dathodi nulli.

Plants terrestrial. Stem long-creeping, 1.5-3 mm

in diameter, densely clothed with linear, rigid, lus-

trous, castaneous scales, these 2-4 mm long, 0.2-

0.4 mm broad, nonclathrate, sparingly to amply

setulose, the setae concolorous with the scale body.

Leaves monomorphic, borne on short (1-2 mm),

often bulbous phyllopodia, usually 1-2 cm apart,

18-34 cm long, 4-10 cm broad. Petiole 10-24 cm

long, 1-2 mm in diameter, lustrous, castaneous to

atropurpureous, essentially glabrous. Lamina sub-

coriaceous, narrowly or broadly deltate, not or

scarcely reduced at base, 1 -pinnate, the segments

adnate, slightly ascending to slightly reflexed, lin-

ear, acute, their margins often with minute, scat-

tered, glandular trichomes, the rachis adaxially with

dense, minute, castaneous, rigid, unicellular tri-

chomes, these 0.1-0.3 mm long, costae with sim-

ilar, but shorter and fewer, trichomes. Veins

completely obscured (even when held to light), 1-

2-forked, often connivent distally, on some spec-

imens frequently anastomosing, hydathodes lack-

ing. Sori superficial, commonly round, essentially

medial between (and at maturity often crowding

both) costa and margin, paraphyses not seen.

TYPE— Peru, Pasco, Prov. Oxapampa, border

Prov. Oxapampa and Pasco, van der Werff et al.

8569 (holotype, uc!; isotype, MO!).

In elfin or cloud forests, in clay or sphagnum,

2700-3000 m, Amazonas, Pasco and Cuzco.

Peru; Bolivia (Tate 349 [NY]).

This may be confused with G. werffii. In addition

to the characters used in the key, this differs in the

rarely (vs. frequently in the latter) anastomosing

veins and in the short, rather swollen, phyllopo-

dium that joins the petiole base to the stem. In G.

werffii the petiole base is articulate and the line of

abscission very evident.

The species is named in honor of the late L. Earl

Bishop, who recognized this as a new species and

tentatively labeled several of the specimens as

"Ceradenia vaga," but until now it has remained

undescribed.

Amazonas: Prov. Bagua, Cordillera Colan, E of La

Peca, Barbour 3264 (MO, USM). Pasco: Prov. Oxapampa,

Dist. Chontabamba, near the summit of San Gutardo,

Leon et al. 526 (USM). Prov. Oxapampa, Cerro Pajonal,

29 km from Oxapampa, D. Smith & Foster 2504 (uc).

Border, Prov. Oxapampa and Pasco, below San Gutardo

(as "Cotardo"), van der Werff et al. 8550 (MO). Cuzco:

Prov. La Convencion, Cordillera Vilcabamba, Dudley

10892 (F, GH), 11145 (GH, US).

14. Grammitis discolor (Hooker) Morton, Amer.

Fern J. 60: 66. 1970.

Polypodium discolor Hooker, Icon. pi. 4: /. 386. 1841.

TYPE: "British Guiana" (Guyana), Schomburgk

1031 (holotype, K; photo, F).

Ctenopteris discolor (Hooker) John Sm., Hist. fil. 185.

1875.

Ceradenia discolor (Hooker) Bishop, Amer. Fern J.

78:4. 1988.

Plants probably terrestrial. Stem stout, erect,

provided with linear, brown scales, these 3-6 mm

long, nonclathrate, entire to sparsely and remotely

setulose. Leaves not articulate, 9-20 cm long, 1-

2 cm broad. Petiole essentially obsolete. Lamina

86

FIELDIANA: BOTANY

subcarnose, linear to narrow-elliptic, strongly and

gradually reduced at base, deeply pinnatisect (about

3/4 to the rachis) the segments 1-2 times as long as

broad, rachis rarely evident (usually immersed in

the tissue), rachis and tissue lacking trichomes,

moderately to densely farinose, the farina white,

mostly disposed in stellate patterns. Veins ob-

scure, free, pinnately branched in the segments,

hydathodes lacking. Sori round, submarginal, with

farina deposited among the sporangia.

In dense forests, apparently terrestrial (habit not

cited on any of the labels seen, but clay particles

evident among the roots), 1 300-2000 m, thus far

found only in Junin.

Guyana; Venezuela; Colombia; Peru; Bolivia.

This is very similar to G. curvata ssp. pearcei,

differing only in the farina pattern and the shape

of segments and may represent merely another

infraspecific variant of the latter.

Junin: Schunke Hacienda above San Ramon, C.

SchunkeA152(GH, us). Chanchamayo Valley, C. Schunke

470 (F, us), 513 (F, us).

15. Grammitis curvata (Sw.) Ching, ssp. pearcei

(Baker) Stolze, stat. & comb. nov.

Polypodiutn pearcei Baker, in Hooker and Baker, Syn.

fil., ed. 2: 508. 1874. LECTOTYPE (designated

here): Ecuador, Cuesta de Puente Grande, Pearce

in June 1865 (K!, photo, F, isolectotype, 2 laminae

on the right, K!)

Ctenopteris amylacea Copel., Philipp. J. Sci. 84: 469.

1956. TYPE: Bolivia, Cargadira, R. S. Williams

1134 (holotype, us!; isotypes, GH!, uc!).

Grammitis amylacea (Copel.) Morton, Contr. U.S.

Natl. Herb. 38: 233. 1973.

Ceradenia pearcei (Baker) Bishop, Amer. Fern J. 78:

4. 1988.

Plants epiphytic or epipetric. Stem stout, erect,

provided with linear, orange-brown to ferruginous

scales, these 3-6 mm long, nonclathrate, sparsely

to amply setose, the setae pale orange or whitish.

Leaves not articulate, 12-30(-35) cm long, (2.5-)

3-8 cm broad. Petiole essentially obsolete. Lamina

subcarnose, rachis and costae rarely evident (usu-

ally immersed in the tissue), narrowly to broadly

elliptic, strongly and gradually reduced at base,

deeply pinnatisect, the segments 3-7 times as long

as broad, rachis and tissue lacking trichomes,

moderately farinose, the farina particles white (of-

ten brown on dried specimens), solitary or dis-

posed in small, amorphous clusters. Veins ob-

scure, free, once-forked from the costa, hydathodes

lacking. Sori round, borne at the tip of the acros-

copic vein branch, submarginal, at maturity ex-

tending beyond the segment margin, with farina

deposited among the sporangia.

In scrub and cloud forests, on tree trunks or in

crevices of rocky cliffs, 2000-2750 m, Amazonas,

Huanuco, Cuzco.

Peru; Bolivia.

This differs from ssp. curvata (West Indies) in

the longer and more abundant setae on the stem

scales. Furthermore, ssp. curvata has minute, ap-

pressed (sometimes glandular?) trichomes scat-

tered on the lamina, in addition to the farina de-

posits. These trichomes are 0.1-0.2 mm long,

cylindrical, castaneous, and are often quite abun-

dant, though very inconspicuous. Such processes

have not been seen in specimens of ssp. pearcei.

Also very similar (and likely another subspecies)

is Grammitis albidula (Baker) Morton, from

southeastern Brazil, but the latter has smaller leaves

and supramedial (vs. submarginal) sori.

There are two sheets of Pearce specimens in the

type folder at Kew. One has the stem and several

leaves and bears a printed label marked "Type

Specimen." The other sheet has two laminae with-

out a stem, and a third lamina from another Pearce

collection with the number 279 and "holotype"

handwritten in blue ink. The two stemless laminae

were annotated by Bishop (1988) as lectotype, but

that typification has not been published. The other

sheet is a much preferable lectotype, since the plant

is complete and not mixed, and is so designated

here.

Amazonas: Prov. Bagua, E of La Peca, Barbour 2767

(F, uc). Prov. Chachapoyas, west of Molinopampa, Wur-

dack 1499 (us), 1543 (us). Huanuco: Cushi, trail to Tam-

bo de Vaca, Bryan 686 (F, us). Cuzco: Prov. Paucartam-

bo, Manu Park, between Pillahuata and Pilcopata, Skog

&Skog5198(vs).

16. Grammitis terrestris (Bishop) Stolze, comb,

nov.

Ceradenia terrestris Bishop, Amer. Fern J. 79: 23.

1989. TYPE: Peru, Amazonas, Prov. Chacha-

poyas, south of Molinopampa-Diosan Pass,

Wurdack 1643 (holotype, us!; isotype, GH!).

Plants terrestrial. Stem stout, erect, provided

with linear-deltate, deep orange to castaneous,

nonclathrate scales, these 3-5 mm long, moder-

ately to amply setulose. Leaves not or scarcely

articulate, 8-18 cm long, 1-2 cm broad, lacking

long, spreading unicellular trichomes. Petiole 1-2

cm long, 0.6-0.9 mm in diameter, minutely glan-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. V.

87