UNIVERSITY OF
ILLINOIS LIBRARY
AT URBANA CHAMPAIGN
Ou ^BIOLOGY
JUL11198*
PIOLOGY
71- -JANA
Botany
Published by Field Museum of Natural History
Volume 39
FERNS AND FERN ALLIES
OF GUATEMALA
PART I
OPHIOGLOSSACEAE THROUGH
CYATHEACEAE
ROBERT G. STOLZE
B 2 3
December 8, 1976
FIELDIANA: BOTANY
A Continuation of the
BOTANICAL SERIES
of
FIELD MUSEUM OF NATURAL HISTORY
VOLUME 39
FIELD MUSEUM OF NATURAL HISTORY
CHICAGO, U.S.A.
FERNS AND FERN ALLIES
OF GUATEMALA
PARTI
FIELDIANA
Botany
Published by Field Museum of Natural History
Volume 39
FERNS AND FERN ALLIES
OF GUATEMALA
PARTI
OPHIOGLOSSACEAE THROUGH
CYATHEACEAE
ROBERT G. STOLZE
Custodiana, Pteridophyte Herbarium
Field Museum of Natural History
December 8, 1976
Publication 1246
Library of Congress Catalog Number: 76-47775
US ISSN 0015-0746
PRINTED IN THE UNITED STATES OF AMERICA
We acknowledge the assistance of the National Science Foundation, program for
Systematic Biology, given to the principal investigator, Louis O. Williams, over a
period of many years for the "Flora of Guatemala." This welcome assistance made
possible both the field work and the research necessary to complete this work.
The National Science Foundation, Office of Science Information Service, granted
the principal investigator funds which permit the publication of the final parts of the
"Flora of Guatemala." The Museum and the principal investigator are most appre •
ciative of this financial aid.
LIST OF ILLUSTRATIONS
1. Botrychium: B. dissectum ssp. decomposition; B. underwoodianum;
B. virginianum var. mexicanum 7
2. Ophioglossum: O. palmatum; O. ellipticum; O. reticulatum; O.
crotalophoroides; O. nudicaule var. tenerum; O. nudicaule var. vulcanicum . 11
3. Danaea: D. cuspidata; D. elliptica; D. nodosa 15
4. Marattia: M. excavata; M. interposita 18
5. Osmunda: O. regalis var. spectabilis ; O. cinnamomea 23
6. Actinostachys germanii 26
7. Anemia: A. cicutaria; A. speciosa; A. pastinacaria; A. hirsuta;
A. hirta; A. oblongifolia 29
8. Lygodium: L. uolubile; L. heterodoxum; L. venustum 37
9. Schizaea elegans 40
10. Dicranopteris: D. flexuosa; D. pectinata 43
11. Gleichenia: G. palmata; G. bancroftii; G. bifida; G. underwoodiana 47
12. Hymenophyllum: H. fucoides; H. polyanthos; H. crispum; H. lanatum .... 57
13. Trichomanes: T. collariatum ; T. radicans; T. reptans; T. godmanii 75
14. Plagiogyria: P. semicordata; P. pectinata 92
15. Cibotium regale: 95
16. Culcita coniifolia 98
17. Dicksonia gigantea 101
18. Alsophila salvinii 104
19. Cnemidaria decurrens 107
20. Cyathea: C. tuerckheimii; C. multiflora; C. tuerckheimii; C. fulua 110
21. Lophosoria quadripinnata 114
22. Metaxya rostrdta 115
23. Nephelea: N. mexicana; N. tryoniana 117
24. Sphaeropteris: S. horrida; S. myosuroides 121
25. Trichipteris: T. schiedeana; T. scabriuscula; T. pansamalana 125
VII
CONTENTS
Ophioglossaceae 4
Botrychium 5
Ophioglossum 9
Marattiaceae 13
Danaea 14
Marattia 16
Osmundaceae 21
Osmunda 21
Schizaeaceae 24
Actinostachys 25
Anemia 27
Lygodium 36
Schizaea 41
Gleicheniaceae 41
Dicranopteris 42
Gleichenia 45
Hymenophyllaceae 51
Hymenophyllum 52
Trichomanes 71
Plagiogyraceae 90
Plagiogyria 91
Dicksoniaceae 93
Cibotium 94
Culcita 97
Dicksonia 99
Cyatheaceae 102
Alsophila 103
Cnemidaria 105
Cyathea 108
Lophosoria 113
Metaxya 115
Nephelea 116
Sphaeropteris 120
Trichipteris 123
Ferns and Fern Allies of Guatemala
The fern flora of Guatemala is exceedingly rich and diverse, and
since such a great number of species is involved, it was thought
more practicable to publish them in several parts of one volume.
Part I treats all of the more primitive families, through the Cyathe-
aceae, and includes a total of 110 species. Subsequent parts will deal
with the Polypodiaceae (sens. lat. ), the water ferns Marsileaceae
and Salviniaceae, and, finally, the "fern allies."
A number of phyletic systems have been suggested in the past 40
years by various authors, but among current pteridologists there
appears to be more disagreement than agreement as to which
scheme is the most "valid." Furthermore, in arranging genera and
families in linear sequence, as is necessary in a floristic treatment,
phyletic relationships are often confused or obscured. Therefore,
there is no attempt here to deal with new problems of phylogeny or
classification. A very conservative system is employed, similar to
that of Christensen (in Verdoorn, Manual of Pteridology , 1938), and
families are arranged in such an order. Within families, genera and
species are arranged alphabetically. This is in keeping with the for-
mat heretofore utilized in all treatments in the "Flora of Guate-
mala."
During the early days of publication of the "Flora," the late
Conrad V. Morton of the U. S. National Museum agreed to do the
research on the pteridophytes, and for a number of years he ex-
pended considerable time and effort on the project. Unfortunately,
the pressures of other research and, eventually, ill health prevented
him from completing the work. Shortly before Mr. Morton's death
in 1972, I was asked by Louis O. Williams to handle the task, and
when I agreed, Morton very graciously offered to lend me all the
notes on his research up to that time. These included species lists, a
number of preliminary keys, and some notes on various genera, all
of which have been extremely useful to me and have expedited the
work considerably. It is with a deep sense of appreciation that I ac-
knowledge his generosity.
2 FIELDIANA: BOTANY, VOLUME 39
I am indebted to Holla Tryon and David Lellinger for reading por-
tions of the manuscript, and for their valuable comments and criti-
cisms on the treatment of many genera. I also wish to thank W. H.
Wagner, David Barrington, Gerald Gastony, and Paul Windisch for
advice pertaining to the genera with which they are especially ac-
quainted. The fine illustrations were produced by Richard Roesener
and Marion Pahl, who contributed not only their expertise, but their
patience and understanding as well. I would also like to express my
appreciation to Dorothy Nash and Laura Schlivek, for their assis-
tance in typing, proofreading, and the countless other tasks essen-
tial to bringing the manuscript into its final form.
Research on Part I of the "Ferns and Fern Allies of Guatemala"
has been based principally on the outstanding fern collection at
Field Museum of Natural History. In addition, several thousands of
specimens have been studied from the following herbaria: United
States National Herbarium, Washington, D.C.; Gray Herbarium,
Cambridge, Massachusetts; Museo de Historia Natural, Guatemala
City, Guatemala; New York Botanical Garden, Bronx, New York;
and the University of Michigan Herbarium, Ann Arbor, Michigan. I
wish to express my deep appreciation to the curators of these collec-
tions for granting me loans of their material, or for allowing me to
examine the specimens at their institutions.
Finally, I wish to thank Louis O. Williams, who has filled the mul-
tiple role of mentor, colleague, and friend throughout the course of
this work. His guidance and encouragement have made it all pos-
sible.
All abbreviations of periodical publications follow the system of
Botanico-Periodicum-Huntianum, Hunt Botanical Library, Pitts-
burgh, Pennsylvania, 1968.
KEY TO FAMILIES OF GUATEMALAN FERNS AND FERN ALLIES
a. Plants homosporous.
b. Leaves ample, broad to grasslike (if very small, then with marginal or super-
ficial sori), circulate in vernation (except Ophioglossaceae); sporangia borne
variously on the margins or abaxial surfaces of leaves or on terminal spikes or
panicles.
c. Sporangia thick-walled; annulus lacking,
d. Sporangia single, borne in panicles or immersed in the tissue of spikes;
leaves not circinate in vernation (erect, or only folded over)
OPHIOGLOSSACEAE.
d. Sporangia fused into synangia, borne on the abaxial surface of leaves;
leaves circinate in vernation. . . . MARATTIACEAE.
STOLZE: FERNS OF GUATEMALA 3
c. Sporangia thin-walled (1 cell thick); annulus partially to fully encircling the
sporangia (or in Osmundaceae rudimentary or lacking).
e. Annulus of sporangia oblique or transverse, uninterrupted by the stalk (or
in Osmundaceae distal and often rudimentary or lacking),
f . Leaves translucent, mostly one cell thick . . . HYMENOPHYLLACEAE.
f. Leaves opaque, more than one cell thick.
g. Leaves dimorphous; sporangia not arranged in definite groups, borne
singly or covering the surface of a segment; and commonly (in ours)
borne on scarcely- or non-foliaceous fertile pinnae, or in rows on special-
ized, non-foliaceous marginal segments,
h. Annulus of sporangia few-celled at the distal end, or essentially
lacking OSMUNDACEAE.
h. Annulus of sporangia complete, oblique or transverse and apical,
i. Sporangia borne in panicles, or in rows on specialized, scarcely- or
non-foliaceous segments; leaves various, linear to flabelliform, or
pinnately or dichotomously branched SCHIZAEACEAE.
i. Sporangia completely covering the abaxial surface of fertile pin-
nae; leaves pinnate topinnatifid PLAGIOGYRIACEAE.
g. Leaves monomorphous; sporangia arranged in definite groups (sori) on
the margins or abaxial surface of leaves,
j. Leaves pseudodichotomously branched, scandent or trailing; spores
120-800 in each sporangium GLEICHENIACEAE.
j. Leaves pinnately divided, erect and arborescent or subarborescent;
spores 16-64 in each sporangium,
k. Sori marginal, the indusia bivalvate; indument consisting only of
trichomes, scales lacking DICKSONIACEAE.
k. Sori borne on the abaxial surface of leaf segments, the indusia
scalelike to cup-shaped or globose, or lacking; indument of scales
or trichomes, or both CYATHEACEAE.
e. Annulus of sporangia vertical, interrupted by the stalk
POLYPODIACEAE.
b. Leaves rudimentary or minute or scalelike, not circulate in vernation; sporan-
gia borne at the axils of leaves or in strobili terminating the branches or the
main axis.
1. Stems hollow and conspicuously jointed; leaves forming a sheath at stem
nodes; branches whorled or none EQUISETACEAE.
1. Stems solid, not jointed; leaves rudimentary or lacking or, if present, not
sheathing at nodes; branches dichotomous or subdichotomous.
m. Leaves numerous, conspicuous, in several ranks; sporangia single, uni-
locular, bivalvate LYCOPODIACEAE.
m. Leaves (in ours) few and rudimentary, or essentially lacking; sporangia
fused into a tri-locular synangium PSILOTACEAE.
a. Plants heterosporous, bearing both microspores (male) and megaspores (female).
n. Plants free-floating on water, the roots trailing SALVINIACEAE.
n. Plants not free-floating (if aquatic, then rooted in mud),
o. Leaves (in ours) 4-foliolate, commonly floating on the water; sporangia borne
in stalked, hardened, nutlike sporocarps near petiole base
MARSILEACEAE.
4 FIELDIANA: BOTANY, VOLUME 39
o. Leaves simple; sporangia borne in the axils of scalelike leaves or imbedded in
the broadened bases of subulate leaves.
p. Stem cormlike, subterranean, bearing simple, long-subulate, quill-like or
rushlike leaves, commonly partly or wholly immersed in water; sporangia
imbedded in the imbricate leaf bases ISOETACEAE.
p. Stem pinnately- or subdichotomously-branched, the branches and main
stem bearing numerous, minute leaves (plants sometimes of wet places, but
not normally immersed in water); sporangia borne in axils of leaves which
are arranged in terminal strobili SELAGINELLACEAE.
OPHIOGLOSSACEAE
References: L. M. Underwood and R. C. Benedict, Ophioglossa-
ceae, in North Amer. Fl. 16: 3-13. 1916. Robert T. Clausen, A mono-
graph of the Ophioglossaceae, Mem. Torrey Bot. Club 19: 1-177.
1938.
Small, terrestrial, or, rarely epiphytic herbs; rhizome hypogean, short, usually
erect, with the thick, fleshy, mycorrhizal roots lacking root hairs; stipular sheaths
present at the base of the leaf; leaves solitary or few, erect or pendent, not circinate
in vernation (or only folded over), dimorphic, or (in ours) with a common stalk bear-
ing a foliaceous sterile lamina and one or several non-foliaceous fertile segments;
sterile lamina stalked or sessile, simple to decompound, with free or reticulate vena-
tion; fertile segments spicate or paniculate; sporangia developed from a group of
epidermal and subepidermal cells, in two rows on the surface of axes of the panicles
or immersed in the tissue of the spike, with walls several cells thick, lacking an an-
nulus, dehiscent into two valves by a transverse slit; spores uniform, trilete, numer-
ous, from 1,500 to 15,000 in each sporangium.
The family Ophioglossaceae, containing three genera and about
50 species, differs from the true ferns (Filicales) in several impor-
tant respects: the erect or curved, rather than circinate, vernation,
the stipular sheaths, the large thick-walled sporangia, and the
fleshy leaf texture, which is due to the complete lack of scleren-
chymatous tissue. The Ophioglossaceae are widely distributed
throughout the world, with representation in arctic as well as tropi-
cal regions. Some of the species have wide and disjunct distribu-
tions, e.g., Botrychium lunaria has been reported in such diverse
areas as Greenland, Argentina, Siberia, and New Zealand. Several
species of Ophioglossum are pan- tropical. The family has no eco-
nomic importance. Two of the three genera are represented in
Guatemala.
a. Sterile lamina ternately decompound; veins free; sporangia borne in a panicle,
free from each other and not immersed in the fertile segment Botrychium.
a. Sterile lamina entire, or palmately or digitately lobed; veins reticulate; sporangia
borne in a spike, fused laterally and immersed in the fertile segment
Ophioglossum.
STOLZE: FERNS OF GUATEMALA 5
BOTRYCHIUM Swartz
References: J. Milde, Botrychiorum monographia, Verb. Zool.
Bot. Ges. Wien 19: 55-190. 1869. F. K. Butters, Botrychium virgini-
anum and its American varieties, Rhodora 19: 207. 1917.
Terrestrial herbs; rhizome hypogean, the bud for the following season partly or
wholly enclosed within the base of the common stalk; leaves one or two, sparsely
pubescent; sterile lamina sessile to long-stalked, ternately decompound or nearly so,
the basal pinnae much longer and broader than the others, sometimes nearly as
large as the rest of the lamina; venation pinnate, free; fertile segment a solitary,
long-stalked, once- to thrice-pinnate panicle; sporangia protruding from the surface
of the panicle, not immersed in the tissue.
The genus Botrychium contains about 25 species, with distribu-
tion in temperate to tropical regions in both hemispheres.
a. Sterile lamina sessile or nearly so
B. virginianum var. mexicanum.
a. Sterile lamina long-stalked,
b. Segment tips rounded or truncate; ultimate segments widely spaced, with
broad, mostly rounded sinuses B. underwoodianum.
b. Segment tips acute or subacute; ultimate segments mostly crowded, joined by
narrow, acute sinuses
B. dissectum ssp. decomposition.
Botrychium dissectum Sprengel ssp. decompositum (Mart. &
Gal.) Clausen, Mem. Torrey Bot. Club 19: 58. 1538. B. decomposi-
tum Mart. & Gal. Nouv. Mem. Acad. Roy. Sci. Bruxelles 15: 51. 1. 1.
1842.
Grassy, open places or partly open forests, 900-2,200 m.; Alta
Verapaz; Baja Verapaz; Chiquimula; Huehuetenango; Jalapa;
Zacapa. United States (Louisiana); Mexico; Honduras; Costa Rica;
Panama.
Mature plants 22-50 cm. high; common stalk 2-4 cm. long; stalk of the sterile seg-
ment 8-30 cm. long; sterile lamina 5-15 cm. long and 9-22 cm. broad, deltoid, or
roughly pentagonal, subternately decompound; pinnae (except the enlarged, more
highly dissected basal pair) pinnatisect to once-pinnate, very rarely pinnate-pinnati-
sect, with their tips commonly elongate; costae diverging from the rachis at a
relatively narrow angle (about 45°); ultimate segments 6-16 mm. long, crowded,
joined by narrow, acute sinuses, ovate, oblong, or lanceolate, obtuse to acute, their
margins irregularly serrate to crenate; fertile stalk 15-32 cm. long, panicle 2- or 3-
pinnate.
In his monograph, Clausen reported B. schaffneri Underwood
(Seler2738) from Guatemala, citing the specimen at the U. S. Natio-
nal Herbarium. However, he also cited the collection with this same
number at the N. Y. Botanical Garden as B. dissectum ssp. decom-
6 FIELDIANA: BOTANY, VOLUME 39
positum. Examination of both specimens now supports the latter
identification; thus there is still no documented record of B. schaff-
neri from Guatemala.
The leaves of ssp. decomposition vary widely in their shape and
cutting throughout the range. Although the typical plant can easily
be distinguished from B. underwoodianum, there are atypical
plants, as well as some with immature leaves, which could be con-
fused. Additional characteristics are discussed under B. underwood-
ianum Maxon.
It is possible that there are two different elements included in B.
dissectum in Guatemala. One of these, presumably typical decom-
positum, cannot be matched by any plants from the United States
or Canada, where B. dissectum occurs in many forms. In contrast,
there is a collection from Guatemala (Hatch & Wilson 264), as well
as another from Honduras, which can be closely matched by speci-
mens from temperate North America. This is a form in which the
segments are sharply angular and rhomboidal, perhaps approaching
forma obliquum (Muhl. ) Fern. Specimens of remarkably similar
appearance were discovered by Dr. W. D. Reese of Southwestern
Louisiana University, growing in more or less dry rises in bayous
west of New Orleans. More material was later gathered by Dr.
Reese and Dr. W. H. Wagner, Jr. (University of Michigan), which I
examined at the herbarium in Ann Arbor. Some of the Louisiana
specimens cannot be distinguished from the collections cited from
Central America.
Furthermore, the identity of a number of specimens of B. dissec-
tum with more highly divided leaves and exceptionally small ulti-
mate segments is still in doubt. ( As indicated above, some Mexican
material has been misidentified as B. schaffneri. ) Much more study
is needed in Section Sceptridium of the genus, particularly study of
mass collections from Mexico and Central America, before a more
precise classification of the group is possible.
Botrychium underwoodianum Maxon, Bull. Torrey Bot. Club 32:
222. 1905.
Forested slopes, 1,150-3,500 m. Huehuetenango; Zacapa. Hon-
duras; Costa Rica; Venezuela; Jamaica.
Mature plants 22-53 cm. high; common stalk 1.5-3.5 cm. long; stalk of the sterile
segment 8-28 cm. long; sterile lamina 8-14 cm. long and 11-16 cm. broad, deltoid or
roughly pentagonal, decompound; pinnae (except enlarged basal ones) pinna te-pin-
natisect to twice-pinnate, divided nearly to their tips, these not particularly elon-
FlG. 1. Botrychium, habit, X 1/z. a, B. dissectum ssp. decompositum; b, B. under-
woodianum ( sterile lamina only ); c,B. virginianum var. mexicanum.
8 FIELDIANA: BOTANY, VOLUME 39
gate; costae, especially of lower pairs of pinnae, diverging from rachis at a broad
angle (60-90°), at least in dried specimens; ultimate segments 7-14 mm. long, widely
spaced, with broad, mostly rounded sinuses, oblong, obovate or subspathulate,
truncate, obtuse or (rarely) subacute, their margins irregularly serrate to crenate;
stalk of the fertile segment 16-29 cm. long, panicle 2- to 3-pinnate.
In addition to the characters used in the key, there are other char-
acters which are often helpful in separating B. underwoodianum
from B. dissectum ssp. decomposition. In the latter the tips of the
ultimate segments are mostly acute or subacute, and the costae
diverge from the rachis at a relatively narrow angle, averaging
about 45°. In B. underwoodianum, the tips of the ultimate segments
are commonly rounded to truncate, and the costae diverge from the
rachis at a broader angle (from 60-90°). Wagner, however, who has
made frequent field studies of Botrychium, cautions against using
the costa angle in determination of dried material, suggesting that
this may be artificially induced in pressing.
Botrychium virginianum (L. ) Swartz var. mexicanum Grev. &
Hook., in Hooker's J. Bot. Kew Card. Misc. 3: 223. 1833 (as B. vir-
ginicum beta mexicanum}. B. cicutarium (Savigny) Sw., Syn. Fil.
171. 1806. B. brachystachys Kunze, Linnaea 18: 305. 1844.
Wet forests and wooded slopes, 900-3,800 m. Alta Verapaz; Chi-
maltenango; Guatemala; Jalapa; El Progreso; Quezaltenango; San
Marcos; Solola; Suchitepequez ; Zacapa. Mexico; Costa Rica; Pan-
ama; Greater Antilles; Venezuela and Colombia south to Bolivia
and Brazil; Old World.
Mature plants 29-55 cm. high; common stalk 19-30 cm. long; sterile lamina sessile
or short-stalked (to 2 mm.) to 21 cm. long and 24 cm. broad, deltoid, ternately di-
vided or essentially so, with the lowest pinnae commonly much longer than the
others; pinnae (except enlarged basal ones) mostly pinnate-pinnatisect; ultimate
segments 4-9 mm. long, more or less crowded, joined by narrow, acute sinuses, ellip-
tic, oblong or obovate, truncate, rounded or subacute, their apices variously
toothed or lacerate; fertile stalk 5-15 cm. long, with panicle 3-12 cm. long, pinnate to
bipinnate.
In his monograph, Clausen suggested that B. cicutarium might
well be considered a subspecies of B. virginianum , a species which
occurs in parts of the Old World and in the western hemisphere from
Canada to Mexico. The two had been traditionally separated on the
basis that the latter has fertile segments greatly exceeding the
sterile blade in length, the blades are more highly dissected, and the
plants produce two leaves in a season; whereas in B. cicutarium the
fertile segments exceed the sterile slightly or not at all, the sterile
blade is not as highly dissected, and only one leaf is produced in a
STOLZE: FERNS OF GUATEMALA 9
season. When studying many collections, one may note that tem-
perate zone plants generally possess such characters thus attrib-
uted to B. uirginianum, while plants from Mexico southward gen-
erally have characters aligned with B. cicutarium. Hence it still may
be prudent to maintain a distinction at this time, even if not at the
species level.
Since Clausen's monograph, more tropical American collections
have been made in many areas, offering further evidence that the
two taxa are conspecific. The characteristics are extremely variable,
so that a number of plants of B. virginianum, in the United States,
for example, have the fertile segments equal to or only slightly ex-
ceeding the sterile blade. Conversely, some plants have been col-
lected in Central America with their fertile segments far exceeding
the sterile. Likewise, the degree of dissection may vary to some
extent throughout the range of both species. Greville and Hooker
chose to assign varietal status to Mexican and Central American
plants of B. uirginianum, a view which was shared by a number of
other authors, including Chamisso, Schlechtendal, Milde, and Tor-
rey. I consider this classification more appropriate and quite consis-
tent with the subspecific status currently recognized in other com-
parable taxa in the genus. As with the B. dissectum complex, fur-
ther field work, especially mass population studies, will be neces-
sary to effect a proper understanding of the relationships of the
taxa.
OPHIOGLOSSUM Linnaeus
Terrestrial or epiphytic herbs ; rhizome hypogean, with the bud for the following
season exposed at the base of the common stalk; leaves solitary or few, glabrous;
sterile lamina sessile or short-stalked, entire or palmately- to digitately lobed; vena-
tion reticulate, the primary areoles enclosing free veinlets and/or smaller, secondary
areoles; fertile segments consisting of one to several, stalked, slender spikes arising
from the apex of the common stalk or from the base of the sterile blade; sporangia
coalescent, immersed in the tissue of the spike.
Approximately 25 species of Ophioglossum are scattered widely
around the earth. Four species are found in Guatemala, one of which
is represented by two varieties. A fifth is also to be expected in
Guatemala.
a. Sterile lamina palmately or digitately lobed; fertile segments several, arising
from the upper part of the stalk or lower part of the sterile lamina. . O. palmatum.
a. Sterile lamina entire; fertile segment solitary, arising at base of sterile lamina.
b. Pale, median band of tissue and midvein extending nearly to apex of sterile
lamina; veins forming areoles which enclose smaller secondary areoles as well
as free veinlets O. ellipticum.
10 FIELDIANA: BOTANY, VOLUME 39
b. Pale, median band and midvein lacking; veins forming areoles not enclosing
secondary areoles, but often with included free veinlets.
c. Rhizome conspicuously globose O. crotalophoroides.
c. Rhizome cylindrical to somewhat swollen, but never globose,
d. Common stalk 3-12 cm. long, lamina usually arising from middle of
plant O. reticulatum.
d. Common stalk not over 2.5 cm. long, lamina arising from base of plant.
e. Base of lamina cuneate to short attenuate
O. nudicaule var. tenerum.
e. Base of lamina truncate to subcordate, usually clasping
O. nudicaule var. vulcanicum.
Ophioglossum crotalophoroides Walter, Fl. Carolin. 256. 1788.
Moist, grassy, open places, 900-3,000 m.; Chiquimula; Huehue-
tenango; [Chimaltenango; Zacapa, fide Morton]. Southern United
States; Mexico; [Honduras, fide Morton]; Colombia; Peru; Bolivia;
Uruguay.
Plants terrestrial; rhizome globose, bearing one or (commonly) two leaves;
mature plant 3.5-13 cm. tall; common stalk 0.5-3.5 cm. long; fertile segment 2-11 cm.
long, with spike 0.5-1.5 cm. long and 1-3.5 mm. thick; sterile lamina 1-4.5 cm. long
and 0.8-2 cm. wide, ovate to deltoid, acute or rarely obtuse, cordate or truncate and
abruptly attenuate at base, lacking a pale, median band of tissue and distinct mid-
vein (or, rarely, a poorly defined midvein present near the base) ; venation indistinct,
diffuse, reticulate, the areoles occasionally with included free veinlets.
Among the diminutive species of Ophioglossum, occasional
plants may be found with rhizomes which are quite swollen. How-
ever, only in O. crotalophoroides is the rhizome so conspicuously
globose.
Ophioglossum ellipticum Hook. & Grev. Icon. Filicum 1: t. 40A.
1831.
Wet, grassy, mostly open places, 1,400-1,600 m.; Jalapa. British
Honduras; Honduras; Costa Rica; the Guianas southward to Brazil
and Bolivia.
Plants terrestrial; rhizome thick-cylindrical, bearing two to several leaves;
mature plant 4-16 cm. tall; common stalk 0.6-3.5 cm. tall; fertile segment 3-14 cm.
long, with spike 1-2.5 cm. long and 1.2-3 mm. thick; sterile lamina 1.5-6 cm. long and
0.6-1.7 cm. wide, elliptic, acute to rarely obtuse, cuneate to short-attenuate at base,
with a distinct midvein and pale, median band of tissue (the latter often becoming
indistinct at the apex) ; venation rather distinct, diffuse, with somewhat raised veins
forming elongate, primary areoles, which contain numerous secondary areoles and
often free, included veinlets.
FIG. 2. Ophioglossum. a, O. palmatum, habit, X 1/z; b-d, O. ellipticum: b, habit, X
Vf, c, portion of lamina, showing venation, X 2Vi; d, rhizome, X I1/?; e, O. reticu-
latum, portion of lamina, showing venation, X 2; f-g, O. crotalophoroides; f, habit,
X %; g, rhizome, X IVi; h, O. nudicaule var. tenerum, lamina outline, X 5; i, O. nudi-
caule var. vulcanicum, base of lamina, X 10.
11
12 FIELDIANA: BOTANY, VOLUME 39
The pale, median band of tissue and the double-areolate venation
are characters which easily distinguish O. ellipticum from all other
species of the genus in Guatemala.
Ophioglossum nudicaule L. f. var. tenerum (Mett. ex Prantl)
Clausen, Mem. Torrey Bot. Club 19: 146. 1938. O. tenerum Mett. ex
Prantl, Ber. Deutsch. Bot. Ges. 1: 352. 1883.
Moist, grassy, open places or open pine barrens, 200-2,100 m.;
Zacapa. Georgia, Florida; Hispaniola; Colombia; French Guiana;
Bolivia; Old World.
Plants terrestrial; rhizome thick-cylindrical to swollen, but never globose, bearing
one to two leaves; mature plant 2-11.5 cm. tall; common stalk 0.5-2.5 cm. long; fer-
tile segment 1.5-9 cm. long, with spike 0.3-1.5 cm. long and 1-2 mm. thick; sterile
lamina 0.5-2.5 cm. long and 0.2-0.8 cm. wide, elliptic to ovate, acute to rarely obtuse,
cuneate to short-attenuate at base, with midvein lacking or very poorly defined;
venation somewhat obscured due to the thickened texture, diffuse, reticulate, the
areoles mostly with free, included veinlets.
Clausen pointed out in his monograph that, on the basis of a few
intermediate plants collected in Africa, the separation of O. nudi-
caule var. tenerum from the typical variety might not be justifiable.
In either case, the plants in Guatemala are separable from those of
var. vulcanicum, as is designated in the key.
Ophioglossum nudicaule L. f. var. vulcanicum Clausen, Mem.
Torrey Bot. Club 19: 150. 1938.
Open meadows on slopes of volcanos, 2,800-3,700 m.; Huehue-
tenango. Panama.
Plants terrestrial; rhizome cylindrical, bearing one or two leaves; mature plant
3-5 cm. tall; common stalk 0.5-2 cm. long; fertile segment 2-4.5 cm. long, with spike
0.5-1 cm. long and 1.5-2.5 mm. thick; sterile lamina 0.8-1.7 cm. long and 0.4-0.7 cm.
wide, ovate to suborbicular, acute or often subapiculate, base truncate to subcor-
date, usually clasping, with midvein lacking; venation somewhat obscured due to
the thickened texture, diffuse, reticulate, the areoles mostly with free, included vein-
lets.
Ophioglossum palmatum L. Sp. PI. 2: 1063. 1753. Cheiroglossa
palmata (L.) Presl, Suppl. Tent. Pterid.: 57. 1845. Ophioderma pal-
mata (L.) Nakai, Bot. Mag. (Tokyo) 39: 193. 1925.
Not yet collected in Guatemala, there is good reason to expect it
there. It is a rather rare fern in all the regions in which it has been
found. Deep shade of forests, on tree trunks, branches, stumps, 0-
2,500 m. Florida; West Indies; Honduras; Costa Rica; Colombia;
British Guiana; Brazil; Ecuador; Peru.
STOLZE: FERNS OF GUATEMALA 13
Plants epiphytic, with fleshy leaves pendent or spreading; rhizome stout, bearing
one to several leaves; rhizome and petiole base densely covered with broad, pale,
tawny or reddish scales, which are long-fringed and hairlike for most of their length;
mature plant 24-100 cm. long; common stalk 15-50 cm. long; fertile segments 1 to 8,
borne on upper stalk or base of leaf, with fertile spikes 1.5-4.5 cm. long and 2-4 mm.
thick, borne on short stalks 0.2-1.8 cm. long; sterile lamina 10-50 cm. long and 9-26
cm. wide, lacking a midrib, palmately or digitately lobed, with apices of lobes obtuse
to acute, leaf base cuneate to attenuate; venation evident, reticulate, with free,
included veinlets and/or secondary areoles enclosed by larger, primary areoles.
Ophioglossum reticulatum L. Sp. PI. 2: 1063. 1753.
Open pastures, moist open slopes and thickets, 1,000-2,500 m.;
Chiquimula; Huehuetenango; Jalapa [Alta Verapaz, fide Morton].
Southern Mexico; Honduras; Costa Rica; West Indies; Colombia
and Venezuela southward to Bolivia; Old World tropics.
Plants terrestrial; rhizome erect, cylindrical, commonly bearing a single leaf;
mature plant 7-34 cm. tall; common stalk 3-12 cm. long with spike 1.2-5.5 cm. long
and 2-3.5 mm. thick; sterile lamina 2-8 cm. long, 1.2-5.2 cm. wide, ovate, deltoid or
subreniform, obtuse to acute, base cordate, truncate, often short- to long- (12 mm.)
attenuate, with midvein lacking; venation evident, diffuse, reticulate, the areoles
commonly with free, included veinlets.
The fact that plants of this species are so much larger, and there-
fore more conspicuous, than those of O. nudicaule and O. crotalo-
phoroides, perhaps accounts for its greater representation in her-
baria. However, more careful searching for all the species of the
genus will certainly reveal a much wider distribution of the indi-
vidual taxa.
MARATTIACEAE
Reference: L. M. Underwood, Marattiaceae, in North Amer. Fl.
16: 15-23. 1916.
Terrestrial plants with fleshy, creeping or erect rhizomes; leaves small to huge,
circulate in vernation, usually compound, occasionally dimorphic, stipulate, petiole
swollen and articulate at the rhizome; lamina coarse, with pinnae jointed to the
rachis, often with swollen nodes; venation free, or rarely reticulate; sporangia borne
on the abaxial surface of the leaves, with walls several cells thick, lacking an annu-
lus, separate and contiguous or coalescing into thick, circular or (in ours) elongate,
double-rowed synangia, and opening by terminal pores or longitudinal slits; spores
uniform, monolete or trilete, numerous (1,500-7,000) in each sporangium.
This distinctive and natural family is pantropic in distribution. Of
the six or seven recognized genera, only two occur in the New
World, and both of these are represented in Guatemala. The family
is of no economic importance.
14 FIELDIANA: BOTANY, VOLUME 39
a. Leaves 1-pinnate, dimorphic; synangia nearly covering the abaxial surface of the
fertile pinnae Danaea.
a. Leaves 3-pinnate, uniform; synangia scattered, borne near the ends of the
veins Marattia.
DANAEA J. E. Smith
Reference: L. M. Underwood, A review of the genus Danaea, Bull.
Torrey Bot. Club 29: 669-679. 1902.
Plants terrestrial, with rhizomes horizontal or obliquely ascending; leaves distich-
ous, dimorphous, simple or (in ours) 1-pinnate; primary axis nodose, the lower nodes
often lacking pinnae, provided with reddish, appressed, peltate, often suborbicular
scales; sterile pinnae subfalcate, sessile or short-stalked (to 5 mm.), slightly undu-
late and entire, or serrate near the acuminate or caudate apex; lamina provided with
scales on the costae and tissue beneath, those of the tissue commonly fimbriate,
widely scattered and extremely minute, veins free, paired or once-forked at or near
the costa; fertile pinnae similar in shape to the sterile ones, but somewhat reduced in
size; sporangia nearly covering the abaxial surface, coalescing into elongate, double-
rowed synangia which open by terminal pores ; spores monolete.
A distinctive genus, confined to the American tropics, easily rec-
ognized by the nodose axes, dimorphous leaves with mostly oppo-
site pinnae, synangia nearly covering the fertile pinnae beneath, and
the usually once-pinnate blades. Of the 30-35 species in the genus,
only three are known from Guatemala.
a. Petiole lacking nodes; larger sterile pinnae 20-40 cm. long; synangia more than
150 pairs on each fertile pinna D. nodosa.
a. Petiole nodose; sterile pinnae commonly less than 20 cm. long; synangia less
than 100 pairs on each fertile pinna,
b. Sterile pinnae 6 pairs or less, larger ones 3-4.5 cm. broad, subentire throughout;
not found over 600 m D. elliptica.
b. Sterile pinnae 10-18 pairs, not more than 2 cm. broad, serrate at apex; 1,200-
2,100 m D. cuspidata.
Danaea cuspidata Liebm., Kongel. Danske Vidensk. Selsk. Skr.
V. (l):307(seors. 155). 1849.
Wet forest floor, often in shaded ravines, 1,200-2,100 m. Alta
Verapaz; Baja Verapaz; Huehuetenango. Southern Mexico; Costa
Rica; Panama.
Plants to 1 m. tall, terrestrial, occasionally climbing; primary axis with several
nodes below the lamina; lamina with 10-18 pairs of pinnae, larger sterile ones 9-12
cm. long, 1.5-2 cm. broad, narrow-elliptic or -oblong, with base obtuse, truncate or
broadly cuneate, apex serrate, acuminate; laminar scales widely scattered along the
veins and on the tissue beneath, minute, suborbicular, entire to erose; fertile pinnae
similar in shape to sterile ones, 4.5-8 cm. long, 0.6-0.8 cm. broad; synangia ca. 100
pairs per pinna.
FIG. 3. Danaea, habit (sterile leaves), a, D. cuspidata, X V* (inset X I'/i); b, D.
elliptica, X Vi; c,D. nodosa, X 1/2.
15
16 FIELDIANA: BOTANY, VOLUME 39
A minute but interesting feature of our species of Danaea may be
found on the veins and tissue of the abaxial side of the pinnae. What
at first appear to be widely scattered, red resin dots are revealed
(under high magnification) to be microscopic scales 0.1 mm. or less.
Reddish, appressed, suborbicular and peltate, they are similar to
the larger scales of the primary axis. Those of D. cuspidata, how-
ever, have subentire to erose margins, whereas those of D. elliptica
and D. nodosa are commonly fimbriate, often so deeply that they
present a stellate appearance.
Danaea elliptica J. E. Sm., in Rees Cycl. 11: Danaea No. 2. 1808.
D. media Liebm., Kongel. Danske Vidensk. Selsk. Skr. V (1): 306.
1849.
Moist ground, in forests, 60-600 m. Alta Verapaz; Huehuetenan-
go. West Indies; southern Mexico; British Honduras; Honduras;
Nicaragua; Costa Rica; Panama; the Guianas; Colombia; Brazil;
Peru; Bolivia.
Plants to 1 m. tall, terrestrial; primary axis with several nodes below the lamina;
lamina with six pairs of pinnae or less, larger sterile ones 12-20 cm. long, 3-4.5 cm.
broad, narrowly oblanceolate to mostly elliptic, margin subentire throughout, with
base cuneate, apex acuminate; laminar scales minute, suborbicular, mostly fimbri-
ate, widely scattered along the veins and on the tissue beneath: fertile pinnae
similar to sterile ones in shape, 9.5-11.5 cm. long, 2.3-2.8 cm. broad; synangia ca.
90-100 pairs per pinna.
Danaea nodosa (L.) J. E. Sm. Mem. Acad. Roy. Sci. (Turin) 5:
420. l793.Acrostichum nodosum L., Sp. PI. 1070. 1753.
Low wet ground, often in ravines, in forests, sea level to 950 m.
Alta Verapaz; Izabal. West Indies; southern Mexico; British Hon-
duras; Honduras; Nicaragua; Costa Rica; Panama; southward to
Peru and Brazil.
Plants to 2 m. tall, terrestrial; petiole lacking nodes; lamina with 7-11 pairs of pin-
nae, larger sterile ones 20-40 cm. long, to 4.5 cm. broad, narrowly elliptic, oblong, or
oblanceolate, margin subentire throughout, broadly cuneate at base, with an acumi-
nate to caudate apex; laminar scales minute, suborbicular, mostly fimbriate, widely
scattered along the veins and on the tissue beneath; fertile pinnae similar to the
sterile ones in shape, to 26 cm. long and 2 cm. broad; synangia ca. 150-170 pairs per
pinna.
MARATTIA Swartz
Plants terrestrial, with stout, erect rhizomes; leaves deltoid, 2- to 3-pinnate, or
rarely 4-pinnate as to some of the larger pinnae, the fertile and sterile leaves alike:
STOLZE: FERNS OF GUATEMALA 17
major axes stramineous to brownish, with sparse, brown, narrow, usually attenuate
scales, the scales becoming larger, broader and abundant toward petiole base; terti-
ary axes ( in ours) irregularly winged, narrowing at the base of each segment so as to
present a scalloped outline, glabrous, or provided with pale or brown attenuate
scales, or with slender, tortuous, reddish-brown trichomes or hairlike scales; veins
free, simple to several times branched; sporangia in two rows, opening by longi-
tudinal slits, coalescent into 2-valved, oval synangia, which are borne near the ends
of the veins on a receptacle, short-stalked or (in ours) sessile, sometimes subtended
by slender, tortuous scales or trichomes; spores monolete.
A pantropical genus of about 50 species, centered chiefly in the
East Indies. The problem of Marattia in Central America requires
much closer examination and more field studies. A meaningful
classification must be based on better and more complete collec-
tions. Most species with sessile synangia apparently differ only in
characters of size and shape, and it is likely there are but one or two
species where four or five have been previously recognized. Even
size and shape are of questionable value in delimiting species, since
tertiary segments vary between specimens in gradual, almost
imperceptible stages, from short, narrow and obtuse, to long, broad,
and acute to acuminate. Furthermore, most specimens currently
available consist merely of a single pinna, with no label data as to
where the pinna was located on the leaf. Obviously, the shape and
size of tertiary segments of basal pinnae can differ greatly from
those of pinnae near the leaf apex.
The following four species are tentatively recognized in Guate-
mala, although the distinctions are admittedly arbitrary.
a. Scales abundant on midribs and veins of tertiary segments M. alata.
a. Scales lacking (or a few hairlike scales or trichomes widely scattered) on midribs
and veins of tertiary segments,
b. Tertiary segments commonly acute to acuminate, krger ones 24-60 mm. long
and 10-20 mm. broad M. interposita.
b. Tertiary segments rounded or truncate to subacute (or rarely a few segments
acute), to 20 (25) mm. long and 10 mm. broad.
c. Larger tertiary segments 15-20 (25) mm. long and (6) 7-10 mm. broad
M. excavata.
c. Larger tertiary segments 1 1-15 mm. long and 4-6 ( 7) mm. broad
M. iL-einmanniifolia.
Marattia alata Sw. Prodr. 128. 1788 (not M. alata Raddi. 1825).
M. laevis J. E. Sm. PL Ic. Ined. 2: t. 47. 1790 (notM. laevis Kaulf.
1824).
Although this is essentially a West Indian species, at least one
specimen (Ghiesbreght 241} has been found in Chiapas; hence it is
FlG. 4. Marattia. a-b, M. excavata: a, habit, X !/?; b, ultimate segments, X 2; c-e,
M. interposita: c, habit, X V?; d, ultimate segments, X 2; e, synangium, X 12.
18
STOLZE: FERNS OF GUATEMALA 19
to be expected in Guatemala. Dense wet forests and wooded slopes,
1,100-1, 800m. Cuba; Jamaica; Hispaniola; southern Mexico.
Rhizome and petiole base not seen; upper petiole, rachis and costae sparsely pro-
vided with broad, mostly attenuate, orange- to red-brown scales; leaves several
meters long, 3- (4-) pinnate, subcoriaceous, tissue glabrous and glands generally
lacking above, glabrous beneath, but costules, midribs and veins rather thickly cov-
ered with rusty brown, broad, attenuate or hairlike scales; ultimate (tertiary) seg-
ments to 20 mm. long and 9 mm. broad, diminishing to 5 mm. long and 2 mm. broad
at pinnule apices, subsessile, oblong or narrowly ovate, with apices rounded to
acute, margin serrate ( commonly deeply so), the bases somewhat obliquely excavate
on the acroscopic side; veins of the tertiary segments simple to once-forked, com-
monly distinct; synangia 8- to 16-locular, submarginal or supramedial (rarely medi-
al) on the segments, usually subtended by slender, brown scales.
Indument is a variable and inconsistent character among species
of Marattia, and generally of little taxonomic value. However,
broad, rusty-brown scales are so abundant on veins and midribs of
segments in M. (data that it can be readily distinguished from all
others in Guatemala, on this character alone. Indument, if any,
found on veins and midribs of other species consists of minute,
widely scattered trichomes or hairlike scales, which are sometimes
detectable only under high magnification.
Marattia excavata Underw. North Amer. Fl. 16: 22. 1909.
Dense, wet montane or cloud forests, 1,300-2,850 m., Alta Vera-
paz; Baja Verapaz; Quezaltenango; El Quiche; San Marcos; Suchi-
tepequez; Zacapa. Mexico (Chiapas); Honduras; El Salvador; Nic-
aragua; Costa Rica.
Rhizome erect, to 40 cm. tall; petiole base and croziers rather thickly invested
with large, broad, yellowish brown to reddish brown scales; upper petiole, rachis
and costae nearly glabrous, provided with scattered, broad, attenuate, pale to light
brown scales or slender, tortuous, hairlike ones; leaves 2-3 m. long, commonly 3-
pinnate, subcoriaceous, tissue glabrous above, sometimes thickly dotted with
minute reddish or yellowish glands, glabrous beneath, or with pale to reddish brown,
hairlike scales or tortuous trichomes scattered along the midribs and veins; ulti-
mate (tertiary) segments to 20 (25) mm. long and 10 mm. broad, diminishing to 8
mm. long and 4 mm. broad at pinnule apices, sessile, oblong to narrowly ovate, often
subfalcate, with apices rounded or truncate to subacute, margin serrate, the bases
obliquely excavate on the acroscopic side; veins of the tertiary segments simple or
once-forked, usually distinct, occasionally somewhat obscure; synangia 10- to 18-
locular, supramedial to submarginal on the segments, sometimes sparsely subten-
ded by hairlike scales.
This species lies midway between M. weinmannii folia and M. in-
terposita in size and shape. All other characters in the three taxa are
highly variable and inconsistent, and it is likely that future mono-
graphic study will reveal that they are conspecific— varieties at
20 FIELDIANA: BOTANY, VOLUME 39
best. The tertiary segments of M. weinmannii folia are relatively
small and rounded, with simple to rarely once-forked veins that are
commonly obscure. In M. excavata, the segments are somewhat
larger, often subacute (rarely acute), with veins sometimes simple,
but commonly once-forked, and usually quite distinct. The seg-
ments of M. interposita are extremely large, acute to long-acumi-
nate, with veins often several-times forked. None of these charac-
ters appear, by themselves, to be taxonomically significant, and are
rather a manifestation of the effect of increased size, due either to
maturity of the individual plant or to ecological or geographical
variation.
Marattia interposita Christ, Bull. Herb. Boissier II (6): 285. 1906.
Casco de burro, maigre maiz (fide Steyermark, Quezaltenango).
Damp, usually dense, montane or cloud forests, or in forested que-
bradas, 1,300-2,800 m.; Baja Verapaz; Quezaltenango; San Marcos.
Honduras; Costa Rica; probably Panama and Colombia.
Rhizome and petiole base not seen; upper petiole, rachis and costae mostly glab-
rous, or sparsely provided with broad to slender, usually attenuate, pale to brown
scales; leaves to 2 m. long, 3- (4-) pinnate, subcoriaceous, tissue glabrous above,
often thickly dotted with minute, reddish or yellowish glands, glabrous beneath, or
with pale to brown hairlike scales or tortuous trichomes widely scattered along the
midribs and veins; ultimate (tertiary) segments to 60 mm. long and 20 mm. broad,
diminishing to 12 mm. long and 6 mm. broad at pinnule apices, subsessile, oblong to
narrowly ovate, often subfalcate, with apices acute to acuminate, margin serrate,
and the bases obliquely excavate on the acroscopic side; veins of the tertiary seg-
ments simple, or more commonly 1- or 2-(3-) forked, distinct to somewhat obscure;
synangia 10- to 18-locular, mostly submarginal on the segments, rarely sparsely
subtended by slender scales.
With this species should probably be included M. chiricana from
Panama, which is said to differ in its greater size and larger, less
marginal synangia.
It is said that the scaly rootstocks of M. interposita are some-
times cooked and eaten in times when the corn crop is poor. Also,
croziers and petiole bases are reportedly cooked and mixed with
tortillas.
Marattia weinmanniifolia Liebm. Kongel. Danske Vidensk. Selsk.
Skr. V. 1: 308 (seors.156). 1849.
Dense, wet forest, 1,200-1,700 m. Alta Verapaz. Southern Mexico;
Honduras; El Salvador. Type from Villa Alta, Oaxaca, Mexico,
Liebmann Fl. Mex. 651.
STOLZE: FERNS OF GUATEMALA 21
Rhizome and petiole base not seen; upper petiole, rachis and costae sparsely to
abundantly provided with broad and attenuate, or hairlike, brown scales; leaves to
2 m. long, commonly 3-pinnate, chartaceous to subcoriaceous, tissue glabrous and
glands generally lacking above, glabrous beneath, or with a few brown trichomes or
hairlike scales widely scattered on veins and midribs; ultimate (tertiary) segments
to 16 mm. long and 6 mm. broad, diminishing to 5 mm. long and 2 mm. broad at pin-
nule apices, sessile or subsessile, oblong to narrowly ovate, often subfalcate, with
apices rounded to subacute, margins subentire to crenulate or serrate, the bases
somewhat obliquely excavate on the acroscopic side; veins of the tertiary segments
simple, or once-forked, faint or obscure (or rarely distinct); synangia 10- to 15-locu-
lar, inframedial to medial, or most commonly submarginal on the segments, some-
times sparsely subtended by hairlike scales.
OSMUNDACEAE
«
Reference: R. C. Benedict, Osmundaceae, in North Amer. Fl. 16:
27-28. 1916.
Coarse, terrestrial plants with woody, usually erect, rhizomes and hard, fibrous
roots ; leaves large, circulate in vernation, densely caespitose, petiole not articulate
at the rhizome, pinnately compound, alike or (in ours) completely dimorphous or
with dimorphous pinnae, with tissue at base of petiole expanded into sheathlike
wings; venation free; sporangia not in definite sori, borne abaxially on the segments
or (in ours) completely replacing the vegetative tissue of some pinnae or entire lami-
nae, exindusiate, short-stalked, globose or pyriform, with walls one cell thick, annu-
lus lacking or of only a few thickened cells near the distal end, longitudinally dehis-
cent; spores uniform, green, trilete, relatively numerous (120-512) in each sporan-
gium.
Considered to be among the most primitive of families, with many
characters evidently intermediate between the leptosporangiate and
eusporangiate ferns; however it is quite distinctive and natural and
has no close affinity with any other living fern family. The three ( to
five) genera of the Osmundaceae, containing 15-20 species, may be
found in both tropical and temperate regions throughout the world,
but the family is represented in the New World by only a few species
of Osmunda.
OSMUNDA Linnaeus
Mature leaves quite coarse and large, pinnately compound, completely dimorphous
or with dimorphous pinnae, arising in two rings from a stout rhizome, the inner ring
fertile and developing first, with leaves mostly erect, the outer one sterile, with the
leaves somewhat spreading; fertile leaves or pinnae lacking green leaf tissue; veins
free, at least once-forked; leaf tissue commonly glabrous, but the axes often sparse-
ly to moderately invested with trichomes on the adaxial side; rachis or costa (at
least in ours) with a thin, green wing of tissue along each side, arising from the adax-
22 FIELDIANA: BOTANY, VOLUME 39
ial edges; sporangia relatively large, developing simultaneously, dark brown to red-
dish brown, densely crowded on the segments of the fertile blades.
The genus contains about 12 species which are generally found in
swampy areas in temperate and tropical regions around the world.
Larger rhizomes of Osmunda, with their dense mats of coarse roots,
are widely used in the culture of orchids and other commercially
popular epiphytes. Two species occur in Guatemala.
a. Sterile laminae pinnate-pinnatisect; ultimate segments entire, the veins common-
ly once-forked; fertile laminae non-foliaceous throughout O. cinnamomea.
a. Sterile laminae bipinnate; ultimate segments serrulate, the veins commonly
twice- forked; fertile laminae non-foliaceous and fertile only in the apical por-
tion O. regalis var. spectabilis.
Osmunda cinnamomea L. Sp. PL 1066. 1753. O. bipinnata L. Sp.
PL 1065. 1753.
Mostly wooded swamps and bogs, 1,250-1,500 m.; Alta Verapaz.
Eastern and central United States and Canada; southern Mexico;
Honduras; Costa Rica; West Indies; Colombia; Venezuela; Brazil;
Peru; Paraguay; Southeast Asia.
Mature leaves to 1.5 m. tall, completely dimorphous; petiole to 50 cm. long, stra-
mineous to reddish brown, darker toward the base, glabrous or with some scattered,
reddish brown, tortuous, pluricellular trichomes; sterile lamina to 80 cm. long and
20 cm. broad, elliptic to broadly lanceolate, pinnate-pinnatisect; texture chartace-
ous to subcoriaceous; rachis stramineous to light brown, with scattered trichomes,
a thin wing of tissue connecting adjacent pinnae; pinnae to 13 cm. long and 2 cm.
broad, spreading to strongly ascending, subopposite, articulate at rachis, reddish
brown trichomes scattered along the costa and in very dense clusters at pinna base;
ultimate segments to 1 cm. long and 0.6 cm. broad, cut nearly to costa, subfalcate,
obtuse to subacute, margins entire and bearing abundant minute trichomes; veins
once-forked; fertile leaves bipinnate due to reduction of leaf tissue, very narrow
(pinnae to 3 cm. long), rachis generally darker-colored than that of sterile pinnae and
with a greater abundance of trichomes; trichomes very thick among the crowded,
dark reddish brown sporangia.
Osmunda regalis L. var. spectabilis (Willd.) Gray, Manual, ed. 2:
600. 1856. O. spectabilis Willd. Sp. PL 5: 98. 1810. O. mexicana Fee,
Mem. Fam. Foug. 9: 43. 1857.
In wet thickets, wooded swamps, and along shaded river banks,
1,200-2,100 m.; Alta Verapaz; Baja Verapaz; Chiquimula; Huehue-
tenango; El Quiche; Santa Rosa. Eastern and central United States
and Canada; West Indies; southern Mexico; Honduras; Costa
Rica; Colombia; Venezuela; Brazil; Ecuador; Peru; Paraguay;
Uruguay; Old World.
FIG. 5. Osmunda. a-b, O. regalis var. spectabilis: a, habit, X Vi; b, ultimate seg-
ment, X 2!/2; c-d, O. cinnamomea: c, habit, X Vz; d, ultimate segment, X 2'/2.
23
24 FIELDIANA: BOTANY, VOLUME 39
Mature leaves to 1.8 m. tall, partly dimorphous, fertile leaves with fertile and
sterile portions combined on the same blade, the fertile pinnae commonly apical;
petiole glabrous, to 75 cm. long, stramineous or light brown to reddish brown, usu-
ally darkest toward the base; sterile blade to 80 cm. long and 35 cm. broad, elliptic
or oblong, bipinnate, chartaceous to subcoriaceous, tissue glabrous; rachis stramin-
eous or light brown, non-alate, glabrous, or with minute trichomes scattered widely
on the adaxial side; pinnae to 30 cm. long, strongly ascending, subopposite, articu-
late at rachis, with trichomes of two kinds, minute (0.1-0.2 mm.), light brown,
mostly unicellular ones scattered along the adaxial side of the costae and costules,
and some longer (to 1 mm.), tortuous, reddish brown, pluricellular ones clustered at
the bases of pinnules; costae narrowly alate (best observed from adaxial side); pin-
nules distant, short-stalked and articulate at the costae, subopposite, 7 to 15 pairs
per pinna, 1.5-5.5 cm. long and 0.5-1.7 cm. broad, oblong to narrow-elliptic, obtuse to
acute, broadly rounded or, more commonly, truncate at base, margins serrulate;
veins mostly twice-forked; fertile pinnae essentially bipinnate, the crowded, reddish
brown sporangia almost completely replacing the tissue.
This variety differs from the typical in its narrower, more coriace-
ous leaves, broader panicles, and rachis glabrous or with dark-col-
ored trichomes only at the bases of pinnules and (rarely) pinnae.
The typical variety, occurring in parts of Eurasia, has black, hair-
like scales more or less persistent along the rachis. Some workers
also separate the South American and southern Central American
representatives of var. spectabilis into yet another variety, palus-
tris Schrad., based on the size of the segments and length of their
stalks. However, these characters appear to be too variable and in-
consistent to provide sufficient varietal distinction.
SCHIZAEACEAE
References: K. Prantl, Untersuch. Morph. Gefasskrypt. II: Schiz-
aceen, W. Engelmann, Leipzig, 1881; W. R. Maxon, Schizaeaceae,
in North Amer. Fl. 16: 31-52. 1909.
Coarse to delicate, terrestrial plants of highly diverse habit and shape, with rhi-
zomes oblique, ascending or horizontal, sometimes short- or long-creeping ( dichoto-
mously branched in Lygodium), rather densely clothed with pale to dark brown,
stiff or lax, one- to several-celled trichomes; leaves small and inconspicuous,
medium-sized, or large and twining, circinate in vernation, closely to widely spaced,
or densely caespitose, petiole not articulate at the rhizome, sterile and fertile ones
alike, or partly to fully dimorphous; lamina absent to scarcely or normally foliace-
ous, linear, dichotomously branched, flabelliform, or pinna tely arranged; veins com-
monly free, or in a few species reticulate; sporangia sessile, each with uninterrupted
distal annulus, dehiscing longitudinally, marginal (though often appearing to be
superficial), not arranged in definite sori, commonly borne in rows on specialized,
scarcely- or non-foliaceous fertile segments, with true indusia lacking or consisting
of modified, sometimes scalelike, marginal tissue; spores monolete or trilete, vari-
ously sculptured, from 32 to 256 in each sporangium.
STOLZE: FERNS OF GUATEMALA 25
An extremely interesting family with cosmopolitan distribution,
composed of three to six genera of highly diverse habit and form.
Some authors prefer to recognize Anemia and Lygodium as fami-
lies; some include Actinostachys and Lophidium merely as sections
of Schizoea. Four genera are recognized from Guatemala in this
treatment.
a. Leaves vinelike, twining and scrambling; secondary pinnae arising in pairs from
very short stalks on the primary rachis Lygodium.
a. Leaves erect or suberect; pinnae not arising as above.
b. Sterile laminae pinnate to tripinnate; sporangia borne on panicles .... Anemia.
b. Sterile laminae simple and grasslike, or several times dichotomous; sporangia
borne on terminal, linear segments, these pinnately or (apparently) digitately
arranged.
c. Sterile laminae simple, grasslike, not lobed or branched; sporangiophores
apparently digitate Actinostachys.
c. Sterile laminae once to several times dichotomous, flabelliform (at least in
outline) ; sporangiophores arranged in pinnate spikes Schizaea.
ACTINOSTACHYS Wallich ex Hooker
References: Olof H. Selling, Studies in the recent and fossil spe-
cies of Schizaea . . . Acta Horti Gothob. 16: 1-112. 1944; David W.
Bierhorst, Leaf development in Schizaea and Actinostachys, Amer.
J. Bot. 56: 860-870. 1969.
Terrestrial, erect ferns; rhizome horizontal to ascending, thickly invested with
one- to several-celled, brownish trichomes; leaves very densely caespitose (or rarely
rather widely spaced); petiole triangular, or plane with a narrow, foliaceous wing on
either side, glabrous, pale green, or brown toward the base; lamina linear, grasslike
and scarcely or not at all foliaceous, glabrous, acute to rounded at apex; sporangio-
phores terminal, apparently digitate, with sporangia crowded in apparently four
rows on the narrowly foliaceous, indusiform segments; spores monolete.
About 15 species, generally distributed in tropical or subtropical
regions of both hemispheres, with a majority occurring in the Pacif-
ic Islands.
Actinostachys has been treated by some authors as one of three
sections ( with Euschizaea and Lophidium ) of the genus Schizaea.
However, as pointed out by Bierhorst (1969), the differences in the
arrangement and development of the fertile segments, reinforced by
distinctions in embryo and gametophyte, appear to warrant a gener-
ic separation at least in Actinostachys.
The following is the only species heretofore reported from Guate-
mala.
26
FIELDIANA: BOTANY, VOLUME 39
FIG. 6. Actinostachys germanii. a, habit, X Vz; b, sporangiophore, X 2Vz; c,
sporangiophore segment, X 25.
Actinostachys germanii Fee, Mem. Fam. Foug. 11: 123. 1866.
Schizaea germani Prantl, Unters. Morph. Gefass. Shiz. II: 132.
1881.
Known in Guatemala from only one collection, Steyermark 39864,
lowland jungle, at sea level, Punta Palma, Dept. Izabal, and one
from British Honduras, Peck 936, mangrove swamp, New Haven.
Southern Florida; West Indies; Trinidad; Colombia; Brazil.
Rhizome ascending, provided with stiff, orange to yellowish brown 1- or 2-celled
trichomes, which occasionally extend to the lower petiole; leaves several, 8-20 cm.
long, erect, caespitose, scarcely foliaceous; petiole generally glabrous, subterete or
triangular at the dark brown base, gradually altering above to a flattened, grasslike
lamina consisting of a rachis flanked by a pair of narrow wings of tissue, rarely more
than 1 mm. broad, subacute at apex; sporangiophores terminal, apparently digitate,
segments 3-8, subequal, slender, erect or recurved, 10-15 mm. long, to 0.7 mm.
broad, with yellowish brown, filiform trichomes arising around and among the
sporangia.
The species of Actinostachys are among the least conspicuous of
all the ferns and undoubtedly have been overlooked repeatedly in
the field. This would explain the apparent disjunct distribution of
A. germanii; hence it is to be expected this species will be found
eventually in other departments of Guatemala, as well as in many
other areas of the American Tropics.
STOLZE: FERNS OF GUATEMALA 27
ANEMIA Swartz
Reference: J. T. Mickel, A monographic study of the fern genus
Anemia, subgenus Coptophyllum, Iowa State Coll. Sci. 36 (4): 349-
482. 1962.
Terrestrial, erect ferns; rhizome horizontal, ascending, or creeping, clothed with
orange, reddish, or blackish, filiform to acicular, pluricellular trichomes; leaves poly-
stichous and caespitose, or distichous and closely to widely spaced, fully dimor-
phous (i.e., either completely fertile or completely sterile), or partly dimorphous,
with only the basal pair of pinnae fertile, to 90 cm. long (including the elongate fer-
tile pinnae); petiole and rachis stramineous to dark brown, often darker at base,
delicate to wiry to rather stout, terete, rounded beneath, sulcate above, glabrate, or
lightly to densely villous with pale to reddish, usually tortuous, trichomes; sterile
lamina or portion of lamina oblong to deltoid, pinnate to tripinnate, firm-membrana-
ceous to coriaceous, glabrous, or lightly to densely villous as on the rachis, the
trichomes often shorter and fewer-celled or lacking on the adaxial side, veins free or
rarely reticulate, commonly oblique and approximate, several to many times dichoto-
mous; fertile pinnae long-stalked, usually elongate and erect, often surpassing the
sterile blade, 2- to 3-pinnate, sporangia borne in 2 rows on the ultimate divisions,
these either slender and scarcely foliaceous, or narrow-foliaceous and indusiform,
but true indusia lacking; spores trilete.
The genus contains 70 to 80 species, occurring in tropical to sub-
tropical regions in both hemispheres, predominantly in the neotrop-
ics. The plants are most often found growing on rocks or in rocky
situations.
a. Leaves completely dimorphous, i.e., either wholly fertile or sterile. . . A. cicutaria.
a. Leaves partly dimorphous, i.e., fertile leaves with only the basal pair of pinnae
soriferous.
b. Fertile pinnae borne below the base of sterile lamina, i.e., sterile lamina short-
stalked; ultimate fertile divisions narrow-foliaceous and indusiform.
c. Sterile pinnae 1-4 pairs, merely serrate A. speciosa.
c. Sterile pinnae 8 to many pairs, deeply incised or compound,
d. Leaves distichous, closely- to widely-spaced; rhizome with dark brown or
blackish trichomes A. adiantifotia.
d. Leaves polystichous, densely caespitose; rhizomes with orange or reddish
trichomes.
e. Sterile lamina bipinnate-pinnatifid to tripinnate; rhizome with orange
trichomes A. guatemalensis.
e. Sterile lamina bipinnate (rarely bipinnate-pinnatifid as to lowermost pair
of pinnae); rhizome with reddish brown trichomes
A. bartlettii.
b. Fertile pinnae borne at base of sterile lamina, i.e., sterile lamina sessile; ulti-
mate fertile divisions slender, not or scarcely foliaceous.
f. Veins reticulate; panicle of fertile pinna as long as the stalk or (usually)
longer A. phyllitidis.
f . Veins free ; panicle of fertile pinna shorter than ( or rarely equalling) the stalk.
28 FIELDIANA: BOTANY, VOLUME 39
g. Pinnae mostly deeply incised, the segments commonly narrow and cune-
ate A. hirsuta.
g. Pinnae merely crenulate or denticulate, or rarely with a few deep lobes on
some lower pinnae.
h. Petiole of fertile leaf usually greatly exceeding the sterile leaf; lamina
gradually tapering to a pinnatifid apex; rhizome horizontal, short-
creeping A. pastinacaria.
h. Petiole of fertile leaf scarcely (if ever) exceeding the sterile leaf; lamina
abruptly terminating in an epongate to acuminate or obovate to fla-
belliform apical segment; rhizome oblique to ascending,
i. Sterile lamina deltoid or ovate-deltoid, terminating in an acuminate or
elongate apical segment; pinnae subacute to acute, mostly short-
stalked; leaves petiolate A. hirta.
i. Sterile lamina obovate-oblong or obovate, terminating in a broadly
obovate or flabelliform apical segment; pinnae broadly rounded,
sessile; sterile leaves often sessile, forming a rosette. . . A. oblongifolia.
Anemia adiantifolia (L.) Sw. Syn. Fil. 157. 1806. Osmunda adian-
tifolia L. Sp. PL 1065. 1753. O. asplenifolia Sav. in Lam. Encycl. 4:
652. 1797. Ornithopteris adiantifolia (L.) Bernh. Neues J. Bot. 1:
50. 1806. Anemia asplenifolia Sw. Syn. Fil. 157. 1806. Cola (fide
Steyermark, Jalapa).
Rocky, grassy, or partly wooded places, usually on limestone,
0-1,900 m. ; Alta Verapaz; Chiquimula; Huehuetenango; Izabal;
Jalapa; Peten; Santa Rosa. Peninsular Florida; West Indies; Mexi-
co; British Honduras; Honduras; Costa Rica; also reported from
Colombia and Brazil, but specimens not seen.
Rhizome creeping, rather densely clothed with dark brown or blackish, acicular
trichomes 2-4 mm. long, these commonly extending to the petiole base; leaves to 65
cm. long (including the erect, fertile pinnae) and 30 cm. broad, distichous, often
widely spaced, partly dimorphous, the fertile pinnae borne 0.5-2 cm. below the sterile
portion of the lamina, sterile leaf similar in size and shape to the fertile, often a little
shorter; petiole of fertile leaf to 45 cm. long, rarely exceeding the sterile leaf, stra-
mineous, darker toward base, or rarely brownish throughout, rather stout, pale to
dark villous, glabrate except at base; sterile portion of lamina ovate-deltoid, bipin-
nate to (commonly) bipinnate-pinnatifid (or tripinnate at base), subcoriaceous, lus-
trous on both sides, with short, stout, pale trichomes widely scattered on veins and
tissue, glabrate above; sterile pinnae anadromous, numerous, ascending, all but
upper ones stalked, basal ones to 11 cm. long and 6 cm. broad, the others gradually
diminishing in size to a pinnatisect apex; pinnules with ultimate segments common-
ly obovate or oblanceolate (or rarely linear-obovate or linear-oblong), cuneate,
rounded to subtruncate at apex, margins erose-denticulate, slightly cartilaginous;
veins free, evident, raised; fertile pinnae to 25 cm. long, panicle commonly longer
than its stramineous or light brown stalk, mostly tripinnate, segments narrow-
foliaceous and indusiform.
FlG. 1. Anemia, a, A. cicutaria, habit, X '/i; b, A. speciosa, habit, X 1/2; c, A. pas-
tinacaria, habit, X %; d, A. hirsuta, portion of sterile lamina, X Va; e, A. hirta, por-
tion of sterile lamina, X 1/2 ; f , A. oblongifolia, portion of sterile lamina, X 1/2.
29
30 FIELDIANA: BOTANY, VOLUME 39
Anemia bartlettii Mickel, Iowa State Coll. J. Sci. 36 (4): 420.
1962.
Ravine, mountain pine ridge, 450 m., El Cayo District, British
Honduras (type, Bartlett 11898). Known only from the type and one
other specimen (Hunt 428) from type locality.
Rhizome horizontal, densely clothed with reddish brown, filiform trichomes;
leaves to 70 cm. long (including the erect, fertile pinnae) and 15 cm. broad, polysti-
chous, caespitose, partly dimorphous, the fertile pinnae borne 1-2 cm. below sterile
portion of lamina, sterile leaf similar in size and shape to fertile, often somewhat
shorter; petiole of fertile leaf to 45 cm. long, exceeding the sterile leaf, stramineous,
atropurpureus toward base, or brownish throughout, rather stout, sparsely to
densely invested with orange or reddish, filiform trichomes, glabrate; sterile portion
of lamina ovate-deltoid, bipinnate, or rarely bipinnate-pinnatifid as to lowermost
pinnae, chartaceous, not lustrous, orange-pilose on both sides; sterile pinnae (at
least the lower ones) anadromous, 11-16 pairs, spreading to ascending, those of
lower half of lamina short-stalked, to 9 cm. long and 4 cm. broad, oblong, obtuse;
pinnules broadly adnate at base, obtuse at apex, crenate or a few on lowermost pin-
nae deeply pinnatifid, margin not or scarcely cartilaginous; veins free, evident,
slightly or not at all raised, a distinct midrib lacking or evident only at base of pin-
nule; fertile pinnae to 27 cm. long, panicle much longer than its brown stalk, mostly
tripinnate, segments narrow-foliaceous and indusiform.
In addition to the differences noted in the key, Anemia bartlettii
may be further distinguished from A. guatemalensis by the color of
the stalks of the fertile pinnae, which are darker than the main
rachis. In A. guatemalensis the stalks of the fertile pinnae are com-
monly stramineous, or rarely brownish, but in either case they are
concolorous with the rachis.
Anemia ci cut aria Kunze, Sprengel Syst. Veg. 4: 31. 1827. A. bi-
pinnata Moore, Index Fil. 66. 1857 (not Swartz, 1806). Ornithop-
teris cicutaria (Kunze) Underw. Mem. Torrey Bot. Club 12: 15.
1902.
Among rocks, in forests, ca. 120 m.; Peten. Mexico (Yucatan
Peninsula and Cozumel Island); Bahama Islands; Cuba.
Rhizome horizontal, short-creeping, with dense tufts of dark- to reddish-brown
trichomes; leaves closely grouped, obscurely distichous, fully dimorphous (i.e.,
either completely fertile or completely sterile), the fertile much longer than the
sterile, petioles delicate, stramineous, or occasionally brown at base; sterile leaf to
15 cm. long and 5 cm. broad, petiole to 8 cm. long, mostly glabrate, lamina ovate to
deltoid-ovate, bipinnate to bipinnate-pinnatifid, firm- herbaceous, pilose (often
sparsely so) on both sides; pinnae 3-7 pairs, anadromous, subequilateral, spreading
to ascending, short-stalked, deltoid-ovate, obtuse to subacute, margins slightly
cartilaginous; pinnules mostly sessile, oblong to obovate to rhomboid, cuneate at
base, entire, or broadly dentate at apex; veins evident to somewhat obscure, slight-
ly raised; fertile leaf erect, to 25 cm. long, petiole to 15 cm. long, commonly much
STOLZE: FERNS OF GUATEMALA 31
longer than the fertile panicle, mostly tripinnate, the segments narrow-foliaceous
and indusiform.
This is the only species of Anemia in Central America with leaves
fully dimorphous. Thus far, it has been reported in Guatemala only
from Peten. However, the plants are relatively small and quite in-
conspicuous, and should eventually be found in other areas.
Anemia guatemalensis Maxon, North Amer. Fl. 16: 46. 1909.
Hemianemia guatemalensis (Maxon) Reed, Bol. Soc. Brot. 21 (2):
161. 1947.
Grassy banks, rocky hillsides, mostly in pine-oak forests, 1,000-
2,000 m.; Baja Verapaz; Chimaltenango; Escuintla; Guatemala;
Huehuetenango; Jalapa; El Quiche; Santa Rosa (type from Cerro
Gordo, Heyde & Lux 4095). Honduras; El Salvador.
Rhizome horizontal, densely clothed with orange trichomes; leaves to 65 cm. long
(including the erect, fertile pinnae) and 25 cm. broad, polystichous, caespitose,
partly dimorphous, the fertile pinnae borne 1-3 cm. below sterile portion of lamina,
sterile leaf similar in shape to fertile, commonly much shorter; petiole of fertile leaf
to 40 cm. long, not or slightly exceeding the sterile leaf, commonly stramineous
throughout, rather stout, sparsely to closely invested with pale to orange, filiform
trichomes, glabrate; sterile portion of lamina ovate-deltoid, bipinnate-pinnatifid to
tripinnate, subcoriaceous, not lustrous, pale- or orange-pilose beneath, sparsely so
or glabrous above; sterile pinnae 8-15 pairs, catadromous, or a few lower ones
anadromous, spreading to ascending, those of lower half of lamina short-stalked, to
12 cm. long and 7 cm. broad, ovate to lanceolate, subacute; pinnules mostly narrow-
adnate at base, obtuse to acute at apex, pinnatifid to pinnatisect, or those of lower
pinnae fully pinnate, margin not or scarcely cartilaginous; veins free, evident,
slightly or not at all raised, a distinct midrib lacking or evident only at base of seg-
ment; fertile pinnae to 21 cm. long, panicle much longer than its usually stramine-
ous stalk, mostly tripinnate, segments narrow-foliaceous and indusiform.
Anemia hirsuta (L.) Sw. Syn. Fil. 155. 1806. Osmunda hirsuta L.
Sp. PL 1064. 1753.
In rocky or grassy places, shaded or open slopes or in forests, 200-
2,200 m.; Guatemala; Huehuetenango; Jalapa; Jutiapa; Santa
Rosa; Solola; Zacapa. Greater Antilles; Trinidad and Tobago;
Mexico to Brazil and Bolivia.
Rhizome horizontal, short-creeping, densely clothed with orange, filiform tri-
chomes; leaves to 35 cm. long (including the erect, fertile pinnae) and 5 cm. broad,
polystichous, caespitose, partly dimorphous, the fertile pinnae and lowermost ster-
ile pinnae arising from the same position on the axis, sterile leaf similar in shape to
the fertile, but with much shorter petiole; petiole of fertile leaf to 18 cm. long, com-
monly much exceeding the sterile leaf, stramineous, often atropurpureus at base,
wiry, sparsely orange- villous, glabrate, sterile portion of lamina oblong-lanceolate
to ovate-oblong, gradually tapering to a pinnatifid apex, pinnate-pinna tifid to bipin-
32 FIELDIANA: BOTANY, VOLUME 39
nate, firm-herbaceous, dull, or somewhat lustrous abope, villous beneath, sparsely
villous or glabrate above; sterile pinnae 7-12 pairs, spreading or slightly ascending,
those of lower half of blade short-stalked, to 2.5 cm. long and 0.8 cm. broad, oblong
to ovate-oblong, obtuse to subacute, commonly inequilateral at base, truncate
above and cuneate below, obliquely and deeply incised into linear or narrowly cune-
ate segments, these irregularly toothed, margins not cartilaginous, a distinct midrib
lacking or evident only at pinna base; veins free, commonly obscure beneath, raised
and distinct above; fertile pinnae to 18 cm. long, panicle much shorter than (or
rarely equalling) the stramineous stalk, mostly tripinnate, ultimate divisions slen-
der and scarcely folia ceous.
Beyond the characters used in the key, there is little to distin-
guish Anemia hirsuta from A. pastinacaria, both of which share the
same general distribution, habitat, and sometimes even the same
herbarium sheet. Mickel (pers. comm. ) reports that he has found -A.
hirsuta, A. pastinacaria, and A. jaliscana growing together in Oaxa-
ca, and all apparently hybridizing readily. He also advises that A.
hirsuta crosses with A. tomentosa, A. karwinskyana, and A. phyl-
litidis in Mexico.
Anemia hirta (L.) Sw. Syn. Fil. 155. 1806. Osmunda hirta L. Sp.
PL 1064. 1753.
In forests, about 400 m. ; Peten. British Honduras; Honduras;
West Indies; Brazil.
Rhizome ascending, clothed at apex with deep orange or reddish, filiform tri-
chomes; leaves to 32 cm. long (including the erect, fertile pinnae) and 10 cm. broad,
polystichous, caespitose, partly dimorphous, the fertile pinnae and lowermost ster-
ile pinnae arising from same position on the axis, sterile leaf similar in shape to the
fertile, but with shorter petiole; petiole of fertile leaf to 20 cm. long, scarcely ( if ever)
exceeding the sterile leaf, stramineous, wiry to rather stout, rather densely rusty-
villous; sterile portion of lamina deltoid or ovate-deltoid, terminating in an acumi-
nate or elongate apical segment, pinnate, firm-membranaceous, dull to slightly lus-
trous, sparsely to densely villous, rachis rusty-villous like the petiole; sterile pinnae
7-12 pairs, spreading to slightly ascending, basal pair often deflexed, mostly short-
stalked, to 5.5 cm. long and 1.5 cm. broad, obliquely lanceolate, acute or subacute,
strongly inequilateral at base, truncate or rounded above and cuneate below, crenu-
late, margin slightly or not at all thickened, midrib distinct nearly to apex; veins
free, distinct, raised on adaxial side; fertile pinnae to 18 cm. long, panicle shorter
than (or rarely equalling) the stramineous stalk, mostly tripinnate, ultimate divi-
sions slender and scarcely foliaceous.
It is difficult to account for the curious disjunction in the distribu-
tion of this species. Among all collections examined, the only South
American representation was southeastern Brazil. Anemia hirta is
not a particularly inconspicuous species and it seems that it should
have been found in at least a few localities between Brazil and the
STOLZE: FERNS OF GUATEMALA 33
Lesser Antilles. However, Mickel noted in his monograph that A.
bartlettii has a similar disjunction with its nearest relatives.
Anemia oblongifolia (Cav.) Sw. Syn. Fil. 156. 1806. Osmunda ob-
longifolia Cav. Icon. Descr. PI. 6: 69. 1801. O. humilis Cav. I.e. Ane-
mia humilis (Cav.) Sw. Syn. Fil. 156. 1806. A. pilosa Mart. & Gal.
Nouv. Mem. Acad. Roy. Sci. Bruxelles 15 (5): 19. 1842. A. see-
mannii Hook. London J. Bot. 7: 564. 1848. A. presliana Prantl, Un-
ters. Morph. Gefass. Schiz. II: 104. 1881.
In forests, on rocks or rocky banks, 200-650 m.; Chiquimula;
Zacapa. Mexico to Panama; Venezuela to Brazil and Bolivia.
Rhizome oblique or ascending; clothed with orange or reddish, filiform trichomes;
leaves to 30 cm. long (including the erect, fertile pinnae) and 3.5 cm. broad, poly-
stichous, caespitose, partly dimorphous, the fertile pinnae and lowermost sterile
ones arising from the same position on the axis, sterile leaf similar in size and shape
to the fertile; petiole of fertile leaf 1-6 cm. long, shorter than the sterile leaves, stra-
mineous, or often light brown and strongly arcuate at base, rather stout, sparsely
villous, glabrate; sterile portion of lamina obovate-oblong or obovate, terminating
in a broadly obovate to flabelliform apical segment, pinnate, coriaceous or subcori-
aceous, slightly lustrous, especially above, pale- or orange-pilose on both sides,
sparsely so on rachis; sterile pinnae 2-9 pairs, spreading to slightly ascending, con-
tiguous or subimbricate, sessile, to 1.8 cm. long and 0.7 cm. broad, oblong, broadly
rounded at apex, strongly inequilateral at base, truncate or rounded above and
cuneate below, crenulate, margin somewhat thickened, a distinct midrib lacking;
veins free, rather evident, raised on adaxial side; fertile pinnae to 18 cm. long, pan-
icle much shorter than the stramineous stalk, mostly tripinnate, ultimate divisions
slender and only slightly foliaceous, becoming less so at maturity.
Anemia oblongifolia is one of the most distinctive of the Central
American species. The coarse, broadly rounded pinnae are often
nearly as broad as long; the slender, fertile pinnae extend far above
their own sterile blades; and the sterile leaves are so short and so
densely caespitose that they sometimes appear rosette-like.
Anemia pastinacaria Moritz, in Prantl Unters. Morph. Gefass.
Schiz. II: 110. 1881. A. longistipes (Liebm.) C. Chr. Index Fil. 53.
1905.
In rocky or grassy places, shaded or open slopes or in forests. 750-
1,650 m.; Alta Verapaz; Baja Verapaz; Chimaltenango; Huehue-
tenango; Izabal; Jalapa; Jutiapa; Peten; Santa Rosa; Zacapa.
Cuba; Trinidad; Mexico to Panama; Venezuela to Brazil and Bo-
livia.
Rhizome horizontal, short-creeping, densely clothed with orange, filiform tri-
chomes; leaves to 45 cm. long (including the erect, fertile pinnae) and 7 cm. broad,
polystichous, caespitose, partly dimorphous, the fertile pinnae and lowermost ster-
34 FIELDIANA: BOTANY, VOLUME 39
ile ones arising from same position on the axis, sterile leaf similar in shape to the
fertile, but with much shorter petiole; petiole of fertile leaf to 22 cm. long, usually
greatly exceeding the sterile leaf, commonly stramineous, often atropurpureus at
base, or brownish throughout, wiry to rather stout, glabrate; sterile portion of lam-
ina ovate-deltoid to ovate-oblong, gradually tapering to a pinnatifid apex, only
rarely with a distinct apical segment, pinnate, firm-membranaceous to subcoria-
ceous, dull to somewhat lustrous, pilose beneath, sparsely so or glabrate above,
rachis sparsely orange- villous ; sterile pinnae 5-10 pairs, spreading or slightly as-
cending, subsessile, to 3.5 (5) cm. long and 1.5 (2) cm. broad, obliquely oblong,
obtuse to acute, inequilateral at base, truncate or subtruncate above, cuneate below,
denticulate or crenate, or rarely with several deep lobes on lower pinnae, margin
slightly or not at all thickened, a midrib lacking or sometimes evident half the
length of the pinna; veins free, distinct and raised on adaxial side; fertile pinnae to
23 cm. long, panicle much shorter than (or rarely equalling) the brownish or stra-
mineous stalk, mostly tripinnate, ultimate divisions slender and scarcely ( or not at
all) foliaceous.
Some rather coarse specimens from Guatemala (Standley 80916),
Chimaltenango; Standley 82373, Huehuetenango) and several from
Honduras have been determined in herbaria as A. phyllitidis, due to
their apparent reticulate venation and their relatively large and
coarse pinnae (to 5 cm. long and 2 cm. broad). However, this anasto-
mosing of veins is infrequent and generally rather casual, and in no
other features do these specimens truly resemble -A. phyllitidis. The
pinnae are strongly inequilateral, with a midrib extending halfway
or less to a subacute apex, and some lower pinnae are deeply lobed.
The fertile panicles are much shorter than their stalks, and the ster-
ile blades gradually taper to a pinnatifid apex. The position of these
particular specimens appears to lie somewhere between A hirsuta
and A. pastinacaria. It is possible they are hybrids between one of
these and -A. phyllitidis. They are provisionally determined here as
A. pastinacaria. See further discussion of this species relationship
under A. hirsuta.
Anemia phyllitidis (L.) Sw. Syn. Fil. 155. 1806. Osmunda phyl-
litidis L. Sp. PL 1064. 1753.
Moist, often rocky, slopes and banks of ravines, in thickets and
edges of forests, 900-2,400 m.; Alta Verapaz; Chimaltenango; Chi-
quimula; El Progreso; Escuintla; Guatemala; Huehuetenango;
Jalapa; Santa Rosa; Solola; Zacapa. Greater Antilles (rare); Trini-
dad; Mexico to Panama; Colombia and Venezuela to Argentina.
Rhizome oblique to ascending, rather densely clothed with long (to 7 mm.), fili-
form, orange to reddish brown trichomes; leaves to 90 cm. long (including the erect,
fertile pinnae) and 25 cm. broad, polystichous, caespitose, partly dimorphous, the
fertile pinnae and lowermost sterile ones arising from the same position on the axis,
STOLZE: FERNS OF GUATEMALA 35
paripinnate or imparipinnate, the terminal segment (if present) subequal to next
pinna below, sterile leaf similar in shape to the fertile, but often shorter; petiole of
fertile leaf to 70 cm. long, frequently exceeding the sterile leaf (but more often not),
stramineous, or dark brown at base, stout, rather densely villous, often glabrate,
especially above; sterile portion of lamina ovate to deltoid, once-pinnate (or rarely
the lowermost pinnae lobed near the base), commonly subcoriaceous, dull to slightly
lustrous, rigidly and sparsely pilose, glabrate; sterile pinnae 3-7 pairs, ascending, or
the lowermost spreading, sessile or short-stalked, to 14 cm. long and 3 cm. broad,
ovate to lanceolate, often subfalcate, acute to acuminate, subequilateral at base and
rounded to truncate (or rarely some upper ones inequilateral), crenulate to dentate,
margin slightly cartilaginous, a distinct midrib extending nearly or fully to apex;
veins copiously anastomosing, rather obscure, or somewhat distinct and raised
above; fertile pinnae to 40 cm. long, panicle as long as the stramineous to reddish
brown stalk or (usually) longer, commonly tripinnate, ultimate divisions slender and
scarcely or not at all foliaceous.
Anemia speciosa Presl, Abh. Bohm. Ges. Wiss. V. 4: 349. 1845. A.
mexicana Kl. var. paucifolia Hook., 2nd Cent. Ferns, t. 65, 1861.
Ornithopteris speciosa (Presl) Reed, Bol. Soc. Brot. 21 (2): 153.
1947.
On bluffs and at forest edges, mostly on limestone, 0-1,350 m. ;
AltaVerapaz; Izabal; Peten. Cuba; Mexico; British Honduras.
Rhizome creeping, closely invested with brown to blackish, acicular trichomes;
leaves to 50 cm. long (including the erect, fertile pinnae) and 15 cm. broad, disti-
chous, crowded, partly dimorphous, the fertile pinnae borne 0.6-2.8 cm. below the
sterile portion of the lamina, sterile leaf similar in size and shape to the fertile; peti-
ole of fertile leaf to 30 cm. long, shorter than the sterile leaf, stramineous to light
brown, darker at base, wiry to rather stout, sparsely pilose, glabrate; sterile portion
of lamina deltoid, ovate or oblong-lanceolate, once-pinnate ( or rarely the lowermost
pinnae lobed at base), coriaceous to subcoriaceous, dull, or somewhat lustrous
above, glabrous; sterile pinnae 1-4 pairs, spreading or more often ascending,
stalked, to 10 cm. long and 3.5 cm. broad (the apical segment commonly larger),
ovate to oblong-lanceolate, subacute to acuminate, subequilateral at base and
rounded to subcordate, margin serrulate or serrate, cartilaginous, a distinct midrib
extending nearly or fully to apex; veins free, impressed above, somewhat raised and
rather distinct beneath; fertile pinnae to 12 cm. long, panicle longer than its
stramineous or light brown stalk, 2- to 3-pinnate, segments narrow-foliaceous and
indusiform.
Although superficially resembling Anemia phyllitidis in its undi-
vided pinnae and the apical segment of the lamina similar to the pin-
nae below, A. speciosa is easily distinguished from the former by its
free venation and its fertile pinnae borne beneath the sterile portion
of the lamina. A. phyllitidis has reticulate venation and the fertile
pinnae and lowermost sterile ones arising together on the axis.
36 FIELDIANA: BOTANY, VOLUME 39
LYGODIUM Swartz
References: A. H. G. Alston & R. E. Holttum, Notes on the tax-
onomy and nomenclature in the genus Lygodium (Schizaeaceae),
ReinwardtiaS: 11-22. 1959.
Vinelike plants with relatively short petioles but indefinitely long, flexuous,
scandent or trailing rachises; rhizome subterranean, branched and creeping, rather
densely clothed with stout, dark trichomes; leaves closely to rather widely spaced,
obscurely distichous, but ostensibly in a single row; rachis bearing alternate pinnae,
these with a minute stalk terminated by an arrested bud and flanked by a pair of
secondary pinnae; pinna stalk commonly a few millimeters long or appearing merely
as a rounded protuberance on the main rachis; secondary pinnae palmate, pinnately
branched, or with lateral pinnules lacking, the pinna stalk thus bearing a dichoto-
mously lobed or subpalmate lamina; veins commonly free (reticulate in L. hetero-
doxum); fertile pinnae similar in size and shape to the sterile, or (in a few species)
greatly constricted; sporangia apparently dorsal, in a single row on each side of the
midvein of contracted segments which are pinnately arranged along the margins of
the pinnules, the tissue serving as an indusium; spores trilete, commonly with
perine.
A pantropic genus, containing about 35 species. It is a natural
group, and quite distinctive due to its vinelike habit. The following
three species occur in Guatemala.
a. Veins reticulate L. heterodoxum.
a. Veins free.
b. Tertiary pinnules (at least the fertile ones) gradually diminishing in size toward
pinna apex, often hastate, not articulate, only slightly swollen (if at all) at
junction with their stalks L. venustum.
b. Tertiary pinnules subequal, not hastate, nodose-articulate at junction with
their stalks L. volubile.
Lygodium heterodoxum Kunze, Farrnkr. 2: 32. 1849.
Scandent or trailing on trees, shrubs and herbs, in forests and
damp thickets, 0-1,000 m.; Alta Verapaz; Huehuetenango; Izabal;
Peten. Southern Mexico to Costa Rica; reported from Venezuela,
but no specimens seen.
Rhizome short-creeping, clothed with stout, pluricellular, lustrous, blackish tri-
chomes; leaves several meters long; petiole and rachis stout, to 3 mm. in diameter,
terete, glabrous (rarely puberulent), stramineous, or the petiole brownish at base;
pinna stalks scarcely evident, 1-3 mm. long, the terminal bud inconspicuous, mostly
covered with pale to whitish, few-celled trichomes; secondary pinnae to 40 cm. long
and 30 cm. broad, pinnately branched, or often dichotomously or palmately cleft,
the pinnules or segments subequal in size and shape; tertiary pinnules 1-4 pairs, to
25 cm. long, simple and lanceolate to oblong ( rarely subhastate) or dichotomously to
subpalmately cleft into elongate segments, their stalks mostly narrow-alate, mar-
gins broadly and obscurely serrulate; costules continuous, not nodose-articulate at
FIG. 8. Lygodium. a-b, L. volubile: a, habit, X '/i; b, base of tertiary pinnule, X 2;
c-d, L. heterodoxum: c, habit, X '/i; d, base of tertiary pinnule, X 2'/2; e-f, L. uenus-
tum : e, habit, X Va ; f , base of tertiary pinnule, X 2.
37
38 FIELDIANA: BOTANY, VOLUME 39
segment base; leaf tissue firm-membranaceous, rather lustrous on both surfaces,
glabrous; midribs and veins glabrous or nearly so; veins oblique, raised, reticulate,
forming 3-5 ranks of areoles between midrib and margin.
Known in southern Mexico as "bejuco de alambre," "palmillo,"
and "yerDa de culebra." The petiole and lower parts of the rachis are
used in basket making in the Usumacinta River region of Huehuete-
nango, and are considered there to be better than "mimbre," the
vining stem ofMonstera or Philodendron sold in the local markets.
Lygodium venustum Sw. J. Bot. (Schrader) 1801 (2): 303. 1803;
Alston & Holttum, Reinwardtia 5: 15. 1959. L. mexicanum Presl,
Rel. Haenk. 1: 72. 1825. L. commutatum Presl, Abh. Bohm. Ges.
Wiss V. 4: 370. 1845 (type from Hacienda de Santa Luca, Guate-
mala?, Friedrichsthal).Alambrillo (Retalhuleu).
Scandent or trailing, in forests and thickets, often on rocky bluffs,
0-1,000 m.; Alta Verapaz; Baja Verapaz; Chiquimula; Escuintla;
Izabal; Jutiapa; Peten; El Progreso; Retalhuleu; San Marcos;
Santa Rosa; Zacapa. West Indies; Mexico and Central America
southward to Brazil and Bolivia.
Rhizome creeping; rhizome and petiole base densely clothed with stout, pluricellu-
lar, lustrous, blackish trichomes; leaves to several meters long; petiole and rachis
stout, terete, to 2.5 mm. in diameter, petiole to 30 cm. long, glabrous or nearly so,
light- or gray-brown, darker at base, rachis stramineous or light brown, sparsely to
densely puberulent; pinna stalks 5-10 mm. long, terminal bud inconspicuous to
scarcely evident, covered with pale to rusty, few-celled trichomes; secondary pinnae
pinnate to bipinnate, to 25 cm. long and 20 cm. broad, the pinnules gradually dimin-
ishing in size toward pinna apex; tertiary pinnules (3) 5-8 pairs, to 8 (10) cm. long,
lanceolate with a broadly cuneate base or elongate deltoid-ovate with a cordate base,
bearing 1-3 distinct pairs of subsessile, auriculiform, deltoid-ovate to flabellate seg-
ments, or (more commonly) with a pair of enlarged acute to obtuse basal lobes, thus
appearing subhastate or halberd-shaped; midrib of tertiary pinnules pilose (usually
densely so), not articulate, though sometimes slightly swollen at the junction with
the stalk, narrow-alate, the wings often continuing down along the axis of the next
order; leaf tissue firm-membranaceous or papyraceous, slightly lustrous, especially
above, glabrous; midribs and veins sparsely to densely pilose; veins oblique, raised,
free, once to several times forked.
Called "yerba buena" in Southern Mexico and "crespillo" in
southern Mexico, El Salvador, and Nicaragua.
This species has been often confused with, and frequently deter-
mined, Lygodium polymorphism HBK. However, the latter name is
a synonym of the Asiatic L. flexuosum (L.) Sw., a species with
which L. venustum is closely allied, but from which it is considered
distinct.
STOLZE: FERNS OF GUATEMALA 39
Lygodium mexicanum has been treated as a distinct species, on
the basis that the tertiary segments are continuous with their
stalks, the basal veins are alternate, the pubescence is less dense,
and the tissue more lustrous. It has been claimed that the segments
of L. venustum are nodose-articulate on their stalks and that the
basal veins are opposite or subopposite. However, these characters
are not consistent. Degree of luster and amount of pubescence
varies. The basal veins of segments can be predominantly opposite
in some specimens, alternate in others, or both conditions may
occur on adjacent segments on the same specimen. Furthermore,
the tertiary segments are never truly articulate on their stalks.
Occasionally there may be a slight swelling at the point from which
the basal veins arise, but this condition varies widely on individual
specimens.
Lygodium volubile Sw. J. Bot. (Schrader) 1801 (2): 304. 1803.
Scandent, in forests, 0-200 m. ; Izabal. British Honduras; Jamai-
ca; Cuba; Trinidad; South America.
Rhizome short-creeping, covered with stout, pluricellular, lustrous, deep brown to
blackish trichomes; leaves to several meters long; petiole and rachis stout, terete, to
2.5 mm. in diameter, glabrous or slightly puberulent, stramineous to light brown or
grayish, darker at base; pinna stalks 1-2 mm. long, terminal bud scarcely evident,
covered with pale to rusty, few-celled trichomes; secondary pinnae pinnate, to 20
cm. long and often as broad, the pinnules subequal in size and shape; tertiary pin-
nules 2-5 pairs, to 15 cm. long and 2.5 cm. broad, oblong-lanceolate, acute or acumi-
nate at apex, base broadly cuneate, truncate or subcordate (or very rarely auricu-
late-hastate), nodose-articulate at junction with the stalk, margins crenulate-ser-
rate; stalks of tertiary pinnules narrow-alate, the wings often continuing down
along the axis of the next order; leaf tissue firm-membranaceous, slightly lustrous,
glabrous to short -pilose; midrib and veins glabrous to pilose; veins oblique, raised,
free, commonly once or twice forked.
EXCLUDED SPECIES
LYGODIUM RADIATUM Prantl, Unters. Morph. Gefass. Schiz. 2:
66. 1881. L. digitatum Eaton, Mem. Amer. Acad. Arts 2. 8: 217.
1860.
L. digitatum was reported by Hemsley (Biol. Cent. Am. Bot. 3:
694. 1886) to be in Guatemala, citing a Godman & Salvin collection
from Izabal. I have not seen this specimen, but it is likely a misiden-
tification, for L. digitatum ( = L. radiatum ) has not been reported
with certainty north of Costa Rica.
FlG. 9. Schizaea elegans. a, habit, X Vt; b, sporangiophore, X 4; c, sporangiophore
segment, abaxial side, X 16; d, sporangiophore segment, adaxial side, X 16.
40
STOLZE: FERNS OF GUATEMALA 41
SCHIZAEAJ.E. Smith
References: Olof H. Selling, Studies in the recent and fossil spe-
cies oiSchizaea . . . Acta Horti Gothob. 16: 1-112, 1944; David W.
Bierhorst, Leaf development mSchizaea andActinostachys, Amer.
J. Bot. 56: 860-870. 1969.
Terrestrial, mostly erect ferns; rhizome minute to stout, horizontal or ascending,
sometimes short-creeping, thickly invested with pale to brown, one- to several-celled
trichomes; leaves commonly distichous, closely spaced or densely caespitose, simple
and linear, grasslike, or once to several times dichotomous with ultimate divisions
scarcely foliaceous, or, in ours, flabelliform (at least in general outline) and foliace-
ous, with the divisions linear-oblong to obovate; petiole generally glabrous, much
longer than the lamina, commonly rounded to obtusely angled abaxially, sulcate
adaxially; sporangiophores in terminal clusters, pinnatifid, with sporangia borne in
2 crowded rows on the narrowly-foliaceous, indusiform segments; spores monolete.
About 30 species occurring generally in tropical or subtropical
regions, in both hemispheres, but most prevalent in South America
and the Pacific Islands. Only one species has been reported from
Guatemala.
Schizaea elegans ( Vahl) Sw. J. Bot. (Schrader) 1800 (2): 103. 1801.
Acrostichum elegans Vahl, Symb. Bot. 2: 104. 1791. Lophidium ele-
gans (Vahl) Presl, Suppl. Tent. Pterid. 77. 1845.
In dense, wet forests, on wooded slopes and ridges, 0-500 m.; Alta
Verapaz; Huehuetenango; Izabal; Peten. Southern Mexico to
Panama; Colombia to Trinidad and southward to Brazil and
Bolivia.
Rhizome horizontal or ascending, densely covered with pale, stramineous or light
brown, 1- to several-celled trichomes which often extend onto the lower petiole;
leaves erect, caespitose, 20-70 cm. long; petiole stout, to 50 cm. long and 2.5 cm.
thick, rounded to obtusely angled on abaxial side, rather broadly sulcate on adaxial
side, mostly glabrous, stramineous, darker brown toward the base; lamina to 20 cm.
long, subcoriaceous, glabrous, slightly lustrous, flabelliform (at least in general out-
line), once to several times dichotomously forked or cleft, the divisions linear-
oblong, oblanceolate or obovate, with margins entire and plane or slightly revolute,
the apices sharply lacerate; costae once to several times forked within the ultimate
divisions; sporangiophores pinnately branched at the tips of the lacerations, with
sporangia borne in 2 crowded rows on the recurved, villous, narrowly foliaceous, in-
dusiform segments.
GLEICHENIACEAE
References: L. M. Underwood, A preliminary review of the North
American Gleicheniaceae, Bull. Torrey Bot. Club 34: 243-262. 1907;
42 FIELDIANA: BOTANY, VOLUME 39
W. R. Maxon, Gleicheniaceae, in North Amer. Fl. 16: 53-63. 1909;
T. Nakai, A new classification of Gleicheniales, Bull. Natl. Sci. Mus.
Tokyo 29: 1-71. 1950; R. E. Holttum, On the taxonomic subdivision
of Gleicheniaceae Reinwardtia4: 257-280. 1957.
Coarse, terrestrial plants, leaves scandent or trailing, often widely spreading and
forming dense thickets, frequently xerophilous; rhizome long-creeping on or just
beneath the surface of the ground, frequently branching, variously provided with
scales or trichomes; leaves commonly large (often to several meters long), circinate
in vernation, widely spaced to (rarely) sub fasciculate, petiole not articulate at the
rhizome, sterile and fertile leaves alike, once or several times pseudodichotomously
branched; petiole bearing a pair of opposite pinnae, with an arrested bud at its apex,
or continuing as the rachis which produces one or two more opposite pairs of pinnae;
pinnae bipinnate, or once to several times pseudodichotomously branched, with
each succeeding axis bearing an arrested apex; indument consisting of scales or of
simple, branched or stellate trichomes; veins free, simple or once to several times
forked; sori abaxial on the veins, circular in shape, exindusiate, each composed of 2
to many sporangia, situated on a barely elevated receptacle; sporangia sessile or
subsessile, pyriform, each with an essentially complete, transverse, usually medial
annulus, dehiscing longitudinally; spores 120 to ca. 800 in each sporangium, mono-
lete or trilete, generally smooth and (in ours) lacking significant ornamentation.
The pseudodichotomous branching of leaves and the usually pec-
tinate penultimate segments make this one of the most distinctive
of fern families. These frequently xerophilous plants are often
weedy, their large, spreading leaves forming dense tangles or thick-
ets along banks of ravines or over low shrubs. Approximately 120
species have been recognized by various authors in as many as eight
genera, these primarily based upon differences in type of indument,
shape of spores, and degree of branching. The following treatment is
based generally on the classification of Holttum ( 1957). Two genera
are recognized in Guatemala.
a. Leaves and rhizomes with trichomes only, scales lacking; veins with 2 or more
forks; sori commonly with 6 or more sporangia Dicranopteris.
a. Leaves and rhizomes with scales and trichomes; veins once-forked; sori with 2 to
5 sporangia Gleichenia.
DICRANOPTERIS Bernhardi
Reference: R. C. Ching, On the genus Gleichenia Smith, Sunyat-
senia 5: 269-289. 1940.
Rhizome provided with rigid, or somewhat lax, pluricellular trichomes; leaves
rather widely spaced; petiole at first erect, stout, rigid, of indeterminate growth,
bearing one to several pairs of opposite pinnae which again branch one or more
times in opposite pairs, or which branch alternately and quite unequally; only the
penultimate segments are pectinate, other axes are naked; a dense tuft of trichomes
and a pair of reduced, stipule-like appendages borne within each fork; penultimate
FIG. lO.Dicranopteris. a,D. flexuosa, habit, X Vi; b-c,D. pectinata: b, habit, X Vi;
c, base of fork, showing tuft of trichomes and pair of appendages borne within, X
3'/2.
43
44 FIELDIANA: BOTANY, VOLUME 39
segments borne in pectinate, subequal, usually ascending pairs; veins several times
forked; sori exindusiate, each composed of 6 to many sporangia; spores monolete or
(more commonly) trilete, smooth-surfaced, commonly lacking significant ornamen-
tation.
An essentially pantropical genus of eight or 10 species, growing
most frequently in dry, open locations, often forming dense mats or
thickets. It is readily distinguished, within the family, by its lack of
scales, the indument consisting solely of trichomes.
a. Penultimate (and other) axes branching in opposite pairs; accessory, mostly re-
duced, leafy segments usually borne in pairs at the base of each fork ( these in
addition to stipule-like segments borne within the forks) D. flexuosa.
a. Penultimate axes borne alternately along the axis of the next order below; acces-
sory, mostly reduced, leafy segments not borne in pairs at the base of each fork
(not to be confused with the stipule-like segments borne within the forks)
D. pectinata.
Dicranopteris flexuosa ( Schrad. ) Underw. Bull. Torrey Bot. Club
34: 254. 1907. Mertensia flexuosa Schrad. Gott. gel Anz. 863. 1824.
Gleichenia flexuosa ( Schrad. ) Mett. Ann. Lugd. Bat. 1: 50. 1863.
Leaves scandent or trailing, on dry, open slopes and ridges, or in
moist, open woods, 1,400-2,500 m. Alta Verapaz; Quezaltenango.
West Indies; Southern Mexico to Panama, southward to Brazil,
Bolivia and Paraguay. (Some isolated collections in the United
States from coastal Alabama. )
Rhizome protostelic, long-creeping, provided with rigid, reddish brown trichomes,
these falh'ng away cleanly, leaving the surface smooth or only slightly scabrous;
petiole terete, stramineous to light brown, smooth and rather lustrous, glabrous
except at the very base; pinnae one to several opposite pairs, these repeatedly pseu-
dodichotomous, with all subsequent branches in opposite pairs; a tuft of reddish
trichomes and a pair of reduced, subentire to pinnatisect, stipule-like appendages
borne within each fork; a pair of reduced, accessory, pectinate segments commonly
produced at the base of each fork, the other axes otherwise naked; penultimate seg-
ments sessile, branched in divergent to strongly ascending, subequal pairs, to 30
cm. long and 6 cm. broad, deeply pinnatisect, lanceolate to oblanceolate, coriaceous,
glabrous, glaucous beneath; ultimate segments mostly linear, retuse, slightly di-
lated toward the base, margins strongly revolute, often folded back and nearly
touching each other beneath the segment; veins 2- to 4-forked, approximate; sori
inframedial between midrib and margin; sporangia yellow- to red-brown, commonly
6-12 per sorus; spores trilete.
Dicranopteris pectinata (Willd. ) Underw. Bull. Torrey Bot. Club
34: 260. 1907. Mertensia pectinata Willd. Kongl. Vetensk. Acad.
Nya Handl. II. 25: 168. 1804. Gleichenia pectinata (Willd.) Presl,
Rel. Haenk. 1: 71. 1825. Gleichenella pectinata (Willd.) Ching, Sun-
yatsenia5: 276. 1940.
STOLZE: FERNS OF GUATEMALA 45
Leaves clambering, often forming dense thickets, on dry, open,
sometimes rocky slopes or moist, shaded river banks, and at edges
of forests, 0-2,300 m. Alta Verapaz; Huehuetenango; Izabal; Que-
zaltenango; Zacapa. West Indies; Mexico to Panama, Colombia to
Trinidad, southward to Bolivia and Brazil.
Rhizome solenostelic, long-creeping, strongly and conspicuously scabrous from
the persistent bases of slender, pluricellular reddish brown trichomes; petiole terete,
stramineous to light brown, smooth and glabrous except near the base; pinnae one
to several opposite pairs, each of these bearing several alternate branches which are
again alternately branched or which terminate in a pair of penultimate segments; a
tuft of reddish trichomes and a pair of reduced, entire to pinnatisect stipule-like
appendages borne within each fork, other axes completely naked and without re-
duced, accessory segments at the base of each fork; penultimate segments sessile,
branched in divergent to strongly ascending, subequal pairs, to 25 cm. long and 5.5
cm. broad, deeply pinnatisect, lanceolate to oblanceolate, chartaceous to coriaceous,
glabrous, glaucous beneath; ultimate segments mostly linear, retuse, slightly di-
lated toward the base, margins revolute; veins 2- to 4-forked, approximate; sori
medial or, more commonly, inframedial between midrib and margin; mature sporan-
gia usually yellowish brown, about 8-20 per sorus; spores monolete.
GLEICHENIA Smith
Reference: R. C. Ching, On the genus Gleichenia Smith, Sunyat-
senia 5: 269-289. 1940.
Rhizome provided with narrow, setose or short-ciliate scales; leaves widely or
(rarely) closely spaced, 2 m. or more in length (in ours); petiole suberect to ascend-
ing, stout, rigid, terete or somewhat flattened on the adaxial side near the basal pin-
nae, bearing a pair of opposite, ascending pinnae, with an arrested bud at its apex,
or continued as the rachis which subsequently produces one or two more opposite
pairs of pinnae; pinnae bipinnate, or (more typically) once to several times pseudo-
dichotomous with the penultimate segments in diverging, subequal, pectinate (in
ours) pairs; a dense tuft of pale to dark, lanceolate to ovate, usually acuminate
scales within each fork, and often flanked by one or two pairs of reduced, stipule-like
appendages; ultimate segments rigidly chartaceous to subcoriaceous, commonly
glabrous above, variously pubescent or tomentose and often whitish or yellowish
granulose beneath; veins simple or (in ours) once-forked; sori exindusiate, each with
2 to 5 sporangia; spores trilete or monolete, commonly (in ours) lacking ornamen-
tation.
More than 100 species are included in this genus, with distribu-
tion in tropical to south temperate areas of both hemispheres.
a. Pinnae bipinnate, not forked; axillary scales entire G. bancroftii.
a. Pinnae once to several times pseudodichotomous, the penultimate segments pec-
tinate-pinnatisect; axillary scales cilia te or setose.
b. Sori mostly inframedial, crowding or touching the midrib; segments densely
and evenly tomentose beneath.
46 FIELDIANA: BOTANY, VOLUME 39
c. Costal scales on abaxial side copious, orange, long-ciliate, 1-2 mm. long, axil-
lary scales orange to reddish brown, lax, mostly long-ciliate G. bifida.
c. Costal scales on abaxial side essentially lacking, or (more often) scattered,
dark brown to blackish or dark-centered, often setose, less than 1 mm. long;
axillary scales dark brown to blackish, rigid, mostly short -ciliate or se-
tose G. brevipubis.
b. Sori mostly medial to supramedial, rarely crowding the midrib (except on
strongly revolute segments); segments not tomentose, or rarely (in G. under-
woodiana ) with tomentum confined to the area along the midrib.
d. Midribs of ultimate segments commonly provided beneath (at least near
the base) with orange to reddish brown, ciliate sometimes hairlike, scales,
and abundant, orange to reddish brown, stellate trichomes
G. underwoodiana.
d. Midribs of ultimate segments lacking scales (or rarely with some scattered
whitish or yellow hairlike ones), but sparsely provided beneath with whit-
ish or yellow stellate trichomes G. palmata.
Gleichenia bancroftii Hook. Sp. Fil. 1: 5. 1844. Mertensia bancrof-
tii Kunze, Linnaea 18: 307. 1844. Dicranopteris bancroftii (Hook.)
Underw. Bull. Torrey Bot. Club 34: 252. 1907. Hicriopteris bancrof-
tii (Hook.) Ching, Sunyatsenia 5 : 278. 1940.
Commonly on slopes and sides of ravines in open forests, with
leaves clambering over shrubs and trees, 1,650-3,200 m. Baja Vera-
paz; Huehuetenango; San Marcos; Zacapa. Southern Mexico; Hon-
duras; El Salvador; Costa Rica; West Indies; Colombia; Venezuela;
Ecuador; Peru; Bolivia.
Rhizome stout, to 6 mm. in diameter, smooth or slightly muricate, long-creeping
on or just beneath the surface of the ground, provided with appressed, yellowish
brown or grayish brown scales; leaves widely spaced; petiole stout, mostly smooth
and lustrous, terete, or somewhat flattened on the adaxial side near the basal pin-
nae, stramineous to light brown and glabrous above, often darker and scaly at the
base; pinnae bipinnate; bud of primary axil thickly beset with entire, whitish or yel-
lowish, lanceolate to ovate, acuminate scales; pinnae 0.5-1.5 m. long, to 40 cm.
broad, with costa glabrous, or with a few narrow scales at bases of penultimate seg-
ments, terete beneath, slightly bicarinate above; pinnules numerous, crowded,
spreading at nearly right angles from the costa, sessile, linear, pectinate nearly or
quite to the costule; ultimate segments linear, chartaceous or subcoriaceous, acute
strongly revolute, slightly dilated at base, often with scales scattered along midrib;
veins once (rarely twice) forked; sori inframedial between midrib and margin, with 2
to 5 sporangia, subtended by a cluster of yellowish trichomes; spores trilete.
Within the family this is the only species in the neotropics having
the pinnae bipinnate and unforked. It should be confused with no
other species in Guatemala.
Gleichenia bifida (Willd.) Sprengel, Syst. Veg. ed. 16, 4: 27. 1827.
Mertensia bifida Willd. Kongl. Vetensk. Acad. Nya Handl. II, 25:
FIG. 11. Gleichenia. a-b, G. palmata: a, habit, X Vi; b, bud of primary axil, showing
scales and stipule-like appendages, X I'/i; c, G. bancroftii, habit, X Vi; d, G. bifida,
ultimate segments, showing sori crowding midrib, X 3; e, G. underwoodiana, ulti-
mate segments, showing tomentum along midrib, and sori distant from midrib, X 3.
47
48 FIELDIANA: BOTANY, VOLUME 39
168. 1804. M. fulva Desv. Mem. Soc. Linn. Paris 6: 200. 1827. Di-
cranopteris fulva (Desv.) Underw. Bull. Torrey Bot. Club 34: 255.
1907. D. bifida (Wllld.) Maxon, North Amer. Fl. 16: 60. 1909. Sti-
cherus bifidus (Willd.) Ching, Sunyatsenia 5: 282. 1940. Chispa
(Baja Verapaz).
In forests and damp thickets or open or brushy slopes, 400-2,300
m. Alta Verapaz; Baja Verapaz; Huehuetenango; Izabal; Quezal-
tenango; San Marcos. West Indies; Mexico to Panama; Colombia
and Venezuela southward to Bolivia and Brazil.
Rhizome long-creeping, often branched, dark brown, provided with reddish brown,
setose or short-cilia te, linear, acuminate scales; leaves widely spaced; petiole stout,
suberect, smooth, dull or somewhat lustrous, terete, yellowish brown to greenish
brown (rarely reddish), mostly glabrous, except sometimes scaly at apex and base;
bud of primary axil thickly covered with orange to reddish, long-ciliate, ovate to
lanceolate, acuminate scales, and often flanked by a pair of reduced, stipule-like
appendages; pinnae once- or twice-pseudodichotomous, with an arrested bud (or
rarely a secondary axis) produced at the apex, scaly as in the primary axil; scales on
the axes orange, long-ciliate, often dense; secondary axes 2-6 cm. long, not pectinate
or only partly so on the acroscopic side; tertiary axes 3-10 cm. long, pectinate, or
partly so on the acroscopic side; penultimate segments (20) 30-60 cm. long, 3-7 cm.
broad, pectinate, cut nearly to the costa, lanceolate, mostly subcoriaceous ; costa
above glabrous or provided with orange or whitish, mostly stellate trichomes, be-
neath with abundant, conspicuous (1-2 mm. long) spreading, orange, ciliate scales;
ultimate segments linear to linear-lanceolate, obtuse to acute, slightly or strongly
revolute, densely and evenly tomentose with orange or yellowish, stellate trichomes
and a few hairlike scales, sometimes whitish or partly deciduous in age; veins 15-40
pairs, once-forked; sori mostly inframedial, crowding or touching the midrib, with 3
to 4 sporangia; spores monolete, bean-shaped.
Gleichenia bifida can usually be easily distinguished from other
species (except G. brevipubis) in our area by the dense tomentum on
the abaxial side of the segments, in which the sori are often quite
deeply immersed. In some specimens of G. underwoodiana, pinna
segments may be quite densely villous along the midrib, but are
much less so toward the margin. Pinna segments of G. bifida ( ex-
cept those glabrescent with age) are densely and evenly tomentose
throughout.
The common name "chispa" apparently refers to the combustible
nature of the spores, which produce a sparkling display when poured
over an open flame.
Gleichenia brevipubis Christ, Bull. Herb. Boissier II. 6: 280. 1906.
Palma cateje; venado del rio (fide Steyermark, Quezaltenango).
STOLZE: FERNS OF GUATEMALA 49
In thickets and on moist, open banks and ravines, 1,200-2,700 m.
Alta Verapaz; El Progreso; Quezaltenango; San Marcos; Suchite-
pequez. Jamaica; Mexico to Costa Rica.
Rhizome branched, long-creeping, dark brown, provided with dark brown, setose
or short-cilia te, linear, acuminate scales; leaves closely to widely spaced; petiole
stout, suberect, smooth, dull or somewhat lustrous, terete, stramineous, or greenish
to reddish brown, glabrous, or with a few scattered scales at base; bud of primary
axil thickly covered with blackish, dark brown or dark-centered, rigid, setose or
short-ciliate, linear-lanceolate to lance-ovate scales (or rarely some with longer,
white cilia) and often flanked by a pair of reduced, stipule-like appendages; pinna
once- or twice-pseudodichotomous, with an arrested bud (or rarely a secondary axis)
produced at the apex, these scaly as in the primary axil; scales on the axes lacking,
or scattered, minute, reddish brown or blackish; secondary and tertiary axes 4-12
cm. long, wholly pectinate, or partly so on the acroscopic side; penultimate seg-
ments (20) 24-40 cm. long, pectinate, cut nearly to the costa, lanceolate, mostly sub-
coriaceous; costa above glabrous, or provided with scattered, blackish scales or
whitish, stellate trichomes, beneath with scattered, minute (less than 1 mm.), often
appressed, dark brown or dark-centered, commonly setose, scales and usually with a
few inconspicuous, orange, often hairlike ones; ultimate segments linear to linear-
lanceolate, obtuse to acute, slightly or strongly revolute, densely and evenly tomen-
tose with whitish or yellowish, stellate trichomes and a few hairlike scales; veins
18-30 pairs, once-forked; sori mostly inframedial, crowding or touching the midrib,
with 3 to 4 sporangia; spores monolete, bean-shaped.
Maxon considered this conspecific with G. bifida, and perhaps it
merits only varietal status. However, the two probably differ from
each other to the same extent that G. palmata differs from G. under-
ivoodiana, especially as to the scales of axes, axils, and costae.
Throughout the range, G. brevipubis varies somewhat in scale char-
acters. In southern Central America the black, setose costal scales
are usually quite numerous, whereas farther north these scales are
often widely scattered to nearly lacking. In addition, the secondary
axes of G. brevipubis are often pectinate, on both sides of the axis,
whereas those of G. bifida are never wholly pectinate. But this is not
always a reliable character, for on some specimens the secondary
axes of both species can be pectinate only on the acroscopic side.
Furthermore, the secondary axes are not always available for exam-
ination, having been omitted by collectors, due to lack of space on
the herbarium sheet. The entire species complex needs a thorough,
updated study to clarify specific and infraspecific relationships.
Gleichenia palmata (Underw.) C. Chr. Index Fil. Suppl. 1: 113.
1913. Mertensia palamata Schaffner in Fee, Mem. Fam. Foug. 9: 40
(seors. 32). 1857. nom. nud. Gleichenia palmata (Schaffner) Moore,
Index Fil. 380. 1862. nom. nud. Dicranopteris palmata Underwood,
50 FIELDIANA: BOTANY, VOLUME 39
Bull. Torrey Bot. Club. 34: 259.1907 (type from Orizaba, Veracruz,
Mexico, Pringle 6129). Sticherus palmata (Underw. ) Copel. Gen.
Fil. 28. 1947.
On mountain slopes and sides of ravines in forest, 1,100-2,600 m.
AltaVerapaz; BajaVerapaz; Chiquimula; Guatemala (?); San Mar-
cos; Zacapa. Cuba; Jamaica; Southern Mexico; Honduras.
Rhizome branched, long-creeping near surface of ground, dark brown to atropur-
pureus, provided with reddish brown, setose or short-ciliate scales; leaves widely
spaced; petiole stout, suberect, smooth but not lustrous, terete, or somewhat flat-
tened on the adaxial side near the basal pinnae, yellowish to greenish and glabrous
above, dark brown, lightly tuberculate and scaly at the base; bud of primary axil
thickly covered with ciliate, reddish brown, lanceolate to ovate, acuminate scales,
and flanked by 1 or 2 pair of reduced, stipule-like appendages; pinnae once to several
times pseudodichotomous, with an arrested bud (or rarely with a secondary axis)
produced at the apex, these scaly and appendaged as in the primary axil; scales on
the axes reddish brown to whitish, long-ciliate, usually confined to the acroscopic
side; secondary axes 4-8 cm. long, not pectinate; tertiary axes 3-4 cm. long, not pec-
tinate, or rarely partly so on the acroscopic side; quaternary axes commonly fully
pectinate as in the penultimate segments; penultimate segments 12-25 cm. long, 2-4
cm. broad, pectinate, cut nearly or fully to the costa, lanceolate, chartaceous to sub-
coriaceous ; costa above glabrescent or provided with scattered to abundant reddish
brown, orange or whitish, often hairlike scales, beneath with abundant, reddish
brown or (rarely) orange, ciliate scales; ultimate segments linear, obtuse to sub-
acute, slightly to strongly revolute, faintly white- or yellow-granulose beneath, the
midribs, veins and tissue beneath lacking scales, but lightly pubescent with white or
yellow, stellate, scattered trichomes (rarely a few pale, delicate, hairlike scales pres-
ent on midrib); veins 10-30 pairs, once-forked; sori medial to supramedial between
midrib and margin, with 3 to 5 sporangia; spores monolete, bean-shaped.
Specimens of Gleichenia palmata can be confused with those of G.
underwoodiana that are not densely pubescent. Segment midribs of
the latter are typically provided with abundant, reddish, stellate tri-
chomes and (at least in the proximal portion) rather conspicuous
reddish brown, ciliate scales; whereas the midribs in G. palmata are
provided merely with a light scattering of white or pale yellow tri-
chomes. But even when pubescence is light in G. underwoodiana,
and scales may be nearly lacking on the midrib, the indument is
decidedly darker in color than the pale trichomes of G. palmata. In
addition, the tertiary axes in G. underwoodiana are typically pec-
tinate, at least on the acroscopic side. In G. palmata, the tertiary
axes are not pectinate or, rarely, have reduced segments on the acro-
scopic side.
Gleichenia underwoodiana (Maxon) C. Chr. Index Fil. Suppl. 1:
44. 1913. Dicranopteris underwoodiana Maxon, North Amer. Fl. 16:
STOLZE: FERNS OF GUATEMALA 51
59. 1909 (type from Chiapas, Ghiesbreght 271). Sticherus under-
woodianus (Maxon) Nakai, Bull. Natl. Sci. Mus. Tokyo 29: 31. 1950.
On moist, shaded, mountain slopes and stream banks, 2,000-3,200
m. Chimaltenango; Huehuetenango; El Quiche; San Marcos;
Zacapa. Mexico; Honduras; El Salvador; Costa Rica.
Rhizome branched, long-creeping, dark brown to atropurpureus, provided with
reddish brown, setose to short-cilia te scales; leaves widely spaced; petiole stout,
suberect, smooth, somewhat lustrous, terete, yellowish brown to greenish brown or
atropurpureus, glabrous, or sometimes scaly at apex and base; bud of primary axil
thickly covered with ciliate, reddish brown, lanceolate to ovate, acuminate scales
and flanked by 1 or 2 pair of reduced, stipule-like appendages; pinna once to several
times pseudodichotomous, with an arrested bud (or rarely a secondary axis) pro-
duced at the apex, these scaly and often appendaged as in the primary axil; scales
on the axes reddish brown to whitish, long-ciliate, usually confined to the acroscopic
side; secondary axes 3-12 cm. long, not pectinate, or (rarely) only partly so on acro-
scopic side; tertiary axes 4-8 cm. long, pectinate, or partly so on acroscopic side;
quaternary axes fully pectinate as in the penultimate segments; penultimate seg-
ments 10-28 cm. long, 1.5-3.2 cm. broad, pectinate, cut nearly or fully to the costa,
lanceolate, chartaceous to subcoriaceous; costa above glabrescent or provided with
scattered to abundant reddish brown, orange or whitish, often hairlike scales,
beneath with abundant, reddish brown or (rarely) orange, ciliate scales; ultimate
segments linear, obtuse to acute, slightly to strongly revolute, white- or yellow-
granulose beneath, the midribs often densely tomentose and scaly, provided (at
least in the proximal third) with orange to reddish brown scales and stellate tri-
chomes, the veins with orange to reddish brown trichomes, the tissue glabrous;
veins 16-22 pairs, once-forked; sori medial to supramedial between midrib and mar-
gin, with 3 to 5 sporangia; spores monolete, bean-shaped.
This species has been combined with G. compacta Christ in some
herbaria. However, a type fragment of the latter at U.S. National
Herbarium has obviously inframedial sori, and blackish and dark
setose costal scales, which indicates closer affinity with G. brevi-
pubis than with G. underwoodiana.
HYMENOPHYLLACEAE
References: C. B. Presl, Hymenophyllaceae, Abh. Bohm. Ges.
Wiss. V. 3: 93-162. 1843. R. van den Bosch, Synopsis Hymenophyl-
lacearum, Ned. Kruidk. Arch. 4: 341-419. 1859, and Supplement op.
cit. 5: 135-185. 1861. K. Prantl, Die Hymenophyllaceen, Untersuch.
Morph. Gefasskrypt. I: 3-14. 1875. E. B. Copeland, Genera Hymen-
ophyllacearum, Philipp. J. Sci. 67: 1-110. 1938. C. V. Morton, The
genera, subgenera and sections of the Hymenophyllaceae, Contr.
U.S. Natl. Herb. 38: 153-214. 1968.
52 FIELDIANA: BOTANY, VOLUME 39
Medium-sized, small or minute, usually delicate, epiphytic plants, or a few terres-
trial; rhizome thin and long-creeping with leaves widely spaced, or (less commonly)
stout, ascending to erect with leaves fasciculate, provided with scattered, simple,
light brown, reddish or blackish trichomes; leaves commonly monomorphous, or
rarely dimorphous in Trichomanes, typically circinate in vernation, not articulate at
the rhizome; lamina flabellate, or pinnatifid to decompound, variously pubescent,
scales lacking (except some marginal scalelike processes present in T. membranace-
um), the tissue 1 cell thick, lacking stomata; venation anadromous, or often cata-
dromous in Trichomanes, free or in a few species reticulate; sori marginal, terminal
on the veins; indusium immersed to short-stalked, bivalvate deeply or fully to base,
or tubular to salverform with an entire or bilabiate mouth; receptacle short and
rudimentary or filiform to thickly cylindrical or narrow-turbinate, not or long-ex-
serted from the indusium at maturity; sporangia sessile, borne on all sides of the
receptacle, each with uninterrupted, oblique annulus, borne on all sides of the
receptacle, each with uninterrupted, oblique annulus, dehiscing more or less longi-
tudinally; spores tetrahedral to subglobose, trilete, white or, more commonly green-
ish, 32-420 (fide Bower) in each sporangium.
The "filmy ferns" constitute a natural and distinctive, predom-
inantly tropical, family containing 550-600 species. Two genera,
subequal in size, are recognized here, although 30 or more have been
proposed by earlier workers. Most species are all to minute, with
laminar tissue rarely more than one cell thick. Sori are always mar-
ginal, with the indusium tubular (cylindrical to salverform) in
Trichomanes or deeply bivalvate inHymenophyllum. They are typi-
cally epiphytes of dark, wet forests and are of no importance to
man, except in an aesthetic sense. With their delicate, filmy leaves
and usually pendent habit, they are among the loveliest and most
graceful of all ferns.
a. Indusium obviously bivalvate (often fully to base), never tubular, the valves con-
stituting the major part of the indusium; receptacle filiform or thickened, not or
slightly exserted; venation anadromous Hymenophyllum.
a. Indusium tubular, obconic or salverform, sometimes bilabiate, the tubular por-
tion constituting an equal or major part of the indusium; receptacle filiform,
somewhat to greatly exserted at maturity; venation anadromous or catadro-
mous Trichomanes.
HYMENOPHYLLUM J. E. Smith
Reference: C. V. Morton, The American species of Hymenophyl-
lum section Sphaerocionum , Contr. U.S. Natl. Herb. 29: 139-201.
1947.
Plants epiphytic or occasionally terrestrial; rhizome long-creeping, commonly
sparsely provided with brown or reddish trichomes; leaves monomorphous, simple
or (in ours) pinnatisect to decompound, determinate or indeterminate, commonly
2-60 cm. long, short- to long-petiolate; petiole delicate, wiry or stout, terete and
STOLZE: FERNS OF GUATEMALA 53
smooth, shorter than the lamina, nonalate or marginate, to broadly alate for most or
part of its length, glabrous, or provided with simple to stellate trichomes; lamina
subflabellate to ovate to linear, glabrous or variously provided with simple to stel-
late trichomes; venation free, subflabellate or (most commonly) pinnate, anadro-
mous, spurious veins absent; ultimate segments usually narrow, often filiform, the
margins entire or serrate; indusium partly or not at all immersed in the tissue, sub-
orbicular to elliptic or obovate, bivalvate deeply or fully to base, the base often conic
but never tubular; receptacle filiform to thickened, cylindrical to narrowly turbi-
nate, rudimentary or short or somewhat exserted at maturity ; spores tetrahedral to
subglobose, trilete, smooth to variously sculptured, white or, more commonly,
greenish.
Approximately 250-300 species of Hymenophyllum are found
throughout the world. Although essentially a tropical genus, some
of its species occur in subtropical latitudes, and a few even in wet
temperate areas. They are mostly epiphytic plants, but have been
occasionally reported from wet, rocky banks in dense forests. The
genus (especially the American species) is much in need of good,
monographic work. Morton's study of section Sphaerocionium was
a beginning, but confusion still exists among species of subgenera
Mecodium and Hymenophyllum. Consequently, a few of the species
concepts herein are provisionally treated.
a. Ultimate segments sharply and deeply serrate throughout, the serrations com-
monly remote (Subgenus: HYMENOPHYLLUM).
b. Indusium entire or serrulate near apex; sori 1-2 at acroscopic base of pinna
(rarely also with 1 on basiscopic side); receptacle often much-exserted at
maturity; elev. 1,250-3,150 m H. fucoides.
b. Indusium laciniate to subciliate in apical half; sori 1-5 along acroscopic side of
pinna, and often 1-2 (3) on basiscopic side; receptacle slightly or not at all
exserted; elev. 300-1,200 m H. ectocarpon.
a. Ultimate segments entire (sometimes undulate to strongly crisped, but never
serrate).
c. Lamina glabrous (Subgenus: MECODIUM).
d. Lamina essentially pinnate or pinnatifid, the primary segments simple to
bifid (or proximal ones sometimes subflabellate or pinnatifid); segments
plane to slightly undulate,
e. Lamina (of mature leaves) 4-14 cm. long; ultimate segments mostly 2-3
mm. broad H. asplenioides.
e. Lamina rarely over 2.5 cm. long; ultimate segments mostly 1-1.5 mm.
broad.
f. Pinnae 3-7 pairs; lamina (of mature leaves) 1.5-2.5 cm. long, ovate to el-
liptic, the basal pinnae either reduced or strongly ascending, thus the
lamina appearing tapered to base H. abruptum.
f. Pinnae 1-2 (3) pairs; lamina 0.5-1.2 cm. long, subflabellate, basal ones
spreading, thus the lamina truncate at base H. brevifrons.
d. Lamina bipinnatifid to tripinnatifid; ultimate segments plane, undulate, or
strongly crispate.
54 FIELDIANA: BOTANY, VOLUME 39
g. Ultimate segments conspicuously crispate, often so strongly as to appear
dentate or deeply lobed H. contortum.
g. Ultimate segments plane, undulate or slightly crispate.
h. Petiole .01-.02 mm. thick, neither alate nor marginate; lower half of rach-
is nonalate or essentially so; lower 2-4 pinnae short-stalked
H. undulatum.
h. Petiole (of mature leaves) 0.3-0.7 mm. thick, often marginate or alate;
rachis alate throughout; pinnae adnate or basal one short-stalked.
i. Lamina not or scarcely narrowed at base; the lowermost pair of pinnae
somewhat or not at all reduced; lamina 1.5-2.5 times longer than broad,
and 1.5-3.5 times longer than the petiole; fertile segments not usually
constricted at apex, the indusium slightly immersed in the tissue
H. polyanthos.
i. Lamina gradually narrowed at base, with the lower 3-5 pairs of pinnae
somewhat to greatly reduced; lamina 3-8 times longer than broad, and
4-7 times longer than the petiole; fertile segments often greatly con-
stricted, thus the indusium appearing stalked H. myriocarpum.
c. Lamina provided with trichomes, at least on margins or veins (Subgenus:
SPHAEROCIONIUM) .
j. Trichomes borne on abaxial and adaxial leaf surfaces as well as on veins
and/or margins (Subsection: HIRSUTA).
k. Leaves 2-6 cm. long and 1-1.5 cm. broad, determinate; petiole delicate, 0.1-
0.2 mm. thick; pinnae simple or bifid, or rarely some basal ones subflabel-
late.
1. Stellate trichomes on segment margins sessile or subsessile; petiole tri-
chomes mostly simple or bifurcate; pinnae mostly bifid H. fragile.
1. Stellate trichomes on segment margins obviously stalked; petiole tri-
chomes mostly stellate; pinnae mostly simple H. lanatum.
k. Leaves (mature) 8-60 cm. long and 2.5-8 cm. broad, indeterminate; petioles
wiry or stout, 0.3-0.9 mm. thick (except 0.2-0.25 mm. in H. elegantulum);
pinnae 1- or 2-pinnatifid.
m. Petiole wiry, 0.2-0.25 mm. thick, sparsely provided with simple (rarely a
few bifurcate) trichomes H. elegantulum.
m. Petiole stout, 0.3-0.9 mm. thick, sparsely to abundantly provided with
mixed, mostly stellate, trichomes.
n. Trichomes of segment margins stellate, obviously stalked, delicate, the
rays spreading; petiole 0.3-0.5 mm. thick, alate at least near apex;
pinnae essentially pinnatifid, the secondary segments mostly simple to
bifid H. sieberi.
n. Trichomes of segment margins stellate, sessile or subsessile, stout, the
rays commonly appressed and directed toward segment tip; petiole
(0.5) 0.6-0.9 mm. thick, nonalate; pinnae essentially bipinnatifid, the
larger secondary segments deeply pinnatifid H. crassipetiolatum.
j . Trichomes lacking between veins and margins ( Subsection: CILIATA ).
o. Petiole alate, at least near apex; rachis alate throughout,
p. Trichomes on segment margins mostly simple, or once-forked at base,
rather widely scattered; trichomes lacking (or rare) on veins adaxially.
q. Petiole delicate, ca. 0.2 mm. thick, rarely more than 3 cm. long; petiolar
trichomes mostly simple, scattered H. crispum.
STOLZE: FERNS OF GUATEMALA 55
q. Petiole stout, commonly 0.5-0.8 mm. thick and 3-9 cm. long; petiolar
trichomes (when present) mostly stellate H. microcarpum.
p. Trichomes on segment margins mostly bifurcate or stellate, usually
abundant; trichomes present (sometimes sparse) on the veins of both
sides.
r. Petiole conspicuously alate one-half to three-quarters to base, with tri-
chomes mostly stellate; larger pinnae with 2-4 pairs of seg-
mnts H. hirsutum.
r. Petiole alate only at apex (or rarely one- third to base), with trichomes
mostly simple or bifurcate; larger pinnae commonly with 4-7 pairs of
segments H. maxonii.
o. Petiole nonalate throughout; rachis nonalate at base, or nearly throughout.
s. Trichomes on veins and margins stellate, usually abundant; rachis non-
alate nearly throughout; leaves indeterminate.
t. Pinnae commonly pinnate, the costae not continuously alate on both
sides; pinnae (some of them) elongate, often becoming frondlike and
greatly exceeding the adjacent ones H. tricophyllum.
t. Pinnae merely pinnatifid, thus the costae continuously alate on both
sides; pinnae (except reduced basal pairs) subequal to adjacent
ones H. pulchellum.
s. Trichomes on margins mostly simple or bifurcate, on veins rare or lack-
ing; rachis alate in the distal portion; leaves determinate,
u. Indusia (most of them) much broader than the ultimate segments;
rachis of mature leaves commonly nonalate in proximal half of
lamina H. lineare.
u. Indusia as broad or scarcely broader than the ultimate segments; rach-
is of mature leaves alate (or pinnae conspicuously decurrent) except
near base of lamina H. elegans.
Hymenophyllum abr upturn Hook., Sp. Fil. 1: 88, t. 31b. 1844.
Leptocionum abruptuhi Presl, Abh. Bohm. Ges. Wiss. V: 336. 1848.
Mecodium abruptum (Hook.) Copel. Philipp. J. Sci. 67: 26. 1938.
In deep shade, on trees or rotting logs in forests, 1,000-2,500 m.;
AltaVerapaz; Chiquimula; Zacapa. Greater Antilles.
Plants commonly epiphytic; rhizome thin, long-creeping, delicate, essentially
glabrous; leaves glabrous, determinate, subdistant on the rhizome, often forming
dense mats, mature ones 2-4 cm. long; petiole delicate, essentially glabrous, 0.5-2
cm. long, ca. 0.1 mm. in diameter, nonalate; lamina 1.5-2.5 (3.3) cm. long, pinnati-
sect, ovate, elliptic, or oblong, the basal pinnae strongly ascending (occasionally re-
duced) thus the lamina tapered to base; rachis narrow-alate (due to the decurrent
pinnae), or nonalate at base, with the basal pinnae subdistant; pinnae 3-7 pairs, sim-
ple or bifid, the ultimate segments 1.5-1.7 mm. broad, entire, plane or slightly undu-
late, the apices retuse; sori 1-4, obturbinate, terminating the apical segments;
indusia entire, as broad as or broader than the segments, deeply immersed, bivalv-
ate one-half to two-thirds to the cuneate base; receptacle narrow-cylindric, some-
what exserted at maturity.
This species and H. brevifrons are easily confused, as both are
minute, delicate, and roughly similar in leaf outline. However, the
56 FIELDIANA: BOTANY, VOLUME 39
basal pinnae of the latter commonly spread from the rachis at nearly
right angles, so that most laminae have a truncate appearance.
Basal pinnae of H. abruption are most often strongly ascending, or
they are sometimes greatly reduced; in either case the lamina ap-
pears to be tapered to base. Although the rachis is characteristically
alate in both species, the wing may be often lacking in H. abruptum
between the two lowest pinnae, and the basal pinna is occasionally
subpetiolulate.
Hymenophyllum asplenioides (Sw.) Sw. J. Bot. (Schrader)
1800(2): 98. 1801. Trichomanes asplenioides Sw. Prodr. 136. 1788.
Mecodium asplenioides (Sw.) Copel. Philipp. J. Sci. 67: 26. 1938.
Pendent from tree trunks and branches, in wet forests, 1,200-
2,000 m.; Alta Verapaz; Baja Verapaz; Chiquimula. Mexico; Hon-
duras; Nicaragua; Costa Rica; Panama; West Indies; Venezuela;
British Guiana; Brazil.
Plants epiphytic; rhizome thin, long-creeping, delicate, glabrous or with a few,
scattered light brown, simple trichomes; leaves glabrous, determinate, distant on
the rhizome, mature ones 5-15 cm. long; petiole glabrous, delicate, 1-5 cm. long, 0.2-
0.3 mm. in diameter, nonalate; lamina 4-14 cm. long, pinnatisect, oblong or linear,
not or slightly reduced at base, the rachis alate throughout, or occasionally nonalate
at the very base; pinnae 4-15 pairs, decurrent, simple to bifid, or lower ones sub-
flabellate or shallowly pinnatifid; ultimate segments mostly 2-3 mm. broad, entire,
plane or slightly undulate; sori numerous, terminating most of the ultimate segm-
ments; indusia entire, as broad or slightly broader than the ultimate segments,
scarcely or not at all immersed, bivalvate to the arcuate or subtruncate base; recep-
tacle narrow-cylindric, short, not exserted.
Hymenophyllum brevifrons Kunze, Bot. Zeit. 185. 1847. H. tabla-
ziense Christ, Bull. Soc. Bot. Geneve II. 1: 216. 1909. Mecodium
tablaziense (Christ) Copel. Philipp. J. Sci. 67: 26. 1938.
On tree trunks, or sometimes on clay or limestone banks, in for-
ests, near sea level; Izabal. British Honduras; Nicaragua; Costa
Rica; Panama; West Indies; the Guianas.
Plants usually epiphytic; rhizome thin, creeping, delicate, essentially glabrous;
leaves glabrous, determinate, subdistant on the rhizome, often forming dense mats,
mature ones 1-2 cm. long; petiole glabrous, delicate, 0.1-1 cm. long, 0.1-0.15 mm. in
diameter, not alate, or scarcely so at apex; lamina 0.5-1.2 cm. long, pinnatisect, sub-
flabellate, the basal pinnae spreading, thus the lamina truncate at the base; rachis
broadly alate throughout, with pinnae crowded or imbricate; pinnae 1-2 (3) pairs,
simple or bifid (or sometimes the basal ones again bifid), the ultimate segments
1-1.5 mm. broad, entire, plane or slightly undulate, the apices retuse; sori 1 or 2 (3),
obturbinate, terminating the apical segments; indusia entire, as broad or broader
than the segments, deeply immersed, bivalvate one-half to two-thirds to the cune-
ate base; receptacle narrow-cylindric, somewhat exserted at maturity.
FlG. 12. Hymenophyllum. a-b, H. fucoides: a, habit, X 1; portion of lamina, show-
ing serrate segments, X 2; c, H. polyanthos, portion of lamina, showing entire seg-
ments, X 2 ; d-e, H. crispum : d, habit, X 1 ; e, portion of lamina, X 4 ; f-g, H. lanatum :
f, habit, X Vi ; g, apex of lamina, X 3'/2.
57
58 FIELDIANA: BOTANY, VOLUME 39
Further comparison of this species with H. abruption appears
under discussion of the latter.
Juvenile or depauperate specimens of H. polyanthos and H.
myriocarpum are frequently determined as H. brevifrons. In these
stages, leaves of the former may be not only minute, but the pinnae
also quite crowded, thus approximating the subflabellate appear-
ance of H. brevifrons ; hence a combination of characters must be
compared for proper identification. In H. brevifrons the petiole is
always glabrous, nonalate (except sometimes at the apex), and very
delicate: 0.1-0.15 mm. in diameter. The pinnae are always simple or
bifid, although the basal pair may be sometimes twice bifid. Each
leaf bears only 1-2 (3) terminal sori. In//, polyanthos and//, myri-
ocarpum petioles (even in depauperate specimens) are often alate or
marginate, at least partially to base, with a few reddish trichomes,
and are commonly about 0.3 mm. (rarely 0.2 mm) in diameter. Most
pinnae are pinnately divided, and sori are 2-several — not neces-
sarily terminal on the blade.
With H. brevifrons perhaps belongs H. brevistipes Liebm. of
Mexico. I have not seen the type of the latter, but Liebmann's
description (Kongel. Danske Vidensk. Selsk. Skr. V. 1: 290 [seors.
138] 1849) indicates no significant differences, nor do specimens so
identified in herbaria.
Hymenophyllum contortum v.d. Bosch, Ned. Kruidk. Arch 5:
170. 1863. Mecodium contortum (v.d. Bosch) Copel. Philipp. J. Sci.
67: 26. 1938.
On tree trunks or moist banks, in forests, 1,200-1,500 m. ; Alta
Verapaz. Southern Mexico; Costa Rica; Colombia; Venezuela; Peru
(?).
Plants usually epiphytic; rhizome thin, delicate, provided with scattered, simple,
reddish trichomes; leaves subdistant on the rhizome, mature ones (in ours) 5-15 cm.
long; petiole 1.2-6 cm. long, 0.15-0.2 mm. in diameter, provided at base with a few
simple, reddish trichomes or glabrescent, nonalate; lamina 3.5-9 cm. long (in ours),
1.2-2 cm. broad, glabrous throughout, 2- to 3-pinnatifid, gradually or not at all nar-
rowed to the base, the lower 1-4 pairs of pinnae somewhat to greatly reduced; rachis
mostly alate, but nonalate in proximal portion of lamina, the wing conspicuously
crispate; pinnae numerous, adnate, or one or two proximal ones short-stalked; ulti-
mate segments 0.5-1 mm. broad, entire, but contorted (often appearing almost
dentate or lobed) due to the strongly crispate tissue; sori several and terminal or
lateral on each pinna; indusia as broad or broader than the ultimate segments, the
valves suborbicular, broadly ovate or elliptic, slightly or not immersed at the cune-
ate or truncate base, often appearing stalked due to the constricted or strongly
crispate segment tip; receptacle narrow-cylindric, short or rudimentary.
STOLZE: FERNS OF GUATEMALA 59
The question of whether or not Hymenophyllum contortum is
truly distinct from H. undulatum must be resolved by future study.
The few collections available from Guatemala and other Central
American areas appear to be sufficiently different from H. undu-
latum, but in some specimens further outside our range the differ-
ences are less obvious. Generally, at least in our area, H. contortum
is a rather small, inconspicuous plant, rarely over 15 cm. long, with
blade only about two or three times the length of the petiole, and
quite appropriately named for the severely contorted tissue of the
rachis wing and ultimate segments. H. undulatum is typically a
slender, graceful plant, festooning tree trunks and branches in long,
delicate strands. The lamina is usually six to eight times as long as
the petiole, and the ultimate segments are rather gently undulating,
rarely (if at all) slightly crispate. However, leaf shape and degree of
crisping of the tissues are about the only significant differences be-
tween the two, and these do not appear to hold up very well
throughout the entire range. There are some South American and
West Indian plants much less elongated than in the typical condi-
tion oiH. undulatum, and, at least in portions of the leaves, the seg-
ments appear more than merely "undulate." Subgenus Mecodium
in the New World is long overdue for good monographic study, and
only at that time can the precise relationship of the two taxa be as-
certained.
Hymenophyllum crassipetiolatum Stolze, Amer. Fern J. 66:
15-18. 1976.
Pendent from tree trunks, or growing on moist banks, in deep
shade in cloud forests, 1,250-3,300 m. ; Alta Verapaz; Baja Verapaz;
Chiquimula; El Progreso; El Quiche; Zacapa. Mexico (Chiapas);
Honduras; El Salvador.
Plants epiphytic or terrestrial; rhizome wiry, long-creeping, provided with simple,
reddish to light brown trichomes; leaves subdistant on the rhizome, indeterminate,
pendent, mature ones 12-42 cm. long, 3-8 (10) cm. broad; petiole 4-11 (16) cm. long,
(0.5) 0.6-0.9 mm. in diameter, nonalate (although basal pinna sometimes short-
decurrent), sparsely to abundantly provided with simple, bifid, or (mostly) stalked,
stellate trichomes; lamina ovate or, more commonly, narrow-lanceolate, not reduced
at base, or the lower 1-4 pairs of pinnae somewhat shorter; rachis broadly alate
throughout, or nonalate at base, the wings plane to slightly crispate, sparsely or
abundantly provided with sessile or subsessile, stellate trichomes; pinnae commonly
bipinnatifid, the costae broadly alate; pinna segments 4-12 pairs, the larger ones
deeply pinnatifid, the apical ones bifid to simple; ultimate segments plane or slight-
ly undulate, entire, veins and leaf surfaces sparsely to abundantly provided with
sessile, stellate trichomes; marginal trichomes abundant, sessile-stellate, most of
60 FIELDIANA: BOTANY, VOLUME 39
the 4-6 stout rays commonly appressed and directed toward the segment tip; indu-
sium as broad or broader than the ultimate segment, not or scarcely immersed in the
segment tip, the valves as broad or broader than long, their margins provided with
simple or bifurcate trichomes.
This plant could be confused in Guatemala only with H. sieberi,
but similarity between the two is superficial. Besides the charac-
teristics listed in the key, H. crassipetiolatum also differs from H.
sieberi in that the valves of most indusia are broader than long and
are provided with simple to bifurcate trichomes. Most indusia ofH.
sieberi have valves slightly longer than broad, with bifurcate to stel-
late trichomes.
Specimens ofH. crassipetiolatum are often found in herbaria mis-
identified as H. trapezoidale Liebm. The latter apparently does not
occur in Guatemala, and differs from H. crassipetiolatum in that
segments are glabrous between veins and margin, petioles are more
delicate (less than 0.6 mm. thick) and generally lack stellate tri-
chomes, and the rachis is often nonalate in the lower portion. The
rachis in H. crassipetiolatum is alate throughout, or is lacking
wings just at the base of the blade.
Hymenophyllum crispum HBK. Nov. Gen & Sp. 1: 26: 1815.
Sphaerocionium crispum (HBK.) Kl. Linnaea 1: 537: 1844. Hymeno-
phyllum schiedeanum v.d. Bosch, Ned. Kruidk. Arch. 4: 414. 1859.
In deep shade, on trees, slopes, and banks of streams and ravines,
1,800-3,000 m. ; Quezaltenango ; El Quiche; San Marcos. Jamaica;
Mexico; Costa Rica; Colombia; Venezuela; Peru; Bolivia; Brazil.
Plants terrestrial or epiphytic; rhizome thin, long-creeping, delicate, sparsely pro-
vided with lax, filiform, light brown trichomes; leaves subdistant on the rhizome,
determinate, pendent or often forming dense mats, mature ones 4-20 cm. long;
petiole delicate, 0.5-3 cm. long, 0.15-0.2 mm. in diameter, inconspicuously alate
toward the apex, sparsely pilose, the trichomes mostly simple; lamina linear to nar-
row-ovate, the lower 1-4 pair of pinnae slightly reduced, leaf surface glabrous be-
tween veins and margin; rachis conspicuously alate throughout, the wing undulate
to crispate, provided with scattered, simple to stellate trichomes; pinnae deeply
pinnatisect, the costae broadly alate; pinna segments 2-5 pairs, simple, or some
basal ones bifid, undulate to crispate, entire, trichomes on the margin scattered,
mostly simple, those on the veins stellate and scattered abaxially, lacking adaxially;
indusium as broad or broader than the ultimate segment, scarcely immersed in the
tissue, the valves suborbicular, ovate or elliptic, their margins provided with scat-
tered, simple trichomes.
Hymenophyllum ectocarpon Fee, Mem. Fam. Foug. 11: 115, t.31.
f.l. 1866.
STOLZE: FERNS OF GUATEMALA 61
On tree trunks in dense forests, 300-1,200 m.; Alta Verapaz; Iza-
bal. British Honduras; Nicaragua; Costa Rica; Lesser Antilles.
Plants epiphytic; rhizome thin, creeping, sparsely provided with simple, reddish
trichomes; leaves subdistant on the rhizome, mature ones 7-15 cm. long; petiole 1-2
cm. long, 0.3-0.4 mm. in diameter, provided with scattered, simple, reddish tri-
chomes, very rarely narrow-alate (though often thin-marginate); lamina of mature
leaves 6-13 cm. long, 2.5-4.5 cm. broad, lanceolate to ovate, glabrous except on the
major axes, commonly tripinnatifid, not or scarcely narrowed at base, occasionally
the proximal 1-2 pairs of pinnae slightly reduced; rachis narrow-alate, or marginate
toward the base, the wing plane; pinnae adnate, or the basal pair short-stalked,
mostly symmetrical, or sometimes asymmetrical at base (truncate acroscopically,
oblique basiscopically); ultimate segments 1-1.8 mm. broad, plane, sharply and
deeply serrate throughout, the apices rounded or retuse; sori 1-5 on the acroscopic
side of each pinna and often 1-2 (3) on the basiscopic side; indusium somewhat or
not at all constricted at the cuneate base, scarcely to partially immersed in the tis-
sue, the valves acute or obtuse, lanceolate to elliptic or obovate, lacinate to subcili-
ate in the apical half; receptacle stout-cylindric or slightly thickened at base.
Hymenophyllum ectocarpon is only provisionally maintained here
as a separate species. It appears to differ from//, fucoides (at least
in Guatemala) merely in the shape and number of indusia. There-
fore, positive identification of sterile material is difficult or impos-
sible. Generally, H. ectocarpon seems to be a more delicate plant,
with petiole a bit shorter and thinner, and the lamina relatively
longer and narrower. Also, from Guatemalan collections examined,
H. ectocarpon apparently prefers lower elevations (300-1,200 m.),
while/?, fucoides is found at 1,250-3,150 m.
Hymenophyllum elegans Sprengel, Syst. Veg. 4: 133. 1827. //.
caudatellum Christ, Bull. Herb. Boissier II. 4: 939. 1904. Sphaero-
cionium elegans (Sprengel) Copel. Philipp. J. Sci. 67: 32. 1938.
Thus far known from only one collection in Guatemala (Maxon &
Hay 3329, tree trunk, trail from Senahu to Actala, Alta Verapaz).
Costa Rica; Panama; West Indies; Colombia; Venezuela; Brazil;
Peru; Bolivia.
Plants epiphytic; rhizome thin, long-creeping, delicate, sparsely provided with
lax, filiform, light brown trichomes; leaves subdistant on the rhizome, determinate,
pendent or often somewhat matted, mature ones 5-25 cm. long; petiole delicate, 0.5-
2 cm. long, 0.15-0.2 mm. in diameter, nonalate, sparsely pilose with simple or bifid
trichomes; lamina commonly linear to linear-lanceolate, rarely ovate, the lowest 1-2
pairs of pinnae slightly reduced, leaf surface glabrous between veins and margin;
rachis (at least on mature leaves) alate except near base of lamina, or most pinnae
conspicuously decurrent, thus the rachis appearing discontinuously alate, provided
with scattered, simple or bifurcate trichomes; pinnae sessile to adnate, pinnatisect
(or rarely some segments cut entirely to the costa), often appearing subdigitate;
pinna segments 1-2 (3) pairs, simple, or the basal acroscopic one bifid, plane or
62 FIELDIANA: BOTANY, VOLUME 39
slightly undulate, trichomes simple to bifid, scattered to abundant on the margins,
rare to lacking on the veins; indusia as broad or broader than the ultimate segments,
scarcely or not immersed in the segment tip, the valves suborbicular or elliptic, their
margins provided with abundant, mostly simple trichomes.
This and H. lineare do not differ significantly. See discussion of
the latter for additional comments.
Hymenophyllum elegant ulum v.d. Bosch, Ned. Kruidk. Arch. 4:
408. 1859. H. pulchellum sensu Hook. Sp. Fil. 1: 91, pro parte, and
t.33A. 1846 (not//, pulchellum Schlecht. & Cham. 1830. Sphaeroci-
onium elegantulum (v.d. Bosch) Copel. Philipp. J. Sci. 67: 32. 1938.
Pendent on tree trunks or occasionally on moist, shaded cliffs, in
deep forests, 2,400-3,000 m. ; Baja Verapaz; Chimaltenango; Hue-
huetenango; El Progreso; Quezaltenango. Mexico; Honduras; El
Salvador; Costa Rica; Panama; Greater Antilles; Venezuela;
Colombia to Bolivia.
Plants commonly epiphytic; rhizome filiform, long-creeping, provided with sim-
ple, light brown trichomes; leaves distant on the rhizome, indeterminate, pendent,
mature ones 8-25 cm. long, 2.5-5 cm. broad; petiole 1.5-4 cm. long, 0.2-0.25 mm. in
diameter, nonalate (although basal pinna sometimes short-decurrent), sparsely pro-
vided with simple (rarely bifurcate) trichomes; lamina linear to broadly lanceolate,
the lower 1-4 pairs of pinnae somewhat or greatly reduced; rachis alate above, non-
alate toward the base, rather abundantly stellate- hirsute; pinnae deeply pinnatisect,
the costae broadly alate; pinna segments 4-6 pairs, mostly simple, occasionally bifid
(or rarely the basal acroscopic ones trifid); ultimate segments plane or slightly
undulate, entire, veins and leaf surfaces rather abundantly provided with short-
stalked, stellate trichomes; marginal trichomes stellate or bistellate, subsessile to
short-stalked; indusium as broad as or slightly narrower than the ultimate segment,
not or slightly immersed, the valves usually longer than broad, their margins pro-
vided with stalked, stellate trichomes.
This and //. pulchellum are sometimes confused in herbaria. Fur-
ther information may be found under discussion of the latter.
Hymenophyllum fragile (Hedw.) Morton, Contr. U.S. Natl. Herb.
29: 172. 1947. Trichomanes fragile Hedw. Fil. Gen. t.18. 1802. Hy-
menophyllum intercalatum Christ, Bull. Herb. Boissier II. 4: 942.
1904.
Growing on tree trunks, often in mats (outside Guatemala some-
times reported found on wet, mossy banks), 750-1,150 m.; Alta
Verapaz. Mexico; Costa Rica; Panama; Greater Antilles; Colom-
bia; Venezuela; Peru; Bolivia; Brazil.
Plants commonly epiphytic; rhizome filiform, long-creeping, provided with sim-
ple, delicate, light brown trichomes; leaves determinate, subdistant on the rhizome,
mature ones (in ours) 3-6 cm. long, ca. 1.5 cm. broad; petiole 0.1-1 cm. long (in ours),
STOLZE: FERNS OF GUATEMALA 63
0.1-0.2 mm. in diameter, nonalate (although basal pinna sometimes short-decurrent),
sparsely to abundantly provided with simple or bifurcate trichomes (or some stel-
late ones at apex) ; lamina ovate to linear-lanceolate, scarcely or not reduced at base;
rachis broadly alate throughout, rather abundantly provided with sessile, stellate
trichomes; pinnae simple to bifid, or rarely some basal ones twice bifid or subflabel-
late; ultimate segments plane or slightly undulate, entire, veins and leaf surface
sparsely to abundantly provided with sessile, stellate trichomes; marginal tri-
chomes sessile or subsessile, stellate, rather abundant; indusium as broad or slight-
ly narrower than the ultimate segment, not or slightly immersed, the valves more or
less as long as broad, their margins rather abundantly provided with stalked, stel-
late trichomes.
From Costa Rica to South America, the species is frequently
found at higher elevations (to ca. 2,000 m.) and plants are generally
larger than those in Guatemala, with leaves to 15 cm. long, petioles
to 2.5 cm. long, and pinnae more commonly once or twice bifid.
Hymenophyllum fucoides (Sw.) Sw. J. Bot. (Schrader) 1800 (2):
99. 1801. Trichomanes fucoides Sw. Prodr. 136. 1788. Hymenophyl-
lum spinulosum HBK. Nov. Gen. & Sp. PI. 1: 26. 1815. Leptocio-
nium fucoides (Sw.) Presl, Hymenoph. 27. 1843.
On tree trunks in dense forests, 1,250-3,150 m.; Alta Verapaz;
Chimaltenango; Huehuetenango; Jalapa; Quezaltenango; El Qui-
che; Zacapa. Southern Mexico; Honduras to Panama; West Indies;
Colombia and Venezuela to Brazil and Bolivia.
Plants epiphytic; rhizome thin, creeping, sparsely provided with simple, reddish
trichomes; leaves subdistant on the rhizome; mature ones 8-20 cm. long; petiole 1.5-
4 cm. long, 0.4-0.7 mm. in diameter, provided with scattered, simple, reddish tri-
chomes, nonalate (although often thin-marginate); lamina of mature leaves 7-17 cm.
long, 1.5-5 cm. broad, ovate to linear-lanceolate, glabrous except on the major axes,
commonly tripinnatifid, somewhat narrowed at base, the lower 1-5 pairs of pinnae
slightly reduced; rachis narrow-alate, or marginate toward the base, the wing plane;
pinnae adnate, or the proximal 1-3 pairs short-stalked, mostly symmetrical, or some-
times asymmetrical at base (truncate acroscopically, oblique basiscopically); ulti-
mate segments 1-1.8 mm. broad, plane, sharply and deeply serrate throughout, the
apices rounded or somewhat retuse; sori 1-2 per pinna, borne acroscopically on the
basal pinnules (very rarely a single sorus may be borne basiscopically); indusium
strongly constricted at base, almost appearing short-stalked, not (or rarely slightly)
immersed in the tissue, the valves acute or obtuse, lanceolate to elliptic or obovate,
entire or serrulate near the apex; receptacle thickened to narrow- turbinate, slightly
to greatly exserted at maturity.
This and other species of section Ptychophyllum are in need of
further study. As Morton suggested, H. fucoides may well be an
aggregate species ; and one of the components is most certainly H.
ectocarpon, under which see additional comments.
64 FIELDIANA: BOTANY, VOLUME 39
Hymenophyllum hirsutum (L.) Sw. J. Bot. (Schrader) 1800 (2):
99. 1801. Trichomanes hirsutum L. Sp. PL 1098. 1753. T. ciliatum
Sw. Prodr. Veg. Ind. Occ. 136. 1788. Hymenophyllum ciliatum (Sw.)
Sw. J. Bot. (Schrader) 1800 (2). 100. 1801. Sphaerocionium ciliatum
(Sw.) Presl, Hymenoph. 34. 1843. S. hirsutum (L.) Presl, loc. cit. H.
microcarpum Fee, Crypt. Vase. Bres. 1: 245. 1869 (not//, microcar-
pum Desv. 1827).
In deep shade, pendent from tree trunks, or growing on moist
banks or rotting logs in wet forests, 150-1,800 m.; Alta Verapaz;
Baja Verapaz; Chiquimula; Huehuetenango; Izabal; Peten; El Qui-
che. West Indies; southern Mexico; Honduras; Nicaragua; Costa
Rica; Colombia to the Guianas, south to Bolivia and Brazil.
Plants epiphytic or terrestrial; rhizome thin, wiry, long-creeping, provided with
simple, reddish to light brown trichomes; leaves subdistant on the rhizome, deter-
minate, pendent or spreading, mature ones 3-12 cm. long; petiole 0.5-2.5 cm. long,
0.2-0.3 (0.4) mm. in diameter, alate one-half to three-quarters to base, abundantly
provided with bifurcate to (mostly) stellate trichomes; lamina ovate to oblong, the
lower 2-3 pairs of pinnae somewhat to greatly reduced, leaf surface glabrous between
veins and margin; rachis conspicuously alate throughout, the wings plane to slight-
ly crispate, abundantly provided with bifurcate to (mostly) stellate trichomes;
pinnae deeply pinnatifid, the costae broadly alate; pinna segments 2-5 pairs, simple,
or some basal ones bifid, plane to very slightly undulate, entire, marginal trichomes
abundant, bifurcate or stellate, those of the veins mostly stellate on both sides; in-
dusium about as broad as the ultimate segment, not or scarcely immersed in the
segment tip, the valves suborbicular, their margins abundantly provided with
simple to bifurcate trichomes.
This is easily confused with H. maxonii, under which see further
discussion.
Hymenophyllum lanatum Fee, Mem. Fam. Foug. 11: 116. 1866.
Sphaerocionium lanatum Copel. Philipp. J. Sci. 67: 31. 1938.
Growing on tree trunks or large, mossy logs, often in dense mats,
900-1,500 m.; Alta Verapaz; Baja Verapaz; Huehuetenango. West
Indies; Surinam; Venezuela(?).
Plants commonly epiphytic; rhizome filiform, long-creeping, provided with sim-
ple, delicate, yellow-brown trichomes; leaves determinate, subdistant on the rhi-
zome, mature ones 2-5 cm. long (in ours), 1-1.5 cm. broad; petiole 0.5-3 cm. long, 0.1-
0.15 mm. in diameter, nonalate, sparsely provided with stalked, stellate trichomes;
lamina ovate to lanceolate, scarcely or not reduced at base, divided nearly to the
rachis into linear segments; rachis broadly alate throughout, rather abundantly pro-
vided with short-stalked, stellate trichomes; primary segments simple, or rarely
some basal ones bifid, essentially plane, entire, veins and leaf surface abundantly
provided with short-stalked, stellate trichomes; marginal trichomes stellate or bi-
stellate, obviously stalked, rather abundant; indusium slightly narrower than the
STOLZE: FERNS OF GUATEMALA 65
segments, not or slightly immersed, the valves about as long as broad, their margins
rather abundantly provided with stalked, stellate trichomes.
Hymenophyllum lineare (Sw.) Sw. J. Bot. (Schrader) 1800 (2):
100. 1801. Trichomanes lineare Sw. Prodr. Veg. Ind. Occ. 137. 1788.
Didymoglossum lineare (Sw.) Desv. Mem. Soc. Linn. Paris 6: 331.
1827. Sphaerocionium lineare (Sw.) Presl, Hymenoph. 34. 1843.
Hymenophyllum elegantissimum Fee, Mem. Fam. Foug. 11: 118.
1866.
In shade, on tree trunks, 600-1,200 m.; Alta Verapaz; Baja Vera-
paz. Costa Rica; West Indies.
Plants epiphytic; rhizome thin, long-creeping, delicate, sparsely provided with
lax, filiform light brown, mostly simple trichomes; leaves commonly pendent,
mature ones 5-15 cm. long; petiole delicate, 0.5-3 cm. long, 0.15-0.2 mm. in diameter,
nonalate, sparsely pilose with simple or bifurcate trichomes; lamina commonly
ovate or narrow-oblong, the lowest 2-3 pairs of pinnae slightly reduced, leaf surface
glabrous between veins and margin; rachis nonalate in proximal half, or often nearly
to apex, provided with scattered, simple to bifurcate trichomes; pinnae mostly
adnate to sessile, or several lower pairs short-stalked, deeply pinnatisect (or some
basal segments cut entirely to costa), the costa commonly alate throughout (often
very narrowly so); pinna segments 3-5 pairs, simple to bifid, plane or slightly undu-
late, trichomes simple or forked at base, scattered on the margins, rare or lacking on
the veins on both sides; indusia commonly much broader than the ultimate seg-
ments (often conspicuously so), not or rarely immersed in the segment tip, the
valves suborbicular to elliptic, their margins provided with scattered, simple
trichomes.
Besides the characters noted in the key, Hymenophyllum lineare
often differs from H. elegans in the shape of the mature leaf, that of
the former mostly broader and ovate, and that of the latter much
narrower. However, the two may be merely ecological variants, the
few distinctions perhaps resulting from the exposed or protected,
dry or wet habitat in which the individual occurs. All their apparent
differences are due to constriction of tissue. Thus, in H. lineare seg-
ments and wings are so narrow that the pinnae often appear elon-
gated and skeletal, and the indusia conspicuously broader, (a paral-
lel condition occurs in two closely related species of Trichomanes: T.
capillaceum andT. angustatum).
Neither H. lineare nor H. elegans are adequately represented in
herbaria. It is hoped that additional collections, more careful obser-
vations, and better field notes will further establish the proper rela-
tionship of these species in the future.
Hymenophyllum maxonii Christ ex Morton, Contr. U.S. Natl.
Herb. 29: 165. 1947. Sphaerocionium maxonii (Christ ex Morton)
66 FIELDIANA: BOTANY, VOLUME 39
Pic. Ser. Webbia 28: 471. 1973. H. maxonii var. angustius Morton,
Contr. U.S. Natl. Herb. 29: 166. 1947.
In deep shade, on tree trunks, in forests or wooded ravines, 1,000-
2,000 m. ; Alta Verapaz (type collection: on trail from Senahu to Ac-
tala, Maxon & Hay 3326); Baja Verapaz; Chiquimula; Huehuete-
nango.
Plants epiphytic; rhizome thin, wiry, long-creeping, provided with simple, reddish
trichomes; leaves subdistant on the rhizome, determinate, commonly pendent,
mature ones 6-30 cm. long; petiole 2.5-7 (10) cm. long, (0.2) 0.3-0.5 mm. in diameter,
rather conspicuously alate at the apex or, rarely, along the upper third, the wing
abruptly terminating, provided with simple to bifurcate trichomes; lamina ovate to
oblong, the lower 2-3 pairs of pinnae somewhat to greatly reduced, the leaf surface
glabrous between veins and margin; rachis conspicuously alate throughout, the
wings plane to slightly crispate, abundantly provided with simple or bifurcate
(rarely stellate) trichomes; pinnae deeply pinna tif id, the costae broadly alate; pinna
segments simple to bifid, the basal acroscopic ones often again bifid, plane to
slightly undulate, entire, marginal trichomes abundant, bifurcate or stellate, those
on the veins mostly stellate on both sides; indusium about as broad as the ultimate
segment, not or scarcely immersed in the segment tip, the valves suborbicular, their
margins abundantly provided with simple to bifurcate trichomes.
Morton further recognized var. angustius from Alta Verapaz,
presumably differing from the typical H. maxonii in its longer and
relatively narrower leaves.
There are few significant differences between H. maxonii and H.
hirsutum. Perhaps H. maxonii may be nothing more than a more
luxuriant variety of the latter, having somewhat more complex pin-
nae, with blade and petiole longer, but with a comparatively shorter
petiole wing. H. hirsutum apparently has petioles with predomi-
nantly stellate trichomes, while those of H. maxonii are simple to bi-
furcate. However, the stellate trichomes are evidently produced
from the wing (rather than from the petiole itself), thus H. hir-
sutum, having a much greater percentage of the petiole alate, has an
abundance of stellate trichomes. Conversely, the petiole oiH. max-
onii appears predominantly simple-hirsute, since only the abbrevi-
ated wing near the apex has stellate trichomes.
Hymenophyllum microcarpum Desv. Mem. Soc. Linn. Paris 6:
333. 1827 (not H. microcarpon Fee, 1869). H. angustifrons Christ,
Bull. Herb. Boissier II. 4: 940. 1904. Mecodium microcarpum
(Desv.) Copel. Philipp. J. Sci. 67: 25. 1938. Sphaerocionium micro-
carpum (Desv.) Copel. torn. cit. 34.
In deep shade, commonly on tree trunks, in forests or wooded
ravines, 900-2,600 m.; Alta Verapaz; Baja Verapaz; Huehuetenan-
STOLZE: FERNS OF GUATEMALA 67
go; Jutiapa; Zacapa. Honduras and El Salvador to Panama;
Greater Antilles; Colombia to the Guianas, southward to Brazil and
Bolivia.
Plants commonly epiphytic; rhizome wiry, long-creeping, rather densely provided
with simple, light brown or reddish trichomes; leaves distant, determinate, mature
ones 10-30 cm. long, petiole 3-9 (12) cm. long, 0.5-0.9 mm. in diameter, rather con-
spicuously alate for nearly half its length, sparsely to abundantly provided with
stellate trichomes, or glabrescent; lamina ovate or broadly lanceolate, only the basal
pair of pinnae scarcely (or not at all) reduced, leaf surface glabrous between veins
and margin; rachis conspicuously alate throughout, the wing undulate to crispate,
sparsely to abundantly provided with stellate trichomes; pinnae deeply pinnatisect,
the costae broadly alate; pinnules 4-8 pairs, deeply pinnatifid; ultimate segments
linear, plane or somewhat undulate, entire, trichomes on the margin scattered,
mostly simple, or once-forked at base, those on the veins stellate and scattered abax-
ially, lacking adaxially; indusia commonly much broader than the ultimate seg-
ments, not immersed in the tissue, the valves ovate or elliptic, their margins pro-
vided with scattered, simple trichomes.
Hymenophyllum myriocarpum Hook. Sp. Fil. 1: 106. 1844. Sphae-
rocionium myriocarpum (Hook.) Kl., Linnaea 18: 536. 1844. Meco-
dium myriocarpum ( Hook. ) Copel. Philipp. J. Sci. 67: 25. 1938.
On tree trunks and branches, in forests, from sea level to 2,800 m.;
AltaVerapaz; Baja Verapaz; Chimaltenango; Chiquimula; Izabal;
San Marcos; Solola. Mexico; Honduras; El Salvador; Costa Rica;
Colombia and Venezuela to Bolivia.
Plants epiphytic; rhizome thin, creeping, provided with scattered, simple, reddish
trichomes; leaves subdistant on the rhizome, mature ones 10-25 (30) cm. long; peti-
oles 1.5-4 (5) cm. long and 0.3-0.7 mm. in diameter, provided with scattered, simple,
reddish trichomes or glabrescent, marginate or alate partly or fully to base (often
deciduous); lamina of mature leaves 8-21 (24) cm. long, 1.5-3.5 cm. broad, commonly
4-7 times as long as the petiole, glabrous throughout, 2- to 3-pinnatifid, gradually
narrowed at base, the lower 3-5 pairs of pinnae somewhat to greatly reduced; rachis
alate or marginate throughout, the wing plane to undulate or revolute (occasionally
somewhat crispate); pinnae numerous, adnate; ultimate segments 1 mm. broad or
less, entire, plane to undulate; sori one to several on most pinnules; indusia much
broader than the greatly constricted segment tips, thus appearing stalked, the
valves mostly suborbicular or broadly ovate; receptacle narrow-cylindric, short to
rudimentary.
This is a highly problematical species, perhaps synonymous with
H. polyanthos. For further information, see discussion of the latter.
Hymenophyllum polyanthos (Sw.) Sw. J. Bot. (Schrader) 1800
(2): 102. 1801. Trichomanes polyanthos Sw. Prodr. 137. 1788. Meco-
dium polyanthos (Sw.) Copel. Philipp. J. Sci. 67: 19. 1938.
68 FIELDIANA: BOTANY, VOLUME 39
On trunks and branches of trees, and on fallen logs, occasionally
reported on clay banks, in deep forests, from sea level to 3,100 m.;
Alta Verapaz; Baja Verapaz; Chimaltenango ; Chiquimula; Escuin-
tla; Guatemala; Huehuetenango; Izabal; Jalapa; El Progreso; El
Quiche; San Marcos; Solola; Suchitepequez; Zacapa. Mexico to
Panama; West Indies; South America; Old World tropics and sub-
tropics.
Plants commonly epiphytic; rhizome thin, creeping, provided with scattered, sim-
ple, reddish trichomes; leaves subdistant on the rhizome, mature ones 7-17 cm. long;
petiole 2-5 cm. long and 0.3-0.7 mm. in diameter, provided with scattered, simple,
reddish trichomes or glabrescent, nonalate, or more commonly marginate or alate
partly or fully to base (wings often deciduous); lamina of mature leaves 5-12 cm.
long, 2-6 cm. broad, about 1.5-3.5 times as long as the petiole, glabrous throughout,
2- to 3-pinnatifid, not or scarcely reduced at base; rachis alate or marginate through-
out, the wing plane to undulate or revolute (rarely somewhat crispate); pinnae
several to many pairs, adnate, or the lower one short-stalked; ultimate segments
1 mm. broad or less, entire, plane to undulate, or rarely slightly crisped; sori often
several on each pinnule; indusium narrower to slightly broader than the segment
tips, the valves mostly ovate to elliptic, somewhat immersed in the tissue; recep-
tacle narrow-cylindric, short or rudimentary.
This is a highly variable species, one of the most widely distrib-
uted ferns of tropical regions throughout the world, which ranges in
altitude from sea level to the montane forests. It is a member of the
subgenus Mecodium , whose species are characterized by entire seg-
ments and glabrous laminae. This very lack of pubescence, coupled
with the sameness of leaf margins, renders it a taxonomically diffi-
cult group. Although Copeland studied the species as it occurs in
parts of the Old World, very little work has been done with it in the
neotropics; consequently the relationship with its New World allies
is not well understood. Monographic work may reveal that dozens
of so-called "species" in tropical America are but synonyms of H.
polyanthos, as Copeland discovered in the Old World. One of these
may be the Guatemalan H. myriocarpum, which is distinguished
primarily by leaf shape, a characteristic notably variable through-
out the Hymenophyllaceae. The differences listed in the key are
mostly quantitative and may be nothing more than products of
varying habitats or availability of moisture.
Hymenophyllum pulchellum Schlecht. & Cham. Linnaea 5: 618.
1830. Sphaerocionium pulchellum (Schecht. & Cham.) Presl, Hy-
menoph. 34. 1843. H. pannosum Christ, Bull. Herb. Boissier II. 5:
250. 1905.
STOLZE: FERNS OF GUATEMALA 69
In shade, in or at edges of forests or in wooded ravines, pendent
from logs, tree trunks, or rocky banks, 250-2,500 m.; Alta Verapaz;
Baja Verapaz; San Marcos; Solola; Suchitepequez. Southern Mexi-
co; British Honduras; Honduras; Costa Rica; southeastern Brazil.
Plants terrestrial or epiphytic; rhizome thin, long-creeping, delicate, sparsely pro-
vided with lax, filiform, light brown or reddish trichomes; leaves subdistant on the
rhizome, indeterminate, commonly pendent, mature ones 6-30 cm. long; petiole deli-
cate, 2.5-4 cm. long, 0.2-0.3 mm. in diameter, nonalate, sparsely provided with sim-
ple to stellate trichomes; lamina linear to oblong, the lower 2-6 pairs of pinnae often
greatly reduced, leaf surfaces glabrous between veins and margins; rachis nonalate
except near apex, sparsely to abundantly provided with stellate trichomes ; pinnae
mostly short-stalked, deeply pinnatifid, the costa continuously alate on both sides;
pinna segments 4-8 pairs, simple, or some basal ones bifid, plane or slightly undu-
late, entire, trichomes on the margin and veins abundant, sessile, stellate; indusium
about as broad as or narrower than the ultimate segment, scarcely immersed in the
tissue, the valves suborbicular or hemispherical, their margins abundantly provided
with stellate trichomes.
Trichomes on the margins and veins in this species are often so
dense and long that they appear to spring from the intervening tis-
sue as well. Consequently, specimens of H. pulchellum are some-
times found in herbaria filed with species of subsection Hirsuta
usually determined as H. elegantulum. Trichomes on the latter,
however, are rarely as densely crowded, and their points of attach-
ment on the laminar surface are generally quite obvious. Moreover,
most of the trichomes on the segment margins of H. elegantulum
are short-stalked, whereas those of H. pulchellum are commonly
sessile.
Hymenophyllum sieberi (Presl) v.d. Bosch, Ned. Kruidk. Arch. 4:
414. 1859. Sphaerocionium sieberi Presl, Hymenoph. 58. 1843. Hy-
menophyllum wercklei Christ, Bull. Herb. Boissier II. 4: 940. 1904.
Sphaerocionium wercklei (Christ) Copel. Philipp. J. Sci. 67: 31.
1938.
Pendent from tree trunks and branches, or on mossy logs, in and
at edges of dense forests, 950-1,600 m.; Alta Verapaz; Huehuete-
nango. Mexico (Chiapas) to Costa Rica; Puerto Rico; Guadeloupe;
Martinique.
Plants commonly epiphytic; rhizome wiry, long-creeping, provided with simple,
reddish or light brown trichomes ; leaves subdistant on the rhizome, indeterminate,
pendent, mature ones 15-60 cm. long, 3-6.5 (10) cm. broad; petiole 1.5-5.5 (7) cm.
long, 0.3-0.5 mm. in diameter, broadly alate near the apex, and often narrowly alate
along the apical half, sparsely to abundantly provided with furcate or stalked, stel-
late trichomes; lamina ovate or, more commonly, linear-lanceolate, gradually but
distinctly reduced at base, the lower 4-10 pairs of pinnae much shorter; rachis
70 FIELDIANA: BOTANY, VOLUME 39
broadly alate throughout (but the wing sometimes fugacious at base of lamina), the
wings plane to slightly crispate, sparsely or abundantly provided with subsessile or
stalked, stellate trichomes; pinnae shallowly or deeply pinna tifid, the costae broadly
alate, pinna segments 3-7 pairs, simple to bifid, or rarely some larger basal ones
twice bifid or subflabellate ; ultimate segments plane or slightly undulate, entire,
veins and leaf surface sparsely to abundantly provided with subsessile or stalked,
stellate trichomes; marginal trichomes stalked, stellate, the 3-4 delicate rays com-
monly spreading; indusium as broad or broader than the ultimate segment, not or
scarcely immersed in the segment tip, the valves usually somewhat longer than
broad, their margins provided with bifurcate or stalked, stellate trichomes.
This may be confused in Guatemala with H. crassipetiolatum,
under which see further discussion. The two are part of a very close-
ly knit species complex which includes H. plumieri Hook. & Grev.,
H. lindenii Hook. and//, interruptum Kunze of South America.
Hymenophyllum tricophyllum HBK. Nov. Gen. & Sp. 1: 27. 1815.
Sphaerocionium tricophyllum (HBK.) Copel. Philipp. J. Sci. 67: 32.
1938.
In shade, growing in dense, tangled mats, commonly on tree
trunks, but (especially in Guatemala) often on moist, clay banks.
2,000-3,200 m.; Zacapa. Costa Rica; Panama; Colombia; Venezuela;
Ecuador; Bolivia.
Plants terrestrial or (outside Guatemala) more commonly epiphytic; rhizome thin,
long-creeping, delicate, sparsely provided with lax, filiform, light brown or reddish
trichomes; leaves subdistant on the rhizome, indeterminate, mature ones 3-12 cm.
long; petiole delicate, 0.5-3 cm. long, 0.1-0.2 mm. in diameter, nonalate, provided
with widely scattered, simple or bifurcate trichomes; lamina commonly linear, the
two lower pairs of pinnae scarcely reduced, leaf surfaces glabrous between veins and
margin; rachis nonalate throughout, provided with widely scattered, simple or
bifurcate trichomes; pinnae mostly short-stalked, pinnate, the costa not continu-
ously alate on both sides, or some of them greatly elongated, often becoming frond-
like, much exceeding adjacent ones; ultimate segments linear, plane or slightly un-
dulate, entire, trichomes on the margins and veins abundant, stellate; indusium
about as broad as or narrower than the ultimate segment, scarcely immersed in the
tissue, the valves suborbicular to hemispherical, their margins abundantly provided
with stellate trichomes.
This is perhaps the most delicate and certainly the most peculiar
species in section Sphaerocionium. The filiform petioles and rach-
ises entwine with adjacent leaves, often forming dense mats on
banks or tree trunks. Occasionally, one pinna becomes greatly elon-
gated and once again pinnate-pinnatifid, and then often equals the
length of the leaf on which it is borne.
Hymenophyllum undulatum Sw. J. Bot. (Schrader) 1800 (2): 101.
1800. Trichomanes undulatum Sw. Prodr. 137. 1788 (not T. undu-
STOLZE: FERNS OF GUATEMALA 71
latum Wall. 1828, nor T. undulatum v.d. Bosch. 1861). Sphaerocio-
nium undulatum (Sw.) Presl, Hymenoph. 35. 1843. Mecodium un-
dulatum (Sw.) Copel. Philipp. J. Sci. 67: 26. 1938.
Pendent from tree trunks, or on wet banks or mossy rocks, in for-
ests, 1,200-2,600 m.; Alta Verapaz; Baja Verapaz (?); Guatemala;
Zacapa. Southern Mexico; Honduras; Costa Rica; Panama; Greater
Antilles; Colombia; Ecuador; Brazil.
Plants usually epiphytic; rhizome thin, delicate, creeping, provided with scat-
tered, simple, reddish trichomes; leaves subdistant on the rhizome, mature ones 7-
35 cm. long; petiole 2-4.5 cm. long, 0.1-0.2 mm. in diameter, provided at base with a
few simple, reddish trichomes or glabrescent, nonalate; lamina of mature leaves 8-
30 cm. long, 1-3 cm. broad, glabrous throughout, 2- to 3-pinnatifid, gradually nar-
rowed to the base, the lower 2 to many pinnae somewhat to greatly reduced; rachis
marginate or narrow-alate in the distal one- half to two-thirds of the lamina (often
narrow-alate on alternate sides due to the decurrent pinna bases), the wing plane or
undulate; pinnae numerous, adnate, or long-decurrent, or one or several proximal
ones short-stalked; ultimate segments 1 mm. broad or less, entire, slightly to
strongly undulate; sori several and terminal or lateral on each pinna; indusia nar-
rower to much broader than the segments, the valves suborbicular, ovate or elliptic,
slightly or not at all immersed at the cuneate or truncate base; receptacle narrow-
cylindric, short or rudimentary.
Perhaps H. contortum should be included here. See further discus-
sion under that species.
TRICHOMANES Linnaeus
Reference: J. G. Wessels Boer, The New World species of Tricho-
manes sect. Didymoglossum and Microgonium, Acta Bot. Neerl.
11: 277-330. 1962.
Plants epiphytic or terrestrial, occasionally saxicolous; rhizome sparsely to
densely provided with brown or blackish trichomes, stout, ascending to erect with
leaves fasciculate, or (more commonly) thin, delicate and long-creeping with leaves
widely spaced; leaves monomorphous, or rarely dimorphous, simple to several times
pinnate, 0.5-80 cm. long (in ours), subsessile to long-petiolate; petiole delicate to
wiry, flattened, terete, or occasionally rounded-trigonous, in a few species sulcate on
the adaxial side, commonly shorter than the lamina and often alate for all or part of
its length; lamina suborbicular to lanceolate to subdeltoid, glabrous, glandular or
hirsute, the trichomes simple, bifid or sessile-stellate; venation pinnate (catadro-
mous or anadromous) or flabellate, the true veins free or (rarely) reticulate near the
margin, spurious veins often present, mostly parallel, but in some species perpen-
dicular, to the true veins; ultimate segments narrow, often linear to filiform, com-
monly entire, obtuse, truncate or emarginate at the apex; indusium fully, partly or
not at all immersed in the tissue, tubular, narrow-cylindrical to salverform, with an
entire or bilabiate mouth which is often widely flaring; receptacle filiform,
commonly long-exserted from the indusium at maturity; spores tetrahedral to sub-
globose, trilete, spinose, tuberculate, or nearly smooth, white or, more commonly,
greenish.
72 FIELDIANA: BOTANY, VOLUME 39
There are 250-300 species in the genus, occurring in tropical to
subtropical regions of both hemispheres. They are typically found in
deep, wet forests, on trees or on the forest floor, on clay banks of
ravines and beside streams and waterfalls; some may grow on
moist, rocky cliffs or even on wet rocks in streams. Many are
pendent from branches, while a few of those with minute leaves are
so tightly clustered as to form dense mats on tree trunks.
Of the 24 species treated here, 23 have been collected in Guate-
mala and another, T. ekmanii, has been found in British Honduras.
It may be presumed that T. ekmanii and other minute species will
be found in the future not only in additional locations in Guatemala,
but in other tropical American regions as well. Such inconspicuous
ferns undoubtedly have been overlooked by collectors in many
areas; thus current distribution records are probably far from com-
plete.
a. Leaves strongly dimorphous, fertile laminae subentire, narrow and spikelike,
sterile ones pinnatisect T. diversifrons.
a. Leaves essentially monomorphous, fertile ones often somewhat longer than the
sterile, but similar in outline.
b. Leaves sub-bipinnate to 4-pinnate; venation anadromous.
c. Mature leaf 16-50 cm. long; rhizome stout, 1 mm. or more in diameter,
d. Rhizome erect or ascending (rarely short-creeping); leaves approximate or
subcaespitose T. rigidum.
d. Rhizome long-creeping; leaves distant,
e. Indusium obviously bilabiate, lips widely flaring; petiole broadly alate
usually to base T. collariatum.
e. Indusium scarcely bilabiate, mouth rarely or never flaring; petiole not
alate, or only slightly so just below the basal pinnae T. radicans.
c. Mature leaf 2.5-15 cm. long; rhizome delicate, 0.2-0.5 mm. in diameter,
f. Tissue of most ultimate segments with elongate, narrow folds parallel to
the veins; tube of the indusium nearly as broad as long . . . . T. pyxidiferum.
f. Tissue of segments essentially plane; tube of the indusium 2 or more times
as long as broad.
g. Rachis alate throughout (in T. hymenophylloides sometimes obscurely
so),
h. Petiole alate nearly or quite to the base; rachis broadly alate, with each
wing about as broad as the rachis T. diaphanum.
h. Petiole nonalate, or (rarely) alate just at the apex; rachis narrow-alate,
with each wing much narrower than the rachis
T. hymenophylloides.
g. Rachis nonalate for most of its length,
i. Ultimate segments 0.5-1 mm. broad; costae alate except at their
very base T. angustatum.
i. Ultimate segments 0.1-0.4 mm. broad; costae of larger pinnae common-
ly nonalate at least in the proximal third T. capillaceum.
b. Leaves simple to pinnate-pinnatifid; venation catadromous or flabellate.
STOLZE: FERNS OF GUATEMALA 73
j. Mouth of indusium deeply divided into two prominent lips, these commonly
dark-margined.
k. Venation pinnate, with midrib distinct at least in lower half of lamina; lam-
ina of mature leaf commonly 1-9 cm. long.
1. Sori mostly to fully immersed in the tissue, or at least flanked on either
side by a wing of tissue; spurious veinlets frequently parallel to and very
near the margin T. krausii.
1. Sori not or scarcely immersed in the tissue; spurious veinlets rarely, if
ever, parallel to and near the margin.
m. Lips of the indusium mostly broader than long; spurious veinlets rath-
er sparse T. hymenoides.
m. Lips of the indusium mostly as long as or longer than broad; spurious
veinlets abundant T. reptans.
k. Venation flabellate, with lamina commonly lacking a distinct midrib; or,
when venation sometimes pinnate (in T. ovale), lamina of mature leaf 0.5-
0.8 cm. long,
n. Fertile leaves 0.5-0.8 cm. long and frequently with distinct midrib; sorus
solitary at lamina apex T. ovale.
n. Fertile leaves 1-3 cm. long and lacking distinct midrib; sori ( 1-) several to
many.
o. Sori borne between the lobes, rarely exserted much beyond the outline
circumscribed by lobe apices; lips of the indusium widely flar-
ing T. punctatum ssp. sphenoides.
o. Sori not borne between lobes, but conspicuously exserted from the
lamina; lips of the indusium scarcely or not at all flaring T. curtii.
j . Mouth of indusium not ( or scarcely ) bilabiate, not dark-margined.
p. Margin of lamina bearing paired, orbicular scales T. membranaceum.
p. Margin of lamina glabrous or bearing simple to stellate trichomes.
q. Spurious veinlets scattered to numerous, parallel or perpendicular to the
true veins.
r. Leaves simple or lobed, mature ones 1-3 cm. long.
s. Leaves with an uninterrupted, spurious vein submarginal along the
perimeter of the lamina; marginal trichomes lacking.
t. Spurious veinlets numerous, both parallel and perpendicular to
true veins, the venation apparently reticulate T. godmanii.
t. Spurious veinlets scattered, none perpendicular to true veins
T. ekmanii.
s. Leaves without an uninterrupted, submarginal spurious vein; mar-
ginal trichomes present, mostly stellate T. petersii.
r. Leaves pinnate or pinnatisect, mature ones commonly 5-70 cm. long,
u. Leaves subcaespitose, long-petiolate; spurious veinlets numerous
throughout each pinna, perpendicular to true veins . . . . T. pinnatum.
u. Leaves distant, subsessile; spurious veinlets few, near the margin,
parallel to true veins T. tuerckheimii.
q. Spurious veinlets absent.
v. Leaves lobed or pinnatifid; rhizome thin, horizontal and long-creeping;
trichomes on the lamina stellate T. polypodioides.
74 FIELDIANA: BOTANY, VOLUME 39
v. Leaves pinnate or pinnatisect; rhizome stout, suberect or ascending;
trichomes on the lamina simple, septate, with the basal cell much short-
ened and broadened.
w. Petiole broadly alate halfway or more to the rhizome; rachis broadly
alate, with each wing as wide or wider than the rachis; mouth of the
indusium rarely wide-flaring T. galeottii.
w. Petiole nonalate, or narrow-alate only at base of blade; rachis (at
least the lower half) narrow-alate, with each wing narrower than the
rachis; mouth of the indusium often widely flaring T. crispum.
Trichomanes angustatum Carm. Trans. Linn. Soc. 12: 513. 1818.
T. tenerum Sprengel, Syst. Veg. 4: 129. 1827., T. schiedeanum Fee,
Crypt. Vase. Bres. 1: 188. 1869 (not T. schiedeanum K. Mull. 1854).
Vandenboschia tenera (Sprengel) Copel. Phillip. J. Sci. 67: 53. 1938.
V. angustata (Carm.) Copel. op. cit. 73: 466. 1941.
In deep shade, on tree trunks or rocky cliffs or banks of ravines,
1,950-3,300 m.; Chimaltenango; El Progreso; Sacatepequez ; San
Marcos. Greater Antilles; Colombia; Brazil; Bolivia; Tristan
d'Acunha.
Plants terrestrial or epiphytic; rhizome thin, long-creeping, delicate, sparsely to
abundantly provided with light or dark brown, flattened trichomes; leaves mono-
morphous, essentially glabrous, subdistant, but commonly forming dense mats,
mature ones 4-12 (16) cm. long and 1.5-5 cm. broad, the laminae much longer than
their petioles; petiole 0.5-3 cm. long, terete to somewhat flattened, nonalate, green
to stramineous or light brown, usually darker at base; lamina linear-lanceolate, lan-
ceolate or oblanceolate, yellow- to gray-green, thin-membranaceous, translucent, bi-
pinnate-pinnatifid or tripinnate, the lower (2) 3-5 pinnae commonly reduced (often
greatly so); rachis nonalate; pinnae 8 to numerous, short-stalked, ascending, crowd-
ed or imbricate, costae alate except below the first pinnule; ultimate segments
linear, 0.5-0.8 mm. broad, emarginate, lateral margins entire, plane; veins free,
anadromous, spurious ones absent; sori 1-4 per pinna, terminating the proximal
veinlets, winged on either side by a narrow band of tissue; indusium narrow-funnel-
form, the tube 2 or more times as long as broad, the mouth flaring but not bilabiate
nor dark-margined; receptacle long-exserted; spores subglobose, densely glandular.
This may be often confused with Trichomanes capillaceum, for
both species are delicate and have very reduced ultimate segments
and nonalate petioles and rachises. Typical T. capillaceum appears
almost skeletal, with tissue present only on the ultimate segments.
However, a number of specimens (at least in Guatemala) have tis-
sue extending along the axes as far down as the distal portion of the
costa. Besides the differences noted in the key, T. angustatum can
frequently be distinguished by the several pairs of greatly reduced
pinnae at the base of the lamina, and by its possession of more than
one sorus on each pinna. T. capillaceum rarely has more than the
FIG. 13. Trichomanes. a-b, T. collariatum: a, habit, X Vi; b, portion of pinna, X 5;
c, T. mdicans, sori, X 5; d-e, T. reptans: d, habit, X 1/i; e, apex of lamina, X 4; f, T.
godmanii, habit, X Vi.
75
76 FIELDIANA: BOTANY, VOLUME 39
basal pair of pinnae reduced and very rarely more than one sorus on
each pinna.
Trichomanes capillaceum L. Sp. PL 2: 1099. 1753. T. trichoideum
Sw. J. Bot. (Schrader) 1800 (2): 98. 1801. T. tenellum Hedw. Fil.
Gen. & Sp. 1799. T. schiedeanum K. Mull. Bot. Zeit. (Berlin) 716.
1854 (not T. schiedeanum Fee, 1869). Vandenboschia capillacea (L.)
Copel. Philipp. J. Sci. 67: 53. 1938.
On trees in forest or on wet, shaded banks of ravines, 300-2,850
m.; AltaVerapaz; BajaVerapaz; El Quiche; Chiquimula; Huehue-
tenango; Izabal; Quezaltenango; San Marcos; Solola; Suchitepe-
quez; Zacapa. Greater Antilles; Southern Mexico; Honduras; El
Salvador; Nicaragua; Costa Rica; Panama; Colombia; Venezuela;
Ecuador; Peru.
Plants terrestrial or epiphytic; rhizome thin, long-creeping, delicate, sparsely to
abundantly provided with light or dark brown, flattened trichomes; leaves mono-
morphous, essentially glabrous, subdistant, mature ones 5-10 (15) cm. long and 1-5
cm. broad, the laminae usually longer than their petioles; petiole 1-5 cm. long, terete,
nonalate, green or drying stramineous, darker at base; lamina lanceolate, ovate or
oblanceolate, yellow- or gray-green, thin-membranaceous, translucent, 3- to 4-pin-
nate, the lower one or two pairs of pinnae somewhat reduced; rachis nonalate; pin-
nae 6 to numerous, short-stalked, ascending, commonly crowded or imbricate, cos-
tae of the larger ones nonalate (at least in the proximal third); ultimate segments
filiform, 0.1-0.3 mm. broad, emarginate, lateral margins entire, plane; veins free,
anadromous, spurious ones absent; sori commonly 1 per pinna (but rarely as many
as 4), terminating the proximal veinlets, mostly stalked and free from the tissue;
indusium narrow-funnelform, the tube 2 or more times as long as broad, the mouth
flaring but not bilabiate nor dark-margined; receptacle long-exserted; spores sub-
globose, densely granular.
See also discussion of Trichomanes angustatum, a species with
which T. capillaceum is frequently confused.
Trichomanes collariatum v.d. Bosch, Ned. Kruidk. Arch. 4: 368.
1859. T. martinezii Rovir. Pterid. Mex. 106. 1909 (type from Chi-
apas, Mexico, Martinez 1103). Vandenboschia martinezii (Rovir.)
Pic. Ser., Webbia 28: 472. 1973.
Wet forest and wooded ravines, climbing and often appressed to
tree trunks, 50-600 m.; Alta Verapaz; Huehuetenango; Izabal;
Peten. Southern Mexico; British Honduras; Honduras; Nicaragua;
Costa Rica; Panama; Colombia; Venezuela; Ecuador; Peru; Brazil.
Plants epiphytic; rhizome stout, long-creeping, provided with dark brown, flat-
tened trichomes; leaves monomorphous, distant, 15-40 cm. long, 4-10 cm. broad,
subsessile to short-petioled; petiole 0-3 cm. long, stout, terete, yellow- to gray-
STOLZE: FERNS OF GUATEMALA 77
brown, broadly alate, commonly with very reduced pinnae at base which overlap the
rhizome; lamina linear, lanceolate, or narrow-elliptic, dark green, often drying black-
ish, firm-membranaceous, opaque, subbipinnate to bipinnate-pinnatifid, subequally
reduced at apex and base; rachis with a few, brownish, filiform trichomes, alate
throughout and prominently so in lower half; pinnae (on mature leaves) 20-35 pairs,
rarely more than 5 cm. long, alternate, sessile to short-stalked, spreading at a broad
angle (commonly ca. 90°) from the rachis, oblong or linear-lanceolate to narrow-
ovate, essentially glabrous; ultimate segments linear or narrow-oblong, obtuse
(sometimes retuse to bifid) plane, margins entire; segments and venation anadro-
mous; veins free, pinnately arranged, strongly ascending, once or twice forked,
spurious ones absent; sori often only one per pinnule, and terminating the proximal
veinlet, not or scarcely immersed in the tissue; indusium salverform, bilabiate, the
lips broad and strongly flaring; receptacle filiform, mostly short-exserted; spores
subglobose, tuberculate to spinulate.
This is often confused in herbaria with Trichomanes radicans. In
addition to the characters noted in the key, the two species usually
can be distinguished by blade shape and petiole length. In T. collari-
atum the petiole is rarely more than 3 cm. long and the blade is
gradually reduced toward the base, with lower pinnae one-third to
one-sixth the length of the central pinnae — a basal pinna frequent-
ly crowding or overlapping the rhizome. The petiole of T. radicans is
commonly 5-12 cm. long, with blade scarcely reduced at base; lower
pinnae are about three-fourths the length of the longest central one.
Trichomanes crispum L. Sp. PL 2: 1097. 1753. T. cristatum Kaulf.
Enum. Fil. 265. 1824.
In forests, on tree trunks, wet banks or cliffs, 600-2,500 m.; Alta
Verapaz; BajaVerapaz; Chiquimula; Guatemala; Huehuetenango;
Santa Rosa; Southern Mexico; Nicaragua; Costa Rica; Panama;
West Indies; Trinidad; Colombia; Venezuela; the Guianas; Brazil;
Peru; Bolivia; Paraguay.
Plants epiphytic, saxicolous or terrestrial; rhizome stout, horizontal and short-
creeping or (in ours) ascending to suberect, provided with dark reddish brown, pluri-
cellular trichomes; leaves monomorphous, subdistant to crowded or (in ours) caespi-
tose, 10-40 cm. long and 2-5 cm. broad; petiole 2.5-12 cm. long, wiry, flattened or
(rarely) subtrigonous, gray- or red-brown, nonalate or alate only at base of lamina,
abundantly provided with reddish brown, septate trichomes which have the basal
cell much shorter and broader than the others; lamina linear, lanceolate or narrow-
ovate, dark green or yellowish green, membranaceous, pinnate to pinnatisect,
scarcely or not at all reduced at base, narrowed gradually to a pinnatifid apex, the
tissue subopaque; rachis in upper half of blade alate, wings lacking in lower half or
each one narrower than the rachis, trichomes abundant as on the petiole; pinnae 15
to many pairs, oblong, subopposite to alternate, spreading at nearly right angles
from the rachis, subdistant, contiguous or imbricate, obtuse at apex, entire or sinu-
ate, repand or crisped, with yellow or orange trichomes scattered along the costa,
78 FIELDIANA: BOTANY, VOLUME 39
veins and margin, lacking spurious veinlets; veins free, catadromous, 1- to 2-forked,
pinnately arranged, spreading obliquely from the costa; sori 2 to 10 at apex of each
pinna, fully immersed in the tissue; indusium narrowly cylindric-turbinate or ob-
conic, not bilabiate, the mouth slightly to strongly flaring; receptacle long-exserted;
spores tetrahedral to subglobose, somewhat granular to spinulate.
This is a highly variable, rather widespread, species, belonging to
section Achomanes. When this section is studied in depth, it will
probably be determined that a number of species now named in
tropical America are conspecific with Trichomanes crispum. One of
these which occurs in Guatemala is T. cristatum, originally de-
scribed by Kaulfuss as differing from T. crispum in leaf size and
shape, and in the degree of crisping of pinna margins. However,
these are hardly reliable characters in that they vary so greatly on
plants throughout the range, even among leaves on the same rhi-
zome, and are therefore a questionable basis for taxonomic distinc-
tion. T. crispum also varies in other characters; the indusium may
be narrow and cylindric to obconic and relatively broad, with mouth
slightly to rather strongly flaring; the rhizome may be horizontal
and short-creeping with leaves subdistant to rather crowded, or
ascending and compact with leaves caespitose. All specimens of T.
crispum seen from Guatemala have caespitose leaves.
Trichomanes curtii Rosenst. in Fedde, Repert. Spec. Nov. Regni
Veg. 22: 5. 1925.Didymoglossum curtii (Rosenst.) Pic. Ser. Webbia
28: 469. 1973.
On rocks or tree trunks, sea level to 75 m. ; Izabal. British Hondu-
ras; Nicaragua, south to Colombia.
Plants saxicolous or epiphytic; rhizome long-creeping, filiform, covered with
brown or blackish, mostly flattened trichomes; leaves monomorphous, approximate
or widely spaced, to 3 cm. long and 1.5 cm. broad, subsessile, or with petioles to 1
cm. long; lamina suborbicular, ovate or spathulate, base rounded to decurrent, mar-
gin entire (on juvenile leaves) or shallowly and irregularly lobed to fissile, essentially
plane, yellowish green, a distinct midrib lacking, tissue firm-membranaceous,
scarcely or not at all translucent, glabrous except for the stellate, marginal tri-
chomes; veins free, flabellate, very crowded, repeatedly forked, with spurious vein-
lets scattered parallel to the true veins; sori several to many, partially or scarcely
immersed in the tissue, commonly extending conspicuously from the margin (not
borne between the lobes); indusium cylindric-turbinate, bilabiate, the lips not (or
seldom) flaring, the edges dark-margined; receptacle scarcely to long-exserted;
spores tetrahedral to subglobose, spinulate.
Trichomanes curtii may be further distinguished from the closely
related T. punctatum ssp. sphenoides by the opaqueness of the tis-
sue (caused by the densely crowded veins) and by the more numer-
ous ( several to many) sori. The tissue of the latter is generally trans-
STOLZE: FERNS OF GUATEMALA 79
lucent, with veins much less crowded, and has sori typically num-
bering two or three.
Trichomanes diaphanum HBK. Nov. Gen. & Sp. 1: 25. 1825. Van-
denboschia diaphana (HBK.) Copel. Philipp. J. Sci. 67: 53. 1938.
On trees, moist banks and (rarely) wet rocks, in deep forests, 300-
3,000 m.; Alta Verapaz; Baja Verapaz; Chiquimula; Quezaltenan-
go; San Marcos; Zacapa. British Honduras; Honduras to Panama;
West Indies; Trinidad; Colombia; Venezuela; French Guiana; Bra-
zil; Ecuador; Peru.
Plants terrestrial or, more commonly, epiphytic; rhizome thin, long-creeping,
rather delicate, sparsely to abundantly provided with dark brown, flattened tri-
chomes; leaves monomorphous, commonly distant, mature ones 3-15 (18) cm. long
and 1-4.5 cm. broad, the blades much longer than their petioles; petiole 1.5-4 cm.
long, light or dark brown, flattened, usually alate to base, each wing commonly as
broad or broader than the petiole (often partially broken away on herbarium speci-
mens); lamina ovate, yellow- or gray-green, membranaceous, translucent, tripinnate
or tripinnate-pinnatifid, scarcely reduced at base, narrowed gradually to a pinnatifid
apex; rachis glabrous or obscurely dotted with minute trichomes on abaxial side,
broadly alate throughout with each wing about as broad as the rachis; pinnae 6-12
pairs, alternate, short-stalked, obliquely ascending, lanceolate to ovate, tissue glab-
rous, but axes and veins often with scattered, minute trichomes, costa and costules
alate; ultimate segments linear, 0.6-0.9 mm. broad, emarginate, lateral margins
entire, the tissue essentially plane; veins free, anadromous, spurious ones absent;
sori 1-5(7) per pinna, terminating the proximal veinlets, both stalk and tube flanked
by a rather broad wing of tissue; indusium narrowly funnelform, the tube 2 or more
times as long as broad, the mouth flaring but not bilabiate nor dark-margined; re-
ceptacle usually long-exserted; spores subglobose, surface densely spinulate.
This and Trichomanes hymenophylloides are doubtfully distinct.
See further comments under discussion of the latter.
Trichomanes diversifrons (Bory) Mett. in Sadebeck, Nat. Pflanz.
1 (4): 108. 1899. T. elegans Rudge, Icon. PI. Guian. 24. 1805 (not T.
elegans L.C. Rich. 1792). Hymenostachys diversifrons Bory, Diet.
Class. Hist. Nat. 8: 462. 1825. Feea diversifrons (Bory) Copel.
Philipp. J. Sci. 67: 74. 1938.
On moist, wooded slopes and ravines, and along shaded stream
and river banks, sea level to 500 m.; Alta Verapaz; Izabal. British
Honduras; Honduras; Nicaragua; Costa Rica; Panama; southward
to Brazil and Bolivia.
Plants terrestrial; rhizome stout, erect, provided with lustrous, dark brown to
blackish, terete or somewhat flattened trichomes which are commonly in dense tufts
at rhizome apex; leaves dimorphous, approximate or subcaespitose, the fertile ones
far exceeding the sterile and with much longer petioles; sterile leaf to 28 cm. long
80 FIELDIANA: BOTANY, VOLUME 39
and 7 cm. broad, petiole 2-8 cm. long, stout to wiry, terete or slightly flattened, yel-
lowish to greenish or brown, commonly narrow-alate (at least toward the apex), with
scattered, dark brown trichomes which extend up the abaxial side of the rachis;
sterile lamina lanceolate, yellow- or gray-green, chartaceous, deeply pinnatisect, nar-
rowing at the base and the pinnatifid apex, the rachis frequently prolonged, flagel-
late and proliferous at the tip, tissue glabrous; pinnae numerous, spreading, acute
or subacute, margins serrulate, cut nearly to the rachis, with sinuses narrow and
acute; veins pinnately arranged, oblique, once- or twice-forked, arising from rachis
as well as costae, free or anastomosing toward pinna margin, spurious ones absent;
fertile leaf to 40 cm. long and 0.7 cm. broad, petiole 12-20 cm. long; fertile lamina
simple, linear, with sori arranged in a nearly continuous line along each margin, com-
pletely immersed in tissue at the forks of veins; receptacle scarcely to slightly
exserted; spores subglobose, tuberculate.
The erect, spikelike, fertile blades of this species are borne on elon-
gate petioles far above the cluster of pinnatisect, sterile blades. In
this character alone Trichomanes diversifrons may be easily distin-
guished from all other species in Guatemala.
Trichomanes ekmanii W. Boer, Acta Bot. Neerl. 11: 319. 1962.
In our area known only from Middlesex, British Honduras, grow-
ing on decayed tree trunks in forest, elev. 60 m., W. A. Schipp 324.
Honduras; Costa Rica; Panama; Greater Antilles ; Colombia; Vene-
zuela; Brazil; Peru; Bolivia.
Plants epiphytic; rhizome long-creeping, filiform, sparsely to densely provided
with flattened, brown trichomes which often extend up the petiole; leaves monomor-
phous, rather widely spaced, 0.5-3 cm. long, 0.5-1.5 cm. broad, subsessile or short-
petiolate; petiole 1-5 mm. long, subterete or flattened; lamina obovate or oblong (or
in some immature leaves suborbicular), gray- or yellow-green, irregularly lobed, or
entire toward the rounded or decurrent base, glabrous except along base of midrib
beneath, the tissue firm-membranaceous, somewhat translucent; veins free, pin-
nately branched from a midrib which is distinct at least in the lower half of the blade,
obliquely ascending, once- or twice-forked, their tips connected by a spurious sub-
marginal vein which is continuous around the perimeter of the lamina, spurious
veinlets present, parallel to the true veins; sori few to 9, situated near apex of lam-
ina, fully immersed in the tissue; indusium cylindric-turbinate, not bilabiate, the
mouth somewhat flaring, not dark-margined; receptacle short- to long-exserted;
spores subglobose, coarsely verruca te.
Trichomanes galeottii Fourn. Bull. Soc. Bot. France 15: 148. 1868.
In forests on tree trunks or on wet rocks along streams or water-
falls, sea level to 1,500 m.; Alta Verapaz; Chiquimula; Izabal;
Peten. Cuba; Puerto Rico; Mexico (Oaxaca); British Honduras;
Honduras; Nicaragua; Costa Rica; Colombia (Choco).
Plants epiphytic or occasionally saxicolous; rhizome stout, ascending or suberect,
provided with dark brown, pluricellular trichomes; leaves monomorphous, caespi-
tose, 5-25 cm. long and 1.5-4 cm. broad; petiole 1.5-8 cm. long, wiry, subterete or flat-
STOLZE: FERNS OF GUATEMALA 81
tened, gray- or red-brown, alate halfway or more to the rhizome, abundantly provided
with reddish brown, septate trichomes which have the basal cell much shorter and
broader than the others; lamina linear, lanceolate or narrow-ovate, dark green or
yellow-green, membranaceous, pinnate to pinnatisect, with the basal pair or two of
pinnae somewhat reduced, narrowed gradually to a pinnatifid apex, the tissue sub-
opaque; rachis broadly alate, with each wing as wide or much wider than the rachis,
trichomes abundant as on the petiole; pinnae 10 to many pairs, oblong, subopposite
to alternate, spreading at nearly right angles from the rachis, contiguous to imbri-
cate (rarely subdistant), obtuse at apex, subentire, slightly repand to crisped, with
trichomes (like those of the rachis) scattered along costa, veins and margin; veins
free, catadromous, 1- to 2-forked, pinnately arranged, spreading obliquely from the
costa, lacking spurious veinlets; sori 1 to 4 at the apex of each pinna, fully immersed
in the tissue; indusium narrowly cylindric-turbinate to obconic, slightly or not at all
bilabiate, the mouth slightly or not at all flaring, not dark-margined; receptacle long
exserted; spores subglobose, surface granular to spinulate.
This is very closely related to Trichomanes crispum, but tends to
be smaller in stature, with fewer sori on each pinna, and with a
broad wing well down the petiole. The petiole of T. crispum is at
most alate just below the base of the lamina; but more typically the
petiole and lower rachis are nonalate. A monographic study, com-
paring both species throughout their entire range, may reveal that
T. galeottii is only varietally distinct.
Trichomanes godmanii Hook, in Baker, J. Linn. Soc. Bot. 9: 337.
1866. (Type from Guatemala, without location, Salvin & Godman
s.n. 1862.)
On tree trunks, 50-700 m.; Alta Verapaz; Izabal; Peten. Cuba;
Mexico; British Honduras; Honduras; Costa Rica; Panama.
Plants epiphytic; rhizome long-creeping, slender, densely provided with flattened,
orange or reddish brown trichomes which extend up the petiole; leaves monomor-
phous, rather widely spaced, 1 to 2.5 cm. long and nearly as broad; petiole 1-7 mm.
long, slender, sub terete or flattened; lamina orbicular to suborbicular, gray- or yel-
low-green, subentire, plane or slightly undulate, truncate or somewhat decurrent at
base, glabrous except along base of midrib beneath, the tissue thin and translucent;
veins pinnately branched from a midrib which is distinct in the lower half of the
blade, subflabellately branched toward the apex, obliquely ascending, once to sev-
eral times forked, their tips connected by a spurious, submarginal vein which is con-
tinuous around the perimeter of the lamina, spurious veinlets present, both parallel
and perpendicular to the true veins (thus the venation apparently reticulate); sori
few to 9, situated near apex of lamina, fully immersed in the tissue; indusium cylin-
dric-turbinate, not bilabiate, the mouth commonly flaring, not dark-margined; re-
ceptacle short- to long-exserted; spores subglobose, coarsely verrucate.
Trichomanes hymenoides Hedw. Fil. Gen. & Spec. t.3. 1799. Didy-
moglossum hymenoides (Hedw.) Desv., Mem. Soc. Linn. Paris 6:
330. 1827.
82 FIELDIANA: BOTANY, VOLUME 39
In forests, on tree trunks, often along rivers or waterfalls, 1,200-
1,600 m.; Alta Verapaz; Baja Verapaz; Chiquimula; San Marcos;
Solola. West Indies; Southern Mexico; British Honduras; Costa
Rica; Panama; Trinidad; Venezuela and Colombia southward to
Uruguay and Argentina.
Plants (in our area) epiphytic; rhizome long-creeping, densely provided with dark
brown or blackish, flattened trichomes which extend up the petiole and occasionally
onto the base of the rachis; leaves monomorphous, widely spaced, often forming
dense mats, (0.5) 1-3.5 cm. long and 0.3-1.5 cm. broad, subsessile to short-petiolate;
petiole 0.1-1 cm. long, subterete or flattened, brown; lamina subreniform, broadly
elliptic or obovate, shallowly lobed to pinnatifid (very rarely bipinnatifid), with
sinuses between pinnae acute to broadly rounded, truncate to cuneate and decurrent
at base, margin plane to undulate (rarely crispate), yellowish green, tissue membra-
naceous, translucent, glabrous except for the blackish, marginal trichomes which
are simple or bifid, or sometimes stellate in the sinuses; veins free, catadromous,
pinnately branched from a midrib which is distinct at least in lower half of lamina,
spurious veinlets few between the true veins, not parallel to the margin; sori solitary
to several, near apex of lamina, not or scarcely immersed in the tissue; indusium
cylindric turbinate, bilabiate, the lips dark-margined, commonly flaring, most of
them broader than long; receptacle short- to long-ex serted; spores tetrahedral to
subglobose, tuberculate.
Trichomanes hymenoides is easily confused with T. rep tans, for
all of the characters may vary from leaf to leaf, even among those on
the same rhizome. Thus, to separate the two it is usually necessary
to use the key and other characters in combination. It is conjectural
whether the two taxa merit more than subspecific distinction.
Trichomanes hymenophylloides v.d. Bosch, Ned. Kruidk. Arch. 5
(3): 209. 1859. Vandenboschia hymenophylloides (v.d. Bosch)
Copel. Philipp. J. Sci. 67: 53. 1938.
On trees and moist banks in deep forests, 750-2,300 m. ; Alta Vera-
paz; Huehuetenango; San Marcos. Southern Mexico; Costa Rica;
Panama; West Indies; Surinam to Colombia; Ecuador; Brazil.
Plants terrestrial or, more commonly, epiphytic; rhizome thin, long-creeping, deli-
cate, sparsely to abundantly provided with dark brown, flattened trichomes; leaves
monomorphous, commonly distant, mature ones 2.5-14 cm. long and 1.5-4 cm. broad,
the laminae much longer than their petioles; petiole 0.5-5 cm. long, light or dark
brown, flattened, narrow wings present on the upper portion or lacking; lamina
ovate or obovate, yellow- or gray-green, membranaceous, translucent, bipinnate-pin-
natifid or tripinnate, scarcely reduced at base, narrowed gradually to a pinnatifid
apex; rachis glabrous or obscurely dotted with minute trichomes on the abaxial side,
narrowly alate throughout (each wing much narrower than the rachis); pinnae 4-10
pairs, alternate, short-stalked, obliquely ascending, lanceolate to ovate, tissue glab-
rous but axes and veins often with scattered, minute trichomes, costa and costules
alate; ultimate segments linear, 0.6-0.9 mm. broad, emarginate, lateral margins
STOLZE: FERNS OF GUATEMALA 83
entire, the tissue essentially plane; veins free, anadromous, spurious ones absent;
sori 1-3(4) per pinna, terminating the proximal veinlets, both stalk and tube flanked
by a narrow wing of tissue; indusium narrowly funnelform, the tube 2 or more times
as long as broad, the mouth flaring but not bilabiate nor dark-margined; receptacle
long-exserted, spores tetrahedral, surface baculate to spinulate.
It is questionable whether this fern is distinct from Trichomanes
diaphanum. Other than the amount of tissue flanking the main axis,
the two hardly differ. Perhaps T. hymenophylloides more often has
merely a single sorus per pinna and is slightly less dissected, but
these conditions vary too often to be of real importance. Also, the
amount of tissue along the axes, even in leaves on the same rhizome,
may be variable. This is particularly true of West Indian plants.
Furthermore, the two taxa share approximately the same geograph-
ic distribution.
Trichomanes krausii Hook. & Grev. Icon. Fil. t.149. 1831. Didy-
moglossum krausii (Hook. & Grev.) Presl, Abh. Bohm. Ges. Wiss.
V: 115. 1843. Hemiphlebium krausii (Hook. & Grev.) Prantl,
Unters. Morph. Gefass. I: Hymenoph. 46. 1875.
In dense, wet forests, often along rivers and streams, on tree
trunks and (outside of our area) on wet, shaded rocks, sea level to
1,650 m.; Alta Verapaz; Chiquimula; Izabal; Peten; Solola; Suchi-
tepequez. Florida; Mexico to Panama; West Indies; Trinidad and
the Guianas to Colombia, and southward to Argentina and Para-
guay.
Plants (in our area) epiphytic; rhizome long-creeping; densely provided with dark
brown to blackish, flattened trichomes which extend onto the petiole and lower
rachis; leaves monomorphous, rather widely spaced, often forming dense mats, to 5
cm. long and 3 cm. broad, subsessile to short-petiolate; petiole 0.1-0.5 cm. long, sub-
terete or somewhat flattened, brownish; lamina narrow-ovate to elliptic to obovate,
deeply pinnatifid or bipinnatifid, with sinuses between pinnae or segments broadly
rounded, abruptly narrowed and slightly decurrent at base, margin slightly to
strongly undulate, yellowish green, tissue membranaceous, translucent (often
scarcely so), glabrous except for the blackish trichomes which are bifid along the
margins, but stellate in the sinuses between pinnae or segments; rachis narrow-
alate; pinnae narrow, ascending, lobed to pinnatifid; veins free, pinnately branched
from the costa at an oblique angle, catadromous, spurious veinlets seldom numer-
ous, parallel to true veins and often parallel to and very near the margin; sori on
each pinna solitary or few, terminating the segments, fully immersed except for the
lips, at least a narrow wing of tissue flanking each side of the indusium; indusium
cylindric-turbinate up to the flaring mouth, bilabiate, lips dark-margined (though
sometimes only scarcely so); receptacle short- to long-exserted; spores subglobose,
densely tuberculate.
84 FIELDIANA: BOTANY, VOLUME 39
Although reported in Guatemala only as an epiphyte, Tricho-
manes krausii has been found growing on wet rocks, often in
streams, from Nicaragua to South America, and in the West Indies.
Trichomanes membranaceum L. Sp. PI. 1097. 1753. Lecanium
membranaceum (L.) Presl, Abh. Bohm. Ges. Wiss. V (3): 104. 1843.
Hemiphlebium membranaceum (L.) Prantl, Unters. Morph. Gefass.
I: Hymenoph. 46. 1875. Lecanolepis membranacea (L.) Pic. Ser.
Webbia28:449. 1973.
On tree trunks, mostly in deep shade in forests, sea level to 160
m. ; Izabal. British Honduras; Nicaragua; Costa Rica; Panama;
West Indies; the Guianas to Colombia and southward to Bolivia.
Plants epiphytic; rhizome long-creeping, slender, densely provided with flattened,
dark brown trichomes; leaves monomorphous, rather widely spaced, in ours 1.5-5
cm. long and nearly as broad, subsessile; lamina suborbicular and entire, or spathu-
late and shallowly to deeply incised into irregular lobes, grayish green, often drying
brown, lacking a distinct midrib, tissue membranaceous, opaque, glabrous, but the
margin bearing numerous, distinctive, cordate, suborbicular scales (which are often
deciduous on mature leaves); veins free, numerous, flabellate, repeatedly forked,
spurious veinlets abundant and parallel with the true veins, but a continuous periph-
eral one lacking; sori several to many on vein tips toward apex of lamina, partially
to fully immersed in the tissue; indusium narrow-cylindrical, scarcely bilabiate, the
mouth not flaring nor dark-margined; receptacle short- to long-exserted; spores
tetrahedral to subglobose, coarsely granular.
Trichomanes ovale (Fourn.) W. Boer, Acta Bot. Neerl. 11: 296.
1962. Didymoglossum ovale Fourn. Bull. Soc. Bot. France 19: 240.
1872.
On tree trunks, sea level to 370 m.; Izabal, Peten; Suchitepequez.
Honduras; Panama; Greater Antilles ; Venezuela; Brazil.
Plants epiphytic; rhizome long-creeping, filiform, densely covered with dark
brown or blackish, flattened trichomes; leaves monomorphous, approximate to
widely spaced, often forming dense mats, to 0.8 cm. long and 0.4 cm. broad, with
petiole half the length of the lamina or shorter; lamina suborbicular, ovate or obo-
vate, base rounded to decurrent, margin entire to sinuate or crenate, essentially
plane, yellow- to gray-green, with midrib distinct (in fertile leaves) or lacking, tissue
membranaceous, translucent, glabrous except for the marginal, stellate trichomes;
veins free, flabellate, or pinnately arranged (especially in fertile laminae), once or
several times forked, spurious veinlets scattered parallel with the true veins; sorus
solitary at lamina apex, partially or fully immersed in the tissue; indusium cylindric-
turbinate, bilabiate, the lips flaring and bicolorous, the edges brownish; receptacle
scarcely or not exserted; spores subglobose, coarsely tuberculate.
This is one of the tiniest of all ferns, and is perhaps frequently
overlooked by collectors. Thus the apparent disjunct distribution,
STOLZE: FERNS OF GUATEMALA 85
reflected by current collections, is probably not indicative of the
true range of Trichomanes ovale.
Trichomanes petersii A. Gray, Amer. J. Sci. 2 (15): 326. 1853. T.
schaffneri Schlecht., Linnaea 26: 368. 1853. Microgonium petersii
(A. Gray) v.d. Bosch, Verh. Kon. Ned. Acad. Wetensch. 9 (6): 7.
1861. Hemiphlebium petersii (A. Gray) Prantl, Unters. Morph. Ge-
fass. I: Hymenoph. 46. 1875. Didymoglossum petersii (A. Gray)
Copel. Philipp. J. Sci. 67: 78. 1938.
Growing on tree trunks in forests, 150-1,600 m.; Alta Verapaz;
Baja Verapaz. Mexico; southeastern United States.
Plants (in our area) epiphytic; rhizome long-creeping, slender, densely provided
with flattened, dark brown trichomes; leaves monomorphous, approximate to wide-
ly spaced, often forming dense mats, to 1 cm. long and 0.4 cm. broad, with petiole
short or nearly as long as the lamina; lamina ovate to elliptic-ovate, apex obtuse,
base rounded to cuneate and long-decurrent, margin entire, sinuate or shallowly
lobed, weakly undulate, yellow- to gray-green, with a distinct midrib, tissue mem-
branaceous, scarcely translucent, glabrous except for the blackish, forked or stellate
trichomes scattered along the margin; veins free, pinnately arranged, obliquely as-
cending, simple or once-forked, spurious veinlets scattered parallel with the true
veins; sorus solitary at lamina apex, fully immersed in the tissue; indusium cylin-
dric-turbinate up to the flaring mouth, not bilabiate nor dark-margined; receptacle
scarcely or not exserted; spores subglobose, densely but minutely granular.
Although heretofore reported only as an epiphyte in Mexico and
Guatemala, Trichomanes petersii occasionally has been found grow-
ing on wet rocks in the southeastern United States.
Trichomanes pinnatum Hedw. Fil. Gen. et Sp. t.4. 1799. T. pen-
natum Kaulf. Enum. Fil. 264. 1824. Neuromanes pinnatum (Hedw.)
Trevis. Herb. Crypt. Trevis. 4. 1851.
In shaded ravines and on ridges and slopes in forests, sea level to
500 m.; Alta Verapaz; Izabal; Peten; Zacapa. West Indies; Trini-
dad; Mexico to Panama and southward to Brazil and Bolivia.
Plants terrestrial; rhizome short-creeping to ascending, provided with lustrous,
dark brown to black, subterete trichomes; leaves essentially monomorphous (al-
though fertile ones are often somewhat larger and with longer petioles), approxi-
mate or subcaespitose, to 70 cm. long and 27 cm. broad, lamina as long as, or slightly
longer than the petiole; petiole stout or wiry, terete or rounded-trigonous, gray- or
red-brown, nonalate (or slightly marginate near base of blade), provided with scat-
tered, filiform, septate trichomes; lamina ovate to subdeltoid, commonly yellowish
green, membranaceous, pinnate, not or scarcely reduced at base, the apex terminat-
ing in a conform, elongate pinna, or the rachis often prolonged, flagellate and prolif-
erous at the tip, the tissue translucent, glabrous; rachis nonalate to narrowly alate,
with scattered to abundant trichomes; pinnae 4-16 pairs, linear-lanceolate, suboppo-
86 FIELDIANA: BOTANY, VOLUME 39
site to alternate, distant (or contiguous on sterile blades), sessile or subsessile, acute
to subacute, truncate at base, or adnate on the proximal side, margin serrate to sub-
spinulose, the costae with scattered to abundant, septate trichomes beneath; veins
pinnately arranged, approximate, spreading from costa at a wide angle, catadro-
mous, simple or once-forked, joined or nearly joined with numerous, perpendicular,
spurious veinlets; sori arranged in a nearly uninterrupted line along each margin,
stalked or subsessile on the tips of veins; indusium cylindric-turbinate, not bilabi-
ate, or only scarcely so, the mouth not flaring or only scarcely so; receptacle long-
exserted; spores tetrahedral to subglobose, tuberculate.
Trichomanes polypodioides L. Sp. PL 1098. 1753. T. sinuosum
Rich., Willd. Sp. PL 5: 502. 1810.
In wet forests, pendent from trunks of trees and tree ferns, 300-
1,800m.; AltaVerapaz; BajaVerapaz; Chiquimula; Huehuetenan-
go; Izabal; San Marcos. Southern Mexico; British Honduras; Hon-
duras; Costa Rica; Panama; West Indies; Trinidad; South America
to Paraguay, Brazil and Uruguay.
Plants epiphytic; rhizome long-creeping, slender, provided with scattered brown,
flattened trichomes; leaves monomorphous, widely spaced, 4-15 cm. long and 1-3
cm. broad; petiole short, commonly less than 0.5 cm. long, delicate, subterete, green-
ish or brown; lamina linear-lanceolate, yellowish green, deeply lobed to pinnatifid
from apex to the decurrent base, the tissue translucent and glabrous except at the
margin, the midrib, veins and margins provided with scattered, brown, stellate tri-
chomes; segments (lobes) 10-25 pairs, oblique, obtuse, often as broad as long, with
sinuses acute to rounded, margins subentire, sinuate or broadly crenate; veins free,
pinnately branched, rather widespread, spreading from the midrib at an oblique
angle, catadromous, spurious veinlets lacking; sori one to a few at segment apex,
fully immersed in the tissue at vein tips; indusium salverform, not bilabiate, mouth
somewhat flaring, not dark-margined; receptacle long-exserted; spores subglobose,
coarsely tuberculate.
Trichomanes punctatum Poiret ssp. sphenoides (Kunze) W.Boer,
Acta Bot. Neerl. 11: 301. 1962. T. sphenoides Kunze, Farrnkr. 216.
1840. Didymoglossum sphenoides (Kunze) Presl, Abh. Bohm. Ges.
Wiss. V: 115. 1943. Hemiphlebium sphenoides (Kunze) Prantl,
Unters. Morph. Gefass. I: Hymenoph. 46. 1875.
On tree trunks, in deep shade, 30-150-m.; Alta Verapaz. Greater
Antilles; Costa Rica; Panama; Colombia; Venezuela; Peru; Bolivia.
Plants epiphytic; rhizome long-creeping, filiform, sparsely to densely provided
with dark brown, flattened, trichomes; leaves monomorphous, approximate or
rather widely spaced, to 1.5 cm. long and 1 cm. broad, subsessile or short-petiolate;
lamina suborbicular (especially when immature) to obovate or spathulate, base
rounded to decurrent, margin essentially plane, entire or irregularly lobed, the lobes
often deep and narrow, yellow- to gray-green, a distinct midrib lacking, tissue mem-
branaceous, commonly translucent, glabrous except for the stellate marginal tri-
STOLZE: FERNS OF GUATEMALA 87
chomes; veins free, flabellate, repeatedly forked, with spurious veinlets parallel to
the true veins; sori several, or often solitary, partially immersed in the tissue,
commonly borne between lobes, thus rarely exserted beyond the outline circum-
scribed by the lobe apices; indusium cylindric-turbinate, bilabiate, the lips mostly
wide-flaring, the edges dark-margined; receptacle not or scarcely exserted; spores
subglobose, coarsely tuberculate.
Trichomanes pyxidiferum L. Sp. PL 2: 1098. 1753. Vandenboschia
pyxidifera (L.) Copel. Philipp. J. Sci. 67: 53. 1938.
On trees and moist, clay banks, in forests, 900-3,000 m. ; Alta
Verapaz; Chiquimula; Quezaltenango; Solola; Suchitepequez. West
Indies; Mexico; Honduras; Nicaragua; Costa Rica; Panama;
Colombia to Bolivia and Brazil; Old World.
Plants epiphytic, rarely terrestrial; rhizome slender, long-creeping, rather abun-
dantly provided with blackish, subterete or flattened trichomes which often extend
onto the lower portion of the petiole; leaves monomorphous, distant, but sometimes
thickly matted, 3-10 cm. long and 1.5-5 cm. broad in maturity, the laminae much
longer than their petioles; petiole 1-3 cm. long, subterete or flattened, narrow-alate
partially or wholly to base, green (concolorous with the tissue) or brownish; lamina
lanceolate to ovate, yellow- or gray-green, membranaceous, translucent or scarcely
so, sub-bipinnate to subtripinnate, scarcely or not at all reduced at base, narrowed
gradually to a pinna tif id apex; rachis alate, glabrous, or on the abaxial side with
scattered, minute trichomes; pinnae 4-10 pairs, alternate, subsessile, anadromous,
obliquely ascending, lanceolate to ovate, tissue glabrous, but axes and veins often
with scattered, minute, glandular trichomes; ultimate segments linear, obtuse, com-
monly retuse, margins plane to undulate, tissue (at least in dried specimens) com-
monly with elongate, narrow folds parallel to the veins; veins free, anadromous,
spurious ones absent; sori one to several per pinna, terminating the proximal veins,
immersed in the tissue (or at least a narrow wing of tissue along each side of the
indusium); indusium broadly turbinate, the tube nearly as broad as long, very shal-
lowly or not at all bilabiate, with mouth scarcely flaring, not dark-margined; recep-
tacle commonly long-exserted; spores tetrahedral to subglobose, sparsely to
densely tuberculate.
Trichomanes radicans Sw. J. Bot. (Schrader) 1800 (2): 97. 1801.
T. scandens Hedw. Fil. Gen. Sp. pi. 6. 1799 (not T. scandens L.
1753). T. kunzeanum Hook. Sp. Fil. 1: 127, t.39d. 1844. T. mexi-
canum v.d. Bosch, Ned. Kruidk. Arch. 5: 164. 1861. Vandenboschia
radicans (Sw.) Copel. Philipp. J. Sci. 67: 54. 1938.
In forests and shaded ravines, on tree trunks, wet cliffs and ravine
banks, or on mossy rocks, 900-3,300 m.; Alta Verapaz; Baja Vera-
paz; El Progreso; Guatemala; Huehuetenango; Jalapa; Quezalte-
nango; Sacatepequez ; San Marcos; Santa Rosa; Solola; Suchitepe-
quez; Zacapa. West Indies; Mexico to Panama; Colombia to the
Guianas and southward to Brazil and Paraguay; Old World.
88 FIELDIANA: BOTANY, VOLUME 39
Plants terrestrial to epiphytic; rhizome stout, long-creeping, densely provided
with dark brown, septate trichomes; leaves monomorphous, distant, 22-50 cm. long,
(6) 10-20 cm. broad; petiole (2) 5-12 cm. long, stout, terete at base but becoming flat-
tened or broadly sulcate above on the adaxial side, yellow- to gray-brown, nonalate
or with minute wings near the base of the lamina; lamina lanceolate to ovate, dark
green or gray-green, often drying blackish, firm-membranaceous, opaque, bipinnate-
to tripinnate-pinnatifid, scarcely reduced at base, narrowed gradually to an acumi-
nate, pinnatifid apex; rachis essentially glabrous, narrow-alate (or not at all in the
lower portion); pinnae 12-18 pairs, commonly 4-10 cm. long, alternate, stalked, as-
cending (most at a 45-60° angle from the rachis), lanceolate to narrow-ovate, glab-
rous; ultimate segments linear to narrow-oblong, subacute, obtuse, or retuse to
bifid, margins entire, essentially plane; veins free, anadromous, pinnately arranged,
strongly ascending, several times forked, spurious ones absent; sori commonly ter-
minating the proximal veinlets of the pinnules, not or scarcely immersed in the tis-
sue; indusium narrow-funnelform or narrow-cylindric, scarcely or shallowly bila-
biate, the mouth rarely or never flaring; receptacle usually long-exserted; spores
subglobose, surface tuberculate to low-spinulate.
This occasionally has been confused in herbaria, and by some
authors, with the West Indian Trichomanes scandens L., which it
superficially resembles. Hemsley (Biol. Cent. Am. 3: 604. 1885) lists
Bernoulli & Carlo 369 from Mazatenango, as T. scandens, but I have
not seen this, nor have I found other collections of the species from
Guatemala.
Trichomanes reptans Sw., Prodr.: 136. 1788. T. quercifolium
Hook. & Grev., Icon. Fil. t.115. 1829. T. montanum Hooker, Icon.
Plant, t.187. 1837. Didymoglossum reptans (Sw. ) Presl, Abh.
Bohm. Ges. Wiss. V: 115. 1843. Hemiphlebium reptans (Sw.)
Prantl, Unters. Gefass. I: Hymenoph. 46. 1875.
In forests, on tree trunks and branches, or on wet rocks, often
along rivers or beneath falls, 800-3,200 m.; Alta Verapaz; Baja
Verapaz; El Progreso; El Quiche; Huehuetenango; Jalapa; Quezal-
tenango; San Marcos; Solola; Suchitepequez ; Zacapa. Jamaica;
Hispaniola; Southern Mexico; Honduras; Nicaragua; Costa Rica;
Panama; Colombia and Venezuela southward to Argentina.
Plants saxicolous or epiphytic; rhizome long-creeping; densely provided with
dark brown to blackish, flattened trichomes which extend up the petiole and lower
rachis; leaves monomorphous, rather widely spaced, often forming dense mats, 2-9
cm. long and 1-4 cm. broad, subsessile to short-petiolate; petiole 0.1-2 cm. long, sub-
terete or somewhat flattened, brownish; lamina lanceolate to ovate or elliptic, pin-
natifid to (rarely) bipinnatifid, with sinuses acute to broadly rounded, truncate or
cuneate and somewhat decurrent at base, margin plane or slightly undulate, yellow-
ish green, tissue membranaceous, translucent (often scarcely so), glabrous except
for the blackish, marginal trichomes which are simple or bifid, or stellate in the
sinuses; rachis narrow-alate; pinnae narrow, ascending, sinuate to shallowly lobed;
STOLZE: FERNS OF GUATEMALA 89
veins free, pinnately branched in the pinnae, catadromous, spurious veinlets numer-
ous, only (rarely) a few parallel and close to the margin: sori on each pinna solitary
or a few, commonly on the distal portion of the lamina, not or scarcely immersed in
the tissue; indusium cylindric-turbinate, bilabiate, the lips dark-margined, exten-
ded, sometimes flaring, most of them as long as or longer than broad; receptacle
short- to long-exserted; spores subglobose, tuberculate.
It is often difficult to distinguish between this and Trichomanes
hymenoides. Further discussion may be found under the latter
species.
Trichomanes rigidum Sw., Prodr. 137. 1788 (not T. rigidum
Hedw. 1802). T. mandiocanum Raddi, PI. Bras. 1: 64. 1825. Seleno-
desmium rigidum (Sw.) Copel. Philipp. J. Sci. 67: 81. 1938.
Damp forests and wooded ravines, 950-1,650 m.; Alta Verapaz;
Baja Verapaz; Huehuetenango; Izabal. West Indies; Southern
Mexico; Honduras; Nicaragua; Costa Rica; Panama; southward to
Brazil and Bolivia; Old World tropics.
Plants commonly terrestrial (rarely low epiphytes); rhizome stout, erect or as-
cending, or rarely short-creeping, provided with lustrous, dark brown, acicular,
septate trichomes which are often densely tufted at rhizome apex; leaves to 35 cm.
long, monomorphous, approximate or subcaespitose, with laminae as long as, or
slightly longer than, their petioles; petiole stout, terete or slightly flattened, yellow-
to gray-brown, nonalate (rarely minutely marginate), with widely scattered tri-
chomes; lamina ovate to subdeltoid, olive green or dark green, firm membranaceous,
subopaque, 3- to 4-pinnate, not or scarcely reduced at the base, narrowed gradually
to a pinnatifid apex; rachis glabrous or with scattered septate trichomes, nonalate,
or only scarcely so; pinnae 12-20 pairs, alternate, stalked, spreading or (more com-
monly) ascending, lanceolate or linear-lanceolate, with dark brown glands or glandu-
lar trichomes scattered along the abaxial side of the axes ; ultimate segments linear,
glabrous, often curled on drying, acute or retuse to bifid, margins entire; veins free,
pinnately arranged, strongly ascending, anadromous, spurious ones absent; sori
short-stalked, commonly terminating the proximal veinlets; indusium narrow-
funnelform, slightly marginate on either side, scarcely or shallowly bilabiate, the
mouth rarely flaring, subentire or erose; receptacle much exserted; spores subglo-
bose, densely but minutely tuberculate.
Trichomanes tuerckheimii Christ, Hedwigia44: 361. 1905.
In dense forest, climbing trees, with leaves closely appressed to
the trunks, 50-500 m. ; Alta Verapaz; Izabal. Southern Mexico;
British Honduras; Costa Rica; Panama; Colombia; Venezuela;
British Guiana; Surinam; Peru.
Plants epiphytic; rhizome long-creeping, stout and wiry, sparsely to abundantly
provided with dark brown, flattened trichomes; leaves monomorphous, widely
spaced, to 16 cm. long and 6 cm. broad (rarely to 30 cm. long), subsessile; petiole 1-3
mm. long, terete, greenish, glabrous or with scattered, brownish trichomes, and
90 FIELDIANA: BOTANY, VOLUME 39
with a small, dense tuft of blackish trichomes at the base; lamina broadly oblong,
dark green to yellow-green, often drying blackish, pinnatisect, truncate at base,
tapering to a gradually pinnatifid apex, the tissue membranaceous and translucent,
the rachis, costa, and often the veins and margins rusty tomentose on the abaxial
side; pinnae commonly 4-18 pairs, adnate and essentially perpendicular to the
rachis, crowded or imbricate, with apices broadly rounded or often retuse, margins
plane, entire to crenate; veins pinnate, free, simple, oblique, catadromous, com-
monly terminating short of the margin, spurious veinlets few, extremely short,
parallel to main veins, predominantly situated near the margin; sori few to several,
short-stalked, borne on vein tips along the lateral margins of pinnae; indusium nar-
row-cylindric or narrow-turbinate, not or scarcely bilabiate, mouth not flaring;
receptacle commonly long-exserted ; spores tetrahedral, coarsely and deeply
spinose.
Trichomanes tuerckheimii has perhaps the most peculiar habit of
any of the Guatemalan species in the genus. It is found climbing
trunks of trees with its leaves tightly appressed to the bark, to
which they adhere by means of rust-colored, prehensile trichomes
abundant along the abaxial side of veins and midribs and often
along the margins.
PLAGIOGYRIACEAE
Plants coarse, terrestrial, glabrous (at least at maturity); rhizome stout, erect,
dictyostelic, woody, provided with hard, fibrous roots, old leaf bases crowded and
persistent; leaves to 1 m. long, circinate in vernation, dimorphous, fertile ones with
longer petioles and narrower, constricted pinnae; petiole subtriangular, sulcate,
commonly flattened and slightly expanded at the sheathlike base, not articulate at
the caudex, with a double row of pneumatophores at base, which are generally with-
ered or worn away on mature plants; laminae pinnate or pinnatifid, herbaceous to
subcoriaceous, lanceolate or narrowly ovate; sterile pinnae entire or (in ours) serru-
late to biserrate; veins free, simple or variously forked; fertile pinnae subentire to
erose, with margins at first reflexed and partially protecting the developing sporan-
gia, the tissue later spreading, or sometimes so strongly retroflexed that both edges
touch on the adaxial side; sporangia exindusiate, covering the abaxial side of fertile
pinnae on both sides of the costa, not arranged in definite sori, each long-stalked (4-6
rows of cells), with an oblique annulus uninterrupted by the stalk, dehiscing lateral-
ly; spores trilete, smooth to tuberculate, 48 in each sporangium.
The family consists of one genus, with about 30 closely related
species which are essentially confined to wet, montane areas, in
Asia and tropical America. The dimorphous leaves are borne on
erect rhizomes approximately 1.5 cm. in diameter, which often
appear 3-4 times as thick due to the crowded, persistent, petiole
bases. Thus they sometimes have the aspect of diminutive tree
ferns. The sterile leaves arise somewhat obliquely from the rhizome
in a circular pattern, from the center of which spring several fertile
STOLZE: FERNS OF GUATEMALA 91
leaves. The habit of the plant resembles that of Osmunda cinna-
momea.
PLAGIOGYRIA (Kunze) Mettenius
References: G. Mettenius, Ueber einige farngattungen, 2: Plagio-
gyria, Abh. Senckenberg. Naturf. Ges., Frankfort, 1858. E. B. Cope-
land, The fern genus Plagiogyria, Philipp. J. Sci. 38: 377-417. 1929.
D. B. Lellinger, The American species of Plagiogyria sect. Cari-
natae, Amer. Frn. J. 61: 110-118. 1971.
Characters are those of the family. The following two species
occur in Guatemala.
a. Sterile pinnae 4-6 mm. wide; veins solitary, simple or once forked at or above the
base P. pectinata.
a. Sterile pinnae 8-9 mm. wide; veins (most of them) paired, or forked from the base,
with branches again forked above the base (or some veins solitary and once
forked) P. semicordata.
Plagiogyria pectinata (Liebm.) Lellinger, Amer. Fern J. 61: 115.
1971. Lomaria pectinata Liebm. Kongel. Danske Vidensk. Selsk.
Skr. V. 1: 233 (seors. 81). 1849. (type from Oaxaca, Mexico, Lieb-
mann). L. arguta Fee, Mem. Fam. Foug. 8: 70. 1857 (type from
Veracruz, Mexico, Schaffner 98). Plagiogyria aequidentata Fourn.
Mex. PI. 1: 133. 1872 (type from Veracruz, Mexico, F. Miiller 723).
P. arguta (Fee) Copel. Philipp. J. Sci. 38: 407. 1929.
In moist, mountain forests, at about 3,100 m.; Huehuetenango.
Mexico.
Rachis and petiole stramineous, the latter dark brown at base; sterile leaf to 58
cm. long and 10 cm. broad, with petiole 4-8 cm. long; sterile lamina deeply pinnati-
sect, lanceolate, tapering to a pinnatifid apex, commonly narrowed at the base (in
ours reduced almost to auricles); pinnae of sterile leaf produced at nearly right
angles to the rachis, 4-6 mm. wide (in ours), chartaceous to subcoriaceous, linear,
often subfalcate, acute, margins subentire to denticulate and slightly revolute,
somewhat dilated at base, with sinuses narrowly to broadly rounded; veins solitary,
simple or once-forked at or above the base; fertile leaf pinnate, longer and narrower
than the sterile leaf, to 75 cm. long and 7 cm. broad, with petiole 12-20 cm. long, the
pinnae about 2 mm. broad, distant, spreading or ascending, often gently curved
toward lamina apex, scarcely or not dilated at base, margins strongly retroflexed at
maturity, often touching each other across the adaxial side; spores essentially
smooth-surfaced, but also provided with scattered to abundant, irregularly disposed
tubercles.
In Guatemala and southern Mexico, the sterile laminae of this
species are gradually but strongly reduced at the base, with the low-
FlG. 14. Plagiogyria. a-c, P. semicordata: a, habit (with fertile leaf), X Vi; b, sterile
leaf, X 1/2 ; c, portion of pinna, X 3 ; d, P. pectinata, portion of pinna, X 3.
92
STOLZE: FERNS OF GUATEMALA 93
ermost pinnae often only 1 cm. long. However, there are plants in
Mexico, north of Oaxaca, with the lowermost pinnae reduced only to
about half the length of the longest pinnae. The characters of pinna
width, margin, and venation, which are used to distinguish this
from P. semicordata, are also found to be somewhat variable when
comparing large numbers of specimens from tropical America. More
exhaustive field studies and greater collecting throughout the range
may well prove the two to be conspecific.
Plagiogyria semicordata (Presl) Christ, Farnkr. 176. 1897. Loma-
ridium semicordatum Presl, Epim. Bot. 155. 1849. Plagiogyria
biserrata Mett. Abh. Senckenberg. Naturf. Ges. II. Plag. 1858. P.
obtusa Copel. Philipp. J. Sci. 38: 413. 1929.
Cool, wet, mountain forests, 2, 200-2, 800m.; Huehuetenango; San
Marcos. Cuba; Jamaica; Mexico; Costa Rica; Panama; Colombia
and Venezuela to Bolivia.
Rachis and petiole stramineous, the latter dark brown at base; sterile leaf to 80
cm. long and 14 cm. broad, with petiole to 10 cm. long; sterile lamina deeply pinnati-
sect to pinnate, lanceolate, tapering to a pinnatifid apex, commonly reduced gradu-
ally but strongly to the base; pinnae of sterile leaf produced at nearly right angles to
the rachis, 8-9 mm. wide (in ours), chartaceous to subcoriaceous, linear-lanceolate,
straight or somewhat subfalcate, acute, margins mostly biserrate, plane or slightly
revolute, often dilated at base on the distal side, with sinuses (on pinnatisect por-
tions) broadly rounded, or pinna bases subdistant, with little or no connecting tissue
along the rachis; veins mostly paired, or forked from the base, with branches again
forked above the base, or, less frequently, some veins solitary and once- forked; fer-
tile leaf pinnate, longer and narrower than the sterile leaf, to 90 cm. long and 10 cm.
broad, with petiole 16-24 cm. long, the pinnae about 2 mm. broad, spreading or
slightly ascending and curved, scarcely or not dilated at base, margins strongly
retroflexed at maturity, often touching each other across the adaxial side; spores
essentially smooth-surfaced, but also provided with scattered to abundant, irregu-
larly disposed tubercles.
DICKSONIACEAE
Large, terrestrial plants with horizontal, ascending, or (most commonly) erect
stems, which are often massive and trunklike and sometimes provided with a dense
covering of adventitious roots; leaves ample to huge, to several meters long, circi-
nate in vernation, monomorphous, or dimorphous (fertile lamina or parts of the lam-
ina slightly to strongly constricted); petiole stout, glabrescent to variously pubes-
cent, at the base often densely lanate, not articulate at the stem; lamina 2- to 4-pin-
nate, chartaceous to subcoriaceous; indument consisting of unbranched trichomes,
scales lacking; veins free, simple to several times forked; sori terminal on the veins,
marginal, cup-shaped or (in ours) bivalvate, the outer valve of the indusium often a
scarcely modified portion of the segment margin; sporangia short- to (in ours) long-
94 FIELDIANA: BOTANY, VOLUME 39
stalked, the annulus oblique, essentially complete, slightly or not at all interrupted
by the stalk, dehiscing horizontally; spores 48-64 in each sporangium, trilete, tetra-
hedral, the exine smooth to granulate or verruca te.
The five genera of the Dicksoniaceae have been considered by
some authors to be part of the Cyatheaceae. Others have grouped
them with a larger number of genera in the Pteridaceae, a somewhat
conglomerate family. Whichever concept one prefers, the more
restrictive family circumscribed here certainly proves to be a quite
natural one, easily defined and recognized.
There are about 35 species in the family, including the monotypic
genera, Thyrsopteris (Juan Fernandez) and Cystodium (Borneo and
the Philippines). The other three genera are represented in Guate-
mala.
a. Lamina 4-pinnate or more (in ours); costa and costules deeply sulcate on the
adaxial side, the furrow flanked by prominent ribs, which are decurrent onto the
axis of the next order Culcita.
a. Lamina tripinnate or subtripinnate; costa and costule raised or only slightly fur-
rowed, the ribs (if any) not decurrent onto the axis of the next order,
b. Outer valve of the indusium similar to the inner, both cartilaginous; trichomes
on costae and costules (in ours) lax and filiform, whitish to stramineous;
paraphyses sparse, minute, concealed by the sporangia Cibotium.
b. Outer valve of the indusium consisting of a scarcely altered, revolute crenation
of the segment; trichomes on adaxial side of costae and costules rigid and
stout, dark brown (in ours); paraphyses abundant, as long or longer than the
sporangia Dicksonia.
CIBOTIUM Kaulfuss
Reference: W. R. Maxon, The American species of Cibotium,
Contr. U.S. Natl. Herb. 16: 54-58. 1912.
Stem stout, prostrate to ascending (in ours) or erect and massive; leaves huge, to
several meters long, decompound; petiole smooth, essentially glabrous except for a
dense matting of reddish trichomes at base; lamina commonly tripinnate, essential-
ly monomorphous, often glaucous abaxially, pinnae stalked, broadly lanceolate;
costae and costules raised or, rarely, shallow-furrowed, the raised ribs (if any) not
decurrent onto the axis of the next order, variously pubescent, but in ours the tri-
chomes delicate, filiform, lax, spreading or somewhat appressed; ultimate segments
more or less symmetric, straight or subfalcate, spreading at broad, nearly right,
angles from the costule; veins free, simple or once or several times forked; sori at
the ends of veins, attached at the margin of the segment; indusium bivalvate, both
valves similar in color and texture, spreading widely apart at maturity; sporangia
long-stalked, the annulus oblique, not interrupted by the stalk; paraphyses various,
but in ours minute and concealed by the sporangia; spores tetrahedral, trilete.
FIG. 15. Cibotium regale, a, habit, showing part of lamina and petiole base, X Vi;
b, secondary and tertiary axes, adaxial side, X 4; c, ultimate segment, showing
position of sori, X 5.
95
96 FIELDIANA: BOTANY, VOLUME 39
The genus contains eight or ten species, all but two in the Old
World, a few with arborescent habit. A single species is recognized
in Guatemala.
Cibotium regale Versch. & Lem. 111. Hort. 15: t. 548. 1868. C.
wendlandii Kuhn, Linnaea 36: 151. 1869 (type from Guatemala,
location unknown, Wendland s.n.). C. guatemalense Kuhn, torn. cit.
152 (type from Guatemala, location unknown, Wendland s.n.).
Dicksonia wendlandii (Kuhn) Bak., in Hook. & Bak. Syn. Fil. ed. 2:
460. 1874. D. guatemalense (Kuhn) Bak., torn. cit. 461. D. regalis
(Versch. & Lem.) Bak. loc. cit.
Forested slopes or ravines, 750-2,850 m.; Alta Verapaz; Chimal-
tenango; Chiquimula; Guatemala; Quezaltenango; San Marcos;
Solola; Zacapa. Chiapas (original description based on cultivated
specimens collected in Mexico by Ghiesbreght; type probably
Ghiesbreght 351 ) ; Honduras.
Plants acaulescent or with stems erect, stout, to 1 m. tall; leaves to 3 m. long, sub-
coriaceous; petiole smooth, drying stramineous to light brown (sometimes darker at
base), essentially glabrous throughout, but with a dense matting of golden brown
trichomes at base; lamina broadly ovate, tripinnate or subtripinnate; rachis
smooth, commonly drying stramineous, essentially glabrous; pinnae stalked (1-6
cm.), lanceolate, acute to long-acuminate; costae and costules sparsely to abun-
dantly villous, or glabrescent, the trichomes simple, delicate, appressed to laxly
spreading, whitish to stramineous, readily deciduous; pinnules sessile or short-
stalked, linear-lanceolate, attenuate to subcaudate, cut nearly to the costule
throughout, or fully so near the base; ultimate segments dark green adaxially
(drying brownish); lighter abaxially and often glaucous, crenate-serrate, the mar-
gins revolute, apex acute (or appearing acuminate or cuspidate due to the inrolled
margins), the midrib and veins sparsely to densely villous abaxially, commonly
glabrous adaxially; veins simple or once- or twice-forked, 6-11 (15) pairs per seg-
ment; sori crowded along the margins, parallel to the midrib or often turned oblique-
ly; paraphyses sparse, minute, concealed by the sporangia.
Three species of Cibotium which traditionally have been recog-
nized in Central America are C. guatemalense, C. regale, and C.
wendlandii. Maxon and earlier authors based their segregation on
various combinations of characters: segments aristate us. acumi-
nate, sori parallel vs. oblique to the midrib, lamina glabrous and
glaucous vs. densely villous and not glaucous. These features, if
consistent, might constitute a valid basis for segregation, but
examination of a large number of specimens from Mexico, Guate-
mala, and Honduras reveals such characters to be highly variable.
Even in individual specimens, the traditional distinctions do not
hold up well. Both the type collection of C. regale and a specimen
(J. D. Smith 2423) which Maxon cited as C. wendlandii are obvious-
STOLZE: FERNS OF GUATEMALA 97
ly villous, yet they are also glaucous. In both, the sori are mostly
parallel to the midrib, yet often they are arranged in a somewhat
oblique pattern. Finally, both specimens show some variation in the
shape of the apices of ultimate segments: on parts of the leaf the
segments may be merely acute, yet on other parts the segments are
quite acuminate (though never truly aristate, as Maxon main-
tained).
Some of the variability of the above characters may be attributed
to the maturation of the leaf, or perhaps to the amount of moisture
available. For example, it appears that both the trichomes and
"bloom" on the abaxial surface often tend to disappear as the plant
ages; and perhaps in age (or desiccation) the segment margins be-
come revolute. The sharp inrolling of the segment margin may turn
the sori obliquely and, near the apex, may often cause an acute seg-
ment to appear acuminate.
Called "lanilla" in Honduras, this fern is sometimes used for
stuffing pillows (fide Standley).
CULCITA Presl
Reference: W. R. Maxon, The genus Culcita, J. Wash. Acad. Sci.
12: 454-460. 1922.
Stem stout, prostrate to ascending, plants not or scarcely arborescent; leaves
broad, commonly 1 m. long, decompound; petiole smooth, sparsely villous above, to
very densely lanate at base; lamina 3- to (in ours) nearly 5-pinnate, essentially mon-
omorphous, glabrous except for the axes and veins; pinnae broadly ovate to subtri-
angular; costae and costules deeply sulcate on the adaxial side, the furrow flanked
by prominent ribs which are decurrent onto the axis of the next order, glabrescent or
sparsely provided with lax, mostly spreading trichomes; ultimate segments some-
what oblique, asymmetrical, and inequilateral at base, shallowly lobed to deeply
dissected; veins free, simple or forked; sori terminating the veins; indusium subglo-
bose, bivalvate, the outer valve consisting of a scarcely altered, revolute portion of
the segment margin, the inner one differing slightly in texture; sporangia long-
stalked, the annulus somewhat oblique and slightly interrupted by the stalk; para-
physes filamentous, numerous, as long or longer than the sporangia; spores tetra-
hedral, trilete.
Culcita may be readily distinguished from the other two genera in
Guatemala, as indicated in the key. However, it is apparently more
closely allied wiihDicksonia, with which it shares a number of simi-
lar, significant characters, notably the paraphyses, and the outer
valve of the indusium.
There are eight species in the genus, several occurring on Pacific
Islands, from Samoa to Australia and north to Taiwan. C. macro-
FIG. 16. Culcita coniifotta. a, habit, showing portion of lamina and petiole base,
X !/2; b, portion of lamina, adaxial side, showing ribs continuous along axes, X SVi;
c, penultimate segment, showing position of sori, X 3.
98
STOLZE: FERNS OF GUATEMALA 99
carpa is found in the Madeira Islands and the Azores. C. conii folia is
the only species in the neotropics.
Culcita coniifolia (Hook.) Maxon, Annual Rep. Board Regents
Smithson. Inst. 1911: 488, t. 13c. 1912. Dicksonia coniifolia Hook.
Sp. Fil. 1: 70, t. 24A. 1844. D. martiana Kl. in Hook. Sp. Fil. 1: 70,
t. 24B. 1844. Culcita schlimensis Fee, Mem. Fam. Foug. 10: 47.
1865. Balantium martianum (Kl. ) Fee, Vase. Crypt. Bres. 1: 155.
1869. B. coniifolium (Hook.) J. Sm. Hist. Fil. 258. 1875.
In dense to open forest, or in bamboo or shrub thickets, 2,000-
3,000 m.; Huehuetenango; Jalapa; El Progreso. Greater Antilles;
southern Mexico; Honduras; El Salvador; Costa Rica; Panama;
Colombia to Surinam, southward to Brazil and Argentina.
Rhizome stout, prostrate or ascending; leaves erect, to 3 m. long; petiole drying
light brown, dark brown (or atropurpureus at base), unarmed, provided with fili-
form, orange or light brown trichomes, these scattered distally, but densely matted
at base; lamina broadly triangular, subcoriaceous, to 4- (to nearly 5-) pinnate, mono-
morphous, or fertile segments slightly constricted; rachis smooth, light or dark
brown, sparsely provided with light brown or nearly colorless trichomes, these most-
ly lax and twisted; pinnae stalked (to 6 cm.) broadly subtriangular, glabrous except
for a few lax trichomes along the costae and costules, the latter adaxially sulcate (as
the rachis); quaternary segments oblique, somewhat asymmetric, shallowly to
deeply lobed or cut almost entirely to the base, the ultimate segments simple and
acute, or bifid; sori terminating the veins, at the tip of a simple segment or on the
acroscopic lobe of a bifid segment.
DICKSONIA L'Heritier
Reference: W. R. Maxon, The North American tree ferns of the
genus Dicksonia, Contr. U.S. Natl. Herb. 17: 153-156. 1913.
Stem erect, commonly massive and trunklike, rarely prostrate to ascending, to 6
m. tall and 40 cm. in diameter (including the dense covering of adventitious roots);
leaves huge, to several meters long, decompound; petiole smooth, or slightly muri-
cate with the persistent bases of trichomes, glabrous, or sparsely villous, but at base
provided with a dense matting of long, golden brown or red-brown trichomes;
lamina bipinnate or (as in ours) tripinnate, often glaucous abaxially, slightly to
strongly dimorphous, the fertile segments somewhat constricted; pinnae commonly
lanceolate, subsessile or stalked; costae and costules raised or, rarely, shallow-
channeled, the ribs (if any) not decurrent onto the axis of the next order, variously
pubescent abaxially, but provided adaxially with a dense covering of stout, light or
(in ours) dark brown, antrorse trichomes; ultimate segments more or less symmet-
ric, straight or subfalcate, spreading at a broad angle from the costule; veins free,
simple to several times forked; sori terminating the veins; indusium bivalvate, the
outer valve consisting of a scarcely altered, revolute portion of the segment margin;
the inner one differing somewhat in texture and color; sporangia long-stalked, the
100 FIELDIANA: BOTANY, VOLUME 39
annulus oblique, not interrupted by the stalk; paraphyses (at least in ours) filamen-
tous, numerous, as long or longer than the sporangia; spores tetrahedral, trilete.
Most species of Dicksonia are true tree ferns, i.e., the stems are
massive, often lofty trunks, and the leaves are huge and spreading
and commonly highly dissected. The habit is shared with a number
of genera in the Cyatheaceae, notably Alsophila, Cyathea, Nephe-
lea, Sphaeropteris, and Trichipteris. Perhaps coincidentally, per-
haps significantly, Dicksonia shares another important character
with these genera: the adaxial side of costae and costules is abun-
dantly to densely covered with stout, antrorse, pluricellular tri-
chomes.
There are 18-20 species in the genus, about a third occurring in
Central and South America. D. arborescens L'Herit. is found on St.
Helena, and the rest are distributed from Australia and New Zea-
land to the Philippines. One species is recognized in Guatemala.
Dicksonia gigantea Karst. Fl. Columb. 2: 177. 1869. D. ghies-
breghtii Maxon, Contr. U.S. Natl. Herb. 17: 155. 1913 (type from
Chiapas, Mexico, Ghiesbreght 353).
Arborescent ferns occurring in deep, wet forests, on slopes or in
ravines, 1,750-3,200 m.; Baja Verapaz; Quezaltenango; San Mar-
cos; Zacapa. Southern Mexico; Honduras and El Salvador to
Colombia.
Stem to 6 m. tall and 20 cm. thick (including a dense covering of adventitious
roots); leaves to 3 m. long, spreading to ascending; petiole commonly 15-30 cm.
long, drying dark brown or stramineous, smooth, or the proximal portion slightly
muricate with the subpersistent bases of trichomes, lightly pubescent, or at base
provided with a thick matting of golden brown trichomes; lamina tripinnate or sub-
tripinnate, broadly lanceolate, reduced at apex and base, subcoriaceous, fertile ones
often somewhat constricted; rachis smooth or slightly muricate, commonly drying
stramineous, lightly pubescent as on the petiole; pinnae subsessile or short-stalked,
acute to acuminate; costae and costules on abaxial side sparsely to abundantly pro-
vided with yellow to brownish trichomes, these mostly rigid, antrorse or spreading,
on the adaxial side densely covered with mostly dark brown trichomes, these ant-
rorse and rigid; ultimate segments crenate-serrate or deeply lobed, glabrous except
for the axes, not or slightly glaucous, slightly to strongly revolute, with apices acute
to nearly cuspidate; veins 5-7 (8) pairs per segment, simple to once-forked, or in the
deeply lobed segments 2- to 3-forked; sori subdistant along the margins, sometimes
crowded, but rarely touching; paraphyses abundant, lax, spreading, as long or
longer than the sporangia.
There seems no reason to maintain this and D. ghiesbreghtii as
separate species. Maxon distinguished the two on rather tenuous
grounds: that the latter had one or two more veins per segment, and
an extra fork in the veins. With D. gigantea perhaps may also be
FIG. 17. Dicksonia gigantea. a, habit, showing portion of lamina and petiole base,
X Vs; b, portion of lamina, adaxial side, X 21/i; c, ultimate segment, showing position
of sori, X 6.
101
102 FIELDIANA: BOTANY, VOLUME 39
included D. karsteniana of Costa Rica and Colombia, which is said
to differ by similar characters.
CYATHEACEAE
Reference: R. M. Tryon, The classification of the Cyatheaceae,
Contr. Gray Herb. 200: 1-53. 1970.
Large, terrestrial plants with horizontal or (most commonly) erect stems which
are typically massive and trunklike and sometimes provided with a dense covering
of adventitious roots; leaves small to usually huge, to several meters long, circinate
in vernation, essentially monomorphous, borne in a crown at stem apex (or spaced
along a creeping rhizome in Metaxya), petioles imperfectly deciduous and breaking
away irregularly (although in some species the older petiole bases eventually falling
away to reveal distinct, regularly spaced scars); lamina 1 -pinnate to decompound
(simple in one species of Ceylon); indument consisting of scales or trichomes, or
both; veins free or, less commonly, the basal ones merging to form costal areoles;
sori abaxial on the veins, round, the indusia sphaeropteroid to hemitelioid or lack-
ing; sporangia numerous, ovoid to pyriform, each with an essentially complete,
oblique annulus which is uninterrupted by the stalk, dehiscing horizontally; spores
16 to 64 in each sporangium, trilete, tetrahedral to subglobose, smooth or variously
sculptured, with or without perine.
Authors have disagreed on the limits of the family. There is
strong evidence that Lophosoria and Metaxya are more closely
allied to the non-squamate Dicksoniaceae, or might even be more
suitably placed in a separate family, Protocyatheaceae. Still others
include in the Cyatheaceae not only Lophosoria, Metaxya, and the
Dicksoniaceae, butDennstaedtia, Hypolepis, and other genera. Our
treatment follows Tryon's recent classification, which includes
Lophosoria and Metaxya with the six squamate genera. Thus delim-
ited, there are approximately 440 species in the family, occurring in
both the neotropics and paleotropics.
Terminology of indusial characteristics is covered in detail in the
discussion of Cyathea.
a. Stem and petioles provided with trichomes, scales absent.
b. Lamina decompound; segment margins scarcely modified Lophosoria.
b. Lamina simply pinnate; segment margins cartilaginous Metaxya.
a. Stem and petioles provided with scales, trichomes may also be present.
c. Pinnae entire to shallowly lobed; veins forming costal areoles . . . Cnemidaria.
c. Pinnae deeply incised or decompound; veins free.
d. Some, usually many, scales of the petiole and other axes bearing dark-
colored apical (and often marginal) setae.
e. Petiole, rachis, and usually the costa blackish Alsophila.
e. Petiole, rachis, and costa brown to stramineous.
STOLZE: FERNS OF GUATEMALA 103
f. Petiole with stout, sharp, squaminate, blackish spines; petiole scales
structurally marginate (marginal cells differentiated from central
ones in shape, size and orientation) Nephelea.
f. Petiole lacking stout, squaminate spines (though sometimes muri-
cate with the persistent bases of scales); petiole scales structurally
conform (marginal and central cells similar in size, shape and ori-
entation) Sphaeropteris.
d. Scales of the petiole and other axes lacking dark-colored setae (though
often with apices filiform and margins erose to fimbriate).
g. Indusium present, hemitelioid, cyathiform or globose (sides often lacer-
ate and fugacious at maturity) Cyathea.
g. Indusium absent Trichipteris.
ALSOPHILA R. Brown
Stem erect, commonly massive and trunklike (in ours to 3 m. tall); leaves to sev-
eral meters long; petiole variously colored but commonly (as in ours) atropurpureus
to blackish, smooth, or slightly muricate with the persistent bases of scales, or
rarely with stout, nonsquaminate, conical spines at base (the spines not bearing
scalelike remnants at their margins or bases); petiole scales mostly appressed, deep
amber to dark brown, often with paler margins, the marginal cells differentiated
from those of the central portion in shape, size and orientation, some or all scales
bearing dark-colored, apical and (rarely) marginal setae; several pairs of aphlebiae
occurring near the petiole base, these (at least in ours) highly dissected, 3-18 cm.
long, the segments often ephemeral, on dried specimens sometimes completely
deciduous from the axes, thus the latter appearing skeletal; lamina bipinnate to tri-
pinnate-pinnatifid; rachis colored as the petiole, glabrous abaxially or with a few
scattered, brown scales, appressed-hairy adaxially; pinnae short- to long-stalked,
the costae and costules bearing abundant, appressed, minute, pluricellular tri-
chomes on the adaxial side, the costae with scattered to abundant scales on the
abaxial side, those of costules and midribs often dark-setose and bullate; veins free,
simple or once-forked, on lobed segments the basal veins reaching the margin at a
point above the base of the sinus; sori disposed on the veins close to the midribs; in-
dusia scalelike or sphaeropteroid or, more commonly (as in ours), lacking; receptacle
commonly elevated and subglobose, and with paraphyses shorter than the sporan-
gia; spores 16 per sporangium (except 64 in A. salvinii), exine with or without aper-
tures of various sizes, but commonly lacking a single large, equatorial pore near the
center of each face, perine provided with long, irregular ridges.
InAlsophila, the dark setae of petiole scales may be often lacking
on herbarium specimens, due to the breaking away of scale tips.
Setae are more easily observed on costules and midribs on the abax-
ial side of pinnules, which are better protected.
Gerald Gastony has recently studied the spores of several genera
of the Cyatheaceae and reports ( in litt. ) that one paleotropical spe-
cies of Alsophila has porate exine indistinguishable from that in
Cnemidaria, i.e., with a single large equatorial pore near the center
of each face. He also states that A. salvinii, with 64 spores per spor-
FIG. 18. Alsophila salvinii. a, habit, pinna, and petiole base, X Vt\ b, ultimate seg-
ments, X 2 !/2 ; c, portion of segment, with bullate scales, X 5 ; d, petiole scale, X 8.
104
STOLZE: FERNS OF GUATEMALA 105
angium, is atypical in a genus which commonly has 16-spored
sporangia.
Alsophila may be further distinguished from other genera by the
presence of aphlebiae (often greatly dissected) at the very base of
the petiole. These are pinna-like but quite distant from the lower-
most pinnae, and often very different in character. A few species of
Nephelea have subaphlebioid basal pinnae, but these do not occur at
the base of the petiole.
Affinities of the genus are with Nephelea, under which see further
discussion. Alsophila contains over 200 species, most of which occur
in the paleotropics. Approximately 12 species are found in the neo-
tropics, but only one in Central America.
Alsophila salvinii Hook, in Hook. & Bak. Syn. Fil. 36. 1866.
Ravines and slopes of cloud forests, 1,300-2,600 m.; Alta Verapaz;
Baja Verapaz (type from Chilasco, Salvin & Godman s.n.); Chiqui-
mula; Huehuetenango; Jalapa; El Quiche; Zacapa. Mexico (Chia-
pas); Honduras; El Salvador. Chispon (Zacapa, fide Steyermark).
Stem to 3 m. tall; leaves to 2 m. long; petiole to 1 m. long, atropurpureus or black,
smooth, slightly muricate due to persistent bases of scales, rarely with a few spines
near the base; lamina tripinnate or tripinnate-pinnatifid, dark green above and
lighter (often appearing silvery on live plants) beneath; rachis atropurpureus to
blackish, essentially smooth, nonalate; pinnae broadly lanceolate, costae dark
brown to blackish, abundantly provided with minute, dark brown, appressed, pluri-
cellular trichomes on the adaxial side of costae and costules, and with dark brown,
appressed, often setose scales scattered on costae abaxially, abundant and bullate
or sub-bullate on costules and midribs; ultimate segments crenate, or lobed halfway
to the midrib; veins simple to once- forked; sori inframedial between midrib and mar-
gin; indusium lacking; spores 64 per sporangium, tetrahedral, without apertures,
surface essentially smooth.
Alsophila salvinii may be readily distinguished from all other spe-
cies of the family in Guatemala by the uniformly blackish petiole,
rachis, and (usually) costae. Other species may be deep brown to
blackish (if at all) only toward the base of the petiole.
CNEMIDARIA Presl
References: W. R. Maxon, The North American species of Hemi-
telia, subgenus Cnemidaria, Contr. U.S. Natl. Herb. 16: 25-49. 1912.
R. G. Stolze, A taxonomic revision of the genus Cnemidaria (Cya-
theaceae), Fieldiana Bot. 37: 1-98. 1974.
106 FIELDIANA: BOTANY, VOLUME 39
Stem ascending to erect, rudimentary to 1.5 m. long (or, in some species, rarely to
3.5 m. long); leaves to 3.5 m. long and 1.5 m. broad, lanceolate to ovate-oblong; peti-
ole smooth to muricate, or provided with stout, conical spines up to 7 mm. long, and
with scales appressed, whitish or bicolorous (dark brown with whitish margins),
their marginal cells differentiated from those of the central portion in size, shape
and orientation; lamina pinnate to pinnate-pinnatisect, never fully bipinnate, tissue
glabrous, scales of the rachis and minor axes lacking or scattered on the abaxial
side, whitish to bicolorous or brown, flat (as in ours) or a few species with bullate
costal scales, trichomes of rachis and costa stiff, terete, recurved, pluricellular, copi-
ous to scattered, or lacking, but always lacking adaxially on costa and costules;
pinnae sessile to short-stalked, margins entire to deeply cleft (in ours merely lobed);
veins free and connivent to the sinus or more typically (as in ours) the basal ones
merging to form costal areoles ; sori disposed on the veins in a single line ( rarely sev-
eral lines) between costule and segment margin; indusium commonly hemitelioid,
semicircular (rarely fully circular, or reduced to one narrow lobe); receptacle ele-
vated, subglobose, with paraphyses rudimentary or lacking; spores 64 per sporan-
gium, provided with one large pore near the center of each face on or near the equa-
tor, and with numerous smaller apertures scattered over the surface, perine lacking.
Cnemidaria is the most distinctive of all the squamate genera of
Cyatheaceae. Its species are characterized by the generally acaules-
cent habit, the comparatively simple leaf architecture, and the basal
veins of segments conniving at the sinus or, more commonly, merg-
ing to form costal areoles. Moreover, the three large, regularly
spaced pores of the spores are practically unique in the family.
Typical of the other squamate genera of Cyatheaceae are the arbo-
rescent habit, the highly dissected leaves, and the free and noncon-
nivent veins. Cnemidaria is a tropical American genus with 23
species, one of which occurs in Guatemala.
Cnemidaria decurrens (Liebm.) Tryon, Contr. Gray Herb. 200: 52.
1970. Hemitelia decurrens Liebm. Kongel. Danske Vidensk. Selsk.
Skr. V. 1: 286 (seors. 134). 1849 (type from Lobani, Chinantla, Oaxa-
ca, Mexico, Liebmann PL Mex. 2089 (no. 912). H. mexicana Liebm.
torn. cit. 287 (seors. 135) (type from Lacoba, Chinantla, Oaxaca,
Mexico, Liebmann PI. Mex. 2105 (3 sheets: nos. 909, 910, 911). H.
guatemalensis Maxon, Contr. U.S. Natl. Herb. 16: 40. 1912 (type
from Alta Verapaz, Saluin s.n. ). H. lucida (Fee) Maxon, torn. cit. 39.
Cyathea guatemalensis (Maxon) Domin, Pteridophyta 264. 1929. C
lucida (Fee) Domin, loc. cit.
On slopes, in forests, 200-1,000 m. ; Alta Verapaz. Mexico (Chi-
apas and Oaxaca); Honduras.
Stem rudimentary to 0.3 m. long; leaves pinnate to pinnate-pinnatifid, to 2.5 m.
long and 0.8 m. broad; petiole to 0.5 m. long, with spines to 2 mm. long, the scales
scattered to abundant, whitish or bicolorous (brown-centered, with whitish mar-
FIG. 19. Cnemidaria decurrens. a, habit, X Vz; b, portion of pinna, showing vena-
tion, X 2; c, sorus and sporangium, greatly enlarged; d, sori, with sporangia re-
moved to show indusium, X 6.
107
108 FIELDIANA: BOTANY, VOLUME 39
gins); rachis with membranous wing to 2 mm. wide running partially to fully down
each side or (rarely) lacking, provided on the abaxial side with sparse to abundant
whitish or bicolorous scales and minute, terete, recurved trichomes; pinnae subses-
sile, subentire or crenate, or cut one-third to the costa; costae on abaxial side with
scattered, broad, amorphous, whitish or bicolorous scales, trichomes lacking; cos-
tules with scales and trichomes lacking; secondary segments or lobes (when
present) obtuse to rarely subapiculate; veins from the midribs simple, basal ones
commonly merging to form costal areoles; indusium semicircular to (rarely) almost
fully circular, subentire to several-lobed.
CYATHEA J. E. Smith
References: W. R. Maxon, Cyatheaceae (Cyathea, in part) in
North Amer. Fl. 16: 65-88. 1909. W. R. Maxon, The North American
species of Hemitelia, section Euhemitelia (in part), Contr. U.S.
Natl. Herb. 17: 414-420. 1914. R. M. Tryon, A revision of the genus
Cyathea, Contr. Gray Herb. 206: 19-101. 1976.
Stem erect, commonly massive and trunklike; leaves to several meters long; peti-
ole smooth to muricate, or provided with nonsquaminate spines ( the spines not bear-
ing scalelike remnants along their margins or at their bases), scales commonly ap-
pressed, brownish, often with paler or whitish margins, the marginal cells differenti-
ated from those of the central portion in shape, size and orientation, lacking
marginal setae; lamina bipinnate or more highly dissected (in ours bipinnate-pin-
natisect), the axes variously provided with scales and trichomes abaxially, and
(especially on costae and costules) with minute, recurved trichomes on the adaxial
side; veins free, simple or branched, in lobed segments the basal veins reaching the
margin at a point above the base of the sinus; indusium hemitelioid to globose
(sphaeropteroid); spores commonly 64 per sporangium, exine with or without
apertures of various size, but lacking a single large equatorial pore near the center of
each face.
Cyathea is very closely related to Trichipteris. Sterile specimens
of some species are difficult to place in either genus, for the chief
character by which the two may be distinguished is presence or ab-
sence of an indusium.
The indusium in Cyathea may be globose (sphaeropteroid) or cup-
shaped (cyathiform), thus subtending and completely encircling the
sorus, or it may be hemitelioid, i.e., borne proximally on the vein
and subtending but only partially encircling the sorus. The hemiteli-
oid indusium is attached by a broad base beneath the receptacle. It
may spread up and somewhat over the sorus on the proximal side,
or it may be low and saucer-shaped, and sometimes fugacious, thus
appearing flattened and scalelike on mature specimens.
A few species of Trichipteris may appear indusiate, due to the
presence of conspicuous scales around the sori. But close examina-
STOLZE: FERNS OF GUATEMALA 109
tion will reveal that these soral scales are attached by very narrow
bases, around the receptacle, and not necessarily on the proximal
side. Even the smallest of the hemitelioid indusia in Cyathea are
very broadly attached, describing an arc of 75-180° around the
receptacle.
Cyathea is a neotropical genus consisting of about 40 species,
three of which are known in Guatemala.
a. Sori medial to supramedial between midrib and segment margin; indusium hemi-
telioid, partially encircling the sorus on the proximal side; most ultimate seg-
ments 1.5 (2) times as long as broad, beyond the sinus C. multiflora.
a. Sori inframedial between midrib and segment margin; indusium at first globose,
later opening and becoming cup-shaped, completely encircling the sorus; most
ultimate segments 2-3 times as long as broad, beyond the sinus.
b. Petiole scales brown, essentially concolorous (often with margins a paler
brown); petiole scurf (minute squamulae) dark brown; pinnules rarely stalked,
or stalks on a few proximal ones to 1 mm C. fulua.
b. Petiole scales bicolorous (dark brown with broad to narrow, whitish margins);
petiole scurf (minute squamulae) tawny to whitish or bicolorous; pinnules on
the proximal third of larger pinnae short-stalked (2-6 mm.). . . . C. tuerckheimii.
Cyathea fulva (Mart. & Gal.) Fee, Mem. Fam. Foug. 9: 34. 1857
(not C. fulva Sod. 1883). Alsophila fulva Mart. & Gal. Nouv. Mem.
Acad. Roy. Sci. Bruxelles 15: 78, t.23. 1842 (type from Oaxaca,
Mexico, Galeotti 6346). C. furfuracea Christ, Bull. Herb. Boissier II.
4: 950. 1904 (not C. furfuracea Baker, 1874). C. onus ta Christ, loc.
cit. C. conspersa Christ, op. cit. 5: 260. 1905. C. underwoodii Christ,
op. cit. 6: 183. 1906. C. delicatula Maxon, Contr. U.S. Natl. Herb.
13: 4. 1909 (type from Alta Verapaz, between Tactic & Coban,
Tuerckheim 11-1629). C. mollis Rosenst. in Fedde, Repert. Spec.
Nov. Regni Veg. 22: 2. 1925 (not C. mollis Copel. 1917).
On wooded slopes and in ravines, in mountain forest, 900-3,300 m.
Alta Verapaz; El Progreso; Quezaltenango; San Marcos; Zacapa.
Southern Mexico; Honduras; Nicaragua; Costa Rica; Panama; Co-
lombia; Venezuela.
Stem to 5 m. tall; leaves to 3 m. long; petiole light or dark brown, darkest at base,
armed with stout, non-squaminate spines to 0.5 cm. long, or becoming aculeate or
muricate near the blade, essentially glabrous, but bearing abundant, appressed
scales at base, these lanceolate or linear-lanceolate, to 3 cm. long, dark brown, essen-
tially concolorous, often interspersed with brownish squamulae (scurf); lamina bi-
pinnate-pinnatisect, the tissue glabrous; rachis light brown or yellow-brown,
smooth to muricate, or aculeate on lower portion, scales essentially lacking, glab-
rous abaxially, minute-strigose adaxially and often brown-scurfy abaxially at pinna
FIG. 20. Cyathea. a-b, C. tuerckheimii: a, habit, X Vi; b, ultimate segments, X 3,
with inset, showing closed and open indusia, greatly enlarged; c, C. multiflora, ulti-
mate segments, X 3, with inset, showing indusia, greatly enlarged, with and without
sporangia; d, C. tuerckheimii, bicolorous petiole scale, X 21/z; e, C. fulva, concolorous
petiole scale, X 21/2.
110
STOLZE: FERNS OF GUATEMALA 111
bases; pinnae mostly sessile, or a few of the basal ones short-stalked (to 1.5cm.), lan-
ceolate, acuminate; pinnules sessile or subsessile (on largest pinnae rarely a few
short-stalked), incised nearly to the costule; costae and costules densely but minute-
ly strigose adaxially, glabrous or sparsely strigose abaxially, scales of the costules
flat, dull brown, rather abundant or (in ours) scattered or lacking; ultimate seg-
ments subfalcate, 2-3 times as long as broad beyond the sinus, subentire to crenu-
late or crenate-serrate, plane or somewhat revolute, obtuse to subacute, midribs
sparsely strigose, provided abaxially with brownish, acuminate, bullate or sub-
bullate scales; veins simple or (more commonly) once-forked, glabrous or sparsely
strigose adaxially; sori borne at or near the vein fork, inframedial between midrib
and segment margin; indusium globose, becoming broadly lacerate and irregularly
cup-shaped to somewhat flattened at maturity; receptacle subglobose or elongate;
paraphyses about as long as or shorter than the sporangia.
The key characters of petiole scales and scurf, which are effective-
ly used to separate this species from Cyathea tuerckheimii, may be
frequently lacking on herbarium sheets (tree fern collections are
typically fragmentary and often consist of only one or two pinnae)
and the other key character (pinnules stalked or sessile) is not con-
sistently reliable. However, the same kind of scurf present on the
petiole may be often found also on the abaxial side of the rachis,
around the base of the pinna stalks, and sometimes on the stalks
themselves. That of C. fulva is dark and dull brown, while that of C.
tuerckheimii is tawny to whitish, or bicolorous.
Cyathea multiflora J. E. Sm. Mem. Acad. Roy. Sci. (Turin) 5:
416. 1793. Hemitelia multiflora (J. E. Sm.) R. Br. Spreng. Syst.
Veg. 4: 126. 1827. Alsophila multiflora (J. E. Sm.) Presl, Tent.
Pterid. 61. 1836. H. nigricans Presl, Epim. Bot. 31. 1849.
In dense, wet forest, on wet slopes and stream banks, 40-900 m.
Alta Verapaz; Izabal. British Honduras; Honduras; Nicaragua to
Colombia; Ecuador; Peru; Bolivia.
Stem to 5 m. tall ( in ours to 2 m. ) ; leaves to 2 m. long; petiole light brown, sparsely
to densely strigose, or glabrescent, armed with stout spines to 5 mm. long, and
toward the base provided with appressed scales to 2 cm. long, these commonly lan-
ceolate, attenuate, dark brown, often with paler margins; lamina bipinnate-pinnati-
sect, the tissue (in ours) glabrous, firm papyraceous; rachis dark brown to yellow-
brown, aculeate to muricate, or smooth on the distal portion, scales essentially
lacking, strigose, or glabrescent abaxially; pinnae subsessile or (especially the more
proximal ones) short-stalked, elliptic to oblanceolate, abruptly acuminate; pinnules
sessile or subsessile; costae with minute, brown scales, few and widely scattered or
lacking, rather densely strigose adaxially, and densely to sparsely strigose or pilose
or glabrescent abaxially; costules densely strigose adaxially, provided with flat or
bullate, attenuate, brown (in ours) scales abaxially; ultimate segments straight or
slightly falcate, 1.5 (2) times as long as broad, beyond the sinus, subentire to broad-
ly dentate or crenate, plane or somewhat revolute, the apices rounded or truncate,
112 FIELDIANA: BOTANY, VOLUME 39
midribs glabrescent or bearing several stout, rigid trichomes adaxially, sometimes
with a few bullate scales abaxially; veins simple or once-forked; sori borne on the
veins, medial to supramedial between midrib and segment margin; indusium hemi-
telioid, subtending and partly encircling the sorus on the proximal side, flattened or
curving upward around the sorus, subentire or deeply lacerate, often fugacious at
maturity; receptacle subglobose; paraphyses about as long as the sporangia.
Cyathea multiflora varies considerably throughout its range,
especially outside of Guatemala. In our area it is a relatively small
tree fern, with a trunk usually less than 2 m. long, growing chiefly
at lower elevations; but in South America it has been often found
above 2,000 m. and with trunks up to 5 m. long. In Guatemala the
trichomes on costule and midrib are brownish, and the tissue is
glabrous, yet some specimens, from Costa Rica to South America,
have whitish pinnule indument, and the tissue sparsely and minute-
ly pilose on the abaxial side. The petiole scales of C. multiflora in
Guatemala are brown and essentially concolorous, but in Colombia
they are often bicolorous (brown with whitish margins) much like
the scales of Cnemidaria.
Cyathea tuerckheimii Maxon, Contr. U.S. Natl. Herb. 13: 4. 1909
(type from Guatemala, without location, 1907, Tuerckheim 11-1645).
In and at edges of wet forests, on slopes and in ravines, 1,300-
3,000 m. Alta Verapaz; Baja Verapaz; Huehuetenango; Jalapa;
Quezaltenango; El Quiche; San Marcos; Zacapa. Southern Mexico.
Stem to 8 m. tall; leaves to 3 m. long; petiole dark brown or olive-brown to stra-
mineous, darkest at base, armed with stout spines to 0.4 cm. long, or becoming
aculeate or muricate near the blade, essentially glabrous, but bearing abundant,
appressed scales at base, these lanceolate or linear-lanceolate, to 3 cm. long, dark
brown, mostly with whitish margins, and often interspersed with minute, whitish,
tawny or bicolorous squamulae (scurf); lamina bipinnate-pinnatisect, the tissue
glabrous; rachis dark brown to yellow-brown or stramineous, muricate, or aculeate
on lower portion, scales essentially lacking, glabrous abaxially, minute-strigose ad-
axially and often pale-scurfy abaxially at pinna bases; pinnae short-stalked (1-3
cm.), or the more distal ones subsessile, deltoid-lanceolate, short-acuminate; pin-
nules subsessile to short-stalked, those of the proximal third of larger pinnae with
stalks 2-6 cm. long, incised nearly to the costule; costae and costules densely but mi-
nutely strigose adaxially, essentially glabrous abaxially, or the costules with a few,
scattered, brown, flat or sub-bullate scales; ultimate segments falcate or subfalcate,
2-3 times as long as broad beyond the sinus, crenulate to crenate-serrate, slightly
revolute, obtuse to subacute, midribs sparsely strigose or glabrescent, provided ab-
axially with brownish, acuminate, bullate or sub-bullate scales; veins commonly
once-forked, or some of them simple (or on a few plants 1- to 3-forked), glabrous, or
sparsely strigose adaxially; sori commonly borne at the vein fork, inframedial
between midrib and segment margin; indusium globose, becoming broadly lacerate
and irregularly cup-shaped at maturity; receptacle subglobose or elongate; para-
physes shorter than the sporangia.
STOLZE: FERNS OF GUATEMALA 113
This and Cyathea divergens Kunze, from South America and
southern Central America are easily confused. Pinnae of the latter
are generally long-stalked, and pinnules are long-stalked and ta-
pered from near the base. Pinnae of C. tuerckheimii are short-
stalked, with the pinnules sessile to short-stalked and commonly
tapered from about the middle. Rolla Tryon, who is currently pre-
paring a monographic study of Cyathea, has stated (pers. comm. )
that the two taxa probably do not merit more than varietal dis-
tinction.
LOPHOSORIA Presl
Plants acaulescent or with erect stems, massive and trunklike, to 1 m. tall; leaves
to several meters long, completely lacking scales; petiole to 1.5 m. long, light or dark
brown, often blackish at very base, smooth, unarmed, densely lanate at base with
brownish or rusty, septate trichomes; lamina subcoriaceous, commonly subdeltoid,
tripinnate to tripinnate-pinnatisect, pale to dark green above, commonly glaucous
beneath; rachis light brown or stramineous, villous or glabrescent; pinnae stalked,
broadly lanceolate to subdeltoid, the costae and costules villous (often densely so)
above and beneath with pale to dark brown trichomes; pinnules stalked, commonly
bearing 18-24 pairs of segments; ultimate segments incised halfway or nearly quite
to the midrib, the latter densely villous beneath (at least the proximal half), but
glabrous above; veins free, pinnately branched in each lobe, the tips of veinlets
reaching the margin at a point well above the sinus; sori one to a vein, borne on the
basal, acroscopic branch, each containing 6-10 sporangia; receptacle scarcely ele-
vated; paraphyses about as long as the sporangia; indusia lacking; spores 64 per
sporangium, whitish to pale yellow, globose-tetrahedral, encircled by a thickened,
uninterrupted, equatorial rim, laesura prominent, surface sparsely dotted with
minute apertures, perine lacking.
Lophosoria is generally considered the most primitive genus in
the family. It is confined to the neotropics and contains a single
species.
Lophosoria quadripinnata (Gmel.) C. Chr. in Skottsb., Nat. Hist.
Juan Fern. East. Islands 2: 16. 1920. Polypodium quadripinnatum
Gmel. Syst. Nat. 2 (2): 1314. 1791. P. pruinatum Sw. J. Bot. (Schra-
der) 1800 (2): 29. 1801. Alsophila pruinata (Sw.) Kaulf. in Kunze,
Linnaea 9: 99. 1834. Lophosoria pruinata (Sw.) Presl, Abh. Bohm.
Ges. Wiss. V. 5: 345. 1848. Alsophila quadripinnata (Gmel.) C. Chr.
Index Fil. 47. 1905.
Open hillsides, road cuts, and in wooded ravines and along
streams in deep forests, 1,500-3,800 m. Huehuetenango; El Pro-
greso; Quezaltenango; San Marcos; Solola; Zacapa. Southern Mex-
ico; Greater Antilles; Colombia and Venezuela to Brazil and Boliv-
ia; Southern Chile; Juan Fernandez Islands.
114
FIELDIANA: BOTANY, VOLUME 39
FIG. 21. Lophosoria quadripinnata. a, habit, X Vi; b, ultimate segments, X 2; c,
portion of ultimate segment, showing rigid trichomes and lax paraphyses, X 8.
Characters are those of the genus.
Lophosoria quadripinnata is abundant in regions in which it
occurs. Scales and indusia are totally lacking, and the highly dis-
sected leaves are often glaucous beneath, the white or silvery color
(especially on live plants) markedly contrasting with the dark green
adaxial surface. No other tree fern possesses this combination of
characters.
STOLZE: FERNS OF GUATEMALA
115
FIG. 22. Metaxya rostrata. a, habit, X Vi; b, base of pinna, X 2; c, portion of pinna,
showing venation, X 1 Vz ; d, sori, one with sporangia removed, greatly enlarged.
METAXYA Presl
Plants erect from stout, creeping rhizomes which are densely provided with fili-
form, septate, yellowish to orange trichomes; leaves to several meters long, com-
pletely lacking scales; petiole to nearly 1 m. long, light brown or stramineous,
smooth, unarmed, essentially glabrous; lamina chartaceous, simply pinnate, with
apical segment similar in shape and size to the lateral pinnae; rachis with scattered
116 FIELDIANA: BOTANY, VOLUME 39
brown trichomes or glabrescent; pinnae stalked, narrowly lanceolate, with margin
cartilaginous, entire, but coarsely dentate at the acuminate apices, glabrous above,
sparsely pilose along costae beneath; veins free, simple or sometimes forked at or
near the base, spreading from the costa at broad angles; sori 1-3 to a vein, crowding
the costa, each bearing numerous sporangia; receptacle flat; paraphyses commonly
longer than the sporangia; indusia lacking; spores 64 per sporangium, yellowish
brown, tetrahedral, with prominent laesura, without apertures, perine present,
essentially smooth.
Metaxya is found at low elevations in the American tropics and
consists of a single species.
Metaxya rostrata (HBK.) Presl, Tent. Pterid. 60. 1836. Poly-
podium rostratum Willd. Sp. PL 5: 193. 1810 (not P. rostratum
Burm. 1768). Aspidium rostratum HBK. Nov. Gen& Sp. 1: 12. 1815
(nom. nov.). Amphidesmium rostratum (Willd.) J. Sm. J. Bot.
(Hooker) 1: 201. 1842. Alsophila blechnoides (Rich.) Hook. Sp. Fil.
1:35.1844.
In ravines and along streams in forests, 40-300 m. Izabal. British
Honduras; Honduras; Nicaragua; Costa Rica; Panama; Trinidad;
Lesser Antilles; northern South America southward to Brazil and
Bolivia.
Characters are those of the genus.
Within the Cyatheaceae, no other species has widely spaced peti-
oles borne on a long-creeping rhizome. This, and the simply pinnate,
non-squamate leaves, easily separates Metaxya rostrata from all
other species in the family.
NEPHELEA Tryon
Reference: Gerald J. Gastony, A revision of the fern genus
Nephelea, Contr. Gray Herb. 203: 81-148. 1973.
Stem erect, commonly massive and trunklike (in ours to 10 m. tall), provided with
stout, blackish spines and (especially near the apex) large scales; unexpanded cro-
ziers densely scaly and bearing stout, blackish, squaminate spines; leaves to several
meters long; petiole variously colored (but not blackish), provided with stout, black-
ish, squaminate, mostly obturbinate spines often to 1.5 cm. long, and with large,
spreading scales which are commonly dark-colored, often with paler margins, the
marginal cells differentiated from those of the central portion in shape, size and ori-
entation, and with a dark apical seta and often some smaller lateral ones; lamina bi-
pinnate to tripinnatifid (in ours bipinnate-pinnatifid), the axes bearing abundant,
appressed, minute, recurved trichomes on the adaxial side; rachis brownish,
glabrous abaxially, or provided adaxially with scattered to abundant, spreading
scales and often with appressed, setose squamulae; pinnae sessile to stalked, the
axes and midribs commonly bearing flat or bullate scales abaxially; veins free,
STOLZE: FERNS OF GUATEMALA
117
FIG. 23. Nephelea. a-d, N. mexicana: a, habit, X Vi; b, ultimate segments, X 6;
c, petiole scales, greatly enlarged; d, petiole spines, greatly enlarged; e, N. tryoni-
ana, ultimate segments, X 6, with bullate scales, greatly enlarged.
simple or branched, in lobed segments the basal veins reaching the margin at a point
above the base of the sinus; sori disposed on the veins near the midrib, on branched
veins situated at the fork; indusium hemitelioid, meniscoid, sphaeropteroid, or lack-
ing; receptacle globose to elongate, and with paraphyses commonly inconspicuous,
shorter than the sporangia; spores 16 per sporangium, exine without apertures of
various size, lacking a single large equatorial pore near the center of each face,
perine provided with long, irregular ridges or folds.
118 FIELDIANA: BOTANY, VOLUME 39
Next to Cnemidaria this is perhaps the most distinctive of the
squamate genera in the Cyatheaceae. The petioles and young cro-
ziers are abundantly beset with stout, blackish, commonly obtur-
binate spines. These spines are squaminate, i.e., small spines bear
the same kind of differentiated marginal cells as do the scales. Al-
though these scaly portions are often caducous, at least some rem-
nants may be observed at the thickened bases of spines. Other
genera include some species with spiny petioles, but the spines are
not squaminate nor obturbinate. Petiole scales in Nephelea are slen-
der and spreading. Only Sphaeropteris has some species with
spreading scales, whereas in all other squamate genera the petiole
scales are commonly broad and appressed.
Nephelea is most closely related to Alsophila. Among the genera
of Cyatheaceae, only Alsophila, Nephelea, and some species of
Sphaeropteris bear setose scales. Gastony (Amer. J. Bot. 61: 672-
680. 1974) has found that most species of Alsophila and all of Neph-
elea have sporangia bearing 16 spores, whereas the rest of the
family typically has 64-spored sporangia. He also points out that
spore perine in Alsophila-Nephelea is characterized by long, irregu-
lar ridges. Other genera apparently have spore perine lacking this
type of ornamentation, or the perine lacking entirely.
Nephelea is a neotropical genus composed of 18 species which
occur principally in cloud forests. Two species are found in Guate-
mala.
a. Costules and midribs with whitish, bullate scales abaxially; indusium lack-
ing N. tryoniana.
a. Costules and midribs lacking bullate scales (some flat ones often present on cos-
tules); indusium subglobose (although often fugacious and vestigial on mature
plants) N. mexicana.
Nephelea mexicana (Schlecht. & Cham.) Tryon, Contr. Gray
Herb. 200: 40. 1970. Cyathea mexicana Schlecht. & Cham. Linnaea
5: 616. 1830. C. patellaris Christ, Annuaire Conserv. Jard. Bot.
Geneve 4: 207. 1900. Nephelea patellaris (Christ) Tryon, Contr.
Gray Herb. 200: 40. 1970. Alsophila costalis Christ, Bull. Herb.
Boissier II. 4: 951. 1904. Cyathea costalis (Christ) Domin, Pterido-
phyta 262. 1929. Alsophila tenerifrons Christ, Bull. Herb. Boissier
II. 4: 959. 1904. Cyathea tenerifrons (Christ) Domin, Pteridophyta
263. 1929. Nephelea tenerifrons (Christ) Tryon, Contr. Gray Herb.
200: 40. 1970. Cyathea gemmifera Christ, in Jimenez, Bol. Fomento
(San Jose) 3: 661. 1913. Palma de montana (fide Steyermark, Que-
zaltenango).
STOLZE: FERNS OF GUATEMALA 119
In forests, on slopes and in ravines, 200-2,800 m.; Alta Verapaz;
Baja Verapaz; Huehuetenango; Izabal; Quezaltenango; San Mar-
cos; Suchitepequez. Mexico; Honduras and El Salvador south to
Panama; Ecuador.
Stem to 10 m. tall, beset with blackish spines and dark brown scales, especially
near the apex; leaves to 4 m. long; petiole about 1 m. long, light to dark brown,
abundantly provided with stout, blackish spines (to about 1 cm. long), especially
toward the base, and also provided with scattered, slender, spreading, brown to bi-
colorous scales, in addition to a dense covering of minute (ca. 0.1 mm.) brown, ap-
pressed squamulae: lamina commonly bipinnate-pinnatifid, the tissue chartaceous
to subcoriaceous, glabrous, abruptly terminating in a pinna-like apex; rachis brown,
with larger scales essentially lacking, but with appressed, minute squamulae often
abundant on abaxial side, and with appressed, minute, recurved trichomes on adax-
ial side, or rachis often glabrescent distally; pinnae sessile to short-stalked, broadly
lanceolate, the costae and costules rather abundantly provided abaxially with
appressed, minute squamulae, and adaxially with minute, recurved, brown tri-
chomes, the costae also with scattered, whitish scales to 1 mm. long, abaxially; pin-
nules commonly deeply pinnatisect; ultimate segments subentire or crenate-ser-
rate, lacking bullate scales; veins simple to forked; sori crowding the midrib, often
at the vein forks; indusium subsphaeropteroid, irregularly cup-shaped at maturity,
or commonly breaking apart at maturity so that merely a remnant is attached to the
receptacle; receptacle globose or subglobose; paraphyses usually shorter than the
sporangia.
Nephelea tryoniana Gastony, Contr. Gray Herb. 203: 118. 1973.
Wet, mixed mountain forests, 1,500-2,500 m. Alta Verapaz; Zaca-
pa (type from Rio Lima, Sierra de las Minas, Steyermark 30009).
Honduras; Nicaragua.
Stem to 4.5 m. tall, densely provided with stout blackish spines and dark brown to
blackish scales (especially near apex); leaves to about 2 m. long; petiole to 0.4. m.
long, light to dark brown, abundantly provided with stout, blackish spines to ca. 1
cm. long (especially on the lower portions), becoming muricate toward base of lami-
na, and also provided with long (to 2 cm.) slender, spreading, pale to brown, often bi-
colorous scales, in addition to a rather dense covering of minute (ca. 0.1 mm.), pale
to brown, appressed squamulae; lamina commonly bipinnate-pinnatifid, the tissue
papyraceous, glabrous, rather gradually reduced to an acute apex; rachis light
brown, provided on both sides with abundant, long scales as on the petiole, and ab-
axially with appressed, minute squamulae, adaxially with appressed, minute,
recurved trichomes; pinnae sessile or subsessile, narrow-ovate to lanceolate, the cos-
tae scaly as on the rachis, and adaxially with appressed, minute, recurved tri-
chomes; pinnules commonly deeply pinnatisect, the costules densely covered
adaxially with recurved trichomes, abaxially with flat, whitish scales, these bullate
in the distal portion; ultimate segments slender, subentire, spreading at broad
angles from the costule, the midribs sparsely strigose, and often with whitish, bul-
late scales distally; veins simple to (more commonly) once-forked; sori commonly at
the forks of veins and inframedial between midrib and margin; receptacle elongate
or subglobose; paraphyses usually shorter than the sporangia; indusium lacking.
120 FIELDIANA: BOTANY, VOLUME 39
In addition to the characters used in the key, Nephelea tryoniana
can also be distinguished from N. mexicana by the abundance of
long scales borne on the rachis. In N. mexicana the rachis often may
be provided with minute, setose squamulae, but the long, conspicu-
ous scales (as on the lower petiole) are sparse or lacking.
SPHAEROPTERIS Bernhardi
Reference: R. M. Tryon, The American tree ferns allied to Sphae-
ropteris horrida, Rhodora73: 1-19. 1971.
Stem erect, commonly massive and trunklike; leaves to several meters long; peti-
ole smooth to muricate (in ours) or provided with nonsquaminate spines (the spines
not bearing scale-like remnants along their margins or at their bases), scales com-
monly spreading or rarely somewhat appressed abaxially, structurally conform, the
marginal cells similar to those of the central portion in shape, size orientation, and
usually bearing apical and marginal setae; lamina typically bipinnate or more highly
dissected, in a few species simply pinnate (in ours bipinnate-pinnatisect to subtri-
pinnate), the axes variously provided with scales and trichomes abaxially, and
(especially on costae and costules) with minute, recurved trichomes on the adaxial
side; veins free, simple or branched, in lobed segments the basal veins reaching the
margin at a point above the base of the sinus; indusium hemitelioid to globose, or
lacking; spores 64 per sporangium, with or without apertures of various size, but
lacking a single, large, equatorial pore near the center of each face, commonly (as in
ours) with perine.
This tropical genus contains over 100 species, with a large major-
ity in the Old World. Most of the neotropical species occur in South
America, and only the following two have been found in Guatemala.
a. Indusium nearly globose, cup-shaped at maturity; most petiole scales whitish or
tawny, with dark-colored setae; rachis and costae abundantly and conspicuously
scaly S. horrida.
a. Indusium lacking; petiole scales lustrous brown, with setae commonly concolor-
ous; rachis and costae with scales sparse and inconspicuous or lacking
S. myosuroides.
Sphaeropteris horrida (Liebm.) Tryon, Contr. Gray Herb. 200: 20.
1970. Cibotium horridum Liebm. Kongel. Danske Vidensk. Selsk.
Skr. V. 1: 279. 1849 (not Cyathea horrida (L.) J. E. Sm. 1793). Cya-
theaprinceps E. Mayer, Gartenfl. 17: 10: 1868. C. bourgaei Fourn.
Mex. PI. 1: 135. 1872. C. glauca Fourn. loc. cit. C. munchii Christ,
Bull. Herb. Boissier II. 7: 413. 1907.
In forests and damp thickets, along rivers and on sides of ravines,
sometimes found on open banks, 500-2,000 m. Alta Verapaz; Baja
Verapaz; Chiquimula; Huehuetenango; Quezaltenango; Suchitepe-
. . •
FIG. 24. Sphaeropteris. a-c, S. horrida: a, pinna, habit, X Vi; b, ultimate segments,
X 5, with one sorus greatly enlarged; c, petiole scales, greatly enlarged; d-e, S. myo-
suroides: d, portion of petiole, X IVi; e, petiole scales, greatly enlarged.
121
122 FIELDIANA: BOTANY, VOLUME 39
quez. Southern Mexico; Honduras. Palma de montana; palmita
(fide Standley, Quezaltenango).
Stem to 15 m. tall and 40 cm. in diameter; leaves to 5 m. long; petiole light to dark
brown, muricate with the persistent bases of scales, the scales abundant (especially
on the lower petiole), appressed or, more commonly, spreading, to 3 cm. long, linear,
most of them whitish to tawny, with rigid, dark, apical and lateral setae, trichomes
lacking; lamina bipinnate-pinnatisect to subtripinnate, the tissue glabrous, charta-
ceous to subcoriaceous, somewhat glaucous beneath; rachis brown or stramineous,
muricate, abundantly scaly, trichomes essentially lacking, or somewhat villous on
the distal portion adaxially; costae abundantly scaly abaxially, appressed-villous
adaxially, costules glabrous or with a few filiform scales abaxially, abundantly pro-
vided with minute, recurved trichomes adaxially; pinnules sessile, narrow-oblong,
cut nearly (or rarely, quite) to the base; ultimate segments subfalcate, obtuse to
subacute, entire to crenulate, with slightly revolute margins, glabrous or the mid-
ribs beneath with scattered, pale trichomes or filiform scales; veins once- or twice-
forked; sori commonly borne at the vein fork, crowding the midrib; indusium nearly
globose, rupturing apically at maturity but the base persistent, conspicuous, cup-
shaped; receptacle subglobose or elongated, the paraphyses relatively short and in-
conspicuous; spores brownish, tetrahedral, surface essentially smooth, without
apertures.
Of all the American species in the family, plants of Sphaerop tens
horrida are among the largest and most striking. The stout trunk
bears a handsome crown of huge, spreading leaves which are sup-
ported by massive, scaly petioles. In Quezaltenango, the trunks are
reportedly used in the construction of houses. In Mexico the fern is
often called "rabo de mico."
Sphaeropteris myosuroides (Liebm. ) Tryon, Contr. Gray Herb.
200: 20. 1970. Alsophila myosuroides Liebm. Kongel. Danske
Vidensk. Selsk. Srk. V. 1: 286. 1849 (type from Chinantla, Oaxaca,
Mexico, Liebmann s.n. , 1842).
In open woods, on slopes and along streams, sea level to 400 m.
Alta Verapaz; Izabal; Peten. Cuba; Southern Mexico; British Hon-
duras; Honduras; Nicaragua.
Stem to 3 m. tall, sometimes aculeate toward apex; leaves to 2.5 m. long; petiole
to 1 m. long, light or dark brown, or stramineous, darkest at base, densely aculeate,
or muricate toward apex, provided with abundant, appressed, minute trichomes,
scales abundant near the base, linear, to 3 cm. long, bright shiny-brown, setose; lam-
ina bipinnate-pinnatisect to subtripinnate, ovate, the tissue glabrous, chartaceous;
rachis light or grayish brown or stramineous, muricate, rather densely though
minutely strigose with terete, recurved trichomes, scales essentially lacking; pinnae
stalked, costae and costules strigose, densely so on the adaxial side, with scales
lacking, or widely scattered on the costae abaxially; pinnules subsessile to short-
stalked, lanceolate to oblong, cut nearly or quite to the base; ultimate segments sub-
falcate, subacute, crenate-serrate, with slightly revolute margins, glabrous or the
STOLZE: FERNS OF GUATEMALA 123
midribs beneath sparsely provided with pale trichomes and a few scattered, brown
scales; veins once or twice forked; sori medial to inframedial, borne at or slightly
above the vein forks; indusium lacking; receptacle small, subglobose or flattened,
the paraphyses longer than the sporangia, conspicuous; spores brownish, tetrahe-
dral, surface essentially smooth, without apertures.
This and Sphaeropteris horrida are quite distinct, separated
easily by strong characters of indusia and scales. The latter is also a
much larger plant, the trunks often attaining lengths of up to 15 m.,
whereas those of S. myosuroides do not exceed 3 m. Furthermore, S.
myosuroides is a lowland fern, commonly growing at or near sea
level and not found (at least in Guatemala) above 400 m. S. horrida
prefers middle elevations, generally 1,000-1,500 m.
TRICHIPTERIS Presl
References: W. R. Maxon, The North American species of Al-
sophila grouped with A. armata, Contr. U.S. Natl. Herb. 24: 33-46.
1922. Ramon Riba, Revision monografica del complejo Alsophila
swartziana Martius (Cyatheaceae), Anales Inst. Biol. Univ. Nac.
Mexico 38: 61-100. 1967.
Stem horizontal to erect, commonly massive and trunklike, leaves to several
meters long; petiole smooth, muricate or provided with nonsquaminate spines (the
spines not bearing scalelike remnants along their margins or at their bases), scales
commonly appressed, brownish, or bicolorous, with pale or whitish margins, the
marginal cells differentiated from those of the central portion in shape, size and ori-
entation, lacking marginal setae ( though sometimes with dark, marginal denticula-
tions); lamina simply pinnate to tripinnate-pinnatifid (in ours pinnate-pinnatisect to
subtripinnate-pinnatifid), the axes variously provided with scales and trichomes
abaxially, and (especially on costae and costules) with minute, recurved, trichomes
adaxially; veins free (rarely, outside our area, anastomosing), simple or branched, in
lobed segments the basal veins reaching the margin at a point above the base of the
sinus; indusium lacking; spores commonly 64 per sporangium, exine with or with-
out apertures of various size, but lacking a single large, equatorial pore near the
center of each face.
Although very closely related to Cyathea the species of Trichip-
teris comprise a separate evolutionary group, distinguished from
the former by lack of an indusium. For further discussion see Cy-
athea.
The genus contains about 55 species, and is confined to the Ameri-
can tropics.
a. Leaves pinnate-pinnatisect T. ursina.
a. Leaves bipinnate-pinnatifid or more complex.
b. Petiole scales abundant throughout, whitish; scales of secondary axes (costae)
frequent, flat, whitish; petiole spines lacking T. mexicana.
124 FIELDIANA: BOTANY, VOLUME 39
b. Petiole scales abundant only at base (essentially lacking above), brown or bicol-
orous; scales of secondary axes (costae) lacking, or sparse, tawny to brown,
flat to bullate; petiole spines abundant ( except lacking in T. costaricensis).
c. Costa and rachis abundantly spiny throughout T. microdonta.
c. Costa unarmed (rarely muricate), rachis usually unarmed, at least in the
distal portion.
d. Ultimate segments subentire, broadly rounded at apex, 2 to 2!/2 times as
long as broad; petiole scales dark brown, concolorous; sori medial to supra-
medial between midrib and segment margin T. schiedeana.
d. Ultimate segments crenate, serrate, or lobed, acute or subacute, 3Vi to 5
times as long as broad; petiole scales bicolorous, brown with whitish mar-
gins; sori mostly inframedial between midrib and segment margin.
e. Primary axis densely hirsute, sometimes scabrous with the persistent
bases of trichomes T. scabriuscula.
e. Primary axis glabrous or sparsely hirsute, the deciduous trichomes often
leaving a dark scar but an essentially smooth surface,
f. Sori subtended by narrowly attached squamulae (thus appearing indu-
siate) ; margins of petiole scales lacking dark denticulations; petiole un-
armed; larger pinnae often stalked T. costaricensis.
f. Sori lacking basal squamulae; margins of petiole scales with scattered
or continuous dark denticulations (at least near scale apex); petiole
spiny; pinnae sessile (or some basal ones short-stalked),
g. Petiole scales dark brown, with narrow, whitish margins and tips, the
margins (toward their apices) provided with abundant, often con-
tinuous, dark brown denticulations. Guatemala
T. pansamalana.
g. Petiole scales whitish, each with a narrow, brown median stripe or
with a dark spot at base, the margins provided with scattered, dark
brown denticulations. Mexico . . . . T. bicrenata.
Trichipteris bicrenata (Liebm.) Tryon, Contr. Gray Herb. 200: 44.
1970. Cyathea bicrenata Liebm. Kongel. Danske Vidensk. Selsk.
Skr. V. 1: 289. 1849 (type from near Totontepec, Oaxaca, Mexico,
Liebmann s.n.). Alsophila bicrenata (Liebm.) Fourn. Mex. PI.
Crypt. 134. 1872.
In wet forests and thickets, 1,000-2,000 m. Known thus far only in
southern Mexico, but since it has been collected in Chiapas it may
be also expected in adjacent areas of Guatemala.
Stem to 10 m. tall; leaves to several meters long; petiole yellow-brown, glabrous
or sparsely hirsute, abundantly provided with sharp, stout spines to 2.5 mm. long,
but the surface otherwise smooth, with appressed scales at the base, these whitish
to bicolorous (whitish with a narrow, dark median stripe), the margins with scat-
tered dark brown denticulations; lamina bipinnate-pinnatisect or subtripinnate-pin-
natifid, the tissue glabrous, thin-herbaceous; rachis spiny on the lower portion,
hirsute or glabrescent, yellow-brown to stramineous, scales essentially lacking; pin-
nae sessile; costae lacking spines and scales, but abundantly provided with orange
to light brown trichomes, appressed adaxially, spreading abaxially ; pinnules sessile,
FlG. 25. Trichiptens. a-b, T. schiedeana: a, pinna, habit, X 1/2; b, ultimate seg-
ments, X 41/2; c, T. scabriuscula, ultimate segments, X 5; d, T. pansamalana, tip of
petiole scale, greatly enlarged.
125
126 FIELDIANA: BOTANY, VOLUME 39
cut nearly to the costule into linear, subfalcate segments, the sinuses acute to broad-
ly angular; costules adaxially provided with tawny or light brown, mostly appressed
trichomes, abaxially hirsute and with brownish, flat or bullate scales at least on
proximal portions; ultimate segments deeply crenate-serrate or lobed, obtuse to
subacute, the midribs sparsely strigose or glabrescent on the adaxial side, abaxially
hirsute and provided with light brown or tawny, bullate scales near the bases of seg-
ments; veins once- or twice-forked in each segment lobe; sori commonly inframedial
between midrib and segment margin; receptacle small, elongated to subglobose;
paraphyses longer than the sporangia.
Trichipteris costaricensis (Kuhn) Barr., Rhodora 78: 1. 1976.
Hemitelia costaricensis Mett. Ann. Sci. Nat. Bot. V. 2: 265. 1864
nom. nud. H. costaricensis Kuhn, Linnaea 36: 159. 1869. Cyathea
costaricensis (Kuhn) Domin, Acta Bot. Bohem. 9: 107. 1930.
In forests, on mountain slopes, river banks and in ravines, also on
open slopes and roadside banks, 600-1,500 m. Chiquimula; Escuin-
tla; Quezaltenango; Retalhuleu; Santa Rosa. Mexico; Honduras
and El Salvador to Panama.
Stem to 4 m. tall; leaves to several meters long; petiole yellow-brown, trichomes
essentially lacking, unarmed (at least in our area), surface smooth, or minutely
tuberculate, with appressed scales at base, these broad and bicolorous, whitish with
dark brown median stripes or tips; lamina bipinnate-pinnatisect, the tissue glabrous,
firm-herbaceous; rachis unarmed, essentially glabrous; pinnae short-stalked; cos-
tae lacking spines and scales, with light or dark brown minute, recurved trichomes
adaxially, essentially glabrous abaxially; pinnules sessile or subsessile, cut nearly to
the costule into narrow, subfalcate segments, the sinuses acute to narrowly round-
ed; costules provided adaxially with abundant light or dark brown, minute, re-
curved trichomes, abaxially glabrous or with scattered, lax, pale brown trichomes
and scattered, broad, tawny or brownish, flat to sub-bullate scales; ultimate seg-
ments acute or subacute, crenulate to deeply crenate, the lobes often bidentate, the
midribs with a few rigid trichomes adaxially, midrib and veins glabrescent or with
rather abundant, whitish or pale brown, lax trichomes abaxially; veins once- or
twice-forked; sori inframedial (rarely medial) between midrib and segment margin,
subtended by one to a few membranaceous squamulae resembling indusial seg-
ments, these greatly narrowed and attached by a thin point at base, narrowed to a
usually filamentous apex; receptacle subglobose; paraphyses about as long as the
sporangia.
Trichipteris mexicana (Mart.) Tryon, Contr. Gray Herb. 200: 44.
1970. Alsophila mexicana Mart. Icon. PL Crypt. Bras. 70. t.45.
1834 (type from San Pablo de Teoxomulco, Oaxaca, Mexico, Kar-
winsky s.n.) (not Cyathea mexicana Schlecht. & Cham. 1830). A.
godmanii Hook, in Hook. & Bak., Syn. Fil. 36. 1866.
STOLZE: FERNS OF GUATEMALA 127
Wet forests, in ravines and along streams, 800-1,800 m. Alta
Verapaz ( type from Coban, Godman & Salvin s. n. ) ; Huehuetenan-
go; Quezaltenango; San Marcos. Southern Mexico; Honduras.
Stem to 10 m. tall; leaves to several meters long; petiole yellow-brown to stramin-
eous, tuberculate to muricate, but lacking spines, densely whitish-hirsute and
copiously provided with appressed, broad, flat scales, these white or very pale yel-
low, concolorous, or with a narrow, brown median stripe; lamina bipinnate-pinnati-
sect to subtripinnate-pinnatifid, the tissue glabrous, thin-herbaceous; rachis and
costae light brown to stramineous, adaxially densely hirsute-strigose, abaxially
hirsute with whitish, septate trichomes, or muricate with their persistent bases,
abundantly provided with broad, flat, whitish to pale yellow scales; pinnae sessile or
short-stalked; pinnules sessile, cut nearly to the costule into pinnatifid segments,
the sinuses rather broadly rounded ; costules adaxially provided with tawny or light
brown, appressed trichomes, abaxially hirsute and with abundant whitish to tawny,
broad and flat, or (more commonly) bullate scales; ultimate segments deeply cre-
nate to pinnatifid, obtuse to subacute, the midribs on both sides with scattered,
rigid trichomes, provided with scattered, whitish, bullate scales abaxially; veins
commonly once- or twice-forked in each segment lobe; sori inframedial between mid-
rib and segment margin, often confined to the proximal third of the segment; recep-
tacle minute, subglobose; paraphyses few, as long as or slightly longer than
the sporangia.
Trichipteris microdonta (Desv.) Tryon, Contr. Gray Herb. 200:
46. 1970. Polypodium microdontum Desv. Ges. Naturf. Freunde
Berlin Mag. Neuesten Entdeck. 5: 319. 1811. Alsophila microdonta
(Desv.) Desv. Mem. Soc. Linn. Paris 6: 319. 1827. A. armata Mart.
Icon. PL Crypt. Brasil 72, t.28. 1834 (not A. armata (Sw.) Presl.
1836). Cyathea microdonta ( Desv. ) Domin, Pteridophyta263. 1929.
In wet forests and thickets, in swamps and along streams, sea
level to 250 m. Alta Verapaz; Izabal. Southern Mexico; Honduras;
Nicaragua; Costa Rica; Panama; Greater Antilles; Trinidad; Co-
lombia; Venezuela; theGuianas; Brazil; Peru.
Stem to 4 m. tall, armed with stout spines; leaves to 2.5 m. long; petiole dark
brown, with slender, conical, often curved spines to 1 cm. long, sparsely to moder-
ately strigose to glabrescent, or provided with scurf composed of brownish squamu-
lae, and toward the base with appressed scales to 1.5 cm. long, these commonly
lanceolate, attenuate, dark brown, concolorous; lamina bipinnate-pinnatisect, tissue
glabrous, firm membranaceous ; rachis and costae dark brown to yellow-brown,
provided throughout with distant, sharp spines, glabrous abaxially, minutely but
often densely strigose adaxially, scales essentially lacking, or costae with a few scat-
tered, brown, filiform scales; pinnae (except apical ones) stalked; pinnules sessile or
subsessile, or some larger ones short-stalked, cut nearly to the costule into narrow
segments; costules densely strigose adaxially, provided abaxially with pale to
brown trichomes and minute, pale to brown, flat or sub-bullate scales; ultimate
segments subfalcate, linear, 3.5-5 times as long as broad (larger ones 2-3 mm. wide),
obtuse to subacute, margin crenulate to serrate, plane or slightly revolute, the si-
128 FIELDIANA: BOTANY, VOLUME 39
nuses acute or narrowly rounded, midribs with scattered, rigid trichomes on both
sides, and with a few minute, brownish bullate scales abaxially; veins once-forked;
sori borne at or near the vein fork, medial to inframedial between midrib and seg-
ment margin; receptacle minute, flattened; paraphyses flaccid, as long as or longer
than the sporangia.
Trichipteris pansamalana (Maxon) Tryon, Contr. Gray Herb.
200: 44. 1970. Alsophila pansamalana Maxon, Contr. U.S. Natl.
Herb. 24: 40. 1922.
At edges of forests, in ravines, along river banks, 900-2,800 m.
Alta Verapaz (type from Pansamala, Tuerckheim [J. D. Smith No.
1008]); Huehuetenango; Quezaltenango; San Marcos.
Stem to 3 m. tall; leaves to several meters long; petiole yellow-brown, glabrous, or
sparsely hirsute on lower portion, abundantly provided with sharp, straight, stout
spines to 4 mm. long, but the surface otherwise essentially smooth, with appressed
scales at base, these bicolorous, with a broad, dark brown median stripe, the mar-
gins with abundant, dark denticulations which are often continuous (at least near
the scale apex); lamina bipinnate-pinnatisect or subtripinnate-pinnatifid, the tissue
glabrous, thin-herbaceous; rachis spiny on the lower portion, sparsely hirsute-stri-
gose, or more densely so toward apex, scales lacking; pinnae sessile or the basal pair
stalked; costae lacking spines and scales, but sparsely hirsute-strigose abaxially,
more densely so adaxially; pinnules sessile, cut nearly to the costule into linear,
subfalcate segments, the sinuses acute to rounded to squared; costules sparsely
hirsute-strigose adaxially, pilose-strigose abaxially and with scattered, tawny,
flattened or bullate scales; ultimate segments deeply crenate-serrate or lobed,
obtuse to subacute, the midribs essentially glabrous on the adaxial side, but abaxi-
ally pilose-strigose and with scattered whitish or tawny bullate scales, especially
near segment base; veins 1- to 3-forked; sori inframedial between midrib and seg-
ment margin; receptacle elongated to subglobose; paraphyses shorter than the
sporangia.
Trichipteris scabriuscula (Maxon) Tryon, Contr. Gray Herb. 200:
44. 1970. Alsophila scabriuscula Maxon, Proc. Biol. Soc. Wash. 32:
125. 1919. A scabriuscula var. guatemalensis Riba, Rhodora 69: 68.
1967 (type from Huehuetenango, Steyermark 49417).
Rich, wet forests, often along stream banks, sea level to 800 m.
Alta Verapaz (type from Cubilguitz, Tuerckheim [J. D. Smith no.
7806]); Huehuetenango; Izabal. Southern Mexico; Honduras;
Nicaragua.
Stem to 6 m. tall; leaves several meters long; petiole yellow-brown, abundantly
pubescent, or scabrous with the persistent bases of trichomes, provided with many
sharp, straight, conical spines to 8 mm. long; and with appressed scales at the base,
these mostly bicolorous, dark brown with a narrow, whitish margin, the margin pro-
vided with a nearly continuous line of dark brown denticulations; lamina subtripin-
nate, the tissue glabrous, herbaceous; rachis yellow-brown to stramineous, aculeate
STOLZE: FERNS OF GUATEMALA 129
(at least on the lower portion), densely hirsute to scabrous as on the petiole, scales
lacking; pinnae sessile; costae lacking spines and scales, but copiously hirsute;
pinnules sessile, cut nearly to the costule into linear, subfalcate segments, the
sinuses acute to quadrangular; costules densely hirsute, sparsely provided with
tawny to dark brown, flat or sub-bullate scales on the abaxial side; ultimate seg-
ments crenate or with deep lobes (these often bidentate), obtuse to subacute, the
midribs sparsely strigose or glabrescent on the adaxial side, abaxially hirsute and
provided with widely scattered tawny, bullate scales; veins commonly once-forked;
sori commonly inframedial between midrib and segment margin; receptacle small,
elongated to subglobose; paraphyses longer than the sporangia.
In his revision, Riba recognizes two varieties of the species, with
var. guatemalensis differing from the typical variety in its smaller
leaf parts and in the acute to narrowly quadrangular (rather than
broadly quadrangular) sinuses between the segments.
Trichipteris schiedeana (Presl) Tryon, Contr. Gray Herb. 200: 44.
1970. Alsophila schiedeana Presl, Tent. Pterid. 62. 1836. Cyathea
schiedeana (Presl) Domin, Pteridophyta 263. 1929.
In dense forests and wet thickets, on slopes and along rivers and
streams, 50-1,400 m. Alta Verapaz; Huehuetenango; Izabal; Que-
zaltenango; El Quiche. Southern Mexico; British Honduras; Hon-
duras; El Salvador; Costa Rica; Panama. Palma de montafia (Que-
zaltenango, fide Steyermark).
Stem to 5 m. tall; leaves to 3 m. long; petiole gray- to yellow-brown, with sharp,
straight, conical spines (to 8 mm. long) throughout, or merely muricate near the
lamina, provided with scurf composed of brownish squamulae, occasionally sparsely
strigose adaxially, and toward the base with appressed scales to 2 cm. long, these
commonly lanceolate, attenuate, dark brown, concolorous (very rarely with a nar-
row, whitish margin); lamina bipinnate-pinnatisect, tissue glabrous, firm-membra-
naceous; rachis yellow-brown to stramineous, sharp-spiny to muricate, strigose
(densely so adaxially) and often somewhat scurfy like the petiole, provided with
scattered dark brown scales, many of these minute; pinnae sessile or some basal
ones short-stalked; costae lacking spines or scurf, strigose (densely so adaxially),
sparsely to abundantly provided with minute, dark brown scales abaxially; pinnules
sessile, cut nearly to the costule; costules provided with abundant, dark brown,
bullate scales, strigose adaxially, occasionally sparse-strigose abaxially; ultimate
segments straight or subfalcate, oblong, 2-2.5 times as long as broad (larger ones 3-
4.5 mm. wide), with broadly rounded apices, margins commonly subentire (some-
times crenulate), essentially plane, the sinuses acute, midribs with a few dark
brown, bullate scales abaxially; veins simple to once-forked; sori borne at or near
the vein fork, medial to supramedial between midrib and segment margin; recep-
tacle small, subglobose; paraphyses much shorter than the sporangia.
Trichipteris ursina (Maxon) Tryon, Contr. Gray Herb. 200: 44.
1970. Alsophila ursina Maxon, J. Wash. Acad. Sci. 34: 48. 1944
(type from Stann Creek Valley, British Honduras, Gentle 3197).
130 FIELDIANA: BOTANY, VOLUME 39
In deep forests, sea level to 100 m. British Honduras; Nicaragua;
Costa Rica.
Stem short, to 15 cm. long; leaves to 1.5 m. long and 0.25 m. broad; petiole to 25
cm. long, brown, densely covered with spreading scales, these to 2 cm. long, lanceo-
late, attenuate, dark, lustrous brown, concolorous or with narrow white margins;
lamina pinnate-pinnatisect, tissue glabrous, firm-membranaceous to chartaceous;
rachis densely scaly throughout (and thickly strigose, but trichomes obscured by
the scales), the scales similar to those of the petiole, but becoming filiform and less
abundant toward the apex; pinnae sessile (or basal ones short-stalked), spreading at
broad angles from the rachis, incised nearly to the costa; costae provided with dark
brown, filiform scales (these often sub-bullate abaxially) and with abundant, stiff,
appressed, pale trichomes adaxially; ultimate segments oblong, subfalcate, broadly
rounded at apex, the margins subentire to crenulate, plane or slightly undulate,
glabrous, or thinly minute-pilose on midrib and veins abaxially; veins simple or,
more commonly, once-forked; sori medial to supramedial between midrib and mar-
gin, on branched veins borne at or near the fork; receptacle flattened; paraphyses
shorter than the sporangia.
Trichipteris ursina is known only from the type (British Hon-
duras) and a few collections in Nicaragua and Costa Rica. It is one
of only a few in the genus with such reduced blade architecture, and
hence should be confused with no other species in Guatemala. In
dissection of blade it superficially resembles Cnemidaria, but can be
easily distinguished from the latter by the lack of indusia and by the
veins, which neither anastomose nor connive at the sinus.
Publication 1246
UNIVERSITY OF ILLINOIS-URBANA