DEC 12

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Botany

NEW SFK

PTERIDOPHYTA OF PERU

Part II

13. Pteridaceae-15. Dennstaedtiaceae

Rolla M. Tryon

Robert (;. Stol/e

SSm

September 29, 1989

Publication 1403

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

•nation for Contributors to Fieldiana

published by Tl

.raged to give journal and booi-

m (1976 et seq.) (botanical pap

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^parison of montane

nd floristics. Journal 51: 567-

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in 143. Bureau of Am

Part II. IV

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THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

FIELDIANA

Botany

NEW SERIES, NO. 22

PTERIDOPHYTA OF PERU

Part II

13. Pteridaceae-15. Dennstaedtiaceae

Rolla M. Tryon

Gray Herbarium

Harvard University

22 Divinity Avenue

Cambridge, Massachusetts 02138

Robert G. Stolze

Associate Curator. Pteridology

Department of Botany

Field Museum of Natural History

Roosevelt Road at Lake Shore Drive

Chicago, Illinois 60605-2496

Accepted June 24, 1988

Published September 29, 1989

Publication 1403

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 1989 Field Museum of Natural History

Library of Congress Catalog Card Number: 88-84027

ISSN 00 15-0746

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

List of Illustrations

INTRODUCTION 1 1 .

ACKNOWLEDGMENTS 1 2.

1 3. PTERIDACEAE 2 3.

Eriosorus 3 4.

Jamesonia 8

Pterozonium 14 5.

Pityrogramma 16 6.

Anogramma 22

Cheilanthes 23 7.

Adiantopsis 34 8.

Notholaena 37 9.

Pellaea 40 10.

Doryopteris 43 11.

Hemionitis 46 12.

Trachypteris 48 13.

Ceratopteris 50 14.

Adiantum 52

Pteris 70 15.

Acrostichum 81 16.

14. VlTTARIACEAE 83

Hecistopteris 84 17.

Antrophyum 84 18.

Vittaria 89 19.

Anetium 92

15. DENNSTAEDTIACEAE 94 20.

Microlepia 95 21.

Dennstaedtia 95 22.

Saccoloma 101

Pteridium 105 23.

Paesia 106 24.

Hypolepis 106

Blotiella Ill 25.

Lonchitis 113 26.

Histiopteris 115 27.

Lindsaea 115 28.

MAP OF PERU 1 24 29.

INDEX TO NAMES . .125 30.

Eriosorus: E. rufescens; E. flexuosus . . 5

Jamesonia alstonii 10

Pterozonium reniforme 15

Pityrogramma: P. tartarea; P. calome-

lanos 17

Anogramma leptophylla 22

Cheilanthes: C. poeppigiana; C. myrio-

phylla 26

Adiantopsis radiata 35

Notholaena: N. ni\ea; N. sulphurea ... 39

Pellaea: P. ternifolia; P. ovata 42

Doryopteris pedata var. palmata 45

Hemionitis palmata 47

Trachypteris: T. pinnata; T. induta ... 49

Ceratopteris pteridoides 51

Adiantum: A. concinnum; A. capillus-ve-

neris\ A. pulvrulentum 53

Pteris: P. biaurita; P. altissima 71

Acrostichum: A. aureum; A. danaeifol-

ium 82

Hecistopteris pumila 85

Antrophyum lineatum 86

Vittaria: V. graminifolia\ V. lineata; V.

stipitata 88

Anetium citrifolium 93

Microlepia speluncae 96

Dennstaedtia: D. dissecta; D. bipinnata;

D. globulifera 97

Saccoloma: S. inaequale; S. elegans ... 102

Pteridium aquilinum var. arachnoideum

104

Paesia glandulosa 107

Hypolepis parallelogramma 108

Blotiella lindeniana 112

Lonchitis hirsuta 114

Histiopteris incisa 116

Lindsaea: L. lancea; L. stricta; L. arcu-

ata\ L. divaricata 119

ui

PTERIDOPHYTA OF PERU

Part II

13. Pteridaceae-15. Dennstaedtiaceae

Introduction

This second part of "Pteridophyta of Peru" con-

tains treatments of the Pteridaceae, Vittariaceae,

and Dennstaedtiaceae, including a total of 30 gen-

era and 171 species known to occur in Peru. It

represents a considerable revision of "The Ferns

of Peru (Dennstaedtiae to Oleandreae)" by Rolla

Tryon (Contr. Gray Herb., 194: 1-253. 1964), due

to newly published information and numerous

collections made in Peru in the last 25 years. These

collections have mostly been made under the aus-

pices of the Universidad Nacional de Trujillo, the

Museo de Historia Natural de Universidad de San

Marcos, the Missouri Botanical Garden, and Field

Museum of Natural History. The treatment has

been prepared by Rolla Tryon aided by the critical

review of Robert G. Stolze.

Prior to 1 944 the Department of Pasco was a

part of Junin and until recently Ucayali was a part

of Loreto. An attempt has been made to account

for these changes, and this is not difficult when

labels cite towns and provinces. However, on older

specimens the labels often contain only sketchy

data, making it impossible to determine in which

part of Loreto or Junin a plant was collected. The

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

partments and indicates the sequence of the col-

lection citations.

New names published here are indicated by

boldface in the Index to Names. Collections have

been cited from each department of Peru from

which material has been seen and further collec-

tions are sometimes cited to include other her-

baria. In general, all collections seen are cited for

rare species and a selection is cited for common

species. Type collections, mentioned in the no-

menclature, are not included in the specimen ci-

tations, although they are included in the Peruvian

range and ecology.

Appended to some of the generic treatments are

portions of text labeled Comments. Herein are in-

cluded species to be expected in Peru, names based

on Peruvian material but of uncertain application,

excluded species (erroneous reports that may have

special significance), and cultivated species that

are possibly adventive.

Besides the original drawings, a number of plates

published in Fieldiana for "The Ferns and Fern

Allies of Guatemala" (Fieldiana, Bot., n.s., 6. 198 1)

have been used for species occurring in Peru.

Voucher specimens cited in the captions are from

Peru unless otherwise indicated.

Abbreviations of periodical publications gen-

erally follow the system of Botanico-Periodicum-

Huntianum (1968), and abbreviations of authors'

names and of books generally follow TL-2 Taxo-

nomic Literature (2nd ed.) by Stafleu and Cowan

(1976 etseq.}.

Acknowledgments

The treatments of the genera Eriosorus and Ja-

mesonia have been prepared with the collabora-

tion of Alice F. Tryon, and the treatment ofLind-

saea has been reviewed by Karl U. Kramer. The

authors appreciate this aid which has improved

the treatments of these technical genera. Special

thanks are extended to Field Museum's scientific

illustrators Zorica Dabich and Clara Richardson

and to volunteer artist Julia A. Liesse for their

artwork, which adds an important dimension to

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

the generic treatments. The collaboration of the

staff and officers of the following institutions has

been especially valuable in making loans available

for study and during visits: Field Museum of Nat-

ural History (F); Harvard University Herbaria,

most Gray Herbarium (GH) and some Arnold Ar-

boretum (A); Herbarium Truxillense, Universidad

de Trujillo, Trujillo, Peru (HUT); Missouri Bo-

tanical Garden, St. Louis, (MO); University of

California, Berkeley (UC); United States National

Herbarium, Smithsonian Institution, Washington,

D.C. (US); and Museo de Historia Natural "Javier

Prado" de Universidad Nacional Mayor de San

Marcos, Lima, Peru (USM).

Earlier work on "The Ferns of Peru," which

provided a considerable basis for the present treat-

ment, was supported by National Science Foun-

dation Grants 1064, 15949, and N-1565 to Rolla

M. Tryon. The present work is currently supported

in part by National Science Foundation Grant BSR-

8516358 to Rolla M. Tryon and Robert G. Stolze.

We appreciate this financial aid which provides

essential assistance. However, any opinions and

conclusions expressed are those of the authors and

do not necessarily reflect the views of the Foun-

dation.

Family 13: PTERIDACEAE

Pteridaceae Reichenb., Hand. Nat. Pfl.-Syst. 138.

1837, as Pteroideae. TYPE: Pteris L.

Stem erect to long-creeping, bearing trichomes

or scales or both. Leaves usually small to large,

ca. 1-6 m long, circinate in vernation, monomor-

phic to dimorphic, usually pinnate, or entire, pin-

nat i tki. pedate, palmate, ternate, or radiate, or the

petiole furcate apically into two strongly recurved

rachises, glabrous, glandular, pubescent, farinose

and (or) scaly. Petiole lacking stipules, not artic-

ulate to the stem. Veins free or fully anastomosing,

the areolae without included veinlets. Sori at the

vein ends, or the sporangia in short to long soral

lines along the veins, or the sporangia on a mar-

ginal commissure, or scattered on the veins, or on

anastomosing veins and between them, exindu-

siate or the recurved margin modified as an in-

dusium. Sporangia with a 2-3-rowed stalk, an-

nulus vertical or rarely oblique, interrupted by the

stalk.

The Pteridaceae are a family of about 33 genera

and 750 species, widely distributed through most

of the World. There are 1 6 genera in Peru. The

family is diverse, including undoubtedly old and

distinctive evolutionary lines, as well as recent and

poorly defined groups. It is characterized by a

chromosome number of n = 29 or 30 or multiples

(there are exceptions), trilete spores, and the spo-

rangia in exindusiate sori or soral lines, or covered

by a marginal, variously modified indusium. Pter-

idaceae is used here since the earlier Parkeriaceae

is always used for a monogeneric family, including

Ceratopteris.

In the following key to genera, the characters

are sometimes restricted, for the sake of conve-

nience, to Andean species.

Reference

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

daceae, pp. 213-354, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Genera of Pteridaceae

a. Margins of the fertile segments not modified, or somewhat modified but not or hardly concealing

the sporangia b

b. Sporangia borne on the anastomosing veins and between them c

c. Sterile lamina densely scaly abaxially XII. Trachypteris

c. Sterile pinnae glabrous to slightly pubescent abaxially XVI. Acrostichum

b. Sporangia borne only on the veins, on a vascular receptacle, or on a vascular commissure . . . d

d. Stem scarcely developed, not bearing leaf bases from previous years V. Anogramma

d. Stem well developed and bearing leaf bases from previous years e

e. Lamina simple, entire III. Pterozonium

e. Lamina deeply lobed or 1 -pinnate or more complex f

f. Stem bearing only trichomes or bristles g

FIELDIANA: BOTANY

g. Pinnae with several lobes, or pinnatifid or more complex; the segment margins not

or hardly enrolled; the lamina usually clearly determinate I. Eriosorus

g. Pinnae entire; the margins strongly enrolled; the lamina usually indeterminate ....

II. Jamesonia

f. Stem bearing only scales, or trichomes also sometimes present h

h. Two or more vascular bundles near the base of the petiole; the lamina farinose,

glandular, densely pubescent, or glabrous abaxially IV. Pityrogramma

h. One vascular bundle near the base of the petiole, or if 2, then the lamina densely

scaly abaxially i

i. Sporangia borne on the veins, entirely or nearly from the costa to the margin;

segments thinly to moderately pubescent abaxially XI. Hemionitis

i. Sporangia borne at the vein ends, or extending on the veins about halfway to the

I costa (rarely more and then glabrous or farinose abaxially), or on a complete or

incomplete marginal commissure j

j. Lamina densely pubescent or scaly abaxially VI. Cheilanthes

j. Lamina glabrous or farinose abaxially k

k. Lamina farinose abaxially, or if not, then glabrous and the petiole and rachis

very dark reddish to atropurpureous and the pinnae stalked

VIII. Notholaena

k. Lamina not farinose, glabrous abaxially; the petiole and rachis straw-colored

to ruddy brown, or if atropurpureous or darker, then the pinnae sessile . .

IX. Pellaea

a. Margins of the fertile segments modified as an indusium, mostly or wholly concealing the sporangia

1

1. Veins extending into the indusium where they bear the sporangia XIV. Adiantum

1. Indusium without veins or sporangia m

m. Paraphyses (trichomes unlike other lamina indument) present among the sporangia

XV. Pteris

m. Paraphyses absent (immature sporangia may be present) n

n. Sporangia borne separately on the veins; plants aquatic or semiaquatic; leaves dimorphic,

the sterile lamina thin, with evident anastomosing veins XIII. Ceratopteris

n. Sporangia adjacent, in sori or soral lines, rarely single at a vein end; plants terrestrial or

rupestral o

o. Lamina pinnate, it and the pinnae with a rather large, entire, stalked terminal segment

nearly as large as the adjacent ones, or pinnae mostly entire or ternate . . IX. Pellaea

o. Lamina pedate, ternate, or radiate, or usually pinnate and the lamina and pinnae with

a pinnatifid apex, or the pinnae pinnately lobed to pinnatifid, or the terminal segment

minute, ca. 1 mm long p

p. Lamina pedate; petiole with 2 vascular bundles near the base and veins free, or with

1 bundle and veins anastomosing, at least along the costa X. Doryopteris

p. Lamina pinnate or radiate, or pedate and then the petiole with 1 vascular bundle

near the base and the veins free q

q. Lamina pedate, or usually pinnate, and the petiole terete or adaxially concolor-

ously (or nearly) 2-ridged; spores cristate, verrucate, granulate, or rugose

VI. Cheilanthes

q. Lamina ternate or radiate, or pinnate and the petiole adaxially with 2 thin, light-

colored ridges; spores echinate VII. Adiantopsis

I. Eriosorus Terrestrial. Stem decumbent or usually creeping

and slender, bearing trichomes or bristles (very

Eriosorus Fee, (Mem. foug. 5) Gen. fil. 1 52, t. IBB, rarely scales). Leaves usually rather closely spaced.

/ 1. 1852. TYPE: Eriosorus scandens Fee = Lamina 1-4- or 5-pinnate (rarely to 6-pinnate),

E. aureonitens (Hooker) Copel. Figure 1. sparsely to usually definitely pubescent, some-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II. 3

times densely so, especially abaxially, veins free.

Sori rather elongate along the veins, usually not

extending to the vein tip, not paraphysate, exin-

dusiate. Spores tetrahedral-globose, trilete, with a

usually prominent equatorial flange, tuberculate

or coarsely ridged, with a basal triangular ridge.

Eriosorus may be separated from the very closely

related Jamesonia by the usually determinate lam-

ina, the stalked or rarely sessile pinnae, and the

not or hardly enrolled margins of the pinnae. Ja-

mesonia has the lamina indeterminate, or if de-

terminate, the pinnae adnate, and the margins of

the pinnae enrolled. Both genera have the unusual

feature of basally slender petioles.

There is clear morphological and cytological

evidence of hybrids; for example, Eriosorus war-

scewiczii x Jamesonia scammanae in Costa Rica

and Eriosorus cheilanthoides x E. brasiliensis on

Mt. Itatiaia, Brazil (A. F. Tryon, 1970). Material

from Peru (see Comments) with abortive spores

undoubtedly represents hybrids, but these are often

difficult to identify with particular parents. Field

studies are needed to assess variation in popula-

tions, spore abortion, and cytology of Peruvian

species and hybrids.

This treatment of Eriosorus follows that of A.

F. Tryon (1970), and has been prepared with her

collaboration.

Reference

TRYON, A. F. 1970. A monograph of the fern

genus Eriosorus. Contr. Gray Herb., 200: 54-

174.

Key to Species of Eriosorus

a. Leaves erect, or if scandent or scrambling then to 3-pinnate; lamina mostly to ca. 50 cm long, not

exceeding 1m b

b. Lamina narrowly linear, usually 2-pinnate; pinnae nearly as long as broad 3. E. cheilanthoides

b. Lamina elongate-triangular, -ovate, or -rhomboid; central or lower pinnae nearly twice as long as

the upper ones c

c. Both surfaces of the pinnae sparsely or moderately pubescent d

d. Pinna stalks, at least of the lower pinnae, descending from the rachis ... 6. E. wurdackii

d. Pinna stalks at right angles to the rachis, or ascending from the rachis e

e. Lamina 2-pinnate to rarely 2-pinnate-pinnatifid; pinnae at right angles to the rachis,

sessile or nearly so 1 . E. rufescens

e. Lamina 2-pinnate-pinnatifid to 3-pinnate; pinnae ascending from the rachis, stalked .

2. E. accrescens

c. Both surfaces of the pinnae tomentose with matted trichomes f

f. Lamina rather elongate-triangular; rachis straight or nearly so; pinnae sessile or subsessile

4. E. stuebelii

f. Lamina broadest above the base; rachis fractiflex; pinnae stalks strongly ascending from the

rachis 5. E. aureonitens

a. Leaves scandent, twining, or scrambling, to 3 m long or more; lamina 4-pinnate or more complex

g

g. Ultimate segments orbicular or cuneate, broad, with 6-20 veins 7. E. orbignyanus

g. Ultimate segments dichotomously lobed to deeply bifid, slender, each lobe with 1 or 2 veins . .

. . 8. E. flexuosus

1 . Eriosorus rufescens (Fee) A. F. Tryon, Rhodora

65: 56. 1963. Figure la.

Gymnogramma rufescens Fee, (Mem. foug. 5) Gen.

fil. 181, l. 19C,f. 3. 1852. TYPE: Peru, "Andi-

bus," Matthews (not located).

Gymnogramma mohriaeformis Mett., Fil. lechl. 1: 9.

1856. TYPE: Peru, Puno, San Gavan (Rio San

Gaban), Lechler 2255 (holotype, B!; photo, GH;

frag., NY!; isotype, GH!).

Gymnogramma mathewsii Hooker, Sp. fil. 5: 128, t.

290. 1 864. TYPE: Peru, Mathews 1814 (holotype,

K!; photo, GH).

Leaves erect, determinate. Lamina elongate-del-

toid, 2-pinnate to rarely 2-pinnate-pinnatifid,

FIELDIANA: BOTANY

FIG. 1. Eriosorus rufescens: a, habit. Eriosorus flexuosus: b, portion of lamina; c, apex of penultimate division,

(a from Macbride 4302, F, b, c from Macbride 4518, F.)

rachis straight, shallowly grooved adaxially. Pin-

nae sessile or nearly so, nearly at right angles to

the rachis, both surfaces pubescent, with rigid, clear

or bicolorous trichomes, ultimate segments broad-

ly lobed to crenulate, the vein ends extending to

the margin.

Among rocks, on mossy banks, in Sphagnum,

often growing in open places in cloud forests, 2300-

3600 m, Cajamarca to Puno.

Venezuela, Colombia, south to Bolivia.

This Andean species has the lamina usually

bearing bicolorous trichomes.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

( ajamarea: Prov. Cutervo, La Pucarilla, Ldpez & Sa-

gdstegui 5456 (GH). La Libertad: Prov. Pataz, Pumatam-

bo, Puerto del Monte, Ldpez & Sagastegui 3436 (GH).

Huanuco: Cushi, Bryan 683 (F, us). Pampayacu, Kane-

hira 166 (OH, us). Pasco: Border Prov. Oxapampa and

Pasco, San Gotardo, van der Werffet al. 8600 (MO, uc).

Ayacucho: Prov. La Mar, eastern massif of Cordillera

Central, Dudley 12032 (GH). Cuzco: Prov. Urubamba,

Vargas 2897 (MO, us). Prov. Paucartambo, Vargas 12192

(GH).

shrubs, often in the cloud forest zone, 2300-4000

m, Piura to Puno.

Ecuador to Bolivia; Mt. Itatiaia, Brazil; Tristan

da Cunha Islands.

This species is characterized by the pinnae being

nearly the same length throughout the lamina. It

is somewhat frequent at higher elevations and ev-

idently is one parent of the hybrids discussed in

Comments.

2. Eriosorus accrescens A. F. Tryon, Rhodora 65:

57. 1963. TYPE: Peru, (Cuzco), Prov. Uru-

bamba, Puyupata to "Yuncapata," Vargas

2921 (holotype, us!; isotype, MO!).

Leaves erect to subscandent, with indeterminate

growth. Lamina elongate-lanceolate or elongate-

ovate, 2-pinnate-pinnatifid or 3-pinnate, rachis

nearly straight to slightly flexuous, plane to slightly

grooved adaxially. Pinnae stalked, ascending, both

surfaces sericeous, ultimate segments bluntly lobed

to crenulate, the vein ends terminating in a sinus

at the margin.

Wooded ravines, 2950-3500 m, Amazonas, San

Martin, Pasco, and Cuzco.

Peru.

Additional material is needed to adequately

characterize this rare species.

Amazonas: Prov. Chachapoyas, entre Leimebamba y

Balsas, Lopez el al 4444 (GH). San Martin: Prov. Mar-

iscal Caceres, Rio Abiseo National Park, Young & Leon

4726 (F). Pasco: Prov. Oxapampa, Santa Barbara, D.

Smith 8129 (F). Cuzco: Valle de Lares, Montana de Col-

ca, Biies 1925 (us). Altura de Chaco, Biies 2135 (us).

3. Eriosorus cheilanthoides (Sw.) A. F. Tryon, Brit.

FernGaz. 9: 271. 1966.

Grammitis cheilanthoides Sw., Syn. fil. 23, 219, 419.

1 806. TYPE: Tristan da Cunha (Mauritius in error)

(holotype, Herb. Swartz, s!; photo, GH).

Leaves erect, indeterminate. Lamina narrowly

linear, pinnate-pinnatifid, rarely 2-pinnate, rachis

straight, plane to slightly grooved adaxially. Pin-

nae stalked, both surfaces more or less pubescent,

or rarely glandular, the ultimate lobes bifid, the

vein ends extending to, or nearly to, the segment

margin.

Rocky soil, at the edge of boulders, among

Piura: Prov. Huancabamba, Cerro La Viuda, Sagas-

tegui el al. 8211 (GH, HUT, MO). C ajamarca: Prov. Ca-

jabamba, Cajabamba-Luchubamba, Sagastegui et al.

11204 (HUT, MO). Prov. Cutervo, La Pucarilla, Lopez &

Sagastegui 5380 (GH, MO). Ancash: Prov. Huaraz, Huas-

caran National Park, Smith et al. 9623, 12406, 12459

(F). Huanuco: Prov. Huanuco, Carpish, Tryon & Tryon

5319 (GH). Pasco: Prov. Oxapampa, Santa Barbara, D.

Smith 8143 (F). Junin: Prov. Conception, Concepcion-

Satipo, Sounders 1078 (GH). Huancavelica: Prov. Tay-

acaja, entre Colcabamba y Paucartambo, Tovar 2026

(USM). Ayacucho: Prov. La Mar, between Tambo and the

Apurimac, Weberbauer 5658 (B, F, GH, us). Pampalca,

Killip & Smith 23249 (F, GH, NY, s, us). Cuzco: Prov.

Paucartambo, Puente Aguila, Vargas 23122 (GH). Prov.

Cuzco, Rio Marcapato, Madison 1035 (GH). Puno: Aga-

pata, Lechler 2035 (B, F, G, K, p, w).

4. Eriosorus stuebelii (Hieron.) A. F. Tryon, Rho-

dora 65: 57. 1963.

Gymnogramma stuebelii Hieron., Hedwigia 48: 219,

t. 9,f. 5. 1909. TYPE: Peru, (Amazonas), between

Pacasmayo and Moyobamba, Stubel 1058 (ho-

lotype, B!; frag., GH!; photo, GH).

Leaves erect, determinate. Lamina rather elon-

gate-deltoid, 2-pinnate, rachis straight, subterete

to slightly grooved adaxially. Pinnae nearly sub-

sessile, both surfaces rather densely sericeous, ul-

timate segments crenulate, the vein ends termi-

nating in a sinus at the margin or protruding in a

tooth beyond the margin.

In woods, 2000-3000 m, Amazonas and Puno.

Peru.

The Vargas collection cited below is not entirely

consistent with the type and may represent another

species.

Amazonas: Chachapoyas, Wurdack 1527 (US). Puno:

Prov. Sandia, bajando a Cachi-cachi, Vargas 1 1834 (GH).

5. Eriosorus aureonitens (Hooker) Copel., Gen.

fil. 58. 1947.

FIELDIANA: BOTANY

Gymnogramma aureonitens Hooker, Hooker's Icon.

PI. 9: /. 520. 1 852 (prior to May, cf. Gard. Chron.

1852: 278). TYPE: Peru, W. Lobb (holotype, K!;

frag., NY!; photo, OH).

Eriosorus scandens Fee, (Mem. foug. 5) Gen. fil. 152,

t. 13B,f. 1. 1852 ("probably December" cf. W.

T. Stearn, Webbia 17: 207-222. 1962). TYPE:

Peru, Ruiz (not located).

Leaves subscandent or pendent, with indeter-

minate growth. Lamina elongate-lanceolate,

broadest above the base, 2-pinnate-pinnatifid,

rachis fractiflex, subterete or shallowly grooved

adaxially. Pinnae stalked, ascending from the

rachis, adaxially, both surfaces densely tomentose,

ultimate segments bluntly lobed to crenulate, the

vein ends usually protruding in a tooth beyond the

margin.

Shrubby areas, especially in cloud forests and

elfin forests, 2700-3400 m, Piura to Cuzco.

Colombia, Ecuador, and Peru.

The fractiflex rachis and dense tomentum on

the lamina are unusual features that readily char-

acterize this species.

Piura: Prov. Huancabamba, Cordillera Chinguala, Sa-

gdstegui el al. 10222 (GH, uc). Amazonas: Prov. Chach-

apoyas, middle eastern Calla Calla slopes, Wurdack 1 738

(GH, us). Prov. Bagua. Cordillera Colan, Harbour 3370

(F, MO). Huanuco: Playapampa, Macbride 4529 (F). Be-

tween Huanuco and Pampayacu, Kanehira 153 (GH, us).

Pasco: Prov. Oxapampa, Cordillera Yanachaga, Foster

9016 (F, MO). Border Prov. Oxapampa and Pasco, San

Gotardo, van der \Verff8506 (MO, uc). Cuzco: Prov. La

Convention, Cordillera Vilcabamba, Dudley 11090 (GH).

6. Eriosorus wurdackii A. F. Tryon, Contr. Gray

Herb. 200: 147. 1970. TYPE: Peru, Dept.

Amazonas, Prov. Chachapoyas, 6-8 km W of

Molinopampa, Wurdack 1541 (holotype, GH!).

Leaves erect, determinate. Lamina elongate-tri-

angular, 2- or 3-pinnate basally, rachis fractiflex,

at least toward the base, strongly grooved adaxi-

ally. Pinnae stalked, at least the lower ones de-

scending from the rachis, glabrous adaxially,

slightly pubescent abaxially, ultimate segments or-

bicular to ovate, veins extending to, or slightly

short of, the margin.

Rock crevices, in the jalca zone, 2200-2300 m,

Amazonas. Only known from the type collection.

Peru.

This species and E. orbignyanus are unusual in

that the sori form a band on the fertile segment.

7. Eriosorus orbignyanus (Kuhn) A. F. Tryon,

Rhodora65: 56. 1963.

Gymnogramma orbignyana Kuhn, Linnaea 36: 70.

1869. SYNTYPES: Bolivia, (La Paz), Yungas,

D'Orbigny 174 (B!; photo, GH; isosyntype, F!; pho-

to, GH); Bolivia, (Cochabamba) "Yorocares" (Yu-

racares), D'Orbigny 299 (B!; photo, GH; isosyn-

type, P!; photo, GH).

Leaves scandent, evidently indeterminate. Lam-

ina elongate, 4-pinnate, rachis fractiflex, strongly

grooved adaxially. Pinnae stalked, descending from

the rachis, both surfaces sparsely pubescent, ulti-

mate segments broadly lobed to crenulate, with

mostly 6-20 veins, the vein ends extending to, or

nearly to, the margin.

Cloud forests, 1500-1750 m, Huanuco to Cuz-

co.

Colombia south to Bolivia.

Huanuco: Rio Lullapichis watershed, Cerros del Sira,

Dudley 13474 (GH). Pasco: Border Prov. Oxapampa and

Pasco, San Gotardo, van der Werffet al. 8585 (MO, uc).

Prov. Oxapampa, Oxapampa-Paucartambo, Smith &

Pretel 1632 (F). Junin: Provenir, Killip & Smith 25947

(NY, us). Cuzco: Chaupimayo, Sues 1949, 1950 (us).

Prov. La Convencion, Dudley 10337, 11240 (GH).

8. Eriosorus flexuosus (H.B.K.) Copel., Gen. fil.

58. 1947. Figure Ib-c.

Gymnogramma flexuosa Desv., Ges. Naturf. Freunde

Berlin Mag. Neuesten Entdeck. Gesammten Na-

turk. 5: 306. 181 1, ex char., not Eriosorus flex-

uosus (H.B.K.) Copel. TYPE: none cited.

Grammitis flexuosa H.B.K., Nov. gen. sp. 1:5. 1816.

TYPE: Venezuela, Humboldt & Bonpland (ho-

lotype, P!; photo, GH).

Grammitis ruiziana Klotzsch, Linnaea 20: 410. 1847.

TYPE: Peru, (Huanuco), Prov. Panatahuas, Ruiz

74 (holotype, B!; photo, GH).

Gymnogramma flexuosa var. peruviana Hieron.,

Hedwigia 48: 220. 1909. TYPE: Peru, (Amazon-

as), Cuesta de Lejia, near Molinobamba, Stubel

1055 (holotype, B!; photo, GH).

Leaves scandent, scrambling, indeterminate,

sometimes more than 3 m long. Lamina elongate,

branching in several planes, 4- or 5-pinnate, rachis

more or less fractiflex, strongly grooved adaxially.

Pinnae stalked, ascending, or nearly at right angles

to the rachis, both surfaces glabrous or sparsely

pubescent, ultimate segments slender, bifid to

deeply lobed, each lobe with 1 or 2 veins, vein

ends extending to, or short of, the segment margin.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

Among shrubs in moist areas, along river banks

or on mossy slopes, scrambling on other vegeta-

tion, 2200-3600 m, Cajamarca to Puno.

Mexico and Central America; Hispaniola; Guy-

ana west to Colombia and south to Bolivia.

This species is characterized by very large and

long scrambling leaves with the pinnae disposed

in several planes, and slender ultimate segments

with each lobe having one or two veins.

Cajamarca: Prov. Cutervo, La Pucarilla, Lopez & Sa-

gdstegui 5458 (GH). NE of Socota, Stork & Morton 10132

(F, uc, us). Amazonas: 9 km below Chachapoyas, Wur-

dack 777 (GH, us). Prov. Bagua, Cordillera Colan, Bar-

hour 3754 (F, MO). San Martin: Prov. Rioja, Pedro Ruiz-

Moyobamba, D. Smith 4827 (GH). Huanuco: Prov.

Huanuco, Bosque Nacional de Iparia, J. Schunke V. 5227

(F, GH, us). Mito, Macbride & Featherstone 1829 (F, GH,

us). Pasco: Prov. Oxapampa, Cordillera Yanachaga,

Foster 9010 (F, MO). Huancavelica: Prov. Tayacaja, E of

Surcubamba, Stork & Norton 10393 (F, GH, us). Cuzco:

Pillahuata. Cerro de Cusilluyoc, Pennell 13941 (F, GH,

NY). Paucartambo, Vargas 1906 (GH, us). Madre de Dios:

Prov. Manu, Shintuya, Chavez 859 (MO). Puno: Prov.

Carabaya, entre Ayapata y Kahualluyoc, Vargas 10750

(GH).

Comments

DUBIOUS SPECIES

Eriosorus lechleri (Kuhn) A. F. Tryon, Rhodora

65: 56. 1963; Contr. Gray Herb. 200: 171.

1970.

Gymnogramma lechleri Kuhn, Linnaea 36: 71. 1869.

TYPE: Peru, Puno, San Gavan (Rio San Gaban),

Lechler 2262 (holotype, B!; photo, GH).

The type is a single leaf that cannot be definitely

identified.

HYBRIDS

A considerable number of collections have been

identified as hybrids on the basis of morphological

irregularities, abortive spores or sporangia, and

characters intermediate between putative parents.

The two most frequent ones are the following.

Eriosorus cheilanthoides • Eriosorus sp.

Gymnogramma flabellata Grev. & Hooker, J. Bot.

(Hooker) 1: 61, /. 120. 1834. TYPE: Ecuador,

(Azuay), near Cuenca, Jameson in 1832 (holo-

type, Herb. Greville, E!; photo, GH; isotype, Herb.

Hooker, K!; photo, GH).

Eriosorus flabellatus (Grev. & Hooker) Copel., Gen.

fil. 58. 1947.

Specimens of this hybrid mostly have abortive

spores. They are similar to Eriosorus cheilan-

thoides in the narrowly elongate lamina; however,

the pinnae are about twice as long as broad. The

other parent may be Eriosorus flexuosus.

Cajamarca: San Juan, Vargas el al. 5512 (USM). La

Libertad: Prov. Sanchez, between Huamachuco and Ca-

jabamba, Smith & Cdceres 2233 (F, MO). Ancash: Prov.

Yungay, Huascaran National Park, Smith et al. 10378,

10379 (GH). Cuzco: Prov. Paucartambo, Puente Aguila,

Vargas 23123 (GH).

Eriosorus cheilanthoides • Jamesonia sp.

Gymnogramma elongata Grev. & Hooker, J. Bot.

(Hooker) 1: 61, t. 119. 1834. TYPE: Ecuador,

(Azuay), near Cuenca, Jameson in 1832 (holo-

type, Herb. Greville, E!; photos, BM, GH; isotype,

Herb. Hooker, K!; photo, GH).

Eriosorus elongatus (Grev. & Hooker) Copel., Gen.

fil. 58. 1947.

Specimens are similar to both parents in the

narrowly linear lamina. Some collections may be

part of the variation of Eriosorus cheilanthoides,

while others with abortive spores are probably hy-

brids with one of the species of Jamesonia. The

collections will key out to E. cheilanthoides, but

the ultimate lobes have three or usually more veins,

rather than one or two veins.

Piura: Prov. Huancabamba, above Huancabamba,

Hutchison 1616 (GH). Prov. Huancabamba, E of Huan-

cabamba, Weberbauer 6096 (B, F, GH, us). Cajamarca:

Prov. San Miguel, El Tingo, Sagdstegui et al. 9516 (F).

Amazonas: Summit of Puma-urcu, Wurdack 1161 (GH,

us). Cerros Calla Calla, Hutchison & Wright 6978 (F, GH,

us). La Libertad: Prov. Bolivar, La Quinuas, Lopez &

Sagdstegui 3348 (GH). Huanuco: Carpish, Stork & Nor-

ton 9910 (F, GH, MO). Carpish, Ferreyra 8172 (GH). Cuzco:

Cerro de Cusilluyoc, Pennell 13850 (F, GH, NY, s, us).

II. Jamesonia

Jamesonia Hooker & Grev., Icon. fil. 2, /. 178.

1830. TYPE: Jamesonia pulchra Hooker &

Grev. Figure 2.

PsilogrammeKuhn, Festschrift 50jahrjub. Real. Ber-

lin 332 (reprint Chaetopterides 12). 1882, nom.

superfl. for Jamesonia and with the same type.

FIELDIANA: BOTANY

Terrestrial. Stem decumbent to usually creep-

ing, bearing trichomes or bristles. Leaves usually

well spaced. Lamina 1 -pinnate, pubescent or glan-

dular abaxially, veins free. Sori rather elongate

along the veins, extending almost to a vein tip,

not paraphysate, exindusiate. Spores tetrahedral-

globose, trilete, coarsely ridged or tuberculate, with

a prominent equatorial flange.

Differences ofJamesonia and the closely related

Eriosorns are discussed under the latter genus.

Specimens that are old or incomplete are often

difficult to identify. The indument on the pinnae

is best observed in younger leaves toward the apex

of the lamina. Several leaves and pinnae should

be examined in order to be certain of a character.

The mature sporangia of fully fertile pinnae may

conceal the indument on the abaxial surface.

This treatment ofJamesonia follows that of A.

F. Tryon (1962) and has been prepared with her

close collaboration.

Reference

TRYON, A. F. 1962. A monograph of the fern

genus Jamesonia. Contr. Gray Herb. 191: 109-

197.

Key to Species of Jamesonia

a. Pinnae asymmetrical, inequilateral at the usually cordate base b

b. Abaxial surface of the pinnae densely tomentose, the trichomes often matted c

c. Lamina not reduced apically, with patent or slightly appressed trichomes d

d. Stem trichomes dark brown to atropurpureous, darker than the stem surface; border of the

pinnae usually broad, indusioid 2. J. pulchra

d. Stem trichomes tan to light brown, lighter than the stem surface; border of the pinnae narrow

to moderately broad 9. J. imbricata

c. Apex of the lamina vermiform, glandular-pubescent, usually vernicose ... 3. J. scammanae

b. Abaxial surface of the pinnae glabrous, glandular, or somewhat sparsely and loosely pubescent

e

e. Pinnae rigidly herbaceous, the abaxial surface pubescent f

f. Pinnae pubescent adaxially, especially on the enrolled margin g

g. Pinnae with rather short trichomes adaxially, these hardly extending to other pinnae .

1 . J. rotundifolia

g. Pinnae with long trichomes adaxially, these usually enveloping several pinnae

4. J. peruviana

f. Pinnae glandular-pubescent abaxially 7. J. boliviensis

e. Pinnae coriaceous, the adaxial surface glabrous or rarely with appressed glands or a few tri-

chomes h

h. Lamina the same width toward the apex; tomentum of apex and rachis concolorous ....

5. J. alstonii

h. Lamina broadened and narrowly clavate toward the apex; tomentum of the apex and rachis

bicolorous, usually with a prominent dark streak 6. J. goudotii

a. Pinnae symmetrical, equilateral at the truncate to cordate base i

i. Pinnae with slender stalks usually 1 mm or more long, these often perpendicular to the rachis;

the pinna border entire to slightly short-ciliate; veins extending to the margin j

j. Pinnae cordate at the base, the abaxial surface glandular or with a few short trichomes

8. J. scalaris

j. Pinnae truncate to subcordate at the base, the abaxial surface usually tomentose

9. J. imbricata

i. Pinnae adnate, sessile or subsessile, or with a stalk usually ca. 1 mm long and at an oblique angle

to the rachis; the pinna border short- to long-ciliate k

k. Stem trichomes dark brown to atropurpureous; veins extending nearly to the margin

10. J. blepharum

k. Stem trichomes tan to golden brown; veins ending well back of the margin

. 11. J. cinnamomea

TRYON & STOLZE: PTER1DOPHYTA OF PERU. II.

5 cm

5 cm

FIG. 2. Jamesonia alstonii: a, habit; b, portion of rachis with pinnae, adaxial side, (a from Killip & Smith 17475,

Colombia, GH; b from Cleef9736, Colombia, GH.)

1. Jamesonia rotundifolia Fee, Mem. foug. 7: 41,

t. 10. 1857. TYPE: Colombia, Schlim 363

(holotype, not located; isotypes, G!, K!, P!; pho-

to, GH of P).

Stem trichomes castaneous to light brown. Lam-

ina about the same width throughout, the bud with

patent or slightly appressed trichomes. Pinnae ro-

tundate-cordate, inequilateral at the base, rigidly

10

FIELDIANA: BOTANY

herbaceous, adaxial surface more or less pubes-

cent, abaxial surface pubescent or glabrous, the

stalk at an oblique angle to the rachis, border mod-

erately broad, extended at the vein ends, irregu-

larly dentate, sometimes little modified and sim-

ilar to the pinna in texture.

Moist, grassy steppes, rocky places, especially

in the jalca of northern Peru, 3200-3500 m, Piura,

Amazonas, and Huanuco.

Costa Rica; Colombia to Peru.

This species is characterized by its rotundate-

cordate, inequilateral pinnae that are pubescent

on the adaxial surface.

Piura: Prov. Huancabamba, San Antonio-Talaneo,

Sagdstegui & Cabanillas 8622 (F, MO, uc). Prov. Huan-

cabamba, E of Huancabamba, Weberbauer 6076 (B, F,

GH, us). Amazonas: Prov. Chachapoyas, Cordillera de

Calla Calla, Ldpez et al 4447 (HUT). Huanuco: Cushi,

Bryan 612 (F).

2. Jamesonia pulchra Hooker & Grev., Icon. ill.

2, /. 178. 1830. TYPE: Ecuador, (Pichincha),

Monte Cayambe (cited as Peru), Jameson (ho-

lotype, E!; photos, BM, GH; isotype, Herb.

Hooker, K!).

Stem trichomes dark brown to atropurpureous,

darker than the stem surface. Lamina usually

slightly broader in the central portion, the bud

with patent or slightly appressed trichomes. Pin-

nae ovate to orbicular, cordate, inequilateral at the

base, rigidly herbaceous, adaxial surface glabrous

or slightly pubescent, abaxial surface densely to-

mentose, the stalk at an oblique angle to the rachis,

border usually broad, often indusioid, entire or

somewhat dentate.

In a Sphagnum bog, 3350 m, Pasco.

Colombia, Ecuador, and Peru.

Jamesonia pulchra has only recently been col-

lected in Peru. It probably also grows in the jalca

of northern Peru where several other species of the

genus occur. The broad, often indusioid, border

of the pinnae is especially characteristic of this

species.

Pasco: Prov. Oxapampa, Cordillera Yanachaga, D.

Smith 7729 (F, GH).

3. Jamesonia scammanae A. F. Tryon, Contr. Gray

Herb. 191: 164. 1962. TYPE: Costa Rica,

Cerro de la Muerte, Scamman & Holdridge

7929 (holotype, GH!).

Stem trichomes lustrous, light to dark brown.

Lamina about the same width throughout, the bud

vermiform, glutinous, often vernicose, with ap-

pressed trichomes. Pinnae ovate, sometimes lobed,

inequilateral or nearly so at the base, adaxial sur-

face glandular and vernicose or crustose, abaxial

surface tomentose, stalk bent, short to nearly ab-

sent, border moderately broad, entire or slightly

undulate, firm-membranous or similar to the pin-

na in texture.

Grassy and rocky places, usually moist sites,

2750-4400 m, Ancash to Puno.

Costa Rica; Colombia to Bolivia.

This species is readily recognized by the small,

vermiform and usually vernicose lamina apex.

Ancash: Prov. Huari, Huascaran National Park, Smith

& Goodwin 8749 (F, GH). Huanuco: Huanuco, Macbride

& Featherstone 2182 (F, GH, us). Lima: Huarochiri,

Sounders 399 (BM). Pasco: Prov. Oxapampa, Santa Bar-

bara, Foster et al. 10427 (F). Junin: Prov. Satipo, Pampa

Hermosa, Sounders 1036 (GH). Apurimac: Bosques de

Ampay, Vargas 1060 (GH). Cuzco: Cerro de Cusilluyoc,

Pennell 13870 (F, GH, us). Achirani, Vargas 11165 (F,

K, uc). Puno: Tabina, Lechler 2032 (B, E, K, LE, p).

4. Jamesonia peruviana A. F. Tryon, Contr. Gray

Herb. 191: 167. 1962. TYPE: Peru, (Huan-

uco), Tambo de Vaca, J. F. Macbride 4404

(holotype, F!).

Stem trichomes castaneous to atropurpureous.

Lamina broadest at the apex, the bud with tan or

light brown, patent or somewhat appressed tri-

chomes. Pinnae reniform or orbicular-cordate,

inequilateral at the base, rigidly herbaceous or

slightly coriaceous, adaxial surface with long tri-

chomes, these usually enveloping several pinnae,

abaxial surface sparsely pubescent, stalk straight,

perpendicular to the rachis, border broad, with

more or less uniform papillae, firm-membranous.

Grassy places, especially in moist and rocky sites,

2800-4300 m, Amazonas to Puno.

Peru and Bolivia.

This species is characterized by the very long

trichomes, borne on the adaxial surface of the pin-

nae, which envelope the adjacent pinnae.

Amazonas: Prov. Chachapoyas, summit of Cerros de

Calla Calla, Wurdack 1216 (GH, us). La Libertad: Las

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

11

Quinuas. Ldpez & Sagdstegui 3346 (F, GH, MO, uc). An-

cash: Huari, above Ponto. Weberbauer 3302 (B, G, us).

Huanuco: Tambo de Vaca, Bryan 648 (F, us). Pasco:

Prov . Oxapampa, Santa Barbara. D. Smith 81 14 (F, GH),

Foster el al. 10433 (F). Junin: Tarma, Nee (F). Cuzco:

Cerro Cusilluyoc. Pennell 13873 (F, GH, us). Puno: Lech-

ler 2153, in part (B. E. c, K. LE, P).

5. Jamesonia alstonii A. F. Tryon, Contr. Gray

Herb. 191: 168. 1962. TYPE: Colombia, del

Valle, Los Farallones, Cuatrecasas21884(ho-

lotype, GH!). Figure 2.

Stem trichomes usually dark brown or blackish,

sometimes light brown. Lamina about the same

width throughout, the bud with straw-colored,

matted, or slightly appressed trichomes, the to-

mentum of the apex and rachis concolorous. Pin-

nae ovate-cordate or orbicular-cordate, inequila-

teral at the base, coriaceous, the adaxial surface

glabrous, rarely glandular, abaxial surface sparsely

pubescent, stalk straight, perpendicular to the

rachis, border moderately broad, extended at the

vein ends, dentate, with sparse, short, bulbous cilia

or entire, rigidly herbaceous.

Wet grasslands, among boulders, 3400-4000 m,

Cajamarca to Puno.

Southern Mexico; Guatemala and Costa Rica;

Colombia to Bolivia.

Jamesonia alstonii is distinguished by the co-

riaceous pinnae and the concolorous tomentum of

the lamina apex and rachis. It is most closely re-

lated to J. goudotii.

Cajamarca: Prov. Contumaza, jalca de Pozo Kuan,

Sagdstegui et al. 10790 (F, HUT). La Libertad: Prov. Pa-

laz, Paseo de Alaska, Lope: & Sagdstegui 8176 (GH, MO).

Huillias, N of Cachicadon, Stork & Morton 10007 (F, G,

K, uc). Ancash: Prov. Bolognesi, Condorcocha, Cerrate

2547 (GH). Huanuco: Mito, Macbride & Featherstone

1883 (F, G, us). Junin: Huayllay, Mathews 979 (E, GL,

K). Cozco: Cordillera Vilcabamba, Dudley 11191 (GH).

Bites 1378 (us). Puno: Lechler 2153, in part (B, E, o, K,

LE, P, s).

6. Jamesonia goudotii (Hieron.) C. Chr., Index til.

373. 1905.

Gymnogramma goudotii Hieron., Bot. Jahrb. Syst. 34:

476. 1904. TYPE: Colombia, Goudot (holotype,

B!; photo, GH; isotype, G!; photo, GH).

Stem trichomes light brown to lustrous black.

Lamina narrowly clavate. broadest at the apex, the

bud with erect or slightly appressed trichomes, the

tomentum of the apex and rachis bicolorous, usu-

ally with a patch or streak of dark trichomes. Pin-

nae reniform or orbicular-cordate, inequilateral at

the base, coriaceous, the adaxial surface usually

glabrous, sometimes glandular, abaxial surface

sparsely pubescent, stalk straight, perpendicular to

the rachis, border usually extended at the vein

ends, irregularly dentate or ciliate, rarely entire or

nearly so, rigidly herbaceous.

In grasslands and elfin forests, sometimes near

glaciers, usually in wet sites, 3200—4800 m, Ama-

zonas to Cuzco.

Colombia to Peru.

Jamesonia goudotii is characterized by the nar-

rowly clavate lamina that is broadest at the apex

and the bicolorous tomentum in that region.

A single collection from Amazonas: Top of di-

vide between Rio Maranon and Rio Utcubamba,

28-31 kmSWofLeimebambajalca, 17 June 1978,

Gentry et al. 23168 (F, MO) is evidently a hybrid

of /. goudotii and J. imbricata.

Amazonas: Prov. Bagua, Cordillera de Colan, Barbour

3166 (MO). Ancash: Huascaran National Park, D. Smith

11301 (GH), Smith et al. 11655 (F), Smith et al. 12345

(F, GH). Junin: near Comas, Weberbauer 6629 (F, GH,

us). Cuzco: Bues 1406 (us).

7. Jamesonia boliviensis A. F. Tryon, Contr. Gray

Herb. 191: 174. 1962. TYPE: Peru, Apuri-

mac, Bosques de Ampay, Vargas 1060a (ho-

lotype, GH!).

Stem trichomes lustrous, atropurpureous or

blackish. Lamina about the same width through-

out, the bud with rust-colored, patent trichomes.

Pinnae orbicular-cordate or ovate-cordate, ine-

quilateral at the base, rigidly herbaceous, both sur-

faces pubescent, stalk straight, perpendicular to

the rachis, border narrow, with capitate cilia.

In woods and elfin woodlands, 3200-3400 m,

Pasco and Apurimac.

Peru and Bolivia.

This species is unique in having the same form

of indument on both surfaces of the pinnae and

on the border.

Pasco: Prov. Oxapampa, Santa Barbara, Foster et al.

10387 (F). Apurimac: Abancay region, Oct. 1935, V.

Santander et al. (uc).

12

FIELDIANA: BOTANY

8. Jamesonia scalaris Kunze., Hot. Zeit. (Berlin)

2: 738. 1844. TYPE: Peru, (Huanuco), Pillao,

Ruiz 49 (holotype, B!; photo, GH).

Stem trichomes light to dark brown or atropur-

pureous. Lamina about the same width through-

out, the bud with tan or rust-colored, patent or

slightly appressed trichomes. Pinnae ovate-cor-

date, usually lobed, equilateral or nearly so at the

base, herbaceous, the adaxial surface with papil-

late glands, abaxial surface pubescent, sometimes

tomentose, stalk straight or slightly bent, perpen-

dicular to the rachis, veins extending to the mar-

gin, border narrow, with capitate cilia.

Shrublands and grasslands, dwarf forests, cloud

forests, especially in open, wet sites, 2700-4300

m, La Libertad to Puno.

Peru and Bolivia.

This species is distinguished from others with

equilateral and stalked pinnae by the cordate pin-

nae with the stalk perpendicular to the rachis.

La Libertad: Prov. Pataz, Pumatambo, Lopez & Sa-

gdstegui 3435 (GH). Ancash: Prov. Yungay, Huascaran

National Park, Smith el al. 9090 (F, GH). Huanuco: Punta

de Panao, Asplund 13718 (GH, s). Playapampa, Macbride

4524 (F, us). Pasco: Prov. Oxapampa, Cordillera San

Gutardo, Leon et al. 533 (USM). Junin: Prov. Huancayo,

arriba de Chilifruta, Cerrate 5649 (USM). Huancavelica:

Prov. Tayacaja, entre Marcavalle y Huachocolpa, Tovar

4230 (GH, USM). Ayacucho: Prov. La Mar. near Cusi-

machay. Madison 10389(GH). Cuzco: Prov. La Conven-

cion, Cordillera Vilcabamba, Dudley 11190 (GH). Prov.

Paucartambo, Vargas 4322 (GH). Puno: Ayapata, Lechler

2036a (B, E, o, K, LE, P).

9. Jamesonia imbricata (Sw.) Hooker & Grev.,

Icon. fil. 1: 2. 1831.

Stem trichomes tan to dark brown. Lamina about

the same width throughout, the bud with tan,

brown, or bicolorous, patent or appressed tri-

chomes. Pinnae orbicular to ovate, equilateral and

truncate or slightly inequilateral and somewhat

cordate at the base, rigidly herbaceous, adaxial

surface glandular or sometimes also with a few

trichomes, rarely glabrous, abaxial surface tomen-

tose, stalk usually bent or twisted and oblique to

the rachis, veins extending to the margin, border

moderately broad, entire or dentate and sparsely

ciliate.

This is a variable species with three varieties

recognized by A. F. Tryon (1962). Among these,

var. imbricata is known in Peru only from two

collections, including the type, and var. glutinosa

is known from several collections, especially from

the Department of Amazonas. The other, var.

meridensis A. F. Tryon, is confined to Venezuela

and Colombia.

Venezuela, Colombia, Ecuador, and Peru.

Key to Varieties

a. Stem trichomes tan to light brown, lighter than the stem surface; pinnae slightly inequilateral at the

base and somewhat cordate 9a. var. imbricata

a. Stem trichomes reddish to dark brown, the same color or darker than the stem surface; pinnae

equilateral at the base, truncate 9b. var. glutinosa

9a. Jamesonia imbricata var. imbricata

Pteris orbiculata Poiret in Lam., Encycl. 5: 710. 1804,

not Houtt., Nat. Hist. 14: 108. 1783. TYPE: "Pi-

rou," J. Jussieu (holotype. Herb. Jussieu no. 1 332,

P!; photo, GH).

Pteris imbricata Sw., Syn. fil. 102. 1806, based on

Pteris orbiculata Poiret.

In the jalca, 3 1 50 m, Lambayeque.

Colombia, Ecuador, and Peru.

Lambayeque: Prov. Ferrenafe, Laguna Tembladera,

Sagdstegui et al. 12796 (uc).

9b. Jamesonia imbricata var. glutinosa (Karsten)

A. F. Tryon, Contr. Gray Herb. 191: 182.

1962.

Jamesonia glutinosa Karsten, Fl. Columb. 2: 85, /.

143. 1865. TYPE: Colombia. (Cundinamarca),

Bogota, Monte Guadeloupe, Karsten (holotype,

LE!; photo, GH).

In grasslands, scrub forests, and rocky places,

3100-3700 m, Amazonas to Cuzco.

Venezuela, Colombia. Ecuador, and Peru.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

13

Ama/onas: Prov. Chachapoyas, Cordillera Calla-Cal-

la, Leimebamba to Balsas, Ldpez el al. 4447, 4448 (GH).

Summit of Puma-urcu, SE of Chachapoyas, Wurdack

1160 (GH, us). 28-31 km SW of Leimebamba on road

to Balsas, Gentry el al. 23167 (F, MO, us). La Libertad:

Prov. Bolivar, Nevada de Cajamarquilla, Ferreyra 1348

(USM). Pasco: Prov. Oxapampa, Santa Barbara, D. Smith

8125 (F, GH). Cuzco: Tinke, Humbert 30735 (GH).

10. Jamesonia blepharum A. F. Tryon, Contr.

Gray Herb. 191: 190. 1962. TYPE: Peru,

Huanuco, Pampayacu, Ryozo Kanehira 148

(holotype, GH!; isotype, us!).

Stem trichomes lustrous, light to dark brown to

atropurpureous. Lamina nearly the same width

throughout or the base and apex more slender, the

bud with dark brown, appressed, glutinous tri-

chomes. Pinnae orbicular, equilateral at the base,

rigidly herbaceous, the adaxial surface vernicose

with appressed glands, abaxial surface pubescent,

stalk oblique (if present), often broad and decur-

rent on the adaxial surface of the rachis, veins

extending nearly to the margin, border narrow,

ciliate.

In woods, in dwarf forests, 2700-2800 m, Huan-

uco, Pasco, and Cuzco.

Colombia to Bolivia.

The characters in the key provide an adequate

separation of this species and Jamesonia cinna-

momea. It is characterized by nearly adnate pin-

nae and the usually long-ciliate pinna-border. The

lamina is sometimes determinate.

Huanuco: Carpish, entre Huanuco y Jingo Maria, Fer-

reyra 6702 (GH). Prov. Huanuco, Carpish, Hutchison el

al. 5961 (GH). Pasco: Border Prov. Oxapampa and Pasco,

San Gotardo, van der Werffel al. 8553, 8565 (uc). Cuzco:

Valle de Lares, Bites 1908, 1824, 1914 (us).

1 1 . Jamesonia cinnamomea Kunze, Bot. Zeit.

(Berlin) 2: 738. 1 844. TYPE: Colombia, Hart-

weg 1516 (holotype, Herb. Shuttleworth, BM!;

photos, F, GH, NY, us; isotypes, B!, BM!, G!,

GH!, K!, LE!, us!).

Stem trichomes lustrous tan to golden brown.

Lamina nearly the same width throughout, the bud

with a tomentum of rather appressed, cinnamo-

meous trichomes. Pinnae orbicular, equilateral at

the base, coriaceous, the adaxial surface with ap-

pressed, glutinous trichomes, often vernicose,

abaxial surface somewhat pubescent, stalk oblique

to the rachis, veins ending well back of the margin,

border usually narrow, ciliate or glandular.

In the jalca area of northern Peru, 3 1 50 m, Lam-

bayeque.

Colombia, Ecuador, and Peru.

The orbicular pinnae and usually large, reddish

brown lamina bud are characteristic of this species.

Rarely the lamina may be determinate.

Lambayeque: Prov. Ferrenafe, Laguna Tembladera,

Sagdstegui et al. 12796 (HUT).

III. Pterozonium

Pterozonium Fee, Mem. Soc. Mus. Nat. Hist.

Strasbourg 4: 202. 1850. Also (Mem. foug. 5)

Gen. fil. 37, 178. 1852. TYPE: Pterozonium

reniforme (Mart.) Fee (Gymnogramma reni-

formis Mart.). Figure 3.

Terrestrial or usually rupestral. Stem small to

stout, prostrate-creeping to erect, bearing tri-

chomes, bristles, or scales. Leaves clustered or

sometimes well spaced. Lamina simple, entire, or

1 -pinnate or partly 2-pinnate, glabrous to pubes-

cent abaxially, veins free. Sori along the veins,

short and near the vein tips to long and along most

of the vein, not paraphysate, but some short tri-

chomes or farina may be among the sporangia,

exindusiate. Spores tetrahedral-globose, trilete,

with a prominent equatorial flange, the surface

nearly smooth to somewhat tuberculate or rugose.

The 1 3 species of Pterozonium center on the

Roraima sandstones of Venezuela, where all of the

species occur. The genus is endemic to northern

South America and Peru where two species grow.

Reference

LELLINGER, D. B. 1967. Pterozonium, in The

Botany of the Guayana Highland— Part 8, Mem.

New York Bot. Gard., 17: 2-23.

Key to Species of Pterozonium

a. Lamina suborbicular to orbicular, with flabellate venation; stem with trichomes . 1. P. reniforme

a. Lamina elongate, with pinnate venation; stem with scales 2. P. brevifrons

14

FIELDIANA: BOTANY

-:-

••

FIG. 3. Pterozonium reniforme: a, habit; b, portion of fertile leaf, abaxial side; c, stem trichome. (From Steyermark

60480, Venezuela, F.)

I . Pterozonium reniforme (Mart.) Fee, (Mem. foug.

5) Gen. fil. 1 78. 1852. Figure 3.

Gymnogramma reniformis Mart., Icon. pi. crypt. 88,

/. 26. 1834. TYPE: Colombia, (Amazonas), Mt.

Cupati, Rio Japura, ("Brasilia"), Martius (holo-

type, M or BR; isotype, HBG; photos, GH, us; frag.,

us!).

San Martin: Monte Guayrapurima, Tarapoto, Spruce

4641 (B, GH, K, NY). Loreto: Cerro Isco, Vie 6887 (B, p).

Cuesta de Yento, near Balsa Puerto, Raimondi 25 (B).

2. Pterozonium brevifrons (A. C. Sm.) Lell., Mem.

New York Hot. Card. 17: 12. 1967.

Stem with yellowish to reddish brown tri-

chomes. Leaf ca. 5-40 cm long, petiole dark red-

dish brown to blackish, with pale ridges extending

downward from the base of the lamina. Lamina

coriaceous, suborbicular to orbicular, cuneate to

deeply cordate at the base, with flabellate vena-

tion. Sori borne along part of a vein, in a band

back of the margin.

Sandstone rocks, ca. 1000 m, San Martin and

Loreto.

Venezuela, Colombia, and Peru.

Syngramma brevifrons A. C. Sm., Bull. Torrey Bot.

Club 57: 178. 1938. TYPE: Venezuela, (Bolivar),

Mount Roraima, Tate 421 (holotype, NY; frag.,

us!).

Stem with dark brown rigid scales. Leaf ca. 10-

50 cm long, petiole dark reddish brown to black-

ish, grooved toward its apex. Lamina coriaceous,

elongate, narrowly ovate to nearly ovate, abruptly

rounded to slightly cordate at the base. Sori ex-

tending along most of a vein from or near the costa

to back of the margin.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

15

In rock crevices and in rocky soil, ca. 2200 m,

Amazonas and San Martin.

Venezuela, Colombia, and Peru.

Amazonas: Prov. Chachapoyas, 3-6 km W of Moli-

nopampa. Wurdack 1408 (F, GH, USM), 1409 (us). San

Martin: Prov. Rioja, Pedro Ruiz-Moyobamba, D. Smith

4796 (GH).

IV. Pityrogramma

Pityrogramma Link, Handbuch 3: 19. 1833. TYPE:

Pityrogramma chrysophylla (Sw.) Link (Ac-

rostichum chrysophyllum Sw.). Figure 4.

Trismeria Fee, (Mem. foug. 5) Gen. fil. 164. 1852.

TYPE: Trismeria aurea Fee = Pityrogramma tri-

foliata (L.) Tryon (Acrostichum trifoliatum L.).

Terrestrial. Stem small to often stout, decum-

bent or erect, bearing scales, especially at the apex.

Leaves borne in a crown or cluster. Lamina deeply

pinnatifid to 4-pinnate-pinnatifid, usually with

white or yellow farinose indument abaxially, less

often pubescent or rarely glabrous, veins free. Spo-

rangia borne along the veins, somewhat back of

the nearly unmodified margin, at maturity often

confluent apparently over most of the surface, in-

dusium and paraphyses absent.

The 1 6 species of Pityrogramma are native to

America, Africa, and Madagascar; there are 1 1

species in America and five in Peru. A few species

are adventive in the Old World, especially the

widely distributed P. calomelanos. Hybrids, some

of them mentioned in Comments, are probably

more frequent than our present information sug-

gests.

Reference

TRYON, R. 1962. Pityrogramma (including Tris-

meria) andAnogramma. Contr. Gray Herb., 189:

52-76.

Key to Species of Pityrogramma

a. Secondary and (if present) tertiary segments moderately or not toothed, or with broad lobes with

several veins b

b. Petiole with 2, or in large leaves more, vascular bundles toward the base, these roundish, oval,

to C-shaped; pinnae with numerous pinnately arranged lobes or segments c

c. Rachis, pinna-rachises, and petiole glabrous, farinose, or thinly and irregularly pubescent . d

d. Rachis and petiole deep reddish brown to blackish e

e. Pinnae equilateral; the pinnules ascending, those on the basiscopic side more strongly

so than those on the acroscopic side; apical pinnae strongly ascending

1 . P. calomelanos

e. Pinnae inequilateral; the pinnules on both sides of the pinna at nearly right angles to the

pinna-rachis (or in large leaves the tertiary segments so disposed on the pinnule-rachis);

apical pinnae nearly or quite at right angles to the rachis 2. P. tartarea

d. Rachis and upper portion of the petiole, or all of it except the base, straw-colored to light

brown 3. P. chrysoconia

c. Rachis and pinna-rachises densely and persistently lanate-pubescent, the petiole similar but

sometimes the pubescence deciduous 4. P. ferruginea

b. Petiole with 2 nearly C-shaped vascular bundles toward the base, these with the central portion

of each curved toward the center of the petiole; pinnae usually entire to 3-(rarely 5-7-)foliolate

5. P. trifoliata

a. Secondary and tertiary segments strongly laciniate into linear lobes, each with 1 vein

6. P. pearcei

1. Pityrogramma calomelanos (L.) Link, Hand-

buch 3: 20. 1833. Figure 4b-c.

Leaf ca. 0.5-1 m long, its axes glabrous to thinly

farinose or thinly pubescent, petiole and rachis

deep reddish brown to blackish. Lamina usually

narrowly lanceolate to ovate-lanceolate, to long-

triangular, 1-pinnate-pinnatifid to 3-pinnate. Pin-

nae equilateral, equally developed on both sides,

pinnules ascending, the abaxial surface rarely gla-

16

FIELDIANA: BOTANY

5 mm

5cm

FIG. 4. Pityrogramma tartarea: a. habit. Pityrogramma calomelanos: b, portion of rachis and 2 pinnae; c, ultimate

segment, abaxial side, (a from Gentry el al. 39771, p, b from LI. Williams 1780, F, c from LI. Williams 2201, F.)

brous, usually white, lemon yellow, bright yellow,

orange-yellow, or pale roseate farinose, or pubes-

cent.

A plant from Dept. Huanuco, Tingo Maria,

Tryon & Tryon 5242 (GH) has some leaves glabrous

abaxially, others white-farinose, and one that is

pubescent abaxially.

Variety austroamericana and var. ochracea usu-

ally grow above 1500 m, and var. calomelanos

most often grows under 1 000 m. The latter is widely

distributed in the American tropics and seems to

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

17

grow more rapidly and is more often an occupant

of disturbed habitats than the other varieties.

Some plants of Pityrogramma calomelanos are

not easily distinguished from P. tartarea, and these

may represent hybrids of the species.

Key to Varieties

a. Lamina farinose abaxially, the color white, pale yellow, or roseate, or rarely the lamina glabrous

abaxially la. var. calomelanos

a. Lamina farinose abaxially, the color bright yellow to orange-yellow, or lamina pubescent abaxially

b

b. Lamina bright yellow to orange-yellow farinose abaxially Ib. var. austroamericana

b. Lamina slightly to densely pubescent abaxially, especially on the fertile segments

. Ic. var. ochracea

la. Pityrogramma calomelanos var. calomelanos

AcrostichumcalomelanosL.,Sp.pl. 1072. 1753. TYPE:

LINN 1245.19; photos, A, GH are this species.

Acrostichum ebeneum L., Sp. pi. 1071. 1753. LEC-

TOTYPE (designated by Tryon, Contr. Gray

Herb. 189: 60. 1962): Sloane, Voy. Jamaica /. 53,

f.l.

Gvmnogramma calomelanos var. denudata Harr., J.

Linn. Soc., Bot. 16: 37. 1877. TYPE: Peru, (Lor-

eto), Pebas, Steere (holotype, K.!; photo, GH; iso-

type, GH!).

Pityrogramma ebenea (L.) Proctor, Brit. Fern Gaz. 9:

219. 1965.

Cleared hillsides, road banks, river banks, thick-

ets, and sometimes in forests or swamps, 100-

2300 m, most commonly below 1000 m, Tumbes

south to Cuzco and Madre de Dios.

Tropical America.

In the Report of the Standing Committee on the

Stabilization of Specific Names (Taxon 24: 1 7 1-

177. 1975), the earlier lectotypification of Pity-

rogramma ebenea was upheld. The latter one by

Proctor (of LINN 1245.14), which would make

the name the correct one for Pityrogramma tar-

tarea, was rejected.

Tumbes: Between Tumbes and Caucho, Coronado 224

(GH, uc). Prov. Zarumilla, Bosque Nacional de Tumbes,

Simpson & Schunke 462 (F, GH). Amazonas: Prov. Bagua,

Aramango, Lopez el al. 4161 (GH). San Martin: 4 mi E

of Tarapoto, Woytkowski 35212 (MO, uc). Lamas, near

Tarapoto, LI. Williams 6360 (F, GH). Loreto: Above Pon-

go de Manseriche, Mexia 6101 (F, GH, MO, uc). Iquitos,

Tryon & Tryon 5167 (BM. F, GH, u, us, USM). Huanuco:

Tingo Maria, Tryon & Tryon 5244 (BM, F, GH, u, us,

USM). Pasco: Puerto Yessup (as Junin), Killip & Smith

26254 (F, us). Junin: La Merced, Killip & Smith 23389

(us). Ucayali: Bosque von Humboldt, Gentry el al. 36354

(F, MO). Ayacucho: Near Kimpitiriki, Killip & Smith 22958

(us). Cuzco: Valle de Marcapata, Herrera 11 99 (us). Ma-

chu Picchu to Quillabamba, Mexia 8089a (uc). Rio Apu-

rimac, between San Martin and Hacienda Luisiana, Da-

vis el al. 1329 (F, GH). Madre de Dios: Prov. Tambopata,

SSW of Puerto Maldonado, Barbour 4902 (F, MO). Par-

que Nacional del Manu, Foster et al. 3409 (GH).

Ib. Pityrogramma calomelanos var. austroamer-

icana (Domin) Farw. Amer. Midi. Naturalist

12:280. 1931.

Pityrogramma austroamericana Domin, Publ. Fac. Sci.

Univ. Charles 88: 7. 1928; also in Bull. Misc.

Inform. 1929: 221. TYPE: Bolivia, Mandon

1549bis (holotype, K!; photo, GH; isotype, GH!).

Open rocky places, grassy slopes, clay banks,

thickets, and rarely along irrigation ditches, 200-

2800 m, Piura south to Puno.

Costa Rica; mountainous regions of South

America, south to Argentina and Brazil.

The formerly accepted name, var. aureoflava

(Hooker) Bailey, has been correctly typified by a

specimen of Pityrogramma chrysophylla: See-

mann 945 (cited as 948), K!. Panigrahi (Kew Bull.

30: 657-667. [1975] 1976) has discussed this and

it agrees with my own later study of the typification

at Kew.

Piura: 37 km E of Olmos, on road to Pucura, Gentry

et al. 22624 (F, MO). Lambayeque: Prov. Lambayeque,

Abra Porculla Pass, Hutchison & Wright 3480 (GH, us).

Cajamarca: Entre Abra de Porculla y Jaen, Ferreyra 13638

(GH, USM). La Libertad: Prov. Trujillo, Hacienda Mo-

chal, Sagdstegui 425 (GH). Huanuco: Between Chinchao

and Puente Durand, Coronado 89 (GH, uc). Yanano,

Macbride 3667 (F, us). Junin: La Merced, Soukup 2368

(F, GH), 3410 (GH, us). Ayacucho: Ayna, between Huanta

and Rio Apurimac, Killip & Smith 22697 (F, GH, us).

Cuzco: Near Quillabamba, Mexia 8045 (F, GH, MO, us).

Potrero, 8 km W of Quillabamba, Tryon & Tryon 5365

18

FIELDIANA: BOTANY

(BM, F, GH, u, us, USM). Puno: Prov. Sandia, abajo de

Sandia, Ferreyra 16657 (GH).

lc. Pityrogramma calomelanos var. ochracea

(Presl) Tryon, Contr. Gray Herb. 189: 61.

1962.

Gymnogramma ochracea Presl, Reliq. haenk. 1: 17.

1825. LECTOTYPE (designated by Tryon, Contr.

Gray Herb. 189: 62. 1962): Peru, Haenke (ho-

lotype, Haenke, PR Herb. 24360; photos, GH, us;

probable isolectotypes, B, K.).

Ceropteris adiantoides var. peruviana Hieron., Hed-

wigia 48: 221. 1909. TYPE: Peru, (San Martin),

Moyobamba to Rio Huallaga, Stiibel 1096 (ho-

lotype, B).

Pityrogramma perelegans Domin, Publ. Fac. Sci. Univ.

Charles 88: 8. 1928. TYPE: Peru, (San Martin),

Tarapoto, 1855-1856, Spruce (holotype, K!; pho-

to, GH).

Pityrogramma ochracea (Presl) Domin, Publ. Fac. Sci.

Univ. Charles 88: 8. 1928.

Open rocky places, river banks and road banks,

cleared land, less often in thickets, in forests or at

forest borders, rarely along irrigation ditches, 400-

3000 m, or in La Libertad at 50 m along an irri-

gation ditch, La Libertad south to Puno.

Honduras; Venezuela and Colombia, south to

Bolivia.

The variation in spores of this variety is unusu-

al; they are often typical of the genus, with well-

defined dark ridges, but sometimes they are smooth

or only slightly roughened.

La Libertad: Huaca del Sol, Sagdstegui 393 (GH). San

Martin: Juan Jui, alto Rio Huallaga, Klug 4177 (F, GH,

MO, uc). Huanuco: Tingo Maria, Tryon & Tryon 5243

(BM, F, GH, u, us, USM). Pasco: Villcabamba, Macbride

4994 (F, GH, us). Junin: La Merced, Macbride 5276 (F,

us); Soukup 3411 (GH); Cerrate2833 (GH, USM). Ucayali:

La Divisoria (as San Martin), Aug. 3, 1942, Ridoutt (GH,

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

rimac, Killip & Smith 22667 (us). Cuzco: Potrero, 8 km

W of Quillabamba, Tryon & Tryon 5366 (BM, F, GH, u,

us, USM). Madre de Dios: Prov. Manu, Salvation to Shin-

tuya, Plowman & Davis 5079 (GH). Puno: Near Puno,

Soukup 449 (F). Prov. Carabaya, Puente Inambari, Var-

gas 18416 (GH).

2. Pityrogramma tartarea (Cav.) Maxon, Contr.

U.S. Natl. Herb. 17: 173. 1913. Figure 4a.

Leaf usually 0.4-1 m long, its axes glabrous or

thinly farinose or thinly pubescent, petiole and

rachis deep reddish brown to blackish. Lamina

long-triangular to deltoid, 1 -pinnate-pinnatifid to

3-pinnate. Pinnae inequilateral, the basiscopic side

better developed, pinnules at nearly right angles

to the pinna-rachis, the abaxial surface usually

white, pale yellow, or bright yellow-farinose, rarely

pubescent.

The incorrect use of Pityrogramma ebenea (L.)

Proctor for this species has been discussed under

P. calomelanos.

Key to Varieties

a. Lamina farinose abaxially, the color white to pale yellow, rarely almost colorless or slightly pubescent

abaxially 2a. var. tartarea

a. Lamina farinose abaxially, the color bright yellow, or pubescent abaxially b

b. Lamina bright yellow-farinose abaxially 2b. var. a u rat a

b. Lamina pubescent abaxially 2c. var. jamesonii

2a. Pityrogramma tartarea var. tartarea

Acrostichum tartareum Cav., Descr. pi. 242. 1802.

TYPE: Peru, (Lima), near "Guamantanga" (Hua-

mantanga), Nee (holotype, seen by C. Chr. at MA,

Dansk Bot. Ark. 9(3): 10. 1937; the upper frag-

ments of Herb. Willd. 19568-3, B!; photos, GH,

us, are probably from the holotype).

Gymnogramma peruviana Desv., Ges. Naturf. Freunde

Berlin Mag. Neuesten Entdeck. Gesammten Na-

turk. 5: 329. 1811. TYPE: Peru, Joseph deJussieu

(holotype, Herb. Jussieu 1009, P!; photos, GH, us).

Pityrogramma peruviana (Desv.) Maxon, Contr. U.S.

Natl. Herb. 17: 173. 1913.

Pityrogramma calomelanos var. peruviana (Desv.)

Farw., Amer. Midi. Naturalist 12: 280. 1931.

Rocky and shrubby hillsides, in crevices of rocks

and on Inca walls, on cliffs, 800-4000 m, Piura

south to Puno.

Tropical America, south to Bolivia and Brazil.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

19

Piura: Prov. Huancabamba, Loma Redondo, Sapa-

lache-Chinguela, Sagdstegui et al. 10181 (F, MO). Arriba

de Canchaque, Ferreyra 3106 (GH, USM). Lambayeque:

29 km from Olmos on road to Jaen, Correll & Smith

P803 (GH). Cajamarca: Prov. Celendin, Guanambra-

Sendamal. Sagdstegui et al. 12200 (F, GH). Amazonas:

Prov. Chachapoyas, Cerros Calla Calla, Hutchison &

Wright 4819(F, GH). I A Libertad: Prov. Pataz, Retamas,

Ldpez & Sagdstegui 3604 (GH). Ancash: Prov. Yungay,

entre Yungay y Llanganuco, Mostacero et al. 1380 (F,

GH). Huanuco: Mito, Macbride & Featherstone 1429 (F,

us). Carpish, Coronado 60 (GH, uc). Lima: Prov. Canta,

4 km from Huamantanga, Sounders 1249 (GH). Pasco:

Prov. Oxapampa, Cordillera San Gotardo, Leon 515

(USM). Junin: Huacapistana, Tryon & Tryon 5429 (BM,

F, GH, u, uc, USM). Cuzco: Prov. Urubamba, Cerro Ma-

chu Picchu, Mexia 8086 (F, GH, us). Puno: Near Puno,

Soukup 450 (F).

2b. Pityrogramma tartarea var. aurata (Moore)

Tryon, Contr. Gray Herb. 189: 65. 1962.

Gymnogramma tartarea var. aurata Moore, Gard.

Chron. 1870: 493. TYPE: Peru, Pearce, cult.

Veitch (holotype, K.!; photo, GH).

Pityrogramma presliana Domin, Publ. Fac. Sci. Univ.

Charles 88: 6. 1928. TYPE: Peru, Matthews 1823

(holotype, K!; photo, GH).

Open places, clay banks, and among rocks, 800-

2200 m, Cajamarca south to Cuzco.

Colombia south to Argentina.

bey (holotype, P!; photo, GH; isotype, B!; photo,

GH; frag., ex B, us!).

Leaf ca. 25-80 cm long, its axes glabrous or

thinly farinose, petiole (at least the apical portion)

and the rachis straw-colored to light brown. Lam-

ina long-triangular, 2-pinnate to 2-pinnate-pin-

natifid. Pinnae inequilateral, the basiscopic side

somewhat better developed, pinnules nearly at right

angles to the pinna-rachis, the abaxial surface

white- or yellow-farinose.

Rocky and shrubby slopes and hillsides, Inca

walls, 2000-3100 m, Piura to Cuzco.

Costa Rica and Panama; Jamaica; Guyana to

Colombia and south to Bolivia.

Plants with white farina on the leaves and those

with yellow farina both occur nearly throughout

the range of the species and there seems to be no

reason to recognize them as varieties.

Piura: Prov. Ayabaca, Ayabaca, Sagdstegui & Caban-

illas 8709 (HUT). Cajamarca: Prov. Celendin, La Tranca-

Gelig, Sagdstegui et al. 12145 (MO, HUT). Amazonas:

Prov. Chachapoyas, Calla Calla, Aguado 6789 (F). Huan-

uco: Mitotambo, arriba de Mito, Ferreyra 10348 (GH,

USM). Mito, Macbride & Featherstone 1393 (us). Huan-

cavelica: Chuspi, Tocas, Tovar 2052 (GH, USM). Cuzco:

Huadquina, Bues 13 19 (us). Prov. Urubamba, Santa Rita,

Vargas 2683 (us).

Cajamarca: Prov. Cutervo, La Achira, Lopez & Sa-

gdstegui 5463 (GH). Huanuco: Villcabamba, Rio Chin-

chao, Macbride 4995 (F, GH, us). Pasco: Prov. Oxapam-

pa, Rio Boqueria, Smith et al. 1816 (MO). Junin:

Chanchamayo valley, Schunke 38 (F, us). Between Hu-

acapistana and San Ramon, Coronado 259 (GH, uc).

Cuzco: Prov. Paucartambo, Tanamayo, Vargas 6479 (uc).

Prov. Paucartambo, near Santa Isabel, Plowman & Davis

4990 (GH).

2c. Pityrogramma tartarea var. Jameson ii (Baker)

Tryon, Contr. Gray Herb. 189: 66. 1962.

Gymnogramma jamesonii Baker, Syn. fil., ed. 2, 516.

1874. TYPE: Ecuador, (Pichincha), Pichinchan

Andes, Jameson (holotype, K!; photo, GH).

Colombia and Ecuador; this variety is expected

in northern Peru.

3. Pityrogramma chrysoconia (Desv.) Domin,

Publ. Fac. Sci. Univ. Charles 88: 10. 1928.

Acrostichum chrysoconium Desv., Mem. Soc. Linn.

Paris 6: 212. 1827. TYPE: Peru, evidently Dom-

4. Pityrogramma ferruginea (Kunze) Maxon,

Contr. U.S. Natl. Herb. 17: 173. 1913.

Gymnogramma ferruginea Kunze, Linnaea 9: 34. 1 834.

TYPE: Peru, (Huanuco), Quebrada de Chinchao,

Jul. 1829, Poeppig, Diar. 1160 (holotype, not lo-

cated; isotype, P!; photos, GH, us).

Eriosorus ruizianus Fee, (Mem. foug. 5) Gen. fil. 152,

t. 13B,f. 2. 1852. TYPE: (Not located).

Leaf ca. 0.25-1 m long, its axes densely and

usually persistently lanate, the petiole sometimes

deciduously so, petiole and rachis atropurpureous

or blackish. Lamina narrowly elliptic, 1-pinnate-

pinnatifid. Pinnae equilateral, the pinnule-lobes

somewhat ascending, the abaxial surface densely

to rarely thinly lanate.

Moist and dry cliffs, rocky banks, 750-2000 m,

Huanuco and Pasco to Ayacucho.

Guatemala to Panama; Peru.

This is the only species of the genus in America

with a markedly disjunct range. Plants of Central

America are more variable than those of Peru where

20

FIELDIANA: BOTANY

the lamina is narrowly elliptic and the pinnae are

subsessile.

Huanuco: Rio Huallaga canon, below Santo Domingo,

Macbride 4261 (F, GH, us). Tingo Maria, Allard 21530

(us). Pasco: Oxapampa, Soukup 1822 (F, us), 3351 (F,

GH). Junin: 8 km SW of San Ramon, Tryon & Tryon

5451 (BM, F, GH, u, uc, us, USM). Ayacucho: Between

Huanta and Rio Apurimac, Killip & Smith 22482 (us).

5. Pityrogramma trifoliate (L.) Tryon, Contr. Gray

Herb. 189: 68. 1962.

Acrostichum trifoliatum L., Sp. pi. 1070. 1753. SYN-

TYPES: LINN 1245.9, photo, A, and Sloane, Voy.

Jamaica /. 45, f. 2 are both this species.

Trismeria microphylla Fee, (Mem. foug. 5) Gen. fil.

165. 1852. TYPE: Peru (not located).

Trismeria trifoliata (L.) Diels, Nat. Pflanzenfam. 1(4):

265. 1899.

Leaf ca. 0.5-1.25 m long, its axes glabrous to

thinly farinose, petiole and rachis light reddish

brown to blackish. Lamina narrowly lanceolate to

usually elongate, 1-2-pinnate. Pinnae equilateral

or inequilateral, the pinnules, when present, as-

cending, the abaxial surface glabrous or usually

white- or yellow- farinose.

Open rocky ground, in gravel, along road bor-

ders, on river banks, along irrigation ditches, rare-

ly in forests, nearly at sea level to 2300 m, Piura

south to Arequipa.

Tropical America.

Hybrids of this species are mentioned in Com-

ments.

Piura: Serran to Canchaque, Ferreyra 10782 (USM).

I.ambayeque: 20 km from Olmos on road to Jaen, Correll

& Smith P796 (GH). Cajamarca: Prov. Contumaza,

alrededores de San Benito, Sagdstegui 3740 (GH). Ama-

/onas: Prov. Bagua, Chiriaco to Puente Venezuela, Bar-

hour 4410 (MO). La Libertad: Trujillo, Osgood & An-

derson 28 (F), 29 (F, us); Worth et al. 8887 (GH, uc, us).

San Martin: Tarapoto, Spruce 4 153 (BM, GH, us). Loreto:

Prov. Alto Amazonas, Andoas, Rio Pastaza, Vdsquez &

Jaramillo 787 (F, MO). Huanuco: Huanuco to Muna,

Mexia 4100 (GH, MO, uc, us). Lima: Chosica, Tryon &

Tryon 5342 (BM, F, GH, MO, NY, u, uc, us, USM). Junin:

La Merced, Killip & Smith 23496 (F, us). Ucayali: Prov.

Coronel Portillo, Bosque von Humboldt, Smith et al.

1339 (F, MO). Ayacucho: Ayna, Killip & Smith 23112

(us). Apurimac: 45 km from Abancay on road to Chal-

Ihuanca, Sounders 764 (GH). Cuzco: Prov. Anta, Sisal to

Cunyacc, Vargas 741 1 (MO, uc). Prov. La Convention,

above Sinechinete, Davis et al. 1286 (F, GH). Arequipa:

Huario, below Chuquibamba, D. Stafford 1 149 (BM).

6. Pityrogramma pearcei (Moore) Domin, Publ.

Fac. Sci. Univ. Charles 88: 9. 1928. Gym-

nogramma pearcei Moore, Card. Chron. 1 864:

340. TYPE: Pearce cult. Veitch (Pearce 274

K! and "Gymnogramma pearcei. n.sp" K!;

photos, GH, are authentic).

Lamina ovate-lanceolate to long-triangular,

highly dissected, to 4-pinnate, with slender ulti-

mate lobes, each with 1 vein, pinnae equilateral

or nearly so, glabrous.

The original material of this species may have

been collected in Peru. It has not been gathered

again. The species has recently been collected in

Costa Rica, Panama, and Colombia along dis-

turbed roadsides.

Comments

Four hybrids have been proposed with Pityro-

gramma trifoliata as one parent (Tryon, 1962).

These hybrids all have the vascular bundles of P.

trifoliata, at least in large leaves, and also the

strongly ascending-arching pinna-stalks of that

species, at least those of the lower pinnae. They

also have an elongate lamina and the pinnae with

numerous pinnules.

Pityrogramma calomelanos var. ochracea x P. tri-

foliata

Petiole and rachis essentially glabrous, segments

somewhat pubescent abaxially.

Junin: Prov. Tarma, 3 km N of La Merced, Tryon &

Tryon 5440 (GH). Cuzco: Bties 1936 (us). Prov. La Con-

vencion, Chaupimayo, Bties 1433 (GH).

Pityrogramma ferruginea x P. trifoliata

Petiole and rachis deciduously lanate, segments

sparsely lanate abaxially, pinnae irregularly pin-

nate or lobed. Known from a single plant.

Junin: Prov. Tarma, 10 km SW of San Ramon, Tryon

& Tryon 5449 (GH).

Pityrogramma calomelanos var. calomelanos x

P. trifoliata and P. calomelanos var. austroamer-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

21

FIG. 5. Anogramma leptophylla: a, habit; b, fertile lamina, abaxial side. (From Tryon & Tryon 5416, F.)

it. ana x P. trifoliata are both known in Argentina

and may be found in Peru.

V. Anogramma

Anogramma Link, Fil. spec. 137. 1841. TYPE:

Anogramma leptophylla (L.) Link (Polypodi-

um leptophyllum L.). Figure 5.

Terrestrial. Stem very small, erect, bearing tri-

chomes or trichomes and scales. Leaves borne in

a cluster. Lamina to 4-pinnate, glabrous (or, in 1

species, pubescent), veins free. Sporangia in rather

elongate sori on the veins, somewhat back of the

unmodified margin, indusium and paraphyses ab-

sent.

Anogramma is a rather geographically disjunct,

nearly pantropical genus of five or six species. A

single species is in Peru and another may well be

discovered there. The species are evidently annual,

new plants arising each growing season by spores

or by the growth of a dormant gametophyte. The

stem is much reduced and bears only leaves of the

growing season, petioles of previous years being

absent. Leaf size and architecture is variable.

Reference

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

gramma, pp. 224-228, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Species of Anogramma

a. Spores dark brown; stem and usually the base of the petiole only with trichomes; lamina usually

narrowly to broadly ovate, to somewhat deltoid, the ultimate lobes somewhat obtuse

1 . A. leptophylla

a. Spores tan; stem and usually the base of the petiole with scales and trichomes; lamina usually ovate

to often deltoid, the ultimate lobes mostly acute A. chaerophylla

22

FIELDIANA: BOTANY

1 . Anogramma leptophylla (L.) Link, Fil. spec. 1 37.

1841. Figure 5.

Polypodium leptophyllum L., Sp. pi. 1092. 1 753. LEC-

TOTYPE (designated by Morton, Amer. Fem J.

60: 103. 1970): LINN 1251.56; photo, A.

Stem very small, bearing a few trichomes, which

are usually also at the base of the petiole. Leaves

to ca. 20 cm long. Lamina herbaceous, narrowly

ovate to somewhat deltoid, often 2-pinnate, with

bifid, obtuse lobes. Spores dark brown.

Among rocks and at the edge of boulders (lomas

of Dept. Lima), 350-600 m, and Inca walls (Ma-

chu Picchu, Dept. Cuzco), 2200-2400 m, Lima

and Cuzco.

Mexico and Central America; scattered in South

America; Old World east to New Zealand.

Lima: Prov. Chancay, Loma Lachay, Tryon & Tryon

5416 (BM, F, OH, u, us, USM), Coronado 14 (GH, uc, us).

Prov. Chancay, Loma Granados, Stork & Vargas 9333

(GH, uc). Prov. Canete, Loma Quilmana, Coronado 26

(GH, uc, us). Cuzco: Prov. Urubamba, Machu Picchu,

Mexia 8083 (F, GH, MO, uc); Coronado 94 (GH, uc, us).

Comments

Anogramma chaerophylla (Desv.) Link is re-

ported from Bolivia and is expected to grow in

southern Peru. It has a wide but rather disjunct

range in tropical America. It is usually a larger

species than A. leptophylla, with leaves to 40 cm

long. Other distinctions of the two species are pre-

sented in the key.

MUdella Trev., Rendic. R. 1st. Lombardo II, 9: 810.

1876. TYPE: MUdella intramarginalis (Link)

Trev. (Pteris intramarginalis Link) = Cheilanthes

intramarginalis (Link) Hooker.

Stem erect to long-creeping, with light brown,

concolorous or bicolorous to wholly dark-sclerot-

ic, linear to ovate-lanceolate, usually thin scales.

Leaves usually monomorphic. Petiole with 1 or 2

vascular bundles near the base, terete to adaxially

sulcate or 2-ridged. Lamina 1-5-pinnate, or less

often pedate and to at least 3-pinnatifid, glabrous,

pubescent, scaly glandular, or white-, yellow-, or

orange-farinose abaxially. Veins free or rarely

anastomosing without included veinlets, ending at

or near the margin, the ends not to clearly enlarged.

Sporangia usually in sori at or near the vein ends,

or rarely single, or in soral lines along an incom-

plete to complete marginal commissure, exindu-

siate, or 1-many sori covered by a slightly to well-

modified marginal indusium, paraphyses absent.

Spores tetrahedral-globose to globose, trilete, usu-

ally cristate, sometimes rugose, verrucate, tuber-

culate or echinate.

Cheilanthes is a nearly worldwide genus of about

1 50 species, or perhaps more. It is especially well

developed in semiarid regions, with centers of

species in Mexico, the Andes, and in southern Af-

rica. It is a diverse genus, with some of its elements

often placed in the genera Pellaea or Notholaena.

Species 1-2 1 of Notholaena in Tryon ( 1 956) and

species 1-10 in Tryon (1964) are now placed in

Cheilanthes.

VI. Cheilanthes

Cheilanthes Sw., Syn. fil. 5, 126. 1806, conserved

name. TYPE: Cheilanthes micropteris Sw.

Figure 6.

Aleuritopteris Fee, (Mem. foug. 5) Gen. fil. 153. 1852.

TYPE: Aleuritopteris farinosa (Forssk.) Fee (Pter-

is farinosa (Forssk.) = Cheilanthes farinosa

(Forssk.) Kaulf.

References

TRYON, R. 1956. A revision of the American

species of Notholaena. Contr. Gray Herb., 179:

1-106.

TRYON, R. 1964. The ferns of Peru. Contr. Gray

Herb., 194: 88-121.

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

lanthes, pp. 249-26 1 , in Ferns and allied plants,

Springer- Verlag, New York.

Key to Species of Cheilanthes

a. Pinnae densely pubescent abaxially, the trichomes concealing the leaf tissue b

b. Trichomes simple c

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

23

c. Pinnae 1 -pinnate, with appressed-ascending trichomes abaxially 8. C. obducta

c. Pinnae pinnatifid or 2-pinnatifid at the base, tomentose abaxially d

d. Stem rather slender, long-creeping; pinnae nearly deltoid; tips of the stem scales light brown

6. C. fraseri

d. Stem short, compact, multicipital; pinnae nearly oblong; tips of the stem scales dark brown

7. C. bonariensis

b. Trichomes stellate 9. C. mollis

a. Pinnae glabrous, glandular, farinose, or thinly pubescent abaxially and the leaf tissue evident, or

densely scaly abaxially and the scales concealing the leaf tissue e

e. Pinnae glabrous, glandular, farinose, or thinly pubescent abaxially f

f. Rachis pubescent or with trichome-like fibrils, at least adaxially g

g. Rachis densely pubescent with short, stout trichomes, or with very short trichome-like

processes 5. C. pruinata

g. Rachis loosely pubescent with long trichomes or trichome-like fibrils h

h. Rachis and pinnae long-pubescent with patent, whitish to light brown trichomes . . . . i

i. Lamina with few, mostly sessile pinnae; stem scales whitish; petioles breaking evenly

near their base 3. C. fractifera

i. Lamina with few to many pinnae, most of the pinnae stalked; stem scales dark brown;

petioles breaking irregularly 4. C. pilosa

h. Rachis, at least on the adaxial side, and the pinnae somewhat pubescent, with brown,

irregularly oriented trichomes and trichome-like fibrils j

j. Pinna-rachises atropurpureous to blackish on the adaxial side, except sometimes

greenish apically 1 . C. moritziana

j. Pinna-rachises, greenish on the adaxial side, except sometimes dark at its base ....

2. C. notholaenoides

f. Rachis, or the primary axis of the lamina, glabrous, glandular, or farinose k

k. Lamina pedate, 3-pinnatifid at the base 15. C. concolor

k. Lamina pinnate 1

1. Pinnae whitish-farinose abaxially 14. C. farinosa

1. Pinnae glabrous or with scattered sessile glands m

m. Indusia confined to the segments n

n. Segments with sessile glands abaxially 11. C. rufopunctata

n. Segments glabrous abaxially o

o. Stem scales brown, concolorous; each sorus covered by a separate indusium

10. C. orbignyana

o. Stem scales bicolorous, with narrow lighter margins; many sori covered by a

long, continuous indusium 13. C. poeppigiana

m. Indusia extending onto the segment stalk 12. C. marginata

e. Pinnae densely scaly abaxially, the scales concealing the leaf tissue p

p. Petiole with 2 vascular bundles at the base; pinnae lobed or shallowly pinnatifid, with an

underlayer of dissected scales on the abaxial side 23. C. sinuata

p. Petiole with 1 vascular bundle at the base; pinnae 1 -pinnate or more complex, rarely pinnatifid,

lacking a pronounced underlayer of dissected scales on the abaxial side q

q. Stem scales bicolorous with a dark center and narrow lighter margins r

r. Ultimate segments small, beadlike, suborbicular, their adaxial surface nearly glabrous

1 6. C. myriophylla

r. Ultimate segments rather large, nearly oblong, often lobed, their adaxial surface sparsely

hirsute with whitish, nearly rigid trichomes 17. C. cantangensis

q. Stem scales light brown or yellowish to reddish brown, concolorous s

s. All ultimate segments small, nearly suborbicular, or at least the basal segments of the

secondary segments suborbicular; stem scales reddish t

t. All segments glabrous above, the scales that more or less cover the adaxial surface

borne on the abaxial surface; all segments suborbicular or nearly so

.18. C. scariosa

24 FIELDIANA: BOTANY

t. Segments with narrow scales and flattened trichomes adaxially to glabrate; some to

several segments oblong, especially apical ones; scales abaxially not clathrate

2 1 . C. incarum

s. Many or most ultimate segments nearly oblong, few or no suborbicular ones u

u. Scales on the segments abaxially moderately dentate-ciliate or short-ciliate v

v. Stem scales yellowish to rarely brown; lamina ca. 8-20 cm long, mostly longer

than the petiole (rarely equal in length); scales on the pinnules abaxially mostly

not clathrate, the surface smooth, or clathrate and the surface roughened only at

the base 19. C. peruviana

v. Stem scales reddish; lamina ca. 1.5-5 cm long, mostly shorter than the petiole

(rarely equal in length); scales on the pinnules abaxially mostly clathrate and the

surface roughened 20. C. arequipensis

u. Scales on the segments abaxially freely long- or somewhat short-ciliate, not clathrate,

the surface smooth; stem scales brown 22. C. lonchophylla

1 . Cheilanthes moritziana Kunze, Linnaea 23: 307.

1850. LECTOTYPE (designated by Tryon,

Contr. Gray Herb. 194: 98. 1964): Venezuela,

(Dist. Federal, Caracas), La Guayra, Moritz

263 (holotype, B!; photo, GH; isotype, GH!).

Stem rather slender, creeping, sometimes knot-

ted, scales lanceolate to lance-ovate, light brown

to brown, concolorous. Leaves ca. 1 5—40 cm long,

petiole terete, sometimes fibrillose, especially on

the adaxial side, the rachis similar, lamina nar-

rowly lanceolate, 2-pinnate-pinnatifid to 3-pin-

nate. Pinnae only slightly pubescent on both sides.

Indusium more or less crispate, continuous.

Sheltered, rocky places, 1600-3400 m, Caja-

marca and Amazonas to Cuzco.

Venezuela to Colombia and Bolivia.

The differences from the closely related C. noth-

olaenoides are discussed under that species.

Cajamarca: Alrededores de San Benito, Sagdstegui

3739 (GH). Celendin, Stiibel 1045 (B). Cantanoe, Celen-

din to Rio Maranon, Lopez & Sagdstegui 3365 (GH, HUT).

Amazonas: Leimebamba, valle de Utcubamba, Stiibel

1021 (B). San Martin: Salinas de Pilluana, Ule 6900 (B).

29 km S of Tarapoto, Gentry & Smith 45015 (F). Huan-

uco: Muna, Macbride 3921 (F, us). Prov. Dos de Mayo,

Chiquia, Huapalla 3649 (USM). Junin: Yungay, Woyt-

kowski 6586 (MO, us). Cuzco: Yucay, Soukup 918 (F).

a darker center. Leaves 10-20 cm long, petiole

terete, pubescent, as is the rachis, lamina narrowly

lanceolate, mostly 2-pinnate, to 2-pinnate-pin-

natifid. Pinnae slightly pubescent to nearly gla-

brous. Indusium rather broad, more or less cris-

pate, continuous.

Cliffs and rocky places, 2600-3300 m, Piura to

Junin.

West Indies; Mexico to Venezuela, Peru and

Argentina.

This species is closely related to the previous,

C. moritziana; the character of the pinna-rachis

given in the key provides a sufficient separation.

In addition, the lamina of C. notholaenoides is

rather regularly 2-pinnate and the pinnae are pin-

nate nearly to the tip, while in C. moritziana the

lamina is 2-pinnate-pinnatifid and the pinnae have

a definite pinnatifid apex.

Piura: China valley, Rauh PI 93 5 (B). Cajamarca: 52

km W of Cajamarca on road to Chilete, Correll & Smith

P838 (GH). Prov. Celendin, above Balsas, road to Ce-

lendin, Hutchison & Wright 5302 (GH). Pasco: Yana-

huanca, Macbride & Featherstone 1242 (F, GH, us). Jun-

in: Ingahuasi, between Huancayo and Izcuchaca, Tovar

3870 (GH, USM). Uspachaca, Macbride & Featherstone

1307 (F, us).

2. Cheilanthes notholaenoides (Desv.) Weath.,

Contr. Gray Herb. 1 14: 34. 1936.

Pteris notholaenoides Desv., Mem. Soc. Linn. Paris

6: 298. 1827. TYPE: "Hispaniola" (holotype, P!;

photo, GH).

Stem rather slender, creeping, more or less knot-

ted, scales lance-subulate, brown, with or without

3. Cheilanthes fractifera Tryon, Rhodora 62: 7,

/. 7252. January, 1960. TYPE: Peru, Dept.

Ayacucho, Correll & Smith PI 69 (holotype,

GH!).

Cheilanthes saundersii Alston, Lilloa 30: 110, /. 6.

August, 1960. TYPE: Peru, Dept. Lima, Huar-

ochiri, Sounders 350 (or 353 according to Saun-

ders, pers. comm.) (holotype, BM; isotype(?),

Sounders 353, us!).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

25

5 mm

Flu 'n ' Cheilanthes Poeppigiana: a, habit; b, basal pinnule and portion of rachis, abaxial side. Cheilanthes my-

nopnytla: c, portion of rachis and pinna, abaxial side, (a, b from Macbride 3672, F, c from Killip & Smith 21796, F).

26

FIELDIANA: BOTANY

Stem short, decumbent, scales narrowly lanceo-

late, light brown, concolorous. Leaves ca. 5-12 cm

long, the petiole more or less grooved on the adax-

ial side, the ridges rounded and concolorous with

the petiole proper, thinly whitish pubescent as is

the rachis, with large whitish scales at the base,

lamina deltoid to broadly ovate, 2-pinnate-pin-

natifid. Pinnae moderately whitish pubescent

abaxially, thinly pubescent adaxially. Indusium

more or less crenate or of small lobes, well mod-

ified.

Rocky hillsides, 1 600-2200 m, Cajamarca south

to Arequipa.

Endemic to Peru.

This species was first discovered in 1954 by S.

G. E. Saunders and since then it has been collected

several times. The petiole that fractures evenly

toward the base, and the large whitish scales at the

base of the petiole combine to make this a dis-

tinctive species among those of Peru.

Cajamarca: Prov. Contumaza, Cascas-Chepate, Sa-

gastegui 4482 (GH). Lima: Dist. Lange, Saunders 1050

(GH). Dist. Surco, Rimac valley, Saunders 219 (BM, GH).

Rimac valley, Rauh & Hirsch PI 43 (B). Arequipa: Cerros

de Caldera, Rauh & Hirsch P570 (B). Entre Characato y

Yarabamba, G. & C. Miiller 12592 (GH).

4. Cheilanthes pilosa Goldm., Nov. Actorum

Acad. Caes. Leop.-Carol. Nat. Cur. 19, suppl.

1 (Meyen, Obs. bot.): 455. 1843. TYPE: Peru,

Meyen (holotype, B!; photos, GH, us; isotype,

S-PA!).

Cheilanthes macleanii Hooker, Sp. fil. 2: 93, t. 11 OB.

1852. TYPE: Andes of Peru, John Maclean (ho-

lotype, K!; photos, GH, us; isotype, GH!).

Cheilanthes andina Hooker, Sp. fil. 2: 115. 1852.

TYPE: Andes of Peru, John MacLean (holotype,

K!; photos, GH, us).

Stem rather slender, short-creeping, scales lance-

subulate, dark brown, rather sclerotic, with or

without paler borders. Leaves ca. 10-40 cm long,

petiole terete, somewhat pubescent, the rachis

similar but sometimes grooved on the adaxial sur-

face, lamina lanceolate to narrowly so, 2-3-pin-

nate. Pinnae pubescent adaxially and more so

abaxially. Indusium broad, deeply crispate or lobed.

On ledges and in crevices of cliffs, 2300—4400

m, Ancash to Puno.

Peru to Argentina.

The soft, spreading trichomes on the leaf and

the rigid, shining stem scales are characteristic of

this species. Cheilanthes andina is a variant of C.

pilosa, and their differences are perhaps related to

the growth of C. andina under drier and more

exposed conditions. The ultimate segments in C.

andina are less expanded and thicker in texture,

and the trichomes are often shorter.

Two collections evidently represent a hybrid be-

tween C. pilosa and C. pruninata: Cuzco, Visa-

chani, Sues 1782 (GH) and Cuzco, Prov. Calca,

Vargas 17409 (GH). These have, among other

characters, the broad indusia of the former species

and mostly the characteristic short-pubescent pet-

iole and rachis of the latter species.

Ancash: Chiquian, Cerrate 1548 (USM). Between Lla-

mac and Jahuacocha, Cerrate 2337 (USM). Prov. Huay-

las, Huascaran National Park, Smith et al. 12141 (F).

Lima: Prov. Huarochiri, Wankalasila, Cerrate et al. 4412

(USM). Junin: Near Huancayo, Killip & Smith 23365 (us),

Saunders 647 (GH). 15 km E of Huancayo, Tryon &

Tryon 5467 (F). Huancavelica: Cerro Santa Barbara, near

Huancavelica, Tovar 3117 (GH, USM). Cuzco: Cerro de

Cusilluyoc, Pennell 13994 (F, GH, us). Velille, Vargas

6541 (cuz). Valle de Lares, Bues 1825 (us). Puno: Puno,

Mexia 7783 (F, GH, MO, uc, us). Araranca, Pennell 13454

(F, GH, us). Sandia, Weberbauer 717 (B).

5. Cheilanthes pruinata Kaulf., Enum. fil. 210.

1824. TYPE: "Peru" (not located).

Cheilanthes mathewsii Kunze, Farrnkrauter 1 : 50, t.

25. 1848. TYPE: Peru, Mathews 605 (holotype,

BM!; photo, GH, this specimen, identified by Kunze,

may be the holotype, or it may serve to replace

the one probably destroyed at LZ).

Cheilanthes fasciculata Goldm., Nov. Actorum Acad.

Caes. Leop.-Carol. Nat. Cur. 19, suppl. 1 (Meyen,

Obs. bot.): 456. 1843. TYPE: Peru, 9000-12000

ft., Meyen (holotype, B!; photo, GH).

Stem moderately stout, creeping, multicipital,

scales dark reddish brown, concolorous or with

very narrow pale borders. Leaves ca. 20-50 cm

long, petiole terete, short-pubescent, usually

densely so, rachis terete or rarely somewhat

grooved, otherwise like the petiole, lamina linear,

2-3-pinnate-pinnatifid. Pinnae deltoid, decidu-

ously pubescent on both sides except along the

axes and midveins abaxially. Indusium moderate-

ly broad, deeply lobed, crispate, or as separate

lobes on small ultimate segments.

In crevices or on ledges of cliffs and in rocky

soil, 2500-4400 m, Cajamarca south to Puno and

Moquegua.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

27

Peru to Argentina.

The dense, very short pubescence of the petiole

and rachis are especially distinctive characters. The

pinnae are usually deltoid and in drying they are

usually shining-varnished above. This is one of

the most common and widely distributed of the

Altiplano ferns.

Cajamarca: Cajamarca to Cajabamba, Ferreyra 3224

(BM, GH, USM). La Libertad: Above Cachicadan, Stork &

Morton 9975 (F, uc). La Manzanas, Lopez 7779 (uc, us).

Ancash: Chiquian, Ferreyra 5717 (BM), 7304 (USM), Cer-

rate 500 (GH, USM), 3350 (GH, USM). Huanuco: Mito,

Macbride & Featherstone 1490 (F, GH, us). Panao, Fer-

reyra 1943 (USM). Lima: Between Parac and Toncuyo,

Coronado 314 (GH, uc). Matucana, Macbride & Feath-

erstone 420 (F, us). Junin: Tarma, Killip & Smith 21807

(F, us). Near Huancayo, Killip & Smith 23362 (F, GH,

us). Huancavelica: SE of Pampas, Stork & Morton 10251

(F, uc, us). Between Conaica and Laria, Tovar 920 (GH,

USM). Ayacucho: Near Puquio, Ferreyra 7214 (GH, USM).

Apurimac: Andahuaylas, Stork & Morton 10717 (GH; F

and uc, in part). Cuzco: Near Anta, Tryon & Tryon 5362

(BM, F, MO, u, us, USM). Puno: Puno, Mexia 7780 (F, GH,

MO, uc, us). Arequipa: Arequipa, Pennell 13199 (F, GH,

us). 1 2 km S of Arequipa, Eyerdam & Beetle 22126 (GH,

MO, uc, us ). Moquegua: Prov. Mariscal Nieto, between

Torata and Carumas, Dillon et al. 3339 (F).

6. Cheilanthes fraseri Kuhn, Linnaea 36: 83. 1 869.

SYNTYPES: Ecuador, Fraser, Wagner, and

Peru, Ruiz & Pavon (Herb. Mett., B!, a sheet

with Pavon and Wagner in a packet; photos,

GH, us).

Not holaena fraseri (Kuhn) Baker, Syn. fil., ed. 2, 83.

1874.

Stem short- to moderately long-creeping, scales

narrowly linear-subulate, entire or remotely ser-

rulate, brownish, concolorous or somewhat bico-

lorous. Leaves to 40 cm long, petiole about half

as long as the lamina, lamina narrowly linear, pin-

nate-pinnatifid or 2-pinnatifid at the base. Pinnae

mostly deltoid, with the abaxial surface densely

tomentose with rather coarse, matted, pale brown

trichomes. Margin slightly modified.

Rocky places, often shaded, 800-3500 m, Piura

south to Cuzco.

Ecuador to Bolivia.

The differences between this species and the

sometimes similar Cheilanthes bonariensis are

presented in the key. Also C. fraseri has the petiole

and rachis more slender than those in C. bonar-

iensis and the petiole rather long, rather than

short, in relation to the length of the lamina.

Piura: Huancabamba, Scolnik 1435 (uc). Cajamarca:

Santa Cruz, Weberbauer 4139 (B). Between San Marcos

and Cajabamba, Correll & Smith P909 (GH, us). Huan-

uco: Huanuco, Macbride & Featherstone 2328 (F, GH,

us). Ambo, Ferreyra 6912 (GH, USM). Lima: Rimac-Tal,

Rauh & Hirsch 146 (USM). Huancavelica: Prov. Tayacaja,

entre Pampas y Salcabamba, Tovar 3848 (GH, USM). Ay-

acucho: Prov. La Mar, San Miquel-Llausa, Velarde 4951

(GH). Apurimac: Prov. Abancay, Quisapata, Vargas 89 10

(MO, uc). Cuzco: Rio Sambray, Mexia 8055 (F, GH, MO,

uc, us). Quillabamba, Stork et al. 10449 (F, GH, uc).

7. Cheilanthes bonariensis (Willd.) Proctor, Bull.

Inst. Jamaica, Sci. Ser. 5: 15. 1953.

Acrostichum bonariense Willd., Sp. pi. ed. 4, 5: 114.

1810. TYPE: "Bonaria" (holotype. Herb. Willd.

19537, B!; photo, GH).

Pteris aurea Poiret in Lam., Encycl. 5: 710. 1804.

TYPE: Peru, Joseph de Jussieu (holotype, Herb.

Jussieu 1333, P!; photos, GH, us), not Cheilanthes

aurea Baker, 1868.

Notholaena aurea (Poiret) Desv., Mem. Soc. Linn.

Paris 6: 219. 1827.

Notholaena bonariensis (Willd.) C. Chr., Index fil. 6.

1905; 459. 1906.

Stem short-creeping, knotted, scales lance-lin-

ear, entire, brownish, somewhat bicolorous. Leaves

to 60 cm long, petiole '/? as long as the lamina or

less, lamina linear-elliptic, long-attenuate at the

base, pinnate-pinnatifid. Pinnae oblong to deltoid-

oblong, with the abaxial surface covered with a

dense tawny (white when young) tomentum of fine,

matted trichomes. Indusium a very narrow sub-

hyaline band.

On soil banks, rocky slopes, shrubby hillsides

or cliffs, 1200-3800 m, Lambayeque and Ama-

zonas south to Puno and Arequipa.

Southwestern United States to Chile and Ar-

gentina; West Indies.

Cheilanthes bonariensis is one of the character-

istic ferns of the Altiplano. It is usually quite dis-

tinctive in the aspect of its pubescent lamina, but

some variations must carefully be distinguished

from C. fraseri. One collection (Cuzco, Ollantay-

tambo, Hitchcock 22541, GH), is evidently a hy-

brid. It has the slender petiole and rachis and the

long petiole of C. fraseri and the short stem and

dark scale tips of C. bonariensis.

Lambayeque: 44 km from Olmos on road to Jaen,

Correll & Smith P821 (GH). Cajamarca: Prov. Chota,

Llama, Lopez & Sagdstegui 5310 (GH, HUT, MO). Ama-

zonas: Prov. Chachapoyas, Chachapoyas-Molinopampa,

Sagdstegui 6033 (GH). La Libertad: Huaranchal, Sagds-

tegui 211 (GH). Ancash: Chiquian, Ferreyra 5786 (USM).

28

FIELDIANA: BOTANY

Huanuco: Between Ambo and Huanuco, Ferreyra 9234,

10394 (GH, USM). Lima: Oroya RR., Safford 991 (F, GH,

p, us). Junin: Vilcabamba, Tryon & Tryon 5421 (BM, F,

MO, u). Huancavelica: 4 km S of Conaica, Tovar 970

(GH, USM). Ayacucho: Pampalca, Killip & Smith 22256

(F. us). Apurimac: Prov. Andahuaylas, 2 km N of An-

dahuaylas. Stork <$ Horton 10716 (us). Cuzco: Calca,

Vargas 3136 (F). Arequipa: Prov. Arequipa, near Las-

pinas, Eyerdam & Beetle 22144 (GH, us). Puno: Sandia,

Weberbauer 721 (B).

On lomas, 90-100 m, Arequipa.

Peru and Chile.

Cheilanthes mollis is amply distinguished by its

indument of stellate trichomes.

Arequipa: Ocona, Ellenberg4153 (GH). Prov. Caraveli,

Lomas de Atico, Hutchison & Wright 7 130 (GH). Mol-

lendo, Weberbauer 1545 (B).

8. Cheilanthes obducta Kuhn, Linnaea 36: 83.

1869. TYPE: Bolivia, D'Orbigny 386 (holo-

type, B!; photo, GH).

Notholaena obducta (Kuhn) Baker, Syn. fil., ed. 2, 5 1 5.

1874.

Stem short-creeping, scales narrowly linear, pale

brown, or with a castaneous central portion and

lighter margins. Leaves to 55 cm long, petiole much

shorter than the lamina, terete, slightly to densely

pubescent, the rachis similar, lamina lanceolate,

2-pinnate. Pinnae elongate, with the abaxial sur-

face densely subappressed pubescent, the tri-

chomes rather straight. Margin unmodified.

Under shrubs and trees, 600 m, Cajamarca.

Venezuela, Colombia, south to Bolivia and Par-

aguay.

The trichomes on the abaxial surface of the pin-

nae are unusual, many of them having short, few-

celled branches. Also, among the densely pubes-

cent species, this is the only one with clearly su-

bappressed pubescence.

Cajamarca: Valley of the Rio Chamaya, Mesones to

Muro, Hutchison 1425 (uc, us).

9. Cheilanthes mollis (Kunze) Presl, Tent, pterid.

160. 1836.

10. Cheilanthes orbignyana Kuhn, Linnaea 36: 82.

1869. TYPE: Bolivia, Prov. La Laguna, D'Or-

bigny 388 (Herb. Mett., B!, is doubtless from

the original material; isotype, P!; photos, GH,

us).

Stem erect, moderately stout, scales lanceolate

or narrowly so, brown to dark brown, concolorous,

semisclerotic. Leaves ca. 25-50 cm long, petiole

shallowly grooved on the adaxial side, at least to-

ward the apex, flat to convex between the small,

rounded ridges that are concolorous with the pet-

iole proper, glabrous or nearly so, rachis similar

but the ridges somewhat lighter in color and rather

sharp, lamina narrowly ovate, 3-pinnate-pinnati-

fid to 4-pinnate. Pinnae glabrous to slightly glan-

dular. Indusium suborbicular to lunate, well mod-

ified.

Among rocks, 2750-2860 m, Cajamarca and

Lima.

Peru and Bolivia.

Evidently this is a rare species but it should be

found in the mountains south of Lima.

Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui

2937 (F, GH, us), 1 1435 (F, MO). Prov. Contumaza, La

Herilla (Guzmango), Sagdstegui et al. 97/0(MO,uc). Prov.

Cajamarca, Las Quinas-Huatum, Sagdstegui etal. 10132

(HUT, MO, uc). Lima: Bosque de Zarate, San Bartolome,

Valencia & Franke 666 (F).

Notholaena mollis Kunze, Linnaea 9: 54. 1834. TYPE:

Chile, (Valparaiso), near Valparaiso, Playa An-

cha, Poeppig (holotype, LZ destroyed).

Stem short-creeping, knotted, scales narrow-lin-

ear, entire, concolorous or bicolorous. Leaves to

30 cm long, petiole usually shorter than the lam-

ina, terete, slightly to densely pubescent, the rachis

similar, lamina lanceolate, to 3-pinnate. Pinnae

with the abaxial surface densely covered with

whitish to ferrugineous, stalked, stellate tri-

chomes, ultimate segments small, many suborbi-

cular. Margin unmodified, strongly revolute.

11. Cheilanthes rufopunctata Rosenst., Meded.

Rijks-Herb. 19: 9. 1913. TYPE: Bolivia, (La

Paz), Araca, Herzog 2366 (holotype, not lo-

cated; frag, ex Rosenst., us!; isotype, L; photo,

GH).

Stem creeping, often knotted, scales lanceolate,

long-triangular, reddish brown, wholly sclerotic.

Leaves ca. 15-35 cm long, petiole grooved on the

adaxial side, at least toward the apex, deeply sul-

cate between the relatively large, rounded ridges

which are concolorous with the petiole, glabrous,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

29

rachis grooved and glabrous, with green wings to-

ward the apex, lamina long-deltoid to broadly lan-

ceolate, 3-pinnate to 3-pinnate-pinnatifid. Pinnae

glabrous on both sides, except for usually numer-

ous reddish, sessile glands, especially abaxially.

Indusium broad, crispate, well modified.

Rock crevices, 2600-4000 m, Lima to Puno.

Peru and Bolivia.

Cheilanthes glauca (Cav.) Mett. of Chile seems

closely related to this species. It has a pentagonal

lamina, and the segments are thickly beset beneath

with short, dark trichomes.

Lima: Prov. Canta, arriba de San Jose, Miiller et al.

12474 (GH). Matucana, Macbride & Featherstone 285 (F,

GH, us). Junin: Huancayo, Kunkel 482lh (GH). I luan-

cavelica: Near Conaica, Tovar 316 (GH, us, USM). Cuzco:

Valle de Lares, Soukup 29 (F, us). Calca, Vargas 4004

(cuz, uc, us). Puno: Near Puno, Vargas 29 (MO), Soukup

29 (uc).

12. Cheilanthes marginata H.B.K., Nov. gen. sp.

1: 22. 1815. TYPE: Ecuador, (Pichincha),

Quito, Penipe, Humboldt & Bonpland (ho-

lotype, P!; photo, GH).

Stem short-creeping, somewhat multicipital,

scales ovate-lanceolate, attenuate, dark brown,

sclerotic. Leaves ca. 8-25 cm long, the petiole sul-

cate, the ridges rounded, concolorous with the pet-

iole, somewhat scaly, rachis sulcate, with green

wings on the adaxial side above the base, lamina

broadly deltoid to lanceolate, mostly 3-pinnate.

Pinnae glabrous, ultimate segments stalked or nar-

rowed at the base. Indusium glandular-fimbriate,

continuous along the margins and decurrent onto

the axes of the pinnules and pinnae.

On cliffs, among rocks, and in rocky soil, 2200-

4000 m, Piura to Puno.

Venezuela and Colombia, south to Argentina.

The unusual condition of the indusium, which

extends from the segments along their stalks and

along the pinna-rachis, mark this as one of the

most distinctive species of the genus in Peru.

Piura: Prov. Ayabaca, Ayabaca, Sagdstegui et al. 7804

(MO). Cajamarca: Cajamarca to Chilete, Ferreyra 3294

(USM). El Puquio, Guzmango, Sagdstegui 3918 (GH). La

Libertad: Huaranchal, Sagdstegui 207, 2664 (GH). An-

cash: E of Huasta. Cerrate 2264 (USM). Huanuco: Mito,

Macbride 3226 (F, GH, us). Pasco: Between Salcachupan

and Cerro de Pasco, Ferreyra 6619 (GH, USM). Junin:

Huancayo, Kunkel 48 1 (GH). Huancavelica: Colcabamba

to Paurcarbamba, Tovar 1969 (USM). Ayacucho: Pam-

palca, Killip & Smith 22234 (us). Cuzco: Machu Picchu,

Tryon & Tryon 5401 (BM, F, MO, u, us, USM). Puno:

Sandia, Soukup 157 (F), Weberbauer 718 (B).

13. Cheilanthes poeppigiana Kuhn, Linnaea 36:

84. 1869. SYNTYPES: Peru, Poeppig, and

Bolivia, Cuming. (Cumingin Herb. Mett., B!,

is determined by Mettenius as Cheilanthes

poeppigiana and is doubtless part of the orig-

inal materials.) Figure 6a-b.

Stem small, short, scales subulate-lanceolate with

a dark brown sclerotic central portion and paler,

thinner borders. Leaves ca. 8-35 cm long, petiole

grooved on the adaxial side, the ridges rounded,

concolorous with the petiole proper, slightly scaly

to glabrate, rachis similar, but with narrow green

wings on the adaxial side, at least toward the apex,

lamina oblong-lanceolate to deltoid, pinnate-pin-

natifid to 2-pinnate-pinnatifid. Pinnae glabrous,

pinnules adnate to somewhat narrowed at the base.

Indusium continuous, intramarginal, well modi-

fied.

Crevices of cliffs or of Inca walls, among rocks

in ravines and on shrubby slopes, 150-3000 m

(usually at 1 700-2200 m), Tumbes to Puno.

Ecuador to Argentina.

Small plants of this species may have a lanceo-

late lamina; large ones have an ovate to broadly

triangular one. The ultimate segments are rela-

tively broad and herbaceous with a broadly at-

tached base.

Tumbes: Between Tumbes and Cancho, Coronado 226

(GH, uc). Prov. Zarumilla, El Cancho, J. Schunke V. 3031

(F, GH, us). Lambayeque: Entre Beatita de Humay y km

38 (Carretera Olmos-Maranon), Lopez et al. 4042 (GH).

Cajamarca: Entre Cascas y Contumaza, Lopez el al. 3725

(F, GH). La Libertad: Huaranchal, Sagdstegui 194 (GH).

Huanuco: Piedra Grande, Macbride 3672 (F). Junin: Hu-

acapistana, Tryon & Tryon 5436 (BM, F, MO, u, us, USM).

Cuzco: Machu Picchu, Tryon & Tryon 5396 (BM, F, MO,

u, us, USM). Puno: Sandia, Weberbauer 566 (B).

14. Cheilanthes farinosa (Forssk.) Kaulf., Enum.

fil. 212. 1824.

Pteris farinosa Forssk., Fl. aegypt.-arab. 187. 1775.

TYPE: Forsskal (holotype, not located; isotype,

BM!).

Aleuritopteris peruviana Saiki, J. Phytogeog. Tax. 32:

89. 1984. TYPE: Peru, (Huanuco), Muna, Bryan

555 (holotype, F!; isotype, us!).

30

FIELDIANA: BOTANY

Stem rather stout, very short-creeping, decum-

bent to nearly erect, scales narrowly long-trian-

gular, brown, semisclerotic, concolorous. Leaves

ca. 10-70 cm long, petiole terete to grooved on

the adaxial side, the ridges rounded, concolorous

with the petiole proper, slightly scaly or not, gla-

brous to farinaceous, rachis grooved on the adaxial

side, glabrous to farinaceous, lamina long-trian-

gular to ovate-triangular or narrowly so, pinnate-

pinnatifid to 2-pinnate-pinnatifid. Pinnae gla-

brous to somewhat farinose on the adaxial surface,

conspicuously to densely white-farinose abaxially.

Indusium crispate, lobed, more or less continuous,

well modified.

Rocky places, ca. 2000 m, Huanuco, Junin, and

Cuzco.

West Indies; Mexico to Peru; Old World.

This is a most distinctive species because of the

white-farinose covering of the segments which is

especially evident on their under surface.

Saiki (op. cit.) recognized segregate species from

Cheilanthes farinosa sens. lat. in the genus Aleu-

ritopteris, along with some species of Notholaena.

The scattered distribution of C. farinosa in Amer-

ica suggests that it was introduced from cultivated

plants. It probably does represent a complex but

this needs more careful study, especially of cyto-

logical and chemical characters.

Junin: near Huacapistana, Tryon & Tryon 5435 (F).

Cuzco: Nevada Sallcantay, Bites 986 (us).

15. Cheilanthes concolor (Langsd. & Fisch.) R. &

A. Tryon, Rhodora 83: 133. 1981. var. con-

color.

Pteris concolor Langsd. & Fisch., Icon. fil. 19, /. 21.

1810. TYPE: Archipelago Marquesas, Nucahiva

Island (Nukuhiva or Nuku Hiva), Langsdorff(ho-

lotype, LE?; isotype, Herb. Willd. 19961-1, u!;

photo, GH).

Doryopteris concolor (Langsd. & Fisch.) Kuhn in

Decken, Reisen Ost-Afrika 3(3): 19. 1879.

Stem rather short and stout, scales usually ovate-

lanceolate, brown, strongly bicolorous. Leaves ca.

5-35 cm long, petiole usually sulcate or with lateral

wings on the adaxial side, especially toward the

apex, glabrous or very slightly pubescent, lamina

pedate, pentagonal to suborbicular, usually 2-3-

pinnatifid. Pinnae essentially glabrous on both

sides. Indusium well modified, rather continuous

along the fertile margin.

Rocky places and clay banks, 750-2000 m, Ca-

jamarca, San Martin, and Cuzco.

Tropical America; Old World.

The pedate lamina is a distinctive character of

this species among the Peruvian ones. All Amer-

ican plants belong to var. concolor. Variety kirkii

is in Africa, India, and Ceylon.

Cajamarca: Jaen, Feb. 17, 1954, R. Cerron (GH, uc,

us). San Martin: Tarapoto, Li Williams 5451 (F, GH),

Spruce 4045 (GH, K). Cuzco: Valley of the Vilcanota,

Mexia 8053 (GH, MO, uc, us). Potrero, Tryon & Tryon

5375 (BM, F, GH, MO, u, uc, us, USM).

16. Cheilanthes myriophylla Desv., Ges. Naturf.

Freunde Berlin Mag. Neuesten Entdeck. Ges-

ammten Naturk. 5: 328. 1811. TYPE: Peru

(holotype, Herb. Desvaux, P!; photos, GH, us).

Figure 6c.

Cheilanthes elegans Desv. Ges. Naturf. Freunde Ber-

lin Mag. Neuesten Entdeck. Gesammten Naturk.

5: 328. 1811. TYPE: "Chile" (holotype, Herb.

Desvaux, P!; photo, GH; a portion of a pinna,

probably from Peru: J. Jussieu, Herb. Jussieu

1438, P!; photos, GH, us).

Stem moderately stout, short-creeping, multi-

cipital, scales lance- subulate, with a dark, sclerotic

central portion and paler margins. Leaves ca. 1 5-

40 cm long, petiole terete, scaly, as is the rachis,

lamina lanceolate to lanceolate-oblong, or narrow-

ly so, to 4-pinnate. Pinnae densely scaly abaxially,

nearly glabrous adaxially, the ultimate segments

small, suborbicular. Indusium more or less mod-

ified, rather continuous.

Rocky soil, shrubby hillsides, and cliffs, 1550-

3250 m, Piura to Puno and Arequipa.

Hispaniola; Mexico to Colombia and Venezue-

la, south to Chile; Brazil.

This species is characterized by its small, bead-

like ultimate segments that are readily visible on

the adaxial side of the lamina and usually not com-

pletely concealed by the scales on the abaxial side.

The narrow scales on the petiole and rachis su-

perficially resemble matted trichomes. It is one of

the most common and widely distributed of the

ferns on the Altiplano.

Piura: Huancabamba, Scolnik 1436 (uc, us). Caja-

marca: Prov. Contumaza, Jalca de Nanra, Sagdstegui

3891 (GH). Amazonas: Prov. Chachapoyas, Rio Sonche,

near Sonche, Wurdack 1544 (F, GH, uc, us). La Libert ad:

Trujillo to Otusco, Sounders 896 (F, GH). Ancash: Chi-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

31

quian, Cerrate 546 (USM). Huanuco: Acomayo, Tryon &

Tryon 5328 (BM, F, MO, u, us, USM). Lima: Chancay, E

of Sayan, Goodspeed 33032 (GH, MO, uc, us). Pasco:

Solomon 3036 (F, MO). Junin: Vilcambamba, Tryon &

Tryon 5420 (BM, F, GH, MO, u, us, USM). Huancavelica:

N of Pampas, Stork & Horton 10242 (F, GH, MO, uc).

Ayacucho: Ayacucho, West 3641 (MO, uc). A pur i mac:

Challhuanca, Sounders 772 (GH). Cuzco: Anta, Tryon &

Tryon 5363 (BM, F, MO, u, us, USM). Arequipa: 1 4 km N

of Arequipa, Eyerdam & Beetle 22164 (GH, MO, uc, us).

Puno: Sandia, Weberbauer 724 (B).

17. Cheilanthes cantangensis (Tryon) Tryon,

comb. nov.

Notholaena cantangensis Tryon, Rhodora 63:81. 1 96 1 .

TYPE: Peru, Dept. Cajamarca, Prov. Celendin,

Cantange, Lopez & Sagdstegui 3366 (holotype,

GH!).

Stem short, scales lanceolate, with a dark, scle-

rotic central portion and brown, pectinate-serru-

late margins. Leaves 10-15 cm long, petiole terete

or subterete, somewhat scaly, the rachis similar,

lamina lanceolate-elliptic, 2-pinnate-pinnatifid.

Pinnae with the abaxial surface covered with nar-

rowly ovate-lanceolate scales, the adaxial surface

moderately pubescent with rather short, thick, and

rigid trichomes. Margin slightly modified.

Among rocks, 1450 m, Cajamarca.

Endemic to Peru.

This rare species is characterized by the short,

rigid trichomes on the adaxial surface of the seg-

ments. It is known only from the type collection.

reddish brown. Leaves ca. 8-15 cm long, petiole

densely scaly, terete, as is the rachis. Lamina linear

to rather narrowly elliptic, 3-pinnate. Pinnae

densely scaly abaxially, the scales and especially

their tips curving over the glabrous adaxial surface

and concealing it, ultimate segments small, su-

borbicular or nearly so. Margin strongly recurved,

only slightly, if at all, modified.

Exposed calcareous rocky places and cliffs, 2700-

4300 m, La Libertad to Puno.

Peru and Bolivia.

This species is unusual in that the adaxial sur-

face of the ultimate segments is nearly concealed

by the scales on the abaxial surface curving to the

upper surface. Some of the scales on the rachis

have such strongly curved auricles that they give

the appearance of a small scale attached at the

base.

This and the next four species form a distinctive

group, along with Cheilanthes squamosa of Bolivia

and Argentina. They all have rather similar scaly

leaves and prominently verrucate spores.

La Libertad: Yanazara to Huaquil, Lopez & Sagds-

tegui 3395 (GH). Ancash: Mahuay, Cerrate 2200 (USM).

Laguna de Palagatos, Lopez 2388 (HUT). Lima: Matu-

cana, Macbride & Featherstone 424 (F, us), Goodspeed

11347 (GH, uc). Pasco: 95 km S of Huanuco, road to

Cerro de Pasco, Gentry et al. 37495 (F, MO). Junin: Near

Tarma, Tryon & Tryon 5457 (BM, F, GH, MO, u, us, USM).

Huancavelica: SE of Conaica, Tovar 113 (GH, MO, us).

Apurimac: Prov. Andahuaylas, Vargas 8792 ( cuz, uc).

Cuzco: Pisaj, Vargas 23239 (GH). Puno: Pucara, Weber-

bauer 425 (B).

18. Cheilanthes scariosa (Sw.) Presl, Reliq. haenk.

1: 65. 1825.

Acrostichum lanuginosum Willd., Schrift. Acad. Er-

furt, 1802: 31, /. 3,f. 4, not Desf. 1800. TYPE:

Peru, Malaspina Exped. (holotype. Herb. Willd.

19554-1, B!; photos, GH, us).

Acrostichum scariosum Sw., Syn. fil. 16. 1806, nom.

nov. for A. lanuginosum Willd. (not Desf.) and

with the same type.

Cheilanthes ornatissima Maxon, Smithsonian Misc.

Coll. 65(8): 3. 1915. TYPE: Peru, (Lima), moun-

tains back of Lima, Safford 996 (holotype, us!;

isotypes, F!, GH!, MO!, uc!, us!).

Cheilanthes tripinnata Copel., Univ. Calif. Publ. Bot.

19: 301, /. 58. 1941. TYPE: Peru, Dept. Apuri-

mac, Chincheros to Andahuaylas, West 3724 (ho-

lotype, uc!).

Stem rather stout, very short-creeping, multi-

cipital, scales long and filiform, concolorous, light

19. Cheilanthes peruviana (Desv.) Moore, Index

fil. 250. 1861.

Notholaena peruviana Desv., Mem. Soc. Linn. Paris

6: 220. 1827. TYPE: Peru, Dombey (holotype, P!;

photos, GH, uc, us; frag., B!).

Notholaena sinuata var. bipinnata Hooker, Sp. fil. 5:

108. 1864. TYPE: Peru, (Huanuco), "Andes of

Peru", Banos, Brackenridge (Wilkes, Explor.

Exped.) (holotype, K!; photos, GH, us).

Notholaena brackenridgei Baker, Syn. fil. 371. 1868.

TYPE: the same as for N. sinuata var. bipinnata.

Stem erect, stout, scales linear-ligulate, long-at-

tenuate, margins entire or nearly so, brown, con-

colorous. Leaves to 30 cm long, the petiole terete,

scaly, the rachis similar, lamina narrowly lanceo-

late, not narrowed below, 2-3-pinnate, with some

pinnules lobed. Pinnae with the abaxial surface

densely covered with dark brown, lanceolate to

32

FIELDIANA: BOTANY

ovate, somewhat pectinate-serrulate scales. Mar-

gin modified, narrow.

In soil and on rocks of lomas, and on exposed

rocky slopes, 300-3900 m, Lambayeque to Mo-

quegua.

Endemic to Peru.

This and the next species are closely related and

may be separated by the characters mentioned in

the key. The two perhaps are not distinct species.

Lambayeque: 44 km from Olmos on road to Jaen,

Correll & Smith P822 (GH, us). Cajamarca: El Puquio,

Guzmango, Sagdstegui 3913 (GH). La Libertad: Prov.

Otuzco, Huaranchal, Lopez et al. 2665 (GH). Ancash:

Prov. Huaraz, 10 km from Cachabamba, Smith & Bud-

densiek 10896 (F). Lima: Above San Bartolome, Ferreyra

9742 (GH, USM). Arequipa: Lomas de Capac, Coronado

39 (GH, uc, us). Lomas de Chaparra, Ferreyra 11968

(GH, USM). Moquegua: Ilo, D. Stafford 933 (BM).

20. Cheilanthes arequipensis (Maxon) R. & A.

Tryon, Rhodora 83: 133. 1981.

Notholaena arequipensis Maxon, Smithsonian Misc.

Coll. 65(8): 9. 1915. TYPE: Peru, (Arequipa), near

Arequipa, Rose & Rose 18797 (holotype, us!;

photo, GH).

Stem erect or ascending, scales linear, long-at-

tenuate, distantly denticulate with ascending teeth,

reddish brown, concolorous. Leaves to 8 cm long,

petiole terete, scaly, as is the rachis, lamina del-

toid-oblong, 2-pinnate, the pinnules hardly if at

all lobed. Pinnae with the abaxial surface densely

covered with large, ovate-oblong, long-acuminate,

light reddish brown, erose-denticulate scales.

Margin somewhat revolute, unmodified.

Rocky places and crevices of rocks, 2200-3300

m, Lima and Arequipa.

Central Peru to northern Chile and northwest-

ern Argentina.

This species is close to Cheilanthes peruviana

and it is sometimes difficult to distinguish the two.

The small leaves with strongly clathrate and

roughened scales abaxially and the reddish stem

scales are especially distinctive features.

Lima: Prov. Huarochiri, Dist. San Mateo, Sounders

621 (F). Arequipa: Arequipa, R. S. Williams 2638 (GH,

NY, us), Pennell 13198 (B, F, GH, us). Yura, Vargas 8019

(uc). Quebrada de San Lazaro, Munz 15500 (GH). El

Misti, D. Stafford 570 (BM).

21. Cheilanthes incarum Maxon, Smithsonian

Misc. Coll. 65(8): 5.1915. TYPE: Peru, (Cuz-

co), Cuzco, Rose& Rose 19061 (holotype, us!).

Stem rather stout, short-creeping, multicipital,

scales reddish, subulate, attenuate to a capillary

tip. Leaves 10-25 cm long, the petiole densely

scaly, terete, as is the rachis, lamina 2-3-pinnate,

linear-lanceolate. Pinnae densely covered with

scales abaxially, these not clathrate, with a few

readily deciduous ones adaxially, the apical ulti-

mate segments mostly ovate-deltoid, or oblong,

the basal ones of secondary segments nearly su-

borbicular, their adaxial surface only partially con-

cealed by the tips of the scales from the abaxial

surface. Indusium rather narrow, more or less con-

tinuous.

Exposed rocky places, 2600-4100 m, Huancav-

elica to Puno.

Peru and northwestern Argentina.

In general appearance this species is similar to

Cheilanthes scariosa, but may be separated by the

characters mentioned in the key.

Huancavelica: Mejorada, Tovar 1014 (GH, USM). Apu-

rimac: 5 km N of Huacapistana, West 3903 (uc). Prov.

Ayacha, 70 km N of Mejorada, Hutchison 1690 (GH).

Cuzco: Saxihuaman, Tryon & Tryon 5343 (BM, F, GH,

MO, u, us, USM). Puno: Capachia Peninsula (Lake Titi-

caca), Tut in 1032 (BM).

22. Cheilanthes lonchophylla (Tryon) R. & A.

Tryon, Rhodora 83: 133. 1981.

Notholaena lonchophylla Tryon, Contr. Gray Herb.

179: 19. 1956. TYPE: Peru, Mat hews 610 (ho-

lotype, K!; photo, GH).

Stem short, horizontal or ascending, scales lin-

ear-ligulate, sparingly antrorsely denticulate,

brownish, concolorous. Leaves to 30 cm long, pet-

iole terete, scaly, the rachis similar, lamina lan-

ceolate or narrowly so, pinnate-pinnatifid to 2-

pinnate-lobed. Pinnae with the abaxial surface

densely covered with pale brown to whitish, lin-

ear-subulate, long-acuminate, pectinate-dentate

scales. Margin unmodified.

Rocky places, 2850-2950 m, Cajamarca.

Endemic to Peru.

This rare species is characterized by narrow scales

with relatively long marginal cilia on the pinnae

beneath.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

33

Cajamarca: Canyon of the Rio Maranon, above Bal-

sas, Hutchison & Wright 5297 (uc, frag. GH). Prov. Ce-

lendin, 20 km de Bagua, Aguado 6791 (GH, HUT), Mos-

tacero el al. 881 (HUT). Prov. Contumaza, Lopez &

Sagdstegui 7653 (HUT).

23. Cheilanthes sinuata (Sw.) Domin, Biblioth.

Hot. 85: 133. 1913, var. sinuata.

Acrostichum sinuatum Sw., Syn. fil. 14. 1806. TYPE:

Peru, "Squamaria sinuata" Lagasca Herb, (ho-

lotype, Swarlz Herb., s!).

Notholaena tectaria Desv., Mem. Soc. Linn. Paris 6:

219. 1827. TYPE: Peru (holotype, Desvaux Herb.,

P!; photos, GH, us; isotypes(?): Tarma, Dombey

"N. tectaria Desv." ex Desvaux, B!; Tarma, Dom-

bey, BM!).

Notholaena sinuata (Sw.)Kau\f.,Enum. fil. 135. 1824.

Stem short, horizontal, scales linear to linear-

subulate, pectinate-ciliate to entire, brownish,

concolorous. Leaves to 45 cm long, petiole terete,

scaly, the rachis similar, lamina linear, pinnate-

pinnatifid. Pinnae with the abaxial surface thickly

covered with castaneous or pale brown, deltoid to

lanceolate, acuminate, short-fimbriate scales, these

overlying a tomentum of smaller scales dissected

into long capillary processes. Margin unmodified.

Crevices of rocks, rocky banks and hillsides,

1000-3000 m, Piura to Amazonas and south to

Arequipa.

Southwestern United States south to Argentina;

Hispaniola.

This species is especially distinguished by the

two vascular bundles near the base of the petiole;

other American species of Cheilanthes have a sin-

gle bundle. The var. sinuata occurs throughout the

range of the species; other varieties occur in the

southwestern United States and Mexico.

Piura: Cerro Prieto, 20 miles E of Cape Parinas, Haught

187 (us). Lambayeque: 20 km from Olmos on road to

Jaen, Correll & Smith P795 (GH, us). Cajamarca: Prov.

Contumaza, entre Cascas y Contumaza, Lopez et al. 3687

(GH). Amazonas: Tambo de Carrizal, Stubel 1041, in part

(B). La Libertad: Huaranchal, Sagdstegui 210 (GH). An-

cash: S of Chiquian, Cerrate 7589 (USM). Huanuco: Mito,

Macbride & Featherstone 2310 (F, GH, us). Lima: Santa

Eulalia. Goodspeed 1 1307 (GH, MO, uc, us). Junin: Tar-

ma, Ferreyra 530 (GH, USM). Ayacucho: Nasca to Puquio,

Correll & Smith P171B (LL). Cuzco: Quillabamba, Stork

et al. 10449 (MO). Arequipa: Tiabaya, Pennell 13071 (F,

GH).

Comments

Cheilanthes buchtienii (Rosenst.) Tryon, comb,

nov. (Notholaena buchtienii Rosenst., Repert. Spec.

Nov. Regni Veg. 5: 238. 1908) occurs in Argentina

and Bolivia and may well be discovered in south-

ern Peru. It is similar to Cheilanthes fraseri but is

a more robust species, with larger median pinnae

that are lanceolate to deltoid-lanceolate, while in

C. fraseri the median pinnae are small and deltoid.

Cheilanthes hypoleuca (Kunze) Mett. of Chile

was considered to occur in Peru (as Notholaena

tomentosa Desv.) by Pichi-Sermolli (Webbia 8:

186-190. 1951) on the basis of two collections of

W. Lobb, both at BM!: "Andes of Peru, W. Lobb

and Huanico (Huanuco?) Peru, W. Lobb." There

are no modern collections of this species from Peru

and its presence requires confirmation. The species

may occur in southern Peru. It is similar to Chei-

lanthes fraseri but has a strongly sulcate petiole

and rachis, while C. fraseri has the petiole and

rachis terete.

Cheilanthes rigida (Sw.) Domin was correctly

indicated as from "Nova Hispania" (Mexico) by

Swartz (Syn. fil. 104. 1806, as Pteris rigida) and

erroneously from Peru by Swartz (Syn. fil. 299.

1806).

Cheilanthes squamosa Hooker & Grev. was re-

ported from the Capachica Peninsula, Lake Titi-

caca, Puno by Tryon ( 1 964). Further study of the

specimen (Tutin 1032, BM) indicates that it is Chei-

lanthes incarum. Cheilanthes squamosa of Bolivia

and Argentina is characterized by long, brown,

piliform scales on the adaxial surface of the seg-

ments.

VII. Adiantopsis

Adiantopsis Fee, (Mem. foug. 5) Gen. fil. 1 45. 1 852.

TYPE: Adiantopsis paupercula (Kunze) Fee

(Adiantum pauperculum Kunze). Figure 7.

Terrestrial. Stem erect or decumbent and short-

creeping, rarely slender and long-creeping, bearing

scales. Leaves monomorphic, ca. 10-80 cm long,

usually borne in a cluster, petiole with 1-3 vascular

bundles near the base, it and the lamina axes 2-

ridged adaxially. Lamina usually 1— 4-pinnate, or

ternate, radiate or pedate, glabrous or somewhat

short-pubescent abaxially, veins free. Sori sepa-

rate on the vein ends, 1 (rarely 2) covered by a

34

F1ELDIANA: BOTANY

FIG. 7. Adiantopsis radiata: a, habit, b, ultimate segment, abaxial side; c, portion of axis and bases of segments,

adaxial side, (a from Schmalz s.n., Brazil, F, b, c from Bues s.n., F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

35

lunate to roundish marginal indusium, paraphyses

absent. Spores tetrahedral-globose. trilete, echi-

nate.

Adiantopsis is a small tropical American genus

of about seven species, two of them in Peru. Dis-

tinctive features of the genus are the thin, brown

ridges on each side of the lamina axes on the adax-

ial surface, and the echinate spores. In Tryon ( 1 964)

these species were treated in the genus Chei-

lanthes.

References

TRYON, R. 1 964. The ferns of Peru. Contr. Gray

Herb., 194: 98-99.

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

topsis, pp. 266-270, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Species of Adiantopsis

a. Lamina pinnate

a. Lamina radiate or ternate

1. A. chlorophylla

. . 2. A. radiata

1 . Adiantopsis chlorophylla (Sw.) Fee, (Mem. foug.

5) Gen. fil. 145. 1852.

Cheilanthes chlorophvlla Sw., Kongl. Vetensk. Acad.

Handl. 1817: 76. TYPE: Brazil, (Minas Gerais),

Villa Rica, Freyreis (holotype, Herb. Swartz, s!;

photo, us).

Stem compact to creeping, scales narrowly lan-

ceolate to subulate, bicolorous, with a dark, scle-

rotic central portion and light, thin margins. Leaves

ca. 15-80 cm long, petiole with a light-colored

submembranous ridge on each side of the adaxial

surface, at least toward the apex, glabrate or gla-

brous, the rachis similar. Lamina elongate-trian-

gular, 2-3-pinnate-pinnatifid, pinnae glabrate to

glabrous. Indusium suborbicular to elongate, well

modified.

In shady woods, 850-2800 m, Cuzco.

Guatemala; Colombia south to Argentina, and

Brazil.

This species is evidently rare in the Andes from

Colombia to Peru, although rather common in

Argentina and Brazil. The pinnate lamina archi-

tecture readily separates it from A. radiata, the

other Andean species.

Cuzco: Prov. La Convencion, Rio Chaupimayo, Sou-

kup806 (F). Prov. La Convencion, Hacienda Sahuayaco,

Vargas 1659 (GH). Torontoy, Herrera 1332 (us).

2. Adiantopsis radiata (L.) Fee, (Mem. foug. 5)

Gen. fil. 145. 1852. Figure 7.

Adiantum radiatum L., Sp. pi. 1094. 1753. TYPE:

LINN 1252.1 (photo, A, is this species).

Cheilanthes radiata (L.) John Sm., J. Hot. (Hooker) 4:

159. 1841.

Adiantopsis ternata Prantl, Gartenfl. 32: 101. 1893.

TYPE: Venezuela, Orinoco-gebiet, Humboldt

(holotype, B!; photo, GH).

Stem erect to decumbent, often rather stout,

scales subulate, bicolorous, with a dark, sclerotic

central portion and lighter, thin margins. Leaves

ca. 10-50 cm long, petiole glabrous, usually terete,

to shallowly grooved toward the apex, axes of pri-

mary segments (pinnae) glabrous or nearly so, with

a light-colored, submembranous ridge on each side

on the adaxial surface. Lamina ternate, or usually

radiate with up to 7 (rarely to 9) pinnae, asym-

metrically circular, 2-pinnate, pinnae glabrous or

nearly so. Indusium suborbicular to oblong, well

modified.

In dense forests, on wooded slopes, or in rocky

places, 200-2800 m, Cajamarca and Amazonas,

south to Cuzco.

Tropical America.

The radiate arrangement of the usually five to

seven pinnae make this a highly distinctive species.

Rarely there are more pinnae or only three or four.

Cajamarca: Jaen, Feb. 17, 1954, Cerron (GH, uc).

Amazonas: Rio Utcubamba, 40 km S of Bagua Grande,

Hutchison 1469 (GH). Prov. Bagua, Pongo de Retema-

Aramango, Sagdstegui 5944 (HUT). San Martin: Juan

Jui, Alto Rio Huallaga, Klug 4170 (F, GH, MO, uc, us,

USM). Huanuco: Prov. Leoncio Prado, al este de Tingo

Maria, J. Schunke V. 10478 (F). Junin: La Merced, Mac-

bride 5375 (F, us). Huancavelica: SE of Tintay, Tovar

36

FIELDIANA: BOTANY

4705 (GH). Cuzco: Torontoy, Herrera 1299 (us). Prov.

Convention, Echarate, Vargas 1135 (GH).

VIII. Notholaena

NotholaenaR.Br.,Prodr. 145. 1 8 1 0. TYPE: Noth-

olaena trichomanoides (L.) Desv. (Pteris tri-

chomanoides L.), not Notholaena marantae

(L.) Desv. which is Paraceterach marantae

(L.) Tryon. Figure 8.

Notholaena sect. Argyrochosma John Sm., J. Bot.

(Hooker) 4: 50. 1841. TYPE (designated by C.

Chr., Index fil. xl. 1906): Notholaena nivea (Poir-

et) Desv. (Pteris nivea Poiret).

Argyrochosma (John Sm.) Windham, Amer. Fern J.

77: 38. 1987.

Terrestrial or rupestral. Stem small, decumbent

to suberect, short- to rarely rather long-creeping

and slender, bearing scales. Leaves monomorphic,

usually ca. 5—45 cm long, usually borne in a cluster,

petiole with 1 vascular bundle near the base, usu-

ally terete, rarely adaxially flattened, sulcate or

slightly ridged. Lamina usually 1-4-pinnate, or

rarely pedate and 2-pinnatifid or 2-pinnate-pin-

natifid, white- or yellow-farinose abaxially and

often with scales and (or) trichomes, or rarely gla-

brous, veins free. Sori usually on the somewhat

modified vein ends, sometimes of only 1 or 2 spo-

rangia, to rarely along most of the vein, the margin

flat to slightly recurved and not or only slightly

modified, paraphyses absent. Spores globose to

tetrahedral-globose, trilete, usually cristate or

somewhat rugose to plain and with a granulate

deposit.

Notholaena is an American genus of 39 mostly

Mexican species, with three in Peru. Species of

Cheilanthes have previously often been placed in

Notholaena as by Tryon (1956). In that treatment

only species 22-58 are to be retained in Notho-

laena; the others properly are species of Chei-

lanthes. There are two sections: Argyrochosma John

Sm. and Notholaena.

References

TRYON, R. 1956. A revision of the American

species of Notholaena. Contr. Gray Herb., 179:

1-106.

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

olaena, pp. 270-278, in Ferns and allied plants,

Springer- Verlag, New York.

WINDHAM, M. D. 1987. Argyrochosma, a new

genus of cheilanthoid ferns. Amer. Fern J., 77:

37^1.

Key to Species of Notholaena

a. Stem scales bicolorous, with a dark, sclerotic center and lighter margins; ultimate segments mostly

adnate (sect. Notholaena) 1 . N. sulphurea

a. Stem scales concolorous, brown; ultimate segments mostly stalked or sessile (sect. Argyrochosma)

b

b. Ultimate segments mostly orbicular or nearly so; stem scales oily 2. N. stuebeliana

b. Ultimate segments mostly ovate to oblong; stem scales dry 3. N. nivea

1. Notholaena sulphurea (Cav.) John Sm. in See-

mann, Bot. voy. Herald 1: 233. 1854. Figures

8b-c.

1864. TYPE: Peru, (Huanuco), Huanuco, Ma-

thews 981 (holotype, K.!).

Notholaena sulphurea var. JJava Kuhn, Abh. Naturf.

Ges. Halle 11: 32. 1869. TYPE: Peru (location

unknown).

Pteris sulphurea Cav., Descr. pi. 269. 1802. TYPE:

Mexico, (Hidalgo), Chimapan (Zimapan), Nee

(holotype, MA, seen by C. Chr., Dansk Bot. Ark.

9(3): 23. 1937; isotypes, F!, G!).

Cheilanthes borsigniana Koch, Wochen. Grtn. Plan-

zenkr. 1: 2. 1858. TYPE: Peru, Warszewicz (ho-

lotype, B!; photos, GH, us).

Notholaena Candida var. lutea Hooker, Sp. fil. 5: 111.

Stem short-creeping to nearly erect, scales lance-

ovate, attenuate, dark sclerotic with narrow

brownish and deciduously glandular-ciliate mar-

gins. Leaves to 20 cm tall, petiole usually much

longer than the lamina. Lamina pentagonal or

somewhat elongate, 2-pinnate or usually 2-pin-

nate-pinnatifid at the base, 1-pinnate-pinnatifid

above the basal pinnae, ultimate segments adnate,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

37

the abaxial surface densely yellow- to whitish yel-

low-farinose. Sporangia borne on the vein ends,

the margin thick, otherwise unmodified, some-

what recurved.

Open rocky places, 1500-2000 m, Piura and

Amazonas south to Arequipa.

Mexico south to Chile.

This is the only species of section Notholaena

that occurs in the Andes. In addition to the char-

acters mentioned in the key, the section is char-

acterized by a black, granulate deposit on the

spores.

Piura: Prov. Huancabamba, Cerro Colorado, Fosberg

27694 (uc), Fosberg 27709 (us). Cajamarca: Cantange,

Celendin to Rio Maranon, Lopez & Sagdstegui 3367 (GH,

us), Amazonas: Sunibamba, Utcubamba valley, Stiibel

1048, in part (B). Ancash: Caraz, Weberbauer 3007 (B).

Huannco: Piedra Grande, Macbride& Feather stone 20 51

(GH, us). Lima: Mts. back of Lima, Safford 993 (us).

Huancavelica: Valle de Montaro, entre Pampas y Sal-

cabamba, Tovar 3840 (GH, USM). Apurimac: 80 km from

Abancay, on road to Challhuanca, Sounders 769 (GH).

Arequipa: Arequipa, Rauh P567 (B).

2. Notholaena stuebeliana (Hieron.) Tryon, Rho-

dora63: 83. 1961.

Pellaea dealbata var. stuebeliana Hieron., Hedwigia

48: 225, /. 72, / 75. 1909. TYPE: Peru, Dept.

Amazonas, Sunibamba, Stubel 1048, in part (ho-

lotype, B!).

Argyrochosma stuebeliana (Hieron.) Windham, Amer.

Fern J. 77: 41. 1987.

Stem short, erect or decumbent, scales linear,

brown, oily, concolorous, margins entire. Leaves

5-30 cm long, petiole as long as or longer than the

lamina. Lamina deltoid to long-triangular, 3^4-

pinnate, the abaxial surface densely white- to very

pale yellow-farinose, ultimate segments mostly or-

bicular or nearly so. Sporangia borne on the ter-

minal 'A or more of the vein, margin unmodified,

nearly flat or recurved.

Among boulders and on ledges on shrubby

slopes, moist ravines and in rocky soil, 1400-3500

m, Cajamarca and Amazonas south to Junin.

Endemic to Peru.

This species was first known from a single leaf

collected in the latter half of the 19th century.

Recently it has been collected at several localities

and its range in Peru will probably be further ex-

tended.

Cajamarca: 40 km from Cajamarca on road to Chilete,

Correll & Smith P841 (GH, LL, us). Prov. Celendin, can-

yon of Rio Maranon, above Balsas, Hutchison & Wright

5288 (GH, us). Prov. Celendin, Celendin, Lopez & Sa-

gdstegui 3104 (GH, HUT). Amazonas: Prov. Bongara, S

from Pte. Ingenio to Chachapoyas, Plowman 5551 (GH).

La Libertad: Retamas, Lopez & Sagdstegui 3606 (GH).

Huanuco: Mito, Bryan 192 (F). Junin: Prov. Jauli, Saa-

vecha 6318 (GH).

3. Notholaena nivea (Poiret) Desv., J. Bot. Appl.

1:93. 1813. Figure 8a.

Pellaea nivea (Poiret) Prantl, Bot. Jahrb. Syst. 3: 417.

1882.

Argyrochosma nivea (Poiret) Windham, Amer. Fern

J. 77:41. 1987.

Stem short, erect or ascending, scales linear-sub-

ulate, dry, concolorous, margins entire. Leaves 1 0-

30 cm tall, petiole shorter than the lamina or about

the same length. Lamina lanceolate, deltoid-lan-

ceolate or ovate, 2-3-pinnate, the ultimate seg-

ments stalked, the abaxial surface densely white-

or yellow-farinose or glabrous. Sporangia borne

on the terminal 'A-'/z (rarely more) of a vein, mar-

gin unmodified, flat to somewhat recurved.

Rocky hillsides and crevices of rocks, 500—4000

m, Cajamarca south to Puno and Arequipa.

Colombia south to Argentina and Brazil.

This is a complex South American species, con-

sisting of the following three varieties.

Key to Varieties

a. Ultimate segments white-farinose abaxially

a. Ultimate segments glabrous abaxially

a. Ultimate segments yellow-farinose abaxially

. 3a. var. nivea

3b. var. tenera

. 3c. var. flava

38

FIELDIANA: BOTANY

5 cm

5mm

FIG. 8. Notholaena nivea: a, habit. Notholaena sulphurea: b, leaf; c, central pinna, abaxial side, (a from Stork &

Morton 10718, r, b, c from Macbride 3707, F.)

3a. Notholaena nivea var. nivea

Pteris nivea Poiret in Lam., Encycl. 5:718.1 804. TYPE:

Peru, Joseph de Jussieu (holotype, Herb. Jussieu

no. 1047, P!; photos, GH, us).

Notholaena nivea var. oblongata Griseb., Abh. Kon-

igl. Ges. Wiss. Gottingen 24 (Symb. fl. argent.):

342. 1879. TYPE: Argentina, (Salta), Salta, Hi-

eronymus & Lorentz "142 and 161 " (one collec-

tion) (holotype, B!; photo, GH).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

39

La Libertad south to Puno and Arequipa.

Ecuador south to western Argentina.

The sometimes recognized var. oblongata is not

as distinctive as the other varieties and intergrades

with var. nivea.

La Libertad: Otuzco to Huamchirco, Ferreyra 2987

(USM). Ancash: Chiquian, Cerrate 240, 564, 2106 (USM).

Huanuco: Huanuco, Macbride 3207 (F). Lima: San Ma-

teo, Ferreyra 7028 (GH, USM). Oroya RR., Safford 990

(GH, NY, P, us). Junin: Vilcambamba, Tryon & Tryon

5422 (BM, F, MO, u, us, USM). Huancavelica: Huancav-

elica. Stork & Horton 10823 (F, uc). Ayacucho: Ayacu-

cho, West 3642 (uc). Apurimac: Andahuaylas, Stork &

Horton 10718 (F, uc, us). Cuzco: Saxihuaman, Tryon &

Tryon 5344 (BM, F, GH, MO, u, us, USM). Arequipa: Ar-

equipa, Pennell 13245 (F). Puno: Baja Isla, Lake Titicaca,

Mexia 7788 (F, GH, MO, uc, us).

3b. Notholaena nivea var. tenera (Hooker) Gri-

seb., Abh. Konigl. Ges. Wiss. Gottingen 24

(Symb. fl. argent.): 342. 1879.

Notholaena tenera Hooker, Bot. Mag. t. 3055 and text.

1831. TYPE: Argentina, spec. cult, ex Mendoza,

Gillies (holotype, K.!; photo, GH).

Pellaea tenera (Hooker) Prantl, Bot. Jahrb. Syst. 3:

417. 1882.

Pellaea peruviana Copel., Univ. Calif. Publ. Bot. 19:

302. 1941. TYPE: Peru, Apurimac, Abancay re-

gion, Pupils of V. Santander C. (holotype, uc!).

La Libertad south to Puno.

Peru south to Argentina.

La Libertad: Prov. Otuzco, Arriba de Casmiche, Lopez

et al. 3962 (GH). Prov. Otuzco, Jamobamba, Aguado

6743 (HUT). Ancash: Cerro de Huasta, Cerrate 2456 (GH,

USM). Huanuco: Between Huanucay and Ambo, Ferreyra

6566 (USM). Lima: Entre Matucana y San Mateo, Fer-

reyra 5306 (GH, USM). Junin: Palca, Correll & Smith P758

(GH). Huancavelica: Mejorada, Tovar 1000 (GH, USM).

Apurimac: S of Chalhuanca, I his et al. 554 (GH). Cuzco:

Cuzco, Tryon & Tryon 5353 (BM, F, MO, u, us, USM).

Puno: Tequena, Feb. 19, 1948, Aguilar (USM).

Cajamarca south to Cuzco.

Colombia south to Argentina and Brazil.

The type indicated for var.flava is the only col-

lection, among those cited by Hooker and at Kew,

that has yellow indument abaxially.

Cajamarca: Prov. Santa Cruz, Cumbil-Catache, Lopez

& Sagdstegui 5162 (GH). Huanuco: Piedra Grande, Mac-

bride & Featherstone 3706 (F, us). Huancavelica: Valle

de Mantaro, entre Pampas y Salcabamba, Tovar 3847

(GH, USM). Cuzco: Potrero, Tryon & Tryon 5378 (BM, F,

MO, u, us, USM).

IX. Pellaea

Pellaea Link, Fil. spec. 59. 1 84 1 , conserved name.

TYPE: Pellaea atropurpurea (L.) Link (Pteris

atropurpurea L.). Figure 9.

Terrestrial or rupestral. Stem short and decum-

bent or rather slender and short- to long-creeping,

bearing scales. Leaves monomorphic or nearly so,

5 cm to 1.25 m long, borne in a cluster or spaced,

petiole with 1 vascular bundle near the base, terete

to adaxially sulcate or ridged. Lamina 1-4-pin-

nate, usually glabrous, to rarely pubescent or with

a few scales abaxially, veins free or rarely anas-

tomosing. Sori at the vein ends or on their apical

portion, partly to wholly covered by a slightly to

well-modified, recurved marginal indusium, pa-

raphyses absent or rarely short, often farinaceous

glands among the sporangia. Spores globose or

tetrahedral-globose, trilete, usually somewhat ru-

gose or cristate, rarely tuberculate or verrucate.

Pellaea is a nearly pantropical, sometimes tem-

perate, genus of about 35 species. Twenty-two

species are American and there are three in Peru.

Four sections are recognized, two of them in

America with sect. Pellaea the only one repre-

sented in Peru. The other section in America, Or-

mopteris (John Sm.) R. & A. Tryon, is primarily

Brazilian.

3c. Notholaena nivea var. flava Hooker, Sp. fil. 5:

112. 1855. TYPE: Ecuador, (Loja), Loxa

(Loja), Seemann (holotype, K!).

Acrostichum tereticaulon Desv., Ges. Naturf. Freunde

Berlin Mag. Neuesten Entdeck. Gesammten Na-

turk. 5: 310. 1811. TYPE: probably Peru, Dom-

bey (holotype, P!; photos, GH, us).

Notholaena chrysophylla Klotzsch, Allg. Gartenzei-

tung 23: 265. 1855. TYPE: spec. cult, ex Peru,

Warszewicz (holotype, B!).

References

TRYON, A. F. 1957. A revision of the fern genus

Pellaea section Pellaea. Ann. Missouri Bot.

Gard., 44: 129-193.

TRYON, R. M., AND A. F. TRYON. 1 982. Pellaea,

pp. 284-293, in Ferns and allied plants, Spring-

er-Verlag, New York.

40

FIELDIANA: BOTANY

Key to Species of Pellaea

a. Petiole and rachis atropurpureous to black; the pinnae mostly ternately divided ... 1 . P. ternifolia

a. Petiole and rachis ruddy-brown to straw-colored; the pinnae mostly pinnate with more than 3 segments

b

b. Rachis straight; stem moderately stout, compact; scales of the stem, base of the petioles, and leaf

buds tan, concolorous 2. P. sagittata

b. Rachis flexuous; stem mostly slender, cordlike, creeping; scales of the stem, base of the petioles,

and leaf buds bicolorous, tan to dark brown, the central portion often lustrous, sclerotic

. 3. P. ovata

1. Pellaea ternifolia (Cav.) Link, Fil. spec. 59.

1 84 1 , var. ternifolia. Figure 9a.

Pteris ternifolia Cav., Descr. pi. 266. 1802. TYPE:

Peru, (Lima), "Valle de Guarimaya dos leguas de

Guamantanga", (Huamantanga), Nee (holotype,

MA, seen by C. Chr., Dansk Bot. Ark. 9(3): 22.

1937).

Pteris peruviana Poiret in Lam., Encycl. 5: 718. 1804.

TYPE: Peru, Joseph de Jussieu (holotype, Herb.

Jussieu no. 1334A, P!; photos, OH, us).

Conaica, Tovar 976 (GH, USM). Ayacucho: Prov. Lucanas,

Pampa Galeras, Tovar 67 16 (USM). Apurimac: 2 km N

of Andahuaylas, Stork & Morton 10720 (F, uc). Cuzco:

Saxihuaman, Tryon & Tryon 5358 (BM, F, MO, u, us,

USM). Arequipa: Tiabaya, Pennell 13075 (B, F, GH, NY,

us). Puno: Juliaca, R. S. Williams 2633 (GH, NY, us).

2. Pellaea sagittata (Cav.) Link, Fil. spec. 60. 1 84 1 ,

var. sagittata.

Stem moderately stout, elongate, decumbent,

multicipital, scales straight or falcate, bicolorous

with a slender sclerotic stripe narrower than the

borders. Leaves 4-50 cm long, stiff, erect, petiole

and rachis plane on the adaxial surface or sulcate,

atropurpureous to black. Lamina linear to narrow-

ly lanceolate, 1 -pinnate, the pinnae ternate or en-

tire, sessile or subsessile, without a pinna-rachis.

In crevices of igneous rock or on Inca walls, in

sun and semishade, 1 800-4600 m, Cajamarca and

Amazonas south to Puno and Arequipa.

Southwestern United States south to Central

America; Hispaniola; Venezuela and Colombia

south to Argentina; Hawaiian Islands.

Pellaea ternifolia var. wrightiana (Hooker) A.

F. Tryon (Pellaea wrightiana Hooker) occurs in

the southwestern United States and adjacent

northern Mexico. The rest of the extensive range

of the species is occupied by var. ternifolia. Pellaea

weddelliana of Bolivia and Argentina has mostly

simple pinnae and nearly concolorous scales. It

represents a variation off. ternifolia that may be

found in Peru.

Cajamarca: El Gavilan. Cajamarca, Ferreyra 3269

(USM). Amazonas: Carrizal, Stiibel 1025 (B). La Libertad:

Cachicadan, Stork & Morton 9997 (F, uc), Sagdstegui el

al. 11919 (HUT). Ancash: Chiquian, Cerrate 622 (USM),

786 (USM). Huanuco: Mito, Bryan 198 (F). Lima: Cerca

Tupe, Cerrate 1089 (USM). Junin: Huacapistana, Tryon

& Tryon 5424 (BM, F, MO, u, us, USM). Huancavelica:

Pteris sagittata Cav., Descr. pi. 267. 1802. TYPE:

Mexico, Cerro de Guadeloupe, Nee (holotype, MA,

seen by C. Chr., Dansk Bot. Ark. 9(3): 23. 1937).

Stem moderately stout, compact, decumbent,

multicipital, scales straight or nearly so, concol-

orous, tan to rust-colored, lanceolate-triangular,

usually cordate. Leaves 1 5-80 cm long, erect, stiff,

petiole and rachis convex or plane on the adaxial

side, puberulous, the rachis straight or somewhat

flexuous, straw-colored to ruddy-tan, or mottled,

scales at the base of the petiole and on the leaf

buds concolorous. Lamina 1-2-pinnate, rarely 3-

pinnate, the pinnae ascending at acute angles to

the rachis, pinnae entire or of 3-numerous seg-

ments, long-stalked.

On dry banks, in open sun or shade, among

rocks and on stone walls, 1 700-3000 m. La Lib-

ertad south to Cuzco.

Mexico to Guatemala and Colombia, south to

Bolivia.

In addition to var. sagittata, there is var. cordata

(Cav.) A. F. Tryon in the southwestern United

States south to southern Mexico (Pellaea cordifolia

(Sesse & Mocino) A. R. Sm.).

La Libertad: Prov. Otuzco, Motil-Chota, Lopez & Sa-

gdstegui 8035 (HUT). Ancash: Callejon de Huallas, 8 km

SSE of Yungay, Tryon & Tryon 6552 (GH). Lima: Ma-

tucana, Macbride & Featherstone 422 (F, GH, us). Pasco:

Entre Salcahupan y Cerro de Pasco, Ferreyra 6621 (GH,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

41

5 cm

FIG. 9. Pellaea ternifolia: a, habit. Pellaea ovata: b, portion of lamina, abaxial side; c, pinnule, abaxial side, (a

from Davis et al. 1650, F; b, c from Bryan 438, F.)

42

FIELDIANA: BOTANY

USM). Junin: Huacapistana, Tryon & Tryon 5431 (BM, F,

GH, MO, u, us, USM). Huancavelica: Conaica, Tovar 978

(USM). Apurimac: Prov. Abancay, upper Rio Marino,

Stork el al. 10648 (F, uc). Cuzco: Valle de Urubamba,

Herrera 1195 (F, us).

3. Pellaea ovata (Desv.) Weath., Contr. Gray Herb.

114: 34. 1936. Figure 9b-c.

Pteris ovata Desv., Mem. Soc. Linn. Paris 6: 301.

1827. TYPE: Peru (holotype, Herb. Desvaux, P!;

photos, GH, uc).

Stem slender, cordlike, creeping, dichotomously

branched, scales straight or somewhat falcate, bi-

colorous, the sclerotic central portion usually lus-

trous. Leaves 10 cm to 1.25 m long, straight or

frequently twining, subscandent, petiole and rach-

is convex or plane on the adaxial side, the rachis

strongly flexuous, straw-colored to ruddy brown,

scales at the base of the petiole and on the leaf

buds bicolorous. Lamina elongate-triangular, 2-4-

pinnate, usually 3-pinnate, the pinnae descending

from the rachis, with 5-many segments, long-

stalked.

In open sun, brushy grasslands, on shaded banks,

among rocks and on old stone walls, 1 200-2900

m, Cajamarca and Amazonas south to Puno.

Southern Texas to Costa Rica; Jamaica and His-

paniola; Venezuela and Colombia south to north-

ern Argentina.

The leaves of Pellaea ovata often scramble

among the branches of low shrubs and may be

over 1 m long.

Cajamarca: 52 km W of Cajamarca on road to Chilete,

Correll & Smith P839 (GH, us). Prov. Contumaza, El

Tunel, Sagastegui et al. 12642 (HUT). Amazonas: Prov.

Chachapoyas. Chachapoyas-Molinopampa, Sagastegui

6031 (GH). La Libertad: Retamas, Lopez & Sagastegui

3608 (GH). Huanuco: Entre Ambo y Huanuco, Ferreyra

9232 (GH, USM). Lima: Prov. Canta, Km. 82 Lima-Canta,

Sounders 808 (GH). Junin: Huacapistana, Tryon & Tryon

5423 (BM, F, GH, MO, u, us, USM). Huancavelica: La Me-

jorada, Kunkel 452 (GH). Apurimac: Andahuaylas, Stork

& Morton 10712 (F, uc, us). Cuzco: Yucay, Soukup 754

(F, GH). Puno: Sandia, Weberbauer 841 (B).

X. Doryopteris

Doryopteris John Sm., J. Bot. (Hooker) 4: 162.

1841. TYPE: Doryopteris palmata (Willd.)

John Sm. (Pteris palmata Willd.) = Doryop-

teris pedata var. palmata (Willd.) Hicken.

Figure 10.

Terrestrial or rupestral. Stem decumbent and

creeping to suberect, small to stout, bearing scales.

Leaves monomorphic to dimorphic and the fertile

more erect and with narrower segments than the

sterile, borne in a cluster or rather spaced, petiole

with 1 or 2 vascular bundles near the base. Lamina

of various architecture, usually pedate, glabrous,

veins free or anastomosing without included vein-

lets. Sori marginal, the sporangia on a nearly con-

tinuous vascular commissure and covered by a

well-modified indusium, paraphyses absent. Spores

tetrahedral-globose, trilete, the surface nearly

smooth to prominently cristate.

Doryopteris is primarily an American genus of

25 species, with four of them in Peru. The genus

centers in southeastern Brazil and only a few species

extend beyond that region. The species Doryop-

teris concolor is now placed in Cheilanthes.

Reference

TRYON, R. M. 1942. A revision of the genus

Doryopteris. Contr. Gray Herb., 143: 1-80.

Key to Species of Doryopteris

a. Venation free, single areolae present by rare exception; petiole with 2 vascular bundles, at least toward

the base b

b. Soral lines discontinuous between the pinna-segments and most other segments of the lamina

1 . D. crenulans

b. Soral lines continuous between the pinna-segments and most other segments of the lamina ....

2. D. lomariacea

a. Venation completely areolate, or areolae only along the midveins of the segments; petiole with 1

vascular bundle . . . c

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

43

c. Fertile and sterile lamina with partly areolate venation, the areolae mostly along the costae or

midveins; proliferous buds absent at the base of the lamina 3. D. lorentzii

c. Fertile and sterile lamina with completely areolate venation; proliferous buds commonly present

at the base of the lamina 4. D. pedata var. palmata

1 . Doryopteris crenulans (Fee) Christ in Schwacke,

PI. nov. mineiras. 2: 26. 1900.

Pellaea crenulans Fee, Crypt, vase. Bresil 2: 27, /. 87,

f. I. 1872-1873. TYPE: Brazil, (Guanabara), Ti-

juca. Glaziou 5345 (5343 is excluded by Tryon,

1942) (holotype, p; photo, GH; isotypes, GH!, K!).

Stem scales long and narrow. Petiole usually

densely and minutely verrucose, rarely smooth,

with 2 vascular bundles near the base. Fertile and

sterile leaves quite dimorphic, the sterile to 35 cm

long, the fertile to 50 cm long, ultimate segments

with free venation. Soral lines broken by all or

nearly all of the sinuses.

On hillsides in humus or clay soil, ca. 3200 m,

Cajamarca and Amazonas.

Peru to Argentina; southern Brazil.

Cajamarca: Prov. Cutervo, 10 km NW of Socota, Stork

& Horton 10127 (F, uc, us). Amazonas: Chachapoyas,

Soukup 4926 (us).

doba, Lorentz 79; Argentina, (Tucuman), Tucu-

man, Lorentz & Hieronymus (B!), both collections

on one sheet; photo, GH.

Stem scales usually ovate-lanceolate. Petiole

smooth or slightly short-pubescent at the apex,

with 1 vascular bundle near the base. Fertile and

sterile leaves moderately dimorphic, the sterile to

20 cm long, the fertile to 35 cm long, ultimate

segments with areolae along the midveins, the veins

free toward the margin. Soral lines broken by the

sinuses.

Rocky places, 1200-2500 m, Cuzco.

Peru to southern Brazil.

The lamina with only partly areolate venation

is distinctive of this species.

Cuzco: Potrero, Tryon & Tryon 5377 (BM, F, MO, u,

us, USM), Coronado 115 (GH, uc, us). Yucay, Soukup

921 (F). Dist. Santa Ana, Herrera 87 la (us).

2. Doryopteris lomariacea Klotzsch, Linnaea 20:

343. 1847. TYPE: "British Guiana", Schom-

burgk 1197 (holotype, B?; isotypes, BM!, K!).

Stem scales long and narrow. Petiole usually

smooth, with 2 vascular bundles near the base.

Fertile and sterile leaves strongly dimorphic, the

sterile to 50 cm long, the fertile to 120 cm long,

ultimate segments with free venation. Soral lines

continuous around the sinuses.

Open Sphagnum swamp, 1 700 m, Pasco.

Southern Brazil and Paraguay; Peru; Guyana.

Pasco: Enefias, Pichis Trail, (as Junin), Killip & Smith

2 5709 (F, us).

3. Doryopteris lorentzii (Hieron.) Diels, Nat.

pflanzenfam. 1(4): 270. 1899.

Pellaea lorentzii Hieron., Bot. Jahrb. Syst. 22: 392.

1896. SYNTYPES: Argentina. (Cordoba), Cor-

4. Doryopteris pedata (L.) Fee var. palmata

(Willd.) Hicken, Revista Mus. La Plata, Secc.

Bot. 15: 253. 1908. Figure 10.

Pteris palmata Willd., Sp. pi. ed. 4, 5: 357. 1810.

TYPE: Venezuela, (Dist. Federal. Caracas). Bre-

demeyer (holotype. Herb. Willd. 19957, B!; pho-

tos, GH, NY, us).

Dorvopteris palmata (Willd.) John Sm., J. bot. (Hook-

er) 4: 163. 1841.

Stem scales usually ovate-lanceolate. Petiole

usually very short-pubescent, with 1 vascular bun-

dle near the base. Fertile and sterile leaves mod-

erately dimorphic, both to 40 cm long, and usually

with proliferous buds at the base of the lamina,

ultimate segments with fully areolate venation.

Soral lines usually broken by the sinuses.

Rocky woods, hillsides, crevices of stone walls,

1000-2200 m, Cajamarca and Amazonas south to

Cuzco.

Mexico south to Bolivia.

Buds are usually present at the base of the lam-

ina and these will grow into small plants, especially

on the older leaves and those that have reclined

44

FIELDIANA: BOTANY

5 mm

FIG. 10. Doryopteris pedata var. palmata: a, habit; b, young plant from proliferous bud at base of lamina; c,

portion of indusia interrupted at segment sinus, (a, b from Tryon & Tryon 5369, F, c from Hutchison & Wright 3839,

F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

45

to the ground. Adult plants then may develop from

these small ones.

Cajamarca: Prov. Santa Cruz, Bosque de Monteseco,

Sagdstegui et al. 12398 (F, MO). Prov. Santa Cruz, ca. 5

km above Monte Seco, Dillon et al. 4424 (F, GH). Ama-

zonas: Prov. Bongara, Hutchison & Wright 3839 (F, GH).

Junin: Yaupi, Woytkowski 6424 (MO, us). Huancavelica:

Prov. Tayacaja, SE of Tintay, Tovar 4598 (GH). Cuzco:

Prov. La Convencion, Potrero, Tryon & Tryon 5369 (BM,

F, GH, MO, u, us, USM). Quillabamba, Coronado 114 (GH,

uc, us). Yucay, Soukup 920 (F).

XI. IU- mi mi it is

Hemionitis L., Sp. pi. 1077. 1753; Gen. pi. ed. 5,

485. 1754. TYPE: Hemionitis palmata L. Fig-

ure 11.

Gymnopteris Bernh., J. Bot. (Schrader) 1799 (1): 297.

1 799. TYPE: Acrostichum rufum L. (Gymnopteris

rufa (L.) Underw.) = Hemionitis rufa (L.) Sw.

Terrestrial or less often rupestral. Stem short-

creeping to erect, small, bearing scales intergrading

to trichomes. Leaves monomorphic to dimorphic

and the fertile ones longer and more erect, borne

in a cluster, petiole with 1 vascular bundle near

the base. Lamina of various architecture, usually

pedate or pinnate, pubescent, veins free to wholly

anastomosing without included veinlets. Sporan-

gia in long lines along free veins or along the anas-

tomosing veins, indusium and paraphyses absent.

Spores tetrahedral-globose, trilete, cristate, echi-

nate, or tuberculate.

Hemionitis is a tropical American genus of sev-

en species, with three of them in Peru. Hemionitis

rufa and H. tomentosa are distinctive in Peru.

However, in Argentina to southern Brazil and

rarely in Peru is H. tomentosa var. subrufa Ro-

senst., which seems similar to H. rufa. It has entire

pinnae but is separable from H. rufa by the long

stalks and cordate base, especially of the basal pin-

nae.

Reference

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

ionitis, pp. 278-284, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Species of Hemionitis

a. Lamina pinnate, with free venation b

b. Some pinnae 1 -(rarely 2-)pinnate, or several pinnae lobed, or all pinnae entire and then the basal

ones cordate and long-stalked 1 . H. tomentosa

b. All pinnae entire, rarely 1 lobe present, the basal ones broadly cuneate to truncate and short-

stalked 2. H. rufa

a. Lamina deeply lobed, with areolate venation 3. H. palmata

1 . Hemionitis tomentosa (Lam.) Raddi, Opusc. Sci.

284. 1819.

Asplenium tomentosum Lam., Encycl. 2: 308. 1786.

SYNTYPES: Brazil, Commerson, Dombey. A

sheet in Herb. Lamarck, P!; photo, GH: "Asplen-

ium tomentosum e Brasilia," without collector, is

this species.

Gvmnopteris tomentosa (Lam.) Underw., Bull. Torrey

Bot. Club 29: 627. 1902.

Leaves ca. 20-75 cm long. Petiole long, it and

the rachis reddish brown to atropurpureous, pu-

bescent with short, acicular trichomes and fewer,

long, multicellular ones. Lamina suboblong to sub-

deltoid, 1 -pinnate and the pinnae entire, or a few

lobed, cordate or subcordate at the base, or 2-

(rarely partly 3-)pinnate, pinnae stalked, the basal

ones with longer stalks than those above, the dark

color of the stalk often continued into the segment,

both surfaces more or less pubescent with acicular,

multicellular, nearly appressed trichomes.

Forest slopes, stony clearings, and in woods,

700-1600 m, Amazonas to Cuzco.

Peru to Argentina, Paraguay, and Brazil.

Amazonas: Prov. Bagua. valley of Rio Utcubamba,

Hutchison 1476 (GH). San Martin: Tarapoto, Spruce 3992

(B, GH), Woytkowski 35228 (uc). Prov. San Martin, Tar-

apoto-Yurimaguas, Sagdstegui 6857 (F). Huanuco: Near

Pozuzo, Macbride 4788 (F, us). Pasco: Rio Paucartambo,

46

FIELDIANA: BOTANY

1O mm

FIG. 1 1 . Hemionitis palmata: a, habit, b, lamina apex, abaxial side. (From Klug 4288, F.)

near Perene Bridge, (as Junin), Killip & Smith 25345

(us). Junin: La Merced, Macbride 5303 (F). Rio Penedo,

N of La Merced, Killip & Smith 23592 (GH, us). Cuzco:

Quillabamba, Coronado 122 (GH, uc, us). Santa Ana,

Cook & Gilbert 1537 (GH, us). Potrero, Tryon & Tryon

5374 (GH, u, us, USM).

2. Hemionitis rufa (L.) Sw., J. Hot. (Schrader) 1 800

(2): 16. 1802.

Acrostichum rufum L., Syst. nat. ed. 10, 1320. 1759.

(Not Pteris ruffa L., Sp. pi. 1074. 1753, which is

maintained in the Syst. nat. and probably typified

by a Hort. Cliff, specimen in Herb. Banks, BM.)

TYPE: Sloane, Voy. Jamaica /. 45, f. 1 or LINN

1245.6; photo, A.

Gymnopleris rufa (L.) Underw., Bull. Torrey Bot. Club

29:627. 1902.

Leaves ca. 20-60 cm long. Petiole usually long,

it and the rachis dark reddish brown to atropur-

pureous, pubescent with short, acicular trichomes

and fewer, long, multicellular ones. Lamina nar-

rowly elliptic to lanceolate, 1 -pinnate, the pinnae

entire, or rarely a lobe present, usually cuneate at

the base, short-stalked, the dark color of the stalk

not continued into the segment, both surfaces more

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

47

or less pubescent with acicular, multicellular, near-

ly appressed trichomes.

In forests, 400-800 m, San Martin.

Southern Mexico to Panama; Greater Antilles;

Surinam to Colombia and south to Peru.

San Martin: Juan Jui, Klug 4174 (F, GH, uc, us).

3. Hemionitis palmata L., Sp. pi. 1077. 1753.

TYPE: Plumier, Descr. pi. Amer. /. 33, f. 1

or LINN 1248.3; photo, A. Figure 11.

Leaves dimorphic, the sterile ca. 3-10 cm long,

3-lobed or pedately 5-lobed, borne in a cluster, the

fertile ca. 10-25 cm long, few, erect, pedately 5-

lobed. Petiole brown to dark brown, pubescent

with multicellular, slender-tipped trichomes.

Principal lobes shallowly lobed or crenate, with

pubescence similar to that of the petiole.

In forests, 120-900 m, San Martin, Loreto, and

Cuzco.

Mexico and Central America; West Indies;

Guianas to Colombia, south to Bolivia; Brazil.

The leaves bear small dormant buds in the si-

nuses along the margin and these may serve to

propagate the plants vegetatively.

San Martin: Tarapoto, Spruce 3993 (GH, us). Juan Jui,

Klug 4288 (F, GH, MO, uc, us, USM). Near Tarapoto, LI.

Williams 6793 (F, us). Loreto: Iquitos and vicinity, LI.

Williams 3559 (us). Cuzco: Santa Ana, Cook & Gilbert

1477 (us).

XII. Trachypteris

Trachypteris Christ, Schweiz. Naturfors. Ges. 36

(Monogr. Elaphoglossum): 150. 1899. TYPE:

Trachypteris aureonitens (Hooker) Christ (Ac-

rostichum aureonitens Hooker) = Trachyp-

teris pinnata (Hooker f.) C. Chr. Figure 12.

Saffordia Maxon, Smithsonian Misc. Coll. 61(4): 1.

1913. TYPE: Saffordia induta Maxon = Tra-

chypteris induta (Maxon) R. & A. Tryon.

Terrestrial or rupestral. Stem decumbent or

erect, small, bearing scales. Leaves monomorphic

or dimorphic with the sterile entire, in a basal

rosette, and the fertile erect and pinnate, densely

scaly abaxially, glabrous to thinly scaly adaxially,

veins anastomosing without included veinlets.

Sporangia borne in a narrow to broad band, or

from the costa to the margin, on and between the

veins, indusium and paraphyses absent. Spores

globose, trilete, strongly cristate.

This is a small genus of three species. Two of

them are in America and Peru and the third is in

Madagascar.

Reference

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

chypteris, pp. 302-306, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Species of Trachypteris

a. Sterile leaves entire, in a basal rosette; fertile leaves erect, pinnate 1 . T. pinnata

a. Sterile and fertile leaves pedate 2. T. induta

1 . Trachypteris pinnata (Hooker f.) C. Chr., Index

fil. 634. 1906. Figure 12a.

Hemionitis pinnata Hooker f., Trans. Linn. Soc. Lon-

don 20: 167. 1847. LECTOTYPE (designated by

Porter, J. Linn. Soc. Bot. 81: 94. 1980): Ecuador,

Galapagos Islands. Charles Island, Darwin (ho-

lotype, CGE).

Acrostichum aureonitens Hooker, Hooker's Icon. PI.

/. 933 (Cent. Ferns t. 33). 1854. LECTOTYPE

(designated by Tryon. Contr. Gray Herb. 1 94: 86.

1964): Ecuador, Galapagos Islands Cuming 109

(holotype, K!; photo, GH; LECTOPARATYPE:

Ecuador, Galapagos Islands, Chatham Island,

Capt. Wood, K.!; photo, GH).

Sterile leaves several, ca. 5-8 cm long, more or

less prostrate, spatulate to obovate, sessile or near-

ly so, entire or rarely with a few lobes, with scales

on the abaxial surface that are appressed and im-

bricate. Fertile leaves few, ca. 1 2-25 cm long, erect,

subdeltoid to short-oblong, with a long, dark-col-

48

FIELDIANA: BOTANY

5 cm

.."'" \

. 1

••

f^ti-

(,-:•: ^".ni

V •-<--' »*<•'.•

"- '-^i' *>*

.-^^o ?>>-. r

, e.^.

FIG. 12. Trachypteris pinnata: a. habit. Trachypteris induta: b, leaf, abaxial side; c, leaf margin, adaxial side, (a

from Tovar 4603, OH; b, c from Madison 1126, GH.)

ored, often scaly petiole, scales on the abaxial sur-

face similar to those on the sterile leaf.

In rocky woods and in forests, 680-2750 m,

Cajamarca, San Martin, Huancavelica, and Cuzco.

Ecuador and Archipielago de Colon (Galapagos

Islands); Peru to Argentina and Brazil.

A critical study of the American material may

show that two or three geographic varieties are

present.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

49

Cajamarca: Dist. Espinal, Puente El Papayo, Vdsquez

3077 (USM). Rio Crisnejas, Sanchez 45 (GH). Jaen, Rauh

P2160 (B). Prov. Jaen, Rio Chamaya, Hutchison 1424

(F, GH, uc). San Martin: Tambo de Carrizal, Stubel 1016

(B). Huancavelica: SE of Tintay, Tovar 4603 (GH). Cuzco:

Santa Ana, Cook & Gilbert 1480 (us). Huadquina, Biies

1339 (us). Sahuayacu. Biies 837 (us). Prov. La Conven-

cion, Rosariomayo, Vargas 22343 (GH).

2. Trachypteris induta (Maxon) R. & A. Tryon,

Rhodora 83: 137. 1981. Figure 12b-c.

Saffordia induta Maxon, Smithsonian Misc. Coll.

61(4): 2, /. 1-2. 1913. TYPE: Peru, (Lima), along

the Arroya (La Oroya) Railway in the mountains

back of Lima, Safford 989 (holotype, us!; iso-

types, B!, GH!, K.!, uc!, us!).

Sterile and fertile leaves usually few, mono-

morphic or nearly so, ca. 1 5-30 cm long. Petiole

dark in color, usually somewhat scaly. Lamina

more or less suborbicular, pedate, densely scaly

abaxially.

Among rocks, 750-2900 m, Cajamarca and

Amazonas to Lima.

Endemic to Peru.

The scales at the base of the petiole are pink

which is an unusual color among ferns.

Cajamarca: Prov. Contumaza, Cascas-Contumaza,

Sagdstegui 7670 (F, MO), Lopez et al. 2936 (GH, us). Prov.

Contumaza, Contumaza-Toledo, Sagdstegui et al. 12604

(MO), 12608 (HUT). Prov. Contumaza, between Santiago

and Guzmango, Sagdstegui 2936 (GH). Amazonas: Prov.

Chachapoyas, Cerros Calla Calla, Balsas to Leimebam-

ba, Hutchison & Wright 4966 (GH). Rio Maranon, at

Balsas, Madison 1126 (GH). La Libertad: Prov. Otuzco,

Huaranchal, Sagdstegui 205 (GH). Lima: Dist. Surco, km.

68, Carretera Central, Sounders 218 (BM, GH). Matucana,

Rauh P186 (B). Prov. Huarochiri, Molle, Valencia &

Franke 559 (USM).

XIII. Ceratopteris

Ceratopteris Brongn., Bull. Sci. Soc. Philom. Paris

III, 8: 186. 1821. TYPE: Ceratopteris thalic-

troides (L.) Brongn. (Acrostichum thalictroides

L.). Figure 13.

Terrestrial, palustral, or aquatic. Stem small,

erect, bearing a few scales. Leaves dimorphic, the

fertile more erect than the sterile, longer, and the

lamina usually more complex, petiole with 4-many

vascular bundles. Lamina deeply lobed in some

sterile leaves, or 1-5-pinnate, glabrous, veins anas-

tomosing without included veinlets. Sporangia

solitary, borne well apart on the veins, covered by

a well-differentiated marginal indusium, paraph-

yses absent. Spores globose, trilete, with promi-

nent mostly parallel ridges.

Ceratopteris is a predominantly aquatic genus

of three species. It is widely distributed in the trop-

ics, with a single species in the Amazon basin of

Peru.

Reference

LLOYD, R. M. 1974. Systematics of the genus

Ceratopteris Brongn. II. Taxonomy. Brittonia,

26: 139-160.

1. Ceratopteris pteridoides (Hooker) Hieron., Bot.

Jahrb. Syst. 34: 561. 1905. Figure 13.

Parkeria pteridoides Hooker, Exot. fl. 2, t. 147. 1825.

TYPE: "British Guiana", District of Essequibo,

Parker (holotype, K!; photo, GH; isotype, GH!).

Sterile leaves more or less prostrate, ca. 3-20

cm long, with a broad, fleshy, usually inflated pet-

iole, lamina deltoid, thin-herbaceous, with broad

segments, old leaves often with adventitious buds.

Fertile leaves more or less erect, ca. 5-50 cm long,

with a large inflated petiole, with linear segments.

In slow streams, in lagoons, and along rivers,

ca. 100 m, Loreto.

Tropical and subtropical America, south to Bo-

livia, Argentina, and Brazil.

This species is typically a floating aquatic, the

inflated petiole providing buoyancy to the plant.

The adventitious buds on the leaves frequently

develop into separate plants.

The other species in America, Ceratopteris thal-

ictroides (L.) Brongn., may occur in Peru. It typ-

ically roots in mud or in shallow water and lacks

inflated petioles. Ceratopteris richardii Brongn. is

sometimes separated from C. thalictroides, pri-

marily by its spore number of 16 in a sporangium

rather than 32, which does not seem to be a species

character.

Loreto: Rio Itaya, Iquitos, Asplund 14626 (us). Iqui-

tos, Killip & Smith 27427 (us). Prov. Maynas, Isla Ro-

difia, opposite Leticia, Plowman et al. 6380 (F, GH, us).

Prov. Maynas, above Bombabaje, McDaniel & Rimachi

1 7 329 (F, MO).

50

FIELDIANA: BOTANY

FIG. 1 3. Ceratopteris pteridoides: a, sterile leaf; b, fertile leaf; c, fertile segments, abaxial side, (a from Kiltip &

Smith 14791, Colombia, F, b from Austin et al. 7040, Brazil, F, c from Holm-Nielsen el al. 2654, Ecuador, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

51

XIV. Adiantum

Adiantum L., Sp. pi. 1094. 1753; Gen. pi. 485.

1754. TYPE: Adiantum capillus-veneris L.

Figure 14.

Terrestrial or less often rupestral. Stem small

and suberect, or stout and short-creeping, or slen-

der and long-creeping, bearing scales. Leaves small

to very large, usually pinnate (1-6-pinnate), usu-

ally glabrous, less often sparingly scaly, pubescent

or farinose abaxially, veins free or rarely anasto-

mosing without included veinlets. Sori marginal,

not paraphysate, the sporangia borne along, or also

between, the vein ends that extend into the in-

dusium which is formed by a reflexed, modified

lobe, or continuous modified margin. Spores tet-

rahedral-globose or globose, trilete, rather smooth

or irregularly cristate, or rugose.

Adiantum is a large, widely distributed genus of

about 1 50 species. Tropical America is the richest

region for species, and there are 34 in Peru. A

single species, Adiantum capillus-veneris is con-

sidered to be adventive. The genus is exceptionally

well defined by the sporangia that are borne on

the modified, reflexed portions of the margin that

serve as indusia.

In species with cuneate to broadly flabellate or

nearly orbicular ultimate segments, the sori are

borne on the convex side (apical side) of the ulti-

mate segments. In species with rather to definitely

elongate ultimate segments, the sori may be borne

on the upper side (the acroscopic side of the seg-

ment), on the lower side (the basiscopic side of the

segment), on the distal side (the apical side of the

segment), or on the inner side (the side adjacent

to the axis that bears the segment).

Some species are quite distinctive, while others

are ill-defined members of complexes. Hybridiza-

tion is probably rather frequent, tending to obscure

the characters of otherwise distinctive taxa. The

species have been arranged, insofar as possible, to

bring evidently related species together, although

primitive and advanced characters as well as evo-

lutionary relations are uncertain.

References

KRAMER, K. U. 1978. The pteridophytes of Sur-

iname. Uitgaven Natuurw. Studiekring Suri-

name Ned. Antillen, 93: 1-198.

TRYON, R. 1964. The ferns of Peru. Contr. Gray

Herb., 194: 1-253.

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

tum, pp. 319-332, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Species of Adiantum

a. A few to many, orbicular to long indusia on the fertile segment b

b. Lamina with 2 equal, strongly recurved rachises, these bearing 1 -pinnate branches on only one

side, or with the rachises unequal and 1 bearing a well-developed terminal pinna

1 2. A. patens

b. Lamina with a single rachis bearing lateral pinnae c

c. Lamina lacking a 1 -pinnate terminal pinna (either gradually reduced to the apex, or with

a nearly entire to lobed terminal pinna), or with a 1 -pinnate terminal pinna definitely larger

than the adjacent lateral ones d

d. Ultimate segments mostly rather broadly flabellate, many or most of them ca. 2 cm

or more long, usually deeply cleft into spreading lobes; lamina 2- to usually 3- or 4-

pinnate 11. A. digitatum

d. Ultimate segments of various shape, if mostly broadly flabellate then either entire or

nearly so, or ca. 1 cm long, or the lamina 1 -pinnate e

e. Rachis or pinna-rachises glabrous, puberulent, or pubescent; most ultimate seg-

ments definitely, although sometimes shortly, stalked, lacking a distinct midvein

or with one and very long-stalked f

f. Most ultimate segments small, ca. 1 cm in length, or rarely to ca. 2.5 cm and

then the fertile without a definite apex and the indusia continuous along the

convex side g

g. Stalk of the ultimate segments blackish, pubescent, the color ending

abruptly at the base of the segment 9. A. imbricatum

52

FIELDIANA: BOTANY

FIG. 14. Adiantum concinnum: a, habit; b, base of pinna, abaxial side. Adiantum capillus-veneris: c, ultimate

segment. Adiantum pulverulentum: d, portion of rachis with fertile segment, abaxial side. (Adapted from Stolze, Ferns

& fern allies of Guatemala, 1981.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

53

g. Stalk of the ultimate segments light-colored to reddish brown, or if black-

ish then glabrous h

h. Most or many ultimate segments articulate at the junction with the

stalk, deciduous by a sharp, clean break, the distal portion of the

stalk slightly enlarged; the dark color of the stalk ending abruptly at

the base of the segment i

i. Lamina 2— 4-pinnate, many pinnae 1 -pinnate or more complex;

indusia orbicular to orbicular-reniform . 10. A. orbignyanum

i. Lamina 1 -pinnate, the pinnae nearly entire to rather deeply in-

cised; indusia mostly oblong to lunate 14. A. deflectens

h. Segments not articulate, deciduous by the irregular fracture of the

stalk, the color of the apex of the stalk usually continuing into the

segment base j

j. Lamina axes more or less whitish puberulent or pubescent, at

least adaxially k

k. The distal 1 -pinnate pinnae with the inner, upper pinnule

strongly overlaying the rachis; basal pinnae short-stalked

7. A. henslovianum

k. The distal 1 -pinnate pinnae with the inner, upper pinnule

not or hardly overlaying the rachis; basal pinnae long-stalked

8. A. lohatum

j. Lamina axes glabrous 1

1. Basal pinnae definitely stalked, or if very short-stalked then

the ultimate segments overlaying the rachis beyond the base

of the lamina nearly cuneate m

m. Sterile margins of the ultimate segments with each vein

ending in a tooth; segments cuneate with a pronounced

cartilaginous border 4. A. capillus-veneris

m. Sterile margins of the ultimate segments with each vein

ending in a sinus between the definite to indefinite teeth,

or the margin nearly entire n

n. Indusia orbicular to suborbicular; ultimate seg-

ments flabellate, or symmetrically or asymmetri-

cally cuneate o

o. Lamina usually 3- or 4-pinnate, or 2-pinnate

well beyond the base 1 . A. raddianum

o. Lamina usually 1 -pinnate with large, long-

stalked, flabellate pinnae, rarely 2-pinnate ba-

sally 13. A. ruizianum

n. Some to many indusia oblong to lunate; many ul-

timate segments orbicular to suborbicular . . . . p

p. Scales borne at the apex of the stem (not those

of a leaf-base or bud) with a long, filiform,

tortuous tip, the margins denticulate or ciliate

2. A. poiretii

p. Scales borne at the apex of the stem (not those

of a leaf-base or bud) with a short, nearly

straight or curved tip and entire margins . . .

3. A. chilense

1. Basal pinnae nearly subsessile, very short-stalked; the ul-

timate segments overlaying the rachis beyond the base of

the lamina nearly suborbicular q

q. Inner, upper pinnule of lower pinnae divided into 2 or

54 FIELDIANA: BOTANY

more ultimate segments; not on lomas

5. A. concinnum

q. Inner, upper pinnule of the lower pinnae simple; mostly

on lomas 6. A. subvolubile

f. Most ultimate segments large, ca. 3 cm or more in length, rarely 2 cm long

and the fertile with a definite apex which, beyond the lower pinnules of a

penultimate segment, is usually sterile or if fertile then subacute r

r. Dark color of the segment stalk entering the base of the segment abaxially;

segments glaucous abaxially; indusia long, often arcuate

28. A. platyphyllum

r. Dark color of the segment stalk not entering the base of the segment

abaxially; segments green abaxially; indusia roundish to lunate s

s. Ultimate segments mostly bluntly acute to rounded apically

29. A. peruvianum

s. Ultimate segments mostly ovate-cuneate, concavely acuminate,

sometimes with an acuminate auricle 30. A. anceps

e. Rachis and pinna-rachises scaly or rarely glabrous; ultimate segments (often pin-

nae) sessile or subsessile, or mostly some basal ones short-stalked, usually with a

distinct midvein well beyond the base t

t. Sterile margins regularly serrate; segments dull, usually glaucous abaxially;

false veins not or hardly evident abaxially 24. A. petiolatum

t. Sterile margins irregularly biserrate; segments green, usually shining abaxially;

false veins somewhat to usually very evident abaxially . . . 25. A. obliquum

c. Lamina with a 1 -pinnate terminal pinna more or less equal in size to the adjacent lateral

ones u

u. Rachis and pinna-rachises glabrous, puberulent, or pubescent v

v. Pinnules acute or subacute, usually spaced; pinna-rachises puberulent on all sides,

or sparsely so or not on the abaxial side 17. A. urophyilum

v. Pinnules broadly obtuse, mostly imbricate; pinna-rachises glabrous, puberulent,

or pubescent only on the adaxial side, or sometimes also on the lateral side . . w

w. Pinna-rachises puberulent or pubescent on the adaxial or also lateral sides

16. A. tomentosum

w. Pinna-rachises glabrous on all sides x

x. Upper side of the fertile segments entire or nearly so; most indusia or-

bicular to suborbicular; penultimate segments (usually pinnae) gradually

reduced to a slender, acuminate apex, with small segments near the apex

18. A. macrocladum

x. Upper side of the fertile segments shallowly to rather deeply cleft; indusia

usually oblong to lunate; penultimate segments (usually pinnae) rather

abruptly reduced to a large, acute apex, with large segments near the apex

27. A. mathewsianum

u. Rachis and pinna-rachises scaly, sometimes also with trichomes y

y. Lamina commonly 4-5-pinnate at the base; basal pinnae very long-stalked; fertile

segments cleft on the upper side; indusia nearly orbicular ... 15. A. pectinatum

y. Lamina 2-pinnate, very rarely 3-pinnate at the base; basal pinnae rather short-

stalked z

z. Segments dull abaxially, slightly to strongly glaucous aa

aa. Stem rather short-creeping, with crowded petioles; ultimate segments

usually abundantly to rarely very slightly pubescent abaxially; pinnae

usually acuminate, most of the distal segments (not lobes of the apex)

less than half the length of the longest on the pinna

20. A. terminatum

aa. Stem often very slender and long-creeping, with the petioles well spaced;

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II. 55

ultimate segment usually glabrous abaxially or somewhat pubescent and

(or) scaly; pinnae usually acute, most with the distal segment (not lobe

of the apex) not much reduced, about half or more the length of the

longest on the pinna 23. A. latifolium

z. Segments green, usually shining abaxially bb

bb. Segments usually coriaceous; many ultimate segments less than twice as

long as broad; sterile margins evenly and usually finely serrate

21. A. serratodentatum

bb. Segments usually herbaceous; many ultimate segments more than twice

as long as broad; sterile margins unevenly and coarsely serrate

19. A. tetraphyllum

a. One usually long indusium on each side of the fertile pinna or pinnule, or only on the upper side,

or a few additional short ones present cc

cc. Lamina 2-pinnate, with a terminal pinna about equal in size to the lateral ones which are

gradually reduced to a usually acuminate apex 22. A. pulverulentum

cc. Lamina 1 -pinnate, or if 2-pinnate, then the apical segment of the 1 -pinnate pinnae large, about

equal to the simple pinnae, or the 1 -pinnate terminal pinna much larger than the 1 -pinnate

lateral pinnae dd

dd. Rachis with scales or trichomes, or both; pinnae alternate, rarely subopposite ee

ee. Stem scales iridescent; rachis with simple or often stellate trichomes, and often with

a few scales on the adaxial side, usually glabrous abaxially; lamina 1- or 2-pinnate

26. A. alarconianum

ee. Stem scales dull; rachis with scales and often also trichomes on all sides, rarely glabrate

ff

ff. Rachis with trichomes and scales; lamina 1- or 2-pinnate; segments glabrous or

slightly scaly or pubescent abaxially; ultimate segments (usually pinnae) usually

ovate, lanceolate or narrowly to broadly elliptic gg

gg. The dark color of the segment stalk not or hardly continuing into the base of

the segment abaxially 3 1 . A. lucidum

gg. The dark color of the segment stalk continuing well into the base of the segment

abaxially 32. A. poeppigianum

ff. Rachis densely scaly, the scales freely ciliate; lamina 1 -pinnate; the pinnae with

long trichomes abaxially, very narrowly long-triangular, long-attenuate

33. A. scalare

dd. Rachis glabrous; lamina 1 -pinnate; the pinnae usually opposite . . 34. A. macrophyllum

1. Adiantum raddianum Presl, Tent, pterid. 158.

1836, based on Raddi, PI. bras. nov. gen. 1,

/. 78, f. 2. 1825.

Adiantum cuneatum Langsd. & Fisch., Icon. fil. 23, /.

26. 1 8 1 0, not Forster, 1 786. TYPE: Brazil, (Santa

Catarina), "Ins. St. Catharina," Langsdorff (ho-

lotype, LE!; photo, GH; isotype, BM!).

Adiantum tinctum Moore, Gard. Chron. 1862: 932.

TYPE: Peru, Hort. Veitch (holotype, K.!; photo,

GH).

Adiantum amabile Moore, Gard. Chron. 1868: 1090,

not Liebm. 1849. TYPE: Peru, Pearce (holotype,

K!; photo, GH).

Adiantum decorum Moore, Gard. Chron. 1 869: 582.

TYPE: Peru, Pearce (holotype, K!; photo, GH).

Adiantum moorei Baker, Gard. Chron. 1873: 811,

based on A. amabile Moore and with the same

type.

Stem rather stout, very short-creeping, multi-

cipital. Leaves ca. 15-55 cm tall, petiole glabrous

or slightly scaly at the base, rachis glabrous. Lam-

ina elongate-triangular to lanceolate-ovate, broad-

ly ovate or deltoid, commonly 3-pinnate, less often

2- or 4-pinnate. Pinnae stalked, ultimate segments

cuneate and symmetrical to broadly cuneate-fla-

bellate and asymmetrical, glabrous or slightly far-

inose, or with sessile glands abaxially, not artic-

ulate, the color of the apex of the stalk usually

passing into the base of the segment, sterile mar-

gins with each vein ending in a more or less well-

56

FIELDIANA: BOTANY

developed sinus. Indusia few to several, roundish

to suborbicular-reniform.

In forests, open woods, thickets, on rocky banks

and cliffs, borders of irrigation ditches, 400—4000

m, Tumbes south to Puno.

Tropical America.

Adiantum raddianum may be separated from

the closely related A. poiretii by its orbicular to

suborbicular indusia.

The ultimate segments vary from usually gla-

brous abaxially to rather glandular or sometimes

slightly farinose. These are sporadic variations and

relatively rare within the species.

The stalks of the ultimate segments are usually

dark in color, including the apex, and this color

passes into the base of the ultimate segment. In

some collections, however, the dark color does not

extend to the apex of the stalk and in others it

does, but does not extend into the base of the

segment. The latter variation is similar to Adian-

tum orbignyanum, although there is no articula-

tion at the apex of the stalk and the segments break

off irregularly.

Tumbes: El Cancho to Cotrina, Coronado 233 (uc).

Cajamarca: Prov. Contumaza, Sagdstegui et al. 10879

(GH, MO). Prov. Celendin, 4 km E of Celendin, Hutchison

& Wright 5 189 A (F, GH, us). El Puquio, Guzmango, Sa-

gastegui 3917 (GH, HUT). Amazonas: 1-2 km W of Mol-

inopampa, Wurdack 1507 (F, GH, uc). La Libertad: Prov.

Otuzco, Samne, Angulo 1246 (GH). San Martin: Tara-

poto. Spruce 4664 (K). Ancash: Prov. Huaraz, 10 km

from Cachabamba, Smith & Buddensiek 10903 (F). Prov.

Bolognesi, Huasta, Cerrate 2442 (GH, USM). Huanuco:

Mito, Macbride & Featherstone 1435 (F, GH, us). Car-

pish, Coronado 80 (GH, uc, us). Lima: San Buenaventura,

Pennell 14559 (F, GH, NY, PH, us). Junin: La Merced,

Soukup 2570 (F, us, USM). 12 km SW of San Ramon,

Tryon & Tryon 5446 (BM, F, MO, u, us, USM). Huancav-

elica: A 4 km de Conaica, Tovar 979 (GH, USM). Apu-

rimac: Chincheros, West 3694 (MO, uc). Cuzco: Potrero,

near Quillabamba, Tryon & Tryon 5368 (BM, F, GH, MO,

u, us, USM), 5376 (BM, F, GH, MO, u, us, USM). Puno: Juli,

near Lake Titicaca, Shepard 14 (NY).

2. Adiantum poiretii Wikstrom, Kongl. Vetensk.

Akad. Handl. 1825: 443. 1826, nom. nov. for

Adiantum crenatum Poiret, not Willd., March,

1810.

Adiantum crenatum Poiret in Lam., Encycl. suppl. 1 :

137, Sept. 1810. TYPE: Tristan d'Acugha (Tris-

tan da Cunha), Petit-Thouars (holotype, Herb.

Jussieu 1427, P!).

Adiantum thalictroides Schlecht., Adumbr. pi. 5: 53.

1 832. TYPE: Mauritius, Petit- Thouars (holotype,

Herb. Willd. 20101, B!; photo, GH).

Stem slender, rather long-creeping, with tufts of

petioles borne at intervals, or sometimes more

compact. Leaves ca. 20-50 cm. long, petiole gla-

brous or slightly scaly at the base, rachis glabrous

or rarely slightly ceraceous. Lamina elongate-del-

toid, to broadly ovate-deltoid, 3-pinnate (rarely 2-

or 4-pinnate). Pinnae stalked, ultimate segments

cuneate- to subcordate-flabellate, mostly subor-

bicular, rather symmetrical, glabrous or occasion-

ally somewhat farinose or glandular-pubescent

abaxially, not articulate, the color of the apex of

the stalk usually passing into the base of the seg-

ment, sterile margins with each vein ending in a

more or less well-developed sinus. Indusia few,

oblong to long-lunate, or the smallest ones round-

ish.

Wooded hillsides, open woods, thickets, lomas,

and rocky places, 1200-4200 m, Cajamarca and

Amazonas, south to Puno and Arequipa.

Mexico and the West Indies, south to Argentina;

mid-Atlantic islands and Africa and adjacent is-

lands.

The many orbicular to suborbicular ultimate

segments, with at least some oblong to lunate in-

dusia, and the usually creeping stem with the leaves

borne at intervals in clusters, distinguish this

species from Adiantum raddianum.

It differs from Adiantum chilense principally in

the tufts of scales at the apex of the stem. These

scales are ciliate or denticulate and have a long,

filiform, tortuous tip in A. poiretii, while they are

entire-margined and have a short, straight or curved

tip in A. chilense. Rarely the scales of A. poiretii

have a rather short, nearly straight tip although

the margins are ciliate-denticulate.

Pichi-Sermolli (Webbia 12: 693-695. 1957)

considered the plants of Tristan da Cunha to rep-

resent a distinct species and restricted the name

Adiantum poiretii to it. The widespread American

and African species is then known as Adiantum

thalictroides. However, a thorough study of the

variations of this widespread species needs to be

made, including Adiantum chilense, before an ap-

propriate classification of the group can be made.

Rare specimens that have the ultimate segments

somewhat farinose or pubescent abaxially occur

scattered through the range of A. poiretii and are

considered as within the variation of the species.

Yellow farina occurs among the sporangia rather

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

57

frequently and this also seems to be a variant of

the species.

Cajamarca: Prov. Celendin, 4 km E of Celendin,

Hutchison & Wright 5189 (F, GH). Prov. Contumaza,

Guzmango, Sagdstegui el al. 10391 (GH, MO, uc). Ama-

zonas: Conila, Soukup 4166 (us). La Libertad: Between

Huamachuco and Cajabamba, Correll & Smith P918

(GH, us). Prov. Santiago de Chuco, Cachicadan, Sagds-

tegui et al. 10391 (F). Ancash: Cerca a Llamac, Cerrate

2371 (GH, USM). Huanuco: Muna, Macbride 3932 (F, us).

Mitotambo, Ferreyra 10382 (GH, USM). Lima: Churin,

Ferreyra 5356 (GH, USM). Pasco: Paucartambo (as Junin),

Woytkowski 6728 (MO). Junin: 10 km E of Huancayo,

Tryon & Tryon 5468 (BM, F, MO, u, us, USM). Huancav-

elica: Andaimarca, Tovar 1815 (GH, USM). Ayacucho:

Between Huanta and Rio Apurimac, Killip & Smith 22320

(F, NY, us). Cuzco: Near town of Machu Picchu, Tryon

& Tryon 5403 (BM, F, MO, u, us, USM). Arequipa: 14 km

S of Arequipa, Eyerdam & Beetle 22123 (GH). Puno:

Salcedo, Soukup 1 (F, GH, uc, us).

3. Adiantum chilense Kaulf. var. hirsutum Hook-

er & Grev., Icon. fil. 2, t. 173. 1830. TYPE:

Chile, Gillies (holotype, K! or E, Herb. Gre-

ville?; photo, GH of K).

Adiantum poiretii var. hirsutum (Hooker & Grev.)

Tryon, Amer. Fern J. 47: 141. 1957.

Stem slender, rather long-creeping, with tufts of

leaves borne at intervals, or sometimes more com-

pact. Leaves ca. 10-30 cm long, petiole glabrous

or pubescent or slightly scaly at the base, rachis

glabrous or pubescent. Lamina elongate-deltoid to

broadly ovate-deltoid, 2-(rarely 3-)pinnate. Pin-

nae stalked, ultimate segments cuneate to subcor-

date-flabellate, mostly suborbicular, rather sym-

metrical, pubescent or also farinose abaxially, not

articulate, the color of the stalk passing into the

segment base or not, sterile margins with each vein

ending in a rather well-developed sinus. Indusia

few, oblong or lunate, or the smaller ones round-

ish.

Terrestrial, lomas of Capac and Atiquipa, 400-

600 m, Arequipa.

Southern Peru and Chile.

Adiantum chilense var. hirsutum is pubescent

and sometimes also somewhat farinose on the

abaxial side of the ultimate segments. Variety chi-

lense is glabrous abaxially and another variety of

Chile (Adiantum suphureum Kaulf.) is densely

bright yellow-farinose abaxially. The species

Adiantum chilense is distributed from southern

Peru to Argentina and Chile; also in the Juan Fer-

nandez Islands. The distinctions from the closely

related Adiantum poiretii are discussed under that

species.

The materials of var. hirsutum of Peru, and in

some cases of Chile, may be rather intermediate

between the two species.

Arequipa: Lomas de Capac, Coronado 42 (GH). Lomas

de Atiquipa, Coronado 31 (GH).

4. Adiantum capillus-veneris L.. Sp. pi. 1 096. 1753.

LECTOTYPE (designated by Pichi-Sermolli

inWebbia 12:678. 1957): LINN 1252.9; pho-

to, A. Figure 14c.

Stem rather slender, short- to long-creeping.

Leaves ca. 15-45 cm long, petiole glabrous, or

slightly scaly at the base, rachis glabrous. Lamina

elongate-deltoid to ovate to elongate-ovate, 2-3-

pinnate. Pinnae stalked, ultimate segments cu-

neate-flabellate to suborbicular, rather symmet-

rical or not, glabrous abaxially, not articulate, the

color of the apex of the stalk passing into the base

of the segment, sterile margins with a cartilaginous

border, each vein ending in a tooth. Indusia few,

nearly squarish to oblong.

Damp, gravelly sea cliffs, banks of irrigation

ditches, and local seepage areas, sea level to 2000

m, Lima and lea.

Tropical and temperate America; Old World.

This species is distinct among those of Peru by

the pronounced cartilaginous border on the ma-

ture segments, the light grayish brown stem scales,

and the veins that end in a tooth of the sterile

margin. Its close correlation with city, suburban,

and resort areas suggests that it is adventive as an

escape from cultivation.

Lima: Laguna de Villa, Coronado 4 (GH, MO, uc, us).

Miraflores, Tryon & Tryon 5216 (BM, F, GH, MO, u, uc,

us, USM). Chorillos, Gentry & Prance 16461 (F, USM).

lea: Huamani (USM).

5. Adiantum concinnum Willd., Sp. pi. ed. 4, 5:

451. 1810. TYPE: Venezuela, (Dist. Federal),

Caracas, Humboldt & Bonpland (holotype,

B!, Herb. Willd. 20099; photo, GH). Figure

14a b.

Stem moderately slender, rather short-creeping,

often multicipital. Leaves ca. 20-80 cm long, pet-

iole glabrous or slightly scaly at the base, rachis

58

FIELDIANA: BOTANY

glabrous. Lamina ovate-oblong to linear-lanceo-

late, 2-pinnate (except at the very base) to gener-

ally 3-pinnate. Pinnae subsessile, the basal pinnule

divided into 2 ultimate segments, at least at the

base of the lamina overlaying the rachis, ultimate

segments cuneate-flabellate to broadly so to su-

borbicular, rather symmetrical or not, glabrous

abaxially, not articulate, the color of the apex of

the stalk passing into the base of the segments,

sterile margins with each vein ending in a sinus.

Indusia few to several, roundish or reniform to

lunate.

In woods and along irrigation ditches, 200-2000

m, Tumbes to Ancash.

Mexico and West Indies, south to Ecuador and

adjacent Peru.

In Peru, especially in Tumbes, there is variation

within plants in the number of ultimate segments

on the basal pinnules of the basal pinnae. Coron-

ado 222 has one to four ultimate segments, and

229 has two to three. With rare exception, A. con-

cinnum has two such ultimate segments, and A.

subvolubile, one.

Tumbes: Between Tumbes and Cancho, Coronado229

(GH, uc). Between Cancho and Cotrina, Coronado 222

(GH, uc). Piura: Hills of Chiarnique, Barbour 2154 (MO).

Cerca a Chanchaque, Ferreyra 10813 (GH, USM). Lam-

bayeque: 20 km from Olmos on road to Jaen, Correll &

Smith P791 (GH). Cajamarca: Prov. Contumaza, alrede-

dores de San Benito, Sagdstegui 3742'/2 (GH). Ancash:

Prov. Santa, Lampanin, Mostacero et al. 1819 (F, GH).

6. Adiantum subvolubile Kuhn, Linnaea 36: 77.

1869. TYPE: Ecuador, (Azuay), Puente de

Han os. Spruce 5318 (holotype, B!; isotypes,

BM!, GH!, K!).

Stem rather slender, moderately long-creeping,

or rather compact. Leaves ca. 20-60 cm long, pet-

iole glabrous or slightly scaly at the base, rachis

glabrous. Lamina more or less elongate-ovate, 2-

or 3-pinnate in the center. Pinnae subsessile, the

basal pinnule undivided, overlaying the rachis,

basal pinnae often reduced or withered, ultimate

segments symmetrical or not, glabrous abaxially,

not articulate, the color of the apex of the stalk

passing into the base of the segment, sterile mar-

gins with each vein ending in a usually well-de-

veloped sinus. Indusia few, roundish or nearly re-

niform to lunate.

Usually growing on the coastal lomas, in crev-

ices of rocks or at the base of rocks, less often on

the ground, 50-3300 m, Piura to Cuzco and Mo-

quegua.

Ecuador and Peru.

On the lomas Atacongo and Amancaes, Dept.

Lima, a few collections vary toward Adiantum

concinnum in having some leaves with two ulti-

mate segments on the basal pinnule of the basal

pinnae: Cerrate 2721, Soukup 3136, Coronado 13

(GH). Considering the distance from A. concinnum

in northern Peru, these are regarded as variants

within A. subvolubile.

Piura: Olmos to Pucara, Gentry et al. 22630 (F, MO,

us). Canchaque, Ferreyra 3095 (USM). Cajamarca: Sum-

mit of Cerro Prieto, Haught 280 (us). La Libertad: Lo-

mas de Viru, Coronado 283 (GH, uc). Ancash: Lomas

de Mongon, Coronado 303 (GH, uc). Huanuco: Gorge of

Rio Chinchao, Tryon & Tryon 5315 (BM, F). Lima: Loma

de Atacongo, Pennell 14762 (F, GH, NY, PH), Ferreyra

2440 (BM, GH, USM). Loma de Amancaes, Coronado 13

(GH, MO, uc, us), Tryon & Tryon 5214 (BM, F, MO, u, us,

USM). Cuzco: Prov. Urubamba, quebrada de Pamahuan-

ca, Vargas 16017 (GH). Arequipa: Molendo, Johnston

3573 (GH, us). Lomas de Capac, Coronado 38 (uc, us).

Moquegua: Ilo, D. Stafford 931 (BM).

7. Adiantum henslovianum Hooker f., Trans. Linn.

Soc. London 20: 169. 1847. LECTOTYPE

(designated by Porter, J. Linn. Soc., Bot. 81:

95. 1980): Ecuador, Galapagos Islands,

Charles Island, Sept. 1835, Darwin (CGE). IS-

OLECTOTYPE: Herb. Hooker, K!; photo, GH.

Adiantum sessilifolium Hooker, Sp. fil. 2: 44. 1851.

TYPE: Peru, Chachapoyas, Mathews 1855 (ho-

lotype, K!; photos, GH, us).

Adiantum laetum Kuhn, Linnaea 36: 76. 1 869. TYPE:

Peru, (Amazonas), Chachapoyas, Mathews 3295

(holotype, B; isotype, K!; photos, GH, us of K).

Adiantum henslovianum var. macrosorum Hieron.,

Hedwigia 48: 238. 1909. SYNTYPES: Peru,

(Amazonas), near Leimabamba, Stiibel 1931 , B!,

(Cajamarca), near Celendin, Stiibel 1046, B!.

Stem rather small, very short-creeping. Leaves

ca. 1 5-90 cm long, petiole glabrate to more or less

persistently short-pubescent, the rachis puberulent

to short-pubescent. Lamina ovate to elongate-del-

toid, 2-pinnate. Pinnae with the pinna-rachis

whitish-puberulent to short-pubescent, ultimate

segments whitish-pubescent abaxially, or slightly

so, sterile ones moderately lobed, those nearest the

acute to broadly rounded terminal segment greatly

reduced or not, the basal pinnule of at least the

upper pinnae overlaying the rachis. Indusia few,

roundish, reniform to lunate, borne on the upper

and distal sides.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

59

Rocky slopes and Inca walls, 1400-2700 m,

Lambayeque to Cuzco.

Venezuela to Ecuador and Peru.

Adiantum sessilifolium has been separated from

A. henslovianum primarily on the basis of its pu-

bescent pinna-rachises while in the latter species

these are puberulent. However, there is intergra-

dation in this character and the distinction of the

species cannot be maintained.

Lambayeque: 31 km from Olmos on road to Jaen,

Correll & Smith P807 (GH). Cajamarca: W of Balsas,

Osgood & Anderson 67 (F), Osgood 67 (us). 40 km from

Cajamarca on road to Chilete, Correll & Smith P842

(GH). La Libertad: Prov. Pataz, between Alpamarca and

Retamas, Lopez & Sagdstegui 3634 (GH). Huanuco: Muna,

Bryan 428 (F, GH). Cuzco: Cuzco, 1854, Lobb (BM).

8. Adiantum lobatum Presl, Reliq. haenk. 1: 62,

t. 10, f. 4. TYPE: Ecuador, (Guayas), Gua-

yaquil, 1790, Haenke (holotype, PR; photo,

GH).

Stem rather slender, short-creeping. Leaves to

60 cm long, petiole atropurpureous, glabrous.

Lamina deltoid to long-triangular, 2-3-pinnate,

rachis and pinna-rachis whitish-puberulent, es-

pecially on the adaxial side. Pinnae with the ul-

timate segments glabrous, mostly oblong to tra-

peziform, the basal ones more or less flabellate,

entire to sometimes rather strongly incised, stalks

very short to usually short, the dark color entering

the base of the segment, sterile vein tips end be-

tween indistinct marginal crenulations. Indusia or-

bicular to suborbicular or a few sublunate, borne

on the upper and distal sides.

Brushy and rocky hillsides, 150-1400 m,

Tumbes and Lambayeque.

Ecuador and Peru.

Adiantum lobatum is a rare species, perhaps re-

lated to A. patens or A. henslovianum. The present

limited material makes any assessment of it un-

certain.

Tumbes: El Cancho, Coronado 227 (uc). Lambayeque:

Prov. Lambayeque, Dist. Olmos, Vdsquez et al. 4952

(USM). 27 km from Olmos on road to Jaen, Correll &

Smith P80I (GH, LL, us). 31 km from Olmos on road to

Jaen, Correll & Smith P808 (GH, LL).

9. Adiantum imbricatum Tryon, Amer. Fern J. 47:

142, t. 15. 1957. TYPE: Peru, Cuzco, La

Tranca, Sues 1377 (holotype, us!; isotypes,

cuz!, F!, GH!).

Stem not seen. Leaves ca. 1 2-30 cm long, petiole

brownish short-pubescent or glabrate, rachis

brownish short-pubescent. Lamina long-oblong to

elongate-triangular, 2-pinnate, basal pinnae

stalked, those above short-stalked, the innermost

pinnule overlaying the rachis. Pinna axes brown-

ish short-pubescent, ultimate segments broadly

cuneate-flabellate to suborbicular, rather to quite

symmetrical, mostly imbricate, pubescent abaxi-

ally, subarticulate at the junction with the stalk,

the apical portion of the stalk slightly enlarged, its

color abruptly distinct from that of the segment

base, sterile margins not seen. Indusia several to

numerous, roundish to orbicular-reniform, often

closely spaced.

Rather rocky places, 1600-2000 m, Cuzco.

Endemic to Peru.

This is a distinctive species, adequately distin-

guished from others in Peru by the characters pro-

vided in the key.

Cuzco: Puente de Collpani, Biies 1303 (us). La Tranca,

Rio Mapillo, Bites 1305 (us).

10. Adiantum orbignyanum Kuhn, Linnaea 36: 78.

1869. LECTOTYPE (designated by Tryon,

Contr. Gray Herb. 194: 175. 1964): Bolivia,

Mandon 52, B!; photo, GH. LECTOPARA-

TYPES: Bolivia, La Laguna, D'Orbigny 412,

B!; Bolivia, Mandon 1577, B!; Peru, Poeppig,

not seen. ISOLECTOPARATYPE: Bolivia,

Mandon 1577, GH!.

Stem slender, long-creeping, often branched, or

more compact. Leaves ca. 1 5-40 cm long, petiole

glabrous or slightly scaly at the base, rachis gla-

brous. Lamina lanceolate to narrowly ovate-del-

toid, 2-(rarely 3-)pinnate. Pinnae stalked, ultimate

segments cuneate-flabellate to suborbicular, rather

symmetrical or not, glabrous abaxially, articulate

at the junction with the stalk, the apical portion

of the stalk enlarged, its color abruptly distinct

from that of the segment base, the segment decid-

uous with a sharp clean break, sterile margins with

each vein ending in a usually well-developed sinus.

Indusia few to several, roundish to orbicular-re-

niform.

60

FIELDIANA: BOTANY

Most commonly in open, rocky places, 2 1 00-

4000 m, Cajamarca to Cuzco.

Peru, Bolivia, and northwestern Argentina.

The articulate segments of this species are best

observed in older leaves that are fully developed,

and especially in leaves in which some of the seg-

ments have fallen off. In these the stalks with a

slightly enlarged apex may easily be observed. In

some cases the stem is rather long-creeping, bear-

ing clusters of leaves at intervals, as often in A.

poiretii and A. chilense.

Cajamarca: San Miguel, July 30, 1952, Dias (USM).

Huanuco: Pachachupam, Feb., 1 940, Ridoult (USM). Jun-

in: Below Palca, Correll & Smith P766 (GH, us). Apu-

rimac: Cachora to Huillcayocc, Vargas 9104 (uc). Cuzco:

Yucay, Herrera 714 (us), 718 (us), 7/95 (us), Coronado

146 (GH, uc). San Sebastian, Pennell 13621 (F, GH, NY,

PH). 10 km SE of Cuzco, Solomon 3010 (F, MO).

11. Adiantum digitatum Hooker, Sp. til. 2: 38.

1851 (Presl, Tent, pterid. 159. 1836, nomen

nudum). TYPE: Brazil, Sello (holotype, K!;

photo, GH).

Adiantum speciosum Hooker, Sp. fil. 2: 45, t. 85C.

1851. LECTOTYPE (Designated by Tryon, Contr.

Gray Herb. 194: 178. 1 964): Ecuador, Sasaranga,

Seemann 953, K.!; photo, GH.

Adiantum palmatum Moore, Gard. Chron. 1877: 40,

/ 5. TYPE: Peru, Roezl ex hort. B. S. Williams

(holotype, K!; photo, GH).

Stem moderately stout, long-creeping. Leaves

ca. 45-1 50 cm long and the long ones subscandent,

petiole tawny short-pubescent or glabrate, some-

times slightly scaly toward the base, rachis similar

but not scaly. Lamina deltoid, 3-4-pinnate or less

often 2-pinnate. Pinnae stalked, the axes tawny

short-pubescent or rarely glabrate, ultimate seg-

ments usually more or less suborbicular, deeply

cleft into about 4-7 spreading lobes, quite sym-

metrical or not, usually flabellately veined, pu-

bescent abaxially, rarely glabrate, the color of the

apical portion of the stalk abruptly distinct from

that of the segment base or not, sterile margins

with each vein ending in a tooth. Indusia few to

several, oblong, straight or nearly so.

In rocky places or at the base of rocks, in woods,

or on brushy hillsides, 100-4000 m, Tumbes to

Puno.

Ecuador to Argentina, Uruguay, and Brazil.

Adiantum digitatum is a distinctive species with

its flabellate venation, deeply cleft segments, and

frequently much elongated leaf that is up to 1.5 m

long. Most of the axes, especially those of the pin-

nules, are at right angles to the axis that bears

them. In the northern half of Peru this species is

frequent on lomas and a characteristic part of their

flora.

Tumbes: Between Cancho and Cotrina, Coronado 217

(GH, uc). Piura: W of Canchaque, Stork 11396 (GH, us).

Canchaque, Ferreyra 10894 (GH, USM). Lambayeque: Ol-

mos to Jaen, Correll & Smith P829 (GH). Cajamarca: El

Puquio, Guzmango, Sagdstegui 3915 (GH). La Libertad:

Lomas Campana, Coronado 280 (GH, uc). Lomas de

Viru, Coronado 287 (GH). Ancash: Huasta, Cerrate 2478

(USM). Prov. Santa, arriba de Lampanin, Mostacero et

al. 18 18 (F, GH). Huanuco: Muna, Macbride 3919(F, GH,

us) Lima: Lomas de Chancayllo, Hutchison 6296 (F, GH,

us). Lomas de Patavilca, Coronado 306 (GH, uc). Lomas

de Chancay, Ferreyra 8700 (GH, USM). Junin: 10 km

below Palca, Walden 3 (BM), 6 (BM). Cuzco: Vilcambam-

ba, Vargas 4018 (uc, us). Chincheros, Davis et al. 1802

(F). Puno: Ollachea, Vargas 6917 (MO, uc, us).

12. Adiantum patens Willd., Sp. pi. ed. 4, 5: 439.

1810. TYPE: Venezuela, (Dist. Federal), Ca-

racas, Bredemeyer (holotype, B!, Herb. Willd.

20078; photo, GH; isotype, w!).

Stem rather small, short-creeping. Leaves ca.

20-45 cm long, petiole glabrous or slightly puber-

iilent, rachises similar. Lamina deltoid-ovate to

nearly circular, with 2 equal, strongly recurved

rachises, these bearing 1 -pinnate branches on only

1 side, or with the rachises more or less unequal

and one bearing a well-developed terminal pinna.

Penultimate segments with the axes glabrous to

puberulent, rarely short-pubescent, ultimate seg-

ments glabrous abaxially, or rarely whitish short-

pubescent, sterile ones moderately lobed, or evenly

serrate, those nearest the broadly rounded termi-

nal segment reduced or not, the basal ultimate

segments hardly or not overlaying the adjacent

axis. Indusia few, roundish, lunate or reniform,

borne on the upper and distal sides.

In woods and on shaded rocks, 150-2050 m,

Tumbes south to Cuzco.

Mexico and Central America; Venezuela and

Colombia south to Bolivia.

The architecture of the lamina is distinctive in

this species and similar to that of Adiantum pe-

datum of North America.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

61

Tumbes: Between Cancho and Cotrina, Coronado 233

(GH). I 'i ura: Talara, H aught 92 (us). Huancabamba, Fer-

reyra 10877 (GH, USM). Lambayeque: Prov. Lambayeque,

Ldpez el al. 4045 (GH, HUT). Olmos to Jaen, Correll &

Smith P806 (GH, us ). Amazonas: Prov. Bagua, entre

Montenegro y Lumba, Lopez et al. 4228, in part (HUT).

Cuzco: Machu Picchu, Vargas 3352 (MO, us).

13. Adiantum rui/ianum KJotzsch, Linnaea 18:

551. 1845. TYPE: Peru, Ruiz 26 (holotype,

B!).

Adiantum veitchianum Moore, Gard. Chron. 1868:

1090. TYPE: Peru, (Huanuco), Mufia, Pearce

(holotype, K.!; photo, GH).

Adiantum steerei Harr., J. Linn. Soc., Bot. 16: 34.

1877. LECTOTYPE (designated by Tryon, Contr.

Gray Herb. 194: 176. 1964): Peru, (Amazonas),

Poma Cocha, Steere , K!. ISOLECTOTYPES: GH!,

MO!, us!.

Adiantum microsorium C. Chr., Index fil. 30. 1905,

based on A. veitchianum but an unnecessary new

name as Ballard (Kew Bull. 1954[4]: 560) has

pointed out.

Stem slender, long-creeping. Leaves 1 5-45 cm

long, petiole glabrous or slightly scaly at the base,

rachis glabrous. Lamina oblong or oblong-linear

or oblong- tapering, 1-2-pinnate. Pinnae mostly

long-stalked, ultimate segments broadly cuneate-

flabellate to suborbicular, usually rather or quite

symmetrical, glabrous, not articulate, the color of

the apex of the stalk passing into the base of the

segment, sterile margins with each vein ending in

a more or less well-developed sinus. Indusia sev-

eral to numerous, orbicular or nearly so, usually

closely spaced.

Damp or shaded rocky places, 1600-2900 m,

Amazonas, Huanuco, Pasco, and Junin.

Endemic to Peru.

Most of the material seen is 1 -pinnate with rath-

er large, flabellate pinnae on long, slender stalks.

Rarely, the lamina may be 2-pinnate and then the

segments may be smaller, more elongate and with

shorter stalks.

Amazonas: Prov. Chachapoyas, Chachapoyas-Men-

do/a. Lopez & Aldave 8523 (HUT). Prov. Chachapoyas,

Calla Calla, Hutchison & Wright 4798 (F, GH), Wurdack

1182 (F, GH, us). Prov. Chachapoyas, encima de Lei-

mabamba. Ferreyra 15598 (uc, us). Huanuco: Muna,

Bryan 515 (F, us), Macbride 3942 (F, us). Woytkowski

5223 (GH, MO). Pasco: Prov. Oxapampa, Palcazu, van

der Werffet al. 8378 (MO). Dos de Mayo (as Junin), Killip

& Smith 25846 (BM. F, GH. NY, us). Junin: Huacapistana,

Tryon & Tryon 5434 (F).

1 4. Adiantum deflectens Mart., Icon. pi. crypt. 94.

1834. TYPE: Brazil, Prov. Para, Santarem,

Martius (not located).

Stem small, short-creeping to nearly erect. Leaves

ca. 1 5-30 cm long, petiole very slender, glabrous,

rachis similar. Lamina 1 -pinnate, often of two

kinds, one oblong to very elongate-triangular, with

a terminal segment, the pinnae cuneate-flabellate

to broadly so, asymmetrical, the other kind linear,

with a long, naked, rooting rachis tip, the pinnae

cuneate-flabellate to cuneate-ovate, nearly or quite

symmetrical. Pinnae stalked, entire or more or less

cleft into 2-7 close lobes, glabrous, the segment

articulate at the junction with the stalk, the apical

portion of the stalk enlarged, abruptly distinct in

color from the base of the segment, sterile margins

finely and sharply serrate, with each vein ending

in a tooth. Indusia few, oblong to linear, nearly

straight to long-arcuate.

In damp, shady places, 820-1700 m, Cuzco.

Mexico and Central America; Guianas to Co-

lombia, south to Peru, Paraguay, and Brazil.

This is one of the most distinctive of the Pe-

ruvian species. The 1 -pinnate lamina in some

leaves ends in a terminal segment, while in other

leaves the lamina has a long, naked, rooting tip.

The correct name for this species depends upon a

careful study of its group throughout tropical

America and the types of the several names in-

volved. Among these are Adiantum delicatulum

Mart., A. dolabriforme Hooker, A. rhizophyllum

Mart., A. filiforme Hooker, A. phillipense L., A.

lunulatum Burm., A. flagellum Fee, and A. subar-

istatum Fee.

Cuzco: Prov. La Convencion, Hacienda Sahuayaco,

Bues 822 (us), Vargas 1658 (GH), 7660 (GH). Puente de

Collpani, Bites 1298 (us). Machu Picchu to Quillabamba,

Mexia 8088a (F, GH, MO, uc, us). Yanayco Grande, Bues

1035 (us). Prov. La Convencion, Quellouno, Vargas

13553 (GH).

1 5. Adiantum pectinatum Ettingsh., Farnkr. Jetzw.

85, /. 45, figs. 14-16. 1865. TYPE: Brazil,

Goyaz, Pohl 1481 (holotype, w!; isotypes, B!

(s.n.), GH!).

Stem very stout, moderately creeping. Leaves

very large, up to 2 m long or more, petiole very

stout (to nearly 1 cm in diameter at the base),

deciduously pubescent-scaly, the rachis rather per-

62

FIELDIANA: BOTANY

sistently so. Lamina broadly deltoid-ovate to del-

toid-circular, 4-5-(rarely 6-)pinnate, the basal pin-

nae largest. Pinnae widely alternate, pinna-rachis

pubescent-scaly, ultimate segments glabrate or

slightly scaly abaxially, sterile ones deeply incised-

lobed, those nearest the rather obtuse to acute ter-

minal segment of the penultimate segments mod-

erately to greatly reduced. Indusia several, orbi-

cular or mostly so, borne on the upper and

sometimes also a few on the distal side.

In woods and dense forests, 700-2000 m, San

Martin to Cuzco and Madre de Dios.

Costa Rica to Argentina and Brazil.

This species is characterized by the ultimate seg-

ments that are incised on the upper edge, and the

complex lamina that is commonly 4-5-pinnate at

the base. It is the largest of the adiantums, some

leaves reaching a length of 2 m or more.

San Martin: Moyabamba to Huallaga, Stubel 1091 (B).

Tarapoto, Spruce 4781 (K). Huanuco: Pampayacu, Ka-

nehira 130 (GH). Junin: La Merced, Soukup 1082 (F),

Tryon & Tryon 5437 (BM, F, MO, u, us, USM). Cuzco:

Torontoy. Herrera 1297 (us). Prov. La Convention, ba-

jios de Rosalina, Vargas 12289 (GH). Madre de Dios:

Parque National de Manu, M. S. Foster P-84-58 (MO).

16. Adiantum tomcntosum KJotzsch, Linnaea 18:

553. 1845. TYPE: "British Guiana", Schom-

burgk 1202, (holotype, B!; frag., GH!, us!).

Stem moderately stout, rather long-creeping.

Leaves to ca. 1.25 m long, more or less deciduously

puberulent, the rachis persistently so. Lamina

ovate-deltoid or broadly so to broadly ovate-ob-

long, 2-pinnate. Pinnae with the pinna-rachis pu-

berulent to pubescent only on the adaxial side or

also sometimes on the lateral sides, glabrous abax-

ially, the terminal pinna reduced at the base, pin-

nules glabrous, usually imbricate, sterile ones rath-

er coarsely but evenly serrate, those nearest the

usually acute terminal segment moderately re-

duced. Indusia numerous, roundish to short-ob-

long, borne on the upper and distal sides, some-

times a few also on the inner side.

In dense forests, 1 00-650 m, Amazonas to Madre

de Dios.

Guianas to Colombia, south to Peru and Brazil.

This is one of the more distinctive species. The

pinna-rachises are puberulent or pubescent, but

not on the abaxial side, and the ultimate segments

are obtuse and usually closely imbricate. In Peru

it is most common in the Amazon basin.

Amazonas: Prov. Bagua, entre Montenegro y Lumba,

Ldpez et al. 4228 in part (HUT). Prov. Bagua, near Cas-

cadas de Mayasi, Wurdack 1940 (GH, us). San Martin:

Palo Blanco, J. Schunke V. 5629 (F, us). Loreto: Iquitos,

Killip & Smith 27415 (NY, us). Prov. Maynas, Rio Na-

nay, Gentry et al. 22368 (F, MO, uc). Prov. Maynas, Rio

Momon, Sounders 1366 (F, GH). Huanuco: Tingo Maria,

Tryon & Tryon 5295 (BM, F, GH, MO, NY, u, uc, us, USM).

Pasco: Puerto Bermudez (as Junin), Killip & Smith 26549

(GH, NY, us). Madre de Dios: Prov. Tambopata, Tam-

bopata Nature Reserve, Barbour 5181 (F, MO).

17. Adiantum urophyllum Hooker, Sp. Ml. 2: 24,

/. 84B. 1858. SYNTYPES: Colombia, Island

of Gorgona, Barclay, Hinds; Ecuador, Sal-

ango, Seemann; all presumably at K.

Adiantum pilosum Baker, Ann. Bot. (London) 5: 207.

1 89 1 , not Fee, 1 852. TYPE: Colombia, Kalbreyer

956 (holotype, K!; photo, GH).

Adiantum kalbreyeriC. Chr., Index fil. 28. 1905, nom.

nov. for Adiantum pilosum Baker and with the

same type.

Adiantum ceciliae Alston, Lilloa 30: 109. /. 4-5. 1960.

TYPE: Peru, Dept. Junin, south bank of Rio Per-

ene, within 15 miles of confluence of Rio Chan-

chamayo and Rio Paucartambo, Gascoyne-Cecil

50 (holotype, BM; paratype, same data and col-

lector, 47 BM).

Stem moderately stout, moderately long-creep-

ing. Leaves ca. 60-90 cm long, petiole deciduously

puberulent, the rachis persistently so. Lamina

ovate, 2-pinnate. Pinnae few, pinna-rachis puber-

ulent, pinnules glabrate, nearly subacute, mostly

spaced, sterile ones coarsely and unevenly serrate,

those nearest the long-acuminate terminal seg-

ment greatly reduced. Indusia several to numer-

ous, roundish, borne on the upper and distal sides.

One collection seen from a forested slope, 1 200

m, Huanuco.

Costa Rica to Peru.

The pinna-rachises that are puberulent on all

sides make this a distinctive species. If the pub-

erulence is eroded from the abaxial side, then the

subacute and mostly spaced pinnules will separate

it from Adiantum tomentosum.

Huanuco: Prov. Huanuco, Dist. Churubamba, Ha-

cienda Mercedes, Mexia 8175 (BM, F, GH, MO, uc, us).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

63

18. Adiantum macrocladum KJotzsch, Linnaea 18:

554. 1845. TYPE: Peru, Poeppig 1147, (ho-

lotype, B!; photo, BM; isotypes, GH!, LE!, us!).

Adiantum mexiae Copel., Univ. Calif. Publ. Bot. 19:

303. 1941. TYPE: Peru, Huanuco, Mexia 8292

. (holotype, uc; isotypes, GH!, us!).

Stem moderately stout, rather long-creeping.

Leaves up to 1 m long, petiole glabrous, the rachis

similar. Lamina more or less circular to broadly

ovate, 2-pinnate above the basal pinnae, where it

is 3-pinnate. Pinnae with the pinna-rachis gla-

brous, pinnules glabrous, often imbricate or nearly

so, sterile ones mostly evenly serrate, those nearest

the long-acuminate terminal segment greatly re-

duced. Indusia numerous, mostly orbicular or

nearly so, borne on the upper side.

In dense forests, 300-1500 m, San Martin to

Madre de Dios.

Endemic to Peru.

The wholly glabrous pinna-rachises separate this

species from Adiantum tomentosum which also

has obtuse and usually closely imbricate pinnules.

San Martin: Prov. Mariscal Caceres, 2 km abajo de

Tocache Nuevo, J. Schunke V. 3435 (GH, MO, us). Huan-

uco: Tingo Maria, Stork & Morton 9531 (F, uc, us). Pas-

co: Prov. Oxapampa, W side of Cordillera de San Matias,

D. Smith 2013 (F). Cahuapanas on Rio Pichis, (as Junin),

Killip & Smith 26790 (NY, us). Ucayali: Prov. Coronel

Portillo, Bosque Nacional von Humboldt, J. Schunke V.

10404 (F, MO). Madre de Dios: Cocha Cashu station, Rio

Manu, Foster 2565 (GH), Foster el al. 6905 (F).

19. Adiantum tetraphyllum Willd., Sp. pi. ed. 4,

5: 441. 1810. LECTOTYPE (designated by

Lellinger, Proc. Biol. Soc. Wash. 89: 704.

1977): Venezuela, (Sucre), Caripe, Humboldt,

B!, Herb. Willd. 20082-2; photo, GH. LEC-

TOPARATYPE: America, Vahl, B!, Herb.

Willd. 20082-1; photo, GH.

Stem long-creeping and slender to short-creep-

ing and moderately stout, or compact. Leaves ca.

40-150 cm long, petiole persistently to decidu-

ously pubescent-scaly, the rachis densely and per-

sistently pubescent-scaly to moderately so. Lam-

ina deltoid-ovate, to ovate-oblong, 2-pinnate.

Pinnae with the pinna-rachis densely to moder-

ately pubescent-scaly, pinnules green, often shin-

ing abaxially, glabrous, somewhat scaly or pubes-

cent abaxially, the sterile margins unevenly and

coarsely serrate. Indusia few to many, roundish to

long-oblong, borne on the upper and distal sides,

or sometimes also on the inner side.

In forests and on forested slopes, sometimes in

seasonally inundated forests, 100-1 500 m, Tumbes

to Madre de Dios.

Tropical America.

Adiantum tetraphyllum is treated as a variable

species in Peru. It may be distinguished from re-

lated species by its 2-pinnate lamina with a ter-

minal 1 -pinnate pinna about the same size as the

adjacent lateral ones, the pubescent-scaly rachis

and pinna-rachises, the ultimate segments that are

green and often shining abaxially, and most of

them more than twice as long as broad with the

sterile margins unevenly and coarsely serrate.

In Tryon ( 1 964) several species were recognized

in the A. tetraphyllum complex, but the material

of these is now regarded as part of the variation

of A. tetraphyllum. Some of these species may be

distinctive in other regions of tropical America (for

example, A. villosurri) and the Peruvian specimens

were erroneously associated with their name. In

other cases they evidently represent one extreme

or another of the variations within A. tetraphyl-

lum. These species include Adiantum cayennense

Klotzsch, A. fructuosum Sprengel, and A. villos-

issimum Kuhn. Another species of this complex,

Adiantum fuliginosum Fee (A. hirtum Splitg. 1 840,

not (Sw.) Poiret, 1810), is sometimes credited to

Peru.

The variation in Adiantum tetraphyllum in Peru

includes the following characters: (a) stem long-

creeping and slender to short-creeping and mod-

erately stout, or compact; (b) lamina axes densely

pubescent-scaly to moderately so; (c) pinnae acute

with the distal segments about half as long as the

longest on the pinna to long-acuminate with the

distal segments much reduced; (d) ultimate seg-

ments glabrous, somewhat scaly, or pubescent

abaxially; (e) apex of the ultimate segments obtuse

to acute; and (/) indusia few to many on a fertile

segment, roundish to long-oblong.

Tumbes: Prov. Zarumilla, Dist. Matapola, Campo

Verde, J. Schunke V. 2363 (F, GH, us). Between Cancho

and Cotrina, Coronado 223 (GH, uc). Amazonas: Prov.

Bagua, above Cascadas de Mayasi. Wurdack 1972 (F,

GH, us). San Martin: Tarapoto, Spruce 4665 (K). Prov.

Mariscal Caceres, Tocache Nuevo, J. Schunke V. 3876

(F). Loreto: Rio Itaya. near Iquitos, Tryon & Tryon 5197

(BM, F, GH, MO, USM). Rio Nanay, near Iquitos, Tryon &

Tryon 5177 (BM, F, GH, MO, u, us, USM). Huanuco: Tingo

Maria, Tryon & Tryon 5294 (BM, F, GH, MO, u, uc, us,

USM). Pasco: Prov. Oxapampa, cerca de Puerto Ber-

mudez, Leon et al. 297 (USM). Prov. Oxapampa, Cor-

64

FIELDIANA: BOTANY

dillera de San Matias, D. Smith 2013A (MO). Junin: La

Merced, Tryon & Tryon 5438 (BM, F, GH, MO, u, uc, us,

USM). Ucayali: Prov. Coronel Portillo: Parque Nacionai

von Humboldt to Puerto Inca, Croat 51054 (MO). Ay-

acucho: Between Huanta and Rio Apurimac, Killip &

Smith 22623 (F, NY, us). Cuzco: Prov. La Convencion,

below Puerto Capiro, Davis el al. 1312 (F, GH). Prov.

Quispichanci, entre Machichi y Inambari, Vargas 16458

(GH). M ad re de Dios: Prov. Tambopata, Tambopata Na-

ture Reserve, Harbour 5306 (MO). Prov. Tambopata, Rio

Piedras, Vargas 18626 (GH).

20. Adiantum terminatum Miq., Het. Instit. Versl.

Meded. Ned. Instit. Wet. 1842: 3. 1843. (Cit-

ed by C. Chr., Index fil., as Diar. Inst. Reg.

Bat.) TYPE: Surinam, Bergendall, Focke (ho-

lotype, u!).

Stem rather slender, short- to rather long-creep-

ing. Leaves ca. 20—45 cm long, petiole deciduously

pubescent-scaly, the rachis persistently so. Lamina

broadly deltoid to oblong or oblong-ovate, 2-pin-

nate. Pinnae with the pinna-rachis pubescent-sca-

ly, pinnules dull green to faintly glaucous, with

numerous simple trichomes abaxially, sterile ones

mostly coarsely and evenly serrate, those nearest

the acuminate terminal segment reduced. Indusia

several to numerous, very short-oblong, borne on

the upper and often also on the distal and inner

sides.

Caripe, Humboldt & Bonpland (holotype,

B!, Herb. Willd. 20088; the two sterile leaves,

B!; photos, BM, GH).

Stem slender, very long-creeping. Leaves ca. 35-

65 cm long, petiole deciduously pubescent-scaly,

the rachis persistently so. Lamina deltoid-ovate to

oblong-ovate, 2-pinnate. Pinnae with the pinna-

rachis pubescent-scaly, pinnules coriaceous, not

glaucous, glabrate or slightly scaly abaxially, sterile

ones evenly serrate, those nearest the usually ob-

tuse terminal segment of the pinna not much re-

duced. Indusia few to several, roundish to oblong

or sublunate, borne on the strongly recurved mar-

gin on the upper and distal, rarely the inner, sides.

In forests and in open habitats among grasses,

750-1500 m, Piura to Cuzco.

Mexico; West Indies; tropical South America.

This species has coriaceous and usually small,

oblong pinnules. The sterile margins are usually

finely and evenly serrate.

Piura: Cerro de Chiris, Panojal, Bues 17 11 (us). San

Martin: Zepelacio, near Moyobamba, Klug 3462 (F, GH,

MO, NY, us). Loreto: Prov. Requena, Rio Ucayali, En-

carnacion 25032, in part (MO). Huanuco: Pampayacu,

Tump (USM). Cuzco: Prov. La Convencion, Vargas 17279

(GH).

In primary, or less often in secondary forests,

100-800 m, San Martin and Loreto to Puno.

Southern Mexico and Guatemala; Trinidad;

tropical South America.

This species is characterized by usually many

indusia on the fertile segment, the gradually re-

duced pinna-apex, and the long, simple trichomes

on the segments abaxially.

San Martin: Prov. Mariscal Caceres, Palo Blanco, J.

Schunke V. 57 15 (F, us). Loreto: Prov. Maynas, Rio

Momon, Sounders 1370 (F, GH). Near Iquitos, Klug 1270

(NY, us). Nauta, Ferreyra 5134 (USM). Huanuco: Tingo

Maria, Tryon & Tryon 5327 (BM, F, GH, MO, u, uc, us,

USM). Pasco: Prov. Oxapampa, Rio Palcazu, Leon 705

(F). Prov. Oxapampa, Paujil, Leon 303a (USM). Junin:

Puente Perene, Coronado 262 (GH, uc). Ucayali: Bosque

Nacionai von Humboldt, Narita 8 (USM). Cuzco: Prov.

Paucartambo, Rio Tono, Wachter et al. 218 (F). Madre

de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al.

10699 (F). Prov. Tambopata, Tambopata Nature Re-

serve, Barbour 5192 (MO). Puno: San Gaban, Lechler

2319, 2319a (B).

21. Adiantum serratodentatum Willd., Sp. pi. ed.

4, 5: 445. 1810. TYPE: Venezuela (Sucre),

22. Adiantum pulverulentum L., Sp. pi. 2: 1096.

1753. LECTOTYPE (designated by Proctor,

Flora Lesser Antilles 2: 185. 1977): Plumier,

Descr. pi. Amer. /. 47. 1693. Figure 14d.

Stem moderately stout, rather short-creeping.

Leaves ca. 30-90 cm long, petiole more or less

persistently pubescent-scaly, the rachis usually

persistently so. Lamina short-oblong to oblong-

ovate, 2-pinnate. Pinnae with the pinna-rachis pu-

bescent-scaly, pinnules glabrate to somewhat scaly

abaxially, sterile ones coarsely serrate, those near-

est the acute to long-acuminate terminal segment

greatly reduced. Indusium long, 1 (rarely 2) borne

on the upper side, rarely a shorter one also on the

distal side.

In dense or partially open forests or in secondary

growth, 100-1800 m, Loreto south to Cuzco and

Madre de Dios.

Tropical America.

The usually single, long indusium is sufficient

to characterize this 2-pinnate species. It is espe-

cially common in the Amazon basin.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

65

Loreto: Mouth of Rio Santiago, Mexia 6133a (GH, NY,

uc, us). Rio Itaya, Killip & Smith 29431 (F, GH, NY, us).

Near Iquitos, Tryon & Tryon 5194 (BM, F, GH, MO, u,

USM). Huanuco: Prov. Pachitea, Dist. Puerto Inca, D.

Smith 1261 (MO). Pasco: Prov. Oxapampa, Villa Rica-

Yesu, Smith et al. 5461 (GH). Junin: Polonia, Satipo,

1940, Ridoutt (us). Cuzco: Prov. La Convention, Rio

Sam bray, Mexia 8066 (F, GH, MO, uc, us). Madre de

Dios: Prov. Manu. Cerro de Pantiacolla, Foster et al.

10717 (F).

23. Adiantum latifolium Lam., Encycl. 1:43. 1783.

Type: uncertain, there is no specimen in Herb.

Lamarck; Brazil, Commerson (Herb. Jussieu

no. 1408) P!; photos, GH, us, is evidently a

hybrid. The name is applied in its traditional

sense.

Adiantum humile Kunze, Linnaea 9: 80. 1834. TYPE:

Peru, (San Martin), Huallaga, Mission Tocache,

June, 1830, Poeppig (holotype, LZ destroyed), ex

char., the description suggests that the name is a

synonym of A. latifolium.

Adiantum killipii Maxon & Weath., Amer. J. Bot. 19:

166. 1932. TYPE: Panama, Canal Zone, Ancon

Hill, Killip 2752 (holotype, us!).

Stem slender, very long-creeping to moderately

stout and short-creeping. Leaves ca. 30-70 cm long,

petiole deciduously pubescent-scaly, the rachis

persistently so. Lamina deltoid to ovate-deltoid or

ovate-oblong, 2-pinnate. Pinnae with the pinna-

rachis pubescent-scaly, pinnules herbaceous, glau-

cous abaxially, glabrate, or with simple trichomes

abaxially, with the sterile ones evenly serrate, those

nearest the usually acute terminal segment of the

pinna usually not much reduced. Indusia several,

usually oblong, and on the lower as well as the

upper side.

In woods and in dense forests, 100-1000 m,

Amazonas and San Martin to Cuzco and Madre

de Dios.

Tropical America.

This 2-pinnate species has the segments glau-

cous abaxially and the sterile margins finely and

evenly serrate. It is especially common in the Am-

azon basin.

Amazonas: Prov. de Bagua, above Cascadas de May-

asi, Wurdack 1899 (us). San Martin: Prov. Mariscal

Caceres, Tocache Nuevo, J. Schunke V. 3410 (F). Loreto:

Rio Putumayo, Peru-Colombia boundary, Klug 1631 (F,

GH, MO, NY, us). Dist. Iquitos, Fundo Indiana, Mexia

6389 (BM, F, GH, MO, NY, PH, uc, us). Rio Itaya, near

Iquitos, Tryon & Tryon 5170 (BM, F, MO, u, us, USM).

Huanuco: Prov. Pachitea, Bosque Nacional de Iparia, J.

Schunke V. 1139 (F, GH, us ). Junin: Near La Merced,

Killip & Smith 24003 (NY, us). Ucayali: Bosque Nacional

von Humboldt, Gentry et al. 36219 (F, MO). Ayacucho:

Rio Apurimac valley, near Kimpitiriki, Killip & Smith

22925 (GH, NY, us). Cuzco: Prov. Paucartambo, Kosni-

pata, Vargas 10202 (MO, uc). Madre de Dios: Prov. Tam-

bopata, Vargas 18577 (GH), 18630 (GH).

24. Adiantum petiolatum Desv., Ges. Naturf.

Freunde Berlin Mag. Neuesten Entdeck. Ges-

ammten Naturk. 5: 326. 1811. TYPE: uncer-

tain, not seen at p.

Adiantum kaulfussii Kunze, Linnaea 21: 221. 1848.

LECTOTYPE (designated by Tryon, Contr. Gray

Herb. 194: 149. 1964): Martinique, Sieber, Fl.

Martin. 371 (holotype, LZ destroyed; isotype, B!).

Stem slender, long-creeping to somewhat stout

and more compact. Leaves ca. 20—40 cm long,

petiole glabrate, rachis similar. Lamina more or

less oblong and 1 -pinnate or deltoid and 2-pin-

nate, with the pinnate pinnae few, entire pinnae

ovate-lanceolate to oblong, strongly asymmetrical

at the base. Pinnae and pinnules dull, usually glau-

cous abaxially, sterile margins evenly serrate. In-

dusia several, oblong-arcuate to long-arcuate or

nearly straight, more numerous on the upper than

on the distal side.

In forests, disturbed forests, secondary growth,

or on wet, shaded cliffs, 100-600 m, Amazonas to

Madre de Dios.

Tropical America.

One of the variations of Adiantum petiolatum

has a glabrous rachis. This may be distinguished

from other species by the 1 -pinnate lamina with

elongate pinnae with the sterile margins evenly

serrate.

Plants of Adiantum petiolatum with the lamina

2-pinnate may be, at least in part, hybrids with

Adiantum latifolium. Jermy and Walker (Bull. Brit.

Mus. (Nat. Hist), Bot. 13: 254. 1985) described

Adiantum x variopinnatum as a hybrid between

diploid and 1 -pinnate A. petiolatum and tetraploid

and 2-pinnate A. latifolium. Some 2-pinnate col-

lections may represent this hybrid, while others

may involve another 2-pinnate species, or simply

be a 2-pinnate variation of A. petiolatum. Speci-

mens that have the apical pinna much larger than

the few adjacent lateral ones are most likely to

represent hybrids.

Amazonas: Prov. Bagua, above Cascadas de Mayasi,

Wurdack 2055 (F, GH, MO, uc, us). San Martin: Prov.

66

FIELDIANA: BOTANY

Mariscal Caceres, Rio Chiote, J. Schunke V. 4307 (F,

GH, us). Loreto: Near Iquitos, Tryon & Tryon 5191 (BM,

F, GH, MO, u, uc, us, USM). Salinas, Rio Mazan, Schunke

377 (F, GH, NY, uc, us, USM). Huanuco: Prov. Pachitea,

Bosque Nacional de Iparia, J. Schunke V. 2965 (F, GH,

us). Pasco: Prov. Oxapampa, valle del Palcazu, Ledn

677 (F). Junin: Rio Satipo, 1 940, Ridoutt (USM). Ucayali:

Bosque Nacional von Humboldt, Gentry et al. 36382

(MO). Cuzco: Prov. Quispicanchi, Punkiri, Vargas 15412

(GH). Madre de Dios: Culpa, Rio Manu, A/. 5. Foster P-

84-36 (uc).

25. Adiantum obliquum Willd., Sp. pi. ed. 4, 5:

429. 1810. LECTOTYPE (designated by Lei-

linger, Mem. New York Bot. Card. 23: 13.

1972.): Venezuela, (Dist. Federal), Caracas,

Bredemeyer, B!, Herb. Willd. 20067-1; pho-

tos, BM, GH. LECTOPARATYPE: Portorico

(Puerto Rico), Ventenat, B!, Herb. Willd.

20067-2; photos, BM, GH.

Stem slender, long-creeping to compact, often

branched. Leaves ca. 20-60 cm long, petiole de-

ciduously or persistently pubescent-scaly, rachis

similar. Lamina broadly linear to elongate-trian-

gular and 1 -pinnate or deltoid to deltoid-ovate and

2-pinnate, with the pinnate pinnae few. Entire pin-

nae and pinnules oblong to long-triangular, strong-

ly asymmetrical at the base, green, usually shining

abaxially, sterile margins unevenly biserrate. In-

dusia several, short to long-oblong, straight or ar-

cuate, more numerous on the upper than on the

distal side.

In dense forests or on wooded slopes, 250-1 500

m, San Martin to Cuzco and Madre de Dios.

Tropical America.

Plants with 2-pinnate leaves may be within the

variation of Adiantum obliquum, or they may be,

at least in part, hybrids of A. obliquum with 2-

pinnate species. Those with a very large terminal

pinna are the most likely to be hybrids.

San Martin: Prov. Mariscal Caceres, Tocache Nuevo,

J. Schunke V. 7046 (F, MO). Loreto: Rio Itaya, near Iqui-

tos, Tryon & Tryon 5201 (BM, F, MO, u, us, USM). Huan-

uco: Tingo Maria, Tryon & Tryon 5253 (BM, F, MO, u,

us, USM). Pasco: Prov. Oxapampa, cerca a Puerto Ber-

mudez, Leon et al. 294 (GH, USM). Junin: Chanchamayo

valley, C. Schunke 101 (F), 763 (us). Cuzco: Prov. La

Convencion, Puerto Capiro, Rio Apurimac, Davis et al.

1294 (F, uc). Madre de Dios: Prov. Tambopata, Tam-

bopata Nature Reserve, Barbour 4763 (F, MO). Maldon-

ado, Lopez 4585 (GH).

26. Adiantum alarconianum Gaud., Voy. Bonite

Bot. t. 99. 1 846. TYPE: Ecuador, (Guayas),

Guayaquil, April, 1836, Gaudichaud (holo-

type, P!; isotype, B!; frag., us!; photo, us of FI).

Adiantum incisum Presl, Reliq. haenk. 1: 61, /. 10, /

3. 1825, not Forssk. 1775. LECTOTYPE (des-

ignated here): Ecuador, Guayaquil, Haenke 10,

PR or PRC?, not seen; ISOLECTOTYPES: BM!, w!.

LECTOPARATYPE: Haenke 9 (not seen).

Stem moderately stout, short-creeping, scales ir-

idescent. Leaves ca. 30-50 cm long, petiole gla-

brous or slightly pubescent-scaly on the adaxial

side, rachis similar but definitely pubescent-scaly

on the adaxial side, usually glabrous elsewhere.

Lamina linear to broadly linear and 1 -pinnate or

broadly ovate-oblong and 2-pinnate. Entire pin-

nae or pinnules oblong-falcate to long-triangular,

strongly asymmetrical at the base, alternate, gla-

brate abaxially, sessile or subsessile. Indusium sin-

gle, long, borne on the upper side.

In woods, ca. 650 m, Tumbes.

Ecuador and adjacent Peru.

This is a distinctive species with iridescent stem

scales and the indument of the rachis usually con-

fined to the adaxial side. In leaves with a 2-pinnate

lamina, only the basal or lower pinnae are 1 -pin-

nate.

Tumbes: Prov. Zarumilla, Bosque Nacional de Tumbes,

J. Schunke V. 2385 (F, GH, us). Between Cancho and

Cotrina, Coronado 218 (GH, uc). Tumbes to Cancho,

Coronado 228 (uc).

27. Adiantum mathewsianum Hooker, Sp. fil. 2:

35. 1851. TYPE: Peru, (Amazonas), Chach-

apoyas, Mat hews 3296 (holotype, K!; photo,

us; frag, ex K, NY!).

Stem stout, short-creeping. Leaves usually large,

to 1 m or more long, petiole glabrous, rachis sim-

ilar. Lamina broadly ovate-triangular, to 4-pin-

nate. Pinnae widely alternate, the basal very large,

ultimate segments mostly oblong to trapeziform,

bluntly acute (the terminal one acute to acuminate)

glabrous, short-stalked, articulate, the dark color

of the stalk not entering the segment, the inner

edge often overlaying the axis. Indusia many, borne

on the upper and often the distal side, short-oblong

to lunate.

In forests, 450-1 300 m, Amazonas, San Martin,

and Junin.

Peru to Paraguay.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

67

This is a glabrous species with mostly blunt ul-

timate segments. The stalks of the segments are

short and articulate.

Amazonas: Prov. Bagua, above Cascadas de Mayasi,

Wurdack 2024 (OH, us). Prov. Bagua, ca. 5 km S of La

Peca, Barbour4315 (MO). Prov. Bongara, Dist. Sipabam-

ba, Young & Eisenberg 320 (F). San Martin: Tabalosos,

Stiibel 1084 (B). Junin: La Merced, Soukup 1116 (F).

Chanchamayo, 1918, Esposto (USM).

28. Adiantum platyphyllum Sw., Kongl. Vetensk.

Akad.Handl. 1817: 74, /. 3,f. 6. 1817. TYPE:

Brazil, Minas Gerais, Villa Rica, Freyreis (ho-

lotype, Herb. Sw., S-PA!; frag., us!).

Stem moderately stout, short-creeping. Leaves

up to ca. 0.7 m long, petiole glabrous, rachis sim-

ilar. Lamina broadly triangular to ovate-triangu-

lar, 2-pinnate, or small ones oblong, 1 -pinnate.

Pinnae widely alternate, the basal largest, ultimate

segments ovate-lanceolate, cuneate, long-acute to

long-acuminate, glabrous, decidedly glaucous

abaxially, long-stalked, not articulate, the dark col-

or of the stalk entering the base of the segment

beneath, large. Indusia borne on the upper and

distal sides (the inner and lower sides not well

denned), numerous, mostly long-oblong, often ar-

cuate.

In forests, 350-1700 m, San Martin to Madre

de Dios.

Ecuador to Bolivia and Brazil.

The long-stalked, large, glaucous and long-acu-

minate ultimate segments combine to make this

a distinctive species. The dark color of the stalk

enters the base of the segments, while in the fol-

lowing two species, the stalk is articulate at its apex

and the dark color does not enter the segment base.

San Martin: Zepelacio, near Moyobamba, Klug 3490

(F, GH, MO, NY, us). San Roque, LI. Williams 7446 (F,

us). Huanuco: Pozuzo, Macbride 4570 (F, us). Junin:

Chanchamayo valley, C. Schunke 4, 1366, 1368 (F, us).

Madre de Dios: Prov. Tambopata, Vargas 18674 (GH).

29. Adiantum peruvianum Klotzsch, Linnaea 1 8:

555. 1845. SYNTYPES: Peru, (Junin), Vitoc,

Ruiz 25 (B!); Peru, Chachavani, Ruiz 27 (B!).

Frag, ex B, Ruiz & Pavon 25 (us!).

Stem large and massive, very short-creeping.

Leaves to 1 m long, petiole glabrous, rachis sim-

ilar. Lamina broadly ovate-triangular, to 3-pin-

nate, or small ones elongate-triangular or oblong,

1 -2-pinnate. Pinnae widely alternate, the basal

largest, ultimate segments mostly ovate-trapezi-

form, bluntly acute or rounded (the terminal bluntly

acute), glabrous, long-stalked, articulate, the dark

color of the stalk not entering the segment, large.

Indusia many, borne on the upper and distal sides,

few if any on the inner side, roundish to short-

oblong or lunate.

In rich humus in forests, or on rocky slopes or

canyon sides in forests, 750-2800 m, Huanuco to

Cuzco.

Ecuador to Bolivia.

The bluntly acute ultimate segments distinguish

this species from the related Adiantum anceps.

Small fertile leaves may be 1 -pinnate, in larger

ones they are 2-3 -pinnate.

Huanuco: Puente Durand, north of Huanuco, Stork &

Morton 9448 (F, GH, uc, us). Gorge of Rio Chinchao,

Tryon & Tryon 5313 (BM, F, MO, u, us, USM). Pasco:

Prov. Oxapampa, a Oxapampa, Leon 475 (USM). Junin:

Puente San Felix, Cerrate 2869 (GH, USM). 10 km W of

San Ramon, Tryon & Tryon 5450 (BM, F, MO, USM).

Ucayali: Boqueron de Padre Abad, 1943, Ridoutt (USM).

Cuzco: Santa Rosa, Urubamba valley, Cook & Gilbert

1714 (us).

30. Adiantum anceps Maxon & Morton, Amer.

Fern J. 24: 15. 1934. TYPE: Peru, Junin, La

Merced, Killip & Smith 24069 (holotype, us!;

isotype, NY!).

Stem large and massive, short- or very short-

creeping. Leaves to 1.5 m long, petiole glabrous,

rachis similar. Lamina broadly ovate-triangular,

to 3-pinnate, or small ones elongate-triangular or

oblong, 1- or 2-pinnate. Pinnae widely alternate,

the basal largest, ultimate segments mostly ovate-

cuneate concavely acuminate, glabrous, long-

stalked, articulate, the dark color of the stalk not

entering the segment, large. Indusia borne on the

upper and distal sides (the inner and lower sides

not well denned), numerous, roundish to short-

oblong or lunate.

In forests, on steep forested slopes, less often at

the edge of forests or among wet rocks, 200-2000

m, Amazonas to Cuzco.

Colombia, Ecuador, and Peru.

The concavely acuminate, rather than bluntly

acute, ultimate segments separate this species from

the related Adiantum peruvianum.

68

FIELDIANA: BOTANY

Amazonas: Prov. Bagua. above Cascadas de Mayasi,

Wurdack 1885 (F, GH, us). San Martin: Juan Jui, Alto

Rio Huallaga, Klug 4255 (BM, F, GH, MO, NY, uc, us,

USM). Pongo de Cainarachi, Rio Cainarachi, Klug 2688

(BM, F, GH, MO, NY, us). Huanuco: Tingo Maria, Tryon

& Tryon 5288 (BM, F, GH, MO, NY, u, uc, us, USM). Pasco:

Prov. Oxapampa, near confluence of Rio Palcazu and

Rio Iscozacin, D. Smith 1957 (MO). Junin: La Merced,

Killip & Smith 23478 (F, GH, NY, us). Ucayali: Vicinity

of Aguaytia, Croat 20997 (MO), Mathias & Taylor 3589

(F). Cuzco: Prov. La Convencion, Quellouno, Vargas

13556 (GH).

31. Adiantum lucidum (Cav.) Sw., Syn. til. 121.

1806.

Pteris lucida Cav. Descr. pi. 266. 1802. TYPE: Ec-

uador, (Bolivar), Guaranda. Nee (holotype, MA?;

isotype, s!).

Stem moderately stout, short-creeping. Leaves

mostly 30-60 cm long, petiole deciduously pu-

bescent-scaly, rachis similar. Lamina oblong, 1- or

2-pinnate at the base. Entire pinnae asymmetri-

cally ovate-lanceolate to long-triangular, unequal-

ly cuneate, alternate, glabrous or slightly scaly

abaxially, not articulate, the dark color of the stalk

not or hardly continuing into the base of the pinna,

midvein indistinct or lacking in the apical half.

Indusia single, long, on each side of the pinna.

In forests, 300-700 m, San Martin to Madre de

Dios.

Panama; Trinidad and Tobago; Guianas to Co-

lombia and south to Peru.

Jermy and Walker (Bull. Brit. Mus. (Nat. Hist.),

Bot. 13: 173-175. 1985) consider Adiantum lu-

cidum to be a 1 -pinnate sexual diploid species in

Trinidad. Peruvian plants that have a 2-pinnate

lamina may be hybrids with a 2-pinnate species,

or they may be part of the variation of A. lucidum.

San Martin: Prov. Mariscal Caceres, Tocache Nuevo,

J. Schunke V. 4320 (F, GH, us). Huanuco: Prov. Pachitea,

Bosque Nacional de Iparia, J. Schunke V. 1373 (F). Jun-

in: Prov. Chanchamayo, Pichanaki, van der Werffel al.

8620 (MO). Rio Satipo, 1 940, Ridoutt (us, USM). Madre

de Dios: Prov. Tambopata, Rio Piedras, Vargas 18628

(GH).

32. Adiantum poeppigianum (Kuhn) Hieron.,

Hedwigia48: 231. 1909.

Adiantum lucidum var. poeppigianum Kuhn, Jahrb.

Kongl. Bot. Gart. Berlin, 1: 340. 1881. TYPE:

Peru, (Loreto), Prov. Maynas, Poeppig 2268 (ho-

lotype, B!; isotypes, BM!, GH!, LE!, us!).

Stem rather slender, long-creeping. Leaves ca.

30-50 cm long, petiole deciduously pubescent-sca-

ly, rachis similar. Lamina roundish-ovate to ovate-

oblong, usually 1 -(rarely 2-)pinnate at the base.

Entire pinnae lanceolate-cuneate to ovate-lanceo-

late and subcordate, broadest at or near the base,

especially the terminal one, subopposite to alter-

nate, glabrous or slightly scaly abaxially, not ar-

ticulate, the dark color of the stalk continuing into

the base of the pinna, midvein distinct to the apex.

Indusia single, very long, on each side of the pinna

or pinnule.

In dense, or sometimes in rather open, forests,

135-500 m, Amazonas and Loreto to Madre de

Dios.

Endemic to Peru.

This species is distinguished from the somewhat

similar Adiantum lucidum by the dark color of the

stalk of the segment continuing well into the base

of the segment. It is perhaps closer to Adiantum

phyllitidis, which is not known from Peru but may

occur there. Characters of the two species are men-

tioned in Comments.

Amazonas: Prov. Bagua, entre Aramango y Monte-

negro, Lopez et al. 4227 (GH). San Martin: Prov. Mar-

iscal Caceres, Dist. Campanilla, J. Schunke V. 4222 (F,

us). Juan Jui, Ferreyra 4525 (GH, USM). Loreto: Puerte

Arturo, below Yurimaguas, Killip & Smith 27724 (F, NY,

us). Yurimaguas, Killip & Smith 28013 (F, NY, us). Huan-

uco: Prov. Pachitea, Bosque Nacional de Iparia, J.

Schunke V. 1243 (F, GH, us), 1911 (F, GH, us). Junin:

Satipo, 1 940, Ridoutt (GH, USM). Rio Negro to Satipo,

IVoytkowski 5843 (uc, us). Madre de Dios: Parque Na-

cional de Manu, M. S. Foster P-84-97 (MO).

33. Adiantum scalare Tryon, Amer. Fern J. 47:

141, /. 15. 1957. TYPE: Peru, Loreto, Rio

Santiago, Mexia 6162 (holotype, uc!; iso-

types, BM!, F!, GH!, K.I, MO!, PH!).

Stem moderately stout, short-creeping. Leaves

ca. 60-75 cm long, petiole deciduously scaly, rach-

is persistently scaly. Lamina ovate or oblong-ovate,

1 -pinnate, the basal pinnae rarely with a large bas-

al, basiscopic segment. Pinnae closely alternate,

very narrowly long-triangular, entire, often auri-

culate, with long brown trichomes abaxially. In-

dusia single, very long, on each side of the pinna.

In dense forests, 200-400 m, Amazonas to Madre

de Dios.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

69

Endemic to Peru.

This is a very distinctive species, with long, nar-

rowly triangular pinnae that are pubescent abax-

ially and usually have a small, basal, acroscopic

auricle.

Amazonas: Prov. Bagua, above Cascadas de Mayasi,

Wurdack 2000 (GH, us). San Martin: Prov. Mariscal

Caceres, Dist. Campanilla, /. Schunke V. 4253 (F, GH,

us). Loreto: Prov. Maynas, Pebas and vicinity, Plowman

et al. 7034 (F, GH). Huanuco: Prov. Pachitea, Bosque

National de Iparia, Puerto Inca, J. Schunke V. 2975 (F,

GH). Pasco: Cahuapanas, on Rio Pichis, (as Junin), Killip

& Smith 26741 (F, GH, NY, us). Ucayali: Prov. Coronel

Portillo, San Alejandro, Espinoza 281 (USM). Prov. Co-

ronel Portillo, Bosque Nacional de Iparia, Iparia, /.

Schunke V. 2647 (F, GH). Madre de Dios: Manu National

Park, Gentry et al. 27208 (MO).

34. Adiantum macrophyllum Sw., Prodr. 135.

1788. TYPE: Jamaica, Swartz (holotype,

Herb. Swartz, s!).

Stem moderately stout, short to moderately

creeping. Leaves mostly 30-60 cm long, petiole

glabrous, rachis similar. Lamina oblong-acute or

elongate-triangular, 1 -pinnate. Pinnae essentially

opposite, rarely some alternate, entire, long-tri-

angular to ovate-triangular, broadly cuneate, oc-

casionally auriculate or biauriculate, acute to acu-

minate, glabrous, glaucous or dull abaxially,

subsessile, not articulate, the dark color of the short

stalk entering the base of the pinna, large. Indusia

single, long, borne on each side of the pinna, rarely

2 or 3 on some pinnae.

In woods, in dense forests, on shaded banks,

and in rocky places, especially along streams, or

in disturbed vegetation, 200-2500 m, Amazonas

to Puno.

Tropical America.

This is one of the distinctive species of Adian-

tum. It has a 1 -pinnate lamina, commonly op-

posite pinnae that are glabrous or glaucous abax-

ially, and the fertile pinna usually has a single long

indusium on the upper and lower sides.

Amazonas: Prov. Bagua, 5 km S of La Peca, Barbour

4316 (F, MO, USM). San Martin: Tarapoto, Woytkowski

35247 (MO, uc), Soukup 5127 (GH). Loreto: Rio Mara-

non, above Pongo de Manseriche, Mexia 6179a (uc).

Huanuco: Hacienda Mercedes, Mexia 8186 (F, GH, MO,

uc, us). Pasco: Pichis Trail, between San Nicolas and

Azupizu, Killip & Smith 26112 (F, NY, us). Junin: La

Merced, Soukup 1111 (F). Ayacucho: Estrella, between

Huanta and Rio Apurimac, Killip & Smith 23056 (F, GH,

NY, us). Cuzco: Potrero, Tryon & Tryon 5388 (BM, F, MO,

u, us, USM). Madre de Dios: Alto Madre de Dios, Rauh

PI 667 (B). Puno: Prov. Sandia, Chunchosmayo, Weber-

bauer 1267 (B).

Comments

Adiantum phyllitidis John Sm., London J. Hot. 1:

197. 1842. TYPE: British Guiana, Schom-

burgk 300 (BM or K).

This species occurs in the Guianas where it is

rather common, westward to Colombia, and it may

occur in Peru. It is very close to Adiantum poep-

pigianum and the following key will serve to dis-

tinguish these two similar species:

Terminal pinna entire, ovate-acuminate to rather narrowly ovate-acute, subcordate to broadly cuneate

at the base, broadest at or near the base; the lateral entire pinnae similar; or sometimes the apical

pinna with 1 or 2 prominent basal auricles A. poeppigianum

Terminal pinna entire, lanceolate to elliptical, cuneate to narrowly rounded at the base, broadest well

above the base; the lateral entire pinnae similar A. phyllitidis

XV. Pteris

Pteris L., Sp. pi. 1073. 1753; Gen. pi. 484. 1754.

TYPE: Pteris longifolia L. Figure 15.

Terrestrial. Stem erect or short- to long-creep-

ing, bearing scales. Leaves usually monomorphic,

borne at intervals or in a cluster, the petiole with

1-2 (more?) vascular bundles near the base. Lam-

ina 1-5 -pinnate, with pinnate or pedate architec-

ture, the basal pinnae often with enlarged basis-

copic segments, usually glabrous or somewhat scaly

or pubescent abaxially, veins free to wholly anas-

tomosing without included free veinlets. Sori on

a marginal commissure connecting the vein ends,

covered by a well-differentiated marginal indu-

sium, paraphyses absent or rarely present. Spores

tetrahedral-globose or globose, usually with an

equatorial flange, tuberculate to reticulate.

70

FIELDIANA: BOTANY

FIG. 15. Pteris biaurita: a, habit; b, portion of fertile pinna, abaxial side. Pteris altissima: c, portion of pinna,

adaxial side. (Adapted from Stolze, Ferns & fern allies of Guatemala, 1981.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

71

Pteris is a pantropical genus of about 200 species,

a few of them temperate. Twenty-three species are

in Peru, and three of these, Pteris cretica, P. tri-

partita, and P. vittata, are adventive.

It is difficult to place the species of Pteris into

natural groups. In this treatment the species with

less derived lamina architecture (more complex

and pinnate) are followed by those with a derived

architecture (less complex or not pinnate), and free-

veined species precede those with areolate veins.

The stem and leaves of species with very large

leaves such as Pteris Hvida, P. tripartita, and P.

podophylla are poorly represented in herbaria, and

consequently fragmentary specimens may be dif-

ficult or impossible to determine with certainty.

Adequate specimens of these species are much de-

sired.

Most species of Pteris have awns on the adaxial

side of the costa of the penultimate segments at

the base of each costule. A few species such as

Pteris lechleri and P. decurrens lack these awns,

and some such Pteris speciosa have them on some

costae and not on others. The genus is notable for

the often enlarged basal pinnae with elaborated

segments on the basiscopic side.

The treatment of Pteris in Tryon ( 1 964) has been

somewhat modified through the study of new col-

lections.

References

TRYON, R. 1964. Pteris, pp. 188-210, in The

ferns of Peru. Contr. Gray Herb., 194: 1-253.

TRYON, R. M., AND A. F. TRYON. 1982. Pteris,

pp. 332-341, in Ferns and allied plants, Spring-

er-Verlag, New York.

Key to Species of Pteris

a. Veins free (rarely areolae may be present) b

b. All pinnae, or those above the base, entire c

c. Lamina with the basal pinnae usually with a single large pinnule, rarely entire; pinnae 1-5

pairs, cuneate or decurrent, the basal largest or nearly so 22. P. cretica

c. Lamina 1 -pinnate; the pinnae 10 to many pairs, subcordate, the basal ones reduced

23. P. vittata

b. All pinnae pinnatifid or more complex, rarely reduced apical ones are entire d

d. Basal pinnae 1 -pinnate or more complex beyond the basal pinnules e

e. Penultimate segments 1 -pinnate, except sometimes toward the apex; ultimate segments

mostly narrowed at the base, at least on the acroscopic side f

f. Basal pinnae the largest but each much smaller than the remaining portion of the lamina

which is 1-3-pinnate at the base; ultimate segments very coriaceous ... 3. P. coriacea

f. Basal pinnae very large, lamina nearly tripartite, 5-pinnate at the base; ultimate segments

herbaceous 18. P. bakeri

e. Penultimate segments deeply pinnatifid or 1 -pinnate only at the base; ultimate segments

broad at the base g

g. Costa of the penultimate segments (and other axes) smooth abaxially, with deciduous

trichomes, rarely scales; segments usually herbaceous 1 . P. deflexa

g. Costa of the penultimate segments shortly muricate to muricate-spiculate abaxially (the

other axes usually similar), with a caducous scale, rarely a trichome, borne at the apex

of each process; segments coriaceous 2. P. muricata

d. Basal pinnae deeply pinnatifid beyond the 1 (rarely 2 or more) enlarged, basal, inferior, pin-

natifid pinnule(s) h

h. Veins arising from the costules of the ultimate segments, or near their base

4. P. quadriaurita

h. One or 2 veins, especially in the basal portion of the penultimate segment, arising from the

costa of the penultimate segment about halfway between adjacent costules

5. P. pungens

72

FIELDIANA: BOTANY

Venation areolate or predominantly so, or at least a row of areolae along the costa of the penultimate

segment i

i. Pinnae irregularly lobed or some of them entire 17. P. petiolulata

i. Pinnae all entire or regularly pinnatifid or pinnate j

j. Nearly all ultimate segments (usually pinnae or pinnules) stalked, entire, mostly 1 5-40 cm long

k

k. Costal areolae very long and narrow, diverging from the costa; lamina 1 -pinnate; margins

of the sterile pinnae entire back of the apex 15. P. grandifolia

k. Many costal areolae not, or not much, longer than their breadth along the costa 1

1. Lamina 2-(rarely 3-)pinnate at the base, or 1 -pinnate in small leaves; most or many of

the longer areolae 4 times or more longer than broad; margins of the sterile pinnae serrate

back of the apex 14. P. haenkeana

1. Lamina 1 -pinnate, or rarely with an enlarged basal inferior pinnule on the basal pinnae;

most of the longer areolae about twice as long as broad; margins of the sterile pinnae

entire back of the apex 16. P. splendens

j. Ultimate segments joined at the base, or sessile, or lobed m

m. Leaves pinnate, the basal pinnae pinnatifid or pinnate or with a single (rarely more) enlarged

basal inferior pinnule n

n. Leaves dimorphic, the sterile with segments ca. 3 times as broad as the fertile; ultimate

segments mostly more than 10 cm long; few stalked pinnae below the large, deeply

pinnatifid apical portion 13. P. fraseri

n. Leaves monomorphic or nearly so; ultimate segments mostly 2-5 cm long, sometimes

a few to ca. 10 cm long; several to many stalked pinnae below the pinnatifid apical

portion o

o. Two or more costal areolae with their long axis parallel to the costa of the penultimate

segment between adjacent costules; the costa awned on the adaxial side at the base

of the costule 7. P. altissima

o. A single costal areola, or none, with its long axis parallel to the costa of the penultimate

segment between adjacent costules, or if with 2 long areolae then the costa not awned

on the adaxial side p

p. Basal pinnae 1-pinnate-pinnatifid 6. P. propinqua

p. Basal pinnae pinnatifid or with 1 (rarely more) enlarged basal inferior pinnule

q

q. Lamina glabrous or very slightly pubescent r

r. Veins mostly free, only basal areolae present 10. P. biaurita

r. Veins copiously areolate s

s. Apical segment (pinna) of the lamina long-decurrent

11. P. decurrens

s. Apical segment (pinna) of the lamina cuneate or broadly so at the base,

not decurrent t

t. Petiole and rachis reddish to dark reddish; leaf tissue coriaceous;

apex of sterile segments sharply serrate 8. P. speciosa

t. Petiole and rachis stramineous; leaf tissue herbaceous; apex of sterile

segments nearly entire to crenulate-serrate 9. P. transparent

q. Lamina hirsute on both surfaces, especially on the costa of the penultimate

segments 12. P. lechleri

m. Leaves ternate, or pedate and the basal pinnae twice branched, with 3 main axes, each axis

1-pinnate-pinnatifid beyond a branch u

u. Lamina ternate; apex of ultimate segments crenulate to crenulate-serrate; ultimate seg-

ments usually ca. 6-10 cm long 19. P. livida

u. Lamina pedate; ultimate segments ca. 2 cm long or less v

v. Apex of ultimate segments sharply serrate 20. P. podophylla

v. Apex of ultimate segments crenulate 21. P. tripartita

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II. 73

1. Pteris deflexa Link, Hort. berol. 2: 30. 1833.

TYPE: Brazil, hort. hot. berol. (holotype, Link

Herb., B!; photo, GH).

Pteris polita Link, Hort. berol. 2: 30. 1833. TYPE:

hort. bot. berol. (holotype, Link Herb., B!; photo,

GH).

Stem moderately to very stout, to 6-8 cm in

diameter, short-creeping or ascending. Leaves 0.5-

3 m long. Lamina pinnate or ternate, 2-3-pinnate-

pinnatifid at the base, the basal pinnae the largest,

regularly 1 -pinnate-pinnatifid or sometimes with

1 or 2 enlarged basal pinnules, pinnae deeply pin-

natifid beyond the lower or basal ones. Penulti-

mate segments deeply pinnatifid, glabrous, or

slightly pubescent or scaly abaxially. Veins free.

In dense forests, 800-2300 m, Cajamarca and

Amazonas south to Cuzco.

Tropical South America.

Differences between this species and the next,

Pteris muricata, are discussed under that species.

Morton (Contr. U.S. Natl. Herb. 38: 75. 1967)

erroneously considered specimens at Hamburg to

typify Pteris polita Link, and consequently that

name was used for Pteris propinqua. However, the

holotype of Pteris polita at Berlin is clearly Pteris

deflexa.

Cajamarca: Prov. Cutervo, Gruta San Andres, Mos-

tacero et al. 1676 (F, GH). Amazonas: Ca. 10 km W of

Leimebamba. Edwin & Schunke 3701 (F). Prov. Bagua,

E of La Peca, Barbour 2701 (MO, uc). San Martin: Tar-

apoto, Spruce 4326 (BM, GH, K, us). Pasco: Prov. Oxa-

pampa, vicinity of Chequitavo, D. Smith 5273 (GH).

Junin: Prov. Chanchamayo, Rio Rondayacu, Smith et

al. 2608 (F). Cerca a Huacapistana, Ferreyra 503 (GH,

USM). Ucayali: Prov. Coronel Portillo, Sinchono, entre

Tingo Maria y Pucallpa, Aguilar 898 (GH, USM). Ayac-

ucho: Ccarrapa, between Huanta and Rio Apurimac, Kil-

lip & Smith 22408 (F, GH, us). Cuzco: Quillabamba, Pey-

ton & Peyton 1207 (GH).

2. Pteris muricata Hooker, Sp. fil. 2: 193, /. 123B.

1858. TYPE: Colombia, Antioquia, Jervise

(holotype, K!).

Stem moderately stout, short- to long-creeping,

decumbent to ascending. Leaves 0.5-2 m or more

long. Lamina pinnate, ternate or pedate, 2-4-pin-

nate-pinnatifid at the base, basal pinnae the larg-

est, usually regularly 1 -pinnate-pinnatifid or with

1 or more enlarged basal pinnules, deeply pinna-

tifid to 2-pinnate-pinnatifid beyond the lower or

basal pinnae. Penultimate segments deeply pin-

natifid, rarely 1 -pinnate at the base, shortly mur-

icate to muricate-spiculate abaxially, deciduously

scaly and rarely pubescent beneath. Veins free.

In forests and cloud forests, in moist ravines, at

the edge of woods, and in moist, rocky places,

1 700-4000 m, Cajamarca south to Puno.

Mexico south to Bolivia.

This species may be related to Pteris bakeri by

the muricate processes on the costa beneath. It is

evidently closer to the previous species, Pteris de-

flexa, from which it differs in the firmer texture

of the lamina segments and the muricate to mur-

icate-spiculate costae abaxially. In addition, the

processes of Pteris muricata bear caducous scales,

while in P. deflexa there are deciduous trichomes

abaxially. Rarely there are specimens that appear

to be intermediate between the two species.

Cajamarca: Prov. San Miguel, Cerro Quillon, Mos-

tacero et al. 1297 (F). Huanuco: Mito, Macbride & Feath-

erstone 76/9 (F, us). Muna, Bryan 465 (F, GH). Pasco:

Quillasu, Soukup 3283 (GH). Junin: Between Palca and

San Ramon, Correll & Smith P785 (GH). Huacapistana,

Killip & Smith 24191 (us). Huancavelica: Prov. Taya-

caja, cerca a Tocas, Tovar 2053 (USM). Apurimac: Prov.

Abancay, arriba de Abancay, Ferreyra 9808 (GH, USM).

Cuzco: Machu Picchu, Tryon & Tryon 5398 (BM, F, GH).

Tres Cruces, Vargas 1055 (GH, us). Puno: Prov. Cara-

baya, entre Ayapata y Kahualluyoc, Vargas 10751 (GH).

Tabina, Lechler 2030 (B, K).

3. Pteris coriacea Desv., Mem. Soc. Linn. Paris

6: 300. 1 827. TYPE: Peru, Dombey (holotype,

Desvaux Herb., the complete leaf, P!; photo,

GH; isotype, Herb. Gen., P!; photo, GH).

Stem moderately stout, short-creeping. Leaves

30-75 cm or more long. Lamina pinnate, 2-3-

pinnate at the base, basal pinnae the largest, reg-

ularly 1 -pinnate beyond the often 1 or more en-

larged basal pinnules, pinnae mostly 1 -pinnate be-

yond the lower or basal ones. Penultimate segments

mostly 1 -pinnate, smooth to muricate-spiculate

and deciduously scaly abaxially. Veins free.

In woods and in rocky places, 2800-3900 m,

Ancash south to Apurimac.

Ecuador and Peru.

The separate ultimate segments mostly nar-

rowed at the base are characteristic of this species.

It is related to the previous species, Pteris muricata,

by the muricate-spiculate costae abaxially and may

be a high-altitude derivative of that species. Most

collections are from 3100-3900 m, while those of

P. muricata are mostly from 3000 m or lower.

74

FIELDIANA: BOTANY

Ancash: Prov. Bolognesi, cercaa Llamac, Cerrate2372

(GH, USM). Huanuco: Chasqui, Macbride & Featherstone

1755 (F, us). Lima: Between San Mateo and Parac, Co-

ronado 312 (GH, uc). Junin: Prov. Huancayo, entre

Huaytapallana y Chilifruta, Tovar 4290 (GH). Tarma,

Kunkel 406 (GH). Apurimac: Prov. Abancay, Bosque de

Ampay, Vargas 1062 (GH, us).

4. Pteris quadriaurita Retz., Observ. hot. 6: 38.

1791. TYPE: Ceylon, Konig (holotype, Ret-

zius Herb., LD).

Pteris edentula Kunze, Linnaea 9: 75. 1834. (as Pteris

biaurita van? P. edentula) TYPE: Peru, (Hua-

nuco), Pampayacu, Poeppig233 (Diar. 1106) (ho-

lotype, LZ destroyed; isotype, B!; photo, GH; frag.,

us!).

Stem moderately stout, decumbent or erect.

Leaves ca. 0.5-2 m long. Lamina pinnate, 2-pin-

nate-pinnatifid at the base, basal pinnae regularly

deeply pinnatifid beyond the usually single en-

larged basal inferior pinnule, pinnae usually deep-

ly pinnatifid beyond the basal ones. Penultimate

segments deeply pinnatifid, glabrous or usually

slightly pubescent abaxially. Veins free.

Dense forests, woods and thickets, sometimes

in rocky places, 650-2200 m, Lambayeque south

to Cuzco.

Tropical America; Old World.

In Pteris quadriaurita the veins of an ultimate

segment arise from the costule, or near it, while

in Pteris pungens there are one or two veins that

arise from the costa, distant from the costule, es-

pecially in the basal portion of a penultimate seg-

ment.

Lambayeque: Km 32 on road from Olmos to Jaen,

Correll & Smith P830 (GH). Amazonas: Prov. Bagua,

Cordillera Colan, SE of La Peca, Barbour 4192 (MO).

Huanuco: Rio Huallaga, below Rio Santo Domingo,

Macbride 4233 (F, us). Pampayacu, Macbride 5043 (F,

us). Pasco: Prov. Oxapampa, SE of Oxapampa, D. Smith

2898 (MO). Junin: La Merced, Killip & Smith 23473 (F,

us), 23686 (F, us), Cerrate 2832 (GH, USM). Cuzco: Po-

trero, Tryon & Tryon 5367 (BM, F, GH, u, us, USM), Vargas

8236 (uc).

5. Pteris pungens Willd., Sp. pi. ed. 4, 5: 387.

1810. LECTOTYPE (designated by Proctor,

Flora Lesser Antilles 2: 144. 1977): Hispan-

iola, Plumier, Traite foug. Amer. t. 14. 1705.

Stem moderately stout, decumbent or erect.

Leaves ca. 0.5-2 m long. Lamina pinnate, 1-pin-

nate-pinnatifid to usually 2-pinnate-pinnatifid at

the base, basal pinnae regularly deeply pinnatifid

beyond the usually single enlarged, basal inferior

pinnule, pinnae deeply pinnatifid beyond the basal

ones. Penultimate segments deeply pinnatifid, gla-

brous or nearly so abaxially. Veins free.

Dense forests, 100-900 m, San Martin and Lo-

reto south to Madre de Dios.

Mexico and the West Indies; northern South

America, south to Bolivia.

The difference in the venation between Pteris

quadriaurita and this species is mentioned under

the former species.

San Martin: Tarapoto, Spruce 4750 (K). Prov. San

Martin, E of Tarapoto, Woytkowski 35215 (MO). Loreto:

Yurimaguas, Ll. Williams 3865 (F, us). Balsapuerto, Kil-

lip & Smith 28517 (GH, us). Huanuco: Rio Llullapichis,

Dudley 12483 (GH), 12485 (GH). Prov. Pachitea, Bosque

Nacional de Iparia, J. Schunke V. 1755 (F, GH). Madre

de Dios: Prov. Tambopata, Rio Tambopata, Barbour

4880 (MO).

6. Pteris propinqua Agardh, Recens. spec. pter.

65. 1 839. LECTOTYPE (designated here): Ja-

maica, Bancroft (holotype, K!). LECTO-

PARATYPE: Jamaica, MacFadyn, K.

Stem moderately stout to stout, erect. Leaves

0.5-1.5 m long. Lamina pinnate, 2-pinnate-pin-

natifid at the base, basal pinnae the largest, rather

regularly 1-pinnate-pinnatifid, pinnae deeply pin-

natifid beyond the lower or basal ones. Penulti-

mate segments deeply pinnatifid, the base narrow-

ly decurrent on its stalk or onto the next axis,

glabrous or minutely pubescent abaxially. Veins

partly areolate, 1 costal areola with its long axis

parallel to the costa between adjacent costules.

In forests and along forest borders, 1 00-700 m,

San Martin south to Pasco.

Tropical America.

The misapplication of the name Pteris polita to

this species has been mentioned under Pteris de-

flexa.

The base of the penultimate segments is decur-

rent onto the stalk or onto the next axis.

San Martin: Prov. Mariscal Caceres, Tocache Nuevo,

J. Schunke V. 3378 (F, GH, us), 7038 (MO). Loreto: Gam-

itanacocha, Rio Mazan, J. Schunke 282 (F, GH, uc, us).

Paraiso, upper Rio Itaya, Ll. Williams 3356 (F, us).

Huanuco: Tingo Maria, (as San Martin), Allard 21920

(GH, us). Pasco: Puerto Bermudez, (as Junin), Killip &

Smith 26639 (us).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

75

7. Pteris altissima Poiret in Lam., Encycl. 5: 722.

1804. TYPE: Porto Ricco (Puerto Rico), Led-

ru (holotype, Lamarck Herb., P!; photo, GH).

Figure 15c.

Pteris kun:eana Agardh. Recens. spec. pter. 62. 1839.

SYNTYPES: Peru, Poeppig 225 is one of several

collections cited, B!; photo, GH, as is Peru, Ma-

thews 1802, K!; frag., NY.

Stem erect in young plants, becoming moder-

ately stout and short- to rather long-creeping.

Leaves 0.8-2.5 m long. Lamina pinnate or nearly

ternate, 2-pinnate-pinnatifid at the base, basal pin-

nae the largest, 1-pinnate-pinnatifid toward the

base, deeply pinnatifid beyond, pinnae pinnatifid

beyond the lower or basal ones. Penultimate seg-

ments pinnatifid to deeply pinnatifid, glabrous or

minutely pubescent abaxially. Veins copiously

areolate, 2 or more costal areolae with their long

axis parallel to the costa between adjacent costules.

In rain forests, montane forests, and in wet ra-

vines, 1 00-1 500 m, Amazonas south to Cuzco and

Madre de Dios.

Tropical America.

This species is characterized by having two or

more of the costal areolae with their long axis

parallel to the costa between adjacent costules, and

with awns on the costa above.

Amazonas: Prov. Bagua, Rio Utcubamba, 40 km S of

Bagua Grande, Hutchison 1486 (GH, us). San Martin:

Tarapoto, Spruce 4061 (BM, K). Zepelacio, near Moy-

obamba, Klug 3704 (F, GH, K, us). Loreto: Mouth of Rio

Santiago, above Pongo de Manseriche, Mexia 6125 (F,

GH, K, MO, uc, us), 6125a (GH, K, uc, us). Huanuco:

Tingo Maria, Tryon & Tryon 5229 (BM, F, GH, u, USM).

Pasco: Prov. Oxapampa, vicinity of Chequitavo, D. Smith

5258 (GH). Junin: La Merced, Soukup 1098 (F). Ucayali:

Prov. Coronel Portillo, Bosque Nacional Humboldt,

Vdsquez 3387 (F). Ayacucho: Near Kimpitiriki, Rio Apu-

rimac valley, Killip & Smith 23039 (us). Cuzco: Prov.

Paucartambo, Kosnipata, Vargas 11260(GH). Madre de

Dios: Prov. Manu, Atalaya, Foster & Wachter 7435 (MO).

8. Pteris speciosa Kuhn, Linnaea 36: 91. 1869.

LECTOTYPE (designated here): Peru, St.

Gavan (Rio San Gaban), Lechler 2326 (ho-

lotype, B!, Herb. Mett.; photo, GH; frags., NY!,

us!. ISOLECTOPARATYPES: Peru, (San

Martin), Tarapoto, Spruce 4759, K.!; photo,

GH; Ecuador, Spruce 4757, BM!, K!; photo, GH

ofK).

Pteris reticulata Kuhn, Linnaea 35: 91. 1869, exchar.,

not Desv. 1811. SYNTYPES: Peru, Pavon, Peru,

Spruce 4752 (4725?). Material of these collec-

tions at B! may be authentic).

Pteris reticulalovenosa Hieron., Hedwigia 48: 243.

1909, nom. nov. for Pteris reticulata Kuhn, not

Desv., and with the same type.

Stem moderately stout, creeping. Leaves ca.

0.75-1.75 m long. Lamina pinnate, shallowly to

deeply 1 -pinnate-pinnatifid. Pinnae with the ster-

ile or partly fertile ones rather shallowly pinnatifid

and the fully fertile ones shallowly to deeply pin-

natifid, glabrous or minutely pubescent abaxially,

the ultimate segments falcate, coriaceous. Veins

copiously areolate, 1 costal areola with its long

axis parallel to the costa between adjacent costules.

In forests, ca. 1000-1800 m, San Martin south

to Puno.

Colombia to Peru.

Kuhn cited Spruce 4757 as from Ecuador, while

the specimen at K has a label indicating Peru. The

specimen of Spruce 4759 at K has a pencil note

indicating it to be 4757. How many separate col-

lections are involved is uncertain.

Pteris speciosa is especially separated from P.

transparens by the reddish petiole and rachis of P.

speciosa rather than the stramineous petiole and

rachis of P. transparens. In addition, the apex of

sterile segments is sharply serrate in P. speciosa,

while it is nearly entire to crenulate-serrate in P.

transparens. Some specimens of P. speciosa lack

awns on the costa adaxially, while these are present

in P. transparens.

Small portions of a leaf, usually pinnae, may be

similar in P. speciosa and P. livida. These may be

separated as follows. In P. speciosa the costule of

the ultimate segments is prominently raised adax-

ially and rounded, or sometimes narrowly grooved

with the two edges rounded. In P. livida the costule

of the ultimate segments is grooved adaxially with

thin, sometimes narrow wings, which may be

somewhat appressed on either side of the costule.

Pasco: Dos de Mayo (as Junin), Killip & Smith 25812

(us). Cuzco: Entre Quincemil y San Lorenzo, Vargas

11724(GVL).

9. Pteris transparens Mett., Ann. Sci. Nat. Bot.

V, 2: 221. 1864. TYPE: Colombia, (Cundi-

namarca) "Andes Bogotenses," 1400 m,

Triana, evidently 622 (holotype, LZ de-

76

FIELDIANA: BOTANY

stroyed; isotypes, B!, BM!, COL; frag., GH; pho-

tos, GH of B, BM, COL).

Pteris nuda Copel., Univ. Calif. Publ. Bot. 19: 302.

1941. TYPE: Peru, Huanuco, Riochuela Chun-

talagua, Mexia 8299 (holotype, uc; isotypes, F!,

GH!, K!).

Stem rather stout, evidently short-creeping.

Leaves ca. 1-1.5 m long. Lamina pinnate, 1-pin-

nate-pinnatifid at the base, or the basal pinnae

regularly deeply pinnatifid beyond the sometimes

present single enlarged basal inferior pinnule. Pin-

nae, above the basal ones deeply pinnatifid, es-

sentially glabrous abaxially, ultimate segments

patent or nearly so, nearly straight, herbaceous.

Veins copiously areolate, 1 costal areola with its

long axis parallel to the costa between adjacent

costules.

Forested banks and rocky slopes, ca. 600 m,

Huanuco and (probably) Junin.

Colombia and Peru.

The petiole and rachis is stramineous, rather

than reddish as in Pteris speciosa. Also the apex

of sterile segments is nearly entire to crenulate-

serrate rather than sharply serrate. This species is

evidently very rare, and although distinct among

the Peruvian ones, it is a member of a complex in

northern South America which requires clarifi-

cation. Awns are present on the costa adaxially,

these being absent in the otherwise similar Pteris

consanguinea of Venezuela.

Huanuco: Pozuzo, Macbride 4718 (us). Junin: Pichita

Caluga, Walden 56 (GH), and 80 (GH) are young plants,

probably of this species.

10. Pteris biaurita L., Sp. pi. 1076. 1753. LEC-

TOTYPE (designated by Proctor, Flora Less-

er Antilles 2: 145. 1977): Plumier, Descr. pi.

Amer. /. 14. 1693, and Traite foug. Amer. t.

15. 1705. LINN 1246.19 is this species. Fig-

ure 15a-b.

Stem moderately stout, decumbent to erect.

Leaves ca. 0.5-1 m long. Lamina pinnate, 2-pin-

nate-pinnatifid at the base, basal pinnae regularly

deeply pinnatifid beyond the single enlarged basal

inferior pinnule, pinnae deeply pinnatifid beyond

the basal ones. Penultimate segments deeply pin-

natifid, glabrous or minutely pubescent abaxially.

Veins free, except for 1 costal areola with its long

axis parallel to the costa between adjacent costules,

or a few other basal veins joined.

In forests, in clearings and at the edge of forests,

sometimes in rocky places, 100-1 500 m, Huanuco

south to Cuzco.

Tropical America; Old World.

The species is characterized by the areolae only

along the costa, although there may be a few else-

where.

Huanuco: Tingo Maria, Tryon & Tryon 5246 (GH, u,

us). Pozuzo, Macbride 4596 (F). Pasco: Rio Paucartam-

bo, (as Junin), Killip & Smith 25277 (F, us). Junin: Chan-

chamayo valley, C. Schunke 24 (F). Ucayali: Boqueron

Padre Abad, (as San Martin), Allard 22115 (us). Cuzco:

Prov. La Convencion, Chacanares, Vargas 13680 (us).

11. Pteris decurrens Presl, Delic. prag. 1: 183.

1822. TYPE: Brazil, (Guanabara), Rio de Ja-

neiro and vicinity, collector and location not

known.

Pteris horizontalis (Fee) Rosenst., Meded. Rijks-Herb.

19: 10. 1913.

Litobrochia horizontalis Fee, Crypt, vase. Bresil 1 : 48,

t. 12, f. 1. 1869. TYPE: Brazil, Glaziou 2314

(holotype, P?; frag, ex Fee, NY!; photo, GH).

Stem stout, evidently erect. Leaves ca. 1-2.5 m

long. Lamina pinnate, 2-pinnate-pinnatifid at the

base, basal pinnae regularly deeply pinnatifid be-

yond the single enlarged basal inferior pinnule,

pinnae deeply pinnatifid beyond the basal ones.

Penultimate segments deeply pinnatifid, glabrous

or minutely pubescent abaxially, the upper ones

decurrent onto the rachis. Veins copiously areo-

late, 1 or 2 costal areolae with their long axis par-

allel to the costa between adjacent costules.

In wet woods and cloud forests, ca. 1800 m,

Amazonas and Cuzco.

Venezuela, Peru, Bolivia, and Brazil.

The adaxial side of the costa lacks awns and the

apical pinna of the lamina is long decurrent on its

stalk.

Amazonas: Serrania de Bagua, E of La Peca, Gentry

el al. 22864 (us). Cuzco: Prov. Paucartambo, valle de

Pillahuata, Herrera 1621 (us).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

77

12. Pteris lechleri Men., Fil. lechl. 2: 13. 1859.

TYPE: Peru, Dept. Puno, Prov. Carabaya,

Tatanara, Lechler 2533 (holotype, B!; photo,

GH).

Pteris vestita Baker, Syn. fil. 169. 1867. TYPE: Peru,

(San Martin). Tarapoto, Spruce 4063 (holotype,

K!; frag. & photo, us).

Pteris killipii Maxon, Amer. Fern J. 23: 107. 1933,

nom. super/1, for Pteris vestita Baker and with the

same type.

Stem small to moderately stout, erect. Leaves

ca. 0.5-1.25 m long. Lamina pinnate, 2-pinnate-

pinnatifid at the base, basal pinnae regularly deep-

ly pinnatifid beyond the single enlarged basal in-

ferior pinnule, pinnae deeply pinnatifid beyond

the basal ones. Penultimate segments deeply pin-

natifid, hirsute on both surfaces, especially on the

costa. Veins copiously areolate, 1 costal areola with

its long axis parallel to the costa between adjacent

costules.

Montane forests, 850-1 500 m, San Martin south

to Puno.

Peru and Bolivia.

This is a very distinctive species because of the

hirsute lamina and the lack of awns on the adaxial

side of the costa.

San Martin: Zepelacio, near Moyabamba, Klug 3680

(F, GH, K, us). Huanuco: Tingo Maria, (as San Martin),

Allard21195 (us), 27207 (us), 27594 (us). Prov. Leoncio

Prado, al este de Tingo Maria, J. Schunke V. 10222 (F,

MO, us). Junin: Near La Merced, Killip & Smith 23962

(F, GH, us). La Merced, Macbride 5713 (F, us). Ayacucho:

Prov. La Mar, Cordillera Central, Dudley 11946 (GH,

us). Cuzco: Tocate. Biies 1740 (us). Puno: Prov. Cara-

baya, a Sanari, Vargas 17548 (GH).

1 3. Pteris fraseri Kuhn, Bot. Zeit. (Berlin) 26: 4 1 .

1868. LECTOTYPE (designated here): Ec-

uador, Fraser, Herb. Mett. B!. LECTOPAR-

ATYPES: Ecuador, Spruce 5139 and coll.

Hohenacker 42 (not located).

Stem stout, evidently erect. Leaves ca. 2 m long,

dimorphic, the fertile taller than the sterile and

with narrower, well-spaced segments. Lamina ter-

nate, 2-pinnate-pinnatifid at the base, basal pinnae

the largest, regularly deeply pinnatifid beyond the

usually present enlarged basal inferior pinnule,

pinnae deeply pinnatifid beyond the basal ones.

Ultimate segments elongate, glabrous or minutely

pubescent abaxially. Veins copiously areolate, 2

or more costal areolae with their long axis parallel

to the costa between adjacent costules.

Rain forests and in a coffee plantation, 1500-

1 800 m, San Martin, Pasco, and Junin.

Ecuador and Peru.

This species is distinguished by its dimorphic

leaves, the unusual lamina architecture with the

ultimate segments long and entire and mostly

widely spaced, and the lack of awns on the adaxial

side of the costa. It seems close to Pteris macrop-

tera Link of Brazil but that species is hardly di-

morphic.

San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba, D.

Smith 4421 (GH). Pasco: Prov. Oxapampa, Villa Rica,

van der Werffet al. 8296 (MO, uc). Junin: Prov. Chancha-

mayo, S of San Ramon, Smith & Palacios 2645 (F).

14. Pteris haenkeana Presl, Reliq. haenk. 1: 55.

1825. TYPE: Haenke, probably Peru, Dept.

Huanuco (holotype, PR?).

Pteris ampla Kunze, Linnaea 9: 74. 1834. TYPE: Peru,

(Huanuco), Pampayacu, 1829, Poeppig (Diar.

1154) (holotype, probably LZ destroyed. ISO-

TYPES: K!, MO!).

Stem ca. 1 cm in diameter, long-creeping. Leaves

1-2 m long. Lamina pinnate, fully 2-pinnate at or

toward the base, or rarely 3-pinnate, basal pinnae

the largest, 1 -pinnate beyond the lower or basal

pinnae. Pinnae (or pinnules) large, simple, entire,

short- to usually long-stalked, glabrate or with mi-

nute, straight, appressed trichomes abaxially. Veins

copiously areolate, the costal areolae short and

broad.

In forests, 100-1400 m, Amazonas south to

Cuzco.

Colombia south to Bolivia.

This species has very large, entire ultimate seg-

ments. The differences from Pteris grandifolia, with

which it is sometimes confused, are mentioned

under that species.

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

que: 4949 (GH). San Martin: Juan Jui, Klug 4256 (F, GH,

K, MO, uc, us). Zepelacio, near Moyabamba, Klug 3492

(F, GH, K, MO, us). Huanuco: Pampayacu, Kanehira 147

(GH, us). Cushi, Bryan 724 (F, GH, us). Pasco: Prov.

Oxapampa, Villa Rica, van der Werffet al. 8289 (MO).

Junin: La Merced, Soukup 1091 (F). Perene, Kunkel 505

(GH). Ayacucho: Ayna, between Huanta and Rio Apu-

rimac, Killip & Smith 22717 (F, GH, us). Cuzco: Cco-

chayoc, Biies 1726 (us).

78

FIELDIANA: BOTANY

1 5. Pteris grandifolia L., Sp. pi. 1 073. 1 753. LEC-

TOTYPE (designated by Proctor, Flora Less-

er Antilles 2: 143. 1977): Plumier, Descr. pi.

Amer. t. 105 not 106, 1693; LINN 1246.1;

photo, A, is this species.

Pteris grandifolia var. campanae Rosenst., Repent.

Spec. Nov. Regni Veg. 7: 291. 1909. TYPE: Peru,

(San Martin), Mt. Campana, Tarapoto, Spruce

4668 (holotype, location unknown; isotype, K!).

Stem ca. 1 cm in diameter, rather extensively

creeping. Leaves 1-5 m long. Lamina pinnate, 1-

pinnate or rarely 2-pinnate at the base, 1 -pinnate

beyond. Pinnae large, simple, entire, short- to long-

stalked, glabrate to minutely tortuous-pubescent

abaxially, especially on the veins. Veins copiously

areolate, the costal areolae very long, with their

long axis divergent from the costa.

In forests, in thickets, along forest borders, and

on stream banks, 1 35-1 1 00 m, Lambayeque south

to Cuzco.

Tropical America.

This species is similar to Pteris haenkeana but

can be distinguished by the usually 1 -pinnate lam-

ina, the sterile margins of the pinnae that are entire

back of the apex, and the tortuous minute pubes-

cence on the pinnae abaxially. In Pteris haenkeana

the lamina is 2- or 3-pinnate at the base, the sterile

margins are sharply serrate back of the apex, and

the minute pubescence abaxially is straight.

Lambayeque: Puerto Nazareth, 5 km to Olmos, £7-

lenberg 3429 (GH). Amazonas: Prov. Bagua, entre Ara-

mango y Montenegro, Lopez el at. 4214 (GH). San Mar-

tin: Chazuta, Rio Huallaga, Klug 4013 (F, GH, K, MO,

uc). I.oreto: Rio Maranon, above Pongo de Manseriche,

Mexia 6182 (F, GH, K, MO, uc, us). Huanuco: Tingo

Maria, Tryon & Tryon 5287 (BM, F, GH, u, us, USM).

Pasco: Oxapampa, (as Junin), Soukup 2349 (F, GH). Jun-

in: La Merced, Killip & Smith 23693 (F, GH, us). Cerca

a La Merced, Cerrate 2825 (GH, USM). Ucayali: Bosque

Nacional von Humboldt, Gentry et al. 41412 (MO). Cuz-

co: Kosnipata, Peralta 12 (USM).

16. Pteris splendens Kaulf, Enum. fil. 186. 1824.

TYPE: Brazil, Chamisso (holotype, probably

LZ destroyed; isotype, LE!; photo, GH).

Stem stout, short-creeping or nearly erect. Leaves

to ca. 2 m long. Lamina pinnate, 1 -pinnate, the

lower pinnae not or not much reduced, or the basal

pinnae with an enlarged basal inferior pinnule, 1-

pinnate beyond the base. Pinnae very long, large,

simple, entire, short- to long-stalked, glabrous

abaxially. Veins copiously areolate, the costal ar-

eolae short and broad, the longer ones about twice

as long as broad.

In forests, 780 m, Huanuco.

Peru and Brazil.

While this species has only recently been col-

lected in Peru, it probably occurs south to Puno

and also in Bolivia. The few collections from Peru

have the pinnae about half as broad as in most

Brazilian specimens. The pinnae are long and en-

tire and have very conspicuous areolate venation.

Huanuco: Prov. Leoncio Prado, al este de Tingo Mar-

ia, J. Schunke V. 5211 (F, GH, us), 10113 (F, uc), E of

Tingo Maria, Croat 21 192 (MO, uc).

17. Pteris petiolulata Tryon, Rhodora 62: 9. 1960.

TYPE: Peru, (Junin), La Merced, Macbride

5714 (holotype, us!; isotype, F!).

Stem rather small, erect. Leaves 0.75-1 .5 m long.

Lamina pinnate, 1-2-pinnate-pinnatifid at the base,

with long, entire simple pinnae and those with

irregularly spaced pinnatifid or lobed portions var-

iously placed. Pinnae somewhat pubescent on both

surfaces, at least on the costa and costules. Veins

copiously areolate, 2 or more costal areolae with

their long axis parallel to the costa between ad-

jacent costules, when present.

In forests, 500-1500 m, Huanuco, Junin, and

Cuzco.

Venezuela, Colombia, and Peru.

The lamina architecture of this species is very

irregular, and differs among the several collections.

This is indicative of a hybrid origin, possibly be-

tween Pteris splendens and Pteris lechleri. It has

the prominent venation and often long, entire pin-

nae, or entire portions of pinnae, of the former

species and the hirsute costa and lack of awns on

the costa of the latter one. However, it is sparsely

but rather widely distributed and at the present

time its putative parents are not known from sev-

eral of its localities.

Huanuco: Prov. Leoncio Prado, al este de Tingo Mar-

ia, J. Schunke V. 5211 (F, GH, MO), 10113 (MO), both

numbers are also P. splendens. Junin: San Ramon, Killip

& Smith 24696 (F, GH, us). Chanchamayo, Schunke 965

(F). Prov. Tarma, Perene, Esposto (USM). Cuzco: Rio Pa-

chiri, Bues 1767 (us). Prov. Paucartambo, Kosnipata-

Pilcopata, Vargas 14755 (GH).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

79

18. Pteris bakeri C. Chr., Index fil. 593. 1906.,

nom. nov. for Pteris decomposita Baker, not

Gaud., and with the same type.

Pteris decomposita Baker, Syn. fil., ed. 2, 479. 1874.

TYPE: Peru, Dept. Huanuco, Muna, Pearce (ho-

lotype, K!; frag., NY!; Hooker's, Icon. pi. /. 1641.

Stem moderately stout, creeping. Leaves ca. 1

m long. Lamina pinnate, tripartite or nearly so, to

5-pinnate at the base, the basal pinnae much the

largest, pinnae often 2-pinnate-pinnatifid beyond

the basal ones. Penultimate segments deeply pin-

natifid to 1 -pinnate, muricate-spiculate and de-

ciduously scaly abaxially. Ultimate segments usu-

ally lobed, often deeply so. Veins free.

In forests, 2300-3000 m, Huanuco, Pasco, Jun-

in, and Cuzco.

Endemic to Peru.

The lamina is much divided with small ultimate

segments. The muricate-spiculate axes suggest a

relation to Pteris muricata.

Huanuco: Pozuzo, Pearce 534 (K). Pasco: Prov. Oxa-

pampa, Rio San Alberto, Foster et al. 10298 (F), van der

Werffet al 8480 (MO, uc). Junin: Carpapata, above Hu-

acapistana, Killip & Smith 24488 (us). Cuzco: Prov. Uru-

bamba, Skog & Skog 5176 (us).

19. Pteris livida Mett., Ann. Sci. Nat. Bot. V, 2:

222. 1864. LECTOTYPE (designated here):

Colombia, (Cundinamarca), Bogota, Choachi,

Lindig 179, B!, Herb. Mett.; photo, GH. IS-

OLECTOTYPES: BM!, GH!, P!, us!. LECTO-

PARATYPES: Colombia (Tolima), Quindio,

Triana, and Linden 1089 (not located).

Stem very stout in large plants. Leaves 2-6 m

long. Lamina ternate, basal pinnae the largest, reg-

ularly 2-pinnate-pinnatifid, pinnae deeply pinna-

tifid beyond the lower or basal ones. Penultimate

segments deeply pinnatifid or 1 -pinnate at the base,

glabrate to minutely pubescent abaxially, the ul-

timate segments long, entire. Veins copiously are-

olate, 1 costal areola with its long axis parallel to

the costa between adjacent costules.

In wet forests and cleared forests, 900-2500 m,

Amazonas south to Cuzco.

Costa Rica to Colombia, south to Bolivia.

This species has a very large, ternate lamina in

large plants. The costae may bear awns adaxially

at the base of the costules, or these may be lacking.

The ultimate segments are usually large and often

narrowed toward, but not at, the base. The iden-

tification of small portions of the lamina (usually

upper pinnae) has been discussed under Pteris spe-

ciosa.

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

2497 (F, MO). San Martin: Cerro de Escalera, between

Yurimaguas and Moyobamba, Vie 6885 (B). Prov. Mar-

iscal Caceres, N of Pulcache, Plowman & Schunke 11630

(F, GH). Huanuco: Prov. Huanuco, Hacienda Mercedes,

Afexia 8219 (F, GH, K, uc, us). Prov. Huanuco, Mirador,

Feb. 23, 1940, Ridoutt (GH, USM). Pasco: Prov. Oxapam-

pa, Rio San Daniel, Smith et al. 7813 (F). Cuzco: Prov.

La Convencion, Cordillera Vilcabamba, Dudley 1 1266

(GH).

20. Pteris podophylla Sw., J. Bot. (Schrader)

1800(2): 67. 1802, nom. nov. for Lonchitis

pedata L. 1763, not Pteris pedata L. 1753,

and with the same type.

Lonchitis pedata L., Sp. pi., ed. 2, 1536. 1763. TYPE:

Jamaica, P. Browne (holotype, LINN. 1249.1;

photo, A).

Stem very stout, short-creeping. Leaves to 2 m

or more long. Lamina pedate, 2-5-pinnate-pin-

natifid, basal pinnae much the largest, the central

portion 1-pinnate-pinnatifid. Penultimate seg-

ments shallowly to deeply pinnatifid, ultimate seg-

ments glabrate, minutely pubescent or hirsute

abaxially. Veins copiously areolate, 1 costal areola

with its long axis parallel to the costa between

adjacent costules.

Forests, cloud forests, and forest borders, 700-

3000 m, Amazonas south to Cuzco.

Mexico and Central America; Greater Antilles;

Venezuela and Colombia south to Bolivia.

The sharply serrate apex of the ultimate seg-

ments distinguishes this species from the adven-

tive and also pedate Pteris tripartita.

Amazonas: Prov. Chachapoyas, eastern Calla Calla,

Wurdack 1754 (GH, us). Prov. Bagua, SE of La Peca,

Barbour 3746 (F, MO), 4193 (F). San Martin: Zepelacio,

near Moyobamba, Klug 3495 (F, GH, K, MO, us). Huan-

uco: Prov. Huanuco, Carpish, entre Huanuco y Tingo

Maria, Ferreyra 1831 (GH, USM). Pasco: Oxapampa, (as

Junin), Soukup 2344 (F, GH). Junin: San Ramon, Killip

& Smith 24686 (F, us). Ucayali: Prov. Coronel Portillo,

(as Loreto), Sinchono, cerca a La Divisoria, Aguilar 844

(GH, USM). Cuzco: Prov. La Convencion, Yupanqui to

Rio Apurimac, Davis et al. 1228 (F, GH). San Miguel,

Cook& Gilbert 77 74 (us).

80

FIELDIANA: BOTANY

2 1 . Pteris tripartite Sw., J. Bot. (Schrader) 1 800(2):

67. 1802. TYPE: evidently Java, Thunberg

(holotype, UPS).

23. Pteris vittata L., Sp. pi. 1074. 1753. TYPE:

China, Osbeck (holotype, LINN 1246.3; pho-

to, A).

Stem stout, short-creeping. Leaves 0.5-2 m long.

Lamina pedate, 2-5-pinnate-pinnatifid, basal pin-

nae much the largest, the central portion

1 -pinnate-pinnatifid. Penultimate segments shal-

lowly to deeply pinnatifid, glabrous to somewhat

pubescent abaxially. Veins copiously areolate, 1

costal areola with its long axis parallel to the costa

between adjacent costules.

Forests and partly disturbed forests, ca. 200-

1000 m, Loreto and Ucayali.

Sporadically adventive in the American tropics;

Old World.

The character of the apex of the ultimate seg-

ments that serves to distinguish this from the na-

tive Pteris podophylla has been mentioned under

that species.

Stem stout, short-creeping. Leaves 0.05-1.5 m

or more long. Lamina pinnate, 1 -pinnate, nar-

rowed at the base. Pinnae elongate, simple, entire,

very short-stalked, glabrous or sparsely pubescent

abaxially. Veins free.

In locally moist places, among rocks, and on sea

cliffs, 10-1000 m, six localities in Lima.

Adventive in tropical and subtropical America;

Old World.

The wholly 1 -pinnate lamina with free veins

distinguishes this species. It is sometimes confused

with Pteris longifolia of the West Indies and ad-

jacent regions. That species has articulate pinnae

with the base somewhat enlarged, while in Pteris

vittata the short pinna-stalk departs evenly and

gradually from the rachis.

Loreto: Maynas, near Ramon Castilla, McDaniel 1 1827

(GH). Quebrada Tahuayo, above Tamishiyaco, Croat

1 97 38 (MO). Ucayali: Prov. Coronel Portillo, Dist. Padre

Abad, Gdmez 2 (us).

Lima: Miraflores, Tryon & Tryon 5217 (BM, F, GH, u,

uc, us, USM). Santa Eulalia, in Rimac river, Coronado 2

(GH, uc, us). Choisica, Soukup 2047 (F, us). Km 75,

Carretera Central, Sounders 371 (BM).

22. Pteris cretica L., Mant. pi. 130. 1767. TYPE:

Crete (holotype, LINN 1246.7; photo, A).

Stem small to moderately stout, short-creeping.

Leaves 0. 1 5-1 m long. Lamina pinnate, usually 2-

pinnate at the base, and sometimes beyond, the

pinnae with 1 or rarely more enlarged pinnules,

1 -pinnate beyond the lower or basal pinnae. Pin-

nae (and pinnules) elongate, simple, entire, sessile

to very short-stalked, glabrous or sometimes

slightly pubescent, especially on the costa abaxi-

ally. Veins free.

In forests or in mesic rocky places, ca. 1000-

2200 m, Amazonas.

Adventive in scattered localities in tropical

America; Old World.

The free veins and lamina architecture readily

distinguish this species.

Amazonas: Poma Cocha, Steere (GH). Prov. Chacha-

poyas, 5 km below Chachapoyas, Wurdack 776 (GH, us).

Leimebamba, Woytkowski 7677 (GH, MO). Prov. Chach-

apoyas, Rio Utcubamba valley, Smith & Vdsquez 4974

(GH).

XVI. Acrostichum

Acrostichum L. Sp. pi. 1067. 1753; Gen. pi. ed. 5,

484. 1754. TYPE: Acrostichum aureum L.

Figure 16.

Palustral. Stem erect or decumbent-ascending,

bearing scales. Leaves borne in a cluster, slightly

dimorphic, the fertile pinnae usually narrower and

shorter than the sterile, borne in a cluster, the pet-

iole with many vascular bundles near the base.

Lamina 1 -pinnate, the pinnae entire, glabrous or

somewhat pubescent abaxially, veins anastomos-

ing, without included free veinlets. Sporangia borne

over the surface of the fertile pinnae, on and be-

tween the veins, interspersed with paraphyses.

Spores tetrahedral-globose, trilete, minutely pap-

illate or tuberculate.

Acrostichum is a pantropical genus of three, or

perhaps more, species, with two of them in Amer-

ica and in Peru. It is known in Peru only from salt

or brackish swamps in Tumbes at the southern

limit of its range along the Pacific coast.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

81

FIG. 16. Acrostichum aureum: a, portion of lamina; b, portion of sterile pinna, abaxial side. Acrostichum dan-

aeifolium: c, sterile pinna; d, portion of sterile pinna, abaxial side. (From Stolze, Ferns & fern allies of Guatemala,

1981.)

82

FIELDIANA: BOTANY

References

ADAMS, D. C, AND B. P. TOMLINSON. 1979. Ac-

rostichum in Florida. Amer. Fern. J., 69: 42-46.

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

tichum, pp. 348-354, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Species of Acrostichum

a. Pinnae spaced, glabrous abaxially, coriaceous; fertile pinnae only at the apical portion of the lamina;

apical portion of the petiole with much reduced pinnae or spinelike processes 1. A. aureum

a. Pinnae crowded, usually pubescent abaxially, chartaceous; fertile pinnae extending to the base of the

lamina or nearly so; petiole lacking reduced pinnae or spinelike processes .... 2. A. danaeifolium

1. Acrostichum aureum L., Sp. pi. 1069. 1753.

LECTOTYPE (designated by A. R. Smith, Fl.

Chiapas 2: 16. 1981): Plumier, Descr. pi.

Amer. /. 7. 1693. Figure 16a-b.

Leaves 1-3 m long, the petiole with large, brown,

nearly bicolorous scales at the base, the apical por-

tion with much reduced pinnae or spinelike pro-

cesses, lamina 1 -pinnate, with fertile pinnae only

toward the apex. Pinnae large, simple, entire, usu-

ally well and often irregularly spaced, glabrous

abaxially, coriaceous. Paraphyses mostly with

many lobes.

Among mangroves, near sea level, Tumbes.

Florida; Mexico and Central America; West In-

dies; mostly coastal in South America, south to

northern Peru and Brazil.

Tumbes: Prov. Tumbes, near Rio Tumbes, Simpson

& Schunke 605 (F, GH, us). Margen del Rio Tumbes,

Angulo 596 (HUT).

2. Acrostichum danaeifolium Langsd. & Fisch.,

Icon. fil. 5, /. 7. 1810. TYPE: Brazil, (Santa

Catarina), St. Catharina Island, Langsdorff

(holotype, LE). Figure 16c-d.

Leaves 1.5-4 m long, petiole with large, brown,

nearly bicolorous, often fimbriate, scales at the

base, the apical portion lacking reduced pinnae or

spinelike processes, lamina 1 -pinnate, the fertile

with a few sterile pinnae at the base or none. Pin-

nae large, simple, entire, closely spaced, often im-

bricate, usually somewhat pubescent abaxially,

chartaceous. Paraphyses mostly with few or no

lobes.

Swamps, near sea level, Tumbes.

Florida; Mexico and Central America; West In-

dies; mostly coastal in South America, south to

northern Peru and southern Brazil.

Tumbes: Prov. Tumbes, Corrales, Lopez & Sagdstegui

4034 (GH, HUT).

Family 14: VITTARIACEAE

Vittariaceae (Presl) Ching, Sunyatsenia 5: 232.

1940.

Vittarieae Presl, Tent, pterid. 164. 1836, as Vittari-

aceae. TYPE: Vittaria Sm.

Stem erect to rather long-creeping, bearing scales.

Leaves small to ca. 1 m long, circinate in verna-

tion, monomorphic, usually entire (furcate in He-

cistopteris and 1 -pinnate in Rheopteris of New

Guinea), glabrous. Petiole lacking stipules, not ar-

ticulate to the stem, veins free to usually anasto-

mosing, the areolae without included free veinlets.

Sori abaxial, usually long and more or less sunken,

on a vein or on a submarginal commissure, except

in Anetium with the sporangia mostly in scattered

groups on and between the veins and superficial,

exindusiate. Sporangia with a rather short, 1- or

2-rowed stalk, the annulus vertical, interrupted by

the stalk.

The Vittariaceae are a family of six genera and

about 1 00 species. Most species are epiphytes with

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

83

pendent leaves. The family has clathrate stem scales

and unusual sclerids (spicular idioblasts) in the

epidermis. The key to the genera and to the species

of Vittaria have been adapted from Tryon and

Tryon (1982).

Reference

TRYON, R., AND A. F. TRYON. 1982. Vittari-

aceae, pp. 354-370, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Genera of Vittariaceae

a. Veins free; lamina more or less dichotomously lobed or furcate I. Hecistopteris

a. Veins anastomosing; lamina entire b

b. Sporangia in short to long soral lines along the veins or along a submarginal commissure . . . . c

c. Two-several, or many, soral lines on each side of the costa II. Antrophyum

c. One very long soral line, or line of son, on each side of the costa III. Vittaria

b. Sporangia single, or in scattered groups, on the veins and between them IV. Anetium

I. Hecistopteris

Hecistopteris John Sm., London J. Bot 1: 193.

1842. TYPE: Hecistopteris pumila (Sprengel)

John Sm. (Gymnogramma pumila Sprengel).

Figure 17.

Epiphytic. Stem very small, short-creeping, sca-

ly, the scales clathrate, bearing the leaves in a clus-

ter. Leaves very small, furcate, glabrous, veins free.

Sporangia superficial, borne along a portion of a

vein, indusium absent, paraphyses present. Spores

tetrahedral-globose, trilete, somewhat granulate.

An American genus of one species. The roots

may bear proliferous buds that produce new plants.

1. Hecistopteris pumila (Sprengel) John Sm.,

London J. Bot. 1: 193. 1842. Figure 17.

Gymnogramma pumila Sprengel, Tent, suppl. Syst.

veg. 31. 1828. TYPE: Surinam, Weigelt (holo-

type, not located; isotypes, B!, P!).

Stem scales brown, linear or sublinear, slightly

iridescent. Leaves ca. 1-2 cm long, lamina thin,

narrowly to rather broadly flabellate, variously di-

chotomously or subdichotomously branched or

toothed, gradually tapering at the base. Sporangia

borne in short to rather long lines on the veins.

Epiphyte, or on fallen trunks or branches of trees,

270-625 m, Huanuco and Ucayali.

Tropical America; widespread but not com-

monly collected.

This small species is represented by rather few

specimens throughout its range. This is probably

a reflection of its size rather than of its true dis-

tribution.

Huanuco: Tingo Maria, Allard 20490 (us). Ucayali:

Bosque von Humboldt, Leon 172 (USM), Gentry et al.

36355 (F, MO).

II. Antrophyum

Antrophyum Kaulf., Enum. fil. 197. 1824. TYPE:

Antrophyum plantagineum (Cav.) Kaulf.

(Hemionitis plantaginea Cav.). Figure 18.

Polytaenium Desv., Mem. Soc. Linn. Paris 6: 218.

1827. TYPE: Polytaenium lanceolatum (Sw.)

Desv. (Vittaria lanceolata Sw.) = Antrophyum li-

neatum (Sw.) Kaulf.

Epiphytic. Stem small, short-creeping, scaly, the

scales clathrate, bearing the leaves in a loose clus-

ter. Leaves rather small, the lamina simple, entire,

glabrous, veins anastomosing, forming a few to

many rows of areolae on each side of the costa.

Sporangia more or less sunken, borne in several

long lines, or along most of the veins, indusium

absent, paraphyses present or absent. Spores tet-

rahedral-globose, trilete, or ellipsoidal and mono-

lete, smooth to somewhat echinate.

The Mexican and Central American Antro-

phyum ensiforme Hooker is a member of subgenus

Antrophyum. All other American species belong

to subgenus Polytaenium (Desv.) Benedict, which

84

FIELDIANA: BOTANY

2cm

5 mm

FIG. 17. Hecistopteris pumila: a, habit; b, apical portion of fertile leaf. (From Haught 5328, Colombia, F.)

has the lamina with a costa usually extending to

the apex and lacks paraphyses. In subgenus An-

trophyum a costa is absent or only poorly devel-

oped and paraphyses are present. In spite of these

differences, there are some intermediate species,

and the two groups are not clearly separable.

The generic name was spelled correctly as Po-

lytaenium by Desvaux on p. 1 74 of his publication,

although in the formal place of publication, on p.

218, it was misspelled Polyaetnium.

Antrophyum is a pantropical genus of about 40

species, with 1 8 in America and three in Peru.

References

BENEDICT, R. C. 1 907. The genus Antrophyum —

1. Synopsis of subgenera, and the American

species. Bull. Torrey Bot. Club, 34: 445-458.

— . 1911. The genera of the fern tribe Vit-

tarieae: Their external morphology, venation,

and relationships. Bull. Torrey Bot. Club, 38:

153-190.

TRYON, R. 1 964. Taxonomic fern notes. IV. Some

American vittarioid ferns. Rhodora, 66: 110-

117.

Key to Species of Antrophyum

a. Soral lines immersed between prominent lateral flanges of leaf tissue, mostly long and straight, 2-6,

very rarely 1 , on each side of the costa 1 . A. lineatum

a. Soral lines superficial or somewhat immersed, lacking lateral flanges, more or less following the

anastomosing veins, often short, or if long then rather flexuous and at an angle to the costa . . . . b

b. Lamina elliptical or with nearly parallel sides, usually papyraceous; petiole stramineous abaxially,

narrowly alate 2. A. guayanense

b. Lamina narrowly to broadly oblanceolate, papyraceous to coriaceous; petiole usually dark brown

abaxially, usually strongly alate 3. A. cajenense

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

85

FIG. 18. Antrophyum lineatum: a, habit; b, portion of lamina, abaxial side. (From H. H. Smith 1119, Colom-

bia, F.)

1. Antrophyum lineatum (Sw.) Kaulf., Enum. fil.

199. 1824. Figure 18.

Hemionitis lineata Sw., Prodr. 129. 1788. TYPE: Ja-

maica, Swart: (holotype, not located; isotype,

B!, Herb. Willd. 20033; photo, GH).

Vittaria lanceolata Sw., Ges. Naturf. Freunde Berlin

Mag. Neuesten Entdeck. Gesammten Naturk. 2:

133. 1799, based on Hemionitis lineata Sw., not

Vittaria lineata (L.) Sm.

Polytaenium lanceolatum (Sw.) Desv., Mem. Soc. Linn.

Paris 6: 218. 1827.

Leaves ca. 8-40 cm long. Petiole strongly alate,

stramineous to light green abaxially or drying

darker, nearly concolorous with the adjacent leaf

86

FIELDIANA: BOTANY

tissue. Lamina linear to very narrowly elliptic-lin-

ear, more or less coriaceous. Soral lines few, long

and parallel to the costa.

Epiphyte in woods, forests, and cloud forests,

400-2400 m, Cajamarca and Amazonas to Cuzco.

Tropical America.

The lamina of Antrophyum lineatum is narrow

and tapers gradually to the apex and base. It is

broadest in the central portion where it is usually

1-1.5 cm wide, or not more than 2 cm wide and

then the lamina is more than 25 cm long. In the

other two species the lamina is 3-4 cm wide in

the broadest portion, or somewhat less in small

leaves with the lamina less than 25 cm long.

Cajamarca: Prov. Cutervo, San Andres, Lopez & Sa-

gdstegui 5427 (F). Amazonas: Prov. Bagua, ca. 20 km E

of La Peca, Barbour 2806 A (F, MO). Prov. Chachapoyas,

5 km below Chachapoyas, Wurdack 780 (F, GH, uc, us).

San Martin: Prov. Mariscal Caceres, between Mirador

and La Playa, Young & Leon 4928 (F). Huanuco: Mufia,

Bryan 529 (F, us). Prov. Huanuco, Hacienda Mercedes,

Mexia 8220b (uc). Pasco: Oxapampa, (as Junin), Soukup

2352 (GH). Junin: Near La Merced, Killip & Smith 23859

(us). Ayacucho: Near Kimpitiriki, Killip & Smith 22869

(us). Cuzco: 2.5 km from Machu Picchu, Peyton & Peyton

383 (GH, MO). Prov. La Convencion, Huayopata, Peyton

6 Peyton 891 (GH).

2. Antrophyum guayanense Hieron., Hedwigia 57:

212. 1915. LECTOTYPE (designated by

Tryon, Rhodora 66: 113. 1964): Trinidad,

Fendler 151 (holotype, B!; isotype, GH!).

Polytaenium guayanense (Hieron.) Alston, Bull. Misc.

Inform. 1932: 314.

Leaves ca. 10-30 cm long. Petiole narrowly alate,

stramineous abaxially (or darker in drying but

lighter than the adjacent leaf tissue). Lamina rather

narrowly elliptical or with nearly parallel sides,

subcoriaceous to usually papyraceous. Soral lines

many, following nearly all of the veins.

Epiphyte in forests, 135-800 m, San Martin,

Loreto, and Huanuco.

Southern Mexico; Guianas and Trinidad to Co-

lombia and Peru; adjacent Brazil.

This species has often been called Polytaenium

or Antrophyum cajenense (Tryon, 1964). The lam-

ina shape distinguishes it from that species.

San Martin: Tarapoto, Spruce 3993 (P). Loreto: Rio

Huallaga, below Yurimaguas, Killip & Smith 28951 (GH,

us). Prov. Maynas, Mishana, Rio Nanay, Diaz el al. 425

(MO). Prov. Maynas, Estacion Biologica Callicebus-Mis-

hana, Rio Nanay, Vdsquez et al. 661 (F, MO). Huanuco:

Rio Monzon, near Tingo Maria, Tryon & Tryon 5300

(GH, u, us, USM).

3. Antrophyum cajenense (Desv.) Sprengel, Syst.

veg. 4: 67. 1827.

Hemionitis cajenensis Desv., Ges. Naturf. Freunde

Berlin Mag. Neuesten Entdeck. Gesammten Na-

turk. 5: 311. 1811. TYPE: French Guiana (ho-

lotype, Herb. Desv., P!).

Hemionitis brasiliana Desv., Mem. Soc. Linn. Paris

6:216.1 827. TYPE: Brazil (holotype, Herb. Desv.,

P!; photos, GH, us).

Antrophyum brasilianum (Desv.) C. Chr., Index fil.

59. 1905.

Polytaenium cajenense (Desv.) Benedict, Bull. Torrey

Bot. Club 38: 169. 1911.

Polytaenium brasilianum (Desv.) Benedict, Bull. Tor-

rey Bot. Club 38: 169. 1911.

Leaves ca. 10-35 cm long. Petiole usually

strongly alate, stramineous to greenish abaxially

(drying to light or usually dark brown), concol-

orous with, or darker or lighter, than the adjacent

leaf tissue. Lamina narrowly to broadly oblanceo-

late, papyraceous to usually coriaceous. Soral lines

many, following nearly all of the veins.

Epiphyte in dense forests, 100-2040 m, Ama-

zonas to Cuzco.

Southern Mexico to Panama; Greater Antilles;

Guianas to Colombia, south to Bolivia; Brazil.

Antrophyum brasilianum is so similar to A. ca-

jenense that it is evidently a variation of that

species. The oblanceolate lamina is characteristic

of A. cajenense.

Amazonas: Prov. Bagua, Quebrada Mirana, Wurdack

1916 (F, GH, us). San Martin: Ad Saposoa, Woytkowski

7322 (GH, MO, us). Prov. Mariscal Caceres, Tocache

Nuevo, J. Schunke V. 7589 (F, MO). Loreto: Gamitani-

cocha, Rio Mazan, Schunke 283 (F, GH, us, USM). Huan-

uco: Tingo Maria, Ferreyra 10230 (GH, USM), Woyt-

kowski 1097 (GH, USM). Pasco: Paujil, near Puente

Bermudez, Ledn 289 (USM). Ucayali: Prov. Coronel Por-

tillo, Rio Chino, J. Schunke V. 9204 (MO). Cuzco: Ha-

cienda Potrero, near Quillabamba, Coronado 120 (GH,

us). Prov. La Convencion, Tupitari, Vargas 3446 (uc).

Comments

Antrophyum lanceolatum (L.) Kaulf., Enum. til.

198. 1824.

Hemionitis lanceolata L. Sp. pi. 1077. 1753. LEC-

TOTYPE (designated by Proctor, Ferns of Ja-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

87

FIG. 19. Vinaria graminifolia: a, habit; b, portion of lamina, abaxial side; c, stem scale. Vittaria lineata: d, stem

scale. Vittaria stipitata: e, stem scale. (From Stolze, Ferns & fern allies of Guatemala, 1981.)

88

FIELDIANA: BOTANY

maica 258. 1985): St. Vincent, Plumier, Traite

foug. Amer, t. 127, f. C. 1705.

This species is reported from northern South

America and may occur in Peru. It has a long, very

narrow lamina that gradually tapers to the apex

and base, as in Antrophyum lineatum, but has su-

perficial soral lines lacking lateral flanges.

III. Vittaria

Vittaria Sm., Mem. Acad. Roy. Sci. (Turin) 5:413.

1793. TYPE: Vittaria lineata (L.) Sm. (Pteris

lineata L.). Figure 19.

Ananthacorus Underw. & Maxon, Contr. U.S. Natl.

Herb. 10: 487. 1908. TYPE: Ananthacorus an-

gustifolius (Sw.) Underw. & Maxon (Pteris an-

gustifolia Sw.) = Vittaria costata Kunze.

Epiphytic. Stem small, erect or short-creeping,

scaly, the scales clathrate, bearing the leaves in a

cluster or somewhat spaced. Leaves small or of

medium size, the lamina simple, entire, glabrous,

veins anastomosing, forming 1 row, or 2 to a few

rows, of areolae on each side of the costa. Spo-

rangia borne, on each side of the costa, in a con-

tinuous, more or less sunken line along a sub-

marginal commissure, indusium absent,

paraphyses present. Spores tetrahedral-globose,

trilete, or ellipsoidal and monolete, smooth or

slightly granulate.

A pantropical genus of 50 or more species, with

perhaps 10 in America, and eight in Peru. The

sometimes accepted genus Ananthacorus, repre-

sented by Vittaria costata, differs only in the ve-

nation of two or a few rows of areolae on each side

of the costa, rather than a single row as in other

species of Vittaria.

Some of the characters of the species are pro-

vided in the descriptions; other special characters

are mentioned in the discussions.

References

BENEDICT, R. C. 1914. A revision of the genus

Vittaria J. E. Smith. I. The species of the sub-

genus Radiovittaria. Bull. Torrey Bot. Club, 41:

391^*10.

TRYON, R. 1964. Taxonomic Fern Notes, IV.

Some American vittarioid ferns. Rhodora, 66:

110-117.

Key to Species of Vittaria

a. Lamina with 2 or more rows of areolae on each side of the costa 1 . V. costata

a. Lamina with a single row of areolae on each side of the costa b

b. Petiole wholly or mostly narrowly alate, flattened throughout, firm to rather soft, often irregularly

wrinkled when dry c

c. Paraphyses slender, the apical cell not or hardly enlarged; soral lines in deep grooves, more or

less covered by a flange of leaf tissue on each side; stem scales usually with a long, uniseriate,

filiform tip 2. V. lineata

c. Paraphyses with an enlarged, usually dark-colored apical cell d

d. Petiole lighter in color than the green lamina or concolorous with the brownish lamina when

darkened by age or drying 3. V. graminifolia

d. Petiole darker than the lamina, usually dark brown to atropurpureous e

e. Stem scales mostly dark brown to atropurpureous, definitely clathrate with dark, sclerotic

lateral walls; lamina narrow or gradually tapering to the apex f

f. Lamina narrowly elliptical to linear-elliptical, often falcate 4. V. gardneriana

f. Lamina narrowly linear or narrower, rather straight 5. V. ruiziana

e. Stem scales light brown, subclathrate; lamina elliptical, rather abruptly reduced to the

apex 6. V. latifolia

b. Petiole not alate, usually terete or oval, or flattened only at the base, indurated g

g. Stem scales mostly 6 or more cells wide; each soral line in a shallow groove

7. V. moritziana

g. Stem scales 2 cells wide or sometimes 3 cells wide at the base; each soral line in a deep groove

8. V. stipitata

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

89

1. Vittaria costata Kunze, Linnaea 9: 77. 1834.

TYPE: Peru, (San Martin), Huallaga, To-

cache, Jun. 1830, Poeppig (holotype, LZ de-

stroyed; isotype, w!). At B!, Kunze, Analecta.

pteridogr. /. 18, f. 2 is mounted in the her-

barium and has the same data as the holotype

written on it; probably the figure was prepared

from the holotype at LZ.

Pteris angustifolia Sw., Prodr. 129. 1788. TYPE: Ja-

maica and Hispaniola (Fl. Ind. Occid. 3: 1599),

Swartz (holotype, not located), not Vittaria an-

gustifolia Blume, Enum. pi. Javae 199. 1828.

Ananthacorus angustifolius (Sw.) Underw. & Maxon,

Contr. U.S. Natl. Herb. 10: 487. 1908.

Stem scales brown to atropurpureous, usually

short-ciliate. Leaves ca. 10-50 cm long, lamina

narrow, tapering gradually to the apex and base.

Petiole narrowly alate, rather flattened, firm, dark-

er than, or nearly concolorous with the adjacent

lamina tissue when dry. Soral lines sunken, con-

tinuous or irregularly discontinuous.

Epiphyte in forests, 500-650 m, San Martin,

Huanuco, and Cuzco.

Tropical America.

This species is characterized by two or more

rows of areolae on each side of the costa. All of

the other species have a single row.

San Martin: Prov. Mariscal Caceres, Puerto Pizana,

J. Schunke V. 4600 (F, us). Prov. Mariscal Caceres, Dist.

Tocache Nuevo, J. Schunke V. 7797 (MO). Huanuco:

Supte river, N of Tingo Maria, Stork & Morton 9598 (F,

uc, us). Junction of Rio Monzon and Rio Huallaga,

Asplund 12678 (us). Cuzco: Prov. Quispicanchi, entre

Machiche y Inambari, Vargas 16456 (GH).

2. Vittaria lineata (L.) Sm., Mem. Acad. Roy. Sci.

(Turin) 5: 421. 1793. Figure 19d.

Pteris lineata L., Sp. pi. 2: 1073. 1753. TYPE (des-

ignated by Tryon, Contr. Gray Herb. 194: 213.

1964): Santo Domingo. Plumier, Traite foug.

Amer. /. 143.

Vittaria filiformisCav., Descr. pi. 270. 1802. TYPE:

Peru, Nee , MA; photos, F, NY, us; seen by C. Chr.

(Dansk Bot. Ark. 9(3): 24. 1937) and placed here

with some doubt, the type has no stem or petiole

base.

Stem scales dark brown to atropurpureous,

shortly dentate. Leaves ca. 10-100 cm long, lam-

ina very narrow. Petiole narrowly alate, rather flat-

tened, firm, lighter than the lamina or concolorous

with it. Soral lines sunken, continuous.

Epiphyte in forests and cloud forests, or on rot-

ting tree trunks, 1 00-2600 m, Amazonas to Junin.

Tropical America.

The stem scales have unusually long, uniseriate

tips and the soral lines are in deep grooves with a

pronounced laminar flange on each side. The light-

colored paraphyses with the apical cell not or hard-

ly enlarged also serve to distinguish this species.

The combination V. lineata is sometimes cred-

ited to Swartz (Syn. fil. 109. 1806); however, J. E.

Smith cited Pteris lineata L. under Vittaria and

Vittaria lineata in the caption to Figure 5.

Amazonas: Prov. Bagua, 25 km E of La Peca, Barbour

2976 (MO). Serrania de Bagua, Gentry et al. 22895 (F,

MO, us). San Martin: Tarapoto, Ll. Williams 6321 (F,

us). Prov. Mariscal Caceres, Puerto Pizana, J. Schunke

V. 4695 (F, GH, MO, us). Loreto: Prov. Maynas, Rio Itaya,

Revilla et al. 2590 (F, MO). Near mouth of Rio Itaya,

Croat 19670 (F, uc). Huanuco: Tingo Maria, Asplund

12356 (us), Croat 19670 (F, uc). Pasco: Prov. Oxapam-

pa, Rio San Alberto Valley, Smith & Pretel 8036 (F).

Junin: La Merced, Weberbauer 1914 (B).

3. Vittaria graminifolia Kaulf., Enum. fil. 192.

1824. TYPE: Brazil (holotype, LZ destroyed;

isotype, Herb. Greville, Kaulfuss misit, 1827,

E!). Figure 19a-c.

Vittaria filifolia Fee, Mem. foug. 3: 20, t. 3,f. 6. 1852.

LECTOTYPE (designated by Tryon, Rhodora 66:

114. 1964): Guadeloupe, L'Herminier, Herb.

Cosson, P!.

Stem scales brown, shortly dentate or ciliate or

not. Leaves ca. 10-50 cm long, lamina very nar-

row. Petiole usually narrowly alate, rather flat-

tened, soft, lighter in color than the lamina or

concolorous with it dry. Soral lines hardly sunken,

continuous.

Epiphyte in forests, in cloud forests, sometimes

on fallen trunks or on cliffs, 1200-3100 m, Caja-

marca to Cuzco.

Tropical America.

Vittaria graminifolia may be distinguished from

all other Peruvian species by its trilete spores; the

others have monolete spores.

Cajamarca: Prov. Contumaza, Cascas Contumaza,

Lopez et al. 9088 (GH). Prov. San Miguel, Taulis Re-

corco, Mostacero et al. 1 178 (F). Amazonas: 1-2 km W

of Molinopampa, Wurdack 1491 (F, GH, uc, us). Serrania

de Bagua, Gentry et al. 23017 (F, MO). Huanuco: Prov.

Leoncio Prado, Dist. Hermilio Valdizan, Plowman &

Schunke 1 1744 (F). Junin: La Merced, Macbride 5637

90

FIELDIANA: BOTANY

(F, us). Cuzco: Prov. Urubamba, Machu Picchu, lids el

al. 1097 (GH).

4. Vittaria gardneriana Fee, Mem. foug. 3: 15, /.

3,f. 1. 1852. LECTOTYPE (Designated by

Benedict, Bull. Torrey Bot. Club 41: 401.

1914): Brazil, Gardner 147 (holotype, not lo-

cated; isotypes, B!, BM!, K!, P!, us!; photo, GH

ofB).

Vittaria remota Fee, Mem. foug. 7: 26, /. 20. f. 1.

1857. TYPE: Colombia, (Norte de Santander),

Prov. Ocana, Schlim 611 (holotype, not located).

Stem scales brown to atropurpureous, shortly

dentate or long-ciliate. Leaves ca. 5-30 cm long,

lamina linear-elliptical or somewhat broader,

gradually tapering to the apex and base, often fal-

cate. Petiole narrowly alate, flattened, rather soft,

darker than the lamina. Soral lines hardly sunken,

continuous.

Epiphyte in dense forests, in cloud forests, or on

fallen, rotting tree trunks, 850-2400 m, Amazonas

to Cuzco.

Costa Rica and Panama; Greater Antilles; Sur-

inam to Colombia, south to Bolivia; Brazil.

Although sometimes recognized as a distinct

species, Vittaria remota grades into V. gardner-

iana. The venation characters, among others, em-

ployed to separate the two are not constant in the

newer collections studied. Some specimens with a

rather broad lamina have been misidentified as V.

latifolia, but the lamina gradually tapers to the

apex as in V. gardneriana rather than being rather

abruptly reduced as in V. latifolia.

Amazonas: Prov. Bagua. Cordillera Colan, Barbour

3977 (MO, USM), 3596 (MO). Huanuco: Cerros del Sira,

Dudley 13050 (GH), 13221 A (GH). Pasco: Yapas, Pichis

Trail, (as Junin), Killip & Smith 25552 (F, GH, us). Prov.

Oxapampa, van der Werffet al. 8331 (MO, uc), 8424 (MO,

uc). Cuzco: Prov. Urubamba, Machu Picchu, Peyton &

Peyton 1303b (GH, MO). Prov. Paucartambo, Aguada de

Asuncion, Vargas 7353 (uc).

5. Vittaria ruiziana Fee, Mem. foug. 3: 16, /. 3,f.

3. 1852. TYPE: Peru, Ruiz (holotype, not lo-

cated; authentic specimen, Peru. Dombey, det.

Fee, B!; photo, GH).

Pteropsis vittarioides Desv., Mem. Soc. Linn. Paris, 6:

219. 1827. TYPE Peru, (holotype, Herb. Desv.,

P!, a single leaf evidently taken from [Junin] Hu-

asi-Huasi, 1779, Dombey P!). A duplicate of Dom-

bey is in Herb. Kunth, B!; photo, GH).

Vittaria vittarioides (Desv.) Weath., Contr. Gray Herb.

1 14: 34. 1936, not (Thouars) C. Chr., Index fil.

655. 1907.

Stem scales dark brown to atropurpureous,

shortly dentate to entire. Leaves ca. 15-100 cm

long, lamina narrow to very narrow, rather straight.

Petiole mostly narrowly alate, flattened, firm, usu-

ally darker than the lamina. Soral lines sunken or

hardly so, continuous.

Epiphyte in forests or rarely on rocks, 1800-

3450 m, Amazonas to Cuzco.

Venezuela to Colombia, south to Bolivia.

The lamina is narrowly linear or narrower and

rather straight, in contrast to that of the previous

species Vittaria gardneriana in which the lamina

is narrowly elliptical to linear-elliptical and often

falcate.

Amazonas: Cerros Calla Calla, Hutchison & Wright

5688 (F, GH). San Martin: Prov. Mariscal Caceres, Rio

Abiseo, Young 1622 (USM). Huanuco: Near Muna, Mac-

bride 4136 (B, F, GH, us). Cuzco: Prov. Paucartambo,

near Achirani, Vargas 1 1 157 (F, GH, us). Cerro Chuyapi,

Bues A44 (GH, us).

6. Vittaria latifolia Benedict, Bull. Torrey Bot.

Club 41: 403, t. 17. 19 14. Type: Bolivia, (Franz

Tamayo), Santa Barbara, R. S. Williams 1337

(holotype, NY!; isotypes, GH!, us!).

Stem scales light brown, long-ciliate. Leaves ca.

5-30 cm long, lamina elliptical, rather abruptly

reduced to the apex, gradually narrowed to the

base. Petiole narrowly alate, flattened, firm, usu-

ally darker than the lamina. Soral lines hardly

sunken, continuous.

Pasco.

Peru and Bolivia.

Although this rare species is rather close to Vit-

taria gardneriana, it is distinguished by the ellip-

tical lamina that is rather abruptly reduced to the

apex. The scales are subclathrate and their light

brown color sometimes also occurs in V. gard-

neriana, as do the long-ciliate scale margins.

Pasco: Oxapampa, (as Junin), Soukup 2351 (GH).

7. Vittaria moritziana Mett., Ann. Sci. Nat. Bot.

V, 2: 207. 1864. TYPE: Colombia, Canoas,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

91

Lindig 319 (holotype, LZ destroyed; isotype,

B!).

Stem scales dark brown to atropurpureous,

mostly shortly dentate. Leaves ca. 20- 100 cm long,

lamina narrow. Petiole terete or oval, or flattened

only at the base, not alate, indurated. Soral lines

hardly sunken, continuous.

Epiphyte in forests, 2400-3100 m, Amazonas

and Pasco.

Costa Rica; Hispaniola; Venezuela to Colombia

and south to Bolivia.

Amazonas: Prov. Chachapoyas, 2 km W of Molino-

pampa, Wurdack 1480 (F, uc, us). Prov. Chachapoyas,

Calla Calla slopes, Wurdack 1344 (GH, us). Pasco: Prov.

Oxapampa, Cordillera Yanachaga, D. Smith 7665 (F).

8. Vittaria stipitata Kunze, Linnaea 9: 77. 1834.

TYPE: Peru, (Huanuco), Pampayacu, Jul.

1829, Poeppig (Diar. 1121) (holotype, LZ de-

stroyed; evidently illustrated in Kunze, An-

alecta pteridogr. t. 18, f. 1; isotype, P!; au-

thentic specimen, Poeppig, B!, K!; photo, GH

of B). Figure 19e.

Stem scales dark brown to atropurpureous,

somewhat shortly dentate to entire. Leaves ca. 10-

75 cm long, lamina narrow. Petiole terete or oval,

or flattened only at the base, not alate, indurated.

Soral lines sunken, continuous.

Epiphyte in forests and cloud forests, rarely on

rocks, 600-2000 m, Amazonas to Cuzco.

Guatemala to Panama; Greater Antilles; Ven-

ezuela to Colombia and south to Bolivia; Brazil.

This species and the previous, Vittaria morit-

ziana, are characterized by their indurated, terete

or oval and nonalate, usually dark-colored petiole.

They may be distinguished by their stem scales,

which are short and mostly two cells wide in V.

stipitata, while they are longer and mostly six or

more cells wide in V. moritziana.

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

2566 (F, MO, uc). Mendoza, Woytkowski 8312 (GH, MO,

uc). San Martin: Mt. Guayrapurina, near Tarapoto,

Spruce 4773 (K,P). Huanuco: Tingo Maria, Allard 21932

(us). Pasco: Prov. Oxapampa, Chequitavo, D. Smith 5184

(GH). Junin: Above San Ramon, SchunkeA146 (GH, us),

Killip & Smith 24587 (F, us). Cuzco: Prov. La Conven-

cion. Ichiquiato. Vargas 14492 (GH).

IV. Anetium

Anetium (Kunze) Splitg., Tijdschr. Natuurl. Gesch.

Physiol. 7: 395. 1840. Figure 20.

Acrostichum sect. Anetium Kunze, Beibl. Flora 1839:

1: 47. TYPE: Acrostichum citrifolium L. = Ane-

tium citrifolium (L.) Splitg.

Pteridanetium Copel., Gen. fil. 224. 1947, based on

Anetium Splitg., not Anetia Endl. which is not a

homonym as Copeland considered it to be.

Epiphytic. Stem slender, rather long-creeping,

scaly, the scales clathrate, bearing the leaves at

intervals. Leaves small to rarely large, the lamina

simple, entire, glabrous, veins anastomosing,

forming many rows of areolae on each side of the

costa. Sporangia borne superficially and often

sparingly along the veins and also between them,

indusium and paraphyses absent. Spores tetra-

hedral-globose, trilete, with irregular strands.

Anetium is an American genus of one species.

1. Anetium citrifolium (L.) Splitg., Tijdschr. Na-

tuurl. Gesch. Physiol. 7: 395. 1840. Figure

20.

Acrostichum citrifolium L.. Sp. pi. 1067. 1 753. TYPE:

Martinique, Plumier, Traite foug. Amer. t. 116.

1705.

Pteridanetium citrifolium (L.) Copel., Gen. fil. 224.

1947.

Stem scales brownish, mostly broadly ovate-at-

tenuate, iridescent. Leaves herbaceous-fleshy (pa-

pyraceous in drying), pendent, 10 to usually 15-

30 to 100 cm long, the lamina elliptical to oblan-

ceolate to ligulate, narrow to broad, nearly or quite

sessile to short-petioled (especially in large leaves),

apex obtuse to acute or abruptly acuminate.

Epiphyte in dense forests, on fallen tree trunks,

or rarely on wet rocks, 1 00-800 m, San Martin to

Cuzco and Madre de Dios.

Tropical America.

San Martin: Prov. Mariscal Caceres, Quebrada de

Challuayacu, J. Schunke V. 10752 (GH, MO, uc). Prov.

Mariscal Caceres, Nuevo Progresso, J. Schunke V. 3162,

3203 (F, GH, us). Loreto: Puerto Arturo, Rio Huallaga.

below Yurimaguas, Killip & Smith 27900 (F, GH, us).

Rio Mazan, Schunke 301 (F, GH, uc, us). Huanuco: Rio

Mon/an. near Tingo Maria, Tryon & Tryon 5300'/i (GH).

Cuzco: Rio Apurimac, below Puerto Capiro, Davis et al.

1290 (GH). Madre de Dios: Prov. Tambopata, Leon &

Bahorquez888 (USM). Alto Madre de Dios, Rauh & Hirsch

1644 (USM).

92

FIELDIANA: BOTANY

FIG. 20. Anetium citrifolium: a, habit; b, portion of lamina near base, adaxial side; c, portion of fertile lamina,

abaxial side; d, stem scale. (Adapted from Stolze, Ferns & fern allies of Guatemala, 1981.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

93

Family 15: DENNSTAEDTIACEAE

Dennstaedtiaceae Pic.-Ser., Webbia 24: 704. 1 970.

TYPE: Dennstaedtia Moore.

Stem erect, to short- or very long-creeping, bear-

ing trichomes or scales or both. Leaves small or

to 7 m long, circinate in vernation, monomorphic

or rarely somewhat dimorphic, usually pinnate,

rarely simple, glabrous, glandular, pubescent, or

rarely slightly scaly. Petiole lacking stipules, not

articulate to the stem. Veins free to fully anasto-

mosing, the areolae without included free veinlets.

Sori marginal, submarginal, or rarely abaxial, at

a vein end or on a vascular commissure connecting

vein ends, indusiate, the indusium cup- or purse-

shaped, or formed by the modified recurved mar-

gin, or abaxial, or (in Hypolepis) sometimes ex-

indusiate. Sporangia with a short to long, 1-3-

rowed stalk, annulus vertical to slightly oblique,

the indurated portion interrupted by the stalk.

The Dennstaedtiaceae are a family of about 20

genera and 175 species. It is a widely distributed

family through the World and, although basically

pantropical, some elements are in boreal or south

temperate regions. The family is diverse in its

morphology and cytology and therefore is evi-

dently old. It may be characterized by long-creep-

ing stems bearing trichomes and large, decom-

pound leaves, indusiate and marginal or

submarginal sori, and the frequent presence of epi-

petiolar buds.

The key to genera has been adapted from Tryon

and Tryon (1982).

Reference

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

nstaedtiaceae, pp. 370-432, in Ferns and allied

plants, Springer- Verlag, New York.

Key to Genera of Dennstaedtiaceae

a. Indusium cup- or purse-shaped, the adaxial and abaxial portions joined II. Dennstaedtia

a. Indusium abaxial, the opposed margin not or hardly modified, flat or nearly so; or indusium formed

by the modified margin or a separate abaxial indusium may also be present; or indusium absent . .

b

b. Indusium abaxial, the opposed margin not or hardly modified, flat or nearly so c

c. Stem with trichomes and no scales I. Microlepia

c. Stem with scales or sometimes also with trichomes d

d. Each sorus served by 1 vein; indusium nearly cuneate, attached on both sides

III. Saccoloma

d. Each sorus elongate, served by 2 or more veins, these connected by a vascular commissure;

indusium elongate, attached on the inner side X. Lindsaea

b. Indusium formed by the modified margin, or a separate abaxial indusium may also be present;

or indusium absent e

e. Each sorus served by 1 vein VI. Hypolepis

e. Each sorus served by few to many veins f

f. Sterile segments and sterile portions of fertile segments with a modified margin similar to

the indusium but not as broad IV. Pteridium

f. Only the fertile portion of a segment with a modified marginal indusium g

g. Abaxial indusia present V. Paesia

g. Abaxial indusia absent h

h. Lamina more or less pubescent, not glaucous abaxially i

i. Stem decumbent to erect, indurated; veins fully anastomosing . . . VII. Blotiella

i. Stem short-creeping, succulent; veins free or casually anastomosing

VIII. I onchit is

h. Lamina glabrous or nearly so, usually glaucous abaxially; stem long-creeping, slender,

indurated IX. Histiopteris

94

FIELDIANA: BOTANY

I. Microlepia

Microlepia Presl, Tent, pterid. 124. 1836. TYPE:

Microlepia polypodioides (Sw.) Presl (Dick-

sonia polypodioides Sw.) = Microlepia spe-

luncae (L.) Moore. Figure 21.

Dennstaedtia Bernh., J. Bot. (Schrader) 1800(2): 124.

1802. TYPE: Dennstaedtia flaccida (Forster)

Bernh. (Trichomanes flaccida Forster) = Micro-

lepia flaccida (Forster) Fee.

Terrestrial. Stem usually slender, long-creeping,

bearing trichomes or rarely bristles. Leaves usually

widely spaced, to 7 m long, sometimes scandent

on other vegetation. Lamina 1-4-pinnate, usually

pubescent, veins free. Sori abaxial to nearly mar-

ginal, sometimes paraphysate, covered by an

abaxial indusium that is attached at its sides or

base, the leaf tissue beyond the sorus unmodified.

Spores tetrahedral-globose, trilete, the surface finely

echinate.

The nomenclature of this genus is discussed un-

der Dennstaedtia. It is not known from Peru, but

the single American species Microlepia speluncae

undoubtedly occurs there, since it is known from

Ecuador and Bolivia. It should be sought in Peru

in moist or wet forests on mountain slopes.

1 . Microlepia speluncae (L.) Moore, Index til. xciii.

1857. Figure 21.

Polypodium speluncae L., Sp. pi. 1093. 1753. TYPE:

Ceylon, Hermann (holotype, BM).

Stem trichomes rather soft, several cells long,

mostly catenate. Leaves ca. 0.5-1.5 m long, long-

petioled. Lamina lanceolate to broadly lanceolate,

pinnae broadest at or near the base, acuminate,

softly pubescent beneath and less so above. In-

dusium pubescent.

Greater Antilles; widely scattered in South

America; Old World.

The only species in America; to be expected in

Peru.

The sorus and indusium of Microlepia speluncae

are similar to those in Saccoloma inaequale, but

M. speluncae has a creeping stem bearing tri-

chomes, while S. inaequale has an erect stem bear-

ing scales. The lamina architecture and pubescence

of M. speluncae are similar to those of Dennstaed-

tia cicutaria but the sori of M. speluncae are back

of, or close to, the margin, while in D. cicutaria

they are truly marginal.

II. Dennstaedtia

Dennstaedtia Moore, Index fil. xcvii. 1859. TYPE:

Dennstaedtia cicutaria (Sw.) Moore (Dickson-

ia cicutaria Sw.). Figure 22.

Sitobolium Desv., Mem. Linn. Soc. Paris 6: 262. 1 827.

TYPE: Sitobolium punctilobulum (Michx.) Desv.

= Dennstaedtia punctilobula (Michx.) Moore.

Stem usually slender and widely creeping, bear-

ing trichomes. Leaves large to very large, to 7 m

long, sometimes scandent on other vegetation.

Lamina 1 -pinnate to 4-pinnate-pinnatifid, gla-

brous or pubescent, veins free. Sori marginal,

sometimes paraphysate, the adaxial and abaxial

indusia joined to form a cylindrical, globose, or

purse-shaped indusium that is bilabiate or not.

Spores tetrahedral-globose, trilete, the surface ver-

rucate, tuberculate, reticulate, or ridged.

Dennstaedtia is a tropical and extratropical ge-

nus of about 45 species, with 12 in America and

all but two of them in Peru: Dennstaedtia distenta

of Mexico, Central America, and the Greater An-

tilles, and D. punctilobula of eastern North Amer-

ica.

The name Dennstaedtia is used here from a later

publication than that of Bernhardi (see Microlepia)

and with a different type, as in Tryon and Tryon

(1982). The type species of Dennstaedtia Bernh.

is D. flaccida, which has the echinate spores of

Microlepia. We are unwilling to make about 90

new combinations and to designate Microlepia as

Dennstaedtia and Dennstaedtia as Sitobolium, as

would be required by the International Code of

Botanical Nomenclature.

References

TRYON, R. 1960. A review of the genus Den-

nstaedtia in America. Contr. Gray Herb., 187:

23-52.

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

nstaedtia, pp. 377-383, in Ferns and allied

plants, Springer- Verlag, New York.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

95

5 mm

1mm

FIG. 2 1 . Microlepia speluncae: a, pinna; b, penultimate segment, abaxial side; c, laminar trichomes. (From Clute

227 , Jamaica, F.)

96

FIELDIANA: BOTANY

FIG. 22. Dennstaedtia dissecta: a, stem and one pinna; b, base of pinnule, abaxial side. Dennstaedtia bipinnata:

c, portion of rachis and pinna-rachis, adaxial side. Dennstaedtia globulifera: d, base of pinna, abaxial side. (Adapted

from Stolze, Ferns & fern allies of Guatemala, 1981.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

97

Key to Species of Dennstaedtia

a. Axis of the penultimate segments on the adaxial surface bordered on each side by a pronounced

herbaceous wing perpendicular to the plane of the segment, the wing on the basiscopic side decurrent

onto the axis of the next order, either as an herbaceous wing or as a pronounced ridge; trichomes on

the abaxial surface of the pinnules wholly clear brown or tan (rarely subopaque or whitish) b

b. Basal segments of the pinnules of the central pinnae definitely alternate, quite unequal in size, the

inferior one ascending to strongly ascending; indusium cylindrical to subcylindrical, rarely globular;

lamina lanceolate-ovate to ovate 3. D. bipinnata

b. Basal segments of the pinnules of the central pinnae usually subopposite to nearly opposite, rather

or quite equal in size, the inferior one not or slightly ascending; indusium globular or less often

subglobular or rarely cylindrical; lamina deltoid 4. D. globulifera

a. Axis of the penultimate segments on the adaxial surface lacking perpendicular herbaceous wings, or

these present but the one on the basiscopic side not decurrent onto the axis of the next order; tri-

chomes on the abaxial surface of the pinnules, when present, whitish to brownish and usually sub-

opaque c

c. Sterile vein tips on the adaxial surface slender; pinnules abaxially glabrous or with sparse to dense,

rather spreading trichomes d

d. Many or most of the sori borne in a sinus; pinnules abaxially more or less pubescent; lamina

deltoid 1 . D. cicutaria

d. All or most of the sori terminal on a lobe; pinnules abaxially glabrous or nearly so; lamina

ovate- to deltoid-lanceolate 2. D. glauca

c. Sterile vein tips on the adaxial surface enlarged, clavate to punctate (rarely slender in D. dissecta)\

pinnules abaxially glabrate to usually with subappressed, ascending trichomes e

e. Lamina 1 -pinnate-pinnatifid or more complex f

f. Sterile veins ending well away from the glabrous margin g

g. Pinnules 1 -pinnate to 1 -pinnate-pinnatifid; apical segments of the pinnae separate or

nearly so to the prolonged tip, the separate ones closest to the tip obtuse

5. D. dissecta

g. Pinnules entire to deeply pinnatifid; apical segments of the pinnae confluent back of the

prolonged tip, the separate ones closest to the tip acute 6. D. arborescens

f. Sterile veins nearly reaching the persistently pubescent margin 7. D. spruce!

e. Lamina 1 -pinnate, the pinnae entire 8. D. wercklei

1 . Dennstaedtia cicutaria (Sw.) Moore, Index fil.

xcvii. 1857.

Dicksonia cicutaria Sw., J. Bot. (Schrader) 1800(2):

91.1801. TYPE: Jamaica, Swart z (holotype, s!;

frag., us!; photo, us!; isotype, B!, Herb. Willd.

20156; photo, GH).

Dicksonia rubiginosa Kaulf., Enum. fil. 226. 1824.

TYPE: Brazil, (Guanabara), Rio de Janeiro, Mer-

tens (holotype, LE!; photo, GH).

Dennstaedtia rubiginosa (Kaulf.) Moore, Index fil.

xcvii. 1857.

Leaves ca. 1—4 m long. Lamina deltoid, 3-pin-

nate to 4-pinnate-pinnatifid. Axes of the penulti-

mate segments without herbaceous ridges or wings

on the adaxial surface, sterile vein tips on the adax-

ial surface slender, ending well away from the gla-

brous to sparingly pubescent margin. Pinnules

more or less pubescent adaxially and abaxially,

with whitish to brownish usually dimorphic tri-

chomes, some 1 -celled, short, acicular, rigid, oth-

ers (rarely the only kind) multicellular, longer,

sometimes lax and subtortuous. Sori mostly borne

in a sinus, mature indusia saucer- to purse-shaped.

In dense or open forests, at the borders of clear-

ings, along roads through forests, 200-1700 m,

Amazonas south to Puno.

Central Mexico to Panama; Greater Antilles;

Venezuela and Colombia to Bolivia and southern

Brazil.

In addition to the characters mentioned in the

key, this species and the following one, Dennstaed-

tia glauca, may often be separated by the color of

the major axes. In D. cicutaria they are usually

brownish, and in D. glauca they are usually straw-

colored.

98

FIELDIANA: BOTANY

Amazonas: Prov. Bagua, ca. 5 km S of La Peca, Bar-

hour 4312 (MO). San Martin: Near Tarapoto, Spruce

4338 (GH, K.). Zepelacio, near Moyobamba, Klug 3553

(F, GH, K, us). Loreto: Near mouth of Rio Santiago, Mexia

6128 (BM, F, GH, K, MO, uc, us). Huanuco: Dist. Chu-

rubamba, Cotirarda, Mexia 8218 (F, GH, MO, uc, us).

Pozuzo, Macbride 4605 (F, us). Pasco: Quillasu, Soukup

3513 (GH, us). Junin: La Merced, Killip & Smith 23699

(F, GH, us). Satipo, Aug., 1940, Ridoutt (GH, us). Ayac-

ucho: Estrella, between Huanta and Rio Apurimac, Killip

& Smith 22440 (GH, us). Cuzco: Potrero, 8 km W of

Quillabamba, Tryon & Tryon 5372 (BM, F, GH, u, us,

USM). Puno: Churumayo, Soukup 871 (F). Prov. Cara-

baya, Vargas /6/29(GH).

2. Dennstaedtia glauca (Cav.) Looser, Rev. Hist.

Geog. Chile 69: 184. 1932.

Davallia glauca Cav., Descr. pi. 278. 1802. TYPE:

Chile, Cordillera de Planchon, Nee (holotype, MA;

frag., BM!; Looser, op. cit., and C. Chr. in Dansk.

Bot. Ark. 9(3): 28. 1937, discuss the identity of

the type).

Leaves ca. 0.5-2 m long. Lamina ovate- to del-

toid-lanceolate, 3-4-pinnate-pinnatifid. Axes of the

penultimate segments with an herbaceous ridge on

the adaxial surface, sterile vein tips, on the adaxial

surface, slender, ending well away from the gla-

brous margin. Pinnules glabrous or nearly so. Sori

predominantly terminal on lobes, mature indusia

usually purse-shaped.

In moist, shrubby or rocky ravines and on banks,

2900-3700 m, Cuzco and Puno.

Chile, northeast to adjacent Argentina, to Bo-

livia and southern Peru.

Specimens of Dennstaedtia cicutaria that are only

slightly pubescent may be confused with D. glau-

ca. All of the characters mentioned in the key

under heading d must be employed for certain

identification.

Cuzco: Prov. Quispicanchi, Hacienda Chiraura, Her-

rera 2623 (uc, us). Prov. Paucartambo, near Paucartam-

bo. Plowman & Davis 4915 (F, GH). San Jeronimo, Var-

gas 2320 (MO, uc, us). Puno: Cuyocuyo, Weberbauer 934

(B). Prov. Sandia, cerca a Cuyocuyo, Ferreyra 16614

(GH).

3. Dennstaedtia bipinnata (Cav.) Maxon, Proc.

Biol. Soc. Wash. 61: 39. 1938. Figure 22c.

Dicksonia bipinnata Cav., Descr. pi. 1 74. 1 802. TYPE:

Porto Rico, Ventenat (holotype, MA; isotype, B!,

Herb. mild. 20165-1; photo, GH; frag., us!).

Leaves ca. 1-3 m long. Lamina ovate to lanceo-

late-ovate, 3-4-pinnate-pinnatifid. Axes of the

penultimate segments bordered on each side, on

the adaxial surface, by a pronounced herbaceous

wing perpendicular to the plane of the segment,

the wing on the basiscopic side decurrent onto the

axis of the next order, sterile vein tips on the adax-

ial surface usually slender, ending well away from

the glabrous margin. Pinnules usually slightly pu-

bescent abaxially, the trichomes often nearly con-

fined to the base of the tertiary segments, tri-

chomes wholly clear brown to tan (rarely

subopaque or whitish), rather straight and rigid.

Sori mostly borne in a sinus, mature indusia cy-

lindrical to rarely globular.

In forests, 135-1800 m, San Martin and Loreto

to Cuzco and Madre de Dios.

Southern Florida; Mexico and Central America;

Greater Antilles; Trinidad to Colombia, south to

Peru and Bolivia.

The lamina in Dennstaedtia bipinnata is usually

coriaceous or firmly herbaceous and shining abax-

ially, and the ultimate segments are acutely to sub-

acutely dentate. In D. globulifera the lamina is

usually softly herbaceous and dull abaxially, and

the ultimate segments are entire to usually bluntly

dentate.

San Martin: Monte Campana, Tarapoto, Spruce 4690

(A, K). San Roque, LI. Williams 7/79(F). Loreto: Mouth

of Rio Santiago, Mexia 6127 (BM, F, GH, K, MO, uc, us).

Huanuco: Sinchono, Aguilar 940 (USM). Junin: Satipo,

Aug., 1940, Ridoutt (GH, USM). Chanchamayo valley, C.

Schunke 947 (F, us). Ucayali: Prov. Coronel Portillo,

Bosque Nacional de Iparia, J. Schunke V. 2713 (F, GH,

us). Ayacucho: Near Kimpitirki, Apurimac valley, Killip

& Smith 22875 (F, us), 22957 (BM, us). Cuzco: Prov. La

Convention, Dudley 10239 (GH). Madre de Dios: Prov.

Tambopata, Tambopata Nature Reserve, Barbour 5331

(F, MO). Prov. Manu, Rio Palotoa, Terborgh & Foster

6796 (F).

4. Dennstaedtia globulifera (Poiret) Hieron., Bot.

Jahrb. Syst. 34: 455. 1904. Figure 22d.

Polypodium globuliferum Poiret in Lam., Encycl. 5:

554. 1804. TYPE: Santo Domingo, Plumier,

Traite foug. Amer. /. 30. 1705.

Leaves ca. 1-3 m long. Lamina deltoid, 3-pin-

nate to nearly 4-pinnate. Axes of the penultimate

segments bordered on each side on the adaxial

surface by a pronounced herbaceous wing perpen-

dicular to the plane of the segment, the wing on

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

99

the basiscopic side decurrent onto the axis of the

next order, sterile vein tips on the adaxial surface

usually slender, ending well away from the gla-

brous margin. Pinnules more or less pubescent

abaxially, the trichomes wholly clear brown or tan

(rarely subopaque or whitish), rather straight and

rigid. Sori borne in a sinus or at the apex of a lobe,

mature indusia globular to rarely cylindrical.

In forests, 1000-2800 m, Cajamarca to Cuzco.

Texas; Mexico and Central America; Greater

Antilles; Venezuela and Colombia south to Boliv-

ia, Argentina, and southern Brazil.

Additional characters, useful for the separation

of Dennstaedtia globulifera and D. bipinnata are

discussed under the latter species.

trichomes subappressed, ascending, more or less

curled. Sori borne in a sinus or at the apex of a

lobe, mature indusia purse- to cup-shaped.

In forests and cloud forests, 500-2100 m, Ama-

zonas south to Cuzco and Madre de Dios.

Mexico and Central America; Antilles; Trinidad

to Colombia, south to Bolivia, Paraguay, and

southern Brazil.

Although Dennstaedtia obtusifolia is sometimes

recognized as a distinct species, it is very similar

to D. dissecta and the two are probably variations

of one species. The shape of the mature indusia is

from purse- to cup-shaped, and this variation in-

cludes the usually cup-shaped indusia of D. ob-

tusifolia.

Cajamarca: Prov. Hualgayoc, Soukup 3809 (F, us).

Amazonas: Prov. Bagua, Cordillera Colan, Barbour 3981

(MO). Pasco: Oxapampa, Soukup 2357 (F). Pichis Trail,

Yapas, (as Junin), Killip & Smith 25473 (F, GH, us).

Junin: Above San Ramon, Killip & Smith 24648 (F, us).

Chanchamayo valley, C. Schunke 12 (F, us). Cuzco: Valle

de Occobamba, Biies 879 (us). Prov. Urubamba, Chavez

3424 (GH).

5. Dennstaedtia dissecta (Sw.) Moore, Index fil.

305. 1861. Figure 22a-b.

Polypodium dissectum Sw., Prodr. 134. 1788, not

Forster, 1786.

Dicksonia dissecta Sw., J. Bot. (Schrader) 1800(2):

1802. TYPE: Jamaica, Swartz (holotype, s!; frag.,

us!; photo, us).

Dicksonia obtusifolia Willd., Sp. pi. ed. 4, 5: 483. 1810.

TYPE: Venezuela, (Dist. Federal), Caracas, Bre-

demeyer (holotype, B!, Herb. Willd. 20163; pho-

tos, GH, us; frag., LE!).

Dicksonia erosa Kunze, Linnaea 9: 88. 1834. TYPE:

Peru, (Huanuco), Pampayacu, July, 1829, Poep-

pig (Diar. 1127), (holotype, LZ destroyed; iso-

types?, Poeppig 169, B!, P!, LE!; photo, GH of B).

Dennstaedtia obtusifolia (Willd.) Moore, Index fil. 306.

1861.

Dennstaedtia erosa (Kunze) Moore, Index fil. 306.

1861.

Leaves ca. 2-3 m long. Lamina deltoid to lan-

ceolate, 2-pinnate-pinnatifid to 4-pinnate. Axes of

the penultimate segments without perpendicular

herbaceous wings on the adaxial surface, or with

them but the wing on the basiscopic side not de-

current onto the axis of the next order, sterile vein

tips on the adaxial surface clavate to punctate

(rarely slender), ending well away from the gla-

brous margin. Pinnules glabrate to usually pubes-

cent abaxially, especially on the axes, the brownish

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

2498 (F, GH, MO, uc). San Martin: Near Tarapoto, Spruce

4246 (GH). Prov. Rioja, Pedro Ruiz-Moyobamba, Smith

& Vdsquez 4693 (GH). Huanuco: Puente Durand to Exito,

Mexia 8248 (F, GH, uc, us). Pasco: Pichis Trail, (as Junin),

Killip & Smith 25550 (us). Junin: Near La Merced, Killip

& Smith 23915 (BM, F, us). Ucayali: La Divisoria, (as

Huanuco), Aguilar 851 (GH, USM). Ayacucho: Estrella,

between Huanta and Rio Apurimac, Killip & Smith 22647

(F, GH, us). Cuzco: Prov. La Convention, Cordillera Vil-

cabamba, Dudley 11268 (GH). Madre de Dios: Prov.

Manu, Cerro de Pantiacolla, Foster et al. 10909 (F).

6. Dennstaedtia arborescens (Willd.) Maxon, Proc.

Biol. Soc. Wash. 43: 88. 1930.

Davallia arborescens Willd., Sp. pi. ed. 4, 5: 470. 1810.

TYPE: Santo Domingo, Plumier, Traite foug.

Amer. /. 6, 1705.

Davallia concinna Presl, Reliq. haenk. 1: 66. 1825,

not Schrader, 1818. SYNTYPES: "Chile", 1790,

Haenke (2 sheets, PR; photos, GH, us). The species

is not known from Chile, and Presl was undoubt-

edly correct in later (Epim. hot. reprint 1 02) citing

the Haenke collections from Peru.

Deparia mathewsii Hooker, Sp. fil. 1: 85, t. 30B. 1844.

TYPE: Peru, Mathews 1782 (holotype, K!; frag.,

P!, us!; isotypes, BM!, us!).

Dennstaedtia concinna (Presl) Moore, Index fil. xcvii.

1857.

Dennstaedtia mathewsii (Hooker) C. Chr., Index fil.

218. 1905.

Leaves ca. 1.5-3 m long. Lamina ovate to lan-

ceolate, 2-3-pinnate. Axes of the penultimate seg-

ments without perpendicular herbaceous wings on

the adaxial surface, or with them but the wing on

the basiscopic side not decurrent onto the axis of

the next order, sterile vein tips on the adaxial sur-

face clavate to punctate, ending well away from

the glabrous margin. Pinnules glabrate to usually

100

FIELDIANA: BOTANY

pubescent beneath, the trichomes brownish, su-

bappressed, ascending, more or less curled. Sori

mostly borne at a nearly entire margin, often con-

tiguous, mature indusia purse-shaped to usually

cup-shaped.

In forests, 600-2 1 50 m, Amazonas to Puno.

Mexico and Central America; Greater Antilles;

Venezuela to Colombia, south to Bolivia.

The shape and size of the ultimate segments are

quite variable in this species. Sometimes it has

proliferous buds in the axils of the pinnae, a char-

acter also present in Dennstaedtia dissecta.

Amazonas: Prov. Bagua, Cordillera Colan, Barbour

4138 (F, MO). San Martin: Tarapoto, Spruce 4346 (BM,

GH, us). Prov. Rioja, Pedro Ruiz-Moyobamba, Smith &

Vdsquez 4699 (GH). Huanuco: Huacachi, near Muna,

Macbride 4176 (F, us). Cushi, Macbride 4842 (F, us).

Junin: Above San Ramon, Killip & Smith 24642 (F, GH,

us). Cuzco: Prov. La Convencion, Amaibamba, Vargas

9805 (uc). Prou. La Convencion, Cordillera Vilcabam-

ba, Dudley 10165 (GH). Madre de Dios: Prov. Manu,

Atalaya. Foster & Wachter 7425 (MO). Puno: San Gaban,

Lechler 2157 (B).

7. Dennstaedtia spruce! Moore, Index til. 308.

1861. TYPE: Ecuador, Spruce 5350 (holo-

type, Herb. Hooker, K!; photo, us; isotypes,

GH!, P!; frag., us!; photo, us of c).

Leaves ca. 1-2 m long. Lamina lanceolate, pin-

nate-pinnatifid to 2-pinnate-pinnatifid. Axes of the

penultimate segments without herbaceous wings

on the adaxial surface, sterile vein tips on the adax-

ial surface clavate to punctate, nearly reaching the

persistently pubescent margin. Pinnules and pin-

nae pubescent abaxially, especially on the axes,

the trichomes brownish, curled. Sori borne at the

nearly entire margin, mostly contiguous, mature

indusia purse- to cup-shaped.

Dense forests, 1700-1900 m, Pasco.

Ecuador and Peru.

Dennstaedtia sprucei is evidently a rare species

and adequate material has not been available to

determine variation, if there is any, in the shape

of the lamina. The persistently pubescent margin

with the vein ends extending very nearly to the

margin are distinctive characters of D. sprucei.

This species was originally described and figured

by Hooker (Sp. fil. 1: 74, /. 26 A) under the name

Dicksonia pavonii. That name, however, is tech-

nically a nom. nov. for Davallia arborescens Willd.,

not Dicksonia arborescens L 'Her.

Pasco: Oxapampa, (as Junin), Soukup 1826 (GH, us).

Dos de Mayo, (as Junin) Killip & Smith 25848 (BM, GH,

us). Dept. Unknown: Perou (F no. 809472).

8. Dennstaedtia wercklei (Christ) Tryon, Contr.

Gray Herb. 187: 50. 1960.

Saccoloma wercklei Christ, Bull. Herb. Boissier, II, 4:

1 100. 1904. TYPE: Costa Rica, Werckle320(ho-

lotype, P!; photo in Amer. Fern J. 48: /. 13-14.

1958).

Leaves ca. 0.5-1.5 m long. Lamina lanceolate,

1 -pinnate. Pinnae entire, sometimes subauricu-

late, sterile vein tips on the adaxial surface clavate

to punctate, ending near the glabrous margin,

slightly pubescent beneath, especially on the costa,

the trichomes brownish, subappressed, ascending,

more or less curled. Sori contiguous along the en-

tire margin, mature indusia purse-shaped, contig-

uous, and often joined.

In forests and cloud forests, 1700-2040 m,

Amazonas, San Martin, and Huanuco.

Costa Rica, Colombia, and Peru.

The 1 -pinnate lamina makes this a very dis-

tinctive species. It is rarely collected and has an

unusual, disjunct distribution.

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

2572 (F, MO, uc). San Martin: Prov. Rioja, Pedro Ruiz-

Moyobamba, Smith & Vdsquez 45.93 (GH). Huanuco:

Pozuzo, Bryan 674 (us).

III. Saccoloma

Saccoloma Kaulf., Berlin Jahrb. Pharm. 1820: 51.

TYPE: Saccoloma elegans Kaulf. Figure 23.

Terrestrial. Stem moderately stout, erect to de-

cumbent, bearing scales. Leaves borne in a cluster,

to ca. 2.5 m long. Lamina 1-5-pinnate, glabrous

or nearly so, veins free. Sori marginal or nearly

so, not paraphysate, covered or not by an abaxial

indusium attached at its sides, the leaf-tissue be-

yond the sorus unmodified or slightly modified.

Spores tetrahedral-globose, trilete, the surface with

long, nearly parallel ridges.

Saccoloma is a tropical genus of about 1 0 species,

with three of them in America, and two in Peru.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

101

FIG. 23. Saccoloma inaequale: a, apical portion of lamina; b, portion of pinnule, abaxial side. Saccoloma elegans:

c, silhouette of leaf apex; d, fertile pinna margin, abaxial side. (Adapted from Stolze, Ferns & fern allies of Guatemala,

1981.)

102

FIELDIANA: BOTANY

In the 1 -pinnate Saccoloma elegans the sori are

often confluent at maturity or laterally joined, the

margin may be recurved and somewhat modified,

and the abaxial indusium is sometimes poorly de-

veloped.

Reference

TRYON, R. 1962. The genus Saccoloma Kaulf.,

in Taxonomic fern notes, III. Contr. Gray Herb.,

191: 100-106.

Key to Species of Saccoloma

a. Lamina 2-pinnate or more complex

a. Lamina 1 -pinnate, the pinnae entire

1. S. inaequale

. . 2. S. elegans

1. Saccoloma inaequale (Kunze) Mett., Ann. Sci.

Nat. Bot. IV, 15: 80. 1861. Figure 23a-b.

Davallia inaequalis Kunze, Linnaea 9: 87. 1 834. TYPE:

Peru, (Loreto), Maynas, Yurimaguas, Dec. 1830,

Poeppig Diar. 2113 (holotype, LZ destroyed; is-

otypes, B!, GH!; frag., ex B, us!; photo, GH of B).

Microlepia inaequalis var. nigrescens Mett., Fil. lechl.

1: 22. 1856. TYPE: Peru, St. Gavan, (Rio San

Gaban), Lechler 2293 (holotype, probably LZ de-

stroyed; isotype, B!; photos, GH, us).

Saccoloma inaequale var. caudal a Hieron., Hedwigia

47: 207. 1908. TYPE: Peru, (Loreto), near "Yqui-

tos" (Iquitos) and Cocha Stubel 1 129 (holotype,

B).

Stem quite erect. Leaves ca. 1-1.5 m long, long-

petioled. Lamina 2^4-pinnate, deltoid to long-tri-

angular, the apex gradually reduced, veins more

or less evident beneath. Sori distant, indusium

glabrous.

In woods, dense forests, and cloud forests, 1 00-

1 760 m, Amazonas to Puno.

Tropical America.

This is a rather commonly collected species.

There is considerable variation in the size of the

ultimate segments and the degree of dissection of

the lamina.

Amazonas: Prov. Bagua, above Cascadas de Mayasi,

Wurdack 1968 (GH, us). Prov, Bagua, 12 km E of La

Peca, Barbour 2626 (F, MO, uc). San Martin: Prov. Mar-

iscal Caceres, Dist. Tocache Nuevo, /. Schunke V. 3925

(F, GH, us). Tarapoto, LI. Williams 6007 (F, GH). Loreto:

Prov. Maynas, Dist. Iquitos, McDaniel & Rimachi 17091

(F, GH). Mishana, Rio Nanay, Solomon 3574 (F, MO).

Iquitos, Killip & Smith 30687 (F, us). Huanuco: Tingo

Maria, Tryon & Tryon 5223 (BM, F, GH, u, us, USM),

Ferreyra 10320 (GH, USM). Pasco: Prov. Oxapampa, cer-

ca de Puerto Bermudez, Leon 304 (USM). Puerto Yessup,

(as Junin), Killip & Smith 26368 (F, us). Junin: Near La

Merced, Killip & Smith 23988 (F, GH, us). Above San

Ramon, C. Schunke A232 (GH, us). Ucayali: Prov. Co-

ronel Portillo, Parque Nacional von Humboldt, Puerto

Inca, Croat 51053 (MO, uc). Cerro de Canchyuaya, Rio

Ucayali, (as Loreto), McDaniel 14155 (F, GH). Ayacucho:

Prov. La Mar, Cordillera Central, Dudley 11940 (GH).

Cuzco: Prov. La Convention, Cordillera Vilcabamba,

Dudley 10604 (GH), 70/47 (GH). Prov. Quispicanchi, entre

Machiche y Inambari, Vargas 16464 (GH). Madre de

Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al.

10700 (GH). Prov. Tambopata, Tambopata Nature Re-

serve, Barbour 5169 (F, MO). Puno: Sandia, Weberbauer

1263 (B).

2. Saccoloma elegans Kaulf, Berlin Jahrb. Pharm.

1820: 51. TYPE: None cited, but in Enum.

fil. 224. 1824, Kaulfuss cited Sello, Brazil,

which may be accepted as the type; two sheets

of this collection are at B!; photo, GH. Figure

23c-d.

Stem erect to decumbent with age. Leaves ca.

1-2.5 m long, long-petioled. Lamina 1 -pinnate,

ovate-oblong, apex with a conform terminal pin-

na, pinnae simple, long- to oblong-elliptical, veins

prominent beneath. Sori very numerous along the

margin, mostly contiguous, indusium glabrous.

In woods and dense forests, 100-1700 m, San

Martin to Puno.

Central America; Greater Antilles; tropical South

America.

This is a very distinctive species and not closely

related to any others in the genus.

San Martin: Prov. Lamas, Lamas, Belshaw 3429 (GH,

us). Prov. San Martin, Dist. Tarapoto, Rimachi 5766

(F). Loreto: Iquitos, Killip & Smith 27121 (F, NY, us).

Huanuco: Prov. Pachitea, Bosque Nacional de Iparia, J.

Schunke V. 2856 (F, GH, us). Pasco: San Nicolas, Pichis

Trail, (as Junin), Killip & Smith 26072 (F, GH, NY, us).

Junin: Above San Ramon, Killip & Smith 24574 (GH,

NY, us). La Merced, Soukup 1090 (F). Puno: San Gaban,

Lechler 2416 (B).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

103

FIG. 24. Pteridium aquilinum var. arachnoideum: a, portion of stem with petiole base; b, portion of pinna; c,

ultimate segment, abaxial side, (a from Macbride & Featherstone 1671, F, b, c, from Macbride 4507, F.)

104

FIELDIANA: BOTANY

IV. Pteridium

Pteridium Scop., Fl. carniol. 169. 1760. TYPE:

Pteridium aquilinum (L.) Kuhn (Pteris aquil-

ina L.). Figure 24.

Terrestrial. Stem slender, long-creeping, fre-

quently branched, bearing trichomes. Leaves

widely spaced, to ca. 7 m long, sometimes scan-

dent on other vegetation. Lamina 2-pinnate-pin-

natifid to 4-pinnate, pubescent to rarely glabrous,

sterile margins with an indusium similar to that

of the fertile ones, veins free. Sori marginal, not

paraphysate, covered by a well-modified marginal

indusium, the sporangia borne on a long vascular

commissure connecting the vein ends, an abaxial

indusium present or absent or poorly developed.

Spores tetrahedral-globose, trilete, irregularly

granulate.

Pteridium is a genus of one species and 1 2 geo-

graphic varieties, distributed nearly throughout the

world. There are six of the varieties in America

and two of them in Peru.

A special feature of the genus is the development

of a sterile indusium on the sterile margins of seg-

ments. This is similar to the indusium that covers

the sorus but is usually narrower.

Reference

TRYON, R. M. 1 94 1 . Revision of the genus Pteri-

dium. Rhodora, 43: 1-31, 37-67, and Contr.

Gray Herb., 134: 1-70.

1 . Pteridium aquilinum (L.) Kuhn in Decken, Re-

isen Ost-Afrika 3(3): 11. 1879.

Pteris aquilina L., Sp. pi. 1075. 1753. TYPE: Figure

ofFilixfemina Fuchs, Hist, stirp. 596 (misprinted

as 569. 1542).

The two varieties in Peru may be separated by

the following key. However, some specimens are

rather intermediate between the varieties, while

rarely others depart from the varieties in one char-

acter or another.

Key to Varieties

a. Free lobes absent on the axis between the ultimate or penultimate segments, the base of the ultimate

segments, especially toward the apex of a penultimate segment, definitely decurrent

la. var. caudatum

a. Free, or nearly free, lobes present on the axis between the ultimate or penultimate segments

. Ib. var. arachnoideum

la. Pteridium aquilinum var. caudatum (L.) Sad-

ebeck, Jahrb. Hamb. Wiss. Anst. 14, Beiheft

3: 5. 1897.

Pteris caudata L., Sp. pi. 1075. 1753. TYPE: LINN

1246.15!; photo, A.

Pteridium caudatum (L.) Maxon, Proc. U.S. Natl. Mus.

23: 631. 1901.

In woods and borders of pastures, 1250-2300

m, Cajamarca, Amazonas, and San Martin.

Mexico and Central America; West Indies; Ven-

ezuela, Colombia to Peru, and northern Brazil.

Cajamarca: Prov. San Miguel, Niepos, Llatas 1546

(F). Amazonas: Prov. Bagua, 5 km S of La Peca, Barbour

4311 (F, MO). Prov. Chachapoyas. entre Ingenio y Po-

macocha, Lopez et al. 4313 (GH). San Martin: Prov.

Lamas, Dist. Lamas, Belshaw 3449 (GH, uc, us).

Ib. Pteridium aquilinum var. arachnoideum

(Kaulf.) Brade, Zeitschrift Deut. Ver. Wissen.

Kunst, Sao Paulo 1:56. 1 920. Figure 24.

Pteris arachnoidea Kaulf., Enum. fil. 190. 1824. TYPE:

Brazil, Chamisso (holotype or isotype, LE!; photo,

GH).

Pteridium arachnoideum (Kaulf.) Maxon, J. Wash.

Acad. Sci. 14: 89. 1924.

Open slopes, pastures, thickets, rocky places,

and cleared land, 400-3000 m, Piura to Puno.

Primarily in South America; less common in

Mexico, Central America, and the West Indies.

The species Pteridium aquilinum is a serious

weed in many parts of the world, and in Peru var.

arachnoideum is often an unwelcome pioneer. It

is especially a serious pest in newly cleared land

intended for agricultural crops. The main stem is

TRYON & STOLZE: PTERIDOPHYTA OF PERU. II.

105

usually deep in the soil, and it sends forth branches

that bear the leaves. These also bear latent buds,

and cutting the leaves or burning only has the effect

of increasing the number of leaves.

Piura: Prov. Huancabamba, Huancabamba-Cuellodel

Indio, Lopez et al. 8916 (F). Cajamarca: Prov. San Miguel,

Llapa-Uchuquina, Sagdstegui et al. 8887 (F, HUT, MO).

NW of Hualgayoc, Stork & Morton 10027 (F, uc). Ama-

zonas: Prov. Chachapoyas, alrededores de Chachapoyas,

Lopez et al. 4342 (GH). La Libertad: Prov. Trujillo, Cerro

Campana, Sagdstegui & Cabanillas 8352 (MO). San Mar-

tin: Tarapoto, LI. Williams 5641, 5971 (F). Huanuco:

Between Chinchao and Puente Durand, Coronado 92

(GH, us). 5 km NE of Acomayo, Tryon & Tryon 5224

(BM, F, MO, u, us, USM). Lima: Prov. Chancay, Lachay,

Ferreyra 9767 (GH, USM). Pasco: Prov. Oxapampa, near

confluence of Rio Palcazu and Rio Iscozacin, Smith &

Franzen 1942 (F). Junin: Huacapistana, Ferreyra 307

(GH, USM). Carpapata, Cerrate 2796 (GH, USM). Ayacu-

cho: Estrella, Killip & Smith 23095 (F, NY, us). Cuzco:

Cerro de Cusilluyoc, Pennell 13936 (F, GH, NY, us). Prov.

Urubamba, Machu Picchu, Ferreyra 2705 (GH, USM).

Madre de Dios: Planchon, Gutte & Af tiller 8307 (USM).

Puno: Prov. Sandia, San Juan del Oro, Soukup & Lopez

14699 (GH).

V. Paesia

Paesia St.-Hil., Voy. distr. diam. 1: 381. 1833.

TYPE: Paesia viscosa St.-Hil. = Paesia glan-

dulosa (Sw.) Kuhn. Figure 25.