OF

ILLINOIS LIBRARY

AT URBANA-CHAMPA1GN

BIOLOGY

FIEL

Botany

NEW SERIES, NO. 27

PTERIDOPHYTA OF PERU

Part IV

17. Dryopteridaceae

Rolla M. Tryon

Robert G. Stolze

With the collaboration of:

John T. Mickel

Robbin C. Moran

"Mr 21

February 28, 1991

Publication 1424

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i>. 1978. Flora of Barro Colorado Island. Stanford University Press, Stanford, Calif., 943 pp.

i., J. R. Lloyd, and T. D. Pennington. 1963. A comparison of montane and lowland rain forest in Ecuador.

le forest structure, physiognomy, and floristics. Journal of Ecology, 51: 567-'

Langdon, E. J. M. 1979. Yage among the Siona: Cultural patterns in visions, pp. 63-80. In Browman, D. L.,and R. A.

Schwarz, eds., Spirits, Shamans, and Stars. Mouton Publishers, The Hague, Netherlands.

a, J. 1946. The historic tribes of Ecuador, pp. 785-821. In Steward, J. H., ed., Handbook of South American

Indians. Vol. 2, The Andean. Civilizations. Bulletin 143, Bureau of American Ethnology, Smithsonian

tution, Washington, D.C.

Stolze, R. G. 1981. Ferns and fern allies of Guatemala. Part II. Polypodiaceae. Fieldiana: Botany, n.s., 6: 1-522.

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FIELDIANA

Botany

NEW SERIES, NO. 27

PTERIDOPHYTA OF PERU

Part IV

17. Dryopteridaceae

Rolla M. Tryon

Harvard University Herbaria

22 Divinity Avenue

Cambridge, Massachusetts 02138

and

Department of Biology

University of South Florida

Tampa, Florida 33620-5150

Robert G. Stolze

Associate Curator

Department of Botany

Field Museum of Natural History

Roosevelt Road at Lake Shore Drive

Chicago, Illinois 60605-2496

With the collaboration of:

John T. Mickel, New York Botanical Garden

and

Robbin C. Moran, Missouri Botanical Garden

Accepted August 10, 1990

Published February 28, 1991

Publication 1424

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 1 99 1 Field Museum of Natural History

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ISSN 00 15-0746

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

List of Illustrations

INTRODUCTION

ACKNOWLEDGMENTS . .

17. DRYOPTERIDACEAE

Ctenitis . .

1

1

2

5

Megalastrum 11

Triplophyllum 19

Tectaria 21

Cyclopeltis 29

Rumohra 31

Lastreopsis 31

Dryopteris 35

Cyrtomium 38

Didymochlaena 40

Stigmatopteris 42

Cyclodium 47

Polystichum 49

Olfersia 55

Polybotrya 57

Diplazium 65

Athyrium 88

Hemidictyum 90

Cystopteris 92

Woodsia 94

Oleandra 96

Bolbitis 98

Lomariopsis 105

Lomagramma 109

Elaphoglossum Ill

Peltapteris 167

MAP OF PERU 171

INDEX TO NAMES . .172

1. Ctenitis: C. submarginalis; C. ampla . . 6

2. Megalastrum biseriale 12

3. Triplophyllum funestum var. funest um 20

4. Tectaria incisa var. incisa 22

5. Cyclopeltis semicordata 28

6. Rumohra adiantiformis 30

7. Lastreopsis: L. effusa; L. killipii 32

8. Dryopteris: D. paleacea; D. denticulata 34

9. Cyrtomium dubium 39

10. Didymochlaena truncatula 41

1 1 . Stigmatopteris: S. heterophlebia; S. pel-

lucidopunctata 43

12. Cyclodium: C. trianae; C. meniscioides

var. meniscioides 46

1 3. Polystichum: P. pycnolepis; P. platyphyl-

lum; P. montevidense var. montevi-

dense 50

14. Olfersia cervina 56

15. Polybotrya: P. fractiserialis; P. osmun-

dacea; P. polybotryoides 58

16. Diplazium: D. cristatum; D. roemeri-

anum; D. striatum 66

1 7. Athyrium dombeyi 89

18. Hemidictyum marginatum 91

19. Cystopteris fragilis 93

20. Woodsia montevidensis 95

21. Oleandra: O. articulata; O. lehmannii . . 97

22. Bolbitis: B. serrata; B. serratifolia; B.

lindigii 99

23. Lomariopsis: L. fendleri; L. japurensis;

L. nigropaleata 104

24. Lomagramma guianensis 108

25. Elaphoglossum: E. paleaceum; E. eri-

naceum; E. piloselloides; E. squamipes ..110

26. Peltapteris peltata f. peltata 168

111

PTERIDOPHYTA OF PERU

Part IV

17. Dryopteridaceae

Introduction

This fourth part of the "Pteridophyta of Peru" is

devoted exclusively to the family Dryopterida-

ceae. The general style, typography, form of ci-

tations, and so forth follow the previously pub-

lished parts I and II. These matters are adequately

dealt with in part I (Fieldiana: Botany, n.s. No.

20, 1989), and it is not necessary to repeat them

here.

Stigmatopteris, Olfersia, and Polybotrya have

been contributed by Robbin C. Moran, and Elaph-

oglossum and Peltapteris have been contributed

by John T. Mickel. Other genera have been a joint

effort of Rolla M. Tryon and Robert G. Stolze,

each critically reviewing the treatments prepared

by the other.

The Dryopteridaceae are the largest family of

Pteridophyta in Peru, with 26 genera and about

200 species. Many of the genera are taxonomically

difficult because of considerable variation and the

lack of recent critical work on the Andean ele-

ments that would provide guidance. Among these

may be mentioned Ctenitis, Megalastrum, Tec-

tana, Polystichum, Diplazium, and Elaphoglos-

sum. In these genera there is clearly a maximum

of variation in the Andes from Bolivia north to

Colombia, and also extensive intergradation be-

tween many of the variations of a species. The

taxonomy is often less complicated to the east and

north, especially in Mexico and Central America.

Accordingly, a number of commonly recognized

species are placed as synonyms for Peru or they

are maintained with some doubt. Although the

treatments for Peru are based on all materials from

South America, or from tropical America, there

is need for monographic study of many genera, so

that the species characters are better known and

intergradation and its geographic basis are more

fully understood.

Type collections from Peru are mentioned in

the nomenclature but are not repeated in the spec-

imen citations. They are, however, included in the

Peru range and ecology. The nomenclature of the

genera and species is not intended to be complete.

It includes all names based on Peru material and

other names that are considered useful to mention.

C. V. Morton (Amer. Fern J. 62: 57-64. 1972)

has argued that the Cryptogamae Vasculares Qui-

tenses of Sodiro was published over a period of

years, most of it in the Anales Univ. Quito, 1890-

1895, and a small part in the Anales Univ. Central

Rep. Ecuador, 1897. He proposed that the book

was not published until 1897. This conclusion may

be correct, although Luis Mille, who was certainly

in a position to know, indicated (Nov. Rec. Crypt.

Vase. Ecuador, page 1 and page 5, 1927) that the

book was published in 1893. We accept that date

and consequently only one of the names cited for

the Pteridophyta was published earlier in the An-

ales: Acrostichum setigerum Sodiro, 1890 =

Elaphoglossum setigerum (Sodiro) Diels.

Abbreviations of periodicals generally follow the

system of Botanico-Periodicum-Huntianum

(1 968), while those of books and authors generally

follow the system of Taxonomic Literature (TL-

2, 1976 et seq.) The acronyms for herbaria follow

Index Herbariorum and are also provided below.

Acknowledgments

The authors are extremely grateful to Dr. John

T. Mickel (Elaphoglossum, Peltapteris) and Dr.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

Robbin C. Moran (Olfersia, Polybotrya, Stigma-

topteris) for contributing treatments of genera that

are their particular specialties. Their having lent

their expertise to this Flora has provided our best

understanding of the species of these genera in

Peru.

We would like to extend special thanks to Dr.

Abundio Sagastegui (HUT) and Blanca Leon (USM)

for their invaluable assistance in preparing loans

and arranging for packing and shipment of spec-

imens from those two important Peruvian her-

baria. The original drawings were contributed by

Field Museum scientific illustrators Zorica Dabich

and Clara L. Richardson, and by volunteer illus-

trator Julia A. Liesse. Their careful study, pains-

taking efforts, cooperation, and patience have pro-

duced the splendid drawings which add so much

to the descriptions. We are also grateful to Dr.

David S. Harrington (Polystichurri) and Dr. Alan

R. Smith (Cyclodium) for their critical suggestions

on treatments of these two genera. Thanks are also

extended to reviewers of the manuscript for their

valuable comments.

We are also grateful to the officers of the follow-

ing institutions for granting loans of their material

or allowing us to examine specimens in their her-

baria: Herbarium Amazonense, Universidad Na-

tional de la Amazonia Peruana, Iquitos, Peru

(AMAZ); Herbarium Jutlandicum, Aarhus Univer-

sitet, Denmark (AAU); Botanischer Garten und Bo-

tanisches Museum, Berlin-Dahlem, Berlin (B);

British Museum (Natural History), London (BM);

Herbario Nacional Colombiano, Museo de His-

toria Natural, Bogota, Colombia (COL); Royal Bo-

tanic Garden, Edinburgh (E); Field Museum of

Natural History, Chicago (F); Harvard University

Herbaria, Cambridge, Mass.— most Gray Herbar-

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

Truxillense, Universidad Nacional de Trujillo,

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

England (K); Herbario Nacional de Bolivia, Uni-

versidad Mayor de San Andres, La Paz, Bolivia

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

New York Botanical Garden, New York (NY); Mu-

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

bario del Institute de Ciencias Naturales, Univer-

sidad Central, Quito, Ecuador (Q); Institute de

Ciencias, Pontificia Universidad Catolica del Ec-

uador, Quito, Ecuador (QCA); University of Cali-

fornia, Berkeley (uc); Institute Botanico, Caracas,

Venezuela (VEN); United States National Herbar-

ium, Smithsonian Institution, Washington, D.C.

(us); Herbario San Marcos, Universidad Nacional

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

turhistorisches Museum, Vienna (w).

This project has been supported in part by grant

BSR-85- 16358 from the National Science Foun-

dation, Systematic Biology Program. The work

would not have been possible without this assis-

tance. However, any opinions and conclusions ex-

pressed are those of the authors and do not nec-

essarily reflect the views of the Foundation.

Family 17. DRYOPTERIDACEAE

Dryopteridaceae Herter, Rev. Sudam. Bot. 9: 15.

1949; description from Baker in Mart., Fl.

brasil. 1(2): 458, Polypodiaceae, Tribe Aspi-

deae. TYPE: Dryopteris Adans.

Stem stout to slender, often branched, erect,

rarely arborescent, decumbent to long-creeping or

scandent, usually indurated, bearing scales. Leaves

ca. 1 cm to 3 m long, circinate in vernation, mono-

morphic to dimorphic, entire to usually pinnate,

rarely to flabellate, glabrous, pubescent, or scaly

abaxially. Petiole lacking stipules, not articulate to

the stem. Veins usually free, to anastomosing. Sori

abaxial, round to very elongate, at the tip of a vein,

or along it, indusiate or exindusiate, or the spo-

rangia generally distributed over the surface of the

fertile segments; sometimes paraphysate, usually

not. Sporangia with a 2- or 3-rowed stalk and a

vertical or nearly vertical annulus which is inter-

rupted by the stalk.

The Dryopteridaceae are a family of perhaps

1,000 species and over 50 genera. There are 26

genera in Peru. Is is characterized by monolete

spores and a chromosome number based on n =

40 or 41. The groups of genera are sometimes

considered as subfamilies or families. Tryon and

Tryon (1982) treated the Peruvian genera as in the

following synopsis.

Key to Tribes of Dryopteridaceae in Peru

a. Sporangia in sori, or sporangia acrostichoid and then the fertile and sterile leaves very strongly

dimorphic, the fertile with much reduced green tissue and soon withering after the spores are shed

. b

FIELDIANA: BOTANY

b. Petiole continuous c

c. Three vascular bundles, or more, near the base of the petiole

Tribe Dryopterideae, genera I-XV

c. Two vascular bundles near the base of the petiole .... Tribe Physematieae, genera XVI-XX

b. Petiole jointed Tribe Oleandreae, genus XXI

Sporangia acrostichoid and the fertile and sterile leaves more or less dimorphic, the fertile with green

tissue and more or less persistent Tribe Bolbitideae, genera XXII-XXVI

The key to genera has been simplified, as much segments) may have a different configuration on

as possible, by including in a genus only the char- the adaxial (upper) side from that of the pinna-

acters of species known from Peru or those likely rachis.

to occur there. For example, the species of Wood-

sia with an articulate petiole are only far north of

Peru and so the genus is keyed out as having a Reference

continuous petiole. In addition, genera are some-

times keyed out more than once to avoid long TRYON, R. M., AND A. F. TRYON. 1982. Dryop-

headings. Especially in species with decompound teridaceae, pp. 454-627, in Ferns and allied

leaves, the costa (an axis bearing adnate or joined plants, Springer- Verlag, New York.

Key to Genera of Dryopteridaceae

a. Sporangia borne in well-defined sori, sometimes along the veins b

b. Indusium scalelike and attached at the base and arching over the sorus, or attached around the

base of the receptacle, or elongate and attached at the center or on one side c

c. Indusium scalelike and attached at the base and arching over the sorus, or attached around

the base of the receptacle d

d. Indusium scalelike and attached at the base and arching over the sorus

XIX. Cystopteris

d. Indusium attached around the base of the receptacle XX. Woodsia

c. Indusium elongate, attached at the center or usually along one side e

e. Lamina 2-pinnate, the pinnules dimidiate, 3 or more vascular bundles near the base of the

petiole X. Didymochlaena

e. Lamina entire to 5-pinnate, if 2-pinnate then the pinnules nearly equilateral; 2 vascular

bundles near the base of the petiole f

f. Veins free, except as the tips are connected by a marginal strand, or veins copiously

anastomosing and the lamina entire to pinnatisect, at least beyond the basal pinnae . . . g

g. Sori often on both sides of a vein (especially on the basal veins of a vein-system) and

then each sorus and indusium separate distally; lobes and ultimate segments (when

present) entire, crenulate, or slightly and obtusely dentate; lamina entire to 3-pinnate-

pinnatifid XVI. Diplazium

g. Sori single on a vein, sometimes partly on the other side and then the sorus and

indusium continuous distally; ultimate segments linear or strongly and sharply dentate;

lamina 2-pinnate to 5-pinnate XVII. Athyrium

f. Veins freely anastomosing toward the margin; lamina fully 1 -pinnate

XVIII. Hemidictyum

b. Indusium peltate or with a sinus and nearly orbicular to reniform, or indusium fugacious, nearly

obsolete, or truly absent h

h. Petiole articulate (jointed) or pinnae articulate to the rachis i

i. Pinnae articulate, entire, with a large, basal, basiscopic auricle that overlies the rachis . . .

V. Cyclopeltis

i. Petiole articulate; lamina entire XXI. Oleandra

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 3

h. Petiole continuous (not articulate); pinnae (when present) continuous with the rachis j

j. Adaxial side of the costa raised (axes of lower order, e.g., pinna-rachis or rachis, may be

sulcate adaxially) k

k. Veins anastomosing, or if free then the lamina pinnatifid (2-pinnatifid to 3-pinnatifid)

IV. Tectaria

k. Veins free and the lamina fully pinnate 1

1. Adaxial side of the pinna-rachis raised, or if sulcate then the groove not open to admit

the groove of the rachis m

m. Stem decumbent to erect; leaves caespitose; scales of the costae and rachis ample

to abundant n

n. Trichomes on the adaxial side of the axes reddish brown, with 2—4 cells, less

than 0.4 mm long, catenate and usually with a blunt tip when dry; vein tips

slender; basal basiscopic vein of distal pinnules arising from the costule (or

very close to its juncture with the costa) I. Ctenitis

n. Trichomes on the adaxial side of the axes whitish, usually with more than 4

cells, 0.5 mm or more long, terete or flat and with a pointed tip when dry;

vein tips clavate; basal basiscopic vein of the distal pinnules definitely arising

from the costa II. Megalastrum

m. Stem long-creeping, bearing leaves at intervals; scales of the costae and rachis

few or none III. Triplophyllum

1. Adaxial side of the pinna-rachis sulcate, the groove continuous with that of the rachis

o

o. Adaxial side of the rachis with a central ridge and a glabrous groove on each side

VI. Rumohra

o. Adaxial side of the rachis with lateral ridges on each side of a pubescent groove

VII. Lastreopsis

j. Adaxial side of the costa sulcate or nearly flat (there may be a ridge on each side of the

groove) p

p. Sterile and fertile leaves strongly dimorphic; each meristele of the stem surrounded by

a dark sclerenchymatous sheath; indusium absent XV. Polybotrya

p. Sterile and fertile leaves monomorphic or nearly so; each meristele not enclosed by a

dark sclerenchymatous sheath; indusium often present q

q. Lamina tissue usually pellucid-punctate (the "dots" especially visible with transmitted

light), or if the tissue (rarely) lacking visible "dots" then the lamina 1 -pinnate, the

indusium absent, and the pinna-base better developed on the basiscopic side

XI. Stigmatopteris

q. Lamina tissue uniform, not pellucid-punctate, the lamina entire to 5-pinnate, if

1 -pinnate the indusium present, or if absent then the pinna-base better developed

on the acroscopic side r

r. Veins mostly to wholly free; lamina pinnatifid (2-pinnatifid to 3-pinnatifid) and

indusium present, or veins anastomosing and orbicular indusia often present and

these with a sinus IV. Tectaria

r. Veins mostly or wholly free and the lamina pinnate, or veins anastomosing and

orbicular indusia (if present) peltate s

s. Petiole with 2 vascular bundles near its base XVI. Diplazium

s. Petiole with 3 or more vascular bundles near its base t

t. Lamina 2- to 3-pinnate; ultimate segments sharply dentate, or at least the

apex mucronate, or if the apex only slightly so then the margins strongly

revolute XIII. Polystichum

t. Lamina 1- to 5-pinnate; if 2- to 3-pinnate then the ultimate segments with

nearly flat margins that are smooth to obtusely or subacutely dentate . . . u

u. Veins free or mostly free v

v. Indusium with a sinus, reniform to more or less orbicular, persistent

VIII. Dryopteris

FIELDIANA: BOTANY

v. Indusium peltate, orbicular, or if with a sinus then fugacious

XII. Cyclodium

u. Veins anastomosing w

w. Indusium absent, or if present then peltate, the pinnae coriaceous

with a notably thickened margin IX. Cyrtomium

w. Indusium peltate, pinnae herbaceous, the margins not thickened . .

XII. Cyclodium

a. Sporangia distributed over the surface of the fertile lamina or fertile segments (acrostichoid); fertile

and sterile leaves dimorphic, usually strongly so x

x. Pinnae articulate (jointed) y

y. Veins free; paraphyses lacking XXIII. Lomariopsis

y. Veins anastomosing; usually a few paraphyses scattered among the sporangia

XXIV. Lomagramma

x. Pinnae, when present, continuous with the rachis z

z. Most pinnae or pinna-segments 3 cm long or longer, or the lamina entire and the veins

anastomosing with included free veins aa

aa. Veins free and close, simple or forked only near the costa; the vein tips connected by a

marginal strand; sterile lamina 1 -pinnate, with the terminal segment conform with the

lateral ones (pinnae) XIV. Olfersia

aa. Veins free or anastomosing, a marginal strand absent, or if present then the veins fully

anastomosing; sterile lamina with a pinnatifid apex to rarely with a conform apical segment

bb

bb. Fertile lamina more complex than the sterile, 1 -pinnate-pinnatifid or more complex,

or if 1 -pinnate then with a marginal strand in the sterile segments; veins usually free

XV. Polybotrya

bb. Fertile and sterile lamina of equal complexity, 1 -pinnate or less complex, or if 1 -pinnate-

pinnatifid then so only in the basal portions of the lamina; veins anastomosing ....

XXII. Bolbitis

z. Pinnae or pinna-segments ca. 0.5-1.5 cm long, or the lamina entire and the veins free, or if

anastomosing then without included free veinlets cc

cc. Sterile lamina simple, entire, pinnately veined, or in one rare species deeply pedately lobed

XXV. Elaphoglossum

cc. Sterile lamina flabellate or pinnate, or if entire then flabellately veined

XXVI. Peltapteris

I. Ctenitis

Ctenitis (C. Chr.) C. Chr. in Verd., Man. pterid.

543. 1938. Figure 1.

Dryopteris subgenus Ctenitis C. Chr., Biol. Arbejder

tilegnede Eug. Warming 77. 191 1. TYPE: Dryop-

teris Ctenitis (Link) Kuntze (Aspidium Ctenitis Link)

= Ctenitis distans (Brack.) Ching.

Plants terrestrial or sometimes epipetric. Stem

decumbent to erect, provided with usually many,

fibrous roots, the petiole base with a dense cluster

of clathrate scales, these 6-30 mm long (although

shorter and less abundant in C. nigrovenid). Leaves

monomorphic or essentially so, ca. 0.2-2.5 m long,

caespitose, not articulate to the stem. Lamina

1 -pinnate-pinnatifid to 4-pinnate, reduced to a

pinnatifid apex, provided especially on axes with

scales and "Ctenitis hairs," the latter being reddish

brown, unbranched, articulate trichomes, with 2-

4 cells and 0.1-0.4 mm long. Rachis not sulcate

or, if so, the adaxial groove not open to admit the

groove of the pinna stalk, the adaxial ridges (if

any) not continuous with the ridge of the pinna

axis, scales ample to abundant. Veins free (or anas-

tomosing in a few Old World species), their tips

slender and usually terminating at or near the seg-

ment margin, basal basiscopic vein of distal pin-

nules arising from the costule or very close to its

juncture with the costa. Sori roundish, borne along

the veins or at their tips, receptacle somewhat el-

evated, lacking paraphyses. Indusia reniform, or

nearly circular and attached at the center, with a

narrow sinus, persistent, fugacious, or lacking.

Spores ellipsoidal, monolete, surface saccate or

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

1cm

1mm

FIG. 1 . Ctenitis submarginalis: a. stem and portion of petiole; b, portion of lamina; c, base of pinna, abaxial

side. Ctenitis ampla: d, portion of pinna segment, abaxial side; e, marginal trichome. (a, c from Macbride 4241,

F, b from Penland & Summers 262, Ecuador, F, d, e, from Killip & Smith 24072, F.)

FIELDIANA: BOTANY

echinate, in some species with broad folds that

might appear winglike under the light microscope.

Ctenitis is an essentially pantropical genus of

70-80 species, with about half of these in the Neo-

tropics. Its species grow primarily on the floor of

rain forests, at lower or middle elevations, and are

characterized by large decompound leaves and

small, roundish sori with (when present) circular

to reniform indusia. The genus has been confused

with the closely related Lastreopsis, but it can be

readily separated by the configuration of the ad-

axial axes. In Lastreopsis ridges are continuous

with the ridges on the axes of the next order above

or below; but in Ctenitis these ridges are lacking

or, when present, not continuous onto adjacent

axes. Christensen's revisions (1913,1 920) provid-

ed great insight into the characters and relation-

ships of the genus, but much more work is still

required to produce a satisfactory analysis of the

many American species.

Recently a number of species were removed from

Ctenitis and placed in the genera Triplophyllum

and Megalastrum. The evidence supporting this

separation is even stronger than that which

prompted the separation of Lastreopsis. For com-

parison of these features, see discussions under

treatments of Triplophyllum and Megalastrum.

References

CHRISTENSEN, C. 1913. A monograph of the ge-

nus Dryopteris, part 1. The tropical American

pinnatifid-bipinnatifid species. Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 7,

10: 55-282.

CHRISTENSEN, C. 1920. A monograph of the ge-

nus Dryopteris, part 2. The tropical American

bipinnate-decompound species. Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8,

6: 3-132.

Key to Species of Ctenitis

a. Lamina 1 -pinnate-pinnatifid, at least above the basal pinnae b

b. Ultimate segments (at least of proximal pinnae) crenate or crenate-serrate, lacking marginal tri-

chomes; veins of crenate segments often 1 -forked 4. C. nigrovenia

b. Ultimate segments entire, with minute trichomes scattered to abundant on their margins (at least

in segment sinuses); veins unbranched c

c. Indusia pubescent, subpersistent; tissue between veins minutely pubescent abaxially and/or

adaxially; basal veins terminating at segment margin, well above the sinus

3. C. microchlaena

c. Indusia lacking, or minute and fugacious; tissue between veins glabrous (glandular in C. re-

fulgens); basal veins terminating short of segment margin or reaching it at or near the sinus

d

d. Pinnae incised %-y2(-2/3) to the costa, with minute yellow or reddish glands scattered on

(and often between) the veins abaxially; rachis and costa scales entire, not or obscurely

clathrate, often with a sub-bullate base 1. C. refulgens

d. Pinnae incised % or more to the costa, eglandular; rachis and costa scales remotely dentic-

ulate, conspicuously clathrate, more or less flat throughout 2. C. submarginalis

a. Lamina 2-pinnate-pinnatifid to 4-pinnate e

e. Indusia conspicuous and subpersistent; veins and costules eglandular or with conspicuous reddish

glands 5. C. ampla

e. Indusia lacking, or minute and fugacious; veins and (often) costules with minute yellow glands

. . 6. C. sloanei

Ctenitis refulgens (Mett.) (credited in error to

C. Chr. by) Vareschi, Flora Venezuela 1 : 404.

1969.

Phegopteris refulgens Mett., Ann. Sci. Nat. Hot., ser.

5, 2: 240. 1864. LECTOTYPE (designated by C.

Chr., p. 89. 1913): Guyana (as British Guiana),

Schomburgk 1183 (B!; frag., B!). Mettenius cited

four collections, two were cited as the type by

Christensen, one of them, Schomburgk 1183, was

also cited by Mettenius.

Dryopteris refulgens (Mett.) C. Chr., Index fil. 288.

1905.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

Dryopteris refulgens var. peruviana C. Chr., Kongel.

Danske Vidensk. Selsk. Skr. Naturvidensk. Afd.

Ser. 7, 10: 90. 1913. SYNTYPES: Peru, Monte

Campana prope Tarapoto (San Martin), Spruce

4657 (B, L,); Peru, In Monte Guayrapurima, prope

Tarapoto (San Martin), Spruce 4712 (w); Peru,

prope Tabalosas inter urbem Moyobamba et flu-

vium Rio Huallaga, Stiibel 1097 (B). ISOSYN-

TYPES: Spruce 4657 (BM!, GH!, P!); Spruce 4712

(MO!, us!).

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with orange to reddish brown,

linear scales, these obscurely clathrate, about 1 cm

long, with margins entire. Leaves caespitose, to 1

m long and 30 cm broad, the axes adaxially and

the segment margins amply provided with "Cten-

itis hairs." Lamina 1 -pinnate-pinnatifid, the tissue

between the veins glabrous, or usually with scat-

tered, yellow to reddish glands, the rachis and cos-

tae amply provided abaxially with orange to red-

dish brown, filiform, entire scales, these not or

obscurely clathrate, and usually with a sub-bullate

base. Pinnae incised %-!4(-2/3) to the costa, the

segment entire, or sometimes serrulate at the ob-

tuse or truncate apex. Veins simple, 5-8(-10) pairs

per segment, typically reaching the segment mar-

gin, but 1-4 basal ones terminating at or near the

segment sinus (rarely merging to form a costal

areole). Sori mostly inframedial between costule

and margin. Indusia lacking.

In and at edges of wet forests, 135-900 m, San

Martin, Loreto, Huanuco.

Southern Mexico (Chiapas); French Guiana to

Colombia; Peru; Brazil.

Christensen distinguished his var. peruviana

from C. refulgens on the basis of several quanti-

tative characters: pinnae less deeply incised, fewer

veins running to the sinus and basal ones some-

times united. He also said that the costa scales

were fewer, small, and ovate. However, there are

no such scales on the syntypes examined, and none

of the other variable characters are correlated.

Costa and rachis scales are described above as

usually having a sub-bullate base. Although some

of the scales are flat throughout as in the closely

related C. submarginalis, the majority have their

basal portion incurved so that the edges meet; hence

the scale is essentially filiform with a narrowly

cylindrical base, yet not with the expanded, in-

flated appearance which is commonly termed bul-

late.

San Martin: Prov. Lamas, Alonso de Alvarado, J.

Schunke V. 5941 (F). Loreto: Puerto Arturo, Rio Hua-

llaga below Yurimaguas, Killip & Smith 27821 (us).

Huanuco: Prov. Pachitea, Dist. Puerto Inca, D. Smith

1299 (GH, MO).

2. Ctenitis submarginalis (Langsd. & Fisch.) Ching,

Sunyatsenia 5: 250. 1940. Figure la-c.

Polypodium submarginale Langsd. & Fisch., Icon. fil.

12, t. 13. 1810. TYPE: Brazil, "insula Cathari-

nae," Langsdorff(ho\otype, LE).

Nephrodium tarapotense Hooker, Sp. fil. 4: 107. 1862.

TYPE: Peru, "in Monte Campana prope Taro-

poto" (San Martin), Spruce "conf. 4016" (holo-

type, K!). Type not to be confused with Spruce

4016 (K, P) which is C. microchlaena.

Nephrodium lagerheimii Sodiro, Crypt, vase. Quit. 252.

1893. TYPE: Ecuador, "Las pendientes occiden-

tales del Tungurahua," Lagerheim (holotype, not

located; isotypes, c, s!).

Dryopteris submarginalis (Langsd. & Fisch.) C. Chr.,

Index fil. 296. 1905.

Dryopteris submarginalis var. tarapotensis (Hooker)

C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Na-

turvidensk. Afd., ser. 7, 10: 98. 1913.

Dryopteris submarginalis var. lagerheimii (Sodiro) C.

Chr., Kongel. Danske Vidensk. Selsk. Skr. Na-

turvidensk. Afd., ser. 7, 10: 98. 1913.

Plants terrestrial. Stem erect to decumbent, this

and the petiole base densely clothed with orange

or reddish brown, clathrate, linear to lanceolate

scales, these 1-2 cm long, with margins entire or

sparsely denticulate. Leaves caespitose, to 1 m long

and 30 cm broad, the axes adaxially and the seg-

ment margins abundantly provided with "Ctenitis

hairs." Lamina 1 -pinnate-pinnatisect, the tissue

between the veins glabrous, the rachis and costae

amply provided abaxially with orange or brown,

clathrate, flat, linear to filiform, remotely dentic-

ulate scales. Pinnae (at least proximal ones) deeply

incised to within 1-2 mm of the costa, the seg-

ments entire, or serrulate at the obtuse to subacute

apex. Veins simple, 8-15 pairs per segment, typ-

ically reaching the segment margin, basal ones ter-

minating at or near the segment sinus (or often

ending short of the sinus). Sori inframedial to su-

pramedial between costule and margin. Indusia

lacking, or very rarely minute and fugacious.

In deep, wet forests, on rocky slopes, 1 200-1 500

m, Amazonas, San Martin, and Huanuco.

United States (Florida); Hispaniola; southern

Mexico to Panama; Venezuela and Colombia south

to Argentina and Uruguay.

This is a variable species that was separated by

Christensen (1913) into six infraspecific taxa. He

based distinctions on shape of segments and num-

ber of their veins, indusia present or lacking, po-

FIELDIANA: BOTANY

sition of son, and the tissue between veins gla-

brous or glandular-pubescent. These characters

appear to be too variable and uncorrelated for the

recognition of taxa. Soral position, for example,

varies considerably, even on the same lamina, from

inframedial to supramedial. Often son form an

elongated "V", being remote from the costule near

the segment base but crowding it toward the apex.

Specimens in southern South America tend to

have longer, narrower segments, with acute tips

and more veins; elsewhere segments are relatively

broad and obtuse, yet these tendencies are too

inconsistent to merit even infraspecific status.

Amazonas: Prov. Bongara, Dist. Sipabamba, along

Quebrada Fortuna, Young & Eisenberg 322 (MO, uc).

Huanuco: Rio Huallaga Canon, below Rio Santo Do-

mingo, Macbride 4241 (F, us). "[I]n Panatahuac Pro-

vincia," Ruiz 68 (B).

3. Ctenitis microchlaena (Fee) Stolze, comb. nov.

Aspidium microchlaena Fee, Mem. foug. 8: 102. 1857.

TYPE: Mexico, Orizaba, Schaffner 459 (holo-

type, presumably P, but not found; isotype, K!;

photos, F & GH of K).

Dryopteris microchlaena (Fee) C. Chr., Index fil. 278.

1905.

Aspidium karstenii A. Braun, Ind. sem. hort. Berol.

app. 3. 1867 (also published in Ann. Sci. Nat.

Bot. [5] 9: 379. 1868). TYPE: Cultivated Hort.

Berol. from spores of plant collected in Venezue-

la, Karsten (B; frag., BM!; isotype, K!; photos, BM

of B, F & GH of K).

Dryopteris karstenii (A. Braun) C. Chr., Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 7,

10: 98. 1913.

Southern Mexico; Costa Rica; Colombia; Ven-

zuela; Peru.

Alan Smith (Pteridophytes of Venezuela, an an-

notated list. 1985, ined.) placed Dryopteris kar-

stenii as a synonym of C. microchlaena of Costa

Rica and Colombia. There are minor characters

that differ, in the original descriptions of both spe-

cies, such as glabrous, caducous indusia, and gla-

brous laminar tissue in the latter species, vs. per-

sistent, ciliate indusia and puberulent tissue in D.

karstenii. However, an examination of type and

other material of each indicates that Smith was

correct. There is variation throughout the range

in relative size and pubescence of indusia, and

relative abundance of laminar pubescence, but

none of this correlates with any other diagnostic

feature.

Amazonas: Prov. Bagua, along Rio Utcubamba, on

Cerro Tapir, Hutchison 1482 (uc, us). San Martin: Prov.

Lamas, Dist. Lamas, near Rio Chupisena, Belshaw 3503

(F, GH, MO, uc, us). Monte Guayrapurima, near Tara-

poto, Spruce 4016 (K, P); this is a much different plant

than Spruce "conf. 4016" (K), which is the type ofNeph-

rodium tarapotense (= C. sub marginal is).

4. Ctenitis nigrovenia (Christ) Copel., Gen. fil. 1 24.

1947.

Nephrodium nigrovenium Christ, Bot. Gaz. (Craw-

fordsville) 20: 545. 1895. TYPE: Honduras, San-

ta Barbara, San Pedro Sula, Thieme (holotype, P?;

isotype, us!).

Dryopteris nigrovenia (Christ) C. Chr., Index fil. 279.

1905.

Plants terrestrial. Stem erect, this and petiole

base densely clothed with orange, clathrate, linear

to lanceolate scales, these to 1 cm long, with mar-

gins entire or sparsely and minutely denticulate.

Leaves caespitose, to 90 cm long and 25 cm broad,

the margins and the axes on both sides abundantly

provided with "Ctenitis hairs." Lamina 1-pinnate-

pinnatisect, minutely glandular-pubescent on one

or both of the sides, the rachis and costae provided

with filiform to lanceolate scales like those of the

stem. Pinnae deeply incised to within 1-2 mm of

the costa. Veins simple, rarely 1 -forked, basal ones

reaching the margins well beyond the segment si-

nus. Sori medial to supramedial between costule

and margin. Indusia orange to reddish brown, more

or less pubescent, commonly persistent.

On floor of wet, dark forests, ca. 1400 m, Ama-

zonas and San Martin.

Plants terrestrial. Stem decumbent to erect, this

and the petiole abundantly provided with dark

brown, clathrate, lanceolate scales, these to 4-7

mm long, subentire. Leaves caespitose, to 70 cm

long and 20 cm broad, the margins glabrous, the

axes adaxially provided with ''''Ctenitis hairs."

Lamina 1 -pinnate-pinnatisect, abaxially provided

(usually sparsely) with minute yellow or orange

glands, the rachis and costae amply provided with

reddish brown or blackish, linear to filiform scales,

these 1-3 mm long with an expanded or sub-bul-

late base. Pinnae incised near or to the costa, ul-

timate segments entire or proximal ones crenate-

serrate. Veins simple, or those of crenate segments

usually 1 -forked, basal ones reaching the margin

well beyond the segment sinus. Sori medial to in-

framedial between costule and margin. Indusia

minute, reddish brown, glabrous, and (in Peru)

subpersistent.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

Thus far represented in Peru by one specimen:

on shady river bank, 890 m, San Martin.

Southern Mexico to Panama; Trinidad; Vene-

zuela; Colombia to Peru.

Although C. nigrovenia grows from southern

Mexico to Peru, nowhere has it been collected in

abundance. Distinctive features are the sparsely

glandular abaxial surface and the lack of marginal

trichomes. The great majority of species in the

genus are amply provided with " Ctenitis hairs"

on segment margins, especially at the sinus. The

small indusium is usually persistent in South

American representatives of this species, but is

often fugacious in those from Central America.

San Martin: Near Moyobamba, banks of Mayo River,

Woytkowski 35268 (uc).

5. Ctenitis ampla (Willd.) Ching, Sunyatsenia

5: 240. 1940. Figure Id.

Polypodium amplum Willd., Sp. pi. ed. 4, 5: 207. 1810.

TYPE: "America meridionale prope Caripe"

(Venezuela), Humboldt (holotype, B, Herb. Willd.

19722; photos, F, GH), not Dryopteris ampla of

authors (= C. sloanei (Sprengel) Morton).

Aspidium catocarpum Kunze, Linnaea 9: 95. 1834.

TYPE: Peru, between Cassapi and Pampayacu

(Huanuco), Poeppig, coll. July 1829 (holotype, LZ,

destroyed).

Aspidium nemophilum Kunze, Linnaea 9: 95. 1834.

TYPE: Peru (Huanuco), Pampayacu, Poeppig, coll.

July, 1 829 (holotype, LZ, destroyed).

Dryopteris catocarpa (Kunze) Kuntze, Revis. gen. pi.

2: 812. 1891.

Dryopteris nemophila (Kunze) C. Chr., Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8.

6: 57. 1920.

Ctenitis nemophila (Kunze) Ching, Sunyatsenia 5: 250.

1940.

Ctenitis catocarpa (Kunze) Morton, Fieldiana, Bot.

28: 12. 1951.

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with orange to reddish brown,

linear to filiform scales, these subclathrate, 1.5-

2.5 cm long, with margins entire. Leaves caespi-

tose, to 2.5 m long and 80 cm broad, the axes

adaxially and the margins amply provided with

^Ctenitis hairs," and abaxially with orange to red-

dish brown, lanceolate, clathrate scales. Lamina

deltoid, 2-pinnate-pinnatifid to (at base) nearly

4-pinnate, veins and costules abaxially provided

with reddish, cylindric to elliptic glands, or eglan-

dular. Veins simple to 1 -forked in the ultimate

segments. Indusia large or small, but conspicuous

and persistent, often with elliptic, reddish glands,

or short-pubescent or glandular-pubescent.

In dry, open woods or montane rainforests, on

slopes, or in ravines, 380-2100 m, Amazonas to

Junin and Ucayali.

Venezuela; Colombia to Argentina.

This is part of a species complex that has long

been confused; for detailed discussion, see C.

sloanei below. It is possible that C. ampla, as treat-

ed here, contains two separate entities. A number

of specimens in Peru, and one seen from Bolivia,

have large, reddish brown, quite persistent indu-

sia, these lacking marginal trichomes, but with

several reddish brown, elliptic glands scattered on

the surface. These glands are also conspicuous along

the veins and costules abaxially. Typical C. ampla,

occurring throughout the range, including Peru,

has smaller indusia which are usually lighter in

color, with short-pubescent or glandular-pubes-

cent indusia. However, elliptic, reddish brown

glands are lacking on the segments and indusia.

Since the taxonomy of the entire C. sloanei/ ampla

complex is in need of revision, it seems unwise to

create another species or variety at this time.

Amazonas: Prov. Chachapoyas, Ingenio-Pomacocha,

Lopez et al. 431 1 (GH, MO, USM). San Martin: Tarapoto,

Spruce 3942 (K, us). Huanuco: Muna, Bryan 530 (F, GH,

us). Pasco: Prov. Oxapampa, Canyon de Huancabamba,

Leon 666 (F, USM). Junin: La Merced, Killip & Smith

24072 (F, GH, us). Ucayali (as Loreto): Prov. Coronel

Portillo, Boqueron, Ferreyra 16057, in part (USM).

6. Ctenitis sloanei (Sprengel) Morton, Amer. Fern

J. 59: 66. 1969.

Polypodium sloanei Sprengel, Syst. veg. ed. 16, 4: 59.

1827. TYPE: "In sylvis densioribus humidis Cu-

bae ad Cahoba," Poeppig (holotype, LZ, de-

stroyed; frag., BM!; isotypes, B!, HBG, L, P!; frag.,

us!; photo, us of HBG).

Ctenitis ampla of authors, not Polypodium amplum

Willd. 1810, = C. ampla (Willd.) Ching.

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with orange to (en masse)

reddish brown, linear to filiform scales, these sub-

clathrate, 1.5-3 cm long, with entire margins.

Leaves caespitose, to 1.5m long and 50 cm broad,

the axes adaxially and the margins amply provided

with "Ctenitis hairs," and abaxially with orange

to reddish brown, lanceolate, clathrate scales.

Lamina deltoid, 2-pinnate-pinnatifid to (at base)

nearly 4-pinnate, provided abaxially on veins, and

usually costules, with minute (to 0.5 mm) yellow-

ish, cylindrical glands. Veins simple to 1 -forked

in the ultimate segments. Indusia lacking, or oc-

casionally minute and fugacious, short-pubescent.

10

FIELDIANA: BOTANY

In lowland forests, 100-400 m, Cuzco and Ma-

dre de Dios.

United States (Florida); West Indies; southern

Mexico (Chiapas); Nicaragua to Panama; Vene-

zuela; Colombia; Ecuador; Peru.

This is one of a variable complex of neotropical

species whose relationships Christensen (1920) and

Morton (Amer. Fern J. 59: 66. 1969) tried to clar-

ify. Despite their efforts a good revision is still

needed. Only a comparison of types against mass

collections will suffice to delineate species and as-

certain the correct application of names. There are

few good characters with which to separate C. am-

pla, C. sloanei, C. catocarpa, and C. nemophila,

and it is not surprising that Christensen was un-

certain whether all were conspecific, or were close-

ly related taxa with few intermediates. Separation

of C. ampla and C. sloanei has been attempted on

a suite of characters: color and shape of scales,

presence or lack of glands and indusia, shape of

apex and stalk length of pinnules. Examination of

specimens from many areas of the Neotropics in-

dicates that perhaps only the indusia and laminar

glands are reliable and relatively constant, but even

these are at times questionable characters.

Throughout the species complex indusia may be

large and persistent to small and fugacious or lack-

ing, and even persistent indusia can be folded and

inconspicuous within a large sorus of fully ex-

panded sporangia.

Large, red glands on the veins can easily be seen

under low magnification in many specimens of C.

ampla, but the smaller yellowish ones blend with

the tissue in C. sloanei and frequently can be lo-

cated only with diligence and higher (30 x ) mag-

nification. As treated here, C. sloanei is wide-

spread in the Neotropics, yet it is rare in Peru.

Ctenitis ampla is rather common in Peru, but not

especially abundant elsewhere. Ctenitis sloanei

tends to prefer lower elevations throughout its range

(less than 500 m in the two Peruvian collections),

whereas C. ampla is mostly, although not always,

found above 800 m.

Cuzco: Prov. La Convention, Rio Apurimac below

San Martin, Davis et al. 1332 (F). Madre de Dios: Prov.

Manu, Parque National del Manii, Foster 9802 (F, MO).

II. Megalastrum

Megalastrum Holttum, Gard. Bull. Straits Set-

tlem. 39: 161. 1986. TYPE: Megalastrum vil-

losum (L.) Holttum, (Polypodium villosum L.).

Figure 2.

Plants terrestrial. Stem decumbent to erect, oc-

casionally subarborescent to 2 m tall, provided

with many fibrous roots, the petiole usually with

a dense cluster of clathrate scales 6-30 mm long.

Leaves monomorphic, ca. 0.5-3.0 m long, caes-

pitose, not articulate to the stem. Lamina 1-pin-

nate-pinnatifid to 4-pinnate-pinnatifid, reduced to

a pinnatifid apex, often provided on the margins

and/or abaxial surface with "Ctenitis hairs," the

latter being reddish brown, articulate trichomes

with 2-4 cells and 0.1-0.4 mm long, but the ad-

axial axes (except in one Brazilian species) bearing

trichomes with usually more than 4 cells and typ-

ically over 0.5 mm long, these drying terete or flat,

and with pointed tips. Rachis not sulcate or, if so,

the adaxial groove not open to admit the groove

of the pinna stalk, the adaxial ridges (if any) not

continuous with the ridge of the pinna axis, scales

ample to abundant. Veins free, their tips conspic-

uously clavate, ending short of the segment mar-

gin, basal basiscopic vein of distal pinnules arising

from the costa. Sori roundish, borne along the

veins, receptacle somewhat elevated, lacking pa-

raphyses. Indusia reniform, or nearly circular and

attached at the narrow sinus, or lacking. Spores

monolete, roundish to ellipsoidal, echinate or cris-

tate.

Megalastrum is a tropical genus of 45-50 spe-

cies, most of these in America. It was separated

from Ctenitis by Holttum (1986), a decision sub-

stantiated by Smith and Moran (1987) in a paper

that added new combinations for 39 species. The

different trichomes on the adaxial axes is the most

effective character for distinguishing the two gen-

era (see key to genera of Dryopteridaceae), but

there are a number of other corroborating features.

In Megalastrum, the clavate vein tips terminate

obviously short of the segment margin, whereas

in Ctenitis the tips are not or scarcely enlarged and

terminate at or very near the margin. In Ctenitis

the sporangia stalks, and sometimes indusia and

veins, bear unicellular, glandular trichomes, but

these are lacking in Megalastrum.

Although the two genera now may be effectively

separated as outlined above, many problems still

exist in delineating their components. Both con-

tain variable species complexes in which taxa are

distinguished merely by length and abundance of

indument. Detailed examination of specimens

throughout the Neotropics will very likely prove

that a number of taxa currently recognized as spe-

cies are merely varieties or forms of others.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

11

FIG. 2. Megalastrum biseriale: a, habit; b, ultimate segment, adaxial side; c, ultimate segment, abaxial side, (a,

c from J. Schunke A227, us; b from Soukup 1032, F.)

12

FIELDIANA: BOTANY

References

CHRISTENSEN, C. 1913. A monograph of the ge-

nus Dryopteris, part 1. The tropical American

pinnatifid-bipinnatifid species. Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 7,

10: 55-282.

CHRISTENSEN, C. 1920. A monograph of the ge-

nus Dryopteris, part 2. The tropical American

bipinnate-decompound species. Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8,

6: 3-132.

HOLTTUM, R. E. 1 986. Studies in the fern-genera

allied to Tectaria Cav. 6. A conspectus of genera

in the Old World regarded as related to Tectaria.

Gard. Bull. Straits Settlem., 39: 153-167.

SMITH, A. R., AND R. C. MORAN. 1987. New

combinations in Megalastrum (Dryopterida-

ceae). Amer. Fern J., 77: 124-130.

Key to Species of Megalastrum

a. Lamina 1 -pinnate-pinnatifid to (proximally) 2-pinnate; pinnules entire to crenate, or proximal ones

shallowly lobed b

b. Trichomes on the abaxial side of axes and veins 1-2 mm long, moderate to abundant; trichomes

on segment margins to 1 mm long; stem scales often with narrow blackish margins

6. M. hirsutosetosum

b. Trichomes on the abaxial side of axes sparse, 0.1-0.8 mm long, or lacking, or if abundant and to

1 mm long, then mixed with many other trichomes ca. 0.2 mm long; trichomes on segment

margins less than 0.3 mm long; stem scale margins not blackish c

c. Lamina chartaceous; most veins obscure; tissue on abaxial side commonly with minute (0. 1

mm), cylindrical glands d

d. Sori mostly inframedial; rachis scales filiform, to 6 mm long; trichomes lacking or sparse

on the axes abaxially 1 . M. honestum

d. Sori mostly supramedial; rachis scales lanceolate, 2-3 mm long; trichomes abundant on the

axes abaxially 2. M. yungense

c. Lamina firm-herbaceous; most veins evident; tissue eglandular e

e. Rachis scales scattered, 1-1.5 mm long 4. M. platylobum

e. Rachis scales moderate to abundant, 2-4 mm long f

f. Axes abaxially densely covered with trichomes to 0.2 mm long; abaxial surface of lamina

abundantly provided with minute trichomes 3. M. microsorum

f. Axes abaxially glabrous, or with scattered trichomes to 0.6 mm long; abaxial surface

glabrous 5. M. biseriale

a. Lamina 2-pinnate-pinnatisect to 4-pinnate-pinnatifid; pinnules regularly pinnatisect to 1 -pinnate or

more complex g

g. Indusia large, persistent 10. M. andicola

g. Indusia lacking h

h. Lamina short-pubescent, the trichomes (at least abaxially) 0.1-0.5 mm long i

i. Costules, costae, and rachis abaxially glabrous, or the trichomes scattered and varying in

length from 0.3 to 0.6 mm long 7. M. subincisum

i. Costules, costae, and usually the rachis, densely and regularly puberulent abaxially, the

trichomes 0.05-0.2 mm long 8. M. vastum

h. Lamina abaxially long-pilose, especially on costae and veins, the trichomes moderate to abun-

dant, mostly 0.7-1.5 mm long 9. M. pulverulentum

1 . Megalastrum honestum (Kunze) Smith & Mor-

an, Amer. Fern J. 77: 128. 1987.

Polypodium honestum Kunze, Linnaea 9: 49. 1834.

TYPE: Peru (Huanuco), Pampayaco (Pampaya-

cu), Poeppig 22 (holotype, LZ, destroyed).

Polypodium fibrillosum Baker, Syn. fil. 307. 1867.

TYPE: Peru (San Martin), Tarapoto, Spruce 47 42

(holotype, K!; isotypes, BM!, BR, GH!; photo, us of

BR).

Dryopteris fibrillosa (Baker) C. Chr., Index fil. 264.

1905.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

13

Dryopteris honesta (Kunze) C. Chr., Index fil. 271.

1905.

Ctenitis honesta (Kunze) R. & A. Tryon, Rhodora 84:

127. 1982.

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with orange, linear scales,

these obscurely clathrate, to 1.5 cm long, the mar-

gins entire or rarely with a few, minute, scattered

setae. Leaves to 1.2 m long and 40 cm broad, the

axes on the abaxial side sparsely provided with

trichomes to 0.3 mm long or, more commonly,

glabrous. Lamina chartaceous, the tissue and veins

abaxially often with minute, scattered, yellow to

reddish cylindrical glands (ca. 0.1 mm), 1-pinnate-

pinnatisect, to 2-pinnate near the base, the rachis

and costae abaxially provided with abundant, red-

dish brown, filiform, setulose scales, these 2-6 mm

long (to 4 mm on costae). Pinnae incised nearly

or quite to the costae, the ultimate segments entire

to crenate, with scattered marginal trichomes 0. 1-

0.2 mm long. Veins obscure. Sori mostly infra-

medial. Indusia lacking.

In forests, 400-1700 m, San Martin, Huanuco,

Junin, Madre de Dios.

Peru; Bolivia.

In this and Megalastrum yungense the tissue is

so dense that most veins can usually be seen only

by transmitted light. The lamina of other species

in the complex is much thinner and veins are

mostly quite evident. Axes on the abaxial side

typically lack trichomes, although filiform scales

are quite abundant. On only one of all Peruvian

collections examined (Ucayali, cited below) was a

sparse covering of minute trichomes detected.

Huanuco: Pampayacu, Kanehira 179 (us). La Divi-

soria, 25 km NE of Tingo Maria, Moran 3707 (USM).

Junin: Schunke Hacienda above San Ramon, Killip &

Smith 24659 (us). Chanchamayo Valley, C. Schunke 957

(F). Ucayali: Fundo Chela, Sinchono (as Loreto or Hua-

nuco), Aguilar, Aug. 3, 1948 (GH, USM). Madre de Dios:

Prov. Manu, Rio Sotileja, Parque Nacional Manu, Foster

etal. 11598(F).

2. Megalastrum yungense (Christ & Rosenst.)

Smith & Moran, Amer. Fern J. 77: 1 29. 1987.

Dryopteris yungensis Christ & Rosenst., Repert. Spec.

Nov. Regni Veg. 5: 234. 1908. TYPE: Bolivia,

Sirapuya near Yanacachi, 2000 m, Buchtien 493

(holotype, P!; isotype, us).

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with brown (sometimes mot-

tled with black), linear scales, these scarcely or

obscurely clathrate, ca. 6 mm long, the margins

setulose. Leaves to 80 cm long and 25 cm broad,

the axes on the abaxial side amply provided with

trichomes 0.2-0.6 mm long. Lamina chartaceous,

the tissue and veins abaxially often with minute,

scattered, cylindrical glands (ca. 0.1 mm), 1-pin-

nate-pinnatisect, to 2-pinnate near the base, the

rachis and costae abaxially amply provided with

reddish brown, setulose, lanceolate scales, these

2-3 mm long (less on costae). Pinnae incised near-

ly or quite to the costae, the ultimate segments

entire, obtuse, with scattered marginal trichomes

0.1 mm long. Veins obscure. Sori mostly supra-

medial. Indusia lacking.

Thus far known in Peru from one collection, in

high montane forest, 1 800 m, San Martin.

Peru; Bolivia.

Rachis scales in this species complex are typi-

cally filiform, many times longer than broad; but

those of M. yungense are lanceolate to narrow-

deltoid. Scales of the stem and petiole base are

also distinctive, at least in the specimen from Peru:

essentially dark brown, but with scattered black

cells or groups of cells. Because the type consists

only of a leaf broken off above the stem, it is not

possible to ascertain if this color pattern is con-

sistent.

San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba

Rd., km 390, D. Smith 4424 (uc).

3. Megalastrum microsorum (Kuntze) Stolze,

comb. nov.

NephrodiummicrosorumHoo]/ieT,Sp.fil.4: 106. 1862,

nom. illeg., not Endl. 1833. TYPE: Ecuador, at

the foot of Chimborazo, Spruce (holotype, K!; frag.,

BM!; photos, F & GH of K).

Dryopteris microsora Kuntze, Revis. gen. pi. 2: 813.

1891, nom. nov. for Nephr odium microsorum

Hooker and with the same type.

Dryopteris leptosora C. Chr., Index fil. 274. 1905, nom.

nov. for Nephrodium microsorum Hooker and with

the same type.

Megalastrum leptosorum (C. Chr.) Smith & Moran,

Amer. Fern J. 77: 128. 1987.

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with orange to brown, linear

scales, these narrowly clathrate, to 2 cm long, the

margins conspicuously setose, the setae often bi-

furcate. Leaves to 1 m long and 30 cm broad, the

axes on the abaxial side abundantly provided with

whitish, pluricellular trichomes 0.1-0.2 mm long.

14

FIELDIANA: BOTANY

Lamina firm-herbaceous, the abaxial surface

abundantly and minutely pubescent, 1-pinnate-

pinnatisect to 2-pinnate, the rachis and costae

abaxially provided with abundant reddish brown,

filiform, setulose scales, these 2-4 mm long. Pin-

nae incised nearly or quite to the costae, the ul-

timate segments entire to crenate, with scattered

marginal trichomes ca. 0.1 mm long. Veins evi-

dent, although sometimes indistinct. Sori supra-

medial to somewhat inframedial. Indusia lacking.

Thus far known in Peru from the single speci-

men cited below: beneath a moist rock wall, 2800

m, Junin.

Ecuador; Peru.

Megalastrum microsorum can be distinguished

from its nearest relatives by the abundant, though

minute, pubescence of the abaxial surface, on and

between the veins. An interesting feature is the

character of the stem, petiole, and (sometimes)

rachis scales. These are conspicuously setulose as

in many species of the genus, but the setae are

commonly bifid at the apex. This condition also

has been observed on petiole scales of M. platy-

lobum, although only rarely.

Junin: Paucartambo, Woytkowski 6742 (us).

4. Megalastrum platylobum (Baker) Smith &

Moran, Amer. Fern J. 77: 128. 1987.

Polypodium platylobum Baker, Syn. fil. 307. 1867.

TYPE: Mt. Guayrapurima, near Tarapoto (San

Martin), Spruce 4656 (holotype, K!; isotypes, BM!,

K!, P!).

Polypodium tarapotense Baker, Syn. fil. ed. 2:505. 1 874.

TYPE: based on one sheet of type of P. platylo-

bum (K.!, see discussion below).

Dryopteris platyloba (Baker) C. Chr., Index fil. 285.

1905.

Dryopteris tarapotensis (Baker) C. Chr., Index fil. 297.

1905.

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with orange to brown, linear

scales, these narrowly and obscurely clathrate, to

1 cm long, the margins setulose, the setae occa-

sionally bifid at apex. Leaves to 1.5 m long and

40 cm broad, the axes on the abaxial side densely

covered with whitish trichomes of mixed lengths,

many ca. 1 mm long, and many more only 0.1

mm long. Lamina firm-herbaceous, 1-pinnate-

pinnatisect, or 2-pinnate near the base, occasion-

ally bearing some scattered, minute (0.05 mm)

trichomes on the tissue abaxially, the rachis abax-

ially provided with scattered, reddish brown, fi-

liform scales, these 1-1.5 mm long and with se-

tulose margins. Pinnae incised nearly or quite to

the costae, the ultimate segments entire to crenate

or rarely shallowly lobed, their margins amply cil-

iate with trichomes ca. 0.5 mm long. Veins evi-

dent. Sori supramedial. Indusia lacking.

Thus far known in Peru only from the type col-

lections, San Martin.

Venezuela; Peru.

At Kew there are five sheets of Spruce 4656.

Three, from Hooker's herbarium, represent the

holotype of Polypodium platylobum; the two oth-

ers, in another folder and marked "sheet 1 " and

"sheet 2," are isotypes. After having described P.

platylobum, Baker seven years later described P.

tarapotense, based on one of the sheets of Spruce

4656, which contains the distal portion of the leaf,

plus the petiole of a species of Cyatheaceae (Cnem-

idaria speciosal). For a full discussion, see C.

Christensen's monograph (1913, p. 110).

Megalastrum platylobum can be further distin-

guished from closely related species by the mixed

trichomes on the axes abaxially, especially on the

costae: many spreading pluricellular ones ca. 1 mm

long, plus a dense covering of 1 (-2)-cellular ones

ca. 0.1 mm long. Axes trichomes of M. hirsuto-

setosum are all long (1-2 mm), and those of others

in the species complex are short (commonly 0.3-

0.5 mm).

5. Megalastrum biseriale (Baker) Smith & Mo-

ran, Amer. Fern J. 77: 127. 1987. Figure 2.

Polypodium biseriale Baker, Syn. fil. 309. 1867. TYPE:

Ecuador, Mt. Tungurahua, Spruce (holotype, K!).

Dryopteris biserialis (Baker) C. Chr., Index fil. 254.

1905.

Ctenitis biserialis (Baker) Lell., Fern gaz. 11: 108. 1975.

Plants terrestrial. Stem erect, this and the petiole

base densely provided with orange or light brown,

linear scales, these obscurely clathrate, to 1.5 cm

long, the margins setulose. Leaves to 1 m long and

30 cm broad, the axes on the abaxial side sparsely

provided with whitish trichomes 0.2-0.6 mm long,

or glabrate. Lamina firm-herbaceous, the tissue

glabrous and eglandular, 1 -pinnate-pinnatisect to

2-pinnate, the rachis and costae abundantly pro-

vided on the abaxial side with reddish brown, fi-

liform scales, these 3-4 mm long (on costae 1-2

mm), margins remotely setulose. Pinnae incised

nearly or quite to the costae, ultimate segments

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

15

entire to crenate or shallowly lobed, with scattered

marginal trichomes 0. 1 mm long. Veins evident.

Sori mostly inframedial. Indusia lacking.

In forests and wooded ravines, 600-1800 m,

Pasco, Junin, Madre de Dios.

Panama; Colombia; Ecuador; Peru.

Pasco: San Nicolas, Pichis Trail (as Junin), Killip &

Smith 26021 (us). Junin: Chanchamayo Valley, C.

Schunke 45, 87 (F, us). Schunke Hacienda, above San

Ramon, C. Schunke A227 (us). La Merced, Chancha-

mayo, Soukup 1032 (F). Madre de Dios: Prov. Manu,

Atalaya, Foster & Wachter 7423 (F, MO).

6. Megalastrum hirsutosetosum (Hieron.) Smith

& Moran, Amer. Fern J. 77: 128. 1987.

Dryopteris hirsute- setosa Hieron., Hedwigia 46: 343,

t. 6. 1907. TYPE: Ecuador, plateau above All-

payacu between Banos and Jivaria de Pintuc, Stu-

bel 903 (holotype, B!; frag. & photo, BM).

Ctenitis hirsuto-setosa (Hieron.) Lell., Proc. Biol. Soc.

Wash. 89: 709. 1977.

Plants terrestrial. Stem erect, provided at apex

with a dense cluster of deep orange to reddish

brown scales, these obscurely clathrate, to 1 cm

long, with narrow, often blackish, densely setulose

margins. Leaves caespitose, to 1.2 m long and 30

cm broad, the axes and veins on both sides mod-

erately to abundantly provided with rigid, pluri-

cellular trichomes 1-2 mm long. Lamina 1-pin-

nate-pinnatisect to 2-pinnate, the axes lacking

scales, or the rachis with a few, scattered, filiform

ones. Pinnae incised nearly or quite to the costae,

the ultimate segments subentire, crenate or very

shallowly lobed, the margins with abundant

spreading trichomes to 1 mm long. Veins indis-

tinct to obscure, 1 -several-forked. Sori usually me-

dial between costule and margin. Indusia lacking.

In forests, 1000-1500 m, Amazonas, Huanuco,

Pasco, Junin, Ucayali.

Colombia; Ecuador; Peru.

This can be easily distinguished from all the less

dissected species of Megalastrum by the abundant,

spreading trichomes which clothe the axes and

veins, abaxially and adaxially. There is a single

specimen from San Martin (Knapp & Mallet 7022,

MO) that closely resembles M. hirsutosetosum, in

that the axes and margins have abundant, long

trichomes. However, the tissue is glandular abax-

ially, and the axes are conspicuously filiform-scaly.

The few scattered sporangia all appear to be bar-

ren, so the plant is probably a hybrid, perhaps

involving M. platylobum. j-v

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

2499 (F, MO). Huanuco: Prov. Leoncio Prado, Tingo

Maria, Aguilar 25 (USM). Pampayacu, Kanehira 184 (GH).

Pasco: Pichis Trail, San Nicolas (as Junin), Killip & Smith

26037 (us). Junin: La Merced, Chanchamayo, C. Schunke

22 (uc). Ucayali: Prov. Coronel Portillo (as Loreto), Sin-

chono, between Tingo Maria and Pucallpa, Aguilar 865

(USM).

7. Megalastrum subincisum (Willd.) Smith &

Moran, Amer. Fern J. 77: 129. 1987.

Polypodium subincisum Willd., Sp. pi. ed. 4, 5: 202.

1810. TYPE: Venezuela, Caracas, Bredemeyer

(holotype, B, Herb. Willd. 19701; photos, F, us).

Dryopteris subincisa (Willd.) Urban, Symb. antill.

4: 19. 1903.

Ctenitis subincisa (Willd.) Ching, Sunyatsenia 5: 250.

1940.

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with bright brown to casta-

neous, linear to filiform scales, these clathrate, 1.5-

2.5 cm long, the margins setose. Leaves to 3 m

long and 1.2m broad, the axes adaxially provided

with a dense covering of terete, curved trichomes

mostly 0.5-0.8 mm long. Lamina 2-pinnate-pin-

natisect to 3-pinnate-pinnatifid, the costules, cos-

tae, and rachis abaxially glabrous, or with tri-

chomes scattered and varying in length from 0.3

to 0.6 mm long, the scales of the rachis and costae

dark brown, clathrate, linear or filiform (from a

sometimes dilated base) and mostly 2-6 cells wide,

the margins commonly setose. Pinnae with tissue

between the veins typically glabrous abaxially, but

occasionally sparsely to abundantly puberulent.

Sori medial to inframedial on the segments. In-

dusia lacking.

In rain forests and wooded ravines, 500-2 1 00

m, Amazonas, San Martin, Junin, Cuzco.

West Indies; southern Mexico to Panama; Ven-

ezuela; Colombia to Bolivia.

This is one of the most widely distributed spe-

cies in the genus, and is part of a complex that is

rather variable in laminar indument. This group

includes M. vastum from Peru and the Andes and

M. spectabile (Kaulf.) Smith & Moran from south-

ern South America. Christensen stated (1920, p.

68) that M. vastum might be merely a variety of

M. subincisa; the other species appears to differ in

no greater degree. The only diagnostic features, as

16

FIELDIANA: BOTANY

noted in the key, are those of size and abundance

of trichomes and shape and margin of scales, and

there is some variability even in these.

Although pubescence is generally confined to the

axes and veins on the above species, some speci-

mens are sparsely to abundantly puberulent on the

tissue between the veins. This condition has been

noted in both M. subincisum and M. vastum. The

entire species complex is in need of study through-

out its range; meanwhile the two species are main-

tained as distinct for purposes of this treatment.

Amazonas: Prov. Bagua, valley of Rio Maranon above

Cascadas de Mayasi, Wurdack 1851 (us). San Martin:

"In Monte Campana, prope Tarapoto," Spruce 4340 (K).

Junin: Schunke Hacienda, above San Ramon, C. Schunke

A225 (us). Cuzco: Prov. Urubamba, along Rio Urubam-

ba near town of Machu Picchu, Tryon & Try on 5412 (F,

GH). Prov. Paucartambo, Kosnipata-Pilcopata, Vargas

11284(GH).

. Megalastrum vastum (Kunze) Smith & Moran,

Amer. Fern J. 77: 129. 1987.

Polypodium vastum Kunze, Linnaea 9: 50. 1834.

TYPE: Peru (Huanuco), "inter Pampayaco (Pam-

payacu) et Cocheros et ad Ventanilla de Cassapi,"

Poeppig 217 (holotype, B!; isotypes, B!, L; photos,

F & us of L).

Dryopteris vasta (Kunze) Hieron., Hedwigia 46: 347.

1907.

Dryopteris mollicoma C. Chr., Kongel. Danske Vi-

densk. Selsk. Skr. Naturvidensk. Afd. ser. 8, 6:

75. 1920. TYPE: Ecuador, "in silv. suband. ori-

ent." Oyacachi, Sodlro (holotype, P; frag., BM; iso-

type, A!; photo, F of BM).

Ctenitis mollicoma (C. Chr.) Ching, Sunyatsenia 5:

250. 1940.

Megalastrum mollicomum (C. Chr.) Smith & Moran,

Amer. Fern J. 77: 128. 1987.

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with bright brown to casta-

neous, lanceolate to filiform scales, these clathrate,

1-3 cm long, the margins denticulate to setulose.

Leaves to 2 m long and 0.8 m broad, the axes

adaxially provided with curved trichomes mostly

0.5-0.8 mm long or glabrescent. Lamina 2-pin-

nate-pinnatisect to (proximally) 3-pinnate-pin-

natifid, the costules, costae, and (often) the rachis

densely and regularly puberulent abaxially, the tri-

chomes 0.05-0.2 mm long, the scales of the rachis

and costae dark brown, clathrate or subclathrate,

lanceolate to filiform and mostly 3-10 cells wide,

the margins entire to denticulate or sometimes

setose. Pinnae with tissue between the veins typ-

ically glabrous abaxially, but sometimes sparsely

to abundantly puberulent, or occasionally hirsute,

adaxially. Sori typically medial. Indusia lacking.

Dense forests, 750-1850 m, Amazonas, Hua-

nuco, Junin, and Ayacucho.

Costa Rica; Colombia to Bolivia; Paraguay.

Killip & Smith 22779, cited below, differs from

typical M. vastum in the dense puberulence abax-

ially on the tissue between the veins. In the M.

subincisum complex, some scattered trichomes are

often evident on the laminar tissue, but never so

conspicuously as seen on this particular specimen.

This character should be considered, along with

other conditions of indument, in future studies of

the various species complexes of both Ctenitis and

Megalastrum.

Probably M. vastum should be recognized mere-

ly as a variety of M. subincisum, under which see

further discussion. Only the type and a few other

collections have been found in Peru; the species

seems to be most abundant in Ecuador.

Amazonas: Prov. Bagua, 1 2 mi E of La Peca, Barbour

2504 (F, MO). Huanuco: Cushi, Macbride 4846, 4855 (F,

us). Junin: Prov. Chanchamayo, Chilpex, 26 km S of

San Ramon, Smith & Palacios 2639 (F, MO). Ayacucho:

"Aina" (Ayna), between Huanta and Rio Apurimac, Kil-

lip & Smith 22779 (us).

9. Megalastrum pulverulentum (Poiret) Smith &

Moran, Amer. Fern J. 77: 129. 1987.

Polypodium pulverulentum Poiret in Lam., Encycl. 5:

555. 1804. TYPE: Plumier, Traite foug. Amer. t.

34. 1705, based on a plant from Hispaniola.

Polypodium karstenianum Klotzsch, Linnaea 20: 390.

1847. TYPE: "Colombia" (Venezuela), Karsten

II, 3 (holotype, B; probable isotype, HBG; photo,

F & us of HBG).

Dryopteris karsteniana (Klotzsch) Hieron., Hedwigia

46: 348. 1907.

Dryopteris pulverulenta (Poiret) C. Chr., in Urban,

Symb. antill. 9: 305. 1925.

Ctenitis pulverulenta (Poiret) Copel., Gen. fil. 124.

1947.

Ctenitis karsteniana (Klotzsch) Vareschi, Flora Ven-

ezuela 1: 405. 1969.

Plants terrestrial. Stem erect, arborescent, in Peru

to 1 m tall, this and the petiole base densely clothed

with orange to brown, linear scales, these sub-

clathrate, 1-2.5 cm long, the margins entire to

remotely setulose. Leaves to 3 m long and 1.5 m

broad, the axes on both sides moderately to abun-

dantly pilose. Lamina 3-pinnate to nearly 4-pin-

nate-pinnatifid, the rachis deciduously pilose and

with scattered scales 1-2 mm long, these with con-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

17

spicuously setose margins. Pinnae commonly

2-pinnate-pinnatisect, but basal ones often basi-

scopically enlarged and more highly dissected, the

costae and veins moderately to abundantly pro-

vided with spreading pluricellular trichomes mostly

0.7-1.5 mm long or a little shorter adaxially. Sori

inframedial, medial, or rarely supramedial. In-

dusia lacking.

In forests, 540-1800(-2700) m; San Martin,

Cuzco.

Greater Antilles; southern Mexico to Costa Rica;

Venezuela; Colombia to Bolivia.

This, like M. hirsutosetosum, is distinguished by

the long (1-2 mm), spreading trichomes on the

axes and veins. However, leaves of the latter are

never more than 2-pinnate, and scales of the pet-

iole base have blackish, densely setulose margins.

Leaves of M. pulverulentum are often 4-pinnate-

pinnatisect at base, and the petiole scales are con-

colorous, with subentire to remotely denticulate

margins.

The few specimens found thus far in Peru are

var. pulverulentum, a common variety widely dis-

tributed throughout the Neotropics. Variety hey-

dei (C. Chr.) Smith & Moran, with essentially gla-

brous laminae, is confined to Guatemala (or

possibly also in Ecuador).

Young & Leon 4958, cited below, appears to be

intermediate between M. pulverulentum and the

Central American M. pansamalense (C. Chr.)

Smith & Moran, the latter differing chiefly in its

denser covering of long trichomes and its broad,

entire laminar scales. The Young & Leon collec-

tion has much smaller leaves, and abundant lan-

ceolate to ovate scales with entire margins, but is

similar in every other character to M. pulverulen-

tum. Although fully mature, the cited specimens

have leaves only 50 cm long and 18 cm broad,

whereas both of the above species have leaves from

1 to 3 m long. It is possible that Young & Leon

4958 (2650-2750 m) represents a depauperate

specimen or a high elevation variant; it is uncom-

mon to find M. pulverulentum above 1500 m.

San Martin: Tarapoto, Spruce 47 18 (BM, K, p). Prov.

Mariscal Caceres, Rio Abiseo National Park, Young &

Leon 4958 (F, GH). Cuzco: San Miguel, Urubamba Val-

ley, Cook & Gilbert 1128 (us). Prov. Paucartambo, Ha-

cienda Villa Carmen, Vargas 14685 (GH). Department

unknown: Peru, undesignated locality, Soukup 296 (F).

10. Megalastrum andicola (C. Chr.) Smith &

Moran, Amer. Fern J. 77: 127. 1987.

Plants terrestrial. Stem erect, this and the petiole

base densely clothed with orange to brown, linear

to lanceolate scales, these subclathrate, 1-2.5 cm

long, the margins setulose. Leaves to 1.5 m long

and 40 cm broad, the axes on both sides amply to

abundantly pubescent. Lamina 2-pinnate-pinnati-

sect to 3-pinnate-pinnatifid, the rachis and costae

sparsely to amply scaly, the scales orange to brown,

linear, 4-5 mm long, their margins denticulate to

subentire. Pinnae mostly 1 -pinnate-pinnatisect, but

basal ones more deeply divided and enlarged at

the base basiscopically, the ultimate segments mi-

nutely pubescent on both sides on the veins, and

sometimes on the intervening tissue. Indusia per-

sistent, light to dark brown, 1 mm or more in

diameter, often glandulose.

This is part of a variable complex of neotropical

species that is in need of further study. It is one

of the few indusiate species of Megalastrum, and

the only indusiate one in Peru. A number of va-

rieties and forms have been described, based chief-

ly on differences in length and abundance of in-

dument. Megalastrum andicola is probably only

varietally distinct from the West Indian M. vil-

losum (L.) Holttum; the latter apparently differs

only in its larger size, greater dissection of lamina,

and in the fewer and early deciduous scales on the

axes. Pending future study of the species complex,

M. andicola and one other form are tentatively

recognized in Peru.

Key to Forms of Megalastrum andicola

a. Trichomes on axes, veins, and leaf tissue ca. 1 mm long lOa. f. andicola

a. Trichomes on axes, veins, and leaf tissue 0.2-0.3 mm long lOb. f. lehmannianum

18

FIELDIANA: BOTANY

lOa. Megalastrum andicola f. andicola

Nephrodium villosum var. opacum Hieron., Bot. Jahrb.

Syst. 34: 446. 1904, LECTOTYPE (designated

here): Colombia, Fusagasuga, Lindig 159 (B).

Nephrodium villosum f. spruceanum Hieron., Bot.

Jahrb. Syst. 34: 446. 1904. LECTOTYPE (des-

ignated here): Ecuador, Spruce 5295 (B!; isotypes,

c, K!).

Dryopteris andicola C. Chr., Kongel. Danske Vidensk.

Selsk. Skr. Naturvidensk. Afd. ser. 8, 6: 88. 1920.

TYPE: Ecuador, Spruce 5295 (holotype, B!; iso-

types, c, K!).

Dryopteris andicola f. spruceana (Hieron.) C. Chr.,

reference as above, p. 89.

Ctenitis andicola (C. Chr.) Ching, Sunyatsenia 5: 250.

1940.

In rain forests and cloud forests, 750-2300 m,

Amazonas, San Martin, Junin, and Pasco.

Colombia; Ecuador; Peru.

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

2587 (F, uc). Prov. Bongara, hills 1-5 km SSE of Yam-

brasbamba, Wurdack 1030 (GH, us). San Martin: Tara-

poto, LI. Williams 5985 (F). Pasco: Oxapampa, Soukup

3354 (GH, us). Junin: Prov. Tarma, Valle del Rio Chan-

chamayo, Esposto 663 (USM).

lOb. Megalastrum andicola f. lehmannianum

(Hieron.) Stolze, comb. nov.

Nephrodium villosum f. lehmannianum Hieron., Bot.

Jahrb. Syst. 34: 446. 1904. LECTOTYPE (des-

ignated here): Colombia, Lehmann 7369 (holo-

type, B; isolectotypes, K!, us!).

Dryopteris andicola f. lehmanniana (Hieron.) C. Chr.,

Bot. Jahrb. Syst. 34: 89. 1904.

Thus far represented in Peru by the single col-

lection cited below: Podocarpus forest, 1 880-1 950

m, Junin.

Colombia; Ecuador; Peru.

Hieronymus originally distinguished his new

form by size of indusia and number of sori on a

segment. It has been seen that these characters are

too variable and inconsistent to be of taxonomic

value. The only other difference observed between

f. lehmannianum and f. andicola is the length of

trichomes on the lamina, but at least this character

is consistent and conspicuous. The tissue, veins,

and minor axes of f. andicola are densely covered

on both sides with whitish, spreading trichomes

ca. 1 mm long. In f. lehmannianum equivalent

axes are as densely pubescent, but the trichomes

are never more than 0.3 mm long, and those on

the tissue abaxially are noticeably less abundant.

Junin: Prov. Chanchamayo, Rio Rondayacu, 45 km

from San Ramon, D. Smith el al. 2615 (F, GH, MO).

III. Triplophyllum

Triplophyllum Holttum, Kew Bull. 41: 239. 1986.

TYPE: Triplophyllum protensum (Sw.) Holt-

tum (Aspidium protensum Sw.). Figure 3.

Plants terrestrial. Stem creeping, sparsely to

moderately provided with narrow, nonclathrate

scales, these commonly 1—4 mm long. Leaves

1 -pinnate-pinnatifid to nearly 4-pinnate (as to bas-

al portion), borne at approximate to subdistant

intervals, monomorphic, to ca. 1 .6 m long and 0.8

m broad, long-petiolate, not articulate to the stem.

Lamina deltoid-pentagonal (to somewhat elongate

in a few Old World species), the basal pinnae con-

spicuously larger than adjacent ones, and more

than half as long as the rest of the lamina, provided

especially on the axes with scales and ''''Ctenitis

hairs," the latter being reddish brown, un-

branched, articulate trichomes, with 2-4 cells and

0.1-0.4 mm long. Rachis not sulcate or, if slightly

so, the adaxial groove not open to admit the groove

of the pinna stalk, the adaxial ridges (if any) not

continuous with the ridge of the pinna axis, scales

few or lacking. Veins free, or in a few species rarely

anastomosing, but without free veins in the are-

oles. Sori roundish, borne along the veins or at

their tips, receptacle somewhat elevated, lacking

paraphyses. Indusia reniform (in Peru species) or

rarely lacking. Spores monolete, roundish to ellip-

soidal, the surface bearing thin, winglike ridges.

Triplophyllum is a genus of 20 tropical species,

five of them in America, with but a single variety

occurring in Peru, and one species perhaps to be

expected. The more obvious characters separating

it from Ctenitis are seen in the key to genera of

Dryopteridaceae. In addition, Ctenitis often has

cylindric glands on indusia, and on axes and tissue

between the veins. If glands are present on the

lamina or indusia in Triplophyllum, they are

spherical.

According to Holttum ( 1 986) there are two oth-

er varieties of T.funestum: var. perpilosum Holtt.

of Colombia, with many trichomes between the

veins on both surfaces, and var. hirsutum Holtt.

of Guyana, with trichomes to 1 mm long on the

axes.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

19

5 cm

FIG. 3. Triplophyllum funestum var. funestum: a, habit; b, petiole scale; c, ultimate segment, abaxial side (a

b from Petelot s.n., Brazil, F; c from Klug 174, F.)

20

FIELDIANA: BOTANY

Reference

HOLTTUM, R. E. 1986. Studies in the fern-genera

allied to Tectaria Cav., 5. Triplophyllum, a new

genus of Africa and America, Kew Bull., 41:

237-260.

1 . Triplophyllum funestum (Kunze) Holttum var.

funestum, Kew Bull. 41: 256. 1986. Figure 3.

Aspidiumfunestum Kunze, Linnaea 9: 96: 1 834. TYPE:

"ad Ega, Brasiliae, lect. 1 832," Poeppig(ho\otype,

presumably w).

Nephrodium funestum (Kunze) Hooker, Sp. fil. 4: 129,

t. 259. 1862.

Dryopteris protensa var.funesta (Kunze) C. Chr., Kon-

gel. Danske Vidensk. Selsk. Skr. Naturvidensk.

Afd., ser. 8, 6: 91: 1920.

Ctenitis protensa (Sw.) Ching, var. funesta (Kunze)

Proctor, Rhodora 63: 34. 1961.

Plants terrestrial. Stem creeping, this and the

petiole and rachis sparsely to moderately provided

with reddish brown, lanceolate scales, these non-

clathrate, 1—4 mm long. Leaves approximate to

somewhat spaced along the stem, to 1 m long and

50 cm broad, with ''''Ctenitis hairs" ample on the

axes and few on the veins and segment margins.

Lamina deltoid-pentagonal, 1 -pinnate-pinnatifid

to 2-pinnate-pinnatifid, but 3-pinnate-pinnatifid

as to the greatly enlarged basal pinnae. Central

pinnae commonly 1 -pinnate-pinnatifid, the basal

pinnules more strongly developed acroscopically,

ultimate segments obtuse, the basal pinnae often

nearly as large as the rest of the lamina, much

more strongly developed basiscopically. Veins

branched in the ultimate segments, terminating at

or near the margin, with slender tips. Sori borne

near the segment margins. Indusia orange to red-

dish brown, conspicuous, persistent, with minute

pluricellular trichomes.

In wet soil of rain forests, sea level to 700 m,

San Martin, Loreto, Pasco, and Madre de Dios.

Puerto Rico; Lesser Antilles; Nicaragua to Pan-

ama; Trinidad; the Guianas to Colombia, and

southward to Brazil and Bolivia.

Some specimens from Peru approach T. acuti-

lobum Holttum, a large species of Brazil, with bas-

al pinnae 44 cm long or more, axes adaxially gla-

brous, and segments and lobes mostly acute. Allard

22003 and Vargas 18629 (cited below) lack ad-

axial trichomes and have some segments acute,

but both have small leaves with basal pinnae about

1 5 cm long. Other specimens examined have one

or the other of these features, but not both. Ob-

viously these characters are too variable to serve

as a basis for the further division of var. funestum.

San Martin: On ridge east of Tingo Maria, Allard

22003 (GH, us). Loreto: Mishuyacu, near Iquitos, King

174 (F, us). Prov. Maynas, Rio Momon, Sounders 1365,

1367, 1368 (GH). Pasco: Prov. Oxapampa, Quebrada

Castilla on Omaiz River, Leon & Young 1020 (GH). Ma-

dre de Dios: Prov. Tambopata, Tambopata Nature Re-

serve, Harbour 4921, 5189 (F). Prov. Tambopata Vargas

18629 (GH).

Comments

Triplophyllum dicksonioides (Fee) Holttum, Kew

Bull. 4: 257. 1986.

Aspidium dicksonioides Fee, Crypt vase. Bresil 1 : 143,

t. 49. 1869. TYPE: Northern Brazil, Rio Negro

near San Gabriel, Spruce 2129 (holotype, not lo-

cated; isotypes, BM, K), cited in error (fide Chris-

tensen) as Glaziou 2129.

Dryopteris protensa (Sw.) C. Chr. var. dicksonioides

(Fee) C. Chr., Kongel. Danske Vidensk. Selsk.

Skr. Naturvidensk. Afd., ser. 8, 6: 93: 1920.

This species occurs from the Guianas to Colom-

bia, and in northwest Brazil, so there is good rea-

son to believe it may be found in Amazonian Peru,

perhaps in Loreto where T. funestum grows. It is

distinguished from the latter by the tiny, spherical

glands on the indusia and lamina, by the sori,

which are mostly apical on the veins, and by the

more highly dissected lamina (up to 4-pinnate)

with much smaller ultimate segments. In contrast,

T. funestum var. funestum is minutely pubescent

on the veins and indusia, but eglandular; sori are

seldom apical on the veins, and the lamina is

1 -pinnate-pinnatifid to (in basal pinnae) 3-pin-

nate-pinnatifid.

IV. Tectaria

Tectaria Cav., Anales Hist. Nat. 1: 115. 1799.

TYPE: Tectaria trifoliata (L.) Cav. (1802)

(Polypodium trifoliatum L.). Figure 4.

Aspidium Sw., J. Bot. (Schrader) 1800(2): 4, 29. 1802,

nom. superfl. for Tectaria and with the same type.

Stem erect or decumbent, usually stout, to mod-

erately long-creeping and slender, bearing scales.

Leaves ca. 1 0 cm to 2 m long, petiole continuous.

Lamina simple and entire to deeply lobed, or to

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

21

FIG. 4. Tectaria incisa var. incisa: a, portion of stem and petiole; b, lamina; c, portion of pinna, abaxial side,

(a from Tryon & Tryon 5206, F, b from Alfaro s.n., Costa Rica, F, c from Moran 3653, F.)

22

FIELDIANA: BOTANY

3-pinnate-pinnatifid, glabrous, scaly, or pubes-

cent, usually monomorphic, or dimorphic with the

fertile usually more erect, longer and less expanded

than the sterile, pinnae not articulate, adaxial side

of the costa raised. Veins usually anastomosing,

usually with included free veinlets, rarely free. Sori

(in Peru) usually roundish, borne on the veins and

covered by reniform to peltate (rarely some elon-

gate) indusia, or exindusiate and roundish or rarely

some elongate, not paraphysate (elsewhere the son

may be linear, rarely on a marginal projection or

the sporangia partly acrostichoid). Spores ellip-

soidal, monolete, ridged, cristate or echinate.

Tectaria is a large genus of about 150 species,

some 20 of them in America and six in Peru. It is

pantropical and sometimes subtropical. The sorus

and lamina architecture is unusually variable in

the genus, and several segregate genera have been

recognized on the basis of these variations. Tryon

and Tryon (1982) list the ones that occur in Central

America, the Greater Antilles, and northern South

America.

Tectaria is a difficult genus in the Andes because

of a lack of critical studies, the uncertainty of the

application of some names, the probability of un-

recognized hybrids, and the lack of cytological in-

formation. This treatment, based largely on Pe-

ruvian materials, will need to be refined, especially

as field studies of populations and cytological in-

formation on the different morphological kinds

become available. The cytological complexities of

American Tectaria, with diploids, triploids, and

tetraploids, are partly indicated by Jermy and

Walker (1985 and references).

Soral variation is shown in Mexia 8217, MO (T.

antioquoiand) and Smith et al. 1709, MO (T. li-

zarzaburui); in both, the sorus and the indusia

vary from roundish to reniform to quite elongate.

References

JERMY, A. C., AND T. G. WALKER. 1985. Cyto-

taxonomic studies of the ferns of Trinidad. Bull.

Brit. Mus. (Nat. Hist.) Bot., 13: 133-286.

MORTON, C. V. 1966. The Mexican species of

Tectaria. Amer. Fern J., 56: 120-137.

TRYON, R. M., AND A. F. TRYON. 1 982. Tectaria,

pp. 470-48 1 , in Ferns and allied plants, Spring-

er-Verlag, New York.

Key to Species of Tectaria

a. Veins mostly or wholly free 1 . T. brauniana

a. Veins fully anastomosing, except for included free veinlets b

b. Lamina 1 -pinnate or more complex, or the rachis fully alate, or if the lamina simple then with a

pair of basal lobes and an abrupt base c

c. Sori of the lateral pinnae or large pinna-segments in 2 series between the costules, each series

extending from the costa to the margin in a usually regular line, or a third series present only

near the costa; sori indusiate, most or all of the indusia with a sinus, often attached laterally

d

d. Lamina simple, with 2 basal lobes, or with a single pair of stalked or sessile pinnae, or with

2 or more such pairs and then the basal pinnae entire to shallowly lobed or pinnatifid, these

and the usually entire pinnae distally mostly with obtuse lobes 2. T. incisa

d. Lamina with 2 or more pairs of stalked or sessile pinnae, the basal pinnae deeply pinnatifid

or more complex, with 2 or more large, basiscopic lobes, these pinnae and distal ones mostly

regularly lobed with acute to acuminate, sometimes subacute or obtuse lobes

3. T. lizarzaburui

c. Sori of the lateral pinnae or large pinna-segments in 3-6, usually 4 or 5, series between the

costules, each series extending from the costa toward the margin in a regular or sometimes

irregular line; sori indusiate or not e

e. Basal pinnae stalked or sessile, the rachis immediately above them not alate; sori indusiate,

or most exindusiate, or a few small indusia present; proliferous buds sometimes present at

the base of the pinnae 4. T. antioquoiana

e. Basal pinnae joined to the pair above by the alate rachis; sori exindusiate; proliferous buds

absent 5. T. draconoptera

b. Lamina entire, long-decurrent at the base; sori exindusiate 6. T. plantaginea

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

23

1 . Tectaria brauniana (Karsten) C. Chr., Index fil.

suppl. 3: 177. 1934.

Aspidium braunianum Karsten, Fl. columb. 1: 63, /.

31. 1859. TYPE: Colombia, "Villavizencio" (Vi-

llavicencio). Karsten (not located).

Stem rather long-creeping. Leaves ca. 25-70 cm

long. Lamina mostly 2-pinnatifid, to 3-pinnatifid

at the base, gradually reduced at the apex, pu-

bescent on the axes on both sides but especially

abaxially; basal pinna-segments pinnatifid, more

developed on the basiscopic side; lateral pinna-

segments obtusely to sometimes acutely lobed;

proliferous buds absent. Veins mostly to usually

wholly free. Leaf-tissue glabrous to somewhat pu-

bescent. Sori indusiate, in a single series between

the costules and margin. Indusium with a broad

to narrow sinus, usually glabrous.

Primary forests, wet ravines, one label indicates

that the plant was an epiphyte, 250-800 m, Huan-

uco, Cuzco, and Madre de Dios.

Costa Rica to Bolivia.

The essentially free venation is distinctive for

this species, as is the 2-3 -pinnatifid lamina archi-

tecture.

Huanuco: Ca. 1 5 km N of Tingo Maria, Woytkowski

1363 (uc). Cuzco: Prov. Paucartambo, Cosnipata Valley,

N of Patria, Wachteret al. 196 (F). Madre de Dios: Prov.

Tambopata, Dist. Tambopata, Yung 21 (MO, uc).

2. Tectaria incisa Cav., Descr. pi. 249. 1802.

TYPE: Puerto Rico, comm. (and probably

collected by) Ventenant (holotype, MA, seen

by C. Chr. [Dansk Bot. Ark. 9(3): 14. 1937]).

Figure 4a-c.

Aspidium martinicense Sprengel, Anleit. Kenntn. Gew.

3: 133. 1804. TYPE (cited by Morton, 1966):

Dominican Republic, Poiteau (not located). Plu-

mier, Traite foug. Amer, t. 145, also cited by

Sprengel, is this species.

Tectaria martinicensis (Sprengel) Copel., Philipp. J.

Sci. 2C:410. 1907.

Stem ascending to erect. Leaves ca. 20 cm to 2

m long. Lamina simple, with 2 basal lobes, or with

2 basal entire pinnae and an entire terminal seg-

ment, or usually with 2 or more stalked or sessile

pinnae, then the basal pinnae entire or usually with

1 large, basiscopic lobe on 1 or both of the pinnae,

or with 2 lobes on 1 or both of the pinnae, or less

often with 2 or more large, basal lobes and smaller

lobes beyond on both sides; lateral pinnae usually

entire, sometimes lobed, the stalked terminal seg-

ment entire to variously lobed; proliferous buds

sometimes present at or near the base of the pin-

nae, or only 1 bud present; the stalked terminal

segment and some distal pinnae may be long-de-

current on the rachis. Veins fully anastomosing,

except for the included free veinlets. Leaf-tissue

glabrous to very short-pubescent to pilose on both

surfaces. Sori indusiate, the indusia mostly or all

with a sinus, often attached laterally, in 2 series

between costules from the costa to the margin, or

a third series present only near the costa, in usually

regular lines. Indusium glabrous or sometimes

shortly glandular-pubescent on the margin and

surface.

In primary forests, in cloud forests, on hillsides,

in ravines, often among rocks, along roadsides,

and sometimes on wet, shady cliffs, 100-2100 m,

Cajamarca south to Madre de Dios.

Tropical America.

In Peru Tectaria incisa is a highly variable spe-

cies in the architecture of the basal pinnae of the

lamina and of the stalked terminal segment, in the

presence or absence of proliferous buds at or near

the base of the pinnae, and in the pubescence or

lack of it on the leaf tissue. Population studies,

including information on developmental stages, as

well as on cytology and hybridization are needed

to determine the basis for the extreme variation

and the evolutionary validity of some of the vari-

ations.

The characters employed to define taxa in this

complex all vary from one extreme to another,

and sometimes a single collection will consist of

more than one named taxon. For these reasons the

species Tectaria incisa is treated broadly. The

principal variations with their names are distin-

guished below and by our annotations. This treat-

ment serves to organize the variation of the species

so that it may be studied more readily, and at the

same time avoids the formal recognition of groups

that may not be evolutionary units. Although these

groups are not recognized as taxa in Peru, in regions

to the north the circumstances may be different.

a. Typical Tectaria incisa has both basal pinnae,

or at least one of them, with a large, basiscopic

lobe, the two sides beyond are entire or nearly so,

the stalked terminal segment is usually lobed, of-

ten with two, sometimes one, large basal lobe(s),

the pinnae have the leaf-tissue glabrous or essen-

tially so above and beneath, small plants may have

the lamina with two basal lobes, or with two basal

24

FIELDIANA: BOTANY

pinnae and a large terminal segment. Specimens

have been annotated as Tectaria incisa.

This variation (var. incisa) occurs throughout

the range of the species in Peru, except for Caja-

marca, where it is not yet known.

Amazonas: Prov. Bagua, Rio Maranon, above Cas-

cadas de Mayasi, Wurdack 1861 (us). San Martin: Cha-

zuta, Rio Huallaga, Klug 4038 (F, GH, MO). Loreto: Prov.

Maynas, 10 km S of Iquitos, Tryon & Tryon 5206 (F,

GH, us). Above Pongo de Manseriche, Mexia 6343 (F,

GH, MO, uc, us). Huanuco: Prov. Huanuco, near conflu-

ence of Rio Huallaga and Rio Cayumba, Mexia 8304

(BM, F, GH, MO, uc, us). Cerca Tingo Maria, Aguilar 307

(USM). Pasco: Yapas, Pichis Trail (as Junin), Killip &

Smith 25584 (us). Junin: Prov. Tarma, 4 km N of La

Merced, Tryon & Tryon 5443 (GH, us). Ucayali: Prov.

Coronel Portillo, Bosque Nacional von Humboldt, Vas-

quez 3882 (F, MO). Ayacucho: Near Kimpitiriki, Killip &

Smith 22975 (GH, us). Cuzco: Prov. La Convention, Rio

Apurimac, above Boca de Tigre rapids, Davis et al. 1315,

1 317 (F, GH). Madre de Dies: Parque Nacional del Manu,

Cocha Cashu Station, Foster & Terborgh 6613 (GH).

b. Plants with leaves that are similar to typical

T. incisa may have the leaf tissue pubescent above

and beneath. These have sometimes been named

Tectaria incisa var. pilosa (Fee) Morton. The in-

dument varies from pilose to very short-pubescent

and from quite dense to very sparse. Specimens

have been annotated as Tectaria incisa-pilose. They

are scattered through the range of the species and

do not seem to represent a taxon.

This variation ranges in Peru from Cajamarca

south to Madre de Dios. A few collections are the

following:

Cajamarca: Woytkowski 6905 (us). Huanuco: Allard

21868 (us), 27577 (us), Ferreyra 10238 (GH). Loreto:

Wurdack2146 (GH), Mexia 6354 (GH, MO, uc, us). Pasco:

Killip & Smith 26516 (F, us), Leon 307 (USM). Madre de

Dios: Chavez 1846 (MO).

c. Proliferous buds are present at or near the

base of the pinnae on some or all leaves of a plant.

This variation has been named Tectaria incisa f.

vivipara (Jenm.) Morton, and as a species, Tectaria

vivipara Jermy & Walker. There may be tight, scaly

buds or the buds may have developed into small

plantlets; sometimes there is a single bud on the

lamina. This variation is sporadic in its distribu-

tion and does not seem to be a taxon. Collections

have been annotated as Tectaria incisa-vivipa-

rous.

In Peru it is known from Loreto, Huanuco, Pas-

co, and Madre de Dios. The following document

the occurrence of this variation in Peru:

Loreto: J. Schunke V. 2688 (F, GH, us), Mexia 6250

(F, GH, MO, uc, us). Huanuco: Gentry et al. 41435 (MO).

Pasco: (as Junin), Killip & Smith 26690 (F, us). Madre

de Dios: Foster et al. 3401 (F), Nunez 5728 (MO).

d. The basal pinnae, and sometimes those above,

are definitely lobed beyond the one or more large,

basal basiscopic lobes, and the stalked terminal

segment is lobed. This variation has been named

Tectaria incisa ssp. transiens Morton, and has been

treated as a species, Tectaria transiens (Morton)

A. R. Sm. The leaf-tissue may be pilose above and

beneath or glabrous. It does not seem to merit

recognition as a taxon in Peru. Specimens have

been annotated as Tectaria incisa "transiens."

It occurs from Cajamarca south to Madre de

Dios. The following represent this variation:

Cajamarca: Soukup 3813 (us). Amazonas: Barbour

2496 (MO). Loreto: /. Schunke 325 (F, GH, us). Madre

de Dios: Foster & Wachter 7420 (F, MO).

e. All of the pinnae and the stalked terminal

segment may be entire or nearly so, and proliferous

buds usually occur near or at the base of the pin-

nae, or a single bud may be present. This is Tec-

taria andina (Sodiro) C. Chr. The leaf-tissue is

glabrous on both surfaces. It occurs in Ecuador

and Peru and may represent a taxon. However, it

seems to intergrade with typical Tectaria incisa

and so is not treated formally here. Specimens

have been annotated as Tectaria incisa "andina."

It is known from Amazonas, Loreto, and Hua-

nuco in Peru. A few of the collections are:

Amazonas: Soukup 6611 (GH), Wurdack 1823 (GH, us).

Loreto: Mexia 6131a, in part: (GH, us). Huanuco: Skog

et al. 5141 (us), Gentry et al. 36148 (MO).

3. Tectaria lizarzaburui (Sodiro) C. Chr., Index

fil. suppl. 3: 181. 1934.

Nephrodium lizarzaburui Sodiro, Recens. crypt, vase.

Quit. 55. 1883. TYPE: Ecuador, Atacazo, Ha-

cienda Bolona, Sodiro (holotype, not located). Ec-

uador, Sodiro (K.!), photo GH may be authentic, it

is not wholly typical.

Stem ascending to erect. Leaves to 2.5 m long.

Lamina mostly 1 -pinnate, more complex at the

base; terminal stalked segment acutely lobed on

each side, with two large, acuminate basal lobes,

somewhat decurrent on the rachis (cuneate at the

base); basal pinnae usually deeply pinnatifid on

the basiscopic side with 2 to several large, acu-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

25

minate lobes, sometimes to 2-pinnate-pinnatifid,

the acroscopic side mostly acutely to subacutely

lobed; lateral pinnae mostly regularly, acuminate-

ly to obtusely lobed; proliferous buds absent. Veins

fully anastomosing, except for included free vein-

lets. Leaf-tissue glabrous to sometimes short-pu-

bescent abaxially. Sori indusiate, in 2 series be-

tween the costules from the costa to the margin,

in usually regular lines, or a third series present

only near the costa. Indusium with a sinus, often

attached laterally, glabrous to slightly pubescent

on the surface, the edge glabrous to ciliate.

In primary forests, in cloud forests, and in sec-

ondary growth, 1000-2000 m, Amazonas south to

Ayacucho.

Venezuela and Colombia, south to Peru.

This species is close to Tectaria incisa, especially

to variation "d" with the basal pinnae lobed be-

yond the large, basal lobe. That variant of T. incisa

differs, however, in having obtuse lobes and the

leaf is usually 1 m or less long. Tectaria lizarza-

burui has most lobes acuminate to subacute and

a large leaf to 2.5 m long.

Cajamarca: Prov. Santa Cruz, 3.5 km ENE of Mon-

teseco, Santisteban & Guevara 130 (F). Amazonas: Prov.

Bagua, 12 km E of La Peca, Harbour 2692 (F, MO, uc).

Prov. Bongara, 5 km N of N end of Lake Pomacocha,

on road to Rioja, Hutchison & Wright 6787 (GH, uc, us).

Pasco: Prov. Oxapampa, Rio El Tungui, Smith et al.

1709 (MO). Junin: Above San Ramon, C. SchunkeA217

(us), Killip & Smith 24639 (F, us). Chanchamayo Valley,

C. Schunke 36 (F, us), 576, 950 (F). Ayacucho: Prov. La

Mar, eastern Massif of the Cordillera Central, Dudley

11907 (GH).

4. Tectaria antioquoiana (Baker) C. Chr., Index

fil. suppl. 3: 177. 1934.

Nephrodium sodiroi Baker, J. Bot. 15: 16. 1877. TYPE:

Andes of Ecuador, Sodiro (holotype, Ecuador, fl.

Pilaton, Sodiro, K!; photo, GH), nom. rejec. (see

discussion).

Nephrodium antioquoianum Baker, J. Bot. 19: 205.

1881. TYPE: Colombia, Antioquia, Kalbreyer

1806 (holotype, K!; photo, GH). The epithet al-

tered to antioquianum by Baker, Ann. Bot. (Lon-

don) 5: 329. 1891.

Polypodium haynaldii Sodiro, Recens. crypt, vase.

Quit. 61. 1883. TYPE: Ecuador, San Miguel de

los Colorados, Sodiro (holotype, not located; iso-

type, K!; photo, GH).

Tectaria sodiroi (Baker) Maxon, Proc. Biol. Soc. Wash.

43: 88. 1930, nom. rejec.

Tectaria haynaldii (Sodiro) C. Chr., Index fil. suppl.

3: 180. 1934.

Stem ascending to erect. Leaves ca. 50 cm to 1 .5

m long. Lamina 1 -pinnate, with 1 to 2 pairs of

stalked or sessile pinnae; stalked terminal segment

and adnate large pinna-segments, when present,

long-decurrent on the rachis, the stalked terminal

segment usually lobed, sometimes entire; basal

pinnae entire to usually with 1, rarely 2, large,

basal, basiscopic lobes, not connected to the pair

above by an alate rachis; proliferous buds rarely

present, usually absent. Veins fully anastomosing,

except for included free veinlets. Leaf-tissue gla-

brous above and beneath. Sori usually indusiate,

with a sinus, rarely a few peltate, in 3-6 series

between costules from the costa toward the mar-

gin, or well beyond the costa, in regular or often

irregular lines. Indusium glabrous, rarely some very

small indusia present.

In dense forests, primary forests, somewhat open

or shrubby woods, often on hillsides or in ravines,

100-1500 m, Amazonas south to Cuzco.

Colombia south to Peru.

This species differs from the next, Tectaria dra-

conoptera, in not having the rachis alate to the

basal pair of pinnae, and from Tectaria incisa in

having three to six, usually four or five, series of

sori between adjacent costules.

Tectaria sodiroi has been commonly applied to

this species, and we have annotated sheets with

this name. However, Dr. Robbin Moran has

pointed out to us that the holotype (seen by him

at K) consists of a part of a stem with lomariopsid

vascular pattern and an attached sterile leaf of

Bolbitis nicotianifolia (Sw.) Alston. There is also

a fertile pinna of Tectaria sodiroi. Since the fertile

pinna provides an inadequate holotype, the name

Tectaria sodiroi is rejected.

Amazonas: Prov. Bagua, below Montenegro, 1 8 km E

of Olmos, Hutchison & Wright 3649 (GH, MO, uc, us).

Prov. Bagua, Montenegro-Chiriaco, Sagdstegui 5922

(GH). San Martin: Zepelacio, near Moyobamba, Klug

3666 (GH, MO, us). Loreto: Santa Rosa, below Yurima-

guas, Killip & Smith 28985 (us). Prov. Maynas, Yano-

mono, Vdsquez & Jaramillo 4100 (MO). Huanuco: Prov.

Leoncio Prado, E de Tingo Maria, /. Schunke V. 10169

(F, MO, uc). Prov. Huanuco, Dist. Churubamba, Mexia

8217 (BM, F, GH, MO, uc, us). Tingo Maria (as San Martin),

Allard 21564, 21565 (us). Pasco: Prov. Oxapampa, Rio

Alto Iscozacin, Foster & d'Achille 10062 (F). Cuzco: Prov.

Paucartambo, Atalaya, Foster et al. 3074 (GH). Prov. La

Convention, Rio Mapitunuari, Dudley 11428 (GH, MO).

5. Tectaria draconoptera (D. C. Eaton) Copel.,

Philipp. J. Sci. 2 C: 410. 1907.

26

FIELDIANA: BOTANY

Aspidium draconopterum D. C. Eaton, Mem. Amer.

Acad. Arts n.s. 8: 211. 1860. TYPE: "Turbo in

sinu Uraba, Novae Granadae" (Colombia), A.

Schott 19 (not located).

Stem ascending to erect. Leaves ca. 80 cm to 1 .5

m long. Lamina deeply pinnatisect, the large pin-

na-segments all connected by the broadly to nar-

rowly alate rachis; basal pinna-segments entire or

with 1 large, basal, basiscopic lobe; lateral pinna-

lobes entire; proliferous buds absent. Veins fully

anastomosing, except for included free veinlets.

Leaf-tissue glabrous above and beneath. Sori ex-

indusiate, in 4-7 series between costules from the

costa to near the margin, the series in regular or

mostly irregular lines.

In primary forests and in forested ravines, 350-

700 m, Amazonas south to Cuzco and Madre de

Dios.

Central America; Colombia, Ecuador, and Peru.

Tectaria draconoptera is a distinctive species,

with very many small, exindusiate sori, and with

the basal pinnae connected to the pair above by

the alate rachis.

Amazonas: Valley of Rio Maranon, above Cascadas

de Mayasi, Wurdack 1862 (us). Loreto: Veradero de Ma-

zan, Rio Amazonas to Rio Napo, Croat 19530 (F, MO).

Huanuco: Prov. Leoncio Prado, E of Tingo Maria, /.

Schunke V. 10171 (F, MO, us). Pasco: Prov. Oxapampa,

Paujil, Leon 296 (USM). Junin: 2 km S of Satipo, Solomon

3274 (MO, uc). Ucayali: Vicinity of Aguaytia (as Loreto),

Croat 209 19 (MO). Cuzco: Prov. Paucartambo, Hacienda

Villa Carmen, Vargas 14682 (GH). Prov. Paucartambo,

Cosnipata, Vargas 15779 (GH). Rio Alto, Urubamba,

Bites 1760 (us). Madre de Dios: Parque Nacional del

Manu, Foster & Terborgh 6549 (F).

6. Tectaria plantaginea (Jacq.) Maxon, Contr. U.S.

Natl. Herb. 10: 494. 1908.

Polypodium plantagineum Jacq., Collectanea 2: 104,

t. 3,f. 1. 1788. TYPE: Martinique, probably Jac-

quin (holotype, probably w or BM, not seen). The

illustration is definitive.

Stem short-creeping. Leaves ca. 20-60 cm long.

Lamina entire, long-decurrent at the base; a pro-

liferous bud present at the apex of the lamina.

Veins fully anastomosing, except for included free

veinlets. Leaf-tissue glabrous above and beneath.

Sori exindusiate or rarely indusiate, in 2 series

between the costules, or in 3 or 4 series from the

costa to the margin, the lines regular when in 2

series, often irregular and only near the costa when

in 3 or 4 series.

In ravines and on stream banks in primary wet

forests, 350-750 m, San Martin, and Loreto, south

to Huanuco and Madre de Dios.

Central America and West Indies, south to Peru

and Brazil.

This is a very distinctive species with an entire

lamina that is long-decurrent at its base and with

an apical bud. The species is usually exindusiate,

but in the Guianas T. plantaginea var. macrocarpa

(Fee) Morton is indusiate. Tectaria plantaginea

var. conjluens Morton, with especially the lower

sori confluent and the sporangia on an elongate

receptacle, seems of sporadic occurrence and not

a taxon.

San Martin: Prov. Mariscal Caceres, Palo Blanco,

above Rio Tocache, Plowman & Schunke 7451 (F). Prov.

Mariscal Caceres, Nuevo Progreso, J. Schunke V. 3135

(F, GH, us). Prov. Mariscal Caceres, Santa Rosa de Mi-

shollo, J. Schunke V, 6813 (F, us). Loreto: Balsapuerto,

Killip & Smith 28535 (F, GH, us). Huanuco: Tingo Maria

(as San Martin), Allard 20884, 20887, 20894, 21550 (us).

Madre de Dios: Prov. Manu, Casa Erika, Rio alto Madre

de Dios, Foster & Baldeon 12887 (F).

Comments

Two species have been excluded from the Pe-

ruvian Tectaria flora.

Tectaria heracleifolia (Willd.) Undenv., Bull. Tor-

rey Bot. Club 33: 200. 1906.

Aspidium heracleifolium Willd., Sp. pi. ed. 4, 5: 217.

1810. LECTOTYPE (designated by Underw., Bull.

Torrey Bot. Club 33: 200. 1906): Hispaniola, Plu-

mier, Traite foug. Amer. /. 147, 1705.

This species occurs in southern Florida, the West

Indies, Mexico and Central America, and northern

South America. It evidently does not occur in Peru.

It has a centrally attached, peltate indusium and

in this character differs from other species in

America. While a few Peruvian collections have

some peltate indusia, some to most of the indusia

have a sinus and the specimens are referred to

other species.

Tectaria trifoliata (L.) Cav., Descr. pi. 249. 1802.

Polypodium trifoliatum L., Sp. pi. 1087. 1753. LEC-

TOTYPE (designated by Underw., Bull. Torrey

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

27

FIG. 5. Cyclopeltis semicordata: a, habit; b, pinna base, abaxial side. (From Stolze, Ferns & fern allies of

Guatemala, 1981.)

28

FIELDIANA: BOTANY

Bot. Club 33: 199. 1906): Martinique, Plumier,

Traite foug. Amer. /. 148. 1705.

This species occurs in the West Indies, and in

Surinam, Venezuela, and northern Brazil. The in-

dusia have a sinus and the sori are in several series

between costules. It differs from Tectaria antio-

quoiana, to which materials may key out, in hav-

ing the lamina only lobed, or if pinnate then the

terminal stalked segment is abrupt at the base rath-

er than strongly decurrent.

Two names of uncertain application are based

on Peruvian collections. While they both probably

represent Tectaria antioquoiana, accurate identi-

fication depends on a study of authentic materials:

Tectaria kunzei (Hieron.) C. Chr., Index fil.

suppl. 3: 181. 1934. is based on Aspidium kunzei

Hieron., Hedwigia 46: 353, 15 July, 1907., which

in turn is based on Aspidium macrophyllum var.

decurrens Kunze (not Tectaria decurrens (Presl)

Copel., 6 Nov. 1907). Aspidium macrophyllum var.

decurrens Kunze, Linnaea 9: 89. 1834 is based on

two collections of Poeppig: Diar. 1138 from Pam-

payaco, and Diar. 2288 from Prov. Maynas. The

latter is evidently the type collection, because Diar.

1138 was segregated as a distinct species by Presl

in 1851, leaving Diar. 2288 to represent var. de-

currens.

Tectaria poeppigii (Presl) C. Chr., Index fil. suppl.

3: 183. 1934, is based on Aspidium poeppigii Presl,

Epim. Bot. 62. 1851. Presl cites only Pampayaco,

Poeppig, which, from the locality, must be Poeppig

1138.

V. Cyclopeltis

Cyclopeltis John Sm., Companion, Bot. Mag. 72,

III, 2: 36 (first of two pages). 1846. TYPE:

Cyclopeltis semicordata (Sw.) John Sm. (Poly-

podium semicordatum Sw.). Figure 5a-b.

Stem decumbent, rather stout, short-creeping,

bearing scales. Leaves ca. 20 cm to 1.5 m long,

petiole continuous. Lamina 1 -pinnate, the pinnae

entire or nearly so, nearly glabrous to somewhat

scaly and pubescent, monomorphic, pinnae artic-

ulate, adaxial side of the costa raised. Veins free

or slightly anastomosing. Sori roundish, on the

veins in 1-3 series on each side of the costa, not

paraphysate, covered by fugacious to persistent,

peltate indusia. Spores spheroidal, monolete, with

prominent folds, more or less spinulose.

Cyclopeltis is a tropical genus of four to six spe-

cies, with one of them in American and in Peru.

1. Cyclopeltis semicordata (Sw.) John Sm., Com-

panion Bot. Mag. vol. 72, III, 2: 36 (first of

two pages). 1846. Figure 5a-b.

Polypodiwn semicordatum Sw., Prodr. 132. 1788.

TYPE: Jamaica, Swartz (not seen); "Jamaica, ex

Vahl" B!, Herb. Willd. 19742, photo GH is doubt-

less authentic.

Stem short-creeping, the apex densely scaly with

soft, long, linear, brown scales. Leaves mostly 50

cm to 1 m long, the petiole more or less persistently

scaly especially toward the base. Lamina 1 -pin-

nate, usually with a conform apical segment, pin-

nae entire or rarely slightly lobed, sessile or nearly

so, auriculate at the base of the basiscopic side,

the large auricle overlapping the rachis, articulate

to the rachis and deciduous with age. Veins free

or slightly anastomosing. Sori borne in 2 series

(rarely 1 or 3) on each side of the costa, indusium

peltate.

Primary forests, hillside forests, thickets, partly

disturbed forests, and river banks, 100-750 m,

mostly 400 m or lower, Amazonas to Madre de

Dios.

Southern Mexico and the Greater Antilles, south

to Bolivia and the Amazon Basin of Brazil.

This is a very distinctive species, with a 1 -pin-

nate lamina, articulate pinnae with a large basal,

basiscopic auricle, and peltate indusia. The veins

are usually free but sometimes they are casually

anastomosing.

Amazonas: Prov. Bagua, 3-5 km above mouth of Rio

Santiago, Wurdack 2180 (F, GH, us). E of Huampami,

Rio Cenepa, Berlin 301 (MO). San Martin: Prov. Maris-

cal Caceres, Dist. Campanilla, Mashuyaca, J. Schunke

V. 4218 (F, GH, MO, us). Juan Jui, Klug4166 (MO). Loreto:

Gamitanacocha, Rio Mazan, C. Schunke 194 (F, GH, uc,

us). Mouth of Rio Santiago, above Pongo de Manseriche,

Mexia 6112 (F, GH, MO, uc, us). Huanuco: Prov. Pachi-

tea, Bosque Nacional de Iparia, J. Schunke V. 1247 (F,

GH, us). Prov. Huanuco, Tulumayo, cerca a Tingo Maria,

Ferreyra 2174 (GH, USM). Prov. Tingo Maria, Aldave &

Fernandez 5603 (HUT). Pasco: Quillasu, Soukup 3328

(F, GH). Junin: Prov. Tarma, La Merced, Cerrate 2828

(F, GH). Ucayali: Prov. Coronel Portillo (as Loreto), Bos-

que Nacional de Iparia, J. Schunke V. 2673 (F, GH, us).

Prov. Coronel Portillo, Bosque von Humboldt, Young

& Salazar 1017 (MO). Ayacucho: Near Kimpitiriki, Rio

Apurimac, Killip & Smith 22881 (GH, us). Madre de

Dios: Parque Nacional de Manu, Cocha Casha, Foster

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

29

2 cm

2 cm

FIG. 6. Rumohra adiantiformis: a, habit; b, portion of rachis with bases of costae, adaxial side; c, pinnule,

abaxial side. (From Anderson et al. 35820, Brazil, F.)

30

FIELDIANA: BOTANY

& Terborgh 6636, 6665 (F). Prov. Tambopata, Rio Ma-

dre de Dios, Alfaro 1851 (MO).

VI. Rumohra

Rumohra Raddi, Opusc. Sci. 3: 290. 1819. TYPE:

Rumohra aspidioides Raddi = Rumohra

adiantiformis (Forster) Ching. Figure 6a-c.

Stem rather stout, long-creeping, bearing scales.

Leaves ca. 10 cm to 1 m long, petiole continuous.

Lamina 2-pinnate-pinnatifid to 4-pinnate, gla-

brous or somewhat scaly, monomorphic, adaxial

side of the rachis with a central ridge and glabrous

grooves on each side, pinnae not articulate. Pinna-

rachis sulcate on the adaxial side, the groove con-

tinuous with that of the rachis. Veins free. Sori

roundish, on the veins or at the vein tips, not

paraphysate, covered by peltate indusia that, with

the slender central stalk, is often fugacious. Spores

rather ellipsoidal, monolete, with short to long

ridges, or saccate.

Rumohra is a small genus consisting of the wide-

ly spread R. adiantiformis and a few other species.

In America there is the former species and R. ber-

teriana (Colla) Duek & Rodriguez of the Juan Fer-

nandez Islands.

1 . Rumohra adiantiformis (Forster) Ching, Sinen-

sia 5: 70. 1934. Figure 6a-c.

Polypodium adiantiforme Forster, Prodr. 82. 1786.

TYPE: "Ins. austral." G. Forster (holotype or iso-

type, BM, GOET).

Stem long-creeping, the leaves spaced, densely

covered with brown, broad, thin scales. Leaves to

ca. 1 m long, the petiole usually scaly, especially

toward the base. Lamina 2-pinnate-pinnatifid to

4-pinnate, glabrous or somewhat scaly, coria-

ceous, the basal pinnae the largest. Sori roundish,

indusium peltate, often fugacious.

On rocks near river, ca. 2400 m, Amazonas.

Bermuda, south to southern Chile and the Falk-

land Islands; Old World.

Rumohra adiantiformis is uncommon in the

Andes. It is distinguished from species of other

genera by the peltate indusia, stalked pinnae, and

the entire to obtusely dentate ultimate segments.

If the indusia are absent, the adaxial side of the

rachis, with a central ridge and a groove on each

side, is distinctive. The adaxial side of the penul-

timate segments has the costa prominently raised,

rather than flattened or grooved as in Dryopteris.

Amazonas: Prov. Chachapoyas, Rio Ventilla, 1-2 km

W of Molinopampa, Wurdack 1472 (GH, us).

VII. Lastreopsis

Lastreopsis Ching, Bull. Fan Mem. Instil. Biol.,

Bot. 8:157.1938. TYPE: Lastreopsis recedens

(Moore) Ching (Lastrea recedens Moore) =

Lastreopsis tenera (R. Br.) Tindale. Figure

7a-b.

Stem erect to short- or long-creeping, usually

moderately stout, bearing scales. Leaves ca. 50 cm

to 2 m or rarely 3 m long, petiole continuous.

Lamina 2-pinnate to 5-pinnate-pinnatifid, pubes-

cent, glandular-pubescent, glandular, and often

scaly, monomorphic, adaxial side of the rachis with

a ridge on each side of a pubescent groove, pinnae

not articulate, the pinna-rachis sulcate adaxially,

the groove continuous with that of the rachis. Veins

free. Sori roundish, borne on the veins or on the

vein tips, not paraphysate, covered by reniform,

rarely subpeltate indusia or exindusiate. Spores

rather ellipsoidal, monolete, strongly ridged, or

saccate.

Lastreopsis is a genus of about 35 species, with

five in America and two in Peru. It is pantropical

and south temperate in its distribution. It is readily

separated from Dryopteris, Stigmatopteris, and

Cyclodium by the raised costa on the adaxial side.

Those other genera, which may have some species

that resemble some of Lastreopsis, all have the

costa grooved on the adaxial side.

Reference

TINDALE, M. D. 1 965. A monograph of the genus

Lastreopsis Ching. Contr. N.S.W. Natl. Herb.,

3: 249-339.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

31

FIG. 7. Lastreopsis effusa ssp. divergens: a, portion of lamina. Lastreopsis killipii: b, portion of rachis and pinna

base, adaxial side; c, tertiary segment, abaxial side, (a from R. Williams 1273, Bolivia, GH; b, c from Little 9401,

Colombia, F.)

FIELDIANA: BOTANY

Key to Species of Lastreopsis

a. Lamina catadromic, at least above the basal pinnae; sori exindusiate; rachis glabrous or nearly so

1 . L. of f usa

a. Lamina anadromic throughout; sori indusiate; rachis prominently scaly 2. L. killipii

1 . Lastreopsis effusa (Sw.) Tindale, Viet. Nat. 73:

184. 1957. Figure 7a.

Polypodium effusum Sw., Prodr. 134. 1788. LEC-

TOTYPE (designated by Tindale, 1957): Jamai-

ca, Swartz(ho\oiype,Bl,Herb. Willd. 1 97 24; pho-

to GH; isotypes, c, UPS).

Stem decumbent, short-creeping, to nearly erect,

bearing brown, lanceolate scales. Leaves ca. 1-2.5

m long; the petiole glabrous, or slightly scaly, es-

pecially toward the base. Lamina 3 -pinnate to

4-pinnate-pinnatifid, catadromic, at least above

the basal pinnae, the apex gradually reduced, usu-

ally somewhat pubescent and/or glandular, es-

pecially on the adaxial side of the axes; usually

bearing a proliferous bud toward the apex of the

rachis; pinnae stalked, the basal the largest. Veins

free. Sori borne on the veins, exindusiate.

Dense forests, disturbed forests, and on moun-

tain slopes, 135-1 800 m, Cajamarca to Cuzco and

Madre de Dios.

Tropical America.

Of the four more or less sympatric subspecies

recognized by Tindale (1957), only ssp. divergens

(Willd.) Tindale is in Peru. Lastreopsis effusa is

one the few species of the genus that has the lamina

usually with a scaly bud toward the apex of the

rachis.

Dryopteris killipii Maxon, Amer. Fern J. 18: 4. 1928.

TYPE: Panama, Chiriqui, W of El Boquete, Killip

5360 (holotype, us; isotype, GH!).

Stem short-creeping, decumbent, bearing brown,

narrow scales. Leaves ca. 2-5 m long; the petiole

rather persistently scaly, with mostly brownish,

narrow scales. Lamina 4-pinnate to nearly 5-pin-

nate, anadromic, the apex gradually reduced, var-

iously scaly, pubescent, and/or glandular, especial-

ly on the axes; lacking a proliferous bud; pinnae

stalked, the basal the largest. Veins free. Sori borne

on the veins, covered by a reniform, persistent

indusium.

Dense forests and cloud forests, 2080-2700 m,

Cuzco.

Costa Rica, Panama, Colombia, and Peru.

The leaves are up to 3.5 or even 5 m long.

Cuzco: Prov. La Convencion, altura de Pintobamba,

Vargas 3552 (us). Prov. La Convencion, Cordillera Vil-

cabamba, Dudley 11243 (GH, us).

Comments

Except for the widespread Lastreopsis effusa,

other species of South America are not commonly

collected, but two of them may be found in Peru.

Cajamarca: Prov. Hualgayoc, Soukup 3812 (F, us).

Prov. Santa Cruz, Sagdstegui et al. 12391 (F, GH, HUT).

Loreto: Prov. Maynas, Rio Nanay, across from Bellavis-

ta, McDaniel & Rimachi 18794 (GH, MO). Santa Rosa,

below Yurimaguas, Killip & Smith 28733, 28853 (us).

Huanuco: Pampayacu, Rio Chinchao, Macbride 5036 (F,

us). Pasco: Prov. Oxapampa, Valle del Palcazu, Leon

679 (F). Junin: Above San Ramon, C. Schunke A226

(us). Chanchamayo Valley, C. Schunke 50, 90 (F, us).

Prov. Satipo, Leon 186 (MO, USM). Cuzco: Prov. La Con-

vencion, Rio Apurimac, Davis et al. 1310 (F). Madre de

Dios: Prov. Manu, Cocha Cashu uplands, Nunez 5815

(MO).

2. Lastreopsis killipii (Maxon) Tindale, Viet. Nat.

73: 185. 1957. Figure 7b-c.

Lastreopsis amplissima (Presl) Tindale, Viet. Nat.

73: 185. 1957.

Polystichum amplissimum Presl, Epim. Bot. 58. 1851.

SYNTYPES: Brazil, Sellow; Brazil, Serra d'Es-

trella, Beyrich (neither located); ISOSYNTYPE:

Sellow (B).

This species has a wholly anadromic lamina with

indusiate sori and is closely related to L. killipii.

It has the ultimate segments mostly sharply point-

ed to mucronate apically, while L. killipii has them

mostly obtuse to subacute. This species ranges from

Venezuela south to southeastern Brazil and Par-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

33

FIG. 8. Dryopteris paleacea: a, habit; b, pinna segment, abaxial side; Dryopteris denticulata: c, lamina, (a, b

from Killip & Smith 18824, Colombia, F, c adapted from Stolze. Ferns & fern allies of Guatemala, 1981.)

34

FIELDIANA: BOTANY

aguay; it is also in Bolivia and may be expected

to grow in southern Peru.

Lastreopsis exculta (Mett.) Tindale, Viet. Nat. 73:

185. 1957.

Aspidium excultum Mett., Uber einige Farngatt.

IV. Phegopt. Aspid. 69, t.!7,f. 9. 1859. (Abh.

SenckenbergNaturf. Ges. 2: 353. 1858. TYPE:

Venezuela, Caracas, Moritz 433 (holotype, B?;

isotype, GH!).

This species has a catadromic lamina, indusia,

and a short-pubescent costa adaxially. It grows in

Mexico south to northern South America and Ec-

uador and probably also grows in northern Peru.

VIII. Dryopteris

Dryopteris Adans., Fam. plantes 2: 20. 551. 1763,

nom. conserv. TYPE: Dryopteris filix- mas (L.)

Schott (Polypodium filix- mas L.). Figure 8.

Arachniodes Blume, Enum. pi. Javae 24 1 . 1 828. TYPE:

Arachniodes aspidioides Blume = IDryopteris ar-

istata (Forster) Kuntze.

Lastrea subsection Ipolystichopsis John Sm., Hist. fil.

217. 1875. LECTOTYPE (designated by C. Chr.,

Index fil. xxi. 1906): Lastrea pubescens(Sv/.) Presl

(Polypodium pubescens (L.) = Dryopteris pubes-

cens (L.) Kuntze.

Dryopteris subgenus Polystichopsis (John Sm.) Ch. Chr.,

Danske Vidensk. Selsk. Skrift. Nat. Math. VII,

6(1): 101. 1920.

Polystichopsis (John Sm.) Holtt., Flora Malaya 2 (Ferns

Malaya): 484. 1955.

Byrsopteris Morton, Amer. Fern J. 50: 149. 1960.

TYPE: Byrsopteris aristata (Forster) Morton

(Polypodium aristatum Forster) = Dryopteris ar-

istata (Forster) Kuntze.

Stem decumbent, small or stout, or erect to ca.

30 cm tall, rarely long-creeping, bearing scales.

Leaves ca. 1 0 cm to 2 m long, petiole continuous.

Lamina 1 -pinnate, usually 2-pinnate-pinnatifid, to

rarely 6-pinnate, glabrous, glandular, pubescent,

or scaly, monomorphic or slightly dimorphic, ad-

axial side of the costa sulcate to nearly flat. Veins

free. Sori roundish, borne on the veins or on the

vein tips, not paraphysate, covered by usually ren-

iform, rarely subpeltate, indusia, or exindusiate.

Spores rather ellipsoidal, monolete, prominently

ridged, saccate, or cristate, often spinulose.

Dryopteris is a large genus of about 1 50 species.

It is worldwide in its distribution. The center of

species diversity is in China and adjacent regions.

There are about 25 species in America, and five

in Peru.

Dryopteris sensu stricto is closely allied to the

sometimes recognized genera Arachniodes and

Polystichopsis. Some species, usually placed in one

of the three groups, are divergent in their char-

acters toward another group. The whole assem-

blage is closely allied and relations of a number

of species are unclear. For these reasons all species

are treated here in the single genus Dryopteris.

Two subgenera may be recognized: Dryopteris

subgenus Dryopteris has the stem with internal

glands, the lamina often catadromic, and the basal

pinnae not, or not much, enlarged. Peruvian spe-

cies are Dryopteris patula, D. paleacea, and D.

saffordii. Dryopteris subgenus Polystichopsis (in-

cluding Arachniodes) has the stem lacking internal

glands, the lamina anadromic, and the basal pin-

nae enlarged. Peruvian species are Dryopteris den-

ticulata and D. ochropteroides.

Reference

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

teris, pp. 496-509, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Species of Dryopteris

a. Rachis definitely scaly with long, narrow, dark brown or darker scales, these grading into fibrils;

pinnae sessile or nearly so, deeply pinnatifid to usually pinnatisect; secondary segments approximate,

nearly entire 1 • D- paleacea

a. Rachis with a few scattered scales, or none, long-pubescent, minutely glandular, or glabrous . . . . b

b. Lamina minutely glandular, especially on the rachis, at the base of the pinnae, on the pinna-

rachis, costa, and usually elsewhere c

c. Pinnae stalked, or rarely short-stalked, the basal ones not or hardly reduced, usually the largest

on the lamina; basal secondary segments on the central pinnae narrowed at the base, or stalked,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

35

usually connected to the next pair of segments by a narrow wing of the pinna-rachis

2. D. patula

c. Pinnae sessile or nearly so, the basal ones reduced, usually much reduced, shorter than the

central pinnae; basal secondary segments on the central pinnae usually joined to the next pair

of segments by their decurrent base 3. D. saffordii

b. Lamina eglandular, glabrous, or with a few fibrils, or long-pubescent, especially on the axes d

d. Rachis and other axes glabrous or with a few fibrils 4. D. denticulata

d. Rachis and other axes long-pubescent, especially on the abaxial side, or partly so

5. D. ochropteroides

1. Dryopteris paleacea (Sw.) Hand.-Mazz., Ver-

handl. Zool.-Bot. Gesells. Wein 58: (100).

1908. Figure 8a-b.

Aspidium paleaceum Sw., Syn. fil. 52. 1806. TYPE:

Peru, Lagasca misit (holotype, s, Herb. Swartz;

photos, GH, us).

Aspidium parallelogramma Kunze, Linnaea 13: 146.

1839. SYNTYPES: Mexico, Hegewisch, Mexico,

Karwinsky (not located).

Dryopteris parallelogramma (Kunze) Alston, Amer.

Fern J. 47: 92. 1957.

Stem erect, stout. Leaves borne in a crown, ca.

30 cm to 1.5m long. Lamina 1 -pinnate-pinnatifid

to usually 1 -pinnate-pinnatisect, more or less scaly,

especially on the rachis, which is rather densely

scaly, with mostly long, narrow, reddish brown to

dark brown or partly blackish scales abaxially and

often fibrillose adaxially. Pinnae sessile or nearly

so, the basal not much shorter than the central

ones; the secondary segments approximate, nearly

entire to somewhat dentate.

In dense forests or thickets, disturbed forests, in

moist ravines, on wet, shaded road banks, and at

the base of rocks, 2400-4000 m, Cajamarca to

Puno.

Mexico and Greater Antilles, south to Argentina

and southern Brazil; primarily montane.

The correct taxonomy of this species is not clear.

There may be one, essentially pantropical, species,

or there may be two, one American and the other

Old World. The American element is here treated

as a separate species, pending a definitive reso-

lution of this problem.

The correct nomenclature depends upon the in-

terpretation of the publication of Aspidium pale-

aceum by Don (Prodr. fl. nepal. 4. 1825). If this

is a new species (Alston, Amer. Fern J. 47: 91-92.

1957; A. R. Smith and Fraser-Jenkins, Taxon 31:

326-329. 1982) then the correct name for the col-

lective species is Dryopteris wallichiana (Sprengel)

Hylander. This is also the correct name for the

Old World element, as a species, while the Amer-

ican element is Dryopteris parallelogramma

(Kunze) Alston.

If Don's name is a later use of Aspidium pale-

aceum Sw. (the conclusion adopted here; also by

Fraser-Jenkins, Taxon 29: 610-611. 1980) then

the correct name for the collective species is

Dryopteris paleacea (Sw.) Hand.-Mazz. This is also

the correct name for the American element, as a

species, while the Old World element is Dryopteris

wallichiana (Sprengel) Hylander.

Cajamarca: Prov. San Miguel, Vista Alegre, Llatas Q.

1574 (F). Prov. Celendin, entre Celendin y Cajamarca,

Lopez et al. 4462 (GH, HUT). Amazonas: Prov. Chacha-

poyas, Cerros Calla Calla, Hutchison & Wright 5753 (F,

GH, MO, uc, us). Prov. Chachapoyas, Puma-urcu, SE of

Chachapoyas, Wurdack 559 (F, GH, uc, us). La Libertad:

Prov. Pataz, above Pampa Rosas, Young 3 105 (F). Prov.

Santiago de Chuco, Dist. Cachicadan, Sounders 884 (F,

GH). Between Huamachuco and Cajabamba, Correll &

Smith P925 (GH). San Martin: Dist. Huallaga, valley of

Rio Apisoncho, Hamilton & Holligan 925 (uc). Prov.

Mariscal Caceres, Rio Abiseo National Park, Young &

Leon 4982 (F). Huanuco: Mito, Macbride & Featherstone

1512 (F, us). Prov. Huanuco, 32 km from Huanuco on

Huanuco-La Union Road, Smith et al. 2187 (MO). Junin:

Carpapata, above Huacapistana, Killip & Smith 24465

(F, us). Huancavelica: Entre Huschocolpa y Surcubamba,

Tovar 4237 (GH). Ayacucho: Prov. La Mar, Dudley 9051

(GH). Prov. La Mar, between El Tambo and Ayna, Plow-

man & Davis 4684 (GH). Apurimac: Dist. Abancay,

Sounders 761 (GH). Cuzco: Machu Picchu, Peyton & Pey-

ton 759 (GH, MO). Prov. La Convention, Dist. Vilcabam-

ba, Davis et al. 7279 (F, GH). Puno: Prov. Carabaya, entre

Ayapata y Kahualluyac, Vargas 10735 (GH).

2. Dryopteris patula (Sw.) Underw., Our native

ferns, ed. 4, 117. 1893.

Aspidium patulum Sw., Kongl. Vetensk. Acad. Handl.

1817: 64. TYPE: Brazil, Minas Gerais, Freyreiss

(holotype, s, seen by C. Chr. Monogr. genus

Dryopteris 1: 19. 1913).

36

FIELDIANA: BOTANY

Stem erect or nearly so, rather stout. Leaves

borne in a cluster, ca. 25 cm to 1 m long. Lamina

2-pinnate to 3-pinnate-pinnatifid, minutely glan-

dular, especially on the rachis, at the base of the

pinnae, on the pinna-rachis, and often elsewhere,

the rachis sometimes with scattered light brown

scales. Pinnae stalked, rarely short-stalked to near-

ly sessile, the basal not or hardly reduced, usually

the largest, secondary segments usually spaced,

usually shallowly to sometimes deeply lobed or

more complex.

In woods, among shaded rocks, and on rocky

slopes, 2000-2700 m, Huanuco, Junin, and Cuz-

co.

Mexico and the Greater Antilles, south to Ar-

gentina and southern Brazil; primarily montane.

This species has a more complex lamina than

Dryopteris saffordii. The basal segments of the cen-

tral pinnae are narrowed at the base, or stalked

and connected to the next pair of segments by a

narrow wing along the pinna-rachis.

Huanuco: Chinchao to Puente Durand, Coronado 82

(us). Mima, Bryan 419 (F). Yanano, Macbride 3821 (F,

us). Junin: Carpapata, Kunkel 603 (GH). Cuzco: Machu

Picchu, Leon 453 (F, GH, HUT), Coronado 95 (uc, us).

Prov. Paucartambo, Pillawata, Vargas 16688 (GH).

3. Dryopteris saffordii C. Chr., Amer. Fern J. 1:

94. 1911. TYPE: Peru, mountains back of

Lima, Arroya Railroad, W. E. Safford 994

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

Stem usually small, decumbent to erect. Leaves

borne in a cluster, ca. 15-60 cm long. Lamina

1 -pinnate-pinnatifid to rarely 2-pinnate-pinnati-

fid, minutely glandular, especially on the rachis,

at the base of the pinnae, on the pinna-rachis, the

costa, and often elsewhere, the rachis sometimes

with scattered light brown scales. Pinnae sessile or

nearly so, the basal ones much reduced, secondary

segments approximate or spaced, nearly entire to

lobed or rarely pinnatifid.

Brushy, rocky slopes, open rocky places, at the

base of rocks, 400-4000 m (400-600 m on Loma

Lachay, otherwise at 2200-4000 m), Cajamarca

to Puno.

Peru and Bolivia.

Dryopteris saffordii is very close to D. patula

and might better be treated as a subspecies or va-

riety of it. Soukup 2961, Huancayo, and Fiebrig

3133, in part, Bolivia (GH) have large leaves with

the lamina 2-pinnate-pinnatifid and nearly sessile

pinnae, the basal definitely reduced. These collec-

tions are rather intermediate between the two

species. Apparently intermediate specimens, how-

ever, are few and its seems best to maintain two

species at the present time.

Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui

2938 (F, GH). 29 km from Cajamarca on road to Chilete,

Correll & Smith P849 (GH), P852 (GH, us). Prov. Caja-

marca, cumbre El Galivan, Ferreyra 3276 (USM). La Li-

bertad: Prov. Otuzco, Huaranchal, Lopez et al. 2663

(GH). Between Huamachuco and Cajabamba, Correll &

Smith P922 (GH). Prov. Sanchez, Carrion, Sagdstegui

9445 (HUT). Ancash: Prov. Bolognesi, E of Huasta, Cer-

rate 2460 (GH, USM). Prov. Bolognesi, arriba de Chiqui-

an, Cerrate 1550 (GH). Lima: Prov. Chancay, Loma de

Lachay (as Luchay), Tryon & Tryon 5417 (F, GH, us),

Coronado 22 (GH, uc, us). Prov. Huarochiri, Dist. Surco,

Saunders 657 (F, GH, us). Junin: Huancayo, Soukup 2961

(F, GH, MO, us). Cuzco: Prov. Urubamba, Chupani, Var-

gas 11125 (GH). Puno: Near Puno, Soukup 92 (F).

4. Dryopteris denticulata (Sw.) Kuntze, Revis. gen.

pi. 2: 812. 1891. Figure 8c.

Polypodium denticulatum Sw., Prodr. 134. 1788.

TYPE: Jamaica, Swartz (holotype, s, seen by C.

Chr., Monogr. genus Dryopteris 2: 113. 1920).

Arachniodes denticulata (Sw.) Ching, Acta Bot. Sinica

10: 260. 1962.

Stem decumbent to erect, small to rather stout.

Leaves borne in a cluster, ca. 25 cm to 1 m long.

Lamina 3 -pinnate-pinnatifid to 4-pinnate-pinnati-

fid, or 5 -pinnate, glabrous or glabrate. Pinnae

stalked, the basal ones enlarged, the longest, sec-

ondary segments 1 -pinnate or more complex.

Moist forests, and roadside banks, 2000-3000

m, Cajamarca, south to Puno.

Southern Mexico and the Greater Antilles, south

to northern South America and Bolivia; south-

eastern Brazil.

The lamina is very strongly and wholly anad-

romic, and the ultimate segments are mostly

sharply dentate, especially the apical ones.

Cajamarca: Prov. Cutervo, Llipa, Mostacero et al. 1749

(F, HUT). Amazonas: Prov. Chachapoyas, 5 km below

Chachapoyas, Wurdack 778 (F, GH), 779 (F, USM). San

Martin: Leon 2144, 2162 (F). Prov. Bongara, El Ingenio-

Pomacochas, Sagdstegui 5968 (GH). Huanuco: Cushi,

Bryan 685 (F). Pasco: Prov. Oxapampa, Rio San Alberto

valley, E of Oxapampa, Smith & Pretel 7598 (F). Puno:

Sandia, Vargas 11859 (on).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

37

5. Dryopteris ochropteroides (Baker) C. Chr., In-

dex fil. 280. 1905.

Nephrodium ochropteroides Baker, Ann. Bot. (Lon-

don) 5: 325. 1891. TYPE: Jamaica, Fox's Gap,

Hart (holotype, K; isotype, u).

Polystichopsis ochropteroides (Baker) Morton, Amer.

Fern J. 50: 155. 1960.

Arachniodes ochropteroides (Baker) Lell., Amer. Fern

J. 77: 101. 1988.

Stem rather slender and long-creeping in small

plants, becoming rather stout and decumbent.

Leaves clustered to spaced, ca. 40 cm to 1 m long.

Lamina 3-pinnate to 4-pinnate-pinnatifid, with the

axes long-pubescent, especially on the abaxial side,

usually densely so but sometimes the pubescence

scattered. Pinnae stalked, the basal ones enlarged

and the longest, secondary segments 1 -pinnate or

more complex.

Dense cloud forests, ca. 1290 m, Huanuco; a

single collection from Peru.

Jamaica, Surinam, Venezuela, and Peru; range

uncertain.

The lamina is definitely anadromic and the ul-

timate segment are subacute to acute.

This species is separated from the similar

Dryopteris macrostegia (Hooker) O. Kuntze, of

Venezuela, Colombia, and northern Brazil, by the

axes of the lamina that are long-pubescent. The

latter species has the axes of the lamina glabrous.

The pubescence of Dryopteris ochropteroides may

be scattered and sparse. Both species are seldom

collected and additional material is needed in or-

der to understand their relation.

long, petiole continuous. Lamina 1 -pinnate, or

rarely entire, or 2-pinnate, glabrate to more or less

scaly, monomorphic, adaxial side of the costa sul-

cate, pinnae not articulate, more or less inequi-

lateral at the base. Veins free to regularly anas-

tomosing and with or without included free

veinlets. Sori roundish, borne on the veins or at

the vein tips, in 2 to several (rarely 1) series on

each side of the costa, not paraphysate, covered

by persistent to fugacious, peltate indusia or rarely

exindusiate. Spores ellipsoidal to spheroidal,

monolete, ridged, rugose or saccate.

Cyrtomium is a genus of about 25 species, most

of them in eastern Asia. In America the nine or

ten species are concentrated in Mexico and Central

America. A single native species, C. dubium, oc-

curs in Peru.

Although the predominantly free-veined Amer-

ican species are sometimes recognized as a distinct

genus Phanerophlebia, the two groups are not

wholly distinctive.

References

CHING, R. C. 1936. On the genus Cyrtomium

Presl. Bull. Chinese Bot. Soc., 2: 87-106.

CHRISTENSEN, C. 1930. The genus Cyrtomium.

Amer. Fern J., 20: 41-52.

MAXON, W. R. 1912. Notes on the North Amer-

ican species of Phanerophlebia. Bull. Torrey Bot.

Club, 39: 23-28.

UNDERWOOD, L. M. 1899. American ferns, II.

The genus Phanerophlebia. Bull. Torrey Bot.

Club, 26: 205-216.

Huanuco: Rio Llullapichis watershed, ascent of Cerros

del Sira, Dudley 13062 (GH, us).

1. Cyrtomium dubium (Karsten) R. & A. Tryon,

Rhodora 83: 134. 1981. Figure 9a-b.

IX. Cyrtomium

Cyrtomium Presl, Tent, pterid. 86. 1836. TYPE:

Cyrtomium falcatum (L. f.) Presl (Polypodium

falcatum L. f). Figure 9.

Phanerophlebia Presl, Tent, pterid. 84. 1836, cited as

a synonym of Cyrtomium by Moore, Index fil.

Ixxxii. 1857. TYPE: Phanerophlebia nobilis

(Schlect. & Cham.) Presl (Aspidium nobile Schlect.

& Cham.) = Cyrtomium nobile (Schlect. & Cham.)

Moore.

Stem decumbent, short, rather stout, to nearly

erect, bearing scales. Leaves ca. 15 cm to 1.5 m

Phegopteris dubia Karsten, Fl. columb. 1: 169, t. 84.

1861. TYPE: Colombia, Bogota, Tequendama,

Karsten (not located).

Phegopteris dictyophylla Kuhn, Linnaea 36: 106. 1869.

TYPE: Ecuador, Spruce 5263 (holotype, B?; iso-

type, as Spruce 5265, p, Herb. Bonaparte. Not

Spruce 5263, P, Herb. Bonaparte which is Polys-

tichum Bonapartii Rosenst.; holotype, p, Herb.

Bonaparte; isotype, GH!).

Polystichum dubium (Karsten) Diels, Nat. Pflanzen-

fam. 1(4): 194. 1899.

Stem decumbent, very short-creeping, to erect,

bearing dark brown, lustrous scales. Leaves ca. 30

cm to 1 m long, the petiole usually sparingly scaly.

Lamina usually 1 -pinnate, to rarely partly or fully

38

FIELDIANA: BOTANY

FIG. 9. Cyrtomium dubium: a, habit; b, portion of pinna, abaxial side, (a from Peyton & Peyton 1275, GH; b

from Killip & Smith 20366. GH.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

39

2-pinnate, the apex gradually reduced, pinnae usu-

ally entire or nearly so, herbaceous, usually rather

sharply dentate, often with a usually prominent

basal auricle on the acroscopic side, veins mostly

free to freely anastomosing, with or without in-

cluded free veinlets. Sori in usually 3 (2 to 4) series

on each side of the costa or costule, exindusiate.

Dense hillside forests, scrubby forests, some-

times among rocks, or rarely a low epiphyte, 1450-

3400 m, Cajamarca to Cuzco.

Costa Rica south to Bolivia.

Cyrtomium dubium is a morphologically rather

isolated species that is not clearly placed in either

Polystichum or Cyrtomium. It has, however, the

usually anastomosing veins and multiseriate sori

of the latter genus and seems best placed there.

It is a variable species in lamina architecture

and venation. Spruce 5263 is evidently a mixed

collection of 2 species and Spruce 5265 is a mix-

ture of 1 -pinnate and 2-pinnate Cyrtomium du-

bium.

The pinnae vary from usually entire with an

obtuse to acuminate apex, to more or less serrately

lobed, or the pinnae may be basally mostly

1 -pinnate or fully 1 -pinnate, as in Barbour 2875.

The venation varies from fully anastomosing, ex-

cept at the margins, to mostly free. Although some

of this variation may be related to hybridization,

especially with species of Polystichum, spores ap-

pear to be normal and the lamina architecture is

regular. The 1 -pinnate and 2-pinnate lamina forms

are therefore considered to be within the normal

range of variation of the species, as in various

species of Lindsaea.

Cajamarca: Prov. Cutervo, San Andres, Lopez & Sa-

gdstegui 5401 (GH, HUT). Amazonas: E of La Peca, Bar-

bour 2875 (MO). San Martin: Between Mirador and La

Playa, Young & Leon 4927 (F, USM). Huanuco: Mima,

Bryan 554 (F, us). Huacachi, near Mima, Macbride4l26

(F, us). Junin: Carpapata, above Huacapistana, Killip &

Smith 24464 (BM, F, us). Ayacucho: Between Huanta and

Rio Apurimac, Killip & Smith 22362 (us). Cuzco: Uru-

bamba, Machu Picchu, Peyton & Peyton 1328 (GH, MO),

1504 (MO). Quillabamba, Santa Teresa, Peyton & Peyton

1275 (GH, MO). Prov. Paucartambo, Pillahuata, Aldave

5021 (HUT).

Comments

Cyrtomium falcatum (L. f.) Presl, Tent, pterid. 86.

1836.

Polypodium falcatum L. f., Suppl. pi. syst. veg. ed. 13,

446. 1782. TYPE: Japan, Nagasaki, Thunberg

(holotype, LINN?; isotype, UPS).

This is a commonly cultivated species of eastern

Asia. It has been collected on the sea-cliffs near

Lima and it may be adventive elsewhere in Peru.

It diners from Cyrtomium dubium in being in-

dusiate, and having very coriaceous pinnae with

thickened margins.

Lima: Prov. Lima, Magdalena del Mar, cerca a Lima,

Julio 14, 1942, Zuniga (GH).

X. Didymochlaena

Didymochlaena Desv., Ges. Naturf. Freunde Ber-

lin Mag. Neuesten Entdeck. Gesammten Na-

turk. 5: 303. 1811. TYPE: Didymochlaena

sinuosa Desv. = Didymochlaena truncatula

(Sw.) John Sm. Figure lOa-c.

Stem decumbent or usually erect, rather stout,

bearing scales. Leaves ca. 75 cm to 2 m or rarely

3 m long, petiole continuous. Lamina 2-pinnate,

nearly glabrous to scaly and pubescent, mono-

morphic, pinnules dimidiate. Veins free. Sori

elongate, borne on the veins, not paraphysate, cov-

ered by elongate indusia on each side of a vein.

Spores ellipsoidal to spheroidal, monolete, with

short ridges or saccate, spinulose.

Didymochlaena is a monotypic genus of pan-

tropical distribution. The single species is variable,

especially in its lamina architecture.

1. Didymochlaena truncatula (Sw.) John Sm., J.

Bot. (Hooker) 4: 196. 1841. Figure lOa-c.

Adiantum lunulatum Houtt., Nat. Hist. 2, 14: 209. t.

100, f. 1. 1783, not Burm. 1768. TYPE: uncer-

tain, the figure may serve for identification.

Aspidium truncatulum Sw., J. Bot. (Schrader) 1800(2):

36. 1802, nom. nov. for Adiantum lunulatum

Houtt., and with the same type.

Didymochlaena lunulata Desv., Mem. Soc. Linn. Par-

is 6: 282. 1827, nom. superfl. tor Aspidium trun-

catulum Sw., and with the same type.

Stem stout, bearing dense, linear, brown scales.

Leaves to 2 m or rarely 3 m long, the petiole mod-

erately scaly to usually densely so toward the base.

Lamina 2-pinnate, the ultimate segments (pin-

nules) short-stalked, dimidiate, entire, obtuse, gla-

40

FIELDIANA: BOTANY

FIG. 10. Didymochlaena truncatula: a, stem and portion of lamina; b, pinnules, abaxial side; c, portion of

costule with pinnule bases, adaxial side. (From Stolze, Ferns & fern allies of Guatemala. 1981.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

41

brous or slightly scaly, the rachis with long pro-

cesses at the base of the pinnae on the adaxial side.

Sori covered by an elongate indusium, on each

side of a vein, and continuous distally.

Wet forests and rain forests, 1 1 0-900 m, Ama-

zonas to Cuzco and Madre de Dios.

Tropical America; Old World.

This is a highly distinctive species, unusual in

its pantropical distribution. In Peru, it has been

most commonly collected in Loreto.

Amazonas: Rio Maranon above Cascadas de Mayasi,

Wurdack 2015 (GH, us). Serrania de Bagua, above La

Peca, Harbour 2390 (MO). San Martin: Prov. Mariscal

Caceres, Tocache Nuevo, /. Schunke V. 6905 (F, MO,

us). Prov. Mariscal Caceres, Cerro Santa Cruz, /. Schunke

V. 8036 (MO, USM). Prov. Rioja, Pedro Ruiz-Moyabam-

ba, D. Smith 4430 (MO). Loreto: Rio Itaya, Soledad,

Killip & Smith 29768 (F, GH, us). Prov. Maynas, Yan-

amona, McDaniel 16195 (GH). Alto Amazonas, Andoas,

Vdsquez 4400 (MO). Huanuco: Tingo Maria, Stork &

Morion 9473 (F, uc, us). Pasco: Prov. Oxapampa, vicin-

ity of Chequitavo, D. Smith 5263 (MO). Junin: Chan-

chamayo Valley, C. Schunke 8 (F, us). Ucayali: Prov.

Coronel Portillo, between La Divisoria and El Boqueron,

Plowman & Kennedy 5768 (F, GH). Cuzco: Prov. Quis-

pichanchi, Punkiri, Vargas 15408, 16108 (GH). Madre

de Dios: Prov. Manu, Shintuya, Vargas 17837 (GH), Fos-

ter et al. 3143 (GH, us).

Plants terrestrial. Stem short-creeping to erect.

Lamina 1 -pinnate to 2-pinnate-pinnatifid, taper-

ing to a pinnatifid apex, glabrous on both surfaces,

with internal, pellucid (sometimes drying black)

punctate glands. Pinnae commonly linear to nar-

rowly oblong, the apices long-acuminate and ser-

rate. Rachis and costae stramineous to brown, scaly,

the scales thin, flaccid, usually ending in a papil-

lose-glandular tip. Groove of the costae pubescent

within, the trichomes less than 0. 1 mm long. Veins

free to irregularly anastomosing, curved-ascend-

ing, ending behind the margin in a clavate tip. Sori

round or slightly oblong near the costae. Indusium

absent, but some species with an indusiumlike

scale.

The genus contains about 25 species, with dis-

tribution from southern Mexico to southeastern

Brazil and the Antilles. The most distinctive fea-

ture of Stigmatopteris is the internal punctate

glands. These glands are most easily seen when

the leaf is held between the eye and a light, but

are also visible under a microscope (oblique light)

as tiny, raised bumps.

References

XI. Stigmatopteris

Contributed by Robbin C. Moran

Stigmatopteris C. Chr., Bot. Tidsskr., 29: 292.

1909. TYPE: Polypodium flavopunctatum

Kaulf. = Stigmatopteris rotundata (Willd.) C.

Chr. (Aspidium rotundatum Willd.). Figure 11.

Dryopteris subgenus Stigmatopteris (C. Chr.) C. Chr.,

Kongel. Danske Vidensk. Selsk. Skr. Naturvi-

densk. Afd., ser. 7, 10: 73. 1913.

CHRISTENSEN, C. 1 909. On Stigmatopteris, a new

genus of ferns with a review of its species. Bot.

Tidssk., 29: 291-304.

CHRISTENSEN, C. 1913. A monograph of the ge-

nus Dryopteris, part 1. The tropical American

pinnatifid-bipinnatifid species. Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 7,

10: 53-282.

CHRISTENSEN, C. 1 920. A monograph of the ge-

nus Dryopteris, part 2. The tropical American

bipinnate-decompound species. Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8,

6: 1-132.

Key to Species of Stigmatopteris

a. Veins free

b. Lamina 1-pinnate-pinnatifid

FIG. 1 1 . Stigmatopteris heterophlebia: a, habit; b, portion of pinna, abaxial side. Stigmatopteris pellucidopunc-

tata: c, portions of pinna segments, adaxial side, (a, b from Moran 3165, Costa Rica, F, c from Brant et al. 1679,

Colombia, F.)

42

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

43

a.

c. Pinnae cut ca. % to the costa 1 . S. pellucidopunctata

c. Pinnae entire or cut less than V4 to the costa 2. S. longicaudata

b. Lamina 2-pinnate-pinnatifid 3. S. lechleri

Veins anastomosing d

d. Rachis and costae densely puberulent 4. S. heterophlebia

d. Rachis and costae lacking trichomes 5. S. opaca

1. Stigmatopteris pellucidopunctata (C. Chr.) C.

Chr., Dot. Tidsskr. 29: 304. 1909. Figure lie.

Polypodium macrophyllum Hooker, Sp. fil. 4: 241.

1 862, nom. illeg., not (Blume) Mett. 1 856. TYPE:

Peru, San Martin, Mt. Guayrapurima, August

1856, Spruce 4720 (holotype, K.!; isotypes, BM!,

o!, P!, us!; photos, F, GH, MO, NY, all of BM).

Dryopteris pellucido-punctata C. Chr., Index fil. 283.

1905, nom. nov. for P. macrophyllum Hooker and

with the same type.

Lamina 1 -pinnate-pinnatifid. Pinnae 20-40 cm

long and 2.5-6 cm broad, 15-20 pairs, cut ca. %

to the costa, sessile or the basal ones short-stalked,

the stalk 2-7 mm long. Segments 4-10 mm broad,

serrate at the apex, the margins entire, serrate or

lobed. Rachis and costae stramineous or brown,

scaly, the scales ovate, lanceolate or filiform. Veins

free, 7-14 per segment, unbranched or 1 -forked

with a short acroscopic branch bearing the sorus.

Sori round.

Wet forests, 200-600 m, Amazonas, San Mar-

tin, Huanuco, Pasco, and Madre de Dios.

Colombia; Ecuador; Peru; Bolivia.

This species closely resembles S. ichtiosma (So-

diro) C. Chr., a species known only from the west-

ern cordillera of Ecuador but which eventually may

be found in Peru. It differs from S. pellucido-

punctata by the presence of a small, scalelike in-

dusium associated with the sorus and having only

lanceolate scales on the axes (i.e., it lacks filiform

scales).

Amazonas: Prov. de Bagua, Quebrada Tambillo (be-

low km 280 of Maranon road), valley of Rio Maranon

above Cascadas de Mayasi, Wurdack 2002 (GH, us, USM).

Huanuco: Fundo Chela, Sinchono, Aguilar 927 (USM).

Pasco: Paujil, near Puerto Bermudez, Leon 311 (USM).

Madre de Dios: Prov. Manu, Atalaya, vicinity of Ha-

cienda Amazonia, 2-3 km W of village, Foster & Wach-

ter 7437 (MO, USM).

2. Stigmatopteris longicaudata (Liebm.) C. Chr.,

Bot. Tidsskr. 29: 300. 1909.

Polypodium longicaudatum Liebm., Kongel. Vidensk.

Selsk. Skr. Naturvidensk. Afd., ser. 5, 1: 209.

1849. LECTOTYPE (designated by A. R. Smith,

Fl. Chiapas, part 2:213. 1981): Mexico, Veracruz,

Baranca de Huitamalco, Liebmann Flora Mex.

737 (c- Folio Herbarium; ISOLECTOTYPE, K!).

Dryopteris longicaudata (Liebm.) Maxon, Contr. U.S.

Natl. Herb. 13: 18. 1909.

Lamina 1 -pinnate. Pinnae mostly 14-20 cm long

and 2.0-2.5 cm broad, linear to narrowly lanceo-

late, 1 2-20 pairs, the margins entire or with cre-

nate lobes cut less than V4 to the costae, the upper

pinnae adnate to the rachis with a conspicuous

decurrent tapering basal wing, lower pinnae free,

subsessile or stalked, the stalk 2-5 mm long. Ra-

chis and costae glabrous to densely scaly, the scales

ovate to lanceolate, or (in Venezuela, Peru, and

Bolivia) linear and fibrillose. Veins free, 3-6 per

segment. Sori round, filiform scales present among

the sporangia in plants from Venezuela, Peru, and

Bolivia.

Wet forests, 220-2 100(-2500) m, San Martin,

Huanuco, Junin, Ucayali, Cuzco, and Madre de

Dios.

Southern Mexico; Guatemala; Costa Rica; Pan-

ama; Venezuela; Peru; Bolivia.

This species is distinctive in its decurrent, ta-

pering wings from the pinna bases, which are usu-

ally confluent in the distal V2 to !/3 of the lamina.

San Martin: Mt. Guayrapurima, near Tarapoto, Spruce

4012 (BM, G, K, P). Huanuco: Tingo Maria, Tryon &

Tryon 5330 (BM, F, GH). Junin: E of Quimiri Bridge, near

La Merced, Killip & Smith 23906 (NY, us). Ridge E of

Tingo Maria, Allard 22558 (us). Ucayali: Prov. Coronel

Portillo (as Loreto), Sinchono, cerca de la Divisoria,

Aguilar 861 (GH, USM). Sinchono (as Loreto), entre Tingo

Maria y Pucallpa, Aguilar 865 (GH). Cuzco: Prov. Pau-

cartambo, Cosnipata Valley, Rio Tono, first foothill ridge

on road N of Patria, Wachter et al. 200 (F). Madre de

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

10-15 km NNW of Shintuya, transect to ridgetop, Foster

et al. 10908 (F). Parque Nacional del Manu, Cocha Cashu

Biological Station, Foster P-84-91 (F).

44

FIELDIANA: BOTANY

3. Stigmatopteris lechleri (Mett.) C. Chr., Kongel.

Danske Vidensk. Selsk. Skr. Naturvidensk.

Afd., ser. 8, 6: 28. 1920.

Phegopteris lechleri Mett., Fil. lechl. 2: 25. 1859. TYPE:

Peru, Puno, "St. Gaban" (San Gaban), Lechler

2497 (holotype, B!; frag., BM!).

Polypodium punctatum Hooker, Sp. fil. 4: 262, 1862,

nom. illeg., not (L.) Swartz, 1802. LECTOTYPE

(designated by Christensen, Kongel. Danske Vi-

densk. Selsk. Skr. Naturvidensk. Afd., ser. 7, 10:

79. 1913): Peru, San Martin, Tarapoto, Mt.

Guayrapurima, August 1856, Spruce 4719 (K.!;

ISOLECTOTYPES, G!, P!; photos, F, MO of K).

Polypodium prasinum Baker, Syn. fil. 312. 1867, nom.

nov. for Polypodium punctatum Hooker and with

the same type.

Dryopteris prasina (Baker) C. Chr., Index fil. 285. 1905.

Stigmatopteris prasina (Baker) C. Chr., Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Adf., ser. 7,

10: 79. 1913.

Stigmatopteris ecuadorensis C. Chr., Kongel. Danske

Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8,

6: 29. 1920. TYPE: Ecuador (Andes of Quito),

Sodiro (holotype, B!; isotypes, BM!, P).

Lamina 2-pinnate-pinnatifid. Pinnae 25-40 cm

long and 5-15 cm broad, widest at the base. Pin-

nules 4-10 cm long and 1.0-2.3 cm broad, nar-

rowly lanceolate-triangular, sessile to broadly ad-

nate and decurrent, the lobes oblong to rectangular,

entire to serrate apically. Rachis and costae stra-

mineous or brown, scaly, the scales linear or nar-

rowly triangular, tortuous, fibrillose. Veins un-

branched or rarely (in the larger segments)

branched, nearly reaching the margin. Sori round.

Wet forests, 200-1700 m, San Martin, Huanu-

co, Pasco, and Puno.

Costa Rica; Venezuela; Colombia; Ecuador;

Peru.

This species, with its 2-pinnate-pinnatifid lam-

ina, is the most finely divided in the genus.

Huanuco: 25 km NE of Tingo Maria, La Divisoria,

Moran 3696 (MO, USM). Pasco: Prov. Oxapampa, Gran

Pajonal, north of Chequitavo, D. Smith 5082 (MO).

4. Stigmatopteris heterophlebia (Baker) R. C.

Moran, comb. nov. Figure lla-b.

Polypodium heterophlebium Baker, J. Bot. 22: 363.

1884. TYPE: Costa Rica, Prov. unknown, La-

guna, 305 m, Harrison 59 (holotype, K!).

Dryopteris heterophlebia (Baker) C. Chr., Index fil.

270. 1905.

Lamina 1 -pinnate. Pinnae 10-16(-18) cm long

and 2. 5-3. 5(— 4.5) cm broad, widest at or near the

base, the margins entire to crenate, the base adnate

and decurrent to or nearly to the next pinna pair,

free pinnae pairs 1-3, the basal pinnae usually

somewhat reduced, widest near the middle, stalked,

the stalk 2—4 mm long. Rachis and costae tan, light

brown, or stramineous, puberulent and scaly, the

hairs less than 0. 1 mm long, erect, capitate-glan-

dular (when fresh), the scales to 3 mm long, lan-

ceolate, not fibrillose. Veins anastomosing irreg-

ularly. Sori discrete or confluent where the veins

anastomose.

Wet forests, 100-1900 m, Loreto.

Nicaragua to Peru.

Differing from the closely related S. alloeoptera

(Kunze) C. Chr., which grows in Venezuela and

Colombia, by the glandular hairs on the axes and

fewer pinna pairs.

Loreto: Santa Rosa, lower Rio Huallaga below Yuri-

maguas, Killip & Smith 28984 (NY).

5. Stigmatopteris opaca (Baker) C. Chr., Kongel.

Danske Vidensk. Selsk. Skr., Naturvidensk.

Afd., ser. 7, 10: 78. 1913.

Meniscium opacum Baker, J. Bot. 166. 1877. TYPE:

Ecuador, Andes of Quito, Sodiro 54/3 (holotype,

K!; photos, BM, GH, us, all of K).

Polypodium oligophlebium Baker, Syn. fil. ed. 2, 506.

1874, nom. illeg., not Kunze, 1850. TYPE: Peru,

San Martin, Tarapoto, Cerro de Guayrapurima,

Spruce 4653 (holotype, K!).

Dryopteris christii C. Chr., Index fil. 257. 1905, nom.

nov. for Meniscium opacum, and with the same

type, not Dryopteris opaca (Don) C. Chr. 1905.

Dryopteris paucinervata C. Chr., Index fil. 283. 1905,

nom. nov. for Polypodium oligophlebium Baker,

and with the same type.

Lamina 1 -pinnate, opaque, the internal glands

not visible. Pinnae 12-27 cm long and 3-5 cm

broad, widest at or just above the base, the margins

entire to crenate, the base sessile, the upper pinnae

with a short decurrent base, the basal pinnae near-

ly equalling the above pinnae, widest near the mid-

dle, sessile. Rachis and costae lacking hairs, scaly,

the scales lanceolate to linear or amorphous. Veins

anastomosing, those of the sterile leaf with a single

excurrent vein projecting from the apex where the

two lateral veins join. Sori round, oblong, or ar-

cuate.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

45

2mm

""Oaf

5cm

FIG. 1 2. Cyclodium trianae: a, lamina; b, stem and petiole base. Cyclodium meniscioides var. meniscioides: c,

lamina apex; d. fertile pinna; e, sori. (a, b from Ellenberg 3257, Ecuador, GH; c, d from Fiebrig 6344, Paraguay, F,

e from Schunke V. 5610, F.)

46

FIELDIANA: BOTANY

Wet forests, 250-600 m.

Ecuador and Peru.

In Peru known only from Spruce's type collec-

tion ofPolypodium oligophlebium from San Mar-

tin.

XII. Cyclodium

Cyclodium Presl, Tent, pterid. 85. 1836. LEC-

TOTYPE (designated by John Sm., Hist. fil.

203. 1875): Aspidium confertum Kaulf. = Cy-

clodium meniscioides (Willd.) Presl. Figure

12a-b.

Peltochlaena Fee, (Mem. foug. 5) Gen. fil. 289. 1852,

nom. provis., illegit.

Stem usually rather short-creeping and moder-

ately stout, to long-creeping (to 5 m when climb-

ing), bearing scales. Leaves ca. 30 cm to 2 m long,

petiole continuous. Lamina 1 -pinnate to 2-pin-

nate-pinnatifid or rarely 3 -pinnate at the base, or

rarely simple, usually slightly short-pubescent or

scaly, monomorphic to dimorphic and then the

fertile more erect, longer, and with less expanded

segments. Veins free or anastomosing, with or usu-

ally without included free veinlets. Sori roundish,

borne on the veins, not paraphysate, covered by

often fugacious indusia which are peltate or or-

bicular-reniform with a narrow sinus. Spores el-

lipsoidal to spheroidal, monolete, slightly to

strongly ridged, usually finely papillate.

Cyclodium is a tropical American genus of ten

species. It was formerly treated in Stigmatopteris

(Tryon & Tryon, 1982), but the revision by Smith

(1986) clearly shows that it is distinct from, and

perhaps not closely related to, that genus.

There are two species in Peru, and an additional

one may occur there (see Comments).

This treatment has been adapted from that of

Smith (1986) and has been prepared with the col-

laboration of Dr. A. R. Smith.

References

SMITH, A. R. 1986. Revision of the Neotropical

fern genus Cyclodium. Amer. Fern J., 76: 56-

98.

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

matopteris, pp. 519-524 in Ferns and allied

plants, Springer- Verlag, New York.

Key to Species of Cyclodium

a. Veins all free; lamina 1 -pinnate-pinnatifid to 3-pinnate at the base, the apex gradually reduced . . .

l.C. trianae

a. Veins regularly anastomosing; lamina 1 -pinnate, with a conform (or nearly) apical segment

. 2. C. meniscioides

1. Cyclodium trianae (Mett.) A. R. Sm., Amer.

Fern J. 76: 92. 1986. Figure 12a-b.

Aspidium trianae Mett., Ann. Sci. Nat. Bot. 5, 2: 243.

1864. TYPE: Colombia, Prov. Barbacoas (Nari-

no), via de Tuquerres, Triana "32" (holotype, B;

isotype, BM; both seen by A. R. Smith; photos of

BM at uc, us).

Nephrodium firmifolium Baker, Syn. fil., ed. 2, 501.

1874. TYPE: Peru (San Martin), Mt. Guayapu-

rima, near Tarapoto, Spruce 4662 (holotype, K;

isotypes BM, BR, p; all seen by A. R. Smith).

Terrestrial, stem creeping. Leaves monomor-

phic or only slightly dimorphic, gradually reduced

at the apex. Lamina 1 -pinnate-pinnatifid to usu-

ally 2-pinnate or 2-pinnate-pinnatifid, or 3-pin-

nate only at the base. Veins free. Sori with orbic-

ular-reniform indusia with a narrow sinus, or rarely

peltate.

In dense primary forests or partly cleared for-

ests, 300-900 m, Lambayeque and Loreto, south

to Junin.

Panama, south to Peru.

Var. trianae occurs throughout the range of the

species. In South America it is generally confined

to the Amazonian side of the Andes. Var. cho-

coense A. R. Sm. is only in Panama and the Pacific

side of the Andes in Colombia. The only repre-

sentative in Peru is var. trianae.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

47

Lambayeque: Puerto Nazareth, 5 km from Olmos, El-

lenberg 3449 (GH). Loreto: Balsapuerto, Killip & Smith

28540 (GH, NY, us). Pasco: Prov. Oxapampa, Rio Pal-

cazu, cerca de Iscozacin, Leon 712 (GH). Prov. Oxapam-

pa, Palcazu valley, Iscozacin, Foster 9498 (MO). Junin:

Prov. Satipo, above Pichanaki, Leon 216 (GH).

the fertile then more erect and with shorter and

narrower pinnae. Lamina 1 -pinnate or rarely sim-

ple, with a conform (or nearly) apical segment, the

pinnae entire to strongly crenulate. Veins anas-

tomosing, usually with included free veinlets. Sori

with peltate indusia.

2. Cyclodium meniscioides (Willd.) Presl, Tent,

pterid. 85. 1836.

Terrestrial or epiphytic and short-climbing to 2

m. Leaves subdimorphic to strongly dimorphic,

Trinidad; and widely distributed in tropical

South America.

In addition to the two varieties in Peru, there is

var. rigidissimum (C. Chr.) A. R. Sm., which rarely

occurs in Guyana and perhaps in Venezuela.

Key to Varieties

a. Margins of the pinnae entire, sinuate, crenulate to crenate 2a. var. meniscioides

a. Margins of the pinnae, especially toward the apex of the sterile ones, sharply serrate

2b. var. paludosum

2a. Cyclodium meniscioides var. meniscioides.

Figure 12c-e.

Aspidium meniscioides Willd., Sp. pi. ed. 4, 5: 218.

1810. TYPE: Brazil, Hoffmannsegg (holotype, B!,

Herb. Willd. 19737; photo, GH).

Stigmatopteris meniscioides (Willd.) Kramer, Proc.

Kon. Nederl. Akad. Wetensch. C, 71: 521. 1968.

In dense primary forests, in swampy forests

where sometimes epiphytic, or in partly disturbed

forests, rarely on fallen logs or epiphytic to 2 m,

100-650 m, Amazonas south to Madre de Dios.

Range of the species.

Amazonas: Near Pongo Mori, Rio Comain, Berlin 949

(MO, uc). Prov. Bagua, 34 km NE of Chiriaco, Barbour

4401 A (MO). San Martin: Prov. Mariscal Caceres, To-

cache Nuevo, J. Schunke V. 5610 (F, us). 23 km S of

Nuevo San Martin, Gentry et al. 37573 (MO). Loreto: 7

km SW of Iquitos, Croat 18579 (GH, MO, uc). Above

Pongo de Manseriche, Mexia 6 193 a (GH, uc, us). Mishu-

yacu, near Iquitos, Klug 241 (F, us). Pasco: Puerto Ber-

mudez (as Junin), Killip & Smith 26566 (us). Prov.

Oxapampa, Palcazu valley, D. Smith 3733 (MO, uc).

Ucayali: Vicinity of Aguaytia, Mathias & Taylor 5138

(F). Madre de Dios: Prov. Manu, Cerro de Pantiacolla,

Foster et al. 10704 (F).

2b. Cyclodium meniscioides var. paludosum

(Morton) A. R. Sm., Amer. Fern J. 76: 87.

1986.

Dryopteris paludosa Morton, Bull. Torrey Bot. Club

66: 50. 1939. TYPE: Colombia, Antioquia, Puer-

to Berrio, Pennell 3723 (holotype, NY; frag., us,

both seen by A. R. Smith).

Stigmatopteris paludosa (Morton) R. & A. Tryon,

Rhodora83: 136. 1981.

160 m, Loreto.

Colombia and Peru.

Loreto: Bersalles-Iquitos, Vargas 11476 (GH).

Comments

Cyclodium guianense (Klotzsch) Gomez, Phyto-

Iogia60: 371. 1986.

Aspidium guianense Klotzsch, Linnaea 20: 364. 1847.

TYPE: "British Guiana," Schomburgk 1157 (ho-

lotype; B?; isotypes, K, uc, seen by A. R. Smith).

Stigmatopteris guianensis (Klotzsch) C. Chr. Ind. fil

suppl. 3: 174. 1934.

This species occurs in Trinidad, northern South

America, and in the Amazon basin of Colombia,

and may occur in Peru. It has all of the veins free,

as in C. trianae, but has a 1 -pinnate lamina rather

than a 2-pinnate to 2-pinnate-pinnatifid one, and

peltate indusia rather than orbicular-reniform ones,

these with a narrow sinus. Only rarely does C.

trianae have peltate indusia.

48

FIELDIANA: BOTANY

XIII. Polystichum

Polystichum Roth, Tent. fl. germ. 3: 31, 69. 1799.

TYPE: Polystichum lonchitis (L.) Roth (Poly-

podium lonchitis L.). Figure 13.

Plecosorus Fee, (Mem. fam. foug. 5) Gen. fil.: 150.

1852. TYPE: Plecosorus mexicanus Fee = Ple-

cosorus speciosissimus (Kunze) Moore = Polys-

tichum speciosissimum Kunze) R. & A. Tryon.

Stem decumbent to erect, often small, to stout,

bearing stramineous to blackish scales. Leaves 5

cm to often 1 m or rarely 3 m long, petiole con-

tinuous. Lamina usually 1 -pinnate, 2-pinnate, or

to 3-pinnate-pinnatifid, rarely entire to pinnatifid,

more or less scaly, often densely so, monomorphic

or rarely dimorphic, adaxial side of the costa sul-

cate to nearly flat. Veins free. Sori round, borne

on the veins, not paraphysate, covered by per-

sistent to fugacious, peltate indusia, or (in all Pe-

ruvian species) exindusiate. Spores ellipsoidal to

spheroidal, monolete, saccate, cristate, or ridged,

often spinulose and frequently perforate.

Polystichum is a large, nearly worldwide genus

of about 1 50 species. There are perhaps 25 species

in America and four in Peru. The species of An-

dean Polystichum are not well known, and the

following treatment is tentative. Species of Polys-

tichum are evidently variable genetically and cy-

tologically and they also vary in different envi-

ronments. Population studies, cytological studies,

and broad morphological comparisons are needed

in order to recognize and define the taxa (Bar-

rington, 1985). Currently unrecognized hybrids

may be an important element in the apparent in-

tergradation of taxa.

Although characters of the scales of the leaf axes

are emphasized in the key to species by Smith

(1985), among others, these have been found to

be usually variable in Peru and are not generally

employed in this treatment. However, these struc-

tures need further investigation and may be more

important than now considered.

Among the synonyms, only those names based

on Peruvian material and those that are in general

use are included. The names of many species rec-

ognized, especially from Ecuador, Bolivia, and

southeastern Brazil, have not otherwise been in-

cluded.

Although Christensen (Index fil. 727. 1906) in-

dicated that the first publication of the generic

name Polystichum was in Arch. Bot. (Leipzig) 2( 1 ).

1 799, Taxonomic Literature-2 indicates that Roth's

publication was in 1 799 while Arch. Bot. (Leipzig)

2(1) was in 1800.

This treatment has been prepared with the per-

spective and critical collaboration of Dr. David S.

Barrington, who has seen some of the types cited.

The key will be adequate for the identification

of many complete collections. However, in some

cases, a concordance of the majority of the char-

acters mentioned may be necessary for identifi-

cation, and rarely materials may represent a vari-

ation that is not accounted for in the key.

References

BARRINGTON, D. S. 1985. The present evolu-

tionary and taxonomic status of the fern genus

Polystichum: The 1984 Botanical Society of

America Pteridophyte Section Symposium.

Amer. Fern J., 75: 22-28.

SMITH, A. R. 1985. Pteridophytes of Venezuela,

an annotated list, pp. 1 89-1 92, published by the

author, Department of Botany, University of

California, Berkeley.

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

tichum, pp. 524-538, in Ferns and allied plants,

Springer- Verlag, New York.

Key to Species of Polystichum

a. Petiole with scales few or not persistent above its base, varying to densely scaly, scales mostly small,

to ca. 0.5 cm long, elongate, rarely larger and broad; rachis with flattish scales and/or fibrils; pinnules

with revolute margins or plane (flat) b

b. A scaly bud usually present in the axil of a pinna near the apex of the lamina, or more than one

present; apical portion of the lamina usually prolonged, with a 1 -pinnate portion 5-10 cm long

with pinnae mostly 1-2.5 cm long, or longer 2. P. platyphyllum

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

49

FIG. 13. Polystichum pycnolepis: a, habit. Polystichum platyphyllwn: b, habit; c, distal pinnae, with proliferous

bud; Polystichum montevidense var. montevidense: d, pinnule, (a from Madison 1034, GH; b, c from Macbride

4034, F, d from Soukup 4329, F.)

50

FIELDIANA: BOTANY

b. Apex of the lamina lacking a scaly bud, usually gradually reduced, if prolonged the 1 -pinnate

portion short with most pinnae less than 1 cm long c

c. Pinnules definitely mucronate, usually plane (flat), or if the margins revolute then the apical

mucro plane; lamina 2-pinnate; if the lamina less than 8 cm broad then the basal pinnae reflexed

and/or the lamina apex prolonged, the apex of the pinnae acute to attenuate; or if the lamina

over 8 cm broad then the basal pinnae patent or reflexed 1 . P. montevidense

c. Pinnules sometimes definitely mucronate and plane, usually not or hardly mucronate, often

with the margins and apical mucro revolute; lamina 2-pinnate to 3-pinnate; the lamina usually

less than 8 cm broad and the basal pinnae patent or ascending, the lamina apex not or hardly

prolonged, and the apex of the pinnae obtuse to subacute; or if the lamina over 8 cm broad,

then the basal pinnae ascending 3. P. orbiculatum

a. Petiole with scales usually persistent above its base, mostly large, ca. 1-2 cm long and sometimes to

1 cm broad; rachis, at least at the base, usually with some brown or darker, involute or twisted scales,

these often also on the petiole; pinnules with revolute margins; apex of the lamina lacking a scaly

bud 4 P. pycnolepis

1 . Polystichum montevidense (Sprengel) Rosenst.,

Hedwigia46: 111. 1906.

Leaves ca. 10 cm to 1.5 m long. Petiole scales

sparingly persistent or persistent, especially at or

near the base, or petiole nearly devoid of scales;

scales brown to dark brown or darker, less than 1

cm long or rarely longer. Lamina 2-pinnate to rare-

ly 3-pinnate at the base of some pinnae; ca. 2—40

cm broad; the apex lacking a bud, gradually to

sometimes rather abruptly reduced, or prolonged.

Rachis bearing a few to many light brown to dark

brown fibrils and narrow scales, or sometimes

broader ones, or nearly devoid of scales. Pinnae

mostly patent, the apex acute to usually attenuate

and deeply pinnatifid to the tip; basal pinnae re-

duced or not, patent or reflexed; pinnules plane

(flat) or nearly so.

Key to Varieties of Polystichum montevidense

a. Leaves ca. 50 cm to 1.5 m, mostly 60-80 cm long; lamina ca. 10 to 40 cm broad, mostly 2 to 2'/2

times as long as broad, the apex gradually to sometimes rather abruptly reduced, the portion beyond

the 2-pinnate portion shorter than the longest pinna; basal pinnae not or slightly reduced, patent or

somewhat ascending la. var. montevidense

a. Leaves ca. 10-90 cm, mostly ca. 30 cm long; lamina ca. 2-1 1 cm, mostly ca. 6 cm broad, mostly 3

to 4 times as long as broad, the apex usually prolonged, the portion beyond the 2-pinnate portion

longer than the longest pinna; basal pinnae reduced and/or reflexed 1 b. var. nudicaule

1 a. Polystichum montevidense var. montevidense.

Figure 13d.

Polypodium montevidense Sprengel, Syst. veg., ed. 16,

4: 59. 1827. TYPE: Uruguay, Monte Video, Sello

(not located).

Polystichum lehmannii Hieron., Hot. Jahrb. Syst. 34:

452. 1904. TYPE: Colombia, Prov. Cauca, Los

Motilones, Lehmann 3674 (holotype: B; isotype,

us!).

Polystichum mexiae Copel., Univ. Calif. Publ. Bot.

19: 299. 1941. TYPE: Ecuador, Prov. Pichincha,

Nono to Mindo, Mexia 7679 (holotype, uc;

isotypes, F!, GH!).

In dry rocky places and on open hillsides, more

often in forests in rocky places, and in cloud for-

ests, 1800-3600 m, Piura to Puno.

Venezuela and Colombia, south to Bolivia, east

through Argentina to Uruguay and southeast Bra-

zil; possibly in Central America.

Recent studies have usually confined Polysti-

chum montevidense to northwestern Argentina and

eastward to southeastern Brazil. However, Peru

materials are not separable, and the species is best

regarded as an Andean one, with an extension east-

ward. Intergradation between var. montevidense

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

51

and var. nudicaule indicates that the two are not

distinct species. In spite of the several characters

presented in the key, there is considerable overlap

between the two taxa.

Plants with the leaves especially scaly on the

rachis and pinna-rachises have been referred to

Polystichum montevidense var. squamulosum (Hi-

eron.) Hieron., but this seems appearance to be a

part of the normal variation of the species in Peru.

Polystichum yungense Rosenst. of Bolivia is prob-

ably a large, well-developed variation of this spe-

cies.

Piura: Ayabaca, Soukup 4329 (F, us). Cajamarca:

Prov. Contumaza, Cascas-Contumaza, Lopez et al. 9109

(GH). Prov. Hualgayoc, Monte Seco, Soukup 3816 (F,

us). Prov. San Miguel, Cerro Quillon, Mostacero et al.

1293 (F, GH). Amazonas: Prov. Bagua, SE of La Peca,

Barbour 3696 (uc), 3984 (MO). San Martin: Lamas, Bel-

shaw 3428 (GH, uc, us). Prov. Mariscal Caceres, Rio

Abiseo National Park, Young & Leon 4586 (F). Huanuco:

Mito, Macbride & Featherstone 1698 (F, GH, us). Yana-

no, Macbride 3827 (F, us). Pasco: Prov. Oxapampa, San

Alberto, van der Werff et al. 8433 (MO, uc). Ayacucho:

Between Huanta and Rio Apurimac, Killip & Smith 23239

(GH). Cuzco: Prov. Urubamba, Chupani, Vargas 11 126

(GH). Prov. La Convention, between Rumichurco and

Alcobamba, Davis et al. 1227 (F, GH). Puno: Prov. San-

dia, entre Sandia y Cuyocuyo, Ferreyra 16818 (GH, USM).

Ib. Polystichum montevidense var. nudicaule

(Rosenst.) Tryon, comb. nov.

Polystichum nudicaule Rosenst., Repert. Spec. Nov.

Regni Veg. 11: 56. 1912. SYNTYPES: Bolivia,

Unduavi, 3400 m, Buchtien 2657, 2658, 2659,

2662 (not located); ISOSYNTYPES, Buchtien

2657 (us), 2655 (uc!, us), 2659 (us).

Rocky crevices, on cliffs, at base of rocks, and

in boggy places in heath, 1 800-4000 m, Piura to

Puno.

Venezuela and Colombia, south to Bolivia; per-

haps in southeastern Brazil and Chile.

This variety is distinguished especially by its

prolonged lamina apex and the usually reflexed

and reduced basal pinnae. It intergrades with var.

montevidense and perhaps represents a variation

of it growing under less favorable conditions. Po-

lystichum wolfii Hieron. from Bolivia is probably

this variety.

A few collections have some leaves on the stem

with closely spaced and nearly entire pinnules and

also some that have widely spaced and rather la-

ciniate pinnules.

Piura: Prov. Huancabamba, above Cachaque on road

to Huancabamba, Hutchison 1643 (GH, uc). Cajamarca:

Prov. Cajamarca, Cajamarca-Bambamarca, Smith &

Vdsquez 3477 (uc). Prov. Contumaza, Las Achiras, Sa-

gdstegui 3887 (GH, HUT). Amazonas: Prov. Chachapoyas,

NNE of Diosan, Wurdack 1596 (F, GH, uc, us). La Liber-

tad: Prov. Santiago de Chuco, Laguna El Toro, D. Smith

2300 (GH). San Martin: Rio Abiseo National Park, Young

& Leon 4401 (F). Huanuco: Cani, Macbride 3459 (F, us).

Tambo de Vaca, Macbride 4420 (F, us). Lima: Prov.

Huarochiri, Dist. San Mateo, Sounders 324 (F, uc). Rui-

nas de Cantamarca, Coronado 297 (GH, uc). Junin: In-

cahuasi, Soukup 3177 (F). Near Huancayo, Killip & Smith

23366 (us). Ayacucho: Between Huanta and Rio Apu-

rimac, Killip & Smith 22254 (us). Apurimac: Prov. An-

dahuaylas, Dist. Chincheros, Sounders 737 (GH). Prov.

Abancay, Curahuaci, Marin 1961 (F). Cuzco: Prov. Cuz-

co, Saxihuaman, Cuzco, Tryon & Tryon 5359 (GH, us).

Taucca, Davis et al. 1540 (F). Puno: Granja Salcedo,

Soukup 66 (F). Have, Soukup 872 (F).

2. Polystichum platyphyllum (Willd.) Presl, Tent,

pterid. 84. 1836. Figure 13b-c.

Aspidium platyphyllum Willd., Sp. pi., ed. 4, 5: 255.

1810. TYPE: Venezuela, Caracas, Bredemeyer

(holotype; B!, Herb. Willd. 19879; photo, GH).

Leaves ca. 40 cm to 1.5 m long. Petiole scales

sparingly or not persistent, light brown to brown,

or especially near the base dark brown; less than

1 cm long, rarely longer. Lamina 2-pinnate, or

rarely nearly 3 -pinnate at the base of some pinnae,

ca. 10-30 cm broad; the apex with the axil of a

pinna bearing a scaly bud, or more than 1 axil

bearing a bud, the apex usually prolonged, very

gradually, or sometimes abruptly reduced. Rachis

bearing mostly few, sometimes many, light brown

to brown fibrils and narrow scales, some broader

scales also often present. Pinnae patent, apex acute

or to attenuate and deeply pinnatifid to the tip;

basal pinnae the longest to somewhat reduced,

usually slightly ascending or patent, rarely re-

flexed; pinnules plane (flat).

In primary forests, on steep forested slopes, in

ravines and along stream banks, and on open,

rocky, moist road banks, 600-2800 m, Cajamarca

and Amazonas to Puno.

Southern Mexico and Central America; Jamai-

ca; Venezuela and Colombia, south to northwest-

ern Argentina and east to southeastern Brazil.

Nearly all leaves of this species have a bud near

the apex of the lamina, and perhaps all plants have

at least one leaf with a laminar bud. In a collection

52

FIELDIANA: BOTANY

from a large population (Try on & Try on 5404, GH)

all leaves except one very small one have a bud.

In old leaves, this bud may develop into a small

plantlet, with roots and small leaves. Other dis-

tinctive characters of this species are mentioned

in the key.

Cajamarca: Prov. Santa Cruz, Monte Seco, Sagdstegui

et al. 12407 (F, MO). Prov. Hualgayoc, Hacienda Taulis,

Hutchison & Bismark 6331 (F, GH, MO, uc, us). Ama-

zonas: Prov. Chachapoyas, Quebrada Molino, Wurdack

771 (F, GH, uc, us). Prov. Bagua, 12 km E of La Peca,

Barbour 2400 (F, MO, uc). San Martin: Near Tarapoto,

Spruce 3965 (BM, GH, us). Between Tarapoto and Yu-

rimaguas, Croat 58100 (F). Huanuco: Mima, Bryan 548

(F, GH, us), Macbride 4034 (F, GH, us). Prov. Huanuco,

near Riachuela Contalagua, Mexia 8297 (F, GH, MO, uc,

us). Pasco: Pichis Trail (as Junin), between San Nicolas

and Azupizu, Killip & Smith 26120 (us). Prov. Oxapam-

pa, 5 km SE of Oxapampa, D. Smith 2915 (GH). Junin:

Chanchamayo valley, C. Schunke 13 (F, us). Near La

Merced, Killip & Smith 23911 (GH, us). Prov. Chancha-

mayo, cerca a Monobamba, Fernandez & Vargas 204

(USM). Ayacucho: Between Huanta and Rio Apurimac,

Killip & Smith 22443 (us). Apurimac: Prov. Andahuay-

las, Alavarado (USM). Cuzco: Prov. Urubamba, near town

of Machu Picchu, Tryon & Tryon 5404 (BM, F, GH, uc,

us). Prov. Urubamba, camino a Winayhuayna (Huinay-

huayna), Chavez 3440 (GH, MO). Puno: Prov. Carabaya,

Vargas 17529 (GH).

3. Polystichum orbiculatum (Desv.) Remy, in Gay,

Hist. Fis. Pol. Chile, Bot. (Fl. Chilena) 6:515.

1853 (as Polystichum orbiculare by Christ,

Ark. f. Bot. 4[12]: 3. 1905).

Leaves ca. 1 5 cm to 1 m or more long. Petiole

scales persistent or sparingly so; usually brown to

dark brown near the base and brown to whitish

beyond; scales less than 1 cm long, rarely longer.

Lamina 2-pinnate to 3-pinnate, or rarely small

ones 1 -pinnate-pinnatifid, ca. 2-25 cm broad; the

apex lacking a bud, shortly acute to gradually re-

duced and attenuate. Rachis bearing a few to many

brown to whitish scales, these broad or sometimes

narrow and grading into fibrils. Pinnae mostly as-

cending, less often patent, the apex obtuse to less

often subacute; basal pinnae somewhat reduced or

not, mostly patent, sometimes ascending; pinnules

plane (flat) or usually with revolute margins and

apex.

The new names of Pteridophyta in Gay, Flora

Chilena 6: 470-549 have been variously attributed

to Gay, to Fee, to Remy, to Fee & Remy, and to

Remy & Fee. All of the new nomenclature is ev-

idently to be attributed to Remy (as Remy, in Gay.

Fl. Chil. 6:) and we have followed this citation. In

his Cat. method, chron. public. A. L. A. Fee, 1 863,

Fee does not mention any contribution to the Flo-

ra Chilena of Gay, and in 1858, Sturm in Abhandl.

natur.-hist. ges. Nurnberg 2: 151-202 (reprint 1-

52) nine of the 1 1 new species are credited to Remy.

Looser in Rev. Universit. (Santiago, Chile) 1 5(7):

694-717. 1930, credits the treatments of pterido-

phytes to Remy, although in later publications, on

Blechnum for example, he cites species authors as

Remy & Fee. It seems that the original treatment

was prepared by Remy and that Fee had some

editorial role with the manuscript.

Key to Varieties of Polystichum orbiculatum

a. Lamina usually 2-pinnate, rarely 1 -pinnate-pinnatifid or 3-pinnate only at the base of some pinnae

3a. var. orbiculatum

a. Lamina 3-pinnate well beyond the base of the pinnae 3b. var. boboense

i. Polystichum orbiculatum var. orbiculatum

Aspidium orbiculatum Desv., Ges. Naturf. Freunde

Berlin Mag. Neuesten Entdeck. Gesammten Na-

turk. 5: 321. 1811. TYPE: "America australi"

(holotype, P!, Herb. Desvaux; photo, GH, us).

Nephrodium polyphyllum Presl, Reliq. haenk. 1: 37.

1825. TYPE: Peru, Haenke (holotype, PR! or PRC;

photo, BM of PR).

Nephrodium trapezoides Presl, Reliq. haenk. 1: 37, /.

6,f. 1. 1825. TYPE: Peru, Haenke (holotype, PR

or PRC?; isotype, K!).

Polypodium rigidum Hooker & Grev., Icon. fil. /. 163.

1830, not Lam. & DC. 1805. TYPE: "Andium

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

53

Peruvianorum" (Ecuador), Monte Pichincha,

Jameson (holotype, K!).

Polystichum polyphyllum (Presl) Presl, Tent, pterid.

83. 1836.

Polystichum haenkeanum Presl, Tent, pterid. 83. 1 836,

nom. nov. for Nephrodium trapezoides Presl, not

Polystichum trapezoides (Sw.) Presl, and with the

same type.

Jamesonia paleacea Kunze, Hot. Zeit. 1844: 739, not

Polystichum paleaceum (Borkh.) Schwarz, 1949.

TYPE: Venezuela, Prov. Merida (also "Cara-

cas"), Linden 505 (holotype, B; isotypes, E!, GH!,

K!; photos, GH of BR and BM may be this species,

or they and the holotype may be P. pycnolepis).

Polypodium saxatileK\o\zsch, Linnaea 20: 383. 1847.

TYPE: Peru, Herb. Ruiz 65 (holotype, B?). Hi-

eronymus (Hedwigia, 46: 358. 1907) treats this

as a variety of Polystichum orbiculatum.

Polypodium crenatodentatum Klotzsch, Linnaea 20:

384, 1847. TYPE: Peru, Ruiz 73 (holotype, B?)

Hieronymus (op. cit., p. 357) treats this as a va-

riety of Polystichum orbiculatum.

Commonly growing in rocky, shaded places, at

the edge of rocks, or on cliffs, on sandstone and

on limestone, less often on grassy or shrubby hill-

sides, or at high altitudes in the puna, 2700-4900

m, mostly 3200-^*000 m, Lambayeque to Puno.

Southern Mexico; Costa Rica; Venezuela and

Colombia south to Bolivia.

Polystichum orbiculatum is a highly variable

species with most of the more distinctive varia-

tions included in var. orbiculatum. One extreme,

especially characteristic of the puna, has erect leaves

with strongly ascending pinnae that are closely set

and the pinnules have strongly revolute margins.

Another extreme has spreading leaves with mostly

patent and spaced pinnae and the pinnules are flat.

Another variation has the lamina relatively broad

and the pinnae and pinnules sharply acute to acu-

minate. These all intergrade to an extent that im-

plies they are variations within one taxon.

The label on Soukup & Guillen 5386 (us): "Iqui-

tos" (Loreto) is undoubtedly an error. The species

grows at much higher altitudes than the ca. 100

m of Iquitos.

Lambayeque: 7 km NW of Incahuasi, Sillon A Skill-

man 4110(r, GH, uc). Cajamarca: Prov. Hualgayoc, cer-

ca a Hualgayoc, Ferreyra 8539 (GH, USM). Prov. Con-

tumaza, Casabamba. Sagdstegui et al. 10721 (F).

Amazonas: NNE of Diosan. Wurdack 1597 (F, GH, uc,

us). La Libertad: Prov. Santiago de Chuco, above Cachi-

cadin. Stork & Morton 9976 (F, uc, us). Prov. Pataz,

entre Retamas y La Paccha, Lopez & Sagdstegui 3592

(GH). San Martin: Prov. Mariscal Caceres, Rio Abiseo

National Park, Young & Leon 4402 (F). Ancash: Prov.

Pallasca. Conzuzo, Lopez 1165 (GH, us). Prov. Yungay,

above Yungay, Tryon A Tryon 6557 (F, GH). Lima; Be-

tween Parac and Toncuyo, Coronado 310 (GH, uc). Prov.

Huarochiri, Mina Perdida, Cerrate 2040 (USM). Pasco:

Puna de Huaron, Humbert 30894 (GH). Prov. Pasco,

Huayllay, Boeke 1086 (AAU, MO). Jonin: Near Huancayo,

Killip A Smith 23363 (GH, us). Ca. 1 8 km SSE of Tarma,

Iltis et al. 129 (GH, uc, us). Huancav elica: Prov. H uan-

cavelica. westward from Huancavelica, Stork A Horton

10851 (F, uc, us). Prov. Tayacaja, arriba de Toca, Tovar

2018 (GH, USM). Ayacucho: Pumaccuanca pass, West 3658

(uc). Prov. Huanta, Ayacucho-Huanta, Sounders 727

(GH). Apurimac: Bosques de Ampay, Vargas 1061 (GH).

Cuzco: Prov. Urubamba. Machu Picchu, Peyton A Pey-

ton 1055 (GH, MO). Prov. Espinar, Hacienda Chachachi.

Vargas 10812 (GH). Arequipa: Cailloma, Mutter A Mul-

ler 2151 (GH). Puno: Salcedo, Soukup 275 (F, GH, uc).

Capachica Peninsula, Tutin 1071 (A, BM).

3b. Polystichum orbiculatum var. boboense i Hi-

eron.) Tryon, comb. nov.

Polystichum boboense Hieron., Hedwigia 46: 358. /.

8,f. 25. 25a. 1907. TYPE: Ecuador, Rio Bobo,

near Tulcan. Stubel 353, in part (holotype, B).

Polystichum sodiroi Christ, Ark. f. BoL 4( 1 2): 4. 1 905,

TYPE: Ecuador, Quito, Sodiro (not located).

In ravines, 2800-3000 m, Piura.

Ecuador and Peru.

This variety is the most distinctive among the

variations of Polystichum orbiculatum. The lam-

ina is fully 3-pinnate and often the leaves are large,

up to 1 mm or more long. Intermediates between

the two varieties, with the lamina 3-pinnate only

at or near the base of some pinnae, are present

throughout most of the range of var. orbiculatum.

Piura: Prov. Huancabamba, above Huancabamba.

Hutchison 1608 (uc). Prov. Huancabamba, Huanca-

bamba-Cuello del Indio, Sagdstegui et al. 8256 (F, uc;

at MO this collection is toward var. boboense).

4. Polystichum pycnolepis (Klotzsch) Moore, In-

dex fil. 92. 1858. Figure 13a.

Aspidium pycnolepis Klotzsch, Linnaea 20: 365. 1847.

SYNTYPES: Venezuela, Merida, Paramo de Cu-

lata, Aforitz 295, Moritz 296; Karasten "Coll. II,

a, b". (B?); ISOSYNTYPES: Moritz 295 (E!), (BM!,

E!, K!; photo, GH of P), Moritz s.n. (GH!).

Aspidium gelidum Klotzsch, Linnaea 20: 365. 1847.

TYPE: Venezuela, Merida, Paramo de Culata.

Moritz 294 (holotype, B?; isotypes, BM, K!; photo,

GH of BM).

Polypodium cochleatum Klotzsch, Linnaea 20: 388.

1847. TYPE: Peru, Ruiz 63 (holotype, B?).

Polystichum gelidum (Klotzsch) Fee, (Mem. foug. 5)

Gen. fil. 278. 1852.

Polystichum cochleatum (Klotzsch) Hieron., Jahrb. BoL

Syst. 34: 452. 1904.

54

FIELDIANA: BOTANY

Leaves ca. 35 cm to 1 m long. Petiole scales

more or less persistent, especially toward the base,

dark brown to atropurpureous, or some light

brown, often many ca. 2 cm long and to 1 cm

broad. Lamina 2-pinnate, very rarely nearly

3-pinnate at the base of some pinnae, ca. 8-20 cm

broad, rarely 5-7 cm broad; the apex lacking a

bud, gradually to rather abruptly reduced. Rachis

bearing usually very many brown fibrils and brown

or darker, long, often involute or twisted, narrow

scales, some lanceolate scales and broader ones

usually also present. Pinnae patent to strongly as-

cending, the apex subacute to acuminate and deep-

ly pinnatifid to the tip; basal pinnae reduced or

not, usually ascending, sometimes patent or re-

flexed; pinnules with revolute to strongly revolute

margins, the tip usually revolute.

Most commonly growing on open, rocky hill-

sides, in rock crevices, and at the edge of large

rocks, also on grassy or shrubby slopes and in

heath lands, 1 700 to usually 3000-4400 m, Ama-

zonas to Puno.

Venezuela and Colombia south to Bolivia.

Aspidium pycnolepis is evidently the correct ba-

sionym, among those names proposed by Klotzsch

in the same publication. Aspidium gelidum was

reduced to Phegopteris pycnolepis by Mettenius

(Abh. SenckenbergNaturf. Ges. 2; 295. 1858; Uber

einige Farngattugen, 4; 1 1. 1859) and Polypodium

cochleatum (as Phegopteris cochleata Mett.) was

reduced to Polypodium pycnolepis by Hooker (Sp.

fil. 4: 248. 1862).

This species is characterized by dense, brown

fibrils on the rachis and narrow, involute or twist-

ed, usually brown or darker scales, especially to-

ward the apex of the petiole and on the rachis.

The usually large petiole scales are persistent.

Amazonas: Prov. Chachapoyas, Cerro Campanario,

Wurdack 1589 (F, GH, uc, us). La Libertad: Between

Huamachuco and Cajambamba, Correll & Smith P924

(GH). Huanuco: Tambo de Vaca, Macbride 4369 (F, GH,

us). Mito, Macbride & Featherstone 1699 (F, us). Apo-

rimac: Prov. Abancay, Dist. Abancay, Sounders 762 (GH).

Cuzco: Prov. Calca, Vargas 15583 (GH). Prov. Urubam-

ba, Chincheros, Taucca, Davis et al. 1572 (F). Puno: Prov.

Sandia. S of Limbani, Metcalf 30436 (F, GH, MO, uc, us).

Prov. Huancane, Occa Pampa, Shepard 74 (GH).

Comments

Polystichum muricatum (L.) Fee, (Mem foug. 5)

Gen fil. 278. 1852.

Polypodium muricatum L., Sp. pi. 1093. 1753. TYPE:

Haiti, Petiver, Pteri-graphia Amer., no. 53, /. 1,

f. 6. 1712. (evidently a poor copy of Plumier,

Traite foug. Amer. t. 39).

Polystichum moritzianum (Klotzsch) Hieron., Hed-

wigia46: 354. 1907.

This species has been commonly recorded from

Peru; however, the collections are mostly either P.

platyphyllum or P. montevidense. Polystichum

muricatum is a species of Mexico and Central

America, the Greater Antilles, and the northern

part of South America.

It is characterized by large leaves, to 1 m or

more long, attenuate pinnae that are deeply pin-

natifid to the tip, flat pinnules, large, and brown

or darker petiole scales that are often short-ciliate,

and usually persistent well above the base, and

especially by the rachis bearing only fibrils and

very narrow scales, rather than some broad scales.

The species is commonly indusiate, although Smith

(1985) indicates that in Venezuela it is both in-

dusiate and exindusiate.

XIV. Olfersia

Contributed by Robbin C. Moran

Olfersia Raddi, Opusc. Sci. 3: 283, /. lib. 1819.

TYPE: Olfersia corcovadensis Raddi = O. cer-

vina (L.) Kunze. Figure 14.

Plants terrestrial or low-climbing. Stem short-

creeping, in cross-section with the meristeles ar-

ranged irregularly and each one not surrounded

by a dark sclerenchymatous sheath. Sterile and

fertile leaves strongly dimorphic. Sterile lamina

pinnate with a conform terminal pinna. Veins

forking near or at the base, parallel, ca. 1 mm

apart, connected by a submarginal vein (this some-

times difficult to see in dried material). Fertile

leaves 2-pinnate, rarely 1 -pinnate. Son exindu-

siate, linear to oblong. Spores monolete, echinu-

late, ca. 50 microns long.

Olfersia is a monotypic genus that occurs from

southern Mexico to southeastern Brazil and in the

Antilles. It has been included in Polybotrya be-

cause of its strongly dimorphic leaves and creep-

ing, scaly stem; however, Olfersia differs in stem

anatomy, venation, and shape of the leaf apex.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

55

FIG. 14. Olfersia cervina: a, stem; b, apex of sterile leaf; c, portion of fertile leaf; d, portion of pinna, abaxial side,

(a from Pabst 4771, Brazil, F, b, c from H. H. Smith 1075, Colombia, F; d from Madison et al, 4616, Ecuador, F.)

56

FIELDIANA: BOTANY

Reference

XV. Polybotrya

MORAN, R. C. 1986. The neotropical fern genus

Olfersia. Amer. Fern. J., 76: 161-178.

1 . Olfersia cervina (L.) Kunze, Flora 7: 3 1 2. 1 824.

Figure 14.

Osmunda cervina L., Sp. pi. 2: 1065. 1753. LECTO-

TYPE (designated by Proctor, Flora Lesser An-

tilles 223. 1977): Plumier, Traite foug. Amer. t.

154. 1705, illustrating a plant from Martinique.

Olfersia corcovadensis Raddi, Opusc. Sci. 3: 283, t.

lib. 1819. TYPE: Brazil, Rio de Janeiro, Mt.

Corcovado, Raddi (holotype, presumably n).

Polybotrya cervina (L.) Kaulf, Enum. fil. 55. 1824.

Stem scales to 2.5 cm long and 0.1 cm broad,

linear, golden-brown, entire. Sterile leaves 0.5-1 .2

m long and 0.3-0.5 m broad, subcoriaceous, gla-

brous. Petiole shorter than the lamina, scaly at

base. Pinnae 15-30 cm long and 3-8 cm broad,

ovate-lanceolate to lanceolate, 4-12 pairs, short-

stalked, entire, the basiscopic side excavate. Ra-

chis and costae not or only faintly grooved, gla-

brous. Fertile leaves produced freely on terrestrial

as well as scandent stems, more erect than the

sterile leaves and taller due to the longer petiole,

soon withering after the spores are shed. Petiole

40-85 cm long. Pinnae 7-20 cm long, subdistant.

Pinnules 0.5-1 cm long and 0.1-0.3 cm broad,

pectinately arranged, broadly adnate and joined

by a narrow wing of tissue.

Shaded forests, ravines, swamps, mostly terres-

trial or on dead logs, only occasionally climbing,

0-2000 m, Amazonas, Huanuco, Cuzco, Madre

de Dios, Puno.

Entirely neotropical, West Indies; southern

Mexico south to Bolivia and southeastern Brazil.

Amazonas: Prov. Bagua, 43 km by road NE of Chi-

riaco, Barbour 4514 (MO). Huanuco: Ridge E of Tingo

Maria, Allard 22301 (GH, us). Cuzco: Prov. La Conven-

tion, Dudley 10324 (GH). Madre de Dios: Pantiacolla,

serrania across Rio Alto Madre de Dios from Shintuya,

Gentry et al. 27366 (MO). Puno: Prov. Carabaya, Vargas

18936 (GH).

Contributed by Robbin C. Moran

Polybotrya Willd., Sp. pi., ed. 4, 5: 99. 1810. TYPE:

Polybotrya osmundacea Willd. Figure 15.

Soromanes Fee, Mem. foug. 2: 16. 1845. TYPE: So-

romanes serratifolium Fee = Polybotrya serrati-

folia (Fee) Klotzsch.

Plants hemiepiphytic or (in 2 species) terrestrial.

Stem 1-3 cm thick, densely scaly, long-creeping

in the hemiepiphytic species, short-creeping in the

terrestrial ones, in cross-section with 4-10 circu-

larly arranged meristeles, each surrounded by a

dark sclerenchymatous sheath. Sterile and fertile

leaves strongly dimorphic. Sterile lamina 1— 4-pin-

nate, the apex pinnatifid or (in P. polybotryoides)

subconform. Veins free or (in 4 of the 1 -pinnate

species) anastomosing. Indusia absent. Sporangial

capsules glabrous or (in P. pubens) setose. Spores

monolete, echinate.

Polybotrya contains 35 species and is entirely

neotropical, occurring from Mexico to southeast-

ern Brazil and the West Indies. Its center of di-

versity is in the Andes, where 23 species occur, 12

of them in Peru, with four more expected there.

The genus is divided into three subgenera, all

of them in Peru. Subgenus Sorbifolia Moran

(1987a), type, P. sorbifolia Kuhn, is represented

in Peru by species 1. P. fractiserialis and 2. P.

crassirhizoma. Subgenus Soromanes (Fee) Moran

(1987a), type, P. serratifolia (Fee) Klotzsch, is rep-

resented in Peru by 3. P. polybotryoides, 4. P. sub-

erecta, and 5. P. andina. The other species, 6-16,

belong to subgenus Polybotrya.

The genus is readily identified by its strong ster-

ile-fertile leaf dimorphy, unique stem cross-sec-

tion, and (in most species) hemiepiphytic habit.

References

MORAN, R. C. 1987a. Monograph of the Neo-

tropical fern genus Polybotrya (Dryopterida-

ceae). Bull. Illinois Nat. Hist. Surv., 34: 1-138.

MORAN, R. C. 1987b. Sterile-fertile leaf dimor-

phy and evolution of soral types in Polybotrya

(Dryopteridaceae). Syst. Bot., 12: 617-628.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

57

58

FIELDIANA: BOTANY

Key to Species of Polybotrya

a. Sterile lamina 1 -pinnate b

b. Veins of sterile pinnae free (subgenus Sorbifolid) b

c. Plants terrestrial; stem scales brown, concolorous; fertile pinnules commonly round or oblong,

usually less than 1 cm long 1 . P. fractiserialis

c. Plants hemiepiphytic; stem scales orange or reddish brown, with a dark central stripe and

lighter borders; fertile pinnules usually more than 1 cm long 2. P. crassirhizoma

b. Veins anastomosing (subgenus Soromanes) d

d. Lamina apex a terminal segment similar to the lateral pinnae 3. P. polybotryoides

d. Lamina apex gradually reduced, evenly pinnatifid, unlike the lateral pinnae e

e. Sterile lamina glabrous or rarely pubescent abaxially, the trichomes 0.1-0.3 mm long,

colorless, erect, 1-3-celled; pinnae 4-5 cm broad, 6-13 pairs 4. P. suberecta

e. Sterile lamina densely pubescent abaxially, the trichomes 0.5-1.2 mm long, tawny, spread-

ing, 5-12-celled; pinnae 5.7-10 cm broad, 4-7 pairs 5. P. andina

a. Sterile lamina 1 -pinnate-pinnatifid to 4-pinnate (subgenus Polybotrya) f

f. Pinnules of medial pinnae anadromous, i.e., the acroscopic pinnule is closest to the rachis . . . .g

g. Stem scales yellowish to dark golden brown, long, tortuous, and intricate; the basal basiscopic

veinlet often arising between the main pinnate groups of veins 6. P. altescandens

g. Stem scales dark brown, rather straight, spreading or ascending; the basal basiscopic veinlet

not arising between the main pinnate groups of veins 7. P. osmundacea

f. Pinnules of medial pinnae catadromous, i.e., the basiscopic pinnule is closest to the rachis . . h

h. Axes pilose, the trichomes 1-2.5 mm long, acicular i

i. Stem scales usually denticulate, translucent, cream-colored to castaneous; lamina broadest

at the base or nearly so, the petiole more than 15 cm long; fertile pinnules caudate, 3-15

mm broad; sporangial capsules setose 8. P. pubens

i. Stem scales entire, opaque, dark brown; lamina base tapered to a short (less than 4 cm long)

petiole; fertile pinnules 1-3 mm broad, botryoid; sporangial capsules glabrous

9. P. glandulosa

h. Axes glabrous or, if pubescent, the trichomes less than 1 mm long j

j. Tertiary segments or lobes of the sterile lamina 0.5-1.5 mm wide and with one vein ....

10. P. lechleriana

j. Tertiary segments or lobes of the sterile lamina more than 2 mm broad and with more than

one vein k

k. Stem scales yellowish or dark golden brown, membranous, translucent, often long, tor-

tuous and intricate, the margins erose to denticulate 1

1 . Lamina pubescent on both surfaces; veins prominulous adaxially

1 1 . P. puberulenta

1 . Lamina glabrous on both surfaces; veins not prominulous m

m. Adaxial groove of costa pubescent within; basal basiscopic veinlet often arising

between the main pinnate groups of veins; pinnae pinnatifid in the distal portion

6. P. altescandens

m. Adaxial groove of costa glabrous within; basal basiscopic veinlet arising from the

costule; pinnae divided nearly to the apex 12. P. aequatoriana

k. Stem scales dull brown, thick, opaque, spreading to curved and ascending, the margins

entire or nearly so n

n. Sterile lamina mostly 2-pinnate-pinnatifid; tertiary segments or lobes entire or nearly

so

o

o. Leaf margins sparsely and minutely ciliate; plants usually growing at 0-600 m

. 13. P. caudata

FIG. 15. Polybotrya fractiserialis: a, habit, fertile and sterile leaf. Polybotrya osmundacea: b, sterile pinna. P.

polybotryoides: c, portion of pinna, abaxial side, (a from Irwin et al. 54784, Surinam, F, b from Schunke V. 5785, F,

c from Moran 3593, Ecuador, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 59

o. Leaf margins glabrous; plants growing at 1000-2100 m 14. P. appressa

Sterile lamina mostly 3-pinnate-pinnatifid; tertiary segments lobed p

p. Basal tertiary segments less than 2.5(-3) times longer than broad

15. P. alfredii

p. Basal tertiary segments 3-6 times longer than broad q

q. Costa evenly puberulent abaxially; groove of the costules on the adaxial side

truncated by the ridges of the costal groove and therefore not decurrent, glabrous

or nearly so within 1 6. P. hickeyi

q. Costa glabrous or with only scattered trichomes abaxially; groove of the costules

on the adaxial side decurrent into the groove of the costa, filled with reddish

to brownish trichomes . . 7. P. osmundacea

1 . Polybotrya fractiserialis (Baker) John Sm., Hist,

fil. 133. 1875. Figure 15a.

Acrostichum fractiseriale Baker, Syn. fil. 414. 1868.

TYPE: Peru, San Martin, Mt. Campana, Spruce

4337 (holotype, K; color slide, MO of K; photos,

GH, US Of K.).

Acrostichum plumbicaule Baker, Syn. fil. 413. 1868.

TYPE: Peru, San Martin, Tarapoto, Spruce 4090

(holotype, K; photo, GH; photo and frag., P!, us!).

Polybotrya plumbicaulis (Baker) John Sm., Hist. fil.

133. 1875.

Plants terrestrial. Stem scales dull brown,

opaque, the margins entire or rarely denticulate.

Sterile lamina 1 -pinnate, glabrous. Pinnae (11-)

15-25(-28) cm long and (2.1-)2.5-4(-4.5) cm

broad. Axes glabrous, or rarely with fine pubes-

cence, the trichomes 0.1-0.2 mm long, adaxial

grooves shallow, glabrous. Veins free. Fertile lam-

ina 2-pinnate. Pinnules 1-3 mm broad, round,

oblong or linear, pectinately arranged.

Wooded talus slopes, usually along rocky

streams, 200-1 500 m, Amazonas, San Martin and

Loreto to Cuzco and Madre de Dios.

The range of P. fractiserialis consists of two

widely separated regions, the eastern slopes of the

Andes from Ecuador and Bolivia, and the Guian-

as.

Amazonas: Prov. Bagua, Valley of Rio Maranon above

Cascadas de Mayasi near Campamento Sta. Montenegro

(Kms. 280-284 of Maranon road), Wurdack 1854 (us).

San Martin: Prov. Lamas, between Tarapoto and Moy-

obamba, ca. 10 km NW ofTabalosas, Croat 51154 (MO).

Loreto: Above Pongo de Manseriche, Mexia 6246a (uc,

us). Aguaytia, Croat 21014 (MO). Huanuco: Prov. Huan-

uco, Tingo Maria, Tryon & Tryon 5221 (GH). Previsto,

antes de Aguaytia, Aguilar 947 (USM). Pasco: Iscozacin,

Foster 7937 (F, USM). Prov. Oxapampa, Palcazu, Rio

Alto Iscozacin, Foster 9996 (F). Junin: Prov. Satipo, Pi-

chanaki, Leon 226 (USM). Chanchamayo Valley, C.

Schunke 165 (F, us). Cuzco: Prov. Paucartambo, Vargas

11280 (GH). Madre de Dios: Prov. Manu, town of Ata-

laya, 2-3 km W of village, Foster 7455 (F, MO, USM).

Pantiacolla, serrania across Rio Alto Madre de Dios from

Shintuya, Gentry et al. 27365 (MO). Valle de Marcapata,

Herrera 1200 (us).

2. Polybotrya crassirhizoma Lellinger, Amer. Fern

J. 62: 49, / 1, 8. 1972. TYPE: Peru, Loreto,

Gamitanacocha, Rio Mazan, /. Schunke 268

(holotype, us!; isotypes, F!, GH!, NY!, uc!).

Polybotrya macbridei Lellinger, Amer. Fern J. 62: 5 1,

/ 2, 9. 1972. TYPE: Peru, Junin, Schunke

Hacienda, La Merced, Macbride 5602 (holotype,

us!; isotype, F!).

Plants hemiepiphytic. Stem scales dull orange

or reddish brown, rarely shiny and dark brown,

with a dark central stripe and lighter borders, the

margins denticulate to erose. Sterile lamina 1 -pin-

nate, lanceolate, glabrous. Pinnae ( 1 1-) 1 5-20(-24)

cm long and 2-4 cm broad, the margins crenate

to dentate. Axes sparsely scaly and pubescent, the

trichomes ca. 0.1 mm long, colorless, subulate,

adaxial grooves glabrous within. Veins free, often

with a single basiscopic veinlet arising from the

costa between the main 1 -pinnate groups. Fertile

lamina 2-pinnate, occasionally 2-pinnate-pinnati-

fid in large leaves. Pinnues 1-3 mm broad, oblong

to linear, pectinately arranged.

Wet forests, 100-1500 m, Loreto, Huanuco,

Pasco, and Junin.

Colombia to Bolivia and western Brazil; most

frequent and abundant in Amazonia, but also on

the eastern slopes of the Andes.

Loreto: Prov. Maynas, Quebrada Yanomona, Explo-

rama tourist camp, Rio Amazonas above the mouth of

60

FIELDIANA: BOTANY

Rio Napo, Moran 3640, 3641 (F, Q, QCA). Huanuco: Prov.

Pachitea, Rio Pozuzo, Foster 9284 (F, MO, USM). Pasco:

Prov. Paucartambo, Cosnipata Valley, Rio Tono, road

N of Patria to first foothill ridge, Foster 10627 (F). Junin:

Chanchamayo Valley, C. Schunke 157, 158 (F). Schunke

Hacienda, above San Ramon, Killip & Smith 24605 (NY,

us).

3. Polybotrya polybotryoides (Baker) Christ, Bull.

Herb. Boissier, ser. 2, 1: 70. 1901. Figure 15c.

Acrostichum polybotryoides Baker, J. Bot. 19: 207.

1881. TYPE: Colombia, Norte de Santander,

Ocana, Kalbreyer 1254 (holotype, K; color slide,

MO of K; photos, GH, MO of K).

Acrostichum juglandifolium Baker, J. Bot. 19: 207.

1881, nom. illeg., not Kaulfuss, 1 824. TYPE: Co-

lombia, Antioquia, Kalbreyer 1798 (holotype, K;

color slide, MO of K.; photos, GH, MO of K).

Polybotrya juglandifolia Christ, Bull. Herb. Boissier,

ser. 2, 4: 965. 1905. nom. nov. tor Acrostichum

juglandifolium Baker, and with the same type.

Polybotrya kalbreyeri C. Chr., Index fil. 504. 1906,

nom. super/1, for Polybotrya juglandifolia Christ.

Plants hemiepiphytic. Stem scales lustrous,

golden yellow when fresh, turning yellow-brown

to purple-brown upon drying, ascending with

spreading tips, the margins denticulate. Sterile

lamina 1 -pinnate, the tissue between the veins gla-

brous on both surfaces. Pinnae 1 2-20(-30) cm long

and 3-6(-8) cm broad, 5-12 pairs, the margins

entire, crenate or broadly serrate. Axes glabrous

or nearly so, adaxial grooves glabrous within. Veins

anastomosing, the tips connected by a faint intra-

marginal connecting strand. Fertile leaves 2-pin-

nate. Pinnules linear, entire or lobed toward the

base.

Wet forests, 0-1400(-1850)m, Huanuco, Pasco.

Southern Mexico to Central Peru.

Huanuco: SW slope of Rio Llullapichis watershed, as-

cent of Cerros del Sira, Dudley 13290D (GH). Pasco:

Prov. Oxapampa, Abra los Mellizos, 4-8 km from Enen-

as, Skog et al. 5036 (us).

Plants hemiepiphytic. Stem scales dark brown,

linear, stiff and ascending to spreading, denticu-

late. Sterile lamina 1 -pinnate, glabrous or rarely

with uncinate trichomes abaxially. Pinnae (10-)1 5-

21(-25) cm long and (3-)4-5(-6.5) cm broad, 6-

13 pairs, the margins entire or shallowly serrate.

Axes glabrous or rarely pubescent, the trichomes

less than 0. 1 mm long, uncinate, the adaxial grooves

glabrous within. Veins anastomosing, the tips oc-

casionally uniting to form a discontinuous sub-

marginal strand. Fertile leaves 2-pinnate to 2-pin-

nate-pinnatifid. Pinnules linear, entire, or lobed at

the base, pectinately arranged.

Wet forests, 600-1800 m, Junin.

Colombia to Peru.

Junin: Prov. Tarma, Chanchamayo, Esposto 10928

(USM).

5. Polybotrya andina C. Chr., Index fil. 504. 1 906,

nom. nov. for Acrostichum insigne Baker, not

Fee, 1872-73, and with the same type.

Acrostichum insigne Baker, J. Bot. 15: 167. 1877, nom.

illeg., not Fee, 1872-1873. TYPE: Ecuador, Pi-

chincha, "Andes of Quito," Sodiro (holotype, K;

photo, GH, us of K; isotypes, AAU!, GH!, Q!, uc!).

Plants hemiepiphytic. Stem scales dark casta-

neous, linear, shiny or dull, opaque, denticulate.

Sterile lamina 1 -pinnate, the apex pinnatifid, pu-

bescent abaxially, the trichomes spreading, tawny,

uncinate, 4-10-celled. Pinnae 17-33 cm long and

5.7-10 cm broad, 4-7 pairs, the margins entire.

Axes pubescent, the adaxial grooves glabrous.

Veins anastomosing. Fertile lamina 2-pinnate.

Pinnules linear, pectinately arranged, entire to lob-

ulate.

Wet forests, 1 000-1 200 m, not known from Peru

but expected to occur there.

Ecuador.

4. Polybotrya suberecta (Baker) C. Chr., Index fil.

506. 1906.

Acrostichum suberectum Baker, J. Bot. 19:207. 1881.

TYPE: Colombia, Antioquia, Kalbreyer 1877

(holotype, K; color slide, MO of K; photos, GH, us

ofK).

Acrostichum hackelianum Sodiro, Crypt Vase. Quit.

491. 1893. TYPE: Ecuador, Pichincha, banks of

the Rio Pilaton, Sodiro (holotype, Q!)-

6. Polybotrya altescandens C. Chr., Index fil. 7.

1905, nom. nov. tor Acrostichum chrysolepis

Sodiro, not Fee, 1 869, and with the same type.

Acrostichum chrysolepis Sodiro, Crypt vase. Quit. 485.

1893, nom. illeg., (not Fee, 1869). TYPE: Ecua-

dor, Pichincha, "ad Pilaton-Toachi," Sodiro (ho-

lotype, P!).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

61

Plants hemiepiphytic. Stem scales yellow to dark

golden brown, linear, mostly concolorous, shiny,

long, tortuous and intricate, denticulate, the base

cordate and darkened around point of attachment.

Sterile lamina to 2-pinnate-pinnatifid or rarely

3-pinnate at the base, lanceolate, glabrous. Pinnae

14_25(-35) cm long and 3-10(-17) cm broad, not

deeply cut, pinnatifid above the base in the distal

portions. Pinnules of the medial pinnae 6-10 cm

long and 1-3.5 cm broad, catadromous, sessile.

Axes pubescent and sparsely scaly abaxially, the

trichomes 0.1-0.2 mm long, erect, the scales scat-

tered, denticulate, adaxial grooves slightly pubes-

cent within. Veins free, with the basal basiscopic

veinlet often arising between the main pinnate

groups of veins. Fertile lamina 3-pinnate. Pinnules

narrowly oblong to linear, pinnate, with the sori

obovate or oblong.

Wet forests, (800-) 1200-2500 m, Loreto, Hua-

nuco.

Colombia; Ecuador; Peru.

Loreto: Prov. Coronel Portillo, Dist. Padre Abad, divi-

soria cerca al Rio Chino, /. Schunke 10200 (MO). Hua-

nuco: La Divisoria, ca. 25 km NE of Tingo Maria, Moran

3688 (F, G, GH, MO, USM).

7. Polybotrya osmundacea Willd., Sp. pi. ed. 4, 5:

99. 1810. TYPE: Venezuela, Monagas, Cari-

pe, Humboldt 459a (fertile), 459b (sterile) (ho-

lotype, B, Herb. Willd. 19507-1, 19507-2;

photos, F, GH). Figure 15b.

Acrostichum osmundaceum (Willd.) Hooker, Sp. fil.

5: 246. 1864.

Plants hemiepiphytic. Stem scales variable, with

plants from Amazonia tending to have thick,

opaque, dark brown, subentire, somewhat squar-

rose scales and plants from elsewhere tending to

have shiny, ascending, spreading scales, often with

a dark central stripe and lighter borders, margins

erose to denticulate. Sterile lamina 2-pinnate-pin-

natifid to 3-pinnate-pinnatifid, deltate, ovate, or

lanceolate, the tissue between the veins glabrous

or rarely glandular, the glands round, sessile, res-

inous, the margins glabrous. Pinnae 20-50 cm long

and 10-25 cm broad, anadromous, deeply cut to

the apex. Pinnules of the medial pinnae to 1 4 cm

long and 4 cm broad, anadromous, or sometimes

in plants from Amazonia, catadromous, the base

prolonged acroscopically, stalked, the stalk 1-5

mm long. Tertiary segments lobed. Axes glabrous

or pubescent abaxially, the trichomes 0.2-0.5 mm

long, whitish, the scales absent or few and incon-

spicuous, adaxial grooves pubescent within. Veins

free. Fertile lamina to 3-pinnate-pinnatifid. Pin-

nules lanceolate, pinnate.

Wet forests, 0-2100 m, San Martin and Loreto

to Cuzco and Madre de Dios.

One of the most common and widespread spe-

cies of Polybotrya, from Guatemala to French Gui-

ana, Bolivia and northern Brazil; Antilles.

San Martin: Camino a Pushurumbo, 7-8 km E del

puente de Palo Blanco, Mariscal Caceres, Tocache Nue-

vo, J. Schunke 5785 (COL, NY, us). Loreto: Prov. Maynas,

Peter Jensen's Explorama Lodge, 50 mi downriver from

Iquitos, Moran 3642 (AMAZ, F, USM). Huanuco: Tingo

Maria, Allard 21609, 21997 (us). Pasco: Pichis Trail,

Yapas, Killip & Smith 25452 (NY, us). Prov. Oxapampa,

Palcazu, Rio Alto Iscozacin, Ozuz, Foster 9994 (F). Ju-

nin: E of Quimiri bridge, near La Merced, Killip & Smith

23991 (F, NY, us). Ucayali: Rio Aguaytia above mouth

of Quebrada Yurac-Yacu, Croat 20851 (MO). Cuzco: Prov.

Paucartambo, arriba de Tono, cerca de Patria, Fernan-

dez et al. 1013 (F). Madre de Dios: Prov. Manu, Vargas

17743 (GH).

8. Polybotrya pubens Mart., Icon. pi. crypt. 87, /.

25. 1834. LECTOTYPE (designated here):

Brazil, ad flum. Amazonas prope Serpa, Mar-

tius (holotype, M!).

Polybotrya pubens Kunze, Linnaea 9: 22. 1 834, not

Martius, 1834. TYPE: Peru, Loreto, Prov. May-

nas, ad Yurimaguas, Poeppig (holotype, P!, one

of the syntypes of P. pubens Mart.).

Polybotrya decorata Lellinger, Amer. Fern J. 62: 54,

/ 5, 6, 11. 1972. TYPE: Peru, Loreto, Iquitos,

Killip & Smith 26955 (holotype, us!; isotype, NY!).

Plants hemiepiphytic. Stem scales variable, on

large mature stems whitish to coffee-brown, con-

colorous, ovate, membranous, denticulate, or on

young or terrestrial stems narrowly lanceolate, cas-

taneous. Sterile lamina mostly 1 -pinnate-pin-

natifid, the base occasionally 2-pinnate-pinnatifid,

pubescent throughout or rarely glabrous adaxially,

the trichomes 1-2 mm long, acicular, whitish. Pin-

nae 9-19 cm long and 2.5-3.5 cm broad, often

with the lowest pinnae having a few free proximal

segments or pinnules. Pinnules or segments catad-

romous, oblong, slightly falcate. Veins free. Axes

conspicuously pubescent, adaxial grooves pubes-

cent. Fertile lamina mostly 1 -pinnate-pinnatifid,

sometimes 2-pinnate at base. Pinnae mostly en-

tire, long-caudate. Sporangial capsules setose.

62

FIELDIANA: BOTANY

Wet forests, primarily in Amazonia, 0-1400 m,

Amazonas, San Martin, Loreto, Pasco, Junin, Ma-

dre de Dios.

Colombia to Bolivia; western Brazil.

This species is characterized by its relatively

uncut lamina, long, dense, whitish pubescence, and

long-caudate fertile pinnae. In addition, it is the

only species of Polybotrya with setose sporangial

capsules. The name Polybotrya pubens was pub-

lished twice in the same year. The name of Martius

is accepted as correct, since Kunze took his name

from a specimen annotated by Martius.

Amazonas: Ridge crest of Quebrada Chuivi (above

Km 278 of Maranon road), valley of Rio Maranon, near

Cascadas de Mayasi, Wurdack 1933 (us). San Martin:

Monte Campana, Spruce 4740 (photo, GH, us). Camino

a Pushurumbo, 7-8 km E del Puente de Palo Blanco,

M ari seal Caceres, Tocache Nuevo, J. Schunke 5789 (NY,

us). Loreto: Above Tamishuyacu, Croat 19771 (AMAZ,

MO). Pasco: Prov. Oxapampa, Iscozacin, Foster 7858 (F,

USM). Junin: E of Quimiri bridge, near La Merced, Killip

& Smith 23196 (NY, us). Puerto Bermudez, Killip & Smith

26543 (NY, us). Madre de Dios: Prov. Manu, Cerro de

Pantiacolla, Rio Palotoa, 10-15 km NNW of Shintuya,

Foster 10821 (F).

9. Polybotrya glandulosa Kuhn, Linnaea 36: 65.

1869. TYPE: Brazil, Amazonas, San Gabriel,

Spruce (holotype, B!).

Polybotrya subelliptica Lellinger, Amer. Fern J. 62:

56. 1972. TYPE: Peru, Loreto, Mishuyacu, near

Iquitos, Klug 1390 (holotype; us!; isotypes, F!,

NY!).

Plants hemiepiphytic. Stem scales dull brown,

thick, opaque, concolorous to bicolorous, curved-

appressed, the margins entire, the base expanded,

the dorsal surface sometimes with a medial groove.

Sterile lamina 2-pinnate-pinnatifid, subelliptic,

gradually tapered to a short, less than 4 cm long,

petiole, pilose on both surfaces. Pinnae 14-16 cm

long and 3-3.5(-4) cm broad, sessile or nearly so.

Pinnules of the medial pinnae 15-20 cm long and

5-7 cm broad, catadromous, crenate or lobed. Axes

pilose, the trichomes 1-2.5 mm long, whitish, acic-

ular, adaxial grooves pilose. Veins free. Fertile

lamina 2-pinnate-pinnatifid, botryoid.

Wet lowland forests, 1 00 m, Loreto.

Southern Venezuela; northeastern Peru; north-

western Brazil.

This species is known from only three collec-

tions (in Peru only from the type of P. subelliptica)

and is apparently endemic to the northeastern

Amazon basin. It is easily recognized by its lam-

ina, which gradually tapers to a short, less than 4

cm long, petiole, and the long-pilose pubescence.

10. Polybotrya lechleriana Mett., Fil. lechl. 1:1:

4. /. /,/ 7-5. 1854. TYPE: Peru, Puno, San

Gaban, Lechler 2156 (holotype, LZ destroyed;

isotypes, B!, K, L!; frag., F!, us!; photo, GH of

K).

Acrostichum lechlerianum (Mett.) Hooker, Sp. fil. 5:

246. 1864, nom. illeg. (not Mettenius, 1856).

Plants hemiepiphytic. Stem scales lanceolate,

cream to dull brown, thin, appressed, the margins

denticulate to entire. Sterile lamina 3-4 pinnate,

lanceolate to elliptic, pubescent on both surfaces

and especially so along the veins. Pinnae (7-) 1 0-

15 cm long and (2-)3-7(-12) cm broad, deeply

and finely cut to the apex. Pinnules of the medial

pinnae 1.5-5 cm long and 1-2 cm broad, sessile

or nearly so. Ultimate segments 0.5-1 mm broad,

single- veined, falcate. Axes pubescent, the junc-

tures often provided with a brown, ovate scale,

adaxial grooves pubescent or glabrous. Fertile

lamina 3-4-pinnate. Pinnules oblong, botryoid.

Wet forests, (1 00-) 1000- 1500 m, San Martin,

Junin, Cuzco, Puno.

Colombia to Bolivia; Guyana (Mt. Roraima).

This species has the most finely divided lamina

of any in Polybotrya. The narrow, 1 -nerved ulti-

mate segments readily distinguish the species.

San Martin: Mt. Guayrapurima, near Tarapoto, Spruce

4744 (L, P, frag., us). Junin: Prov. Satipo, Mapiri, ca. 12

km SW of Chequitavo, Smith 61713 (MO, USM). Cuzco:

Prov. La Convention, Cordillera Vilcabamba, Camp 2.5,

Dudley 10325 (GH). Puno: La Pampa, Rio Tavara, Wat-

kins (us).

11. Polybotrya puberulenta R. C. Moran, Bull.

Illinois Nat. Hist. Surv. 34: 90, / 41. 1987.

TYPE: Ecuador, Napo, Cordillera Oriental,

camino Baeza-Tena, 5 km S de Cosanga, Mo-

ran 3528 (holotype, F!; isotypes, COL!, GH!,

LPB!, MO!, NY!, Q!, OCA!, uc!, us!, YEN!).

Plants hemiepiphytic. Stem scales yellow to sor-

did orange, darker in the center with light yellow

borders, membranous, spreading, erose to dentic-

ulate. Sterile lamina to 2-pinnate-pinnatisect, del-

tate to broadly ovate, puberulent on both surfaces,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

63

the trichomes ca. 0. 1 mm long, spreading. Pinnae

to 34 cm long and 1 8 cm broad, deeply cut to the

apex. Pinnules of the medial pinnae to 10 cm long

and 3 cm broad, catadromous, the basiscopic mar-

gin thickened, decurrent on the costa. Axes pu-

berulent, adaxial grooves pubescent within. Veins

free, prominulous adaxially. Fertile leaves 4-pin-

nate. Pinnules lanceolate, pinnate, botryoid.

Wet forests, 1 100-1 500 m, not known from Peru

but expected to occur there.

Ecuador; Bolivia.

12. Polybotrya aequatoriana R. C. Moran, Bull.

Illinois Nat. Hist. Surv. 34: 94, / 43. 1987.

TYPE: Ecuador, Napo, Cordillera Oriental,

camino Baeza-Tena, 34 km S de Baeza, Mo-

ran 3512 (holotype; F!; isotypes, AAU!, COL!,

GH!, LPB!, MO!, NY!, Q!, QCA!, uc!).

Plants hemiepiphytic. Stem scales yellowish or-

ange, membranous, long, tortuous and intricate,

the margins minutely denticulate. Sterile lamina

2-pinnate-pinnatifid, lanceolate, moderately to

slightly pubescent below and often glandular, the

trichomes 0.1-0.3 mm long, fine, appressed, the

glands resinous, round. Pinnae 25-35 cm long and

1 3-1 8(-20) cm broad, divided nearly to the apex,

prolonged acroscopically, catadromous, the base

stalked, the stalk 1-3 mm long, the margins slight-

ly thickened and lighter colored, often revolute

upon drying. Axes pubescent and scaly below, the

trichomes 0. 1-0.2 mm long, erect, the scales lin-

;ar, flexuous, dark, often appressed, adaxial grooves

pubescent within. Veins free, the basal basiscopic

veinlet arising from the costule. Fertile leaves

3-pinnate. Pinnules linear, pinnate, botryoid.

Cloud forests, 2100-2400 m, not known from

Peru but expected to occur there.

Ecuador; Bolivia.

13. Polybotrya caudata Kunze, Linnaea 9: 23.

1834. TYPE: Peru, Huanuco, Pampayacu,

Poeppig (B!).

Olfersia caudata (Kunze) Kunze, Linnaea 21: 206.

1848.

Acrostichum caudatum (Kunze) Hooker, Sp. fil. 5: 244.

1864, nom. illeg., not Hooker, 1840.

Plants hemiepiphytic. Stem scales dull brown,

opaque, appressed, entire, the base elevated, thick-

ened, curved. Sterile lamina 2-pinnate-pinnatifid

(rarely 3-pinnate at the base), ovate to lanceolate,

the tissue between the veins pubescent or (rarely)

glabrous, the trichomes 0.2-1 .5 mm long, acicular,

the margins minutely and sparsely ciliate. Pinnae

up to 20-45 (-60) cm long and 7-20(-30) cm broad.

Pinnules of medial pinnae 7-15 cm long and 1-3

cm broad, catadromous, the base slightly pro-

longed acroscopically. Tertiary segments or lobes

entire or nearly so. Axes glabrous or pubescent to

various degrees, the trichomes 0.2-1.5 mm long,

acicular, whitish, adaxial grooves usually pubes-

cent within, often densely so at the junctures. Veins

free. Fertile lamina 2-pinnate. Pinnules 0.4-1 cm

broad, linear-caudate, entire or lobed at the base.

Wet forests, 0-1900 m, Amazonas and Loreto

to Cuzco and Puno.

Eastern and southern Mexico to French Guiana,

northern Brazil and Bolivia.

Amazonas: Prov. Bagua, left bank of Rio Maranon

opposite Quebrada Mirana (opposite Km 277 of Ma-

ranon road above Cascadas de Mayasi), Wurdack 201 1

(GH, us, USM). San Martin: San Roque, LI. Williams

7681 (F, us). Loreto: Prov. Maynas, Peter Jensen's Ex-

plorama Lodge, 50 mi downriver from Iquitos at Yano-

mona Ck. Moran 3663 (AMAZ, USM). Huanuco: 10 km S

of Tingo Maria, Stork & Morton 9509 (F, uc, us). Pasco:

Puerto Bermudez, Killip & Smith 26637 (NY, us). Madre

de Dios: Parque Nacional de Manu, Cocha Cashu Bio-

logical Station, M. Foster P-84-42 (F). Prov. Manu, Cerro

de Pantiacolla, Rio Palotoa, 5-10 km NNW of Shintuya,

Palotoa Control Post, Foster et al. 11045 (F). Cuzco:

Prov. La Convention, at Camp Zero, Dudley 11513 (GH,

us). Puno: San Gaban, Lechler 2321, 2329 (B).

14. Polybotrya appressa R. C. Moran, Bull. Il-

linois Nat. Hist. Surv. 34: 94, / 44. 1987.

TYPE: Ecuador, Napo, Cordillera Oriental,

camino Baeza-Tena, 34 km al sur de Baeza,

Moran 3586 (holotype, F!; isotypes, GH!, MO!,

NY!, Q!, QCA!, us!).

Plants hemiepiphytic. Stem scales dull brown,

thick, opaque, entire, appressed or curved and as-

cending. Sterile lamina mostly 2-pinnate-pinnati-

fid, dark green adaxially, lighter below, both sur-

faces and the margins glabrous. Pinnae to 37 cm

long and 18 cm broad. Pinnules of the medial

pinnae catadromous, the base prolonged acroscop-

ically, stalked, the stalk 1-5 mm long. Tertiary

segments entire or nearly so. Axes evenly pubes-

cent abaxially, the trichomes less than 0. 1 mm

long, the scales absent or few and inconspicuous,

64

FIELDIANA: BOTANY

linear to lanceolate, appressed, light brown, ad-

axial grooves sparsely pubescent within. Veins free.

Fertile lamina unknown.

Cloud forests, 1000-2250 m, Cajamarca.

Ecuador and Peru.

Cajamarca: Lower edge of Cutervo National Park, 10-

1 5 km N of San Andres de Cutervo, Gentry et al. 61498

(MO).

15. Polybotrya alfredii Brade, Bradea 1: 12, t. 1,

f. 2. 1969. TYPE: Costa Rica, Prov. San Jose,

Tablazo, Brade & Brade 98 (holotype, HB; iso-

type, NY!).

Plants hemiepiphytic. Stem scales dull brown,

opaque to translucent, spreading, the margins den-

ticulate or more, commonly entire. Sterile lamina

2-pinnate-pinnatifid to 4-pinnate, ovate, the tissue

between the veins glabrous. Pinnae ( 1 5-)20-45 cm

long and 10-22 cm broad, deeply cut to the apex.

Pinnules of the medial pinnae up to 12 cm long

and 5 cm broad, catadromous, the base stalked,

the stalk 1-3 mm long. Tertiary segments lobed.

Axes pubescent abaxially, the trichomes 0.1-0.2

mm long, usually tawny, adaxial grooves pubes-

cent within. Veins free. Fertile leaves 3-pinnate-

pinnatifid. Pinnules lanceolate, pinnate.

Wet forests, 700-1900 m, Huanuco.

Costa Rica to Bolivia.

A thick mucilage often invests the petiole bases

and distal portion of the stem. This mucilage dis-

appears after drying and leaves no vestige of its

former presence.

Huanuco: SW slope of the Rio Lullapichis watershed

on the ascent of Cerro del Sira, Dudley 13258, 18265 A

(GH).

1 6. Polybotrya hickeyi R. C. Moran, Bull. Illinois

Nat. Hist. Surv. 34: 88, / 40. 1987. TYPE:

Bolivia, Cochabamba, Prov. Chapare, road

from Cochabamba to Villa Tunari, Hickey

801 (holotype, GH!; isotype, MU).

Plants hemiepiphytic. Stem scales dark choco-

late-brown, mostly opaque, concolorous, or the

apex with thinner, lighter borders, the margins

denticulate to entire, the base thickened, attached

across its full width. Sterile lamina to 3-pinnate-

pinnatifid, deltate, the abaxial surface sightly pu-

berulent, the trichomes up to 0. 1 mm long, cylin-

drical, appressed. Pinnae to 46 cm long and 25 cm

broad, deeply cut to the apex. Pinnules of the me-

dial pinnae 5-10 cm long and 2-4 cm broad, ca-

tadromous, the base slightly prolonged acroscop-

ically, the basiscopic margins thickened and

decurrent on the costa. Tertiary segments lobed.

Axes puberulent and scaly, the trichomes ca. 0. 1

mm (?) long, erect, the scales appressed, denticu-

late, flexuose, adaxial groove glabrous or nearly so

within, not prominent. Fertile lamina 4-pinnate.

Pinnules lanceolate to oblong, pinnate, botryoid.

Wet forests, 1700-2400 m, Pasco.

Colombia; Peru; Bolivia.

Pasco: Prov. Oxapampa, Canyon de Huanca-

bamba, Leon 622 (GH).

Comments

Polybotrya fulvastrigosa Christ, Bull. Herb. Bois-

sier, ser. 2, 1: 70. 1901. TYPE: Peru, Loreto,

Cerro de Canchahuaya, Huber 1448.

I have not seen the type and am uncertain from

the description to what species this name belongs.

Polybotrya lomarioides Mett., Fil. lechl. 2: 5. 1 858.

TYPE: Peru, Puno, San Gaban, Lechler.

I have not seen the type and am uncertain from

the description if this name applies to a species of

Polybotrya or to another genus.

Polybotrya nutans Kunze, Linnaea 9: 24. 1834.

TYPE: Peru, "Sylvae flor. Peruv. ad Pam-

payaco . . . ," Poeppig, in July 1829 (B!, P!).

The fertile leaf of the type specimen is a Poly-

botrya, but I do not know which species. The ster-

ile leaf of the type specimen is a tree fern, perhaps

a species of Trichipteris.

XVI. Diplazium

Diplazium Sw., J. Bot. (Schrader) 1800 (2): 61.

1 802. TYPE: Diplazium plantagineum (L.) Sw.

(Hemionitis plantaginea Sm., Asplenium

plantagineum L., nom. super jl. for A. plan-

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

65

FIELDIANA: BOTANY

taginifolium L.) = Diplazium plantaginifoli-

um (L.) Urban. Figure 16.

Anisogonium Presl, Tent, pterid. 115. 1836. TYPE:

Anisogonium fraxinifolium (Presl) Presl = Dipla-

zium fraxinifolium Presl.

Diplazium subgenus Anisogonium (Presl) C. Chr., In-

dex fil. xxxi. 1906.

Plants terrestrial, rarely epipetric or epiphytic.

Stem commonly erect, in some species to 1 m tall,

often decumbent, rarely long-creeping, sparsely to

abundantly scaly, with not or scarcely clathrate

scales, and usually bearing many long fibrous roots.

Leaves essentially monomorphic, ca. 25 cm to 3

m long, caespitose to (occasionally) subdistant, not

articulate to the stem. Lamina simple and entire

to 3-pinnate-pinnatifid, glabrous to sparsely pu-

bescent or somewhat scaly on abaxial axes, on

some species bearing a proliferous bud distally on

the rachis, axes adaxially sulcate and usually am-

ply and minutely puberulent within the sulci. Ra-

chis deeply sulcate adaxially, the costae less so but

the edges of the sulci often thin and raised, inter-

rupted and enlarged at the junction with the axis

of the next order, in a few species appearing as

perpendicular, herbaceous wings. Veins free, or in

a few species anastomosing and the areoles with-

out included veinlets. Sori elongate, elliptic to

(more commonly) linear, borne on one or often

both sides of the vein (diplazioid), receptacle

scarcely or slightly raised, lacking paraphyses. In-

dusia commonly very narrow, often delicate and

hyaline, attached along the vein, sometimes shriv-

eling at maturity, rarely fugacious or lacking. Spo-

rangia on stalks with 2 or 3 files of cells, annular

cells (12-) 15-20. Spores more or less ellipsoidal,

monolete, laesura with winglike folds, the surface

smooth, or papillate to echinate.

Diplazium is a pantropical genus of over 300

species, and although a number of regional treat-

ments have appeared in recent years, it is sorely

in need of monographic work. Little has been done

to delineate true relationships of species or infra-

generic groups. The following key, as in previous

floristic works, artificially separates many species

according to degree of laminar dissection. Al-

though this character frequently is helpful in key-

ing out certain taxa, there is evidence to suggest it

is a poor indicator of species relationships. Char-

acters that more accurately note true affinities seem

to be those of venation, indument, and perhaps

soriation.

There is ample evidence of hybridization within

the genus in Peru, especially in the species with

decompound leaves. A number of intermediate

specimens have been examined, many with abor-

tive sporangia.

A relatively small group of species with areolate

venation and unusual stem scales perhaps merits

recognition as subgenus Anisogonium. At least one

species (D. lechleri) is distinguished by otherwise

free veins connected by an inframarginal vein.

Certain species complexes are characterized by

coarse, rigid scales on stem and axes, others have

thin, flaccid ones, and still others entirely lack lam-

inar scales. Minute, 1-2-celled trichomes occur on

rachis and costae in varying frequency and posi-

tion, but this feature is more variable and hence

probably less reliable as a group indicator than are

scales.

Sori borne "back-to-back" on both acroscopic

and basiscopic sides of a vein were termed dipla-

zioid by early authors, and this has become the

trademark of the genus. There are rare examples

of this in Asplenium. These sori appear more com-

monly in some species than others, but the fre-

quency of their occurrence seems not to delineate

groups as well as the color, texture, and shape of

indusia. More critical study of sori and indusia

would be advisable in future studies of the genus.

References

PROCTOR, G. R. 1985. Diplazium, pp. 392-409,

in Ferns of Jamaica. Brit. Mus. (Nat. Hist.).

London.

SMITH, A. R. 1976. Diplazium delitescens and

the Neotropical species of Asplenium sect. Hy-

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

STOLZE, R. G. 1 98 1 . Diplazium, pp. 1 82-1 99, in

Ferns and fern allies of Guatemala, Part II, Poly-

podiaceae. Fieldiana, Bot. n. s., 6.

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

zium, pp. 543-554, in Ferns and allied plants,

Springer- Verlag, New York.

FIG. 16. Diplazium cristatum: a, habit, D. roemerianum: b, apex of lamina. Diplazium striatum: c, base of pinna

segment; d, portion of rachis and pinnae bases, adaxial side, (a from Schunke V. 2733, F, b from Marling et al. 19758,

Ecuador, F; c from /. Schunke 295, F, d from LI. Williams 702, F.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

67

Key to Species of Diplazium

a. Veins copiously anastomosing b

b. Leaves pinnatifid to pinnatisect, often pinnate at base, conspicuously petiolate

29. D. pinnatifidum

b. Leaves simple, subentire, petiole short or lacking 30. D. praestans

a. Veins free or, in D. lechleri, connected only at their tips by a slightly inframarginal vein c

c. Leaves simple, subentire 28. D. plantaginifolium

c. Leaves pinnate to decompound d

d. Pinnae deeply pinnatifid to nearly 3-pinnate, or if subentire to shallowly lobed then conspic-

uously inequilateral at base (cuneate or excavate basiscopically, truncate and strongly produced

and often auriculate acroscopically) e

e. Pinnae, at least several proximal pairs, 1-3-pinnate f

f. Costae essentially glabrous abaxially, or sometimes with spreading trichomes 0.4-0.5

mm long or filiform scales, but never regularly and minutely puberulent g

g. Pinnae and pinnules strongly reflexed; axes flexuous 13. D. flexuosum

g. Pinnae and pinnules spreading or ascending; axes essentially straight (costae occa-

sionally somewhat sinuous near apex) h

h. Scales of axes, especially of costae and costules, frequent to abundant, conspicu-

ously setose i

i. Scales of axes bicolorous, the marginal cells and teeth castaneous to blackish,

the teeth mostly bifid; sori linear, mostly 3-4 mm long; pinnae remote

9. D. remotum

i. Scales of axes concolorous, the marginal teeth not bifid; sori elliptic, less than

2 mm long; pinnae approximate 7. D. moritzianum

h. Scales of the axes, if any, with margins entire (rarely remotely denticulate) . . . . j

j. Indusia thick, bicolorous, dark brown, but blackish near line of attachment

8. D. bicolor

j. Indusia thin, essentially concolorous throughout k

k. Lamina 2-pinnate, pinnules of larger pinnae crenate to lobed, cut halfway

or less to the costules, free portion of segment (if any) commonly as broad

as long 3. D. ambiguum

k. Lamina 2-3-pinnate-pinnatisect, pinnules of larger pinnae cut % or more to

the costule (or tertiary segments free and again pinnatifid), free portion of

segment 1 .5-4 times as long as broad 1

1. Costules conspicuously alate adaxially, the wings often interrupted at

segment bases and expanded into cristate lamellae; indusia margins erose-

ciliolate 5. D. buchtienii

1 . Costules not or slightly alate, the wings (if any) not or scarcely expanded;

indusia subentire (in D. hians irregularly splitting at maturity) m

m. Indusia tumid (inflated); veins commonly 4 or 5 pairs per ultimate

segment; free portion of ultimate segments about 1.5 times as long

as broad 1 . D. hians

m. Indusia flat; veins 6-8 pairs in larger segments; free portion of ulti-

mate segments 2-4 times as long as broad n

n. Free portion of ultimate segments twice as long as broad; costae

and costules either moderately scaly or slightly pubescent with

spreading trichomes about 0.4 mm. long 2. D. divergens

n. Free porton of ultimate segments 3—4 times as long as broad;

costae and costules completely lacking indument . 6. D. vastum

f. Costae moderately to abundantly puberulent on abaxial side with minute trichomes,

these 0. 1-0.2 (0.3) mm long and sometimes partly obscured by scurf o

o. Tissue between the veins on abaxial surface minutely puberulent 10. D. expansum

o. Tissue between the veins glabrous p

68 FIELDIANA: BOTANY

p. Lamina (2-)3-pinnate-pinnatisect; sori less than 2 mm long; pinnules 1.5-4(-5)

cm long 4. D. alienum

p. Lamina 2-pinnate (pinnules entire to deeply lobed); sori 4-10 mm long; larger

pinnules (4-)5-12 cm long q

q. Pinnae with 12-16 pairs of pinnules; larger pinnules 4-6 times as long as broad,

entire or crenate-serrate; indusia dark brown with a blackish base

11. D. venulosum

q. Pinnae with less than 8 free pairs of pinnules; larger pinnules 3-3.5 times as

long as broad, most of them deeply lobed to pinnatifid (sometimes subentire);

indusia light brown 1 2. D. tungurahuae

e. Pinnae subentire to pinnatisect (rarely fully pinnate at the base of basal pinnae) r

r. Pinnae commonly truncate and subequilateral at base, or sometimes more strongly pro-

duced basiscopically s

s. Proliferous buds commonly in the axils of distal pinnae; proximal pinnae deltoid or

subdeltoid, larger ones 9-22 cm broad, their ultimate segments with attenuate or

acuminate apices 14. D. macrophyllum

s. Proliferous buds lacking; proximal pinnae ovate, elliptic or oblong (rarely deltoid-

lanceolate), larger ones 3-5.5(-7) cm broad, their ultimate segments obtuse to acute

t

t. Ultimate segments of proximal pinnae narrowly acute, their margins crenate to

shallowly lobed; mature leaves 75 cm long and 22 cm broad

15. D. subobtusum

t. Ultimate segments obtuse, occasionally subacute, their margins subentire to ser-

rulate; mature leaves 1-2 m long and 30-50 cm broad u

u. Indusia vestigial or lacking; sori mostly 1.5-2.5 mm long; lamina chartaceous

to subcoriaceous; ultimate segments of larger pinnae more than 20 pairs ....

16. D. lindbergii

u. Indusia persistent; sori mostly 3-7 mm long; lamina firm-membranaceous;

ultimate segments of pinnae 15 pairs or fewer 17. D. stria turn

r. Pinnae (most of them) conspicuously inequilateral at base, cuneate or excavate basiscopi-

cally, truncate and much more strongly produced (often auriculate) acroscopically . . v

v. Pinna base attenuate or excavate basiscopically, opposed acroscopically by a greatly

elongated auricle; basiscopic pinna margin subentire, crenate, or shallowly lobed

(sometimes pinnatifid on the basal pinnae) 21. D. bombonasae

v. Pinna base cuneate to shallowly or deeply lobed basiscopically, opposed acroscopically

by a slightly enlarged basal lobe; basiscopic margins of (at least) proximal pinnae

deeply lobed to pinnatisect w

w. Rachis bearing a proliferous bud on the adaxial side towards the apex; free pinnae

16-20 pairs; proximal pinnae subequilateral at base 18. D. caracasanum

w. Rachis lacking proliferous buds; free pinnae 8-15 pairs; proximal pinnae (and

others) conspicuously inequilateral x

x. Proximal pinnae incised nearly to costa, their basal acroscopic segments free

and usually short-stalked; indusium usually with a very narrow, whitish margin;

scales of stem apex somewhat flaccid, dull medium-brown, 3—4 mm long . .

19. D. stuebelianum

x. Proximal pinnae lobed % to % to the costa, their ultimate segments never free;

indusium concolorous; scales of stem apex coarse, castaneous to blackish, often

sublustrous, 1-3 mm long 20. D. cristatum

d. Pinnae entire to crenate, or lobed V4 or less to costa, essentially subequilateral at base y

y. Lamina gradually reduced to a pinnatifid apex z

z. Pinnatifid apical portion of leaf about !/2 the length of the lamina; free pinnae 2-4 pairs

25. D. paucijugum

z. Pinnatifid apical portion of leaf '/5 (or commonly less than) the length of the lamina; free

pinnae on mature leaves 7-25 pairs aa

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 69

y.

aa. Rachis and costae sparsely to moderately provided on abaxial side with appressed

(often filiform) scales; a proliferous bud often borne distally on the rachis

24. D. celtidifolium

aa. Rachis and costae lacking scales; proliferous bud lacking bb

bb. Pinnae 11-17 pairs, most of them lobed about L4 to the costa, acute at apex,

broadest near the center 22. D. cuneifolium

bb. Pinnae commonly less than 10 pairs, with entire to shallowly crenate margins

and acuminate apex, broadest at or near the base 23. D. grandifolium

Lamina terminating abruptly in a conform or subconform apical segment (or rarely non-

conform: with 1 or 2 basal lobes) cc

cc. Veins 1-2 forked, not connected by an inframarginal vein; sori arching, the indusia

thin, mostly borne on one side of a vein 26. D. roemerianum

cc. Veins simple, or paired at the costa, connected at their tips by a slightly inframarginal

vein; sori straight, the indusia firm, rigid, mostly double 27. D. lechleri

1. Diplazium hians Klotzsch, Linnaea 20: 361.

1847. TYPE: Venezuela, Merida, Moritz 289,

in part (holotype, B, frag., us!; isotypes, BM!,

K!, frags., us! of BM & K; photos, us of BM &

K).

Plants terrestrial, rarely on bases of tree trunks.

Stem erect, this and the very base of the petiole

provided with blackish, lanceolate scales to 1.5

cm long. Leaves to 2 m long, 2-pinnate-pinnatifid.

Lamina chartaceous or subcoriaceous, to 80 cm

broad, glabrous abaxially, the costae and costules

moderately to abundantly scaly. Pinnae contigu-

ous, patent or ascending, short-stalked or sessile,

costae on abaxial side with scales orange to cas-

taneous, filiform to lanceolate or ovate and atten-

uate. Pinnules moderately to deeply lobed, cos-

tules adaxially with parallel, cartilaginous ribs,

abaxially scaly as on the costae. Ultimate segments

8-10 pairs, obtuse, subentire, the free portion about

1.5 times as long as broad. Veins free, commonly

4 or 5 pairs on a segment. Sori elliptic, all except

basal ones less than 2 mm long. Indusia thin, gray-

brown, tumid (inflated) and subentire until ma-

turity, then usually irregularly splitting, with only

a narrow portion persistent.

In forests, 1200-3100 m, San Martin, Pasco,

Junin.

Venezuela; Colombia; Ecuador; Peru; Brazil.

This species has been poorly understood— it is

often a name under which several species have

been placed. The type collection is a mixture; Mo-

ritz 289 at Paris is a new species called D. mor-

itzianum from Colonia Tovar, Edo. Aragua; the

other isotypes are from Edo. Merida. With D. hians

probably should be included D. bogotense (Kar-

sten) Hieron. of Colombia, and perhaps also D.

altissimum (Jenm.) C. Chr. from the Greater An-

tilles. The species included as D. hians in Proctor's

"Ferns of Jamaica" (1985, p. 405) is something

else, for the minor axes are described as puberu-

lous, with scales few or lacking. Furthermore, Field

Museum specimens from the Greater Antilles that

are annotated as D. hians have linear, persistent

indusia (not elliptic, tumid, and evanescent). Taxa

allied with this species are in great need of revision.

Also see D. divergens for further discussion.

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

& Young 2164 (F, uc, USM). Pasco: Prov. Oxapampa,

Dist. Oxapampa, Rio Alberto, Leon 649 (USM). Prov.

Oxapampa, 4-5 km N of Mallampampa, D. Smith &

Canne 5804 (MO, uc). Prov. Oxapampa, San Alberto,

Cordillera de Yanachaga, van der Werffet al. 8479 (MO,

uc). Junin: Carpapata, above Huacapistana, Killip &

Smith 24462 (GH, us). Chanchamayo Valley, C. Schunke

144, 147, 166 (F).

2. Diplazium divergens Rosenst., Repert. Spec.

Nov. Regni Veg. 12: 471. 1913. TYPE: Bo-

livia, Yungas, Polo-Polo, near Coroico, Buch-

tien 3393 (holotype, B?; isotypes GH!, P!, us!;

photos, F & us of P).

Plants terrestrial. Stem erect, this and the petiole

base sparsely scaly. Leaves to 2 m long, 2-pinnate-

pinnatifid. Lamina firm-membranaceous, to 80 cm

broad, the costae and costules moderately scaly,

the scales orange, filiform to linear and attenuate,

glabrous, or rarely the costae with scattered

spreading trichomes about 0.4 mm long. Pinnae

contiguous to subdistant, proximal ones conspic-

uously petiolate. Pinnules deeply pinnatisect. Ul-

70

FIELDIANA: BOTANY

timate segments 12-15 pairs, obtuse, free portion

twice as long as broad. Veins free, commonly 6-

8 pairs on a segment. Sori linear, most of them

1.5-2.5 mm long. Indusia thin, gray-brown, flat,

subentire.

In forests, 1000-2100 m, Huanuco, Pasco, Ju-

nin.

Peru and Bolivia.

This rare species is closely related to, and often

confused with, D. hians, but the lamina is thinner

in texture, ultimate segments are more numerous

and relatively narrower, and the indusia are linear

and flat, not elliptic and tumid. The Bryan and

Smith specimens cited below are intermediate be-

tween the two in most characters. They have abor-

tive sporangia and are apparently hybrids. The

Macbride collection seems to be a hybrid involv-

ing D. divergens and a pubescent species, as it has

few scales on the axes, but has some spreading

trichomes on the costae abaxially, and also has

many abortive sporangia.

Hunuco: Muna, Bryan 550 (F, us), Macbride 4036 (F,

GH, us). Junin: Chanchamayo Valley, C. Schunke 44,

708 (F). Pasco: Prov. Oxapampa, Rio El Tunqui, D.

Smith et al. 1721 (F, MO).

3. Diplazium ambiguum Raddi, Opusc. Sci. 3: 292.

1819. TYPE: Brazil, Mandiocca, Raddi (ho-

lotype, PI?; isotype, FI; photo, us of FI).

Athyrium ambiguum (Raddi) Milde, Bot. Zeit. (Berlin)

28: 350. 1870.

Diplazium ambiguum var. pubescens Rosenst., Hed-

wigia 46: 108. 1906. TYPE: Brazil, Estado Santa

Catharina, Blumenau, Passo Mansa, Haerchen 96

(holotype, s; possible isotype, us).

Plants terrestrial. Stem erect, this and the petiole

base scaly, the scales firm, dark brown to blackish,

lanceolate or linear-lanceolate, the margins sub-

entire. Leaves to 1.4 m long, 2-pinnate. Lamina

firm-herbaceous to chartaceous, to 70 cm broad,

abaxially glabrous, or the costae and costules

sparsely to moderately pubescent with spreading,

articulate trichomes, these mostly 0.4-0.5 mm long.

Pinnae approximate, or proximal ones subdistant,

patent or slightly ascending, stalked, costae sparse-

ly scaly (rarely naked), the scales light brown, ovate

and attenuate to filiform, their margins entire. Pin-

nules sessile, often partially adnate, crenate or lobed

about halfway to the costule, costules abaxially

scaly as on the costae, adaxially with low, parallel,

herbaceous or cartilaginous wings, these some-

times interrupted and expanded near segment bas-

es, but not produced into conspicuous lamellae.

Ultimate segments (if any) with free portion com-

monly broader than long. Veins free, 3-6 pairs to

a segment. Sori linear, commonly 3-5 mm long.

Indusia linear, persistent, thin, light or dark brown,

flat, the margins subentire or, very rarely, erose-

ciliolate.

In forests, 1 50-2400 m, Amazonas and Loreto

to Ayacucho.

Venezuela and Colombia, south to Brazil and

Paraguay.

Diplazium ambiguum is a fitting name for the

specimens assigned to this species. There is some

question that it is correctly applied, since the type

has not been examined. However, all specimens

so determined from Peru match the simple orig-

inal description and the excellent set of six isotype

photos at us. It is distinguished from the other

decompound species in that leaves are essentially

2-pinnate. The pinnules of larger pinnae are ses-

sile, many of them at least partly adnate, and are

crenate or dissected halfway or less to the costule.

Other decompound species are at least deeply

2-pinnate-pinnatisect and either glabrous on the

axes abaxially or regularly and minutely puberu-

lent, with stout, rigid 1-3-celled trichomes mostly

0.1-0.2 mm long. Most specimens of D. ambig-

uum are glabrous abaxially, but occasionally the

costae and costules are sparsely to moderately pu-

bescent with spreading, delicate, multicellular tri-

chomes about 0.5 mm long.

This species is most closely related to D. dipla-

zioides (Klotzsch & Karsten) Alston, of northern

South America, Central America, and the Lesser

Antilles, but the latter differs in that the pinnules

are cut nearly or quite to the costule, the sori are

much shorter, the indusia are always erose-cilio-

late, and the axes are never abaxially pubescent.

Diplazium ambiguum further lives up to its name

in the characters of indusia and indument. Mar-

gins of the indusia are predominantly entire but

occasionally are erose-ciliolate, as in D. dipla-

zioides. Axes are most commonly glabrous abax-

ially, but sometimes they are pubescent with

spreading, multicellular trichomes. (On Spruce

4344 from Monte Campana, San Martin, axes of

different leaves vary from glabrous to pubescent.)

Further uncertainty is introduced by Rosen-

stock's D. ambiguum var. pubescens, which was

said to differ from the typical in the glandular-

puberulent axes and fimbriate indusia. Although

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

71

the type has not been seen, a topotype, Haerchen

35a (F), does have erose-ciliolate indusia, but the

axes are glabrous abaxially. Costae are puberulent

adaxially, which perhaps is what Rosenstock was

referring to, but this condition is quite common

throughout the genus, so the character is of little

importance.

Obviously this species and its allies are very

much in need of further study, throughout their

range.

Amazonas: Mouth of Rio Santiago, Tessmann 4390

(us). San Martin: Prov. Mariscal Caceres, Dist. Tocache

Nuevo, Quebrada Saule Chico, /. Schunke V. 4362 (F,

GH, us). Monte Campana, near Tarapoto, Spruce" 4 344"

(BM, BR, K, w). Mt. Guayrapurima, Spruce "4344" (p).

Loreto: Prov. Maynas, Yanamono Tourist Camp, van

der Werffet al 9855, 9868 (MO). Pasco: Oxapampa, 4-

5 km N of Mallampampa, D. Smith & Canne 5801 (MO).

Junin: Prov. Chanchamayo, Chilpes, D. Smith & Pa-

lacios 2637 (F, MO). Ayacucho: "Aina" (Ayna), between

Huanta and Rio Apurimac, Killip & Smith 22727 (us).

4. Diplazium alienum (Mett.) Hieron., Hedwigia

59: 336. 1918.

Asplenium alienum Mett., Fil. lechl. 2: 1 8. 1 859. LEC-

TOTYPE (designated here): Peru (Puno), San

"Gavan" (Gaban), Lechler 2320 (B!; ISOLEC-

TOTYPE, P!; photos, BM & F of B, GH & us of P.).

PARATYPE: Lechler 2173 (B!).

Asplenium fuscopubescens Hooker, Sp. fil. 3: 264. 1860.

LECTOTYPE (designated here): Peru, (San Mar-

tin), Monte Campana, near Tarapoto, Spruce 4759

(K!; ISOLECTOTYPES, BM!, P!; photos, GH & us

of BM). Schlim 69 of Colombia, also cited by

Hooker, is Diplazium gracilescens.

Diplazium fuscopubescens (Hooker) Moore, Index fil.

329. 1861.

Diplazium lehmannii Hieron., Bot. Jahrb. Syst. 34:

458. 1904. TYPE: Colombia, "prope Ricaurte ad

rivulum Cuaiquer," Lehmann 5064 (holotype, B;

frag., BM!; isotypes, F!, us!).

Plants terrestrial, or occasionally growing on the

base of tree trunks. Stem erect, this and the petiole

base provided with a few broad, brown scales.

Leaves to 2 m long, 2-3-pinnate-pinnatisect. Lam-

ina firm-membranaceous, to 50 cm broad, the cos-

tae, and often the costules and rachis, moderately

to abundantly puberulent abaxially, the trichomes

spreading, septate, 0.1-0.3 mm long. Pinnae trun-

cate and subequilateral at the base. Pinnules 1 0-

20 pairs on larger pinnae, 1.5-4(-5) cm long, the

costules adaxially alate, the wings perpendicular

to the lamina and often expanded into crestlike

lamellae. Ultimate segments 8-12 pairs, those of

larger pinnae usually free, lobed to pinnatisect.

Veins free. Sori less than 2 mm long, usually single,

occasionally diplazioid. Indusia light or (more

commonly) dark brown, subentire to lightly erose

or ciliolate, persistent.

In dense, wet forests and wooded valleys, 700-

2700 m, San Martin to Cuzco and Puno.

Colombia; Ecuador; Peru; Bolivia.

Hooker and Baker (Syn. fil. 242. 1867) treated

this as conspecific with Asplenium sandwichianum

(= Diplazium sandwichianum (Presl) Diels) of Ha-

waii. The type of the latter has not been examined,

but herbarium specimens so determined are less

dissected and puberulent, and costules are scarce-

ly winged. The disjunction is also unlikely.

Diplazium buchtienii is possibly synonymous;

see treatment of the latter for further discussion.

A distinctive feature of this species and D. alienum

is the conspicuous wings on the adaxial side of the

costules, which are membranaceous and borne

perpendicular to the lamina surface. Often these

are expanded, near the bases of tertiary segments,

into crestlike lamellae similar to those in certain

species groups of Hymenophyllum.

Huanuco: Slope of Rio Llullapichis watershed, ascent

of Cerros del Sira, Dudley 13252 (GH), 13253 (MO, us).

Pasco: Prov. Oxapampa, Palcazu Valley, Rio San Jose,

D. Smith 3949 (MO, uc). Junin: Prov. Satipo, Pichanaki,

Leon 230 (USM). Cuzco: La Convencion, Valle San Mi-

guel, Bues 2147 (GH, us). Prov. La Convencion, Cordil-

lera Vilcabamba, Dudley 10326B (GH). Madre de Dios:

Prov. Manu, Cerro de Pantiacolla, Foster et al. 10702

(F).

5. Diplazium buchtienii Rosenst., Repert. Spec.

Nov. Regni Veg. 6:312.1 909. TYPE: Bolivia,

San Carlos, near Mapiri, Buchtien 1136 (ho-

lotype, B?; isotype, us!).

Plants terrestrial. Stem erect, this and the very

base of the petiole provided with broad, firm, cas-

taneous to blackish scales. Leaves 60-1 50 cm long,

2-3-pinnate-pinnatisect. Lamina firm-membra-

naceous to chartaceous, to 40 cm broad, essentially

glabrous abaxially, scales lacking, or rare and scat-

tered. Pinnae approximate, slightly ascending, ses-

sile, or proximal ones short-stalked. Pinnules 1 0-

1 6 pairs per pinna, perpendicular to the costa, the

costules adaxially alate, the wings often interrupt-

ed at segment bases and there expanded into crest-

like lamellae. Tertiary segments 8-15 pairs, entire

and adnate to free and again pinnatifid. Veins free,

4(5) pairs on a segment, or less. Sori elliptic, less

72

FIELDIANA: BOTANY

than 2 mm long. Indusia light or dark brown, thin,

erose-ciliolate.

In wet forests, 1200-2500 m, Huanuco to Cuz-

co.

Peru; Bolivia.

This species probably should be included with

D. alienum, as it seems to differ from the latter

primarily in its glabrous, rather than regularly pu-

berulent, axes on the abaxial side. Although the

occurrence of laminar trichomes and/or scales is

presumed to be a strong diagnostic character in

Diplazium, it seems to be less consistent in some

species groups. Monographic study is needed.

Huanuco: Prov. Huanuco, km 452, Lima-Tingo Ma-

ria road, Young & Sullivan 557 (MO, uc). Pasco: Prov.

Oxapampa, San Alberto, Cordillera de Yanachanga, van

der Werff et al. 8456 (MO, uc). Junin: Chanchamayo

Valley, C. Schunke 49, 143, 171 (F). Cuzco: Prov. Pau-

cartambo, Valle de Pillahuata, Herrera 1612 (us).

In dense forests, 500-1800 m, San Martin, Ju-

nin, Cuzco.

Colombia; Ecuador; Peru.

Among the decompound species of the genus,

this species is marked by its naked lamina, which

is dark green adaxially and quite pale green abax-

ially, the stramineous axes, the stalked, subdistant

pinnae and pinnules, and the ascending, com-

monly obtuse, ultimate segments with parallel

sides.

Peyton & King 1422 from Prov. La Convention

in Cuzco (GH, MO) is a hybrid involving D. vastum.

The lamina has the typical light green color abax-

ially, and the pinnules have numerous, narrow,

ascending segments, but the axes are amply pro-

vided with attenuate, spreading scales, and the in-

dusia are short and elliptic. The sporangia are all

abortive.

Junin: Prov. Tarma, Chanchamayo, Esposto

(Cat. No. 11011, USM), C. Schunke 79, 447, 843

(F, us). Cuzco: Prov. Paucartambo, Hacienda Villa

Carmen, Vargas 14681 (GH).

6. Diplazium vastum (Mett.) Diels, Nat. Pflanzen-

fam. 1(4): 228. 1899.

Asplenium vastum Mett., Ann. Sci. Nat. Bot., ser. 5,

2: 237. 1864. TYPE: Colombia, Bogota, Alto Tri-

go, Lindig 349 (holotype, B, frag., us!; iso types,

BM, K!, P!; photos, F, GH, us of P).

Diplazium tarapotense Rosenst., Repert. Spec. Nov.

Regni Veg. 7: 295. 1909. TYPE: Peru, San Mar-

tin, Tarapoto, Mt. Campana, Spruce 4682 (ho-

lotype, P!; isotypes, BM!, GH!, K!; frag., us! of BM;

photos, GH & us of P).

Plants terrestrial. Stem not seen, probably erect.

Leaves huge, probably to 2 m long, 2-pinnate-

pinnatifid, occasionally 3-pinnate as to base of

larger pinnae. Lamina firm-membranaceous to

chartaceous, to 80 cm broad, pale green and gla-

brous abaxially, lacking scales, dark green adaxi-

ally. Pinnae subdistant, somewhat ascending, long-

stalked. Pinnules incised nearly or quite to the

costule, numerous, subdistant, proximal ones

short-stalked, costules adaxially with low, thick-

ened wings, these often interrupted and slightly

expanded near segment bases, but not produced

into lamellae. Ultimate segments 10—15 pairs, nar-

row-oblong, with parallel sides, commonly obtuse,

ascending, entire or occasionally crenulate, larger

ones 3—4 times as long as broad. Veins free, 6-8

pairs on larger segments. Sori linear, 2-3 mm long.

Indusia thin, light brown, subentire, persistent.

7. Diplazium moritzianum Stolze, sp. nov.

Squamae caulis nigellae, ovato-lanceolatae, margini-

bus integris vel denticulatis; folium usque ad 2 m Ion-

gum, 2-pinnato-pinnatisectum; lamina in pagina abax-

iali glabra, axibus moderate vel abundanter paleaceis;

pinnae contiguae, sessiles vel petiolulatae; costao et cos-

tulae squamis fuscis vel castaneis, dentatis vel setosis;

segmenta ultima 6-10-jugata, obtusa, saepe crenato-ser-

rata; venae liberae, 3-5-jugatae; sori elliptic!, minus quam

2 mm longi; indusium tenue, plerumque planum, eroso-

ciliatum.

Plants terrestrial. Stem erect to ascending, this

and the petiole base scaly, the scales coarse, black-

ish, sublustrous, ovate-lanceolate, the margins en-

tire to remotely denticulate. Leaves to 2 m long,

2-pinnate-pinnatisect. Lamina chartaceous, to 70

cm broad, abaxially glabrous, the axes moderately

to abundantly scaly. Pinnae approximate, patent

or ascending, sessile, or proximal one stalked, cos-

tae with scales dark brown or castaneous, narrow,

attenuate, the margins conspicuously dentate to

setose. Pinnules deeply incised, often nearly to the

costule, costules abaxially scaly as on the costae,

adaxially with low herbaceous parallel wings, these

often interrupted and expanded near segment bas-

es, but not produced into conspicuous lamellae.

Ultimate segments 6-10 pairs, obtuse, larger ones

crenate-serrate and about twice as long as broad.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

73

Veins free, 3-5 pairs to a segment. Sori broadly

elliptic, less than 2 mm long. Indusia persistent,

thin, light or dark brown, flat or occasionally

slightly tumid, the margins erose-ciliolate.

TYPE— Venezuela, Colonia Tovar, Merit z 289,

in part (holotype, P!, 2 sheets; frag., F!; photos, F

& GH of P).

Montane rain forests, 2600-3100 m, San Mar-

tin, Huanuco, Pasco.

Venezuela; Peru; southeastern Brazil (Lanna

1708 [F]).

This has the same type number as D. hians, but

type specimens of the latter are designated as Me-

rida, "Colombia" [Venezuela], whereas the two

sheets at Paris are designated as Colonia Tovar,

which is in Edo. Aragua. The Paris specimens rep-

resent a very different species, with scales on axes

conspicuously dentate to setose, the ultimate seg-

ments relatively narrower and usually crenate-ser-

rate, and the indusia commonly flat, with margins

erose-ciliolate. In comparison, D. hians has scales

with entire margins, ultimate segments broader

and subentire, and indusia strongly tumid.

Diplazium moritzianum is more closely related

to D. pedatum Klotzsch of Venezuela, which also

has indusia with fimbriate margins. Among other

characters, the latter differs from D. moritzianum

in the very narrow, often subulate and entire rhi-

zome and petiole scales.

San Martin: Prov. Mariscal Caceres, near Mirador,

Rio Abiseo National Park, Leon 2164 (F, USM). Huanuco:

Carpish Divide, Sandeman 5117 (K). Pasco: Prov. Oxa-

pampa, Rio San Alberto Valley, D. Smith & Pretel8033

(GH, MO).

8. Diplazium bicolor Stolze, sp. nov.

Folium 2-pinnato-pinnatifidum, usque ad 1.5 m Ion-

gum; lamina glabra, squamis absentibus; pinnae subdis-

tantes, petiolulatae; pinnulae numerosae, subdistantes,

ad basin truncatae, ad apicem attenuatae; segmenta ul-

tima lata, obtusa; venae liberae, 4-5-jugatae; sori linea-

res; indusia crassa, integra, bicoloria, fusca et nigella.

Plants terrestrial. Stem erect to decumbent, pro-

vided with stout, appressed, ovate-lanceolate

scales, these blackish brown, to 1.5 cm long, with

margins entire. Leaves to 1.5 m long, 2-pinnate-

pinnatifid, petiole essentially glabrous and not

scaly. Lamina chartaceous, to 70 cm broad, gla-

brous except for sparse, minute puberulence in the

sulci of axes adaxially. Pinnae subdistant, some-

what ascending, stalked. Pinnules pinnatifid, nu-

merous, subdistant, gradually tapering from a

truncate base to a pinnatifid, attenuate apex, cos-

tules adaxially with low, herbaceous wings per-

pendicular to the plane of the lamina. Ultimate

segments about 1 0 pairs, obtuse, the free portion

often nearly as broad as long. Veins free, 4-5 pairs

on a segment. Sori linear, commonly extending

from the midrib nearly to the segment margin, at

least the basal acroscopic ones diplazioid. Indusia

thick, the margins entire, bicolorous, essentially

dark brown, but blackish near the line of attach-

ment, commonly persistent.

TYPE— Peru, Pasco, Prov. Oxapampa, Dist.

Oxapampa, Rio San Alberto, Leon 643 (holotype,

F!; iso types, GH!, USM).

Wet forests, 1200-2000 m, Pasco, Junin.

Peru; Bolivia (Beck 3059 [F]).

This is one of a few species in Peru, including

D. celtidifolium, with thick, bicolorous indusia.

Indusia in the genus are typically thin-textured and

delicate, and may be light to dark brown, whereas

in D. bicolor they are thicker, quite firm, and dark

brown, becoming blackish in a line along the point

of attachment to the vein. Also characteristic is

the "herringbone" soral pattern. The four or five

pairs of sori are straight, ascending and crowded,

and nearly fill the space between midrib and seg-

ment margin.

Junin: Chanchamayo Valley, C. Schunke 662 "A"' (F).

9. Diplazium remotum Fee, Crypt, vase. Bresil, 1 :

8l,t.24,f. 1. 1869. TYPE: Brazil, Glaziou

2332 (holotype, P).

Plants terrestrial. Stem erect, to 50 cm long,

stout, subarborescent, provided with coarse, dark

brown to blackish scales to 1 cm long. Leaves to

2 m long, 2-pinnate-pinnatifid. Lamina charta-

ceous, to 70 cm broad, glabrous, or axes minutely

and sparsely puberulent abaxially, axes also mod-

erately to abundantly scaly, the scales mostly bi-

colorous, the main body light brown, the margins

castaneous to blackish and conspicuously dentate,

the dark, elongated, teeth commonly bifid. Pinnae

remote, stalked, patent to slightly ascending. Pin-

nules 8-12 free pairs (on larger pinnae), deeply

incised. Ultimate segments 8-12 pairs, mostly ob-

tuse and entire, or the apices crenulate-serrate.

Veins free. Sori mostly 3—4 mm long and single,

or those on basal veins diplazioid. Indusia linear,

74

FIELDIANA: BOTANY

bicolorous, brown or gray-brown, but blackish near

point of attachment, firm, persistent, margins sub-

entire or slightly erose.

In dense, wet forests, often on stream banks,

1 700-3000 m, Amazonas to Cuzco.

Venezuela; Peru; Brazil.

The scales of the axes are most distinctive. They

are the same as those of the putative areolate sub-

genus A nisogonium, i.e., with dark brown to black-

ish margins contrasting with the lighter brown of

the main body of the scale, and elongated, setalike,

bifid, marginal teeth. The type has not been ex-

amined, and the protologue mentions nothing of

these singular scales. However, Fee's illustration

does indicate the abundantly paleaceous axes, and

an enlarged drawing of a scale depicts the long,

marginal, apparently dark, teeth. Furthermore, Dr.

Badre of the Laboratoire de Phanerogamic at Paris

has kindly examined the holotype and confirmed

the presence of these scales.

Probably conspecific is D. rostratum Fee, of Bra-

zil, described in the same publication and illus-

trated as Figure 2 of the same plate. The former

is described as having pinnule apices entire, the

segment apices rostrate, and the rachis bicanalicu-

late, whereas in D. remotum the pinnule apices are

supposed to be serrate, the segment apices dentate,

and the rachis simply canaliculate. Dr. Badre has

stated (in litt.) that the scales of the holotype (Gla-

ziou 2331) are identical to those of D. remotum.

Skog & Skog 5210 (us), from Cuzco, apparently

is a hybrid involving D. remotum and another

species. The specimen has proliferous buds scat-

tered all over the lamina, dentate scales (with teeth

that are not always dark or bifid), and axes often

puberulent abaxially. Most sporangia are abortive.

Amazonas: Prov. Bagua, 20 km E of La Peca, Barbour

2763 (F, MO, uc). San Martin: Prov. Mariscal Caceres,

Rio Abiseo National Park, Leon 2160 (F). Pasco: Prov.

Oxapampa, San Alberto, Cordillera de Yanachaga, van

der Werffet al. 8457 (MO). Cuzco: Prov. La Convention,

Cordillera Vilcabamba, Dudley 10460 (GH, MO). Prov.

Paucartambo, Valle de Pillahuata, Herrera 1623 (us in

part).

10. Diplazium expansum Willd., Sp. pi. ed. 4, 5:

354. 1 8 1 0. TYPE: Venezuela, Caracas, Brede-

meyer (holotype, B!, Herb. Willd. 19948; pho-

tos, F, GH, US).

Asplenium expansum (Willd.) Presl, Reliq. haenk. 1 :

46. 1825.

Allantodia asplenioides Kunze, Linnaea 9: 72. 1834.

TYPE: Peru, Huanuco, Cuchero, Poeppig, Aug.

1829 (holotype, w; isotypes, B!, BM; prob. isotype,

MO!; photos, BM, of w; GH, p & us of BM).

Diplazium asplenioides (Kunze) Presl, Tent, pterid

114. 1836.

Athyrium expansum (Willd.) Milde, Bot. Zeit. 28: 353.

1870, not Moore, 1860.

Diplazium bonapartii Rosenst., Repert. Spec. Nov.

Regni Veg. 7: 295. 1909. TYPE: Peru (San Mar-

tin), Monte "Guayrapurina" (Guayrapurima),

Spruce 4683 (holotype, P!; frag., us!; photos, F,

GH, us).

Plants terrestrial. Stem erect, this and the petiole

naked, or with a few dark brown scales. Leaves to

2 m long, 2-pinnate-pinnatifid. Lamina firm-

membranaceous, to 70 cm broad, gradually re-

duced to a pinnatifid apex, minutely puberulent

on costae, costules, veins, and leaf tissue abaxially,

trichomes of the axes reddish brown, septate, 0. 1-

0.3 mm long, those between the veins mostly uni-

cellular and whitish. Pinnae truncate and subequi-

lateral at base, mostly stalked. Pinnules 10 pairs

or more on larger pinnae, deeply incised (some

basal ones nearly to the costule), truncate and sub-

equilateral at base. Ultimate segments 8-12 pairs,

mostly obtuse or truncate. Veins free. Sori mostly

2 mm long or less and single, or those of basal

veins diplazioid. Indusia linear, thin, dark brown,

erose-ciliolate, subpersistent.

In dense, wet forests, 100-2000 m, Loreto and

San Martin to Cuzco.

Greater Antilles; southern Mexico; Guatemala;

Venezuela; Colombia to Peru; Brazil.

This is distinguished from most species in the

genus by the abundant minute trichomes on the

tissue between the veins abaxially. Diplazium bon-

apartii has numerous, rather than sparse, ap-

pressed scales on the costules, but is otherwise

identical, therefore conspecific. Diplazium mela-

nosorum (Sodiro) C. Chr., of Ecuador and Colom-

bia, is similar, but the lamina is subcoriaceous,

and indusia are broadly elliptic instead of linear.

San Martin: Near Tarapoto, Spruce 4124 (BM, K). Lo-

reto: Prov. Maynas, 50 mi from Iquitos, Moran 3655

(MO). Huanuco: Prov. Huanuco, Rio Huallaga, above

Rio Cayumba, Mexia 8316 (K, us). Junin: Prov. Tarma,

Chanchamayo, Esposto (USM), C. Schunke 35, 43, 499

(F). Ucayali: Prov. Coronel Portillo, Boqueron, Ferreyra

16057 (GH, USM in part). Cuzco: Prov. Quispicanchi,

Valle de Marcapata, Herrera 1595 (us).

11. Diplazium venulosum (Baker) Diels, Nat.

Pflanzenfam. 1(4): 226, 228. 1899.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

75

Asplenium venulosum Baker, Syn. fil. 238. 1 867. TYPE:

Ecuador, Mt. Tungurahua, Spruce 5343 (holo-

type, K.!; isotypes, BM!, P!, w!; photos, F & GH of

K).

Plants terrestrial. Stem erect, provided with co-

pious, linear, brown scales, these to 1.5 cm long,

their margins remotely ciliolate or denticulate.

Leaves to 3 m long, 2-pinnate, the petiole densely

scaly as on the stem, but marginal teeth of scales

elongated, more numerous and crowded. Lamina

chartaceous, to 85 cm broad, often with a prolif-

erous bud distally on the rachis, indument of the

axes abaxially a mixture of minute trichomes, scurf

(appressed squamules), and filiform brown, den-

tate scales. Pinnae narrowly acute at apex, sub-

equilateral and truncate at base, mostly stalked,

patent or slightly ascending. Pinnules remote, en-

tire to crenate-serrate, free ones of larger pinnae

12-16 pairs, larger ones 6-12 cm long, 4-6 times

as long as broad, costules adaxially provided with

low wings of tissue perpendicular to the plane of

the lamina, but these not abruptly expanded or

cristate. Veins free. Sori linear, 5-10 mm long.

Indusia dark brown, becoming blackish at base,

firm, persistent, the margins subentire.

Rare, in dense, wet forests, usually along stream

banks, 1700-1900 m, Amazonas and San Martin.

Colombia; Ecuador; Peru.

In addition to the key characters, this species

also can be separated from D. tungurahuae by the

dense covering of linear, attenuate, tortuous, scales

on the petiole. Few species ofDiplazium have such

an abundance of petiole scales. The few (if any)

scales on the petiole base of D. tungurahuae are

broad, rigid and appressed, with margins suben-

tire.

Amazonas: Prov. Bagua, Cordillera Colan SE of La

Peca, Barbour 4189 (USM). San Martin: Prov. Rioja,

Venceremos, D. Smith 4456 (F, MO, uc).

1 2. Diplazium tungurahuae (Sodiro) C. Chr., Ind.

fil. suppl. 1: 28. 1913.

Asplenium tungurahuae Sodiro, Anal. Univ. Central

Quito 22: 97 (Sert. fl. Ecuad. 2: 20) 1908. TYPE:

Ecuador, Tungurahua, Sodiro, in 1 904 (holotype,

Q?; isotype, P!; frag., us!; photos, GH & us of P).

Asplenium crassifolium Sodiro, Anal. Univ. Central

Quito 22: 97 (Sert. fl. Ecuad. 2: 21) 1908. TYPE:

Ecuador, Chillanes, Mt. Chimborazo, Sodiro, in

1891 (holotype, Q?; isotype, P!; photos, F, GH &

us of P).

Diplazium crassifolium (Sodiro) C. Chr., Ind. fil. suppl.

1: 26. 1913.

Plants terrestrial. Stem erect or decumbent, pro-

vided at apex with coarse, dark brown scales about

1 cm long. Leaves to 1.5 m long, 2-pinnate. Lam-

ina chartaceous, to 50 cm broad, often with pro-

liferous buds on rachis or costae, indument of the

axes abaxially a mixture of minute, 1- or 2-celled

trichomes, broad, appressed scales, and scurf (ap-

pressed squamules). Pinnae subequilateral and

truncate at base, mostly stalked. Pinnules suben-

tire to pinnatifid, sessile to short-stalked, broadest

at the base, free ones 8 pairs or fewer, larger ones

4-9 cm long, 3-3.5 times as long as broad, costules

adaxially provided with low wings of tissue per-

pendicular to the plane of the lamina, but these

not abruptly expanded or cristate. Veins free. Sori

linear, commonly 4-6 mm long. Indusia light

brown, persistent, with margins subentire or light-

ly erose.

In dense, wet forests, often in ravines or along

stream banks, 100-2300 m, Cajamarca, Amazo-

nas, and Loreto to Junin.

Colombia; Ecuador; Peru.

This species is a coarse fern, the broad pinnae

and pinnules mostly triangular from a truncate

base. As in most of the puberulent group of de-

compound species, the trichomes on the costae

and costules abaxially are minute (in D. tungura-

huae mostly 0.1 mm long and 1-2-celled). The

minor axes also have few to many broad, ap-

pressed scales, and the costa and rachis also often

have a sparse to ample covering of appressed squa-

mules, that sometimes partially obscure the tri-

chomes. Not all specimens examined have prolif-

erous buds, but on those that do the buds often

are on the costae as well as the rachis. This may

be only a variant of D. striatum, under which see

further discussion.

Cajamarca: Prov. Cutervo, road to San Andres Na-

tional Park, Lopez & Sagastegui 5430 (GH, HUT, uc).

Amazonas: Prov. Bongara, Shillac, D. Smith & Vasquez

4886 (MO, uc). Loreto: Balsapuerto, lower Rio Huallaga

basin, Killip & Smith 28506 (us). Huanuco: Fundo Chela,

Sinchono, Aguilar 921 (USM). Tingo Maria, Tryon & Tryon

5283 (BM, F, GH, us). Pasco: Prov. Oxapampa, Cordillera

de San Matias, D. Smith 2004 (F, MO). Junin: Chancha-

mayo Valley, C. Schunke 960 (F, us).

13. Diplazium flexuosum Presl, Tent, pterid. 1 14.

1836, nom. nov. for Asplenium flexuosum Presl

and with the same type.

76

FIELDIANA: BOTANY

Asplenium flexuosum Presl, Reliq. haenk. 1: 46, t. 7,

f. 1. 1825, not Schrader, 1818. TYPE: Peru, "in

vallibus Cordillerarum," Haenke (holotype, PR or

PRC; isotype, K).

Athyrium flexuosum (Presl) Milde, Hot. Zeit. (Berlin)

28: 353. 1870.

Diplazium preslianum C. Chr., Index fil. 237. 1905,

nom. super/1, for Diplazium flexusosum Presl and

with the same type.

Plants terrestrial, probably clambering. Stem

decumbent, provided at the apex with coarse,

blackish scales to 6 mm long. Leaves to 2 m long,

2-pinnate-pinnatisect. Lamina chartaceous, essen-

tially glabrous, with some scattered trichomes on

the rachis, but not regularly puberulent, the axes

conspicuously flexuous. Pinnae strongly reflexed,

proximal ones long-stalked, to 60 cm long. Pin-

nules strongly reflexed, lobed to pinnatisect, in-

equilateral at base, broadly cuneate acroscopically,

truncate and more strongly produced basiscopi-

cally. Veins free. Sori linear, 2-4 mm long, those

of basal veins mostly diplazioid, the rest usually

single. Indusia light brown or gray-brown, per-

sistent, the margins entire to lightly erose.

In forests, leaves probably clambering on shrubs,

about 1 600 m, Huanuco and Ucayali.

Rare and endemic.

This cannot be confused with any other species

of the genus in Peru. The alternate pinnae and

pinnules are strongly reflexed and abruptly bent

at the axils, which results in a conspicuous, flex-

uous aspect to the axes.

Huanuco: Pampayacu, Kanehira 181 (GH, us). Ucayali

(as Loreto): Prov. Coronel Portillo, NE of the pass at La

Divisoria, Skoget al. 5151 (F, us). Department unknown:

Matthews 1818 (BM, K, us), Poeppig, in 1829 (K, p).

14. Diplazium macrophyllum Desv., Prodr. 280.

1827. TYPE: "Hab. in America calidiori,"

collector and locality not cited, but possibly

Dombey or Poeppig, Peru (holotype, p; pho-

tos, GH, us).

Asplenium desvauxii Mett., Abh. Senckenberg. Na-

turf. Ges. 3: 225, / 4, 1859, nom. nov. for Dipla-

zium macrophyllum Desv., not Asplenium mac-

rophyllum Mett. 1856.

Asplenium procerum Sodiro, Anal. Univ. Central Qui-

to 22: 96 (Sert. Fl. Ecuad. 2: 19). 1908. TYPE:

Ecuador "in silvis subtropic. val. Nanegal," So-

diro, 1 89 1 (holotype, Q?; isotype, P!, 3 sheets; pho-

tos, F, GH).

Plants terrestrial. Stem erect, provided at apex

with sublustrous, brown scales, these 8-14 mm

long. Leaves to 2.5 m long, 1-pinnate-pinnatisect

(at least proximally), petiole sparsely scaly toward

base. Lamina to nearly 1 m broad, terminating

abruptly in a pinnatifid apex, essentially glabrous

(rachis rarely sparsely puberulent), but sparsely

provided abaxially on costae and rachis with dull,

light brown scales, these filiform or attenuate,

mostly flaccid and appressed, and distal pinna ax-

ils commonly provided abaxially with proliferous

buds. Pinnae subequilateral, proximal ones del-

toid or subdeltoid, stalked, incised nearly (rarely

quite) to the costa, distal ones subentire, larger

pinnae 9-22 cm broad, their ultimate segments

with attenuate or acuminate apices. Veins free,

pinnately branched in the segments. Sori single,

or often diplazioid on the basal acroscopic vein

branch. Indusia commonly persistent, light to dark

brown, or often blackish near their attachment to

the vein.

In dense, wet forests, often in ravines and along

stream and river banks, 400-1800 m, Amazonas

to Cuzco.

Venezuela; Colombia to Bolivia.

Specimens of D. macrophyllum are often deter-

mined as D. costale (Sw.) Presl of Jamaica, which

differs in the darker, larger, very abundant scales

on the costae, the deeply serrate segment apices,

and the short (ca. 5 mm) sori, which extend only

halfway or less to the segment margin. There are

several other taxa in this species complex, which

is in need of more detailed examination: Dipla-

zium appolinaris Fee (Lesser Antilles) belongs here,

as well as D. oxylobum Sodiro (Ecuador). The lat-

ter, with short sori, appears to be intermediate

between D. macrophyllum and D. costale.

A collection from San Martin, Knapp & Alcorn

7749 (F, MO), is similar to D. macrophyllum in

every way except for its exceedingly dense indu-

ment. Leaves of the latter are essential glabrous,

and with a few scales, but in this specimen the

petiole and rachis are covered with attenuate,

brown scales with setulose margins, these to 2 cm

long toward the stem, becoming shorter and less

abundant in the distal portion of the lamina. In

addition, the rachis distally and the costae and

veins on both sides are beset with copious septate

trichomes up to 1 mm long. A number of shorter

trichomes also are borne adaxially on the tissue

between the veins. Although the specimen is ro-

bust (leaf 1.5 m long), it is sterile, and may be

merely a monstrous form or hybrid. At any rate,

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

77

it is inappropriate here to describe a new form or

variety on the basis of a single, sterile specimen.

Ama/onas: Prov. Bagua, valley of Rio Maranon above

Cascadas de Mayasi, Wurdack 1860 (us). San Martin:

Mt. Campana, near Tarapoto, Spruce 4336 (BM, K, p).

Huanuco: Prov. Huanuco, Dist. Churubamba, Mt. Santo

Toribio, Mexia 8249 (BM, F, GH, MO, uc, us). Pasco:

Prov. Oxapampa, Palcazu Valley, Iscozacin, Foster et

al. 7881 (F, MO). Junin: Schunke Hacienda, above San

Ramon, C. Schunke A- 194 (GH, us). Ucayali (as Loreto):

Along Rio Aguaytia above mouth of Quebrada Yurac-

Yacu, Croat 20888 (F, MO, uc). Cuzco: Prov. La Con-

vention, Cordillera Vilcabamba, Rio Klause, Dudley

10182 (GH, us). Puno: Near "San Gavan" (San Gaban),

Lechler 2158 (K, p).

1 5. Diplazium subobtusum Rosenst., Repert. Spec.

Nov. Regni Veg. 7: 296. 1909. TYPE: Peru,

San Martin, Mt. Guayrapurima (as Ecuador,

"Monte Guayrapurina"), Spruce 4019 (ho-

lotype, P!; frag., us!; photos, F, GH, us).

Plants apparently terrestrial. Stem erect, pro-

vided at apex with a few broad, castaneous scales.

Leaves 75 cm long, 1 -pinna te-pinnatisect above

the 2-pinnate basal pinnae, petiole moderately pu-

bescent, lacking scales. Lamina 22 cm broad, grad-

ually reduced to a pinnatifid apex, axes and veins

moderately puberulent abaxially with spreading,

flexuous trichomes to 0.3 mm long, the interven-

ing tissue also with scattered, shorter trichomes,

costae abaxially with a few scattered, appressed,

light brown scales, rachis lacking proliferous buds.

Pinnae subequilateral, short-stalked, incised near-

ly (the basal pair quite) to the costa, the ultimate

segments of proximal pinnae narrowly acute, their

margins crenate to shallowly lobed. Veins free.

Sori single, or frequently diplazioid. Indusia per-

sistent, thin, light brown, the margins entire to

slightly erose.

Known thus far in Peru only from the type col-

lection, San Martin.

Costa Rica; Panama; Peru.

This is one of the questionable species related

to D. striatum, from which it differs principally in

the great degree of pinna dissection, a feature that

is variable in the latter species. Apparently D. sub-

obtusum has not been collected again in Peru since

the type, nearly 130 years ago. It may be merely

a variant ofD. striatum. However, no matter how

deeply lobed the pinnae of the latter, the ultimate

segments are essentially entire, with obtuse to sub-

truncate apices (rarely subacute). Most of the seg-

ments in proximal pinnae of D. subobtusum are

not only narrowly acute, but their margins are con-

spicuously lobed. In this respect, they are some-

what intermediate between D. striatum and D.

macrophyllum. Furthermore, the leaf is only 75

cm long and 22 cm broad, whereas most leaves of

D. striatum are 1-2 m long. Tentatively, then, D.

subobtusum is maintained here as distinct.

16. Diplazium lindbergii (Mett.) Christ, Prim. Fl.

Costar. 3: 27. 1901.

Asplenium lindbergii Mett., Ann. Sci. Nat. Bot. ser. 5,

2: 236. 1864. LECTOTYPE (designated by Lei-

linger, Proc. Biol. Soc. Wash. 89: 707. 1977): Bra-

zil, Caldes, Lindberg 543 (B).

Plants terrestrial. Stem erect or decumbent,

sparsely scaly. Leaves to 1.5 m long, 1-pinnate-

pinnatisect, petiole dark reddish brown to atro-

purpureous, toward the base provided with dull

brown, linear or lanceolate scales about 1.5 cm

long. Lamina chartaceous to subcoriaceous, to 40

cm broad, generally reduced to a pinnatifid apex,

some sparse, filiform scales on the apex, glabrous

except for some scattered, septate trichomes abax-

ially on the costa and segment midribs, lacking

proliferous buds. Pinnae subequilateral, linear or

linear-lanceolate, subsessile or short-stalked, in-

cised % to % (or rarely quite) to the costa, costae

awned adaxially near the base of costules, ultimate

segments more than 20 pairs on larger pinnae,

obtuse to truncate, often strongly revolute. Veins

free. Sori 1 .5-2.5 mm long (slightly longer on basal

veins), often diplazioid. Indusia vestigial or lack-

ing.

Rare in Peru, in wet forests, 1 800-2000 m, San

Martin, Pasco, and Cuzco.

Southern Mexico to Panama; Venezuela and

Colombia, south to Bolivia and Brazil.

In this species, the raised, adaxial edges of costae

are strongly produced, appearing usually as per-

pendicular, herbaceous wings. Where these are in-

terrupted near the costule bases, the ends com-

monly separate from the costa, as short, subacute

awns. A similar condition is evident in D. stria-

tum, but the wings are less pronounced and the

awns are less conspicuous or lacking in that spe-

cies.

Several other neotropical species probably

should be included here, among them D. grande

78

FIELDIANA: BOTANY

(Baker) C. Chr. (Colombia, Ecuador), D. indura-

tum Diels (Costa Rica), and D. subnudum (Kar-

sten) Alston (Colombia). Each has been separated

from D. lindbergii chiefly on the degree of pinna

dissection. In Diplazium, this highly variable fea-

ture is of little value when unsupported by other

characters.

Specimens of D. lindbergii are often determined

as D. brasiliense Rosenst., which is similar in as-

pect and texture. However, the type of the latter

(Brazil, Haerchen 91, s!) has broader and fewer

segments (to 1 2 pairs), the indusia are subpersis-

tent, and the axes are minutely, but densely, pu-

berulent abaxially.

San Martin: Prov. Rioja, Venceremos, D. Smith &

Vasquez 4995 (F, MO, uc). Pasco: Prov. Oxapampa, SE

of Oxapampa, D. Smith 2909 (F, MO). Cuzco: Prov. Pau-

cartambo, Valle de Pilcopata, Herrera 1623 (us).

17. Diplazium striatum (L.) Presl, Tent, pterid.

114. 1836. Figure 16c-d.

Asplenium striatum L., Sp. pi. 2: 1082. 1753. TYPE:

Petiver, Pter. Amer. t. 3, f. 3, 4, copied from

Plumier, Traite foug. Amer. /. 18, 19, based on

a specimen from Martinique.

Diplazium obtusum Desv., Prodr. 281. 1827. TYPE:

"Crescit in Peruvia" ("Habitat in America cali-

diori" on label), collector unknown (holotype, P!;

photos, BM, GH, us).

Diplazium tabalosense Hieron., Hedwigia 47: 214.

1 908. TYPE: Peru (San Martin), above Tabalosos

on the way to Moyobamba, Huallaga River val-

ley, Stubel 1089 (holotype, B!; photos, BM, F, OH,

us).

Plants terrestrial. Stem erect, near the apex

sparsely provided with dark brown scales to 1 cm

long. Leaves to 2 m long, 1 -pinnate-pinnatifid to

nearly 2-pinnate, petiole brown at base, lighter

brown to stramineous distally, scaly at base. Lam-

ina firm-membranaceous, to 50 cm broad, lacking

proliferous buds, reduced to a pinnatifid apex,

abaxially glabrous to minute puberulent, and axes

sparsely to moderately provided with appressed,

attenuate scales. Pinnae (2.5-)3.5-5(-6) cm broad,

lanceolate or deltoid-lanceolate, sessile to short-

stalked, incised V2 to % (or sometimes quite) to the

costa, costae with perpendicular herbaceous wings

adaxially, these interrupted at base of costules,

there sometimes ending as short awns. Ultimate

segments 10-15 pairs or fewer, their apices obtuse

to subtruncate (rarely subacute), their margins

plane. Sori mostly 3-7 mm long, often diplazioid.

Indusia thin, light brown, often shriveling (but

persistent) at maturity.

In dense forests, often in wet ravines or on river

banks, 100-1800 m, Cajamarca, Amazonas, and

Loreto south to Cuzco and Madre de Dios.

West Indies; southern Mexico to Panama; Ven-

ezuela and Colombia to Bolivia.

This species varies considerably in leaf dissec-

tion. Further study may prove that a number of

related taxa are conspecific. Larger pinnae of D.

striatum vary from 2 cm broad and cut halfway

to the costa, to 6 cm broad and dissected nearly

or quite to the costa. Narrower pinnae are more

common toward the north of the range, whereas

specimens from Peru typically have broader and

deeply dissected pinnae, as in the type of D. ta-

balosense. The leaves of D. tungurahuae are even

more highly dissected. It is similar in most other

characters and may better be considered a variety

of D. striatum. In fact, some sheets of Schunke

295 (below) are intermediate between the two. A

species with narrower, less dissected pinnae is D.

angelipolitanum Rosenst. (Colombia), which

probably should be included with D. striatum.

Cajamarca: Prov. Cutervo, Dist. Cujillo, Tambillo,

Jelski 1053 (GH, us). Amazonas: Prov. Bagua, between

Aramango and Montenegro, Lopez et al. 4223 (GH, HUT).

San Martin: Mt. Campana, near Tarapoto, Spruce 4685

(K, P, w). Loreto: Mouth of Rio Santiago, Pongo de Man-

seriche, Mexia 6729 (B, F, GH, K., MO, uc, us). Gamitan-

acocha, Rio Mazan, /. Schunke 295 (F, GH, uc, USM).

Pasco: Pozuzo, Hacienda Ballisteros, Bryan 697, 698 (F).

Junin: Chanchamayo Valley, C. Schunke 1344 (F). Cuz-

co: Prov. Paucartambo, Villa Carmen, Vargas 11246,

14683 (GH). Madre de Dios: Parque Nacional del Manu,

M. Foster P-84-57, P-84-86 (uc).

18. Diplazium caracasanum (Willd.) Moore, In-

dex fil. 324. 1861.

Asplenium caracasanum Willd., Sp. pi. ed. 4, 5: 338.

1810. TYPE: Venezuela, Caracas, Bredemeyer

(holotype, B!; Herb. Willd. 19919; photos, F, GH,

us).

Asplenium striatum var. caracasanum (Willd.) D. C.

Eaton, Mem. Amer. Acad. Arts 8: 206. 1860.

Diplazium shepherdii var. proliferum Rosenst., Re-

pert. Spec. Nov. Regni Veg. 7: 294. 1909. SYN-

TYPES: Peru (San Martin), "in silvis secus flu-

men Mayo," Spruce 4755 (B!, GH!, K!, P!); Spruce

4785 (BM!, P!, us!).

Plants terrestrial. Stem erect, provided at apex

with a few medium to dark brown scales, these

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

79

coarse, often sublustrous, 3-6 mm long. Leaves

50-90 cm long, 1 -pinnate-pinnatifid, petiole na-

ked or with a few dark brown scales at base. Lam-

ina 10-20 cm broad, gradually reduced to a pin-

natifid apex, glabrous and with scales rare or

lacking, bearing a proliferous bud adaxially on the

rachis toward the apex. Pinnae 1 6-20 pairs, all but

the distal ones cut 34 or nearly to the costa, mostly

inequilateral at base, but proximal ones subequi-

lateral, segments subacute, larger ones 2-3 times

as long as broad. Veins free, pinnately branched

in the segments. Sori single, or sometimes dipla-

zioid on the basal acroscopic vein branch. Indusia

thin-textured, dull brown, but sometimes with a

very fine white margin.

In forests, usually along streams or in ravines,

ca. 700 m, San Martin, Huanuco.

Lesser Antilles; Venezuela; Peru.

Some taxa in the D. cristatum complex may not

merit species status, as they seem to differ only in

the degree of pinna dissection. D. caracasanum,

however, is distinguished by additional characters,

as noted in the key. Furthermore, indusia of the

latter often bear a fine white line along the margin,

as in the next species, D. stuebelianum. This char-

acter, a very rare one in the genus, was not noted

in the original descriptions of the two species.

Huanuco: Prov. Huanuco, near Tingo Maria, Tryon

& Tryon 5230 (BM, F, GH, uc, us).

19. Diplazium stuebelianum (Hieron.) Stolze, stat.

et comb. nov.

Diplazium shepherdii var. stuebeliana Hieron., Hed-

wigia 47: 212. 1908. LECTOTYPE (designated

here): Peru (Amazonas?), between Pacasmayo and

Moyobamba, Stiibel 70726 (B!; photos, F, GH).

PARATYPE: Ecuador, between Banos and Ji-

varia de Pintuc, valle Pastaza, Stiibel 986 (B!; pho-

tos, F, GH).

Plants terrestrial. Stem erect to decumbent, pro-

vided at apex with dull, medium brown scales,

these somewhat flaccid, 3-4 mm long. Leaves 42-

58 cm long, 1 -pinnate-pinnatisect, petiole naked

or with a few dark brown scales at base. Lamina

8-16 cm broad, gradually reduced to a pinnatifid

apex, glabrous and without scales, proliferous buds

lacking. Pinnae 10-15 pairs, strongly inequilateral

at base, proximal ones (at least) incised nearly to

the costa, their basal acroscopic segments free and

usually short-stalked. Veins free, pinnately branched

in the segments. Sori single, or sometimes dipla-

zioid on the basal acroscopic vein branch. Indusia

thin-textured, dull brown, with a very fine white

margin.

In forest and thickets, 1200-1700 m, Amazo-

nas?, Junin, Ucayali.

Ecuador; Peru; Bolivia.

Like several other taxa in the species complex

(e.g., D. lonchophyllum and D. werckleanum), D.

stuebelianum differs from D. cristatum in degree

of pinna dissection. However, unlike these others,

D. stuebelianum is also distinguished by some good

qualitative characters. Scales at the stem apex in

D. cristatum and nearest allies are coarse, blackish,

sublustrous, and usually less than 3 mm long,

whereas in D. stuebelianum they are nearly flaccid,

dull brown, and 3-4 mm long. The indusium color

is also a very subtle but distinctive character. In

fully expanded indusia, there is a very fine, chalky

white line along the margin. Indusia of D. crista-

tum are concolorous, although a white-edged mar-

gin is often present in D. caracasanum.

Junin: Schunke Hacienda, above San Ramon, C.

Schunke A- 189 (us). La Merced, Chanchamayo, C.

Schunke (A, P). Chanchamayo Valley, C. Schunke 112

(us), 729 (F, us). Ucayali: Prov. Coronel Portillo, Sin-

chono, Aguilar 892 (GH, USM).

20. Diplazium cristatum (Desr.) Alston, J. Hot.

74: 173. 1936. Figure 16a.

Meniscium cristatum Desr. in Lam., Encycl. 4: 94.

1797. TYPE: Martinique, Joseph Martin (holo-

type, P!, Herb. Lamarck).

Asplenium arboreum Willd., Sp. pi. ed. 4, 5: 320. 1810.

TYPE: Venezuela, Caracas, Bredemeyer (holo-

type, B!, Herb. Willd. 19892-1; photos, F, GH).

Asplenium denticulosum Desv., Ges. Naturf. Freunde

Berlin Mag. Neuesten Entdeck. Gesammten Na-

turk. 5: 323. 1811 (not Gaud. 1827). TYPE: "In

America calidiore," Dombey (holotype; P!; photo,

GH).

Asplenium shepherdii Sprengel, Nova Acta Phys.-Med.

Acad. Caes. Leop.-Carol. Nat. Cur. 10:231.1821.

TYPE: Jamaica, Henry Shepherd (holotype, LZ,

destroyed; isotype, PH).

Diplazium shepherdii (Sprengel) Link, Hort. berol. 2:

70: 1833.

Diplazium arboreum (Willd.) Presl, Tent, pterid. 1 14.

1836.

Diplazium denticulosum (Desv.) C. Chr., Index fil.

231. 1905.

Plants terrestrial. Stem erect, provided at the

apex witih a few castaneous to blackish scales,

80

FIELDIANA: BOTANY

these coarse, often somewhat lustrous, 1-3 mm

long. Leaves 25-80 cm long, 1-pinnate-pinnatifid,

petiole naked or with a few dark brown scales at

base. Lamina to 30 cm broad, gradually reduced

to a pinnatifid apex, glabrous and lacking scales,

proliferous buds absent (or very rarely present).

Pinnae 8-12 free pairs, mostly lobed V4 to % to the

costa (rarely more deeply), inequilateral at base,

i.e., cuneate basiscopically, truncate acroscopical-

ly, the basal acroscopic lobe or segment much more

strongly produced than the opposing basiscopic

one, segments obtuse, 1 or 2 times as long as broad.

Veins free, pinnately branched in the segments.

Sori commonly diplazioid on the basal acroscopic

vein branch, single on the others. Indusia thin-

textured, dull brown, the margin subentire.

In dense wet or dry forests, often in ravines or

along stream banks, 300-1700 m, Amazonas and

Loreto to Ayacucho and Madre de Dios.

West Indies; Mexico to Colombia and Vene-

zuela, south to Argentina and Paraguay.

This and near allies form a neotropical species

complex that is as taxonomically difficult as that

ofAsplenium auritum/ cuspidatum. Because of the

variability of leaf dissection it has been divided

into a number of species, varieties, and forms. The

group is characterized by glabrous, medium-sized,

pinnate to pinnate-pinnatisect leaves that taper to

a pinnatifid apex, pinnae with conspicuously in-

equilateral base (i.e., excavate to cuneate basi-

scopically, truncate and much more strongly pro-

duced acroscopically), and stem apices bearing only

a few, small, coarse, blackish scales. With pinnae

deeply lobed to pinnatifid, D. cristatum is at the

center of the complex. Taxa with entire to shal-

lowly lobed pinnae are D. werckleanum Christ

(Mesoamerica), D. unilobum (Poir.) Hieron. (West

Indies), and D. bombonasae. The species D. lon-

chophyllum Kunze (Mesoamerica) and D. drepa-

nolobium A. R. Smith (southern Mexico) have

pinnae dissected nearly or quite to the costa. Ul-

timately, monographic treatment will probably

determine that most of these names should be

synonyms of D. cristatum. Two specimens of the

latter seen from Peru could easily key out to D.

werckleanum, and several others to D. loncho-

phyllum in the keys of Stolze (1981) and other

recent neotropical Floras. See D. bombonasae and

D. stuebelianum for further comparisons.

C. Schunke 500 and 944 (F) from Junin, with

proximal pinnae deeply dissected, are practically

identical to Central American specimens of D. lon-

chophyllum. Both specimens also bear proliferous

buds distally on the rachis, a condition extremely

rare in D. cristatum but a diagnostic feature in D.

caracasanum.

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

2500 (MO, uc). San Martin: River Mayo, near Tarapoto,

Spruce 4758 (BM, GH, K, us). Loreto: "Flumen Maranon

inf. fluv. Huallaga," Spruce 3911 (K). Pasco: Pozuzo,

Hacienda Ballisteros, Bryan 691, 693 (F). Junin: Colonia

Perene, Killip & Smith 24925 (F, GH, us). Ucayali: Prov.

Coronel Portillo, Dist. Iparia, Bosque Nacional de

Imparia, /. Schunke V. 2733 (F, GH, us). Ayacucho: Prov.

San Miguel-La Mar, Teresita, Barron (USM). Madre de

Dios: Rio Manu, Cocha Cashu Station, Parque Nacional

del Manu, Foster et al. 7197 (F).

2 1 . Diplazium bombonasae Rosenst, Repert. Spec.

Nov. Regni Veg. 7. 294. 1909. TYPE: Ec-

uador, Rio Bombonasa, Spruce (holotype, P!,

Herb. Bonaparte 10027; photos, GH & us).

Plants terrestrial. Stem erect to decumbent, pro-

vided at the apex with a few dark brown to blackish

scales, these coarse, sometimes sublustrous, 2-3

mm long. Leaves 35-65 cm long, 1 -pinnate, pet-

iole gray-brown, with a few dark brown scales at

the base. Lamina to 1 2 cm broad, firm-herbaceous

to chartaceous, gray-green, gradually reduced to a

subattenuate, pinnatifid apex, essentially glabrous

and lacking scales, lacking proliferous buds. Pin-

nae 15-24 pairs, approximate to subdistant, fal-

cate, attenuate, conspicuously inequilateral at base

(attenuate to excavate basiscopically, truncate ac-

roscopically and with a pronounced basal auricle),

basiscopic margins beyond the basal auricle sub-

entire, crenate, or shallowly lobed, or sometimes

pinnatifid as to basal pinnae. Veins free, com-

monly 1-3-forked. Sori single or diplazioid, borne

on the acroscopic branch of each vein. Indusia

thin-textured, dull brown, the margins subentire

to erose.

In forests, on stream banks or on slopes of ra-

vines, 180-600 m, Loreto, Huanuco, Madre de

Dios.

Ecuador; Peru; Bolivia.

As in other taxa related to D. cristatum, pinna

dissection in D. bombonasae can vary widely. Be-

yond the enlarged basal auricle, pinna margins may

be essentially entire throughout the lamina (as in

the type), or shallowly to deeply lobed. In deeply

lobed pinnae the acroscopic auricle is sometimes

incised nearly to the costa.

South American specimens are sometimes iden-

tified as D. unilobum (Poir.) Hieron. of the West

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

81

Indies, a similar taxon with subentire, but strongly

auriculate, pinnae. However, stem scales of the

latter are 2-3 times longer, the lamina is lighter

in color (yellow-green) and often subcoriaceous,

and the petiole and rachis are stramineous. Al-

though monographic study may prove that both

taxa are only variants of D. cristatum, D. bom-

bonasae is tentatively maintained here as a distinct

species.

Loreto: Near Rio Santiago above Pongo de Manse-

riche, Mexia 6217 (BM, GH, K, MO, uc). Huanuco: Prov.

Leoncio Prado, Dist. Rupa Rupa, 5 km from Tingo Ma-

ria, J. Schunke V. 3267 (F, GH, us). Madre de Dios: Prov.

Manu, Atalaya, vicinity of Hacienda Amazonia, Foster

& Wachter 7445 (F, MO).

22. Diplazium cuneifolium Rosenst, Repert. Spec.

Nov. Regni Veg. 12: 470. 1913. TYPE: Bo-

livia, North Yungas, Polo-Polo near Coroico,

Buchtien 3390 (holotype, s; isotypes, GH!, P!,

us!; photo, BM of s, F, & us of P).

Plants terrestrial. Stem erect, provided with

coarse, broad, dark gray-brown to blackish scales

about 5 mm long. Leaves to 80 cm long, 1 -pinnate.

Lamina to 24 cm broad, membranaceous, grad-

ually reduced to a short, pinnatind apex, lacking

scales, sparsely and minutely puberulent on the

axes and adjacent tissue abaxially, costae (toward

the base) and rachis moderately and minutely pu-

berulent within the adaxial sulci. Rachis deeply

sulcate adaxially, lacking proliferous buds. Pinnae

11-17 pairs, well-spaced, at least proximal ones

short-stalked, broadest near the center, narrowly

acute at apex, narrowly to broadly cuneate at the

subequilateral base, margins lobed about one-

quarter to costa, larger pinnae 10-13 cm long and

2-3 cm broad. Veins free, pinnately branched. Sori

borne on most of the vein branches, extending

from the costa nearly to the pinna margin, usually

diplazioid on the basal branches, the other sori

usually single. Indusia thin, dull brown, margin

subentire to erose or irregularly fimbriate.

In forests, 700-1000 m, Junin, Cuzco, Madre

de Dios.

Peru and Bolivia.

This is a rarely collected species, probably con-

fined to lower mountain slopes of Bolivia and the

southern half of Peru. In the type collection, pin-

nae in the proximal half of the lamina have rather

narrowly cuneate bases, but in all other specimens

examined the pinna bases are much more broadly

cuneate. Considering the variability typical of oth-

er species of Diplazium, it seems likely that the

narrowly cuneate base seen in the type will be a

character more exceptional than representative.

Junin: Prov. Satipo, San Francisco de Satipo, Solomon

3315 (F, MO). Prov. Satipo, Reserva Forestal de Universi-

dad Central near Satipo, van der Werff et al. 8633 (MO,

uc). Cuzco: Prov. La Convention, Palma Real, Vargas

17296 (GH). Madre de Dios: Prov. Manu, Atalaya, vi-

cinity of Hacienda Amazonia, Foster & Wachter 7447

(F).

23. Diplazium grandifolium (Sw.) Sw. var. andi-

cola Stolze, var. nov.

Varietas haec a varietate typica differt indusiis 0.2-

0.4 mm latis, tenuibus, brunneolis, planis (nee usque ad

1 mm latis, nee crassis, nee bicoloribus, nee involutis in

maturis).

Plants terrestrial. Stem erect or decumbent, pro-

vided with coarse, broad, blackish scales about 5

mm long. Leaves to 1.2 m long, 1 -pinnate. Lamina

to 28 cm broad, firm-herbaceous to chartaceous,

gradually to abruptly reduced to a short, pinnatifid

apex, lacking scales, glabrous to minutely puber-

ulent on the axes abaxially, rachis and bases of

costae puberulent adaxially within the sulci. Ra-

chis deeply sulcate adaxially, lacking proliferous

buds. Pinnae commonly 7-10 pairs, well-spaced,

not or shortly stalked, broadest at or near the base,

acuminate at the apex, broadly cuneate to truncate

at the subequilateral base, margins entire to broad-

ly crenulate, larger pinnae 8-15 cm long and 2-

3.5 cm broad. Veins free, pinnately branched. Sori

borne on most of the vein branches, extending

from the costa to a few millimeters from the mar-

gin, usually diplazioid on the basal acroscopic

branch, the other sori single or diplazioid. Indusia

dull light brown, flat, about 0.2-0.4 mm broad,

the margins subentire to erose.

TYPE— Peru, Loreto, Gamitanacocha, Rio Ma-

zan, J. Schunke 281 (holotype, us!; isotypes, F!,

GH!, USM!).

In deep, wet forests, often in ravines, 100-1500

m, San Martin and Loreto to Madre de Dios and

Puno.

Venezuela; Ecuador; Peru; Bolivia; western Bra-

zil.

This variety differs from D. grandifolium var.

grandifolium in the delicate, narrow, dull light

brown indusia that do not change essentially in

color and form, whereas developing and mature

82

FIELDIANA: BOTANY

indusia of var. grandifolium are strikingly differ-

ent. Immature sporangia are protected by a broad

(often to 1 mm) bicolorous indusium, dark brown

to blackish and very firm at its attachment along

the vein, but thin and often hyaline toward the

margin. As sporangia mature, the indusium rolls

back tightly to reveal a dark, usually lustrous, un-

derside. This is nearly identical with the indusium

of D. celtidifolium, whereas that of var. andicola

may be compared with D. centripetale (Baker)

Maxon, of Venezuela, Ecuador and the West In-

dies.

Diplazium grandifolium var. grandifolium is

common in the West Indies, and from Central

America to Colombia, whereas var. andicola is

primarily Andean, although a single specimen has

been seen from Brazil, near the Peru-Bolivia bor-

der. The ranges overlap in Venezuela and Ecuador,

where a few specimens of each have been found.

San Martin: Prov. Mariscal Caceres, Tocache Nuevo,

/. Schunke V. 8489 (F, MO). Loreto: Santa Rosa, lower

Rio Huallaga below Yurimaguas, Killip & Smith 28911

(us). Huanuco: Prov. Huanuco, Tingo Maria, Tryon &

Tryon 5224 (BM, GH, us). Pasco (as Junin): Cahuapanas,

on Rio Pichis, Killip & Smith 26781 (us). Junin: Chan-

chamayo Valley, C. Schunke 142, 451 (F, us). Ucayali:

Prov. Coronel Portillo, Dist. Iparia, Bosque Nacional

de Iparia, /. Schunke V. 2687 (F, GH, us). Madre de Dios:

Prov. Tambopata, Tambopata Nature Reserve, Barbour

4963 (F, MO, uc). Puno: Prov. Carabaya, San Gaban,

Vargas 18934 (GH).

24. Diplazium celtidifolium Kunze, Bot. Zeit.

(Berlin) 3: 285. 1845. TYPE: Venezuela, Ca-

racas, Linden 544 (holotype LZ, destroyed;

isotypes, FI, K!, P!; photos, s of FI & K).

Plants terrestrial. Stem erect to decumbent, this

and petiole base provided with coarse, dark brown,

linear or linear- lanceolate scales 4-14 mm long.

Leaves 1-2 m long, 1 -pinnate. Lamina 30-60 cm

broad, herbaceous to subcoriaceous, terminating

in a short, pinnatifid apex, the axes glabrous or

densely puberulent and sparsely to amply provid-

ed on the abaxial side with flaccid, appressed,

brown scales, these mostly linear to filiform, es-

pecially abundant on the costae. Rachis deeply

sulcate adaxially, sometimes bearing proliferous

buds at the bases of distal pinnae. Pinnae 8-12

pairs, well spaced, conspicuously stalked, long-

acuminate at apex, broadly cuneate to truncate at

the subequilateral base, margins entire to shallow-

ly crenate-dentate, larger pinnae 12-26 cm long

and 2.5-7 cm broad. Veins free, pinnately

branched. Sori borne on most of the vein branch-

es, extending from the costa to about 5 mm from

the margin, always diplazioid on the basal acro-

scopic vein, the other sori single or diplazioid.

Indusia bicolorous, brown and blackish, at ma-

turity rolled back to the blackish, often lustrous

base.

Two varieties are recognized here.

Key to Varieties

a. Rachis and costae glabrous; leaves to 2 m long and 60 cm broad, chartaceous to subcoriaceous; larger

pinnae 1 7-26 cm long, 4-7 cm broad 24a. var. celtidifolium

a. Rachis and costae minutely but densely puberulent; leaves to 1 m long and 35 cm broad, herbaceous;

larger pinnae 12-17 cm long, 2.5-3.5 cm broad 24b. var. puberulum

24a. Diplazium celtidifolium var. celtidifolium.

Diplazium callipteris Fee, (Mem. foug. 5) Gen. fil. 2 1 4.

1852. PROBABLE TYPE: Venezuela, Funck &

Schlim 233, in 1845-1846 (holotype, not located;

isotype, BM!); originally cited in error as Linden

233 from Cuba (see discussion above).

Asplenium callipteris (Fee) Baker, Syn. fil. 231. 1867.

Asplenium celtidifolium (Kunze) Baker, Syn. fil. 232.

1867.

Athyrium celtidifolium (Kunze) Milde, Bot. Zeit. (Ber-

lin) 28: 353. 1870.

In wet forests, 500-1050 m, Amazonas, San

Martin, Puno.

Trinidad; French Guiana; Surinam; Venezuela;

Colombia; Peru; Brazil.

There has been confusion in the nomenclature,

synonyms, and typification of names associated

with D. celtidifolium. Apparently Fee erred when

citing the type of D. callipteris as Linden 233 from

Cuba. During studies of Diplazium in Peru, Lin-

den 233 could not be located. However, there is a

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

83

Funck & Schlim 233 from Venezuela (BM), marked

as an isotype, that matches the original description

in every way. Linden, half-brother to Schlim, col-

lected with him in Venezuela, and it is possible

names and numbers became mixed, bringing about

Fee's mistake in citing the type. Therefore, if fur-

ther search does not produce Linden 233, the Brit-

ish Museum specimen must be considered at least

an isotype. Furthermore, since this specimen does

not differ significantly from the type of D. celti-

difolium, the two names must be synonymous.

A number of specimens from the Lesser Antilles

are found in herbaria identified as D. callipteris.

However, all of these are apparently D. legalloi

Proctor, a similar species with coarser leaves, usu-

ally conspicuously crenate pinna margins, and del-

icate, light brown indusia.

Amazonas: Prov. Bagua, Montenegro-Chiriaco, Sa-

gdstegui 5923 (GH). San Martin: Mt. Guayrapurima,

Spruce 4760 (K, p, us). Puno: Prov. Carabaya, San Ga-

ban, Vargas 18933 (GH).

24b. Diplazium celtidifolium var. puberulum

Stolze, var. nov.

Varietas haec a varietate typica differt foliis usque ad

1 m longis et 35 cm latis, herbaceis, pinnis grandioribus

12-17 cm longis et 2.5-3.5 cm latis, rachidi et costis

dense puberulis.

Leaves to 1 m long and 35 cm broad, herbaceous

(not chartaceous to subcoriaceous). Larger Pinnae

12-17 cm long and 2.5-3.5 cm broad. Rachis and

costae densely puberulent on both sides.

TYPE— Peru, Pasco (as Junin), Pichis Trail, "Ya-

pas" [Yapaz], dense forest, 1350-1600 m, Killip

& Smith 25508 (holotype, us!; isotype, GH!; pho-

tos, F & GH of us).

Thus far known only from the type, Pasco, and

one paratype, cited below.

Although the most obvious difference between

this and var. celtidifolium is in the smaller and

thinner-textured leaves, this is not merely a de-

pauperate form of D. celtidifolium, for it differs

also in the puberulent axes. In the typical variety,

there is no laminar indument other than the scales

on the rachis and costae abaxially; but in var. pu-

berulum the axes are densely covered on both sides

with minute trichomes about 0.1-0.2 mm long.

Such trichomes are found with greater or lesser

frequency in others of the species complex, notably

such taxa as D. grandifolium, D. centripetale (Ba-

ker) Maxon of the West Indies and northern South

America, and D. eggersii Sodiro of Ecuador (the

latter most likely conspecific with D. centripetale,

see Comments). This is a character that needs more

attention in future monographic work on the ge-

nus.

Pasco (as Junin): Pichis Trail, "Yapas" [Yapaz], Killip

& Smith 25457 (us).

25. Diplazium paucijugum Stolze, sp. nov.

Caulis paleis crassis, nigellis; folium usque ad 65 cm

longum et 24 cm latum, 1-pinnatum, longipetiolatum,

parte apicali pinnatifida cerca dimidia longiores quam

lamina; rachis et costa in pagina adaxiali paleis flaccidis,

appressis, linearibus vel nliformibus; pinnae 2-4-juga-

tae, apicibus acuminatis, basibus subaequilateribus,

truncatis vel subcordatis; venae liberae, pinnatiramosae;

indusia crassa, bicoloria, brunnea et nigella, maturitatate

involuta.

Plants terrestrial. Stem erect to decumbent, pro-

vided with coarse, broad, blackish scales about 5

cm long. Leaves to 65 cm long, 1 -pinnate, on a

petiole nearly the length of the lamina. Lamina to

24 cm broad, firm-herbaceous, the pinnatifid api-

cal portion of the leaf about half the total length

of the lamina, essentially glabrous, but axes sparse-

ly puberulent abaxially, to glabrate, also provided

on the abaxial side with flaccid, appressed, brown

scales, these mostly linear to filiform, especially

abundant on the costae. Rachis deeply sulcate

adaxially, lacking proliferous buds. Free pinnae 2-

4 pairs at base of lamina, approximate to subdis-

tant, acuminate at apex, truncate to subcordate at

the subequilateral base, basal pair short-stalked,

margins entire, or basal pinnae broadly crenate,

larger pinnae 10-13 cm long, 3.5-4 cm broad. Veins

free, pinnately branched. Sori borne mostly on the

basal acroscopic vein branches, extending from

near the costa to about 5 mm short of the margin,

mostly single, occasionally diplazioid. Indusia

coarse, bicolorous, brown and blackish, at matu-

rity rolled back to the blackish base.

TYPE— Peru, "In monte Campana, prope Tara-

poto, Peruviae Orientalis" (Dept. San Martin),

Spruce 4339 (holotype, K!; isotype, P!; photos, F &

GH Of K & P).

Thus far known only from the type and one

other collection from Huanuco, in jungle on ridge

east of Tingo Maria, 625-1 100 m, cited below.

This species is distinguished from others in Peru

by the unusual configuration of the lamina. On a

84

FIELDIANA: BOTANY

petiole nearly as long as the lamina are borne a

few pairs of discrete pinnae, and the rest (half or

more) of the lamina is gradually reduced to a pro-

longed, pinnatifid apex. A similar condition occurs

in D. riedelianum (Kuhn) C. Chr. of Brazil and

(probably a synonym) D. verapax (Donn.-Sm.) Hi-

eron. of Mesoamerica. However, those taxa have

narrower laminae, much smaller pinnae with cu-

neate bases and long-tapering tips, and several

proliferous buds in the axils of proximal pinnae.

Diplazium paucijugum is closely related to D.

celtidifolium, especially in the narrow, flaccid scales

on the abaxial axes and the glabrous adaxial axes.

However, it differs from the latter species es-

pecially in the unusual lamina shape, the few and

smaller pinnae, the lack of proliferous buds and

by the minutely puberulent (to glabrate) axes abax-

ially. A proliferous bud is often found distally on

the rachis in D. celtidifolium, and axes are totally

lacking in indument except for the abaxial scales

on rachis and costae.

Huanuco (as San Martin): E of Tingo Maria, Allard

22334 (GH, us).

26. Diplazium roemerianum (Kunze) Presl, Tent,

pterid. 113. 1836. Figure 16b.

Asplenium roemerianum Kunze, Linnaea 9: 62. 1834.

TYPE: Peru, "in argillosis humidis ad Pampaya-

co" (Pampayacu, Huanuco), Poeppig 166, July,

1829 (holotype, B!; isotypes, B!, K!, L, P!; photos,

F, GH, MO & us of L).

Asplenium Jlavescens Mett., Ann. Sci. Nat. Bot., ser.

5, 2: 234. 1864, sp. nov. based on Hooker, Fil.

exot. /. 100. 1859, not Asplenium juglandifolium

Lam. TYPE: Cultivated at Kew from a Venezu-

elan plant collected by Wagener, this evidently

illustrated in /. 100 (holotype, K; isotype, BM!).

PARATYPE: Venezuela, Colonia Tovar, Fendler

498 (K). ISOPARATYPE: (GH!). Sloane, Hist. Jam.

"Filix maxima . . ." 82, t. 371 is excluded. Jen-

man, Bull. Bot. Dept. Jamaica n.s. 1(5): 77. 1894

says that the Sloane specimen at BM is a sterile

leaf of Acrostichum cervinum (= Olfersia cervina).

Diplazium flavescens (Mett.) Christ, Farnkr. Erde 217.

1897.

2-8 pairs, well-spaced, short-stalked, acuminate

at apex, broadly to narrowly cuneate at the sub-

equilateral base, the margins entire to crenulate-

serrate. Veins 1 -forked, free, the branching vein-

lets spreading from the costa at 50- to 70-degree

angles and gently arching to the pinna margin. Sori

mostly borne on the acroscopic vein branch, thus

gently arched like the veinlet. Indusia borne most-

ly along one side (occasionally on both sides) of a

veinlet, light to reddish brown, the margins erose

to subentire.

In wet forests, 700-2300 m, Loreto and Ama-

zonas south to Cuzco.

Greater Antilles; Guadeloupe; Venezuela; Co-

lombia; Ecuador; Peru; Brazil.

This and D. flavescens have been separated on

the basis of relative number of pinnae and serrate

vs. entire pinna margins: typical D. flavescens with

6-8 pairs of pinnae and crenulate margins; D. roe-

merianum with 1-2 pairs of pinnae and entire

margins. The two characters are inconsistent, and

there are no other significant differences. Pinnae

with entire margins are often serrate toward the

apex, and when many specimens are compared

throughout the range it becomes apparent that

margins are highly variable, whether on laminae

with few or many pinnae. It would appear that the

type of D. Jlavescens simply represents a more ro-

bust form of the species.

The Colombian Asplenium caucense Karsten and

A. ocanniense Karsten also surely belong here;

however the Central American taxon once clas-

sified as D. flavescens var. proliferum Christ has

been recognized (Stolze, 1981) as a quite distinct

species, D. obscurum Christ.

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

2647 (F, MO, uc). San Martin: Eastern Peru, near Tarapo-

to, Spruce 4674 (BM, P). Loreto: Sierra del Pongo, Mexia

627 3a (GH, K., uc, us). Huanuco: Fundo Chela, Sinchono,

Aguilar 920 (USM). Pasco: Pozuzo, Hacienda Ballisteros,

Bryan 672 (F, us). Junin: Chanchamayo Valley, C.

Schunke 8 (P), 72 (F, us), 73, 802 (F). Cuzco: Prov. La

Convencion, Guayanay, Vargas 13238 (GH).

Plants terrestrial. Stem erect or decumbent, this

and the petiole base provided with 3-7 mm long,

blackish, linear to lanceolate, subentire scales.

Leaves to over 1 m long, 1 -pinnate, long-petiolate.

Lamina to 35 cm broad, firm-herbaceous, termi-

nating abruptly in a nearly conform apical segment

(this occasionally with a large basal lobe), glabrous,

with scattered filiform scales on the rachis. Pinnae

27. Diplazium lechleri (Mett.) Moore, Index fil.

141. 1859.

Asplenium lechleri Mett., Fil. lechl. 1: 16, /. 2. 1856.

TYPE: Peru (Puno), "St. Gavan" (San Gaban),

"in sylvis montanis," Lechler 2269a (holotype,

B; isotypes, K!, L; photos, F, GH & us of L, us of

K).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

85

Plants terrestrial. Stem erect or decumbent, this

and the petiole base scaly, the scales 7-12 cm long,

dark brown, rigid, linear to lanceolate, attenuate,

subentire. Leaves to 2 m long, 1 -pinnate, long-pet-

iolate. Lamina to 35 cm broad, chartaceous to

subcoriaceous, terminating abruptly in a conform

apical segment, subglabrous, but axes (and some-

times the laminar surface) sparsely provided on

abaxial side with filiform scales that grade into

dark pluricellular trichomes. Pinnae 6-9 pairs,

subdistant, short-stalked, acuminate at apex,

rounded to cuneate at the subequilateral base (al-

though basal pinnae sometimes narrow-cuneate at

the basiscopic base). Veins simple, or paired at the

costa, straight, spreading at broad (75- to 80-de-

gree) angles, free, except connected at their tips by

a slightly inframarginal vein. Sori straight, crowd-

ed, and strictly parallel, mostly diplazioid. Indusia

dark brown, firm and persistent.

San Martin to Puno.

Costa Rica; Surinam; Venezuela; Colombia;

Peru; Brazil.

This species is easily distinguished by an apical

segment that is nearly identical to the lateral pin-

nae, the crowded, straight sori borne at nearly right

angles to the costa, and the mostly simple veins

that are free almost to the margin, where they join

an inframarginal vein.

San Martin: Mount Guayrapurima, Spruce 4687 (K,

p). Huanuco: SW slope of Rio Llullapichis watershed,

ascent of Cerros del Sira, Dudley 13085, 13381 (GH).

Junin: Pichis Trail, Porvenir, Killip & Smith 25922 (us).

Cuzco: Prov. Paucartambo, Cosnipata Valley, Rio Tono,

Wachter et al. 166 (F). Madre de Dies: Prov. Manu,

Cerro de Pantiacolla, Rio Palotoa, Foster et al. 107 05 -A

(F).

28. Dipla/ium plantaginifolium (L.) Urban, Symb.

antill. 4: 31. 1903.

Asplenium plantaginifolium L., Syst. nat. ed. 10, 2:

1323. 1759. TYPE: P. Browne, Jamaica (not lo-

cated). Proctor, Ferns of Jamaica, p. 394. 1985,

designated a (necessary?) neotype: Maxon 1949,

Jamaica, St. Catharine, Mt. Diablo (us!; isoneo-

type, BM!).

Asplenium plantagineum L., Sp. pi. ed. 2: 1537. 1763,

nom. superfl. for A. plantaginifolium L. and with

the same type.

Diplazium plantagineum (L.) Sw., J. Bot. (Schrader)

1800(2). 62. 1802.

Plants terrestrial. Stem erect or decumbent, this

and the petiole base sparsely provided with deep

reddish brown or blackish scales, these linear or

narrow-deltoid, 1-3 mm long. Leaves to 60 cm

long, simple, entire to sinuate, or crenate-serrate

apically, rarely bearing a proliferous bud at base

of lamina. Petiole nearly as long as, or longer than,

the lamina. Lamina to 6 cm broad, glabrous, lan-

ceolate to oblong-lanceolate, acuminate at apex,

broadly cuneate to truncate at base. Veins spread-

ing from the costa at 50-75 degrees, with 2-4 pairs

of strongly ascending branches running parallel

with the primary vein, to the leaf margin. Sori

(many of them) extending nearly from rachis to

margin and borne on both sides of a vein. Indusia

very narrow, delicate and often partly deciduous.

Southern Mexico to Panama; West Indies; Ven-

ezuela; Peru; Bolivia; Brazil.

In rain forests, 800-1800 m, Junin.

Only a few American species of Diplazium have

simple leaves, two of which occur in Peru; but

they are easily distinguished by the characters used

in the key.

Junin: East of Quimiri Bridge, near La Merced, Killip

& Smith 23971 (GH, us). Chanchamayo Valley, C.

Schunke 457 (F).

29. Diplazium pinnatifidum Kunze, Linnaea 9: 72.

1834. TYPE: Peru (Huanuco), "In crepidi-

nibus aquosis, lutosis, umbrosissimus," Cu-

chero, Poeppig diar. 1149 July, 1829 (holo-

type; w!; probably isotype, K!; photo, BM of w).

Anisogonium pinnatifidum (Kunze) Presl, Tent, pte-

rid. 116. 1836.

Asplenium kunzei Mett., Fil. hort. hot. Lips. 74. 1 856,

nom. nov. for Diplazium pinnatifidum Kunze, and

with the same type. Not Asplenium pinnatifidum

Nutt. 1818.

Plants terrestrial. Stem erect or decumbent, this

and the petiole base sparsely paleate, the scales 3-

4 mm long, dull brown, ovate or lanceolate, com-

monly subentire, but sometimes with castaneous

to blackish marginal cells that develop elongated

teeth that are bifid at apex. Leaves to 1 m long,

mature ones pinnatifid or pinnatisect, often pin-

nate at base, long-petiolate, juvenile ones suben-

tire to shallowly lobed. Lamina to 28 cm broad,

coriaceous, tapering gradually to a pinnatifid apex,

abruptly reduced at base, cut deeply to the rachis,

the rachis and costae usually concealed abaxially

by copious, amorphous, brown scales, and some-

times with a few, spreading, dark-toothed ones like

86

FIELDIANA: BOTANY

those of the stem. Segments 3-7 pairs, joined by

an acute to broadly rounded sinus (or the basal 1-

2 pairs sometimes fully discrete), the apex obtuse

to acute, the surfaces glabrous. Veins copiously

anastomosing from costa to margin. Sori dipla-

zioid, commonly branching to follow the veins.

Indusia dull brown, narrow, erose to fimbriate,

very delicate and early deciduous.

In forests, in wet, shady places, often on hillsides

and ravine banks, 250-2500 m, Amazonas and

Loreto to Cuzco and Madre de Dios.

Colombia; Ecuador; Peru; Bolivia.

The protologue ofDiplazium pinnatifidum con-

tains the exact citation of locality and habitat as

on the type label (above), except that the former

reads "Pampayaco" instead of Cuchero. Both sites

are in Huanuco.

The two areolate species of Diplazium in Peru

not only are easily distinguished by the characters

in the key; they differ also in their scales. In D.

praestans these are filiform and entire; in D. pin-

natifidum they are broad and often conspicuously

dentate, with dark setiform teeth. The areolate ve-

nation and these singular scales indicate close re-

lationship with D. aberrans Maxon & Morton and

D. pactile Lell. (Colombia and Ecuador) and D.

chimborazense (Baker) Christ and D. macrodic-

tyon (Baker) Diels (Ecuador). Some authors be-

lieve the characters help define a natural species

group or subgenus: Diplazium subgenus Aniso-

gonium.

Diplazium pactile differs from D. pinnatifidum

in the thin-textured and pubescent leaf tissue; D.

chimborazense has larger, thinner leaves, attenu-

ate pinnae, and the veins merge only toward the

pinna margin. Diplazium aberrans differs in its

simple, oblanceolate leaves, with petiole short or

lacking; D. macrodictyon has stellate trichomes on

the abaxial surface and veins. Another taxon, D.

fuscum (Baker) C. Chr. of Ecuador, seems to be

merely a robust form of D. pinnatifidum; an iso-

type and two authentic specimens at Kew are like

the latter in every respect, except that their larger

leaves have six or seven discrete pinnae, one pair

of which has lobed margins.

Amazonas: Prov. Bagua, E of La Peca, Barbour 2588,

2814 (F, MO, uc). San Martin: Mt. Guayrapurima, near

Tarapoto, Spruce 4686 (B, BM, GH, p). Loreto: Pumayacu,

between Balsapuerto and Moyobamba, Klug3218 (F, GH,

K, MO, us). Huanuco: Tingo Maria, Tryon & Tryon 5280

(BM, F, GH, uc, us). Pasco: Prov. Oxapampa, Puerto La-

guna, D. Smith 8448 (uc). Junin: Chanchamayo Valley,

C. Schunke 515 (F, us). Ucayali: Prov. Coronel Portillo,

Distr. Calleria, J. Schunke V. 3080 (F, GH, us). Cuzco:

Santa Isabel, Valle Cosnipata, Scolnik 930 (us). Madre

de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al.

10718 (F).

30. Diplazium praestans (Copel.) Morton, Contr.

U.S. Natl. Herb. 38: 41. 1967.

Athyrium praestans Copel., Amer. Fern J. 38: 132.

1948. TYPE: San Martin, Chazuta, Rio Huallaga,

Klug 4002 (holotype, us!; isotypes, F!, GH!, MO!,

P!, uc!).

Plants terrestrial. Stem erect or decumbent, oc-

casionally short-creeping, provided with medium

to dark brown, filiform scales, these 2-3 mm long

and only several cells broad, their margins entire.

Leaves to 35 cm long, simple, entire to broadly

sinuate, lacking a proliferous bud. Petiole 0-5 cm

long, with scales like those of the stem, but longer

and broader. Lamina to 9 cm broad, glabrous,

elliptic to (commonly) oblanceolate, subacute at

apex, long-attenuate at base. Veins copiously anas-

tomosing. Sori linear, gently arching, up to 5 cm

long, mostly single (rarely double). Indusia nar-

row, firm, persistent.

In dense forests, in wet places in deep shade,

often along banks of rivers and streams, 100-900

m, San Martin and Loreto to Junin and Madre de

Dios.

Peru; Bolivia; Amazonian Brazil.

Another species with simple leaves and copi-

ously anastomosing veins is Diplazium aberrans

Maxon & Morton of Colombia and Ecuador, but

this differs from D. praestans in its larger (to 60

cm) leaves and longer and narrower areoles. It

differs even more significantly in its peculiar scales,

which are borne along the costa, and sometimes

veins, abaxially, as well as on the stem and petiole.

These scales (at least of the stem) have castaneous

to blackish marginal cells that develop elongated,

bifid teeth. Scales on D. praestans (confined to the

stem and petiole) are entire and concolorous. For

further discussion of these scales see D. pinnati-

fidum.

San Martin: Prov. Mariscal Caceres, Distr. Campani-

lla, Quebrada de Mashuyacu, J. Schunke V. 4250 (F, GH,

us). Loreto: Rio San Alejandro, Woytkowski 5118 (GH,

MO, uc, us). Huanuco: Prov. Leoncio Prado, Rio Mon-

zon, near Bella, Plowman 5875 (GH). Junin: Rio Pinedo,

N of La Merced, Killip & Smith 23621 (GH, us). Ucayali:

Prov. Coronel Portillo, Distr. Iparia, Bosque Nacional de

Iparia, J. Schunke V. 2752 (GH, us). Madre de Dios:

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

87

Prov. Manu, Parque Nacional Manu, R. Foster 11722

(F).

Comments

Diplazium centripetale (Baker) Maxon, Pterido-

phyt. Porto Rico 441. 1926.

Asplenium centripetale Baker, Syn. fil. ed. 2: 490. 1874.

LECTOTYPE (designated by Proctor, Ferns of

Jamaica): Jamaica, Macfayden (K).

Asplenium eggersii Sodiro, Crypt, vase. Quit. 188.

1 893. TYPE: Ecuador, between Bodegas and Bal-

sapamba, Sodiro, (holotype, not located; probable

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

Diplazium eggersii (Sodiro) C. Chr., Index fil. 231.

1905.

This is very similar to D. celtidifolium var. pu-

berulum, especially in the pinna size and the pu-

berulent axes. It occurs in the West Indies, Ven-

ezuela, and Ecuador, and might be expected in

Peru. The most obvious differences from the latter

species are in the characters of sori and pinnae. In

D. centripetale, sori extend only halfway to % to

the pinna margin, and indusia are thin, flat, and

dull light brown throughout. The 15-25 pairs of

pinnae are commonly sessile. In D. celtidifolium,

sori extend from the costa to within 5 mm of the

pinna margin, and indusia are firm, bicolorous

(brown distally and blackish proximally), and at

maturity are rolled back to the blackish, often lus-

trous, base. There are only 8-12 pairs of pinnae,

most of which are conspicuously stalked.

In the protologue, the type of Asplenium eggersii

was said to be collected in Ecuador, between Bo-

degas and Balsapamba. Thus far no specimen has

been found exactly designating this locality. How-

ever, there are two sheets at Paris collected by

Sodiro in January, 1891 "in reg. tropic, inter Bo-

degas et Pisagua," one of which is inscribed in

Sodiro's hand as "Asplen. (Dipl.) eggersii, Nov.

Sp." Given the confusion that usually attends So-

diro specimens, descriptions, and type localities,

this is likely to be an isotype (if not the holotype)

of the name. In any case, these specimens at Paris,

and other Ecuador collections made in adjacent

areas and determined as D. eggersii, quite match

specimens of D. centripetale from the West Indies.

Key to Species of Athyrium

a. Lamina 2-pinnate to 2-pinnate-pinnatifid; ultimate segments rather sharply dentate

1 . A. dombeyi

a. Lamina 4- to 5-pinnate; ultimate segments entire and elongate, bifid, or lobed . . 2. A. ferulae-emu

XVII. Athyrium

Athyrium Roth, Tent. fl. germ. 3: 58. 1 799. TYPE:

Athyrium filix-femina (L.) Roth (Polypodium

filix-femina L.). Figure 17.

Stem usually decumbent to nearly erect, usually

short-creeping and moderately stout, or long-

creeping and slender, bearing scales, these not or

obscurely clathrate, brown, concolorous. Leaves

ca. 20 cm to 2 m long, petiole continuous. Lamina

1 -pinnate-pinnatifid to 5-pinnate, nearly glabrate,

somewhat scaly, or slightly pubescent or glandular,

monomorphic or nearly so. Veins free. Sori round-

ish or usually elongate on one side of a vein, or

sometimes also on the other side distally, not pa-

raphysate, usually covered by reniform to elongate

or hook-shaped indusia, or essentially exindusiate.

Spores rather ellipsoidal, monolete, variously

ridged.

Athyrium is a large genus of about 100 species,

most of them in eastern and southeastern Asia.

There are two species in the Andes and in Peru.

The genus is technically separated from Dipla-

zium by its chromosome number of n = 40 or

multiples, and the indusia either only on one side

of a vein, or if on the other side then only distally

and continuous around the apex of the sorus. Di-

plazium has a chromosome number of n = 41 and

multiples, and usually some sori are on both sides

of a vein and distinct distally.

Athyrium ferulaceum and related species of Cen-

tral America are placed here on the basis of the

chromosome number of n = 40 (Gomez, Atti 1st.

Bot. Univ. Pavia 7: 30. 1971, as "Athyrium bra-

dearum") (= Diplazium bradeorum), and the in-

dusia that are commonly only on one side of a

vein, or rarely partly on the other side and con-

tinuous distally. The group is of uncertain affinity

and the spores resemble species of Diplazium more

than those of Athyrium. However, it seems closer

to Athyrium in its chromosome number and more

obvious characters.

88

FIELDIANA: BOTANY

FIG. 17. Athyrium dombeyi: a, habit; b, pinnule, abaxial side. (From Tryon & Tryon 6029, Columbia, GH.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

89

1. Athyrium dombeyi Desv., Mem. Soc. Linn.

Paris 6: 266. 1827. (as dombei). TYPE: "Pe-

ruvia (Herb. Mus. Paris)," presumably Peru,

Dombey (holotype, P). Figure 17a-b.

Stem stout, decumbent to erect, bearing brown

to dark brown, elongate scales. Leaves ca. 20 cm

to 1 m long, the petiole mostly glabrous except

for usually persistent scales near the base. Lamina

2-pinnate to 2-pinnate-pinnatisect, glabrous or

very slightly scaly, pinnae sessile to usually short-

stalked, the basal usually somewhat reduced, ul-

timate segments rather sharply dentate. Sori more

or less elongate, on one side of a vein, rarely some

on the other side distally, covered by an indusium

of similar extent.

In ravines, on damp banks, along streams, and

at the edge of aqueducts, 2500-3300 m, Lamba-

yeque to Cuzco.

In South America southward to southeastern

Brazil and Argentina; perhaps also northward to

Central America, Mexico, and Hispaniola.

Athyrium dombeyi is a member of the Athyrium

filix-femina complex which is mostly temperate

and boreal in distribution and it is perhaps a va-

riety of that species: Athyrium filix-femina var.

dombeyi (Desv.) Hieron.

Lambayeque: Prov. Ferrenafe, 4 km NW of Incahuasi,

Dillon & Skillman 4157 (F, GH). Cajamarca: Prov. Ca-

jamarca, Dist. Encanada, Sanchez 318 (GH). Amazonas:

Prov. Bagua, E of La Peca, Barbour 2759 (F). La Libertad:

Prov. Otuzco, above Agalpampa, Sounders 894 (GH),

595 (F, GH). San Martin: Dist. Huallaga, 30 km above

Jucusbamba, Hamilton & Holligan 929 (us). Prov. Mar-

iscal Caceres, between Mirador and La Playa, Young &

Leon 4932 (F). Ancash: Above Yungay on road to Huan-

ganuco, Correll & Smith P963 (GH). Huanuco: Mima,

Macbride 4327 (F, us). Cani, Macbride 3404 (F, us). Cuz-

co: Prov. Paucartambo, vicinity of Achirani, Vargas

11139(F,vc).

pinnae sessile to short-stalked, the basal the larg-

est, ultimate segments entire and elongate, bifid,

or lobed. Sori elongate, on one side of a vein, rarely

some on the other side distally, covered by an

indusium of similar extent.

A single collection known from Peru: in humus,

1450 m, Cuzco. Elsewhere in forests, on ravine

banks, and rarely on sandstone cliffs, 100-2700 m.

Central America south to Peru.

Cuzco: Prov. La Convention, Choquellowanca, Var-

gas 12928 (GH).

XVIII. Hemidictyum

Hemidictyum Presl, Tent, pterid. 1 10. 1836. TYPE:

Hemidictyum marginatum (L.) Presl (Asple-

nium marginatum L.). Figure 18.

Stem erect to decumbent, stout, bearing scales.

Leaves ca. 1 to 3 m (or more) long, petiole con-

tinuous. Lamina 1 -pinnate, the pinnae entire, gla-

brous. Veins free near the costas, fully anasto-

mosing toward the margin, without included free

veinlets, the vein ends connected by a marginal

vascular strand. Sori elongate along the veins, not

paraphysate, covered by elongate indusia. Spores

rather ellipsoidal, monolete, prominently ridged

or saccate.

Hemidictyum is a monotypic, morphologically

and cytologically isolated genus of the American

tropics. The chromosome number of n = 31 may

raise doubts as to its proper inclusion in the

Dryopteridaceae, which have n = 40 or 4 1 . How-

ever, in other characters, the alliance of the genus

seems to be with that family and with the Tribe

Physematieae.

2. Athyrium ferulaceum (Hooker) Christ, Bull.

Herb. Boissier, ser. 2, 4: 968. 1904.

Asplenium ferulaceum Hooker, Sp. fil. 3: 216. 1860.

SYNTYPES: Colombia (New Grenada), Hartweg

1519; Ecuador, Quito, Jameson (both K).

Diplazium ferulaceum (Hooker) Lell., Proc. Biol. Soc.

Wash. 98: 376. 1985.

Stem small to stout, decumbent to erect, bearing

broad, brown scales, especially at the apex. Leaves

ca. 30 cm to 1 m long, the petiole essentially gla-

brous. Lamina 4-5 pinnate, glabrous or nearly so,

1. Hemidictyum marginatum (L.) Presl,

pterid. 111. 1836. Figure 18a-c.

Tent.

Asplenium marginatum L., Sp. pi. 1082. 1753. TYPE:

Either LINN 1250.20 or Petiver, Pteri-graph.

Amer. /. 12, f. 2, both of which are this species,

the latter based on Plumier, Traite foug. Amer.

t. 106.

Stem apex densely scaly, with brown, linear

scales. Leaves ca. 1 to 3 m (or more) long, the

petiole glabrous or somewhat scaly near the base.

Lamina 1 -pinnate, with a conform apical segment,

90

FIELDIANA: BOTANY

FIG. 18. Hemidictyum marginatum: a, leaf apex; b, portion of pinna, abaxial side; c, pinna margin, adaxial

side. (From Stolze, Ferns & fern allies of Guatemala, 1981.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

91

pinnae large, entire, thin. Veins with the ends con-

nected by a nearly marginal vascular strand. Sori

elongate, often very long, borne along part of a

free vein, toward the costa from the anastomosing

veins, indusium of similar extent, very thin.

Steep banks in forests, in dense forests and in

cloud forests, 260-1 700 m, Amazonas to Ucayali.

Southern Mexico and the Greater Antilles, south

to Bolivia and southeastern Brazil.

Amazonas: Prov. Bagua, Quebrada Mirana, Rio Ma-

ranon, above Cascadas de Mayasi, Wurdack 1890 (GH,

us). Prov. Bagua, 12 km E of La Peca, Barbour 2582

(MO). San Martin: Tarapoto, Spruce 4783 (GH, us). Cha-

zuta, Rio Huallaga, Klug 3932 (GH, us). Loreto: Between

Rio Amazonas and Rio Napo, Croat 19556 (MO). Hua-

nuco: Prov. Huanuco, Tingo Maria, Tryon & Tryon 5254

(GH, us). Prov. Huanuco, Gasa, Stork & Morton 9874

(F, us). Pasco: Pichis Trail, Enenas, (as Junin), Killip &

Smith 25765 (F, us). Prov. Oxapampa, Paujil, Leon 285

(USM). Prov. Oxapampa, Villa Rica, van der Werffet al.

8304 (MO). Junin: Near La Merced, Killip & Smith 23875

(F, GH, us). Near Perene Bridge, Killip & Smith 25291

(F, us). Ucayali: Prov. Coronel Portillo, La Divisoria,

(as Loreto), /. Schunke V. 10199 (F, MO, us). Madre de

Dios: Prov. Manu, Cocha Cashu uplands, Nunez 5841

(MO).

XIX. Cystopteris

Cystopteris Bernh., Neues J. Bot. (Schrader) 1(2):

26. 1806, nom. conserv. TYPE: Cystopteris

fragilis (L.) Bernh. (Polypodium fragile L.).

Figure 19.

Stem short and decumbent to rather long-creep-

ing, bearing scales. Leaves ca. 10-80 cm long,

petiole continuous. Lamina 2-pinnate to 3-pin-

nate-pinnatifid, glabrous, glandular or slightly pu-

bescent, monomorphic. Veins free. Sori round,

borne on the veins, not paraphysate, more or less

covered by scalelike to half-cup shaped indusia.

Spores rather ellipsoidal, monolete, echinate, ver-

rucate, slightly rugose, or with inflated processes.

Cystopteris is a small genus of about six species,

nearly worldwide in distribution, with a single spe-

cies, C. fragilis, in the American tropics. The An-

dean Cystopteris is highly variable and may even-

tually be classified as various taxa representing

different cytotypes. At this time it seems best to

treat it as a single species. BlasdelFs interpretation

of variation in C. fragilis as being due to wide-

spread hybridization seems to lack evidence and

merit; accordingly the morphological variations

are treated as within the natural variation of one

species.

There are two subgenera, Cystopteris (including

C. fragilis) and submenus Acystopteris(Nakai) Bias-

dell, the latter often recognized as a genus.

Reference

BLASDELL, R. F. 1963. A monographic study of

the fern genus Cystopteris. Mem. Torrey Bot.

Club, 21(4): 1-102.

1. Cystopteris fragilis (L.) Bernh., Neues J. Bot.

(Schrader) 1(2):27, t. 2, f. 9. 1806. Figure

19a-b.

Polypodium fragile L., Sp. pi. 1091. 1753. TYPE:

Plunkenet, Phytographia t. 180, f. 5. 1691.

Cystopteris translucens Desv., Mem. Soc. Linn. Paris

6: 264. 1827. TYPE: Peru, (holotype, P, Herb.

Desvaux; photos, GH, us).

Athyrium fumaroides Presl, Reliq. haenk. 1: 39, /. 6,

f. 2. 1825. TYPE: Peru, Haenke (holotype, PR or

PRC).

Stem scales light to dark brown, usually rather

broad. Leaves ca. 10-50 cm long, the petiole usu-

ally glabrous, sometimes slightly scaly. Lamina

1-pinnate-pinnatisect to 3-pinnate-pinnatifid,

gradually reduced at the apex, essentially glabrous,

pinnae nearly sessile to short-stalked, the basal

somewhat reduced or not. Veins free. Sori round-

ish, borne on the veins, covered by a basally at-

tached, scalelike indusium, its apex often some-

what elongate.

Usually in rocky, locally moist places: on ledges,

in crevices of rocks, on rock walls, and on rocky

roadside banks, also along streams, on shrubby

hillsides, and banks of irrigation ditches, rarely in

wet forests or rarely epiphytic, 300—4500 m, Ca-

jamarca to Puno. This is a widely distributed spe-

cies in Peru, especially at 3000-3500 m; the lowest

altitude for the species, 300-700 m, is on the Lo-

mas de Atiquipa, Department of Arequipa.

Widely distributed in both hemispheres; pri-

marily montane in the tropics.

Cajamarca: Arriba de Incahuasi, Sagdstegui et al. 12894

(F, GH, HUT). Prov. Contumaza, Lleden, Sagdstegui 9392

(MO). Amazonas: Rio Ventilla, near Molinopampa, Wur-

dack 1548 (GH, us). La Libertad: Prov. Santiago de Chu-

co, Motil to Shorey, Sounders 891 (F, GH). Prov. Sanchez

Carrion, Huayllides, D. Smith 2275 (MO). Ancash: Prov.

92

FIELDIANA: BOTANY

FIG. 19. Cystopteris fragilis: a, habit; b, pinnule, abaxial side. (Adapted from Stolze, Ferns & fern allies of

Guatemala, 1981.)

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

93

Huaylas, Huascaran National Park, Smith et al. 9786

(F). Prov. Bolognesi, Chiquian, Mostacero et al. 1342

(MO). Huancuco: Chaglla, Macbride 3638 (F, GH, us).

Lima: Viso, Macbride & Featherstone 613 (F, us). Prov.

Canta, Dist. Huamantanga, Sounders 1149 (f, GH). Prov.

Yauyos, Tupe, Cerrate 1071 (USM). Pasco: Prov. Oxa-

pampa, Cordillera de Yanachaga, van der Werff et al.

8470 (MO, uc). Junin: Prov. Tarma, 5 km SW of Hu-

acapistana, Tryon & Tryon 5432 (GH, us). Huancavelica:

Entre Conaica y Manta, Tovar827 (GH). Ayacucho: Nas-

ca to Puquio, Correll & Smith PI 56 (GH). Apurimac:

Prov. Abancay, Vargas 2296 (uc). Cuzco: Prov. Cuzco,

between Cuzco and Sagsaywaman, Stevens 22070 (F, MO).

Machu Picchu, Leon 465 (USM). Yucay, Coronado 144

(GH). Arequipa: Lomas de Atiquipa, Coronado 37 (GH,

uc, us); ca. 100 km N of Arequipa, Treacy 694 (GH).

Puno: 10 km from Puno, Correll & Smith PI 87 (GH).

XX. Woodsia

Woodsia R. Br., Prodr. 158. obs. IV sub Alsophila.

18 10, as Woodia; also and corrected in Trans.

Linn. Soc. London 1 1 : 1 7 1-1 74. 1 8 1 5. TYPE:

Woodsia ilvensis (L.) R. Br. (Acrostichum il-

vense L.). Figure 20.

Stem decumbent to erect, usually small and

short-creeping, bearing scales. Leaves ca. 5-50 cm

long, the petiole joined in a few (not South Amer-

ican) species. Lamina 1 -pinnate or 1 -pinnate-pin-

natisect or rarely to 2-pinnate-pinnatifid, glabrous,

glandular, scaly, and/or pubescent. Veins free. Sori

roundish, borne on the veins, not paraphysate,

more or less covered by an indusium surrounding

the receptacle that is globose, or of a few large

segments, or of a few trichomelike cilia. Spores

ellipsoidal to spheroidal, monolete, with short to

long winglike ridges and somewhat echinate.

Woodsia is mostly a circumboreal genus of about

25 species, with a single species in South America

and Peru.

Cheilanthes crenata Kunze, Linnaea 9: 84. 1834.

TYPE: Peru, (Huanuco), Huanuco, Poeppig in

1830 (holotype, LZ, destroyed; isotype, not locat-

ed).

W oodsia peruviana Hooker, Sp. fil. 1 : 6 1 , /. 2 IB. 1 844.

TYPE: Peru, (Lima), Huamantanga, Mathews 602

(holotype, K; isotype, GH!).

Woodsia crenata (Kunze) Hieron., Bot. Jahrb. Syst.

34: 440. 1904.

Stem with light brown to dark brown, usually

broad scales. Leaves ca. 8-40 cm long, the petiole

glabrous or often slightly; scaly, especially at the

base. Lamina 1-pinnate-pinnatifid to usually

1 -pinnate-pinnatisect, rarely 2-pinnate-pinnatifid,

gradually reduced at the apex, glandular-pubes-

cent and pubescent, pinnae sessile or nearly so,

the basal ones reduced. Veins free. Sori roundish,

borne on a vein, at first covered by a nearly glob-

ular indusium that later splits into segments.

In rocky places, on cliffs, in crevices of rocks, at

the bases of large rocks, or on Inca walls, also on

brushy rocky slopes, on pastured hillsides, grassy

slopes, or stream banks, rarely in wet woods or on

wooded hillsides, 200-4300 m, Lambayeque to

Arequipa and Puno. The species grows at 200-600

m on coastal lomas and usually at 3000—4000 m

in the Altiplano.

Hispaniola; Venezuela and Colombia, south to

Argentina and to southeastern Brazil; southern Af-

rica.

Woodsia motevidensis is a highly variable spe-

cies in the size of the lamina and the extent to

which the margin is flat or revolute. At high al-

titudes, especially in exposed sites, the leaves may

be less than 1 0 cm long and the ultimate segments

have revolute margins. Especially on lomas, which

are mostly cloudy and foggy during the growing

season, the leaves may be to 40 cm long and the

ultimate segments of the lamina are expanded and

flat.

Reference

BROWN, D. F. M. 1964. A monographic study

of the fern genus Woodsia. Beih. Nova Hed-

wigia, 16: 1-154.

1 . Woodsia montevidensis (Sprengel) Hieron., Bot.

Jahrb. Syst. 22: 363. 1896. Figure 20a-d.

Dicksonia montevidensis Sprengel, Syst. veg. ed. 16,

4: 122. 1827. TYPE: Uruguay, "Monte Video,"

Sello (a specimen at B is authentic).

Lambayeque: 42 km from Olmos on road to Jaen,

Correll & Smith P816 (GH). Cajamarca: Prov. Contu-

maza, arriba de Contumaza, Lopez et al. 37 13 (F, GH).

Prov. Contumaza, alrededores de Guzmango, Sagdste-

gui 9675 (MO). La Libertad: Lomas de Viru, Coronado

284 (GH, uc), Lopez et al. 3638 (GH). Ancash: Lomas de

Mongon, Coronado 299 (GH). Cerro al E de Chiquian,

Ferreyra 7343 (GH). Huanuco: Piedra Grande, Macbride

3671 (F, GH, us). Muna, Woytkowski 5252 (GH, MO).

Lima: Loma de Amancaes, Tryon & Tryon 5215 (F, GH,

us), Coronado 12 (MO). Matucana, Macbride & Feath-

erstone 427 (F, us). Junin: Near Huancayo, Killip & Smith

22127 (GH, us). Huacapistana, Cerrate 2884 (GH). Huan-

cayo, Soukup 3549 (MO). Huancavelica: Entre Colca-

bamba y Paucarbamba, Tovar 2008 (GH). Ayacucho: Be-

94

FIELDIANA: BOTANY

a \

FIG. 20. Woodsia montevidensis: a, habit; b, stem scale; c, pinna segment, abaxial side. (From Bryan 375, F.)

tween Huanta and Rio Apurimac, Killip & Smith 22485 Amparaes, Nunez 6642, 6649 (MO). Slopes of Sicuani,

(GH, us). Apurimac: Prov. Aymaraes, Dist. Cotarosi, Vargas 9831 (F, GH, uc). Arequipa: Arequipa, Pennell

Saunders 775 (GH). Cuzco: Machu Picchu, Tryon & Tryon 13197 (GH, us). Puno: Puno, Graja Salcedo, Mexia 7781

5402 (GH, us), Coronado 96 (GH, uc, us). Prov. Calca, (GH, uc, us). Chucuito, Coronado 165 (GH, uc).

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

95

XXI. Oleandra

Oleandra Cav., Anales Hist. Nat. 1:115. 1799.

TYPE: Oleandra neriiformis Cav. Figure 21.

Stem erect and branching, or climbing and long-

creeping, slender, to 5 m long, bearing scales.

Leaves ca. 10-50 cm long, petiole articulate. Lam-

ina simple, entire, glabrate, pubescent, and/or scaly,

monomorphic or very rarely dimorphic. Veins free.

Sori roundish, borne on the veins, not paraphy-

sate, covered by a reniform to peltate indusium.

Spores rather spheroidal, monolete, more or less

prominently ridged and echinate.

Oleandra is a pantropical genus of about 35 spe-

cies. There are at least six species in America and

three in Peru. It is one of the few ferns that have

some species with an erect, branched stem that

results in a shrublike habit.

The petiole is articulate and the portion of it

below the joint is called a phyllopodium.

Reference

MAXON, W. R. 1914. The American species of

Oleandra. Contr. U.S. Natl. Herb., 17: 392-398.

Key to Species of Oleandra

a. Stem widely creeping, its scales spreading; phyllopodia mostly 5-30 mm long, naked, like the petiole

above the joint; lamina and indusia glabrous 1. O. articulata

a. Stem ascending, erect or climbing, its scales closely appressed b

b. Lamina glabrous or with some scales, mostly 1-2 cm wide, rarely some to 2.5 cm or less than 1

cm wide; phyllopodia mostly 10-20 cm long, or some as short as 3 mm, or as long as 30 mm,

naked, like the petiole above the joint; indusia glabrous 2. O. lehmannii

b. Lamina pubescent, at least in part, especially on or near the costa, mostly 3-3.5 cm wide, rarely

some 2 cm or more than 3.5 cm wide; phyllopodia 1-3 mm long, or some to 6 mm long, at first

scaly, like the stem; indusia slightly pubescent on the surface and with ciliate margins

3. O. pilosa

1. Oleandra articulata (Sw.) Presl, Tent, pterid.

78. 1836. Figure 21a-b.

Aspidium articulation Sw., J. Bot. (Schrader) 1800(2):

38. 1 802. TYPE: Martinique, Plumier, Traite foug.

Amer. t. 136.

Aspidium nodosum Willd., Sp. pi. ed. 4, 5: 21 1. 1810,

nom. superfl. for Aspidium articulatum and with

the same type.

Oleandra nodosa Presl, Tent, pterid. 78. 1836, nom.

nov. for Aspidium nodosum Willd. and with the

same type.

Stem widely creeping, with spreading, usually

widely spreading, scales. Phyllopodia 5-30 mm

long, slender, without scales. Lamina glabrous, ca.

15-40 cm long and 2.5-8 cm wide. Indusia gla-

brous.

Epiphytic in dense forests or cloud forests, 1 330

to ca. 1 700 m, Huanuco, pasco, and Junin.

Guatemala to Panama; West Indies; northern

South America south to Peru and northern Bra-

zil; Cocos and Galapagos Islands.

Rarely the scales of the stem are mostly only

slightly spreading, rather than widely spreading.

Maxon chose Plumier t. 136, the only element

included in the original publication of Aspidium

articulatum Sw., as the type of the name. He in-

vestigated material from Mauritius, included in

Aspidium articulatum later by Swartz (Syn. fil. 42,

236. 1805) in the Swartz herbarium, Stockholm,

and decided that it could not be type material of

Aspidium articulatum. Morton (Amer. Fern J. 58:

105-107. 1968) reached a contrary conclusion and

applied the name Aspidium nodosum to the Amer-

ican species and Aspidium articulatum to the spe-

cies of Africa and adjacent islands. Joncheere

(Taxon 18: 538-541. 1969) argued persuasively

that Maxon's original typification was correct and

we confidently agree with his conclusion. The cor-

rect name for the African species is Oleandra dis-

tenta Kunze (Pic.-Ser., Webbia 20: 755. 1965).

Huanuco: Rio Llullapichis watershed, ascent of Cerros

del Sira, Dudley 13243 (GH). Pasco: Pichis Train, Enenas

(as Junin), Killip & Smith 25756 (F, NY, us). Prov. Oxa-

96

FIELDIANA: BOTANY

rid

•• t*

i , s.mm,

2cm

FIG. 21. Oleandra articulata: a, habit; b, portion of lamina, abaxial side. Oleandra lehmannii: c, stem and

phyllopodium. (a, b from Ollgaard & Balslev 9085, Ecuador, F; c from D. Smith 5395, F.)

pampa, Cordillera San Matias, Leon 313 (GH, USM).

Junin: Above San Ramon, Killip & Smith 24878 (NY,

us).

2. Oleandra lehmannii Maxon, Contr. U.S. Natl.

Herb. 17:395. 1 9 14. TYPE: Colombia, Antioquia,

above Amalfi, Lehmann XLII (holotype, us!; iso-

type, K!; photo, GH). Figure 21c.

Stem ascending to erect and climbing, with

closely appressed scales. Phyllopodia (3-)10-

20(-30) mm long, mostly slender, without scales.

Lamina glabrous or with some scales, ca. 10-30

cm long and l-2(-2.5) cm, rarely less than 1 cm,

broad. Indusia glabrous.

Epiphytic or rarely terrestrial in rich organic

matter, montane rainforests, elfin forests, rarely in

Sphagnum bogs, 1480-2500 m, Huanuco and Pas-

co.

Colombia south to Peru; northern Brazil.

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

97

Huanuco: Rio Llullapichis watershed, ascent of Cerros

del Sira, Dudley 13306B, 13453, 13489 (OH), 1 3554 (GH,

us). Pasco: Pichis Trail, Dos de Mayo (as Junin), Killip

& Smith 25870 (F, GH, NY, us). Prov. Oxapampa, 20 km

W of Oxapampa, D. Smith 5395 (GH). Chontabamba,

Ledn et al. 968 (F, GH).

capata, de Lantreppe (NY). Prov. Paucartambo, San Pe-

dro, Vargas 6801 (uc, us). Puno: Tatanara, Lechler2539

(B, K).

Comments

3. Oleandra pilosa Hooker, Gen. fil. t. 45B. 1840.

TYPE: "British Guiana," Berbice, Schom-

burgk 416 (holotype, K!; photo, GH).

Aspidium pendulum Splitg. Tijdschr. Natuurl. Gesch.

Physiol. 7: 412, 1840, not Raddi, 1819. SYN-

TYPES: Surinam, Berlijn plantation, Splitgerber

(L; photo, GH); Schomburgk 416 (K, L; photo GH

ofK).

Oleandra micans Kunze, Bot. Zeit. (Berlin) 9: 346.

1851. TYPE: Peru, (San Martin), Mission To-

cache, (sterile), PoeppigM. Aug. 1830(Z)/ar. 7955)

(holotype, LZ, destroyed). Data from Linnaea 9:

89. 1834. Tracing in B! Herb. Mett. of "Oleandra

micans Kunze, Peruvia Poeppig."

Stem ascending to erect and climbing, with

closely appressed scales. Phyllopodia l-3(-6) mm

long, stout, at first scaly. Lamina pubescent, es-

pecially on or near the costa, ca. 10-40 cm long,

and (2-)3-3.5(-6) cm broad. Indusia slightly pu-

bescent on the surface and with ciliate margins.

Epiphytic in forests, 325-1000 m, San Martin

to Puno.

Trinidad and northern South America, south to

Peru.

Oleandra pilosa and O. lehmannii are the only

species in Peru with appressed scales on the stem.

The variation in these species makes it impossible

to determine the identity of the type of Oleandra

micans from the original description and the trac-

ing at B. No isotype has been seen. However, col-

lections of O. pilosa in Peru are from 325-1000

m, while those of O. lehmannii are from 1480-

2500 m. Since Tocache is at an altitude of about

500 m, O. micans is treated as a synonym of O.

pilosa.

Huanuco: Tingo Maria (as San Martin), Allard 21215,

21999 (GH). Prov. Leoncio Prado, E of Tingo Maria, J.

Schunke V. 5194 (GH, MO, us). Pasco: Prov. Oxapampa,

Palcazu valley, D. Smith 3807 (GH). Cuzco: Rio Mar-

Oleandra hirta Brack., U.S. Expl. exped. 16 (fil.):

214. 1854. Atlas, t. 29. 1855. TYPE: Brazil,

Organ Mountains, U.S. Expl. Exped. (us).

This species is found primarily in Brazil, but it

has been collected in Bolivia (Tate 1152, NY) and

may occur in southern Peru. The stem and scales

are like those of O. articulata but the lamina is

pubescent and the indusia are usually slightly pu-

bescent on the surface or somewhat ciliate. The

shape of the lamina is variable, as it is in O. pilosa

and other species of the genus. It may be gradually

attenuate basally and more or less acuminate api-

cally (type of O. hirta), or it may vary to narrowly

cuneate to broadly rounded basally and acute to

caudate apically.

XXII. Bolbitis

Bolbitis Schott, Gen. fil. 1. 13 (fasc. 3). 1835. TYPE:

Bolbitis serratifolia (Kaulf.) Schott (Acrosti-

chum serratifolium Kaulf.). Figure 22.

Plants terrestrial to epipetric, sometimes be-

coming hemiepiphytic. Stem short- to long-creep-

ing, or scandent well up tree trunks, slender to

rather stout, moderately to densely provided with

light brown to blackish, subclathrate scales and

with few to many fibrous roots. Leaves strongly

to weakly dimorphic, ca. 25 cm to 2 m long, ap-

proximate to widely spaced, petiole continuous

with the stem. Lamina simple and entire to

1 -pinnate-pinnatifid, sometimes imparipinnate,

the pinnae not articulate to the rachis, glabrate to

somewhat scaly, the rachis or apical segment often

bearing a proliferous bud. Veins areolate, with or

without included free veinlets, or (in a few species

not in Peru) free. Sporangia commonly borne over

the abaxial surface of the segment, lacking pa-

FIG. 22. Bolbitis serrata: a, habit; b, portion of pinna, abaxial side. Bolbitis serratifolia: c, portion of pinna,

abaxial side. B. lindigii: d: portion of pinna, abaxial side, (a, b from Killip & Smith 28495, us; c from C. Schunke

161, F, d from Killip & Smith 25184, F.)

98

FIELDIANA: BOTANY

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

99

raphyses. Indusium lacking. Spores monolete, more

or less ellipsoidal.

Bolbitis is a pantropical genus of 44 species, in-

cluding 1 4 in America. It is allied to Lomagramma

and Lomariopsis, especially in the large, dimor-

phic leaves, which are borne on a stem that some-

times climbs well up the trunks of trees, and the

acrostichoid pattern of sporangia. Venation is typ-

ically areolate, although veins of several Asian spe-

cies are free. It differs from both allied genera in

that the pinnae are not articulate to the rachis.

Species in Peru can be further distinguished from

Lomariopsis by their anastomosing veins. This

treatment is adapted in large part from the mono-

graph of Hennipman (1977).

Reference

HENNIPMAN, E. 1977. A monograph of the fern

genus Bolbitis (Lomariopsidaceae). Leiden Bot.,

ser., 2: 1-331.

Key to Species of Bolbitis

a. Sterile lamina pinnate and gradually reduced to a nonconform, usually pinnatifid, apex

1 . B. aliena

a. Sterile lamina simple, or pinnate and abruptly reduced to a discrete (usually conform) apical segment

b

b. Free veinlets abundant, spreading in various directions within the areoles c

c. Sterile lamina lacking proliferous buds; adult sterile lamina commonly with an apical segment

and 3-8 pinnae (very rarely simple or with 2 pinnae) 4. B. nicotianifolia

c. Sterile lamina either with a subterminal or an axillary proliferous bud; adult sterile lamina

simple, subternate or ternate d

d. A proliferous bud borne in the pinna axils (or on a pinna stalk); sterile lamina ternate or

subternate; pinnae often to 20 cm long and 12 cm broad 5. B. oligarchica

d. A proliferous bud borne at or near the lamina apex; sterile leaf usually simple, occasionally

subternate; pinnae (when present) to 8 cm long and 4 cm broad 7. B. pandurifolia

b. Free veinlets lacking within the areoles or, when present, directed either distally (toward the pinna

margin) or proximally (toward the costa), but not or rarely in various directions e

e. Pinnae with only costal areoles (or rarely 1 or 2 more produced beyond the costal one); lateral

pinnae commonly 4-6, less than 2 cm broad; rachis distinctly alate throughout

2. B. serrata

e. Pinnae amply areolate beyond the costal areole; lateral pinnae of mature leaves 8-36, 2-5 cm

broad; rachis nonalate, or scarcely alate toward the apex f

f. Areoles mostly isomorphic and slightly elongated, free included veinlets lacking or rare;

veins commonly distinct and abaxially raised; stem scales mostly 8-20 mm long and linear

(sometimes caducous) 6. B. lindigii

f. Areoles mostly heteromorphic and rather elongated, free included veinlets lacking to fre-

quent and directed distally; secondary and tertiary veins becoming indistinct between pri-

mary ones, not or scarcely raised; stem scales 2-5 mm long, ovate to broadly lanceolate

. 3. B. serratifolia

1. Bolbitis aliena (Sw.) Alston, Bull. Misc. In-

form. 1932: 310.

Acrostichum alienum Sw., J. Bot. (Schrader) 1800 (2):

13. 1802. TYPE: Collector unknown, Jamaica

(holotype, s, Herb. Swartz; isotype, s, Herb.

Swartz).

Gynnopteris aliena (Sw.) Presl, Tent, pterid. 244. 1836.

Leptochilus alienus (Sw.) C. Chr., Bot. Tidsskr. 26:

285. 1904.

Plants terrestrial or epipetric. Stem long-creep-

ing. Sterile leaves well spaced, pinnate or deeply

pinnatisect, 25-90 cm long, 14—30 cm broad.

Lamina broadest at or near the base, reduced grad-

ually to a deltate, pinnatifid apex. Rachis broadly

alate throughout, or nonalate toward base, lacking

proliferous buds. Pinnae commonly 3-6 pairs, ad-

nate, or proximal ones stalked, entire to shallowly

lobed, or basal pair with a greatly expanded basal

100

FIELDIANA: BOTANY

lobe. Veins copiously areolate, lacking included

free veinlets. Fertile leaves commonly smaller than

the sterile, with pinnae less deeply lobed.

In rain forests and wooded ravines, creeping on

earth or rocks or rocky cliffs, 100-1000 m, Ama-

zonas to Pasco and Ucayali.

Southern Mexico; Honduras; Nicaragua; West

Indies; Venezuela and Colombia, southward to

Bolivia.

This should not be confused with other species

in Peru, as it is the only one lacking a discrete

apical segment. It is sometimes confused with B.

portoricensis (Sprengel) Hennipman, a similar

species with a pinnatifid apex whose range extends

to Ecuador. However, the areoles of B. portori-

censis have included free veinlets, whereas these

are lacking in B. aliena.

Amazonas: Prov. Bagua, Rio Maranon opposite Que-

brada Mirana, Wurdack 2033 (GH, us). San Martin: Prov.

Mariscal Caceres, Dist. Tocache Nuevo, Schunke V. 3891

(F, GH, MO, us). Huanuco: Prov. Huanuco, Tingo Maria,

Tryon & Tryon 5281 (F, GH, uc, us). Pasco: Prov. Oxa-

pampa, Palcazu, Rio Alto Iscozacin, Foster & d'Achille

10129 (F). Ucayali: Km 86 on Pucallpa-Tingo Maria

road, along trail to Arboretum of Bosque von Humboldt

Experimental Station, D. Smith 1225 (MO).

2. Bolbitis serrata (Kuhn) C. Chr., Index fil. suppl.

3: 50. 1934. Figure 22a-b.

Chrysodium serratum Kuhn, Linnaea 36: 63. 1869.

TYPE: Peru, San Martin, near Tarapoto, Spruce

4123 (holotype, B; isotypes, BM!, GH!, K!, NY, p,

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

Acrostichwn serratum (Kuhn) Baker, Syn. fil., ed. 2:

524. 1874, nom. illeg. (not Poiret, 1810).

Leptochilus serratus (Kuhn) C. Chr., Bot. Tidsskr. 26:

289. 1904.

Plants terrestrial or epipetric. Stem long-creep-

ing, slender, to 5 mm thick. Sterile leaves lacking

a proliferous bud, subdistant, pinnate, to 40 cm

long and 1 0 cm broad. Lamina with 4-6(-8) pin-

nae and a conform apical segment, this narrow-

lanceolate or narrow-elliptic and with a narrowly

cuneate base. Rachis distinctly alate throughout,

the wing with a cartilaginous margin. Pinnae 0.8-

2(-2.3) cm broad, short-stalked, the margin entire

to crenate-serrate, the base narrow-cuneate. Veins

areolate only along the costa, or rarely with 1 or

2 more areoles beyond the costal one, the areoles

lacking included free veinlets. Fertile leaves com-

monly longer than the sterile, the lamina smaller,

but the petiole much longer.

In rain forests, creeping among wet rocks or on

soil, 100-550 m, San Martin and Loreto.

Colombia; Ecuador; Peru.

The smaller leaves of B. serrata, with fewer pin-

nae and more slender stems, usually distinguish it

from B. lindigii and B. serratifolia, but the vena-

tion pattern is a more diagnostic character. In most

pinnae there is merely a costal areole, but some-

times one or two areoles may be produced beyond

this. None of the areoles bear included free vein-

lets. The veins of B. lindigii regularly anastomose,

are commonly distinct and raised abaxially, and

the areoles beyond the costal one are nearly iso-

morphic (pentagonal or hexagonal, not greatly

elongated), they lack included free veinlets, or the

rare ones are minute and often directed toward

the costa. Veins of B. serratifolia rather freely anas-

tomose, but are not or scarcely raised, and sec-

ondary and tertiary veins become indistinct be-

tween the primary ones. The areoles are quite

irregular in shape and many are greatly elongated.

Free included veinlets may be frequent to sparse

or lacking, but when present they are directed to-

ward the pinna margin.

San Martin: Tarapoto, near road to Yurimaguas, Mar-

tin & Plowman 1815 (GH, us). Loreto: Balsapuerto, lower

Rio Huallaga basin, Killip & Smith 28495 (F, GH, NY,

us). Prov. Alto Amazonas, above Pongo de Manseriche,

Wurdack 2110 (F, GH, NY, s, u, uc, us).

3. Bolbitis serratifolia (Kaulf.) Schott, Gen. fil. t.

13. 1835. Figure 22c.

Acrostichum serratifolia Kaulf., Enum. fil. 66. 1824.

TYPE: Brazil, Rio de Janeiro (B?, LE?).

Poecilopteris crenata Presl, Epim. bot. 174. 1851.

TYPE: Brazil, near Rio de Janeiro, Mikan (ho-

lotype, PR or PRC).

Leptochilus serratifolius (Kaulf.) C. Chr., Bot. Tidsskr.

26: 289. 1904.

Leptochilus crenatus (Presl) C. Chr., Index fil., suppl.

1: 48. 1913.

Bolbitis crenata (Presl) C. Chr., Index fil., suppl. 3: 47.

1934.

Plants terrestrial, epipetric, or hemiepiphytic.

Stem short- to long-creeing, sometimes low-scan-

dent, to 1 .5 cm thick, sparsely provided with ovate

or lanceolate scales 2-5 mm long. Sterile leaves

occasionally with a proliferous bud (or primor-

dium) subapically on the apical segment (or pin-

nae), approximate on the stem, pinnate, to 1.2 m

long and 30 cm broad, the rachis, costae and veins

sparsely provided abaxially with castaneous, fila-

mentous scales. Lamina with 8-36 pinnae and a

TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV.

101

conform or subconform apical segment, this

broadly lanceolate and with a broadly cuneate or

rounded base. Rachis not alate, or only scarcely

so near the apex. Pinnae 2—4 cm broad, short-

stalked, the margin entire to crenate-serrate. Pri-

mary veins distinct, but secondary and tertiary ones

usually indistinct between the primary ones, not

or scarcely raised, amply anastomosing, with a

broad costal areole and many heteromorphic and

often elongated areoles beyond, free included vein-

lets lacking, or few to many and then directed

distally (toward pinna margin). Fertile leaves sub-

equal to the sterile but the petiole proportionately

longer and the pinnae much shorter and narrower.

In dense forests or wooded ravines, on moist

ground, often low-scandent on bases of tree trunks,

occasionally on rocks, 150-1600 m, San Martin

and Loreto south to Ayacucho and Cuzco.

Southern Mexico (Chiapas); Costa Rica; Pana-

ma; Surinam to Colombia, southward to Brazil

and Argentina.

This species bears a superficial resemblance to

B. lindigii. See treatments of the latter and B. ser-

rata for detailed comparison.

San Martin: Near Tarapoto, Spruce 4736 (BR, G, GH,

K., P). Loreto: Puerto Arturo, below Yurimaguas, Killip

& Smith 27797 (F, us). Huanuco: Prov. Huanuco, Tingo

Maria, Tryon & Tryon 5234 (F, GH, us). Junin: Schunke

Hacienda above San Ramon, C. Schunke A-215 (GH,

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

rimac, Killip & Smith 22626 (F, us). Cuzco: Prov. La

Convention, Cordillera Vilcabamba, Rio Klause, Dud-

ley 10187 (GH).

4. Bolbitis nicotianifolia (Sw.) Alston, Bull. Misc.

Inform. 1932: 310.

Acrostichum nicotianifolium Sw., Syn. fil. 13. 199.

1806. TYPE: Virgin Islands, St. Thomas, Ven-

tentat (holotype, s).

Gymnopteris nicotianifolia (Sw.) Presl, Tent, pterid.

244. 1836.

Leptochilus nicotianifolius (Sw.) C. Chr., Bot. Tidsskr.

26: 285. 1904.

Leptochilus killipii Maxon, Amer. Fern J. 21: 138.

1931. TYPE: Panama, above Juan Diaz, Killip

2778 (holotype, us!; isotypes, B, P, s).

Bolbitis killipii '(Maxon) Lell., Fern Gaz. 11: 107. 1975.

Plants terrestrial or hemiepiphytic. Stem short-

to long-creeping or scandent, moderately provided

with lanceolate, brown to blackish scales 3-5 mm

long. Sterile leaves lacking a proliferous bud, ap-

proximate to subdistant on the stem, to 1.2m long

and 40 cm broad. Lamina pinnate (rarely simple

or subternate), with 3-8 pinnae and a conform or

somewhat enlarged apical segment, this elliptic and

with a cuneate or shortly decurrent base. Rachis

nonalate. Pinnae commonly to 28 cm long and 8

cm broad, subsessile or short-stalked, the margin

essentially entire, the base cuneate or often obtuse

basiscopically. Veins copiously areolate, with many

free included veinlets spreading in various direc-

tions. Fertile leaves pinnate (rarely simple), sim-

ilar to sterile ones, but with fewer and smaller

pinnae.

In rain forests or in wooded ravines, on wet

ground or on bases of tree trunks, 100-540 m, San

Martin, Loreto, Huanuco, Cuzco, Madre de Dios,

and Puno.

Guatemala to Panama; West Indies; the Guia-

nas to Colombia and south to Peru.

The rare specimens with simple leaves or only

with two pinnae might be confused with B. oli-

garchica or B. pandurifolia. See treatments of these

two species for further discussion.

Bolbitis nicotianifolia and B. oligarchica are very

closely related and may hybridize, as suggested by

Hennipman (1977). There is a specimen from Ta-

rapoto, San Martin, LI. Williams 6064 (us), con-

taining a portion of a sterile leaf which seems to

be intermediate between the two species. Although

there are nine pinnae (typical of B. nicotianifolia),

these are more strongly produced basiscopically

and only 1 1/2 times as long as broad (characteristic

of B. oligarchica). Furthermore, there is a prolif-

erous bud in the axil of a subapical pinna, a con-

dition common in the latter species, but not in the

former. Both species have been found in San Mar-

tin.

San Martin: "In monte Guayrapurima," Spruce 4636

(K). Loreto: Salinas, Rio Mazan, Schunke 372 (F, GH, uc,

us). Huanuco: Prov. Pachitea, Comunidad Nativa Santa

Marta, D. Smith 1244 (MO). Cuzco: Prov. Paucartambo,

Hacienda Villa Carmen, Vargas 14680 (GH). Madre de