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The Ferns and Fern-allies
of Costa Rica, Panama, and the Choco

(Part 1: Psilotaceae through Dicksoniaceae)

by

David B. Lellinger

p^eRi6oLo6ia

a pueLicatioa of
cfieanieRicari fepin sociecy

Number 2A . . . . 1989

Pteridologia

Editor

David B. Lellinger, Dept, of Botany, NHB-166, Smithsonian Institution,
Washington, DC 20560.

Associate Editors

Donald R. Farrar, Dept, of Botany, Iowa State University, Ames, lA 50011.

Alan R. Smith, Dept, of Botany, University of California, Berkeley, CA 94720.

Terry R. Webster, Biological Sciences Group, University of Connecticut, Storrs,
CT 06268.

PTERIDOLOGIA is a series of monographs devoted to the publication of
longer works dealing with a broad range of topics concerning ferns and fern-allies.
It is published irregularly as separate numbers of 32 pages or more, independently
paged. PTERIDOLOGIA is owned by the American Fern Society, Inc.

Matters for publication should be addressed to the Editor. Standing orders,
pre- and post-publication orders, changes of address, and claims for missing issues
should be sent to the Editor.

PTERIDOLOGIA issues are announced several months in advance of
publication and will be available on standing order at 20% discount or on
individual pre-publication order at 10% discount. Dealer inquiries welcome.

52 h
■ tgL‘4-5

The Ferns and Fern-allies
of Costa Rica, Panama, and the Choco

(Part 1: Psilotaceae through Dicksoniaceae)

by

David B. Lellinger

Department of Botany, NHB-166
National Museum of Natural History
Smithsonian Institution
Washington, DC 20560

American Fern Society, Inc.

Illustrations (c) 1989 by the

Smithsonian Institution

Library of Congress Cataloging- in-Publication Data

Lellinger, David B.

The ferns and fern-allies of Costa Rica, Panama, and the Chocd /
by David B. Lellinger.

p. cm. -- (Pteridologia ; 2-)

Includes bibliographies.

Contents: pt. 1. Psilotaceae through Dicksoniaceae .

ISBN 0-933500-01-7 : $32.00 (v. 1)

1. Fems--Costa Rica. 2. Ferns- -Panama. 3. Ferns- -Colombia-
-Chocd (Dept.) 4. Pteridophyta- -Costa Rica. 5. Pteridophyta-
-Panama. 6. Pteridophyta- -Colombia- -Chocd (Dept.) I. Title.

II. Series: Pteridologia ; 2, etc.

QK526.C8L45 1989

587'.097286--dcl9 89-6461

CIP

ISBN 0-933500-01-7

Printed in the United States of America

CONTENTS

Introduction 5

List of the families, genera, and subgenera 13

Key to the families 15

Floristic treatments

Psilotaceae 21

Lycopodiaceae 24

Selaginellaceae 44

Isoetaceae 70

Equisetaceae 72

Ophioglossaceae 76

Marattiaceae..... 82

Osmundaceae 91

Plagiogyriaceae 94

Schizaeaceae 95

Parkeriaceae 107

Adiantaceae 109

Vittariaceae 160

Pteridaceae 173

Loxsomaceae 184

Hymenophyllaceae 185

Gleicheniaceae 229

Polypodiaceae 24 1

Cyatheaceae 334

Dicksoniaceae 363

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INTRODUCTION 5

INTRODUCTION

My interest in the pteridophytes of Costa Rica, Panama, and the Choc6 began
as a result of helping to teach a fern course in Costa Rica offered by the
Organization for Tropical Studies in the summer of 1967. In preparing for the
course, I compiled most of a checklist of the pteridophytes of Costa Rica based on
specimens present in the U. S. National Herbarium. Later, in attempting to
identify specimens collected during the course, it became obvious that the
checklist was incomplete, that few adequate monographic works were available to
aid in identification, and that a modern pteridophyte Flora was vitally needed for
the region.

. The pteridophytes of the area are as diverse as the landscape; I recognize about
1100 taxa in 117 genera, and new species or species new to the area still are
discovered from time to time. Species that became known to me after the plates
were assembled (e.g., 137a Llavea cordifolia) are given "a" numbers. They and a
few others are not illustrated. Judging by the rate of discovery, the Flora appears
to be at least 90% complete. The relatively little-known Atlantic slope of the
mountains from Central Costa Rica to central Panama is most likely to yield new
discoveries and should have first priority in future exploration.

The Nicaraguan border with Costa Rica was chosen as the northern boundary
for the Flora because Nicaragua lacks the high mountains that characterize Costa
Rica and western Panama. As Costa Rica is phytogeographically continuous with
Panama, so is eastern Panama with adjacent Colombia. To find a natural
southern limit for the Flora, Prof. Elias R. de la Sota, of the Museo de La Plata,
La Plata, Argentina, and I undertook a survey of Choco pteridophytes in
January -March, 1971. We gathered what data we could from other pteridophyte
collections and made several collecting trips to diverse sites, mostly in the
northern half of the Depto. del Choco. We found that the Choco pteridophyte
flora changed rather abruptly at an elevation of 1000 m in the Andes Mountains to
the east of the Choco basin, and so this was taken (with the Golfo de Urabd and
the drier regions to the east and the perpetually wet regions to the south beyond
the Rio Calima in Depto. El Valle) as the limit of the Flora (Lellinger, 1975).
Although these boundaries do not coincide precisely with those of the Depto.
Choco, they do come fairly close. Tallies of the species of pteridophytes found in
the Choco and in Panama and other countries and island groups of the western
hemisphere also show that the pteridophytes of Costa Rica, Panama, and the
Choc6 are a coherent phytogeographic group (Lellinger, 1975, 1985). Cocos
Island, a possession of Costa Rica far from the mainland, is not treated in this
Flora (see G6mez P., 1975a, b).

Many species of pteridophytes are quite restricted in the elevations at which
they will grow; coastal and lowland habitats versus montane habitats is a
fundamental and useful distinction to make. The lowland ferns (up to 500 m
elevation) tend to be widespread, ranging as far as southern Mexico and Brazil.
The montane ferns (500-1500 m or greater elevation) are often more restricted in
distribution and are more likely to be endemics. The richest pteridophyte areas
are the undisturbed saddles between mountains from central Costa Rica to
western Panama at mostly 1000-1500 m elevation, where cloud-borne moisture

6 INTRODUCTION

from the Caribbean refreshes the epiphytic ferns almost daily, even during the dry
season. Any lengthy dry season, as in the northern half of the Choco, makes it
impossible for most epiphytes to grow, and so severely limits their diversity.
Throughout the Flora area, species diversity is greatly diminished in secondary
forests, even those in good condition that are often thought to be primary forests.
The diversity of ferns and fern-allies in primary forests is at least twice that in old
secondary forests in good condition. Many of the rare epiphytes in such genera as
Elaphoglossum are known only from old collections. It may well be that some did
not survive cutting of the primary forests and are now extinct. The best hope for
rare species lies in preserving large tracts of undisturbed forest, as is being carried
out on an unprecedented scale by the people of Costa Rica.

Map of Costa Rica

INTRODUCTION 7

My aim has been to produce a Flora especially useful for identification and
curation. Therefore, I emphasized the keys, generally using two or more
characters in each couplet to make each choice more certain. In addition, I tried
to include all names based on New World types, and so the synonymies are
extensive and will be useful widely. Specimens wrongly named or placed under
synonyms in herbaria are an impediment to rapid and careful taxonomic work.

This book is arranged in part taxonomically, with few exceptions according to
the generic arrangement of Crabbe, Jermy, and Mickel (1975), so that related
families and genera fall close together (see List of Families, Genera, and
Subgenera or Sections). The accepted taxa are given serial numbers and virtually
all, except for the late additions, are illustrated to enable the reader, after using
the keys, to confirm the identification without reference to a herbarium, although
a herbarium should be consulted if one is available. The taxa are arranged
alphabetically within the genus (or in large genera, within subgenera) in order to
group related taxa and their illustrations together. I have tried to adopt generic
concepts that are useable by non-specialists and to avoid both microgenera that
are difficult to distinguish (e.g., in the Hymenophyllaceae) and composite genera
that contain readily distinguishable and presumably related groups of species that
can stand as genera (e.g., in the Polypodiaceae). The family descriptions are not
inclusive but concentrate on the characteristics of the genera and species of each
family that are found in the Flora area. I have adopted the name Tectariaceae to
replace the illegitimate and unconservable Aspidiaceae, which has in addition
been used formerly in a much broader and confusing circumscription; the name
Pteridaceae, which was incorrect when published because it included types of
earlier published family names, but which is correct in a narrow circumscription;
and the names Adiantaceae, Dicksoniaceae, and Dryopteridaceae, all of which
have recently been accepted for conservation by the Committee for Pteridophyta.

The genera are numbered sequentially throughout the Flora. These numbers
are used in the keys to the families and to the genera and also appear in the
running heads so the reader can turn readily from the keys to the generic
treatments. Likewise, all species and infraspecific taxa are numbered sequentially
so that the reader can turn readily from the keys to the text to the illustrations.
Those species and infraspecific taxa with a letter following the number are mostly
not illustrated.

Illustrations are based on specimens in the U. S. National Herbarium (US),
unless otherwise noted by another herbarium acronym. They are based on
specimens from the Flora area, unless another countiy is noted, ’^^en the country
is designated as Colombia, the specimens illustrated are not from the Choco. All
bar scales are 1 cm long, except as noted. Scales referring to two or more
illustrations are centrally placed and have full cross-bars at the ends. Those
referring to a single figure are placed near that figure and have half bars at the
ends.

Many pteridophytes, because of wide ranges, have accumulated many scientific
names. To save space and to prevent this Flora from being unwieldy, I have cited
only basionyms in the synonymies, but not transferred names having parenthetical
authors. Transferred names, at least for most species, are listed in the “Index
Fiiicum” (Christensen, 1905 “1906, 1913, 1917, 1934; Pichi Sermolli et ah,

8 INTRODUCTION

Map of Panama

INTRODUCTION 9

1965; Jarrett et al., 1985). Furthermore, I have replaced descriptions of the
species and infraspecific taxa with augmented keys; one or more phrases
describing the taxon may follow the strictly contrasting information in each portion
of the key that leads to a species or other infraspecific taxon. For the same
reason, specimens are cited only of the rarest taxa, in order to document their
presence in the Flora area. Some of the uncited specimens I have seen are in an
unpublished, largely typewritten data base that I used in preparing the Flora.

Literature cited in synonymies conforms largely to standard usage (Lawrence et
al., 1968; Stafleu & Cowan, 1976-1988), with some amplifications so that the
citations are intelligible without recourse to the standard works. I have included
page numbers (but not dates, even if they are different) for some widely held
reprints (Mettenius in Triana & Planchon, 1864; K. Presl, 1845a, 1851; Sodiro,
1897) and postprints (Presl, 1843, 1845b, 1847). I have also included some widely
held alternate titles, such as Fee’s memoir series, Jenman’s treatment of the ferns
and fern allies of Jamaica, Spring’s monograph of Lycopodium, and Domin’s
treatment of the pteridophyta of Dominica.

Types and lectotypes of many early names are based on plates published in
three pre-Linnaean works. Plunder’s “Traite” (1693), his “Description” (1705),
and Sloane’s “Voyage” (1707-25). The full titles of these works are cited below.

In the citation of specimens (mostly types), the word photo followed by a
number refers to the numbered series of photographs taken by C. V. Morton to be
found at US and distributed to other herbaria. For the type specimens of many
collectors, the usual herbarium of deposit is given, even when I have not seen the
specimens.

Because collectors have so frequently revisited a few choice collecting localities,
I have placed these localities and the provinces of Costa Rica and Panama on
maps (pp. 6, 8, and 10). For these localities, the province is not cited in the
discussions but can be learned by examining the maps. Localities in the Flora area
are cited from northwestern Costa Rica through eastern Panama to the southern
Choco.

The ranges cited in the discussion are based on herbarium specimens, both old
and new. Insofar as collecting has been adequate, they reflect the original range
of each taxon in the Flora area. In the more settled areas of this region, the
landscape has been virtually entirely altered by agricultural and other activities,
and most of the pteridophytes can no longer be Wnd. Their preservation in large,
well protected parks and nature preserves appears to be their best chance of
survival.

Special literature, mostly relatively recent monographs and revisions, is cited
under each genus. Since this work was begun, several general works have been
published that I have consulted repeatedly in matters of taxonomy, nomenclature,
and plant distribution. ITiese are Kramer (1978), Mickel and Beitel (1988),
Proctor (1977, 1985), Smith (1981, 1985), and Stolze (1976, 1981, and 1983).

My fieldwork in the Flora area was supported by the Smithsonian Research
Foundation, the National Geographic Society, and the Organization for Tropical
Studies. Herbarium visits (to A and GH, CR, DUKE, F, MO, NY, PMA, and UC)
were supported by the Smithsonian Institution. Preparation of the illustrations
was entirely supported by the Smithsonian Research Foundation and by the

10 INTRODUCTION

Map of the Choco

INTRODUCTION 11

Walcott Fund of the Department of Botany. Publication was generously
supported by the Smithsonian Institution. I deeply appreciate the support of all
these organizations. I am indebted to my colleagues D. S. Barrington (for the
treatment of Polystichum), L. E. Bishop, M. D. Correa A., F. R. Fosberg, L. D.
Gomez P., R. J. Hickey, R. E. Holttum, J. T. Mickel (for the treatment of
Elaphoglossum), J. D. Montgomery, R. C. Moran, the late C. V. Morton, D. H.
Nicolson, B. 011gaard, R. L. Petersen, G. R. Proctor, H. E. Robinson, A. R. Smith
(especially for his meticulous reading of the entire manuscript), R. G. Stolze, A. F.
Tryon, R. M. Tryon, Jr., W. H. Wagner, Jr., H. van der Werff, P. G. Windisch, and
J. J. Wurdack, for lending specimens or for help with problems of taxonomy and
nomenclature. Mary Monsma prepared the illustrations over a period of many
years, except for the drawings of Elaphoglossum scales done by Alice Tangerini;
and Brian Kahn, Gregory McKee, and Alice Tangerini assembled and lettered the
plates and prepared the maps. I am grateful for their work, which has added so
much to the usefulness of the Flora. Ellen Farr and Robert Sims provided much
needed advice on data bases and desk-top publishing, and Gregory McKee also
assisted with the desk-top publishing, for which I am most thankful.

CHRISTENSEN, C. 1905 - 1906. Index Filicum. H. Hagerup, Copenhagen.

— . 1913. Index Filicum. Supplementum (1906-1912). H. Hagerup, Copenhagen.

— — . 1917. Index Filicum. Supplement preliminaire pour les aim6es 1913, 1914, 1915, 1916.

Triers Bogtrykkeri, Copenhagen.

. 1934. Index Filicum. Supplementum Tertium pro Annis 1917-1933. H. Hagerup,

Copenhagen.

CRABBE, J. A., A. C. JERMY and J. T. MICKEL. 1975. A new generic sequence for the
pteridophyte herbarium. Fern Gaz. 11:141-162.

GOMEZ P., L. D. 1975a. The ferns and fern-allies of Cocos Island, Costa Rica. Amer. Fern J.
65:102-104.

. 1975b. Contribuciones a la pteridologia costarricense. VII. Pteridofita de la Isla de Cocos.

Brenesia 6:33 - 48.

JARRETT, F. M., et al. 1985. Index Filicum. Supplementum Quintum pro annis 1961-1975.
Clarendon Press, Oxford.

KRAMER, K. U. 1978. The Pteridophytes of Suriname. Uitgaven Natuurw. Studiekring Suriname
Nederl. Antillen 93:1-198.

LAWRENCE, G. H. M., et al. 1968. Botanico-Periodicum-Huntianum. Hunt Botanical Library,
Pittsburgh, PA.

LELLINGER, D. B. 1975. A phytogeographic analysis of Choc6 pteridophytes. Fern Gaz.
11:105-114.

1985. The distribution of Pancuna’s pteridophytes. Pp. 43 - 47 in W. G. D’Arcy & M.

Correa A., eds. The Botany and Natural History of Panama. Missouri Botanical Garden,
St. Louis, MO.

METTENIUS, G. in J. TRIANA and G. PLANCHON. 1864. Ann. Sci. Nat. Bot. V, 2:193-271.

Reprinted as Prodromus Florae Novo-granatensis, vol. 2, pp. 275 - 353. Masson, Paris.
MICKEL, J. T. and J. BEITEL. 1988. Pteridophyte Flora of Oaxaca, Mexico. New York Botanical
Garden, Bronx, NY.

PICHI SERMOLLI, R. E. G., et al. 1965. Index Fihcum. Supplementum Quartum pro Annis
1934- 1960. International Bureau for Plant Taxonomy and Nomenclature, Utrecht.
PLUMIER, C. 1693. Description des plantes de I’Amerique. L’imprimerie royale, Paris.

. 1705. Traite des fougeres de l’Am6rique. L’imprimerie royale, Paris.

12 INTRODUCTION

PRESL, K. B. 1843. Hymenophyllaceae. Pp. 1-70. Gottlieb Haase, Prague. Postprinted as Abh.
Konigl. Bohm. Ges. Wiss. V, 3:92- 162. 1845.

. 1845a. Botanische Bemerkungen. Abh. Konigl. Bohm. Ges. Wiss. V, 3:431-584.

Reprinted as Botanische Bemerkungen. Pp. 1-154. 1846. Gottlieb Haase, Prague.

.1845b. Supplementum Tentaminis Pteridographiae. Pp. 1-119. Amadei Haase, Prague.

Postprinted as Abh. Konigl. Bohm. Ges. Wiss. V, 4:261-380. 1847.

. 1847. Die Gefassbiindel im Stipes der Farm. Pp. 1-48, t. 1-7. Gottlieb Haase, Prague.

Postprinted as Abh. Konigl. Bohm. Ges. Wiss. V, 5:307 - 356, 1. 1-7. 1848.

. 1851. Epimeliae Botanicae. Abh. Konigl. Bohm. Ges. Wiss. V, 6:361-624, t. 1-15.

Reprinted as Epimeliae Botanicae. Pp. 1-264, 1. 1-15. 1851. Amadei Haase, Prague.
PROCTOR, G. R. 1977. Flora of the Lesser Antilles, vol. 2. Pteridophyta. Arnold Arboretum,
Jamaica Plain, MA.

. 1985. Ferns of Jamaica. British Museum (Natural History), London.

SLOANE, H. 1707 - 25. A Voyage to the Islands Madera, Barbados, Nieves, S. Christophers and
Jamaica. British Museum, London.

SMITH, A. R. 1981. Flora of Chiapas, part 2. Pteridophytes. California Academy of Sciences, San
Francisco, CA.

. 1985. Pteridophytes of Venezuela, an Annotated List, Privately published by the author.

SODIRO, A. 1897. Cryptogamae Vasculares Quitenses. Universidad Central, Quito, Ecuador.
[Title page date 1893.]

STAFLEU, F. A. and R. S. COWAN. 1976-1988. Taxonomic Literature, ed. 2. 7 vols. Bohn,
Scheltema & Holkema, Utrecht; all but vol. 1 with W. Junk, The Hague.

STOLZE, R. G. 1976. Ferns and Fern Allies of Guatemala, part 1. Ophioglossaceae through
Cyatheaceae. Fieldiana, Bot. 39:i-v + 1-130.

. 1981. Ferns and Fern Allies of Guatemala, part II. Polypodiaceae. Fieldiana, Bot. N. S.,

6:i-ix + 1-522.

. 1983. Ferns and Fern Allies of Guatemala, part III. Marsileaceae, Salviniaceae, and the

Fern Allies. Fieldiana, Bot. N. S., 12:i-iii + 1-91.

LIST OF FAMILIES 13

LIST OF FAMILIES, GENERA, AND SUBGENERA OR SECTIONS

PSILOTACEAE

1. Psilotum
LYCOPODIACEAE

2. Lycopodium
subg. Lycopodium
subg. Cernuistachys
subg. Selago
SELAGINELLACEAE

3. Selaginella
ISOETACEAE

4. Isoetes

EQUISETACEAE

5. Equisetum
OPHIOGLOSSACEAE

6. Botrychium

7. Ophioglossum
MARATTIACEAE

8. Marattia

9. Danaea
OSMUNDACEAE

10. Osmunda
PLAGIOGYRIACEAE

11. Plagiogyria
SCHIZAEACEAE

12. Schizaea

13. Actinostachys

14. Lygodium

15. Anemia
PARKERIACEAE

16. Ceratopteris
ADIANTACEAE

17. Llavea

18. Cheilanthes

19. Aleuritopteris

20. Adiantopsis

21. MildeUa

22. Notholaena

23. Pellaea

24. Doryopteris

25. Anogramma

26. Pityrogramma

27. Hemionitis

28. Gymnopteris

29. Bommeria

30. Jamesonia

31. Eriosorus

32. Adiantum

VITTARIACEAE

33. Antrophyum

34. Anetium

35. Hecistopteris

36. Ananthacorus

37. Vittaria
PTERIDACEAE

38. Pteris

39. Neurocallis

40. Acrostichum
LOXSOMACEAE

41. Loxsomopsis
HYMENOPHYLLACEAE

42. Hymenophyllum
subg. Hymenophyllum
subg. Leptocionium
subg. Mecodium

43. Trichomanes
subg. Achomanes
subg. Didymoglossum
subg, Pachychaetum
subg. Trichomanes
GLEICHENIACEAE

44. Gleichenia

45. Dicranopteris
POLYPODIACEAE

46. Pleopeltis

47. Pseudocolysis

48. Microgramma

49. Neurodium

50. Dicranoglossum

51. Niphidium

52. Hyalotrichopteris

53. Campyloneurum

54. Phlebodium

55. Polypodium

subg. Goniophlebium
subg. Marginaria
subg. Polypodium

56. Pecluma

57. Grammitis
sect. Cryptosorus
sect. Grammitastrum
sect. Grammitis
sect. Xiphopteris

58. Glyphotaenium

59. Cochlidium

60. Loxogramme

LIST OF FAMILIES 14

CYATHEACEAE

61. Metaxya

62. Lophosoria

63. Cnemidaria

64. Cyathea

65. Alsophila

66. Sphaeropteris
DICKSONIACEAE

67. Dicksonia

68. Culcita

DENNSTAEDTIACEAE

69. Dennstaedtia

70. Costaricia

71. Hypolepis

72. Paesia

73. Pteridium

74. Histiopteris

75. Lonchitis

76. Blotiella

77. Saccoloma

78. Lindsaea

79. Ormoloma

80. Odontosoria
THELYPTERIDACEAE

81. Thelypteris

subg. Amauropelta
subg. Cyclosorus
subg. Goniopteris
subg. Macrothelypteris
subg. Meniscium
subg. Steiropteris
ASPLENIACEAE

82. Asplenium
sect. Asplenium
sect. Sphenopteris

83. Loxoscaphe

84. Schaffneria
ATHYRIACEAE

85. Athyrium

86. Diplazium

subg. Anisogonium
subg. Diplazium

87. Hemidictyum
WOODSIACEAE

88. Cystopteris

89. Woodsia
TECTARIACEAE

90. Ctenitis

sect. Amplae
sect. Ctenitis
sect. Hirtae

91. Megalastrum

92. Triplophyllum

93. Lastreopsis

94. Tectaria

95. Cyclopeltis
DRYOPTERIDACEAE

96. Didymochlaena

97. Polystichum

98. Cyrtomium

99. Arachniodes

100. Polybotrya

101. Olfersia

102. Maxonia

103. Dryopteris

104. Cyclodium

105. Stigmatopteris
LOMARIOPSIDACEAE

106. Bolbitis

107. Lx)magramma
118. Lomariopsis

109. Elaphoglossum
sect. Amygdalifolia
sect. Decorata
sect. Elaphoglossum
sect. Eximia

sect. Lepidoglossa
sect. Polytrichia
sect. Squamipedia
sect. Setosa
sect. Undulata
DAVALLIACEAE

110. Oleandra

111. Nephrolepis
BLECHNACEAE

112. Blechnum
subg. Blechnum
subg. Lomaria

113. Salpichlaena

114. Woodwardia
MARSILEACEAE

115. Marsilea
SALVINIACEAE

116. Salvinia
AZOLLACEAE

117. Azolla

KEY TO THE FAMILIES 15

KEY TO THE FAMILIES

1. Plants bearing expanded fronds (megaphylls) with several vascular strands (reduced to or nearly
to often winged stipes with terminal, sporangium-bearing sorophores in some genera of the
Schizaeaceae and to minute, crowded fronds in the aquatic families Azollaceae and Salviniaceae), if
grass-like, not bearing a single, large sporangium in a swollen base..6.

1. Plants bearing needle-like or scale-like leaves (microphylls) with a single vascular strand, or the
leaves rush-like.. 2.

2(1). Stems and branches obviously jointed, usually fluted, hollow, often harsh or brittle from silica
deposited in the cells; leaves borne in a whorl above a slightly swollen sheath at each node, the tips
often scarious and deciduous.

EQUISETACEAE— 5. Equisetum

2(1). Stems and branches not obviously jointed, rarely fluted or hollow, lacking silica; leaves
spirally arranged along the stem or with 2 lateral and 2 smaller, dorsal rows of leaves.. 3.

3(2). Leaves inconspicuous, scale-like, borne on nearly naked, repeatedly dichotomously branched
stems; sporangia tripartite, borne in the axil of a minute sporophyll.

PSILOTACEAE— 1. Psilotum

3(2). Leaves conspicuous, not scale-like, more or less covering the stems; sporangia not
tripartite.. 4.

4(3). Leaves grass-like, borne in a cluster on a compact, 2- or 3-lobed corm, bearing a single, large
sporangium in a swollen base.

ISOETACEAE— 4. Iso^es

4(3). Leaves not grass-like or borne on a corm, not bearing a single, large sporangium in a swollen
base..5.

5(4). Sporangia borne in leaf axils along distal portions of the stems or in cylindrical strobili at
branch tips; leaves in most species acicular to narrow and monomorphic (equally disposed around
the stems), often coriaceous.

LYCOPODIACEAE“2. Lycopodium

5(4). Sporangia borne in flattened or quadrangular strobili at branch tips; leaves in most species
broad and dimorphic (in 2 lateral and 2 smaller, dorsal rows of leaves), usually membranaceous.

SELAGINELLACEAE— 3. Sela^nella

6(1). Plants epiphytic, epipetric, or terrestrial; if rooted in mud, then more than 30 cm long.. 10.

6(1). Plants floating in water or sometimes rooted in mud at the edge of ponds or streams or in
dried-up pools; plants less than 30(50) cm long.. 7.

7(6). Laminae at least 6 cm long, pinnately lobed or divided; stipes commonly inflated; laminae
dimorphic, the fertile laminae with narrow segments held above the broader, shorter sterile laminae.

PARKERIACEAE— 16. Ceratopteris

7(6). Laminae less than 6 cm long, not pinnately lobed or divided; stipes obsolete or filiform;
l2miinae monomorphic. .8.

8(7). Stipes filiform, several cm long; laminae quadrifid; sporocarps conspicuous, up to ca. 5 mm
long, hard, short-stalked, inserted at or near the base of the stipe.

MARSILEACEAE-"115. Marsilea

8(7). Stipes obsolete; laminae simple, entire; sporocarps inconspicuous, concealed by the
laminae..9.

9(8) Laminae on the water’s surface oval, mostly 5-15 mm long, with obvious, short hairs on the
surface exposed to the air, usually bright green.

SALVINIACEAE— 116. Salvinia

9(8). Laminae all more or less ovate, mostly less than 1 mm long, lacking hairs on the surface
exposed to the air, often reddish, always crowded along repeatedly branched stems.

AZOLLACEAE— 117. Azolla

16 KEY TO THE FAMILIES

10(6). Sporangia fused laterally into elongate or linear, brownish or gray, short-stalked or sessile
synangia opening by a double row of pores.

MARATTIACEAE — 8. Marattia and 9. Danaea

10(6). Sporangia discrete, not fused into synangia.. 11.

11(10). Sporangia small, mostly less than 0.3 mm in diam., dehcate, thin-walled, usually brownish
at maturity and in age, usually opening into 2 deciduous, irregular portions by the action of a vertical,
oblique, or rarely terminal, thick- walled annulus, usually containing 64 spores (leptosporangia),
usually on expanded lamina tissue, usually borne in separate groups (sori)..13.

11(10). Sporangia large, mostly more than 0.3 mm in diam., firm, thick-walled, yellowish at
maturity, sometimes becoming brownish in age, usually opening into 2 persistent, hemispherical or
clamshell-shaped halves without the action of an annulus, usually containing more than 100 spores
(eusporangia), usually scattered on naked axes.. 12.

12(11). Laminae soft, with little supporting tissue, up to 40 cm long, the fertile portion inserted
near the base of the sterile portion; sporangia producing thousands of spores.

OPHIOGLOSSACEAE — 6. Botrychium and 7. Ophioglossum

12(11). Laminae firm, with supporting tissue, 50-150 cm long, the fertile portion apical or fertile
fronds separate from the sterile ones; sporangia bearing hundreds of spores.

OSMUNDACEAE-10. Osmunda

13(11). Plants lacking both hairs and scales. Laminae pinnatisect, dimorphic, the sterile pinnae
linear, serrulate toward the acute to acuminate apex, the fertile piimae narrower, the sporangia in a
marginal coenosorus protected by a broad, scarious, continuous, revolute false indusium.

PLAGIOGYRIACEAE— 11. Plagiogyria

13(11). Plants, or some parts of them, bearing hairs or scales or both.. 14.

14(13). Sporangia borne in 1 or 2 rows on sorophores terminating grass-like or fan-like fronds (in
Actinotstachys and Schizaed) or on the lateral margins of the pinnules of high-climbing fronds (in
Lygodium) or in 2 rows on dimorphic, branched, elongate, erect pairs of basal pinnae {in Anemia);
sporangia ovoid or pear-shaped with an apical or oblique, patch-like annulus.

SCHIZAEACEAE— 12. Schizaea, 13.Actinostachys, Id Lygodium ^
and 15. Anemia

14(13). Sporangia not borne on sorophores or specialized basal pinnae..l5.

15(14). Laminae membranaceous (1 cell thick between the veins in most species); indusia
(involucres) tubular or equally bivalvate.

HYMENOPHYLLACEAE — 42. Hymenophyllum and 43. Trichomanes

15(14). Laminae mostly papery to leathery (at least (2)3 cells thick between the veins); indusia not
tubular (except in Loxsomopsis) or equally bivalvate.. 16.

16(15). Indusia borne at the ends of broad, marginal teeth, cylindrical or urn-shaped and often
slightly constricted at the mouth, ca. 1 mm in diam. and 1.5 mm long.

LOXSOMACEAE — 41. Loxsomopsis

16(15). Indusia not as above.. 17.

17(16). Sori long or rarely short, in a single continuous or discontinuous line on each side of the
costa or costule, protected by a continuous, usually ample, often deciduous, introrse indusium facing
the costa or costule.

BLECHNACEAE — 112. Blechnum, 113. Salpichlaena, and 114. Woodwardia

17(16). Indusia not as above..l8.

18(17). Sori not elongate along the lateral veins or, if so, then not protected by a narrow, usually
persistent indusium..20.

18(17). Sori elongate along the lateral veins, on one or both sides of the vein or on one side and
hooking over to the other side, protected by a usually persistent, narrow indusium.. 19.

19(18). Rhizome scales strongly clathrate, the clear lumina readily visible with a hand lens, the
cells mostly rather short, 1-2 times longer than wide; plants mostly epiphytic.

ASPLENIACEAE— 82.yl5/7/enm/n, 83. Loxoscaphe, and 84. Schaffheria

KEY TO THE FAMILIES 17

19(18). Rhizome scales not clathrate, the obscure lumina not readily visible with a hand lens, the
cells mostly very long, 3-6 times longer than wide; plants mostly terrestricd.

ATHYRIACEAE™ 86. Diplaziuniy and 87. Hemidictyum

20(18). Sporangia organized into discrete sori visible as points or lines, indusiate or exindusiate,
dorsal or marginal, the fronds usually monomorphic..25.

20(18). Sporangia apparently or truly acrostichoid, not visible as points or lines, exindusiate (except
Cydodium and Maxonia in the Dryopteridaceae), partially or fully dimorphic (except Anetium in the
Vittariaceae and Cydodium in the Dryopteridaceae, and Llavea, with a false marginal indusium, in
the Adiantaceae), the fertile area confined to specialized fronds or pmnae..21.

21(20). Laminae simple, elliptic or elliptic-oblanceolate, very thin, with venation reticulate; fronds
monomorphic, the sporangia scattered thinly over the abaxial surface of the fertile laminae.

VITTARIACEAE — 34, Anetium

21(20). Laminae pinnate or more divided; fronds dimorphic, the sporangia scattered thickly over
the usually reduced abaxial surface of the fertile laminae..22.

22(21). Sterile laminae coriaceous, the veins a reticulum of elongate hexagons lacking included
veinlets; plants terrestrial, usually in swamps or along estuaries or coasts.

PTERIDACEAE— 39. Neurocallis and 40. Acrostichum

22(21). Sterile laminae papyraceous, the veins free, casually anastomosing, or forming an irregular
reticulum with included veinlets; plants mostly hemiepiphytic or epiphytic, usually in forests..23.

23(22). Fertile segments linear, incurved, bearing scarious, continuous, marginal false indusia;
sterile fronds 2-3-pinnate, the segments ovate-lanceolate, long-stalked.

ADIANTACEAE— 17. Llavea

23(22). Plants not as above..24.

24(23). Pinnae articulate to a usually narrowly alate rachis or, if the laminae simple or
dichotomously branched, the stipes usually articulate near the rhizome; rachises and other axes
terete, flattened or grooved, the grooves never provided with short hairs.

LOMARIOPSIDACEAE— 106. Bolbitis, 107. Lomagramma,

108. Lomariopsis, and 109. Elaphoglossum

24(23). Pinnae not articulate, the rachises exalate; rachises and other axes mostly grooved, the
grooves usually provided with short hairs.

DRYOPTERIDACEAE— 100. Pofybotrya, 101. Olfersia, and 102. Maxonia

25(20). Sori surficial, usually round to elongate, rarely linear (if linear, not in a submarginal
groove), not protected by a false indusium..36.

25(20). Sori marginal or nearly so, often elongate or linear, often protected by a reflexed and
modified segment margin (false indusium). .26.

26(25). Sporangia borne only on the abaxial surface of the continuous or discontinuous false
indusia.

ADIANTACEAE— 32. Adiantum

26(25). Sporangia never borne on the abaxial surface of the false indusia..27.

27(26), Indusia cup-like, formed by a conspicuous, extrorse true indusium and a modified or
unmodified portion of the frond margin.

DICKSONIACEAE— 67. Dicksonia and 68. Culdta

27(26). Indusia not as above..28.

28(27). Indusia extrorse, cup-like or shallow and elongate laterally, marginal or slightly
submarginal; frond margins not modified, rarely reflexed.

DENNSTAEDTIACEAE— 69. Dennstaedtia, 70. Costarida, 77. Saccoloma,

78. Lindsaea, 79. Omtoloma, and 80. Odontosoria

28(27). Indusia not as above..29.

29(28). Fronds not simple and lmear..32.

29(28). Fronds simple, linear or nearly so. Sori marginal and linear, often in a groove parallel to

the margin (the margin sometimes reflexed). .30.

18 KEY TO THE FAMILIES

30(29). Sori protected by a linear, scarious, extrorse indusium.

TECTARIACEAE — 94. Tectaria
30(29). Sori not protected by an extrorse indusium. .31.

31(30). Rhizome scales strongly clathrate.

VlTIARlAC^AE-—2)6.Ananthacoms and 37. Vittaria
31(30). Rhizome scales not strongly clathrate.

POLYPODLACEAE — 49. Neurodium
32(29). Plants in aquatic habitats, rooted in mud.

PARKERIACEAE — 16. Ceratopteris
32(29). Plants not as above..33.

33(32). Fronds several times dichotomously divided, epiphytic.

POLYPODLACEAE— 50. Dicranoglossum
33(32). Fronds pinnately or pedately divided, usually terrestrial or epipetric..34.

34(33). Fronds usually less than 0.5(0.75) m long, often linear or oblong; stipes commonly wiry,
stramineous, dark brown, reddish-brown, or blackish; rhizomes thin, usually short-creeping or
multicipital, scaly.

ADIANTACEAE— 18. Cheilanthes, 19. Aleuritopteris, Ud.Adiantopsis
21. Mildella, 23. Pellaea, 24. Doryopteris, and 30. Jamesonia
34(33). Fronds usually at least (0.5)1 m long, commonly deltate and 2-3-pinnate; stipes commonly
thick, pale brown; rhizomes thick, long-creeping (short-creeping to erect in Pteris), hairy (mixed with
scales in Histopteris).. 35.

35(34). Rhizomes long-creeping; lamina axes usually provided with short, stout hairs; laminae
often dark green (glaucous in Histiopteris); sori discontinuous (subcontinuous in Lonchitis,
continuous in Histiopteris).

DENNSTAEDTIACEAE— 71. Hypolepis, 72. Paesia^ 73. Pteridium,

74. Histiopteris, 15. Lonchitis, and 76. Blotiella

35(34). Rhizomes short-creeping to erect; lamina axes mostly glabrous; laminae often pale to
bright green; sori continuous, usually from the sinus to near the apex of the segments.
PTERIDACEAE— 38. Pteris
36(25). Sori round or nearly so, indusiate or not..43.

36(25). Sori elongate to linear, exindusiate..37.

37(36). Fronds simple, entire to shallowly lobed; sori linear to elongate, often in grooves..40.

37(36). Fronds deeply lobed to pinnate or more divided; sori elongate, rarely linear, never in
grooves.. 38.

38(37). Adaxial surface of the costae (and often the rachis) grooved and provided with short,
acicular hairs.

THELYPTERIDACEAE— 81. Thelypteris
38(37). Adaxial surface of the costae not provided with short, acicular hairs.. 39.

39(38). Fronds simple or with up to 3 pairs of elliptic lobes; stipes narrowly alate for much of then-
length.

POLYPODLACEAE— 47. Pseudocolysis
39(38). Fronds not as above.

ADIANTACEAE — 22. Notholaena, 25. Anogramma, 26. Pityrogramma,

21. Hemionitis, 28. Gymnopteris, 29. Bommeria, and 31. Eriosorus
40(37). Sori forming elongate, polygonal areolae without included veinlets.

VITTARIACEAE — 33. Antrophyum
40(37). Sori elongate or linear, but not forming polygonal areolae..41.

41(40). Laminae dimorphic, the fertile ones linear, the sterile ones elliptic-lanceolate.

POLYPODLACEAE— 46. Pleopeltis
41(40). Laminae monomorphic, linear, oblong, or oblanceolate..42.

KEY TO THE FAMILIES 19

42(41). Laminae linear with the sori in several approximate grooves or laminae subflabellate and
lacerate at the apex with the sori not in grooves.

VITTARIACEAE-— 33. Antrophyum and 35. Hecistopteris

42(41). Laminae not as above.

POLYPODIACEAE— 58. Glyphotaenium, 59. Cochlidium, and 60.

Loxogramme

43(36). Rhizome an erect, massive (up to ca. 10 cm in diam.) caudex often forming a tree-like
trunk, bearing large fronds in a whorl or compressed spiral; sporangia short-stalked, the capsule
angular, the annulus oblique.

CYATHEACEAE~61. Metaxya, 62. Lophosoria, 63. Cnemidaria,

64. Cyathea, 65. Alsophila, and 66. Sphaeropteris

43(36). Rhizome erect or not, not massive, never forming a tree-like trunk; sporangia long-stalked,
the capsule subspherical, the annulus vertical.. 44.

44(43). Sori exindusiate..49.

44(43). Sori indusiate, the indusia if deciduous leaving a gap among the sporangia or present as
scale-like shreds..45.

45(44). Indusia cup-shaped or scale-like, usually fragmenting or deciduous in age.

WOODSIACEAE--88. Cystopteris and 89. Woodsia

45(44). Indusia peltate or reniform..46.

46(45). Pinnae articulate to the rachis, often deciduous in age, or laminae simple, the stipes long,
wiry, articulate to the rhizome, often with an elongate phyllopodium.

DAVALLIACEAE — 110. Oleandra and 111. Nephrolepis

46(45). Pinnae not articulate to the rachis or deciduous, and laminae not simple, stipes not long,
wiry, or articulate to the rhizome.. 47.

47(46). Rachises adaxiaUy sulcate, the sulcus continuous with the adaxially sulcate costae of the
lateral pinnae; edges of the sulcae usually glabrous.

DRYOPTERIDACEAE— 96. Didymochlaena, 97. Polystichum, 98. Cyrtomium,
99.Arachniodes, 103. Dryopteris, 104. Cyclodium, and 105.

Stigmatopteris

47(46). Rachises adaxially terete or, if sulcate, the sulcus not continuous with any adaxially sulcate
costae of the lateral pinnae; edges of the sulcae with short to rarely long, multicellular, septate hairs
or short, unicellular, needle-like hairs..48.

48(47). Hairs on the adaxial surface of the rachis usually short, unicellular, and acicular; laminae
mostly pinnate-pinnatifld, sometimes merely pinnate; vascular bundle above the stipe base one, U-
shaped.

THELYPTERIDACEAE— 81. Thelypteris

48(47). Hairs on the adaxial surface of the rachis rcU'ely short, unicellulcu", and aciculair, often long,
multicellular, often septate, rarely absent; laminae often not pinnate-pinnatifid, commonly 2-pinnate
or more divided; vascular bundles above the stipe base several.

TECTARIACEAE--"90. Ctenitis, 91. Megalastrum, 92. Triplophyllum,

93. Lastreopsis, 94. Tectaria, and 95. Cyclopeltis

49(44). Laminae usually appearing to be dichotomously branched, with pinnatisect or rarely
retroflexed ultimate divisions of the fronds and resting buds in the axils of at least some dichotomies.
Sporangia always very few (commonly ca. 5 - 12) in each sorus.

GLEICHENIACEAE— 44. Gleichenia and 45. Dicranopteris

49(44). Laminae rarely appearing to be dichotomously branched, never with resting buds in the
axils of some dichotomies.. 50.

50(49). Fronds pinnate, elongate, recumbent or scrambling on other vegetation, commonly in
sunny locations; rachises, and often the costae and costules, sparingly spiny and/or sparsely to
densely pubescent and rough to the touch.

DENNSTAEDTIACEAE— 71. Hypolepis

20 KEY TO THE FAMILIES

50(49). Fronds rarely greatly elongate and recumbent or scrambling; axes never spiny..51.

51(50). Laminae simple, piimatifid, or pinnatisect, often less than 30 cm long.

POLYPODIACEAE— 46. Pleopeltis, 48. Microgramma, 50. Dicranoglbssum,

51. Niphidium, 52. Hyalotrichopteris, 53. Campy loneurum, 54. Phlebodium,
55. Polypodium, 56. Pecluma, 57. Grammitis, 58. Glyphotaenium, and
59. Cochlidium

51(50). Laminae pinnate or more divided, usually more than 30 cm long. .47.

POLYPODIACEAE— Genera 55. Polypodium and 57. Grammitis

PSILOTACEAE: 1.PS1LOTUM 21

FLORISTIC TREATMENTS
PSILOTACEAE

Monotypic in the Flora area; see description oiPsilotum.

1. PSILOTUM Swartz

Plants epiphytic or less commonly epipetric, terrestrial, or on old masor^;
rhizomes subterranean, short-creeping, branching, lacking true roots but bearing
short, brownish rhizoids; aerial stems loosely clustered, simple at the base, 3-7
times dichotomous distally, flattened or triangular in cross-section; appendages on
the stems minute, few, alternate, scale-like, the sterile ones acicular, the fertile
ones forked; synangia sessile or nearly so, globular, usually tripartite, yellowish,
dehiscing vertically; spores of 1 kind.

Pantropical; 2 species. A primitive genus (and family) not closely related to the
true ferns.

COOPER-DRIVER, G. 1977. Chemical evidence for separating the Psilotaceae from the Fihcales.
Science 198:1260-1261.

WAGNER, W. H., Jr. 1977. Systematic implications of the Psilotaceae. Brittonia 29:54 - 63.

1. Stems strongly flattened in cross-section, 1.5 -2.5 mm wide. Fronds 3-5 times dichotomous,
15 - 40 cm long.

1. P. complanatum

1. Stems round or angled in cross-section, 0.5 -1.5 mm in diam. Fronds 3-7 times dichotomous,
10 - 35 cm long.

2. P. nudum

1. Psilotum complanatum Swartz, J. Bot. (Schrader) 1800(2):110. 1801; Syn. Fil.

188, 414, t. 4, f. 5. 1806.

Psilotum complanatum var. latissimum Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:271.
1849. TYPE LOCALITY: Mexico.

Bemhardia califomica K. Muell. Bot. Zeitimg (Berlin) 14:222, f. 3 - 5. 1856. TYPE: California
[presumably Mexico!, Deppe (B not seen).

Bemhardia schiedeana K. Muell. Bot. 2^itung (Berlin) 14:222, f. 9-11. 1856. TYPE: Hacienda
de la Laguna, Jalapa, Edo. Veracruz, Mexico, Schiede (B not seen).

TYPE: Jamaica, Swartz (B-Hb. Willd. 19437 not seen microfiche S. I. Library),
according to Proctor (Ferns Jamaica 15. 1985).

Plants epiphytic, in the Flora area known only from the Rio Sucio, selvas de
Sarapiqui, Pcia. Heredia, 125 m {Gomez 473, CR) and from an unlocalized
collection made in Costa Rica (March 1908, Ridgway, US). Also from Cuba,
Jamaica, Hispaniola, Mexico to El Salvador, and Venezuela, Colombia, and
Peru.

2. Psilotum nudum (L.) Pal. Beauv. Prodr. Aeth6ogram. 112. 1805.

Lycopodium nudum L. Sp. PI. 2:1100. 1753. TYPE: Based on a specimen with “nudum 2” in the
hand of Linnaeus (LINN 1257.1 not seen microfiche S. I. Library).

Hoffmannia aphylla Willd. Mag. Bot. (Romer & Usteri) 2(6):17. 1789, nom. superfl. An

illegitimate renaming of L. nudum L., and so based on the type of that name.

Psilotum triquetrum Swartz, J. Bot. (Schrader) 1800(2): 109. 1801, nom. superfl. TYPE: A

renaming of L. nudum L., cmd so based on the type of that name.

Bemhardia dichotoma Willd. ex Bernh. J. Bot. (Schrader) 1800(2): 132. 1801, nom. superfl,

TYPE: An illegitimate renaming of L. nudum L., and so based on the type of that name.

22 PSILOTACEAE: 1. PSILOTUM

PSILOTACEAE: 1.PSILOTUM 23

Psilotum floridanum Michx. FI. Bor.-Amer. 2:281. 1803. TYPE: Florida, Michaux (P not seen),
examined by Morton (Amer. Fern J. 57:180. 1967).

Bemhardia antillarum K. Muell. Bot. Zeitung (Berlin) 14:234. 1856. SYNTYPES: Jcimaica,
Bertero in 1821 (B not seen); Martinique, Sieber FI Mart. 58 (B not seen); Trinidad, Sieber FI. Trin.
200 (B not seen); and Est. Rio de Janeiro, Brazil, Langsdorjf (B not seen).

Bemhardia deppeana K. Muell. Bot. Zeitung (Berlin) 14:235. 1856. TYPE: California

[presumably Mexico], Deppe (B not seen).

Psilotum domingense Gand. Bull. Soc. Bot. France 66:306. 1919. TYPE: Sto. Domingo,

Hispaniola, Bory (P not seen).

Plants epiphytic or epipetric, at 0-100 m elevation, from the Atlantic coastal
plain of Costa Rica and Panama. Also throughout tropical America except the
Guianas.

FIGS. 1“10. Psilotum djid Lycopodium. FIG. 1. Frond ofF. complanatum, Ridgway. FIG. 2. Frond
of F. nudum, Hart 7a. FIG. 3. Pl^mt of L. clavatum war. aristatum, Lellinger 955. FIG. 4. Plant of L.
contiguum, Allen 5416. FIG. 5. Plant of L. jussiaei, Mickel 3243. FIG. 6. Plant of L. magellanicum,
Ekman H-13666, Hispaniola. FIG. 7. Plant of L. thyoides. Woodruff 29. FIG. 8. Plant of L.
alopecuroides var. integerrimum, Proctor 25304, Guatemala. FIG. 9. Plant of L. carolinianum var.
meridionale, Lehmann 8911, Colombia. FIG. 10. Branch apex of L. cemuum var. cemuum, Maxon
8351.

24 LYCOPODIACEAE: 2. LYCOPODIUM

LYCOPODIACEAE

Monotypic in this treatment; see description oi Lycopodium.

2. LYCOPODIUM L.

Plants terrestrial or epiphytic; rhizomes very short and inconspicous or wide-
creeping and conspicuous, provided with tufts of dichotomously branched roots at
the nodes and with sparse leaves; aerial stems simple, dichotomously branched, or
apparently pinnately branched, usually glabrous, seldom hairy, the leaf-like
appendages arranged in a series of spirals, minute or not, 1-nerved, mostly linear,
narrowly lanceolate, or narrowly lanceate, all alike or the fertile ones slightly to
greatly different, often forming discrete strobili; sporangia axillary, sessile or short-
stalked, yellow, reniform or subglobose; spores of 1 kind.

Pantropical with some temperate species; ca. 500 species. The infrageneric
taxonomy of this genus has not yet been settled, and many tropical species are
poorly known. The subgenera adopted here follow 011gaard’s concepts, which
emphasize branching patterns and spore morphology.

0LLGAARD, B. 1975. Studies in Lycopodiaceae, I. Observations on the structure of the
sporangium wall. Amer. Fern J. 65:19-27.

. 1979. Studies in Lycopodiaceae, II. The branching patterns and infrageneric groups of

Lycopodium sensu lato. Amer. Fern J. 69:49-61.

ROLLERI, C. 1981. Sinopsis de las especies de Lycopodium L. (Lycopodiaceae: Pteridophyta) de la
seccion Crassistachys Herter. Revista Museo La Plata, Secc. Bot., 13:61 - 113.

WILCE, J. H. 1972. Lycopod spores, I. General spore patterns and the generic segregates of
Lycopodium. Amer. Fern J. 62:65-79.

1. Rhizomes not wide-creeping; erect or pendent stems regularly dichotomous, usually not forming
distinct strobili; spores pitted or grooved (subg. Selago).. 13.

1. Rhizomes wide-creeping, rooted and bearing erect stems at intervals; erect stems unbranched,
pinnately branched, or subdichotomous, usually forming distinct strobili; spores broadly ridged or
sharply reticulate..2.

2(1). Erect stems subdichotomously branched, the strobili usually stalked; spores reticulate (subg.
Lycopodium). .9.

2(1). Erect stems pinnately branched or unbranched, rarely subdichotomously branched, the
strobili usually sessile; spores broadly ridged (subg. Cemuistachys)..3.

3(2). Erect branches unbranched or subdichotomously branched, up to 20(40) cm long..4.

3(2). Erect branches piimately branched, often tree-like, up to ca. 100 cm long.. 5.

4(3). Rhizomes appressed to the soil; strobili solitary on subdichotomously branched erect
branches up to 30 cm long; fertile leaves wider than the sterile ones, the strobili stalked, distinct.

9. L. carolinianum var. meridionale

4(3). Rhizomes arching above the surface of the soil; strobili solitary on unbranched erect
branches up to 40 cm long; fertile leaves only shghtly different from the sterile ones, the strobili
indistinct.

8. L. alopecuroides var. integerrimum

5(4). Leaves and/or stems decidedly setose; leaves on main stems appressed..7.

5(4). Leaves and stems not setose; leaves on main stems spreading..6.

6(5). Leaves fine, flexible, ca. 0.2 mm wide. Ultimate branches visible between the distant leaves;
stems including the leaves 3-6 mm in diam.

10. L. cemuum var. cemuum

6(5). Leaves coarse, stiff, ca. 0.5 mm wide. Ultimate branches usually obscured by the crowded
leaves; stems including the leaves 3-5(7) mm in diam.

12. L. pendulinum

LYCOPODIACEAE: 2. LYCOPODIUM 25

7(5). Leaves nearly appressed, imbricate, the stems not visible; ultimate fertile branches 3-8 cm
long.

13. L. riofrioi

7(5). Sterile leaves spreading and incurved, distant, the stems visible; ultimate fertile branches
0.5 -2(3) cm long..8.

8(7). Leaves densely ciliate; largest branchlets including the leaves mostly less than 3 mm wide.

14. L. trianae

8(7). Leaves sparsely or not ciliate; largest branchlets including the leaves more than 3 mm wide.

11. L. cemuum var. curvatum

9(2). Sterile leaves all in many longitudinal rows equally arranged around the stems..ll.

9(2). Sterile leaves, at least the larger ones, in 2 longitudinal rows in a single plane.. 10.

10(9). Leafy branches up to 3.5 mm wide; strobili usually 4 in a group. Fertile leaves in the strobili
broadly ovate-lanceolate, 1.5 - 2 times longer than wide.

7. L. thyoides

10(9). Leafy branches 3-6 mm wide; strobili sohtary. Fertile leaves in the strobili lanceolate to
almost diamond-shaped, 3-4 times longer than wide.

5. L. jussiaei

11(9). Leaves acuminate at the apex; rhizomes hypogeous. Upright portions of the plants up to 15
cm long; strobili solitary or paired.

6. L. magellanicum

11(9). Leaves with a prolonged, filiform apex; rhizomes hypogeous or epigeous..l2.

12(11). Aerial branches widely dichotomous, spreading; strobili several, long-stalked; rhizomes
predominently epigeous.

3. L. clavatum var. aristatum

12(11). Aerial branches narrowly dichotomous, strictly erect; strobili solitary, sessile; rhizomes
predominently hypogeous.

4. L. contiguum

13(1). Sterile leaves not coriaceous, lax to somewhat stiff, ascending, spreading, or recurved away
from the stem..21.

13(1). Sterile leaves coriaceous, stiff, strongly ascending (spreading in L. goudotii), appressed or
incurved along the stem. Sterile leaves ca. (0.5)1 cm long.. 14.

14(13). Plants terrestrial, erect; leafy stems 5-15 mm in diam., up to 50 cm long..l6.

14(13). Plants epiphytic, pendent..l5.

15(14). Leafy stems ca. 2 mm in diam., up to 0.5 m long, branched 3-6 times. Sterile leaves ca.
4-5 mm long, 0.2 mm wide, linear; fertile leaves lanceolate, at least 3 times longer than wide.

40. L. tortile

15(14). Leafy stems 5 mm in diam., up to 1 m long. Leaves very thick, acicular, appressed or
slightly incurved, ca. 8 mm long, 0.75 mm wide, the midrib obscure.

25. L. funifomte

16(14). Leaves 4-5 mm long, 3 mm wide, ascending, reddish.

26. L. goudotii

16(14). Leaves ca. 1 cm long, usually less than 3 mm wide, strongly ascending, appressed or
incurved toward the stem..l7.

17(16). Leaves not completely appressed, ciuwed, not crowded, with the stems visible toward the
base of the plant, at least in larger specimens..l9.

17(16). Leaves completely appressed, straight, crowded, the stems nowhere visible.. 18.

18(17). Leaves with very small, longitudinal wrinkles on the abaxial surface and with none or only a
few low, clear, specialized marginal cells. Leafy stems 10-25 cm long, 5-9 mm wide including the
leaves; leaves in probably 8 longitudinal rows, ca. 7.5-10 mm long, 1-1.25(1.5) mm wide, thick, stiff,
not keeled, narrowly lanceolate, greenish, yellowish, or reddish.

20. L. crassum

26 LYCOPODIACEAE: 2. LYCOPODIUM

18(17). Leaves smooth on the abaxial surface and with occasional, tooth-like, clear, specialized
marginal cells. Leafy stems 5-15 cm long, 6-10 mm wide including the leaves; leaves in 12
longitudinal rows, ca. 5-6 mm long, 1 mm wide, rather thick, stiff, not keeled, lanceolate, greenish or
yellowish.

19. L. chiricanum

19(17). Leaves broadly lanceolate, ca. 6 mm long, 2.5 mm wide, usually distinctly keeled, decidedly
spreading, dark ohve green, with a wide band of specialized, bulbous marginal cells. Leafy stems
30 - 40 cm long, 10 - 17 mm wide including the leaves; leaves apparently in 8 longitudinal rows, rather
thick, stiff.

33. L. pflanzii

19(17). Leaves narrowly lanceolate, ca. 7-10 mm long, 1-1.5 mm wide, not keeled, somewhat
spreading, pale yellowish or greenish, with scattered, specialized, clear, bulbous or tooth-like
marginal cells..20.

20(19). Leafy stems 5-15 mm in diam.; leaves in 8-12 longitudinal rows, with few to many
specialized, clear, bulbous marginal cells. Leaves narrowly lanceolate, ca. 10 mm long, 1 mm wide,
ascending or usually rather spreading and incurved at the tip, thick, stiff, sUghtly and broadly keeled
towards the leaf base, greenish or yellowish, rarely reddish at the apex.

37. L. saurums

20(19). Leafy stems 4-6 mm in diam. leaves in ca. 6 longitudinal rows, with a few specialized,
clear, tooth-like marginal cells. Leaves lanceolate, ca. 7 mm long, 1.5 mm wide, ascending and nearly
straight at the tip, rather thick, stiff, not keeled, yellowish to greenish and usually reddish at the apex.

15. L. attenuatum

21(13). Sterile leaves linear to narrowly lanceolate or narrowly lanceate..24.

21(13). Sterile leaves narrowly oblanceolate, obovate, or elhptical..22.

22(21). Fertile and sterile leaves monomorphic; plants erect. Leaves thin but firm, narrowly
oblanceolate, 10 - 17 mm long, 3 mm wide, the midrib abaxially prominent.

24. L. foliaceum

22(21). Fertile and sterile leaves strongly dimorphic; plants pendent..23.

23(22). Stems red; sterile leaves 2.5-5 mm long. Sterile leaves thin, almost translucent, obovate.

21. L. cuneifolium

23(22). Stems not red; sterile leaves 5-8 mm long. Sterile leaves thin but firm, nearly eUiptical.

22. L. dichaeoides

24(21). Largest leafy branches including the leaves 1-2(3) mm wide. Apex of the sterile leaves
curved away from the stem, the leaves ca. 3 mm long, 0.2 mm wide, linear; fertile leaves linear-
lanceolate.

42. L. verticillatum war. parvifolium

24(21). Largest leafy branches including the leaves (3)5 - 30(40) mm wide..25.

25(24). Plants epiphytic (except L. hippurideum)\ leafy branches (3)5 - 30(40) mm wide..28.

25(24). Plants terrestrial; leafy branches 5-12 mm wide..26.

26(25). Sterile leaves linear, ca. 0.2 mm wide, entire, very crowded, the stems concealed. Sterile
leaves thick, not expanded at the base, the midrib obscure.

18. L. chamaeleon

26(25). Sterile leaves lanceolate or linear-lanceolate, ca. 0.2-1 mm wide, obscurely toothed,
distant, the stems mostly visible.. 27.

27(26). Stems ca. 1-2 mm in diam. excluding the leaves; leaves 0.5-0.75 mm wide; plants usually
10-25 cm long. Leaves reflexed, spreading, sparingly toothed, the midrib prominent abaxially.

36. L. reflexum

27(26). Stems ca. 2-3 mm in diam. excluding the leaves; leaves 1-1.25 mm wide; plants usually
25-40 cm long. Leaves strongly reflexed, very sparsely toothed, the midrib usually not prominent
abaxially.

28. L. hojfmannii

LYCOPODIACEAE: 2. LYCOPODIUM 27

28(25). Sterile leaves lanceolate to linear-lanceolate, 1-3 mm wide; leaves weakly to strongly
dimorphic, the fertile shorter than the sterile ones..36.

28(25). Sterile leaves linear, 0.1-1 mm wide; leaves monomorphic (slightly dimorphic in L.
dichotomum ) . .29.

29(28). Sterile leaves 0.2- 1 mm wide. Leaves thin, often shghtly revolute at the margins and with
usually obscure midribs.. 32.

29(28). Sterile leaves ca. 0.1 mm wide..30.

30(29). Plants (15)25 - 50 cm long, lax and pendent. Branches mostly more than 5 cm distant along
the stems; leaves thin, distant, shghtly expanded at the base, the midrib prominent.

43. L. watsonianum

30(29). Plants 15(20) cm long, erect..31.

31(30). Leaf bases red. Leaves straight, ca. 10-12 mm long; sporangia 1-1.2 mm wide.

44. L. wilsonii

31(30). Leaf bases not red. Branches less than 3 cm distant along the stems; leaves thin, distant,
not expanded at the base, curved, ca. 7-8 mm long, the midrib obscure; sporangia ca. 0.8 mm wide.

35. L. polycarpum

32(29). Sterile leaves ca. 10 mm long; stems 1-2 mm in diam. excluding the leaves..34.

32(29). Sterile leaves 15-20 mm long; stems ca. 3 mm in diam. excluding the leaves..33.

33(32). Stems red; plants epiphytic. Leaves thin, 20 - 30 mm long, 0.75 mm wide.

34. L. pithyoides

33(32). Stems not red; plants terrestrial. Leaves thin, 15-20 mm long, 1 mm wide.

27. L. hippurideum

34(32). Sterile leaves crowded, ascending; fertile leaves shghtly dimorphic; stems at least 1 mm in
diam. Plants stiff, erect or perhaps occasionahy pendent; branches diverging at a 30 - 40° angle;
leaves thin, linear, ca. 10 mm long, 0.75 mm wide.

23. L. dichotomum

34(32). Sterile leaves distant, spreading; fertile leaves monomorphic, stems less than 0.5 mm in
diam.; plants lax and decidedly pendent..35.

35(34). Stems branched 1-4 times at ca. a 45° angle; stems with the leaves mostly 10-15 mm wide.
Leaves thin, arcuate, ca. 10 mm long. 0.75 mm wide.

17. L. capillare

35(34). Stems branched 3-6 times at ca. a 20° angle; stems with the leaves mostly 5-10 mm wide.
Leaves very thin, straight, keeled, ca. 8 mm long, 0.75 mm wide.

31. L. mollicomum

36(28). Sterile leaves at the base of the plant ca. 2-3 mm wide..39.

36(28). Sterile leaves at the base of the plant ca. 1- 1.5(2) mm wide..37.

37(36). Sterile leaves ascending at a 30 - 45° angle to the stem. SporophyUs weakly or rarely
strongly dimorphic; sterile leaves linear, crowded, often revolute, 8-12 mm long, 0.3 mm wide.

41. L. tubulosum

37(36). Sterile leaves spreading at ca. a 90° angle to the stem..38.

38(37). Plants pendent, 15-60 cm long; leaves strongly dimorphic, the fertile ones shorter; stems
often pale pink. Sterile leaves linear, few and distant, 8-12 mm long, 0.75-1 mm wide.

38. L. subulatum

38(37). Plants erect, 5-10(15) cm long; leaves monomorphic; stems stramineous. Leaves linear,
thick, somewhat distant, slightly revolute at the margins, the midribs prominent abaxially, obscure
adaxially.

16. L. bradeorum

39(36). Lowest sterile leaves imbricate, stiff, somewhat spreading to strongly ascending. Fertile
leaves slightly to strongly dimorphic; sterile leaves thin but stiff, the midribs obscure or prominent
only near the base of the leaf.

39. L. taxifolium

28 LYCOPODIACEAE: 2. LYCOPODIUM

39(36). Lowest sterile leaves not imbricate, thin, spreading to somewhat ascending..40.

40(39). Fertile leaves ca. 0.5 mm wide; proximal sterile leaves 15-25 mm long, linear-lanceolate.
Leaves somewhat dimorphic, the sterile ones thin and flexible, the midribs prominent to the apex.

30. L. linifolium

40(39). Fertile leaves ca. 1 mm wide; proximal sterile leaves 7-10 mm long..41.

41(40). Leaves strongly dimorphic; sterile leaves lanceolate, 7-9 mm long, with obscure midribs.
32. L. myrsinites

41(40). Leaves essentially monomorphic; sterile leaves elliptic-lanceolate, 8-10 mm long, the
midribs prominent nearly to the leaf apex.

29. L. lancifolium

LYCOPODIUM subg. LYCOPODIUM

3. Lycopodium clavatum van aristatum (Humb. & Bonpl. ex Willd.) Spring, Flora

21(1):173. 1838.

Lycopodium aristatum Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:17. 1810. TYPE: Silla de
Caracas, Distr. Fed., Venezuela, Humboldt & Bonpland (B-Hb. Willd. 19351 not seen; isotype P not
seen Tyron photo GH, US).

Lycopodium trychophyllum Desv. Mem. Soc. Linn. Paris 6:184. 1827. TYPE LOCALITY: Brazil.
Synonymized by Proctor (FI. Less. Antill. 2:31. 1977).

Lycopodium aristatum var. incurvum Hook. & Grev. Bot. Misc. 2:376. 1831. TYPE: Martinique,
Sieber FI. Mixt. 327 (K not seen).

Lycopodium trichiatum var. desvauxianum Spring, Nouv. M6m. Acad. Roy. Sci. Bruxelles 15:92.
1842. TYPE: A renaming of L. trychophyllum, and so based on the type of that name.

Lycopodium eriostachys F6e, Crypt. Vase. Bresil 1:224. 1869. TYPE: Serra dos Orgaos, Est. Rio
de Janeiro, Brazil, Glaziou 1788 (P or RB not seen; isotype BR not seen photo 5215).

Plants terrestrial, at 600-3300 m elevation, on wet banks and cliffs and
occasionally in swamps, from the Cordillera Central to Pcia. Chiriquf. Also from
the Antilles, Mexico to Venezuela, Bolivia, and Brazil.

I separate as var. aristatum the tropical American specimens, which are large
and coarse, compared to the north temperate specimens of var. clavatum. Variety
aristatum hybridizes with L. contiguum in the northern half of the Cordillera de
Talamanca and on Volcan Chiriquf. The hybrids have one or more sessile strobili
terminating upright branches, and the rhizomes may be hypogeous or epigeous.

4. Lycopodium contiguum Klotzsch, Linnaea 18:519. 1845.

Lycopodium clavatum var. pseudocontiguum Christ in Pitt. Prim. FI. Costaric. 3(1):55. 1901.
TYPE: Cerro de las Vueltas, Buena Vista massif, Pcia. S. Jos6, 3000 m, Pittier 10467 (P not seen;
isotype US).

Lycopodium mayoris Rosenst. M6m. Soc. Sci. Nat. Neuchatel 5:56, f. 11. 1912. TYPE: Trail from
Bogota to the P^amo Cruz Verde, Depto. Cundinamarca, Colombia, ca. 2900 m. Mayor 17 (S not
seen; isotype US).

SYNTYPES: Silla de Caracas, Distr. Fed., Venezuela, Moritz 224 (B not seen;
isotype US); and Pcia. Quito, Ecuador, Hartweg 1474 (K not seen fragm NY).

Plants terrestrial, at 2200-3500 m elevation, in paramos and occasionally in
open areas of cloud forests, from the Cerro de la Muerte to Cerro Copete (Pcia.
Chiriquf) and Volcdn Chiriquf. Also from Venezuela and Colombia to Bolivia.

This species hybridizes with L. clavatum var. aristatum; see notes under that
variety.

LYCOPODIACEAE: 2. LYCOPODIUM 29

5. Lycopodium jussiaei Desv. in Poir. Encyc. M^th. Suppl. 3:543. 1814.

Lycopodium haenkei K. Presl, Reliq. Haenk. 1:78. 1825. TYPE: Hudnuco, Pcia. Huanuco, Peru,
Haenke (PRC not seen).

TYPE: Peru, Jussieu (P-Hb. Juss. 658 not seen photos 3247, 3248).

Plants terrestrial, at 2500-2900 m elevation, in bogs and open areas in montane
forests, from the Cerro de la Muerte and vicinity. Also from Jamaica, Hispaniola,
Venezuela, and Colombia to Bolivia and Brazil.

6. Lycopodium magellanicum (Pal. Beauv.) Swartz, Syn. Fil. 180. 1806.

Lepidotis magellanica Pal. Beauv. Prodr. Aethdogram. 102. 1805. TYPE: Straits of Magellan,
Chile, Commerson (persumably G-Hb. Delessert not seen). A specimen was communicated to
Palisot by A. L. de Jussieu, and so there is presumably an isotype in the Jussieu herbarium.
However, Hb. Juss. no. 668, labelled “magellan... Commerson” is L. confertum Willd., whereas no.
669, without collector or locality and labelled both L. magellanicum and L. confertum, is true L.
magellanicum. I suspect that Palisot was sent part of 669, which I assume should bear the same
locality and collector as 668.

Plants terrestrial, at 3300-3800 m, in paramos, in the Flora area known only
from Cerro Chirripo, Pcia. S. Jose {Evans & Lellinger 182, US). Also from
Hispaniola {Ekman H13666, US), Argentina, and Chile.

Tbe North American specimens have been confirmed by Dra. Cristina Rolleri,
who has made extensive morphological and anatomical studies of the group. This
species differs only slightly in aspect from the Andean L. spurium Willd.

7. Lycopodium thyoides Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:18. 1810.

Lycopodium complanatum var. tropicum Spring in Mart. FI. Bras. 1(2):116. 1840. SYNTYPES:
Serra d’Estrella, Est. Rio de Janeiro, Brazil, Schott (W not seen); and S. Gongalo d’Amarante, Est.
Piam, Brazil, Martius (M not seen).

Lycopodium complanatum var. adpressifolium Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles
15:102. 1842, nom. superfl. TYPE: A renaming of L. complanatum var. tropicum, and so based on
the type of that name.

TYPE: Silla de Caracas, Distr. Fed., Venezuela, Humboldt (B-Hb. Willd. 19352
not seen fragm NY; isotype P not seen).

Plants terrestrial, at 1800-3400 m elevation, from the Cordillera Central and
the Cordillera de Talamanca to Cerro Chirripo. Also from Venezuela and
Colombia to Bolivia and Brazil.

I agree with Wilce (Beih. Nova Hedwigia 19:156-158. 1965) that the

specimens from the Antilles and those from Mexico to El Slavador represent
other, closely related taxa, rather than typical L. thyoides.

LYCOPODIUM subg. CERNUISTACHYS Herter

8. Lycopodium alopecuroides var. integerrimum Spring, Nouv. M6m. Acad. Roy.

Sci. Bruxelles 15:75. 1842.

Lycopodium longipes Hook. & Grev. Bot. Misc. 2:372. 1831. TYPE: Sta. Catarina Island, Est.
Sta. Catarina, Brazil, Macrae (K not seen).

TYPE: A renaming of L. longipes Hook. & Grev., and so based on the type of
that name.

Plants terrestrial, at 1300 m elevation, along wet roadsides, in the Flora area
known only from Tapantf {Saenz & Nassar 1450, CR). Also from Cuba, Mexico,
Guatemala, Venezuela, Colombia, Ecuador, and Brazil.

30 LYCOPODIACEAE: 2. LYCOPODIUM

LYCOPODIACEAE; 2. LYCOPODIUM 31

9. Lycopodium carolinianum van meridionale (Underw. & Lloyd) Nessel &
Hoehne, Arch. Bot. Sao Paulo 1(4):431. 1927.

Lycopodium meridionale Underw. & Lloyd, Bull. Torrey Bot. Club 33:121. 1906. TYPE: Puerto
Rico, Wilson 94 (NY not seen).

Plants terrestrial, at 0-100 m elevation, in swampy areas, in the Flora area
known only from Quibdo and vicinity. Also from the Antilles, southern Mexico,
Guatemala, Belize, Honduras, Nicaragua, and tropical South America.

10. Lycopodium cernuum L. Sp. PI. 2:1103. 1753, van cernuum.

Lycopodium capillaceum WiUd. ex Spring, Flora 21(1):165. 1838, nom. illeg. BASED ON:

Guanaguana, Edo. Monagas, Venezuela, Humboldt 473 (B-Hb. Willd. 19429 not seen microfiche S. I.
Library). Invalid because published in synonymy.

Lycopodium cernuum var. capillaceum Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:80. 1842.
TYPE: Based on L. capillaceum Willd. ex Spring, and so based on the type of that name.

Lycopodium cernuum vm . panamense Werckle ex Nessel, Revista Sudamer. Bot. 6:173, t. XIX, f.
102. 1940. TYPE: Environs of Balboa, Pcia. Panama, Powell 1921 (BONN not seen photo US;
isotypes P-Hb. Bonaparte, Hb. Werckle neither seen).

Lycopodium cernuum var. watsonianum Nessel, Revista Sudamer. Bot. 6:173, t. XIX, f. 101. 1940.
TYPE: Panama, Biolley (P-Hb. Beukert, Hb. Christ neither seen).

TYPE: Locality and collector unknown (LINN 1257.13 not seen microfiche S. I.
Library).

Plants terrestrial, at 0-2700 m elevation, in disturbed places, throughout Costa
Rica and western Panama, but not so common in Pcia. Darien or the Choco. Also
from throughout tropical America.

11. Lycopodium cernuum var. curvatum (Swartz) Hook. & Grev. Bot. Misc. 2:369.
1831.

Lycopodium curvatum Swartz, J. Bot. (Schrader) 1800(2):116. 1801; Syn. Fil. 178, 402. 1806.
TYPE: West Indies, Swartz (S not seen).

Plants terrestrial, at 500-2000 m elevation, from Cerro Chompipe (Pcia.
Heredia), Alto S. Juan on the road to Dominical (Pcia. S. Jose), and the summit of
Cerro Jefe. Also from Venezuela to Bolivia.

This variety resembles Lycopodiella camporum 011gaard & Windisch.

12. Lycopodium pendulinum Hooker, Icon. PI. l:t. 90. 1837.

Lycopodium cernuum f. macrostachyum Christ in Pitt. Prim. FI. Costaric. 3(1) :55. 1901. TYPE:
Achiote, Volcan Poas, Pcia. Heredia, 2300 m, Tonduz 10692 (P? not seen; isotypes CR, US).

Lycopodium cernuum var. pendulinum (Hook.) Nessel, Barlappgewachse 367. 1939, as

''pendulum'' and incorrectly attributed to Hooker.

TYPE: Casapf, Pcia. Huanuco, Peru, Mathews 1776 (K not seen fragm US).
Plants terrestrial, at 2200-2700 m elevation, from the Cordillera Central and
the Cordillera de Talamanca to Cerro Arizona above Sta. Fe. Also from
Venezuela and Colombia to Bolivia.

FIGS. 11-19. Lycopodium. FIG. 11. Branch apices of L. cernuum var. curvatum, Standley &
Valerio 52083. FIG. 12. Branch apices of L. pendulinum, Tonduz 10692. FIG. 13. Branch apex of L.
riofrioi. Woodruff. FIG. 14. Branch apices of L. trianae, Killip 35353. FIG. 15. Branch portion of L.
attenuatum, Scamman 7804. FIG. 16. Branch portion of L. bradeomm, Standley & Valerio 50389.
FIG. 17. Branch portion of L. capillare, Johnson 917, Guatemala. FIG. 18. Branch apex of L.
chamaeleon, Standley & Valerio 52083. FIG. 19. Br^mch portion of L. chiricanum, Maxon 5364.

32 LYCOPODIACEAE: 2. LYCOPODIUM

This species and L. riofrioi have pendulous fertile branch tips.

13. Lycopodium riofrioi Sodiro, Anal. Univ. Quito 12(81):337 (repr. 582). 1895.

Lycopodium pensum Lellinger & Mickel in Lellinger, Proc. Biol. Soc. Wash. 89:717, f. 2. 1977.
TYPE: 6 mi. from S. Rafael de Heredia on the slopes of Volcan Barba, Pcia. Heredia, McAlpin 216
(DUKE fragm GH).

TYPE: Gualea, Pcia. Pichincha, Ecuador, Sodiro (not seen).

Plants terrestrial, at 1700-2000 m elevation, from Volcan Barba and the
Cordillera de Talamanca to Pcia. Chiriqui. Also from Venezuela, Colombia, and
Ecuador.

14. Lycopodium trianae Hieron. Bot. Jahrb. Syst. 34:574. 1905.

Lycopodium cemuum var. trianae (Hieron.) Nessel, Barlappgewachse 358. 1939, attributed

erroneously to Hieronymus.

LECTOTYPE: Arrostradero, Depto. Choco, 2500 m, Triana 234 (B not seen;
isolectotypes NY, US), chosen by Nessel (Barlappgewachse 359. 1939).

Plants terrestrial, at 0-800 m elevation, in disturbed areas in lowland rain
forests and second-growth forests, from the Rfo S. Juan basin. Also from Pacific
coastal Colombia south to Depto. Nariho.

LYCOPODIUM subg. SELAGO Baker

15. Lycopodium attenuatum Spring, Nouv. M^m. Acad. Roy. Sci. Bruxelles 24:8.
1849.

Lycopodium tobari Sodiro, Anal. Univ. Quito 12(81):320 (repr. 565). 1895. SYNTYPES: Cerro de
Puntas, Ecuador, Sodiro (Hb. Sodiro not seen) and Mt. Pichincha, Ecuador, 3400 m, Sodiro (Hb.
Sodiro not seen).

LECTOTYPE: Mt. Pichincha, Pcia. Pichincha, Ecuador, 11,500 ft, Hartweg 1470
(US; isolectotype B not seen fragm NY, K not seen fragm NY), chosen by
Lellinger (Proc. Biol. Soc. Wash. 89:717. 1977).

Plants terrestrial, at 3100-3400 m elevation, in paramos, from the Cerro de la
Muerte. Also from Colombia and Ecuador.

16. Lycopodium bradeorum Christ, Repert. Spec. Nov. Regni Veg. 8:20. 1910.

ILycopodium brauseanum Herter in Urban, Symb. Antill. 7:165. 1912. TYPE: Mt. S. Juan,

Venezuela, 600 m, /. R. Johnston 156 p. p. (GH not seen).

TYPE: La Palma, Pcia. S. Jose, 1400 m. Erode & Erode 282 (P not seen photo
22346; isotypes UC, US).

Plants epiphytic, at 1000-2400 m elevation, from the Cordillera Central and
from near Boquete. Also from Guatemala, El Salvador, Nicaragua, Venezuela,
and Ecuador.

This is a species that is very distinct in aspect, resembling a young Toxus or
Cryptomerio plant.

17. Lycopodium capillare Sodiro, Recens. Crypt. Vase. Quit. 90. 1883.

Lycopodium sarmentosum var. rubescens Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:13.
1849. TYPE: Depto. Cauca, Colombia, Hartweg 1464 (BR not seen; isotype fragment US).

Lycopodium underwoodianum Maxon, Contr. U. S. Natl. Herb. 13:41, t. 9. 1909. TYPE:

Coliblanco, on the lower slopes of Volcan Turrialba, Pcia. Cartago, ca. 1950 m, Maxon 213 (US;
isotypes CR, NY).

LYCOPODIACEAE: 2. LYCOPODIUM 33

Lycopodium guatemalense Maxon, Contr. U. S. Natl. Herb. 17:177, t. 9A. 1913. TYPE:

Pansamal^ Depto. Alta Verapaz, Guatemala, ca. 1200 m, von Tuerckheim 957 (US).

TYPE: Hacienda Guanaxilia, Pcia. Pichincha, Ecuador, 400 m, Sodiro (K not
seen).

Plants epiphytic, at (1000)1500-2400 m elevation, from the Cordillera Central,
the vicinity of Boquete, and Cerro Mali (Pcia. Darien). Also from southern
Mexico, Guatemala, Venezuela, Colombia, Ecuador, and southern Brazil.

18. Lycopodium chamaeleon (Herter) Morton, Amer. Fern J. 54:72. 1964.

Urostachys chamaeleon Herter, Amer. Fern J. 48:82. 1958. TYPE: Barranca Grande, near S.
Marcos, Pcia. S. Jose, Feb 1926, O. Jimenez (US).

Plants terrestrial, at ca. 1500-2400 m elevation, on wet banks, from between
Volcan Barba and Volcan Turrialba and from the type locality.

This species could be mistaken for L. reflexum, but its leaves are narrower and
more crowded and completely conceal the stems, except at the base of the plant.
When found growing together, these two species do not hybridize (R. Faden in litt.
22 Dec 1965).

19. Lycopodium chiricanum Maxon, Contr. U. S. Natl. Herb. 17:176, t. 8. 1913.

TYPE: Summit of Volcan Chiriqui, Pcia. Chiriqui, ca. 3370 m, Maxon 5364 (US
fragm NY).

Plants terrestrial, at 3000-3300 m elevation, in montane forests and open areas,
from Volcan Turrialba and Volcan Chiriqui.

20. Lycopodium crassum Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:50. 1810.

Lycopodium erythraeum Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:7. 1849. SYNTYPES:
Sierra Nevada, Edo. Merida, Venezuela, 10,000 ft. Linden 569 (BR not seen); Andes of Ecuador,
14,000 ft, Jameson (K not seen); and Mt. Antisana, Pcia. Pichincha, Ecuador, Hartweg (BR? not
seen).

Urostachys crassus var. herteri Nessel, Repert. Spec. Nov. Regni Veg. 36:180, t. 171. 1934. TYPE:
Mt. Antisana, Pcia. Pichincha, Ecuador, Hartweg 1472 (Hb. Lawory, Hb. Beukert not seen; isotypes
BM? and BR? neither seen).

Urostachys kuppen Nessel, Repert. Spec. Nov. Regni Veg. 36:180, t. 176. 1934. TYPE: Volcan de
Pasto, Depto. Narino, Colombia, 3500 - 4000 m, Hartweg 1474b (P not seen).

Urostachys moritzii Herter ex Nessel, Revista Sudamer. Bot. 6:160, t. 9, f. 29. 1940. TYPE: Sierra
Nevada, Edo. Merida, Venezuela, Greven 61 in 1898 (Hb. Greven not seen).

Urostachys ononis Herter, Ind. Lycopod. 73. 1949, nom. superfl. TYPE: A renaming of U. moritzii
Greven ex Herter, and so based on the type of that ncune.

Urostachys bonae-voluntatis Herter, Revista Sudamer. Bot. 10:112. 1953. TYPE: Mt. Mojanda,
Pcia. Pichincha, Ecuador, Mille 95 (US).

Urostachys bonae-voluntatis var. minor Herter, Revista Sudamer. Bot. 10:113. 1953. TYPE:

Between Quebrada de S. Paulino and Quebrada de Lopez, Lagunilla de las Casitas, western slopes of
the Cordillera Central, Depto. Cauca, Colombia, Cuatrecasas 19085a (US).

Lycopodium nesselianum Duek & Lellinger, Amer. Fern J. 68:120. 1978. TYPE: A renaming of
Urostachys moritzii Greven ex Herter non Lycopodium moritzii MueU. (Bot. Zeit. 19:65. 1861), and
so based on the type of that name.

TYPE: Mt. Antisana near Chusslongo, Pcia. Pichincha, Ecuador, 2218 hexap,
Humboldt 2263 (B-Hb. Willd. 19417 not seen Tryon photo).

Plants terrestrial, at 3000-3700 m elevation, in paramos, from the Cerro de las
Vueltas and Cerro Chirripo. Also from Venezuela to Peru.

34 LYCOPODIACEAE: 2. LYCOPODIUM

22 L. dichaeoide:

LYCOPODIACEAE; 2. LYCOPODIUM 35

21. Lycopodium cuneifolium Hieron. Bot. Jahrb. Syst. 34:572. 1905.

Lycopodium aqualupianum var. ohtusifolium Spring, Nouv. M6m. Acad. Roy. Sci. Bruxelles 24:31.
1849. TYPE LOCALITY: Colombia.

SYNTYPES: Volcan Barba, Pcia. Heredia, Hojfmann 50 (B not seen fragm
US); and near Merida, Edo. Merida, Venezuela, Moritz 577 (B not seen).

Plants epiphytic, at 1500-2400(3000) m elevation, mostly in lower montane
forests, from the Cordillera Central to near Copey de Dota (Pcia. S. Jose) and
from the Cerro de la Horqueta (Pcia. Chiriqui). Also from Venezuela and
Colombia.

This species resembles L. dichaeoides, but has smaller sterile leaves and reddish
stems.

22. Lycopodium dichaeoides Maxon, Proc. Biol. Soc. Wash. 18:231. 1905.

TYPE: Near Finca Sepacuite, Depto. Alta Verapaz, Guatemala, Cook & Griggs
251 (US).

Plants epiphytic, at (800)1100-2100 m elevation, in forests, from the Cordillera
Central to the Serrania del Darien. Also from Guatemala, Honduras, Nicaragua,
Colombia, and Ecuador.

This species resembles L. cuneifolium ^ but has larger sterile leaves and
stramineous stems. It may hybridize with that species in Pcia. Darien.

23. Lycopodium dichotomum Jacq. Enum. Stirp. Vindob. 314. 1762.

Lycopodium gramineum Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:19. 1849. TYPE:
Guayaquil, Pcia. Guayas, Ecuador, 29 (K not seen fragm NY).

Lycopodium barbatum Christ, Bull. Herb. Boissier II, 5:254. 1905, non Kaulf., 1824. TYPE: Costa
Rica, Werckle in 1904 (P not seen; isotype US).

Lycopodium chamaepeuce Herter, Bot. Jahrb. Syst. 43, Beibl. 98:50. 1909. TYPE LOCALITIES:
Guadeloupe and French Guiana.

Urostachys chamaepeuce var. urbanicus Herter, Repert. Spec. Nov. Regni Veg. 19:164. 1923.
SYNTYPES: Grenada, Broadway 3457 p. p. (B not seen; isotype fragm US as 3275) and 3458b (B not
seen).

Urostachys mortonii Herter in F. Morton, Repert. Spec. Nov. Regni Veg. 28:108. 1930.

SYNTYPES: Hacienda de Chocola, Depto. Mazatenango, Guatemala, 820 m, Morton 225a and 284a
(B neither seen).

Urostachys lindeneri Herter ex Nessel, Revista Sudamer. Bot. 6:164, t. 50. 1940. TYPE: Chilion,
near Mazatenango, Depto. Suchitepequez, Guatemala, Bemouilli & Cario 1866 (P-Hb. Christ not
seen).

Urostachys schlechtendalii Nessel, Revista Sudamer. Bot. 6:164, t. 11, f. 51. 1940. TYPE:

Guaveyma, Mexico, 5000 ft, Wagner 1849 (Hb. Beukert, Hb. Merson neither seen).

LECTOTYPE: Plate 45 in volume 3 of Jacquin’s “Hortus Botanicus
Vindobonensis,” which illustrates a plant from Martinique, chosen by G. R.
Proctor (FI. Less. Antill. 2:28. 1977).

FIGS. 20-27. Lycopodium. FIG. 20. Sterile branch portion of L. crassum, Evans & Lellinger 143.
FIG. 21. Sterile and fertile branch portion of L. cuneifoliumy Brade & Brade (Ros. Fil. Costar. Exs.
125). FIG. 22. Sterile and fertile branch portion of L. dichaeoides , Cook & Doyle 70, Guatemala.
FIG. 23. Branch tip of L. dichotomum, Standley 26116. FIG. 24. Branch portion of L. foliaceum,
Standley & Valerio 49706. FIG. 25. Branch portion of L. funiforme, Skutch 3731. FIG. 26. Branch
portion of L. goudotii, Goudot 3, Colombia. FIG. 27. Branch portion of L. hippurideum, Allen 4890.

36 LYCOPODIACEAE: 2. LYCOPODIUM

Plants epiphytic, at 0-1200(2500?) m elevation, in forests, from throughout the
Flora area, but mostly from the Cordillera de Tilardn, the Cordillera Central, and
central Panama. Also from Florida, the Antilles, and Mexico to Brazil.

24. Lycopodium foliaceum Maxon, Smithsonian Misc. Collect. 56(29) :1, t. 1.
1912.

TYPE: Upper Caldera River, Holcomb’s Trail above Boquete, Pcia. Chiriqui,
ca. 1650 m, Maxon 5628 (US; isotype NY).

Plants epiphytic, at 1000-2400 m elevation, from the Cordillera Central to
Boquete.

25. Lycopodium funiforme Cham, ex Spring, Nouv. M6m. Acad. Roy. Sci.
Bruxelles 15:50. 1842.

SYNTYPES: “California,” Chamisso (P-Hb. Bory not seen), presumably an
incorrect reference; and Guadeloupe, UHerminier (P not seen).

Plants epiphytic, at 0-2000 m elevation, in forests, from scattered localities in
Costa Rica and Panama, and from La Trojita, Rio Calima, Depto. Valle
{Cuatrecasas 16553, US). Also from the Antilles and Venezuela to Guyana.

The earlier publication of L. funiforme Bory ex Brongn. (Hist. Veg. Foss. 2:10,
18, t. 7, f. 9. 1837) is often cited, but is not valid. There was no intent to describe
a new species; the description, in a discussion of phyllotaxy, is incidental and not
diagnostic, and the illustration lacks analyses.

26. Lycopodium goudotii Herter, Bot. Jahrb. Syst. 43, Beibl. 98:47. 1909.

TYPE: Bogota, Distr. Esp., Colombia, Goudotl in 1844 (G not seen; isotype P

not seen 011gaard photo).

Plants terrestrial, at 3600 m elevation, in pdramos, in the Flora area known only
from Cerro Crestones, Chirripo Massif, Pcia. S. Jose {Gomez 7270, CR). Also
from Colombia and Peru.

^ 27. Lycopodium hippurideum Christ in Pitt. Prim. FI. Costaric. 3(1) :56. 1901.

Lycopodium lechleri Hieron. Bot. Jahrb. Syst. 34:571. 1905. LECTOTYPE: Near Tabina, Depto.
Puno, Peru, Lechler2012 (B not seen photo US).

TYPE: El Paramo, Buena Vista massif facing east, Pcia. Cartago, Pittier 10619
(P not seen; isotypes CR, US fragm NY).

Plants terrestrial, at (1200)2300-3300 m elevation, in montane (mostly oak)
forests and occasionally in paramos, from the Cordillera Central, the Cordillera de
Talamanca to Cerro Chirripo, and Volcdn Chiriqui. Also from Hispaniola,
Mexico, Guatemala, El Salvador, Venezuela, and Colombia to Bolivia.

This species closely resembles L. pithyoides in its habit, but not its habitat.

28. Lycopodium hoffmannii Maxon, Contr. U. S. Natl. Herb. 17:423. 1914.

TYPE: Volcan Barba, Pcia. Heredia, Hoffmann 85 (US; isotype B not seen
fragm NY).

Plants terrestrial, at 2000-3300 m, on wet banks, from the Cordillera Central,
Cerro de las Vueltas, and Pcia. Chiriqui.

This species resembles L. reflexum in habit, but as a rule has larger, usually
longer stems and wider leaves. It may hybridize with L. reflexum.

LYCOPODIACEAE: 2. LYCOPODIUM 37

29. Lycopodium lancifolium Maxon, Contr. U. S. Natl. Herb. 17:177. 1913.

TYPE: Upper Caldera river, Holcomb’s Trail above Boquete, Pcia. Chiriqui, ca.
1650 m, Maxon 5627 (US fragm NY).

Plants epiphytic, at 1500-2000 m elevation, in the Flora area known only from
the type locality and from El Alto de La Palma, Pcia. S. Jose {Horich U. C. Bot.
Gard. 58.201, US).

This species is related to L. linifolium and L. taxifolium but has essentially
monomorphic, wider leaves than those species.

30. Lycopodium linifolium L. Sp. PL 2:1100. 1753.

Lycopodium linifolium var. laxum Fee, Hist. Foug. Antill. [Mem. Foug. 11]:129. 1866. TYPE:
Mexico, Schajfner 124 (P? not seen).

Lycopodium pittieri Christ, Bull. Soc. Bot. Geneve II, 1:236. 1909. TYPE: Cocos Island, Pittier
12357 (P not seen; isotype US).

Lycopodium linifolium var. subaristatum Christ, Bull. Soc. Bot. Geneve II, 1:236. 1909.

SYNTYPES: Costa Rica, Werckli (P not seen); and Tablazo, Pcia. S. Jos6, 1900 m, Brade & Brade
228 (P not seen), synonymized by G6mez (Brenesia 12113:16. 1977).

LECTOTYPE: Plate 166, figure C, of Plunder’s “Traite...”, chosen by Proctor
(FI. Less. Antill. 2:26. 1977).

Plants epiphytic, at 0-900(2200) m elevation, in mostly lowland forests
throughout the Flora area. Also throughout tropical America.

This species is said to be called “Ciprecillo” and to be a remedy for snake bites
(Standley & Valerio 45143, US).

31. Lycopodium mollicomum (Mart, ex Spring) Spring in Mart. FI. Bras.
1(2):113. 1840.

Lycopodium dichotomum subsp. mollicomum Mart, ex Spring, Flora 21(1): 162. 1838. TYPE
LOCALITY: Brazil.

Lycopodium flaccidum sensu F6e, Crypt. Vase. Br6sil 2:92, t. 106, f. 1. 1873, non Bory in Bel.,
1834. TYPE: Alto Macah6, Est. Rio de Janeiro, Brazil, Glaziou 4677 (presumably P not seen).

Urostachys neptunei Herter, Ind. Lycopod. 72. 1949. TYPE: A renaming of L. flaccidum sensu
Fee, and so based on the type of that name.

Lycopodium williamsii Underw. & Lloyd, Bull. Torrey Bot. Club 33:112. 1906. TYPE: New
Brazil, Bolivia, 5500 ft, Williams 1393 (NY not seen).

Plants epiphytic, at 1200-1900 m elevation, from the Cordillera Central and the
valley of the Rio Grande de Orosi (Pcia. Cartago). Also from tropical South
America.

32. Lycopodium myrsinites Lam. Encyc. M^th. 3:654. 1789.

Lycopodium roraimense Underw. & Lloyd, Bull. Torrey Bot. Club 33:115. 1906. TYPE: Forest
slopes near Roraima, Guyana., Jenman (NY not seen).

Lycopodium skutchii Maxon, Proc. Biol. Soc. Wash. 46:159. 1933. TYPE: Chichavac, Depto.
Chimaltenango, Guatemala, 2400-2700 m, Skutch 243 (US).

TYPE: Hispaniola, comm. J. Martin (P-Hb. Lam not seen).

Plants epiphytic, at (300)1600-3000 m elevation, in forests, from the Cordillera
Central, the northern end of the Cordillera de Talamanca, and the Fila de Cedral.
Also from Hispaniola, Mexico to Honduras, and Venezuela to Peru and Guyana.

This species is related to L. dichaeoides, but differs in its lanceolate leaves that
are acute at the apex.

38 LYCOPODIACEAE: 2. LYCOPODIUM

33 L. pfianzii

32 L. myrsinites

30 L. linifolium

LYCOPODIACEAE: 2. LYCOPODIUM 39

33. Lycopodium pflanzii (Nessel) Lellinger, Proc. Biol. Soc. Wash. 89:717. 1977.

Urostachys pflanzii Nessel, Repert. Spec. Nov. Regni Veg. 36:181, 1. 171. 1934. SYNTYPES: Near
Corocoro, Depto. La Paz, Bolivia, Pflanz in 1920 (Hb. Nessel not seen; isotype fragm US ex Hb.
Bodino, Milan); and S. Benito, Sacaba, Depto. Cochabamba, Bolivia, 3000 - 4000 m, 14 Nov 1921,
Steinbach (B not seen).

Plants terrestrial, at 3100-3200 m elevation, in paramos, in the Flora area
known only from the Cerro de la Muerte ca. 22 km north of S. Isidro del General,
Pcias. Cartago and S. Jose {Tryon & Tryon 7055, GH). Also from Bolivia.

It is possible, according to B. 011gaard (pers. comm.), that this plant is not
conspecific with Bolivian L. pflanzii.

34. Lycopodium pithyoides Schlechtend. & Cham. Linnaea 5:623. 1830.

Lycopodium mandioccanum var. mexicanum Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:45.

1842. TYPE: A renaming of L. pithyoides Schlechtend. & Cham., and so based on the type of that
name.

Lycopodium gigas Herter, Bot. Jahrb. Syst. 43, Beibl. 98:50. 1909. SYNTYPES: Southern Mexico,
collector unknown (not seen); and St. Yago, Libani, Cuba, Linden 1996 (P not seen; isosyntype K not
seen), synonymized by A. R. Smith (FI. Chiapas 2:254. 1981).

TYPE: Near Jalapa, Edo. Veracruz, Mexico, Schiede & Deppe (B or HAL not
seen).

Plants epiphytic, at 1200-1900(2700) m elevation, in forests, from the Fila de
Cedral, Las Concavas (Pcia. Cartago), and the northern end of the Cordillera de
Talamanca. Also from (Tuba, Hispaniola, Mexico to Honduras, and Venezuela.

35. Lycopodium polycarpum Kunze, Linnaea 9:5. 1834.

Urostachys cuatrecasasii Herter, Revista Sudamer. Bot. 10:123. 1953. TYPE: Rio Digua, Depto.
El VaUe, Colombia, 900-1180 m, Cuatrecasas 15143A (US).

TYPE: Near Casapi, Depto. Hudnuco, Peru, PoeppigDiar. 1124 (LZ destroyed).
Plants epiphytic, at (100)600-2000(2400) m elevation, from the Cordillera de
Tilaran, the Cordillera Central, the vicinity of Guapiles (Pcia. Limon), the
Cordillera de Talamanca to Pcia. Veraguas, and the Rio Salaquf (Depto. Choco).
Also from Jamaica, Puerto Rico, the Lesser Antilles, and Colombia.

This species differs from L. wilsonii in having shorter, falcate-ascending leaves
and more crowded, thinner, shorter stems.

36. Lycopodium reflexum Lam. Encyc. M6th. 3:653. 1789, var. reflexum.

Lycopodium squarrosum Swartz, Nov. Gen. Sp. PI. Prodr. 137. 1788, non Forst., 1786, nom. illeg.
TYPE: Jamaica, Swartz (S not seen).

Lycopodium reflexum Willd. Sp. PI. ed. 4, 5:52. 1810, non Lam. 1789, nom. illeg. TYPE: Volcan
Tungurahua, Pcia. Tungurahua, Ecuador, Nee (B-Hb. Willd. 19419 not seen Tryon photo).

Lycopodium bifldum Humb. & Bonpl. ex Willd. Sp. PI. ed 4, 5:53. 1810. TYPE: Cuchilla de
Guanaguana, Edo. Monagas, Venezuela, 3288 ft, Humboldt 474 (B-Hb. Willd. 19421 not seen
microfiche S. I. Library).

FIGS. 28 - 36. Lycopodium. FIG. 28. Branch apex of L. hojfmannii, Valerio 2. FIG. 29. Branch
portion of L. lancifolium, Maxon 5627. FIG. 30. Branch portion of L. linifolium, Svenson 324. FIG.
31. Branch portion of L. mollicomum, Maxon 8280. FIG. 32. Sterile and fertile branch portion of L.
myrsinites, Skutch 1054, Guatemala. FIG. 33. Branch portion of L. pflanzii, Tryon & Tryon 7055
(GH). FIG. 34. Branch portion of L. pithyoides, von Tuerckheim 551, Guatemala. FIG. 35. Branch
portion of L. polycarpum, Allen 2070. FIG. 36. Branch apex of L. reflexum, Scamman 7532.

40 LYCOPODIACEAE: 2. LYCOPODIUM

Lycopodium eversum Poir. Encyc. Meth. Suppl. 3:556. 1813. TYPE: A renaming of L. reflexum
Willd., and so based on the type of that name.

Lycopodium reversum K. Presl, Reliq. Haenk. 1:82. 1825. TYPE: Guayaquil, Pcia. Guayas,

Ecuador, Haenke (PR not seen).

Lycopodium reflexum var. densifolium Baker, Handb. Fern-allies 11. 1887. SYNTYPES:
Colombia, Hartweg 1480 (K not seen); Venezuela, Moritz 2266 (K not seen); and Brazil, Glaziou
15797 (K not seen).

TYPE: Martinique, J. Martin (P-Hb. Lam. 442 not seen), cited by Underwood
and Lloyd (Bull. Torrey Bot. Club 33: 104. 1906).

Plants terrestrial, at 600-3300 m elevation, on wet banks or occasionally on
rotting tree trunks, mostly in open areas, from the Cordillera Central, near
Puriscal (Pcia. S. Jose), and the Cordillera de Talamanca to Pcia. Code. Also
from the Antilles, Trinidad, Mexico to Bolivia, Brazil, and Paraguay.

This species is related to L. hojfmannii and may hybridize with it. Proctor
(Rhodora 68:464. 1966) recognized L. reflexum var. rigidum (J. F. Gmel.) Proctor
from Jamaica, Hispaniola, and several islands in the Lesser Antilles.

37. Lycopodium saurunis Lam. Encyc. Mdh. 3:653. 1789.

Lycopodium andinum Rosenst. Repert. Spec. Nov. Regni Veg. 5:239. 1908. TYPE: Murusata
Mountains 40 km from La Paz, Depto. La Paz, Bohvia, Buchtien 173 (S not seen; isotype US).

TYPE: Bourbon [Reunion], Commerson (P not seen).

Plants terrestrial, at 2500-3400 m elevation, in paramos, from the Cordillera de
Talamanca to Pcia. Chiriquf. Also from Venezuela, Colombia to Bolivia, and
Argentina.

Rolleri (Revista Museo La Plata, Secc. Bot. 13:79. 1981) has ascribed L.

capellae (Herter) Morton to Costa Rica on the basis of one specimen from Cerro
Asuncion, Pcia. S. Jose {de la Sola 5048, LP), which I have not seen. Lycopodium
capellae differs from this species in having sporophylls shorter and wider than the
sterile leaves and in having the leaves thinner and less appressed to the stems.
011gaard (pers. comm.) is of the opinion that true L. saurunis does not occur north
of Peru. If so, this species may be without a correct name in the Flora area.

38. Lycopodium subulatum Desv. in Poir. Encyc. M6th. Suppl. 3:544. 1814.

Lycopodium congestifolium Spring, Nouv. M6m. Acad. Roy. Sci. Bruxelles 15:70. 1842. TYPE:
Peru, Dombey (LG not seen; isotypes G-Hb. Delessert, P neither seen).

Urostachys subulatus var. polymorphus Nessel, B^lappgewachse 243. 1939, attributed erroneously
to “(Willd.) Nessel.” TYPE: Ecuador, Humboldt (B-EO?. Willd. 19434a not seen Tryon photo GH,
US).

TYPE: Ecuador?, Hb. Desvaux (P not seen photo US).

Plants epiphytic, at (900)1800-2700 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Pcia. Chiriquf. Also from Colombia
to Argentina and Brazil.

39. Lycopodium taxifolium Swartz, Nov. Gen. Sp. PI. Prodr. 138. 1788.

Lycopodium passerinoides H.B.K. Nov. Gen. Sp. 1:41 (fol. 33). 1816. TYPE: Near Olleras and
Aipate, Depto. Puno, Peru, 747 hexap, Humboldt (B-Hb. Willd. 19410 not seen microfiche S. I.
Library).

Lycopodium stmthioloides K. Presl, Reliq. Haenk. 1:82. 1825. TYPE: “Hab. in Nootka-Sund,”
Vancouver Island, British Columbia, Canada. The specimen must be from tropical America,
collected by Haenke (PRC not seen).

LYCOPODIACEAE: 2. LYCOPODIUM 41

Lycopodium nitens Schlechtend. & Cham. Linnaea 5:623. 1830. PROBABLE TYPE: Near
Jalapa, Edo. Veracruz, Mexico, Schiede 831 (B not seen fragm US).

Lycopodium herminieri Spring, Bull. Acad. Roy. Sci. Bruxelles 8:514. 1841. TYPE: Guadeloupe,
L'Herminier (LG not seen).

Lycopodium picardae Christ ex Krug in Urban, Bot. Jahrb. Syst. 24:148. 1897. TYPE: Haiti, 1200
m, Picarda 978 (P? not seen fragm & photo US).

Lycopodium schwendeneri Herter, Bot. Jahrb. Syst. 43, Beibl. 98:50. 1909. TYPE: Haiti, 10 Sept
1908, Bogemann (B? not seen fragm US).

lUrostachys panamensis Nessel, B^lappgewachse 189. 1939, attributed to “(Humb. & Bonpl.)
Herter.” TYPE: Based on Panama, Bonpland (B-Hb. Willd. 19411 not seen Tryon photo US). This
is a new species because the name L. panamense Willd. is unpublished, but it is also superfluous
because L. brongniartii Spring is cited in synonymy.

Urostachys costaricensis Herter, Amer. Fern J. 48:83. 1958. TYPE: Costa Rica, Mar 1908,

Ridgway (US).

TYPE: Jamaica, Swartz (S not seen).

Plants epiphytic, at 1000-2200(2900) m elevation, in forests, from the
Cordillera Central to Pcia. Chiriquf. Also from the Antilles and Trinidad to
Surinam and from Mexico to Venezuela and Peru.

The dimorphism varies in this species from slight (usually in young and short
plants) to extreme (usually in old and long plants).

40. Lycopodium tortile Christ, Bull. Soc. Bot. Geneve 11, 1:235. 1909.

Lycopodium comans Christ in Schwacke, PI. Nov. Mineir. 2:40. 1900, non Hook., 1844.

SYNTYPES: Serra de Itatiaia, Est. Rio de Janeiro, Brazil, 2300 m, Ule 255 and 3537 (P or B neither
seen).

Urostachys comans Herter, B^lappgewachse 127. 1939, attributed erroneously to “(Christ) Hert.”
TYPE: In effect a renaming of L. comans Christ in Schwacke, and so based on the type of that name.

TYPE: Navarro, Pcia. Cartago, Werckle in 1905 (B-Hb. Christ not seen; isotype
US).

Plants epiphytic, at 1000-2900 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Pcia. Chiriquf. Also from Colombia,
Ecuador, and Brazil.

This species differs only slightly from L. tuerckheimii Maxon (Contr. U. S. Natl.
Herb. 13:23. 1909). The only difference I have seen is that the sporophylls of L.
tortile are often much shorter than the sterile leaves, a condition much less marked
in L. tuerckheimii. If the two are united, L. tortile has priority by a few days.

41. Lycopodium tubulosum Maxon, Contr. U. S. Natl. Herb. 17:178, 1. 10. 1913.
TYPE: Pacayas, at the foot of Volcdn Turrialba, Pcia. Cartago, 1400 m, Biolley

fil. 17398 (US fragm NY).

Plants epiphytic, at 600-2000 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, and the Cordillera de Talamanca to Pcia. Chiriquf.

42. Lycopodium verticillatum var. parvifolium (Werckle ex Nessel) Lellinger,
Proc. Biol. Soc. Wash. 89:719. 1977.

Urostachys verticillatus war. parvifolius Werckle ex Nessel, Revista Sudamer. Bot. 6:163, f. 43. 1940.
TYPE: Volcdin Barba, Pcia. Heredia, 2100 m, Brade & Brade 283 (HB not seen).

Plants epiphytic, at 1100-2400 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, and the Cordillera de Talamanca to Pcia. Chiriquf.
Also from Mexico to Nicaragua.

42 LYCOPODIACEAE: 2. LYCOPODIUM

LYCOPODIACEAE; 2. LYCOPODIUM 43

This variety is slightly distinct from typical material from Reunion in its shorter,
more spreading sterile leaves. It differs from L. tortile in having thinner, laxer, less
appressed sterile leaves, particularly at the base of the stems. 011gaard in Stolze
(Fieldiana, Bot. N.S., 12:32. 1983) considers this to be a species distinct from the
Old World L. verticillatum and calls it L. acerosum Swartz. Specimens from Sitio
Birds, Pcia. Cartago [?] {Pittier 191, US) and Volcan Chiriqui {Davidson 1021, US)
look much like L. verticillatum var. parvifolium but may prove to be a variety of L.
tenue Willd., according to B. 011gaard (in litt.).

43. Lycopodium watsonianum Maxon, Smithsonian Misc. Collect. 56(29) :3, t. 3.
1912.

TYPE: Upper Caldera river, Holcomb’s Trail above Boquete, Pcia. Chiriqui, ca.
1600 m, Maxon 5712 (US fragm NY).

Plants epiphytic, at 900-2700 m elevation, in forests, from Los Angeles de S.
Ramon (Pcia. Alajuela), the Cordillera Central, and the Cordillera de Talamanca
to Pcia. Chiriqui.

This species has pendent stems up to 10-times branched. The sterile leaves are
more spreading than are those of L. verticillatum W3X. parvifolium, which are clearly

ascending.

44. Lycopodium wilsonii Underw. & Lloyd, Bull. Torrey Bot. Club 33:111. 1906.

Lycopodium tiichodendron Herter, Bot. Jahrb. Syst. 43, Beibl. 98:35, 49. 1909. TYPE:

Guadeloupe, Bory 103 (P not seen).

Lycopodium stamineum Maxon, Smithsonian Misc. Collect. 56(29):2, t. 2. 1912. TYPE: Upper
Rio Caldera watershed, Holcomb’s Trail above Boquete, Pcia. Chiriqui, ca. 1750 m, Maxon 5636
(US).

Lycopodium arcanum Maxon ex Yuncker, Field Mus. Nat. Hist., Bot. Ser. 17:310, t. 3. 1938.
TYPE: Above El Achote, Depto. Comayagua, Honduras, 1800 m, Yuncker, Dawson & Youse 6149
(US).

TYPE: Luquillo Mountains, Puerto Rico, Wilson 271 (NY not seen).

Plants epiphytic, at 1700-2700 m elevation, from the Cordillera Central,
Palmira (Pcia. Heredia), the Fila de Cedral, Cerro Carpintera, and the Cordillera
de Talamanca to Pcia. Chiriqui. Also from Jamaica, Puerto Rico, Guadeloupe,
Martinique, and Mexico to Venezuela and Ecuador.

011gaard in Stolze (Fieldiana, Bot. N. S., 12:43-44. 1983) has united L.

stamineum with L. wilsonii. The plants of the former species tend to have
narrower leaves with reddish bases, but the coloration may be inconstant and
specimens of the latter with narrow leaves matching those of L. stamineum are
known.

FIGS. ?n-4A. Lycopodium. FIG. 37. Branch portion of L. saumms, Williams 20052. FIG. 38.
Branch portion of L. subulatum, Tonduz 1958. FIG. 39. Branch portion of L. taxifolium, Allen 1568.
FIG. 40. Branch apex of L. tortile, Standley 35410. FIG. 41. Branch portion of L. tubulosum, Skutch
2389. FIG. 42. Branch portion of L. verticillatum var. parvifolium, Terry 1021. FIG. 43. Branch
portion of L. watsonianum, Woodson & Schery 690. FIG. 44. Branch portion of L. wilsonii, Standley
& Valerio 49898.

44 SELAGINELLACEAE: 3. SELAGINELLA

SELAGINELLACEAE

Monotypic; see description of Selaginella.

3. SELAGINELLA Pal. Beauv.

Plants terrestrial, occasionally epipetric, rarely hemiepiphytic or epiphytic;
stems mostly prostrate and long-creeping, ascending, or erect, rarely caespitose,
branched, ca. 0.5-3 mm in diam., often articulate, usually stramineous or
greenish, often sulcate, bearing often long, wiry, dichotomously branched roots
from the axils of the branches and bearing numerous, small, 1-nerved, usually
ovate or lanceolate, often aristate and ciliate or denticulate leaves, the leaves
monomorphic, crowded, and spirally arranged around the stems or dimorphic,
usually not crowded, and arranged in 2 small, dorsal, appressed ranks and 2
ventral, larger, usually spreading ranks; sporophylls usually monomorphic, slightly
modified, aggregated into usually rather loose terminal strobili; sporangia solitary
in the axils of the sporophylls, the megasporangia bearing 1-4 large, pale, trilete
megaspores, the distal microsporangia bearing many, much smaller, often orange,
trilete microspores.

Pantropical with some temperate and xerophytic species; ca. 700 species.
Selaginella is usually divided into subg. Selaginella, with uniform, spirally arranged
leaves, and subg. Stachygynandrum (Pal. Beauv.) Baker, with dimorphic leaves
arranged in 4 ranks. (Only the latter is represented in the Flora area.) Hellwig
(1969) found that spore characters support the subgeneric classification and point
toward a classification below that level. However, additional characters, such as
habit and stelar structure, need to be taken into account. Some species of
Selaginella are attractive plants in tropical gardens and in greenhouses, especially
S. pallescens, with white-margined leaves, S. umbrosa, with red stems, and S.
exaltata, a large, scrambling species with ample, feathery branches.

ALSTON, A. H. G. 1955. The heterophyllous Selaginellae of continental North America. Bull. Brit.

Mus. (Nat. Hist.), Bot. 1:221-274, t. 5, 6.

, A. C. JERMY and J. M. RANKIN. 1981. The genus Selaginella in tropical South America.

Bull. Brit. Mus. (Nat. Hist.), Bot. 9:233 - 330.

HELLWIG, R. L. 1969. Spores of the heterophyllous Selaginellae of Mexico and Central America.

Ann. Missouri Bot. Gard. 56:444 - 464.

1. Stems not articulate; aerial roots emanating from the abaxial side (with lateral leaves) of the
stems..l7.

1. Stems articulate; aerial roots emanating from the adaxial side (with median leaves) of the stems.
Megaspores often solitary in the basal sporophylls of the strobili; microspores beige..2.

2(1). Branches glabrous..5.

2(1). Branches pubescent..3.

3(2). Plants scandent, 1-2 m long; lateral leaves of the ultimate branches up to 2.5 mm long.
Branches of the main stems pinnate.

55. S. exaltata

3(2). Plants suberect, less than 1 m long; lateral leaves of the ultimate branches 3-7 mm long..4.

4(3). Branches of the main stems dichotomous; lateral leaves entire, with a small, curved auricle at
the acroscopic base.

47. S. articulata

4(3). Branches of the main stems pinnate; lateral leaves distantly denticulate, not auriculate at the
acroscopic base.

83. S. tomentosa

SELAGINELLACEAE: 3. SELAGINELU\ 45

5(2). Main stems branched from near the base, prostrate to erect; leaves dimorphic and the lateral
ones spreading throughout..?.

5(2). Main stems simple, erect, the plants arborescent; leaves subequal and uniformly decreasing
in size towards the base of the plant..6.

6(5). Basiscopic base of lateral leaves truncate. Plants usually ca. 30 cm long; all leaves entire or
the lateral ones shghtly denticulate toward the base; median leaves acute, with 1 large auricle.

46. S. arthritica

6(5). Basiscopic base of lateral leaves cordate-auriculate. Plants 30-60 cm long; leaves all entire
or the lateral ones shghtly denticulate toward the base; median leaves acute, with 2 unequal auricles.

58. S. geniculata

7(5). Lateral leaves ca. 3-4 mm long..9.

7(5). Lateral leaves up to 2(3) mm long. .8.

8(7). Leaves white-margined, the lateral ones long-cihate at the base. Plants prostrate in dense
mats, the branches ascending at the tips; leaves ovate-lanceolate, ca. 2 times longer than wide, acute
at the apex, cihate on the unequal auricles.

48. S. atirrensis

8(7). Leaves not white-margined, the lateral ones entire at the base. Plants prostrate in dense
mats, the branches ascending at the tips; lateral leaves 1.5-2 times longer than wide, acute or
somewhat round at the apex; median leaves ca. 3 times longer than wide, very unequally biauriculate.

46. S. arthritica

9(7). Median leaves with 1 auricle or with 2 unequal auricles, one twice as large as the other.. 12.

9(7). Median leaves with 2 subequal auricles.. 10.

10(9). Median leaves acuminate, their auricles large, 1/4- 1/6 as long as the leaf; lateral leaves
ovate-oblong; plants erect, their stems rooting in the proximal 1/3. Plants usually less than 30 cm
long; stems 1(2) mm in diam. at the base.

57. S. galeottii

10(9). Median leaves aristate, their auricles very short; lateral leaves oblong; plants scandent, then-
stems rootmg in the proximal 2/3.. 11.

11(10). Stems distinctly articulate, trailing; stolons absent. Plants usually 30 - 50 cm long; stems
2-3 mm in diam. at the base.

80. S. silvestris

11(10). Stems obscurely articulate, erect; stolons usually present. Plants ca. 35 cm long; stems
0.75-2 mm in diam. at the base, bearing thin, elongate, trailing soboliferous stems producing an
elongate tuber at their apex.

84. S. tuberosa

12(9). Median leaves biauriculate.. 14.

12(9). Median leaves uniauriculate..l3.

13(12). Main stems commonly flagelliform at the apex; lateral leaves of the main stems ascending.
Axillary leaves biauriculate, the small auricles short-ciliate; median leaves subaristate, evenly
denticulate throughout, not white-margined.

79. S. sertata

13(12). Main stems never flagelliform; lateral leaves of the main stems spreading, parallel-sided.
Axillary leaves biauriculate, the auricles equal, entire or sparsely ciliate; median leaves sparsely
ciliate proximally, denticulate distally, not white-margined.

54. S. eurynota

14(12). Lateral leaves of the main stems ca. 1 mm wide, ca. 8 mm distant, the stems 8-14 mm
wide including the leaves. Axillary leaves exauriculate, denticulate, slightly white-margined; median
leaves acuminate, unevenly biauriculate, entire or denticulate, shghtly white-margined.

64. S. kunzeana

14(12). Lateral leaves of the main stems mostly more than 1.5 mm wide, no more than 5(8) mm
distant, the stems 6-8 mm wide including the leaves.. 15.

46 SELAGINELLACEAE: 3. SELAGINELLA

15(14). Lateral leaves nearly symmetrical, almost exauriculate, entire or sparingly denticulate at
the base. Median leaves aristate, very unequally biauriculate.

80. S. silvestris

15(14). Lateral leaves asymmetrical, with a curving, acroscopic auricle, copiously ciliate at the
base..l6.

16(15). Lateral leaves of the ultimate branches ovate-lanceolate like those of the main stems;
median leaves acuminate, often ciliate. Leaves coriaceous, ciliate at the base, entire distally; midrib
of the median leaves prominent.

52. S. diffusa

16(15). Lateral leaves of the ultimate branches parallel-sided, unlike those of the main stems;
median leaves aristate, entire or denticulate. Plants in prostrate mats, with the fertile branches erect.

61. S. horizontalis

17(1). Stems forming rosettes, curling inwards when dry; leaves usually subcoriaceous. Leaves
white-margined, ciliate, dark green adaxially, the leaves of the main stem pale brown abaxially.

73. S. pallescens

17(1). Stems not forming rosettes; leaves usually membranaceous..l8.

18(17). Stems erect, ascending, or prostrate, not reddish..21.

18(17). Stems erect and reddish, at least at the base..l9.

19(18). Leaves of the main stems proximal to the branched portion dimorphic and the lateral ones
spreading, except at the base of the stem; stems pale reddish. Leaves narrowly white-margined,
faintly denticulate, the median ones aristate, the lateral ones subacute, oblong.

60. S. hoffmannii

19(18). Leaves of the main stems proximal to the branched portion monomorphic, ascending and
appressed to the stem; stems bright red..20.

20(19). Lateral leaves ciliate only at and near the base, the acroscopic margin entire. Plants
usually less than 30 cm long, the frond-like portion deltate, not exceeding 15(20) cm long, 15 cm
wide.

85. S. umbrosa

20(19). Lateral leaves not cihate at the base, the acroscopic margin usually minutely and evenly
denticulate. Plants 20-60 cm long, the frond-like portion broadly lanceate, 15-40 cm long, 15-30
cm wide.

59. S. haematodes

21(18). Leaves on main stems proximal to the branched portion dimorphic, sometimes smaller at
the base of the main stem; most species not arborescent..30.

21(18). Leaves on main stems proximal to the branched portion monomorphic; most species
arborescent..22.

22(21). Branches (including the leaves) up to 10(11) mm wide; strobili always terminal on leafy
branches..24.

22(21). Branches (including the leaves) 13-15 mm wide; strobili mostly appearing lateral on a
leafy branch.. 23.

23(22). Median leaves shortly aristate, denticulate proximal to the apex.

49. S. bombycina

23(22). Median leaves acuminate, entire proximal to the apex.

81. 5. tanyclada

24(23). Basiscopic margin of the lateral leaves round at the base; plants terrestrial, up to 30(35) cm
long (75 cm in S. wolfii) or hemiepiphytic, up to 2 m long. .27.

24(23). Basiscopic margin of the lateral leaves truncate at the base; plants terrestrial, (15)30-75
cm long..25.

25(24). Plants forming scrambling vines. Median leaves evenly biauriculate, acute; lateral leaves
biauriculate, ascending, acute to round at the apex; all leaves entire, lacking cilia.

74. S. plana

SELAGINELLACEAE: 3. SELAGINELLA 47

25(24). Plants arborescent, not scranibling..26.

26(25). Leaves of the main stem ca. 3 mm long, appressed to the stem up to the third to sixth
branch; lateral leaves strongly ciliate.

45. S. anceps

26(25). Leaves of the main stems 5-6 mm long, usually spreading above the first or second
branch; lateral leaves denticulate.

72. S. oaxacana

27(23). Plants hemiepiphytic, up to 2 m long, the main stems at maturity adherent to tree trunks.

65. S. longissima

27(23). Plants terrestrial, up to 75 cm long. .28.

28(27). Branches (including the leaves) 7-12 mm wide; leaves entire, those of the main upright
stems the largest.

87. S. wolfii

28(27). Branches (including the leaves) 2-3 mm wide; leaves denticulate to ciliate, those of the
main upright stems not larger than those of the branches.. 29.

29(28). Lateral leaves of the main stem ovate, more or less appressed, imbricate, sparingly cilate at
the base.

73. S. pallescens

29(28). Lateral leaves of the distal half of the main stem oblong and 2 times longer than wide,
spreading, not imbricate, densely ciliate on the acroscopic base.

86. S. viticulosa

30(21). Lateral leaves with a false vein on either side of the midrib. Lateral leaves distant, broadly
ovate-lanceolate, acute at the apex, the adaxial surface usually pubescent towards the leaf base;
branches sometimes flagelliform and terminated by a bud.

75. S. porphyrospora

30(21). Lateral leaves without false veins..31.

31(30). Main stems (including the leaves) no more than 7(8) mm wide..34.

31(30). Main stems (including the leaves) ca. 10 mm wide..32.

32(31). Median leaves long-ciliate; lateral leaves long-ciliate at the base, denticulate toward the
apex, subacute, round at the acroscopic base, trimcate or cordate at the basiscopic base.

53. S. estrellensis

32(31). Median leaves denticulate; lateral leaves denticulate at the base, entire toward the apex,
obtuse, round at the acroscopic and basiscopic base..33.

33(32). Median leaves aristate; strobili terminal on a leafy branch.

67. S. martensii

33(32). Median leaves acute-acuminate; strobili appearing lateral on a leafy branch.

50. S. chrysoleuca

34(31). Stems suberect; aerial roots confined to the proximal half of the stem..41.

34(31). Stems prostrate or nearly so, rooting throughout..35.

35(34). Branches (including the leaves) mostly ca. 2 mm wide; plants less than 10 cm long. Lateral
leaves broadly ovate-lanceolate; median leaves elliptic-acuminate.

68. S. microphylla

35(34). Branches (including the leaves) more than 3 mm across; plants 10-25 cm long..36.

36(35). Lateral leaves long-ciliate at the acroscopic base; aerial roots ca. 0.25 mm in diam., flexible;
main stems ca. 0.5 mm in diam..39.

36(35). Lateral leaves denticulate at the acroscopic base.. 37.

37(36). Lateral leaves less than 2 mm long; plants 10-15 cm long. Aerial roots ca. 0.25 mm in
diam., flexible; main stems ca. 0.5 mm in diam.

54a. S. euclimax

37(36). Lateral leaves 2.5-4 mm long; plants mostly at least 15 cm long. Aerial roots ca. 0.5 mm
in diam., stiff; main stems ca. 0.75 mm in diam..38.

48 SELAGINELLACEAE: 3. SELAGINELLA

38(37). Lateral leaves ca. 1 mm wide, white-margined at the acroscopic base. Lateral leaves ovate-
lanceolate; median leaves elliptic-aristate, denticulate throughout.

82. S. tarapotensis

38(37). Lateral leaves ca. 1.5-2 mm wide, not white-margined at the acroscopic base. Lateral
leaves asymmetrically ovate; median leaves elliptic-aristate, denticulate only toward the base.

76. S. producta

39(36). Median leaves denticulate toward the apex; leaves membranaceous, curling laterally when
dry. Lateral leaves ovate-lanceolate, round at the apex, sometimes sparsely pubescent adaxially;
median leaves broadly elliptic-aristate.

77. S. revoluta

39(36). Median leaves regularly ciliate toward the apex; leaves not curling when dry..40.

40(39). Median leaves elliptic-aristate; lateral leaves white-margined abaxially, the cilia at the leaf
base ca. 1/4- 1/3 as wide as the leaf; lateral leaves distant on the main stems.

62. S. huehuetenangensis

40(39). Median leaves orbicular-aristate; lateral leaves not white-margined abaxially, the cilia at
the leaf base ca. 1/2 as wide as the leaf; lateral leaves approximate on the main stems.

63. S. idiospora

41(34). Lateral leaves acute and usually slightly acuminate at the apex..44.

41(34). Lateral leaves nearly obtuse to round and not acuminate at the apex..42

42(41). Stems ca. 1 mm in diam.; median leaves ovate-acuminate, denticulate.

78. S. seemannii

42(41). Stems 0.5-0.75 mm in diam.; median leaves elliptic-aristate..43.

43(42). Lateral leaves elliptic, denticulate to entire; sporophylls monomorphic. Median leaves
elliptic-aristate, less than half as wide as the lateral leaves, denticulate; main stems sometimes
flagelliform.

51. S. cladorrhizans

43(42). Lateral leaves obliquely oblong, ciliate to denticulate; sporophylls dimorphic. Median
leaves elliptic-aristate, more than half as wide as the lateral leaves, denticulate.

66. S. lychnuchus

44(41). Lateral leaves entire or ciliolate-denticulate. Main stems flagelliform and rooting at the
apex; lateral leaves ovate, acuminate; median leaves lanceolate-aristate, weakly uniauriculate.

56. S. flagellata

44(41). Lateral leaves ciliate (rarely denticulate in S. novae-hollandiae).A5.

45(44). Plants less than 5 cm long; stems lacking reduced, vegetatively reproductive stolons at the
base. Lateral leaves ovate, acute, long-ciliate at the base; median leaves lanceolate.

69. S. minima

45(44). Plants at least (5)7 cm long; stems often with reduced, vegetatively reproductive stolons at
the base..46.

46(45). Median leaves long-aristate, exauriculate; lateral leaves obliquely oblong, strongly ciliate at
the base.

70. S. mollis

46(45). Median leaves acuminate to weakly aristate, very unevenly biauriculate; lateral leaves
broadly ovate, weakly ciliate almost to the apex.

71. S. novae-hollandiae

45. Selaginella anceps (K. Presl) K. Presl, Abh. Konigl. Bohm. Ges. Wiss. V, 3:581
(repr. 151). 1845.

Lycopodium gracile Desv. ex Poir. Encyc. M6th. Suppl. 3:551. 1814. TYPE LOCALITY: Peru.

Lycopodium anceps K. Presl, Reliq. Haenk. 1:80. 1825. TYPE: “Hab. in insula Luzon” [actually
Peru], Haenke (PRC not seen).

Sela^nella gracilis (Desv. ex Poir.) Hieron. Hedwigia 58:292. 1917, non Moore, 1886, nom. illeg.

SELAGINELLACEAE: 3. SEU\GINELLA 49

Plants terrestrial, at 0-1000(1700) m elevation, in or at the edges of forests,
often covering large areas, from throughout the Flora area. Also from Venezuela
to Bolivia.

46. Selaginella arthritica Alston, Arch. Bot. Sist. 11:43. 1935.

TYPE: Golfo Dulce, near Puerto Jimenez, Pcia. Puntarenas, 30 m, Cufodontis
218 (BM not seen fragm US: isotype W not seen).

Plants terrestrial, at 0-900(1200) m elevation, in forests, throughout Costa Rica
and Panama and from the Rio Nuqui, Depto. Choc6 (Haught 5460, BM, US).
Also from Nicaragua and Colombia.

This species has both a prostrate, juvenile, usually sterile phase with upright
branches up to ca. 8 cm long and an arborescent, mature, fertile phase with
upright branches over 30 cm long. Although the two phases are completely unlike
in habit, the leaves of both are similar in their details. Occasionally the lateral
branches of S. arthritica become abnormally long and bifurcate, bud, or root at the
apex.

47. Selaginella articulata (Kunze) Spring, Flora 21:182. 1838.

Lycopodium articulatum Kunze, Linnaea 9:10. 1834. TYPE: Tocache Mission, at the Rio

Huallaga, Depto. Loreto, Peru, Jul- Aug 1830, Poeppig (LZ destroyed).

Plants terrestrial, at 500-1000 m elevation, in forests, from around El Valle,
Sta. Rita ridge (Pcia. Colon), and Campo Tres (Pcia. Panama). Also from
Venezuela to Bolivia.

48. Selaginella atirrensis Hieron. in Engl. & Prantl, Nat. Pflanzenfam. 1(4):711.
1901.

LECTOTYPE: Atirro, Pcia. Cartago, 2000 ft,/. D. Smith 5103 (US; isolectotype
NY), chosen by Alston (Bull. Brit. Mus. (Nat. Hist.), Bot. 1:259. 1955).

Plants terrestrial, at 0-2400 m elevation, in forests, from scattered localities in
the Cordillera Central and the Atlantic coastal plain through the Choco. Also
from Venezuela to Peru.

This species exists in a low-altitude (up to ca. 600 m), rather erect growth form
and in a high-altitude, creeping form. The leaves in both forms are variable,
especially in marginal cilia length and in the width of the whitish, marginal band
around the leaves. Neither of these characters correlates with habit. Bruce
McAlpin (pers. comm.) found that the two forms maintained themselves under
conditions of uniform culture. Although it is possible that two species are
involved, I have not been able to separate the specimens by any character other
than habit.

49. Selaginella bombycina Spring, Nouv. M6m. Acad. Roy. Sci. Bruxelles 24:191.
1849.

TYPE: Peru, Mathews (K not seen).

Plants terrestrial, at 300-800 m elevation, mostly in wet forests, from the
Atlantic side of the Cordillera Central, western Panama, and scattered localities in
the Choco. Also from Colombia to Peru.

50. Selaginella chrysoleuca Spring, Bull. Acad. Roy. Sci. Bruxelles 10:226. 1843.

Selaginella sprucei Hook. Sec. Cent. Ferns t. 83. 1861. TYPE: Mt. Campana, near Tarapoto,
Depto. S. Martm, Peru, Spruce 4623 (K not seen; isotypes BM, CGE neither seen, US).

50 SELAGINELLACEAE: 3. SELAGINELLA

SELAGINELLACEAE: 3. SELAGINELLA 51

Seiagineila centipediformis Gomez, Phytologia 52:153. 1982. TYPE: Fortuna hydroelectric project,
Pcia. CMriqm, 1200-14430 m, Hammei 2121 (MO).

TYPE: Bolivia, D'Orbigny (P not seen).

Plants terrestrial, at 1100“ 1400 m elevation, in the Flora area known only from
the Fortuna hydroelectric project area. Also from Venezuela to Bolivia.

This species, like S. bombydna, is unusual in bearing the strobili apparently
laterally, rather than at the apex of leafy branches.

51. Seiagineila cladorrhizans A. Braun, Ann. Sci. Nat. Bot. V, 3:282. 1865.

Seiagineila ambigua A. Braun, Ann. Sci. Nat. Bot. V, 3:286. 1865. TYPE: Qnebrada del

Imposible, Cuman^ Edo. Sucre, Venezuela, Moritz 221 (B not seen; isotype BM not seen Morton
photo 6522).

SYNTYPES: Colonia Tovar, Edo. Aragua, Venezuela, Moritz 448 (B not seen);
and Fendier 324 p. p. (B not seen; isosyntype NY).

Plants terrestrial, at 100-800 m elevation, in swamps and forests, from along
the Aiajuela - Grecia Highway and the cliffs of the R. Rosales (Pcia. Alajuela) and
scattered localities throughout Panama. Also from Mexico to Honduras,
Nicaragua, Trinidad, Tobago, and Venezuela.

The median leaves of this species are unusually small in proportion to the
lateral leaves.

52. Seiagineila diffusa (K. Presl) Spring, Bull. Acad. Roy. Sci. Bruxelles 10:143.
1843.

Lycopodium diffitsum K. Presl, Reliq. Haenk. 1:78. 1825. TYPE: Panama, Haenke (PRC 24294
not seen photo at F).

Lycopodium mnioides Sieber ex Hook. & Grev, Bot. Misc. 2:394. 1831. TYPE: Trinidad, Sieber
FL Mba. 325 (K? not seen).

Seiagineila ciliauricula Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:219. 1849. SYNTYPES:
Merida, Edo. Merida, Venezuela, Moritz 380 (B not seen; isosyntype LG not seen); and Chivada de
Sta. Marta, Depto. Magdalena, Colombia, Purdie (K not seen), synonymized by Alston (J. Bot. Brit.
For. 72:36. 1934).

ISelaginella cirrhipes Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:221. 1849. TYPE: Galip^,
Distr. Fed., Venezuela, Moritz 379 (LG not seen).

Sela^nella pofycephaia Baker, J. Bot. Brit. For. 21:332. 1883. SYNTYPES: Mountains of Ocana,
Depto. Norte de Santander, Colombia, 5000 - 6000 ft, Schlim 493 (K not seen fragm NY) and
Colombia, Holton 82 (K not seen).

Plants terrestrial, at (200)800-1700 m elevation, in forests and pastures on wet
banks and tree bases, from the Cordillera Central, the northern foothills of the
Cordillera de Talamanca, the Peninsula de Osa, El Valle, north of Gamboa
(Canal Zone), and the Serrama de Baudo (Depto. Choco). Also from Trinidad
and Venezuela to Ecuador.

53. Seiagineila estrellensis Hieron. Hedwigia 41:200. 1902.

LECTOTYPE: Estrella, Pcia. Cartago, 4400 ft, Cooper 6062 (US; isolectotype B
not seen), chosen by Alston (Bull. Brit. Mus. (Nat. Hist), Bot. 1:249. 1955).

FIGS. 45-48. Seiagineila. FIG. 45. Branch portions of S. anceps, Lewis 2067. FIG. 46. Branch
portions of S. arthritica, Standley 45278. FIG. 47. Branch portions of S. articulata, Hunter & Allen
308. FIG. 48. Branch portions of S. atirrensis^ Stork 4629.

52 SELAGINELLACEAE: 3. SELAGINELLA

50 S. chrysoleuca

SELAGINELLACEAE: 3. SELAGINELLA 53

Plants terrestrial, at 1200-2400(3200) m elevation, in forests, from the
Cordillera Central and the Cordillera de Talamanca to Pda. Chiriqui. Also from
Honduras to Nicaragua.

53a. Selaginella eedimax Alston ex Crabbe & Jermy, Fem Gaz. 11:259. 1976.

TYPE: West of Tambo, Depto. Cauca, Colombia, 2300 m, Haught 5220 (BM
not seen; isotypes COL not seen, US).

Plants terrestrial, at 1200 m elevation, in forests, in the Flora area known only
from the trail west of the continental divide on the road from Fortuna Lake to
Chiriqui Grande, Pda. Chiriqui (Hampshire & Whitefoord 8456, BM, CR neither
seen, US). Also from Colombia.

54. Selaginella euiynota A. Braun, Ann. Sci. Nat. Bot. V, 3:293. 1865.

TYPE: Near Aguacate, Pda. S. Jose, Hoffmann 907 (B not seen; isotype K not
seen).

Plants terrestrial, at 0-1000 m elevation, in wet and moist forests, from the
Cordillera de Tilaran to the Canal Zone, and also in the central Choco. Also from
Guatemala and Nicaragua.

55. Selaginella exaltata (Kunze) Spring, Bull. Acad. Roy. Sci. Bruxelles 10:234.
1843.

Lycopodium exaltatum Kunze, Linnaea 9:8. 1834. TYPE: From Uchiza to Tocache, Depto.
Loreto, Peru, Poeppig 1953 (LZ destroyed; isotype LG not seen).

Selaginella strobiiifem Christ, Bull. Herb. Boissier 11, 1:72. 1901. TYPE: Between Ucayali and
Huallaga, Depto. Loreto, Peru, Huber 1515 (P not seen).

Plants terrestrial, at 0-200(400) m elevation, in wet forests, from the Peninsula
de Osa (Pcia. Puntarenas) and the lowlands of Panama and the Choco. Also from
other areas in Colombia and from Ecuador to Bolivia.

This species is always erect when fertile, but may be prostrate when
sterile.

56. Selaginella flagellata Spring, Bull. Acad. Roy. Sci. Bruxelles 10:228. 1843.

Selaginella rhizophora Baker, J. Bot. Brit. For. 22:244. 1884. SYNTYPES: Near the towns of S.
Juan and Panama, Pcia. Panama, Seemann 29 and 32 (both K neither seen fragms NY).

TYPE: Upper Oyapok River, French Guiana, Leprieur (LG not seen; isotypes
B, G, P none seen).

Plants terrestrial or epipetric, at 0-300(600) m elevation, in forests and stream
beds, from the Atlantic and Pacific lowlands of Costa Rica to the Canal Zone and
Pda. Panama. Also from Mexico, Guatemala, Nicaragua, Trinidad and Tobago,
and Venezuela to Bolivia, Brazil, and the Guianas.

57. Selaginella galeottii Spring, Bull. Acad. Roy. Sci. Bruxelles 10:230. 1843.

? Sela^nella poeppigiana var. mexicana Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:218.
1849. TYPE: Near Jalapa, Edo. Veracruz, Mexico, Schiede (B not seen; possible isotype BM not
seen).

FIGS. 49 - 52. Selaginella. FIG. 49. Branch portions of S. bombycina, Hart 61. FIG. 50. Branch
portions of S. chrysoieuca, Hammel 2121. FIG. 51. Branch portions of S. cladorrhizans, Standley
31032. FIG. 52. Branch portions of S. diffusa, Maxon 7935.

54 SELAGINELLACEAE; 3. SELAGINELLA

SELAGINELLACEAE: 3. SELAGINELLA 55

LECTOTYPE: Edo. Veracruz, Mexico, Galeotti 6606 (P not seen photo BM not
seen; isolectotype LG not seen), chosen by Alston (Bull. Brit. Mus. (Nat. Hist.),
Bot 1:259. 1955).

Plants terrestrial, at (600)1000” 1700 m elevation, in forests, from north of S.
Ramon, the Cordillera Central, the northern foothills of the Cordillera de
Talamanca, the Fortuna dam site (Pcia. Chiriqui), and the Rio dos Bocas valley
northwest of Sta Fe. Also from Mexico, Guatemala, Belize, and Honduras.

58. Selaginella geniculata (K. Presl) Spring, Bull. Acad. Roy. Sci. Bruxelles
10:230. 1843.'

Lycopodium genicuiatum K. Presl, Reiiq. Haenk. 1:80. 1825. TYPE: “In insula Luzon” [probably
Guayaquil, Ecuador or Depto. Huanuco, Peru], Haenke (PRC not seen).

Selagineiia ferruminata Spring, Bull. Acad. Roy. Sci. Bruxelles 10:231. 1843. TYPE: Pangoa,
Depto. JuniE, Peru, Mathews 1083 (P not seen; isotypes FI not seen, K not seen, US).

Seiaginella elongata Klotzsch, Linnaea 18:522. 1845. TYPE: Peru?, Poeppig 189 (not seen),

according to Reed (Mem. Soc. Brot. 18:105. 1965-1966).

Plants terrestrial, at 0-100 m elevation, in wet forests, from the Depto. Choco.
Also from other areas in Colombia and Ecuador to Pern.

59. Selaginella haematodes (Kunze) Spring in Mart. FI. Bras. 1(2): 126. 1840.

Lycopodium haematodes Kunze, Linnaea 9:9. 1834. TYPE: Tocache Mission, upper Rio

Huallaga, Depto. S. Martm, Peru, Jun.e 1830, Poeppig (LZ destroyed).

Seiaffneiia fiticina Spring, Nouv. M6m. Acad. Roy. Sci. Bruxelles 24:189. 1849. SYNTYPES:
Between Puerto Cabello and Valencia, Edo. Carabobo, Venezuela, 2000 - 3000 ft, Linden (BR not
seen); “Prov. Caracas,” Venezuela, 3000 ft, Linden 3313 (LG not seen); and Peru, Mathews (K not
seen).

Plants terrestrial, at 0-600 m elevation, in wet forests, from scattered localities
in central Panama through the Choco. Also from Venezuela to Bolivia.

60. Selaginella hoffmannii Hieron. Hedwigia 41:184. 1902.

SYNTYPES: Near Aguacate, Pcia. S. Jose, Hoffmann 905, 905a, and 909 (all B
none seen), and others from Mexico and Guatemala.

Plants terrestrial, at 100-1000 m elevation, in moist or seasonally dry places,
from Nicoya (Pcia. Guanacaste), the western slopes of the Cordillera Central, the
valley of the Rio Reventazon near Turrialba (Pcia. Cartago), and the vicinity of
Boquete. Also from Mexico to Nicaragua.

61. Selaginella horizontalis (K. Presl) Spring, Bull. Acad. Roy. Sci. Bruxelles
10:226. 1843.

Lycopodium horimntale K. Presl, Reiiq. Haenk. 78. 1825. TYPE: “Ex vallibus Cordilieris

Peruanis,” Haenke (PRC not seen photo US).

Selagnella fendleri Baker, J. Bot. Brit. For. 21:334. 1883. TYPE: Chagres, Canal Zone, Fendler
382 (K not seen fragm NY; isotype MO),

Selagineiia sylvatica Baker, J. Bot. Brit. For. 22:25. 1884. TYPE: Near Panama City, Pcia.
Panama, Seemann 31 (K not seen fragm NY).

Plants terrestrial, at 0-800(1 100) m elevation, in forests, from scattered
localities in the Cordillera Central, near Platanillo (Pcia. Cartago), and

FIGS. 53-56. Selagineiia. FIG. 53. Branch portions of S. estrellensis, Standley 35620. FIG. 54.
Branch portions of S. eurynota, Standley & V^erio 47175. FIG. 55. Branch portions of S. exaltata,
Stern et al. 129. FIG. 56. Branch portions of S. flagellata, Killip 39970.

56 SELAGINELLACEAE: 3. SELAGINELLA

.25 cm

SELAGINELLACEAE: 3. SELAGINELLA 57

throughout Panama and the Choc6. Also from Venezuela and other localities in
Colombia.

Singer et al. (Science 172:1341-1342. 1971) reported that this plant serves as
food for the larvae of the butterfly Euptychia westwoodi Butler.

62. Selaginella huehuetenangensis Hieron. Hedwigia 43:32. 1904.

SYNTYPES: Between Sta. Cruz Almor and Ixcan, Depto. Huehuetenango,

Guatemala, Bernoulli & Cario 178 (B not seen); and between Dolores and
Jachicha, Depto. Alta Verapaz, Guatemala, Bernoulli & Cario 183 (B not seen).

Plants terrestrial, often on roadside banks, at 0-700(1400) m elevation, mostly
in lowland forests, from the Cordillera de Tilardn, the Cordillera Central, and the
Llanuras de S. Carlos to Pcia. Darien. Also from Mexico to Honduras, and
Nicaragua.

63. Selaginella idiospora Alston, Bull. Brit. Mus. (Nat. Hist.), Bot. 1:246. 1955.
TYPE: Along the Saklak River below Secanquim, Depto. Alta Verapaz,

Guatemala, 550 m, Pittier 191 (BM not seen; isotype US).

Plants terrestrial, at 600-800 m elevation, in forests, from the Cordillera de
Tilaran to the western slopes of the Cordillera Central. Also from Mexico to
Honduras.

64. Selaginella kunzeana A. Braun, Ann. Sci. Nat. Bot. V, 3:296. 1865.

TYPE: Braun cites several syntypes, some from Peru and some with a query.
The others, which are from the Prodromus area, are: Near La Palma, Depto.
Cundinamarca, Colombia, Triana 3 (B not seen); and Muzo, Minas, Depto.
Boyaca, Colombia, 700 m, Lindig 1513 (B not seen).

Plants terrestrial, at 0-600(1000) m elevation, mostly in lowland forests, often
on exposed banks, from the Cordillera Central, the Cordillera de Talamanca, the
Fila Costeha, west of Cerro Pando (Pcia. Chiriqui), and the Choco. Also from
Mexico, Honduras, Venezuela, and other localities in Colombia to Peru.

This species is closely allied to S. eurynota and S. sertata.

65. Selaginella longissima Baker, J. Bot. Brit. For. 19:208. 1881.

TYPE: Depto. Antioquia, Colombia, 3000 ft, Kalbreyer 1815 (K not seen).

Plants hemiepiphytic, at 0-1800 m elevation, from the Choc6. Also from other
areas in western Colombia.

66. Selaginella lychnuchus Spring, Nouv. M6m. Acad. Roy. Sci. Bruxelles 24:247.
1849.

Selaginella lychnuchus var. flaccida Spring, Nouv. M6m. Acad. Roy. Sci. Bruxelles 24:248. 1849.
TYPE: Galipdn, Distr. Fed., Venezuela, Moritz 71 (LG not seen; isotype BM not seen photo 6523).

Selagnella lychnuchus var. rigdiuscula Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:248. 1849,
nom. superfl. TYPE: A renaming of the typical variety, and so based on the type of the species
name.

Selaginella lychnuchus var. pusilla A. Braun, Ann. Sci. Nat. Bot. V, 3:288. 1865. TYPE: Caracas,
Distr. Fed., Venezuela, Gollmer (B not seen).

FIGS. 57 - 60. Selaginella. FIG. 57. Branch portions of S. galeottii, Pittier 1820. FIG. 58. Branch
portions of S. geniculata, Haught 5495. FIG. 59. Branch portions of S. haematodes, Burch et al. 1079.
FIG. 60. Branch portions of S. hoffmannii, Cornman 1167.

58 SELAGINELLACEAE: 3. SELAGINELLA

.25 cm

SELAGINELLACEAE: 3. SELAGINELLA 59

TYPE: Merida, Edo. Merida, Venezuela, Moritz 378 (B not seen; isotype BM
not seen photo 6525).

Plants terrestrial, at 1200-1700 m elevation, from near S. Ramon, La Hondura
and Cerro Tablazo (both Pda. S. Jose), and Tapanti. Also from Venezuela.

67. Selaginella martensii Spring, Nouv. M6m. Acad. Roy. Sci. Bruxelles 24:129.
1849.

Selaginella solmsii Baker, Handb. Fern-allies 56. 1887. TYPE: Guatemala, Bernoulli & Cario 181
(K not seen).

LECTOTYPE: Near Jalapa and Mirador, Edo. Veracruz, Mexico, Galeotti
6606bis (LG not seen; isolectotype ''6606*' BR not seen), chosen by Gregory and
Riba (FI. Veracruz 6:16. 1979).

Plants terrestrial, at (700)1300-2500 m elevation, in forests, from the
Cordillera Central, the north end of the Cordillera de Talamanca, around
Boquete, from La Mesa to El Valle, and the Serrama de Pirre near Cana (Pda.
Darien). Also from Mexico to Nicaragua.

The median leaves of this species have longer aristae than do those of the
superficially similar 5. oaxacana.

68. Selaginella microphylla (H.B.K.) Spring, Bull. Acad. Roy. Sci. Bruxelles
10:234. 1843.

Lycopodium microphyllum H.B.K. Nov. Gen. Sp. 1:39 (fol. 32). 1816. TYPE: Near Quilquase,
Depto. Cauca, Colombia, 1004 hexap, Humboldt & Bonpland (B not seen; presumable isotype P not
seen photo 3347).

Selagnella thujifolia Spring in Mart. FI. Bras. 1(2):120. 1840, as '‘thujaefolia.” TYPE:

Montevideo, Uruguay, Sellow (BR? not seen).

Selagnella jamesonii Baker, J. Bot. Brit. For. 21:97. 1883. SYNTYPES: Near Quito, Ecuador,
Jameson 312 (K not seen) and Sodiro (not seen).

Selaginella schmidtchenii Hieron. Hedwigia 43:40. 1904. TYPE: Colombia, Schmidtchen in 1882
(B not seen).

Plants terrestrial or epipetric, at 0-500 m elevation, in lowland wet to moist
forests, from the S. Ramon -S. Mateo road (Pcia. Alajuela), the Peninsula de Osa,
and S. Jose Island (Pcia. Panama). Also from Venezuela, Ecuador and Colombia
to Argentina, Paraguay, Uruguay, and Brazil.

69. Selaginella minima Spring, Bull. Acad. Roy. Sci. Bruxelles 10:139. 1843.

TYPE: Mt. Eigre, He de Cayenne, French Guiana, Leprieur 158 (LG not seen;
isotypes G, K, NY, and P none seen, US).

Plants terrestrial, often on banks, at 0-1300 m elevation, in grassy areas, from
the Rio Rosales, Grecia, and La Palma de S. Ram6n (all Pcia. Alajuela) and S.
Jose Island (Pcia. Panama). Also from French Guiana.

70. Selaginella mollis A. Braun, Ann. Sci. Nat. Bot. V, 3:276. 1865.

Selaginella illecebrosa Alston, Bull. Brit. Mus. (Nat. Hist.), Bot. 1:239, t. 5, f. A-E. 1955. TYPE:
Along lower part of stream tributary to the Rio Frio, Depto. Izabal, Guatemala, 75-150 m,
Steyermark 41579 (BM not seen; isotypes NY, US).

FIGS. 61-64. Selagnella. FIG, 61. Branch portions of S. horizontalis, Aviles 2. FIG. 62. Branch
portions of 5. huehuetenangensis, Chrysler 5172. FIG. 63. Branch portions of S. idiospora, Burger &
Stolze 5110. FIG, 64. Branch portions of S. kunzeana, Tonduz 4678.

60 SELAGINELLACEAE; 3. SELAGINELLA

SELAGINELLACEAE: 3. SEU\GINELLA 61

TYPE: “Prov. Ocana,” Depto. Norte de Santander, Colombia, 4000-6000 ft,
Schlim 1029 (G not seen; isotype K not seen).

Plants terrestrial, often on banks, at 0-600 m elevation, in wet forests, from the
lowlands and foothills of Costa Rica, Panama, and the northern Choco. Also from
Mexico, Nicaragua, and Colombia adjacent to the Choco.

71. Selaginella novae-hollandiae (Swartz) Spring, Bull. Acad. Roy. Sci. Bruxelles
10:234. 1843.

Lycopodium novae-hollandiae Swcirtz, Syn. Fil. 184, 410. 1806. TYPE LOCALITY: “Australia”
[probably western South America].

Plants terrestrial, at 0 - 900 m elevation, often on banks in open areas, from the
Atlantic lowlands, the Cordillera Central, and the Meseta Central to Pcia. Darien.
Also from Nicaragua, Venezuela, Colombia to Bolivia, and Argentina.

72. Selaginella oaxacana Spring, Nouv. M^m. Acad. Roy. Sci. Bruxelles 24:177.
1849.

Lycopodium flabellatum var. strictum Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles
15:12. 1842. TYPE: Chinantla, Edo. Oaxaca, Mexico, Galeotti 6608 (BR not seen).

Sela^nella wendlandii Hieron. in Engl. & Prantl, Nat. Pflanzenfam. 1(4):683. 1901.

LECTOTYPE: Near S. Miguel, Costa Rica, Wendland 771 (B not seen), chosen by Alston (Bull.
Brit. Mus. (Nat. Hist.), Bot. 1:277. 1955).

ISela^nella costaricensis Hieron. in Engl. & Prantl, Nat. Pflanzenfam. 1(4):683. 1901. TYPE
LOCALITY: Costa Rica. Synonymized by Alston (Bull. Brit. Mus. (Nat. Hist.), Bot. 1:237. 1955).

TYPE: Chinantla, Edo. Oaxaca, Mexico, Galeotti 6608bis (LG? not seen;
isotypes ''6608'' BR not seen, P not seen).

Plants terrestrial, at 0-1600 m elevation, on banks, often along streams, from
the Cordillera de Tilaran and the Cordillera Central to El Valle, the Serram'a de
Pirre above Cana, and the central Choco. Also from Mexico to Honduras,
Nicaragua, other localities in Colombia, and Ecuador.

73. Selaginella pallescens (K. Presl) Spring in Mart. FI. Bras. 1(2): 132. 1840.

Lycopodium pallescens K. Presl, Reliq. Haenk. 1:79. 1825. TYPE: Mexico, Haenke (PRC not
seen).

Lycopodium cuspidatum Link, Hort. Reg. Bot. Berol. 2:161. 1833. TYPE: From material

cultivated at the Botanical Garden in Berlin, originally from Mexico (B not seen).

Selaginella cuspidata var. elongata Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:67. 1849.
TYPE LOCALITY: Guatemala.

Selagnella incana Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:157. 1849. TYPE: Realejo,
Nicaragua, Sinclair (K not seen).

Selaginella emmeliana van Geert, R6v. Hort. 10:220, f. 20. 1884. TYPE: Quinquina [Cinchona?],
South America, Emmel (BR? not seen).

Selagnella microdendron Baker, J. Bot. Brit. For. 23:116. 1885. TYPE: Cuba, Wright 3910 (K not
seen; isotypes BM not seen, US).

Selagnella harrisii Underw. & Hieron. in Urban, Symb. Antill. 7:162. 1912. TYPE: Jamaica,
Harris 7587 (NY not seen; isotypes BM, IJ, K none seen).

FIGS. 65-69. Selaginella. FIG. 65. Branch portions of S. longissima. Archer 1999. FIG. 66. Branch
portions of S. lychnuchus, Biolley. FIG. 67. Branch portions of S. martensii, Killip 5251. FIG. 68.
Branch portions of S. microphylla, Erlanson 164. FIG. 69. Branch portions of S. minima, Johnston
878.

62 SELAGINELLACEAE: 3. SELAGINELLA

SELAGINELLACEAE: 3. SELAGINELLA 63

Plants epipetric or terrestrial, at 100-1300(2000) m elevation, in moist to dry,
shady to open places, often on walls or clay banks, from the Pacific side of the
Cordillera de Tilaran, Cartago and Orosi (Pcia. Cartago), Cerro de Piedra Blanca
(Pcia. S. Jose), the west and south sides of the Cordillera de Talamanca to Pcia.
Veraguas, and the vicinity of Cerro Jefe. Also from Jamaica, Mexico to
Nicaragua, Venezuela, and Colombia.

This species is variable in its habit, and includes both arborescent and rosette
forms. At lower elevations it is commonly associated ^ih Adiantum concinnum.

74. Selaginella plana (Desv. ex Poir.) Hieron. in Engl. & Prantl, Nat. Pflanzenfam.
1(4):703. 1901.

Lycopodium planum Desv. ex Poir. Enc. Meth. Suppl. 3:554. 1814. TYPE: India orientalis,
collector unknown (presumaby P-Hb. Desv. not seen).

Plants terrestrial, at 0-700 m elevation, naturalized in waste places and
plantations, from Limdn and Moin (Pcia. Limon) and Cerro Campana (Pcia.
Panama). Also naturalized in Jamaica, Puerto Rico, the Lesser Antilles, Trinidad,
and Brazil.

According to B. McAlpin (pers. comm.), the similar S. uncinata (Desv.) Spring
occurs at the Summit Botanical Garden in the Canal Zone, but apparently has not
yet escaped.

75. Selaginella porphyrospora A. Braun, Ann. Sci. Nat. Bot. V, 3:286. 1865.

Sela^nella bulbifera Baker, Card. Chron. 1867: 783, 950. 1867. TYPE: Presumably based on
material cultivated at Kew (K not seen).

Sela^nella binervis Liebm. ex Baker, J. Bot. Brit. For. 22:112. 1884. TYPE: Dos Puentes, S.
Antonio Huatusco, Edo. Veracruz, Mexico, Liebmann (K not seen; isotype C not seen).

Selaginella bemoullii Hieron. Hedwigia 41:192. 1902. SYNTYPES: Between Cubulco and

Joyabaj, Depto. Quiche, Guatemala, Bernoulli 1121 (B not seen) and Bernoulli & Cario 160 (B not
seen).

TYPE: Edo. Veracruz, Mexico, Sartorius (B not seen fragm BM not seen).

Plants terrestrial or epipetric, at (400)1200-2100 m elevation, often on banks,
from the Cordillera Central and the Cordillera de Talamanca to Pcia. Chiriqui.
Also from Mexico to Nicaragua, and Venezuela.

Some specimens from the Flora area have atypical, 1-nerved leaves.

76. Selaginella producta Baker, J. Bot. Brit. For. 21:243. 1883.

Sela^nella tobagensis Hieron. in Urban, Symb. Antill. 3:524. 1903. TYPE: Morne d’Or, Tobago,
1200 ft, E^ers 5815 (B not seen; isotype US).

SYNTYPES: British Guiana, Appurty Drake; Amazon Valley, Est. Amazonas,
Brazil, Spruce 2043; Est. Minas Gerais, Brazil, Lindberg (all K none seen).

Plants terrestrial, at 0-100 m elevation, in rain forests, in the Flora area known
only from La Concepcion, 15 km E of Quibdo, Depto. Choco {Archer 1996, US).
Also from other localities in Colombia, Venezuela, Trinidad, Tobago, Brazil and
the Guianas, and Peru.

FIGS. 70 - 75. Selaginella. FIG. 70. Branch portions of S. mollis, Wagner. FIG. 71. Branch portions
of S. novae-hollandiae, Burch et al. 1088. FIG. 72. Branch portions of S. oaxacana, Lellinger 909.
FIG. 73. Branch portions of S. pallescens, Haupt 73. FIG. 74. Branch portions of 5. plana, McAlpin
2434. FIG. 75. Branch portions of S. porphyrospora, Brown CR-233.

64 SELAGINELLACEAE; 3. SELAGINELLA

SELAGINELLACEAE: 3. SELAGINELU\ 65

77. Selaginella revoluta Baker, J. Bot. Brit. For. 21:141. 1883.

Sela^nella platybasis Baker, J. Bot. Brit. For. 21:242. 1883. TYPE: Near Panure, Est. Amazonas,
Brazil, Spruce 2502 (K not seen).

Sela^nella demissa Christ, Bull. Herb. Boissier II, 1:75. 1900. TYPE: Cerro de Canchahuaya,
Ocayali, Depto. Loreto, Peru, Huber 1421 (P not seen).

TYPE: Near Maupures, along the Rio Ornioco, Edo. Amazonas, Venezuela,
Spruce 3621 (K not seen; isotypes BM not seen, CGE not seen, US).

Plants terrestrial, at 0-1000 m elevation, in forests, from along the Pan
American Highway above S. Isidro del General, El Valle, and the Canal Zone.
Also from Venezuela to the Guianas and Brazil, Colombia, and Peru.

78. Selaginella seemannii Baker, J. Bot. Brit. For. 21:244. 1883.

Selaginella barbacosensis Hieron. Hedwigia 43:46. 1904. TYPE: Near Barbacoas, Depto. Narino,
Colombia, Lehmann 89 (B not seen).

TYPE: Cachual Island, mouth of the Rio S. Juan, Seemann 1006 (K not seen;
isotype BM not seen photo 6517).

Plants terrestrial, at ca. 0 m elevation, in the Flora area known only from the
type locality. Also from Colombia and Peru.

79. Selaginella sertata Spring, Nouv. M^m. Acad. Roy. Sci. Bruxelles 24:104.
1849.

Selaginella nicaraguensis Baker, J. Bot. Brit. For. 21:333. 1883. TYPE: Grenada, Nicaragua, Levy
360 (K not seen fragm NY).

TYPE: Panama [Nicoya, Pcia. Puntarenas, Costa Rica], Sinclair (K not seen
fragm B not seen).

Plants terrestrial, at 0-300(900) m elevation, in moist and wet forests, from the
Pacific-facing and coastal slopes of central Costa Rica, the Peninsula de Nicoya, S.
Felix to Cerro Flor (Pcia. Chiriqui), and the Canal Zone. Also from Mexico to
Nicaragua and Colombia.

80. Selaginella silvestris Aspl. Ark. Bot. 20A(7): 30, f. 3-5. 1926.

TYPE: El Chaco, S. Yungas, Depto. La Paz, Bolivia, 1900 m, Asplund 1140
(UPS not seen).

Plants terrestrial, at 0-2400 m elevation, in forests, from the Cordillera de
Tilaran to the northern foothills of the Cordillera de Talamanca, the Atlantic
lowlands of Bocas del Toro to the Canal Zone, the mountains beyond La Pintada
(Pcia. Code), and between Bahia Solano and El Valle (Depto. Choco). Also from
Mexico to Nicaragua and other localities in Colombia to Bolivia.

81. Selaginella tanyclada Alston ex Crabbe & Jermy, Amer. Fern J. 63:143. 1973.

TYPE: La Concepcion, 15 km E of Quibdo, Depto. Choco, ca. 75 m. Archer
1966 (US fragm BM not seen).

Plants terrestrial, at 0-1000 m elevation, in forests, from Campo Tres (Pcia.
Panama), the type locality, and Alto del Buey (9 June 1940, von Sneidem, S not
seen).

FIGS. 76-79. Selaginella. FIG. 76. Branch portions of S. producta. Archer 1996. FIG. 77. Branch
portions of S. revoluta^ Steyermark & Allen 17469. FIG. 78. Branch portions of S. seemannii, Killip
& Smith 29638, Peru. FIG. 79. Branch portions of S. sertata, Lankester 603.

66 SEU\GINELLACEAE: 3. SELAGINELLA

SELAGINELLACEAE: 3. SELAGINELLA 67

82. Selaginella tarapotensis Baker, J. Bot. Brit. For. 21:98. 1883.

Selaginella faucium Liebm. ex Baker, Handb. Fern-allies 58. 1887. TYPE: Huitamalco and

Hacienda de Jovo, Edo. Veracruz, Mexico, Liebmann PI. Mex. 2048 (K not seen fragm NY).

TYPE: Mt. Guayapurima, near Tarapoto, Depto. S. Martin, Peru, Spruce 4625
(K not seen; isotypes BM, CGE, LE none seen; US).

Plants terrestrial, at 500-2700 m elevation, in forests, from the Cordillera
Central, Cerro Tute (Pda. Veraguas), and Pda. Darien. Also from Colombia to
Bolivia.

83. Selaginella tomentosa Spring, Nouv. M^m. Acad. Roy. Sci. Bruxelles 24:231.
1849.

TYPE: Gorgona Island, Colombia, Hinds (LG not seen; isotype K not seen).
Plants terrestrial, at 0-300 m elevation, in forests, from the Depto. Choco.
Also from other areas of western Colombia.

84. Selaginella tuberosa McAlpin & Lellinger, Brenesia 24:409. 1986.

TYPE: Las Alturas de Coton, Pcia. Puntarenas, McAlpin 2505 (US; isotypes
CR, DUKE, NY).

Plants terrestrial, at ca. 1500 m elevation, in moist forests, known only from the
type locality.

This species is peculiar in having large, elongate, annual tubers borne at the end
of underground stems. The tubers form at the end of the wet season, the plants
die down at in the dry season, and new growth appears from the tubers at the
beginning of the following wet season.

85. Selaginella umbrosa Lemaire ex Hieron. in Engl. & Prantl, Nat. Pflanzenfam.
1(4):683, f. 404. 1901.

Sela^nella erythropus var. major Spring, Nouv. Mem. Acad. Roy. Sci. Bruxelles 24:156. 1849.
TYPE: Guatemala, Skinner (K not seen).

Selagjnella lemairei Hieron. Hedwigia 58:287. 1917, nom. illeg. TYPE: A renaming of S. umbrosa
Lemaire ex Hieron., and so based on the type of that name.

TYPE: Not stated; perhaps best considered a renaming of S. erythropus var.
major, and so based on the type of that name.

Plants terrestrial, at 0-700 m elevation, often in open areas, from the Llanura
de Tortuguero, the valley of the Rio General, the Atlantic coastal plain of Panama
to the Canal Zone, Pcia. Colon, and adjacent Pcia. Panama, and around Quibdo.
Also from the Lesser Antilles, Mexico to Honduras, Nicaragua, Venezuela, and
Colombia.

86. Selaginella viticulosa Klotzsch, Linnaea 18:524. 1845.

SYNTYPES: La Guiara, Distr. Fed., Venezuela, Otto 460 (B not seen) and
Moritz 70 (B not seen; isotype BM not seen photo 6512).

Plants terrestrial, at 0-500(1000) m elevation, from near S. Ramon, the
Peninsula de Nicoya, the Peninsula de Osa, and scattered localities in lowland
Panama. Also from Venezuela and Colombia.

FIGS. 80 - 82. Selaginella. FIG. 80. Branch portions of S. silvestris, Biolley 7507. FIG. 81. Branch
portions of S. tanyclada, Archer 1966. FIG. 82. Branch portions of S. tarapotensis, Pittier 5639.

68 SELAGINELLACEAE: 3. SELAGINELLA

SELAGINELUVCEAE: 3. SELAGINELLA 69

87. Selaginella wolfii Sodiro, Anal. Univ. Quito 12(83); 490 (repr. 620). 1895.

TYPE: Cordillera Occidental, Ecuador, 1800 m, Sodiro (P not seen).

Plants terrestrial, at 0-1500 m elevation, from BaMa Solano and the central
Depto. Choco. Also from Ecuador and Peru.

This species is close to S. bombycma and to S. oaxacana.

FIGS. 83 “87. Selaginella. FIG. 83. Branch portions of S. tomentosa, Killip & Garcia 33151. FIG. 84.
Branch portions of S. tuberosa, McAlpin 2505. FIG. 85. Branch portions of S. umbrosa. Hart 11.
FIG. 86. Branch portions of S. viticuiosa. Burger & Stolze 5411. FIG. 87. Branch portions of S.
wolfii, Hau^t 5525.

70 ISOETACEAE: 4. ISOETES

ISOETACEAE

Monotypic; see description of Isoetes.

4. ISOETES L.

Plants aquatic, amphibious, or terrestrial in vernal pools, rooted in mud or
sometimes gravelly soil; roots and leaves borne on 2- or 3-lobed, erect, usually
subterranean, indeterminate, globular, corm-like stems; roots rather fleshy, borne
in longitudinal lines on the outer and basal faces of the corms; leaves rush-like,
erect or spreading, crowded, borne on the apical and outer surfaces of the corms,
triangular or terete, with an unbranched midrib, 0- several longitudinal strands of
thickened supporting cells, and longitudinal air chambers divided by lateral
septae; fertile leaves swollen at the base, bearing an adaxial, embedded
sporangium often partially or entirely covered by a scarious velum and a usually
triangular ligule just distal to the sporangium; sporangia of 2 types, the
megasporangia bearing a few large, spherical, white, often highly ornamented
megaspores, the microsporangia bearing many minute, usually ellipsoid, tan
microspores.

Widespread in temperate and tropical regions; ca. 100-200 species. The genus
is the only extant representative of the Lycopsida subclass Isoetidae, which has a
long fossil record. Tlie peculiar morphology of Isoetes has made it the object of
much study (Karrfalt, 1980; Paolillo, 1963). Some of the species show
physiological adaptations to their hot and sunny habitat (Keeley, 1981). The
taxonomy of this genus is difficult but is made easier through examination of
mature spores with the scanning electron microscope. The tropical American
species of the genus are currently under study by H. P. Fuchs-Eckert (1982) and R.
J. Hickey (1986). According to Hickey (pers. comm.), the upland populations of
southern Central America all differ slightly from one another, as do the lowland
ones, and it is best to treat each of the two groups as a single species, rather than
to name several populations of each as separate species.

FUCHS-ECKERT, H. P. 1982. Zur heutigen Kenntnis von Vorkommen imd Verbreitung der
sudamerikanischen Isoetes- Arten. Proc. K. Nederl. Akad. Wetensch. C, 85:205-260.
HICKEY, R. J. 1986. Isoetes megaspore surface morphology; nomenclature, variation, and
systematic importance. Amer. Fern J. 76:1-16.

KARRFALT, E. E. 1980. A further comparison of Isoetes roots and stigmarian appendages. Canad.
J. Bot. 58:2318-2322.

KEELEY, J. E. 1981. Diurnal acid metabohsm in vernal pool Isoetes (Isoetaceae). Madrono
28:167-171.

PAOLILLO, D. J. 1963. The developmental anatomy of Isoetes. Illinois Biol. Monogr. 31:1-130.
PFEIFFER, N. E. 1922. Monograph of the Isoetaceae. Aim. Missouri Bot. Card.
9:79-232.

1. Leaves 30 - 55 cm long, the ala 2-5 mm wide distal to the leaf base, hyaline or whitish,
sometimes faintly brown-edged, 9 - 12 cm long; hgule deltate, 3-6 mm long; velum covering less than
half of the sporangium; mature megaspores 0.35 - 0.60 mm in diam., tuberculate.

88. /. panamensis

1. Leaves 5-30 cm long, the ala up to 1 mm wide distal to the leaf base, brown-edged or
occasionally pale, 1-5 cm long; hgule deltate to lanceolate, 1.5-4 mm long; velum small to
complete; mature megaspores 0.40 - 0.70 mm in diam., not tuberculate.

89. /. storkii

ISOETACEAE: 4. ISOETES 71

88. Isoetes panamensis Maxon & Morton in Woodson & Seibert, Ann. Missouri
Bot. Gard. 26:272. 1939.

Isoetes pacifica Svenson, Amer. Fern J. 34:123, 1. 10. 1944. TYPE: East of Chanduy, Pda. Guayas,
Ecuador, Svenson 11002 (BKL not seen), synonymized by Stolze (Fieldiana, Bot. N.S., 12:66. 1983).

Isoetes savannarum G6mez, Phytologia 49:339. 1981. TYPE: Near Tanque, La Cruz, Pcia.

Guanacaste, ca. 200 m, Gdmez 7350 (CR; isotype US? not seen).

TYPE: Vicinity of Bejuco, Pcia. Panama, Woodson, Allen & Seibert 1685 (US;
isotype NY).

Plants aquatic, at 0-200 m elevation, in temporary pools in grasslands, from
northwestern Costa Rica and Pcia. Panama. Also from Guatemala, Ecuador, and
Peru.

89. Isoetes storkii Palmer, Amer. Fern J. 22:136. 1932.

Isoetes montana Palmer, Amer. Fern J. 21:134. 1931, non Weber, 1922, nom. illeg. TYPE: Poas
Lake, Pcia. Alajuela, 8500 ft. Stork 2361 (US; isotypes GH, NY).

Isoetes tryoniana Gomez, Revista Biol. Trop. 17:108, f. 8. 1970. TYPE: Large lake on Chirrip6
Grande, Pcia. S. Jose, 3300 m, Gomez PtC-1032 (CR not seen).

TYPE: A renaming of /. montana Palmer, and so based on the type of that
name.

Plants aquatic, at 2500-3700 m elevation, in pools in paramos and montane
bogs, from Volcdn Pods, and the Cordillera de Talamanca to Pcia. Bocas del Toro.

72 EQUISETACEAE: 5. EQUISETUM

EQUISETACEAE

Monotypic; see description of Equisetum.

5. EQUISETUM L.

Plants terrestrial, often in wet places; rhizomes subterranean, blackish, wide-
creeping with wiry roots, branched, bearing erect stems at their nodes; erect stems
cylindrical, hollow, unbranched or with few to many branchlets in a whorl at some
or all nodes; erect stems and branchlets gray-green, hollow between the nodes,
slightly fluted, tough and harsh from silica deposited in the cell walls; leaves
reduced, connate laterally to form nodal sheaths with often scarious and deciduous
distal teeth; sporophylls highly modified (peltate) and aggregated to form strobili
terminal on the branches or branchlets; sporangia ca. 6 per sporophyll, dehiscent
adaxially; spores of 1 size, spherical, greenish, bearing 4 hygroscopic, coiled bands
(elaters).

Widespread in temperate and to some extent tropical regions; 15 species plus
many interspecific hybrids. The genus is the only extant representative of the
ancient class Equisetopsida, which was especially conspicuous in the
Carboniferous. The taxonomy of the genus has been well worked out by Hauke,
but the identification of individual specimens remains difficult because of the
minute characters and the frequent interspecific hybrids.

HAUKE, R. L, 1963. A taxonomic monograph of the genus Equisetum subgenus Hippochaete,
Beih. Nova Hedwigia 8:1-123, 1. 1-22.

. 1966. A systematic study of Equisetum arvense. Nova Hedwigia 13:81-109, 1. 1-9.

. 1978. A taxonomic monograph of Equisetum subgenus Equisetum. Nova Hedwigia

30:385 - 455.

1. Erect stems 1-1.5 mm in diam., usually less than 0.5 m long, irregularly branched with 0-3(4)
branches at a node.

90. E. bogotense

1. Erect stems 3.5-24 mm in diam., usually 2-8 m long, regularly branched with (0)5-30 or more
branches at a node..2.

2(1). Ridges of branches bearing square tubercles; stomata (revealed by a drop of alcohol on the
stem) in bands (2)3 -4(5) rows wide. Stems up to 5 m long, 3.5-24 mm in diam.; strobili decidedly
apiculate.

91. E. giganteum

2(1). Ridges of branches serrate, bearing triangular tubercles or the tubercles irregular; stomata in
single rows or in bands 2(3) rows wide..3.

3(2). Stomata in single rows or in bands 2(3) rows wide; tubercles irregular to triangular. Stems
up to 4.5 m long, 3-22 mm in diam.; strobili acute or slightly apiculate; spores abortive.

93. E. xschaffneri

3(2). Stomata always in single rows; tubercles triangular. Stems up to 8 m long, 6-18 mm in
diam.; strobili blunt or shghtly apiculate.

92. E. myriochaetum

90. Equisetum bogotense H.B.K. Nov. Gen. Sp. PI. 1:42 (fol. 35). 1816.

Equisetum bogotense vai.flagelliforme Kunze, Linnaea 9:4. 1834. TYPE: Antuco, Chile, Feb 1829,
Poeppig (LZ destroyed).

Equisetum bogotense var. nudum Milde, Verh. K. K. Zool.-Bot. Ges. Wien 12:1245. 1862. TYPE:
Six syntypes from Guatemala to Chile cited.

Equisetum bogotense var. polystachum Milde, Verh. K. K. Zool.-Bot. Ges. Wien 13:226. 1863.
TYPE: Caracas, Distr. Fed., Venezuela, Moritz (Hb. F6e not seen).

EQUISETACEAE 5. EQUISETUM 73

Equisetum stipulaceum Vauch. M6m. Soc. Phys. Gen6ve 1(2):377. 1872. TYPE: Peru, Dombey (P
not seen).

Equisetum rinihuense Kunkel, Ber. Schweiz. Bot. Ges. 74:59. 1964. TYPE: Lago Rinihue, Pcia.
Valdivia, Chile, Kunkel 6436 (Hb. Kunkel? not seen; isotype K not seen).

TYPE: Near Sta. Fe de Bogota and Alto del Roble, Distr. Esp., Colombia, 1360
hexap, Humboldt & Bonpland (P-Hb. H.B.K. not seen microfiche S. I. Library).

Plants terrestrial, at 1000 - 2900 m elevation, in pastures, roadside ditches, and
on banks, from the Cordillera Central, the Fila de Cedral, Cerro Tablazo, and the
Cordillera de Talamanca to Pcia. Chiriqui. Also from Venezuela and Colombia to
Argentina and Chile.

91. Equisetum giganteum L. Sp. PL ed, 2, 2:1517. 1763.

Equisetum ramosissimum H.B.K. in Willd. Sp. PI. ed. 4, 5:9. 1810, non Desf., 1799, nom. illeg.
TYPE: Caracas, Distr. Fed., Venezuela, Humboldt & Bonpland (B-Hb. Willd. 19337 not seen
microfiche S. I. Library).

Equisetum caracasanum DC. FI. France 6:244, nota. 1815. TYPE: Caracas, Distr. Fed.,
Venezuela, Vargas 49 in 1829 (P-Hb. DC. not seen).

Equisetum humboldtii Poir. Enc. Meth. Suppl. 4:549. 1816. TYPE: A renaming of E.

ramosissimum H.B.K. in Willd., and so based on the type of that name.

Equisetum pyramidale Goldm. in Meyen, Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19,
Abh. 1:469. 1843. TYPE: Melipilla, Pcia. Santiago, Chile, Meyen (B? not seen).

Equisetum scandens Remy ex Gay, Hist. Ffs. Polit. Chile, Bot. 6:471. 1854. TYPE: Quillota, Pcia.
Valparaiso, Chile Gay (BR? or P? not seen).

Equisetum xylochaetum Mett. Fd. Lechl. 2:34. 1859. TYPE: Near Arica, Depto. Tarapaca, Chile,
Lechler 1556 (B not seen). The nom. nud. Equisetum poeppig^anum A. Braun ex Mett. (Fil. Lechl.
1:27. 1856) is cited in synonymy.

Equisetum lechleri Milde, Verb. K. K. Zool.-Bot. Ges. Wien 11:350. 1861. TYPE: Near Arica,
Pcia. Tarapaca, Chile, Lechler 1556 (B not seen).

Equisetum martii Milde, Verb. K. K. Zool.-Bot. Ges. Wien 12:1258. 1862. SYNTYPES: Salgado,
Est. Minas Gerais, Brazil, Martius in 1818 (M not seen photo 7777); and Caldas, Est. Minas Gerais,
Brazil, 12 Sept 1854, Lindberg (BR not seen).

Equisetum martii var. minus Milde, Verb. K. K. Zool.-Bot. Ges. Wien 12:1261. 1862. TYPE: Peru,
Gaudichaud 33 in 1834 (P-Hb. DC. not seen).

Equisetum brasiliense Milde, Verb. K. K. Zool.-Bot. Ges. Wien 12:1262. TYPE: Brazil, Weddell
921 in 1858 (P-Hb. DC. not seen).

Equisetum elongatum var. scaberrimum Milde, Verb. K. K. Zool.-Bot. Ges. Wien 12:1264. 1862.
TYPE: Valparaiso, Pcia. Valparaiso, ChUt, Jelinek (W not seen).

Equisetum brasiliense f. nudum Milde, Verb. K. K. 2Lool.-Bot. Ges. Wien 13:228. 1863, as '‘nudal’
TYPE: Brazil, Weddell (P-Hb. DC. not seen). Presumably a superfluous name because based on the
type of E. brasiliense", the correct name is f. brasiliense.

Equisetum brasiliense f. ramosum Milde, Verb. K. K. Zool.-Bot. Ges. Wien 13:228. 1863, as
‘"ramosa.” TYPE: Sto. Domingo [Hispaniola], de Tussac (Hb. F6e not seen).

Equisetum giganteum var. chilense Milde, Verb. K. K. Zool.-Bot. Ges. Wien 13:229. 1863, as
''chilensisl" TYPE: Chile, Gay (Hb. Fee not seen).

Equisetum elongatum var. ajfine Milde, Ann. Mus. Bot. Lugduno-Batavum 1:68. 1864. TYPE:
Santiago de Chile, Pcia. Santiago, Chile, Philippi PI. Chil. ed. Hohenacker 70 in 1859 (L? not seen).

Equisetum elongatum var. dolosum Milde, Ann. Mus. Bot. Lugduno-Batavum 1:245. 1864. TYPE:
Copiap6, Pcia. Atacama, Chile, Phliippi PI. Chil. ed. Hohenacker 701 (B not seen).

Equisetum bolivianum Gand. Bull. Soc. Bot. France 66:304. 1920. TYPE: Yungas, Bolivia, Bang
in 1901 (LY? not seen; isotype US).

Equisetum philippii Gand. Bull. Soc. Bot. France 66:304. 1920, as ''phillipi.” TYPE: Chile,
Philippi (LY? not seen).

74 EQUISETACEAE: 5. EQUISETUM

EQUISETACEAE 5. EQUISETUM 75

Equisetum giganteum van. digitaliferum Pastore, Physis (Buenos Aires) 15:249. 1939. TYPE:
Estancia Grande, S. Luis, Argentina, Pastore 94 (SI not seen).

TYPE: Plate 125, f. 2 of Plumier’s “Plantarum Americanum Fasciculus
Quintus,” which is presumably based on a specimen collected by Plumier on
Hispaniola. The plate is clearly drawn from a specimen of E. giganteum, and the
reference on p. 115 to E. fluviatile L. is a misidentification on Plumier’s part.

Plants terrestrial, at 600-1700 m elevation, on river banks and in wet places,
from the Rio Reventazdn valley and the Cordillera de Talamanca to Pcia.
Chiriqui. Also from Cuba, Jamaica, Hispaniola, Mexico to El Salvador,
Nicaragua, and South America except for the Guianas.

92. Equisetum myriochaetum Schlechtend. & Cham. Linnaea 5:623. 1830.
Equisetum myriochaetum var. densum Milde, Verb. K. K. Zool.-Bot. Ges. Wien 11:355. 1861.

TYPE: Mexico, Schajfner (B not seen).

Equisetum myriochaetum var. laxum Milde, Verb. K. K. Zool.-Bot. Ges. Wien 11:355. 1861.

TYPE: Mexico, Schaffner (B not seen).

Equisetum mexicanum Milde, Verb. K. K. Zool.-Bot. Ges. Wien 12:1256. 1862. TYPE: Mexico,
Karwinski (M not seen).

Equisetum elongatum var. mueiieri Milde, Ann. Mus. Lugduno-Batavum 1:244. 1864. TYPE: Rio
Blanco, Eugenio, Mexico, Aug 1853, Mueller (L? not seen).

Equisetum myriochaetum f. sprucei Milde, Nov. Actorum Acad. Caes. Leop.-Carol. German. Nat.
Cur. 32(2) :499. 1867. TYPE: Banos, Rio Pastaza, Pcia. Oriente, Ecuador Spruce 5370 (B not seen).

TYPE: Misantla, Edo. Veracruz, Mexico, Schiede & Deppe 833 (HAL? not seen;
isotypes B not seen, LZ destroyed).

Plants terrestrial, at 200-1300 m elevation, on river banks and in wet places,
from the northern and western slopes of the Cordillera de Talamanca. Also from
Mexico to Honduras, Venezuela, Colombia, Ecuador, and Peru.

93. Equisetum xschaffneri Milde, Verb. K. K. Zool.-Bot. Ges. Wien 11:345. 1861.

Equisetum elongatum var. annuliferum Milde, Ann. Mus. Bot. Lugduno-Batavum 1:244. 1864.
TYPE: Sierra de la Cruz, Mexico, Aug 1853, Mueller (L? not seen).

TYPE: Near Orizaba, Edo. Veracruz, Mexico, Schajfner 315 (B not seen).

Plants terrestrial, at 1100-1400 m elevation, in marshes and pastures, from the
eastern end of the Meseta Central, around Cartago, and near Sta. Maria de Dota
(Pcia. S. Jose). Also from Mexico to El Salvador, Venezuela, Colombia, Ecuador,
and Peru.

This species is the hybrid between E. giganteum and myriochaetum.

FIGS. 88 - 93. Isoetes and Equisetum. FIG. 88. Leaf of I. panamensis, Woodson et al. 1685. FIG. 89.
Leaf base of I. storkii, Latbrop 5560. FIG. 90. Main axis with brancblets of E. bogotense, Burger &
Liesner 6792. FIG. 91. Portion of ultimate branch of E. giganteum, Mickel 3380. FIG. 92. Portion of
ultimate branch ofE. myriochaetum, Hauke & L6pez 336(2b). FIG. 93. Portion of ultimate branch
of E. xschaffneri, Mickel 2412.

76 OPHIOGLOSSACEAE: 6. BOTRYCHIUM

OPHIOGLOSSACEAE

Rhizomes subterranean, short and erect or rarely globular, glabrous, bearing
fleshy roots lacking root hairs and 1-few fronds; fronds small, brittle, not
circinate, partially dimorphic; stipes fleshy; laminae papyraceous to subcoriaceous,
the sterile portion simple or forked to 3-pinnate, the usually erect fertile portion
greenish or yellowish, unbranched and bearing 2 rows of sunken sporangia or
branched several to many times and bearing 2 rows of short-stalked sporangia;
sporangia globular, lacking an annulus, dehiscing transversely; spores trilete,
numerous (more than 1,000 per sporangium).

1. Sterile portion of the laminae 2-3-pinnate, more or less deltate, with free veins; fertile portion
of the laminae pinnately divided.

6. Botrychium

1. Sterile portion of the laminae simple or furcate, not deltate, with anastomosing veins; fertile
portion(s) of the laminae simple.

7. Ophioglossum

6. BOTRYCHIUM Swartz

Plants terrestrial, in meadows and pastures or occasionally in forests; rhizomes
subterranean, erect, short, bearing fleshy roots at the base and usually a single
frond near the apex; buds of new fronds protected by the sheathing base of the old
frond, the laminae when young folded, not circinate; fronds glabrous, partially
dimorphic; sterile portion of the laminae expanded, 2-3-pmnate, spreading,
yellow-green or bright green, the lamina subternate, the venation free; fertile
portion of the laminae contracted, 1-3-pinnate, erect, yellow, bearing short-
stalked, globular sporangia.

Widespread in temperate regions, with some tropical representatives; ca. 35
species. The species of Botrychium are subtle and their characters tend to be
variable, all of which causes considerable difficulty in identifying them. The genus
is being monographed by W. H. Wagner, Jr., who has combined herbarium and
field studies to arrive at far better species concepts than were formerly known.

CLAUSEN, R. T. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club
19(2):1-177.

WAGNER, W. H., Jr. and F. S. WAGNER. 1983. Genus communities as a systematic tool in the
study of New World Botrychium (Ophioglossaceae). Taxon 32:51-63.

1. Fertile portion of the laminae inserted several cm proximal to the base of the sterile portion;
sterile portion papyraceous to subcoriaceous..3.

1. Fertile portion of the laminae inserted at the base of the sterile portion; sterile portion
membranaceous to papyraceous..2.

2(1). Fertile portion of the laminae about as long as the sterile; ultimate segments acute, often with
a few shallow incisions. Stipes 6-38 cm long; sterile laminae deltate, 6-28 cm long, 12-38 cm wide,
3-pinnate-pinnatifid, the ultimate segments oblong.

96. B. viipnianum var. mexicanum

2(1). Fertile portion of the laminae usually greatly exceeding the sterile; ultimate segments round,
usually lacking incisions. Stipes 25 - 30 cm long; sterile laminae broadly deltate, 10-13 cm long,
12-14 cm wide, 2-pinnate-pinnatifid to 3-pinnate, the ultimate segments elliptic to obovate.

97. B. virginianum var. virginianum

3(2). Pinnae obtuse or round at the apex; basal basiscopic pinnule of the lateral pinnae 1-2 times
longer than the basal acroscopic pinnule; segment margins irregularly serrate-lacerate.

95. B. underwoodianum

OPHIOGLOSSACEAE: 6. BOTRYCHIUM 77

3(2). Pinnae acute at the apex; basal basiscopic pinnule of the basal pinnae 2-3 times longer than
the basal acroscopic pinnule; segment margins entire to evenly crenate.

94. B. decompositum

94. Botrychium decompositum Martens & Galeotti, Nouv. M^m. Acad. Roy. Sci.
Bruxelles 15:51, 1. 1. 1842.

TYPE: Orizaba, Edo. Veracruz, Mexico, 5000-6000 ft, Galeotti 6452 (BR not
seen photo 5061).

Plants terrestrial, at 1200-2200 m elevation, from Volcan Barba and Las
Concavas (Pcia. Cartago). Also from Louisiana, Mexico, Guatemala, and
Honduras.

95. Botrychium underwoodianum Maxon, Bull. Torrey Bot. Club 32:222, t. 6.
1905.

Botrychium tematum var. daedaleum Christ in Pitt. Prim. FI. Costaric. 3(1):46. 1901. TYPE:
Cerros de Velirla, Copey, Pcia. S. Jos6, 2600 - 2700 m, Tonduz 11864 (BR or P not seen; isotype CR
not seen photo US).

TYPE: Jamaica, Ze/ima/t in 1874-1879 (NY not seen; isotype US).

Plants terrestrial, at 2200-3500 m elevation, in pastures, from La Palma de S.
Ramon, the Cordillera Central, the Fila de Cedral, and around Sta. Maria de Dota
(Pcia. S. Jose). Also from Jamaica, Hispaniola, Guatemala, Honduras, Venezuela,
and Colombia.

96. Botrychium virginianum var. mexicanum Hook. & Grev. Bot. Misc. 3:223.
1833.

Osmunda cicutaria Sav. ex Poir. in Lam. Encyc. Meth. 4:650. 1798. TYPE: Based on plate 159 of
Plumier’s “Traitd...”, which is based on specimens collected by Plumier on Hispaniola.

Botrychium brachystachys Kunze, Linnaea 18:305. 1844. TYPE: Mexico, Leibold 35 (LZ

destroyed).

Botrychium dichronum Underw. Bull. Torrey Bot. Club 30:45. 1903. TYPE: Morce’s Gap,
Jamaica, 1500 m, Clute 96 (NY not seen; isotype US).

Botrychium virginianum var. meridionale Butters, Rhodora 19:213. 1917. TYPE: Edo. Chiapas,
Mexico, Ghiesbreght 252 (GH not seen).

SYNTYPES: Jalapa, Edo. Veracruz, Mexico, Chamisso (K not seen); and Rigla,
Mexico, Veitch (K not seen).

Plants terrestrial, at 1000-2700 m elevation, in forests, from the Cordillera
Central, Cerro Piedra Blanca (Pcia. S. Jose), Cerro Carpintera, the northern
portion of the Cordillera de Talamanca, and around Boquete. Also from Cuba,
Jamaica, Hispaniola, Mexico, Guatemala, El Salvador, Colombia to Bolivia, and
Brazil.

According to W. H. Wagner, Jr. (pers. comm.), intermediates with var.
virginianum are known.

97. Botrychium virginianum (L.) Swartz, J. Bot. (Schrader) 1800(2):111. 1801,
var. virginianum.

Osmunda virginiana L. Sp. PI. 2:1064. 1753. TYPE: North America, Kalm (LINN 1244.3 not seen
microfiche S. I. Library).

Botrychium gracile Pursh, FI. Amer. Sept. 2:656. 1814. TYPE: Virginia, Pursh (PH? not seen).
Botrychium virginianum var. intermedium Butters, Rhodora 19:210. 1917. TYPE: Canton, New
York, Phelps 47 (GH not seen), synonymized by Clausen (Mem. Torrey Bot. Club. 19(2) :98. 1938).

78 OPHIOGLOSSACEAE: 6. BOTRYCHIUM

OPHIOGLOSSACEAE 7. OPHIOGLOSSUM 79

Botrychium virg^nianum var. occidentale Butters, Rhodora 19:213. 1917. TYPE: South end of
Lake Pend d’Oreille, Idaho, Sandberg et al 762 (GH not seen; isotype US).

Plants terrestrial, at 1600-2400 m elevation, in forests and open areas, from
south of Sta. Maria de Dota (Pcia. S. Jose), and Boquete and the Volcan de
ChiriquL Also from Canada, the United States, Mexico, and Guatemala.

7. OPHIOGLOSSUM L.

Plants terrestrial, in meadows and pastures (subg. Ophioglossum) or epiphytic
on tree trunks in wet forests (subg. Cheiroglossa); rhizomes subterranean, erect,
short, rarely globose, bearing fleshy roots at the base and 1-few fronds near the
apex; fronds glabrous, not circinate, partially dimorphic; sterile portion of the
laminae expanded, simple (furcate in subg. Cheiroglossa), pale to dark green, the
venation irregularly areolate, sometimes with finer veins dividing the major
areolae into yet smaller ones; fertile portion of the laminae (1 in subg.
Ophioglossum, several in subg. Cheiroglossa) contracted, simple, erect (in subg.
Ophioglossum) or pendulous (in subg. Cheiroglossa), bearing 2 rows of globular
sporangia partially embedded in the tissue.

Widespread in temperate and tropical regions, but nowhere common; ca. 25
species. These reduced plants have few characters, and they are variable. The
sterile, expanded portion of the laminae is peculiar in lacking a midrib. Species
delimitation in Ophioglossum may be difficult; size is not a reliable character, and
venation is sometimes influenced by size.

CLAUSEN, R. T. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club
19(2):1~177.

1. Plants 25 - 60 cm long; sterile portion of the laminae palmately lobed, usually with several
elongate, short-stalked fertile portions along the lateral edges of the sterile portion and the stipe
apex; plants epiphytic (subg. Cheiroglossa).

101. O. palmatum

1. Plants less than 15(30) cm long; sterile portion of the laminae entire and unlobed, with 1
elongate, long-stalked fertile portion inserted on the common stalk at or just proximal to the sterile
portion; plants terrestrial (subg. Ophioglossum)..!.

2(1). Rhizomes erratically spherical, ca. 3-5 mm in diam.

98. O. crotalophoroides

2(1). Rhizomes cylindrical or ovoid, usually less than 3 mm in diam. ..3.

3(2). Primary areolae of the sterile portion of the laminae with included secondary areolae.
Laminae 4-6 cm long, 1-2 cm wide.

99. O. ellipticum

3(2). Primary areolae of the sterile portion of the laminae with occasional included veinlets, but
lacking secondary areolae..4.

4(3). Sterile portion of the laminae 2-6 cm long, (2)2.5 -4.5 cm wide; rhizomes cylindrical.

102. O. reticulatum

4(3). Sterile portion of the laminae 0.4- 1.5 cm long, 0.3 -0.8 cm wide; rhizomes ovoid.

100. O. nudicaule

FIGS. 94-102. Botrychium and Ophioglossum. FIG. 94. Median pinna of B. decompositum, Wagner
81001. FIG. 95. Median pinna of B. underwoodianum, Tonduz 11864. FIG. 96. Median pinna of B.
virginianum var. mexicanum, Standley 34153. FIG. 98. Plant of O. crotalophoroides, Skutch 604,
Guatemala. FIG. 99. Plant of O. ellipticum, Williams & Molina 14153, Honduras. FIG. 100. Plant
of O. nudicaule, Pringle 7086, Mexico. FIG. 101. Plant of O. palmatum, Skutch 3427. FIG. 102.
Plant of O. reticulatum, Tonduz 8788.

80 OPHIOGLOSSACEAE: 7. OPHIOGLOSSUM

98. Ophioglossum crotalophoridcs Walter, FI. Carol. 256. 1788.

Ophioglossum bulbosum Michx. FI. Bor.-Amer. 2:276. 1803. TYPE: “In sabulosis Carolinae
inferioris,” Michaux (P not seen Morton photo 3373).

Ophioglossum stipatum CoUa, Mem. Reale Acad. Sci. Torino 39:52, t. 75, f. 1. 1836. TYPE
LOCALITY: Chde.

TYPE: South Carolina, Walter (not seen).

Plants terrestrial, often among mosses, at 3300-3400 m elevation, in exposed
areas and pastures, in the Flora area known only from the Valle de los Conejos,
near El Canon, Pda. Cartago {Gomez & Burger, Gomez 3411, both CR). Also
from the southern United States, Hispaniola, Mexico to Honduras, Venezuela,
Colombia, Peru, Bolivia, Uruguay, Argentina, and Chile.

This species is unique among New World Ophioglossums for its globular
rhizomes, which were studied in detail by Mesler (Amer. Fern J. 63:28-33. 1973).

99. Ophioglossum ellipticum Hook. & Grev. Ic. Fil. l:t. 40A. 1827.

Ophioglossum vulgatum var. surinamense Luerss. J. Mus. Godeffroy 3:246, f. 119, 120, 131, 132.
1875. TYTE: Surinam, Weigelt (B? not seen).

TYPE: Demarera, Guyana, Parker (K not seen).

Plants terrestrial, at 0-900(2100) m elevation, in savannas, meadows, and open
ground, from the Cordillera Central, the Cordillera de Talamanca near Copey
(Pda. S. Jose), the basin of the Rio General, and various localities in Pcia.
Panama. Also from Mexico to Honduras and the Guianas to Brazil.

According to Wagner, Allen, and Landry (Castanea 49:99-110. 1984), this
species may be a large form of O. nudicaule, at least as to material from the
southeastern United States. Ophioglossum usterianum Christ in Usteri (FI. Sao
Paulo 137. 1911) and O. valdivianum Phil. (Linnaea 33:306. 1865) may be

synonyms of this species; I have seen neither types nor other specimens of either.

100. Ophioglossum nudicaule L. fil. Suppl. PI. Syst. Veg., ed 13. 443. 1782.

Ophioglossum ypanemense Mart. Icon. PI. Crypt. Bras. 39, 130, t. 73, f. 1. 1834. TYPE: Ypanema,
Est. S. Paulo, Brazil, 1700 ft, Martius in 1818 (BR not seen; isotype M not seen photo 7769).

Ophioglossum macrorhizum Kunze, Farrnkrauter 1:57, t. 29, f. 1. 1840. TYTE: French Guiana,
Leprieur (LZ destroyed?).

Ophioglossum melipillense Remy in Gay, FI. FIs. Polit. Chile, Bot. 6:542. 1853. TYPE

LOCALITY: Melipilla, Chile.

Ophioglossum spruceanum Fee, Crypt. Vase. Bresil 1:218, t. 52, f. 3. 1869. TYPE: Near S. Gabriel
de Cachoeira, Rio Negro, Est. Amazonas, Brazil, Spruce 2041 (RB not seen; isotypes BM not seen
photo 7770, US).

Ophioglossum tenemm Mett. ex Prantl, Ber. Deutsch. Bot. Ges. 1:352. 1883. TYPE LOCALITY:
Georgia, United States.

Ophioglossum nudicaule var. minus Clausen, Mem. Torrey Bot. Club 19(2):148, f. 29. 1938.
TYPE: Vicinity of Ft. Myers, Lee County, Florida, Standley 354 (US; isotypes GH, NY).

Ophioglossum nudicaule var. vulcanicum Clausen, Mem. Torrey Bot. Club 19(2): 150, f. 31, 32.
1938. TYPE: Llanos del Volc^, Pcia. Chiriquf, Panama, Martin 2798 (US).

Ophioglossum dendroneuron E. St. John in Small, Ferns S.E. States 364, fig. 1938. TYPE: Crystal
River, Lecanto, Citrus Co., Horida, E. St. John 568 (NY not seen).

TYPE: Cape of Good Hope, South Africa, Thunherg (UPS not seen microfiche
S. I. Library).

Plants terrestrial, at 0-1600 m elevation, from Tarbaca (Pcia. S. Jose), Llanos
del Volcan (Pcia. Chiriquf), near Gatun (Canal Zone), and east of Pacora and S.

OPHIOGLOSSACEAE 7. OPHIOGLOSSUM 81

Jose Island (both Pda. Panama). Also from the southeastern United States, Cuba,
Hispaniola, Mexico, Guatemala, Honduras, Nicaragua, Venezuela, Colombia,
Bolivia, Guyana, French Guiana, Brazil, and Argentina.

101. OpMoglossum palmatum L. Sp. PL 2:1063. 1753.

TYPE: Plate 163 of Plumier’s “Traite...”, which is based on a specimen
collected by Plumier on Hispaniola.

Plants epiphytic, at 1400-2200 m elevation, in forests, from the vicinity of S.
Ramon, the Cordillera Central, Orosi (Pda. Cartago), and Cerro Tacarcuna. Also
from Florida, the West Indies, Mexico, Honduras, Nicaragua, Venezuela,
Colombia to Bolivia and Brazil, Guyana, and Surinam.

102. Ophioglossum reticulatum L. Sp. PL 2:1063. 1753.

Ophioglossum pemvianum K. Presl, Suppl. Tent. Pterid. 52 (postpr. 313). 1845. SYNTYPES:
Pampayaco, Depto. Hu^uco, Peru, Poeppig 1150 (PRC not seen); and Sion Mission, Depto. S.
Martin, Peru, Poeppig in 1830 (PRC not seen).

TYPE: Plate 10, £ 4, of Petiver’s “Pterigraphia Americana,” which is a
transposed re-drawing of a portion of t. 164 of Plumief s “Traite...”, which is in
turn based on a specimen collected by Plumier on Hispaniola.

Plants terrestrial, at 700-2100 m elevation, in pastures, on roadbanks, and on
riverbanks, from the Cordillera de Tilaran, the Cordillera Central, Tarbaca (Pda.
S. Jose), Cerro Tablazo, and the Cordillera de Talamanca to Pda. Chiriqui. Also
from the Antilles, Mexico to Honduras, Nicaragua, Venezuela to Bolivia, Brazil,
the Guianas, and Argentina.

82 MARATTIACEAE: 8. MARATTIA

MARATTIACEAE

Rhizomes fleshy, creeping or erect and often massive, bearing deciduous or
persistent scales; fronds monomorphic or slightly dimorphic in some species,
brittle; stipes articulate to the rhizome, enlarged at the base and these bearing ala-
like outgrowths; laminae simple to 3(4)-pinnate, sometimes sparsely scaly or hairy,
especially the axes fleshy and swollen at the point of insertion of the opposite
pinnae; rachis (and costae if present) commonly alate; synangia tan to brown,
borne on the abaxial surface of the laminae, the sporangia fused in 2 rows, lacking
an annulus, dehiscent by a pore or slit; spores monolete or trilete, numerous
(more than 1,000 per sporangium).

1. Fronds usually more than 1 m long, 2-3(4)-pinnate, monomorphic; synangia short, not covering
the abaxial surface of the segment, dehiscent by shts.

8. Marattia

1. Fronds usually less than 1 m long, simple to pinnate, shghtly dimorphic; synangia long, nearly
covering the abaxial surface of the frond or pinna, dehiscent by pores.

9. Danaea

8. MARATTIA Swartz

Plants terrestrial, usually in forests, often in very dense shade; rhizomes erect,
short, massive, scaly, the scales thin, tan; fronds large; stipes fleshy, stipulate,
stramineous to brown at the base, scaly at the base; laminae 2-3(4)-pmnate,
especially the axes fleshy; pinnae alternate, the costae exalate, sparsely scaly,
slightly inflated and blackish at the base; costules alate, the alae narrowly
oblanceolate between the pinnules or segments; ultimate segments often oblong
or ovate, usually glabrous, with free, unbranched or forked veins; sporangia in 2
rows, fused into sessile or short-stalked, subterminal, long-oval, brownish synangia
dehiscent by a longitudinal slit.

Widespread but infrequent in the tropics, especially in the Old World; ca. 60
species.

UNDERWOOD, L. M. 1909. Marattiaceae. North Amer. FI. 16(l):15-23.

1. Costules not spinulose on the adaxial surface; synangia sessile or nearly so; brown hairs and
narrow scales present on the abaxial surface of the veins..3.

1. Costules sparingly spinulose on the adaxial surface; synangia distinctly stalked; brown hairs and
narrow scales absent from the abaxial surface of the veins..2.

2(1). Sterile ultimate (tertiary) segments crenate to lobed; lobes of pinnatifid pinnules 0.2 -0.8 cm
long.

105. M. laevis

2(1). Sterile ultimate (tertiary) segments entire to slightly crenate; lobes of pinnatifid pinnules
0.4- 1 cm long.

106. M. pittieri

3(1). Laminae 2-pinnate, the acute-acuminate pinnae 6-30 cm long, 1.5--2 cm wide.

105a. M. laxa

3(1). Laminae predominantly 3-pinnate (smaller pinnae 2-pinnate), the pinnae or pinnules 1.2-6
cm long, 0.6- 1.5(2) cm wide..4.

4(3). Ultimate (tertiary) segments round or subacute at the apex, 8 - 20(25) mm long, 4-10 mm
wide. Laminae predominantly 3-pinnate.

103. M. excavata

MARATTIACEAE; 8. MARATTIA 83

103. Marattia excavata Underw. North Amer. FI. 16(1) :22. 1909.

TYPE: Coliblanco, Pda. Cartago, ca. 2000 m, Maxon 272 (NY; isotype US).

Plants terrestrial, at 1200-1900(2700) m elevation, in wet forests, from the
Cordillera de Tilardn, the Cordillera Central, the Cordillera de Talamanca to
Pda. Chiriqui, and the Fila Costeha near S. Vito. Also from Mexico to Nicaragua.

This species and M. interposita are not markedly distinct anywhere in their
largely coextensive ranges. Detailed field and laboratory study may show that this
species should be no more than a variety of M. interposita.

104. Marattia interposita Christ, Bull. Herb. Boissier II, 6:285. 1906.

Marattia chiricana Maxon, Contr. U. S. Natl. Herb. 17:421. 1914. TYPE: Cuesta de Las Palmas, S
slope of Cerro de la Horqueta, Pcia. Chiriqui, 1700 - 2100 m, Maxon 5525 (US; isotypes GH, US).

TYPE: Navarro, Pcia. Cartago, Werckle (P not seen fragm NY; isotype US).

Plants terrestrial, at (1100)1400-2000 m elevation, in dense, damp to wet
forests, from the Cordillera de Tilaran, the Cordillera Central, Cerro Carpintera,
and Cerro de la Horqueta (Pcia. Chiriqui). Also from Guatemala to Nicaragua.

For a comment on this species, see M. excavata.

105. Marattia laevis J. E. Smith, PL Icon. Ined. 2:t. 47. 1790.

Marattia kaulfussii J. Smith in Hook. Gen. Fil. ad t. 26. 1839. TYPE: Possibly Brazil, collector
unknown (presumably LZ destroyed), according to Kaulfuss (Enum. Fil. 32. 1824), or even Brazil, ^4.

Cunningham in 1816 (BM not seen photo 7750).

TYPE: Hispaniola, Thierry-Th. 90 (LINN not seen), examined by J. M. Camus
(in litt. ad R. G. Stolze 27 June 1985).

Plants terrestrial, at 1500-2000 m elevation, in wet forests, in the Flora area
known only from the flanks of Volcan Barba and south of Tapanti. Also from
Cuba, Hispaniola, Puerto Rico, Venezuela to Bolivia, and Brazil.

105a. Marattia laxa Kunze, Linnaea 18:306. 1845.

TYPE: “In humidis region, temper.,” Mexico, Leibold 62 (LZ destroyed; isotype
Hb. Roemer not seen).

Plants terrestrial, at ca. 1500 m elevation, in the Flora area known only from
near Monteverde, Pcia. Puntarenas {Burger & Antonio 10848, F fragm US; Michel
3582, US). Also from Mexico.

106. Marattia pittieri Maxon, Contr. U. S. Natl. Herb. 17:421. 1914.

TYPE: Holcomb’s trail above Boquete, Pcia. Chiriqui, 1625 m, Maxon 5704 (US
fragm NY; isotype US).

Plants terrestrial, at 1600-1700 m elevation, in wet forests, known only from
the type collection.

This species probably will prove to be synonymous with M. laevis, from whch it
differs by the characters given in the key. The other differentiating characters
listed by Maxon also characterize large specimens of M. laevis that have been
collected in recent years in Colombia.

84 MARATTIACEAE: 8. MARATTIA

MARATTIACEAE: 9. DANAEA 85

9. DANAEA J. E. Smith

Plants terrestrial, in the deepest shade of dense forests; rhizomes short-creeping
or ascending, bearing thick, fleshy roots; fronds small to medium-sized,
subdimorphic, the fertile ones larger but with the pinnae somewhat contracted;
stipes fleshy, stipulate, nodose, stramineous to pale brown, often scaly, the scales
small, thin, dark brown; laminae pinnate or rarely simple, firm, rather fleshy;
rachises similar to the stipes; lateral pinnae mostly elliptic-oblong, opposite,
inserted on the rachises at the swollen nodes, the veins paired or forked near the
costae, closely spaced, bearing nearly throughout elongate, brownish synangia
dehiscent by 2 rows of pores.

Solely tropical America; ca. 30 species.

UNDERWOOD, L. M. 1909. Marattiaceae. North Amer. H. 16(1):15- 23.

1. Sterile fronds simple or with 1 pair of minute basal lobes or pinnae. Sterile fronds with stipes
8 “ 10 cm long, laminae ca. 14 cm long, 3.5 cm wide, cuneate at the base, acuminate and short-serrate
at the apex; fertile fronds with stipes 15 cm long, laminae ca. 15 cm long, 0.9 cm wide.

107. D. carillensis

1. Sterile fronds pinnate, with at least several pairs of lateral pinnae..2.

2(1). Sterile pinnae less than 2.5 cm wide..4.

2(1). Sterile pinnae (2.5)3 cm or more wide..3.

3(2). Sterile pinnae 7-11(15), elliptic, mostly obtuse with an acuminate apex; stipes with 1-5
nodes. Sterile laminae broadly lanceolate, the pinnae up to 15 cm long, 4(5) cm wide, linear- elliptic
to oblanceolate, acute to almost round at the base, acuminate-cuspidate at the apex.

110. D. elliptica

3(2). Sterile pinnae 15 or more, nearly parallel-sided, mostly acute with a caudate apex; stipes
usually without nodes. Sterile laminae lanceate to oblong, the pinnae up to 27(32) cm long, 4.5(5)
cm wide, cuneate to almost round at the base, acuminate-cuspidate at the apex.

112. D. nodosa

4(2). Terminal pinna present, the laminae odd-pinnate; laminae lacking an apical bud..6.

4(2). Terminal pinna absent or deciduous, the laminae even-pinnate; laminae with an apical bud..5.

5(4). Laminae crisped, the margins erose. Sterile fronds with stipes 4-9 cm long with usually 1
node, the laminae ovate-lanceolate, 10-21 cm long, 4-9 cm wide.

108. Z>. crispa

5(4). Laminae not crisped, the margins entire to serrate. Sterile fronds with stipes 4-15 cm long
with 1 or 2 nodes, the laminae linear or linear-lanceolate, 10-30 cm long, 4- 10(15) cm wide.

115. D. wendlandii

6(4). Terminal pinna 2-3 times longer than the longest lateral pinnae. Sterile fronds with stipes 9
cm long with 2 or 3 nodes, the laminae lanceolate, 20 cm long, 12 cm wide.

113. D. plicata

6(4). Terminal pinnae shorter than the longest lateral pinnae..7.

7(6). Largest sterile pinnae 4-5.5 cm long, 0.8- 1.2 cm wide, parallel-sided, distinctly falcate at the
apex. Sterile fronds with stipes 15-20 cm long with 3 or 4 nodes, the laminae linear, 25 - 30 cm long,
5.5-8 cm wide.

114. D. semilata

7(6). Largest sterile pinnae (4)7-18 cm long, 1.2 -2.5 cm wide, only the larger ones parallel-sided,
not or slightly falcate..8.

FIGS. 103-106. Marattia. FIG. 103. Tertiary segments of M. excavata, Valerio 119. FIG. 104.
Tertiary segments of M. interposita, Killip 5145. FIG. 105. Tertiary segments of M. laevis, Pittier
10031, Venezuela. FIG. 106. Tertiary segments of M. pittieri, Maxon 5704.

86 MARATTIACEAE: 9. DANAEA

109 D. cuspidata

MARATTIACEAE: 9. DANAEA 87

8(7). Sterile pinnae inequilateral and obtuse to round at the base, the veins much darker than the
lamina tissue, usually paired at origin. Sterile fronds with stipes 11-45 cm long, with 0-3 nodes;
sterile laminae linear-lanceate, not much tapered at the base, 15-40(55) cm long, (7)10-24 cm wide.

109. D. cuspidata

8(7). Sterile pinnae inequilateral and acute to rarely round at the base, the veins not or only
slightly darker than the lamina tissue, usually not paired at origin. Sterile fronds with stipes 15-31
cm long, with 1 or 2 nodes; sterile laminae usually linear-lanceolate, tapered at the base, 15 - 60 cm
long, (5)10-30 cm wide.

111. D. moritziana

107. Danaea carillensis Christ, Bull. Soc. Bot. Geneve II, 1:234. 1909.

TYPE: Carrillo, Pda. S. Jose, 400 m, Werckle 365 (P not seen).

Plants terrestrial, at 400-1400 m elevation, in wet forests, from 18 km north of
S. Ramon and Volc^n Cacho Negro (both Pda. Alajuela), Virgen del Socorro
(Pda. Heredia), and Carrillo (Pda. S. Jose).

108. Danaea crispa Endres in Reichenb. f. Bot. Zeitung (Berlin) 30:489. 1872.
TYPE: Costa Rica, Endres (B? not seen).

Plants terrestrial, at 300-1200 m elevation, mostly in very dark forests, from
near S. Ramon, the Cordillera Central, the Cordillera de Talamanca (Pda.
Chiriqui), northwest of Sta Fe, and El Cupe (Pda. Code).

109. Danaea cuspidata Liebm. Kongel. Danske Vidensk. Selsk. Skr.,
Naturvidensk. Afd. V, 1:307 (repr. 155). 1849.

Danaea pterorachis Christ, Bull. Soc. Bot. Gendve 11, 1:235. 1909. TYPE: Costa Rica, Werckle in
1903 (P not seen).

Danaea muenchii Christ in Rosenst. Repert. Spec. Nov. Regni Veg. 22:23. 1925. TYPE: S. Pablo,
Edo. Chiapas, Mexico, Muench 159 (probably P not seen; isotypes DS not seen, US).

TYPE: Chinantla, between Lobani and Petlapa, Edo. Oaxaca, Mexico, 2500 ft,
Liehmann (C not seen).

Plants terrestrial, at (600)1000-2300 m elevation, in forests usually near
streams, from the Cordillera Central, Cerro Tablazo, the Cordillera de Talamanca
to Pda. Chiriqui, the Fila Costena near S. Vito, the Peninsula de Osa, the valley of
the Rio Dos Bocas (Pda. Veraguas), Cerro Pilon (Pda. Code), near the Altos de
Pacora (Pda. Panama), and above Cana. Also from Mexico and Guatemala.

This species may prove to be conspecific with D. moritziana, which is an earlier
name. Both species have the minute scales on the abaxial lamina surface noted by
Stolze (Fieldiana, Bot. 39:16. 1976).

110. Danaea elliptica J. E. Smith in Rees, Cycl. 11: Danaea no. 2. 1808.

Danaea geniculata Raddi, Opusc. Sci. 3:281. 1819. TYPE: Mandiocca, Est. Rio de Janeiro, Brazil,
Raddi (FI not seen).

Danaea media Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:306 (repr.
154). 1849. TYPE: Near Hacienda de Jovo, Edo. Veracruz, Mexico, Liehmann (C not seen photos
5578, 5625, 5625a).

FIGS. 107-111. Danaea. FIG. 107. Plant of D. carillensis, Brade & Brade 303. FIG. 108. Frond of
D. crispa, Brade & Brade 328. FIG. 109. Sterile median pinna of D. cuspidata, Scamman 6992. FIG.
110. Frond of D. elliptica, Stern et al. 681. FIG. 111. Sterile median pinna of D. moritziana, Lellinger
& de la Sota 247.

88 MARATTIACEAE: 9. DANAEA

MARATTIACEAE: 9. DANAEA 89

Danaea oligosora Fourn. ex Baker, Ann. Bot. (London) 5:499. 1891. TYPE: Guadeloupe,

Fournier (P? not seen).

Danaea polymorpha Leprieur ex Baker, Ann. Bot. (London) 5:499. 1891. SYNTYPES:

Guadeloupe, Mazi (P? not seen); and Grenada, Sherring (K not seen).

Danaea grandifoUa Underw. North Amer. FI. 16(1):18. 1909. TYPE: Valparaiso, Sta. Marta,
Depto. Magdalena, Colombia, Smith 992 (NY not seen).

Danaea elliptica var. crispula Rosenst. Repert. Spec. Nov. Regni Veg. 6:310. 1909. TYPE: Rio
Mayo, near Tarapoto, Depto. S. Martin, Spruce 4770 (S not seen; isotypes L not seen photo 839a,
US).

LECZTOTYPE: Mt. Diablo, Jamaica, Sloane Herb. 1:85 (BM not seen), chosen
by Proctor (FI. Less. Antill. 2:48. 1977).

Plants terrestrial, at 0-600(1200) m elevation, in forests, from Puerto Viejo
(Pcia. Heredia), Carrillo (Pcia. S. Jose), Pejivalle (Pcia. Cartago), Barro Colorado
Island (Canal Zone), S. Jose Island and Juan Didz (Pcia. Panama), the vicinity of
Cana, and Cupica (Depto. Choc6). Also from the Antilles, Trinidad, Mexico to
Honduras, Nicaragua, Venezuela, Colombia, Peru, Bolivia, the Guianas, and
Brazil.

111. Danaea moritziana K. Presl, Suppl. Tent. Pterid. 35. 1846.

TYPE: Colonia Tovar, Edo. Aragua, Venezuela, Moritz 257 (PRC? not seen;
isotypes B not seen fragm NY, L not seen photo 842).

Plants terrestrial, at 700-1800 m elevation, in forests, from below the Alturas
de Nique (Pcia. Darien) and Alto del Buey and the Ciudad Bolivar - Quibdo road
(Depto. Choc6). Also from Venezuela, Colombia, Peru, and Bolivia.

This species may prove to be conspecific with D. cuspidata and is the earlier
name, or the Colombian coastal material alone may be D. cuspidata.

112. Danaea nodosa (L.) J. E. Smith, M6m. Acad. Roy. Sci. (Turin) 5:420, t. 9, f.
11. 1793.

Acrostichum nodosum L. Sp. PI. 2:1070. 1753. LECTOTYPE: Plate 108 of Plumier’s “Traite...”,
which is based on a specimen collected by Plumier near Port-de-Paix, Hispaniola, chosen by
Underwood (Bull. Torrey Bot. Club 29:671. 1902).

Danaea lon^folia Desv. Ges. Naturf. Freunde Berlin Mag. 5:307. 1811. TYPE: Antilles, collector
unknown (P-Hb. Desv. not seen), synonymized by Weatherby (Contr. Gray Herb. 94:23. 1936).
Danaea angustifolia K. Presl, Suppl. Tent. Pterid. 35. 1846. TYPE: Antilles, Bertero (P not seen).
Danaea elata Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:306 (repr.
154). 1849. TYPE: Hacienda de Jovo, Edo. Veracruz, Mexico, 1500 ft, Liebmann PI. Mex. 2199, FI.
Mex. 848 (C not seen), synonymized by A. R. Smith (FI. Chiapas 2:89. 1981).

Plants terrestrial, at 0-1700 m elevation, in forests, from the Cordillera Central,
the Cordillera de Talamanca, the Atlantic lowlands of Costa Rica and Panama,
and scattered localities in Pcia. Darien and the northern half of the Choco. Also
from the Antilles, Trinidad, Mexico to Honduras, Nicaragua, Venezuela to Peru,
Surinam, and Brazil.

113. Danaea plicata Christ, Repert. Spec. Nov. Regni Veg. 8:19. 1910.

TYPE: Carrillo, Pcia. S. Jose, 400 m, Erode & Erode 327 (P not seen).

FIGS. 112-115. Danaea. FIG. 112. Frond of D. nodosa, Mickel 3504. FIG. 113. Laminae of D.
plicata, McAlpin 1291. FIG. 114. Plant of D. serrulata, Killip 35263. FIG. 115. Plant of D.
wendlandii, Mickel 3526.

90 MARATTIACEAE: 9. DANAEA

Plants terrestrial, at 400-600 m elevation, in dense, wet forests, known only
from the type locality and from about 30 km above S. Ramon on the road to Los
Angeles Norte, Pcia. Alajuela {McAlpin 1291, DUKE, US).

114. Danaea serrulata Baker, J. Bot. Brit. For. 19:208. 1881.

TYPE: Colombia, Kalbreyer 1352 (K not seen).

Plants terrestrial, at 200-300 m elevation, in forests, in the Flora area known
only from the Corcovada region, upper Rio S. Juan, Depto. Choco (Killip 35263,
US).

This species may be conspecific with the earlier D. humilis Moore from Peru.
Both species are small, odd-pinnate, with falcate pinnae that are strongly
inequilateral at the base and somewhat serrate at the acute to acuminate apex.
Juvenile specimens of D, moritziana resemble adult specimens of D. serrulata, but
differ in having 15(18) or fewer, rather than more than 15(18), lateral pinna pairs
and in tending to have the pinnae obtuse, rather than acute, at the apex.

115. Danaea wendlandii Reichenb. f. Bot. Zeitung (Berlin) 30:490. 1872.

TYPE: Piedragas[?], Costa Rica, Wendland 744 (B not seen Alston photo).
Plants terrestrial, at 1800 m elevation, in forests, from above S. Ramon on the
road to Los Angeles Norte (Pcia. Alajuela), the Atlantic coastal plain of Costa
Rica, the Cordillera de Talamanca, Chagres (Pcia. Panama), between Cana and
Cuasi (Pcia. Darien), and the northwestern part of the Depto. Choc6. Also from
Nicaragua, Colombia, and Ecuador.

The stipes of this species are often alate and have but one node. The Jamaican
D. jenmanii Underw. is related.

OSMUNDACEAE: 10. OSMUNDA 91

OSMUNDACEAE

Monotypic in the Flora area; see description of Osmunda.

10. OSMUNDA L.

Plants terrestrial, in swamps, roadside ditches, and open areas; rhizomes erect,
stout, bearing fibrous roots; fronds medium-sized to large, caespitose, partially or
fully dimorphic, the fertile fronds (or portions of fronds) longer than the sterile
ones; stipes closely set in shallow spirals, subalate distal to the base (diamond-
shaped m cross-section), nearly woody and persistent at the base; rachises
stramineous; laminae pinnate-pinnatifid to 2(3)~pinnate, villous when young, often
glabrous at maturity or nearly so; sterile laminae or pinnae free-veined, the veins
forked; fertile fronds or pinnae without an expanded lamina, the sporangia borne
in clusters on the veinlets; sporangia thick-stalked, globular, with a rudimentary
annulus, bearing usually several hundred chlorophyllous, trilete spores.

Widely distributed in temperate to tropical regions; ca. 10 species.

BENEDICT, R. C. 1916. Osmundaceae. North Amer. FI. 16(l):27 - 28.

1. Fronds partially dimorphic, with an apical fertile portion; laminae glabrous, 2-pinnate. Laminae
ovate to lanceolate, 30 - 90 cm long, 9-30 cm wide.

117. O. regalis VM.palustris

1. Fronds entirely dimorphic, the sterile and fertile ones separate; laminae tomentose, especially in
the pinna axils, pinnate-pinnatifid. Sterile laminae narrowly elliptic, 30 - 65 cm long, 10-15(25) cm
wide.

116. O. dnnamomea var. imhricata

116. Osmunda cinnamomea var. imbricata (Kunze) Milde, Monogr. Osmund. 95.

1868.

Osmunda bipinnata L. Sp. PL 2:1065. 1753. TYPE: Plate 155 of Plumier’s ‘Traite...”, which is
based on a specimen collected by Plumier on Hispaniola.

Osmunda imbricata Kunze, Farrnkrauter 2:29, t. 112. 1849. TYPE: Edo. Merida, Venezuela,
Funck & Schlim 1221 (LZ destroyed).

Plants terrestrial, at 1200=- 1800 m elevation, in swamps, roadside ditches, and
open areas, in the Flora area known only from Cerro Tablazo, and the vicinity of
Juan Vihas and the Rio Macho dam (both Pcia. Cartago). Also from Bermuda,
the Greater Antilles, Mexico to Honduras, Venezuela, Colombia, Peru, Brazil,
and Paraguay.

According to Fernald (Rhodora 32:75. 1930), this variety differs from var.

dnnamomea in its narrower (1-2.3 cm wide), more coriaceous, and opaque
pinnae.

117. Osmunda regalis var. brasiliensis (Hook. & Grev.) Pic. Ser. Webbia 31:247.

1977.

Osmunda palustris Schrad. Goett. Gel. Anz. 1824:866. 1824. TYPE: Brazil, Prince Maxmillian
von Wied-Neuwied (LE? or M not seen).

Osmunda spectabilis var. brasiliensis Hook. & Grev. Bot. Misc. 3:230. 1833. TYPE: Serra do
Orgaos, Est. Rio de Janeiro, Brazil, Swainson (K not seen).

Plants terrestrial, at 1100-1800 m elevation, in swamps, roadside ditches, and
open areas, from Cerro Tablazo, Cerro Carpintera and Tapanti, and the
Cordillera de Talamanca to Pcia. Chiriqm. Also from El Salvador, Venezuela,
Colombia to Peru, and Brazil.

92 OSMUNDACEAE: 10. OSMUNDA

OSMUNDACEAE: 10. OSMUNDA 93

This variety differs from var. spectabilis (Willd.) A. Gray, which occurs from
North America to Honduras, in having short-petiolulate (4-6 mm long) pinnae
and pinnules, rigid and coriaceous laminae, and pinnules that are round or slightly
excised at the upper base.

FIGS. 116-121. Osmunda, Plagiogyria, and Schizaea. FIG. 116. Lamina apices of O. cinnamomea
var. imbricatay Yuncker et al. 5979, Honduras. FIG. 117. Lamina apex of O. regalis var. brasiliensis,
Pittier 7911. FIG. 118. Sterile frond of P. costaricensis, Mickel 3194. FIG. 119. Sterile frond of P.
semicordatay Pittier 2970. FIG. 120. Frond of S. eleganSy Valerio 1819. FIG. 121. Sterile and fertile
fronds of S. poeppigianay Pittier 10580.

94 PLAGIOGYRIACEAE: 1 1. PLAGIOGYRIA

PIAGIOGYRIACEAE

Monotypic; see description of Plagiogyria.

11. PLAGIOGYRIA (Kunze) Mett.

Plants terrestrial, in wet, usually open areas; rhizomes erect or nearly so, stout,
lacking hairs at maturity, bearing fibrous roots; fronds medium- to large-sized,
dimorphic, the fertile fronds with contracted pinnae, narrower and often shorter
than the sterile ones; stipes closely set in shallow spirals, subalate distal to the base
(diamond-shaped or broady triangular in cross-section), blackish, nearly woody
and persistent at the base; rachises stramineous, glabrous; laminae pinnatisect to
barely pinnate, glabrous, herbaceous; sterile laminae free-veined, the veins simple
or 1 -forked, often paired at origin, the margins distinctly serrate; fertile laminae
greatly reduced, with very wide, usually erose margins that are reflexed when
young, free-veined, the veins 1-forked, the branches apparently joining a
commissural vein, the sporangia borne on the forked veins, especially distally,
thick-stalked, subglobular, the annulus oblique; spores trilete, 48 per sporangium.

Tropical and subtropical, mostly Old World; ca. 35 species. A genus of
uncertain position resembling, but perhaps not related to, Osmunda.

LELLINGER, D. B. 1971. The American species of Plagiogyria sect. Carinatae. Amer. Fern J.

61:110-118.

1. Sterile laminae decidedly tapered at the base, the basal pinnae no more than half as long as the
longest pinnae. Sterile laminae elliptic, 30- 60(90) cm long, 8- 14 cm wide.

119. P. semicordata

1. Sterile laminae not much tapered at the base, the basal pinnae ca. 2/3 as long as the longest
pinnae. Sterile laminae lanceolate, 30 - 70 cm long, (9)12-15(30) cm wide.

118. P. costaricensis

118. Plagiogyria costaricensis Mett. ex Kuhn, Linnaea 36:149. 1869.

Plagiogyria anisodonta Copel. Philipp. J. Sci. 38:409, t. 9. 1929. TYPE: Volcan Poas, Pcia.
Alajuela, 2800 m, O. Jimenez 1018 (US).

Pla^ogyria denticulata Copel. Philipp. J. Sci. 38:412, t. 12. 1929. TYPE: Cani, 7 mi NE of Mito,
Depto. Huanuco, Peru, ca. 8500 ft, Macbride 3432 (US; isotype F).

TYPE: Volcdn Barba, Pcia. Heredia, Wendland 1066 (B or GOET not seen).
Plants terrestrial, at 1800-3300 m elevation, in open areas and subparamos,
from the Cordillera Central and the Cordillera de Talamanca to Cerro Chirripo.
Also from Venezuela, Colombia, Peru, and Bolivia.

119. Plagiogyria semicordata (K. Presl) Christ, Famkr. Erde 176. 1897.

Lomaridium semicordatum K. Presl, Abh. Konigl. Bohm. Ges. Wiss. V, 6:515 (repr. 155). 1851.
TYPE: Colombia, collector unknown (PRC not seen).

Plagiogyria biserrata Mett. Abh. Senckenberg. Naturf. Ges. 2:272, t. 15, f. 1-18. 1858.

LECTOTYPE: Colonia Tovar, Edo. Aragua, Venezuela, Moritz 400 (B not seen photo 10211;
isolectotypes HBG not seen photo 5339, US), chosen by Copeland (Phil. J. Sci. 38:409. 1929).

Plagiogyria obtusa Copel. Philipp. J. Sci. 38:413, t. 13. 1929. TYPE: Near Pico Turquino, Cuba,
Leon 11126 (US; isotype NY).

Plants terrestrial, at 2000-3000 m elevation, in open areas and subpdramos,
from the Cordillera Central and the Cordillera de Talamanca to Pcia. Chiriquf.
Also from Cuba, Jamaica, Mexico, Guatemala, Venezuela, and Colombia to
Bolivia.

SCHIZAEACEAE: 12. SCHIZAEA 95

SCHIZAEACEAE

Rhizomes ascending, short-creeping, multicipital, or long-creeping; hairy, the
hairs soft and lax to stiff and momliform; fronds monomorphic to partially or fully
dimorphic, small to medium-sized (to ca. 1 m) or very large and twining; stipes
sulcate or sometimes terete, stramineous to castaneous or atropurpureous, often
dark only at the base, glabrous or sometimes with an indument like that of the
rhizome, not articulate; laminae absent, reduced or flabellate in Schizaea and
Actinostachys, pinnately organized and determinate mAnemiay and indeterminate,
twining, and very large in Lygodium; ultimate segments with free, rarely casually or
fully anastomosing veins; sori marginal but sometimes appearing superficial, borne
on specialized, nonfoliose ultimate frond segments, protected by a modified
segment margin (or exindusiate in Anemia); sporangia usually few, large, sessile,
symmetrical or eccentric, the thickened cells of the terminal annulus 1-ca. 20,
distinct from the remainder of the thin-walled sporangium; spores monolete or
trilete, 32-256 per sporangium, distinctly ornamented.

1. Fronds indeterminate, twining, climbing and scrambling on other vegetation. Pinnules or
ultimate segments often rather palmate.

14. Lygodium

1. Fronds determinate, not twining, climbing, or scrambling..!.

2(1). Sterile laminae 1-3-pinnate, with expanded segments, never flabellate or grass-like.

15. Anemia

2(1). Sterile laminae unbranched and grass-like or dichotomously branched to flabellate, never
pinnately organized.. 3.

3(2). Fertile segments (sorophores) pinnate spikes somewhat resembling feathers; sterile portions
of fronds dichotomous to flabellate.

12. Schizaea

3(2). Fertile segments (sorophores) digitate; sterile portions of fronds undivided, flattened or
triangular in cross-section.

13. Actinostachys

12. SCHIZAEA J. E. Smith

Plants terrestrial, often in sandy, sterile soil; rhizomes short-creeping or
ascending, occasionally branched, densely bristly, the hairs brown, conspicuously
septate; fronds small to medium-sized, usually with reduced laminae; stipes
approximate, adaxially sulcate, often angular, pilose at the base with hairs like
those of the rhizome, longer than the laminae; laminae simple or dichotomously
divided, sometimes flabellate, papyraceous, thickened at the margin, glabrous or
inconspicuously and thinly pilose, the simple or dichotomously branched veins
usually visible; sporangia borne in 2 rows on the narrow, exindusiate segments of
the triangular to oblong, pinnatisect to subpinnately branched, laxly hairy
sorophores that are borne terminally at the apex or apical margins of the laminae;
sporangia pyriform or oblong.

Southern Hemisphere and North America, often in sandy soils; ca. 30 species.
LELLINGER, D. B. 1969. Schmeaceae (Filicales). In B. Maguire and collaborators. The botany
of the Geayana Highland -Part VllL Mem. New York Bot. Gard. 18:2-11.

1. Laminae almost flabellately divided into usually 8 or 16 segments (2)3 - 12 mm wide.

120. 5'. elegans

1. Laminae dichotomously divided into usually 32 or 64 linear segments 1-2 mm wide.

121. S. poeppigiana

2 cm

96 SCHIZAEACEAE: 12. SCHIZAEA

SCHIZAEACEAE: 13. ACTINOSTACHYS 97

120. Schizaea elegans (Vahl) J. E. Smith, M4m. Acad. Roy. Sci. (Turin) 5:419.
1793.

Acrostichum elegans Vahl, Symb. Bot. 2:104, t. 50. 1791. TYPE: Trinidad, von Rohr (C not seen).
Lophidium latifolium Rich. Actes Soc. Hist. Nat. Paris 1:114. 1792. TYPE: French Guiana, Le
Blond (P? not seen).

Schizaea flabellum Mart. Icon. PI. Crypt. Bras. 115, t. 55, f. 2. 1834. TYPE: Prov. Rio Negro, Est.
Amazonas, Brazil, Martins (M not seen; isotypes B not seen photo 10333, BR not seen photo 4755).

Schizaea pacificans Mart. Icon. PI. Crypt. Bras. 116, t. 56, f. 1. 1834. TYPE: Rio Pureos, Est.
Amazonas, Brazil, Martins (M not seen; isotype BR not seen).

Schizaea spectabilis Mart. Icon. PI. Crypt. Bras. 117. 1834. TYPE: Region of the Rio Japura, Est.
Amazonas, Brazil, Martins (M not seen).

Schizaea attennata Beyrich ex Sturm in Mart. FI. Bras. 1(2): 184. 1859. TYPE: “Prope Rio de
Janeiro...prope Botofogo,” Est. Rio de Janeiro, Brazil, Beyrich in 1824 (B not seen photo 10328;
isotype L not seen photo 2247).

Schizaea elegans var. amazonica Christ, Hedwigia 44:370. 1905. TYPE: Manaus, Est. Amazonas,
Brazil, Ule 5441 (B not seen photo 10332).

Plants terrestrial, at 0-1200 m elevation, in moist and dryish forests, from
Navarro (Pcia. Cartago), the valley of the Rio General, the Pacific lowlands of
western Panama, the Canal Zone and adjacent Pcia. Panama, and throughout the
Choc6. Also from Jamaica, Mexico to Honduras, Nicaragua, Trinidad, Venezuela,
Colombia to Bolivia, the Guianas, and Brazil.

121. Schizaea poeppigiana Sturm in Mart. FI. Bras. 1(2): 181. 1859.

Schizaea occidentalis Griseb. Cat. PI. Cub. 273. 1866. TYPE: Near Monte Verde, eastern Cuba,
Wright 926 (probably GOET not seen; isotype US).

ISchizaea digitata var. orbicnlaris Baker, J. Bot. Brit. For. 19:208. 1881. TYPE: Depto. Antioquia,
Colombia, 5000 - 6000 ft, Whyte (K not seen).

LECTOTYPE: “Ad Ventanilla de Cassapillo,” Peru, Poeppig (BR or M not
seen), chosen by Underwood (North Amer. FI. 16(1):38. 1909).

Plants terrestrial, at 600-1200 m elevation, in savannas, from Cerro Negro
(Pcia. Limon), Los Palmares (Pcia. S. Jose?), the valley of the Rfo Diqufs (Pcia.
Puntarenas), and Boquete. Also from the Bahamas, the Greater Antilles,
Venezuela, Colombia, Peru, Bolivia, and Guyana.

13. ACTINOSTACHYS Wall, ex Hook.

Plants terrestrial, often in sandy, sterile soil; rhizomes short-creeping or
ascending, branched, densely bristly, the hairs dark brown, conspicuously septate;
fronds medium-sized, without expanded laminae; stipes approximate, triangular in
cross-section, bristly at the very base with hairs like those of the rhizome;
sporangia borne in 4-6 rows on the narrow, exindusiate segments of the digitately
branched, laxly hairy sorophores that are borne terminally at the apex of the
fronds; sporangia pyriform or oblong.

Tropical regions, mostly Old World; ca. 10 species.

LELLINGER, D. B. 1969. Schizaeaceae (Filicales). In B. Maguire and collaborators. The botany
of the Guayana highland -Part VIII. Mem. New York Bot. Gard. 18:2- 11.

FIGS. 122-176. Actinostachys and Lygodinm. FIG. 122. Plant oiA. pennnla, Pittier 10555. FIG. 123.
Sterile pinnule of L. heterodoxnm, Bunting & Licht 668, Nicaragua. FIG. 124. Sterile pinnule of L.
micanSy Hayes 6. FIG. 125. Sterile pinnule of L. radiatanty Maxon 4735. FIG. 126. Sterile pinnule of
L. vennstnniy Lellinger 698.

98 SCHIZAEACEAE: 14. LYGODIUM

122. Actinostachys pennula (Swartz) Hook. Gen. Fil. t. IIIA. 1842.

Schizaea pennula Swartz, Syn. Fil. 150, 379. 1806. TYPE: “Habitat in America calidiore,”

collector unknown (S not seen).

Schizaea trilateralis Schkuhr, Vier Zwanzigste Kl. Linn. Pfl.-Syst. 2:137, t. 136. 1805. TYPE: Near
the Essequibo River, Guyana, Meyer (HAL not seen).

Schizaea penicillata Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:86. 1810, nom. superfl. TYPE: A
renaming of S. pennula, and so based on the type of that name.

Actinostachys germanii Fee, Hist. Foug. Antill. [Mem. Foug. 11]: 123, t. 29, f. 3. 1866. TYPE:
Sainte-Rose, Guadeloupe, Germain (P not seen), synonymized by Wagner and Quevedo (Abstr. Bot.
Soc. Amer. 927. 1985).

Plants terrestrial, at 200-700 m elevation, in open areas, from Los Palmares
(Pda. S. Jose?), the valley of the Rio General, and Alto de Pure, Baja Talamanca
(Pda. Limon). Also from Puerto Rico, the Lesser Antilles, Trindad, Venezuela,
Peru, Guyana, Surinam, Brazil, and Uruguay.

14. LYGODIUM Swartz

Plants terrestrial, scrambling or sometimes high-climbing vines often at forest
margins; rhizomes short-creeping, ca. 1 cm in diam., bristly, the hairs stiff,
blacosh, septate; stipes approximate, blackish and bristly at the base, flexible;
rachises elongate, much longer than the stipes, indeternunate; pinnae alternate,
reduced to a short or obsolete stalk bearing a single pair of lateral (secondary)
branches and terminated by a dormant bud; pinnae or pinnules dichotomously
divided or simple and with basal lobes, papyraceous, thickened at the margin,
commonly sparsely pilose at least on the veins, the veins free and forked or
forming elongate areolae without included veinlets; sporangia borne in 2 rows on
the narrow, elongate, simple sorophores, protected by highly modified, scale-like,
antrorse folds of the sorophore surface.

Mostly tropical and mostly in the Old World; ca. 40 species.

DUEK, J. J. 1978. A taxonomic revision of Lygodium (Filicinae) in America. Feddes Repert.
89:411-423.

LELLINGER, D. B. 1969. Schizaeaceae (Filicales). In B. Maguire and collaborators. The botany
of the Guayana highland -Part VIII. Mem. New York Bot. Gard. 18:2- 11.

1. Veins of the laminae reticulate. Lateral (secondary) branches each with 1-3(5) stalked, simple
or dichotomous pinnules, each with two equal basal lobes.

123. L. heterodoxum

1. Veins of the laminae free..2.

2(1). Lateral (secondary) branches usually ca. 3 cm long, bearing a single palmate or dichotomous
pinnule. Ultimate segments (10)15-30 cm long, 1.5 -2.5 cm wide.

125. L. radiatum

2(1). Lateral (secondary) branches usually ca. 5-15 cm long, bearing several simple or pinnate
pinnules.. 3.

3(2). Ultimate segments not articulate, without a node-like swelling at the apex of the stalk; simple
pinnules 3-5(8) cm long, 1(2) cm wide distal to the base, lobed at the base, the lobes with a costa.

126. L. venustum

3(2). Ultimate segments articulate, with a node-like swelling at the apex of the stalk; simple
pinnules (4)9-13 cm long, 1.5-2 cm wide, not or scarcely lobed at the base, the lobes without a
costa.

124. L. micans

SCHIZAEACEAE: 14. LYGODIUM 99

123. Lygodium heterodoxum Kunze, Faimkrauter 2:32, 1. 113. 1849.

Hydroglossum spectabile Liebm. Kongel. Danske Vidensk, Selsk. Skr., Naturvidensk. Afd. V, 1:299
(repr. 147). 1849. LECTOTYPE: S. Pedro Tepinapa, Edo. Oaxaca, Mexico, Liebmann PI. Mex.
2634, FI. Mex. 936 (C not seen; isolectotype US), chosen by A. R. Smith (FI. Chiapas 2:145. 1981).

Hydroglossum mexicanum F6e, Cat. Foug. Mex. [M6m. Foug. 9]: 42. 1857. SYNTYPES: Edo.
Oaxaca, Mexico, Galeotti 6419 (P or RB not seen); and Mexico, Linden & lungers 763 (P or RB not
seen).

TYPE: Mexico, Galeotti 6419bis (LZ destroyed; isotypes BR as ''6419 ^ the
logical lectotype).

Plants terrestrial, at 0-300(800) m elevation, from the Pacific coastal plain and
adjacent foothills of Costa Rica to the Peninsula de Burica of Panama. Also from
Mexico to Hondur2LS and Nicaragua.

124. Lygodium micans Sturm in Mart. FI. Bras. 1(2): 178. 1859.

Lygodium scandens sensu Schkuhr, Vier Zwanzigste Kl. Linn. Pfl.-Syst. 2:138, t. 138. 1805, nom.
illeg., non Swartz, 1801. TYPE LOCALITY: Essequibo, Guyana.

Hydroglossum heptaphyllum Schrad. Goett. Gel. Anz. 1824:863. 1824. TYPE: Brazil, Prince
Maxmillian von Wied-Neuwied (M not seen).

Lygodium acuminatum Sturm in Mart. FI. Bras. 1(2):174, t. 14, f. 12. 1859. TYPE: Mt.

Corcovado, Agua de Serra, Est. Rio de Janeiro, Brazil, Pohl 3859 (W? not seen; isotype BR not seen
photo 4765).

TYPE: Guyana, Schomburgk 399 (B not seen; isotype BM not seen).

Plants terrestrial, at ca. 100 m elevation, in the lowlands of central Panama.
Also from Nicaragua, Venezuela, Trinidad, Colombia, Peru, Guyana, and Brazil.

This species is closely related to (and some think conspecific with) L. volubile
Swartz. It differs in having subcordate to truncate sterile pinnules and veins of the
fertile segments strigose abaxially, rather than usually oblique-truncate sterile
pinnules and veins and surface of the fertile segments densely sericeous abaxially.

125. Lygodium radiatum Prantl, Unters. Morph. Ge^sskiypt. 2:66. 1881.

Lygodium di^tatum D. C. Eaton, Mem. Amer. Acad. Arts N.S., 8:217. 1860, non K. Presl, 1825,
nom. illeg. SYNTYPES: Panama Railway at Gatun, Pcia. Colon, Hayes 25 (YU not seen; isotypes
GH, US); and Falls of the Rfo Truando, Depto. Choc6, Schott 77 (YU not seen; isosyntypes B? not
seen, MO).

TYPE: Based on L. digitatum D. C. Eaton, and so based on the type of that
name.

Plants terrestrial, at 0 - 400(900) m elevation, in lowland forests and swamps
from the valley of the Rfo General, the Peninsula de Osa, the lowlands of Panama,
and the northern half of the Depto. Choco. Also from Colombia and Peru.

126. Lygodium venustum Swartz, J. Bot. (Schrader) 1801(2): 303. 1804.

Hydroglossum hastatum Willd. Sp. PI. ed. 4, 5:79. 1810, nom. superfl. TYPE: A renaming of L.

venustum Swartz, and so based on the type of that name.

Hydroglossum hirsutum Willd. Sp. PI. ed. 4, 5:80. 1810. TYPE: Cumanacoa, Edo. Sucre,

Venezuela, Humboldt & Bonpland (B-Hb. Willd. 19479 not seen microfiche S. I. Library).

Lygodium polymorphum H.B.K. Nov. Gen. Sp. 1:25. 1816, non Ugena polymorpha Cav., 1801.
TYPE: Mt. Quetepe, below Cuman4 and Cumanacoa, Edo. Sucre, Venezuela, Humboldt &
Bonpland (P-Hb. Humb.? not seen).

Lygodium mexicanum K. Presl, Reliq. Haenk. 1:72. 1825. TYPE: Mexico, Haenke (PRC not seen
fragm NY).

100 SCHIZAEACEAE: 15. ANEMIA

Lygodium pohlianum K. Presl, Suppl. Tent. Pterid. 105 (postpr. 366). 1845. TYPE: Serra de S.
Jezabel and S. Luzia, Est. Minas Gerais, Brazil, Pohl (PRC not seen; isotype BR not seen photo
4763).

Lygodium commutatum K. Presl, Suppl. Tent. Pterid. 110 (postpr. 371). 1845. SYNTYPES:
Hacienda S. Luca, Depto. Chontales, Nicaragua, Friedrichsthal (PRC? not seen); Porto Bello, Pcia.
Col6n, Panama, Billberg (PRC? not seen); Peru, Poeppig (PRC? not seen); and Chile, Cuming
(PRC? not seen).

Lygodium schiedeanum K. Presl, Suppl. Tent. Pterid. 110 (postpr. 371). 1845. TYPE: Based on
Lygodium mexicanum sensu Schlechtend. & Cham. (Linnaea 5:620. 1830), and ultimately on the
basis of that name, which is Papantla, Edo. Veracruz, Mexico, Schiede (HAL? not seen).

Lygodium schiedeanum var. tenuilobum K. Presl, Suppl. Tent. Pterid. 110 (postpr. 371). 1845.
TYPE: Papantla, Edo. Veracruz, Mexico, Schiede (PRC? not seen).

Lygodium palmatilobum Sturm in Mart. FI. Bras. 1(2): 170, t. 14, f. 6. 1859. TYPE: Rio de
Janeiro, Est. Rio de Janeiro, Brazil, Doellinger (BR not seen photo 4762).

Lygodium mucronulatum Sturm in Mart. FI. Bras. 1(2):171, t. 14, f. 9. 1859. SYNTYPES: Near
Paramaribo, Surinam, Ehrhard (BR? not seen); and near Caracas, Distr. Fed., Venezuela, Moritz (B
not seen).

Lygodium venustum vdx.granatense Christ, Bot. Jahrb. Syst. 24:145. 1897. TYPE: Grenada, Eggers
6293 (P not seen; isotype US).

TYPE: Brazil, Breynius (S? not seen), according to Proctor (FI. Less. Antill.
2:51. 1977).

Plants terrestrial, at 0-600 m elevation, in forests, river banks, and savannas,
from the lowlands of Costa Rica and Panama. Also from Cuba, Jamaica,
Hispaniola, the Lesser Antilles, Mexico to Nicaragua, and tropical South America.

15. ANEMIA Swartz

Plants terrestrial or sometimes epipetric, often in open areas; rhizomes short-
creeping or ascending, hairy, the hairs abundant, long, straight, soft; fronds mostly
small, partially or rarely completely dimorphic, the basal pinna pair upright, highly
branched, elaminate; stipes approximate, sparsely pilose; rachises stramineous,
sparsely pilose; sterile portion of the laminae or sterile fronds oblong or lanceolate
to triangular, spreading, 1-3-pinnate, sparsely pilose; fertile portion of the
laminae or fertile fronds subglabrous, stramineous, bearing sporangia in 2 rows on
the ultimate segments of the fertile portion; sporangia ovoid to globular.

Tropical and subtropical regions, mostly New World, especially Brazil; ca. 90
species.

MICKEL, J. T. 1962. A monographic study of the fern genus Anemia, subgenus Coptophyllum.
Iowa State J. Sci. 36:349 - 482.

. 1981. Revision of Anemia subgenus Anemiorrhiza (Schizaeaceae). Brittonia 33:413-429.

1. Veins entirely anastomosing (partially so in hybrids with A. hirta), except at the frond margins.
Sterile laminae ovate-deltate, (6)8-22 cm wide, (7)10-30 cm long, the pinnae simple, unlobed,
(2)3 -4(7) lateral pairs; stipes 10-45 cm long.

133. A. phyllitidis
1. Veins entirely free..2.

2(1). Sterile portion of the laminae pinnate, the pinna margins entire to shallowly crenate, never
lacerate..7.

2(1). Sterile portion of the laminae or sterile fronds pinnate-pinnatifid to 3-pinnate..3.

3(2). Fronds completely dimorphic, the fertile ones separate from the sterile ones; sterile laminae
1 - 2.5 cm long, dissected into linear segments ca. 0.5 mm wide.

130. A. millefolia

SCHIZAEACEAE: 15. ANEMIA 101

3(2). Fronds partially dimorphic, the fertile portion erect, usually long-stalked and exceeding the
sterile portion; sterile portion of the laminae at least 3 cm long, not dissected into linear segments.. 4.

4(3). Sterile portion of the laminae pinnate-pinnatifid to barely 2-pinnate; rhizomes erect or
ascending, the hairs reddish-brown; segments entire or crenately lobed at the apex..6.

4(3). Sterile portion of the laminae 2-3-pinnate; rhizomes short- to long-creeping, the hairs
blackish to brown; segments sharply serrate or crenate at the apex..5.

5(4). Sterile segments cuneate at the base, almost stalked, sharply serrate at the apex; sterile
laminae dr^g pale green, often subglabrous. Stipes 4-60 cm long; sterile portion of the laminae
lanceolate to deltate, 5-30 cm long, 2-25 cm wide; ultimate segments with nearly parallel veins ca.
0.25 mm distant.

127. A. adiantifolia

5(4). Sterile segments not cuneate at the base, broadly attached, crenate at the apex; sterile
laminae medium to dark green, drying brown, sparsely pilose. Stipes 4-10(20) cm long; sterile
portion of the laminae ovate, 5-8(12) cm long, 2.5-8(10) cm wide; ultimate segments with diverging
veins ca. 0.5 mm distant.

134.^. tomentosa var. mexicana

6(4). Fertile portion of the laminae exceeding the sterile portion, upright. Stipes 2-18(30) cm
long; sterile portion of the laminae oblong-lanceate, 3-11 cm long, 1.5-4 cm wide; ultimate
se^ents widest distal to the base, oblanceolate or liguliform, entire, with nearly parallel veins ca.
0.25 mm distant.

129. A. hirsuta

6(4). Fertile portion of the laminae shorter than or equalling the sterile portion, not upright.
Stipes 5-12 cm long; sterile portion of the laminae elliptic- lanceolate, 6-11 cm long, 3-5 cm wide;
ultimate segments widest at the base, oblong, crenately lobed, with diverging veins ca. 0.5 mm
distant.

128^. cUnata

7(2). Sterile fronds eiBtipitate or nearly so, the stipes 1-3(5) cm long, forming a basal rosette;
pinnae wide, round at the apex. Sterile laminae oblong to obovate-lanceolate, 2-6 cm long, 1.3 -2.5
cm wide; stipes of fertde fromk 1-5(9) cm long; sterile portion of fertile laminae similar to the
sterile laminae.

131. ^. obiongifoiia

7(2). Sterile fronds with obvious stipes 2-6 cm long, not forming a basal rosette; pinnae narrow,
mostly acute to obtuse at the apex. Sterde lammae lanceate, widest at the base, acute at the apex,
3- 10 cm long, 2-5.5 cm wide; sterile portion of fertile laminae similar to the sterile lammae.

132. A. pastinacaria

127. Anemia adiantifolia (L.) Swartz, Syn. Fil. 1S7. 1806.

Osmunda adiantifolia L. Sp. PI. 2:1065. 1753. LECTOTYPE: Plate 158 of Plumier’s “Trait6...”,
which is based on a specimen collected by Plumier on Hispaniola, chosen by Proctor (FI. Less. Antill.
2:53. 1977).

Osmunda asplenifolia Sav. in Lam. Encyc. Meth. 4:652. 1798. TYPE: Santo Domingo,

Hispaniola, Martin (P-Hb. Lam. not seen photo 2845).

Anemia carvifoiia K. Presl, Reliq. Haenk. 1:74. 1825. TYPE: Mexico, Haenke (PRC not seen).

Anemia adiantifolia var. helveola F6e, Cat. Foug. Mex. [M6m. Foug. 9]:41. 1857. TYPE: Villa
Alta, Edo. Oaxaca, Mexico, Gaieotti 6S85bis (P not seen).

Anemia adiantifolia var. distans Fee, Cat. Foug. Mex. [M6m. Foug. 9j:41. 1857. TYPE: Talca,
Edo. Oaxaca, Mexico, Gaieotti 6567bis (P not seen).

Attemia adiantifolia var. pumila Christ, Bull. Herb. Boissier II, 7:794. 1907. TYPE: Between
Cayey and Cidra along the Rio Plata, Puerto Rico, Sintenis 2194 (P not seen; isotype US).

Anemia adiantifolia var. subaunta Christ, Bull. Herb. Boissier II, 7:794. 1907. TYPE: Utuado, Los
Angeles, Puerto Rico, Sintenis 5948 (P not seen).

102 SCHIZAEACEAE: 15. ANEMIA

135 C. pteridoides
136 C. pteridoides x richardii

SCHIZAEACEAE: 15. ANEMIA 103

Plants terrestrial, at ca. 100 m elevation, along rivers, from the Inter american
Highway along the Rio Ahogados, 14 km northwest of Liberia (Pcia. Guanacaste).
Also from Florida, the Bahamas, the Antilles, Trinidad, Mexico to Honduras, and
Venezuela, Ecuador, and Brazil.

128. Anemia clinata Mickel, Amer. Fern J. 56:58, t. 4. 1966.

TYPE: Along the Rio Perene near Hacienda 3, Colonia Perene, Depto. Jurun,
Peru, ca. 600 m, Killip & Smith 25194 (US; isotypes F not seen, NY not seen).

Plants terrestrial, at ca. 800 m elevation, in the Flora area known only from
between S. Felix and Cerro Flor, Pcia. Chiriqui {Allen 1925, GH not seen, MO not
seen, UC, US). Also from Venezuela, Colombia, Peru, and Bolivia.

129. Anemia hirsuta (L.) Swartz, Syn. Fil. 156. 1806.

Osmunda hirsuta L. Sp. PI. 2:1064. 1753. LECTOTYPE: Plate 162 of Plumier’s “Traite...”,
chosen by Lellinger (Proc. Biol. Soc. Wash. 98:366. 1985).

Osmunda filiformis Sav. in Lam. Encyc. M6th. 4:652. 1798. TYPE: “Ex Carolina meridionah”
[locality false], collector unkonwn (P-Hb. Lam. not seen photo 2844).

lAnemia repens Raddi, Opusc. Sci. 3:282. 1819; PI. Bras. Nov. Gen. 1:71, t. 9, f. 2b. 1825. TYPE:
Mt. Corcovado, Est. Rio de Janeiro, Brazil, Raddi (FT not seen).

Anemia dissecta K. Presl, Rehq. Haenk. 1:74, t. 11, f. 4. 1825. TYPE: Mexico, Haenke (PRC not
seen probable isotype BR not seen photo 4790).

Anemia hirsuta var. achilleifolia Martens & Galeotti, Nouv. M6m. Acad. Roy. Sci. Bruxelles 15:20.
1842, as “achilleaefolia.’' TYPE: Near Zacatep6que and Juquila, Edo. Oaxaca, Mexico, 2000 - 6000
ft, Galeotti 6363 (BR not seen photo 4777; isotype LE not seen).

Anemia hirsuta var. inteffifolia Fourn. Mexic. PI. 1:139. 1872. TYPE: Ultimately a renaming of
Osmunda filiformis Sav. in Lam., and so based on the type of that name.

Anemia hirsuta var. dissecta Fourn. Mexic. PI. 1:139. 1872, nom. superfl. TYPE: A renaming of
A. hirsuta var. achilleifolia Martens & Galeotti, and so based on the type of that name.

Anemia hirsuta var. schwackeana Christ, Bull. Herb. Boissier II, 2:695. 1902. SYNTYPES: Sta.
Luzia do Rio das Velhas, Est. Minas Gerais, Brazil, Schwacke (P not seen); S. Joao d’El Rey, Brazil,
Silveira 403 (P not seen); Chapadao, Est. Goids, Brazil, Ule 536, 537, 3204, and 3205 (all P none
seen).

Anemia hirsuta var. subfiliformis Christ, Bull. Herb. Boissier II, 2:695. 1902. SYNTYPES:

Presumably Chapadao, Est. Goias, Brazil, Ule 535 (P not seen) and 3203 (P not seen).

Anemia hirsuta var. humboldtiana Hieron. Bot. Jahrb. Syst. 34:566. 1905. LECTOTYPE:

Between Caripe and Sta. Cruz, Edos. Monagas- Sucre, Venezuela, Humboldt 459 (B-Hb. Willd.
19495-2 not seen Tryon photo), chosen by Lellinger (Proc. Biol. Soc. Wash. 98:366. 1985).

Plants terrestrial or epipetric, at 0-1200(1700) m elevation, in open areas, often
on roadside banks, from the Cordillera Central, the Fila de Cedral, the Meseta
Central, the Pacific lowlands of Pcia. Guanacaste to the Canal Zone and adjacent
Pcia. Panama, the mountains south of Sta. Fe, and between Las Margarita and El
Valle (Pcia. Code). Also from the Antilles, Mexico to Nicaragua, Trinidad,
Venezuela, Colombia, Peru, Bolivia, and Brazil.

FIGS. Ill -131 . Anemia and Ceratopteris. FIG. 127. Frond of A. adiantifolia, Lellinger 699. FIG.
128. Frond of A. clinata, Allen 1925. FIG. 129. Frond of A. hirsuta, Stevens 153. FIG. 130. Plant of
A. millefolia, Allen 1166. FIG. 131. Plant of A. oblongifolia, Duke 8641. FIG. 132. Frond of A.
pastinacaria, Erlanson 582. FIG. 133. Frond of A. phyllitidis, Alfaro 16295. FIG. 134. Plant of^.
tomentosa var. mexicana, Shreve 6738, Mexico. FIG. 135. Sterile frond of C. pteridoides, Standley
31624. FIG. 136. Sterile frond of C. pteridoides x richardii, Pittier 6852. FIG. 137. Sterile frond of C.
thalictroides, Luetzelburg 12912, Brazil.

104 SCHIZAEACEAE: 15. ANEMIA

Anemia hirsuta is known to hybridize with several other species oi Anemia in the
Flora area, including ^4. oblongifolia from Nuestro Amo (Pcia. S. Jose) and S. Jose
Island, Pcia. Panama (Erlanson 528, GH, US). Gomez (Brenesia 18:156. 1980)
has found hybrids with^. phyllitidis {A. xdidicusana Gomez) along the road to El
Rodeo, Villa Colon, Pcia. S. Jose {Gomez 7130, CR, MICH not seen) and at
Navarro, Pcia. Cartago {Gomez 7183, CR); with A. hirta from Boruca, Pcia.
Puntarenas {Tonduz 4439, CR); and withal, pastinacaria from near Puriscal, Pcia.
S. Jose {Gomez 853, CR) and from near El Rodeo {Gomez 7131, 7169, both CR).

130. Anemia millefolia (Gardn.) K. Presl, Suppl. Tent. Pterid. 80 (postpr. 341).
1845.

Coptophyllum millefolium Gardner, Lx)ndon J. Bot. 1:133. 1842. TYPE: Near Villa de Arayas,
Est. Goias, Brazil, Gardner 4083 (K not seen; isotypes F not seen, GH not seen, NY not seen, P not
seen photo 3564, U not seen photo 125).

Plants terrestrial, at 600-1000 m elevation, in open areas, from Laguna La
Yeguada (Pcia. Veraguas), the vicinity of El Valle, and Cerro Camapana (Pcia.
Panama). Also from Venezuela, Colombia, and Brazil.

131. Anemia oblongifolia (Cav.) Swartz, Syn. Fil. 156. 1806.

Osmunda oblon^folia Cav. Icon. Descr. PI. 6:69, t. 592, f. 2. 1801. TYPE: Panama, Nie (MA not
seen), examined by Christensen (Dansk Bot. Ark. 9(3):31. 1937).

Osmunda humilis Cav. Icon. Descr. PI. 6:69, t. 592, f. 3. 1801. TYPE: Taboga, Pcia. Panama, N6e
(MA not seen).

Anemia pilosa Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:19, t. 2, f. 1. 1842.
TYPE: Juquila, Edo. Oaxaca, Mexico, 2000 - 6000 ft, Galeotti 6353 (BR not seen photos 4785, 4786).

Anemia pumila Klotzsch, Linnaea 18:526. 1845. TYPE: Caripe, Edo. Monagas, Venezuela, Moritz
159 (B not seen; isotypes L not seen photo 344, P not seen photo 3512).

Anemia seemannii Hook. London J. Bot. 7:564, t. 16. 1848. TYPE: Taboga, Pcia. Panama,
Seemann (K not seen fragm NY not seen).

Anemia cornea Prantl, Unters. Morph. Gef^krypt. 2:104. 1881. TYPE: Trapiche de la

Concepcion, Edo. Oaxaca, Mexico, Liebmann 2622 (C not seen; isotype BM not seen photo 7717,
fragm US).

1 Anemia presliana Prantl, Unters. Morph. Gefasskrypt. 2:104. 1881. SYNTYPES: Vella Boa, Est.
Goi^s, Brazil, Pohl (M and W neither seen); Est. Goi^s, Brazil, Gardner 4087 (W not seen); and
Tijuca, Mariana, Est. Minas Gerais, Brazil, Martius (M not seen).

Anemia ulei Christ in Schwacke, PI. Nov. Mineir. 2:36. 1900. SYNTYPES: Serra dos Pyreneos,
Est. Goias, Brazil, Ule 386 (P not seen photo 3582) and 3198 (P not seen).

Plants terrestrial or epipetric, at 0-1200 m elevation, in savannas and open
areas, from the Pacific coastal plain of northwestern Costa Rica and the adjacent
foothills, the lower reaches of the Meseta Central, the mountains of Panama from
Boquete to El Valle, between Las Minas and Pese (Pcia. Herrera), and the Canal
Zone and adjacent Pcia. Panama. Also from Mexico to Nicaragua, Venezuela to
Bolivia, Guyana, and Brazil.

132. Anemia pastinacaria Moritz in Prantl, Unters. Morph. Gefasskrypt. 2:110.
1881.

Anemia pilosa var. longistipes Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V,
1:301 (repr. 149). 1849. TYPE: Between Trapiche de la Concepcibn and Tepitongo, Edo. Oaxaca,
Mexico, Liebmann (C not seen).

LECTOTYPE: “In convalli del Tigre,” Venezuela or Colombia, Moritz 26 (B
not seen), chosen by Lellinger (Proc. Biol. Soc. Wash. 89:367. 1985).

SCHIZAEACEAE; 15. ANEMIA 105

Plants terrestrial, at 0-1000 m elevation, in savannas and thickets, from the
Pacific coastal plain of Costa Rica and its adjacent foothills, near David and
Boquete (Pcia. Chiriqm), near Las Margaritas (Pcia. Code), Ocu (Pcia. Herrera),
the Canal Zone, and Cerro Campana and S. Jose Island (Pcia. Panama). Also
from Jamaica, Mexico to Honduras, Nicaragua, Trinidad, Venezuela, Colombia,
Peru, Bolivia, Surinam, and Brazil.

See A. hirsuta for a discussion of hybrids with that species.

133. Anemia phyllitidis (L.) Swartz, Syn. Fil. 155. 1806.

Osmunda phyllitidis L. Sp. PI. 2:1064. 1753. TYPE: Based on plate 156 of Plumier’s “Traite...”,
which is based on a specimen collected by Plumier on Hispaniola.

Anemia fraxinifolia Raddi, Opusc. Sci. 3:282. 1819; PL Bras. Nov. Gen. 1:69, t. 8bis. 1825. TYPE:
Near Rio de Janeiro, Est. Rio de Janeiro, Brazil, Raddi (FI not seen).

Anemia sorbifolia Schrad. Goett. Gel. Anz. 1824:864. 1824. TYPE: “Pr. Bah. et Sebastianop.,”
Brazil, Maxmilian von Wied-Neuwied (LE or M not seen; isotype BR not seen photo 4784).

Anemia obliqua Schrad. Goett. Gel. Ajiz. 1824:864. 1824. TYPE: Brazil, Maxmilian von Wied-
Neuwied (LE or M not seen).

Anemia longifolia Raddi, PI. Bras. Nov. Gen. 1:69, t. 8. 1825. TYPE: Near Rio de Janeiro, Est.
Rio de Janeiro, Brazil, Raddi (FI not seen).

Anemia cordifolia K. Presl, Rehq. Haenk. 73, t. 11, f. 3. 1825. SYNTYPES: “Hab. in Mexico et in
vallibus Cordillerarum Peruviae,” Haenke (PRC neither seen).

Anemia haenkei K. Presl, Reliq. Haenk. 1:74. 1825. TYPE: Cordillera de Peru, Haenke (PRC not
seen).

Anemia densa Link, Hort. Reg. Bot. Berol. 2:142. 1833. TYPE: Cultivated in the garden at Berlin,
originally from Brazil, Link (B not seen; authentic material HBG not seen photo 5430, L not seen
photo 345).

Anemia langsdorffiana K. Presl, Suppl. Tent. Pterid. 89 (postpr. 350). 1845. TYPE: Ilha Sta.
Catarina, Est. Sta. Catarina, Brazil, Langsdorff (JlE not seen).

Anemia phyllitidis var. caryotidea Christ, Bull. Herb. Boissier II, 2: 692. 1902. TYPE: Brazil,
Glaziou (P not seen photo 3573).

Anemia phyllitidis f. subtripinnatifida Rosenst. Hedwigia 43:233. 1904. TYPE: Toledo, Est. S.
Paulo, Brazil, Ulbricht 74 (S not seen).

Anemia phyllitidis f. auritolobata Rosenst. Hedwigia 46:159. 1907. SYNTYPES: Porto Alegre,
Mun. Sta. Cruz, Est. Rio Grande do Sul, Brazil, Juergens & Steir 6b (S not seen); and Lages, Est. Sta.
Catarina, Brazil, Spannagel (S not seen).

Anemia phyllitidis f. transitoria Rosenst. Hedwigia 46:159. 1907. TYPE: Porto Alegre, Mun. Sta.
Cruz, Est. Rio Grande do Sul, Brazil, Juergens & Steir 6c (S not seen).

Anemia lancea Christ, Bull. Herb. Boissier II, 7:791. 1907. TYPE: Est. Mato Grosso, Brazil, H.

Smith 143 (P not seen; isotype R not seen).

Anemia phyllitidis var. pygmaea Christ in Wettst. Denkschr. Kaiserl. Akad. Wiss., Math.-
Naturwiss. Kl. 79:52, t. 7. 1907. SYNTYPES: Near Yporanga, valley of the Rio Ribeira, Est. S.
Paulo, Brazil, 130 m, Wettstein (P not seen); and Apiahy, Est. S. Paulo, Brazil, Puiggari (P not seen).

Plants terrestrial, at 0 - 1700 m elevation, in pastures and on banks, from the
Cordillera Central, the lower reaches of the Meseta Central, the Fila de Cedral,
Cerro Carpintera, S. Isidro del General (Pcia. S. Jose), the Cordillera de
Talamanca (Pcia. Chiriqm), Tonosi (Pcia. Los Santos), Penonome and El Valle,
La Yeguada (Pcia. Veraguas), and Aquarubia (Pcia. Panama). Also from Cuba,
Jamaica, Hispaniola, Mexico to Nicaragua, Trinidad, Venezuela, Colombia to
Bolivia, Guyana, Surinam, Brazil, Paraguay, Uruguay, and Argentina.

This is the only species of Anemia in the Flora with anastomosing veins, which
are uncommon in the genus. See A. hirsuta for comments on hybrids with that

106 SCHIZAEACEAE: 15. ANEMIA

species. Hybrids with A. hirta (L.) Swartz are known from Finca dos Padres (Pcia.
S. Jose) and El Llano (Pcia. Cartago).

134. Anemia tomentosa var. mexicana (K. Presl) Mickel, Iowa State J. Sci. 36:427.
1962.

Anemia fulva var. mexicana K. Presl, Suppl. Tent. Pterid. 84 (postpr. 345). 1845. TYPE: Eastern
Mexico, Leibold (LZ destroyed).

Plants epipetric, at ca. 750 m elevation, in the Flora area known only from along
a stream flowing into Laguna La Yeguada, Pcia. Veraguas (Armond 505, DUKE).
Also from Hispaniola, Mexico, Venezuela, and Colombia.

PARKERIACEAE: 16. CERATOPTERIS 107

PARKERIACEAE

Monotypic; see description of Ceratopteris.

16. CERATOPTERIS Link

Plants aquatic, rooted or floating; rhizomes usually erect, obsolete, scaly, the
scales sparse, tan, wide, flaccid; fronds medium-sized, dimorphic, the fertile ones
longer than the sterile ones, with proportionally longer stipes and with contracted
laminae, glabrous, herbaceous; stipes approximate, glabrous, fleshy and collapsing
when dry, bearing many long, thick, smooth roots near the base; sterile laminae
simple (pinnately lobed) to 3-pinnate, the segments wide, often lanceate, the
venation reticulate, the areolae rounded-polygonal, irregular, without included
veinlets, sometimes bearing proliferous buds at the pinna or segment bases; fertile
laminae 2 ~ 3-pinnate, the segments narrow, linear, the venation reduced to a
single, central vein and a commissural vein near each margin, not reticulate, the
margins wide, revolute, protecting the submarginal sporangia; sporangia borne in
1-4 rows, globular, nearly sessile, the annulus irregularly developed, sometimes
absent or vestigial; spores 16 or 32 per sporangium, globular, trilete.

Pantropical and subtropical; 4 species.

LLOYD, R. M. 1974. Systematics of the genus Ceratopteris Brongn. (Parkeriaceae) II. Taxonomy.

Brittonia 26:139-160.

1. Sterile laminae with the ultimate segments acute-attenuate. Plants rooted or floating; sterile
laminae 2-pinnate-pinnatifid.

136. C. pteridoides x richardii

1. Sterile laminae with the ultimate segments round, obtuse, or acute, or the laminae simple.. 2.

2(1). Sterile laminae pinnate to 3-pinnate, the basal lobes or pinnae alternate; lamina buds usually
developing into plantlets only on senescent laminae; plants usually rooted.

137. C. thalictroides

2(1). Sterile laminae simple (rarely pinnate or 2-pinnate), the basal lobes or pinnae mostly
opposite; lamina buds usually developing into plantlets on actively growing laminae; plants usually
floating.

135. C. pteridoides

135. Ceratopteris pteridoides (Hook.) Hieron. Bot. Jahrb. Syst. 34:561. 1905.

Parkeria pteridoides Hook. Exot. FI. 2:t. 147. 1825. LECTOTYPE: Essequibo District, Guyana,
Parker (K not seen), chosen by Lloyd (Brittonia 26:156. 1974).

Parkeria lockhartii Hook. & Grev. Icon. Fil. l:ad t. 97. 1828. TYPE: About 4 mi SW of Arima,
Trinidad, Lockhart (K not seen).

Ceratopteris parkeri J. Smith, J. Bot. (Hooker) 4:70. 1841, nom. superfl. TYPE: A renaming of C.
pteridoides, and so based on the type of that name.

Plants aquatic (usually floating), at 0 - 100 m elevation, in streams and ponds,
from near S. Bernardino (Pcia. Guanacaste), the vicinity of Nievecita (Pcia. Bocas
del Toro), Madden Lake (Pcia. Colon), the Canal Zone, and the lower Rio
Truando. Also from Florida, Louisiana, (Tuba, Hispaniola, Guatemala, El
Salvador, Nicaragua, Trinidad, Venezuela to Peru, the Guianas, Brazil, Paraguay,
and Argentina.

136. Ceratopteris pteridoides (Hook.) Hieron. x richardii Brongn.

Plants aquatic (rooted or floating), at 0-200(600) m elevation, in streams and
ponds, from Salinas (Pcia. Puntarenas), Bagaces and north of Upala (Pcia.
Guanacaste), the IICA lake near Turrialba (Pcia. Cartago), the vicinity of Ft.

108 PARKERIACEAE: 16. CERATOPTERIS

Sherman (Canal Zone), and Matias Hernandez and the vicinity of El Llano (Pcia.
Panama). Also from Mexico and El Salvador.

137. Ceratopteris thalictroides (L.) Brongn. Bull. Sci. Soc. Philom. Paris III,
8:186. 1821.

Acrostichum thalictroides L. Sp. PI. 2:1070. 1753. TYPE: Ceylon, Hermann (BM-Hb. Hermann
III, 42, specimen 377, not seen), according to Lloyd (Brittonia 26:148. 1974).

Ellobocarpus oleraceus Kaulf. Enum. Fil. 148. 1824, nom. superfl. TYPE: A renaming of C.
thalictroides (L.) Brongn., and so based on the type of that name.

Ceratopteris froesii Brade, Arq. Jard. Bot. Rio de Janeiro 18:31, t. 1, f. 4. 1964. TYPE: Campos
Gerais de Andorinha, Est. Par^ Brazil, Froes 19906 (IAN not seen).

Plants aquatic (usually rooted), at 0 - 200 m elevation, in streams and ponds,
from Matapolo and the Hacienda Taboga (Pcia. Guanacaste), Las Sabanas (Canal
Zone), the vicinity of Old (Pcia. Cocld), and between Panama and Chepo (Pcia.
Panama). Also from Florida, Jamaica, Mexico, Guatemala, El Salvador,
Nicaragua, Trinidad, Venezuela, Ecuador, Surinam, and Brazil.

ADIANTACEAE: 17. LLAVEA 109

ADIANTACEAE

Rhizomes usually erect or short-creeping, less commonly long-creeping, rarely
glabrous or merely viscid, usually scaly, the scales concolorous or bicolorous, not
clathrate, sometimes reduced to bristles, or hairy, the hairs firm to lax, usually
brown or reddish-brown, rarely stramineous; fronds small to medium-sized,
usually stiffly erect, sometimes lax and pendent; stipes terete or sometimes sulcate,
usually brown or atropurpureous, variously scaly, soft-hairy, viscid, or glabrous, not
articulate to the rhizome; laminae usually less than 0.5 m long, simple and entire
or lobed or up to 4-pimiate; ultimate segments either roundish and often
somewhat contracted and bead-like or angular and sometimes also dimidiate and
not bead-like, always many-veined; sori marginal or nearly so or distinctly surficial
or confined to the false indusium; false indusia absent or thin, usually entire, pale
brown, a reflexed and often modified segment margin; sporangia usually
numerous, sometimes few per soms, short-stalked, the stalk usually of 3 rows of
ceils, the thickened cells of the vertical annulus sharply distinct from the thin-
walled cells or not; spores trilete.

1. Fronds or pinnae monomorpMc or slightly dimorphic (the sterile and fertile differing slightly in
shape). .3.

1. Fronds or pinnae strongly dimorphic (the sterile and fertile differing strongly in shape or in
stature).. 2.

2(1). Laminae pentagonal, dimorphic, the fertile fronds much longer than the sterile ones.

27. Hemionitis

2(1). Laminae ovate, the pinnae dimoprhic, the proHmal ones sterile and with triangular-
lanceolate segments, the distal ones fertile and with nearly linear segments.

17. Llavea

3(1). Sporangia confined to the abaHal surface of the fake indusia; ultimate segments commonly
oblong and dimidiate or trapezoidal to reniform and not dimidiate.

32. Adimtum

3(1). Sporan^a marginal to surficial, never on the false indusia.. 4.

4(3). Laminae linear, indeterminate, the pinnae approximate, more or less round, less than 5 mm
in diameter, the rachises usually hairy, especially at the apex of the lamina.

3Q. Jamesonia

4(3). Laminae not linear, determinate, the pinnae not round.. 5.

5(4). Sori exindusiate, surficial along the veins, sometimes protected by hairs, scales, or a wax-like
farina.. 11.

5(4). Sori protected by a false indusium, marginal or nearly so, hairs and scales few or absent,
farina absent (except mAleuritopteris)..6.

6(5). Laminae bearing copious, white to yellowish farina on the abaxial surface.

19. Aleuritopteris

6(5). Laminae not farinose, sometimes glaucous..?.

7(6). Laminae palmate, the nearly equal pinnae aU radiating from the apex of the stipe; sori short,
discontinuous.

l^.Adiantopsis

7(6). Laminae pinnate, with at least two pairs of lateral lobes or pinnae, the basal lobes or pinnae
sometimes basiscopically lobed..8.

8(7). Adarial surface of the stipes and rachises bearing abundant cylindrical to slightly clavate,
short hairs.

21. Mildeila

8(7). Adajdal surface of the stipes and rachises glabrous, scaly, or hairy, not bearing abundant
cylindrical to slightly clavate, short hairs.. 9.

110 ADIANTACEAE: 17. LLAVEA

9(8). Laminae pentagonal, pinnatifid or bipinnatifid, at least the basal lobes basiscopically lobed.

24. Doryopteris

9(8). Laminae not pentagonal, pinnate or more divided, the basal pinnae basiscopically lobed or
not..l().

10(9). Stipes dark reddish-brown, dark brown, or blackish.

18. Cheilanthes

10(9). Stipes stramineous to pale brown (except atropurpureous in P. temifolia, with mostly ternate
pinnae and linear segments).

23. Pellaea

11(5). Fronds glabrous, up to ca. 15 cm long; rhizomes rudimentary, the plants annual.

25. Anogramma

11(5). Fronds usually hairy, scaly, or with a wax-like farina, rarely glabrous, mostly more than 15
cm long; rhizomes well developed, the plants perennial.. 12.

12(11). Laminae pentagonal, sparsely pilose on both surfaces, lacking a wax-like farina on the
abaxial surface.

29. Bommeria

12(11). Laminae not as above..l3.

13(12). Laminae pinnate, the pinnae oblong or lanceolate, entire.

28. Gymnopteris

13(12). Laminae not as above..l4.

14(13). Fronds borne in a line along a usually long-creeping rhizome; laminae sparsely pilose
abaxially and often adaxially.

31. Eriosorus

14(13). Fronds fasiculate or caespitose from a short-creeping, multicipital, ascending, or erect
rhizome; laminae scaly, bearing a wax-like farina, or densely hairy on the abaxial surface, the adaxial
surface often glabrous..l5.

15(14). Laminae long-triangular or nearly so, widest near the base.

26. Pityrogramma

15(14). Laminae oblong, narrowly elliptical, or pentagonal (triangular only in N. incana, with
ovate, plane, short-stalked ultimate segments), widest near the middle of the lamina.

22. Notholaena

17. LLAVEA Lag.

Plants terrestrial or sometimes epipetric or in soil over rocks; rhizomes thick,
short-creeping, densely scaly, the scales linear-lanceate, black at the base, reddish-
brown changing to stramineous toward the apex, the margins entire; fronds large,
membranaceous, strongly dimorphic, essentially glabrous; stipes and rachises
stramineous or sometimes pale reddish-brown, deeply sulcate; sterile fronds 2-4-
pinnate, the pinnae and pinnules long-stalked, the segments mostly ovate, the
veins mostly 2-forked, the margins thickened and closely and irregularly spinulose;
fertile fronds 2- or 3-pinnate, the pinnae long-stalked, the segments linear,
membranaceous, sparsely pilosulous along their costules, the margins revolute
with a continuous false indusium; veins 1-forked; sori elongate along the lateral
veins of the segments; sporangia short-stalked.

A morphologically isolated, monotypic genus of Central America.

137a. Llavea cordifolia Lag. Gen. Sp. PI. 33. 1816.

TYPE: “Nova Hispania,” collector unknown (probably MA or FI not seen).

Plants terrestrial, at 1800 m elevation, in the Flora area known only from Alto
de Morizal, La Palma {Gomez 3112, CR). Also from Mexico and Guatemala.

ADIANTACEAE: 18. CHEILANTHES 111

The divided laminae and pale stipes are reminiscent of a few species of Pellaea,
but the extremely dimorphic pinnae show that there is no relationship.

18. CHEILANTHES Swartz

Plants terrestrial or epipetric, often in seasonally dry, open areas; rhizomes
ascending or short-creeping, rarely long-creeping, densely scaly, the scales
narrowly lanceate, often bicolorous, tan with a dark brown central stripe, the
margins entire, erose, ciliate or toothed; fronds small, scaly (glandular or hairy in a
few species); stipes usually brown, reddish-brown, or atropurpureous, often shiny,
scaly especially and often exclusively at the base, terete or subsulcate; rachises
resembling the stipes in color and indument; laminae pinnate-pinnatifid to 5-
pinnate (mostly 2-3-pinnate), commonly lanceolate, rather coriaceous, often at
least slightly hairy and/or scaly, the basal pinnae often inequilateral and
basiscopically produced, the ultimate segments small, often bead-like, rarely
linear, with usually strongly revolute margins and continuous, narrow to wide,
membranaceous, marginal false indusia; veins free, simple or forked, hidden
within the lamina tissue; sori usually round but sometimes laterally confluent, at
the tips of veins; sporangia short-stalked, often few per sorus.

This morphologically diverse genus of tropical to temperate xerophytes is found
in both hemispheres. Some of the ca. 170 species probably can be placed in
segregate genera. The taxonomic value of some traditionally generic characters is
in doubt because of convergence due to harsh habitats.

KNOBLOCH, I, 1976. Morpholo,gical characters in Cheilanthes together with a key to North and
Central American species. Flora 165:507 ” 522.

1. Laminae bearing viscid glands; stipes at least 3 times longer than the laminae. Laminae
compact, ovate-triangular to subpentagonal; fronds cm long.

143. C. kaulfussii

1. Laminae lacking \iscid glands; stipes usually less than 3 times longer than the laminae..!.

2(1). Stipes, rachises, and costae glabrous or nearly so, not villous, or only the stipe bases scaly..5.

2(1). Stipes, rachises, and costae villous, with many reddish or tan hairs and some wide or narrow,
hair-like scales..3.

3(2). Laminae predominantly 2-pimiate; ultimate segments oblong, ca. 3 mm long, 1.5 mm wide;
rhizomes compact, the stipes approximate.

147. C notholaenoides

3(2). Laminae predominantly 3-pinnate; ultimate segments bead-like, ca. 1 mm long and wide.. 4.

4(3). Rhizomes long-creeping, the stipes distant; laminae lanceolate, widest near the base.

144. C lendigera

4(3). Rhizomes short-creeping, the stipes approranate; laminae nearly linear, usually widest near
the middle.

139. C castmea

5(2). Indusia entire, erose, or incised, not decurrent on the segment axes (except in C.
membranacea)..! .

5(2). Indusia ciliate or toothed, slightly to greatly decurrent on the segment axes; segments obtuse
or roimd..6.

6(5). Laminae about twice as long as wide; segment margins often sparingly provided with stubby,
conical trichomes; ultimate segments undi\4ded or with elongate lateral segments; terminal segments
4-20 times longer than wide; pinnae distant.

142. C hirsuta

112 ADIANTACEAE: 18. CHEILANTHES

6(5). Laminae about as long as wide; segment margins rarely provided with trichomes; ultimate
segments mostly with round or only shghtly elongate lateral segments; terminal segments typically
2-3(4) times longer than wide; pinnae usually crowded or imbricate.

145. C. mar^nata

7(5). Segments acute to apiculate at the apex..9.

7(5). Segments round at the apex. .8.

8(7). Laminae 4-pinnate; terminal pinna and pinnule segments not much longer than the lateral
ones; indusia strongly decurrent on the segment margins, (0.5)1 mm wide.

146. C. memhranacea

8(7). Laminae 3-pinnate or 3-pinnate-pinnatifid; terminal pinna and pinnule segments much longer
than the lateral ones; indusia scarcely decurrent on the segment margins, 0.5 mm wide.

141. C hanisii

9(7). Laminae 3-pinnate or less divided (the basal basiscopic pinnules of the basal pinnae pinnate).
Fronds 15 - 20(30) cm long, narrowly lanceolate, 2-4 times longer than wide.

138. C. angustifolia

9(7). Laminae 3-pinnate-pinnatifid or more divided (the basal basiscopic pinnule of the basal
pinnae pinnate-pinnatifid or 2-pinnate). Fronds 30 - 45 cm long, ovate-lanceolate, ca. 2 times longer
than wide.

140. C. cuneata

138. Cheilanthes angustifolia H.B.K. Nov. Gen. Sp. 1:21 (fol. 17). 1816.

Allosoms angustifolius var. minus Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd.
V, 1:219 (repr. 67). 1849. LECTOTYPE: Near Yavesia, Edo. Oaxaca, Mexico, Liebmann PI. Mex.
2349, FI Mex. 231 (C not seen), chosen by A. R. Smith (FI. Chiapas 2:71. 1981).

TYPE: Joullo, Mexico, 1134 m, Humboldt (B not seen).

Plants terrestrial, at 800 - 1600(2000) m elevation, in forests, often among rocks,
from near S. Ramon and Alajuela (Pcia. Alajuela), the mountains around the
Meseta Central, and the Cordillera de Talamanca to Pcia. Chiriqui. Also from
Mexico to Nicaragua.

139. Cheilanthes castanea Maxon, Proc. Biol. Soc. Wash. 32:111. 1919.

TYPE: 12-14 leagues south of Saltillo, Edo. Coahuila, Mexico, Palmer 1390
(US).

Plants probably terrestrial, at 2000 m elevation, in light forests, in the Flora area
known only from near the Hotel La Georgina, Cordillera de Talamanca, Pcia.
Cartago (Gomez 683, CR). Also from the United States and Mexico.

140. Cheilanthes cuneata Kaulf. ex Link, Hort. Reg. Bot. Berol. 2:40. 1833.

TYPE: From cultivated material; a specimen labelled “Cheilanthes cuneata
Kaulf.” in the hand of Kaulfuss may be the type (B not seen Tryon photo).

Plants terrestrial, at ca. 1600 m elevation, in the Flora area known only from
1-3 km southeast of Jorco, Pcia. S. Jose (Mickel 2408, NY, US). Also from
Mexico and Venezuela.

141. Cheilanthes harrisii Maxon, Contr. U. S. Natl. Herb. 24:51. 1922.

TYPE: Old England, below Cinchona, St. Thomas Parish, Jamaica, 1050 m,
Harris 12902 (US; isotypes BM, U, K none seen, NY).

Plants terrestrial, at 1500-2200 m elevation, often in Oak forests, in the Flora
area known only from the vicinity of Sta. Maria de Dota, Pcia. S. Jose (Standley

ADIANTACEAE: 18. CHEILANTHES 113

41668, US); and from between La Sierra and El Empalme, Pda. S. Jose {Jimenez
M. 2637, CR, F). Also from Cuba and Jamaica.

142. Cheilanthes hirsuta Link, Hort. Reg. Bot. Berol. 2:40. 1833.

lAllosorus ciliatus var. minor K. Presl, Reliq. Haenk. 1:59. 1825. TYPE: Mexico, Haenke (PRC
not seen).

Cheilanthes nifescens Link, Hort. Reg. Bot. Berol. 2:39. 1833. PROBABLY AUTHENTIC
SPECIMENS: From cultivated material in the botanical garden at Berlin (B not seen fragm US; L

not seen photo 755).

Cheilanthes crenulata Link, Hort. Reg. Bot. Berol. 2:41. 1833. PROBABLY AUTHENTIC

SPECIMENS: From cultivated material in the botanical garden at Berlin (B not seen fragm US, L
not seen photo 754).

Allosorus decompositus Martens & Galeotti, Nouv. M6m. Acad. Roy. Sci. Bruxelles 15:48, t. 10, f.
2. 1842. TYPE: Juquila, Edo. Oaxaca, Mexico, 4500 - 5500 ft, Galeotti 6362 (BR not seen).

Cheilanthes pyramidalis Fee, M6m. Soc. Sci. Hist. Nat. Strasbourg 5 [M6m. Foug. 7]: 38, t. 25, f. 3.
1857. SYNTYPES (all from Edo. Mexico, Mexico): Valley of Mexico, Schajfner 88; S. Agostin,
Schajfner 305; Guatimalpan, Las Cruces Mountains, ca. 2700 m, Schaffner304 (all P? none seen).

Pellaea angustifolia var. elongata Rovirosa, Pteridogr. Sur M6xico 130. 1910. TYPE: Mesa de
Coapilla, Edo. Chiapas, Mexico, Rovirosa 1059 (PH not seen fragm US, photo US).

TYPE: From material cultivated in the botanical garden at Berlin, presumably
originally from Mexico. The probable holotype is a specimen, doubtless collected
by Link, bearing the handwritten label “H. b. Link . . . Cheilanthes hirsuta” (B not
seen Tryon photo).

Plants terrestrial or epipetric, at 1500-3000 m elevation, in open areas and light
forest, often on banks, from Volcdn Irazu, Cerro Carpintera, Cerro Tablazo, Jorco
(Pcia. J. Jose), and the Cordillera de Talamanca to Pda. Chiriquf. Also from
Mexico to Honduras and Venezuela and Colombia.

This species is closely related to C. marginata; depauperate specimens are
especially difficult to identify.

143. Cheilanthes kaulfussii Kunze, Linnaea 13:145. 1839.

Cheilanthes viscosa Kaulf. ex Link, Hort. Reg. Bot. Berol. 2:43. 1833, non Carm., 1818, nom. illeg.
PROBABLY AUTHENTIC SPECIMEN: From cultivated material in the botanical garden at
Berlin, originally from Mexico (L not seen photo 753).

Cheilanthes glandulifera Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:258
(repr. 106). 1849. LECTOTYPE: Between Talea and Hacienda de Sta. Gertrudis, Edo. Oaxaca,
Mexico, 4000 - 5000 ft, Liebmann PI. Mex. 2354, the sheet bearing Prantl’s annotation (C not seen;
isolectotype K not seen), chosen by A. R. Smith (FI. Chiapas 2:72. 1981).

TYPE: A renaming of C. viscosa, and so based on the type of that name.

Plants terrestrial or epipetric, at 1000-1400(18(X)) m elevation, on dry banks
and in exposed areas, from the Cordillera Central, Cerro Tablazo, and Jorco
(Pcia. S. Jose). Also from the southwestern United States to Honduras and
Colombia.

144. Cheilanthes lendigera (Cav.) Swartz, Syn. Fil. 128, 328. 1806.

Pteris lendigera Cav. Descr. PI. 268. 1802. SYNTYPES: Ixmiquilpan, Edo. Hidalgo, Mexico, Nee
(MA not seen fragm US); and Guaranda, Pcia. Bolivar, Ecuador, Nee (MA not seen). See
Christensen (Dansk Bot. Ark. 9(3) :23. 1937).

Cheilanthes lanuginosa Martens & Galeotti, Nouv. M6m. Acad. Roy. Sci. Bruxelles 15:75, t. 20, f. 2.
1842. TYPE: Near Moran and R6al del Monte, Edo. Hidalgo, Mexico, 7500 - 8500 ft, Galeotti 6450
(BR not seen photo 5095).

114 ADIANTACEAE: 18. CHEILANTHES

ADIANTACEAE: 18. CHEILANTHES 115

Cheilanthes minor Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:75, t. 21, f. 1.
1842. TYPE: Edge of the Rio Capulalpan, Cordillera Oriental de Oaxaca, Edo. Oaxaca, Mexico,
6500 ■=7500 ft, Galeotti 6464 (BR not seen photo 5101).

Myriopteris marsupianthes Fee, Gen. Fil. [Mem. Foug. 5]:149, t. 12A, f. 1. 1852. TYPE: Mt.
Orizaba, Edo. Veracruz, Mexico, 3400 m. Martens & Galeotti 6256 (BR or P not seen).

Myriopteris villosa Fee, Gen. Fil. [Mem. Foug. 5]:149, t. 28, f. 3. 1852. TYPE: Mexico, 2000 m,
Galeotti 6478 (BR or P not seen).

Cheilanthes frigida Linden ex T. Moore, Gard. Chron. 1857:772. 1857. TYPE: A specimen

labelled “C. frigida Hort. Linden . . . (original).” (K-Hort. Veitch not seen Weatherby photo).

Pomatophytum pocillatum M. E. Jones, Contr. W. Bot. 16:12. 1930. TYPE: Ramsey Canon,
Huachuca Mountains, Cochise County, Arizona, 24690 (POM not seen; isotype US).

Plants epipetric, on stone walls, or terrestrial, at 1100-2300(3400) m elevation,
often in exposed areas, from the Fila de Cedral, Volcan Irazu, Volcan Turrialba,
Cerro Chirripd, and Volcan ChiriquL Also from Hispaniola, the southwestern
United States to Honduras, and Venezuela to Ecuador.

145. Cheilanthes marginata H.B.K. Nov. Gen. Sp. 1:22 (fol. 18), fol. t. 669. 1816.

Allosorus ciliatus K. Presl, Rehq. Haenk. 1:59. 1825, nom. superfl. TYPE: A renaming of C.
marpnata H.B.K., and so based on the type of that name.

TYPE: Near Penipe, Pcia. Chimborazo, Ecuador, 2340 m, Humboldt (B not
seen).

Plants terrestrial or epipetric, at 1500-3400 m elevation, in low forests and
open areas, from the Cordillera Central, Cerro Carpintera, Cerro Tablazo and the
Fila de Cedral, and the Cordillera de Talamanca to Volcdn Chiriqui. Also from
Mexico to Honduras and Venezuela to Bolivia.

This species is similar to C. membranacea but is far more delicate and has less
divided fronds.

146. Cheilanthes membranacea (Davenp.) Maxon, Amer. Fern J. 8:119. 1918.

Pellaea membranacea Davenp. Bot. Gaz. (Crawfordsville) 21:262, t. XVIII, f. 5, 6. 1896. TYPE:
Sierra de S. Felipe, Edo. Oaxaca, Mexico, Pringle 5963 (VT not seen).

Plants terrestrial or epipetric, at 2900-3200 m elevation, from Cerro Vueltas
and Cerro Buenavista (Pcia. S. Jose), and Cerro Chirripo. Also from Mexico and
Guatemala.

This species is similar to C chaerophylla (Martens & Galeotti) Kunze and to C.
marginata but is at least fully 4-pinnate, usually has proportionally longer stipes,
and has entire indusia.

FIGS. 138-150. Cheilanthes, Aleuritopteris, Adiantopsis, and Mildella. FIG. 138. Basal pinna of C
angustifolia, Pahner 177, Mexico. FIG. 139. Median pinna of C castanea, Correll 23285, Mexico.
FIG. 140. Basal pinna of C. cuneata, Mickel 2408. FIG. 141. Basal pinna of C. harrisii, Standley
41668. FIG. 142. Suprabasal pinna of C. hirsuta, Standley 29057, Honduras. FIG. 143. Median pinna
of C. kaulfussii, Brown CR-237. FIG. 144. Median pinna of C. lendigera. Stork 411. FIG. 145.
Median pinna of C. marginata, Brade & Brade (Ros. Fil. Costar. Exs. 115). FIG. 146. Basal pinna of
C. membranacea, Kellerman 5791, Guatemala. FIG. 147. Median pinna of C. notholaenoides, von
Tuerckheim 1166, Guatemala. FIG. 148. Plant of Al. farinosa, Beaman 3952, Guatemala. FIG. 149.
Plant of Ad. radiata, Ebinger 416. FIG. 150. Plant of M. intramarpnalis. Smith 48/133a.

116 ADIANTACEAE: 19. ALEURITOPTERIS

147. Cheilanthes notholaenoides (Desv.) Maxon ex Weath. Contr. Gray Herb.

114234. 1936.

Pteris notholaenoides Desv. Mem. Soc. Linn. Paris 6:299. 1827. TYPE: Hispaniola, collector
unknown (P-Hb. Desv. not seen Tryon photo). Desvaux’s description of the rachis as tomentose
could have come neither from Plumier’s t. 58 in the “Trait6...” or from his description.

Cheilanthes micromera Link, Hort. Reg. Bot. Berol. 2:36. 1833. TYPE: Based on material

cultivated at the botanical garden in Berlin, originally from Mexico. The presumable holotype bears
a label “H. b. Link . . . Cheilanthes micromera LK.” (B-Hb. Link not seen Tryon photo).

Cheilanthes aspidioides F6e, Gen. Fil. [M6m. Foug. 5]:157. 1852. TYPE: San Pedro Nolasco, Edo.
Oaxaca, Mexico, ca. 2600 m, Galeotti 6557 (P or RB not seen).

Plants terrestrial, at 100? and 3000 m elevation, in the Flora area known only
from the Pdramo del Jaboncillo, Buena Vista, Pcia. S. Jose {Gomez 884, CR); and
from the woods of Boquete near David (8 Dec 1909, Helion, GH, US). The latter
locality may be above, and at higher elevation, than David. Also from Texas,
possibly Cuba, Hispaniola, Mexico to Belize, and Venezuela.

19. ALEURITOPTERIS Fee

Plants terrestrial or epipetric, in seasonally dry, open areas; rhizomes erect or
ascending, scaly, the scales linear-lanceate, weakly bicolorous, the margins entire;
fronds medium-sized, largely lacking hairs or scales; stipes brown, reddish-brown,
or atropurpureous, shiny, slightly scaly, especially at the base, terete; rachises
resembling the stipes in color; laminae 1-2-pinnate- pinnatifid, commonly
lanceolate, rather coriaceous, copiously farinose on the abaxial surface, the basal
and suprabasal pinnae strongly basiscopically produced; pinnae lanceate, the
pinnules elongate, lobed, with continuous, wide, membranaceous, brownish
marginal indusia; veins free, simple or forked, hidden within the lamina tissue; sori
laterally confluent, at the tips of veins; sporangia short-stalked.

Tropical and subtropical subxerophytes, all but one exclusively in the Old
World; ca. 15 species. This genus is sometimes considered to be a synonym of
Cheilanthes, and some apparent species of Cheilanthes do resemble species of
Aleuritopteris, but the similarities presumably are due to convergence.

148. Aleuritopteris farinosa (Forssk.) F6e, Gen. Fil. [M6m. Foug. 5]: 153, t. 12B, f.

1. 1852.

Pteris farinosa Forssk. H. Aegypt.-Arab. 187. 1775. TYPE: Yemen, Forssk&l (isotype BM not
seen photo 6695).

Cheilanthes pulverulacea K. Presl, Reliq. Haenk. 1:64. 1825. TYPE: Mexico, Haenke (PRC not
seen).

Aleuritopteris mexicana F6e, Gen. Fil. [M6m. Foug. 5]:154. 1852. TYPE: Llano Verde, Edo.
Oaxaca, Mexico, 2000 m, Galeotti 6551 (presumably P not seen; isotype BR not seen).

Plants terrestrial or epipetric, at 100-2000 m elevation, from Hacienda Palo
Verde, Cerro Guyacdn (Pcia. Guanacaste), Volcan Irazu, and scattered localities
in the Cerro de Talamanca from above Copey de Dota (Pcia. S. Jose) to Cerro
Punta and near David (both Pcia. Chiriquf). Also from Jamaica, Hispaniola,
Mexico, Guatemala, and Peru.

This species, which is fairly common in Mexico, is rare in the Flora area.
Apparently it is becoming more widespread and frequent as drier habitats appear
due to deforestation and climatic change.

ADIANTACEAE: 20. ADIANTOPSIS-21. MILDELLA 117

20. ADIANTOPSIS Fee

Plants terrestrial among rocks or epipetric, usually not in open areas; rhizomes
erect, scaly, the scales narrowly lanceate, weakly bicolorous, paler at the entire
margins; fronds medium-sized, palmately divided, glabrous; stipes blackish to dark
brown, shiny, scaly only at the base, terete; costae like the rachises except
uniformly alate, the alae brown, membranaceous; pinnules dimidiate, oblong,
faintly acroscopically auriculate, the margins slightly revolute; veins free, simple or
forked; sori round, discrete, protected by discrete, scarious, whitish, entire indusia;
sporangia short-stalked, few per sorus.

MAXON, W. R. 1908. The Cuban species of Adiantopsis. Contr. U. S. Natl. Herb. 10:485 - 486.

Tropical subxerophytes, all but one in the New World; ca. 6 species. The
species on Madagascar may be only distantly, if at all, related, and is more likely to
be related to species of Cheilanthes from Africa.

149. Adiantopsis radiata (L.) F^e, Gen. Fil. [M^m. Fong. 5]: 145. 1852.

Adiantum radiatum L. Sp. PI. 2:1094. 1753. LECTOTYPE: A specimen labelled “Adiant. 3
radiatum” in the hand of Linnaeus (LINN 1252.1 not seen microfiche S. I. Library), perhaps
collected by Sloane in Jamaica, chosen by Lellinger (Mem. New York Bot. Card. 23:3. 1972).

Plants terrestrial, at 0-400 m elevation, from the vicinity of Lx)s Chiles (Pcia.
Alajuela), Ancon Hill (Canal Zone), and Cerro Azul (Pcia. Panama). Aso from
Cuba, Jamaica, Hispaniola, the Lesser Antilles, Mexico to Honduras, Nicaragua,
Trinidad, Venezuela, Colombia, Peru, Bolivia, Guyana, Brazil, Paraguay, and
Argentina.

21. MILDELLA Trev.

Plants terrestrial or epipetric, usually on dry banks in open areas; rhizomes
branched, creeping to ascending, the fronds clustered near the rhizome apices,
densely scaly, the scales linear-lanceate, strongly bicolorous to concolorous; fronds
small, glabrous; stipes and rachises chestnut-colored to dark brown, adaxially
grooved, the groove provided with bicellular, short, stiff hairs; laminae pinnate-
pinnatifid to 2-pinnate, long-lanceolate to ovate-triangular, the proximal pinnae
mequilaterally triangular, the distal segments simple, adnate, linear to triangular;
veins free, simple or forked; indusia scarious, vaulted, inframarginal at maturity;
sporangia short-stalked, typically 2 per sorus.

Mostly Old World subtropics of China with extensions to Tibet and India; 8
species. The New World species are more tropical, with M intramarginalis locally
abundant on dryish, roadside banks and perhaps readily cultivated in terraria.
HALL, C. C. & D. B. LELLINGER. 1967. A revision of the fern genus Mildella. Amer. Fern J.

57:113-134.

150. Mildella intramarginalis (Kaulf. ex Link) Trevis. Rendiconti 1st. Lombardo,
Milano II, 9:810. 1876, var. intramarginalis.

Pteris intramarginalis Kaulf. ex Link, Hort. Reg. Bot. Berol. 2:34. 1833, as ‘'inframargfnalis”

LECTOTYPE: Cultivated in the Botanical Garden at Berlin, Link (B), chosen by Hall and Lellinger
(Amer. Fern J. 57: 119-120. 1967).

Plants terrestrial, at 900-2000 m elevation, on banks and roadsides, from the
Cordillera Central, Cerro Tablazo, and the Cordillera de Talamanca to Pcia.
Chiriquf. Aso from Mexico to Nicaragua.

118 ADIANTACEAE: 22. NOTHOLAENA

22. NOTHOLAENA R. Br.

Plants terrestrial or epipetric, often in seasonally dry, open areas; rhizomes
short-creeping to ascending, branched, scaly, the scales concolorous or bicolorous,
lax to stifi, sometimes bearing spine-like teeth; fronds small- to medium-sized,
scaly, hairy, or farinose; stipes terete, scaly at least at the base, usually reddish-
brown to dark brown; rachises resembling the stipes in color and mdument;
laminae mostly pinnate-pinnatifid to 3-pinnate, commonly oblong, sometimes
ovate or pentagonal, rather coriaceous, the pinnae entire to pinnatifid, pinnate, or
more divided, commonly scaly or farinose on the abaxial surface, the margins
usually plane or slightly revolute; veins free, simple or forked, hidden within the
lamina tissue; sori submarginal, commonly oblong; sporangia short-stalked, many
per sorus.

Tropical and subtropical (rarely temperate) xerophytes, mostly New World; ca.
35 species, but properly subject to division.

TRYON, R. 1956. A revision of the American species of Notholaena. Contr. Gray Herb.

179:1-106.

1. Laminae scaly, not farinose.. 5.

1. Laminae farinose, the farina white, cream, or yellow, not scaly..2.

2(1). Farina white..4.

2(1). Farina yellow or yellowish..3.

3(2). Laminae subpentagonal, 1-2 times longer than wide. Laminae 3-7 cm long, 2-6 cm wide,
2-pinnate-pinnatifid, the basal basiscopic pinnules of the basal piimae greatly elongate.

158. N. sulphurea

3(2). Laminae linear, 5-20 times longer than wide. Laminae 5-15 cm long, 1- 1.5(2) cm wide,
pinnate-pinnatifid, the basal basiscopic pinnules of the basal pinnae not elongate.

151. N. affinis

4(2). Laminae 3-pinnate; ultimate segments ovate, short-stalked; rhizome scales concolorous; stipe
scales few, only at the base. Laminae 5-15 cm long, 1.5-6 cm wide, the basal basiscopic pinnules
not elongate.

156. N. incana

4(2). Laminae 2-pinnate-pinnatifid; ultimate segments oblong, sessile, broadly attached at the
base; rhizome scales bicolorous; stipe scales many, scattered along the stipe. Laminae 5- 15 cm long,
2-7 cm wide, the basal basiscopic pinnules greatly elongate.

154. N. Candida var. Candida

5(1). Laminae lanceolate to oblanceolate, widest at or distal to the middle; fronds nearly
exstipitate; all pinnae sessile. Laminae 7-32 cm long, 3-6 cm wide, pinnate-pinnatifid; stipe base
and rhizome scales dark brown.

153. N. brachypus

5(1). Laminae linear; fronds decidedly stipitate; at least the lower pinnae not sessile..6.

6(5). Pinnae pinnatisect, bipinnatifid, or pinnate-pinnatifid; stipe base scales blackish. Laminae
8-25 cm long, 1.5-4 cm wide.

155. N. galeottii

6(5). Pinnae lobed to pinnatifid; stipe base scales not blackish..7.

7(6). Rhizome scales concolorous, reddish at the margins, dark brown at the center; scales of the
abaxial surface of the pinnae cream-colored in mass. Laminae 15-30 cm long, 1-2.5 cm wide.

152. N. aurea

7(6). Rhizome scales bicolorous, reddish to nearly stramineous; scales of the abaxial surface of the
pinnae brown or reddish-brown in mass. Laminae 20-30(60) cm long, 2-4 cm wide.

157. N. sinuata var. sinuata

ADIANTACEAE: 22. NOTHOLAENA 119

151. Notholaena aMnis (Mett.) T. Moore, Ind. Fil. 233. 1861.

Cheilanthes affinis Mett. Abh. Senckenberg. Naturf. Ges. 3:63. 1859. TYPE: Mexico, Aschenbom
(B not seen photo GH not seen).

Plants epipetric, at 0-100 m elevation, in dry to moist forest areas on peridotite
cliffs, from along the Pacific coast from Playa Panama near Playa del Coco to
Punta Guiones (both Pcia. Guanacaste). Also from Mexico to Honduras and
Nicaragua.

152. Notholaena aurea (Poir.) Desv. M6m. Soc. Linn. Paris 6:219. 1827.

Pteris aurea Poir. Encyc. Meth. 5:710. 1804. TYPE: Peru, /. de Jussieu (P-Hb. Juss. 1333 not seen
photo 3098).

Acrostichum bonariense Willd. Sp. PI. ed. 4, 5:114. 1810. TYPE LOCALITY: “Bonaria” [Buenos
Aires, Argentina, but perhaps actually from Mexico] (B not seen photo GH).

Cheilanthes ferru^nea Willd. ex Link, Enum. Hort. Berol. Alt. 2:463. 1822. TYPE LOCALITY:
“America meridionali” (B not seen photo GH).

Notholaena chiapensis Rovirosa, Pteridogr. Sur Mexico 229, t. 48, f. 1-6. 1910. TYPE: Near El
Sacramento, Edo. Chiapas, Mexico, 2000 m, Rovirosa 1077 (PH not seen fragm and photo US;
isotype GH).

Plants epipetric, at 1200 m elevation, in the Flora area known only from Volcan
Tenorio, Pcia. Guanacaste (Gomez 1050, CR) and from an unlocalized collection
from Costa Rica (Erode 136, G not seen, P not seen). Also from the southwestern
United States, Jamaica, Hispaniola, Mexico to Honduras, and Colombia to
Argentina and Chile.

153. Notholaena brachypus (Kunze) J. Smith, Ferns Brit. For. 172. 1866.

Cheilanthes squamosa var. brachypus Kunze, Linnaea 18:340. 1845. TYPE: Mexico, Leibold 52 (B
not seen photo GH; isotype K not seen photo GH).

Plants terrestrial, at 0-1000 m elevation, in moist to dry forests and open areas,
from Pcia. Guanacaste and the lower reaches of the Meseta Central. Also from
Mexico to Nicaragua.

154. Notholaena Candida (Martens & Galeotti) Hook. Sp. Fil. 5:110. 1864.

Cheilanthes Candida Martens & Galeotti, Nouv. M6m. Acad. Roy. Sci. Bruxelles 15:73, t. 20, f. 1.
1842. TYPE: Guadalajara, Edo. Jalisco, Mexico, Galeotti 6442 (BR not seen photo 5092).

Ceropteris monosticha Fee, M6m. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 7]:44, t. 22, f. 2.
1857. TYPE: Near Orizaba, Edo. Veracruz, Mexico, Schaffner 155 (P or RB not seen).

Cheilanthes furfuracea K. Presl ex W. Koch, Wochenschr. G^tnerei Pflanzenk. 1:3. 1858, nom.
illeg. TYPE: A renaming of C. Candida Martens & Galeotti, and so based on the type of that name.

Notholaena sulphurea var. alba Mett. ex Kuhn, Abh. Naturf. Ges. Halle 11:32. 1869. TYPE
LOCALITY: Mexico.

Plants epipetric, at 700 m elevation, in the Flora area known only from the
Brasil hydroelectric plant, Sta. Ana, Pcia. S. Jose (Gomez 7193 and 7243, both
CR). Also from Mexico, Guatemala, Honduras, and Nicaragua.

155. Notholaena galeottii F6e, Gen. Fil. [M6m. Foug. 5]: 159. 1852.

Notholaena arsenii Christ, Notul. Syst. (Paris) 1:232. 1910. LECTOTYPE: Tetela, Edo. Puebla,
Me?dco, 2170 m. Arsine 2046 (P not seen), chosen by Tryon (Contr. Gray Herb. 179:55. 1956).

Notholaena hyalina Maxon, Amer. Fern J. 5:4. 1915. TYPE: S. Jos6 Pass, Edo. S. Luis Potosi,
Mexico, Pringle 3297 (US).

120 ADIANTACEAE: 22. NOTHOLAENA

ADIANTACEAE: 22. NOTHOLAENA 121

TYPE: Caputalpan, Edo. Oaxaca, Mexico, 3000 m, Galeotti 6565 (P? not seen;
isotypes BM not seen photo 7526, BR not seen, K not seen).

Plants epipetric, at 3000 m elevation, in the Flora area known only from Volcan
Turrialba (Gdmez 899, CR). Also from southern Mexico and Guatemala.

156. Notholaena incana K. Presl, Reliq. Haenk. 1:19, 1. 1, f. 2. 1825.

Gymnogramma Candida Mett. Abh. Senckenberg. Naturf. Ges. 3:50. 1859. LECTOTYPE: Valle
de Mexico, Mexico, Schmitz 231 (B not seen Weatherby photo), chosen by Maxon and Weatherby
(Contr. Gray Herb. 127:5. 1939).

LECTOTYPE: Mexico, Haenke (PRC not seen photo GH not seen), chosen by
Maxon and Weatherby (Contr. Gray Herb. 127:5, 7. 1939).

Plants epipetric, at 1400-1500 m elevation, in the Flora area known only from
near Prusia de Cartago, Pcia. Cartago {Gdmez 7156, CR). Also from Hispaniola,
Mexico, and Guatemala.

157. Notholaena sinuata (Swartz) Kaulf. Enum. Fil. 135. 1824, var. sinuata.

Acrostichum sinuatum Lag. ex Swartz, Syn. Fil. 14. 1806. TYPE LOCALITY: “Peru.” If the type
specimen was collected by Lagasca, the locahty must surely be Mexico. Lagasca’s original herbarium
was destroyed, but type material may be in FI, MA, MEXU, or S.

Notholaena tectaria Desv. Mem. Soc. Linn. Paris 6:219. 1827. TYPE: Peru, collector unknown (P
not seen photo GH).

Notholaena chalcolepis A. Braun ex Kunze, Linnaea 13:135. 1839. TYPE: Mexico, Karwinsky in
1827 (B not seen photo GH; isotype P not seen photo GH).

Notholaena laevis Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:46. 1842. TYPE:
South of Sola, Edo. Oaxaca, Mexico, Galeotti 6350 (BR not seen; isotypes K, B neither seen).

Notholaena sinuata var. integra Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd.
V, 1:213 (repr. 61). 1849. TYPE: A renaming of N. laevis Martens & Galeotti, and so based on the
type of that name.

Notholaena crassifolia T. Moore & Houlst. Gard. Mag. Bot. 3:20. 1851. TYPE: From cultivated

material, Henderson (not seen).

Notholaena pruinosa Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:78. 1857.

TYPE: Valleys of Toluca and Orizaba, Mexico, Schaffher 167c (P or RB not seen).

Notholaena deltoidea Baker, Syn. Fil. ed. 2, 514. 1874. TYPE: Mexico, Leibold (K not seen photo
GH).

Notholaena sinuata var. pinnatifida Farw. Amer. Midi. Naturahst 12:283. 1931. TYPE: Bear
Mountains, New Mexico, Rusby B1 (MICH not seen).

Plants terrestrial or epipetric, up to 2000 m elevation, from Volcan Poas and the
vicinity of Hotel Robert on Volcan Irazu, and from two unlocalized specimens
collected in Costa Rica. Also from Hispaniola, the southwestern United States to
Honduras, Nicaragua, and Venezuela and Colombia to Argentina.

158. Notholaena sulphurea (Cav.) J. Smith in Seem. Bot. Voy. Herald 1:233.
1854.

Pteris sulphurea Cav. Descr. PI. 269. 1802. TYPE: Chimapan [Zimapan], Edo. Hidalgo, Mexico,
Nee (MA not seen; isotypes F, G not seen).

FIGS. 151-158. Notholaena. FIG. 151. Frond of N. affinis, St2mdley 74006, Guatemala. FIG. 152.
Frond of N. aurea, Tonduz 756, Guatemala. FIG. 153. Frond of N brachypus, Jimenez L. 347. FIG.
154. Frond of N Candida, Maxon 3368, Guatemala. FIG. 155. Frond of N. galeottii, Pringle 3297,
Mexico. FIG. 156. Plant of N. incana. Hatch & Wilson 335, Guatemala. FIG. 157. Frond of N.
sinuata var. sinuata, Williams et al. 21855, Guatemala. FIG. 158. Frond of N sulphurea, Copeland,
Mexico.

122 ADIANTACEAE: 23. PELLAEA

Notholaena cretacea Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:216
(repr. 64). 1849. TYPE: S. Lorenzo, Tehuac^, Edo. Puebla, Mexico, Liebmann (C not seen;
isotypes K not seen, P not seen, US).

Cheilanthes borsi^ana Reichenb. f. & Warsz. ex W. Koch, Wochenschr. G^tnerei Pflanzenk. 1:2.
1858. TYPE: Peru, Warszewicz (B not seen Weatherby photo).

Notholaena Candida var. lutea Hook. Sp. Fil. 5:111. 1864. TYPE: Depto. Hu^uco, Peru,
Mathews 981 (K not seen).

Cheilanthes lepida Phil. Anales Univ. Chile 94:355. 1896. TYPE: Copiapo, Pcia. Atacama, Chile,
Geisse (SGO not seen photo GH not seen).

Plants terrestrial or epipetric, at 100-3000 m elevation, in the Flora area known
only from Volcan Turrialba {Gomez 900, CR), the Pacific side of Aguacate, Pcia.
Puntarenas {Gomez 1082, CR), and Panama City, Pcia. Panama {Seemann in 1847,
BM not seen). Also from Mexico, Guatemala, Nicaragua, Colombia, Peru, and
Chile.

23. PELLAEA Link

Plants epipetric or terrestrial, often on cliffs or boulders in pastures or open
areas; rhizomes creeping, ascending, or erect, scaly, the scales very narrow,
concolorous or bicolorous, tan and sometimes with a blackish central stripe, the
margins entire to erose; fronds small- to medium-sized, stipes and rachises pale or
dark, even atropurpureous, terete or adaxially sulcate; laminae l-3(4)-pinnate,
firmly herbaceous to coriaceous, usually glabrous, sometimes glaucous, deltate-
lanceate to oblong, the ultimate segments in most species elliptical or nearly so,
with revolute margins; veins free, usually forked, hidden within the lamina tissue
in most species; sori round to elongate, borne at the apex of the veins, often
laterally confluent; sporangia short-stalked, many per sorus.

Tropical and subtropical, mostly New World; ca. 80 species.

TRYON, A. F. 1957. A revision of the fern genus Pellaea section Pellaea. Ann. Missouri Bot. Gard.

44:125-193.

1. Veins visible on both abaxial and adaxial surfaces of the laminae; ultimate segments broadly
attached to the costae or costules; laminae herbaceous. Laminae lanceolate to deltate-lanceolate,
8-20 cm long, 5-18 cm wide, 2-3-pinnate-pinnatifid; axes glabrous or with a few pale, narrow, hair-
like scales; ultimate segments lanceolate to oblong.

160. P. skinneri

1. Veins not visible on either surface of the laminae; ultimate segments stalked or sessile, narrowly
attached to the costae or costules; laminae coriaceous.. 2.

2(1). Laminae oblong; pinnae ternate; stipes atropurpureous, glabrous. Laminae 7-22 cm long,
1.5 - 3.5 cm wide; ultimate segments oblong to linear.

161. P. temifolia

2(1). Laminae lanceate; pinnae pinnately divided; stipes stramineous, densely pilosulous. Laminae
30 - 60 cm long, 10-15 cm wide; rachises, costae, and longer costules flexuous; ultimate segments
ovate.

159. P. ovata

159. Pellaea ovata (Desv.) Weath. Contr. Gray Herb. 114:34. 1936.

Pteris ovata Desv. M6m. Soc. Liim. Paris 6:301. 1827. TYPE: Peru, collector unknown (P-Hb.
Desv. not seen), examined by Weatherby (Contr. Gray Herb. 114:34. 1936).

Pteris flexuosa Kaulf. ex Schlechtend. & Cham. Linnaea 5:614. 1830. TYPE: Near Jalapa, Edo.
Veracruz, Mexico, Aug 1828, Schiede & Deppe (B not seen).

ADIANTACEAE: 24. DORYOPTERIS 123

Plants epipetric or occasionally terrestrial, at 1400-2000(3100) m elevation, in
open and lightly forested areas, often on slopes and banks, from the Cordillera
Central and the Fila de Cedral. Also from the southwestern United States,
Hispaniola, Mexico to Honduras, Nicaragua, Venezuela to Bolivia, and Argentina.

160. Pellaea skinneri Hook. Sp. Fil. 2:141, t. 118B. 1858.

Pellaea flavescens Fourn. Mexic. PI. 1:119. 1872, nom. illeg., non Fee, 1869. TYPE: Cuernavaca,
Edo. Morelos, Mexico, Bourgeau (P not seen).

TYPE: Guatemala, Skinner (K not seen Weatherby photo).

Plants terrestrial, at 0-100(600) m elevation, from Palmira de Guanacaste and
Playa Hermosa (both Pcia. Guanacaste), and Pte. de la Garita (Pda. Alajuela).
Also from Mexico, Guatemala, El Salvador, Nicaragua, and Colombia.

161. Pellaea temifolia (Cav.) Link, Fil. Sp. 59. 1841.

Pteris temifolia Cav. Descr. PI. 266. 1802. LECTOTYPE: Guarimaya Valley near Guamantanga,
Depto. Lima, Peru, Nee (MA not seen; isolectotype F not seen), chosen by Christensen (Dansk Bot.
Ark.9(3):22. 1937).

Pteris pemviana Poir. Encyc. M6th. 5:718. 1804. TYPE: Peru, Jussieu (P not seen photo 3099;
isotype B-Hb. Willd. 19980 not seen microfiche S. I. Library).

Pteris subverticillata Swartz, Syn. Fil. 103. 1806, nom. superfl. TYPE: A renaming of P. temifolia,
and so based on the type of that name.

Pteris triphylla Bertero ex CoUa, Herb. Pedem. 6:199. 1836. TYPE: Chile, Bertero in 1828 (TO not
seen; isotypes GH not seen, SGO not seen).

Pellaea weddelliana Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 81:74. 1857.

LECTOTYPE: “Province de Tomina,” Bolivia, Weddell 3778 (K not seen; isolectotypes P not seen,
YU not seen), chosen by A. F. Tryon (Arm. Missouri Bot. Gard. 44:150. 1957).

Adiantum tematum Sess6 & Moc. PI. Nov. Hisp. 182. 1890; ed. 2. 169. 1893, nom. illeg., non
Humb. & Bonpl. ex Willd., 1810. TYPE: Near Sta. Rosa, Edo. Guanajuato, Mexico, Sesse & Mocino
(MA not seen).

Pellaea temifolia var. petiolulata C. Chr. in Aspl. Ark. Bot. 20A(7): 18. 1926. TYPE: Chiguana,
Depto. Potosi, Bolivia, 4000 m,Asplund 3071 UPS: isotype US).

Pellaea brandegeei C. C. Hall, Amer. Fern J. 37:111. 1947. TYPE: Cape region, Edo. Baja
California Sur, Mexico, Brandegee in 1893 (UC).

Plants epipetric, at 1500-2000 m elevation, in the Flora area known only from
Prusia de Cartago, Pcia. Cartago {Gomez 7139, CR) and from Pico Blanco, Pcia. S.
Jose {Gomez 882, CR). Also from the southwestern United States, Hispaniola,
Mexico to Honduras, Nicaragua, Venezuela to Bolivia, Argentina, and Chile.

24. DORYOPTERIS J. Smith

Plants terrestrial in forests and on roadside banks; rhizomes short-creeping to
erect, scaly, the scales narrowly lanceate, usually bicolorous, tan with a narrow,
blackish central stripe, the margins entire; fronds medium-sized, often slightly
dimorphic with the fertile monds taller than the sterile ones; stipes
atropurpureous, shiny, much longer than the laminae, usually terete, mostly
glabrous; laminae pentagonal, broadly debate, the basal pinnae basiscopically
lobed or produced, herbaceous, glabrous, with revolute margins and continuous,
narrow, marginal indusia; veins anastomosing without included veinlets or free in
some species; sori laterally confluent, continuous around the frond margin;
sporangia long-stalked.

Tropics and subtropics, mostly New World; ca. 35 species.

TRYON, R. M., Jr. 1942. A revision of the genus Doryopteris. Contr. Gray Herb. 143:1-80.

124 ADIANTACEAE: 24. DORYOPTERIS

ADIANTACEAE: 25. ANOGRAMMA 125

162. Doiyopteris pedata van palmata (Willd.) Hicken, Revista Mus. La Plata
15:253. 1908.

Pteris palmata Willd. Sp. PI. ed. 4, 5:357. 1810. TYPE: Caracas, Distr. Fed., Venezuela,

Bredemeyer (B-Hb. Willd. 19957 not seen Tryon photo).

Pteris pedata var. gemmipara Sodiro, Anales Univ. Quito 8(55): 68 (repr. 99). 1893. TYPE: Rio
Guallabamba near Los Reales, Pcia. Pichincha, Ecuador, 1600-1800 m, Sodiro (Hb. Sodiro not

seen).

Doryopteris mayoris Rosenst. M6m. Soc. Sci. Nat. Neuchatel 5:51, t. 2, f. 2. 1912. TYPE:

Medellm, Depto. Antioquia, Colombia, Mayor 180 (S not seen; isotype US).

Plants terrestrial, at 100-1900 m elevation, in forests and on roadside banks,
from the upper Rio Reventazon valley (Pcia. Cartago), Cerro Tablazo, the vicinity
of S. Isidro del General, above Cahazas and La Yeguada (Pcia. Veraguas), and
Barro Colorado Island (Canal Zone). Also from Mexico, Nicaragua, Venezuela,
and Colombia to Bolivia.

25. ANOGRAMMA Link

Plants terrestrial or epipetric, often on banks among mosses or on wet rocks in
or near streams and waterfalls; rhizomes small or obsolete, erect, hairy and/or
scaly, the trichomes tan to brown; fronds small to minute, stipes thin, darkened at
the base, sparsely scaly; laminae 3-pinnate, oblong-lanceolate, membranaceous,
glabrous, rather finely divided, the ultimate segments narrowly oblanceolate to
linear, sometimes cleft at the apex, plane; veins free, simple, obvious; sori
elongate, lateral on the veins near the apex; sporangia short-stalked.

Subtropical to tropical regions of both hemispheres, but absent from North
America north of Mexico; 6 species.

TRYON, R. 1%2. Taxonomic fern notes. 11. Pityrogramma (including Trismeria) and Anogramma.
Contr. Gray Herb. 189:52 - 76.

163. Anogramma leptophylla (L.) Link, Fil. Sp. 137. 1841.

Polypodium leptophyllum L. Sp. PI. 2:1092. 1753. LECTOTYPE: A specimen labelled

“heterophyllum 46” in the hand of Linnaeus, annotated “leptophyllum” by J. E. Smith (LINN
1251.56 not seen microfiche S. I. Library), chosen by Morton (Amer. Fern J. 60:101-103. 1970).

Gymnogramma leptophylla var. mexicana Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci.
Bruxelles 15:27. 1842. TYPE: Mirador, Edo. Veracruz, Mexico, Galeotti 6294 (BR not seen photo
5136).

Gymnogramma chaerophylla var. cryptoff-ammoides Bommer in Dur. & Pitt. Bull. Soc. Roy. Bot.
Belgique 35, Mem. 234. 1896. TYPE: Cerro du Durazno, Irazu Massif, Pcia. Cartago, 2200 m, Pittier
637 (BR not seen; isotype CR).

Pityrogramma guatemalensis Domin, Spisy Pfir. Fak. Karlovy Univ. 88:9. 1928. TYPE: [Volcan]
Fuego, Guatemala, Salvin & Godman 172 (PRC? not seen; isotypes K not seen Tryon photo, US).

FIGS. 159-173. Pellaea, Doryopteris, Anogramma, and Pityrogramma. FIG. 159. Basal pinnules of
median pinna of P. ovata, Valerio 196. FIG. 160. Basal pinnules of median pinna of P. skinned,
Chaves 8, Nicaragua. FIG. 161. Plant of P. temifolia, Howard 82, Nicaragua. FIG. 162. Sterile and
fertile fronds of D. pedata var. palmata, Standley 41475. FIG. 163. Plant of A. leptophylla, Valerio
244. FIG. 165. Median pinna of P, calomelanos var. calomelanos, Skutch 4382. FIG. 166. Frond of
P. chrysoconia, Lehmann 3673, Colombia. FIG. 167. Median pinna of P. dealbata, Elmore H49.
FIG. 168. Median pinnule of P. dukei, Duke 11200. FIG. 170. Median pinna of P. ebenea var. ebenea,
Werckl6. FIG. 171. Subbasal pinna of P. ferrugnea, Maxon 4974. FIG. 172. Pinnule of P. pearcei.
Mayor 144, Colombia. FIG. 173. Median pinna of P. trifoliata, Skutch 2295.

126 ADIANTACEAE: 26. PITYROGAMMA

Plants terrestrial, at 600-2900 m elevation, on banks and slopes, often among
mosses, from the Cordillera Central, the Meseta Central, and the Cordillera de
Talamanca to Pcia. Chiriqui. Also from Mexico, Guatemala, El Salvador,
Nicaragua, and Peru to southern Brazil.

26. PITYROGRAMMA Link

Plants terrestrial, often somewhat weedy, common on roadside banks; rhizomes
erect to ascending, scaly, the scales narrowly lanceolate, concolorous, tan to
brown, the margins entire; fronds medium- or large-sized; stipes brown to
atropu^ureous, shiny, scaly at the base, usually sulcate; rachises resembling the
stipes in color; laminae 1-3-pinnate, usually lanceolate, firmly herbaceous to
subcoriaceous, slightly to more usually copiously farinose on the abaxial surface,
the farina usually white or yellow, or pubescent in a few species; pinnae usually
narrowly lanceate, equilateral (the basal ones inequilateral in a few species), the
ultimate segments triangular to elongate, nearly plane, not revolute; veins free,
forked, hidden within the lamina tissue; indusia absent; sori elongate, irregular,
lateral on the veins near the apex; sporangia short-stalked.

Mostly New World tropics, but widely naturalized in the Old World tropics; ca.
15 species.

TRYON, R. 1962. Taxonomic fern notes. II. Pityrogramma (including Trismeria) and Anogramma.

Contr. Gray Herb.l89;52-76.

1. Pinnae palmate with usually (2)3(7) lobes. Fronds up to 1.5 m long; stipes and rachises
atropurpureous; laminae linear, slightly dimorphic, the sterile pinnules linear, entire, 2-12 cm long,
ca. 1 cm wide, often with a pale yellow farina on the abaxial surface, the fertile pinnules laterally
contracted.

173. P. trifoliata

1. Pinnae pinnate, the lateral lobes or pinnules many..2.

2(1). Rachises and costae densely lanate abaxially, the hairs stramineous, with obvious cross-walls.
Fronds up to 1 m long; stipes and rachises atropurpureous, the surface often hidden by the abundant
hairs; laminae lanceate, up to 25 cm wide, pinnate-pinnatifid to 2-pinnate, the pinnae hnear-
lanceolate, the pinnules or segments up to 1 cm long, 4 mm wide.

171. P. femiginea

2(1). Rachises and costae glabrous or glabrescent at maturity, usually farinose abaxially..3.

3(2). Scales small, generally inconspicuous on the rhizome and stipe bases, their apices long, thin,
one cell wide..6.

3(2). Scales large, conspicuous on the rhizome and stipe bases, several cells wide nearly to then-
apices.. 4.

4(3). Laminae with white farina, coriaceous, 2-pinnate or 2-pinnate-pinnatifid; ultimate segments
wide, with several veins. Fronds 20 - 35 cm long; stipes sulcate, atropurpureous at the base, tan to
stramineous distally; laminae broadly lanceolate, 10-25 cm long, 8-15 cm wide, coriaceous, the
segments often revolute.

166. P. chrysoconia

4(3). Laminae with yeUow farina or not farinose, papyraceous, 3-pinnate or more compound;
ultimate segments narrow, with 1 or 2(3) veins..5.

5(4). Laminae not farinose; ultimate segments with 1 vein. Stipes 10-20 cm long; laminae ovate-
lanceolate to debate, 10-25 cm long, 5-25 cm wide.

172. P. pearcei

5(4). Laminae usually with yellow or white farina; ultimate segments with 1 or 2(3) veins. Stipes
14-31 cm long; laminae debate to nearly triangular, 15-21 cm long, (7)11-20 cm wide.

168. P. dukei

ADIANTACEAE: 26. PITYROGRAMMA 127

6(3). Ultimate segments obovate or oval; small fronds 2-pmnate-pinnatifid; large fronds 3-pinnate
or 3-pinnate-pinnatifid, Stipes 8 - 38 cm long; laminae narrowly lanceate to ovate-lanceolate, 9-33
cm long, 6-20 cm wide.

167. P. dealbata

6(3). Ultimate segments elliptical, lanceolate, lanceate, deltate, or oblong; small fronds pinnate-
pinnatifid to 2-pinnate at the base; large fronds 2-3-pinnate..7.

7(6). Pinnules or ultimate segments and subapical pinnae at right angles to their axes, obtusely
lobed or pinnatifid, obtuse or blunt at the apex; basal and suprabasal pinnae inequilateral,
basiscopically developed. Fronds 10 - 60(75) cm long, 2.5 - 20(30) cm wide, linear to lanceate,
pinnate-pinnatifid to 2-pinnate-pinnatifid..9.

7(6). Pinnules or ultimate segments and subapical pinnae oblique to their axes, serrate or acutely
pinnatifid, acute at the apex; basal and suprabasal pinnae equilateral. Fronds 4-50(75) cm long,
8 - 20(30) cm wide, narrowly to broadly lanceolate, 2-pinnate to 3-pinnate-pinnatifid..8.

8(7). Farina white, pale yellow, or nearly absent.

165. P. calomelanos var. calomelanos

8(7). Fcirina brilliant yellow, rarely orangish.

164. P. calomelanos var, aureoflava

9(7). Farina white.

170. P. ebenea var. ebenea

9(7). Farina brilliant yellow.

169. P. ebenea var. aurata

164. Pityrogramma calomelanos van aureoflava (Hook.) Weath. ex Bailey, Man.

Cult. PI. 64. 1924.

Gymnogramma calomelanos var. aureoflava Hook. Card. Ferns t. 50 text. 1862. LECTOTYPE:
Ecuador, Seeman 948 (K not seen), chosen by Tryon (Contr. Gray Herb. 189:61. 1962).

Pityrogramma austroamericana Domin, Kew Bull. 1929:221. 1929. LECTOTYPE: Challapampa,
Depto. Oruro, Bolivia, 2550 - 27(X) m, Mandon 1549bis (K not seen Tryon photo not seen; isotype
GH not seen).

Pityrogramma hybrida var. broadwayana Domin, Rozpr. Ceske Akad. Ved. Tf. II, Vedy Mat. Pnr.
51(15):3. 1941. TYPE: Cultivated at the St. Clair Experiment Station, Trinidad, Broadway 4678 (US;
probable isotype NY not seen).

Plants terrestrial, at 0-1400 m elevation, along roads and trails, from the
adjacent lowlands, foothills, and slopes of the Cordillera de Tilaran, the Cordillera
Central, and the northern end of the Cordillera de Talamanca. Also from
Honduras, Nicaragua, Venezuela, Ecuador to Bolivia, Brazil, and Argentina.

F. R. Fosberg (pers. comm.) has pointed out that Bailey always suppressed
parenthetical authors in order to avoid imposing on those who had to write out the
names adopted in his book. His citation of “var. aureoflava Weatherby” is not
clearly a transfer of Hooker’s name, but Weatherby gives the intended citation in
full in the Gray Herbarium Card Index.

165. Pityrogramma calomelanos (L.) Link, Handb. 3:20. 1833, var. calomelanos.

Acrostichum calomelanos L. Sp. PI. 2:1072. 1753. TYPE: A specimen, locality and collector
unknown, labelled “24 Acrostic, calomelcmos” in the hand of Linnaeus (LINN 1245.19 not seen
microfiche S. I. Library).

Gymnogramma bidentata K. Presl, Reliq. Haenk. 1:18, t. 2, f. 3. 1825. TYPE: Panama, Haenke
(PRC not seen), synonymized by Tryon (Contr. Gray Herb. 189:60. 1962).

ICeropteris serrata Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:81. 1857. TYPE:
Orizaba, Edo. Veracruz, Mexico, Schaffner 162 (P or RB not seen).

128 AD!ANTACEAE: 26. PITYROGAMMA

Gymnogramma calomelanos var. denudata Harr. J. Linn. Soc., Bot. 16:37. 1877. TYPE: Pebas,
Pda. Loreto, Peru, Steere (MICH not seen; isotype K not seen Tryon photo not seen), examined by
Price (Contr. Univ. Michigan Herb. 15:203 - 204. 1982).

Acrostichum album Veil. FI. Flumin. ll:t. 91. 1831; Arq. Mus. Nac. Rio de Janeiro 5:453. 1881.
TYPE : t. 91, which is based on a collection made by Vellozo from around Rio de Janeiro, Brazil.

Ceropteris calomelanos var. gracilis Hieron. Hedwigia 48:222. 1909. TYPE: Near Coetando on
road between La Plata and Mt. Huila, Depto. Huila, Colombia, 1700 m, Stuebel 1271 (B not seen).

Pityrogramma chamaesorbus Domin, Spisy Pfir. Fak. Karlovy Univ. 88:6. 1928. TYPE: Guyana,
Schomburgk 390 (K not seen Tryon photo not seen).

Pityrogramma calomelanos f. major Domin, Rozpr. Ceske Akad. Ved. Tf. II, Vedy Mat. Prir.
51(18):11. 1941. TYPE: Antahuacana, Depto. Cohcabamba, Bohvia, 750 m, Buchtien 2182 (US).

Pityrogramma calomelanos var. subintegra Domin, Rozpr. Ceske Akad. Ved. Tf. II, Vedy Mat.
Pfir. 51(18):11. 1941. TYPE: Antahuacana, Depto. Cochabamba, Bolivia, 750 m, Buchtien 2181
(US).

Pityrogramma calomelanos var. ochrophylla Domin, Rozpr. Ceske Akad. Ved. Tf. II, Vedy Mat.
Pfir. 51(18):12. 1941. SYNTYPES: Riverside to Minas, Pcia. Camaguey, Cuba, Shafer 1174 (K not
seen); and The Pinales, SE of Paso Estancia, Pcia. Oriente, Cuba, Shafer 1749 (K not seen;
isosyntype US).

Pityrogramma calomelanos var. venezuelae Domin, Rozpr. Ceske Akad. Ved. Tf. II, Vedy Mat.
Pfir. 51(18):12. 1941. TYPE: Colonia Tovar, Edo. Aragua, Venezuela, Fendler 298 (K not seen;
isotype US).

Pityrogramma martinicensis Domin, Vestn. Krai. Ceske Spolecn. Nauk, Tf. Mat.-Pfir. 1941(15): 1.
1942. TYPE: Martinique, Duss 1509 (US).

Plants terrestrial, at 0-1400(2000) m elevation, often on river and roadside
banks, from throughout the Flora area. Also from throughout tropical America,
including Florida.

See P. trifoliata for a discussion of hybrids between it and this variety.

166. Pityrogramma chrysoconia (Desv.) Maxon ex Domin, Spisy Prir. Fak.
Karlovy Univ. 88:10. 1928.

Acrostichum chrysoconium Desv. Mem. Soc. Linn. Paris 6:212. 1827. TYPE: Huasahures

[Huasahuasi?], Depto. Junm, Peru, Dombey (P not seen Cintract photo; isotypes B not seen Tryon
photo, fragm US).

Gymnogramma guianensis Klotzsch, Lirmaea 20:412. 1847. TYPE: Guyana, Rich. Schomburgk
1154 (B not seen Tryon photo not seen fragm NY; isotype K not seen fragm NY not seen).

Gymnogramma omithopteris Klotzsch, Linnaea 20:413. 1847. TYPE: Paramo de Mucuches, Edo.
Merida, Venezuela, Moritz 288 (B not seen Tryon photo fragm US; isotype K not seen Tryon photo).

Gymnogramma flexilis Klotzsch, Linnaea 20:414. 1847. TYPE: Merida, Edo. Merida, Venezuela,
Moritz 287 (B not seen; isotype K not seen fragm NY).

Allosorus farinosus Kunze, Farrnkrauter 2:5, t. 103. 1848, non K. Presl, 1836, nom. illeg. et superfl.
TYPE: A rencuning of Gymnogramma omithoperis Klotzsch, and so based on the type of that name.

Ceropteris obtusa Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:80. 1857. TYPE:
“Province Rio de la Hacha” Sierra Nevada, Comis. La Guajira, Colombia, Schlim 873 (P or RB not
seen; isotype P not seen Tryon photo not seen).

Gymnogramme tartarea var. pallidipes Hook. Sp. Fil. 5:149. 1864. TYPE: Not stated.
Pityrogramma subnivalis Domin, Spisy Pfir. Fak. Karlovy Univ. 88:5. 1928. TYPE: Santa Marta,
Depto. Magdalena, Colombia, Purdie in 1844 (K not seen Tryon photo fragm NY).

Pityrogramma omithopteris f. seminigripes Domin, Lilloa 2:32. 1938. TYPE: Ecuador, Sodiro 352
in 1871 (K not seen).

Pityrogramma omithopteris f. elatior Domin, Lilloa 2:33. 1938. TYPE: Vicinity of Vetas, Depto.
Santander, Colombia, 3100-3250 m, Killip & Smith 17912 (US).

ADIANTACEAE: 26. PITYROGRAMMA 129

Pityrogramma omithopteris f. denudata Domin, Lilloa 2:33. 1938. TYPE: Colombia, Purdie (K not
seen).

Pityrogramma omithopteris var. decomposita Domin, Lilloa 2:34. 1938. TYPE: Near Facatativa,
Depto. Cundinamarca, Colombia, Ariste-Joseph A400 (US), cited by Domin as A400bis, to
distinguish it from another sheet oiA400.

Pityrogramma omithopteris var. sanctae-martae Domin, Lilloa 2:34. 1938. TYPE: Santa Marta,
Depto. Magdalena, Colombia, Smith 1061 (US).

Plants terrestrial, at 20007-3400 m elevation, in the Flora area known only
from along the Interamerican Highway between Cartago and S. Isidro del General
(Scamman 6085, GH) and the summit of Volcan Chiriqui (Maxon 5356, US).
Also from Jamaica, Venezuela to Bolivia, and Guyana.

The fronds of this species may resemble white farinose fronds of F. ehenea var.
ebenea, but the stipes are atropurpureous only at the base in P. chrysoconia and
not throughout, as they are in P. ebenea.

167. Pityrogramma dealbata (K. Presl) Domin, Rozpr. Cesk6 Akad. Ved. Tf. II,
Vedy Mat. Pnr. 51(15):7. 1941.

Gymnogramma dealbata K. Presl, Reliq. Haenk. 1:18, t. 3, f. 1. 1825. LECTOTYPE: Panama,
Haenke (PRC 24358a not seen Prague photo), chosen by Tryon (Contr. Gray Herb. 189:67. 1962).

Ceropteris schaffneri Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:80. 1857.

SYNTYPES: Near Orizaba, Edo. Veracruz, Mexico, Schaffner 165a and 165b (P or RB neither
seen).

ICeropteris plicata Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:81. 1857. TYPE:
Near Tolutla [Totutla], Edo. Veracruz, Mexico, Schaffner 164 (P or RB not seen).

Plants terrestrial or epipetric, at 0-1100(1600) m elevation, on cliffs, road cuts,
and river banks, from Pta. Guapilol and Isla de Cano (Pcia. Puntarenas), the
slopes of the Cordillera Central and the Fila de Cedral, and the Pacific slopes and
lowlands of west-central Panama. Also from Mexico to Nicaragua.

Young, sterile fronds of this species, in which the ultimate segments have not
yet contracted to their final shape, may be confused with fronds .of P. calomelanos
var. calomelanos.

168. Pityrogramma dukei Lellinger, Proc. Biol. Soc. Wash. 89:721, f. 4. 1977.

TYPE: Near the Rio Truando, 3-5 km above the airport at La Teresita, Depto.
Choco, Duke 11200 (US; isotype NY).

Plants terrestrial, at 0-100 m elevation, in disturbed areas along roads and
trails, from the northern half of the Depto. Choco.

This species is most closely related to P. pearcei. It is now known to bear white,
as well as brilliant yellow, farina.

169. Pityrogramma ebenea var. aurata (T. Moore) Lellinger, Proc. Biol. Soc.
Wash. 98:387. 1985.

Gymnogramma tartarea var. aurata T. Moore, Card. Chron. 1870(15):493. 1870. TYPE:

Cultivated by Veitch, originally from Peru (not seen); Tryon (Contr. Gray Herb. 189:66. 1962) noted
that Peru, Pearce 218 (K not seen photo GH not seen) is authentic, and may be the type.

Pityrogramma praestantissima Domin, Spisy Prir. Fcik. Karlovy Univ. 88:6. 1928. TYPE: Depto.
Antioquia, Colombia, Kalbreyer 1909 (K not seen photo GH not seen).

Pityrogramma presliana Domin, Spisy Pfir. Fak. Karlovy Univ. 88:6. 1928. TYPE: Peru, Mathews
1823 (K not seen photo GH not seen).

130 ADIANTACEAE: 26. PITYROGAMMA

Pityrogramma presliana Domin, Spisy Pfir. Fak. Karlovy Univ. 88:6. 1928. TYPE: Peru, Mathews
1823 (K not seen photo GH not seen).

Pityrogramma presliana var. herzogii Domin, Vestn. Krai. Ceske Spolecn. Nauk, Tr. Mat.-Prir.
1941(15):3. 1942. TYPE: Tres Cruces, Cordillera de Sta. Cruz, Depto. Sta. Cruz, Bolivia, 1500 m,
Herzog 1532 (US).

Pityrogramma galapagoensis Domin, Vestn. Krai. Ceske Spolecn. Nauk, Tf. Mat.-Pfir. 1941(15):4.
1942. TYPE: Bindloe Island, Galapagos Islands, Ecuador, Snodgrass & Heller 782 (US)-.

Plants terrestrial, at 2500-3000 m elevation, on roadside banks, in the Flora
area known only from the Cerro de la Muerte, Pcia. S. Jose (Gomez 6759, CR)
and from Providencia de Copey, Pcia. S. Jose (Gomez 6715, CR). Also from
Colombia, the Galapagos Islands, and Ecuador to Argentina.

170. Pityrogramma ebenea (L.) Proctor, Brit. Fern Gaz. 9:219. 1965, var. ebenea.

Acrostichum ebeneum L. Sp. PI. 2:1071. 1753. LECTOTYPE: A specimen, locality and collector
unknown, labelled “Acrostichum 20 ebeneum” in the hand of Linnaeus (LINN 1245.14 not seen
microfiche S. I. Library), chosen by Proctor (Brit. Fern Gaz. 9:219. 1965).

Acrostichum tartareum Cav. Descr. PI. 242. 1801. TYPE: Near Guamantanga, Depto. Lima, Peru,
Nee (MA not seen fragm B-Hb. Willd. 19568-3 not seen Tryon photo).

Hemionitis dealbata Willd. Sp. PI. ed. 4, 5:131. 1810, nom. superfl. TYPE: A renaming of
Acrostichum tartareum Cav., and so based on the type of that name.

Gymnogramma peruviana Desv. Ges. Naturf. Freunde Berlin Mag. 5:329. 1811. TYPE: Peru, /.
de Jussieu (P-Hb. Juss. 1009 not seen Cintract photo).

Ceropteris pallescens Fee, Cat. Foug. Mex. [Mem. Foug. 9]: 14. 1857, nom. nud. TYPE: Valley of
Orizaba, Edo. Veracruz, Mexico, Schaffner 159 (P or RB not seen).

Ceropteris stuebelii Hieron. Hedvdgia 48:223, t. 10, f. 9. 1909. TYPE: Near Ibague, Depto.
Tohma, Colombia, Stuebel 12 (B not seen fragm US; isotype GH).

Pityrogramma tripinnata Domin, Spisy Prir. Fak. Karlovy Univ. 88:7. 1928. TYPE: Region of S.
Luis Potosi, Edo. S. Luis Potosi, Mexico, Parry & Palmer 1004 (K not seen Tryon photo; isotype US).

Pityrogramma amgibua Domin, Vestn. Krai. Ceske Spolecn. Nauk, Tf. Mat.-Pnr. 1941(15): 3.
1942. TYPE: Laguna de Chiriquf and vicinity, Pcia. Bocas del Toro, Hart 24 (US). This is said by
Domin to be a hybrid, but is more hkely just a highly divided form of P. tartarea.

IPityrogramma tartarea var. mathewsii Domin, Vestn. Krai. Ceske Spolecn. Nauk, Tf. Mat.-Pffr.
1941(15):6. 1942. TYPE: Quebrada de Pariahuanca, Colombia, Mathews 970 (K not seen).

IPityrogramma tartarea var. kalbreyeri Domin, Vestn. Krai. Ceske Spolecn. Nauk, Tf. Mat.-Pnr.
1941(15):6. 1942. TYPE: Ocana, Depto. Norte de Santander, Colombia, 5000 ft, Kalbreyer 341 (K
not seen).

IPityrogramma tartarea var. pteridoides Domin, Vestn. Krai. Ceske Spolecn. Nauk, Tf . Mat.-Pffr.
1941(15):7. 1942. SYNTYPES: Edo. Chiapas, Mexico, Ghiesbreght Fil Austro-Mex. 276 (K not
seen); and Mexico, Graham 408 in 1830 (K not seen).

IPityrogramma tartarea var. delicatula Domin, Vestn. Krai. Ceske Spolecn. Nauk, Tf. Mat.-Pffr.
1941(15):7. 1942. TYPE: Loja, Pcia. Loja, Ecuador, collector unknown (K not seen).

Plants terrestrial or rarely epipetric, at 0-2700(3200) m elevation, in forests and
open areas, often on roadside banks, from the Cordillera Central, the Meseta
Central, the Fila de Cedral, the Cordillera de Talamanca, the Canal Zone, and
near Quibdo (Depto. Choco). Also from the Greater Antilles, Mexico to
Nicaragua, Venezuela to Bolivia, and Brazil.

171. Pityrogramma ferruginea (Kunze) Maxon, Contr. U. S. Natl. Herb. 17:173.
1913.

ADIANTACEAE: 26. PITYROGRAMMA 131

Pityrogramma lanata Klotzsch ex A. Braun, App. Ind. Sem. Hort. Berol. 17. 1854. TYPE:

Cultivated in the garden at Berlin, originally from Pcia. Veraguas, Panama, Warscewicz (presumably
B not seen).

Gymnogramma hommeri Christ in Dur. & Pitt. Bull. Soc. Roy. Bot. Belgique 35, Mem. 237. 1896.
SYNTYPES: Between the Llanuras de S. Carlos and the Cuesta de la Vieja, Pcia. Heredia, 750 m,
Biolley 1066 (BR not seen); and Carrillo, Pcia. S. Jos6, 300 m, Pittier 1169 (BR not seen).

Plants terrestrial, along road and river banks, at 100-2000 m elevation, from
the Cordillera de Tilardn, the Cordillera Central, the Cordillera de Talamanca to
Pcia. Chiriqm, and the Fila Costeha near S. Vito. Also from Guatemala, El
Salvador, Nicaragua, and Peru.

172. Pityrogramma pearcei (T. Moore) Domin, Spisy Pnr. Fak. Karlovy Univ.

88:9. 1928.

Gymnogramma pearcei T. Moore, Card. Chron. 1864:340. 1864. TYPE: Cultivated by Veitch
from Andean specimens collected by Pearce, according to Tryon (Contr. Gray Herb. 189:73. 1962),
who provided details of the authentic material (K not seen).

Gymnogramma decomposita Baker, Gard. Chron. 1872:1587. 1872. TYPE: Cultivated by Veitch
from Andean specimens [probably collected by Pearce], according to Tryon (Contr. Gray Herb.
189:73. 1962), who provided details of the authentic material (K not seen Tryon photo).

Gymnogramma elegantissima Pynaert, Rev. Hort. Beige Etrangere 15:211. 1898, non Bull, 1889,
nom. illeg. TYPE: Unknown.

Gymnogramma fumarioides Rosenst. M6m. Soc. Sci. Nat. Neuchatel 5:45, t. 6, f. 10. 1912. TYPE:
Lake above Medellin, Depto. Antioquia, Colombia, Mayor 144 (S not seen; isotypes P not seen Tryon
photo, US).

Plants terrestrial along streams in open areas, at 1200-1500 m elevation, from
Tapanti and Cerro Colorado 20-28 mi from S. Felix (Pcia. Chiriqui). Also from
Colombia.

173. Pityrogramma trifoliata (L.) Tryon, Contr. Gray Herb. 189:68. 1962.

Acrostichum trifoliatum L. Sp. PI. 2:1070. 1753. LECTOTYPE: Presumably t. 45, f. 2 of Sloane’s
“Voyage....” The description in Linnaeus’ “Amoenitates” could not have been drawn from Sloane’s
description alone, but the decreasing division of the pinnae mentioned in the description in the
“Amoenitates” is shown in Sloane’s illustration. The specimen in the Linnaean Herbarium (LINN
1245.9 not seen microfiche S. I. Library) is labelled “Acrostichum 15 trifoliatum” in Linnaeus’ hand.
It might have been from Jamaica, received from Houstoun prior to 1753, in which case it should be
the lectotype. According to Jackson’s “Index,” however, it and LINN 1245.10, to which 1245.9 is
pinned, were not present in the Linnaean herbarium in 1753, but were present in 1767.

Trismeria argentea Fee, Gen. Fil. [M6m. Foug. 5]:165, t. 14A, f. 1. 1852. SYNTYPES: Caracas,
Distr. Fed., Venezuela, Mofitz (P or RB not seen); and Est. Minas Gerais, Brazil, Pohl (P or RB not
seen).

Trismeria aurea F6e, Gen. Fil. [M6m. Foug. 5]: 165. 1852, nom. superfl. TYPE: A renaming of
Acrostichum trifoliatum L., and so based on the type of that name.

Trismeria microphylla Fee, Gen. Fil. [Mem. Foug. 5]: 165. 1852. TYPE LOCALITY: Peru.

Plants terrestrial, at 0-1800 m elevation, along rivers and roadsides in
disturbed areas, from the Cordillera de Tilaran, the Cordillera Central, Cerro
Tablazo, the Cordillera de Talamanca to Pcia. Chiriqui, the Peninsula de Burica
(Pcia. Chiriqm), and the Rio Yape (Pcia. Darien). Also from Florida, the Greater
Antilles, Mexico to Nicaragua, Venezuela to Bolivia, Brazil, Paraguay, Argentina,
and Chile.

This species is known to hybridize with P. femiginea and with several varieties of
P. calomelanos, but only the hybrid with var. calomelanos has been found as yet in

132 ADIANTACEAE: 27. HEMIONITIS

the Flora area, from the Rio Pejivalle, Pda. Cartago {Gomez 6788, CR), and from
road banks 4 km before the bridge over the Rio Pacuarito, Pda. Limon (Gdmez
6805, CR). Gomez and Wollenweber (Amer. Fern J. 68:121. 1978) have shown
that Costa Rican material of this species is variable in pinna division, with simple
and bipartite pinnae as or more common than tripartite pinnae.

27. HEMIONITIS L.

Plants terrestrial or occasionally epipetric, often on banks or cliffs; rhizomes
short-creeping to erect, scaly, the scales narrowly lanceate with a long, hair-like
apex, concolorous, tan, thin, the margins entire; fronds small- to medium-sized,
monomorphic or slightly dimorphic with a basal rosette of sterile fronds and taller,
upright fertile fronds; stipes atropurpureous, shiny, scaly and pilose, especially at
the base, the hairs tan, multicellular; laminae lobed, usually pentagonal,
herbaceous, with the color of the stipe extending into the midribs of the lamina
lobes, slightly pilose on both surfaces, the hairs thin, straight, tan, multicellular
with the walls of adjoining cells obvious and sometimes dark in color; lamina lobes
blunt to elongate, the basal pair usually bearing a single, basiscopic lobe, entire to
broadly crenate, plane at the margin; venation of elongate areolae lacking free
included veinlets; indusia absent; sporangia scattered along the veins, short-
stalked.

Central America and the West Indies; 7 species.

MICKEL, J. T. 1974. A redefinition of the genus Hemionitis. Amer. Fern J. 64:3 - 12.

1. Laminae broadly ovate to nearly round in outline, entire, scarcely lobed, with linear hairs 1- or
2-celled distal to the slightly bulbous base, the articulations pale and inconspicuous. Stipes 1.5-7 cm
long; laminae 1-4 cm long, 1-4 cm wide.

m.H.leyyi

1. Laminae more or less pentagonal in outline, mostly shallowly to deeply crenate, deeply lobed,
with acicular hairs 2-5-celled distal to the bulbous base, the articulations dark and conspicuous.. 2.

2(1). Laminae with a single lateral pair of lobes each usually having a single basiscopic lobe; fertile
lamina margins usually deeply crenate. Fertile stipes 2.5-10 cm long; laminae 2.5-10 cm long,
2-14 cm wide.

175. H. palmata

2(1). Laminae with usually 2 lateral pairs of lobes, the basal lobes each usually having a single
basiscopic lobe; fertile lamina margins shallowly crenate to entire. Fertile stipes 1-4 cm long;
laminae 3- 10 cm long, 2- 13 cm wide.

176. H. pinnatifida

174. Hemionitis levyi Foum. Bull. Soc. Bot. France 17:237. 1870.

Hemionitis otonis Maxon, Contr. U. S. Natl. Herb. 17:171. 1913. TYPE: Along road from Ojo de
Agua to Brasil (Santa Ana), Pcia. S. Jos6, ca. 800 m, O. Jiminez 333 (US).

TYPE: Isla de Omotepe, Depto. Rivas, Nicaragua, 40 m. Levy 1197 not seen;
isotypes HBG not seen photo 5536, US).

Plants terrestrial, at ca. 800 m elevation, in rocky places, in the Flora area
known only from the type of H. otonis and from Garrobillos, Canton Puriscal, Pcia.
S. Jose {Gdmez 617, CR). Also from Mexico, El Salvador and Nicaragua.

175. Hemionitis palmata L. Sp. PI. 2:1077. 1753.

LECTOTYPE: Plate 33, f. 1 of Plunder’s “Description...”, which is based on a
specimen collected by Plumier on Martinique, chosen by Proctor (Ferns Jamaica
213.1985). The specimen of this species annotated by Linnaeus (LINN 1248.3 not

ADIANTACEAE: 28. GYMNOPTERIS 133

seen microfiche S. I. Library) was not in the Linnaean Herbarium prior to 1756,
according to Jackson’s “Index,” and so cannot be the type.

Plants terrestrial, at 500-1900 m elevation, on roadside banks and in pastures,
from the Peninsula de Nicoya, the vicinity of S. Ramon, the Atlantic coastal plain
of northern Costa Rica, the upper Rio Reventazdn valley, and the upper Rio
General valley. Also from the Antilles, Mexico to Nicaragua, Trinidad,
Venezuela, Colombia, Peru, Bolivia, Surinam, and Brazil.

176. Hemionitis pinnatifida Baker, Syn. Fil. 399. 1868.

TYPE: Central America [presumably Costa Rica], Wendland 438 (presumably K
not seen).

Plants terrestrial, at ca. 1000 m elevation, in rocky places, in the Flora area
known only from Garrobillos, Canton Puriscal, Pcia. S. Jose {Gomez 617, CR, a
mixed collection also containing H. levyi). Also from Mexico to Nicaragua.

Although both this species and H. palmata are dimorphic, with longer fertile
fronds, H. pinnatifida shows incomplete dimorphism occasionally, with one or
more of the usually sterile rosette leaves partially or entirely fertile.

28. GYMNOPTERIS Bernh.

Plants terrestrial; rhizomes short-creeping to erect, scaly, the scales linear-
lanceate with a long, hair-like apex, bicolorous, tan with a blackish central stripe,
the margins entire; fronds medium-sized, monomorphic; stipes stramineous,
castaneous, or atropurpureous and shiny, scaly and pilose or pilose and pilosulous,
the scales linear, tan, entire, the large hairs straight, tan, multicellular, with the
wall of adjoining cells obvious, the small hairs apparently unicellular, acicular,
minute, tan; laminae 1-2-pinnate, debate to oblong, the pinnae ovate-lanceolate,
herbacous, short-stalked, the margins entire, slightly hairy on both surfaces, the
hairs thin, straight, tan, multicellular with the walls of adjoining cells obvious and
sometimes dark in color; venation free or areolate with free veins at the margins,
the areolae elongate, lacking free included veinlets; indusia absent; sporangia
scattered along the veins, short-stalked.

Tropical America and Asia; 5 species.

LELLINGER, D. B. 1969. The taxonomic position of Coniogramme americana. Amer. Fern J.

59:61-65.

MICKEL, J. T. 1974. A redefinition of the genus Hemionitis. Amer. Fern J. 64:3-12.

1. Venation entirely free; stipes and rachises castaneous to atropurpureous. Laminae oblong,
pinnate, 20-40 cm long, 4.5-14 cm wide; lateral pinnae 6-12 pairs, ovate-lanceolate to oblong,
equilateral at the base, abruptly acute to cuneate at the apex, 2-8 cm long, 1-2.5 cm wide; spores
tuberculate.

177. G. rufa

1. Venation partially areolate, free only at the margins; stipes and rachises stramineous. Laminae
debate to lanceate, piimate or rarely pinnate-pinnatifid or 2-pinnate, 7-23 cm long, 6.5-18 cm wide;
lateral pinnae 1-6 pairs, ovate-lanceolate, shghtly inequilateral at the base, gradually acute to
cuneate at the apex, 4- 10 cm long, 2-4.5 cm wide; spores cristate.

178. G. subcordata

177. Gymnopteris nifa (L.) Bemh. ex Underw. Bull. Torrey Bot. Club 29:627.

1902.

Pteris rufa L. Sp. PI. 2:1074. 1753, as LECTOTYPE: Plate 45, fig. 1 of Sloane’s

“Voyage...”, chosen by Linnaeus (Syst. Nat. ed. 10, 2:1320. 1759).

134 ADIANTACEAE: 28. GYMNOPTERIS

ADIANTACEAE: 29. BOMMERIA 135

Plants terrestrial, at 0-400 m elevation, in forests, from the valley of the Rio
General and the slopes and coastal plain from near Puntarenas to Juan Diaz
(Pcia. Panama). Also from Cuba, Jamaica, Mexico to Honduras, Nicaragua,
Venezuela, Colombia, Ecuador, Peru, Surinam, and Brazil.

178. Gymnopteris subcordata (D. C. Eaton & Davenp. in Rose) Underw. Bull.
Torrey Bot. Club 29:628. 1902.

Gymnogramme subcordata D. C. Eaton & Davenp. in Rose, Contr. U. S. Natl. Herb. 5:138, t. 16.
1897, non Coniogramme subcordata Copel., 1910. LECTOTYPE: Yamala, Edo. Sinaloa, Mexico,
Palmer 1416 (US; isolectotypes NY, UC neither seen), chosen by Mickel & Beitel (Mem. New York
Bot. Card. 46:209. 1988).

Coniogramme americana Maxon, Contr. U. S. Natl. Herb. 17:607. 1916. TYPE: A renaming of
Gymnogramma subcordata D. C. Eaton & Davenp. in Rose, and so based on the type of that name.

Plants terrestrial, at ca. 500 m elevation, on shady banks, in the Flora area
known only from above the old hotel plaza on Volcdn Poas {Gomez 3025, Hb.
Gomez) and from the valley of the Rio General {Gomez 255, Hb. Gomez). Also
from Mexico, Guatemala, and Nicaragua.

29. BOMMERIA Fourn. in Baill.

Plants epipetric or terrestrial; rhizomes short-creeping, branched, scaly, the
scales narrowly lanceate, weakly bicolorous, dark brown with paler, reddish-brown
margins, the margins entire; fronds small- to medium-sized; stipes much longer
than the laminae, reddish-brown to dark brown or atropurpureous, rather shiny,
scaly at the base, subterete; laminae pentagonal, subternate, the basal pinnae
strongly basiscopically developed, bipinnatifid proximally, bipinnatifid or
pinnatifid distally, herbaceous, pilose on both surfaces and on the rachis or midrib,
especially abaxially, the hairs pluricellular with obscure cross-walls, rather stiff,
pale tan or reddish-tan; veins free and forked or forming polygonal areolae lacking
included veinlets; sori elongate, irregular, the sporangia lateral on the veins,
especially near and at the plane or revolute segment margins; sporangia short-
stalked.

Central America and adjacent United States; 4 species.

HAUFLER, C. R. 1979. A biosystematic revision of Bommeria. J. Arnold Arbor. 60:445 - 476.

179. Bommeria pedata (Swartz) Foum. Bull. Soc. Bot. France 27:327. 1880.

Hemionitis pedata Swartz, Syn. Fil. 20, 209, t. 1, f. 3. 1806. TYPE: Without locality or collector, ex
Herb. Cavanilles (presumably S not seen), perhaps collected by N6e in Mexico, according to Haufler
(1979, p. 471).

1 Gymnogramme pedata var. palmata Kunze, Linnaea 13:130. 1839. TYPE: Barranca Teocelo,
Edo. Veracruz, Mexico, Aug 1829, Schiede (presumably LZ destroyed).

FIGS. 174-182. Hemionitis, Gymnopteris, Bommeria, aaid Jamesonia. FIG. 174. Frond of H levyi,
Jim6nez L. 333. FIG. 175. Frond of H. palmata, Scamman 6080. FIG. 176. Frond of H. pinnatifida,
Maxon 7700, Nicaragua. FIG. 177. Median pinna of G. mfa, Pittier 5212. FIG. 178. Subapical pinna
of G. subcordata, McVaugh 15925, Mexico. FIG. 179. Plant of B. pedata, Pringle 11781, Mexico.
FIG. 180. Median portion of frond of /. alstonii, Lellinger 873. FIG. 181. Median portion of frond of
/. rotundifolia, Williams 20044. FIG. 182. Median portion of frond of /. scammanae. Carpenter 289.

136 ADIANTACEAE: 30. JAMESONIA

Plants epipetric, at 1400 m elevation, in sunny places, in the Flora area known
only from Volcdn Tenorio, Pcia. Guanacaste {Gomez 69867, CR). Also from
Mexico, Guatemala, Honduras, El Salvador, and Nicaragua.

30. JAMESONIA Hook. & Grev.

Plants terrestrial in pdramos and subpdramos; rhizomes short- to long-creeping,
branched, bearing bristles or bristle-like hairs, these mostly dark brown,
pluricellular, acicular, 1 cell wide distal to the base or throughout; fronds small- to
medium-sized; stipes short, densely pilose, bristly at the base, terete, trigonous, or
sulcate; laminae pinnate, linear, coriaceous, indeterminate, with a permanent,
indeterminate crozier, the apex glutinous or protected by hairs; pinnae short-
stalked, simple, imbricate, minute (mostly less than 1 cm long), round or slightly
elongate, often villous abaxially, strongly revolute; veins free, forked, usually
mostly hidden within the lamina tissue; sori elongate, lateral on the veins,
exindusiate but protected by the revolute pinna margins; sporangia long-stalked.

Highlands of Central and South America, especially in the Andes; ca. 20
species.

TRYON, A. F. 1962. A monograph of the fern genus Jamesonia. Contr. Gray Herb. 191:109-197.

1. Pinnae 1-2 mm wide; laminae up to 3 mm wide; hairs on the rachis and abaxial surface of the
pinnae shorter than the pinnae, the plants appearing nearly glabrous, even at the frond apex.

182. /. scammanae

1. Pinnae 2-9 mm wide; laminae 3-10 mm wide; hairs on the rachis and abaxial surface of the
pinnae often longer than the pinnae, the plants obviously hairy, especially at the frond apex. .2.

2(1). Pinnae sparsely villous on the adaxial surface, especially near the pinna margins; pinnae
(2)4-9 mm in diam., up to (2)3-5 mm distant, herbaceous; apical bud of the laminae with its hairs
usually wider than the pinnae just proximal to the apex.

181. /. rotundifolia

2(1). Pinnae usually glabrous on the adaxial surface, occasionally very sparsely villous; pinnae 2-5
mm in diam., 2 mm distant, coriaceous; apical bud of the lamina with its hairs usually no wider than
the pinnae just projdmal to the apex.

180. /. alstonii

180. Jamesonia alstonii A. F. Tryon, Contr. Gray Herb. 191:168, f. 11. 1962.

TYPE: La Laguna, Cerro de la Torre, Los Farallones, Cordillera Occidental,

Depto. del Valle, Colombia, 3500-3550 m, Cuatrecasas 21884 (GH not seen;
isotypes COL not seen, US).

Plants terrestrial, at 3400-3800 m elevation, in paramos and subparamos, from
the Cerro de la Muerte and Cerro Chirripo. Also from Mexico, Guatemala,
Nicaragua, and Colombia to Bolivia.

This species apparently hybridizes on the Cerro de las Vueltas with a species of
Eriosorus {Gdmez 686, CR).

181. Jamesonia rotundifolia F6e, M6m. Soc. Sci. Hist. Nat. Strasbourg 5 [M6m
Foug.7]:41,t. 10,f.3. 1857.

TYPE: Ocaha, Depto. Norte de Santander, Colombia, 3300 m, Schlim 363 (RB
not seen; isotypes G not seen, K not seen, P not seen).

Plants terrestrial, at 3000-3500 m elevation, in paramos and subparamos, from
the Cerro de las Vueltas, the Cerro de la Muerte, and Cerro Chirripo. Also from
Colombia to Peru.

ADIANTACEAE: 31.ERI0S0RUS 137

182. Jamesonia scammanae A. F. Tryon, Contr. Gray Herb. 191:164, f. 9. 1962.

TYPE: Cerro de la Muerte, Pda. Cartago?, Scamman 7929 (GH; isotype US).
Plants terrestrial, at 3000-3800 m elevation, in paramos and subparamos, from
the Cerro de las Vueltas, the Cerro de la Muerte, and Cerro Chirripo. Also from
Colombia to Bolivia.

31. ERIOSORUS

Plants terrestrial in montane and pdramo areas; rhizomes short- to long-
creeping, branched, bearing pluricellular hairs, bristles, or rarely scales; fronds
medium- to large-sized, the larger ones indeterminate and scrambling; stipes
commonly pilose or glandular, mostly castaneous to atropurpureous, terete or
nearly so, at least at the base; laminae (l)2-3-^innate, rhombic or lanceate to
linear, herbaceous, in most species less than 10(20) cm wide; rachises similar to
the stipes but more often glabrous or nearly so and sulcate, sometimes flexuous;
ultimate segments orbicular, obovate, ovate, bifid, or narrowly elliptic, often
pilose, plane or revolute; veins free, commonly forked, usually visible; sori
exindusiate, elongate, lateral on especially the ultimate branches of the veins,
sporangia short-stalked.

Middle to high elevations in the tropics of the New World, especially the Andes,
and on a few Atlantic Islands; ca. 25 species.

TRYON, A. F. 1970. A monograph of the fern genus Eriosorus. Contr. Gray Herb. 200:54-174.

1. Fronds scandent or recumbent; rachises usually markedly flexuous; laminae up to ca. 2.5 m long;
pinnae mostly 12-25 cm long..5.

1. Fronds erect; rachises not or only slightly flexuous; laminae 10 - 45(60) cm long; pinnae mostly
(1)3-8 cm long..2.

2(1). Pinnae irregular, ca. 1 cm long, the ultimate segments almost bead-like.

188. Eriosorus warscewiczii x Jamesonia scammanae

2(1). Pinnae regular, usually 3-8 cm long, the ultimate segments not bead-like..3.

3(2). Rhizomes short-creeping, the stipes congested, long- villous; laminae long- villous on both
surfaces. Fronds 15 - 45 cm long, 3-pinnate-pinnatifid, membranaceous, the margins plane.

183. E. congestus

3(2). Rhizomes long-creeping, the stipes distant, glabrous or nearly so..4.

4(3). Fronds 2-pmnate-pinnatifid, subcoriaceous, the margins revolute. Laminae glabrous on the
adaxial surface, short-villous abaxially; fronds 10 - 40(60) cm long.

187. E. warscewiczii

4(3). Fronds pinnate-pinnatifid or subbipinnate, chartaceous, the margins nearly plane. Laminae
sparsely pilose on both surfaces; fronds 15-28 cm long.

185a. E. novogranatensis

5(1). Rachises castaneous to very dark brown throughout; pinnule axes not flexuous, slightly
retrorse, strongly arcuate-ascending; laminae glabrous.

185. E. glaberrimus

5(1). Rachises stramineous or pale brown towards the apex; pinnule axes slightly flexuous, slightly
retrorse, not ascending; laminae glabrous to densely pubescent, especially on the axes..5.

6(5). Lamina tissue glabrous to sparsely pubescent; ultimate segment lobes usually (1)1.5 -3 mm
long and widely divergent.

184. E. flexuosus wai.flexuosus

6(5). Lamina tissue densely short-villous on the abaxial surface, less so adaxially; ultimate segment
lobes usually 0.5- 1(1.5) mm long and not widely divergent.

186. E. villosulus

138 ADIANTACEAE: 31. ERIOSORUS

187 E. warscewiczii

186 E. villosulus

189 A. andicola

ADIANTACEAE: 31.ERI0S0RUS 139

183. Eriosoms congestus (Christ) Copel. Gen. Fil. 58. 1947.

Gymnogramma congesta Christ, Bull. Herb. Boissier II, 4:1098. 1904. LECTOTYPE: La Palma,
Pda. S. Jose, 1459 m, Tonduz 12575 (P not seen; isotypes CR, G not seen, US fragm NY), chosen by
Maxon (Bull. Torrey Bot. Club 42:81. 1915).

Plants terrestrial, at 1300-2600 m elevation, on road banks and in pastures and
forest clearings, from the Cordillera Central, the Cordillera de Talamanca to near
S. Isidro del General, and above Cerro Punta ca. 2 mi east of Guadalupe (Pcia.
Chiriqui).

A. F. Tryon (1970, p. 170) reported hybrids of this species with E. glaberrimus
from La Palma, which have slightly flexuose rachises, non-arcuate pinnae, ultimate
segments similar to those of E. congestus, and abortive spores.

184. Eriosoms flexuosus (H.B.K.) Copel. Gen. Fil. 58. 1947, var. flexuosus.

Gymnogramma flexuosa Desv. Ges. Naturf. Freunde Berlin Mag. 5:306. 1811. TYPE: “Am.
Aequin.,” collector unknown (P-Hb. Desv. not seen Tryon photo).

Grammitis flexuosa H.B.K. Nov. Gen. Sp. 1:5 (fol. 4). 1816. TYPE: Near Caracas, Distr. Fed.,
Venezuela, Humboldt & Bonpland (P-Hb. Bonpl. not seen photo 4447; isotype B-Hb. Willd. 19597
not seen Tryon photo).

Gymnogramma refracta Kunze ex Klotzsch, Linnaea 20:410. 1847. TYPE: Sierra Nevada, Edo.
Merida, Venezuela, Moritz 359 (B not seen Tryon photo; isotypes BM not seen photo 7492, P not
seen Weatherby photo, US).

Gymnogramma miziana Kunze ex Klotzsch, Linnaea 20:410. 1847. TYPE: Pcia. Panatahuas,
Depto. Hu^uco, Peru, Ruiz 74 (B not seen Tryon photo).

Gymnogramma haematodes Christ, Bull. Herb. Boissier II, 4:1097. 1904. TYPE: Summit of
Volc^ Poas, Pcia. Alajuela, 2644 m, Tonduz 10713 (P not seen Weatherby photo; isotype US),
chosen by Maxon (Bull. Torrey Bot. Club 42:84. 1915).

Gymnogramma flexuosa var. linearis Christ, Bull. Herb. Boissier II, 4:1096. 1904. TYPE: Las
Vueltas, Tucurrique, Pcia. Cartago, Tonduz 12796 (P not seen; isotype G not seen).

Gymnogramma flexuosa wax. pemviana Hieron. Hedwigia 48:220. 1909. TYPE: Cuesta de Lejia,
near Molinopampa, Depto. Amazonas, Peru, 2300 m, Stuebel 1055 (B not seen Tryon photo).

Gymnogramma platytrichia C. Chr. Kongl. Svenska Vetensk. Akad. Handl. 16(2) :58. 1937.

TYPE: Les Roseaux, Massif de la Hotte, Haiti, 2300 m, Ekman 10657 (S not seen; isotypes B not
seen, US).

Plants terrestrial, at (600)1500-2800(3300) m elevation, in forests, clearings,
and on road banks, from the Cordillera Central, the Fila de Cedral, the Cordillera
de Talamanca, and the summit of the west peak of the Cerro Tacarcuna massif.
Also from Hispaniola, Mexico, Guatemala, El Salvador, Venezuela, Colombia to
Peru, Guyana, and Brazil.

A. F. Tryon (1970, p. 163) reported hybrids of this species with E. warscewiczii
on Volcan Pods.

185. Eriosoms glaberrimus (Maxon) Scamman, Contr. Gray Herb. 191:85. 1962.

Psilogramme glaberrima Maxon, Bull. Torrey Bot. Club 42:82. 1915. TYPE: La Palma, Pcia. S.
Jose, 1459 m, Tonduz 12531 (US fragm NY; isotypes CR, P not seen).

FIGS. 183-191. Eriosoms and Adiantum. FIG. 183. Median pinna of E. congestus, Maxon 8159.
FIG. 184. Pinna of E. flexuosus, de la Sota 5044. FIG. 185. Pinna of E. glaberrimus, Standley 39424.
FIG. 186. Median pinna of E. villosulus, Pittier 10502. FIG. 187. Median pinna of E. warscewiczii,
Tonduz 10712. FIG. 188. Pinna of E. warscewiczii x /. scammanae, Tryon & Tryon 7048 (GH). FIG.
189. Median pinnule of basal pinna of A. andicola, Standley 34699. FIG. 190. Suprabasal pinnule of
suprabasal pinna of^. braunii, Tonduz 8802. FIG. 191. Pinnae of A. caryotideum, Killip 2676.

140 ADIANTACEAE: 32. ADIANTUM

Plants terrestrial, at 1200-2300 m elevation, in forests and along road banks,
from the Cordillera Central and the Cordillera de Talamanca to S. Isidro del
General. Also from Nicaragua.

185a. Eriosoms novogranatensis A. F. Tryon, Contr. Gray Herb. 200:120, f. 19.
1970.

TYPE: Boqueron de Quindiu, Depto. Tolima, Colombia, 3400 m, Alston 7728
(GH not seen; isotype BM not seen).

Plants epiphytic, at 1600-2200 m elevation, from Cerro Pata de Macho and
Cerro Punta (both Pcia. Chiriqui). Also from Colombia.

186. Eriosoms villosulus (Maxon) Scamman, Contr. Gray Herb. 191:88. 1962.

Psilogramme villosula Maxon, Bull. Torrey Bot. Club 42:83. 1915. TYPE: Cerro de las Vueltas,
Pcia. S. Jos6, 3000 m, Pittier 10502 (US).

Plants terrestrial, at (2100)2700-3400 m elevation, in high montane and
paramo regions, from the Cordillera de Talamanca.

A. F. Tryon (1970, p. 162) considered this to be a variant of E. flexuosus, which it
may be, although it differs from typical material of that species in the shape of its
ultimate segments as well as in its pubescence. Scamman (Contr. Gray Herb.
191:88. 1962) maintained E. villosulus as a distinct species, principally because it
usually grows at higher elevations than does typical E. flexuosus. Some or all of
the specimens of E. villosulus may be hybrids between E. flexuosus and E. congestus
or, because they resemble E. flexuosus in lamina architecture, may be backcrosses
between such a hybrid and E. flexuosus.

187. Eriosoms warscewiczii (Mett. in Tr. & Planch.) Copel. Gen. Fil. 59. 1947.

Gymnogramma warscewiczii Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:211 (repr. 293). 1864.
LECTOTYTE: Volcan Turrialba, Pcia. Cartago, 11000 ft, Warscewicz 20 (B not seen Tryon photo
fragm NY), chosen by Maxon (Bull. Torrey Bot. Club 42:80. 1915).

Psilogramme jimenezii Maxon, Amer. Fern J. 18:3. 1928. TYPE: Crater of Volcan Poas, Pcia.
Alajuela, 2800 m, O. Jimenez 1034 (US fragm NY).

Plants terrestrial, at (1700)2000-3400 m elevation, on banks, often along
roadsides, from the Cordillera Central, Cerro Tablazo, and the Cerro de la
Muerte. Also from Colombia.

188. Eriosoms warscewiczii (Mett.) Copel. x Jamesonia scammanae A. F. Tiyon

Gymnogramma kupperi Losch, Mitt. Bot. Staatssamml. Miinchen 1:21. 1950. TYPE: Chirripo
Grande, Pcia. S. Jose?, 3500 m, Kupper 1223 (M not seen photo 7484), placed tentatively by A. F.
Tryon (Contr. Gray Herb. 200:120. 1970).

Plants terrestrial, at 3100-3400 m elevation, in paramos, from the Cordillera de
Talamanca.

If hybrids between Eriosorus and Jamesonia are assigned to a separate genus,
the correct name for this hybrid is Eriosonia kupperi (Losch) Pic. Ser.

32. ADIANTUM L.

Plants terrestrial or sometimes epipetric; rhizomes mostly ascending or short-
creeping, occasionally long-creeping, scaly especially at the apex, the scales small,
lanceate, obscurely clathrate or eclathrate, brown to blackish, usually concolorous;
fronds medium-sized; stipes long, wiry, shiny, atropurpureous or blackish,
occasionally dark brown, often obscurely scaly or pubescent; laminae 1-3(5)-

ADIANTACEAE: 32. ADIANTUM 141

pinnate, rarely simple, usually oblong or lanceolate with a pinnatifid apex, the
apex conform in a few species, membranaceous to firmly herbaceous; ultimate
segments sessile to short-stalked, articulate and deciduous in some species,
flabellate, oblong, or trapezoidal, sometimes dimidiate, commonly glaucous,
sparsely pilosulous or with small, dissected, even stellate scales on the abaxial
surface; veins free, forked, rarely anastomosing, usually visible; sori elongate,
superficial on the reflexed, marginal, usually elongate to linear false indusia;
sporangia long-stalked.

Temperate to tropical regions world-wide, especially at low and middle
elevations in tropical America; ca. 200 species.

SCAMMAN, E. 1960. The Maidenhair ferns (Adiantum) of Costa Rica. Contr. Gray Herb.

187:3-22.

1. Veins free; laminae 1-3(5) -pinnate.. 3.

1. Veins copiously anastomosing, the areolae elongate with short-tapered ends, lacking included
veinlets..2.

2(1). Laminae simple. Rhizomes short-creeping, knotted, the stipe bases approximate; stipes
10-35 cm long; laminae cordate at the base, acute at the apex, 7-13 cm long, 5-7 cm wide; sori
continuous along the lateral margins of the lamina.

193. A. cordatum

2(1). Laminae pinnate. Rhizomes long-creeping, the stipe bases 2-5 mm distant; laminae broadly
triangular to broadly ovate, 10-15(20) cm long, 10-13(18) cm wide; lateral pinnae 1-3(5)
subopposite to alternate pairs; terminal pinna conform; sori continuous along the lateral margins of
the pinnae.

219. A. wilsonii

3(1). Laminae digitate-pedately divided, the basal pinna usually 4-pinnate, basiscopically
developed, forming almost half the lamina. Laminae obhquely deltate, 15-20 cm long, 15-30 cm
wide, usually sparsely puberulent; sori reniform to orbicular, 4-8 on the distal and outer edge of the
ultimate segments.

204. A. patens

3(1). Laminae pinnately divided, the basal pinnae not basiscopically developed and forming half
the lamina..4.

4(3). Ultimate segments several times longer than wide, partially to entirely dimidiate, sessile or
very short-stalked; axes scaly or hairy in most species.. 18.

4(3). Ultimate segments usually no more than 1.5(2) times longer than wide, not dimidiate (except
in A. trapeziforme and partially so in A. caryotideum), short- to long-stalked; axes glabrous in most
species..5.

5(4). Dark color of the stalk abruptly terminating at the base of the ultimate segment, the
segments articulate. Costule apex swoUen in some species.. 10.

5(4). Dark color of the stalk gradually passing into the base of the ultimate segment, the segments
never articulate..6.

6(5). Laminae pinnate. Laminae oblong, 15 - 25(30) cm long, 8-16(18) cm wide; pinnae glaucous,
subsessile, lanceate, the distal ones excavate basiscopically; sori usually continuous on both margins.

7Q1.A. macrophyllum

6(5). Laminae 2-3-pinnate..7.

7(6). Laminae 2-piimate, the basal 2-4 pinna pairs pinnate, the many distal pinnae all simple.
Laminae ovate-lanceolate, 15-35 cm long, 15-25 cm wide, the ultimate segments 2-4(5) cm long,
trapeziform or ovate and excavate at the lower base.

191.^. caryotideum

7(6). Laminae (2)3-pinnate, almost all the pinnae l-2-pimiate..8.

8(7). Pinnae subsessile, the basal acroscopic pinnule 1-3 mm from the rachis; rhizome scales ca. 2
mm long, 0.2 mm wide. Rhizomes compact; laminae lanceolate, (10)20-50 cm long, (6)8-25 cm

142 ADIANTACEAE: 32. ADIANTUM

wide, (2)3-pmnate; basal acroscopic pinnules larger than the distal pinnules or segments, overlapping
the rachis throughout the lamina.

192.^. concinnum

8(7). Pinnae decidedly stalked, the basal acroscopic pinnule 5-25 mm from the rachis; rhizome
scales ca. 2.5-5 mm long, 0.4 mm wide..9.

9(8). Rhizomes stout, compact, the stipes fasciculate; ultimate segments cuneate to cuneate-
flabellate, lobed to incised along the outer margin; sori orbicular to suborbicular. Laminae
lanceolate, (10)15-45 cm long, (5)10-15 cm wide, (2)3-pinnate.

210. A raddianum

9(8). Rhizomes slender, short- to long-creeping, the stipes up to 5 mm distant; ultimate segments
mostly orbicular to suborbicular, entire to lobed along the outer margin; sori oblong to lunulate.
Laminae lanceolate, 10-35 cm long, 5 - 20(23) cm wide, 3-pinnate.

2JQ1.A. poiretii

10(5). Laminae 2-pinnate or more divided.. 13.

10(5). Laminae pinnate..!!.

11(10). Lateral pinnae much longer than wide, 6-9 cm long, broadly ovate-lanceolate, glaucous
abaxially; laminae with a conform apical pinna, never flagelliform and rooting at the apex.

211. ^. seemannii

11(10). Lateral pinnae no longer than wide, less than 2 cm long, lunulate, obovate, or nearly
triangular, not glaucous abaxially; laminae usually flagelliform and rooting at the apex, sometimes
with a conform apical pinna.. 12.

12(11). Laminae 4-8 cm wide; sterile pinna margins entire; lateral pinnae lunulate, (1.5)2-3 times
wider than long; rhizome scales ca. 0.4 mm wide, concolorous.

200.^. lunulatum

12(11). Laminae 2-4 cm wide; sterile pinna margins sharply toothed with a vein ending in each
tooth; lateral pinnae obovate or nearly triangular, usually 1-1.5 times wider than long; rhizome
scales ca. 0.15 mm wide, weakly bicolorous.

195. A. deflectens

13(10). Costule apices not swollen, the segments shghtly articulate..l5.

13(10). Costule apices swollen, the segments readily articulate.. 14.

14(13). Lateral pinnules or ultimate segments of pinnules mostly obtuse at the base (up to a 150°
angle); margins of sterile segments entire or nearly so; rhizome scales up to 5(8) mm long, 0.75 mm
wide, concolorous, yellow-brown, sparsely short-ciliate. Laminae ovate-lanceolate, (15)25-60 cm
long, (5)10-45 cm wide.

208. A. princeps

14(13). Lateral pinnules or ultimate segments of pinnules mostly acute at the base; margins of
sterile segments finely serrulate, especially at the apex; rhizome scales up to ca. 3 mm long, 0.5 mm
wide, weakly bicolorous, usually rather dark brown, densely short- to long-ciliate. Laminae ovate-
lanceolate, 10 - 30 cm long, 10 - 25 cm wide.

213.^1. tenenim

15(13). Ultimate segments mostly 2.5-5 cm long, acute or acuminate at the apex.. 17.

15(13). Ultimate segments mostly less than 2 cm long, truncate or round at the apex.. 16.

16(15). Costae and costules without an axillary pale area at the point of their insertion. Laminae
lanceolate or ovate-lanceolate, 3-4-pinnate, 20 - 45 cm long, 10-25 cm wide, distinctly glaucous.

189. ^. andicola

16(15). Costae and costules with an axillary pale area at the point of their insertion. Laminae
lanceolate, 3(4)-pinnate, 20-40 cm long, 15 - 25(30) cm wide, slightly glaucous.

190. A. braunii

17(15). Laminae 1- 2-pinnate; sori commonly 10 mm wide; pinnule margins entire. Laminae
lanceolate, longer than wide, 20 - 30 cm long, 15 - 25 cm wide.

211. >1. seemannii

ADIANTACEAE: 32. ADIANTUM 143

17(15). Laminae 3-4-pmnate; sori 1-2 mm wide; pinnule margins lobed. Laminae very broadly
ovate, about as long as wide, 20 - 40 cm long, 20 - 35 cm wide.

216. >4. trapeziforme

18(4). Laminae 3-5-pinnate at the base. Laminae broadly ovate, sometimes wider than long,
40 - 60 cm long, 40-90 cm wide; axes with rather stiff, multicellular, yellowish hairs and similar

narrow scales.

205. A. pectinatum

18(4). Laminae 1- or 2-pinnate at the base.. 19.

19(18). Rachis covered, mostly adaxially, with short, rather stiff, straight, glandular hairs; scales
absent; sori about as wide as long, on low lobes. Laminae 2-pinnate, broadly ovate, 25 - 35 cm long,
20 - 25 cm wide, usually with 2 pairs of lateral, compound pinnae.

217. A. urophyllum

19(18). Rachis covered, mostly abaxially, with narrow scales often with expanded, laxly toothed or
hairy bases; lax, tv^ted hairs also sometimes present; sori mostly much longer than wide, not on low
lobes..20.

20(19). Rachis scales narrow, ca. 3 mm long, decidedly spreading. Laminae 2-pinnate, ovate to
oblong, 20 - 45 cm long, 20-30 cm wide, with (3)4-7 pairs of lateral, compound pinnae.

194.^. decoratum

20(19). Rachis scales narrow, 1-2.5 mm long, usually not strongly spreading. .21.

21(20). Laminae abaxially glabrous or sparsely covered with narrow scales with expanded and/or

ciliate bases, glaucous or not..23.

21(20). Laminae abaxially sparsely covered with jointed hairs, these not expanded cmd ciliate at the
base..22.

22(21). Laminae glaucous abaxially; pinnules more than half their width distant. Laminae 2-
pinnate, broadly lanceate to broadly triangular, (15)20-30(40) cm long, (7)10 - 25(30) cm wide, with
1-3(4) pairs of lateral, compound pinnae.

197. ^. humile

22(21). Laminae not glaucous abaxially; pinnules less than half their width distant. Laminae 2-
pinnate, ovate to oblong, 15-20 cm long, 12-20 cm wide, with 2-4 pairs of lateral, compound
pinnae.

214.^1. terminatum

23(21). Mature laminae always 2-pinnate with the terminal pinna about the same size as the
(2)3-8(12) pairs of lateral, compound pinnae and usually tapered at the base..29.

23(21). Mature laminae pinnate or 2-pinnate with the terminal pinna larger than the 1-2(3) pairs
of lateral, compound pinnae and usually not tapered at the base..24.

24(23). Sori oblong, several on the acroscopic, outer, and sometimes basiscopic edge of the
rounded or acute pinnae or pinnules..26.

24(23). Sori linear, 1(3) each on the acroscopic and outer or basiscopic edge of the acute to
acuminate pinnae or pinnules..25.

25(24). Pinnae or pinnules lanceate, uniformly tapering to an acute or acmninate apex; laminae
l(2)-pinnate, oblong to lanceate, (15)20-35 cm long, (5)10-20 cm wide, with 0-1(2) pairs of lateral,
compound pinnae.

199.^. lucidum

25(24). Pinnae or pinnules oblong, usually with a distinct outer edge; laminae 2-pinnate, oblong,
ovate-lanceolate, or triangular, 20 - 45 cm long, 20 - 30 cm wide, with 1-4 pairs of lateral, compound
pinnae.

218. ^4. villosum

26(24). Laminae 2-pinnate with (1)2 -3(4) pairs of lateral, compound pinnae; rhizomes long-
creeping with the fronds (8)10-20 mm distant, 1.5 -2.5 mm in diam. including the stipe bases.
Laminae broadly lanceate to broadly ovate, (12)15 - 25(40) cm long, 20-30 cm wide.

198. ^. latifolium

144 ADIANTACEAE: 32. ADIANTUM

Laminae pinnate or 2-pinnate with 1 pair of compound pinnae; rhizomes short-creeping
with the fronds 1-3(5) mm distant, (1)2-5 mm in diam. including the stipe bases..27.

27(26). Proximal simple pinnae truncate and not overlapping the rachis at the acroscopic base,
2.5 - 3.5 times longer than wide; sterile lamina margins biserrate; rhizome scales appressed. Laminae
pinnate and lanceate or 2-pinnate and triangular, 10 - 20(22) cm long, (3.5)6-10(12) cm wide, with
0-1 pair of lateral, compound pinnae, sparsely scaly.

202. A. obliquum

27(26). Proximal simple pinnae cordate to subcordate and overlapping the rachis at the acroscopic
base, (1.5)2 -2.5(3) times longer than wide; rhizome scales spreading.. 28.

28(27). Laminae not glaucous, scaly, the scales cihate at the base; sterile lamina margins usually
slightly biserrate. Laminae pinnate and lanceate or rarely 2-pinnate and triangular, 9 - 26 cm long,
5 - 15 cm wide, with 0 or 1 pair of lateral, compound pinnae.

203. ^1. obliquum X petiolatum

28(27). Laminae glaucous, not scaly; sterile lamina margins uniformly serrate. Laminae pinnate,
ovate, lanceolate, or oblong, 10 - 20(25) cm long, (3)5 - 12 cm wide.

206. A. petiolatum

29(23). Lateral pinnae 1- 2(2.5) cm wide. Laminae ovate-lanceolate to oblong, 20 - 30(40) cm
long, 10 - 25(30) cm wide, with (2)3-6(10) pairs of lateral, compound pinnae; pinnules oblong,
obtuse to round at the apex, fertile on the acroscopic and outer margins, the sori 1- several, the
fertile margin often strongly revolute; rhizomes creeping, 2-5 mm in diam.; stipes 1-10 mm distant.

212. A. serratodentatum

29(23). Lateral pinnae more than (2.2)2.5 cm wide..30.

30(29). Sori on the acroscopic margin of the pinnules (3)5 -many, elongate, usually 2-5 times
longer than wide.. 32.

30(29). Sori on the acroscopic margin of the pinnules 1(3), linear, usually more than 10 times
longer than wide..31.

31(30). Pinnules with a single linear or shghtly curved sorus on the acroscopic margin, almost
completely dimidiate, the midrib close to the basiscopic margin throughout the length of the pinnule.
Laminae broadly ovate-lanceolate to oblong, 20-35 cm long, 15 - 25(30) cm wide, with (2)4-8 pairs
of lateral, compound pinnae.

209. A. pulverulentum

31(30). Pinnules with linear sori on the acroscopic, outer, and often apical portion of the basiscopic
margin, excavate at the basiscopic base but not dimidiate, the midrib central in the pinnule. Laminae
oblong, broadly ovate-lanceolate, or triangular, 20 - 45 cm long, 20-30 cm wide, with 1-4 pairs of
lateral, compound pinnae.

218.^. villosum

32(30). Subapical pinnules ca. 1/2 as long as the median ones; stipes (8)10-20 mm distant.
Laminae broadly lanceate to broadly ovate, (12)15 - 25(40) cm long, 20 - 30 cm wide, with (1)2 -3(4)
pairs of lateral, compound pinnae.

198. A. latifolium

32(30). Subapical pinnules ca. 1/4 as long as the median ones; stipes 1-3 mm distant.. 33.

33(32). Most fertile pinnules with the sterile apex turned toward the pinna apex and acute, the
midrib distinct more than half way to the pinnule apex, submarginal distally, the veins distinctly
prominulous; rhizomes long-creeping, the stipes usually distant. Laminae triangular to broadly
ovate, (15)20 - 35(40) cm long, (12)20-30(40) cm wide, with 2-4 pairs of lateral, compound pinnae.

215. A. tetraphyllum

33(32). Most fertile pinnules with the sterile apex straight and obtuse, the midrib distinct only at
the pinnule base, the veins slightly prominulous; rhizomes short-creeping, the stipes approximate.
Laminae broadly ovate, 25 - 40 cm long, 20-30(40) cm wide, with 3-8(12) pairs of lateral,
compound pinnae.

196. A. fmctuosum

ADIANTACEAE: 32. ADIANTUM 145

189. Adiantmn andicola Liebm. Koegel. Danske Vidensk. Selsk. Skr., Naturvid.
Md. V, 1:266 (repr. 114). 1849.

Adimtum cuneatum var. m^stifoiium Martens & Galeotti, Nouv. M6m. Acad. Roy. Sci. Bruxelles
15:70. 1842, as ''m^stifolia” LECTOTYPE: Pk d’Orizaba, Edo. Veracruz, Mexico, 9500 ft,

Galeotti 6266 (BR not seen; isolectotype K not seen Tryon photo), chosen by Morton (Contr. U. S.
Natl. Herb. 38:216. 1973).

Adiantum tenemm var. dissectum Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles
15:71. 1842. TYPE: Edo. Oaxaca, Mexico, 8000 ft, Galeotti 6361 (BR not seen photo 5063).

Adiantum amabile Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturv. Afd. V, 1:265 (repr. 113).
1849. LECTOTYPE: Between Tonaguia and Roayaga, Edo. Oaxaca, Mexico, Liebmann FI. Mex.
826 (fo/.) (C), chosen by A. R. Smith (H. Chiapas 2:19. 1981).

Adiantum glaucophyllum Hook. Sp. Fil. 2:40. 1851. LECTOTYPE: Chiriqm, “Veraguas” [Pcia.
Chiriqui], Feb 1849, Seemann (K not seen Kew photo), chosen by Leilinger (Proc. Biol. Soc. Wash.
89:704. 1977). This is the specimen illustrated later by Hooker (Icon. Plant. 10:t. 961. 1854).

Adiantum coopen Baker, J. Bot. Brit. For. 25:25. 1887. TYPE: Costa Rica, Cooper (K not seen
Tryon photo fragm NY).

Adiantum palmense Christ, Bull. Soc. Bot. Geneve II, 1:230. 1909. TYPE: La Palma, Pcia. S. Jose,
1800 m, Werckle 17063 (P-Hb. Christ not seen T^on photo).

Adiantum amplum var, concolor Rosenst. Repert. Spec. Nov. Regni Veg. 22:6. 1925. TYPE:
Candelaria, Pcia. S. Jos6, 17(X) m, Brade & Brade 207 (S not seen; isotype UC).

TYPE: Cerro de Zempoaltepec, Edo. Oaxaca, Mexico, 9000 ft, Liebmann PL
Mex, 2203 j FI Mex. 434 (C not seen fragms NY and US).

Plants terrestrial, at 1200 "1800 m elevation, in forests and ravines, from
Monteverde, the Cordillera Central, the Cerro de Piedra Blanca and Cerro
Tablazo (Pcia. S. Jose), and the Cordillera de Talamanca. Also from Mexico to
Nicaragua, Venezuela, and Colombia.

190. _ Adiantum braunii Mett. ex Kuhn, Linnaea 36:75. 1869.

Adiantum braunii var. cuneatum Mett. ex Kuhn, Linnaea 36:75. 1869, as “cuneata.” TYPE:
Mexico, Berlandier 1151 (B not seen).

Adiantum convolutum Fourn. Mexic. PI. 1:127, t. 6. 1872. SYNTYPES: Meaco, Ghiesbreght 406
(P not seen) and Orizaba, Edo. Veracruz, Mexico, Botteri 18 (P not seen).

Adiantum heteroctitum Christ, Bull- Herb. Boissier II, 4:1094. 1904. TYPE: Costa Rica, Werckle in
1903 (P-Hb. Christ not seen photo 2597).

TYPE: Mexico, Kaiwinslq (B not seen; isotype BR not seen).

Plants terrestrial, at 1000" 1100 m elevation, along rivers and streams, from the
Meseta Central. Also from Mexico, Guatemala, El Salvador, Nicaragua,
Venezuela, Colombia, and Ecuador.

191. Adiantum caiyotideum Christ, Bull. Soc. Bot. Gendve II, 1:230. 1909.

TYPE: Costa Rica, Werckle in 1904 (P-Hb. Christ not seen Tryon photo).

Plants terrestrial, at 0 = 100 m elevation, on river banks, from the unlocalized
type locality, the upper Rio Pedregal (Pcia. Los Santos), and near Pacora, the
Tocumen river, and the vicinity of Juan Diaz (all Pcia. Panama). Also from
Mexico and Guatemala.

192. Adiantum condnnum Willd. Sp. PL ed. 4, 5:451. 1810.

Adiantum lutescens Moug. ex Fee, Gen. Fil. [M6m Foug. 5]:119. 1852. TYPE: Edo. Oaxaca,
Mejdco, collector unknown (Hb. Mougeot not seen). Synonymized by Christensen (Ind. Fil. 29.
1905).

146 ADIANTACEAE: 32. ADIANTUM

ADIANTACEAE: 32. ADIANTUM 147

Adiantum condnnum var. integrum T. Moore, Ind. Fil. 23. 1857. TYPE: Based on A. concinnum
var. 0 Hook. (Sp. Fil. 2:42. 1851), nom. illeg., and so based on the basis of that name, which is
Andes of Quito, Pcia. Pichincha, Ecuador, Jamew/i 16 (K not seen).

Adiantum concinnum var. laxum T. Moore, Ind. Fil, 23. 1857. TYPE: Chachapoyas, Depto.
Amazonas, Peru, Mathews 1850 (K not seen). See Hooker (Sp. Fil. 2:43. 1851) for a brief but valid
description and statement of the type.

lAdiantum concinnum var. subscandens Baker, Syn. Fil, 123. 1867. TYPE: Mt. Chimborazo,
Pcia. Chimborazo, Ecuador, Spruce 5706 (K not seen).

LECTOTYPE: Qimana, Edo. Sucre, Venezuela, Humboldt & Bonpland (B-Hb.
Willd. 20099-2 not seen Tryon photo fragm GH), chosen by Tryon (Contr. Gray
Herb. 194:168. 1964).

Plants terrestrial or epipetric, at 0- 1700(2100) m elevation, in disturbed, open
areas on banks, cliffs, and roadsides, from throughout Costa Rica and western
Panama. Also from Cuba, Jamaica, the Lesser Antilles, Mexico to Nicaragua,
Trinidad, Venezuela, and Colombia to Bolivia.

193. Adiantum cordatum Maxon, Amer. Fern J. 21:136. 1931.

TYPE: Near Puerto Obaldia, Com. S. Bias, 0-50 m, Pittier 4297 (US; isotypes
GH, NY).

Plants terrestrial, at O- 100 m elevation, from the Caribbean lowlands of central
Panama. Also from. French Guiana.

194. Adiantum decoratum Maxon & Weath. Amer. J. Bot. 19:165. 1932,

TYPE: Rio Tapia, Pcia. Panama, Standley 26216 (US).

Plants terrestrial, at 0-800 m elevation, along streams, from the Peninsula de
Nicoya and the Pacific lowlands of central Costa Rica, the Atlantic and Pacific
lowlands of Panama, and near Riosucio (Depto. Choco). Also from Mexico to
Honduras and Nicaragua.

195. Adiantum deflectens Mart. Icon. PL Crypt. Bras. 94. 1834.

Adiantum dolabriforme Hook. Icon. PI. 2:t. 191. 1837. TYPE: Brazil, Gardner 55 (K not seen
photo 15142).

Adiantum filiforme Gardn. in Hook. Icon. Pi. 6:t. 503. 1843. TYPE: Near Oeiras, Est. Piaui,
Brazil, Gardner 2391 (K not seen photo 15148).

Adiantum flagellum Fee, Gen. Fil. [Mem. Foug 5|:117. 1852. TYPE: Brazil, collector unknown
(Hb. Mougeot not seen).

Adiantum subaristatum Fee, Crypt. Vase. Br6sil 1:33, t. 8, f. 2. 1869. TYPE: Est. Bahia, Brazil,
Blmchet 2373 (RB not seen).

Adiantum deflectens var. tremulum Hieron. Bot. Jahrb. Syst. 34:487. 1904, as ''tremulaP

LECTOTYPE: Near Colonia Tovar, Edo. Aragua, Venezuela, Fendler 81 (US), chosen by Lellinger
(Proc. Biol. Soc. Wash. 89:703. 1977).

FIGS. 192-2Q3. Adiantum. FIG. 192. Basal portion of pinna of^. concinnum, Smith 48/109. FIG.
193. Frond ofyl. cordatum, Pittier 4297. FIG. 194. Basal portion of median pinnae of A. decoratum,
BioUey M17. FIG. 195. Median pinna of A. deflectens, Lellinger 701. FIG. 196. Median pinnules of
median pinna of^. fructuosum, Biolley. FIG. 197. Basal portion of median pinna of A. humile, Killip
2834. FIG. 198. Basal portion of median pinna of A. latifolium, Maxon 4770. FIG. 199. Median
pinnae of A. lucidum, Correll 12276. FIG. 200. Median pinnae of A. lunulatum, Bartlett & Lasser
16842. FIG. 201. Pinnae of .<4. macrophyilum, Smith 6869. FIG. 2J02. Median pinnae of A. obliquum,
Killip 2902. FIG. 203. Median pinnae of^. obliquum X petiolatum, Pittier 5208.

148 ADIANTACEAE: 32. ADIANTUM

TYPE: Santarem, Est. Pard, Brazil, Martins (M not seen).

Plants epipetric or terrestrial, at 0-100 m elevation, in open areas, from the
Pacific coastal plain of northern Costa Rica, the Canal Zone, and S. Jose Island
(Pcia. Panama). Also from Mexico, Guatemala, Honduras, El Salvador,
Nicaragua, Venezuela, Colombia to Peru, French Guiana, Brazil, and Paraguay.

196. Adiantum fructuosum Poepp. ex Spreng. Syst. Veg. ed. 16, 4:113. 1827.

Adiantum rigidum Link, Fil. Sp. 69. 1841, non K. Presl, 1836. TYPE: Based on material cultivated
in the botanical garden at Berlin, according to Link sent by Poeppig from Chile (B not seen), an
origin which Hooker (Sp. Fil. 2:22. 1858) rightly doubted.

Adiantum pachysorum Reichenb. ex Kunze, Linnaea 21:224. 1848. TYPE: Near Poelebantje,
Surinam, Kappler 1732o (LZ destroyed?).

Adiantum tetraphyllum var. majus Fourn. Mexic. PI. 1:129. 1872. TYPE: Essentially a renaming
of A. fructuosum Kunze, and so based on the type of that name. See Lellinger (Proc. Biol. Soc.
Wash. 89:704. 1977) for details of the typification.

TYPE: Cuba, Poeppig in 1822 (LZ destroyed; isotype B not seen fragm US, L
not seen photo 259).

Plants terrestrial, at 0-1000 m elevation, from throughout the Flora area. Also
from Cuba, Hispaniola, Trinidad, Venezuela, Colombia to Peru, and Brazil.

The distinctions between this species and A. tetraphyllum mentioned in the key
seem to hold for specimens from the Flora area, but specimens from the West
Indies are more difficult to distinguish.

197. Adiantum humile Kunze, Linnaea 9:80. 1834.

Adiantum killipii Maxon & Weath. Amer. J. Bot. 19:166. 1932. TYPE: Anc6n Hill, Canal Zbne,
Killip 2752 (US).

LECTOTYPE: Tocache, Depto. S. Martin, Peru, Aug 1830, Poeppig (W not
seen), chosen by Tryon (Contr. Gray Herb. 194:161. 1964).

Plants terrestrial, at 0-800 m elevation, in forests and ravines, from the Canal
Zone and vicinity and eastern Panama. Also from Belize, Trinidad, Colombia,
Peru, and Surinam.

This species is unique among its group in the Flora area for having true hairs,
rather than narrow scales, on the abaxial surface of the laminae.

198. Adiantum latifolium Lam. Encyc. M6th. 1:43. 1783.

Adiantum denticulatum Swartz, Nov. Gen. Sp. PI. Prodr. 135. 1788, non Burm., 1768, nom. illeg.
TYPE: Jamaica, Swartz (S not seen Maxon photo).

Adiantum datum Desv. Ges. Naturf. Freunde Berlin Mag. 5:327. 1811. TYPE: “Habitat in
Brazil,” Dombey (P-Hb. Juss. Cat. 1421 not seen photo 3153).

Adiantum triangulatum Kaulf. Enum. Fil. 204. 1824. TYPE: Trinidad, Ryan (C not seen Maxon
photo).

Adiantum fovearum Raddi, PI. Bras. Nov. Gen. 1:58, t. 77. 1825. TYPE: Vicinity of Rio de
Janeiro, Est. Rio de Janeiro, Brazil, Raddi (FI not seen).

Adiantum haenkeanum K. Presl, Rehq. Haenk. 1:62. 1825. TYPE: Guayaquil, Pcia. Guayas,
Ecuador, Haenke (PRC not seen; isotypes B not seen, K not seen).

Adiantum laxum Kunze, Linnaea 9:79. 1834. TYPE: Cuba, Poeppig in 1822 (LZ destroyed;
isotype B not seen fragm US).

Adiantum argutum Splitg. Tijdschr. Natuurl. Gesch. Physiol. 7:427. 1840. SYNTYPES: Near
Blaauwe Berg, Surinam, Splitgerber 290 (L not seen photo 193) and 891 (L not seen photo 194).

ADIANTACEAE: 32. ADIANTUM 149

Adiantum lucidum var. bipinnatum Mett. ex Fourn. Mexic. PI. 1:129. 1872. LECTOTYPE:
Teapa, Edo. Tabasco, Mexico, Linden (P not seen photo 2606), chosen by Lellinger (Mem. N. Y.
Bot. Card. 23:13. 1972).

Adiantum triangulatum var. acuminatum Jenm. Bull. Bot. Dept. Jamaica 33:7. 1892. TYPE:
Jamaica, Jenman (NY? not seen).

Adiantum intermedium var. medioximum Christ, Bot. Jahrb. Syst. 24:93. 1897. TYPE: Castleton,
Jamaica, Harris 1554 (P not seen photo 2604).

Adiantum fovearum var. reductum Jenm. Ferns Brit. W. Ind. Guiana 87. 1899. TYPE: Corentyn

River, Guyana, Jenman (NY not seen).

Adiantum fovearum var. major Rosenst. Repert. Spec. Nov. Regni Veg. 21:345. 1925. TYPE:
Morro das Pedros, Munic. Iguape, Est. S. Paulo, Brazil, Erode 7764 (S not seen).

Adiantum latifolium wdx. grisebachianum Domin, Rozpr. Kr^il. Cesk6 Spolecn. Nauk, Tf. Mat.-Pfir.
[Pterid. Dominica] 2: 139. 1929. SYNTYPES: Roseau, Dominica, Domin (PRC not seen); and
Mayagiiez, Puerto Rico, Sintenis PL Portoric. 436b (PRC? not seen).

TYPE: Uncertain, but possibly Brazil, Commerson (P-Hb. Juss. Cat. 1408 not
seen photo 3151), according to the note accompanying Morton’s photograph.
Proctor (FI. Less. Antill. 186. 1977) has designated as the neotype Guadeloupe,
Proctor 20110 (A not seen; isoneotype US).

Plants terrestrial, at 0-100 m elevation, in forests, from the Atlantic coastal
plain of Costa Rica, Panama, and the northern part of the Choco. Also from
throughout tropical America.

One specimen {Tonduz 14561, US) seems to be intermediate between this
species and A. obliquum. It has rhizomes and frond division like those of A.
latifolium, but pinnae and pinnules like those oiA. obliquum.

199. Adiantum lucidum (Cav.) Swartz, Syn. Fil. 121. 1806.

Pteris lucida Cav. Descr. PI. 266. 1801. TYPE: Guaranda, Pcia. Pichincha, Ecuador, Nie (isotype
S not seen). The holotype was not seen by Christensen at MA (Dansk Bot. Ark. 9(3) :22. 1937).

Pteris aspera Poir. Encyc. M6th. 5:713. 1804. TYPE: French Guiana, Leblond (P-Hb. Lam. not

seen).

Adiantum vaiium Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:435. 1810. TYPE: Between Caripe
and Guardia de S. Agustm, Edo. Monagas, Venezuela, Humboldt 453 (B-Hb. Willd. 20106-1 and -2
not seen Tryon photos).

Plants terrestrial, at 0-300 m elevation, in forests, from the lowlands of Panama
and the northern portion of the Choco. Also from Nicaragua, Trinidad and
Tobago, Venezuela, Colombia to Peru, and French Guiana.

200. Adiantum lunulatum N. L. Burm. FI. Indica 235. 1768.

Adiantum philippense L. Sp. PI. 2:1094. 1753, nom. dub. See Verma (Nova Hedwigia 3:463 - 468.
1%1) and Morton (Contr. U. S. Natl. Herb. 38:371. 1974) for the arguments concerning the status of
this name.

Adiantum arcuatum Swartz, Syn. Fil. 122. 1806. TYPE: Based on^. lunulatum sensu Cav. (Descr.
PI. 272. 1801), non L., 1753, which is based on a specimen from the “Marianas, the Philippines and
Mexico,” Nie (MA? not seen). Synonymized by Christensen (Dansk Bot. Ark. 9(3) :25. 1937).

LECTOTTPE: Malabar, India, Burmann (G not seen photo 16782), chosen by
Morton (Contr. U. S. Natl. Herb. 38:370. 1974).

Plants terrestrial or epipetric, at 0-400(800) m elevation, in forests and open
areas, from the Cordillera de Tilardn, the Pacific coastal plain of Costa Rica and
Panama, and the Canal Zone. Also from Cuba, Mexico to Nicaragua, Venezuela,
and Colombia.

150 ADIANTACEAE: 32. ADIANTUM

201. Adiantum macrophyllum Swartz, Nov. Gen. Sp. PI. Prodr. 135. 1788.

Adiantum macrophyllum var. glaucum T. Moore, Card. Chron. Ill, 3:260, f. 126, 127. 1875. TYPE
LOCALITY: Colombia and Peru.

TYPE: Jamaica, Swartz (S not seen; isotype SET not seen).

Plants terrestrial, at 0-900(1400) m elevation, on wet banks and in ravines,
from the wetter parts of Costa Rica, Panama, and the adjacent Choc6. Also from
the Antilles, Trinidad, Mexico to Nicaragua, and tropical South America except
Guyana and French Guiana.

202. Adiantum obliquum Willd. Sp. PI. ed. 4, 5:429. 1810.

Adiantum macropus Schrad. Goett. Gel. Anz. 1824:872. 1824. TYPE: Brazil, Prince Maxmillian
zu Wied-Neuwied (GOET or LE not seen).

Adiantum macrodus Kaulf. ex Kimze, Flora 22(1), Beibl. 42. 1839. TYPE: Ilheus, Est. Bahia,
Brazil, Martiusl or Prince Maxmillian zu Wied-Neuwiedl (not seen).

Adiantum obliquum var. majus Hook. Sp. Fil. 2:8, t. 79A, f. 1. 1851. SYNTYPES: Colombia,
Cuming 1202 (K not seen); Montagne de Mahurie, French Guiana, Leprieur (K not seen); Tanaii,
near Para, Est. Par^, Brazil, Spmce 39 (K not seen); and Guadeloupe, UHerminier (K not seen).

Adiantum obliquum var. bipinnatum Christ, Bull. Herb. Boissier II, 4:1094. 1904. TYPE: Costa
Rica, Werckli (P not seen).

LECTOTYPE: Caracas, Distr. Fed., Venezuela, Bredemeyer (B-Hb. Willd.
20067-1 not seen Tryon photo), chosen by Lellinger (Mem. New York Bot. Gard.
23:13. 1972).

Plants terrestrial, at 0-700(1500) m elevation, in forests, from the Cordillera
Central, the Atlantic coastal plain of Costa Rica, the lowlands of Panama, and the
northern half of the Choco. Also from throughout tropical America.

The rhizome scale cells of this species are shorter than are those of A.
petiolatum or A. obliquum x petiolatum, the rhizome scales are thinner, laxer,
and larger, and the veins of the laminae are more prominulous abaxially.

203. Adiantum obliquum Willd. x petiolatum Desv.

Plants terrestrial, at 0-200 m elevation, from the Canal Zone and vicinity.
Although these specimens have 64 apparently normal spores per sporangium
and are known to be sexual tetraploids, they are intermediate between their
putative parents, sharing the lamina scales of A. obliquum and the pinna shape and
rhizome scales of A. petiolatum. They are also intermediate between these two
species with respect to pinna number and marginal lamina serrations.

204. Adiantum patens Willd. Sp. PI. ed. 4, 5:439. 1810.

TYPE: Caracas, Distr. Fed., Venezuela, Bredemeyer (B-Hb. Willd. 20078 not
seen Tryon photo fragm NY; isotype W not seen).

Plants terrestrial, at 1000-2100 m elevation, in forests and along roadsides and
trails, from the Meseta Central, Cerro Tablazo, and the Cordillera de Talamanca
to Cerro Chirripo. Also from Mexico, Guatemala, Honduras, El Salvador,
Venezuela, and Colombia to Bolivia.

205. Adiantum pectinatum Kunze ex Baker, Syn. Fil. 120. 1867.

SYNTYPES: Brazil, Burchell 7416 (K not seen; isosyntype P not seen photo
2616); and Tarapoto, Depto. S. Martin, Peru, Spruce 4781 (K not seen; isotype P
not seen photo 2615).

ADIANTACEAE: 32. ADIANTUM 151

Plants terrestrial, at 0-500 m elevation, from the Pacific lowlands of central
Costa Rica. Also from Peru, Bolivia, Brazil, and Argentina.

The name A. pectinatum Kunze ex Ettingsh. (Farnkr. Jetztw. 85, t. 45, f. 14-16.
1864) must be considered a nomen nudum because there is no description and the
three figures are coordinate, each being a pinna; therefore there is no “illustration
with analysis” to validate the name.

206. Adiantum petiolatum Desv. Ges. Natuif. Freunde Berlin Mag. 5:326. 1811.

Adiantum oblongatum Schrad. Goett. Gel. Anz. 1824:872. 1824. TYPE: Brazil, Prince Maxmillian
zu Wied-Neuwied (GOET or LE not seen).

Adiantum kaulfussii Kunze, Linnaea 21:221. 1848. LECTOTYPE: Martinique, Sieber FI. Mart.
371 (B not seen), chosen by Tryon (Contr. Gray Herb. 194:149. 1964). The other syntype is:
Beekhuizen Plantation, Surinam, Kegel (LZ destroyed).

Adiantum lucidum var. pinnatum Fourn. Mexic. PI. 1:129. 1872. LECTOTYPE: Teapa, Edo.
Tabasco, Mexico, Linden (P not seen photo 2605), chosen by Lellinger (Proc. Biol. Soc. Wash.
89:704. 1977).

TYPE LOCALITY: French Guiana and Brazil. No specimen at P was seen by
Weatherby or Tryon.

Plants terrestrial, at 0-600 m elevation, in forests, from throughout the Flora
area. Also throughout tropical America.

This species occasionally has long-creeping rhizomes with rather widely spaced
stipes (ca. 1 cm); in this character it mimics A. latifolium. Intermediates exist
between the specimens with ovate or lanceolate fronds bearing few lateral pinnae
and those with oblong laminae bearing many pairs of pinnae. The latter slightly
resembled, ohliquum, which see for a further comment on this species.

207. Adiantum poiretii Wikstr. Kongl. Vetensk. Acad. Handl. II, 12:443. 1826.
Adiantum crenatum Poir. Encyc. M6th. Suppl. 1:137. Sept 1810, non Willd., Mar 1810, nom. illeg.

TYPE: Tristan d’ Acuna, Petit-Thouars (P-Hb. Juss. 1427 not seen).

Adiantum pellucidum Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:72. t. 19.
1842. TYPE: Near Tanetse and Talea, Edo. Oaxaca, Mexico, Galeotti 6461 (BR not seen; isotypes
BR, C, K, LE none seen).

Adiantum gratum F6e, Gen. Fil. [Mem. Foug. 5]:119. 1852. Near Nolasco, Edo. Oaxaca, Mexico,
Galeotti 6542 (presumably P not seen).

TYPE: A renaming of A. crenatum Poir., and so based on the type of that name.
Plants terrestrial or epipetric, at 100-1500(2100) m elevation, in forests and
ravines, from the Cordillera Central, the Meseta Central, and the Cordillera de
Talamanca to Pcia. Chiriqui. Also from Hispaniola, Mexico, Guatemala, El
Salvador, Venezuela, Colombia to Bolivia, Brazil, Paraguay, Uruguay, Argentina,
and Chile.

This species has the stalks of the ultimate segments longer than those of A.
raddianum.

208. Adiantum princeps T. Moore, Gard. Chron. Ill, 4:197, f. 43, 44. 1875.

1 Adiantum amplum K. Presl, Reliq. Haenk. 1:63. 1825. TYPE: Mexico, Haenke (PRC not seen).
Adiantum subtrapezoideum Christ, Bull. Herb. Boissier II, 4:1094. 1904. TYPE: Nicoya, Pcia.
Guanacaste, Pittier 13768 (P-Hb. Christ not seen Tryon photo).

TYPE: From material cultivated by Veitch originally from Colombia (K not
seen Maxon photo).

152 ADIANTACEAE: 32. ADIANTUM

205 A. pectinatum

208 A. princeps

209 A. pulverulentum

ADIANTACEAE: 32. ADIANTUM 153

Plants epipetric or terrestrial, at 0 - 700(1100) m elevation, in forests and
ravines, from the Peninsula de Nicoya, the Cordillera de Tilaran, the Pacific
coastal plain of northern Costa Rica, the Meseta Central, Penonome and vicinity,
and the Canal Zone. Also from Mexico to Nicaragua, Colombia, and Venezuela.

This species may be difficult to distinguish from A. tenerum; the angle of the
pinnule or segment base varies in both species. In Mexican specimens, the
segments of^. princeps are larger than are those of^. tenerum, but this difference
is less marked in Costa Rican material.

209. Adiantem pulveralentum L. Sp. PL 2:1096. 1753, var. pulverulentum.

Adiantum monosoratum Willd. Sp. PI. ed. 4, 5:445. 1810. TYPE: Caracas, Distr. Fed., Venezuela,
Bredemeyer (B-Hb. WiHd. 2CK)87 not seen Tryon photo fragm NY).

Adiantum umbrosum Willd. Sp. PL ed. 4, 5:447. 1810. TYPE: Caracas, Distr. Fed., Venezuela,
Bredemeyer (B-Hb. Willd. 20090 not seen Tryon photo).

Adiantum apiculatum Schrad. Goett. Gel. Anz. 1824:872. 1824. TYPE:Brazil, Prince Maxmillian
zu Wied-Neuwied (M not seen). Synonymized by Christensen (Ind. Fil. 23. 1905).

Adiantum kunzeanum K. Presl, Tent. Pterid. 157. 1836. TYPE: Based on A. pulverulentum sensu
Kunze, which is based on .<4. monosoratum Willd., and so based on the type of that name.

Adiantum pulverulentum var. camptocarpon F6e, Gen. Fil. [M6m. Foug. 5j:114. 1852. TYPE:
“Habitat in America meridionali,” collector unknown (Hb. Mougeot not seen).

Adiantum claussenii Fee, Gen. Fil. [Mem. Foug. 5]: 115. 1852. TYPE: Novo Friburgo, Est. Rio de
Janeiro, Brazil, Claussen (P not seen photo 2618).

Adiantum pulverulentum var. biserratum Hieron. Bot. Jahrb. Syst. 34:489. 1904, as “biserrataP
SYNTYPES: Rio Dagua, Buenaventura region, Depto. El Valle, Colombia, Lehmann 5027 (B not
seen fragm NY); Mount Tolima, Depto. Tolima, Colombia, Mar 1882, Schmidtchen (B not seen);
and near Tacares, Pcia. Alajuela, Nov 1854, Hoffinann (B not seen).

Adiantum pulverulentum var. crenatoserratum Hieron. Hedwigia 48:232. 1909, as ''crenato-

serrataP TYPE: Based on^. monosoratum Willd., and so based on the type of that name.

LECTOTYPE: Plate 47 of Plumieris “Description...”, which is based on a
specimen collected by Plumier on Hispaniola or Martinique, chosen by Proctor
(FL Less. AntilL 2:185. 1977).

Plants terrestrial, at 0-1600(1800) m elevation, from throughout the moist to
wet regions in the Flora area. Also from throughout tropical America.

Proctor (Ferns Jamaica 242. 1985) recognized A. pulverulentum var. caudatum
Jenm. in Jamaica.

210. Adiantum raddianum K. Presl, Tent. Pterid. 158. 1836.

Adiantum cuneatum Langsd. & Fisch. PL Voy. Russes Monde 1:23, t. 26. 1810, non Forst., 1786,
nom. ileg. TYPE: Ilha de Sta. Catarina, Est. Sta. Catarina, Brazil, Langsdorjf (LE not seen Tryon
photo not seen; isotypes B-Hb. Willd. 200% not seen Tryon photo, BM not seen).

Adiantum cuneatum var. angustifolium Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles
15:70. 1842. SYNTYPES: Edo. Veracruz and Edo. Oaxaca, Mexico, Galeotti 6266 (BR not seen)
and 6359 (BR not seen).

Adiantum tinctum T. Moore, Gard. Chron. 1862:932. 1862. TYPE: From material cultivated by
Veitch, originally from Peru (K not seen Tryon photo not seen).

FIGS. 7M-21Q. Adiantum. FIG. 204. Lamina ofA. patens, Valerio 205. FIG. 205. Basal portion of
basal pinnule of median pinna of A. pectinatum, Jimenez 835. FIG. 206. Median pinnae of A.
petiolatum, Brade & Brade 319. FIG. 207. Pinnule of A. poiretii, Chrysler 5339. FIG. 208. Median
pinna of A. princeps, Hinton 15883, Mexico. FIG. 209. Basal portion of median pinna of A.
pulverulentum, Werckle. FIG. 210. Pinnule of^. raddianum, Werckle.

154 ADIANTACEAE: 32. ADIANTUM

Adiantum colpodes T, Moore, Card. Chron. 1865:530. 1865. TYPE: From material cultivated by
Veitch, originally collected by Pearce in Ecuador (K-Hb. T. Moore not seen Tryon photo not seen).

Adiantum rubellum T. Moore, Card. Chron. 1^:866. 1868. TYPE: From material cultivated by
Veitch, originally from Bolivia (K not seen Tryon photo not seen).

Adiantum amabile T. Moore, Card. Chron. 1868:1090. 1868, non Liebm., 1849, nom. illeg. TYPE:
From material cultivated by Veitch, originally collected by Pearce in Peru (K not seen).

Adiantum decorum T. Moore, Card. Chron. 1869:582. 1869. TYPE: From material cultivated by
Veitch, originally collected by Pearce in Peru (K not seen Tryon photo not seen).

Adiantum wagneri Mett. ex Kuhn, Linnaea 36:11. 1869. TYPE: Mt. Pichincha, Pcia. Pichincha,
Ecuador, Wagner (B? not seen).

Adiantum cuneatum var. major Baker in Mart. FI. Bras. 1(2):594. 1870, as ''majus.'" SYNTYPES:
Lagoa Santa, Est. Minas Gerais, Brazil, Warming (K not seen); Caldas, Est. Minas Gerais, Brazil,
Regnell 1 488 (K not seen); Est. Sta. Catarina, Brazil, F. Mueller 249 (K not seen photo 11495).

Adiantum moorei Baker, Gard. Chron. 1873:811. 1873. TYPE: A renaming of A. amabile T.
Moore, and so based on the type of that name.

Adiantum mfopunctatum Mett. ex Kuhn, Jahrb. Konigl. Bot. Gart. Berlin 1:350. 1881. TYPE
LOCALITY: Yungas, Bolivia. The type evidently is D’Orbigny 165 (B not seen; isotype P not seen
Tryon photo not seen), according to Tryon (Contr. Gray Herb. 194:169. 1964).

Adiantum cuneatum var. vastum Rosenst. Hedwigia 43:217. 1904. TYPE: Toledo, Est. Sta.

Catarina, Brazil, Ulbricht 16 (S not seen).

Adiantum werckleanum Christ, Bull. Herb. Boissier II, 4:1093. 1904. TYPE: Costa Rica, Werckle
in 1903 (P-Hb. Christ not seen photo 2648).

Adiantum cuneatum f. elongatum Rosenst. Hedwigia 46:82. 1906, as “elongata.” TYPE: Munic.
Blumenau, Est. Sta. Catarina, Brazil, Haerchen 2a (S not seen).

Adiantum boliviense Christ & Rosenst. in Rosenst. Repert. Spec. Nov. Regni Veg. 5:230. 1908.
TYPE: La Florida near Yanacachi, South Yungas, Depto. La Paz, Bolivia, 1600 m, Buchtien 459 (S
not seen; isotypes P not seen, UC, US).

Adiantum baenitzii Rosenst. Repert. Spec. Nov. Regni Veg. 5:230. 1908. TYPE: Sirupaya near
Yanacachi, S. Yungas, Depto. La Paz, Bolivia, Buchtien 462 (S not seen; isotype UC).

Adiantum cuneatum var. subintegrum Hieron. Hedwigia 48:240, t. 11, f. 14. 1909, as "'subintegral'
TYPE: Near Purac6, Depto. Cauca, Colombia, Stuebel 112 (B not seen).

Adiantum decorum var. quadripinnatum Rosenst. Meded. Rijks-Herb. 19:8. 1913, as

“quadripinnata.” TYPE: Yungas of S. Mateo, Depto. Cochabamba, Bolivia, 1500 m, Herzog 1995 (S
not seen).

Adiantum remyanum Espinosa, Bol. Mus. Nac. Hist. Nat. 15:96, t. 3, 5g, 5h. 1936. TYPE: Based
on^. formosum sensu Remy in Gay (Hist. Fis. PoUt. Chile, Bot. 6:487. 1853), non R. Br., 1810, and
so based on the type of that name, which is Topocalma, Chile, Gay (P not seen).

Adiantum cuneipinnulum Nair & Ghosh, Acta Bot. Indica 2:78. 1974, nom. superfl. TYPE: A
renaming of^. cuneatum Langsd. & Fisch., and so based on the type of that name.

TYPE: A renaming of A. cuneatum sensu Raddi, and so based on t. 78, f. 2 of
Raddi’s “Plantamm Brasiliensium Nova Genera,” which illustrates a specimen
collected by Raddi on Mt. Estrella, Est. Rio de Janeiro, Brazil.

Plants terrestrial, at 1200-1900(2300) m elevation, from the Cordillera Central,
the Fila de Cedral, and the Cordillera de Talamanca. Also from Jamaica,
Hispaniola, the Lesser Antilles, Trinidad, Venezuela, Colombia to Bolivia, Brazil,
Paraguay, and Uruguay.

211. Adiantum seemannii Hook. Sp. Fil. 2:5, t. 81A. 1851.

TYPE: S. Lorenzo, Pcia. Veraguas, Seemann 1124 (K not seen Tryon photo).
Plants terrestrial, at 0-700 m elevation, in forests, from the Peninsula de
Nicoya, the Cordillera de Tilardn, the Cordillera Central, the Meseta Central, the

ADIANTACEAE: 32. ADIANTUM 155

Atlantic lowlands of Costa Rica and Panama, the Peninsula de Osa, the Pacific
lowlands of Panama, and Bahia Solano (Depto. Choc6). Also from Mexico,
Guatemala, Honduras, and Colombia.

Large, mature plants of this species may have either pinnate or 2-pinnate
fronds. The pinnae of the pinnate fronds are larger than the pinnules of the 2-

pinnate fronds.

212. Adiantum serratodentatum Humb. & BonpL ex Willd. Sp. PL ed. 4, 5:445.
1810.

Adiantum obtusum Desv. Ges. Naturf. Freunde Berlin Mag. 5:327. 1811. TYPE: “Habitat in
Guyana?,” collector unknown (P not seen photo 2624).

Adiantum rhomboideum H.B.K. Nov. Gen. Sp. 1:20 (fol. 16). 1816, non Schkuhr, 1805, nom.
superfl, et ffleg. TYPE: A renaming of A. serratodentatum Humb. & Bonpl. ex Willd., and so based
on the type of that name.

Adiantum bonpiandii Desv. M6m. Soc. Linn. Paris 6:309. 1827. TYPE: A renaming of A.
rhomboideum H.B.K., and so b^ed on the type of that name.

Adiantum cassioides Desv. Mem. S(k. Linn. Paris 6:309. 1827. TYPE LOCALITY: “Habitat in
America calidiori.” No type was seen at P by Weatherby (Contr. Gray Herb. 114:17. 1936).

Adiantum proximum Gaud, in Freyc. Voy. Uranie Bot. 403. 1829. TYPE: Rio de Janeiro, Est.
Rio de Janeiro, Brazil, Gaudichaud (P not seen).

Adiantum obtusum var. revolutum Miq. Inst. Versl. Meded. Kon. Nederl. Inst. Wetensch.
1842:188. 1843. TYPE: Surinam, 6ii (U not seen photo 197).

Adiantum kunzei Miq. Inst. Versl. Meded. Kon. Nederl. Inst. Wetensch. 1842:189, fig. 1843.
TYPE: Para, Surinam, Focke 86 (U not seen), seen by Kramer (Acta Bot. Neerl. 3:483. 1954).

Adiantum rhomboideum var. lamm Klotzsch, Linnaea 18:551. 1845. TYPE: Essequibo River,
Guyana, Rich. Schomburgk 266 (B not seen).

Adiantum rhomboideum var. strictum Klotzsch, Linnaea 18:551. 1845. SYNTYPES: Guanaguana,
Edo. Monagas, Venezuela, Moritz 46b (B not seen); Plain of Aragua, Edo. Aragua, Venezuela,
Moritz 163 (B not seen); and Canuku Mountains, Guyana, Rich. Schomburgk 1184 (B not seen).

Adiantum jacobinae F6e, Gen. Fil. [M6m. Foug. 5]:115. 1852. TYPE: Jacobina, Est. Bahia, Brazil,
Blanchet (P not seen photo 2625).

Adiantum obtusum var. majus T. Moore, Ind. Fil. 32. 1857. SYNTYPES: St. Vincent, Guilding (K
not seen); French Guiana, Lepiieur (K not seen); Brazil, Gardner 3350 (K not seen; isosyntype US);
and Bay of Choco, Depto. Narino, Cauca, or Valle, Colombia, Hinds (K not seen).

Adiantum obtusum var. mkrophyllum Fee, Crypt. Vase. Br6sil 1:38. 1869. TYPE: Novo Friburgo,
Est. Rio de Janeiro, Brazil, Claussen (P or RB not seen).

Adiantum rectmgulare Lindm. Ark. Bot. 1:204, t. 9, f. 3. 1903, nom. superfl. TYPE: A renaming,
in effect, of A. kunzei Miquel, and so b^ed on the type of that name.

TYPE: Caripe, Edo. Monagas, Venezuela, Humboldt 450 (B-Hb. Willd. 20088
not seen Tryon photo).

Plants terrestrial, at O-SOO m elevation, in thickets and forests, from Volcan
Orosi (Pcia. Guanacaste), Buenos Aires (Pcia. Puntarenas), and lowland western
Panama. Also from Trinidad, Venezuela, Colombia, Peru, Bolivia, and the
Guianas.

Kramer (Acta Bot. Neerl. 3:483. 1954) considered A, kunzei Miquel to be

varietally distinct from A. serratodentatum, but the presence of intermediates
makes this separation somewhat doubtful, in my opinion.

213. Adiantum teneram Swartz, Nov. Gen. Sp. PL Prodr. 135. 1788.

Adiantum tenerum var. dissectum Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles
15:71. 1842. TYPE: Mexico, Galeotti 6361 (BR not seen).

156 ADIANTACEAE: 32. ADIANTUM

ADIANTACEAE: 32. ADIANTUM 157

Adiantum trapezoides F6e, Gen. Fil. [M6m. Foug. 5]:117. 1852. TYPE: Puente Nacional, Edo.
Veracruz, Mexico, Galeotti 6317 (P not seen).

lAdiantum extensum F6e, M6m. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:72. 1857.

SYNTYPES: Near Orizaba and Huatusco, Edo. Veracruz, Mexico, Schajfner 40 and 41 (P or RB
neither seen).

Adiantum littorale Jenm. Ferns Brit. W. Ind. Guiana 96. 1899. TYPE: Jamaica, Jenman (NY not
seen).

Adiantum tenerum var. obtusissimum Christ, Bull. Herb. Boissier II, 7:266. 1907. SYNTYPES:
Navarro, Pcia. Cartago, 1100 m, Werckle in 1905 (P not seen); and La Palma, Pcia. S. Jose, 1100 m,
Werckle in 1905 (P not seen).

TYPE: Jamaica, Swartz (S not seen Maxon photo; isotype B-Hb. Willd. 10097
not seen Tryon photo).

Plants terrestrial or epipetric, at 0-100 and 1000-1100 m elevation, from the
Rio Navarro 2 km west of the bridge at Troya (Pcia. Cartago), 2 km north of
Puerto Limdn (Pcia. Limdn), near Guaniquito (Pcia. Los Santos), and Juan Diaz
(Pcia. Panama). Also from Florida, the Antilles, Mexico to Honduras, Trinidad,
and Venezuela.

For an additional comment on this species, see A. princeps. All occurrences of
this species may be escapes, rather than naturalized or native populations.

214. Adiantum terminatum Kunze ex Miq. Inst. Versl. Meded. Kon. Nederl. Inst.
Wetensch. 1842:187. 1843.

Adiantum hirtum Klotzsch, Linnaea 18:553. 1845, non Poir., 1810, nom. illeg. TYPE: Kanuku
Mountains, Guyana, Rich. Schomburgk 1444 (K? not seen).

TYPE: Bergendaal, Surinam, Focke (U not seen).

Plants terrestrial, at ca. 100 m elevation, in the Flora area known only from the
Finca Gebrueder Hundrisser, Pcia. Limon {Erode & Erode 746, NY). Also from
Guatemala, Nicaragua, Venezuela, Colombia to Bolivia, Guyana, Surinam, and
Brazil.

The indument of sparse hairs on the abaxial lamina surface in this species is
exactly like that of^. humile; however,^, terminatum is a plant of smaller stature
with smaller pinnules that are less glaucous on the abaxial surface.

215. Adiantum tetraphyllum Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:441. 1810.

Adiantum tematum Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:436. 1810. TYPE: Caripe, Edo.
Monagas, Venezuela, Humboldt 448 (B-Hb. Willd. 20075 not seen Tryon photo).

Adiantum acuminatum Desv. Ges. Naturf. Freunde Berlin Mag. 5:327. 1811. TYPE: Puerto

Rico, Ledm (P not seen Weatherby photo).

Adiantum prionophyllum H.B.K. Nov. Gen. Sp. 1:20 (fol. 16). 1816, nom. superfl. TYPE: A
renaming of A. tetraphyllum Humb. & Bonpl. ex Willd., and so based on the type of that name.

Adiantum tetraphyllum var. microphyllum Sphtg. Tijdschr. Natuurl. Gesch. Physiol. 7:427. 1840.
TYPE: Near Paramaribo, Surinam, Splitgerber (L not seen).

FIGS. 211-219. Adiantum. FIG. 211. Lamina apex of A. seemannii, Skutch 4673. FIG. 212. Basal
portion of median pinna of A. serratodentatum, Hart 35. FIG. 213. Median pinna of A. tenerum,
Burger & Ramirez 4107. FIG. 214. Basal portion of median pinna of A. terminatum, Cuatrecasas
7551, Colombia. FIG. 215. Median pinnules of A. tetraphyllum, Skutch 3633. FIG. 216. Median
pinnules of basal pinna of A. trapeziforme, Alfaro 46. FIG. 217. Basal portion of median pinna of^.
urophyllum, Seemann. FIG. 218. Basal portion of median pinna of A. villosum, Cornman 566. FIG.
219. Frond of^. wilsonii, Wedel 288.

158 ADIANTACEAE: 32. ADIANTUM

lAdiantum tetraphyllum var. macropterum Kunze, Bot. Zeitung (Berlin) 3:284. 1845. TYPE:
Caracas, Distr. Fed., Venezuela, Moritz (LZ destroyed?).

?Adiantum prionophyllum var. curtum T. Moore, Ind. Fil. 35. 1857. SYNTYPES: Pcia.

Esmeraldas, Ecuador, Seemann (K not seen); and Fernando Poo, Vogel (K not seen).

lAdiantum prionophyllum var. subcoriaceum T. Moore, Ind. Fil. 35. 1857. TYPE: Guadeloupe,
UHerminier 4 (K not seen).

Adiantum falcatum Veil. FI. Flumin. ll:t. 96. 1831; Arq. Mus. Nac. Rio de Janeiro 5:455. 1881,
non Swartz, 1802, nom. illeg. TYPE LOCALITY: Vicinity of Rio de Janeiro, Est. Rio de Janeiro,
Brazil.

Adiantum tetraphyllum var. suhsimplex Christ in Krug in Urban, Bot. Jahrb. Syst. 24:95. 1897.
SYNTYPES: Grenada, Eggers 6141b (P not seen) and 6469 (P not seen photo 2638).

Adiantum tetraphyllum var. costaricense Christ in Pitt. Prim. FI. Costaric. 3(1):19. 1901. TYPE:
Tm"s, Pcia. Cartago, 670 m, Tonduz 11313 (BR or P not seen; isotype US).

LECTOTYPE: Near Guanaguana and Caripe, Edo. Monagas, Venezuela,
Humboldt & Bonpland (B-Hb. Willd. 20082-2 not seen Tryon photo), chosen by
Lellinger (Proc. Biol. Soc. Wash. 89:704. 1977).

Plants terrestrial, at 0-1600 m elevation, in forests and ravines, from
throughout the Flora area. Also throughout tropical America.

This species is often confused withal, fructuosum.

216. Adiantum trapeziforme L. Sp. PI. 2:1097. 1753.

Adiantum rhomboideum Schkuhr, Vier Zwanzigste Kl. Linn. Pfl.-Syst. 2:114, t. 122. 1805. TYPE:
Plate 59 of Sloane’s “Voyage...”, which is based on a specimen from Jamaica.

Adiantum eminens K. Presl, Tent. Pterid. 158. 1836. TYPE: Based on^. trapeziforme sensu K.
Presl (Reliq. Haenk. 1:63. 1826), non L., and so based on the type of that name, which is Panama,
Haenke (PRC not seen fragm NY).

Adiantum formosissimum Klotzsch, Liimaea 18:556. 1845, nom. superfl. TYPE: A renaming of A.
rhomboideum Schkuhr, which was cited in synonymy, and so based on the type of that name.

Adiantum trapeziforme var. oblongatum T. Moore, Ind. Fil. 40. 1857. TYPE: Cordillera de

Veracruz, Edo. Veracruz, Mexico, Galeotti 6338 (K? not seen).

Adiantum trapeziforme var. plumieri T. Moore, Ind. Fil. 40. 1857. TYPE: Near Veracruz, Edo.
Veracruz, Mexico, Linden 70 (K? not seen).

LE(2TOTYPE: Plate 59 of Sloane’s “Voyage...”, chosen by Lellinger (Proc. Biol.
Soc. Wash. 89:704. 1977).

Plants terrestrial, at 0-300(1100) m elevation, from the northern and central
Pacific lowlands of Costa Rica and S. Pedro de S. Ramon (Pcia. Alajuela). Also
from C!uba, Jamaica, the Lesser Antilles, and Mexico to Nicaragua.

217. Adiantum urophyllum Hook. Sp. Fil. 2:24, t. 84B. 1851.

Adiantum pilosum Baker, Ann. Bot. (London) 5:207. 1891, non Fee, 1852, nom. illeg. TYPE:
Ocana to Pamplona, Depto. Norte de Santander, Colombia, Kalbreyer 956 (K not seen Tryon photo).

Adiantum kalbreyeri C. Chr. Ind. Fil. 28. 1905. TYPE: A renaming of A. pilosum Baker, and so
based on the type of that name.

Adiantum orosiense Christ, Repert. Spec. Nov. Regni Veg. 8:17. 1910. SYNTYPES: Valverde
Finca, Orosi, Pcia. Cartago, Brade & Brade 16816 (P-Hb. Christ not seen Tryon photo); and Carrillo,
Pcia. S. Jos6, 400 m, Brade & Brade 314 (P not seen).

SYNTYPES: Gorgona Island, Colombia, Barclay (K not seen). Hinds (K not
seen); and Salango, Pcia. Guayas, Ecuador, Seemann (K not seen).

Plants terrestrial, at (200)900-1500 m elevation, in forests, from the Atlantic
slopes of the Cordillera Central, between S. Isidro del General and Dominical
(Pcia. S. Jose), the Cordillera de Talamanca near Boquete, near Sta. Fe (Pcia.

ADIANTACEAE: 32. ADIANTUM 159

Veraguas), and the Mojarras de Tad6 and the Rio Calima (Depto. Choco). Also
from Venezuela and Colombia to Peru.

218. Adiantum villosum L. Syst. Nat. ed. 10, 2:1328. 1759.

Adiantum falcatum Swartz, J. Bot. (Schrader) 1800(2) :82. 1801, non Buchoz, 1785, nom. illeg.
LECTOTYPE: Plate 55, f. 1, of Sloane’s “Voyage...”, chosen by Proctor (Ferns Jamaica 17. 1984),
which is based on a specimen from Jamaica. Buchoz’s name (Jard. Univ. t. 158. 1785) may prove to
be a nom. nud. if the illustration lacks an analysis showing essential characters.

Pteris dolabriformis Poir. Encyc. M6th. 5:722. 1804. TYPE: Santo Domingo, collector unknown
(holotype or fragm P not seen photo 2645).

Adiantum monotis Linnaea 19:684. 1847. TYPE: Mexico, 548 (B? not seen).
Adiantum lanceolatum F6e, Gen. Fil. [M6m. Foug. 5]:115. 1852. TYPE: French Guiana, Leprieur
(Hb. Mougeot not seen).

Adiantum villosum var. macrosorum T. Moore, Ind. Fil. 42. 1857. TYPE: Trinidad, Lockhart (K
not seen).

Adiantum obliquetruncatum F6e, Hist. Foug. Antill. [M6m. Foug. 11]: 18, t. 7, f. 3. 1866.

SYNTYPES: Guadeloupe, L'Herminier (P or RB not seen; isotype P not seen photo 2646); and
Trinidad, Germain in 1862 (P or RB not seen).

NEOTYPE: Near Spanish Town, Jamaica, Sloane, (BM-Hb. Sloane not seen),
chosen by Proctor (FI. Less. Antill. 185. 1977). No specimen of A. villosum

currently is extant in the Linnaean Herbarium. The specimen labelled “Adiant A
villosum” in the hand of Linnaeus (LINN 1252.10 not seen microfiche S. I.
Library), which usually is cited as the type, is A. pulverulentum; according to
Jackson’s “Index” it was added after 1755, and so cannot be a type.

Plants terrestrial, at 0-700 m elevation, in forests and ravines, from the
Peninsula de Nicoya, the western slopes of the Cordillera de Tilaran and the
Cordillera Central, Caldera (Pcia. Puntarenas), the Atlantic coastal plain near
Limon (Pcia. Limon), around Sta. Fe (Pcia. Veraguas) and Penonome, and central
Panama. Also from the Antilles, Mexico to Nicaragua, Trinidad and Tobago,
Venezuela, Colombia, and Brazil.

One specimen {Maxon 5757, US) with atypical pinnules and discontinuous sori
may be a hybrid with >4. ohliquum.

219. Adiantum wilsonii Hook. Sp. Fil. 2:6, t. 72A. 1851.

Adiantum wilsonii var. semicordatum Rosenst. Repert. Spec. Nov. Regni Veg. 22:6. 1925. TYPE:
Llanuras de S. Carlos, Pcia. Alajuela, 200 m, Brade & Erode 473 (S not seen; isotypes NY, UC, US).

TYPE: Near Bath, Jamaica, Wilson (K not seen Tryon photo).

Plants terrestrial, at 0-800 m elevation, in forests, from the Atlantic coastal
plain of Costa Rica and Panama, the Canal Zone and adjacent Pcia. Panama,
Cerro Tacarcuna and central Pcia. Darien, and the northern half of the Choco.
Also from the Greater Antilles, Mexico to Belize, Nicaragua, and Colombia.

Rosenstock’s variety refers to the large form with four or five pairs of lateral
pinnae. This variant is scattered throughout the range of the species, but in
Central America it is predominantly in the southern portion of the range. There
seem to be no substantial characters to separate it from var. wilsonii.

160 VITTARIACEAE: 33. ANTROPHYUM

VITTARIACEAE

Rhizomes bearing abundant, crowded roots, the roots densely covered with
long, matted root hairs, the rhizomes erect to long-creeping, scaly, the scales
narrowly lanceolate, often long-attenuate at the apex, concolorous or nearly so,
strongly clathrate; fronds minute to small, usually lax, thin, and often pendent;
stipes terete to flattened, often obsolete, not articulate; laminae less than 0.5 m
long, simple, often linear or nearly elliptic, entire (lacerate at the apex in
Hecistopteris), lacking sclerenchyma but with inconspicuous spicular, cells present
in the epidermis; veins free and divergent or parallel, or anastomosing m long,
polygonal areolae lacking free included veinlets; sori surficial to submarginal;
mdusia absent; sporangia superficial or immersed in shallow to deep grooves
associated with the veins, rather short-stalked, mixed with paraphyses or the
paraphyses wanting; spores monolete or trilete.

BENEDICT, R. C. 1911. The genera of the fern tribe Vittarieae, their external morphology,
venation, and relationships. Bull. Torrey Bot. Club 38:153-190, t. 2-8.

1. Sporangia acrostichoid, scattered over the abaxial surface of the elliptic to oblanceolate, lax
laminae.

34. Anetium

1. Sporangia in discrete lines following the veins, often in grooves..2.

2(1). Laminae irregularly forked or lacerate at the apex, less than 5 cm long.

35. Hecistopteris

2(1). Laminae entire, more than (5)10 cm long. .3.

3(2). Sori in several lines or in rows following the polygonal areolae on each side of the midrib.

33.Antrophyum

3(2). Sori in a single marginal or submarginal line on each side of the midrib..4.

4(3). Areolae in two or more series parallel to the midrib on each side of the midrib; stipes and
midribs at the base green or stramineous.

36. Ananthaconis

4(3). Areolae in a single series oblique to the midrib on each side of the midrib; stipes and midribs
at the base green or blackish.

37. Vittaria

33. ANTROPHYUM Kaulf.

Plants epiphytic; rhizomes short-creeping, the roots bearing brownish, matted
root hairs, scaly, the scales distinctly clathrate, blackish, sometimes brownish,
sometimes blaclash with brownish margins, often with an extended, filiform apex;
fronds small to medium-sized; stipes narrowly alate or obsolete; laminae simple,
linear to narrowly rhombic, narrowly ovate, or narrowly obovate, often attenuate
at the base, acute or acuminate at the apex, firmly herbaceous to subcoriaceous or
even fleshy; midribs narrow to wide, absent or obscure toward the frond apex in
some species; veins reticulate in long, polygonal areolae without included veinlets,
with short, free, marginal veins in some species, immersed in the lamina tissue;
sori elongate on the veins, forming lines or discontinuous polygons, superficial or
immersed in grooves, exindusiate; sporangia long-stalked, mixed with branched-
clavate paraphyses inv4. ensiforme, otherwise lacking paraphyses.

Tropical and subtropical, mostly in the Old World; ca. 50 species, including
those often included in Poly taenium.

BENEDICT, R. C. 1907. The genus Antrophyum - 1. Synopsis of subgenera, and the American
species. Bull. Torrey Bot. Club 34:445 - 458. 1907.

VITTARIACEAE: 33. ANTROPHYUM 161

TRYON, R. M., Jr. 1964. Taxonomic fern notes. IV. Some American vittarioid ferns. Rhodora
66:110-117.

1. Paraphyses present among the sporangia; spores monolete; soral lines diverging at a 30° -60°
angle to the midrib. Laminae nearly linear, ca. 10 times longer than wide, 10-35 cm long,
10 - 25(30) cm wide; stipes obsolete; midribs prominulous only in the proximal 2/3 -4/5 of the
lamina.

223. ^. ensiforme

1. Paraphyses absent among the sporan^a; spores trilete; soral lines parallel to the midrib or
diverging up to 30° from the midrib.. 2.

2(1). Sporangia in (1)2 -3(4) long, simple lines on each side of and parallel to the midrib, always in
rather deep grooves. Laminae linear, 7-42 cm long, (1)2-11 mm wide; stipes obsolete; midribs
blackish at the base, pale and prominulous to the frond apex.

226. >1. lineatum

2(1). Sporangia in (2)4-10 series of short, angular lines or polygonal areolae on each side of and
not parallel to the midrib (except predominantly parallel in A. lanceolatum), either in shallow
grooves or superficial.. 3.

3(2). Longitudinal axes of the areolae predominantly parallel to the midrib. Laminae linear, 6-31
cm long, (5)7-13(15) mm wide, attenuate at the base, acuminate at the apex; stipes obsolete;
midribs pale and prominulous throughout.

225. >1. lanceolatum

3(2). Longitudinal axes of the areolae predominantly oblique to the midrib.. 4.

4(3). Soral lines all superficial; areolae mostly obtuse at the ends; marginal veins mostly or all free,
not united to form marginal areolae, lacking hydathodes; stipes obsolete..6.

4(3). Soral lines slightly sunken; areolae mostly truncate at the ends; marginal veins mostly
anastomosing in short areolae, with few or no free marginal veinlets, lacking hydathodes; stipes
obsolete or not. Rhizome scales with a long, filiform apex; midribs prominulous nearly to the frond
apex..5.

5(4). Laminae narrowly to rarely broadly oblanceolate, widest distal to the middle, strongly alate to
the base, the ala wider than the width of the stipe. Laminae (6)10-38 cm long, 1.5-3.5(4.5) cm
wide; stipes obsolete; frond margins decidedly revolute at maturity.

221. A. cajenense

5(4). Laminae narrowly elliptical, widest at the middle, scarcely alate to the base, the ala narrower
than the width of the stipe. Laminae 7-31 cm long, (1.5)2 -5 cm wide; stipes obsolete or nearly so;
frond margins slightly or not revolute at maturity.

224. yl. guayanense

6(4). Laminae elliptic-lanceolate; midribs prominulous only in the proximal Il2>i2l?>) of the
laminae. Laminae 9-31 cm long, 2.5 -4.5 cm wide.

220.^. anetioides

6(4). Laminae oblong or narrowly oblanceolate; midribs prominulous in the proximal 9/10(2/3) of
the lamina. Laminae (3)6-20 cm long, 1.2 -3(3.5) cm wide.

222. A. chlorosporum

220. Antrophyum anetioides Christ, Bull. Herb. Boissier II, 5:12. 1905.

LECTOTYPE: Rio Las Vueltas, Tucurrique, Pcia. Cartago, 635 m, Tonduz
12757 (P not seen; isotypes NY not seen, US), chosen by Benedict (Bull. Torrey
Bot. Club 34:455. 1907).

Plants epiphytic, at 500-700 m elevation, in forests, from the valley of the Rio
Reventazon.

162 VITTARIACEAE: 33. ANTROPHYUM

VITTARIACEAE: 33. ANTROPHYUM 163

221. Antrophyum cajenense (Desv.) Spreng. Syst. Veg. ed. 16,4:67. 1827.

Hemionitis cajenensis Desv. Ges. Naturf. Freimde Berlin Mag. 5:311. 1811. TYPE: French
Guiana, collector unknown (P-Hb. Desv. not seen photo 3597; possible isotype [French Guiana, /.
Martin] B-Hb. Willd. 19560 not seen microfiche S. I. Library).

Antrophyum discoideum Kunze, Bot. Zeitung (Berlin) 6:702. 1848. LECTOTYPE: Colombia,
Karsten 30 (B not seen Tryon photo GH not seen), chosen by Tryon (Rhodora 66: 112. 1964).

Antrophyum subsessile var. elongatum Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:208 (repr.
290). 1864, as "'elongata^ LECTOTYPE: Based on Antrophyum discoideum Kunze, and so based
on the lectotype of that name.

Antrophum lacantunense Rovirosa, Pteridogr. Sur M6xico 240, t. 38A, f. 10-11. 1910. TYPE: Rio
Lacanja, Edo. Chiapas, Mexico, Martinez (Hb. Rovirosa 1097 not seen; isotype GH).

Plants epiphytic, at 0-1700 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, the Atlantic coastal plain of Costa Rica and
Panama, the Cordillera de Talamanca, the vicinity of S. Isidro del General, the
Fila Costeha near S. Vito, the Peninsula de Osa, and the northern part of the
Choco. Also from Cuba, Jamaica, Mexico, Guatemala, Honduras, Nicaragua,
Venezuela, Colombia to Bolivia, Surinam, and Brazil.

222. Antrophyum chlorosponim Mickel & Beitel, Pterid. FI. Oaxaca 41, f. 42K,L.
1988.

TYPE: Ridge between Yetzelalag and Lovani, Edo. Oaxaca, Mexico,
3600-5800 ft, Hallberg 1546 (NY not seen; isotype UC).

Plants epiphytic, at 1300-1700 m elevation, from the Cordillera Central, the
vicinity of Tapanti, and the Cordillera de Talamanca near the Panama border
(Pcia. Limon). Also from Mexico.

223. Antrophyum ensiforme Hook, in Benth. PL Hartw. 73. 1841.

Antrophyum falcatum Martens & Galeotti, Nouv. M6m. Acad. Roy. Sci. Bruxelles 15:49, t. 12.
1842, non Blume, 1828, nom. illeg. TYPE: Llano Verde, Edo. Oaxaca, Mexico, Galeotti 6385 (BR
not seen).

Antrophyum camosum Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Aid. V, 1:161
(repr. 9). 1849. TYPE: A renaming oi Antrophyum falcatum Martens & Galeotti, and so based on
the type of that name.

Antrophyum galeottii Fee, Hist. Antroph. [Mem. Foug. 4]:51, t. 5, f. 4. 1852. TYPE: Comatlepec,
Edo. Oaxaca, Mexico, Galeotti 6385 (BR not seen).

TYPE: Mt. Totontepeque, Edo. Oaxaca, Mexico, 10000 ft, Hartweg 522 (K not
seen).

Plants epiphytic, at (700)1000-2500 m elevation, in forests, from the Cordillera
de Tilaran, the Cordillera Central, Cerro Tablazo, Cerro Carpintera, the northern
end of the Cordillera de Talamanca, and around Cerro Punta and Boquete. Also
from Mexico to Nicaragua.

224. Antrophyum guayanense Hieron. Hedwigia 57:212. 1915.

LECTOTYPE: Trinidad, Fendler 151 (B not seen; isolectotype US), chosen by
Tryon (Rhodora 66:113. 1964).

FIGS. 220 -226. Antrophyum. FIG. 220. Plant of A. anetioides, Tonduz 12757. FIG. 221. Plant of A.
cajenense^ Alfaro 75. FIG. 222. Plant of A. chlorosporum, Standley 37897. FIG. 223. Plant of A.
ensiforme^ Lellinger 1673. FIG. 224. Plant of A. guayanense^ LeUinger & de la Sota 420. FIG. 225.
Plant of^. lanceolatum, de la Sota 5229. FIG. 226. Plant of .,4. lineatum, Maxon 315.

164 VITTARIACEAE: 34. ANETIUM

Plants epiphytic, at 100 - 300 m elevation, in forests, in the Flora area known
only from the Mojarras de Tado, 8.5 km east of Istmina, Depto. Choco (Lellinger
Sl de la Sota 420, COL, CR, HUA, LP, US). Also from Trinidad, Venezuela,
Colombia to Peru, Guyana, French Guiana, and Brazil.

225. Antrophyum lanceolatum (L.) Kaulf. Enum. Fil. 198. 1824.

Hemionitis lanceolata L. Sp. PI. 2:1077. 1753, non Polytaenium lanceolatum (Swartz) Desv., 1827.
TYPE: Plate 127C of Plumier’s “Traite...”, which illustrates a plant collected by Plumier on St.
Vincent.

Antrophyum feel Schaffn. ex F6e, M6m. Soc. Sci. Hist. Nat. Strasbourg 5 [M6m. Foug. 7]:42, t. 22, f.

I. 1857. TYPE: Near Huatusco, Edo. Veracruz, Mexico, Schajfner 133 (P or RB not seen; isotype K
not seen).

Antrophyum stenophyllum Rovirosa, Pteridogr. Sur M6xico 242. 1910, non Baker, 1898, nom. illeg.
TYPE: Cerro de Moyos, Rio Lacanj^ region, Edo. Chiapas, Mexico Martinez (Hb. Rovirosa 1104 not
seen).

Plants epiphytic, at 0-700(1400) m elevation, in forests, from the Cordillera de
Tilaran, Vara Blanca (Pcia. Heredia), the Atlantic coastal plain of Costa Rica, the
Peninsula de Osa, the Canal Zone, and Alto del Buey. Also from the Antilles,
Mexico, Guatemala, Honduras, Nicaragua, Trinidad, Colombia, Ecuador,
Surinam, French Guiana, and Brazil.

226. Antrophyum lineatum (Swartz) Kaulf. Enum. Fil. 199. 1824, var. lineatum.

Hemionitis lineata Swartz, Nov. Gen. Sp. PI. Prodr. 129. 1788, non Vittaria lineata (L.) J. E. Smith,
1793. TYPE: Jamaica, Swartz (SBT not seen; isotype B-Hb. Willd. 20033 not seen microfiche S. I.
Library).

Vittaria lanceolata Swartz, Ges. Naturf. Freunde Berlin Neue Schriften 2:133, t. 7, f. 2. 1799.
TYPE: A renaming of Hemionitis lineata Swartz, and so based on the type of that name.

Plants epiphytic, at 0-2400 m elevation, in forests, from the Cordillera Central,
Cerro Tablazo, Cerro Carpintera, the Cordillera de Talamanca, near Sta. Fe, near
El Valle, and the Canal Zone and adjacent Panama. Also from the Greater
Antilles, Mexico, Guatemala, Belize, Nicaragua, Venezuela, Colombia to Bolivia,
Brazil, and Argentina.

Panamanian specimens from near sea level are very small. Proctor (Amer. Fern

J. 72:114. 1982) has distinguished specimens from Jamaica as Polytaenium

lineatum var. intramarginale (Baker ex Jenm.) Proctor.

34. ANETIUM Splitg.

Plants epiphytic; rhizomes long-creeping, slender, clothed with roots bearing
abundant, brownish, matted root hairs and abundantly scaly, the scales dark
brown, iridescent, clathrate; fronds small; stipes short to obsolete; laminae distant,
simple, elliptic to oblanceolate, nearly attenuate at the base, acute to acuminate at
the apex, papyraceous to herbaceous; midribs narrow, not extending to the lamina
apex; veins reticulate in long, polygonal areolae without included veinlets, but with
short, free, marginal veins scarcely or not hidden in the lamina tissue; sporangia
superficial, scattered mostly along but also between the veins, not orgamzed into
sori, exindusiate; sporangia long-stalked.

New World lowland tropics; monotypic.

VITTARIACEAE; 35. HECISTOPTERIS-36. ANANTHACORUS 165

227. Anetium citrifolium (L.) Splitg. Tijdschr. Natuurl. Gesch. Physiol. 7:395.

1840.

Acrostichum citrifolium L. Sp. PI. 2:1067. 1753. TYPE: Plate 116 of Plumier’s “Trait6...”, which
illustrates a specimen collected by Plumier on Martinique.

Hemionitis parasitica L. Syst. Nat. ed. 10, 2:1322. 1759. TYTE: Jamaica, Browne, a specimen
labelled “Hemionitis A parasitica” in the hand of Solander (LINN 1248.2 not seen microfiche S. I.
Library).

Antrophyum pendulum LePrieur in F6e, Hist. Antroph. [M6m. Foug. 4]:51. 1852. SYNTYPES:
French Guiana, LePrieur 1830 (P or RB not seen) and 1859 (P or RB not seen).

Plants epiphytic, at 0-700 m elevation, in forests, from the Cordillera de
Tilar^n, the valley of the Rfo Reventazdn, the Atlantic lowlands of Costa Rica and
Panama, the Peninsula de Osa, the Canal Zone, and the foot of Alto del Buey and
near La Teresita (Depto. Choc6). Also from Jamaica, Hispaniola, Puerto Rico,
the Lesser Antilles, Mexico to Honduras, Nicaragua, Trinidad and Tobago,
Venezuela, Colombia, Peru, Bolivia, the Guianas, and Brazil.

35. HECISTOPTERIS J. Smith

Plants epiphytic; rhizomes erect or ascending, minute, borne at intervals along
long-creeping, proliferous roots bearing conspicuous root hairs, densely scaly, the
scales dark brown, clathrate; fronds very small; stipes obsolete; laminae simple,
roughly spathulate, glabrous, irregularly forked or lacerate at the apex, the lobes
somewhat lacerate; veins free, flabellate; sori elongate, superficial on the veins in
the middle portion of the lobes, exindusiate; sporangia short-stalked, mixed with
reddish-brown, multicellular, catenate paraphyses.

New World tropics; monotypic.

228. Hecistopteris pumila (Spreng.) J. Smith, London J. Bot. 1:193. 1842.

Gymnogramme pumila Spreng. Tent. Suppl. Syst. Veg. ed. 16. 31. 1828. TYPE: Surinam, Weigelt

(LZ destroyed; isotype B not seen; probable isotype UC).

Hecistopteris pumila var. ohtusa Maxon & Morton, Bull. Torrey Bot. Club 765: 75:76. 1948.

TYPE: Tafelberg Creek, Saramacca River headwaters, Surinam, Maguire 24101 (NY).

Plants epiphytic, at 0-300(1000) m elevation, in forests, from Volcan Arenal,
the Atlantic coastal plain of Costa Rica and Panama, near S. Isidro del General,
the Peninsula de Osa and vicinity, the Canal Zone and adjacent areas, and the
northern part of the Choc6. Also from the Antilles, Mexico to Honduras,
Trinidad, Venezuela, Colombia to Peru, Surinam, French Guiana, and Brazil.

Christ (Hedwigia 44:366. 1905) named three subspecies of H. pumila, but the
names are all illegitimate because they were proposed in binomial form, contrary
to the “International Code of Botanical Nomenclature.”

36. ANANTHACORUS Underw. & Maxon

Plants epiphytic; rhizomes short-creeping, densely clothed with roots bearing
abundant, brownish, matted root hairs, scaly, the scales brown or rarely blackish,
clathrate; fronds medium-sized; stipes approximate, nearly exalate, often nearly
obsolete; laminae simple, linear, glabrous, coriaceous, the margins plane to
subrevolute, the midrib prominent to the frond apex on the abaxial surface of the
laminae; veins obscure, anastomosing, forming elongate, polygonal areolae
without included veinlets in a few series on each side of the miorib; sporangia
forming a continuous, submarginal coenosorus in a shallow groove along a

166 VITTARIACEAE: 36. ANANTHACORUS

.1 cm

VITTARIACEAE: 37. VITTARIA 167

marginal commissural vein, exindusiate; sporangia short-stalked, mixed with
reddish-brown, branched-clavate paraphyses.

MAXON, W. R. 1908. A new genus allied to Vittaria. Contr. U. S. Natl. Herb. 10:486 - 487.

New World tropics; monotypic, but related closely to Vittaria.

229. Ananthaconis angustifolius (Swartz) Underw. & Maxon, Contr. U. S. Natl.
Herb. 10:487. 1908.

Pteris angustifolia Swartz, Nov. Gen. Sp. PI. Prodr. 129. 1788, non Vittaria angustifolia Blume,
1828. TYPE: Probably Jamaica, Swartz (S not seen; cf. FI. Ind. Occ. 3:1599. 1806).

Vittaria costata Kunze, Linnaea 9:77. 1834. TYPE: Tocache, Huallaga, Depto. S. Martin, Peru,
June 1830, Poeppig (LZ destroyed; isotype W not seen).

Plants epiphytic, at 0-1000(2100) m elevation, in forests, from the Cordillera de
Tilardn, the Cordillera Central, the Atlantic coastal plain of Costa Rica and
Panama, the valley of the Rio Reventazdn, the Peninsula de Osa, northwest of Sta.
Fe, the Canal Zone and vicinity, S. Jose Island (Pcia. Panama), near Cana, and the
Rio Ciego, Cabo Corrientes, the Rio Tmando, and the Rio Baudo (all Depto.
Choco). Also from throughout tropical America.

This species resembles Vittaria remota, but has thicker laminae with the
venation more hidden and with several series of areolae between the midrib and
the frond margin and parallel to the midrib. This genus differs from Vittaria in
having areolate venation that is akin to the venation oiAntrophyum.

37. VITTARIA J. E. Smith

Plants epiphytic; rhizomes erect or short-creeping, covered with roots bearing
conspicuous, abundant, brownish root hairs, scaly at the apex, the scales
lanceolate, strongly clathrate, often iridescent; fronds mostly medium-sized; stipes
short to obsolete, blackish or pale at the base; laminae simple, narrow, linear in
most species, pendent, mostly firmly herbaceous to coriaceous; midribs mostly
prominulous; veins forming a single series of usually oblique areolae between the
midrib and the margin, immersed in the lamina tissue; sori forming a single
submarginal line on each side of the midrib, in a shallow to deep groove,
exindusiate; sporangia long-stalked, mixed with paraphyses.

Tropics and subtropics, mostly in the Old World; ca. 50 species.

BENEDICT, R. C. 1914. A revision of the genus Vittaria J. E. Smith. I. The species of the subgenus
Radiovittaria. Bull. Torrey Bot. Club 41:391-410.

TRYON, R. M., Jr. 1964. Taxonomic fern notes. IV. Some American vittarioid ferns. Rhodora
66:110-117.

1. Laminae (2.5)3-17 mm wide; stipes brown or black toward the base; soral paraphyses bulbous
or swollen at the apex, not twisted; rhizomes radial, the fronds arising from all sides..5.

1. Laminae l-2.5(3.5) mm wide; stipes pale toward the base; soral paraphyses slender to clavate,
usually twisted; rhizomes flattened, the fronds all arising from the sides.. 2.

FIGS. 227 - 233. Anetium, Hecistopteris, Ananthacorus, and Vittaria. FIG. 227. Plant of Anet.
citrifolium, Lankester. FIG. 228. Plant of H. pumila, Lellinger 687. FIG. 229. Plant and lamina
venation of Ananth. angustifolius^ Mickel 3342. FIG. 230. Plant and rhizome scale detail of V.
costaricensis^ Standley & Valerio 44805. FIG. 231. Plant of V. dimorpha, Maxon 60. FIG. 232. Plant
of V. gardneriana, Maxon 5676. FIG. 233. Plant and rhizome scale detail of V graminifolia, Svenson
330.

168 VITTARIACEAE: 37. VITTARIA

2(1). Rhizome scales ca. 0.75-1 mm (8-20 cells) wide, the central cells 1.5-3 times longer than
wide..4.

2(1). Rhizome scales ca. 0.25-0.5 mm (4-6 cells) wide, the central cells 2-10 times longer than
wide. Rhizome scales with all clathrate walls of equal thickness, blackish in mass; stipes obsolete or
nearly so; laminae 1-2.5 mm wide, 15-60 cm long; paraphyses slender, often branched with the
branches linear to slightly clavate, tan to reddish brown.. 3.

3(2). Spores monolete.

234. V. lineata

3(2). Spores trilete.

231. V. dimorpha

4(2). Laminae 1-2.5 mm wide, usually strongly revolute at maturity; central cells of the rhizome
scales strongly clathrate, often the walls of the marginal cells less clathrate than those of the central
ones; rhizome scales several cells wide to the apex or with a short tip 1 cell wide, grayish or dark
golden brown in mass. Stipes obsolete or nearly so; laminae 6-30 cm long, paraphyses stout,
distinctly clavate, becoming spoon-shaped by collapse of the terminal cell, reddish-brown or darker;
spores trilete.

233. V. graminifolia

4(2). Laminae 3-3.5 mm wide, slightly revolute at maturity; central cells of the rhizome scales
weakly and evenly clathrate throughout; rhizome scales 1 cell wide at and proximal to the apex,
reddish-brown in mass. Stipes obsolete; laminae ca. 60 cm long, the midrib prominulous toward the
base, obscure distally, the soral line supramedial; paraphyses slender, slightly clavate; spores
monolete.

230. V. costaricensis

5(1). Fronds 3-6 cm long, 4-8 mm wide, narrowly oblanceolate, exstipitate, membranaceous;
soral lines medial.

235. V. mm/ma&a+20V

5(1). Fronds 8-70 cm long, 3-17 mm wide, linear to narrowly elliptic, short- to long-stipitate,
papyraceous; soral lines submarginal.. 6.

6(5). Laminae (8)10-17 mm wide, linear to narrowly elliptic, the areolae obhque to the midrib.
Rhizome scales several cells wide nearly throughout, narrowly lanceolate, ca. 1-2 mm long, 0.3 -0.5
mm wide; stipes 0.5-2 cm long; midribs prominulous.

236. V. remota

6(5). Laminae (3)4-8 mm wide, linear, the areolae more or less parallel to the midrib.. 7.

7(6). Rhizome scales 1 cell wide except at the base, almost hair-like. Rhizome scales ca. 1-3 mm
long, 0.1 mm wide; stipes 2-6 cm long; fronds 20 - 60(75) cm long, 3-8 mm wide; midribs not
prominulous.

237. V. stipitata

7(6). Rhizome scales several cells wide nearly throughout, linear to lanceolate.. 8.

8(7). Rhizome scales narrowly lanceolate, ca. 2-3 mm long, 0.25-0.5 mm wide, appearing
toothed, the end walk of marginal ceUs laterally produced; stipes 0.2- 1.5 cm long; fronds 10-20(30)
cm long, (3)4-7 mm wide; midribs not or scarcely prominulous.

232. V. gardneriana

8(7). Rhizome scales linear, ca. 2-5 mm long, 0.2 -0.3 mm wide, nearly entire; stipes 5-8 cm
long; fronds 15-60 cm long, 4-6 mm wide; midribs not prominulous.

235a. V. moritziana

230. Vittaria costaricensis Lellinger, Proc. Biol. Soc. Wash. 98:389, 1. 18. 1985.

TYPE: El Silencio near Tilaran, Pcia. Guanacaste, ca. 750 m, Standley & Valerio
44805 (US).

Plants epiphytic, at 700-800 m elevation, in forests, known only from the type.

VITTARIACEAE: 37. VITTARIA 169

231. Vittaria dimorpha K. Muell. Bot. Zeitung (Berlin) 12:547, 1. 13, f. 3. 1854.

Vittaria bradeorum Rosenst. Repert. Spec. Nov. Regni Veg. 22:18. 1925. TYPE: Tablazo, Pcia. S.
Jose, Brade & Brade 165 p. p. (S not seen photo 6104; isotypes NY, UC).

TYPE: Near Jalapa, Edo. Veracruz, Mexico, Schiede 791 (B not seen; isotype
NY not seen).

Plants epiphytic, at (0)600-2300 m elevation, in forests, from Cerro Tablazo,
the vicinity of Cartago, the Rio Matina (Pcia. Limon), and the vicinity of Boquete.
Also from Mexico, Guatemala, El Salvador, and Nicaragua.

This species seems to differ from V. lineata only in having trilete, rather than
monolete, spores, and so its status as an independent species needs to be
investigated.

232. Vittaria gardneriana F^e, Hist. Vittar. Pleurogr. [M^m. Foug. 3]:15, t. 3, f. 1.
1852.

Vittaria karsteniana Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:207 (repr. 289). 1864.

LECTOTYPE: San Pedro, Pcia. Ocana, Colombia, Schlim 318 (B not seen; isolectotype US), chosen
by Lellinger (Mem. New York Bot. Card. 23:18. 1972).

Vittaria gracilis Moritz ex Kuhn, Linnaea 36:67. 1869. TYPE: Colonia Tovar, Edo. Aragua,
Venezuela, Moritz 464 (B not seen; isotype L not seen photo 2366).

Vittaria bommeri Christ, Bull. Herb. Boissier II, 5:11. 1905. SYNTYPES: Costa Rica, Werckle 217
(P not seen); and the valley of the Rio Socio, Costa Rica, Pittier 576a (P not seen).

Vittaria gardneriana var. stenolepis Rosenst. Hedwigia 46:149. 1907. SYNTYPES: JoinviUe, Est.
Sta. Catarina, Brazil, E. O. Mueller 3a (S not seen); and Pirabeiraba, Est. Sta. Catarina, Brazil,
Schmalz 41.1 (S not seen).

LECTOTYPE: Organ Mountains, Est. Rio de Janeiro, Brazil, Gardner 147 (BM
not seen; isolectotypes B not seen, US), chosen by Benedict (Bull. Torrey Bot.
Club 41:401. 1914).

Plants epiphytic, at 1400-2000 m elevation, in forests, from the Cordillera
Central, Cerro Carpintera, above Boquete, and the Serrarua de Pirre (Pcia.
Darien). Also from Hispaniola, Venezuela, Colombia to Peru, and Brazil.

233. Vittaria graminifolia Kaulf. Enum. Fil. 192. 1824.

Vittaria filifolia Fee, Hist. Vittar. Pleurogr. [Mem. Foug. 3]:20, t. 3, f. 6. 1852. LECTOTYPE:
Guadeloupe, UHerminier (P-Hb. Cosson not seen; isolectotype NY not seen), chosen by Tryon
(Rhodora 66:114. 1964).

7 Vittaria setacea Christ, Bull. Herb. Boissier II, 6:47. 1906. TYPE: Navarro, Pcia. Cartago, Werckle
in 1905 (presumably P-Hb. Christ not seen photo 22050; presumable isotypes B, NY neither seen).

TYPE: Brazil, comm. Kaulfuss in 1827 (LZ destroyed; isotype E not seen),
examined by Tryon (Contr. Gray Herb. 194:215. 1964).

Plants epiphytic, at 0-2900 m elevation, in forests, from Monteverde, Alajuela
(Pcia. Alajuela), the Cordillera Central, the Atlantic coastal plain of Costa Rica,
Cerro Carpintera, the Cordillera de Talamanca to Pcia. Chiriqui, Barro Colorado
Island (Canal Zone), Cana, and Bahia Solano. Also from throughout tropical
America.

Vittaria setacea may be a synonym of V. lineata; cf. Mickel & Beitel (Pterid. FI.
Oaxaca 400. 1988).

234. Vittaria lineata (L.) J. E. Smith, M^m. Acad. Roy. Sci. (Turin) 5:421. 1793.

Pteris lineata L. Sp. PI. 2:1073. 1753. LECTOTYPE: Plate 143 of Plumier’s “Traite...”, which
illustrates specimens from Hispaniola, chosen by Tryon (Contr. Gray Herb. 194:213. 1964).

1 cm

170 VITTARIACEAE: 37. VITTARIA

242 P. grandifolia

*

VITTARIACEAE: 37. VITTARIA 171

VittariafiliformisC&v. Descr. PL 270. 1801. TYPE: Peru, (MA not seen photo US).

Vittaria angustifrons Michx. FI. Bor.-Amer. 2:261. 1803. TYPE: Asiahatcha River, Florida,

Michaux (P not seen photo 3356; isotypes B-Hb. Willd. 20025-2 not seen Tryon photo, P-Hb. Juss.

Cat. 1351 not seen photo 3107).

Vittaria schkuhrii Raddi, PL Bras. Nov. Gen. 1:51. 1825. TYPE: Corcovado, Est. Rio de Janeiro,
Brazil, Raddi (FI not seen).

Vittaria costata Kunze, Linnaea 9:77. 1834; Analecta Pteridogr. 29, 1. 18, f. 2. 1837. TYPE: Mision
Tocache, upper Rio Huallaga, Depto. S. Martin, Peru, June 1830, Poeppig (LZ destroyed).

Vittaria deppeana K, Muell. Bot. Zeitung (Berlin) 12:547. 1854. TYPE: Jalapa, Edo. Veracruz,
Mexico, Schiede 790 (B not seen fragm NY not seen; isotype NY not seen).

Vittaria pachydictyon K. Muell. Bot. Zeitung (Berlin) 12:547, t. 13, f. 4. 1854. TYPE: Surinam,
Weigelt (B not seen).

Plants epiphytic, at 0-3300 m elevation, in forests, from the Cordillera de
Tilar^n, the Cordillera Central, the Atlantic coastal plain of Costa Rica and
Panama, the Cordillera de Talamanca to Villa Mills (Depto. Cartago), Golfo
Duke (Pcia. Puntarenas), the Canal Zone, around Porto Bello (Pcia. Colon), S.
Jose Island (Pcia. Panama), near Cana, and La Teresita (Depto. Choco). Also
from throughout tropical America.

235. Vittaria minima (Baker) Bened. Bull. Torrey Bot. Club 38ri64. 1911.

Antrophyum minimum Baker, Ann. Bot. (London) 5:488. 1891. TYPE: Mountains of Costa Rica,
4000 ” 5000 ft, Endres in 1869 (K not seen fragm NY).

Antrophyum werckleanum Christ, Bull. Herb. Boissier II, 5:11. 1905. TYPE: Costa Rica, Werckle
in 1903 (P not seen fragm NY).

Plants epiphytic, at 900-1900 m elevation, in forests, from the Cordillera
Central, Cerro Carpintera, the vicinity of S. Isidro del General, the Fila Costeha
near S. Vito de Java, and the Fortuna dam site (Pcia. Chiriqui). Also from
Nicaragua.

235a. Vittaria moritziana Mett. in Tr. & Planch. Ann. Sd. Nat. Bot. V, 2:207
(repr.289). 1864.

TYPE: Canoas [Canaos, Depto. Cundinamarca], Colombia, 2500 m, Lindig 219
(B not seen).

Plants epiphytic, at 2300 - 2400 m elevation, in forests, in the Flora area known
only from the ridge above the Rio Terbi, Valle de Silencio, Cordillera de
Talamanca, Pcia. Limon (Davidse et al 28730, UC). Also from Hispaniola and
Venezuela to Bolivia.

236. Vittaria remota F6e, M6m. Soc. Sci. Hist. Nat. Strasbourg 5 [M6m. Foug.
7];26,t.20,f. 1. 1857.

TYPE: Pcia. Ocana, Colombia, Schlim 611 (RB not seen).

Plants epiphytic, at 0-1700 m elevation, in forests, from the Cordillera Central,
Guapiles (Pcia. Limon), the upper Rio Reventazon valley, near Sta. Fe, La Mesa

FIGS. 234 - 242. Vittaria and Pteris. FIG. 234. Plant and rhizome scale detail of V. lineata, Lellinger
711. FIG. 235. Plant of V. minima, Lellinger 817. FIG. 236. Plant of V. remota, de la Sota 5236.
FIG. 237. Plant of V. stipitata, Valerio 157. FIG. 238. Basal portion of pinna of P. altissima, de la
Sota 5188. FIG. 239. Basal portion of pinna of P. biaurita, Pittier 7009. FIG. 240. Basal portion of
pinna of P. cretica, Heyde & Lux 3215, Guatemala. FIG. 241. Basal portion of pinna of P. daguensis,
Lellinger & de la Sota 660. FIG. 242. Basal portion of pinna of P. grandifolia, Biolley fil. 17389.

172 VITTARIACEAE: 37. VITTARIA

and Cerro Pilon (Pda. Code), Cerro Jefe, Cerro Pirre, Cerro Mali (Pda. Darien),
Alto del Buey, the Rio Calima, and the Rio Nuqui (Depto. Choco). Also from
Jamaica, Hispaniola, Puerto Rico, Guatemala, and Venezuela to Bolivia.

This species resembles Ananthacorus angustifolius, but has thinner laminae with
the veins slightly obvious and only one series of areolae between the midrib and
the frond margin. According to D. R. Farrar (pers. comm.), young fronds of A.
angustifolius are distinctly reddish, whereas those of V. remota are not; this
distinction may be of generic significance.

237. Vittaria stipitata Kunze, Linnaea 9:77. 1834.

TYPE: Pampayacu, Depto. Hudnuco, Peru, Poeppig Diar. 1121 (LZ destroyed;
isotype P not seen), seen by Tryon (Contr. Gray Herb. 194:215. 1964).

Plants epiphytic, at 0-1700 m elevation, in forests, from the Cordillera Central,
Finca La Selva (Pda. Heredia), the valley of the Rio Reventazon, the valley of the
Rio General, the Fila Costeha near S. Vito de Java, the valley of the Rio Dos
Bocas near Sta. Fe, around Porto Bello (Pda. Colon), Cerro Jefe, the Serrania de
Pirre (Pcia. Darien), Alto del Buey, and the lower Rio S. Juan. Also from Cuba,
Jamaica, Hispaniola, Guatemala, Nicaragua, Venezuela, Colombia to Bolivia, and
Brazil.

PTERIDACEAE: 38. PTERIS 173

PTERIDACEAE

Rhizomes stout, short-creeping, ascending, or erect, scaly, the scales linear-
lanceate to lanceolate, concolorous to bicolorous; fronds medium- to large-sized,
monomorphic to slightly or greatly dimorphic; stipes usually thick, commonly
sulcate; laminae oblong (or pedate in some species of Pteris), pinnate (to 3(4)-
pinnate-piimatifid in some species of Ptem), glabrous or thinly pilose, the margins
often thickened; veins free and commonly forked, anastomosing only in a series of
narrow, costal areolae, or completely anastomosing without included veinlets; sori
borne over the abaxial surface of the pinnae (Acrostichum), confined to a
submarginal band (Neurocallis), or along a submarginal commissural vein and
protected by the commonly scarious, renexed frond margins (Pteris); sporangia
long-stalked, the capsules globular; paraphyses present; spores trilete.

1. Fronds greatly dimorphic, the fertile longer than the sterile and with much narrower pinnae; sori
in a submarginal band.

39. Neurocallis

1. Fronds monomorphic or slightly dimorphic, the fertile fronds with somewhat narrower pinnae
or segments; sori acrostichoid or submarginal.. 2.

2(1). Sori acrostichoid; fron^ pinnate, the fertile pinnae apical or borne on separate fronds from
the fertile ones.

40. Acrostichum

2(1). Sori marginal; fronds l“3(4)-pinnate.

38. Pteris

38. PTERIS L.

Plants terrestrial, occasionally weedy; rhizomes stout, short-creeping to erect,
often woody, bearing scales at the apex, the scales lanceate or lanceolate,
concolorous, brownish, the margins entire; fronds medium- to large-sized; stipes
scaly at the base, mostly brown, often thick, commonly sulcate; laminae 1-3(4)-
pinnate-pinnatifid, often oblong, sometimes distinctly pedate or the basal pinnae
only slightly basiscopically elaborated, firmly herbaceous to coriaceous, usually
pale green, glabrous or sometimes sparsely hairy, the margins often thickened, the
axes sulcate adaxially and often remotely spiny on the ridges; veins free, partially
anastomosing (with costal areolae), or totally anastomosing without included
veinlets; sori elongate on a submarginal commissural vein, in most species not
extending to the segment apices or the sinuses between segments, protected by the
reflexed, commonly scarious frond margin; sporangia long-stalked; paraphyses
present.

Tropics and subtropics world-wide; ca. 250 species.

SCAMMAN, E. 1%1. The genus Pteris of Costa Rica. Rhodora 63:194 - 205.

1. Supra-basal pinnae lobed with 2 or more lobes, pinnatifid or more divided..3.

1. Supra-basal pinnae absent, entire, or with a single, basiscopic lobe, otherwise narrowly linear,
not pinnatifid or more divided. .2.

2(1). Veins anastomosing toward the pinna margin; fronds usually at least 1 m long; pinnae all
simple, unlobed, 15-45 cm long, 2-3 cm wide, acute or round at the base, acuminate at the apex.

242. P. ^andifolia

2(1). Veins entirely free; fronds usually less than 1 m long; at least the basal pinnae lobed or more
divided (or the laminae merely ternate); supra-basal pinnae 10-20 cm long, 1-2 cm wide, decurrent
at the base, serrate-acuminate at the apex.

240. P. cretica

3(1). Veins anastomosing, at least along the piimule costules..8.

174 PTERIDACEAE: 38. PTERIS

3(1). Veins entirely free..4.

4(3). Fronds not or scarcely pedate, the basal pinnae only pinnatifid beyond the basal pinnules..6.

4(3). Fronds decidedly pedate, the basal pinnae pinnate-pinnatifid beyond the basal pinnules..5.

5(4). Segments round or with blunt teeth, not mucronate; fronds herbaceous, dark green. Laminae
up to ca. 50 cm long, 30 cm wide; basal pinnae 2-3-pinnate-pinnatifid, strongly inequilateral; distal
pinnae pinnatifid, 2-3 cm wide.

244. P. muricella

5(4). Segments usually mucronate, the midrib prolonged into the segment apex; fronds coriaceous,
gray-green. Laminae up to ca. 100 cm long, 50 cm wide; basal pinnae 2-3-pinnate-pinnatifid,
strongly inequilateral; distal pinnae pinnatifid, 3-4 cm wide.

243. P. muricata

6(4). Veins all arising from the pinnule costules, none from the pinna costae running to the sinuses
between the pinnules. Laminae 15-100 cm long; basal pinnae bipartite; distal, pinnatifid pinnae 3-4
cm wide, often caudate; segments adnate.

250. P. plumula

6(4). Veins not all arising from the pinnule costules, some from the pinnae costae running to the
sinuses between the pinnules..?.

7(6). Lateral pinnae 1-3(5) pairs; apical pinnae about as long as the subtending lateral ones;
stipes minutely and distantly spiny; sinuses between the segments mostly symmetrical. Fronds ca.
0.5 - 1 m long.

251. P. pungens

7(6). Lateral pinnae (except in juveniles) 5-7 pairs; apical pinnae 2-3 times longer than the
subtending lateral ones; stipes smooth; sinuses between the segments mostly asymmetrical. Fronds
ca. 0.5- 1.5 m long.

247. P. paucinervata

8(3). Veins free, 1-forked, except for the narrow, uniarcuate areolae along the costae between the
costules. Laminae 0.5- 1.5 m long, 0.25-0.4 m wide, pinnate-pinnatifid, except the basal pinnae with
a prolonged, pinnatifid, basiscopic lobe.

239. P. biaurita

8(3). Veins partially or completely anastomosing, in addition to the narrow or wide, uni- or
polyarcuate areolae along the costae between the costules..9.

9(8). Costal areolae 1 between the costules (uniarcuate)..12.

9(8). Costal areolae 2-4(6) between the costules (polyarcuate), often with 1 long and 1 or 2 short
areolae.. 10.

10(9). Costae and costules spiny on the abaxial surface; laminae dark green. Laminae up to 0.5 m
long, 0.2 m wide; apical pinnae similar to the narrowly ovate-lanceolate, subtending lateral pinnae;
veins anastomosing in 1 or 2 series.

246. P. obscura

10(9). Costae and costules not spiny on the abaxial surface; laminae gray-green..ll.

11(10). Lateral pinnae (except the basal and occasionally the suprabasal ones) 5-10(15) cm wide,
up to 30 cm long, linear-lanceolate, with up to ca. 12(18) pairs of distant, falcate segments.

238. P. altissima

11(10). Lateral pinnae (except the basal ones) 2.5 -3.5 cm wide, up to 25 cm long, linear-
lanceolate, with up to ca. 18 pairs of crowded, falcate segments.

241. P. daguensis

12(9). Ultimate segments (2)5-10 cm long (less in apical portions); rachises and costae sometimes
sparingly spiny. Laminae pedate, up to 3 m long, 1.5 m wide, sparsely pilosulous abaxially, especially
on the veins.

245. P. navarrensis

12(9). Ultimate segments less than 3 cm long; rachises and costae not spiny..l3.

PTERIDACEAE: 38. PTERIS 175

13(12). Fronds 2(3)-pmnate-pinnatifid, slightly pedate with the basal basiscopic pinnule usually
only pinnatifid; fronds up to 1(1.25) m long. Fertile segments mostly with a single row of areolae
along the costule.

249. P. polita

13(12). Fronds at least 4-pinnate-pinnatifid at the base, decidedly pedate with the basal basiscopic
pinnule 2-pinnate-pinnatifid; fronds up to 2 m long.. 14.

14(13). Veins (and sometimes the laminae) sparsely pilose on the abaxial surface; sterile segments
toothed; ultimate segments usually mucronate, falcate; indusia usually continuous around the round
sinuses.

248. P. podophylla

14(13). Veins and laminae glabrous on the abaxial surface; sterile segments entire; ultimate
segments usually round, not falcate; indusia usually discontinuous, only approaching the acute to
obtuse sinuses.

252. P. tripartita

238. Pteris altissima Poir. Encyc. M^th. 5:722. 1804.

Pteris macroptera Link, Hort. Reg. Bot. Berol. 2:32. 1833. TYPE: Based on material received
from Brazil and cultivated in the botanical garden in Berlin (B not seen). See Morton (Contr. U. S.
Natl. Herb. 38:72. 1967).

Pteris kunzeana J. Agardh, Recens. Sp. Pterid. 62. 1839. TYPE: Based on P. podophylla sensu
Kunze, excluding all synonymy, and, therefore, on Pampayacu, Depto. Hu^uco, Peru, Poeppig Diar.
1137 (LZ destroyed); and Mission Tocache, Rio Huallaga, Depto. S. Martin, Peru, July 1830, Poeppig
(LZ destroyed).

Pteris elata J. Agardh, Recens. Sp. Pterid. 63. 1839. TYPE: Panama, Cuming 1267 (K not seen
fragm NY; isotype GH).

Pteris arborescens Martens & Galeotti, Nouv. M6m. Acad. Roy. Sci. Bruxelles 15:54. 1842.

TYPE: Chinantla, Edo. Oaxaca, Mexico, Galeotti 6375 (BR not seen). Several isotypes labelled P.
arborescens and bearing number 6575 corrected to 6376 are truly the latter number and are isotypes
of Litobrochia mexicana Fee, which is a synonym of P. pulchra Schlechtend. & Cham.

Pteris protea Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd., V, 1:228 (repr. 76).
1849. TYPE: Between Colipa and Misantla, Edo. Veracruz, Mexico, Liebmann 757 (C not seen
fragm US; isotypes GH not seen Tryon photo, K not seen).

Litobrochia grandis Fee, M6m. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:75. 1857.

SYNTYPES: Barranca de S. Francisco, near Mirador, Edo. Veracruz, Mexico, Schaffher 144 (RB
not seen); Cuba, Morelet (P or RB not seen); and Hispaniola, de Tussac (P or RB not seen).

Pteris schiedeana Ettingsh. Farnkr. Jetztw. 99, f. 52-54, t. 65, f. 9., t. 66, f. 1. 1864. TYPE: none
cited.

TYPE: Puerto Rico, LeDru in 1797 (P-Hb. Lam. not seen Tryon photo; isotype
P-Hb. Juss. 1315 not seen photo 3083).

Plants terrestrial, at 0“2000 m elevation, at forest margins, along trails, and
sometimes in open areas, from throughout the Flora area. Also from the Antilles,
Central America, Venezuela to Bolivia, Surinam, and Brazil.

See P. podophylla for a comment on a presumed hybrid with that species.

239. Pteris biaurita L. Sp. PL 2:1076. 1753.

Litobrochia galeottii Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:75. 1857. TYPE:
Oaxaca and Teotalingo, Edo. Oaxaca, Mexico, Galeotti 6485 (RB not seen).

Pteris biaurita var. subpinnatifida Jenm. Bull. Bot. Dept. Jamaica 41:7. 1893; Ferns Brit. W. Ind.
Guiana 123. 1900. TYPE: Jamaica, /e/iman (NY? not seen).

LECTOTYPE: Plate 14 of Plumier’s “Description...”, chosen by Proctor (FI.
Less. Antill. 2:145. 1977).

176 PTERIDACEAE: 38. PTERIS

Plants terrestrial, at 0-1100 m elevation, in open areas in forests, from the
Atlantic and Pacific coastal plains of Costa Rica, the valley of the Rio General,
between S. Felix and the Cerro Colorado copper mine (Pcia. Chiriqm), El Valle,
and near Pacora (Pcia. Panama). Also from the Antilles, Mexico to Nicaragua,
Venezuela to Peru, Guyana, and Brazil.

240. Pteris cretica L. Mant. PL 130. 1767.

Pteris triphylla Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:51, t. 14, f. 1. 1842,
non J. Agardh, 1839. TYPE: Jalapa, Edo. Veracruz, Mexico, Galeotti 6393 (BR not seen photos
5119, 5120).

Pteris trifoliata F6e, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 81:114. 1857. TYPE: A
renaming of P. triphylla Martens & Galeotti, and so based on the type of that name.

TYPE: Crete, collector unknown (LINN 1246.7 not seen microfiche S. I.
Library), according to Tryon (Contr. Gray Herb. 194:192. 1964).

Plants terrestrial, at 1600-1900 m elevation, in open areas, often among rocks,
from La Banderilla and Reventado (Pcia. Cartago), and from Chilibre (Pcia.
Panama). Also from the southeastern United States, the Bahamas, Jamaica,
Mexico, Guatemala, Peru, Brazil, and Argentina.

This Old World species is widely escaped in the New World.

241. Pteris daguensis (Hieron.) Lellinger, Amer. Fern J. 67:59. 1977.

Pteris orizabae var. daguensis Hieron. Bot. Jahrb. Syst. 34:495. 1904. TYPE: Banks of the Rio
Dagua, Depto. El Valle, Colombia, 200 - 500 m, Lehmann 8933 (B not seen; isotype US).

Plants terrestrial, at 0-100 m elevation, in disturbed forests and waste places, in
the Flora area known only from near Loma del Cuchillo, Depto. Choco {Lellinger
& de la Sota 660, COL, CR, HUA, LP, US). Also from adjacent areas in
Colombia.

242. Pteris grandifolia L. Sp. PI. 2:1073. 1753.

Pteris socorrensis Karst. FI. Columb. 1(3): 118, t. 58. 1860. TYPE: Near Socorro and Chaparral,
Depto. El Valle, Colombia, 800-1100 m, Karsten (LE not seen Tryon photo).

Pteris grandifolia var. campanae Rosenst. Repert. Spec. Nov. Regni Veg. 7:291. 1909. TYPE: Mt.
Campana near Tarapoto, Depto. S. Martm, Peru, Spruce 4668 (S not seen; isotype K not seen).

LECTOTYPE: Plate 105 of Plumier’s ‘Traite...’’, chosen by Proctor (FI. Less.
Antill. 2:143. 1977). Although Linnaeus cited t. 106 in error for t. 105, he also
cited t. 8 of Plumier’s “Description...”, which is identical with t. 105. Both plates
are based on a specimen collected by Plumier on Hispaniola.

Plants terrestrial, at 0-600(1500) m elevation, in wet forests, from La Palma
and the Atlantic and Pacific lowlands of Costa Rica and Panama. Also from
Florida, the Antilles, Trinidad, Mexico to Nicaragua, Venezuela, and Colombia to
Peru.

243. Pteris muricata Hook. Sp. Fil. 193, t. 123B. 1858.

TYPE: Depto. Antioquia or Cundinamarca, Colombia, Jervise (K not seen).
Plants terrestrial, at 1300-3000 m elevation, along streams and in forests, often
in open areas, from the Cordillera Central, Cerro Carpintera, and the Cordillera
de Talamanca. Also from Mexico, Guatemala, Nicaragua, and Colombia to
Bolivia.

PTERIDACEAE: 38. PTERIS 177

244. Pteris muricella F^e, M4m. Soc. Sci. Hist. Nat. Strasbourg 5 [M^m. Foug.
8]:73. 1857.

Pteris mollis Bommer & Christ, Bull. Herb. Boissier 4:658. 1896. TYPE: S. M2ircos, Pcia. S. Jose,
1355 m, Tonduz 7565 (BR not seen; isotypes P not seen Tryon photo, US).

SYNTYPES: Near Cordoba and Huatusco, Edo. Veracruz, Mexico, Schajfner
143 (RB not seen) and 1854 (P or RB not seen).

Plants terrestrial, at 1000-2600 m elevation, in forests and in open areas, from
the Cordillera Central and the Cordillera de Talamanca to Pcia. Chiriqui. Also
from Mexico, Guatemala, El Salvador, Nicaragua, and Venezuela.

245. Pteris navarrensis Christ, Bull. Soc. Bot. Geneve II, 1:227. 1909.

TYPE: Valley of the Rio Navarro, Pcia. Cartago, 1400 m, Werckle 16761 (P not
seen; isotype US).

Plants terrestrial, at 1000-2700 m elevation, in forests and at forest margins,
from the Cordillera Central and the Cordillera de Talamanca to Pcia. Chiriqui.

This variable species may have aculeate axes at high elevations and may be
identical to P. livida Mett. var. livida, but probably not to \2LT.ferox Mett.

246. Pteris obscura Mett. ex Kuhn, Linnaea 36:90. 1869.

TYPE: Venezuela, Punch & Schlim 599 (B not seen).

Plants terrestrial, at 1300-1400 m elevation, in forests, in the Flora area known
only from Cerro Pirre {Goldman 1890, US). Also from Colombia.

247. Pteris paucinervata F6e, M6m. Soc. Sci. Hist. Nat. Strasbourg 5 [M^m. Foug.
8]:73. 1857.

TYPE: Barranca de S. Martin, near Mirador, Edo. Veracruz, Mexico, Schajfner
152 (RB not seen).

Plants terrestrial, at 1000-1800 m elevation, in forests, from Vara Blanca (Pcia.
Heredia), Cerro Tablazo, and around Boquete. Also from Mexico, Guatemala,
and Nicaragua.

248. Pteris podophylla Swartz, J. Bot. (Schrader) 1800(2) :67. 1801.

Lonchitis pedata L. Sp. PL ed. 2, 2:1536. 1763, non Pteris pedata L., 1753. TYPE: Jamaica,
Browne (LINN 1249.1; isotype S not seen).

Litobrochia camptocarpa Fee, Gen. Fil. [M6m. Foug. 5]:137. 1852. TYPE: Colonia Tovar, Edo.
Aragua, Venezuela, Moritz 47 (P or RB not seen).

Litobrochia setifera F6e, Gen. Fil. [M6m. Foug. 5]: 138. 1852. TYPE: Mt. S. Martin, Mexico,
Galeotti 6571 (RB not seen).

Pteris trialata Sodiro, Anales Univ. Quito 8(55):76 (repr. 107). 1893. TYPE: Niebli, Pcia.

Pichincha, Ecuador, Sodiro (Hb. Sodiro not seen; isotypes P not seen, US).

Pteris fermginea Bommer in Dur. & Pitt. Bull. Soc. Roy. Bot. Belgique 35, Mem. 191. 1896, nom.
illeg. PubUshed in synonymy.

Pteris pedata Kuhn in Schenck, Hedwigia 35:161. 1896, non L., 1753, nom. illeg. A renaming of P.
podophylla Swartz, and so based on the type of that name.

Pteris podophylla var. rufopubescens Hieron. Hedwigia 48:244. 1909. TYPE: Between Bogota and
Muzo, Depto. Cundinamarca or Boyadi, Colombia, Stuebel 538 (B not seen).

Pteris inflexa Copel. Univ. Calif. Publ. Bot. 19:289, t. 41. 1941. TYPE: Barranca near Ojo de
Agua, Orizaba, Edo. Veracruz, Mexico, 1200 m, Copel. Mex. Ferns 79 (MICH not seen; isotype UC).

TYPE: A renaming of Lonchitis pedata L., and so based on the type of that
name.

178 PTERIDACEAE: 38. PTERIS

244 P. muricella

252 P. tripartita

PTERIDACEAE: 38. PTERIS 179

Plants terrestrial, at (400)1000 - 2500 m elevation, at forest edges and along
trails in forests, from north of S. Ram6n, the Cordillera Central, the Cordillera de
Talamanca to Pda. Chiriqui, near S. Vito de Java, and Curandd below Novita
(Depto. Choc6). Also from Cuba, Jamaica, Hispaniola, Mexico, Guatemala, El
Salvador, Nicaragua, and Venezuela to Bolivia.

Pteris costaricensis Rosenst. probably is a hybrid between this species and P.
altissima. A presumable isotype {Erode & Erode 469, UC) has irregular spores,
some of them abortive, and probably 16 or 32 per sporangium. The lamina is
yellow-green as in P. podophyllo, but the costal veins are biarcuate like those of P.
oltissimo. Rosenstock (Repert. Sp. Nov. Fedde 22:7. 1925) cited 461 as the type
number; either it or 469 could be in error.

249. Pteris polita Link, Hort. Reg. Bot. Berol. 2:30. 1833.

Pteris propinqua J. Agardh, Recens. Sp. Pterid. 65. 1839. LECTOTYPE: Jamaica, Bancroft (K not
seen), chosen by Tryon (Contr. Gray Herb. 194:202. 1964).

Pteris propinqua var. cumingiana J. Agardh, Recens. Sp. Pterid. 65. 1839. TYPE: Panama, Cuming
1182bis (K not seen fragm US).

Pteris biformis Splitgb. Tijdschr. Natuurl. Gesch. Physiol. 7:422. 1840. TYPE: Merveille

Plantation, Surinam, Splitgerber (L not seen photos 187- 189).

Pteris hostmanniana Ettingsh. Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss. Kl. 23:55. 1864.
TYPE LOCALITY: Surinam.

TYPE: Based on material from Brazil cultivated in the botanical garden at
Berlin (authentic specimens B not seen photos 5449, 5450), according to Morton
(Contr. U. S. Natl. Herb. 38:75. 1967).

Plants terrestrial, at 0-700 m elevation, in forests, from the Atlantic coastal
plain of Costa Rica and western Panama, near Sta. Fe, the Canal Zone and
adjacent Pda. Panama, near Cana, and near La Teresita (Depto. Choco). Also
from Jamaica, Mexico to Nicaragua, Venezuela to Bolivia, Surinam, and Brazil.

250. Pteris plumula Desv. M^m. Soc. Linn. Paris 6:297. 1827.

Pteris pectinata Desv. Ges. Naturf, Freunde Berlin Mag. 5:324. 1811, non Cav., 1802, nom. Uleg.
TYPE: Antilles, collector unknown (P not seen), discussed by Weatherby (Contr. Gray Herb. 124:18.
1939).

Pteris swartziana J. Agardh, Recens. Sp. Pterid. 34. 1839, nom. illeg. TYPE: A renaming of P.
biaurita sensu Swartz (Syn. Fil. 98. 1806), and so based on the basis of that name, which itself is
superfluous because P. quadriaurita Retz. was cited in synonymy by Swartz.

Pteris repandula Link, Fil. Sp. 56. 1841. TYPE: A renaming of P. biaurita sensu Link (Hort. Reg.
Bot. Berol. 2:28. 1833), and so based on the basis of that name.

Pteris nemoralis var. major Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:53.
1842. TYPE: Zacuapan, Edo. Veracruz, Mexico, Galeotti 6291 (BR not seen).

Pteris felosma J. Smith, Bot. Mag. (Curtis) 72, Comp.:24. 1846. TYPE: Cultivated at Kew,
originally from Jamaica, collector unknown (K not seen; isotypes BM not seen, L not seen photo
2177).

FIGS. 243 - 252. Pteris. FIG. 243. Pinnule base of P. muricata, Maxon 314. FIG. 244. Pinnule base of
P. muricella. Smith 48/141. FIG. 245. Pinna base of P. navarrensis, Lankester 711. FIG. 246. Pinna
base of P. obscura, Goldman 1890. FIG. 247. Pinna base of P. paucinervata, Killip 5204. FIG. 248.
Pinnule base of P. podophyllo, Maxon 8083. FIG. 249. Basal portion of basal basiscopic pinnule of
basal pinna of P. polita. Stern 684. FIG. 250. Pinna base of P. plumula, Wilbur & Stone 9788. FIG.
251. Pinna base of P. pungens, Scamman & Holdridge 7916. FIG. 252. Pinna base of P. tripartita,
Feddema 2076, Colombia.

180 PTERIDACEAE: 39. NEUROCALLIS

Pteris quadriaurita var. affluentior Jenm. J. Bot. Brit. For. 19:53. 1881, as "'ajfluentiusP TYPE:
Jamaica, Jenman 12 in 1879 (K not seen).

Pteris quadriaurita var. curtidens Christ in Dur. & Pitt. Bull. Soc. Roy. Bot. Belgique 35, Mem. 189.
1896. TYPE: S. Francisco de Guadalupe, Pcia. S. Jos6, Tonduz 8494 (BR not seen; isotype CR).

Pteris quadriaurita var. asperula Christ in Pitt. Prim. FI. Costaric. 3(1) :22. 1901. TYPE: Canas
Gordas, Pcia. Puntarenas, 1100 m, Pittier 10963 (P or RB not seen; isotypes CR, US).

Pteris quadriaurita var. strigulosa Christ, BuU. Herb. Boissier II, 5:160. 1905, nom. illeg. Based on
P. quadriaurita var. asperula Christ, non P. asperula J. Smith, 1841.

TYPE: A renaming of P. pectinata Desv., and so based on the type of that name.
Plants terrestrial, at 0-1900 m elevation, in forests, along trails and streams,
from the Cordillera de Tilardn, the Cordillera Central, the Meseta Central, Cerro
Carpintera, the Cordillera de Talamanca to Pcia. Chiriqui, and Cerro Azul. Also
from throughout most of tropical America, except for the Guianas.

The well known name P. quadriaurita Retz., which applies to sexual diploid
material from Ceylon and southern India, probably should not be used for
American material or for Old World material that is apogamous and triploid,
judging by counts reported by Love, Love, and Pichi Sermolli (Cytotax. Atlas
Pterid. 124. 1977).

251. Pteris pungens Willd. Sp. PI. ed. 4, 5:387. 1810.

Pteris macroura Willd. Sp. PI. ed. 4, 5:380. 1810. TYPE: Plate 13 of Plumier’s “Trait6...”, which
was based on a specimen collected by Plumier on Martinique or Hispaniola.

Pteris acuminata Desv. Ges. Naturf. Freunde Berlin Mag. 5:324. 1811. TYPE: Puerto Rico,
collector unknown (P-Hb. Desv. not seen), examined by Proctor (pers. comm.).

Pteris longicauda Christ in Pitt. Prim. FI. Costaric. 3(1):21. 1901. TYPE: Sto. Domingo de Golfo
Dulce, Pcia. Puntarenas, Tonduz 10011 [cited as 10071] (P not seen; isotype US).

Pteris pungens var. shimekii Rosenst. M6m. Soc. Sci. Nat. Neuchatel 5:37, f. 3. 1912. TYPE: Trail
from Fresno, near Mariquita, Depto. Tolima, Colombia, 800 m. Mayor 101 (S not seen; isotype US).

LECTOTYPE: Plate 14 of Plumier’s “Traite...”, which was based on a specimen
collected by Plumier along the Grande Riviere in the Leogane section of
Hispaniola, chosen by Proctor (FI. Less. Antill. 2:144. 1977).

Plants terrestrial, at 0-800(1100) m elevation, from throughout the Flora area.
Also from Jamaica, Hispaniola, Puerto Rico, the Lesser Antilles, Trinidad, Mexico
to Nicaragua, Venezuela to Bolivia, and Surinam.

252. Pteris tripartita Swartz, J. Bot. (Schrader) 1800(2) ;67. 1801.

TYPE: Java, Thunherg (UPS not seen), cited by Swartz (Syn. Fil. 100, 293.
1806).

Plants terrestrial, at 0-700(1100) m elevation, from Tapantf, near Limon (Pcia.
Limon), the Peninsula de Osa, Cerro Jefe and Cerro Brewster (Pcia. Panama),
and the Rio Tamana below Novita (Depto. Choc6). Also from Florida, Cuba,
Jamaica, Puerto Rico, the Lesser Antilles, Nicaragua, Venezuela, and Colombia.

This species is naturalized from the Old World and is spreading rapidly in the
New World tropics.

39. NEUROCALLIS

Plants terrestrial; rhizomes stout, erect or short-ascending, woody, scaly at the
apex, the scales small, linear-lanceate, sharply bicolorous, the margins slightly
erose; fronds large, strongly dimorphic, the fertile ones exceeding the sterile ones;

PTERIDACEAE: 40. ACROSTICHUM 181

stipes thick, sulcate, scaly and atropurpurous at the base, stramineous distally;
laminae oblong, pinnate, firmly herbaceous, glabrous, the veins anastomosing
without included veinlets in several series on each side of the costae, the size of
the areolae diminishing toward the frond margin; sterile pinnae elliptic-oblong,
acute at the base, acuminate at the apex; fertile pinnae linear, bearing sporangia
in a wide, submarginal band scarcely protected by the sterile, revolute pinna
margin; sporangia long-stalked, the capsules deciduous, the stalks appearing
among the sporangia like paraphyses.

Antilles, Costa Rica, and Venezuela; monotypic.

253. Neurocallis praestantissima van subcaudata Gdmez, Revista Biol. Trop.

20:183. 1972.

Pteris macrodictya Christ, Bull. Herb. Boissier II, 7:267. 1907. TYPE: Costa Rica, Werckle in 1904
(P not seen).

TYPE: Patillo, Pcia. Cartago, Gomez & Kennedy 3345 (CR not seen).

Plants terrestrial, at 900-2400 m elevation, in wet forests, from Patillo, Tapanti,
near Pejivalle (Pcia. Cartago), and the Cerro de las Caricias and near Vara Blanca
(both Pcia. Heredia).

The typical variety from the Antilles and Venezuela differs in having the sterile
pinnae sessile or nearly so.

40. ACROSTICHUM L.

Plants terrestrial in marshes and usually mangrove swamps; rhizomes stout,
short-creeping or erect, bearing copious, spongy roots, scaly at the apex, the scales
lanceolate, slightly bicolorous, the margins entire to erose; fronds large,
caespitose, slightly dimorphic, the fertile pinnae distal or on separate fronds from
the sterile pinnae; stipes thick, sulcate, stramineous to brownish, scaly at the very
base; laminae oblong, pinnate, chartaceous to coriaceous, glabrous or thinly pilose
on the abaxial surface; veins anastomosing in many series of about uniform size,
without included veinlets; sporangia long-stalked, borne over the abaxial surface
of the pinnae, not organized into sori; paraphyses present.

Pantropical, mostly in swamps; 3 species.

ADAMS, D. C. and P. B. TOMLINSON. 1979. Acrostichum in Florida. Amer. Fern J. 69:42 - 46.

1. Veins of sterile pinnae immersed abaxially, glabrous; fertile fronds spreading like the sterile
ones, with only 1-4 apical pinna pairs fertile; pinnae 10-14 pairs, distant, coriaceous; paraphyses
among the sporangia minute, capitate-stellate. Fronds 1.5-3 m long; laminae oblong, 1-2 m long,
20 - 40 cm wide; pinnae elliptic-spathulate, 15-30 cm long, 3-6 cm wide.

254. ^1. aureum

1. Veins of sterile pinnae prominulous abaxially, usually thinly pilose; fertile fronds strictly erect
and taller than the spreading sterile ones, with all the pinnae fertile; pinnae ca. 15 or more pairs,
imbricate or nearly so, chartaceous; paraphyses among the sporangia with a large, sausage-shaped
terminal cell. Fronds 1.5 -3.5 m long; laminae oblong, 1-2.5 m long, 25 - 60 cm wide; pinnae
narrowly lanceolate to spathulate, 15-40 cm long, 3-5 cm wide.

255. A. danaeifolium

254. Acrostichum aureum L. Sp. PI. 2:1069. 1753.

Acrostichum marginatum Schkuhr, Vier Zwanzigste Kl. Linn. PfI.-Syst. 2:185, t. 3b. 1805, non L.,
1753, nom. illeg. TYPE: Essequibo River, Guyana, Meyer (HAL not seen).

Acrostichum formosum K. Presl, Delic. Prag. 1:160. 1822. TYTE: Rio de Janeiro, Est. Rio de
Janeiro, Brazil, Langsdorjf (LE or PRC not seen).

182 PTERIDACEAE: 40. ACROSTICHUM

PTERIDACEAE: 40. ACROSTICHUM 183

Acrostichum juglandifolium Kaulf. Enum. Fil. 65. 1824. TYPE: A renaming of A. marginatum
Schkuhr, and so based on the type of that name.

Chrysodium vulgare F6e, Hist. Acrostich. [M6m. Foug. 2]:97. 1845, nom. superfl. TYPE: A
renaming of^. aureum L., and so based on the type of that name.

Chrysodium cayennense F6e, Hist. Acrostich. [M6m. Foug. 2]: 100, t. 59. 1845. TYPE: French
Guiana, LePrieur 1837 (P-Hb. Bory not seen).

Chrysodium scalpturatum F6e, Hist. Acrostich. [M6m. Foug. 2]: 100, t. 61. 1845. SYNTYPES:
Near Lazareto, Panama, Bonpland (RB not seen); and New Ireland, Richard (P-Hb. Bory not seen).

LECTOTYPE: Plate 7 of Plumier’s “Description...”, which is based on a
specimen collected by Plumier at Ft. Royal, Martinique, chosen by Lellinger
(Proc. Biol. Soc. Wash. 98:366. 1985).

Plants terrestrial, at 0-100(300) m elevation, in swamps, from the Atlantic and
Pacific coasts and coastal plains of Costa Rica and Panama, the Canal Zone, and
the Rio Baud6. Also from Florida, the Antilles, Mexico to Nicaragua, Trinidad,
Venezuela, Colombia, Ecuador, the Guianas, and Brazil.

Seey4. danaeifolium for a discussion of hybrids with that species.

255. Acrostichum danaeifolium Langsd. & Fisch. PL Voy. Russes Monde 1:5, t. 1.

1810.

Chrysodium hirsutum Fee, Hist. Acrostich. [Mem. Foug. 2]:99, t. 62, f. 2. 1845. SYNTYPES:
Brazil, Pohl (MA? not seen), Martius 365 (MA not seen); Guyana, LePrieur (P or RB not seen);
Guadeloupe, L*Herminier (P or RB not seen); Hispaniola, Ritter (P or RB not seen); Guatemala,
Friedrichsthal 231 (W not seen); and South Africa, Schott (W not seen).

Chrysodium lomarioides Jenm. Timheri 4:314. 1885, non Acrostichum lomarioides Bory, 1833.
TYPE: Guyana, /e/imon (NY? not seen).

Acrostichum excelsum Maxon, Proc. Biol. Soc. Wash. 18:224. 1905. TYPE: A renaming of

Chrysodium lomarioides Jenm., and so based on the type of that name.

Chrysodium lomarioides var. hastatum Christ, Bull. Herb. Boissier II, 5:725. 1905. TYPE: Near El
Carmen, Edo. Chiapas, Mexico, Muench (P not seen; isotypes DS not seen, US).

Acrostichum excelsum f. lobatum A. A. Eaton, Bull. Torrey Bot. Club 33:464. 1906. TYPE:
Florida, yl. A. Eaton (GH? not seen).

Acrostichum danaeifolium f. incanum Domin, Rozpr. Kr^. Cesk6 Spolecn. Nauk, Tf. Mat.-Pffr. 2
[Pterid. Dominica]: 111. 1929. TYPE LOCALITY: Florida.

TYPE: Ilha Sta. Catarina, Est. Sta. Catarina, Brazil, Langsdorjf (LE not seen
Nicolson photo).

Plants terrestrial, at 0-200 and 1100-1200 m elevation, in coastal swamps and
rarely upland marshes, from the Atlantic coast of Costa Rica and Panama, near
Desamparados (Pcia. S. Jose), and Barro Colorado Island and the vicinity of Juan
Mina, Chagres River (Canal Zone). Also from tropical America except Peru.

R. L. Petersen (pers. comm.) has found a hybrid between this species and A.
aureum in lowland Panama. ITie fertile fronds and paraphyses are intermediate
between the parents, and the spores are abortive.

FIGS. 253 - 259. Neurocallis, Acrostichum, Loxsomopsis, and Hymenophyllum. FIG. 253. Sterile and
fertile pinnae of N. praestantissima var. subcaudata, Skutch 3753 and Standley & Valerio 46929. FIG.
254. Abajdal surface of the sterile lamina of A. aureum, Mickel 2761. FIG. 255. Abaxial surface of
the sterile lamina of^. danaeifolium, Bartlett & Lasser 16318. FIG. 256. Basal portion of suprabasal
pinnule of L. costaricensis, Mickel 3001. FIG. 257. Median pinna of H. tunbiigense. Stork 1290. FIG.
258. Median pinna of H. fucoides, Maxon 5447. FIG. 259. Median pinna of H. consanguineum,
Standley & Valerio 50300.

184 LOXSOMACEAE: 41. LOXSOMOPSIS

LOXSOMACEAE

Monotypic; see description of Loxsomopsis.

41. LOXSOMOPSIS Christ

Plants terrestrial; rhizomes long-creeping, branched, bearing a few fibrous roots
and numerous dark brown bristles; fronds mostly large, scattered along the
rhizomes; stipes brown, sparsely bristly at and near the base, slightly sulcate;
laminae lanceolate to oblong, 2-pinnate-pinnatisect, herbaceous, provided with
catenate hairs on the axes and especially the abaxial lamina surface; veins free,
forked, reaching the margin, the sterile vein tips slightly swollen; pinnae
asymmetrical, gradually reduced toward the basiscopic base, the basiscopic basal
pinnules reduced nearly to auricles; sporangia borne at the apex of an elongate,
marginal receptacle, protected by an urn-shaped, entire indusium; paraphyses
present.

Tropical America, but very rare; 3 species.

256. Loxsomopsis costaricensis Christ, Bull. Herb. Boissier II, 4:399. 1904.

TYPE: Costa Rica, Werckle & Brune 279 (P not seen).

Plants terrestrial, at 1900-2600 m elevation, in wet, open forests and scrub
forests, from the Cordillera Central, near Tapanti, and the Cordillera de
Talamanca along the Interamerican Highway south of El Empalme (Pcia. S. Jose).

According to F. S. Wagner (Amer. J. Bot. 67:737. 1980), this species has 46
pairs of chromosomes, which is a dennstaedtioid number.

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 185

HYMENOPHYLLACEAE

Rhizomes very thin and long-creeping or short and erect to ascending, hairy, the
hairs multicellular, often bristle-like; fronds minute to medium-sized, rarely
hemiepiphytic and indefinitely long; stipes wiry or thin, often alate at least distally,
obsolete in a few species; laminae simple to several times pinnate, usually 1 cell
thick between the veins, rarely nearly skeletonized, simple to several times
pinnate, glabrous or minutely hairy, the hairs simple, forked, stellate, or more
complex; veins free or anastomosing without included veinlets, sometimes with
false veins parallel or perpendicular to the true veins; sori marginal, terminal on
ultimate segments or along the entire to lobed lamina margin; indusia (involucres)
bilabiate or tubular; sporangia relatively few per sorus, subspherical, the annulus
transverse to slightly oblique; paraphyses none.

1. Involucres bilabiate, circular, semicircular, or angled or conical at the base; sporangia-bearing
receptacles low or short, rarely protruding from the involucre.

42. Hymenophyllum

1. Involucres long-conical to tubular, truncate at the apex or with short involucral lips; sporangia-
bearing receptacles long, setiform, commonly protruding from the involucre in age.

43. Trichomanes

42. HYMENOPHYLLUM J. E. Smith

Plants epiphytic or occasionally epipetric; rhizomes long-creeping, almost
thread-like, hairy, the hairs multicellular, brown or reddish; fronds small to
minute; stipes filiform, sometimes partially alate, scattered along the rhizomes,
glabrous or hairy, the hairs simple or variously stellate; laminae simple to
decompound, ovate to oblong, for the most part only 1 cell thick, the rachis
commonly partially or entirely alate, glabrous or the lamina surface and/or
margins hairy, the hairs simple, forked, or variously stellate, the sterile ultimate
segments with a single vein not reaching the margin; sporangia borne on usually
low marginal receptacles, protected by an involucre formed of two usually nearly
round or ovate flaps of tissue; sporangia long-stalked.

Mostly pantropical in perpetually moist localities at middle elevations; ca. 310
species.

LELLINGER, D. B. 1984. Hymenophyllaceae (Filicales) in B. Maguire and collaborators. The
botany of the Guayana Highland -Part XII. Mem. New York Bot. Card. 38:9 - 46.
MORTON, C. V. 1947. The American species of Hymenophyllum section Sphaerocionium. Contr.
U. S. Natl. Herb. 29:139-199.

MORTON, C. V. 1968. The genera, subgenera, and sections of the Hymenophyllaceae. Contr. U. S.
Natl. Herb. 38:153 - 214.

1. Segment margins entire or, if rarely slightly denticulate, the teeth tipped by 1-ceUed or stellate
hairs; involucres usually round or obtuse, rarely conical at the base..3.

1. Segment margins toothed, the teeth not tipped by hairs; involucres often rather conical at the

base (subg. Hymenophyllum)..!.

2(1). Involucres ovate to obovate, not more than 2 times longer than wide; only 1(2) adaxial
acroscopic pinna segments fertile; rachises glabrous; laminae (1)1.5 -2 cm wide.

258. H. tunhrigense

2(1). Involucres ovate to linear, 2-5 times longer than wide; (1)2-5 adaxial acroscopic pinna
segments fertile; rachises usually sparsely pilose; laminae 2-4(6) cm wide.

257. H. fucoides

3(1). Laminae glabrous (subg. Mecodium).. 23.

186 HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM

3(1). Laminae hairy on the lamina tissue and/or the veins on the abaxial surface (subg.
Leptocionium).A.

4(3). Hairs absent on the lamina surfaces, borne only on the veins and margins of the laminae. .11.

4(3). Hairs present on the lamina surfaces, as well as on the veins and margins..5.

5(4). Stipes 0.15-0.25 mm in diam.; fronds mostly less than 15 cm long, determinate or
subdeterminate..8.

5(4). Stipes (0.25)0.4-0.6 mm in diam.; fronds mostly more than 15 cm long, indeterminate.. 6.

6(5). Rachises not alate; laminae densely covered with stellate hairs, the surface obscure. Laminae
grayish adaxially, cinnamomeous abaxially; pinnae up to 4 cm long, 1 cm wide, the fronds tapered
toward the base.

211. H. plumosum

6(5). Rachises alate; laminae sparsely covered with stellate hairs, the surface visible..?.

7(6). Proximal pinnae equilateral at the base, acuminate at the apex; brownish, stellate, commonly
4- armed hairs few between the veins; veins of the laminae not alate.

275. H. sieberi

7(6). Proximal pinnae inequilateral at the base, excavate at the basiscopic base, obtuse at the apex;
cinnamomeous, stellate, mostly 5 -7-armed hairs many between the veins; veins of the laminae
sometimes shghtly alate on the abaxial surface.

266. H. horizontale

8(5). Pinnae all simple, undivided. Stipes 1-3 cm long; laminae ca. 2 times longer than the stipes;
rachises alate throughout; ultimate segments 1-1.75 mm wide.

267. H. lanatum

8(5). Pinnae, or most of them, pinnatifid or more divided..9.

9(8). Rachises alate throughout; laminae 4-8 times longer than the stipes, narrowly lanceate,
usually widest at the base; stipes 0.5 - 1.5 cm long. Ultimate segments 1.25 - 1.75 mm wide.

263. H. fragile

9(8). Rachises partially exalate; laminae 1-2(4) times longer than the stipes, ovate to ovate-
lanceolate, usually widest distal to the base; stipes 1-4 cm long..l0.

10(9). Stipe hairs stellate; basal basiscopic pinnules usually simple; ultimate segments 0.8-1 mm
wide; lowest pinnae petiolulate.

264. H. hemipteron

10(9). Stipe hairs simple (1-3-celled) or a few forked with 1- or 2-celled, elongate rays; basal
basiscopic pinnules l(2)-forked; ultimate segments 1-1.5(2.25) mm wide; basal pinnae sessile or
rarely petiolulate.

262. H. elegantulum

11(4). Stipes (exclusive of any ala) usually less than 0.3 mm in diam., usually less than 5 cm
long.. 14.

11(4). Stipes (exclusive of any ala) 0.3 mm in diam. or more, usually more than 5 cm long; hairs on
the lamina margins simple or forked at the base.. 12.

12(11). Rachises not entirely alate. Stipes (3)5-10 cm long; laminae ovate-lanceolate, 6-18 cm
long, 2.5-9 cm wide; lumina of marginal cells rather large and clear.

276. H. subrig^dum

12(11). Rachises alate throughout..l3.

13(12). Stipes alate at the apex; rhizomes 0.3 -0.5 mm in diam. Stipes 3-9 cm long; laminae ovate
to ovate-lanceolate, rarely linear, 7.5-20 cm long, (2)4- 10 cm wide.

269. H. microcarpum

13(12). Stipes not alate at the apex; rhizomes 0.7-1 mm in diam. Stipes 5-10 cm long; laminae
ovate to ovate-lanceolate, 8- 15 cm long, 6- 10 cm wide.

259. H. consanguineum

14(11). Rachises partially exalate..l9.

14(11). Rachises entirely alate..l5.

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 187

15(14). Marginal hairs mostly simple or forked at the base..l7.

15(14). Marginal hairs mostly stellate..l6.

16(15). Laminae less than 10 cm long; pinna lobes 2-4 mm long; stellate marginal lamina hairs
definitely stalked. Stipes 0.75 - 3 cm long, broadly alate toward the apex; laminae linear-lanceolate to
lanceolate, 0.8 -2.5 cm wide, the broad rachis ala often rather crisped, the stellate hairs abundant.

265. H. hirsutum

16(15). Laminae 13-23 cm long; pinna lobes less than 2 mm long; stellate marginal lamina hairs
subsessile. Stipes less than 1 cm long, broadly alate toward the apex; laminae linear to linear-
lanceolate, 1.5 - 2.5 cm wide, the broad rachis ala plane, the stellate hairs few.

270. H. mortonianum

17(15). Laminae (especially the stipe ala) slightly crisped. Stipes 1-2.5 cm long; laminae
lanceolate, 7 - 15 cm long, 1 - 2(4) cm wide; lumina of lamina cells large and clear.

260. H. crispum

17(15). Laminae plane..l8.

18(17). Ultimate segments ca. 0.6-1.25 mm wide; rachis ala not uniform, 0-0.25 mm wide. Stipes
1-3.5 cm long; laminae lanceolate, 2.5 -6.5 cm long, 1-2.5 cm wide.

274. H. semigiabrum

18(17). Ultimate segments 1-2 mm wide; rachis ala uniformly ca. 0.5 mm wide. Stipes up to 1 cm
long; laminae linear (in juveniles) to ovate, up to 3 cm long, 2 cm wide.

273. H. saenzianum

19(14). Pinnae, at least the proximal ones, petiolulate..21.

19(14). Pinnae sessile or adnate..20.

20(19). Marginal hairs stellate. Stipes 0.2 -2(3) cm long; laminae narrowly lanceolate to oblong,
2.5-7.5(11) cm long, 0.5 -2(3) cm wide.

261. H. elegans

20(19). Marginal hairs mostly simple. Stipes 1-3.5 cm long; laminae lanceate, 2.5 -6.5 cm long,
1 - 2.5 cm wide.

274. H. semigiabrum

21(19). Marginal hairs simple or forked at the base; veins glabrous; fronds ovate or oblong,
determinate. Stipes 1-3 cm long; laminae 4-9 cm long, 1.5-4 cm wide.

268. H. lineare

21(19). Marginal hairs stellate; veins with stellate hairs; fronds linear, indeterminate..22.

22(21). Stipe hairs mostly stellate; pinnae with 5-7 pairs of segments; costae completely alate.

272. H. pulchellum

22(21). Stipe hairs mostly simple; pinnae with 1-3 pairs of segments, or some of these overgrown
and frond-like; costae partially exalate.

277. H. trichophyllum

23(3). Laminae 2-pinnate or more divided..26.

23(3). Laminae pinnate-pinnatifid or less divided..24.

24(23). Laminae suborbicular, obtuse to truncate at the base, almost ternately divided with only
the basal lobes themselves lobed. Fronds less than 2.5 cm long; stipes 0.2 -0.8 cm long; ultimate

segments 1-2 mm wide.

279. H. brevifrons

24(23). Laminae triangular to lanceolate, acute to obtuse at the base, not ternately divided, with
several pairs of lobes themselves lobed..25.

25(24). Ultimate segments 2-2.5 mm wide; involucres ovate, obtuse at the base, round at the apex;
fronds 3 - 12(20) cm long.

278. H. asplenioides

25(24). Ultimate segments ca. 1.5 mm wide; involucres nearly trapezoidal, acute at the base and
apex; fronds 2-4 cm long.

277a. H. apiculatum

188 HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM

26(23). Laminae usually plane (except the segments often folded in H. nigrescens), truncate at the
base, ovate or lanceolate, widest near the base, often erect; rachises 1-4 times longer than the
stipes..29.

26(23). Laminae undulate or crisped (except in some H. myriocarpum), gradually attenuate toward
the base, narrowly elliptical or linear, widest near the middle, often pendent; rachises (2)3-7 times
longer than the stipes.. 27.

27(26). Ala uniform along the rachis between the pinnae; laminae plane or somewhat undulate.
Stipes alate to the base; ultimate segments constricted proximal to the involucre.

283. H. myriocarpum

27(26). Ala narrower distally between the pinnae; laminae undulate to crisped..28.

28(27). Laminae undulate, not strongly crisped, 5-30 cm long; lateral branches occasionally
overgrown and frond-like; ultimate segments constricted proximal to the involucre; involucres not
undulate, entire at the apex.

287. H. undulatum&ai+ low

28(21). Laminae strongly crisped, up to 7(10) cm long; lateral branches not overgrown and frond-
like; ultimate segments not constricted proximal to the involucre; involucres undulate, irregular at
the apex.

280. H. contortum

29(26). Stipes not at all alate. Stipes 1-8 cm long; laminae ovate to linear-lanceolate, 3.5-23 cm
long, 1.5-9 cm wide; piimae occasionally overgrown and somewhat frond-like.

281. H. costaricanum

29(26). Stipes alate at least at the apex and often to the base..30.

30(29). Involucres acute at the base; rachises usually with an ala ca. 0.1 mm wide..32.

30(29). Involucres obtuse to truncate at the base; rachises usually with an ala 0.25-0.5 mm
wide..31.

31(30). Segments and alae usually plane, sometimes the segments slightly folded when dry; pinnae
often not imbricate; involucres orbicular. Stipes (1.5)4 -8 cm long; laminae 3-pinnate-pinnatifid to 4-
pinnate, often ovate.

285. H. polyanthos

31(30). Segments and alae folded; piimae usually imbricate; involucres wider than long. Stipes
1-5(6) cm long; laminae 2-3-pinnate, often linear-lanceolate.

284. H. nigrescens

32(30). Fronds up to 12(23) cm long, 3-4-pinnate; involucres acute at the apex.

286. H. siliquosum

32(30). Fronds mostly 15-25 cm long, 3-5-pinnate; involucres obtuse to round at the apex.

282. H. farallonense

HYMENOPHYLLUM subg. HYMENOPHYLLUM

257. Hymenophyllum fucoides (Swartz) Swartz, J. Bot. (Schrader) 1800(2) :99.
1801.

Trichomanes fucoides Swartz, Nov. Gen. Sp. PI. Prodr. 136. 1788. TYPE: Jamaica, Swartz (S not
seen photo 6174; isotypes B-Hb. Willd. 20227-1 not seen Tryon photo, BM not seen photo 6590).

Hymenophyllum spinulosum H.B.K. Nov. Gen. Sp. 1:26 (fol. 21). 1816. TYPE: Mt. Avila, between
La Guaira and Caracas, Distr. Fed., Venezuela, 750 hexap elev., Humboldt & Bonpland (P not seen
microfiche S. 1. Library).

Hymenophyllum peruvianum Hook. & Grev. Icon. Fil. 2:t. 208. 1831. TYPE: Esmeraldas,

Ecuador, 5000 ft, Jameson (K not seen).

Hymenophyllum blepharodes K. Presl, Hymenophyllaceae 51 (postpr. 143). 1843. TYPE:

Martinique, Kohaut (PRC not seen).

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 189

Hymenophyllum fucoides yar.frigidum Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk.
Aid. V, 1:291 (repr. 139). 1849, as ''pigidaJ' TYPE: Cerro de Zempoaltepec, Edo. Oaxaca, Mexico,
8000 - 9000 ft, Liehmann (C not seen).

Hymenophyllum pedicellatum Kunze ex Klotzsch, Linnaea 20:439. 1847. TYPE: Merida, Edo.
Merida, Venezuela, Moritz 346 (B not seen; isotypes BR not seen photo 4834, US). Leptocionium
fucoides sensu Klozsch (Linnaea 18:533. 1844) is cited in synonymy.

Hymenophyllum podocarpon Fee, Crypt. Vase. Br6sil 1:196, t. 71, f. 3. 1869. TYPE: Organ
Mountains, Est. Rio de Janeiro, Brazil, Glaziou 3350 (P not seen photos 4579, 4580).

Hymenophyllum fucoides var. integrum Kuhn ex Christ in Pitt. Prim. FI. Costaric. 3(1):3. 1901.
TYPE: Guatemala, Bernoulli 340 (P not seen).

Plants epiphytic or rarely epipetric, at 400-3000 m elevation, in forests, from
the Cordillera Central, Cerro Carpintera, the Cordillera de Talamanca to Pcia.
Chiriqui, the Fila Costena near S. Vito de Java, Cerro Jefe, Cerro Campana (Pcia.
Panama), and Cerro Pirre above Cana. Also from Cuba, Jamaica, Hispaniola, the
Lesser Antilles, Mexico to Nicaragua, Trinidad and Venezuela to Bolivia and
Brazil.

Plants with the longest teeth on the valves and lamina margin have been
segregated as H. ectocarpon Fee, a species only dubiously distinct from H. fucoides.

258. Hymenophyllum tunbrigense (L.) J. E. Smith, M^m. Acad. Roy. Sci. (Turin)
5:418. 1793.

Trichomanes tunbrigense L. Sp. PI. 2:1098. 1753. TYPE: Uncertain; not Britain, collector

unknown (LINN 1253.5 not seen microfiche S. I. Library), which according to Proctor (Ferns Jam. 5.
1985) was not in LINN until after 1755.

Plants epiphytic, at 1500-3800 m elevation, in forests and scrub forests, from
the Cordillera Central and the Cordillera de Talamanca to Cerro Chirripo. Also
from the southeastern United States, Jamaica, Hispaniola, Mexico, Guatemala, El
Salvador, Venezuela, Colombia, Ecuador, Bolivia, Argentina, and Chile.

HYMENOPHYLLUM subg. LEPTOCIONIUM (K. PresI) Christ

259. Hymenophyllum consanguineum Morton, Contr. U. S. Natl. Herb. 29:163.
1947.

TYPE: Holcomb’s Trail above Boquete, 1450-1650 m, Maxon 5624 (US;
isotype NY).

Plants epiphytic on tree trunks, at 1200-2400 m elevation, in forests, from
Monteverde, the Cordillera Central, Cerro Carpintera, the Cordillera de
Talamanca to Pcia. Chiriqui, Cerro Tute (Pcia. Veraguas), and above Cana. Also
from Venezuela.

260. Hymenophyllum crispum H.B.K. Nov. Gen. Sp. 1:26 (fol. 22). 1816.

Sphaerocionium schiedeanum K. Presl, Hymenophyllaceae 60 (postpr. 152). 1843. TYPE: Mexico,
Schiede (PRC not seen).

Sphaerocionium crispum var. pilosum Klotzsch, Linnaea 18:537. 1845, nom. superfl. TYPE: A
renaming of Hymenophyllum crispum H.B.K., and so based on the type of that name.

1 Hymenophyllum ciliatum var. nudipes Kunze, Linnaea 18:351. 1845. TYPE: Mexico, Leibold 72
p. p. (LZ destroyed). This mixed collection may be partially referable to H. hirsutum.

1 Hymenophyllum divaricatum v. d. Bosch, Nederl. Kruid. Arch. 5(3): 190. 1863. TYPE:

Venezuela, Moritz (B not seen).

Hymenophyllum orbignianum v. d. Bosch, Nederl. Kruid. Arch. 5(3): 191. 1863. TYPE: Yuracare,
Bolivia, D’Orbigny 304 (P? not seen).

190 HYMENOPHYLLACEAE; 42. HYMENOPHYLLUM

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 191

Hymenophyllum crispum var. brasilianum F6e, Crypt. Vase. Br6sil 1:195, t. 71, f. 2. 1869, as
''brasiliana.” TYPE: Serra dos Orgaos, Est. Rio de Janeiro, Brazil, Glaziou 3347 (P not seen photo
4566).

Hymenophyllum constrictum Christ, Bull. Herb. Boissier II, 4:939. 1904. TYPE: Costa Rica,
Werckl^ in 1903 (P not seen; isotype US).

Hymenophyllum nitens Werckld ex Christ, Bull. Herb. Boissier II, 4:940. 1904, non R. Br., 1810.
SYNTYPES: Costa Rica, Werckli 249 (P not seen photo 17045) and 252 (P not seen); and Serra
Itatiaia, Brazil, 2100 m, Dusen 161 (P not seen photo 17047).

TYPE: Silla de Caracas, Distr. Fed., Venezuela, 990 hexap elev., Humboldt &
Bonpland (P not seen photo 4567).

Plants epiphytic, at 1500-2000 m elevation, in forests, from the Cordillera
Central. Also from Jamaica, Mexico, Guatemala, Venezuela, Colombia, Peru,
Bolivia, and Brazil.

261. Hymenophyllum elegans Spreng. Syst. Veg. ed. 16, 4:133. 1827.

Hymenophyllum cruegeri K. Muell. Bot. Zeitung (Berlin) 12:722. 1854. TYPE: Mt. Tocuche,
Trinidad, Purdie & Cmeger (B not seen fragm US).

Hymenophyllum delkatissimum Fee, Crypt. Vase. Bresil 2:86, t. 105, f. 1. 1873. TYPE: Serra dos
Orgaos, Est. Rio de Janeiro, Brazil, Glaziou 3591 (P not seen photo 4595; isotype US).

Hymenophyllum caudatellum Christ, Bull. Herb. Boissier II, 4:939. 1904. TYPE: Costa Rica,
Werckle in 1903 (P not seen; isotype US fragm NY).

Hymenophyllum elegans f. minor Morton, Contr. U. S. Natl. Herb. 29:154. 1947. TYPE: Sierra del
Libano, Sta. Marta mountains, Depto. Magdalena, Colombia, 1650 m, H. H. Smith 1100 (NY).

TYPE: Brazil, Sellow (LZ destroyed).

Plants epiphytic, at 1000-2800 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Pcia. Chiriqui. Also from Cuba,
Hispaniola, the Lesser Antilles, Guatemala, Trinidad, Venezuela, Colombia to
Bolivia, and Brazil.

262. Hymenophyllum elegantulum v. d. Bosch, Nederl. Kruid. Arch. 4:408. 1858.

Hymenophyllum buchtienii Rosenst. Repert. Spec. Nov. Regni Veg. 5:229. 1908. TYPE: Unduavi,

North Yungas, Depto. La Paz, Bolivia, Buchtien 897 (S not seen; isotype US).

Hymenophyllum elegantulum var. petiolulatum Morton, Contr. U. S. Natl. Herb. 29:171. 1947.
TYPE: Jayuya, Puerto Rico, 1200 m, Sargent 3144 (US).

TYPE: Plate 33A of Hooker’s “Species Filicum,” which was based on a
specimen collected at Pillzhum, Pcia. Canar, Ecuador {Jameson, K not seen;
isotype NY).

Plants epiphytic, at 1500-3300 m elevation, in forests, from the Cordillera
Central, Cerro Chirripd, and Cerro Horqueta (Pcia. Chiriqui). Also from Cuba,
Hispaniola, Puerto Rico, Mexico, Guatemala, Honduras, El Salvador, and
Venezuela to Bolivia.

This species has rather variable fronds 6-30 cm long; often the larger
specimens have petiolulate proximal pinnae. In my opinion such specimens
should not be recognized taxonomically.

FIGS. 260 - 265. Hymenophyllum. FIG. 260. Median pinna of H. crispum, Hinton 14306a, Mexico.
FIG. 261. Median pinna of H. elegans, Killip 5340. FIG. 262. Median pinna of H. elegantulum, Torres
R. 6. FIG. 263. Pinna of H. fragile, Skutch 3559. FIG. 264. Frond of H. hemipteron, Maxon 8464.
FIG. 265. Median pinna of H hirsutum, Lellinger 827.

192 HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM

263. Hymenophyllum fragile (Hedw.) Morton, Contr. U. S. Natl. Herb. 29:172.
1947.

Trichomanes fragile Hedw. Fil. Gen. Sp. t. 18. 1802. TYPE: Not stated; Hedwig’s original
herbarium is at G. Plate 18 is labelled but the accompanying, unpaged text is headed rigidum,
which must be taken as T. rigidum sensu Hedw., non Swartz, 1788.

Trichomanes hispidum Poir. Encyc. Meth. 8:71. 1808. TYPE: A renaming of T. rigidum sensu
Hedw., and so based on the type of T. fragile.

Hymenophyllum intercalatum Christ, Bull. Herb. Boissier II, 4:942. 1904. TYPE: Turrialba, Pcia.
Cartago, Pittier 13259ter (P not seen; isotype CR not seen).

Plants epiphytic, at (700)900-2300(3300) m elevation, in forests, from the
Cordillera Central, Cerro Carpintera, the Cordillera de Talamanca to Pcia.
Chiriqui, near the Escuela Agricola Alta Piedra, Sta. Fe, near Cana, and the upper
slopes of Alto del Buey. Also from Cuba, Jamaica, Hispaniola, Mexico,
Guatemala, Venezuela to Bolivia, and Brazil.

264. Hymenophyllum hemipteron Rosenst. Repert. Spec. Nov. Regni Veg. 22:4.
1925.

Hymenophyllum hemipteron f. acropteron Rosenst. Repert. Spec. Nov. Regni Veg. 22:5. 1925, as
''acroptera.” TYPE: La Palma, Pcia. S. Jose, Brade & Brade 851 (S not seen fragm US).

Hymenophyllum hemipteron f. minor Rosenst. Repert. Spec. Nov. Regni Veg. 22:5. 1925. TYPE:
Cisma, near S. Geronimo, Pcia. Alajuela, 2100 m, Brade & Brade 620 (S not seen fragm US; isotype
NY).

Hymenophyllum palmense Rosenst. Repert. Spec. Nov. Regni Veg. 22:5. 1925. TYPE: La Palma,
Pcia. S. Jose, 1400 m, Brade & Brade 600 (S not seen; isotype UC).

LECTOTYPE: Cerro Tablazo, Pcia. S. Jose, Brade & Brade 621 (S not seen
fragm US), chosen by Morton (Contr. U. S. Natl. Herb. 29:169. 1947).

Plants epiphytic on trunks, at 1000 - 2500(2800) m elevation, in forests, from the
Cordillera Central, Cerro Tablazo, and the Cordillera de Talamanca to Pcia.
Chiriqui.

The ultimate segments of this species are longer in relation to their width than
are those of H. elegantulum, and the plants are smaller.

265. Hymenophyllum hirsutum (L.) Swartz, J. Bot. (Schrader) 1800(2) :99. 1801,
van hirsutum.

Trichomanes hirsutum L. Sp. PI. 2:1098. 1753. TYPE: Plate 50B of Plumier’s “Traite...”, which is
based on a specimen collected by Plumier in the West Indies.

Trichomanes ciliatum Swartz, Nov. Gen. Sp. PI. Prodr. 136. 1788. TYPE: Jamaica, Swartz (S not
seen; isotypes B-Hb. Willd. 20222 not seen Tryon photo, BM not seen photo 6587, US).

Sphaerocionium vestitum K. Presl, Hymenophyllaceae 58 (postpr. 150). 1843. LECTOTYPE: Rio
de Janeiro, Est. Rio de Janeiro, Brazil, Beyrich (PRC? not seen), inferentially chosen by Morton
(Contr. U. S. Natl. Herb. 29:115. 1947).

Sphaerocionium grevilleanum K. Presl, Hymenophyllaceae 34 (postpr. 126). 1843. TYPE: Based
on H ciliatum sensu Hook. & Grev. (Icon. Fil. l:t. 35. 1827), which is based on a specimen collected
on St. Vincent (Guilding, K not seen).

Sphaerocionium commutatum K. Presl, Hymenophyllaceae 34 (postpr. 126). 1843. TYPE: Based
on H. horyanum sensu Raddi (PI. Bras. t. 79, f. 4. 1825), which is based on a specimen collected near
Rio de Janeiro {Raddi, FI not seen).

1 Hymenophyllum ciliatum var. nudipes Kunze, Linnaea 18:351. 1845. TYPE: Mexico, Leibold 72
p. p. (LZ destroyed; isotype B not seen). This mixed collection may be partially referable to H.
crispum.

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 193

IHymenophyllum ciliatum var. omifolium Kimze, Linnaea 21:240. 1848. TYPE: Surinam, Weigelt
(LZ destroyed).

Hymenophyllum remotum v. d. Bosch, Nederl. Kruid. Arch. 4:413. 1858. TYPE: A renaming of H.
ciliatum sensu Hook. & Grev. (Icon. Fil. l:t. 35. 1827), which is based on a specimen collected on St.
Vincent (Guilding, K not seen).

Hymenophyllum surinamense v. d. Bosch, Nederl. Kruid. Arch. 4:414. 1858, nom. superfl. TYPE:
Based ultimately on Trichomanes ciliatum Swartz, and so based on the type of that name.

Hymenophyllum gardnerianum Sturm in Mart. FI. Bras. 1(2):297. 1859. TYPE: Rio de Janeiro,
Est. Rio de Janeiro, Brazil, Gardner 213 (BR not seen; isotype P not seen photo 4563).

Hymenophyllum atrovirens Fee & L’Herminier in Fee, Hist. Foug. Antill. [Mem. Foug. 11]:120, t.
30, f. 4. 1866, non Colenso, 1844, nec Christ, 1904, nom. illeg. TYPE: Pitou Caraibe, Guadeloupe,
Germain in 1864 (P or RB not seen).

Hymenophyllum microcarpon Fee, Crypt. Vase. Br6sil 1:245, t. 69, f. 3. 1869, non H. microcarpum
Desv., 1827, nom. illeg. SYNTYPES: Rio de Janeiro, Est. Rio de Janeiro, Brazil, Glaziou 2268 (P
not seen photo 4558) and 33S6 (P not seen).

Hymenophyllum caulopteron Fee, Crypt. Vase. Br6sil 1:197, t. 70, f. 3. 1869. SYNTYPES: Serra
de Estrella, Est. Rio de Janeiro, Brazil, Glaziou 1713 (P not seen photo 4561) and 920 (P not seen
photo 4560; isosyntype NY not seen); and Rio de Janeiro, Est. Rio de Janeiro, Brazil, Glaziou 2269
(P not seen) and 2270 (P not seen photo 4559).

Hymenophyllum ulei Christ & Giesenh. Flora 86:85, f. 6, 7. 1899. TYPE: Teresopolis, Serra dos
Orgaos, Est. Rio de Janeiro, Brazil, 1000 m, Ule 4510 (presumably P not seen; isotypes L not seen
photo 2528, US). The locality stated in the original description is in error.

Hymenophyllum elatius Christ in Schwacke, PI. Nov. Mineir. 2:13. 1900. TYPE: S. Antonio, Est.
Sta. Catarina, Brazil, Ule 206 (P not seen).

Hymenophyllum dimorphum Christ, Bull. Herb. Boissier II, 4:941. 1904. TYPE: Alto de Mano
Tigre, basin of the Rio Diqms, Pcia. Puntarenas, 700 m, Pittier (P not seen).

Hymenophyllum ciliatum f. tuberosum Rosenst. Hedwigia 46:74. 1906, as '‘tuberosa.” TYPE: A
renaming of H. ulei Christ & Giesenh., and so based on the type of that name.

Hymenophyllum ciliatum var. abbreviatum Rosenst. Hedwigia 56:360. 1915, as ''abbreviataP

TYPE: Ribeira, Est. S. Paulo, Brazil, Brade 5169 (S not seen).

Plants epiphytic on trunks, at 0-2000 m elevation, in forests, from the
Cordillera de Tilaran, the Cordillera Central, the Cordillera de Talamanca, the
Fila Costeha, Loma Prieta (Pcia. Los Santos), Cerro Jefe, and Punta S. Francisco
Solano and La Trojita (Depto. Choco). Also from the Antilles, Mexico to
Nicaragua, Trinidad, Venezuela to Bolivia, the Guianas, and Brazil.

266. Hymenophyllum horizontale Morton, Contr. U. S. Natl. Herb. 29:181. 1947.
TYPE: La Palma, ca. 1600 m, Standley 38114 (US fragm GH).

Plants epiphytic, at (300)1300-1700 m elevation, in forests, from near S.
Ramon, La Palma, La Hondura, and Carrillo (all Pcia. S. Jose), and the vicinity of
Orosi and Juan Vihas (both Pcia. Cartago).

267. Hymenophyllum lanatum F6e, Hist. Foug. Antill. [M6m. Foug. 11]:116, t. 31,
f.3. 1866.

TYPE: Sainte-Rose, Guadeloupe, UHerminier in 1861 (presumably P not seen
photo 4592; isotype MO not seen photo 13016).

Plants epiphytic, at 1400-1800 m elevation, in the Flora area known only from
the upper slopes of Alto del Buey {Lellinger & de la Sota 299, COL, CR, LP, US).
Also from the Antilles, Mexico, Guatemala, Venezuela, Colombia, and Surinam.

194 HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 195

268. Hymenophyllum lineare (Swartz) Swartz, J. Bot. (Schrader) 1800 (2): 100.
1801.

Trichomanes lineare Swartz, Nov. Gen. Sp. PI. Prodr. 137. 1788. TYPE: Jamaica, Swartz (S not
seen photo 6191; isotype B-Hb. Willd. 20221 not seen Tryon photo).

Hymenophyllum elegantissimum F6e, Hist. Foug. Antill. [Mem. Foug. 11]:118, t. 29, f. 2. 1866.
TYPE: Guadeloupe, UHerminier (P not seen photos 4572, 4573).

Hymenophyllum catherinae Hook, ex B2iker, Syn. Fil. 67. 1867. TYPE: Catherine’s Peak, Jamaica,
Wilson 573 (K not seen fragm US).

Hymenophyllum durandii Bommer & Christ, Bull. Herb. Boissier 4:657. 1896. TYPE: La Palma,
Pcia. S. Jos6, 1500“ 1700 m, Tonduz 9693 (BR not seen photo 4831; isotype CR).

Hymenophyllum lineare var. dussii Christ ex Duss, Foug. Antill. Frang. 9. 1903. TYPE:

Guadeloupe, 1115” 1350 m, Duss 4387 (B or P not seen; isotype US).

Hymenophyllum ceratophylloides Christ, Bull. Herb. Boissier II, 4:942. 1904. LECTOTYTE:
Costa Rica, Werckl6 280 (P not seen), chosen by Lellinger (Proc. Biol. Soc. Wash. 89:716. 1977).

Plants epiphytic, at 900-1600 m elevation, in forests, from Monteverde, the
Cordillera Central, the Fila Costeha, and the upper slopes of Alto del Buey. Also
from Cuba, Jamaica, Puerto Rico, the Lesser Antilles, and Guatemala.

269. Hymenophyllum microcarpum Desv. Mdm. Soc. Linn. Paris 6:333. 1827.

Hymenophyllum beyrichianum Kunze, Linnaea 9:108. 1834. TYPE: Pampayacu, Depto. Huanuco,
Peru, July 1829, Poeppig (LZ destroyed).

Hymenophyllum organense Hook. Sp. Fil. 1:90, t. 32B. 1844. TYPE: Organ Mountains, Est. Rio
de Janeiro, Brazil, Gardner 210 (K not seen; isotype P not seen photo 4597).

Hymenophyllum pteropodum v. d. Bosch, Nederl. Kruid. Arch. 5(3):187. 1863. SYNTYPES:
Quito, Pcia. Pichincha, Ecuador, Cuming 23 (B not seen) and Jameson 109 (K not seen).

Hymenophyllum lindigii Mett. in Tr. & Planch. Ann. Sci, Nat. Bot. V, 2:1% (repr. 278). 1864.
TYPE: Fusagasuga, Depto. Cundinamarca, Colombia, Lindig 122 (B not seen).

Hymenophyllum notabile F6e, Crypt. Vase. Br6sil 1:193, t. 69, f. 2. 1869. SYNTYPES: Brazil,
Glaziou 1714 (P not seen photo 4602) and 2271 (P not seen photo 4603).

Hymenophyllum contractile Sodiro, Anales Univ. Quito 6(45):151 (repr. 22). 1892. TYPE:

Nanegal, Pcia. Pichincha, Ecuador, Sodiro (Hb. Sodiro not seen fragm US).

Hymenophyllum kaieteurum Jenm. Ferns Brit. W. Ind. Guiana 15. 1898. TYPE: “Forests of the
Potaro river and other regions,” Guyana, /e/imc/i (NY).

Hymenophyllum an^stifrons Christ, Bull. Herb. Boissier II, 4:940. 1904. SYNTYPES: Canas
Gordas, Pcia. Pimtarenas, Pittier 10984 (P not seen; isotype US); and P^amo de Guanacas, Depto.
Cauca, Colombia, 2000-2500 m, Lehmann 6414 (P? not seen; isotypes UC, US).

Hymenophyllum microcarpum var. lanceolatum Morton, Contr. U. S. Natl. Herb. 29:163. 1947.
TYPE: Vara Blanca de Sarapiquf, Pcia. Heredia, 1340 m, Skutch 3639 (US).

TYPE: Hispaniola, collector unknown (P-Hb. Desv. not seen photo BM not
seen).

Plants epiphytic on trunks or rarely epipetric, at (400)900-2000 m elevation, in
forests, from the Cordillera Central, the Cordillera de Talamanca to Pcia.
Chiriquf, the Fila Costena, near Sta. Fe, Cerro Pirre, and Alto del Buey and the
upper Rio S. Juan region. Also from the Greater Antilles, Dominica, Mexico,
Guatemala, Honduras, Nicaragua, Venezuela to Bolivia, Guyana, and Brazil.

FIGS. 266 - 272. Hymenophyllum. FIG. 266. Proximal median pinna of H. horizontale, Brade &
Brade 627. FIG. 267. Frond of H lanatum, Sherring, Grenada. FIG. 268. Median pinna of H.
lineare, Brade & Brade 61. FIG. 269. Median pinna oiH. microcarpum, Stork 4822. FIG. 270. Pinna
of H. mortonianum, Lellinger & de la Sota 425. FIG. 271. Median pinna of H. plumosum, Maxon
5693. FIG. 272. Median pinna of H. pulchellum, Lankester 648b.

196 HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 197

270. Hymenophyllum mortonianum Lellinger, Proc. Biol. Soc. Wash. 98:385, f. 15.
1985.

TYPE: Mojarras de Tado, 8.5 km east of Istmina, Depto. Choco, 150-250 m,
Lellinger & de la Sola 425 (US; isotypes COL, CR, HU A, LP).

Plants epiphytic, at 100-300 m elevation. Known only from the type.

271. Hymenophyllum plumosum Kaulf. Enum. Fil. 267. 1824.

Sphaerocionium aureum K. Presl, Hymenophyllaceae 57 (postpr. 149). 1843. TYPE; Serra da
Estrella, Est. Rio de Janeiro, Brazil, Beyrich (Hb. Beyrich not seen).

TYPE: Brazil, Chamisso (LE? not seen).

Plants epiphytic, at (1000)1200-2300 m elevation, in forests, from the
Cordillera Central, the valley of the Rio Grande de Orosi (Pda. Cartago), and the
Cordillera de Talamanca to Pda. Chiriqm. Also from Colombia to Bolivia, and
Brazil.

272. Hymenophyllum pulchellum Schlechtend. & Cham. Linnaea 5:618. 1830.

Hymenophyllum chrysothrix Sturm in Mart. FI. Bras. 1(2):298. 1859. TYPE; Serra da Estrella,
Est. Rio de Jcineiro, Brazil, Martins (BR not seen photo 4828).

Hymenophyllum silveirae Christ in Schwacke, PI. Nov. Minier. 2; 14. 1900. SYNTYPES (all from
Est. Minas Gerais, Brazil); Serra do Campestre, Silveira 2332 (P not seen); Serra das Camarinhas,
Schwacke 11085 (P not seen); and Itacolumi, Schwacke 12528 (P not seen photo 4619).

Hymenophyllum pannosum Christ, Bull. Herb. Boissier II, 5;250. 1905. TYPE; Costa Rica,

Werckle (P not seen; isotype US fragm NY).

Hymenophyllum lineare var. brasiliense Rosenst. Hedwigia 46;74. 1906. TYPE; Sta. Cruz, Est.
Rio Grande do Sul, Brazil, Juergens 137 {Ros. Fil Austrobras. Exs. 175) (S not seen; isotypes NY,
US).

Hymenophyllum lineare f. tuberosum Rosenst. Hedwigia 46;74. 1906, as ''tuberosa” TYPE; Rio
Grande, Est. S. Paulo, Brazil, Wacket {Ros. Fil Austrobras. Exs. 175) (S not seen; isotype US).

Hymenophyllum lineare f. pseudocarpum Rosenst. Hedwigia 46;74. 1906, as ''pseudocarpa.”

TYPE; Rio Grande, Est. S. Paulo, Brazil, Wacket {Ros. Fil Austrobras. Exs. 176) (S not seen; isotype
US).

TYPE: Cuesta Grande, Jalacingo, Edo. Veracruz, Mexico, Schiede & Deppe 807
(B-Hb. Willd. 20239 not seen microfiche S. I. Library; isotypes BM, LE neither
seen).

Plants epiphytic, at 1600 - 1900 m elevation, in the Flora area known only from
Las Nubes, Pda, S. Jose {Knight, US) and from Pacayas, Pda. Cartago {Lankester
648b, US). Also from Mexico, Guatemala, Belize, Honduras, and Brazil.

273. Hymenophyllum saenzianum L. D. Gdmez, Revista Biol. Trop. 17:107, f. 7.
1970.

TYPE: Madreselva, Cordillera de Talamanca, Pda. Cartago, 2800 m, Gomez
PtC-47568 (CR).

Plants epiphytic on trunks, at 1200-2800 m elevation, in forests, from the
northern end of the Cordillera de Talamanca.

FIGS. 273 - 277. Hymenophyllum. FIG. 273. Frond of H. saenzianum, Mickel 3452. FIG. 274.
Median piima of H. semiglabmm, Cornman 1175. FIG. 275. Proximal median pinna of H. sieberi,
Muench 11, Mexico. FIG. 276. Median pinna of H. subrigidum, Scamman 5853. FIG. 277. Median
pinna of H. trichophyllum, Farrar CR-225.

198 HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM

The type specimen is juvenile; more fully developed fronds are ovate, with ca. 6
alternate, simple to 2-forked pinnae. The ultimate segments are unusual in being
up to 2 mm wide.

274. Hymenophyllum semiglabrum Rosenst. Repeit. Spec. Nov. Regni Veg. 9:67.
1910.

TYPE: La Palma, Pcia. S. Jose, 1400 m, Brade & Brade 395 (Ros. Fil Costar.
Exs. 174) (S not seen photo 6193 fragm US; isotypes NY, UC).

Plants epiphytic, at 1000-1500 m elevation, in forests, known only from the
type, from Sta. Clara de Cartago, Pcia. Cartago (Maxon 8178 p. p.y US), and from
the vicinity of Boquete (Comman 1175 , US).

275. Hymenophyllum sieberi (K. Presl) v. d. Bosch, Nederl. Kruid. Arch 4:414.
1858.

Sphaerocionium sieberi K. Presl, Hymenophyllaceae 58 (postpr. 150). 1843. TYPE: Martinique,
Kohaut (Sieber exs. 71) (PRC not seen).

Hymenophyllum wercklei Christ, Bull. Herb. Boissier II, 4:940. 1904. LECTOTYPE: Costa Rica,
Werckli 247 (P not seen; possible isolectotype US fragm NY), chosen by Morton (Contr. U. S. Natl.
Herb. 29:179. 1947).

Plants epiphytic, at (300)900 - 1700 m elevation, in forests, from the Cordillera
Central, the valley of the Rio Reventazon, and Platanillo (Pcia. Cartago). Also
from Puerto Rico, the Lesser Antilles, Mexico to Nicaragua, and Venezuela.

276. Hymenophyllum subrigidum Christ, Bull. Herb. Boissier H, 5:260. 1905.

Hymenophyllum atrovirens Christ, Bull. Herb. Boissier II, 4:941. 1904, non Colenso, 1844, nec Fee
& L’Herminier in Fee, 1866, nom. illeg. TYPE: Costa Rica, Werckle in 1903 (P not seen; isotype US
fragm NY).

TYPE: A renaming of H. atrovirens Christ, and so based on the type of that
name.

Plants epiphytic on trunks, at (800)1400-2400 m elevation, in forests, from the
Cordillera Central, the Cordillera de Talamanca to Pcia. Chiriqui, and Cerro
Pirre.

This species is peculiar in that the sinuses between adjacent segments or
pinnules are not so deep as is usual in Hymenophyllum , and so the dichotomies of
the lamina have a webbed appearance.

277. Hymenophyllum trichophyllum H.B.K. Nov. Gen. Sp. 1:27 (fol. 22). 1816.

Hymenophyllum procerum v. d. Bosch, Nederl. Kruid. Arch. 4:409. 1858, nom. nud. Based on H
pulchellum sensu Mett. (Fil. Lechl. 1:25. 1859), which lacks a description.

Hymenophyllum moritzianum Sturm in Mart. FI. Bras. 1(2):295. 1859. TYPE: Near Merida, Edo.
Merida, Venezuela, Moritz 344 p. p. (BR? not seen).

Hymenophyllum eriophorum v. d. Bosch, Nederl. Kruid. Arch. 5(3):180. 1863. TYPE: “Andes of
Quito,” Pcia. Pichincha, Ecuador, /amcjo/i 82 (not seen).

Hymenophyllum trichophyllum var. contractum Hieron. Hedwigia 45:226. 1906, as ''contractu.”
TYPE: Mt. Tolima, Colombia, Stuebel 43 (B not seen).

TYPE: Between Cocollar and El Guardia de S. Agustin, probably Edo.
Monagas, Venezuela, Humboldt & Bonpland (P not seen).

Plants epiphytic, at 1500-3500 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Pcia. Chiriqui. Also from Guatemala,
Venezuela to Bolivia, and Guyana.

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 199

HYMENOPHYLLUM subg. MECODIUM Copel.

277a. Hymenophyllum apiculatum Mett. ex Kuhn, Linnaea 35:391. 1868.

TYPE: Near Colonia Tovar, Edo. Aragua, Venezuela, Fendler 32 (B not seen;
isotype US).

Plants epiphytic, at 1000 m elevation, in the Flora area known only from Cerro
Jefe (Valdespino, Aranda & Rios 344b, PMA not seen, US; Valdespino & Aranda
171, PMA not seen, UC). Also from Venezuela and Colombia.

278. Hymenophyllum asplenioides (Swartz) Swartz, J. Bot. (Schrader) 1800(2) :98.
1801.

Trichomanes asplenioides Swartz, Nov. Gen. Sp. PI. Prodr. 136. 1788. TYPE; Jamaica, Swartz (S
not seen photo 6160; isotype B-Hb. Willd. 20217 not seen Tryon photo).

Plants epiphytic on trunks, at 1000-2000 m elevation, in forests, from the
Cordillera Central, Cerro Carpintera, and the Cordillera de Talamanca from
Copey de Dota (Pcia. S. Jose) to the vicinity of Boquete. Also from the Greater
Antilles, Mexico, Guatemala, Honduras, Nicaragua, Trinidad, Venezuela, Guyana,
and Brazil.

279. Hymenophyllum brevifrons Kunze, Bot. Zeitung (Berlin) 5:185. Mar 1847.

Hymenophyllum tablaziense Christ, Bull. Soc. Bot. Gen6ve II, 1:216. 1909. TYPE: Cerro Tablazo,
1900 m, Brade 155 (P not seen; isotypes UC, US).

TYPE: French Guiana, Leprieur 235 (LZ destroyed; isotype B not seen, isotype
fragms NY, US). See Kunze (Farrnkrauter 1:236, t. 96, f. 2. Sept 1847) for a
statement of the type.

Plants epiphytic, at 100-1000(1900) m elevation, in forests, from the Cordillera
de Tilardn, north of S. Ramon, Finca La Selva (Pcia. Heredia), Cerro Tablazo, the
vicinity of S. Isidro del General, and the Canal Zone and adjacent Pcia. Panama.
Also from Cuba, Hispaniola, Guatemala, Belize, Guyana, and French Guiana.

This species is allied to H. brevistipes Liebm. of Mexico and Honduras, which
has the stipes narrowly alate almost to the base, and to H. abruptum Hook, of the
Greater Antilles, which has fronds that are a little larger and less divided.

280. Hymenophyllum contoitum v. d. Bosch, Nederl. Kruid. Arch. 5(3): 170. 1863.
TYPE: Aguacate, Pcia. S. Jose, Hoffmann (B not seen).

Plants epiphytic, at 900-1500 m elevation, in forests, from Aguacate, Peralta,
and La Palma (Pcia. S. Jose), Vara Blanca (Pcia. Heredia), and Platanillo and
near Juan Vinas (both Pcia. Cartago). Also from Hispaniola, Mexico, and
Guatemala.

This species is closely related or possibly conspecific with the South American
H. fendlerianum Sturm in Mart. (FI. Bras. 1(2):291. 1859), which has longer and
narrower fronds with even more contorted segments that seem invariably to be the
host of a black, bristle-like fungus.

281. Hymenophyllum costaricanum v. d. Bosch, Nederl. Kruid. Arch. 5(3):161.
1863.

Hymenophyllum alfredii Rosenst. Report. Spec. Nov. Regni Veg. 22:4. 1925. TYPE: Cisma, near
S. Jer6nimo, Pcia. Alajuela, 2100 m, Brade & Brade 601 (S not seen; isotypes NY, UC, US).

200 HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 201

Hymenophyllum costaricanum var. emarginatum Rosenst. Repert. Spec. Nov. Regni Veg. 22:4.
1925. TYPE: La Palma, Pcia. S. Jose, 1450 m, Brade <& Brade 608 (S not seen photo 6163; isotypes
UC, US).

TYPE: Volcan Barba, Pcia Heredia, Hoffmann (B not seen; isotypes L not seen
photo 2478, P not seen photo 4564, UC, US).

Plants epiphytic on trunks, at (400)1000-2900 m elevation, in forests, from the
Cordillera Central, Cerro Tablazo, the Cordillera de Talamanca to Pcia. Chiriqui,
and northwest of Sta. Fe.

This species is often misidentified as H. polyanthos, but differs in having entirely
exalate stipes. However, it is possible that a few specimens from the Flora area
could be referable to H. polyanthos var. protrusum (see a comment under H.
polyanthos). The pinnae of H. costaricanum are less crowded and often its laminae
are longer and more linear than are those of H. polyanthos var. polyanthos. Two
small, sterile specimens found at 200 - 900 m elevation in the Choco {Killip 35339
and Lellinger & de la Sota 195, both US) appear to be a distinct species related to
H. costaricanum, but are insufficiently known to be described.

282. Hymenophyllum farallonense Hieron. Bot. Jahrb. Syst. 34:430. 1904.

TYPE: Farallones de Cali, Depto. Valle, Colombia, 2000 m, Lehmann 1980 (B
not seen; isotype US).

Plants epiphytic, at 900-1800 m elevation, in forests, in the Flora area known
only from Alto del Buey {Lellinger & de la Sota 263, LP, US, and 319, COL, LP,
US).

This species is closely related, and may be conspecific with, H. trianae Hieron.
(Bot. Jahrb. Syst. 34:429. 1904), which is said to have longer and wider ultimate
segments and larger involcures.

283. Hymenophyllum myriocarpum Hook. Sp. Fil. 1:106, t. 37D. 1844.

TYPE: Colombia, Hartweg 1530 (K not seen; isotypes BM not seen, P not seen
photo 4601).

Plants epiphytic, at 1500-3300 m elevation, in forests and open areas, from S.
Pedro de S. Ramon (Pcia. Alajuela), the Cordillera Central, and the Cordillera de
Talamanca to Pcia. Chiriqui. Also from Mexico, Guatemala, Honduras to
Nicaragua, and Venezuela to Bolivia.

284. Hymenophyllum nigrescens Liebm. Kongel. Danske Vidensk. Selsk. Skr.,
Naturvidensk. Afd. V, 1:292 (repr. 140). 1849.

Sphaerocionium nigricans K. Presl ex Klotzsch, Linnaea 18:536. 1845, non Hymenophyllum

nigricans Colla, 1836. SYNTYPES: Colonia Tovar, Edo. Aragua, Venezuela, Moritz 268 (B not seen)
and 268b (B not seen); and Peru, Dombey 87 (B? not seen; isotype P not seen).

Hymenophyllum atrosanguineum v. d. Bosch, Nederl. Kruid. Arch. 5(3): 165. 1863. TYPE:

Aguacate, Pcia. S. Jose, Hoffmann (B not seen).

Hymenophyllum laciniosum Christ, Bull. Herb. Boissier II, 4:938. 1904. TYPE: Costa Rica,
Werckle (P not seen). A paratype from Volcan Sotara, Depto. Cauca, Colombia {Lehmann 3671, P
not seen; isoparatype US) is neither H nigrescens nor H. polyanthos.

FIGS. 278 - 281. Hymenophyllum. FIG. 278. Frond of H. asplenioides, Killip 5399. FIG. 279. Frond
of H. brevifrons, Brade 155. FIG. 280. Median pinna of H. contortum, Lankester 592. FIG. 281.
Median pinna of H. costaricanum, Maxon 5576.

202

HYMENOPHYLLACEAE; 42. HYMENOPHYLLUM

HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM 203

Hymenophyllum camosum Christ, Bull. Herb. Boissier II, 4:938. 1904. TYPE: Costa Rica,

Werckle in 1903 (P not seen photo 17048).

1 Hymenophyllum nigrescens var. gracile Rosenst. Meded. Rijks-Herb. 19:4. 1913, as ''gracilis”
TYPE: Cordillera de Sta. Cruz, Depto. Sta. Cruz, Bolivia, Herzog 3559/a (B? not seen). I have not
seen any undoubted specimens of H. nigrescens from Colombia to Bolivia, and so this variety may be
a synonym of H. polyanthos.

Hymenophyllum contextum Rosenst. Repert. Spec. Nov. Regni Veg. 22:3. 1925. LECTOTYPE:
La Palma, Pcia. S. Jose, 1400 m, Brade & Brade 5 (US; isolectotype S not seen), chosen by Lellinger
(Mem. New York Bot. Card. 38:14. 1984).

LECTOTYPE: Near Chinautla, Edo. Puebla, Mexico, 7000 ft, Liebmann FI
Mex. 537 (C not seen; isolectotypes BM, K, P none seen), chosen by A. R. Smith
(FL Chiapas 2:133. 1981).

Plants epiphytic on trunks and branches, at (500)1200-2600(3300) m elevation,
in forests and open areas, from the Cordillera de Tilaran, the Cordillera Central,
Cerro Carpintera, Cerro Tablazo, the Fila de Cedral, the Cordillera de Talamanca
to Cerro Chirripo, and northwest of Sta. Fe. Also from Mexico, Honduras,
Venezuela, and perhaps Colombia, Peru, Bolivia, and Brazil.

For a comment on the taxonomy of this species, see H. polyanthos.

285. Hymenophyllum polyanthos (Swartz) Swartz, J. Bot. (Schrader) 1800(2): 102.

1801, var. polyanthos.

Trichomanes polyanthos Swartz, Nov. Gen. Sp. PI. Prodr. 137. 1788. TYPE: Jamaica, Swartz, (S
not seen photo 6178; isotypes B-Hb. Willd. 20235, 20237 not seen Tryon photos, BM not seen photo
6586).

ITrichomanes clavatum Swartz, Nov. Gen. Sp. PI. Prodr. 137. 1788. TYPE: Jamaica, Swartz (SBT
not seen). Most of the Jamaican material determined as H. polyanthos has exalate stipes, in contrast
to the Central and South American specimens.

Hymenophyllum jalapense Schlechtend. & Cham. Linnaea 5:619. 1830. TYPE: Near Jalapa, Edo.
Veracruz, Mexico, Schiede [812] (B not seen; isotypes BM not seen, L not seen photo 2514, LE not
seen).

Hymenophyllum millefolium Schlechtend. & Cham. Linnaea 5:620. 1830. TYPE: Jalacingo, Edo.
Veracruz, Mexico, Schiede & Deppe [513] (B not seen; isotypes BM not seen, HBG? not seen, fragm
US).

Hymenophyllum g^evilleanum K. Presl, Hymenophyllaceae 32 (postpr. 124). 1843. TYPE: Plate
128 of Hooker and Greville’s ‘Tcones Filicum,” which illustrates specimens collected on Jamaica
(Swartz, S not seen) and on St. Vincent (Guilding, K not seen).

Hymenophyllum fratemum K. Presl, Hymenophyllaceae 54 (postpr. 146), t. 12B. 1843. TYPE:
Jamaica, collector unknown (PRC not seen).

Hymenophyllum poeppigianum K. Presl, Hymenophyllaceae 54 (postpr. 146). 1843. TYPE:

Pampayacu, Depto. Huanuco, Peru, Poeppig (PRC not seen).

Sphaerocionium breutelii K. Presl, Gefassbundel Farrne 29 (postpr. 357), t. 5, f. 19. 1847. TYPE:
Mt. Miseri, St. Kitts, Breutel (PRC not seen).

Hymenophyllum imbricatum Kunze ex Sturm in Mart. H. Bras. 1(2):287. 1859, non Blume, 1828,
nom. illeg. SYNTYPES: Prov. Sebastianopolis, Est. Rio de Janeiro, Brazil, Martius & Langsdorff
(BR not seen); and Serra do Cubatao, Est. S. Paulo, Brazil, Lindberg (BR not seen).

FIGS. 282 - 287. Hymenophyllum. FIG. 282. Median pinna of H. farallonense, Lehmann 1980,
Colombia. FIG. 283. Median pinna of H. myriocarpum, Chrysler 5113. FIG. 284. Median pinna of
H nigrescens, Knight. FIG. 285. Median pinna of H. polyanthos var. polyanthos, Stork 4823. FIG.
286. Median pinna of H. siliquosum, Maxon 7984. FIG. 287. Median pinna of H. undulatum,
Standley 35202.

204 HYMENOPHYLLACEAE: 42. HYMENOPHYLLUM

Hymenophyllum pusillum Schott ex Sturm in Mart. FI. Bras. 1(2):289. 1859. SYNTYPES: Ilha
Sta. Catarina, Est. Sta. Catarina, Brazil, Chamisso (BR not seen); Serra do Cubatao, Est. S. Paulo,
Brazil, Sellow (BR not seen); Est. Goias, Brazil, Pohl 1419 (BR not seen); and Mt. Kanuku, Guyana,
Rich. Schomhurgk 1180 (B not seen).

Hymenophyllum sturmii v. d. Bosch, Nederl. Kruid. Arch. 5(3):152. 1863. SYNTYPES: Serra dos
Orgaos, Est. Rio de Janeiro, Brazil, ‘'Gaudichaud, Vauthier, Beyrich alii” (none seen).

Hymenophyllum fecundum v. d. Bosch, Nederl. Kruid. Arch. 5(3): 153. 1863. TYPE: Aguacate,
Pcia. S. Jos6, Hoffmann (B not seen).

Hymenophyllum botryoides v. d. Bosch, Nederl. Kruid. Arch. 5(3): 160. 1863. TYPE: Mexico,
Galeotti 6394 (BR or K not seen photo BM not seen; isotype US).

1 Hymenophyllum matthewsii v. d. Bosch, Nederl. Kruid. Arch. 5(3): 162. 1863. SYNTYPES: Peru,
Matthews (L? not seen; isosyntypes P not seen photos 4607, 4611); and Quito, Pcia. Pichincha,
Ecuador, Cuming (B not seen). Pinnules atypical for H. polyanthos.

Hymenophyllum viridissimum F6e, Crypt. Vase. Br6sil 1:194, t. 49, f. 3. 1869. SYNTYPES: Brazil,
Glaziou 1718 and 2050 (both P not seen photo 4609).

Hymenophyllum mazei Fourn. ex Christ in Krug in Urban, Bot. Jahrb. Syst. 24:85. 1897.

SYNTYPES: Guadeloupe, Maze 461 (P? not seen) and 651 p. p. (P? not seen).

Hymenophyllum polyanthos var. reductum Jenm. Ferns Brit. W. Ind. Guiana 8. 1898. TYPE: Not
stated.

Hymenophyllum subdeltoideum Fee ex Christ, Bull. Herb. Boissier II, 2:322. 1902. TYPE: Est.
Sta. Catarina, Brazil, Ule 3351 (P not seen).

Hymenophyllum trichomanoides var. subalatum Rosenst. Repert. Spec. Nov. Regni Veg. 22:4.
1925. TYPE: Cerro Tablazo, 1900 m, Brade & Brade 396 (S not seen; isotype US).

Mecodium mexiae Copel. Univ. Calif. Publ. Bot. 19:294, t. 48. 1941. TYPE: Near confluence of
the Rio Cayumba with the Rio Huallaga, Depto. Huanuco, Peru, 875 m, Mexia 8282 (MICH not
seen; isotype MO).

Plants epiphytic on trunks and branches, at (600)900-3300 m elevation, in
forests and open areas, from the Cordillera Central, Cerro Tablazo, the Cordillera
de Talamanca to Pcia. Chiriquf, the valley of the Rfo General, the Peninsula de
Osa, Cerro Jefe, Cerro Pirre, and the northern half of the Choco. Also from
throughout tropical America.

This species and H. nigrescens are not strongly separated, and are part of a large
and variable complex that is much in need of critical study. Proctor (FI. Less.
Antill. 2:76-77. 1977) distinguished H. polyanthos V2S. protrusum (Hook.) Farw.,
with pendent, elongate, apparently indeterminate laminae, as seen in Valerio 214
(US) from Volcdn Barba and in Michel 3220 (US) from the Cerro de la Muerte.
The tendency toward indeterminateness occurs sporadically in Costa Rican
material, and in my opinion these specimens are not worthy of taxonomic
recognition (although the Antillean ones very well may be). Therefore, I have
excuded synonyms of wdiV.protrusum from the synonymy.

286. Hymenophyllum siliquosum Christ, Bull. Herb. Boissier II, 4:938. 1904.

LECTOTYPE: Costa Rica, Werckle 289 (P not seen), chosen by Lellinger (Proc.
Biol. Soc. Wash. 89:716. 1977). The other syntype, Costa Rica, Werckle 307 (P not
seen; isosyntype CR) is H. lineare, a species of subg. Leptocionium.

Plants epiphytic, at (1100)1500-1700 and 2800-3200 m elevation, in forests,
from La Palma and vicinity (Pcia. S. Jose), the Cerro de la Muerte, and Cerro
Chirripo. Also from Mexico.

HYMENOPHYLLACEAE: 43. TRICHOMANES 205

287. Hymenophyllum undulatum (Swartz) Swartz, J. Bot. (Schrader) 1800(2);101.

1801.

Trichomanes undulatum Swartz, Nov. Gen. Sp. PI. Prodr. 137. 1788. TYPE: Jamaica, Swartz (S
not seen photo 6172; isotypes B-Hb. Willd. 20238 not seen Tryon photo, BM not seen, P not seen
photo 4623).

Hymenophyllum flaccidum v. d. Bosch, Nederl. Kruid. Arch. 4:400. 1858, nom. superfl. TYPE:
Based on H. undulatum sensu Hedw. (Fil. Gen. Sp. t. XI. 1799), which was based on T. undulatum
Swartz.

Hymenophyllum undulatum var. regenerans C. Chr. in Aspl. Ark. Bot. 20A(7):8. 1926. TYPE:
Between S. Felipe and El Chaco, S. Yungas, Depto. La Paz, Bolivia, 2500 m, Asplund 1439 (C or S
not seen).

Plants epiphytic, at 1200-3300 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca. Also from the Greater Antilles, Mexico
to Honduras, Venezuela, Colombia, Ecuador, perhaps Bolivia, and Brazil.

43. TRICHOMANES L.

Plants epiphytic, epipetric, or terrestrial; rhizomes erect or ascending and
strongly rooted or long-creeping and with rather inconspicuous roots, hairy, the
hairs multicellular, often bristle-like, dark brown, borne especially at the apex;
fronds minute (in subg. Didymoglossum) to small or rarely medium-sized, usually
monomorphic; stipes obsolete to longer than the laminae, commonly alate distal
to the base, sometimes glabrous except for hairs at the base, or with pale, simple
hairs (in subg. Achomanes) or blackish, simple, bifid, or stellate hairs (in subg.
Didymoglossum); laminae simple to decompound, orbicular or flabellate to ovate
or oblong, for the most part only 1 cell thick, the rachis commonly entirely alate or
the laminae flabelliform and the midrib obsolete, usually glabrous on the lamina
surface and often on the veins, or with hairs like those of the stipes; veins
pinnately or flabellately branched, sometimes connected by or accompanied by
thinner, false veins parallel to the lamina margin or to the true veins; sori
marginal, borne at pinna apices or laterally on the pinnae; sporangia borne on a
long, exserted setiform receptacle surrounded by a tubular, sometimes 2-lipped
involucre that is free to almost wholly immersed in the lamina tissue; sporangia
short-stalked.

Mostly pantropical in wet lowlands; ca. 320 species.

LELLINGER, D. B. 1984. Hymenophyllaceae (Filicales) in B. Maguire and collaborators. The
botany of the Guayana Highland - Part XII. Mem. New York Bot. Gard. 38:9 - 46.
MORTON, C. V. 1968. The genera, subgenera, and sections of the Hymenophyllaceae. Contr. U. S.
Natl. Herb. 38:153 - 214.

WESSELS BOER, J. G. 1962. The New World species of Trichomanes sect. Didymoglossum and
Microgonium. Acta Bot. Neerl. 11:277 - 330.

1. Fronds simple to pinnate-pinnatifid; venation catadromous; sori terminating terminal segments
of pinnae and pinnules; segment margins often ciliate or with sessile, stellate hairs..3.

1. Fronds 2-pinnate to 4-pinnate-pinnatifid; venation anadromous; sori terminating lateral
segments of pinnae and pinnules; segment margins usually not ciliate or with stellate hairs..2.

2(1). Rhizomes long-creeping, the fronds distant; plants usually epiphytic; stipes 0.5- 1(1.5) mm in
diam. (subg. Trichomanes). .6.

2(1). Rhizomes erect or short-creeping, the fronds clustered; plants usually terrestrial; stipes
0.75-2.5 mm in diam. (subg. Pachychaetum)..4.

3(1). Fronds less than 5(8) cm long, simple to bipinnatifid but not truly pinnate; false veins present
between the true veins or submarginad; plcmts epiphytic or rarely epipetric (subg.
Didymoglossum). .30.

206 HYMENOPHYLLACEAE: 43. TRICHOMANES

3(1). Fronds more than 5 cm long, deeply pinnatifid to 2-pinnate-pinnatifid; false veins absent
(except in T. pinnatum and apparently in T. ankersii and T. tuerckheimii); plants often terrestrial
(subg. Achomanes).. 13.

4(2). Rachises entirely alate, the ala more than 0.5 mm \vide; stipes 1-2.5 mm in diam.; laminae 3-
pinnate, 15 - 25 cm long, iridescent blue-green in life, ca. 3 cells thick.

320. T. elegans

4(2). Rachises not or slightly alate, the ala less than 0.5 mm wide; stipes 0.25-1.25 mm in diam.;
laminae 3-pinnate to 4-pinnate-pinnatifid, 5-15(18) cm long, 1 cell thick..5.

5(4). Stipes 0.75-1.25 mm in diam., usually not alate; cells of the lamina with occluded lumina;
involucres conical, 1-1.5 mm long, 0.75-1 mm wide.

321. T. rigidum

5(4). Stipes 0.5 -0.6 mm in diam., always alate; cells of the lamina with clear lumina; involucres
conical, 0.75-1 mm long, 0.4 -0.5 mm wide.

322. T. spnicei

6(2). Segments normally expanded, mostly more than 0.5 mm wide, the cells of the lamina in more
than 6 rows on each side of the midrib..8.

6(2). Segments narrow to filiform, up to 0.5 mm wide, the cells of the lamina usually in 2-6 rows
on each side of the midrib.. 7.

7(6). Segments filiform, the lamina cells in often only 2 or 3 rows on each side of the midrib;
involucres alate and not on naked axes. Laminae reduced nearly to the veins; fronds 3-4-pinnate;
ultimate segments mostly less than 0.5 cm long.

323. T. capillaceum

7(6). Segments narrow, the lamina cells usually in 4-6 rows on each side of the midrib; involucres
exalate and on naked axes. Fronds 3 - 4-pinnate; ultimate segments usually less than 0.5 cm long.

322a. T. angustatum

8(6). Fronds 5- 15 cm long; rhizomes ca. 0.5 mm in diam. .11.

8(6). Fronds (10)15-50 cm long; rhizomes 1-2 mm in diam..9.

9(8). Sterile lamina lobes usually broad and low, (1)2-4 mm wide; laminae pinnate-piimatifid to 2-
pinnate-pinnatifid. Stipes 0.2 -1(2) cm long; involucres flared at the apex.

329. T. mpestre

9(8). Sterile lamina lobes narrow and long, 0.25-0.5 mm wide; laminae 2-pinnate to 4-pinnate..l0.

10(9). Involucres not or scarcely flared at the apex; laminae broad, usually truncate at the base;
stipes 3 - 12 cm long.

328. T. radicans

10(9). Involucres decidedly flared at the apex; laminae narrow, usually tapered at the base; stipes
1 - 6 cm long.

324. T. collariatum

11(8). Lamina segments slightly folded parallel to the veins; involucres ca. 2 mm long, 1 mm wide,
conical, only slightly flared at the apex.

327. T. pyxidiferum

11(8). Lamina segments plane, not folded parallel to the veins; involucres ca. 1.5 mm long, 0.5 mm
wide, tubular, greatly flared at the apex.. 12.

12(11). Stipes mostly alate throughout; pinnae approximate to imbricate; ala of the rachis 0.25-0.5
mm wide; fertile fronds mostly 3-pinnate, usually 2-6(8) sori per pinna.

325. T. diaphanum

12(11). Stipes not alate or alate only at the apex; pinnae distant; ala of the rachis 0.1-0.25 mm
wide; fertile fronds mostly 2-pinnate-pinnatifid, usually 1(3) sori per pinna and only the basal
acroscopic pinna segments fertile.

326. T. hymenophylloides

13(3). Laminae not cihate, the hairs of the costae and stipe bases simple or stellate and coarse..23.

13(3). Laminae ciliate, at least on the midrib and veins.. 14.

HYMENOPHYLLACEAE: 43. TRICHOMANES 207

14(13). Cilia stellate or forked at the base. Laminae pinnatifid, acuminate at the base, usually
exstipitate, 5- 15 cm long, 1.5-2 cm wide.

301. T. polypodioides

14(13). Cilia simple..l5.

15(14). Cilia with a dark, bulbous base; laminae 2.5 -8.5 cm long, 1-1.5 cm wide, pinnate-
pinnatifid, truncate at the base, decidedly stipitate.

290. T. crinitum

15(14). Cilia without a markedly bulbous, dark base; laminae pinnate or pinnatifid.. 16.

16(15). Laminae pinnatifid or pinnate, the sterile lobes or pinnae entire or nearly so.. 18.

16(15). Laminae bipinnatifid or 2-pinnate, the sterile lobes or pinnae crenate, serrate, or more
dissected.. 17.

17(16). Laminae 2-pinnate. Stipes and laminae densely sericeous with reddish hairs; laminae 6-30
cm long, (1.5)3 -8 cm wide, pendent; stipes 2-5(8) cm long, not alate.

297. T. lucens

17(16). Laminae bipinnatifid, the lobes crenate or serrate. Laminae 3.5-15(25) cm long, 1.5-3
cm wide, slightly narrowed at the base; stipes 1.5 -3.5(6) cm long, alate to the base.

293. T. delicatum

18(16). Stipes narrowly alate to the base; rachis ala on each side as wide or wider than the rachis.
Plants 3-15(25) cm long; laminae lanceate, usually truncate at the base, pinnatifid, the lobes never
imbricate.

295. T. galeottii

18(16). Stipes not alate or narrowly alate only at the apex; rachis ala on each side narrower than
the rachis or absent.. 19.

19(18). Involucres 1/3 -2/3 immersed in the lamina tissue..22.

19(18). Involucres fully to 2/3 immersed in the lamina tissue..20.

20(19). Laminae not much narrowed at the base; proximal pinnae not deflexed or sur current.
Plants 5 - 30(40) cm long; involucres 2-4 times longer than wide, tubular, somewhat to greatly flared
at the apex, 1-10 per pinna.

291. T. crispum

20(19). Laminae decidedly narrowed at the base, and the proximal piimae deflexed and
surcurrent..21.

21(20). Stipes entirely exalate. Plants 40 - 60 cm long; pinnae nearly acute at the apex.

300a. T. plumosum

21(20). Stipes very narrowly alate. Plants 25 - 45 cm long; pinnae nearly round at the apex.

299a T. pellucens

22(19). Involucres flared at the apex; rhizomes very long-creeping, branched, the fronds scattered;
pinnae obtuse or round at the apex, truncate or rarely cordate at the basiscopic base. Plants
(15)25 - 60 cm long.

298. T. ludovicinum

22(19). Involucres not flared at the apex; rhizomes erect or ascending, unbranched, the fronds
crowded; pinnae acute at the apex, cordate and overlapping the rachis at the basiscopic base. Plants
15-30 cm long.

292. T. dactylites

23(13). Fertile and sterile fronds monomorphic..26.

23(13). Fertile and sterile fronds dimorphic, the linear fertile fronds exceeding the sterile ones..24.

24(23). Rachis of fertile fronds broadly alate; involucres immersed in the lamina tissue. Plants
15 - 30(45) cm long; rachis of sterile fronds occasionally flagelliform and with a terminal, proliferous
bud.

294. T. diversifrons

24(23). Rachis of fertile fronds not alate; involucres free..25.

208 HYMENOPHYLLACEAE: 43. TRICHOMANES

25(24). Sterile laminae pectinate; sterile rachis never prolonged or with a terminal, proliferous
bud. Plants 7-15(25) cm long.

299. T. osmundoides

25(24). Sterile laminae pinnate; sterile rachis often prolonged and with a terminal, proliferous bud.
Plants 3 - 10 cm long.

289. T. botryoides

26(23). Laminae piimate to tripinnatisect, not flagelliform at the apex; false veins absent or parallel
to the true veins; plants hemiepiphytic (except T. kalbreyeri)..7S.

26(23). Laminae pinnate, occasionally flagelliform and with a proliferous bud at the apex; plants
not hemiepiphytic.. 27.

27(26). False veins present at right angles to the true veins. Plants terrestrial; rhizomes erect.

300. T. pinnatum

27(26). False veins absent between the true veins. Plants terrestrial; rhizomes erect.

296. T. hostmannianum

28(26). Rhizomes erect, thick; laminae coarsely pinnate-pinnatifid; rachis hairs at least partially
stellate; plants low trunk epiphytes.

296a. T. kalbreyeri

28(26). Rhizomes very long-creeping; laminae pinnate to tripinnatisect; rachis hairs simple; plants
hemiepiphytic with the laminae and long-creeping rhizomes appressed to smooth tree trunks..29.

29(28). Pinnae 7-12 mm wide, almost entire to crenate, (6)10-50 pairs per frond.

302. 7. tuerckheimii

29(28). Pinnae 3-6 mm wide, serrately lobed, 5-25 pairs per frond.

288. T. ankersii

30(3). Fronds with paired, pale, marginal, scale-like projections. Plants 2.5 -6.5 cm long; stipes
obsolete; laminae simple, spathulate to irregularly lobed.

311. 7. membranaceum

30(3). Fronds without paired, marginal, scale-like projections. Laminae simple to bipinnatifid..31.

31(30). Fronds with blackish, stellate marginal hairs, lacking a submarginal false vein; hps of the
involucres usually with blackish margins..34.

31(30). Fronds glabrous on the margin, with or without a submarginal false vein; involcures
scarcely with two hps, lacking a dark margin.. 32.

32(31). False veins between the true veins reticulate, with cross veins; well-developed fertile fronds
obovate to nearly orbiculate. Laminae up to ca. 2 cm in diam.

306. 7. godmanii

32(31). False veins between the true veins linear, without cross veins; well developed fertile fronds
oblong or more elongate..33.

33(32). Fronds with a band of marginal cells about as wide as the continuous, submarginal false
vein; distal part of fronds irregularly and often deeply lobed.

305. 7. ekmanii

33(32). Fronds with a bcmd of marginal cells narrower than the discontinuous, submarginal false
vein; distal part of fronds only slightly lobed.

309. 7. kapplerianum

34(31). Venation of sterile fronds distinctly pinnate, with the costae extending to the frond
apex. .41.

34(31). Venation of sterile fronds flabellate, with the costae absent or not extending beyond the
middle of the frond..35.

35(34). Fronds mostly more than 1 cm long, firmly membranaceous, usually with several sori..37.

35(34). Fronds mostly less than 1 cm long, thin and translucent, usually with a single, apical
sorus..36.

36(35). Involucres without dark-edged hps; costa absent in sterile fronds.

312. 7. nummularium

HYMENOPHYLLACEAE: 43. TRICHOMANES 209

36(35). Involucres with dark-edged lips; costa present at the base of sterile laminae.

313. T. ovale

37(35). Marginal hairs stellate; veins usually with no more than 5 rows of cells between them..39.

37(35). Marginal hairs simple or forked at the base; veins usually with at least 12 rows of cells
between them. .38.

38(37). Laminae usually wider than long, nearly round, cordate to obtuse at the base, entire or
slightly crenate distally and at the round apex; cells between the veins elongate parallel to the true
veins.

319. T. rhipidophyllum

3S(31). Laminae usually longer than wide, lanceolate, elliptic, or oblanceolate, obtuse at the base,
deeply lobed to bipinnatifid distally (except nearly entire in juveniles); cells between the veins
isodiametric.

308. T. hymenoides

39(37). Fronds irregularly split along the veins into several irregular segments; sori (5)6 or more
per lamina; involucres entirely exerted beyond the lamina tissue.

304. T. curtii

398(37). Fronds entire or only slightly split at maturity; sori 4(6) or fewer per lamina; involucres
partially immersed in the lamina tissue (except in T. punctatum subsp. labiatum).. 40.

40(39). Lips of the involcures as wide as the tube, rimmed with 1 or 2 rows of darkened cells.

316. T. punctatum subsp. sphenoides

40(39). Lips of the involcures wider than the tube, rimmed with 2-4 rows of dsu'kened cells.

315. T. punctatum subsp. labiatum

41(34). Lips of the involucres lacking rows of darkened marginal cells.

314. T. petersii

41(34). Lips of the involucres with 1 or more rows of darkened marginal cells (except in juveniles
of T. hymenoides).. 42.

42(41). Fronds pinnatifid, bipinnatifid, or pinnate-pinnatifid with stellate hairs mainly in the
sinuses and simple or forked hairs on the margins..44.

42(41). Fronds entire or irregularly lobed distally with stellate hairs throughout and simple or
forked hairs absent..43.

43(42). Fronds 0.5 -1.5 cm long, 0.1 -0.3 cm wide, usually linear to hnear-oblong; lips of the
involucres with a single row of darkened cells.

303. T. angustifrons

43(42). Fronds 1.5 -2.5 cm long, 0.4- 1.2 cm wide, linear with shght lobes or obovate, irregularly
lobed distally; lips of the involcures with several rows of darkened cells.

317. T. pusillum

44(42). Lips of the involcures ovate-lanceolate, at least twice as long as wide; laminae densely hairy
on the rachis. Laminae usually bipinnatifid or pinnate-pinnatifid, tapered gradually at the base.

307. T. gourlianum

44(42). Lips of the involucres subcircular, often wider than long; laminae not densely hairy on the
rachis..45.

45(44). Ftdse veins submarginal, absent between the true veins; lips of the involucres usually with
2-4 rows of darkened marginal cells.

310. T. krausii

45(44). False veins not submarginal, present between the true veins; lips of the involcures usually
with 1 row of darkened marginal cells..46.

46(45). Sori 3-8(16) per frond; laminae 2-5.5 cm long; false veins (1)2 -3(4) between adjacent
true veins.

318. T. reptans

46(45). Sori 1-3 per frond; laminae 1-3 cm long; false veins 0- 1(2) between adjacent true veins.

308. T. hymenoides

ymwr-

210 HYMENOPHYLLACEAE; 43. TRICHOMANES

HYMENOPHYLLACEAE: 43. TRICHOMANES 21 1

TRICHOMANES subg. ACHOMANES K, PresI

288. Trichomanes ankersii Parker ex Hook. & Grev. Icon. Fil. 2(ll):t. 201. 1831.

Trichomanes commutatum Sturm in Mart. FI. Bras. 1(2);261. 1859. SYNTYPES: Tanaii, Rio
Acara, Est. Par^, Brazil, Spruce 414 (B not seen); and Mt. Canuku, Guyana, Rich. Schomburgk 1215
p. p. (B not seen).

Trichomanes guianense Sturm in Mart. FI. Bras. 1(2):262. 1859. TYPE; Demerara, Guyana, Rich.
Schomburgk 1215 p. p. (B not seen).

TYPE: Demerara, Guyana, Ankers (K not seen Kramer photo).

Plants epiphytic on trunks, at 0-300 m elevation, in forests, from the Atlantic
coastal plain of Costa Rica, the Peninsula de Osa, Sta. Rita ridge (Pcia. Colon),
the El Llano - Carti road (Pcia. Panama), the Rio Tuquesa (Pcia. Darien), and the
Choco. Also from Nicaragua, Venezuela to Bolivia, Brazil, and the Guianas.

289. Trichomanes botryoides Kaulf. Enum. Fil. 263. 1824.

Feea nana Bory, Diet. Class. Hist. Nat. 6:446. 1824; 17:t. 47, f. 1. 1831. TYPE: French Guiana,
Poiteau (P not seen).

TYPE LOCALITY: Guyana.

Plants epipetric, at 300-400 m elevation, in the Flora area known only from the
trail from Rio Blanco del Norte to Cano Sucio, Pcia. Code (Knapp 3743, MO,
US). Also from Venezuela, Colombia, Guyana, and Surinam.

290. Trichomanes crinitum Swartz, Nov. Gen. Sp. PL Prodr. 136. 1788.

Trichomanes Themtinieri Fee, Hist. Foug. Antill. [Mem. Foug. 11]: 107, t. 29, f. 1. 1866.

SYNTYPES: Ste. Rose, St. Luis, and Goyave, Guadeloupe, L’Herminier in 1861 (all P or RB none
seen).

TYPE: Jamaica, Swartz (SBT not seen fragm US photo 6198; isotypes B-Hb.
Willd. 20198 not seen Tryon photo, BM not seen photo 6592, S not seen).

Plants epiphytic or occasionally terrestrial on rotting logs, at (700)1000-2400 m
elevation, in forests, from north of S. Ramon, the Cordillera Central, the
Cordillera de Talamanca above Cartago and above Boquete, near Sta. Fe, Cerro
Jefe, between Cerro Jefe and La Eneida (Pcia. Panama), and Cerro Pirre. Also
from Cuba, Jamaica, the Lesser Antilles, Venezuela, Colombia, Ecuador, and
Peru.

291. Trichomanes crispum L. Sp. PL 2:1097. 1753.

Trichomanes haenkeanum K. Presl, Hymenophyllaceae 36, 65 (postpr. 128, 157). 1843. TYPE:
Based on T. crispum sensu K. Presl (Reliq. Haenk. 1:69. 1825), and so based on the type of that
name, which is Mountains of Hu^uco, Depto. Hudnuco, Peru, Haenke (PRC not seen).

Trichomanes schomburgkii v. d. Bosch, Nederl. Kruid. Arch. 5(2):148. 1861. TYPE: Berbice,
Guyana, Schomburgk 442 (L not seen; isotypes K not seen photo 19069, US).

Trichomanes crispum var. remotum Fee, Hist. Foug. Antill. [Mem. Foug. 11]: 106. 1866. TYPE:
Guadeloupe, VHerminier [792?] in 1864 (P or RB not seen).

FIGS. 288 "-297. Trichomanes. FIG. 288. Basal portion of pinna of T. ankersii, Cuatrecasas 16607.
FIG. 289. Plant of T. botryoides, Elias 5, Colombia. FIG. 290. Frond of T. crinitum. Lent 2088. FIG.
291. Median segments of T. crispum, Pittier 4211. FIG. 292. Median pinnae of T. dactylites,
Cuatrecasas 17381, Colombia. FIG. 293. Plant of T. delicatum, Killip & Cuatrecasas 39022,
Colombia. FIG. 294. Fertile and sterile fronds of T. diversifrons, Maxon 5735. FIG. 295. Median
pinnae of T. galeottii, Lellinger 1234. FIG. 296. Frond of T. hostmannianum, Haught 1335,
Colombia. FIG. 297. Frond of T. lucens, Valerio III.6.

212 HYMENOPHYLLACEAE: 43. TRICHOMANES

Trichomanes badium Fourn. Bull. Soc. Bot. France 15:144, 147, 148. 1868. LECTOTYPE: Cuba,
Wright 900 (P not seen Weatherby photo; isotypes GH not seen, US), chosen by Proctor (FI. Jamaica
127. 1985).

Trichomanes lindi^i Fourn. Bull. Soc. Bot. France 15:144, 147, 148. 1868. SYNTYPES:

Venezuela, Fendler 289 (P not seen) and 389 p. p. (P not seen); and Muzo, Depto. Boyaca,
Colombia, 700 m, Lindig 140 (P not seen; isosyntype US).

Trichomanes mode Fourn. Bull. Soc. Bot. France 15:147, 148. 1868. TYPE: Guadeloupe, Est.
Bahia, Brazil, Blachet (P not seen).

Trichomanes cristatum var. macrothrix F6e, Crypt. Vase. Br6sil 1:186. 1869. TYPE: Rio de

Janeiro, Est. Rio de Janeiro, Brazil, Glaziou 2246 (P or RB not seen).

Trichomanes imbricatum Sodiro, Anal. Univ. Quito 6(44):70. 1892. TYPE: Near Sto. Domingo de
los Colorados, Pcia. Pichincha, Ecuador, 400 m, Sodiro (Hb. Sodiro not seen; isotypes P, GH neither
seen).

LECTOTYPE: Plate 86 of Plumier’s “Traite...”, which is based on a specimen
collected by Plumier on Martinique, chosen by Proctor (FI. Less. Antill. 2:95.
1977).

Plants epiphytic on trunks or rarely terrestrial, at 0-1700 m elevation, in
forests, from Monteverde, north of S. Ramon, the Cordillera Central, the
Cordillera de Talamanca, the valley of the Rio General, Sta. Fe, Pcia. Panama
east of the Canal Zone, Cana and Cerro Pirre, and the northern part of the Choco.
Also from the Antilles, Mexico, Guatemala, Nicaragua, Trinidad, Venezuela,
Colombia, Peru, Bolivia, Brazil, the Guianas, Uruguay, and Paraguay.

292. Trichomanes dactylites Sodiro, Anales Univ. Quito 6(44):70 (repr. 12). 1892.

Trichomanes elatum Desv. M6m. Soc. Linn. Paris 6:327. 1827, non Forst., 1786, nom. illeg.
TYPE: “Habitat in America calidiori,” collector unknown (P not seen Cintract photo). Synonymized
by Weatherby (Contr. Gray Herb. 114:35. 1936).

Trichomanes digitatum Sodiro, Recens. Crypt. Vase. Quit. 4. 1883, non Swartz, 1806, nom. illeg.
TYPE: Near the confluence of the Rio Napa with the Rio Pilat6n, Pcia. Pichincha, Ecuador, 900 m,
Sodiro (Hb. Sodiro not seen; presumable isotype US).

TYPE: A renaming of T. digitatum Sodiro, and so based on the type of that
name.

Plants epiphytic on trunks, at 900-1800 m elevation, in forests, from Cerro Jefe,
Cerro Tacarcuna, Alto del Buey, and near the suspension bridge at ca. Km. 141 of
the Ciudad Bolivar - Quibdo road (Depto. Choco). Also from Venezuela,
Colombia, and Ecuador.

293. Trichomanes delicatum v. d. Bosch, Nederi. Kruid. Arch. 5(2): 145. 1861.

Trichomanes daguense Weath. Contr. Gray Herb. 95:36. 1931. TYPE: Dagua Valley, Cordoba,
Depto. El Valle, Colombia, 80-100 m, Killip 5251 (US; isotypes GH, NY).

TYPE: Quito, Pcia. Pichincha, Ecuador, Cuming 21 (B not seen fragm L not
seen).

Plants epiphytic, usually on tree trunks, at 0-300 m elevation, in forests, from
throughout the Choco. Also from coastal Colombia south of the Choco.

294. Trichomanes diversifrons (Bory) Mett. ex Sadeb. in Engl. & Prantl, Nat.
Pflanzenfam. 1(4): 108. 1899.

Hymenostachys diversifrons Bory, Diet. Class. Hist. Nat. 8:462. 1825; 17:t. 42, f. 2. 1831. TYPE:
French Guiana, Poiteau (P not seen).

HYMENOPHYLLACEAE: 43. TRICHOMANES 213

Trichomanes elegans Rudge, PI. Guian. 1:24, t. 35. 1805, non Rich., 1792, nom. illeg. TYPE:
French Guiana, Martin (BM ex Hb. Rudge not seen; isotype NY not seen).

Feea boryi v. d. Bosch, Nederl. Kruid. Arch. 4:347. 1858, nom. superfl. TYPE: A rencuning of
Hymenostachys diversifrons Bory, and so based on the type of that name.

Trichomanes dimorphum Mett. ex Sadeb. in Engl. & Prantl, Nat. Pflanzenfam. 1(4): 108. 1899.
TYPE: Not stated.

Plants terrestrial, at 0-300(1000) m elevation, in forests, from the Atlantic and
Pacific lowlands of Costa Rica and Panama, and the Choco. Also from Mexico,
Guatemala, Honduras, Belize, Nicaragua, and tropical South America.

295. Trichomanes galeottii Foum. Bull. Soc. Bot. France 15:148. 1868.

Trichomanes killipii Weath. Contr. Gray Herb. 95:36, t. 8. 1931. TYPE: Buenaventura, Depto. El
Valle, Colombia, 0~ 10 m, Killip 11727 (GH not seen; isotypes NY not seen, US).

LECTOTYPE: Edo. Oaxaca, Mexico, 3000 ft, Galeottii 6530 (BM not seen;
isolectotypes BR not seen photos 4801 and 19847, K not seen photo 19063, P not
seen Weatherby photo), chosen by A. R. Smith (FI. Chiapas 2:242. 1981).

Plants epiphytic on trunks, at (0)300-1900 m elevation, in forests, from the
Cordillera de Tilardn, the Cordillera de Talamanca, the valley of the Rio General,
the Peninsula de Osa, above Cahazas (Pda. Veraguas), Cerro Jefe and vicinity,
and the mouth of the Rio S. Juan. Also from Cuba, Mexico to Honduras, and
Nicaragua.

296. Trichomanes hostmannianum (Klotzsch) Kunze, Bot. Zeitung (Berlin) 5:352.
1847.

Neurophyllum hostmanninanum Klotzsch, Linnaea 18:532. 1845. TYPE: Surinam, Hostmann 75
(B not seen; isotypes K not seen photos 19086, 19087, NY, U not seen photo 126, US).

Neuromanes abruptum v. d. Bosch, Nederl. Kruid. Arch. 4:350. 1858, nom. superfl. TYPE: A
renaming of Neurophyllum hostmannianum Klotzsch, and so based on the type of that name.
Neurophyllum abruptum Fee, which was also cited, is a nomen nudum.

Trichomanes huberi Christ, Bull. Herb. Boissier 6:992. 1898. TYPE: Cununy [Cunanf], Terr.
Amapa, Brazil, Huber 1099 (P not seen Weatherby photo).

Plants terrestrial, at 0 - 100 m elevation, in forests, in the Flora area known only
from the lower Rio S. Juan region of the Choco {Forero 3943, 4842, COL, MO,
US). Also from Venezuela, Colombia, Peru, Brazil, and the Guianas.

This species resembles the more common T. pinnatum but lacks the cross-veins
between the main veins that characterize T. pinnatum.

296a. Trichomanes kalbreyeri Baker, J. Bot. Brit. For. 19:203. 1881.

TYPE: Depto. Antioquia, Colombia, 6500 ft, Kalbreyer 1857 (K not seen;
isotype US).

Plants epiphytic, on fallen logs, at 1100-1500 m elevation, in the Flora area
known only from the Rio S. Rafael, Atlantic slopes of Volcan Barba {Grayum et al
7753, MO not seen, UC), and the Fortuna dam site, Pcia. Chiriqui {van der Werff
& van Hardeveld 67580, MO not seen, UC). Also from Colombia.

297. Trichomanes lucens Swartz, Nov. Gen. Sp. PI. Prodr. 136. 1788.

Trichomanes lambertianum Hook. Sp. Fil. 1:139, t. 41B. 1846. TYPE: Pillao, Depto. Huanuco,
Peru, Ruiz & Pavon (OXF not seen; isotype K not seen photo 19075).

Trichomanes splendidum v. d. Bosch, Nederl. Kruid. Arch. 4:360. 1858. SYNTYPES: Bogota,
Distr. Esp., Colombia, Hartweg 1531 (L not seen); and Peru, Lechler (L not seen).

214 HYMENOPHYLLACEAE: 43. TRICHOMANES

HYMENOPHYLLACEAE: 43. TRICHOMANES 215

Trichomanes auratum F6e, Crypt. Vase. Br6sil 1:186, t. 67, f. 1. 1869. TYPE: Brazil, Glaziou 2465
(P not seen).

TYPE: Jamaica, Swartz (SET not seen; isotypes B-Hb. Willd. 20197 not seen
Tryon photo, BM not seen photo 6593S not seen photo 6200, S not seen photo
6200).

Plants epiphytic on trunks, at 1500-2700 m elevation, in forests, from
Monteverde, the Cordillera Central, and the Cordillera de Talamanca to Pcia.
Chiriqui. Also from Jamaica, Venezuela, Colombia to Bolivia, and Brazil.

298. Trichomanes ludovicinum Rosenst. Repert. Spec. Nov. Regni Veg. 22:6.
1925.

TYPE: La Palma, Pcia. S. Jose, Brade & Brade 64 (S not seen; isotypes HB not
seen, NY, P not seen Weatherby photo, S not seen photo 6201, US).

Plants epiphytic on trunks or terrestrial, at (700)1000-2000 m elevation, in
forests, from the Cordillera de Tilaran, the Cordillera Central, Moravia (Pcia.
Limon), the Rio Cacao in the valley of the Rio General, the Cordillera de
Talamanca to Pcia. Chiriqui, near Sta. Fe, and Cerro Pajita, north of El Valle.

299. Trichomanes osmundoides DC. ex Poir. Encyc. M^th. 8:65. 1808.

Trichomanes spicisonim Desv. Ges. Naturf. Freunde Berlin Mag. 5:329, t. 7, f. 7. 1811. TYPE:

French Guiana, collector unknown (holotype or isotype FI not seen photo 16577), synonymized by
Desvaux (Mem. Soc. Linn. Paris 6:326. 1827).

Feea polypodina Bory, Diet. Class. Hist. Nat. 6:446. 1824; 17:t. 43. 1831. TYPE LOCALITY:
Guadeloupe.

Trichomanes spicatum Hook. Sp. Fil. 1:114. 1846. LECTOTYPE: Portland, Jamaica, Purdie (K
not seen), chosen by Proctor (Ferns Jamaica 8. 1985).

Trichomanes mougeotii v. d. Bosch, Nederl. Kruid. Arch. 4:378. 1858, as “mougeoti.” TYPE:
French Guiana, Leprieur (L not seen; possible isotype US).

Trichomanes platyrachis Domin, Rozpr. Krai. Ceske Spolecn. Nauk, Tf. Mat.-Prir. 2 [Pterid.
Dominica]:45, t. V, f. 1-3. 1929. TYPE: Retreat Plantation, John Crow Hill near Bath, Jamaica,
3500 ft, N. Wilson (K not seen; isotype BM).

TYPE: Guyana, collector unknown (P-Hb. DC. not seen; probable isotypes B-
Hb. Willd. 20192 not seen Tryon photo, FI not seen photo 16578).

Plants terrestrial, at (0)200-700 m elevation, in forests, from the valley of the
Rio General, the Peninsula de Osa, Santa Rita ridge (Pcia. Colon), Cerro Jefe,
Cerro Azul, and near La Eneida (all Pcia. Panama), and the Rio Atrato basin.
Also from Cuba, Jamaica, Hispaniola, the Lesser Antilles and Trinidad, French
Guiana, Venezuela, and Colombia.

299a. Trichomanes pellucens Kunze, Linnaea 9:104. 1834.

TYPE: Tocache, upper Rio Huallaga, Depto. Huanuco, Peru, Jun 1830, Poeppig
(LZ destroyed; isotype BR not seen).

FIGS. 298 - 307. Trichomanes. FIG. 298. Median pinna segments of T. ludovicinum, Standley &
Valerio 49060. FIG. 299. Fertile £md sterile fronds of T. osmundoides, Lellinger & de la Sota 471.
FIG. 300. Frond of T pinnatum, Erlanson 63. FIG. 301. Plant of T. polypodioides, Skutch 4596.
FIG. 302. Basal portion of pinna of T. tuerckheimii, Skutch 3079. FIG. 303. Frond of T. angustifrons,
Gomez 6565. FIG. 304. Plant of T. curtii, Mickel 2790. FIG. 305. Plant of T. ekmanii, Wagner 31.
FIG. 306. Plant of T. godmanii, Maxon 6871. FIG. 307. Frond of T. gourlianum, Liesner 1171.

216 HYMENOPHYLLACEAE: 43. TRICHOMANES

Plants terrestrial or epipetric, rarely epiphytic on low trunks, at 500? and 1300 m
elevation, in the Flora area known only from the slopes of Cerro Kikirchabata,
Pcia. Limon {Gomez et al 23802, CR not seen, UC) and from the hills near
Moravia, Pcia. Limon (L. O. Williams 16185, US). Also from Guatemala,
Venezuela and Guyana, Colombia to Bolivia, and Brazil.

300. Trichomanes pinnatum Hedw. Fil. Gen. Sp. t. 5, f. 1. 1799.

Trichomanes rhizophyllum Cav. Descr. PI. 279. 1801. TYPE: Panama, Nee (MA not seen),
synonymized by Christensen (Dansk Bot. Ark. 9(3) :28. 1937).

Trichomanes floribundum Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:505. 1810, nom. superfl.
TYPE: A renaming of T. pinnatum Hedw., and so based on the type of that name.

Trichomanes longifolium Desv. Ges. Naturf. Freimde Berlin Mag. 5:328. 1811. TYPE: “Habitat
in America calidiore,” collector unknown (P-Hb. Desv. not seen photo 22053).

Trichomanes pennatum Kaulf. Enum. Fil. 264. 1824. TYPE: French Guiana, collector unknown
(LZ destroyed).

Neurophyllum hedwigii v. d. Bosch, Nederl. Kruid. Arch. 4:348. 1858, nom. superfl. TYPE: A
renaming of T. pinnatum Hedw., and so based on the type of that name.

Neuromanes kaulfussii v. d. Bosch, Nederl. Kruid. Arch. 4:348. 1858, nom. superfl. TYPE: A
renaming of T pennatum Kaulf, and so based on the type of that name.

Neuromanes immersum v. d. Bosch, Nederl. Kruid. Arch. 4:349. 1858. TYPE: Sto. Domingo, de
Tussac (P or RB not seen; isotype L not seen).

Trichomanes schomburgkianum Sturm in Mart. FI. Bras. 1(2):249. 1859. SYNTYPES: Near
Osembo, Surinam, Kappler 1751 (BR not seen); Berbice, Guyana, Rich. Schomburgk 299 (BR not
seen; isosyntype K not seen photo 19088); Essequibo River and the Canuku Mountains, Guyana,
Rich. Schomburgk 255 p. p. (BR not seen), 1135 [not 1155 as cited] (BR not seen photo 4818;
isosyntypes BM not seen, K not seen).

Neurophyllum hedwigianum Fee, Icon. Esp. Nouv. [M6m. Foug. 10]: Appendix. 1865, nom. superfl.
TYPE: A renaming of Neurophyllum pinnatum (Hedw.) K. Presl, and so based on the type of
Trichomanes pinnatum Hedw.

Neurophyllum thecaphyllum Fee, Hist. Foug. Antill. [Mem. Foug. 11]:104, t. 28, f. 1. 1866. TYPE:
Sto. Domingo, de Tussac (P or RB not seen).

Trichomanes pinnatum f. stipitatum Domin, Rozpr. Kr^. Ceske Spolecn. Nauk, Tf. Mat.-Pnr. 2
[Pterid. Dominica]:55. 1929. SYNTYPES: Windward Coast, Dominica, /. Jones 1922 (PRC not
seen); and near Laudat, Eggers 737 (PRC not seen). Although Domin cited Neurophyllum hedwigii in
synonymy, this form is not based on that name, which is illegitimate and without nomenclatural
standing.

Trichomanes pinnatum var. kaulfussii Domin, Rozpr. Krai. Ceske Spolecn. Nauk, Tf. Mat.-Pfir. 2
[Pterid. Dominica] :55. 1929. TYPE: Based on T. pennatum Kaulf., and not on Neuromanes

kaulfussii v. d. Bosch, nom. superfl., the other name cited by Domin.

TYPE: “Jamaica,” ex Herb. Banks (G? not seen); or, if the specimen is not
extant, t. 5, fig. 1 of Hedwig’s “Filicum Genera et Species” will serve as a
lectotype. Proctor in Howard (FL Less. Antill. 89. 1977) noted that T. pinnatum
has not been collected in Jamaica, except supposedly for the type material. It is
likely that the type material is from elsewhere in the Antilles.

Plants terrestrial, at 0-700(1200) m elevation, in forests, from near S. Ramon,
Sarapiqui (Pcia. Heredia), the valley of the Rfo General, throughout lowland
Panama, and the Choco. Also from Puerto Rico, the Lesser Antilles, Mexico to
Nicaragua, and tropical South America.

This wide-ranging and usually common species is infrequently found in Costa
Rica, except for the Peninsula de Osa. It is common in the Choco and the wetter
lowlands of Panama.

HYMENOPHYLLACEAE: 43. TRICHOMANES 217

300a. Trichomanes plumosum Kunze, Linnaea 9:104. 1834.

Trichomanes undulatum v.d. Bosch, Nederl. Kruid. Arch. 5(2):147. 1861, non Swartz, 1788, nom.
illeg. SYNTYPES: Near Sangari, Peru, Lechler 2548 (L not seen; isosyntypes BR not seen photo
4800, K not seen photo 19066); near Tatanara, Peru, Lechler 2505 and 2571 (both L neither seen).

TYPE: Pampayacu, Depto. Huanuco, Peru, Poeppig Diar. 1107 (LZ destroyed;
isotypes BR not seen photo 4819, K not seen photo 19070, U not seen Kramer
photo).

Plants probably epiphytic, at 1300-1400 m elevation, in the Flora area known
only from Cerro Pirre {Goldman 1889, US). Also from Venezuela to Peru.

301. Trichomanes polypodioides L. Sp. PI. 2:1098. 1753.

Trichomanes sinuosum Rich, ex Willd. Sp. PI. ed. 4, 5:502. 1810. TYPE: Guadeloupe, comm.
Richard (B-Hb. Willd. 20191 not seen Tryon photo).

Trichomanes quercifolium Desv. Ges. Naturf. Freunde Berlin Mag. 5:328. 1811. SYNTYPES:
Cayenne, St. Thomas, Puerto Rico, collectors unknown (all P-Hb. Desv. none seen). Synonymized by
Weatherby (Contr. Gray Herb. 114:35. 1936).

Trichomanes incisum Kaulf. Enum. Fil. 261. 1824, non Thunb., 1800, nom. illeg. TYPE: Brazil,
Chamisso (B? or LE? not seen).

Trichomanes cognatum K. Presl, Hymenophyllaceae 41 (postpr. 133). 1843. TYPE: Serra de
Estrella, Est. Rio de Janeiro, Brazil, Beyrich (PRC not seen).

Trichomanes serricula F6e, Crypt. Vase. Br6sil 1:189, t. 68, f. 3. 1869. TYPE: Barra [Manaus], Est.
Amazonas, Brazil, collector unknown (P or RB not seen).

Trichomanes ujhelyii Kiimmerle, Ann. Mus. Nat. Hung. 10:540. 1912. TYPE: Tagua, Sierra S.
Lorenzo, Colombia, 2200 m, Ujhelyi (BP not seen).

NEOTYPE: Montserrat, Proctor 19068 (A not seen; isoneotype US), designated
by Proctor (FI. Less. Antill. 2:93. 1977).

Plants epiphytic on trunks, often of tree ferns, at (0)300-1700 m elevation, in
forests, from the Cordillera de Tilaran, the Cordillera Central, the Cordillera de
Talamanca to Pcia. Chiriqui, the valley of the Rio General, the Peninsula de Osa,
the Atlantic coastal plain of Costa Rica and Panama, the Canal Zone, Cerro
Campana (Pcia. Panama), the El Llano - Carti-Tupile road (Com. S. Bias),
southwestern Pcia. Darien, and the northern Choco. Also from the Antilles,
Trinidad, Mexico to Honduras, and throughout tropical South America.

302. Trichomanes tuerckheimii Christ, Hedwigia 44:361. 1905.

Trichomanes torotumani Vareschi, Acta Bot. Venez. 1:88, t. 3B. 1966. TYPE: Rio Torotumani,
Terr. Amazonas, Venezuela, Vareschi & Maegdefrau 6663 (VEN not seen).

SYNTYPES: Leticia, Depto. Loreto, Peru, Ule 6228 (P not seen; isosyntype K
not seen Kramer photo); and Cubiliquitz, Depto. Alta Verapaz, Guatemala, 350
m, von Tuerckheim 8348 (P not seen; isosyntypes K not seen lO’amer photo, US).

Plants hemiepiphytic on trunks, at 0 - 900 m elevation, in forests, from the valley
of the Rio Reventazon, the valley of the Rio General, the Peninsula de Osa and
vicinity, Isla de Colon (Pcia. Bocas del Toro), Sta. Rita ridge (Pcia. Colon), the
trail from El Llano to Carti-Tupile (Com. S. Bias), and the Rio Atrato basin. Also
from Mexico, Guatemala, Belize, Colombia to Peru, and Surinam.

This species and the closely related but much rarer T. ankersii are hemiepiphytic
on smooth-barked tree trunks. The rhizomes and appressed fronds apparently are
able to absorb water that flows down the trunk when it rains.

218 HYMENOPHYLLACEAE: 43. TRICHOMANES

Jib I. punctatum

313 T. ovale subsp. sphenoides 317 T. pusillum 319 T. rhipidophyllum 318 T. reptans

HYMENOPHYLLACEAE: 43. TRICHOMANES 219

TRICHOMANES subg. DIDYMOGLOSSUM (Desv.) C. Chr.

303. Trichomanes angustifrons (F^e) W. Boer in Kramer, FI. Netherlands Antill.
l(Pterid.):17. 1962.

Didymoglossum angustifrons Fee, Hist. Foug. Antill. [Mem. Foug. 11]:113, t. 28, f. 5. 1866. TYPE:
Guadeloupe, L’Herminier (P or RB not seen).

Trichomanes setiferum Baker ex Jenm. J. Bot. Brit. For. 19:52. 1881. TYPE: Jamaica, Jenman 14
(K not seen).

Trichomanes mosenii Lindman, Ark. Bot. 1:46, f. 25D, E, 27. 1903. TYPE: Sorocaba, near Santos,
Est. S. Paulo, Brazil, Mosen 3810 (S not seen; isotype NY).

Plants epiphytic on trunks or epipetric, at 0-200(600) m elevation, in forests,
from the Atlantic lowlands of Costa Rica, the lower valley of the Rio Reventazon,
and the Canal Zone. Also from the Antilles, Trinidad, the Netherlands Antilles,
Mexico, Venezuela, Peru, the Guianas, Brazil, and Paraguay.

304. Trichomanes curtii Rosenst. Repert. Spec. Nov. Regni Veg. 22:5. 1925.

TYPE: Finca Hundriesser, Pcia. Limon, Erode (Sc Erode 398o {Ros. Fil Costor.
Exs. 222) (S not seen; isotype UC).

Plants epiphytic on trunks, at 0-300(1100) m elevation, in forests, from the
Atlantic lowlands of Costa Rica and Panama, the vicinity of S. Isidro del General,
the Peninsula de Osa, the Canal Zone, Sta. Rita ridge (Pcia. Colon), near Cana,
and the Choco. Also from Guatemala, Belize, Nicaragua, and Colombia.

305. Trichomanes ekmanii W. Boer, Acta Bot. Neerl. 11:319, f. 33. 1962.

TYPE: La Cumbre, Cordillera Central, Dominican Republic, Ekmon H 14342
(U; isotypes F, G not seen, GEE, MO, S not seen, UC, US).

Plants epiphytic on trunks and lianas, at 0-200 m elevation, in forests, from
Finca La Selva (Pcia. Heredia), the Peninsula de Osa, the Canal Zone, near Cana,
and west of Acandi and at Cabo Corrientes (both Depto. Choco). Also from
Cuba, Hispaniola, Belize, Honduras, Venezuela, Colombia, Peru, Bolivia,
Surinam, and Brazil.

306. Trichomanes godmanii Hook, in Baker, J. Linn. Soc., Bot. 9:337. 1866.

LECTOTYPE: Guatemala, Solvin (Sc Godmon in 1862 (K not seen fragm US),
chosen by W. Boer (Acta Bot. Neerl. 11:319. 1962).

Plants epiphytic on trunks, at 0-200 m elevation, in forests, from the Atlantic
coastal plain of Costa Rica, the Peninsula de Osa, and the Canal Zone. Also from
Cuba, Mexico to Honduras, and Nicaragua.

307. Trichomanes gourlianum Grev. in J. Smith in Seemann, Bot. Voy. Herald
240. 1854.

Trichomanes lehmannii Hieron. Bot. Jahrb. Syst. 34:420. 1904. TYPE: Rio Timbiqm, Depto.
Cauca, Colombia, Lehmann 8918 (B not seen; isotype US).

FIGS. 308 - 319. Trichomanes. FIG. 308. Plant of T. hymenoides, Lellinger 1107. FIG. 309. Plant of
T. kapplerianum, Mickel 3462. FIG. 310. Plant of T. krausii, Mickel 3510. FIG. 311. Plant of T.
membranaceum, Mickel 3536. FIG. 312. Plant of T. nummularium, Nee 6913. FIG. 313. Plant of T.
ovale, Standley 55777, Honduras. FIG. 314. Plant of T. petersii, Taylor, South Carolina. FIG. 316.
Frond of T. punctatum subsp. sphenoides, Pittier 2375. FIG. 317. Plant of T. pusillum, Fendler 24,
Venezuela. FIG. 318. Plant of T. replans, Skutch 3268. FIG. 319. Frond of T rhipidophyllum. Cook
& Doyle 236.

220 HYMENOPHYLLACEAE: 43. TRICHOMANES

TYPE: Pda. Darien, Gourlie (not seen). According to W. Boer (Acta Bot.
Neerl. 11:316. 1962), the type is not present in the Greville Herbarium in

Edinburgh.

Plants epiphytic on trunks, at 0-300(1300) m elevation, in forests, from the
Peninsula de Osa, the El Llano - Carti road (Pda. Panama), and the Choco. Also
from other localities in Colombia and from Ecuador.

308. Trichomanes hymenoides Hedw. Fil. Gen. Sp. t. 6, f. 3. 1799.

Trichomanes muscoides Swartz, J. Bot. (Schrader) 1800(2):95. 1801. TYPE: Jamaica, Swartz (S
not seen; isotype SBT not seen). Morton (Contr. U. S. Natl. Herb. 38:77. 1967) considered tlds
species to be a renaming of T. hymenoides Hedw., and so based on the type of that name.

Trichomanes apodum Hook. & Grev. Icon. Fil. l:t. 117. 1829. TYPE: Barbados, Parker (K not
seen).

Trichomanes pabstianum K. Mueller, Bot. Zeitung (Berlin) 12:738. 1854. TYPE: Ilha de Sta.
Catarina, Est. Sta. Catarina, Brazil, Pahst 267 (B not seen; isotype fragm US).

Didymoglossum sociale Fee, Crypt. Vase. Br6sil 2:85, t. 85, f. 3. 1873. TYPE: Tijuca, Est. Rio de
Janeiro, Brazil, Glaziou 5242 [not 5249, as cited] (P or RB not seen; isotype US).

Trichomanes orbiculare Christ, Bot. Jahrb. Syst. 19, Beibl. 47: 26. 1894. TYPE: Blumenau, Est.
Sta. Catarina, Brazil, Moeller (not seen).

Trichomanes fraseri Jenm. Gard. Chron. Ill, 20:266. 1896. TYPE: Grenada, Fraser (K not seen).
Trichomanes hymenoides i. pseudoreptans Rosenst. Hedwigia 46:75. 1906. TYPE: Serra de Joao
Rodriguez, Est. Rio Grande do Sul, Brazil, Juergens & Slier 140 (S not seen photo 6152; isotypes NY,
UC).

TYPE: Not stated, but according to Morton (Contr. U. S. Natl. Herb. 38:77.
1967), presumably Jamaica, Swartz (G not seen; isotypes B-Hb. Willd. 20188 not
seen Tryon photo, FI not seen photo 16574, L not seen photo 2416). Wessels Boer
(Acta Bot. Neerl. 11:306. 1962) considered Hedwig’s plate to be the type.

Plants epiphytic on trunks and epipetric, at 500-1500(2100) m elevation, in
forests, from the Cordillera de Tilaran, the Cordillera Central, Cerro Tablazo,
Cerro Carpintera, the Fila Costena near S. Vito, the Cordillera de Talamanca to
Pcia. Chiriquf, Cerro Campana (Pcia. Panama), and Cerro Pirre. Also from Cuba,
Jamaica, Hispaniola, the Lesser Antilles, Mexico, Guatemala, Belize, Trindad,
Venezuela, Colombia, Peru, Bolivia, Brazil, Paraguay, Uruguay, and Argentina.

309. Trichomanes kapplerianum Sturm in Mart. FI. Bras. 1(2):276. 1859.

Didymoglossum cordifolium Fee, Hist. Foug. Antill. [Mem. Foug. 11]: 113, t. 28, f. 4. 1866. TYPE:
Martinique, Rivoire (P or RB not seen).

Trichomanes muscoides var. minor Jenm. Bull. Bot. Dept. Jamaica 20: 8. 1890. TYPE: Not
stated. Authentic material may be in the Jenman Herbarium (NY).

Didymoglossum palmarum Vareschi, Acta Bot. Venez. 1:86. 1966. TYPE: Near La Fria, Edo.
Tachira, Venezuela, Medina 1108 (VEN).

TYPE: Surinam River near Station Victoria, Surinam, Kappler 1760 (W not
seen).

Plants terrestrial on trunks and epipetric, at 0-1500 m elevation, in forests,
from Finca Hundriesser (Pcia. Limon), Virgen del Socorro (Pcia. Heredia),
behind the Instituto Interamericano de Ciencias Agrfcolas near Turrialba (Pcia.
Cartago), Barro Colorado Island (Canal Zone), Cerro Gaital Caracoral (Pcia.
Code), Bahfa Solano, and Lx)ma del Cuchillo (Depto. Choco). Also from Puerto
Rico, the Ixsser Antilles, Trinidad, Colombia, Peru, the Guianas, and Brazil.

HYMENOPHYLLACEAE: 43. TRICHOMANES 221

310. Trichomanes krausii Hook. & Grev. Icon. Fil. 2:t. 149. 1830.

Didymoglossum fructuosum F6e, Hist. Foug. Antill. [Mem. Foug. 11]:112, t. 28, f. 3. 1866. TYPE:
Guadeloupe, L’Herminier in 1861 (P or RB not seen).

Trichomanes acrocarpon Fee, Crypt. Vase. Bresil 1:188, t. 70, f. 1. 1869. TYPE: Rio Grande do
Ajiro, Est. Rio de Janeiro?, Brazil, Glaziou 2248 (RB? not seen; isotype P not seen).

TYPE: Dominica, Kraus (E not seen).

Plants epipetric or epiphytic on trunks and branches, at 0-900(1200) m
elevation, in forests, from the Cordillera de Tilaran, the Cordillera Central, the
Cordillera de Talamanca to Pcia. Chiriqui, the Atlantic lowlands of Costa Rica
and Panama, the valley of the Rio General, the Canal Zone, Cerro Campana and
Juan Diaz (Pcia. Panama), and the vicinity of Cana. Also throughout tropical
America.

311. Trichomanes membranaceum L. Sp. PI. 2:1097. 1753.

TYPE: “America,” collector unknown (LINN 1253.1 not seen Maxon photo).
Plants epiphytic or epipetric, at 0-900(1200) m elevation, in forests, from the
Cordillera de Tilaran, the Atlantic coastal plain of Costa Rica and Panama, the
valley of the Rio General, the Peninsula de Osa, near the Fortuna reservoir site
(Pcia. Chiriqui), near Sta. Fe, the Canal Zone, around Porto Bello and Sta. Rita
ridge (both Pcia. Colon), southeast of Puerto Obaldia (Com. S. Bias), and the
northern half of the Choc6. Also from the Antilles, Mexico, Guatemala, Belize,
Nicaragua, Trinidad and Tobago, Venezuela, Colombia to Bolivia, and Surinam.

312. Trichomanes nummularium (v. d. Bosch) C. Chr. Ind. Fil. 645. 1906.

Didymoglossum nummularium v. d. Bosch, Nederl. Kruid. Arch. 5(3): 135. 1863. TYPE: Barra
[Manaus], along the Rio Negro, Est. Amazonas, Brazil, Spruce (L not seen).

Trichomanes goebelianum Giesenh. Flora 76:179, f. 3A, B. 1892. TYPE: Paraiso, near S.
Esteb^, Edo. Carabobo, Venezuela, Goebel in 1890 (B not seen).

Plants epiphytic, at 0-200 m elevation, in forests, from near Siquirres (Pcia.
Limon), near Piedras Blancas (Pcia. Puntarenas), the Peninsula de Osa, Isla de
Bastimentos (Pcia. Bocas del Toro), and the Rio Masambi Grande near Summit
Garden (Canal Zone). Also from Venezuela and Brazil.

313. Trichomanes ovale (Fourn.) W. Boer, Acta Bot. Neerl. 11:296. 1962.

Didymoglossum ovale Fourn. Bull. Soc. Bot. France 19:240. 1872. TYPE: Tovar, Edo. Aragua,
Venezuela, Fendler25 (P not seen; isotypes B not seen, G not seen, GH not seen, MO).

Trichomanes solitarium Jenm. Gard. Chron. Ill, 16:592. 1894. TYPE: Ugly River, St. Mary
Parish, Jamaica, Syme (K not seen; isotype IJ not seen).

Trichomanes fontanum Lindm. Ark. Bot. 1:44, t. 24D, F, 25C, 26. 1903. TYPE: Sto. Angelo, Est.
Rio Grande do Sul, Brazil, Lindman A1043 (S not seen; isotype US).

Trichomanes sphenoides var. minor Rosenst. Hedwigia 46:75. 1906. TYPE: Municipio Rio Pardo,
Est. Rio Grande do Sul, Brazil, & Stier207 (S not seen).

Plants epiphytic or epipetric, at 0-100 m elevation, in forests, from the
Peninsula de Osa, Laguna de Chiriqui and vicinity (Pcia. Bocas del Toro), the
Canal Zone, and Punta Guayabo Chiquita (Pcia. Darien). Also from the Greater
Antilles, Mexico to Honduras, Venezuela, Colombia, Surinam, and Brazil.

314. Trichomanes petersii A Gray, Amer. J. Sci. Arts II, 15:326. 1853.

Trichomanes schajfneri Schlechtend. Linnaea 26:368. 1854. TYPE: Huatusco, Edo. Veracruz,
Mexico, Schaffher (B not seen).

222 HYMENOPHYLLACEAE: 43. TRICHOMANES

TYPE: Near the Sipsey River, Winston County, Alabama, 8 Jan 1853, Peters
(GH not seen; presumable isotype US).

Plants epiphytic, at 1700-1800 m elevation, in the Flora area known only from
Cerro Bure, Pcia. Heredia (Gomez PtC-23867, CR). Also from the southeastern
United States, Mexico, and Guatemala.

315. Trichomanes punctatum subsp. labiatum (Jenm.) W. Boer, Acta Bot. Neerl.
11:302. 1962.

Trichomanes labiatum Jenm. Card. Chron. II, 24:7. 1885. TYPE: Bartica Grove, Essequibo
River, Guyana, Jenman 2109 (K not seen).

Trichomanes fruticulosum Jenm. Gard. Chron. Ill, 15:71. 1894. TYPE: Kykoverall Island, mouth
of the Mazaruni River, Guyana, Jenman 2351 (K not seen).

Trichomanes myrioneuron Lindm. Ark. Bot. 1:48, t. 25F-I, 28A, B, 29. 1903. TYPE: French
Guiana, Sagot 847 p. p. (S not seen).

Plants terrestrial on trunks and epipetric, at 0-600 m elevation, in forests, from
between Pto. Jimenez and the Rio Nuevo (Pcia. Guanacaste), the valley of the Rio
Reventazon southeast of Turrialba, and Finca La Selva (Pcia. Heredia). Also
from Venezuela, the Guianas, and Brazil.

316. Trichomanes punctatum subsp. sphenoides (Kunze) W. Boer, Acta Bot.
Neerl. 11:301. 1962.

Trichomanes sphenoides Kunze, Linnaea 9:102. 1834. TYPE: Cuchero, near Pampayacu, Depto.
Huanuco, Peru, Poeppig in 1829 (W not seen; isotype MO).

Plants epiphytic on trunks and epipetric, at 0-500 m elevation, in forests, from
Peralta (Pcia. Cartago), the vicinity of Guapiles (Pcia. Limon), Laguna de Chiriqui
and vicinity (Pcia. Bocas del Toro), the Canal Zone, Tumba Vieja (Pcia. Colon),
S. Jose Island (Pcia. Panama), between Paya and Pucro (Pcia. Darien), and the
region between the Rio Atrato and Rio S. Juan basins. Also from the Greater
Antilles, Guatemala, Honduras, and Venezuela to Bolivia.

317. Trichomanes pusillum Swartz, Nov. Gen. Sp. PI. Prodr. 136. 1788.

TYPE: Jamaica, Swartz (S not seen; isotypes B not seen Tryon photo, BM not
seen photo 6594, SBT not seen).

Plants epiphytic on trunks, at 1000-1500 m elevation, in forests, from north of
S. Ramon, above Platanillo (Pcia. Cartago), and between Cartago and S. Isidro del
General (Pcia. Cartago or S. Jose). Also from Jamaica, Hispaniola, Puerto Rico,
Venezuela, and Brazil.

318. Trichomanes reptans Swartz, Nov. Gen. Sp. PI. Prodr. 136. 1788.

Trichomanes quercifolium Hook. & Grev, Icon. Fil. l:t. 115. 1829. TYPE: Esmeraldas, Pcia.
Esmeraldas, Ecuador, 8000 Jameson (E not seen).

Trichomanes montanum Hook. Icon. PI. 2:t. 187. 1837. TYPE: Near La Sierra on the road to
Esmeraldas, Pcia. Esmeraldas, Ecuador, 8500 ft, Jameson (K not seen).

Trichomanes pusillum var. macropus Christ in Schwacke, PI. Nov. Mineir. 2:15. 1900.

LECTOTYPE: Sta. Rosa du Copey, Pcia. S. Jose, Tonduz 12250 (G? not seen; isolectotypes BR not
seen photo 4815, US), inferentially chosen by Christ (Prim. FI. Costar. 3(1):5. 1901) when he

described the variety as new a second time.

TYPE: Jamaica, Swartz (S not seen; isotype SBT not seen).

Plants epiphytic on roots and trunks and epipetric, at (100)1000-1800 m
elevation, in forests, from S. Pedro de S. Ramon (Pcia. Alajuela), the Cordillera

HYMENOPHYLLACEAE: 43. TRICHOMANES 223

Central, the Cordillera de Talamanca to Pcia. Chiriqui, above Sta. Fe, Sta. Rita
ridge (Pcia. Col6n), and Cerro Pirre. Also from Jamaica, Hispaniola, Mexico,
Guatemala, Honduras, Nicaragua, Venezuela to Peru, Brazil, and Argentina.

319. Trichomanes rhipidophyllum Slosson, Bull. Torrey Bot. Club 40:687, t. 26, f.
1-3. 1913.

TYPE: Near Onaca, Sta. Marta, Depto. Magdalena, Colombia, H. H. Smith
2445 (NY; isotype US).

Plants epiphytic on trunks, at 800-1000 m elevation, in forests, from Juan Vihas
(Pcia. Cartago) and Villa Colon (Pcia. S. Jose). Also from Colombia.

TRICHOMANES subg. PACHYCHAETUM K. PresI

320. Trichomanes elegans Rich. Actes Soc. Hist. Nat. Paris 1:114. 1792.

Trichomanes millefolium Desv. M6m. Soc. Linn. Paris 6:329. 1827. TYPE: Brazil, collector
unknown (P not seen Cintract photo).

Trichomanes prieurii Kunze, Analecta Pteridogr. 48. 1837. SYNTYPES: French Guiana, Leprieur
(LZ destroyed); Rio Japura, Est. Amazonas, Brazil, Martius (M not seen); and Guyana, Rudge (K
not seen).

Trichomanes lastreoides K. Presl, Hymenophyllaceae 15 (postpr. 107). 1843. TYPE: Brazil, Sellow
(PRC not seen).

Trichomanes anceps Hook. Sp. Fil. 1:135, t. 40C1. 1846, var. anceps. SYNTYPES: Hooker cited
several syntypes, including two from the Old World that should not be chosen lectotype.

Trichomanes weddellii v. d. Bosch, Nederl. Kruid. Arch. 5(2):174. 1861. SYNTYPES: Bolivia,
Weddell (L? not seen); Guyana, Schomburgk (K or L? not seen); and Brazil, Sellow (B not seen).

Trichomanes opacum v. d. Bosch, Nederl. Kruid. Arch. 5(2):175. 1861. TYPE: S. Gavan, Depto.
Puno, Peru, Lechler 2175 p. p. (L? not seen).

Trichomanes leprieurii Hook. Gard. Ferns t. 11. 1862, nom. superfl. TYPE: A renaming of T
prieurii Kunze, and so based on the type of that name.

TYPE: French Guiana, Le Blond (P not seen).

Plants terrestrial, at 0-900(1700) m elevation, in forests, from the Atlantic
lowlands and adjacent foothills of Costa Rica and Panama, Cana, and the Choco.
Also from the Lesser Antilles, Nicaragua, and tropical South America.

321. Trichomanes rigidum Swartz, Nov. Gen. Sp. PL Prodr. 137. 1788.

Trichomanes bifidum Vent, ex Willd. Sp. PI. ed. 4, 5:511. 1810. TYPE LOCALITY: West Indies.
Trichomanes compressum Desv. Ges. Naturf. Freunde Berlin Mag. 5:329. 1811. TYPE

LOCALITY: Hispaniola, according to Weatherby (Contr. Gray Herb. 114:35. 1936).

Trichomanes mandioccanum Raddi, PI. Bras. Nov. Gen. 1:64, t. 79, f. 2. 1825. TYPE: Serra da
Estrella, Est. Rio de Janeiro, Brazil, Raddi (FI not seen).

Trichomanes firmulum K. Presl, Hymenophyllaceae 46 (postpr. 138). 1843. TYPE: Serra da
Estrella, Est. Rio de Janeiro, Brazil, Beyrich (PRC not seen; isotype L not seen photo 2455).

Trichomanes daucoides K. Presl, Abh. Konigl. Bohm. Ges. Wiss. V, 6:372 (repr. 12), t. 7. 1851.
SYNTYPES: Isla de Ometepe, Nicaragua, Friedrichsthal (PRC not seen); and Sto. Tomas,
Guatemala, Friedrichsthal (PRC not seen).

Trichomanes krugii Christ ex Krug in Urban, Bot. Jahrb. Syst. 24:90. 1897. SYNTYPES: Eastern
Cuba, Wright 903 (K not seen; isosyntype US); Martinique, Duss 1531 (not seen); St. Vincent, Smith
485 (K not seen); Grenada, Eggers 6145b, 6175b (neither seen); and Grenada, Sherring 129 (K not
seen; isosyntype US).

TYPE: Jamaica, Swartz (S not seen photo 6184; isotype B-Hb. Willd. 20202-1
not seen Tryon photo).

224 HYMENOPHYLLACEAE: 43. TRICHOMANES

328 T. radicans

HYMENOPHYLLACEAE: 43. TRICHOMANES 225

Plants terrestrial, at 500-1700(2500) m elevation, in forests, from Lx)s Angeles
de S. Ramon (Pda. Alajuela), Volcdn Pods, the Cordillera de Talamanca to Pda.
Chiriqui and its foothills, the mountains of Panama, and the Serrama de Baudo.
Also from the Antilles, Trinidad, Mexico to Nicaragua, and tropical South
America.

322. Trichomanes sprucei Baker, Syn. Fil. 87. 1867.

Trichomanes subdeltoideum C. Chr. Ind. Fil. 650. 1906, nom. superfl. TYPE: A renaming of T.
sprucei Baker, non T. spruceanum Hook., 1854, and so based on the type of T. sprucei.

SYNTYPES: Brazil, Spruce 1399 (K not seen); and Panure, Est. Amazonas,
Brazil, Spruce 2838 (K not seen; isosyntype US).

Plants terrestrial, at 0-100 m elevation, in forests, in the Flora area known only
from Los Pedregales, below Cerro Tife, Pda. Code, 500 m {Knapp & Dressier
3795, MO) and from La Concepcion, 15 km east of Quibdo {Archer 2000, US).
Also from other localities in Colombia, Venezuela, Peru, and Brazil.

TRICHOMANES subg. TRICHOMANES

322a. Trichomanes angustatum Carm. Trans. Linn. Soc. London 12:513. 1818.

Trichomanes tenerum Spreng. Syst. Veg. ed. 16, 4:129. 1827. TYPE: Brazil, collector unknown
(LZ destroyed).

Trichomanes fulvum Klotzsch in Sturm in Mart. FI. Bras. 1(2):272, t. 18, f. 6. 1859. TYPE: Brazil,
Sello (W not seen; isotype K not seen photo 19095).

TYPE: Tristan da Cunha, Cunningham (K not seen; isotype BM not seen).

Plants epiphytic, at ca. 2000 m elevation, in forests, in the Flora area known only
from the southern slope of Volcan Barba, Pda. Heredia {Montgomery & Huttleston
85-94, US). Also from Cuba, Jamaica, Hispaniola, Mexico to Honduras, and
Colombia to Argentina and Brazil.

323. Trichomanes capillaceum L. Sp. PI. 2:1099. 1753.

Trichomanes tenellum Hedw. Fil. Gen. Sp. ad t. 6, f. 1. 1799. TYPE: Jamaica, Swartz (LZ

destroyed?; isotype FI not seen photo 16562).

Trichomanes trichoideum Swartz, J. Bot. (Schrader) 1800(2) :98. 1801, nom. superfl. TYPE: A
renaming of T. tenellum Hedw., and so based on the type of that name. Swartz cited T pusillum in
publishing T. trichoideum, but when later (Syn. Fil. 144. 1806) he renamed T. trichoideum as T
trichodes, he did cite T. tenellum.

Trichomanes angustissimum K. Presl, Abh. Konigl. Bohm. Ges. Wiss. V, 6:378 (repr. 18), t. 8A.
1851. SYNTYPES: Jamaica, Swartz (B-Hb. Willd. 20200 not seen microfiche S. I. Library); and
Colonia Tovar, Edo. Aragua, Venezuela, Moritz 151 (PRC? not seen).

Trichomanes schiedeanum K. Muell. Bot. Zeitung (Berlin) 12(41) :716. 1854. TYPE: Mexico,
Schiede (B not seen).

Trichomanes capillaceum var. subclavatum Christ in Pitt. Prim. H. Costaric. 3(1):4. 1901. TYPE:
Tuis, Pcia. Cartago, 650 m, Tonduz 11308 (P not seen; isotypes CR, US).

FIGS. 320 - 329. Trichomanes. FIG. 320. Median pinna of T. elegans, Cuatrecasas 16547. FIG. 321.
Median pinna of T. rigdum, Williams 907. FIG. 322. Median pinna of T. spmcei, Lellinger & de la
Sota 463. FIG. 323. Median pinna of T. capillaceum, Jimenez M. 1458. FIG. 324. Distal median
pinna of T. collariatum, Wilbur & Stone 9735. FIG. 325. Median pinna of T. diaphanum, Killip 5210.
FIG. 326. Median pinna of T. hymenophylloides, Lellinger 784. FIG. 327. Median pinna of T.
pyxidifemm, Skutch 2315. FIG. 328. Median pinna of T. radicans, Lellinger 1101. FIG. 329. Median
pinnule of T. mpestre, Skutch 2529.

226 HYMENOPHYLLACEAE: 43. TRICHOMANES

Trichomanes hypnoides Christ, Bull. Herb. Boissier II, 5:725. 1905. TYPE: Zonehuitz, Edo.
Chiapas, Mexico, Muench 108 (P? not seen; isotypes DS, US).

TYPE: Plate 99D of Plumiefs “Traite...”, which is based on a specimen
collected by Plumier on Hispaniola.

Plants epiphytic, often on tree fern trunks, at 600-2000(2700) m elevation, in
forests, from the Cordillera de Tilardn, the Cordillera Central, the Cordillera de
Talamanca to Pcia. Chiriqui, and Cerro Tacarcuna. Also from the Greater
Antilles, Mexico to Nicaragua, and Venezuela to Peru.

The laminae of T. capillaceum var. subclavatum are not so completely
skeletonized as are those in the more extreme forms of this species, but the
variation seems continuous and taxonomically inconsequential.

324. Trichomanes collariatum v. d. Bosch, Nederl. Kruid. Arch. 4:368. 1859.

Trichomanes repens Schott ex Sturm in Mart. FI. Bras. 1(2):264. 1859, TYPE: Locality unknown,
Herb. Mus. Pal. Vindob. 4480 (W not seen photo BM not seen), synonymized by A. R. Smith (FI.
Chiapas 2:241. 1981).

Trichomanes repens var. stipitatum Sturm in Mart. FI. Bras. 1(2):264. 1859. TYPE: Near S.
Gabriel, Rio Negro, Est. Amazonas, Brazil, Spmce 2181 (W not seen; isotypes K not seen photo
19049, UC, US).

Trichomanes martinezii Rovirosa, Pteridogr. Sur Mexico 106, t. 74A, f. 1-3. 1910. TYPE: Rio
Lacanja, Edo. Chiapas, Mexico, L. Martinez (Hb. Rovirosa 1103 not seen).

TYPE: Teapa, Edo. Tabasco, Mexico, 300 ft. Linden 1508 (L not seen; isotypes
GH not seen, K not seen photo 19047).

Plants epiphytic on trunks, at 0-1500(2000) m elevation, in forests, from the
Cordillera de Tilardn, the Cordillera Central, the valley of the Rio Reventazon,
the Atlantic coastal plain of Costa Rica and adjacent Panama, near the Fortuna
Reservoir (Pcia. Chiriqui), near Sta. Fe, the Canal Zone and adjacent Panama,
southeast of Puerto Obaldia (Com. S. Bias), the Cana-Cuasi trail (Pcia. Darien),
and the slopes of Alto del Buey. Also from Mexico to Honduras, Nicaragua, and
Venezuela and Colombia to Peru and Brazil.

325. Trichomanes diaphanum H.B.K. Nov. Gen. Sp. 1:25 (fol. 16). 1816.

Trichomanes eximium Kunze ex Sturm in Mart. FI. Bras. 1(2):271. 1859, nom. superfl. TYPE: A
renaming of T. diaphanum H.B.K., which was cited in synonymy, and so based on the type of that
name.

Trichomanes debile v. d. Bosch, Nederl. Kruid. Arch. 5(2):154. 1861. TYPE: Carabobo, Edo.
Carabobo, Venezuela, Punch & Schlim 596 (L? not seen).

Trichomanes bradei Christ, Bull. Soc. Bot. Geneve II, 1:217. 1909. TYPE: La Palma, Pcia. S. Jose,
1400 m, 14 Mar 1908, Brade (P? not seen).

Trichomanes diaphanum var. subalatum Rosenst. Repert. Spec. Nov. Regni Veg. 7:291. 1909, as
“subalata.'' TYPE: Mt. Pichincha, Pcia. Pichincha, Ecuador, Spruce 5654 (S not seen).

Trichomanes eximium var. crispulum Rosenst. Repert. Spec. Nov. Regni Veg. 20:89. 1924.

LECTOTYPE: Organ Mountains, Est. Rio de Janeiro, Brazil, Luetzelburg 6196 (M not seen;
isolectotype US), chosen by Lellinger (Mem. New York Bot. Gard. 38:27. 1984).

TYPE: Venezuela, Humboldt & Bonpland (B not seen).

Plants epiphytic on trunks and branches, at (0)5(X)- 2300(2700) m elevation, in
forests, from the Cordillera Central, the Cordillera de Talamanca to Pcia.
Chiriqui, the Peninsula de Osa, the vicinity of Laguna de Chiriqui (Pcia. Chiriqui),
Cerro Gaital Caracoral and La Mesa (Pcia. Code), near Alto Piedra near Sta. Fe,

HYMENOPHYLLACEAE: 43. TRICHOMANES 227

Cerro Jefe, and Loma del Cuchillo (Depto. Choc6). Also from Guatemala,
Honduras to Nicaragua, Venezuela to Guyana, and Colombia to Peru and Brazil.

326. Trichomanes hymenophylloides v. d. Bosch, Nederl. Kniid. Arch. 5(3):209.

1863.

Trichomanes pyxidiferum Hook. & Grev. Icon. Fil. 2:t. 206. 1831, non L., 1753, nom. illeg. TYPE:
Based on St. Vincent, Guilding [194] (K not seen; isotype GH not seen; presumable isotype L not
seen photo 2451).

Trichomanes leptophyllum v. d. Bosch, Nederl. Kruid. Arch. 4:363. 1859, non A. Cunn. in Hook.,
1836, nom. illeg. TYPE: Based on T. pyxidifemm Hook. & Grev., and so based on the type of that
name.

TYPE: A renaming of T. leptophyllum v. d. Bosch, and so based on the type of
that name.

Plants epiphytic on trunks and branches, at 1400 - 2000 m elevation, in forests,
from Monteverde, Porrosati (Pcia. Heredia), Sta. Clara de Cartago (Pcia.
Cartago), and above Boquete. Also from the Antilles, Mexico, Guatemala,
Honduras, Trinidad, Venezuela to Ecuador, Guyana, Surinam, and Brazil.

327. Trichomanes pyxidiferum L. Sp. PL 2:1098. 1753.

Trichomanes brasiliense Desv. Mem. Soc. Linn. Paris 6:328, t. VII, f. 4. 1827. TYPE: Brazil,
collector unknown (P not seen).

Trichomanes olivaceum Kunze ex Klotzsch, Linnaea 20:437. 1847. TYPE: Merida, Edo. Merida,
Venezuela, Moritz 341 (B not seen; isotype BR not seen photo 4808).

Trichomanes emarpnatum K. Presl, Gefassbiindel Farm 24 (postpr. 332). 1847, non Poir., 1808,
nom. illeg. SYNTYPES: Serra da Estrella, Est. Rio de Janeiro, Brazil, Beyrich (PRC not seen;
isotype L not seen photo 2449), and Pohl (PRC not seen).

Trichomanes cavifolium K. Muell. Bot. Zeitung (Berlin) 12(43) :753. 1854. TYPE: Based on T.
pyxidiferum sensu Schlechtend. & Cham. (Linnaea 5:618. 1830), hence on the basis of that name,
which is the specimen Cuesta grande de Chiconquiaco, Mexico, Schiede (B not seen).

Trichomanes lechleri v. d. Bosch, Nederl. Kruid. Arch. 4:363. 1859. TYPE: Based on T.

pyxidifemm sensu Kunze (Linnaea 9:106. 1834), hence on the basis of that name, which is the
specimen Near Pampayacu, Depto. Huanuco, Peru, Poeppig 1105 (LZ destroyed).

Trichomanes tenellum Mart, ex v. d. Bosch, Nederl. Kruid. Arch. 5(2): 154. 1861, non Hedw., 1799,
nom. illeg. TYPE: Brazil, Martius 146 (L not seen photo 2452).

Trichomanes tranninense Fee, Crypt. Vase. Bresil 1:187, t. 69, f. 1. 1869. TYPE: Brazil, Glaziou
2251 (P or RB not seen).

Trichomanes junceum Christ, Bull. Herb. Boissier II, 4:944. 1904. TYPE: Costa Rica, Werckle (P
not seen; isotype US fragm NY).

Trichomanes pyxidifemm var. organense Rosenst. Repert. Spec. Nov. Regni Veg. 20:89. 1924.
SYNTYPES: Serra dos Orgaos, Est. Rio de Janeiro, Breizil, Luetzelhurg 93 (M not seen); and
Itarmaraty, Serra dos Orgaos, Est. Rio de Janeiro, Brazil, Luetzelhurg 92 (M not seen photo 6204;
isosyntype NY).

Trichomanes pyxidifemm f. g^acile Rosenst. Repert. Spec. Nov. Regni Veg. 20:90. 1924. TYPE:
Corcovado, Est. Rio de Jemeiro, Brazil, Luetzelhurg 262 (M not seen; isotype NY).

TYPE: Plate 50E of Plumier’s “Traite...”, which was based on a plant collected
by Plumier on Hispaniola.

Plants epiphytic on tree trunks and branches, rarely epipetric, at
400-1300(1800) m elevation, in forests, from near S. Ramon, the Cordillera
Central, Cerro Carpintera, the Cordillera de Talamanca to Pcia. Chiriquf, the
valley of the Rfo General, and the Fila Costeha near S. Vito. Also from the

228 HYMENOPHYLLACEAE: 43. TRICHOMANES

Greater Antilles, Mexico to Nicaragua, and Venezuela and Colombia to Guyana
and Brazil.

328. Trichomanes radicans Swartz, J. Bot. (Schrader) 1800(2) :97. 1801, var.

radicans.

Trichomanes kunzeanum Hook. Sp. Fil. 1:127, t. 39D. 1844. SYNTYPES: Near Pampayacu,
Depto. Huanuco, Peru, Poeppig 1132 (K not seen photo 19050; isotype LZ destroyed); Pangoa,
Depto. Junrn, Peru, Mathews 1088 (K not seen photo 19051); and Truxillo and Merida, Edo.
Merida, Venezuela, Linden 176 (K not seen; isosyntypes BR not seen, FI not seen photo 16586).

Trichomanes mexicanum v. d. Bosch, Nederl. Kruid. Arch. 5(2):164. 1861. SYNTYPES: Mexico,
Schiede 806 (B not seen fragm L not seen); and Mexico, Schaffher 7 (P or RB not seen fragm L;
probable isosyntype K not seen photo 19052).

Trichomanes latevirens Fee, Crypt. Vase. Bresil 1:191, t. 67, f. 2. 1869. TYPE: Brazil, Glaziou 1677
(P or RB not seen).

TYPE: Jamaica, Swartz (S not seen photo 6205).

Plants epiphytic on trunks, at (100)400-2800(3200) m elevation, in forests, from
the Cordillera de Tilaran, the Cordillera Central, Cerro Carpintera, the Cordillera
de Talamanca to Pcia. Chiriqui, Cerro Campana (Pcia. Panama), and Cana. Also
throughout tropical America.

Proctor (Amer. Fern J. 72:108. 1982) distinguished specimens with more
skeletonized fronds and larger, broadly winged involucres growing on calcareous
rocks in Jamaica as T. radicans var. antillarum (v. d. Bosch) Proctor.

329. Trichomanes rupestre (Raddi) v. d. Bosch, Nederl. Kruid. Arch. 4:370. 1859.

Hymenophyllum rupestre Raddi, PI. Bras. Nov. Gen. 1:67, t. 80. 1825. SYNTYPES: Gavia, Tijucca,
Est. Rio de Janeiro, Brazil, Raddi (FI not seen); and Corcovado, Est. Rio de Janeiro, Brazil, Raddi
(FI not seen).

Trichomanes venustum Desv. Mem. Soc. Linn. Paris 6:328. 1827. TYPE: Brazil, collector unknown
(P not seen).

Trichomanes luschnathianum K. Presl, Hymenophyllaceae 45 (postpr. 137). 1843. TYPE: Rio de
Janeiro, Est. Rio de Janeiro, Brazil, Luschnath (PRC not seen).

Trichomanes pellucidum Goldm. in Meyen, Nov. Actorum Acad. Caes. Leop.-Carol. Nat. Cur. 19,
Abh. 1:466. 1843. TYPE: Corcovado, Est. Rio de Janeiro, Brazil, Martiusl (not seen).

Trichomanes frondosum Fee, Crypt. Vase. Bresil 1:190, t. 68, f. 1, 2. 1869. SYNTYPES: Brazil,
Glaziou 917 (P or RB not seen), 1647 (P or RB not seen), 1715 (P or RB not seen; isosyntype K not
seen photo 19054), and 2254 (P or RB not seen).

Trichomanes venustum var. monomorphum Christ, Bull. Herb. Boissier II, 4:944. 1904. TYPE:
Costa Rica, Werckle (P not seen).

Plants epiphytic on trunks, at 100-1600 m elevation, in forests, from the
Cordillera Central, the Cordillera de Talamanca to Pcia. Chiriqui, the valley of the
Rio General, and the Fila Costeha near S. Vito. Also from Venezuela, Colombia,
Peru, Bolivia, and Brazil.

This species is rather variable; the more dissected forms resemble T. collariatum
or T. radicans, but the pinnae are more inequilateral in T. rupestre.

GLEICHENIACEAE: 44. GLEICHENIA 229

GLEICHENIACEAE

Rhizomes very long-creeping, dichotomously branched, mostly dark brown,
bearing pluricellular, lax or bristle-like hairs, narrow, mostly brownish, often
ciliate or setose scales, or both; fronds medium-sized to large, commonly
scrambling over other vegetation; stipes distant along the rhizomes, wiry; laminae
pinnately or usually pseudodichotomously divided, with at least the ultimate
segments pectinate, occasionally scaly, the pseudodichotomies bearing a hairy or
scaly bud, this sometimes breaking dormancy and producing a prolonged axis
bearing a pair of segments and a resting bud at its apex; penultimate and more
proximal axes naked, partially pectinate, or pectinate; laminae glabrous, glaucous,
or hairy abaxially; veins always free, simple or 1 -3-forked; sori round, surficial on
the abaxial surface of the laminae; indusia absent; sporangia a small (2-ca. 15)
and often constant number per sorus, short-stalked, pear-shaped; paraphyses
absent; spores more than 100 per sporangium, monolete or trilete.

1. Veins in the ultimate segments simple or 1-forked; sporangia usually 2-ca. 5 per sorus;
rhizomes and resting buds scaly.

44. Gleichenia

1. Veins in the ultimate segments 2- or 3-forked; sporangia usually 8-15 per sorus; rhizomes and
resting buds hairy.

45. Dicranopteris

44. GLEICHENIA J. E. Smith

Plants terrestrial, often in disturbed places like roadbanks and landslides;
rhizomes long-creeping, thin, occasionally forked, scaly, the scales lanceate or
lanceolate, commonly weakly bicolorous, pale to dark brown, ciliate to setose;
fronds mostly large, often sprawling over other vegetation; stipes long, terete or
nearly so, rigid, often glabrous; laminae debate to oblong, the rachis bearing
distant, opposite pairs of pinnae flanking a terminal bud that elongates to continue
the rachis or remains dormant at the frond apex, the bud and pinna bases
variously scaly; pinnae mostly 1-3 times pseudodichotomously divided (pinnate in
G. bancroftii) with a dormant bud at each division, at least the ultimate divisions
laminate, these laminae pectinate, linear to elliptic, glabrous adaxially, often hairy
or glaucous abaxially and with scales on the axes, mostly subcoriaceous or
coriaceous, often somewhat revolute at the margins; veins 1-forked; sori round,
exindusiate, composed of a rather constant, low (ca. 2-5) number of subsessile
sporangia, paraphyses absent.

Pantropical and subtropical, with a few species in temperate regions of the
Southern Hemisphere; ca. 130 species, including Diplopterygium and Sticherus.
MAXON, W. R. 1909. Gleicheniaceae. North Amer. FI. 16(l):53-63.

1. Primary branches bipinnate, not dichotomously forked. Plants large, with 1(3) pair(s) of pinnae
up to 1.5 m long, 50 cm wide.

330. G. bancroftii

1. Primary branches once to several times pseudodichotomously forked, not pinnately divided.. 2.

2(1). Bud scales linear-lanceolate to narrowly triangular, usually ciliate, whitish or pale reddish
brown to dark brown, indurate; laminae and axes usually scaly or pubescent..?.

2(1). Bud scales broadly triangular to roundish, usually eciliate, pale to dark brown, not indurate;
lammae and axes glabrous (except in G. pteridella).3.

3(2). Laminae and axes with broad, ciliate scales; primary branches (l)2(3)-forked; central frond
axes sometimes prolonged. Primary and secondary branches not pectinate.

342. G. pteridella

230 GLEICHENIACEAE: 44. GLEICHENIA

3(2). Laminae and axes glabrous; primary branches l(2)-forked; central frond axes not
prolonged..4.

4(3). Segments strongly reflexed, mostly ca. 8 mm distant. Plants up to 50 cm long; primary
branches not pectinate.

344. G. retroflexa

4(3). Segments not reflexed..5.

5(4). Axes shghtly flexuous, especially toward the apex of the pinnae. Plants up to ca. 1 m long;
central frond axes and pinna axes prolonged; primary branches pectinate; base of segments fully
adnate to the rachis, more surcurrent than decurrent.

340. G. orthoclada

5(4). Axes not flexuous.. 6.

6(5). All branches pectinate; segments approximate, fully adnate to the axis. Plants up to 2 m long;
central frond axes and pinna axes prolonged.

337. G. intermedia

6(5). Primary branches not pectinate; segments somewhat distant, the base hastulate, the
acroscopic base free and overlapping the axis. Plants probably less than 1 m long; central frond axes
prolonged.

336. G. hastulata

7(2). Branches with their segments up to 3.5 mm across, at a 30° angle to one another, 3(5) times
equally pseudodichotomously forked.

335. G. gnidioides

7(2). Branches with their segments more than 8 mm across..8.

8(7). Bud scales broadly to narrowly lanceolate, with long, lax, twisted, marginal cilia.. 14.

8(7). Bud scales narrowly lanceolate to linear, with short, stiff, straight, marginal cilia (cilia absent
in G. nitidula)..9.

9(8). Laminae densely tomentose on the abaxial surface, sometimes glabrescent. Rachises with
deciduous, atropurpureous, shiny, lanceolate scales.

332. G. brevipubis

9(8). Laminae glabrous or scaly on the abaxial surface, not tomentose.. 10.

10(9). Branches with their segments l-2.5(3.5) cm wide.. 12.

10(9). Branches with their segments (3)4-9 cm wide..ll.

11(10). Margins and abaxial surface of the laminae with stellate scales and stellate hairs. Plants
large; central frond axes prolonged; primary branches not pectinate; segments (1.5)2 -3 cm long,
usually less than their width distant.

333. G. compacta

11(10). Margins and abaxial surface of the laminae with lanceolate, deeply toothed or ciliate scales.
Plants large; central frond axes prolonged; primary branches not pectinate; segments 2-4 cm long,
more than their width distant.

343. G. remota

12(10). Laminae glabrous on the pinna axes. Plants ca. 0.5 m long; central frond axes elongate in
the largest specimens; primary rachises partially or not pectinate, rarely fully so; ultimate branches
with their segments ca. 10 mm wide, glaucous abaxially.

339. G. nitidula

12(10). Laminae scaly on the piima axes.. 13.

13(12). Veins 18-26 pairs per segment; segments linear, round at the apex; branches with their
segments (10)15-35 mm wide.

345. G. rubiginosa

13(12). Veins 9-15 pairs per segment; segments lanceate, obtuse to acute at the apex; branches
with their segments 10 - 25 mm wide. Plants 30(45) cm long.

346. G. strictissima

GLEICHENIACEAE: 44. GLEICHENIA 231

14(8). Scales of the buds and axes whitish or stramineous, sometimes with a reddish-brown,
indurate central area. Laminae glaucous on the abaxial surface; segment axes glabrous; primary
branches not pectinate; central frond axes not prolonged.

341. G. pallescens

14(8). Scales of the buds and axes reddish-brown, sometimes with a slightly darker, but not
indurate central area..l5.

15(14). Laminae not tomentose on the abaxial surface..!?.

15(14). Laminae densely tomentose on the abaxial surface, sometimes glabrescent..l6.

16(15). Segments 1.5 -2.5 mm wide, 10-18 mm long. Plants 0.5-2 m long.

331. G. bifida

16(15). Segments ca. 2.5 mm wide, 6 mm long. Plants ca. 0.5 m long.

338. G. maritima

17(15). Branches with their segments 1-2 cm wide; segments strongly revolute, coriaceous, the
veins obvious on the adaxial surface. Primary branches partially pectinate; central frond axes
repeatedly prolonged.

334. G. costaricensis

17(15). Branches with their segments (1)1.5 -3 cm wide; segments slightly revolute, chartaceous,
the veins almost hidden on the adaxial surface. Primary branches not or partially pectinate, or with
accessory pinnae; central frond axes repeatedly prolonged.

333. G. compacta

330. Gleichenia bancroftii Hook. Sp. Fil. 1:5, t. 4A. 1844.

Mertensia bancroftii var. vitellina Kunze, Linnaea 18:307. 1845. TYPE: Caracas, Distr. Fed.,
Venezuela, Moritz 11 (LZ destroyed; isotype B not seen).

Mertensia grandis Fee, Hist. Foug. Antill. [Mem. Foug. 11]: 120. 1866. SYNTYPES: Matouba,
Guadeloupe, UHerminier in 1861 (P not seen; isosyntypes IJ not seen, US); and Huatusco, Mirador,
Edo. Veracruz, Mexico, Schafiher 230 (RB not seen).

Gleichenia bancroftii var. ^acilis Jenm. Bull. Bot. Dept. Jamaica 5:276. 1898. TYPE: Jamaica,
1500-1800 m,Jenman (NY? not seen).

Gleichenia brunei Christ, Bull. Herb. Boissier II, 5:13. 1905. TYPE: El Desengano, Pda.

Heredia?, 1400 m, Brune 317 (P not seen).

LECTOTYPE: Jamaica, Bancroft (K not seen), chosen by Proctor (FI. Less.
Antill. 60. 1977).

Plants terrestrial, at ca. 1800 m elevation, in the Flora area known only from the
type of G. brunef one unlocalized collection from Costa Rica (Werckle in 1907,
US), and Vara Blanca, Pda. Heredia {Chrysler 4986, MO, US). Also from Cuba,
Jamaica, Hispaniola, the Lesser Antilles, Mexico to El Salvador, and Venezuela
and Colombia to Bolivia.

331. Gleichenia bifida (Willd.) Spreng. Syst. Veg. ed. 16, 4:27. 1827.

Mertensia bifida Willd. Kongl. Vetensk. Akad. Nya Handl. 25:168, t. 5, f. B. 1804. TYPE:
Caracas, Distr. Fed., Venezuela, Bredemeyer (B-Hb. Willd. 19468 not seen Tryon photo).

Mertensia pubescens Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:73. 1810. TYPE: Sta. Cruz,
Cuman^, Edo. Sucre, Venezuela, Humboldt & Bonpland (B not seen Morton photo).

Mertensia femignea Desw. Ges. Naturf. Freunde Berlin Mag. 5:307. 1811. TYPE: French Guiana,
collector unknown (P not seen photo 4481).

Mertensia immersa Kaulf. Enum. Fil. 38. 1824. TYPE: Ilha de Sta. Catarina, Est. Sta. Catarina,
Brazil, Chamisso (LE not seen fragm M not seen photo 8860).

Plants terrestrial, at 600-1700(2000) m elevation, at forest margins, along
roadsides, and on streambanks and landslides, from throughout the Flora area.
Also from the Antilles, Central America, and tropical South America.

232 GLEICHENIACEAE: 44. GLEICHENIA

331 G. bifida 335 G. gnidioides

GLEICHENIACEAE: 44. GLEICHENIA 233

This species differs slightly from specimens from the Greater Antilles and
Mexico to El Salvador that have been called Dicranopteris cubensis Underw. or
Mertensia flilva Desv. in having broad, pale reddish-orange, long-ciliate bud scales.
Gleichenia bifida is not so much a pioneer on clay banks, and often follows
Dicranopteris pectinata.

332, Gleichenia brevipubis Christ, Bull. Herb. Boissier II, 6:280. 1906.
LECTOTYPE: Valley of the Rio Navarro, Pcia. Cartago, 1400 m, Werckle in

1905 (P not seen photo 4478; isolectotypes CR, GH, UC, US fragm NY), chosen
by Lellinger (Proc. Biol. Soc. Wash. 89:713. 1977).

Plants terrestrial, at (600) 1400 -1900(2700) m elevation, in forests and on open
banks and roadsides, from La Palma de S. Ramon (Pcia. Heredia), the Cordillera
Central, the Cordillera de Talamanca to Pcia. Chiriqui, the Fila Costeha near S.
Vito, and Cerro Jefe. Also from Puerto Rico, Mexico, and Honduras.

This species has been confused with G. bifida, from which it differs in having an
extremely dense, compact, whitish tomentum on the abaxial surface of the laminae
and deciduous, atropurpureous, short-ciliate scales on the abaxial surface of the
costae.

333, Gleichenia compacta Christ, Bull. Herb. Boissier II, 5:254. 1905.

Gleichenia mellifera Christ, BiiU. Herb. Boissier II, 6:281. 1906. TYPE: Valley of the Rio

Navarro, Pcia. Cartago, 1400 m, Werckle (P not seen; isotjpe US fragm NY).

Gleichenia bradeomm Rosenst. Repert. Spec. Nov. Regni Veg. 10:274. 1912. TYPE: La Palma,
Pcia. S. Jose, 1400 m, Brade & Brade 501 (S not seen photo 6128; isotypes B not seen photos
10420-10422, UC, US fragm NY).

TYPE: La Palma, Pcia. S. Jose, 1600 m, Werckle (P not seen photos 4483-4485
fragm NY, fragm US).

Plants terrestrial, at 600-1400(3300) m elevation, in forests, ravines, and
exposed places, from the Cordillera Central, the Cordillera de Talamanca to Pcia.
Chiriqui, near Muheco (Pcia. Cartago), above El Valle, Cerro Azul, Cerro Jefe,
and Cerro Pirre.

As shown by Moore 6620 (BH, US), the scales and hairs on the abaxial lamina
surface are variable in this species. Gleichenia bradeomm and part of G. compacta
are terminal portions of laminae, whereas G. mellifera is a basal portion. The
scales on terminal portions are reddish-brown with rather long, lax setae; those of
the basal portions are darker brown and sometimes paler toward the center of the
scale, with fewer, shorter, and stiffer setae.

334, Gleichenia costaricensis (Underw.) C. Chr. Ind. Fil. Suppl. 1:44. 1913.

Dicranopteris costaricensis Underw. Bull. Torrey Bot. Club 34:253. 1907. TYPE: Volcan Poas,
Pcia. Alajuela, 2600 m,/. D. Smith 6859 (NY; isotypes B not seen, CR, GH, MO, US).

FIGS. 330 -337. Gleichenia. FIG. 330. Pinnules of G. bancroftii, Molina R. 11502, Honduras. FIG.
331. Ultimate segment of G. bifida, Molina, Williams et al. 17412. FIG. 332. Ultimate segments of
G. brevipubis, Skutch 3514. FIG. 333. Ultimate segments of G. compacta, Standley 33860. FIG. 334.
Ultimate segment of G. costaricensis, Brown CR-162. FIG. 335. Ultimate segments of G. gnidioides,
Lellinger & de la Sota 491. FIG. 336. Portion of lateral branch of G. hastulata, Brade & Brade 502.
FIG. 337. Portion of lateral branch of G. intermedia, Standley & Valerio 50023.

234 GLEICHENIACEAE: 44. GLEICHENIA

.25 cm

GLEICHENIACEAE: 44. GLEICHENIA 235

Plants terrestrial, at (1000)1500-3000 m elevation, in open areas and on road
banks, from La Balsa de S. Ramon (Pcia. Alajuela), the Cordillera Central, and
the Cordillera de Talamanca to the Cerro de la Muerte. Also from Colombia.

This species is very closely related to G. revoluta which has stiffer

marginal setae on the lamina scales and rhizome scales borne on low
protuberances.

335. Gleichenia gnidioides Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:266
(repr. 348). 1864.

TYPE: Acostadero, Depto. Choco, 2400 m, Triana (B not seen photo 10335;
isotype US).

Plants terrestrial, at 0-100 m elevation, in disturbed places, from the Caribbean
side of the divide at El Cope (Pcia. Code) and the central Choco. Also from high
elevations in the Cordillera Occidental bordering the Choco.

This species is distinct in having very short segments and a narrow angle
between the branches.

336. Gleichenia hastulata Rosenst. Repert. Spec. Nov. Regni Veg. 10:274. 1912.
TYPE: La Palma, Pcia. S. Jose, 1400 m. Erode & Erode 502 (S not seen photo

6140; isotype fragm NY, fragm US).

Plants terrestrial, at 1700-1800 m elevation, on banks, from La Palma and
Cerro Colorado above S. Felix (Pcia. Chiriqui).

337. Gleichenia intermedia Baker, J. Bot. Brit. For. 25:24. 1887.

Gleichenia axialis Christ, Bull. Herb. Boissier II, 5:14. 1905. TYPE: Costa Rica, Werckle in 1903
(P not seen photos 4494 - 4495; isotype US).

TYPE: Costa Rica, Cooper (K not seen; isotypes GH, US).

Plants terrestrial, at 1600-2400 m elevation, in forests, thickets, and roadsides,
from the Cordillera Central, the northern end of the Cordillera de Talamanca,
and Cerro Punta (Pcia. Chiriqui).

338. Gleichenia maritima Hieron. Bot. Jahrb. Syst. 34:562. 1905.

LECTOTYPE: Near Buenaventura, Depto. El Valle, Colombia, Lehmonn 4432

(B not seen photo 10337; isolectotype US), chosen by Lellinger (Proc. Biol. Soc.
Wash. 89:714. 1977).

Plants terrestrial, at 700 - 800 m elevation, in disturbed forests, in the Flora area
known only from near Km. 141 of the Ciudad Bolivar - Quibdo road, Depto.
Choco {Lellinger & de la Soto 883, COL, LP, US). Also from Pacific coastal
Colombia south of the Choc6.

339. Gleichenia nitidula Rosenst. Repert. Spec. Nov. Regni Veg. 10:275. 1912.

TYPE: S. Carlos to Buena Vista, Pcia. Alajuela, Erode {Ros. Fil Costar. Exs.
184) 503 (S not seen photos 10388-10389, M not seen photo 8863, NY, UC, US).

Plants terrestrial, at 600-900(1100) m elevation, in forests and along roadsides,
from La Balsa de S. Ramon (Pcia. Heredia), the Atlantic side of the Cordillera

FIGS. 338 - 342. Gleichenia. FIG. 338. Ultimate segment of G. maritima, Lellinger & de la Sota 883.
FIG. 339. Ultimate segment of G. nitidula. Cook & Doyle 30. FIG. 340. Portion of lateral branch of
G. orthoclada. Cooper. FIG. 341. Ultimate segment of G. pallescens, Brown CR-246. FIG. 342.
Base of ultimate branches of G. pteridella, Maxon 8473.

236 GLEICHENIACEAE: 44. GLEICHENIA

345 G. rubignosa

346 G. strictissima

347 D. flexuosa

348 D. pectinata

GLEICHENIACEAE: 44. GLEICHENIA 237

Central, the Fortuna Dam site (Pda. Chiriqui), west of Sta. Fe, north of El Valle,
and Cerro Jefe (Pda. Panama). Also from Colombia and Ecuador.

340. Glekhenia orthoclada Christ, Bull. Herb. Boissier II, 5:16. 1905.

Gleichenia intemtedia var. dissitifolia Baker, J. Bot. Brit. For. 25:24. 1887. TYPE: Valle de la
Estrella, Pda. Cartago, 1600” 1800 m, Cooper (K not seen; isotypes GH, US).

TYPE: Costa Rica, Werckle 212 (P not seen).

Plants terrestrial, at 1500 “1800 m elevation, on banks, from the Cordillera
Central. This species is said by Gomez (pers. comm.) to be the hybrid between G.
intermedia and G. retroflexa.

341. Gleichenia pallescens Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:267
(repr.349). 1864.

Gleichenia bicolor Christ, Bnll. Herb. Boissier II, 6:279. 1906. SYNTYPES: Valley of the Rio
Navarro, Pda. Cartago, 1400 m, Werckli (P not seen; isos3mtypes S not seen photo 6127); and
Chemin du Cascajal, WerckM (P not seen). The Goebel collection cited by Christ should be
considered a parat>pe.

TYPE: Velez, Edo. Merida?, Venezuela, 2100 m, Lindig 273 (B not seen;
isotypes K not seen Maxon photo 316 and fragm US, P not seen photo 4519).

Plants terrestrial, at 1300 “2100 m elevation, along roadside banks, from La
Palma de S. Ramon (Pda. Alajuela), the Cordillera Central, Tapanti, and above
Sta. Fe. Also from Venezuela and Colombia.

342. Gleichenia pteridella Christ, Bull. Herb. Boissier II, 6:284. 1906.

TYPE: Costa Rica, WerckM in 1903 (P not seen photos 4526 “4527; isotype US).
Plants terrestrial, at 1400-2000 m elevation, in open places and roadsides, from
the Cordillera de Tilaran, the Cordillera Central, and Cerro Tablazo. Also from
Nicaragua.

343. Gleichenia remota (Kaulf.) Sprang. Syst. Veg. ed. 16, 4:27. 1827.

Mertensia remota Kaulf. Enum. Fil. 39. 1824. TYPE: Ilha de Sta. Catarina, Est. Sta. Catarina,
Brazil, Chamisso (LE not seen).

Gleichenia trachyrhizoma CMst, Bull. Herb. Boissier 11, 6:280. 1906. TYPE: Valley of the Rio
Navarro, Pcia. Cartago, 1400 m, Werckle in 1905 (P not seen photo 4541; isotypes CR, UC, US).

Dicranopteris wiiliamsii Maxon, Amer. Fern J. 2:21. 1912. TYPE: Near Cana, Pcia. Darien, R. S.
Williams 917 (US; isotype NY).

Gleichenia aequilateralis Jemn. Ferns Brit. W. Ind. Guiana 353. 1909, as “aequilateralel’ TYPE:
Upper Demerara River, Guyana, 4149 (NY? not seen).

Plants terrestrial, at 0-1400 m elevation, in forests and along trails and
roadsides, from the valley of the Rio Navarro (Pcia. Cartago), the Cordillera de
Talamanca above S. Isidro del General, the Peninsula de Osa, north of El Valle,
near Gamboa (Canal Zone), Cerro Jefe, southeast of Puerto Obaldfa (Com. S.
Bias), and along the Ciudad Bolivar - Quibdo road at ca. Km. 141 (Depto.
Choco). Also from Cuba, Trinidad to Brazil, Venezuela, and Colombia.

FIGS. 343 ” 348. Gleichenia and Dicranopteris. FIG. 343. Ultimate segments of G. remota, Mickel
2757. FIG. 344. Lateral branches of G. retroflexa, Maxon 510. FIG. 345. Ultimate segment of G.
mbipnosa, Lellinger & de la Sota 744. FIG. 346. Ultimate segment of G. strictissima, Chrysler 4990.
FIG. 347. Lateral branches of D. flexuosa. Smith 2107, Mexico. FIG. 348. Lateral branch of D.
pectinata, Scamman 6987.

238 GLEICHENIACEAE: 45. DICRANOPTERIS

344. Gleichenia retroflexa Bomm. & Christ, Bull. Herb. Boissier 4:657. 1896.

Gleichenia intermedia var. flexuosa Baker, J. Bot. Brit. For. 25:24. 1887. TYPE: Costa Rica,
Cooper (K not seen; isotypes GH, US).

TYPE: La Palma, Pda. S. Jose, Pittier 1471 (BR not seen photo 4751; isotype
US).

Plants terrestrial, at 600-2500 m elevation, often on exposed day banks, from
the Cordillera Central, Cerro Tablazo, and the northern end of the Cordillera de
Talamanca.

345. Gleichenia nibiginosa Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:267
(repr. 349). 1864.

Gleichenia rubiginosa f. virescens Hieron. Bot. Jahrb. Syst. 34:561. 1905. SYNTYPES: Peru,
Mathews 1092 (B not seen); and Colombia, Schmidtchen in 1882 (B not seen).

TYPE: Puente Nacunol, Depto. Santander, Colombia, 1900 m, Lindig 71 (B not
seen photo 10394; isotypes P not seen photo 4504, K not seen Maxon photo 317,
fragm US).

Plants terrestrial, at 1400-1800 m elevation, in forests, from Pico Mali and Alto
del Buey (both Depto. Choco). Also from Venezuela to Peru.

This species is doubtfully distinct from G. strictissima, and differs in being larger
and in having longer segments with more pairs of veins.

346. Gleichenia strictissima Christ, Bull. Herb. Boissier H, 5:13. 1905.

Gleichenia glaucina Christ, Bull. Herb. Boissier II, 6:283. 1906. SYNTYPES: Navarro, Pcia.
Cartago, Werckli in 1905 (P not seen photo 4536); and La Palma, Werckle (P not seen). A
presumable unlocalized isosyntype is at US.

TYPE: Costa Rica, Werckle 215 in 1903 (P not seen photo 4537; isotype US).
Plants terrestrial, at 800 - 1800 m elevation, on hillsides and wet banks, from S.
Ramon, the Cordillera Central, near Tapanti, and Cerro Horqueta (Pcia.
Chiriqui).

This species may only be a small form of G. rubiginosa.

45. DICRANOPTERIS Bemh.

Plants terrestrial, usually in disturbed places like roadbanks and landslides,
often a pioneer on bare soil; rhizomes long-creeping, thin, occasionally forked,
hairy, the hairs pluricellular, rather lax to bristle-like, brown; fronds medium to
large-sized, often sprawling over other vegetation or forming thickets; stipes long,
terete, slightly hairy at the base; laminae debate to oblong, the rachis bearing
distant, opposite pairs of pinnae flanking a terminal bud that elongates to continue
the rachis or remains dormant at the frond apex, the bud protected by dark,
reddish-brown, pluricellular, straight hairs; pinnae pseudodichotomously divided
(appearing nearly pinnately divided in D. pectinata), with only the ultimate
division laminate, these laminae pectinate, oblong to narrowly elliptic, glabrous
adaxially, hairy or glaucous abaxially, mostly subcoriaceous, the axes bearing
simple or stellate, brownish hairs or glabrous; veins 2-forked; sori round,
exindusiate, composed of usually 8-15 subsessile sporangia, paraphyses absent.

Pantropical, mostly at low elevations; 10 species.

MAXON, W. R. 1909. Gleicheniaceae. North Amer. FI. 16(l):53-63.

GLEICHENIACEAE: 45. DICRANOPTERIS 239

1. Reflexed, pirmatifid accessory pinnae absent from each fork of the lamina; pinnae unequally
dichotomous, appearing pinnate, with the pinnules (ultimate divisions) equally dichotomous or nearly
so; plants usually large, the laminae glaucous on the abaxial surface.

348. D. pectinata

1. Reflexed, pinnatifid accessory pinnae present at ail but the ultimate forks of the lamina; pinnae
equally dichotomous throughout with the central axis of the pinnae often prolonged above the basal
dichotomy; plants often medium-sized, not glaucous on the abaxial surface.

347. D. flexuosa

347. Dicranopteris flexuosa (Schrad.) Underw. Bull. Torrey Bot. Club 34:254.
1907.

Mertensia flexuosa Schrad. Gott. Gel. Anz. 1824:863. 1824. TYPE: Brazil, Prince Maxmillian von
Wied-Neuwied (BR not seen photo BM not seen), cited by Proctor (Ferns Jamaica 82. 1985).
Usually Schrader types are to be sought at GOET or LE.

Mertensia rigida Kunze, Linnaea 9:16, 1834. TYPE: Near Chibangata, Peru, Poeppig 1153 (LZ
destroyed; possible isotype L not seen photo 1410).

Mertensia pumila Mart. Icon. PI. Crypt. Bras. Ill, t. 60, f. 2. 1834. TYPE: Serra da Estrella, Est.
Rio de Janeiro, Brazil, Martius (M not seen photo 8868).

Mertensia scalpturata Fee, Crypt. Vase. Bresil 1:199, t. 72, f. 1. 1869. SYNTYPES: Brazil,
Claussen 102a (P not seen), Glaziou 364 (P not seen photo 4502), Rio de Janeiro, Est. Rio de
Janeiro, Brazil, Glaziou 1695 (P not seen photo 4503).

Plants terrestrial, at 0"1800 m elevation, on hillsides and roadside banks, from
the vicinity of Tarbaca (Pcia. S. Jose), Buenos Aires (Pcia. Puntarenas), Cerro
Horqueta (Pcia. Chiriqui), between Canazas and the foot of the Cordillera Central
(Pcia. Veraguas), between Las Minas and Pese (Pcia. Hererra), Barro Colorado
Island (Canal Zone), and Taboga Island and Cerro Azul (both Pcia. Panama).
Also from throughout most of tropical America.

348. Dicranopteris pectinata (Willd.) Underw. Bull. Torrey Bot. Club 34:260.
1907.

Mertensia pectinata Willd. Kongl. Vetensk. Acad. Nya Hand!. 25:168, t. 4. 1804. TYPE: Caracas,
Distr. Fed., Venezuela, Bredemeyer (B-Hb. Willd. 19465 not seen Tryon photo).

Mertensia glaucescens Humb. & Bonpi. ex Willd. Sp. PL ed. 4, 5:72. 1810. TYPE: Cumana, Edo.
Sucre, Venezuela, Humboldt & Bonpland (B not seen Morton photo; isotype P not seen photo 4525).

Mertensia brasiliana Desv. Ges. Naturf. Freunde Berlin Mag. 5:329. 1811. TYPE: Brazil, collector
unknown (P not seen photo 4522).

Mertensia emofginata Raddi, PL Bras. Nov. Gen. 1:72, t. 6. 1825, nom. superfl. TYPE: A renaming
of Mertensia glaucescens Humb. & BonpL ex W'dld., which was cited in synonymy, and so based on
the type of that name.

Giekhenia nitida K. Presl, Reliq. Haenk. 1:70. 1825. TYPE: Mexico, Haenke (PRC not seen
fragms NY not seen, US).

Mertensia elata Desv. Mem. Soc. Linn. Paris 6:201. 1827. TYPE: Jamaica, collector unknown (P-
Hb. Desv. not seen Cintract photo at UC).

Mertensia glaucescens var. cubense Fee, Hist. Foug. Antill. [Mem. Foug. 11]:121. 1866. TYPE:
Cuba, collector unknown (not seen).

Mertensia glaucescens var. madcana Fee, Hist. Foug. Antill. [Mem. Foug. 11]: 121. 1866.

SYNTYPES: Mexico, Galeotti 6402 (P not seen); and Hispaniola, de Tussac (P? not seen).

Mertensia dichotoma var. Uherminieri Fee, Hist Foug. Antill. [M6m. Foug. 11]: 121. 1866. TYPE:
Calumet, Guadeloupe, UHerminier 97 (presumably P-Hb. Bory not seen; isotype US).

Giekhenia linearis var. depauperata Christ, Bull. Herb. Boissier II, 5:14. 1905. TYPE: Costa Rica,
Werckle (P not seen).

240 GLEICHENIACEAE: 45. DICRANOPTERIS

Gleichenia pectinata var. sublinearis Christ, Bull. Herb. Boissier II, 6:282. 1906. LECTOTYPE:
Navarro, Pcia. Cartago, 3500 ft, /. D. Smith 4994 (US; isolectotype P not seen), chosen by Lellinger
(Proc. Biol. Soc. Wash. 89:714. 1977).

Plants terrestrial, at 0-1700 m elevation, often on stream banks or along roads,
from the Cordillera Central, the Cordillera de Talamanca, throughout Panama
and the Choc6. Also from throughout the Antilles, Central America, and tropical
South America.

This species commonly is a pioneer on road cuts.

POLYPODIACEAE: 46. PLEOPELTIS 241

POLYPODIACEAE

Rhizomes long-creeping, short-creeping, ascending, or erect, ca. 1-10 mm in
diam., scaly (the scales rarely absent), the scales entire or erose, often bearing
surficial hairs, marginal hairs, or marginal setae; fronds minute to medium-sized;
stipes articulate (the phyllopodia low) or not; laminae simple and entire or lobed,
pinnatifid, pinnatisect, pinnate, or rarely more divided, usually glabrous,
sometimes softly pubescent or bearing stiff, usually dark setae, the midribs or
rachises sometimes also scaly; veins free to fully anastomosing without sterile
included veinlets in most genera (free included veinlets present in Microgramma
and Pleopeltis); sori superficial or sunken into the lamina tissue, rarely elongate or
linear, usually round and borne at the apex of a vein or at the junction of several
veins, this sometimes ring-shaped; indusia absent; sporangia long-stalked, globose;
linear to clavate or branched paraphyses often present (caducous, peltate, scale-
like paraphyses present in Pleopeltis); spores monolete or trilete.

1. Laminae narrowly spathulate, acuminate at the apex, attenuate at the base, the stipes obsolete;
indusia elongate to linear and nearly parallel to the midrib; rhizome scales strongly clathrate with
iridescent lumina.

60, Loxogramme

1. Plants not as above.. 2.

2(1). Stipes usually variously sulcate, rarely wiry (terete and often wiry in Pecluma), articulate to
the rhizome, the older portions of the rhizome bearing low, usually closely set phyllopodia (except in
Hyalotrichopteris)', rhizomes generally short- to long-creeping and conspicuous, often branched;
laminae glabrous or scaly or soft hairy, at least on the rachis or midrib,. 5.

2(1), Stipes always terete, usually wiry, not or apparently not articulate to the rhizome, the
rhizomes lacking phyllopodia: rhizomes generally erect or ascending and inconspicuous, unbranched;
Icuninae glabrous or commonly bearing stiff, dark, setiform hairs on the surface, margins, or rachis or
in the sori..3.

3(2), Sporangia in linear coenosori, 1 on each side of the midrib of the fertile lamina or fertile
portion of the lamina.

59. Cochlidium

3(2). Sporangia in round or elongate sori, several on each side of the midrib.. 4.

4(3). Laminae entire, repand, or shallowly lobed, commonly thick and spongy; sori irregular in
shape, mostly at least slightly elongate, often in pits in the lamina tissue.

58. Glyphotaenium

4(3). Laminae entire, deeply lobed, pinnatifid, pinnatisect, or rarely pinnate or more divided, rarely
thick and spongy; sori regular, round, rarely in pits in the lamina tissue.

57. Grammitis

5(2). Laminae pinnatisect with narrow, closely set segments; stipes terete, mostly dark brown or
blackish; rhizome scales attached at the base.

56. Pecluma

5(2). Laminae entire, shallowly to deeply lobed, pinnatifid, pinnatisect, pinnate, or more divided, if
pinnatisect, the segments relatively broad and distant; stipes variously sulcate, often pale brown;
rhizome scales peltately attached.. 6.

6(5). Sori superficial on the abaxial surface, round or occasionally elongate..8.

6(5), Sori marginal, linear, rarely interrupted,.?.

7(6). Fronds several times dichotomously branched, usually bearing minute scales on the abaxial
surface; stipes and axes blackish.

50. Dicranoglossum

7(6). Fronds simple, entire, glabrous on the abaxial surface; stipes and axes pale brown.

49. Neurodium

242 POLYPODIACEAE: 46. PLEOPELTIS

8(6). Laminae simple or shallowly lobed (pinnate wth unlobed pinnae in a few species of
Campyloneumm, with prominent primary veins and two rows of sori between the veins); rhizomes
mostly very long-creeping, wire- or cord-like, occasionally thick and short-creeping (as in

Niphidium)..!!.

8(6). Laminae deeply lobed, pinnatifid, pinnatisect, or more divided; rhkomes mostly ascending or
short-creeping, rather thick..9.

9(8). Venation free (1- to several times forked), casually anastomosing, or with 1 to several rows of
areolae with a single unbranched excurrent veinlet in each areola.

55. Poiypodium

9(8). Venation mostly of polygonal areolae lacking excurrent sterile veinlets, the excurrent fertile
veinlets double and fused under the round sori or irregularly disposed under the elongate sori.. 10.

10(9). Sori round; rhizomes ca. 5 mm thick, contorted, bearing orangish scales; veins decidedly
prominulous on the abaxial surface of the laminae.

54. Phlebodium

10(9). Sori elongate to oval, rarely round; rhizomes ca. 3 mm thick, not contorted, bearing pale
brown scales; veins scarcely prominulous on the abaxial surface of the laminae.

47. Pseudocolysis

11(8). Laminae with decidedly prominulous main and transverse veins, the tertiary veins many and
irregular; sori in a single row between the main veins.

51. Niphidium

11(8). Laminae not as above.. 12.

12(11). Venation partially or entirely of polygonal areolae, lacking straight or flexuous,
prominulous or occult main side veins.. 14.

12(11). Venation of straight or flexuous, prominulous or occult main side veins and transverse
secondary veins bearing (1)2(3 or more) unbranched excurrent fertile and sterile veinlets.. 13.

13(12). Laminae thinly pilose, the hairs ca. 1 mm long, multicellular, clear, with prominent cross-
walls; plants epipetric; fronds 3-9 cm long.

52. Hyalotrichopteris

13(12). Laminae lacking hairs (except thinly pilosulous, the hairs ca. 0.1 mm long in
Campyloneumm occultum); plants usually epiphytic; fronds mostly longer than 10 cm.

53. Campyloneumm

14(12). Sori (1)2 per fertile areola; laminae glabrous or sometimes sparsely scaly on the abaxial
surface of the midribs. Midrib scales narrowly lanceolate or narrowly lanceate.

53. Campyloneumm

14(12). Sori 1(2) per fertile areola; laminae glabrous to densely scaly on both surfaces..l5.

15(14). Laminae dull, sparsely to densely scaly on both surfaces, the scales orbicular to lanceolate;
midribs usually blackish on the abaxial surface, darker than the laminae; sori round to oval, usually
protected by peltate paraphyses when very young.

46. Pleopeltis

15(14). Laminae shiny, glabrous to sparsely scaly on both surfaces, the scales adcular from an
orbicular base; midribs not blackish on the abaxial surface; sori round, mixed with usually
inconspicuous non-peltate paraphyses.

48. Microgramma

46. PLEOPELTIS Humb. & Bonpl.

Plants epiphytic; rhizomes wiry, very long-creeping, scaly, the scales peltate or
lanceolate with a peltate attachment, weakly clathrate or not clathrate, often
irregular at the margin; stipes articulate to the rhizome or to low phyllopodia,
often (with the midrib) blackish; fronds small, simple; laminae usually rather
narrowly elliptic, chartaceous to subcoriaceous, dull, sparsely scaly, the scales
small to minute, peltate or lanceolate; veins irregularly anastomosing with free

POLYPODIACEAE: 46. PLEOPELTIS 243

included veinlets; sori large, round or sometimes elongate, protected when young
by overlapping scales (except P, percussa, with abundant dark, reddish-brown
sporangiasters), exindusiate; sporangia long-stalked.

Pantropical; ca. 40 species.

WEATHERBY, C. A. 1922. The group of Polypodium lanceolatum in North America. Contr. Gray
Herb. 65:3-14.

1. Rhizome scales not clathrate or only the central cells so, with small and rather obscure lumina
and with the pale, marginal cells often worn away in age leaving only the central cells..4.

1. Rhizome scales entirely clathrate or only one row of marginal cells not so, with large, clear
lumina and with the marginal cells not worn away in age..2.

2(1). Sori confluent, forming a single coenosorus on each side of the midrib; sterile laminae 6-10
times wider than the fertile ones. Sterile laminae lanceolate or elliptic, 3- 11 cm long, 1-3 cm wide;
fertile laminae linear, 4- 10 cm long, 3-5 mm wide.

356. P. wiesbaurii

2(1). Sori oval or elongate, but not confluent; sterile laminae (1)1.5 -4 times wider than the fertile
ones..3.

3(2). Sterile laminae 2-4 times wider than the fertile ones; rhizomes 2-3 mm in diam.; midribs of
the sterile fronds stramineous abaxially. Sterile laminae elliptic-lanceolate, round or acute at the
apex, acute at the base, 4- 11 cm long, 0.8 -2.8 cm wide; fertile laminae linear, 4- 12 cm long, 0.4-1
cm wide.

350. P. fructuosa

3(2). Sterile laminae (1)1.5 -2(2.5) times wider than the fertile ones; rhizomes 1 mm in diam.;
midribs of the sterile fronds blackish abaxially. Sterile laminae elliptic, acute to acuminate at the
base, acute at the apex, 4-8 cm long, 1.5-3 cm wide; fertile laminae linear, 5-9 cm long, 1(1.5) cm
wide.

354. P. panamensis

4(1). Rhizome scales orbiculate, peltate, ca. 0.5 mm in diam., blackish at the center, with pale,
ciliate-toothed margins, bearing on their surface abundant, reddish-brown, unicellular trichomes up
to ca. 1 mm long; sori oval to oblong. Stipes 0.5-2 cm long; laminae usually linear, 2.5-9(10) cm
long, 0.4- 1(1.6) cm wide.

349. P. astrolepis

4(1). Rhizome scales lanceolate, peltately attached near the base, 1-4 mm long, bicolorous
(except in P. percussa), not concealed by reddish-brown trichomes; sori round to oval..5.

5(4). Paraphyses in sori abundant, dark reddish-brown, with a few yellowish sporangia scattered
among them. Stipes 0.5-4 cm long; laminae linear, acuminate to attenuate at the base, often
caudate at the apex, 10 - 20(25) cm long, 1- 1.5 (2.5) cm wide.

355. P. percussa

5(4). Paraphyses in sori few to none. Laminae 6-20 cm long, 1- 1.5(2) cm wide..6.

6(5). Laminae shallowly to deeply and irregularly lobed. Laminae lanceolate, 12-14 cm long,
1.5-3 cm wide.

351. P. leucospora

6(5). Laminae entire..?.

7(6). Stipes compressed; sori 1-3 mm wide; rhizome sc2iles 1-2 mm long. Stipes 0.5-6 cm long;
laminae 8-15(20) cm long, 0.5- 1.5(2) cm wide; sori submarginal.

352. P. macrocarpa var. complanata

7(6). Stipes terete; sori 2.5-5 mm wide; rhizome scales (1.5)2 -4 mm long. Stipes 1.5-9 cm long;
laminae 10-20 cm long, 1 - 2(2.5) cm wide; sori supramedi£d to submarginal.

353. P. macrocarpa var. macrocarpa

244 POLYPODIACEAE: 46. PLEOPELTIS

POLYPODIACEAE: 46. PLEOPELTIS 245

349. Pleopeltis astrolepis (Liebm.) Foum. Mexic. PI. 1:87. 1872.

Grammitis lanceolata Schkuhr, Vier Zwanzigste Kl. Linn. Pfl.-Syst. 1:9, t. 7 p.p. 1804, non
Pleopeltis lanceolata. Kaulf., 1824. TYPE: Jamaica, Swartz (HAL not seen). Schkuhr’s citation of G.
lanceolata Swartz refers to an Old World specimen and is in error.

Grammitis elongata Swartz, Syn. Fil. 22, 213. 1806, non Pleopeltis elongata Kaulf., 1824. TYPE:
Based on G. lanceolata sensu Schkuhr excl. syn.

Grammitis revoluta Spreng. ex Willd. Sp. PI. ed. 4, 5:139. 1810, non Pleopeltis revoluta v. A. v. R.,
1909. TYPE: Without locality, collector unknown (B-Hb. Willd. 19584 not seen microfiche S. I.
Library).

Grammitis squamulosa Sphtg. Tijdschr. Natuurl. Gesch. Physiol. 7:398. 1840, non Pleopeltis

squamulosa (Kaulf.) K. Presl, 1836, nom. illeg. TYPE: Surinam, collector unknown (L not seen).

Polypodium astrolepis Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:185
(repr. 33). 1849. LECTOTYPE: Trapiche de la Concepcion, Edo. Oaxaca, Mexico, Liebmann FI.
Mex. 87 in 1841, lower plant (C not seen), chosen by A. R. Smith (FI. Chiapas 2:177. 1981).

Drynaria prieurii Fee, Gen. Fil. [Mem. Foug. 5]:271. 1852. SYNTYPES: French Guiana, Leprieur
(RB not seen; possible isotype US); and Martinique, Rivoire (P or RB not seen).

Pleopeltis angustifolia D. C. Eaton, Amer. J. Sci. Arts II, 27:198. 1859. TYPE: Based ultimately on
Grammitis elongata Swartz, and so based on the type of that name.

Taenitis swartzii Jenm. J. Bot. Brit. For. 17:263. 1879, nom. superfl. TYPE: A renaming of
Grammitis elongata Swartz, and so based on the type of that name.

Plants epiphytic, at 0 - 1400(3000) m elevation, in forests and on trees in open
areas, from the Cordillera Central, the Meseta Central, Cerro Carpintera, the
Atlantic coastal plain of Costa Rica, the Cordillera de Talamanca to Pcia.
Chiriqui, El Valle, and Alcalde Diaz (Pcia. Panama). Also from throughout
tropical America.

350. Pleopeltis fructuosa (Maxon & Weath. in Weath.) Lellinger, Proc. Biol. Soc.
Wash. 89:722. 1977.

Polypodium fmctuosum Maxon & Weath. in Weath. Contr. Gray Herb. 65:12. 1922. TYPE:
Upper Caldera River, Holcomb’s trail above Boquete, 1450-1650 m, Maxon 5689 (US).

Plants epiphytic, at 1000-2500 m elevation, in forests, from the Cordillera
Central, Cerro Carpintera, and the Cordillera de Talamanca to Pcia. Chiriqui.

351. Pleopeltis leucospora (Klotzsch) T. Moore, Ind. Fil. 77. 1857.

Polypodium leucosporum Klotzsch, Linnaea 20:404. 1847. TYPE: Paramo de Mucuchfes, Edo.
Merida, Venezuela, Moritz 306 (B not seen).

Polypodium lanceolatum var. elisabethae Jenm, Bull. Bot. Dept. Jamaica N.S., 4:199. 1897.

TYPE: Near Cinchona, Jamaica, collector unknown \Jenmanl] (IJ not seen U. S. Natl. Herb, photo
3038).

Plants epiphytic, at 2000-2100 m elevation, in forests and in open areas, in the
Flora area known only from the Laguna de Zarcero, Pcia. Alajuela {A. Smith 1432,
NY). Also from Jamaica, Hispaniola, Venezuela, Colombia, and Peru.

Wagner (Fern Gaz. 11:125-135. 1975) demonstrated that this is the hybrid
between Polypodium (Marginaria) thyssanolepis A. Braun and Polypodium
lanceolatum L., which is a synonym of Pleopeltis macrocarpa (Bory ex Willd.)

FIGS. 349-356. Pleopeltis. FIG. 349. Plant of P. astrolepis, Alfaro 6026. FIG. 350. Plant of P.
fmctuosa, Standley & Valerio 49243. FIG. 351. Plant of P. leucospora, Smith 1432 (NY). FIG. 352.
Plant of P. macrocarpa var. complanata, Knight. FIG. 353. Plant of P. macrocarpa var. macrocarpa,
Maxon 8486. FIG. 354. Plant of P. panamensis, Maxon 5784. FIG. 355. Plant of P. percussa. Stork
2282. FIG. 356. Plant of P. wiesbaurii, Mickel 2993.

246 POLYPODIACEAE: 46. PLEOPELTIS

Kaulf. var. macrocarpa. I place this hybrid in Pleopeltis because in frond and
rhizome form it seems more allied to the other species of that genus than it does
to the species of Polypodium subg. Marginalia. If hybrids between Pleopeltis and
Polypodium are assigned to a separate genus, the correct name for this hybrid is
Pleopodium xleucosporum (Klotzsch) Mickel & Beitel.

352. Pleopeltis macrocarpa var. complanata (Weath.) Lellinger, Proc. Biol. Soc.
Wash. 89:722. 1977.

Polypodium lanceolatum var. complanatum Weath. Contr. Gray Herb. 65:8. 1922. TYPE: Juan
Vinas, Pcia. Cartago, Pittier 1855 (US).

Plants epiphytic, at 1200-2200(2500) m elevation, in forests and open areas,
from the Cordillera de Tilaran, the Cordillera Central, and the Cordillera de
Talamanca to Pcia. Chiriqui.

353. Pleopeltis macrocarpa (Bory ex Willd.) Kaulf. Berlin Jahrb. Pharm. 21:41.
1820, var. macrocarpa.

Polypodium lanceolatum L. Sp. PI. 2:1082. 1753, non Pleopeltis lanceolata Kaulf., 1824. TYPE:
Plate 6, f. 2 of Petiver’s “Pterigraphia Americana,” which is a transposed redrawing of t. 137 of
Plumier’s “Traite...”, which illustrates a specimen collected by Plumier on Hispaniola. See also Pichi
Sermolli (Webbia 20:349. 1965) for a discussion of the typification of this species.

Polypodium macrocarpum Bory ex Willd. Sp. PI. ed. 4, 5:147. 1810. TYPE: Mauritius, Bory (B-
Hb. Willd. 19629 not seen; isotype P not seen).

Plants epiphytic, at (600)1600-3100 m elevation, in forests and open areas,
from the Hacienda Santamaria in the Cordillera de Tilaran (Pcia. Alajuela), the
Cordillera Central, and the eastern slopes of Volcan Chiriqui. Also from Cuba,
Jamaica, Hispaniola, Mexico to Honduras, Nicaragua, Venezuela, Colombia to
Chile, Guyana, Brazil, Uruguay, and Argentina.

354. Pleopeltis panamensis (Weath.) Pic. Ser. Webbia 23:189. 1968.

Polypodium panamense Weath. Contr. Gray Herb. 65:13. 1922. TYPE: Around Portobelo, Pcia.
Colon, 5 - 200 m, Maxon 5784 (US; isotypes F, GH, NY).

Plants epiphytic, at 0-200 m elevation, from Bismarck (Pcia. Code), Ft.
Sherman (Canal Zone), Portobelo and the Rios Medio and Guanche (all Pcia.
Colon).

355. Pleopeltis percussa (Cav.) Hook. & Grev. Icon. Fil. t. 67. 1828.

Polypodium percussum Cav. Descr. PI. 243. 1801. TYPE: Peru, Nee (MA not seen Killip photo).
Polypodium rostratum Cav. Descr. PI. 244. 1801, non Burm., 1768, nom. illeg. TYPE: Mt. S.
Antonio, Ecuador, Nee (MA not seen), examined by Christensen (Dansk Bot. Ark. 9(3):11. 1937).

Polypodium avenium Desv. Ges. Naturf. Freunde Berlin Mag. 5:314. 1811. TYPE LOCALITY:
Brazil. Synonymized by Desvaux (Mem. Soc. Linn. Paris 6:227. 1827).

Polypodium cuspidatum K. Presl, Reliq. Haenk. 1:20, t. 1, f. 3. Jun-Nov 1825, non Don, 1 Feb
1825, nom. illeg. TYPE: Panama, Haenke (PRC not seen).

Polypodium stigmaticum K. Presl, Reliq. Haenk. 1:20, t. 3, f. 2. 1825. TYPE: Mountains at
Huanuco, Depto. Huanuco, Peru, Haenke (PRC not seen).

Polypodium microlepidum Desv. Mem. Soc. Linn. Paris 6:228. 1827. TYPE: Peru, collector
unknown (P-Hb. Desv. not seen), synonymized by Weatherby (Contr. Gray Herb. 114:32. 1936).

Polypodium elongatum Desv. Mem. Soc. Linn. Paris 6:228. 1827, non Aiton, 1789, nom. illeg.
TYPE: Brazil, collector unknown (P-Hb. Desv. not seen), examined by Weatherby (Contr. Gray
Herb. 114:30. 1936).

POLYPODIACEAE: 47. PSEUDOCOLYSIS 247

Polypodium haenkeanum Spreng. Syst. Veg. ed. 16, 4:46. 1827, nom. superfl. TYPE: A renaming
of P. stigmaticum K. Presl, and so based on the type of that name.

Polypodium percussum var. squamosum Christ in Dur. & Pitt. Bull. Soc. Roy. Bot. Belgique 35,
Mem. 232. 1896. TYPE: Carrillo, Pcia. S. Jose, 300 m, Pittier 1164 (BR not seen).

Plants epiphytic, at 0-1200(1700) m elevation, in forests and open areas, from
throughout the Flora area. Also throughout tropical Central and South America.

356. Pleopeltis wiesbaurii (Sodiro) Lellinger, Proc. Biol. Soc. Wash. 89:723. 1977.

Drymoglossum wiesbaurii Sodiro, Anales Univ. Quito 11(75):312 (repr. 419). 1894. TYPE: Rio
Chimbo, Pcia. Guayas, Ecuador, 300 - 500 m, Sodiro (Hb. Sodiro not seen).

IPolypodium bonapartii Rosenst. Repert. Spec. Nov. Regni Veg. 7:309. 1909. TYPE: Rio

Chasuan, Mt. Chimborazo, Pcia. Chimborazo, Ecuador, Spruce 5731 (S? or P? not seen Bonaparte
photo).

Pteropsis underwoodiana Maxon, Contr. U. S. Natl. Herb. 16:51, t. 28. 1912. TYPE: Near Suerre,
Llanuras de Sta. Clara, Pcia. Limon, 300 m,/. D. Smith 6941 (US; isotype NY).

Plants epiphytic, at 0-1200 m elevation, in forests, from the Cordillera de
Tilaran, the foothills of the Cordillera Central, the Meseta Central, the Atlantic
coastal plain of Costa Rica, the valley of the Rfo General, and above El Valle and
El Cope (Pcia. Code). Also from Ecuador.

This species is probably related to, or may even be conspecific with, the little-
known Ecuadorian Polypodium chionolepis Sodiro. The only character that sets
this species, which is the type of the genus Marginariopsis, apart from other species
of Pleopeltis is its coenosori. In venation and chromosome number (jc = 35,
according to F. Wagner, pers. comm.), it is a Pleopeltis; its rhizome scales are like
those of P. fructuosa and P. panamensis. Either of the latter species could be
considered Marginariopsis, but both lack coenosori, and P. panamensis is not
always dimorphic. Since a continuum of characteristics occurs in these three
species, generic or even infrageneric separation is not warranted for any of them.

47. PSEUDOCOLYSIS Gomez

Plants epiphytic; rhizomes thin, long-creeping, scaly, the scales linear-
lanceolate, attached at the base, weakly clathrate at the center, with pale, erose
margins; stipes articulate to the rhizome, narrowly alate for most of their length;
fronds medium-sized; laminae deeply pinnatifid (juveniles often simple),
herbaceous, dull, scaly, the scales minute, peltate; veins irregularly anastomosing
with a few included veinlets; sori linear, ascending, exindusiate, lacking
paraphyses; sporangia long-stalked.

Central America; 2 species.

GOMEZ P., L. D. 1977. Contribuciones a la pteridologia Centroamericana II. Novitates. Brenesia
10/11:115-119.

357. Pseudocolysis bradeorum (Rosenst.) G6mez, Brenesia 10/11:116. 1977.

IPolypodium guatemalense Kloztsch, Allg. Gartenzeitung 23:33. 1855. TYPE: Guatemala,

Warsczewiczl (B? not seen).

Polypodium bradeorum Rosenst. Repert. Spec. Nov. Regni Veg. 10:279. 1912. TYPE: Llanuras
de S. Carlos, Pcia. Alajuela, 200 m, Brade & Brade 460 (S not seen photo 5957; isotypes UC, US).

Plants terrestrial, at 100-200 m elevation, in forests, from the northern Atlantic
coastal plain of Costa Rica and Changuinola (Pcia. Bocas del Toro). Also from
Mexico, Belize, and Nicaragua.

248 POLYPODIACEAE: 47. PSEUDOCOLYSIS

POLYPODIACEAE: 48. MICROGRAMMA 249

The epithet guatemalense is earliest, if P. guatemalense is synonymous.

48. MICROGRAMMA K. PresI

Plants epiphytic; rhizomes usually wiry, long-creeping, scaly, the scales
lanceolate, sometimes narrowly so with a hair-like apex, not clathrate except for
the peltate attachment, usually entire; stipes obsolete or short and alate, articulate
to the rhizome or to low phyllopodia; fronds small, simple, subdimorphic or
dimorphic (the fertile fronds narrow); laminae ovate or elliptic to linear,
chartaceous, shiny, mostly glabrous, the midribs pale; veins irregularly
anastomosing with free included veinlets; sori usually small to large, round
(elongate in a few species); paraphyses often present; sporangia long-stalked.

Neotropics and tropical Africa and adjacent islands; ca. 20 species.

1. Laminae glabrous on both surfaces or with a few narrow scales on the abaxial surface of the
midrib..3.

1. Laminae sparsely to densely scaly on both surfaces. Fertile laminae much narrower than the
sterile ones.. 2.

2(1). Sterile fronds usually lanceolate or elliptic, rarely suborbiculate, (8)15-40(55) mm long,
sparsely covered on both surfaces with linear scales less than 1 mm long having a peltate base much
wider than the apical portion of the scale.

362. M. reptans

2(1). Sterile laminae usually suborbiculate to ovate, rarely lanceolate, 3-15(18) mm long, densely
covered on both surfaces with narrowly lanceolate scales ca. 1 mm long having a peltate base only
sUghtly wider than the apical portion of the scale.

363. M. tecta var. nana

3(1). Fertile laminae much narrower than the sterile ones; spherical tubers 2-3 cm in diam.
present on the rhizomes. Sterile laminae 3 - 12 cm long, 5-13 mm wide, acute or acuminate at the
apex, attenuate and exstipitate at the base.

358. M. brunei

3(1). Fertile laminae only shghtly narrower than the sterile ones; spherical tubers absent. .4.

4(3). Rhizomes (including the scales) 2-4(5) mm in diam. Sterile laminae broadly to narrowly
elliptical, acute at the base, 6- 11 cm long, (1.2)2 -4 cm wide.

360. M. nitida

4(3). Rhizomes (including the scales) 1-2(3) mm in diam...5.

5(4). Sori round; sterile laminae 2-9.5 cm long, 0.8- 1.7 cm wide; rhizome scales reddish, ca. 0.5
mm wide.

359. M. lycopodioides

5(4). Sori shghtly elongate, at ca. a 45® angle to the midrib; sterile laminae 7-17 cm long, 1.5 -2.5
cm wide; rhizome scales whitish, ca. 1 mm wide.

361. M. persicariifolia

358. Microgramma brunei (Werckl^ ex Christ) Lellinger, Amer. Fern J. 67:59.

1977.

Polypodium brunei Werckle ex Christ, Bull. Soc. Bot. Geneve II, 1:221. 1909. TYPE: Carrillo,
Pcia. S. Jose, Werckle (P).

Plants epiphytic, at 700 - 1300 m elevation, in forests, from the Atlantic slopes of
the Cordillera Central, the northern end of the Cordillera de Talamanca,

FIGS. 357 - 363. Pseudocolysis and Microgramma. FIG. 357. Plant of P. bradeorum, Mickel 3484.
FIG. 358. Plant of M. brunei, Correa & Dressier 176. FIG. 359. Plant of M. lycopodioides, Mickel
2341. FIG. 360. Plant of M. nitida, Morley 790. FIG. 361. Plant of M. persicariifolia. Stern et al. 231.
FIG. 362. Plant of M. reptans, Schubert 1323. FIG. 363. Plant of M. tecta var. nana, Holm & litis 193.

250 POLYPODIACEAE: 48. MICROGRAMMA

northeast of the Fortuna Camp reservoir site (Pda. Chiriqm), north of El Valle,
Cerro Jefe and Cerro Campana (Pda. Panama), and the Tutunendo-El Carmen
road near Camp 12 (Depto. Choco). Also from other localities in Colombia.

359. Microgramma lycopodioides (L.) Copel. Gen. Fil. 185. 1947.

Polypodium lycopodioides L. Sp. PI. 2:1082. 1753. TYPE: Lx)cality and collector unknown (LINN
1251.2 not seen microfiche S. I. Library).

Polypodium salicifolium Willd. Sp. PI. ed. 4, 5:149. 1810, non Vahl, 1807, nom. illeg. TYPE:
Brazil, Hojfmannsegg (B-Hb. Willd. 19603 not seen Tryon photo).

Polypodium venosum Desv. Ges. Naturf. Freunde Berlin Mag. 5:314. 1811, non Lour., 1790, nom.
illeg. TYPE: French West Indies, collector unknown (P-Hb. Desv. not seen; probable isotype P-Hb.
Juss. 1061 not seen photo 2924). Synonymized by Weatherby (Contr. Gray Herb. 114:33. 1936).

Polypodium pellitum Willd. ex Kaulf. Enum. Fil. 89. 1824. TYPE: Brazil, Commerson (P-Hb.
Willd. 19604 not seen Tryon photo). Synonymized by Morton (Contr. U. S. Natl. Herb. 38:64. 1967).

Polypodium salicinum Wikstr. Kongl. Vetensk. Acad. Handl. II, 12:443. 1826. TYPE: A

renaming of P. salicifolium Willd., and so based on the type of that name.

Polypodium funiculosum Desv. Mem. Soc. Linn. Paris 6:226. 1827. TYPE: A renaming of P.
lycopodioides sensu Poir. (Enc. Meth. 5:509. 1804), syn. excl., and so based on the type of that name,
which is Hispaniola, collector unknown (P not seen).

Polypodium venulosum Desv. Mem. Soc. Liim. Paris 6:226. 1827. TYPE: A renaming of P.
venosum Desv., and so based on the type of that name.

Polypodium lycopodioides var. subdimorphum Christ in Dur. & Pitt. Bull. Soc. Bot. Belgique 35,
Mem. 233. 1896. TYPE: Alto del Roble, Pcia. Heredia, 1700 m, Pittier 11 in 1888 (G-Hb. Boissier
not seen).

Phymatodes prominula Maxon, Contr. U. S. Natl. Herb. 10:501. 1908. TYPE: S. Juan trail, Isla de
Margarita, Edo. Nueva Esparta, Venezuela, 500 Johnston 155 (US).

Polypodium lycopodioides var. longipes Hassl. Trab. Inst. Bot. Farm. Buenos Aires 45:78. 1928.
TYPE: Cordillera Central, Paraguay, Hassler 6733 (G not seen).

Polypodium lycopodioides var. longipes Domin, Rozpr. Kr^. Ceske Spolecn. Nauk, Tf. Mat.-Prfr. 2
[Pterid. Dominica]: 137. 1929, non Hassl., 1928, nom. illeg. TYPE: Near Barahona, Hispaniola, von
Tuerckheim 2736 (PRC? not seen).

Polypodium lycopodioides f. obtusum Domin, Rozpr. Krai. Ceske Spolecn. Nauk, Tr. Mat.-Prfr. 2
[Pterid. Dominica]: 137. 1929, as ''obtusaP TYPE: Trinidad, Fendler 50 in 1877-1880 (PRC? not
seen; isotype US).

Polypodium lycopodioides var. stipitatum Bosco, Nuovo Giorn. Bot. Ital. N.S., 45:151. 1938.

TYPE: Plan de Sapote, Pcia. Santiago-Zamora, Ecuador, 2100 m, Crespi (TO not seen).

Plants epiphytic, at 0-2500 m elevation, in forests, from throughout the wetter
portions of Costa Rica, Panama, and the Choco. Also from Puerto Rico, Mexico
to Honduras, Nicaragua, Trinidad, Tobago, and tropical South America.

Gomez (Brenesia 6:50-51, f. 3. 1975) has found a hybrid apparently between
this species a.nd Polypodium fuscopetiolatum, which he has mmtdM. moraviana.

360. Microgramma nitida (J. Smith ex Hook.) A. R. Smith, Proc. Calif. Acad. Sci.
IV, 40:230. 1975.

Phlebodium nitidum J. Smith, Bot. Mag. (Curtis) 72, Comp.: 13. 1846. NEOTYPE: '‘‘Collect. Hort.
Kew, 185ip in the hand of J. Smith (BM not seen), chosen by A. R. Smith (FI. Chiapas 2:149. 1981).

Polypodium palmeri Maxon, Contr. U. S. Natl. Herb. 17:600. 1916. TYPE: Near G6mez Farias,
Edo. Tamaulipas, Mexico, ca. 350 m. Palmer 308 (US; isotype UC).

Plants epiphytic, at 0-100 m elevation, in forests, from the Atlantic coastal
plain of Costa Rica near Limon (Pcia. Limon), the Canal Zone, and adjacent Pcia.
Colon. Also from Jamaica, Barbados, Mexico to Honduras, and Nicaragua.

POLYPODIACEAE: 49. NEURODIUM 251

361. Microgramma persicariifolia (Schrad.) K. Presl, Tent. Pterid. 214. 1836.
Polypodium persicariifolium Schrad. Goett. Gel. Anz. 1824:867. 1824. TYPE: Brazil, Maxmillian

von Wied-Neuwied (M not seen).

Plants epiphytic, at 0-100 m elevation, in forests, from the Com. S. Bias, the
Pcia. del Darien, and throughout the Choco. Also from Venezuela, Colombia,
Peru, Bolivia, Guyana, Surinam, and Brazil.

362. Microgramma reptans (Cav.) A. R. Smith, Proc. Calif. Acad. Sci. IV, 40:230.
1975.

Acrostichum reptans Cav. Anales Hist. Nat. 1:104. 1799. TYPE: Guayaquil, Pcia. Guayas,

Ecuador, Nee (MA not seen). Synonymized by Christensen (Dansk Bot. Ark. 9(3) :9. 1937).

Polypodium ciliatum Willd. Sp. PI. ed. 4, 5:144. 1810. TYPE: Est. Para, Brazil, Hojfmannsegg (B-
Hb. Willd. 19601 not seen Tryon photo).

Polypodium cajanense Desv. Ges. Naturf. Freunde Berlin Mag. 5:314. 1811. TYPE: French
Guiana, collector unknown (P-Hb. Desv. not seen; isotype FI not seen photo 16022).

Craspedaria lanceolata Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 7]:65. 1854.
TYPE: Surinam, Hostmann 324 (RB not seen).

Polypodium piloselloides var. moniliforme Hook. Sp. Fil. 5:34. 1863, as '' moniliformis P TYPE:
Not stated.

Plants epiphytic, at 0-200(1000) m elevation, in forests, from the Atlantic
coastal plain of Costa Rica and Panama, Golfito (Pcia. Puntarenas) and the
Peninsula de Osa, the Canal Zone, Cana, and the Choco. Also from Mexico to
Honduras, Nicaragua, Trinidad, Tobago, and tropical South America.

363. Microgramma tecta var. nana (Liebm.) Mickel & Beitel, Pterid. FI. Oaxaca
251. 1988.

Acrostichum nanum Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Aid., V, 1:171
(repr. 19). 1849. TYPE: Between S. Miguel Coatlan and La Galera, Edo. Oaxaca, 7000 - 8000 ft,
Liebmann PL Mex. 2655, FI. Mex. 36 (C not seen).

Polypodium blandulum Christ, Bull. Herb. Boissier II, 7:259. 1907. TYPE: Rio Grande, Pcia. S.
Jose, 600 m, Werckle in 1906 (P not seen).

Plants epiphytic, at 0-1300 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, Finca La Selva (Pcia. Heredia), the Meseta
Central, the vicinity of S. Isidro del General, and the Peninsula de Osa. Also from
Mexico, Honduras, Venezuela to Bolivia and adjacent Brazil, and Surinam.

This differs from M tecta (Kaulf.) Alston var. tecta of southeastern Brazil in
having narrower, hairlike scales on the abaxial surface of the fertile laminae.

49. NEURODIUM Fee

Plants epiphytic; rhizomes short-creeping, bearing copious roots provided with
dark brown root hairs, scaly, the scales lanceolate, strongly clathrate, bicolorous,
toothed, the pale margins worn away in age; stipes short, subterete, blackish,
articulate to the rhizome, slightly alate at the apex; fronds medium-sized, simple,
subdimorphic (the fertile fronds contracted laterally at and proximal to the apex);
laminae narrowly rhombic, coriaceous, shiny, glabrous, the adaxial surface
wrinkled when dry, the midribs greenish abaxially; veins irregularly anastomosing
with simple or branched and recurved included veinlets; sori in a continuous,
submarginal coenosorus, partially protected by the reflexed lamina margin,
exindusiate; paraphyses highly branched filaments; sporangia long-stalked.

Caribbean; monotypic.

252 POLYPODIACEAE: 49. NEURODIUM

367 Niphidium nidulare

368 H. anetioides

POLYPODIACEAE: 50. DICRANOGLOSSUM-51. NIPHIDIUM 253

CHRISTENSEN, C. 1929. Taxonomic fern-studies I. Revision of the polypodioid genera with
longitudinal coenosori... Dansk Bot. Ark. 6(3):1-102, 1. 1 -XIII.

364. Neurodium lanceolatum (L.) F6e, Hist. Vittar. Pleurogr. [Mdm. Foug. 3]:28.

1852.

Pteris lanceolata L. Sp. PI. 2:1073. 1753. TYPE: Plate 40 of Plumier’s “Description...”, which
depicts a plant collected by Plumier on Hispaniola.

Plants epiphytic, at 0 - 100 m elevation, at forest margins, along roadsides, and
in pastures, from the slopes of Volcan Sta. Maria, the Rio Flores, and near
Guayabo (all Pcia. Guanacaste). Also from Florida, the Antilles, Mexico to
Honduras, and French Guiana.

50. DICRANOGLOSSUM J. Smith

Plants epiphytic; rhizomes short-creeping, bearing copious roots provided with
brown root hairs, scaly, the scales lanceolate, strongly clathrate, bicolorous, the
margins worn away in age; stipes articulate to the rhizome, obsolete, narrowly
alate to the base; fronds medium-sized, monomorphic; laminae oblanceolate in
outline, pseudodichotomously forked a few times, attenuate at the base, the
segments long, narrow and attenuate at the apex, herbaceous, glabrous or bearing
small, lanceolate scales abaxially, the midribs prominulous abaxially; veins
obscure, simple, forked, or anastomosing without included veinlets; sori in a
continuous or discontinuous submarginal coenosorus or discrete, partially
protected by the reflexed lamina margin, exindusiate; paraphyses absent;
sporangia long-stalked.

Neotropical; 6 species.

CHRISTENSEN, C. 1929. Taxonomic fern-studies I. Revision of the polypodioid genera with
longitudinal coenosori... Dansk Bot. Ark. 6(3):1- 102, 1. 1 -XIII.

365. Dicranoglossum panamense (C. Chr.) Gdmez, Brenesia 8:46. 1976.

Eschatoff-amme panamensis C. Chr. Dansk Bot. Ark. 6(3) :37. 1929. TYPE: Around the Agua
Clara Reservoir near Gatun, Canal Zone, Maxon 4642 (US; isotype NY).

Plants epiphytic, at 0-200(900) m elevation, in forests and occasionally
plantations, from the Atlantic and Pacific coastal plains of Costa Rica and
Panama, the valley of the Rio General, and throughout eastern Panama and the
Choco. Also from Honduras, Nicaragua, and Colombia to Peru.

51. NIPHIDIUM J. Smith

Plants epiphytic or rarely epipetric or terrestrial; rhizomes short-creeping or
sometimes long-creeping, the younger portions bearing copious roots provided
with brown root hairs, scaly, the scales lanceolate-acuminate, concolorous to
sharply bicolorous, the margins often worn away in age; stipes short, articulate to
the rhizome or to phyllopodia, scaly at the base; fronds medium-sized to large,
simple, monomorphic; laminae narrowly elliptic-lanceolate to oblong, attenuate at
the base, round, acute, or acuminate at the apex, coriaceous, glabrous, or with a
few scales on the abaxial surface of the midrib, often glaucous, with usually

FIGS. 364 - 368. Neurodium, Dicranoglossum, Niphidium, and Hyalotrichopteris. FIG. 364. Plant of
Neurodium lanceolatum, Johnson 1038, Guatemala. FIG. 365. Plant of D. panamense, Mickel 3474.
FIG. 366. Plant of N. crassifolium, Biolley 40. FIG. 367. Plant of N nidulare, Torres R. 42. FIG. 368.
Plant of H. anetioides, Skutch 2753.

254 POLYPODIACEAE: 52. HYALOTRICHOPTERIS

conspicuous hydathodes; principal side veins prominulous, ascending, the
secondary veins immersed, forming inconspicuous, irregular, transverse areolae
with numerous minor areolae containing numerous free veinlets; sori in 1 row
between the main veins, exindusiate; sporangiasters sometimes numerous;
sporangia long-stalked, the capsules sometimes setose.

Neotropical; 10 species.

LELLINGER, D. B. 1972. A revision of the fern genus Niphidium. Amer. Fern J. 62:101-120.

1. Rhizome scales grayish in mass, entire or shghtly repand, sometimes shghtly erose in age, the
cells mostly isodiametric or fusiform, not contorted, those of the central band (2)3-5 times longer
than wide. Laminae narrowly oblanceolate, sometimes narrowly lanceolate, attenuate or acuminate
at the base, acute to round and sometimes cuspidate at the apex, (30)45 - 80(105) cm long, 6-11(18)
cm wide, with brown hydathodes on the adaxial surface; paraphyses common among the sporangia.

366. N. crassifolium

1. Rhizome scales blackish in mass, shallowly toothed, the cells mostly isodiametric, slightly
contorted, those of the central band 1.5-3 times longer than wide. Laminae narrowly lanceolate,
attenuate, obtuse, or round at the base, acute at the apex, (25)55-105 cm long, 3 -6.5(9) cm wide,
with inconspicuous hydathodes on the adaxial surface; paraphyses rare among the sporangia.

367. N. nidulare

366. Niphidium crassifolium (L.) Lellinger, Amer. Fern J. 62:106. 1972.

Polypodium crassifolium L. Sp. PI. 2:1083. 1753. LECTOTYPE: Plate 6, f. 1 of Petiver’s

“Pterigraphia Americana,” chosen by Lelhnger (Amer. Fern J. 62:106. 1972).

Polypodium porrectum Willd. Sp. PI. ed. 4, 5:161. 1810. TYPE: Caracas, Distr. Fed., Venezuela,
Bredemeyer (B-Hb. Willd. 19622 not seen Tryon photo fragm US; isotype W).

Polypodium coriaceum Raddi, Opusc. Sci. 3:286. 1819; PI. Bras. Nov. Gen. 1:16, t. 25. 1825, non
Swartz, 1788, nom. illeg. TYPE: Corco-secco, Est. Rio de Janeiro, Brazil, Raddi (FI, labelled
“Polyp: coriaceum nob: e Brasilia”).

Pleuridium angustum Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:98. 1857.

TYPE: “Province d’Ocana,” Depto. Norte de Santander, Colombia, ca. 2400 m, Schlim 610 (BR;
isotypes HB, L, W).

Plants epiphytic, occasionally epipetric or terrestrial, at 0 - 1600 m elevation, in
forests, from throughout the Flora area. Also from throughout tropical America.

367. Niphidium nidulare (Rosenst.) Lellinger, Amer. Fern J. 62:112. 1972.

Polypodium crassifolium var. nidulare Rosenst. Repert. Spec. Nov. Regni Veg. 22:18. 1925.

TYPE: La Palma, Pda. S. Jose, 1400 m, Brade & Brade 113 (Ros. Fil Costar. Exs. 300) (S photos
5969, 20635; isotypes B, NY, UC, US).

Plants epiphytic, at 1300-2000 m elevation, in forests, from Monteverde,
Zarcero (Pcia. Alajuela), La Palma (Pcia. S. Jose), and La Fortuna and Estrella
(Pcia. Cartago).

52. HYALOTRICHOPTERIS W. H. Wagner

Plants epipetric; rhizomes creeping, branched, densely scaly, the scales
lanceolate, strongly clathrate, concolorous; stipes obsolete; fronds small or minute,
monomorphic, not articulate; laminae spathulate, herbaceous, sparsely hairy, the
hairs pluricellular, filiform, with a short basal cell also bearing a short glandular
cell; veins anastomosing, forming 1(2) rows of roundish, polygonal areolae, with
short, free veins near the margin and a single included veinlet in each areola;
hydathodes present on the adaxial surface of the lamina; sori round, exindusiate;
paraphyses present; sporangia long-stalked.

Nicaragua and Costa Rica; monotypic.

POLYPODIACEAE: 53. CAMPYLONEURUM 255

WAGNER, W. H., Jr. and D. R. FARRAR. 1977. The Central American fern genus Hyalotricha
and its family relationships. Syst. Bot. 1:348-362.

368. Hyalotrichopteris anetioides (Christ) Wagner, Taxon 27:548. 1978.

Pofypodium anetioides Christ, Bull. Soc. Bot. Geneve II, 1:219. 1909. TYPE: Candelaria

Mountains, Pcia. S. Jose, 1450 m, Brade & Brade 177 (Ros. Fit Costar. Exs. 88) (P not seen; isotypes
GH, MICH? not seen, NY, S not seen photo 5941, DC).

Plants epipetric, at 1000 -2000 m elevation, on cliffs in dense forests, from the
Fila de Cedral, Orosi (Pcia. Cartago), and the Cordillera de Talamanca to Pcia.
Chiriqui. Also from Nicaragua.

S3. CAMPYLONEURUM K. PresI

Plants epiphytic or occasionally epipetric or terrestrial; rhizomes short-creeping
and robust to long-creeping and wiry, often glaucous, bearing rather few, long
roots without conspicuous root hairs, scaly especially at the apex, the scales
peltate-lanceolate, sometimes clathrate, usually brownish; fronds small to large,
monomorphic; stipes short to sometimes long, articulate to the rhizome or to short
phyllopodia, scaly at the base; laminae simple (pinnate in a few species), oblong to
elliptic, herbaceous to coriaceous, glabrous or with a few scales on the abaxial
surface of the midrib, often shiny, with usually conspicuous hydathodes; principal
side veins parallel, prominulous in most species, ascending, the secondary veins
immersed or not, forming rather regular, transverse areolae, sometimes with 2(3
or more) minor areolae usually containing a single, unbranched included veinlet,
or parallel side veins absent and the veins uniform and anastomosing with 1 or 2
sori in each fertile areola; sori (1)2 in major areolae, usually forming double rows
between the main veins, exindusiate; paraphyses absent; sporangia long-stalked.

Neotropical; ca. 50 species.

LELLINGER, D. B. 1988. Some new species of Campyloneurum and a provisional key to the genus.

Amer. Fern J. 78:14-35.

1, Rhizomes compact to short-creeping, rarely long-creeping, 3-10 mm in diam.; stipes 0.2- 1(2)
cm distant. .6.

1. Rhizomes very long- creeping, 1-2(3) mm in diam.; stipes usually 1-4 cm distant. .2.

2(1). Fertile areolae unisoriate with the fertile veins not branching from the midpoint of the
transverse vein, occasionally bisoriate. Rhizome scales lanceolate, entire, clathrate with narrow
lumina; stipes 1-5.5 cm long, 0.25-0.75 mm in diam.; laminae linear-lanceolate, 6-16 cm long,
5-18 mm wide, acute to cuneate at the base, acuminate at the apex; sori in 2 or 3 series on each side
of the midrib.

374. C falcoideum

2(1). Fertile areolae 2(3)-soriate..3.

3(2). Adaxial surface of the laminae very shiny; laminae elliptic, 2-3 times longer than wide;
fronds 5-20(25) cm long. Rhizome scales lanceolate, 3-5 mm long, with slightly toothed margins,
golden brown, spreading; stipes 2-10 cm long, 0.25-1 mm in diam.; laminae elliptic, 3.5-15 cm
long, 1.5 -5.5 cm wide, acuminate at the base and apex; sori in 4-6 series on each side of the midrib.

382. C sublucidum

3(2). Adaxial surface of the laminae dull; laminae linear to long-elliptic, at least 3 times longer
than wide; fronds 12-50 cm long..4.

4(3). Stipes 1-3 cm long; laminae elliptic-oblong, (2)2.5 -3.5(5) cm wide, acuminate at the base.
Stipes 1-1.5 mm in diam.; sori in 4-5(6) series on each side of the midrib.

381. C serpentinum

256 POLYPODIACEAE: 53. CAMPYLONEURUM

4(3). Stipes 2-12(18) cm long; laminae elliptic to elliptic-oblong, (2)3-8(11) cm wide, acute to
attenuate at the base..5.

5(4). Stipes (3)4-12(18) cm long; laminae acute, cuneate, or attenuate at the base, (1)2- 3(6)
times longer than the stipes, coriaceous. Stipes ca. 1 mm wide; laminae elliptic-lanceolate, eUiptic-
oblong, or elliptic, 10 - 30(40) cm long, (1.5)2-8(11.5) cm wide, acuminate to caudate at the apex;
sori in 4-8(15) series on each side of the midrib.

383. C. wercklei

5(4). Stipes 2-5(7) cm long; laminae cuneate to attenuate at the base, (4)6-8(12) times longer
than the stipes, papyraceous. Stipes ca. 1 mm wide; laminae elliptic-oblong, (12)15-40(50) cm long,
(2.5)4 -8.5 cm wide, acute to acuminate at the apex; sori in 4- 13 series on each side of the midrib.

380. C. repens

6(1). Laminae not linear, more than 3 cm wide.. 10.

6(1). Laminae linear or nearly so, less than 3(4.5) cm wide..?.

7(6). Rhizome scale cells predominantly 2-4 times longer than wide. .9.

7(6). Rhizome scale cells predominantly 1-2 times longer than wide.. 8.

8(7). Cells of the rhizome scales at least in part contorted. Stipes 2-4 cm long, 1-1.5 mm wide
distal to the base; laminae linear, 10-30 cm long, (3)8-10(14) cm wide, shghtly revolute, attenuate
at the base and apex; venation mostly obscure abaxially; sori in 1(2) series on each side of the midrib.

374a. C. irre^lare

8(7). Cells of the rhizome scales not contorted. Stipes 1-5 cm long, 1-1.5 mm wide distal to the
base; laminae linear, 30 - 45 cm long, 9-18 mm wide, often revolute, attenuate at the base and apex;
venation mostly obscure abaxially; sori in (1)2 series on each side of the midrib.

372. C. cooperi

9(7). Stipes 1- 1.5(2) mm in diam. near the base, 0-5(7) cm long; laminae (0.5)l-2.5 cm wide;
sori usually less than 2 mm in diam. Laminae linear, (10)20 - 45(60) cm long, attenuate at the base
and apex; venation usually entirely obscure abaxially; sori in 1 - 3 series on each side of the midrib.

370. C. angustifolium

9(7). Stipes 2-4(5) mm in diam. near the base, (5)10-15(20) cm long; laminae (1)2.5 -4(5) cm
wide; sori usually more than 2 mm in diam. Laminae linear, 15 - 45(60) cm long, cuneate to long-
attenuate at the base, acuminate to attenuate at the apex; venation usually visible, often prominulous;
sori in 2-3(4) series on each side of the midrib.

369. C. amphostenon

10(6). Laminae pinnate. Stipes 0.3- 1(1.5) m long; laminae ovate-lanceolate, ca. 0.5 -1(1.5) m
long, 40 - 80 cm wide; pinnae elliptic-lanceolate, 20 - 40 cm long, 6-10 cm wide, usually with 3
alternate, lateral pinna pairs plus a conform terminal pinna.

376. C magnificum

10(6). Laminae simple.. 11.

11(10). Primary and transverse veins all hidden; stipes 1-2 cm long, 1-2 mm in diam.; laminae
narrowly oblanceolate. Laminae (6)10 - 30(35) cm long, 1.5-5 cm wide, attenuate at the base,
apiculate at the apex; transverse veins arcuate with 2 included, acroscopic veinlets; sori in 3 - 5 series
on each side of the midrib.

378. C. occultum

11(10). Primary and transverse veins mostly visible (transverse veins mostly obscure in C. costatum
and xalapense); stipes (1)2-30 cm long, 1.5-4 mm in diam.; laminae not narrowly oblanceolate.. 12.

12(11). Fertile transverse veins always with only 2 free, acroscopic, included veinlets, both of them
fertile. Laminae 30 - 45 cm long, 9 - 13 cm wide, elliptic, acute-attenuate at the base, apiculate at the
apex; sori in ca. 12 series on each side of the midrib.

371. C. brevifolium

12(11). Fertile transverse veins with 3 or more free, acroscopic, included veinlets or the venation
irregular.. 13.

13(12). Transverse veins mostly visible on the abaxial surface.. 15.

POLYPODIACEAE: 53. CAMPYLONEURUM 257

13(12). Transverse veins mostly not visible on the abaxial surface.. 14.

14(13). Stipes 1-4(5) cm long; laminae usually with obvious lime dots on the adaxial surface.
Ltiminae oblong, 15-40(45) cm long, 2 -4(5.5) cm wide, attenuate at the base, apiculate at the apex.

384. C xalapense

14(13). Stipes (2)4-14 cm long; laminae lacking lime dots on the ad2ixicd surface. Laminae
oblong-elliptic, 15-30(35) cm long, 2-6 cm wide, attenuate to cuneate at the base, apiculate at the
apex.

373. C. costatum

15(13). Laminae 2 -5(6.5) cm wide..l7.

15(13). Laminae 5- 14 cm wide.. 16.

16(15). Venation regular, the fertile transverse veins usually with 3 free, acroscopic, included
veinlets, the lateral 2 fertile, the middle one sterile and prolonged, sometimes forming a pair of
areolae; sori in 2(3) regular rows between the lateral veins. Stipes 6-14(23) cm long; laminae
narrowly elliptic to oblong, (20)35-75 cm long, 5- 11 cm wide, cuneate at the base, round to acute at
the apex; midrib scales dark brown, up to 1.5 mm long, deciduous; lateral veins 5-6 mm distant.

379. C. phyllitidis

16(15). Venation irregular with the primciry areolae usually divided by several irregular veinlets not
parallel to the primary veins; sori in 2-4 irregular rows between the primary veins. Stipes 10-23 cm
long; laminae oblong, (30)45 - 90 cm long, 6 - 14 cm wide, cuneate at the base, acute to subapiculate
at the apex; midrib scales pale brown, 1-3 mm long, subpersistent; primary veins (4)5-10 mm
distant.

375. C. latum

17(15). Laminae dull, plane and entire at the margin; stipes 4-15 cm long; rhizomes ca. 5 mm in
diam. L2uninae linear to oblong, 30 - 50 cm long, 3 -5(6.5) cm wide, attenuate at the base, acute at
the apex, with weak lime dots on the adaxial surface; primary veins 4-5 mm distant.

377. C. multipunctatum

17(15). Laminae shiny, somewhat wrinkled and irregularly crenate at the margin; stipes 1-4(5) cm
long; rhizomes ca. 3 mm in diam. Laminae oblong, 15-40(45) cm long, 2 -4(5.5) cm wide, apiculate
at the apex, with obvious lime dots on the adaxial surface; primary veins 5-6 mm distant.

384. C. xalapense

369. Campyloneurum amphostenon (Kunze ex Klotzsch) F^e, Gen. Fil. [M^m.

Foug.5]:258. 1852.

IPolypodium dimorphum Link, Hort. Reg. Bot. Berol. 2:88. 1833. TYPE: Not stated; presumably
a specimen collected by Link from a plant cultivated at the Botanical Garden in Berlin, originally
from tropical America (B? not seen).

Polypodium amphostenon Kunze ex Klotzsch, Linnaea 20:399. 1847. TYPE: Merida, Edo.

Merida, Venezuela, Moritz 120b (B not seen; isotype BM not seen fragm and Maxon photo 121 US,
K not seen).

Polypodium angustifolium var. monstrosum Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:258
(repr. 340). 1864, as ''monstrosaP TYPE: Cipac6n, Depto. Cundinamarca, Colombia, 2800 m,
Lindig241 (B not seen; isotype US).

Polypodium pittieri Christ in Pitt. Prim. FI. Costaric. 3(1):16. 1901. TYPE: El Paramo toward the
east of Cerro Buena Vista, Pcia. S. Jose, 3000 m, Pittier 10479 (BR; isotypes BM not seen photo GH,
US).

Plants epiphytic or terrestrial, at 2000-3100 m elevation, in forests and in open
areas, from the Cordillera Central and the Cordillera de Talamanca to Pcia.
Chiriquf. Also from Cuba, Jamaica, Hispaniola, Mexico, Guatemala, Venezuela,
and Colombia to Bolivia.

258 POLYPODIACEAE: 53. CAMPYLONEURUM

POLYPODIACEAE: 53. CAMPYLONEURUM 259

370. Campyloneumm angustifolium (Swartz) F^e, Gen. Fil. [M^m. Foug. 5]:257.
1852.

Polypodium angustifolium Swartz, Nov. Gen. Sp. PI. Prodr. 130. 1788. TYPE: Jamaica, Swartz (S
not seen Maxon photo 101; isotypes B-Hb. Willd. l%ll-2 not seen Tryon photo, BM not seen photo
6729).

IPolypodium calaguala Ruiz, Mem. Calaguala c. tab. 1805. Reference not seen; copy at G.
Polypodium crassifolium f. angustissimum Rosenst. Mem. Soc. Sci. Nat. Neuchatel 5:45. 1912, as
“angustissima.” TYPE: Above Facatativa, Savane de Bogota, Depto. Cundincunarca, Colombia, 2600
m. Mayor 40 (S not seen; isotype US).

Plants epiphytic, at (100)500-3300 m elevation, in forests, pastures, and on
roadsides, from the Cordillera de Tilaran, the Cordillera Central, Cerro
Carpintera, the Cordillera de Talamanca to Pcia. Chiriqui, the Fila Costeha near
S. Vito, near Sta. Fe, near El Valle, and the upper Rfo Truando (Depto. Choco).
Also throughout tropical America.

371. Campyloneumm brevifolium (Lodd. ex Link) Link, Fil. Sp. 124. 1841.

Polypodium brevifolium Lodd. ex Link, Hort. Reg. Bot. Berol. 2:90. 1833. TYPE: Origin

unknown; cultivated by Loddiges, collector unknown (B? not seen).

Polypodium caespitosum Lodd. ex Link, Hort. Reg. Bot. Berol. 2:91. 1833. TYPE: Origin

unknown; cultivated by Loddiges, collector unknown (B? not seen).

Plants epiphytic or terrestrial, at 600-1400 m elevation, in forests, from the
Reventazon and Navarro valleys (both Pcia. Cartago), the Fila Costeha near S.
Vito, the vicinity of Boquete, and Cana and the Serrania del Darien between
Cerro Mali and the Rfo Pucro (Pcia. Darien). Also from Venezuela, Colombia,
and Peru.

372. Campyloneumm cooperi Lellinger, Amer. Fern J. 78:19, f. 3, 9. 1988.

TYPE: Cartago, Pcia. Cartago, 4250 ft. Cooper 6053 (US; isotype US).

Plants epiphytic, at 900 - 1300 m elevation, in forests, known only from the type
and from Turrialba, Pcia. Cartago (Werckle, US).

373. Campyloneumm costatum (Kunze) K. Presl, Tent. Pterid. 190. 1836.

Polypodium costatum Kunze, Linnaea 9:38. 1834. TYPE: Limonar, Cuba, Aug 1822, Poeppig (LZ
destroyed; isotypes BM not seen, K not seen, L not seen photo 1853).

Campyloneumm immersum J. Smith in Seem. Bot. Voy. Herald 231. 1854. TYPE: Bay of Utria,
Depto. Choco, Seemann 61 (K not seen Maxon photo 246).

Plants epiphytic, at 0-200 m elevation, in forests, from the Atlantic coastal
plain of Panama, near El Valle, and the Canal Zone and adjacent Pcia. Panama.
Also from Florida, Cuba, Jamaica, Hispaniola, Puerto Rico, Mexico to Belize,
Nicaragua; Colombia and Venezuela to Trinidad, the Guianas, and central Brazil.

This species resembles C. occultum in having partially obscure venation, but
differs from that species in having longer stipes and in lacking minute hairs on the
abaxial surface of the laminae.

FIGS. 369-376. Campyloneumm. FIG. 369. Plant of C. amphostenon, Evans & Lellinger 92. FIG.
370. Plant of C. angustifolium, L2mkester 647. FIG. 371. Plant of C. brevifolium, Mickel 2105. FIG.
372. Plant of C. cooperi, Werckle. FIG. 373. Plant of C. costatum, Maxon 4872. FIG. 374. Plant and
portion of lamina of C. falcoideum. Stork 4841 and Seibert 284. FIG. 375. Plant and portion of
lamina of C. latum. Cook & Doyle 287. FIG. 376. Base of median pinna of C magnificum,
Cuatrecasas 13667, Colombia.

260 POLYPODIACEAE: 53. CAMPYLONEURUM

374. Campyloneurum falcoideum (Kuhn ex Hieron.) M. Meyer ex Lellinger, Proc.
Biol. Soc. Wash. 89:708. 1977.

Polypodium falcoideum Kuhn ex Hieron. Bot. Jahrb. Syst. 34:533. 1904. SYNTYPES: Rio Sucio,
Costa Rica, 800 m, Lehmann 1741 (B not seen; isosyntype US); and near Desengano, Pcia. Heredia?,
Wendland 876 (B not seen).

IPolypodium rodriguezianum Gomez, Revista Biol. Trop. 17:107, f. 5-6. 1970. TYPE: Cerro
Carpintera, Pcia. Cartago, 1800 m, Gomez PtC-2063 (CR not seen).

Plants epiphytic, at 0-2000 m elevation, in forests, from the Atlantic plain of
Costa Rica, the Cordillera Central, Cerro Tablazo, Cerro Carpintera, and the
Cordillera de Talamanca to Pcia. Chiriqui.

An earlier epithet may be available from Polypodium schnittspahnii Christ (Bull.
Herb. Boissier 6:836. 1898), which is based on “Andes, MoritzT' (presumably P
not seen).

374a. Campyloneurum irregulare Lellinger, Amer. Fern J. 78:24, f. 6, 12. 1988.

TYPE: Vicinity of Quito, Pcia. Pichincha, Ecuador, 3000 m, Holdridge 1580
(US).

Plants epiphtytic or occasionally epipetric, at 800-1500 and 3200-3300 m
elevation, in forests, from near S. Ramon, Angel Falls (Pcia. Alajuela), Cartago,
and the Cordillera de Talamanca, especially in Pcia. Chiriqui. Also from
Venezuela and Colombia to Bolivia.

This species resembles C. angustifolium in lamina size and shape, but differs in
having rhizome scales with contorted cells.

375. Campyloneurum latum T. Moore, Ind. Fil. 225. 1861.

ICampyloneumm moritzianum Fee, Gen. Fil. [Mem. Foug. 5]:258. 1852. TYPE: Caracas, Distr.
Fed., Venezuela, Moritz 3 (RB not seen).

LECTOTYPE: St. Vincent, Guilding (K not seen), chosen by Proctor (FI. Less.
Antill.342. 1977).

Plants epiphytic, at 0- 1400 m elevation, in forests, from the Cordillera Central,
the Atlantic coastal plain of Costa Rica and Panama, the Cordillera de Talamanca
to Pcia. Chiriqui, around El Valle and Penonome (both Pcia. Code), Chagres
(Canal Zone), the Rio Fato (Pcia. Colon), and Loma del Cuchillo and Andagoya
(both Depto. Choco). Also throughout tropical America except the Guianas.

The epithet moritzianum will displace the well-known epithet latum if the two
species prove to be synonymous.

376. Campyloneurum magnificum T. Moore, Ind. Fil. 226. 1861.

Polypodium fendleri D. C. Eaton, Mem. Amer. Acad. Arts N.S., 8:199. 1860, non Campyloneurum
fendleri Moore, 1861. TYPE: Colonia Tovar, Edo. Aragua, Venezuela, Fendler 410 (YU not seen;
isotypes E? not seen, MO).

TYPE: Colonia Tovar, Edo. Aragua, Venezuela, Fendler 410 (E? not seen;
isotypes MO, YU not seen).

Plants terrestrial or hemiepiphytic, at 800 - 1200 m elevation, in forests, in the
Flora area known only from the headwaters of the Rio Tuquesa ca. 2 km from the
continental divide, Pcia. Darien {Croat 27143, MO), and from near the Ciudad
Bolivar - Quibdo road at ca. Km. 139, Depto. Choco {Lellinger & de la Sota 916,
LP, US). Also from Venezuela and Colombia to Bolivia.

POLYPODIACEAE: 53. CAMPYLONEURUM 261

377. Campyloneurum multipunctatum (Christ) Lellinger, Proc. Biol. Soc. Wash.
89:708. 1977.

Polypodium phyllitidis var. elongatum Hieron. Bot. Jahrb. Syst. 34:534. 1904, as ''elongata.”

SYNTYPES: Rio Paez, Depto. Tolima, Colombia, 800-1300 m, Lehmann 5721 (B not seen;
isosyntype US); and Cerro Yanghuan near Pindilic, Cordillera Oriental de Cuenca, Pcia. Azuay,
Ecuador, 2500 - 2800 m, Lehmann 7679 (B not seen; isosyntypes GH, US).

Polypodium phyllitidis f. multipunctatum Christ, Bull. Herb. Boissier II, 5:7. 1905. TYPE: Costa
Rica, Werckle 174 (P not seen).

Plants epiphytic, at 600-1300 m elevation, in forests, from the Cordillera
Central and Tuis (Pcia. Cartago). Also from Colombia and Ecuador.

378. Campyloneurum occultum (Christ) G6mez, Brenesia 8:46. 1976.

Polypodium occultum Christ, Bull. Herb. Boissier II, 5:7. 1905. TYPE: Rio de las Vueltas,
Tucurrique, Pcia. Cartago, 365 m, Tonduz 12756 (P not seen photo BM not seen). Tonduz 12752
(US) is also this species from the same locality, and actually may be the type number if the published
number is in error.

Polypodium trichiatum Rosenst. Repert. Spec. Nov. Regni Veg. 7:148. 1909. TYPE: Cordillera
Occidental near Riobamba, Pcia. Chimborazo, Ecuador, 600 m, Rimbach 87 (S not seen photo 6089).

Plants epiphytic or rarely epipetric, at 0 - 600(900) m elevation, in forests, from
the Cordillera de Tilaran, the Atlantic coastal plain of Costa Rica, the valley of
the Rio Reventazon, near Sta. Fe, the Canal Zone and adjacent Pcia. Colon and
Pcia. Panama, near Cana, and near Bahia Solano and Acandi (Depto. Choco).
Also from Belize, Nicaragua, Colombia, Peru, Bolivia, and Brazil.

379. Campyloneurum phyllitidis (L.) K. Presl, Tent. Pterid. 190. 1836.

Polypodium phyllitidis L. Sp. PI. 2:1083. 1753. TYPE: Plate 38 of Plumier’s “Description...”,

which illustrates a plant from the Antilles.

Polypodium comosum L. Sp. PI. 2:1084. 1753. TYPE: Plate 131 of Plumier’s “Traite...”, which
illustrates a plant collected by Plumier on Hispaniola.

Polypodium simplicifolium Salisb. Prodr. Stirp. Chap. AUerton 403. 1796, nom. superfl. TYPE: A
renaming of P. phyllitidis L., and so based on the type of that name.

Polypodium conjugatum Poir. Encyc. Meth. 5:516. 1804. TYPE: West Indies, Houston in 1732 (P-
Hb. Juss. Cat. 1071 not seen photo 2934).

IPolypodium parallelinerve Desv. Mem. Soc. Linn. Paris 6:230. 1827. TYPE LOCALITY:

“Habitat in ins. Africanis?” presumably in error.

Polypodium gladiatum Veil. FI. Flumin. ll:t. 59. 1831; Arq. Mus. Nac. Rio de Janeiro 5:445.
1881, non Kunze, 1834, nom. illeg. TYPE LOCALITY: Brazil.

Plants epiphytic, at 0-700 m elevation, in forests, from near S. Ramon, the
Cordillera Central, the Atlantic coastal plain of Costa Rica and Panama, the
Meseta Central, the Cordillera de Talamanca to Pcia. Chiriquf, near Guaniquito
(Pcia. Los Santos), La Mesa del Valle (Pcia. Code), the Canal Zone and adjacent
Pcia. Panama, near Cana, and the northern half of the Choco. Also throughout
tropical America.

The cross veins are sometimes rather hidden in this species on the abaxial
surface of the laminae, as in C. costatum, but the laminae are relatively wider and
taper more abruptly at the base than in C. costatum.

380. Campyloneurum repens (Aubl.) K. Presl, Tent. Pterid. 189. 1836.

Polypodium repens Aubl Hist. PI Guiane 2:962. 1775. TYPE: Plate 134 of Plumier’s “Traite...”,
which illustrates a specimen collected by Plumier on Martinique.

262 POLYPODIACEAE: 53. CAMPYLONEURUM

379 C. phyllitidis

POLYPODIACEAE: 53. CAMPYLONEURUM 263

IPolypodium nitidum Kaulf. Enum. Fil. 92. 1824. TYPE: Ilha Sta. Catarina, Est. Sta. Catarina,
Brazil, Chamisso (LE not seen).

1 Polypodium crispum Fee, Gen. Fil. [Mem. Foug. 5]:259. 1852. TYPE: Brazil, Martins 303 (P or
RB not seen).

Plants epiphytic, at 0-900(1600) m elevation, in forests, from the Atlantic and
Pacific coastal plains of Costa Rica, the Cordillera Central, the Fila Costeha near
S. Vito, near Puerto Armuelles (Pcia. Chiriqm), Loma Prieta (Pcia. Los Santos),
the vicinity of El Valle, the Canal Zone and adjacent Pcia. Panama, Cana and
Paca (Pcia. Darien), and throughout the Choco. Also from Jamaica, Puerto Rico,
the Lesser Antilles, Mexico, Guatemala to Honduras, Nicaragua, Venezuela,
Colombia to Bolivia, the Guianas, and Brazil.

381. Campyloneurum serpentinum (Christ) Ching, Sunyatsenia 5:263. 1940.

Polypodium serpentinum Christ, Bull. Herb. Boissier 11, 6:51. 1906. TYPE: Navarro, Pcia.

Cartago, Werckle (P not seen).

Plants epiphytic, at 0-1700 m elevation, in forests, from near S. Ramon, the
Cordillera Central, the Meseta Central, the Reventazon and Navarro valleys (Pcia.
Cartago), the Cordillera de Talamanca to Pcia. Chiriqui, near El Valle,
Masarganti and vicinity (Com. S. Bias), Cerro Pirre and between Paya and Palo de
Las Letras (Pcia. Darien), and Loma del Cuchillo and La Teresita (both Depto.
Choco). Also from Mexico, and Venezuela to Peru.

382. Campyloneurum sublucidum (Christ) Ching, Sunyatsenia 5:263. 1940.

Polypodium sublucidum Christ, Bull. Herb. Boissier II, 7:261. 1907. TYPE: La Palma, Pcia. S.
Jose, 1500 m, Werckle 17051 (P not seen).

Plants epiphytic or epipetric, at 900-1400 m elevation, in forests, from La
Hondura (Pcia. S. Jose), Orosi and near Pejivalle (both Pcia. Cartago), and
between Cana and the Alturas de Nique (Pcia. Darien).

383. Campyloneurum wercklei (Christ) Lellinger, Amer. Fern J. 78:28. 1988.
Polypodium wercklei Christ, Bull. Herb. Boissier II, 5:7. 1905. SYNTYPES: Costa Rica, Werckle

(P not seen); and Rio Sucio, Costa Rica, 800 m, Lehmann 1741 (P not seen; isosyntype US).

Plants epiphytic, at 100-2000(2400) m elevation, in forests, from the Cordillera
de Tilaran, the Cordillera Central, the Atlantic coastal plain of Costa Rica and
adjacent Panama, the Cordillera de Talamanca to Pcia. Chiriqui, the Fila Costeha
near S. Vito, El Valle, and the Pacific coastal plain of southern Costa Rica. Also
from Ecuador.

384. Campyloneurum xalapense F6e, Gen. Fil. [M6m. Foug. 5] :258. 1852.

Campyloneurum caudatum Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:96. 1857,
non Raddi, 1819, nom. illeg. TYPE: C6rdoba and Huatusco, Edo. Veracruz, Mexico, Schajfner 176
(RB? not seen; isotype K not seen), synonymized by A. R. Smith (FI. Chiapas 2:68. 1981).

Polypodium weatherbyanum F. Seym. Phytologia 31:171. 1975. TYPE: A renaming of

Campyloneurum caudatum, and so based on the type of that name.

FIGS. 377 - 384. Campyloneurum. FIG. 377. Plant of C. multipunctatum, Tonduz 14223. FIG. 378.
Plant of C. occultum, Kilhp 2659. FIG. 379. Portion of lamina of C. phyllitidis, Wilhams 1028. FIG.
380. Plant of C. repens, Kennedy 1924. FIG. 381. Plant of C. serpentinum, Maxon 665. FIG. 382.
Plant of C sublucidum, Standley & Valerio 47108. FIG. 383. Plant of C. wercklei, LeUinger 1393.
FIG. 384. Plant of C. xalapense, Werckle.

264 POLYPODIACEAE: 54. PHLEBODIUM

TYPE: Jalapa, Edo. Veracruz, Mexico, Galeotti 6723 (P or RB not seen fragm
ex K at NY).

Plants epiphytic, at 500-1200 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, the Meseta Central, the Navarro valley (Pcia.
Cartago), the Fila Costeha, and Barro Colorado Island (Canal Zone). Also from
Mexico to El Salvador.

54. PHLEBODIUM (R. Br.) J. Smith

Plants epiphytic or epipetric; rhizomes creeping, thick, often knotted, densely
scaly, the scales lanceolate, usually reddish-brown, ciliate-toothed; stipes rather
thick, articulate to short phyllopodia, scaly at the base; fronds medium-sized to
large, monomorphic; laminae ovate to oblong, pinnatisect or pinnatifid with a
broadly alate rachis, herbaceous to subcoriaceous, glabrous, often glaucous; veins
anastomosing in polygonal areolae, generally lacking included veinlets, except the
fertile areolae with an included polygonal areola bearing a sorus distally; sori
small, round, exindusiate, lacking paraphyses; sporangia long-stalked.

Neotropical; ca. 4 species.

1. Sori in 1(2) rows on each side of the segment midrib; laminae slightly to copiously glaucous on
the abaxial surface; veins forming a relatively uniform network of areolae, without prominent
primary lateral veins; rhizomes with their scales ca. 1 cm in diam.

385. P. pseudoaureum

1. Sori in 3-6 rows on each side of the segment midrib; laminae not or only shghtly glaucous on
the abaxial surface; primary lateral veins prominent, with less prominent, arcuate secondary veins
forming areolae; rhizomes with their scales ca. 2 cm in diam.

386. P. decumanum

385. Phlebodium pseudoaureum (Cav.) Lellinger, Amer. Fern J. 77:101. 1988.

Polypodium pseudoaureum Cav. Descr. PI. 247. 1801. TYPE; Without locality. Nee (MA not
seen), examined by Christensen (Dansk Bot. Ark. 9(3):12. 1937).

Polypodium sporadocarpum Willd. Sp. PI. ed. 4, 5:171. 1810, nom. superfl. TYPE: A renaming of
Polypodium pseudoaureum Cav., and so based on the type of that name.

Polypodium areolatum Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:172. 1810. TYPE: Cumana,
Edo. Sucre, Venezuela, Humboldt & Bonpland (B-Hb. Willd. 19645 not seen mfc. S. I. Library).

Polypodium pulvinatum Link, Hort. Reg. Bot. Berol. 2:99. 1833. TYPE: Cultivated in the

Botanical Garden at Berlin, originally from Brazil, Link (B not seen).

Chrysopteris trilobata Fee, Gen. Fil. [Mem. Foug. 5]:266. 1852. TYPE: South America, Pamplin (P
or RB not seen).

Chrysopteris grandis Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:96. 1857. TYPE:
Ocana, Depto. Norte de Santander, Colombia, Schlim 598 (P or RB not seen).

Chrysopteris lanosa Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:97. 1857. TYPE:
S. Angel, Mexico, Schajfner 311 (P or RB not seen).

Chrysopteris microdictya Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:97. 1857.
TYPE: Near Orizaba, Edo. Veracmz, Mexico, Schajfner 203 (RB not seen ).

Polypodium auratum Vellozo, FI. Flumin. ll:t. 74. 1831; Arq. Mus. Nac. Rio de Janeiro 5:448.
1881. TYPE: Plate 74, illustrating material collected around Rio de Janeiro, Br^izil, by Vellozo.

Polypodium areolatum var. loreum Bomm. in Dur. & Pitt. Bull. Soc. Roy. Bot. Belgique 35, Mem.
219. 18%. SYNTYPES: Rio Virilla, La Uruca, Pcia. S. Jose, 1100 m, Pittier 8017 (BR not seen),
Biolley 131 (BR not seen); S. Franciso de Guadalupe, Pcia. S. Jose, Tonduz 99 (BR not seen); and
Pico Piedra Blanca, west of Volcan Irazu, Pcia. S. Jose, Tonduz 418bis (BR not seen).

Polypodium aureum var. reductum Jenm. Bull. Bot. Dept. Jamaica N.S., 4:138. 1897. TYPE: Not
stated; probably Jam2uca, /cnman (NY? not seen).

POLYPODIACEAE: 55. POLYPODIUM 265

Plants epiphytic, at 100-2300(2600) m elevation, in forests and pastures, from
the Cordillera de Tilaran, the Atlantic coastal plain of Costa Rica, the Cordillera
Central, the Meseta Central, the Cordillera de Talamanca to Pcia. Chiriqui, near
Sta. Fe, and the vicinity of Cana. Also from Florida, the Antilles, Mexico to
Nicaragua, Venezuela to Bolivia, Brazil, the Guianas, and Argentina.

386. Phlebodium decumanum (Willd.) J. Smith, J. Bot. (Hooker) 4:59. 1841.

Polypodium decumanum Willd. Sp. PI. ed. 4, 5:170. 1810. TYPE: Brazil, Hoffmannsegg (B-Hb.
Willd. 19640 not seen microfiche S. I. Library).

Chrysopteris dictyocallis Fee, Gen. Fil. [Mem. Foug. 5]:265. 1852. TYPE: French Guiana, Poiteau
(P not seen).

Phlebodium multiseriale T. Moore & Houlst. Gard. Chron. 1855:469. 1855. TYPE: Not stated;
from cultivated material.

Plants epiphytic, at 0-200 m elevation, in forests, from Tortuguero (Pcia.
Limon), near Unguia and the Parque Natural Nacional Los Catios (both Depto.
Choco). Also from Jamaica, Hispaniola, Puerto Rico, the Lesser Antilles,
Trinidad, Mexico to Honduras, and tropical South America.

Hybrids of this species with P. pseudoaureum have been reported from
Tortuguero, Pcia. Limon, by L. D. Gomez (pers. comm.).

55. POLYPODIUM L.

Plants epiphytic, sometimes epipetric or rarely terrestrial; rhizomes creeping,
scaly, the scales concolorous or bicolorous, usually not more than weakly clathrate,
mostly tan or brownish; stipes relatively long, thin, articulate to low phyllopodia,
mostly scaly at the base, otherwise glabrous, a few (with the rachises) pilosulous;
fronds small to large, monomorphic or rarely subdimorphic with the fertile
laminae somewhat contracted; laminae pinnatisect or pinnate, rarely simple or
decompound, mostly oblong to lanceolate, glabrous or scaly, rarely pilosulous;
veins simple, forked, or pinnate, mostly or entirely free (subg. Polypodium and
Marginalia) or re^larly anastomosing, with primary lateral veins and secondary
transverse veins forming areolae each bearing a single, unbranched, excurrent
veinlet (subg. Goniophlebium and Marginalia); sori surficial, lateral to terminal on
a free veinlet, round or rarely elongate, exindusiate; paraphyses filamentous or
absent; sporangia long-stalked.

Cosmopolitan; ca. 100 species.

EVANS, A. M. 1969. Interspecific relationships in the Polypodium pectinatum - plumula complex.
Ann. Missouri Bot. Gard. 55:193-293.

MAXON, W. R. 1916. Studies of tropical American ferns -No. 6. Contr. U. S. Natl. Herb.
17:541-608.

SOTA, E. R. de la 1966. Revisi6n de las especies Argentinas del grupo “Polypodium squamatum” L.
“Polypodiaceae” (s. str.). Revista Mus. La Plata, Secc. Bot. 10:69-186, 1. 1- VII.

1. Laminae sparsely to densely scaly throughout the abaxial surface (subg. Marginaria).22.

1. Laminae glabrous or pubesecent on the abaxial surface, occasionally scaly only the the abaxial
surface of the rachis and other axes.. 2.

2(1). Veins free and forked or occasionally anastomosing in a single, irregular series (subg.
Polypodium ) . .40.

2(1). Veins regularly anastomosing in 1- several series, the areolae with a single free, included
veinlet (subg. Goniophlebium). 3.

3(2). Laminae simple, not lobed. Sori in 2 or often more rows on either side of the midrib, usually
glaucous on the abaxial surface, oblong or narrowly ovate-lanceolate, obtuse to acute at the base.

266 POLYPODIACEAE: 55. POLYPODIUM

8-23 cm long, (3)4-6 cm wide, the midribs with ovate scales abaxially; rhizome scales orbiculate,
deciduous.

394. P. laevigatum

3(2). Laminae piimatifid to pinnate..4.

4(3). Sori elongate, usually 1.5-2 times longer than wide. Rhizomes compact, their scales linear,
reddish-brown; laminae narrowly to broadly lanceolate or elliptic-lanceolate, 40-100(120) cm long,
20 - 35(50) cm wide, with 30 - 50 or more pairs of segments; rachises lacking a dark brown abaxial
stripe.

398. P. plectolepidioides

4(3). Sori round or nearly so..5.

5(4). Laminae pinnatifid with all the segments broad at the base and fully adnate to the rachis..ll.

5(4). Laminae pinnate with at least the proximal pinnae short-petiolulate to sessile, tapered at the
base, and not adnate..6.

6(5). Distal pinnae short-petiolulate to sessile, not adnate..9.

6(5). Distal pinnae adnate, not short-petiolulate to sessile..?.

7(6), Sori in 1 row on each side of the costa; proximal pinnae obtuse to truncate at the base.
Laminae ovate-lanceolate, 10 - 40 cm long, 8-25 cm wide; rhizome scales linear-lanceolate, ca. 1 cm
long, 1 mm wide, thin, clathrate with large, clear lumina.

401. sessilifolium

7(6). Sori in 2-6 rows on each side of the costa, sometimes partially in 1 row on subfertile
laminae; proximal pinnae cuneate to acute at the base..8.

8(7). Sori in 2 or 3 rows on each side of the costa; rhizomes short-creeping; rhizome scales
spreading, with pale margins absent or nearly so; pinnae (1)2-3 cm wide. Laminae ovate-lanceolate,
15-50(75) cm long, 20-50(70) cm wide; pinnae 3-10(15) pairs.

403. P. triseriale

8(7). Sori in 4-6(10) rows on each side of the costa; rhizomes long-creeping; rhizome scales
appressed to the rhizome, with broad, pale margins; pinnae (3)4-6 cm wide. Laminae ovate-
lanceolate, 30 - 75 cm long, 25 - 40 cm wide; pinnae 4-10 pairs.

391.P. giganteum

9(6). Sori in 1 or 2 rows on each side of the costa; proximcil pinnae cordate at the base, the auricles
imbricate and overlapping the rachis. Laminae ovate-lanceolate, ca. 50 cm long, 30 cm wide; pinnae
ca. 10 subopposite, linear-lanceolate, falcate-ascending pairs, the terminal pinna slightly longer than
the subtending lateral ones.

393. P. kunzeanum

9(6). Sori in 3-6(10) rows on each side of the costa; proximal pinnae acute at the base..l0.

10(9). Sori in 3-5 rows on each side of the costa; pinnae l-2.5(3.5) cm wide. Laminae long-
lanceolate to oblong, 20 - 60(75) cm long, 12-30 cm wide; pinnae (1)4-16(19) linear-lanceolate to
oblong pairs.

389. P fraxinifolium

10(9). Sori in 4-6(10) rows on each side of the costa; pinnae (3)4-6 cm wide. Laminae ovate-
lanceolate, 30-75 cm long, 25 - 40 cm wide; pinnae 4-10 pairs.

391. P. giganteum

11(5). Laminae truncate or only slightly tapered at the base (with a few basal auricles in P. wagneri,
with puberulent rachises); stipes exalate.,13.

11(5). Laminae abruptly tapered at the base, the basal segments reduced to auricles; stipes
partially or entirely alate..l2.

12(11). Laminae slightly pilose on the abaxial surface of the veins, glabrous on the adaxial surface;
segments slightly and irregularly crenate, the lateral margins usually not parallel, the apices usually
acute. Laminae linear, 25 - 40 cm long, 5-7(9) cm wide.

396. P. loriciforme

POLYPODIACEAE: 55. POLYPODIUM 267

12(11). Laminae rather densely pilose on the veins on both surfaces; segments entire, the lateral
margins mostly parallel, the apices usually round. Laminae linear, 15-40 cm long, 3-7(8) cm wide.

402. P. subviride

13(11). Rhizome scales spreading, linear to lanceolate, usually persistent. .16.

13(11). Rhizome scales appressed, lanceolate to orbiculate, often deciduous and leaving a blackish
scar.. 14.

14(13). Sori in (1)2 rows on each side of the costa; segments usually round at the apex. Laminae
linear-lanceolate or lanceate, (15)30 - 70 cm long, (6)9-22 cm wide, the segments not or only slightly
falcate.

397. P. maritimum

14(13). Sori in 1 row on each side of the costa; segments usually acute at the apex.. 15.

15(14). Rhizomes (1)1.5 -2.5 mm in diam.; laminae lanceate to triangular, 6-24 cm long, 4- 12 cm
wide, with 10-22 non-falcate pairs of segments.

399. P. ptilorhizon

15(14). Rhizomes 3-5 mm in diam.; laminae lanceate, 20 - 45 cm long, 12-26 cm wide, with
20 - 35 pairs of usually falcate segments.

395. P. loriceum

16(13). Sori in 2 or 3 rows on each side of the costa. Laminae (10)25 - 40(100) cm long, 5 - 20(25)
cm wide, with 15-30(33) pairs of segments, the bases cordate; costae, veins, and laminae pilose on
both surfaces.

387. P. dissimile

16(13). Sori in 1 row on each side of the costa.. 17.

17(16). Rhizome scales dark brown to black and clathrate at least at the center, weakly to strongly
bicolorous..20.

17(16). Rhizome scales stramineous to reddish-brown throughout, not clathrate, concolorous..l8.

18(17). Laminae 1-1.5 times longer than wide; segments usu2dly 1.5-3 times their width distant,
4-7(10) pairs per pinna. L^mlinae oblong to broadly ovate, (10)15 - 30(45) cm long, (8)12-30 cm
wide.

388. P. flagellare

18(17). Laminae 2-4 times longer than wide; segments usually 0.5 -1(1.5) times their width
distant, (8)15-30 pairs per pinna.. 19.

19(18). Proximal pinnae somewhat contracted distal to the base; stipes and rachises usually with a
dark brown abaxial stripe; rhizome scales narrowly laneolate to acicular with a dark central area at
the base. Laminae lanceate, (5)8-30(45) cm long, (4)7-16 cm wide, with (5)8-15(30) pairs of
segments.

390. P. fuscopetiolatum

19(18). Proximal pinnae not contracted distal to the base; stipes and rachises lacking a dark brown
abaxial stripe, the rachises often reddish; rhizome scales broadly ovate-lanceolate without a dark
central area at the base. Laminae lanceate, (7)12-32 cm long, (3.5)7-13(16) cm wide, with
(8)15-33 pairs of segments.

400. P. rhodopleuron

20(17). Laminae at least 20 cm wide; pinnae at least (11)15 mm wide. Laminae ovate-lanceolate,
(15)25-65(115) cm long, 20 - 30(42) cm wide.

392. P. kuhnii

20(17). Laminae no more than 10(16) cm wide; pinnae no more than 8(10) mm wide. .21

21(20). Rhizome scales nearly Hnear distal to their expanded base, bicolorous, reddish-brown
nearly throughout to blackish at the center and pale along the mcU’gins; costae and laminae usually
sparsely to densely pilosulous on both surfaces. Laminae hnear, (12)25-45(55) cm long,
(2)3.5-7(14) cm \vide.

404. P. wagneri

268 POLYPODIACEAE: 55. POLYPODIUM

21(20). Rhizome scales broadly lanceolate, concolorous, blackish throughout; costae and laminae
glabrous on both surfaces. Laminae lanceate, (7)10 - 30(50) cm long, (3)5-15(21) cm wide.

405. P. wiesbaueri

22(1). Laminae pinnatifid or piimatisect, the segments entire..28.

22(1). Laminae pinnate-piimatifid or more divided, the segments or pinnae lobed. Veins free..23.

23(22). Rhizomes 0.5-1 mm in diam., very long-creeping; fronds 2-6 cm long. Laminae piimate-
pinnatifid, ovate-lanceolate, 1-3.5 cm long, 1-2 cm wide, the ultimate segments 0.5-1 mm wide;
rhizome scales ca. 0.5 mm long, bearing a few hairs on their surface.

m.P.fallax

23(22). Rhizomes 2-5 mm in diam., creeping; fronds 6-40 cm long..24.

24(23). Pinna lobes all short except the basal ones. Lobes usually 2-3(4) mm wide; rhizome scales
ca. 3-5 mm long, 0.75 mm wide, bicolorous with a dark, sharply defined median stripe, clathrate, the
cells large; laminae lanceate, 5-18(22) cm long, 3-10 cm wide, the pinna lobes few and short on
small specimens.

416. P. murorum

24(23). Pinna lobes mostly long, the basal ones often shorter than the median ones..25.

25(24). Laminae pinnate-pinnatifid, the lobes usually ca. 1 mm wide..27.

25(24). Laminae very deeply pinnate-pinnatifid or 2-pinnate to quadripinnatisect, the lobes usually
0.5-2 mm wide; rhizome scales ca. 4 mm long, 1 mm wide, concolorous, not clathrate, the cells
minute..26.

26(25). Laminae 2-pinnate-pinnatifid to quadripinnatisect; ultimate segments ca. 0.5 mm wide, not
lobed toward the apex; pinnae petiolulate (the stalks 2-5 mm long). Laminae lanceolate to linear-
lanceate, 9 - 25(40) cm long, (2)4-6(10) cm wide.

411. P. friedrichsthalianum

26(25). Laminae pinnate-piimatifid to 2-pinnate; ultimate segments 1-2 mm wide, conspicuously
lobed toward the apex; pinnae subsessile (the stalks 0.5-2 mm long). Laminae linear-lanceate,
10 - 25(28) cm long, (2.5)3.5-7 cm wide.

413. P. lindenianum

27(25). Pinnae up to 15 pairs; rhizome scales slightly bicolorous, the central area brown; laminae
triangular. Laminae 7- 15 cm long, (3)4-7(10) cm wide.

406. P. 'xaspidiolepis

27(25). Pinnae 35-45 pairs; rhizome scales concolorous, reddish-stramineous throughout; laminae
narrowly triangular. Laminae 9- 15 cm long, 1.5-5 cm wide.

418. P. xpinnatisectum

28(22). Laminae truncate or abrupt at the base..30.

28(22). Laminae long- attenuate at the base. Veins anastomosing.. 29.

29(28). Scales on the adaxial surface of the laminae 1.5 -2.5 mm long, narrowly lanceate from a
short-stellate base; rhizome scales 3-8 mm long. Laminae nearly linear, attenuate at the base and
apex, 12-26 cm long, 1.6 -3.5 cm wide, densely covered on the abaxial surface with reddish, very
narrowly lanceolate scales with ciliate bases.

408. P. bombycinum

29(28). Scales on the adaxial surface of the laminae up to 1 mm long, orbiculate-stellate or with
one 1 acicular ray; rhizome scales less than 3 mm long. Laminae nearly linear, attenuate at the base
and apex, 15-21 cm long, ca. 3.5 cm wide, densely covered on the abaxial surface with reddish,
broadly ovate-lanceolate, long-ciliate scales.

407. P. balaonense

30(28). Rhizome scales 1.5-3 mm long, lanceolate or with an expanded base and a long-acuminate
to sub-acicular apex, spreading (except appressed in P. montigenum).32.

30(28). Rhizome scales 0.25-1 mm long, roundish to triangular-ovate, very closely appressed to
the rhizome..31.

POLYPODIACEAE: 55. POLYPODIUM 269

31(30), Rhizomes 2.5 -3.5 mm in diam.; scales on the adaxial surface of the lamina orbiculate and
ca. 0.5 mm in diam. or with a short, linear-lanceolate ray and ca. 0.2 mm long. Laminae linear,
truncate at the base, acute at the apex, 10-40(55) cm long, 4-9(10) cm wide, with 25-40 pairs of
lateral segments.

417. P. myriolepis

31(30). Rhizomes 4-7 mm in diam.; scales on the adaxial surface of the lamina linear-lanceolate
with an expanded base, ca. 1 mm long. Laminae oblong, truncate at the base, acute to obtuse at the
apex, 20 - 35 cm long, 9-22 cm wide, with 13-32 pairs of lateral segments.

409. P. collinsii

32(30). Laminae mostly or entirely covered on the abaxial surface with orbiculate to lanceolate
scales, the latter with an expanded base; rhizome scales sharply bicolorous, the central stripe black or
dark brown, the margins pale, repand-crisped (except concolorous and whitish in P. furfuraceum);
small sinuses absent along the lamina margins..35.

32(30). Laminae subglabrous on the abtixial surface with lanceate scales; rhizome scales
concolorous or partially or entirely bicolorous with plane, ciliate margins; small sinuses present along
the lamina margins..33.

33(32). Rhizomes very long-creeping, 2-3 mm in diam.; stipes 2-5 cm distant; rhizome scales
appressed. Laminae (11)15-30 cm long, (5)10-14(18) cm wide, ovate-lanceolate to oblong.

415. P. montigenum

33(32). Rhizomes creeping, (2)3-5(10) mm in diam.; stipes 0.2-4 cm distant; rhizome scales
spreading.. 34.

34(33). Stipes 2-4 cm distant; laminae subdimorphic, oblong to triangular, the fertile ones
narrower and longer than the sterile ones; laminae (2)2.5 -3(4) times longer than wide, with
9 - 18(20) pairs of lateral segments, the fertile ones narrower than the sterile ones, obtuse to round at
the apex. Laminae 8-35 cm long, (3)4-11(13) cm wide.

423. P. remotum

34(33). Stipes 0.5-3 cm distant; laminae monomorphic, ovate-lanceolate or rarely oblong; laminae
1.5 -2.5(3) times longer than wide, with (12)16-24 pairs of lateral segments, the fertile and sterile
ones equally wide, usually acute to acuminate at the apex. Laminae 8-20(25) cm long, 4-8(12) cm
wide.

419. P. pleheium

35(32). Stipes mostly longer than the rachises. Rhizomes long-creeping; laminae broadly
triangular, 2-13 cm long, 2-8 cm wide, densely covered on the abaxial surface with lanceolate,
toothed-ciliate scales; pinnae 3-10 mm wide, 2-9 lateral p2urs per frond; veins anastomosing.

424. P. thyssanolepis

35(32). Stipes mostly shorter than or equalling the rachises..36.

36(35). Rhizomes 3-5 mm in diam.; rhizome scales concolorous or weakly bicolorous, without a
blackish central stripe. .39.

36(35). Rhizomes 1-1.5 mm in diam.; rhizome scales strongly bicolorous, with pale margins and a
blackish central stripe. Rhizomes long-creeping; laminae narrowly triangular to Unear, (2)3-10(13)
cm long, 1- 3.5(5) cm wide; pinnae 1-3 mm wide, (4)9-15 lateral pairs per frond; veins
anastomosing..37.

37(36). Larger scales of the abaxial lamina surface (but not the margins) acicular from an
orbiculate, expanded, entire to erose base, the apical portion entire to remotely toothed.

420. P. polypodioides var. aciculare

37(36). Larger scales of the abaxial lamina surface (but not the margins) lanceolate with an acute
to acuminate apex, the base not expanded, the apical portion subentire, erose-toothed, or densely
fimbriate-ciliate..38.

38(37). Larger scales of the abaxial lamina surface (but not the margins) entire or erose-toothed in
the apical region.

421. P. polypodioides var. michauxianum

270 POLYPODIACEAE: 55. POLYPODIUM

38(37). Larger scales of the abaxial lamina surface (but not the margins) strongly fimbriate-ciliate.

422. P. polypodioides wai. polypodioides

39(36). Scales of the abaxial lamina surface usually lanceolate, strongly toothed, at least 0.5 mm
wide, whitish with a sharply defined, black or brown center; rhizomes short-creeping and knotted, the
stipes approximate; rhizome scales concolorous, very pale reddish-brown, broadly lanceolate with an
acuminate apex, entire or weakly erose at the margin. Laminae linear or narrowly triangular,
10 - 30(40) cm long, 2-6(9) cm wide; sori superficial; veins free.

412. P. furfuraceum

39(36). Scales of the abaxial lamina surface usually orbiculate, ciliate to entire, mostly ca. 0.3 mm
wide, pale brown with a poorly defined, dark brown center; rhizomes long-creeping, the stipes 1-6
cm distant; rhizome scales reddish-brown, weakly bicolorous, linear-lanceolate with an attenuate
apex, sparsely ciliate at the margin. Laminae oblong, (10)12 - 30(35) cm long, (3.5)5-10 cm wide;
sori deeply sunken; veins anastomosing.

414. P. macrolepis

40(2). Pinnae acute to attenuate at the base. Laminae 15-25 cm long, 12-20 cm wide, oblong-
lanceolate; pinnae 5-10(15) pairs; sori medial, usually oval.

426. P. fratemum

40(2). Pinnae obtuse to truncate at the base or adnate to the rachis..41.

41(40). Laminae distantly serrate; sori submedial, round; rhizome scales linear distal to the
expanded base, somewhat revolute, the margins toothed. Laminae broadly lanceolate, 15-45 cm
long, 10-35 cm wide; pinnae (8)14-24 pairs; sori submedial.

425. P. echinolepis

41(40). Laminae entire or irregularly and shallowly crenate; sori usually medial or supra-medial
and shghtly elongate; rhizome scales lanceolate, entire..42.

42(41). Pinnae or segments 6-10 mm wide, 20-30(40) pairs per frond; rachises puberulous, the
hairs up to 0.2(0.5) mm long. Laminae (10)15-45(55) cm long, (6)7-13(15) cm wide, narrowly
lanceolate, (2)3-4 times longer than wide; sori up to 1.5 mm long, 1.25 mm wide.

428. P. ursipes

42(41). Pinnae or segments 10-25 mm wide, 10-15(20) pairs per frond; rachises glabrous or
sparsely hairy, the hairs 0.2 -0.5 mm long. Laminae (10)15 - 40(50) cm long, 8-20(25) cm wide,
lanceolate, 1.5 -2.5(3) times longer than wide; sori up to 2 mm long, 1 mm wide.

427. P. sororium

POLYPODIUM subg. GONIOPHLEBIUM (Blume) C. Chr.

387. Polypodium dissimile L. Syst. Nat. ed. 10, 2:1325. 1759.

Polypodium chnoodes Spreng. Neue Entdeck. 3:6. 1822. TYPE: Martinique, Kohaut (Sieber Exs.)
(LZ destroyed).

Polypodium retrofractum Desv. Mem. Soc. Linn. Paris 6:237. 1827. TYPE: “Habitat in America
calidiori,” collector unknown (P not seen), examined by Weatherby (Contr. Gray Herb. 114:32.
1936).

Polypodium chnoodes var. minus Christ, Bull. Herb. Boissier II, 6:49. 1906. TYPE: Navarro, Pcia.
Cartago, Werckle (P not seen).

TYPE: Jamaica, Browne (LINN 1251.24 not seen microfiche S. 1. Library).

Plants epiphytic, at 500-1500 m elevation, in forests, from west of Tilaran
(Pcia. Guanacaste), the Cordillera Central, the Atlantic coastal plain of Costa
Rica, the Reventazon and Orosi valleys (Pcia. Cartago), the Rio General valley,
the Fila Costeha near S. Vito, the Fortuna Reservoir site (Pcia. Chiriqui), near
Sta. Fe, above Cana, and La Esperanza on the Rio Calima (Depto. El Valle).
Also from the Antilles, Mexico, Guatemala, Honduras, Nicaragua, Trindad,
Venezuela, and Colombia.

POLYPODIACEAE: 55. POLYPODIUM 271

388. Polypodium flagellare Christ in Bomm. & Christ, Bull. Herb. Boissier 4:660.
1896.

Goniophlebium patens J. Smith, Bot. Voy. Herald 230. 1854, non Polypodium patens Swartz, 1788.
TYPE: Hacienda de Madre, Panama, Seemann (BM not seen).

TYPE: S. Mateo, Plain of Surubres, Pda. Puntarenas, Biolley 2671 (BR not seen
photo 4989; isotypes CR, US).

Plants epiphytic, at 0-700 m elevation, in forests, from near Surubres (Pcia.
Puntarenas), S. Luis de Turrubares (Pcia. S. Jose), around El Valle, near
Guaniquito (Pcia. Los Santos), and the Canal Zone and adjacent Pcia. Panama.

389. Polypodium fraxinifolium Jacq. Collectanea 3:187. 1791.

Polypodium rhizocaulon Willd. Sp. PI. ed. 4, 5:196. 1810. TYPE: Caracas, Distr. Fed., Venezuela,
Bredemeyer (B-Hb. Willd. 19690 not seen Tryon photo).

Polypodium distans Raddi, PI. Bras. Nov. Gen. 1:21, t. 31. 1825, non Kaulf., 1824, nom. illeg.
TYPE: Brazil, Raddi (FI not seen).

Polypodium polystichum Link, Hort. Reg. Bot. Berol. 2:101. 1833. TYPE: Cultivated in the
botanical garden at Berlin, originally from Brazil, Link (B not seen).

Polypodium mutabile Kunze, Linnaea 9:46. 1834. TYPE: Merida, Edo. Merida, Venezuela, Moritz
351 (B not seen).

Polypodium cymatodes Kunze, Linnaea 23:277, 317. 1850. SYNTYPES: Cultivated in the

botanical gardens at Leipzig and Berlin, originally from Venezuela, Kunze (LZ destroyed).

Goniophlebium deflexum T. Moore & Houlst. Card. Mag. Bot. 3:61. 1851. TYPE: Cultivated in
the garden at Kew, originally from Brazil, collector unknown (K? not seen).

Polypodium fluminense Veil. FI. Flumin. ll:t. 66. 1831; Arq. Mus. Nac. Rio de Janeiro 5:447.
1881. TYPE: Plate 66, which illustrates a plant collected from the region of Rio de Janeiro by
Vellozo.

Polypodium fraxinifolium \dx . fraxinellum Christ, Bull. Herb. Boissier II, 5:5. 1905. TYPE: Costa
Rica, Werckle & Bmne (P not seen).

TYPE: Caracas, Distr. Fed., Venezuela, Bredemeyerl (BM? not seen). The
ample illustration supplied by Jacquin (Icon. PI. Rar. t. 637. 1793) makes it likely
that the type was collected by Bredemeyer, rather than by Jacquin himself, for
Stafleu (Tax. Lit. 1:230. 1967) noted that Jacquin’s own material is scrappy.

Plants epiphytic, at (200)500-2100 m elevation, in forests, from the Cordillera
de Tilaran, the Cordillera Central and adjacent Atlantic lowlands of Costa Rica,
the Cordillera de Talamanca to Pcia. Chiriquf, between Cerro Pilon and El Valle,
Cerro Campana (Pcia. Panama), the Alturas de Nique (Pcia. Darien), and Pico
Mali, Alto del Buey, and the Rio El Salto (all Depto. Choco). Also from
Guatemala, Nicaragua, Trinidad, Venezuela, Colombia to Bolivia, and Brazil.

Polypodium scutulatum Sodiro (Sert. FI. Ecuad. 2:29. 1908) is a closely allied
species. A possible hybrid between P. fraxinifolium and P. triseriale is known from
near Boquete, Pcia. Chiriquf {Killip 5171, Comman 928, both US).

390. Polypodium fuscopetiolatum A. R. Smith, Amer. Fern J. 70:24, f. 13-15.
1980.

TYPE: 6-8 km west-northwest of Soyalo, Edo. Chiapas, Mexico, Breedlove
37155 (DS not seen).

Plants epiphytic or epipetric, at 800-1800 m elevation, in forests, from the
Cordillera de Tilaran, the Cordillera Central, the Meseta Central, the Fila de
Cedral, east of Copey (Pcia. S. Jose), the Fila Costeha near S. Vito, and Cerro

272 POLYPODIACEAE: 55. POLYPODIUM

385 Phlebodium pseudoaureum

388 Polypodium flagellare

POLYPODIACEAE: 55. POLYPODIUM 273

Punta (Pda. Chiriqm). Also from Mexico, Guatemala, El Salvador, Nicaragua,
Venezuela, and Colombia.

This species is related to P. rhodopleuron and less closely to P. plesiosorum
Kunze, which is known from Nicaragua but not from the Flora area. From the
latter species it differs in having a usually dark, abaxial stipe and rachis stripe,
narrowly lanceolate-attenuate, often denticulate rhizome scales ca. 0.5 - 0.75 mm
wide, and usually segments that are contracted distal to the base. Gomez
(Brenesia 6:50-51, f. 3. 1975) has found a hybrid apparently between P.

fuscopetiolatum and Microgramma lycopodioides, which he has named M.
xmoraviana.

391. Polypodium giganteum Desv. M6m. Soc. Linn. Paris 6:236. 1827.

Polypodium fraxinifolium subsp. luridum Christ, Bull. Herb. Boissier II, 6:48. 1906. TYPE:

Navarro, Pcia. Cartago, 1400 m, Werckle (P not seen; isotype US fragm NY).

Polypodium fraxinifolium subsp. articulatum Christ, Bull. Herb. Boissier II, 6:49. 1906.

LECTOTYPE: Turrialba, Pcia. Cartago, 550 m, Pittier 9061 (US; isolectotype CR), chosen by
Lellinger (Proc. Biol. Soc. Wash. 98:387. 1985).

Polypodium adnatum var. glaucescens Mille, Revista Col. Nac. Vicente Rocafuerte 9:208. 1927.
TYPE: Esmeraldas, Pcia. Esmeraldas, Ecuador, Sodiro in 1904 (QCA? not seen).

TYPE LOCALITY: Brazil. No specimen was seen in the Desvaux Herbarium
by Weatherby (Contr. Gray Herb. 114:30. 1936).

Plants terrestrial, at 0-1300(1700) m elevation, in forests, from north of S.
Ramon, the Atlantic coastal plain of Costa Rica and adjacent slopes of the
Cordillera Central, the Reventazon and Navarro valleys (Pcia. Cartago), the valley
of the Rio General, the vicinity of the Laguna de Chiriqui (Pcia. Bocas del Toro),
near the Fortuna reservoir site (Pcia. Chiriqui), El Valle, Sta. Rita ridge (Pcia.
Colon), Cerro Jefe, the El Llano -Carti road (Pcia. Panama), near Cana, and the
central Choco. Also from Venezuela, Colombia to Peru, and Brazil.

This species commonly has been identified as large P. fraxinifolium, but differs
from that species in having adnate distal pinnae, or as P. adnatum Kunze ex
Klotzsch, but differs from that species in lacking adnate proximal pinnae. It is also
close to the Andean P. caeceresii Sodiro, but differs in having broader, distinctly
clathrate, brownish rhizome scales and apical pinnae that are more acute at the
base.

392. Polypodium kuhnii Fourn. Bull. Soc. Bot. France 19:251. 1872.

TYPE: Isla de Omotepe, Lago de Nicaragua, Depto. Rivas, Nicaragua, Levy
1161 (P not seen).

Plants epiphytic, epipetric, or terrestrial, at 0-500(900) m elevation, in forests,
from the vicinity of Los Chiles (Pcia. Alajuela), northeast of Turrialba (Pcia.
Cartago), S. Luis de Turrubares (Pcia. S. Jose), the Peninsula de Osa and Golfito
(Pcia. Puntarenas), near Madden Dam (Canal Zone), Uva Island (Pcia.
Veraguas), Isla Taboga, Isla S. Jose, and the Rio de la Maestra (Pcia. Panama).
Also from Nicaragua, Venezuela, Colombia, and Ecuador.

FIGS. 385 - 390. Phlebodium and Polypodium. FIG. 385. Plant of Phlebodium pseudoaureum, Stern
et al. 1072. FIG. 386. Plant of Phlebodium decumanum, Johnson 1065, Guatemala. FIG. 387.
Median pinnae of P. dissimile, Weber 6176. FIG. 388. Plant of P. flagellare, Killip 2570. FIG. 389.
Median pinna of P. fraxinifolium, Cooper 6049. FIG. 390. Plant of P. fuscopetiolatum, Stork 2987.

274 POLYPODIACEAE: 55. POLYPODIUM

POLYPODIACEAE: 55. POLYPODIUM 275

393. Polypodium kunzeanum C. Chr. Ind. Fil. 536. 1906.

Polypodium cordatum Kunze, Linnaea 9:44. 1834, nom. illeg., non Desv., 1827. TYPE: Casapi,
Depto. Huanuco, Peru, PoeppigDiar. 1148 in July 1829 (LZ destroyed).

TYPE: A renaming of P. cordatum Kunze, and so based on the type of that
name.

Plants terrestrial, at 0 - 1000 m elevation, in forests, from Calera de S. Ramon,
S. Luis de Turrubares (Pcia. S. Jose), the vicinity of El Valle, the Laguna la
Yeguada (Pcia. Veraguas), El Valle, and Maje (Pcia. Panama). Also from
Colombia, Peru, and Brazil.

394. Polypodium laevigatum Cav. Descr. PI. 244. 1801.

Polypodium glaucophyllum Kunze ex Klotzsch, Linnaea 20:393. 1847. SYNTYPES: Venezuela?,
Otto 570 (B not seen); and Merida, Edo. Merida, Venezuela, Moritz 305 (B not seen; isosyntype BR
not seen photo 4999).

TYPE: Montana de S. Antonio, Quito, Pcia. Pichincha, Ecuador, Nee (MA not

seen Killip photo 484).

Plants epiphytic, at 800 - 1900 m elevation, in forests, from La Palma, Cartago
to the Orosi valley (Pcia. Cartago), the vicinity of El Valle, and Cana and Cerro
Tacarcuna. Also from the Lesser Antilles, Venezuela, and Colombia to Peru.

A possible hybrid of this species with P. wieshaueri is known from west of
Molinopampa, Depto. Amazonas, Peru {Wurdack 1363, US). Christensen (Dansk
Bot, Ark. 9(3): 11. 1937) seems to be in error in equating P. laevigatum with P.
semipinnatifidum (Fee) Mett., even if the latter species is a hybrid having P.
laevigatum as one parent.

395. Polypodium loriceum L. Sp. PI. 2:1086. 1753.

Polypodium duke Poir. Encyc. Meth. 5:523. 1804. TYPE: Martinique, collector unknown (P-Hb.
Juss. 1085 not seen microfiche S. I. Library).

Polypodium elegans Cav. ex Swartz, Syn. Fil. 35. 1806, non Poir., 1804, nom. illeg. TYPE: A plate
in Cavanilles’ “Elenchus Plantarum Horti Regii Matritensis.” Reference not seen.

Polypodium ramosum Lodd. ex Link, Hort. Reg. Bot. Berol. 2:97. 1833. TYPE: Cultivated in the
garden at London, origin unknown, Loddiges (K? not seen).

Polypodium falcaria Kunze, Linnaea 18:316. 1845. TYPE: Mexico, Leibold 90b (LZ destroyed;
isotypes Hb. Roemer, B, P none seen).

Goniophlebium cala^ala Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:93. 1857.
TYPE: Valley of Mexico, Mexico, Schaffner (P or RB not seen).

Goniophlebium invertens Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 7]:33, t. 37, f. 3.
1865. TYPE: Mt. Orizaba, Edo. Veracruz, Mexico, Schajfner424 (RB not seen).

1 Polypodium chartaceum Baker, J. Bot. Brit. For. 15:166. 1877. TYPE: Near Niebh, Pcia.

Pichincha, Ecuador, Dec 1871, Sodiro 48/43 (K not seen fragm US).

Polypodium loriceum var. heterolepis Sodiro, Anales Univ. Quito 11(72): 28 (repr. 352). 1894.
TYPE: Not stated.

Polypodium loriceum var. oligomemm Christ in Dur. & Pitt. Bull. Soc. Bot. Belgique 35, Mem.
221. 1896. TYPE: La Verbena, Pcia. S. Jose, Tonduz 8804 (BR not seen).

FIGS. 391-398. Polypodium. FIG. 391. Distal pinna of P. giganteum, Seemaum. FIG. 392. Median
piima of P. kuhnii, Erlanson 473. FIG. 393. Median pinna of P. kunzeanum, Allen 748. FIG. 394.
Plant of P. laevigatum, Lankester 619. FIG. 395. Median segments of P. loriceum, Standley &
Valerio 46989. FIG. 3%. Median segments of P. loriciforme, Mickel 3533. FIG. 397. Median
segments of P. maritimum, Correa A. 343. FIG. 398. Pinna of P. plectolepidioides, Brade 168.

276 POLYPODIACEAE: 55. POLYPODIUM

Polypodium loriceum var. obscurum Rosenst. Repert. Spec. Nov. Regni Veg. 6:308. 1909, as
'‘obscuraP TYPE: Mt. Abitagua, Andes of Quito, Pcia. Pichincha, Ecuador, Spruce 5234 (S not
seen).

Polypodium loriceum var. squamuligera Rosenst. Repert. Spec. Nov. Regni Veg. 6:308. 1909.
TYPE: Mt. Tungurahua, Pcia. Tungurahua, Ecuador, Spruce 5233 (S not seen).

Polypodium beyerianum Rosenst. Repert. Spec. Nov. Regni Veg. 22:17. 1925. LECTOTYPE:
Turrialba, Pcia. Cartago, 650 m. Erode & Erode 21 (S photo 5953; isolectotypes NY, UC), chosen by
Lellinger (Proc. Biol. Soc. Wash. 98:387. 1985).

Polypodium loriceum f. duplisorum Domin, Rozpr. Krai. Ceske Spolecn. Nauk, Tr. Mat.-Pnr. 2
[Pterid. Dominica]: 130. 1929, as "‘duplisoroP LECTOTYPE: Heights of Aripo, Trinidad, Eroodwoy
9957o (NY; isolectotype US), chosen by Lellinger (Proc. Biol. Soc. Wash. 89:727. 1977).

LECTOTYPE: Plate 7, f. 10 of Petiver’s “Pterigraphia Americana,” which is a
left-to-right transposition of t. 78 of Plunder’s “Traite...”, which illustrates a
specimen collected by Plumier on Martinique, chosen by Proctor (FI. Less. Antill.
2:331. 1977).

Plants epiphytic, at (500)1000-3000 m elevation, in forests, from the Cordillera
de Tilaran, the Cordillera Central, the Cordillera de Talamanca to Pcia. Chiriqui,
Sta. Rita ridge (Pcia. Colon), Cana and vicinity, and the Ciudad Bolivar - Quibdo
road at Km. 141 and the Rio Tamana (both Depto. Choco). Also from the
Antilles, Mexico to Nicaragua, Venezuela, Colombia to Bolivia, and Brazil.

396. Polypodium loriciforme Rosenst. Repert. Spec. Nov. Regni Veg. 22:17. 1925.

TYPE: Llanuras de S. Carlos, Pcia. Heredia, 200 m, Brade & Erode 493 (S not
seen photo 6015; isotypes NY, UC).

Plants epiphytic, at 0-1500 m elevation, in forests, from the Atlantic coastal
plain of Costa Rica, the Atlantic slope of Volcan Barba, the valley of the Rio
Reventazon, and the ridge above Platanillo (Pcia. Cartago), near Fortuna (Pcia.
Chiriqui), northwest of Sta. Fe, El Cope (Pcia. Code), and Cerro Pirre.

397. Polypodium maritimum Hieron. Bot. Jahrb. Syst. 34:527. 1904.

SYNTYPES: Rio Dagua near Buenaventura, Depto. El Valle, Colombia,
Lehmann 5035 (B not seen; isotype US); and Rio Micay, Depto. Cauca, Colombia,
Lehmann 8922 (B not seen).

Plants epiphytic, at 0-900 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, the Atlantic coastal plain of Costa Rica and
western Panama, near Sta Fe, near El Valle, around Portobelo (Pcia. Colon),
Cerro Campana and Sta. Rita ridge (both Pcia. Panama), and throughout the
northern half of the Choco. Also from Nicaragua, Venezuela, Colombia, and
Ecuador.

398. Polypodium plectolepidioides Rosenst. Repert. Spec. Nov. Regni Veg. 10:278.
1912.

Polypodium loriceum var. umbroticum Christ, Bull. Herb. Boissier II, 5:5. 1905. TYPE: Costa
Rica, Werckle & Prune (P not seen; possible isotype US).

TYPE: Turrialba, Pcia. Cartago, 650 m, Brade & Brade 361 {Ros. Fil Costar. Exs.
168) (S; isotypes GH, NY, UC, US).

Plants epiphytic, at 600-1000 m elevation, from the upper Rio Reventazon
valley (Pcia. Cartago).

POLYPODIACEAE: 55. POLYPODIUM 277

399. Polypodium ptilorhizon Christ, Bull. Herb. Boissier II, 5:6. 1905.

TYPE: Costa Rica, Werckle in 1904 (P not seen).

Plants epiphytic, at 1000-2500 m elevation, in forests and open areas, from the
Cordillera de Tilaran, the Cordillera Central, the Cordillera de Talamanca to
Pcia. Chiriqm, near Sta. Fe, Cerro Pilon (Pcia. Code), and Cerro Pirre. Also from
Nicaragua.

400. Polypodium rhodopleuron Kunze, Linnaea 18:315. 1845.

SYNTYPES: Xalapa, Edo. Veracruz, Mexico, Schiede & Deppe 745 (LZ
destroyed; probable isosyntype P not seen); and Mexico, Leibold 91 p. p. (LZ
destroyed).

Plants epiphytic, at 400-1900 m elevation, in forests, from the Cordillera de
Tilaran, Cerro Carpintera, the Reventazon and Orosi valleys (Pcia. Cartago), and
the Cordillera de Talamanca to Pcia. Chiriqui. Also from Mexico, Guatemala, El
Salvador, and Nicaragua.

This species is sometimes confused with P. plesiosorum Kunze, which occurs
from Mexico to Nicaragua. It differs in having narrower laminae 2-4 times
longer than wide and 6.5-10(13) cm wide, versus laminae 1.5 -2.5 times longer
than wide and 8-18 cm wide in P. plesiosorum, which also lacks reddish rachises
and has very broadly ovate, subappressed rhizome scales and hairs on the adaxial
surface of the rachis and costae.

401. Polypodium sessilifolium Desv. M6m. Soc. Linn. Paris 6:238. 1827.

Polypodium sumcuchense Hook. Icon. PI. l:t. 69. 1837. TYPE: Surucucho, near Cuenca, Pcia.
Azuay, Ecuador, (K not seen).

Polypodium andinum Karst. FI. Columb. 1:171, t. 85. 1861, nom. illeg., non Hook., 1860. TYPE:
Near Bogota, Depto. Cundinamarca, Colombia, Karsten (LE or W not seen).

Polypodium remotum Baker, Ann. Bot. (London) 5:470. 1891, nom. illeg., non Desv., 1827.

TYPE: Salazar, Depto. Norte de Santander, Colombia, Kalbreyer 843 (K not seen Maxon photo 308).

Polypodium uniseriale C. Chr. Ind. Fil. 572. 1906. TYPE: A renaming of P. remotum Baker, and
so based on the type of that name.

Polypodium sessilifolium var. papallactensis Mille, Revista Col. Nac. Vicente Rocafuerte 9:211.
1927. TYPE: Ecuador, Mille 2002 (QCA? not seen; isotype P not seen).

Polypodium pseudofratemum A. C. Smith, Bull. Torrey Bot. Club 58:307. 1931. TYPE: Cerro
Duida, Edo. Amazonas, Venezuela, Tate 645 (NY not seen fragm US).

TYPE: Peru, collector unknown (P not seen Cintract photo).

Plants epiphytic, at 2000-2700 m elevation, in forests, from the Cordillera
Central and Madre de Selva (Pcia. Cartago). Also from Colombia to Bolivia.

Specimens from Mexico named P. sessilifolium Liebm. (Kongel. Danske
Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:192 (repr. 401). 1849) are another
species; Liebmann’s name is a later homonym. Polypodium antillense Maxon
(Proc. Biol. Soc. Wash. 43:83. 1930) is an allied species.

402. Polypodium subviride Lellinger, Proc. Biol. Soc. Wash. 98:389, f. 17. 1985.

TYPE: La Eneida region near Cerro Jefe, Pcia. Panama, Kennedy 1117 (US).
Plants epiphytic, at 600-1200 m elevation, in forests, from near Sta. Fe, El
Valle and Bismark (Pcia. Code), near Colon (Pcia. Colon), Cerro Jefe, and Cerro
Pirre. Also from Colombia and Ecuador.

278 POLYPODIACEAE: 55. POLYPODIUM

401 P. sessilifolium

405 P. wiesbaueri 406 P. Xaspidiolepis

POLYPODIACEAE: 55. POLYPODIUM 279

403. Polypodium triseriale Swartz, J. Bot. (Schrader) 1800(2) :26. 1801.

Polypodium brasiliense Poir. Encyc. Meth. 5:525. 1804. LECTOTYPE: Plate 40 from vol. 1 of
Sloane’s “Voyage...”, chosen by Proctor (Ferns Jamaica 531. 1985).

Polypodium neriifolium Schkuhr, Vier Zwanzigste Kl. Linn. Pfl.-Syst. 1:14, t. 15. 1804. TYPE:
Locality and collector unknown (Hb. Breyne not seen).

Polypodium attenuatum Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:191. 1810. TYPE: Caracas,
Distr. Fed., Venezuela, Humboldt & Bonpland (B-Hb. Willd. 19683-1 to -3 not seen Tryon photos).

Polypodium variolatum Willd. Sp. PI. ed. 4, 5:192. 1810. TYPE: Brazil, Sieberl, comm.

Hoffmannsegg (B-Hb. Willd. 19685-1 to -3 not seen Tryon photos), synonymized by Morton (Contr.
U. S. Natl. Herb. 38:66. 1967).

Polypodium albopunctatum Raddi, Opusc. Sci. 3:287. 1819. TYPE: Brazil, Raddi (FI not seen).
Polypodium longifolium K. Presl, Dehc. Prag. 1:167. 1822, nom. illeg., non Cav., 1802. TYPE: Rio
de Janeiro, Est. Rio de Janeiro, Brazil, collector unknown (PRC not seen).

Polypodium elatius Schrad. Goett. Gel. Ajiz. 1824:868. 1824. TYPE: Brazil, Prince Maxmillian
von Wied-Neuwied (LE? not seen).

Polypodium lucens Schrad. Goett. Gel. Anz. 1824:868. 1824. TYPE: Brazil, Prince Maxmillian
von Wied-Neuwied (LE? not seen).

Polypodium preslianum Spreng. Syst. Veg. ed. 4. index. 1828; Link, Hort. Reg. Bot. Berol. 2:102.
1833. TYPE: Presumably a renaming of P. longifolium K. Presl, and so based on the type of that

name.

Polypodium gladiatum Kunze, Linnaea 9:45. 1834, nom. illeg., non Veil., 1827. TYPE: Cuba,
Poeppig (LZ destroyed; probable isotype BR not seen photo 4981).

Polypodium campylopodum Klotzsch, Linnaea 20:395. 1847. TYPE: Merida, Edo. Merida,

Venezuela, Moritz 352 (B not seen).

IPolypodium xiphophoron Kunze ex Mett. Abh. Senckenberg. Naturf. Ges. 2:73. 1856.

SYNTYPES: Surinam, Kegel (B not seen); Brazil, Pohl (B not seen); and French Guiana, Leprieur (B
not seen).

Goniophlebium intermedium Fee, Hist. Foug. Aiitill. [Mem. Foug. 11]:69, t. 18, f. 3. 1866. TYPE:
Guadeloupe, UHerminier (RB not seen).

1 Goniophlebium gauthieri Fee, Crypt. Vase. Bresil 1:112, t. 34, f. 3. 1869. SYNTYPES: Brazil,
Glaziou 2406 (P or RB not seen); and Ilha Sta. Catarina, Est. Sta. Catarina, Brazil, Gauthier (RB not
seen).

Polypodium palma Vellozo, FI. Flumin. ll:t. 69. 1831; Ajq. Mus. Nac. Rio de Janeiro 5:447. 1881.
TYPE: Plate 69, illustrating material collected around Rio de Janeiro by Vellozo; apparently no
specimens were made or are now extant.

Polypodium neriifolium var. acuminatissimum Kuntze, Revis. Gen. PI. 2:819. 1891. TYPE:

Between Turrialba and Cartago, Pcia. Cartago, Kuntze [Hb.l] 2256 (not seen; isotype NY).

Polypodium neriifolium var. heterophyllum Kuntze, Revis. Gen. PI. 2:819. 1891. TYPE

LOCALITY: Trinidad.

Goniophlebium ampliatum Maxon, Contr. U. S. Natl. Herb. 10:492. 1908. TYPE: A renaming of
Polypodium gladiatum Kunze, and so based on the type of that ncune.

Polypodium preslianum var. immsersum Rosenst. Repert. Spec. Nov. Regni Veg. 6:314. 1909, as
'Hmmersa.” TYPE: S. Antonio near Mapiri, Depto. La Paz, Bolivia, 800 m, Buchtien 1062 (S not
seen; isotype US).

FIGS. 399 - 407. Polypodium. FIG. 399. Median segments of P. ptilorhizon, Burger & Stolze 5331.
FIG. 400. Plant of P. rhodopleuron, Standley & Valerio 44662. FIG. 401. Median pinna of P.
sessilifolium, Alfaro 122. FIG. 402. Median segments of P. subviride, Correa & Dressier 160. FIG.
403. Median pinna of P. triseriale, Stork 2473. FIG. 404. Plant of P. wagneri, Allen 1960. FIG. 405.
Plant of P. wiesbaueri, Weber 6229. FIG. 406. Frond of P. aspidiolepis, Scamman 6124. FIG. 407.
Adaxial surface of pinna base of P. balaonense, Haught 2868.

280 POLYPODIACEAE: 55. POLYPODIUM

Polypodium brasiliense var. pleiosorum Rosenst. ex Hassl. Trab. Inst. Bot. Farm. Buenos Aires
45:71. 1928. SYNTYPES: Paraguay, Hassler 10139, 10139a, 11615 (all G none seen; possible

isosyntypes NY not seen); and Paraguay, Rojas 3960 (G not seen; possible isosyntype NY not seen).

TYPE: “India orientalis,” collector unknown (UPS-Hb. Thunberg not seen).
Plants epiphytic, at 0-1500 m elevation, in forests, from throughout the Flora
area except the drier portions of northwestern Costa Rica and western Panama.
Also from the Antilles, Mexico to Nicaragua, and tropical South America.

A possible hybrid with P. fraxinifolium is known from near Boquete (Killip 51 71,
Comman 928, both US).

404. Polypodium wagneri Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:255
(repr.337). 1864.

Goniophlebium pectinatum J. Smith, Bot. Voy. Herald 230. 1854, nom. illeg., non J. Smith, 1841.
TYPE: New city of Panama, Pcia. Panama, Seemann 14 (BM not seen; isotype K not seen Maxon
photo 245 fragm US).

Polypodium costaricense Christ in Bommer & Christ, Bull. Herb. Boissier 4:660. 1896. TYPE:
Plain of Surubres south of Puntarenas, Pcia. Puntarenas, Biolley 2677 (BR not seen photo 4985;
isotypes BR not seen photo 4986, US fragm NY).

SYNTYPES: Ocaha, Depto. Norte de Santander, Colombia, Schlim 636 (B not
seen); Panama, Wagner (B not seen); and Panama, Hayes (B not seen).

Plants epiphytic, at 0-1000(1800) m elevation, in forests, from Naranjo de
Alajuela (Pcia. Alajuela), the lower end of the Meseta Central, the valley of the
Rfo General, the Pacific side of the mountains of Panama from Boquete to El
Valle, Guaniquito (Pcia. Los Santos), the Canal Zone and adjacent Pcia. Panama,
near Cana and near El Real (Pcia. Darien), and the Serrama del Darien east of
Ungufa (Depto. Choco). Also from Venezuela and Ecuador.

405. Polypodium wiesbaueri Sodiro, Recens. Crypt. Vase. Quit. 65. 1883.

TYPE: Eastern slopes of Mt. Pichincha, Pcia. Pichincha, Ecuador, above 3000
m, Sodiro (Hb. Sodiro not seen).

Plants epiphytic or occasionally epipetric, at 2000-3000 m elevation, in forests,
pastures, and on roadsides, from the Cordillera Central and the Cordillera de
Talamanca to Pcia. Chiriquf. Also from Colombia and Ecuador.

This species can be divided into small, compact forms and large, lax ones. The
differences presumably are due to an exposed versus a protected habitat, and are
not of taxonomic significance.

POLYPODIUM subg. MARGINARIA (Bory) C. Chr.

406. Polypodium xaspidiolepis Baker, J. Bot. Brit. For. 25:26. 1887.

Polypodium thyssanolepis var. bipinnatifidum Christ in Bommer & Christ, Bull. Herb. Boissier
4:661. 1896. TYPE: S. Rafael de Cartago, Pcia. Cartago, 1600 m, Pittier 9721 (BR not seen; isotype
US).

TYPE: Costa Rica, Cooper (K not seen).

Plants epiphytic or sometimes epipetric, at 1400 - 2400 m elevation, in pastures
and along roads, from the southern slope of Volcan Irazu.

This species is a hybrid between P. friedrichsthalianum and P. thyssanolepis,
according to Wagner, Wagner, and Gomez (Brenesia 12/13:81-101. 1977).

POLYPODIACEAE: 55. POLYPODIUM 281

Apparent backcrosses between this hybrid and P. thyssanolepis are known from
Sanitorio Duran and vicinity {Scamman 7223, Weber 6042, both GH).

407. Polypodium balaonense Hieron. Bot. Jahrb. Syst. 34:529. 1904.

TYPE: Near Balao, Pcia. Guayas, Ecuador, Eggers 14286 (B not seen; isotypes F
not seen, US).

Plants epiphytic, in forests, in the Flora area known only from Tres Bocas on the
Rio Coasi, Pcia. Darien (Kirkbride & Duke 1385, NY) and from the “Isthmus of
Darien” (Seemann, US). Also from Colombia to Peru.

408. Polypodium bombycinum Maxon, Contr. U. S. Natl. Herb. 17:592. 1916.
TYPE: Rio Dagua, Depto. El Valle, Colombia, 300-1000 m, Lehmann 7666

(US; isotype GH not seen).

Plants epiphytic, at 100-1100 m elevation, in the Flora area known only from S.
Francisco, Pcia. Veraguas {Powell, US), near El Cope, Pcia. Code {Davidse &
Hamilton 23682, UC), and near Cana {Goldman 1915, US). Also from Venezuela,
Colombia, Bolivia, Guyana, and Brazil.

409. Polypodium collinsii Maxon, Contr. U. S. Natl. Herb. 17:583, t. 41. 1916.

Polypodium mickelii de la Sota, Revista Mus. La Plata, Secc. Bot. 10:163. 1966. TYPE: Along
ridge 3-5 km south of Villa Alta, Edo. Oaxaca, Mexico, Mickel 1130 (ISC; isotypes LP, NY).

TYPE: Near Pantepec, Edo. Chiapas, Mexico, Collins & Doyle 227 (US).

Plants epiphytic, in the Flora area known only from specimens grown at the
University of California Botanical Garden, Berkeley, received from C. H.
Lankester, Las Concavas, Pcia. Cartago, Costa Rica. Also from Mexico.

This species is included on the somewhat dubious presumption that Lankester’s
material, the only known from Costa Rica, originated in the wild and was not sent
to Lankester from Mexico and grown by him.

410. Polypodium fallax Schlechtend. & Cham. Linnaea 5:609. 1830.

Polypodium margaritifenim Christ, Bull. Herb. Boissier II, 5:2. 1905. TYPE: Sto. Domingo de
Golfo Dulce, Pcia. Puntarenas, 0 m, Tonduz 11257 (P not seen; isotype US).

Polypodium tuerckheimii Christ, Bull. Herb. Boissier II, 5:254. 1905. TYPE: Qubilquitz, Depto.

Alta Verapaz, Guatemala, 350 m, von Tuerckheim 7721 (P not seen Weatherby photo US).

TYPE: Misantla region, Edo. Veracruz, Mexico, Schiede & Deppe (B? not seen).
Plants epiphytic, at 0 - 800 m elevation, in forests, from the Rio General valley
and Sto. Domingo de Golfo Dulce (Pcia. Puntarenas). Also from Mexico to

Honduras, and Nicaragua.

411. Polypodium friedrichsthalianum Kunze, Farrnkrauter 2:55, 1. 123. 1850.

TYPE: Guatemala, Friedrichsthal PL Guat. Exs. 1322 (LZ destroyed; isotype W
not seen).

Plants epiphytic or rarely epipetric, at 800 - 1900 m elevation, in pastures and
along roadsides, from the Cordillera Central, the Mesta Central, Cerro
Carpintera, and the northeastern slopes of the Cordillera de Talamanca. Also
from Guatemala.

412. Polypodium furfuraceum Schlechtend. & Cham. Linnaea 5:607. 1830.

Polypodium nivosum Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:89. 1857.

TYPE: Huatusco, Edo. Veracruz, Mexico, Schafjner 193 (RB not seen).

282 POLYPODIACEAE: 55. POLYPODIUM

412 P. furfuraceum

409 P. collinsii

POLYPODIACEAE: 55. POLYPODIUM 283

Polypodium macbridense Shimek, Ferns Nicaragua 199, t. 20, f, 6-9. 1897. TYPE: Opposite El
Castillo, Depto. Rio S. Juan, Nicaragua, Shimek (ISC not seen).

Polypodium margallii Rovirosa, Pteridogr. Sur Mexico 206, t. 38A, f. 1-5. 1910. TYPE: Mesa de
Coapilla, Edo. Chiapas, Mexico, 1700 m, Rovirosa 1061 (K not seen; isotype PH not seen Morton
photo).

TYPE: Near Jalapa, Edo. Veracruz, Mexico, Schiede & Deppe [750] (B not seen;
isotypes LE not seen, MO, P not seen).

Plants epiphytic or rarely epipetric or terrestrial, at 0-2400 m elevation, in
forests, pastures, and along roadsides, from the Cordillera de Tilaran, the
Cordillera Central, the Meseta Central, the Atlantic coastal plain of Costa Rica,
the Cordillera de Talamanca to Pcia. Chiriqui, and Cana and vicinity. Also from
Mexico to Nicaragua.

413. Polypodium lindenianum Kenze, Farrnkrauter 2:83, 1. 134. 1851.

Polypodium cancellatum Fee, Gen. Fil. [Mem. Foug. 5]:242. 1852. TYPE: Cuba [Edo. Chiapas,

Mexico, according to A. R. Smith (FI. Chiapas 2:188. 1981)], Linden (RB not seen; isotype P not
seen).

SYNTYPES: Near S. Bartolo, Edo. Chiapas, Mexico, March, Linden (LZ
destroyed); Yerba Buena, Edo. Chiapas, Mexico, Linden (LZ destroyed); Edo.
Chiapas, Mexico, 7000 ft, March 1840, Linden (LZ destroyed); and Edo. Chiapas,
Mexico, Linden s. n. and 1539 (LZ destroyed; isosyntypes P not seen). The
numbered collection surely is Linden 1539, according to Kunze (Farrnkrauter 2:56.
1851).

Plants epiphytic, at 600-1500 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, and the Cachi reservoir (Pcia. Cartago). Also
from Mexico to Honduras, and Nicaragua.

414. Polypodium macrolepis Maxon, Contr. U. S. Natl. Herb. 17:584. 1916.

TYPE: Near Camp Aguacatal, east slope of Volcdn Chiriqui, Pcia. Chiriqui,

2100-2300 m, Maxon 5278 (US).

Plants epiphytic, at 2000-3200 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Pcia. Chiriqui.

This species differs from the similar P. myriolepis in having spreading, rather
than appressed, rhizome scales.

415. Polypodium montigenum Maxon, Publ. Field Mus. Nat. Hist., Bot. Ser.
17:306. 1938.

IPolypodium plebeium var. cooperi Baker, J. Bot. Brit. For. 25:25. 1887. TYPE: Costa Rica,
Cooper (K not seen; isotype US).

TYPE: Between Vara Blanca and La Concordia, Pcia. Heredia, 1600-1950 m,
Maxon & Harvey 8479 (US; isotypes GH, NY).

FIGS. 408 - 414. Polypodium. FIG. 408. Adaxial surface of pinna base of P. bombycinum, Uribe U.
1910, Colombia. FIG. 409. Adaxial surface of pinna base of P. collinsii, Lankester (cultivated,
originally from Costa Rica). FIG. 410. Plant of P. fallax, Skutch 3060. FIG. 411. Median pinna of P.
fiiedrichsthalianum, Stork 4794. FIG. 412. Abaxial surface of pinna base of P. furfuraceum, A. Smith.
FIG. 413. Median pinnae of P. lindenianum, Standley & Valerio 44449. FIG. 414. Abaxial surface of
pinna base of P. macrolepis, Mickel 3326.

284 POLYPODIACEAE: 55. POLYPODIUM

POLYPODIACEAE: 55. POLYPODIUM 285

Plants epiphytic or rarely epipetric or terrestrial, at 1500-2800 m elevation, in
forests, open places, and roadsides, from the Cordillera Central and the Cordillera
de Talamanca to Pcia. Chiriqui. Also from Mexico to Nicaragua.

Positive determination of P. plebeium var. cooped requires rhizome material;
the isotype is only a single, detached frond.

416. Polypodium murorum Hook. Icon. PI. l:t. 70. 1837.

TYPE: Vicinity of Quito, Pcia. Pichincha, Ecuador, Jameson 49 (K not seen).
Plants epipetric or epiphytic, at 2700 - 3400 m elevation, in forests, pastures, and
roadsides, from Volcan Irazu and the Cordillera de Talamanca from the Cerro de
la Muerte (Pcia. Cartago) to Cerro Chirripo (Pcias. Cartago and S. Jose). Also
from Hispaniola, Venezuela, and Colombia.

417. Polypodium myriolepis Christ in Bommer & Christ, Bull. Herb. Boissier
4:661. 1896.

Polypodium costaricanum Hieron. Bot. Jahrb. Syst. 34:530. 1904, non P. costaricense Christ, 1896.
LECTOTYPE: Volcan Irazu, Pcia. Cartago, Wendland 644 (B not seen), chosen by de la Sota
(Revista Museo La Plata, Secc. Bot. 10:169. 1966).

Potypodium wendlandii Hieron. Hedwigia 44:180. 1905, nom. superfl. TYPE: A renaming of P.
costaricanum Hieron., and so based on the type of that name.

TYPE: La Palma, Pcia. S. Jose, 1500-1700 m, Tonduz 9692 (BR not seen photo
5030; isotype US).

Plants epiphytic, at 1000-3000 m elevation, in forests, from the Cordillera
Central, Cerro Tablazo, and the Cordillera de Talamanca to Pcia.
Chiriqui.

418. Polypodium xpinnatisectum (Brade) G6mez, Amer. Fern J. 66:28. 1976.
Potypodium furfuraceum f. pinnatisectum Brade, Bradea 1:16, t. 5. 1969, as ''pinnatisectaP TYPE:

Cerro Carpintera, Pcia. Cartago, 1800 m, Brade & Brade 16 (HB not seen).

Plants epiphytic, at 1200-1800 m elevation, in forests, from Monte de la Cruz
(Pcia. Heredia), Cerro Carpintera, and the Cachi reservoir (Pcia. Cartago).

According to Gomez (Amer. Fern J. 66:28. 1976), this species is a hybrid

between P. fneddchsthalianum and P. furfuraceum.

419. Polypodium plebeium Schlechtend. & Cham. Linnaea 5:607. 1830.

Potypodium karwinskyanum A. Braun ex Kunze, Linnaea 23:279, 319. 1850. TYPE: Guatemala,
Friedrichsthal 76 (LZ destroyed?).

Potypodium cheilostictum Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:87. 1857.
TYPE: Orizaba, Edo. Veracruz, Mexico, Schajfner 453 (RB not seen).

Potypodium cartilagineum K. Presl ex Ettingsh. Denkschr. Kaiserl. Akad. Wiss., Math.-Naturwiss.
Kl., 22:79. 1864; Farnkr. Jetztw. 38, 1. 15, f. 8. 1864. TYPE: Not stated.

TYPE: Near Jalapa, Edo. Veracruz, Mexico, Schiede & Deppe [746] (B not seen;
isotype LE not seen).

FIGS. 415-422. Potypodium. FIG. 415. Plant of P. montigenum, de la Sota 5074. FIG. 416. Median
pinnae of P. muromm, Mickel 3253. FIG. 417. Adaxial surface of pinna base of P. myriolepis,
Standley & Valerio 50229. FIG. 418. Frond of P. pinnatisectum (after Brade). FIG. 419. Plant of P.
plebeium, Stork 4761. FIG. 420. Abaxial surface of pinna base of P. potypodioides var. aciculare,
Standley 33312. FIG. 421. Abaxial surface of pinna base of P. potypodioides var. michauxianum.
Carpenter 329. FIG. 422. Abaxial surface of pinna base of P. potypodioides var. potypodioides,
Johnston 123.

286 POLYPODIACEAE: 55. POLYPODIUM

Plants epiphytic, at 1300-2500 m elevation, in forests, from the Cordillera
Central, Cerro Carpintera, Cerro Piedra Blanca (Pcia. S. Jose), the Cordillera de
Talamanca to Pcia. Chiriqui, and near Sta. Fe. Also from Mexico, Guatemala,
and Honduras to Nicaragua.

420. Polypodium polypodioides var. adculare Weath. Contr. Gray Herb. 124:33.
1939.

TYPE: Rio Torres near S. Francisco de Guadalupe, Pcia. S. Jose, 1170 m,
Tonduz 8476 (GH fragm US).

Plants epiphytic or rarely epipetric, at 1000-2500 m elevation, in forests, open
areas, and roadsides, from the Cordillera de Tilaran, the Cordillera Central, and
the Cordillera de Talamanca to Pcia. Chiriqui. Also from Mexico, Guatemala, El
Salvador, and Nicaragua.

This variety is largely altiudinally distinct from the others of this species. The
acicular scales that characterize it are found on the abaxial surface of the laminae.
Non-diagnostic, similar scales are found on the frond margins of all of the P.
polypodioides varieties found in the Flora area.

421. Polypodium polypodioides var. michauxianum Weath. Contr. Gray Herb.
124:31. 1939.

TYPE: Kilby, Nansemond County, Virginia, U.S.A., Femald et al 4703 (GH not
seen).

Plants epiphytic, at 0-1100(1700) m elevation, in forests, from the Cabo Blanco
Nature Reserve and Nicoya (Pcia. Puntarenas), the Cordillera de Tilaran, the
Meseta Central, the Atlantic lowlands of Costa Rica and Panama, near the
Fortuna reservoir site (Pcia. Chiriqui), and Isla Taboga (Pcia. Panama). Also
from the United States, Mexico, Guatemala, Honduras, and Nicaragua.

422. Polypodium polypodioides (L.) D. Watt, Canad. Naturalist & Quart. J. Sci.
N.S., 3:158. 1867, var. polypodioides.

Acrostichum polypodioides L. Sp. PI. 2:1068. 1753. LECTOTYPE: Jamaica, collector unknown
(BM-Hb. Plukenet not seen), chosen by Weatherby (Contr. Gray Herb. 124:28. 1939).

Acrostichum ferruginosum L. Sp. PI. ed. 2, 2:1525. 1763. TYPE: Plate 89, f. 9 of Plukenet’s
“Phytographia,” as cited by Browne (Civ. Nat. Hist. Jamaica, ed. 2:105. 1789). No specimen that
could be a type is present in the Linnaean Herbarium.

Acrostichum ferrugineum L. Syst. Nat. ed. 12, 2:686. 1767, nom. superfl. TYPE: An inadvertent
renaming of ferruginosum L., and so based on the type of that name.

Polypodium incanum Swartz, Nov. Gen. Sp. PI. Prodr. 131. 1788, nom. superfl. TYPE: A

renaming oiA. polypodioides L., and so based on the type of that name.

Polypodium ceteraccinum Michx. FI. Bor.-Amer. 2:271. 1803, nom. superfl. TYPE: A renaming
of polypodioides L., and so based on the type of that name.

Polypodium velatum Schkuhr, Vier Zwcmzigste Kl. Linn. Pfl.-Syst. 2:188, t. 11b. 1805, nom.

superfl. TYPE: A renaming oiA. polypodioides L., and so based on the type of that name.

Polypodium chrysoconion Spreng. Syst. Veg. ed. 16, 4:51. 1827. TYPE: Plate 77 of Plumier’s
“Traite...”, which illustrates a specimen collected by Plumier on Hispaniola.

IPolypodium incanum var. umbrosum Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk.
Aid. V, 1:212 (repr. 60). 1849. TYPE: Chinantla District, Edo. Oaxaca, Mexico, 4000 - 5000 ft,
Liebmann (C? not seen).

IPolypodium incanoides Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:88. 1857.
TYPE: Huatusco, Edo. Veracruz, Mexico, Schaffner 199 (RB not seen).

POLYPODIACEAE: 55. POLYPODIUM 287

Wolypodium incanum var. oblongum Fourn. Mexic. PI. 1:83. 1872, as ""oblonga^ TYPE: A
renaming of P. incanoides Fee, and so based on the type of that name.

'^Polypodium mesetae Christ, Bull. Herb. Boissier II, 6:49. 1906. TYPE: Meseta Central, Pcia. S.
Jos6, 2000 m, Alfaro 16907 (P not seen).

Plants epiphytic, at 0-700 m elevation, in forests and open areas, from the
Peninsula de Nicoya (Pcia. Guanacaste), the Meseta Central, the Atlantic coastal
plain of Costa Rica, near Sta. Fe, El Valle, Penonome, the Canal Zone, and
throughout eastern Panama. Also from the Bahamas, the Antilles, Mexico,
Guatemala, Belize, Nicaragua, Trinidad, and Venezuela.

423. Polypodium remotum Desv. M^m. Soc. Linn, Paris 6:232. 1827.

Polypodium leucosticton Kunze ex Klotzsch, Linnaea 20:380. 1847. LECTOTYPE: Merida, Edo.

Merida, Venezuela, Moritz 336 (LZ destroyed?), inferentially chosen by Hieronymus (Bot. Jahrb.
Syst. 34:521. 1904; Hedwigia 48:260. 1909).

Polypodium plebejum var. columbiense Kuhn, Abh. Naturf. Ges. HaUe 11:40 (repr. 18). 1869, as
''columbiensisP TYPE: A renaming of P. leucosticton Kunze ex Klotasch, and so based on the type
of that name.

Polypodium microchasmum Baker, J. Bot. Brit. For. 25:44. 1887. TYPE: Tweeside, St. Andrew
Parish, Jamaica, Mrs. Baker (K not seen Maxon photo 168; isotype IJ not seen).

Polypodium plebejum var. palmense Christ, Bull. Herb. Boissier II, 5:4. 1905. SYNTYPES: La
Palma, Pcia. S. Jose, 1495 m, Tonduz 12571 (P not seen); and Volcan Turrialba, Pcia. Cartago, 2500
m, Pittier 13257 (P not seen).

Polypodium leucosticton var. hartwegianum Hieron. Hedwigia 48:260. 1909, as ''hartwegiana.”
SYNTYPES: Rio Blanco near Popayan, Depto. Cauca, Colombia, Stuebel 76 (B not seen); Near
Silvia, Depto. Cauca, Colombia, Stuebel 98 (B not seen); Mt. Tungurahua and near Banos, Pastaza
Valley, Pcia. Tungurahua, Ecuador, Stuebel 823, 830, 834 (all B none seen); and between Banos and
Jivarfa de Pmtuc, Pastaza Valley, Pcia. Tungurahua, Ecuador, Stuebel 975a (B not seen).

TYPE: “Habitat in America calidiori,” collector unknown (P not seen Cintract
photo).

Plants epiphytic, at (400)1100-2400 m elevation, in forests, from the Cordillera
de Tilaran, the Cordillera Central, the Corillera de Talamanca to Pcia. Chiriquf,
and Cerro Tute (Pcia. Veraguas). Also from Cuba, Jamaica, Hispaniola,
Guatemala, Venezuela, and Colombia to Bolivia.

424. Polypodium thyssanolepis A. Braun ex Klotzsch, Linnaea 20:392. 1847.

Polypodium rhagadiolepis Fee, Gen. Fil. [Mem. Foug. 5]:237. 1852. TYPE: Yerba Buena, Edo.
Chiapas, Mexico, Linden (RB not seen).

Polypodium purpusii Christ, Bull. Herb. Boissier II, 7:416. 1907. TYPE: Pachuca, Distr.

Zacoapan, Edo. Hidalgo, Mexico, 6000 - 7000 ft, Purpus in 1904 (P not seen; isotype US).

Polypodium incanum var. fimbriatum Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles
15:36. 1842. LECTOTYPE: Barranca de Guadalajara, Edo. Jalisco, Mexico, Galeotti 6438 (BR not
seen; isolectotype P not seen), chosen by Morton (Contr. U. S. Natl. Herb. 38:257. 1973).

SYNTYPES: Venta Grande, Colombia, Moritz 22 (B not seen; isosyntype P not
seen); and Colombia, Otto 896 (B not seen; isosyntype P not seen).

Plants epipetric or rarely terrestrial, at 1000-1700 m elevation, in forests, along
streams, and in open areas, from La Verbena (Pcia. S. Jose), Cerro Carpintera,
the southern slopes of Volcan Irazu, and the valley of the Rio Navarro (Pcia.
Cartago). Also from the southwestern United States, Cuba, Jamaica, Hispaniola,
Mexico, Guatemala, Honduras, Nicaragua, Venezuela, and Colombia to Bolivia.

288 POLYPODIACEAE: 55. POLYPODIUM

427 P. sororium 428 P. ursipes

POLYPODIACEAE: 55. POLYPODIUM 289

Wagner, Wagner, and Gomez (Brenesia 12/13:81 - 103. 1977) indicate that this
species usually is diploid through much of its range, but is commonly tetraploid
between Tres Rios and Cartago, which accounts for its ability to hybridize with P.
friedrichsthalianum to form P. aspidiolepis in that area. Wagner & Wagner (Fern
Gaz. 11:125-135. 1975) have found that this species also hybridizes with

Pleopeltis macrocarpa var. macrocarpa to form Pleopeltis xleucospora. In the Flora
area this hybrid is known from the crater of Volcan Irazu (O. Jimenez 1155, US).

POLYPODIUM subg. POLYPODIUM

425. Polypodium echinolepis F^e, M^m. Soc. Sci. Hist. Nat. Strasbourg 5 [M^m.
Foug.8]:87. 1857.

Polypodium sessilifolium Liebm. Kongel. Danske Vidensk. Selsk, Skr., Naturvidensk. Afd. V,
1:192 (repr. 40). 1849, non Desv., 1827, nom. illeg. TYPE: Barranca de Huitamalco, Edo. Veracruz,
Mexico, Liebmann FI Mex. 697 (C not seen), examined by A. R. Smith (FI. Chiapas 2:185. 1981).

Polypodium insigne Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:193
(repr. 41). 1849, non Blume, 1828, nom. illeg. LECTOTYPE: Hacienda de Mirador, Edo. Veracruz,
Mexico, Liebmann FI Mex. 700 (C not seen), chosen by A. R. Smith (FI. Chiapas 2:185. 1981).

Goniophlebium plectolepis Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:95. 1857.
TYPE: Near Orizaba, Edo. Veracruz, Mexico, Schaffner 187 (P not seen).

Goniophlebium serraUim Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:95. 1857.
TYPE: Orizaba, Edo. Veracruz, Mexico, Schaffner 495 (P not seen).

Polypodium macrodon Hook. Sp. Fil. 4:218. 1862. TYPE: Coban, Depto. Alta Verapaz,

Guatemala, Salvin (K not seen).

Polypodium legionarium Baker, Syn. Fil. 337. 1867, nom. superfl. TYPE: A renaming of P.
macrodon Hook., and so based on the type of that name.

Polypodium subareolatum Christ, Bull. Soc. Bot. Geneve II, 1:220. 1909. TYPE: Costa Rica, 2200
m, Werckle (P not seen).

TYPE: Cordoba, Edo. Veracruz, Mexico, Schaffner 188 (RB not seen).

Plants epiphytic or on fallen trunks, at 1200-1800 m elevation, in forests, from
Monteverde, near Zarcero (Pcia. Alajuela), and around Boquete. Also from
Mexico, Guatemala, Honduras, and Nicaragua.

426. Polypodium fraternum Schlechtend. & Cham. Linnaea 5:608. 1830.

IPolypodium henchmannii J. Smith ex T. Moore & Houlst. Gard. Mag. Bot. 3:18. 1851. TYPE:
Not stated.

Goniophlebium revertens Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:94. 1857.
SYNTYPES: Huatusco, Edo. Veracruz, Mexico, Schaffner 189 (P or RB not seen; isosyntype K not
seen); and Orizaba, Edo. Veracruz, Mexico, Schaffner 450 (P or RB not seen; isosyntype K not seen).

Polypodium petiolatum Davenp. Bot. Gaz. (Crawfordsville) 19:394. 1894. TYPE: Las Canoas,
Edo. S. Luis Potosi, Mexico, Pringle 4001 (GH not seen; isotype VT not seen Maxon photo).

TYPE: Near Jalapa, Edo. Veracruz, Mexico, Schiede & Deppe [753] (B not seen;
isotype LE not seen).

FIGS. 423-428. Polypodium. FIG. 423. Plant of P. remotum, von Tuerckheim, Guatemala. FIG.
424. Abaxial surface of pinna base of P. thyssanolepis, Torres R. 36. FIG. 425. Median pinna of P.
echinolepis, Maxon 5724. FIG. 426. Medium pinna of P. fraternum. Roe, Roe & Mori 584, Mexico.
FIG. 427. Median pinna of P. sororium, von Wedel 807. FIG. 428. Median pinna of P. ursipes,
Chrysler 5555.

290 POLYPODIACEAE: 66. PECLUMA

Plants epiphytic, at 200 m elevation, in the Flora area known only from La
Estrella, Pcia. Limon {Gomez P. 897, CR). Also from Mexico, Guatemala, Belize,
and Honduras.

427. Polypodium sororium Humb. & Bonpl. ex Willd. Sp. PL ed. 4, 5:191. 1810.

IGoniophlebium elatum Fee, Gen. Fil. [Mem. Foug. 5]:256. 1852, TYPE: Cuba, Linden 1890 (P
or RB not seen).

Polypodium glycirrhiza Fee, Hist. Foug. Antill. [Mem. Foug. 11]:49. 1866, non D. C. Eaton, 1856,
nom. illeg. TYPE: Guadeloupe, L’Herminier in 1861 (P or RB not seen).

Polypodium endresii Baker, Ann. Bot. (London) 5:468. 1891. TYPE: Costa Rica, Endres (K not
seen Maxon photo 240, fragm NY).

TYPE: Near Caripe, Edo. Monagas, Venezuela, Humboldt 424 (B-Hb. Willd.
19684-1 not seen microfiche S. I. Library).

Plants epiphytic or rarely epipetric, at 0-800(1300) m elevation, in forests, from
La Palma de S. Ramon and the Zarcero region (Pcia. Alajuela), the northern
slopes of the Cordillera Central, the Atlantic lowlands of Costa Rica and Panama,
the Fila Costeha near S. Vito, between La Pintada and Coclecito (Pcia. Code),
near Cerro Campana (Pcia. Panama), near Puerto Obaldia (Com. S. Bias), and
the northern half of the Choco. Also from the Antilles, Trinidad, Mexico,
Guatemala, Venezuela, Colombia to Peru, and Surinam.

This species is rather variable in frond form throughout its extensive range.
Specimens from the Choco differ from those of the remainder of the range in
being sparsely hairy on the axes, veins, and lamina surfaces, like P. chnoophonim
Kunze. But they differ from that species in having laminae that are dark green,
truncate, and fully pinnate at the base.

428. Polypodium ursipes Moritz ex C. Chr. Ind. Fil. 572. 1906.

Polypodium ursipes Moritz ex Kuhn, Linnaea 36:135. 1869, in synon.

Polypodium ambiguum Mett. ex Kuhn, Linnaea 36:134. 1869, non Desv., 1827, nom. illeg.

LECTOTYPE: Near Colonia Tovar, Edo. Aragua, Venezuela, Fendler 254 (US), chosen by Lellinger
(Proc. Biol. Soc. Wash. 98:387. 1985).

TYPE: A renaming of P. ambiguum Mett. ex Kuhn, and so based on the type of
that name.

Plants epiphytic or on fallen logs, at (800)1800-3300 m elevation, in forests,
from the Cordillera Central, Cerro Tablazo, the Cordillera de Talamanca to Pcia.
Chiriqui, near Sta Fe, and near El Valle. Also from El Salvador and Venezuela.

This species is very close to the Mexican P. hartwegianum Hook, in Benth., from
which it differs in having densely puberulous stipes with the hairs all less than 0.2
mm long. The valid description on which P. ursipes is based is that of P. ambiguum
Mett. ex Kuhn, under which “F. ursipes Moritz msc.” is listed in synonymy.

56. PECLUMA M. G. Price

Plants epiphytic or terrestrial; rhizomes short-creeping, often with proliferous
roots, scaly, the scales lanceolate, not clathrate, concolorous; stipes terete,
articulate to low phyllopodia, glabrous or sparsely pubescent; fronds small to
medium-sized, monomorphic; laminae pinnatisect, mostly elliptical to oblong or
lanceolate, glabrous or sparsely or rarely densely pubescent on the abaxial surface,
the rachis sparsely scaly in some species; segments long, narrow, usually linear,
closely spaced, fully adnate and often decurrent or sur current on the rachis; veins
free, mostly forked, sometimes casually to almost completely anastomosing; sori

POLYPODIACEAE: 56. PECLUMA 291

sori usually medial on the segments, round, exindusiate; paraphyses sometimes
present; sporangia long-stalked.

Neotropical; ca. 35 species.

EVANS, A. M. 1969. Interspecific relationships in the Polypodium pectinatum - plumula complex.
Ann. Missouri Bot. Gard. 55:193 ”293.

PRICE, M. G. 1983. Pecluma, a new tropical American fern genus. Amer. Fern J. 73:109-116.

1. Rachises not scaly or with inconspicuous, hair-like scales..4.

1. Rachises sparsely to densely scaly with lanceolate to ovate scales,.2.

2(1). Rachises black; rhizome scales narrowly lanceolate, acuminate; rachis scales inconspicuously
toothed or fimbriate; sporangia setose. Laminae linear, 15 - 40(50) cm long, 3 - 7.5 cm wide, cuneate
at the base or rarely subtruncate; segments 2-3 mm wide; veins l(2)-forked, free.

438. P. plumula

2(1). Rachises brown or reddish-brown; rhizome scales broadly ovate, acute; rachis scales entire;
sporangia not setose..3.

3(2). Laminae greatly tapered at the base, the basal segments gradually reduced to auricles,
strongly deflexed; rachis scales narrowly elliptic-lanceolate, flat, few. Laminae narrowly oblong-
elliptic, 8” 35(40) cm long, 2-5(6) cm wide, cuneate to attenuate at the base; segments 2-3.5 mm
wide, not uniformly long, the lamina outline often slightly irregular; veins 1-forked, free.

429. P. alfredii

3(2). Laminae slightly tapered at the base, the basal segments slightly reduced, but not to auricles,
slightly or not deflexed; rachis scales broadly ovate-lanceolate, cordate, subbullate, many. Laminae
narrowly oblong-elliptic, (6)15-45(50) cm long, (1.5)3-6(10) cm wide, cuneate to subtruncate at the
base; segments 1.5 - 3 mm wide, not uniformly long, the lamina outline often slightly irregular; veins
1-forked, free.

433. P. cupreolepis

4(1). Laminae abaxially glabrous or nearly so..8.

4(1). Laminae abaxially pubescent throughout or in an oblong area surrounding each sorus..5.

5(4). Laminae pilose mostly in an oblong patch around the sorus. Laminae narrowly rhombic,
attenuate at the base, acuminate at the apex, 20 - 90 cm long, (3)6-18 cm wide; segments 4-10 mm
wide; veins 2- or 3-forked, free.

439. P. ptilodon var. caespitosa

5(4). Laminae evenly pubescent on the abaxial surface..6.

6(5). Veins 1-forked, free; laminae up to 25(35) cm long, pilose on the adaxial surface. Laminae
narrowly elliptic, elliptic-lanceolate, or linear, truncate to acute at the base, 5-25(35) cm long,
1.5 -4(6) cm wide; segments 2-5 mm wide; veins free.

436. P. hyff-ometrica

6(5). Veins (l)2-forked, partially to almost completely anastomosing; laminae usually at least 30
cm long, puberulous or glabrous on the adaxial surface..7.

7(6). Laminae abaxially sparsely puberulent with hairs shorter than 0.2 mm or glabrous; segments
tapering from base to apex, the sinuses between them narrowly V-shaped. Laminae linear-lanceolate
to linear-elliptic or narrowly rhombic, acuminate to occasionally subtruncate at the base, 20 - 50(70)
cm long, 4-6(10) cm wide; segments 3-7 mm wide.

437. P. pectinata

7(6). Laminae abaxially rather densely pubescent with hairs 0.35-0.50 mm long; segments linear
from distal to the base to the apex, the sinuses between them narrowly U-shaped. Laminae linear,
obtuse or with a few auricles at the base, 15-40 cm long, 4-7 cm wide; segments 4-8 mm wide.

431. P. camptophyllaria var. camptophyllaria

8(4). Veins entirely free.. 10.

8(4). Veins partially to almost completely anastomosing. .9.

292 POLYPODIACEAE: 56. PECLUMA

9(8). Laminae (7)10-25 cm wide, truncate to acute at the base; veins only occasionally
anastomosing. Laminae lanceate or sometimes elliptic-lanceolate, (25)35-100 cm long; segments
4-9 mm wide.

430. P. divaricata

9(8). Laminae 4-6(10) cm wide, acuminate to occasionally subtruncate at the base; veins often
almost completly anastomosing. Laminae linear-lanceolate to linear-elliptic or narrowly rhombic,
20 - 50(70) cm long; segments 3-7 mm wide.

437. P. pectinata

10(8). Laminae 7-25 cm wide.. 12.

10(8). Laminae 3-7(9) cm wide.. 11.

11(10). Segments 2-3.5 mm wide; veins 1-forked; rhizome scales ovate-lanceolate, ca. 0.5-1 mm
wide, not clathrate. Laminae narrowly oblong, 15-55 cm long, attenuate to cuneate at the base, the
proximal segments reflexed.

429. P. alfredii

11(10). Segments 3.5-6 mm wide; veins (l)2-forked; rhizome scales linear-lanceolate, ca. 0.1 -0.2
mm wide, decidedly clathrate. Laminae narrowly lanceolate with several pairs of auricles at the base,
25-50 cm long, cuneate to attenuate at the base, the proximal segments or auricles usually not
reflexed.

432. P. consimilis var. consimilis

12(10). Rachis hairs on the adaxial surface ca. 0.2 mm long; rhizome scales densely fimbriate.
Laminae lanceate or sometimes elliptic-lanceolate, (25)35-100 cm long; segments 4-9 mm wide;
veins 2- or 3-forked.

430. P. divaricata

12(10). Rachis hairs at on the adaxial surface at least 0.5 mm long; rhizome scales entire or
sparsely hirsute. Laminae narrowly lanceolate, 20 - 80 cm long; segments 3-9 mm wide; veins 1- or
2-forked.. 13.

13(12). Rachises abaxially rather densely villous, the surface somewhat obscure.

435. P. eurybasis var. villosa

13(12). Rachises abaxially sparsely pilose, the surface clearly visible.

434. P. eurybasis var. glabrescens

429. Pecluma alfredii (Rosenst.) M. G. Price, Amer. Fern J. 73:113. 1983.

Polypodium alfredii Rosenst. Repert. Spec. Nov. Regni Veg. 22:15. 1925. TYPE: Turrialba, Pcia.
Cartago, 650 m, Brade & Brade 697 (S not seen photo 5936; isotype US).

Polypodium alfredii var. curtii Rosenst. Repert. Spec. Nov. Regni Veg. 22:15. 1925. TYPE: Rio
Chis region, near Juan Vinas, Pcia. Cartago, 1200 m, Brade & Brade 695 (S not seen; isotype US).

Polypodium tablazianum Rosenst. Repert. Spec. Nov. Regni Veg. 22:14. 1925. LECTOTYPE:
Carpintera, Pcia. Cartago, 1500 m, Brade & Brade 149 (S not seen photo 6080; isolectotypes UC,
US), chosen by Lellinger (Proc. Biol. Soc. Wash. 89:727. 1977).

Polypodium cyathicola Copel. Univ. Calif. Publ. Bot. 19:292, t. 43. 1941. TYPE: Cuautlancillo, 10
km north of Orizaba, Edo. Veracruz, Mexico, 1600 m, Copeland Mex. Ferns 127 (MICH not seen;
isotypes GH, UC, US).

Plants epiphytic, at 400-1300(1700) m elevation, in forests, from the Cordillera
de Tilaran, the Cordillera Central, the Meseta Central, Cerro Tablazo, and the
foothills at the northern end of the Cordillera de Talamanca. Also from Mexico
to Honduras.

430. Pecluma camptophyllaria (F^e) M. G. Price, Amer. Fern J. 73:113. 1983, var.
camptophyllaria.

Polypodium camptophyllarium Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]:86.
1857. TYPE: Near Ocana, Depto. Norte de Santander, Colombia, Schlim 128 p. p. in 1850 (P not
seen; isotypes BR not seen, G not seen, K not seen photo US).

POLYPODIACEAE: 56. PECLUMA 293

Polypodium cinerascens Lindm. Ark. Bot. 1:238, t. 11, f. 6. 1903. LECTOTYPE: Rio de Janeiro,
Est. Rio de Janeiro, Brazil, Regnell IA99 (S not seen; isolectotypes BR, C, S none seen, US), chosen
by Evans (Ann. Missouri Bot. Card. 55:252. 1968).

Plants terrestrial or occasionally epiphytic, at 300 - 1000 m elevation, in forests
and along roadsides, from near S. Ramon, the vicinity of Guapiles (Pcia. Limon),
and El Valle. Also from the Greater Antilles, Nicaragua, Venezuela, Colombia,
Peru, Bolivia, and Brazil.

431. Pecluma consimilis (Mett. in Tr. & Planch.) M. G. Price, Amer. Fern J.

73:113. 1983, var. consimilis.

Polypodium consimile Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:253 (repr. 335). 1864.
TYPE: Ocana, Pcia. Norte de Santander, Colombia, 6000 - 7000 ft, Schlim 633 (B not seen; isotypes
BR not seen, G not seen, L not seen photo 1849).

Polypodium consimile var. minus Hieron. Bot. Jahrb. Syst. 34:519. 1904, as ''minors
LECTOTYPE: Rfo Ambica, Depto. Tolima, Colombia, 2000 m, Lehmann 2353 (B not seen;
isolectotypes LE not seen, US), chosen by Evans (Ann Missouri Bot. Gard. 55:261. 1968).

Polypodium pityrolepis Rosenst. Report. Spec. Nov. Regni Veg. 22:16. 1925. TYPE: Rfo Chis,
near Juan Vinas, Pcia. Cartago, 1200 m, Brade & Brade 694 (S not seen; isotypes NY, US).

Plants terrestrial or on the bases of trees, at 1000-2000 m elevation, in forests,
from the Cordillera Central, Cerro Carpintera, and the Cordillera de Talamanca
to Pcia. Chiriquf. Also from Jamaica, Hispaniola, Mexico, Guatemala, Venezuela,
and Colombia.

432. Pecluma cupreolepis (Evans) M. G. Price, Amer. Fern J. 73:113. 1983.

Polypodium cupreolepis Evans, Ann. Missouri Bot. Gard. 55:224, f. 17. 1968. TYPE: Hills of
Patzcuaro, Edo. Michoacan, Mexico, Pringle 3353 (US; isotypes B not seen, GH, MICH not seen,
MO, NY, UC).

Plants epiphytic or epipetric, occasionally terrestrial, at 600-2000 m elevation,
from Las Nubes (Pcia. S. Jose), between Pacayas and Cot (Pcia. Cartago), and the
vicinity of Pejivalle (Pcia. Cartago). Also form Mexico to Nicaragua.

This species is closely related to P. alfredii, and the two species may only merit
varietal status.

433. Pecluma divaricata (Fourn.) Mickel & Beitel, Pterid. FI. Oaxaca 269. 1988.

Polypodium divaricatum Fourn. Mexic. PI. 1:80. 1872. TYPE: Zacuapan, Edo. Veracruz, Mexico,
Galeotti 6287 (P not seen; isotypes B not seen, G not seen photo 16671).

Polypodium bolivianum Rosenst. Repert. Spec. Nov. Regni Veg. 5:236. 1908. TYPE: Sirupaya,
near Yanacachi, South Yungas, Depto. La Paz, Bolivia, 2300 m, Buchtien 481 (S not seen photo 5958;
isotypes P not seen, US).

1 Polypodium bolivianum var. brevipes Rosenst. Repert. Spec. Nov. Regni Veg. 12:473. 1913.
TYPE: Polo-Polo, near Coroico, North Yungas, Depto. La Paz, Bohvia, 900 m, Buchtien 3497 (S not
seen).

Polypodium carpinterae Rosenst. Repert. Spec. Nov. Regni Veg. 22:16. 1925. TYPE: La

Carpintera, Pcia. Cartago, 1850 m, Brade & Brade 14 p. p. (S not seen).

Plants epiphytic or rarely terrestrial, at (200)900-2600 m elevation, in forests,
from the Cordillera Central, Cerro Carpintera, the Cordillera de Talamanca to
Pcia. Chiriquf, the Fila Costeha near S. Vito de Java, above Sta. Fe, Cerro
Campana (Pcia. Panama), Cana, and Alto del Buey. Also from Mexico to
Honduras, Venezuela, and Colombia to Bolivia.

294 POLYPODIACEAE: 56. PECLUMA

POLYPODIACEAE: 56. PECLUMA 295

In South America this species is distinct from P. eurybasis in its 2- or 3-forked
veins and basal lamina segments 1/2 or more as long as the longest segments. In
Central America, these characters distinguish the species less clearly.

434. Pecluma euiybasis van glabrescens (Rosenst.) Lellinger, Amen Fem J. 74:59.
1984.

Polypodium lachniferum var. glabrescens Rosenst. Repert. Spec. Nov. Regni Veg. 11:57. 1912.
TYPE: Unduavi, Depto. La Paz, Bolivia, 3400 m, Buchtien 2770 (S not seen; isotype US).

Polypodium lachniferum f. incurvatum Rosenst. Repert. Spec. Nov. Regni Veg. 11:57. 1912, as
'‘incurvataP TYPE: Unduavi, North Yungas, Depto. La Paz, Bolivia, 3400 m, Buchtien 2769 (S not
seen; isotypes UC, US).

Plants epiphytic, at (800)1200-2400 m elevation, in forests, from La Palma de
S. Ramon, between Moravia and Quebrada Platanillo (Pcia. Cartago), the Cerros
de Zurqui (Pcia. Heredia), Las Nubes and the Laguna de la Chonta (Pcia. S.
Jose), Cerro Horqueta (Pcia. Chiriqui), and the Rio Mutata at the foot of Alto del
Buey. Also from Venezuela, Colombia, Ecuador, and Bolivia.

For a comment on this species, see P. divaricata.

435. Pecluma eurybasis van villosa (Evans) Lellinger, Amen Fern J. 74:59. 1984.

Polypodium eurybasis var. villosum Evans, Ann. Missouri Bot. Card. 55:245. 1968. TYPE:

Foothills above Bogota, just north of the mouth of the Quebrada El Obispo, Depto. Cundinamarca,
Colombia, 2700 m, Fosberg 19688 (US).

Plants epiphytic, at (800)1700-2600 m elevation, in forests, from the Cordillera
de Talamanca to Pcia. Chiriqui, near Sta. Fe, and Cerro Tacarcuna. Also from

Venezuela and Colombia to Bolivia.

436. Pecluma hygrometrica (Splitg.) M. G. Price, Amen Fem J. 73:115. 1983.

Polypodium hygrometficum Splitg. Tijdschr. Natuurl. Gesch. Physiol. 7:409. 1840. TYPE:

Surinam, Splitgerber 1069 (L not seen photo 1890).

Polypodium pectinatum var. caliense Hieron. Bot. Jahrb. Syst. 34:517. 1904. TYPE: Near Las
Juntas del Dagua, Depto. El Valle, Colombia, 300-1000 m, Lehmann 7668 (B not seen; isotypes LE
not seen, US).

Polypodium tmncatulum Rosenst. Repert. Spec. Nov. Regni Veg. 9:343. 1911. TYPE: Valley of
the Rio Espiritu Santo, Antahuacana, Depto. Cochabamba, Bolivia, 750 m, Buchtien 2168 (S not
seen; isotype US).

Plants epiphytic, at 0-900 m elevation, in forests, from the Cordillera Central,
the Atlantic coastal plain of Costa Rica, the valley of the Rio General, the Canal
Zone and vicinity, Cana, and the northern and central Choco. Also from Mexico
to Honduras, Nicaragua, Venezuela, Colombia, Peru, Bolivia, and Guyana.

437. Pecluma pectinata (L.) M. G. Price, Amer. Fern J. 73:115. 1983.

Polypodium pectinatum L. Sp. PI. 2:1085. 1753. TYPE: Plate 37 of Plumier’s “Description...”,
which illustrates a specimen collected by Plumier on Martinique.

FIGS. 429 - 435. Pecluma. FIG. 429. Abaxial surface of portion of lamina of P. alfredii, Alfaro (J. D.
Smith) 6056. FIG. 430. Segments of P. camptophyllaria var. camptophyllaria, Standley 37291. FIG.
431. Abaxial surface of portion of lamina of P. consimilis var. consimilis, Torres R. 54. FIG. 432.
Abaxial surface of portion of lamina of P. cupreolepis, Standley 60040, Guatemala. FIG. 433. Abaxial
surface of portion of lamina of P. divaricata^ Maxon 5248. FIG. 434. Abaxial surface of portion of
lamina of P. eurybasis var. glabrescens, Standley & Valerio 50346. FIG. 435. Abaxial surface of
portion of lamina of P. eurybasis var. villosa, Maxon 5300.

296 POLYPODIACEAE: 56. PECLUMA

'ySikiuW4iMi^

I \,

436 P. hygrometrica

438 P. plumula

437 P. pectinata

441 G. alsophilicola

439 P. ptilodon var. caespitosa

440 G. alfarii

443 G. ambigens

POLYPODIACEAE: 56. PECLUMA 297

Plants epiphytic, at 0-2100 m elevation, in forests, from the Cordillera Central,
the Atlantic coastal plain of Costa Rica, the foothills of the northern end of the
Cordillera de Talamanca, the valley of the Rio General, Bajo Mona along the Rio
Caldera (Pcia. Chiriqui), Penonome and vicinity, the Canal Zone, near Cana, and
the central Choco. Also from the Antilles, Venezuela, and Colombia to Peru.

438. Pecluma plumula (Humb. & Bonpl. ex Willd.) M. G. Price, Amer. Fern J.
73:115. 1983.

Polypodium plumula Humb. & Bonpl. ex WiUd. Sp. PI. ed. 4, 5:178. 1810. LECTOTYPE:
Caracas, Distr. Fed., Venezuela, Bredemeyer (B-Hb. Willd. 19655-1 not seen microfiche S. I.
Library), chosen by Evans (Ann. Missouri Bot. Card. 55:233. 1968).

Polypodium schkuhrii Raddi, Opusc. Sci. 3:287. 1819, as ‘'schkuriiP TYPE: Based on P.

pectinatum sensu Schkuhr (Vier Zwanzigste Kl. Linn. Pfl.-Syst. 2:189, t. 17b. 1805), syn. excl., and so
based on the basis of that name, which is the plate or specimen (now at HAL) from which the plate
was drawn.

Polypodium elasticum Rich, ex Desv. Mem. Soc. Linn. Paris 6:233. 1827, non Bory ex Willd., 1810,
nom. superfl. TYPE: A renaming of P. plumula Humb. & Bonpl. ex Willd., and so based on the type
of that name. See Evans (Ann. Missouri Bot. Gen’d. 55:229. 1968) for an explanation of the

nomenclatural complexities of this name.

Polypodium pulchrum Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:41, t. 8, f. 2.
1842. TYPE: Jalapa, Edo. Veracruz, Mexico, 4000 ft, Galeotti 6332 (BR not seen photo 5023;
isotypes BR not seen, K not seen photo 15430bis).

IPolypodium inversum Velloso, FI. Flumin. 11:11, t. 72. 1831; Arq. Mus. Nac. Rio de Janeiro
5:448. 1881. TYPE: Based solely on Velloso’s plate, there being no type specimens preserved. Of
the four species Evans (Ann. Missouri Bot. Gard. 55:265. 1968) suggested, in stature, segmentation,
and phytogeography, this is most likely a synonym of P. plumula, and certainly not of P. filicula Kaulf.
nor of P. sicca Lindm.

Plants epiphytic, at 0 - 1900 m elevation, in forests, from the Meseta Central,
Volcan Barba, Cerro Tablazo, and Bahia Honda (Pcia. Veraguas). Also from
Florida and throughout most of tropical America.

439. Pecluma ptilodon van caespitosa (Jenm.) Lellinger, Amen Fern J. 77:101.
1988.

Polypodium pectinatum var. caespitosum Jenm. Bull. Bot. Dept. Jamaica N.S., 4:125. 1897.

TYPE: Old England, St. Andrew Parish, Jamaica, 4000 ft, Jenman (NY not seen).

Plants terrestrial or occasionally epiphytic, at 0 - 100 m elevation, in forests, in
the Flora area known only from the vicinity of Chiriqui lagoon, Pcia. Chiriqui (von
Wedel 2611, MO, US) and Tres Bocas on the Rio Coasi, Pcia. Darien (Kirkbride &
Duke 1383, NY). Also from Florida, Cuba, Jamaica, Hispaniola, and Mexico to
Honduras.

FIGS. 436 - 444. Pecluma and Grammitis. FIG. 436. Abaxial surface of portion of lamina of P.
hygrometrica, Lellinger 671. FIG. 437. Segments of P. pectinata, Standley & Valerio 48712. FIG. 438.
Abaxial surface of portion of lamina of P. plumula, Hitchcock. FIG. 439. Abaxial surface of portion
of lamina of P. ptilodon var. caespitosa, von Wedel 2611. FIG. 440. Median portion of lamina of G.
alfarii, Lellinger 998. FIG. 441. Median portion of lamina of G. alsophilicola, Lellinger & de la Sota
198. FIG. 442. Median portion of lamina of G. alsopteris, Killip & Garcia 33706, Colombia. FIG.
443. Median portion of lamina of G. ambigens, Standley 34907. FIG. 444. Plant of G. anfractuosa,
Maxon 8487.

298 POLYPODIACEAE: 57. GRAMMITIS

57. GRAMMITIS Swartz

Plants mostly epiphytic, occasionally epipetric or on fallen logs; rhizomes mostly
short-creeping to erect, conspiculously to inconspicuously scaly or rarely glabrous,
the scales lanceolate, clathrate or not; stipes obsolete to longer than the laminae,
wiry, inconspicuously articulate, not scaly or scaly only at the base, often long-
setose throughout; fronds minute to medium-sized, monomorphic or rarely slightly
dimorphic; laminae simple to pinnatisect (rarely more divided in sect.
Cryptosorus), mostly linear, oblong, or elliptic, glabrous, sparsely setose, pilose, or
pilosulous, or rarely glandular, the setae stiff to lax, short to long, mostly reddish-
brown, sometimes confined to the sori; veins unbranched, forked, or pinnate,
rarely irregularly anastomosing with a few included veinlets; sori terminal on a
vein or vein branch, superficial or occasionally impressed, round or rarely oblong,
exindusiate; paraphyses present or absent; sporangia long-stalked; spores trilete,
commonly greenish.

Pantropical; ca. 400 species in the broad sense, including ''Cryptosorus'' and
''Xiphopteris.'' The name ''Grammitis sect.. Xiphopteris (Kaulf.) Presl” used below
is the traditional name applied to this group of plants. This name is based on
Xiphopteris Kaulf., which is a synonym of Cochlidium, and so cannot be correctly
used in Grammitis. I do not wish to coin a new sectional name because Grammitis
is being divided into several genera by L. E. Bishop and the current sectional
names and concepts are not likely to continue in use.

BISHOP, L. E. 1977. The American species of Grammitis sect. Grammitis. Amer. Fern J.
67:101-106.

. 1988. Ceradenia, a new genus of Grammitidaceae. Amer. Fern J. 78:1-5.

MAXON, W. R. 1914. Notes upon Polypodium duale and its allies. Contr. U. S. Natl. Herb.
17:398 - 406.

. 1916. Polypodium trichomanoides and its American allies. Contr. U. S. Natl. Herb.

17:542-557.

MORTON, C. V. 1967. The genus Grammitis in Ecuador. Contr. U. S. Natl. Herb. 38:85-123.
. 1971. Supplementary notes on Grammitis in Ecuador. Phytologia 22:71 - 82.

1. Fronds pinnatifid or pinnate, rarely only deeply lobed..3.

1. Fronds simple or shallowly lobed..2.

2(1). Laminae not black-bordered (sect. Grammitastrum)..9.

2(1). Laminae black-bordered (sect. Grammitis). A.

3(1). Veins in the segments simple or 1-forked; laminae pinnatifid to pinnatisect, the segments
about as long as wide Xiphopteris"').

3(1). Veins in the segments pinnate, with several branches; laminae pinnatifid to pinnate-
pinnatifid, the segments generally much longer than wide (sect. Cryptosoms)..23.

4(2). Laminae up to 3 mm wide; rhizome scales 2-6 cells wide at the base.

489a. G. paramicola

4(2). Laminae more than 3 mm wide; rhizome scales 8-12 cells wide.. 5.

5(4). Laminae sparsely setose on the surface and margins, the setae ca. 0.5 mm long. Fronds
narrowly oblanceolate, 3-9 cm long including the stipes, 3-6 mm wide; sori discrete.

489. G. marginella

5(4). Laminae glabrous or nearly so..6.

6(5). Fronds 2.5-5 cm long; laminae spathulate, obtuse to round at the apex, the sterile veins
forked. Laminae 3-8 mm wide, attenuate toward the base; sori discrete.

488. G. leptopoda

6(5). Fronds 5-20 cm long; laminae linear, acute or apiculate at the apex, the sterile veins
simple..7.

POLYPODIACEAE: 57. GRAMMITIS 299

7(6). Midrib of the fronds greenish on the abaxial surface in the basal half of the lamina, the
sclerenchymatous sheath not exposed. Laminae 8-10(14) mm wide, attenuate toward the base; sori
discrete to somewhat confluent laterally.

487. G. bufonis

7(6). Midrib of the fronds brownish on the abaxial surface in the basal half of the lamina, the
sclerenchymatous sheath exposed; laminae 3-8 mm wide..8.

8(7). Border of the laminae 0.1 -0.2 mm wide; midribs lacking many-branched hairs. Laminae
3-8 mm wide, attenuate toward the base; sori confluent or discrete.

486. G. bryophila

8(7). Border of the laminae usually up to 0.1 mm wide; midribs bearing minute, many-branched
hairs, each branch multicellular, the cells pale except for the end walls. Laminae ca. 3 mm wide,
attenuate at the base; sori confluent.

488a. G. limbata

9(2). Midrib obvious on the surface abaxially; laminae 2-4 mm wide; stipes 5-15 mm long;
lamina setae apparently unicellular; rhizome scales absent. Laminae hnear, attenuate at the base,
2-11 cm long.

484. G. jungermannioides

9(2). Midrib immersed in the lamina abaxially; laminae (3)4-5 mm wide; stipes up to 5 mm long;
lamina setae obviously multicellular; rhizome scales ca. 1 mm long, 0.2 mm wide, setose hke the
laminae. Laminae linear, attenuate at the base, 3 - 9 cm long.

485. G. sprucei

10(3). Veins in the fertile segments 1-forked, with 2 hydathodes on the adaxial surface of the fertile
segments.. 17.

10(3). Veins in the fertile segments simple, with 1 hydathode on the adaxial surface of the fertile
segments..!!.

11(10). Laminae not cut to the midrib, the segments laterally connate.. 13.

11(10). Laminae cut to the rachis, the segments discrete.. 12.

12(11). Laminae glabrous on the abaxial surface, except for a few setae on the rachis, the setae
much shorter than the segments; rhizome scales clathrate, the lateral walls black. Laminae 2.5-11
cm long, 2-4(6) mm wide.

496. G. limula

12(11). Laminae strongly setose on the abaxial surface, mostly in the sori, the setae as long as the
segments; rhizome scales not clathrate, all walls tan or yellowish. Laminae 2.5-9 cm long, 2-4 mm
wide.

502. G. zurquina

13(11). Laminae monomorphic, all portions equally deeply lobed; rhizome scales not clathrate,
brown or golden, or absent. .15.

13(11). Laminae at least slightly dimorphic, the fertile portions less deeply lobed than the sterile
ones; rhizome scales clathrate, ecihate..l4.

14(13). Hairs on the abaxial surface of the rachis dark at maturity, setiform, 1-celled distal to a
very short basal cell. Stipes up to 1 cm long; laminae 4- 10 cm long, 2-3 mm wide.

498. G. myosuroides

14(13). Hairs on the abaxial surface of the rachis pale at maturity, lax, of 2-4 equally short cells.
Stipes up to 1 cm long; laminae 4- 10 cm long, 4-5 mm wide.

494. G. delitescens

15(13). Lateral margins of the segments parallel, the segments hnear. Rhizome scales golden,
ecihate. Stipes 1 - 2 cm long; laminae linear, attenuate at the base, acute at the apex, 5 - 14 cm long,
5-8 mm wide; fertile segments not gibbous; setae ca. 6 per segment.

492. G. cookii

15(13). Lateral margins of the segments not parallel, the segments rather triangular.. 16.

300 POLYPODIACEAE: 57. GRAMMITIS

16(15). Lamina setae atropurpureous, thick and stiff; rhizome scales pale brown with
atropurpureous setae; laminae subcoriaceous; axis of the segments at ca. a 75® angle to the midrib.
Laminae 5-15 cm long, 4-7 mm wide.

491. G. caucana

16(15). Lamina setae stramineous, thin and flexible; rhizome scales absent; laminae
membranaceous; axis of the segments at ca. a 45° angle to the midrib. Laminae 2-6 cm long, 3-4
mm wide.

499. G. pseudomitchellae

17(10). Laminae lobed or pinnatifid, not cut completely to the midrib; rhizome scales ecihate or
with pale ciha (except ciha dark in G. flabelliformis).. 19.

17(10). Laminae pinnatisect, cut completely to the rachis; rhizome scales with dark cilia.. 18.

18(17). Fertile pinnae strongly gibbous; sterile pinnae spaced about their width apart, 5-7(9) per
cm; pinnae strongly decurrent. Laminae 4-14(18) cm long, (3)4-8(10) mm wide.

490. G. blepharodes

18(17). Fertile pinnae not gibbous or weakly so; sterile pinnae spaced less than their width apart,
7-9 per cm; pinnae weakly decurrent. Laminae(2.5)4- 12 cm long, (3)5-7 mm wide.

493. G. daguensis

19(17). Laminae lobed usually ca. 1/2 the way to the midrib, with a broad, uncut, central portion;
stipes obsolete. Rhizome scales clathrate, setose; laminae linear, 3-7 cm long, 5-10 mm wide,
finely setose on both surfaces, the setae reddish-brown.

501. G. truncicola

19(17). Laminae lobed usually more than 3/4 the way to the midrib, with a narrow, uncut, central
portion; stipes obvious..20.

20(19). Laminae lacking long, dark setae; sporangia with many short, pale setae. Laminae 2-7 cm
long, 3-5 mm wide; segments ascending at ca. a 30 - 45° angle to the midrib.

497. G. mitchellae

20(19). Laminae with long, dark setae; sporangia glabrous..21.

21(20). Segments at a 75 - 90° angle to the midrib. Segments approximate to subimbricate.

494a. G. flabelliformis

21(20). Segments at a 60 - 75° angle to the midrib..22.

22(21). Laminae 2-8(13) cm long, 3-5(7) mm wide; segments ascending at ca. a 60° angle to the
midrib.

500. G. setulosa

22(21). Laminae (7)10-15 cm long, 6-9 mm wide; segments sHghtly ascending, at ca. a 75° angle
to the midrib.

495. G. hyalina

23(3). Stipes and laminae (or their hairs) glandular (also thinly pilose in G. nubigena), the glands
sessile, spherical or elongate, whitish (G. jamesonioides group). .34.

23(3). Stipes not glandular, the laminae rarely so..24

24(23). Stipes broadly alate nearly to the base, the ala 1-2(3) mm wide at the stipe apex, tapered
gradually below (G. fucoides group).. 35.

24(23). Stipes exalate or with a narrow ala usually only at the apex..25.

25(24). Fronds pilose or setose, at least in the sori or on the stipes..28.

25(24). Fronds glabrous, the stipes sometimes hirsutulous..26.

26(25). Sori in shallow depressions and supramedial or in rather deep cavities and submarginal (G.
subcapillaris group). .37.

26(25). Sori superficial and medial..27.

27(26). Laminae papyraceous to membranaceous, the margins plane; segments at least (2)3 times
longer than wide (except shorter in G. phlegmaria) (G. suprasculpta group). .41.

27(26). Laminae coriaceous, the margins revolute; segments no more than 2(2.5) times longer than
wide (G. moniliformis group).. 46.

POLYPODIACEAE: 57. GRAMMITIS 301

28(25). Fronds pilose to weakly sericeous, with thin, flexible, pale, long hairs; fronds papyraceous
to membranaceous, flexible, often entirely pendent..33.

28(25). Fronds setose, with rather thick, stiff, dark brown to reddish-brown setae; fronds
coriaceous to herbaceous, stiff, erect or the laminae pendent from stiff, spreading stipes..29.

29(28). Laminae coriaceous; segment margins strongly revolute; rhizome scales clathrate, entire,
often iridescent; rhizomes very long-creeping in most species (G. moniliformis group). .46.

29(28). Laminae subcoriaceous to herbaceous; segment margins not or scarcely revolute; rhizome
scales not clathrate or clathrate with whitish setae (except clathrate and lacking setae in G. firma)..30.

30(29). Stipes equalling or exceeding the rachises (except in G. ambigens and mathewsii); laminae
not or only slightly tapered at the base, truncate; segments mostly 4 mm wide or more, tapering from
the base to the apex. Midribs not black; rhizome scales golden, brown, or reddish-brown, not
clathrate, often of several files of bulging cells and bristle-tipped (G. meridensis group). .51.

30(29). Stipes much shorter than the rachises; laminae tapered at the base, obtuse to acute;
segments mostly less than 3.5(5) mm wide, linear (except in G. pminosa and staheliana).31.

31(30). Segments often tapered, 1-3 times longer than wide; midribs and veins occult; lime dots
not present on the adaxial surface; laminae less than 10 cm long (G. staheliana group).. 57.

31(30). Segments linear, many times longer than wide..32.

32(31). Midribs and veins occult; lime dots absent on the adaxial surface of the laminae (G. pilipes
group).

467b. G. pilipes

32(31). Midribs and often the veins black or dark brown; lime dots common on the adaxial surface;
laminae mostly more than 10 cm long (G. taxifolia group). .58.

33(28). Stipes more than 5 cm long; laminae mostly abrupt at the base, truncate to obtuse; laminae
barely cut to the midrib, the segments mostly slightly connate laterally (G. asplenifolia group). .63.

33(28). Stipes less than 2 cm long; laminae mostly tapered at the base, acute to attenuate; laminae
mostly cut to the rachis, the segments not at all connate laterally (G. cultrata group). .66.

34(23). Stipes and laminae lacking setae; glands usually elongate, abundant. Rhizome scales ovate,
not clathrate, brownish-orange, entire or sparsely setose; laminae linear, attenuate at the base,
acuminate or acute at the apex, 5-18(31) cm long, 0.5-1.8(2.7) cm wide, the segments ascending,
entire to shallowly lobed,

456. G. jamesonioides

34(23). Stipes and laminae sparsely setose; glands usually spherical, sparse. Rhizome scales
lanceate, not clathrate, brown or reddish-brown, entire; laminae elliptic, acute at the base and apex,
8 - 12 cm long, 2.5 - 5 cm wide, the segments ascending, shallowly crenate-lobed.

467. G. nubigena

35(24). Sori supramedial; stipe ala strongly undulate; rhizome scales clathrate, black with
abundcmt, short, whitish setae. Laminae elliptic-oblanceolate, 7-15(26) cm long, 3-7.5(10) cm
wide; segments decurrent, sometimes surcurrent, 1-5 cm long, 2-4 mm wide.

476. G. subsessilis

35(24). Sori nearly marginal; stipe ala entire; rhizome scales not clathrate, brownish to golden with
none or few, long, dark setae..36.

36(35). Sori not on marginal teeth; segments entire or furcate, sometimes repeatedly so. Rhizome
scales lanceolate, brownish; laminae linear to narrowly elliptic, up to 60 cm long, 4-12(25) cm wide;
segments 2-3.5 mm wide.

454. G, fucoides

36(35). Sori on prominent, marginal teeth; segments entire or sometimes furcate, rarely repeatedly
so. Rhizome scales ovate-lanceolate, golden; laminae linear to elliptic-lanceolate, up to 45 cm long,
4-15 cm wide.

469. G. podocarpa

37(26). Rhizome scales ciliate; stipes at least (2)3 cm long, densely hirsutulous..39.

37(26). Rhizome scales eciliate; stipes up to 2(3) cm long, glabrous..38.

302 POLYPODIACEAE: 57. GRAMMITIS

38(37). Segments mostly spreading, the acroscopic segment margins inserted at a 90° angle to the
rachis. Laminae linear-elliptic, attenuate at the base, acuminate at the apex, 5 - 25(30) cm long,
1-2.5 cm wide.

475. G. suhcapillaris

38(31). Segments mostly shghtly ascending, the acroscopic segment margins inserted at a 75° angle
to the rachis (the largest segments often somewhat gibbous). Laminae linear-elliptic, attenuate at
the base, acuminate at the apex, 4- 17 cm long, (0.5)1- 1.5(2.2) cm wide.

479. G. suspensa

39(37). Laminae pinnatifid, not cut quite to the midrib, the midrib subglabrous abaxially, not
hirsutulous. Laminae linear-elliptic, 4- 14 cm long, 1-2 cm wide, attenuate at the base, acuminate at
the apex; segments strongly decurrent, ascending at a 60° angle to the rachis.

455. G. isidrensis

39(37). Laminae pinnatisect, cut to the rachis, the rachis hirsutulous abaxially.. 40.

40(39). Segments linear, decurrent, 1-2 cm long; laminae 5-15(21) cm long, 1.2 -3.8 cm wide,
apiculate at the apex. Laminae linear-eUiptic, shghtly tapering and obtuse at the base; segments

(1) 1.5 -2(2.5) mm wide, decurrent, at a 75 - 90° angle to the rachis.

445. G. apiculata

40(39). Segments oblong, shghtly decurrent, 0.5-1 cm long; laminae 2-5 cm long, 0.5-1 cm wide,
acute at the apex. Laminae linear, obtuse to acute at the base; segments 1.5 -2.5 mm wide, at a
45 - 60° angle to the rachis.

452. G. epiphytica

41(27). Segments entire or shghtly repand..43.

41(27). Segments crenately or serrately lobed..42.

42(41). Rhizome scales ecihate. Laminae hnear, 7 - 30(40) cm long, (1)1.5 -3(6) cm wide,
attenuate at the base, acute at the apex; segments decurrent, lanceate, serrately lobed.

478. G. suprasculpta

42(41). Rhizome scales with long, pale ciha. Laminae hnear, 3-23 cm long, 0.7-3 cm wide,
acuminate at the base, acute to acuminate at the apex; segments decurrent, oblong, deeply serrately
lobed.

463. G. micula

43(41). Segments nearly hnear, 4-10 times longer than wide; rhizome scales clathrate with
occluded lumina; rachises often flexuous toward the apex. Laminae hnear, 8-21(32) cm long,
0.6 - 2.5 cm wide, attenuate at the base, acute at the apex; segments decurrent, ascending at ca. a 45°
angle to the rachis.

481. G. tmesipteris

43(41). Segments rather triangular, mostly less than (4)5 times longer than wide; rhizome scales
clathrate with rather clear lumina; rachises not flexuous toward the apex.. 44.

44(43). Rachises with bifid or stellate hairs on the abaxial surface. Laminae linear, 2-5 cm long,
0.4 -0.8 cm wide, attenuate at the base, obtuse at the apex.

470. G. randallii

44(43). Rachises glabrous on the abaxial surface or with simple hairs.. 45.

45(44). Rhizome scales distantly setose, the setae straight, sharp, ca. 0.2 mm long; segments mostly

(2) 3 -5(8) times longer than wide. Laminae linear, 4-14 cm long, 1-2 cm wide, attenuate at the
base, acute at the apex; segments spreading at a 60 - 75° angle to the rachis.

448. G. barbensis

45(44). Rhizome scales not setose; segments mostly ca. as long as wide. Laminae linear, 10-15
cm long, 0.5 - 1.2 cm wide, attenuate at the base, obtuse or broken at the apex.

467a. G. phlegmaria

46(27,29). Abaxial lamina surface setose, especially in the sori, the setae long, reddish; segments
more than 2 times longer than wide (except sometimes in G. anfractuosa)..^.

46(27,29). Abaxial lamina surface glabrous; segments less than 2 times longer than wide..47.

POLYPODIACEAE: 57. GRAMMITIS 303

47(46). Rhizome scales more than 12 cells wide, 3-5(8) mm long, 0.5- 1(2) mm wide, iridescent,
the lateral cell walls rather thin, reddish. Stipes glabrous or occasionally sparsely setose; laminae
linear, 4- 15(20) cm long, 0.5- 1.4 cm wide.

465. G. moniliformis

47(46). Rhizome scales up to 6 cells wide, 1.5-2 mm long, 0.2 -0.3 mm wide, not iridescent, the
lateral cell Wcdls very thick, blackish. Stipes glabrous; laminae linear, 4-13 cm long, 0.5 -1.2 cm
wide.

461. G. melanosticta

48(46). Rhizomes ca. 1 mm in diam., short, erect, distant along wide-creeping, proliferous roots.
Stipes sparsely setose; laminae linear-elliptic, 2-8(11) cm long, 0.4-1 cm wide.

444. G. anfractuosa

48(46). Rhizomes 1-3(5) mm in diam., long-creeping to erect- ascending, without proliferous
roots.. 49.

49(48). Rhizome scales ovate-lanceolate, 1-2 mm (ca. 24 cells) wide; rhizomes very long-creeping.
Lateral and end clathrate cell walls of the rhizome scales equciUy thin; Icuninae linear-elliptic, 8-20
cm long, 1.2- 1.9 cm wide, not curling when dry.

471. G. rigescens

49(48). Rhizome scales linear-lanceolate, 0.5-1 mm (6-10 cells) wide; rhizomes short-creeping to
erect-ascending..50.

50(49). Segments spreading, obtuse to round at the apex; sterile segments glabrous abaxially;
lateral clathrate cell walls of the rhizome scales thicker than the end walls; laminae often curling
when dry. Laminae linear-eUiptic or linear-lanceolate, 5-15(23) cm long, (0.7) 1-2.5 cm wide.

482a. G. xiphopteroides

50(49). Segments ascending, acute at the apex; sterile segments long-setose abaxially; lateral
clathrate cell walls of the rhizome scales as thick as the end walls. Laminae (5)10-16 cm long,
(0.8)1 -2(2.3) cm wide.

468. G. pilosissima

51(30). Segments strongly ascending, at a (20)30 - 45° angle to the rachis, 5-15(25) cm long.
Rhizome scales golden to brown, entire; laminae ovate, triangular, or occasionally oblanceolate,
10-25 cm long, 5- 15 cm wide; segments 4-6 mm wide, the venation partially areolate.

472. G. sectifrons

51(30). Segments spreading, at a 75-90° angle to the rachis, 0.5 -4.5(6) cm long..52.

52(51). Rhizome scales reddish-brown or rarely brown, with copious, short, pale setae or fewer,
long, reddish-brown setae,.55.

52(51). Rhizome scales brown, entire or with a few brown setae..53.

53(52). Lamina segments (2)3-5 mm wide; rhizome scales usually sparsely setose; sori with
branched glands having enlarged, terminal cells; venation free. Laminae triangular to oblong,
6-19(23) cm long, 2-8(10) cm wide.

457. G. kalbreyeri

53(52). Lamina segments (3)5-8 mm wide; rhizome scales entire; sori eglandular; venation
areolate..54.

54(53). Laminae not setose abaxially or on the margins; stipes mostly as long as the rachises.
Laminae narrowly oblong, (4)8-25 cm long, (4)6-9(12) cm wide.

450. G. comuta

54(53). Laminae setose abaxially and on the margins; stipes mostly 0.5-0.75(1) times as long as
the rachises. Laminae narrowly lanceolate, 15-45 cm long, 5- 10 cm wide.

459a. G. mathewsii

55(52). Rhizome scales sparsely setose with long, dark, marginal setae; laminae usually less than 10
cm long; stipes usually shorter than the laminae. Laminae narrowly lanceolate, 3.5-10 cm long,
1.2-3 cm wide, with long, soft, reddish setae on both surfaces.

443. G. amhigens

304 POLYPODIACEAE: 57. GRAMMITIS

55(52). Rhizome scales densely setose with short, pale, marginal setae; laminae usually more than
10 cm long; stipes usually longer than the laminae..56.

56(55). Segments entire, obviously decurrent, spaced less than their width apart. Laminae oblong
or lanceate, 4-23 cm long, 1-5.5 cm wide.

462. G. meridensis

56(55). Segments crenate, not decurrent, often spaced more than their width apart. Laminae
oblong, 5 - 37 cm long, 1 - 5 cm wide.

460. G. mayoris

57(31). Laminae pinnatifid, cut almost to the midrib, the lobes narrowly triangular to oblong; setae
on the sporangia conspicuous, much longer than the capsules; laminae 3-10 cm long, 0.6-2 cm
wide; rhizome scales pale brown with abundant setae; stipes and rachises sparsely long-setose;
laminae elhptic to linear-elhptic, acute to acuminate at the base, acuminate at the apex.

474. G. staheliana

57(31). Laminae lobed, with a broad, uncut central portion, the lobes broadly triangular; setae on
the sporangia few and inconspicuous, not longer than the capsules. Laminae 2-6 cm long, 0.7-1 cm
wide; rhizome scales pale brown with scattered setae; stipes and midribs sparsely long-setose;
laminae linear-elliptic, acuminate at the base, acute at the apex.

469a. G. pruinosa

58(32). Laminae pinnate-pinnatifid. Laminae oblong, 12-15 cm long, 3.5-4 cm wide.

466. G. myriophylla

58(32). Laminae pinnatifid or pinnatisect..59.

59(58). Laminae rather densely hirsutulous on the abaxial surface. Rhizome scales reddish-brown
with a terminal and sometimes a few lateral setae ca. 1 mm long; stipes and rachises sparsely long-
setose; laminae linear-elliptic, 18-30 cm long, 2-5 cm wide, acuminate at the base, caudate at the
apex, segments linear, spreading to shghtly ascending, with adaxial lime dots, the costa visible on the
abaxial surface; sporangia glabrous.

442. G. alsopteris

59(58). Laminae glabrous on the abaxial surface (sometimes circumsorally sparsely hirsutulous in
G. taxifolid)..Gd.

60(59). Segment costae, but not the veins, clearly visible abaxiaUy; laminae caudate at the apex.
Rhizome scales blackish, clathrate, entire; stipes and rachises sparsely setose; laminae linear-elliptic,
3 - 18 cm long, 0.6 - 4.5 cm wide, acuminate at the base, caudate at the apex.

453. G. firma

60(59). Segment costae and the veins both clearly visible abaxially; laminae not caudate at the
apex..61.

61(60). Segment margins entirely glabrous. Rhizome scales ecihate; stipes setose, more densely so
toward the base; laminae linear-elhptic, 7-31 cm long, 2-6(8) cm wide, acuminate at the apex, acute
or rarely obtuse or acuminate at the base.

483. G. zeledoniana

61(60). Segment margins sparsely setose toward the apex..62.

62(61). Rhizome scales sparsely long-cihate; stipes setose, more densely so at the base. Stipes
3- 15 cm long; laminae linear-elliptic, 6 - 30(40) cm long, 2-5 cm wide.

473. G. semihirsuta var. semihirsuta

62(61). Rhizome scales entire; stipes evenly setose throughout. Stipes 2-7 cm long; laminae
linear-elliptic, 6-16(20) cm long, 2 -3.5(5) cm wide.

480. G. taxifolia

63(33). Rhizome scales dull, reddish-brown to stramineous, with obscure cells about as long as
wide and dense setae often paler than the body of the scales; sporangia glabrous (except in G.
lehmanniana).. 65.

63(33). Rhizome scales shiny, reddish-brown, with obvious cells several times longer than wide and
setae about the same color as the body of the scales; sporangia setose..64.

POLYPODIACEAE: 57. GRAMMITIS 305

64(63). Stipes 0.3 -0.8 mm in diam., atropurpureous, sparsely setose; laminae 2-3(4) cm wide.
Stipes (5)9-21 cm long, often horizontal with the laminae pendent; laminae lanceolate, (6)10-40 cm
long, obtuse at the base, acute to acuminate at the apex.

446. G. asplenifolia

64(63). Stipes 1-1.5 mm in diam., gray or brown, densely setose; laminae (3.5)4 -7 cm wide.
Stipes 10-26 cm long; laminae lanceolate, 15-50 cm long, acute at the base, acuminate at the apex.

449. G. chrysleri

65(63). Segments with short, pale setae on the abaxial surface (these sometimes partly deciduous),
usually 5 mm or less wide; sporangia setose; stipes 0.25-0.75 mm in diam. Laminae linear,
10-35(50) cm long, 2-5(8) cm wide, obtuse to truncate at the base.

459. G lehmanniana

65(63). Segments glabrous on the abaxial surface, usually more than 5 mm wide; sporangia not
setose; stipes 1-1.5 mm in diam. Laminae linear, 8-35(65) cm long, 3.5-9 cm wide, obtuse to
truncate at the base.

447. G. atroviridis

66(33). Sericeous hairs along the segment margins all simple or absent..70.

66(33). Sericeous hairs along the segment margins partially or entirely paired or sessile-stellate..67.

67(66). Hairs on the abaxial surface of the laminae sericeous, mostly sessile-stellate, those on the
rachis often reddish-brown. Rhizome scales ciliate; laminae linear, (8)15 - 40(60) cm long, (1.5)4 -6
cm wide, acuminate at the base, acute to acuminate at the apex; segments (3)4 -5(6) cm wide, gray-
green, broadly adnate, the acroscopic margins often nearly surcurrent, the mairginal hairs mostly
sessile-stellate; sporangia short- to long-setose.

458. G. lanigera var. lanigera

67(66). Hairs on the abaxial surface of the laminae simple, entirely pale..68.

68(67). Hairs on the segment margins simple or sometimes paired; segments more than 4 times
longer than wide. Rhizome scales absent; laminae linear or in small fronds elliptic, 8-40 cm long,
2-7 cm wide, acute to acuminate at the base and apex; segments linear, 2-4 mm wide, not or
slightly decurrent, not surcurrent; sporangia short-setose or glabrous.

482. G. turrialhae

68(67). Hairs on the segment margins sessile-stellate or paired; segments ca. 2-2.5 times longer
than wide..69.

69(68). Rhizome scales ciliate. Laminae linear, (3.5)6-55 cm long, 0.7 -3.5(4) cm wide,
acuminate to attenuate at the base, acute at the apex; segments 2-6.5 mm wide, yellow-green or pale
green, decurrent at the base, the acroscopic margin curving proximally to meet the rachis; sporangia
short- to long-setose.

451. G. cultrata

69(68). Rhizome scales absent. Laminae linear, 3-22 cm long, 0.5- 1.5 cm wide, acute at the base
and apex; segments 2-3 mm wide, pale green, decurrent at the base, the acroscopic margin curving
proximally to meet the rachis; sporangia short- to long-setose.

464. G. mollissima

70(66). Launinae decidedly sericeous on the abaxial surface and veins. Rhizome scales ca. 1 mm
long, 0.3 mm wide, dark brown with whitish setae; laminae 2- 12 cm long, 5-10(12) mm wide, linear,
acuminate to attenuate at the base, acute at the apex, with adaxial lime dots; segments not or only
slightly gibbous, often less than 3 times longer than wide, parallel-sided, membranaceous; sporangia
glabrous.

477. G. subtilis

70(66). Laminae glabrous or nearly so on the abaxial surface..71.

71(70). Segments strongly gibbous, almost triangular, glabrous on the abaxial surface; rhizome
scales absent. Laminae 3-25 cm long, (0.5) 1-2(3) cm wide, linear, acuminate at the base, acute at
the apex; segments membranaceous; sporangia not setose.

440. G. alfarii

306 POLYPODIACEAE: 57. GRAMMITiS

71(70). Segments not or only slightly gibbous, essentially parallel-sided, sparsely sericeous on the
abaxial surface; rhizome scales ovate-lanceolate, ca. 1 mm long, 0.5 mm wide, gray with reddish-
brown setae 1-2 mm long. Laminae (5)10-30 cm long, 1.5 -2.5 cm wide, linear, obtuse or acute at
the base, acute at the apex; segments papyraceous; sporangia decidedly long-setose.

441. G. alsophilicola

GRAMMITIS sect. CRYPTOSORUS (Fee) Tryon & Tryon

440. Grammitis alfarii (J. D. Smith) Morton, Contr. U. S. Natl. Herb. 38:103.
1967.

Polypodium oligosorum Mett. ex Kuhn, Linnaea 36:132. 1869, non Klotzsch, 1847, nom. illeg.
SYNTYPES: Colonia Tovar, Edo. Aragua, Venezuela, Moritz 460 (B not seen fragm US); and
Venezuela, Fendler208 (B not seen), Karsten 10 (B not seen).

Polypodium alfarii J. D. Smith, Bot. Gaz. (Crawfordsville) 33:262. 1902. TYPE: Sierra Alta de
Navarro, Pcia. Cartago, 2000 m, Alfaro 73 (/. D. Smith 8063) (US).

Plants epiphytic, at (800)1000-2700 m elevation, in forests, from the Cordillera
Central, Cerro Carpintera, the vicinity of S. Isidro del General, the Cordillera de
Talamanca to Pcia. Chiriqui, and Cerro Campana. Also from Venezuela,
Colombia, and Ecuador.

441. Grammitis alsophilicola (Christ) F. Seym. Phytologia 31:176. 1975.

Polypodium alsophilicola Christ, Bull. Soc. Bot. Geneve II, 1:219. 1909, as ''alsophilicolumP

TYPE: Vallee du Copal, Pcia. Puntarenas, Pittier 10971 (P not seen; isotype US).

Plants epiphytic, at 0-900 m elevation, in forests, from the valley of the Rio
General and coastal southern Costa Rica, El Valle, near El Llano on the road to
Cartf (Pcia. Panama), Cana and the Sambu basin (Pcia. Darien), and Alto del
Buey and environs. Also from other localities in Colombia.

442. Grammitis alsopteris Morton, Contr. U. S. Natl. Herb. 38:112, t. 4. 1967.

TYPE: Hacienda La Mascota, Canton Mera, Pcia. Napo-Pastaza, Ecuador, 900
m, Mexia 7013 (US; isotype UC).

Plants epiphytic, at (100)500-1700 m elevation, in forests, from the
northwestern slopes of the Cordillera Central, the vicinity of S. Isidro del General,
the Fila Costeha near S. Vito de Java, near El Valle, Cerro Campana, Cerro
Tacarcuna, and Alto del Buey and the upper Rfo Truando (Depto. Choco). Also
from Colombia and Ecuador.

443. Grammitis ambigens (Copel.) F. Seym. Phytologia 31:176. 1975.

Ctenopteris ambigens Copel. Phihpp. J. Sci. 84:460, 1. 13. 1956. TYPE: Volc^ Turrialba, near the
Finca del Volcan de Turrialba, Pcia. Cartago, ca. 2000 - 2400 m, Standley 35215 (US).

Ctenopteris knightii Copel. Phihpp. J. Sci. 84:419, t. 6. 1956. TYPE: Las Nubes to La Palma, Pcia.
S. Jose, Knight 29359 (US).

Plants epiphytic, at 1800-2900 m elevation, in forests, from the Cordillera
Central.

In the new genus Ceradenia, the correct name of this species is C. knightii
(Copel.) L. E. Bishop.

444. Grammitis anfractuosa (Kunze ex Klotzsch) Proctor, Rhodora 63:35. 1961.

Polypodium anfractuosum Kunze ex Klozsch, Linnaea 20:375. 1847. TYPE: Merida, Edo. Merida,
Venezuela, Moritz 330 (B not seen; isotype US).

POLYPODIACEAE: 57. GRAMMITIS 307

Polypodium monticola Klotzsch, Linnaea 7PS\7>11. 1847. SYNTYPES: Panao, Villao, Muiia, Depto.
Huanuco, Peru, Ruiz 58 (B not seen); and M6rida, Edo. Merida, Venezuela, Moritz 383 (B not seen

fragm and photo US).

Polypodium saxicola Baker, J. Bot. Brit. For. 15:264. 1877, non Swartz, 1817, nom. illeg. TYPE:
Jamaica, Jenman 84 in 1877 (K not seen).

Polypodium induens Maxon, Bull. Torrey Bot. Club 32:75. 1905. TYPE: A renaming of

Polypodium saxicola Baker, and so based on the type of that name.

Plants epiphytic, at (700)1000-2500 m elevation, in forests, from the Cordillera
de Tilaran, the Cordillera Central, the Cordillera de Talamanca to Pcia. Chiriquf,
near Sta. Fe, and Alto del Buey. Also from Cuba, Jamaica, Hispaniola, Mexico,
Guatemala, Honduras, Venezuela, Colombia to Peru, and Mt. Roraima, Brazil.

445. Grammitis apiculata (Kunze ex Klotzsch) F. Seym. Phytologia 31:176. 1975.

Polypodium apiculatum Kunze ex Klotzsch, Linnaea 20:378. 1847. SYNTYPES: Guyana, Rich.
Schomburgk 1213 (B not seen); and Colonia Tovar, Edo. Aragua, Venezuela, Moritz 247 (B not seen;
isosyntype US).

Polypodium pecten Fee, Gen. Fil. [Mem. Foug. 5]:240. 1852. TYPE: Merida, Edo. Merida,
Venezuela, Linden 239 (P or RB not seen).

Polypodium pectinellum Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 7]:61, t. 27, f. 2.
1857. TYPE: Merida, Edo. Merida, Venezuela, Moritz 239 (P not seen).

IPolypodium confluens Fee, Crypt. Vase. Bresil 1:89, t. 26, f. 3. 1869, non R. Br., 1810, nom. illeg.
TYPE: Brazil, Glaziou 2413 (P or RB not seen).

Plants epiphytic, at 500-1500 m elevation, in forests, from the Cordillera
Central and the valley of the Rfo General. Also from Mexico, Honduras,
Venezuela, Colombia, Peru, and Brazil.

446. Grammitis asplenifolia (L.) Proctor, Brit. Fern Gaz. 9:76. 1962.

Polypodium asplenifolium L. Sp. PL 2:1084. 1753, var. asplenifolia. TYPE: Plate 7, f. 16 of
Petiver’s “Pterigraphia Americana,” which is a transposed redrawing of t. 102A of Plumier’s
“Traite...”, which illustrates a plant collected by Plumier on Martinique. Proctor (Brit. Fern Gaz.
9:76. 1962) correctly excluded Linnaeus’ variety p from the concept of this species; it is G. cultrata.

Polypodium laxifrons Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Aid. V, 1:204
(repr. 52). 1849. LECTOTYPE: Bcirranca de S. Francisco near Mirador, Edo. Veracruz, Mexico,
Liebmann PI. Mex. 2713, FI. Mex. 2321 (C not seen; isolectotype US), chosen by A. R. Smith (FI.
Chiapas 2:120. 1981).

Polypodium repandum Fee, Crypt. Vase. Bresil 1:87, t. 29, f. 1. 1869, non Lour., 1790, nom. illeg.
SYNTYPES: Brazil, Glaziou 2409 (P or RB not seen); and Serra do Couto, Est. Rio de Janeiro,
Brazil, Glaziou 3170 {? or RB not seen).

Polypodium laxifrons var. lividum Kuhn ex Krug in Urban, Bot. Jcihrb. Syst. 24:126. 1897.

LECTOTYPE: Mt. Guaraguao, Adjuntas, Puerto Rico, Sintenis 4328 (US), chosen by Lellinger
(Proc. Biol. Soc. Wash. 89:727. 1977).

Plants epiphytic, at 1400-2100 m elevation, in forests, from La Palma de S.
Ramon, the Cordillera Central, Cerro Carpintera, and the Cordillera de
Talamanca to Pcia. Chiriquf. Also from the Antilles, Mexico to Nicaragua,
Trinidad, Venezuela, Colombia, Ecuador, and Bolivia.

447. Grammitis atroviridis (Copel.) F. Seym. Phytologia 31:176. 1975.

Ctenopteris atroviridis Copel. Philipp. J. Sci. 84:461. 1956. TYPE: Volcan Turrialba, Pcia.

Ciirtago, 3000 m, Torres R. 195 (US).

308 POLYPODIACEAE: 57. GRAMMITIS

456 G. jamesonioides

455 G. isidrensis

454 G. fucoides

POLYPODIACEAE: 57. GRAMMITIS 309

Plants epiphytic, at (700)1000-2300 m elevation, in forests, from Monteverde,
the Cordillera Central, Cerro Carpintera, Tapanti, the Cordillera de Talamanca to
Pcia. Chiriqui, near Sta. Fe, Cerro Campana, and Cana.

448. Grammitis barbensis Lellinger, Proc. Biol. Soc. Wash. 98:379, f. 11. 1985.

TYPE: Volcan Barba, Pcia. Heredia, 3000 m, Valeno 209 (US).

Plants epiphytic, at 3000 m elevation, in forests, known only from the type.

449. Grammitis chrysleri (Copel.) Proctor, Brit. Fern Gaz. 9:218. 1965.

Ctenopteris chrysleri Copel. Philipp. J. Sci. 84:448. 1956. TYPE: Morces Gap to Vinegar Hill,
Jamaica, 3950 - 5000 ft, Maxon 1524 (US).

Plants epiphytic, at 1300-1500(2300) m elevation, in forests, from Tapanti,
Orosi, and Muneco (all Pcia. Cartago), Cana, and Cerro Pirre. Also from
Jamaica, Hispaniola, Venezuela, Colombia, and Bolivia.

450. Grammitis comuta Lellinger, Proc. Biol. Soc. Wash. 98:381, f. 12. 1985.

TYPE: Las Nubes, Pcia. S. Jose, ca. 1500-1900 m, Standley 38843 (US).

Plants epiphytic, at 1500-2300 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Pcia. Chiriqui.

The anastomosing venation doubtless correlates with the wide segments.

451. Grammitis cultrata (Willd.) Proctor, Rhodora 63:35. 1961.

Polypodium asplenifolium var. P L. Sp. PI. 2:1085. 1753. TYPE: Plate 88 of Plumier’s “Traite...”,
which illustrates a specimen collected by Plumier on Martinique. Linnaeus’ citation of the Petiver
reference is in error, and should be Polypodium pendulum subtus pilosum, 1. 12, f. 3.

Polypodium cultratum Willd. Sp. PI. ed. 4, 5:187. 1810. TYPE: Jamaica, Swartz (B-Hb. Willd.
19674 not seen Tryon photo).

Polypodium senile Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 7]:60, t. 25, f. 1. 1857.
TYPE: Ocana, Depto. Norte de Santander, Colombia, Schlim 364 (RB not seen).

Polypodium antioquanum Baker, J. Bot. Brit. For. 19:205. 1881. TYPE: Depto. Antioquia,

Colombia, Kalbreyer 1203 (K not seen photo and fragm US).

Plants epiphytic, at 0-2700 m elevation, in forests, from near S. Ramon, the
Cordillera Central, the Cordillera de Talamanca to Pcia. Chiriqui, Cerro
Tacarcuna, and the Choco. Also from the Greater Antilles, Mexico to El
Salvador, Venezuela, and Colombia to Bolivia.

452. Grammitis epiphytica (Copel.) Lellinger, Proc. Biol. Soc. Wash. 89:714.
1977.

Ctenopteris epiphytica Copel. Philipp. J. Sci. 84:436. 1956. TYPE: La Trojita, Rio Calima, Depto.
El Valle, Colombia, 5-50 m, Cuatrecasas 16723 (US).

Plants epiphytic, at 0 - 100 m elevation, in forests, known only from the type.

FIGS. 445 - 456. Grammitis. FIG. 445. Median portion of lamina of G. apiculata, Skutch 4361. FIG.
446. Median portion of lamina of G. asplenifolia, Lellinger 650. FIG. 447. Median portion of lamina
of G. atroviridis, Lellinger 755. FIG. 448. Median portion of lamina of G. barbensis, Valerio 209.
FIG. 449. Median portion of lamina of G. chrysleri, R. S. Williams 846. FIG. 450. Median portion of
lamina of G. comuta, Maxon 5509. FIG. 451. Median portion of lamina of G. cultrata, Lellinger
1019. FIG. 452. Median portion of lamina of G. epiphytica, Cuatrecasas 16723. FIG. 453. Median
portion of lamina of G. firma. Hatch & Wilson 250, Guatemala. FIG. 454. Median portion of lamina
of G. fucoides, Skutch 3379. FIG. 455. Medium portion of lamina of G. isidrensis, Standley & Valerio
50474. FIG. 456. Median portion of lamina of G. jamesonioides, Mickel 3204.

310 POLYPODIACEAE: 57. GRAMMITIS

453. Grammitis firma (J. Smith) Morton, Contr. U. S. Natl. Herb. 38:110. 1967.

Polypodium firmum Klotzsch, Linnaea 20:378. 1847, non Kaulf., 1827, nom. illeg. LECTOTYPE:
Guyana, Rich. Schomburgk 1170 (B not seen; isolectotype K not seen), chosen by Looser (Rev. Univ.
ChUe 36(1):75. 1951).

Ctenopteris firma J. Smith, Hist. Fil. 184. 1875. TYPE: A renaming of Polypodium firmum
Klotzsch, and so based on the type of that name.

Polypodium aromaticum Maxon, Proc. U. S. Natl. Mus. 27:743. 1904. TYPE: Blue Mountain
Peak, Jamaica, 1950-2225 m, Underwood 1449 (NY not seen fragm US).

Polypodium herzogii Rosenst. Repert. Spec. Nov. Regni Veg. 6:176. 1908. TYPE: Incacorral,
Pcia. Cochabamba, Bolivia, 2200 m, Herzog 783 (S not seen; isotypes UC, US).

Plants epiphytic, at 1000-2500 m elevation, in forests, from the Cordillera
Central. Also from Jamaica, Mexico, Guatemala, Venezuela, Colombia, Peru,
Bolivia, and Guyana.

454. Grammitis fucoides (Christ) Morton, Phytologia 22:78. 1971.

Polypodium fucoides Christ, Bull. Herb. Boissier II, 5:2. 1905. TYPE: Costa Rica, Werckle 172 (P
not seen; isotype US).

Polypodium crassulum Maxon, Contr. U. S. Natl. Herb. 17:598. 1916. TYPE: Tablazo, Pcia. San
Jose, 1900 m, Brade (US fragm NY; isotype UC).

Plants epiphytic, at (1000)1500-2700 m elevation, in forests, from the
Cordillera Central and the north end of the Cordillera de Talamanca.

455. Grammitis isidrensis (Copel.) F. Seym. Phytologia 31:178. 1975.

Ctenopteris isidrensis Copel. Phihpp. J. Sci. 84:441. 1956. TYPE: Yerba Buena, northeast of S.
Isidro, Pcia. Heredia, ca. 2000 m, Standley & Valerio 49881 (US).

Plants epiphytic, at (1000)2000-2500 m elevation, in forests, from La Palma de
S. Ramon (Pcia. Alajuela) and the southern flanks of Volcan Barba.

456. Grammitis jamesonioides (F6e) Morton, Contr. U. S. Natl. Herb. 38:108.
1967.

Polypodium jamesonioides Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 7]:59, t. 21, f.
4. 1857. TYPE: “Provincia Ocanae,” Depto. Norte de Santander, Colombia, Schlim 399 (P or RB
not seen).

1 Ctenopteris nudipes Copel. Philipp. J. Sci. 84:405. 1956. TYPE: Volcan Chiriqui, Pcia. Chiriquf.
10400 ft, Davidson 1022 (US; isotype MO? not seen).

Plants epiphytic, at 2700-3200 m elevation, in forests, from the Cordillera de
Talamanca to Pcia. Chiriquf. Also from Hispaniola, Venezuela, and Colombia to
Peru.

Ctenopteris nudipes is known only from the type, some fronds of which are more
lobed than is typical for G. jamesonioides, but the specimen probably falls within
the range of variation of that species.

457. Grammitis kalbreyeri (Baker in im Thurn) Morton, Amer. Fern J. 60:66.
1970.

Polypodium longipes Fee, Crypt. Vase. Bresil 2:53, t. 95, f. 3. 1873, non Link, 1850, nom. illeg.
TYPE: Serra dos Orgaos, Est. Rio de Janeiro, Brazil, Glaziou 4414 (P or RB not seen).

Polypodium kalbreyeri Baker in im Thurn, Timehri 5:215. 1886. SYNTYPES: Mt. Roraima, Edo.
Bolivar, Venezuela, im Thurn 186 (K not seen); and “Prov. of Ocana,” Depto. Norte de Santander,
Colombia, 6500 ft, Kalbreyer (K not seen).

Polypodium transiens Lindm. Ark. Bot. 1:235, t. 11, f. 7. 1903. TYPE: A renaming of P. longipes
Fee, and so based on the type of that name.

POLYPODIACEAE: 57. GRAMMITIS 311

Polypodium roraimense Brause, Notizbl. Konigl. Bot. Gart. Berlin 6:110. 1914, non Baker, 1887,
nom. illeg. TYPE: Mt. Roraima, Edo. Bolivar, Venezuela, Ule 8520 (B not seen fragm US; isotypes
K not seen Maxon photo, L not seen photo 1984, UC).

Polypodium pressum Brause, Repert. Spec. Nov. Regni Veg. 13:294. 1914. TYPE: A renaming of
P. roraimense Brause, and so based on the type of that name.

Plants epiphytic, at 1000-2300 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, Cerro Tablazo, the upper Rfo Reventazon and
Orosi valleys and Platanillo (all Pcia. Cartago), and around Boquete. Also from
Guatemala, Venezuela, Colombia, Peru, Guyana, Surinam, and Brazil.

This species is polymorphic. Virtually all specimens from Central America have
elongate laminae and slightly ciliate rhizome scales; these match the illustration of
P. longipes Fee. Most specimens from the Guyana Highlands have more debate
laminae and entire rhizome scales. The photograph of the isotype of P. roraimense
at Kew shows that both forms can occur on the same plant.

458. Grammitis lanigera (Desv.) Morton, Contr. U. S. Natl. Herb. 38:105. 1967,
var. lanigera.

Polypodium lanigerum Desv. Ges. Naturf. Freunde Berlin Mag. 5:316. 1811. TYPE: Peru,

Dombey (P not seen Cintract photo).

Polypodium seiiceo-lanatum Hook. Sp. Fil. 4:221. 1864. LECTOTYPE: Ravines of Pichincha,
Pcia. Pichincha, Ecuador, 11000-12000 it, Jameson 235 (K not seen fragm US), chosen by Morton
(Contr. U. S. Natl. Herb. 38:105. 1967).

Polypodium altemifolium Hook. Sp. Fil. 4:222, t. 277A. 1864, non Willd., 1810, nom. illeg.

LECTOTYPE: Near Esmeraldas and between Cuenca and Guayaquil, Ecuador, 3000-10000 ft,
Jameson (K not seen Maxon photo fragm US), chosen by Morton (Contr. U. S. Natl. Herb. 38:105.
1967).

Polypodium longum C. Chr. Ind. Fil. 541. 1906. TYPE: A renaming of Polypodium altemifolium
Hook., and so based on the type of that name.

Ctenopteris fabispora Copel. Philipp. J. Sci. 84:457. 1956, as 'fabaesporaP TYPE: Between Alto
de Las Palmas and the top of Cerro Horqueta, Pcia. Chiriqm, 2100 - 2268 m, Maxon 5479 (US).

Plants epiphytic, at 1200-2700(2900) m elevation, in forests, from the
Cordillera Central and the Cordillera de Talamanca to Pcia. Chiriquf, and the Fila
Costeha near S. Vito de Java. Also from Hispaniola, the Lesser Antilles, and
Colombia to Bolivia.

459. Grammitis lehmanniana (Hieron.) Morton, Contr. U. S. Natl. Herb. 38:104.
1967.

Polypodium lehmannianum Hieron. Bot. Jahrb. Syst. 34:513. 1904. LECTOTYPE: Near

Cuaiquer and S. Pablo, Cordillera de Pasto, Depto. Narino, Colombia, 1000-1300 m, Lehmann 19
(US), chosen inferentially by Maxon (Proc. Biol. Soc. Wash. 57:18. 1944).

Polypodium pastoense C. Chr. Ind. Fil. 551. 1906, nom. superfl. TYPE: A renaming of P.
lehmannianum Hieron., non P. lehmannii Mett., 1857, and so based on the type of P. lehmannianum.

Polypodium sublongipes Christ, Bull. Soc. Bot. Geneve II, 1:218. 1909. TYPE: Costa Rica,

Werckle in 1904 (P not seen), synonymized by Maxon (Proc. Biol. Soc. Wash. 57:18. 1944).

Plants' epiphytic, at 200-1100 m elevation, in forests, from the road from S.
Ramon to Los Angeles (Pcia. Alajuela), the Atlantic lowlands of northern Costa
Rica, the Fortuna reservoir site (Pcia. Chiriquf), near Sta. Fe, and Cerro Campana
(Pcia. Panama). Also from Guatemala and Ecuador.

312 POLYPODIACEAE: 57. GRAMMITIS

466 G. myriophylla

POLYPODIACEAE: 57. GRAMMITiS 313

459a. Grammitis mathewsii (Kunze ex Mett.) Morton, Amer. Fern J. 60:66. 1970.

Polypodium mathewsii Kunze ex Mett. Abh. Senckenberg. Naturf. Ges. 2:74. 1856. SYNTYPES:
Chachapoyas, Depto. Amazonas, Peru, Mathews 1811 (B-Hb. Mett. not seen; isosyntype K? not
seen), 1837 (BM not seen), and 3281 (K not seen).

Goniophlebium villeminianum Fee, M6m. Soc. Hist. Nat. Strasbourg 5 [Mem. Foug. 7]:63, t. 27, f.
3. 1857. TYPE: Ocana, Depto. Norte de Santander, Colombia, 3400 - 3700 m, Schlim 1009 (RB not
seen; isotype BR not seen photo 5043).

Plants epiphytic or occasionally epipetric or terrestrial, in paramos, at
2000-2800 m elevation, in the Flora area known only from the Interamerican
Highway 21-22 km southeast of El Empalme, Pcia. S. Jose {Smith & Belt 1993,
UC) and from Cerro Punta, Pcia. Chiriqui (van der Werjf & Herrera 6252, UC).
Also from Colombia to Bolivia.

460. Grammitis mayoris (Rosenst.) Lellinger, Amer. Fern J. 74:59. 1984.

Polypodium mayoris Rosenst. Mem. Soc. Sci. Nat. Neuchatel 5:53, t. 4, f. 6. 1912. TYPE: Paramo
del Ruiz, Depto. Antioqeia, Colombia, ca. 3700 m. Mayor 69 (S not seen; isotype US).

Plants epiphytic, at 2000-3400 m elevation, in forests, from Volcan Turrialba
and the Cordillera de Talamanca to Pcia. Chiriqui. Also from Colombia.

461. Grammitis melanosticta (Kunze) F. Seym. Phytologia 31:179. 1975.

Polypodium melanostictum Kunze, Linnaea 9:44. 1834. TYPE: Cuesta de Carpis, Depto.

Huanuco?, Peru, Poeppig in 1829 (LZ destroyed; isotype W not seen).

Polypodium calvum Maxon, J. Wash. Acad. Sci. 12:440. 1922. TYPE: Sierra Maestra, Pcia.
Oriente, Cuba, 11131 (US).

Plants epiphytic, at 1200-1500 m elevation, in forests, in the Flora area known
only from above Platanillo, ca. 22 km east of Turrialba, Pcia. Cartago, 1200-1450
m {Michel 3394, NY, US). Also from Cuba, Hispaniola, Guatemala, Venezuela,
Colombia, Peru, and Bolivia.

462. Grammitis meridensis (Klotzsch) F. Seym. Phytologia 31:179. 1975.

Polypodium meridense Klotzsch, Linnaea 20:380. 1847. TYPE: Merida, Edo. Merida, Venezuela,
Moritz 335 (B not seen).

Polypodium spixianum Mart, ex Mett. Abh. Senckenberg. Naturf. Ges. 2:57. 1856. TYPE

LOCALITY: Brazil. “P. nidulans Beyr. herb.” is cited in synonymy.

Polypodium radicale Moritz ex Baker, Syn. Fil. 332. 1867. SYNTYPES: Southern Brazil, Gardner
128 (B or K not seen); and Venezuela, Fendler 216 (B or K not seen).

Polypodium meridense f. undulatum Hieron. Bot. Jahrb. Syst. 34:519. 1904, as “undulata.” TYPE:
Mt. Bordoncillo near Pasto, Depto. Nariho, Colombia, 3000 m, Lehmann 653 (B not seen).

Plants epiphytic, at 1500-3300 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Pcia. Chiriqui. Also from Hispaniola,
El Salvador, Venezuela, Colombia, Peru, Bolivia, and Brazil.

FIGS. 457 - 466. Grammitis. FIG. 457. Median portion of lamina of G. kalhreyeri, Lellinger 1063.
FIG. 458. Median portion of lamina of G. lanigera var. lanigera, Maxon 8168. FIG. 459. Median
portion of lamina of G. lehmanniana, Contreras 4781, Guatemala. FIG. 460. Median portion of
lamina of G. mayoris, Evans & Lellinger 168. FIG. 461. Plant of G. melanosticta, Mickel 3394. FIG.
462. Median portion of lamina of G. meridensis, Evans & Lellinger 65. FIG. 463. Median portion of
lamina of G. micula, Skutch 3015. FIG. 464. Median portion of lamina of G. molhssima, Pittier 4479.
FIG. 465. Plant of G. moniliformis, L. O. Williams 20054. FIG. 466. Median portion of lamina of G.
myriophylla, Gomez et al. 21892.

314 POLYPODIACEAE: 57. GRAMMITIS

463. Grammitis micula Lellinger, Proc. Biol. Soc. Wash. 98:381, f. 13. 1985.

TYPE: Vicinity of S. Isidro del General, 1000 m, Skutch 3015 (US; isotype UC).
Plants epiphytic, at 1000-1200(1800) m elevation, in forests, known only from
the type and possibly from Alto del Buey {Lellinger & de la Sota 261, US).

This species was thought to be Grammitis hombersleyi (Maxon) Lellinger, a
species from Trinidad differing in rhizome scales and segment shape.

464. Grammitis mollissima (F6e) Proctor, Rhodora 63:35. 1961.

Polypodium mollissimum Fee, Hist. Foug. Antill. [Mem. Foug. 11]:47, t. 12, f. 2. 1866. TYPE:
Guadeloupe, UHerminier in 1861 (P not seen fragms NY and US),

Polypodium cultratum var. elasticum Jenm. Bull. Bot. Dept. Jamaica N.S., 4:119. 1897. TYPE:
Presumably Jamaica, (NY? not seen).

Plants epiphytic, at 100-2600 m elevation, in forests, from the Cordillera
Central, the Cordillera de Talamanca probably to Pda. Chiriqm, Barro Colorado
Island (Canal Zone), the upper Mamoni River (Pcia. Panama), and above Cana.
Also from the Antilles, Mexico to Honduras, Colombia, and Venezuela to French
Guiana.

465. Grammitis moniliformis (Lag. ex Swartz) Proctor, Brit. Fern Gaz. 9:219.
1965.

Polypodium moniliforme Lag. ex Swartz, Syn. Fil. 33. 1806. TYPE: Peru, collector unknown (S not
seen photo 6020).

Polypodium subcrenatum Hook. Icon. Plant. 8:t. 719. 1845. TYPE: Andes of Quito, Pcia.

Pichincha, Ecuador, Jameson 215 (K not seen; isotype FI not seen photo 16012).

Polypodium moniliforme var. major Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk.
Afd. V, 1:1% (repr. 44). 1849. LECTOTYPE: Pico de Orizaba, Edo. Veracruz, Mexico, Liebmann
PI Mex. 2533 (C not seen), chosen by A. R. Smith (FI. Chiapas 2:123. 1981).

Jamesonia adnata Kunze, Farrnkrauter 2:80, t. 133, f. 1. 1851. TYPE: Paramo de Tolima, Depto.
Tolima, Colombia, Linden 1006 (LZ destroyed; isotypes B not seen Tryon photo, FI not seen photo
16010).

Polypodium angustissimum Fee, Crypt. Vase. Bresil 2:55, t. %, f. 3. 1873. TYPE: Brazil, Glaziou
5297 (P or RB not seen).

Polypodium subdicarpon Fee, Crypt. Vase. Bresil 2:55, t. %, f. 4. 1873. TYPE: Brazil, Glaziou
4410 (P or RB not seen).

Polypodium moniliforme var. bogotense Hieron. Hedwigia 48:249. 1909, as ''bogotensis P TYPE:
Bogota region, Depto. Cundinamarca, Colombia, Stuebel 431 p. p. (B not seen).

Polypodium moniliforme var. minus Christ, Bull. Soc. Bot. Geneve II, 1:217. 1909. SYNTYPES:
Crater of Volcan Irazu, 3380 m, Pittier 179 (P not seen; isotype US); and Serra do Itatiaia, Est. Rio
de Janeiro, Brcizil, 2200 m, Ule 3785 (P not seen).

Polypodium moniliforme var. culebriliense Bosco, Nuovo Giorn. Bot. Ital. N.S., 45:149. 1938.

TYPE: Paramo de Culebrilla, Pcia. Chimborazo?, Ecuador, 3500 m, Crespi (TO not seen).

Plants epiphytic or rarely terrestrial or on fallen logs, at (600)2000-3400(3600)
m elevation, in forests, from the Cordillera Central, the Cordillera de Talamanca
to Pcia. Chiriquf, and north of Cerro Torra, west of the Rfo Surama, Depto. Choco
{Forero 3159, MO). Also from Jamaica, Hispaniola, Mexico to Honduras,
Venezuela, Colombia to Bolivia, and Brazil.

466. Grammitis myriophylla (Mett. ex Baker) Morton, Contr. U. S. Natl. Herb.
38:108. 1867.

IPolypodium longisetosum Hook. Sp. Fil. 4:225. 1864. TYPE: Andes of Quito, Pcia. Pichincha,
Ecuador, Jameson 79 (K not seen).

POLYPODIACEAE: 57. GRAMMITIS 315

Polypodium myriophylium Mett. ex Baker, Syn. Fil. 338. 1868. TYPE: Tatanera, Depto. Puno,
Peru, Lechler 2567 (K not seen; isotype L not seen fragm US).

Plants epiphytic, at 2500-3000 m elevation, in forests, in the Flora area known
only from the upper Rio Colubre, Pcia. Bocas del Toro {Gomez et al 21892,
21935, both CR not seen, MO). Also from Ecuador, Peru, and Bolivia.

This species is most unusual in Grammitis in having pinnate-pinnatifid laminae.

467. Grammitis nuMgena (Maxon) Proctor, Brit. Fern Gaz. 9:219. 1965.

Polypodium nubigenum Maxon, Contr. U. S. Natl. Herb. 17:599. 1916. TYPE: Blue Mountain
Peak, St. Thomas Parish, Jamaica, 2200 m, Maxon 1477 (US).

Plants epiphytic, at 2500 m elevation, in the Flora area known only from Volcan
Barba {Valerio 53, CR, US). Also from Jamaica.

467a. Grammitis pMegmaria (J. Smith) Proctor, Rhodora 68:467. 1966, van

phlegmaria.

Polypodium phlegmaria J. Smith, London J. Bot. 1:195. 1842. TYPE: Near Mt. Roraima, Edo.
Bolivar, Venezuela, Schomburgk 161 (K not seen Maxon photo).

Polypodium tovarense Klotzsch, Lirmaea 20:374. 1847. TYPE: Colonia Tovar, Edo. Aragua,
Venezuela, Moritz 251 (B not seen; isotype L not seen photo 1948).

Polypodium subdimidiatum Baker, Syn. Fil. 324. 1867. SYNTYPES: Venezuela, Fendler 207 (K
not seen); Andes [Pcia. Pichincha?], Ecuador, Jameson 2122 (K not seen); and Guyana, Appun 1130
(K not seen).

Plants epiphytic, at 2500-2800 m elevation, in forests, in the Flora area known
only from the Pan American Hwy. 21-22 km southeast of El Empalme, Pcia. S.
Jose {Smith & Beliz 2080 and 2104, both UC). Also from Venezuela, Colombia,
and Pern,

Proctor (Rhodora 68:467. 1966) distinguished Lesser Antillean material as var.
antiliana, which differs from continental material in having narrower and more
oblong but less gibbous (almost entire) segments.

467b. Grammitis pilipes (Hook.) Morton, Phytologia 22:77. 1971.

Polypodium pilipes Hook. Icon. PI. 3:t. 221. 1840. TYPE: Chachapoyas, Depto. Amazonas, Peru,
Mathews in 1838 (K not seen photo 15425).

Polypodium blandum Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg [Mem. Foug. 7]:59, t. 22, f. 5.
1857. TYPE: South America, collector unknown (RB not seen).

Polypodium decipiens Hook. Sp. Fil. 4:231, t. 279B. 1862. TYPE: ‘‘Colombia” [Venezuela],
Moritz 337 (K not seen; isotype US not seen), examined by Maxon (in herb.) and found to be the
basis of Hooker’s illustration.

Polypodium pozuzoense Baker, Icon. PI 17:t. 1672. 1886 - 87. TYPE: Pozuzo, Pcia. Huanuco,
Peru, 80CM3 ft, July 1863, Pearce (K not seen photo 15429).

Plants epiphytic, at 2800-2900 m elevation, in forests, in the Flora area known
only from Volcan Barba {Grayum & Quesada 7420, MO). Also from Venezuela
and Pern.

468. Grammitis pilosissima (Martens & Galeotti) Morton, Contr. U. S. Natl.
Herb. 38:114. 1967.

Polypodium pilosissimum Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:39, t. 9, f.
2. 1842. LECTOTYPE: Zacuapan, Edo. Veracruz, Mexico, Galeotti 6379 (BR not seen photo 5017
fragm US; isolectotype K not seen), chosen by A. R. Smith (FI. Chiapas 2:124. 1981).

Polypodium pilosissimum var. glabriusculum Mett. Abh. Senckenberg. Naturf, Ges. 2:42. 1856, as
“glabriusculaP SYNTYPES: Mexico, Leibold 97 (B not seen); Merida, Edo. Merida, Venezuela,

316 POLYPODIACEAE: 57. GRAMMITIS

471 G. rigescens

473 G. semihirsuta

470 G. randallii

POLYPODIACEAE: 57. GRAMMITIS 317

Moritz 216 (B not seen; isosyntype US); and Caracas, Distr. Fed., Venezuela, Funck & Schlim 965 (B

not seen).

Polypodium piiosissimum var. hirsutum Mett. Abh. Senckenberg. Naturf. Ges. 2:42. 1856, as
'"hirsutaP TYPE: Venezuela, Funck & Schlim 1848 (B not seen).

IPolypodium ieptophyllum Moritz ex Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:250 (repr.
332). 1864, non L., 1752, nom. illeg. SYNTYPES: Sosiego, Colombia, 2400 - 2600 m, Lindig 300 (B
not seen); Colonia Tovar, Edo. Aragua, Venezuela, Moritz 437 p. p. (B not seen; isosyntype L not
seen photo 1830), Fendier 217 (B not seen); Peru, Lechler (B not seen); Guyana, Schomburgk (B not
seen). The Moritz specimen is G. apiculata, and does not agree with the original description.

Ctenopteris megaloura Copel. Philipp. J. Sci. 84:391. 1956. TYPE: Coban, Depto. Alta Verapaz,
Guatemala, 1350 m, von Tuerckheim II 1855 (US).

Plants epiphytic or epipetric, at 2900-3500 m elevation, in forests and on trees
in pastures, from Cerro de la Muerte (Pda. Cartago), Cerro Chirripo, and Volcan
Chiriqui. Also from Mexico and Guatemala.

This species is known only from high elevations. Rather similar specimens from
lower elevations (1200-2500 m) are G. xiphopteroides.

469. Grammitis podocarpa (Maxon) F. Seym. Phytologia 31:180. 1975.

Polypodium podocarpum Maxon, Smithsonian Misc. Collect. 56(24):2, t. 1-3. 1911. TYPE:

Upper Caldera watershed between Camp I and the Divide, Holcomb’s trail above Boquete, Pcia.
Chiriqui, 1650-1925 m, Maxon 5640 (US; isotype NY).

Plants epiphytic, at 1600-2200 m elevation, in forests, from the Cordillera de
Talamanca above Boquete.

469a. Grammitis pruinosa (Maxon) Morton, Contr. U. S. Natl. Herb. 38:262.
1973.

Polypodium pminatum Baker, Syn. Fil. ed. 2. 508. 1874, non Swartz, 1802, nom. illeg. TYPE:
Chontales, Depto. Chontales, Nicaragua, Tate 44 (K not seen photo 15430 fragm US).

Polypodium pminosum Maxon, Proc. Biol. Soc. Wash. 52:117. 1939. TYPE: A renaming of P.
pminatum Baker, and so based on the type of that name.

Grammitis kirkbridei Mickel, Amer. Fern J. 74:117, f. 3C. 1984. TYPE: Tres Bocas on the Rio
Cuasi, Pcia. Darien, Panama, Kirkbride & Duke 1381 (NY not seen; isotype MO).

Plants epiphytic, at ca. 100-1000 m, in forests, in the Flora area known only
from Cerro Jefe and from Tres Bocas on the Rio Cuasi (Pcia. Darien). Also from
Nicaragua and Venezuela.

470. Grammitis randallii (Maxon) Proctor, Brit. Fern Gaz. 9:219. 1975.

Polypodium randallii Maxon, Amer. Fern J. 18:46. 1928. TYPE: Crest of Gossamer Peak, St.
Thomas Parish, Jamaica, 800 - 900 m, Maxon 9280 (US fragm NY; isotype IJ not seen).

Plants epiphytic, at 1000-1400 m elevation, in forests, from Cerro Jefe, Cerro
Mali (Pcia. Darien), and east of the Alturas de Nique (Pcia. Darien). Also from
Jamaica.

This species looks very much like a diminutive specimen of G. barbensis, but
differs in its bifid to stellate stipe hairs.

FIGS. 467 - 474. Grammitis. FIG. 467. Median portion of lamina of G. nubigena, Valerio 53. FIG.
468. Median portion of lamina of G. pilosissima, Maxon 5341. FIG. 469. Median portion of lamina
of G. podocarpa, Maxon 5460. FIG. 470. Median portion of lamina of G. randallii, Gentry & Mori
13811. FIG. 471. Median portion of lamina of G. rigescens, Pittier 10464. FIG. 472. Median portion
of lamina of G. sectifrons, Skutch 3298. FIG. 473. Median portion of lamina of G. semihirsuta,
Mickel 3200. FIG. 474, Median portion of lamina of G. staheliana, Lellinger 1319.

318 POLYPODIACEAE: 57. GRAMMITIS

471. Grammitis rigescens (Boiy ex Willd.) Lellinger, Proc. Biol. Soc. Wash.
89:383. 1985.

Polypodium rigescens Bory ex Willd. Sp. PI. ed. 4, 5:183. 1810. TYPE: Reunion, Bory (probably
P-Hb. Juss. 1098-C not seen photo 2951).

Polypodium rigescens var. major Hieron. Bot. Jahrb. Syst. 34:503. 1904. TYPE: Mt. Ruiz, Depto.
Caldas or Tolima, Colombia, 3000 m, Apr 1882, Schmidtchen (B not seen).

Plants epiphytic, at 2100-3400 m elevation, from between Cerro Gallito and
Volcdn Barba (Pcia. Heredia), and Pdramo Buena Vista, Villa Mills, and vicinity
(Pcias. S. Jose and Cartago). Also from Hispaniola, Venezuela, Colombia to Peru,
and Brazil.

472. Grammitis sectifrons (Kunze ex Mett.) F. Seym. Phytologia 31:180. 1975.

Polypodium sectifrons Kunze ex Mett. Abh. Senckenberg. Naturf. Ges. 2:99, t. 2, f. 3-4. 1856.
LECTOTYPE: Puerto Rico, Schwanecke (GH), chosen by Proctor (Ferns Jamaica 585. 1985).

Drynaria elastica Fee, Hist. Foug. Antill. [Mem. Foug. 11]:72, t. 20, f. 2. 1866, non Polypodium
elasticum Bory ex Willd., 1810. TYPE: Guadeloupe, UHerminierl (RB not seen).

Polypodium petraefolium Jenm. Bull. Bot. Dept. Jamaica N.S., 4:139. 1897. TYPE: A renaming
of Drynaria elastica Fee, and so based on the type of that name.

Plants epiphytic, at 1000-2500 m elevation, in forests, from the Cordillera
Central, the Reventazon and Orosi valleys (Pcia. Cartago), and the Cordillera de
Talamanca to Pcia. Chiriquf. Also from Jamaica, Hispaniola, Puerto Rico, the
Lesser Antilles, Venezuela, and Colombia.

473. Grammitis semihirsuta (Klotzsch) Morton, Contr. U. S. Natl. Herb. 38:113.
1967, var. semihirsuta.

Polypodium semihirsutum Klotzsch, Linnaea 20:379. 1847. TYPE: “Panatahuas Provinciae,” Peru,
Ruiz 9 (B not seen fragm NY; isotype US).

Polypodium leucolepis Gilbert, Bull. Torrey Bot. Club 24:260. 1897. TYPE: Blue Mountains,
Jamaica, Moore (GH not seen).

Polypodium pectinatum var. hispidum Christ, Prim. FI. Costaric. 3(1): 15. 1901, as ''hispida.”

TYPE: El Paramo, Buena Vista massif, Pcia. Cartago, 3000 m, Pittier 10474 (BR or P not seen;
isotypes CR, US).

Plants epiphytic, at (1100)3000-3500 m elevation, in forests, from La Palma de
S. Ramon, the Cordillera de Talamanca to Pcia. Chiriquf, and Alto del Buey. Also
from Jamaica, Hispaniola, Mexico to El Salvador, Venezuela, and Colombia to
Bolivia.

Specimens with the costae and laminae setose abaxially have been called G.
semihirsuta var. fuscosetosa (Hieron.) Lellinger. Although Hieronymus (Bot.
Jahrb. Syst. 34:515. 1904) ascribed this variety to Guatemala, Colombia, and Peru,
I have found only specimens from the highlands of Colombia to be referable to it.

474. Grammitis staheliana (Posth.) Lellinger, Proc. Biol. Soc. Wash. 89:716. 1977.

Polypodium stahelianum Posth. Recueil Trav. Bot. Neerl. 23:401. 1928. TYPE: Emmaketen,
Surinam, 700 m, Gonggrijp & Stahel 5765 (U not seen photo 23).

Plants epiphytic, at (100)600-1200 m elevation, in forests, from west of S.
Ramon, Loma Prieta (Pcia. Los Santos), above Penonome, near Cerro Jefe, and
the upper Rfo Truando. Also from Nicaragua, Venezuela, Colombia, and
Surinam.

POLYPODIACEAE: 57. GRAMMITIS 319

475. Grammitis subcapillaris (Christ) F. Seym. Phytologia 31:180. 1975.

Polypodium subcapillare Christ, Bull. Herb. Boissier II, 5:1. 1905. TYPE: La Palma, Werckle
17033 (P not seen fragm and photo US).

Plants epiphytic, at 500-2000 m elevation, in forests and on trees in open areas,
from the Cordillera de Tilaran, the Cordillera Central, the vicinity of Pejivalle
(Pcia. Cartago), La Mesa (Pcia. Code), the Canal Zone, Cerro Jefe, and Cerro
Tacarcuna and the Alturas de Nique (Pcia. Darien).

476. Grammitis subsessilis (Baker) Morton, Contr. U. S. Natl. Herb. 38:107.
1967.

Polypodium pteropus Hook. Sp. Fil. 4:192, t. 275B. 1862, non Blume, 1828, nom. illeg.

SYNTYPES: Andes of Quito, Pcia. Pichincha, Ecuador, Jameson 348 (K not seen). Spruce 5712 (K
not seen); Mt. Abitagua, Pcia. Tungurahua?, Ecuador, Spmce (K not seen); Mt. Roraima, Edo.
Bolivar, Venezuela, Schomburgk (K not seen); and “New Grenada,” Hartweg 1495 (K not seen).

Polypodium subsessile Baker, Syn. Fil. 329. 1867. TYPE: A renaming of Polypodium pteropus
Hook., and so based on the type of that name.

Polypodium euchlomm Kunze ex Hieron. Bot. Jahrb. Syst. 34:509. 1904, non Sodiro, 1883, nom.
superfl. et illeg. TYPE: Although ostensibly a validation of the nomen nudum P. euchlomm Kunze
ex Klotzsch (Linnaea 20:375. 1847), the name must be based on P. subsessile Bak. in Hook. & Bak.,
which was cited in synonymy.

Polypodium chiricanum Maxon, Contr. U. S. Natl. Herb. 17:597. 1916. TYPE: Between Alto de
las Padmas and Cerro de la Horqueta, Pcia. Chiriqm, 2100 - 2268 m, Maxon 5478 (US).

Polypodium pendulum var. boliviense Rosenst. Repert. Spec. Nov. Regni Veg. 25:60. 1928.

SYNTYPES: S. Carlos, Mapiri region, Depto. La Paz, Bohvia, 850 m, Buchtien 210 (S not seen;
isosyntype US); amd Hacienda Simaco above the road to Tipuami, Depto. La Paiz, Bolivia, 1400 m,
Buchtien 5256 (S not seen; isosyntype US).

Ctenopteris obovata Copel. Philipp. J. Sci. 84:442. 1956. TYPE: A renaming of P. pendulum var.
boliviense Rosenst., amd so based on the type of that name.

Plants epiphytic, at 200-2400 m elevation, in forests, from the Cordillera
Central, the Cordillera de Talamanca around Boquete, west of Sta. Fe, La Mesa
(Pcia. Code), Cerro Jefe, Cerro Pirre, the Serranfa de Darien, the Cerro de
Garagara (Pcia. Darien), and the northern half of the Choco. Also from
Venezuela and Colombia to Bolivia.

The specimens from Pcia. Darien have appreciably narrower and more distant
segments than do those from the cordilleras farther north.

477. Grammitis subtilis (Kunze ex Klotzsch) Morton, Contr. U. S. Natl. Herb.
38:104. 1967.

Polypodium subtile Kunze ex Klotzsch, Linnaea 20:375. 1847. TYPE: Merida, Edo. Merida,
Venezuela, Moritz 325 (B not seen fragm NY; isotype US).

Plants epiphytic, at 1000-2400 m elevation, in forests, from the Cordillera
Central, Cerro Carpintera, Tapanti, the vicinity of S. Isidro del General, and
around Boquete. Also from Mexico, Guatemala, Venezuela, Colombia, and
Ecuador.

478. Grammitis suprasculpta (Christ) F. Seym. Phytologia 31:181. 1975.

Polypodium suprasculptum Christ, Bull. Herb. Boissier II, 5:3. 1905. SYNTYPES: Volcan

Turrialba, Pcia. Cartago, 2400 m, Pittier 14145 (P not seen); and Costa Rica, Werckle (P not seen).

Plants epiphytic, at 1200-2400 m elevation, in forests, from La Palma de S.
Ramon, the Cordillera Central, Alto de La Estrella (Pcia. Cartago), and Boquete.

320 POLYPODIACEAE: 57. GRAMMITIS

POLYPODIACEAE; 57. GRAMMITIS 321

479. Grammitis suspense (L.) Proctor, Brit. Fern Gaz. 9:77. 1962.

Polypodium suspensum L. Sp. PI. 2:1084. 1753. TYPE: Plate 87, right hand figure, of Plumier’s
“Traite...”, which illustrates a specimen collected by Plumier on Martinique.

Polypodium jubiforme Kaulf. Flora (Regensburg) 6:364. 1823, as “iubaeformeP TYPE:

Mairtinique, Sieber FI. Mart. 3S3 (PRC? not seen; isotypes FI not seen photo 15988, GH not seen).

Polypodium confusum J. Smith, Lx)ndon J. Bot. 1:194. 1842. TYPE: Guyana, Schomburgk 51 (K
not seen fragm US).

Polypodium saccatum Fee, Gen. Fil. [Mem. Foug. 5]:239. 1852. SYNTYPES: Guadeloupe,

Perrottet (P or RB not seen); and Martinique, Rivoire (P or RB not seen).

Polypodium jubiforme var. delicatulum Christ in Krug in Urban, Bot. Jahrb. Syst. 24:126. 1897.
TYPE: Guadeloupe, Maze 1088 (P not seen).

Polypodium suspensum f. minor Kuhn ex Krug in Urban, Bot. Jahrb. Syst. 24:127. 1897. TYPE:
Puerto Rico, Sintenis 420 (B not seen).

Ctenopteris rhizophorae Copel. Philipp. J. Sci. 84:425. 1956. TYPE: Along the Rio Dagua,
Buenaventura, Depto. El Valle, Colombia, Killip 5334 (US).

Plants epiphytic, at 200-1500 m elevation, in forests, from the lower Rio S.
Juan basin, and Alto del Buey. Also from the Antilles, Trinidad, Venezuela,
Colombia, Guyana, and Surinam.

480. Grammitis taxifolia (L.) Proctor, Rhodora 63:35. 1961.

Polypodium taxifolium L. Sp. PI. 2:1086. 1753. TYPE: Plate 89 of Plumier’s “Traite...”, which
illustrates a specimen collected by Plumier on Martinique.

Polypodium Vherminieri Fee, Gen. Fil. [Mem. Foug. 5]:238. 1852. TYPE: Guadeloupe, Perottet (P
or RB not seen).

Polypodium Vherminieri var. costaricense Rosenst. Repert. Spec. Nov. Regni Veg. 22:17. 1925.
TYPE: La Palma, 1400 m, Brade & Brade 216 (S not seen photo 6086; isotypes UC, US).

Plants epiphytic, at 1000-1800(3000) m elevation, in forests, from north of S.
Ramon and the Cordillera Central. Also from Hispaniola, Puerto Rico, the
Lesser Antilles, Trinidad, Venezuela, Ecuador, Surinam, and Brazil.

481. Grammitis tmesipteris (Copel.) F. Seym. Phytologia 31:181. 1975.

Ctenopteris tmesipteris Copel. Phihpp. J. Sci. 84:410. 1956. TYPE: Cerro de Las Vueltas, Pcia. S.
Jose, 3000 m, Standley 43755 (US).

Plants epiphytic, at 2700-3300 m elevation, in forests, from the Cordillera de
Talamanca.

482. Grammitis turrialbae (Christ) F. Seym. Phytologia 31:181. 1975.

Polypodium turrialbae Christ in Dur. & Pitt. Bull. Soc. Roy. Bot. Belgique 35: Mem. 226. 1896.
TYPE: Volc^ Turrialba, Pcia. Cartago, 2750 m, Pittier 847 (BR not seen photo 5042; isotype US).

1 Polypodium crispulum Christ, Bull. Herb. Boissier II, 4:1002. 1904. TYPE: Volcan Turrialba,
Pcia. Cartago, 1400 m, Pittier 14147 (P not seen).

IPolypodium exsudans Christ, Bull. Herb. Boissier II, 4:1103. 1904. TYPE: Costa Rica, Werckle in
1903 (P not seen).

FIGS. 475-485. Grammitis. FIG. 475. Median portion of lamina of G. subcapillaris, Standley &
Valerio 47022. FIG. 476. Frond of G. subsessilis, Standley & Valerio 52254. FIG. 477. Median
portion of lamina of G. subtilis, Kilhp 5289. FIG. 478. Median portion of lamina of G. suprasculpta,
Mickel 3027. FIG. 479. Median portion of lamina of G. suspensa, Kilhp 35302. FIG. 480. Median
portion of lamina of G. taxifolia, Skutch 3751. FIG. 481. Median portion of lamina of G. tmesipteris,
Evans & LeUinger 171. FIG. 482. Mediam portion of l2miina of G. turrialbae, Skutch 3421. FIG. 483.
Median portion of l2unina of G. zeledoniana, Maxon 8148. FIG. 484. Plant of G. jungermannioides,
Maxon 8163. FIG. 485. Plant of G. spmcei, Skutch 2832.

322 POLYPODIACEAE: 57. GRAMMITIS

Plants epiphytic, at 1500-2800 m elevation, in forests, from the Cordillera
Central, and the Cordillera de Talamanca at Madre de Selva (Pcia. Cartago) and
near the Panama border (Pcia. Limon).

482a. Grammitis xiphopteroides (Liebm.) A. R. Smith, Amer. Fern J. 70:26. 1980.

Polypodium xiphopteroides Liebm. K. Dankse Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:196
(repr. 44). 1849. LECTOTYPE: Hacienda de Mirador, Edo. Veracruz, Mexico, Liebmann PI. Mex.
2548 (C not seen), chosen by A. R. Smith (FI. Chiapas 2:125. 1981).

Polypodium rigens Maxon, Proc. U. S. Natl. Mus. 27:741. 1904. TYPE: John Crow Peak, Jamaica,
1650-1800 m, Maxon 1346 (US).

Ctenopteris vemicosa Copel. Philipp. J. Sci. 84:452, t. 9. 1956. TYPE: Alto de Estrella, Pcia.
Cartago, Standley 39140 (US), indicated as type by Copeland on the specimen, but not in the
publication.

Plants epiphytic, at 1200-2500 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, the Cordillera de Talamanca to Pcia. Chiriqui, the
Fila Costeha near S. Vito de Java, and Cerro Jefe (Pcia. Panama). Also from
Cuba, Jamaica, Hispaniola, Mexico, Guatemala, Honduras, Venezuela, Colombia
to Peru, Surinam, and Brazil.

483. Grammitis zeledoniana Lellinger, Proc. Biol. Soc. Wash. 89:383. 1985.

Polypodium taxifolium var. fragillimum Christ, Bull. Herb. Boissier II, 4:1103. 1904, non

Grammitis fragillima (Copel.) Morton, 1973. TYPE: Costa Rica, Werckle in 1903 (P not seen).

TYPE: A renaming of Polypodium taxifolium var. fragillimum Christ, and so
based on the type of that name.

Plants epiphytic, at (1000)1500-2500 m elevation, in forests, from the
Cordillera Central and the Cordillera de Talamanca to Pcia. Chiriqui.

GRAMMITIS sect. GRAMMITASTRUM (Fourn.) Morton

484. Grammitis jungermannioides (Klotzsch) Ching, Bull. Fan Mem. Inst. Biol.
Bot. 10:15. 1940.

Polypodium jungermannioides Klotzsch, Linnaea 20:373. 1847. TYPE: Merida, Edo. Merida,
Venezuela, Moritz 312 (B not seen; isotype BM not seen).

Grammitis repanda Kunze ex Mett. Fil. Lechl. 1:9. 1856, nom. superfl. TYPE: A renaming of P.
jungermannioides Klotzsch, and so based on the type of that name.

Polypodium sprucei var. furcativenosa Hieron. Bot. Jahrb. Syst. 34:499. 1904. SYNTYPES: Rio
Balsayacu near Pasto, Depto. Narino, Colombia, Lehmann 654 (B not seen; isosyntype US); and
near Chilascb, Depto. Baja Verapaz, Guatemala, Salvin & Godman (K not seen).

Polypodium sprucei var. costaricensis Christ, Repert. Spec. Nov. Regni Veg. 8:17. 1910. TYPE:
Volc^ Barba, Pcia. Heredia, 2200 m, Brade & Brade 296 (P? not seen; isotypes NY not seen, UC).

Plants epiphytic, at 600-2700 m elevation, in forests, from the Cordillera
Central, Tapantf, the Cordillera de Talamanca to Pcia. Chiriqui, and S. Isidro del
General. Also from Mexico to Honduras, Venezuela, and Colombia.

485. Grammitis sprucei (Hook.) J. Smith, Hist. Fil. 181. 1875.

Polypodium sprucei Hook. Sec. Cent. Ferns t. 10. 1860. TYPE: Near Tarapoto, Depto. S. Martin,
Peru, Spruce 4746 (K not seen; isotype US).

Polypodium dendricola Jenm. Card. Chron. Ill, 16:467. 1894, as ''dendricolum.'' TYPE: Port
Royal Mountains, Jamaica, Hart (IJ not seen; isotype NY not seen).

Polypodium rosulatum Christ in Bommer & Christ, Bull. Herb. Boissier 4:662. 1896. TYPE: Rio
Naranjo, Costa Rica, Pittier 7953 (BR not seen photo 5027).

POLYPODIACEAE: 57. GRAMMITIS 323

Polypodium yarumalense Hieron. Bot. Jahrb. Syst. 34:499. 1904. TYPE: Near Yarumal, Depto.
Antioquia, Colombia, 2000-2200 m, Lehmann 7390 (B not seen; isotype US).

Plants epiphytic, at 1200-2300 m elevation, in forests, from the Cordillera
Central, Platanillo (Pcia. Cartago), and the Cordillera de Talamanca to Pcia.
Chiriqui. Also from Jamaica, Hispaniola, Colombia, and Peru.

GRAMMITIS sect. GRAMMITIS

486. Grammitis bryophila (Maxon) F. Seym. Phytologia 31:172. 1975.

Polypodium byrophilum Maxon, Amer. Fern J. 16:7. 1926. TYPE: Above La Palma on the road to
La Hondura, Pcia. S. Jose, 1700 m, Maxon & Harvey 7980 (US).

Polypodium haplophlehicum A. C. Smith, Bull. Torrey Bot. Club 58:37. 1931. TYPE: Mt. Duida,
Terr. Fed. Amazonas, Venezuela, Tate 5S3 (NY not seen; isotype US).

Plants epiphytic, at 1400-2700 m elevation, in forests, from the Cordillera
Central, the Cordillera de Talamanca to Villa Mills (Pcia. Cartago), and Cerro
Jefe. Also from Venezuela, Colombia, and Bolivia.

487. Grammitis bufonis G6mez, Phytologia 52:154. 1982.

TYPE: Cerro Sapo, Pcia. Darien, 1085 m, Hammel 1171 (MO).

Plants epiphytic, at 1000-1100 m elevation, in forests, from the Fortuna dam
site (Pcia. Chiriqui), the hills north of El Valle, Cerro Jefe, and Cerro Sapo (Pcia.
Darien).

488. Grammitis leptopoda (C. H. Wright) Copel. Philipp. J. 80:255. 1952.

Polypodium leptopodon C. H. Wright, Trans. Linn. Soc. London, Bot. II, 6:83. 1901. TYPE:
Summit of Mt. Roraima, Edo. Bolfvair, Venezuela, 8600 ft, McConnell & Quelch 569 (K not seen
photo US; isotypes NY, US).

Plants epiphytic, in the Flora area known only from Volcan Poas {Stork 2337 p.
p.y UC). Also from Venezuela.

This species is very close to G. bryophila, but the differences in lamina shape
and venation remain constant in specimens of all sizes, demonstrating that they
are not correlated with lamina size and indicating that the two species appear to
be distinct.

488a. Grammitis limbata F6e, Gen. Fil. [M^m. Fong. 5] :233. 1852.

Polypodium nigrolimbatum Jenm. Bull. Bot. Dept. Jamaica N.S., 4:69. 1897, nom. superfl. TYPE:
A renaming of Grammitis limbata Fee, and so based on the type of that name.

Polypodium marginellum var. brasiliense Rosenst. Hedwigia 46:135. 1906. TYPE: Serra Ikerim,
Est. Sta. Catarina, Brazil, Schmalz 163 (S not seen; isotype UC).

Polypodium hessii Maxon, Bull. Torrey Bot. Club 42:223. 1915. TYPE: Sierra de Nagubo, Puerto
Rico, Hess 312 (US).

Plants epiphytic, at 800-900 m elevation, in forests, in the Flora area known
only from Cerro Jefe {Sytsma 1469, Folsom et al. 7112, both MO). Also from the
Antilles, Bolivia, and Brazil.

489. Grammitis marginella (Swartz) Swartz, J. Bot. (Schrader) 1800(2): 17. 1801.

Polypodium marginellum Swartz, Nov. Gen. Sp. PI. Prodr. 130. 1788. TYPE: Jamaica, Swartz (S
not seen photo 6018; isotype UPS-Hb. Thunb. 24445 p. p. not seen).

Plants epiphytic, at (100)1400-2700 m elevation, in forests, from the Cordillera
Central, Cerro Tablazo, the northern end of the Cordillera de Talamanca,

POLYPODIACEAE: 57. GRAMMITIS 325

between Ft. S. Lorenzo and Ft. Sherman (Canal Zone), and Cerro Jefe. Also
from Jamaica and Hispaniola.

489a. Grammitis paramicola L. E. Bishop, Amer. Fern J. 67:105. 1977.

TYPE: 25 km southeast of Gigante near the Huila - Caqueta divide, Depto.
Huila, Colombia, 11000 ft. Little 8663 (US).

Plants epiphytic, at 2400-2800 m elevation, in forests, in the Flora area known
only from the Atlantic slope of the Cordillera Talamanca between the Rio Terbi
and the Rio Sini, Pcia. Limon (Davidse et al 29020, MO). Also from Colombia
and Peru.

“GRAMMITIS sect. XIPHOPTERIS (Kaulf.) Presl”

490. Grammitis blepharodes (Maxon) F. Seym. Phytologia 31:173. 1975.

Polypodium blepharodes Maxon, Contr. U. S. Natl. Herb. 17:407. 1914. TYPE: Near La Palma,
Pcia. S. Jose, 1450-1550 m, Maxon 406 (US; isotype NY).

Polypodium blepharodes var. microlepis Rosenst. Repert. Spec. Nov. Regni Veg. 22:14. 1925.
TYPE: Cerro Tablazo, Pcia. S. Jose. 1900 m, Brade & Brade 80b (S not seen photo 5955a; isotypes
NY, UC).

Plants epiphytic, at 500-1700 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, Cerro Carpintera, the Cordillera de Talamanca to
Pcia. Chiriqui, the Fila Costena near S. Vito de Java, Cerro Tute (Pcia. Veraguas),
El Valle and Penonome, Cerro Azul (Pcia. Panama), Cerro Pirre, Cerro Mali, and
Cerro de Garagara (all Pcia. Darien), and Alto del Buey. Also from Mexico,
Guatemala, Nicaragua, Venezuela, and Colombia.

This species is closely related to, and may even be conspecific with, G.
blepharolepis (C. Chr.) Morton, of which I have seen no authentic material. It is
also similar to G. daguensis, which is a lowland species.

491. Grammitis caucana (Hieron.) Morton, Contr. U. S. Natl. Herb. 38:96. 1967.

Polypodium caucanum Hieron. Bot. Jahrb. Syst. 34:503. 1904. TYPE: Near the Rio Dagua,
Depto. Cauca, Colombia, 2300 m, Lehmann 3257 (B not seen).

Plants epiphytic, at (600)1500-2100 m elevation, in forests, from north of S.
Ramon, the Cordillera Central, the Cordillera de Talamanca to Pcia. Chiriqui,
and El Valle. Also from Nicaragua, Venezuela, Colombia, and Guyana.

492. Grammitis cookii (Underw. & Maxon) F. Seym. Phytologia 31:173. 1975.

Polypodium cookii Underw. & Maxon, Contr. U. S. Natl. Herb. 17:408. 1914. TYPE: Near Finca
Sepacuite, Depto. Alta Verapaz, Guatemala, Cook & Griggs 80 (US fragm NY).

Plants epiphytic, at 1000-1700 m elevation, in forests, from Monteverde and
the Cordillera Central. Also from Guatemala.

493. Grammitis daguensis (Hieron.) Morton, Contr. U. S. Natl. Herb. 38:98. 1967.

Polypodium daguense Hieron. Bot. Jahrb. Syst. 34:504. 1904. TYPE: Rio Dagua, Depto. Cauca,
Colombia, Lehmann 1951 (B not seen fragm and photo US).

FIGS. 486 - 492. Grammitis. FIG. 486. Plant of G. bryophila, Maxon 7980. FIG. 487. Plant of G.
fluminensis, Lellinger & de la Sota 778. FIG. 488. Plant of G. leptopoda, Wright, Guyana. FIG. 489.
Plant of G. marginella, Standley & Valerio 48209. FIG. 490. Median portion of lamina of G.
blepharodes, Standley 33123. FIG. 491. Median portion of lamina of G. caucana, Brade & Brade 80.
FIG. 492. Median portion of lamina of G. cookii, Lellinger 773.

326 POLYPODIACEAE: 57. GRAMMITIS

POLYPODIACEAE: 57. GRAMMITIS 327

Plants epiphytic, at 0-300 m elevation, in forests, from the Rio S. Juan basin
and Boca Curiche (both Depto. Choco). Also from Venezuela and Colombia to
Bolivia.

This species is similar to G. blepharodes, which is an upland species, and to G.
blepharolepis (C. Chr.) Morton, which has shorter, wider, and more distant
segments. Some of the segments of G. daguensis are gibbous, and so that
character does not separate the species from G. blepharolepis very well. Most of
the Choco specimens of G. daguensis are slightly intermediate between that
species and G. blepharolepis', they have relatively shorter, less parallel-sided
segments than do specimens of G. daguensis from Pcias. El Valle and Cauca.

494. Grammitis delitescens (Maxon) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5:32.
1953.

Polypodium delitescens Maxon, Bull. Torrey Bot. Club 32:74. 1905. LECTOTYPE: Summit of
Blue Mountain Peak, Jamaica, Maxon 1513 (US), chosen by Copeland (Amer. Fern J. 42:52. 1952).

Plants epiphytic, at ca. 900 m elevation, in forests, in the Flora area known only
from one collection in Costa Rica {Gomez 776, CR) and from Cerro Jefe
{Valdespino et al 268, US). Also from Cuba, Jamaica, Mexico, Guatemala,
Honduras, and Nicaragua.

This species is closely related to G. myosuroides.

494a. Grammitis flabelliformis (Poir.) Morton, Contr. U. S. Natl. Herb. 38:57.
1967.

Polypodium flabelliforme Poir. Encyc. Meth. 5:519. 1804. TYPE: Plate 87, left hand, of Plumier’s
“Traite...”, which illustrates a specimen collected by Plumier on Martinique.

Polypodium taenifolium Jenm. Bull. Bot. Dept. Jamaica N.S., 4:114. 1897. TYPE: At or near Mt.
Moses, St. Andrew Parish, Jamaica, Syme (IJ not seen).

Polypodium sintenisii Hieron. Hedwigia 44:101. 1905. SYNTYPES: Mt. Hymene, Sierra de
Luquillo, Puerto Rico, Sintenis 1796 (B not seen; isosyntype US); Mt. Guaraguao near Adjuntas,
Puerto Rico, Sintenis 4327b (B not seen); Mt. Piedra in the Sierra de Naguabo, Puerto Rico, Sintenis
5462b (B not seen; isosyntype US); Guadeloupe (various localities). Maze 18, 328, 673, 709 (all B
none seen); and Grenada, Sherring 156 p.p. (B not seen).

Plants epiphytic, at 1000 m elevation, in forests, in the Flora area known only
from Cerro Jefe {Valdespino et al 329 and 354, both US). Also from Jamaica,
Hispaniola, Puerto Rico, the Lesser Antilles, Trinidad, Venezuela, Colombia, and
Surinam.

495. Grammitis hyalina (Maxon) F. Seym. Phytologia 31:174. 1975.

Polypodium hyalinum Maxon, Contr. U. S. Natl. Herb. 17:406. 1914. TYPE: Upper slopes of

Volcan Barba, Pcia. Heredia, Pittier 1928 (US fragm NY).

Plants epiphytic, at 1800-2500 m elevation, in forests, from the Cordillera
Central.

FIGS. 493 " 500. Grammitis. FIG. 493. Median portion of lamina of G. daguensis, Cuatrecasas,
Colombia. FIG. 494. Plant of G. delitescens, Williams & Molina 13690, Honduras. FIG. 495.
Median portion of lamina of G. hyalina, Valerio A66. FIG. 496. Median portion of lamina of G.
limula, Lellinger 1091. FIG. 497. Median portion of lamina of G. mitchellae, Maxon 3213,
Guatemala. FIG. 498. Plant of G. myosuroides, Proctor 4363, Jamaica. FIG. 499. Median portion of
lamina of G. pseudomitchellae, KiUip 5289a. FIG. 500. Median portion of lamina of G. setulosa,
Evans & Lellinger 66.

328 POLYPODIACEAE: 57. GRAMMITIS

496. Grammitis limula (Christ) G6mez, Brenesia 8:47. 1976.

Polypodium limulum Christ, Bull. Soc. Bot. Geneve II, 1:218. 1909. SYNTYPES: La Palma, Pcia.
S. Jose, 1500 m, Pittier 708 (P not seen), 1459 m, Tonduz 12595 (P not seen; isosyntype US), Brade 79
(P not seen), and Werckle in 1903 (P not seen).

Plants epiphytic, at (500)700-2400 m elevation, in forests and on trees in
pastures and along roadsides, from the Cordillera de Tilaran, the Cordillera
Central, Cerro Tablazo, the Fila de Cedral, Cerro Carpintera, and the Cordillera
de Talamanca to Pcia. Chiriqui. Also from Guatemala, Honduras, Venezuela, and
Colombia.

497. Grammitis mitchellae (Baker in Hemsl.) F. Seym. Phytologia 31:174. 1975.

Polypodium mitchellae Baker in Hemsl. Biol. Centr.-Amer., Bot. 3:664. 1885. TYPE: Orange
Walk, Belize, Mitchell (K not seen).

Plants epiphytic, at ca. 500-800 m elevation, in forests, in the Flora area known
only from along Sta. Rita ridge, Pcia. Colon {Correa & Dressier 1043, US). Also
from Guatemala, Belize, and Nicaragua.

498. Grammitis myosuroides (Swartz) Swartz, J. Bot. (Schrader) 1800(2): 18.
1801.

Polypodium myosuroides Swartz, Nov. Gen. Sp. PI. Prodr. 131. 1788. TYPE: Jamaica, Swartz (S
not seen photo 6024).

Polypodium jamesonii Jenm. Bull. Bot. Dept. Jamaica N.S., 4:112. 1897, non Mett., 1883, nom.
illeg. TYPE: Jamaica, /e/imc/i (NY? not seen).

Plants epiphytic, at 2600-3000 m elevation, in forests, from the Cordillera de
Talamanca to Villa Mills (Pcia. Cartago). Also from Cuba, Jamaica, Puerto Rico,
and Mexico.

499. Grammitis pseudomitchellae Lellinger, Proc. Biol. Soc. Wash. 89:383, f. 14.
1985.

TYPE: 10 km north of S. Rafael de Heredia on Volcan Barba, Pcia. Heredia,
1950 m,Mickel3033{NY).

Plants epiphytic, at 1800-2000 m elevation, from Volcan Barba, Tapanti, and
Holcomb’s trail above Boquete.

In frond outline this species closely resembles G. mitchellae from Guatemala,
but that species has forked veins in the segments, setose sporangia, fewer and
shorter marginal setae, and more and shorter setae on the abaxial surface of the
laminae.

500. Grammitis setulosa (Rosenst.) F. Seym. Phytologia 31:175. 1975.

Polypodium setulosum Rosenst. Repert. Spec. Nov. Regni Veg. 10:277. 1912. TYPE: S. Isidro del
General, Pcia. S. Jose, 656 m, Tonduz (Hb. Jimenez 214) (S not seen; isotype fragm US).

Plants epiphytic, at (600)1500-2800 m elevation, in forests, from the Cordillera
Central, Tapanti, the Cordillera de Talamanca to Cerro Chirripo, S. Isidro del
General, the Peninsula de Osa, the Fila Costeha near S. Vito de Java, and Cerro
Jefe. Also from Mexico, Guatemala, Nicaragua, and Colombia.

501. Grammitis truncicola (Klotzsch) Morton, Contr. U. S. Natl. Herb. 38:98.
1967.

Polypodium tmncicola Klotzsch, Linnaea 20:374. 1847. TYPE: Colonia Tovar, Edo. Aragua,
Venezuela, Moritz 252 (B not seen; isotype US).

POLYPODIACEAE: 58. GLYPHOTAENIUM 329

Polypodium tmncicola var. major Klotzsch, Linnaea 20:374. 1847, nom. superfl. TYPE: The
typical variety as described by Klotzsch, and so based on the type of P. tmncicola.

Polypodium tmncicola var, minor Klotzsch, Linnaea 20:374. 1847. TYPE: Merida, Edo. Merida,
Venezuela, Moritz 333 (B not seen; isotypes NY, US).

Polypodium andinum Hook. Sec. Cent. Ferns t. 6. 1860. LECTOTYPE: Rio Hondacha, Andes of
Quito, Pcia. Pichincha, Ecuador, Jameson 780 (K not seen), chosen by Morton (Contr, U. S. Natl.
Herb. 38:98. 1%7).

Plants epiphytic, at 1200 - 1600 m elevation, in forests, from Monteverde, and
La Palma and La Hondura (Pcia. S. Jose). Also from Venezuela to Peru.

502. Grammitis zurquina (Copel.) F. Seym. Phytologia 31:175. 1975.

Xiphopteris zurquina Copel. Amer. Fern J. 42:99, t. 9. 1952. TYPE: Cerros de Zurqui, northeast
of S. Isidro, Pcia. Heredia, 2000 - 2400 m, Standley & Valerio 50495 (US).

Plants epiphytic, at (1200)1900-2400(3300) m elevation, in forests, from S.
Ramon, the southeastern slopes of Volcan Barba, and Cerro Asuncion (Pcia.
Cartago).

58. GLYPHOTAENIUM J. Smith

Plants epiphytic; rhizomes short-creeping, scaly, the scales linear-lanceate, not
clathrate, often setose or glandular at least on the margin; stipes usually about
equalling the laminae, sometimes somewhat shorter, scaly only at the base, long-
setose, the setae thin, flexible, reddish-brown; fronds small to medium-sized,
monomorphic; laminae simple, entire to lobed, mostly oblong, thick and spongy,
setose on the margins and sparsely so on the lamina surface, the setae long,
reddish-brown; veins usually at least partially anastomosing with 1 row of long,
narrow areolae on each side of the midrib; sori lateral on the veins, elliptical to
linear, rarely round, often in several series, usually immersed in cavities in the
lamina, exindusiate; paraphyses obscure; sporangia long-stalked.

Neotropical; 9 species.

1. Sori mostly linear, 2-5 mm long, deeply immersed in the lamina tissue. Rhizome scales entire,
eglandular or with a few gland-tipped hairs along the margins; stipes 3 - 14 cm long, brown or gray;
laminae lanceate, 5-15 cm long, 2-4 cm wide, obtuse at the base, acute at the apex, lobed 1/3 -2/3
of the way to the midrib, the lobes 1-2 times longer than wide.

505. G. spongiosum

1. Sori mostly round or elliptical to elongate, up to 2(3) mm long, superficial to deeply
immersed..2.

2(1). Rhizome scales densely glandular, the glands whitish, cylindrical; laminae undulate to entire,
rarely lobed. Stipes 1.5-5 cm long; laminae linear, 2-11.5 cm long, 5-11(14) mm wide, round at
the apex, acute at the base.

504. G. percrassum

2(1). Rhizome scales eglandular, entire or distantly ciliate-toothed; laminae lobed to undulate,
rarely entire.. 3.

3(2). Rhizome scales distantly ciliate-toothed, contorted near the apex, gray to yellowish; stipes
atropurpureous, smooth; sori often more than 1 mm wide. Stipes 2-11 cm long; laminae linear or
linear-lanceate, (6)8 - 20(25) cm long, (1.5)2 -4 cm wide, acute to obtuse at the base, all but the
juveniles crenately or serrately lobed.

506. G. trifurcatum

3(2). Rhizome scales entire, not contorted near the apex, yellowish to orange; stipes brown or gray,
minutely fluted; sori usually less than 1 mm wide. Stipes 3-5 cm long; laminae linear, (3)5-15(38)
cm long, (0.6)1- 1.5 cm wide, acute to obtuse at the apex, all but the juveniles crenately lobed.

503. G. crispatum

330 POLYPODIACEAE: 58. GLYPHOTAENIUM

506 G. trifurcatum

POLYPODIACEAE: 58. GLYPHOTAENIUM 331

503. Glyphotaenium crispatum (J. Smith in Seem.) J. Smith, Hist. Fil. 188.
1875.

Ctenopteris crispata J. Smith in Seem. Bot. Voy. Herald 227, t. 48. 1854, non Polypodium

crispatum L., 1753. SYNTYPES: Isla de Cacahual, Depto. Choco, Seemann (BM not seen); and Bay
of Choco, Colombia, Seemann 995 (BM not seen fragm US).

Polypodium goniopteroides C. Chr. Ind. Fil. 530. 1906. TYPE: A renaming of Ctenopteris crispata
J. Smith in Seem., and so based on the type of that name.

Polypodium enterosoroides Christ, Bull. Herb. Boissier II, 7:260. 1907. TYPE: La Palma, Pcia. S.
Jose, 1500 m, Werckle 17095 [or 170451] (P not seen; isotypes NY, US).

Plants epiphytic, at 0-600 m elevation, in forests, from La Palma, Pejivalle
(Pcia. Cartago), and the Bay of Choco and Isla de Cacahual (Depto. Choco). Also
from Colombia and Ecuador.

Apparently this species grows only in the wettest forests.

504. Glyphotaenium percrassum (Baker) Copel. Gen. Fil. 212. 1947.

Polypodium percrassum Baker, J. Bot. Brit. For. 25:26. 1887. TYPE: Costa Rica, Cooper (K not

seen fragm NY; isotype US).

Polypodium repletum Christ, Bull. Herb. Boissier II, 7:260. 1907. TYPE: Costa Rica, Werckle in
1905 (P not seen).

Plants epiphytic, at 1300-2600 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Pcia. Chiriqui.

505. Glyphotaenium spongiosum (Maxon) Copel. Gen. Fil. 212. 1947.

Enterosora spongiosa Maxon, Proc. Biol. Soc. Wash. 52:113. 1939. TYPE: Vicinity of La Palma,
on the road to La Hondura, Pcia. S. Jose, ca. 1600 m, Maxon & Harvey 7975 (US).

Plants epiphytic, at 1500-2400 m elevation, in forests, from the Cordillera de
Tilaran, the Cordillera Central, and the Cordillera de Talamanca to Pcia. Chiriqui.

506. Glyphotaenium trifurcatum (L.) Lellinger, Amer. Fern J. 75:31. 1985.

Polypodium trifurcatum L. Sp. PI. 2:1084. 1753. TYPE: Plate 138 in Plumier’s “Traite...”, which
illustrates a specimen collected by Plumier on Martinique.

Polypodium trifurcatum var. brevipes Hieron. Bot. Jahrb. Syst. 34:500. 1904. SYNTYPES:

Quebrada de Imbi, Rio Cuaiquer, Cordillera de Pasto, Depto. Narino, Colombia, Lehmann 77 (B
not seen); Farallones de Cali, Depto. El Valle, Colombia, Lehmann 1977 (B not seen); and
Cordillera west of Cali, Depto. El Valle, Colombia, Lehmann 7664 (B not seen; isosyntype US).

Plants epiphytic, at 700-2500 m elevation, in forests, from north of S. Ramon,
the Cordillera Central, Pejivalle (Pcia. Cartago), the mountains of western
Panama, Campo Tres northeast of Altos de Pacora (Pcia. Panama), Cerro de
Garagara, Cana, and Cerro Pirre (all Pcia. Darien), and the Serrarua del Darien
and the Ciudad Bolivar - Quibdo road at ca. Km. 141 (Depto. Choco). Also from
the Antilles, Venezuela, Colombia to Bolivia, Guyana, and Surinam.

FIGS. 501-506. Grammitis and Glyphotaenium. FIG. 501. Median portion of lamina of Gr.
truncicola, Maxon 392. FIG. 502. Median portion of lamina of Gr. zurquina, Standley & Valerio
51599. FIG. 503. Plant of Gl. crispatum, Skutch 2760. FIG. 504. Plant of Gl. percrassum, Standley &
Torres 47929. FIG. 505. Plant of Gl. spongiosum, Skutch 4958. FIG. 506. Plant of Gl. trifurcatum,
Standley & Valerio 51633.

332 POLYPODIACEAE: 59. COCHLIDIUM

59. COCHLIDIUM Kaulf.

Plants epiphytic or epipetric; rhizomes erect or ascending, scaly, the scales
lanceate, concolorous; stipes short to obsolete, not articulate, brown, glabrous;
fronds small to minute, monomorphic or subdimorphic; laminae simple, linear,
entire (or the sterile portions serrately lobed in C. serrulatum), subcoriaceous,
glabrous (or bearing short, cylindrical hairs in C. serrulatum), often bearing
obvious hydathodes near the apex on the adaxial surface; veins free (unbranched
or 1 -forked) or sometimes casually anastomosing to form a partial series of
areolae along the midrib; sori lateral on the veins, usually elongate and often
confluent into a linear coenosorus, usually in a single, deep groove, exindusiate;
paraphyses lacking; sporangia long-stalked.

Neotropical with 1 species extending to Africa; 16 species.

BISHOP, L. E. 1978. Revision of the genus Cochlidium (Grammitidaceae). Amer. Fern J.
68:76 - 94.

CHRISTENSEN, C. 1929. Taxonomic fern-studies I. Revision of the polypodioid genera with
longitudinal coenosori... Dansk Bot. Ajk. 6(3):l-93, 1. 1 -XIII.

1. Sori superficial, round or broadly elliptical, often confluent at maturity, confined to a usually
broad, entire terminal portion of the narrow, otherwise usually serrately lobed laminae; receptacles
discrete, easily visible when the sporangia are removed. Rhizomes erect or ascending; laminae
hnear, 2-8 cm long, 0.5 -2.5 mm wide.

509. C. serrulatum

1. Sori immersed in a long, deep groove, aggregated into a linear coenosorus along the uniformly
wide, entire laminae; receptacles linearly confluent, closely parallel to the costa.. 2.

2(1). Hydathodes obscure on the adaxial surface of the laminae, usually not visible under 20 x
magnification; sporangia more than 0.25 mm long. Rhizomes erect or ascending; laminae hnear,
3 - 15 cm long, 1.5 - 2.5 mm wide.

508. C. rostratum

2(1). Hydathodes usually conspicuous on the adaxial surface of the laminae; sporangia less than
0.25 mm long. Rhizomes erect or ascending; laminae linear, (2.5)4-11 cm long, (1)2-4 mm wide.
507. C linearifolium

507. Cochlidium linearifolium (Desv.) Maxon ex C. Chr. Dansk Bot. Ark. 6(3) :23.
1929.

Monogramma linearifolia Desv. Ges. Naturf. Freunde Berlin Mag. 5:302, t. 7, f. 5. 1811. TYPE:
French Guiana, collector unknown (P-Hb. Desv. not seen).

Pleurogramme immersa Fee, Hist. Vittar. Pleurogr. [Mem. Foug. 3]:37, t. 4, f. 5. 1852, nom.
superfl. TYPE: A renaming of Pleurogramma linearifolia Desv., and so based on the type of that
name.

Pleurogramme gyroflexa Christ in Pitt. Prim. FI. Costaric. 3(1):10. 1901. TYPE: Valley of the Rio
General, Pcia. S. Jose, 700 m, Pittier 12061 (BR or P not seen; isotype CR).

Plants epiphytic, at 0-600 m elevation, in forests, from the Atlantic coastal
plain of Costa Rica, the valley of the Rio General, the Peninsula de Osa, Cerro
Jefe, Sta. Rita ridge (Pcia. Colon), above Cana, and the Corcovada region and
Bahia Solano (Depto. Choco). Also from Guatemala to Nicaragua, Colombia,
Guyana, and Brazil.

508. Cochlidium rostratum (Hook.) Maxon ex C. Chr. Dansk Bot. Ark. 6(3) :23.
1929.

Monogramme rostrata Hook. Sp. Fil. 5:122, t. 288B. 1864. TYPE: Island in Lake Omotepec,
Depto. Rivas, Nicaragua, Wright (K not seen fragm NY; isotype US).

POLYPODIACEAE: 60. LOXOGRAMME 333

Cochlidium rostratum var. areolatum C. Chr. Dansk Bot. Ark. 6(3) :25. 1929. TYPE: Morne
Colombeau, Anse-a-Foleur, Haiti, Ekman H-4376 (C not seen; isotype US).

Plants epiphytic, at 1500-2600(3000) m elevation, in forests, from the
Cordillera de Tilardn, the Cordillera Central, and the Cordillera de Talamanca to
Pcia. Chiriqm. Also from Cuba, Jamaica, Hispaniola, the Lesser Antilles, Mexico
to Nicaragua, Venezuela, and Colombia.

509. Cochlidium serrulatum (Swartz) L. E. Bishop, Amer. Fern J. 68:80. 1978.

Acrostichum serrulatum Swartz, Nov. Gen. Sp. PI. Prodr. 128. 1788, non Polypodium serrulatum
Mett., 1856. TYPE: Jamaica, Swartz (S not seen fragm US; isotypes B-Hb. Willd. 19589-1, 3 not seen
Tryon photos).

Xiphopteris extensa Fee, Hist. Foug. Antill. [Mem. Foug. 11]:14, t. 19, f. 3. 1866. TYPE:

Guadeloupe, L’Herminier in 1864 (P or RB not seen).

Polypodium duale Maxon, Contr. U. S. Natl. Herb. 16:61. 1912. TYPE: A renaming of

Acrostichum serrulatum Swartz, and so based on the type of that name.

Xiphopteris auyantepuiensis Vareschi, FI. Venez. 1(2):879. 1969. TYPE: Auyan-tepui, Edo.

Bohvar, Venezuela, 1800 m, Vareschi & Foldats 4806 (VEN not seen).

Plants epiphytic, at (0)300-2500 m elevation, in forests and open areas, from
the Cordillera de Tilaran, the Cordillera Central, the Cordillera de Talamanca to
Pcia. Chiriquf, near Sta. Fe, Cerro Jefe, the Cerro de Garagara (Pcia. Darien), and
Alto del Buey and the lower Rfo S. Juan. Also throughout tropical America.

60. LOXOGRAMME (Blume) Presl

Plants epipetric or epiphytic; rhizomes creeping, they and the roots bearing
abundant, matted hairs, scaly, the scales lanceate, distinctly clathrate; stipes
obsolete, subarticulate, scaly at the base, the scales like those of the rhizomes;
fronds small, monomorphic; laminae simple, linear-elliptic, entire, attenuate at the
base and apex, chartaceous, yellow-green, sparsely pilosulous, the hairs short,
relatively thick, cylindrical, pluricellular, brown, deciduous; veins forked and
copiously anastomosing with included veinlets; sori lateral on the veins, elongate,
in a single series on each side of the midrib, slightly impressed, exindusiate;
paraphyses apparently lacking, but sporangium stalks often present; sporangia
long-stalked.

Paleotropics; ca. 40 species with 1 in Central America.

510. Loxogramme mexicana (F6e) C. Chr. Ind. Fil. Suppl. 3:125. 1934.

Selliguea mexicana Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 7]:43, 1. 10, f. 4. 1857.
TYPE: Sierra de S. Pedro Nolasco, Talea, Edo. Oaxaca, Mexico, Juergensen 945 (P or RB not seen;
isotype BM not seen).

Grammitis salvinii Hook. Sec. Cent. Ferns t. 71. 1861. TYPE: Vera Paz [Depto. Alta Verapaz?],
Guatemala, 3500 - 5000 ft, Salvin (K not seen; probable isotype BM not seen).

Plants epipetric or epiphytic, at 1000 - 1900 m elevation, in forests, from the Fila
de Cedral, Cerro Tablazo, and the Cordillera de Talamanca to Pcia. Chiriquf.
Also from Mexico, Guatemala, El Salvador, and Nicaragua.

334 CYATHEACEAE: 61. METAXYA-62. LOPHOSORIA

CYATHEACEAE

Rhizomes often stout and clothed at least at the base with a thick covering of
adventitious roots, forming an erect, tree-like trunk (usually short in Cnemidaria,
ascending in Lophosoria, and creeping in Metaxya), scaly and sometimes spiny
(only densely hairy at the apex in the latter genera), the scales concolorous or
more commonly weakly to sharply bicolorous and the margins sometimes
fimbriate or bearing minute, spine-like cells; fronds mostly large, borne at the
apex of the erect or ascending rhizomes (scattered along the creeping rhizomes in
Metaxya); stipes thick, often spiny at the base, mostly deciduous and revealing
patterned scars in regular spirals on the rhizome; laminae l-3(4)-pinnate,
glabrous, hairy, or scaly, commonly with short, curved hairs on the adaxial surface
of the costae and costules and with scales usually like those of the rhizome,
sometimes dissimilar, often confined to the abaxial surface of the rachis, costae,
and sometimes costules; veins free, simple or forked (and re^larly forming costal
areolae in Cnemidaria); sori round, surficial on the laminae; indusia absent, scale-
like, shallowly or deeply cup-like, or mostly or completely enclosing the sporangia;
sporangia with a short, thick stalk, angular capsules, and oblique annulus;
paraphyses often abundant; spores trilete.

1. Caudices and stipe bases scaly, sometimes hairs also present; plants mostly arborescent (except
in Cnemidaria).3.

1. Caudices and stipe bases hairy, scales never present; plants not arborescent.. 2.

2(1). Laminae pinnate, not glaucous or pruinose abaxially; rhizomes creeping.

61. Metaxya

2(1). Laminae decompound, often glaucous or pruinose abaxially; rhizomes erect to ascending.

62. Lophosoria

3(1). Basal basiscopic vein of each vein group arising from the costa or nearly so, not distal along
the costule; cells of the body of the stipe scales (the protruding marginal and apical spine-hke or
fimbriate cells excepted) entirely uniform, the cells along the margin not different from the central
ones. Stipe scales whitish to brown, hnear, 0.5 -1(1.5) mm wide, 3-5(6) cm long; indusia fragile,
completely surrounding the sporangia, shghtly apiculate at the apex.

66. Sphaeropteris

3(1). Basal basiscopic vein of each vein group arising from the costule markedly distal to the costa;
cells of the body of the stipe scales (any protruding marginal and apical spine-like or fimbriate cells
excepted) not entirely uniform, the cells along the margin different from the central ones in size,
shape, wall thickening, or orientation..4.

4(3). Stipe scales (and often those of the rachises and costae) provided with a dark (rarely pale)
apical spine-like cell and sometimes similar, distant, spreading lateral cells, the scales concolorous to
strongly bicolorous with the dark central band often several cells thick at the base and the pale to
whitish margins often erose.

65. Alsophila

4(3). Stipe scales (and those of the rachises and costae) lacking apical or lateral spine-like cells
(except in C poeppigii, with approximate, ascending spine-like cells and in Cyathea stipularis and
trichiata, with rather thin, fimbriate ceils at entirely right angles to the margins), the scales
concolorous to weakly bicolorous, the central band usually only 1 cell thick at the base..5.

5(4). Mature laminae usually 2-pinnate-pinnatifid, rarely 2-pinnate (merely pinnate or pinnate-
pinnatifid in a few small species), the pinnules (or pinnae) regularly deeply pinnatifid (occasionally
merely deeply lobed); costae and costules usually strigose adaxially, the hairs stiff and recurved;
laminae often scaly or hairy abaxially, at least on the costae and costules; basal veins free and not
connivent (except a transverse costal vein joining adjacent pinnate veins groups in C petiolata and
williamsii); indusia present or absent; rhizomes usually forming an obvious, erect trunk.

64. Cyathea

CYATHEACEAE: 61. METAXYA-62. LOPHOSORIA 335

5(4). Mature laminae 1-2-pinnate, the pinnae or pinnules sometimes crenately lobed, rarely
deeply lobed; costae and costules usually not strigose adaxially; laminae usually glabrous abaxially,
the costules and costae occasionally scaly or hairy; basal veins of each vein group forming regular
areolae along the costae or the basal veins connivent to the base of the sinus or occasionally meeting
the sinus just distcd to the base; sori usually in 1 medicd, sinuous series; indusia present; rhizome
usually not forming a trunk.

63. Cnemidaria

61. METAXYA K. Presl

Plants terrestrial; rhizomes creeping, woody, 8-15 mm in diam., hairy, the hairs
pale tan to golden, multicellular, soft, rather straight; stipes stout, long, sparsely
hairy, articulate to low phyllopodia; fronds large, monomorphic; laminae oblong,
pinnate, papyraceous, glabrous, the rachis and costae adaxially sulcate; pinnae
linear, acute at the base, acute-caudate at the apex; veins free, unbranched or 1-
forked, at the apex arcuate and confluent with the cartilaginous pinna margin; sori
surficial, irregularly disposed on the proximal portion of the veins, round or
slightly elongate, exindusiate; paraphyses abundant, wooly, pluricellular; sporangia
short-stalked.

Neotropical; monotypic.

ROY, S. K. and R. E. HOLTTUM. 1965. Cytological and morphological observations on Metaxya
rostrata (H.B.K.) Presl. Amer. Fern J. 55:158-164.

TRYON, R. M., Jr. 1970. The classification of the Cyatheaceae. Contr. Gray Herb. 200:3-53.

511. Metaxya rostrata (H.B.K.) K. Presl, Tent. Pterid. 60. 1836.

Polypodium rostratum Humb. & Bonpl. ex Willd. Sp. PI. ed. 4, 5:193. 1810, non Burm., 1768, nom.
illeg. TYPE: Near S. Antonio de Yavita, Terr. Amazonas, Venezuela, Humboldt 966 (B-Hb. Willd.
19691 not seen Tryon photo).

Aspidium rostratum H.B.K. Nov. Gen. Sp. 1:12 (fol. 10). Jan 1816. TYPE: Effectively a renaming
of Polypodium rostratum Humb. & Bonpl. ex Willd., and so based on the type of that name.

Polypodium humboldtii Poir. Encyc. Meth. Suppl. 4:497. Dec 1816. TYPE: A renaming of
Polypodium rostratum Humb. & Bonpl. ex Willd., and so based on the type of that name.

Polypodium parkeri Hook. & Grev. Icon. Fil. 2:t. 232. 1831. TYPE: Mazaruni River, Guyana,
Parker (K not seen).

Plants terrestrial, at 0-600 m elevation, in forests, from the Atlantic coastal
plain of Costa Rica and Panama, Los Palmares and Cerro Turrubares (Pcia. S.
Jose), the Peninsula de Osa and vicinity, the Canal Zone, Sta. Rita ridge (Pcia.
Colon), La Campana (Pcia. Panama), and the northern part of the Choco. Also
from the Lesser Antilles, Mexico to Honduras, Nicaragua, Trinidad, Venezuela,
Colombia to Bolivia, the Guianas, and Brazil.

62. LOPHOSORIA K. Presl

Plants terrestrial; rhizomes erect to ascending, forming a low caudex, hairy, the
hairs pale tan to whitish, multicellular, rather stiff and straight; stipes stout, long,
densely hairy at the base; fronds very large, monomorphic; laminae ovate, 4-
pinnate, chartaceous, villous on the axes, the hairs whitish, pluricellular, soft,
somewhat contorted, the rachis and costae (but not the costules) sulcate; pinnae
oblong; pinnules oblong to lanceate, truncate at the base, acute-acuminate at the
apex, the lateral segments deeply pinnatifid to pinnate, farinose on the abaxial
surface, the farina deposited as a surficial coating or as a tomentum of minute

336 CYATHEACEAE: 62. LOPHOSORIA

516 Cn. ewanii

517 Cn. horrida

518 Cn. mutica var. chiricana

CYATHEACEAE: 62. LOPHOSORIA 337

rods; veins free, 1-forked; sori lateral on the veins, round, exindusiate; paraphyses
few to abundant, catenate, lax; sporangia short-stalked.

Neotropical; monotypic.

TRYON, R. M., Jr. 1970. The classification of the Cyatheaceae. Contr. Gray Herb. 200:3 - 53.

512. Lophosoria quadripinnata (J. F. Gmel.) C. Chr. in Skottsb. Nat. Hist. Juan
Fernandez, Bot. 1:16. 1920.

Polypodium glaucum Swartz, Nov. Gen. Sp. PI. Prodr. 134. 1788, non Thunb., 1784, nom. illeg.
TYPE: Jamaica, Swartz (S not seen; isotype B-Hb. Willd. 19723 not seen microfiche S. I. Library).

Polypodium quadnpinnatum J. F. Gmel. Syst. Nat. ed. 13, 2:1314. 1791. TYPE: A renaming of
Polypodium glaucum Swartz, and so based on the type of that name.

Polypodium cinereum Cav. Descr. PI. 248. 1801. TYPE: Chiloe, Nee (MA not seen), seen by
Christensen (Dansk Bot. Ark. 9(3): 13. 1937).

Polypodium pruinatum Swartz, J. Bot. (Schrader) 1800(2) :29. 1801; Syn. Fil. 41, 70. 1806, nom.
superfl. TYPE: A renaming of Polypodium cinereum Cav., and so based on the type of that name.

Polypodium griseum Schkuhr, Vier Zwanzigste Kl. Linn. Pfl.-Syst. 2:25, t. 25b. 1805, nom. superfl.
TYPE: A renaming of Polypodium pminatum Swartz, and so based on the type of that name.

IPolypodium caesium K. Presl, Rehq. Haenk. 1:27. 1825. TYPE: Huanuco, Depto. Huanuco,
Peru, Haenke (PRC not seen).

Alsophila millefolia Desv. Mem. Soc. Linn. Paris 6:320. 1827, as ''millefolium P TYPE: Plate 33
of Plumier’s “Traite...”, which is based on a specimen collected by Plumier on Hispaniola.

Cyathea discolor Bory in Duperrey, Voy. Coquille, Crypt. 6:281. 1829. TYPE: Chile, D’Urville (P-
Hb. Bory not seen photo 3448; isotype L not seen photo 316).

Alsophila monticola M£U't. Icon. Plant. Crypt. Bras. 75. 1834. TYPE: Villa Rica, Est. Minas
Gerais, Freireiss (M not seen).

Lophosoria affinis K. Presl, Gefassbiindel Farm 37 (postpr. 345). 1847. TYPE: Caracas, Distr.
Fed., Venezuela, Karsten 53 (PRC not seen).

Lophosoria polypodioides K. Presl, Gefassbiindel Farm 37 (postpr. 345). 1847. TYPE: Not stated.
Trichosoms glaucescens Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:283
(repr. 131). 1849. LECTOTYPE: Between Talea and the Hacienda de Sta. Gertrudis, Edo.

Veracruz, Mexico, Aug 1842, Liebmann PL Mex. 2095, FL Mex. 875 (C not seen photo 5590;
isolectotype US), chosen by A. R. Smith (FI. Chiapas 2:144. 1981).

Trichosorus glaucescens var. major Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk.
Afd. V, 1:283 (repr. 131). 1849. TYPE: S. Yago Amatlan, Edo. Oaxaca, Mexico, 6000 ft, July 1842,
Liebm. PL Mex. 2073 (C not seen photo 5591; isotype US).

Trichosoms densus Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:284
(repr. 132). 1849. LECTOTYPE: Cerro Chinautla, Edo. Puebla, Mexico, 7000 ft, Aug 1841,

Liebmann PL Mex. 2083, FL Mex. 881 (C not seen photo 5592; isolectotype US), chosen by A. R.
Smith (FI. Chiapas 2:144. 1981).

FIGS. 507-518. Cochlidium, Loxogramme, Metaxya, Lophosoria, and Cnemidaria. FIG. 507. Plant of
Co. linearifolium, Stern et al. 533. FIG. 508. Plant of Co. rostratum, Werckle (Hb. Jimenez 552).
FIG. 509. Plant of Co. sermlatum, Brade 71. FIG. 510. Plant of Lox. mexicana, Brade & Brade 201.
FIG. 511. Median portion of pinna of M. rostrata, Standley 27526. FIG. 512. Pinnule of Lop.
quadripinnata, Killip 5256. FIG. 513. Median portion of median pinna of Cn. chocoensis, Lellinger &
de la Sota 895. FIG. 514. Median portion of median pinna of Cn. choricarpa, Chrysler 5235. FIG.
515. Median portion of median pinna of Cn. cocleana, Wilbur 11111. FIG. 516. Median portion of
median pinna of Cn. ewanii, Lellinger & de la Sota 178. FIG. 517. Median portion of median pinna
of Cn. horrida, Pittier 1679. FIG. 518. Median portion of median pinna of Cn. mutica var. chiricana,
Killip 5297.

338 CYATHEACEAE: 63. CNEMIDARIA

Trichosorus frigidus Liebm. Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. V, 1:284
(repr. 132). 1849. LECTOTYPE: Cerro de Zempoaltepec, Edo. Oaxaca, Mexico, 9000 ft, June 1842,
Liebmann PL Mex. 2082, FI. Mex. 884 (C not seen; isolectotype US), chosen by A. R. Smith (FI.
Chiapas 2:144. 1981).

lAlsophila schafjheriana F6e, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]: 109. 1857.
TYPE: S. Martin near Huatusco, Edo. Veracruz, Mejdco, Schaf/her 232 (RB not seen).

IPlecosorus leptocladon Fee, Icon. Esp. Nouv. [Mem. Foug. 10]: 19, t. 34, f. 1. 1865. TYPE:
Ocana, Depto. Norte de Santander, Colombia, Schlim 438 (RB not seen).

Lophosoria prostrata Fee, Crypt. Vase. Br6sil 1:173. 1^9. TYPE: Serra do Couto, Est. Rio de
Janeiro, Brazil, Glaziou 3165 (P not seen photo 3446; isotype RB not seen).

Lophosoria acaulis Fee, Crypt. Vase. Br6sil 1:174. 1869. TYPE: Serra do Couto, Est. Rio de
Janeiro, Brazil, Glaziou 3164 (P not seen photo 3447; isotype RB not seen).

Alsophila pruinata var. conspicua Sodiro, Amales Univ. Quito 12(80):232 (repr. 538). 1895. TYPE:
Confluence of the Rfo Pilat6n with the Toachi, Pcia. Pichincha, Ecuador, Sept 1892, Sodiro (Hb.
Sodiro not seen; isotype P not seen photo 3396).

Alsophila contracta Hieron. Hedwigia 45:236. 1906, non Fee, 1869, nom. illeg. SYNTYPES:
Between Pacasmayo and Moyobamba near Challuayacu and Tambo Centamala, Depto. S. Martin?,
Peru, Stuebel 1066 (B not seen); and near Ines and Callecalle, Depto. Amazonas, Peru, 2600 - 3450
m, Stuebel 1067 (B not seen).

Alsophila pruinata var. tenuis Christ, Bull. Herb. Boissier II, 6:185. 1906. TYPE: Costa Rica,
Werckle (P not seen).

Alsophila bilineata Sodiro, Anales Univ. Centr. Ecuador 22:90 (repr. 13). 1908. TYPE: Mt.
Pichincha, Pcia. Pichincha, Ecuador, April 1907, Sodiro (Hb. Sodiro not seen; isotype US).

Alsophila revoluta C. Chr. Ind. Fil. Suppl. 1:5. 1913. TYPE: A renaming of Alsophila contracta
Hieron., and so based on the type of that name.

Plants terrestrial, at 1500-3000 m elevation, along roadsides, forest margins
and in open areas, from the Cordillera de Tilaran, the Cordillera Central, the
Cordillera de Talamanca to Pcia. Chiriqui, the Fila Costeha near S. Vito de Java,
and the summit of Alto del Buey. Also from the Greater Antilles, Mexico to
Nicaragua, Trinidad, Venezuela, Colombia to Bolivia, Brazil, Argentina, and
Chile.

63. CNEMIDARIA K. Presl

Plants terrestrial; caudices ascending or erect, woody, rudimentary in most
species or up to ca. 1.5 (3. 5) m long, often with a thick covering of adventitious
roots, especially at the base, scaly, the scales mostly bicolorous, the central stripe
dark brown, the margins whitish, commonly erose; stipes stout, long, smooth or
spiny, scaly especially at the base, the scales like those of the rhizomes; fronds
large, monomorphic; laminae oblong, pinnate or pinnate-pinnatifid, often with a
conform terminal pinna, obtuse to truncate at the base, acute or obtusish at the
apex, papyraceous or chartaceous, sparsely scaly and sometimes hairy on the
rachis and costae, the lamina surface usually glabrous, the rachis and costae
adaxially sulcate; pinnae oblong, obtuse to truncate at the base, acute, acuminate,
or caudate at the apex, commonly broadly crenate; veins 1- 4-forked or in pinnate
groups, often with 1 or more lateral pairs anastomosing, confluent, or terminating
at or distal to the sinus between lobes; sori lateral on the veins, in 1-4 sinuous
series between the costules and the margin, the indusia saucer- to cup-shaped or
subglobose; paraphyses rudimentary or lacking; sporangia short-stalked.

Neotropical; 25 species.

STOLZE, R. G. 1974. A taxonomic revision of the genus Cnemidaria (Cyatheaceae). Fieldiana,
Bot. 37:1-98.

CYATHEACEAE: 63. CNEMIDARIA 339

1. Basal veins united; costal areolae present. Sori in 1(2) series..6.

1. Basal veins free; costal areolae absent. Sori in 1-4 series..2.

2(1). Pinnae entire, minutely glandular abaxially; pinnae linear-lanceolate, subobtuse at the base,
acuminate at the apex.

516a. C. glandulosa

2(1). Pinnae distinctly lobed, glabrous abaxially..3.

3(2). Pinna segments obtuse to round; rachises pilose on the adaxial and abaxial surfaces, the hairs
curved, subappressed. Stipe base spines absent; pinna segments subentire, separated by narrow,
acute sinuses; veins 1- or 2-forked; sori in 1 series.

518. C mutica var. chiricana

3(2). Pinna segments acute to acuminate; rachises glabrous (except sometimes sUghtly pilose on
the abaxial surface in C. mutica var. grandis).A.

4(3). Lateral margins of the segments of the larger pinnae crenate to shallowly lobed; supra-basal
sinuses round or the pinnae sometimes cut to the costa. Stipe base spines up to 2.5 mm long or
rarely absent; veins 3-forked or pinnately branched; sori in 1-4 series.

520. C. mutica var. grandis

4(3). Lateral margins of the segments of the larger pinnae entire or nearly so; supra-basal sinuses
acute..5.

5(4). Pinna segments more than 2 times longer than wide; pinnae cut more than 3/4 of the way to
the costa. Stipe base spines up to 2.5 mm long or rarely absent; veins 1- 4-forked; sori in 1 or 2
series.

521. C. mutica var. mutica

5(4). Pinna segments up to 1.5 times longer than wide; pinnae cut 1/2 -3/4 of the way to the costa.
Stipe base spines obsolete to absent; veins 1- or 2-forked; sori in 1 series.

519. C. mutica var. contigua

6(1). Pinnae entire to broadly serrate, narrowly elliptic-lanceolate; lateral pinnae obtuse to cuneate
at the base, acute-aciuninate at the apex.

522a. C stolzeana

6(1). Pinnae decidedly lobed..?.

7(6). Pinna segments acute to acuminate at the apex; median pinnae cut 3/4 or more of the way to
the costa; lateral pinnae up to 14 cm wide. Sori in 1 submarginal series.

517. C. horrida

7(6). Pinna segments round at the apex; median pinnae cut no more than 1/2 (2/3 in C. cocleana)
of the way to the costa; lateral pinnae less than 8 cm wide..8.

8(7). Segment apices entire; laminae gradually reduced to a non-conform, narrowly tri2mgular
apex, the subapical lateral pinnae about 1/2 as long as the apex. Lateral pinnae 15-28 cm long,

2.5 - 6 cm wide.

514. C. choricarpa

8(7). Segment apices usually finely serrate; laminae abruptly reduced to a subconform, nearly
oblong apex commonly bearing a pair of long, basal lobes at the base, the subapical lateral pinnae
about as long as the apex. .9.

9(8). Stipe bases lacking spines.. 11.

9(8). Stipe bases decidedly spiny.. 10.

10(9). Pinna segments approximate, the sinuses narrow (1-2 mm wide); indusia semicircular to
almost circular, not completely surrounding the receptacle; costae and costules with broad, brown,
deciduous scales on the abaxial surface. Lateral pinnae 25 - 40 cm long, 4,5 - 6 cm wide.

522. C. spectabilis var. colombiensis

10(9). Pinna segments distant, the sinuses U-shaped (3-5 mm wide); indusia circular, completely
surrounding the receptacle; costae and costules lacking scales on the abaxial surface. Lateral pinnae
ca. 40 cm long, 6 cm wide.

515. C. cocleana

340 CYATHEACEAE: 63. CNEMIDARIA

11(9). Pinnae shallowly pinnatifid, incised ca. 1/4 of the way to the costa; veins from the costule
simple or 1-forked, in larger specimens commonly laterally anastomosing to form an irregular row of
costal areolae; scales on the axes entirely pale or with pale margins and a dull brown center. Lateral
pinnae (10)15-45 cm long, (3)4-6 cm wide.

516. C ewanii

11(9). Pinnae deeply pinnatifid, incised ca. 1/2 of the way to the costa; veins from the costule 1-3-
forked, in larger specimens not laterally anastomosing, the costal areolae absent; scales on the axes
with pale margins and a glossy brown center. Lateral pinnae (10)15-35 cm long, (2)3.5 -7.5 cm
wide.

513. C. chocoensis

513. Cnemidaria chocoensis Stolze, Fieldiana, Bot. 37:59, t. 20. 1974.

TYPE: Principal ridge and slopes 2 km east of S. Jose del Palmar, Depto.
Choco, 1550-1650 m, Lellinger& de la Sota 763 (US; isotypes COL, CR, F, HU A,

LP).

Plants terrestrial, at 700-800 m elevation, in the Flora area known only from
0.3 km east of the road across the suspension bridge at ca. Km. 141 along the
Ciudad Bolivar - Quibdo road, Depto. Choco, 750 m (Lellinger & de la Sota 895,
COL, CR, F, HU A, LP, US). Also from other localities in Colombia.

514. Cnemidaria choricarpa (Maxon) Tryon, Contr. Gray Herb. 200:51. 1970.

Hemitelia choricarpa Maxon, Contr. U. S. Natl. Herb. 16:40, t. 24d. 1912. TYPE: Buenos Aires,
Pcia. Puntarenas, Pittier 4835 (US fragm NY).

Plants terrestrial, at 0-1800 m elevation, in forests, from 1 km north of La
Hondura (Pcia. S. Jose), above Platanillo (Pcia. Cartago), the valley of the Rio
General, the Fila Costena near S. Vito, the Peninsula de Osa, Cerro Jefe, Cerro
Pirre, and the Choco. Also from other localities in Colombia.

515. Cnemidaria cocleana Stobce, Fieldiana, Bot. 37:52, f. 17. 1974.

TYPE: La Mesa, 5 mi north of El Valle, Pcia. Code, Tyson, Godfrey et al 2452
(FSU not seen Field Mus. photo F not seen GH not seen).

Plants terrestrial, at 500-1000(1200) m elevation, in forests, from Fortuna Lake
(Pcia. Bocas del Toro), near Sta. Fe, El Valle, above El Cope (Pcia. Code), and
the upper Rio Tuquesa (Pcia. Darien).

516. Cnemidaria ewanii (Alston) Tryon, Contr. Gray Herb. 200:52. 1970.

Cyathea ewanii Alston, J. Wash. Acad. Sci. 48:231. 1958. TYPE: Between Mocoa and Urcusique,
Depto. Putamayo, Colombia, 900 m, Ewan 16729 (BM not seen; isotypes GH not seen, MO not seen,
UC, US).

Plants terrestrial, at 0-800 m elevation, in forests, from the foot of Alto del
Buey and Istmina (Depto. Choco). Also from Colombia and Ecuador.

516a. Cnemidaria glandulosa Stolze, Amer. Fern J. 74:101, f. 1. 1984.

TYPE: North of S. Felix on Cerro Colorado copper mine road, Pcias.
Chiriqui- Bocas del Toro, 5000-5500 ft, Mori & Kallunki 5908 (US; isotype MO
not seen).

Plants terrestrial, at 1500-1600 m elevation, known only from the type.

517. Cnemidaria horrida (L.) K. Presl, Tent. Pterid. 57. 1836.

Polypodium horridum L. Sp. PI. 2:1092. 1753. LECTOTYPE: Plate 4 of Plumier’s

“Description...”, chosen by Stolze (Fieldiana, Bot. 37:42. 1974).

CYATHEACEAE: 63. CNEMIDARIA 341

Cyathea commutata Spreng, Anleit. Kenntn. Gew. 3:146, t. 3, f, 32. 1804, TYPE: Santo Domingo
[Hispaniola], collector unknown (LZ destroyed).

Hemitelia hookeri Fee, Gen. Fil. [Mem. Foug. 5]:349. 1852. TYPE: Based on H. horrida sensu
Hooker (Sp. Fil. 1:30, t. XV. 1866) syn. excl., and so the plate (or a specimen from which the plate
was drawn) must be the type.

Hemistegia repanda Fee, Gen. Fil. [Mem. Foug. 5]:351. 1852. TYPE: Cuba, Linden (RB not
seen).

Plants terrestrial, at 700-1400 m elevation, in forests, from along the Rio
Sarapiqui between Cariblanco and S. Miguel (Pda. Alajuela) and Pies de Sta.
Barbara (Pcia. Heredia). Also from the Greater Antilles, St. Thomas, Venezuela,
and Colombia to Peru.

518. Cnemidaria mutica var. chiricana (Maxon) Stolze, Fieldiana, Bot. 37:35.
1974.

Hemitelia chiricana Maxon, Contr. U. S. Natl. Herb. 16:33, t. 20. 1912. TYPE: Between Alto de
las Palmas and the top of Cerro de la Horqueta, Pcia. Chiriqm, 2100 - 2268 m, Maxon 5519 (US).

Plants terrestrial, at 1200-2200 m elevation, in forests, from near Tapantf,
around Boquete, and near Sta. Fe. Also from Venezuela.

519. Cnemidaria mutica var. contigua (Maxon) Stolze, Fieldiana, Bot. 37:37.
1974.

Hemitelia contigua Maxon, Contr. U. S. Natl. Herb. 16:32, t. 18. 1912. TYPE: 5 mi south of
Cartago, Pcia. Cartago, ca. 18(X) m, Maxon 523 (NY fragm US).

Plants terrestrial, at 1200-2100 m elevation, in forests, from El Socorro de S.
Ramon (Pcia. Alajuela), the Cordillera Central, and the north end of the
Cordillera de Talamanca.

520. Cnemidaria mutica var. grandis (Maxon) Stolze, Fieldiana, Bot. 37:36. 1974.

Hemitelia grandis Maxon, Contr. U. S. Natl. Herb. 16:37, t. 23. 1912. TYPE: Vicinity of

Coiiblanco, Pcia. Cartago, ca. 1950 m, Maxon 307 (US; isotype NY).

Hemitelia mdis Maxon, Contr. U. S. Natl. Herb. 17:413, t. 16. 1914. TYPE: Holcomb’s Trail
above Boquete, Pcia. Chiriqm, ca. 1750 m, Maxon 5682 (US; isotypes CR, GH not seen, NY).

Plants terrestrial, at 900 - 1900 m elevation, in forests, from north of S. Ramon,
the Cordillera Central, Tuis (Pcia. Cartago), the Cordillera de Talamanca to Pcia.
Chiriqm, and near Sta. Fe.

521. Cnemidaria mutica (Christ) Tryon, Contr. Gray Herb. 200:52. 1970, var.
mutica.

Hemitelia mutica Christ, Bull. Soc. Bot. Geneve II, 1:233. 1909. LECTOTYPE: Turrialba, Pcia.
Cartago, 850 m, Werckle (P not seen fragm US), chosen by Stolze (Fieldiana, Bot. 37:34. 1974).

Hemitelia horrida var. hetersora Rosenst. Repert. Spec. Nov. Regni Veg. 10:275. 1912. TYPE:
Rio Chis near Juan Vinas, Pcia. Cartago, ca. 1200 m, Brade & Brade 451 (S not seen fragm US).

Hemitelia pittieri Maxon, Contr. U. S. Natl. Herb. 16:32, t. 19a. 1912. TYPE: Valle de Agua
Buena, Canas Gordas, Pcia. Puntarenas, Pittier 10969 p. p. (US fragm F not seen). The type
collection is partly C. choricarpa (CR not seen).

Hemitelia arachnoidea Maxon, Contr. U. S. Natl. Herb. 16:34, t. 21a, b. 1912, non Cyathea
arachnoidea Hook., 1865. TYPE: Vicinity of La Palma, Pcia. S. Jose, 1450-1550 m, Maxon 451 (US;
isotype NY).

Cyathea suharachnoidea Domin, Rozpr. Kr^. Ceske. Spolecn. Nauk, Tf. Mat. Pfir. 2 [Pterid.
Dominica]: 264. 1929. TYPE: A renaming of Hemitelia arachnoidea Maxon, and so based on the
type of that name.

342 CYATHEACEAE: 63. CNEMIDARIA

CYATHEACEAE: 64. CYATHEA 343

Plants terrestrial, at (400)1000-2100 m elevation, in forests, from the
Cordillera de Tilaran, the Cordillera Central, the Cordillera de Talamanca to
Pcia. Chiriqui, and northwest of Sta. Fe. Also from Venezuela.

522. Cnemidaria spectabilis var. colombiensis Stolze, Fieldiana, Bot. 37:63, f. 21.

1974.

TYPE: Cordoba, Depto. Valle, Colombia, 80-100 m, Killip 5039 (US; isotypes
GH not seen, NY).

Plants terrestrial, at 0-500 m elevation, in forests, from the central Choco.

522a. Cnemidaria stolzeana G6mez, Phytologia 50:457. 1982.

TYPE: Beyond sawmill on lumber road above El Cope, Pcia. Code, 900 m,
Hammel 1036 (CR not seen; isotype MO).

Plants terrestrial, at 900 -ca. 1500 m, in forests, known only from the type and
from beyond the R. Tres Brazos 9 km from Sta. Fe {Croat 25601, MO).

64. CYATHEA J. E. Smith

Plants terrestrial, arborescent; caudices erect, woody, sometimes with a thick
covering of adventitious roots, especially at the base, scaly, the scales concolorous
or bicolorous, sometimes erose or toothed; stipes stout, short to long, smooth or
spiny, scaly and sometimes hairy, the scales concolorous or bicolorous, the
marginal cells different in shape and orientation from the central ones, rarely
bearing spine-like marginal cells, the hairs minute, the stipes also scurfy with
minute, appressed hairs or amorphous scales; fronds large, monomorphic;
laminae ovate to oblong, rarely pinnate, usually 2-pinnate-pinnatifid or
occasionally more divided, tapered to a non-conform, pinnatifid apex, or 1- or 2-
pinnate and often with a conform apex, chartaceous to coriaceous; veins free or
rarely forming costal areolae, simple to 2-forked; sori lateral or rarely terminal on
the veins, round, exindusiate or indusiate, the indusium an entire or partial,
evanescent sphere, a firm, cup- or saucer-like structure, or a firm to thin, obvious
to obscure scale sometimes hidden by the sporangia; paraphyses present or absent,
or rarely the sporangia mixed with a few scales; sporangia short-stalked.

Neotropical with a few species in the Western Pacific; ca. 120 species.
BARRINGTON, D. S. 1978. A revision of the genus Trichipteris. Contr. Gray Herb. 108:1-93.
LELLINGER, D. B. 1988. The disposition of Trichopteris (Cyatheaceae). Amer. Fern J. 78:90-94.
TRYON, R. M., Jr. 1976. A revision of the genus Cyathea. Contr. Gray Herb. 206:19 - 98.
WINDISCH, P. G. 1977. Synopsis of the genus Sphaeropteris (Cyatheaceae) with a revision of the
neotropical exindusiate species. Bot. Jahrb. Syst. 98:176-198.

FIGS. 519-532. Cnemidaria and Cyathea. FIG. 519. Median portion of median pinna of Cn. mutica
var. conti^a, Maxon 8220. FIG. 520. Median portion of median pinna of Cn. mutica var. grandis,
Lellinger 111. FIG. 521. Median portion of median pinna of Cn. mutica var. mutica, Killip 5156.
FIG. 522. Median portion of median pinna of Cn. spectabilis var. colombiensis, Killip ?>5?>11. FIG.
523. Pinnule of Cy. acutidens, Lellinger 804. FIG. 524. Basal segment of pinnule of Cy. aterrima,
Cuatrecasas 15190, Colombia. FIG. 525. Ultimate segment of Cy. caracasana var. maxonii, Scamman
& Holdridge 7868. FIG. 526. Pinna apex of Cy. conformis. Stern & Chambers 188. FIG. 527. Basal
portion of median pinna of Cy. costaricensis, Mickel 2861. FIG. 528. Pinna of Cy. decorata, Killip &
Cuatrecasas 38914, Colombia. FIG. 529. Basal portion of pinnule of Cy. delgadii. White & Lucansky
196823. FIG. 530. Pinnule of Cy. divergens var. divergens, Maxon 351. FIG. 531. Median pinna of Cy.
falcata, Sytsma, Andersson & Brenner 4415 (MO). FIG. 532. Pinnule of Cy. fulva, Wilbur & Stone
10623.

344 CYATHEACEAE: 64. CYATHEA

1. Sori exmdusiate..l5.

1. Sori indusiate, the indusia various, from fully enclosing the sori to a small, basal, scale-like
structure sometimes hidden by the sporangia..2.

2(1). Indusia completely enclosing the immature sori, fragile and usually evanescent, often
persisting only as an irregular, basal fragment.. 10.

2(1). Indusia partially surrounding the immature sori, commonly firm, often splitting into a few
fragments at maturity or remaining as an entire or partial basal cup or small scale-like structure
sometimes hidden by the sporangia..3.

3(2). Pinnae pinnate to 2-pinnate..5.

3(2). Piimae entire to pinnatifid..4.

4(3). Rhizome and stipe base scales whitish; laminae elliptic-lanceolate, acute at the base and
apex, ca. 1 m long, 25 cm wide; rachises and costae densely pilose.

528. C. decorata

4(3). Rhizome and stipe base scales brown with white margins; laminae lanceolate, obtuse at the
base, the basal pinnae deflexed, narrowly acute at the pinnatifid apex, ca. 30 - 35 cm long, 12-15 cm
wide; rachises densely pilose; costae sparsely scaly.

545. C. pseudonanna

5(3). Pinnules, or most of them, entire to lobed..8.

5(3). Pinnules pinnatifid to pinnate..6.

6(5). Pinnules stalked 1-4 mm; ultimate segments more than 1 cm long, entire, the basal ones
usually slightly tapered at the base and not fully adnate. Stipe base and lamina scales pale, the larger
ones with a reddish-brown center; basal pinnule segments smaller than the distal ones.

534. C. holdridgeana

6(5). Pinnules sessile or stalked no more than 1 mm; ultimate segments conspicuously toothed, or
if entire then less than 1 cm long, the basal ones broadly adnate..?.

7(6). Costae and veins conspicuously pilose on both surfaces; veins (l)2-forked; scales on the
abaxial surface of the costae bullate or plane, whitish; ultimate segments decidedly serrate;
receptacle with stout paraphyses.

523. C. acutidens

7(6). Costae and veins sparsely strigose or lacking hairs on both surfaces; veins l(2)-forked; scales
on the abaxial surface of the costae bullate or plane, pale brown, golden, or rarely dark brown;
ultimate segments often entire or crenate; receptacles lacking stout paraphyses.

538. C multiflora

8(5). Laminae gradually or abruptly tapered to a pinnatifid apex. Caudex up to 3 m long; pinnae
lanceate, tapered to a non-articulate, pinnatifid apex; pinnules up to 2.5 cm wide, not articulate;
medial veins often anastomosing to form a subcontinuous series of costal areolae; sori submarginal.

541. C. petiolata

8(5). Laminae with a conform terminal pinna, not a pinnatifid apex..9.

9(8). Lateral pinnules cordate at the base; apical pinnules articulate; laminae glabrous on the
abaxial surface; sori submarginal. Caudex up to 8 m long; pinnae linear; pinnules up to 1.5 cm wide;
medial veins occasionally anastomosing to form a discontinuous series of costal areolae.

526. C. conformis

9(8). Lateral pinnules obtuse at the base; apical pinnules not articulate; laminae short-pubescent
abaxially; sori nearly medial. Pinnules ca. 1 cm wide; medial veins not anastomosing.

535. C. impar

10(2). Basal and suprabasal pinnules sessile or nearly so.. 12.

10(2). Basal and suprabasal pinnules stalked (1.5)2 mm or more.. 11.

11(10). Rachises and stipes stramineous; costae stramineous; basal and suprabasal pinnules stalked
2-10 mm; stipe base scale margins whitish. Pinnules up to 3.5 cm wide, glabrous except for a few
deciduous scales on the abaxial surface of the costules; stipes up to 3 m long.

530. C. divergens var. divergens

CYATHEACEAE: 64. CYATHEA 345

11(10). Rachises and stipes atropurpureous; costae atropurpureous to brown; basal and suprabasal
pinnules stalked (1.5)2 -8 mm; stipe base scale margins golden. Pinnules up to 3(3.5) cm wide,
glabrous abaxially except for a few deciduous scales on the costules; stipes up to 2 m or more long.

533. C gracilis

12(10). Adaxial surface of the segments bearing subdeciduous, short, lax, irregularly twisted hairs
on and sometimes between the veins. Stipe base scales nearly linear, concolorous, medium brown
with shghtly paler margins; segments sparsely pilosulous on the abaxial surface of the veins and with
bullate and plane, whitish to tan scales on the costules.

548. C. suprastrigosa

12(10). Adaxial surface of the segments glabrous or with short to long, stiff hairs only on the
veins.. 13.

13(12). Veins of the segments on the abaxial surface with slightly curved, whitish, stiff hairs ca. 0.75
mm long. Stipe base scales nearly linear, concolorous, medium brown with shghtly paler margins;
golden brown, bullate or plane scales present on the abaxial surface of the costae and costules.

529. C. delgadii

13(12). Veins of the segments on the abaxial surface glabrous (translucent hairs sometimes present
on the costules).. 14.

14(13). Costules of the segments lacking hairs on the abaxial surface; pinnules of median pinnae
(1)1.5 “2 cm wide; stipe base scales mostly bicolorous with a dark, central area and wide to narrow,
pale margins.

525. C. caracasana var. maxonii

14(13). Costules of the segments with lax, translucent hairs on the abaxial surface; pinnules of
median pinnae 1 - 1.5(2) cm wide; stipe base scales concolorous, medium brown with paler margins.

532. C.fulva

15(1). Fronds pinnate or pinnate-pinnatifid..l8.

15(1). Fronds 2-pinnate-pinnatifid..l6.

16(15). Costules usually with many, long, straight or curved hairs 0.25-1.25(2) mm long on the
abaxial surface.. 22.

16(15). Costules with a few short, straight hairs less than 0.5 mm long or with cobwebby hairs on
the abaxial surface. Short, stiff hairs absent between the veins on the abaxial surface of the

laminae.. 17.

17(16). Rachises and costae abaxially glabrous or nearly so..26.

17(16). Rachises and costae abaxially with a close covering of matted hairs and small scales..28.

18(15). Fronds pinnate-pinnatifid or rarely more divided, the pinnae deeply lobed, usually
pinnatifid..21.

18(15). Fronds pinnate, the pinnae entire to shallowly lobed.. 19.

19(18). Plants diminutive, the fronds ca. 0.5 m long, ca. 5- 10 cm wide; sori in 1 series on each side
of the costae; pinnae entire to shallowly lobed.

531. C falcata

19(18). Plants medium-sized, the fronds ca. 1 m long, more than 10 cm wide; sori in 2 or 3 series
on each side of the costae; pinnae entire to repand..20.

20(19). Vein groups often anastomosing to form costal areolae; laminae glabrous abaxially.
Laminae ca. 60 cm long, 20 cm wide; sori in 2 rows on each side of the costae.

553. C. williamsii

20(19). Vein groups not anastomosing or forming costal areolae; laminae sparsely castaneous-
glandular abaxially. Laminae ca. 40 - 50 cm long, 20 cm wide; sori in 2 or 3 rows on each side of the
costae.

547a. C. stolzei

21(18). Stipes predominantly with narrow, brownish scales; laminae 1- 1.5 m long, 30 - 45 cm wide;
basal pinnae less than 1/2 as long as the longest pinnae.

550. C. ursina

346 CYATHEACEAE: 64. CYATHEA

21(18). Stipes predominantly with narrow, whitish hairs; laminae 30 - 75 cm long, 15-25 cm wide;
basal pinnae 2/3 as long as the longest pinnae.

542. C. phalaenolepis

22(16). Veins on the adaxial surface of the laminae glabrous..25.

22(16). Veins on the adaxial surface of the laminae with straight to curved hairs..23.

23(22). Segments entire or faintly crenate; scales on the costules longer than the hairs, not bullate,
irregularly ciUate. Paraphyses very twisted, forming a mat, not exceeding the sporangia.

536. C. lockwoodiana

23(22). Segments decidedly crenate-serrate; scales on the costules or veins shorter than the hairs,
bullate, not ciliate..24.

24(23). Stipes spiny, the scales few and deciduous; pinnae caudate at the apex. Fronds 2-3.5 m
long; stipe bases spiny; largest pinnules 15-25 mm wide; scales on the adaxial surface of the costules
lanceolate, white, ca. 2 mm long; scales on the costules of the ultimate segments bullate, white.

549. C. trichiata

24(23). Stipes not spiny, the scales abundant, linear, distantly toothed, reddish-tan, persistent;
pinnae acute to acuminate at the apex. Paraphyses nearly straight, exceeding the sporangia; scales of
the costae linear, whitish, plane.

524. C. aterrima

25(22). Laminae abaxially hairy only on the costules and veins. Fronds 2-4 m long; largest
pinnules 13-15(20) mm wide; costules abaxially with broadly lanceolate, reddish-brown scales.

547. C. stipularis

25(22). Laminae abaxially hairy between the veins, as well as on the costules and veins. Fronds up
to 2.5 m long; laminae ovate, lanceolate, or oblanceolate, ca. 1.5-1.75 m long, 0.8-1 m wide;
costules of the ultimate segments with bullate, whitish scales.

543. C pilosissima

26(17). Sori bearing 1-few, ovate-acuminate, whitish scales at the base of the receptacle. Stipe
bases very shghtly spiny; fronds ca. 3 m long; rachises and costae pale brown to nearly stramineous;
piimules acute-acuminate; ultimate segments up to 2.5 mm wide, crenate-serrate, obtuse to acute at
the apex; scales on the costules and veins plane or nearly so, whitish to pale brown, fimbriate.

527. C. costaricensis

26(17). Sori not bearing basal scales.. 27.

27(26). Costules and veins glabrous or sparsely hairy on the abaxial surface, the hairs commonly
irregularly branched, appearing somewhat stellate; pinnules spreading.

544. C. poeppigii

27(26). Costules and veins villous on the abaxial surface; pinnules ascending. Stipe base scales
linear, ca. 25 mm long, 1.5 mm wide.

551. C. villosa

28(17). Costae sparingly spiny. Costules sparingly scaly on the abaxial surface, the scales lanceate
or acicular; paraphyses pale; pinnules 10 - 18(20) mm wide.

537. C microdonta

28(17). Costae not spiny..29.

29(28). Costae and costules abaxially with dark brown, predominantly bullate scales ca. 0.3 -0.5
mm long and wide. Stipes spiny, ca. 1 m long; laminae lanceolate or ovate-lanceolate, ca. 1.5 m long,
1.2 m wide; largest pinnules ca. 15 mm wide.

546. C. schiedeana

29(28). Costae and costules abaxially with predominantly lanceolate to Unear, plane or slightly
bullate scales more than 0.5 mm long..30.

30(29). Pinnules 2-2.5 cm wide; rachises and costae abaxially with small, irregular, brown,
subbuUate scales with a long, bristle-like apex. Pinnules gradually attenuate at the apex, the costules
abaxially with brown, plane, fimbriate scales.

552. C. wendlandii

CYATHEACEAE: 64. CYATHEA 347

30(29). Pinnules less than 2(2.5) cm wide; rachises and costae lacking such scales on the abaxial
surface..31.

31(30). Laminae pale brown abaxially; narrow, linear scales 2-5 mm long present on the costae,
especially at the pinnule bases; hairs absent. Stipes spiny, ca. 0.5 -0.7 m long; laminae ovate or
ovate-lanceolate, 1.2- 1.5 m long, 0.7 -0.8 m wide.

539. C. nigripes var. brunnescens

31(30). Laminae pale green abaxially; narrow, rather stiff, straight hairs present on the abaxial
surface of the costae; scales absent. Stipes spiny, 0.6 -0.8 m long; laminae ovate-lanceolate, 1.4- 1.7
m long, ca. 1 m wide.

540. C. nigripes var. nignpes

523. Cyathea acutidens (Christ) Domin, Pteridophyta 262. 1929.

Alsophila leucolepis wm.puhescens ChihimViii. Prim. FI. Costaric. 3(1):42. 1901. SYNTYPES:
Cerro Canas Gordas, Pcia. Puntarenas, 1100 m, Pittier 10981 (BR or P not seen; isosyntypes CR, UC,
US), 10989 (BR not seen photo 4899 fragm NY; isosyntypes CR, NY, US), and 10992 (BR not seen
photo 4898; isosyntypes CR, P not seen, US).

Alsophila acutidens Christ, Bull. Herb. Boissier II, 6:186. 1906. TYPE: Based on Alsophila

leucolepis \ar.pubescens Christ in Pitt., and so based on the syntypes of that name.

Plants terrestrial, at 1100-1200 m elevation, in forests, from the Fila Costeha.

524. Cyathea aterrima (Hook.) Domin, Pteridophyta 262. 1929.

Alsophila aterrima Hook. Syn. Fil. 38. 1866. TYPE: Near Tarapoto, Depto. S. Martin, Peru,
Spruce 4713 (K not seen; isotypes BM not seen, P not seen, US).

Alsophila mollicula Maxon, J. Arnold Arbor. 27:440. 1946. TYTE: Between Sta. Marta and

Marsella, Depto. Caqueta, Colombia, Woronov & Juzepczuk 6415 (US; isotypes LE not seen Tryon
photo not seen).

Dryopteris macarenensis Alston, Mutisia 7:5. 1952. TYPE: North escarpment of the Sierra de la
Macarena, Intend. Meta, Colombia, 8(X) m, Philipson 2281 (BM not seen; isotype COL not seen).

Alsophila scopulina Tryon, Rhodora 62:2, f. 2. 1960. TYPE: Summit of Cerro Isibukuri, Rio
Kananari, Depto. Vaupes, Colombia, Schultes & Cabrera 13411 (GH not seen; isotypes B not seen,
BM not seen, NY not seen, U not seen, US).

Alsophila lechria Tryon, Rhodora 62:4, t. 1251. 1960. TYPE: Mesa de Los Santos, Depto.

Santander, Colombia, 1500 m, Killip & Smith 15202 (GH not seen; isotypes NY not seen, US).

Plants terrestrial, at 1400-1800 m elevation, in forests, in the Flora area known
only from the northwest side of Alto del Buey (Lellinger & de la Sot a 246, BM,
COL, CR, HU A, LP, US). Also from Venezuela, Colombia and Peru.

525. Cyathea caracasana var. maxonii (Underw. in Maxon) Tryon, Contr. Gray
Herb. 206:83. 1976.

Cyathea membranulosa Christ, Bull. Herb. Boissier II, 7:271. 1907. SYNTYPES: La Palma, Pcia.
S. Jose, 1500 m, Werckle 17080 (P not seen; isotype CR not seen), and S. Pascon, Costa Rica, 1500 m,
Werckle 17024 (P not seen; isotype CR not seen).

Cyathea maxonii Underw. in Maxon, North Amer. FI. 16:82. 1909, as ''maxoni.” TYPE:

Mountains 5 mi south of Cartago, Pcia. Cartago, 1800 m, Maxon 524 (NY; isotype US).

Plants terrestrial, at 1100-2900 m elevation, in forests, from north of S. Ramon,
the Cordillera Central, the Cordillera de Talamanca to Pcia. Chiriquf, and the Fila
Costeha.

526. Cyathea conformis (Tiyon) Stolze, Fieldiana, Bot. 37:80. 1974.

Hemitelia conformis Tryon, Rhodora 62:1, f. 1. 1960. TYPE: Pinas Bay, Pcia. Darien, Stem &
Chambers 188 (GH; isotypes A not seen, MO, US, Y not seen).

348 CYATHEACEAE: 64. CYATHEA

Plants terrestrial, at 0-100 m elevation, in forests, from the Pacific coast of
Pcia. Darien. Also from the Pacific coast of Colombia south of the Choco.

527. Cyathea costaricensis (Klotzsch ex Kuhn) Domin, Acta Bot. Bohem. 9:107.
1930.

Hemitelia costaricensis Klotzsch ex Kuhn, Linnaea 36:159. 1869. LECTOTYPE: “Costa Rica et
Veragua,” Warscewicz 36 (B not seen fragms NY, US), chosen by Barrington (Contr. Gray Herb.
208.76. 1978).

Plants terrestrial, at 100-1500(2400) m elevation, in forests, from the Pacific
slopes of the Cordillera de Tilaran, La Palma, near El Empalme (Pcias.
Cartago-S. Jose), near Remedios (Pcia. Chiriqm), and Cerro Galera ca. 2 km
from the Pacific Ocean (Canal Zone). Also from Mexico to Nicaragua.

528. Cyathea decorata (Maxon) Tryon, Contr. Gray Herb. 200:50. 1970.

Hemitelia decorata Maxon, J. Arnold Arbor. 21:439, t. 1. 1946. TYPE: Along the Rio

Yurumangui, Depto. El Valle, Colombia, 5-50 m, Cuatrecasas 15737 (US; isotype F not seen).

Plants terrestrial, at 0-300 m elevation, from the upper Rio S. Juan area. Also
from Depto. El Valle, Colombia.

529. Cyathea delgadii Sternb. Vers. FI. Vorwelt 1:47, t. B. 1820.

Cyathea vestita Mart. Denkschr. Konigl.-Baier. Bot. Ges. Regensburg 2:146. 1822. SYNTYPES:
“Prov. Sebastianopohtanae” [Est. Rio de Janeiro, Brazil], Martins (M not seen) “et Pauliae” [Est. S.
Paulo, Brazil], Martins (M not seen); and Serra dos Pilaoes, Est. Goias, Brazil, Pohl (M? not seen),
according to Martins (Icon. PI. Crypt. Bras. 75. 1834).

Cyathea oligocarpa Kunze, Linnaea 9:101. 1834. TYPE: Pampayacu, Depto. Huanuco, Peru,
Poeppig Diar. 1101 (LZ destroyed; isotypes Poeppig 218, B not seen, MO, P not seen). Poeppig’s
diary numbers and his specimen numbers are not identical.

Cyathea hirtnla Mart. Icon. PI. Crypt. Bras. 76, t. 53. 1834. TYPE: Alamada, Serra do Mar, Est.
Bahia, Brazil, Maxmillian von Wied-Nenwied (LE? or M? not seen photo BR, fragm BM not seen).

Cyathea schanschin Mart. Icon. PI. Crypt. Bras. 77, t. 54. 1834. SYNTYPES: Presumably based
on several Martins collections from Brazil (M none seen), Peru, Haenke (BR? not seen), and Peru,
Poeppig (BR? not seen).

Cyathea denticnlata Goldm. in Meyen, Nov. Actorum Acad. Caes. Leop.-Carol. Nat.Cur. 19, Abh.
1:466. 1843. TYPE: Corcovado, Est. Rio de Janeiro, Brazil, Meyen (B not seen).

Cyathea feei Glaziou in Fee, Crypt. Vase. Bresil 1:179, t. 66, f. 2. 1869. LECTOTYPE: Rio de
Janeiro, Est. Rio de Janeiro, Brazil, Glazion 2286 (P not seen photo GH not seen fragm NY not
seen), chosen by Tryon (Contr. Gray Herb. 206:72. 1976).

Cyathea sphaerocarpa Fee, Crypt. Vase. Bresil 1:180, t. 53, f. 2. 1869. TYPE: Rio de Janeiro, Est.
Rio de Janeiro, Brazil, Glazion 2283 (P not seen Tryon photo GH not seen).

Cyathea abmptecandata F6e, Crypt. Vase. Br6sil 1:183, t. 62, f. 2. 1869. TYPE: Rio de Janeiro,
Est. Rio de Janeiro, Brazil, Glazion 2284 (P not seen photo GH not seen fragm NY; isotype US).

Cyathea pilosa Baker, Syn. Fil. ed. 2. 19. 1874. TYPE: Tarapoto, Depto. S. Martin, Peru, Spmee
4729 (K not seen photo 8110; isotypes GH not seen, P not seen photo GH not seen, fragm F not
seen; fragm US).

Cyathea copelandii Kuhn & Luerssen in R. Copel. Abh. Naturwiss. Vereine Bremen 7:278. 1882.
TYPE: Ilha da Trindade, Brazil, R. Copeland (K? not seen; isotype fragm P not seen Tryon photo
GH not seen).

Cyathea hypotricha Christ, Bull. Herb. Boissier II, 4:947. 1904. TYPE: Costa Rica, Werckle (P not
seen fragm NY).

Cyathea bahiensis Rosenst. Repert. Spec. Nov. Regni Veg. 20:90. 1924. TYPE S. Gomala, Rio
Femmeas region, Est. Bahia, Brazil, Lnetzelbnrg 12534 (M not seen photo BM not seen).

CYATHEACEAE: 64. CYATHEA 349

Cyathea micromera Rosenst. Repert. Spec. Nov. Regni Veg. 20:90. 1924. TYPE: Apertada-hora,
Est. Goids, Brazil, Luetzelburg 12841 (M not seen photo BM not seen).

Cyathea trindadensis Brade, Arq. Inst. Biol. Veg. 3:1, t. 1, f. 1. 1936. TYPE: Ilha da Trindade,
Brazil, Campos Porto 579 (RB not seen; isotype F not seen).

TYPE: Gancho do Generale Delgado along the road to Caldas Novas, Est.
Goids, Brazil, Pohl (PRC not seen fragms GH not seen and probably US).

Plants terrestrial, at 500-1500(2900) m elevation, in forests, from north of S.
Ram6n, the Cordillera Central, Tuis and Platanillo (both Pcia. Cartago), the
Cordillera de Talamanca to Villa Mills (Pcia. Cartago), the Fila Costeha near S.
Vito, above Sta. Fe, El Valle, Cerro Campana and Cerro Jefe (both Pcia.
Panama), and Cerro Pirre. Also from Venezuela, Colombia to Bolivia, Guyana,
Brazil, Paraguay, and Argentina.

530. Cyathea divergens Kunze, Linnaea 9:100. 1834, var. divergens.

Cyathea equestris Kunze, Linnaea 9:100. 1834. TYPE: Cerro de Cristobal near Pampayacu,
Depto. Huanuco, Peru, July 1829, Poeppig (LZ destroyed fragm K not seen Tryon photo GH not
seen), synonymized by Mettenius in Tr. & Planch. (Ann. Sci. Nat. Bot. V, 2:265 (repr. 347). 1864).

Cyathea glohularis K. Presl, Epim. Bot. 30. 1849. TYPE: “Novae-Grenadae,” Linden (PRC? not
seen).

Cyathea equestris var. boconensis Karsten, Linnaea 28:456. 1857. TYPE: Paramo de Bocono,
Sierra de Merida, Edo. Trujillo, Venezuela, Karsten (LE? not seen).,

Cyathea petiolulata Karsten, FI. Columb. 2:163, t. 185. 1869. TYPE: Merida, Edo. Merida,

Venezuela, 2000 - 2500 m, Karsten (LE? not seen).

Cyathea calva Karsten, FI. Columb. 2:175, t. 192. 1869. TYPE: Near Escuque, Sierra de Merida,
Edo. Trujillo, Venezuela, 1000 m, Karsten (LE? not seen).

Cyathea firma Kuhn, Linnaea 36:163. 1869. TYPE: Merida, Edo. Merida, Venezuela, Funck &
Schlim 1228 (B not seen fragm US).

Alsophila subaspera Christ in Pitt. Prim. FI. Costaric. 3(1):43. 1901. SYNTYPES: Forests of
Copey, Pcia. S. Jose, Tonduz 11787 (BR or P not seen; isosyntypes A, CR, GH, NY, US), 11802 (BR
not seen photo 4943 fragm NY; isosyntypes B not seen, CR, P not seen), and 12183 (BR or P not
seen; isosyntypes CR, NY).

Cyathea petiolulata var. pastoensis Hieron. Bot. Jahrb. Syst. 34:437. 1904. TYPE: Near Altaquer
and S. Pablo, Cordillera de Pasto, Depto. Narino, Colombia, 900-1400 m, Lehmann 81 (B not seen).

Cyathea pelliculosa Christ, Bull. Herb. Boissier II, 4:946. 1904. SYNTYPES: Costa Rica, Werckle
45 (P not seen) and 50 (P not seen).

Alsophila latisecta Christ, Bull. Herb. Boissier II, 6:185. 1906. TYPE: Valley of the Rio Navarro,
Pcia. Cartago, Werckle 16767 (P not seen).

Cyathea divergens var. minor Rosenst. Repert. Spec. Nov. Regni Veg. 22:2. 1925. LECTOTYPE:
La Palma, Pcia. S. Jose, 1400 m, Brade & Brade 108 (S not seen; isolectotypes NY, UC), chosen by
Tryon (Contr. Gray Herb. 206:54. 1976).

Cyathea divergens var. hirta Losch, Mitt. Bot. Staatssaml. Miinchen 1(1):20. 1950. TYPE: Chirrip6
Grande, Pcia. S. Jose, 2000 m, Kupper 1265 (M not seen), synonymized by Tryon (Contr. Gray Herb.
206:55. 1976).

TYPE: Near Pampayacu, Depto. Huanuco, Peru, Jul 1869, Poeppig Diar. 1163
(LZ destroyed).

Plants terrestrial, at (400)1200-2700(3200) m elevation, from Volcan Orosi
(Pcia. Guanacaste), the Cordillera Central, the Fila de Cedral and Cerro Tablazo,
the Cordillera de Talamanca to Pcia. Chiriquf, and the valley of the Rfo General.
Also from Venezuela, Colombia to Peru, and Guyana.

350 CYATHEACEAE: 64. CYATHEA

533 Cy. gracilis

543 Cy. pilosissima

542 Cy. phalaenolepis

535 Cy. impar

CYATHEACEAE: 64. CYATHEA 351

Alsophila latisecta Christ is a 3-pinnate-pinnatifid form whose description
matches a specimen collected above Purisil, Pcia. Cartago, not far from the type
locality {Lellinger 1537, CR, US).

531. Cyathea falcata (Kuhn) Domin, Pteridophyta 262. 1929.

Alsophila falcata Kuhn, Linnaea 36:155. 1869. TYPE: Gorgona Island, Depto. Narino, Colombia,
Seemann (K not seen fragms NY not seen US).

Nephrodium kuhnii Hieron. Bot. Jahrb. Syst. 34:440. 1904. TYPE: Cordillera de Pasto, Depto.
Narino, Colombia, 1000 m, Lehmann 16 (K not seen; isotype US).

Plants epipetric, at 600 m elevation, in the Flora area known only from near
Llano Grande, Pcia. Code (Sytsma, Andersson & Brenner 4415, MO). Also from
Pacific coastal Colombia, although not yet recorded from the Choco.

532. Cyathea fulva (Martens & Galeotti) F6e, Cat. Foug. Mex. [M6m. Foug. 9]:34.
1857.

Alsophila fulva Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15:78, t. 23. 1842.
TYTE: Talea, Cordillera Occidental, Edo. Oaxaca, Mexico, Galeotti 6346 (BR not seen photos 4877,
4905 fragm US; isotype P not seen fragm UC).

Cyathea schlechtendalii Kunze, Bot. Zeitung (Berlin) 3:288. 1845. TYPE: Venezuela,

Schlechtendal (LZ destroyed), synonymized by Tryon (Contr. Gray Herb. 206:66. 1976).

Cyathea aurea Klotzsch, Allg. Gsu'tenzeitung 24:105. 1856. AUTHENTIC MATERIAL: Between
Petaquire and Colonia Tovar, Venezuela, Karsten 7 (B not seen fragm NY), synonymized by Tryon
(Contr. Gray Herb. 206:66. 1976).

Cyathea aurea var. squamosa Karsten, Linnaea 28:459. 1857. TYTE: Between Petaquire and
Colonia Tovar, Venezuela, Karsten (LE? not seen; isotype B not seen).

Cyathea furfuracea Christ, Bull. Herb. Boissier II, 4:950. 1904, non Baker, 1874, nom. illeg.

TYTE: Costa Rica, Werckle (P not seen; isotype NY fragm US).

Cyathea onusta Christ, Bull. Herb. Boissier II, 4:950. 1904. TYTE: Costa Rica, Werckle 41 (P not
seen; isotype US).

Cyathea papyracea Christ, Bull. Herb. Boissier II, 4:946. 1904. LECTOTYTE: Costa Rica,

Werckle 52, second sheet (P not seen), chosen by Gastony (Contr. Gray Herb. 203:147. 1973).

Cyathea conspersa Christ, Bull. Herb. Boissier II, 5:260. 1905. TYTE: A renaming of Cyathea
furfuracea Christ, and so based on the type of that name.

Cyathea underwoodii Christ, Bull. Herb. Boissier II, 6:183. 1906. SYTTrYPES: Navarro, Pcia.
Cartago, Werckle in 1903 (P not seen; isosyntype NY), Werckle in 1901 - 1905 (P not seen; isosyntypes
NY, US), and Werckle in 1905 (P not seen).

Cyathea delicatula Maxon, Contr. U. S. Natl. Herb. 13:4. 1909. TYPE: Between Tactic and
Coban, Depto. Alta Verapaz, Guatemala, von Tuerckheim II 1629 (US; isotype GH not seen).

Cyathea mollis Rosenst. Repert. Spec. Nov. Regni Veg. 22:2. 1925, non Copel, 1917, nom. illeg.
LECTOTYTE: La Pahna, 1400 m, Brade & Brade 631 (S not seen; isolectotypes B, US fragm NY),
chosen by Tryon (Contr. Gray Herb. 206:68. 1976).

FIGS. 533-543. Cyathea. FIG. 533. Median pinnule of upper pinna of Cy. gracilis, Brade 853. FIG.
534. Piimule of Cy. holdridgeana, Nisman 104. FIG. 535. Apex of lateral pinna of Cy. impar, Tyson,
Dwyer & Blum 3264. FIG. 536. Ultimate segment of Cy. lockwoodiana. Stern et al. 528. FIG. 537.
Pinnule base of Cy. microdonta, Kilhp 2540. FIG. 538. Ultimate segment of Cy. multiflora. Burger &
Stolze 5876. FIG. 539. Basal portion of median pinnule of Cy. nigripes var. brunnescens, Lelhnger &
de la Sota 428. FIG. 540. Basal portion of median pinna of Cy. nigripes var. nigripes, Kilhp 35132.
FIG. 541. Apex of lateral pinna of Cy. petiolata, Kilhp 2802. FIG. 542. Pinna of Cy. phalaenolepis,
Kilhp 35347. FIG. 543. Basal portion of median pinnule of Cy. pilosissima, Lehinger & de la Sota
714.

352 CYATHEACEAE: 64. CYATHEA

Cyathea molliuscula Domin, Acta Bot. Bohem. 9:138. 1930. TYPE: A renaming of Cyathea mollis
Rosenst., and so based on the type of that name.

Plants terrestrial, at 700-3100 m elevation, in forests, from Monteverde, the
Cordillera Central, the Cordillera de Talamanca to Pda. Chiriqm, La Mesa (Pda.
Code), Cerro Campana and Campo Tres (both Pda. Panama), Cerro Pirre, and
Alto del Buey. Also from Mexico, Guatemala, Honduras, Nicaragua, Venezuela,
and Colombia.

533. Cyathea gracilis Griseb. FI. Brit. W. Ind. 704. 1864.

TYPE: Fox’s Gap, Jamaica, Purdie (K not seen; isotype BM not seen).

Plants terrestrial, at 1000-1500 m elevation, in forests, from La Palma and
vicinity and the Finca Haberle near Cerro Tablazo. Also from Jamaica and
Colombia.

534. Cyathea holdridgeana Nisman & Gdmez, Amer. Fern J. 61:168, t. 26. 1971.
TYPE: La Chonta, at Km. 55 of the Interamerican Highway, Pcia. Cartago, 2200

m, Nisman S. 104 (CR; isotypes F not seen, GH not seen).

Plants terrestrial, at 2200 m elevation, in the Flora area known only from the
central northeastern slopes of the Cordillera de Talamanca.

This species is thought by Tryon (Contr. Gray Herb. 206:90. 1976) to be a
hybrid between C. divergens and C. stipularis.

535. Cyathea impar Tryon, Contr. Gray Herb. 206:42, f. 59, 60. 1976.

TYPE: Northeast slope of Cerro Jefe, on the road to Buenos Aires, Pcia.
Panama, 2600 ft, Tyson, Dwyer & Blum 3264 (MO).

Plants terrestrial, at 700-800 m elevation, in forests, from Cerro Jefe, the Sta.
Rita ridge road (Pcia. Colon), and north of Cerro Mali (Pcia. Darien).

536. Cyathea lockwoodiana (Windisch) Lellinger, Amer. Fern J. 74:57. 1984.

Sphaeropteris lockwoodiana Windisch, Bradea 2:57. 1976. TYPE: Cerro Chiribiquete, Rfo

Macaya (tributary of the Rio Apaporis), Comis. Vaupes, Colombia, Schultes & Cabrera 5635 (GH
not seen; isotype US).

Plants terrestrial, at 0-800 m elevation, in forests, from Las Cascadas (Canal
Zone), the Rfo La Maestra (Pcia. Panama), the vicinity of Cana, and Mamey
(Pcia. Darien). Also from Venezuela and Colombia.

537. Cyathea microdonta (Desv.) Domin, Pteridophyta 263. 1929.

Polypodium microdontum Desv. Ges. Naturf. Freunde Berlin Mag. 5:319. 1811. TYPE: “Am.
calidiore,” collector unknown (P not seen photo 3427).

Polypodium aculeatum Raddi, Opusc. Sci. 3:288. 1819; PI. Bras. Nov. Gen. 1:27, t. 42. 1825, non
L., 1753, nom. iUeg. TYPE: Rio de Janeiro, Est. Rio de Janeiro, Brazil, Raddi (FI not seen).

Alsophila armata Mart. Icon. PI. Crypt. Bras. 72, t. 28, 48. 1834. TYPE: A renaming of P.
aculeatum Raddi, and so based on the type of that name.

Alsophila ferox K. Presl, Tent. Pterid. 62. 1836. TYPE: A renaming of P. aculeatum Raddi, and so
based on the type of that name.

Plants terrestrial, at 0-1100 m elevation, in forests, from the foothills of the
Cordillera Central and the Atlantic coastal plain of Costa Rica and Panama, the
valley of the Rfo General, Chorrera (Pcia. Panama), the Canal Zone, and the

CYATHEACEAE: 64. CYATHEA 353

central Choco. Also from Cuba, Jamaica, Hispaniola, Trinidad, Mexico to
Nicaragua, Colombia, Peru, Brazil, and the Guianas.

538. Cyathea multiflora J. E. Smith, M^m. Acad. Roy. Sci. (Turin) 5:416. 1793.

Hemitelia nigricans K. Presl, Abh. Konigl. Bohm. Ges. Wiss. V, 6:391 (repr. 31). 1851, non

Cyathea nigricans Mett., 1863. TYPE: Rio S. Juan, Guatemala, Friedrichsthal (W not seen fragms
NY, US).

Hemitelia atrovirens Trevis. Atti 1st. Veneto II, 2:164. 1851. TYPE: Rio S. Juan, Guatemala,
Friedrichsthal 639 (FI? not seen).

Hemitelia obscura Mett. in Tr. & Planch. Ann. Sci. Nat. Bot. V, 2:264. 1864, non Cyathea obscura
(Bedd.) Copel, 1909. TYPE: “Prov. de Barbacoas, via de Tuquerres,” Depto. Narino, Colombia,
1600 m, Triana 189 (B not seen fragm NY; isotypes K not seen fragm NY, P not seen).

Hemitelia denticulata Hook. Syn. Fil. 31. 1865. TYPE: Presumably Panama, Cuming 1360 (K not
seen), according to Tryon (Contr. Gray Herb. 206:32. 1976).

Hemitelia lindigii Bdcer, Syn. Fil. ed. 2. 454. 1874. TYPE: Alto del Trigo, Depto. Cundinamarca,
Colombia, Lindig 310 (K not seen fragm US; isotype P not seen).

Hemitelia hartii Baker, J. Bot. Brit. For. 24:243. 1886. TYPE: Chiriqui Lagoon, Pcia. Bocas del
Toro, Hart 43 (K not seen; isotypes NY not seen, P not seen, US).

Alsophila leucolepis wax. pubescens Christ, Prim. FI. Costaric. 3(1):42. 1901. SYNTYPES: Cahas
Gordas, Pcia. Puntarenas, Pittier 10981 (BR or P not seen; isosyntype US), Pittier 10982 (BR or P not
seen); and Pittier 10992 (P not seen; isosyntype US).

Alsophila decussata Christ in Pitt. Prim. FI. Costaric. 3(1):41. 1901. TYPE: El General, Diquis
basin, Pcia. Puntarenas, 600 m, Pittier 12027 (BR not seen photo 4903; isotypes CR, NY, P not seen,
US).

Alsophila pinnula Christ in Pitt. Prim. FI. Costaric. 3(1) :43. 1901. TYPE: Not stated, but

presumably Marais de la Rio Matina, Pcia. Limon, 20 m, Pittier 10267 (P not seen photo 3438;
isotype US), according to Barrington (Contr. Gray Herb. 208:81. 1978).

Hemitelia squarrosa Rosenst. Repert. Spec. Nov. Regni Veg. 22:2. 1925. TYPE: Finca

Hundriesser, Pcia. Limon, Brade & Brade 405 {Ros. Fil Costaric. Ejcs. 195) (S not seen; isotypes B
not seen, UC, US fragm NY).

Cyathea austroamericana Domin, Pteridophyta 263. 1929. TYPE: A renaming of H. nigricans K.
Presl, and so based on the type of that name.

Cyathea columbiana Domin, Pteridophyta 263. 1929. TYPE: A renaming of Hemitelia obscura
Mett., and so based on the type of that name.

Hemitelia squamulosa Losch, Mitt. Bot. Staatssaml. Miinchen 1:20. 1950. TYPE: Orosi, Pcia.
Cartago, 1100 m, Kupper 798 (M not seen photo BM not seen), synonymized by Tryon (Contr. Gray
Herb. 206:32. 1976).

Cyathea leucolepismata Alston, J. Wash. Acad. Sci. 48:231. 1958. TYPE: Near S. Diego de
Colorado, between Umbria and Puerto Asis, Depto. Putumayo, Colombia, Ewan 16748 (BM not
seen; isotypes UC, US).

TYPE: Tropical America, Shakespeare (BM not seen Tryon photo fragms NY,
US).

Plants terrestrial, at 0 - 1600 m elevation, in forests, from the Cordillera Central,
the Cordillera de Talamanca to Pcia. Chiriqui, S. Isidro del General, the Atlantic
and Pacific lowlands from Costa Rica to the Canal Zone, the mountains of Pcias.
Veraguas and Code, and the Andean slopes of the Depto. Choco. Also from
Guatemala to Honduras, Nicaragua, Colombia to Bolivia, and Brazil.

This species is extremely variable in frond form, with differences in the ultimate
segments and even in the indument, which doubtless accounts for the large
number of nomenclatural synonyms.

354 CYATHEACEAE: 64. CYATHEA

539. Cyathea nigripes van brunnescens (Barn) Lellinger, Amen Fern J. 77:101.
1988.

Trichopteris nigripes var, brunnescens Barr, Rhodora 78:4, f. 5-6. 1976. TYPE: Rio Yurumangui,
Depto. El Valle, Colombia, 5-50 m, Cuatrecasas 16155-C (US; isotype GH).

Plants terrestrial, at 0-300(1400) m elevation, from the Serrania del Darien,
the summit of Cerro Pirre, Alto del Buey, and the Rio S. Juan basin. Also from
Colombia, Eucador, and Peru.

540. Cyathea nigripes (C. Chr.) Domin, Pteridophyta 263. 1929, var. nigripes.

Alsophila melanopus Hook. Syn. Fil. 37. 1865, non Hassk., 1855, nom. illeg. LECTOTYPE: Mt.
Chimborazo, Pcia. Chimborazo, Ecuador, 3000 ft. Spruce 5742 (K not seen), chosen by Barrington
(Contr. Gray Herb. 208:41. 1978).

Alsophila nigripes C. Chr. Ind. Fil. 45. 1905. TYPE: A renaming of A. melanopus Hook., and so
based on the type of that name.

Plants terrestrial, at 0-1500 m elevation, from north of S. Ramon, the
Cordillera Central, the Cordillera de Talamanca to S. Isidro del General, near El
Valle, Cerro Pirre and the Cerro de Garagara (Pcia Darien), the central Choco,
and the basin of the Rio S. Juan. Also from Colombia and Ecuador.

541. Cyathea petiolata (Hook.) Tryon, Contr. Gray Herb. 206:42. 1976.

Hemitelia petiolata Hook. Sp. Fil. 1:31, t. 16. 1844. TYPE: “Isthmus of Panama,” Sinclair (K not
seen).

Hemistegia marginalis K. Presl, Gefassbiindel Farm 47 (postpr. 355). 1847, nom. superfl. TYPE:
A renaming of Hemitelia petiolata Hooker, and so based on the type of that name.

Cyathea panamensis Domin, Pteridophyta 264. 1929, nom. superfl. TYPE: A renaming of

Hemitelia petiolata Hook., and so based on the type of that name.

Hemitelia woronovii Maxon & Morton, Amer. Fern J. 36:91. 1946. TYPE: Penas Blancas, Depto.
Antioquia, Colombia, Woronow & Juzepczuk 4549 (LE not seen; isotype US).

Plants terrestrial, at 0-200(600) m elevation, in forests, from the Canal Zone
through the Choco. Also from the Pacific coastal slopes of Colombia south of the
Choco.

542. Cyathea phalaenolepis (C. Chr.) Domin, Pteridophyta 263. 1929.

Alsophila phalaenolepis C. Chr. Repert. Spec. Nov. Regni Veg. 10:213. 1911. TYPE: Pcia.
Esmeraldas, Ecuador, Aug 1904, Sodiro (P not seen photo 3437).

Plants terrestrial, at 0-300 m elevation, in forests, from the southern half of the
Choco. Also from adjacent Colombia and Ecuador.

543. Cyathea pilosissima (Baker) Domin, Pteridophyta 262. 1929.

Alsophila pilosissima Baker, Syn. Fil. ed. 2. 457. 1874. TYPE: Mount Campana, Tarapoto,
Depto. S. Martin, Peru, Spmce 4322 (K not seen Tryon photo fragms NY, US).

Plants terrestrial, at 0-200(1000) m elevation, from Cerro Pirre and the Alturas
de Roque (Pcia. Darien), near Puerto Obaldfa (Com. S. Bias), Acandf, Loma del
Cuchillo, the Rio Truando, Bahia Solano, and Puerto Utria (all Depto. Choco).
Also from Ecuador and Peru.

544. Cyathea poeppigii (Hook.) Domin, Pteridophyta 263. 1929.

Alsophila poeppigii Hook. Sp. Fil. 1:43. 1844. TYPE: Peru, Poeppig in 1829 (K not seen; isotype
fragm US).

Alsophila elongata Hook. Sp. Fil. 1:43. 1844. TYPE: Colombia, Hartweg 1528 (K not seen;
isotype fragms NY, US).

CYATHEACEAE: 64. CYATHEA 355

Alsophila sprucei Hook. Syn. Fil. 37. 1865, non Cyathea sprucei Hook., 1865. TYPE: Chimborazo,
Pda. Chimborazo, Ecuador, 2500 ft. Spruce 5740 (K not seen; isotype P not seen).

Alsophila impressa Fee, Crypt. Vase. Bresil 1:165, t. 58, f. 1. 1869. TYPE: Brazil, Glaziou 983 (P
not seen).

Alsophila tijucensis F6e, Crypt. Vase. Bresil 1:171, t. 63, f. 1. 1869. LECTOTYPE: Tijuca, Est.
Rio de Janeiro, Brazil, Glaziou 1707 (P not seen fragm NY), chosen by Windisch (Bot. Jahrb. 98:189.
1977).

Alsophila hakeri Sodiro, Anales Univ. Quito 12:134 (repr. 532). 1895, nom. illeg., non Zeiler, 1885.
TYPE: Near Sardinas, Valle de Pallatanga, Pcia. Chimbrazo, Ecuador, 1040 m, Sodiro (Hb. Sodiro
not seen; isotype NY).

Alsophila quitensis C. Chr. Ind. Fil. 47. 1905. TYPE: A renaming of A. hakeri Sodiro, and so
based on the type of that name.

Alsophila coriacea Rosenst. Mem. Soc. Sci. Nat. Neuchatel 5:50, t. 2, f. 1. 1912. TYPE: Between
Valparaiso and Supfa, Depto. Antoquia - Caldas, Colombia, 1500 m. Mayor 86 (S not seen; isotype
fragm P not seen, isotype UC).

Cyathea oreites Domin, Pteridophyta 263. 1929. TYPE: A renaming of Alsophila sprucei Hook.,
and so based on the type of that name.

Cyathea anacampta Alston, J. Wash. Acad. Sci. 48:230. 1958. TYPE: Confluence of the Rio
Ticuanayoy and the Rfo Caqueta, Depto. Putumayo, Colombia, 1100 m, Ewan 16081 (BM not seen;
isotype US).

Plants terrestrial, at 0 - 1300 m elevation, in forests, from the Atlantic coastal
plain of Costa Rica and Panama, the Cordillera Central, the Peninsula de Osa, the
southern Pacific coastal plain of Costa Rica and adjacent Panama, and El Valle.
Also from Venezuela and Colombia to Bolivia and Brazil.

545. Cyathea pseudonanna (G6mez) Lellinger, Proc. Biol. Soc. Wash. 98:376.
1985.

Trichopteris pseudonanna G6mez, Phytologia 50:69. 1981. TYPE: Cerro Tuti, Edo. Veraguas,
Panama, Folsom & Edwards 3370 (MO).

Plants terrestrial, at 900 - 1300 m elevation, known only from the type locality.
This species appears to be most closely related to C. decorat a.

546. Cyathea schiedeana (K. Presl) Domin, Pteridophyta 263. 1929.

Alsophila schiedeana K. Presl, Tent. Pterid. 62. 1836. TYPE: Based ultimately on Between

Huitamalco and Cuapa, Mexico, Schiede (B or HAL not seen).

Alsophila chnoodes Christ, Bull. Herb. Boissier II, 4:958. 1904. TYPE: Costa Rica, Werckle (P not
seen photo 3395; isotypes NY, US).

Alsophila crassifolia Werckle ex Christ, Bull. Herb. Boissier II, 6:184. 1906. TYPE: Luna,

Quebrada Pulanca, Costa Rica, Werckle (P not seen; isotype US).

Plants terrestrial, at (0)600 - 1400 m elevation, in forests, from the Cordillera
Central, the Cordillera de Talamanca and its eastern foothills to Pcia. Chirquf, the
Fila Costeha near S. Vito, the vicinity of Chiriquf Lagoon (Pcia. Bocas del Toro),
Cerro Tute (Pcia. Veraguas), Cerro Pilon (Pcia. Code), and the Rfo Guanche
near the Portobelo Road (Pcia. Colon). Also from Mexico to El Salvador.

547. Cyathea stipularis (Christ) Domin, Pteridophyta 263. 1929.

Alsophila stipularis Christ, Bull. Herb. Boissier II, 4:958. 1904. TYPE: Costa Rica, Werckle (BR
or P not seen; isotypes GH, US).

Cyathea aphlebioides Christ, Bull. Herb. Boissier II, 6:179. 1906. TYPE: Navarro, Pcia. Cartago,
1400 m, Werckle (P not seen; isotype US).

356 CYATHEACEAE: 64. CYATHEA

552 Cy. wendlandii

CYATHEACEAE: 64. CYATHEA 357

Alsophila ichthyolepis Churist, Bull. Herb. Boissier 11, 6:186. 1906, as ‘"ichtyolepis” LECTOTYPE:
La Palma, Pcia. S. Jos6, 1459 m, Tonduz 12527 (P not seen photo 3458; isolectotypes GH, NY, US),
chosen by Lellinger (Proc. Biol. Soc. Wash. 98:366. 1985).

Alsophila ochroleuca Christ, Bull. Soc. Bot. Geneve II, 1:232. 1909. TYPE: La Palma, Pcia. S.
Jose, Werckle 17055 (P not seen; isotype BR not seen Tryon photo).

Plants terrestrial, at 1100-1700(2400) m elevation, in forests, from the
Cordillera Central, the Cordillera de Talamanca to Pcia. Chiriqui, and the Fila
Costeha near S. Vito. Also from Nicaragua.

547a. Cyathea stolzei A. R. Smith ex Lellinger, Amer. Fern J. 77:101. 1988.

Trichopteris pinnata Stolze, Amer. Fern J. 74:103, f. 2. 1984, non Cyathea pinnata Roxb. ex Clarke,
1874. TYPE: Sta. Rita Ridge road 21-26 km from the Transisthmian Hwy., Pcia. Colon, 500-550
m, Knapp 5881 (MO not seen; isotype F).

TYPE: A renaming of Trichopteris pinnata Stolze, and so based on the type of
that name.

Plants terrestrial, at 0-500 m elevation, from west of Pto. Viejo (Pcia.
Heredia), the Sta. Rita ridge road (Pcia. Colon), Cerro Azul (Pcia. Panama),
Cerro Jefe, between the continental divide and the Atlantic ocean opposite Card
(Com. S. Bias), and between El Valle and Bahia Solano (Depto. Choco).

Smith and Grayum (Amer. Fern J. 78:105-108. 1988) reported hybrids between
this species and C. ursina from Cerro Sardinal northeast of Volcan Barba (Smith et
al 1780, CR, MO, UC).

548. Cyathea suprastrigosa (Christ in Pitt.) Maxon, North Amer. FI. 16:83. 1909.

Hemitelia suprastrigosa Christ in Pitt. Prim. FI. Costaric. 3(1):44. 1901. TYPE: “Forets de

I’Achiote,” VoIcm Poas, Pcia. Alajuela, 2000 m, Tonduz 10701 (BR or P not seen; isotypes CR, NY,
P not seen, US).

Cyathea conspicua Christ, Bull. Herb. Boissier II, 6:178. 1906. TYPE: Volc^ Turrialba, Pcia.
Cartago, Werckle in 1905 (P not seen; isotypes US fragm NY).

Plants terrestrial, at 1400-3000 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Cerro Chirripo.

549. Cyathea trichiata (Maxon) Domin, Acta Bot. Bohem. 9:166. 1930.

Alsophila trichiata Maxon, Contr. U. S. Natl. Herb. 24:44, t. 15. 1922. TYPE: Near Cana, Pcia.
Darien, 660 m, R. S. Williams 928 (US; isotype NY? not seen).

Plants terrestrial, at 0-1400(1600) m elevation, in forests, from the Cordillera
Central and the north end of the Cordillera de Talamanca and their adjoining
lowlands, the Fila Costeha, the Peninsula de Osa, and the lowlands of Panama to
the central Choco. Also from Nicaragua, Venezuela, Colombia, and Ecuador.

550. Cyathea ursina (Maxon) Lellinger, Amer. Fern J. 77:101. 1988.

Alsophila ursina Maxon, J. Wash. Acad. Sci. 34:48. 1944. TYPE: Antelope Ridge, Stann Creek
Valley, Stann Creek Distr., Belize, Gentle 3197 (US; isotype MICH not seen).

FIGS. 544-553. Cyathea. FIG. 544. Ultimate segment of Cy. poeppigii, Skutch 17. FIG. 545. Pinna
of Cy. pseudonanna, Folsom & Edwards 3370 (MO). FIG. 546. Pinnule base of Cy. schiedeana, de la
Sota 5182. FIG. 547. Pinnule base of Cy. stipularis, Tonduz 12527. FIG. 548. Ultimate segment of
Cy. suprastrigosa, Pittier 4175. FIG. 549. Pinnule base of Cy. trichiata, R. S. Wilhams 928. FIG. 550.
Median pinna of Cy. ursina, Scamman 7438. FIG. 551. Piimule base of Cy. villosa, Killip & Smith
15309, Colombia. FIG. 552. Basal portion of median pinnule of Cy. wendlandii, Allen 2783. FIG.
553. Median pinna of Cy. williamsii, R. S. Williams 850.

358 CYATHEACEAE: 65. ALSOPHILA

Plants terretrial, at 100-300 m elevation, from Cerro Sardinal, Timbirina, and
Finca La Selva (all Pda. Heredia). Also from Belize and Nicaragua.

For a comment on hybrids with C. stolzei, see that species.

551. CyatheavillosaHumb.&Bonpl.exWilld. Sp. PI. ed. 4, 5:495. 1810.

Chnodphora humboldtii Kaulf. Enum. Fil. 250. 1824, nom. illeg. TYPE: A renaming of C villosa
Humb. & Bonpl. ex Willd., and so based on the type of that name.

Alsophila ri^dula Mart. Icon. PI. Crypt. Bras. 74, t. 51. 1834. TYPE: Est. S. Paulo, Brazil, Martins
(M not seen).

Alsophila vemicosa Mett. ex Kuhn, Linnaea 36:155. 1869. TYPE: Venezuela, Fendler 344 (B not
seen fragm NY not seen; isotypes GH not seen, MO not seen, US).

Alsophila villosa var. dusenii Christ in Dusen, Ark. Bot. 9(15):2. 1910. SYNTYPES: Serrinha, Est.
Parana, Brazil, Dusen 3437 (S not seen; isotype US); and Rio Guavirova, Est. Parana, Brazil, Dusen
8049 (S not seen).

Alsophila cipoensis Sehnem, FI. Ilustr. Catar. 1(CIAT):96, t. 39. 1978. TYPE: Serra do Cipo,
Jaboticatuba, Est. Minas Gerais, Brazil, Hatschbach 29880 (ASSL not seen; isotype UC).

LECTOTYPE: Sta. Cruz, Edo. Sucre, Venezuela, Humboldt 436 (B-Hb. Willd.
20175-2 not seen Tryon photo), inferentially chosen by Barrington (Contr. Gray
Herb. 208:60. 1978).

Plants terrestrial, at 300-800 m elevation, in forests, from Buenos Aires (Pcia.
Puntarenas), and S. Felix to Cerro Flor (Pcia. Chiriqui). Also from Venezuela,
Colombia, and Brazil.

552. Cyathea wendlandii (Mett. ex Kuhn) Domin, Pteridophyta 263. 1929.

Alsophila wendlandii Mett. ex Kuhn, Linnaea 36:158. 1869, as ''wendlandi.” TYPE: San Miguel,
Costa Rica, Wendland 761 (B not seen fragm US).

Alsophila latisecta Christ, Bull. Herb. Boissier II, 6:185. 1906. TYPE: Valley of the Rio Navarro,
Pcia. Cartago, 1400 m, Werckle 16767 (P not seen fragm NY; isotype US).

Plants terrestrial, at 500-1500 m elevation, in forests, from La Marina (Pcia.
Alajuela), west of Sta. Fe, north of El Valle, and near Cerro Jefe.

553. Cyathea williamsii (Maxon) Domin, Acta Bot. Boehm. 9:171. 1930.

Alsophila williamsii Maxon, Contr. U. S. Natl. Herb. 24:46, t. 17. 1922. TYPE: Mountains above
Cana, Pcia. Darien, 1800 m, R, S. Williams 850 (US; isotype NY).

Plants terrestrial, at 800-1800 m elevation, in forests, from Sta. Fe, near Sta.
Rita (Pcia. Colon), Cerro Jefe, the El Llano -Carti road (Com. S. Bias), Cerro
Pirre and above Cana, and the slopes of Alto del Buey. Also from Venezuela.

65. ALSOPHILA R. Br.

Plants terrestrial, arborescent; caudices usually erect (prostrate or decumbent in
some species), woody, often with a thick covering of adventitious roots, especially
at the base of erect rhizomes, scaly; stipes stout, obsolete to long, smooth to spiny,
scaly, the scales concolorous to bicolorous, with usually black marginal spine-like
cells and/or an apical cell, the marginal cells different in shape and orientation
from the central ones; laminae pinnate-pinnatifid to 3-pinnate-pinnatifid,
chartaceous to subcoriaceous; veins simple or more commonly forked; sori lateral
on the veins, usually borne close to the costa, round, exindusiate or indusiate (the
indusia various), sometimes paraphysate; sporangia short-stalked, bearing 16
spores.

Pantropical; over 200 species, with ca. 30 plus hybrids in the New World.

CYATHEACEAE: 65. ALSOPHIU\ 359

CONANT, D. S. 1983. A revision of the genus Alsophila (Cyatheaceae) in the Americas. J. Arnold
Arbor. 64:333 - 382.

GASTONY, G. J. 1973. A revision of the fern genus Nephelea. Contr. Gray Herb. 203:81 - 148.

1. Indusia whitish to tan, thin and delicate, deciduous to persistent, rupturing irregularly..3.

1. Indusia dark brown, thick and fu-m, persistent, rupturing to 2-4 lobes..2.

2(1). Costae not alate between the more distal, sessile pinnules, usually pale brown abaxially;
pinnule segments of median pinnae mostly 3-4 mm wide; indusia cup- to urn-shaped or rarely
spherical with an apical pore.

555. yl. erinacea var. erinacea

2(1). Costae alate between the more distal, sessile pinnules, usually atropurpureous abaxially;
pinnule segments of median pinnae mostly 2-3 mm wide; indusia urn-shaped (rarely cup-shaped or
spherical without an apical pore).

554. A. cuspidata

3(1). Scales on the costae and costules bicolorous, the central band atropurpureous, terminating in
an apical spine-like cell, the margins pale; costae not alate between the more distal, sessile pinnules.

557. A. imrayana var. basilaris

3(1). Scales of the costae and costules concolorous, pale to brownish, bearing 1- several pale to
dark brown, marginal spine-like cells; costae alate between the more distal, sessile pinnules.. 4.

4(3). Laminae uniformly 2-pinnate-pinnatifid; pinnules up to 2.5 cm wide; ala of the costae ca.
0.25 " 0.5 mm wide.

556. A.firma

4(3). Laminae 2-pinnate-pinnatifid to 3-pinnate-pinnatifid; pinnules up to 4.5 cm wide; ala of the
costae ca. 0.4 -0.7 mm wide.

558. A. polystichoides

554. Alsophila cuspidata (Kunze) Conant, J. Arnold Arbor. 64:371. 1983.

Cyathea cuspidata Kunze, Linnaea 9:101. 1834. TYPE: “Prov. Maynas,” Peru, Poeppig Diar. 2286
(LZ destroyed; isotypes B not seen, NY not seen, P not seen, fragm US).

Cyathea oyapoka Jenm. Ferns Brit. W. Ind. Guiana 58. 1898. TYPE: Upper Oyapok river,
French Guiana, June 1883, Leprieur (NY; isotypes F, GH, NY, P none seen, US).

Cyathea punctifera Christ in Pitt. Prim. FI. Costaric. 3(1):40. 1901. TYPE: Tms, Pcia. Cartago, 650
m, Tonduz 11307 (BR not seen; isotypes GH, fragm NY, P not seen, US).

Cyathea hassleriana Christ, Bull. Herb. Boiss. II, 7:926. 1907. TYPE: Near Caacupe, La

Cordillera, Paraguay, Hassler 120 (P? not seen; isotype S not seen).

Plants terrestrial, at 0-1100(1800) m elevation, in forests, in the foothills of the
Cordillera Central and the Cordillera de Talamanca to Pcia. Chiriquf, Cerro Pilon
(Pcia. Code), the Canal Zone, El Llano -Carti road (Pcia. Panama), between
Paya and Boca de Paya (Pcia. Darien), and the southern Choco. Also from
Nicaragua, Colombia to Bolivia, French Guiana, Brazil, and Paraguay.

555. Alsophila erinacea (Karst.) Conant, J. Arnold Arbor. 64:371. 1983, var.

erinacea.

Cyathea erinacea Karst. Linnaea 28:453. 1857. TYPE: Merida, Edo. Merida, Venezuela, 2000 m,
Karsten (W or LE not seen; isotype B not seen).

Cyathea aureonitens Christ, Bull. Herb. Boiss. II, 4:948. 1904. LECTOTYPE: Rio Navarrito, Pcia.
Cartago, 1400 m, Pittier 2413 (BR not seen; isolectotypes NY not seen, B not seen, P not seen),
chosen by Gastony (Contr. Gray Herb. 203:121. 1973).

Cyathea cuspidata var. ri^da Rosenst. Meded. Rijks-Herb. 19:6. 1913. TYPE: Espiritu Santo,
Depto. Cochabamba, Bohvia, 1600 m, Herzog 2237 (L not seen fragm US; isotypes M not seen, S not
seen, UC not seen, US).

360 CYATHEACEAE: 65. ALSOPHILA

CYATHEACEAE: 65. ALSOPHILA 361

Plants terrestrial, at (100)800-2000 m elevation, in forests, from the Cordillera
de Tilaran, the Cordillera Central, and the Cordillera de Talamanca to Pcia.
Chiriqui, La Mesa (Pcia. Code), and the Rio S. Juan between Dipurdu and S.
Miguel (Depto. Choco). Also from Venezuela and Colombia to Bolivia.

556. Alsophila firma (Baker) Conant, J. Arnold Arbor. 64:372. 1983.

Cyathea mexicana Schlechtend. & Cham. Linnaea 5:616. 1830, non Alsophila mexicana Mart.,
1834. TYPE: Jalapa, Edo. Veracruz, Mexico, Schiede (B not seen; isotypes BM, K neither seen).

Cyathea articulata Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 8]: 111. 1857, non
Alsophila articulata J. Smith ex T. Moore & Houlst., 1852. TYPE: Villa Alta and Talea, Edo.
Oaxaca, Mexico, Galeotti 6531 (BR not seen; isotypes B not seen fragm NY, P not seen, US).

Hemitelia firma Baker, J. Bot. Brit. For. 15:161. 1877, non Cyathea firma Mett. ex Kuhn, 1869.
TYPE: Andes of Quito, Pcia. Pichincha, Ecuador, Aug 1875, Sodiro (K not seen; isotypes P not seen,
US).

Cyathea patellaris Christ, Annuaire Conserv. Jard. Bot. Geneve 4:207. 1900. TYPE: Rio de Las
Vueltas, Tucurrique, Pcia. Cartago, 631 m, Tonduz 12783 (G not seen photos 16879-16882; isotypes
F not seen, GH, M not seen, NY, P not seen, US).

Alsophila costalis Christ, Bull. Herb. Boissier II, 4:951. 1904. TYPE: Near Turrialbaf?], Pcia.
Cartago, Werckle & Brune (P not seen; isotype fragm NY).

Alsophila furcata Christ, Bull. Herb. Boissier II, 4:957. 1904. TYPE: Azul Quelle, Turrialba, Pcia.
Cartago, Werckle & Bmne (P not seen; probably isotype US).

Alsophila tenerifrons Christ, Bull. Herb. Boissier II, 4:959. 1904. TYPE: Jimenez, Pcia. Limon,
200 m, Alfaro 16509 (P not seen USNH photo 3150 fragm NY; isotypes GH not seen, P not seen,
fragm US).

Cyathea trejoi Christ, Bull. Herb. Boissier II, 5:733. 1905. TYPE: S. Pablo, Edo. Chiapas, Mexico,
1500 m, Muench in 1903 (P not seen fragm NY), synonymized by Tryon (Contr. Gray Herb. 206:92.
1976).

Cyathea arida Christ, Bull. Herb. Boissier II, 6:180. 1906. SYNTYPES: Navarro, La Luna, Pcia.
Cartago, 1400 m, Werckle 16757 (P not seen; isosyntype US) and 16780 (P? not seen); and Alto de
Mano de Tigre, Diquis, Pcia. Puntarenas, 700 m, Pittier 12097 (P? not seen; isosyntype US).

Cyathea gemmifera Christ in Jimenez, Bol. Fomento (San Jose) 3:664, f. 1-6. 1913. TYPE:
Cerros del Tremedal near S. Ram6n, Pcia. Heredia, 1300-1400 m, Tonduz (Hb. Jimenez 913) 17563
(CR not seen; isotypes A, F not seen, P not seen, UC, US).

Cyathea firmula Domin, Acta Bot. Boehm. 9:115. 1930. TYPE: A renaming of Hemitelia fimia
Baker, and so based on the type of that name.

Plants terrestrial, at 0-1700 m elevation, in forests, from the slopes of the
Cordilleras and adjacent ridges and wet lowlands throughout the Flora area. Also
from Mexico to Nicaragua and Ecuador.

557. Alsophila imrayana var. basilaris (Christ) Lellinger, Amer. Fern J. 75:31.
1985.

Cyathea basilaris Christ, Bull. Herb. Boissier II, 4:949. 1904. LECTOTYPE: Costa Rica, Werckle
(P not seen; isotypes NY, US), chosen by Gastony (Contr. Gray Herb. 203:127. 1973).

FIGS. 554-562. Alsophila, Sphaeropteris, Dicksonia, 2md Culcita. FIG. 554. Pinnule base of A.
cuspidata, Idrobo & Schultes 922, Colombia. FIG. 555. Piimule base of A. erinacea var. erinacea,
Werckle 584. FIG. 556. Pinnule base of A. firma, Killip 2692. FIG. 557. Pinnule base of A. imrayana
VcU". basilaris, Grant 10879, Colombia. FIG. 558. Pinnule base of A. polystichoides, Tonduz 12528.
FIG. 559. Ultimate segment of S. bmnei, Maxon 332. FIG. 560. Base of median pinnule of median
pinna of D. gigantea, Standley & Torres 51059. FIG. 561. Base of median pinnule of median pinna of
D. karsteniana, Scamman & Holdridge 7892. FIG. 562. Pinnule of C coniifolia. Burger & Stolze
5231.

362 CYATHEACEAE: 66. SPHAEROPTERIS

Cyathea reticulata Werckle ex Christ, Bull. Herb. Boissier 11, 5:251. 1905. TYPE: Volc^ Irazu,
Pcia. Heredia, 1800 m, Werckle 6 (P not seen; isotypes NY, P not seen, US).

Plants terrestrial, at (400)1600-2000 m elevation, in forests, from the
Cordillera Central and near Boquete. Also from Venezuela, Colombia, and
Ecuador.

558. Alsophila polystichoides Christ in Dur. & Pitt. Bull. Soc. Roy. Bot. Belgique
35,M^m. 177. 1896.

Alsophila mucronata Christ in Dur. & Pitt. Bull. Soc. Roy. Bot. Belgique 35, Mem. 178. 1896.
TYPE: Confluence of the Rio Puerto Viejo and the Rio Sarapiqui, Pcia. Heredia, Pittier 7484 (BR
not seen photo 4924; isotype US).

Cyathea hastulata Christ, Bull. Herb. Boissier II, 4:945. 1904. TYPE: Arias Agua Punta, Costa
Rica, Werckle (P not seen; isotype NY).

Cyathea werckleana Christ, BuU. Herb. Boissier II, 6:181. 1906. LECTOTYPE: La Palma, Pcia. S.
Jose, 1459 m, Tonduz 12677 (P not seen; isolectotypes B not seen, US), chosen by Gastony (Contr.
Gray Herb. 203:144. 1973).

Cyathea hemiotis Christ, BuU. Herb. Boissier II, 6:182. 1906. TYPE: Navarro, La Luna, Pcia.
Cartago, Werckle (P? not seen; authentic material BM not seen, NY, P not seen, UC, US).

Cyathea caduca Christ, BuU. Herb. Boissier II, 7:271. 1907. TYPE: Jardin de Carmiol, San Jose,
Pcia. S. Jose, Werckle in 1906 (P not seen; isotype fragm US).

TYPE: Above Aragon, Pcia. Cartago, 600 m, Pittier 9017 (BR not seen photo
4934 fragm NY; isotype US).

Plants terrestrial, at 700-2000 m elevation, in forests, from the Cordillera
Central and the Cordillera de Talamanca to Pcia. Chiriquf.

66. SPHAEROPTERIS Bernh.

Plants terrestrial, arborescent; caudices erect, woody, with a thick covering of
adventitious roots, especially at the base, scaly, the scales concolorous but with
dark, marginal, spine-like cells; stipes stout, long, smooth or nearly so, not spiny,
scaly, the scales concolorous, whitish or tan, with all cells alike in orientation,
thickness, and color, except for the marginal spine-like cells, the stipes also
bearing a scurf of minute, branched, darkened hairs; laminae 2(3)-pinnate-
pinnatifid to 2-pinnate-pinnatisect, the segments entire, coriaceous; veins rarely
simple, usually 1-forked; sori lateral at the fork of a vein, indusiate, the indusium
cup-shaped or globose, thin to firm; paraphyses long, clavate; sporangia short-
stalked.

Paleotropical (many species), with 6 species in the Neotropics.

TRYON, R. M., Jr. 1971. The American tree ferns alUed to Sphaeropteris horrida. Rhodora
73:1-19.

559. Sphaeropteris brunei (Christ) Tryon, Contr. Gray Herb. 200:20. 1970.

Cyathea brunei Christ, BuU. Herb. Boissier II, 4:947. 1904. TYPE: Costa Rica, Prune & Werckle
(P not seen; isotype GH, presumable isotype US).

Cyathea caesia Christ, BuU. Herb. Boissier 11,7:272. 1907. LECTOTYPE: La Palma, Pcia. S. Jose,
1500 m, 24 Nov 1905, Werckle 17008 (P not seen; isotype fragm US), chosen by Tryon (Rhodora
73:13. 1971).

Plants terrestrial, at 800-2000 m elevation, at forest margins and on cleared
slopes, from the Cordillera Central, the Fila Las Cruces near S. Vito, and Pcia.
Chiriquf. Also from the Depto. Magdalena, Colombia.

DICKSONIACEAE: 67. DICKSONIA 363

DICKSONIACEAE

Rhizomes prostrate, ascending, or erect and stout, with adventitious roots at the
base and densely covered with long, soft, golden hairs at the apex; fronds large;
stipes persistent, short to long, smooth, hairy at the base, the hairs long, straight,
pale to dark brown; laminae 2-pinnate-pinnatifid to 5-pinnate, subcoriaceous,
glabrous, glaucous, or sparsely hairy abaxially, hairy on the adaxial surface of the
major axes; veins free, simple or forked 1- several times; sori submarginal or
marginal at the end of a vein; indusia double, appearing globose or nearly so, the
outer (false) indusium poorly to well developed from the margin of a lobe, the
inner (true) indusium well developed, extrorse; sporangia long-stalked, the stalks
thick, the capsules angular-globose; paraphyses catenate, at least slightly
exceeding the sporangia; spores trilete.

1. Laminae up to 3-pimiate; major 2ixes hairy on the adaxial surface; sori globose.

67. Dicksonia

1, Laminae at least 4-pinnate; major axes glabrous on the adaxial surface; sori subglobose,
elongate laterally.

68. Culcita

67. DICKSONIA L’Her.

Plants terrestrial, arborescent; caudices erect, woody, with a thick covering of
adventitious roots, especially at the base, hairy at the apex, the hairs soft, golden;
stipes persistent, short to long, smooth, hairy, the hairs long, straight, pale to dark
brown; laminae mostly 2-pinnate-pinnatifid or 3-pinnate, the segments entire (in
sterile fronds) to lobed, coriaceous; veins mostly 1-forked; sori submarginal at the
end of a vein, indusiate, the outer indusium prolonged from the margin of the
lobe, the inner indusium tan, scarious, ciliate; paraphyses somewhat longer than
the sporangia, catenate, with or without a glandular terminal cell; sporangia short-
stalked.

Mostly Malaysia and the Pacific Islands; ca. 30 species.

MAXON, W. R, 1913. The North American ferns of the genus Dicksonia. Contr. U. S. Natl. Herb.

17:153-156.

1. Costae smooth to slightly rough and mostly glossy on the abaxial surface; fertile segments
(2.5)3 -3.5(4.5) mm wide; costae and costules glabrous to sparsely covered with lax, appressed,
deciduous, somewhat catenate hairs and sometimes also with a few stiff, spreading, subdeciduous or
persistent, usuaOy terete, bristle-like hairs; soral paraphyses catenate without a glandular terminal

cell.

560. D. gigantea

1. Costae slightly to decidedly rough and scarcely or not glossy on the abaxial surface; fertile
segments 2-3 mm wide; costae and costules usually densely covered with lax, appressed, deciduous,
somewhat catenate hairs, but bristle-like hairs absent; soral paraphyses catenate, often with a dark
red, non-swoUen, glandular terminal cell.

561. D. karsteniana

560. Dicksonia gigantea Karsten, FI. Columb. 2:177. 1869.

Dicksonia lobulata Christ, Bull. Herb. Boissier II, 6:187. 1906. TYPE: Cerros de Velirla, Copey
de Dota, Pcia. S. Jose, 2600 - 2700 m, Tonduz 11789 (P not seen; isotype US).

Dicksonia navarrensis Christ, Bull. Herb. Boissier II, 6:188. 1906. TYPE: Valley of the Rio
Navarro, Pcia. Cartago, Werckle in 1905 (P not seen; isotype BM not seen photo 7918).

Dicksonia ghiesbreghtii Maxon, Contr. U. S. Natl. Herb. 17:155. 1913. TYPE: Edo. Chiapas,
Mexico, Ghiesbreght 353 (US fragm NY).

364 DICKSONIACEAE: 68. CULCITA

TYPE: Mt. Guadalupe, Andes of Bogota, Depto. Cundinamarca, Colombia,
2600 m, Karsten (LE or W not seen).

Plants terrestrial, at 1200-2000(2700) m elevation, in forests and shaded
ravines, from the Cordillera Central, Cerro Tablazo, Cerro Carpintera, the upper
Rio Reventazon valley, the Cordillera de Talamanca to Pcia. Chiriqui, the Fila
Costeha near S. Vito de Java, and Cerro Pirre and the Alturas de Nique (Pcia.
Darien). Also from Mexico to El Salvador, Colombia, and Ecuador.

561. Dicksonia karsteniana (Klotzsch) T. Moore, Ind. Fil. 190, 313. 1860.

Balantium karstenianum Klotzsch, Linnaea 20:444. 1847. TYPE: Colonia Tovar, Edo. Aragua,
Venezuela, Karsten II 9 (B not seen; isotype HBG not seen photos 5372, 5374).

Dicksonia sellowiana var. arachneosa Sodiro, Anales Univ. Quito 7(49) :83 (repr. 44). 1892.

TYPE: Volcan Corazon, Pcia. Pichincha, Ecuador, 3300 m, Sodiro (Hb. Sodiro not seen).

Plants terrestrial, at 1900-2300 m elevation, in forests and ravines, from the
Cordillera Central and the Cordillera de Talamanca. Also from Venezuela and
Colombia to Bolivia.

The American members of Dicksonia are in need of additional study; new,
perhaps chemical, data would be especially welcome. This species and D. gigantea
are not always easily separated; the largely southeastern Brazilian D. sellowiana
Hook, apparently differs in having soral paraphyses shorter than the sporangia.

68. CULCITA K. Presl

Plants terrestrial, not arborescent; rhizomes prostrate or ascending, rarely erect
at the apex, woody, with adventitious roots, especially at the base, hairy at the
apex, the hairs soft, twisted, golden; stipes long, smooth, hairy, the hairs long,
straight, pale to rarely dark brown; laminae 4-pinnate-pinnatifid or 5-pinnate, the
segments subcoriaceous, lobed, the sterile lobes acute at the apex; sori marginal at
the end of a vein, indusiate, the indusia wider than long, the outer indusium
scarcely prolonged from the margin of the lobe, the inner indusium opposed, tan,
firm, eciliate; paraphyses much longer than the sporangia, catenate, without a
glandular terminal cell; sporangia short-stalked.

One species of the Atlantic islands and southwestern Europe and one in the
neotropics. Other species formerly placed here belong in Calochlaena.

MAXON, W.R. 1922. The genus Culcita. J. Wash. Acad. Sci. 12:454 - 460.

TURNER, M. D. and R. A. WHITE. 1988. Calochlaena, a new genus of Dicksonioid ferns. Amer.
Fern J. 78:86 - 95.

562. Culcita coniifolia (Hook.) Maxon, Annual Rep. Board Regents Smithsonian
Inst. 1911:488. 1912.

Dicksonia coniifolia Hook. Sp. Fil. 1:70, t. 24A. 1844. TYPE: Merida, Edo. Merida, Venezuela,
Linden S38 (K not seen; isotype BR not seen photo 4847).

Dicksonia martiana Klotzsch ex Hook. Sp. Fil. 1:70, t.24B. 1844. TYPE: Southern Brazil, Sellow
(B or K not seen).

Culcita schlimensis Fee, Icon. Esp. Nouv. [Mem. Foug. 10]: 47, t. 36, f. 3. 1865. TYPE: Ocana,
Depto. Norte de Santander, Colombia, Schlim 322 (RB not seen).

Plants terrestrial, at 2000-3300 m elevation, in forests, thickets, and clearings,
from the Cordillera Central and the Cordillera de Talamanca to Pcia. Chiriqui.
Also from Cuba, Jamaica, Hispaniola, Mexico, El Salvador, Venezuela, Colombia
to Peru, and Brazil.

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