Illinois Natural History Survey bulletin


\

Monograph of the

Neotropical Fern Genus
Polybotrya (Dryopteridaceae)

RobbinC. Moran

Volume 34, Article 1

November 1987





Monograph of the Neotropical Fern Genus

Polybotrya (Dryopteridaceae)

RobbinC. Moran

Illinois Natural History Survey

Illinois Department of Energy and Natural Resources

Department of Plant Biology

University of Illinois at Urbana-Champaign

Illinois Natural History Survey Bulletin

Volume 34. Article 1

November 1987



This work is dedicated to my parents, Livia Ann Moran and John Howard Moran.

Dr. Robbin C. Moran currently works at the Missouri Botanical Garden in St. Louis.

Illinois Natural History Survey, Lorin I. Nevling, Chief

A Division of the Illinois Department of Energy and Natural Resources

Printed by Authority of the State of Illinois

(62997—1,200—11-87)

No charge is made for most publications of the Illinois Natural History Survey, and a list of those

publications is available upon request. Single copies of most publications are available to anyone request-

ing them. Requests for multiple copies should be made in writing and should explain the use to be

made of the publications. Address correspondence to the Office of the Chief at the address below.

Illinois Natural History Survey

Natural Resources Building

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Champaign, Illinois 61820

Citation:

Moran, R.C. 1987. Monograph ofthe Neotropical Fern Genus /'o/)*o/r>'<'(Dryopteridaceae). Illinois

Natural History Survey Bulletin 34 (I): U1.18.

US ISSN 0073-4918



Contents

Acknowledgments v

Part One: Introduction and Discussion

Materials and Methods 1

Taxonomic History of the Genus 3

Ecology 6

Geography 7

Morphology and Anatomy II

Chromosome Numbers 27

Cladistic Analysis of the Species 27

Subdivision of the Genus 31

Relationships with Other Groups 34

Part Two: Taxonomic Treatment

Notes on the Presentation of Data 37

Description of the Genus 38

Key to the Species of Polybotrya 39

Species Descriptions 43

Polybotrya subgenus Soromanes

1

.

Polybotrya serratifolia (Fee) Klotzsch 43

2. Polybotrya polybotryoides (Baker) Christ 46

3. Polybotrya suberecta (Baker) C. Chr 50

4. Polybotrya andina C. Chr 50

Polybotrya subgenus Sorbifolia

5. Polybotrya sorbifolia Kuhn 53

6. Polybotrya fractiserialis (Baker) J. Smith 54

7. Polybotrya crassirhizoma Lellinger 58

8. Polybotrya espiritosantensis Brade 60

Polybotrya subgenus Polybotrya

9. Polybotrya caudata Kunze 60

10. Polybotrya goyazensis Brade 66

1 1

.

Polybotrya pubens Martius 68

12. Polybotrya glandulosa Kuhn 71

13. Polybotrya lechleriana Mettenius 71

14. Polybotrya attenuata Moran 74

15. Polybotrya stolzei Moran 76

16. Polybotrya alfredii Brade 78

17. Polybotrya botryoides (Baker) C. Chr 80
18. Polybotrya lourteigiana Lellinger 82

19. Polybotrya pittieri Lellinger 84

20. Polybotrya cylindrica Kaulfiiss 86

2 1

.

Polybotrya hickeyi Moran 88

22. Polybotrya puberulenta Moran 90
23. Polybotrya alata Moran 92



24. Polybotrya aequatoriana Moran 94

25. Polybotrya appressa Moran 94

26. Polybotrya altescandens C. Chr 97

27. Polybotrya gomezii Moran 99

28. Polybotrya osmundacea Willd 101

29. Polybotrya cyathifolia Fee 106

30. Polybotrya latisquamosa Moran 108

31. Polybotrya sessilisora Moran 108

32. Polybotrya canaliculata Klotzsch Ill

33

.

Polybotrya semipinnata Fee 113

34. Polybotrya speciosa Schott 115

35. Polybotrya pilosa Brade 117

Names of Uncertain Application 119

Excluded Taxa 120

Literature Cited 122

Taxa and Distribution of Polybotrya 1 24

Distribution Maps 124

Index to Collectors' Numbers 132

Index to Taxonomic Names 137



Acknowledgments

No one helped me more than the five pteri-

dologists who carefully reviewed every page

of this monograph: Dr. David B. Lellinger,

Smithsonian Institution, Washington, D.C.;

Dr. Alan R. Smith, University of California,

Berkeley; Mr. Robert G. Stolze, Field Mu-

seum of Natural History, Chicago, Illinois;

Dr. Rolla M. Tryon, Harvard University,

Cambridge, Massachusetts; and Dr. Warren

H. Wagner, Jr., University of Michigan, Ann

Arbor. Not only did they spend long hours

reading my rough manuscripts, but they also

patiently answered my numerous queries

about the intricacies of fern taxonomy.

My first field trip—to Costa Rica—was

made on funds awarded by the Tinker Foun-

dation. Doctoral Dissertation Improvement

Grant # 83-06990 from the National Science

Foundation provided most of my financial

support, primarily for three additional col-

lecting trips in Latin America and for a semes-

ter of study at Harvard University.

Two people were particularly helpful dur-

ing my fieldwork in Latin America. First,

Mr. Luis D. Gomez P. of the Museo Nacional

de Costa Rica, San Jose, gave me lodging in

his house and extended numerous other cour-

tesies while I worked in Costa Rica. Second,

Professor Francisco Ortega, UNELLEZ,
Guanare, Venezuela, gave me the use of his

herbarium and plant-drying facilities and took

me on collecting trips through the Andes of

western Venezuela. I thank them both for

their hospitality and for sharing their knowl-

edge of ferns.

During my semester of study at Harvard

University, Dr. Alice F. Tryon was extremely

helpful in showing me how to obtain high

quality scanning electron microscope photo-

graphs of spores. Dr. Rolla M. Tryon helped

with nomenclatural, systematic, and other

matters.

I learned much about pteridophyte biol-

ogy from two summer field courses with Drs.

Warren H. Wagner and Rorence S. Wagner,

the first at Flathead Lake Biological Station,

Montana, and the second at Mountain Lake

Biological Field Station, Virginia. I thank

them for these memorable summers.

Dr. Kenneth R. Robertson of the Illinois

Natural History Survey, Champaign, was my
major advisor and provided much help and

encouragement. Others who served on my
doctoral committee were Dr. J. Leland

Crane, Illinois Natural History Survey,

Champaign; Drs. Johannes M. J. de Wet,

Zane B. Carothers, and Malcolm L. Sargent,

University of Illinois at Urbana-Champaign;

Dr. Rolla M. Tryon, Harvard University,

Cambridge; and Dr. Warren H. Wagner, Uni-

versity of Michigan, Ann Arbor. Dr. Tom
L. Phillips, University of Illinois at Urbana-

Champaign, provided numerous helpful com-

ments, esp)ecially on the Ecology and Geog-

raphy sections.

1 completed most of the work for this

monograph in the herbarium at the Illinois

Natural History Survey, and I am greatly in-

debted to that institution for its support and

for the use of its facilities. Several biologists

at the Survey helped me in various ways. Mr.

John Taft and Ms. Mary Kay Solecki provid-

ed useful discussion and patiently tested my
keys. Dr. David Swofford was helpful with

the cladistic analysis. Mr. Bill N. McKnight

was a constant friend and critic throughout

the study and assisted with the reproduction

of the illustrations. 1 am also indebted to the

editorial staff at the Survey, especially Patty

Duzan and Eva Steger who typeset the man-

uscript, Audrey Hodgins who edited it. and

Molly Scott who helped with the production.

I am grateful to the directors and curators

of the following herbaria for the large quan-

tities of valuable material they made available

to me, often for a considerable period of time.

The abbreviations used in the text follow the

names of the herbaria; Arnold Ariwretum,

Harvard University (A), Cambridge, Mas-

sachusetts; Herbarium Jutlandicum, Univer-



sity of Aarhus (AAU), Risskov, Denmark;

Herbarium Amazonense, Universidad Na-

cional de la Amazonia Peruana (AMAZ),
Iquitos, Peru; Botanischer Garten und

Botanisches Museum (B), Berlin, Federal

Republic of Germany; British Museum of

Natural History (BM), London, England;

Herbier (CAY), Cayenne Cedex, French

Guiana; Herbario Nacional Colombiano,

Museo de Historia Natural, Universidad Na-

cional (COL), Bogota, Colombia; Herbario

Nacional de Costa Rica, Museo Nacional

(CR), San Jose, Costa Rica; John G. Searle

Herbarium, Field Museum of Natural History

(F), Chicago, Illinois; Conservatoire etJardin

botaniques de la Ville de Geneve (G), Swit-

zerland; Gray Herbarium, Harvard Univer-

sity (GH), Cambridge, Massachusetts;

Rijksherbarium (L), Leiden, Netherlands; In-

stituto Miguel Lillo de la Fundacion Miguel

Lillo (LIL), Tucuman, Argentina; Facultad

de Ciencias Naturales y Museo, Division

Plantas Vasculares, Universidad Nacional de

La Plata (LP), La Plata, Argentina; Herbario

Nacional de Bolivia, Universidad Mayor de

San Andres (LPB), La Paz, Bolivia; Her-

barium of the University of Michigan

(MICH), Ann Arbor, Michigan; Missouri

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

New York Botanical Garden (NY), Bronx,

New York; Museum National d'Histoire

Naturelle (P), Paris, France; Academy of

Natural Sciences of Philadelphia (PH),

Philadelphia, Pennsylvania; Herbario Uni-

versitario, Universidad Nacional Experimen-

tal de los Llanos Occidentals "Ezequiel

Zamora" (PORT), Portuguesa, Venezuela;

Herbario del Instituto de Ciencias Naturales,

Universidad Central (Q), (Juito. Ecuador; In-

stituto de Ciencias, Pontificia Universidad

Catolica del licuador (C?CA), Quito, Ecua-

dor; Jardim Botanico do Rio de Janeiro (RB),

Rio de Janeiro, Brazil; Department of Bot-

any, University of California (UC), Berke-

ley, California; United States National Her-

barium, Smithsonian Institution (US), Wash-

ington, D.C.; Herbario San Marcos, Museo

de Historia Natural, Universidad Nacional

Mayor de San Marcos de Lima (USM). Lima,

Peru; Instituto Botanico (VEN), Caracas,

Venezuela; Pringle Herbarium, University of

Vermont (VT), Burlington, Vermont; Institut

fiir systematische Botanik der Universitat

Zurich (Z), Zurich, Switzerland.

This work was originally part of a disser-

tation submitted to the Graduate College of

the University of Illinois at Urbana-Cham-

paign in partial fulfillment of the require-

ments for the degree of Doctor of Philosophy

in plant biology.



Monograph of the Neotropical Fern Genus Polybotrya

(Dryopteridaceae)

Robbin C. Moran

Part One:

Introduction and Discussion

The need still is for more monographic work

on tropical species, which represent the great

majority of all ferns.

R.E. Holttum(1982)

The genus Polybotrya (Dryopteridaceae) in-

cludes 35 species of neotropical ferns. It is

distinguished by 1 ) strongly dimorphic leaves

with fertile leaves that resemble skeletons of

the sterile, photosynthetic ones; 2) usually

high-climbing stems that are covered with

scales; and 3) a unique stem anatomy with 5

to 12 circularly arranged meristeles, each sur-

rounded by a black sclerenchymatous sheath,

with numerous tiny leaf traces arching be-

tween adjacent meristeles. The center of di-

versity of the genus is the Andes, where 23

sjjecies occur, 12 of which are endemic. Tlie

coastal mountains of southeastern Brazil are

notable because they contain 5 species, all

endemic. The range of Polybotrya is from

Chiapas, Mexico, southward through Central

America; the West Indies; northern South

America southward along the Andes to

Bolivia and Paraguay and eastward to the

Guiana Highlands; the Amazon River basin

and the Matto Grosso; and southeastern

Brazil. Species of the genus typically inhabit

wet, shaded, primary tropical forests from

sea level to 2500 m, most often occurring at

middle altitudes between 500 and 2000 m.

Polybotrya is divided into three subgen-

era: 1 ) Soromanes , leaves simply pinnate and

veins anastomosing; 2) Sorhifolia, leaves

simply or twice pinnate and veins free, close,

and parallel; and 3) Polybotrya, leaves de-

compound and veins free. Polybotrya cer-

vina, a species usually included in Polybot-

rya, is removed to the monotypic genus Ol-

fersia (Moran 1986). Polybotrya is related to

dryopteroid genera such as Arachniodes,

Cyclodium, Maxonia. Olfersia, and Poly-

stichopsis. Carl Christensen, the father of

modem fern taxonomy, observed (1916) that

Polybotrya may have arisen from Maxonia

because both have high-climbing stems and

strongly dimorphic leaves. The morphologi-

cal and anatomical evidence presented here

suggests that Polybotrya may have evolved

instead from a Cyclodium-Wkt ancestor.

I chose Polybotrya for study because two

aspects of the genus immediately intrigued

me: its strongly differentiated sterile and fer-

tile leaves and its long, creeping hemiepiphy-

tic stem (Fig. 1 ). Since these features evolved

separately in unrelated fern genera, studying

Polybotrya might well provide insight into

broader questions of fern evolution. Polybot-

rya was suited to monographic study because

the number of species (35), all of which are

neotropical, was manageable. Finally, no

previous monographic work had been done

on Polybotrya and many problems of

nomenclature and identification remained to

be solved.

Materials and Methods

This monograph is based on the study of

about 25(X) herbarium sheets, which repre-

sent approximately 8(X) individual collections

from 30 herbaria (see acknowledgments).

Unfortunately, Polybotrya at Kew could not

be examined because their pyolicy precludes

loaning specimens for use by graduate stu-

dents (G.L. Lucas, in litt. 1983; pers.

comm.). Kew, however, did send color slides

of several critical type specimens.



Illinois Na iural History Survky Vol.34. Art. 1

FiGliRE 1 . Habit sketches for two species of Polyhoina. Above. P. cr(issirhi:om(2. a climbing species;

below. P. sorbijolia. a teirestnal species. Note that the fcnile leaves an; skeleionlikc and that iheir

orientation is more erect than that of the spreading sterile leaves.



November 1987 Monograph of Polybotra

I spent seven months observing and col-

lecting 1 8 of the 35 species of Polyhotrya in

Costa Rica, Ecuador, Peru, and Venezuela

and was able to make such important obser-

vations about the biology of the species as

presence of aerophores and mucilage on

stems, duration and habit of sterile versus

fertile leaves, and variation within and be-

tween populations. During field work, her-

barium, cytological, and anatomical mater-

ials were also collected.

Anatomical cross sections were prepared

by freehand sectioning, and staining was

rarely needed to determine cell types or tissue

layers. Leaf segments were cleared by soak-

ing them in 10 percent NaOH in a warm oven

for 5 to 6 days. Several changes of clearing

solution were usually needed to replace so-

lution that had become darkly stained by

phenolic substances. Since this procedure did

not remove all the dark color from the leaf,

the segments were placed in full-strength

Clorox bleach from 10 to 60 minutes. This

treatment usually rendered even the most

stubborn leaves translucent for microscopic

study.

Taxonomic History of the Genus

Polyhotrya was first described by Willdenow

in his fourth edition of Linnaeus' s Species

Plantarum (1810), as distinguished by "Cap-

sules sessile, globose, aggregated in naked

paniculate spikes. Nonindusiate" [my transla-

tion]. Polyhotrya osmundacea, the sole

species described in the new genus, received

its specific epithet because of the fancied re-

semblance of its fertile leaf with the fertile

leaf apex oi Osmunda regalis. Since strongly

contracted, nonindusiate fertile leaves were

considered the principal feature of the new

genus, other species with similar leaves were

eventually placed in Polyhotrya. Since fully

differentiated fertile leaves have arisen sepa-

rately along many phyletic lines, this proce-

dure resulted in a highly unnatural, poly-

phyletic genus. Species placed in Polyhotrya

at one time and then excluded are represented

today in as many as 12 genera (see Excluded

Taxa).

Past Generic Concepts

Each of the nineteenth-century pteridologists

who wrote about Polyhotrya had a slightly

different concept of the genus (Table I).

Biume (1S28) placed many unrelated acros-

tichoid ferns, including species of Loma-

gramma. Stenosemia, and Bolhilis, in Poly-

hotrya. SchotI (1834-1836) later restricted

Pohhotrxa to the American species allied to

P. osmunducea. The subsequent recognition

of the Asian acrostichoids as distinct from

the American genus Polyhotrya testifies to

Schott's keen perception. Although Schott's

classification was not immediately adopted

by his colleagues, it was eventually revived

by Smith (1875) and Christensen (1934).

PresI (1836) relied heavily on venation

patterns and anatomical characters in defining

his genera and did not accept Schott's circum-

scription of Polyhotrya. He merged species of

Bolhitis sect. Egenolfianae with Polyhotrya

because both had free veins, but he removed

the anastomosing-veined species placed in

Polyhotrya by Blume (1828). Smith (1841)

largely followed this classification in his ar-

rangement of fern genra.

Fee (1845) was the first pteridologist to

try to make sense out of the increasing

number and diversity of acrostichoid ferns.

He included the species related to P. osmim-

dacea in his subgenus Eupolybotrya and

created two subgenera of Polyhotrya (see

Excluded Taxa) in which he put many species

now placed in Bolhitis, Blechnum, Lomariop-

.v/.v, and Teraiophyllum. Fee also erected the

segregate genus Soromanes for species of

Polyhotrya with anastomosing veins and 1-

pinnate leaves. In my treatment. Soromanes

is a subgenus of Polyhotrya.

The German pteridologist Mettenius had

a wide circumscription of Polyhotrya. He de-

scribed several new species in the genus,

some of which are here retained. The major-

ity, however, have been placed in Bolhilis,

Eluphoglossiiin , and Leptochilus.

Hooker (1864; 194-195) and Baker in

Hooker and Baker ( 1 874: 399ff . ) subsumed

both Polyhotrya and Soromanes as subgenera

in their large, eclectic genus Acrostichum.



Illinois Natural History Survey Vol.34. An. 1

which consisted of any fern with acrostichoid

sori. This decision was essentially a reversion

to Swartz's (1806) concept of Acrostichum,

and although both genera were recognized as

subgenera, Polybotrya continued to include

unrelated species that have since been placed

in Atalopleris, Elaphoglossum, Psomio-

carpa, and Teratophyllum. Hooker and

Baker's classification was accepted by sev-

eral pteridologists, especially by flora writ-

ers, until the hegemony of Hooker's concepts

was supplanted by the more natural views of

Smith (1875). Smith accorded generic status

to Polybotrya, restricting it to P. osmundacea

and its immediate allies; however, he kept

Soromanes as a distinct genus.

Unfortunately, Smith's views were not

immediately accepted by pteridologists.

Christ (1897) and Diels (1899) classified

many of the acrostichoids with free veins in

Polybotrya, a decision that again resulted in

an amalgamation of unrelated sfjecies. Chris-

tensen ( 1905) in Index Filicum used Polybot-

rya at the rank of genus, but it had essentially

the same circumscription as it had had under

Hooker. Unlike the writers cited above, he

placed Olfersia cervina in Polybotrya. His

classification was used by Schumann (1915)

in her important work on the vascular supply

in fertile leaves of acrostichoid ferns. Chris-

tensen (1934) later removed the discordant

species from Polybotrya and placed most of

them in the genera where they are found today

(Atalopteris , Egenolfia. Lomagramma. and

Psomiocarpa)

.

Copeland's Genera Filicum (1947)

adopted Christensen's earlier ( 1905) concept.

Pichi-Sermolli (1977) treated Polybotrya.

Table 1. Comparative treatment oi Polybotrya.

Reference

Polybotrya

subgenus

Soromanes*

Polybotrya

subgenus

Sorbifolia*

Polybotrya

subgenus

Pol\holr\a*

Genera formerh

included in

Pol\botr\a

WilidenowdSlO)

Kault'uss{1824)

Blume(1828)

Schott( 1834-1836)

Presl(l836)

Smith (1841)

Fee (1845)

Hooker (1864)

Smith (1875)

Christcnscn(1905)

Christcnscn(l934)

Copeland(1947)

Pichi-Scniiolli(1977)

Tryon& rryon(1982)

Soromanes

Acro.^tichutn

subgenus

Soromanes

Soromanes

Pohholrxa

Polybotrya

Polybotrya

Soromanes

Polxbotr\a

Acrostichum

subgenus

Polybotrya

Polybotrya

Pohbotrxa

Polybotrya

Polybotrya

Polybotrya

Pol\botr\a

Polybotrya

Polybotrya

Polybotrya

Polybotrya

Polybotrya

Polybotrya

Polybotrya

subgenus

Eupolybotrya

Acrostichum

subgenus

Polybotrya

Polybotrya

Pohbotrxa

Polybotrya

Polybotrya

Polybotrya

Pohbotrxa

Olfersia

Bolbitis. Lomma-

gramma. Stenosemia

Bolbitis

Bolbitis

Blcchnum. Bolbitis.

Lomariopsis.

Teratophyllum

.Atalopleris.

Psomiocarpa.

Teratophyllum

.Atalopteris. Bolbitis.

Olfersia. Psomio-

carpa. Teratophyllum

Olfersia

Olfersia

* refers to the treatment of the cenus in this work.



November 1987 Monograph of Poi.ybotra

Soromanes, and Olfersia as distinct genera

but placed Soromanes and Polyhotrya on

separate branches of his phylogenetic dia-

gram. Tryon and Tryon (1982) are the most

recent authors to discuss Polybotrya and re-

lated genera; their concept is like that used

in the present work, except that they include

Olfersia cervina in Polybotrya. In the present

monograph, 1 restrict Polybotrya to the

species allied closely to P. osmundacea,

make Soromanes a subgenus of Polybotrya.

and keep Olfersia cervina. which has been

placed in Polybotrya by many recent

pteridologists, in its own monotypic genus.

1 recognize 35 species of Polybotrya. all of

which are neotropical and 10 of which are

new.

Work at the Species Level

The species of Polybotrya have received less

study over the years than those of most other

fern genera. Fee (1845), Hooker (1864), and

Hooker and Baker (1874) have been the only

monographers of the genus. Although they

attempted to identify all of the then-known

species of Polybotrya. they did not always

see types, overlooked several published

names, put some species in synonymy with-

out adequate study, and did not make detailed

observations on the genus because they were

studying hundreds of other ferns at the same

time. Despite these shortcomings, the works

of these three men have been the best source

for identifying specimens of Polyhotrya in

many regions of tropical America.

The difficulty of identifying specimens

of Polybotyra has been partially alleviated

by local or regional floras, but these cover

only a limited portion of the geographic range

of the genus and are often incomplete. In

many of these floras, types were rarely

examined and names, therefore, were often

applied incorrectly. Only Sodiro's (1897)

treatment of the ferns of Ecuador is available

for the Andean region; yet he was aware of

only 6 of the 23 species that occur in that

region. Vareschi's (1969) treatment for Ven-

ezuela and Brade's (1971) for Brazil have

helpful illustrations, but the names are often

misapplied. The best treatment of Polybotrya

for Mesoamerica is Stolze's ( 1981 ) excellent

work for the Flora of Guatemala, but

Guatemala lacks several species found in

Mesoamerica. In brief, adequate keys and

descriptions are lacking for identifying

Polybotrya throughout most of its range and,

as a result, many specimens have been mis-

identified.

In this century, most of the research on

Polybotrya has focused on the specific level.

Brade (1935, 1948) described two new

species of Polybotrya in papers that included

other ferns as well. Later, in a series of three

papers published in the Brazilian journal

Bradea ( 1969a, b,c), he described ten addi-

tional new species from Brazil, Costa Rica,

and Venezuela. These papers were based on

field studies in the American tropics, and

most of the types were plants he had collected

by himself or with his brother Alfred. Brade

(1971) published a synopsis providing keys

and illustrations of the Brazilian species of

Polybotrya. Unfortunately, he died before

completing the descriptions, habitats, and

distributional information. Only five of

Brade's names are accepted in this mono-

graph as representing valid species; the re-

mainder are relegated to synonymy.

More recently, Lellinger (1972, 1977)

published two important papers on Polybot-

rya. In the first, he described five new species

from South America and offered interesting

ecological notes on the genus. In the second,

which deals with other ferns as well, he de-

scribed two new species that are endemic to

Colombia. Only three of these seven species

are accepted here. About half of the species

described by Brade and by Lellinger have,

therefore, been relegated to synonymy in the

present work, largely because I found older,

obscure names during my research. In some

cases, these names had not been u.sed since

their original publication. The proliferation

of names is one of the pitfalls faced by

taxonomists when synthetic or revisionary

works are unavailable for a group and in this

case indicates how much Polybotrya has

needed a monograph.



Illinois Natural History Survey Vol.34. Art. 1

Ecology

Polyholrya inhabits wet, shaded, tropical

forests. These may be hot, humid, lowland

rain forests or cool, cloud forests at high ele-

vations. The genus rarely occurs in disturbed

forests, and when it does, it is represented

by only a few isolated plants. However, small

scale disturbances within mature forest, such

as light gaps created when a huge tree dies,

appear to be important in establishing young

plants (Lellinger 1972). During fieldwork in

tropical America, I never observed young

sporophytes or their associated gametophytes

establishing themselves on roadbanks or on

shaded, disturbed slopes adjacent to roads.

Furthermore, all herbarium specimens that

contained habitat information listed only pri-

mary forest as the habitat. The restriction of

Polyhotrya to wet, shaded, primary, tropical

forest is reflected by its geographic distribu-

tion (Fig. 2a). The genus is absent from such

arid regions as central Mexico and the west-

em coast of Peru and from such nonforested

regions as the Llanos of Venezuela and Co-

lombia (Figs. 2a & 3). The altitudinal range

of Polyholrya is from sea level to 25(X) m.

with most plants collected from 500-2000 m
(Fig. 4). This range certainly corresponds to

my field observations that Polyhotrya is most

diverse, frequent, and abundant at middle

elevations.

Figure 2. Distribution of Polyholrya. a. composite of Maps 1-21; b. number of species (left bar),

number of endemics (right bar). Countries or areas are outlined; those with only one bar have no

endemics. See Table 3 for numbers of taxa in each countn,



November 1987 Monograph of Polybotra

Geography

In the following discussion, the distribution

of Pohhotrya is given according to the re-

gional centers defined by Tryon (1972).

These regional centers, all mountainous,

were determined by their high endemism and

species richness.

The most important regional center in the

diversification of Pohhotrya is the Andean,

from Venezuela and Colombia south to

Bolivia (Fig. 2b, Table 2). The Andean Re-

gion contains 23 species, 12 of which are

endemic, and I suspect that new species of

Pohhotrya will be found there. Within this

region. Colombia contains the richest

Pohhotrya flora: 17 species, 6 of which are

endemic (Table 3). The coastal mountains of

northern Venezuela contain P. canaliculata

and P. serratifolia, both occurring in cloud

forests from 1200-2400 m. Pohhotrya

canaliculata has a very limited distribution

and is a true endemic to the coastal mountains

(Map 19); it occurs outside of the Andean

Region as defined by Tryon (1972). Polyhot-

r\a serratifolia is considered a near-

endemic, extending southwest into the Andes

around Merida and eastward to Trinidad

(Map 1).

The Serra do Mar Mountains of the South-

eastern Brazilian Region contain five species

Figure 3. Distribution of Polybotrya species within regional centers of diversity and endemism for

tropical American ferns defined by Tryon (1972) Primary centers arc indicated with dark shading:

secondary centers, with light shading. The number on the left is the total number of species; the number

in parentheses indicates how many of those species are endemic. Sec Table 4 for comparisons.



Illinois Natural History Survey Vol. 34, Art. 1

of Polyhotrya, all of which are endemic (Fig.

3, Tables 2 & 3). I find it remarkable that

this region has no species of Polybotrya in

common with those of the other regional cen-

ters (Table 4) or with intervening areas. This

distinctiveness of the Polybotrya flora is ac-

centuated by its lack of the simply pinnate

Sf)ecies, that is, subgenera Soromanes and

Sorhifolia. The biogeography of Polybotrya

strongly supports data from many other

sources that suggest that the isolated Serra

do Mar Mountains are a distinctive biogeo-

graphic center (Tryon 1972).

Only one species, P. goyazensis, occurs

in the Matto Grosso and intervening area be-

tween the Andean and Brazilian regional cen-

ters (Map 6). This species is closely related

to the widespread P. caudata and is probably

an example of the peripheral "budding" of a

new species from the southern part of the

range of P. caudata.

The Amazon basin contains one endemic,

P. glandulosa, which has been collected only

three times (Map 7). Most species of Polybot-

rya in the Amazon basin also grow in the

Andean Region. In contrast, the coastal

mountains of .southeastern Brazil have

contributed no species to the Amazonian low-

lands. My field experience in the Amazon

basin of Ecuador and Peru showed that in

forests on wet, sandy soils P. caudata was

common; forests on lateritic soils, however,

had P. crassirhizoma. P. caudata. P. osmun-

dacea. and P. puhens. All five species were

abundant in the western Amazon basin where

I collected, and I suspect that they extend

farther eastward into the Amazon basin than

is shown on their range maps.

The Guiana Highlands Region does not

contain a distinctive Polybotrya flora. Only

six species occur there and most of these are

widespread elsewhere. The region, therefore,

has a high fioristic affinity with the Andean

and Central American regional centers (Table

4). Polybotrya sessilisora, which occurs in

the Guiana Highlands and in the adjacent

Amazon basin (Map 20), is probably more

common in the Highlands and its range may

be centered there. Polybotrya has two notable

examples of Andean-Guianan range disjunc-

2500 -

2000 -

2 1500
Ul

I-
UJ

2 1000

500 -

2 3

31 33

I

Figure 4. Altitudinal ranges of Polyholrya species. The numbers ab*ive the bars correspond to the

species numbers assigned in the taxonomic treatment.



November 1987 Monograph of Polybotra

tions. TTie first example, P. lechleriana, is

known in the Guianas from a single collection

at Mount Roraima in Guyana. 1900 km from

its nearest locality in Colombia (Map 7). This

disjunction is probably best explained by

long-distance spore dispersal from Andean

populations. The second example, P. frac-

tiserialis. occurs primarily in the foothills of

the Andes and is disjunct in the southern

Pakaraima Mountains of British Guiana,

2000 km from the nearest Andean population

in Ecuador. Unlike P. lechleriana, P. frac-

tiserialis is common and widespread in the

Guianas (Map 3).

The Central American Region contains

seven species, including one endemic (Fig.

3, Table 2). All seven species occur in Costa

Rica (Table 3). My fieldwork there showed

that P. polyhotryoides, P. osmundacea. P.

alfredii and P. gomezii (endemic) were par-

ticularly common in the mountains and that

P. caudata was frequent in the costal low-

lands. Polyhotrya alfredii and P. gomezii

were particularly abundant in cloud forests,

often growing together. In the mountains of

central Panama, just outside the Central

American Region as defined by Tryon

(1972), one endemic occurs, P. alata.

The Greater Antilles Region, otherwise

quite rich in fern species, has only one

Polxhoirya species, the widespread P. os-

mundacea (Figs. 2 & 3, Table 2). Puerto

Rico lacks Polyhotrya but certainly has suit-

able forest habitat. In the Lesser Antilles, P.

osmundacea is found on Grenada. Martinique

and Guadeloupe have the closely related en-

demic P. cyuthifolia. The Caribbean islands

have played a minor role in the diversification

of Polyhotrya.

Polyhotrya caudata and P. osmundacea

clearly have the largest geographical and al-

titudinal ranges of any species in the genus

(Maps 5 & 18). With the exception of south-

eastern Brazil, these two species extend

nearly throughout the entire area covered by

all of the other species in the genus.

Polyhotrya occurs on only one oceanic

island—Cocos Island, about 500 km (310

mi) southwest of Costa Rica in the Pacific

Ocean. Three species occur there: P.

Table 2. The geographic regions shown in Figure 3 and the species oi Polyhotrya that occur in them.



10 Imjnois Na riKAi History Survry Vol. 34, Art. 1

Table 3. Distribution of Polyhoina by country. Endemics are in boldface. Numbers in parentheses

refer to the species numbers assigned in the taxonomic treatment.

Mexico: 2 species, no endemics, polyholryoides (2), caudata (9).

Belize: 2 species, no endemics, polybotryoides (2), caudata (9).

GuATEMAt,A: 3 species, no endemics, polybotryoides (2), caudata (9), osmundacea (28).

Honduras: 3 species, no endemics, polybotryoides (2), caudata (9), osmundacea (28).

Nicaragua: 3 species, no endemics, caudata (9), alfredii (16), osmundacea (28).

Co.STA Rica: 6 species, 1 endemic, polybotryoides (2), sorbifolia (5), caudata (9), alfredii (16),

gomezii (27), osmundacea (28).

Cocos Island: 3 species, no endemics, polybotryoides (2). caudata (9), osmundacea (28).

Panama: 5 species, 1 endemic, polybotryoides (2), caudata (9), alfredii (16). alata (23),

osmundacea (28).

Cliba: 1 species, not endemic, osmundacea (28).

Jamaica: 1 species, not endemic, osmundacea (28).

Haiti: 1 species, not endemic, osmundacea (28).

Guadeloupe & Martinique: 1 endemic, cyathifolia (29).

Grenada: 1 species, not endemic, osmundacea (28).

Trinidad: 3 species, no endemics, serratifolia (1), caudata (9), osmundacea (28).

French Guiana: 3 species, no tr\Atm\cs. fractiserialis (6), caudata (9), osmundacea (28).

Surinam: 2 species, no endemics. /r«(r/.?er/a//.v (6), caudata (9).

Guyana: 4 species, no cnAetmcsfractiserialis (6), caudata (9), lechleriana (13). osmundacea (28).

Venezuela: 6 species, 2 endemics, serratifolia (I), sorbifolia (5), caudata (9), glandulosa (12),

osmundacea (28), canaliculata (32).

Colombia: 17 species, 6 endemics. /7()/v/wrno((yf.?( 2 ).ii(/)<'rf(7a( 3 ),iorfe//o/ia (5), crojiir/i/roma (7),

caudata (9), pubens (11), lechleriana (13), attenuata (14). stolzei (15). botryoides (17),

lourteigiana (18), pittieri (19). hickcyi (21), altescandcns (26), osmundacea (28),

latisquamosa (30), sessilisora (31).

Ecuador: 14 species, 3 endemics, polybotryoides (2), suberecia (3), andina {4). fraciiserialis (6).

crassirhizoma (7), caudata (9), pubens (11), lechleriana (13), alfredii (16), puberulenta (22),

aequatoriana (24), appressa (25), altescandcns (26), osmundacea (28).

Peru: 1 1 species, no endemics, polybotryoides (2). suherecta (i).fractiserialis (6).

crassirhizoma (7), caudata (9), pubens (11), glandulosa (12). lechleriana (13), alfredii (16).

altescandcns (26). osmundacea (28).

Bolivia: 10 species, no endemics. /rac/i.vfn'u//.? (6), crassirhizoma (7), caudata (9),

pubens (11), lechleriana (13), alfredii (16). hirkcyi (21). puberulenta (22). aequatoriana (24).

osmundacea (28).

PARACiUA"!': 1 species, not endemic, goyazensis (10).

Brazil: 1 3 species, 5 endemics, sorhifrlia (5), crassirhizoma (7), cspiritosantensis (8). caudata (9),

goyazcnsis (10), pubens (11), glandulosa (12). cylindrica (20), osmundacea (28), sessilisora (31).

seniipinnata (33), speciosa (34), pilosa (35).



November 1987 Monograph of Poi.ybotra

caudata. P. osmundacea. and P. polyhot-

ryoides. As noted in the previous paragraph.

P . caudala and P . osmundacea have the most

extensive range of any species in the genus.

Although not as wide ranging, P. polyhot-

ryoides has a long north-south distribution

from Chiapas, Mexico, to Pasco, Peru, and

is the most far-ranging species of the sub-

genus Soromanes. This distribution supports

Tryon's (1970) findings that the ferns of

oceanic islands tend to be those that are wide-

spread beyond the source area for the island

(the source area for Cocos Island is defined

by Tryon as Central America and Colombia).

According to Tryon (1970), widely distrib-

uted species have a broader ecological

amplitude and therefore establish themselves

more often on oceanic islands because of their

ability to grow undera wide range of environ-

ments.

Morphology and Anatomy

Information on the morphology and anatomy

of Polyhotrya has been extremely useful for

assessing affinities of related genera and for

understanding functional aspects of the biol-

ogy of the genus. Every phylogenetic argu-

ment made here relies heavily on evidence

from stem anatomy (along with other fea-

tures, of course) in determining how Polyhot-

rya relates to other fern genera and families.

Anatomical studies have also answered other

biological questions about the genus. For

example, studies of the diplodesmic venation

system showed how the sori are efficiently

supplied with water and nutrients. Studies of

sterile-fertile transitional leaves established

that the diplodesmic veins are homologous

with the veinlets of the sterile leaves. In short,

anatomical studies of root, stem, and leaf

have been valuable in understanding diverse

questions about the biology of Polyhotrya.

Roots

A dense tangle of dark, tough, fibrous,

adventitious roots ari.ses from the ventral side

of climbing stems and from all sides of ter-

restrial stems. No positional relationship

exists between the leaf bases and the roots;

roots are apparently borne at random. Roots

arising directly from the stem measure about

I mm in diameter, but their terminal branches

may be only 0.3 mm wide. Roots branch at

irregular intervals, increasing the number of

tiny supportive rootlets. On climbing plants,

these tenacious rootlets penetrate the outer

layers of tree bark and firmly anchor the stem.

This firm attachment hinders pulling the stem

away from the tree. When a stem is forcibly

pulled away from a trunk, the large roots of

Polyhotrya remain attached to the stem,

clinging to shredded pieces of bark, lichens,

and mosses.

I studied the root anatomy of six spe-

cies— P. alfredii, P. caudata. P. gomezii,

P. osmundacea, P. polybotryoides , and P.

sorhifolia. All were similar. Figure 5a shows

Table 4. Affinities offo/yiofrya floras between regional centers (.see Fig. 3). Floristic affinity is the

percent species in common (C) of the total species in the two regions (A + B), i.e., 1(X)C/A + B.

Regional Center

(Tryon 1972)

Total Species Species in Common Floristic Affinity

(A + B) (C) (%)

Andean &Guianan

Central American & Andean

Andean & Southeast Brazilian

Central American & Guianan

Guianan & Southeast Bra/.ilian

Central American & Southeast Brazilian

29

30

28

13

11

5



12 Illinois Natural Hlstory Survey Vol.34. Art. I

a root in cross section taken about 0.5 cm
from the ventral surface of a scandent stem

of P. osmundacea. The stele is weakly ellip-

tic, protostelic, and comprises about one-fifth

of the root's diameter. Two protoxylem poles

occur; each is exarch and consists of four to

seven protoxylem elements. The center of

the stele contains three to five large

metaxylem cells. Surrounding the xylem are

phloem elements— minute, angular, thin-

wailed cells about the same diameter as the

protoxylem elements. They are not shown in

Figure 5a because they were poorly pre-

served.

Surrounding the stele is the innermost

layer of the cortex— the endodermis. It con-

sists of a single, continuous, light-colored

layer composed of thin-walled, rectangular

Jg$k^

QA^

0.1 mm

0.1 mm
Figure 5. Root anatomy and various epidermal features of Polyhoirya. a.b. and c are the same size.

a. root cross section of P. osmundacea: b. close-up of a stem meristele in cross section, showing root

trace diverging about midway between the laterally diverging leaf traces: c. phloroglueinol-containing

gland from the stem epidermis of P. cauduta; d. stomata on stem aerophore of P. caudata: e. fertile

segment of P. stolzei, adaxial surface. Note different shapes of epidemial cells and the two kinds of

hairs; the flaccid hair represents the type that intergrades with scales. EN endodermis; EP epidermis;

LT leaf trace; PH phloem; RT root trace; SC sclerenchyma: SH sclerenchyma sheath surrounding a

meristele; T tracheid; X xylem.



November 1987 Monograph of Poi.ybotra 13

cells. Although the endodermis is narrow, it

contrasts sharply with the dark, thicker-

walled cells on either side. The anticlinal

walls have casparian strips that are easily vis-

ible even in unstained material.

Outside the endodermis is a ring of dark,

thick-walled fibers (Fig. 5a). This ring is the

most conspicuous feature of Polyhotrya root

anatomy. Unstained, the walls of the fibers

appear orangy or reddish and may become

so thick that they nearly occlude the entire

lumen of the cell. The walls thicken evenly

on all sides, unlike many angiosperms and

some ferns where the deposition of cell walls

occurs centripetaily (Bierhorst 1971; Wagner

1952),

Outside the ring of dark fibers, the corti-

cal cells take on a different appearance. In

contrast to the cells of the inner ring of fibers,

the outer cortical cells are parenchymatous

with thin, irregular, dark brown cell walls.

Cells of the epidermis are compact and

quadrangular, in contrast to the shape of the

cortical cells immediately below. The most

conspicuously differentiated cells of the

epidermis are the root hairs, generally 0.2-

1 .0 mm long and straight, narrow, and unicel-

lular. Their color normally varies from brown

to amber, but brilliant yellow hairs occur in

several species. Unfortunately, the root

anatomy of Polyhotrya cannot be compared

with that of other genera of dryopteroid ferns

because of inadequate information for those

genera.

Stem

Habit. The stem is climbing in all species

of Polyhotrya except for P . fractiserialis and

P. sorbifolia, where it is terrestrial (Fig. I).

Climbing stems grow horizontally about 0.5-

2.0 cm beneath the forest floor until they

contact a tree. The stem then turns upward,

anchoring itself to the tree by adventitious

roots produced on the ventral surface. Stems

usually climb to a height of 1 to 5 m, but the

highest scandent stem I saw was just under

15 m. The longest underground stem that I

saw was traced back 8 m from where it en-

tered the soil at the base of its support tree.

As a stem begins to climb, it grows wider.

The diameter of a climbing stem may reach

3.5 cm; terrestrial stems seldom grow wider

than I cm. The intemode length on climbing

stems is generally 10-15 cm.

Stems of the two terrestrial species of

Polyhotrya differ in overall appearance from

stems of the climbing species. Stems of ter-

restrial species are about 1
5 -20 cm long with

intemodes 1-3 cm long. These short inter-

nodes impart a basketlike aspect to the ar-

rangement of leaves instead of the linearly

spaced aspect typical of the leaves of climb-

ing species (Fig. I). The amount of scale

cover is another difference. Stems of the ter-

restrial species are less scaly than stems of

climbing species due to the sloughing off or

rotting away of scales, presumably because

the scales are buried in moist, tropical soils

where decay is rapid. Furthermore, the

petiole bases hide the few scales that remain

in terrestrial species; the best place to find

scales is the stem apex, where there is less

decay and fewer petiole bases. Despite these

differences, both types of stems show affinity

by sharing the distinctive anatomy described

below.

Branching is rare in stems oi Polyhotrya.

I never saw dichotomy of the stem apex, and

lateral branching was exceedingly rare in

scandent stems, although occasional in ter-

restrial portions. I did not investigate the re-

lationship, if any, of branching to leaf posi-

tion, but that examination might provide

further evidence for comparisons with other

genera of dryopteroid ferns.

Anatomy. The stems of all dryopteroid

fern genera have radially symmetric dictyo-

steles; however, the stems of Polyhotrya are

distinctive (Fig. 6). The most noticeable fea-

ture in cross section is the vascular bundles

—

terete, circularly arranged, each bundle en-

sheathed by a jet-black ringof sclerenchyma.

The 5 to 12 vascular bundles are positioned

around the central portion of the stem, their

dark outlines contrasting vividly with the

matrix of bright white ground parenchyma.

Numerous tiny leaf traces emanate from the



14 III iNois Natural Hisiory Survey Vol.34. An. 1

PH -r^c,-^^

1

Figure 6. Stem anatomy of Polyhoirya osmumlucca. typical of the genus. Lower left is a cross section

of the stem with a mcristclc enlarged above. Note "xylary arm" in 2. AP aerophore: EN end^xlemiis;

GP ground parenchyma: l.T leaf trace: MS meristele: P parenchyma: F^ pericycle: PH phloem:

RT root trace: 1" trachcid: SH sclcrenchyma sheath consisting of fibers: X xylem: XP protoxylem.



November 1987 Monograph of Polybotra 15

sides of the bundles, collectively forming a

C-shaped line that connects adjacent bundles.

This pattern is unmistakable (Fig. 6), and no

other fern genus yet studied has this kind of

stem anatomy. (For comparison of stem

anatomy with closely related groups, see Re-

lationships with Other Groups.)

The epidermis of the stem is darkly

sclerotized and one cell layer thick; it pro-

duces numerous scales (described separately

below). The cell walls of the epidermis are

relatively straight compared to the sinuous

ones of the leaf epidermis (cf. Figs. 5c & 9).

The stems of Polyhotrya have minute

glandular hairs on the epidermis. These short-

stalked hairs have, in relation to their length,

exceptionally large, round heads, 54-56

microns in diameter (Fig. 5c). Because these

glands are tiny and few, they had been over-

looked until found by Widen et al. (1983),

who were also the first to report phloro-

glucinols in Polyhotrya. The phloroglucinols

are apparently produced and stored in these

glands. In other dryopteroid ferns, such

glands are known to secrete an oleoresin,

along with various lipids, between the cuticle

and the outer layer of the cell wall. The

phloroglucinols are, presumably, located in

this secretion (Widen et al. 1983). Using ex-

tracts from the stem and petiole bases of P.

caudata. Widen et al. found that aspidin BB
was the principal compound, along with

smaller amounts of albaspidin BB,desaspidin

BB, and flavaspidic acid BB. Internal glandu-

lar hairs, which probably also contain

phloroglucinols, occur in the stems and

petioles of other dryopteroid ferns (Mehra &
Mittal 1961 ). I did not, however, find internal

glands in Polyhotrya.

Aerophores occur on the stems of most

species of Polyhotrya but are apparently ab-

sent in Sf)ecies having a mat of thick, woollike

scales, such as in P. aequatoriana, P. alte-

scandens, and P. crassirhizoma. On living

stems, aerophores appear as yellowish white

lines 1-3 mm wide that connect with

aerophores on the petiole (for description of

petiolar aerophores, see section below on

petioles). Aerophores usually extend 2-6 cm
down the stem from the base of the petiole.

In cross section, they project about 0.5-1

mm above the surrounding surface and con-

sist of thin-walled parenchyma cells that in-

terrupt the otherwise continuous, thickened

epidermis (Figs. 6 & I6g). Stomata abound

and can be seen with a binocular microscope

(Fig. 5d). Presumably, aerophores allow oxy-

gen into the stem that is otherwise surrounded

by compact, oxygen-impermeable, scleren-

chyma fibers. Because aerophores shrink and

darken upon drying, thereby matching the

contour and color of the stem's scleren-

chyma, they cannot be seen on herbarium

material.

Below the epidermis is a well-developed

hypodermis, usually three to eight cells thick

and darker than the epidermis that forms a

conspicuous ring around the white ground

parenchyma. The cells of the hypodermis are

thick-walled sclerenchyma fibers that thwart

cutting the stem, .sometimes even with a hefty

machete, and I had problems cutting thin,

even, cross sections for microscopic study.

A ring of hypodermal sclerenchyma also en-

circles the stem in closely related genera such

as Arachniodes, Cyclodium, Maxonia, Olfer-

sia, and Stigmatopteris (pers. obs.).

Below the hypodermis lies the cortex,

which consists of scattered clusters of

brachysclereids (stone cells) in a matrix of

whitish parenchyma. The parenchyma cells

are generally rounded and contain abundant

amyloplasts. In a freshly cut stem, the white

color of these cells contrasts vividly with the

darkly sclerotized hypodermal and meristelar

sheaths. Sclereids never abound, as in Olfer-

sia (Fig. 16i), but are scattered in clusters

throughout the ground parenchyma. The dep-

osition of secondary walls is extremely un-

even; most of the wall is laid down on the

side of the cells facing the inside of the

sclereid cluster. A small, cuplike lumen can

usually be seen on the side of the cell facing

away from the center of the sclereid cluster.

The thick, dark cell walls contain numerous

simple pits and a lesser number of ramiform



16 M.LiNois Natural HrsxoRY Survey Vol. 34. An. I

pits. Many Dryopteridaceae and Lomariop-

sidaceae also have sclereid clusters in the

ground parenchyma of their stems (pers.

obs.).

Inside the scierenchyma sheath that sur-

rounds each meristele lies a band of paren-

chyma two to four cells wide (Fig. 6). The

cells of this layer are more compact and

smaller than the cortical parenchyma, but like

the cortical parenchyma, they contain amylo-

plasts. This inner parenchymatous layer stops

abruptly at the endodermis, which stands out

as a single row of clear, narrow, rectangular

cells interrupting the darker parenchyma on

either side of it ( Fig . 6) . The next layer toward

the center is the pericycle, which consists of

compact, orangy cells that form a continuous

band around the xylem and phloem.

The phloem consists mostly of clear, an-

gular, thin-walled sieve cells (Fig. 6.2 &
6.4). Phloem is broadest on the distal side

of the stele but forms a thin band elsewhere.

This band is broken in two locations by arms

of the xylem that extend to the pericycle (Fig.

6.2). Scattered among the clear cells of the

phloem are parenchymatous cells.

Inside the ring of phloem is the xylem,

which has large, conspicuous metaxylary

tracheids (Fig. 6.3). These tracheids occur

singly and in clusters within a matrix of dark

brown (when unstained) parenchyma. The

tracheids are generally rounded or somewhat

angular; when adjacent to another tracheid,

the walls are straight. Numerous pits occur

between all the cells of the xylem. Pro-

toxylem elements occur in the xylary "arms"

(Fig. 6.2). Each meristele is, therefore,

diarch and exarch.

Root traces arise between the xylary arms

of the meristele (Fig. 5b). The roots diverge

through the cortex to the ventral surface of

the stem. In contrast to the root traces, the

leaf traces originate from the xylary arms.

Four to six leaf traces are usually produced

at each leaf gap from adjacent meristeles.

During their passage from the cortex to the

petiole, the leaf traces bifurcate and anas-

tomose, forming a reticulum like that in

Maxonia (Chandra 1975). Fach leaf trace is

surrounded by a single layer of sclerotized

cells (Fig. 5b). TTie walls of these cells are

brownish and slightly thickened on the inner

tangential surface. The distance from the in-

ception of a leaf gap to the point where the

leaf diverges from the stem is about 10-15

cm.

Stem scales. Many spec\es of Polybotrya

have distinctive stem scales. Some species,

in fact, can be identified solely on the basis

of their stem scales (thus the importance of

always collecting part of the stem). Color is

the easiest feature of the scales to use in iden-

tifying the species of Polybotrya. Botanists

should have no difficulty identifying P. alte-

scandens, with its bright golden or yellow

scales; P . crassirhizoma, with its dull orange

or red scales; and P. latisquamosa. with its

shiny, castaneous scales. Depending on the

species, scales range from concolorous to

slightly darker in the center to bicolorous with

a dark central stripe.

Scale habit is another helpful identifica-

tion feature. Some species, such as P. alte-

scandens and P . crassirhizoma. have narrow,

densely tangled scales that impart a woolly

appearance to the stem. Other species, such

as P. appressa. P. caudata. and P. cylin-

drica, have thick, dark, massive scales that

are conspicuously appressed to the stem. Be-

cause of their great width and spreading habit,

the scales of P. serratifolia are distinct from

those of other species of the subgenus

Soromanes . Most species oi Polyhatrya have

ascending and slightly spreading scales—

a

characteristic that is not particularly distinc-

tive because of its prevalence in the genus.

The shape of stem scales is usually not

helpful in identification because most species

have lanceolate to linear-lanceolate stem

scales. Two species, however, P. alata and

P. altcscandcns (Figs. 42 & 45). have ex-

tremely long, attenuate scales that differ strik-

ingly from those of the other species. Petiole

scales are generally similar to those of the

stem but tend to be shorter and wider. The

petiolar scales of P. latisquamosa have be-

come, as the specific epithet implies, greatly

widened .so that this characteristic distin-



November 1987 Monograph of Polybotra 17

guishes the sfiecies from all others in the

genus (Fig. 50d).

Despite variation in color, habit, and

shape, stein scales are of two general types,

here termed simply Type 1 and Type 2. Scales

of Type 1 are thin and translucent and have

easily visible cell walls (Fig. 7). Most of the

scale is one cell layer thick, although the

center may be thicker and darker. The mar-

gins vary from denticulate to ero.se, with the

teeth formed from the projecting ends of two

adjacent cells (Fig. 7). Marginal teeth in

0.1 mm

Figure 7. Stem scales of various Polybotrya species, a, b.

i. P. crassirhizoma.

P. altescandens; c. P. boiryoides;



18 Ii I iNois Natlrai. History Survey Vol.34. An. I

Polyhotrya are never glandular, as in Bolhitis

(Hennipman 1977) and Stigmatopteris (pers.

obs.). Scales of Type I are attached basal I y

at a single point or. more rarely, across the

full length of the base. This point or line of

attachment is always darkly sclerotized (Fig.

7a). Representative species with scales of

Type 1 include P . crassirhizoma. P. goyazen-

sis, P. puherulenta, and P. serratifoUa (Figs.

26, 29, 41, & 19).

Scales classified as Type 2 are thick,

opaque, and composed of many cell layers.

The margins are entire or subentire. The base

is greatly thickened, sometimes almost

stipelike, and is curved and ascending. When
the scale is removed, a circular or oval scar

remains on the stem. Scales of this type are

usually appressed, but they are squarrose in

P. osmundacea. Representative species with

scales of Type 2 include P. appressa, P.

caudata, and P. cylindrica (Figs. 44, 28, &
39). In contrast to scales of Type I, which

show tremendous variety in habit, color, and

shape, those of Type 2 are relatively

homogeneous.

Evidence suggests that morphological

transition occurs easily between the two types

of scales. Species such as P. lechlehana. P.

lourteigiana. P. osmundacea. and P. puhcns

have both types of scales but on different

individuals. Even on a single plant, scales

belonging to Type 1 can occur on the climb-

ing portion, with scales of Type 2 and/or

intermediates on the terrestrial portion. If

morphological transition is easily ac-

complished, then evolutionary change in

scale type could be expected to occur in

closely related species pairs. This hypothesis

is supported by two closely related species

P. caudata and P. goyazensis (Figs. 28 &
29); the former has scales of Type 2 whereas

the latter has scales of Type 1 . Clearly, the

stem scales of Polyhotrya have been mor-

phologically and evolutionarily plastic.

The evolutionary advantage of scales in

Polyhotrya is unknown. Their contribution

to protecting the stem from mechanical dam-

age is probably minimal since the stem is

surrounded by a tough, resistant, scleren-

chymalous sheath. Scales seem unnecessary

on the older, hardened parts of the stem. Dur-

ing fieldwork in Latin America. I found sev-

eral plants of Polyhotrya that had lost all of

their scales on the lower, older part of the

climbing stem, yet the plants appeared vigor-

ous. Perhaps the scales afford protection to

the stem apex , where young tissues have not

yet become sclerified.

Another hypothesis is that the scales re-

tain water by capillary action for later use by

the roots. Retention would be advantageous

to a climbing plant that has the leaves on the

scandent portion of the stem separated by

several meters from the roots in the soil.

Stems collected in the field, however, never

seemed to hold water between the scales and

plants never appeared water stressed. The

bark of the support tree, which is usually

covered with water-retaining mosses,

lichens, and organic debris, probably could

provide most of the water needed by the plant.

Further, the retention of capillar)' water be-

tween stem scales might be harmful in a hot,

humid, tropical environment that promotes

decay. No satisfacton,' hypothesis, therefore,

has been offered to explain the adaptive sig-

nificance of scales in Polyhotrya.

Even more difficult to explain is the

adaptive advantages of the numerous minor

modifications of scales. What could be the

function of a dark, central stripe, of reddish

color, of an appressed habit, or of a linear

shape? How these and other minor modifica-

tions of scales are significant in evolution, if

indeed they are. is baffling.

Leaves

Petiole. The petiole base in some ferns

is specialized to accumulate food and to per-

sist as a storage organ long after the rest of

the leaf to which it was attached has withered

and decayed. These speciali7ed petiole bases,

called "trophopods." are a new source of sys-

tematic data in ferns (Wagner & Johnson

1983). Polyhotrya. however, lacks troph-

opods. and the food-storing function of the

trophopod is perfomied b)' the stem in the



November 1987 Monograph of Polybotra 19

amyloplast-containing cells of its ground

parenchyma.

A cross section of the petiole of Polyhot-

r\a reveals leaf traces that are arranged in a

mushroomlike outline, with the base of the

mushroom oriented adaxially (Fig. 8a). This

pattern also occurs in Cyclodium trianae, a

species closely related to Polyhotrya. and in

some species of the similar family Lomariop-

sidaceae (Hennipman 1977; Holttum 1978).

Other closely related dryopteroid genera,

however, such as Cyrtomium, Dryopteris,

and Polystichum, display a C-shaped arrange-

ment of leaf traces in the petiole (pers. obs.).

These genera also have fewer leaf traces than

Polyhotrya, probably as a consequence of

their narrower petioles. Further study of

petiole anatomy in dryopteroid genera may

provide helpful information for assessing

their relationships.

In Polyhotrya. the two vascular bundles

closest to the adaxial surface of the petiole

differ from the lower vascular bundles in sev-

eral respects. The most obvious difference is

their shape: elongate and slightly curved in-

stead of round (cf. Fig. 8b, d). Inside each

of the two adaxial bundles, the xylem is hook-

shaped in outline, in contrast to the lenticular

shape seen in the lower traces. Only these

uppermost vascular bundles, not the lower,

yield the lateral traces that supply the pinnae.

As they traverse the rhachis and petiole, all

traces remain separate; they do not anas-

tomose to form a reticulum within the petiole

.

Aerophores run down the sides of the

petiole and join below with those on the stem.

On living petioles, they appear narrow,

linear, and light green. The surface of the

aerophores bears stomata, and beneath them

is parenchyma that contains intercellular

spaces. This parenchyma, when seen in cross

section (Fig. 8c), interrupts the otherwise

continuous band of collenchyma that encir-

cles the petiole. Upon drying, the aerophores

darken and collapse, leaving a sulcus on

either side of the central, adaxial sulcus of

the petiole. Drying, therefore, makes the

petiole trisulcate; in living petioles only a

central sulcus is present.

Petiole and stem mucilage. Mucilage

has apparently never been reported in

Polyhotrya or any other genus of Dryop-

teridaceae. During fieldwork in Costa Rica

and Ecuador, however, I found that stems

and petiole bases of P. alfredii usually had

a thick coat of translucent mucilage similar

to that on the stems and/or petioles of certain

Blcchnum and Thelypteris species. Upon dry-

ing, the mucilage disappeared completely,

although sometimes the scales exhibited a

matted or flattened appearance, as if stuck

together. Attempts to rehydrate the mucilage

failed.

Other species of Polyhotrya probably

have mucilage. Several dried specimens of

P. lechleriana and P. lourteigiana exhibited

matted, flattened scales that suggested the

earlier presence of mucilage. These two

species are closely related to P. alfredii, an

observation that suggests that mucilage is

probably confined to the P. alfredii group

(species nos. 13-20). I did not see matted,

flattened scales in any other species of

Polyhotrya. The mechanism of mucilage se-

cretion and its adaptive significance, if any,

is unknown.

Nectaries. Koptur et al. (1982) were the

first to report the existence of nectaries in

Polyhotrya. They found them in P. osmun-

dacea on the rhachis of unfurling leaves, near

the yellowish, lateral, linear aerophores. Sev-

eral times during fieldwork in Costa Rica, I

noticed ants on young leaves, but I never saw

them feeding on nectar. The nectar itself

would certainly be nutritious because it con-

tains sucrose, glucose, and fructose in con-

centrations up to 35-75 percent by weight

(Koptur et al. 1982). Amino acids also occur

in the nectar, and Koptur et al. list those

present. Although ants may benefit from the

nectaries, additional fieldwork is needed to

establish whether or not the plants benefit.

Apparently, ants do not defend the plants— at

least I was never attacked by ants while col-

lecting Polyhotrya.

Costa-costule architecture Costa-cos-

tule architecture refers to the morphology of

the juncture of these axes. Several kinds of



20 Illinois Na turai- His iory Survey Vol. 34, An. 1

0.1 mm

FiouRE 8. Petiole anatomy of Potybotrya alfredii. a. cross section of petiole 2 cm above the stem

.showing the arrangement of vascular bundles in the shape of a mushRxim: b. adaxial bundle showing

hooked xylcm; c. aerophore; d. abaxial bundle. COL collenchyma; EN endodermis; EPI epidermis;

T tracheid.



November 1987 Monograph of Poi.ybotra 21

costa-costule architecture occur; the most

thorough review of them is by Holttum

(1959); more than any other pteridologist, he

has shown the importance of this feature in

the taxonomy of ferns. According to Holttum

(1984), the dryopteroid genera have "midribs

of ultimate leaflets grooved, the groove of

the rachis bearing the leaflets being open to

admit the leaflet-groove, the margin of the

lamina of the leaflet being decurrent (but not

prominent) down the side of the rachis; hairs

of ctenitoid type lacking." This type of ar-

chitecture is illustrated in Figure 47h.

Holttum (1984) describes the tectarioid gen-

era as having "midribs of ultimate leaflets

more or less prominent (in Tectaria some-

times slightly grooved) and bearing ctenitoid

hairs, usually many." In Polybotrya. the

basic type of architecture is dryopteroid, a

finding that supports its classification among

the dryopteroid ferns.

Within Polybotrya. however, modifica-

tions are found in the dryopteroid pattern of

costa-costule architecture. Typical dryop-

teroid costa-costule architecture is shown in

the most primitive group in the genus— the

group of species related to P . osmundacea.

In contrast, the most advanced species in the

genus

—

P. altemuita, P. lechleriana, and P.

stolzei— always have the costular groove

truncated by the ridges of the costa; therefore,

the groove is not decurrent (Figs. 33b, 32b,

& 34f). This truncated costa-costule architec-

ture is considered to have been derived from

the decurrent-grooved condition, as evi-

denced from outgroup comparison with other

dryopteroid ferns. Intermediates, such as P.

aequatorianu and P. alfredii. have costular

grooves that gradually fill up near the costa,

so that the ridges of the costa are only slightly

interrupted by the shallow, weakly decurrent

groove of the costule (Fig. 43d). In P. alfredii

and P. lourteigiana, these three types of

costa-costule architecture can be found on

different parts of the same leaf, although the

intermediate type predominates. In summary,

most species of Polybotrya have dryopteroid

costa-costule architecture, but distinctive

modifications of this basic type occur.

Anatomy of the lamina. I studied leaf

clearings and cross sections for 17 species of

Polybotrya to determine differences between

species or subgenera. Leaf clearings revealed

that stomata were absent from the adax-

ial epidermis but abundant on the abaxial sur-

face between the veins. The long axes of the

stomata ran almost parallel to the veins. The

mean length of the stomata (for all species)

was 56 microns, but means for the species

ranged from 48 to 74 microns. Subsidiary

cells surrounded the distal one-half to two-

thirds of the guard cells. The shape of the

subsidiary cells can be like the sinuous epi-

dermal cells or slightly more rounded and

circular (Fig. 9d,f)-

In face view, the epidermal cell walls fit

together like the pieces of a jigsaw puzzle

(Fig. 9). This interlocking is best exemplified

by the abaxial epidermal cells, which have a

more highly irregular outline than the adax-

ial ones. The cells above and below the veins,

however, have straight walls. The slightly

elongate axes of the cells are oriented parallel

to the veins and point in the direction of the

leaf margin.

In cross section, the abaxial epidermis

and adaxial epidermis form a single, compact

layer of relatively thick-walled cells (Fig.

9g,h). Palisade parenchyma is absent from

the mesophyll, which is composed mostly of

large intercellular air spaces surrounded by

long, cylindrical, parenchymatous cells. An

arm of collenchyma extends from the abaxial

epidermis to the vein, where it forms a sheath

around the vein but does not continue to the

adaxial side of the leaf. Leaf anatomy was

similar in all species of Polybotrya. probably

because the entire genus occupies a similar

habitat.

Hair types. Two fundamentally different

types of hairs occur in Polybotrya. One type

intergrades with scales, the other does not.

The nonintergrading type is erect or spread-

ing, usually cylindrical, and whitish or

tawny. The size of hairs of this type varies

tremendously, from unicellular hairs that are

less than 0. 1 mm long and not visible to the

naked eye to readily visible acicular hairs



22 Illinois Natural History Survey Vol. 34. An. 1

that have up to 15 cells and are up to 2 mm
long (Fig. 10 a-i,k). The apical cell is point-

ed, rarely rounded. All but five species of

Polyhotrya have this kind of hair.

Hair of the second type is fundamentally

different from hair of the first type because

each hair is a tiny, uniseriate scale and. there-

fore, by definition, a hair. Every intermediate

stage exists between these hairs and the

scales, as evidenced by studying their change

in form from the apex to the base of the costa

(Figs. lOj & 36a,b). At the apex of the costa.

^cpZ9^r

FiGLiRF. 9. Leaf anatomy of Polyhotrya. a-f. face views of the abaxial leaf epidermis The elongalcd

cells with straight walls lie above the veins, g, h. cross sections of the leaf. a. Polyhotrya serratifolia:

h. P. puhcns: c. P . fractiserialis: d. P. canalUulata: e, g. P. crassirhizoma: f. h. P. speciosa.



November 1987 Monograph of Polybotra 23

only the tiny uniseriate scales, the "hairs,"

occur. These are readily distinguished from

other hairs by their flattened cells, appressed

habit, and darkened transverse walls. To-

wards the base of the costa, the hairs become

longer and are composed of more cells; soon

are found "hairs" that are two cells wide near

their base, that is, scales. These small scales

grade imperceptibly into the larger scales on

the rhachis. The scales on the rhachis and

costae are, therefore, developmentally

homologous with this second kind of hair.

FiCiiiRF 10 Hair types in Polyhotrya taken from the abaxial surface of the eo.sta. a. P. puhens;

b. P. semipinnata; c. P. altescandens: d. P. pillicri. e. P. alfrejii, f. P. ^kmdulosa: g. P. speciosa;

h. P. osmundacea; i. P. caudata; j. P. lechleriana (reduced scale type); k. P. cylindrica.



24 iLi.iNors Natural History Survey Vol. 34. Art. 1

These hairs are called proscales since they

are developmentally, and probably evolu-

tionarily, precursors to scales (Moran 1986).

Proscales occur on the lamina and axes

of all species of Polybotrya and all genera

of dryopteroid ferns. Proscales are, however,

so minute that pteridologists often overlook

them or do not bother to mention them. Pro-

scales are smallest on the distal parts of the

lamina (Fig. 49h) and at least 30x is needed

to see them. Their flattened cells, often hav-

ing reddish cross walls, occasionally twist

upon drying, imparting the appearance of a

Ctenitis-\\a\v. These are not true Ctenitis-

hairs, however, because such hairs never in-

tergrade with scales (Christensen 1913). This

nonintergrading feature is rarely mentioned

in the definition of Ctenitis-\\&\rs.

Fertile Leaves

Moran (1987) has presented a detailed study

of the sterile and fertile leaves (trophophylls

and sporophylls) of Polybotrya. Below is a

synopsis of the subject.

The fertile leaves oi Polybotrya resemble

a skeleton of the sterile ones because their

green lamina has been reduced to a narrow

wing above the veins. This extreme dimor-

phy, termed "holodimorphy," is distin-

guished from other types of dimorphy by the

nearly total loss of photosynthesis in the fer-

tile leaf (Wagner & Wagner 1977). The re-

duction of green tissue is the most conspicu-

ous aspect of dimorphy, but differences in

seasonality, duration, orientation, insertion

on the stem, size, color, texture, and venation

are also involved. Thus, sterile-fertile leaf

dimorphy is an ensemble of characteristics

and not merely a single feature of the leaf.

Pteridologists previously described the sori

of Polybotrya as acrostichoid, but three types

of sori occur in the genus: botryoid,

coenosoric, and a new type here termed "am-

phiacrostichoid." Botryoid sori, which are

round and discrete, are considered primitive

on the basis of outgroup comparison with

other dryopteroid ferns (Fig. 38b). The

coenosoric type, which are oblong to linear,

probably evolved from the botryoid type by

basipetal fusion of the sori (Fig. 35h). The

amphiacrostichoid type developed from a

coenosoric ancestor by the expansion of the

margins of the receptacle beyond the original

adaxial surface; as a result, the receptacle

assumed its own adaxial surface and sporan-

gia appeared on both surfaces of the leaf (Fig.

30g). Coenosoric and amphiacrostichoid sori

contain diplodesmic veins that are developi-

mentally homologous with veins of the sterile

leaf.

Sporangia. The sporangial stalk of

Polybotrya is three-celled at its apex, im-

mediately below the capsule. Below the apex,

however, only two rows of cells occur, each

three to four (rarely 5) cells long (Fig. 1 1 ).

Where the two rows of the base join the three

cells at the apex, a paraphysis occurs; it is

always uniseriate. unbranched, and multicel-

lular. About three-fourths of the species of

Polybotrya have paraphysate sporangial

stalks, but paraphyses occur on only 40-80

percent of the stalks from a single sample.

In dried or rehydrated material, the apical

cell of the paraphysis is smaller and shriveled

compared to the ceils below. In fresh mater-

ial, the apical cell appears swollen and glan-

dular. Sporangial stalks of P. fractiserialis

are unique because they have lightbulb-

shaped, glandular cells at the base of the

paraphyses (Fig. I Ig). The functions of these

glandular cells and paraphyses are unknown.

Sporangial capsules of Polybotrya are

typical of those found in related dryopteroid

ferns. The mean number of annular cells for

sf>ecies of Polybotrya ranges from 12 to 24,

with most species having between 14 and 18.

Both the epistomium and hypostomium con-

sist of three or four thin-walled, transversely

elongated cells. All species of Polybotrya

have glabrous sporangial capsules, except P.

imbcns. which has short, subulate hairs at

the top of the capsule near the annulus (Fig.

30e,n. Usually two of these hairs occur on

either side of the annulus. creating the

impression that the capsule has "homs.""

Spores. Spore sizes, measured by the

longest axis, are given (when available) at

the end of each species description. Numbers



November 1987 Monograph of Poi.ybotra 2S

are for spwres measured immediately after

being placed in distilled water because this

medium is readily available to taxonomists.

Fifteen spores were measured per specimen.

Spores of most Polybotrya species range

from 45-65 microns in length, with extremes

of 32 and 80 microns. According to Tryon

and Tryon (1982). Polybotrya has relatively

large spores compared to those of other gen-

era in their tribe Dryopteroideae. The spores

appyear dark brown when viewed with trans-

mitted light under a compound microscope

but deep orange when viewed with reflected

light under a dissecting microscope.

Spores of Polybotrya are fairly uniform

within the genus compared to the variation

encountered in other genera of dryopteroid

ferns (e.g., Dryopieris and Polystu hum).

Polybotrya spores are monolete, with the

aperture linear and one-third to three-fourths

the length of the long axis. The aperture is

often obscured by the broad [lerispore folds

and smaller spines. The exospore appears

smooth (Fig. 12), as it does in spores of most

genera of dryopteroid ferns. The perispore

consists of two layers: the lower layer is thin

and appressed to the exospore; the upper layer

is thicker, with inflated folds and echinate to

various degrees (Fig. 12). No constant differ-

ences were found between the three subgen-

era or smaller species groups.

The principal differences between species

appear in the prominence and density of peri-

sporal folds and spines. Typically, the in-

FiciURF. 1 1 . Sporangia of various Polybotrya species, a. P. allenuata: b. P. sorhifolia. showing both

sides of the same sporangium; c. P. espirilosanlensis; d. P. speciosa: c. P. osmundacea; f,g. P.fracli-

seriatis (note in g the globose, glandular cell al the base of the paraphysis).



Illinois Natural History Survey

Figure 12. Spovcs oU\>tyhotrva. a. P. semipinnaki-.b. P. osmumtacea-.c. P. scrralifolio.d. P.pinien.

c. P. gomezii: f. P. alfredii: g. P. crassirhizoma: h. P. cylindrica: i. P. speciosa. a: Duane et al.

6W->-' (F) bC. Schiinkc .?S0 (GH). c: Femller 261 (GH). d: Lellingcr & de la Sola 251 (US), e:

Moran 3241 (CR). f: M,mw 3I6S (CR). g; Plowman et al. 4025 (GH). h: Handro 222S (GH). i: Webh

26 (GH). c and h arc x 5(XK1, all others are x 1(XX).



November 1987 Monograph of Polybotra 27

flated folds are well developed and the spines

are so numerous that they impart a "fuzzy"

appearance. However, the folds may be rela-

tively low. as in P. goyazensis and P. sor-

hifolia. or the spines may be reduced in height

and/or density, as in P. speciosa (Fig. I2i).

Species may also differ in spore size. This

range is clearly shown by the two terrestrial

species in the subgenus Sorbifoliu: P. sor-

bifolia and P. fractiserialis differ greatly in

spore size (40-47 and 52-56 microns long,

respectively), a difference that probably re-

flects different ploidy levels.

Comparison of the spores shown in Figure

12 with the spores of other dryopteroid ferns

illustrated by Tryon and Tryon (1982) shows

that the spores of Polyhotrya are most like

those of Maxonia and Stigmaiopteris (Tryon

and Tryon include Cyclodium in Stigmatnp-

leris). Cyclodium. which is most closely re-

lated to Polyhotrya, has spores similar to

those of Polyhotrya but less spiny (A.R.

Smith 1986). This observation supports evi-

dence from external morphology that these

genera form a closely related group.

Chromosome Numbers

The only previous reports of chromosome

numbers in Polyhotrya came from two sepa-

rate counts of P. osmundacea. one from

Jamaica and the other from Trinidad; both

gave a chromosome number of n = 41

(Walker 1966; Smith & Mickel 1977). To
add to the information about chromosome

numbers in the genus. I collected meiotic

material during fieldwork in Latin America.

Young fertile segments were placed for 3-5

hours in distilled water saturated with

paradichlorobenzene. The segments were

then removed, blotted gently, and placed in

a fixative of 3; 1 ethyl alcohol to glacial acetic

acid. The material was stored in a freezer

until it could be examined. Sporangia were

squashed in a drop of aceto-carmine and then

photographed

.

This work recorded new chromosome

number counts for four species

—

P. alfredii.

P. altescandens , P. polyhotryoides, and P.

serratlfolia (Fig. 13). Each of the four counts

had a chromosome number of h = 4 i . a find-

ing that argues, along with morphological

features, for classifying Polyhotrya among

the dryopteroid ferns, all of which have

n = 4\. Chromosome counts are still needed

from the other species of Polyhotrya. espe-

cially where polyploidy may be involved in

the evolution of one species from another,

as in P. fractiserialis and P. sorhifolia.

Cladistic Analysis of the Species

A cladogram of species relationships was

constructed using the PAUP program, ver-

sion 2.3 (Swofford 1985; PAUP is an ac-

ronym for Phylogenetic Analysis Using Par-

simony). Table 5 shows the input data used

in the analysis, and Table 6 gives the charac-

ters and character states used in the input data

matrix. The character state trees (hy-

pothesized evolutionary pathways) for the

characters (Table 6) are shown in Figure 14.

Further information on the characters and jus-

tification of their postulated evolutionary

pathways is given in the Morphology and

Anatomy section.

Because of homoplasy in the data set,

PAUP found a large number of equally par-

simonious cladograms. Therefore, a consen-

sus cladogram (Fig. 15) was printed for the

first 50 trees to determine the branching pat-

terns they all had in common. The 50 trees

agreed on the branching patterns for 12

species— about one-third of the genus. The

groups that had congruent branching patterns

were subgenera Soromanes and .Sorhifolia

(Fig. 15, .serratlfolia— espiritosantcnsis) and

the group of species related to P. caudata

(Fig. 15, caudata—puhens). Homoplasy in

the remaining species accounted for the

numerous, equally parsimonious clado-

grams. Nevertheless, distinct groups of

species are shown on the consensus clado-

gram, a result that supports the following

subdivision of the genus.



28 Ii.i-iNOis Naiurai. History Survey Vol. 34. An. 1

"f

Figure 13. Chromosome squashes of four sfwcies of Polybotrya. All squashes have n = 4l. My
interpretations are at the right, a. P. altescimdem. Ecuador. Pichincha. Moron .^559 [CV\): b. P.

polyhotryaides, Costa Rica, Cartago. Moran 2I7S (MO); c. P. serralijhlia, Venezuela. Trujillo. Moran

3709 (MO); d. P. alfredii, Costa Rica. Cartago. Moran 2442 (CR).



November 1987 Monograph of Polybotra 29

Table 5. Data matrix for cladistic analysis of 35

character states and polaiily. Ancestor = hypothetica!

? = unknown character state. NA = not applicable

species of Polybotrya. Sec text for discussion of

ancestor possessing all primitive character states.



30 Illinois Nai ural History Survey Vol. 34, An. 1

Tabi F. 6. Characters and character states used in the cladistic analysis of 35 species of Polyhotrya.

The numbers given to each character state correspond with those shown on one of the cladograms in

Figure 14.

1

.

Dissection of sterile leaves (Fig. 14a). = 4-pinnate; 1 = 3-pinnate-pinnatifid; 2 = 3-pinnate; 3 = 2-

pinnate-pinnatifid; 4 = 2-pinnate; 5 = 1 -pinnate.

2. Venation (Fig. 14b). = close and long-parallel; 1 = obliquely ascending: 2 = anastomosing.

3. Type of fertile leaf (Fig. 14c). 1 = botryoid, but with lamina not completely reduced; 2 = fully

botryoid; 3 = coenosoric; 4 = caudate.

4. Pinnule arrangement (Fig. 14d). 0= anadromic; 1 =catadromic.

5. Symmetry of pinnule base (Fig. 14b). = symmetrical and truncate; 1 = prolonged acroscopically

and truncate; 2 = symmetrical and cuneate.

6. Submarginal connecting strand (Fig. 14e). = absent; 1 = several connections; 2 = present.

7. Stem scale base (Fig. 14d).0= attached by a single point; 1 = attached across the width of the base.

8. Hair type (Fig. 14b). O = long, acicular; 1 = small, jointed; 2= uncinate.

9. Pubescence of laminar surface (Fig. 14b). O = both surfaces pubescent; 1 = glabrous; 2 = abaxial

surface pubescent.

10. Pubescence of laminar margin (Fig. I4d). = glabrous; 1 =ciliate.

1 1. Pubescence of costae (Fig. 14b). 0= uniform and dense pubescence; 1 = moderately pubescent

or with scattered hairs; 2 = glabrous.

12. Lamina base (Fig. I4b). = deltate; 1 = reduced; 2 = cuneate.

13. Scale color (Fig. I4f). = golden or yellow; 1 = brown; 2 = reddish or bnghl castaneous; 3 = cream

or whitish.

14. Receptacular hairs (Fig. 14d). = unbranched; 1= branched.

15. Size of lamina (Fig. 14d). 0= > 1 meter; 1 = < 1 meter.

16. Costal scale type (Fig. 140- = flaccid and ovate; 1 = linear to narrowly lanceolate; 2 = linear

and tortuous; 3 = caducous.

17. Apex of .sterile leaf (Fig. !4d). = pinnatifid; 1 = subconform

.

18. Stem habit (Fig. I4d). = hemiepiphytic; 1 = terrestrial

.

19. Shape of the tertiary pinnules (Fig. 14b). = ligulate; 1 = oblong or ovate; 2 = obovate.

20. Spore size (Fig. I4d). = 40-50 microns long; I =52-56 microns long.

\.

4

/



November 1987 Monograph of Polybotra 31

Figure 15. Consensus tree for 50 equally parsimonious cladograms of species relationships mPolybot-

rya. The number before the decimal refers to the character; the number after the decimal refers to the

character state (Table 6).

Subdivision of the Genus

I subdivide Polybotrya into three easily rec-

ognized subgenera: 1) Soromanes, sterile

leaves I -pinnate, veins anastomosing; 2) Sor-

bifolia. sterile leaves 1-2-pinnate, veins free,

close, and parallel; 3) Polybotrya, sterile

leaves decompound, veins free.

Subgenus Soromanes ranges throughout

Central America and the Andes, primarily in

mountainous areas, but it is conspicuously

ab.sent from the Amazon basin and from

southeastern Brazil (Maps 1 & 2). This sub-

genus consists of four well-defined species

(species nos. 1-4) that occur in montane

forests, primarily from 500-2000 m (Fig. 4).

The only contemporary pteridologist who

has maintained Soromanes at the generic

level is Pichi-Sermolli (1977), who also

placed Soromanes and Polybotrya on sep-

arate branches of his phylogenetic diagram,

associated with different generic groups.

Other pteridologists, such as Christensen

(1905), Copeland (1947), and Tryon and

Tryon (1982) have subsumed Soromanes in

Polybotrya. My research on both genera has

shown that Soromanes is the closest genus

to Polybotrya. Both genera share a unique

stem anatomy: a circular grouping of meri-

steles with each meristele surrounded by a

dark, sclerenchymatous sheath (Figs. 6 &
16g). No other fern genus has this unmistak-



32 Illinois Natural History Survey Vol. 34, Art. I

1 cm

Figure 16. Polybotrya and closely related genera, a. venation of sterile pinna. Cyclodium meniscioides;

b. venation of sterile pinna, Polybotrya polyhotryoides: c. pinna of Cyclodium triamie var. trianae

(compare to P. sessilisora and P. osmundacea): d. sterile pinna of Olfersia cenina. showing venation

entirely unlike any Polybotrya; e-i. stem cross sections (dorsal surface is up) of e. Maxonia apiifolia,

f. Lomuriopsis fendleri . g. Polybotrya caudata. h. Bolbilis lindigii. i. Olfersia cervina. AP aerophone;

LT leaf trace; RT root trace; SH sclerenchyma sheath; SC sclereid or stone cell.



November 1987 Monograph of Poi.ybotra 33

able stem anatomy, one of the hallmarks of

Poh'hotryu. Other compelling similarities,

such as holodimorphic leaves, long-creeping

and densely scaly stems, petiole anatomy,

and spore morphology, further demonstrate

that Soromanes and Polybotrya are closer

genealogically to each other than to any other

genus; that is, they are sister groups. In my
opinion, these similarities justify including

Soromanes in Polybotrya.

Subgenus Sorhifolia is primarily South

American (Maps 3 & 4) and consists of four

species (species nos. 5-8) that occur in low-

elevation forests primarily from 0-I2(X)

(1500) m (Fig. 4). Two species, P. sorhifolia

and P . fractiserialis , are very closely related

and contrast sharply with the rest of the genus

because of their terrestrial stems. These two

species also have distinctive venation: the

veins are long, parallel, and sharply ascend-

ing (Figs. 24 & 25). The third species, P.

crassirhizoma, is dissimilar from the others

and has dull orange stem scales and slightly

more spreading veins (Fig. 26). It is an abun-

dant fern in the western Amazon basin, from

Colombia south to Bolivia. Polybotrya es-

piritosanlensis is included in this subgenus,

even though its leaves are 2-pinnate because

its venation (Fig. 27b) is exactly like that of

P. fractiserialis (Fig. 25e); this similarity

suggests a close relationship.

The species having decompound leaves,

ail free veined (species nos. 9-35), make up

the subgenus Polybotrya, which is further

subdivided into four species groups. Since I

see no purpose in giving these groups formal

taxonomic names, I informally refer to them

with the name of a typical, widespread

species for each group, e.g., the "P. caudata

group." The first three groups described

below have catadromic pinnules; the fourth

has anadromic pinnules. Pinnule arrange-

ment, however, cannot be used to classify

the species groups at a higher level because

no other characters correlate with it.

TheP. c«(«/«/« group (species nos. 9-12)

is the most distinctive because it has peculiar,

caudate fertile pinnules that are soriferous on

both surfaces, that is, amphiacrostichoid. Ac-

tually, what looks like the adaxial soriferous

surface is the expanded, thickened margin of

the receptacle; the true, phylogenetic, adaxial

surface is reduced to a thin green line (see

Morphology and Anatomy section). In addi-

tion to this diagnostic sorus, the group is

further united by the presence of whitish,

septate, acicular hairs (Fig. IOa,f,i). Poly-

botrya glandulosa is tentatively assigned to

this group because of similarities in leaf cut-

ting and pubescence; however, it has botryoid

fertile leaves. This character is constant in

P. pubens. but both P. caudata and P. goya-

zensis can be glabrous. The P. caudata group

is most frequent and abundant at low eleva-

tions from 0-l(X)0 m, although P. caudata

itself occasionally occurs up to 1900 m (Fig.

4).

The second species assemblage, the P.

alfredii group, consists of eight primarily An-

dean species (species nos. 13-20). This

group has a characteristic appearance, al-

though it is difficult to describe because few

features are constant. In general, the pinnae

are short-stalked and crowded near the costa

and rhachis. The pinnae bases are more or

less equilateral and not acroscopically pro-

longed as in the rest of the genus. The tertiary

.segments are often oval or rounded, never

narrow or strap-shaped. The grooves of the

costules are either truncated by the ridges of

the costa or very weakly admitted to the

groove of the costa (see Morphology and

Anatomy section). Most species in the P.

alfredii group have botryoid fertile leaves.

The third species group, the P. altescan-

dens group, consists of seven species (species

nos. 21-27). These species, with the excep-

tion of one endemic to the mountains of Costa

Rica, grow in the Andes at slightly higher

altitudes than the rest of the genus (Fig. 4).

This group has pinnule ba.ses slightly to

strongly asymmetric, a characteristic that

helps to distinguish it from the P. alfredii

group.

The fourth species group centers on the

type of the genus, P. osmundacea. and differs

from the rest of the genus by having anadrom-

ically arranged pinnules. As shown in the

illustrations, the species (species nos. 28-35)

are very similar in dissection of the leaf. The



34 Illinois Natural History Survey Vol.34, Art. 1

pinnule base ascends obliquely on the basi-

scopic side and prolongates conspicuously on

the acroscopic side. This asymmetry imparts

a distinctive appearance to the pinnule base

(Fig. 48).

Relationships with Other Groups

Pteridologists have always classified Poly-

hotrya with Dryoptehs and such closely re-

lated genera as Arachniodes, Cyclodium.

Cyrtomium, Maxonia, Olfersia, Polystichop-

sis, Polystichiim. and Stigmaiopteris. These

genera share the following features with Poly-

botrya: base chromosome number .v = 41,

dryopteroid costa-costule architecture, spores

monolete with prominently inflated perispo-

rial folds, more than three vascular bundles

in the petiole, and petioles and petiolules not

articulate at their bases.

Familial Relationships

The dryopteroid genera are most closely re-

lated to the tectarioid ferns, such as Ctcnitis.

Lastreopsis, and Tectaria. These genera con-

stitute a natural group on the basis of their

costa-costule architecture and special

"C/fvi/V/.s-hairs." As noted in the Morphology

and Anatomy section of this monograph,

Polyhotrya lacks Crt'/;/7/.v-hairs, and their ab-

sence argues strongly against a close relation-

ship with the tectarioid genera. Furthermore,

Polyhotrya has costa-costule architecture of

the dryopterioid type rather than the tectarioid

type.

Polyhotrya might possibly be construed

as a member of the Lomariopsidaceae. Three

genera of Lomariopsidaceae

—

Lomagram-

ma, Lomariopsis, and Teratophylltim—
closely resemble Polyhotrya in overall habit

because they have densely scaly, high-

climbing stems, strongly differentiated sterile

and fertile leaves, and nonindusiate, acros-

tichoid sori. Furthermore, at the anatomical

level, the three genera have diplodesmic ve-

nation and dark, sclerenchymatous sheaths

surrounding each meristele in the stem. Al-

though no pteridologist has ever classified

Polyhotrya with the Lomariopsidaceae, these

similarities prompt me to consider the possi-

bility.

Stem anatomy is tremendously important

in the taxonomy of Polyhotrya and the

Lomariopsidaceae. The stem anatomy of

Polyhotrya is unique and has been thoroughly

described in the Morphology and Anatomy

section. The stem anatomy of the Lomariop-

sidaceae is also unique among ferns because

it has a broad, strap-shaped, ventral meristele

that differs from the remaining circular or

oblong meristeles (Fig. 16f,h). \jkt Polyhot-

rya. each meristele is surrounded by a dark,

sclerenchymatous sheath. This elongated

ventral meristele creates a dorsiventral dic-

tyostele. Presumably, this ventral meristele

was formed during phylogeny by the fusion

of two, once-distinct meristeles like the upper

ones (Holttum 1978). In fact, the two ventral

meristeles have not completely fused in

Lomariopsis, as evidenced by the shallow

indentation in the ventral band (Fig. I6f).

The ancestor of the Lomariopsidaceae, there-

fore, had a radially symmetrical dictyostele

of several to many, circularly arranged meri-

steles with each meristele surrounded by a

dark, sclerenchymatous sheath. In other

words, the ancestral stem anatomy was like

that of Polyhotrya. If one looks at Figure

16g. a cross section of a Polyhotrya stem,

and imagines what it would look like if the

two or three ventral meristeles were fused

into a single meristele, the result would be a

replica of the stem anatomy of the Lomariop-

sidaceae. This compelling similarity in stem

anatomy suggests that Polyhotrya and the

Lomariopsidaceae arose from a similar an-

cestral slock among the dryopteroid ferns.

The possibility also exists, however, that

parallel evolution accounts for these

similarities in stems adapted for climbing.

The leaf architecture of the two differs,

hovve\er. and most species of Lomariop-

sidaceae have siniph pinnate leaves. The

leaves of Polyhotrya. like those of most

drNopteroid ferns, are primitively decom-

pound. All Lonianopsidaceae, except B()/-

hitis. have articulate leaves and/or pinnae. In



November 1987 Monograph of Poi.ybotra 35

contrast. Polyhutrya and other dryopteroid

genera have nonarticulate leaves and/or pin-

nae. In the climbing genera Lomuriopsis and

Teratophyllum. leaves on the terrestrial por-

tion of the stem (bathyphylls) differ greatly

from those on the scandent portion (ac-

rophylls). In Lomahopsis, the bathyphylls

are less divided (usually simple and entire)

than the acrophylls, and in Teratophyllum

they are more dissected than the acrophylls.

In Polybotrya . however, the bathyphylls and

acrophylls are about the same size and shape.

Venation is yet another difference. Sev-

eral kinds of venation occur in the three gen-

era of Lomariopsidaceae most similar to

Polybotrya. Teratophyllum has simple or

forked veins, as does Lomariopsis. except

that the veins of the former unite with the

cartilaginous margin (Holttum 1978).

Polybotrya also has free veins, but the pattern

is entirely different from that of these two

genera, as can be seen by comparing the il-

lustrations in this monograph with those

.shown by Holttum (1978, Figs. 1-8). The

veins ofLomagramma differ completely from

those of these three genera; its veins form a

network of three or more rows of oblique

areoles without main veins. Venation, there-

fore, does not support a relationship between

Polybotrya and the Lomariopsidaceae.

In summary, Polybotrya and some genera

of Lomariopsidaceae share the following

characteristics; climbing stems, dimorphic

sterile and fertile leaves, diplodesmic veins,

and dark sclerenchymatous sheaths surround-

ing each meristele. The two groups, however,

contrast sharply in such features of leaf ar-

chitecture as the amount of dissection, pinnae

articulation, acrophylls versus bathyphyll dif-

ferences, venation patterns, and stem

anatomy. This conflicting evidence is dif-

ficult to assess. Certainly, the climbing habit,

sterile-fertile leaf dimorphy, and diplodesmic

veins have arisen many times in ferns, but

the similarity in stem anatomy is less easily

explained. I suspect, however, that the

Lomariopsidaceae may have had a separate

origin among the dryopteroid ferns, apart

from Polybotrya. Strong evidence exists that

Polybotrya was derived from a Cyclodium-

like ancestor (see below).

Relation to Similar Dryopteroid Genera

Maxonia. This monotypic genus was first

described by Christensen (1916), who ob-

served that his new genus "must stand be-

tween Polybotrya and certain species of

Dryopteris grouped with D. ampli.s.sima

[Arachniodes]." Copeland (1947) also held

that Ma.xonia apiifolia (Swartz) C. Chr. rep-

resented a phylogenetic intermediate between

Arachniode.s and Polybotrya (Fig. 17), and

most later pteridologists have agreed that

Ma.xonia is intimately related to Polybotrya.

Maxonia and Polybotrya appear very

similar because both have highly differen-

tiated sterile and fertile leaves and densely

scaly, climbing stems. Nevertheless, a com-

parison of stem anatomy shows some impor-

tant differences (Fig. I6e,g). The most evi-

dent dissimilarity is that each meristele of

Ma.xonia is not surrounded by a dark scleren-

chymatous sheath as in Polybotrya. Also, the

dictyostele of Ma.xonia is dorsiventral

(Chandra 1975; Walker 1972), unlike that of

Polybotrya, which is radially symmetrical.

Clearly, Mcixonia lacks the stem anatomy that

characterizes Polybotrya, thus weakening the

hypothesis that these genera are closely

related.

Strong evidence exists that Maxonia is

most closely related to the American species

of Arachniodes, in particular y4. macrostegia

(Hooker) Tryon and Conant. Indeed, if

Maxonia apiifolia had monomorphic leaves

it would without doubt be placed in

Arachniode.s. The similarities between the

two genera are striking and are best seen in

the sterile leaves. Both genera have pinnules

arranged anadromically, basal pinnae elon-

gated basiscopically, and laminae broadened

notably at the base and of similar thickish

texture. In addition, the pinnules and smaller

segments have cuneate bases and acute

apices, thereby imparting a characteristic

"streamlined" appearance to the lamina that



36 Illinois Natural History Survey Vol. 34. Art. 1

contrasts sharply with the truncate or acro-

scopically prolonged bases and rounded

apices in Polyhotrya. Both Maxonia and

Arachniodes have costa-costule architecture

of the dryopteroid type (grooves decurrent

into each other; Holttum 1984). Finally, both

genera have remarkably similar brown, thick-

ish, round-reniform indusia.

The evolution of the climbing stem of

Maxonia poses no problem \i Arachniodes is

accepted as the ancestor. Since the American

species oi Arachniodes have creeping stems,

the potential to evolve a fully hemiepiphytic

stem, as in Maxonia, was initially present

and easily achieved.

In short, Maxonia is similar to Polyhotrya

because it arrived at a comparable evolution-

ary grade or level, but it came from a different

source (Fig. 17). Polyhotrya was probably

derived from Cyclodium (see below), and

Maxonia from Arachniodes, probably from

an ancestor close to -4. macrostegia. Maxonia

should not, however, be subsumed with

Arachniodes because it is derived from it. It

is convenient, and certainly in harmony with

past taxonomic practice, to distinguish

Maxonia generically on the basis of its leaf

dimorphism and climbing stem.

Olfersia. This genus consists of a single

species, O. cervina (L.) Kunze, the place-

ment of which has been controversial. Most

recently, pteridologists have placed Olfersia

in Polyhotrya because both have strongly di-

morphic sterile and fertile leaves, nonin-

dusiate sori, similar perispore morphology,

and densely scaly, creeping stems. Yet Olfer-

sia differs from Polyhotrya by its venation,

conform apical pinna, and stem anatomy.

Moran ( 1 986) studied Olfersia and concluded

that it and Polyhotrya are sister taxa best

maintained in separate genera.

Cyclodium (sensu Smith 1986). Polyhot-

rya was probably derived from an ancestor

which, if it were alive today, would be placed

in Cyclodium. This genus has two important

prerequisites for the evolution of Polyhotrya.

First, the fertile leaves of Cyclodium are

slightly to strongly dimorphic. Second, the

stem is creeping in all species of Cyclodium,

and in several species it becomes fully

hemiepiphytic. Cyclodium. therefore, has the

genetic capacity to evolve two cardinal fea-

tures of Polyhotrya: holodimorphic sterile

and fertile leaves and a climbing stem.

Moreover, Cyclodium contains species

that look remarkably like certain sp)ecies of

Polyhotrya. The 2-pinnate-pinnatifid leaf of

C. trianae (Mett.) A.R. Smith var. trianae

is strikingly like some species of Polyhotrya

with respect to cutting and venation (cf. Fig.

16c to P. caudata. P. osmundacea. and P.

sessilisora. Figs. 28, 47, & 51 . respectively).

In fact, several times during fieldwork in

Ecuador, I mistook sterile leaves of C.

trianae for terrestrial leaves of P. caudata.

Similarity is also seen in the simply pinnate

lamina of C. meniscioides (Willd.) Presl,

which has anastomosing venation notably

like that found in Polyhotrya subgenus

Soromanes and is also simply pinnate (Fig.

16a,b). This similarity of venation was noted

long ago by Hooker and Baker (1874). I am

not suggesting that these two species of Cy-

clodium gave rise to Polyhotrya. but within

both genera, species have evolved with simi-

lar characteristics, and this homoloaous vari-

Polytx)trya

T
Maxonia

T
Arachniodes

Maxonia Polyhotrya

T t
Arachniodes Cyclodium

T T
Ancestor {Dryopteris?)

FiciURi 17. Two phylogcnctic hypotheses concerning the relationships of Maxonia and Polyhotrya:

left, the hypothesis proposed by Christensen (1916): right, the hypothesis proposed here. Sec text for

explanation of characters involved.



November 1987 Monograph of Poi.ybotra 37

ation implies a close genetic relationship. In

conclusion, the close relationship between

Cyclodium and Polybotrya is evidenced by

their mutual possession of dimorphic leaves,

creeping stems, similar evolutionary tenden-

cies, and, at least in some species, remarka-

bly similar leaf dissection and venation.

Part Two:

Taxonomic Treatment

//; evohitionary biology almost all phenom-

ena and processes are explained through in-

ferences based on comparative studies.

These, in turn, are made possible by very

careful and detailed descriptive studies. It is

sometimes overlooked how essential a com-

ponent in the methodology of evolutionary

biology the underlying descriptive work is.

Ernst Mayr (1982)

Notes on the Presentation of Data

In most cases, the key can be used with speci-

mens lacking fertile leaves. Fertile leaves,

however, are often helpful in identification

and are given as additional characters in many

couplets. The key will generally not work

when leaves are less than 45 cm long. To

facilitate the comparison of similar species,

1 have arranged the species phylogenetically

rather than alphabetically. The synonymy

given for each species is complete. 1 have

kept the descriptions short by not repeating

characteristics that are constant for the genus.

For example, all species of Polybotrya have

a pinnatifid leaf apex; this characteristic is

mentioned only in the generic description and

is not repeated in each species description.

The distribution maps (see appendix) were

compiled from the specimens listed in the

Specimens Examined section in each species

treatment. In some cases, I could not find

localities despite searching various atlases,

indices, and gazetteers.

The terms basiscopic and acroscopic are

used frequently in the key and in descriptions

to refer to the sides of pinnules (Fig. 18).

The acroscopic side of a pinnule is the side

directed toward the apex of the pinna bearing

the pinnule. Conversely, the basiscopic side

of a pinnule is that side directed toward the

pinna base and rhachis. Two other frequently

used terms, catadromic and anadromic , refer

to pinnule arrangement. The pinnule arrange-

ment is catadromic when the pinnule closest

to the rhachis is basiscopic; when the pinnule

closest to the rhachis is acroscopic, the ar-

rangement is anadromic (Fig. 18). I have de-

basiscopic

side

costule

rhachis

PINNULE

acroscopic

side

tertiary
segment

costa

catadromic

anadromic

PINNAE
FiuuRK 18. Terms frequently used to describe the leaves of /'o/yfcofrya. See text for further cxplantions.



38
Illinois Natural History Survey Vol. 34. Art. 1

fined the specialized terms that refer to the

fertile leaves, such terms as botryoid.

coenosoric, and amphiacrostichoid, in the

Morphology and Anatomy section and in the

Description of the Genus section.

Description of the Genus

POLYBOTRYA WILLD

Polybolrva Willd., Species Plantarum. ed. 4.

5;99. 1810. Type species; Polyhotrya osmun-

dacea Willd.

Soromanes Fee. Mem. Fam. Foug. 2 (Hist.

Acrost.) 16. 1845.Type species; Soromanes

serratifolium Fee = Polyhotrya serratifolia

(Fee) Klotzsch.

Bom'olhallus Klotzsch. Bot. Zeit. 4;104. 1846.

nom. nudum. Type species; Botryothallus

kunzei Klotzsch nom. nudum = Polyhotrya

serratifolia (Fee) Klotzsch.

Stem hemiepiphytic, long-creeping, or (two

species) terrestrial and short-creeping, 1 .0-3.5 cm

wide (excluding scales), the scandent portions un-

branched. the terrestrial portions occasionally

branched; dictyostelic. in cross section with a

characteristic pattern of 4-10 circularly arranged

meristeles. each meristele surrounded by a dark

sclerenchyma sheath; aerophores present in most

(all?) species, continuous and decurrent from the

lateral aerophores of the petiole, darkening upon

drying and not visible in herbarium specimens, in

fresh material appearing yellowish, linear, 1-3

mm wide, slightly raised and bearing stomata;

rootlets produced only on the ventral surface;

scales numerous, densely covering the stem,

spreading or appressed, primarily of two general

types; 1) thick, dark brown, opaque, margms en-

tire , attached across the length of the curved .
thick-

ened base, and 2) thin, variously colored (ranging

from yellow, orange, red to brown), translucent,

margins denticulate or highly erose. anached at a

central, basal point. Sterile leaves up to 2.0 m

long, reclining when mature, intcmodes generally

10-15 cm apart, bulbils lacking;/?*'//*'/*- with 8-16

vascular bundles arranged in a mushroomlike out-

line, the base decurrent for a short distance on the

stem, scaly, with scales shorter and wider than

those on the stem, the aerophores present laterally

as thin yellowish green lines, the adaxial surface

slightly tlattcned, trisulcate in dried material; color

stramineous, light green or darkened abaxially;

rhachis trisulcate below becoming unisulcate

above, the groove pubescent within; lamina

papyraceous to subconaceous. 1 -pinnate (sub-

genera Soromanes and Sorbifolia) or 1-pinnate-

pinnatifid to 4-pinnate (subgenus Polyhotrya),

usually lanceolate, the base rarely deltate or

cuneate, the apex pinnatifid; pinnae not articulate

to the rhachis. linear, lanceolate or deltate. sym-

metrical or slightly prolonged acroscopically. a

few species having the basal pair slightly pro-

longed basiscopically; pinnules arranged caia-

dromically or anadromically, symmetncal or

asymmetrical at the base, if asymmetrical then

with the basiscopic side reduced and oblique and

the acroscopic side prolonged; grooves ofcosiules

decurrent into the the grooves of the costa or trun-

cated by the ridges of the costa and therefore not

decurrent (intermediates exist); veins free or (in

subgenus Soromanes) anastomosing, ending close

to the margin, hydathodes absent: hairs of two

types: 1 ) all species have tiny, reddish, appressed.

jointed, flattened hairs on the lamina that grade

into the scales (these actually represent reduced

scales), and 2) most species have hairs that do not

intergrade with the scales, these are whitish and

cylindrical, ranging from unicellular and less than

0.1 mm long to 15 cells and 2.0 mm long, or

uncinate. Fertile leaves appearing as a skel-

etonized version of the sterile, more ephemeral

than the sterile, produced only on the scandent

portion of the stem in hemiepiphytic species: son

nonindusiate and of three types; 1 ) round, discrete,

usually at the tips of pinnately arranged stalks

(botryoid type). 2) oblong or linear formed by the

fusion of several different sori (coenosoric type),

and 3) sori apparently occupying both surfaces of

a caudate pinnule (amphiacrostichoid type); di-

plodesmic veins present except in botryoid sori:

sporangia with 64 spores per capsule, the capsule

glabrous except in P. puhens which has setose

hairs at the apex near the annulus: sporangial stalks

with three rows of cells at the apex, two-rowed

below, paraphvsate in most species by lateral

hairs, these multicellular and unbranched

(branched onlv in P. speciosa): annulus of 15-22

indurated cells: spores (32)45-65(80) microns

long, dark brown when viewed with transmitted

light under a compound microscope, deep orange

w'hen viewed with retlected light under a dissect-

ing microscope, monolete. aperaturc linear and

i/,-y4 the length of the long axis, exospore

smooth, pcrispore bilayered with inflated folds

and cchinate to various degrees. .v = 41.



November 1987 Moncxiraph of Polybotra 39

Key to the Species of Polybotrya

1. Sterile leaf 1-pinnate.

2. Veins of sterile leaf anastomosing (subgenus Saromanes).

3. Plants of Mcsoamerica; vein tips joined into a faint, continuous, submarginal connecting

strand 2. P. potyhotryoides. p. 80.

3. Plants of South America and Trinidad; vein tips usually, but not always, free.

4. Lamina pubescent abaxially with uncinate hairs. Western Cordillera of Ecuador.

5. Hairs on abaxial surface 0.1-0.3 mm long, colorless, erect, 1-3 celled; free pinnae pairs

6-13 below the pinnatifid apex, mostly 15-21 x 4-5 cm
3. P. suberecia, p. 50.

5. Hairs on abaxial surface 0.5-1 .2 mm long, tawny, spreading, 5-12 celled; free pinnae

pairs 4-7 below the pinnatifid apex, mostly 17-33 x 5.7-10 cm
4. P. andinii. p. 50.

4. Lamina glabrous abaxially.

6. Apex evenly pinnatifid, not similar to the lateral pinnae (Fig. 20a); vein tips free or in-

completely anastomosing.

7. Stem scales linear, stiffish, mostly 0.3-1.0 mm wide, usually opaque and concol-

orous; fertile pinnules more than 2.0 cm long. Colombia, Ecuador, and Peru. ...

3. P. suberecia, p. 50.

7. Stem scales lanceolate, flaccid, mostly 1-2 mm wide, often with a dark central stripe

and lighter borders; fertile pinnules generally less than 2.0 cm long. Andes of Vene-

zuela; Trinidad 1. P. serralifolia, p. 43.

6. Apex subconform, similar to lateral pinnae, or with one or a few small basal lobes

(Fig. 20b, c); vein tips joined into a faint continuous submarginal connecting vein (see

text for observation of this character). Southern Mexico to Peru

2. P. polybotryaides, p. 80.

2. Veins of sterile leaf free (subgenus Sorbifolia).

8. Stem terrestrial and short-creeping; scales brown, essentially concolorous; fertile pinnules com-

monly round or oblong, usually less than 1 cm long.

9. Largest pinnae 7-10 times longer than broad, abaxial surface often with white or light

brown, sessile, globose glands; apex of 3-7 pinnalike lobes (Fig. 24a), these with

long decurrent bases; stem scales shiny brown, mostly transparent and denticulate; spores

40-47 microns long 5. P. sorbifolia, p. 53.

9. Largest pinnae 5-7 times longer than broad, abaxial surface without glands; apex of 2 or 3

lobes, their bases not long decurrent (Fig. 25b, c); stem scales dull brown, mostly opaque

with entire or subentire margins; spores mostly 52-56 microns long

6. P. fractiserialis. p. 54.

8. Stem hcmicpiphytic and long-creeping; scales reddish brown or orange, with a prominent cen-

tral stripe and lighter borders; fertile pinnules linear and usually more than 1 cm long

1. P. crassirhizoma, p. 58.

1. Sterile leaves 1-pinnatc-pinnatifid to 4-pinnatc (subgenus Polyholrya).

10. Pinnules of medial pinnae catadromic (Fig. 18).

11. Rhachis and costae pubescent, hairs 1.0-2.5 mm long. Amazonian lowland forests.

12. Lamina more than 15 cm wide at the base; petiole more than 15 cm long; largest pinna

lobes with entire margins; stem scales usually denticulate, translucent, cream to casta-

neous; fertile pinnules caudate. 3-15 mm wide, sori acrostichoid, apparently covering

both surfaces; sporangial capsules setose (Fig. 30e,0 \\. P. puhens, p. 68.

12. Lamina up to 10 cm wide at the base; petiole up to 4 cm long; largest pinna lobes with ser-

rate margins; stem scales entire, dark, and opaque; fertile pinnules botryoid, sori round,

di.scretc, 1-3 mm wide; sporangial capsules glabrous

12. P. f>lundulosa . p. 71.

1 1 . Rhachis and costae glabrous, or if pubescent, hairs shorter than I mm long. Amazonia and

elsewhere.



40 Illinois Natural HrsTORV Survey Vol. 34. Art. 1

1 3 . Ultimate segments or lobes of sterile leaf 0.5 - i . 5 mm wide and only one-nerved; lamina often pubes-

cent on both surfaces. Andes of Colombia to Bolivia, Guyana 13. P. lechleriana. p. 71.

13. Ultimate segments or lobes of sterile leaf more than 1 mm wide and with several nerves; lamina rarely

pubescent on both surfaces.

14. Stem scales bright golden or yellowish a/u/ the pinnatifid portions of the pinnae or pinnules with

a single veinlet running towards the sinus arising directly from the costa or costule between the

main lobes or vein groups (Fig. 45d). Andes of Colombia to Peru

26. P. aliescandens. p. 97.

14. Plants without the above combination of characters.

15. Laminar margins sparsely ciliate, hairs minute, less than 0.1 mm long (Fig. 28a) d/u^ fertile

pinnules caudate, son amphiacrostichoid, covering both surfaces of the leaf.

16. Stem scales dull brown, opaque, appressed-ascending, margins subentire, base curved

and thickened (Fig. 28h). Widespread 9. P. caudata, p. 60.

16. Stem scales shiny reddish to castaneous. membranous, spreading, margins dentic-

ulate, base usually cordate at point of attachment (Fig. 29f). Paraguay and Brazil.

10. P. goyazensis. p. 66.

15. Laminar margins glabrous, or if sparsely ciliate, fertile pinnules not caudate and sori not am-

phiacrostichoid.

17. Abaxial surface of sterile leaf pubescent, hairs fine, erect, whitish, less than 0. 1 mm
long and costae scaly with numerous, golden brown, tortuous scales. Panama

23. P. alata. p. 92.

17. Plants without the above combination of characters. Panama and elsewhere.

18. Plants nearly glabrous throughout, even within the grooves and pinnule mar-

gins cut less than Vi of the way to the costule. Costa Rica

27. P. gomezii, p. 99.

18. Plants usually pubescent on the major axes and within the grooves; pinnules

cut more than % of the way to^e costule.

19. Lamina margins sparsely ciliate, hairs 0.1 mm long and stem scales

opaque, appressed, the base curved and thickened. Southeastern Brazil.

20. P. cylindrka. p. 86.

19. Lamina margins glabrous; stem scales as above or thin, spreading, and trans-

lucent. Plants not from southeastern Brazil.

20. Pinnule bases more or less symmetrical (Fig. 35); tertiary segments often

inserted at about right angles to the costule. ovate to oblong, usually

less than 2.5 (3.0) times longer than broad.

21 . Pinnules up to 2.5 X 1 .2 cm. with only 5-7 segments and;or lobes;

lamina up to 55 x 26 cm. apex long-attenuate; major axes with nar-

row, dark, tortuous scales. Colombia

14. P. attenuata. p. 74.

21. Pinnules larger than 2.5 x 1.2 cm, with more than 5 segments

and/or lobes; lamina often larger than 55 x 26 cm, apex rarely long-

attenuate; major axes rarely with narrow, dark, tortuous scales.

22. Sterile lamina slightly reduced at the base; basal pinnae less than

3.5 times longer than broad, usually elongated basiscopically;

fertile leaves coenosone; sori not stipitatc but placed close to

the midrib.

23. Sterile lamina pubescent adaxially. Colombia

15. P stohei. p. 76.

23. Sterile lamina glabrous adaxially.

24. Margins of tertiary segments entire to crcnate. serrate

or lobcd; pinnules of medial pinnae cut more than half

way to the costule for most of their length in pinnae 25

cm or longer, Mcsoamcrica. Ecuador, and Bolivia.

16. P. alfredii. p. 78.



November 1987 Moncxiraph of Polybotra 41

24. Margins of tertiary segments entire, never crenate, serrate, orlobcd; pinnules of medial

pinnae cut less than half way to the costulc for most of their length. Colombia. ...

18. P. lourleigiana, p. 82.

22. Sterile lamina broadest at the base; basal pinnae 3.5-4.0 times longer than broad, not

strongly elongated basiscopically; fertile leaves botryoid; son stipitate, stalks 1-4 mm long.

Colombia 19. P. pittieri, p. 84.

20. Pinnule bases acroscopically prolonged (Fig. 44); tertiary segments and lobes usually inserted

obliquely to the costule, somewhat elongate and falcate, more than 2.5 times longer than broad.

25. Costal scales scattered, dark, ovate to lanceolate, flaccid (Fig. 52b); fertile leaves botryoid

(Fig. 52c). Cloud forests of northern Venezuela 32. P. canaliculala. p. 111.

25. Costal scales absent, or if present, then linear, tortuous; fertile leaves rarely botryoid.

26. Sterile lamina 3-4-pinnate-pinnatisect, ultimate segments or lobes 1-2 mm wide;

major axes on all sides with numerous, narrow (1-3 cells wide), reddish brown, tortu-

ous, spreading scales; fertile leaves botryoid. Colombia

17, P. botnoides, p. 80.

26. Sterile lamina to only 3-pinnate-pinnatifid, ultimate segments or lobes more than 1-2

mm wide; major axes lacking numerous scales, or if scales present, then not as above.

27. Lamina puberulent on both surfaces; veins prominulous. Andes of Ecuador and

Bolivia 22. P. puberulenia, p. 90.

27. Lamina always glabrous adaxially and usually so abaxially; veins not promin-

ulous.

28. Stem scales dull brown, thick and opaque, attached across the length of the

curved and thickened base, margins subentire.

29. Tertiary segments lobed, lamina usually 3-pinnate-pinnatifid.

30. Costae evenly pilosulous, hairs less than 0.1 mm long; costular

grooves truncated by the ridgesof the costae and therefore notdecur-

rent; grooves glabrous or nearly so within. Andes of Colombia and

Bolivia 21. P. hickeyi, p. 88.

30. Costae glabrous or with only scattered hairs; costular grooves decur-

rent into the costal grooves; grooves filled with reddish or brownish

hairs. Widespread species 28. P. osmundacea, p. 101.

29. Tertiary segments entire or only minutely serrate at the apex; lamina

mostly 2-pinnate-pinnatifid. Andes of Ecuador

25. P. appressa, p. 94.

28. Stem scales reddish brown or golden, thin and translucent, attached at a cen-

tral point, base not curved or thickened, margins denticulate to erose.

31. Stem scales reddish brown; lamina with punctate, resinous glands; costal

grooves packed with protruding hairs 0.3-0.8 mm long. Guadeloupe

and Martinique 29. P. cyathifolia, p. 106.

31. Stem scales golden; lamina lacking punctate, resinous glands; costal

grooves glabrous. Andes of Ecuador

24. P. aequatoriana. p. 94.

10. Pinnules of medial pinnae anadromic (Fig. 18).

32. Stem scales yellowish to golden; lamina usually with a single veinlet springing directly from the

costa or costule between the main lobes or vein groups (Fig. 45d). Andes of Colombia, Ecuador,

and Peru 26. P. altescandens. p. 97.

32. Stem scales brown, castaneous, or reddish; lamina lacking a veinlet as described above.

33. Sterile leaves 2-pinnate; pinnules entire, bases symmetrical andcuneate; fertile pinnules cy-

lindrical and entire. Known only from the state of Espirito Santo, southeastern Brazil.

8. P. cspiritosantensis. p. 60.

33. Sterile leaves 2-4-pinnate; pinnules divided, lobcd, or serrate, the bases usually prolonged

acroscopically; fertile pinnules lobed and/or divided.

34. Scales of the petiole base 5-6 mm wide, broadly ovate, castaneous. Colombia. ...

30. P. lalisi/uanwsa, p. 108.



42 Illinois Na I URAi. HisiOKY Survey Vol.34. Art. 1

34. Scales of the petiole base less than 4 mm wide, usually narrowly lanceolate, brown, castaneous, or

reddish.

35. Abaxial surface of lamina evenly pubescent; stem scales reddish. Southeastern Brazil.

36. Costal hairs less than 1 mm long 34. P. speciosa, p. 115.

36. Costal hairs 1-2(2.5) mm long 35. P. pilosa, p. Ml.

35. Abaxial surfaccof lamina glabrous; stem scales variously colored. Southeastern Brazil and else-

where.

37. Fertile leaves botryoid, i.e.. all ultimate soriferous segments round and discrete, not fusing

to form an oblong or linear sorus (Fig. 52c).

38. Sterile lamina to 4-pinnate; stem scales dull to unaided eye. dark brown, with entire or

subcntire margins; costal scales broadly ovate to lanceolate; costae glabrous or more

rarely pubescent; hairs short, reddish, less than 0.1 mm long; sori short-stalked (Fig.

52c). Cloud forests, Andes of Venezuela 32. P. canaliculaia. p. 111.

38. Sterile lamina to 3-pinnate; stem scales shiny, dark castaneous, with denticulate-erose

margins; costal scales narrowly lanceolate to linear; costae often pubescent, hairs 0.5-

1 .0 mm long, whitish; sori sessile (Fig. 51c). Lowland forests, northern Amazon basin

and Guiana Highlands 3\. P. sessilisora. p. 108.

37. Fertile leaves coenosoric, i.e., some or most of the ultimate soriferous segments, especially

those near the apex, fusing to form an oblong to linear sorus (Fig. 47g).

39. Abaxial surface of costulcs evenly hirsute, hairs less than 0. 1 mm long, or if glabrous,

stem scales reddish. Southeastern Brazil.

40. Stem scales reddish; margins of tertiary segments or lobes crenate to lot>ed. ..

34. P. speciosa. p. 115.

40. Stem scales bright castaneous; margins of tertiary segments or lobes entire. ...

33. P. semipinnata. p. 113.

39. Abaxial surface of costulcs glabrous or variously pubescent by soft whitish hairs more

than 0.1 mm long; stem scales rarely reddish. Plants not of southeastern Brazil.

41

.

Costal grooves glabrous within; sterile lamina mostly 2-pinnate-pinnatifid. Costa

Rica 27. P. gomezii. p. 99.

41 . Costal grooves pubescent within, hairs reddish to brownish; sterile lamina mostly

3-pinnate-pinnatifid.

42. Basal acroscopic segment of pinnules usually with a slight basal gibbosity on

both margins (Fig. 49c); lamina membranaceous, almost always with reddish

punctate glands abaxially. Guadeloupe and Martinique

29. P. cyathifolia. p. 106.

42. Basal acroscopic segment of pinnules lacking basal gibbosity, usually slight-

ly reduced or oblique (Figs. 47 & 48); lamina thicker, papyraceous to charta-

ceous, rarely with reddish punctate glands abaxially Widespread

28. P. osmundacea. p. 101.



November 1987 Monograph of Poi.ybotra 43

Species Descriptions

POLYBOTRYA subgenus SOROMANES
(Fee) Moran comb. & stai. now

Soromanes Fee, Mem. Fam. Foug. 2 (Hist.

Acrost.) 16. 1845. Typf. sPF.crEs: Soromunei-

serraiifolium Fee = Polybotrya serratifolia

(Fee) Klotzsch.

Polybotrya section Soromanes (Fee) Klotzsch,

Linnaea 20:430. 1847.

Polybotrya subgenus Soromanes (Fee), attri-

buted incorrectly to Klotzsch by Fee, Genera

Filicum 50, invalid.

Acrostichum subgenus Soromanes Hooker,

Species Filicum 5:256. 1864. Type species:

Acrostichum caenopieris Hooker = Polybotrya

serratifolia (Fee) Klotzsch.

Sterile leaves I -pinnate; veins from adjacent

costules anastomosing, 4-6 pairs, curved ascend-

ing, the basal pair joining 'A- 'A the distance from

the margin to the costa, then with an excurrent

vein to the base of the above anastomosing pair,

the distal vcinlcts strongly curved towards the

apex, vein tips at the margin free or, in P. pohbot-

ryoides. joined by a submarginal vein. Fertile pin-

nae pectinate.

1. Polybotrya serratifolia (Fee) Klotzsch (Fig.

19, Map 1).

Polybotrya serratifolia (Fee) Klotzsch, Lin-

naea 20:430. 1847.

Soromanes serratifolium Fee, Mem. Fam.

Foug. 2 (Hist. Acrost.) 82, tab. 43. 1845.

Type: P! (3 sheets), in Bory Herbarium, cited

by Fee as "Polybotrya serrala, Galeotti, in

herb. Bory, Habitat in Mcxicana rcpublica,

Lagunetta (Galeotti). - V.S. in herb. Bory."

but probably collected by Linden in Ven-

ezuela and later distributed by Galeotti (sec

text discussion). I have selected as the Icc-

totypc that sheet which Fee illustrated for his

tab. 43.

Soromanes dentatum Fee, Mem. Fam. Foug.

2 (Hist. Acrost.) tab. 43. 1845. Fee's tab. 43

was erroneously labeled as "S. dentatum" in-

stead of 5. serratifolium . His tab. 43 is a

precise illustration of the lectotypc of P. ser-

ratifolia.

Soromanes integrifolium Fee, Mem. Fam.

Foug. 2 (Hist. Acrost.) 82, tab. 42. 1845.

Fertile leaf only, the sterile one is that of

Cyctodium meniscioides. Type: Alexander

Braun Herbarium (B).

Botryothallus kunzei Klotzsch, Bot. 2^it.

104. 1846. nom. nudum, cited by Ettings-

hauscn (1864).

Polybotrya kunzei Ettingshausen, Denkschr.

Akad. Wien. 22:66. fig. 2. 1864. Type: Only

"in Colombia" was mentioned.

Acrostichum caenopteris Hooker, Species

Filicum. 5:256. 1864. nom. nov. for Soro-

manes serratifolium Fee, non Kaulf. 1824,

with same type.

Polybotrya coenopteris (Hooker) Christ,

Famkr. 44. 1897.

Soromanes coenopteris (Hooker) Christ,

Bull. Herb. Boissier, II. 3:613. 1903.

Acrostichum hartii Baker, J. Bot. 371 . 1881

.

Type: Trinidad, War/ 22S(holotypc:K, photo

at US!; isotype: NY!).

Polybotrya hartii (Baker) C. Chr., Index

Filicum. 504. 1906.

Polybotrya crassa Morton, Fieldiana Bot.

28:13. 1951. Type: Venezuela. Monagas:

south-facing forested slopes above lime-

stone bluffs, northeast of Guacharo, alt. 1300

-1400 m, 1 1 April 1945, Sleyermark 61991

(holotype:US!; isotypes: F!, VEN!).

5/emhemiepiphytic, 1 .5-3.0cmthick;scu/ej

0.8-1.5 mm long, 0.5-2 mm wide, narrowly

lanceolate, reddish brown to light orange, with or

without a prominent dark central stripe, membran-

ous, lustrous, spreading, margins highly erose to

merely denticulate at the apex. Sterile leaves up

to 0.8 m long, dull green adaxially, pale green

abaxially, chartaccous to subcoriaceous; petiole

scaly at base, up to 30 cm long, trisulcate and

flattened adaxially; lamina up to 50 x 27 cm, I-

pinnate; pinnae 6-12 pairs, mostly (7)10-

20(23) X (2)3-4.5(5.5) cm wide, lanceolate, base

rounded, cuncatc to truncate, margins entire to

serrate, apex acuminate; veins in pinnated groups

3-6 mm apart, the tips arcuate, free; axes with a

few scattered scales, rarely pubescent, the hairs

tiny, less than 0. 1 mm long, unicellular, whitish;

grooves nearly glabrous within. Fertile leaves 2-

pinnate, often about the same length as the sterile



44 Illinois Natural His iory Survky Vol. 34. Art. 1

I cm
F.GURF 19 PoWhcn'a serranfolia (Fde) Klo.^sch. a. habit; b. stem scales (note ca.se margins and

d°k central stnpes,; c. petiole scale; d. s.enlc pmna; e-g. fertUe p.nnae show.ng vanat.on .n cutung^

oarK central sinptM, y^
j ,^ w -rff ^ WiiH'flfUl UW {MO) d: Aristei;uieU2 .^963

h. sporangium with paraphysis. a-c; van der Wcrjf & Ww^JwUi -^•^- " *'^'y'

(VEN). c Fendler 261 (MO), f.h: Broadway 9947 (GH). g: type, probably Luuicn (P).



November 1987 Monograph of Poi ybotra 45

leaves; pinnae linear to narrowly triangular,

evenly long-tapered to apex, up to 14x2.5 cm;

pinnules entire to occasionally slightly lobate,

linear, oblong or sometimes clavate . 4 - 1 5( 20) mm
long, 2-3 mm wide, the adaxial margins folding

together at maturity, giving the pinnules a cylindri-

cal appearance; sori coenosoric. continuously cov-

ering the pinnule, occasionally lobcd at the base

or botryoid; sporangial stalks paraphysate; spores

(44)50-60(63) microns long. n = A\.

Other illustrations: Fee's tab. 43 is an excel-

lent, precise illustration of the lectotype; Vareschi.

Fl. Venez.. Helechos. vol. 1, tab. 71. 1969 (as

P. crassa).

Polybotrya serratifolia grows in undisturbed,

wet, premontane and cloud forests from 1200-

2400 m elevation. It is known only from the moun-

tains of northern Venezeula and Tnnidad (Map

I). Van der Werff and Smith (1980) report this

species (as Polybotrya sp., aff. crassa) from the

state of Falcon, Venezuela, where it grows in wet

premontane forest with another Venezuelan en-

demic, P . canaliculata.

The distinctive, wide, flaccid, crose scales of

P. serratifolia (Fig. 19b,c) distinguish it from the

other species in subgenus Soromanes . Polybotrya

serratifolia differs from the similar P. polybot-

ryoides by its submarginal connecting vein. This

vein, however, is not easily seen because the con-

necting vein of P. polybotryoides is very faint and

the vein tips of P. serratifolia. though free, are

arcuate and simulate a submarginal connecting

vein.

Polybotrya hartii and P. crassa are placed in

synonymy, although at first sight they look differ-

ent. Both were originally distinguished from P.

serratifolia by their more dissected (botryoid) fer-

tile leaves. But these botryoid leaves represent a

break-up of the coenosorus— a phenomenon that

I interpret as atavistic. These highly divided fertile

leaves arc not uncommon; even the type at Paris

contains a sheet with a 3-pinnatc fertile leaf (Fig.

19g). As might be expected, intermediates exist

between coenosoric and botryoid fertile leaves

(Fig. 190- Still, the most common type of fertile

leaf in P. serratifolia is 2-pinnate (coenosoric)

with relatively short pinnules (Fig. 19a, c).

In addition to its finely divided fertile leaf,

Polybotrya crassa was distinguished by having an

acroscopic auricle and several lobes at the pinna

base. This condition represents nothing more than

a part-fertile, part-sterile leaf; such transitional

forms arc commonly found in other species of

subgenus Soromanes and frequently cause

taxonomic confusion. Accordingly. P. crassa is

here placed in synonymy.

Fee's citation of the type locality and collector

("Mexicana republica. Lagunctta (Galeotti). -V.S.

in herb. Bory")seemstobe the result of confusion.

One of the labels on the type specimen reads (my

translation): "Lagunetta, sent from Galeotti, Oc-

tober 1845." The word Lagunetta appears without

mention of Mexico. Fee probably thought that

Galeotti had collected the specimen in Mexico,

the principal American country in which Galeotti

collected (Morton 1971:63). Galeotti returned to

Europe in 1840, where he sold many duplicates

of his own and other collections. Morton (1971)

observed: "Galeotti collected only in Mexico and

a few specimens in Cuba but is often cited as the

collector of plants from Brazil, Venezuela, and

Colombia, but these plants were actually collected

by Linden, and Galeotti was merely the distributor

of the Linden plants." Presumably, Jean Jules Lin-

den collected the type of P. serratifolia, which

was later distributed by Galeotti. Although Linden

collected in Cuba, Mexico, and Guatemala, his

last and most profitable trip was to Venezuela and

Colombia (1841-1844) where, based on other col-

lections, this species is known to occur. Polybot-

rya serratifolia has not been collected in Mexico.

I cannot find a town called "Lagunetta" in Mexico,

but a town with that name is located in the state

of Lara, Venezuela, and I suspect that is where

Linden collected the type.

Specimens examined: Trinmjaij. Presioe 1491

(MO). 1492 (MO); heights of Aripo. Broadway 9947

(GH. US). 9949 (F, NY. US). Fay H59 (BM),

Venezuki.a. Aragua: Cordillera Inierior. Cerro

El Pauji. TopoF.I Pauji.al surdcElConsejo.iVcvt'rmart

cS Sloddari IIS051 (GH. VEN);ccrcaTcjerias.V«r«eW

7764 (YEN); Parque Nacional "H. Pillicr." bosque dc

Rancho Grande. Tschudi 167 (YEN); Colonia Tovar.

Fendler 235 (BM). 261 (GH. K. MO. NY. PH. US);

Colonia Tovar. Moriiz 277 (BM). Falcon: Curimagua

(Core), van Cotthem 1327 (UC); Sierra de San Luis.

Montana de Paraguariba. van der Werff & Winnfield

3430 (MO) Distrito P'ederal: between El Junquito and

Colonia Tovar. Sleyermark 91756 (GH. YEN); El Jun-

quito. Schnee 615 (YEN); off road Caracas-Colonia

Tovar. in Ibresl below Club Jundolandia. Rirry 948

(YEN). Lara: Lagunetta. (jtdeolli \.ii. (P); Dito Moran,

8.7 mi SE of Sanare. Parquc Nacional Yacambu. A.R.

Smith 1259 (PORT. UC); Dito. Moran. Rivero el at.

I60fi (PORT). Dlto. Iribarrcn, Parquc Tercpaina, Iriarie

52 (PORT); Dtio Andres Elroy Blanco. 7 km de Sanare.



46 Illinois Natural History Survey Vol.34, Art. I

Rivera 5I3A (PORT); sclva arriba de Sanare, Aris-

leguiela 396J (US, VEN); 22(X) m sobre los bafios de

Ri'o Claro, A.R. Smith 4055. Merida: verlientes del Rio

Capaz, arriba de La Azulila, Sleyermark & Ruhe 97133

(VEN); rich forest above Hacienda Agua Blanca, above

La Azulita, Sleyermark 56114 (F, US) Monoagas:

south-facing forested slopes above limestone bluffs,

northeast of Guacharo, all. 1300-1400 m. II Apnl

1945, Sleyermark 61991 (F, US. VEN). Portuguesa:

15 km E of Chabasquen, 67 km NNW of Guanare,

Sleyermark et al . 726675 (PORT). /266W) (UC, VEN).

Trujillo: arriba dc Escuquc, entre Estuque y La Mesa

de San Pedro, Sleyermark 104717 (MO, VEN); Dtto.

Bocono, ca. 10 mi SW of Batalal on road to Bocono,

Laguna dc Aguas Negras, A.R. Smilh etal. 922 (PORT,

UC); 2 km NW of Caserio Cerros de Guaramarcal, 42

km SE of Bocono, Moran 3709 (F, MO, PORT, VEN).

Yaracuy: El Amparo hacia Candelaria, a 7-10 km al

Norte dc Salom, Sleyermark el al. 106758 (NY, MO.
PORT); Distnto Bruzual, Serram'a de Aroa, 1 1-15 km
NNE of Urachichc, 3 km NE of Caserio Buenos Aires,

Sleyermark et al. 124749 (PORT, UC, VEN); Dtto.

Bolivar, entre las Parchitas, Tierra Fria y Ojo de Agua,

Onega <$ Smilh 2491 (PORT), 2510 (PORT).

2. Polybotrya polybotryoides (Baker) Christ

(Figs. 20& 21, Map 2).

Polybotrya polybotryoides (Baker) Christ,

BulL Herb. Boissier, U. 1:70. 1901.

Acrostichum polybotryoides Baker, J. Bot.

207. 1881. Type: Colombia. Norte de San-

tander: Ocaiia, on trees in the forest, 70(X)

ft., Kalbreyer 1254 (holotypc: K, color slide

at MO!; photo GH!, MO!).

Acrostichum juglandifolium Baker, J. Bol.

207. 1881. nom. illeg.. turn Kaulfuss, 1824.

Type: Colombia. Antioquia: Kalbreyer I79S

(holotypc: K, color .slide at MO!; photo GH!,

MO!).

Polybotrya juglandifolia Christ, Bull. Herb.

Boissier, II. 4:96.S. 1904. nom. now for

Acrostichum juglandifolium Baker, non Kaul-

fuss, with same type.

Polyholrya juglandifolia Christ var. lobata

Christ, Bull. Herb. Boissier, II. 6:168. 1906.

Type: Costa Rica. Cartago: Rio Navarro,

Coll. Inst. Costaricensis no. 16769. Werckle

s.n. (P?).

Polybotrya kalbreyeri C. Chr., Index

Filicum. 504. 1906. nom. superfl. for

Polybotrya juglandifolia Christ; with the

same type.

Polybotrya aucuparia Christ, Bull. Herb.

Boissier, II. 6:166. 1906. Type: Costa Rica.

Cartago: Valley of the Ri'o Navarro, 1400 m,

Werckle 16770 (P!; photos F!, NY!, UC!).

S/em 0.5 -2.0cm thick, hemiepiphytic; 5ca/«

linear, mostly 9-15 mm long, 0.5-1 .0 mm wide,

ascending with spreading tips, lustrous, golden

yellow when fresh, turning yellow brown to purple

brown upon drying, margins denticulate. Sterile

leaves up to 1 .45 m long; petioles up to 45 cm
long; lamina up to 1 .0 m. 1-pinnate. but occasion-

ally becoming pinnate-pinnatifid to 2-pinnate in

transitional sterile-fertile leaves, subcoriaceous,

dark green adaxially, pale green abaxially. nearly

glabrous on both surfaces: pinnae up to 12 pairs,

mostly 12-20(30) X 3-6(8) cm, lanceolate, the

base round, cuneate or subtruncate, with the acro-

scopic side slightly prolonged, the margins entire,

crenate or serrate, the apex acuminate; veins in

pinnate groups 5-12 mm apart, the side branches

ascending and anastomosing at acute angles with

the adjacent ones (rarely, the veins fail to anas-

tomose locally), the vein tips connected by a faint

intramarginal connecting strand; axes nearly gla-

brous or with scattered hairs, these unicellular and

colorless. Fertile leaves 2-pinnate, 12-28 cm
broad; pinnules oblong to linear, 0.9-2(2.5) x 1-

2 mm; sori coenosoric, continuously covering the

abaxial surface of the pinnules; sporangial stalks

paraphysate; spores mostly (45)50-65(68) mi-

crons long. ;? = 41

.

Other illustrations: Ettingshausen, Famkr.

fig. 2. 1864 (as P. kunzei, portion of pinnae show-

ing vein); Hooker. Icones Plant. 1877. pi. 1690

(is Acrostichum polybotryoides) x\A p\. 1691 (as

A. juglandifolium): Rovirosa. Pterid. Sur. Mex.

pi. 60. 1909 (as A. coenopteris): Smith. Flora of

Chiapas, part II. fig. 71. 1981 (as P. aucuparia):

Stolz-c. Fieldiana Bot. II. fig. 60d. I98I (portion

of pinna with venation, as P. aucuparia).

Polyholrya polybotryoides grows in wet.

shaded, tropical forests from sea level to 1400

(1850)ni. It has the largest range of any species

in subgenus Soromanes, even occurring on Cocos

Island (Map 2). Gome? (1976) reported this

species from Nicaragua, but I have not seen a

specimen from that country .

The shape of the leaf apex varies clinally from

southern Mexico to Peru (Fig. 21 ). From southern

Mexico to Costa Rica, the apex is pinnatifid, hav-



November 1987 Monograph of Polybotra 47

FiciURE 20. Polybotrya polybotryoides (Baker) Christ, a. northern apex form. Guatemala; b. sterile

leaf with intermediate apex form, Panama; c. southern apex form, Ecuador; d. fertile leaf; c. sterile

pinna; f. stem scales, a: Sleyermark 37320 (F). b; Croal 12143 (MO), c.d; Moran 3569 (F). c,f; Moran

2I7H (CR).



48 Illinois Natural History Survey Vol. 34. Art. 1

ing two to four basal lobes; this apex does not

resemble the lateral pinnae. South of Costa Rica,

the apex resembles the lateral pinnae because it

has only one or two basal lobes with the remainder

of the apex relatively narrow and the margins en-

tire to crenatc. Since apex shape varies clinally

and no other characters correlate with it, I interpret

the two extreme shapes as geographic variations

of the same species and do not believe that these

should be named. Plants having an intermediate

apex shape do not appear to be hybrids, since they

do not have aborted spores. In South America,

the nearly conform apex of P. polybotryoides is

taxonomically useful because the three other

species of subgenus Soromanes , which might pos-

sibly be confused with this species, have strongly

and evenly pinnatifid apices. The subconform

apex in southern populations of P. polybotryoides

is derived within the genus, as evidenced by com-

parison to all other species of Polybotrya and to

all other genera of dryopteroid ferns.

Polybotrya polybotryoides is most like P. sub-

erecta. Another morphological character separat-

ing these two, besides apex shape, is the submar-

ginal connecting vein present in P. polybotryoides

(Fig. 20e)butonly partially formed in P. suberecta

(Fig. 22d). Unfortunately, the connecting vein is

faint and difficult to see because of the thick lamina

and, in some specimens, margins that have be-

come revolute upon drying and thereby hide the

region just inside the margin. The connecting

strand is best seen with magnification on the abax-

ial surface of the pinna or with the unaided eye

and the pinna held up to strong light.

Specimens examined: Mbxico Chiapas: Tum-

bala, Rtnirosa 972 (GH. PH); Finca Mexiquito, Piirpus

6761 (BM, F, GH, MO, NY, UC, US), 7245 (BM,

GH, US); 18-20 km N of Ocozocoautla de Espinoza,

along road to Mai Paso, Breedlove & Smith 21886

(MICH, NY, UC).
Bhlizk. Cayo: Hwy 28.5 mi S of Belmopan, Croat

24566 (CR. MO); Toledo district, Maya Mts , between

Rio Caraval and Union Camp, Boutin & ScMosser 5902

(MO).

Gliatkmala. Alta Verapaz: 7 mi up ihc road to

Oxec along road off Hwy 7F. bclwecn Tucuru and El

Estor ca. 6 km NE of Pan/os, Croat 4I6J7 (CR, MO).

Izabal: along Rio Bonita, Cerro San Gil, Steyermark

41698a (F, US); Cerro San Gil, Steyermark 41870 (F).

Quezaltenango: near Calahuache. Standley 67135 (F);

Finca Pirencos. below Santa Maria de Jesus, Standley

6820J (F). San Marcos: above Finca F.l Porvenir, Vol-

can Tajumuico, up I.onia Bandera Shac, Steyermark

37320 (F).

Honduras. Atlantida: Lancelilla Valley, near

Tela, Standley 53955 (F. US); Montana U .Manga, 30

km SE de La Ceiba, Nelson el al. 3296 (MO); Cordillera

Nombrc de Dios, Gome: 7027 (CR). Comayagua:

Quebrada Potrero, Cerro Azul dc Meambar, Cdme:69l4

fCR). Cortes: mountains on N side of Lake Yojoa,

Morton 7629 (US).

Costa rica. Cartago: Valley of the Rio Navarro.

1400 m. Werckle 16770 (P. photos F. NY. UC); along

Camino Rai? de Hule. SE of Plantanillo, Croat 36800

(CR. MO); Raiz de Hule. Moravia de Chirripo. Ocampo
727 (CR); Chitaria. forest near old jailhouse. Moran

2170. 2171. 2173. 2176. 2178 (CR. F. MO. NY):

Chitaria. Valeria 329 (US), 33133 (CR ); Finca Navarro.

Maxon 639 (NY). Cocos Island: Wafer Bay. Gomei

3324 (CR, F, US); Wafer Valley, Pittier 16232 (CR,

US); Wafer Bay river valley. Holdridge 5153 (GH).

Klawe 1545 (US); trail between Chatham and Wafer

bays. Gome: 18064 (CR); Chalham Bay. Jimene: M.

3209. 3210 (CR. F. GH. MICH); Chatham Bay. Four-

nier 357 (NY). Puntarenas: Osa Peninsula, on ridge

9,.S km W of Rincon de Osa. Mickel 2742 (NY); above

San Vito at Finca Wilson. NE of home. Evans & Bowers

3152 (MO); vicinity of biological field station at Finca

Wilson, 5 km S of San Vito de Java. Mickel3l80 (NY).

San Jose: Carrillo. Brade 372 (UC).

Panama . Cocle: Continental divide on road to Coc-

lesito. Hammel 3503 (CR) Darien: El Llano-Carti

road. Churchill <4 de Nevers 4993 (MO); Serrania de

Pirre, along ascent of Serrania de Pirrc above Cana Gold

Mine between Rio Cana and Rio Escucha Ruido. Croat

37757 (MO). 37794 (MO) Panama: El Llano-Carti

road, 13.7 km N of Pan-American Highway. Folsom

3590 (MO); Cerro Campana. ca. 10 km SW of Capira,

trail to summit, Mori & Kallunki 3574 (MO. NY); I mi

upstream from Frizzel's Finca Indio. on slopes of Cerro

Jefe, Foster & Kennedy I8I4{V. MO); Cerro Jefe, Web-

ster et al. 16467 (UC); along road to Cerro Campana.

Croat 14687 (F. MO. NY); trail to Cerro Campana.

Kirkhride & Hayden 274 (MO. NY); Cen-o Campana.

FSU Field Station. Kennedy et al. 2074 (MO); cloud

forest on Cerro Campana above Su Lin Motel. Crixj/

14742 (MO); Cerro Campana. above Su Lin Motel.

Croat 4266 (MO); Cerro Campana. near FSU building.

Croat 121 14 (F. MO). 12143 (MO. US). 14786 (MO);

8-12 km N of El Llano, along new El Llano-Carti

road. Nee et al. 8768 (MO); Cerro Campana. lower

slopes abtne FSU cabin, Cr<xif227S9(MO);CcrroCam-

pana along trail to summit. Croat I7I6I (MO. US);

high point of ridges S of Ipeti. 5-6 hr walk from Chocii

village. Serrania dc Majc. Knapp et al. 4542 (MO);

along trail to top of Cerro Campana. SW slope from

road. Nee & Stockwell 1 1610 (MO); Cerro Campana.

Armond 298 (MO). Veraguas: 5 mi NW of Santa F^.

slope above Rio Primeai Brazo. below E.scuela Agricola

Alia Piedra. Croat 23244 (MO). Uesner 820 (MO);

Valley of Rio Dos Bocas along road between Escuela

.\gncola .Mto Picdra and Calovebora. 15 6 km NW of

Santa Fe. Croat 27648 (MO); 0.6 mi beyond Escuela

J



November 1987 Monograph of Poiybotra 49

Agricola Alto Piedra, Croat <S Folsom .U05I (MO);

vicinity of Escuela Agricuitura Alto Piedra near Santa

Fe along trail to top of Cerro Tute, Aniomo 4019 (MO);

ridge of Cordillera de Tute. trail to Cerro Tute. above

Escuela Agricola Alto de Piedra. just W of Santa Fe.

Knapp & Dressier 5461 (MO).

Colombia Antioquia: Kalbreyer J79S (K. color

slide at MO; photo GH. MO); vie. Planta Provedencia.

28 km SW of Zaragoza, Alverson et al. 342 (MO, US);

cerca de Porcesito en el valle de Rio Medellin, Hodge

67H0 (GH); Barbosa. Henri-Stanislas 1709 (US).

Choco: Trail to Miniqui'a E of Puerto Mutis. Leilinger

& de la Sola 39 (US); Ri'o Mutata ca. .^ km above

junction with Rfo El Valle. NW of Alto del Bucy. Lei-

linger <S de la Sola 190 (US) Norte de Santander:

Ocana, on trees in the forest, alt. 7(XX) ft., Kalbreyer

1254 (K. color slide at MO; photo GH, MO).

EcuAiX)R. Bolivar: Limon. cstrivaciones in-

fcriores de la Cordillera Occidental. Aeosia Soils 6358

(F). Napo: 7.1 km al norcste de Baeza, Cascada de San

Rafael. Moran 3593 (Q. QCA). Foster H5-I62 (UC);

camino Bacza-Tena. \5 km al nortc de Tena. Moran

3530 (Q. QCA). Morona-.Santiago: Cordillera de

Cutucu. western slopes, trail from Logrono to Yaupi,

Madison el al. 3356 (GH) Pastaza: 6 5 km W of Mera,

Moran 3590 (F, Q, QCA). Pichincha: Hotel Tinalandia,

bosque primario al lado norte del Rio Toachi, Moran

3562 (Q, QCA) Tungurahua: 25 km al este dc Banos,

detrasel pueblo Ri'o Negro, Moran 3569 iP. Q. QCA),

Pkru. Huanuco: SW slope of Rio Lulla Pichis

watershed, a.scent of Cerros del Sira. Dudley I3290D

(GH) Pasco: Prov. Oxapampa. Abra los Mellizos, 4-8

km from Enanas, Skog et al. 5036 (US).

Figure 21. Clinal vaiiation in apex shape of Polyholrya polyhoiryoides (Baker) Christ. Plants from

the northern portions of the range have deeply and evenly pinnatifid apices. Southward, this shape

gradually changes into a subconfonm apex . Intermediates arc found primarily in Costa Rica and Panama.



50 Iij iNois Naturai. History Survey Vol.34. Art. I

3. Polybofrya suberecta (Baker) C. Chr. (Fig.

22, Map 1).

Pohhdirya suherecui (Baker) C. Chr., Index

Filicum 506. 1906.

Acroslichum suherectum Baker. J Bot. 207

1881. Type; Colombia. Aniioquia: 4(XX)-

4500 ft., Kalbreyer 1877 (holotypc: K. color

slide at MO!: photo GH!, US!).

Acroslichum hacketiunum Sodiro, Anal.

Univ. Quito XII (78):21 . 1895. (Crypt. Vase.

Quit. 491. 1893.) Typf: Ecuador. Pichincha:

banks of the Rfo Pilaton, Sodiro s.n. (holo-

typc: Q!).

Stem 1-2 cm thick, heniiepiphytic; scales

dark brown, narrow, linear, 0.2-0. 4( 1 .0) x 8-15

mm, stiff to spreading. Sterile leaves up to 1 .6 m
long: petioles about Vi to Mi the length of the

lamina, scaly at base, becoming less so distally:

liimimi 1-pinnatc, mostly 0.5-1 .()( 1 .2) ni long,

lanceolate: pinnae oblong-lanceolate, (10)15-

21(25) X (3)4-5(6.5) cm, 6-13 free pairs, gla-

brous to densely pubescent beneath, the hairs un-

cinate, multicellular, reddish or clear, 0.1-0.6

mm long, the margins entire, or (more rarely)

shallowly and inconspicuously serrate, the base

cuneatc to broadly rounded, the apex acuminate:

veins anastomosing, the tips sporadically uniting

to form a discontinuous, submarginal connecting

strand: axes with a few narrow, scattered scales,

usually glabrous, rarely pubescent by uncinate

hairs, or rarely by short (less than 0.2 mm), sub-

ulate, clear hairs. Fertile leaves 2-pinnate to 2-pin-

nate-pmnatifid. 10-30 cm wide, pectinate: mod-

erately scaly, the scales appressed, Wnear; pinnules

linear, 25-55 x 1-3 mm; sori coenosoric. con-

tinuously covering the abaxial surface of the pin-

nule: sporanfiial stalks paraphysate; spores (43)

46-55(58) microns long.

Other illustrations: Hooker. Iconcs Plant. 17,

pi. 1692. \^11 [an Acroslichum).

Polybolrya suberecta grows in montane

forests from 600 to 1800 m elevation. Most of the

specimens are from the western slopes of the

Andes (Map I ) These locations correspond to my
field experience in Hcuad<ir. where this species

was common on the weslcrn side of the Andes

hul absent from the eastern side.

The range of this species overlaps with P

polyhotryoides. a similar species The two species

are most easily distinguished b\ the shape of the

leaf apex: P. polyhotryoides has a subciinform

apex in Colombia and ticuador. but P. suherecia

has an evenly tapering, pinnatifid apex. Other dis-

tinguishing characters of P. suherecia are the un-

cinate hairs that occur in some specimens (Fig.

22a) and the discontinuous, submarginal connect-

ing vein (Fig. 22d). Polybolrya serralifolia also

resembles P. suberecta but differs by its wide,

flaccid stem scales and a distribution that is re-

stricted to high altitude forests in Venezuela and

Trinidad (Map I). Polybolrya suberecta looks

very much like P. andina— a species with which

it grows in the western Andes of Ecuador. See P.

andina for comparison.

Specimens examined: Colombia. Antioquia:

4000-4500 ft . Kalbreyer 1877 (K. color slide at MO;
photo GH. US) Choco: hills above Rfo Capa and Rfo

Mumba. up river from LXoro. Juncosa /467( MO. COL);

NW side of Alio del Buey. Lellinger & de la Sola 24S

(COL. LPB. US); 0.3 km E of the Ciudad Bolfvar-

Quibdo road, across the suspension bridge at km 141.

Lellinger & de la Sola 894 (COL. US ) Cundinamarca:

Ojodc Agua. S sideof RfoGuavio. 32 km Eof Gachala.

Grant 10556 {\JS). Magdalena: below Valparaiso.//.//.

Smith 98J (F. GH. NY); "Cincinnati." lower slopes of

Ml San Lorenzo, near Sta. Marta, 1300 m. Seifriz 24

(US) Santander: along highway between Pamplona

and Bucaramanga, Munic Tona, Corregimento Cor-

cova, Vereda la Marina, Croat 56510 (MO, UC).

Ecliador Los Rios: Patricia Pilar. Dodson 7380

(COL. MO. US). Dodson el al. 8679 (MO. US).

Pichincha: banks of the Rfo Pilaton. Sodiro s.n. (Q);

2.5 km E of Comejo Astorga. Moran 3546.5 (F. GH.

Q. QCA); 3 km from El Paraiso. road El Paraiso-

Saguangal. Ollgaard el al. 37823 (AAU. Q. QCA); 3

km from La Armenia, road Gualea-La Armenia.

Ollgaard et al. 37859 (AAU); about 40 km WNW' of

Quito. 3 5 km N of Mindo. ,V/<>ra;i 3565 (F. GH. Q.

QCA); Tinalandia Resort. N side of Rfo Toachi. about

25 km E of Sto. Domingo. Moran 3561 (F. GH. Q.

QCA); ca. 2 km N of Mindo. Hacienda San Vicente.

Fo.sler 85-37 (UC).

Pfrl'. Junin: Pro\ Tamia. Chanchamayo. Esposlo

10928 (USM).

4. Polybolrya andina C Chr (Fig. 23. Map 2).

Polybolrya andina C. Chr.. Index Filicum 7.

1905. noin. nov. for Acroslichum insigne

Baker, mm Fee 1872-73. with same type.

.Acroslichum insigne Baker. J. Bot. 167.

1877. nom. illeg.. mm Fee 1872-73. Typf:

Ecuador. Pichincha: "Andes of Quito." So-

diro (holotypc: K, photo GH!, US!: isotypes:

AAU!, CW.. Q!. UC!).



November 1987 Monograph of Polybotra SI

rrrrf E
u

5 cm
FrouRF 22. Polyhotrya suherecta (Baker) C. Chr. a. uncinate hairs from abaxial leaf surface; b. stem
scales; c. sterile leaf; d. sterile pinna; e.f. fertile pinnae, a: Ollgaard ct al. 37H23 (AAU). b-d: Moran
3546.5 (F). e: Smith 983 (NY), f: Dodson 7380 (US).



52 Illinois Natural History Survey Vol.34, Art. 1

FiGURF. 23. Polyholryti iimUtui C. Chr. a. fertile pinna; b. sterile pinna; c. sterile leaf: d. multicellular

uncinate hairs from aba.\ial surface of the leaf; e. stem scales, a: Sodiros.n. (AAU). b-e; Moran3563 (F).



November 1987 Monograph of Polybotra 53

Stem 1-1.5 cm thick, hemiepiphytic; scales

mostly 8-13 mm long, dark castancous. shiny or

dull, opaque, margins denticulate. Sterile leaf up

to 1 .0 m long; petiole up to 30 cm long, Vi to as

long as the lamina, scaly at base, becoming less

so upwards; lamina mostly 60-80 x 30-50 cm,

deltatc or broadly ovate-lanceolate, subcoria-

ceous, glabrous above, below densely pubescent

with spreading, tawny, uncinate, 4-to-IO-cellcd

hairs, these 0.5-1.2 mm long; pinnae 17-33 x

5.7-10 cm, mostly less than 3.5 times as long as

broad, 4-7 pairs below the pinnatitld apex, mar-

gins entire, base cuneate-rounded, petiolulate,

gradually becoming sessile distally; veins anas-

tomosing, the tips uniting discontinuously; axes

pubescent, the hairs like those on the lamina. Fer-

tile leaves slightly smaller, 40-80 x 20-32 cm,

pubescent with hairs like those of the sterile

lamina; pinnae pectinate; pinnules proximally

lobulate, becoming entire apically, 3-6 mm wide;

sori coenosoric, covering most of the abaxial sur-

face of the pinnule; spore length unknown.

Sodiro (1897) noted that this species "grows

in the tropical and subtropical region up to 1200

m in the forests around Sto. Domingo de los Co-

lorados and in the Nanegal and Mindo valleys"

(translation mine). This species is very rare and

restricted in range and has been collected only

twice from the western cordillera of Ecuador west

of Quito (Map 2). I saw only three plants at the

Tinalandia site. Polyhotryu siiherecta and P. poly-

hotryoides also occurred at this locality and were

more abundant.

This species is most closely related to P. suh-

erecta. from which it differs by its longer and

wider pinnae, fewer (4-7) free pinnae pairs below

the apex, and spreading, multicellular, tawny hairs

on the abaxial surface (Fig. 23d). Pubescent plants

of P. suherecta occur in the region of Ecuador

where P. andina grows. Within this region, the

hairs of both species are uncinate, but in P . suh-

erecta they are much shorter (0.1-0.3 mm), 1-

or 2-celled, erect, and easily overlooked by the

naked eye (Fig. 22a). Only these two species of

Polyhotryu have uncinate hairs. Because the fertile

leaf of ^. andina. known only from Sodiro's type

collection, had very few spores, I could not make

an adequate measurement of spore length.

Specimens examined: Ecuador Pichincha:

Hotel Tinalandia, casi 25 km al cste dc Slo Domingo

lie los Colorados, bosque primario arriba la monlana al

lado norte del Rio Toachi, HKK) m, Mnrim .i563 (F. Q.

QCA); "Andes of Quito." Sodiro s.n. (K, photo GH,
US; AAU. GH, Q, UC).

POLYBOTRYA subgenus SORBIFOLIA
Moran, suhiien. now

Type spixiks: P. sorhifolia Kuhn, l.innaca

34:64. 1869.

Folia l-pinnata. pinnae lineares vel lanceo-

latae, 6-10 sexies vel decies longiores quam

lutiores. apice longiacuminato : venae sihi paral-

lelae pinnatae. 3-5 in quoque lurmae, liherae.

Stem terrestrial and short-creeping (2 spp.) or

hemiepiphytic and long-creeping (I sp.). Sterile

leaves 1-pinnate; pinnae lincarto lanceolate, 6-10

times as long as broad, apices long-acuminate;

veins in pinnate groups of 3-5, strongly ascend-

ing, free all the way to the margin. Fertile leaves

pectinate or more rarely, moniliform.

5. Polybotrya sorhifolia Kuhn (Fig. 24, Map 3).

Polyholrya sorhifolia Kuhn, Linnaca 36:64.

1869. Based on var. salicifolium Hooker and

with same type.

Acrostichum caenopteris Hooker var. salici-

folium Hooker, Species Filicum 5:257. 1864.

Typr: Brazil. Pcrnambuco; Scrrado Araripe,

Gardner 1901 (holotypc: K, photo GH!; frag-

ment NY!).

Polyholrya salicifolia Leilinger, Amcr. Fern

J. 62:54. 1972. Type: Colombia. Santander:

vicinity of Puerto Berrio, between Carare and

Magdalena River, 100-700 m, 8 June 1935,

Haui;ht 1757 (ho\otype: US!; isotypc: BM!).

Stem terrestrial, 1-2.5 cm thick; stem scales

shiny brown, linear, 0.4-1 x 10-20 mm. mem-
branaceous, spreading, the margins denticulate.

Sterile leaves up to 1.3 m; petiole 15-50 cm long;

lamina lanceolate to oblanceolate, 0.3-1.5 x

0.15-0.50 m, chartaccous, with 12-18 pinnae

pairs, the apex abruptly acuminate, its lowermost

lobes decurrcnt; pinnae lanceolate-acuminate,

(7.3)10-22(25.5) X (0.7)1-3(3.5) cm, the base

truncate on its acroscopic side, cuncate on the

basiscopic side, the margins entire, crenate, usu-

ally scirate at the apex, the abaxial surface gla-

brous or glandular, the glands scattered, round,

whitish to reddish, sessile glands; veins slender,

sharply ascending, long parallel, usually oblique

to the costa, the tips free; (uo with a few scattered,

apprcssed .scales. Fertile leaves lanceolate, 0.3-

1 .3 X 15-40 m, pinnatc-pinnalifid;/)m/i(;(' linear,

4-15x0.7-1.0 cm, monilltorm because of the

round to oblong pinnules, these 1 3 mm wide;



54 Illinois Natural Hlstory Survey Vol. 34. Art. I

sori coenosoric, completely covering the abaxial

surface of the pinnules; sporangial stalks paraphy-

sate; spores mostly 40-47 microns long.

Other illustrations: Murillo. Cat. Illustrado

de las Plantas de Cundinamarca, 2; 10.^. 1966 (as

P. scrrotifolici): Varcschi, Flora Vcncz.,

Helcchos, vol. 1, lab. 7.V 1968 (as P. serratifolia):

Brade, Bradea 1(9):62, fig. 3. 1971 (as P. ser-

ratifolia); Lellinger, Amer. Fern J. 62, figs. 3. 4.

10. 1972 (as P. salicifolia).

The distribution of P. sorbifolia is spotty but

extensive (Map 3). This species looks like P.frac-

liserialis; see that species for comparison. Unlike

all other species in the genus, both P. sorbifolia

and P. fractiserialis are terrestrial

—

never climb-

ing—and it is doubtful that they could climb be-

cause their stems are only short-creeping. Both

species grow in shaded, rocky habitats along

streambanks, from 0-1200 m. 1 collected P. sor-

bifolia in Costa Rica at the El Rodeo site. It is

common at this locality, growing on talus along

a stream in the bottom of a humid, limestone ravine

that is surrounded by dry uplands.

Unfortunately, the specific epithet "satici-

fiilia" cannot be used for this species even though

its pinnae bear a strong resemblance to leaves of

certain Salix species, most notably 5. nigra and

S. amygdaloides. This resemblance was noted by

Hooker (1864) and Lellinger (1972).

Specimens examined: CosrA Rica. San Jose:

Alajuelila. Alfaro S07.i (US); El Rodeo. Moran 3145

(CR. F. GH. MO. PORT). Knighl v n (US), Gome:

7122 (CR). Hunnewelt 16514 (MICH)

Vhnl/uki.a. Aragua: Parquc Nacional "H. Pit-

tier," Rancho Grande, Tscluuli 162 (VEN). Steyermark

elaL 95827(1 (US) Portuguese: Dtto Araure, FilaSan

Jose, al oeste de Sla, Lucia, Onega cS Grimann 2707

(PORT). Yaracuy: "La Enjalma" al sur de Chivacoa.

Vareschi & Pannier 26M (US. VEN).

Co(.()Mi)iA, Boyaca: vallc del Rio Cusiana, entre

Pajarilo y Gua/ul. Murill,> 1457 (COL). 1491 (COL).

Cundinamarca: entre Nilo y la qucbrada de Agua de

Diosilo. Miirillo el al. 2H9 (COL. US) Magdalena:

Santa Maria, near Jiracasaca. H.H Smilh 1052 (1-. GH.

L. MICH. MO, NY, PH, US). Meta: along Cono Rosa

Blanca, a small stream outside of Villavicencio,

Kirkbride 404 (MO. NY). Santander: vicinity of Puerto

Berrio, between Carare and Magdalena River. Haught

/7.57(BM. US).

Bra/m Goias: .Serra Dourada. 17 km (straight

line) S of Goias Velho, h km NE of Mossaniedes, An-

derson 10152 (NY). Para: Serra dos Carajas. Serra

Norte, near waterfall near AMAZ Exploration Camp.

Silvu et al. BG 526 (AAU. F. GH. MICH. NY. UC).

Pernambuco: Serra de Araripe. Gardner 1901 (NY.

plioio of K specimen at GH). Roraima: Indian trail

from Surucucu to Uaica, Maita Mts. Prance etal. 10466

(NY).

6. Polybotrya fractiserialis (Baker) J. Smilh

(Fig. 25. Map 3).

Polybotrya fractiserialis (Baker) J. Smilh.

Hist. Filicum 133. 1875.

Acrostichum fractiseriale Baker. Synopsis

Filicum 414. 1868. Type: Peru. San Mani'n;

"in sylvis monlis campana, terrestre."" Dec.

1855, Spruce 4337 (holotype: K. color slide

at MO!; photo GHI. USD.

Acrostichum plumbicaule Baker. Synopsis

Filicum 413. 1868. Type: Peru. San Martin:

Terapoto, Ad rupes secus rivularum. Aug.

1855, Spruce 4090 acaoiype. K. photo GH!.

photo and fragment P!. US!).

Polybotrya plumbicaulis (Baker) J. Smith.

Hist. Filicum 133. 1875.

Stem terrestrial. 1-1 .5 cm thick, short-creeping

with inlemodes 1-3 cm long, apex scaly, behind

the apex usually naked or only sparsely scaly;

scales dull brown, opaque, 0.3-0.9 x 8-17 mm,
ascending, margins entire to more rarely denticu-

late. Sterile leaves to 1.4 m tall; petiole about

equaling the lamina, stramineous to lead-gray;

lamina slightly reduced or broadest at the base,

chartaceous to subcoriaceous, apex with one or

two major basal lobes, merely crenate-lohulate

above; pinnae mostly 9-15 pairs, linear to oblong.

(1 1)15-25(28) X (2.1)2.5-4.0(4.5) cm, margins

serrate, especially towards the acuminate apex,

base rounded to cunealc. w ith the acroscopic mar-

gin usually more oblique; veins numerous, fine.

and in closely parallel pinnate groups, occasion-

ally with a simple vein springing directly from the

cosla; axes usually glabrous or with fine, subulate,

0.1-0.2 mm hairs Fertile leaves creel. 2-pinnatc.

commonly taller than the sterile, to 1.5 m long;

pinnae linear, mostly 7-12(17) x 5-10(15) cm;

pinnules round, oblong, or linear. 1-3 mm wide;

.son coenosoric. completely covering the abaxial

surface of the pinnule; sporangial stalks paraphy-

sale. often with a bulbous glandular cell al base

of the paraphysis; spores (48)52-56(60) microns

long.



November 1987 Monograph of Polybotra 55

Figure 24. Polyhoirya sorhifolia Kuhn. a. sterile leaf; b. stem; c. ferlllc leaf; d. sterile pinna; e. stem

scales, a-e: Moran 3145 (F).



56 Ii.i.iNois Natural History Survey Vol. 34. Art. 1

Fic-.URE 25. Polyhotrya fracliserialis (Baker) J. Smith, a. fertile leaf: b. apex of sterile leaf; c. sterile

leaf; d. stem scales; e. sterile pinna; f. stem, a: C. Schunke 666 (F). b.d.c: Moran 3536 (F). c: Gonggrijp

& Stohel 3300 (MO), f: Tn.on & Tryon 5221 (GH).



November 1987 Monograph of Polybotra 57

Other illustnitions: Brade. Bradca 1(9):62.

fig. 4. 1971 (as P. polyhotryoides).

This species, like P. sorhifoUa. is terrestrial

and grows in wooded talus slopes, usually along

rocky strcamsides. The altitudinal range of P. frac-

lixerialis is from 200-1500 m, but about 80 per-

cent of the collections come from the 400- 8(X) m
range. Although principally a forest species, 1 have

seen vigorous fertile plants growing along sunny

streamsides in Ecuador, an observation which

suggests that this species has more tolerance to

higher light conditions than its cogenerics.

The range of P. fracliserialis occupies two

discontinuous regions: I ) the Andes from Bolivia

to Ecuador, and 2) the Guianas (Map 3). A dis-

tance of 900 km separates the northernmost popu-

lation in Ecuador from the westernmost population

in British Guiana. All populations of this species

occur on the eastern side of the Andes. Their ab-

sence from the western side is probably due to the

extreme dryness that characterizes that side of the

Andes south of the equator.

Polyholrya fracliserialis and P. sorbifolia

look very much alike and can be easily confused.

Several characters, however, distinguish the two.

The easiest feature to recognize is the shape of

the medial pinnae; those of P. fracliserialis are

relatively shorter and broader than those of P.

sorbifolia (see key and descriptions); this stoutness

is accentuated just below the apex (Fig. 25). The

apex of P. sorbifolia is another difference; it has

more numerous, decurrent, narrower pinnae (Fig.

24). Usually, the veins of P. sorbifolia are at a

more oblique angle to the costa than those of P.

fracliserialis. Another difference, albeit more rec-

ondite, is found in the sporangial stalks of P.

fracliserialis. which have a lightbulb-shaped gland

at the base of the paraphysis (Fig. 1 If.g); P. sor-

bifolia lacks such a gland (Fig. I lb). In fact, no

other species of Polybolrya has a glandular cell

on the paraphysis or elsewhere on the sporangium.

Polyholrya sorbifolia, however, has round, sessile

glands on the lamina; such glands are lacking on

P. fracliserialis.

Spore size is yet another diffencc; P. frac-

liserialis has much larger spores than P. sorbifolia.

This difference suggests that P . fracliserialis may

be a p<ilyploid, but chromosome counts are still

wanting for the two species. Polyploidy has not

yet been documented cytologically in Polybolrya.

Polybolrya fracliserialis and P. plumhicaulis

were described at the same time, in the same pub-

lication, and were distinguished on the basis of

leaf size, number of pinnae pairs, and two versus

three or four veinlets in each pinnate group. These

characters, however, intergrade completely even

on leaves from a single plant. I chose the name

P. fracliserialis. rather than the simultaneously

published name P. plumhicaulis. because it was

used by Posthumus (1928) in his treatment of

Surinam ferns.

Specimens examined: Frknch Guiana. (Juy-

ane: Cayenne, 1847, Leprieur s.n. (P), Inini: Cocao,

La Comic Riv.. 1847, Leprieur s.n. (P); Tumuc Aumac,

Maul Itany, Homk s.n. (P); ca. 45 km SE of Saul,

Granville i674 (CAY, Z); region de Paul Isnard, Mon-

tagne Lucifer, Granville 5219 (CAY, Z); Saul, Monts

du Fumec. Granville 5121 (CAY. Z).

Surinam. 3 km S of Juliana Top. 12 km N of

Lucie Riv., Maguire et al. 54384 (MO, NY). Irwin el

at. 54669 (NY). 547K4 (F. MO. NY); Brownsberg.

Gonggrijp & Slahel S300 (MO. US)

Guyana Rssequibo: Southern Pakaraima Mts..

Kopinang Falls. Maguire et al. 46068-A (NY).

Ecuador. Napo: casi 10 km SE de Tena. .^ km
E de Puerto Nuevo por camino a Puerto Misahuallf.

Moran J5J6 (F. GH. MO. Q. QCA. US). Pastaza: Rio

Capihuari. tributary of Rfo Pastaza. 0llgaard et al.

35124 (AAU).

Phru. Amazonas: Prov. de Bagua. Valley of Ri'o

Maranon above Cascadas de Mayasi near Campamento

Sta. Montenegro (kms. 280-284 of Maranon rd.), Wur-

dack IH54 (US). Cuzco: Prov. Paueartambo, Vargas

1 12H0 (GH). Huanuco: Prov. Huanuco. Tingo Maria.

Tryon & Tryon 5221 (GH); Tingo Marfa. Allard 20661

(US). 22305 (GH. US). 22526 (US); Previsto. antes de

Aguaitia. Aguilar 947 (USM); Prov. Leonicio Prado.

Distrito Rupa Rupa. al este de Tingo Maria, cerca al

Cerro Quemado. J. Schunke 10173 (MO); hills E of

Tingo Mana. Croat 21 152 (MO) Junin: Prov Sallpo,

Pichanaki. Leon 226 (USM); Schunke Hacienda, above

Ramon. C Schunke A2I4 (US); Chanchamayo Valley.

C. Schunke 165 (F. US). 666 (F), 1396 (F). Loreto:

above Pongo de Manseriche, Mc.xia 6246a (UC. US);

Aguaytia. Croat 2I0I4 (MO). Madre de Dios: Prov.

Manu, town of Atalaya. 2-3 km W of village. Foster

7455 (F. MO. USM); Pantiacolla. serrani'a across Rio

Alto Madre de Dios from Shintuya. Gentry et al. 27365

(MO); prov. Manu. km 139 on road between Cabo dc

homas and Shintuya. Faster 4020 (F. USM); Parque

Nacional de Manu. Cocha Cashu Biological Station.

Fouer P-84-9/ (F); vallc de Marcapata. Herrera 1200

(US) Oxapampa: Prov Pasco. Iscoazin, Faster 7937

(I-, USM) San Martin: Mt Campana. Spruce 4337

(color slide of K specimen at MO; photo GH. US);

Tarapoto. Ad rapes secus rivularam. Aug 1855, Spruce

4090 (photo of K specimen at GH; photo and fragment

P. US); Prov de Lamas, beween Tarapoto and

Moyobamba. ea. 1 km NW of Tabalosas. Croat 5 1 154

(MO).



58 Illinois Natural History Survey Vol. 34. An. 1

Bolivia. La Paz: Cordillera Real, Rfo Chimate,

Tate 502 (NY), 513 (NY); Mapin, Williams 1188 (NY,

US); Mapiri, Buchiien 1015 (US), 2138 (US); Mapiri

region, San Carlos, Buchtien 13 (US); Mapiri,

Charopampa, Buchiien II (MICH, P). Prov. unknown:

Challana Riv., Cardenas 1249 (LIL, US).

7. Polybotrya crassirhizoma Lellinger (Fig.

26, Map 4).

Polybotrya crassirhizoma Lellinger, Amer.

Fern J. 62.49, figs. 1, 8. 1972. Type: Peru.

Loreto; Gramitanacocha, Rfo Mazan, alt.

100-125 m, J. Schunke 268 (holotype: US!;

isotypes: F!, GH!, NY!, UC!).

Polybotrya macbridei Lellinger, Amer. Fern

J. 62;51, figs. 2, 9. 1972. Type: Peru. Juni'n:

Hacienda Schunke, La Merced, Macbride

5602 (holotype: US!; isotype: F!).

Stem l-2cm thick, hemiepiphytic;5fa/ei 10-

25 X 0.2-1 . 1 mm, dull orange or reddish brown,

rarely shiny and dark brown, with a dark central

stripe and lighter borders, rarely concolorous,

margins denticulate to erose. Sterile leaves up to

1 .35 m; petiole Vi to 'A the length of the lamina,

scaly at base; lamina lanceolate, to 1 .0 x 0.44 m;

pm«ae 6-20 pairs, (11)15-20(24) x 2-4 cm, lan-

ceolate to oblong-lanceolate, the base unequal,

broadly rounded or truncate on the acroscopic side,

slightly excavate or tapering on the basiscopic

side, the margins crenate-dentate or more rarely

lobate-serrate, apex acuminate, often serrate; veins

free, often with a single veinlet springing from

the costa between the main pinnate groups; axes

with a few, scattered, narrow, appressed scales,

occasionally pubescent, the hairs short, less than

0. 1 mm, erect, colorless, subulate, unicellular.

Fertile leaves 2-pinnate, occasionally 2-pinnate-

pinnatifid in large individuals, 'A to 'A smaller

than the sterile leaves; axes similar to those of the

sterile leaves but with larger more numerous

scales; sori coenosoric, completely covering the

abaxial surface of the pinnules, occasionally dis-

crete and round at the pinnule base; sporant;ia

nonparaphysate; spores (48)52-56(61) microns

long. Ai = 41.

Other illustrations: See original descriptions

of P. crassirhizoma and P. macbridei. cited

above.

Polybotrya crassirhizoma grows primarily in

lowland forests of the weslem Amazon basin and

in prcmontane forests of the eastern Andes (Map

4) from 100 to 1500 m. The cluster of dots in

eastern Ecuador reflects recent collecting activity

in that area and this species' true abundance there.

I saw this species in nearly every forest that I

visited in the Amazon basin of Ecuador, and it

was one of the most frequent and abundant ferns.

This abundance suggests that P. crassirhizoma

occurs farther east in the Amazon basin than is

shown in Map 4. Apparently. P. crassirhizoma

flourishes on clayey soils and does not grow in

sterile, sandy soils.

I chose the name P. crassirhizoma over P.

macbridei because of the more widely distributed

type collection. Polybotrya crassirhizoma and P.

macbridei were said to differ by stem thickness,

scale luster and color, and the presence versus the

absence of lobing at the base of the sterile pinnae

(Lellinger 1972). These characters, however, in-

tergrade completely as shown by a series of speci-

mens collected near Coca in the Amazon basin of

Ecuador (Moran 3616). The most obvious charac-

ter that supposedly distinguishes P. macbridei is

the basally lobed pinnae, but this character repre-

sents the lobing that typically occurs in transitional

sterile-fertile leaves. This example illustrates how

part-fertile leaves can cause taxonomic confusion.

Accordingly, I placed P. macbridei in synonymy.

Specimens examined: Colombia .\mazonas:

Loreto- Yaco. Schultes & Black 46-266 |GH>

Ecuador Napo: 27 km SE of Coca, petroleum

well Auca 4, Moran 3616 (F. GH. NY. Q. (JCA. US);

20 km NE of Coca, 5 km N of Joya de las Sachas,

Moran 3615 (F, GH, NY. PORT. Q. (JCA, US); about

10 km SE of Tena. 3 km E of Puerto Nuevo on road to

Puerto Mishahuallf. Moran 3535 (F. GH. NY. Q. (JCA.

US); Bimbino, on the Rfo Pacuno, lOhrW of confluence

with Rfo Napo, Whilmore 752 (BMl; Rfo Cuyabeno.

about 2 km upstream from Puerto Bolfvar. Brandbyge

el al. 33684 (AAU); San Pablo de los Sccoyas.

Brandbyge & Asanza C 32881 (AAU); San Pablo de

los Sccoyas. Rfo Wai si aya. Brandbyge ei al. 32617

(AAU); San Pablo de los Secoyas. on path to

Shushufindi . Brandbyge el al. 32544 (AAU); .SO km NE
of Coca. Lugo S. 3293 (GH); Canton Putumayo. Rfo

Aguanco, town of Dureno, Plowman el al. 4025 (GH);

Anagua. Parquc Nacional Yasuni. SEF forest project

area. Ollgaard el al. 38S.t6 (AAU). 39084 (AAU) Pas-

taza: Curaray. Vallc dc la Mucrte, Holm-Sielsen el al.

22484 [AAV. Q. QC.\): basin otRio Pasta/a. Pacayacu-

Sarayacu region. Gill 47 (NY); village of Rio Chico. 8

km from Puyo. Shemluck 280 (F); Rfo Bobonaza, be-

tween oil exploration camp ChichiaMa and Dcstacamcnio

Caho Po70. Ollgaard el al. 34855 (AAU. Q. QCA);

Rio Bobonaza. hclween Cachilama and the outlet of Rio

Bulco. Ollgaard el al .U748 l..\.\V): Rio Pastaza. near

the Peruvian border, around Destacamento Ishpingo,



November 1987 Monograph of Poi.ybotra 59

Figure 26. Polyhotrya crassirhizoma Ullingcr. a. fertile leaf; b. sterile leaf; c. fertile pinna; d. sterile

pinna; e. stem scales, a: 0llgaard el al. 34H55 (AAU). b-e: Moran 3615 (F).



60 Illinois Natural History Survey Vol.34. Art. 1

0llgaard el al. 34970 (AAU); 2 km W of Yuralpa, S

border of Ri'o Napo. Holm-Nielsen & Jeppsen 9H7

(AAU).

Peru. Huanuco: Prov Pachilae, Ri'o Pozouso,

Foster 9284 (F, MO, USM). Junin: Chanchamayo Val-

ley, C. Schunke 157 (F), I5H (F), 661 (F), SI2 (F);

Schunke Hacienda, above San Ramon. KilUp & Smith

24605 (NY, US); E of Quimiri Bridge, near La Merced,

Killip & Smith 23876 (NY, US); Hacienda Schunke. La

Merced, Macbnde 5602 (F, US); La Merced, Chan-

chamayo, Soukup 1065 (F). Loreto: Prov. Maynas,

Quebrada Yanomono, Explorama tourist camp, Rio

Amazonas above mouth of Rio Napo, Transect 6, Gentry

et al. 27880 (MO), Moran 3640. 3641 (F, Q, QCA);

Mishuyacu, near Iquitos, Klug 1386 (NY, US);

Gamitanacocha, Rio Mazan, J Schunke 268 (F. GH.

NY, UC, US), 14285 (US); Veradcra de Mazan, Croat

20763 (MO); Rio Napo near Entrada de Isla Inayuga,

Croat 20551 (MO).

Bolivia. Cochabamba: Prov. Carrasco, conflu-

ence of Rio Leehe with Rio Isarsama, Beck 1635 (LPB).

Brazil. Acre: Cruzeiro do Sul, vicinity of Serra

da Moa, Prance et al. 12180 (NY).

8. Polybotrya espiritosantensis Brade (Fig. 27,

Map 21).

Polybotiya espiritosantensis Brade, Rodri-

guesia 10:28, tab. 3. 1948. Type: Brazil. Es-

pi'iito Santo: Municipality of Itaguagu. Jati-

boca, virgin forest, 700-8(X) m, A. C. Brade

18224 (holotype: RB!).

Stem 1-2 cm wide, hemicpiphytic; scales

mostly 10-15 x 1.0-2.5 cm, bright reddish brown,

spreading, membranous, the center often darkened

to varying degrees, margins highly erose-dcnticu-

late, occasionally fimbriate. Sterile leaves up to

1.2 m long; petiole to 30 cm long, about half as

long as the lamina, scaly at base; lamina ovate,

2-pinnate proximally, becoming 1 -pinnate dis-

tally, 60-90 x 50-70 cm, subcoriaceous, pale

green, nearly glabrous on both surfaces, the apex

subconform and pinnalike but with one or two

basal lobes; pinnae alternate, 6-8 pairs, the apex

conform, like the lateral pinnules, the upper pinnae

simple and resembling the pinnules of the lower;

pinnules anadromic throughout, 3-4 pairs, ellip-

tic, 10-13 X 2-3 cm, the margins entire, the apex

acuminate, the base cuneatc, the proximal pin-

nules with stalks 2-4 mm long, the distal pinnules

sessile: veins slender, none conspicuously thicker

than the others, long, parallel. 1-2 branched, the

tips free; axes with a few scattered, appressed,

narrow .scales, otherwise glabrous. Fertile leaves

2-pinnate, amphiacrostichoid; pinnules caudate,

entire, appearing cylindrical when malure: sporan-

gial stalks paraphysate; spores (52)54-60(62)

microns long.

Other illustrations: Brade's original descrip-

tion has an excellent photograph of the tyf)e:

Brade, Bradea 1:67. tab. 6. fig. 5. 1971 (stem

scales only).

Polybotrya espiritosantensis is endemic to the

state of Espirito Santo in southeastern Brazil (Map

21 ). 1 expect this species will eventually be found

in other parts of mountainous coastal Brazil.

Polybotrya espiritosantensis has the most distinc-

tive laminar cutting of any species in the genus.

No other species has the simple, entire pinnules

that evenly taper at the base and apex (Fig 27b).

Unlike all other species of Polybotrya that have

pinnatifid apices, the leaf and pinnae apices of

this species are conform or nearly so (Fig. 27a).

The numerous, close, long, parallel veins (Fig.

27b) are like those of the 1 -pinnate species P.

fractiserialis and P. sorbifolia and probably indi-

cate a close relationship.

Specimens examined: Brazil. Espirito San-

to: Santa Thereza, 900 m, Foster & Foster 854 (GH,

US); Municipality of Itagua^u. Jatiboca. virgin forest.

700-800 m. A. C. Brade 18224 (RB).

POLYBOTRYA subgenus POLYBOTRY.\

Type species: P. osmundacea Willd., Sp. PI.

ed. 4. 5:99. 1810.

Polvhotrya subgenus Eupolybotrya Fee, Mem.

Fam. Foug. (Hist. Acrost.) 2:16. 1845.

Acrostichum section Polybotrya Hooker,

Species Filicum 5:244. 1864.

Stem hemicpiphytic; sterile leaves 1-pinnate-

pinnatifid to 4-pinnate; veins free.

9. Polybotrya caudataKunze (Fig. 28. Map 5).

Polybotrya caudala Kunzc, Linnaea 9:23.

1834. T>pe: Peru. Huanuco: Pampayaco,

July 1829. Poeppig s.n. (B!).

Polypodium adianloides Aublet. Hist. PI.

Guiane Franijoisc 2:962. 1775. nomen itieg..

non Bumi. (1768). Type: Guyana. Aublet

s.n. (holotype: BM!, Morton photo 6626 at

B!. F!. GH!).



November 1987 Monograph of Polybotra 61

E
o

FiGURF, 27. Polyhotrya espiritosantensis Bradc. a. sterile leaf; b. sterile pinnule; c. stem scales d fertile
pinna, ad; Brade 18224 (RB).



62 Illinois Naturai History Survey Vol.34. Art. 1

Olfersia caudala (Kunze) Kunze, Linnaca

21:206. 1848.

Psomiocarpa caudala (Kunzc) Presl, Epim.

Bot. 162. 1849.

Acroslichum cuudulum (Kunze) Hooker,

Species Filicum 5:244. 1864. nomen illeg.

mm Hooker (1840).

Polyhotrya acuminata Kaulfuss var. v'dlosa

Christ, Prim. Fl. Costar. 3( 1 ):9. 1901 . Type:

Costa Rica. Limon: Llanuras de Santa Clara,

500 m, Biolky 10688 (lectotype: US!;

isotype: CR!).

Polyhotrya villosula Christ, Bull. Herb. Bois-

sier, II. 6:168. 1906. Type: Costa Rica. Santa

Clara, Las Delicias. 1897, 500 m, Pittier

10688 (P?).

Polyhotrya costaricensis Brade, Bradea 1:11,

tab. 1 , fig. 1 . 1969. Type: CcstaRica. Limon:

Hundrisser Ranch, Atlantic shore, Sept.

1909, Brade & Brade 374 (holotype: HB;

isotypes: NY!, UC!).

Stem 0.5-2.5 cm thick, hemiepiphytic; scales

linear-lanceolate to narrowly triangular, 8-20 x

1-2 mm, dull brown, opaque, entire to denticulate,

the base elevated, thickened, curved, and ap-

pressed. Sterile leaves up to 2 x 1 m; petioles

30-70 cm long, about 'A the length of the lam-

ina; lamina mostly 2-pinnatc-pinnatirid, rarely 3-

pinnate at the base, firm-chartaccous. glabrous to

pilose, the hairs 0.2-1.5 mm long, acicular. the

margins sparsely ciliate. the hairs minute, less

than 0.2 mm long; pinnae up to 20-45(60) x 7-

20(30) cm. subdcltate. acuminate, subcquilateral.

slightly more developed on the acroscopic side;

pinnules slightly prolonged acroscopically, the

ba.sc truncate to slightly cordate, catadromically

arranged in the medial pinnae; veins free, some-

times with a single vein springing from the costa

between the pinnate groups; axes nearly glabrous

or pubescent to various degrees, the hairs acicular,

whitish; grooves usually pubescent within, often

densely so at the junctures. Fertile leaves similar

in size to the sterile, mostly 2-pinnale. but with

some of the larger pinnules lobed at base; pinnae

caudate. 4-8(1 1) X 0.5-1. 5 cm, apparently sorif-

erous on both surfaces; spores (43)46-50(53) mi-

crons long. ;i = 41

.

Other illustrations: Fee, Mem. Fani. Foug

(Hist. Acrost), Atlas, tab. 34, 1845; Vareschi.

Fl. Vcncz., vol. 1. tab. 73. 1968; Brade. Bradea

1(9):63. fig. I; p. 67. fig. 6.; Croat. Fl. Barro

Colorado Is., figs. 28 & 29. 1978 (as P. villosula);

Stolze, Fieldiana Bot. n.s. 6. fig. 60a,b. 1981;

Tryon & Tryon. Ferns & Allied Plants, figs. 80.8,

80.9, 80.22, 80.23. 1982.

Polyhotrya caudata is one of the most widely

distributed species in the genus (Map 5). It grows

primarily in lowland forests from sea level to 800

m, but specimens from the Andes have occasion-

ally been collected as high as 1900 m. I found

this species at many of the sites where I collected

in Costa Rica, Ecuador, and Peru, but not in Ven-

ezuela. Polyhotrya caudata is one of only three

species in the genus that occur on an oceanic is-

land—Cocos Island, about 500 km (310 mi)

southwest of Costa Rica.

Polyhotrya villosula was said to differ from

P. caudata only by its long and villous pubes-

cence, but specimens are usually glabrous or fully

pubescent, with few intermediate forms. Since

pubescence does not correlate with any other

character and I found no difference in geographical

range between the two forms, I regard P. villosula

as a synonym of P. caudata.

The juvenile leaves in glabrous plants of this

species may be difficult to distinguish from P.

osmundacea. A good character to separate the two

species is the minutely and sparsely ciliate leaf

margin oi P . caudata (Fig. 28a); that off. osmun-

dacea is always glabrous (Fig. 47). The thick,

dull brown, curved, and appressed stem scales

(Fig. 28b, h) also help distinguish P. caudata from

many other Polyhotrya species.

Specimens examined: Mexico. Chiapas: easi-

em highlands near Guatemalan border. Breedlove 34101

(CAS).

Guatemala. Alta Verapaz: near Rio Icvolay.

near Hacienda Yaxcbnal. 5 mi NW ofCubilquitz. 250-

.^()0 m, Steyermark 44675 (F); CubilquiU. 350 m,

Tuercklu-im I4S (US). SUS (P. Z). 7SI2 (US). 8040B

(US), S04I (GH, NY). Izabal: between Bananera and

"La Prcsa" in Montana del Mico. Steyermark 3S229 (F),

.i.S27l (F). .<9/97 (F): Rio Chacon. 30 m. Johnson 1221

(US); between Los Amales and Izabal. Sierra del Mico,

Kellerman 7J54 (F. Wl); along Rio Fno. 65 m, Sleyer-

mark .<y«95 (F. US); 2.5 mi N of Rio Dulcc on gravel

road to Pclcn. Dunn & LeDoux 22005 (MO. NY): vi-

cinity EXMIBAL Camp 2 (La Gloria). NW of Lake

Izabal , 400-500 m. Jones & Facey 3246 (NY); vicinity

of Quirigua. Sitimlley 24195 (GH); vicinity of Puerto

Bamos. SlandUy 250S5 (GH. I'S)

Bhi 1/l F.I Cayo: Vallcntin. Lundcll 6416 (GH.

US) Stann Creek: along road and stream al Dr\ Creek.

near Dist of Cayo Border, CriHil 24541 (CR, MO):

Middlesex. Gentle 2947 (GH. MICH. US); Schipp 273

(BM, F, GH. MO. UC, US, Z).



November 1987 Monograph of Polybotra 63

Figure 28. Polyboirya caudata Kunze. a. leaf margin showing minute cilia; b. stem and basal pinna

(note apprcsscd scales— same scale as 0; c-f. acroscopic pinnules, pinnae apex toward the right m

all; g. fertile leaf; h. stem scales, abaxial view at left, side view in center, adaxial view at right.

a,c: Costa Rica, Scamman 7156 (GH). b; French Guiana, Cremers 7369 (CAY). d,g,h: Costa Rica,

Moran 2186 (F). e: Bolivia, Sleinhach 7499 (MO), f: Trinidad, Fendler 105 (GH).



M Illinois Natural History Survey Vol. 34. Art. 1

Honduras. Atlantida: Lancctilla Valley, near

Tela, Slandley 53983 (F. US); near Ceiba, 400 m. Dyer

A225 (US).

Nkaragu A. Chontales: 4 km al oeste de La Liber-

tad, El Comatillo, Gome: & Cittar 6731 (CR); vicinity

of La Libertad, .500-700 m, Slandley 9047 (F). Zclaya:

road to Mina Nueva Americana, Stevens d Krukoff

12666 (CR, MO); along new road from Ri'o Blanco to

Ri'o Copalar, Stevens & Krukoff 12056 (CR, MO); Caiio

Aamora on Rio Rama, Stevens & Krukoff HS27 (CR,

MO); base camp 3.6 km SECerro San Isidro, Ri'o Kama,

Ri'o Escondido, 0-65 m. Proctor et al. 27091 (F. NY,

VEN); Mosquito Coast, Schramm s.n. (US); Bluefields,

Niell 2610 (CR); area de la Bahi'a de Bluefields, Ri'o

Escondido, 0-30 m, Molina R 1907 (F. US); near

Bluefields. Danneherger s.n. (US); Braggman's Bluff,

Englesing 291 (F, US); Mina Nueva Americana, 11.3

km N of main road, Pipoly 5299 (CR, MO); 13 mi

above Kururia, Pipoly 3H24: Colonia Kururia, Pipoly

3880 (CR, MO), 3901 (CR, MO); Neptune Mining Co.,

Stevens & Krukoff 13005 (CR, MO); near junct. of road

to Alimidkanba with road between El Empalme and

Limbiaka, Stevens & Krukoff 12741 (CR, MO); Apz. 5

km de Rama sobre el Ri'o Rama, Gome: & Citlar 6409

(CR); El Salto along Rio Pis Pis, Pipoly 3542 (CR.

MO); 13 mi above Kururia on road to San Jeronimo.

Pipoly 3817 (CK, MO).

Costa Rica. Cartago: near Turrialba. slope of

the Rio Reventazon behind the Instituto Interamericano

de Ciencias Agricolas. 600 m. Mickel3368 (NY); Valley

of Rio Reventazon 9 km ENE of Turrialba near Pavones,

650m. Holm & litis 200(F. NY). Cocos Island: Barclay

2199 (BM); Wafer Bay, Gome: 3354 (CR. F, US, Z);

Comfr-/52,S(CR); Chatham Bay . Jimene:M . 3200{CR.

GH). Heredia: Holdridge's Finca La Selva, Rio Puerto

Viejo at Quebrada El Sura and Q. El Salto, Rossbach

3710 (GH), Scamman 7482 (GH), Scamman & Hold-

ridge 7998 (CR, GH); 2 km upstream from confluence

of Ri'o Puerto Viejo with Ri'o Sarapiqui. Finca "La

§elva," Burger & Stohe 5865 (CR, F, GH, US), Mickel

3570 (NY) IJmon: Llanuras de Santa Clara, Biolley

10688 (CR, US); Hundrisser Ranch, Atlantic slope,

Brade & Brade 374 (NY, UC); 7 km SW of Bnbn,

100-250 m, Gomez et al. 20423 (CR, MO. UC); N end

of Tortuguero Natl. Park. Burger & Anlonio 11263 (CR,

F, NY); near Guapiles. at bridge over Rio Guacimo, 50

m, Moran 2182. 2185. 2186 (CR, F, P, PORT); near

banana plantation and Pandora, near Rio Estrclla.

Rossbach 3624. 3625 (GH); La Lola, a cacao finca,

near Ri'o Madre de Dios, Scamman 7156 (GH); Finca

Monlccristo, on the Rio Rcventazi^n below Cairo, 25

m, Standlcy & Valerio 48622 (US). Puntarenas: Osa

Peninsula, Mickel 1944 (CR, NY); hills N of Palmar

Norte, along trail to Jalisco. Croat 35176 (MO); Osa,

40 km W of lA rt. 2. Gome: 19489 (CR, MO).
Panama Rocas del Torn: in l^iguna dc Chiriqui

and its neighborhood, Han 53 (US) Canal Zone: "Isth-

mo Panama," Hayes 8 (B. GH, Y. US); hills N of

Frijolcs, Slandley 27597 (MO, US); Cen-o Azul. Ty.um

2109 (MO); along hogback ridge S of Fuertes Cove.

Pearson Peninsula, Croat 8153 (MO); headwaters of the

Ri'o Chinilla, above Nuevo Limon, Maxon 6895 (US);

ravine near Frijoles, Killip 2929 (US); along riverbank

between Fort Sherman and Fort Lorenzo, Croat <S Porter

15436 (MO); hilly forest around the Agua Clara Reser-

voir, near Gatun, 20-30 m, Maxon 4641 (GH, NY,

US); Orange River Trail. Cornman 544 (\JC. US); Bano

Colorado Island, Gatun Lake, McLxon el al. 6817 (GH),

Seaverns 56 (F), Starry 91 (F, GH), Croat 5114 (MO).

6850 (MO), 7362 (MO), 8025 (MO), 9000 (MO, NY),

9004 (MO, UC), 9103 (COL, MO, UC), 10804 (MO),

/5256 (MO), 17367 {?. MO), Bailey & Bailey 541 (GH).

Chrysler 4836 (UC); 10 mi from main gate, near Rio

Frijolito, Croat 15081 (F, MO, US); Allison Armour

Trail, Welmore <« Woodwonh 130 (GH). Chiriqui:

above Rio San Felix near town of San Felix, ca. 13 mi

N of Rio San Felix bridge, 8(X)-1200 m, Croat 33452

(MO); forest behind Vivero forestal, 12 km N of Los

Planes de Homito, IRHE Fortuna Hydroelectric Project.

1200-1300 m, Knapp 4966 (MO) Colon: Santa Rita

Ridge, end of road from Transisthmian Highway, ca.

10 mi from road. Porter el al. 4763 (MO); Santa Rita

Ridge, 2.8 mi from Boyd -Roosevelt Hwy.. Croat <S

Porter 15342 (F, MO); forest along Portobelo-Nombre

de Dios road. 10 km W of Nombre de Dios. Knapp <S

Mallet 5402 (MO) Darien: vicinity of Cana, summit

of knoll above Cana, Stern et al. 523 (GHl Panama:

2.5 mi N of Goofy Lake on road to Cerro Azul, Croat

11544 (F. MO); upper Mamoni River, 150-400 m, Pil-

lier 4492 (US); Orange River Valley, E of Juan Diaz,

Killip 2544 (GH, US); along Rio Pirati. foothills of the

Serrania de Maji, Knapp & Mallet 5133 (MO); in high

ridges of the Serrania de Maji, S of the Choco village

of Ipeti, Knapp el al. 4J79 (MO); along Juan Diaz River,

4 mi above Juan Diaz, 0-75 m. Killip 2845 (US). San

Jose Island: Perlas archipelago. Gulf of Panama, 55 mi

SSE of Balboa. Johnston 452 (GH. US) Veraguas: Isia

de Coiba. Mende: 75 (MO).

Trinidad. Aripo road via Anma. Broadway 5717

(F. Z); without IcKality. Fendler 105 (GH. MICH. MO.
NY. P, PH, UC); without locality, Broadway 5358{^\y.

forest, Brazil. Britlon et at. 2144 (GH. NY); Mount

Tamana. Rriiion el al. 1935 (NY); Oropuche, local road

via Valencia, Broadway 9207 (MO, UC); without local-

ity, Crueger 139 (B); La Sierra, Maraval, Hombersley

s.n. (MO); Arima Ward, Guanapo River N'alley, ca. I

mi SE of La Lcja village. Jenny 3122 (BNI); Cumaca

Road, Fay 472 (BM); Bnckl'ield's Tea Plantation, 3 mi

5 of Forestry Rest House, Jermy 2137 (BM); V'alencia

Ward, near Valencia. Jermy 10826 (BM); Hollis Reser-

voir Road. Mickel 9472 (UC).

Frlnch GL'IAN,^. Saint Jean du Mcanini. Bcnoist

1271 (P); Central Guyana, l.epricur s.n. (P); Sommcl

Tabiilairc. 650-750 m. ca 45 km SE of Saul, Crcmers

6374 (CAY), 6523 (CAY. Z); Bassin du Haul Inipi. 7

km WSW du Pic Coudreau (Moms Bakra). Granville

4012 (CAY. Z); Haut Oyaopivk. W of Trois Sauts

Cnque Eulcupousing, rive gauche du Saut Cambrouse,



November 1987 Monograph of Polybotra 65

Granville I/4J. 1165 (CAY, NY. Z); S of Tampoc:

Saut Koutnakou Soula. Cremers 44f>0 (CAY. Z); Saul

Tampoc. Granville -fH-f I (CAY); Haule Riv. Mana: Saut

Grous Tigrc. Cremers 7552 (CAY, Zl; Haute Crique

Baboune. affluent de la Riv Mana, Cremers 7369

(CAY, Z); Haute Crique Waamahpann. au depart du

chemin indien, Granville 971 (CAY); Frontierc Guyanc-

Surinam, Tuma Humac, Granville 99 1 (CAY, Z); Crique

Gabaret. 25 km de rembouchurc. layon N-W, Oldeman

1933 (CAY, NY); Cnque "Roche Fende" (aftleunl de

la Comte) a 1 km environ de son embouchure, Granville

B.47II (CAY, Z); W of Saul on trail to Monts Galbao.

Boom & Mori 1856 (CAY, NY); Piste allant de Cilron

vers le Massif du Decou Decou, Billiel & Jailin I6S3

(CAY); Region de Paul Massff du Decou Decou, Crem-

ers 7951 (CAY, Z).

SuRtNAM. West Rivier, 2-5 km SW of Juliana

Top, 275-300 m, Imin elal. 54896 (NY); no locality,

Hostmann s.n. (NY); 45 km S of Paramaribo, new road

to Hanover, N of Zanderij, Tryon & Kramer 561 1 (GH,

MICH, NY); about 25 km S of Paramaribo, l.imieman

4570 (GH), Kramer 1954 (MO); 3 km S of Juliana Top,

12 km N of Lucie River, 300-325 m. /mm 55/6/ (MO,

NY); Para Dist . Limleman & Teimissen 15291 (Z);

Suriname River. Plantage "Accaribo." d'An^renwnd

s.n. (Z).

Guyana. Kamuni Ck.. Groete Ck., Essequibo

River, Maguire <4 Fan.thawe 22855 (GH, NY); Es-

sequibo River. Persatid 372 (F); Rockstone. Gleason

582 (GH. NY); Demerara. Jenman s.n. (NY); Barima

River. Jenman s.n. (NY); 3 mi S of Chodikas, Guppy

462 (BM); Essequibo River, Moraballi Ck., near Bar-

tica, Richards 803 (BM); lower Cuyuni River, Sandwith

1561 (BM).

Venezukla. Bolivar: Dtto. Heres, Campamenio

Gun, Fernandez 951 (PORT, UC); Roraima, Schom-

hurgks.n. (B, NY); without locality, anno l843,Sf/iom-

burgk 1659 (B); F of Cerro El Picacho, N of l.as Nieves

and Las Chicharras, 45 km N of Tumeremo, vicinity of

Beborah. Altiplanicic de Nuria, 6(X)- 650 m, S/cvcrmorA:

89120 (NY, VEN); a lo largo de la Quebrada Acarabisi,

limiles del Eslado Bolivar con la zona dereclamacion,

Aymard el al. 952 (PORT, UC) Portuguesa: Depio

Paez, Pozo Blanco, enire Acarigua y Payara, 190 m,

Onega 636 {PORT. UC, VEN). Sucre: selvadcl Rcctan-

gulo N del Lago dc Tuanoco, Lasser & Vareschi 3926

(VEN); vicinity of Cristobal Colon, Avicagua, Broad-

way 560 (GH. NY, US) Territorio Federal Delta

Amacuro: Depto. Antonio Diaz, 9°I5'N, 6()"57'W,

upper reaches of riverine forest of Cano Atoiba tributary

of Boca Araguao. Sleyermark el al. 115033 (VEN);

Depto Antonio Diaz, vicinity of Caho Jotajana ( = ticrra

alta). tributary of Caiio Guiniquina. NW of Epana near

boundary with Depto. Tucupita, 9°I5'N. ai'lO'W,

50 m. Sleyermark elal. 1 15021 (MO. UC. VEN); Depto.

Pcdemalcs (boundary with Depto Tucupita): Caiio

Simoina. west of Isia Coucuina. S of Barra de Coucuina.

.50 m, Sleyermark el al. II433I (MO, VEN). I I43.Q

(MO, UC, VEN).

Colombia. Amazonas: Trapecio Amazonico,

Loretoyacu River, 100 m, Schulles <$ Black 8467 (GH,

US). Cauca: Cali, Lehmann 2998 (BM); en la orilla

opuesla Puerto Lim6n-Rio Caqueta, Mora 4430 (COL);

Gorgona Island, off Nariiio. Taylor 1223 (MICH, NY);

Rio Timbiqui, B.T. 443 (GH). Choc6: slopes and ridge

of Loma del Cuchillo. ca. 15 km WSW of Chigorod6,

150-400 m. Leilinger & de la Sola 643 (LLP. US);

upper Rio Truando. 2 km SSW of the confluence of

Rio Nercua near the MADUREX Camp. 100 m. Ul-

linger & de la Sola 589 (COL, LP, US); trail to Miniquia

E of Puerto Mulis (Bahia Solano), 20-1 20 m. Leilinger

& de la Sola 26 (CR. COL. LP. US); Rio San Juan,

3.5 km SW of Andagoya. just NE of the mouth of the

Rio Suruco. 60 m. Leilinger & de la Sola 496 (COL,

LP); Municipio de Rio Sucio, Parque Nacional Los

Catios. alrededor del Campamento de Tilupo. 250-320

m. Forero & Jaramillo 1745 (COL, MO). Cun-

dinamarca: Cordillera Oriental, MesaNegra.Gazuguan

Valley. 6 km NW of Medina. 580 m. Grant 10436 (F,

US). El Valle: La Cumbre, Cordillera Occidental,

1700-2200 m, Killip 11343 (GH, NY, US); Finca La

Pradera ca. 6 km SW of El Cairo on trail to Rio Blanco,

between EI Brillanle and Boquer6n, Leilinger & de la

Sola 790 (COL, US). Narino: Pambana, between Rio

Pimbi and Rio Cuembi, on Rio Telembi, above Bar-

bacoas, 50 m, Ewan 16846 (BM, GH, UC).

EcuAtxjR. Los Rios: Rio Palenque biological sta-

tion, km 56 Quevedo-Sto. Domingo road, Evoy 104

(NY). Morona-Santiago: Pachicutza, at "Escuela is-

comisional [sic] Cardinal Dofner," km 140 on road

Loja Gualaquiza, 900- 1 000 m, Holm-Nielsen el al. 4489

(AAU, Q, QCA), 4495 (AAU, Q, (JCA); Sucua,

Swingle el al. 70-02-05-2 (UC, US). Napo: ca.si 10 km
SE de Tena, 3 km este de Puerto Nuevo por camino a

Puerto Misahualli, 300 m, Moran 3534 (F, Q, QCA);

10 km al sur de Tena a Puerto Misahualli, Moran 3588

(Q, QCA); Anangu, Parque Nacional Yasuni, SEE proj-

ect, 260-350 m, 0llgaard el al . 38932 (AAU, Q, QCA);

27 km SE of CiKa, Moran 3617 (MO, C, QCA). Pas-

taza: Rio Bufeo, northern tributary of Rio Bobonaza,

300 m, Ollgaard el al. 43798 (AAU, Q, QCA); Rio

Bobonaza, around houses between Huagraeachi and

Cachitama, below Montalvo, 300 m, Ollgaard el al.

34634 (AAU, Q, QCA); oil exploration camp Chi-

chirota, on the Rio Bobonaza, 300 m, 0llgaard el al.

35290 (AAU, Q, QCA) Sur de SanUnder: vicinity of

Barran ca Bermeja, Haughl 1325 (MO)

PlRU. Ama/.onas: Prov dc Bagua, left bank of

Rio Maraiion opposite Quebrada Mirana (opposite km
277 of Maraiion road above Cascadas de Mayasi), 425-

450 m, Wurdack 2011 (GH, US, USM) Cuzco: Prov,

La Convcnciiin, 73°40'W, 12°30'S, at Camp Zx;ro,

l\Om. Dudley //5/.? (GH, US); Camp I, 910-940 m,

in J. Knox's quadrate, Dudley 10168 (GH); Prov.

Paucariambo, Mautainiza (?], 80()-9(X) m. Vargas

17800 (GH) Hu^nuco: Tingo Maria, 615-1100 m,

Allard 22593 (US); Cuchcro, Poeppig s.n. (BM); hills



66 Illinois Natural History Survey Vol.34. An. 1

E of Tingo Mari'a, Croat 21191 (MO); 10 km S of Tingo

Maria, Slork & Norton 9509 (F. UC. US); Pampayaco,

Poeppig 201 (B). Loreto: Prov. Mayna,s. Peler Jensen's

Explorama Lodge, 50 mi downriver from Iquitos at

Yanamono Ck., Moran 3663 (AMAZ, USM); Prov.

Maynas. about 10 km SW of Iquitos at zoological park,

Moran 3671 (AMAZ, USM); Pebas on the Amazon

River, Williams 1739 (F); Caballo-Cocha on the Ama-

zon River, Williams 2137 (F); Ri'o Mazan, 100-125 m,

C. Schunke 380 (GH, NY); Prov. Maynas, Rio Itaya,

10 km S of Iquitos, Tryon (S Tryon 5204 (GH); lower

Rio Huallaga, 155-210 m, Williams 3999 (F, US);

Quebrada Nawampa, Croat 17620 (MO); Puerto Arturo,

lower Rio Huallaga below Yurimaguas, 135 m, Killip

& Smith 30690 (NY, US); Mishuyacu, near Iquitos, 100

m, Klug 1166 (F, NY, US); Quebrada Tahuayo above

Tamishiyaco, Croat 197/7 (MO); Rio Itaya above

Iquitos, Croat 19220 (MO); pnmary forest 17 km SW
of Iquitos on road to Puerto Almendara, Croat 18388

(MO); 1 2 km SW of Iquitos, Croat 18219 (MO). Madre

de Dios: 12°49'S, 69°17' W, Prov. Tambopala, Tam-

bopata Nature Reserve, ca. 30 air or 70-80 river km

SSW of Puerto Maldonado at effluence Rio Torre/Rio

Tambopata, 260 m, Barbour 4764. 4790 (F, MO); Par-

que Nacional de Manu, Cocha Casha Biological Station,

Foster P-84-42 (F). Pasco: Puerto Bermudez, 375 m,

Killip & Smith 26637 (NY, US). Puno: San Gavan,

Lechler 2.?2 /, 2i29 (B ). San Martin: San Roque , 1 350

-

1500 m, Williams 7681 (F, US); Cerro de Campana,

Spruce 4634 (P).

Bolivia. La Paz: Prov. Sud Yungas, Limite de

los Deptos. La Paz/Beni, Ri'o Quiquibey, 320 m. Beck

8037 (LPB); San Carlos, Mapiri, Mapiri River, Tate

422 (LPB, NY); Mapiri, Buchtien 35 (B), 290 (NY,

UC). Santa Cruz: Prov. Sara, Ri'o Yapaicani, 400 m,

Steinbach 7499 (B, F, GH, MO, Z); bosque del Rio

Surutii, 400 m, Steinbach 3032 (US).

Brazil. Acre: Cruziero do Sul vicinity, Rio Moa

between Igarape Ipiranga and Aquidaba. Prance et al.

12069 (F, LP, NY, UC). Amazonas: prope Sao Gabriel

da Cachoeira, ad Rio Negro, Spruce 21 16 (B. BM, P.

RB); Larges, on the Amazon River 1 km below mouth

of Rio Negro, Conant 940 (F, GH, NY). Para: Serra

dos Carajas, AMAZ camp AZUL, Sperling 5914 (GH,

NY); Bclem, Huber 2599 (BM), 7235 (BM).

10. Polybotrya goyazensis Brade (Fig. 29, Map 6).

Polybottya goyazensis Brade, Bradea 1:24.

tab. 1, fig. 1. 1969. Type: Brazil. Goias:

Goiania, piimai^ forest, December 1936.

A. C Brade 15373 (holotype: RB!; isotype:

NY!).

Polybotrya macedoi Brade, Bradea 1:24.

1969. TvpF,: Brazil. Goias: IVlunicipio Jatai.

Localidad Balsans. 1 November 1950,

Macedo 2682 (holotype: HB; isotypes: HB,

MO!. SP. US!; paratypcs; Brazil. Goias:

Fazenda Queixado, 8 December 1948,

Macedo 1447 (RB. SP); Fazenda Balsamo.

15 December 1948, Macedo 1521 (SP).

Stem 1-2.5 cm thick, hemiepiphytic; scales

spreading, membranaceous, generally 6-12(16)

mm long, dark castaneous or dark reddish with

lighter borders, margins strongly denticulate to

erose, the base cordate to various degrees, often

black and sclerified at the point of attachment.

Sterile leaves up to 1 .45 m long; petiole 'A to V2

as long as the lamina; lamina mostly 2-pinnate-

pinnatifid, lanceolate to ovate, 1.2 x 0.9 m, the

abaxial surface nearly glabrous to densely pubes-

cent, the hairs whitish, acicular, up to 1 mm long,

glands occasionally present, these reddish, sessile,

resinous; pinnae up to 45 x 23 cm, lanceolate to

narrowly deltate, the suprabasal ones soon becom-

ing pinnatifid; pinnules narrowly deltate, mostly

falcate, catadromically arranged above the base.

4-8( 10) X 1-2(3) cm, the acroscopic side slightly

prolonged, the margins entire to crenate. ciliate,

the hairs minute, 4-10 celled; axes nearly glabrous

abaxially or densely pubescent, by tiny, unicellu-

lar hairs, scales very small or lacking, adaxially

pubescent in the central groove with reddish hairs,

0.3-0.8 mm long; grooves usually ciliate on the

ridges. Fertile leaves 2-pinnate, amphiacrosti-

choid: pinnules caudate, often lobed or undulate

at the base; sporangial stalks paraphy sate; spores

(44)48-62(70) microns long.

Other illustrations: See original description

of P. goyazensis: Sehnem, Fl. llust. Catarinense,

pi. 18, 1979.

Polybotrya goyazensis occurs in Paraguay

—

the only species of Polybotrya known from that

country— and in the southern half of Brazil but

not in the coastal mountains (Map 6). This range

is unusual because it comprises regions apparently

not occupied by other species of Polybotrya. I

suspect that P. govazcnsis evolved from south-

ernmost, isolated populations of P. caudata.

Polybotrya goyazensis has stem scales that

are spreading, shiny, membranaceous, translu-

cent, darkened in the center with lighter borders,

margins strongly denticulate to crose, and the base

cordate around the darkened p<iint of attachment

(Fig. 29f). In contrast, P. caudata has stem scales

that are cur\'ed, appressed. thick, dull brown, con-

colorous. margins entire, and attached across the

length of the thickened base (Fig 28h). The mi-

nute cilia on the margins of the lamina ser^e to

distinguish these two species from almost all other

species of Polybotrya (Fig. 28a).



November 1987 Monograph of Polybotra 67

Fiourf; 29. Polyholrya goyazensis Bradc. a. fertile leaf; b. sterile leaf; c-c. proximal acroscopic
pinnules; f. stem scales. a,b: Rojas 10431 (B). c; Brade 15373 (RB). d: Frame & Selva 59677 (NY)
e.f; Argent el al. 6336 (UC).



68 Illinois Natural History Survey Vol.34. Art. I

Brade (1969c) distinguished P. macedoi from

P. goyazensis by the former's pilose hairs on the

axes and its slightly darker scales with lighter bor-

ders and a longer acuminate tip. These differences

are not constant. Various degrees of pubescence

exist, from densely pilose to nearly glabrous. This

intergradation is shown by the type ai P. goyazen-

sis: the holotype at RB is nearly glabrous but the

isotype at NY is densely hirsute. Examination of

the stem scales shows a complete gradation of

color, sometimes even on a single specimen.

Specimens examined: Brazil. Goias: Goiania.

primary forest. December 1936, A. C. Brade 15373

(RB, NY); Serra do Caiapo, 40 km S of Caiaponia,

Prance & Silva 59677 (NY); Municipio Jataf. Locaiidad

Balsans. Mucedo 26S2 (MO, US). Federal District:

"Brasilia." Glaziou 14456 (B. P). Mate Grosso: 270

km N of Xavantina. 12°54'S, 51°52'W. Ratter et al.

2047 (MO); Municipio Bnlhante. Rio Anhanduf.

Hatschbach 251 18 (UC); Expedition Base Camp. Brejo,

12°49'S, 5I°46'W, Argent & Richards 6651 (NY);

49.5 km N along road from base camp, Harley et al.

10922 (NY); Xavantina, ca. 4 km N of ferry. Argent et

al. 6336 (RB. UC) Para: no locality. 1914. Petelot

.v.n. (F).

PARACiUAY. Southern Paraguay. Sierra de Amam-
bay. Rojas 10182 (B. BM). 10431 (B. L. LIL, M,

MICH. P). IOH04 (B. BM. L); no locality. Hassler

y/6/«(MICH).

11. PolybotryapubensMartius(Fig. 30,Map6).

Polybotrya puhens Martius, Icon. Plant.

Crypt. Brasil. 87, tab. 25. 1834. Type:

Brazil, ad flum. Amazonas prope Serpa, Sep-

tember, Martius s.n. (holotype: M).

Polybotrya pubens Kunze, Linnaea 9:22.

1834. non Martius. Type: Peru. Loreto: Prov.

Maynas, ad Yurimaguas, September 1830,

Poeppig s.n. (lectotype: P!).

Acrostichum pubens (Kunze) Hooker &
Baker, Synopsis Filicum 414. 1868.

Acrostichum caudatuin (Martius) Hooker var.

pubens (Martius) Baker, Fl. Brasil. 1(2):586.

tab. 38, figs. 1,2. 1870.

Dryopteris guenlheri Rosenstock. Fcddes

Repert. 25:59. 1928. Type: Bolivia. La Paz:

region of Mapiri, San Carlos, 15 November

1926, Buchticn 260 (holotype: UC!).

Polybotrya crespiana Bosco. Nouvo Giom.

Bet. Ital., 11. 45:145, tab. 9, fig. 1. 1938.

Type: Fcuador. Santiago-Zamora: Indanza,

Crespi s.n. (type not found, known only from

Bosco's fig. 1).

Polybotrya decorata Lellinger, Amer. Fern

J. 62:54, figs. 5, 6, 11. 1972. Type: Peru.

VxiTtXo: Iquitos, alt. ca. 1(X) m, Killip & Smith

26955 (holotype: US!; isotype: NYl).

Stem 1 .0-1 .5 cm thick, hemiepiphytic; scales

of large mature stems mostly 10-15 x 2-5 mm,
flaccid, membranaceous, concolorous, whitish to

coffee brown or castaneous, spreading or more

commonly matted, margins fimbriate, young or

terrestrial stems with narrowly lanceolate, castane-

ous, denticulate scales. Sterile leaves up to 1.2 m
long; petiole up to 21 cm long, about 'A to 'A the

length of the lamina; lamina to 1 m long, narrowly

deltate to elliptic, tapered evenly to the apex,

mostly 1-pinnate-pinnatifid, the base occasionally

2-pinnate-pinnatifid, pubescent throughout or

rarely glabrous adaxially, rarely with punctate,

reddish glands; pinnae narrowly triangular, up to

9-19 X 2.5-3.5 cm, serrate to pinnatifid, often

with only the lowest pinnae having a few free

proximal segments; pinnules or segments cata-

dromic, slightly falcate, oblong, the basal ones

slightly longer than the suprabasal and more

deeply cut; veins free, 6-9 per segment; axes con-

spicuously pubescent, the hairs 1-2 mm long, un-

branched, acicular, whitish, 6-15 celled, the sep-

tae often reddish, polyseriate scales lacking, but

scales present as reduced, appressed, reddish, flat-

tened, uniseriate "hairs." Fertile leaves 2-pinnate,

to 0.8 m long, amphiacrostichoid; axes pubescent

on both surfaces, the hairs ca. 0.1 mm long, col-

orless, subulate; proximal pinnae deeply lobed or

fully pinnate, long-caudate; sporangia with the

capsule walls setose at the apex near the annulus

by two hairs, these 1 -3 celled; sporangial stalks

paraphysate; spores (44)48 - 57(63) microns long.

Other illustrations: Martius. Icones Plant.

Crypt. Brasil. 87. tab. 25. 1834; von Ettings-

hausen. Denkschr. Ak. Wien., tab. 7, figs. 3, 10,

11. 1864; Martius, Fl. Brazil, U2):586. tab. 38,

figs. 1, 2. 1870; Brade, Bradca 1(9): 63, fig. 4;

Ibid. 67. fig. 7. 1971.

Polvhotrya puhens grows primarily in low-

land forests of the Amazon basin but occurs in

premontanc forests up to 1400 m in the Andes

(Map 6). During my fieldwork in Ecuador. 1 found

this species in well-drained forest soils that were

slightly elevated above the surrounding swamp

forest.

This is one of the most distinct species of

Polybotrya. readily distinguished by its combina-

tion of stem scales, lamina shape and cutting.



November 1987 Monograph of Polybotra 69

FiGURF 30. Polyhotrya puhens Martius. a. pinna of a large, highly divided leaf: b. pinna of typically

cut leaf: c. sterile leaf; d. fertile leaf; e. setose sporangium with paraphysis on stalk; f. setose sporangium;
g. cross section of fertile pinna apex; h. stem scales; i. hairs from sterile lamina, a; Killip & Smith
26955 (US), b: Killip & Smith 26543 (NY), c.h.i: hahlev 47fi7 (CR). d.g: Killip & Smith 26194 (NY),
e.f: Holm-Nielsen & Jeppsen 663 (AAU).



70 Illinois Natural History Survry Vol.34, Art. I

pubescence, and long-caudate fertile pinnae. In

addition, P. pubens has, unlike any other species

of Polybotrya, setose sporangial capsule walls

(Fig. 30e,0. The sterile lamina is usually 1-

pinnate-pinnatifid or almost 2-pinnate at the base,

but large leaves that are 2-pinnate-pinnatifid for

much of their length are occasionally collected

(Fig. 30a). These large, more highly divided

leaves appear quite distinct but there are inter-

mediate forms between the extremes of lamina

cutting. The 2-pinnate-pinnatifid form was named

as P. decorata.

Polybotrya pubens and P. glandulosa might

be confused because both have similar laminar

cutting, long, acicular hairs, and both grow in

lowland forests of the Amazon basin. But the

lamina of P. glandulosa is, unlike any other

species of Polybotrya, cuneatc and the petiole is

extremely short, only 2.5 cm long. Furthermore,

its fertile leaf is botryoid instead of amphiacros-

tichoid, and its sporangial capsules are glabrous

instead of setose.

Juvenile stems and thin terrestrial stems have

narrow, castaneous, spreading scales that differ

from those on the scandent stems. Dryopteris

guentheri represents a juvenile form of P. pubens

with this scale type. The transition from juvenile

to mature plants and the changes in scale and

lamina morphology are shown in an excellent

series of specimens collected in Ecuador by

Balslcv (#4787).

The pubescence of the fertile leaf consists of

colorless, subulate hairs about 0.1 mm long—
these are much reduced in comparison to the hairs

on the sterile leaves. The setae on the sporangial

capsule walls are unique in Polybotrya. They

occur only at the apex of the sporangium near the

annulus (Fig. 30e,f); it is odd that these setae are

not shown on the sporangia illustrated in 1834 by

Martius in his otherwise excellent plate of P.

pubens. The adaptive value (if any) of the hairs

on the capsules is not known.

Unlike all other species of Polybotrya. P.

pubens lacks wide, polyseriate scales on the axes.

Rather, scales are present only in ihcir reduced

form, that is, as tiny, apprcssed. reddish, jointed,

uniseriate "hairs."

Both Martius and Kun^c published the name

P. pubens in 1834. Kun/e's work appeared in the

first issue oi l.innaea. which was published 22-28

June 1834 (Statlcu l%7). Slatlcu and Cowan

(1981) report that Martius's second fascicle of

"Icones plantarum crytogamicarum" was pub-

lished in 1834. Since no month or day is given.

1 cannot determine which work was published

first. I consider Martius the author of P. pubens

because Kunze cleariy attributed this name to Mar-

tius.

Specimens examined: Colombia Amazonas:

Ri'o Apaporis, Soratama (above mouth Rio Kananaril

and vicinity, Schulles & Cabrera 16053 (US).

Ecuaoor Morona-Santiago: Taisha. 3-4 km

ESE of the militar> camp, Brandbyge & Asanza C.

}IH70 (AAU) Napo: Reseria Faunistica Cuyabeno. al

lado nortc de Laguna Grande. Balslev 4787 (AAU, CR.

Q, QCA). 4802 (AAU, Q, QCA). Laegaard 51136

(AAU.Q,QCA),5//50(AAU,Q.QCA),5/224(AAU,

Q, QCA); Parque Nacional Yasunf, in the area of the

SEP project. Ollgaard el al. 38969 (AAU). 39039

(AAU). 39040 (AAU); 6 km along Ri'o Pano. Hotm-

Nielsen & Jeppsen 663 (AAU). Pastaza: Rfo Bolxjnaza.

oil exploration camp Chichirota and Destacamento Cabo

Pozo, Ollgaard el at. 34894 (AAU. Q. (JCA); Curaray.

SE of the airstrip, Holm-Nielsen el al. 22121 (.AAU);

oil exploration camp Chichirota. on the Rio Bobonaza.

0llf;aard el at. 35350 (AAU); Ri'o Bobonaza. between

Cachitama and the outlet of Ri'o Bufeo. Otlgaard el at.

34703 (AAU). Santiago-Zamora: Taisha. Cazalet &
Pennington 7720 (BM. F. K. NY, US).

Peru. Amazonas: ridge crest of Quebrada Chuivi

(above km 278 of Mararion road), valley of Rio Maraiion

near Cascadas dc Mayasi. W'urdack 1933 (USi Junin:

E of Quimiri Bridge, near La Merced. Kittip <S Smith

23916 (NY. US); Pichis Trail, Santa Rosa. Killip <S

Smith 26194 (NY. US); Pichis Trail. San Nicolas. Killip

& Smith 25972 (NY. US); Pueno Bermudez. Kdlip <t

Smith 26543 (NY, US). I^orelo: above Tamishuyacu.

Croat 19771 (AMAZ. MO); primar\ forest 17 km SW
of Iquitos. Croat 18508 {MOy. Iquilos. ca. 100 m. Killip

& Smith 26955 (NY. US); December 1830. Poeppig

s.n. (photo of specimen al \'. BM); Prov. Maynas. ad

Yuriniaguas, September 1 830. Poeppig i.n. (P); La Vic-

toria on the Amazon River. Williams 3136 (F); lower

Rio Huallaga. Williams 4797 (F, US); Yurimaguas,

Spruce 3880 (P, fragment NY); along Quebrada

Nawampa, Croat 17689 (MO); Santa Rosa, lower Rio

Huallaga below Yurimaguas. Killip & Smith 28760 (NY

,

US) Pasco: Pro\ Oxypampa. Iscoazin. Foster 7858

(F, USM) San Martin: Monte Campana. Spruce 4740

(pholo GH. US); camino a F^i>hurumbo. 7-8 km al esle

del Puente de Palo Blanco. Mariscal Cacercs. Tivachc

Nucvo. J. Schunke 5789 (NY. US); San Roque. Wil-

liams 7620 (F).

Bolivia. La Paz: Prov, Larecaja. Consata 7 km

linvards Mapiri, Beck 4924 (F. LPB); region of Mapiri.

San Amonio. fi/«/im';i.«5(US), //2.*(US), II24{K\):

region of Mapin. San Carlos, Buchticn 260 (VC). 299

(NY); Mapin. Rushy 442 (NY).

Br \/ii .Amazonas: Rio Negro. \S\9.Martiuss.n.

(photo ofM specimen aiBM) Para: Martius s.n. (photo

ot M specimen al BM)



November 1987 Monograph of Polybotra 71

12. PolybotryaglandulosaKuhn(Fig. 31, Map?).

Polyhotrya glandiilosa Kuhn, Linnaea 36:65.

1869. Type: Brazil. Amazonas: San Gabriel,

no date. Spruce s.n., not 2116 as indicated

on Morion negative, (holotype: B!),

Polyhotrya suhelliplica Lcllinger, Amcr. Fern

J. 62:56. 1972. Type: Peru. Loreto: Mis-

huyacu, near Iquitos, 100 ni, Kliif; 1390

(holotype: US!; isotypes: F!, NY).

Stem 1-2 cm thick, hemiepiphytic; scales dull

brown, thick, opaque, concolorous to bicolorous,

curved-appres,sed, 8-15 x 1.5-2.5 mm, the mar-

gins entire, the base thickened and expanded,

slightly elevated, the dorsal surface sometimes

with a medial groove. Sterile leaves up to 75 cm
long; petiole very short, up to 2.5 cm long; lamina

2-pinnate-pinnatifid, subelliptic, 65-73 x 22-28

cm, the base cuneate, about 8 cm wide, the apex

long-attenuate, the abaxial surface with or without

sessile, red, resinous, round glands, the margins

ciliate with hairs similar to those of the veins and

axes; pimiae ca. 35 pairs, the medial ones 14-16

X 3-3.5(4) cm, sessile or short-stalked, oblong,

the base truncate; pinnules catadromic, oblong,

falcate at the apex, those of the medial pinnae

15-20 X 5-7 mm, the margin entire, crenate or

lobed, generally with 6-10 veins, the basal basi-

scopic margin decurrent; axes and veins densely

pilose on both surfaces, the hairs tawny, pluricel-

lular, acicular, 1.0-2.5 mm long, the scales few

or absent. fer///e /eavM narrowly elliptic, 45 x 15

cm, 2-pinnatc-pinnatirid, botryoid; medial pinnae

6-8 X 1.5-2 cm; axes sparsely pilose, with scat-

tered, linear, appressed, dark scales; sporangial

stalks paraphysate.

Pohbotrxa glandulosa, which has been col-

lected only three times, is one of the rarest species

of Polyhotrya. It is the only species in the genus

that is endemic to the Amazon basin (Map 7);

however, 1 suspect it will be found in the adjacent

Guiana Highlands as that region becomes better

explored. I failed to find this species during my
fieldwork in the Iquitos area, in part because the

location of "Mishuyacu" is unknown. The eleva-

tional range for this speties is 100-140 m.

This species is unlikely to be confused with

any other Polyhotrya since, unlike other species

in the genus, the lamina tapers gradually to an

extremely short petiole (Fig. 31c). The long,

acicular hairs on the axes and veins further distin-

guish this species from most other congenerics.

Polyhotrya puhens. however, has long acicular

hairs and leaf cutting similar to P. glandulosa;

since it grows in Amazonian forests, the two

species could be confused. Polyhotrya glan-

dulosa, however, can be distinguished from P.

puhens because the former has a botryoid, instead

of an amphiacrostichoid, fertile leaf and a short

petiole.

The specific epithet refers to the red, .sessile,

round, resinous glands on the abaxial suface of

the type specimen (Fig. 31d). The two other col-

lections, however, lack such glands. This variabil-

ity is not unusual, since glands are variably present

in all gland-bearing species of Polyhotrya.

Specimens examined: Venezuei.a. Territorial

Federal Amazonas: Dept. Ri'o Negro, 0-3 km N of

Cerro dc Neblina Base Camp, on the Rfo Mawarinuma,

140 m, Ue.mer 16293 (MO, UC).

Peru. I.oreto: Mishuyacu, near Iquitos, 100 m,

Klug 1390 (F, NY. US).

Brazil. Amazonas: Sao Gabriel, no date. Spruce

s.n. (B).

13. Polyhotrya lechleriana Mettenius (Fig. 32,

Map 7).

Polyhotrya lechleriana Mettenius, Filices

Uchler. 1:4, tab. 1, figs. 1-5. 1854. Type:

Peru. Puno: San Gavan, Lechler 2156 (lec-

totype: B!; isotype: L!; fragments F!, US!;

photo GH! of K).

Acrostichum lechleriamim (Mettenius)

Hooker, Species Filicum 5:246. 1864, nom.

illeg., non Mettenius 1856.

Stem to 1.5 cm thick, hemiepiphytic,

mucilaginous?; .scales lanceolate, up to 15x3

mm, cream to dull brown, thin, appressed, the

margins denticulate to entire. Sterile /ca/ pubes-

cent throughout, the hairs 3-12 celled, 0.3-1.2

mm long, lax, spreading, colorless; /jc/iV)/? 1/1-1/2

as long as the lamina, stramineous; lamina fine-

ly divided, (4)3-pinnate-pinnatisect, lanceolate,

both surfaces pubescent, especially along the axes

and veins, eglandular, the apex acute, not long-

attenuate; /^mmyc lanceolate to ovate, (7)10-15 x

(2)3-7(12) cm, the base truncate, short-stalked,

crowding the rhachis;/?(>if!M/e.v catadromic, oblong

with subparallcl sides, the base truncate, nearly

sessile, crowding the costa, the apex acute to

rounded; ultimate segments single veined, talcatc,

0.5-1 mm wide; (i.vc.v pubescent abaxially, usually

with a single scale at the juncture of the costa and



72 Illinois Natural History Survey Vol.34. Art. 1

Fir.URF. 31. Polybotrya i>landiilo.ia Kuhn. a. stem scales, b. fertile leaf; c. stcnle leaf: d. aba.xial

surface of sterile pinna showing sessile reddish glands (as dots) and acicular hairs; e, medial pinna.

a-c,e: Klug 1390 (US, F). d: Spruce s.n. (B).



November 1987 Monograph of Poi.ybotra 73

1 cm

Figure 32. Polyhotrya lechteriana Mcltcnius. a. sterile leaf; b. rhachis-costa juncture of fertile leaf;

c. costal hairs; d. rhachis-costa juncture of sterile leaf; e. stem scales. a,c; Spruce 4744 (P). b,d: Stiihel

914 (B). e: Dudley 10325 (GH).



74 Illinois Naturai, History Survey Vol.34, Art. 1

costulc, the scale ovate, apprcssed. thin, brown;

grooves glabrous or puberulent within, truncated

by the ridges of the next lower order, usually

becoming shallower near the juncture. Fertile leaf

3-4-pinnate, botryoid. pubescent with hairs like

those on the sterile leaf; sori distinct, round. 0.5-

0.8 mm wide, on short stalks, these 1-2 mm long,

the receptacle moundlike; sporangial slalks

paraphysate; spores {50)54-60(65) microns long.

Other illiistralions: Hooker. Second Century

of Ferns, lab. 97. 1861; Mettenius's original dc-

.scription has an excellent illustration of the type

at Berlin.

Polybolrya lechleriana occurs in rich mon-

tane forests from (100)1000-1500 m. This species

is primarily Andean but has a notable range dis-

junction in the Guiana Highlands at Mt. Roraima

in Guyana. 1900 km from the nearest known popu-

lation in Colombia (Map 7). This disjunction is

best explained as an example of long distance

spore dispersal. Another significant Andean-

Guianan range disjunc'ion occurs in P. frac-

tisericitis (Map 3).

This species is easily recognized by its finely

divided sterile lamina with very narrow, single-

veined segments and lobes (Fig, 32). The finely

cut leaves, so unlike others in the genus, arc prob-

ably what prompted Copeland (1947) to remark

that this species "looks out of place here |in

Polyhotrya]" But P. lechleriana has all the fea-

tures of a typical Polybolrya including the unique

stem anatomy of the genus.

Some forms of this plant resemble Polybolrya

stolzei an endemic to the Andes of Colombia, but

P. lechleriana differs by the narrower width of

its ultimate segments or lobes, each of which is

one-veined (see key). Polybotrya lechleriana

probably evolved from a less dissected ancestor

in the P. alfredii group by cessation of the growth

of the marginal meristem to produce a more finely

divided sterile lamina. This species is named in

honor of WilibaldLechler( 1814-1856). a Gemian

pharmacist, botanist, and explorer, who sent many

of his collections to Mettenius.

Specimens examined: Guyana. Mi Roraini.i.

Waruina Trail. Pcrsaiul 114 (NY).

CoLOMHiA C'auca: forests of Rio Timbiqui.

l.chmann H T4I.< ((iH) Choco: alredcdorcs dc San Jose

del Palmar. Ccrro S dc la pohlaciiin. Forcrti <S Juromillo

2469 (COL, MO, NY); NW side of Alto del Bucy,

Leilinger & de la Sota 281 (COL. US); Rio Nuqui.

Haughi 5498 (COL. F, NY, US) Narino: Municipio

dc Allaquer. enlrc Junin y Bucnavisla. t50 m. Morn

42S7 (COL).

Ecuador. Chimborazo: Rfo Palora. eastern cor-

diliera of Riobatnba. l4fK) m. Rimhach 31 (GH. PH.

US). Pastaza: Mera, remnants of primary forest in hills

above town. Plowman & Davis 4542 (GH). Tun-

gurahua: along road between Puyo and Banos at second

bridge W of Mera, ca. 3 km W of Mera, 1160 m. Croat

49718 (MO. UC); Bancs-Pintuc. Siiibel 913a. 914 (B.

BM).

Peru Cuzco: Prov. La Convencion, Cordillera

Vilcabamba, Camp 2 5. 1760 m. 73°38 ' W. 12°38'S.

Dudley 10325 (GH); Paucartambo. Valle de Pillahuata.

Gerrera 1624 (US). Junin: Prov. Satipo. "Mapiri" ca.

12 km SW of Chequilavo. AC. Smith 61713 (MO.

USM); Pichis Trail. Porvenir, 1500-1900 m. Killip A
Smith 25921 (NY. US). Puno: San Gavan. Uchler2l56

(B. F. GH. L. US). 2/76 (B); La Pampa. Ri'o Tavara.

Watkins s.n. (US). San Martin: in monte Guay-

rapurima. prope Tarapoto. Spruce 4744 (L, P; fragment

US). Dept. unknown: enlre Los Rios Tintiuiguiato del

PachinyTapeciadelTocate. 1500m. Bu« /7J.^(US).

Bolivia La Paz: Hacienda Simaco sobre el

camino a Tipuani-Tale. 1400 m. Buchtien 5308 (BM.

GH, MO. NY. UC, US. Z).

14. Polybotrya attenuata Moran. sp. nov. (Fig.

33, Map 7).

Polybolrya attenuata Moran, sp. nov. T>pe:

Colombia. Antioquia: Guatape, forests on

slopes above river, 1/77, McAlpin <$ Kuhn

77-12 (holotype: NY!). The plant from which

the type specimen was taken is cultivated in

the fern greenhouse. New York Botanical

Garden (accession no. 332/78).

Planta hemiepiphytica: caulis ca. I cm diam.

:

squamulis membranaceis. linearibus. acutis. 9-

12 mm longis. 0.3-0.7 mm latis. in margine den-

ticulatis vel integris: laminae steriles lanceolatae

vel anguste dellalae, 3-pinnatae. apice longiat-

tenuatae, usque ad 55 cm longae. 26 cm latae.

glabrae vel punctatae . glandulis resinaceis. rotun-

dis: pinnulae catadromicae . 2.0-2.5 cm longae.

0.8-1 .2 cm latae. apice rotundatac: segmenta ter-

tiaria obovata. ad basim cuneata: costae et

rhachides sulcalae puhescenies in sulco. squamis

angustis, fuscatis. denticulatis: folia fertilia bot-

ryoidca: sori rotundi. ca. I mm tongi. pedicellati

.

Stem about I cm wide, hemiepiphytic; scales

light to dark brown, mosth concolorous or rarely

with a narrow hyaline border, thin, linear, 9-12 >^

0.3-0.7 mm, the margins subcntire to denticulate.

Sterile leaf up to 8 m long; petiole V\-Va as

long as the lamina; lamina to 55 x 26 cm,

3-pinnatc, narrowly lanceolate or triangular, the

apex long-attenuate, the tissue with or without



November 1987 Monograph of Polybotra 75

FiciiiRF-; 33. Pnlyhoirya altcniiala Moran. a. sterile leaf; b. pinnules and eosla-rhachis juncture (note

the abundance of narrow, dark scales on the axes); c. stem scales; d. two pinnules of fertile leaf;

e. fertile leaf, a-c: McAlpin & Kuhn 77-12 (NY).



76 Illinois Natural History Survey Vol. 34, Art. 1

punctate, resinous, spherical, sessile glands; pin-

nae 10-13 X 3.5-4.0 cm long, lanceolate to ob-

long, the base truncate, short-stalked, crowding

the rhachh; pinnules catadromic, 2.0-2.5 x 0.8-

1.2 cm, with only 5-7 lobes or segments, the

apex acute or rounded, the base truncate, sessile

or with a short stalk less than 1 mm long; tertiary

segments obovate, entire to slightly lobed, con-

taining several vein branches, the base cuneate;

axes pubescent within grooves and especially ab-

axially, the hairs 4-12 celled, 0.2-0.3 mm long

jointed, with reddish cross-walls; costal scales

numerous, narrow, dark, denticulate, tortuous, ap-

pressed; grooves pubescent within, the hairs red-

dish, multicellular. Fertile leaves botryoid,

3-pinnate, densely scaly, the scales grading into

jointed hairs (the hairs are actually reduced scales);

sori stipitate, the stalks 1-3 mm long; sporangial

stalks paraphysate.

Polybotrya altenuata, named for its distinc-

tive, long-attenuate leaf apices, is endemic to the

Andes of Colombia (Map 7). This species differs

from P. lechleriana. a similar species, by its long-

attenuate apex, oval tertiary segments, resinous

glands on the lamina, and the darker, narrower

scales on the axes. The pinnules, furthermore, are

shorter and stubbier, having only 5-7 lobes and/or

segments, and these have a cuneate base (Fig.

33a,b).

The presence of glands on the lamina is not

constant. The living plant from which the holotype

was made has always produced leaves having

round, sessile, resinous glands on the undersur-

face. The Lehmann specimen, however, lacks

such glands. Similar glands are also variably pres-

ent in other species, such as P. osmundacea and

P. glandulosa.

Specimens examined: Colombia. Antioquia:

Guatape, McAlpin & Kuhn 77-12 (NY). Nariho: along

river Pipulquer. west slopes of the western Andes,

Lehmann 500B (US).

15. Polybotrya stolzei Moran, sp. nov. (Fig. 34,

Map 9).

Polybotrya stolzei Moran, sp. nov. Type:

Colombia. El Valle: Santa Rosa, Dagua Val-

ley, forest along Rio Cabellcte, 200-300 m,

22 September 1922, Killip 11549 (holotype:

US!; isotypes: GH!, NY!, PH!).

Planta hemiepiphytica: caulis 0.6-1.5 cm
diam., Imucilaginus?): squamae fuscatae,

opacae, lineares, integrae, usque ad 12 mm lon-

gae. 0.4 mm latae: lamina sterilis 3-pinnata.

plerumque pubescens in superftciebus ambabus:

rhachis et costae pubescentes. squamis ovatis

tenuibus aliquot; pinnulae catadromicae , pro

parte maxima sessiles, usque ad 33 "X- 9 cm; seg-

menta tertiaria ovalia vel oblonga Integra, (5)6-

8(10) X (2)3-4(5) mm. Folia fenilia ignota.

Stem 0.6-1.5 cm thick, hemiepiphytic,

mucilaginous (?); scales dull brown, appressed,

opaque, linear, 0.4-12.1 x 0.3-1. mm, margins

subentire. Sterile leaves up to I m long; petiole

'A to 1/2 as long as the lamina; lamina lanceolate

orsubdeltate, 30-65(80) x 19-32(60) cm, 3-pin-

nate, usually pubescent on both surfaces, always

so along the veins beneath, the hairs whitish to

tawny, articulated, 0.1-0.4 mm long, spreading;

pinnae to 33 x 9 cm, narrowly triangular to lan-

ceolate, equilateral or, in the basal pair, with the

basiscopic side slightly prolonged, the base trun-

cate, short-stalked, crowding the rhachis; p/nnu/ej

catadromic, oblong, the base truncate, nearly ses-

sile, the apex rounded to acute; tertiary segments

ovate to oblong, (5)6-8(10) x (2)4(5) mm. with

several vein branches, the margins entire to

slightly lobed; axes pubescent abaxially, with a

few thin, ovate, brown scales, especially at the

pinnule junctures; grooves nearly glabrous to mod-

erately puberulent within, never packed with long-

protruding hairs, truncated by the ridges of the

next lower order. Fertile /fa/ unknown.

This new species is named for Robert G.

Stol/e, pteridologist at the Field Museum of Nat-

ural History, Chicago. Illinois. He onginally

suggested this genus to mc and has given much

help and encouragement dunng the preparation of

this monograph.

Polybotrya stolzei is endemic to the western

Andes of Colombia (Map 9). occurring in wet.

shaded forests from 2(X)-1750 m. The laminar

cutting of P. stolzei is intermediate between P.

lechleriana and P. alfredii. Extremeh large pin-

nules of P. stolzei look like those of P. alfredii

but are distinguished by the pubescence on both

surfaces of the lamina and by the blunter apices

of the medial pinnules. Smaller, more delicate

forms of P. stolzei may look like P. lechleriana

but are distinguished by their oblong tertian, seg-

ments with more than one vein f>er segment. Un-



November 1987 Monograph of Poi ybotra 77

FiciURF. 34. Polybotrya stolzei Moran. a. costal scales from the pinnule-costa junctures; b. abaxial

view of medial pinnules of "c"; c. medial pinna; d. stem scales; e. basal pinna of large leaf; f. adaxial

view of rhachis-costa juncture, a-d: Killip 1 1549 (GH). e,f: Leilinger & de la Sola 2H4 (COL).



78 III iNois Natural Hisidry Survky Vol.34. Art. 1

fortunately, none of the collections contain fertile

leaves and this structure, therefore, cannot be com-

pared to the distinctive botryoid fertile leaf of P.

lechlerianii.

Specimens examined: Colombia. Choco: NW
side i)f Alto del Buey, LeUinger & de la Sola 2fiO (COL,

US), 2H4 (COL, US); principal ridge and slopes 2 km

E of San Jose del Palmar, 15.5()-I6()() m, LelUnger &

de la Sola 747 (COL, LP, US), 74« (COL, LP, US).

El Valle: Santa Rosa, Dagua Valley, forest along Rio

Caballele, 200-3(X)m, /fi7/(p //549 (GH, NY, PH, US).

16. Polybotrya alfredii Brade (Fig. 35, Map 8).

Polybotryci cdfredii Bradc, Bradca 1:12, tab.

1, fig. 2. 1969. Type; Costa Rica. San Jose;

Tablazo, 1900 m, July 1908, (Brade's origi-

nal description states August, apparently an

error.) Brade & Brade 98 (holotypc; HB;

isotypc; NY!).

Polybotrya alfredii Brade f. carpinterae

Brade, Bradea 1:13. 1969. Type: Costa Rica.

San Jose: Carpintera, terrestrial, 1500 m, 25

November 1908, Brade & Brade s.n.

(holotype: HB).

Polybotrya gracilis Brade, Bradca, 1:14, tab.

1, fig. 3. 1969. Type: Costa Rica. Cartago:

Tablazo, 1900 m, 28 August 1908, Brade

554 (holotypc: HB).

Stem 1-2 cm wide, hemiepiphytic, the apex

in living plants usually covered by a thick mucilage

that disappears upon drying; scales dull, dark to

light brown, opaque to translucent, 8-16(22) x 1-

1.5(2.0) mm wide, narrowly lanceolate, spread-

ing, attached across the width of the slightly thick-

ened base, the margin denticulate or more com-

monly entire. Sterile leaves up to 1 .4 m x 0.7 ni;

petiole up to 35 cm long; lamina ovate, up to

3-pinnate-pinnatifid, subchartaceous, often drying

greyish green, the proximal margins of the pinnule

bases often ciliatc, the hairs less than 0.1 mm
long; pinnae up to 45 X 22 cm, apex acuminate:

pinnules up to 12 X 5 cm, ovate to narrowly trian-

gular, the base truncate and symmetrical, those

of the basal pinnae anadron)ic or subcqual, those

of the medial pinnae catadromous; ullimute seg-

ments ovate to oblong, entire, crenulate, serrate

or lobed, the apex obtuse or acute; axes evenly

pilosulous abaxially, the hairs 0.1-0.2 mm long.

usually tawny; grooves pubescent within, very

shallow or even disappearing within about 1 cm
of the rhachis so that the costa is almost terete

where it joins the rhachis. Fertile leaves up to

0,8 X 0.35 m, up to 3-pinnate-pinnatirid, ovate,

coenosoric; axes pubescent, the hairs short, 0.1-

0.2 mm long, usually with ovate to lanceolate,

appressed, thin, denticulate to fimbiiate scales;

sporungial stalks paraphysate; spores (50)55-

62(66) microns long.

Other illustrations: See the original descrip-

tions cited above.

The altitudinal range of Polybotrya alfredii

is 7(X)-190O m. In Ecuador, this species grows

in rich montane forests in the eastern Andes. In

Costa Rica, this species occurs primarily in wet.

shaded cloud forests, usually growing alongside

P. gomezii. Polybotrya alfredii produces numer-

ous terrestrial leaves that often predominate the

forest floor. Twice in Costa Rica and once in

Ecuador, 1 saw fertile leaves produced from the

terrestrial portion of the stem. These are the only

instances in which I saw fertile leaves produced

tertestrially by a scandent species.

Polybotrya alfredii looks like P. lourteigiana

but can be distinguished by its more highly cut

lamina. Since leaves lower on the stem tend to be

smaller and less divided than those higher on the

stem, cutting should be compared using pinnae

over 25 cm long (sec couplet 24 in the key). The

two species also differ in the shape of the ultimate

segments or lobes; P. alfredii has relatively more

ovate and shorter segments than P. lourteigiana

(compare Figs. 35 & 37).

Unlike all other species of Polybotrya. P.

alfredii has the stem apex and the lowermost 5 to

1 5 cm of the petiole covered w ith a thick, translu-

cent mucilage. Mucilage is totally lost upon drying

and therefore not visible on herbarium specimens.

The function of this mucilage is unknown. More

fieldwork is needed to determine if this unusual

mucilage occurs in two closely related species. P.

lechleriana and P. lourteigiana. Nectaries are

another unusual feature of the morphology of P.

alfredii During tleldwork in Costa Rica, I saw

nectaries on the rhachis of P. alfredii like those

described by Koptur et al. (1982) for P. osmun-

dacea. Sec the Morphology and .Anatomy section

of this monograph for further information concern-

ing nectaries.



November 1987 MONCWRAPH of POI.YBOTRA 79

15 cm

FiciURF. 35. Polyhoina alfredii Bradc. a. sterile leaf; b. slcm scales, the middle shown sideways;

c-g. pinnules, acroscopic side to the right; h. fertile pinnule, adaxial view; i. adaxial surface of costa

and pinnule base, a.b.f.h.i: Moran 2442 (CR). c; Moran 3532 (F). d: Buchtien 5164 (US), e: Moran

3214 (CR). g: Dwyer 8337 (MO).



80 iLLiNors Natural History Survry Vol. 34. An. 1

I put P. gracilis in synonymy with P. alfredii,

although the former appears distinct because the

leaf is smaller, only 2-pinnate, and has obtuse

pinnule apices (Fig. 35e). Polyhotrya gracilis re-

sembles P. alfredii, however, in such characteris-

tics as scales, pubescence, stem mucilage, habitat,

and range. Furthermore, at several localities in

Costa Rica, I collected plants that exhibited com-

plete intergradation between the typical large

leaves of P. alfredii and the smaller ones of P.

gracilis. Polybotrya gracilis, therefore, represents

a small-leaf form of P. alfredii that has well-

rounded pinnule apices. I chose the name P. al-

fredii over P. gracilis because an isotype has been

seen. This species is named in honor of Alexander

Brade's brother and field companion, Alfred

Brade.

Specimens examined: Nicaragua. Rivas: Isla

de Ometepe, NW slope of Volcin Maderas, Stevens <S

Krukoff65I8 (CR); Graytown, Camp Murroeal, Shimek

& Smith s.n. (MICH).

Costa Rica. Alajuela: Univ. of San Ram6n

biological field station. Moran 3214 (CR, F, GH, MO);

12 km NNW of San Ram6n by road on way to San

Lorenzo. 1 km S of Balsa, Liesner & Judziewicz 14855

(CR. MO); San Jer6nimo. Werckle 559 (US); Suerre.

Ilanuras de Santa Clara. Donnell Smith 6939 (US); Santa

Clara. Cooper 10240 (US); NW of Zarcero, ca. 2 km
W of Zapote on dirt road to Sta. Elena. Lellinger <S

White 1361 (US); San Antonio de Zarcero. AC. Smith

48/293 (US); N of San Ram6n, ca. 4 km N of Balsa

along road to Colonia Palmerena, Lellinger & While

1244 (US); Santiago de San Ramon, Brenes 21980 (F),

14248 (GH); Alto de la Raima de San Ram6n. Brenes

11676 (F, NY); 7 miles N of San Ramon square along

ridge road, Evans & Bowers 2944 (MO); Finca La Selva.

San Rafael de Vara Blanca, pendiente N del VolcSn

Barba, Jimenez 803 (F); vicinity of La Palma, MtLxon

412 (NY); 11 km N of San Ramon, Mickel 2958 (LP,

NY. UC); San Luis de Zarcero, AC. Smith 778 (NY).

Cartage: road between La Suiza and Tuis, DeWolf385

(CR); Pacayas, Lankester 653 (US); along Inter-Amer-

ican Hwy. between Cartago and San Isidro del General,

Scamman 5985 (CR), 5986 (CR); Orosi, Finca del Sr.

Valverde, Brade 16824 (CR); El Muneco, S of Navarro,

Standley 33648 (LIS); Sanui Clara de Cartago, Maxon

& Harvey 8242 (US); SE of Plalanillo, along Camino

Raiz de Hule, Croat 36761 (MO); Platanillo, de la Sota

5252 (LP, US); foolof Volcdn Turrialba, Pacayas, Scam-

man 7155 (CR); along road from Moravia to Quebrada

Plalanillo (Tsipiri), Croat 36669 (MO); ca. 22 km E of

Turrialba, high ridge above Platanillo, Mickel 3426 (LP,

NY); near the town of Estrella, Moran 2237 (CR). 2241

(CR); near Alto Patillos, Moran 2442 (CR); Reserva e

Tapanlf, Gdmez 18906 (CR, MO, UC); Tapantf,

Grayum & Sleeper 3448 (MO); 1 2- 1 6 km S of Tapantf,

along the road above the Rio Grande de Orosi, Stolze

1488 (F. UC). Heredia: 7.5 km N of Vara Blanca, in

vicinity of Ri'o La Paz Grande, Croat 36007 (MO); Vara

Blanca de Sarapiqui, N slope of Central Cordillera,

Skutch 3236 (GH, US); Santo Domingo de Vara Blanca,

Valerio 2356 (US); Vara Blanca, between Volcan Poas

and Barba, Chrysler & Roever 5095 (US); 4 km N of

Vara Blanca, Croat 35608 (MO); near Rio Las Vuellas,

NE of Volcan Barba, Lent 2642 (F); Virgen del Socorro,

Moran 3168 (CR). Puntarenas: 5 km NE Finca las

Cruces, San Vilo de Java, Burch 4505 (NY): near field

station, Rincon de Osa, Burch 4414 (NY); 5 km S of

San Vito de Java, in vicinity of biological field station

at Wilson finca. Mickel 2003 (NY); 5 km S of San Vito

de Java. 1-4 km SW of biological station at Finca Wil-

son, Mickel 3126 (NY). San Jos*: Tablazo. Valerio 63

(US); Biolley 73 (US, Z); Tablazo. Brade & Brade 67

(P), 98 (NY); Alto U Palma, Moran 2325 (CR).

Panama Chiriquf: Palo Alto, just E of Boquete.

Stem et al. 1034 (CR. US); Holcomb trail. Common
927 (MICH. UC, US); vicinity of El Boquete, Common
1178 (US), 1233 (US), Maurice 699 (US); Rfo Caldera,

from El Boquete to die Cordillera, Killip 5202 (CR.

US); valley of Rio Piamasta. about 5 mi E of Boquete.

Killip 5163 (US), 5386 (US), 5429 (CR, MICH. US).

Cocl*: Valle de Ant6n. Ocampo 877 (CR); Cerro Pilon.

Dwyer 8337 (MO); cloud forest at La Mesa above El

Valle. Croat 13431 (MO. UC).

EcUAiXJR. Napo: camino Baeza-Tena. 15 km al

norte de Tena, 1 100 m, Moran 3532 (F, GH. PORT.

Q, (JCA). Pastaza: village of Ri'o Chico. 8 km from

Puyo, Shemluck 304 (F). Tungurahua: casi 25 km al

este de Banos. detras el pueblo Rio Negro, por Ri'o

Topo (malempresado "Rfo Toro" en el mapa). Moran

3570 (F, GH, Q. QCA).

Peru . Hu^nuco: SW slope of the Ri'o Llulla Pichis

watershed on the ascent of Cerro del Sira. Dudley 13258

(GH), I8265A (GH).

Bolivia. La Paz: Hacienda Simaco sobre el

camino a Tipuana, Buchtien 5164 (GH, US).

17. Polybotrya botryoides (Baker) C. Chr. (Fig.

36, Map 8).

Polybotrya botryoides (Baker) C. Chr. . Index

Filicum 504. 1906.

Acroslichum botryoides Baker. J. Bot.

19:206. 1S81. Typf: Colombia. Antiixiuia:

on trees in the forest, 6000-7000 ft, 22 July

1880, Kalhreyer 1873 (hololype: K: photo

and fragment GH!. US!).

Stem to 2 cm thick, hcmicpiphytic: scales dull

brown, about 1 cm long, with the margins subcn-

tire to erose, the base curved and thickened. Sterile

leaves up to 1.5 m (?): petiole densely scaly al



November 1987 Monograph of Poi ybotra 81

Figure 36. Polyhotrya hotryoides (Baker) C. Chr. a. costal scales, the one on the right enlarged to

show the biseriate apex; b. uniseriate "hair" from the costa. Such hairs arc actually reduced scales as

shown by their intergradation with the scales and by their characteristics which differ from those of

true hairs; c. sterile pinnule base; d. botryoid fertile pinnule; e. sterile pinnule; f. stem scale, at right

is a side view showing the thickened base; g. petiolar scale, medial portion darkened and margins

crosc. a-g; Juncosa 1369 (MO).



Illinois Natural History Survky Vol. 34, An. I

base, the scales similar to those on the stem, but

with wider pale borders and more strongly erose;

lamina 4-pinnate, nearly glabrous on both sur-

faces; pinnae lanceolate, up to 35 x 20 cm, finely

divided all the way to the apex; pinnules up to

10x3 cm. catadromic or subequal. lanceolate,

the base often overlapping the costa, subequilat-

eral, i.e.. not prolonged acroscopically. short-

stalked (1-2 mm long); tertiary sef;menl.t oblong,

sessile, up to 2 x 0.9 cm, apex acute or rounded,

base equilateral; quaternary seamenls oblong to

ovate, sessile, up to 5 x 3 mm, the margins entire

or crcnatc; axes scaly, the scales numerous, tortu-

ous, thin, reddish, most of these only 2-5 cells

wide, intergrading with hairs, these articulate, red-

dish, 4-8 celled. 0.1-0.3 mm long; grooves

pubescent within by reddish hairs. Fertile leaves

botryoid, 3-4-pinnate, with the ultimate soriferous

segments globose. 1-2 mm diameter; receptacle

paraphysate by articulate hairs, these often buried

among the sporangia; sporangial stalks paraphy-

sate; spore size unknown.

Polybotrya botryoides is endemic to the

Andes of Colombia (Map 8), occurring in forests

from 1830-2130 m. The above description is

based primarily on the Juncosa collection, which

was the only dried specimen I had available for

study. I could not find enough spores on this speci-

men to make an adequate measurement of spore

size.

This rare species can be separated from all

Dther species of Polybotrya by its distinctive com-

bination of botryoid fertile leaf. 4-pinnatc sterile

leaf, and axes covered by numerous, linear, tortu-

.lus, reddish scales (Fig. 36a). No other species

.if Polybotrya is as finely divided as this one.

Polybotrya botryoides looks like P. alfredii be-

;ause of its large decompound lamina, catadrom-

ically arranged pinnules, subequal pinnule bases,

pubescence of the axes, and dull brown stem

»cales. The Venezuelan endemic P. canaliculata

also has botryoid fertile leaves, but it lacks hairs

.)n the axes, has very different axial scales, and

IS less finely cut (Fig. 52).

Specimens examined: Coiomhia .Antioquia:

Kalhrcyer IS7.i (photn and Iragrncnl GH. I'S). main

nwy. Mcdcllin-Pucrlii Triunlb. Municipality Cocoma,

;a. 5 km E of Cocoma Pcajc. Qucbrada El Biadal , I S.^0

11, Juncosa 1.^69 (MO, UC).

18. Polybotrja lourteigiana Lellinger (Fig. 37,

Map 10).

Polybotrya lourteigiana Lellinger, Proc.

Biol. Soc. Washington 89:723, fig. 5. 1977.

Type; Colombia. Choco: Trail along ridge

from the confluence of the forks of the Rio

Mutata above the Ri'o Dos Bocas to the top

of AltodelBuey.ca. \450-\150m, Lellinger

& de la Sola 251 (holotyf)e: US!; isotypes:

COL!. LP!).

Stem to 2 cm thick, hemiepiphytic. mucilagin-

ous (?); scales dark brown, concolorous. and en-

tire, or cream-colored with a darker center and

ciliate margins, mostly 8-17x0.5-2.0 mm.
Sterile leaves up to 1.6 m long; petiole up to Vz

as long as the lamina; lamina glabrous on both

surfaces, up to 3-pinnate but mostly 2-pinnate-

pinnatifid throughout, deltate to lanceolate; lower

pinnae mostly 30-45 x 10-18. elliptic, lanceo-

late or narrowly triangular: pinnules arranged

catadromically, subequilateral at base, subsessile,

narrowly oblong, generally 10-15 free pairs, 5-

8(11) x 1.5-2.0(3.0) cm. the apex slightly fal-

cate; tertiary segments 5-12(15) x 2-4(6) mm.
arranged catadromically. oblong, the ap)ex

rounded to truncate, the margins entire; axes

pubescent abaxially , the hairs either tawny or red-

dish, 0.5-1.2 mm long, multicellular, lax, or with

the hairs shorter, stiffer, 0. 1-0.3 mm long, costal

scales present, inconspicuous, often at the

junctures with the costules. ovate to orbicular,

flaccid, brown; grooves pubescent within, espe-

cially at the junctures, truncated by the ridges of

the next lower order. Fertile leaves 4-pinnate,

coenosoric, or subbotryoid, if the son discrete,

then not on long stalks; costa pubescent, with or

without ovate, denticulate scales: sporangial

stalks paraphysate; spores (50)55-67(70) microns

long.

Other illustrations: See Lellinger's original

description (cited above) which contains a photo-

graph of a portion of the holotype.

This plant occurs in mid-elevation forests

from 750 to 2250 m in the western Andes of Co-

lombia (Map 10). See comments under Polybotrya

alfredii and P. pittieri for comparisons with those

species. Considerable morphological variation

exists in the specimens cited below, but I found



November 1987 Monograph of Poi.ybotra 83

Figure 37. Polyhotrya lourteigiana Lcllingcr. a. basal pinnae; b. pinnule bases, apex to the left;

c,d. sterile pinnules, apex to the left; c. stem scales; f. fertile pinnules. a,b,c,e,f: Letlinger & tie la

Sola 251 (US), d: Cuatrecasas 15527 (US).



84 Illinois Nm ural History Survey Vol.34. Art. 1

no constant characters to separate the extreme

forms. F^ubescence varies tremendously; the Lel-

linger and de la Sota specimens are mostly long

pilose on the axes whereas the Cuatrecasas speci-

mens have shorter, stiffer hairs. Intermediates

exist, and no other characters correlate with pubes-

cence.

The type collection consists of six sheets, of

which US #2748016 is here excluded; it actually

represents P. stolzei.

Specimens examined: Colombia. Choco: Trail

along ridge from the confluence of the forks of the Ri'o

Mutata above the Ri'o Dos Bocas to the top of Alto del

Buey. ca. 1450-1750 m, LeUinger & de la Sota 251

(COL, LP, US); principal ridge and slopes 2 km E of

San Jose de! Palmar, LeUinger A de la Sota 768a (COL,

LP. US); 0.3 km E of the Ciudad Boli'var-Quibdo road

across the suspension bridge at ca. km 141, LeUinger

& de la Sota 899 (COL, LP, US); Mojarras de Tado,

8.5 km E of Istmina, LeUinger & de la Sota 387 (CR,

COL, LP, US). Choc6-Antioquia: principal ridge W
of La Mansa, at ca. km 105.5 of the Ciudad Boli'var-

Quibdo road, LeUinger & de la Sota 948 (COL, LP,

US) Choc6-EI Valle: Serrania de Los Paraguas, along

the trail from El Cairo to Rio Blanco, ca. 8 km SW of

El Cairo. LeUinger & de la Sota 843 (COL. LP, US).

El Valle del Cauca: Hoya del Ri'o Call, La Margarita,

Duque-Jaramillo 1868 (COL); Cordillera Occidental,

vertienle occidental, del Ri'o Sanquinini, La Laguna.

Cuatrecasas 15527 (US); Cordillera Occidental, ver-

tienle occidental, monte La Guarida, filo de la Cordillera

sobre La Carbonera, entre Las Brisas y AlbSn, Cuatre-

casas 22143 (F, US). Narino: Municip. de Altaquer,

Carretera enffe Altaquer y Juni'n, Guyambe. Mora 4160

(COL).

19. Polybotrya pittieri LeUinger (Fig. 38, Map

11).

Polybotrya pittieri LeUinger, Proc. Biol. Soc.

Washington 89;725, fig. 6. 1977. Type: Co-

lombia. Cauca: Dagua Valley, C6rdoba,

Pacific Coastal Zone, 30-100 m, Pittier 587

(holotype: US!).

Stem 1 cm thick, hemiepiphytic; scales 9-14

X 1.0-1.4 mm, spreading, reddish, membrana-

ceous, the margin denticulate. Sterile leaves up

to 1.3 m long; petiole to 38 cm long; lamina trian-

gular at base, to 60 x 90 cm, 3-pinnatc at the base,

2-pinnate-pinnatifid medially, nearly glabrous on

both surfaces, the apex long-tapered; pinnules ar-

ranged catadromically, up to 5 x 2 cm, relatively

short and stubby, the base subcquilatcral, the apex

obtuse or slightly acute; tertiary segments up to

1.0 X 0.5 cm, oblong; axes pubescent on the ab-

axial surface, the hairs short, stiff, ca. 0.2 mm
long; grooves truncated by the ridges of the next

lower order or only indistinctly admitted to the

groove, pubescent within, the hairs reddish or

tawny. 0.2-0.5 mm long. Fertile leaves subtri-

angular, 4-pinnate, botryoid, the apex long-

attenuate; pinnules to 2.1 X 1 cm; sari stipitale,

round, the stalks 1-4 mm long, arranged pin-

nately; sporangial stalks paraphysate; spores

(62)65-70(77) microns long.

Other illustrations: Lellinger's original de-

scription has a photograph of the type.

Polybotrya pittieri, known from only two col-

lections, is endemic to the western Andes of Co-

lombia (Map 1 1 ). This species occurs at low ele-

vations, from 30-100 m. unlike its most closely

related species, P. attenuata and P. botryoides.

which occur at higher elevations. It is named after

Henry F. Pittier (1857-1950), a Swiss botanist

and civil engineer who collected prolifically in the

American tropics.

Polybotrya pittieri looks most like P. lour-

teigiana. LeUinger (1977) observed only that P.

pittieri had a rather stiffly pilose indument on the

rhachis and costae, whereas P. lourteigiana had

a lax, hirtous indument. From the greater number

of specimens of P. lourteigiana available to me,

however, pubescence type does not appear con-

stant. Nevertheless, P. pittieri seems to be a dis-

tinct species, and the best way to distinguish it is

by its botryoid fertile leaves with prominently

stipitate sori (Fig. 38b). P. lourteigiana. on the

other hand, has coenosoric fertile leaves and sub-

sessile sori (Fig. 37f). Another difference, al-

though one that may prove inconsistent when more

specimens are known, is the broadly dellate lamina

of P. pittieri: P. lourteigiana is slightly reduced

at the base. Using these characters. I find that the

paratypcs of P. pittieri actually belong to P. lour-

teigiarui

.

Polybotrya pittieri might also be confused

with P. camiliculata since both have botryoid fer-

tile leaves, but P. pittieri differs by its broadly

deltatc lamina, pinnule cutting, and pubescent

axes that lack scales. Its spores are extremely large

compared to others in the genus, suggesting that

P. pittieri is a polyploid.

Specimens examined: Colombia Cauca:

Dagua Valley. C6rdoba. Pacific Coa.stal Ztme. 30-100

m. Pimer587^VS): El Tambo, von Sneidem /590(US)



November 1987 Monograph of Poi.ybotra 85

v-rn/^-^.-'V

V • -^2?
;_-i3

1 cm

FrouRF. 38. Polyholrya piltieri Lcllingcr. a. sterile leaf; b. fertile leaf; c. stem scales; d. sterile basi-

scopic pinnule from basal pinna, a-d: Piiiier 587 (US).



86 Illinois Natlkai History Survey Vol.34. An. 1

20. Polybotrya cylindrica Kaulfuss (Fig. 39,

Map 16).

Polybotrya cylindrica Kaulfuss, Enum.

Filicum 56. 1824. Type: Brazil. Santa

Catarina: Island of St. Catharina, Chamisso

s.n. (LE).

Polybotrya incisa Link, Hort. Berol. 2; 1 35.

1833. Type: Brazil. No collector or date

given (holotype; B!).

Acrostichum incisum (Link) Hooker, Species

Filicum 5:245. 1864.

Acrostichum cylindricum (Kaulfuss) Hooker,

Species Filicum 5:246. 1864.

Polybotrya frondosa Fee, Crypt. Vase. Bre-

sil. 1:15. 1869. Type: Brazil. No exact lo-

cality given, Glaziou 2428 (holotype: K;

photo GH!).

Polybotrya osmundacea var. cylindrica

(Kaulfuss) Luetzelb. Estudo Bot. Nordeste

Brazil 3:245. 1923.

Polybotrya osmundacea var. frondosa (Fee)

Luetzelb. Estudo Bot. Nordeste Brazil 3:245.

1923,

Polybotrya osmundacea var. incisa (Link)

Luetzelb. Estudo Bot. Nordeste Brazil 3:245.

1923.

Stem 1-2 cm thick, hemiepiphytic; scalesduW

brown, opaque, concolorousorwith a very narrow

hyaline border, entire, generally 6-12 mm long,

the base thickened, curved, appressed. Sterile

leaves up to 1.7 m long; petiole 30-50 cm long;

lamina to 1.3 m long, 3-pinnatc-pinnatifid, gla-

brous or pubescent with small, unicellular, whitish

hairs, the margins sparsely to moderately ciliate

by jointed hairs, these ca. 0.1 mm long; pinnae

narrowly lanceolate to subdeltate, the lowermost

to 52 X 28 cm; pinnules generally 9-14 x 3.5 x

5.5 cm, anadromic proximally, becoming cata-

dromic distally, the acroscopic side slightly pro-

longed, proximal ones with a short 3-5 mm stalk;

tertiary segments ca. 7-1 1 per pinnule, ovate to

oblong, the largest usually pinnatisect, the apex

acute or rounded; vein tips slightly clavate and

prominulous; axes abaxially sparsely to densely

pubescent, the hairs colorless, clear, stiff, subulate

ca. 0.1-0.3 mm long, with the cross walls not

easily visible, mixed with appressed, linear,

scraggly, inconspicuous scales; groovr.v filled with

reddish hairs. Fertile leaves to 90 x 62 cm, 3-4-

pinnate; ultimate segments often oblong and ap-

pearing cylindrical due to folding back of the seg-

ment margins; sporangial stalks paraphysate;

spores (55)60-66(70) microns long.

Other illustrations: Fee, Mem. Fam. Foug.

2 (Hist. Acrost.), tab. 36. 1846; op. cit. tab. 35

(as P. incisa); Brade, Bradea, 1(9):66, figs. 1 &
2;67,figs. 16&17. 1969; Sehnem, Flora Ilustrada

Catarinense, plate 19; plate 20 (as P. cylindrica

\w. frondosa). 1979.

Polybotrya cylindrica, endemic to the coastal

mountains of southeastern Brazil (Map 16). is the

largest and most finely divided species of Polybot-

rya in Brazil. It occurrs in wet, shaded, primary-

forests from to 775 m. This species differs further

from other Polybotrya species in southeastern

Brazil by its stem scales that are dull brown with

entire margins and the thickened and curved bases.

The curved scale base orients the rest of the scale

parallel to the surface of the stem, so that the

scales are appressed (Fig. 39f,g). Another helpful

feature in identification is the laminar margin,

which is sparsely ciliate by jointed hairs less than

0.1 mm long.

Brade ( 1971 ) distinguished P. frondosa from

P. cylindrica by the amount of lamina dissection.

But lamina dissection exhibits complete intergra-

dation from 2-pinnate to 3-pinnate-pinnatifid and

no other characterisitcs correlate with it (Fig.

39a-d).

Although I have not seen the type of this

species, the original description by Kaulfuss is

sufficient to ascertain his plant's identity. He spe-

cifically mentions the ciliate margins that distin-

guish this plant from others in southeastern Brazil.

Furthermore, this species is the only Polybotrya

that grows on the Island of Santa Catarina, where

the type was collected. The specific epithet prob-

ably refers to the fertile segments which, upon

drying, reflex so that the sori appear on all sur-

faces, thereby imparting a cylindrical shape to the

segments.

Specimens examim'd: Bra/ii Espirito Santo:

Mumcipio dc Cachocira dc ll.ipomirim-Vargcni {"!).

Rniiti- 19^7: (HB. MO. NVV Parana: Mun Guaraluba.

Pcdra Branca dc ,\raraquara. Hiiischhiich S14I (US);

Pcdra Branca dc .Araraquara. Hmsihhiich 7424 (US);

mi locality. Huischhach 10749 {>i\). no iiK-ality. Dii.fcn

UbSfi (GH); Porto Dom Pedro II, Dusen 44:.hBS\):



November 1987 Monograph of Polybotra 87

FiouRF. 39. Polyhotrya cylindrica Kaulfuss. a. sterile pinna; b-d. sterile pinnules showing variation

in amount of dissection; e. fertile pinnules; f. stem scales, top view; g. stem scale, side view.

a,e: Goedas 217 (NY), b: Mosen 3050 (B). c.f,g: Schmah I5S (UC). d: Du.sen 6926 (F).



88 Illinois Natural History Survey Vol. 34, Art. I

Vdlla Grande, Dusen 6984 (BM); Porto de Cima, Du.sen

6926 (F, GH, LIL, MO, NY, PH, UC, Z); Tacarehy.

Dusen 15351 (BM, F, PH), 15353 (GH, MO, PH, UC.

Z) Rio de Janeiro: without locality, Hunnewell 1851

1

(MICH); Distiito Federal, Guanabara, Represa Camori,

Brade 12576 (RB); Rezende, 80 mi WNW of Rio, on

Paraiba River, Rio Palmital, Castellanns 25710 (F);

Serra de Itatiaia, Brade J 2614 (NY, RB); Brene. Glaziou

955(RB); vicinity of Meio da Serra, Smith & Brade 2286

(GH); Estrada Velha de Barrira a Teresopolis, Duarie

& Pereira 65323 (F, LP); Corcovado, Miers s.n. (NY);

Teresopolis, Brade 9365 (NY, UC); Serra da Carioca,

Brade 13982 (RB). Rio Grande do Sui: S. Leopoldo.

Lehnun 2562 (LIL); Porto Alegre, Fargens 251 (UC).

Santa Catarina: Municip Ibirama, Horto Florestal.

Smith & Klein 7546 (US); Horto Florestal, I.N. P.,

Ibirama, Rett: & Klein 1 101 (US), 1102 (US). 2665

(US), 3466 (US); Blumenau, Wamow, Gnedas 217

(NY, UC, US); Itajai. Reit: 151 (LIL); Municipality

Brusque, Azambruja, Smith and Reitz 6135 (COL, GH,
MO, US); ReseiA'a Florestal, dos Piloes, Duarie 3179

(LP, NY); Mina Velha, Garuva, Sao Francisco do Sul,

Reil: & Klein 4635 (US), 4682 (US): Sabia, Vidal

Ramos, Reit: & Klein 5108 (US); Passo Mansa, Haer-

chen 124 (NY); Joinville, Schmal: 78 (MO), 158 (NY,

UC); Tres Barras, Garuva, Sao Francisco do Sul, Reit:

(4 Klein 5549 (US); Sta. Catherine 1834, Caudichaud

73 (P); Azambuja-BrtJsgui, Reit: 2825 (LIL, P);

Blumenau, Mu//fr60.?9 (MICH). Sao Paulo: Sao Bcnlo.

Luetzelhurg 254 (MICH, US); Sen-a da Bocaina. Brade

20931 (LP); Morro das Pedras, Iguape, Brade s.n. (US);

Paranapicaba, Biological Station, Handro 1229 (GH,

US); Ague funda, Handro 2228 (GH, US); Serra do

Mar, Wackel 21556 (GH); Campo Grande to Alto da

Serra, 40 km SE of Sao Paulo, Tryon & Tryon 6591

(GH); Santos, 1874, Mosen 3050 (B, P). State un-

known: Gla:iou 2428 (photo of K specimen at GH); no

locality, Claussen s.n. (P); Micken s.n. (NY); Capao do

Fradie, Schneus 3412 (GH); Morro do Aniao. Sehnem

3092 (GH).

21. Polybotrya hickeyi Moran, sp. nov. (Fig.

40, Map 13).

Polybotrya hickeyi Moran, sp. nov. Type;

Bolivia. Cochabamba: Prov. Chapare, road

from Cochabamba to Villa Tunari, in thick

rain forest, very heavily shaded, on trunk of

Ficus 1700 m, 23 March 1980. Hickey 801

(with Eshbaugh) (holotype: GH!; isotype:

MU).

Caulis hemiepiphyticus. 1-2 cm diam.: squa-

mae alrohrunneae, vulgo opacae. leviter nilidae,

concolores vel marginihus tenuiorihus el pal-

lidioribus ad apicem: lamina usque ad 3-pinnata-

pinnatifida. deltaia, usque ad 1 m longa. leviter

pubens inferne. pilis cylindricis. appressis, usque

ad 0. 1 mm longis: pinnae usque ad 46 ^ 25 cm.

deltalae. paribus proxirrmlibus acroscopice pro-

longatis; pinnulae catadromicae supra basin, del-

tatae. ohlongae vel lanceolatae: costae sine sulcis

prominentibus: venae prominulae superne. apice

leviter incrassatae. Sori discreti. rotundi. oppositi

vel suboppositi.

Stem 1-2 cm thick, hemiepiphytic; scales

dark chocolate brown, mostly opaque, concolor-

ous, or the apex with thinner, lighter borders, the

margins denticulate to entire, the base thickened,

attached across its full width. Sterile leaves up to

1,5 m (?); petiole with thin, ovate to lanceolate

scales; lamirui to 3-pinnate-pinnatifid, deltate. to

about I m long, the abaxial surface slightly pu-

berulent, the hairs cylindrical, appressed, up to

0.1 mm long; pinnae up to 46 x 25 cm, deltate.

the proximal pair basiscopically prolonged, the

distal ones gradually becoming acroscopically pro-

longed; pinnules catadromic above the base, del-

tate to oblong-lanceolate, the basiscopic margins

thickened and decurrent on the costa, the base

slightly prolonged acroscopically, the apices

acuminate; tertiary segments also with a thick-

ened, decurrent, light-colored basiscopic margin;

veins prominulous adaxially. the tips thickened

and ending just behind the margin; costules bor-

dered by a raised, erect flap or wing of tissue;

axes often not prominently grooved, pubescent by

ca. 0. 1 mm long, colorless, erect hairs, the scales

appressed, denticulate, flexuose. Fertile leaves

botryoid; sori with few fusions, arranged mostly

oppositely or suboppositely.

Polybotrya hickeyi grows in high elevation

forests in the Andes of Colombia and Bolivia; it

has not yet been collected from Ecuador or Peru

(Map 13). This species looks most like P.

puberulenta— see that species for comparison.

This plant is named for one of the collectors. Dr.

R. James Hickey. who has made impc>nant con-

tnbutions to the study oflsoetes and Lycopodium.

The Colombian specimens look like the Bo-

livian type specimen except for the stem scales.

The Bolivian type has dark, chocolate brown

scales with denticulate margins and lighter borders

apically (Fig. 40g). The Colombian specimens

have merely dull brown, concolorous scales with



November 1987 Monograph of Poi.ybotra 89

Figure 40. Polyhotrya hickeyi Moran. a. fertile pinnules; b,c,d. sterile pinnules, acroscopic side to

the right; c. medial pinna; f. basal pinna (note that the two proximal basal pinnules arc fertile); g. stem

scales (Bolivia); h. stem scale (Colombia); i. abaxial surface of distal pinnules, a.d: Ludwif; 262 (NY).

b.h; Grant 10283(F). c: Killip & Smith 202/0 (US), c.f.g.i: holotypc. HickeyHOI .wnh Eshbaugh (GH).



90 Illinois Natural History Survey Vol.34, Art. I

entire margins and a prominently curved and thick-

ened base; therefore, they appear appressed (Fig.

40h). This thickened and curved scale base appears

somewhat elevated, like those of P. caudata and

P. cylindrica . The lamina of P. hickeyi turns a

peculiar yellowish green upon drying.

Specimens examined: Colombia Cauca: W
slope of W Cordillera, above Carpinteria. Alston S278

(BM) Cundinamarca: Cordillera Oriental, Toquiza.

Gazaunta Valley, Cordillera de Helicona, 15 km NW
of Medina, 2200 m. Grant 10283 (F, US); Fusagasuga,

Ludwig 262 (BM, MO, NY) Norte de Santander: Pica-

Pica Valley, above Tapata (N of Toledo) 2100-2400

m, KUlip <* Smith 20210 (GH, NY, US).

Bolivia. Cochabamba: Prov. Chapare, road from

Cochabamba to Villa Tunari, 1700 m, Hickey 801 with

Eshbaugh (GH).

22. Polybotrya puberulenta Moran, sp. now

(Fig. 41, Map 13).

Polybotrya puberulenta Moran, sp. nov.

Type: Ecuador. Napo; camino Baeza-Tena,

5 km al sur de Cosanga. Cordillera Oriental,

bosque perturbado, 1 100 m, 4 de enero 1984,

Moran 3528 (holotype; F!; isotypes: COL!,

GH!, LPB!, MO!, NY!, Q!, QCA!, UC!,

US!, VEN!).

Caulis hemiepiphyticus: squamae aurantiacae

sordidae vel luteae, memhranaeae. late patenies.

centroleviterfuscatae. marginibus denticulatis vel

erosis: lamina 2-pinnata-pinnatisecta perlate

ovata, crassa.flavovirenles paltide, pubents ulrin-

que, pilispatentihus.flexuosis, ca. O.l mm longis:

pinnae usque ad 34 x 18 cm, infimus hasiscopice

prolongatae: pinnulae catadromicae oblongae. in

margine incrassatae et decurrentes: lohi tantum

crenali. apice truncati vel rotundati: venae pro-

minulae superne. apice leviter incrassatae. Folia

fertilia 4-pinnata. tuibus dense pubescentibus.

pilis 3-5 mm longis et squamis linearibus. appres-

sis vel leviter patentibus: segmenta opposita vel

subopposita. receptaculo pubescenii , pilis 1-2 mm

longis. articulatis.

Stems 1-2 cm thick, hemiepiphytic; .scales

yellow to sordid orange, darker in the center with

light yellow borders, membranous, spreading, 1-

.3 mm wide, the margins crosc to denticulate, at-

tached across the width of the narrowed base, in

fresh material somewhat rugose transversely.

Sterile leaves up to 1 .2 m long; petiole up to 27

cm long; lamina to 2-pinnate-pinnatisect, broadly

ovate, thick and somewhat fleshy in living mater-

ial, color light yellowish green, puberulent on both

surfaces, the hairs spreading, flexuous, ca. 0.1

mm long, intergrading with the scales on the lower

suri'ace; pinnae up to 34 x 18 cm, the proximal

pairs basiscopically prolonged, the distal ones grad-

ually becoming equilateral or slightly acroscopi-

cally prolonged; pinnules catadromic throughout,

oblong, the basiscopic margin thickened, decur-

rent on the costa. the apices obtuse to acute, base

cuneatc and slightly prolonged acroscopically;

coslules bordered by a raised flap of lighter colored

tissue; lobes of the largest pinnules merely crenate.

the margins thickened, the apices obtuse or trun-

cate; veins prominulous adaxially, the tips slightly

thickened and ending just behind the margin; axes

evenly pubescent abaxially , the hairs 0. 1 mm long,

tawny, erect, scales caducous, usually lacking;

grooves pubescent within, becoming shallow and

flat at the junctures, not admitted to the groove

of the next lower order. Fertile leaves 4-pinnate.

the axes densely pubescent, the hairs jointed. 3-5

mm long, the scales 5-10 mm long, linear, ap-

pressed to slightly spreading; sort opposite or sub-

opposite; sporangial stalks paraphysate; recepta-

cle pubescent by jointed hairs, 1-2 mm long, that

protrude beyond the sporangia; spore size un-

known.

Polybotrya puberulenta grows at high eleva-

tions in the Andes of Ecuador, Bolivia, and prob-

ably Peru (Map 13). The specific epithet refers to

the sterile lamina, which is puberulent on both

surfaces. Another distinctive feature of this species

is the stem scales, which are relatively wide (1-3

mm), thin, spreading and yellowish when viewed

with transmitted light. The lamina is relatively

thick and fleshy; the mesophyll cells apparently

collapse when dry and the veins become promin-

ulous. The lamina, which is always broadest at

the base, has a peculiar light greyish green color

when dry.

This species differs from P. hickeyi in several

characteristics of the stem scales, pinnule arrange-

ment, lamina dissection, pubescence, paraphysis

length, and scalyness of the major axes. Both

species, however, have thickened, decurrent.

basiscopic margins. The medial, less cut pinnules

of P. hickcvi may approach in outline those of P.

puberulenta. but the largest pinnules of the basal



November 1987 Monograph of Polybotra 91

Figure 41. Polybolrya puberulenta Moran. a. pinnules; b. stem scales; c. sterile leaf; d. abaxial

surface of costa and pinnules, a-d: Moran 3528 (F).



92 Ii.r.iNois Natural History Survey Vol. 34. Art. 1

pinnae are much more highly divided in P. hickeyi

(compare Figs. 40f and 41a).

Unfortunately, Rusby's Bolivian collection

consists only of the fertile leaf, but I believe it

belongs to this species because of its cutting, in-

dument type, and yellowish stem scales. The long,

protruding paraphyses are easily seen with a hand

lens and offer an excellent character to distinguish

this species from P. hickeyi.

Specimens examined: Ecuador. Napo: camino

Baeza-Tena, 5 km al sur de Cosanga, Cordillera Orien-

tal, bosque perlurbado, 1 100 m, Moran 3528 (COL, F,

GH, LPB, MO, NY, Q, QCA, UC, US, VEN).

Bolivia. La Paz: Yungas, 6000 ft, "only the fertile

frond collected," Rusby 443 (NY).

23. Polybotrya alata Moran, sp. nov. (Fig. 42,

Map 14),

Polybotrya alata Moran, sp. nov.. Type:

Panama. Panama; Campo Tres, 3 mi NE of

Altos de Pacora, primary forest, alt 500-800

m, epiphytic, the caudex closely appressed,

sterile, 10 March 1973, Croat 22706

(holotype: MO!; isotype: L!).

Caulis hemiepiphyticus usque ad 2.5 cm
diam.i squamae lineares. 15-25 x 0.8-1 .2 mm,
chrysohrunneae, concolores vel centris leviterfus-

catis. appressis, ascendentibus, marginihus den-

ticulatis. Lamina 2-pinnata-pinnatifida, superne

glabra, inferne pubens, pilis circa 0. 1 mm longis,

niveis, suhulatis: pinnae usque ad 30 x 8 cm, an-

guste triangulares, apicibus attenuatis, zonis

mediis alatis secus costam: pinnulae catad-

romicae, usque ad 6x2 cm, lobis catadromicis,

crenatis: rhachis el costae pubescentes et

squamatae. Folia fertilia ignota.

Stem 2.5 cm thick, hemiepiphytic; scales

linear, 15-25x0.8-1.2 mm, membranaceous,

golden brown, concolorous or with a slightly dark-

ened central stripe, appressed-ascending, margins

denticulate. Sterile leaves up to 1.3 m long; petiole

to 45 cm long, scaly, pubescent with 0.2 mm long,

unicellular, colorless hairs; lamina 2-pinnate-

pinnatifid, the adaxial surface glabrous, the ab-

axial surface lightly to moderately pubescent, the

hairs unicellular, less than 0. 1 mm long, whitish,

subulate; pinnae up to 30 x 8 cm, narrowly trian-

gular with a long tapering apex, 3-4 times as long

as broad, the lowest becoming pinnatifid in the

apical one-third, the transition zone between pin-

nate and pinnatifid marked by a narrow, straight,

decurrent, green wing parallel to the costa: pin-

nules catadromically arranged in the medial pin-

nae, lobed almost to the costule, inequilateral at

base, up to 6 x 2 cm, the proximal acroscopic

ones longer than the proximal basiscopic; pinnule

lobes catadromically arranged, the margins cre-

nate, the apices rounded; axes pubescent ab-

axially, the hairs 0. 1-0.2 mm long, subulate, col-

oriess, often difficult to see on the lamina, inter-

mixed on the costae with long, scraggly. subclath-

rate, denticulate scales; groove usually glabrous,

abruptly truncated by the raised ridge of the costal

groove, the junctures short-pubescent. Fertile

leaves unknown.

Polybotrya alata has been collected only

twice, both times in the mountains of Panama

(Map 14), but 1 suspect it also occurs in the moun-

tains of southern Costa Rica. Distinctive features

of this species are the narrowly triangular pinnae

with long attenuate apices and the pinnules that

are lobed almost to the costule. thereby making

the lamina 2-pinnate-pinnatifid (Fig. 42a). The

transition zone from the free pinnules to the pin-

natifid apex differs from all other species of

Polybotrya by the thin, basally decurrent. green

wing, which imparts an alate appearance to the

costa (Fig. 42c). The stem scales are also distinc-

tive by their long length (15-25 x 0.8-1.2 mm)
and shiny, golden brown color.

The short, even pubescence on the abaxial

surface of the leaf separates P. alata from all other

Central American Polybotrya. except P. caudata.

which can also have pubescent leaves. The laminar

hairs of P. caudata. however, are longer ( l-3mm)

than those of P. alata and are 4-10 celled. The

two species also differ sharply by their stem scales.

The only other Central American species with

which P. alata could be confused is P. osmun-

dacea. but that species is easily distinguished by

its anadromic pinnule arrangement and nonalate

costae. Furthermore, the costular groove of P.

alata is truncated by the ridges of the costa; in

contrast. P. osmundacca has uninterrupted, decur-

rent grooves (Fig. 47h).

Specimens examined: Panama Panami: Cam-

po Tres. .1 mi NE of Alios dc Pacora. Croat 2270ft (MO.

I.) Veraguas: ^ mi W of Santa Fc on ro-id past Escucia

Agncola Alto Picdra on Pacific side of divide, 800-1200

m. Croat 23011 (MO).



November 1987 Monograph of Polybotra
93

E
u

^ 1
Ficn^RR 42. Polyho.ry^a alota Moran. a. sterile pinna; b. margin of stem scale; c abaxial surface ofdistal p,nnules (note pubescence and decurrent p.nnule bases); d. stem scale, a-d: CroaTlTm (MO)



94 Illinois Natural Hisiory Slrvky Vol.34. An. 1

24. Polybotrya aequatoriana Moran, sp. nov.

(Fig. 43, Map 15).

Polybotrya aequatoriana Moran, sp. nov.

Typr: Ecuador. Napo: Cordillera Oriental,

camino Baeza-Tena, 34 km a! sur de Baeza,

2100 m, 30 de diciembre 1983, Moran 3512

(holotypc: F!; isotypcs: AAU!, COL!, GH!,

LPB!, MO!, NY!, Q!, QCA!. UCi).

Caulis hemiepiphyticus. 1 .5-3.4(4.0) cm
diam.; squamis sordide aurantiacn-luteis. nitidis,

memhranaceis, imple.xis. lanatis. in marline mi-

nute denticulatis, 1-2(3) X 12-20(25) mm: lamina

lanceolata, usque ad 1.4x0.7 m, 2-pinnata-

pinnatifida, modice vel sparse puberula subter,

pilis appressis, 0. 1-0.3 m longis; pinnae longidel-

tatae. 25-35 x 13-18(20) cm: pinnulae catad-

romicae, acroscopice prolongatae, basiscopice

valde obliquae: lobi integri vel vade crenati:

rhachis et costae pubescentes, pilis rigidis albidis.

sutcis profundis pubescentibus. Pinnulae fertile

s

lineares, vulgo 4-6 x 0.5-1 .0 cm, segmentis op-

positis vel suboppositis circularihus vel oblongis,

3-5 X 1-2 mm.

Stem 1.5-3.5(4) cm thick, hemiepiphytic;

scales dingy orange yellow, mostly darkening

upon drying, shiny, membranous, tangled and

woollike, the margins minutely denticulate, 1-

2(3) X 12-20(25) mm. Sterile leaves up to 1.6 m
long; lamina lanceolate, to 1 .4 x 0.7 m, 2-pinnate-

pinnatifid, moderately to lightly pubescent below,

the hairs fine, appressed, 0. 1-0.3 mm long, grad-

ing into scales along the axes, often with a few

scattered resinous, circular glands (use at least

30 x ), the margins somewhat thickened and

lighter colored, often becoming revolute upon dry-

ing; pinnae long-triangular, 25-35 x 13-18(20)

cm; pinnules arranged catadromically above the

base, prolonged acroscopically, reduced and very

oblique basiscopically; lobes entire or rarely shal-

lowly crenate; axes pubescent below with un-

evenly scattered, whitish, erect hairs 0. 1-0.2 mm
long, scaly by linear, flexuous, darkened, often

appressed scales, these most abundant at the

junctures; grooves deep and di.slinct, filled with

hairs, these sometimes clavatc, ca. 0.1 mm long,

reddish, the ridges well detlned. stramineous, con-

tinuous with those of the costules but truncated

by the ridges of the rhachis. Fertile leaves bot-

ryoid; pinnules linear, mostly 4- 6 x 0.5-1 .0 cm;

sort arranged oppositely or suboppositely, round

to oblong; sporangial stalks paraphysate; spore

size unknown.

Polybotrya aequatoriana occurs in the cloud

forests of Ecuador and Bolivia (Map 15), from

2100-2410 m, the highest known altitudinal range

of any species of Polybotrya. Polybotrya aequa-

toriana is named for Ecuador, where I collected

the type.

Polybotrya aequatoriana can be easily sepa-

rated from P. hickeyi and P. appressa by its stem

scales, which are thin, yellowish orange, and

membranaceous (Fig. 43b); see P. appressa for

further comparison with that species. Polybotrya

aequatoriana has highly reduced basiscopic lobes

on the sterile pinnules (Fig. 43c), and the fertile

pinnules are much narrower than those of other

congenerics.

Specimens examined: EcuAtxsR Napo: Cor-

dillera Oriental, camino Baeza-Tena. 34 km a! sur de

Baeza. 2100 m, Moran 3512 (AAU, COL, F, GH, LPB,

MO, NY. Q, QCA. UC), 3585 (F. Q. QCA); road

Baeza-Lago Agrio. ca 114 km from Lago Agrio. 1750

m. OUgaard et at. 35776 (AAU. QCA).

Bolivia La Paz: Prov Sud Yungas. Huancane,

7 5 km hacia el sud sobre el camino nuevo. 2410 m.

Beck 3108 (LPB); same locality, but at 6 5 km. 2280

m. Beck 3060 (LPB, NY).

25. Polybotrya appressa Moran. sp. nov. (Fig.

44, Map 17).

Polybotrya appressa Moran, sp. nov. Type:

Ecuador. Napo: Cordillera Oriental, camino

Baeza-Tena. 34 km al sur de Baeza. bosque

virgen, 2100 m, 24 de enero 1984. Moran

3586 (holotypc: F!; isotypcs: GH!, MO!,

NY!. Q!. QCA!. US!).

Caulis hemiepiphyticus. 1-2 cm diam.:

squamae concotores, obscure brunneae. integrae.

appressae, vulgo 1-2 x 0. 1-0.2 cm; lamina ovata

vel late deltata. 2-pinnaia-pinnatifida. superne

glabra, inferne pilosa, pilis dispersis. ca. 0.1 mm
longis: pinnae usque ad 37 x 18 cm: pinnulae pro-

longatae acroscopice. latis basiscopicis reductae:

lobis rotundatis vel plusminu.'ive truncatis: rhachis

et costae acqualiier pubescentes, pilis usque ad

0. 1 mm longis. sulcis glabris. Foliafertilia ignota.

Stem 1-2 cm thick, hemiepiphytic; scales

concolorous, dull brown, thick, opaque, entire.

*



November 1987 Monograph of Polybotra 95

FicjURF. 43. Polvboirya cwquatoriana Moran. a. sterile leaf; b. stem scales; c. costa and pinnule bases,

abaxial view (note only scattered hairs); d. rhachis, costa, pinnule bases, adaxial view, groove minutely

pubescent, thin, a-d; Moran 3512 (F).



96 Ii.i.iNois Natural History Survey Vol.34. An. I

»^»lj«'v

Figure 44. Polyhotrya appressa Moran. a. sterile leaf; b. stem scales: c. eosta and pinnule bases,

abaxial view; d. costa and pinnule bases, adaxial view, a-d: Moran 3586 (F).

I



November 1987 Monograph of Polybotra 97

appressed, mostly 1-2x0,1-0.2 cm. Sterile

leaves up to 1.5 m long; lamina ovate to broadly

deltate, dark green above, slightly paler below,

2-pinnate-pinnatirid, up to 1.3x0.74 m, gla-

brous adaxially , abaxially with a few scattered, ap-

pressed hairs, these less than 0. 1 mm long, inter-

grading with scales on the costae; pinnae to 37 x

18 cm, widest at the base, the apex long-tapering,

the acroscopic basal pinule usually prolonged

beyond the rest, pinnules arranged catadromically,

prolonged acroscopically and reduced basiscopi-

cally at the base, the proximal ones with acuminate

apices, the distal ones with falcate and truncate

apices: lobes rounded to somewhat squarish, entire

or rarely minutely serrate at the apex; axes evenly

pubescent abaxially, the hairs short, less than 0.1

mm long, the costal scales present or caducous

and absent, linear to lanceolate, appressed, light

brown, thin; grooves sparsely pubescent within,

not very deep or prominent, bordered by broad,

shallow ridges, not admitted to the groove of the

next lower order. Fertile leaf unknown.

Polyhotrya appressa is known only from the

eastern Andes of Ecuador (Map 17), where it

grows in cloud forests at high elevations. This

new species is named for its distinctive, appressed

stem scales (Fig. 44b), which help separate it from

similar species, many of which have spreading,

membranous stem scales. Polyhotrya caudata has

similiar dull, thick, opaque stem scales, but that

species does not grow at high elevations.

Polyhotrya appressa differs from P. aequa-

toriana. a closely allied Ecuadorian species, by

its stem scales, groove architecture and pubes-

cence, and lamina shape. Polybotyra hickeyi is

also very closely related, especially by its short,

even, abaxial, costal pubescence, and the indis-

tinct adaxial grooves that are glabrous within; but

it differs pnmarily by its more finely cut lamina.

In addition, the lamina of P. appressa is dark

green, in contrast to the pale yellowish green

laminae of P. hickeyi and P. aequatoriana.

Specimens examined: Ecuador. Napo: Cor-

dillera Orienlal, camino Bacza-Tena. .14 km a! sur dc

Baeza, Bosque virgcn, 2100m. 24deenero \9M.Morun
i5«6 (F, GH, MO, NY. Q, QCA, US); Cordillera Orien-

tal, 5 km al sur de Cosanga. camino Bacza-Tena, 1(X)0

m. 4 de enero 1984. Moran iSll (Q, QCA); Cerro

Huacamayos, on road Baeza-Tena, ca. 34 km from

Baeza, 0llgaard ei al. 53824 (AAU, QCA).

26. Polyhotrya altescandens C. Chr. (Fig. 45,

Map 121.

Polyhotrya alte.uandens C. Chr.. Index

Filicum 7. 1905. nam. nov. for Acroslichum

chrysolepis Sodiro, non Fee 1869.

Acrostichum chrysolepis Sodiro, [Anal.

Univ. Quito (Xl)77:56l. 1894.|Crypt. Vase.

Quit. 485. 1893. mm. illeg., non Fee 1869.

Typr: Ecuador. Pichincha: selva tropical, de

Pilaton-Toachi, scptiembre 1892. Sodiro s.n.

(holotypc; PI).

Stem (1)2-3(4) cm thick, hemiepiphytic;

scales golden to yellowish, mostly concolorous,

shiny, tangled and forming a dense "woor" around

the stem, linear. 0.6-1.3 x 15-25(32) mm, base

cordate and darkened at the point of attachment.

Sterile leaves up to 2 m long; lamina to 1 .65 m
long, lanceolate, to 2-pinnate-pinnatifid or rarely

3-pinnatc, usually moderately pubescent, the hairs

fine, appressed, more or less cylindrical, multicel-

lular, 0.2-0.4 mm long, inlcrgrading with scales

on the costa; pinnae evenly tapering to a truncate

base, the acroscopic side slightly prolonged, espe-

cially in the distal pinnae. 14-25(35) x 3-10(17)

cm. soon becoming pinnatifid in the distal por-

tions; pinnules anadromous or subequal proxi-

mally. catadromous distally, entire tocoarsely ser-

rate, the basal acroscopic one almost always con-

spicuously prolonged beyond the others; axes

pubescent abaxially, the hairs subulate, colorless,

0.1-0.2 mm long, unicellular, the scales few,

scattered, denticulate; grooves slightly pubescent

within, those of the costulc dceurrent into those

of the costa, costal groove gradually becoming

shallow and expanded before the juncture with the

rhachidial groove; veins in pinnate groups, in large

pinnae and pinnules the lowermost basiscopic

veinlet often springing directly from the costa or

coslulc. appearing as a single isolated veinlet be-

tween the main pinnate groups. Fertile leaves 3-

pinnate, pubescent by jointed hairs 0.5-1.0 mm
long, these grading into linear, tortuous scales;

sori obovatc or oblong; sporangiul stalks para-

physate; spores (54)59-69(72) microns long.

« = 4 1

.

This species has an altitudinal range of

(8(K)) 1200-2500 m. Most of the collections come

from the western cordillcra of Ecuador, but two

apparently disjunct locations occur in Colombia



98 Illinois Natural History Slrvey Vol. 34. Art. 1

FiuuRi- 45. Polybolrya allescimdens C. Chr. a. stem scales; b. fertile leaf; c. fenile pinnae; d. sterile

pinnule (note isolated veinlet between the main pinnate groups); e. fertile pinnule (note obovate seg-

ments), a.b: Moran 3559 (CR). c.e: Rimbach 91 (F). d: Steyermark 52S12 (F).



November 1987 Monograph of Polybotra 99

and Peru (Map 12). I suspect that this species

occurs in many other valleys of the Colombian

and Peruvian Andes, and that lack of collecting

accounts for the apparent disjunctions. My field

experience in Ecuador showed this species to be

quite common in the western cordillera but absent

from the eastern cordillera.

Polyhotrya allescandens can be immediately

recognized by its bright golden to yellowish scales

that thickly cover the stem. These scales are longer

and narrower than those of any other species of

Pohbotrya (Fig. 45a). The lamina is less cut than

that of similar Andean species, and the pinnae

soon become pinnatifid in their distal portions, a

characteristic that further accentuates this less-

divided look. The sori are usually clavate or short-

oblong, in contrast to the more elongated sori of

other species (Fig. 45e). An unusual tendency of

the venation further distinguishes this species: in

large pinnae and pinnules the lowermost basi-

scopic vein migrates onto the costa or costule and

therefore appears as a single isolated veinlct be-

tween the main pinnate groups (Fig. 45d).

Specimens examined: Colombia Antjoquia:

bosquc bajii la cumbrc cerca dc Boqucron. camino cntre

Medellfn y Palmilas. 2.100-:.'i(X) m. HoJi:cf<?92 (OH I.

Ecuador Azuay: rich dense jungle between

Chacanceo and Rio Blanco, on road to Moileluro. be-

tween Ri'i) Putucay and Ri'o Norcay, 1 220 m.Steyenntirk

52812 (F). Carchi: environs of Maldonado, 1450 m.

Madison el al. 4851 (F). Chimborazo: mounlains in

Sacramento area, H7.i;,i;//i.v J J073 (NY), Cotopaxi: road

between Quevcdo and El Corazon, 6.4 km NW of El

Corazon. 67.5 km SE of Quevcdo, Croal 55844 (MO).

Guayas: valley of Ri'o Chimbo. K(K) m. Rimbuch 91 (!'.

US) Imabahura: in the vicinity of the Rfo Verge, ca.

5 km SW Irom the village of Mani, Rio Cachaco, 1.100

m, Sperling & Bleiweiss 5034 (GH, Q, QCA)

Pichincha: tropical forest de Pilaton-Toachi, September

1892, Sodiro s.n. (P); road El Paraiso-Saguangal. 1

1

km from El Paraiso. 0llnaard et al. i7702 (AAU).

}7867 (AAU); road El Paraiso-Saguangal. 3 km from

El Paraiso. 1500 m. Ollgaartl el al .1782(HAA\Jr. selva

topical. Valle de Nanegal. Sadirii .v./i (P. US); Mindo.

Svdow M9 (US); road from Slo Domingo dc los Co-

lorados to Aloag. 2 5 km F, of Comejo Astorga. I2(X)

m. Moran 3544 (F, GM, Q, QCA); Tinalandia Resort,

ca. 25 km E of Slo Domingo dc los Colorados. N side

of Rio Toachi. 1000 m. Moran 3559 (CR, F. GH. NY.

0, QCA); ca. 30 km WNW of Quito, 67 km on road

to Mmdo. 2200 m, Moran 3564 (F, Q, QCA, US)

Province unknown: western cordillera, 800 m, forest

region, Rimhath 312 (US); without locality, Jameson

33 (P)

Pt Ru Huanuco: La Divisoria, ca. 25 km NE of

Tmgo Maria, Moran 3688 {V. GH, MO, USM). Loreto:

Prov. CoroncI Porlillos, Dtto. Padre Abad , divisoria

cerca al Rio Chino, J. Schunke 10200 (MO).

27. Polybotrya gomezii Moran, sp. nov. (Fig.

46. Map 14).

Pohhotrxa gomezii Moran, sp. nov. Type:

Costa Rica. Alajucla: ca. 20 km N of San

Ramon, at the Univ. of San Ramon's Biolog-

ical Field Station, 1100 m, cloud forest, 17

July 1983, Moran 3241 (holotype: CR!;

isotypes: F!. GH!. MO!, NY!, UC!. US!).

Cciulis hemiepiphylicus. 1-2 cm diam.;

squamae appressae vel ascendentes, brunneae,

obscurae, opacae, anguste lanceolatae, vulgo 10-

20 X 1-2 mm, marginibus vulgo integris; lamina

usque ad 1.5x^0.75 m, ovata vel deltata. ad-

modum glabra. 2-pinnata-pinnalifida. coria-

cea, inferne anadromica. superne catadromica:

pinnae usque ad 45 x 20 cm, anguste deltalae,

apicibus longiacuminatis: pinnulae usque ad 12 x

2.5 cm, acroscopice leviler prolongata. Foliafer-

tilia usque ad 0.75 m, 3-pinnata: sporae (50)54-

60(64) micrometralae.

Stem 1-2 cm thick, hemiepiphytic; scales ap-

pressed to spreading, dull brown, opaque, thick-

ened in the middle, narrowly lanceolate, 10-20 x

1-2 mm, margins entire or with a few apical den-

ticulations. Sterile leaf up to 1.5 m long; lamina

up to 1,2 X 0.75 m, 2-pinnate-pinnatifid or very

rarely 3-pinnate, coriaceous, ovate-triangular,

nearly glabrous, usually anadromic below, becom-

ing catadromic in the distal one-third, the change

from anadromic to catadromic marked by a small,

reduced, basiscopic pinnule or segment; pinnae

up to 45 X 20 cm, narrowly triangular, the apex

long acuminate, the lower ones pinnatifid in the

apical third, giving a broadened appearance to the

pinnae; pmnu/e.? up to 12 x 2.5 cm, apex of prox-

imal ones acuminate to long acuminate, gradually

becoming curved and then truncate in the pin-

natifid apex, the base inequilateral with the acro-

scopic side slightly more developed, the margins

serrate to pinnatifid; axes usually glabrous abax-

ially or with unicellular, fine, whitish hairs, scaly,

the scales caducous, appressed. up to 2 mm long,

light brown, with a long narrow apex; grooves

mostly glabrous within or nearly so, sometimes



100 Illinois Natural History Survey Vol 34. An 1

FiauRF. 46. Polybotrya gomezii Moran. a. acroscopic basal pinnule, pinnae apex toward the left (note

catadromous venation); b, sterile leaf (note that the transition zone from anadromic to catadroniic is

marked by reduced basiscopic pinnules); c. fertile leaf; d. stem scales; e. basal pinna. a,d; Moran J241

(CR). b.e; Moran 2441 (F). c: l.enl i5M (F).



November 1987 MONOGRAPI{ ofPoi YBOIRA 101

pubescent at the junctures, the hairs short, less

than 0. 1 mm long and inconspicuous. Fertile

leaves up to 0.75 m long, 3-pinnate, coenosoric;

axes with scattered scales similar to those on the

sterile leaf, junctures pubescent, the hairs numer-

ous, short, less than 0.2 mm long; sporangial

stalks paraphysate; spores (50)54-60(64) microns

long.

This species is named in honor of Luis D.

Gomez P., pteridologist, formerly at the Museo

Nacional de Costa Rica, who has encouraged my
study oi Polybotrya and extended much hospitality

during my thesis research in Costa Rica. Polybot-

rya gomezii is endemic to Costa Rica (Map 14),

where it occurs from sea level up to 1500 m,

reaching its greatest frequency and abundance in

cloud forests. Most of the specimens were col-

lected in the mountains around San Jos6. I found

P. gomezii at five localities, always with P. al-

fredii; no hybrids were found.

This species can be distinguished from others

in Central America by its nearly glabrous, only

2-pinnate-pinnatifid leaves, and the nearly gla-

brous axes. The pinnae soon become pinnatifid in

the apical half, which also gives a more broad,

less finely cut appearance to the leaf (Fig. 46b).

The opaque, dull brown stem scales with entire

or sparsely denticulate margins (Fig. 46d) also

help separate this species from others found in

Costa Rica. The veinletsof the pinnules are always

arranged catadromically, even in pinnae having

the pinnules arranged anadromically. This order-

ing is unusual because the disposition of the

veinlets usually reflects the pinnule arrangement;

that is, if the pinnules are anadromic, the veinlets

are also anadromic.

Specimens examined: Costa Rica. Alajuela:

ca. 20 km N of San Ramon, at the Univ. of San Ram6n's

Biological Field Station, cloud forest, Moran 3241 (CR,

F, GH. MO. NY, UC, US); forest between Quebrada

Quicuyal and Quebrada Arrayancs, Cariblanco, Lent

3537(F). Cartago: near AltoPalillos, M»ran2-/4/ (CR,

F, GH, MO, NY, US); cloud forest near the entrance

to Parque Nacional Tapanti, 1270 m, Moran 3338 (CR,

F, GH, NY, US); 3 km SE of Tapanti, lower slopes of

Alto Patillos, Lent 1084 (F); La Hondura, 1400 m, Val-

eria s.n.. (CR); Navarro, 1400 m. Werckle s.n. (GH,

P). Heredia: about 10 km on the road towards Virgen

de Socorro, l()(X) m. Moran 3160 (CR) Puntarenas:

Monteverde Cloud Forest Reserve. I.S60 m. Fieiller &
Koplur 51 (UC) San Josi: Parque Nacional Braulio

Carrillo, ca. I km along road from entrance, Moran

3271 (CR).

28. Polybotrya osmundacea Willd. (Figs. 47 &
48, Map 18).

Polybotrya osmundacea Willd., Species PL,

ed. 4, 5;99. 1810. Typf: Venezuela.

Monagas: Caripe, Humboldt 459a (fertile),

^59/) (sterile). Herb. Willd. 19507-1. 19507-

2 (holotypc: B; photos F!, GH!).

Acrostichum osmundaceum (Willd.) Hooker,

Species Filicum 5:246. 1864.

Polybotrya aristeguietae Bradc, Bradea 1:19,

tab. I. 1969. Typf: Venezuela. Miranda:

Santa Teresa-Altogracia de Orituco, June

1953. Aristeguieta 17H0 (holotype: VHN!).

Polybotrya vareschii Brade, Bradea 1 ; 20, tab.

2. 1969. Typi;: Venezuela. Aragua: Rancho

Grande, Dependiente Norte, selva nublada,

800 m, Vareschi & Gessner 1875 (holotype:

VEN!).

Stem 1-2.5(4) cm thick, hemiepiphytic;

scales extremely variable, with plants from

Amazonia tending to have thick, opaque, dark

brown, subentire, somewhat squarrose scales and

plants from elsewhere tending to have shiny, as-

cending, spreading, bicolorous scales, often with

a dark central stripe and lighter borders, margins

highly erose or, less commonly, denticulate,

mostly 1.0-2.2 x 0.»-\ .9i,23) cm. Sterile leaves

up to 1 .8 m long; lamina deltate, ovate, or lanceo-

late, to 3-pinnatc-pinnatifid, chartaceous, rarely

with scattered, round, sessile, resinous glands on

the abaxial surface, the margins glabrous; pinnae

narrowly deltate, equilateral, (7)13-18 pairs; pin-

nules arranged anadromically or rarely catadrom-

ically (Peru), up to 14x4 cm, the largest deeply

cut at the base, apex acuminate; basal acroscopic

tertiary segments ovate, lanceolate or rhombic,

margins entire, crenate or lobed, the apex often

serrulate; axes glabrous or pubescent abaxially,

the hairs whitish. 0.2-0.5 mm long, the scales

absent or few and inconspicuous; grooves dccur-

rcnt into those of the next lower order, usually

filled with short, less than 0.7 mm long, reddish

or brownish hairs. Fertile leaves to 3-pinnatc-

pinnatifid, deltate. coenosoric; sporangial stalks

paraphysate;.\/wr<'.v( 50)54 62(66) microns long.

Other illustrations: See original descriptions

of P. aristeguietae and P. vareschii; Humboldt et

al.. Nov, Gen. Sp. PI., tab. 2. 1815; Proctor,

Ferns of Jamaica, fig. 107. 1985.



102 Illinois Natural History Survey Vol.34. An. 1

T'

(;li*^

id.



November 1987 Monograph of Polybotra 103

Polybotrya osmuiuiaceti , the type species of

the genus, is one of the most widely distributed

species of Polyholrya (Map 18). It occurs in wet,

shaded forests from sea level to 2100 m. The

specific epithet refers to the fancied resemblance

of the fertile leaves to the fertile apex of Osmunda

regafo (Willdenow 1810).

This species varies tremendously in scales,

cutting, and pubescence. In plants from

Amazonia, the scales are usually thick, dark,

opaque, more or less squarrose, and not thickly

investing the stem so that the yellowish stem

aerophores are easily visible in fresh material (Fig.

47b). Plants from other parts of the range may be

similar, but more often the scales are thin, lighter

colored, transparent, often bicolorous, and more

thickly investing the stem (Fig. 47c). The stem

scales, however, do not correlate with any other

characteristics.

Laminar cutting is another variable charac-

teristic. The type specimens of P. aristegiiielae

and P. vareschii. both collected in northern Ven-

FiGURF. 48. Geographical variation in the pinnule cutting oi Polyholrya osmundaiea Willd. The elon-

gated side of the pinnule is always acroscopic. Clockwise, starting from Bolivia: Buchlien 298 (UC);

Killip & Smith 23WI (F); Moran 3642 (F); Moran 36IH (PORT); LeUinner & de la Sola 21 J (COL);

Moran 2167 (CR); H. Smilh 1050 (US); Vareschi & Pannier I6H6 (US); Kdlip & Lasser 37756 (US);

Murillo 2580 (F); Broadway 5589 (F); Steyermark 107148 (MO); Granville 3865 (CAY); Berg el al.

PI8I38 {N\).



104 Illinois Natural History Survey Vol.34. Art. 1

ezucla, scarcely differ from each other in cutting.

These two specimens, however, seem to differ

from P. osmundacea elsewhere in its range by

their less-cut leaves (Fig. 47d; Fig. 48, pinnules

from northern Venezuela). From the greater

number of specimens available to me, it appears

that these extremes of cutting are connected by

intermediates and that no other characters correlate

with these extremes. Accordingly, 1 have placed

P. arisleguietae and P. vareschii in synonymy

with P. osmundacea.

Specimens examined: Guatemala. Izabal: vi-

cinity of Quirigua. Slanclley 24195 (NY, US).

HoNnuRAS. Prov. unknown: near Lake Yojoa,

Sleeves & Ray 504 (GH, US); near Cockscomb Mts.,

Schipp 8101 (US); Maya Mounds, Schipp 8108 (F).

Nicaragua. Comarca del Cabo: 40-50 km SW
of Waspan, AiH'ood 3711 (VT). Zelaya: Cerro la

Pimienta. no. 1 , summit area, P:poly5l46 (CR);costado

S del Cerro La Pimienta, Grijulva 300 (CR).

Costa Rica. Cartago: forests of Las Vueltas.

Tucurrique, Tonduz 13337 (US), 18879 (US); forests of

Tuis, Pittier 12416 (CR, US); Aragon, Pinier 9076 (Z);

Chitaria, Valeria 328 (US); Chitaria, forest near old

jailhouse, Moran 2167 (CR, F); Turrialba, Aragon, Ton-

duz 9006 (US), 9016 (US); vicinity of Pejivalle, Skutch

4637 (CR. US); Peralla, Lankester s.n. (US); vicinity

of Pejivalle, Standley & Valeria 47127 (US); Turrialba,

Instituto Interamericana, Ganaderia, Croat 690 (MO),

738 (MO); Florecia, Turrialba, Jimenez 3299 (F), 3304

(CR, F); Reventazon below Turrialba, Hatch 90 (F);

Turrialba, Pinier 9016 (Z); 3 km W of Turrialba, Mickel

2624 (LP, NY); Turrialba, near the Interamerican Inst.,

Scamman 7152 (GH), 7687 (GH); Turrialba, Unt 299

(GH); valley of Ri'o Reventazon, 3 km SE of Turrialba,

Holm & litis 200 (MICH). Cocos Island: Stewart 241

(US); Fisher s.n. (US); Klawe 1474 (US). 1504 (US).

Lim6n: SW of Siquirres, on road to Turrialba between

Moravia and Guayacin I km of Guayac^, Leilinger &
White 1436(¥, US); near banana plantation and Pandora.

Ri'o Estrella. Rosshach 3628 (GH); Los Diamantes,

USDA Rubber Plant Station, Scamman 5987 (GH). 7153

(GH); hills of Adantic slope, Tonduz 14568 (P); vicinity

of Guipiles, Standley 37096 (US). Puntarenas: Colo,

at sea level, Valeria 333 (F); San Vito, de la Sata 5181

(LP); Finca Loma Linda I mi SW of Canas Gordas.

Croat 22260 (MO); road to Puerto Jimenez. Osa. 40

km W of LA. route 2, Gomez 19489 (CR, UC); Osa

Peninsula, 15 km S of Tropical Science Center field

station and ca. 20 km S of Rincon de Osa, Mickel 2803

(NY). 2817 (NY); Finca Las Cruces, San Vito de Java,

Surch 4613 (MO. NY); ca. 10 mi. SE of RinciSn dc Osa

along road to Pacific, F.vans & Bowers 2792 (MO). San

Jos<: vicinity of El General, Skutch 3018 (F, MO); San

Isidro dc El General. Scamman 5984 (GH).

Panama Chiriquf: 10-1 1 miles W of Puerto Ar-

mucllcs in vicinity of San Bartolo Limitc, Croat 21987

(MO); vicinity of Gualaca ca. 10.7 miles from Planes

de Homito, La Fortuna on road to dam site, Antonio

5133 (MO, UC); 4.5-5.0 km N of dam over Fortuna

Lake. Croat & Grayum 60022 (MO); Fortuna Dam area,

Quabrada Bonito to N of reservoir, Churchill 5776

(MO). Darien: vicinity of airstrip at Cana gold mine,

Croat 38047 (MO). Panama: Cerro Jefe. near summit,

Croat 22682 (MO).

Cuba Oriente: Sierra Maestfa on ascent from Ri'o

Yara, Ekman 14210 (US); Bahi'a de Taco. on Uie ridge

below Santa Maria and Rio Jiguani on the path to the

"Iberia" mines, Ekman 3769 (NY); Trail Navas to Camp
Buena Vista, Shafer 4458 (NY); La Peria. Maurel <t

Maurel 3816 (NY); U Peria, Shafer 8889 (NY), near

Monte Verde. Wright 786 (BM, F, GH, MO. NY, PH,

UC, US); Eggers 5324 (F); Finca Guadalupe, Placetas.

Las Villas, Acuna 17531 (US); Sevilla Estate, near San-

tiago, trail from Magdalena lo Sierra Maestra, Taylor

447 (NY); Loma del Galo and vicinity. Sierra Maestra.

Hioram & Clement 6424 (GH. US); La Prenda. Hioram

2495 (COL. P, US, UC); Santiago, Loma del Gato-

Cobre, Clement 725 (US).

Jamaica. Clarendon: "Second Breakfast Spring
"

below Tweedside, Undenvood 1612 (NY). Portland:

Dollwood, Watt 160 (GH, P, US). 7276 (BM); Mabess

River, below Vinegar Hill, Underwood 1323 (NY); trail

from Vinegar Hill to Mabess River, Underwood 1246

(NY). /250 (NY); Mabess. Fisher 132 (NY); Blue Mts..

Stony River to Macungo River. Marely <& Whitefoord

655 (BM), 675 (BM); Blue Mts , Stony River Base

Camp. Morely& Whitefoord 695 ^UM. MO);JohnCrow

Mts., E slope, 1.5 mi SW of Ecclesdown. Wilson &
Webster 549 (MICH); above Moore Town, Clute 259

(NY, US); valley of the Rio Grande, 8 air miles S of

Port Antonio, Gastany 43 (GH); vicinity of Thomsons

Gap, Maxon & Killip 756 (F. GH, NY, US); Spur of

John Crow Mts. opposite Mill Bank. Maxon 9366 (GH,

NY, PH); vicinity of Mill Bank. Maton <t Killip 148

(F, GH. NY. US); along tfail to waterfall N of Hardwar

Gap. Proctor 16506 (MO). 22269 (GH) St. .\ndrew:

Blue Mts. near Cinchona, 15 km from Kingston. Fisher

s.n. (P); Catherine Peak. Faull 12583 (GH) St. Ann:

Blue Mts.. Trafalgar, Perkins 1163 (GH). St.

Catharine: vicinity of Hollymount, Mount Diablo,

Maxan 2293 (NY. US) St. Thomas:Mansrield and ad-

joining properties, near Bath. Maxon 2470 (NY. US);

Manchester Blue Ml., Day s.n. (N\). forested ridge E

of Cuna Cuna Gap, Maxon 9464 {>i\. US); Bath, Gilbert

s.n. (GH); upper soulhem slopes and summit of Mac-

casucker Bump, Maxon 9522 (GH. NY. PH); SE slopes

of Stone Hole Bump, Maxon 8978 (GH. NY, PH); Com
Puss Gap and vicinity, trail W over Blue Mis.. Wilson

& Murray 564 (BM. GH. MICH); Cora Puss Gap, Proc-

tor 3969 (PH. US).

HAm. Massif du Nord. Chavar>, Ekman 4734

(US); Camp Pcrrin. Ekman 5214 (US)

Grenada. Grand Eiang, Beard 1252 (UC. US);

no localitv , Fraser .v. n . ( P); in Mirabeau Mts. . Broadway



November 1987 MONCXjRAPU of POIYBOIRA 105

2520 (Z); St. Georges, Azimas. 1896, Broadway s.n.

(NY); without locality, Sherring 13 (BM).

TRINIDAI3. Pass, Arima Valley. Fleming <S Fleming

52 (NY); no locality. Fendler 69 (BM, F, GH, MICH,

MO, NY, P, PH, UC); Hart 229 (P); Mount Tocuche,

Brinon et al. 1267 (GH, NY, US); Tacarigua Ward, El

Tocuche, Walker TI0995 (BM); Mome Bleu, Britton el

al. 2276 (GH, NY, US); St. George. Blanchisseuse,

Las Lapas trace, Barnard et al. 411 (BM, MO); Blan-

chisseuse saddle, Richardson 2037 (US); Blanchisseuse

road, 1
1 -mile post, Broadway 6902 (F, MO, Z); Blan-

chisseuse road, 10-mile post, Broadway 5589 (F, MO);

Heights of Anpo. Broadway 9948 (F, NY, US), 9950

(F, GH, NY, US), 9951 (GH, NY. US); Las Lapas

road. Broadway 6459 (BM); Arima-Blanchisseuse

road, 13-mile post, Jermy 1 1 195 (BM), 9-10-mile post,

Jermy 2368 (BM); Mome Bleu ridge, Jermy 2846 (BM);

Arima-Blanchisseuse road, 10-mile post. Fay 373

(BM); Maracas Valley. Las Cuevas trail, FavJ45 (BM);

10.5 mi N of Arima, Crosby 76 (MICH).

French Guiana. Sommettabulaire, zonecentrale,

about 40 km SE of Saul, Granville 3865 (CAY, Z).

Guyana. Region of Mt. Raywa, Jenman s.n.

INY).

Venezuela. Anzoategui: Dtto Bolivar. Fila El

Gacharo, ridge of the fila above Los Chorros and El

Cielo, Serrani'a de Turumiquire, Davidse & Gonzalez

19444 (MO). Apure: Reserva Forestal San Camilo, SW
of caserio San Camilo (El Nula), Sleyermark et al.

101552 (US, VEN). Aragua: selva nublada de Rancho

Grande, Dependiente Norte, Vareschi & Gessner 1875

(VEN); Parque Nacional, Dos Ritos, Killip & Lasser

37756 (US, VEN). Barinas: Dno. Bolivar, along road

from Barinitas to Merida, near one land bridge at San

Isidro. 30 km NW of Barinitas, Moran 3718 (PORT,

VEN); Dtto. Bolivar, San Isidro, ca. 5 mi NW of La

Soledad along Barinas-Sto. Domingo road, A.R. Smith

et al. 1388 (PORT, UC, Z). Distrito Federal: virgin

wet forest on slopes along old road between "Por-

tachuelo" and "Penita" (Petaquire) and Carayaca, be-

tween Colonia Tovar-Junquito road and Hacienda El

Limon, 6-8 mi below junction of Junquito-Colonia

Tovar road, Steyermark & Nevling 95930 (GH, VEN).

Falc6n: Cerro Azul, Wmgfield 6914 (VEN). Falcdn/

Lara: Cerro Socopa, Liesner et al. 8359 (MO, VEN).

Lara/Yaracuy: Dtto Urdaneta y Bolivar, la fila Azul

y Hda. El Jaguar. Ortega & Smith 2387 (PORT); Sierra

de Aroa, 10-13 mi NW of Urachiche (Edo. Yaracuy)

along din road leading NW from Urachiche to Duaca

(Edo. Lara), A.R Smith el al. 1347 (UC, PORT, Z).

Miranda: Santa Teresa-Altogracia de Orituco. Aris-

leguieta 1780 (VEN); Cerros del Bachiller, near E end

of virgin evergreen forest, above Quebrada Corozal, S

of Santa Cruz, 10 km (by air) W of Cupira, Steyermark

& Davidse 116499 (MO, UC, VEN). Monagas: Caripc.

Humboldt 459a.b (P, photo F, GH); Cerro de Giucharo,

of Guicharo, Steyermark 62015 (F). Nueva Rsparta:

Cerro Copcy, Sugden 1151 (UC). Territorio Federal

Amazonas: Sierra Parima, vecinidadesde Simarawochi,

Rio Matacuni, a unos 6-7 km al ocste de la frontera

Venezolana-Brasilcra, Steyermark 107148 (MO, NY.

VEN); Depto. Rio Negro. 0-2 km E of Cerro La Neblina

Base Camp on Rio Mawarinuma. Liesner 16133 (MO.

UC); environs of Neblina Base Camp, Plowman &
Thomas 13678 (F. UC). Sucre: Peninsula de Paria, Dtto.

Mariiio, camino Mundo Nuevo-Manacal. 18-20 km N

de Irapa. Dumonl el al. 7439 (NY, VEN); alrededore

de Manacal, Murillo 2580 (F, NY, VEN); Peninsula de

Paria, Cerro de Humo, NE de Irapa, Steyermark 94952

(F, GH, VEN); Peninsula de Paria, arriba de Mundo

Nuevo, oeste de Cerro de Humo, Steyermark <& Rahe

96145 (GH, VEN); Peninsula de Paria. vicinity of Ma-

nacal 15 km (by air) NW of Irapa. Steyermark & Liesner

120634 (MO, UC, VEN); Peninsula de Paria, arriba de

Mundo Nuevo, oeste de Cerro de Humo, Steyermark &
Rabe 71756 (VEN). Tachira: Dtto. Uribante. along road

from La Siberia to entrance to Las Cuevas Represa, van

der Werff& Gonzalez 5202 (MO, UC). Yaracuy: Dtto.

Bruzual, selva siempreverde, Montaiia de Maria

Lionoza, Quebrada Quibayo, desde abajo hasta casi la

cumbre, Sleyermark et al. 125039 (UC, VEN); Cerro

"Chimborazo," Sierra de Aroa, Vareschi & Pannier

2715 (US, VEN); en la selva que cubrc la fila "La

Enjalma" al sur de Chivacoa, Vareschi & Pannier 2660

(VEN). Zulia: Parija, Vareschi 3 147 {VEH); Dist. Mara,

NW wooded slopes of Cerro Negro, 5.5 km SW of

Rancho 505, ^oiV.\oG\iAf,aie, Sleyermark el al. 122814

(MO, VEN); Dtto. Bolivar, Cuenca del Embalse Burro

Negro (Pueblo Viejo), sector entre Quiros-EI Pensado

y el pie de Cerro Socopo. en el area aprox 10 km en

li'nea recta al este de Chunigauarita. Bunting 9516

(VEN); 15 km de El Vigi'a, carretera Panamericana,

Vare.fchi & Pannier 1686 (US).

Colombia. Antioquia: Rio Leon. Bendix site,

Ca/>i 74 (MICH); carretera al mar cerca de Villa Arleaga,

Gutierrez & Barkley 1 70109 (GH); Municip. Anori, Pro-

videncia, Soejarto2805 (COL) Boyaci: Muzo, Lindsay

262 (BM). Cauca: Agua Clara, along hwy from Buena-

venturea to Cali, Killip <S Cuatrecasas 38902 (F, US);

Costa del PaciTico, Ri'o Micay, brazo Noanamito, orilla

derecha. El Chachajo, Cuatrecasas 14246 (US). Choc6:

0.5-2.5 km N of the INDERENA Camp on Rio Truando

near Caserio La Teresita, Lellinger & de la Sola 553

(COL, US); NW side of Alto del Buey, Lellinger & de

la Sola 213 (COL, US), 250 (COL, LP, US); trail from

Ri'o Mecana to Alto de Mecana, Gentry & Juncosa4l02l

(MO, UC) Magdalena: Sierra Nevada de Santa Marta,

regi6n del Campano, Barkley & Gutierrez V. 1897

(MICH); Santa Marta. near Us Partidas, 3500 ft, H.H

Smith 1050 (B, F, NY, MICH, MO. PH, US, VT);

Forest Boca Toma, El Recuerdo, 2500 ft, Bennett 23

(F); Santa Marta Mts., trail beyond falls. El Recuerdo,

Niemeyer 44 (US). Meta: Villavicencio, Alston 7641

(BM). SanUnder: Batbosa, Henri-Stanislas 1710 (US);

Mesa de los Santos, Killip & Smith 15341 (COL, GH,

NY, US); between Ubrija and San Vicente, ,4/.s/on 7341



106 Illinois Natural History Survey Vol.34. Art. 1

(BM) Prov. unknown: Municip de Marsella, Vereda

La Nona, Finca Palermo, cerca al caserio Caracas, Cor-

dillera Central, vertiente occidental, Idrobo etal. 101 16

(COL).

EcuAiXiR. Los Rfos: Ri'o Palenque Biological Sta-

tion, km marker 56 N of Quevedo, Moran 3600 (Q,

QCA). Napo: 27 km SE de Coca, alrededor de pozo de

petroleo Auca 4, Moran 3618 (PORT, Q, (JCA); 12 km

SW of Coca, por el camino se llama "Los Zorros,"

Moran 3612 (Q, QCA); 73 km NE de Baeza, propiedad

de Inecel, "Cascada de San Rafael," Moran 3592 (F,

MO, Q, (JCA); Puerto Francisco de Orellana (Coca).

17 km SW of the town al road along Ri'o Napo (Los

Zorros), Balslev & Madsen 10649 (AAU, Q, QCA);

Ri'o Napo. Panachocha (Oasis), Hurling el al. 7535 (F,

GH); Anangu. Parque Nacional Yasuni, SEE project

area, 0llgaard et al. 38845 (AAU, Q, QCA), 38894

(AAU, Q, QCA), 39086 (AAU, Q, (JCA). Pastaza:

Lorocachi, zone oeste del campamento militar a 3 km

del Rio Curaray, Jaramillo et al. 30783 (AAU, Q.

QCA). Pichincha: Pululahua, Sodiro s.n. (US); Chim-

borazo. Spruce 5685 (P); Los Colorados, Sodiro s.n.

(P); 15 km E of Sto. Domingo de los Colorados, road

behind Brasilia a Toachi. along RioToachi. Moran J547

(F, Q, QC/K). Santiago-Zamora: Cordillera Cutucij,

ridge just S and W of Rio Itzintza, Camp 1298 (NY),

1359 (NY). Tungurahua: Banos-Jivaria de Pintuc,

Stiibel 875 (B). Prov. unknown: Junganza, Crespi s.n.

(US); San Miguel, Sodiro 81 (UC).

Pkru . Huanuco: SW slope of the Rio LIulla Pichis

watershed, on the ascent of Cerros del Sira. Camp 3

(Laguna), Dudley 13005 (GH); Tingo Marfa, Allard

21609 (US), 21997 (US). Junin: Pichis Trail, Yapas,

Killip & Smith 25452 (NY, US); Chanchamayo Valley,

C. Schunke 164 (F), 705 (F), 134/ (F). 1395 (F), 1451

(F); E of Quimiri bridge, near La Merced, Killip <S

Smith 23991 (F, NY, US). Loreto: Prov. Maynas, Peter

Jensen's Explorama L<.)dge, 50 mi downriver on the

Amazon at Yonamono Ck., Moran 3642 (AMAZ. F,

USM); Altura Tuta Pishco on Rio Napo, Croat 20287

(MO); Gamitanacocha, Rfo Mazan, J . Schunke 275 (F.

GH. NY. UC, US), 380 (F, GH, UC, US); Pumayacu,

between Balsapucrto and Moyobamba, Klug 3208 (F.

GH. MO. US); Veradera de Mazan, Croat 20786 (MO);

17 km SW of Iquilos, Croat 18476 (MO); Rfo Napo

near Entrada dc lslaInayuga.Cr()o/205'/.^(MO). Madre

de Dios: Prov. Manu. Vargas 1 7743 (GH) San Martin:

Camino a Pushurumbo. 7-8 km E del pucnie dc Palo

Blanco. Mariscal Caceres, Tocache Nuevo. J Schunke

5785 (COL. NY. US) Ucayali: Rio Aguaytia above

mouth of Qucbrada Yurac-Yacu, Croat 20851 (MO).

Bolivia. La Paz: Region de Mapiri. Buchiien 298

(NY, UC, US); Mapin. San Carlos. Buchiien 1066 (LP);

Yungas. Mururata. 1839. Penlland s.n. (P).

Brazil Amazonas: Manaus-Caracarai road. km.

148. Berg et al. PI8I38 (F, NY. VEN) Parfi: Scrra

dos Carajis. AMAZ Camp Azul, Sperling et al. 5915

(GH. NY).

29. Polybotrya cyathifolia Fee (Fig. 49, Map
19).

Polybotrya cyathifolia Fee. Mem. Fam.

Foug. 6. (Hist. Acrost.) tab, 2. 1866. Type:

Guadeloupe, "near the house of Mr. Bovie,"

L'Herminier s.n. (hoiotype; P!; other L'Her-

minicr specimens that are probably types are

atGH!, L!, fragment NY!).

Stem 1-2 cm thick, hemiepiphytic; scales

dull, light reddish brown, concolorous, spreading,

the margins subentire to denticulate. Sterile leaves

up to 1.5 m long; lamina to 3-pinnate. ovate to lan-

ceolate, up to 1.2 X 0.8 m. membranaceous, al-

most always with round, sessile, reddish, punctate

g\ar\ds; pinnae up to 40 x 23 cm, narrowly deltate,

pinnatifid to the very apex, stalked 5-10 mm;
pinnules catadromous to anadromous but mostly

subequal, narrowly deltate, stalked 1-3 mm. the

base subequi lateral, slightly prolonged acroscopi-

cally but not oblique or cuneate on the basiscopic

side; tertiary segments serrate to entire, the proxi-

mal ones of the largest pinnules oblong, slightly

gibbous at the base on both the basiscopic and

acroscopic sides; axes abaxially sparsely to mod-

erately pilose with hairs similar to those above;

grooves densely packed with multicellular reddish

or tawny hairs, these 0.3-0.8 mm long and con-

spicuously exserted from the grcwve. evident to

the unaided eye, especially at the pinnae junctures.

Fertile leaves 3-pinnate-pinnatirid. coenosoric:

sporangial stalks paraphysate; spores (47)50-

59(63) microns long.

Other illustrations: See original description:

Plumier. Tr. Foug. Amer. tab. 32. 1705.

Polybotrya cyathifolia is endemic to the Car-

ribcan islands ofGuadeloupe and Martinique (Map

19). Few collections have been made of this fern,

but the original collection by L'Herminier is well

prepared and has many duplicates.

Polybotrya cyathifolia is very much like P.

osmundacea. but it differs by the shape of its

tertiary segments, the resinous punctate glands,

and by the long-pilose hairs on the axes. The best

way to distinguish P. cyathifolia is by the costal

groove hairs that copiously fill and protrude from

the groove, especially at the junctua^s (Fig. 49c).

Polvbotrva osmwuiacca. on the other hand, usu-

ally has very short hairs (0.1-0.2 mm long) in

the groove. An important tendency in lamina cut-

ting is for the basal tertiary segments to be slightly



November 1987 Monograph of Poi ybotra 107

FiGURK 49. Polyholryci cyalhifolia Fee. a. stem scales; b. basal pinna; c. rhachis-costa-costulc

junctures; d. fertile pinnule; e-g. sterile pinnules; h. abaxial view of lamina showing resinous glands,

a.c.h: LHerminier s.n. (P). b,g: L'Herminier s.n (GH). d: collector unknown (GH). c,f: collector

unknown (B).



108 Illinois Natural History Survey Vol. 34. Alt. )

gibbous at its base on both sides (Fig. 49c). This

gives the pinnules a characteristic appearance, al-

though it is somewhat difficult to describe.

Polybotrya osmundacea rarely has these lobules

and, if so, they are developed only on the acro-

scopic side. Most specimens off. cyathifoUa have

the abaxial surface of the lamina covered with

punctate, resinous glands (Fig. 49h); P. osmun-

dacea rarely has such glands.

Specimens examined: Guadeloupe, "near the

house of Mr. ho\ie, " L'Herminier s.n. (P, GH. L, frag-

ment NY).

Martinique. Duss 1503 (NY, US), 3897 (NY),

47/9 (F, US); Hahn s.n. (P); Belanger s.n. (F).

30. Polybotrya latisquamosa Moran, sp. nov.

(Fig. 50, Map 19).

Polybotrya latisquamosa Moran, sp. nov.

Type: Colombia. Meta: Cordillera La Maca-

rena (extremo nordeste), macizo Renjifo, al-

rededores, alt. 1300-19(X) m, 6-20 de enero

1951 , Idrobo & Schultes 1106 (holotype: US!;

isotype: COL?).

Caulis 2 cm diam.; squamae castaneae.

lineares vel anguste lanceolatae, 7-12 x 0.8-2.0

mm; petiolus ad basim squamatus, squamis cas-

taneis. longioribus, 12-16 x 5-6 mm, deltatisvel

late ovatis, marginihus pallidis erosis; lamina late

deltata, 3-pinnata-pinnatifida, glabra in super-

ficiebus utrinque, usque ad 13 .5 x 5 cm. petiolulis

usque ad 8 cm: segmenta tertiaria valde ascenden-

tia basiscopice. Folia fertilia ignota.

Stem 2 cm thick, hemiepiphytic; scales cas-

taneous, linear to narrowly lanceolate, ascending-

spreading, darker and opaque in the center with

lighter borders, margins erose, 7-12x0.8-2.0

mm. Sterile leaves up to 1.5 m (?); petiole scaly

at the base with scales similar to those of the stem

but much larger, 12-16x5-6 mm, deltate to

broadly ovate; lamina broadly dcltatc (?), 3-pinnatc-

pinnatifid, glabrous on both surfaces; pinnae ca.

10-12 free pairs (?), alternate, the largest 45 x 26

cm; pinnules arranged anadromically, the proxi-

mal ones with stalks up to 8 mm long, the base

strongly prolonged acro.scopically, basiscopically

shortened and strongly ascending, up to 13.5 x 5

cm; tertiary segments up to 3.5 x L3 cm, lanceo-

late, cuneatc at the base, pinnatifid, 3-5 free be-

low the pinnatifid apex, the basiscopic side strong-

ly oblique, often with the first proximal 1-3 lobes

completely suppressed; costae glabrous abaxially

or with a few scattered, inconspicuous, less than

0.1 mm long, whitish, subulate hairs; grooves

filled with reddish, inconspicuous hairs less than

0.1 mm long. Fertile /ea/ unknown

.

The type location is a remote area in south-

central Colombia, the Cordillera de Macarena,

separated from the main Andean chain (Map 19).

Since the holotype consists of only a portion of

the stem and several incomplete pinnae, additional

collections of this species would add greatly to

the incomplete description.

Polybotrya latisquamosa resembles P. osmun-

dacea because of its large decompound leaves with

anadromically arranged pinnules but differs by its

exceedingly large, broad, castaneous scales at the

base of the petiole (Fig. 50d); thus the specific

epithet. The stem scales differ from those of P.

osmundacea by their deep castaneous color The

holotype is larger and more finely cut than the

average P. osmundacea specimen, and the basi-

scopic sides of the pinnules and tertiary segments

are extremely reduced and oblique-ascending (Fig.

50a). Finally, the major axes of P. latisquamosa

are glabrous (Fig. 50e), unlike those of P. osmun-

dacea, which are often pubescent.

31. Polybotrya sessilisora Moran, sp. nov. (Fig.

51, Map 20).

Polybotrya sessilisora Moran. Type: Colom-

bia. Vaupes: Rio Vaupcs, Mitii y alrededores,

250 m, 8 September 1951, Schultes & Ca-

brera 13963 (holotype: US!; isotypes: COL!.

GH!).

Caulis hemiepiphyticus. 5-10 mm diam.:

squamae appressae, ascendentes, lineares, 8-12

mm longae. atrocastaneae . marginibus pallidis et

valde vehementer denticulatis: lamina usque ad

3-pinnata-pinnatifuia. glabra, coriacea. pallide

viridis: pinnae anguste deltatae vulgo 17-36 x 7-

16 cm: pinnulae vulgo 3.5-10 x 1 .2-3.0 cm,

anadromicae. apicibus persaepe obtusis: rhachis

et costae sparsim pubescentes. squamis fuscatis.

appressis. Pe.xuosis: venae conspicuae et pro-

minulae. Folia fertilia usque ad 3-pinnata:

lamina anguste alata viridis. marginibus leviter

incras.satis: sori ca. I mm longi. discreti. circu-

lares.



November 1987 Monograph of Polybotra 109

5 cm

E
u

FrouRE 50. Polybotrya latisquamosa Moran. a. sterile pinnule; b. apex of sterile leaf; c. stem scale;

d. scale from petiole base; e. pinnule from medial pinna, a-c: Idroho & Schulies 1106 (US).



110 Illinois Natural History Survey Vol.34, Art. 1

15 cm

FiciURF. 5 1 . Polybotrya sessilisora Moran. a. distal half of sterile leaf; b. ahaxial surface of costa and

pinnules (note the thickened, lighter colored margin and dark tortuous scales); c. fertile pinnule, abaxial

view (note the thickened margins and discrete son); d. stem showing straight, appressed scales;

e. medial pinnules of basal pinna, acroscopic side is up. a,c,d,c: Schultcs & Cabrera 1396.^ (COL),

b: Prance et al. I5.1J2 (NY).



November 1987 Monograph of Polybotra 111

Stems 5-10 mm thick, hemicpiphytic; scales

appressed, ascending, linear, 8-12 mm long, dark

castaneous with lighter narrow borders and

strongly denticulate margins. Sterile leaves up to

75 cm long; petiole V2 to as long as the lamina,

scaly with scales similar to those of the stem, but

more tortuous and spreading, the base mostly cor-

date; lamina to 40 cm long, up to 3-pinnate-

pinnatifid, narrowly deltate, the tissue glabrous,

coriaceous, light green, the margins glabrous; pm-

nae narrowly deltate, 17-36 x 7-1 6 cm, the prox-

imal acroscopic pinnule or segment slightly pro-

longed above the re^l; pinnules to 3.5-10 x 1 .2-3

cm, arranged anadromically, the basiscopic mar-

gin thickened and decurrent on the costa, the apex

of the less cut medial pinnules merely acute or

obtuse; veins conspicuous and prominulous abax-

ially; wees moderately to sparsely pubescent abaxi-

ally, the hairs colorless, tawny, 0.5-1 .0 mm long;

grooves of axes decurrent on those of lower order,

not interrupted, pubescent within by tiny, less than

0.2 mm long, reddish, jointed hairs, covered with

dark, spreading, tortuous, denticulate scales like

those of the petiole. Fertile leaves 3-pinnate, bot-

ryoid, the margins slightly thickened; sort sessile,

round, about 1 mm long; spores (44)46-52(56)

microns long.

Polybotrya sessilisora grows in lowland

forests of the northern Amazon basin (Map 20).

It is probably more common than the number of

collections suggests, since the northern Amazon

basin is poorly collected. I suspect that this species

also occurs in the adjacent Guiana Highlands.

TTie specific epithet refers to the botryoid son

that are sessile instead of short-stalked as in other

species of Polybotrya. Moreover, the sori are em-

bedded in the lamina, which is not completely

reduced to the axis (Fig. 51c). As evidenced by

outgroup comparison to other dryopteroid ferns,

the ancestor to Polybotrya surely had fertile leaves

with discrete, round sori from which the lamina

was reduced. Because the lamina of P. sessilisora

is not fully reduced, I interpret its fertile leaf as

the most primitive in the genus. No other species

in the genus has this distinct kind of fertile leaf.

Another distinctive feature of P. sessilisora

is the dark castaneous scales that contrast sharply

with the light green lamina. The scales of the axes

are spreading and tortuous, whereas those of the

stem are appressed and straighter (Fig. 51b.d).

These scales become smaller and narrower in the

distal parts of the lamina until they become uni-

seriate, appressed hairs. The lamina is always

broadest at the base, in contrast to the closely

related P. osmundacea. which is usually reduced

at the base. Pol\botr\a osmundacea also tends to

be much more highly dissected when leaves of

equal size are compared.

Specimens examined: Colombia Vaupes: Rio

Vaupcs. Mitu y alrededores, 250 m. Schulles & Cabrera

tJ96Jt (COL, GH, US).

Bra/.u . Amazonas: Tapuruquara, beside road to

airpon. Prance et at. 15332 (NY); Reserva Experimen-

tal, km 6(), Manaus-Caracarai road. Conant 1016 (GH),

I4H2 (GH); Reserva Ducke, km 26, Manaus-Itacoatiara

road, Cormni 1080 (GH).

Brazil-Guyana Boundary; Akarai Moun-

tains, height of land between drainage of Rio Mapuera

(Trombelas tributary) and Shodikar Creek (Essequibo

tnbulary), dense forest 600-800 m, A.C. Smith 2984

(GH, NY).

32. Polybotrya canaliculata Klotzsch (Fig. 52,

Map 19).

Polybotrya canaliculata Klotzsch, Linnaea

20:429. 1847. Lf.ctotypf.: Venezuela. Ar-

agua: Colonia Tovar, 1846, Moritz 27H (lec-

totype: B!; isolectotypes: F!, GH!. L!. NY!,

P!, US!; photo GH! of L). Lectoparatype:

Colonia Tovar, 1846, Karsten (Coll. II) no.

13 (B!; isolectoparatypes; BM!, L!; photo of

L specimen, NY!).

Acrostichum canaliculatum (Klotzsch) Hook-

er, Species Filicum 5:247. 1864.

Stems 0.5-2 cm thick, hemicpiphytic; scales

ascending with spreading tips, 12-20x0.7-1.2

mm, dull brown to dark purple brown, concolor-

ous or rarely with a very narrow hyaline border,

the margins entire to denticulate, opaque to

slightly thickened at the center and base. Sterile

leaves up to 1 .8 m long; petiole up to 45 cm long,

shorter than the lamina, scaly at the base or

throughout, the scales becoming progressively

smaller and more ovatc-lanccolate and erosc-

dcnticulatc upwards, with a blackened basal point

of attachment; lamina glabrous, lanceolate to

triangular or subtriangular, 50-80 cm wide, to

(4)3-pinnate-pinnatifid. subcoriaceous. the abax-

ial surface lighter in color than the adaxial; ba.wl



112 Illinois Natural History Survey Vol.34. Art. I

FiouRF. 52. Polyhotrya camiliculata Klotzsch. a. fertile leaf; b. abaxial surface of costa and pinnule:
e. adaxial view of fertile pinnule (the main axis is the costa— same scale as b); d. stem scales: e-f. pin-
nules: g. basal pinna: h. medial pinna, from same leaf as in g. a-d: Morii: 27S (B). e.f: Kursten I3{B).
g.h: v,w der Werjf 3486 (MO).



November 1987 MONCXiRAPH of POLYBOTRA 113

pinnae triangular to subtriangular, up to 40 x 28

em. the lowermost pinnate throughout except pin-

natifid at the extreme apex; pinnules arranged

catadromically or anadromically. up to 16 x 6.5

cm, triangular-lanceolate, the acroscopic side usu-

ally prolonged; coslules adaxially deeply grooved

and bordered by a raised, erect to spreading tlap

of tissue; tertiary segments entire to pinnatifid

(deeply so in large leaves), equilateral at the base

with a rounded to acute apex; axes scaly, the scales

scattered, flaccid, membranaceous, ovate to lan-

ceolate, denticulate, 1-3.5 mm long, the abaxial

surface usually glabrous; grooves pubescent

within, and especially at segment junctures, the

hairs tiny, less than 0.2 mm long, reddish. Fertile

leaves botryoid, 0.8 x 0.6 m, 4-pinnate; axes with

numerous septate hairs that often grade into narrow

scales; pinnae up to 30 cm long, subtriangular;

son round to obovate, distinctly stalked, the stalk

about 1 mm long; spores (50)55-66(70) microns

long.

This species grows in cloud forests in the

Andes of northern Venezuela from 1200 to 1600

m (Map 19). Van der Werff and Smith (1980)

state that this species can be abundant locally, but

that the plants rarely bear fertile leaves.

The sf>ecific epithet refers to the raised flap

af tissue that borders the costules and accentuates

the depth of the central groove. This character is

not diagnostic, since other decompound Polyhot-

rya species also have raised costular ridges, al-

though in P. canaliculata these ridges tend to be

relatively taller and more well developed.

When available, the botryoid fertile leaves

[Fig. 52a,c) distinguish this species from P. os-

mimdacea and its allies. Distinguishing vegetative

-haracters arc the dull brown, concolorous stem

scales, and the major axes with prominent, ovate

to lanceolate, flaccid scales (Fig. 52b). The pin-

fiule arrangement of P. canaliculata may be either

anadromic or catadromic. This variation is pecu-

liar because the pinnule arrangement is usually

jonslant within most species of Polyhotrya. A
similarly variable species in this respect is P.

^omezii, an endemic from Costa Rica. I interpret

P. canaliculata as a primitive species of Polyhot-

n,a because of its decompound lamina and bot-

ryoid fertile leaf.

I have chosen the Moritz collection as the

lectotype because of its wide distribution in her-

baria and the excellent quality of the specimens.

Specimens examined: Vknk/uki.a. Aragua:

Colonia Tovar, 1846, Moriiz 278 (B, BM. F-, GH, L,

NY. P. US); Colonia Tovar. 1846. Karsten 13 (B. BM.

L; photo of L specimen. NY): Parque Nacional Henri

Pittier. bosque de Rancho Grande. Tschudi 166 (VEN);

Colonia Tovar, 1854-5, Fendler 262 (GH, MO, NY,

P, PH. US) Falc6n: Sierra de San Luis, arriba de Sta.

Maria, alt. I2(X) m. 5 June 1979, van der Werff 3486

(MO. UC); Sierra de San Luis, selva nublada. entre La

Chapa y Una, Steyermark 99185 (VEN). Yaracuy:

Dito. Bolivar, entre las Parchitas. Tierra Fria y Ojo de

Agua. Onega & Smith 2498 (PORT). 2511 (PORT).

State unknown: Andes of Venezuela, 1889, Coebel

s.n. (P).

33. Polybotrya semipinnata Fee (Fig. 53, Map
18).

Polybotrya semipinnata Fee, Crypt. Vase.

Bresil. 1:16. 1869. Type: Brazil. Rio de

Janeiro: Yacuacanga, 15 June 1869, Glaziou

2427 (K, P!, RB!, US!; photo of K specimen

at US!).

Aspidium scandens Raddi, Plant. Brasil.

1:34, tab. 49. 1825. Type: Brazil. Raddi s.n.

(FI; isotype: K, photo GH!).

Polyhotrya scandens (Raddi) Christ, Bull.

Herb. Boissier, IF 4:965. 1904. nom. illegil.,

non F6e 1852.

Stem 1-1.5 cm thick, hemiepiphytic; scales

membranous, spreading, mostly 9-12 x 0.5-1.2

mm, bright castaneous, concolorous or with a dark

central stripe and lighter borders, the margins

denticulate to strongly erose. Sterile leaves up to

1 m long; petiole 'A to 'A the length of the lamina;

lamina ovate to lanceolate to 3-pinnate but mostly

2-pinnate-pinnatifid throughout, generally 45-

60(75) X 30-55(64) cm, coriaceous, glabrous ex-

cept on major axes; pinnae broadly triangular to

ovate , mostly l5-30x Il-I7cm, divided at base

and soon becoming pinnatifid distally; pinnules

5-10(13) X 2-3 cm, lanceolate, anadromic

throughout, the bases mostly cuneate, unequal,

the basiscopic side more narrowly cuneate and the

acroscopic side slightly prolonged, proximal ones

with a 3-5 mm long stalk, rarely with the lower-

most acroscopic segment cut to the costule; axes

pubescent abaxially with hairs less than 0.1 mm
long, stiff, colorless, subulate, the scales few or

absent; grtmves glabrous or nearly so within, decur-

rent on those of the next lower order. Fertile leaves

smaller than the sterile, 3-pinnate (-pinnatifid),

cocnosoric; tertiary segments oblong, generally

3-7 mm long; sporangial stalk paraphysate;

spores (48)50-62(66) microns long.



Iii.iNOis Na ruRAi. History Survey Vol.34. Art. I

Figure 53. Polyholrya scmipiiiiuiui Fee. a. sterile pinnules; b. sterile leaf; c middle pinnae of fertile

leaf; d. stem scales, a-c: BraJc S()6I (PH). d: Diuirlc ct al. 65322 {¥).



November 1987 Monograph of Polybotra lis

Other illustrations: Raddi, Plant. Brasil. tab.

49. 1825; Mettenius, Filices Hon. Lips., 23, tab.

2, figs. 1-6. 1856 (as P. acuminata); Brade,

Bradea 1:64, fig. 1; 67, fig. 9. 1971 (as P. scan-

dens).

Polybotrya semipinnata is one of five

Polybotrya species that are endemic to the coastal

mountains of southeastern Brazil (Map 18, Table

2), a distribution that emphasizes the biogeo-

graphic distinctness of the Serra do Mar Moun-

tains. The altitudinal range of P. semipinnata is

from 800 to 1000 m.

This plant, easily recognized by its lamina

cutting, is not readily confused with other species

of Polybotrya. The pinnule bases are distinctive

because they are stalked and more narrowly

cuneate on the basiscopic side. The pinnae are

stouter and much less divided compared to those

of other decompound Polybotrya species; the dis-

tal portions soon become pinnatifid, a characteris-

tic that accentuates this less-cut appearance.

An earlier published name, P. acuminata

Link, has often been applied to this species. Since

the type specimen cannot be located and I cannot

discern from Link's description the species he had

in mind, 1 treat P. acuminata as a name of uncer-

tain application.

Specimens examined: Brazil Rio De Janeiro:

Yacuacanga, Glaziou 2427 (P, RB, US; photo of K

specimen at US); Estrada Veiha da Bocaina, Teresopolis,

Duarle el al. 65322 (F, LP); Serra dos Orgaos,

Teres6polis can. Quebrafrasco, 1000 m, Brade 16456

(RB). Sao Paulo: Piruhyba. Loefyren & Duvall 36217

(RB); propc Rio Grande ad Sao Paulo Railway, 800 m,

Wett.':lein & Schiffner .<^.n. (P); Alto da Serra, Lueder-

waldl s.n. (BM, NY), 21547 (NY); Iguape. morro das

Pedras, Semnha Peroupara, Brade 8061 (NY, PH, UC,

US); Pilar, Gerdes 102 (NY, UC). SU. Catarina: with-

out locality, Schwacke s.n. (P).

34. Polybotrya speciosa Schott (Fig. 54, Map
21).

Polybotrya speciosa Schott, Genera Filicum

tab. 7. 1834. NEOTYlfE: (here chosen) Schott,

Genera Filicum tab. 7, based on material from

"Brasiliae provincia Scbastianopolitana."

Polybotrya tomentosa Brade, Arq. Inst. Biol.

Veg. Rio de Janeiro 1:224, fig. 2, plates 2

and 3. 1935. Type: Brazil. Minas Gerais:

Serra do Itatiaia Maromba, 25 June 1930,

Brade 1035 1 (holotypc: RB!),

Polybotrya osmundacea Willd. var. cris-

popaleacea Rosenst., FeddesRepert. 21:349.

1925. Type: Brazil. Sao Paulo: Alto da Serra,

11 February 1925, Brade 5838 (S; isotype:

UC!).

Polybotrya litoralis Brade, Bradea 1:26, tab.

1 , fig. 2. 1969. Type: Brazil. Rio de Janeiro:

Angra dos Reis, Serra do Mar, 29 June 1935,

Brade 14943 (holotype: RB!).

Polybotrya rosenstockiana Brade, Bradea

1:27, tab. 1, fig. 3. 1969. Type: Brazil. Rio

de Janeiro: Serra dos Orgaos, Corrego Beija-

flor, Brade 16579 (holotype: RB!; isotype:

LP!).

Stem 1-3 cm thick; scales commonly brick

red, rarely dull brown, concolorous or with a dark

central stripe, generally 8-20(27) x 0.5-1 .5(2.0)

mm, spreading, membranous, the margins den-

ticulate to strongly erose. Sterile leaves up to 1.4

m long; petiole 'Ao- Vi the length of the lamina;

lamina to 1.2x0.8 m, lanceolate to ovate, to

3-pinnate but mostly 2-pinnate-pinnatifid through-

out, tomentose to glabrous, the margins sparsely

ciliate to glabrous; pinnae to 40 x 15 cm, free

pinnules usually 5-9; pinnules acroscopic, short

to long triangular, mostly 4.5-8.0(10.0) x 1.5-

3.5 cm, the base stalked, the stalk 2-4 mm long,

the acroscopic side prolonged, the basiscopic side

oblique; tertiary segments generally oblong, the

margins entire to crenulate or dentate; a.xes tomen-

tose to glabrous or subglabrous, the hairs usually

0.1-0.5 mm long, the scales few, appressed, tor-

tuous, narrow; grooves pubescent within, the hairs

reddish. Fertile leaves coet)osor\c , 3-pinnate; jrpo-

rangial stalks paraphysate; receptacle glabrous or

with multicellular, branched paraphyses, these as

long as or slightly longer than the sporangia;

spores (56)60-75(82) microns long.

Other illustrations: See original descriptions

cited above; Brade, Bradea, tab. 3, fig. 4 (as P.

littoralis); tab. 4, fig. 1 (as P. rosenstockiana);

tab. 4, fig. 3; tab. 6, figs. 12, 13 & 15. 1971.

Polybotrya speciosa is endemic to the Serra

do Mar Mountains along the coast of southeastern

Brazil (Map 21). It differs from the four other

species of Polybotrya there by its combination of

strongly denticulate, red stem scales, finely cut

lamina, and multicellular, branched paraphyses

(Fig. 54). The pubescence of the abaxial surface

varies from densely tomentose to nearly glabrous.



116 Illinois Natural History Survey Vol 34. Art. 1 1

Figure 54 Pohbotna speciosa Schott. a. stenle and fertile leaves; b. branehed paraphyscs; c. pin-

nules; d.e. stem scales, a.b.e; Brade 16579 (RB). c.d; BroJe 1035
1
(RB).



November 1987 Monograph of Polyboira 117

Five of the specimens examined had the adaxial

surface slightly pubescent whereas the remaining

were completely glabrous. No other species of

Polybotrya, except P. pilosa, has branched para-

physes.

The spore size varies greatly, perhaps reflect-

ing different ploidy levels. The averages from

eight collections, with 15 spores measured from

each, are as follows (in microns): 56. 57, 57, 58,

69, 69, 79, 82. This problem needs further study

of additional collections andcytological samples.

I have been unable to locate Schott's type,

but I feel quite certain that Schott's excellent plate

represents this species. Schott's specimens be-

came part of the Cardinal Hynald herbarium now
located in Budapest, Hungary (BP). The curator

of the fern collection at Budapest, Mr. Tibor Szer-

dahelyi, informed me (in litt.) that much of

Schott's type material had been destroyed during

World War II and that he could not find the type.

My placement of the three species described

by Brade in synonymy with P. speciosa requires

comment. I find no differences between the types

of P. tomentosa and P. liloralis— the two might

well have been collected from the same individual.

Given this likeness, I find it odd that Brade ( 1 969c)

did not mention P. tomentosa in his discussion

after the description of P. litoralis. I also place

P. rosenstockiana and P. osmundacea var. cris-

popaleacea in synonymy, although they differ

slightly from most specimens of P. speciosa by

their somewhat broader stem scales (Fig. 54, com-

pare d & e). They also differ by their leaves,

which are less pubescent, primarily so along the

axes and veins. Both these characters, however,

intergrade and do not correlate with any others.

Specimens examined: Brazil. Rio de Janeiro:

Serra dos Orgaos, Corrego Beijaflor, Brade 16579 (LP,

RB); Serra do Taquaral, Brade 17464 (MO. NY);There-

zopolis. Brade 977.? (BM. UC). 9M3 (NY); Parque

Nacional de Serra dos Orgaos. de la Sola 234i (LP);

Serra dos Orgaos. Morro Assu. Luelzelhurg 6H5H (US);

Organ Mts . Rose & Russell 20790 (US); Corcavado,

collected by the U.S. South Pacific Exploring Expedi-

tion. 1838-42 (NY. US); Mt. Tijuca. Cuyler4796{\}S):

Angra dos Reis. Serra do Mar. Brade 14943 (RB);

Guanabara. Estrada do Sumaro, Pahsl el al 6772 (LP).

Minas (ierais: Serra do Itatiaia. Maromba. Brade 1035

1

(RB); Itatiaia. Maromba. Brade 20214 (F. LP. MO.
NY); same locality. Brade 12616 (BM); Tijuca. Alston

,S99/(BMl Sao Paulo: Alto da Serra. flra<yf5«JS(UC).

State unknown: "Brazil," Webb 26 (GH).

35. Polybotrya pilosa Brade (Fig. 55, Map 20).

Polybotrya pilosa Brade, Bradea 1:27, tab.

1, fig. 4. 1969. Type: Brazil. Rio de Janeiro:

Teresopolis, Varesea, 1000 m, 27 October

1929, Brade 9787 (holotype: RB!; isotypes:

NY!, UC!).

Differing from P. tomentosa only by charac-

teristics of the pubescence; hairs 1-2(2.5) mm
long, pilose, acicular, pluricellular. whitish or

tawny, disposed along the abaxial surface of the

veins and major axes, rarely on the intervening

laminar tissue between the veins, al.so occurring

on the adaxial surface and protruding from the

costal and rhachidial grooves. Paraphyses present,

of both branched and unbranched types.

Polybotrya pilosa is endemic to southeastern

Brazil (Map 20). This species may be subject to

reinterpretation in the future, as it differs from P.

speciosa only by its pubescence, which shows no

intergradation between the two species. The hairs

of P. speciosa are less than 1 mm long, somewhat

tortuous, and occur on the veins and intervening

laminar tissue. The hairs of P. pilosa are more

than 1 mm long, straightish, and acicular; they

occur only along the veins, not on the intervening

laminar tissue (Fig. 55a). These differences are

readily apparent to the unaided eye. The rein-

terpretation of P . pilosa will require fieldwork

designed to study intra- and interpopulational vari-

ation in southeastern Brazil.

Other illustrations: See Brade's original de-

scription, cited above.

Specimens examined: Brazu,. Rio de Janeiro:

"near Rio de Janeiro," collected by U.S. Exploring Ex-

pedition. 1838-42 (US); Tijuca. Brade 20733 (NY.

UC); "Rio Janeiro." 1851. Andersjon s.n. (LP); Organ

Mts.. Wagner s.n. (GH); Teresopolis. Varesea, Brade

9787 (NY. RB. UC); Serra Estrela. Weddell 952 (P);

"Bra.silia." Riedel HI (GH).



118 Ilunois Natural History Survey Vol.34, Art!

FiciiiRH 55. Polxholna pilosit Bradc. a. costa and pinnule with characteristic long, acicular hairs;

b. sterile leaf; c. stem scales, a: Wagner s.n. (GH). b.c; hnide 97S7 (RB).



November 1987 Monograph of Polybotra 119

Names of Uncertain Application

Pohhotrya acuminata Link, Hort. Bcrol. 2:135.

1833. Psomiocarpa acuminata (Link) PrcsL

Epim. Bot. 162. 1849. Type: Brazil. Collector?

1 have been unable to locate the type and cannot

discern from the original description the taxon to

which this name appMcs. The type specimen may

have been lost during World War II; 1 have, how-

ever. Link's other type specimens from Berlin.

Mettenius (1856, tab. 2) has an excellent illustra-

tion of P. semipinnala. which he refers to as P.

acuminata, but 1 do not know whether he saw

type material.

Polybotnafulvostrigosa Chnsl. Bull. Herb. Bois-

sier, II. 1:70. 1901. Type: Peru. Loreto: Cerro

de Canchahuaya, Huher 1448. 1 cannot find the

type and am uncertain from Christ's description

to what species this name belongs.

Pohhotna lomarioides Mettenius, Filices Lech-

ler. 2:5. 1858. Type: Peru. Puno: San Gavan,

Lechler s.n.. I have not seen the type and cannot

be certain from the description if this name applies

to a Polybotrya species or to another genus.

Polybotrya nutans Kunze, Linnaea 9:24. 1834.

Type: Peru. "Sylvae flor. Peruv. ad Pampayaco

in cortice arborum vetustarum parasitica, Jul.

1829," Poeppif; s.n. (B!. PI). The fertile leaf of

the type specimen came from a species of Polybot-

rya. but I do not know which one. The sterile leaf

of the type specimen is a tree fern, perhaps a

species of Trichipteris.

Polybotrya scandens Fee, Genera Filicum 47.

1852. Type: Venezuela. Lagunetta, Galeotti. I

have not seen the type and cannot ascertain from

Fee's description the species to which this name

applies. 1 suspect it is P. osmundacea. Windisch

( 1982) reported Fee's specimen cii P. scandens to

be at the Botanical Gardens in Rio de Janeiro

(RB). Although 1 received other Fee specimens

from RB, P. scandens was not among them. Fee

gives the location as "Lagunetta, Mexico," but

this site is certainly an error: see the discussion

under P. serratifolia

.

Polybotrya trapezoides Link, Filicum Species

164. 1841. Type: Cultivated plant at Beriin of

unknown origin (B!). I received a specimen from

Beriin (B) labelled as species but cannot be sure

that this is the type. This specimen, however, is

certainly not a species of Polybotrya. Because it

is so young, I am uncertain what genus it repre-

sents.



120 Illinois Natural History Survey Vol.34. Art. 1

Excluded Taxa

Excluded Subgenera

Polyholrya subg. Ectoneura Fee, Mem. Fam.

Foug. (Hist. Acrost.) 75. 1845 {nomen illegit.).

= Bolbitis (fide Hcnnipman 1977).

Polyhotrya subg. Egenolfia Fee, Mem. Fam.

Foug. (Hist. Acrost.) 14. 1845. = Bolbitis (fide

Hennipman 1977).

Excluded Sections

Polyhotrya sect. Arthrobotrya v.A.v.R., Handb.

725. 1908. — Lomariopsis sect. Polyseriatae

(fide Holttum 1978).

Polyhotrya sect. Egenolfia Diels, E&P Nat. Pf].

Fam. 1:195. 19(X). = Bolhitis (fide Hennipman

1977).

Polyhotrya sect. Lomagramma Kuhn, Ann. Mus.

Bot. Ludg.-Bat. 4:198. 1899. = Lomagramma

(fide Holttum 1978).

Polyhotrya sect. Teratophyllum Christ, Famkr.

Erde 42. 1897. = Lomariopsis sect. Poly-

seriatae (fide Holttum 1978).

Excluded Species

Polyhotrya acrostichoides Kuhn, Fil. Afr. 52.

1868. = Bolhitis acrostichoides (Sw.) Ching

(fide Hennipman 1977).

Polyhotrya apiifolia Kunze, Famkr. 1:142, tab.

62. 1844. = Psomiocarpa apiifolia (Kunze)

Presl.

Polyhotrya appendiculata (Willd.) J. Smith, Jour.

Bot. 4:150. 1841. = Bolhitis appemiicuUita

(Willd.) Iwatsuki, (fide Hennipman 1977).

Polyhotrya arfakensis Gibbs, Arfak 71. 1917.

= Alsophila hiformis Rosenstock (fide Holttum

1963).

Polyhotrya articulata Fee, Mem, Fam. Foug.

(Hist. Acrost.) 74, tab. 37. 1845. = Teratophyl-

lum articulaliim (Fee) Kuhn (fide HoUtum 1978).

Polyhotrya aspidioides Grisebach, Cat. PI. Cub.

276. 1866. = Atalopteris aspidioides (Gn^chach)

Maxon & C. Chr.

Polyholrya asplenifolia (Belanger) Prcsl, Tent.

Ptcrid. 231. 1836. = Bolhitis appendiculata

(Willd.) Iwat.suki (fide Hennipman 1977).

Polyhotrya aurita Blume, Fl. Jav. Fil. 15, tab. 1.

1828. = Stenosemia aurita (Sw.) IVesl (Jldc

Christenscn 1905).

Polyhotrya hifurcata (L.f.) J. Smith, Jour. Bot.

4:150. 1841. = Elaphoglossum bifurcalum

(Jacq.) Mickel (fide Mickel 1980).

Polyhotrya blumeana (Fee) Mettenius, Fil. Lips.

24, tab. 2, fig. 10. 1856. = Leptochilus lomari-

oides Blume.

Polyhotrya cenina (L.) Kaulf. , Enum. Fil. 55.

1824. =Olfersia cenina (L.) Kunze, Flora

7:312. 1824.

Polyhotrya cicutaria Blume, Enum. Fil. 100.

1828. = Stenosemia aurita (Sw.) Presl (fide

Christenscn 1905).

Polyhotrya duplicato-serrata Hayata. Ic. Fl. For-

mosa 5:305, fig. 123. 1915. = Bolbitis rhizo-

phylla (Kaulf.) Hennipman (fide Hennipman

1977).

Polyhotrya exaltata Brackenridge, U.S. Expl.

Exp. 16:78. 1854. = Bolhitis rhizophylla

(Kaulf.) Hennipman (fide Hennipman 1977).

Polyhotrya filiculifolia (L.) Farw., Amer. Midi.

Nat. 12:303. 1931. = Anemia?

Polyhotrya flabellata (Willd.) Mettenius, Ann.

Sci. Nat., Paris, V, 2:206. 1864. = Peltapteris

peltata (Sw.) Morton.

Polyhotrya fraxinifolia (Presl) Mettenius. Fil.

Lechler. 2:8. 1959. = Bolbitis serratifolius

(Kaulf.) Schott (fide Hennipman 1977).

Polyhotrya furcata (L.f.) Kuhn, Fil. Afr. 52.

1 868 .
= Elaphoglossum hifurcata (Jacq. ) Mickel

(fide Mickel 1980).

Polyhotrya gaudichaudiana (Gaudich.) Moore.

Index Fil. XV. 1857. = Bolhitis appendiculata

(Willd.) Iwatsuki (fide Hennipman 1977).

Pohhotrya hamiltoniana (Wallich) F^e, Mem.
Fam. Foug. (Hist. Acrost.) 78. 1845. = Bolhitis

appendiculata (Willd.) Iwatsuki (fide Hennipman

1977).

Polyhotrya helfcriana Kunze, Famkr. Erde 2:35,

tab. 94. 1848. = Bolbitis appendiculata (WxUd.)

Iwatsuki (fide Hennipman 1977).

Polyhotrya intermedia Fee, Mem. Fam. Foug.

(Hist. Acrost.) 76, tab. 40, fig. I. 1845. = Bol-

hitis rhizophylla (Kaulf.) Hennipman (fide Hen-

nipman 1977).

Polyhotrya lomarioides (Blume) Kuhn. .-^nn.

Mus. Ludg.-Bat. 4:295. 1869. = Lomagramma

lomarioides (Blume) J. Smith (fide Holttum

1978).



November 1987 Monograph of Polybotra 121

Polybotrya marallioides Braekenridge, U.S.

Expl. Exf)ed. 16:79. 1854. = Asplenium maral-

lioides (Braekenridge) C. Chr. (fide Christensen

1905).

Polvbotrva marginataB\umc, Enum. Pi. Jav. 100.

1 828. = Bolhilis appendiculato ( Wiljd. ) Iwat.suki

{fide Hennipman 1977).

Polybotrya montana (Gaudich.) Moore, Index Fil.

350. 1862. =Bolbitis appendicukitu (Willd.)

Iwatsuki (fide Hennipman 1977).

Polybotrya nana Fee, Mem. Fam. Foug. (Hist.

Acrost.) 75, tab. 38, fig. 1. 1845. = Blechmm

lanceolalum (R. Br.) Sturm, {jide Christensen

1905).

Polybotrya neglecta Fee, Mem. Fam. Foug. (Hist.

Acrost.) 75, tab. 39, fig. 2. 1845. = Bolbilis

rhizophylla (Kauif.) Hennipman (fide Hennipman

1977).

Polybotrya nieuwenhuisenii Raciborski, Bull. Int.

Ac. Cracovie 57. 1902. = Heterogonium steno-

semioides (Baker) Christ {fide Holttum 1975).

Polybotrya iwdiflora Belanger, Voy. Ind. Or. Bot.

2:17. 1833. = Bolbilis appendiiidata (Willd.)

Iwatsuki (fide Hennipman 1977).

Polybotrya orientalis Blume, Enum. PI. Jav. 99.

1828. = Stenosemia aitrita (Sw.) Presl (fide

Christensen 1905).

Polybotrya peltata (Sw.) J. Smith, Jour. Bot.

4:150. 1841. = Peltapteris peltata (Sw.)Monon.

Polxbotrya plumieri (Fee) Moore, Index Fil. 13.

1857. = Elaphoglossum plumieri Moore (fide

Proctor 1977).

Polybotrya polyphylla (Braekenridge) C. Chr.,

Index Fil. 505. 1906. = Lomagramma polyphylla

Braekenridge.

Polybotrya proiifera Mettenius, Fil. Lips. 24, pi.

2, fig. 11. 1856. = Bolbilis subcrenala (Hooker

& Grev.) Ching (fide Hennipman 1977).

Polybotrya pleroides (i . Smith) Kuhn, Ann. Mus.

Bot. Lugd.-Bat. 4:295. 1869. = Lomagramma

pteroides i. Smith (fide Holttum 1978).

Polybotrya quercifi>lia (Ret/.) Mettenius, Fil.

Lcchler. 2:12. 1859. = Leplochilus zeylandicus

(Houtt.) C. Chr. (fide Christensen, 1905).

Polybotrya rhizophylla (Kaulf.) Presl, Tent.

Pterid. 231. 1836, = Bolbilis rhizophylla

(Kaulf.) Hennipman (fide Hennipman 1977).

Polybotrya serrulala Fee, Mem. Fam. Foug.

(Hist. Acrost.) 76, pi. .39, fig. ii. 1845. = Bol-

hilis rhizophylla (Kaulf.) Hennipman (Jlde Hen-

nipman 1977).

Polybotrya sinensis (Baker) C. Chr., Index Fil.

57. 1913. = Bolbilis sinensis (Baker) Iwatsuki

(fide Hennipman 1977).

Polybotrya sorbifolia (L.) Keyserling, Pol. Cyath.

Herb. Bung. 32. 1873. nomen illegil.

= Lomariopsis sorbifolia (L.) Fee (fide Proctor

1977).

Polybotrya stenosemioides (Baker) Copel.,

Polypod. Philipp. 40. 1905. = Helerogonium

stenosemioides (Baker) C. Chr. (Jlde Holttum

1975).

Polybotrya subqiiinqitefida (Fee) Mettenius, Fil.

Lechler. 2:12. 1859. =lLeptochilus laiifolium

(Meyen) C. Chr. (fide Christensen 1905).

Polybotrya laccaefolia (J. Smith) Mettenius, Fil.

Leehler. 2:12. 1859. = Leplochilus latifolius

(Meyen) C. Chr. (Jlde Christensen 1905).

Polybotrya lenuifolia (Desv.) Kuhn, Fil. Afr. 52.

1868. = Slenochlaena lenuifolia (Desv.) Moore

(fide Christensen 1905).

Polybotrya leysmanniana (Baker) Posthumus,

Ree. Trav. Bot. Neeri. 33:872. 1930. =

Stenosemia leysmanniana (Baker) Diels (fide

Christensen 1905).

Polybotrya trilohaia (J. Smith) Mettenius, Fil.

Lips. 24. 1856. = Leptochiltts latifolius (Mcycn)

C. Chr. (Jlde Christensen 1905).

Polybotrya tripartita (Hooker & Grev.) J. Smith,

Jour. Bot. 4:150. 1841. - Peltapteris tripartita

(Hooker & Grev.) Morton (fide Morton 1955).

Polybotrya vivipara Hooker, Exotic Flora, 2: pi.

107. 1825. = Bolhilis appendiculala (Willd.)

Iwatsuki subsp. vivipara (Hooker) Hennipman

(fide Hennipman 1977).

Polybotrya wilkesiana Braekenridge, U.S. Expl.

Exped. 16:80, tab. 10. 1854. =Teralophyllum

wilkesianaum (Braekenridge) Holttum (fide

Holttum 1978).



122 iLLfNois Natural History Survey Vol 34. Art. 1

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124 Illinois Natural History Survey Vol.34. Art. I

Taxa and Distribution of

Polybotrya

Numbers correspond to the species numbers as-

signed in the taxonomic treatment.

1

.

Polybotrya serratifolia (F6e) Klotzsch: Trini-

dad, Venezuela.

2. Polybotrya polybotryoides (Baker) Christ:

Mexico, Belize, Guatemala, Honduras, Costa

Rica, Panama, Colombia, Ecuador, Peru.

3. Polybotrya suberecta (Baker) C. Chr.: Co-

lombia, Ecuador, Peru.

4. Polybotrya andina C. Chr.; Ecuador.

5

.

Polybotrya sorbifolia Kuhn; Costa Rica, Ven-

ezuela, Colombia, Brazil.

6. Polybotrya fractiserialis (Baker) J. Smith:

French Guiana, Surinam, Guyana, Ecuador,

Peru, Bolivia.

7. Polybotrya crassirhizoma Lellinger: Colom-

bia, Ecuador, Peru, Bolivia, Brazil.

8. Polybotrya espiritosantensis Brade: Brazil.

9. Polybotrya caudata Kunze: Mexico, Guate-

mala, Belize, Honduras, Nicaragua, Costa

Rica, Panama, Trinidad, French Guiana,

Surinam, Guyana, Venezuela, Colombia,

Ecuador, Peru, Bolivia, Brazil.

10. Polybotrya goyazensis Brade: Brazil, Para-

guay.

1 1

.

Polybotrya pubens Martins: Colombia, Ecua-

dor, Peru, Bolivia, Brazil.

12. Polybotrya glandulosa Kuhn: Venezuela,

Peru, Brazil.

13. Polybotrya lechleriana Mettenius: Guyana,

Colombia, Ecuador, Peru, Bolivia.

14. Polybotrya attenuata Moran: Colombia.

15. Polybotrya stohei Moran: Colombia.

16. Polybotrya alfredii Brade: Nicaragua, Costa

Rica, Panama, Ecuador, Peru, Bolivia.

17. Polybotrya botryoides (Baker) C. Chr.: Co-

lombia.

18. Polybotrya lourteigiana Lellinger: Colombia.

19. Polybotrya pittieri Lellinger: Colombia.

20. Polybotrya cylindrica Kaulfuss: Brazil.

21. Polybotrya hickeyi Moran: Colombia. Bo-

livia.

22. Polybotrya puberulenta Moran: Ecuador,

Bolivia.

23. Polybotrya alata Moran: Panama.

24. Polybotrya aequatoriana Moran: Ecuador,

Bolivia.

25. Polybotrya appressa Moran: Ecuador.

26. Polybotrya altescandens C. Chr.: Colombia,

Ecuador, Peru.

27. Polybotrya gomezii Moran: Costa Rica.

28. Polybotrya osmundacea Willd.: Guatemala.

Honduras, Nicaragua, Costa Rica. Panama,

Cuba, Jamaica. Haiti. Grenada, Trinidad,

French Guiana, Guyana, Venezuela. Colom-

bia, Ecuador. Peru. Bolivia. Brazil.

29. Polybotrya cyathifolia Fee: Guadeloupe,

Martinique.

30. Polybotrya latisquamosa Moran: Colombia.

31. Polybotrya sessilisora Moran: Colombia,

Brazil.

32. Polybotrya canaliculata Klotzsch: Vene-

zuela.

33. Polybotrya semipinnata ¥€t: Brazil.

34. Polybotrya speciosa Schott: Brazil.

35. Polybotrya pilosa Brade: Brazil.

Distribution Maps

The small open dots on the twenty-one distribution

maps that follow indicate towns. Other symbols

are explained in the legends.



November 1987 Monograph of Polybotra 125

Map 1. Distribution of Polyhotrya suherecta (Baker) C. Chr. (diamond) and P. serratifolia (Fde)
Klotzsch (circle). Map 2. Distribution of Polybolrya polyhoinoides (Baker) Christ (circle) and P
andina C. Chr. (square).



126 Illinois Natural History Survey Vol. 34. Art. 1

Map 3. Distribution of Potybolna sorhifolia Kuhn (diamond) and P. fractiseriaUs (Baker) J. Smith

(circle). Map 4. Distribution of Polybolrya crassirhizoma Lellinger.



November 1987 Monograph of Polybotra 127

Map 5. Distribution of Polybotrya caudata Kunze. Map 6. Distribution of Polyboirya pubens Mat-
tius (circle) and P. goyazensis Brade (diamond).



128 Illinois Natural History Survey Vol.34. Art. 1

•) - -^ '

Map 7. Distribution of Potybotrya lechleriana Mettcnius (circle), P. attenuata Moran (triangle), and

P. glandulosa Kuhn (square). Map 8. Distribution of Polybotna atfredii Brade (circle) and P.

boiryoides (Baker) C. Chr. (triangle). Map 9. Distnbution of Polyholrya siohei Moran. Map 10.

Distribution of Polybotrya lourteigiana Lellinger. Map 1 1 . Distribution of Potybotrya pittieri Lel-

linger.



November 1987 Monograph of Polybotra 129

Map 12. Dislnbution of Polybotryaaltescandens C.Chr. Map 13. DhlribMion of Polyholrya hickeyi

Moran (circle) and P. puberulenta Moran (triangle). Map 14. Distribution of Polyhotrya gomezii

Moran (circle) and P alata Moran (triangle). Map 15. Distribution of Polyhotrya aequatoriana

Moran. Map 16. Distribution of /'o/>'b<>/o'ac>'/iminca Kaulfuss. Map 17. Distribution of Po/>*o/-

rya appressa Moran.



1 30 iLLtNois Natural History Survey Vol. 34, Art. 1

18

Map 18. Distribution of Polyhotrya osmumlacea Willd. (circle) and P. semipinnaUi Fdc (triangle).

Map 19. Distribution of P. latisquamosa Moran (solid square). P. cimaUcuhua Klotz.sch (triangle),

and P. lyathifolia Fee (open square).



November 1987 MONCXJRAPH of POLYBOTRA 131

Map 20. Distribution of Polyhotrya sessilisora Moran (circle) and P. pilosa Bradc (triangle).

Map 21 . Distribution of Polybotrya speciosa Schott (circle) and P. espiriiosaniensis Bradc (triangle).



132 Illinois Natural History Survey Vol. 34. An. 1

Index to Collectors' Numbers

Numbers in parentheses refer to the species num-

bers assigned in the taxonomic treatment.

Acosta Solis 6358 (2).

Acuna 17531 (28).

Aguilar 947 (6).

Alfaro 8073 (5).

Allard 20661 (6); 21609 (28); 21997 (28);

22305 (6); 22526 (6); 22593 (9).

Alston 7341 (28); 7641 (28); 8278 (21);

8991 (34).

Alverson et al. 342 (2).

Anderson 10152 (5).

Antonio 4019 (2); 5133 (28).

Argent & Richards 6651(10).

Argent eta! . 6336 (10).

Aristeguieta 1780 (28); 3963 (1).

Armond 298 (2).

Atwood 3711 (28).

Aymard et al. 952 (9).

B.T. 443 (9).

Bailey & Bailey 541 (9).

Balslev 4787 (11); 4802 (11).

Balslev & Madsen 10649 (28).

Barbour 4764 (9); 4790 (9).

Barclay 2199 (9).

Barkley & Gutierrez V. 1897(28).

Barnard et al. 411 (28).

Beard 1252 (28).

Beck 1635 (7); 3060 (24); 3108 (24);

4924 (11); 8037 (9).

Bennett 23 (28).

Benoist 1271 (9).

Berget al. P18138 (28).

Berry 948 (1).

Billict & Jadin 1683 (9).

Biolley 73 (16); 10688 (9).

Boom & Mori 1856 (9).

Boutin & Schlosser 5902 (2).

Brade 372 (2); 554 (16); 5838 (.34);

8061 (33);9365 (20); 9773 (34); 9787 (35);

9843 (34); 10351 (34); 12576 (20);

12614 (20); 12616 (34); 13982 (20);

14943 (34); 15373 (10); 16456 (33);

16579 (.34); 16824 (16); 17464 (34);

18224 (8); 19972 (20); 20214 (34);

20733 (35); 20931 (20).

Bradc & Brade 67 (16); 98 (16); 374 (9).

Brandbygc & Asanza C. 3 1 870 ( 1 1 );

32881 (7).

Brandbygc ct al. 32544 (7); 32617 (7);

33684 (7).

Breedlove 34101 (9).

Breedlove & Smith 21886 (2).

Brenes 11676 (16); 14248 (16); 21980 (16).

Britton et al. 1267 (28); 1935 (9); 2144 (9);

2276 (28).

Broadway 560 (9); 2520 (28); 5358 (9);

5589 (28); 5717 (9); 6459 (28);

6902 (28); 9207 (9); 9947 (1);

9948 (28); 9949 (1);9950 (28);

9951 (28).

Buchtien 11 (6); 13 (6);35 (9&ll);260 (11);

290 (9); 298 (28); 299 (11); 1015 (6);

1066 (28); 1123 (11); 1124 (11); 2138 (6);

5164 (16); 5308 (13).

Bues 1743 (13).

Bunting 9516 (28).

Burch 4414 (16); 4505 (16); 4613 (28).

Burger & Antonio 11263(9).

Burger & Stoize 5865 (9).

Cain 74 (28).

Camp 1298 (28); 1359 (28).

Cardenas 1249 (6).

Castellanos 25710 (20).

Cazalet & Pennington 7720 (11).

Chrysler 4836 (9).

Chrysler & Roever 5095(16).

Churchill 5776 (28).

Churchill & de Nevers 4993 (2).

Clement 725 (28).

Clute 259 (28).

Conant 940 (9); 1016 (31): 1080 (31);

1482 (31).

Cooper 10240 (16).

Comman 544 (9); 927 (16); 1178 (16);

1233 (16).

Crcmers 4460 (9); 6374 (9); 6523 (9);

7369 (9); 7552 (9); 7951 (9).

Croat 690 (28); 738 (28); 4266 (2);

5114 (9); 6850 (9); 7362 (9);

8025 (9); 8153 (9); 9000 (9);

9004 (9); 9103 (9); 10804 (9);

11544 (9); 12114 (2); 12143 (2):

13431 (16); 14687 (2); 14742 (2);

14786 (2); 15081 (9); 15256 (9);

17161 (2); 17367 (9); 17620 (9);

17689 (11); 18219 (9); 18388 (9);

18476 (28); 18508 (11); 19220 (9);

19717 (9); 19771 (11); 20287 (28);

20543 (28); 20551 (7); 20763 (7):

20786 (28); 20851 (28); 21014 (6):

21152 (6); 21191 (9); 21987 (28);

22260 (28); 22682 (28); 22706 (23);

22789 (2); 23011 (23): 23244 (2);



November 1987 Monograph of Polybotra 133

24541 (9): 24566 (2); 27648 (2):

33452 (9); 35176 (9); 35608 (16);

36007 (16); 36669 (16); 36761 (16);

36800 (2); 37757 (2); 37794 (2);

38047 (28); 41637 (2); 49718 (13);

51154 (6); 55844 (26); 56510 (3).

Croat & Folsom 34051 (2).

Croat & Grayum 60022 (28).

Croat & Porter 15342 (9); 15436 (9).

Crosby 76 (28).

Crueger 139 (9).

Cuatreeasas 14246 (28); 15527 (18);

22143 (18).

Cuyler 4796 (34).

Davidse & Gonzalez 19444 (28).

DcWolf 385 (16).

Dodson 7380 (3).

Dodson et al. 8679 (3).

Donnell Smith 6939 (16).

Duarle 3179 (20).

Duarte & Pereira 65323 (20).

Duarte et. al. 65322 (33).

Dudley 10168 (9); 10325 (13); 11513 (9);

13005 (28); 13258 (16); 13290D (2);

I8265A (16).

Dumont et al. 7439 (28).

Dunn & LeDoux 22005 (9).

Duque-Jaramillo 1868 (18).

Dusen 4423 (20); 6926 (20); 6984 (20);

13658 (20); 15351 (20); 15353 (20).

Duss 1503 (29); 3897 (29); 4719 (29).

Dwyer 8337 (16).

Dyer A225 (9).

Eggers 5324 (28).

Ekman 3769 (28); 4734 (28); 5214 (28);

I42I0 (28).

Englesing 291 (9).

Esposto 10928 (3).

Evans & Bowers 2792 (28); 2944 (16);

3152 (2).

Evoy 104 (9).

Ewan 16846 (9).

Fargens 251 (20).

Fault 12583 (28).

Fay 345 (28); 373 (28); 472 (9); 859 (1).

Fendlcr 69 (28); 105 (9); 235 (1); 261 (1);

262 (32).

Fernandez 951 (9).

Fiedler* Koptur 51 (27).

Fisher 132 (28).

Fleming & Fleming 52 (28).

Folsom 3590 (2).

Forero & Jaramillo 1745 (9); 2469 (13).

Foster P-84-42 (9); P-84-91 (6); 85-37 (3);

85-162 (2); 4020 (6); 7455 (6); 7858 (11);

7937 (6); 9284 (7).

Foster & Foster 854 (8).

Foster & Kennedy 1814 (2).

Foumier 357 (2).

Gardner 1901 (5).

Gastony 43 (28).

Gaudichaud 73 (20).

Gentle 2947 (9).

Gentry & Juncosa 41021(28).

Gentry et al. 27365 (6); 27880 (7).

Gerdes 102 (33).

Gerrera 1624 (13).

Gill 47 (7).

Glaziou 955 (20); 2427 (33); 2428 (20);

14456 (10).

Gleason 582 (9).

Goedas 217 (20).

Gomez 3324 (2); 3354 (9); 4528 (9);

6914 (2); 7027 (2); 7122 (5);

18064 (2); 18906 (16);

19489 (9 & 28).

Gomez &Cittar 6409 (9); 6731 (9).

Gomez et al. 20423 (9).

Gonggrijp & Stahcl 3300 (6).

Grant 10283 (21); 10436 (9); 10556 (3).

Granville 971 (9); 991 (9); 1143 (9);

1165 (9); 3674 (6); 3865 (28); 4012 (9);

B.4711 (9); 4841 (9); 5121 (6); 5219 (6).

Grayum & Sleeper 3448 (16).

Grijalva 300 (28).

Guppy 462 (9).

Gutierrez & Barkley 170109(28).

Haerchen 124 (20).

Hammel 3503 (2).

Handro 1229 (20); 2228 (20).

Harley et al. 10922 (10).

Marling et al. 7535 (28).

Hart 53 (9); 228 (1); 229 (28).

Hassler 11618 (10).

Hatch 90 (28).

Hatschbac-h 7424 (20); 8141 (20); 10749 (20);

25118 (10).

Haught 1325 (9); 1757 (5); 5498 (13).

Hayes 8 (9).

Henri-Stanislas 1709 (2); 1710 (28).

Herrcra 12(X) (6).

Hickey 801 (21).

Hioram 2495 (28).

Hioram & Clement 6424 (28).

Hodge 6592 (26); 6780 (2).

Holdridgc 5153 (2).



134 Illinois Natural History Survey Vol.34. Art. I

Holm & litis 200 (9 & 28).

Holm-Nielsen & Jeppsen 663 (11); 987 (7).

Holm-Nielsen et al. 4489 (9); 4495 (9);

22121 (11); 22484 (7).

Huber 2599 (9); 7235 (9).

Humboldt 459a, b (28).

Hunnewell 16514 (5); 18511 (20).

Idrobo & Schultcs 1106 (30).

Idroboct al. 10116 (28).

Inarte 52 (1).

Irwin 55161 (9).

Irwin et al. 54669 (6); 54784 (6); 54896 (9).

Jameson 33 (26).

Jaramillo et al. 30783 (28).

Jenny 2137 (9); 2368 (28); 2846 (28);

3122 (9); 10826 (9); 11195 (28).

Jimenez M. 803 (16); 3200 (9); 3209 (2);

3210 (2); 3299 (28); 3304 (28).

Johnson 1221 (9).

Johnston 452 (9).

Jones & Facey 3246 (9).

Juncosa 1369 (17); 1467 (3).

Kalbreyer 1254 (2); 1798 (2); 1873 (17);

1877 (3).

Karsten 13 (32).

Kellerman 7354 (9).

Kennedy et al. 2074 (2).

Kilhp 2544 (9); 2845 (9); 2929 (9);

5163 (16); 5202 (16); 5386 (16);

5429 (16); 11343 (9); 11549 (15).

Killip & Cuatrecasas 38902 (28).

Killip & Lasscr 37756 (28).

Killip & Smith 15341 (28); 20210 (21);

23876 (7); 23916 (11); 23991 (28);

24605 (7); 25452 (28); 25921 (13);

25972 (11); 26194 (11); 26543 (II);

26637 (9); 26955 (11); 28760 (11);

30690 (9).

Kirkbridc 404 (5).

Kirkbride & Hayden 274 (2).

Klawe 1474 (28); 1.504 (28); 1545 (2).

King 1166 (9); 1386 (7); 1390 (12);

3208 (28).

Knapp 4966 (9).

Knapp & Dressier 5461 (2).

Knapp & Mallet 5133 (9); 5402 (9).

Knapp etal. 4479 (9); 4542 (2).

Kramer 1954 (9).

l-acgaard 511.36 (I1);5I150 (11);5I224 (II).

Lankester 653 (16).

Lasser & Vareschi 3926 (9).

Lechler 2156 (13); 2176 (13); 2321 (9);

2329 (9).

Lehmann B.T413 (13);500B (14); 2998 (9).

Lehnun 2562 (20).

Lellinger&delaSota 26 (9);39 (2); 190 (2);

213 (28|; 248 (3); 250 (28); 251 (18);

280 (15); 281 (13); 284 (15); 387 (18);

496 (9); 553 (28); 589 (9); 643 (9);

747 (15); 748 (15); 768a (18); 790 (9);

843 (18); 894 (3); 899 (18); 948 (18).

Lellinger & White 1244 (16); 1361 (16);

1436 (28).

Lent 299 (28); 1084 (27); 2642 (16);

3537 (27).

Leon 226 (6).

Liesner 820 (2); I6I33 (28); 16293 (12).

Liesner & Judziewicz 14855 (16).

Liesner et al. 8359 (28).

Lindeman 4570 (9).

Lindeman & Teunissen 15291 (9).

Lindsay 262 (28).

Loefgren & Duvall 36217 (33).

Ludwig 262 (21).

Luederwaldt 21547 (33).

Luetzelburg 254 (20); 6858 (34).

Lugo S. 3293 (7).

Lundell 6416 (9).

Macbride 5602 (7).

Macedo 1447 (10); 1521 (10); 2682 (10).

Madison etal. 3356 (2); 4851 (26).

Maguire & Fanshawe 22855 (9).

Maguire et al. 46068-A (6); 54384 (6).

Maurel & Maurel 3816 (28).

Maurice 699 (16).

Maxon 412 (16); 639 (2t; 2293 (28);

2470 (28); 4641 (9); 6895 (9);

8978 (28); 9366 (28); 9464 (28);

9522 (28).

Maxon & Har\ey 8242 (16).

Maxon & Killip 148 (28); 756 (28).

Maxon et al. 6817 (9).

McAlpin & Kuhn 77-12 (14).

Mendez 75 (9).

Mexia 6246a (6).

Mickel 1944 (9); 2003 (16); 2624 (28);

2742 (2); 2803 (28); 2817 (28); 2958 (16);

3126 (16); 3180 (2): 3368 (9); 3426 (16);

3570 (9); 9472 (9).

Molina R. 1907 (9).

Mora 4160 (18); 4287 (13); 4430 (9).

Moran 2167 (28); 2170 (2); 2171 (2);

2173 (2); 2176 (2); 2178 (2); 2182 (9);

2185 (9); 2186 (9); 2237 (16); 2241 (16);

2325 (16); 2441 (27); 2442 (16); 3145 (5);

3160 (27);3168 (16);3214 (16);3241 (27);



November 1987 Monograph of Polybotra 135

3271 {27);3338 (27);35I2 (24);3527 (25);

3528 (22); 3530 (2); 3532 (16); 3534 (9);

3535 (7); 3536 (6); 3544 (26); 3546.5 (3);

3547 (28); 3559 (26); 3561 (3); 3562 (2);

3563 (4); 3564 (26); 3565 (3); 3569 (2);

3570 (16); 3585 (24); 3586 (25); 3588 (9);

3590 (2); 3592 (28); 3593 (2); 3600 (28);

3612 (28); 3615 (7); 3616 (7); 3617 (9);

3618 (28); 3640 (7); 3641 (7); 3642 (28);

3663 (9); 3671 (9); 3688 (26); 3709 (1);

3718 (28).

Morcly & Whitefoord 655 (28); 675 (28);

695 (28).

Mori & Kallunki 3574 (2).

Moritz 277 (1);278 (32).

Morton 7629 (2).

Mosen 3050 (20).

Mullcr 6039 (20).

Murillo 1457 (5); 1491 (5); 2580 (28).

Murilloet al. 289 (5).

Nee & Stockwell 11610 (2).

Nee et al. 8768 (2).

Nelson et al. 3296 (2).

Nieil 2610 (9).

Niemcyer 44 (28).

Ocampo 727 (2); 877 (16).

Oldcman 1933 (9).

011gaard et al. 34634 (9); 34703 (11);

34748 (7); 34855 (7); 34894 (11);

34970 (7); 35124 (6); 35290 (9);

35350 (11); 35776 (24); 37702 (26);

37820 (26); 37823 (3); 37859 (3);

37867 (26); 38836 (7); 38845 (28);

38932 (9); 38894 (28); 38969 (11);

39039 (11); 39040 (11); 39084 (7);

39086 (28); 43798 (9); 53824 (25).

Onega 636 (9).

Ortega & Grimann 2707 (5).

Ortega & Smith 2387 (28); 2491 (1);

2498 (32); 2510 (1); 2511 (32).

Pabst el al. 6772 (34).

Perkins 1163 (28).

Pcrsaud 114 (13); 372 (9).

Pipoly 3542 (9); 38 17 (9); 3824 (9); 3880 (9);

3901 (9); 5146 (28); 5299 (9).

Pittier 587 (19); 4492 (9); 9016 (28);

9076 (28); 10688 (9); 12416 (28);

16232 (2).

Plowman & Davis 4542(13).

Plowman & Thomas 13678(28).

Plowman et al. 4025 (7).

Pocppig 201 (9).

Porter et al. 4763 (9).

Prance & Silva 59677 (10).

Pranceetal. 10466 (5); 12069 (9); 12180 (7);

15332 (31).

Prestoe 1491 (1); 1492 (1).

Proctor 3969 (28); 16506 (28); 22269 (28).

Proctor et al. 27091 (9).

Purpus 6761 (2); 7245 (2).

Ratter et al. 2047 (10).

Rcitz 151 (20); 2825 (20).

Reitz& Klein 1101 (20); 1102 (20);

2665 (20); 3466 (20); 4635 (20);

4682 (20); 5108 (20); 5549 (20).

Richards 803 (9).

Richardson 2037 (28).

Riedel 81 (35).

Rimbach 31 (13); 91 (26); 312 (26).

Rivero 513A (1).

Riveroet al. 1608 (1).

Rojas 10182 (10); 10431 (10); 10804 (10).

Rose & Russell 20790 (34).

Rossbach 3624 (9); 3625 (9); 3628 (28);

3710 (9).

Rovirosa 972 (2).

Rusby 442 (II); 443 (22).

Sandwith 1561 (9).

Scamman 5984 (28); 5985 (16); 5986 (16);

5987 (28);7152 (28);7153 (28);7155 (16);

7156 (9); 7482 (9); 7687 (28).

Scamman & Holdridge 7998 (9).

Schipp 273 (9); 8101 (28); 8108 (28).

Schmalz 78 (20); 158 (20).

Schnee 615 (1).

Schneus 3412 (20).

Schomburgk 1659 (9).

Schultes & Black 46-266 (7); 8467 (9).

Schultes & Cabrera 13963 (31); 16053 (11).

Schunke, C. 157 (7); 158 (7); 164 (28);

165 (6); 380 (9); A214 (6); 661 (7); 666

(6); 705 (28); 812 (7); 1341 (28); 1395

(28); 1396 (6); 1451 (28).

Schunke, J. 268 (7); 275 (28); 380 (28);

5785 (28); 5789 (11); 10173 (6);

10200 (26).

Seavems 56 (9).

Sehnem 3092 (20).

Seifriz 24 (3).

Shafer 4458 (28); 8889 (28).

Shemluck 280 (7); 304 (16).

Shcrring 13 (28).

Silva et al. BG526 (5).

Skogetal. 5036 (2).

Skutch 3018 (28); 3236 (16); 4637 (28).



136 Illinois Natural History Survey Vol. 34. Art. I

Smith, A.C. 48/293 (16); 778 (16);

2984 (31); 61713 (13).

Smith, A.R. 1259 (1);4055 (1).

Smith, A.R. et al. 922 (1); 1347 (28);

1388 (28).

Smith, H.H. 983 (3); 1050 (28); 1052 (5).

Smith & Brade 2286 (20).

Smith & Klein 7546 (20).

Smith & Reitz 6135 (20).

Sodiro 81 (28).

Soejarto 2805 (28).

Sola, de la 2343 (34); 5181 (28); 5252 (16).

Soukup 1065 (7).

Sperling 5914 (9).

Sperling & Bleiweiss 5034 (26).

Sperling et al. 5915 (28).

Spruce 2116 (9); 3880 (11); 4090 (6);

4337 (6); 4634 (9); 4744 (13); 4740 (II);

5685 (28).

Standley 9047 (9); 24195 (9 & 28);

25085 (9); 27597 (9); 33648 (16);

37096 (28); 53955 (2); 53983 (9);

67135 (2); 68203 (2).

Standley & Valerio 47127 (28); 48622 (9).

Starry 91 (9).

Sleeves & Ray 504 (28).

Steinbach 3032 (9); 7499 (9).

Stemetal. 523 (9); 1034 (16).

Stevens & Krukoff 6518 (16); 8827 (9);

12056 (9); 12666 (9); 12741 (9);

13005 (9).

Stewart 241 (28).

Steyermark 37320 (2); 38229 (9); 38271 (9);

39197 (9); 39895 (9); 41698a (2);

41870 (2); 44675 (9); 52812 (26);

56114 (1); 61991 (1); 62015 (28);

89120 (9); 91756 (1); 94952 (28);

99185 (32); 104717 (1); 107148 (28).

Steyermark & Davidse 116499 (28).

Steyermark & Liesner 120634 (28).

Steyermark & Nevling 95930 (28).

Steyermark & Rabe 71756 (28); 96145 (28);

97133 (1).

Steyermark & Stoddart 1 1 805 1 ( 1 )

.

Steyermark et al. 95827a (5); 101552 (28);

106758 (1); 114331 (9); 114332 (9);

115021 (9); 115033 (9); 122814 (28);

124749 (I); 125039 (28); 126675 (1);

126680 (1).

Stolze 1488 (16).

Stork & Horton 9509 (9).

Stubel 875 (28); 913a (13): 914 (13).

Sugden 1151 (28).

Swmgle et al. 70-02-05-2 (9).

Sydow 339 (26).

Tate 422 (9); 502 (6); 513 (6).

Taylor 447 (28); 1223 (9).

Tonduz 9006 (28); 9016 (28); 13337 (28);

14568 (28); 18879 (28).

Tryon & Kramer 5611 (9).

Tryon&Tryon 5204 (9); 5221 (6); 6591 (20).

Tschudi 162 (5); 166 (32); 167 (I).

Tuerckheim 148 (9): 358 (9); 7812 (9);

8040B (9); 8041 (9).

Tyson 2109 (9).

Underwood 1246 (28); 1250 (28); 1323 (28):

1612 (28).

Valerio 63 (16): 328 (28): 329 (2); 333 (28);

2356 (16); 33133 (2).

van Cotthem 1327 (1).

van der Werff 3486 (32).

van der Werff & Gonzalez 5202 (28).

van der Werff & Wingfieid 3430 (1 ).

Vareschi 3147 (28); 7764 (I).

Vareschi & Gessner 1875(28).

Vareschi & Pannier 1686 (28); 2636 (5):

2660 (28): 2715 (28).

Vargas 11280 (6); 17743 (28); 17800 (9).

von Sneidem 1590 (19).

Wacket 21556 (20).

Walker T10995 (28).

Watt 160 (28): 7276 (28).

Webb 26 (34).

Webster et al. 16467 (2).

Weddell 952 (35).

Wcrckic 559 (16); 16770 (2).

Wetmore & Woodworth 130(9).

Whitmorc 752 (7).

Wiggins 11073 (26).

Williams 1188 (6): 17.39 (9): 2137 (9);

3136 (11); 3999 (9); 4797 (11);

7620 (ID; 7681 (9).

Wilson & Murray 564 (28).

Wilson & Webster 549 (28).

Wingficld 6914 (28).

Wright 786 (28).

Wurdack 1854 (6): 1933 (11): 2011 (9).



November 1987 Monograph of Polybotra 137

Index to Taxonomic Names

Accepted names are set in roman type; new taxa

and combinations appear in bold type; ail other

names are italicized. A number after a name refers

to the number of the accepted taxon . Abbreviations

refer to sections of this monograph: ex (excluded

names), unc (names of uncertain application), sub

(subdivision of the genus).

Acrostichum

botryoides 1

7

caenopteris 1

var. salicifolium 5

canaliculatum Til

caudatum 9

var. pubens 1

1

chrysolepis 26

cylindricum 20

fracliseriale 6

hackelianum 3

hanii I

incisum 20

insigne 4

juglandifoUum 2

lechlerianum 1

3

osmundaceum 28

plumbicaule 6

polybotryoides 2

pubens I 1

suhereclum 3

Aspidium

scanJens 33

Botryothallus see Polybotrya

kunzei 1

Dryopteris

guentheri 1

1

Olfersia

caudata 9

cervina ex

Polybotrya

sect. Arthrobotrya ex

sect. Egenolfia ex

sect. Lomagramma ex

sect. Teraiophyllum ex

subg. Ectoneura ex

subg. Egenolfia ex

subg. Polybotrya sub

subg. Sorbifolia sub

subg. Soromanes sub

acroslichoides ex

acuminata unc.

var. villosa 9

aequatoriana 24

alaU 23

alfredii 16

forma carpinlerae

altescandens 26

andina 4

apiifolia ex

appendiculaia ex

appressa 25

arfakensis ex

aristeguielae 28

arliculala ex

aspidioides ex

asplenifolia ex

attenuata 14

aucuparia 2

aurita ex

bifurcata ex

blumeana ex

botryoides 1

7

canaliculata 32

caudata 9

cervina ex

cicutaria ex

coenopteris I

costaricensis 9

crassa 1

crassirhizoma 7

crespiana 1

1

cyathifolia 29

cylindrica 20

decorata 1

1

duplicato-serrata ex

espiritosantensis 8

exallaia ex

filiculifolia ex

flabellala ex

fractiserialis 6

fraxinifolia ex

frondosa 20

fulvostrigosa unc

furcata ex

gaudichaudiana ex

glandulosa 12

gomezii 27

goyazensis 10

gracilis 1

6

hamiltoniana ex

/lar/// I

helferiana ex

hickeyi 2

1

incisa 20



138 Iij iNois Natural History Survey Vol. 34. Art. 1

intermedia ex

juglundifolia 2

var. lobata 2

kalhreyeri 2

kunzei I

latisquamosa 30

lechlcriana 13

litoralis 34

lomarioides (Blume) Kuhn ex

lomarioides Mettenius unc

lourteigiana 18

macbridei 7

macedoi 9

marattioides ex

marginata ex

monlana ex

nana ex

neglecia ex

nieuwenhuisenii ex

nodiflora ex

nutans unc

orientalis ex

osmundacea 28

var. crispopaleacea 34

var. cylindrica 20

var. frondosa 20

var. incisa 20

peltata ex

pilosa 35

pittieri 1

9

plumbicaulis 6

plumieri ex

polybotryoides 2

polyphylla ex

prolifera ex

pteroides ex

pubcns 1

1

puberulenta 22

quercifolia ex

rhizophytla ex

rosenstockiana 34

salicifolia 5

scandens Fee unc

scandens (Raddi) Christ

semipinnata 33

serratifolia 1

serrulata ex

sessilisora 3 1

i/neni/j ex

sorbifolia Kuhn 5

sorbifolia (L.) Keys, e;

speciosa 34

stenosemioides ex

stolzei 1

5

subelliptica 1

2

suberecta 3

subquinquefida ex

taccaefoliii ex

tenuifolia ex

teysmanniana ex

tomentosii 34

trapezoides unc

trilobata ex

ex

28

9

ex

ex

33

tripartita

vareschii

villosula

vivipara

wilkesiana

Polypodium

adiantoides 9

Psoniiocarpa

acuminata unc

caudata 9

Soromanes see Polybotrya

coenopteris I

dentatum 1

integrifolium 1

serratifolium 1

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