Pteridophytas de Peru 1 PDF
Pteridophytas de Peru 1 PDF
Pteridophytas de Peru 1 PDF
ILLINOIS LIBRARY
AT URBANA-CHAMPAIGN
NATURAL HIST. SURVEY
FIELD
Botany
NEW SERIES NO. 20
UBMW
PTERIDOPHYTA OF PERU
Parti
1. Ophioglossaceae-12. Cyatheaceae
Rolla M. Tryon
Robert G. Stolzc
'
PTERIDOPHYTA OF PERU
Parti
1. Ophioglossaceae-12. Cyatheaceae
Rolla M. Tryon
Gray Herbarium
Harvard University
22 Divinity Avenue
Cambridge, Massachusetts 02138
Robert G. Stolze
Collection Manager, Fern Herbarium
Department of Botany
Field Museum of Natural History
Roosevelt Road at Lake Shore Drive
Chicago, Illinois 60605-2496
INTRODUCTION 1 1 .
Botrychium virginianum 7
ACKNOWLEDGMENTS 2 2. Ophioglossum: O. palmatum; O. crotal-
KEY TO FAMILIES OF PTERIDOPHYTA IN ophoroides; O. reticulatum 10
PERU 3 3. Marattia laevis 14
1 . OPHIOGLOSSACEAE 5 4. Danaea: D. moritziana; D. nodosa .... 16
Ophioglossum 8 O. cinnamomea 22
2. MARATTIACEAE 13 6. Anemia: A. pastinacaria; A. phylli-
Marattia 13 tidis 26
Danaea 15 7. Lygodium: L. venustum; L. volubile ... 31
3. OSMUNDACEAE 20 8. Schizaea elegans 35
Osmunda 21 9. Gleichenia bifida 40
4. SCHIZAEACEAE 23 10. Dicranopteris: D. pectinata; D. flex-
Anemia 24 uosa 48
Lygodium 30 11. Hymenophyllum: H. fucoides var. fu-
Schizaea 33 coides\ H. polyanthos; H. crispum 51
5. GLEICHENIACEAE 37 12. Trichomanes: T. radicans; T. pinnatum;
Gleichenia 37 T. hymenoides 77
Dicranopteris 46 13. Loxsomopsis pearcei 100
6. HYMENOPHYLLACEAE 49 14. Plagiogyria semicordata 102
Hymenophyllum 50 15. Culcita coniifolia 104
Trichomanes 76 16. Dicksonia sellowiana 106
7. LOXOMATACEAE 98 17. Lophosoria quadripinnata var. quadri-
Loxsomopsis 99 pinnata 108
8. PLAGIOGYRIACEAE 99 18. Metaxya rostrata 110
Plagiogyria 101 19. Sphaeropteris: S. elongata; S. quindiu-
9. DlCKSONIACEAE 101 ensis 113
Culcita 103 20. Alsophila engelii 117
Dicksonia 105 21. Nephelea cuspidata 119
10. LOPHOSORIACEAE 107 22. Trichipteris pubescens 121
Lophosoria 107 23. Cyathea caracasana var. boliviensis ... 134
11. METAXYACEAE 109 24. Cnemidaria speciosa 137
Metaxya 109
1 2. CYATHEACEAE Ill
Sphaeropteris 112
Alsophila 116
Nephelea 118
Trichipteris 1 20
Cyathea 129
Cnemidaria 1 36
111
PTERIDOPHYTA OF PERU
Parti
1. Ophioglossaceae-12. Cyatheaceae
Introduction
The pteridophytes form a large and conspicuous their knowledge of particular genera, but several
element of the Peruvian flora, including 96 genera treatments will be contributed by specialists cur-
and about 1 ,000 species. Peru, which encompasses rently involved in monographic studies. The re-
one of the world's richest biotas, occupies a central sults will be published in five parts, divided as
position in the Andes, and accordingly a knowl- equally as practicable, beginning with the present
edge of its flora is basic to understanding the plant volume on the Ophioglossaceae through Cyathea-
life of the Andes as a whole. The Andes form a ceae.
largely tropical mountain chain with an essentially Studies are based primarily on the collections
north and south orientation, which is of special Museum, Harvard University Herbaria,
at Field
significance to its biogeography and the underlying and United States National Herbarium (Smith-
processes of speciation. sonian Institution), but specimens at many other
A portion of the Ferns of Peru was published United States and European institutions have been
by Rolla M. Tryon in 1964 (Contr. Gray Herb., examined. The extensive collections made under
encouragement and active sup-
vol. 194), with the the current Flora of Peru project, a joint under-
port of Theodore K. Just of Field Museum of Nat- taking of Field Museum and Missouri Botanical
ural History. A commitment was made at that Garden, are fully utilized. A close collaboration
time to complete the project after undertaking a has also been established with Universidad Na-
major work on the genera of pteridophytes in cional de Trujillo, Trujillo, Peru, and Museo de
America. The present work is a cooperative proj- Historia Natural "Javier Prado" de Universidad
ect of Harvard University Herbaria and Field Mu- Nacional Mayor de San Marcos, Lima, Peru, in-
seum of Natural History to present an account of cluding loans of their specimens. The type of each
all of the Pteridophyta of Peru. This is particularly name has been determined when possible, and an
important in relation to the Flora of Peru, initiated effort has been made to see the holotype, or at
at Field Museum over 50 years ago, and thus far least type photographs, or authentic material.
encompassing some 8,500 published pages. It is Original drawings illustrate the diagnostic features
the only comprehensive treatment of Pacific coast- of each genus and, where possible, some of the
al, Andean, and Amazonian plants, and is closer number of plates published
species. In addition, a
to completion than any flora of the larger South Botany, Ferns and fern allies of Gua-
in Fieldiana:
American countries. temala, have been used for species occurring in
The taxonomy is based not only on Peruvian Peru. Voucher specimens cited in the legends are
materials, but includes an assessment of the An- from Peru unless otherwise indicated.
dean species of a genus and, where appropriate, Prior to 1 944 the Department of Pasco was a
of all tropical American species. Most of the work part of Junin, and until recently Ucayali was a
will be accomplished by the authors, according to part of Loreto. An attempt has been made to ac-
specimens the labels often contain only sketchy Field Museum scientific illustrators, Zorica Da-
data,making it impossible to determine in which bich and Clara L. Richardson, and by volunteer
part of Loreto or Junin a plant was collected. The illustrators, Julia A. Liesse, Rosemarie Seitz, and
map of Peru at the end of the text shows the de- Lisa M. Thorns. To these fine artists we wish to
partments and indicates the sequence of the col- express our sincere appreciation, not only for their
lection citations. painstaking work, but also for their patience and
In the present work, Part I, the treatments of understanding. Special thanks are extended to sev-
the Ophioglossaceae through the Plagiogyriaceae eral persons who have been particularly helpful to
have been prepared by Robert G. Stolze, and those by providing special aid with the ex-
us, either
of the Dicksoniaceae through the Cyatheaceae, by amination of specimens at their institutions, or
Rolla M. Tryon. However, each author has re- through stimulating discussions or correspon-
viewed and edited the manuscript of the other, to dence concerning problems with various genera:
the extent that the treatments are a joint effort. Dr. David Lellinger (US), Dr. Alan R. Smith (UC),
New names published here are indicated by Dr. Henk van der Werff (MO), Dr. W. H. Wagner
boldface in the Index to Names. Collections have (MICH), and Dr. Paulo Windisch (Dept. Botanico,
been cited from each department of Peru from Rio Preto, Brazil). We also appreciate comments
which material has been seen, and further collec- on the manuscript by several reviewers.
tions are sometimes cited to include other her- We extend our thanks to the officers of the fol-
baria. In general, all collections seen are cited for lowing institutions for granting loans of their ma-
rare species, and a selection is cited for common terial or allowing us to examine specimens in their
species. Type collections, mentioned in the syn- herbaria: Botanischer Garten und Botanisches
onymy, are not repeated in the specimen citations, Museum, Berlin-Dahlem, Berlin (B); British Mu-
although they are included in the Peruvian range seum (Natural History), London (BM); Jardin Bo-
and ecology. tanique National de Belgique, Meise (BR); Royal
The nomenclature of the genera and species is Botanic Garden, Edinburgh (E); Field Museum of
not intended to be complete. Synonyms are listed Natural History, Chicago (F); Harvard University,
when they are considered useful and when the type Cambridge, Mass. most Gray Herbarium (GH),
of the name of a species or infraspecific taxon is some Arnold Arboretum (A); Institut fur Allge-
from Peru. meine Botanik und Botanischer Garten der Uni-
Appended to some of the generic treatments are versitat, Hamburg (HBG); Herbarium Truxil-
portions of text labeled Comments. Herein are in- lense,Universidad Nacional de Trujillo, Trujillo,
cluded species to be expected in Peru, names based Peru (HUT); Royal Botanic Gardens, Kew, En-
on Peruvian material but of uncertain application, gland (K); Rijksherbarium, Leiden, The Nether-
excluded species (erroneous reports that may have lands (L); Botanische Staatssammlung, Munich
special significance), and cultivated species that (M); University of Michigan, Ann Arbor (MICH);
are possibly adventive. Missouri Botanical Garden, St. Louis (MO); New
Abbreviations of periodical publications gen- York Botanical Garden, New York (NY); Museum
system of Botanico-Periodicum-
erally follow the National d'Histoire Naturelle, Paris (P); Univer-
Huntianum (1968), and abbreviations of authors' sityof California, Berkeley (UC); United States
names and of books generally follow TL-2 Taxo- National Herbarium, Smithsonian Institution,
nomic Literature by Stafleu and Cowan (1976 et Washington, D.C. (US); Museo de Historia Nat-
seq.). ural "Javier Prado" de Universidad Nacional
FIELDIANA: BOTANY
Key to Families of PTERIDOPHYTA in Peru
The following key is restricted to the families that occur in South America and to the genera that are
known from Peru or are likely to occur there. Several families are keyed out more than once in order
to simplify the headings and to provide for more accurate identification.
a. Sporangia separate d
d. Sporangia borne on the inner side of a peltate sporangiophore in an apical strobilus; leaves much
reduced, forming a sheath at the apex of each elongate, ridged interode Goint) of the stem
25. Equisetaceae
d. Sporangia single in or near the axil of a leaf, or on 1 lobe of a 2-lobed leaf, or several to many
sporangia borne on a leaf: at the margin, on the abaxial surface, on a specialized portion, or on
a specialized leaf e
e. A singlesporangium borne in or near the axil of a leaf f
f. Plants homosporous; leaves lacking a ligule 26. Lycopodiaceae
f. Plants heterosporous, with megasporangia and microsporangia; leaves with a ligule g
g. Leaves less than 1 cm long, borne along an elongate stem, fertile leaves in an apical
strobilus 27. Selaginellaceae
g. Leaves 2 cm long or usually longer, clustered at the apex of a compact to slightly elongate
stem; all leaves usually fertile 28. Isoetaceae
e. Several to many sporangia borne on a leaf, or a single sporangium borne on 1 lobe of a 2-
lobed leaf h
h. Plants heterosporous; the leaf bearing megasporangia and/or microsporangia, these enclosed
in small, specialized structures i
i. Plant with stem rooted in wet soil or underwater; leaves filiform, or with 4 leaflets at
the apex of the petiole 22. Marsileaceae
i. Plant floating on water; the floating leaves entire, oblong to suborbicular, or unequally
2-lobed with 1 lobe submerged 23. Salviniaceae
h. Plants homosporous, the isomorphic sporangia exposed at the margin or on the abaxial
surface of a leaf, or on a specialized portion of a leaf, or on a specialized leaf, sometimes
enclosed prior to maturity within an indusium j
j. Sporangia sessile or subsessile, or with a stalk of 4 or more rows of cells, annulus absent,
or if present then apical, lateral, or oblique, not interrupted by the stalk k
k. Annulus lacking, or sporangia with a poorly differentiated annulus 1
j. Sporangia with a 1- to 3 -rowed stalk; the annulus vertical or nearly so, interrupted by
the stalk u
u. Petiole articulate at or very near the stem 21. Polypodiaceae
u. Petiole continuous with the stem, or articulate well above the base of the petiole v . . .
ff. Stem with trichomes, or with scales and an abaxial indusium present
15. Dennstaedtiaceae
.
FIELDIANA: BOTANY
ff. Stem with scales and an abaxial indusium absent gg
gg. Spores trilete 13. Pteridaceae
gg. Spores monolete 15. Dennstaedtiaceae
bb. Spores monolete and the sori or sporangia abaxial hh
hh. Sori elongate, adjacent to and parallel to the segment axis
20. Blechnaceae
hh. Sori roundish, or elongate, and most or all of them neither adjacent to
nor parallel to the segment axis, or sporangia acrostichoid ii
jj. Spores with chlorophyll (green); lamina or main veins more or less
dichotomous (furcate) 21. Polypodiaceae
jj .
Spores lacking chlorophyll (not green) and lamina pinnately veined
or branched kk
kk. Petiole, at least basally, with 2 vascular bundles 11
17. Dryopteridaceae
ii. Pinnae articulate to the rachis mm
mm. Sporangia acrostichoid, or in indusiate sori and the lamina 2-
pinnate, or 1 -pinnate and the pinnae with a large, basal, bas-
iscopic auricle 17. Dryopteridaceae
mm. Sporangia in indusiate sori, lamina 1 -pinnate, the pinnae cor-
date, or the basal, basiscopic side less developed
. 19. Davalliaceae
a. Sterile lamina ternately decompound; veins free; sporangia borne in a panicle, separate from each
other and not immersed in the fertile segment I. Botrychium
a. Sterile lamina entire or palmately or digitately lobed; veins anastomosing; sporangia joined laterally
in a synangial spike II. Ophioglossum
BUTTERS, F. K. 1917.
Botrychium virginianum
The genus contains about 25 species, with dis- and its American Rhodora, 19: 207.
varieties.
tribution in temperate to tropical regions in both MILDE, J. 1869. Botrychiorum monographia.
hemispheres, but is poorly represented in tropical Verh. Zool. Bot. Ges. Wien, 19: 55-190.
a. Fertile leaf with the sterile lamina long-stalked; leaf bud completely enclosed by the sheath of the
current leaf 1 . B. schafTneri
a. Fertile leaf with the sterile lamina sessile or nearly so; leaf bud partly exposed within the sheath of
the current leaf 2. B. virginianum
1. Botrychium schafTneri Underw., Bull. Torrey shaped sinuses, the margins entire to crenate, very
Bot. Club 30: 51. 1903. TYPE: Mexico, San rarely serrate.
Luis Potosi, Schaffner 9 (holotype, NY; iso-
type, K). Terrestrial, in high scrub forests and open mead-
ows, 2350-3600 m, Amazonas, Cuzco, Madre de
Dios(?).
Mexico; Honduras south to Argentina.
Fertile plants 12-30 cm tall (sometimes to 50 This is often difficult to distinguish from B. un-
cm outside Peru), the common stalk to 4 cm long, derwoodianum (Guatemala to Venezuela), which
the leaf bud completely enclosed by the sheath of is usually a much larger fern with stalks of sterile
the current leaf. Sterile lamina of fertile leaf long- laminae up to 28 cm long. However, Mexican
stalked (3-8 cm), fleshy, ternate, to 3-pinnate-pin- plants of B. schqffneri seem to rival the latter in
natifid,3-10 cm long, 6-10 cm broad. Ultimate size, and the more reliable diagnostic features ap-
segments 0.3-0.7 cm long, mostly obtuse to sub- pear to occur in the ultimate segments. Those of
acute, separated by usually narrow to broadly U- B. underwoodianum are 0.7-1 .4 cm long with mar-
FIELDIANA: BOTANY
FIG.1. Botrychium virginianum: a, lamina with fertile branch; b, stem and part of petiole; c, part of fertile pinna,
(a from Seller 865, El Salvador, GH; b-c from Lyonnet 896, Mexico, GH.)
more recent and numerous collections throughout WAGNER, W. H., ET AL. 1984. Ophioglossum el-
the range of the species indicates that these are lipticum in Louisiana and the taxonomy of O.
variable characters and not wholly correlated with nudicaule. Castanea, 49: 99-1 10.
FIELDIANA: BOTANY
Key to Species of Ophioglossum
a. Sterile lamina deeply and irregularly palmately lobed; fertile spikes several; plants epiphytic
1 . O. palmatum
a. Sterile lamina entire; fertile spike solitary; plants terrestrial b
b. Stem globose 5. O. crotalophoroides
c. Sterile lamina sessile or nearly so, the common stalk and stalks of sterile and fertile segments
d. Primary areoles not enclosing secondary ones or free veinlets; base of sterile lamina narrowly
cuneate to attenuate e
e. Sterile lamina elliptic or oblong, 1.5-3 times as long as broad, cuneate at base; plants of
1 .
Ophioglossum palmatum Sp. pi. 1063. 1 753.
1 ... in deep ravine, Allard21580 (us). Pasco: Prov. Oxapam-
TYPE: based on illustration of plant from pa, 1800 m, van der \Verff8352 (MO). Cuzco: Prov. La
Convention, Tunquimayo, Idma, Vargas 10681 (MICH,
Haiti, Plumier, Traite foug. Amer., /. 163.
MO).
1705. Figure 2a.
5mm
3cm
10 FIELDIANA: BOTANY
From high, rocky meadows to the open forests and sedges, 2800-4000 m, Cajamarca, Lima, Cuz-
of middle elevations, and the grassy soil of coastal co.
lomas, 400-3800 m, Amazonas south to Cuzco. Colombia; Peru; Brazil.
Widely distributed in tropical America; Old In spite of Clausen's work (1938), considerable
World. study is still needed to understand the variation
Ophioglossum reticulatum is variable in the within O. nudicaule. The Peruvian plants here
shape and base of the sterile lamina and in the identified cannot be clearly placed within any of
frequency of secondary areoles and free, included the varieties delineated by Clausen; but, aside from
veinlets. These characters intergrade freely with the fact that their fertile segments do not rise too
those of the often recognized O. petiolatum, so that conspicuously beyond the sterile lamina, they seem
it is impractical to separate the two as distinct taxa. to fit best under var. tenerum. Diagnostic features
Thus circumscribed, O. reticulatum is perhaps the of O. nudicaule are so few and variable that it is
most ubiquitous of any species in the genus. often difficult to discern which are innate differ-
ences and which are merely induced by rigorous
Amazonas: Prov. Bongara, Shilla. Young & Eisenberg habitat or attributed to degree of maturity of the
460 (MO). La Libertad: Prov. Pataz, Pumatambo, Puerta individual plant. that this and other
It is likely
del Monte. Lope: & Sagdstegui 3438 (GH). San Martin:
Monte Campana, near Tarapoto, Spruce 4709 (p, us).
varieties acceptedby Clausen are not supportable,
as evidenced by the studies of Wagner et al. ( 1 984),
Lima: Lomas de Quilmana, Coronado 27 (us). Prov.
Chancay, Lomas de Lachay, Coronado & Velarde 15 (uc, which suggest that they "all should be subsumed
us); Tryon & Tryon 5419 (GH). Pasco: Prov. Oxapampa, under the single binomial, O. nudicaule."
Chontabamba, D. Smith & Brack 3056 (F, MO). Ayac-
ucho: Aina, between Huanta and Rio Apurimac, Killip
& Smith 22809 (GH, NY, us). Apurimac: Prov. Abancay, Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui
et al. 6412 (GH, HUT). Lima: Prov. Huarochiri, Dist. de
Cachora, Huillcayoc, Vargas 9096 (MICH, uc). Cuzco:
Mariatana, Cueva Mortero, Cerrateet al. 4812 (GH, USM).
Prov. Urubamba, Machu Picchu, Leon 452 (GH, USM);
Cuzco: Prov. Acomayo, Quenco Grande, near Acomayo,
Tryon & Tryon 5400 (GH, us), 5419 (GH). Madre de Dios:
Prov. Manu, Parque Nacional Manu, Foster et al. 11486
Vargas 9752 (F, GH, K, ucp.p., us). Prov. Espinar, Yauri,
Vargas 5619 (MICH). Prov. Urubamba, Vargas 14122
(F).
(GH).
scure, areoles small, not enclosing secondary ar- greatly exceeding the lamina, fertile spike 0.6-1.2
eoles or (except rarely) free veinlets. Fertile seg- cm long, on a stalk 1.3-4 cm long.
ment solitary, borne at base of sterile lamina, on
mature leaves somewhat exceeding the lamina, In open, grassy areas, 2840-3850 m, Junin, Cuz-
fertile spike 0.5-1 .2 cm long, on a stalk 0.8-2. 5(-4) co, Puno.
cm long. Peru and Bolivia.
Leaves of this rare fern grow with proximal por-
In high rocky meadows and slopes, with grasses tions of leaves embedded in the soil. This is evi-
On moist, open, grassy slopes, 600-3900 m, iform, fertile stalk, and has thus far been found
Huanuco, Lima, Junin, Apurimac, Cuzco. only at low elevations. However, we have exam-
Southern United States; Mexico to Honduras; ined some Bolivian plants of this subspecies, from
Colombia and Venezuela south to Argentina and elevations to 3000 m, which are more variable:
Chile. sometimes more robust, with stouter fertile stalks,
and with sterile laminae broader and less attenuate
Among the diminutive species of Ophioglos-
to base and apex. Subspecies lusitanicum, from
sum, a few plants are found with stems which are
somewhat swollen. However, only in O. crotalo- Europe, Asia, and Africa, has obtuse sterile lam-
is the stem so conspicuously globose. inae with larger and fewer areoles. Subspecies cal-
phoroides
With this should probably be included O. opacum ifornicum (Prantl) Clausen (California & Mexico)
has laminae larger and broader, with less attenuate
Carmichael, from Tristan da Cunha. It is said to
differ from the former merely in the somewhat bases than in ssp. coriaceum and, according to
shorter fertile segment and slightly smaller globose Clausen (1938), perhaps does not merit distinc-
stem. tion.As with the O. nudicaule complex (q.v.), far
more study is needed to clarify the taxonomy, at
Huanuco: Prov. Huanuco, Huanuco-Tingo Maria road, both inter- and intraspecific levels.
from Bandera Blanca to Carpish Tunnel, Luteyn & Lu-
teyn 5464 (NY). Lima: Prov. Lima, Lomas de Lurin, Lima: Prov. Chancay, Lomas de Lachay, Coronado &
Ferreyra 9566 p.p. (USM). Junin: Huaytapata. near Ha- Velarde 16 (GH, uc, us); Tryon & Tryon 5414 (GH p.p.,
cienda Conocancha, Tiller 14 (USM). Apurimac: Prov. us).Lomas de Quilmana, Coronado 27 (uc, us p.p.).
Abancay, Cachora, Vargas 9054 (MICH p.p., MO). Cuzco: Lomas de Lurin. Ferreyra 9566 (GH, USM p.p.). Lomas
Prov. Calca, Vilcabamba, Vargas 3978 (us). Prov. Can- de Atocongo, Grant 7509 (GH).
as, slopes of San Andres de Checca, Vargas 11010 (F,
uc).
Comments
6. Ophioglossum lusitanicum L. ssp. coriaceum (A.
Ophioglossum ellipticum Hooker & Grev., Icon,
Cunn.) Clausen, Mem. Torrey Bot. Club 19(2):
fil. 1, t. 40A. 1831. TYPE: illustration, based
161. 1938.
12 FIELDIANA: BOTANY
on specimens collected in Demerara, "British to pinnately compound or rarely simple or pal-
Guiana," Parker. mate, monomorphic to somewhat dimorphic. Pet-
iole with an expanded, stipular base, there swollen
This perhapsmay be expected in Peru, as its and sometimes subarticulate, frequently also with
range thus
isfar reported from Guatemala to Costa several swollen and darkened nodes throughout
Rica, and the Guianas southward to Brazil and its Pinnae commonly borne in opposite
length.
(possibly) Bolivia. Although usually a smaller plant pairs from swollen, darkened nodes on the rachis.
than Ophioglossum reticulatum, the aspect of the Veins free or (in Christensenia) reticulate. Spo-
sterile lamina is similar in that the primary areoles rangia borne on the abaxial surface of leaves, with
contain a number of secondary areoles and usually thickened walls, lacking an annulus, separate and
some However, the secondary ar-
free veinlets. contiguous or fused into indurated synangia and
eoles are much more numerous in O. ellipticum, opening by terminal pores. Spores ellipsoidal and
and the walls are thinner and far less distinct than monolete or trilete, rugose or echinate, very nu-
those of the primary ones. Additionally, laminae merous (1,500-1,700) in each sporangium.
in O. ellipticum have a somewhat distinct midvein
and a pale, median band of tissue which persists Of the seven genera (about 1 50 species) in Mar-
to within a short distance of the apex. It is most attiaceae, two are represented in Peru: Danaea,
closely related to O. nudicaule, in fact, the plants confined to the Neotropics, and Marattia, pan-
heretofore recognized as O. ellipticum may be tropical. Next to Ophioglossaceae, it is considered
merely O. nudicaule with larger and more fully to be the most primitive of the living fern families
developed leaves (see Wagner et al., 1984). and can be easily recognized by the character of
the sporangia being fused into thick, elongate, dou-
ble-rowed synangia.
Family 2: MARATTIACEAE
References
Marattiaceae Bercht. & J. S. Presl, Prir. rostlin 1 :
272. 1820. TYPE: Marattia Sw. DEVRIESE, W. H.,ANDP. HARTING. 1853. Mon-
ographic des Marattiacees, Leiden, Nether-
Danaeaceae Agardh, Aphor. hot. 117. 1822. TYPE: lands.
Danaea Sm. TRYON, R. M., AND A. F. TRYON. 1982. Mar-
attiaceae, pp. 39-50, in Ferns and allied plants,
Stem fleshy, stout to massive, creeping or erect, Springer- Verlag, New York.
sometimes scaly. Leaves moderate in size to huge UNDERWOOD, L. M. 1909. Marattiaceae, in N.
(3 m or more), circinate in vernation, 1 -pinnate Amer. fl. 16: 15-23.
a. Leaves 1-3-pinnate or more, monomorphic; synangia scattered, borne near the ends of veins
I. Marattia
a. Leaves 1 -pinnate, somewhat dimorphic (fertile ones commonly longer and narrower and with smaller
pinnae); synangia nearly covering the abaxial surface of fertile pinnae II. Danaea
monomorphic. Lamina with expanded tissue es- are borne near the ends of veins on a receptacle,
14 FIELDIANA: BOTANY
sessile or (in M. laevis) on short, thickened stalks. some Brazilian specimens of M. laevis with "soros
Spores ellipsoid, monolete. ... stipitati," upon which character Hooker (fol-
lowing Smith's unpublished description) partially
This is a pantropical genus of about 40 species, based the new species M. Kaulfussii (1839). Ob-
commonly occurring in dense, wet forests. It is viously Hooker had seen the type of M. laevis, and
readily separated from other Neotropical genera supposed it lacked the stalked synangia; therefore,
by the large, decompound, subternate leaves with he surmised that the Brazilian collections of Kaul-
stipular bases, and the distinctive ovoid synangia. fuss represented a new species. Finally in 1936
A single species is known in Peru. Alston (J. Bot., p. 174) combined the two names,
but failed to specify reasons for doing so. In turn-
Leaves 24 m long, commonly 3-pinnate-pin- (F, GH, us). Junin: Schunke Hacienda above San Ramon,
Occurring in dense forests, principally along the Plants terrestrial, with stems creeping to decum-
Cordilleras Centraland Oriental, in Amazonas, bent or erect. Leaves to 2 m
long, 1 -pinnate (or
Huanuco, Pasco, Junin, and Cuzco, 1400-2400 m. simple in 2 species outside Peru), monomorphic
Costa Rica; Cuba; Hispaniola; Colombia; Ven- to somewhat dimorphic (fertile ones commonly
ezuela; Brazil; Ecuador; Peru; Bolivia. longer and narrower and with smaller pinnae). Pet-
In various herbaria, M.
been confused
laevis has iole (in most species) nodose, scaly, bearing sti-
with M. alata, a species from the Greater Antilles. pules at the base. Rachis nodose, nonalate to con-
Although leaf outline is similar in each, M. laevis spicuously alate at least distally, glabrous or more
is easily distinguished by its stalked synangia and often with appressed scales. Lamina with a distinct
awned penultimate axes, these characters are apical segment, or this replaced by a proliferous
unique in the genus, at least in New World species. bud, provided with scales abaxially, those of the
There has been some confusion as to the identity laminar tissue minute and scattered. Sterile pinnae
of M. laevis, beginning with Smith's original de- opposite or subopposite, straight to subfalcate, ses-
scription, which mentioned nothing about the dis- sile or short-stalked, entire and often undulate, or
tinctive stalked synangia and adaxial awns. Kaul- serrate near the apex. Veins free. Fertile pinnae
fuss (Enum. fil., p. 32, 1824) mentioned seeing similar to the sterile ones, but commonly reduced
16 FIELDIANA: BOTANY
in size, and with vegetative tissue nearly lacking. America may be synonymous with some in the
Sporangia nearly covering the abaxial surface, co- West Indies. Although about 35 species have been
alescing into much elongated, indurated synangia recognized, 20-25 is a more realistic total to expect
and opening by terminal pores. Spores monolete, once a greatly needed revision has been under-
the surface prominently echinate. taken. Six species from Peru are recognized in the
following treatment.
Danaea
confined to the Neotropics. With its
is
nae, number of veins per centimeter, veins paired UNDERWOOD, L. M. 1 902. A review of the genus
in origin versus distinctly forked. Species previ- Danaea. Bull. Torrey Hot. Club, 29: 669-679.
ously thought of as distinct in Central or South
c. Petiole of mature plants with 1-3 nodes; apices of sterile pinnae serrate; larger sterile pinnae
1 .2-2.8 cm broad d
d. Sterilepinnae broadly oblanceolate, broadest above the middle, abruptly acuminate at the
apex; veins predominantly simple, occasionally paired in origin or forked
3. D. oblanceolata
d. Sterile pinnae lanceolate to oblong-lanceolate, broadest near or below the middle, tapering
e. Sterilepinnae subequilateral at the truncate base; veins simple, rarely forked; petiole nodes lacking;
terminal segment of sterile lamina much shorter than larger pinnae, or replaced by a proliferous
bud 6. D. trichomanoides
1 . Danaea nodosa (L.) Sm., Mem. Acad. Roy. Sci. Danaea grandifolia Underw., N. Amer. fl. 16: 18. 1909.
(Turin) 5: 420. 1793 Figure 4b-c TYPE: Colombia, Valparaiso, Santa Marta, //.
H. Smith 992 (holotype, NY; isotypes, BM, F!, GH!,
MO!, us!).
Acrostichum nodosum L., Sp. pi. 1070. 1753. LEC-
TOTYPE (designated by Proctor, Flora Lesser
Antilles 2: 48. 1977): Plumier, Traite foug. Amer., Sterile leaves to 2 m long and 60 cm broad,
/. 108, based on specimen from Martinique or
imparipinnate; petiole lacking nodes, sparsely sca-
Mag. Neuesten Entdeck. Gesammten Naturk. 5: pressed, dark brown, peltate, nearly circular and
307. 1811. TYPE: "in antillis" (p). ca. 1 mm in diameter or to 3 mm
long, margins
merit of separating D. nodosa and D. elliptica. pair of pinnae, these broadest at or above the mid-
While the characters of nodose or non-nodose pet- dle, with margins more crisped. All fall within the
iolesand pinnae number usually serve to distin- natural variation of this species.
guish the two, even these have not been reliable
consistently in specimens seen from Peru. Further Amazonas: Prov. de Bagua, valley of Rio Maranon
study is needed. near Cascadas de Mayasi, Wurdack 1934 (GH, us). San
Martin: On ridge in jungle E of Tingo Maria, Allard
21381 (GH, us). Loreto: Fierro Cano a km 4 del Centre
Ama/onas: E of Huampami, near Pedro Alberto, Ber- al. 1439 (MO).
Forestal Jenaro Herrera, Spichiger et
lin 1524 (MO). San Martin: Prov. Rioja, km 390, Pedro
Huanuco: Churubamba, Hacienda Mercedes, trail
Dist.
Ruiz-Moyobamba road. D. Smith & Vasquez 4726 (F, to Balsa-playa, Mexia 817 Oa (uc). Pasco: Prov. Oxa-
MO). Loreto: Balsapuerto (lower Rio Huallaga basin), Kil-
pampa, Palcazu Valley, Iscozacin, Foster 9502 (MO).
lip & Smith 28410 (GH, us). Huanuco: Prov. Pachitea, Junin: Pichis Trail, Yapas, Killip & Smith 25507 (GH,
Dist. Honoria, en frente a Tournavista, al este del Rio
NY, US).
Pachitea, J. Schunke 1800 (F, GH). Pasco: Puerto Ber-
mudez (as Junin), Killip & Smith 2651 1 (NY, us). Junin:
Prov. Satipo, near Satipo, van der Werffet al. 8644 (MO).
Cuzco: Prov. Quispicanchi, Punkiri Rio Arriba, Vargas 3. Danaea oblanceolata Stolze, Amer. Fern J. 77:
16121 (GH). Madre de Dios: Prov. Tambopata, SSW of 33. 1 987. TYPE: Peru, Dept. Pasco (as Junin),
Puerto Maldonado, Barbour 5178 (F, GH, MO).
Cahuapanas, on Rio Pichis, Killip & Smith
26777 (holotype, us!; frag., F!).
18 FIELDIANA: BOTANY
rate apex, abaxial surface amply provided with Colombia; Venezuela; Peru.
minute, dark brown scales; veins commonly sim- Except for Danaea nodosa, this is the most widely
ple, but sometimes paired at origin or forked. Fer- distributed representative of Danaea in Peru.
tile pinnae 12-14 pairs, larger ones 7-8 cm long Moreover it is likely that the Central American D.
and 0.8-1 cm broad, short-stalked, the apex ob- cuspidata Liebm. also belongs here, along with one
tuse. or two West Indian species. A number of taxa with
pinnae under 2.5 cm broad were separated by Un-
0-500 m, thus far
Terrestrial in dense forests, derwood (1902), merely on the degree of forking
known only from Peru: Pasco and Ucayali. and spacing of veins, an inconsistent character cor-
This is perhaps most closely related to D. alata related rarely or not at all by other features, hence
Sm., of the West Indies and Venezuela, especially a comprehensive revision of the genus should prove
in that both species have predominantly simple many of these to be synonymous. Peruvian spec-
veins. However, in D. oblanceolata pinnae are imens of D. moritziana apparently are more vari-
fewer and relatively shorter and broader, and most able than in other areas, sometimes having smaller
are broadest well above the middle, where the and fewer pinnae as in Venezuela, sometimes with
margins then bend abruptly to a short-acuminate more and larger pinnae as in "D. cuspidata" of
apex. In D. alata, as in all members of the D. Costa Rica. A few very large specimens from Tin-
moritziana complex, pinnae are broadest at or near go Maria have pinnae with rather glossy under-
the middle, from whence they taper gradually to surfaces and extremely attenuate apices. Very
a moderately acuminate or attenuate apex. closely related to this is D. alata Sm. of the Lesser
Antilles, Trinidad, and Venezuela. Although D.
Pasco: Oxapampa, Palcazu Valley, Iscozacin, ./?. Foster alata tends to have more numerous pinnae than
9466 (MO), 10049 (F). Ucayali: Vicinity of Aguaytia, Croat D. moritziana, the only consistent difference be-
20938 (MO).
tween the two is that of simple versus forked veins.
To further add to the confusion, some specimens
of D. moritziana in Peru have been incorrectly
determined as D. stenophylla Kunze, which is
4. Danaea moritziana Presl, Abh. Konigl. Bohm
Ges. Wiss. 5(4): 35. 1845. TYPE: Venezuela, merely a synonym for D. alata.
Underwood (1902) described both of the latter
Colonia Tovar, Moritz 257 (holotype, PR?; is-
as having unforked veins and distinguished them
otype, L; frag., us!; photo, us). Figure 4a.
principally by the degree of crowding of veins, and
that D. stenophylla was a "small" plant. Later
Sterile leaves to 1.2 m long and 35 cm broad
(1909) he reversed himself by describing D. steno-
(somewhat broader than fertile ones), imparipin-
phylla as a "rather tall plant" and as having mostly
nate, or sometimes with the apical segment re-
forked veins. Obviously he had not seen the type
placed by a proliferous bud; petiole with 1-3 nodes,
collection, for the isotype (2 sheets) at Kew con-
sparsely scaly; rachis scarcely alate, or slightly so
tains leaves to nearly a meter long, with veins
in younger plants; pinnae 8-16 pairs, subsessile to
obviously simple (only rarely forked), differing
short-stalked, elliptic to oblong-lanceolate, larger
from D. alata in no way that we can perceive. We
ones (5-)6-20 cm long and 1 .2-2.2 cm broad, ine-
cannot be certain which West Indian plants with
quilateral at base, narrow and rounded to cordate
forked veins were seen by Underwood, and later
basiscopically, broader and cuneate to subtruncate
by Proctor (1977), but perhaps they are D. ja-
acroscopically, gradually tapering to an acuminate
maicensis.
or long-attenuate apex, margin subentire but con-
spicuously serrate at apex, veins (and often the
tissuebetween the veins) amply provided with mi- Cajamarca: Tambillo, Jelski 1073 (P). Amazonas: Bag-
nute scales; veins predominantly forked at or near ua, ca. 25 km E of La Peca, Barbour 2969 (MO). San
the base, occasionally simple, or paired at origin. Martin: Monte Campana, Tarapoto, Spruce 4711 (P).
Loreto: Rio Corrientes, Cachuela, McDaniel 11198 (F,
Fertile pinnae 8-15 pairs, larger ones 4-11 cm GH, MO). Huanuco: Tingo Maria, Tryon & Tryon 5278
long, 0.6-0.8 cm broad, subsessile or short-stalked, (F, GH, uc, us). Pasco: Prov. Oxapampa, San Alberto,
the apex acute to apiculate. Cordillera de Yanachaga, van der Werffet al. 8434 (MO).
Junin: Chanchamayo Valley, C. Schunke 156 (F, us).
Cuzco: Prov. La Convention, ca. 10 km from Hacienda
Terrestrial, in dense forests, often along streams
Luisiana, Dudley 10412 (GH). Madre de Dios: Prov. Manu,
or in ravines, 300-2300 m, Cajamarca to Loreto, Cerro de Pantiacolla, Rio Palotoa 10-15 km NNWof
south to Cuzco and Madre de Dios. Shintuya, Foster 10750 (F).
er than size,however, they share few other similar 272. 1820. TYPE: Osmunda L.
20 FIELDIANA: BOTANY
fined to the Old World. This is a very old fern I. Osmunda
family; only Ophioglossaceae and Marattiaceae are
considered to be more primitive. Like the other Osmunda L., Sp. pi. 1063. 1 753. TYPE: O. regalis
leptosporangiate members of the Filicales, Os- L. Figure 5.
mundaceae have sporangial walls only one cell
thick, yet the family is similar to the Eusporan- Plants terrestrial. Leaves coarse, 1-2 or long- m
giatae in that the annulus is lacking or is composed er, 1-2-pinnate, completely or partially dimor-
of only a few thickened cells. phic, the fertile ones (or portions of them) lacking
green leaf tissue. Sterile lamina commonly gla-
brous at maturity, but the axes often sparsely to
References moderately provided with trichomes. Veins free,
at least 1 -forked. Sporangia relatively
large, de-
BENEDICT, R. C. 1909. Osmundaceae, in N. veloping simultaneously, commonly borne in
Amer. fl., 16: 27-28. clusters on the segments of the fertile leaves. Spores
TRYON, R. M., AND A. F. TRYON. 1982. Os- tetrahedral-globose, trilete, rugose or crested with
mundaceae, pp. 50-57, in Ferns and allied plants, slender echinate processes.
Springer- Verlag, New York.
This is a genus of about 1 species which gen-
erally grow in swampy areas in temperate and
a. Sterile lamina 1-pinnate-pinnatisect; ultimate segments entire, the veins commonly 1 -forked; fertile
lamina nonfoliaceous throughout 1 O. cinnamomea .
a. Sterile lamina 2-pinnate; ultimate segments serrulate, the veins commonly 2-forked; fertile lamina
nonfoliaceous and fertile only in the distal portion 2. O. regalis var. spectabilis
1. Osmunda cinnamomea L., Sp. pi. 1066. 1753. trichomes abundantly interspersed among the
TYPE: United States, Maryland, Kalm (ho- crowded sporangia.
lotype, LINN 1 244. 1 2; photo, F). Figure 5c-d.
In wet places, usually in sphagnum swamps,
Leaves completely dimorphic, vegetative tissue 1700-2800 m, Cajamarca, Amazonas, and Pasco.
Eastern and central United States and Canada;
essentially lacking in fertile ones. Petiole glabrous
or moderately provided with tortuous, filiform, West Indies; southern Mexico to Honduras; Costa
Rica; Colombia; Venezuela; Brazil; Peru; Para-
pluricellular, reddish brown trichomes, sheathed
at the base. Sterile lamina 1-pinnate-pinnatifid, guay; southeast Asia.
tissue glabrous, but with filiform trichomes as on Some earlier authors recognized this in the Neo-
the petiole appearing especially at bases of pinnae, tropics as var. imbricala (Kunze) Milde, distin-
rachis narrowly alate adaxially; pinnae to 10 cm guishing it from North American plants on the
basis of darker-colored tomentum and thicker lam-
long, spreading to strongly ascending, subopposite,
articulate with age at the rachis, deeply incised to
ina. We find little justification for the separation.
22 FIELDIANA: BOTANY
2. Osmunda regalis L. var. spectabilis (Willd.) A. length of their stalks. The characters are too in-
Gray, Manual ed. 2: 600. 1856. Figure 5a-b. significant and inconsistent to merit such a dis-
tinction.
Osmunda spectabilis Willd., Sp. pi. 5: 98. 1810. TYPE:
United States, Pennsylvania, Muhlenberg (holo- Amazonas: Mendoza, on rocky wall of ravine, Woyl-
type, B, Herb. Willd. 19504; photo, GH). kowski 8253 (GH, MO, us). Prov. Bongara, W
& S margins
Osmunda palustris Schrader, Gott. Gel. Anz. 866. of Laguna Pomacocha, Wurdack 896 (us). Pasco: Prov.
1824. TYPE: not located.
Oxapampa, Canyon de Huancabamba, Leon 612 (GH).
Osmunda regalis ssp. palustris (Schrader) Love & Love, Cuzco: Empalizada, Biies 1705 (us).
Taxon 26: 324. 1977.
a. Leaves erect or suberect; pinnae lacking arrested buds in branch axils; sporangia not covered by a
laminar flange b
phyllitidis (L.) Sw. (Osmunda phyllitidis L.). features are often quite variable, such as: relative
Figure 6. length of fertile pinnae and sterile portion of lam-
ina; color of rhizome indument; degree of dissec-
Coptophyllum Gardn., London J. Bot. 1: 133. 1842. tion of pinnae. Color of trichomes is particularly
TYPE: Coptophyllum buniifolium Gardn. = Ane- difficult to assess, inasmuch as each observer sees
mia buniifolia (Gardn.) Moore.
colors in different ways, and definitions of color
Anemirhiza J. Sm., Seemann, Bot. Voy. Herald 243.
1854. TYPE: Anemirhiza adiantifolia (L.) J. Sm.
terms are usually difficult to interpret. Rhizome
(Osmunda adiantifolia L.)
= Anemia adiantifolia indument in Anemia generally grades from orange
(L.) Sw. to reddish orange to red to brownish red; in many
specimens the distinctions are clear, but in others
Plants terrestrial. Stem decumbent to creeping, they are somewhat inconstant. Hybridization is
rarely erect, bearing short to long trichomes and not uncommon in the genus and must be taken
fibrous roots. Leaves ca. 1-75 cm long, erect, par- into consideration when attempting identification
tially (as in Peruvian species) dimorphic, with a of collections. The following key will be more ef-
pair of fertile pinnae at the base of or below the fective if attention is given to combinations of
sterile lamina, or with similar leaves longer and characters, whenever possible.
more erect than the wholly sterile leaves, or wholly
dimorphic, the whole leaf either sterile or fertile.
Sterile lamina 1-2-pinnate (rarely pinnatifid); veins Reference
free, or rarely anastomosing without included free
veinlets. Sporangia borne on fertile segments which MICKEL, J. T. 1962. A monographic study of the
are reduced to mere axes or have narrow borders fern genus Anemia, subgenus Coptophyllum.
of laminar tissue along the axes. Spores tetrahe- Iowa State Coll. J. Sci., 36: 349-482.
dral-globose, trilete.
d. Petiole stout (1-2 mm thick), commonly yellow; fertile pinnae erect, with stalk 1.5-5 cm
long 3. A. flexuosa
c. Apex of stem horizontal, exposed beyond the most recent petiole bases; fertile pinnae commonly
longer than sterile portion of the lamina e
e. Lamina hirsute to glabrate, petiole commonly glabrate and atropurpureous (rarely light
brown to yellow) 4. A. ferruginea
24 FIELDIANA: BOTANY
Lamina and petiole conspicuously lanate, petiole yellow
e. 5. A. myriophylla
h. Sterile pinnae crenulate or denticulate, or rarely with a few deep lobes on several proximal
pinnae 7. A. pastinacaria
g. Petiole less than 6 cm long; lamina abruptly terminating in a broadly obovate or obdeltoid
apical segment; apex of sterile pinnae broadly rounded 8. A. oblongifolia
f. Veins anastomosing; stalk of fertile pinnae usually equaling or shorter than the panicle; sterile
pinnae (larger ones) 5-14 cm long 9. A. phyllitidis
1. Anemia din at a Mickel, Amer. Fern J. 56: 58. 2. Anemia villosa Willd., Sp. pi. ed. 4, 5: 92. 1 8 0.
1
1966. TYPE: Peru, Junin, along Rio Perene, TYPE: "America meridionale," Humboldt &
Killip & Smith 21594 (holotype, us!; isotypes, Bonpland (holotype, B, Herb. Willd. 19496;
F!, NY!). photos, GH, us). Venezuela, Prov. Cumana,
"Between Catuaro and Cariaco": fide HBK.,
Stem with apex ascending and thickly covered Nov. gen. sp. fol. ed. 1: 26. 1815.
with petiole bases, densely provided with orange
trichomes. Leaves 10-25 cm long, 4-9 cm broad, Stem with apex ascending and thickly covered
pilose to hirsute, the suberect fertile pinnae borne with petiole bases, or the apex horizontal and ex-
8-16 mm below the sterile portion of the lamina; posed beyond the most recent petioles, densely
petiole dark brown, less than mm thick. Sterile
1 provided with orange to brownish red trichomes.
lamina 1 -pinnate-pinnatisect to 2-pinnate, with 5- Leaves 10-55 cm long, villous to hirsute, the erect
1 1 pairs of pinnae; pinnae dissected nearly or quite fertilepinnae borne 3-18 mm
below the base of
to the costa, the ultimate segments joined at the the sterile portion of the lamina; petiole yellow to
base or discrete, but broadly adnate, not stalked; light brown, 1-2 mm thick. Sterile lamina 1 -pin-
veins free. Fertile pinnae suberect, usually shorter nate to pinnate-pinnatisect, with 7-19 pairs of pin-
than the sterile portion of the lamina, short-pe- nae; pinnae with rounded lobes, not cut entirely
tiolulate (stalk 0.3-0.8 cm long), the ultimate fer- to the costa (except rarely as to proximal lobes of
tile segments with narrow bands of laminar tissue basal pinnae); veins free. Fertile pinnae erect,
on each side of the axes. slightly shorter than to (more typically) somewhat
longer than the sterile portion of the lamina, their
In thickets and deep forests, 600-1 300 m, Junin. stalks 1-5 cm long, the ultimate fertile segments
Panama; Colombia and Venezuela to Bolivia with narrow bands of laminar tissue on each side
and western Brazil. of the axes.
One of the distinguishing features of clinata A
is the suberect or slightly spreading position of the
On rocky, open slopes, 900-2000 m, Cajamarca,
fertile pinnae, which are rigidly erect in most species
Amazonas, San Martin, Ayacucho, Cuzco.
of Anemia. It is also one of the smaller and more
Surinam to Colombia, south to Peru; east coast
delicate species, with slender, wiry petioles less
of Brazil.
than 1 mm
thick.
This appears to differ only quantitatively from
Anemia flexuosa, the latter with sterile pinnae more
Junin: Near La Merced, E of Quimiri Bridge, Killip &
Smith 23591 (NY, us). Colonia Perene, Killip & Smith strongly dissected and, typically, with fertile pin-
25036 (F, NY, us). La Merced, Kunkel 564 (OH). Chan- nae shorter than the sterile portion of the lamina.
chamayo Valley, C. Schunke 78 (F, us). Fertile pinnae of A. villosa are commonly longer
26 FIELDIANA: BOTANY
than the lamina, but occasionally equal or exceed gas 19836, Prov. La Convencion, alt. 1650 (GH). m
it. Evidently these are the reasons why Prantl con- Leaves are much thinner in texture than typical
sidered it merely a variety of A. flexuosa. See Mickel A. flexuosa, with some anastomosing veins, and
(1962) for more detailed discussion. fertile pinnules are long-petiolulate.
Cajamarca: Prov. Jaen, San Patricio, Chontali, Chi- Amazonas: Rodriguez de Mendoza, Soukup 5023 (us).
moy65 (USM). Amazonas: Prov. Chachapoyas, Cano Santa Huanuco: Vilcabamba, on Rio Chinchao, Macbride4985
Lucia, Wurdack 599 (F, GH, NY, uc, us). San Martin: (F, NY, us), Stork & Morton 9876 (F, uc, us). Junin:
Tarapoto, in monte Campana, Spruce 4708 (GH, NY, p, Huacapistana, Killip & Smith 24152, 24324 (F, NY, us).
us). Ayacucho: Prov. La Mar, Aina, Ldpez & Soukup Ayacucho: Ccarrapa, between Huanta and R;o Apuri-
15.088 (GH). Cuzco: Prov. La Convencion, Urusaiwa, mac, Killip & Smith 22319 (F, GH, NY, us). Cuzco: Prov.
Vargas 22349 (GH). La Convencion, Valley of the Sambray, Mexia 8038 (F,
GH, MO, uc).
laminar tissue on each side of the axes. rocky slopes and clay banks, 850-2200 m, San
Martin, Huancavelica, Cuzco.
and open woods or on moist rocky
In thickets Honduras; Guyana to Colombia, south along
the Andes to Bolivia and Brazil.
slopes or clay banks, 900-3000 m, Amazonas,
Huanuco, Junin, Ayacucho, Cuzco.
The variety ahenobarba (Christ) Mickel, from
Surinam to Colombia, south to Bolivia. Brazil, differs in its more deeply dissected pinnules
Peruvian specimens of Anemia flexuosa with acute to acuminate tips. Both are difficult to
Many
are to be found in various herbaria; especially distinguish from Anemia tomentosa (Savigny) Sw.,
abundant are collections from Huanuco, Junin, especially var. anthriscifolia (Schrader) Mickel,
and Cuzco. This is a highly variable species, par-
from northern Argentina, Bolivia, Paraguay, and
southeastern Brazil. See Mickel (1962) for further
ticularly in the dissection of sterile laminae, some-
times 2-pinnate-pinnatifid, typically 2-pinnate, comments.
occasionally approaching the 1 -pinnate-pinnatifid San Martin: Tarapoto, Spruce 4044 (GH, NY, p, us).
condition of A. villosa (q.v.) and thus is often con- Huancavelica: Prov. Tayacaja, Dist. Huachocolpa, near
28 FIELDIANA: BOTANY
a few specimens in Peru tend to be intermediate these characters are too variable and inconsistent
even in these characters. See A. hirsuta for further to be of real value. We
tentatively maintain it here
discussion. as a synonym, although we have not seen the type
to fully substantiate this.
San Martin: Tarapoto, Spruce 4 134 (p in part). Loreto:
Yanayacu, Biies 1037 (us). Huanuco: Yanano, Macbride Cuzco: Echarate, Camino de Sahuayacu, Bites 845 (us).
3816 (F, us). Junin: Prov. Huancayo, Pariahuanca, Tovar Ccochayoc, Biies 1712, 1719 (us). Prov. Convencion,
7948a (USM). Ayacucho: Aina, between Huanta and Rio Valle de Lucumayo-Pistipata, Vargas 4176 (MO, uc, us).
Apurimac, Killip & Smith 22614 (F, NY, us). Cuzco: Prov.
Convencion, Hacienda Sahuayaco, Vargas 1655 (GH),
1657 (GH, MO).
9. Anemia phyllitidis (L.) Sw., Syn. fil. 155. 1806.
Figure 6c.
ing in an obovate or obdeltoid apical segment which mate fertile segments essentially lacking laminar
is often as broad as long; pinnae to 1.8 cm long tissue.
and 0.7 cm
broad, strongly inequilateral at base,
truncate or rounded acroscopically and cuneate
Moist, often rocky slopes and ravine banks, in
basiscopically, broadly rounded at apex, the mar-
and at edges of forests, 700-1700 m, Amazonas
gins crenulate or subentire; veins free. Fertile pin- and Cuzco.
to Loreto, south to Huancavelica
nae to 1 8 cm long, the stalk 24
times the length
Mexico to Panama; Greater Antilles; Trinidad;
of the panicle, the ultimate fertile segments essen-
Venezuela and Colombia to Argentina and Uru-
tially lacking laminar tissue.
guay.
This is one of the most common species of Ane-
In forests, on rocky slopes, 1000-1 100 m, Cuz-
mia in Peru and, with its large, subentire pinnae
co.
and reticulate venation, is the easiest to recognize.
Mexico to Panama; Venezuela to Brazil and Bo-
livia.
Amazonas: Prov. Bagua, S of Bagua Grande on Rio
Anemiaoblongifolia is rather distinctive in its
Utcubamba, Hutchison 1479 (uc, us). San Martin: Ze-
coarse pinnae which are broadly rounded at their
pelacio, near Moyobamba, Klug 3440 (F, MO, NY, uc,
us). Loreto: Yanayacu, Biies 2005 (us). Huanuco:
tips and often nearly as broad as long. Sterile leaves Prov.
are commonly very short-petiolate (often subses- Huanuco, Puente Durand, Stork & Morton 9432 (F, GH,
MO, uc, us). Pasco: Oxapampa, Soukup 2361 (F, GH).
sile), and typically are so densely caespitose that
Junin: Yucapata, Woytkowski 6642 (GH, MO, us). Huan-
they give the appearance of a rosette. Some authors cavelica: Prov. Tayacaja, SE of Tintay, Tovar 46 11 (GH).
have separated A. humilis on the strength of its Ayacucho: Aina, between Huanta and Rio Apurimac,
villous sterile pinnae with crenulate margins; but Killip & Smith 22719 (NY, us). Cuzco: Prov.
La Con-
repens Raddi (= A. hirsuta ?), but having pinnae er branches of shrubs and trees. It is widely dis-
less incised.Assuming it belongs to subg. Anemia tributed, primarily pantropic, with several extra-
(unsubstantiated by the description), it could be tropical species both in the New and Old World.
A. pastinacaria, but we cannot be certain without There are 30-35 species, but the number could be
examining the type. somewhat reduced by further study, as some ap-
pear to intergrade rather freely. Three species are
recognized in Peru.
II. Lygodium
a. Primary pinna-branch pinnate, the pinnules stalked, except sometimes near the apex of the primary
branch b
b. Pinnules (at least fertile ones) diminishing in size toward the apex of the primary branch, mostly
expanded at base into lobes, the lobes often discrete or even stalked 1 L. venustum .
b. Pinnules subequal, not expanded at base, subentire, very rarely some basal ones lobed at the base
2. L. volubile
a. Primary pinna-branch radiate, the ultimate segments deeply lobed, joined at the base
. 3. L. radiatum
1. Lygodium venustum Sw., J. Bot. (Schrader) lular trichomes. Primary pinna-branches pinnate
1801(2): 303. 1803. TYPE: Jakob Breyne, to (rarely) 2-pinnate, sessile to short-stalked,
Cent. I, t. 96. 1678, not "Brazil, Breynius" broadly to narrowly deltoid. Fertile pinnules 9-
(designated by Proctor, Flora Lesser Antilles 2 1 alternate and commonly widely spaced, all but
,
2: 51. 1977), a specimen which probably does the distal ones stalked, stalk often nodose at its
not exist. Figure 7a. juncture with pinnule base, sparsely to abundantly
pilose on axes, veins and (occasionally) the leaf
Pinna-stalks 2-10 mm moderately to
long, tissue, larger ones 2.5-5 cm long, gradually di-
densely pubescent with tawny to orange pluricel- minishing in size toward the apex of the primary
30 FIELDIANA: BOTANY
FIG. 7. Lygodium venustum: a, primary axis and pair of pinnae. Lygodium volubile: b, pinna; c, base of pinnule,
(a from Irwin et al. 21469, Brazil, F,b from Jeanpert s.n., Brazil, F, c from L. B. Smith 1352, Brazil, F.)
pas: Pteridophytes, Calif. Acad. Sci. p. 145. 1981, in certain regions outside South America: relative
and Stolze, Ferns & Fern Allies of Guatemala: Part size and shape of ultimate segments and their bas-
I, Fieldiana, Bot. 39: 38. 1976. The closely related es; degree of pubescence on abaxial surface; angle
Lygodium volubile readily can be distinguishedby of veins. None of these are consistently correlated
the fewer, larger, and subequal pinnules on each in the many Peruvian specimens examined.
pinna-branch; the distal ones are nearly or
i.e.,
quite as large as the proximal ones. Pinnules (at Amazonas: Quebrada Huampami, Kayap 1066 (MO).
least fertile ones) of L. venustum gradually dimin- San Martin: Prov. San Martin, Laguna Sauce, Ramirez
ish in size and shape so that subapical ones are
& Sotero 092-85 (F, GH, HUT). Loreto: Rio Tacsha Cur-
aray, Croat 20394 (F, GH, MO, NY). Gamitanacocha, Rio
often one-third to one-half the size of those near
Mazan, J. Schunke 234 (F, GH, NY, uc, us). Madre de
the pinna base. The Breyne plate was cited by Dios: El Pilar, Lopez & Soukup 4601 (GH).
Swartz and is clearly this species.
32 FIELDIANA: BOTANY
Amazonas: Al lado de Huampami, Kayap 1217 (MO, dichotoma (L.) Sm. (Acrostichum dichoto-
Santa Rosa, lower Rio Huallaga below Yu-
us). Loreto: mum L.). Figure 8.
rimaguas, Killip & Smith 28937 (F, GH, NY, us). Puerto
Melendez, below Pongo de Manseriche, Tessmann 4735
(NY). Pasco: Prov. Oxapampa, west side of Cordillera de Lophidium Rich., Actes Soc. Hist. Nat. Paris 1: 114.
San Matias, D. Smith 2011 (F, MO). Ucayali: Prov. Co- 1 792. TYPE: Lophidium latifolium Rich. = Schi-
ronel Portillo, km 89 Carretera Federico Basadre, C. zaea elegans (Vahl.) Sw.
Vdsquez 1 (USM). Actinostachys Wall., Numerical list of plants in East
Indies Company Museum, 1829, description
1.
Lygodium oligostachyum (Willd.) Desv., Mem. Soc. Plants terrestrial. Stem erect or ascending, rath-
Linn. Paris 6: 205. 1827. er densely provided with septate trichomes and
with slender, fibrous roots. Leaves ca. 5-50 cm
ed. 4, 5:
Hydroglossum oligostachyum Willd., Sp. pi. long, caespitose, long-petiolate, glabrous or with
81. 1810. TYPE:
Plumier, Traite foug. Amer., t.
scattered, small trichomes, slightly to partially or
92. 1705 (based on specimens from Haiti, near
Lake Miragoan). wholly dimorphic, fertile ones with elongate fertile
segments borne pinnately or subdigitately at the
This is similar in general aspect to Lygodium apex of laminar axes which are simple or dichot-
venustum, but differs in the conspicuously 2-pin- omously branched. Lamina filiform, grasslike and
nate pinnae and the strongly flexuous (zigzag) and scarcely or not at all foliaceous, or fusiform to
flabelliform (in general outline) and foliaceous.
more delicate primary and secondary pinna rach-
Veins free. Sporangia crowded in or more rows 1
ises.Although L. oligostachyum has been consid-
ered by most authors to be confined to Hispaniola,
on the scarcely foliaceous segments (sporangio-
Duek (1978) assigned to it a broad distribution phores). Spores ellipsoidal, monolete.
the Poeppig and Cuming collections in question, genus. About 30 species occur in tropical or ex-
but they are probably L. venustum and not L. oli- tratropical regions of both hemispheres, five of
which are found in Peru.
gostachyum as suggested. All specimens of the lat-
ter which we have examined in various herbaria
have been collected in Hispaniola, so it is unlikely
to be found in Peru.
Reference
a. Leaves with lamina obviously foliaceous, flabelliform in general outline, the divisions linear-oblong
toobovate 1 S. elegans .
a. Leaves not or scarcely foliaceous, but filiform and grasslike or, if flabelliform in outline, then the
divisions filiform b
b. Sporangiophore pinnatifid, its segments bearing sporangia in a single row on each side of the vein
. .c
c. Leaves 3-6 times dichotomous, dimorphic; axis of the sporangiophore strongly curved at
maturity 4. S. poeppigiana
b. Sporangiophore subdigitate, its segments bearing sporangia in 2 or more (sometimes indistinct)
rows on each side of the vein 5. S. pennula
1. Schizaea elegans (Vahl) Sw., J. Bot. (Schrader) 2. Schizaea pusilla Pursh, Fl. Amer. sept. 2: 657.
1800(2): 103. 1 80 1 Figure
. 8. 1814. TYPE: United States, New Jersey, Bur-
lington Co., Quaker Bridge, Pursh? (not lo-
Acrostichum elegans Vahl, Symb. hot. 2: 104, t. 50. cated).
1791. TYPE: Trinidad, von Rohr (c).
Schizaea flabellum Mart., Icon. pi. crypt. 115, /. 55.
1834. TYPE: Brazil, Prov. Rio Negro, Martins Leaves unbranched, 7-10 cm long, linear, grass-
(holotype, BR; isotype, B; photo, us of B). with expanded lamina essentially lacking, pet-
like,
Lophidium elegans (Vahl) Presl, Suppl. tent, pterid.
77. 1845.
iole 0.40.5 mm broad, rather flattened, or sub-
Lophidium flabellum (Mart.) Presl, Suppl. tent, pterid. terete at base, sterile and fertile leaves erect and
77. 1845. ones apically
straight or slightly flexuous, fertile
bearing a pinnatifid, conduplicate, scarcely folia-
Leaves 20-80 cm long, petiole 15-65 cm long, ceous sporangiophore. Sporangiophore 0.5-0.7 cm
1-2.5 mm in diameter, subterete, obtusely angled long, with 5-7 pairs of linear segments,
abundantly
abaxially. Lamina subcoriaceous, essentially gla- provided on the margins and among the sporangia
brous, flabelliform (in general outline), 1 -several with deep orange to reddish brown, flexuous tri-
times-dichotomously forked or cleft, the divisions chomes. Sporangia in a single row of 5-7 on each
oblong to obovate, lateral margins entire, distal side of the vein.
ones strongly and sharply lacerate. Veins dichot-
omously forked within the ultimate divisions.
Sporangiophores borne along the distal margins
Thus far known only from Pasco, "plant colo-
of the lamina, pinnately branched at the tips of nizing sandy, wet landslide; shrubland on white
the marginal lacerations. Sporangia in a single row sandstone, 2700-2800 m."
on each side of the vein. United States (New Jersey, New York); Canada
(Newfoundland, Nova Scotia).
humus on forest floor, or in sandy or rocky
In Heretofore, Schizaea pusilla has been known to
soilon wooded slopes and ridges, 130-2200 m, occur only in a few scattered locations in north-
eastern North America. The recent collection in
Amazonas, San Martin, Loreto, Huanuco, Pasco,
Junin, Cuzco.
Pasco duplicates the northern plants in every fea-
Southern Mexico to Panama; Trinidad to Co- ture (including spores) except that the sterile leaves
are straight to slightly flexuous instead of con-
lombia, and south to Bolivia and Brazil.
Several varieties have been recognized, based spicuously curling. The great disjunction in dis-
tribution is at first startling until one understands
upon number and shape of laminar divisions,
characters which do not appear to merit distinc- the problems of encountering this very inconspic-
34 FIELDIANA: BOTANY
FIG. 8. Schizaea elegans: a, habit; b, sporangiophore; c, sporangiophore segment, abaxial side; d, sporangiophore
segment, adaxial side. (From Stolze, Ferns and fern allies of Guatemala, 1976.)
long, linear, grasslike, with an expanded lamina belliform after pressing), 5-8 times dichotomously
essentially lacking, petiole 0.6-0.8 in diam- mm forked, the divisions linear, scarcely or slightly
convex abaxially, shallowly sulcate adaxially,
eter, foliaceous. Fertile lamina on somewhat longer pet-
somewhat flattened, or terete at base, sterile and ioles than the sterile, 3 or 4 times dichotomous,
fertile leaves straight or slightly flexuous, fertile the divisions linear, nonfoliaceous, bearing spo-
ones apically bearing a pinnatifid, conduplicate, rangiophores at their apices. Sporangiophores 1.5-
scarcely foliaceous sporangiophore. Sporangio- 2 cm long, with 12-25 pairs of linear segments,
phore 1.2-3 cm long, with (10-) 12-2 5 pairs of amply provided on the segment margins and among
linear segments, abundantly provided on the mar- the sporangia with pale to light brown, flexuous
gins and among the sporangia with pale to light trichomes. Sporangia in a single row of 1 5-25 on
brown, flexuous trichomes. Sporangia in a single each side of the vein.
row of (8-) 10- 18 on each side of the vein.
Thus far known in Peru only from one of the
Thus far known from a single collection (dept. syn types.
unknown) in Peru; elsewhere in open woods or Greater Antilles; Mexico (Chiapas); Costa Rica
clearings and savannas, in sand or sandy or rocky to Venezuela and Guyana; Peru; Bolivia.
soil, sea level to 400 m.
Surinam to Venezuela and northern Brazil;
Peru.
Of the many specimens seen by us in various 5. Schizaea pennula Sw., Syn. fil. 150, 379. 1806.
herbaria, only one was (possibly) collected in Peru: TYPE: "America meridionalis ... habitat in
Poeppig (NY). The label is worded simply "In Pe- America calidiore" (holotype, S-PA).
ruvia, diar. Kunze, 1834." However, Kunze (Lin-
naea9: 19. 1834) lists only one species of Schizaea Actinostachys pennula (Sw.) Hooker, Gen. fil., /. Ilia.
from Peru (41. 5". dichotomd), which is S. poep- 1842.
36 FIELDIANA: BOTANY
In sand and sandy soil of open forests and clear- pinnatisect (rarely pinnate) pectinate. Sporangia
ing, sea level to 260 m, San Martin, Loreto. borne in exindusiate sori on the abaxial surface of
Costa Rica; Puerto Rico; Guadeloupe; Trinidad; ultimate segments, with a short many-rowed stalk
Surinam to Colombia, south to Bolivia and Brazil. of cells and a central to oblique or nearly apical
annulus not interrupted by the stalk. Spores
San Martin: Chazuta, Rio Huallaga, Klug 3986 (F, GH, monolete or trilete, lacking chlorophyll, 1 20 to ca.
MO, NY, uc, us). Loreto: San Juan, vicinity of Iquitos, 800 in each sporangium.
Asplund 14414 (F, p, NY, us). Dist. Iquitos, Maynas, Que-
brada Shushuna, Rimachi 3995 (F, NY), 4870 (NY).
The pseudodichotomous branching of leaves and
the usually pectinate penultimate segments make
this one of the most distinctive of fern families.
These ferns frequently are found in dry, open areas
Family 5: GLEICHENIACEAE and species with larger leaves sometimes spread
over low shrubs to form dense thickets. Approx-
Gleicheniaceae (R. Br.) Presl, Reliq. haenk. 1 70. :
recognized in Peru.
Terrestrial. Stem long-creeping, bearing tri-
a. Axils and axillary buds with scales; veins simple to 1 -forked; sporangia 2-4(-5) per sorus
I. Gleichenia
a. Axils and axillary buds with trichomes; veins 2-4-forked; sporangia ca. 8-15 per sorus
II. Dicranopteris
a. Leaves simple (or very rarely with a single fork), deeply pinnatisect to pinnate; lamina linear
2. G. simplex
a. Leaves pseudodichotomously forked, or with 1-3 pairs of 2-pinnate pinnae; lamina broad, never
linear b
b. Pinnae 2-pinnate, not forked; axillary scales with margins entire 1 . G. bancroftii
b. Pinnae once to several times pseudodichotomous; axillary scales conspicuously ciliate or setose
(or entire in G. nitiduld) c
c. Ultimate segments moderately to densely tomentose on midrib and/or veins abaxially, the
whitish to orange tomentum often obscuring the segment tissue (sometimes thinning in age)
d
d. Larger penultimate segments 1.4-2.2 cm broad; larger ultimate segments 1.2 cm long or
less; pinnae usually 3-4 times-forked; branches strongly ascending, usually tightly crowded
and subparallel with adjacent ones 3. G. pennigera
d. Larger penultimate segments 2.5-8 cm broad; larger ultimate segments 1.5-4.5 cm long;
pinnae 1 - or 2-forked; branches spreading to moderately ascending, not crowded e
e. Sori inframedial, crowding or touching the segment midrib; ultimate segments (larger
ones) 1.8-3 mm broad beyond the dilated base, usually strongly revolute 4. G. bifida
e. Sori medial, rarely crowding the midrib; ultimate segments (larger ones) (3-)3.5 4.5 mm
broad, plane to moderately revolute 5. G. tomentosa
c. Ultimate segments naked to scaly abaxially, never tomentose (if trichomes present, these
scattered, short, or rigid) f
f. Penultimate segments less than 2 cm broad; ultimate segments 2-3.5(-4) times longer than
broad g
g. Scales ample abundant on costae and veins abaxially
to 6. G. revoluta
g. Scales essentially lacking on lamina h
h. Axillary scales broad, ciliate; primary axis conspicuously tuberculate to muricate . .
7. G. tuberculata
h. Axillary scales narrow and rigid, entire or (rarely) sparingly dentate or short-setose;
primary axis smooth 8. G. nitidula
f. Penultimate segments (at least larger ones) 2.2-8 cm broad; ultimate segments (4-)5-12
times longer than broad i
j. Ultimate segments (most of them) remote, separated by a space once or twice their
width; sori medial to supramedial 10. G. remota
38 FIELDIANA: BOTANY
k. Costa scales 1.5-3 mm long; midrib scales filiform, the scale body 1-3 cells wide;
axillary scales 2-3 mm long, with dark, rigid setae 1
1. Gleichenia bum-mini Hooker, Sp. til. 1: 5. 1844. long and less than 3 cm broad. Scales of the petiole
TYPE: Jamaica, Bancroft (holotype, K). essentially lacking, but abundant on the rachis
abaxially, these lanceolate, attenuate, orange to
Mertensia bancroftii (Hooker) Kunze, Linnaea 18: 307. reddish brown, the margins long-ciliate. Ultimate
1844.
segments contiguous at the very base or (especially
Dicranopteris bancroftii (Hooker) Underw., Bull. Tor-
proximal ones) discrete and almost their own width
rey Bot. Club 34: 252. 1907.
apart, strongly revolute, on the abaxial side slightly
Hicriopteris bancroftii (Hooker) Ching, Sunyatsenia 5:
278. 1940. to conspicuously pruinose and scaly, scales along
Diplopterygium bancroftii (Hooker) A. R. Smith, Amer. the midrib similar to those of the rachis, but to-
Fern J. 70: 26. 1980. ward the segment apex grading to pluricellular,
stellate trichomes. Sori medial to inframedial.
Leaves with 1-3 pairs of 2-pinnate pinnae, these
to 1.5 m
long and 40 cm broad, the axes naked or Road cuts, banks and open rocky slopes,
with scattered, linear, or filiform scales. Axillary to Cajamarca, south
(1900-)2700-3800 m, Piura
scales lanceolate to ovate, long-acuminate to at- to Huancavelica and Cuzco.
tenuate, yellowish to light brown, with entire mar- Colombia to Bolivia.
gins. Penultimate segments (pinnules) very nu- There has been some confusion as to publication
merous, crowded, at nearly right angles to the costa, of the name Mertensia simplex. In the Diet. Sci.
pectinate, cut nearly or quite to the costa. Ultimate Nat. 1 827 there is no text, but the plate with anal-
segments strongly revolute, midribs abaxially often ysis constitutes a valid publication. A description
provided with scattered, filiform scales. Sori in- was given at approximately the same time by Des-
framedial. vaux in Mem. Soc. Linn. Paris 6: 200. 1827.
In open forests, commonly on slopes or ravine Piura: Prov. Huancabamba, road to Canchaque,
banks, 2400-2800 m, infrequent in Peru: Huan- Hutchison 1617 GH, NY, uc). Lambayeque: Dist. In-
(F,
uco, Pasco, Cuzco. cahuasi, Laguna Tembladera, Sagdsteguiet al. 12774 (F,
Mexico to Panama; West HUT, MO). Cajamarca: Prov. San Miguel, El Tingo, Sa-
Indies; Colombia;
gdstegui et al. 9517 (F, MO, NY). Amazonas: Prov. Chach-
Venezuela; Ecuador; Peru; Bolivia.
apoyas, banks above swamp on summit of Cerros de
Calla-Calla, Wurdack 1212 (F, GH, NY, us). La Libertad:
Huanuco: Huanuco, within 5 km of Carpish, Tryon & Prov. Pataz, Huaylillas, Puerta del Monte, Lopez & Sa-
Tryon 5316 (F, OH, uc, us). Pasco: Border Prov. Oxa- gdstegui 3457 (GH). San Martin: Dist. Huallaga, Valley
pampa and Pasco, van der Werffet al. 8604 (MO). Cuzco: of Rio Apisoncho, Hamilton & Holligan 540 (us). An-
La Convencion, Valle de San Miguel, Biies 2126 (us). cash: Prov. Huari, Huascaran National Park, Quebrada
Pachachaca, D. Smith et al. 12564 (F, MO). Huanuco:
Yanano, stony clay bank, Macbride 4925 (F, GH, us).
2. Gleichenia simplex (Desv.) Hooker, Icon, pi., Huancavelica: Prov. Tayacaja, Chuspi-Tocas, between
/. 92. 1837.
Colcabamba and Paucarbamba, Tovar 2066 (GH, USM).
Cuzco: La Convencion, Biies 2078 (GH, us).
40 FIELDIANA: BOTANY
Leaves pseudodichotomously forked. Pinnae Sticherus bifidus (Willd.) Ching, Sunyatsenia 5: 282.
1940.
(2-)3 4-forked, the branches (at least in Peruvian
Sticherus mathewsii (Hooker) Nakai, Bull. Nat. Sci.
specimens) strongly ascending, often tightly Mus. 29: 22. 1950.
crowded and subparallel with adjacent ones. Ax-
illary scales lanceolate, orange, the margins co- Leaves pseudodichotomously forked. Pinnae 1-
piously beset with long, tortuous pale or whitish or 2-forked, the branches spreading to moderately
cilia. Larger penultimate segments 1.4-2.2 cm
ascending. Axillary scales lanceolate to ovate, at-
broad, the costae abaxially provided with a mix-
tenuate, light brown to pale orange, the margins
ture of orange or whitish long-ciliate scales and a
amply ciliate. Larger penultimate segments (2.5-)3-
scattered whitish tomentum, adaxially naked or 7 cm broad, the costa on the abaxial side amply
sparsely to amply white-tomentose. Ultimate seg- to abundantly provided with orange, long-ciliate
ments slightly to strongly revolute, larger ones to
scales, on the adaxial naked or sometimes with
1.2 cm long, densely covered abaxially with whit-
sparse whitish tomentum. Ultimate segments
ish tomentum (often thinning in age). Sori infra-
moderately to (typically) strongly revolute, larger
medial, mostly crowding or touching the midrib. mm mm
ones 1 5-35 long and 1 .8-3 broad (beyond
the dilated base), densely covered abaxially with
Open places in forests, on slopes or stream banks,
orange to whitish tomentum (sometimes thinning
1000-2100 m, Loreto?, Huanuco, Pasco, Junin,
in age), the midribs also sparsely provided with
Cuzco.
tawny to whitish, filiform, ciliate scales. Sori in-
Colombia; Venezuela; Peru; Bolivia; Brazil.
framedial, mostly crowding or touching the mid-
Special note should be made of the branching rib.
pattern throughout the range of this species.
Branches may be very strongly ascending, thus
and clearings, often on banks
In forests, thickets
quite crowded and even subparallel with each oth- and 150-1800 m (very rarely to 2200 m)
slopes,
er; or they may diverge at broader angles, as seen
Piura to Loreto, south to Madre de Dios and Puno.
in the type. Thus far, all specimens examined from
Mexico to Panama; West Indies; Colombia and
Peru exhibit the crowded pattern, which might
Venezuela south to Bolivia and Paraguay.
tempt one to consider separation at subspecific
level. However, this difference appears uncorre- Piura: Prov. Huancabamba, slopes of Cerro La Viuda,
lated with other characters,and specimens, at least Sagdsteguiet al. 8 209 p.p. (NY). Antazonas: Prov. Bagua,
outside Peru, seem to grade freely from one pattern along roadside, 37 km NE of Chiriaco, Barbour 4486
to the other. (MO). San Martin: Zepelacio, near Moyobamba, Klug
3458 (F, GH, MO, NY, us). Loreto: Pumayacu, between
" Balsapuerto and Moyobamba, Klug 3242 (F, GH, MO, NY,
Loreto?: Altura de Salarayacu" (Sarayacu?), Biies 847
us). Huanuco: Dist. Churubamba, trail Cotirarda to
(us). Huanuco: Between Huanuco and Pampayacu, Ka-
Mercedes, Mexia 8191 (F, GH, MICH, MO, NY, uc, us).
nehira 153 (GH, us). Pasco: Prov. Oxapampa, Gran Pa-
Pasco: Prov. Oxapampa; Palcazii, Rio Alto Iscozacin,
jonal, D. Smith 5073 (F, MO). Junin: Concepcion, Her- Foster & d'Achille 10098 (F). Junin: Huacapistana, be-
rera 142 (us). Cuzco: Valle de Lares, Hda. Huy-huy, Biies
tween Tarma &
San Ramon, Ferreyra 286 (GH, USM).
1826 (us). Prov. Convention, Lucumayo, Vargas 4221
Ucayali: Vicinity of Aguaytia, on steep slopes along Rio
(us).
Aguaytia, Croat 21003 (F, MO, uc). Ayacucho: Ca. 25
km SW from Hacienda Luisiana and Rio Apurimac, ca.
25 km from Tambo, Dudley 11887 (GH). Cuzco: Prov.
4. Gleichenia bifida (Willd.) Sprengel, Syst. veg. Paucartambo, near Asuncion, West 7122 (MICH, uc).
Madre de Dios: Prov. Manii, Rio Inambari, Chavez 1057
4: 27. 1827. Figure 9.
(MO). Puno: San Gaban to Ollachea, Dillon et al. 1243
(F, MO, NY).
Mertensia bifida Willd., Kongl. Vetensk. Acad. Nya
Handl. 25: 168. 1804. TYPE: Venezuela, Czn-
cas,Bredemeyer(ho\olype,B,Herb. Willd. 19468;
photo, GH). 5. Gleichenia tomentosa (Sw.) Sprengel, Syst. veg.
Dicranopteris bifida (Willd.) Maxon, N. Amer. fl. 16:
4: 27. 1827.
60. 1909.
Gleichenia mathewsii Hooker, Sp. fil. 1 9. : 1 844. TYPE:
Peru, Mathews 1092, in part (holotype, K; photo, Mertensia tomentosa Sw., Kongl. Vetensk. Acad. Nya.
us). Handl. 25: 1 77, /. 5J. 4. 1 804. TYPE: Peru, Herb.
Mertensia mathewsii (Hooker) Fee, Mem. Foug. 1 1: Cavanilles (holotype, s; isotype, p?; photos, us of
122. 1866. S&P).
Sticherus velatus (Kunze) Copel., Gen. fil. 28. 1947. below) from Cajamarca has segments strongly re-
volute and 3 mm
broad or less, but sori are medial
and scales frequently are castaneous at base.
Leaves pseudodichotomously forked. Pinnae 1
-
or 2-forked, the branches spreading to moderately Cajamarca: Prov. Cutervo, La Achira (San Andres-
ascending. Axillary scales lanceolate to ovate, at- Socota), growing over bushes,Lopez & Sagdstegui 5464
tenuate, deep to pale orange, or castaneous at base, (GH, HUT). Amazonas: Prov. Bagua, Cordillera Colan SE
of La Peca, Barbour 3980 (MO). San Martin: "In monte
the margins amply long-ciliate. Larger penultimate
Campana prope Tarapoto," Spruce 4707 (p, us). Huan-
segments 4-7 cm broad, the costa abundantly pro- uco: Playapampa, steep banks, Macbride 4509 (F, us).
vided abaxially with orange, long-ciliate scales, and Pasco: Cushi, trail to Tambo de Vaca, Bryan 687 (F).
adaxially with orange to whitish filiform scales Cuzco: La Convention, ca. 12 km NE from Hacienda
which often grade into a pale tomentum (this Luisiana and Rio Apurimac, Dudley 10545 (GH, us).
of the densely tomentose species of Gleichenia, Dicranopteris affinis (Kuhn) Maxon, Contr. U.S. Natl.
Herb. 24: 47. 1922.
which include G. bifida as well as those combined Maxon, Contr. U.S.
Dicranopteris pruinosa (Mart.)
here under G. tomentosa. We have been unable to Natl. Herb. 24: 49. 1922.
obtain the type specimens of the latter, but type Sticherus pruinosus (Mart.) Ching, Sunyatsenia 5: 284.
1940.
photos (us) indicate clearly enough that ultimate
Sticherus revolutus (HBK.) Ching, Sunyatsenia 5: 285.
segments are too broad and soral lines appear too 1940.
farfrom midribs to be G. bifida. The photos, along Sticherus affinis (Kuhn) Nakai, Bull. Natl. Sci. Mus.
with Swartz's original description ( 1 804) of Mer- 29: 13. 1950.
tensia tomentosa and his subsequent Latin de-
scription (Syn. fil. 164, p. 392, 1806) make it ev- Leaves pseudodichotomously forked. Pinnae 2-
ident that G. tomentosa and G. velata are the same. 4-forked. Axillary scales lanceolate to ovate, lax,
All this confusion within the species complex is thin-textured, orange to reddish brown, the mar-
-
warranted, for G. bifida and G. tomentosa do not gins amply ciliate. Larger penultimate segments I
As seen in
differ strongly. the key, the former has 1.8(-2) cm broad, the costa on the abaxial side
narrower, more strongly revolute ultimate seg- amply to copiously provided with orange to red-
ments, and soral lines usually are closer to the dish brown, long-ciliate scales, on the adaxial side
42 FIELDIANA: BOTANY
the costa naked or filiform-scaly. Ultimate seg- Leaves pseudodichotomously forked, petiole and
ments moderately to strongly revolute, larger ones rachis conspicuously tuberculate to muricate. Pin-
2-3.5(-4) times longer than broad, the midrib and nae 2-3-forked. Axillary scales closely imbricate,
veins scaly as on the costa, those of the veins often ovate to lanceolate, castaneous to reddish brown,
filiform or grading to tortuous, stellate trichomes, the margins amply ciliate. Larger penultimate seg-
the veins not or scarcely raised. Sori medial to ments 1 .3-1.5 cm broad, pruinose abaxially, scales
supramedial. essentially lacking. Ultimate segments moderately
revolute, larger ones ca. 3 times longer than broad.
In elfin forests, clearings, and along roadsides, Sori supramedial.
1770-3400 m, Piura to Amazonas, south to Cuzco
and Puno. Thus far represented only by the type collection,
Costa Rica & Panama; Colombia; Venezuela; from Puno. This species appears to differ from
Ecuador; Peru; Brazil; Bolivia. Gleichenia revoluta only in the lack of scales on
This species is locally abundant in middle to the laminar surface and in the tuberculate to mur-
upper elevations and is sometimes confused with icate primary axis. However, the specimens were
Gleichenia bifida, probably because of the often taken from a plant apparently quite advanced in
copious indument borne on the abaxial surfaces. age, for few sporangia remain on the segments,
These scales, especially on the veins, often become and it is possible that any scales once borne on
filiform and even grade into stellate trichomes; the axes may have fallen away as well. Density of
however, the surfaces are never truly tomentose, indument varies within some species of Gleichen-
as in G. bifida. Also, penultimate segments of the ia, depending upon maturity. Furthermore, tu-
latter are muchbroader, with linear ultimate seg- berculate primary axes are not uncommon in the
ments, and the sori crowd the midrib, whereas in genus, although never so conspicuously so as in
G. revoluta sori are medial to supramedial, most G. tuberculata. The species is maintained as dis-
of them quite remote from the midrib. Further- tinct here, but with reservations.
more, G. bifida grows at lower elevations. It is
likely that G. revoluta occurs also in Costa Rica
and Panama [as G. costaricensis (Underw.) C. Chr.]
and in Bolivia [as G. boliviensis(Maxon & Morton) 8. Gleichenia nitidula Rosenst., Repert. Spec. Nov.
Lell.]. Probably G. subandina Sodiro and G. hy-
Regni Veg. 10: 275. 1912. TYPE: Costa Rica,
poleuca Sodiro of Ecuador are also synonymous San Carlos, Brade & Brade 503 (holotype, not
with G. revoluta.
located; isotypes, NY, s).
Playapampa, in sphagnum montana, Macbride 4508 (F, sparingly dentate or short-setose. Larger penulti-
GH, NY, us). Cuzco: Cordillera Vilcabamba, 23 km NE mate segments 1.4-1.8 cm broad, not or scarcely
from Hacienda Luisiana, Dudley 11133 (GH, us). Puno:
pruinose abaxially, scales lacking. Ultimate seg-
Prov. Sandia, near Limbani, Metcalf 30539 (MO, us;
ments slightly revolute, larger ones ca. 3 times
atypical in narrower segments and darker, narrower
scales). longer than broad.
44 FIELDIANA: BOTANY
Ama/onas: Prov. Bagua, roadside, 37 km NE of Chi- us
lotype, K; isotypes, BM, K; frag., us!; photo,
Barbour 4485 (MO), 4486 (USM). San Martin: Tar-
riaco,
ofK).
apoto-Yurimaguas Hwy., km 39, McDaniel 14209 (GH,
MO). Tarapoto, Carretera Tarapoto-Yurimaguas, Ri-
machi 5163, 5251 (MO, NY). Huanuco: Monzon, con- Mertensia longipinnata (Hooker) Klotzsch, Linnaea
fluencia con Huallaga, cerca de Tingo Maria, Ferreyra 18: 537. 1844.
10047 (GH). Dicranopteris longipinnata (Hooker) Maxon, Contr.
U.S. Nail. Herb. 24:48. 1922.
Sticherus longipinnatus (Hooker) Nakai, Bull. Nat. Sci.
Mus. 29: 21. 1950.
11. Gleichenia peruviana (Maxon) 1 ell.. Amer.
Fern J. 74: 57. 1984.
Leaves pseudodichotomously forked, occasion-
ally bearing foliaceous appendages within the ax-
Dicranopteris peruviana Maxon, Amer. Fern J. 33: ils. Pinnae commonly 1 -forked. Axillary scales
133. 1943. TYPE: Peru, Huanuco, Playapampa,
ca. 2700 m, Macbride 4510 (holotype, F!; isotype,
linear to filiform, 2-3 mm
long, lustrous reddish
us!). brown to deep orange, not or rarely tightly im-
bricate, the margins with dark, rigid, ascending
Leaves pseudodichotomously forked, with fo- setae. Penultimate segments (larger ones) 4-9 cm
liaceous appendages often borne within some of broad, costae on abaxial side amply to abundantly
the more proximal axils. Pinnae 1-3-forked. Ax- scaly, the scales 1.5-2 mm
long, lustrous reddish
illary scales ovate or lance-ovate, attenuate, 0.5- brown, setose and filiform or substellate, on adax-
1.5 mm long, castaneous to reddish brown, closely ialside often appressed filiform-scaly. Ultimate
imbricate, their margins amply ciliate, the cilia lax, segments approximate, contiguous at their bases
short and whitish to pale orange. Penultimate seg- (very rarely some proximal ones discrete and
ments (larger ones) 2.5^4(-5) cm broad, costae on somewhat apart), plane to slightly revolute, not or
abaxial side amply to copiously scaly, the scales ones 25-45
slightly pruinose, larger mm
long and
lustrous castaneous to reddish brown, ovate to lan- 2.5-3.5 mmbroad, abaxially bearing filiform or
ceolate, acuminate or attenuate, commonly less substellate scales on midrib and veins and often
than 1 mm long, their margins short-ciliate, costae some pale, delicate trichomes at the very margin.
on adaxial side sparsely to amply provided with Veins not or slightly prominulous. Sori mostly
tortuous trichomes or filiform scales. Ultimate seg- raised.
ments crowded, contiguous at their bases, slightly
to moderately revolute, not or slightly pruinose, on slopes and ridges, 300-1600 m
In forests,
larger ones 2-24 mm
long and 1 .5-2.5
1 broad mm (reported on Dudley 10448 label as also seen at
(beyond the dilated base), abaxially the midribs 3300 m, but probably mistaken for G. peruviana),
and (usually) veins minutely scaly, ciliate- or se- Amazonas, Loreto, Pasco, Huanuco, Cuzco.
tose-scaly, the scales often becoming filiform or Surinam; Peru; Brazil.
grading into stellate, whitish trichomes. Veins often Despite obvious differences, this and Gleichenia
raised. Sori mostly inframedial. rubiginosa share a distinctive type of laminar scale
not found in other species in Peru. The scales along
In thickets, open forests and ravines, often on the costae of penultimate divisions, abaxially, are
exposed rocky ridges, 2000-3400 m, La Libertad, typically long and narrow, the marginal cell walls
Huanuco, Pasco, Cuzco. deep reddish brown and bearing rigid dark setae.
Apparently confined to Peru, Cordilleras Cen- Along midrib and veins, these grade into filiform
tral and Oriental. scales often only two to three cells wide (the setae
then much longer than the width of the scale body)
La Libertad: Prov. Pataz, Pampa de Huayno-huincho, and thence into delicate, pluricellular trichomes.
Huaylillas, Ldpez & Sagdstegui 3517 (GH, HUT). Huan- The Costa Rican G. mellifera Christ also bears this
uco: Wet, dense jungle, 12 mi S of Panao, Macbride & kind of laminar scale, and although we have not
Featherstone 2217 Pasco: Prov. Oxapampa, Dist.
(F).
seen the type, it probably belongs here.
Oxapampa, Rio San Alberto, Foster et al. 10306 (F);
Leon 641 (F, GH); D. Smith & Pretel 7594 (F, MO). Cuzco:
Prov. La Convention, on open exposed ridge, Dudley Amazonas: Prov. Bagua, valley of Rio Maranon above
10710 (GH, MO). Cascadas de Mayasi, Wurdack 2049 (F, NY, uc, us).
Huanuco: SW slope of Rio Llullapichis watershed, Dud-
ley 13193 (GH). Ucayali: Prov. Coronel Portillo, Padre
12. Gleichenia longipinnata Hooker, Sp. til. 1: 9. Abad, Chacra de Cesar Vela, J. Schunke 5465, (F, GH,
1844. TYPE: Surinam, Hostmann 238 (ho- MO, NY, us). Pasco: Prov. Oxapampa, Valle de Palcazii,
In wet forests and thickets, 1 100-2850 m, Piura, This, like Gleichenia, is an essentially pantrop-
Huanuco, Pasco, Ucayali, Cuzco, Puno. ical genus. The two are readily distinguished by
Colombia; Venezuela; Ecuador, Peru. the character of their indument: that of Gleichenia
Although easily distinguished from G. longipin- being scales and trichomes, while scales are com-
nata apparently closely related to the
(q.v.), this is pletely lacking in Dicranopteris. Three species of
latter by virtue of the distinctive laminar scales. the latter are recognized in Peru.
b. Pinna-forks obviously branching unequally; accessory leafy segments lacking at the base of each
46 FIELDIANA: BOTANY
fork (not to be confused with stipule-like segments often borne within the forks); spores monolete
3. D. pectinate
a. Leaves 1 -forked, each branch consisting of a single, sessile, pinnate (to distally pinnatisect) pinna;
ultimate segments mostly well-spaced, not contiguous; lamina abaxially with dense, reddish brown
tomentum 2. D. nervosa .
1. Dicranopteris flexuosa (Schrader) Underw., Rio Chinchao, Macbride 5010 (F, NY, us). Ucayali: Prov.
Bull. Torrey Bot. Club 34: 254. 1907. Figure Coronel Portillo, Obeteni, E ridge of basin, Chrostowski
66-13 (uc). Cuzco: Prov. Convention, Choquallo, Var-
lOb-c.
gas 8153 (MO, uc). Puno: Prov. Sandia, Asalaya, Vargas
14835 (OH).
Mertensia flexuosa Schrader, Gott. Gel. Anz. 863.
1 824. TYPE: Brazil, Maximilian Prinz Neuwied
(i.e., each of the stalks subequal in length and di- Axils bearing a dense tuft of long, lax, reddish
verging at about the same angle), and a pair of brown trichomes and usually a pair of reduced
reduced, accessory, pectinate segments commonly stipule-like appendages.Pinnae (larger ones) 1 5-
produced at the base of each (but rarely the ulti- 40 long, 5-12 cm broad, the axes and laminar
cm
mate) fork. Axils bearing a tuft of stout, rigid, surfaces abaxially covered with a dense reddish
castaneous trichomes and a pair of reduced sti- brown tomentum (this sometimes gray to whitish
pule-like segments. Pinnae and their branches long- in age). Ultimate segments well-spaced, not con-
stalked, the axes and laminar surfaces glabrous, tiguous at base, to 60 mm
long and 3.5 mm
broad,
the forked, penultimate segments sessile, to 30 cm coriaceous, glaucous abaxially, the margins strong-
long and 6 cm broad. Ultimate segments contig- ly revolute. Spores trilete.
uous at base, to 30 mm
long and 4 broad,mm
coriaceous, glaucous abaxially, the margins strong- Thus far represented in Peru by a single collection:
Puno: Region of Rio Inambari, trail from Aricoma Pass
ly revolute. Spores trilete.
to Santo Domingo, 1800 m, McCarroll 125 (MICH).
San Martin: NW
of San Martin, Rioja, Rio Negro, question. Further study of the species complex is
Soukup 5156 (GH). Huanuco: Vilcabamba, hacienda on needed.
48 FIELDIANA: BOTANY
3. Dicranopteris pectinate (Willd). Underw., Bull. Tingo Maria to Pucallpa, Ridoutt s.n. (GH). Cuzco: Prov.
Torrey Hot. Club 34: 260. 1907. Figure lOa. Paucartambo, near Asuncion, West 7121 (MICH p.p., MO,
vcp.p.).
5 mm broad, chartaceous, glaucous abaxially, the bescent variant of D. pectinata (q.v.), although Hi-
margins scarcely to moderately revolute. Spores eronymus certainly should have mentioned the
monolete. distinctive branching pattern of the latter if this
were the case. Pending examination of the type, it
a. Indusium bivalvate, the valves '/2 or more the length of the indusium (rarely less); receptacle not or
and slightly exserted; venation anadromous; ultimate segments sometimes serrate, lacking false
rarely
veins I. Hymenophyllum
a. Indusium tubular and (and then the valves not more than '/a the
entire, or bilabiate, rarely bivalvate
length of the indusium; receptacle commonly exserted at maturity, often strongly so; venation anad-
romous or catadromous; ultimate segments not serrate, with false veins sometimes present
. . II. Trichomanes
I. Hymenophyllum
Hymenophyllum subg. Sphaerocionium (Presl) C. Chr.,
Index fil. Suppl. 3: 5. 1934.
Hymenophyilum Sm., Mem. Acad. Roy. Sci. (Tur- Mecodium(Cope\.)CopeL, Philipp. J. Sci. 67: 17. 1938.
in) 5: 418. 1793. LECTOTYPE (designated Hymenophyllum subg. Mecodium Copel., Philipp. J.
by Presl, Hymenophyllaceae 31. 1843): Hy- Sci. 64: 93. 1937. TYPE: Hymenophyllum po-
lyanthos (Sw.) Sw.
menophyllum tunbridgense (L.) Sm. Figure 11.
menophyllum dicranotrichum (Presl) Sadeb. bearing scattered trichomes and small, delicate
Hymenophyllum subg. Leptocionium (Presl) Christ, roots. Leaves monomorphic, petiolate, commonly
Farnkr. Erde 20. 1897.
1-20(-50) cm long or, in H. speciosum, to nearly
Sphaerocionium Presl, Hymenophyllaceae 33. 1843.
TYPE: Sphaerocionium hirsutum (L.) Presl (Tri- 2 m long. Lamina simple and entire (outside Peru)
chomanes hirsutum L.) = Hymenophyllum hir- or pinnatifid to decompound, glabrous or sparsely
sutum (L.) Sw. to densely pubescent. Ultimate segments entire to
50 FIELDIANA: BOTANY
FIG. 1
Hymenophyllum fucoides \ar.fucoides: a, habit; b, pinna with son. Hymenophyllum polyanthos: c, pinna
1.
with Hymenophyllum crispum: d, pinna with sori. (a from Seller 295, El Salvador, F, b from Plowman 6074, F,
sori.
c from Schunke 462, F, d from Wurdack 1510, F.)
menophyllum, Mecodium, and Leptocionium. A toalatum they are small and scattered, and in H.
good revision of the latter was done by Morton plumosum they are not at all foliose, but instead
( 1 947) under the name Sphaerocionium, but this are low and inconspicuous, and usually obscured
and others still require careful study, and some of by the dense, matted tomentum covering the en-
the species concepts are only provisionally treated tirelamina. In H. mirificum they could be mis-
below. Although there is no attempt here at a for- taken for trichomes, since many of them, although
mal subgeneric classification, the species are so four to six cells wide at the base, abruptly become
designated in the key and their descriptions follow uniseriate and grade to a filiform tip.
in natural order.
To facilitate use of the key to species, some ex-
planation of the diagnostic characters may be help- Reference
ful here, especially in subg. Leptocionium. Caution
is recommended when using the couplets that sep- MORTON, C. V. 1947. The American species of
arate those species having trichomes borne only Hymenophyllum sect. Sphaerocionium. Contr.
on veins and margins of segments (Morton's sub- U.S. Natl. Herb., 29: 139-201.
a. Ultimate segments conspicuously serrate, glabrous, or rarely the serrations tipped by a short, pluri-
trichome [subg. Hymenophyllum]
cellular b
b. Costae and veins lacking lamellae c
c. Pinnae subequilateral at least beyond the base; sori (most of them) borne in the same plane as
the lamina, only occasionally arcuate; petiole usually more than 0.3 mm
in diameter
1 H. fucoides .
c. Pinnae dimidiate secondary segments all borne on acroscopic side); sori (most of them)
(i.e.,
strongly arcuate, thus arranged nearly perpendicular to the plane of the lamina; petiole less
than 0.3 mmin diameter 2. H. peltatum
b. Costae or veins (at least adaxially) bearing few to many lamellae not in the plane of the lamina,
these several to many cells wide at base, becoming uniseriate toward apex d
d. Petiole nonalate; rachis flexuous, nonalate, or slightly so to marginate distally; lamellae scattered
and inconspicuous 3. H. mirificum
d. Petiole broadly alate, often nearly to base; rachis straight, broadly alate throughout; lamellae
frequent and conspicuous 4. H. lamellatum
a. Ultimate segments entire, sometimes undulate to crispate, but never serrate, or rarely minutely
denticulate and then the teeth tipped by unicellular or stellate trichomes e
52 FIELDIANA: BOTANY
e. Lamina glabrous; ultimate segments with margins entire (although often crispate) [subg. Mecodium]
f
f. Leaves 1.5-4(-5) cm long; pinnae 3-6 pairs; rachis strongly flexuous to fractiflex
5. H. apiculatum
f. Leaves (mature ones) 6-30 cm long; pinnae 8-30 pairs; rachis straight or slightly (rarely strongly)
flexuous g
g. Mature indusia ovoid, or rhomboid, most of them somewhat to markedly longer
elliptical,
than broad, apex obtuse to acute, base narrowly to broadly cuneate (occasionally rounded)
and slightly to deeply immersed in the segment tissue; soriferous segment tips not or slightly
constricted, as broad as (or slightly narrower than) mature indusia h
h. Mature indusia broad- or narrow-ovoid, broadest toward the rounded or broadly cuneate
base, slightly immersed in the segment tissue; receptacle filiform to narrow-cylindrical,
short, not or rarely exserted i
i. Petiole marginate or alate near apex or throughout (wings sometimes partly deciduous);
rachis alate throughout; leaves commonly less than 15 cm long; growing at 100-
2000(-2400) m 6. H. polyanthos
i. Petiole neither alate nor marginate; rachis commonly nonalate at base, at least on one
side; leaves (mature ones) commonly 14-30 cm long; growing at 2100-3150 m . . . .
7. H. mathewsii
h. Mature indusia rhomboid, or conical, broadest at or beyond the middle, the base
elliptic,
narrow-cuneate, deeply (often halfway) immersed in the segment tissue; receptacle fili-
form, long, often exserted 8. H. trichomanoides
g. Mature indusia subglobose, circular in outline to broader than long, the apex rounded, the
base rounded or subtruncate, not or scarcely immersed in the segment tissue; soriferous
segment tips somewhat to greatly constricted, narrower than the mature indusia (usually so
much so that indusia appear pedicellate) j
j. Sporangia (5-)6-20 per sorus; petiole marginate or alate, at least distally (wing sometimes
deciduous); rachis alate throughout k
k. Indusium apex erose; 1 -several elongated pinnae intermixed among normal ones . .
9. H. ferax
k. Indusium apex entire; elongated pinnae rare or lacking among normal ones
10. H. myriocarpum
j. Sporangia l-4(-5) per sorus; petiole neither marginate nor alate; rachis commonly non-
alate toward base (or discontinuously alate on either side due to decurrent pinna bases)
1 1 . H. u lid u a
I 1 11 ii i
e. Lamina pubescent (at least on margins or veins), often densely so; ultimate segments entire or
rarely minutely denticulate [subg. Leptocionium} 1
Q
q. Marginal trichomes mostly forked at base or 2-forked (a few simple or stellate),
petiole trichomes simple or forked; leaves determinate 14. H. elegans
base 1 7. H. crispum
m. Petiole (of mature leaves) stout, 0.4-1 .5 in diameter mm s
s. Rachis conspicuously alate throughout; petiole usually alate just below lamina; trichomes
lacking on adaxial side of lamina t
t. Ultimate segments and rachis wing slightly to strongly undulate; stellate trichomes
lacking on petiole 1 8. H. valvatum
t. Ultimate segments and rachis wing not or scarcely undulate; stellate trichomes fre-
quent on petiole among the simple and forked ones 19. H. microcarpum
s. Rachis nonalate, at least in the proximal portion; petiole nonalate; trichomes present
on axes and veins both abaxially and adaxially 20. H. ruizianum
1. Trichomes (few to many) borne on the segment surface between the veins as well as on the
veins and margins u
u. Veins lacking lamellae v
v. Petiole slender, 0.2-0.3 mm
in diameter (or sometimes to 0.4 in H. amabile) mm w . . .
visible y
y. Rachis broadly alate throughout; leaves determinate; petiole trichomes simple
to forked and often stellate 22. H. fragile
Rachis nonalate (sometimes alate distally); leaves indeterminate; petiole tri-
y.
chomes simple or forked, not or rarely stellate 23. H. elegantulum
x. Lamina densely tomentose, the veins and tissue often obscured
24. H. amabile
v. Petiole (of mature leaves) stout, 0.4-1.0 mm
in diameter z
z. Lamina densely tomentose, the veins and tissue mostly obscured by the tomentum;
lanceolate, 8-15 cm broad, not or slightly reduced at base .... 27. H. lindenii
bb. Petiole 2-7 cm long, 0.4-0.6(-0.7) mm in diameter; lamina linear, 3-8 cm broad,
and gradually reduced toward base
often strongly 28. H. plumieri
u. Veins bearing conspicuous lamellae (or, in H. plumosum, the outgrowths low and incon-
spicuous, these and the veins obscured by dense tomentum) cc
cc. Rachis nonalate nearly or wholly throughout dd
dd. Lamellae minute and inconspicuous, these and the veins obscured by dense to-
mentum 29. H. plumosum
dd. Lamellae broad and conspicuous, especially abaxially (if surface densely tomentose,
most lamellae still usually visible among the trichomes) ee
ee. Trichomes frequent, but not forming a dense cover; lamellae conspicuous abax-
54 FIELDIANA: BOTANY
ee. Trichomes densely covering (sometimes obscuring) tissue and veins; lamellae
sometimes lacking adaxially 31. H. tomentosum
cc. Rachis alate nearly or wholly throughout ff
ff. Petiole rather stout, 0.4-0.7 mm in diameter; rachis strongly and regularly alate, at
least above the base; leaves 9-60 cm long; pinnae (larger ones) with 6-24 segments
gg
gg. Pinnae mostly narrow-triangular, narrowly acute to attenuate, larger ones 20-
45 mm long, with (6-)7-l 2 pairs of segments hh
hh. Lamellae on abaxial side mostly consisting of scattered flanges, about as
broad as long, lacking adaxially 32. H. lobatoalatum
hh. Lamellae on abaxial side abundant, conspicuous, elongate, often nearly the
length of the vein, sometimes less abundant and conspicuous adaxially . .
H. pyramidatum 33.
gg. Pinnae oblong to broadly triangular, obtuse or subacute, larger ones 6-20 mm
long, with 3-6(-8) pairs of segments 34. H. verecundum
ff. Petiole slender, 0. 1 0-0. 1 5 mm in diameter; rachis weakly and irregularly alate due
to the decurrent pinna bases; leaves less than 7 cm long; pinnae with 2-4 segments
35. H. tarapotense
1. Hymenophyllum fucoides (Sw.) Sw., J. Hot. (BM) of Hymenophyllum cristatum Hooker Grev. &
(Schrader) 1800(2): 99. 1802. This sheet contains two specimens of//, fucoides,
while pinned to it is an exact copy of/. 148, Icon,
Leaves 4-25(-30) cm long, 1 .5-8 cm broad. Pet- 1829, the illustration of//, cristatum, complete
fil.
iole 0.3-0.8 mm in diameter, nonalate or margin- with globose receptacle and conspicuous lamellae
ate to alate distally, glabrous to sparsely or mod- on the veins, which is an Ecuadorean species of a
erately pubescent with catenate trichomes. Lamina different section, Buesia (Morton) Morton. This
broadly or narrowly oblong, not or rather abruptly led to another error, for the subsequent illustration
reduced at base, 3-4-pinnatisect. Rachis broadly H. fucoides in Hooker's "Century of Ferns" (1854)
to narrowly alate throughout, or nonalate just at was evidently produced from this mixed sheet,
the base, sparsely to moderately provided with thus misleading later investigators about some of
flexuous, catenate trichomes. Pinnae 3-20(-25) the principal diagnostic features (for a full discus-
pairs, adnate, orproximal 1-2 pairs short-stalked, sion see Stolze, Amer. Fern J. 77: 137-140. 1987).
subequilateral except at base, there truncate ac- A second source of confusion has been the high-
roscopically, cuneate basiscopically, with 2-8 pairs ly variable nature of the species, which has
of pinnatisect to 2-pinnatisect segments. Sori 1-8 prompted the recognition of a number of related
per pinna, not or scarcely immersed in the segment taxa at various levels over the years. Morpholog-
tissue, most of them borne in the same plane as ical features are rather constant in the West Indian
the lamina, not or only occasionally arcuate. In- specimens, where leaves are quite small and com-
dusia narrowly to broadly oblong or elliptic, the pact; but variability increases in Central America,
apex obtuse to subacute and entire to laciniate, where some leaves are larger and more elongate,
receptacle narrow-cylindrical or fusiform to ovoid, and apices of indusia are deeply dentate. Even
not exserted or occasionally slightly so. greater differences are found in South American
plants, especially in Peru, where variability is at
The species occurs in the West Indies; southern its peak. It is likely that part of this may be due to
Mexico to Panama; Venezuela and Colombia to the effects of ecological niches and altitudinal levels,
Brazil and Bolivia. but this kind of analysis must wait for a much-
This species has been misunderstood, both in needed revision of this part of the genus.
itsnomenclature and taxonomy, since it was first For purposes of this treatment //. fucoides has
recognized by Swartz in 1788. The source of the been segregated into four varieties, a key to which
greatest confusion is a supposed type collection is provided here.
a. Rachis marginate to narrow-alate, or if broadly alate then alae essentially plane, ultimate segments
plane b
b. Apex of indusium entire, erose or denticulate c
c. Rachis narrow-alate, or sometimes nonalate at base; petiole nonalate, or occasionally marginate
near lamina base; basal pinnae often short-stalked; sori rarely more than 3 to a pinna ......
la. var. fucoides
c. Rachis broadly alate throughout, each wing broader than the rachis; petiole usually alate distally,
at least near lamina base; basal and other pinnae adnate; sori often 5-8 to a pinna
Ib. var. calodictyon
b. Apex of indusium spinulose-dentate to laciniate var. pedicellatum
Ic.
a. Rachis broadly alate its entire length, alae and/or ultimate segments often undulate and their margins
discontinuously conduplicate Id. var. chachapoyense
la. Hymenophyllum fucoides var. fucoides. Fig- Amazonas: Trail east of La Peca in Serrania de Bagua,
ure lla-b. Gentry et al. 23034 (F, MO, us). San Martin: Venceremos,
near Amazonas border, Gentry et al. 45402 (MO). Huan-
uco: Cani, NE of Mito, Bryan 206, 384 (F); Macbride
Trichomanes fucoides Sw., Prodr. 136. 1788. TYPE: 3397 (B, F, us). Cuzco: Cerro Chuyapi, Bues ASS (GH,
Jamaica, Swartz (holotype, s; isotypes, B, BM; us), A56 (us). Puno: Sandia, Weberbauer 713 (B).
phOtOS, F & GH Of B, US of BM).
Leptocionium fucoides (Sw.) Presl, Hymenophyllaceae
27. 1843.
Ib. Hymenophyllum fucoides var. calodictyon
Meringium fucoides (Sw.) Copel., Philipp. J. Sci. 67:
45. 1938. (Bosch) Stolze, comb. & stat. nov.
Petiole (0.3-)0.4 0.8 mm in diameter, nonalate, Hymenophyllum calodictyon Bosch, Ned. Kruidk.
Arch. 5(3): 172. 1863. TYPE: Peru, Ruiz 84 (ho-
occasionally marginate near apex. Rachis narrow-
lotype, B!).
alate or marginate, at least distally, alae plane.
Pinnae adnate, or 1-2 proximal pairs short-stalked,
ultimate segments plane. Sori l-3(-4) pairs to a
Petiole 0.5-0.8 mm in diameter, conspicuously
alate to marginate distally. Rachis broadly alate
pinna, apex of indusia entire to denticulate, rarely
throughout, each of the alae plane and much wider
dentate.
than the rachis. Pinnae all adnate, with ultimate
segments plane. Sori ( l-)2-8 pairs to a pinna, apex
On
trunks and branches of trees in wet forest,
of indusia entire.
rarely on wet rock walls, 1 700-3300 m, Amazonas,
San Martin, Huanuco to Puno.
On trees in wet forests, Cajamarca and Ama-
West Indies; southern Mexico to Panama; Ven-
zonas south to Cuzco, 2100-3300 m.
ezuela and Colombia to Brazil and Bolivia.
Ecuador; Peru.
The most diminutive of these ferns scarcely dif-
In addition to the characters mentioned in the
fer from H. tunbridgense (L.) J. Sm., the type of
key, var. calodictyon commonly may be distin-
the genus, which is said to differ in its smaller
guished from the other varieties by the broader
leaves and in its strongly arched sori. The latter
segments, membranaceous tissue, and the lighter,
species supposedly occurs in the West Indies and
yellow-green color.
in Europe and Africa, but it may be only another
56 FIELDIANA: BOTANY
Ic. Hymenophyllum fucoides var. pedicellatum In undulate to crispate rachis wings and ulti-
its
(Klotzsch) Hieron., Bot. Jahrb. Syst. 34: 435. mate segments,var. chachapoyense resembles Hy-
1904. menophyllum tortuosum Hooker & Grev. of Chile;
however, alae and segments of the latter are much
Hymenophyllum pedicellatum Klotzsch, Linnaea 20: more strongly crispate and are setose-serrate as
439. 1847. TYPE: Venezuela, Merida, Moritz 346 well.
(holotype, B; isotypes, BR, us in part!).
Leptocionium pedicellatum (Klotzsch) Fourn., Bull.
Soc. Bot. France 19: 249. 1872.
Amazonas: Prov. Chachapoyas, Cerros de Calla Calla,
26 km above Leimebamba on road to Balsas, Hutchison
& Wright 6987 in part (F, GH, uc). Huanuco: Tambo de
Petiole 0.3-0.5 mm in diameter, nonalate. Vaca, Macbride 4460 (F, us).
Rachis broadly to narrowly alate, alae plane. Pin-
nae adnate, or basal pair short-stalked, ultimate
segments plane. Sori 1-3 pairs to a pinna, apex of
indusia laciniate to spinulose-dentate.
2. Hymenophyllum peltatum (Poir.) Desv., Mem.
Soc. Linn. Paris 6: 333. 1827.
diagnostic character above the level of species. ises in H. cristatum are obviously alate through-
out, lamellae are numerous and conspicuous, and
Junin: Prov. Huancayo, Dist. Huancayo, Huaytapal- indusia are deeply dentate to laciniate at apex. For
lana. Sounders 1 167 (GH). Cuzco: "Ccarcco" (Ccorca?), further discussion about the confused identity of
Biies 1395 (us). ?Madre de Dios: Pinasniocj, "Panticalla
H. cristatum, see H. fucoides.
Pass" (Pantiacolla?), Cook & Gilbert 1868, 1870 (us).
Puno: Prov. Carabaya, Ayapata, Vargas 10705 (GH).
Cuzco: Prov. La Convencion, Vilcabamba, Biies 1592,
1593, 2104 (us). Prov. Paucartambo, Acanacu, Vargas
29 (GH); West 7144 (uc, us). Prov. Urubamba, Puyu-
3. Hymenophyllum mirificum Morton, Bot. Gaz. pata-Yuncapata, Vargas 2922 (MO, us).
(Crawfordsville) 93: 338. 1932. TYPE: Peru,
Cuzco, Prov. La Convencion, Vilcabamba,
Biies 1600 (holotype, us!; photo, F).
4. Hymenophyllum lamella turn Stolze, sp. nov.
nonalate or, rarely, very narrowly so or marginate adnatae, l-2.5(-3) cm longae, costis et venis lamellatis
praeditae. Segmenta saepe ultima undulata. marginibus
at apex, sparsely provided with brownish, flex- serratis.Indusia sphaeroidea vel ovoidea, brevipedicel-
uous, catenate trichomes. Lamina broadly or nar- lata, integris, receptaculis subglobosis vel
marginibus
rowly oblong, not or abruptly reduced at base, ovoideis.
58 FIELDIANA: BOTANY
Some of the specimens cited below were earlier fewer pinnae, but the character of its sori is dif-
suspected of being new species. On the sheets of ferent. The narrowly cuneate base of the indusium
Bites A 17 & A20 (us) are Morton's penciled re- is deeply immersed in the segment tip, whereas in
marks, "sect. Buesia, n. sp.?"; and Lechler 2568 H. polyanthos the indusium base is broadly cu-
(B) is annotated by Mettenius, "//. latipes Mett." neate to obtuse and only slightly immersed in the
Apparently Mettenius intended to publish, but a tissue.
Mecodium dendritis (Rosenst.) Copel., Philipp. J. Sci. confluence of Rio Huallaga, Mexia 8282 (holo-
67: 26. 1938. type, uc!; isotypes, B!, F!, MO!).
Mecodium apiculatum (Kuhn) Vareschi, Flora Ven- Hymenophyllum mexiae (Copel.) Morton, Contr. U.S.
ezuela, Caracas 1: 198. 1969. Natl. Herb. 38: 173. 1968.
Leaves 1.5-4(-5) cm long, 0.8-1.8 cm broad. Leaves (6-)8-15 cm long, 2-5(-6) cm broad.
Petiole 0.3-1 .5 cm long, 0. 1-0.25 mm in diameter, Petiole 0.5-5 cm long, 0.3-0.6 mm in diameter,
marginate to narrow-alate, glabrous or with a few alate near theapex or throughout, glabrous. Lam-
scattered trichomes, especially at very base. Lam- ina ovate or elliptic, scarcely reduced at base (or
ina ovate or deltoid-ovate, not or scarcely reduced proximal 1-2 pairs of pinnae slightly reduced), 3-
at base, 2-3-pinnatifid, glabrous. Rachis glabrous, 4-pinnatisect, glabrous. Rachis glabrous, straight
strongly flexuous to fractiflex, alate throughout. or slightly flexuous, alate throughout. Pinnae 8-
Pinnae 3-6 pairs, adnate, larger ones 0.5-1.2 cm 14 pairs, essentially adnate, but not strongly over-
long, bearing 14 pairs of secondary segments. Ul- lapping the rachis, larger ones 1 -4 cm long, bearing
timate segments entire, plane, or occasionally the 2-many secondary segments. Ultimate segments
margins involute or conduplicate. Sori 1-4 to a entire, plane or occasionally a few with condupli-
pinna, usually confined to apical portion of lam- cate segment margins, larger ones commonly more
ina. Indusia rhomboid or ovoid, apex entire and than 1 mm
broad, the free portion relatively short,
obtuse to subacute, with broadly cuneate base, usually about twice as long as broad. Sori 2-many
about halfway immersed in the segment tissue, per pinna. Indusia relatively small, usually about
receptacle filiform, not exserted, bearing 6-12 spo- as broad as the segment apex, ovoid, elliptic, or
rangia. lanceolate, broadest toward the base, apex obtuse
or subacute, the broadly cuneate base slightly im-
on branches and trunks of trees
In dense forests mersed in the segment tissue, margins entire, re-
and on fallen logs, 100-1 500(-2000) m, San Mar- ceptacle filiform to narrowly cylindrical or fusi-
tin, Loreto, Huanuco, Pasco, Cuzco. form, not exserted, bearing 6-15 sporangia.
Venezuela; Colombia; Peru; Bolivia.
This would appear to be simply a reduced form In dense forests, on fallen logs, and on trunks
of Hymenophyllum polyanthos; however, it not of trees and tree ferns, 1 00-2000(-2400) m, Ama-
only has smaller leaves, more delicate petioles, and zonas, Loreto, San Martin south to Puno.
pedicellate, for the apices of ultimate segments are glabrous. Rachis glabrous, alate throughout except
strongly constricted; furthermore, mature indusia usually nonalate at base, or there narrow-alate on
are subglobose, or often noticeably broader than only one side. Pinnae 14-18 pairs, adnate and often
long. This contrasts with the essentially ovoid in- overlapping the rachis, or basal ones short-stalked,
dusia of//, polyanthos et al., commonly somewhat larger ones (1.5-)2-3.5 cm long, bearing 4-8 pairs
longer than broad and slightly to strongly im- of secondary segments. Ultimate segments entire,
mersed in the segment tips, which are not or plane or occasionally undulate, or a few with mar-
scarcely constricted. While this character is prob- gins conduplicate. Sori 4-many per pinna. Indusia
ably the most reliable one in subg. Mecodium in relatively large, usually slightly broader than the
Peru, unfortunately even it is not 1 00% constant, segment apex, ovoid, apex obtuse to subacute, the
for on occasional specimens indusia may vary broadly cuneate base slightly immersed in the seg-
somewhat from one shape to another. Conse- ment tissue, margins entire, receptacle short and
quently, most species in the subgenus must be sep- filiform, bearing 6-12 sporangia.
arated by suites of characters. Ultimately, taxo-
nomic problems will be solved only by thorough Cloud forests, commonly on trunks and branch-
study of species groups throughout their entire es of trees, 2 1 00-3 1 50 m, Huanuco, Pasco, Cuzco.
range. Ecuador; Peru.
Two other characters are often helpful in sep- Hutchison 1751 (cited below) has delicate, di-
arating H. polyanthos (at least in Peru) from H. minutive leaves only 6-8 cm long; this depauper-
myriocarpum particularly useful where sterile ate condition probably results from the unusual
specimens are involved. Pinnae of H. myriocar- habitat, "in cave." This species is very closely re-
pum commonly have basal segments strongly Hymenophyllum polyanthos (q.v.) and may
lated to
overlapping the rachis. In H. polyanthos pinnae not merit distinction at the species level. Normal
are adnate and basal segments often crowd or touch specimens of//, mathewsii tend to have larger sori,
the rachis, but rarely do some of them overlap. in relation to the segment apex, and longer, pen-
Also, H. polyanthos occurs at lower elevations in dent leaves, with no trace of tissue along the petiole
Peru, 100-2000(-2400) m; whereas H. myriocar- and, commonly, none between the proximal two
pum is found at ( 1 700-)2000-4200 m. pinnae. Leaves of H. polyanthos are smaller and
60 FIELDIANA: BOTANY
more compact, erect or arching, with rachis dis- guished by the narrower ultimate segments. Larger
tinctly alate. The petiole is marginate or alate ones are commonly 0.6-0.9 mm
broad and linear.
throughout, or at least distally. Often the wing is Other species of subg. Mecodium have relatively
deciduous upon drying, but traces can usually be broader ultimate segments, most of them over 1
found by close scrutiny. mm broad, or if not, then only about twice as long
as broad. This is a rather reliable way to identify
Huanuco: Prov. Huanuco, Carpish, Plowman 6074A sterile specimens.
(F).Pasco: Prov. Oxapampa, San Alberto, Yanachaga, Hymenophyllum trichomanoides differs sharply
van der Werffel al. 8437 (MO). Cuzco: Huadquina, Bues
from all other species of subg. Mecodium in Peru
1346 (us). Prov. La Convention, Cochapata, Valle de
San Miguel, Bues 2176 (us). Prov. Urubamba, summit by the narrowly cuneate sori which are deeply im-
of Huayna Picchu, Hutchison 1751 (atypical) (F, uc). mersed in the segment tissue. Indusia are broadest
at or beyond the middle and, at least in Peru, are
pinna. Indusia relatively large, subrotund, circular Leaves 7-28 cm long, 2.5-5(-6) cm broad. Pet-
in outline or broader than long, apex erose, base iole 1.5-8(-10) cm long, 0.3-0.7 mm in diameter,
rounded, not immersed in the greatly constricted marginate to alate throughout or at least distally
segment apex, receptacle short, filiform, bearing (wings sometimes deciduous), essentially gla-
7-12 sporangia. brous. Lamina lanceolate to ovate, elliptic, or ob-
lanceolate, scarcely to strongly reduced at base, 2-
On trees in wet forests, 2400-3300 m, Cuzco. 3-pinnatisect, glabrous. Rachis glabrous, straight
Venezuela; Colombia to Peru. or slightly flexuous, alate throughout, rachis wings
This is most closely related to Hymenophyllum and those of other axes often conduplicate. Pinnae
myriocarpum var. endiviifolium, from which it dif- 8-30 adnate and basal segments strongly
pairs,
fers in the eroseindusia tips and the presence of overlapping the rachis, larger ones 1-4 cm long,
one to several unusually elongate pinnae which bearing 5-10 pairs of secondary segments. Ulti-
are scattered among the normal ones the latter mate segments entire, plane to undulate, often with
a condition similar to that of H. undulatum. A conduplicate margins. Sori 5-many per pinna. In-
remarkably huge specimen, Lechler 2420 (B; not dusia subglobose or broader than long, much
other 2420 at B, F, K: H. elegans), was collected broader than (and not at all immersed in) the
inPeru at an undesignated site. This is apparently strongly constricted segment apex, rounded at base,
a monstrous form of H. ferax. It matches this rounded and entire at apex, receptacle short to
species in every way, including the erose indusia, nearly obsolete, filiform, bearing 8-20 sporangia.
but the leaf is 85 cm long and the petiole nearly
2 mm
in diameter. It is by far the stoutest specimen This, like H. polyanthos (q.v.), is a widespread
of Hymenophyllaceae we have seen and belies the and greatly misunderstood species which is sorely
appellation "filmy fern." in need of detailed study. It seems to consist of
Key to Varieties
a. Lamina elliptic to oblanceolate, broadest at or beyond the middle; several or many proximal pinnae
somewhat to greatly reduced lOa. var. myriocarpum
a. Lamina ovate to lanceolate, broadest near the base; 1 or 2 proximal pinnae (if any) slightly reduced
b
b. Larger leaves 7-1 5 cm long; pinnae commonly patent and strongly imbricate
1 Oa. Hymenophyllum myriocarpum Hooker, var. Leaves (9-) 12-28 cm long. Lamina elliptic, ob-
myriocarpum. long or oblanceolate, broadest at or beyond the
middle, several proximal pinnae somewhat to
greatly reduced. Rachis and ultimate segments
Mecodium myriocarpum (Hooker) Copel., Philipp. J. plane, some of their margins occasional condu-
Sci. 67: 25. 1938. plicate. Sori 10-20 per pinna.
62 FIELDIANA: BOTANY
In wet forests, usually pendent from tree trunks Skoget al. 5157 (us). Cuzco: Valley of Urubamba, Sues
and branches, or on wet, mossy rocks, rarely on A18,A19(us).
the forest floor, (1 700-)2000-3200 m, Cajamarca,
Amazonas, Huanuco to Cuzco.
Mexico to Costa Rica; Venezuela; Colombia to lOc. Hymenophyllum myriocarpum var. endivi-
Bolivia. ifolium (Desv.) Stolze, stat. & comb. nov.
Cajamarca: Prov. Hualgayoc, Hacienda Taulis, Hymenophyllum endiviifolium Desv., Mem. Soc. Linn.
Hutchison & Bismarck 6415 (F, GH [as Lambayeque], Paris 6: 334. 1827. TYPE: "Peruvia" (holotype,
MO, uc, us). Amazonas: Prov. Bongara, WSW
of Po- p; photos, GH, uc, us).
macocha, Wurdack 866 (GH, us). Huanuco: Mito, Bryan Hymenophyllum multiflorum Rosenst., Meded. Rijks-
206a (F). Pasco: Enenas (as Junin), Pichis Trail, Killip & Herb. 19: 5. 1913. TYPE: Bolivia,Comarapa,
Smith 25700 (F, NY, us). Cuzco: Prov. Urubamba, along Herzog 1951 (holotype, B; isotype, us!).
Rio Urubamba near town of Machu Picchu, Tryon & Mecodium multiflorum (Rosenst.) Copel., Philipp. J.
lOb. Hymenophyllum myriocarpum var. nigres- Leaves (12-) 15-28 cm long. Lamina ovate or
cens (Liebm.) Stolze, stat. & comb. nov. ovate-lanceolate. Rachis and ultimate segments
slightly to strongly undulate, their margins often
Hymenophyllum nigrescens Liebm., Kongel. Danske conduplicate. Sori 1 6-many per pinna.
Vidensk. Selsk. Skr., Naturvidensk. Afd. 5,1: 292.
1849. LECTOTYPE (designated by A. R. Smith, On trees and wet banks in deep forest, 2400-
Flora of Chiapas 2: 133. 1981): Mexico, Puebla,
4200 m, Amazonas, Huanuco, Huancavelica,
Chinantla, Liebmann 537 (lectotype, c; isolec-
Cuzco.
totypes, BM, K, P).
Sphaerocionium nigricans KJotzsch, Linnaea 18: 536. Colombia; Ecuador; Peru; Bolivia.
1 844, not Hymenophyllum nigricans Colla, 1836.
SYNTYPES: Peru, Dombey 87 (syntype, u!; iso-
Amazonas: Prov. Chachapoyas, Calla Calla slopes near
syntype, B); Venezuela, Aragua, Colonia Tovar, km 4 5-4 1 8 of Leimebamba-Balsas road, Wurdack 1750
1
Southern Mexico to Costa Rica; Venezuela; Co- ameter, nonalate, essentially glabrous. Lamina lin-
lombia to Bolivia. ear to narrow-elliptic, often strongly and gradually
reduced at base, 2-3-pinnatisect, glabrous. Rachis
Sterile specimens can easily be confused with
H. polyanthos, but the latter occurs at lower ele- glabrous, marginate to crispate-alate, commonly
nonalate toward base, or discontinuously alate on
vations, and laminae are less compact. See dis-
cussion of H. polyanthos for further comments. alternate sides due to the long-decurrent pinna
bases. Pinnae 1
6-many pairs, adnate, commonly
Amazonas: Prov. Chachapoyas, Cerros de Calla Calla less than 2 cm few of them greatly
long, but often a
above Leimebamba, Hutchison & Wright 6987 (in part: elongate and lamina-like, normal ones bearing 3-
F, GH, uc). San Martin: Prov. Rioja, Pedro Ruiz-Moy- 8(-10) pairs of secondary segments. Ultimate seg-
obamba Road, D. Smith 4376 (F, MO). Pasco: Prov. Ox- ments entire, plane, or the margins conduplicate
apampa, Abra los Mellizos, 4-8 km from Enenas, Skog
in drying, to undulate or markedly crispate. Sori
et al. 5033 (vs). Junin: Huacapistana, Killip & Smith
24164 (us). Ucayali: Plantation Azul, Ridoutt (USM). Prov. 3-many per pinna. Indusia nearly circular, but on
Coronel Portillo (as Loreto), NE of pass at La Divisoria, mature son most of them broader than long, not
monly truncate at base, the margins entire and length. Several species have been described based
plane to crispate, receptacle very short, filiform, on whether the tissue is plane or undulate or
bearing 1^4(-5) sporangia. strongly crispate. However, comparison of indi-
viduals throughout the entire range seems to in-
This species varies greatly as to length and shape dicate that few of these characters are very con-
of lamina and margins of rachis wings and ulti- sistent. Only the conspicuously crispate variants
mate segments. In rocky habitats leaves usually seem to merit recognition, as indicated in the fol-
tend to be less than 10 cm long and narrowly el- lowing key.
liptic, whereas the flaccid, linear leaves pendent
Key to Varieties
a. Ultimate segments and rachis wings plane to undulate, margins often discontinuously conduplicate
in drying; rachis nonalate in the proximal '/3 to /2 or nearly throughout
l
1 la. var. undulatum
a. Ultimate segments and rachis wings markedly crispate, often so strongly as to appear dentate; rachis
commonly alate except near lamina base, or wings sometimes lacking in the proximal 'A or lk of the
rachis . ..lib. var. fendlerianum
1 1 a. H ymenoph y Hum undulatum var. undulatum. cartambo, slopes of Pilahuata, Vargas 4909 (MO, us).
Puno: Sandia, Weberbauer 1293 (B).
64 FIELDIANA: BOTANY
Carabaya, road from San Gaban to Macusani, Maas et 13. Hymenophyilum trichophyllum HBK., Nov.
al. 6109 (MO). Dept. Unknown: Without location, Dom-
gen. sp. 1: 27. 1815.
bey (P).
Key to Varieties
13a. Hymenophyllum trichophyllum var. tricho- Elsewhere, in wet forests, on rocks, clay banks
phyllum. TYPE: Venezuela, Cumana, be- or trees, 2000-3800 m; range the same as the
tween Cocollar and Guardia de San Augustin, species.
Humboldt (holotype, P). The name H. procerum was based on H. pul-
chellum sensu Mett., for which there was no de-
Hymenophyllum procerum Bosch, Ned. Kruidk. Arch. scription. The specimen probably intended as the
4: 409. 1859, nom. nud.
type is the only collection of var. trichophyllum
Sphaerocionium trichophyllum (HBK.) Copel., Phil- thus far known from Peru. It closely matches other
ipp. J. Sci. 67: 32. 1938.
specimens of var. trichophyllum and is cited here.
On shaded rocks, San Martin, Puno, and from Apparently known only from the type collec-
tion. With this probably should be included Hy-
two other collections, localities unspecified, 3000
m. menophyllum sprucei Baker, under which see
Comments.
Elsewhere, in wet forests, on rocks, clay banks
or tree trunks, 800-2400 m; Guatemala; Costa
Rica; Panama; West Indies; Venezuela; Colombia;
Peru; Bolivia; Brazil. 16. Hymenophyllum hirsutum (L.) Sw., J. Bot.
Hymenophyllum tenerrimum was said to differ (Schrader) 1800(2): 99. 1802.
66 FIELDIANA: BOTANY
Trichomanes hirsutum L., Sp. pi. 2: 1098. 1753. TYPE: W side of Cordillera de San Matias, D. Smith 2054 (MO).
based on t. 50b of Plumier, Traite foug. Amer., Cuzco: Prov. La Convencion, Valle San Miguel, San Ja-
apparently representing a plant from the West cinto, Bues 2139 (us). Puno: San Gaban, Lechler 2296
Indies. (B).
Trichomanes ciliatum Sw., Prodr. 136. 1788. TYPE:
Jamaica, Swartz (holotype, s; isotype, B; photo,
us of s).
Hymenophyllum ciliatum (Sw.) Sw., J. Bot. (Schrader) 1 7. Hymenophyllum crispum HBK., Nov. gen. sp.
1800(2): 100. 1802. 1: 26. 1815. TYPE: Venezuela, Silla de Ca-
Sphaerocionium ciliatum (Sw.) Presl, Hymenophyl- racas, Humboldt & Bonpland (holotype, p;
laceae 34. 1843.
photos, GH & us of P). Figure lid.
Sphaerocionium hirsutum (Sw.) Presl, Hymenophyl-
Iaceae34. 1843.
Sphaerocionium crispum (HBK.) Klotzsch, Linnaea
1: 537. 1844.
Leaves determinate, to 12 cm long and 4 cm
broad. Petiole 0.3-2.3 cm long, 0.2-0.3 mm in
Leaves determinate, to 20 cm long and 4 cm
diameter, strongly alate (often nearly to base),
broad. Petiole 0.5-5 cm long, 0.1-0.3 in di- mm
sparsely provided with stellate trichomes. Lamina
ameter, alate near apex, sparsely provided with
2-pinnatisect, sparsely to moderately pubescent,
delicate, simple or 1 -forked trichomes. Lamina
the rachis regularly and conspicuously alate
pinnate to nearly 3-pinnate, sparsely pubescent,
throughout. Pinnae 6-15 pairs, ovate or oblong- the rachis regularly and conspicuously alate
ovate, deeply incised, the 2-5 pairs of segments
throughout, the wing strongly undulate or undu-
plane, simple to bifid, the veins and margins pu-
late-crispate. Pinnae 8-20 pairs, simple (distally)
bescent with stellate or (on the margins) 2-forked
or bifid to nearly 2-pinnate, the 1-8 pairs of seg-
trichomes, but trichomes lacking between veins
ments undulate or crispate, the veins and margins
and margins. Indusia as long as or (usually) longer
sparsely pubescent, marginal trichomes simple or
than broad, somewhat broader than the segment
1 -forked, but trichomes
lacking between veins and
tips.
margins. Indusia as long as or (usually) longer than
broad, Oiten broader than the segment tips.
on branches and trunks of
In dense, wet forests,
trees,500-2000 m, Loreto, Amazonas to Puno. The species occurs in Jamaica, Mexico, Gua-
Mexico to Panama; West Indies; Guianas to temala, Costa Rica, Venezuela, Colombia, Peru,
Venezuela, south to Bolivia and Brazil. Bolivia, and Brazil.
Hymenophyllum crispum might be confused with
Amazonas: Prov. Bagua, E of La Peca, Barbour 2753 H. valvatum, another species in the subgenus with
(MO). San Martin: Tarapoto, Monte Campana, Spruce undulate segments. However, the latter is larger
4698 (K). Loreto: Balsapuerto, lower Rio Huallaga basin,
and coarser with a much thicker petiole, and the
Killip & Smith 28539 (F, GH, NY, us). Huanuco: Prov.
Huanuco, Dist. Churubamba, crest of Santo Toribio, rachis wing and segments are not as strongly un-
Mexia 8145 (B, F p.p., GH, us). Pasco: Prov. Oxapampa, dulate. Two varieties are recognized here.
Key to Varieties
a. Pinnae simple secondary segments 1-2 pairs, simple to bifid
(distally) to trifid or, if pinnatisect, the
1 7a. var. crispum
a. Pinnae (larger ones) 2-pinnatisect, the secondary segments 4-8 pairs, pinnatisect
1 7b. var. bipinnatisectum
1 7a. Hymenophyllum crispum var. crispum. Fig- Rio Lachac, Sues 1813 (GH, us). Prov. Urubamba, Ma-
ure lid. chu Picchu, lit is et al. 1061 (GH, uc, us). Puno: Sandia,
Weberbauer 797 (B).
18. Hymenophyllum valvatum Hooker & Grev., species may be only robust forms of H. hirsutum,
Icon. fil. 2, /. 279. 1831. TYPE: Ecuador, for- which seems to differ chiefly in its smaller lamina
ests of Esmeraldas, on trunks of trees, Jame- and more delicate petiole. The entire complex is
son (holotype, K; possible isotype, K!). in need of detailed study.
Hymenophyllum crispatulum Bosch, Ned. Kruidk. Huanuco: Rio Llullapichis watershed, Cerros del Sira,
Arch. 4: 412. 1859. TYPE: Peru, Tatanara (Puno), Dudley 13339 (GH). Junin: Chanchamayo Valley, C.
Lechler 2530 (holotype, L?; isotypes, B!, P!; pho- Schunke 1491, 1559 (F, us). Puno: "St. Gavan" (San
tos, GH & us of P). Gaban), Lechler (K). Prov. Sandia, Chunchusmayo, We-
Hymenophyllum platylobum Bosch, Ned. Kruidk. berbauer 1237 (B).
Arch. 5:189.1863. TYPE: Peru, Dept. Puno, San
Gaban, Lechler 2489 (apparently cited in error
originally as 2498) (holotype, P; isotype, F!; photo,
us! of P).
Sphaerocionium valvatum (Hooker & Grev.) Copel., 19. Hymenophyllum microcarpum Desv., Mem.
Philipp. J. Sci. 67: 31. 1938. Soc. Linn. Paris 6: 333. 1827. TYPE: His-
paniola (holotype, P; photo, uc).
Leaves determinate, to 20 cm longhand 6 cm
broad. Petiole of mature leaves (2-)3-lOcm long, Hymenophyllum beyrichianum Kunze, Linnaea 9: 108.
0.4-1.0 mm in diameter, commonly alate just be- 1834. TYPE: Peru, Huanuco, Pampayacu, Poep-
low lamina base, pubescent with simple or forked pig (holotype, LZ destroyed).
Mecodium microcarpum (Desv.) Copel., Philipp. J.
trichomes or glabrate. Lamina pinnate-pinnatisect
Sci. 67: 25. 1938.
to nearly 2-pinnate-pinnatifid, sparsely to mod- Sphaerocionium microcarpum (Desv.) Copel., Phil-
erately pubescent, the rachis alate throughout, the ipp. J. Sci. 67: 34. 1938.
gins. Indusia commonly longer than broad, and pinnatisect to nearly 2-pinnate-pinnatifid, sparse-
slightly longer than the segment tips. ly to moderately pubescent, the rachis conspicu-
ously alate throughout. Pinnae 10-16 pairs, ovate-
On wet
tree trunks in forests, 900-1800 m, to elliptic-lanceolate, incised deeply or nearly to
Huanuco, Junin, Puno. the costa into bifid or pinnatifid pinnules, the ul-
Lesser Antilles; Guyana to Colombia, south to timate segments entire or deeply bifid, the veins
Bolivia. (abaxially)and margins sparsely to moderately pu-
It is with some reluctance that this
is maintained bescent, but trichomes lacking on tissue between
as distinct from Hymenophyllum microcarpum, the veins and margins. Indusia commonly longer
from which it seems to differ only in the undulate than broad, and slightly broader than the segment
ultimate segments and (sometimes) rachis wings, tips.
and the character of the petiole trichomes. Even
these features are variable, as only in a few spec- On tree trunks and clay banks in wet forests,
imens examined are the segments and rachis wings 1500-2700 m, Amazonas, Huanuco to Cuzco.
markedly undulate to crispate most are only Greater Antilles; Guatemala to Panama; Guy-
moderately so. Conversely, in a number of spec- ana to Colombia, south to Bolivia and Brazil.
68 FIELDIANA: BOTANY
With this probably might be included H. val- 21. Hymenophyllum simplex Morton, Contr. U.S.
vatum, under which see for further discussion. Natl. Herb. 29. 171. 1947. TYPE: Peru,
Huanuco, Tambillo, Jelski 897 (us!).
Amazonas: Prov. Bagua, E of La Peca, Barbour 2547
(MO). Huanuco: Rio Llullapichis watershed, ascent of Sphaerocionium simplex (Morton) Pic.-Ser., Webbia
Cerros del Sira, Dudley 13290B (MO). Pasco: Prov. Ox- 28: 472. 1973.
apampa, Palmazu, van der Werff et al. 8410 (MO, uc).
Junin: Chanchamayo Valley, C. Schunke 463 (f, us).
Cuzco: Prov. La Convention, Bites 2018, 2035 (us), 2034
Leaves determinate, fragile and small, 5-7 cm
(GH). Prov. La Convention, Alturas de Pintobamba, long. Petiole slender, 0.1-0.15 mm
diameter, in
Vargas 3294 (MO, us). nonalate, glabrate or sparsely pubescent with sim-
ple to stellate trichomes. Lamina simply pinnate
or pinnatisect, moderately pubescent, the rachis
20. Hymenophyllum ruizianum (Klotzsch) Kunze, narrowly alate distally, nonalate at base or in the
Hot. Zeit. (Berlin) 1847: 199. 1847. proximal Pinnae few, ascending, simple, lin-
half.
wings. Indusia about as broad as long, usually nar- be confused with H. fragile. However, the alate
rower than the ultimate segments, the receptacle condition in these atypical leaves is irregular and
not exserted. mostly due to the decurrent bases of pinnae,
whereas in H. fragile the wings are very conspic-
In dense forests, on tree trunks or mossy banks, uous and of nearly consistent width throughout.
800-2700 m, Pasco, Junin, Ucayali, Cuzco.
Mexico to Panama; Greater Antilles; Venezuela Ama/onas: Prov. Bagua, Cordillera Colan NE of La
and Colombia to Bolivia and Brazil. Peca, Barbour 3415 (F, MO). Cuzco: Michihuanuncca,
Biies 720 (us). Puno: Prov. Carabaya, Ayapata, Vargas
This may be confused with Hymenophyllum ele-
1071 7 (GH).
gantulum, under which see for further discussion.
Leaves indeterminate, 12-35 cm long, 2-4 cm Pendent from tree trunks and branches in wet
70 FIELDIANA: BOTANY
Sphaerocionium spectabile (Kuhn) Copel., Philipp. J. to long-stalked stellate trichomes, these mostly
Sci. 67:31. 1938.
spreading, not tightly appressed. Pinnae numer-
ous, larger ones 1-2.3 cm long, lanceolate or lin-
Leaves indeterminate, mature ones 30-180 cm
ear-lanceolate, acute to subattenuate, shallowly or
long, 4-9 cm broad. Petiole (of mature leaves) 0.5-
deeply incised, the 4-7 pairs of segments entire to
0.7 mm in diameter, nonalate, glabrate or sparsely bifid, the tissue and veins mostly obscured by a
pubescent with simple to stellate trichomes. Lam- dense tomentum or orange to tawny, short-stalked,
ina pinnate-pinnatifid or pinnate-pinnatisect,
stellate trichomes. Veins lacking accessory wings.
densely tomentose, the rachis nonalate through- Indusia commonly as broad as long, and as broad
out, commonly flexuous, with a tightly appressed as the segment tips, the receptacle not exserted.
covering of subsessile or short-stalked, stellate tri-
chomes. Pinnae numerous, larger ones (2.5-)3-8
Pendent from
trees or clay banks in wet forests,
cm long, lanceolate or linear-lanceolate, often at- 700-1900 m
throughout the range, but thus far
tenuate, incised deeply or nearly to the costa, the
apparently known in Peru only from one collection
6-14 pairs of segments shallowly to deeply bifid, cited below.
the ultimate divisions entire, the tissue and veins
Venezuela; Colombia; Peru.
mostly obscured by a dense tomentum of orange This and Hymenophyllum speciosum are very
(aging whitish), short-stalked, stellate trichomes.
closely related. See discussion of the latter for ad-
Veins lacking accessory wings. Indusia about as
ditional comments.
broad as long and often broader than the segment
tips, the receptacle not exserted.
San Martin: Near Tarapoto, in Monte Campana,
Spruce 4694 (BR, us).
Pendent from trees or clay banks in wet forests,
1800-3350 m, Amazonas, Pasco, Cuzco, Puno.
Peru and Bolivia.
This might better be treated as a variety ofHy- 27. Hymenophyllum lindenii Hooker, Sp. I'll. 1:
menophyllum karstenianum, as the two differ only 94, t. 34c. 1844. TYPE: Venezuela, Caracas,
quantitatively, e.g.,appressed versus spreading to- Linden 173 (holotype, K!; isotype, P; photos,
mentum, F of K, us of P).
strongly versus slightly flexuous rachis,
relative size and number of segments of pinnae.
Sphaerocionium lindenii (Hooker) Vareschi, Flora
Venezuela 1: 217. 1969.
Amazonas: Prov. Chachapoyas, Hutchison & Wright
5569 (F,GH, MO, uc, us); Sagastegui 7453 (F, HUT). Pasco:
Prov. Oxapampa, Cordillera San Gutardo, Leon et al. Leaves indeterminate, to 35 cm long and 8-15
534(usM). Cuzco: Vallede Lares, Biies 1808, 1817, 1821, cm broad. Petiole (8-) 10-1 5 cm long, 0.7-1 .2 mm
1931 (us). Valle de San Miguel, La Convention, Biies
in diameter, commonly nonalate, pubescent with
2178 (GH, MO, us).
simple to stellate trichomes. Lamina ovate or ovate-
lanceolate, not or slightly reduced at base, nearly
2-pinnate-pinnatisect, moderately pilose, the rachis
26. Hymenophyllum karstenianum J. W. Sturm, abundantly stellate-pubescent, scarcely alate, or
Bot. Zeit. (Berlin) 17: 298. 1859. LECTO- alate distally (or sometimes to base). Pinnae patent
Merida, Moritz 381 p.p. 14 deeply pinnatifid pinnules, the ultimate seg-
(BR).
ments entire to deeply bifid, rarely undulate or
apparently was what prompted Mettenius's orig- nonalate throughout. The entire species complex,
inal annotation on this sheet as "//. valvatum." which also involves H. dependens Morton and H.
Lechler 2489 is a mixed collection, as the other superbum Morton from northern South America,
four specimens of 2489 at Berlin are isotypes of needs closer examination. All may be conspecif-
H. platylobum (= H. valvatum). ic.
trichomes. Lamina linear, usually strongly and late, pinnatifid to pinnatisect, or rarely incised to
gradually reduced at base, often interruptedly fer- the costa, the ultimate segments entire (some prox-
tile (e.g., groups of fertile pinnae separated by some imal ones bifid), the tissue and veins completely
sterile ones), pinnate-pinnatifid to -pinnatisect, obscured by a tomentum of sessile to short-stalked,
moderately pilose, the rachis alate throughout, or stellate trichomes. Veins (especially abaxially) with
occasionally alate only distally or not at all. Pinnae low, inconspicuous lamellae. Indusia about as
patent to ascending, 1 5-many pairs, 1 .5-6 cm long, broad as long, usually slightly broader than the
acute to subattenuate, the 5-12 pairs of segments segment tips.
simple and entire to deeply bifid, the tissue be-
tween the veins sparsely to moderately provided Epiphyte on trunks or branches in dense, wet
with sessile or stalked, stellate trichomes. Veins pendent from wet, clay
forests, or occasionally
lacking accessory wings. Indusia about as broad banks, 550-3100 m, Huanuco, Junin, Cuzco,
as long or usually broader, as broad as the segment Madre de Dios, Puno.
tips. Costa Rica to Colombia; Peru; Bolivia; Brazil.
72 FIELDIANA: BOTANY
The lamellae along the veins in this species are irregularly interrupted, conspicuous lamellae. In-
scarcely "wings" in the sense of being broad and dusia about as broad as long, equaling or slightly
thin flanges of tissue. Instead, they are commonly broader than the segment tips.
long, low outgrowths borne along the abaxial or
(sometimes) adaxial sides of the veins contiguous Epiphyte in dense, wet forests, 2000-3200 m,
to, but not in the same plane with, the segment Amazonas, Huanuco, Pasco, Cuzco.
tissue. They are often difficult to discern, since the Colombia; Ecuador; Peru.
veins and laminar surface are so densely tomen- Shape of lamina and pinnae can be variable and
tose. The best way to observe them is on the older irregular in this species. Typically the leaves are
(proximal) pinnae wheresome of the indument linear, with numerous pinnae 3 or 4 cm long, yet
at times, scattered along the rachis, there may be
may have away. For further discussion see
fallen
also Hymenophyllum tomentosum. a few greatly elongated, attenuate pinnae, occa-
1 5 cm in
Easily confused with Hymenophyllum plumo- sionally reaching length. Although Hy-
sum are H. karstenianum and H. speciosum, which menophyllum multialatum has been thus far re-
have no lamellae on the veins, but are similar to ported from four departments in Peru, the great
the former in most other features, including the majority are represented by Biies's collections in
dense matting of tomentum obscuring the laminar Cuzco. For further discussion see H. tomentosum.
surface. Extreme care and patience must be ex-
Amazonas: Prov. Bagua, Cordillera Colan, Barbour
ercised to determine whether the low outgrowths
3740 (F, MO). Huanuco: Prov. Huanuco, 5 km from Car-
are present or lacking. In fact, the real importance pish, Tryon & Tryon 5318-B (GH). Pasco: Prov. Oxa-
of the presence or absence of lamellae bears more pampa, San Alberto, Cordillera de Yanachaga, van der
During this study of
careful analysis in the future. Werffet al. 8495 (MO). Cuzco: Valley of Rio Urubamba,
Biies A- 16 (F, GH, us). Montana de Calca, Biies 1916 (GH,
Peruvian species of Hymenophyllum, it has been
us), 1919, 1923, 1932 (us). Prov. La Convencion, Cor-
noted that size and frequency of these outgrowths, dillera Vilcabamba, Dudley 1 1 137 (GH, us).
on one or both surfaces, may be more variable
than earlier assumed, and thus may be of much
less taxonomic significance. 31. Hymenophyllum tomentosum Kunze, Lin-
naea 9: 107. 1834. TYPE: Peru, Huanuco,
Huanuco: Vilcabamba, Hacienda on Rio Chinchao, Pampayacu, 1829, Poeppig Diar. 1134 (ho-
Macbride 5146 (F, us). Junin: Schunke Hacienda above lotype, w?; isotype, BM?; frag., B!; probable
San Ramon, Killip & Smith 24840 (F, GH, us). Cuzco:
isotype, MO!; photo, us of MO).
Prov. Paucartambo, above Tambomayo, West 7108 (MO,
uc). Madre de Dies: Prov. Mania, Shintuya, Chavez 860
(MO). Puno: San Gaban, Lechler 2264 (B). Sphaerocionium tomentosum (Kunze) Presl, Hymen-
ophyllaceae 34. 1843.
Hymenophyllum fusagasugense J. W. Sturm, Bot. Zeit.
(Berlin) 1859: 297. 1859 (as "fusugasugense").
30. Hymenophyllum mult ialat urn Morton, Contr. TYPE: Colombia, Cundinamarca, "Fusugasuga"
U.S. Natl. Herb. 29: 185. 1947. TYPE: (Fusagazuga) Karsten (not located).
Peru,
Hymenophyllum fusagasugense var. aberrans Mor-
Cuzco, Prov. La Convencion, Alturas de Sieve,
ton, Contr. U.S. Natl. Herb. 29: 187. 1947 (as
Biies 1575 (holotype, us!; frag., GH!). "fusugasugense"). TYPE: Venezuela, Merida,
Tabay, Gehriger 584 (holotype, us!; isotypes, F!,
Sphaerocionium multialatum (Morton) Pic.-Ser., GH!).
Webbia28: 471. 1973.
Leaves indeterminate, to 50 cm long and 2.5 cm
Leaves indeterminate, to 50 cm long, 3-8(-15) broad. Petiole 0.5-0.6 mm
in diameter, nonalate,
cm broad. Petiole 0.4-0.7 mm in diameter, non- sparsely pubescent with mostly simple trichomes.
alate, sparsely hirsute with mostly simple tri- Lamina pinnate-pinnatifid, densely hirsute, the
chomes. Lamina pinnate-pinnatisect, hirsute, the rachis nonalate throughout. Pinnae numerous, ob-
rachis nonalate throughout. Pinnae numerous, long or narrow-deltoid, pinnatifid (sometimes
narrow-oblong or -deltoid, pinnatisect, sometimes deeply so), the ultimate segments simple or (oc-
irregularly and greatly elongate, the ultimate seg- casionally) bifid, the tissue and veins densely cov-
ments commonly bifid, the tissue and veins mod- ered with (and often obscured by) sessile to short-
erately provided with sessile to short-stalked ses- stalked trichomes. Veins abaxially bearing irreg-
sile trichomes. Veins bearing (especially abaxially) ularly interrupted, conspicuous winglike lamellae,
adaxially. Indusia about as broad as long, equaling late trichomes. Veins abaxially bearing small,
or slightly broader than the segment tips. scattered lamellae, these about as broad as long,
but lacking adaxially. Indusia slightly longer than
On
steep banks and tree trunks, 2750-3100 m, broad, usually as broad as the segment tips, re-
Amazonas, Huanuco. ceptacle not exserted.
Colombia; Venezuela; Peru.
This is distinguished from Hymenophyllum from trunks and branch-
In wet forests, pendent
multialatum and H. plumosum only by the char- es of trees,600-1300 m, Loreto and Huanuco.
acters used in the key. All three appear to be alike, Colombia; Ecuador; Peru.
except for the size and frequency of the lamellae There is little, beyond the key characters, to
along the veins and by the relative density of pu- distinguish this from Hymenophyllum pyrami-
bescence. Of these, H. plumosum (q.v.) is the most datum. Leaves of the latter are perhaps more ro-
distinct, in that the lamellae are merely low and bust, with relatively narrower pinnae and much
inconspicuous outgrowths and the trichomes are denser pubescence. Both also share nearly the same
so densely matted as to completely obscure the range, and more study may prove them to be con-
veins and tissue. In H. tomentosum the indument specific. Also very closely related is H. verecun-
isalso densely matted, but the lamellae are so large dum, which is more easily separated by its much
and conspicuous that many of them emerge through lighter coloredtrichomes and shorter, relatively
the tomentum. Morton (1947) chose to separate broader and more crowded pinnae. But even these
H. fusagasugense and var. aberrans on the pres- characters sometimes tend to be intermediate.
ence or absence of these processes adaxially. How-
ever, the size or lack of lamellae on the adaxial Loreto: Pumayucu, between Balsapuerto and Moy-
side varies considerably in the species, and the obamba, Klug 3248 (B, F, GH, MO, us). Huanuco: Prov.
character is difficult to observe due to the dense
Huanuco, Dist. Churubamba, Mt. Santo Toribio, Mexia
8258 (B, F, GH, uc, us).
surface tomentum. To prove that the lamellae are
completely absent on a given specimen would re-
quire scraping away all the tomentum of each leaf.
33. Hymenophyllum pyramidatum Desv., Mem.
A thorough study of the species complex through- Soc. Linn. Paris 6: 332. 1827. TYPE: "habitat
out its range is needed.
inAmerica" (possibly a Dombey collection
from Peru) (holotype, P; photos, GH, us).
Amazonas: Prov. Chachapoyas, Cerros de Calla Calla,
19 km above Leimebamba, Hutchison & Wright 5570
(uc). Prov. Chachapoyas, near km 422, Balsas-Leime- Sphaerocionium pyramidatum (Desv.) Copel., Phil-
bamba road, Wurdack 1249 (F, GH, uc, us). ipp. J. Sci. 67: 30. 1938.
naea 20: 438. 1847. TYPE: Peru, Panatahua, 1-2 cmnear the apex, pubescent with simple to
Ruiz 83 (holotype, stellate trichomes. Lamina pinnate-pinnatifid,
B!; isotype, MA?).
densely pubescent with simple to stellate tri-
Sphaerocionium lobatoalatum (Klotzsch) Pic.-Ser., chomes, the rachis alate throughout. Pinnae nu-
Webbia28:471. 1973. merous, larger ones 20-45 mm
long, 6-1 1 mm
broad at base, most of them narrow-triangular,
Leaves indeterminate, to 60 cm long and 8 cm the apex narrowly acute to attenuate, deeply lobed
broad. Petiole 0.4-0.7 mm
in diameter, alate, if to pinnatifid, the 6-12 pairs of ultimate segments
at all, only near base of lamina, pubescent with simple to bifid, the tissue and veins usually co-
simple to stellate trichomes or glabrate. Lamina piously provided with orange to dark brown (rare-
pinnate-pinnatifid, pubescent with stellate tri- ly tawny), subsessile to short-stalked, stellate
chomes, the rachis strongly alate throughout. Pin- trichomes. Veins abaxially with numerous, con-
nae numerous, larger ones 20-45 long, 7-1 1 mm spicuous, elongate, winglike lamellae (often nearly
mm broad at base, most of them narrow-trian- the length of the vein), these sometimes less con-
gular, the apex narrowly acute to subattenuate, spicuous and abundant on the adaxial surface. In-
deeply pinnatind, the 7-12 pairs of ultimate seg- dusia slightly longer than broad, usually as broad
ments bifid, the tissue and veins moderately pro- as the segment tips, receptacle not exserted.
74 FIELDIANA: BOTANY
In dense, wet forests, pendent from stumps, tree (us). Madre de Dios: Prov. Manii, Shintuya, Chavez 833
and branches, and on wet banks, 600-1 800 p.p. (MO). Puno: Prov. Sandia, E of Oconeque, Hodge
trunks,
6090 (F, GH, us).
m, along Cordillera Central from San Martin to
Cuzco.
Colombia; Peru; Bolivia. 35. Hymenophyllum tarapotense Stolze, sp. nov.
This belongs to a close-knit complex of species
involving Hymenophyllum verecundum and H. Folia usque ad 6 cm longa et 1.6 cm lata. Petiolus 1-
lobatoalatum. See discussion of the latter for fur- 2 cm loneus. 0.1-0.15 mm
diametro. Lamina pinnato-
ther comments. pinnatifida. Rhachis versus basin non alata. Pinnae 6-
12-jugatae, bifidae vel pinnatifidae, usque ad 1 cm lon-
San Martin: Tingo Maria, Allard 20818 (us). Huan- gae, pinnae basales admodum reductae, marginibus, venis
et interveniis trichomatibus stellatis instructis. Venae
uco: Pampayacu. Kanehira 123 (F, GH, K, us). Pasco:
lamellis cristiformibus numerosis praeditae.
Prov. Oxapampa, 4-8 km from Enenas, Skoget al. 5077
(us). Junin: Chanchamayo Valley, C. Schunke 466, 926
(F.us), 7055 (F). Cuzco: Prov. Paucartambo, between Leaves indeterminate, to 6 cm long and 1.6 cm
Mistiana and Keros, Vargas 7380 p.p. (MO). broad. Petiole 1-2 cm long, 0.1-0.15 in di- mm
ameter, nonalate, sparsely provided with simple
trichomes. Lamina pinnate-pinnatifid, pubescent
34. Hymenophyllum verecundum Morton, Contr. with stellate trichomes, the rachis weakly alate dis-
U.S. Natl. Herb. 29: 183. 1947. TYPE: Peru, nonalate toward the base. Pinnae 6-12 pairs,
tally,
Huanuco, Churubamba, Mexia 8143-A (ho- bifid, or distal ones pinnatifid with 3-4 segments,
lotype, us!; isotypes, GH!, MO!, uc!). larger ones 1 cm long, basal ones often reduced to
mere auricles, the margins, veins, and intervening
Leaves indeterminate, to 50 cm long and 4 cm tissuemoderately to amply provided with tawny
broad. Petiole 0.4-0.6 mm in diameter, alate, if trichomes. Veins bearing numerous, conspicuous
at all, just below the lamina, glabrate or slightly accessory foliar crests not in the plane of the lam-
pubescent with simple to stellate trichomes. Lam- ina, these less frequent on the adaxial surface. In-
ina pinnate-pinnatifid, densely pubescent with dusia about as long as broad, as broad as the seg-
stellate trichomes, the rachis alate throughout, or ment tips, receptacle minute, not exserted.
sometimes nonalate at base. Pinnae numerous,
larger ones6-20 mm
long, 3-7 mm
broad at base, TYPE Peru, Tarapoto (Dept. San Martin) in
oblong to very broadly triangular, the apex obtuse monte Guayrapurima, Aug. 1856, Spruce 4048
to subacute, pinnatifid, the 3-6(-8) pairs of ulti- Another Spruce specimen
(holotype, K!; photo, F).
mate segments simple to retuse or occasionally at K, with notation "conf. 4048," is H. simplex.
bifid, the tissue and veins copiously provided with Known only from the type.
whitish to tawny, subsessile or short-stalked, stel- This tiny fern is unquestionably the smallest of
late trichomes. Veins abaxially with numerous the species in subg. Leptocionium which bear ac-
conspicuous, elongate, winglike lamellae, these less cessory tissue on their veins. These foliar processes
conspicuous and abundant on the adaxial surface. are not long, low wings as seen in Hymenophyllum
Indusia about as long as broad and usually as broad of
pyramidatum, but are separate, crestlike flanges
as the segment tips, receptacle not exserted. tissue similar to those frequently seen in more
delicate specimens of//, verecundum. Superficially
from tree trunks,
In dense, wet forests, pendent //.tarapotense closely resembles H. fragile, in its
branches, and wet banks, (5 50-) 1500-2 700 m, small size, delicate petiole, and reduced, often bi-
Amazonas, Huanuco, Junin, Cuzco, Madre de fid, pinnae. It may seem strange that this distinc-
Dios, Puno. tive species has not been described before, nor
Colombia; Peru; Bolivia. collected again in over 1 00 years. However, many
This is closely related to Hymenophyllum pyr- of these diminutive members of the genus are very
amidatum and H. lobatoalatum. See discussion of easily overlooked by collectors.
the latter for further comments.
er, Syn. fil. 65. 1 867. TYPE: Peru, San Martin, dimorphic, 0.540 cm or longer. Lamina entire to
Tarapoto, Spruce (not located). nearly 5 -pinnate, subsessile to petiolate, with pet-
iole commonly shorter than the lamina, glabrous
Morton (p. 155, 1947) considered that this is or with trichomes borne mostly on costae and
the same as Hymenophyllum adiantoides, and there margins. Veins free or, in Trichomanes diversi-
is nothing in the description to argue otherwise. It frons, reticulate near the lamina margin, pinnate
is possible that Baker was unaware of the publi- (anadromous or catadromous) or sometimes fla-
cation of the latter species, and even that he based bellate, false veins also present in some species,
his new species on the same specimen cited earlier these mostly parallel, but in a few species perpen-
by van den Bosch. Specimens (B, GH, K, L, NY, us) dicular to the true veins. Sori terminal on the veins.
thought to be probable isotypes of H. adiantoides Indusium mostly tubular, the mouth bilabiate or
are those of Spruce from Tarapoto. They bear the entire and often
flaring, or very rarely nearly bi-
notation "conf. 4700" and are conceivably the ones valved, the sporangia borne on an elongate recep-
on which Baker based his "//. sprucei." tacle. Receptacle commonly exserted beyond the
mouth of the indusium, often greatly so.
Hymenophyllum trifidum Hooker & Grev., Icon, There are about 300 species in the genus, oc-
fil. 2, 1. 196. 1 83 1 TYPE: Ecuador (as "Peru"),
.
curring in tropical to subtropical regions of both
Esmeraldas, Jameson (E?). hemispheres. As in Hymenophyllum, most are
76 FIELDIANA: BOTANY
FIG. 12. Trichomanes radicans: a, habit; b, pinna with sori. Trichomanes pinnatum: c, portion of pinna with false
veins and sori. Trichomanes hymenoides: d, apical portion of leaf, (a from Acosta-Solis 13782, Ecuador, F, b from
Malhias & Taylor 5075, F, c from Barbour 5194, F, d from Wurdack 1513, F.)
flaring; tissue of most ultimate segments with elongate, narrow folds parallel to the veins
4. T. pyxidiferum
f. Rachis nonalate nearly to the apex; pinnae stalked, the costae not alate at base h
h. Costae alate, except not below the basal pinnule; ultimate segments 0.5-0.8 broad; mm
sori 1-5 per pinna; indusia commonly alate, the wings several cells wide
6. T. angustatum
h. Costae nonalate except near the apex; ultimate segments 0.1-0.4 mm broad; sori rarely
more than 1 per pinna; indusia not or scarcely alate, wings (if any) 1 cell wide
7. T. capillaceum
b. Stem erect, or occasionally short-creeping; leaves caespitose to (occasionally) approximate [subg.
Pachychaetum] i
78 FIELDIANA: BOTANY
i. Tissue of segments opaque, the cells small and occluded, not evident when observed with
magnification of 8-12 x; leaves (mature ones) 15-90 cm long; petiole (0.6-)0.7-3.5 mm in
diameter j
j. Rachis obtusely quadrangular, alate (often broadly so) at least in the distal half; pinnae
adnate, the costae alate; lamina usually more than 1 cell thick, at least away from margins
8. T. elegans
j. Rachis or marginate to slightly alate near apex; pinnae (at least proximal
terete, nonalate,
ones) short-stalked, costae not alate at base; lamina essentially 1 cell thick throughout . . .
9. T. rigidum
i. Tissue of segments translucent, the cells large and clear, quite evident with magnification of
8-12 x; leaves commonly less than 15 cm long; petiole 0.3-0.6 mm
in diameter
10. T. cellulosum
a. Venation at least partly sometimes flabellate); false veins present or lacking
catadromous (or k . . .
k. False veins present, these submarginal or borne between and parallel with the true veins [subg.
Didymoglossum] 1
1. Leaves with simple to stellate, commonly dark, trichomes along the margins; indusia distinctly
bilabiate and the lips commonly dark-margined m
m. Venation flabellate, a percurrent costa lacking 11. T. punctatum ssp. sphenoides
m. Venation pinnate, or if subflabellate, the costa essentially percurrent n
n. Mature leaves entire, or distally lobed; stellate trichomes borne all along the margin
1 2. T. angustifrons
n. Mature leaves 1-2-pinnatifid; stellate trichomes confined to segment sinuses, with simple
or bifid trichomes borne on outer margin o
o. Son protruding from the leaf tissue, or slightly immersed at base; false veinlets
q. Lamina bearing numerous paired, circular, scalelike processes all along the margin (these
sometimes deciduous in age), but lacking a submarginal false vein; venation flabellate . . .
1 7. T. membranaceum
k. False veins lacking or, in T. pinnatum, perpendicular to the true veins (some short, veinlike lines
in sect. Lacostea); [subg. Achomanes] r
r. Lamina trichomes stellate or forked from the base; stem filiform, 0.3-0.4 mm in diameter
18. T. polypodioides
r. Lamina trichomes simple or lacking; stem relatively stout and wiry, 0.5-1.2 mm in diameter
s
s. Stem long-creeping up tree trunks, leaves subdistant to remote; lamina appressed to tree
trunks by means of rhizoids on axes t
t. Lamina linear, subentire to lobed, 1-1.5 cm broad; indusia mouth rather widely flaring
1 9. T. tanaicum
t. Lamina oblong, pinnatisect or nearly pinnate, mature ones ( 1 .8-)2-9 cm broad; indusium
mouth not or slightly flaring u
u. Segments of sterile laminae entire or shallowly crenate; veins simple, not forked;
indusium mouth not expanded; rare in Peru 20. T. tuerckheimii
u. Segments of sterile laminae deeply crenate or crenate-serrate; veins (some or many)
1 -forked; indusium mouth slightly flaring; locally common 21. T. ankersii
essentially entire, or without laminar tissue and the sori completely free x
x. Fertile leaves with lamina entire to pinnatisect; sori fully immersed in tissue y . .
y. Sterile lamina with 4-8 veins issuing from the rachis between adjacent costae;
mouths of indusia neither flaring nor recurved, strongly indented between the
vein branches 23. T. diversifrons
y. Sterile lamina with 2(-3) veins issuing from the rachis between adjacent costae;
mouths of indusia slightly flaring and recurved, but not indented between the
vein branches 24. T. trollii
x. Fertile leaves without laminar tissue; sori completely free 25. T. botryoides
. . .
w. Stem wiry, long- or short-creeping; leaves subdistant to remote; fertile leaves with
lamina foliose and deeply pinnatisect to nearly pinnate 28. T. humboldtii
v. Leaves monomorphic z
z. Lamina 3-4-pinnatisect; petiole conspicuously alate throughout, the alae broader than
the petiole quite or nearly to base 22. T. bicorne
z. Lamina not more than 1 -pinnate-pinnatisect; petiole nonalate in the proximal half
or throughout, or very narrowly alate in the proximal portion aa
aa. Lamina abruptly terminating in a conform apical segment or the rachis prolonged,
flagellate, and proliferous at tip; indusia fully exserted from segment margin, often
stalked bb
bb. False veins copious and perpendicular to the true
veins; marginal vein strong,
continuous; pinna margins sharply serrate to spinulose .26. T. pinnatum .
bb. False veins rare or lacking; marginal vein lacking or faint and discontinuous,
pinna margins commonly obtusely serrate 27. T. hostmannianum
aa. Lamina gradually reduced to a pinnatifid apex, the rachis not prolonged and
flagellate; indusia at least partially immersed in the laminar tissue, wholly or
partially alate on each side cc
cc. Pinnae regularly and deeply lobed to pinnatisect dd
dd. Pinnae lobed to pinnatifid; lamina l-2(-2.5) cm broad, not at all obscured
by the moderate pubescence 29. T. crinitum
dd. Pinnae pinnatisect (rarely the segments again lobed); lamina (2.5-)3-12
cm broad, partially to completely obscured by the dense covering of
trichomes 31. T. lucens
cc. Pinnae entire to serrate or crenate (or rarely a few with some irregularly
scattered lobes) ee
ee. Veins bearing crestlike lamellae on abaxial side; pinnae in proximal por-
tion of lamina deflexed 30. T. martiusii
ee. Veins lacking lamellae; pinnae patent or a few basal ones deflexed ff . .
ff. Petiole conspicuously alate halfway or more to the stem; stem erect
to decumbent, leaves contiguous to caespitose .... 32. T. pellucens
ff. Petiole not alate, or scarcely so near apex due to decurrent lamina
base; stem short-creeping or decumbent, the leaves subdistant to con-
tiguous (rarely caespitose) gg
gg. Rachis trichomes on abaxial side predominantly dark brown, stout,
rigid, most of them terete toward their base 33. T. plumosum
. .
flattened hh
hh. Lamina mature ones commonly
linear or linear-lanceolate,
5-12 times as long as broad; mature leaves (10-) 15-50 cm
80 FIELDIANA: BOTANY
long; rachis abaxially provided with trichomes predominantly
unicellular beyond the basal one, occasionally with 2-3 cells
34. T. cristatum
hh. Lamina oblong- to ovate-lanceolate, 3-4 times as long as
broad; mature leaves 5-15 cm long; rachis abaxially provided
with trichomes of 2-8 cells beyond basal one, as well as with
many unicellular ones 35. T. vandenboschii
1. Trichomanes radicans Sw., J. Hot. (Schrader) rupestre form a close-knit species complex which
1800(2): 97. 1802. TYPE: Jamaica, Swartz is need of closer study. The three are only pro-
in
(holotype, s; photo, us). Figure 12a-b. visionally maintained here at the species level. In
Central America T. radicans and T. collariatum
iole 0.2-2 cm long, 0.6-0.8 mm in diameter, par- Indusia broadly conical, the tube 1-1.5 times as
tially alate. Lamina commonly 1-pinnate-pinna- long as broad, not or scarcely bilabiate, the mouth
oblong or elliptic-oblong. Rachis broadly alate
tifid, not or slightly flaring, receptacle commonly long-
throughout. Pinnae pinnatifid or pinnatisect, or exserted.
rarely 2-pinnatifid as to basal segments, strongly
adnate to rachis. Ultimate segments relatively short On trees in wet forests, 900-1 200 m, Amazonas,
and broad, not linear, with simple or forked veins, San Martin.
false veinlets lacking. Venation anadromous. In- Pantropical.
dusia conical, gradually widening from base to a A distinctive feature of Trichomanes pyxidifer-
slightly flaring mouth, narrowly alate on each side, um is the rather sharp, longitudinal folding of seg-
receptacle short- to long-exserted. ment tissue parallel to the veins. Other related
species have plane segments, or some with the
Creeping on tree trunks or wet banks in dense, margins undulate, but none exhibit this unique
wet forests, 1 100-1600 m, Huanuco and Ucayali. character. Perhaps it was this which prompted
Venezuela and Colombia; Peru; Bolivia; Brazil Morton's incorrect observation (1968) of false veins
(also reported from Costa Rica by Alan Smith, in. in the species, since the sharper folds of tissue
82 FIELDIANA: BOTANY
sclerenchymatous strands which produce these axes and indusia, crowding of pinnae and dissec-
"false veins." The species thus far is represented tion of lamina. Each of these may appear to be
in Peru by only two collections, even though its important in individual specimens, but when many
full range extends throughout tropical regions of collections are examined throughout the range of
the world. distribution, it becomes evident that they are quite
variable and uncorrelated.
Amazonas: Prov. Chachapoyas, .la/an (Ingenio-
Chachapoyas), Ldpe: el al. 4264 (GH, HUT). San Martin: Amazonas: Prov. Bagua, Cordillera Colan SE of La
Tarapoto, Spruce 4761 (BR, OH, K, L, us). Peca, Harbour 3 904 (MO). I .ore to: Sierra del Pongo, Mex-
ia 6288B (GH, uc, us). Pasco: Prov. Oxapampa, Dist.
Oxapampa, road Oxapampa to Villa Rica, B. Ledn 660
(USM La Merced, E of Quimiri Bridge,
in part). Junin:
5. Trichomanes diaphanum HBK., Nov. gen. sp. Killip & Smith 24006 (F), 24023 (GH, us). Cuzco: Prov.
1:25 (fol. 1 6). 1 8 1 6. TYPE: Venezuela, Hum- La Convencion, "El Dorado," Vargas 3518 (GH). Puno:
boldt & Bonpland (holotype, p; isotype, B!).
San Gaban, Lechler (L).
ameter (excluding indument), sparsely to abun- Vandenboschia angustata (Carm.) Copel., Philipp. J.
dantly provided with castaneous trichomes. Leaves Sci. 73: 466. 1941.
0.5-6 cm long, 0.3-0.8 mm in diameter, flattened, chomes. Leaves well-spaced to remote, 4-14(-18)
or terete at base, nonalate, or narrowly to broadly cm long, glabrous. Petiole 0.5-5 cm long, 0.2-0.4
alate for most of its length, the wings plane or mm in diameter, terete, nonalate. Lamina 2-pin-
hymenophylloides on characters such as number This is very closely related to Trichomanes cap-
of sori per pinna, width of tissue wings flanking illaceum, under which see further discussion.
In deep forests, on tree trunks, 800-1100 m, ler 2175 (holotype, L?; isotype, P!; photo, F, frag.,
Pasco and Cuzco. L!).
Southern Mexico to Panama; Greater Antilles; Davalliopsis elegans (Rich.) Copel., Philipp. J. Sci. 67:
82. 1938.
Venezuela and Colombia to Peru.
With this might possibly be included Trichom-
anes angustatum, but the two are separated here Terrestrial, rarely epiphytic or epipetric. Stem
provisionally, pending much needed monographic Leaves caespitose to
erect or rarely short-creeping.
revision. Typically, T. capillaceum has very little occasionally approximate, 20-90 cm long, rachis,
expanded tissue along the axes, and even the ul- costae, and often costules amply to abundantly
timate segments are rarely more than 0.3 mm provided with castaneous, pluricellular trichomes
broad. Indusia are not or scarcely alate, and son on abaxial side. Petiole 10-40 cm long, 1-3.5 mm
are solitary on each fertile pinna. Conversely, be- in diameter, usually alate distally. Lamina 2-3-
yond the first pinnule in T. angustatum, the costae pinnate-pinnatifid, deltoid, opaque, usually more
and costules are obviously alate throughout, and than 1 cell thick, at least away from the margins,
ultimate segments are often nearly 1 mm broad. the cells small and occluded, not evident even when
The tissue also forms wings on each side of the magnified at 10-15x. Rachis obtusely quadran-
indusia, and sori are frequently four or five per gular, alate (often broadly so) at least in distal half.
pinna. Pinnae adnate, the costae alate to base. Ultimate
Even some of the characters listed in the key are segments laciniate, with linear lobes, each of these
subject to variation in certain geographic regions, bearing a single vein, false veins lacking. Venation
and intermediates are found frequently. Never- anadromous. Sori usually bent down away from
theless there are areas in which only one or the the plane of the lamina. Indusia subconical to
other occur. Only Trichomanes angustatum oc- somewhat urceolate, not bilabiate, the mouth
curs in Bolivia and southern Brazil, whereas in truncate or slightly (rarely strongly) flaring, recep-
Central America it has not been found south of tacle short- to long-exserted.
Guatemala, a region where T. capillaceum is rath-
er common. In Peru, the latter is rare, and even In dense, wet forests or wet, wooded ravines, on
the few specimens located are not typically skeletal ground, or rarely on tree trunks or wet rocks, sea
in their appearance, whereas Trichomanes angus- level to 1300 m, Amazonas and Loreto south to
tatum is and the specimens
rather widespread Madre de Dios and Puno.
rather typical. One specimen from Cuzco, Vargas Lesser Antilles and Trinidad; Nicaragua to Pan-
8653 (MO), is exactly intermediate between the ama; Guianas to Colombia and south to Bolivia
two; significantly, Cuzco is one of the two de- and Brazil.
84 FIELDIANA: BOTANY
With its large,blue green leaves and laciniate essentially in the plane of the lamina. Indusia sub-
pinnae, this is one of the most beautiful species in conical, not bilabiate, the mouth truncate, not or
the family. Its most distinctive features are the scarcely flaring, receptacle commonly long-exsert-
usually strongly arching sori (bent downward out ed.
of the plane of the lamina) and the relatively thick
tissue. Although the lamina in Hymenophyllaceae On slopes and ridges of dense forests and in
is one cell thick, the tissue in Tri- wooded ravines, occasionally on wet rocks or cliffs,
typically only
chomanes elegans is often several cells thick, es- rarely on bases of tree trunks, 350-2500 m, Ca-
pecially away from the segment margins. The jamarca and San Martin, south to Cuzco and Madre
de Dios.
broadly alate rachis traditionally has been used to
Southern Mexico to Panama; West Indies; South
separate this from T. rigidum (with which it often
grows), and although usually it is a good diagnostic
America, to Bolivia and Brazil; probably Old World
feature, occasional specimens of T. elegans can be tropics.
found throughout the range in which rachis alae This is sometimes confused with Trichomanes
are very narrow or lacking in the proximal portion. elegans, with which it often grows. For compari-
son, see discussion under the latter species.
Consequently, supporting key characters should
be employed for accurate identification.
Cajamarca: Tambillo, Jelski 891 (us). San Martin:
Tarapoto, in monte Guayrapurima, Spruce 4047 (BR, K).
Amazonas: Prov. Bagua, left bank of Rio Maranon Huanuco: East of Tingo Maria (as San Martin), Allard
opposite Quebrada Mirana, Wurdack 2031 (F, GH, us). 22356 (GH, us). Pasco: Oxapampa, Cordillera San Ma-
San Martin: Prov. Mariscal Caceres, Tocache Nuevo, J. tias, Leon 326a (USM). Junin: La Merced, Hacienda
Schunke4855 (F, GH, us), 6955 (F, MO in part, us). Loreto: Schunke, Macbride 5633 (F, us). Ucayali: Vicinity of
Sierra del Pongo, Mexia 1 195 (F, GH, MO, us). Huanuco:
Aguaytia, along Rio Aguaytia (as Loreto), Croat 20927
Prov. Pachitea, Dist. Honoria, /. Schunke 2259 (F, GH).
(MO). Cuzco: Prov. Urubamba, base of Huayna Picchu,
Pasco: Prov. Oxapampa, Valle del Palcazu, Iscozacin, Iltis et al. 1062 (GH). Madre de Dios: Prov. Manu, Cerro
Le6n 692 (F, GH). Junin: Puerto Bermudez, Killip & Smith de Pantiacolla, Foster 10903 (F).
26537 (GH, us). Ucayali: Prov. Coronel Portillo (as Lor-
eto), Padre Abad, J. Schunke 5393 (F, us). Cuzco: Bajada
de Tocate, Biies 1748 (us). Madre de Dios: Prov. Tam-
10. Trichomanes cellulosum Klotzsch, Linnaea 18:
bopata, Rio Tambopata Nature Reserve, Barbour 5165
(MO). Puno: Prov. Carabaya, San Gaban, Vargas 18864 531. 1844. TYPE: "British Guiana," (Guy-
(GH). ana). Kanuku Mountains, Rich. Schomburgk
1186 (holotype, B!; photo, F; isotypes, B! 2
sheets, BM!).
9. Trichomanes rigidum Swartz, Prodr. 137. 1788.
TYPE: Jamaica, Swartz (holotype, s; isotype, Epiphytic, terrestrial, or sometimes epipetric.
B, Herb. Willd. 20202- 7; photos, us of s, GH Stem erect to short-creeping. Leaves caespitose to
& us of B). (occasionally) approximate, 6-15 cm long, gla-
brous. Petiole 1-7 cm long, 0.3-0.6 mm
in di-
Selenodesmium rigidum (Sw.) Copel., Philipp. J. Sci. ameter, narrowly alate at least distally. Lamina 4-
67:81. 1938.
5-pinnate, lanceolate to ovate, translucent, the cells
largeand clear and quite evident when magnified
Terrestrial, rarely epiphytic or epipetric. Stem at 8-12x. Rachis terete, alate throughout. Pinna
erect or occasionally short-creeping. Leaves caes- divisions skeleton-like, the costae, costules, and
sometimes approximate, 1 5-35 cm long,
pitose to ultimate segments with bands of tissue only 1-
lamina glabrous, the axes glabrous, or rarely the several rows wide. Ultimate segments linear, 0.3-
rachis or petiole base with a few, scattered tri-
0.7 mm broad, each bearing a single vein, false
chomes. Petiole 5-16 cm long, 0.6-1.6 mm in di- veins lacking. Venation anadromous. Indusia sub-
ameter, nonalate. Lamina 3-4-pinnate-pinnatifid, conical to somewhat urceolate, not bilabiate, the
deltoid, opaque, 1 cell thick, but cells small and mouth truncate or slightly flaring, receptacle short-
occluded, not evident even when magnified at 1 0- to long-exserted.
1 5 x Rachis terete, nonalate, or marginate to
.
slightly alatetoward apex. Pinnae (at least prox- In wet forests, on trees or on the forest floor,
imal ones) short-stalked and the costae not alate sometimes on wet rocks, 300-1 300 m, Amazonas,
at base. Ultimate segments with linear lobes, each Loreto, Huanuco.
of these bearing a single vein, false veins lacking. Surinam to Colombia; Peru; Brazil.
Venation anadromous. Sori (most of them) borne With this perhaps should be included Trichom-
1 1 . Trichomanes punctatum Poiret ssp. sphen- catadromous, pinnate, with false veinlets sparsely
oides (Kunze) Boer, Acta Bot. Neerl. 1 1: 301. to amply distributed between the true veins. Sori
86 FIELDIANA: BOTANY
well-spaced, (0.5-)l-3.5 cm long, to 1.5 cm broad, nuses and simple to bifid on outer margins, lam-
subsessile or short-petiolate. Lamina obovate to inar cells mostly elongate. Venation catadromous,
elliptic, 1-pinnatifid or rarely 2-pinnatifid, the cos- pinnate, false veinlets commonly abundant, ex-
ta percurrent (but juvenile laminae often subren- tending toward lamina margin but not (or very
iform and merely lobed and the costa indistinct rarely) parallel to it. Sori 1 or a few, borne near
distally),glabrous except for the dark, marginal lamina apex, conspicuously protruding from the
trichomes which are stellate in sinuses and simple leaf tissue, or only slightly immersed at base, not
to bifid on outer margin, laminar cells commonly or scarcely alate. Indusia narrow-funnelform, bil-
isodiametric. Venation catadromous, commonly abiate, the lips dark-margined, and sometimes
pinnate (occasionally subflabellate in juvenile flaring, most of them as long as or longer than
leaves), false veinlets sparse, scattered between true broad, receptacle short- to long-exserted.
veins, extending toward lamina margin but not
parallel to it. Sori 1-several borne near lamina On wet rocks in forests, 1900-2750 m, Ama-
apex, conspicuously exserted, or only slightly im- zonas, Huanuco.
mersed at base, not or very narrowly alate. Indusia Southern Mexico to Panama; Greater Antilles;
narrow-funnelform, bilabiate, but the lips quite Venezuela; Colombia; Ecuador; Peru; Brazil; Ar-
short, commonly broader than long, and usually gentina.
(not always) dark-margined, receptacle long- Although found thus far in Peru only on wet
exserted. rocks, Trichomanes reptans may be expected also
on trees, since it frequently has been reported as
In forests on trees, or rarely on rocky or sandy an epiphyte throughout much of its range.
soil, 700-2400 m, Amazonas, Loreto.
Southern Mexico to Panama; West Indies; Trin- Amazonas: Prov. Bagua, Cordillera Colan SE of La
idad; Venezuela and Colombia, south to Argentina Peca, Harbour3903 (MO, USM). Huanuco: Mito, Bryan
392 (F).
and Uruguay.
Juvenile leaves are often nearly reniform and
shallowly lobed, with veins subflabellate and the
costae sometimes indistinct toward the apex. Hence
15. Trichomanes krausii Hooker & Grev., Icon,
fil. 2, /. 149. 1831. TYPE: Dominica, Kraus
they can be confused with Trichomanes puncta-
tum. However, fertile leaves are pinnatifid, with (holotype, E).
88 FIELDIANA: BOTANY
tinuous submarginal false vein lacking. Sori sev- sparsely scattered along the stem, unlike the dense,
eral to many on vein tips toward the lamina apex, blackish indument found on stems of T. reptans
partially to fully immersed in the tissue. Indusia and its relatives.
common in that subgenus, and trichomes are eto), Rio Mazan, J. Schunke 300 (F, GH, uc, us, USM).
long, 2-9 cm broad, subsessile. Lamina broadly segment tissue. Indusia tubular, neither bilabiate
oblong, pinnatisect, the rachis, costae and (some- nor dark-margined, mouth scarcely to slightly ex-
times) veins and margins abaxially provided with panded, receptacle commonly long-exserted.
simple, reddish brown trichomes that adhere to
tree trunks, free portion of larger segments 2.8-
4.5 cm long and 0.8-1.2 cm
broad, entire to shal- In forests, commonly hemiepiphytic, tightly ap-
lowly crenate. Venation catadromous, veins free, pressed to tree trunks, sea level to 1 000 m, Loreto,
simple, false veins lacking, but often some very Huanuco, Pasco, Cuzco.
short,darkened lines scattered between the veins. The Guianas to Colombia, south to Brazil and
Sori few to several on the pinna-lobes, not or Bolivia.
scarcely immersed in the segment tissue. Indusia This, Trichomanes tuerckheimii, and T. tanai-
fil. 2, t. 201. 1831. TYPE: Guyana, Demerara, study may prove them all to be conspecific.
Ankers (holotype, K; photo, us).
Stem long-creeping, relatively stout and wiry, Loreto: Mishuyacu, near Iquitos, Klug 149, 1135 (F,
0.5-1.2 mm in diameter, sparsely to
abundantly us). Sierra delPongo, Mexia 6289 (GH, MO, uc, us).
Huanuco: 5 km NE of Tingo Maria, Stork & Horton
provided with brown rhizoids that extend onto the 9518 (F, GH, uc, us). Pasco: Prov. Oxapampa, Valle del
leaf axes and adhere to tree trunks. Leaves sub-
Palcazu, Leon 706 (F, GH). Cuzco: Prov. Quispicanchi,
distant to remote, larger ones 5-18 cm long, near Inambari, Vargas 16460 (GH).
90 FIELDIANA: BOTANY
22. Trichomanes bicorne Hooker, Icon, pi., t. 892. Trichomanes elegans Rudge, PI. Guian. 24, /. 35. 1805,
1854. LECTOTYPE (designated here): Brazil, nom. illeg. (not L. C. Rich., 1792, or Poiret, 1808).
Hvmenostachys diversifrons Bory, Diet, class, hist. nat.
Amazonas, Barra do Rio Negro, Spruce 1 178 8: 462. 1825. TYPE: "Guiane," Poiteau (holo-
(lectotype, K!; isolectotype, us!). LECTO- type, P?; isotype, L; photos, GH & us of L).
PARATYPE: Brazil, Amazonas, Sao Gabriel, Feea diversifrons (Bory) Copel., Phillip. J. Sci. 67: 74.
1938.
Spruce 2334 (K!, us).
Ptilophyllum bicorne (Hooker) Prantl, Unters. Morph. Stem stout, erect. Leaves crowded to caespitose,
Gefasskrypt. 1: 48. 1875. 10-35 cm long, strongly dimorphic, sterile ones
elliptic or lanceolate, deeply pinnatisect, short-pet-
Stem stout, decumbent or erect, abundantly iolate, fertile ones commonly longer, linear, sub-
provided with lustrous, castaneous, pluricellular lamina (2-)3-8 cm
entire, long-petiolate. Sterile
trichomes. Leaves monomorphic, crowded or broad, deeply divided into subfalcate segments 4-
caespitose, 3-10 cm long, 3-4-pinnatisect, the axes 7 mm
broad, the rachis abaxially provided with
(and sometimes veins) sparsely provided on the dark, simple trichomes and often flagellate and
abaxial side with simple, castaneous, pluricellular proliferous at the tip. Venation catadromous, a few
trichomes. Petiole 0.5-4 cm long, conspicuously to many veins anastomosing toward the segment
alate throughout, the alae broader than the petiole margin, 4-8 of them issuing from the rachis be-
quite or nearly to base. Lamina ovate, the axes tween adjacent costae, false veins lacking. Fertile
broadly alate, alae and segments often undulate. lamina simple, essentially entire, with sori ar-
Venation catadromous, veins free, false veins lack- ranged in a nearly continuous line along each mar-
ing. Sori immersed in the segment tissue. Indusia gin, fully immersed in tissue in the forks of veins.
short-tubular, borne in the forks of veins, each of Indusia subconical, not bilabiate, the mouth nei-
which extend well beyond the indusial mouth, the ther flaring nor recurved, strongly indented be-
mouth not dark-margined, receptacle long-exsert- tween the vein tips, receptacle short- to long-
ed. exserted at maturity.
Engler & Prantl, Nat. Pflanzenfam. 1(4): 108. Feea trollii (Bergdolt) Vareschi, Flora Venezuela 1 :
gellate and proliferous at the tip. Venation catad- commonly 1 -forked, false veins lacking. Fertile
romous, a few to many veins anastomosing toward leaves bearing numerous, stalked or subsessile, as-
the segment margin, 2(-3) of them issuing from cending sori on each side of the primary axis. In-
the rachis between adjacent costae, false veins dusia urceolate, scarcely or not bilabiate, the mouth
lacking. Fertile lamina simple, essentially entire, not flaring, receptacle strongly exserted at matu-
with sori arranged in a nearly continuous line along rity.
each margin, fully immersed in tissue in the forks
of veins. Indusia subconical, not bilabiate, the Thus far known in Peru from a single collection,
mouth very slightly flaring and recurved, but not roadside along a sandy, rocky stream bank, 350-
indented between the vein tips, receptacle strongly 700 m, Amazonas.
exserted at maturity. Elsewhere terrestrial or epipetric, commonly near
banks of streams or rivers, 250-800 m; Panama;
Terrestrial in dense, wet forests, often in wet the Guianas and Colombia.
ravines or along stream banks, 350-850 m, Ama- The small size, strongly dimorphic leaves, and
zonas to Pasco. nonfoliaceous fertile leaves easily distinguish this
Surinam to Colombia; Peru; Bolivia. from others in the genus.
This is closely related to the more robust Tri-
chomanes diversifrons. Besides the characters in Amazonas: Prov. Bagua, ca. 40-43 km NE of Chiriaco,
the key the latter can usually be recognized by its Barbour 4524 (MO, USM).
broader lamina (to 8 cm) and segments (4-7 mm),
these being separated from the rachis by a con-
spicuous wing ca. 2 mm
broad on each side. The 26. Trichomanes pinnatum Hedwig, Fil. gen. sp.,
lamina of T. trollii is rarely more than 4 cm broad 1799. LECTOTYPE
/. 4,f. 1. (designated by
and are cut nearly to the rachis into segments which Flora Lesser Antilles. 89.
Proctor, 1977):
are commonly 3 cm broad or less. Additionally,
Hedwig, /. 4,f. 1, based on specimen allegedly
the fertile lamina of T. diversifrons has dentate
from Jamaica. Figure 12c.
margins, due to the apices of the indusia which
are deeply indented between the vein forks. In-
Trichomanes pennatum Kaulf., Enum. fil. 264. 1824.
dusial mouths of T. trollii are typically flush with TYPE: French Guiana, without collector (holo-
the vein tips, so that the leaf margins are essentially type, LZ destroyed).
entire throughout. Neurophyllum pinnatum (Hedwig) Presl, Hymeno-
phyllaceae 19. 1843.
92 FIELDIANA: BOTANY
except connected at the tips by a continuous mar- proliferous at the tip, the tissue relatively thick,
ginal vein, false veins copious and perpendicular commonly obscure, glabrous, pinna margins ser-
to true veins. Sori arranged in a nearly uninter- rate, the serrations obtuse to subacute. Venation
rupted line along pinna margins at the tips of veins, catadromous, the veins free, a lateral marginal vein
not immersed in the tissue, often stalked. Indusia faint and discontinuous or (more commonly) lack-
narrowly tubular, not or scarcely bilabiate nor dark ing, false veins lacking or a few very rare and faint
margined, mouth not flaring, receptacle long- ones perpendicular to true veins. Sori arranged in
exserted. a nearly uninterrupted line along pinna margins
at tips of veins, not immersed in the tissue, fre-
Ptilophyllum hostmannianum (Klotzsch) Prantl, Un- hostmannianum from species in other sections of
ters. Morph. Gefasskrypt. 1: 49. 1875. the subgenus; yet it serves to further establish its
Feea humboldtii Bosch, Ned. Kruidk. Arch. 4: 347. 5-25 cm cm broad, moderately to
long, l-2(-2.5)
1859, nom. nov., based on Trichomanes hetero- amply pubescent, the trichomes delicate, un-
phyllum Willd. and with the same type. branched, unicellular (or rarely pluricellular) be-
Feea heterophylla Copel., Philipp. J. Sci. 67: 73. 1938,
nom. nov., based on Trichomanes heterophyllum yond a short, enlarged, basal cell. Petiole nonalate,
Willd. and with the same type. or narrowly so toward the lamina. Lamina linear,
cut nearly or quite to the rachis into deeply lobed
to pinnatifid, patent, pinnae, gradually reduced to
long- to short-creeping. Leaves deep-
Stem wiry, a pinnatifid apex, lamellae lacking on veins. Ve-
ly pinnatisect to nearly pinnate, subdistant or re- nation catadromous, the veins free, false veins
mote, to 22 cm long, dimorphic, sterile ones short- on the apical
lacking. Sori terminating the veins
petiolate, to 3 cm broad, fertile ones long-petio-
part of the pinnae. Indusia tubular, deeply im-
late. far exceeding the sterile ones and 1 cm broad mersed in the segment tissue or, if partially im-
or less, the petiole essentially nonalate, sparsely or mouth
mersed, broadly alate on each side, the
amply provided with castaneous, pluricellular tri- bilabiate and somewhat flaring, the lips not dark-
chomes. Sterile lamina oblong or oblong-elliptic,
margined, receptacle exserted.
commonly equaling or longer than the petiole, the
apex pinnatifid. Venation catadromous, veins free
Apparently known thus far from two collections
and dichotomously branched, false veins lacking. on tree trunks, ca.
in Peru, in damp elfin forest,
Fertile lamina linear, much shorter than the pet-
1850 m, Huanuco. Elsewhere epiphytic or epi-
iole, with a few sori terminal on each pinna and
700-2300 m.
petric,
fully immersed in the tissue. Indusia subconical West Indies; Costa Rica; Venezuela; Colombia;
or broadly tubular, not bilabiate, the mouth not
Ecuador; Peru.
(or scarcely) flaring nor dark-margined, the recep-
Trichomes in many species of subg. Achomanes
tacle long-exserted.
are pluricellular, with two to several greatly elon-
gate cells, these often springing from a very short,
Terrestrial, in open to dense forests, commonly widened basal cell. In species most closely related
in wet, sandy places, sea level to 1400 m, San to Trichomanes crinitum, these are often mixed
Martin, Loreto. with a number of unicellular (above the basal cell)
Venezuela; Colombia; Amazonian Peru and trichomes. However, T. crinitum differs from its
Brazil.
Peruvian relatives in having all, or nearly all, of
There has been some confusion with nomen- its laminar trichomes unicellular above the basal
clature of this species. Lellinger's (1984) citing of
cell.
Trichomanes humboldtii "Bosch" (1858) was in
error, since therewas no such name; instead, it SW
Huanuco: slope of Rio Llullapichis watershed, on
was "Feea humboldtif that Bosch had published. the ascent of Cerros del Sira, Dudley 13525 (GH), 13545
However, Lellinger's use of 'T. humboldtii"'' is le- (GH, us).
Woytkowski 6211 (GH, MO, us). Loreto: Prov. Maynas, losum (sensu Martius in part, Icon. pi. crypt.
Rio Nanay, Mishana, Foster & Foster 4103 (GH); 104, t. 68 right) presumably based on "Brazil,
94 FIELDIANA: BOTANY
Arara-coara" (Araracuara, on Rio Caqueta, Iquitos, Klug 196, 51 1 (F, us). Prov. Maynas, 10 km S
of Iquitos, Tryon & Tryon 5179 (GH, us). Timbuchi on
now Colombia), Martius (holotype, M; iso-
Rio Nanay, LI. Williams 957 (F, us).
types, BR, L; photo, us of L).
On tree trunks in forests, or on clay banks along margined, sometimes apparently bilabiate due to
the narrow wings of tissue extending beyond the
roads and streams, sea level to 150 m, thus far
mouth on two sides, receptacle commonly exsert-
found only in Loreto, but locally common there.
ed.
to Colombia; Peru; Amazonian Brazil.
Surinam
The shape of the indusium is somewhat vari-
able. The mouth is commonly truncate and not at and wooded ravines, pendent from
In forests
all wings of tissue along
bilabiate, but frequently the trees or in sphagnum on sandy
wet rock crevices, or
the sides extend beyond the mouth, giving it a banks, 2150-3500 m, Cajamarca, Amazonas,
"horned" appearance, much like Trichomanes bi- Huanuco, Pasco, Cuzco.
corne. This a distinctive species in Peru, because
is Jamaica; Costa Rica; Venezuela and Colombia,
of the crestlike lamellae borne on many of the south to Bolivia; Brazil.
veins on the abaxial side, perpendicular to the With its l-(2-)pinnate-pinnatifid leaves and the
plane of the lamina. These processes are similar copious, long trichomes that often obscure the
to those occurring in several species of Hymeno- lamina surface, this should not be mistaken for
phyllum. On some of the more densely hirsute any other species of subg. Achomanes.
specimens of T. martiusii, the lamellae are par-
tially obscured by the indument and thus they Cajamarca: Prov. Jaen, Paso de Huascarai, 15 km SE
have been mistaken for other species in subg. of Huancabamba, Fosberg 27842 (us). Amazonas: Prov.
Achomanes, particularly T. pilosum Raddi, of Bo- Chachapoyas, Cerros de Calla Calla above Leimebamba,
Hutchison & Wright 5568 (F, GH, MO, uc, us). San Mar-
livia and southern South America. tin: Prov. Mariscal Caceres, Rio Abiseo National Park,
the trichomes unbranched, light to dark brown, patent (or the basal ones deflexed), lacking lamel-
those of the veins and margins unicellular beyond lae, the margins subentire to crenate or serrate,
a very short basal cell, those of primary axes 1-3- often undulate. Venation catadromous, the veins
celled beyond the basal cell. Petiole rather con- free, false ones lacking. Sori borne near and at
spicuously alate halfway or more to the stem. Lam- segment apices, at tips of veins, sometimes flanked
ina narrowly or broadly lanceolate, gradually re- by vein forks. Indusia tubular or narrow-conical,
duced to a pinnatifid apex, cut nearly or quite to deeply to fully immersed in the segment tissue,
the rachis into adnate, linear or linear-oblong, pat- the mouth neither bilabiate nor dark-margined,
ent pinnae (or basal ones deflexed), their margins not or scarcely flaring, receptacle exserted.
96 FIELDIANA: BOTANY
mosum can usually be distinguished from T. cris- to slight projections of tissue along the flanking
tatum Peru) by lamina shape: that of
(at least in vein forks, receptacle exserted.
the latter is commonly linear, rarely more than 4
cm broad, whereas the lamina of T. plumosum is Terrestrial, or occasionally on tree trunks, com-
typically broadly lanceolate, usually more than 4 monly in wet forests on sandy soil or in open
cm broad. sphagnum bogs, sea level to 2000 m, San Martin
and Loreto to Puno.
San Martin: Tarapoto, Spruce 4766 (BR, w). Huanuco: Venezuela and Colombia, south to Brazil and
SW slope of Rio Llullapichis watershed, ascent of Cerros Argentina.
del Sira, Dudley 13051 (GH), 1 3128 (GH, us), 1 3512 (GH). This species very easily confused with Tri-
Pasco: Prov. Oxapampa, 1 9 km W of Oxapampa, D.
is
Smith 2209 (MO). Junin: La Merced, Macbride 5634 (F, chomanes plumosum; characters are discussed
GH). Cuzco: La Convencion, Biies 2028, 2140 (us). Madre above in the treatment of the latter. Trichomanes
de Dies: Prov. Manu, Shintuya, Chavez 804 (MO). Puno: cristatum and T. plumosum share nearly the same
Prov. Carabaya, San Gaban, Vargas 18875 (GH). Dept.
distribution and are often found growing together.
Unknown: "ad saxa humida prope Sangari," Lechler 2548
In Peru, at least, there are frequent intermediates,
(BR, K, w 3 sheets); although this is the type number of
T. undulatum (= T. vandenboschii), these specimens rep- which indicates a strong probability of hybridiza-
resent part of a mixed collection). tion. These problems are not confined to species
of Peru but also occur in other taxa of the group
of T. crispum L. throughout the Neotropics, as
pointed out by Windisch (1988). Obviously there
34. Trichomanes cristatum Kaulf., Enum. fil. 265. is a great need for careful analysis of plants in situ
1824. TYPE: Brazil, collector undesignated as well as further caryological study before ac-
Trichomanes sellowianum Presl, Hymenophyllaceae San Martin: Rio Negro, Woytkowski 6210 (GH, MO,
15, 37. 1843. TYPE: "Brasilia," Sellow 197 (ho- uc, us). Loreto: Prov. Maynas, Dist. Iquitos, Rio Nanay,
lotype, PR?; possible isotypes, 2 unnumbered Sel- McDaniel & Rimachi 18924 (F, MO). Huanuco: Tingo
low sheets from "Brasilia," B!, K!). Maria (as San Martin), Allard 21585 (GH, us), 27559
(us). Pasco: Prov. Oxapampa, Palcazu, van der Werff
el at. 8376 (MO, uc). Junin: E of Quimiri Bridge near La
Stem short-creeping to decumbent, rarely erect.
Merced, Killip & Smith 23952 (F, us). Cuzco: Prov. La
Leaves monomorphic, subdistant to contiguous, Convencion, Valle de Santa Ana, Herrera 3008 (us).
mature ones 15-50 cm long, 2-4(-5) cm broad, Puno: Prov. Sandia, Vargas 1 1832 (GH).
moderately to rather densely pubescent, the tri-
apices, at tips of veins, often flanked by the vein from Peru need to be examined carefully to de-
forks. Indusia urceolate, fully immersed in the seg- termine if T. accedens truly occurs in Peru.
ment tissue, the mouth somewhat flaring, not dark-
Trichomanes delicatum Bosch, Ned. Kruidk. Arch.
margined, not or scarcely bilabiate, receptacle
exserted at maturity. 5(2): 145. 1861. TYPE: Ecuador, Quito, Cum-
ing 21 (holotype, B; frag., L).
This is closely related to Trichomanes cristatum, margins of the indusia. It is to be expected in Peru,
since it has been found in Colombia, Ecuador, and
from which it is distinguished primarily by its
Bolivia.
smaller size and relatively broader lamina. Besides
the key characters, it can usually be separated from Trichomanes haenkeanum Presl, Hymenophylla-
both T. cristatum and T. plumosum by its some- ceae 15, 36, 65. 1843. TYPE: Peru, mountains
what flaring indusium mouth. That of the other of Huanuco, Haenke (holotype, PR?).
two species is scarcely or not at all expanded. The
Trichomanes crispum var. haenkeanum C. Chr., In-
confused identities of species within this complex
dex fil. 641. 1905.
is illustrated by the fact that the number of the
In the protologue, Presl indicated a close rela-
type collection is shared with specimens of T. plu-
mosum, as noted above in the citation of types. tionship with Trichomanes crispum. Characters
used in the description certainly align T. haen-
Cuzco: Maranura, Beatriz, Bites 895 (us). Prov. La keanum with this group of species but are insuf-
Convencion, Sahuayaco, Vargas 6288 (GH). Prov. La ficient to place it precisely. Until the type is located
Convencion, Choquello, Vargas 8184 (uc). Prov. Pau- and compared with other species in the complex,
cartambo, Atalaya, Valle Kosnipata, Vargas 23155 (OH).
nothing more can be determined.
98 FIELDIANA: BOTANY
a more or less urceolate indusium, short-stalked glaucous abaxially. Petiole dark brown to atro-
(ca. 6 rows of cells), with an oblique annulus not purpureous, sublustrous. Lamina subdeltoid,
interrupted by the stalk. sparsely to amply provided on surface, veins and
costules with light to dark brown, tortuous, septate
This small family contains two genera: Loxoma trichomes, glabrous adaxially. Pinnae commonly
of New Zealand, and the Neotropical Loxsomop- well-spaced, subdeltoid, somewhat or slightly re-
sis. duced at base, with pinnules strongly ascending,
segments and venation catadromous. Veins pin-
nately branched, prominulous. Indusia urceolate
I. Loxsomopsis to (especially at maturity) narrow-cyathiform.
Loxsomopsis Christ, Bull. Herb. Boissier II. 4: 399. Scandent or trailing, or arching to pendent from
1904. TYPE: Loxsomopsis costaricensis clay banks, in on exposed ridges,
elfin forests, often
nearly 3-pinnate. Lamina subcoriaceous, glabrous & Brune 279, Costa Rica) has not been seen, but
adaxially, glabrous to pubescent abaxially. Veins the original description and illustrations suggest
free. Sorus marginal, paraphysate, the indusium the two species are synonymous. Loxsomopsis no-
narrow-cyathiform to urceolate, the rim entire, the tabilis Slosson (type from Bolivia, R. S. Williams
elongate receptacle exserted. Spores trilete, glo- 1303) is supposedly distinguished by its glaucous
bose-tetrahedral, the surface coarsely tuberculate. abaxial surface and by the larger pinnae being
greatly reduced at base. Two sheets of isotype (us)
The genus
is probably represented by a single seem to illustrate this clearly, and yet another is-
species. However, two very closely related species otype (P) is only slightly glaucous and pinnae are
have been described (from Costa Rica and Bolivia, not very strongly reduced at base. Lack of consis-
respectively), based on supposed differences in size tency in all these features suggests that there is but
of leaf, color of petiole, and shape of pinnae and one species of Loxsomopsis.
indusia. Two others have been separated on the
basis of other variable and doubtfully significant Huanuco: SW
slope of Rio Llullapichis watershed, as-
characters. Observations of a number of cent of Cerros del Sira, Dudley 13420 (GH). Playapampa,
speci-
Macbride4521 (F, GH. us). Pasco: Prov. Oxapampa, Cord.
mens throughout the range suggest that such char-
San Gutardo, Ledn 532 (USM). Cuzco: Valle San Miguel,
acters vary with age of the plant and maturity of
La Convention, Bues 2119 (us). Prov. La Convention,
sori, and they demonstrate no significant corre- Cordillera de Vilcabamba, Dudley 10713 (GH, MO).
lation.
FIG. 14. Plagiogyria semicordata: a-b, habit; c, apex of fertile leaf; d, apex of fertile pinna, (a from Cuatrecasas
5465, Colombia, F, b from Wurdack 1159, F, c-d from Little 9184, Colombia, F.)
a. Lamina 4-5-pinnate-pinnatifid, broadest at the base, its tertiary axes grooved on the adaxial side
I. Culcita
a. Lamina 2-3-pinnate-pinnatifid (in America), reduced at the base, its tertiary axes ridged (not grooved)
on the adaxial side . II. Dicksonia
m tall, stout. Leaves monomorphic, covering of trichomes at the base, often longer than
erect and to 3
the lamina. Lamina deltoid, to 4- or 5-pinnate-
to ca. 3 m long. Lamina 4-5-pinnate or slightly
pinnatifid at the base, the major segments stalked,
more complex, the tertiary axes grooved adaxially,
thinly pubescent, or glabrous with age, sterile ul-
veins free. Sori marginal, the adaxial indusium
timate segments with bluntly acute lobes. Sorus
joined basally to the thinner abaxial indusium, or
single on each ultimate lobe.
the two separate. Spores tetrahedral-globose, tri-
In wet elfin forests at ca. 2400-3100 m, Ama-
lete, nearly smooth, rugulose, or tuberculate.
zonas south to Cuzco.
Southern Mexico, Central America, Greater
Culcita coniifolia of tropical America and C.
Antilles, Venezuela and Colombia south to Peru;
macrocarpa of the Canary Islands, Azores, and
Mt. Itatiaia, Brazil.
Spain form subg. Culcita. The other five species, and
Culcita coniifolia is a distinctive species
of Malaysia to Samoa and Australia, comprise the
should be mistaken for no others. Although of
subgenus (or, better, the genus) Calochlaena Max-
wide distribution, it has been rarely collected in
on.
Peru.
a. Tertiary segments of the larger fertile pinnae each with few to several sori 1 D. sellowiana.
a. Tertiary (ultimate) segments of the larger fertile pinnae each with a single sorus ... 2. D. stuebelii
1. Dicksonia sellowiana Hooker, Sp. til. 1: 67. In wet woods or cloud forests, at 1 550-2400 m,
1844. TYPE: Brazil, Sellow (lectotype desig- Cajamarca and Amazonas, south to Cuzco.
nated here, K; isolectotype, HBG; photos, GH, Southern Mexico, Central America, Venezuela,
us of HBG; lectoparatype, Brazil, Miers, K). and Colombia south to Bolivia, Paraguay, Uru-
Figure 16. guay and southeastern Brazil.
Dicksonia sellowiana is a widespread and vari-
Balantium karstenianum Klotzsch, Linnaea 20: 444. able species. Several segregates are sometimes rec-
1847. TYPE: "Columbia," Karsten, no. 9 (Coll.
ognized but these evidently represent minor vari-
II) (holotype, B?; isotype, HBG; photo, GH of HBG).
ations. The other American species are D. stuebelii
Dicksonia gigantea Karsten, Fl. columb. 2: 1 77, 1. 193.
1 869. TYPE: Colombia, (Cundinamarca), Andes
of Peru and D. berteriana (Colla) C. Chr. of the
of Bogota, Guadeloupe, Karsten (not located) Juan Fernandez Islands.
(Karsten, HBG; photo, GH, may be authentic).
Dicksonia karsteniana (Klotzsch) Moore, Index fil.
Cajamarca: Prov. Cutervo, La Pucarilla, Ldpez & Sa-
190(1860), 313(1861). gdstegui 5457 (GH, HUT). Amazonas: Prov. Bagua, ca. 20
Dicksonia spruceana Kuhn, Linnaea 36: 153. 1869. km E of La Peca, Harbour 2870 (MO). Huanuco: Cerros
TYPE: Peru, (San Martin), Tarapoto, Spruce 4728 del Sira, Rio Llullapichis watershed, Dudley 13254 (F,
(holotype, B; isotypes, A!, BM, GH!; photos, GH, GH, MO), 75259, 13373 (GH). Pasco: Oxapampa (as Jun-
MO, uc of BM). in), Soukup 2334 (F, GH). Prov. Oxapampa, Canyon de
Huancabamba, Ledn 614 (F). Cuzco: Prov. La Conven-
Stem to 10 m tall, enclosed, at least basally in tion, Cordillera Vilcabamba, Dudley 10439, 10603,
Lophosoriaceae Pic.-Ser., Webbia 24: 700. 1970. Lophosoria quadripinnata is a widely distrib-
TYPE: Lophosoria Presl.
uted and distinctive species of the cloud forest
zone. One high altitude variety that is local in
Stem sometimes branched, massive, decumbent Ecuador and Peru may be recognized.
to erect and arborescent, indurated, densely cov-
ered by long trichomes. Leaves usually large, ca.
1-5 m
long, circinate in vernation, monomorphic,
1. Lophosoria quadripinnata (Gmelin) C. Chr.,
more or Skottsb. Nat. hist. Juan Fernand. 2: 16. 1920.
pinnate, less pubescent. Petiole lacking
stipules, not articulate to the stem. Veins free. Sori
exindusiate, on the abaxial surface of the segments, Stem apex with few leaves. Petiole about as long
as the lamina or somewhat shorter, with 3 con-
paraphysate with slender trichomes. Sporangia
with a short, 6-rowed stalk and a complete, oblique voluted vascular bundles. Lamina very narrowly
annulus. lanceolate to usually broadly ovate, to 3-pinnate-
pinnatisect, with sessile or short-stalked segments,
The family Lophosoriaceae contains a single,
very sparsely to densely pubescent and often glau-
American genus. cous abaxially, ultimate segments obtuse to sub-
acute. Sori single on a fertile vein.
Key to Varieties
a. Leaf ca. 2-4 rn long, pinnae spaced, patent, the central ones ca. 0.5-1.0 m long
la. var. quadripinnata
a. Leaf ca. 0.25-1 m long, pinnae imbricate, ascending, the central ones ca. 0.05-0.20 m long
. Ib. var. contracta
1 b. Lophosoria quadripinnata var. contracta (Hi- Metaxya Presl, Tent, pterid. 59. 1836. TYPE: Me-
eron.) R. & A. Tryon, Rhodora 84: 1 26. 1 982. taxya rostrata (HBK.) Presl (Aspidium rostra-
turn HBK.). Figure 18.
Alsophila contracta Hieron., Hedwigia 45: 236, /. 75,
/ 8. 1906. SYNTYPES: Peru, (Amazonas), near Terrestrial or rarely on tree bases. Leaves to ca.
Inez and Calle-calle, Stitbel 1067 (B!; frag., GH!);
(Amazonas), near Challuayacu and Tambo Cen- 2 m long. Petiole usually with short trichomes at
tamala, Stubel 1066 (B!). the very base. Lamina 1 -pinnate, with simple pin-
nae, veins free. Sori roundish to elongate, the re-
Wet, shrubby areas, 2800-3500 m, in Amazon- ceptacle nearly flat. Spores globose, trilete, gran-
as and La Libertad. ulate.
Ecuador and Peru.
The var. contracta is evidently a high-altitude A monotypic genus ranging from southern Mex-
ecotype. The following collection is intermediate icoand Central America; Guadeloupe; Trinidad;
between the two varieties: Peru, Amazonas, Puma- and French Guiana west to Colombia and south
Urcu, SE of Chachapoyas, Wurdack 791 (us). to Bolivia; Amazonian Brazil.
Woods and dense forests, usually in sandy, moist, importance are characters of the petiole scales.
well-drained soil, 100-800 m, Amazonas to Cuzco However, the scales may be abraded on the petiole
of a mature leaf and their characters are best seen
and Madre de Dios.
on the croziers. The height and diameter of the
Range of the genus.
arborescent stem vary in a species depending upon
Metaxya is unusual in having the 1-pinnate-
the age of the plant and the growing conditions.
pinnatifid leaves of juvenile plants more complex
than the 1 -pinnate leaves of adult plants.
Accordingly, these characters as well as other
quantitative ones are usually not included in the
Amazonas: Prov. Bagua, Rio Maranon, above Cas- species descriptions.
cadas de Mayasi, Wurdack 1872 (us, USM). San Martin: Several special terms that are used in the Cy-
Prov. Mariscal Caceres, Isla de Pucunuchu, J. Schunke atheaceae are the following.
4783 (F, GH). Prov. Mariscal Caceres, Rio Sion, J. Schunke PETIOLE SPINES Corticinate: The spine is an ex-
3524 GH, us). Loreto: Iquitos, Killip & Smith 27489
(F,
tension of the cortex of the petiole; each bears, at
(GH, us). Huanuco: Prov. Pachitea, Bosque Nacional de
least when young, a scale at its apex which is rather
Iparia, J. Schunke 1620, 1833 (F, GH, us). Pasco: Prov.
Oxapampa, Cordillera San Matias, Leon 332 (USM). blunt when the scale falls off. Squaminate: The
Ucayali: vicinity of Aguaytia, Croat 20966 (us). Prov. spine is not an extension of the petiole; it breaks,
Coronel Portillo, Bosque Nacional Humboldt, Vdsquez or can be broken, from the petiole at its base. It
3910 (F, MO). Cuzco: Prov. Quispicanchi, Vargas 16462
has a sharp apex that does not bear a scale. These
(GH). Prov. Paucartambo, Vargas 1 1245 (GH). Madre de
Dios: Prov. Tambopata, Tambopata Nature Reserve, spines have evidently evolved from petiole scales.
Barbour5191 (F). PETIOLE SCALES Conform: All cells, except those
that may be borne on the edge, are similar in ori-
rounding the receptacle, nearly flat or with slightly TRYON, R. M., AND A. F. TRYON. 1982. Cy-
upturned edges. Cyathiform: Cup-shaped. Urceo- atheaceae, pp. 166-212, in Ferns and allied
late: Urn-shaped. Subsphaeropteroid: Enclosing plants, Springer- Verlag, New York.
c. Petiole with squaminate spines, many of these large, black, with a sharp apex; petiole scales
patent, attached at a usually narrowed base; croziers with large, black, sharp spines
III. Nephelea
b. Petiole scales lacking a differentiated apical seta, the apex rounded to filamentous d
d. Indusium absent IV. Trichipteris
d. Indusium present e
e. Spores lacking large pores, sometimes with variously distributed small pits or pores; veins
free, the basal ones of adjacent segments extending to the margin above the sinus, rarely a
few costal areolae present V. Cyathea
e. Spores with three large equatorial pores, often also with smaller pits or pores; veins anas-
tomosing to form costal areolae, or if free then the basal veins of adjacent segments connivent
to the sinus . VI. Cnemidaria .
portion of petiole scale, (a from Soukup 1651, F, b from Soukup 2336, F, c from Plowman 6062, F.)
a. Petiole scales without a dark apical seta and without dark marginal setae, rarely the margins ciliate
2. S. rufescens
Costa and costules lacking scales or with scales various, but not whitish, fimbriate, or
e.
b. Veins regularly anastomosing, especially along the costa, sori with a hemitelioid indusium ....
4. S. bradei
Petiole scales with a dark apical seta and dark marginal setae [subg. Sphaeropteris] f
f. Sori exindusiate 6. S. elongata
f. Sori with a sphaeropteroid indusium 7. S. quindiuensis
1 .
Sphaeropteris aterrima (Hooker) Tryon, Contr. 2. Sphaeropteris rufescens (Kuhn) Windisch,
Gray Herb. 200:20. 1970. Bradea 1: 372. 1973.
Ahophila aterrima Hooker, Syn. fil. 38. 1866. TYPE: Hemitelia rufescens Kuhn, Linnaea 36: 159. 1869.
Peru, (San Martin), Tarapoto, Spruce 4713 (ho- TYPE: Peru, (San Martin), Monte Guairapurima,
lotype, K!; isotypes, P!, us!). Tarapoto, Spruce 4727 (holotype, B!; isotype, K.;
Cyathea aterrima (Hooker) Domin, Pterid. 262. 1 929. photo, GH of K).
Cyathea rufescens (Kuhn) Domin, Pterid. 264. 1 929.
San Martin: Monte Morro de Moyobamba, Stiibel 3. Sphaeropteris macrosora (Baker) Windisch,
/77J(B, NY). Bradea 1: 372. 1973.
At 400-500 m, Pasco.
and finely ciliate, especially toward the apex. Lam-
ina 2-pinnate-pinnatifid to 2-pinnate-pinnatisect,
Guyana and Brazil, west to Colombia and Peru.
pinnules with prominent whitish, flattish scales
Sphaeropteris macrosora represented in Peru
is
abaxially, veins free. Sori with paraphyses about
by var. reginae Windisch which occurs throughout
as long as the sporangia, indusia sphaeropteroid.
the range of the species. Variety macrosora is in
Guiana and Venezuela and var. vaupensis
British
This species has been collected in Peru only at
Windisch is in Colombia and Venezuela.
ca. 3000-3300 m
in the Department of Ama-
zonas.
Pasco: Prov. Oxapampa, Palcazu, Foster & d'Achille
10168 Ecuador and Peru.
(F).
Sphaeropteris atahuallpa is a distinctive and at-
tractive species. The
croziers are completely en-
veloped by large, whitish scales and these form a
4. Sphaeropteris bradei Windisch, Bradea 1: 372. dense covering to the petiole. Some of the scales
1973. TYPE: Colombia, Vaupes, Cerro Mitu, cm
are up to 6 long and several cells thick at their
Schultes, Raffauf& Soejarto 2422 9 (holotype, base.
GH!).
Amazonas: Summit of Puma-Urcu, SE of Chachapoy-
Petiole sparingly aculeate, with rather broad as, Wurdack 1153 (GH, us). Prov. Chachapoyas, Calla-
scales, these usually dense, especially toward the Calla, Smith & Vasquez 5012 (GH).
3 mm
FIG. 20. Alsophila engelii: a, pinna; b, ultimate segment and son; c, apex of petiole scale. (From Dudley 11944,
OH.)
Karsten, not Alsophila elongata Hooker. Fig- Petiole smooth to tuberculate, brown, bearing
ure 20. mostly blackish to light brown scales with 1 apical
seta. Lamina 2-pinnate-pinnatifid, gradually ta-
pering to the apex, rachis light brown to brown,
with usua "y numerous sma11 dissected scales, pin-
>
Cyathea elongata Karsten, Fl. Columb. 2: 1 59. 1 869.
TYPE: Venezuela, (Merida), Merida, Engel 138
nae sessile or nearly so. Indusia deeply cyathiform
(holotype, B!). to sphaeropteroid, glabrous.
Comments
Nephelea is a tropical American genus of 17
species, with three in Peru. It ranges from southern
Mexico and the West Indies, south to northern
Alsophila paucifolia Baker of Colombia and Ec-
uador may be found in Peru. It differs from A. Argentina and southern Brazil, centering in the
Greater Antilles where nine species occur.
engelii in having a 1-pinnate-pinnatifid lamina. Al-
The squaminate spines on the petiole are dis-
sophila capensis (L. f.) John Sm. ssp. polypodioides
(Sw.) Conant of southeastern Brazil may also occur
tinctive of Nephelea. They are blackish, have a
in Peru. It is characterized by the highly dissected very sharp tip, and at least on old leaves they may
be broken off at their base.
aphlebiae at the base of the petiole.
Reference
III. Nephelea
b. Indusium with whitish trichomes; pinna-rachises usually (sometimes narrowly) green-alate be-
tween the more distal, sessile pinnules; petiole scales on the abaxial side with dark lateral setae
2. N. cuspidata
a. Veins lacking indument abaxially, or present and not stellate; pinna-rachises green-alate between the
more distal, sessile pinnules; petiole scales lacking lateral setae, indusium with trichomes
.3. N. incana
1 .
Nephelea erinacea (Karsten) Tryon, Contr. Gray natifid, usually abruptly reduced to a pinna-like
Herb. 200: 40. 1970. apex, pinna-rachises not green-alate between the
more distal sessile pinnules, veins with minute,
Cyathea erinacea Karsten, Linnaea 28: 453. 1857. whitish, stellate squamules abaxially. Indusia cy-
TYPE: Venezuela, (Merida), Merida, 2000 m, athiform to urceolate or rarely subsphaeropteroid,
Karsten (holotype, not located; isotype, B!).
glabrous to densely scaly.
Alsophila erinacea (Karsten) Conant, J. Arnold Arbor.
64: 371. 1983.
apex of petiole scale, (a from Cuatrecasas 14843, Colombia, F, b, d from Davis 1109, Bolivia, F, c from Rimachi
481, F.)
Amazonas: Prov. Bongara, 5 km N of N end of lake 3. Nephelea incana (Karsten) Gastony, Contr.
Pomacocha, on road to Rioja, Hutchison & Wright 6803 Gray Herb. 203: 137. 1973.
(GH, uc, USM). Rio Cenepa, 10 km E of Huampami,
Berlin 219 (F, MO). Huanuco: Macbride 4843 (F, us).
Cyathea incana Karsten, Fl. Columb. 1: 75, t. 37.
Pasco: Prov. Oxapampa, trail to summit of Cordillera
1860. TYPE: Colombia, (Cundinamarca), Andes
Yanachaga, D. Smith et al 7817, 7851 (F, MO). Prov. of Bogota, 2500 m, Lindig(nol located).
Oxapampa, Rio San Alberto, Smith & Pretel 7942 (USM). Alsophila incana (Karsten) Conant, J. Arnold Arbor.
64: 371. 1983.
2. Nephelea cuspidate (Kunze) Tryon, Contr. Gray Scales on the abaxial side of the petiole with a
Herb. 200: 40. 1970. Figure 21. dark apical seta and without lateral setae, or rarely
with an additional apical seta and lateral setae.
Cyathea cuspidata Kunze, Linnaea 9: 101. 1834. Lamina rigidly papyraceous, 2-pinnate-pinnatifid,
TYPE: Peru, (Loreto), Prov. Maynas, Feb. 1831,
abruptly reduced to a pinna-like apex, pinna-rach-
Poeppig diar. 2286 (holotype, LZ destroyed; iso-
ises green-alate between the more distal sessile
types, B!, P; photo, GH of p).
Alsophila cuspidata (Kunze) Conant, J. Arnold Arbor. pinnules, veins glabrous abaxially, or with occa-
64: 371. 1983. sional trichomes. Indusia meniscoid to subsphaer-
opteroid, pubescent with rather soft and often cris-
Scales on the abaxial side of the petiole with a pate trichomes or trichomoid processes.
dark apical seta and usually with numerous, small-
er apical and lateral setae. Lamina usually papyr- In primary forests at ca. 800-2400 m, Caja-
aceous, predominantly 2-pinnate-pinnatifid, re- marca and Amazonas south to Junin and Ucay-
duced to a pinna-like apex, pinna-rachises green- ali.
alate between the more distal sessile pinnules, veins Colombia south to northwestern Argentina.
with trichomes or stellate squamules abax-
stellate
ially. Indusia rarely cyathiform, to urceolate to Cajamarca: Prov. Hualgayoc, Soukup 3810 (F). Prov.
sphaeropteroid, glabrous or with stellate indu- Cutervo, San Andres, Lopez et al. 6684 (GH, HUT). Ama-
ment. zonas: Laguna Pomacocha, NW
of Jumbilla, Soukup
2560a (GH). Pasco: Prov. Oxapampa, Oxapampa, Leon
502 (USM). Junin: Esposto 670 (USM). Prov. Tarma, ca.
In low-elevation rain forests and less often in 3 km SE of San Ramon, Iltis & Iltis 249 (GH, USM).
cloud forests, usually at 1 50-900 m, rarely to 2200 Ucayali: Prov. Coronel Portillo, Cordillera Azul, Young
m. Amazonas and Loreto south to Puno. & Sullivan 668 (F, MO).
a. Lamina 1 -pinnate-pinnatifid b
b. Pinnae sessile to short-stalked c
c. Petiole 15-30 cm long; lamina not or not much reduced at the base 10. T. pubescens
c. Petiole 3-6 cm long; lamina much reduced at the base 11. T. phegopteroides
b. Pinnae, especially below the lamina apex, very long-stalked 4. T. latevagans
a. Lamina 2-pinnate or more complex d
d. Lamina mostly 3-pinnate-pinnatifid; pinnules abundantly short-pubescent adaxially, or petiole
scales with a narrow, dark center stripe and broad white margins e
e. Leaf ca. 1.5 m
long, pinnules with many small, bullate scales abaxially, abundantly short-
pubescent adaxially; petiole scales nearly concolorous, brown with very narrow, lighter margins
7. T. flava
e. Leaf ca. 4 m
long or more, pinnules with a few flattish scales abaxially, glabrate or with a few
long trichomes adaxially; petiole scales with a narrow dark center stripe and broad whitish
margins 9. T. serpens
i. Petiole aculeate, pinnules usually sessile, basal veins end above a sinus, paraphyses
much shorter than the sporangia 1 . T. procera
i. Petiole muricate to tuberculate, pinnules stalked, basal veins connivent to a sinus,
paraphyses longer than the sporangia 13. T. lechleri
h. Fertile and sterile veins forked; ultimate lobes usually crenulate; paraphyses longer than
the sporangia 5. T. nigra
g. Abaxial and
lateral sides of the pinna-rachises usually densely pubescent with usually whitish
trichomes mostly 1.5-2 long mm 6. T. pilosissima
f. Lamina gradually reduced at the apex j
1. Pinnules with prominent, large, flattish scales abaxially; paramo and subparamo (elfin
forest) 8. T. frigida
mate) segments apically entire to coarsely toothed; central and basal pinnae
short- to long-stalked (rarely sessile) 6. T. pilosissima
n. Petiole scales dark brown to atropurpureous, usually with broad lighter mar-
gins, or whitish to light brown and concolorous; tertiary (usually ultimate)
segments apically crenulate or denticulate; central and basal pinnae sessile
o
o. Petiole with long trichomes, soon glabrous, the caducous trichomes leaving
a smooth petiole surface 14. T. conjugate
o. Petiole with long trichomes, these more or less persistent, when deciduous
leaving a hard base and a scabrous petiole surface 15. T. tryonorum
1 .
Trichipteris procera (Willd.) Tryon, Contr. Gray Petiole lacking trichomes, the scales dark brown
Herb. 200: 46. 1970. or with whitish margins. Lamina 2-pinnate-pin-
natifid, abruptly reduced at the apex, central and
basal pinnae short- to long-stalked, their pinnules
Polypodium procerum Willd., Sp. pi. ed. 4, 5: 206. sessile to short-stalked, abaxial and lateral sides
1810. TYPE: Brazil, Hoffmannsegg (holotype, B!,
of the pinna-rachises sparingly short-aculeate or
Herb. Willd. 19717).
Polypodium pungens Willd., Sp. pi. ed. 4, 5: 206. 1810, lacking spines, glabrate or with scattered to rather
asynonym of Trichipteris procera (Willd.) Tryon numerous brownish trichomes. Pinnules obtuse to
by Harrington, Contr. Gray Herb. 208: 23. 1978. acuminate, glabrate or with some small scales and
Alsophila procera (Willd.) Desv., Mem. Soc. Linn. Paris
(or) trichomes abaxially, ultimate segments obtuse
6: 319. 1827.
to subacute, fertile veins simple or rarely forked.
Alsophila dombeyi Desv., Mem. Soc. Linn. Paris 6:
320. 1827. TYPE: Peru, (Huanuco), Cochero, Paraphyses much shorter than the sporangia.
Dombey (holotype, Herb. Desv. P!; isotype, P!).
Alsophila infesta Kunze, Linnaea 9: 98. 1834. TYPE: In original rain forests or cloud forests, often in
Peru, (San Martin), Tocache, Rio Huallaga, Poep-
secondary forests or in other partly disturbed vege-
pig (holotype, LZ destroyed; isotype, B!).
Alsophila armigera Kunze, Linnaea 9: 98. 1 834. TYPE: tation, 100-1500 m, Amazonas and San Martin
Peru, (Huanuco), Ventanilla de Cassapi, Jul. 1829, south to Puno.
Poeppig (holotype, LZ destroyed; isotypes, MO!, P!; West Indies; South America south to northern
photos, GH, us of P). Peru and Bolivia.
Brazil,
Alsophila pycnocarpa Kunze, Linnaea 9: 97. 1834.
TYPE: Peru, (Huanuco), Pampayacu, Jul. 1829, Trichipteris dombeyi, although recognized by
Poeppig (holotype, LZ destroyed; isotype, Poeppig Barrington, is very close to T. procera and is here
201, B!). included within it. Trichipteris procera has an ex-
Alsophila peruviana Klotzsch, Linnaea 20: 441. 1847. tensive distribution and is quite variable as the
TYPE: Peru, (Junin), Tarma, Ruiz Herb. 66 (ho-
many synonyms described from Peru imply.
lotype, B!).
Alsophila nigripes C. Chr., Index fil. 45. 1905, nom. apex, pinnae and most of the pinnules long-stalked,
nov. for Alsophila melanopus Hooker (not Hassk.) pinna-rachises lacking spines, the abaxial and lat-
and with the same type. eral sides glabrous or with some short, brownish
Alsophila melanopus Hooker, Syn. fil. 37. 1866 (not trichomes. Pinnules acute to acuminate, glabrate
Hassk., 1855). TYPE: Ecuador, Chimborazo,
Spruce 5742 (holotype, K!; isotype, P!). abaxially, ultimate segments subobtuse, acute,
Cyathea nigripes (C. Chr.) Domin, Pterid. 263. 1929. acuminate or subapiculate, fertile veins simple or
forked. Paraphyses shorter than the sporangia.
Petiole lacking trichomes, the scales brown to
dark brown, nearly concolorous. Lamina 2-pin- In forests, 1200-1600 m, San Martin.
Amazonas: Prov. Bagua, Dist. Senepa, Tillett 672-115 In pajonal vegetation, 2200 m, San Martin.
(GH). San Martin: Prov. Mariscal Caceres, Puente Palo Colombia and Peru.
Blanco, Rio Tocache, Plowman et al. 1 1354 (GH). Loreto:
Above Pongo de Manseriche, Mexia6191 (F, GH). Huan-
The very long-stalked pinnatifid pinnae and the
uco: Hills E of Tingo Maria, Croat 21153 (MO). Ucayali: atropurpureous petiole and rachis are distinctive
Km 86 on Pucallpa-Tingo Maria road, Smith et al. 1 179 features of this species.
(GH). Cuzco: Prov. Paucartambo, Cosnipata valley,
Wachler et al. 221 (F, GH). Madre de Dios: Prov. Tam-
San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba, D.
bopata. SSW of Puerto Maldonado, Barbour 4934 (MO). Smith 4799 (GH).
3. Trichipteris kalbreyeri (Baker) Tryon, Contr. 5. Trichipteris nigra (Mart.) Tryon, Contr. Gray
Gray Herb. 200: 45. 1970. Herb. 200:46. 1970.
Alsophila kalbreyeri Baker, Summary new ferns 9. Alsophila nigra Mart.. Icon. pi. crypt. 71.1 834. TYPE:
1892, nom. nov. for Alsophila podophylla Baker Brazil, Prov. Rio Negro, Rio Japura, Martius (ho-
(not Hooker) and with the same type. lotype, not located: isotypes, B!, K).
Trichipteris lasiosora (Kuhn) Tryon, Contr. Gray Herb. pinnate-pinnatisect, gradually to abruptly reduced
200: 45. 1970. at the apex, central and basal pinnae short- to long-
Petiole lacking trichomes, the scales brown or ward the base, sessile to usually short-stalked, pin-
dark brown with lighter margins. Lamina 2-pin- na-rachises lacking spines, the abaxial and lateral
sides with abundant long, mostly whitish tri-
nate-pinnatifid to 2-pinnate-pinnatisect, abruptly
reduced at the apex, pinnae short- to long-stalked, chomes. Pinnules obtuse to attenuate, with long,
pinna-rachises lacking spines, the abaxial and lat- mostly whitish trichomes abaxially, ultimate seg-
somewhat pubescent, some-
eral sides glabrate to ments obtuse to subacute, the apex entire to
times with rather numerous, long, mostly brown coarsely toothed, fertile veins simple or usually
to light brown trichomes. Pinnules acuminate to forked. Paraphyses longer than the sporangia.
HOLOTYPE Peru, Dept. Huanuco, above Chin- 9. Trichipteris serpens Tryon, sp. nov.
chao, Tingo Maria to Chinchao, 1 Aug. 1965, D.
Soejarto 1432 (GH 2 sheets). Petiolus deest. Pedum circinatum sine trichomata,
Only known from the type collection from an squamis centralibus fuscatis angustis marginatis late niv-
eis. Lamina ca. 3-4 m longa tripinnato-pinnatifida pinnis
exposed hillside at 2500 m in Huanuco.
pinnulisque longe petiolulatis. Pinnulae acuminatae sub-
Endemic to Peru. ter squamis paucis brunneis complanatis. Venae fertiles
The stem is reported on the label as slender and furcatae. Paraphyses longiores quam sporangia.
ca. 75 cm tall, the leaves ca. 1.5 m long, and the
cut stem exuding yellow sap. Although the collec- Petiole lacking trichomes, the scales with a dark,
tion is sterile, it very probably represents a species narrow central stripe and broad whitish margins
of Trichipteris. The lamina complexity and the (characters from croziers). Lamina 3-pinnate-pin-
abundant short-pubescence on the upper surface natifid nearly throughout, apex absent, pinnae long-
of the segments amply distinguish the species. to very long-stalked, pinna-rachises lacking spines,
the abaxial and lateral sides somewhat appressed
pubescent and scaly. Pinnules short- to long-
8. Trichipteris frigida (Karsten) Tryon, Contr. stalked, acuminate, glabrate or with a few large,
Gray Herb. 200:45. 1970. flattish, brown scales abaxially, fertile veins forked.
1859. TYPE: Colombia, (Cundinamarca), Andes TYPE Peru, Dept. Cuzco, Prov. La Conven-
of Bogota, 2600 m, Karsten (holotype, not locat-
tion, Cordillera Vilcabamba, 10 July 1968, T. R.
ed; isotype, B!).
Cyathea frigida (Karsten) Domin, Pterid. 262. 1929. Dudley 10949 (holotype, NA 2 sheets; isotype, GH
2 sheets).
Petiole lacking trichomes, the scales light to dark Known only from the type collection in elfin
brown usually with rather narrow lighter margins. forest at 2900 m in Cuzco.
Lamina 2-pinnate-pinnatifid, rarely partly 3-pin-
Endemic to Peru.
nate-pinnatifid, gradually reduced at the apex, pin-
The leaves of this new species are unusual: they
nae sessile to long-stalked, pinna-rachises lacking are 4-6 m long, or more, and trail or are scandent
spines, the abaxial and lateral sides with large flat- over low vegetation. The scales of the croziers,
tish scales or theirhard bases, with scattered brown which will in part become petiole scales, are quite
to light brown trichomes or often with dense, ca- different from those of the related Trichipteris fri-
In montane rain forests, cloud forests, or ceja 12. Trichipteris microdonta (Desv.) Tryon, Contr.
de la montana, 850-2200 m, Amazonas south to Gray Herb. 200: 46. 1970.
Puno.
Guyana, Venezuela, and Colombia south to Bo- Polypodium microdontum Desv., Ges. Naturf. Freunde
livia. Berlin Mag. Neuesten Entdeck. Gesammten Na-
turk. 5: 319. 1811. TYPE: "America australis"
This species and the next are the only ones in
(holotype, Herb. Desv. P!; photos, GH, us).
Peru with a 1-pinnate-pinnatifid lamina. Trichip-
Alsophila microdonta (Desv.) Desv., Mem. Soc. Linn.
teris pubescens is widely distributed and, although Paris 6: 319. 1827.
variable, characterized by the features
is men- Cyathea microdonta (Desv.) Domin, Pterid. 263. 1929.
obtuse, fertile veins simple. Paraphyses longer than Urubamba, Tryon & Tryon 541 1 (GH, USM). Machu Pic-
chu, Iltis et al. 1024 (GH, uc).
the sporangia.
veins on an ultimate segment are connivent to a TYPE: Colombia, Cundinamarca, Cuesta "Fu-
sugasuga," (Fusagasuga), Cuatrecasas 8036 (ho-
sinus.
lotype, us!).
Huanuco: Cerros del Sira, Rio Llullapichis watershed, Petiole with long trichomes, these persistent or
Dudley 13047, 13214, 13220 (GH). Junin: Prov. Satipo,
S of Chequitavo, D. Smith 5130 (F).
deciduous and then leaving a hard base and a sca-
brous petiole surface, the scales dark brown to
atropurpureous, usually with broad, lighter mar-
gins, or whitish to light brown and concolorous,
14. Trichipteris conjugata (Hooker) Tryon, Contr.
the margins slightly or not dark denticulate. Lam-
Gray Herb. 200: 45. 1970.
ina 2-pinnate-pinnatifid to 2-pinnate-pinnatisect,
a. Indusia hemitelioid b
b. Sori costal 3. C. vilhelmii
b. Sori medial to submarginal c
c. Basal veins straight or nearly so, ending above a sinus d
d. Ultimate segments with simple fertile veins and usually entire (rarely coarsely dentate), or
with forked fertile veins and coarsely dentate 1 C. multiflora .
d. Ultimate segments with forked fertile veins, entire or finely dentate to strongly crenate . .
2. C. andina
c. Basal veins curved, connivent to a sinus, sometimes forming costal areolae ... 4. C. petiolata
a. Indusia sphaeropteroid, sometimes evanescent and then the apical umbo or a basal remnant is usually
present e
e. Petiole scales either whitish to light brownish and concolorous or nearly so, or discordantly
bicolorous with whitish to brownish margins; petiole scurf whitish to light brown, or rarely brown
at the base of the petiole f
f. Lamina 2-pinnate-pinnatifid or 3-pinnate only at the base of the pinnae g
g. Large scales of the petiole scurf flattish h
h. Pinnae and pinnules mostly long-stalked 5. C. divergens
h. Pinnae and pinnules sessile or nearly so 7. C. ruiziana
g. Large scales of the petiole scurf mostly, or many of them, crested 6. C. pallescens
f. Lamina 3-4-pinnate nearly throughout i
i. Tertiary segments entire or basally 2-lobed; bullate scales on the abaxial side of the pinnule-
rachis dark brown 8. C. microphylla
i. Tertiary segments with up to usually 5 or more segments or lobes; bullate scales on the
brown
abaxial side of the pinnule-rachis light 9. C. multisegmenta
e. Petiole scales eitherbrown and nearly concolorous, or (sometimes narrowly) concordantly bico-
lorous with light brown to brown margins; petiole scurf brown or absent j
j. Lamina 2-pinnate-pinnatifid or 3-pinnate only at the base of central and basal pinnae; larger
ultimate segments usually ca. 1 mm long k
k. Petiole scales light brown to brown, concolorous to narrowly bicolorous; pinnules usually
1. Cyathea multiflora Sm., Mem. Acad. Roy. Sci. Hemitelia andina Karsten, Linnaea 28: 452. 1856.
TYPE: Colombia, "Santa Martha," 2500 m, Kar-
(Turin) 5: 416. 1793. TYPE: "Amer. merid.,"
sten (not located; authentic specimen, Karsten,
R. Shakespeare (holotype, Herb. Banks BM!;
Herb. Men. B!; photo, GH!).
photos, GH, NY, us).
Petiole lacking, scales and scurf unknown. Lam- Petiole short-aculeate to aculeate, the scales very
ina 3-pinnate-pinnatifid, pinnules long-stalked, light brown to dark brown, predominantly dis-
glabrate abaxially, ultimate segments subentire to cordantly bicolorous with lighter margins, scurf
slightly toothed, fertile veins forked. Sori costal, dense, usually persistent, the large scales Hattish.
indusia hemitelioid, small, nearly concealed by the Lamina 2-pinnate-pinnatifid to 2-pinnate-pinna-
opteroid.
4. Cyathea petiolata (Hooker) Tryon, Contr. Gray Montane forests and cloud forests, 200-2000 1
5. Cyathea divergens var. divergens nearly so, glabrate to pubescent and (or) scaly
abaxially, ultimate segments subentire to crenate
or rarely lobed, fertile veins simple or forked. Sori
Cyathea divergens Kunze, Linnaea 9: 1 00. 1 834. TYPE:
Peru, (Huanuco), Pampayacu, Jul. 1 829, Poeppig costal to nearly medial, indusia sphaeropteroid.
In montane forests, primary or open forests, or lighter margins, scurf dense, persistent or ca-
ducous, the large scales flattish. Lamina 2-pinnate-
cloud forests, 850-2880 m, Amazonas, San Mar-
pinnatifid to 2-pinnate-pinnatisect, pinnules ses-
tin, Huanuco, Pasco, and Cuzco.
sile to short-stalked or rarely long-stalked, gla-
Costa Rica and Panama; northern South Amer-
ica south to Bolivia and southeastern Brazil. brous, pubescent and (or) scaly abaxially, some-
times densely so, ultimate segments subentire to
Cyathea delgadii is characterized by the long
trichomes abaxially, which are usually abundant crenate, fertile veins forked, rarely simple. Sori
costal to medial, indusia sphaeropteroid.
and the rather few, if any, scales. It is one of the
most widely distributed species of Cyathea.
The species occurs in the Greater Antilles; Costa
Rica; and Venezuela and Colombia south to Bo-
livia.
Amazonas: Prov. Bagua, Cordillera Colan, SE of La
Peca, Barbour 3612 (F, MO, USM). San Martin: Rioja, Cyathea caracasana is the most variable species
Soukup 5223 (GH). Huanuco: Tingo Maria, Tryon & Tryon among the American Cyatheaceae. Five varieties
5219 (F, GH, us). Cerros del Sira, Rio Llullapichis wa- were recognized by Tryon (1976) in order to fa-
tershed, Dudley 13070 (GH). Pasco: Prov. Oxapampa,
cilitate the study of this complex. In addition to
Yanachaga National Park, Leon 1014 (GH). Cuzco: Prov.
La Convention, Dudley 10058, 10268 (GH). Prov. Uru- the two varieties in Peru, var. maxonii is endemic
bamba, camino a Huinayhuayna, Chavez 3442 (MO). to Costa Rica, var. caracasana is in the Greater
Antilles and northern South America, and var.
chimborazensis is in Venezuela, Colombia, and
Ecuador.
Key to Varieties
a. Pinnules usually tapering to the apex from beyond the middle, abaxially usually with abundant scales
and trichomes 1 la. var. boliviensis
a. Pinnules, especially toward the base of the central and basal pinnae, tapering to the apex from the
base, indument abaxially sparse or absent lib. var. meridensis
1 la. Cyathea caracasana var. boliviensis (Ro- Amazonas: Prov. Chachapoyas, entre Ingenio y Po-
macocha, Ldpez et al. 4312 (GH, HUT). Prov. Chacha-
senst.)Tryon, Contr. Gray Herb. 206: 77.
poyas, Molinopampa-Diosan pass, Wurdack 1653 (F,
1976. Figure 23.
GH, us). Huanuco: Within 5 km of Carpish, Tryon &
Tryon 5326 (F, GH, NY, uc, us). Pasco: Prov. Oxapampa,
Cyathea mexicana var. boliviensis Rosenst., Repert. Rio San Alberto valley. Smith & Pretel 8003 (F, MO).
Spec. Nov. Regni Veg. 25: 56. 1928. TYPE: Bo-
Oxapampa (as Junin), Soukup 2335 (F, GH). Ucayali:
livia, Hacienda Simaco, above Tipuani, Buchtien
Prov. Coronel Portillo, La Divisoria, (as Loreto), Fer-
5140 (holotype, not located; isotypes, F!, GH!, NY!,
reyra 1074 (GH, us), 7696 (us, USM). Huancavelica: Prov.
us!).
Tayacaja, entre Huachocolpa y Tintay, Tovar 4201 (GH).
Cuzco: Prov. Paucartambo, near Santa Isabel, Pilahuata
In wooded ravines, mountain forests, and es-
to Patria, Plowman & Davis 4988 (GH, USM). Prov. Uru-
pecially in cloud forests, 300-2800 m, Amazonas
1
bamba. Puente Ruinas, Machu Picchu, Iltis et al. 1025
south to Puno. (GH, uc, us). Puno: Cerca a San Juan del Oro, Valle del
Venezuela and Colombia south to Bolivia. Alto Tambopata, Ferreyra 16684 (GH, USM).
FIG. 23. Cyathea caracasana \ar. boliviensis: a, rachis and base of pinna; b, sori (one opened); c, portion of petiole
scale. (From Wurdack 1653, F.)
Cyathea castanea Baker, Syn. fil. ed. 2, 451. 1874. Petiole nearly smooth, the scales brown to dark
TYPE: Peru, (San Martin), Tarapoto, Spruce 47 23 brown, nearly concolorous, scurf absent. Lamina
(holotype, K; isotypes, GH!, MO!, P!, us!; photos,
3-pinnate nearly throughout, pinnules sessile to
GH, US Of P).
short-stalked, somewhat scaly and pubescent
abaxially, ultimate segments entire, fertile veins
Petiole nearly smooth, the scales reddish brown
simple. Sori subcostal, indusia sphaeropteroid.
to atropurpureous, concordantly bicolorous with
VI. Cnemidaria with three large equatorial pores. All of the Pe-
ruvian species have regularly anastomosing veins,
Cnemidaria Presl, Tent, pterid. 56. 1836. TYPE: except for C. uleana, which sometimes has free
veins.
Cnemidaria speciosa Presl. Figure 24.
pecially on the croziers and petiole base, that are genus Cnemidaria (Cyatheaceae). Fieldiana,
Bot., 37: 1-98.
marginate and lack a dark apical seta. Petiole
smooth, muricate, or with corticinate spines. Lam-
a. Pinnae (excluding the basal pair and reduced apical ones) deeply pinnatifid or pinnatisect; the segment
sinuses extending %
or more to the costa b
b. Lamina gradually reduced to a nonconform pinnatifid apex; rachis not alate; petiole scales bi-
l.C. horrida
c. Basal basiscopic veins, especially toward the basal and apical portions of pinnae, commonly
arisingfrom the costa; rachis smooth, petiole smooth or tuberculate; scales of costae and costules
whitish 2. C. uleana
b. Lamina abruptly reduced to aconform or subconform apical segment similar to the lateral pinnae;
rachis conspicuously alate, especially distally; petiole scales predominantly whitish
3. C. alatissima
a. Pinnae (excluding the basal pair and reduced apical ones) subentire to shallowly pinnatifid; the segment
sinuses, if present, extending less than halfway to the costa d
d. Pinnae shallowly and obtusely lobed to shallowly pinnatifid; indusia more or less semicircular,
attached on the costular side of the receptacle 4. C. speciosa
d. Pinnae subentire to broadly and coarsely serrate; indusia circular, completely surrounding the
receptacle 5. C. nervosa
Hemitelia subincisa Kunze, Bot. Zeit. (Berlin) 2: 296. abaya, San Gaban, Vargas 18863 (GH).
duced to a conform or subconform apical segment, 309. 1944. TYPE: Peru, Loreto, mouth of Rio
Santiago, Mexia 6291 (holotype, us! 3 sheets; is-
rachis not alate, essentially smooth. Pinnae very
otypes, F!, GH!, NY!, uc!).
deeply crenate to lobed less than '/2 to the costa,
the costa and costules rarely with dull, brown scales
Petiole smooth to muricate, the scales, mostly
abaxially, basal basiscopic veins mostly arising
near the base, lanceolate or linear-lanceolate, bi-
from the costule or its base. Soral lines supra-
colorous, brown with narrow whitish margins.
medial between the costule and segment margin,
Lamina abruptly reduced to a conform or subcon-
indusia more or less semicircular, rather large.
form apical segment, rachis not alate, smooth. Pin-
nae subentire to broadly serrate, the costa and pri-
In forests and at forest borders, along stream
mary veins lacking scales abaxially, basal basiscopic
banks, and on mountain slopes, 1 1 5-1900 m, San vein arising from a primary vein. Soral lines be-
Martin and Loreto south to Puno. tween the primary veins, indusia commonly cir-
Peru and Bolivia.
cular, completely surrounding the receptacle, en-
Three collections from Loreto, Varadero de Ma- tire to lobed.
zan, (Rio Amazonas to Rio Napo), Croat 19500
(MO, uc), 79574 (F, MO) and 20797 (MO, uc) have Rain forests, 300450 m, Amazonas and Lor-
a few broad whitish scales on the costae abaxially, eto.
rather than few small brown ones or none. These Ecuador and Peru.
may represent a distinct variety of the species. The circular indusium is a character in
the genus
shared only by Cnemidaria cocleana of Panama.
San Martin: Prov. Mariscal Caceres, Palo Blanco, To- Cnemidaria nervosa is a rare fern, represented by
cache Nuevo, J. Schunke V. 5682 (F, GH, NY). Loreto:
Prov. Maynas, Varadero, Vdsquez 721 (MO). Huanuco:
one collection from Ecuador and two from Peru.
Prov. Huanuco, Tingo Maria, Tryon & Tryon 5260 (F,
GH, u, us). Pasco: Yapas, Pichis Trail, Killip & Smith Amazonas: Prov. Bagua. valley of the Rio Maranon,
25563 (GH, NY, us). Prov. Oxapampa, Palcazu, Cerro de Wurdack 2059 (NY, us).
1. Tumbes
2. Piura
3. Lambayeque
4. Cajamarca
5. Amazonas
6. La Libertad
7. San Martin
8. Loreto
9. Ancash
10. Hulnuco
11. Lima
12. Pasco Bolivia
13. Junlh
14. Ucayali
15. lea
16. Huancavelica
17. Ayacucho
18. ApurTmac
19. Cuzco
20. Madre de Dios
21. Arequipa
22. Puno
23. Moquegua
24. Tacna Chile
DEPARTMENTS OF PERU
Accepted names are in roman type, synonyms are in italics, and new names are in boldface. A page
number is provided for the principal place, or the only place, where the name occurs.
mbiginosa 46 /ar/pes 59
schomburgkiana 47 lindenii 7 1
seminuda 47 Hemiphlebium
lineare 66
kapplerianum 88
simplex 39 lobatoalatum 74
Hemitelia 129
velata 42 mathewsii 60
andina 130
yungensis 44 mexiae 59
horrida 138
Didymoglossum 76 microcarpum 68
fccAferi 131
angustifrons 86 mirificum 58
multiflora 130
hymenoides 86
FIELDIANA: BOTANY
142
Hymenophyllum Lomaridium Mertensia
molle 65 semicordatum 101 pectinata 49
multialatum 73 Lophidium 33 pennigera 39
multiflorum 63 elegans 34 pruinosa 42
myriocarpum 62 flabellum 34 remota 44
var. endiviifolium 63 latifolium 33 revoluta 42
var. myriocarpum 62 poeppigianum 36
var. nigrescens 63 Lophosoria 107 simplex 37
nigrescens 63 pruinata 109 tomentosa 4 1
nigricans 63 quadripinnata 107
nigricans 63 var. contracta 109 Metaxya 109
pedicellatum 57 var. quadripinnata 109 rostrata 1 1 1
rostratum 1 1 1
medullaris 1 1 2 ciliatum 67
Publication 13*
Publication 1349.
UNIVERSITY OF ILLINOIS URBANA
580 5FBN.S COOS
FIELDIANA BOTANY CHGO
20 1988
30112017564094
UNIVERSITY OF ILLINOIS-URBANA