OF
ILLINOIS
AT URBANA-CHAMPA1GN
BIOLOGY
Botany
NEW SERIES, NO
PTERIDOPHYTA OF PERU
Rolla M. Tryon
Robert G. Stolze
Part III
16. Thelypteridaceae
Contributed by:
Alan R. Smith
April 30, 1992
Publication 1433
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
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Botanic Gardens, Kew," 1984 edition, or 77,
>ra of Barro Colorado Island. Stanford University Press, Stanford, Calif., 943 pp.
iid T. D. Pennington. 1963. A comparison of montane and lowland rain forest in Ecuador,
ructure, physiognomy, and floristics. Journal of Ecology, 51: 567-601.
;ge among the Siona: Cultural patterns in visions, pp. 63-80. In Browman, D. L.,and R, A.
.mans, and Stars. Mouton Publishers, The Hague, Netherlands.
l"he historic tribes of Ecuador, pp. 785-821. In Steward, J. H., ed., Handbook of South American
ns. Vol. 2. The Andean Civilizations. Bulletin 143, Bureau of American Ethnology. Smithsonian
n. Washington, D.C.
ind fern allies of Guatemala. Part II. Polypodiaceae. Fieldiana: Botany, n.s., 6: 1-5
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FIELDIANA
Botany
NEW SERIES, NO. 29
PTERIDOPHYTA OF PERU
Rolla M. Tryon
Department of Biology
University of South Florida
Tampa, Florida 33620-5150
Robert G. Stolze
Associate Curator
Department of Botany
Field Museum of Natural History
Chicago, Illinois 60605-2496
Part III
16. Thelypteridaceae
Contributed by:
Alan R. Smith
University Herbarium
University of California
Berkeley, California 94720
Accepted December 2, 1991
Published April 30, 1992
Publication 1433
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
© 1992 Field Museum of Natural History
Library of Congress Catalog Card Number: 92-81361
ISSN 00 15-0746
PRINTED IN THE UNITED STATES OF AMERICA
Table of Contents
List of Illustrations
INTRODUCTION
ACKNOWLEDGMENTS
16. THELYPTERIDACEAE . 2
Macrothelypteris 3
Thelypteris 5
subg. Thelypteris 6
subg. Amauropelta 9
subg. Cyclosorus 39
subg. Steiropteris 46
subg. Goniopteris 52
subg. Meniscium 66
MAP OF PERU 77
INDEX TO NAMES . .78
1 1. Macrothelypteris:
1 2. Thelypteris, subg.
tris\ Dryopteris (=
3. Thelypteris, subg.
dulosolanosa . . .
4. Thelypteris, subg.
5. Thelypteris, subg.
losa
6. Thelypteris, subg.
7. Thelypteris, subg.
iana .
M. torresiana 4
Thelypteris: T. palus-
•- Thelypteris) tremula . . 1
Amauropelta: T. glan-
8
38
Cyclosorus: T. depilata
Steiropteris: T. valdepi-
Goniopteris: T. biolleyi
Meniscium: T. maxon-
47
53
72
in
Back cover: Thelypteris valdepilosa
PTERIDOPHYTA OF PERU
Part III
16. Thelypteridaceae
Introduction
Due to unforeseen delays, this third part of the
"Pteridophyta of Peru" has been published sub-
sequent to Part IV, Dryopteridaceae, but the for-
mat follows that of all previously published parts.
Format is explained in Part I (Fieldiana: Botany,
n.s. No. 20, 1 989), and it is not necessary to repeat
it here. The publication of two more parts will
complete the project, contrary to the announce-
ment in the Introduction to Part I that the fern
flora would be divided into five parts. Part V,
nearing completion, will contain the Aspleniaceae
and Polypodiaceae. Part VI, just begun, will treat
the heterosporous ferns and the "fern allies," and
also will include a section dealing with the bio-
geography.
The authors are extremely fortunate to have had
the cooperation of Dr. Alan Smith, University of
California, Berkeley, who contributed this com-
prehensive treatment of the Thelypteridaceae. Not
only is Dr. Smith the acknowledged authority on
the family, but he recently produced the treatment
of Thelypteroideae for the Flora of Ecuador (1983);
thus, even before beginning work on the Thelyp-
teridaceae of Peru, he had a good understanding
of Andean species. Two genera and 1 02 species
are recognized in Peru, 1 9 species of which are
described herein as new. In the section labeled
"Comments" following subg. Amauropelta, there
are descriptions of two of these novelties that came
to the author's attention after the manuscript had
been completed.
The type of each name has been determined
when possible, and an effort has been made to see
the holotype or at least type photographs or au-
thentic material. Voucher specimens cited in the
legends are from Peru unless otherwise indicated.
Original drawings illustrate diagnostic features of
genera and subgenera, and some of the species.
Voucher specimens cited in the legends are from
Peru unless otherwise indicated. The map of Peru
at the end of the text shows the Departments and
indicates the sequence of the collection citations.
The nomenclature of the genera and species is
not intended to be complete. Synonyms are listed
when they are considered useful and when the type
of the name of a species or infraspecific taxon is
from Peru.
Abbreviations of periodical publications gen-
erally follow the system of Botanico-Periodicum-
Huntianum (1968), and abbreviations of authors'
names and of books generally follow Taxonomic
Literature by Stafleu and Cowan (TL-2, 1976 et
seq.\
Acknowledgments
The original drawings were contributed by Zori-
ca Dabich, scientific illustrator at Field Museum.
We express our sincere appreciation for her out-
standing work and for her patience and under-
standing. We also appreciate comments on the
manuscript by several reviewers.
We are indebted to Blanca Leon, of the Univer-
sidad Nacional Mayor de San Marcos (USM) in
Lima, for verification of the identity of duplicate
specimens cited herein from HUT and USM. Dr.
Smith has been unable personally to examine these
during the course of his study.
We are deeply grateful to Dr. Rolf Singer, Re-
search Associate in the Department of Botany at
Field Museum, for checking the accuracy of the
Latin descriptions and diagnoses for the 19 new
species published herein.
FIELDIANA: BOTANY, N.S., NO. 29, APRIL 30, 1992, PP. 1-80
We extend our thanks to the officers of the fol-
lowing institutions for granting loans of their ma-
terial or allowing us to examine specimens in their
herbaria: Botanical Institute, University of Aar-
hus, Denmark (AAU); Botanischer Garten und Bo-
tanisches Museum, Berlin-Dahlem, Berlin (B);
Bailey Hortorium, Cornell University, Ithaca (BH);
The Natural History Museum [formerly British
Museum (Natural History)], London (BM); Jardin
Botanique National de Belgique, Meise (BR); Bo-
tanical Museum, Copenhagen (c); Dudley Her-
barium of Stanford University (DS); Field Museum
of Natural History, Chicago (F); Conservatoire et
Jardin Botaniques de la ville de Geneve (G); Bo-
tanical Museum, Goteborg (GB); Harvard Univer-
sity Herbaria, Cambridge, Mass.— most Gray
Herbarium (GH), some Arnold Arboretum (A);
Herbarium Truxillense, Universidad Nacional de
Trujillo, Trujillo, Peru (HUT); Royal Botanic Gar-
dens, Kew, England (K); Rijksherbarium, Leiden,
The Netherlands (L); Botanical Museum, Lund
(LD); Botanische Staatssammlung, Munchen (M);
University of Michigan, Ann Arbor, Mich. (MICH);
Missouri Botanical Garden, St. Louis (MO); New
York Botanical Garden, New York (NY); Museum
National d'Histoire Naturelle, Paris (P); Swedish
Museum of Natural History, Stockholm (s); Her-
barium, University of Tennessee, Knoxville (TENN);
Institute of Systematic Botany, Utrecht (u); Uni-
versity of California, Berkeley (uc); Botanical Mu-
seum, Uppsala (UPS); United States National Her-
barium, Smithsonian Institution, Washington,
D.C. (us); Museo de Historia Natural "Javier Pra-
do" de Universidad Nacional Mayor de San Mar-
cos, Lima, Peru (USM); Herbario Nacional de Ven-
ezuela, Caracas (VEN); Naturhistorisches Museum,
Vienna (w); and Institiit fur Systematische Bota-
nik, Universitat, Zurich (z).
This project has been supported in part by grant
BSR-85- 16358 from the National Science Foun-
dation, Systematic Biology Program. The work
would not have been possible without this assis-
tance. However, any opinions and conclusions ex-
pressed are those of the authors and do not nec-
essarily reflect the views of the Foundation.
Family 16. THELYPTERIDACEAE
Thelypteridaceae Pic.-Ser., Webbia 24: 7 1 1 . 1 970.
TYPE: Thelypteris Schmidel.
Stem erect or decumbent, stout to slender, bear-
ing usually pubescent scales and many fibrous or
sometimes rather few thick roots, dictyostelic.
Leaves small to large (0. 1-5 m), circinate in ver-
nation, monomorphic to less often subdimorphic,
rarely strongly dimorphic, if dimorphic then the
fertile erect and with smaller segments than the
sterile, clustered to widely spaced. Petiole lacking
stipules, not articulate to stem, at base with two
lunate vascular bundles, these fusing distally into
one U-shaped bundle. Lamina simple to often
1 -pinnate or 1 -pinnate-pinnatifid, infrequently
2-pinnate or up to 3-pinnate-pinnatifid. Indument
of trichomes, these commonly acicular, furcate,
stellate, capitate-glandular, or septate, scales un-
commonly also present on axes but never on lam-
inar tissue. Veins free to fully anastomosing, the
areolae lacking included veinlets or with a single
excurrent veinlet. Sori borne on veins on the ab-
axial surface of lamina, round to occasionally ob-
long or elongate, sometimes arcuate at the juncture
of cross- veins, very rarely the sporangia acrosti-
choid; paraphyses absent or occasionally present;
indusia well developed and round-reniform to
small and spathulate, or lacking. Sporangia with
a 3-rowed stalk, annulus vertical, interrupted by
the stalk. Spores bilateral, monolete, surface re-
ticulate or with more or less connected, winglike,
often perforate ridges, or surface variously ver-
rucate to papillate or echinate (spores globose and
trilete in the Old World Trigonospord). x = 27,
29-36.
Genera in this family are variously circum-
scribed, with all species placed in a single genus
Thelypteris, or the family subdivided into about
30 genera. If an extreme view is taken, nine genera
are known from Peru (i.e., Amauropelta, Amphi-
neuron, Christella, Cyclosorus, Goniopteris, Mac-
rothelypteris, Meniscium, Steiropteris, and The-
lypteris sensu strictd), but I adopt a conservative
classification, with one introduced species in the
genus Macrothelypteris and all others from the re-
gion in Thelypteris. The family is subcosmopoli-
tan, with most species in tropical and subtropical
regions, and is characterized by having two lunate
vascular bundles in the petiole, acicular or other
kinds of trichomes on the leaves, generally bilat-
eral spores with a broad perispore, generally round
sori on the veins abaxially, and chromosome base
numbers varying from 27 to 36 (29, 35, and 36
being the only numbers found in native Neotrop-
ical species).
References
CHRISTENSEN, C. 1913. A monograph of the ge-
nus Dryopteris I. The tropical American pin-
FIELDIANA: BOTANY
natifid-bipinnatifid species. Kongel. Danske Vi-
densk. Selsk. Skr., Naturvidensk. Afd., ser. 7,
10: 53-282.
HOLTTUM, R. E. 1971. Studies in the family The-
lypteridaceae III. A new system of genera in the
Old World. Blumea 19: 17-52.
SMITH, A. R. 1983. Polypodiaceae-Thelypteroi-
deae. In Harling, G. and B. Sparre, eds., Flora
of Ecuador, no. 18: 18-148.
Key to Genera of Thelypteridaceae
a. Lamina 2-pinnate-pinnatifid; axes adaxially lacking grooves I. Macrothelypteris
a. Lamina simple to 1 -pinnate-pinnatifid, rarely 2-pinnate; axes adaxially grooved ... II. Thelypteris
I. Macrothelypteris
Macrothelypteris (H. Ito) Ching, Acta Phytotax.
Sinica 8: 308. 1963. Figure 1.
Thelypteris sect. Macrothelypteris H. Ito in Nakai &
Honda, Nov. Fl. Jap. 4: 141. 1939. TYPE: The-
lypteris oligophlebia (Baker) Ching = Macrothe-
lypteris torresiana (Gaud.) Ching.
Thelypteris subg. Macrothelypteris (H. Ito) A. R. Sm.,
Phytologia 34: 233. 1976.
Terrestrial. Stem short-creeping, thick, 10 mm
in diameter. Lamina 2-pinnate-pinnatifid nearly
throughout, broadest at the base, apex gradually
reduced; pinnae 1 -pinnate-pinnatifid, sessile or
stalked; costae not grooved adaxially; buds and
aerophores absent. Veins free, often forked, tips
not reaching the margin. Indument abaxially of
unbranched septate trichomes mostly over 1 mm
long, rachis and costae lacking scales. Sori round,
medial to supramedial; indusia small, less than 0.3
mm in diameter, often obscured in mature sori;
sporangial capsules bearing short-stipitate glands.
Spores with coarse, more or less connected, per-
forate ridges and a finely reticulate surface, x =
31.
There is one widely naturalized species in trop-
ical and subtropical America. It can be confused
in gross aspect more with such dryopteroid genera
as Ctenitis, Megalastrwn, and Lastreopsis than with
Thelypteris. However, Macrothelypteris agrees with
Thelypteris in having acicular trichomes, two vas-
cular bundles in the petiole, and the lower base
chromosome number.
There are ca. 10 species of Macrothelypteris na-
tive to tropical and subtropical Asia, Pacific Is-
lands, Queensland, and Africa.
Reference
HOLTTUM, R. E. 1969. Studies in the family The-
lypteridaceae. The genera Phegopteris, Pseudo-
phegopteris, and Macrothelypteris. Blumea 17:
5-32.
1. Macrothelypteris torresiana (Gaud.) Ching,
Acta Phytotax. Sinica 8: 310. 1963. Figure 1.
Polystichum torresianum Gaud, in Freyc., Voy. Ura-
nie. 333. 1828. TYPE: Mariana Islands, Gaudi-
chaud (holotype, P).
Aspidium uliginosum Kunze, Linnaea 20: 6. 1847.
LECTOTYPE (chosen by Morton, Contr. U.S.
Natl. Herb. 38: 219. 1973): Cult. Hort. Bot. Leip-
zig, origin Java (BR; photo, us).
Dryopteris uliginosa (Kunze) C. Chr., Index fil. suppl.
3: 100. 1934.
Thelypteris torresiana (Gaud.) Alston, Lilloa 30: 111.
1960.
Stem short-creeping, scales brown, shiny, lan-
ceolate, setose on margins and surface. Leaves few,
approximate, 60-150 cm long. Lamina charta-
ceous, 2-pinnate-pinnatifid, proximal pair of pin-
nae the longest or sometimes greatly reduced and
the next pair the longest. Petiole to 85 cm x 3-
12 mm, stramineous or greenish and scaleless
above the base, often glaucous. Rachis glabrous or
with sparse septate trichomes abaxially. Pinnae
sessile to stalked, to 35 x 17 cm, 1 -pinnate-pin-
natifid, the pinnules short-stalked to mostly slight-
ly adnate, 2-8 x 0.8-2.5 cm, incised almost to
costule into oblique segments 2-4 mm wide, these
entire to crenate or pinnatifid. Aerophores absent.
Buds lacking. Veins forked or unbranched. Indu-
ment of costae and veins abaxially of scattered,
spreading, septate trichomes mostly 1-2 mm, axes
and sometimes laminar tissue with inconspicuous
capitate glands 0. 1 mm, costal scales lacking, lam-
ina adaxially with sparse setae to 0.5 mm on cos-
tules and veins. Sori medial, round, indusia 0.2-
0.4 mm in diameter, hidden by mature sporangia,
glabrous or sparsely setose, receptacle glabrous,
sporangia with capitate glands near annulus.
Edges of lowland forests, 200-800 m, Amazo-
nas, San Martin, Loreto, Huanuco, and Madre de
Dios.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
FIG. 1 . Macrothelypteris torresiana: a, portion of stem and petiole; b, lamina; c, portion of costa with pinnule
base, adaxial side, two basal segments; d, distal portion of rachis and pinna base, abaxial side, with basal acroscopic
pinnule, (a, c, d from Anderson 9958, Brazil, uc; b from Stolze 1498, Costa Rica, F.)
FIELDIANA: BOTANY
Southeastern United States; Antilles; Mexico to
Panama; Colombia to Bolivia; southern Brazil;
northern Argentina; Paraguay; tropical and sub-
tropical Africa and Asia; Pacific Islands. Adven-
tive in the New World tropics and subtropics. The
earliest collection seen from Peru was made in
1978.
Amazonas: Prov. Bagua, Chiriaco to Puente Venezue-
la, 43 km NE of Chiriaco, Barbour 4380 (MO, USM), 4404
(MO, USM). San Martin: San Martin, 15 km E of Shapaja
on road to Chazuta, Knapp 7876 (F, MO). San Martin,
Km 28 of Tarapoto-Yurimaguas road, Knapp & Mallet
8395 (MO). Loreto: Prov. Alto Amazonas, Andoas Pas-
taza, Vdsquez & Jaramillo 791 (F, MO), 792 (MO). Huanu-
co: Prov. Leoncio Prado, Dist. Rupa Rupa, Jacintillo,
Rio Monzon, Schunke V. 10367 (F, MO, USM). Along
road from Tingo Maria to Monson (Monzon), Croat
57922 (MO, USM). Madre de Dios: Prov. Manu, Parque
Nacional Manu, Rio Manu, Cocha Cashu Station, R.
Foster 11429 (F, USM).
II. Thelypteris
Thelypteris Schmidel, Icon. pi. (ed. Keller), 3rd
page, t. XI text. 1763 (nom. cons.). TYPE:
Thelypteris palustris Schott (Acrostichum the-
lypteris L.).
Terrestrial or epipetric, rarely epiphytic. Stem
creeping to ascending or erect, rarely scandent.
Lamina 1 -pinnate to 1 -pinnate-pinnatifid, rarely
simple or 2-pinnate, proximal pinnae reduced or
not, apex commonly gradually reduced, infre-
quently abruptly reduced and pinnalike; pinnae
entire to deeply pinnatifid, rarely 1 -pinnate, sessile
or short-stalked; costae grooved adaxially; buds
absent or present in axils of pinnae; aerophores
absent to present at pinna bases, these tuberculi-
form or scalelike. Veins free to regularly anasto-
mosing, commonly simple, rarely 1-2-forked,
reaching the margin. Indument various abaxially,
often of simple or branched trichomes on axes and
lamina, trichomes infrequently septate, rachis and
costae with or usually without scales. Sori round,
oblong, or elongate along veins, commonly infra-
medial to supramedial; indusia round-reniform,
large (ca. 1 mm in diameter) and persistent or
small (less than 0.3 mm), occasionally fugacious,
sometimes absent; sporangial capsule glabrous or
occasionally setose or bearing inconspicuous glands
from the stalk, x = 27, 29, 31-36.
As treated here, Thelypteris comprises ca. 875
species and is subcosmopolitan, with greatest con-
centration in the tropics. Nearly 300 species are
found in the New World. The genus is usually
divided into numerous subgenera or sections that
are sometimes treated as genera. These groups,
although mostly natural and probably monophy-
letic, are definable only by a combination of char-
acters. Some of the characters concern minute de-
tails of glands and trichomes and require 30 x
magnification or greater for observation. Others
require that complete specimens be at hand, in-
cluding the proximal part of the lamina and the
stem. Hybridization between the groups is known
or suspected and transitional species and species
groups occur. Some of the best characters for cir-
cumscribing the groups involve chromosome
number and characters of spores best seen with
the scanning electron microscope. Because of the
practical problems of identification, as well as un-
certainty over degree of relatedness, a conservative
classification is adopted here.
Thelypteris is probably one of the most poorly
collected fern genera in Peru, as many species are
known only from the type or just a few collections.
This is probably because many species are very
similar in lamina dissection, and collectors gen-
erally avoid them after making a few token gath-
erings. It would not be surprising if the number of
species in the country grows by 25% or more as
additional areas become open to collectors and as
specialists visit remote and species-rich areas.
In Peru, subgenera Amauropelta and Thelypteris
s.s. stand somewhat apart from the other subgen-
era by virtue of their distinctive spores, chromo-
some number, and venation; species of the former
also have reduced proximal pinnae, although these
often go unnoticed or uncollected because they are
mere nubbins less than a few mm long. The other
four subgenera form a large natural assemblage
that could comprise a separate genus, Cyclosorus.
The descriptions of species are drawn from Pe-
ruvian specimens except in the cases where the
material is sparse or otherwise inadequate. In those
cases, the descriptions are amplified by material
from elsewhere in the range, particularly Ecuador.
References
Listed under the appropriate subgenus.
Key to Subgenera of Thelypteris
a. At least some of the trichomes on the rachis, costae, and/or stem apex scales forked or stellate . . .
(species 64, 65, 68-81) . . . He. Thelypteris subg. Goniopteris
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
Trichomes all unbranched (fasciculate in 3 species of subg. Amauropeltd), acicular, unicellular or
multicellular, rarely trichomes absent b
b. Lamina 1 -pinnate, the pinnae entire or margin undulate or serrate, or pinnae incised less than
halfway to the costa; veins regularly anastomosing, the areoles formed by uniting cross- veins 3-
25-seriate between costa and margin c
c. Sori mostly discrete; son indusiate
(species 66, 67, 82) . . . He. Thelypteris subg. Goniopteris
c. Sori mostly confluent where the veins anastomose, thus oblong or lunate (except sometimes
in T. lingulata); sori exindusiate (species 83-98) . . . Hf. Thelypteris subg. Meniscium
b. Lamina 1 -pinnate-pinnatifid or rarely 2-pinnate, the pinnae usually incised half their width or
more; veins meeting margin above the sinus, connivent at the sinus, or with 1 or 2 pairs anas-
tomosing below the sinus d
d. Veins 1-2-forked in the ultimate segments; proximal pinnae not or only slightly shorter than
the longest pair; lamina 2-pinnate (species 1) ... Ha. Thelypteris subg. Thelypteris
d. Veins usually simple in the ultimate segments; proximal pinnae much shorter than the longest
or not; lamina usually 1 -pinnate-pinnatifid, rarely 1 - or 2-pinnate e
e. Lamina with 1 or usually several to many pairs of reduced proximal pinnae, lowermost
auriculiform, glanduliform, or hastate; veins free, meeting margin above the sinus or rarely
at the sinus (species 2—47) . . . lib. Thelypteris subg. Amauropelta
e. Lamina without reduced proximal pinnae, or if reduced pinnae present, then veins united
below sinus f
f. Cartilaginous keel or minutely pubescent false vein running from sinus toward costa,
occasionally lacking; indusia present or absent; aerophores peglike or scalelike at pinna
bases (species 57-63) . . . Hd. Thelypteris subg. Steiropteris
f. Cartilaginous keel lacking; indusia present, persistent; aerophores absent at pinna bases
(species 48-56) . . . He. Thelypteris subg. Cyclosorus
la. Thelypteris subg. Thelypteris. Figure 2.
Lamina 1 -pinnate-pinnatifid to 2-pinnate; prox-
nal pinnae not reduced or only slightly reduced
jlative to the longest pair, distal pinnae gradually
hortened, the lamina with a confluent pinnatifid
pex; aerophores absent; buds absent. Veins from
djacent segments meeting margin above the si-
us, simple or often 1-2-forked. Indument of acic-
lar, often septate crisped trichomes, also of tan,
roadly ovate scales along the abaxial costae, often
nth sessile or stipitate glands. Sori round, indu-
iate; sporangia glabrous. Spores various, with a
;ticulate to verrucate, papillate, or echinate peri-
pore, x = 35.
Subgenus Thelypteris comprises only 2 wide-
pread species: T. palustris, north-temperate in
:anada and eastern United States, Europe, and
^sia; and T. confluens (Thunb.) Morton, south-
smperate in Argentina, Africa, and India to New
Zealand and New Guinea.
teferences
rERNALD, M. L. 1929. A study of Thelypteris
palustris. Rhodora 31: 27-40.
TRYON, A. F., R. TRYON, AND F. BADRE. 1980.
Classification, spores, and nomenclature of the
Marsh fern. Rhodora 82: 461-474.
1. Thelypteris palustris Schott, Gen. fil., sub tab.
10. 1834. Figure 2a-c.
Acrostichum thelypteris L., Sp. pi. 2: 1071. 1753. TYPE:
"in Europae septentrionalioris paludibus" (typi-
fication uncertain).
Dryopteris thelypteris (L.) A. Gray, Manual, ed. 1 . 630.
1848.
Stem long-creeping, 1-3 mm in diameter, scales
few, tan, glabrous or glandular on margin. Leaves
usually several cm apart, monomorphic or slightly
dimorphic with the fertile more erect, narrower,
and with somewhat contracted pinnae and seg-
ments, 20-90(-120) cm long. Lamina membra-
naceous to chartaceous, 1 -pinnate-pinnatifid to
2-pinnate, proximal pinnae not or only slightly
reduced. Petiole 9^15(-60) cm x 1-3 mm, tan to
stramineous above the blackened base, at base gla-
brous or with sparse, tan, ovate, glabrous scales.
Rachis with crispate, septate trichomes to 1 mm
abaxially, also with a few deep red glands ca. 0. 1
mm. Pinnae sessile or short-stalked to 1 mm,
FIELDIANA: BOTANY
FIG. 2. Subgenus Thelypteris. Thelypteris palustris: a, stem and leaf; b, costa and base of pinna segment, abaxial
side; c, rachis and base of pinna, adaxial side. Dryopteris (= Thelypteris) tremula: d, lamina, (a, c from Stolze 279,
Illinois, U.S.A., F, b from Camp, in 1897, Michigan, U.S.A., F; d from Arsene, in 1910, Mexico, uc.)
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
FIG. 3. Subgenus Amauropelta. Thelypteris glandulosolanosa: a, stem and petiole bases; b, lamina; c, pinna
segments, abaxial side; d, sorus (sporangia removed), (a, c, d from Woytkowski 172, uc; b, from Feuerer 8 127 a,
Bolivia, F.)
FIELDIANA: BOTANY
2-10(-12) x 0.5-2(-2.5) cm, deeply pinnatifid to
1 -pinnate and the pinnules entire, crenate, or shal-
lowly lobcd. Aerophores absent. Buds lacking.
Veins 4-10 pairs per segment, often 1-2-forked,
especially in sterile fronds. Indument of costae and
veins abaxially of acicular and crispate, septate
trichomes 0.3-1 mm, also with a few deep red,
sessile or short-stalked, pear-shaped glands, costal
scales tan, ovate, with a few similar glands on
margin, adaxially glabrescent or with scattered tri-
chomes on costules and veins, laminar tissue gla-
brous. Sori medial, round, with a usually large
indusium, receptacle and sporangia glabrous or the
latter sometimes glandular.
Growing on floating mats with lacustrine veg-
etation, elsewhere in the range from marshes and
swamps, 2000 m, Amazonas.
Eastern Canada and United States; Cuba; Ber-
muda; perhaps Mexico; north-temperate Europe
and Asia.
The sole Peruvian collection is sterile and with-
out stem, so the precise identity is uncertain. It is
clearly assignable to subg. Thelypteris on the basis
of the crispate pubescence, forked venation, tan,
ovate costal scales, and pear-shaped glands at the
scale margins and along rachis, costae, and cos-
tules abaxially. This specimen is larger (extreme
measurements in description above) and more dis-
sected than most specimens of T. palustris from
the north-temperate areas, being fully 2 -pinnate
with shallowly lobed pinnules on the proximal
pinnae. In these characters, it matches some spec-
imens from southeastern United States and es-
pecially Dryopteris tremula Christ (fig. 2d), known
only from the type and topotypes collected by Ar-
sene in Michoacan, Mexico, in 1910. The Arsene
collections resemble some specimens of Thelyp-
teris palustris from Louisiana, where he also col-
lected, and mislabeling is a possibility. Additional
collections are necessary to determine whether the
Peruvian species is really T. palustris, and if so,
to which of several varieties it is allied.
Amazonas: Bongara Prov., Pomacocha, D. Smith 6020
(MO, USM).
lib. Thelypteris subg. Amauropelta.
Thelypteris subg. Amauropelta (Kunze) A. R. Sm.,
Amer. Fern J. 63: 121. 1973. Figure 3.
Amauropelta Kunze, Farnkr. 1: 109. 1843. TYPE:
Amauropelta breutelii Kunze = Thelypteris lim-
bata (Sw.) Proctor.
Lamina 1 -pinnate to 1 -pinnate-pinnatifid, rare-
ly 2-pinnate; proximal pinnae usually gradually to
abruptly reduced, mostly 4 or more pairs (rarely
only 1), lowermost pairs sometimes glanduliform,
distal pinnae gradually shortened, the lamina with
a confluent pinnatifid apex; aerophores at pinna
bases often present; buds absent in all but 2 of our
species. Veins from adjacent segments usually
meeting margin above the sinus, rarely connivent
at the sinus, never connivent or united below si-
nus. Indument of hamate or acicular trichomes,
the latter sometimes septate. Sori round to less
often oblong or elongate along the veins, indusiate
to exindusiate; sporangia glabrous or rarely setose.
Spores with a usually finely reticulate perispore.
x = 29.
Subgenus Amauropelta comprises over 200
tropical American species, with one species in Ha-
waii and a few in Africa, Madagascar, and the
Mascarene Islands. It has been subdivided by Smith
(1974) into nine sections, based on the following
characters: stem habit; the type and distribution
of trichomes and glands; and the presence or ab-
sence of costal scales, aerophores, and indusia.
The number of new taxa (13 species) is some-
what surprising, especially since the Ecuadorian
species have recently been treated (Smith, 1983).
Less overlap was encountered between the species
of Peru and Ecuador than was expected, with 30
of 47 species of the subgenus in Peru also occurring
in Ecuador and 30 of 60 found in Ecuador also in
Peru. In general, in subg. Amauropelta there are a
number of very widespread species, often occur-
ring from southern Mexico to Bolivia, and an equal
number of highly restricted and poorly known spe-
cies. Thirteen species are known thus far only from
Peru.
Of the Peruvian species, the most difficult group
is sect. Lepidoneuron, comprising those species with
dark scales along the abaxial costae. This group is
in need of monographic revision, and difficulty
may be encountered in species identification.
References
CHRISTENSEN, C. 1907. Revision of the Ameri-
can species of Dryopteris of the group of D. op-
posita. Kongel. Danske Vidensk. Selsk. Skr.,
Naturvidensk. Afd., ser. 7, 4: 249-336.
SMITH, A. R. 1974. A revised classification of
Thelypteris subgenus Amauropelta. Amer. Fern
J. 64: 83-95.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
Key to Species of subg. Amauropelta
a. Trichomes fasciculate on costae and rachis abaxially, appearing stellate; prominent peglike or scalelike
aerophones up to 5 mm long at base of costae and smaller ones at base of costules b
b. Scales scattered, dark brown on costae abaxially 20. T. phacelothrix
b. Scales absent on costae abaxially or if present, tan c
c. Lamina abaxially with red to yellow sessile glands 19. T. thomsonii
c. Lamina abaxially lacking glands 18. T. canadasii
a. Trichomes not fasciculate (evenly spaced, not appearing stellate) or absent; aerophores present or
absent at base of costae, absent at base of costules d
d. Lamina abaxially with hamate trichomes on tissue and/or axes; adaxial surfaces between veins
bearing numerous appressed trichomes ca. 0.1-0.2 mm; lamina eglandular abaxially e
e. Pinnae entire or nearly so, the largest 0.5-1.5 cm long; buds present in axils of distal pinnae
10. T. pusilla
e. Pinnae shallowly to deeply lobed or pinnatifid, largest ones greater than 1.5 cm long; buds
absent (except T. linkiana) f
f. Costae abaxially with persistent, spreading or ascending, castaneous or blackish scales . . . . g
g. Rachis abaxially with numerous, persistent, spreading scales 3-5 mm long
1 2. T. leoniae
g. Rachis abaxially lacking persistent scales, or scales sparse, not noticeably spreading . . h
h. Sori indusiate, indusium setose 34. T. hutchisonii
h. Sori exindusiate, receptacle occasionally setose 28. T. rudis
f. Costae abaxially lacking persistent scales, if scales present then appressed, dull, light tan
i
i. Sori round, indusiate (look carefully in mature sori!) j
j. Proximal pinnae gradually reduced, reduced pinnae numerous, ca. 7-11 pairs; sori
medial 6. T. scalaris
j. Proximal pinnae abruptly reduced, 6 pairs or fewer; sori supramedial to inframarginal
k
k. Glands numerous, short-stipitate (0.05-0.1 mm), present on costae, veins, and
lamina abaxially; indusia glandular 9. T. pt arm id for mis
k. Glands sparse or absent abaxially; indusia setose 5. T. oligocarpa
i. Sori round to oblong or elongate, exindusiate 1
1. Pinnae less than 3(-5) cm long; buds lacking m
m. Sori oblong or elongate along the veins; proximal reduced pinnae 1 or 2 pairs,
these V3-V2 the length of the longest pinnae 2. T. aspidioides
m. Sori round or slightly oblong; proximal reduced pinnae 3 or more pairs, the
smallest often less than '/10 the length of the longest pinnae n
n. Aerophores peglike at pinna bases; abaxial costal trichomes 0.2-0.3 mm long;
petiole base and stem apex scales numerous, shining, with numerous stiff setae
on margin and surface 1 1 . T. brachypus
n. Aerophores absent; abaxial costal trichomes mostly 0.3-0.5 mm long; petiole
base and stem apex scales sparse, dull, sparsely and laxly ciliate
7. T. andicola
1. Pinnae more than 5 cm long; buds lacking or present in axils of distal pinnae . . . o
o. Sori round; sporangia setose; buds lacking 4. T. longipilosa
o. Sori elongate 1.5-3 mm along veins; sporangia usually glabrous; bud(s) in axil(s)
of distal pinnae 3. T. linkiana
d. Lamina abaxially with acicular trichomes or glabrous; adaxial surfaces glabrous or pubescent,
trichomes appressed or not; lamina glandular or eglandular abaxially p
p. Indusia present q
q. Lamina abaxially between the veins with sessile, reddish, orangish, or yellowish resinous,
hemispherical glands, these often also on indusia r
r. Sori supramedial to inframarginal, partially covered by the strongly inrolled segment
margin; veins abaxially sunken and often darkened in dried specimens; young croziers
mucilaginous; aerophores peglike or scalelike at pinna bases ... 43. T. cheilanthoides
!0 FIELDIANA: BOTANY
r. Sori medial to supramedial, not hidden by the segment margin; veins abaxially flush or
slightly raised; young croziers usually not mucilaginous; aerophores absent or peglike
s
s. Fronds 8-20 x 1.5-2.5 cm; lamina adaxially with dense trichomes 0.1-0.2 mm;
texture thin 4 1 . T. micula
s. Fronds larger than 20 x 2.5 cm; lamina adaxially glabrous or with sparse to mod-
erately dense trichomes; texture chartaceous to subcoriaceous t
t. Costae abaxially with scattered, dark, ovate-lanceolate scales to 1 mm and as-
cending unicellular trichomes mostly 0.2 mm; petiole 1-1.3 mm in diameter . . .
42. T. dudleyi
t. Costae abaxially lacking scales, or scales if present tan to stramineous, trichomes
spreading, unicellular or septate, 0.2-2.0 mm; petiole 1-10 mm in diameter . . . u
u. Costae with amorphous, mostly appressed scales; indusia ca. 1 mm in diameter
38. T. pachyrhachis
u. Costae lacking scales; indusia ca. 0.5-0.7 mm in diameter v
v. Segments short-oblong to deltate; segment margins strongly revolute; texture
chartaceous to subcoriaceous; aerophores absent or weakly developed; tri-
chomes of abaxial costae unicellular; veins fewer than 8 pairs per segment
40. T. opposite
v. Segments linear-oblong; segment margins slightly revolute to planar; texture
chartaceous; aerophores tuberculiform or peglike; trichomes of abaxial cos-
tae sometimes septate; veins 10-16 pairs per segment .... 39. T. balbisii
q. Lamina abaxially between the veins lacking sessile glands, or if glands present then these
short-stipitate, light yellowish, or if reddish then confined to costae and veins w
w. Petiole and often rachis atropurpureous, shining, glabrous x
x. Pinnae 3-8 cm, widest at the base; texture chartaceous to subcoriaceous
13. T. furva
x. Pinnae to 1 8 cm, slightly narrowed at the base (proximal segments slightly shortened);
texture thin-chartaceous 24. T. atrorubens
w. Petiole and rachis stramineous to tan or dull brownish, often pubescent y
y. Trichomes of costae and laminar tissue abaxially dense, uniformly short (0. 1 0-0. 1 5
mm); lamina adaxially with short adpressed trichomes; indusium reduced to a few
cells and less than 0. 1 mm in diameter, bearing a tuft of short trichomes
8. T. enigmatica
y . Trichomes of costae and laminar tissue abaxially sparse to dense, commonly longer
(0.2-1.0 mm); lamina adaxially (between veins) glabrous or with sparse trichomes;
indusium at least 0.2 mm in diameter, glabrous or pubescent z
z. Costae abaxially with ovate to lanceolate or amorphous scales; segment margins
strongly inrolled or not; aerophores peglike or scalelike at pinna bases aa
aa. Segment margins neither strongly inrolled nor covering sori
38. T. pachyrhachis
aa. Segment margins strongly inrolled and partially covering sori bb
bb. Costal scales numerous, bicolorous, the center dark brown, margins pale
44. T. furfuracea
bb. Costal scales sparse to moderately dense, concolorous, stramineous to
tan 43. T. cheilanthoides
z. Costae abaxially lacking scales; segment margins planar or weakly inrolled, not
or slightly covering sori; aerophores lacking cc
cc. Indusia with sparse to dense, long silky trichomes up to ca. 1 mm
16. T. glandulosolanosa
cc. Indusia lacking trichomes or if present then usually stiff, not dense, less than
0.4 mm dd
dd. Costal trichomes mostly (0.3-)0.5-1.5 mm; costules and veins adaxially
with similar stout trichomes; indusia stipitate-glandular on margin, oth-
erwise glabrous; petiole bases with at least a few, persistent, spreading,
lanceolate scales; sporangia glabrous 14. T. pilosula
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III. 1 1
dd. Costal trichomes mostly less than 0.5 mm, usually less than 0.3 mm;
costules and veins adaxially glabrous or with sparse trichomes 0.2 mm
long; indusia eglandular, short-pubescent; petiole bases lacking scales or
scales few, ovate-lanceolate; sporangia often setose 15. T. rufa
p. Indusia absent ee
ee. Sporangia minutely setulose, with trichomes less than 0. 1 mm from capsule ff
ff. Costal trichomes mostly 0.1-0.6 mm; costae abaxially with scattered linear-lanceolate
scales; receptacular trichomes 0. 1-0.5 mm gg
gg. Trichomes of rachis abaxially strongly deflexed and appressed, 0.2 mm long; costal
trichomes abaxially 0. 1-0.2 mm 23. T. cotnptula
gg. Trichomes of rachis abaxially patent, mostly more than 0.3 mm long; costal tri-
chomes abaxially mostly 0.3-0.6 mm 22. T. peruviana
fF. Costal trichomes 0. 1 mm or less; costae abaxially without scales; receptacular trichomes
absent, or if present less than 0. 1 mm hh
hh. Pinnae incised within 0.5 mm of costae; segments oblique; lamina abaxially with
sessile resinous glands 36. T. loretensis
hh. Pinnae incised to 0.5-1 mm from costae; segments spreading to suboblique; lamina
glandless 35. T. concinna
ee. Sporangia glabrous ii
ii. Scales lacking on costae abaxially jj
jj. Costae and lamina abaxially with dense, short trichomes 0.1-0.15 mm
8. T. enigmatica
jj. Costae and lamina abaxially glabrous or sparsely to moderately pubescent, the
trichomes 0. 1 5 mm or more kk
kk. Aerophores at pinna bases peglike; reduced proximal pinnae commonly 5 or
more pairs 11
11. Petiole and often rachis atropurpureous; costae abaxially moderately pu-
bescent 2 1 . T. pavoniana
11. Petiole and rachis tan to stramineous; costae abaxially glabrous or nearly
so 47. T. proboscidea
kk. Aerophores lacking or tuberculiform; reduced proximal pinnae 2-4 pairs . . .
mm
mm. Costae moderately pubescent abaxially 17. T. demissa
mm. Costae glabrous or nearly so abaxially 37. T. deflexa
ii. Scales present on costae abaxially nn
nn. Costal trichomes spreading, pluricellular, often over 1 mm; aerophores lacking or
weakly developed; costal scales stramineous to tan, with lateral walls not easily
visible, scales not clathrate oo
oo. Segment margin inrolled and partially covering the sori; sori confluent at
maturity 45. T. ruiziana
oo. Segment margin not or weakly inrolled, not covering sori; sori not confluent
at maturity 46. T. nitens
nn. Costal trichomes spreading to antrorse, unicellular, 0. 1 to occasionally more than
1 mm, or trichomes lacking; aerophores present or absent; costal scales tan to
brown, often with lateral walls darker than surficial walls (scales clathrate or sub-
clathrate) pp
pp. Costal scales tan; aerophores peglike, 1-3 mm; costae and rachis glabrous or
very sparsely pubescent 47. T. proboscidea
pp. Costal scales brown or darker; aerophores absent or tuberculiform, if peglike
less than 1 mm; costae and rachis glabrous to densely pubescent qq
qq. Lamina 2-pinnate proximally, the pinnules constricted at their base; cos-
tae glabrous or sparsely pubescent abaxially 27. T. pteroidea
qq. Lamina 1 -pinnate-pinnatifid proximally, the pinnules connected by a nar-
row to broad wing; costae glabrous to densely pubescent abaxially . . rr
rr. Costae and rachis abaxially completely lacking trichomes ss
12 FIELDIANA: BOTANY
ss. Petiole and rachis atropurpureous, shining .... 25. T. laevigata
ss. Petiole and rachis stramineous to tan, dull .... 26. T. euchlora
rr. Costae and rachis abaxially with at least a few trichomes tt
tt. Lamina relatively small, pinnae mostly less than 5 cm long and
1.5 cm wide; reduced proximal pinnae l-3(— 4) pairs
33. T. caucaensis
tt. Lamina usually large, pinnae greater than (5-) 10 cm long and 1.5
cm wide; reduced proximal pinnae more than 3 pairs (look for
glanduliform pinnae at base of lamina!) uu
uu. Costal scales decidedly clathrate, the lateral walls black and
lumina clear; costules and veins adaxially with stout hispid
trichomes 1 mm long or more 31. T. corazonensis
uu. Costal scales weakly clathrate or uniformly colored; costules
and veins adaxially glabrous or with short trichomes less than
0.5 mm vv
vv. Trichomes on costae abaxially scattered, relatively few;
lamina and veins abaxially glabrous or nearly so
32. T. brausei
vv. Trichomes on costae abaxially dense or moderately so,
relatively numerous; lamina glabrous or pubescent abax-
ially ww
ww. Costae abaxially with antrorse trichomes; lamina
abaxially between veins with a few hamate tri-
chomes below sinuses; proximal pinnae abruptly
reduced with many glanduliform pinnae
28. T. rudis
ww. Costae abaxially with trichomes mostly spreading;
lamina abaxially between veins glabrous or pubes-
cent with straight trichomes; proximal pinnae grad-
ually to abruptly reduced, with or without glandu-
liform pinnae xx
xx. Trichomes of rachis dense, usually slightly re-
flexed; costal trichomes abaxially mostly 0.2-
0.4 mm; veins with short trichomes 0.1-0.3
mm adaxially 29. T. supina
xx. Trichomes on rachis of moderate density,
spreading; costal trichomes abaxially mostly
0.5-1 mm; veins glabrous or with trichomes
mostly 0.4-1 mm adaxially
30. T. pilosohispida
2. Thelypteris aspidioides (Willd.) R. Tryon,
Rhodora69: 5. 1967.
Ceterach aspidioides Willd., Sp. pi. ed. 4. 5: 1 37. 1 8 1 0.
TYPE: Venezuela, Caracas, Bredemeyer (holo-
type, B, Herb. Willd. 19581; microfiche, uc).
Dryopteris aspidioides (Willd.) C. Chr., Index fil. 253.
1905.
Dryopteris aspidioides var. subhastata C. Chr., Kon-
gel. Danske Vidensk. Selsk. Skr., Naturvidensk.
Afd., ser. 7, 4: 287. 1907. LECTOTYPE (chosen
here): Peru, (San Martin), prope Tarapoto, Spruce
3964 (B; isolectotypes, BM!, c, G!, GH!, NY!, us!).
Stem ascending to erect, scales brown, dull,
ovate-lanceolate to lanceolate, setose on margins
and surfaces. Leaves numerous, clustered, 14-30
cm long. Lamina chartaceous, 1 -pinnate to 1 -pin-
nate-pinnatifid, proximal 1 or 2 pairs of pinnae
somewhat reduced, the lowermost 1-2.5 cm long.
Petiole 2-9 cm x 0.7-1.5 mm, brownish to tan
above the base, often with scattered, persistent
scales. Rachis glabrescent to hamate-pubescent
abaxially. Pinnae short-stalked to 1 mm, 2-5 x
0.4-0.8 cm, entire, crenate, or shallowly incised
ca. halfway to costa, often subauriculate at base,
especially acroscopically. Aerophores absent or
peglike to 0.3 mm. Buds lacking. Veins 2-4 pairs
per segment. Indument on costae, veins, and lam-
inar tissue abaxially of moderately dense, spread-
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
13
ing, mostly hamate trichomes 0. 1-0.3 mm, glands
lacking, costal scales lacking, lamina adaxially gla-
brescent or with numerous adpressed acicular tri-
chomes ca. 0.2 mm on costules, veins, and laminar
tissue. Sori medial to supramedial, oblong to elon-
gate along the veins, exindusiate, receptacle gla-
brous, sporangia glabrous.
Along streams and in stream beds, especially on
wet rocks, 200-1900 m, San Martin, Loreto, Pas-
co, Madre de Dios, and Puno.
Costa Rica and Panama; Colombia to Bolivia;
Venezuela.
San Martin: San Roque, LI. Williams 7758 (F). Tara-
poto, ca. 1 5 km from Tarapoto along road to Yurima-
guas, Martin & Plowman 1814 (F, GH). Tarapoto- Yuri -
maguas Hwy, Km 14-17, McDaniel 13811 (GH, USM).
Chazuta, Rio Huallaga, Klug 4085 (BM, F, GH, MO, NY,
uc). Pongo de Cainarachi, Rio Cainarachi, tributary of
Rio Huallaga, Klug 2658 (BM, F, G, GH, K, MO, NY). Prov.
San Martin, Dist. Tarapoto, Carretera Tarapoto-Yuri-
maguas, Km 12-15, Hickok 635 (GH). 17 km NE of
Tarapoto on road to Yurimaguas, Gentry et al. 37866
(MO, uc, USM). Loreto: Pongo de Chilcayo, Tarapoto, Ule
6518 (G). Pasco: Pichis Trail, Porvenir (as Junin), Killip
& Smith 25958 (NY). Madre de Dios: Prov. Manu, Cerro
de Pantiacolla, Rio Palotoa, 10-15 km NNW of Shin-
tuya, R. Foster et al. 10712 (F, USM). Puno: San Gaban
(as St. Gavan), Lechler 2311 (B).
3. Thelypteris linkiana (Presl) R. Tryon, Rhodora
69: 6. 1967.
Gymnogramma diplazioides Desv., Mem. Soc. Linn.
Paris 6: 214. 1827. TYPE: Hispaniola, collector
unknown (holotype, P, Herb. Desvaux).
Gymnogramma polypodioides Link, Hort. Berol. 2:
50. 1833, not Sprengel, 1827. TYPE: Cultivated
specimen, said to be from Brazil, probably er-
roneously (holotype, B?).
Grammitis linkiana Presl, Tent, pterid. 209. 1836,
nom. nov. for Gymnogramma polypodioides Link.
Dryopteris diplazioides (Desv.) Urban, Symb. antill.
4:21. 1903.
Dryopteris linkiana (Presl) Maxon, J. Wash. Acad. Sci.
14: 199. 1924.
Thelypteris diplazioides (Desv.) Proctor, Bull. Inst. Ja-
maica, Sci. Ser. 5: 59. 1953, not Ching, 1936.
Amauropelta diplazioides (Desv.) Pic.-Ser., Webbia
31:251. 1977.
Stem ascending to usually erect, apical scales
light brown, dull, ovate-lanceolate, glabrescent or
setulose on margins and surface. Leaves few, clus-
tered, 50-100(-140) cm long. Lamina herbaceous,
1 -pinnate-pinnatifid, proximal 2-5 pairs of pinnae
gradually to subabruptly reduced, the lowermost
5-15 mm long, never glanduliform. Petiole 12-25
cm x 1.5-4 mm, brownish to tan above the base,
with scattered, appressed scales at base. Rachis
glabrescent or with hamate trichomes 0.2-0.3 mm
abaxially. Pinnae sessile, 5-15 x 1.2-2.6 cm, pin-
natifid to 2-4 mm from costae, segments 3-6 mm
wide. Aerophores tuberculiform or peglike to 0.5
mm. Buds commonly present in axils of distal
pinnae. Veins 5-10 pairs per segment. Indument
on costae, veins, and sometimes laminar tissue
abaxially of sparse to moderately dense, spreading,
hamate trichomes mostly 0.15-0.3 mm, or lami-
nar tissue glabrous, glands lacking, costal scales
lacking or a few appressed linear tan scales present,
lamina adaxially with numerous adpressed acic-
ular trichomes 0. 1-0.2 mm on costules, veins, and
laminar tissue. Sori medial to supramedial, oblong
to commonly elongate 2-3 mm along veins, ex-
indusiate, receptacle glabrous, sporangia glabrous
or rarely setose.
In montane forests, slopes and ravines, 500-
2200 m, San Martin, Huanuco, Junin, Ayacucho,
and Cuzco.
West Indies; Southern Mexico to Panama; Co-
lombia to Bolivia; Guyana; Venezuela.
San Martin: Mt. Campana, near Tarapoto, Spruce 4084
(GH). Huanuco: Prov. Huanuco, Dist. Churubamba, Hda.
Exito, N slope of Rio Cayumba, Mexia 8112 (BH, F, us).
Junin: Chanchamayo Valley, C. Schunke 16 (F), 57 (F,
us), 56a (F), 60 (F, us), 509 (F), 936, (F), 937 (F), 939 (F),
952, (F), 1 352 (F). Ayacucho: Ayna, between Huanta and
Rio Apurimac, Killip & Smith 22726 (NY, us). Cuzco:
Prov. Paucartambo, Kosnipata, Pilcopata-Santa Ines,
Vargas 11314 (GH).
4. Thelypteris longipilosa (Sodiro) Reed, Phyto-
logia 17: 290. 1968.
Nephrodium longipilosum Sodiro, Anales Univ. Centr.
Ecuador 23(160): 103. 1908 [Sert. Fl. Ecuad., Ser.
II. 26]. TYPE: Ecuador, Volcan Corazon, Dec
1907, Sodiro (isotype, P; frag, and photos, us).
Stem ascending to erect, scales brown, some-
what shiny, lanceolate, setose on margins and sur-
face. Leaves few, clustered, 30-90 cm long. Lam-
ina thick-herbaceous, 1 -pinnate-pinnatifid,
proximal 4—6 pairs of pinnae gradually reduced,
the lowermost ca. 10 mm long, not glanduliform.
Petiole 4-8(-15) cm x 2-3 mm, brownish to tan
above the base, with spreading scales to 1 cm.
Rachis with dense, short trichomes 0.1-0.2 mm
mixed with long, stout trichomes 1-1.5 mm abax-
ially, with hyaline to slightly reddish trichomes
adaxially. Pinnae sessile, 5—10 x 1.2-2 cm, pin-
natifid to ca. 1.5-3 mm from costae, segments 3-
14
FIELDIANA: BOTANY
4 mm wide. Aerophores tuberculiform. Buds lack-
ing. Veins 4-8 pairs per segment, lowermost pairs
running to sinus. Indument on costae abaxially a
mixture of dense, short trichomes 0. 1-0.2 mm and
long, stout trichomes 1-2 mm, veins and laminar
tissue with dense, erect, hamate trichomes 0.2-0.3
mm, glands lacking, costal scales lacking, lamina
adaxially with numerous adpressed acicular tri-
chomes 0.2-0.3 mm on costules, veins, and lam-
inar tissue, also with scattered stout spreading or
ascending setae on costules and veins. Sori medial,
round to slightly oblong, exindusiate, receptacle
glabrous or very sparsely setose, sporangia with
numerous setae 0.1-0.2 mm long on capsule.
In cloud forest, 2300-2500 m, rare in Peru: Pas-
co.
Costa Rica and Panama; Colombia to Peru.
Pasco: Prov. Oxapampa, San Alberto, Cordillera de
Yanachuga, van der Werffet al. 8440 (MO).
5. Thelypteris oligocarpa (Willd.) Ching, Bull. Fan
Mem. Inst. Biol., Bot. 10: 253. 1941.
Polypodium oligocarpum Willd., Sp. pi. ed. 4, 5: 201.
1810. TYPE: Venezuela, Caracas, Humboldt (ho-
lotype, B, Herb. Willd. 19699, in part; microfiche,
uc).
Dryopteris oligocarpa (Willd.) Kuntze, Rev. gen. pi.
3: 378. 1898.
Aspidium navarrense Christ, Bull. Herb. Boissier, II,
6: 160. 1906. TYPE: Costa Rica, Navarro, Werckle
(holotype, p).
Dryopteris columbiana C. Chr., Kongel. Danske Vi-
densk. Selsk. Skr., Naturvidensk. Afd., ser. 7, 4:
279. 1907. TYPE: Colombia, Santa Marta, H. H.
Smith 998 (holotype, P; isotypes, MICH!, uc!, us!).
Dryopteris lomatosora Copel., Univ. Calif. Publ. Bot.
19: 298. 1941. TYPE: Peru, Huanuco, Prov.
Huanuco, Dist. Churubamba, Hda. Mercedes,
Poca Perga, Mexia 8187 (holotype, uc!; isotypes,
F!, GH!, MO!, us!).
Thelypteris lomatosora (Copel.) Reed, Phytologia 17:
288. 1968.
Amauropelta oligocarpa (Willd.) Pic.-Ser., Webbia 3 1 :
251. 1977.
Stem ascending to erect, scales dark brown, dull
to somewhat shiny, lanceolate, setose on margins
and surface. Leaves several, clustered, 30-7 5(-l 10)
cm long. Lamina chartaceous, 1 -pinnate-pinnati-
fid, proximal 3-6 pairs of pinnae abruptly reduced,
the lowermost 1-3 mm long, not glanduliform,
sometimes hastate. Petiole 4-15 cm x 1-3 mm,
brownish and scaleless above the base. Rachis with
dense, spreading trichomes 0.1-1 mm abaxially,
sometimes with septate trichomes to 2 mm. Pin-
nae sessile, 4-1 1(-14) x l-2(-2.6) cm, deeply pin-
natifid to within 1 mm of costae, segments 2-3
mm wide. Aerophores lacking. Buds lacking. Veins
6-10 pairs per segment. Indument on costae, veins,
and often laminar tissue abaxially of dense,
spreading, acicular trichomes 0. 1-1 mm, long and
short trichomes often intermixed on costae, some-
times also with septate trichomes to 1.5 mm on
costae and hamate trichomes 0.2-0.5 mm on lam-
inar tissue, glands lacking, costal scales lacking,
lamina adaxially with scattered, stout, ascending,
acicular trichomes 0.5-1 mm on costules and veins,
and adpressed trichomes 0.1-0.2 mm on laminar
tissue. Sori supramedial to inframarginal, round,
indusia 0.2-0.3 mm wide with setae 0.2-0.3 mm,
receptacle and sporangia glabrous.
In montane forests and ravines, road banks, of-
ten in recently cleared areas or in secondary growth,
800-2450 m, Lambayeque, San Martin, Huanuco,
Junin, Ayacucho, and Cuzco.
Greater Antilles; Mexico to Panama; Colombia
to Bolivia; Venezuela; southern Brazil, north-
western Argentina.
The best characters for distinguishing the spe-
cies are the often inframarginal sori with short-
setose indusia, few pairs of abruptly reduced prox-
imal pinnae, spreading, lanceolate petiole base
scales, and the mixture of long, stout trichomes
on the adaxial costules and veins and more nu-
merous short, appressed trichomes on the laminar
tissue.
Thelypteris navarrensis (Christ) Proctor, treated
here as a synonym, differs primarily in having very
long, septate trichomes on the costae and rachis
abaxially. This variant seems to be common in
southern Central America and the Andes, al-
though perhaps absent elsewhere. Typical T. oligo-
carpa has a mixture of long and short, nonseptate
trichomes.
Lambayeque: 42 km from Olmos on road to Jaen,
Correll & Smith P818 (GH). San Martin: Mt. Guayra-
purina, near Tarapoto, Spruce 4015 (BM, w). Huanuco:
Along hwy between Huanuco and Tingo Maria, at Rio
Tulca, vicinity of Km 443.5, 6 km N of Acomayo, Croat
57852 (MO). Junin: La Merced-Chanchamayo, Soukup
1055 (F). Ayacucho: Prov. La Mar, between Tambo San
Miguel, Ayna and Hda. Luisiana, Dudley 1 1885 (GH).
Cuzco: Prov. Urubamba, Dist. Machu Picchu, just before
Machu Picchu Station, Saunders 1231 (F, GH).
6. Thelypteris scalaris (Christ) Alston, J. Wash.
Acad. Sci. 48: 234. 1958.
Aspidium scalare Christ, Bull. Herb. Boissier, II, 6:
159. 1906. LECTOTYPE (chosen by Christen-
TRYON & STOLZE: PTERIDOPHYTA OF PERU. HI.
15
sen, 1907, p. 323): Guatemala, Alta Verapaz,
Cubilquitz, von Tuerckheim (us!; isolectotype, us!).
Dryopteris scalaris (Christ) C. Chr., Kongel. Danske
Vidensk. Selsk. Skr., Naturvidensk. Afd., ser. 7,
4: 323. 1907.
Stem ascending to erect, scales brown, dull to
somewhat shiny, lanceolate, setose on margins and
surface. Leaves several, clustered, 50-1 80 cm long.
Lamina chartaceous, 1 -pinnate-pinnatifid, proxi-
mal ca. 7-1 1 pairs of pinnae gradually reduced,
the lowermost 5 mm long or less, sometimes glan-
duliform. Petiole to 25 cm x 3 mm, tan to stra-
mineous and lacking scales above the base. Rachis
with moderately dense, spreading trichomes most-
ly 0.2-0.4 mm abaxially. Pinnae sessile, 5-12 x
1.1-2.6 cm, deeply pinnatifid to within 1 mm of
costae, segments 2—4 mm wide. Aerophores lack-
ing or nearly so. Buds lacking or occasionally pre-
sent. Veins 8-16 pairs per segment. Indument on
costae, veins, and laminar tissue abaxially of sparse
to moderately dense, mostly hamate trichomes
mostly 0.2-0.3 mm, glands lacking, costal scales
lacking, lamina adaxially with numerous ad-
pressed trichomes 0. 1-0.2 mm on costules, veins,
and laminar tissue. Sori supramedial to inframar-
ginal, round, indusia with acicular and hamate
trichomes 0.1-0.3 mm, receptacle and sporangia
glabrous.
Edges of streams and on floating mat of lacus-
trine vegetation, 2000-2500 m, Cajamarca and
Amazonas.
Southern Mexico to Panama; Colombia to Peru;
Venezuela.
Cajamarca: Prov. Contumaza, La Pampa (Guzman-
go), Sagdstegui & Sagdstegui 14123 (uc). Amazonas: Prov.
Bongara, Pomacocha, D. Smith 6034 (MO).
7. Thelypteris andicola A. R. Sm., sp. nov.
A speciebus ceteris subg. Amauropeltae distinguenda
paleis paucis laxe ciliatis ad basin petiolorum, pinnis
plerumque 1-4 cm longis, basi aerophoris destitutis, seg-
mentis venis 2-6-jugis institutis, sons exindusiatis, et
praesertim trichomatibus numerosis erectis hamatis 0.3-
0.5 mm ad costas et venas et inter venis abaxialiter dis-
positis.
Stem ascending to erect, scales brown, dull, ovate
to ovate-lanceolate, ciliate. Leaves numerous,
densely clustered, 8-35(-55) cm long. Lamina
chartaceous, 1 -pinnate-pinnatifid, proximal 5-10
pairs of pinnae gradually reduced, the lowermost
1-4 mm long. Petiole 1-15 cm x 0.7-1. 5(-3) mm,
tan to stramineous above the base. Rachis nearly
glabrous abaxially. Pinnae sessile, 1-4 x 0.5-1.1
cm, incised ca. 0.4-0.7 mm from costa. Aero-
phores lacking. Buds lacking. Veins 2-6 pairs per
segment. Indument on costae, veins, and laminar
tissue abaxially of moderately dense, spreading,
mostly hamate trichomes 0.3-0.5 mm, glands
lacking, costal scales lacking, lamina adaxially with
scattered ascending, acicular trichomes 0.2-0.3 mm
on costules and veins, laminar tissue glabrous. Sori
medial, round, exindusiate, receptacle glabrous,
sporangia glabrous.
TYPE— Peru, Huanuco, Chavenillo, Woytkow-
ski 1029 (holotype, uc!).
On damp banks and moist rocks, 2400-3900 m,
La Libertad, Huanuco, and Cuzco.
Known only from Peru.
La Libertad: Prov. Santiago de Chuco, Dist. Quiru-
vilca, about halfway from Motil to Shorey, Sounders 888
(F, GH). Prov. and Dist. Otuzco, near Chaullacocha, on
Trujillo-Otuzco road, Saunders 901 (F, GH). Cuzco: Prov.
Paucartambo, Km 130 hacia Kosnipata, Parque Na-
tional del Manu, Nunez et al. 8528 (MO, uc).
8. Thelypteris enigmatica A. R. Sm., sp. nov.
A speciebus ceteris subg. Amauropeltae distinguenda
caule erecto, soris rotundatis trichomatibus caespitosis
0. 1-0.2 mm longis in receptaculo vel in indusio minuto,
sporangiis glabris, laminis adaxialiter trichomatibus 0. 1
mm longis numerosis appressis, et praesertim costis et
venis et spacio inter venas abaxialiter dense puberulis e
trichomatibus erectis acicularibus 0.1-0.15 mm longis.
Stem erect, scales dark brown, somewhat shiny,
lanceolate, setulose on margins and surface. Leaves
few, clustered, ca. 75 cm long. Lamina charta-
ceous, 1 -pinnate-pinnatifid, proximal ca. 8-1 0 pairs
of pinnae subabruptly reduced, the lowermost 2
mm long or less, sometimes glanduliform. Petiole
ca. 10 cm x 2-4 mm, brownish above the base,
with scattered, spreading scales. Rachis with dense,
acicular trichomes abaxially. Pinnae sessile, the
largest 8-10 x 1 .5 cm, deeply pinnatifid to within
1 mm of costae. Aerophores tuberculiform. Buds
lacking. Veins 8-10 pairs per segment. Indument
on costae, veins, and laminar tissue abaxially of
dense, spreading, acicular trichomes 0. 1-0. 1 5 mm,
glands lacking, costal scales lacking, lamina adax-
ially with numerous ascending to adpressed acic-
ular trichomes 0.1-0.15 mm on costules, veins,
and laminar tissue. Sori medial to supramedial,
round, exindusiate or indusia reduced to a few cells
16
FIELDIANA: BOTANY
and less than 0. 1 mm long, receptacle (or indusial
fragment) with a dense tuft of trichomes 0. 1-0.2
mm, sporangia glabrous.
TYPE— Peru, San Martin, Prov. Rioja, Pedro
Ruiz-Moyobamba road, Km 390, Venceremos,
D. Smith & Vdsquez 4594 (holotype, uc!; isotype,
MO!).
Montane rain forest, 2040 m, San Martin.
Known only from the type.
The relationships of this species are unclear,
hence the species epithet. In aspect and in the
adpressed trichomes on the lamina adaxially, it
resembles T. oligocarpa, but T. enigmatica lacks
uncinate trichomes. The pubescence on all parts
of the plant is uniformly short, 0. 1 5 mm or less,
much as in T. concinna. There is a tuft of tri-
chomes from the receptacle or from a very reduced
indusium.
9. Thelypteris ptarmiciformis (Rosenst.) Reed,
Phytologia 17: 307. 1968.
Dryopteris ptarmiciformis Rosenst., Repert. Spec. Nov.
Regni Veg. 12: 472. 1913. TYPE: Bolivia, Polo-
Polo, Buchtien 3435 (isotype, uc!).
Stem short-creeping to ascending, scales brown,
somewhat shiny or dull, lanceolate, glabrescent or
setose on margins and surface. Leaves few, clus-
tered, (25-)40-70 cm long. Lamina chartaceous,
1 -pinnate-pinnatifid, proximal 2-4 pairs of pinnae
abruptly reduced, the lowermost 1-5 mm long, not
glanduliform. Petiole 7-20 cm x 1.5-3 mm, tan
to stramineous and lacking scales above the base.
Rachis glabrescent or with numerous unicellular
or septate trichomes abaxially, often with short-
stipitate glands 0.05 mm. Pinnae sessile, 3.5-10
x 1.0-1.8 cm, deeply pinnatifid to within 1 mm
of costae, segments 2-3 mm wide, rather oblique
and subfalcate. Aerophores absent. Buds lacking.
Veins 5-12 pairs per segment. Indument on costae,
veins, and laminar tissue abaxially of numerous
short-stipitate glands 0.05-0. 1 mm, sometimes also
with unicellular (mostly ca. 0. 1 mm) and septate
(1-2 mm) trichomes, laminar tissue also with a
few hamate trichomes 0. 1 mm, costal scales lack-
ing, lamina adaxially glabrescent or with ascending
to adpressed trichomes 0.1-0.5 mm on costules,
veins, and laminar tissue, also with short-stipitate
glands. Sori supramedial to inframarginal, round,
indusia with numerous short-stipitate glands and
sometimes sparse trichomes, receptacle glabrous,
sporangia glabrous.
Montane rain forests, 500-2250 m, Cuzco and
Puno.
Peru; Bolivia; southern Brazil.
This is very closely related to T. oligocarpa and
may eventually prove to be only a variant of that
species. One of the specimens cited ( Vargas 16440)
bears septate trichomes to 2 mm mixed with short
unicellular trichomes 0. 1 mm on the abaxial rachis
and costae; the other specimen is very sparsely
pubescent and lacks septate trichomes. Both have
abundant short-stipitate laminar glands and a few
minute hamate trichomes on the lamina, thus
agreeing with the isotype seen.
Cuzco: Prov. Quispicanchi, vicinity of Inambari, Var-
gas 16440 (GH). Puno: Prov. Sandia, vicinity of Sandia,
Vargas 14810 (GH).
10. Thelypteris pusilla (Mett.) Ching, Bull. Fan
Mem. Inst. Biol., Bot. 10: 254. 1941.
Aspidium pusillum Mett., Ann. Sci. Nat. Bot., V, 2:
245. 1864. TYPE: Colombia, Fusagasuga, Lindig
92 (holotype, B).
Dryopteris pusilla (Mett.) Kuntze, Rev. gen. pi. 2: 8 1 3.
1891.
Stem ascending, scales brown, somewhat shiny,
lanceolate, densely setose on margins and surface.
Leaves numerous, arching, clustered, up to 20 cm
long. Lamina thin-herbaceous, 1 -pinnate, proxi-
mal 2-5 pairs of pinnae gradually reduced, the
lowermost 1-2.5 mm long. Petiole 1-7 cm x 0.4-
0.7 mm, stramineous above the base. Rachis with
hamate trichomes 0.2-0.3 mm abaxially. Pinnae
sessile, mostly 0.5-1.5 x 0.3-0.6 cm, entire or
faintly crenulate, sometimes slightly auriculate or
hastate at base. Aerophores lacking. Buds or small
plantlets arising from axils of distal pinnae. Veins
up to 7 pairs per pinna, only proximal pair 1 -forked.
Indument on costae, veins, and laminar tissue
abaxially of moderately dense, spreading, hamate
trichomes 0.15-0.3 mm, glands lacking, costal
scales lacking, lamina adaxially with numerous ad-
pressed acicular trichomes 0.2-0.3 mm. Sori me-
dial to supramedial, round, indusia round-reni-
form, 0.6-1 mm wide, faintly greenish and bearing
numerous hamate trichomes, receptacle glabrous,
sporangia glabrous.
On rock walls, 2560 m, rare in Peru: Cuzco.
Costa Rica; Colombia and Venezuela to Bolivia.
This species is one of the smallest in the genus
and usually has fronds arching and rooting from
buds produced near the distal part of the rachis.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
17
Cuzco: Urubamba, Machu Picchu, above Rio Mando,
4 km from Km 1 14 of Urubamba railroad, Peyton &
Peyton 1313 (MO).
1 1. Thelypteris brachypus (Sodiro) R. & A. Tryon,
Rhodora 84: 128. 1982.
Nephrodium brachypus Sodiro, Recens. crypt, vase.
Quit. 43. 1883. TYPE: Ecuador, Prov. Bolivar,
"bosques del Chimborazo cerca del pueblo de
Chillanes," Sodiro (possible isotype, uc!).
Dryopteris brachypus (Sodiro) C. Chr., Kongel. Danske
Vidensk. Selsk. Skr., Naturvidensk. Afd., ser. 7,
10: 135. 1913.
Stem ascending to erect (sometimes with trunk-
like caudex to 50 cm long), scales brown, shiny,
lanceolate, setose on margins and surface. Leaves
few, clustered, 15(-75) cm long. Lamina charta-
ceous, 1 -pinnate-pinnatifid, proximal 3(-12) pairs
of pinnae gradually reduced, the lowermost ca. 5-
10 mm long, never glanduliform. Petiole 5(-15)
cm x 1.5(-3) mm, brown to tan above the base,
with scattered, patent scales. Rachis glabrescent
(hamate-pubescent) abaxially. Pinnae sessile, 2-
3(-9) x 0.8(-2.0) cm, pinnatifid to ca. 1 mm from
costae. Aerophores peglike to 0.5 mm. Buds lack-
ing. Veins 4(-8) pairs per segment. Indument on
costae, veins, and laminar tissue abaxially of mod-
erately dense, spreading, hamate trichomes mostly
0.2-0.3 mm, lacking glands, sometimes with scat-
tered, brownish, deciduous, linear, entire scales
along costae, lamina adaxially with numerous ad-
pressed acicular trichomes 0.2-0.3 mm on cos-
tules, veins, and laminar tissue. Sori medial, round,
exindusiate, receptacle glabrous, sporangia gla-
brous.
Wet banks and mossy rocky areas, 2750-4000
m, Huanuco.
Panama; Colombia to Peru.
The two Peruvian specimens cited lack the long,
erect caudex found in specimens elsewhere and
are much smaller with only about three pairs of
reduced proximal pinnae (vs. 6-12 pairs else-
where). However, in the hamate trichomes and
other minute characters, the Peruvian specimens
seem very close to authentic material of T. brachy-
pus. The parenthetical measurements in the de-
scription are the upper limit in Ecuadorian spec-
imens.
Huanuco: Tambo de Vaca, Macbride 4364 (F). Mito,
Bryan 17 6 A (F).
12. Thelypteris leoniae A. R. Sm., sp. nov.
Inter species subg. Amauropeltae ob paleis rhachidis
T. funckii (Mett.) Alston proxima sed lamina usque ad
15-25 cm lata, pinnis proximalibus minus numerosis
(4-5 paribus) et abruptius reductis, aerophoris tuber-
culiformibus vel elongatis usque ad 1 mm, paleis rhachi-
dis setulosis margine, soris exindusiatis trichomatibus
caespitosis 0.3-0.5 mm recedens.
Stem not seen, presumably ascending or erect,
scales brown, shiny, narrowly lanceolate, densely
setose on margins and surface. Leaves probably
few and clustered, 40—85 cm long. Lamina sub-
coriaceous, 1 -pinnate-pinnatifid, proximal ca. 4-
5 pairs of pinnae subabruptly reduced, the low-
ermost 5-10 mm long, not glanduliform. Petiole
ca. 15 cm x 2-5 mm, brownish to tan above
the base, with dense, spreading, castaneous, linear-
lanceolate scales 3-8 mm long. Rachis with dense,
spreading scales 3-5 mm long and acicular tri-
chomes abaxially, and with reddish trichomes
adaxially. Pinnae sessile, 4-12 x 1-2 cm, pinnati-
fid 2-3 mm from costae. Aerophores tuberculi-
form or peglike to 1 mm. Buds lacking. Veins 6-
9 pairs per segment, lowermost pair running to or
just above the sinus. Indument on costae abaxially
of dense, spreading, acicular trichomes mostly 0.5-
1 mm, veins and laminar tissue with acicular and
hamate trichomes mostly 0.2-0.5 mm, glands
lacking, costal scales dense, spreading, like those
of rachis but shorter, 2-3 mm, lamina adaxially
with numerous spreading or ascending acicular tri-
chomes 0.5-1 mm on costules, veins, and laminar
tissue. Sori medial to supramedial, round to slight-
ly oblong, exindusiate, receptacle with a tuft of
acicular trichomes 0.3-0.5 mm, sporangia gla-
brous.
TYPE— Peru, San Martin, Prov. Mariscal Ca-
ceres, Rio Abiseo National Park, Chochos valley,
3500 m, Leon 1906 (holotype, F!; isotype, USM!).
Cloud forests, 2600-3500 m, San Martin and
Cuzco.
Peru and Bolivia.
This species is most closely related to T. funckii
(Mett.) Alston, which differs in the narrower lam-
ina, greater number of gradually reduced proximal
pinnae (nearly to rhizome), presence of large
indusia, rachis scales lacking marginal setae, and
absence of peglike aerophores at the base of the
pinnae abaxially. Thelypteris funckii occurs from
Costa Rica to Ecuador, Venezuela, and Guyana.
Another close relative is T. frigida (Christ) A. R.
Sm. & Lellinger, from Honduras to Costa Rica
and Venezuela. This species differs primarily in
18
FIELDIANA: BOTANY
being exindusiate and having fewer rachis and pet-
iole scales. The two Peruvian collections of T.
leoniae agree with one from Bolivia: Unduavi, Nov
1910, Buchtien 2647 (uc).
Cuzco: Prov. Paucartambo, Km 1 30 hacia Kosnipata,
Parque Nacional del Manu, Nunez et al. 8493, in part
(uc!).
13. Thelypteris furva (Maxon) R. Tryon, Rho-
dora 69: 6. 1967.
Dryopteris furva Maxon, J. Wash. Acad. Sci. 34: 24.
1944. TYPE: Peru [Huanuco], near Playapampa,
Macbride 4517 (holotype, F!; photo, GH; isotypes,
F!, us!).
Stem ascending, scales brown, dull to shiny,
ovate-lanceolate, setose on margins and surface.
Leaves few, clustered, 55-90 cm long. Lamina
chartaceous to subcoriaceous, 1 -pinnate-pinnatifid,
proximal 6-8 pairs of pinnae gradually reduced,
the lowermost 2 mm long, sometimes glanduli-
form. Petiole ca. 15-25 cm x 1.5-2 mm, atro-
purpureous or purple-brown and scaleless above
the base. Rachis with moderately dense trichomes
0.2 mm abaxially. Pinnae sessile, 3-8 x 1.0-2.3
cm, pinnatifid ca. 1-2 mm from costae, segments
2-3.5 mm wide. Aerophores absent or weakly de-
veloped. Buds lacking. Veins 4-10 pairs per seg-
ment. Indument on costae, veins, and laminar tis-
sue abaxially of sparse to moderately dense,
spreading, acicular trichomes 0. 1-0.2 mm, glands
absent, costal scales lacking, lamina adaxially with
dense, adpressed trichomes 0.2-0.3 mm on cos-
tules, veins, and laminar tissue. Sori medial, round,
indusia with dense trichomes 0. 1 mm, receptacle
and sporangia glabrous.
Wet cloud forests, 2300-2800 m, Huanuco and
Pasco.
Known only from the type and one other col-
lection.
Pasco: Prov. Oxapampa, San Alberto, Cordillera de
Yanachaga, van der Werffet al. 8420 (MO, uc).
14. Thelypteris pilosula (Mett.) R. Tryon, Rho-
dora 69: 7. 1967.
Aspidium pilosulum Mett., Fil. hort. bot. lips. 130.
1856. TYPE: Cultivated plant, said to have orig-
inated from Caracas, Venezuela, perhaps sent by
Gollmer (possible holotype, B!).
Dryopteris pilosula (Mett.) Hieron., Hedwigia 46: 332.
1907.
Dryopteris rimbachii Rosenst., Repert. Spec. Nov.
Regni Veg. 7: 147. 1909. TYPE: Ecuador, Mt.
Tunguragua, Rimbach 119 (isotypes, BM, uc!, us!).
Dryopteris macbridei C. Chr. & Maxon, J. Wash. Acad.
Sci. 34: 25. 1944. TYPE: Peru, (Huanuco), near
Yanano, Macbride 3828 (holotype, us!; isotype,
F!).
Thelypteris macbridei (C. Chr. & Maxon) R. Tryon,
Rhodora 69: 7. 1967.
Amauropelta pilosula (Mett.) Love & Love, Taxon 26:
325. 1977.
Stem short-creeping or ascending, scales brown,
somewhat shiny, lanceolate, setose and sometimes
stipitate-glandular on margins and surface. Leaves
several, approximate, (22-)50-70(-120) cm long.
Lamina chartaceous to subcoriaceous, 1 -pinnate-
pinnatifid, proximal 5-7 pairs of pinnae gradually
reduced, the lowermost ca. 1-2 mm long, some-
times glanduliform. Petiole 6-20 cm x 2-5 mm,
tan and scaleless above the base. Rachis with dense,
acicular trichomes 0.5-1.5 mm abaxially, some-
times with short-stipitate glands. Pinnae sessile,
2.5-8(-15) x 0.6-1. 5(-2.2) cm, deeply pinnatifid
to within 1 mm of costae, segments 1.5-3(-4) mm
wide. Aerophores absent. Buds lacking. Veins 3-
8 pairs per segment. Indument on costae, veins,
and sometimes laminar tissue abaxially of sparse
to dense, spreading, acicular trichomes mostly
(0.3-)0.5-1.5 mm, also with yellowish, short-stip-
itate glands, costal scales lacking, lamina adaxially
with numerous spreading or ascending trichomes
0.5-1 mm on costules, veins, and sometimes lam-
inar tissue. Sori medial to supramedial, round,
indusia glabrous or with marginal setae and often
short-stipitate glands, receptacle and sporangia
glabrous.
Wet montane forests, shaded ravines, slopes,
and ledges, 1800-3300 m, Cajamarca and Ama-
zonas, south to Apurimac and Cuzco.
Greater Antilles; southern Mexico to Panama;
Colombia to Bolivia; Venezuela.
Cajamarca: Trail from Las Huaringas to Huancabam-
ba, Davis & Turner 729 (GH). Amazonas: Cerros Calla
Calla, E side, 5 km above Leimebamba on road to Balsas
at San Miguel, Hutchison & Wright 4816 (uc, USM).
Huanuco: Prov. Huanuco, 32 km from Huanuco on road
to La Union, D. Smith et al. 2189 (F). Junin: Between
Palea and turnoff to San Ramon, Correll & Smith P782
(GH). Ayacucho: Pampalea, between Huanta and Rio
Apurimac, Killip & Smith 2327 1 (NY). Apurimac: Prov.
Abancay, Abancay, Vargas 16602 (GH). Cuzco: Quispi-
canchi, 1 6 km down from Marcapata, Fernandez & Jap-
anese Exp. C-35 (uc).
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
19
5. Thelypteris rufa (Poiret) A. R. Sm., Flora Ec-
uador 18: 77. 1983.
Polypodium rufum Poiret in Lam., Encycl. 5: 532.
1 804. TYPE: Peru, collector not known, possibly
Pavon? (holotype, P!).
Dryopteris rufa (Poiret) C. Chr., Index fil. 290. 1905.
Dryopteris subandina C. Chr. & Rosenst., Repert. Spec.
Nov. Regni Veg. 12: 472. 1913. TYPE: Bolivia,
Cantana ad fl. Ilimano, Buchtien 3120 (holotype,
s).
Dryopteris limaensis Copel., Univ. Calif. Publ. Bot.
19: 298. 1941. TYPE: Peru, Dist. Lima, road
Chosica to Matucana, Mexia 4079 (holotype, uc!;
isotypes, GH!, MO!, us!).
Thelypteris subandina (C. Chr. & Rosenst.) R. Tryon,
Rhodora69: 8. 1967.
Thelypteris limaensis (Copel.) Reed, Phytologia 17:
288. 1968.
Stem creeping to ascending, scales brown, dull
3 somewhat shiny, ovate-lanceolate, glabrous on
largins and surface. Leaves several, approximate,
0-135 cm long. Lamina chartaceous, 1-pinnate-
innatifid, proximal ca. 7-14 pairs of pinnae very
radually reduced, the lowermost 5 mm long or
:ss, occasionally glanduliform. Petiole 4-20 cm
< 2-5 mm, tan to stramineous and lacking scales
bove the base. Rachis glabrescent or with mod-
rately dense, spreading trichomes mostly 0.2-0.3
nm abaxially, sometimes with short-stipitate
lands less than 0.1 mm. Pinnae sessile, 3-12
-15) x 0.7-2.5 cm, deeply pinnatifid to ca. 1.5
im or less from costae, segments 2— 4(-5) mm
ride. Aerophores lacking. Buds lacking. Veins 4-
0(-13) pairs per segment. Indument on costae,
eins, and often lamina abaxially of sparse to dense,
preading, acicular trichomes mostly 0.2-0.3(-0.6)
nm, occasionally also with yellowish short-stipi-
ate glands 0. 1 mm or less, costal scales lacking,
amina adaxially glabrescent or with sparse to
lense, ascending trichomes 0.1-0.2 mm on cos-
ules, veins, and laminar tissue. Sori medial to
upramedial, round, indusia glabrous or usually
vith sparse to dense trichomes 0.1-0.3 mm and
ometimes minute glands, receptacle glabrous,
porangia glabrous or often with setae 0. 1-0.2 mm
rom capsule.
Weedy fern of irrigation ditches, small streams,
md disturbed sites, 1000-2800(-3200) m, Caja-
narca and La Libertad, south to Apurimac and
^uzco.
Ecuador to Bolivia.
This species is closely related to T. glandulo-
•olanosa, but differs in the simple (vs. often forked)
'eins that are often raised abaxially, shorter non-
ieptate trichomes usually less than 0.3 mm long
on the costae and indusia abaxially, absence of
deep red globose or pyriform glands along the cos-
tae and costules abaxially, commonly setose spo-
rangia, and the generally smaller indusia. More
than half of the numerous collections seen have
at least some setose sporangia; however, not all
sporangia are setose.
Cajamarca: Ca. 9 km W of San Juan and 46 km SW
of Cajamarca on road to San Pedro de Lloc, Dillon &
Whalen 4069 (F, uc). Amazonas: Prov. Bongara, Po-
macocha, D. Smith 6021 (MO). La Libertad: Prov. Otuz-
co, Sinsicap, Orga (Yerba Buena-Sinsicap), Lopez 2275
(GH). Lima: Prov. Lima, near Rio Santa Eulalia, Chosica,
40 km E of Lima, Tryon & Tryon 5341 (F, GH, NY, uc).
Ancash: Road Huaras to Casma, Tryon & Tryon 6566
(F, GH, VEN). Huanuco: Prov. Huanuco, 6 km S of Huanu-
co, Stork & Morton 9381 (F, uc). Junin: Tarma, Killip
& Smith 21908 (F, NY). Apurimac: Prov. Andahualles,
Huancarania, Velarde 4948 (GH). Cuzco: Quebrada de
Sappi, Cuzco, Tryon & Tryon 5356 (GH, USM).
16. Thelypteris glandulosolanosa (C. Chr.) R.
Tryon, Rhodora 69: 6. 1967. Figure 3.
Dryopteris glandulosolanosa C. Chr., Dansk. Bot. Ark.
9(3): 61.1937. LECTOTYPE (chosen here): Peru,
Cuzco, Prov. Quispicanchi, Dist. Huasao, Her-
rera 214 (us 1 198732!; isolectotype, us 1 198733!).
Stem short- to long-creeping, scales brown, dull
to somewhat shiny, appressed, ovate-lanceolate,
glabrous or sparingly ciliate on margins, surface
glabrous. Leaves several, approximate to distant,
20-120(-250) cm long. Lamina chartaceous,
1 -pinnate-pinnatifid, proximal (4-)7-12 pairs of
pinnae gradually reduced, the lowermost 2-10 mm
long, sometimes glanduliform. Petiole 4-30(— 45)
cm x 1.5-6 mm, stramineous and lacking scales
above the base. Rachis glabrescent or with mod-
erately dense, spreading trichomes mostly 0.3-1
mm abaxially. Pinnae sessile, 2.5-10(-13) x 0.7-
2.0(-2.5) cm, deeply pinnatifid to within 1 mm of
costae, segments 2-5 mm wide. Aerophores lack-
ing. Buds lacking. Veins 4—12 pairs per segment,
often forked, sunken, and sometimes darkened.
Indument on costae, veins, and often laminar tis-
sue abaxially of sparse to dense, spreading, acic-
ular or often silky, septate trichomes mostly 0.3-
1 .5 mm, usually with deep red to orangish, stalked
or sessile, pyriform glands along costae, costules,
and veins, costal scales lacking, lamina adaxially
with sparse to moderately dense, ascending or ad-
pressed acicular trichomes 0. 1-0.4 mm and some-
times deep red pyriform glands on costules and
veins, laminar tissue glabrescent or with adpressed
trichomes. Sori supramedial to inframarginal,
FIELDIANA: BOTANY
round, indusia with sparse to dense silky tri-
chomes 0.3-1 .0 mm, receptacle and sporangia gla-
brous.
Along streams and irrigation ditches, (2300-)
2800-4100 m, Lambayeque south to Puno.
Southern Ecuador to northwestern Argentina.
This species is highly variable in the density and
length of trichomes on the abaxial axes and in-
dusia. The type and many other collections, es-
pecially from Dept. Cuzco, have very long, silky,
septate trichomes; in other collections, the tri-
chomes may be sparser, shorter, and more cris-
pate. This latter variant is close to T. rufa, under
which the major differences are listed. Still other
variants have only a very small indusium bearing
a tuft of trichomes, e.g., Shepard 74a (GH), from
Puno, and Correll & Smith P858 (GH), from Ca-
jamarca. In general, T. glandulosolanosa occupies
a similar, but slightly more southerly range and
occurs at generally higher elevations than T. rufa.
But the two species probably grow together, and
it would not be surprising if they hybridize. An-
other closely related but still more southern species
in this group is T. argentina (Hieron.) Abbiatti.
Lambayeque: Prov. Lambayeque, Penachi, Llatas
Quiroz 1397 (F). Cajamarca: Ca. 25 km from Cajamarca
on road to Bambamarca, Correll & Smith P858 (GH),
P860 (GH). Ancash: Prov. Bolognesi, Cerro Capillapunta,
sur de Chiquian, Cerrate 146 (F, GH). Huanuco: Mito,
Macbride & Featherstone 1706 (F). Lima: Prov. Huaro-
chiri, Antisha, Muller & Muller 483 (GH). Junin: Prov.
Huancayo, ca. 9 km from Huancayo towards Chame-
seria, Saunders 648 (F, GH, K). Huancavelica: Yzcuchaca,
Kunkel 341c (B). Cuzco: Andahuaylilla, Coronado 150
(GH, uc). Arequipa: Chilena Valley, Stafford 5 59 (F). Puno:
Prov. Sandia, Cuyo-cuyo, Johns 83-143 (F).
mm from costae. Aerophores absent or tubercu-
liform to 0.2 mm. Buds lacking. Veins 3-5 pairs
per segment. Indument on costae and veins abax-
ially of moderately dense, spreading trichomes ca.
0.2 mm, laminar tissue glabrous, glands lacking,
costal scales lacking, lamina adaxially with as-
cending trichomes 0.2 mm on costules, veins, and
laminar tissue. Sori medial to supramedial, round,
exindusiate, receptacle glabrous, sporangia gla-
brous.
TYPE— Peru, Amazonas, Prov. Chachapoyas,
Cerros Calla Calla, east side, 18 km above Lei-
mebamba on road to Balsas, Hutchison & Wright
4870 (holotype, uc!; isotypes, F!, GH!, NY, USM!).
Cloud forest, 3100 m, Amazonas.
Known with certainty only from the type gath-
ering.
This species may be most closely related to T.
rufa, with which it agrees in pubescence and dis-
section, but it differs in the exindusiate sori, gla-
brous sporangia, and smaller fronds with fewer
reduced proximal pinna-pairs. Hutchison & Wright
4871 (F in part, not GH or uc), from the same
locality as the type, appears also to be this species.
1 8. Thelypteris canadasii (Sodiro) Alston, J. Wash.
Acad. Sci. 48: 234. 1958.
Nephrodium canadasii Sodiro, Recens. crypt, vase.
Quit. 48. 1883. TYPE: Ecuador, Pichincha, Pulu-
lahua, Hacienda Niebli, Sodiro (type material, p).
Nephrodium macradenium Sodiro, Recens. crypt, vase.
Quit. 47. 1883. TYPE: Ecuador, Sodiro (possible
type material, P!; possible isosyntype, uc!).
Dryopteris canadasii (Sodiro) C. Chr., Index fil. 256.
1905.
17. Thelypteris demissa A. R. Sm., sp. nov.
A. T. rufae (Poiret) A. R. Sm. et specierum affinium
distinguenda foliis minoribus 25-50 cm longis, pinnis
reductis proximalibus plus minusve 6, pinnis 2-3 x 0.8-
1 .0 cm, venis segmentorum 3-5-jugis, et praesertim soris
exindusiatis, sporangiis glabris.
Stem ascending, scales brown, somewhat shiny,
lanceolate, sparsely setose on margins and surface.
Leaves few, clustered, 25-50 cm long. Lamina
chartaceous, 1 -pinnate-pinnatifid, proximal ca. 3
pairs of pinnae subabruptly reduced, the lower-
most 1-3 mm long or less, not glanduliform. Pet-
iole 5-10(-25) cm x 1-2 mm, brownish to stra-
mineous and lacking scales above the base. Rachis
with sparse trichomes ca. 0.2 mm abaxially. Pin-
nae sessile, 2-3 x 0.8-1.0 cm, pinnatifid to ca. 1
Stem ascending to erect, scales tan to brown,
dull to somewhat shiny, ovate-lanceolate, glabrous
or sparsely setose on margins and surface. Leaves
few, clustered, 100-300 cm long, mucilaginous
when young. Lamina chartaceous, 1 -pinnate-pin-
natifid, proximal pairs of pinnae abruptly reduced,
the lowermost 1 mm long or less, glanduliform.
Petiole 15+ cm x 4-10 mm, tan to stramineous
and with scattered, appressed scales above the base.
Rachis with dense, fasciculate trichomes 0.1-0.2
mm abaxially. Pinnae sessile, 12-25 x i.6-3(-4)
cm, deeply pinnatifid to within 1 mm of costae.
Aerophores scalelike to 5 mm at pinna bases,
shorter ones also present at pinnule bases. Buds
lacking. Veins 10-30 pairs per segment. Indument
on costae and veins abaxially of moderately dense
to dense, fasciculate trichomes mostly 0.5-0.2 mm,
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
21
ften with shorter acicular trichomes 0. 1 mm on
minar tissue, glands lacking, costal scales tan,
jpressed, lamina adaxially with numerous as-
:nding to adpressed trichomes 0. 1 mm on cos-
iles, veins, and laminar tissue. Sori medial, round
• oblong, indusia with trichomes 0. 1 mm, recep-
,cle and sporangia glabrous.
Montane rain forests, 2000-2800 m, Amazonas,
uanuco, Pasco, and Cuzco. Ecuador to Bolivia.
This differs from T. thomsonii primarily in lack-
ig glands on laminar tissue and indusia. More
>mplete collections and field study are needed to
itablish whether this difference is taxonomically
gnificant.
Amazonas: Chachapoyas, 1 7-7 km down Cerro Calla-
alla toward Leimebamba, Edwin & Schunke 3691 (F).
uanuco: Chaglla, Macbride 3648 (F). Mito, Macbride
Featherstone 1618 (F, G). Pasco: Huancayo, Oxapam-
i, Soukup 2348 (F, GH). Cuzco: Urubamba, Machu Pic-
lu, Puncuyoj, 10 km SW of Incatambo, Peyton & Pey-
n 1377 (MO). Urubamba, Machu Picchu, 0.5 km N of
lion of Sayacmarca and Aobamba rivers, Peyton &
?yton 1498 (MO).
dument on costae and veins abaxially of moder-
ately dense to dense, fasciculate (appearing stellate
from base) trichomes 0.05-0.2 mm, with a few
acicular trichomes 0. 1 mm on laminar tissue, also
with numerous red to yellow, sessile glands on
lamina tissue, costal scales lacking or few, tan,
appressed, lamina adaxially with numerous as-
cending trichomes 0. 1 mm on costules, veins, and
laminar tissue. Sori inframedial to supramedial,
round to oblong, indusia with reddish to yellowish
sessile glands, sometimes sparsely setose, recep-
tacle and sporangia glabrous.
Montane rain forests and cloud forests, 680-
3000 m, Cuzco and Madre de Dios.
Jamaica; Hispaniola; southern Mexico to Pan-
ama; Colombia to Peru.
Cuzco: Prov. Paucartambo, entre Pillahuata y La Es-
peranza, Leon 2221, in part (uc, USM). Prov. La Con-
vention, Dist. Vilcabamba, trail Yupanqui to Rio Apu-
rimac, between Rumichurco and Alcobamba, Davis et
al. 1229 (GH). Madre de Dios: Prov. Manu, Carbon-
Salvacion, Vargas 16902 (GH).
9. Thelypteris thomsonii (Jenman) Proctor, Bull.
Inst. Jamaica, Sci. Ser. 5: 65. 1953.
Polypodium thomsonii Jenman, J. Bot. 24: 272. 1886.
TYPE: Jamaica, St. Andrew Parish, New Haven
Gap, Jenman J.P. 254 (holotype, u, according to
Proctor, 1985; isotypes, NY, us).
Dryopteris thomsonii (Jenman) C. Chr., Index fil. 298.
1905.
Dryopteris stuebelii Hieron., Hedwigia 46: 340, t. 6,
f. 13. 1907. SYNTYPES: Colombia, Sttibel 146,
439 (B).
Amauropelta thomsonii (Jenman) Pic.-Ser., Webbia
31: 251. 1977.
Stem ascending to erect, scales brown, some-
hat shiny, ovate-lanceolate, appressed, setose on
targins and surface. Leaves few, approximate,
30-1 50(-250) cm long, mucilaginous when young,
amina chartaceous, 1 -pinnate-pinnatifid, proxi-
lal ca. 6 pairs of pinnae abruptly reduced, the
twermost 1 mm long or less, glanduliform. Pet-
>le ca. 20-40(-100) cm x 5-6(-10) mm, tan to
:ramineous and scaleless above the base. Rachis
ith dense, fasciculate trichomes 0.05-0.2 mm
baxially. Pinnae sessile, 10-20(-30) x 1.8-3.0
-4.5) cm, deeply pinnatifid to within 1 mm of
astae, segments 3-5 mm wide. Aerophores scale-
ke to 6 mm at pinna bases, shorter ones to 1.5
im also usually present at pinnule bases. Buds
icking. Veins (12-) 16-2 5 pairs per segment. In-
20. Thelypteris phacelothrix (Rosenst.) R. Tryon,
Rhodora69: 7. 1967.
Dryopteris phacelothrix Rosenst., Repert. Spec. Nov.
Regni Veg. 11: 56. 1912. TYPE: Bolivia, Un-
duavi, N. Yungas, Buchtien 2709 (holotype, s?;
isotype, uc!).
Stem unknown, probably ascending, scales
brown, dull to somewhat shiny, ovate, appressed,
glabrescent or setose on margins and surface.
Leaves ca. 70 cm long. Lamina chartaceous,
1 -pinnate-pinnatifid, proximal 6-10 pairs of pin-
nae subabruptly reduced, the lowermost 1 mm
long or less, glanduliform. Petiole ca. 20 cm x 3-
5 mm, brownish to stramineous above the base,
with scattered, more or less appressed scales. Ra-
chis with dense, fasciculate trichomes 0.2-0.3 mm
abaxially. Pinnae sessile, 6-7 x 1 .0-1 .5 cm, deeply
pinnatifid to within 1 mm of costae, segments 2-
3 mm wide. Aerophores peglike, to 1 mm. Buds
lacking. Veins 6-8 pairs per segment. Indument on
costae and veins abaxially of dense, fasciculate
trichomes mostly 0.2-0.3 mm, with mostly non-
fasciculate trichomes 0. 1 mm on laminar tissue,
glands lacking, costal scales castaneous, ovate, to
2 mm, lamina adaxially with numerous ascending
to adpressed, mostly nonfasciculate trichomes 0. 1-
0.3 mm on costules, veins, and laminar tissue. Sori
medial, round, sometimes confluent at maturity,
indusia lacking or reduced to a nearly hidden few-
FIELDIANA: BOTANY
celled fragment, receptacle (or indusial fragment)
setose with trichomes 0.2 mm, sporangia glabrous.
Habitat and elevation not known, probably
montane rain forest, Junin.
Peru and Bolivia.
The sole Peruvian specimen seen has the costal
and rachis trichomes not so obviously fasciculate
as in the type. In the castaneous costal scales, small,
setose indusium, and general laminar dissection,
it is very similar.
Junin: [Prov. Huancayo], Cuesta of Huanacabra, Mat-
thews 937 (GH).
21. Thelypteris pavoniana (Klotzsch) R. Tryon,
Rhodora 69: 7. 1967.
Polypodium pavonianum Klotzsch, Linnaea 20: 386.
1847. TYPE: Peruviae Andium nemoribus, Ruiz
& Pavon [Herb. Ruiz no. 55} (holotype, B!; frag.,
us!).
Polypodium crossii Baker, Ann. Bot. 5: 455. 1891.
TYPE: Ecuador, Sierra de Roritroche, Andes of
Loja, Cross (holotype, K!).
Dryopteris pavoniana (Klotzsch) C. Chr., Index fil.
283. 1905.
Dryopteris pavoniana var. contracta Hieron., Hed-
wigia 46: 333. 1907. TYPE: Peru, near Ines be-
tween Pacasmayo and Moyobamba, Stiibel 1033
(holotype, B!).
Stem short- to long-creeping, scales brown, dull,
ovate, sparsely setose on surface. Leaves few, ap-
proximate to distant, 24-50 cm long. Lamina co-
riaceous, 1 -pinnate-pinnatifid, proximal 4-6 pairs
of pinnae subabruptly reduced, the lowermost 1
mm long or less, glanduliform. Petiole 10-25 cm
x 1-2.5 mm, atropurpureous and lacking scales
above the base. Rachis glabrescent or with sparse
trichomes 0.1 mm abaxially. Pinnae sessile, 1.5-
4.5 x 0.5-0.8 cm, pinnatifid to ca. 1 mm from
costae, segments 1.5-2 mm wide, margin strongly
inrolled. Aerophores peglike, 1-3 mm. Buds lack-
ing. Veins 3-6 pairs per segment, deeply im-
mersed. Indument on costae and veins abaxially
of moderately dense, spreading or ascending, acic-
ular trichomes mostly 0. 1-0.3 mm, glands absent,
costal scales lacking, lamina adaxially glabrous.
Sori medial, round, exindusiate, receptacle and
sporangia glabrous.
Wet slopes and banks, 2550-3300 m, Cajamar-
ca, Amazonas, and Huanuco.
Ecuador to Bolivia; Galapagos.
Cajamarca: South edge of Namora, Correll & Smith
P896 (GH). Amazonas: Prov. Chachapoyas, Wurdack 705
(F, GH, uc, USM). Prov. Chachapoyas, Pomacocha (Lei-
mebamba-Balsas), Lopez et al. 4393 (GH). Huanuco: Mi-
totambo, above Mito, Ferreyra 10 348 A (GH), 10364 (GH,
USM). Mito, Macbride & Featherstone 1622 (F, G, GH).
22. Thelypteris peruviana (Rosenst.) R. Tryon,
Rhodora 69: 7. 1967.
Dryopteris peruviana Rosenst., Repert. Spec. Nov.
Regni Veg. 7: 298. 1909. TYPE: Peru [San Mar-
tin], Cerro de Campana, Spruce 4655 (isotypes,
K!, w!; frag., us!).
Stem short-creeping to suberect, scales brown,
somewhat shiny, lanceolate, setose on margins and
surface. Leaves several, clustered, 25-100(-180)
cm long. Lamina chartaceous, 1 -pinnate-pinnati-
fid, proximal 3-6 pairs of pinnae abruptly reduced,
the lowermost 1 mm long or less, glanduliform.
Petiole 3-1 2 cm x 1 .5-4(-6) mm, brownish to tan
and sparsely scaly above the base. Rachis with
dense, spreading, acicular trichomes mostly 0.4-
1.0 mm abaxially. Pinnae sessile, 5-12(-15) x 1-
2(-3) cm, deeply pinnatifid to within l(-2) mm of
costae, segments 2-3(-6) mm wide. Aerophores
tuberculiform to absent. Buds lacking. Veins 8-
1 4(-l 8) pairs per segment. Indument on costae and
veins abaxially of moderately dense to dense,
spreading, acicular trichomes mostly 0.2-0.6 mm,
trichomes of laminar tissue 0.1-0.2 mm, glands
lacking, costal scales dark brown, subclathrate, se-
tose, lamina adaxially with sparse to numerous
ascending acicular trichomes 0. 1-0.2 mm on cos-
tules, veins, and laminar tissue. Sori medial to
supramedial, round, exindusiate, receptacle with
a few setae to 0.5 mm, sporangia with several setae
0. 1 mm on capsule.
Lowland and montane rain forests, on rocks,
380-1500 m, San Martin and Junin.
Ecuador to Bolivia.
Knapp and Mallet 8472 is smaller than other
cited specimens, with leaves less than 35 cm,
strongly oblique and falcate segments, inequilater-
al pinnae, and only 2 or 3 reduced proximal pinna
pairs. It agrees with other specimens in its scales,
trichomes, setose sporangia, and exindusiate sori.
San Martin: Prov. Lamas, Km 47.9 of Tarapoto- Yu-
rimaguas road, Knapp & Mallet 8472 (F, MO). Junin:
Chanchamayo Valley, Schunke 1498 (F). Yunguy, Woyt-
kowski 6604 (MO).
23. Thelypteris comptula A. R. Sm., sp. nov.
A speciebus ceteris subg. Amauropeltae distinguenda
rhachidi abaxialiter dense pubescent!, trichomatibus de-
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
23
flexis 0.1-0.2 mm longis, costis abaxialiter paleis atro-
brunneis vel nigrescentibus usque ad 1 mm, venis seg-
mentorum usque ad 20-jugis, obliquis costis comparatis,
valde falcatis, sori exindusiatis, sporangiis setulosis tri-
chomatibus 0. 1 mm longis.
Stem not known, probably ascending to erect,
scales not known. Leaves incomplete, probably
50-100 cm long. Lamina chartaceous, 1-pinnate-
pinnatifid, proximal pinnae not seen, probably
abruptly reduced with the lowermost 1 mm long
or less and glanduliform. Petiole incomplete, at
least 3 mm wide, tan above the base. Rachis with
dense, deflexed and appressed trichomes 0.1-0.2
mm abaxially. Pinnae sessile or short-stalked to 1
mm, to 1 7 x 3 cm, deeply pinnatifid to within 0.5
mm of costae, segments ca. 4 mm wide, oblique
and strongly falcate, basal pair of largest pinnae
greatly reduced to less than !/3 the length of the
next pair. Aerophores tuberculiform. Buds not seen.
Veins to 20 pairs per segment. Indument on costae
and costules abaxially of dense, slightly antrorse,
acicular trichomes mostly 0.1-0.3 mm, the veins
and laminar tissue glabrous or with sparse tri-
chomes 0. 1 mm, glands lacking, costal scales dark
brown or blackish, lanceolate, to 1 mm, setose,
laminar tissue adaxially with sparse adpressed tri-
chomes 0.1-0.2 mm. Sori medial to supramedial,
round, exindusiate, receptacle with a few setae 0.1-
0.2 mm, sporangia with minute setae 0.1 mm on
capsule.
TYPE— Peru, Cuzco/Madre de Dios, entre 1 5 Mil
y San Lorenzo, Vargas 11698 (holotype, GH!).
Montane forest, 700 m, Cuzco/Madre de Dios.
Peru and Bolivia.
This is most closely related to T. peruviana but
differs in the shorter, strongly deflexed and ap-
pressed trichomes 0.1-0.2 mm on the rachis,
shorter trichomes on the abaxial costae, more
deeply incised pinnae with strongly falcate seg-
ments, and the basal segments of largest pinnae
greatly reduced. The two species agree in the mi-
nutely setulose sporangia and dark costal scales;
both also occur at lower elevations than most other
species of sect. Lepidoneuron.
Paratype: Bolivia, [La Paz], San Josa, 1700 ft,
Williams 1246 (NY, 2 sheets).
24. Thelypteris atrorubens (Kuhn) A. R. Sm.,
comb. nov.
Aspidium atrorubens Kuhn, Linnaea 36: 112. 1869.
TYPE: Peru (Puno), St. Gavan (San Gaban),
Lechler 2267 (holotype, B!>.
Dryopteris atrorubens (Kuhn) C. Chr., Index fil. 253.
1905.
Stem not known. Leaves ca. 1 50 cm long. Lam-
ina thin-chartaceous, 1 -pinnate-pinnatifid, prox-
imal ca. 8 pairs of pinnae abruptly reduced and
glanduliform, 1 mm long or less. Petiole ca. 1 5 cm
x 2-3 mm, atropurpureous, shining, lacking tri-
chomes and scales. Rachis purplish brown, with
sparse spreading or subappressed trichomes 0.2-
0.3 mm. Pinnae sessile, up to 18 x 2.8 cm, deeply
pinnatifid to within 0.5 mm of costae, segments
3-5 mm wide, spreading. Aerophores lacking. Buds
lacking. Veins 8-11 pairs per segment. Indument
on costae and veins abaxially of scattered spreading
trichomes mostly 0.1-0.3 mm, laminar tissue gla-
brous, glands lacking, costal scales lacking, lamina
adaxially with scattered adpressed trichomes 0. 1-
0.2 mm on costules, veins, and laminar tissue. Sori
supramedial, round, indusia less than 0.3 mm,
setose with trichomes 0.1 mm, receptacle gla-
brous, sporangia glabrous.
Montane forests, 1800-2000 m, Cuzco and
Puno.
Known only from Peru.
The Vargas collection is tentatively assigned here.
It differs from the type in lacking trichomes on the
laminar tissue abaxially and in having slightly lon-
ger costal trichomes. Thelypteris atrorubens differs
from most other Peruvian Thelypteris by the atro-
purpureous and shining petiole and proximal part
of the rachis. Affinities are uncertain, but it may
be an aberrant member of sect. Uncinella; how-
ever, uncinate trichomes are lacking. The rachis
is weak and pinnae are rather lax and widely spaced
about 4 cm proximally, 2.5-3 cm distally, sug-
gesting that the leaves may lean on other vegeta-
tion. The species does not seem particularly close
to others with atropurpureous axes, e.g., T. ar-
borea (Brause) A. R. Sm. and T. pavoniana.
Cuzco: Prov. Paucartambo, Chacapampa, Vargas 9871
(uc, 2 sheets).
25. Thelypteris laevigata (Kuhn) R. Tryon, Rho-
dora 69: 6. 1967.
Phegopteris laevigata Kuhn, Linnaea 36: 112. 1869.
TYPE: Peru (Puno), Tatanara, Lechler 2628 (ho-
lotype, B!).
Dryopteris laevigata (Kuhn) C. Chr., Index fil. 273.
1905.
Stem unknown. Leaves ca. 50-100 cm long.
Lamina thick-chartaceous to subcoriaceous,
1 -pinnate-pinnatifid, proximal ca. 4 pairs of pin-
nae abruptly reduced, the lowermost 2 mm long
24
FIELDIANA: BOTANY
or less, glanduliform. Petiole 10-20 cm x 2-3
mm, atropurpureous above the base, shining, scales
lacking. Rachis atropurpureous, glabrous abaxi-
ally. Pinnae sessile, opposite, up to 7 x 2 cm,
pinnatifid to within 2 mm of costae, appearing
articulate. Aerophores consisting of a darkened
swelling subtended by a crescent-shaped ridge.
Buds lacking. Veins 7-10 pairs per segment. In-
dument on costae abaxially of scattered ovate to
lanceolate, brownish, appressed scales, trichomes
and glands lacking, lamina adaxially with a few
scattered, appressed, acicular trichomes 0.2 mm
on veins and laminar tissue. Sori medial, oblong
to elongate, indusia lacking, receptacle and spo-
rangia glabrous.
Without data on habitat or elevation, Puno.
Ecuador and Peru, in the latter, known only
from the type.
26. Thelypteris euchlora (Sodiro) Reed, Phyto-
logia 17: 275. 1968.
Polypodium euchlorum Sodiro, Recens. crypt, vase.
Quit. 58. 1 883. TYPE: Ecuador, Hacienda Niebli,
Sodiro (probable isotype, K!).
Dryopteris euchlora (Sodiro) C. Chr., Index fil. 263.
1905.
Stem long-creeping to ascending, to 1 m long,
scales brown, dull, ovate-lanceolate, setose on
margins and surface, sometimes glabrescent.
Leaves few, distant, 100-250 cm long. Lamina
chartaceous, 1 -pinnate-pinnatifid, proximal ca. 1 0
or more pairs of pinnae greatly reduced, the low-
ermost 1 mm long, glanduliform. Petiole 30-85
cm x 3-8 mm, tan to stramineous above the base,
glabrous. Rachis glabrous abaxially, often with
reddish trichomes adaxially. Pinnae sessile, 1 2-25
x 2.2-3.5 cm, deeply pinnatifid to within 2 mm
of costae, proximal basiscopic segments reduced.
Aerophores tuberculiform or absent. Buds lacking.
Veins 1 2-20 pairs per segment. Indument on cos-
tae, veins, and lamina abaxially lacking trichomes
or of very sparse trichomes, costal scales brown,
linear to lanceolate, subclathrate, ascending or ad-
pressed, lamina adaxially glabrous or with reddish
trichomes along costae and occasionally veins. Sori
medial to supramedial, round to oblong, indusia
lacking, receptacle and sporangia glabrous.
Montane rain forests, 1 500-2500 m, Amazonas,
Huanuco, and Cuzco.
Colombia to Bolivia.
The collection cited from Amazonas is atypical
in the contracted fertile fronds.
Amazonas: Prov. Bagua, Cordillera Colan, SE of La
Peca, Barbour 4104 (MO). Huanuco: SW slope of Rio
LlullaPichis watershed, Cerros del Sira, Dudley 13405
(GH). Cushi, Macbride 4852 (F). Prov. Huanuco, Dist.
Churubamba, crest of Santo Toribio, Mexia 8153 (GH,
uc). Cuzco: Paucartambo, Pillawata, Yanamayo-Tam-
bomayo, Vargas 16701 (GH).
27. Thelypteris pteroidea (Klotzsch) R. Tryon,
Rhodora69: 8. 1967.
Polypodium pteroideum Klotzsch, Linnaea 20: 389.
1847. SYNTYPES: Venezuela, Galipan, Moritz
291 (B); Venezuela or Colombia, Karsten 40 (Coll.
II) (B).
Dryopteris pteroidea (Klotzsch) C. Chr., Index fil. 287.
1905.
Stem short-creeping to ascending or erect, scales
brown, dull to shining, ovate-lanceolate, glabres-
cent or setose on surface and sparsely ciliate on
margins. Leaves few, well separated, commonly
1.5-5 m long, sprawling or scandent and supported
by other vegetation. Lamina chartaceous, 2-pin-
nate proximally, deeply 1 -pinnate-pinnatifid dis-
tally, proximal ca. 7 pairs of pinnae abruptly re-
duced, the lowermost less than 1 mm long,
glanduliform. Petiole up to 1.3 m x 4-8 mm,
stramineous and lacking scales above the densely
scaly base. Rachis glabrous. Pinnae sessile or short-
stalked, articulate, proximal ones reflexed, 15-35
(-50) x 5-13 cm, proximal ones 1 -pinnate. Pin-
nules entire or crenulate, larger ones constricted
at the base, reflexed, proximal ones often reduced
to !/3 the length of the longest, sometimes subaur-
iculate acroscopically. Aerophores absent. Buds
lacking. Veins simple or commonly 1-2-forked,
often 20 or more pairs per segment. Indument on
costae and veins abaxially lacking or of sparse an-
trorse trichomes 0. 1-0.3 mm long, glands lacking,
costal scales ovate to lanceolate, appressed,
brownish, lamina adaxially glabrous. Sori supra-
medial to inframarginal, round or often oblong,
exindusiate, receptacle glabrous, sporangia gla-
brous.
Montane rain forest, middle elevation, Junin.
Hispaniola; Colombia and Venezuela to Peru.
Junin: Villa Amoretti, Kunkel 606 (GH).
28. Thelypteris rudis (Kunze) Proctor, Bull. Inst.
Jamaica, Sci. Ser. 5: 64. 1953.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
25
PolypodiumrudeKunze,Linnaea 13: 133. 1839. TYPE:
Mexico, Jalapa, Schiede (holotype, LZ destroyed).
Dryopteris rudis (Kunze) C. Chr., Index fil. 289. 1 905.
Dryopteris engelii Hieron., Hedwigia 46: 339, t. 6, f.
12. 1907. LECTOTYPE (chosen by Christensen,
1907: Venezuela, Merida, Engel 90 (B).
Dryopteris boqueronensis Hieron., Hedwigia 46: 329,
t. 4,f. 5. 1907. TYPE: Colombia, Boqueron de
Bogota, Stubel 453 (holotype, B!).
Lostrea rudis (Kunze) Copel., Gen. filic. 140. 1947.
Amauropelta rudis (Kunze) Pic.-Ser., Webbia 31:251.
1977.
Stem creeping to ascending, scales brown, dull
3 somewhat shiny, linear- to ovate-lanceolate,
ensely pubescent to glabrescent on margins and
iirface. Leaves several, approximate to distant,
5-200 cm long. Lamina chartaceous to subcori-
ceous, 1 -pinnate-pinnatifid, proximal ca. 5-12
airs of pinnae abruptly reduced, the lowermost
;veral pairs 5 mm long or less, often glanduli-
jrm. Petiole 4-20 cm x 2-6 mm, tan and usually
icking scales above the base, sometimes sparsely
:aly. Rachis with moderate to dense, spreading
D ascending, often reddish trichomes abaxially.
'innae sessile, (5-)10-20 x (i_)i.5_3(-4) cm,
eeply pinnatifid to within 1.5 mm of costa. Seg-
lents mostly 2-5 mm wide, proximal pairs on
irger pinnae often reduced. Aerophores tuber-
uliform to peglike. Buds lacking. Veins 9-20(-25)
airs per segment. Indument on costae, veins, and
ften lamina abaxially of moderately dense to
ense, acicular trichomes mostly 0.2-0.8 mm, those
f costae and costules ascending to adpressed, a
;w on lamina sometimes hamate, costal scales
inceolate, brown, sometimes clathrate, lamina
daxially glabrescent or with sparse to moderately
ense, ascending trichomes 0.1-0.2 mm on cos-
ales, veins, and laminar tissue. Sori medial to
upramedial, round to oblong, indusia absent, re-
eptacle glabrous or sparingly setose, sporangia
labrous.
Somewhat weedy fern of montane rain forests,
Inn forests, especially along road banks and trails,
reas of secondary growth, 1 700-3 1 00 ( 1 300-4000)
i, Cajamarca, Amazonas, Ancash, Huanuco, Pas-
o, Ucayali, and Cuzco.
Greater Antilles; Mexico to Panama; Colombia
o Bolivia; Venezuela; Guyana.
This is the central species of a difficult complex,
xtremely variable in pubescence, scales, and size
>f leaves. Features characterizing the species are
he presence of often subclathrate scales and as-
ending to adpressed trichomes on the costae
ibaxially, exindusiate sori, and abruptly reduced
pinnae proximally, with several pairs of glandlike
pinnae below the lowest developed ones. Usually
the receptacle is glabrous. Trichomes between veins
are either acicular or, especially near the sinus,
hamate.
Cajamarca: Prov. San Miguel, Niepos, Llatas Quiroz
1529 (F). Amazonas: Prov. Chachapoyas, Cerros Calla
Calla, 1 8 km above Leimebamba on road to Balsas, Km
4 1 0, Hutchison & Wright 4871 (F, GH, uc). Ancash: Prov.
Huari, Huascaran National Park, Quebrada Pachachaca,
D. Smith et al. 12616 (uc). Huanuco: Muna, trail to
Tambo de Vaca, Macbride 4293 (GH). Pasco: Pichis Trail,
Dos de Mayo (as Junin), Killip & Smith 25882 (GH).
Ucayali: La Divisoria (as Loreto), Aquilar850 (GH). Cuz-
co: Prov. Paucartambo, San Pedro, Vargas 11343 (GH).
Department Unknown: Between Pacasmayo and Moyo-
bamba, Stubel 1049 (B).
29. Thelypteris supina (Sodiro) A. R. Sm., Fl. Ec-
uador 18: 82. 1983.
Nephrodium supinum Sodiro, Crypt, vase. Quit. 24 1 .
1893. SYNTYPES: Ecuador, "provincia de Qui-
to, Riobamba, Bolivar, etc.," Sodiro (possible
isosyntypes, NY!, P!, uc!).
Dryopteris supina (Sodiro) C. Chr., Index fil. 296. 1905.
Stem short-creeping to ascending, scales brown,
dull to somewhat shiny, lanceolate to ovate-lan-
ceolate, glabrous to sparsely pubescent on margins
and surfaces. Leaves several, approximate to sub-
distant, (60-)7 5-200 cm long. Lamina subcoria-
ceous, 1 -pinnate-pinnatifid, proximal (4-)8-10
pairs of pinnae subabruptly reduced, the lower-
most 2-4 pairs less than 5 mm long. Petiole mostly
1 0-20 cm x 2-6(-8) mm, tan to stramineous above
the base, often with scattered scales. Rachis with
moderately dense to dense, spreading or often de-
flexed, hyaline trichomes mostly 0.2-0.4(-0.6) mm
abaxially. Pinnae sessile, mostly 10-25 x (1_)1.5-
3.5(-5.5) cm. Segments mostly 2-5 mm wide, usu-
ally separated by broad sinuses, proximal ones of
larger pinnae reduced and often overlapping ra-
chis. Aerophores tuberculiform. Buds lacking.
Veins 8-15(-30) pairs per segment. Indument on
costae, veins, and laminar tissue abaxially of usu-
ally dense, spreading or ascending trichomes most-
ly 0.2-0.4 mm, costal scales lanceolate, brown,
sometimes clathrate, lamina adaxially glabrescent
or with sparse to moderately dense ascending tri-
chomes 0.1-0.3 mm on costules, veins, and lam-
inar tissue. Sori medial to supramedial, round to
oblong, confluent with age, exindusiate, receptacle
glabrous or sparingly setose, sporangia glabrous.
F1ELDIANA: BOTANY
Montane forest, ca. 2200 m, Huanuco.
Colombia to Peru.
The description is drawn from the numerous
Ecuadoran collections and the single, unusually
large Peruvian one.
Huanuco: Prov. Leoncio Prado, Km 463 on Lima-
Tingo Maria road, Young & Sullivan 888 (F).
30. Thelypteris pilosohispida (Hooker) Alston, J.
Wash. Acad. Sci. 48: 233. 1958.
Nephrodium pilosohispidum Hooker, Sp. fil. 4: 105.
1862. TYPE: Ecuador, Bolivar, Volcan Chim-
borazo, Spruce (holotype, K.!, 2 sheets; isotype, B).
Nephrodium retrorsum Sodiro, Recens. crypt, vase.
Quit. 51.1883. TYPE: Ecuador, Pichincha, Cerro
Corazon, Sodiro (possible type material at K!, So-
diro 44/20).
Dryopteris retrorsa (Sodiro) C. Chr., Index fil. 288.
1905.
Dryopteris pilosohispida (Hooker) C. Chr., Kongel.
Danske Vidensk. Selsk. Skr., Naturvidensk. Afd.,
ser. 7, 10: 148. 1913.
Dryopteris dumetorum Maxon, J. Wash. Acad. Sci.
34: 26. 1944. TYPE: Peru, [Huanuco], near Mito,
Macbride & Feather stone 1667 (holotype, F!; pho-
to, MO; isotype, us!).
Thelypteris dumetorum (Maxon) R. Tryon, Rhodora
69: 5. 1967.
Thelypteris retrorsa (Sodiro) A. R. Sm., Fl. Ecuador
18: 71. 1983.
Stem creeping, scales brown, somewhat shiny,
lanceolate, sparsely to densely setose on margins
and surfaces. Leaves few, subdistant, 80-120
(-200+) cm long. Lamina subcoriaceous, 1-pin-
nate-pinnatifid to barely 2-pinnate in large forms,
proximal 5-15 pairs of pinnae subabruptly re-
duced, the lowermost several (up to 1 2) pairs less
than 5 mm long, often glanduliform. Petiole 5-20
cm x 4-7 (-10) mm, tan and usually densely scaly
at base, sparsely scaly distally. Rachis with mod-
erately dense to usually dense, spreading tri-
chomes mostly 0.5-1.5 mm, often reddish abax-
ially. Pinnae sessile, 10-20 x 1 .7-3(_5) cm, deeply
pinnatifid to within 1 mm of costae, or incised to
the costae on larger proximal pinnae. Segments
mostly 3-6 mm wide, proximal pairs on larger
pinnae usually reduced and often strongly reflexed.
Aerophores tuberculiform. Buds lacking. Veins 1 2-
20(-25) pairs per segment. Indument of costae and
veins abaxially of moderately dense to dense,
spreading to slightly ascending, acicular trichomes
mostly 0.5-1 mm, laminar tissue glabrous or
sparsely pubescent, costal scales lanceolate, brown,
shiny, not or weakly clathrate, often setose on mar-
gin, lamina adaxially glabrescent or with sparse to
moderately dense, ascending trichomes mostly 0.4-
1 mm on costules and veins, laminar tissue gla-
brous. Sori medial to supramedial, round to ob-
long, partially hidden by revolute segment margin,
exindusiate, receptacle glabrous or sometimes se-
tose, sporangia glabrous.
Cloud forests, along roadsides and banks, 2400-
3100 m, Amazonas and Huanuco.
Hispaniola; southern Mexico to Costa Rica; Co-
lombia to Venezuela and Bolivia.
Thelypteris retrorsa differs primarily in the more
reflexed pinna segments, more deeply incised pin-
nae cut nearly or quite to the costa, and in the
slightly denser pubescence. These differences do
not seem of sufficient importance for recognition
of two taxa and may be partially the result of ex-
posure and size of plants.
Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 1 8
km above Leimebamba on road to Balsas, Hutchison &
Wright 487 IB (uc). Cerro Puma Urco, Soukup 4084
(MO). Prov. Chachapoyas, 17-7 km down Cerro Calla-
Calla toward Leimebamba, Edwin & Schunke 3688 (F,
USM). Huanuco: Muna, trail to Tambo de Vaca, Macbride
4334 (F).
31. Thelypteris corazonensis (Baker) A. R. Sm.,
Fl. Ecuador 18: 38. 1983.
Nephrodium corazonense Baker, J. Bot. 15: 163. 1877.
(as "carazanense"). TYPE: Ecuador, Pichincha,
Cerro Corazon, Jul 1873, Sodiro 44/7 (holotype,
K!).
Dryopteris corazonensis (Baker) C. Chr., Index fil. 258.
1905.
Stem creeping to ascending or suberect, scales
brown, dull to somewhat shiny, lanceolate, setose
on margins and surfaces or glabrescent. Leaves
several, approximate, (60-) 100-250 cm long.
Lamina subcoriaceous, 1-pinnate-pinnatifid,
proximal ca. 5-10 pairs of pinnae abruptly re-
duced, the lowermost several pairs less than 5 mm
long. Petiole to 120 cm x 4-8 mm, stramineous
and lacking scales above the base, or with sparse
scales distally. Rachis with moderately dense to
dense, spreading, hyaline to reddish trichomes 0.5-
2 mm long abaxially. Pinnae sessile, (7-) 15-25 x
(2-)3—4 cm, deeply pinnatifid to within 2 mm of
costae. Segments mostly 3-5 mm wide, proximal
pairs on larger pinnae slightly reduced and usually
reflexed, overlapping rachis. Aerophores tuber-
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
27
culiform or peglike. Buds lacking. Veins (9-)15-
25 pairs per segment. Indument on costae and veins
abaxially of moderately dense, spreading, acicular
trichomes 0.5-1.5 mm, laminar tissue glabrous,
costal scales lanceolate, brown to blackish, setose,
strongly clathrate, lamina adaxially with sparse to
moderate, ascending trichomes 0.5-1.5 mm on
costules and veins, laminar tissue glabrous. Sori
medial to supramedial, round to oblong, exindu-
siate, receptacle glabrous or occasionally sparingly
setose, sporangia glabrous.
Forming trunks to ca. 30 cm, montane rain for-
ests, 2285-2535 m, Cuzco.
Ecuador and Peru.
This differs from T. brausei primarily in the
presence of stiff trichomes on the costules and veins
adaxially, costal scales more decidedly clathrate
(surface walls hyaline), and somewhat longer but
fewer costal trichomes abaxially. These differences
may be insufficient to distinguish the two as spe-
cies.
Cuzco: Quillabamba, Santa Teresa, between Lambras
Pata and Mandornilloc, 0.5 km SW of La Playa, Peyton
& Peyton 1216 (MO). Urubamba, Machu Picchu, 0.5 km
N of union of Sayacmarca and Aobamba rivers, Peyton
& Peyton 1464 (MO).
32. Thelypteris brausei (Hieron.) Alston, J. Wash.
Acad. Sci. 48: 233. 1958.
Dryopteris brausei Hieron., Hedwigia 46: 337, /. 6, f.
11. 1907. LECTOTYPE (chosen by Christensen,
1907, p. 327): Colombia, Rio Paez, itinere ab
Popayan ad montem Huila, Stubel 145 (B!).
Stem short-creeping to ascending or suberect,
scales brown, dull to shiny, lanceolate, sparsely
setose on margins and surfaces or glabrescent.
Leaves several, approximate, (50-) 100-200 cm
long. Lamina chartaceous to subcoriaceous, 1 -pin-
nate-pinnatifid, proximal 5-10 pairs of pinnae
abruptly reduced, the lowermost several pairs less
than 5 mm long, glanduliform. Petiole to 60 cm
x 2-6 mm, stramineous to brownish and lacking
scales above the base. Rachis with moderately
dense to dense, spreading, hyaline to reddish tri-
chomes 0.5-2 mm long abaxially. Pinnae sessile,
6-16(-25) x i.5_3(_4) Cm, deeply pinnatifid to
within 1 mm of costa. Segments mostly 3-5 mm
wide, proximal ones of larger pinnae slightly re-
duced and reflexed, overlapping rachis. Aero-
phores tuberculiform. Buds lacking. Veins 7-20 +
pairs per segment, prominent and raised abaxially.
Indument on costae and costules abaxially of sparse
to moderately dense, usually ascending, acicular
trichomes 0.5-1 mm, veins and laminar tissue gla-
brous or nearly so, costal scales subclathrate, as-
cending, lamina adaxially glabrous on costules,
veins, and laminar tissue. Sori medial to supra-
medial, round to oblong, exindusiate, receptacle
glabrous, sporangia glabrous.
Cloud forests, 2300-3600 m, San Martin, Pasco,
and Cuzco.
Colombia to Bolivia.
This species is closely related to T. corazonensis:
see the discussion under that species. Other close
relatives are T. rudis and T. pilosohispida.
San Martin: Prov. Mariscal Caceres, NW corner of
Rio Abiseo Nat. Park, Chochos valley, Young 3575 (uc),
3661 (uc, USM), 3685 (uc, USM), 3781 (uc, USM). Pasco:
Prov. Oxapampa, San Alberto, Cordillera de Yanachaga,
von der Werff et al. 8435 (MO, uc). Cuzco: Prov. Pau-
cartambo, Tres Cruces, Vargas 12212 (F, GH).
33. Thelypteris caucaensis (Hieron.) Alston, J.
Wash. Acad. Sci. 48: 233. 1958.
Nephrodium caucaense Hieron., Bot. Jahrb. Syst. 34:
444. 1904. LECTOTYPE (chosen by Christen-
sen, 1 907): Colombia, Antioquia, Paramo de Ruiz,
Lehmann 3102 (B!; isolectotype, BM!).
Dryopteris caucaensis (Hieron.) C. Chr., Index fil. 257.
1905.
Dryopteris millet C. Chr., Kongel. Danske Vidensk.
Selsk. Skr., Naturvidensk. Afd., ser. 7, 10: 138.
1913. TYPE: Ecuador, Paluguillo, Mille 125 (ho-
lotype, P; isotype, BM!; photos, uc, us).
Thelypteris millet (C. Chr.) Reed, Phytologia 17: 293.
1968.
Stem short- to long-creeping, scales brown,
somewhat shiny, lanceolate, setose on margins and
surface. Leaves numerous to few, 10-60 cm long.
Lamina subcoriaceous to coriaceous, 1-pinnate-
pinnatifid, proximal l-3(-4) pairs of pinnae sub-
abruptly reduced, the lowermost 1-5 mm long,
sometimes glanduliform. Petiole up to 25 cm x
1-2.5 mm, tan to stramineous above the base.
Rachis glabrescent to moderately pubescent abax-
ially. Pinnae sessile, mostly 2-5 x 0.5-1.5 cm,
deeply pinnatifid to within 1 mm of costae, seg-
ments 2-3 mm wide. Aerophores absent or tuber-
culiform. Buds lacking. Veins 3-6 pairs per seg-
ment. Indument on costae and veins abaxially of
sparse to moderately dense, ascending, acicular
trichomes mostly 0.3-0.5 mm, laminar tissue gla-
28
FIELDIANA: BOTANY
brous or sparsely pubescent, glands lacking, costal
scales brown, clathrate or subclathrate, ovate-lan-
ceolate, lamina adaxially glabrous. Sori medial to
supramedial, round, exindusiate, receptacle gla-
brous or sometimes with a tuft of trichomes, spo-
rangia glabrous.
Near or above timberline in grassland, (2600-)
3300-4100 m, La Libertad, San Martin, Ancash,
Huanuco, and Cuzco.
Guatemala?; Costa Rica; Colombia to Bolivia;
Venezuela.
La Libertad: Prov. Sanchez Carrion, serial Huayllides,
Laguna Negra, D. Smith 2282 (F). San Martin: Prov.
Mariscal Caceras, Puerta del Monte, Young 1751 (uc).
Ancash: W side of mts at Km 3 1 1 , a few km below
Conococha, Correll & Smith P975 (GH). Huanuco: Cha-
vinillo, Coronado 176, in part (uc). Cuzco: Prov. Pau-
cartambo, Km 1 30 hacia Kosnipata, Nunez et al. 8493,
in part (uc).
34. Thelypteris hutchisonii A. R. Sm., sp. nov.
A speciebus ceteris subg. Amauropeltae distinguenda
caule late repente, paleis atrocastaneis nitidis non clath-
ratis ad costas abaxialiter, costis venis et spatiis inter
venosis abaxialiter dense pubescentibus, e trichomatibus
patentibus acicularibus 0.3-1 mm longis, saepe tricho-
matibus hamatis ad intervenia, lamina atroviridi, in-
dusiis dense setosis.
Stem long-creeping to 25 cm or more, scales
dark purple-brown, shiny, lanceolate, setose on
margins and surface. Leaves few, distant to ap-
proximate, 75-100 cm long. Lamina chartaceous,
1 -pinnate-pinnatifid, proximal 4-7 pairs of pinnae
subabruptly reduced, the lowermost 2 mm long or
less. Petiole ca. 25 cm x 2-4 mm, brown to tan
above the base, lacking scales above the base. Ra-
chis with dense trichomes 0.1-1 mm abaxially.
Pinnae sessile, largest 8-10 x 1.3-1.8 cm, deeply
pinnatifid to ca. 1 mm from costae, segments 2.5-
4 mm wide. Aerophores lacking. Buds lacking.
Veins 7-10 pairs per segment. Indument on costae,
veins, and laminar tissue abaxially of dense,
spreading, acicular trichomes mostly 0.3-1 mm,
some laminar trichomes hamate, glands lacking,
costal scales castaneous, not clathrate, lanceolate,
shiny, lamina adaxially with numerous ascending
trichomes 0.3-1.0 mm on costules, veins, and
laminar tissue. Sori supramedial to inframarginal,
round, indusia densely setose, receptacle glabrous,
sporangia glabrous.
TYPE— Peru, Amazonas, Prov. Chachapoyas,
Cerros Calla Calla, W side, 45 km above Balsas
midway on road to Leimebamba, Hutchison &
Wright 5828 (holotype, uc!; isotypes, F!, GH!, NY!,
USM!).
Rocky terrain, 3 100-3400 m, known only from
Prov. Amazonas in Peru.
In aspect, this is most similar to T. pilosula but
bears dark-castaneous scales on the costae abax-
ially. It has characteristics of both sect. Uncinella
(hamate trichomes, indusia) and sect. Lepidoneu-
ron (costal scales, creeping rhizome), and I am
uncertain of its sectional placement.
Amazonas: Prov. Chachapoyas, Calla-Calla, Aguado
[Herb. Truxillensis 6788] (F).
35. Thelypteris concinna (Willd.) Ching, Bull. Fan
Mem. Inst. Biol., Bot. 10: 251. 1941.
Polypodium concinnum Willd., Sp. pi. ed. 4, 5: 201.
1810. TYPE: Venezuela, Caracas, Bredemeyer
(holotype, B, Herb. Willd. 19698).
Dryopteris concinna (Willd.) Kuntze, Rev. gen. pi. 2:
812. 1891.
Amauropelta concinna (Willd.) Pic.-Ser., Webbia 31:
251. 1977.
Stem ascending to erect, scales brown, dull, ap-
pressed, ovate-lanceolate, glabrous on margins and
surface. Leaves few to many, clustered, 45-1 10 cm
long. Lamina chartaceous, 1 -pinnate-pinnatifid,
proximal 7-12 pairs of pinnae gradually reduced,
the lowermost 1 mm long or less, sometimes glan-
duliform. Petiole 5-15 cm x 1.5-4 mm, purplish
brown to brownish and scaleless above the base.
Rachis commonly with dense trichomes 0.05-0. 1
mm abaxially. Pinnae sessile, 4-10 x 0.8-2.0 cm,
deeply pinnatifid to within 1 mm of costae, seg-
ments 2-3 mm wide. Aerophores lacking. Buds
lacking. Veins 6-10 pairs per segment. Indument
on costae and veins abaxially of dense, spreading
trichomes 0.5-2.0 mm, laminar tissue glabrous,
glands lacking, costal scales lacking, lamina adax-
ially with numerous adpressed trichomes 0.05-0. 1
mm on costules, veins, and laminar tissue, occa-
sionally glabrescent. Sori supramedial, round, ex-
indusiate, receptacle glabrous or setulose, sporan-
gia with trichomes 0.05-1.0 mm on capsule.
Along streambanks, roadbanks, and edges of
montane forests, 500-2 1 50 m, common in Peru:
Lambayeque, Cajamarca, San Martin, Huanuco,
Pasco, Junin, Ucayali, Ayacucho, and Cuzco.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
29
Antilles; Mexico to Panama; Colombia to
northwestern Argentina; Venezuela.
Lambayeque: Prov. Lambayeque, road to Kerguer,
Penachi, Quiroz 1474 (F). Cajamarca: Prov. Cutervo,
Cutervo-Socota, Lopez & Sagdstegui 5332 (GH). San
Martin: Mariscal Caceres, 60 km NE of Jingo Maria,
Tryon & Tryon 5266 (F, GH, USM). Huanuco: Prov. Hua-
nuco, Dist. Churubamba, Hda. Exito, slope of Rio Ysa-
bel, Mexia 8184 (F, GH, uc). Pasco: Prov. Oxapampa,
Canyon de Huancabamba, Leon 675 (F, USM). Junin:
Carpapata, Kunkel611 (GH). Ucayali: Road to Aguaytia,
Km 209, Ridoutt (GH, USM). Ayacucho: Ayna, between
Huanta and Rio Apurimac, Killip & Smith 22807 (GH).
Cuzco: La Convention, Potrero, 8 km W of Quillabam-
ba, Tryon & Tryon 5386 (GH).
Known only from the type.
This species was recognized as distinct by Max-
on but has remained unpublished until now; I use
Maxon's epithet from the original labels. Thelyp-
teris loretensis is most closely related to T. con-
cinna, from which it differs in the more deeply
incised pinnae, more oblique segments, smaller
fronds, presence of scattered, sessile, resinous
glands on the abaxial lamina, generally sparser
(but similar) trichomes on the costae abaxially,
and absence or scarcity of trichomes on the lamina
abaxially. Thelypteris concinna occurs at higher
elevations and has not yet been found in Loreto.
36. Thelypteris loretensis A. R. Sm., sp. nov.
Inter species subg. Amauropehae sect. Blepharithecae
ob sporangia minute setulosa et trichomata similia 0.05-
0.1 mm longa T. concinna proxima sed differt pinnis
profundius incisis, segmentis obliquioribus costis com-
paratis, frondibus minoribus, glandibus resinosis dis-
persis ad laminam abaxialiter, trichomatibus plerumque
minus numerosis ad laminam abaxialiter dispositis.
Stem ascending, scales brown, dull, appressed,
ovate-lanceolate, glabrous on margins and surface.
Leaves few to many, clustered, 35^40 cm long.
Lamina thin-chartaceous, deeply 1 -pinnate-pin-
natifid, proximal 3—4 pairs of pinnae subabruptly
reduced, the lowermost 2-8 mm long, not glan-
duliform. Petiole 5-10 cm x 1-2 mm, tan to stra-
mineous and lacking scales above the base. Rachis
with moderate spreading trichomes 0.05-0. 1 mm
abaxially. Pinnae sessile, 4-6 x 0.9- 1.2 cm, deeply
pinnatifid to within 0.5 mm of costae, or the basal
pair incised to costa and free, segments 1.5-2 mm
wide. Aerophores lacking. Buds lacking. Veins 6-
8 pairs per segment. Indument on costae and veins
abaxially of moderately dense to dense, spreading
trichomes 0.05-0. 1 mm, laminar tissue with nu-
merous reddish, sessile, resinous glands, costal
scales lacking, lamina adaxially glabrous or veins
with sparse adpressed trichomes 0.05-0. 1 mm. Sori
supramedial, round, exindusiate, receptacle gla-
brous, sporangia minutely setulose, trichomes
0.05-0.1 mm.
TYPE— Peru, Loreto, above Pongo de Manseri-
che, right bank of Rio Santiago, Mexia 6211 (ho-
lotype, uc!; isotypes, BH!, F!, GH!, MICH!, MO!, us!).
Among rocks in damp sand, abundant locally,
200 m, Loreto.
37. Thelypteris deflexa (Presl) R. Tryon, Rhodora
69: 5. 1967.
Nephrodium deflexum Presl, Reliq. haenk. 1: 36, t. 5,
f. 2. 1825. TYPE: Peru, near Huanuco, Haenke
(holotype, PR).
Dryopteris lindigii C. Chr., Index fil. 275. 1905. Based
on Nephrodium deflexum Presl, not Dryopteris
deflexa (Kaulf.) C. Chr.
Dryopteris assurgens Maxon, J. Wash. Acad. Sci. 34:
24. 1944. TYPE: Peru, [Huanuco], Playapampa,
Macbride 45 17 a (holotype, F!; photos, GH, MO;
isotypes, F!, us!).
Thelypteris lindigii (C. Chr.) Alston, J. Wash. Acad.
Sci. 48: 233. 1958.
Thelypteris assurgens (Maxon) R. Tryon, Rhodora 69:
5. 1967.
Amauropelta deflexa (Presl) Love & Love, Taxon 26:
325. 1977.
Stem suberect to erect, caudex up to 15+ cm
long, scales brown, dull to shining, appressed,
ovate, glabrous on margins and surface. Leaves
clustered, 30-100(-120) cm long. Lamina herba-
ceous to chartaceous, deeply 1 -pinnate-pinnatifid,
proximal 2— 4(-6) pairs of pinnae gradually to sub-
abruptly reduced, the lowermost 2-8 mm long,
often auriculiform. Petiole 6-12 cm x 1-3 mm,
stramineous and lacking scales above the base.
Rachis glabrous abaxially. Pinnae sessile, opposite
to subopposite, 3-10 x 0.7-2 cm, deeply pinnati-
fid to within 1.5 mm of costae, segments 2-3 mm
wide. Aerophores lacking. Buds lacking. Veins 4-
8 pairs per segment. Indument on costae, veins,
and laminar tissue abaxially lacking or of sparse
trichomes 0.2-0.4 mm on costae and costules, cos-
tal scales lacking, lamina adaxially glabrous or
sparsely pubescent along veins. Sori inframedial
to medial, round, exindusiate, receptacle and spo-
rangia glabrous.
30
FIELDIANA: BOTANY
Montane forests, cloud forests, 1800-3200 m,
La Libertad, Huanuco, Pasco, and Junin.
Southern Mexico to Panama; Colombia to Peru;
Venezuela.
Dryopteris sellensis C. Chr., from Hispaniola,
may also be conspecific.
La Libertad: Ca. 3 km W of Huamachuco, Correll &
Smith P939 (GH). Prov. Huamachuco, Yanac, Sagdste-
gui 4533 (GH). Huanuco: Carpis Divide, Sandeman 5078
(BM). Carpish, Coronado 72 (GH, uc). Pasco: Prov. Oxa-
pampa, San Alberto, Cordillera de Yanachaga, van der
Werffet al. 8493 (uc). Junin: Prov. Chanchamayo, Rio
Rondayaco, 45 km from San Ramon, D. Smith et al.
2613 (F).
38. Thelypteris pachyrhachis (Mett.) Ching, Bull.
Fan Mem. Inst. Biol., Hot. 10: 253. 1941, var.
bogotensis (C. Chr.) Alston, J. Wash. Acad.
Sci. 48: 233. 1958.
Nephrodium crassipes Sodiro, Anales Univ. Centr. Ec-
uador 9(64): 323. 1893 [Crypt, vase. Quit. 234.
1893]. TYPE: Ecuador, along road Quito-Mana-
bi, Sodiro (possible isotype, us!).
Dryopteris pachyrhachis (Mett.) Kuntze var. bogo-
tensis C. Chr., Kongel. Danske Vidensk. Selsk.
Skr., Naturvidensk. Afd., ser. 7, 4: 306. 1907.
LECTOTYPE (chosen here): Colombia, Manga-
nos, Lindig 296 (B!).
Stem erect, scales brown, dull to somewhat shiny,
ovate-lanceolate, glabrous on margins and surface.
Leaves few, clustered, mostly 100-200 cm long.
Lamina chartaceous, 1 -pinnate-pinnatifid, proxi-
mal 5-10 pairs of pinnae gradually to subabruptly
reduced, the lowermost 2-10 mm long, often has-
tate. Petiole 8-20(-50) cm x 4-10 mm, strami-
neous to brownish above the base, sometimes mu-
cilaginous. Rachis glabrescent or with trichomes
abaxially. Pinnae sessile, mostly 10-20 x 1.5-3
cm, deeply pinnatifid 1-2 mm from costa. Aero-
phores scalelike or peglike, to ca. 1 mm. Buds
lacking. Veins 7-14 pairs per segment. Indument
on costae and veins abaxially lacking or of mod-
erately dense, spreading, flattened trichomes mostly
0.5-1 mm, also often with numerous yellowish to
orangish, sessile, resinous glands on laminar tis-
sue, costal scales tan to brownish, amorphous, ap-
pressed, lamina adaxially glabrous or with scat-
tered, acicular trichomes 0.5-0.8 mm on costules,
veins, and laminar tissue. Sori medial, round, in-
dusia ca. 1 mm in diameter, persistent, glabrous
or with sessile glands and trichomes to 0.5 mm,
receptacle and sporangia glabrous.
Montane forests, edge of streams, 2200—2750
m, Cajamarca, Amazonas, Pasco, and Cuzco.
Costa Rica to Peru.
Specimens cited from Amazonas and Pasco are
atypical in lacking glands but do not seem to be
the same as var. sprucei (Baker) A. R. Sm. (Co-
lombia and Ecuador), which also lacks glands. In
aspect, they seem closer to var. pachyrhachis (Cos-
ta Rica to Bolivia and southern Brazil, Greater
Antilles), which generally has laminar glands. This
species group is in particular need of revision.
Cajamarca: Prov. Cutervo, Grutas de San Andres,
Llatas Quiroz & Suarez C. 2738 (F). Amazonas: Prov.
Bagua, Cordillera Colan SE of La Peca, Barbour 4107
(MO, USM). Pasco: Prov. Oxapampa, 2-4 km N of Ma-
llampampa, D. Smith & Canne 5825, 5826 (uc). Cuzco:
Prov. Paucartambo, entre Pillahuata y La Esperanza,
Leon 2221 in part (uc, USM).
39. Thelypteris balbisii (Sprengel) Ching, Bull. Fan
Mem. Inst. Biol., Bot. 10: 250. 1941.
Polypodium balbisii Sprengel, Nova Acta Acad. Caes.
Leop. Carol. German. Nat. Cur. 10: 228. 1821.
NEOTYPE (chosen by Proctor, Fl. Less. Antill.
2: 281. 1977): Dominica, Hodge & Hodge 1203
(GH).
Aspidium sprengelii Kaulf., Flora 6: 365. 1823, nom.
super/I., see Morton, Amer. Fern J. 53: 62. 1963.
TYPE: Same as for Polypodium balbisii Sprengel.
Dryopteris sprengelii (Kaulf.) Kuntze, Rev. gen. pi. 2:
813. 1891.
Dryopteris balbisii (Sprengel) Urban, Symb. antill. 4:
14. 1903.
Dryopteris mercurii Hieron., Hedwigia 46: 335, /. 5,
/ 9. 1907. LECTOTYPE (chosen here): Colom-
bia, Santa Marta, Stiibel 363 (B!).
Thelypteris sprengelii (Kaulf.) Proctor, Bull. Inst. Ja-
maica, Sci. Ser. 5: 65. 1953.
Thelypteris mercurii (Hieron.) Reed, Phytologia 17:
292. 1968.
Stem ascending to erect, scales brown, some-
what shiny, ovate-lanceolate, glabrous on margins
and surface. Leaves few, clustered, 50-100 cm long.
Lamina chartaceous, 1 -pinnate-pinnatifid, proxi-
mal ca. 1 0 pairs of pinnae gradually to subabruptly
reduced, the lowermost 5 mm long or less, some-
times glanduliform, often hastate. Petiole 3-5 cm
x 2-6 mm, tan to stramineous above the base,
with scattered, appressed scales. Rachis glabres-
cent or with flexuous, septate trichomes, some-
times with short-stipitate glands abaxially. Pinnae
sessile, 10-15 x 1.8-2.5 cm, deeply pinnatifid to
within 1 mm of costae. Aerophores tuberculiform
or peglike, to 0.5 mm. Buds lacking. Veins 10-16
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
31
pairs per segment. Indument on costae and veins
abaxially of moderately dense to dense, spreading
or flexuous, acicular and often septate trichomes
mostly 0.5-2.0 mm, sometimes with shorter uni-
cellular trichomes 0.2-0.3 mm on laminar tissue,
also with numerous reddish to yellowish, sessile,
resinous glands on laminar tissue, costal scales
lacking, lamina adaxially glabrescent or with nu-
merous ascending acicular trichomes 0. 1-0.2 mm
on costules, veins, and laminar tissue. Sori medial
to supramedial, round, indusia with reddish to
yellowish sessile glands and lacking trichomes, re-
ceptacle glabrous, sporangia glabrous.
Moist banks and along streams and trails, 250-
300 m, San Martin, Huanuco, and Cuzco.
Antilles; southern Mexico to Panama; Colom-
bia to Peru; Venezuela.
San Martin: Tulumayo, on Rio Tulumayo, 23 km
from Tingo Maria on hwy to Pucallpa, Allard 22269
(GH). Huanuco: Prov. Huanuco, Tingo Maria, valley of
Rio Huallaga, "7000" ft [probably erroneous, perhaps
1000 ft], Belshaw 3066 (F, GH, MICH, uc). Cuzco: Prov.
La Convention, ca. 4 km NE from Hda. Luisiana and
Apurimac River, Dudley 11461 (GH). Rio Tambopata,
near Puerto Maldonado, Nunez 6484 (MO).
40. Thelypteris opposite (Vahl) Ching, Bull. Fan
Mem. Inst. Biol., Hot. 10: 251. 1941.
Polypodium oppositum Vahl, Eclog. amer. 3:53.1 807.
TYPE: Monlserrat, Ryan (holotype, c?, not found;
isotype, BM).
Aspidium conterminum Willd., Sp. pi. ed. 4, 5: 249.
1810. TYPE: Martinique, collector not known
(holotype, B, Herb. Willd. 19698).
Polypodium rivulorum Raddi, PI. bras. 1: 23, t. 35.
1825. TYPE: Brazil.
Aspidium coarctatum Kunze, Bot. Zeit. (Berlin) 1845:
287. 1845. TYPE: Venezuela, Caracas, MoritzSO
(holotype, B; isotype, BM!).
Dryopteris contermina (Willd.) Kuntze, Rev. gen. pi.
2: 812. 1891.
Dryopteris coarctata (Kunze) C. Chr., Index fil. 258.
1905.
Dryopteris rivulorum (Raddi) Hieron., Hedwigia 46:
334. 1907.
Dryopteris leucothrixC. Chr., Smithsonian Misc. Col-
lect. 52: 377. 1907. TYPE: Bolivia, near Yungas,
Rusby 432 (holotype, us!).
Thelypteris coarctata (Kunze) R. Tryon, Rhodora 69:
5. 1967.
Thelypteris leucothrix (C. Chr.) R. Tryon, Rhodora
69: 6. 1967.
Thelypteris contermina (Willd.) Reed, Phytologia 17:
269. 1968.
Amauropelta opposita (Vahl) Pic.-Ser., Webbia 31:251.
1977.
Amauropelta rivulorum (Raddi) Pic.-Ser., Webbia 3 1 :
251. 1977.
Stem suberect to erect, short, scales brown, dull,
appressed, ovate-lanceolate, glabrous on margins
and surface. Leaves few to many, clustered, mostly
(1 5-)40-l 10 cm long. Lamina chartaceous to sub-
coriaceous, deeply 1 -pinnate-pinnatifid, proximal
10-20 pairs of pinnae very gradually reduced, the
lowermost 1-5 mm long, often hastate. Petiole 1-
10 cm x 1-4 mm, stramineous to brownish and
lacking scales above the base. Rachis with mod-
erately dense to dense, crispate trichomes 0.3-0.8
mm abaxially. Pinnae sessile, (1.5-)3-9 x 0.5-1.8
cm, pinnatifid to 1-2 mm from costae, segments
(l-)2^4 mm wide, margins strongly revolute.
Aerophores lacking or weakly developed. Buds
lacking or weakly developed. Veins mostly 3-7
pairs per segment. Indument on costae, veins, and
sometimes laminar tissue abaxially of moderately
dense to dense, acicular or crispate trichomes
mostly 0.2-0.6 mm, also with numerous yellowish
to orangish shiny sessile resinous glands on veins
and laminar tissue, costal scales lacking, lamina
adaxially glabrous or usually with scattered tri-
chomes 0.1-0.3 mm. Sori medial to supramedial,
round, indusia tan, 0.5-0.7 mm in diameter, glan-
dular on margin, sometimes also with a few short
trichomes, sporangia and receptacle glabrous.
Lowland and montane forests, especially along
roads, trails, streams, and ditches, 100-1100
(-1800) m, Amazonas, San Martin, Loreto, Hua-
nuco, Pasco, Junin, Ucayali, Ayacucho, Cuzco,
and Puno.
Puerto Rico; Lesser Antilles; Costa Rica and
Panama; Colombia to Bolivia; Venezuela; south-
ern Brazil.
This is by far the most common Thelypteris of
subg. Amauropelta in the lowlands of Peru. It may
grow in partial or full sun or shade, and exhibits
a variety of textures and sizes; it is also variable
in the density of the trichomes on the lamina and
axes, but the trichomes are often crispate or twist-
ed, especially along the rachis and costae abaxially.
Foster et al. 3005 is unusual in the small leaf size
(15x4 cm) and more pubescent adaxial lamina.
Amazonas: Prov. Bagua, Ribera derecha del Maranon,
Lopez et al. 4162 (GH). San Martin: Prov. Lamas, Dist.
Lamas, below English Evangelical Mission, Lamas, Bel-
shaw 3425 (F, GH, uc). Loreto: Pumayacu, between Bal-
sapuerto and Moyobamba, Klug 3245 (F, G, GH, NY).
Huanuco: Prov. Huanuco, Dist. Churubamba, Hacienda
Exito, Rio Ysabel, Mexia 8138 (F, GH, uc, USM). Pasco:
32
FIELDIANA: BOTANY
Prov. Oxapampa, Palcazu valley on Rio Palcazu between
Iscozacin and San Juan de Chuchurras, D. Smith 3926
(uc). Junin: La Merced, Killip & Smith 23680 (GH). Uca-
yali: Bosque von Humboldt Experimental Station, Km
86 on Pucallpa-Tingo Maria road, D. Smith 1224 (F, MO,
uc). Ayacucho: Prov. La Mar, Hacienda Luisiana, Dud-
ley 10006 (GH). Cuzco: Prov. La Convencion, near Camp
Zero, ca. 4 km NE from Hacienda Luisiana and Apu-
rimac River, Dudley 11462 (GH). Prov. Paucartambo,
Atalaya, R. Foster et al. 3005 (GH). Puno: Prov. Sandia,
Asalaya, Vargas 14828 (GH).
41. Thelypteris micula A. R. Sm., sp. nov.
Inter species subg. Amauropeltae sect. Amauropeltae
e glandibus resinosis ad laminam indusiumque, soris
indusiatis, pinnis proximalibus gradatim reductis T. op-
positam proxima sed differt frondibus minoribus, mem-
branaceis, 8-20 cm longis, pinnis usque ad 1.2 x 0.4
cm, segmentorum venis 1-2-jugis, lamina adaxialiter
dense pubescente trichomatibus 0.1-0.2 mm.
Stem erect, scales brown, somewhat shiny, lan-
ceolate, 1-2 mm long, glabrous on margins and
surface. Leaves few, clustered, ca. 8-20 cm long.
Lamina thin-chartaceous, 1-pinnate-pinnatifid,
proximal 5-6 pairs of pinnae very gradually re-
duced, the lowermost ca. 1 mm long, not glan-
duliform. Petiole 2-3 cm x 0.5 mm, tan and lack-
ing scales above the base. Rachis with trichomes
mostly 0.2-0.4 mm abaxially. Pinnae sessile, 0.8-
1.2 x 0.3-0.4 cm, shallowly pinnatifid less than
1 mm from margin, segments ca. 1.5 mm wide,
broader than long. Aerophores lacking. Buds lack-
ing. Veins 1-2 pairs per segment, or merely forked
or simple toward apex. Indument on costae, veins,
and laminar tissue abaxially of moderately dense
to dense, spreading, acicular trichomes mostly 0. 1-
0.3 mm, also with reddish sessile, resinous glands
on laminar tissue, costal scales lacking, lamina
adaxially with dense spreading or ascending tri-
chomes 0.1-0.2 mm on costules, veins, and lam-
inar tissue. Sori medial to supramedial, round,
indusia with reddish sessile glands and sparse tri-
chomes 0.1 mm, receptacle glabrous, sporangia
glabrous.
TYPE— Peru, Junin, Chanchamayo Valley, C.
Schunke 848 (holotype, F!).
Habitat unknown, 1500 m, known only from
the type.
This species is probably most closely related to
T. opposita, but differs in the small, very thin-
textured fronds and dense, short trichomes (0.1-
0.2 mm) on the adaxial surface of the lamina.
42. Thelypteris dudleyi A. R. Sm., sp. nov.
A speciebus ceteris subg. Amauropeltae distinguenda
caulibus paleis glabris ovato-lanceolatis, foliis parvis 25-
65 cm longis, glandibus ad laminam abaxialiter abun-
dantibus sessilibus vel brevistipitatis, paleis paucis
atrobrunneis usque ad 1 mm longis ad costam, indusiis
glandiferis 0.2-0.3 mm diametro.
Stem ascending to erect, scales brown, dull to
somewhat shiny, ovate-lanceolate, glabrous on
margins and surface. Leaves few to numerous,
clustered, 25-65 cm long. Lamina chartaceous,
1-pinnate-pinnatifid, proximal 5-7 pairs of pinnae
subabruptly reduced, the lowermost 1 mm long or
less, sometimes glanduliform. Petiole 4-6 cm x
1-2 mm, tan and lacking scales above the base.
Rachis glabrous or with trichomes 0.2-0.3 mm
abaxially, sometimes with short-stipitate glands
0.1 mm. Pinnae sessile, 2-8 x 0.6-1.5 cm, deeply
pinnatifid to within 1 mm of costae, segments 1—
2.5 mm wide. Aerophores tuberculiform. Buds
lacking. Veins 4-7 pairs per segment. Indument on
costae abaxially of sparse to moderately dense,
slightly ascending, acicular trichomes mostly 0. 1 5-
0.4 mm, veins and laminar tissue lacking tri-
chomes, but with numerous orangish to yellowish,
sessile, resinous or short-stipitate glands, costal
scales dark-castaneous, to 1 mm, lamina adaxially
with adpressed trichomes 0. 1-0.2 mm on costules,
veins, and laminar tissue, also with sessile glands.
Sori supramedial, round, indusia 0.2-0.3 mm wide,
glandular, lacking trichomes, receptacle and spo-
rangia glabrous.
TYPE— Peru, Cuzco, Prov. La Convencion, be-
tween camp 4 and 5, T. R. Dudley 10783 (holo-
type, GH!; isotypes, F!, MO!, USM!).
Common colonizer in full sun on open land-
slide, 2200-2825 m, Cuzco.
This species has the laminar aspect of T. deflexa
but differs from that species in the presence of a
small indusium, abundant sessile glands on the
abaxial lamina, and more numerous reduced prox-
imal pinna-pairs. It is also similar to T. arenosa
A. R. Sm. from Ecuador and Venezuela, but differs
in the weakly ascending or short-erect rhizome (vs.
trunklike and long-erect), smaller indusium, and
ovate, glabrous petiole base scales. The collector
described the petioles as "purplish-black and
glossy," but this appears to be true only at the
base. There are a few castaneous scales on the
costae abaxially, but it does not seem likely that
these indicate an affinity with sect. Lepidoneuron.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
33
The sessile laminar glands suggest that this may
be a member of sect. Amauropelta, but this is un-
certain. Pennell 13942 agrees with the type in the
glands but differs in being more sparsely pubes-
cent, with larger fronds and larger indusia; it lacks
scales on the abaxial costae.
Cuzco: Pillahuata, Cerro de Cusilluyoc, Pennell 13942
(GH, NY).
43. Thelypteris cheilanthoides (Kunze) Proctor,
Bull. Inst. Jamaica, Sci. Ser. 5: 58. 1953.
Aspidium cheilanthoides Kunze, Linnaea 22: 578.1 849.
TYPE: Brazil, Regnell (type material, s).
Lastrea cheilanthoides (Kunze) T. Moore, Index fil.
88. 1858.
Nephrodium resinosofoetidum Hooker, Sp. fil. 4: 105.
1862. LECTOTYPE (chosen by Christensen,
1913): Ecuador, Tunguragua, Spruce 5300 (K, 2
sheets!).
Dryopteris resinosofoetida (Hooker) Kuntze, Rev. gen.
pi. 2: 813. 1891.
Dryopteris cheilanthoides (Kunze) C. Chr., Index fil.
257. 1905.
Dryopteris cheilanthoides var. resinosofoetida (Hook-
er) C. Chr., Kongel. Danske Vidensk. Selsk. Skr.,
Naturvidensk. Afd., ser. 7, 4: 331. 1907.
Thelypteris resinosofoetida (Hooker) Ching, Bull. Fan.
Mem. Inst. Biol., Bot. 10: 254. 1941.
Dryopteris densa Maxon, J. Wash. Acad. Sci. 34: 25.
1944. TYPE: Peru, [Huanuco], Huacachi, near
Muna, Macbride 4175 (holotype, F, 2 sheets!; iso-
type, us!).
Thelypteris cheilanthoides var. resinosofoetida (Hook-
er) Proctor, Bull. Inst. Jamaica, Sci. Ser. 5: 58.
1953.
Thelypteris densa (Maxon) R. Tryon, Rhodora 69: 5.
1967.
Amauropelta cheilanthoides (Kunze) Love & Love,
Taxon 26: 325. 1977.
Stem suberect to erect, scales brown, dull to
shiny, subappressed, ovate-lanceolate, glabrous on
margins and surface. Leaves several, clustered,
mostly 65-250 cm long, usually mucilaginous when
coiled or young. Lamina subcoriaceous to coria-
ceous, deeply 1 -pinnate-pinnatifid, proximal 4-7
pairs of pinnae subabruptly reduced, the lower-
most 1-5 mm, glanduliform. Petiole 10-100 cm
x 4-10 mm, brownish to tan, densely scaly in
proximal third, lacking scales distally. Rachis gla-
brescent to pubescent with trichomes 0.3-2 mm
long abaxially. Pinnae sessile, 10-32 x 1.5-4 cm,
deeply pinnatifid to within l-2(-3) mm of costae,
segments ca. 3—4 mm wide, margin often strongly
revolute. Aerophores peglike to scalelike, to 2 mm
long or more, smaller ones sometimes present at
base of costules. Buds lacking. Veins 1 2-24(-30)
pairs per segment, usually immersed and appear-
ing dark in dried specimens. Indument on costae
and veins abaxially lacking or of moderately dense
to dense simple to pluricellular, acicular or crispate
trichomes 0.2-2 mm, costae also with a few stra-
mineous to tan, amorphous appressed scales, lam-
inar tissue abaxially glabrous, pubescent, or with
sessile yellowish to orangish glands, lamina adax-
ially glabrous or with sparse trichomes to 0.3 mm
on veins and laminar tissue. Sori supramedial to
submarginal, round, confluent at maturity, indusia
often large, subathyrioid, persistent, sometimes
glandular on margin, receptacle and sporangia gla-
brous.
Montane rain forests, along streams and trails,
(400-) 1000-3 500 m, Cajamarca, Amazonas, San
Martin, Loreto, Huanuco, Pasco, Junin, and Cuz-
co.
Jamaica; Hispaniola; Mexico to Panama; Co-
lombia to Bolivia; Venezuela; southern Brazil.
Cajamarca: Prov. San Miguel, La Toma, Niepos, Lla-
tas Quiroz 1583 (F). Amazonas: Carlo Santa Lucia E of
Chachapoyas, Wurdack 741 (F, GH, uc, USM). San Mar-
tin: Prov. Lamas, Dist. Lamas, 2-4 km N of San An-
tonio, along Rio Cumbasa, Belshaw 3545 (GH, uc). Lo-
reto: Pumayacu, between Balsapuerto and Moyobamba,
Klug3231 (F, GH). Huanuco: Carpish, between Huanuco
and Tingo Maria, Ferreyra 1836 (GH, USM). Pasco: Prov.
Oxapampa, Rio El Tunqui, D. Smith & Alban 5541 (F).
Junin: Chanchamayo Valley, Schunke 15 (F), 62 (F). Cuz-
co: Prov. La Convention, Rio Apurimac, mouth of Rio
Pampaconas, above Sinechinete, Davis et al. 1287 (GH).
44. Thelypteris furfuracea A. R. Sm., sp. nov.
Inter species subg. Amauropeltae sect. Blennocaulonis
indusiis usque ad 1 mm diametro, venis immersis, aero-
phoris elongatis T. cheilanthoidem (Kunze) Proctor ap-
proximans sed differt paleis bicoloribus ad rhachim et
costas abaxialiter occurentibus, paleis corpere castaneo
et marginibus fulvis albidisve, trichomatibus non sep-
tatis 0.2-0.4 mm longis abaxialiter ad costas occurren-
tibus.
Stem not seen, probably ascending to erect, scales
dark brown and shiny with narrow pale margins,
linear-lanceolate to 1 cm, glabrous on margins and
surface. Leaves ca. 50 cm long. Lamina subcori-
aceous, 1 -pinnate-pinnatifid, proximal ca. 7 pairs
of pinnae subabruptly reduced, the lowermost 1
mm long or less, glanduliform. Petiole 5-10 cm
x 3-4 mm, brownish and with a few scales above
the base. Rachis with curved or crispate trichomes
34
FIELDIANA: BOTANY
to 0.2-0.5 mm and scattered scales abaxially. Pin-
nae sessile, 4—7 x 1.0—1.4 cm, deeply pinnatifid
to within 1 mm of costae, segments 2-3 mm wide,
strongly inrolled at margin. Aerophores peglike, to
1 mm. Buds lacking. Veins 8-10 pairs per segment,
sunken and darkened abaxially. Indument on cos-
tae abaxially of moderately dense to dense, curved
or crispate trichomes mostly 0.2-0.4 mm, glands
lacking, costal scales numerous, castaneous with
pale margins, lanceolate, 1-3 mm long, lamina
adaxially glabrescent or with sparse, crispate tri-
chomes 0. 1-0.2 mm on laminar tissue, also some-
times with sessile glands. Sori medial, round, in-
dusia to 1 mm, minutely glandular at margin and
lacking trichomes, receptacle and sporangia gla-
brous.
TYPE— Peru, Amazonas, Prov. Chachapoyas,
20-17 km down E slope of Cerro Calla-Calla, Ed-
win & Schunke 3674 (holotype, F!; isotypes, GH!,
NY!).
Along roadside, middle elevation, known only
from the type.
This species differs from other members of sect.
Blennocaulon (T. cheilanthoides group) by the bi-
colorous scales along the rachis and costae abax-
ially. The scales are castaneous with a narrow tan
or whitish margin. Specimens lack long-septate
trichomes found in most specimens of T. cheilan-
thoides.
45. Thelypteris ruiziana (Klotzsch) A. R. Sm.,
comb. nov.
Polypodium ruizianum Klotzsch, Linnaea 20: 385.
1847. TYPE: Peru, Huanuco, ad Panatahua, Herb.
Ruiz no. 70 (holotype, B!; frag., BM!).
Dryopteris ruiziana (Klotzsch) C. Chr., Kongel. Danske
Vidensk. Selsk. Skr., Naturvidensk. Afd., ser. 7,
10: 152. 1913.
Stem probably suberect to erect, scales brown
to tan, dull, appressed, cellular detail not evident
at 30 x , ovate-lanceolate, glabrous on margins and
surface. Leaves several, clustered, 1 30-1 70 cm long,
probably not mucilaginous when young. Lamina
subcoriaceous, deeply 1 -pinnate-pinnatifid, prox-
imal pinnae not seen, probably subabruptly re-
duced and the lowest glanduliform. Petiole ca. 30
cm x 4 mm, tan and with scattered appressed
scales distally. Rachis with numerous septate tri-
chomes 1-1.5 mm long abaxially. Pinnae sessile,
14 x 1.5 cm, deeply pinnatifid to within 1 mm of
costae, segments ca. 2-3 mm wide, margin often
strongly revolute, basiscopic segment of larger pin-
nae often longer and more curved. Aerophores
small, tuberculiform. Buds lacking. Veins ca. 15
pairs per segment, usually immersed and appear-
ing dark in dried specimens. Indument on costae
and veins abaxially of moderately dense to dense,
septate trichomes 0.5-1 mm, costae also with nu-
merous light tan, amorphous, appressed scales,
laminar tissue glabrous, lamina adaxially gla-
brous. Sori medial, oblong, strongly confluent at
maturity, exindusiate, receptacle and sporangia
glabrous.
Montane forest, along roadsides, 1 900 m, Hua-
nuco.
Perhaps also Ecuador.
Huanuco: Huacachi, near Muna, Macbride 4177 (F).
46. Thelypteris nitens (Desv.) R. Tryon, Rhodora
69: 7. 1967.
Polypodium nitens Desv., Mem. Soc. Linn. Paris 6:
240. 1 827. TYPE: Peru, collector not known (ho-
lotype, P!; photos, GH, uc; frag., BM!).
Dryopteris nitens (Desv.) C. Chr., Kongel. Danske Vi-
densk. Selsk. Skr., Naturvidensk. Afd., ser. 7, 10:
142. 1913.
Dryopteris multiformis C. Chr., Kongel. Danske Vi-
densk. Selsk. Skr., Naturvidensk. Afd., ser. 7, 10:
142. 1913. TYPE: Ecuador, Mt. Pichincha, Mille
(holotype, P; isotype, uc!).
Lastrea nitens (Desv.) Copel., Gen. fil. 139. 1947.
Thelypteris multiformis (C. Chr.) Reed, Phytologia 1 7:
294. 1968.
Stem ascending to erect, scales brown, dull, ovate
to lanceolate, glabrous on margins and surface.
Leaves few, 30-75(-100) cm long. Lamina thick-
chartaceous, 1 -pinnate-pinnatifid (sometimes
2-pinnate in Ecuador), proximal l-3(-6) pairs of
pinnae abruptly reduced, the lowermost often less
than 1 mm, glanduliform. Petiole 1 5-30 cm x 2-
5 mm, stramineous and scaleless above the base.
Rachis glabrescent or deciduously pubescent with
flexuous, septate trichomes 1-2 mm abaxially.
Pinnae sessile, 9-15(-20) x (i.3_)2-3(-6) cm, op-
posite to subopposite, pinnatifid to ca. l(-3) mm
from costae, segments 4-6(-8) mm wide, the basal
basiscopic ones sometimes enlarged and dentate.
Aerophores absent or tuberculiform. Buds lacking.
Veins 6-12(-16) pairs per segment. Indument of
costae and veins abaxially of moderate spreading
or flexuous, acicular and often septate trichomes
mostly 1-2 mm, laminar tissue glabrous, glands
lacking, costal scales light brown, appressed,
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
35
amorphous, lamina adaxially glabrous. Sori me-
dial, round to slightly oblong, exindusiate, recep-
tacle and sporangia glabrous.
Montane rain forests and cloud forests, wet cliffs,
2200-3450 m, Cajamarca and Cuzco.
Ecuador and Peru.
Some specimens from Ecuador are much larger
than those from Peru and may even be 2-pinnate
with the pinnules auriculate or hastate at their
base.
Cajamarca: Prov. Contumaza, Quebrada de Canrra
(Contumaza-Toledo), Sagdstegui et al. 12617 (MO). Cuz-
co: Prov. Urubamba, bottom of Pojpoj waterfall, Davis
et al. 1740 (F), 1788 (F), 7797 (F). Prov. Urubamba,
Yucay, Herrera 713 (us, frag. GH).
47. Thelypteris proboscidea A. R. Sm., sp. nov.
A speciebus ceteris subg. Amauropeltae distinguenda
caulibus paleis glabris, costae paleis fulvis non clathratis,
aerophoris elongatis usque ad 3 mm longis ad basin pin-
narum, soris exindusiatis, petiolo et rhachide et costis
abaxialiter sparsim pubescentibus vel glabris, lamina
cglandulosa abaxialiter.
Stem ascending to erect, scales brown, some-
what shiny, ovate-lanceolate, glabrous on margins
and surface. Leaves few, clustered, 60-90 cm long.
Lamina chartaceous, 1 -pinnate-pinnatifid, proxi-
mal ca. 5 or more pairs of pinnae abruptly reduced,
the lowermost 1 mm long or less, glanduliform.
Petiole ca. 5-1 5 cm x 2-5 mm, tan to stramineous
and lacking scales above the base. Rachis glabrous
or very sparsely pubescent. Pinnae sessile, 8-15 x
1.5-2.3 cm, deeply pinnatifid to within 1 mm of
costae, segments 3-5 mm wide. Aerophores peg-
like, 1-3 mm. Buds lacking. Veins 7-12 pairs per
segment. Indument on costae and veins abaxially
lacking or of very sparse, ascending trichomes 0. 1-
0.3 mm, laminar tissue glabrous, glands lacking,
costal scales lacking or sparse, tan, appressed, non-
clathrate, lamina adaxially glabrous or with very
sparse trichomes 0. 1-0.2 mm on costules and veins.
Sori supramedial, round, exindusiate, receptacle
glabrous, sporangia glabrous.
TYPE— Peru, Ancash, Prov. Bolognesi, cerca a
Llamac, Cerrate 2366 (holotype, GH!; isotype, 2
sheets, USM).
Along streams, 3300-3800 m, Lambayeque and
Ancash.
Known only from northern Peru.
The amorphous costal scales and the pro-
nounced peglike aerophores up to 3 mm long at
the base of the pinnae indicate that this may be a
member of sect. Blennocaulon. Important char-
acters distinguishing it from other members of that
section include: exindusiate sori; epilose or nearly
epilose costae, rachis, and petiole; and eglandular
abaxial lamina.
Lambayeque: Prov. Ferrenafe, ca. 4 km NW of Inca-
huasi, below Cerro Punamachay on trail to Laguna Hual-
taco, Dillon & Skillman 4156 (F, uc).
Comments
Following completion and initial editing of the
manuscript, two more new species were discov-
ered, as well as a range extension for a species
previously known only from Ecuador. Because it
was impractical at that point to revise the key and
order of the species, these species are included here
as addenda.
Thelypteris arrecta A. R. Sm., sp. nov.
Inter species subg. Amauropeltae sect. Phacelothricis,
trichomatibus fasciculatis T. canadasii (Sodiro) Alston
similis sed differ! caule valde erecto foliis distantibus,
aerophoris brevioribus ad basin pinnarum, aerophoris
ad basin costularum absentibus, venis segmentis minus
numerosis trichomatibus minus manifeste fasciculatis ad
rhachim et costas abaxialiter, paleis magis numerosis ad
costas.
Stem strictly erect, more than 1 5 cm long, scales
brown, dull to somewhat shiny, ovate-lanceolate,
with scattered trichomes 0. 1-0.25 mm on margins
and surface. Leaves few, distant, ca. 90 cm long.
Lamina chartaceous, 1 -pinnate-pinnatifid, proxi-
mal ca. 3 pairs of pinnae abruptly reduced, the
lowermost pair less than 1 mm long, glanduliform.
Petiole ca. 1 5 cm x 3-4 mm, brownish, with nu-
merous appressed amorphous scales and numer-
ous trichomes in groups of 2-4. Rachis with nu-
merous clustered trichomes mostly 0.1-0.2 mm
and a few appressed scales. Pinnae sessile, to 12
x 1.6 cm, deeply pinnatifid to within 1 mm of
costa. Segments mostly 2-2.5 mm wide, proximal
pairs of lower pinnae not or only slightly reduced.
Aerophores scalelike or peglike at pinna bases, to
2 mm long. Buds lacking. Veins up to ca. 1 2 pairs
per segment. Indument of costae and costules
abaxially of moderately dense, spreading, acicular
trichomes mostly 0. 1-0.2 mm, these often in pairs
or threes, costae also with appressed nonclathrate
36
FIELDIANA: BOTANY
scales, lamina adaxially of ascending to appressed
trichomes 0.1-0.4 mm on costules, veins, and
laminar tissue. Sori inframedial to medial, round
to slightly oblong, indusia absent or minute, less
than 0. 1 mm, receptacle or indusial fragment with
a few trichomes 0. 1 mm, sporangia glabrous.
TYPE— Peru, Cuzco, Paucartambo, carretera
Acjancaco-Pilcopata, quebrada a la altura del se-
gundo puente de Pillahuata a La Esperanza, Par-
que Nacional Manu, Leon 2223 (holotype, uc!;
isotype, USM).
Montane rain forests, 2650 m, Cuzco.
Known only from the type.
The species epithet derives from the strong re-
semblance of T. ctenitoides to some species of
Ctenitis, many of which have similar scales and
often crenulate segments. It is one of the most
distinctive species in subg. Amauropelta, differing
especially by the numerous subclathrate scales
covering the axes abaxially. These scales suggest
placement in sect. Lepidoneuron, but the large in-
dusia cast doubt on this affinity.
Thelypteris exuta A. R. Smith, Fl. Ecuador 18: 45.
1983. TYPE: Ecuador, Tungurahua, San An-
tonio, Tate 571 (holotype, us!).
Thelypteris ctenitoides A. R. Sm., sp. nov.
A speciebus ceteris subg. Amauropeltae differt caule
valde erecto apice paleis glabris, segmentis crenulatis,
soris submarginalibus, indusiis glabris, 0.5-1 mm in dia-
metro, et praesertim petiolo et rhachide et costis cos-
tulisque abaxialiter paleis numerosis ovato-lanceolatis
atrobrunneis subclathratis sed trichomatibus fere nullis.
Stem strictly erect, ca. 12 cm long, scales dark
brown, subclathrate, somewhat shiny, ovate-lan-
ceolate, lacking trichomes. Leaves fasciculate, ca.
50 cm long. Lamina chartaceous, 1-pinnate-pin-
natifid, proximal 2-4 pairs of pinnae abruptly re-
duced, the lowermost pair less than 5 mm long.
Petiole ca. 8-15 cm x 2-3 mm, stramineous to
tan, with numerous appressed ovate-lanceolate
scales and sparse trichomes. Rachis similar to pet-
iole in indument. Pinnae sessile, to 8 x 1.8 cm,
deeply incised to within 1 mm of costa. Segments
mostly 2-3 mm wide, crenulate, proximal pair on
lower pinnae not reduced. Aerophores peglike at
pinna bases, to 5 mm long. Buds lacking. Veins
up to ca. 1 0 pairs per segment. Indument on costae
and costules abaxially of numerous, appressed,
ovate-lanceolate, subclathrate scales, trichomes
lacking or sparse, 0.1-0.2 mm long, lamina adax-
ially with appressed trichomes 0.1-0.2 mm long,
on costules, veins, and a few on laminar tissue.
Sori submarginal, round, indusia large, 0.5-1 mm
in diameter, lacking trichomes, sporangia gla-
brous.
TYPE— Peru, Amazonas, Prov. Bagua, Cordille-
ra Colan NE of La Peca, Barbour 3416 (holotype,
MO!).
Humid pajonal, 2900-3100 m, Amazonas.
Known only from the type.
Stem suberect to erect, scales light brown, dull,
ovate-lanceolate, glabrous or nearly so. Leaves fas-
ciculate, ca. 30-40(-65) cm long. Lamina thin-
chartaceous, 1 -pinnate-pinnatifid, proximal 2-3
pairs of pinnae abruptly reduced, the lowermost
pair ca. 1 mm long. Petiole 10-20 x l-1.5(-3)
mm, stramineous, with a few scales at base, oth-
erwise glabrous. Rachis glabrous. Pinnae sessile,
opposite or subopposite, 4-6(-10) x 1.2-1.5(-2)
cm, deeply incised to ca. 1 mm from costa. Seg-
ments 2-3(-4) mm wide, proximal pair of pinnae
slightly reduced. Aerophores absent. Buds absent.
Veins 6-9 pairs per segment. Indument lacking
abaxially on costae and costules, lamina adaxially
glabrous except for trichomes 0.4-0.6 mm along
costa and costules. Sori medial, round, indusia
0.2-0.3(-0.5) mm in diameter with marginal tri-
chomes 0.2 mm and glands 0.05 mm, sporangia
glabrous.
Montane rain forest, 1 800 m, San Martin.
Ecuador and Peru.
The sole Peruvian collection differs from the
Ecuadorian ones in having much more reduced
proximal pinnae, setose-margined indusia, and
trichomes along the costules adaxially.
San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba
road, Km 390, Venceremos, D. Smith 4426 (MO).
The following collections do not appear to match
any known species of subg. Amauropelta; they may
represent undescribed species or perhaps hybrids.
All belong to large and difficult species groups or
complexes that are in need of revision before names
can be applied with confidence. Additional col-
lections are necessary before adequate descriptions
and diagnoses can be written.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
37
38
FIELDIANA: BOTANY
C. Schunke 59, 454 (F)— Junin, Chanchamayo
Valley, 1000 m. Most similar to T. opposita but
differing from that in the fewer, more widely spaced
reduced proximal pinnae and the presence of flex-
uous, septate trichomes on the abaxial costae. These
differences suggest a relationship to T. balbisii,
which has generally much wider and longer pin-
nae.
Ridoutt (GH)— Camino a Pucallpa, Km 209.
Most similar to T. opposita but differing in the
dense, short pubescence and numerous glands on
the adaxial lamina, the sparser pubescence on the
abaxial costae, the sunken, darkened veins abax-
ially, and the lack of sessile resinous glands abax-
ially. In characters of the adaxial lamina, it is very
similar to T. micula, which is a much smaller,
thinner-textured species.
Dudley 11284A (GH)— Cuzco, Prov. La Con-
vencion, ca. 17 km NE from Hda. Luisiana and
Apurimac River, above Camp 3, 2100 m. This
belongs to sect. Uncinella, as indicated by the
abundant hamate trichomes on the costae and
lamina abaxially and the dense appressed, short
trichomes on the adaxial lamina. Peglike aero-
phores are present at the pinna bases, and there
are persistent scales along the rachis and costae
abaxially. A description of this probable new spe-
cies must await the discovery of fertile material.
He. Thelypteris subg. Cyclosorus.
Thelypteris subg. Cyclosorus (Link) Morton, Amer.
Fern J. 53: 153. 1963. Figure 4.
Cyclosorus Link, Hort. Reg. Hot. Berol. 2: 128. 1833.
TYPE: Cyclosorus gongylodes (Schkuhr) Link =
Thelypteris interrupta (Willd.) Iwatsuki.
Christella Leveille, Fl. Kouy-tcheou 472. 1915. TYPE:
Christella parasitica (L.) Leveille = Thelypteris
parasitica (L.) Tard.
Amphineuron Holttum, Blumea 19: 45. 1971. TYPE:
Amphineuron opulentum (Kaulf.) Holttum =
Thelypteris opulenta (Kaulf.) Fosberg.
Lamina 1 -pinnate-pinnatifid, pinnae pinnatifid
half their width or more; proximal pinnae not re-
duced or up to ca. 6 pairs gradually reduced in a
few spp., distal pinnae gradually shortened and
lamina with a confluent pinnatifid apex or abrupt-
ly shortened and with a subconform terminal pin-
na; aerophores and buds lacking. Veins simple,
those from adjacent segments meeting margin at
or near the sinus, or connivent at or just below
sinus, or one pair united at an obtuse angle below
sinus with an excurrent vein to sinus. Indument
of acicular, nonseptate trichomes. Sori round, with
a persistent round-reniform indusium at least 0.5
mm in diameter; sporangia glabrous or with a stip-
itate gland on the stalk. Spores with a few prom-
inent connected winglike ridges, or many discon-
nected ones, or somewhat echinate. x = 36.
As circumscribed here, this includes Amphineu-
ron (Holttum, 1977) and Christella (Holttum,
1976), two predominantly Old World groups. The
type of Cyclosorus, which is a synonym of T. in-
terrupta, belongs to a small group of marsh-in-
habiting species not closely related to the rest of
the subgenus in Peru.
Subgenus Cyclosorus comprises about 20 New
World species and a much larger number of Old
World ones. In general, species of Cyclosorus are
common and even weedy along roadsides and
ditches. One introduced species, T. dentata, is
widely naturalized in the Neotropics but is so far
rarely collected in Peru. A second naturalized spe-
cies, T. opulenta, has become one of the most
abundant ferns around Iquitos.
Those species with reduced proximal pinnae,
namely T. dentata, T. hispidula, and T. conspersa,
can be distinguished from subg. Amauropelta by
veins that unite below the sinus or are connivent
at the sinus.
References
HOLTTUM, R. E. 1976. Studies in the family The-
lypteridaceae XI. The genus Christella Leveille,
sect. Christella. Kew Bull. 31: 293-339.
HOLTTUM, R. E. 1977. Studies in the family The-
lypteridaceae XII. The genus Amphineuron
Holttum. Blumea 23: 205-218.
SMITH, A. R. 1971. Systematics of the neotrop-
ical species of Thelypteris section Cyclosorus.
Univ. Calif. Publ. Bot. 59: 1-143.
FIG. 4. Subgenus Cyclosorus. Thelypteris depilata: a, stem and portion of petiole, apical portion of lamina; b,
portion of petiole base; c, costa and pinna segments, abaxial side; d, costa and pinna segments, adaxial side. (From
Sounders 570, holotype, F.)
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
39
Key to Species of subg. Cyclosorus
a. Basal veins from adjacent segments united at an obtuse angle below sinus with an excurrent vein to
sinus; costules, veins, and often lamina adaxially pubescent b
b. Proximal pinnae the longest or nearly so; costae abaxially with ovate scales; lamina abaxially with
sessile orangish or reddish, hemispherical glands 48. T. interrupta
b. Proximal pinnae reduced; costae abaxially scaleless; lamina abaxially lacking glands or the glands
short-stipitate, light yellow c
c. Costae abaxially with most trichomes more than 0.3 mm long with some exceeding 0.5 mm;
petiole and rachis stramineous to tan 49. T. hispidula
c. Costae abaxially with predominantly short trichomes uniform in length, mostly 0.1-0.2 mm
long; petiole and rachis usually purplish brown 50. T. dentata
a. Basal veins from adjacent segments connivent at, or running to, sinus; costules, veins, and lamina
adaxially glabrous or pubescent d
d. Rhizome suberect to erect, hidden by old leaf bases; proximal pinnae often auriculate at base . . .
53. T. patens
d. Rhizome creeping, readily visible; proximal pinnae not auriculate 3
e. Several pairs of proximal pinnae greatly reduced; lamina adaxially with numerous trichomes
between veins 51. T. conspersa
e. Proximal pinnae not reduced; lamina adaxially glabrous between veins f
f. Glands sulfur-yellow, sessile, present abaxially along veins and costules, especially toward
segment tips, and among sporangia; costae abaxially scaleless 52. T. opulenta
f. Glands absent along veins and costules abaxially and among sporangia; costae abaxially
with or without scales g
g. Costae abaxially without trichomes 54. T. depilata
g. Costae abaxially sparsely to densely pubescent h
h. Costal trichomes mostly greater then 0.2 mm, dense; costal scales numerous
55. T. clivalis
h. Costal trichomes ca. 0. 1 mm, moderately dense; costal scales relatively few
56. T. grandis
48. Thelypteris interrupta (Willd.) Iwatsuki, Jap.
J. Bot. 38: 314. 1963.
Pteris interrupta Willd., Phytogr. 13, pi. 10, fig. 1.
1794. TYPE: Southern India, Klein (holotype, B,
Herb. mild. 19770; microfiche, uc).
Polypodium tottum Thunb., Prodr. pi. cap. 1 72. 1 800.
TYPE: South Africa, Thunberg Herb. 24724 (ho-
lotype, UPS!).
Aspidium gongylodes Schkuhr, 24. Kl. Linn. Pfl.-Syst.
[Krypt.Gew.] 1: I93,pl.33c. 1804. TYPE: British
Guiana, Essequebo, collector unknown (holo-
type, HAL?; isotype, s).
Cyclosorus gongylodes (Schkuhr) Kuntze, Revis. gen.
pi. 2: 811. 1891.
Thelypteris gongylodes (Schkuhr) Small, Ferns s.e.
States 248. 1938.
Thelypteris totta (Thunb.) Schelpe, J. S. African Bot.
29: 91. 1963.
Stem long-creeping, to 2 m or more, dark brown
to blackish, nearly devoid of scales. Leaves few,
l-5(-9) cm distant, mostly 50-100(-1 80) cm long.
Lamina chartaceous to subcoriaceous, 1 -pinnate-
pinnatifid, proximal pinnae the longest or nearly
so. Petiole to 50(-100) cm x 3^*(-6) mm, tan to
stramineous and lacking scales above the base.
Rachis glabrescent or with scattered trichomes
mostly 0.2-0.3 mm abaxially. Pinnae sessile or
short-stalked, 7-15(-30) x 0.8-1. 3(-2.2) cm, shal-
lowly pinnatifid to ca. 2-4 mm from costae, not
auriculate at base, segments 2.5-4 mm wide. Aero-
phores absent. Buds lacking. Veins 8-1 2(-l 8) pairs
per segment, lowermost pair from adjacent seg-
ments connivent or obtusely uniting below sinus.
Indument on costae, veins, and laminar tissue
abaxially of very sparse to dense, spreading, acic-
ular or crispate trichomes mostly 0. 1-0.3 mm, also
with sessile orange or red shiny globular glands
0.1 mm on costules, veins, and laminar tissue,
costal scales tan, ovate, to 2 mm, adaxially the
lamina glabrous or nearly so. Sori medial, round,
indusia glabrous or usually with trichomes and
40
FIELDIANA: BOTANY
sessile glands, sporangial stalks often with stalked
glands.
Along streams and edges of lakes, marshes, on
floating islands, 100-260 m, Loreto and Madre de
Dios.
Florida; Antilles; Mexico to Panama; Colombia
to Guianas and northern Argentina; Paraguay;
tropical and subtropical Africa, Asia.
Loreto: Prov. Maynas, 4 km S of Iquitos, Tryon &
Tryon 5211 (F, GH, us). Madre de Dios: Prov. Tambo-
pata, ca. 30 air km SSW of Puerto Maldonado at efflu-
ence of Rio La Torre (Rio D'Orbigny)/Tambopata, Bar-
hour 4908 (MO, uc, USM), Young 3 3 (MO, uc, USM). Cocha
Cashu, Rio Manu, between Panagua and Tayakome, R.
Foster et al. 3374 (F), 3375 (GH, USM). Laguna Coco
Cocha, 5.2 km E of lodge, Funk et al. 8399 (F, USM).
49. Thelypteris hispidula (Decne.) Reed, Phyto-
logia 17: 283. 1968.
Aspidium hispidulum Decne., Nouv. Ann. Mus. Hist.
Nat. 3: 346. 1834. TYPE: Timor, Guichenot (ho-
lotype, P).
Nephrodium quadrangulare Fee, (Mem foug. 5) Gen.
fil. 308. 1852. TYPE: French Guiana, Leprieur
182 (isotypes, NY!, P!).
Dryopteris parasitica (L.) var. glanduligera Rosenst.,
Repert. Spec. Nov. Regni Veg. 7: 304. 1909.
TYPE: Peru, (San Martin), Tarapoto, Spruce 4039
(holotype, P!; isotypes, BM!, P!).
Dryopteris quadrangularis (Fee) Alston, J. Bot. 75:
253. 1937.
Thelypteris quadrangularis (Fee) Schelpe, J. S. African
Bot. 30: 196. 1964.
Christella hispidula (Decne.) Holttum, Kew Bull. 31:
312. 1976.
Stem short-creeping to ascending or erect, scales
brown, shining, linear-lanceolate, setose on mar-
gins and surfaces. Leaves several, fasciculate,
mostly 50-100 cm long. Lamina chartaceous,
1 -pinnate-pinnatifid, proximal 1—4 pairs of pinnae
slightly to greatly reduced, the lowermost to 2 cm
or less. Petiole 10-40 cm x 2-5 mm, tan to stra-
mineous and lacking scales above the base. Rachis
pubescent abaxially, trichomes 0.4-1 mm. Pinnae
sessile, 5-15 x l-2(-2.5) cm, deeply pinnatifid
1.5-3 mm from costae, proximal ones often au-
riculate at acroscopic base, segments 1-4 mm wide.
Aerophores absent. Buds lacking. Veins 6—9(-l 1)
pairs per segment, lowermost pair from adjacent
segments united below sinus with an excurrent
veinlet 1-2 mm to sinus. Indument on costae, veins,
and laminar tissue abaxially of moderately dense
to dense, spreading, acicular trichomes 0.3-1 mm,
often also with sessile to short-stipitate, light yel-
lowish glands, costal scales lacking, adaxially with
numerous acicular trichomes 0.3-1 mm and often
glands on costules, veins, and laminar tissue. Sori
medial, round, indusia pubescent, sporangial stalks
obscurely stipitate-glandular.
Lowland and montane rain forests, pastures,
along trails and edges of woods, 100-1500 m,
Lambayeque and Amazonas, south to Cuzco and
Madre de Dios.
Southeastern United States; Antilles; Mexico to
Panama; Colombia to Guianas and Bolivia; north-
ern Argentina; tropical and subtropical Africa and
Asia.
All Peruvian material seen is referable to what
I previously called T. quadrangularis var. quad-
rangularis (Smith, 1971). However, the relation-
ship between var. quadrangularis in the New World
and T. hispidula in the Old World is unstudied,
and I prefer not to make a varietal combination
at this time. Thelypteris hispidula var. inconstans
(C. Chr.) Proctor (Antilles) and var. versicolor (R.
St. John) Lellinger (southeastern United States) are
sufficiently distinct to be recognized. The status of
Thelypteris quadrangularis var. repens A. R. Sm.
(southern Brazil, Uruguay, northeastern Argenti-
na) is also problematic vis-a-vis Paleotropical
specimens.
Lambayeque: Entre Beatita de Humay y Km 38, car-
retera Olmes-Maranon, Lopez et al. 4044 (GH). Ama-
zonas: Prov. Bagua, Chomza and environs, ca. 5 km S
of La Peca, Barbour 4313 (MO, USM). San Martin: Prov.
San Martin, Tarapoto, San Antonio de Cumbaza, Sagds-
tegui 0219 (GH). San Roque, Ll. Williams 7079 (F, us).
Loreto: Gamitanacocha, Rio Mazan, Schunke 203 (F,
GH, uc, us). Mishuyacu, near Iquitos, Klug 1335 (F, NY,
us). Prov. Maynas, Rio Itaya, 10 km S of Iquitos, Tryon
& Tryon 5207 (F, GH, u, USM). Huanuco: Hacienda at
mouth of Rio Chinchao, Macbride 5028 (F, us). Pasco:
Rio Paucartambo valley, near Perene bridge, Killip &
Smith 25308 (NY, us). Quillasu, Soukup 3295 (GH). Ju-
nin: La Merced-Chanchamayo, Soukup 1029 (F). Aya-
cucho: Estrella, between Huanta and Rio Apurimac, Kil-
lip & Smith 22621 (F, GH, NY, us). Cuzco: Pilcopata,
Atalaya, Paucartambo, Nunez 6857 (MO). Madre de Dios:
Prov. Manu, Pinipiiii Defensa, Vargas 11611 (GH), 11612
(GH).
50. Thelypteris dentata (Forssk.) E. St. John,
Amer. Fern J. 26: 44. 1936.
Polypodium dentatum Forssk., Fl. Aegypt.-arab. 185.
1775. TYPE: southwestern Arabia, Yemen,
Forsskal (holotype, C!).
Polypodium mollejacq., Collectanea 3: 188. 1789, not
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
41
Schreb., 1771, not All., 1785. TYPE: cultivated
specimen from gardens at Schoenbrunn (holo-
type, w!).
Dryopteris mollis (Sw.) Hieron., Hedwigia 46: 348.
1907.
Dryopteris dentata (Forssk.) C. Chr., Kongel. Danske
Vidensk. Selsk. Skr., Naturvidensk. Afd., ser. 8,
6: 24. 1920.
Cyclosorus dentatus (Forssk.) Ching, Bull. Fan Mem.
Inst. Biol., Bot. 8: 206. 1938.
Christella dentata (Forssk.) Brownsey & Jermy, Brit.
Fem Gaz. 10: 338. 1973.
Stem short-creeping, scales brown, shining, lin-
ear-lanceolate, setose on margins and surfaces.
Leaves several, slightly dimorphic with the fertile
having longer petioles and narrowed pinnae, most-
ly 50-100(-135) cm long. Lamina chartaceous,
1 -pinnate-pinnatifid, proximal 2-6 pairs of pinnae
slightly to often greatly reduced, the lowermost
often less than 2 cm. Petiole 1 5-45 cm x 2-5 mm,
tan to purplish brown and lacking scales above the
base. Rachis pubescent abaxially, trichomes most-
ly 0.2-0.4 mm. Pinnae sessile, 7-15 x 1-2.5 cm,
deeply pinnatifid 2-4 mm from costae, proximal
ones often auriculate at acroscopic base, segments
2-4 mm wide. Aerophores absent. Buds lacking.
Veins 6-1 1 pairs per segment, basal pair from ad-
jacent segments united below sinus with an ex-
current veinlet 2-3 mm to sinus. Indument on
costae, veins, and laminar tissue abaxially of mod-
erately dense to dense, uniform, spreading, acic-
ular trichomes 0. 1-0.2 mm, glands and costal scales
lacking, adaxially with numerous acicular tri-
chomes 0.1-0.3 mm on costules, veins, and lam-
inar tissue. Sori medial, round, indusia pubescent,
sporangial stalks obscurely stipitate-glandular.
Lowland forests, especially along trails, in ditch-
es, pastures, 200-600 m, Loreto and Madre de
Dios.
Southeastern United States; Antilles; southern
Mexico to Panama; Colombia to Venezuela and
Bolivia; Brazil; Argentina; tropical and subtropical
Africa, Asia, islands of the Pacific.
This is introduced and widely naturalized in the
New World (Strother & Smith, 1970, Taxon 19:
871-874) but has not reached Andean countries
until recently. It appears still to be uncommon in
Peru but is likely to spread rapidly in the future.
Thelypteris dentata can be easily confused with T.
hispidula, with which it has sometimes been com-
bined.
Loreto: Prov. Iquitos, Procedencia Andoas, 23 Oct
1979, Ayala 2124 (uc). Prov. Alto Amazonas, carretera
Oleoducto secundario entre Andoas y Capahuari Sur, 1 3
Sep 1979, Diaz & Jaramillo 1364 (MO). Madre de Dios:
Prov. Manu, Rio Salvation, 6 Dec 1986, Nunez 6588
(MO).
51. Thelypteris conspersa (Schrader) A. R. Sm.,
Univ. Calif. Publ. Bot. 59: 60. 1971.
Nephrodium conspersum Schrader, Gott. gel. Anz.
1 824: 869. LECTOTYPE (chosen by Smith, 1971,
p. 60): Brazil, near Espirito Santo, Barra de Fucu,
lVied-Neuwied(BR\; photo, uc; isolectotypes, BR!,
L).
Dryopteris bangii C. Chr., Kongel. Danske Vidensk.
Selsk. Skr., Naturvidensk. Afd., ser. 7, 4: 333.
1907. TYPE: Bolivia, Yungas, near Coroica, Bang
2321 (holotype, c; isotypes, B!, GH!, LD!, us!).
Stem short-creeping, scales brown, shining, lin-
ear-lanceolate, setose on margins and surfaces.
Leaves several, mostly 75-150 cm long. Lamina
chartaceous, 1 -pinnate-pinnatifid, proximal 3-6
pairs of pinnae gradually reduced, the lowermost
to 3 cm or less. Petiole 30-60 cm x 3-6 mm,
stramineous and lacking scales above the base.
Rachis densely pubescent abaxially, trichomes 0.5-
1.2 mm. Pinnae sessile, (7-) 10-20 x 1.5-2.2 cm,
deeply pinnatifid 1-3 mm from costae, proximal
ones often auriculate at acroscopic base, segments
2—4 mm wide. Aerophores absent. Buds lacking.
Veins 8-12 pairs per segment, lowermost pair from
adjacent segments running to sinus or connivent
just below sinus, occasionally united below sinus
with an excurrent vein up to ca. 1 mm. Indument
on costae, veins, and laminar tissue abaxially of
usually dense, spreading, acicular trichomes most-
ly 0.5-1 mm, sometimes also with short-stipitate,
light yellowish glands, less often the glands more
numerous than trichomes, costal scales lacking,
adaxially with numerous acicular trichomes to 1
mm on costules, veins, and laminar tissue, these
sometimes replaced by glands. Sori medial, round,
indusia densely pubescent, sporangial stalks ob-
scurely stipitate-glandular.
Montane forests, meadows, 700-1300 m, San
Martin, Junin, and Cuzco.
Hispaniola; Panama; Venezuela, Colombia;
Peru; Bolivia; southern Brazil; northern Argenti-
na; Paraguay; Uruguay.
Two collections, Coronado 260 and Soukup
1159, differ in being only sparsely pubescent but
with numerous short-stipitate glands.
San Martin: Hera near Moyobamba, Woytkowski
35312 [U.C. Bot. Gard. ace. 55.080] (GH, uc). Junin:
42
FIELDIANA: BOTANY
Near San Ramon, Coronado 260 (GH). Cuzco: Prov. La
Convention, Ichiquiato, Vargas 22332 (GH). Potrero, 8
km W of Quillabamba, Tryon & Tryon 5391 (F, GH, u).
Empalizaota, Biies 1719 (GH). Department Unknown:
Soukup 1159(F).
52. Thelypteris opulenta (Kaulf.) Fosberg, Smith-
sonian Contr. Dot. 8: 3. 1972.
Aspidium opulentum Kaulf., Enum. fil. 238. 1824.
TYPE: Guam, Chamisso (holotype, LE).
Aspidium extensum Blume, Enum. pi. jav. 156. 1828.
TYPE: Java, Pulu Pinang, collector not stated
(holotype, L!; photo, uc).
Dryopteris extensa (Blume) Kuntze, Rev. gen. pi. 2:
812. 1891.
Thelypteris extensa (Blume) Morton, Amer. Fern J.
49: 113. 1959.
Amphineuron opulentum (Kaulf.) Holtt., Blumea 19:
45. 1971.
Stem long- to short-creeping, scales brown,
shining, linear-lanceolate, setose on margins and
surfaces. Leaves few, mostly 90-200 cm long.
Lamina chartaceous, 1 -pinnate-pinnatifid, proxi-
mal pinnae not reduced or with 1 pair greatly re-
duced, lamina apex somewhat prolonged. Petiole
40-100 cm x 3-6 mm, tan to stramineous and
lacking scales above the base. Rachis sparsely pu-
bescent and stipitate-glandular abaxially, tri-
chomes and glands 0.1 mm or less. Pinnae sessile
or short-stalked to 2 mm, 15-30(-40) x 1.3-3
(-3.5) cm, deeply pinnatifid 2-4 mm from costae,
proximal ones not auriculate at base, segments 2-
4 mm wide. Aerophores absent. Buds lacking. Veins
8-14 pairs per segment, lowermost pair from ad-
jacent segments united below sinus with an ex-
current veinlet to ca. 1 mm or veins connivent
near sinus. Indument on costae, veins, and some-
times laminar tissue abaxially of sparse to mod-
erately dense, spreading, acicular trichomes less
than 0. 1 mm and often a few longer trichomes to
0.5 mm on axes, also with numerous sulfur-yel-
low, sessile glands especially along costules and
veins, often also of short-stipitate, light yellowish
glands, costal scales lacking, adaxially with scat-
tered acicular trichomes to ca. 0.5 mm on costules
and veins. Sori supramedial, round, often confined
to pinna lobes, indusia with marginal glands and
sometimes a few acicular trichomes, sporangial
stalks obscurely glandular.
Lowland rain forests, secondary forests, old
plantations, along trails, and in partially disturbed
areas, 100-650 m, Amazonas, Loreto, San Martin,
Huanuco, Pasco, Ucayali, Cuzco, and Madre de
Dios.
Jamaica; Lesser Antilles; Costa Rica and Pan-
ama; Colombia to Guianas and Peru; native to
Africa and Asia.
This has become an extremely common natu-
ralized fern in parts of Peru, and I have seen 45
collections, 28 of them from Loreto. Only one of
the collections was made prior to 1964, that in
1956.
Amazonas: Prov. Bagua, ca. 40-43 km NE of Chiriaco,
Barbour 4529 (MO, USM). San Martin: San Martin, 7-15
km E of Shapojo on road to Chazuta, Knapp et al. 7264
(F, MO). Loreto: Quebrada Shanuce above Yurimaguas,
Croat 18050 (F, MO, uc, USM). 13 km SW of Iquitos,
Croat 18595 (F, MO, uc). Rio Itaya, 10 km S of Iquitos,
13 Aug 1956, Tryon & Tryon 5202 (F, GH, u). Huanuco:
Road from Tingo Maria to "Monson" (Monzon), Croat
57921 (MO, uc). Pasco: Prov. Oxapampa, Quebrada Cas-
tilla near Villa America, on Omaiz River, Leon & Young
1021a (uc, USM). Ucayali: Arboretum of Bosque von
Humboldt Experimental Station, Km 86 on Pucallpa-
Tingo Maria road, D. Smith 1223 (MO, uc). Cuzco: Prov.
Quispicanchi, Inambari, Vargas 15362 (GH). Madre de
Dios: Prov. Tambopata, ca. 30 air km SSW of Puerto
Maldonado, Barbour 4962 (F, MO, uc). Prov. Manu, Manu
Park, Cocha Cashu uplands, Nunez 5859 (MO).
53. Thelypteris patens (Sw.) Small, Ferns s.e.
States 243. 1938.
Stem suberect to usually erect, scales tan to
brown, dull to shining, lanceolate to ovate-lan-
ceolate, glabrous or setose on margins and sur-
faces. Leaves several, fasciculate, mostly 45-150
cm long. Lamina chartaceous, 1 -pinnate-pinnati-
fid, proximal pinnae the longest or nearly so, or
with one pair slightly to rarely greatly reduced.
Petiole 1 5-60 cm x 3-6 mm, tan to stramineous
and lacking scales above the base. Rachis pubes-
cent abaxially, trichomes 0.4-1 mm. Pinnae ses-
sile, 10-30 x (1-) 1.5-3 cm, deeply pinnatifid 1-
2 mm from costae, proximal ones often auriculate
at acroscopic base or elongate parallel to rachis,
segments 2-4 mm wide. Aerophores absent. Buds
lacking. Veins 8-15(-21) pairs per segment, low-
ermost pair from adjacent segments running to
sinus or distal vein of a pair meeting margin just
above sinus, infrequently lowermost pair conni-
vent just below sinus. Indument on costae, veins,
and laminar tissue abaxially of moderately dense
to dense, spreading, acicular trichomes 0.2-0.8
mm, often also of sessile to short-stipitate, light
yellowish glands, costal scales lacking, lamina
adaxially glabrous except along costae or with acic-
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
43
ular trichomes 0.3-0.5 mm on costules and some-
times veins. Sori medial, round, indusia usually
densely pubescent, trichomes 0.2-0.5 mm long,
sporangial stalks obscurely stipitate-glandular or
lacking glands.
The two varieties encountered in Peru are gen-
erally readily distinguishable and could perhaps
stand as species. A third entity, var. dissimilis
(Schrader) A. R. Sm., is restricted to southern Bra-
zil, Paraguay, and northeastern Argentina.
Key to Varieties
a. Lamina adaxially glabrous on costules and veins; petiole base scales tan, ovate to ovate-lanceolate,
glabrous or nearly so, dull 53a. var. patens
a. Lamina adaxially with trichomes on costules and veins; petiole base scales brown, lanceolate, pu-
bescent, shining 53b. var. smithiana
53a. Thelypteris patens var. patens.
Polypodium patens Sw., Prodr. 133. 1788. LECTO-
TYPE (chosen by Smith, Univ. Calif. Publ. Bot.
59: 72. 1971): Jamaica, Swartz (s!; photo, uc).
Aspidium stipulareWi\\A., Sp. pi. ed. 4, 5: 239. 1810.
TYPE: Plumier, t. 23.
Aspidium macrourum Kaulf., Flora 6: 365. 1823.
TYPE: Martinique, Sieber 354 (isotypes, L, M!,
MO!).
Lastrea scabriuscula Presl, Epim. hot. 35. 1849, not
Davenp., 1896. LECTOTYPE (chosen by Chris-
tensen, 1913): Brazil, ad Rio de Janeiro, Mikan
(PRC).
Dryopteris patens (Sw.) Kuntze, Rev. gen. pi. 2: 813.
1891.
Thelypteris patens var. scabriuscula (Presl) A. R. Sm.,
Univ. Calif. Publ. Bot. 59: 76. 1971.
Lowland and montane forests, roadsides, wet
ditches, 200-1900 m, Piura, Lambayeque, Caja-
marca, San Martin, Loreto, Huanuco, Lima, Pas-
co, Junin, Cuzco, and Puno.
Florida; Antilles; Mexico to Panama; Colombia
and Venezuela to southern Brazil and Bolivia.
Piura: Hills of Chiarnique ca. 20 km ENE of Naupe,
Barbour2161 (F, MO, uc, USM). Lambayeque: Prov. Lam-
bayeque, Quebrada Onda, Llatas Quiroz 1124 (F). Ca-
jamarca: Prov. Cajamarca, ca. 9 km W of San Juan and
46 km SW of Cajamarca on road to San Pedro de Lloc,
Dillon & Whalen 4069 A (F, GH, uc, USM). San Martin:
Prov. San Martin, Tarapoto, Woytkowski 35075 (MO,
uc). Loreto: Lower Rio Huallaga, LI. Williams 5108 (F).
Huanuco: Prov. Leoncio Prado, along road to Monzon
above Rio Huallaga across bridge from Tingo Maria,
Croat 50973 (MO). Lima: Chosica, Bryan 22 (F). Pasco:
Pozuzo, Macbride 4601 (F, NY, us). Junin: Prov. Tarma,
cerca a la Merced, Cerrate 2855 (GH). Cuzco: Prov. La
Convention, Pawac Portage, one hour float below Po-
mobamba, Davis et al. 1264 (GH). Puno: Prov. Sandia,
Yanamayo, Soukup & Lopez [USM 14656] (GH).
53b. Thelypteris patens var. smithiana Ponce,
Darwiniana 28: 373. 1987. TYPE: Argentina,
Prov. Salta, Dpto. Oran, Zanja Honda-Que-
brada Sierra de Aguarague, Abbiatti & Claps
273 (holotype, LP).
Nephrodium schizotis Hooker, Sp. fil. 4: 107. 1862.
TYPE: Peru, (San Martin), near Tarapoto, Spruce
4030 (holotype, K!; isotype, BM!). Another sheet
of this number at GH is T. patens var. patens.
Lowland and montane forests, roadsides, ditch-
es, seepage areas, along trails, 400-2200 m, Lam-
bayeque, Cajamarca, Amazonas, San Martin, Lo-
reto, Huanuco, Lima, Junin, Ayacucho, Cuzco,
and Puno.
Antilles; Costa Rica to Panama; Colombia to
Venezuela and northwestern Argentina.
This taxon was formerly called T. patens var.
scabriuscula (Presl) A. R. Sm., but Ponce has ex-
amined the type of that variety and found it to be
referable to var. patens.
Lambayeque: 20 km from Olmos on road to Jaen,
Correll & Smith P793 (GH). Cajamarca: Ciudad Jaen,
Cerron [Coronado 43] (GH, uc, us). Prov. Santa Cruz,
Dist. Catache, upper Rio Zana valley ca. 1 km above
Monte Seco on road to El Chorro, Dillon et al. 4328 (F,
GH, uc). Amazonas: Prov. Bagua, along roadside between
La Peca and Bagua Chica, Barbour 4282 (MO, USM). San
Martin: Rio Huallaga Canon, below Rio Santo Domin-
go, Macbride 4260 (F, us). Loreto: Pumayacu, between
Balsapuerto and Moyobamba, Klug 3231 A (us). Huanu-
co: Cueva Grande Estacion near Pozuzo, Macbride 4787
(F). Prov. Huanuco, road Huanuco to Muna, Chulque,
Mexia 4099 (GH, MO, uc). Lima: Prov. Lima, near Rio
Santa Eulalia, Chosica, 40 km E of Lima, Tryon & Tryon
5341.5 (GH). Junin: Prov. Tarma, cerca a la Merced,
Cerrate 2840 (GH, USM). La Merced, Killip & Smith 23476
(NY, us), Kunkel 609 (F, GH). Ayacucho: Estrella, between
44
FIELDIANA: BOTANY
Huanta and Rio Apurimac, Killip & Smith 22650 (GH,
NY, us). Cuzco: La Convention, Rio Apurimac, between
San Martin and Hda. Luisiana, Davis el al. 1328 (F, GH).
Puno: Prov. Sandia, between Rio Azata and Colorado,
Nunez & Munoz 5244 (MO).
54. Thelypteris depilata A. R. Sm., sp. nov. Fig-
ure 4.
A speciebus ceteris subg. Cyclosori distinguenda axi-
bus et lamina omnino glabris, pinnis proximalibus ad
basin angustatis non auriculatis, 2-3 infimis paribus ve-
narum ex segmentis contiguis ad sinum conniventibus,
costarum paleis paucis vel nullis.
Stem long-creeping, scales brown, shining, lan-
ceolate, setose on margins and surfaces. Leaves
few, distant, 80-120 cm long. Lamina thick-char-
taceous, 1 -pinnate-pinnatifid, proximal pinnae not
or only slightly reduced. Petiole 30-60 cm x 4-8
mm, tan to stramineous and lacking scales above
the base. Rachis glabrous abaxially. Pinnae sessile,
16-22 x 1.5-2.2 cm, deeply pinnatifid 2-3 mm
from costae, those in lower half of lamina nar-
rowed at the base, not auriculate, segments 3 mm
wide. Aerophores absent. Buds lacking. Veins 1 3—
17 pairs per segment, lowermost 2-3 pairs from
adjacent segments connivent at sinus. Indument
on costae, veins, and laminar tissue abaxially lack-
ing, costal scales lacking or very sparse, adaxially
the lamina glabrous even on costae. Sori medial,
round, indusia glabrous, sporangial stalks without
glands.
TYPE— Peru, Dpto. Pasco, Prov. Oxapampa, ca.
20 km from Oxapampa towards La Merced, 3400
ft, Saunders 570 (holotype, F!; isotype, uc!).
Along streams, 1000-2150 m, Huanuco and
Pasco.
Known only from Peru.
This differs from other Peruvian Thelypteris
subg. Cyclosorus by the completely glabrous lam-
ina. In other characters, it is most similar to T.
clivalis, which is densely pubescent and has nu-
merous costal scales.
Huanuco: Mima, Macbride 3991 (F, 2 sheets).
55. Thelypteris clivalis A. R. Sm., nom. nov.
Dryopteris oligophylla var. aequatorialis C. Chr., Kon-
gel. Danske Vidensk. Selsk. Skr., Naturvidensk.
Afd., ser. 7, 10: 189. 1913. LECTOTYPE (chosen
by Morton, 1967): Ecuador, Banos, Rio Pastaza,
Spruce 5296 (p; isolectotypes, K!, NY!).
Thelypteris grandis var. aequatorialis (C. Chr.) A. R.
Sm., Univ. Calif. Publ. Bot. 59: 98. 1971.
Stem long-creeping, scales brown, dull to shin-
ing, lanceolate, setose on margins and surfaces.
Leaves few, distant, mostly 100-200 cm long.
Lamina thick-chartaceous to subcoriaceous, 1 -pin-
nate-pinnatifid, proximal pinnae not or only
slightly reduced. Petiole 50-100 cm x 5-10 cm,
tan to stramineous and lacking scales above the
base. Rachis densely pubescent abaxially, tri-
chomes 0.2-0.5 mm, also usually with a few scales.
Pinnae sessile or short-stalked less than 1 mm,
16-25 x 2-3 cm, deeply pinnatifid 2-3 mm from
costae, proximal ones narrowed at the base, not
auriculate, segments 3-4 mm wide, basiscopic
ones of proximal pinnae greatly reduced or some-
times wanting. Aerophores absent. Buds lacking.
Veins 14-21 pairs per segment, lowermost 1-2
(-3) pairs from adjacent segments connivent at the
sinus. Indument on costae, veins, and often lam-
inar tissue abaxially of usually dense, spreading,
crispate and acicular trichomes 0.2-0.5 mm, glands
lacking, costal scales numerous, castaneous, shin-
ing, 0.7-2 mm long, densely setulose, adaxially the
lamina glabrescent or with scattered crispate tri-
chomes 0.2-0.4 mm on and sometimes between
veins. Sori medial, round, indusia densely pubes-
cent, sporangial stalks without glands.
Along roads and trails, montane forests, (1000-)
1600-2500 m, Cajamarca, San Martin, Pasco, Ju-
nin, and Cuzco.
Colombia to Bolivia.
The varietal epithet aequatorialis cannot be
transferred to Thelypteris at species rank because
of the earlier T. aequatorialis (Copel.) Reed.
This species is most closely related to T. grandis
and differs primarily by the denser and longer tri-
chomes on the costae abaxially, more numerous
costal scales, generally narrower pinnae, more fal-
cate segments, and smaller fronds. It grows at high-
er elevation than T. grandis var. kunzeana. The
decision to recognize it at species rather than va-
rietal rank is the result of recognition of its ele-
vational separation and the fact that there seem
to be no intermediates between it and varieties of
T. grandis.
Cajamarca: Prov. Celendin, Llanguat, Mostacero 0955
(F, MO). San Martin: Prov. Rioja, Km 385-390 on Moyo-
bamba-Bagua road, Venceremos to Campamento Gar-
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
45
cia, D. Smith 5977 (F, USM). Huanuco: Between Huanuco
and Pampayacu, Kanehira 167 (us). Prov. Huanuco, road
Huanuco to Muna, Chulque, Mexia 4101 (GH, uc). Pas-
co: Quillasu, Soukup 3288 (GH, us). Junin: Between Tar-
ma and San Ramon, vicinity of Huacapistana, Croat
57649 (F, MO, USM). Prov. Tarma, valle del Rio Chan-
chamayo, Esposto 655 (GH, USM). Carpapata, Kunkel 631
(GH). Yucapata, Woytkowski 6748 (MO, TENN, us). Aya-
cucho: Ayna, between Huanta and Rio Apurimac, Killip
& Smith 22553 (NY). Cuzco: Prov. Urubamba, Machu
Picchu, Vargas 17448 (GH).
56. Thelypteris grandis A. R. Sm., Univ. Calif.
Publ. Bot. 59: 96. 1971, var. kunzeana (Hook-
er) A. R. Sm., Univ. Calif. Publ. Bot. 59: 99.
1971.
Aspidium abruptum Kunze, Linnaea 9: 93. 1834, not
Blume, 1828. TYPE: Peru, (Huanuco), Pampaya-
cu, Poeppig (holotype, LZ, destroyed; probable
isotypes, B!, K!, p!).
Nephrodium kunzeanum Hooker, Sp. fil. 4: 102. 1 862.
Nom. nov. for Aspidium abruptum Kunze, not
Blume.
Dryopteris oligophylla var. kunzeana (Hooker) C. Chr.,
Kongel. Danske Vidensk. Selsk. Skr., Naturvi-
densk. Afd., ser. 7, 10: 189. 1913.
Thelypteris invisa var. kunzeana (Hooker) Morton,
Contr. U.S. Natl. Herb. 38: 62. 1967.
Stem long-creeping, scales brown, dull, linear-
lanceolate, setose on margins and surfaces. Leaves
few, distant, mostly ( 1 00-) 1 50-250 cm long. Lam-
ina thick-chartaceous to subcoriaceous, 1 -pinnate-
pinnatifid, proximal pinnae not or only slightly
reduced, apex subabruptly reduced. Petiole 50-
100 cm x 5-10 mm, tan to stramineous and lack-
ing scales above the base. Rachis moderately to
densely pubescent abaxially, trichomes 0.1-0.2
mm. Pinnae sessile or the proximal short- to long-
stalked up to 20 mm, 20-30 x 2-4.5 cm, deeply
pinnatifid 3-6 mm from costae, proximal ones
narrowed at the base, not auriculate, segments 3-
6 mm wide, basiscopic ones of proximal pinnae
greatly reduced or wanting. Aerophores absent.
Buds lacking. Veins ( 1 2-) 1 5-22 pairs per segment,
lowermost 2-3 pairs from adjacent segments con-
nivent at the sinus. Indument on costae and veins
abaxially of moderately dense, spreading, acicular
trichomes 0. 1-0.2 mm, intervenal tissue glabrous,
glands lacking, costal scales scattered, castaneous,
shining, mostly 0.4-1 mm long, densely setulose,
adaxially the lamina glabrous except along costae.
Son supramedial to submarginal, round, indusia
moderately pubescent, trichomes 0.1 mm long,
sporangial stalks without glands.
Lowland and montane forests, 100-700(-1 500)
m, Amazonas, San Martin, Loreto, Huanuco, Ju-
nin, and Cuzco.
Ecuador to Bolivia; southern Brazil.
Variety grandis is restricted to the Antilles and
southern Florida, whereas var. pallescens (C. Chr.)
A. R. Sm. occurs in the Greater Antilles, Central
America, Venezuela and Colombia, Ecuador, and
perhaps Bolivia. Herein, I raise var. aequatorialis
to species rank (as T. clivalis).
Amazonas: Prov. Bagua, bank of Rio Maranon above
Cascadas de Mayasi, Wurdack 1951 (F, GH, NY, uc, us,
USM). San Martin: Prope Tarapoto, Spruce 4066 (K, p).
Loreto: Lower Rio Nanay, Li Williams 308 (F), 372 (F),
379 (F, us). Dist. Iquitos, bank of Rio Maranon, below
Rancho Indiana, Mexia 6462 (BH, F, GH, MO, NY, u, uc,
us). Pasco: Quillasu, Soukup 3292 (F, GH). Junin: La
Merced, Killip & Smith 23542 (NY, us). Colonia del Pere-
ne, Coronado 241 (GH, uc). Cuzco: Prov. La Convention,
Rio Apurimac below San Martin above Hda. Luisiana,
Davis et al. 1334 (F, GH, uc).
lid. Thelypteris subg. Steiropteris.
Thelypteris subg. Steiropteris (C. Chr.) Iwatsuki,
Mem. Coll. Sci. Kyoto, Ser. B. 31: 31. 1964.
Figure 5.
Dryopteris subg. Steiropteris C. Chr., Biol. Arb. til.
Bug. Warming 81. 1911. TYPE: Dryopteris del-
toidea (Sw.) C. Chr. = Thelypteris deltoidea (Sw.)
Proctor
Steiropteris (C. Chr.) Pic.-Ser., Webbia 28: 449. 1973.
Lamina 1 -pinnate-pinnatifid, or pinnatifid with
a few free pinnae in 2 extralimital species; prox-
imal pinnae the longest or nearly so (except the
type), distal pinnae gradually reduced and the lam-
ina with a confluent, pinnatifid apex, or sometimes
abruptly reduced with a subconform terminal pin-
na; aerophores often strongly developed at pinna
bases; buds lacking or rarely present. Veins from
adjacent segments meeting margin above sinus
(sect. Glaphyropteris), or connivent at sinus (sect.
Steiropteris) with a veinlike keel (pseudovein) ex-
tending from sinus toward costa, occasionally the
veins uniting below sinus or with the pseudovein.
Indument of acicular trichomes, these sometimes
septate. Sori round, indusiate or exindusiate; spo-
rangia glabrous, rarely setulose. Spores with few
prominent, connected, winglike ridges or many
shorter disconnected ones, x = 36.
46
FIELDIANA: BOTANY
FIG. 5. Subgenus Steiropteris. Thelypteris valdepilosa: a, stem with fertile (1.) and sterile (r.) leaves; b, juncture of
rachis and costa, abaxial side of fertile leaf, with aerophore; c, costa and sterile segments, adaxial side, (a, b from
Herrera 3249, Costa Rica, uc; b from Foster et al. 8985, uc.)
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
47
Subgenus Steiropteris comprises 2 1 Neotropical Reference
species, from the Antilles, southern Mexico to Bo-
livia, and southern Brazil (Smith, 1980). In general SMITH, A. R. 1980. Taxonomy of Thelypteris
aspect, several species most closely resemble cer- subgenus Steiropteris, including Glaphyropteris
tain species of subg. Goniopteris, and the two are (Pteridophyta). Univ. Calif. Publ. Bot. 76: 1-38.
perhaps more closely related to each other than to
any other subgenus.
Key to Species of subg. Steiropteris
a. Sinus keel (a cartilaginous, raised, often minutely pubescent pseudovein) lacking; veins (17-)20-45
pairs per segment, meeting margin above the sinus; indusia absent; laminar tissue between the veins
often with sessile glands (sect. Glaphyropteris) b
b. Costae abaxially short-pubescent (0.1-0.2 mm) or with short and long nonseptate trichomes
intermixed; costules and veins adaxially glabrous 57. T. decussate
b. Costae abaxially with dense, soft, septate trichomes 1-2 mm long; costules and veins adaxially
with relatively dense, strigose trichomes up to 2 mm 58. T. comosa
a. Sinus keel prominent; veins fewer than 22 pairs per segment, connivent at the sinus or sometimes
uniting below sinus; indusia present or absent; laminar tissue between veins glandless (sect. Steirop-
teris) c
c. Sori exindusiate d
d. Scalelike aerophores at pinna bases; pinnae incised ca. y3-V2 59. T. glandulosa
d. Scalelike aerophores absent at pinna bases (small darkened swellings may be present); pinnae
incised 2/3-% 60. T. pennellii
c. Sori indusiate e
e. Orangish glands present on receptacle; fronds subdimorphic, the fertile long-petioled and with
contracted pinnae 61. T. valdepilosa
e. Glands absent on receptacle; fronds monomorphic or nearly so f
f. Aerophores present at pinna bases, peglike 62. T. leprieurii
f. Aerophores absent at pinna bases g
g. Indusia minute, spatulate, easily overlooked among mature sporangia; segments 5-8 mm
wide 60. T. pennellii
g. Indusia conspicuous; segments 3-5 mm wide 63. T. gardneriana
57. Thelypteris decussata (L.) Proctor, Bull. Inst.
Jamaica, Sci. Ser. 5: 59. 1953.
Stem massive, suberect, scales tan (whitish when
young), dull, ovate, glabrous, appressed. Leaves
(1 00-) 150-350 cm long, croziers mucilaginous.
Lamina chartaceous, 1 -pinnate-pinnatifid, proxi-
mal pinnae not reduced. Petiole ca. 50-1 50 cm x
5-10 mm, brownish to purplish brown, lacking
scales above the base. Rachis pubescent or gla-
brescent abaxially, trichomes mostly 0. 1-0.25 mm.
Pinnae sessile, 18-25(-45) x 1.8-3(-4.5) cm,
deeply pinnatifid within 1 mm of costae, proximal
ones narrowed at base, not auriculate, segments
2-5 mm wide, margins with numerous flexuous
septate trichomes. Aerophores scalelike at pinna
bases, 5-10 mm long, with shorter, threadlike ones
1-3 mm at bases of costules. Buds lacking. Veins
(17-)25-35(-45) pairs per segment, lowermost pair
from adjacent segments meeting margin just above
sinus, sinus keel lacking. Indument on costae abax-
ially of moderately dense to dense, spreading, non-
septate trichomes mostly less than 0. 1 mm, lam-
inar tissue abaxially lacking trichomes but often
with reddish to orangish sessile resinous glands,
costal scales lacking, adaxially with numerous stout
trichomes to ca. 1 mm on costae, a few on costules.
Sori inframedial to medial, round, exindusiate,
often with receptacular glands.
Montane rain forests, 500-1300 m, Pasco, Ju-
nin, Ayacucho, Cuzco, and Madre de Dios.
Guatemala to Panama; Antilles; Colombia to
Guianas and Bolivia; southern Brazil.
Lechler 2356 (G, K, L), from San Gaban, Puno,
was previously (Smith, 1980) and tentatively
named var. velutina (Sodiro) A. R. Sm., but no
additional material has come to my attention, and
I am now uncertain of the varietal identity of this
specimen.
48
FIELDIANA: BOTANY
Key to Varieties
a. Lamina with red to orange, sessile glands abaxially; pinnae greater than 2 cm wide
57a. var. decussata
a. Lamina eglandular abaxially; pinnae less than 2 cm wide 57b. var. mapiriensis
57a. Thelypteris decussata var. decussata.
Polypodium decussatum L., Sp. pi. 2: 1093. 1753.
TYPE: Petiver's Pterigraphia Amer. 61, t. 2,f. 5,
1712, which is a redrawing of Plumier's Tract.
fil. amer. /. 24, 1705, illustrating a plant said to
have come from Martinique, Morne de la Cale-
basse.
Glaphyropteris decussata (L.) Fee, Crypt, vase. Bresil
2: 40. 1873.
Dryopteris decussata (L.) Urban, Symb. antill. 4: 19.
1903.
Montane rain forests, 500-1 100 m, Pasco and
Cuzco.
Antilles; Guatemala to Panama; Colombia to
French Guiana and Peru.
Pasco: Pichis Trail, San Nicolas, (as Junin), Killip &
Smith 26030 (GH, NY, us). Cuzco: Prov. Quispicanchi,
entre Inambari y 15 Mil, Vargas 16503 (GH).
57b. Thelypteris decussata var. mapiriensis (Ro-
senst.) A. R. Sm., Univ. Calif. Publ. Hot. 76:
16. 1980.
Dryopteris mapiriensis Rosenst., Repert. Spec. Nov.
Regni Veg. 6: 313. 1909. TYPE: Bolivia, La Paz,
San Antonio near Mapiri, Buchtien 1131 (holo-
type, not found at s; isotypes, BM!, us!).
Thelypteris mapiriensis (Rosenst.) Alston, J. Wash.
Acad. Sci. 48: 234. 1958.
Montane rain forests, 700-1 300 m, Junin, Aya-
cucho, and Madre de Dios.
Peru; Bolivia.
Junin: Schunke Hda., above San Ramon, Killip & Smith
24650 (NY). Ayacucho: Prov. La Mar, San Jose to Huan-
huachayo, between Tambo San Miguel, Ayna, and Hda.
Luisiana, Dudley 1 1 945 (GH). Madre de Dios: Prov. Manu,
Cerro de Pantiacolla, Rio Palotoa 10-15 km NNW of
Shintuya, R. Foster et al. 10906 (F, USM).
58. Thelypteris comosa (Morton) Morton, Amer.
FernJ. 51: 38. 1961.
Dryopteris comosa Morton, J. Wash. Acad. Sci. 28:
528. 1938. TYPE: Peru, Pasco, Dos de Mayo,
Pichis Trail (as Junin), Killip & Smith 25872 (ho-
lotype, us!; isotype, NY!).
Stem massive, suberect, scales tan (whitish when
young), dull, ovate, glabrous, appressed. Leaves
ca. 110 cm long. Lamina chartaceous, 1 -pinnate-
pinnatifid, proximal pinnae not reduced or 1-2
pairs slightly reduced. Petiole ca. 45 cm x 8 mm,
brownish, lacking scales above the base. Rachis
pubescent abaxially, trichomes mostly 0. 1-0.2 mm
and a few up to 2 mm. Pinnae sessile, 15-25 x
2.5-3.5 cm, deeply pinnatifid within 1 mm of cos-
tae, proximal ones narrowed at base, not auricu-
late, segments 3-4 mm wide. Aerophores scalelike
at pinna bases, to 7 mm long, with shorter thread-
like ones at bases of costules. Buds lacking. Veins
25-30 pairs per segment, lowermost pair from ad-
jacent segments meeting margin just above sinus,
sinus keel lacking. Indument on costae and veins
abaxially of moderately dense to dense, spreading,
sometimes septate trichomes mostly 1-2 mm,
laminar tissue lacking glands or very sparsely glan-
dular, with a few trichomes, costal scales lacking,
adaxially with numerous stout trichomes to 2 mm
on costae and costules. Sori medial, round, exin-
dusiate, lacking receptacular glands.
Montane rain forests, 1700-2100 m, Pasco and
Cuzco.
Known only from Peru.
Cuzco: Prov. La Convention, ca. 1 7 km walking dis-
tance NE from Hda. Luisiana and Apurimac river, above
Camp 3, Dudley 11271 (GH).
59. Thelypteris glandulosa (Desv.) Proctor var.
brachyodus (Kunze) A. R. Sm., Phytologia 34:
233. 1976.
Polypodium brachyodus Kunze, Linnaea 9: 48. 1834.
TYPE: Peru, (Huanuco), "in sylv. umbros. Cu-
chero" (specimen), Pampayacu, Poeppig (holo-
type, LZ, destroyed; isotype, w!).
Dryopteris brachyodus (Kunze) Kuntze, Rev. gen. pi.
2: 812. 1891.
Dryopteris glandulosa (Desv.) C. Chr. var. brachyodus
(Kunze) C. Chr., Kongel. Danske Vidensk. Selsk.
Skr., Naturvidensk, Afd., ser. 7, 10: 172. 1913.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
49
Thelypteris brachyodus (Kunze) Ching, Bull. Fan Mem.
Inst. Biol., Hot. 6: 286. 1936.
Stem short-creeping, scales few, brown, dull,
ovate, sparsely pubescent to glabrous. Leaves few,
mostly 75-150 cm long. Lamina thick-charta-
ceous, 1 -pinnate-pinnatifid, proximal pinnae not
reduced. Petiole 30-75 cm x 4-8 mm, brownish,
lacking scales above the base. Rachis usually pu-
bescent abaxially, trichomes mostly 0.1-0.3 mm
long. Pinnae sessile to short-stalked 1-4 mm,
mostly 10-25 x 2. 5-4 cm, shallowly pinnatifid ca.
!/3-!/2 their width, proximal ones slightly narrowed
at base, not auriculate, segments 5-8 mm wide.
Aerophores commonly scalelike at pinna bases, 1-
2 mm long. Buds lacking. Veins mostly 14-22 pairs
per segment, lower 3-5 pairs from adjacent seg-
ments usually running alongside a raised minutely
pubescent (trichomes 0.1 mm) sinus keel (false
vein) and connivent with each other or with the
sinus keel. Indument on costae and veins abaxially
of dense, spreading, acicular trichomes mostly ca.
0. 1 mm, laminar tissue glabrous and lacking glands,
costal scales tan, appressed, linear-lanceolate, lam-
ina adaxially glabrous except for trichomes ca. 0. 1
mm along costae. Sori inframedial to medial, round
to often oblong, exindusiate, lacking receptacular
glands.
Lowland rain forests, 100-700 m, Amazonas,
Loreto, and Huanuco.
Southern Mexico (Chiapas); Belize and Guate-
mala to Costa Rica; Colombia; Ecuador; Peru.
Three varieties are known, with var. glandulosa
in the Lesser Antilles and the Guianas to Colombia
and var. longipilosa A. R. Sm. from Costa Rica
and Panama to Ecuador, primarily on the Pacific
slope.
The collections by Sagastegui and Moran are
similar to typical material in venation and lamina
dissection but differ in having glabrous costae
abaxially, strictly opposite pinnae, son more elon-
gate along the veins, and apparently in lacking
aerophores at the pinna bases. These two collec-
tions may represent an undescribed species.
Amazonas: Prov. Bagua, Montenegro-Chiriaco, Sa-
gastegui 5925 (GH). Loreto: Prov. Maynas, Yanamono
tourist camp, 50 mi NE of Iquitos, Vasquez & Jaramillo
516 (F, MO). Balsapuerto, Killip & Smith 28478 (NY, us).
Prov. Maynas, ca. 50 miles downriver on the Amazonas
at Jensen's Explorama lodge, Moran 3727 (MO, uc).
60. Thelypteris pennellii A. R. Sm., Univ. Calif.
Publ. Hot. 76: 28. 1980. TYPE: Colombia,
Tolima, "La Virginia," Libano, Pennell 3265
(holotype, NY!; isotype, us!).
Stem not known, probably short-creeping.
Leaves to 1 40 cm long. Lamina thick-chartaceous,
1 -pinnate-pinnatifid, proximal pinnae not re-
duced, apex pinnatifid, confluent or free. Petiole
to ca. 80 cm x 8 mm, stramineous to tan, lacking
scales above the base. Rachis sparsely pubescent
abaxially, trichomes 0. 1 mm long. Pinnae oppo-
site to subopposite throughout, sessile to short-
stalked 1 mm, 15-20 x 2.5-3.5 cm, pinnatifid ca.
2/3-% their width (to 4-6 mm from costae), prox-
imal ones not narrowed at base, not auriculate,
segments 5-8 mm wide. Aerophores absent or re-
duced to a small, darkened swelling. Buds lacking.
Veins up to 18 pairs per segment, lower 2-3 pairs
from adjacent segments running alongside a mi-
nutely pubescent sinus keel (that extends from si-
nus nearly to costa). Indument on costae abaxially
of scattered, acicular trichomes 0. 1 mm, veins and
laminar tissue glabrous, costal scales tan, ap-
pressed, linear, adaxially the lamina glabrous ex-
cept for trichomes ca. 0. 1 mm along costae. Sori
inframedial to subcostular, round, indusia spatu-
late, sparingly ciliate to glabrous, evanescent or
obscured by mature sporangia, lacking recepta-
cular glands.
Montane, rain forest, Huanuco.
Colombia, Peru.
Huanuco: Pampayacu, Kanehira 132 (GH, us), 133 (GH).
61. Thelypteris valdepilosa (Baker) Reed, Phy-
tologia 17: 323. 1968. Figure 5.
Nephrodium valdepilosum Baker, J. Bot. 19: 204. 1881.
LECTOTYPE (chosen by Smith, 1980): Colom-
bia, Dept. Antioquia, Kalbreyer 1871 (K!; isolec-
totype, K!).
Dryopteris valdepilosa (Baker) C. Chr., Index fil. 299.
1905.
Steiropteris valdepilosa (Baker) Pic.-Ser., Webbia 28:
451. 1973.
Stem suberect, scales brown, shining, lanceolate,
densely setulose on surface and margins. Leaves
several, subdimorphic, the fertile with longer pet-
ioles and narrower pinnae than the sterile, 45-75
(-120) cm long. Lamina chartaceous, 1 -pinnate-
pinnatifid, proximal pinnae not reduced or low-
ermost pair slightly reduced. Petiole 1 5-20 cm x
50
FIELDIANA: BOTANY
3 mm, brownish, lacking scales above the base,
densely pubescent with short trichomes 0.1-0.2
mm, the sterile intermixed with long, silky, septate
trichomes 2-4 mm. Rachis densely pubescent
abaxially, trichomes as on petioles. Pinnae sessile,
4-6(-12) x l-1.7(-3) cm, deeply pinnatifid 1-2
mm from costae, proximal ones slightly narrowed
at base, not auriculate, segments 2-5 mm wide,
the fertile narrower. Aerophores peglike, to 1 mm
at pinna bases. Buds lacking. Veins 7-10 pairs per
segment, lower pair from adjacent segments run-
ning to or just above sinus, minutely pubescent
keel running from sinus nearly to costa. Indument
on costae and veins (sterile lamina) abaxially of
dense, acicular trichomes mostly 1-3 mm long,
0.1-0.3 mm on fertile lamina, laminar tissue of
both fronds with adpressed trichomes 0.2-0.3 mm,
also with inconspicuous glands mainly along veins,
costal scales few, linear, appressed, lamina adax-
ially with adpressed trichomes 0.2-0.3 mm on and
between veins, the sterile also with a few long tri-
chomes 1-3 mm on veins. Sori medial, round,
indusium short-setose, trichomes 0.2-0.3 mm, re-
ceptacle with stalked globose orangish glands.
Montane rain forest, 1000 m, Pasco.
Costa Rica and Panama; Colombia to Peru.
The recent finding of this very distinct and rare,
but apparently widespread, species emphasizes the
need for much additional collecting in Peru and
elsewhere in Latin America. The fertile and sterile
fronds show remarkable dimorphism in pubes-
cence.
Pasco: Oxapampa, Pichis Valley, San Matias Ridge,
10-12 km SW of Puerto Bermudez, above Santa Rosa
Key to Varieties
de Chivis, trail to Loma Linda, R. Foster et al. 8985
(MO, uc, USM).
62. Thelypteris leprieurii (Hooker) R. Tryon,
Rhodora 69: 6. 1967.
Stem creeping to suberect, scales brown, shin-
ing, lanceolate, setulose on surface and margins.
Leaves several, 60-140 cm long. Lamina charta-
ceous, 1 -pinnate-pinnatifid, proximal pinnae not
reduced, apex confluent and pinnatifid. Petiole 30-
70 cm x 3-6 mm, tan to brownish, lacking scales
above the base, pubescent with trichomes 0. 1-1.5
mm. Rachis densely pubescent abaxially, tri-
chomes uniformly short, 0.1-0.2 mm, or mixed
with long trichomes up to 1.5 mm, or trichomes
mostly long, 1-2 mm, septate. Pinnae sessile, 8-
17 x 1.5-3 cm, deeply pinnatifid 0.5-2 mm from
costae, proximal commonly narrowed at base (seg-
ments ca. half the length of the longest), not au-
riculate, segments 3-5 mm wide. Aerophores peg-
like, 0.5-2 mm at pinna bases. Buds lacking. Veins
10-16 pairs per segment, lowermost pair from ad-
jacent segments running to or just above sinus,
minutely pubescent keel running from sinus nearly
to costa. Indument on costae and veins (sterile
lamina) abaxially of moderately dense to dense,
acicular trichomes mostly 0.2-1 .5 mm long, longer
trichomes sometimes septate, laminar tissue usu-
ally glabrous, glands lacking, costal scales absent,
lamina adaxially glabrous or with stout trichomes
on veins. Sori inframedial to medial, round, in-
dusium glabrous, short-setose (trichomes 0.2-0.3
mm), or glandular, receptacle lacking glands.
a. Indusia glandular, without trichomes 62b. var. glandifera
a. Indusia eglandular to glandular, with at least a few trichomes b
b. Trichomes on petiole, rachis, and costae abaxially moderately dense, not septate, mostly 0.2-1
mm long 62a. var. leprieurii
b. Trichomes on petiole, rachis, and costae abaxially dense, obviously septate, mostly 1-2 mm long
. . 62c. var. incana
62a. Thelypteris leprieurii var. leprieurii.
Nephrodium leprieurii Hooker, Sp. fil. 4: 106. 1862.
TYPE: French Guiana, Leprieur (holotype, K!;
frag., BM!; isotypes, B!, us!).
Dryopteris leprieurii (Hooker) Kuntze, Rev. gen. pi.
2: 813. 1891.
Steiropteris leprieurii (Hooker) Pic.-Ser., Webbia 28:
451. 1973.
Montane rain forests, ravines, 700-1200 m, San
Martin, Huanuco, and Junin.
Trinidad; Colombia to Bolivia and French Gui-
ana, southern Brazil.
San Martin: Mt. Campana, prope Tarapoto, Spruce
4660 (K). Huanuco: Tingo Maria (as San Martin), Allard
22559 (us). Prov. Huanuco, Tingo Maria, Tryon & Tryon
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
51
5279 (OH, USM). Junin: Chanchamayo valley, Schunke
139 (F), 665 (F).
62b. Thelypteris leprieurii var. glandifera A. R.
Sm., Univ. Calif. Publ. Bot. 76: 25. 1980.
TYPE: Bolivia, La Paz, Prov. Larecaja, Co-
pacabana, Krukoff 11176 (holotype, us!; iso-
types, G!, GH!, NY!, s!).
Lowland and montane rain forests, 340-1700
m, Pasco, Junin, and Puno.
Peru; Bolivia; central Brazil.
Pasco: Prov. Oxapampa, Quebrada Castilla, near Villa
America, Leon & Young 1037 (uc, USM). Prov. Oxa-
pampa, Gran Pajonal, 2-3 km N of Chequitavo, D. Smith
5097 (MO, uc, USM). Pichis Trail, Yapas (as Junin), Killip
& Smith 25497 (NY). Junin: Schunke Hacienda, above
San Ramon, Killip & Smith 24566 (NY). Puno: San Ga-
ban (as St. Gavan), Lechler 2418 (BM).
62c. Thelypteris leprieurii var. int-an a (Christ) A.
R. Sm., Univ. Calif. Publ. Bot. 76: 26. 1980.
Aspidium incanum Christ, Hedwigia 44: 367. 1905.
TYPE: Brazil, Amazonas, Puritisal, Jurua Miry,
Rio Jurua, Ule 5763 (holotype, p; isotypes, B!, G!,
K!; frag., BM!).
Dryopteris incana (Christ) C. Chr., Index fil. 272. 1 905.
Steiropteris incana (Christ) Pic.-Ser., Webbia 28: 45 1 .
1973.
Lowland rain forest, Loreto.
Ecuador; Peru; Amazonian Brazil.
Loreto: Near Pobre Alegre, 40-45 km upriver from
Atalaia do Norte, Brazil, along Rio Javari, Breedlove
35515 (DS, MO).
63. Thelypteris gardneriana (Baker) Reed, Phy-
tologia 17: 278. 1968.
Nephrodium gardnerianum Baker in C. Martius, Fl.
bras. 1(2): 474. 1870. TYPE: Brazil, Rio de Ja-
neiro, Serra do Orgaos, Gardner 190 (holotype,
K.!; isotypes, BM!, P!).
Dryopteris densiloba C. Chr., Index fil. 261. 1905,
nom. nov. for Nephrodium gardnerianum and with
the same type.
Dryopteris supralineata Rosenst., Repert. Spec. Nov.
Regni Veg. 8: 277. 1910. TYPE: Brazil, Sao Pau-
lo, Serra do Mar, Wacket 212 (holotype, not found
in s; isotypes, BM!, NY!, uc!; probable isotypes,
Rosenst. exs. 437, B!, BM!, L!, NY!, us!).
Steiropteris gardneriana (Baker) Pic.-Ser., Webbia 28:
451. 1973.
Stem short-creeping, scales brown, shining, lan-
ceolate, setulose on surface and margins. Leaves
several, mostly 50-100 cm long. Lamina charta-
ceous, 1 -pinnate-pinnatifid, proximal pinnae not
reduced or lowermost pair slightly reduced, apex
confluent and pinnatifid. Petiole 30-60 cm x 2-
3 mm, tan to brownish, lacking scales above the
base, pubescent with trichomes 0.1-0.2 mm. Ra-
chis moderately pubescent abaxially, trichomes
0.1-0.3 mm. Pinnae sessile, 5-12 x 1.6-2.5 cm,
deeply pinnatifid 1.5-2.5 mm from costae, prox-
imal pinnae commonly narrowed at base (seg-
ments ca. half the length of the longest), not au-
riculate, segments 3-5 mm wide. Aerophores
absent. Buds lacking. Veins 6-12 pairs per seg-
ment, lowermost pair from adjacent segments run-
ning to or just above sinus, minutely pubescent
keel running from sinus nearly to costa. Indument
on costae and veins abaxially of scattered, acicular
trichomes mostly 0. 1-0.3 mm long, laminar tissue
glabrous, glands lacking, costal scales absent, lam-
ina adaxially glabrous except for trichomes 0.1-
0.3 mm along costae. Sori medial, round, indu-
sium short-setose (trichomes 0.1-0.2 mm), recep-
tacle lacking glands.
Rocky slopes, 2400-2800 m, Cuzco.
Colombia; Venezuela; Ecuador; Galapagos;
Peru; Bolivia; southern Brazil. Rare everywhere
except in southern Brazil.
Cuzco: Prov. Paucartambo, Pillahuata, B. Aires, Var-
gas 16740 (GH).
He. Thelypteris subg. Goniopteris.
Thelypteris subg. Goniopteris (Presl) Duek, Adan-
sonia, II. 11: 720. 1971. Figure 6.
Goniopteris Presl, Tent, pterid. 181. 1836. TYPE:
Goniopteris crenata (Sw.) Presl [= Thelypteris poi-
teana (Bory) Proctor].
Lamina simple to usually 1 -pinnate or 1 -pin-
nate-pinnatifid; proximal pinnae the longest or
nearly so, not reduced, rarely a few slightly re-
duced, distal pinnae gradually shortened and the
lamina with a confluent, pinnatifid apex, or the
lamina with a conform apical pinna; aerophores
lacking or rarely present (T. lugubriformis); buds
often present in axil(s) of distal pinnae, rarely in
axil(s) of proximal pinnae (T. poiteand). Veins
commonly connivent at the sinus or uniting below
sinus, or forming a series of aeroles each with a
single excurrent vein (veins meniscioid). Indument
52
FIELDIANA: BOTANY
1mm
1mm
FIG. 6. Subgenus Goniopteris. Thelypteris biolleyi: a, portion of stem and petiole; b, portion of lamina; c, pinnules,
abaxial side; d, indument of segment base; e, indument at edge of rachis. (a, c, d, e from Mexia 6184, uc; b from
Fisher 35395, Mexico, F.)
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
53
of acicular and usually furcate, stellate, or anchor-
shaped trichomes (lacking in a few spp.), these
most evident on stem apex scales and in adaxial
grooves, but also often on axes and lamina abax-
ially, rarely branched trichomes lacking. Sori
round, indusia round-reniform, up to ca. 1 mm in
diameter or reduced to a small fragment, or sori
exindusiate; sporangia glabrous, setose, or with
stellate or furcate trichomes from capsules or stalk.
Spores with few to many prominent connected or
disconnected wings, x = 36.
Subgenus Goniopteris comprises 80-100 Neo-
tropical species, from Florida, the Antilles, and
southern Mexico to northern Argentina and Par-
aguay. Species are most numerous in lowland and
montane rain forests and are virtually absent at
elevations above 2000 m. A few species are semi-
weedy in shaded plantations (coffee, cacao). Many
Peruvian species produce small buds in the axils
of distal pinnae. These buds often remain unde-
veloped on mature fronds but perhaps are impor-
tant in establishment of new plants when the fronds
are senescent and decaying.
Several Peruvian species, i.e., T. macrotis, T.
semihastata, and T. clypeata, completely lack the
stellate or furcate trichomes which are the most
constant characteristic of the subgenus. The last
species has venation very much like subg. Menis-
cium, and could be evolutionarily transitional to
that subgenus; however, T. clypeata also has large
indusia, which are unknown in Meniscium. The
loss of branched trichomes in T. macrotis and T.
semihastata seems likely to have occurred inde-
pendently from T. clypeata. The most primitive
element in the subgenus may be T. lugubriformis,
which has a lamina aspect much like subg. Stei-
ropteris; it is also the only species in the subgenus
with well-developed aerophores.
References
See CHRISTENSEN (1913) and SMITH (1983) under
the family references.
Key to Species of subg. Goniopteris
a. Lamina gradually narrowed distally into a pinnatifid apex b
b. Lamina pinnatifid, or 1 -pinnate-pinnatifid with a few pinnae proximally c
c. Pinnae entire or subentire, not auricled; lamina glabrous between veins adaxially
64. T. pinnatitida
c. Pinnae lobed, at least the proximal ones, also auricled at acroscopic base; lamina with adpressed
trichomes between the veins adaxially 65. T. jamesonii
b. Lamina 1 -pinnate to just short of the apex d
d. Lamina between veins on both sides with appressed, sessile stellate trichomes bearing 3-5
arms; acicular trichomes lacking on axes abaxially; sori exindusiate 76. T. biolleyi
d. Lamina between veins glabrous or with acicular trichomes, or trichomes furcate or stellate,
arms not appressed; acicular trichomes present on axes abaxially; sori indusiate or exindusiate
e
e. Trichomes on axes and lamina abaxially anchor-shaped, i.e., with two retrorse hooks at
tip; sori exindusiate; veins uniting at an obtuse angle below sinus with an excurrent vein
to sinus 77. T. ancyriothrix
e. Trichomes on axes and lamina abaxially acicular, furcate, or stellate; sori exindusiate or
indusiate, indusium usually at least 0.2 mm in diameter; veins connivent at sinus f
f. Stellate trichomes absent from all parts of plant; pinnae auriculate at the acroscopic base
g
g. Buds present in axil(s) of distal pinnae; lamina deltate; proximal pinnae not reduced
66. T. macrotis
g. Buds absent; lamina narrowly lanceate; several pairs of proximal pinnae reduced . .
67. T. semihastata
f. Stellate and/or furcate trichomes present on axes and on rhizome apex scales; pinnae
nonauriculate (except sometimes T. jamesonii) h
h. Pinnae shallowly lobed to V3(-V2) their width, lobes often broader than long; pinnae
slightly auriculate at the acroscopic base; buds lacking 65. T. jamesonii
h. Pinnae lobed at least '/3 or more of their width, not auriculate at the acroscopic base;
bud(s) commonly present in axils of distal pinnae i
54
FIELDIANA: BOTANY
i. Adaxial surface of lamina between veins with adpressed trichomes; aerophores
peglike at pinna bases 69. T. lugubriformis
i. Adaxial surface of lamina between veins glabrous; aerophores lacking j
j. Rachis and costae abaxially densely stellate-pulverulent, lacking acicular tri-
chomes 70. T. eggersii
j. Rachis and costae abaxially with stellate, furcate, and acicular trichomes in-
termixed, or trichomes mostly acicular k
k. Sori exindusiate; trichomes on both sides of rachis deep red; buds lacking
71. T. erythrothrix
k. Sori indusiate; trichomes of rachis hyaline or mixed with reddish ones; buds
commonly present in axils of distal pinnae 1
1. Distal pinnae subabruptly reduced, lamina with a hastate, pinnatifid apex;
costal trichomes abaxially mostly 0.2-1.5 mm long, rather dense, many
forked or stellate 72. T. biformata
1. Distal pinnae gradually reduced; costal trichomes mostly 0.1-0.2(-0.3)
mm long, sparse to moderately dense, mostly simple . . 68. T. abrupta
a. Lamina with a terminal pinna that resembles that lateral ones m
m. Pinnae entire to very shallowly crenate, 3-5 pairs; areoles 1 2-1 7-seriate between costa and margin;
indusia large, vaulted, 1-1.5 mm in diameter 82. T. clypeata
m. Pinnae crenate or more divided, usually more than 4 pairs; areoles less than 8-seriate, or lacking
with veins connivent at the sinus; indusia commonly much less than 1 mm in diameter, or
lacking, not vaulted n
n. Lamina between veins on both sides with sessile, stellate trichomes with 3-5 appressed arms;
trichomes of abaxial costae all stellate, none acicular o
o. Pinnae shallowly incised less than V4 the distance to costae; veins uniting in 3-6 pairs
between costa and margin 78. T. schunkei
o. Pinnae incised ca. V3-l/2 the distance to costae; veins uniting in 1-3 pairs between costa
and margin 79. T. pennata
n. Lamina between veins on both sides glabrous or with acicular or stalked-furcate trichomes,
lacking appressed stellate trichomes; trichomes of abaxial costae a mixture of acicular, furcate,
and stellate trichomes, at least some stalked p
p. Pinnae crenate or very shallowly lobed less than !/5 the distance to costae; buds in axils of
proximal pinnae; trichomes of costae and rachis abaxially acicular; sori exindusiate ....
81. T. poiteana
p. Pinnae shallowly to deeply lobed V3 or more the distance to costae; buds lacking or in axils
of distal pinnae; at least some trichomes of costae and rachis abaxially furcate or stellate;
sori indusiate or exindusiate q
q. Pinnae incised !4-2/5 the distance to costae r
r. Indusia present, small, often hidden in mature sori; costae lacking scales abaxially
74. T. juruensis
r. Indusia absent; costae abaxially with tan, narrow, appressed scales
75. T. tryonorum
q. Pinnae incised V2 or more the distance to costae s
s. Proximal pair of veins from adjacent segments uniting below the sinus at an obtuse
angle, with an excurrent vein to sinus; sori exindusiate; buds lacking
80. T. tetragona
s. Proximal pair of veins running to sinus, sometimes connivent just below sinus at
an acute angle; sori indusiate or seemingly exindusiate; buds commonly present in
axils of distal pinnae t
t. Proximal pinnae cuneate at base; costal trichomes abaxially mostly acicular, a
few furcate 73. T. tristis
t. Proximal pinnae truncate at base; costal trichomes abaxially a mixture of stellate,
furcate, and acicular trichomes u
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III. 55
u. Lamina between veins adaxially with numerous, appressed trichomes; peglike
aerophores present at base of largest pinnae abaxially . . 69. T. lugubriformis
u. Lamina between veins glabrous adaxially; aerophores absent
. 72. T. biformata
64. Thelypteris pinnatifida A. R. Sm., Fl. Ecuador
18: 110. 1983. TYPE: Ecuador, Prov. Pas-
taza, Rio Bobonaza, Spruce 5293 (holotype,
NY!; isotypes, B!, K!, P!).
Stem short-creeping to ascending or suberect,
caudex ca. 1 cm or less in diameter, scales light
brown, dull, ovate-lanceolate, glabrous or mi-
nutely and sparingly setulose with trichomes 0.1
mm. Leaves several, clustered, mostly 12-30 cm
long. Petiole 3-12 cm x 0.6-1.0 mm, tan, gla-
brescent or with long acicular trichomes and a few
short stellate or furcate ones. Lamina herbaceous,
not verrucose, pinnatifid or with l-2(-3) pairs of
subentire free pinnae proximally. Buds lacking.
Rachis pubescent abaxially, trichomes hyaline or
red-tipped, stiff, acicular, 0.5-1 mm, furcate or
stellate trichomes absent or relatively few adaxi-
ally, 0. 1 mm long. Segments (pinnae in more di-
vided leaves) mostly 0.8-1.8 cm x 4-7 mm, en-
tire, rounded or truncate at tip, lamina incised
nearly or quite to rachis at base of lamina, ca. V2
or less in distal half, proximal free pinnae, if pre-
sent, often subauriculate at acroscopic base. Veins
8-13 pairs per segment, simple or l-(2)-forked,
proximal pair (or l'/2 pairs) from adjacent seg-
ments connivent at sinus. Indument abaxially on
costae and veins of moderately dense to dense,
spreading, often red-tipped, acicular trichomes 0.3-
1 mm, furcate and stellate trichomes lacking, lam-
inar tissue glabrous, veins and laminar tissue gla-
brous adaxially. Sori medial, indusia reddish
brown, setose along margin; sporangia glabrous.
Along stream banks, lowland rain forests, 200-
400 m, infrequent in Peru: Loreto, Ucayali, and
Madre de Dios.
Ecuador to Bolivia.
Loreto: Above Pongo de Manseriche, mouth of Rio
Santiago, Mexia 6293 (BH, F, o, GH, MO, NY, uc). Ucayali:
Prov. Coronel Portillo, Pucallpa-Lima Hwy, Km 85,
McDaniel 13944 (GH, MO, USM). Prov. Coronel Portillo,
Bosque von Humboldt, along Quebrada Tahuahillo,
Young 949 (F). Madre de Dios: Prov. Manu, Manu Park,
Cocha Cashu uplands, Nunez 6119 (MO).
65. Thelypteris jamesonii (Hooker) R. Try on,
Rhodora69: 6. 1967.
Nephrodium jamesonii Hooker, Sp. fil. 4: 66. 1 862.
LECTOTYPE (chosen by Christensen, 1913, p.
227): Ecuador, Rio Napo, Jameson 761 (K!).
Dryopteris jamesonii (Hooker) C. Chr., Kongel. Danske
Vidensk. Selsk. Skr., Naturvidensk. Afd., ser. 7,
10:227. 1913.
Dryopteris warmingii C. Chr., Kongel. Danske Vi-
densk. Selsk. Skr., Naturvidensk. Afd., ser. 7, 10:
227. 1913. TYPE: Brazil, Minas Gerais, Lagoa
Santa, Warming 1864 (holotype, c).
Dryopteris macrotis var. nephrodioides Rosenst., Re-
pert. Spec. Nov. Regni Veg. 7: 298. 1909. TYPE:
Peru, Mt. Campana, Spruce 4658 (isotype, P).
Stem suberect to erect, caudex up to 2 cm in
diameter, scales light brown, dull, ovate-lanceo-
late, with simple and furcate trichomes 0.1-0.2
mm. Leaves several, clustered, (15-)20-60(-100)
cm long, sometimes slightly dimorphic with fertile
longer-petioled and with slightly contracted pin-
nae. Petiole 5-60 cm x 1-4 mm, tan to brown,
glabrescent or with acicular and short-stellate or
furcate trichomes. Lamina herbaceous, not ver-
rucose, 1-pinnate-pinnatifid to about the middle
(and then with a long-pinnatifid apex) or to near
the tip (and with a short-pinnatifid apex). Buds
lacking. Rachis with a mixture of stiff, acicular,
sometimes red-tipped trichomes 0.5-1 mm and
furcate or stellate ones 0.1-0.2 mm long. Pinnae
2-10 x l-2(-2.5) cm, incised V3 their width or
less, sometimes merely shallowly crenate, often
auricled at acroscopic base, proximal 1-3 pairs
deflexed and slightly shortened. Segments mostly
3-5 mm wide, rounded or truncate at tip. Veins
mostly 3-7 (-1 1) pairs per segment, mostly simple,
proximal l-2(-2'^) pairs from adjacent segments
connivent at or below sinus with an excurrent vein
to sinus. Indument abaxially on costae and veins
of moderately dense to dense, spreading, acicular
trichomes 0. 1-1 mm, also with furcate and stellate
trichomes on costae, laminar tissue commonly with
acicular trichomes 0.1-0.2 mm or minute glands,
veins and laminar tissue adaxially with rather
dense adpressed trichomes 0.2-0.4 mm. Sori me-
dial, indusia reddish brown, densely setose along
margin; sporangia glabrous or with setae on spo-
rangial stalks.
Along streams or in shaded ravines, terrestrial,
in rock crevices, or epipetric, lowland and mon-
tane rain forests, 200-1300 m, San Martin, Hua-
56
FIELDIANA: BOTANY
nuco, Pasco, Junin, Ucayali, Ayacucho, Cuzco,
and Madre de Dios.
Ecuador to Bolivia, southern Brazil.
A specimen from Pasco (Smith 2900, F, MO) is
atypical in having pinnae incised nearly half their
width and in the presence of buds in axils of distal
pinnae. It was collected at higher elevation (1850
m) than all other specimens.
San Martin: Prope Tarapoto, Spruce 3946 (o, GH, K,
p, syntypes of Nephrodium jamesonii). San Martin, 5-
1 5 km E of Shapaja on road to Chazuta, Knapp & Mallet
7023 (F, MO). Huanuco: Along Rio Monzon near Rio
Huallaga at Tingo Maria, Croat 21203 (MO, uc). Pasco:
Prov. Oxapampa, 5 km SE of Oxapampa, D. Smith 2900
(F, MO, USM). Junin: La Merced, Killip & Smith 23728
(GH). Ucayali: Prov. Coronel Portillo, Km 328, camino
a Pucallpa, Ridoutt (GH). Ayacucho: Ayna, between
Huanta and Rio Apurimac, Killip & Smith 22725 (F).
Cuzco: La Convention, Potrero, 8 km W of Quillabam-
ba, Tryon & Tryon 5379 (F, GH, us, USM). Madre de Dios:
Prov. Manu, Atalaya, vicinity of Hda. Amazonia, 2-3
km W of village, Foster & Wachter 7444 (MO, USM).
66. Thelypteris macrotis (Hooker) R. Tryon,
Rhodora 69: 7. 1967.
Nephrodium macrotis Hooker, Sp. fil. 4: 86, t. 242B.
1862. TYPE: Peru, Tarapoto, Spruce 3979 (ho-
lotype, K; isotypes, A!, BM!, GH!, L, p, uc!).
Dryopteris macrotis (Hooker) Kuntze, Rev. gen. pi. 2:
813. 1891.
Stem suberect to erect, caudex 0.8-2 cm in di-
ameter, scales light brown, dull, ovate-lanceolate,
glabrous or with simple trichomes 0.1-0.2 mm.
Leaves several, clustered, 25-105 cm long, usually
somewhat dimorphic with fertile longer-petioled
and with contracted pinnae. Petiole 1 5-60 cm x
1.5—4.5 mm, tan to brown, glabrescent or with
acicular trichomes. Lamina herbaceous, not ver-
rucose, 1 -pinnate, with a confluent, pinnatifid apex.
Buds and sometimes plantlets often present in axils
of distal pinnae. Rachis with stiff, acicular tri-
chomes 0.3-1.5 mm, lacking furcate or stellate
ones. Pinnae 3.5-15 x 0.7-1. 6(-2) cm, subentire,
crenate, or very shallowly incised V5 their width or
less, with a narrow-triangular auricle to 1 cm long
at acroscopic base, proximal 2—4 pairs strongly
reflexed and slightly shortened, often with pinnae
to middle of blade reflexed and curved upward at
tip. Crenations mostly 3—4 mm wide, rounded or
truncate. Veins 3-7 pairs per vein group, mostly
simple, proximal 1-2 pairs from adjacent seg-
ments connivent at sinus or lowermost united be-
low sinus with an excurrent vein to sinus. Indu-
ment abaxially on costae and veins of moderately
dense to dense, spreading, acicular trichomes 0.2-
1 .5 mm, furcate and stellate trichomes completely
lacking, laminar tissue commonly with acicular
trichomes 0.2-0.3 mm, these sometimes ad-
pressed, veins and laminar tissue adaxially usually
with rather dense adpressed trichomes 0.2-0.4 mm.
Sori medial, indusia reddish brown, densely se-
tose; sporangia glabrous or often with setae on
sporangial stalks.
Along streams and shaded cliffs in wet tropical
and montane forests, 180-1000 m, Amazonas, San
Martin, Loreto, Huanuco, Junin, and Madre de
Dios.
Endemic to Peru.
This is one of the few species of subg. Goniop-
teris that completely lacks stellate or furcate tri-
chomes. Apparently, the lack of such trichomes is
derived in this species. As suggested by the ad-
pressed adaxial pubescence and similar blade dis-
section, T. macrotis is closely allied to, and pos-
sibly derived from T. jamesonii, which has
branched trichomes. Pinnae of T. macrotis are
more shallowly incised, more reflexed, and with
more pronounced auricles than in T. jamesonii.
Amazonas: Prov. Bagua, bank of Rio Maranon above
Cascadas de Mayasi, Wurdack 1973 (F, GH, uc, USM).
San Martin: Prov. Mariscal Caceres, Dist. Campanilla,
Cachihunushca, Rio Huallaga, Schunke V. 4284 (F). Lo-
reto: Pongo de Manseriche, Mexia 6383 (BH, F, GH, MO,
uc, USM). Huanuco: Prov. Pachitea, Dist. Honoria, 1 km
arriba del pueblo de Tournavista, Schunke 1250 (F, GH).
Junin: Hacienda Perene, Coronado 238, 248 (GH, uc).
Madre de Dios: Parque Nacional del Manu, Cocha Ca-
shu Biological Station, M. Foster P-84-69 (uc).
67. Thelypteris semihastata (Kunze) Ching, Bull.
Fan Mem. Inst. Biol., Bot. 10: 254. 1941.
Aspidium semihastatum Kunze, Linnaea 9: 91. 1834.
TYPE: Peru, (Huanuco), ad flum. Pampayacu,
Poeppig(ho\otype, LZ, destroyed; drawing of type,
B; possible isotype, MO!).
Lastrea poeppigiana Presl, Epim. bot. 40. 1849. TYPE:
Peru [erroneously, "Cuba"], Poeppig (holotype,
presumably PR).
Dryopteris semihastata (Kunze) Kuntze, Rev. gen. pi.
2: 291. 1891.
Stem short-creeping to suberect, ca. 5 mm in
diameter, scales light brown, dull, ovate-lanceo-
late, with simple trichomes 0.1-0.2 mm. Leaves
numerous, up to ca. 40 cm long, monomorphic.
Petiole ca. 5 cm x 1 mm, tan, glabrescent or with
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
57
acicular trichomes. Lamina herbaceous, not ver-
rucose, 1 -pinnate with a confluent, pinnatifid apex.
Bud sometimes present in axils of distal pinnae.
Rachis with stiff, acicular trichomes 0.3-1.5 mm,
lacking furcate or stellate ones. Pinnae 2-4 x 0.6-
1 cm, crenate or incised up to V2 their width or
less, with a slightly enlarged triangular auricle to
5 mm long at acroscopic base, proximal 3-6 pairs
reflexed and slightly shortened, sometimes curved
upward at tip. Crenations mostly 2-2.5 mm wide,
rounded. Veins 2-4 pairs per vein group, simple,
proximal pair from adjacent segments connivent
at or running towards sinus. Indument abaxially
on costae and veins of moderately dense to dense,
spreading, acicular trichomes 0.2-1 .5 mm, furcate
and stellate trichomes completely lacking, laminar
tissue commonly with acicular trichomes 0.2-0.3
mm, veins and laminar tissue adaxially usually
with rather dense adpressed trichomes mostly 0.2-
0.5 mm. Son medial, indusia reddish brown,
densely setose; sporangia glabrous or often with
setae on sporangial stalks.
Lower montane rain forest, 670 m, Huanuco.
Endemic to Peru.
This is closely related to T. macrotis and differs
by the narrower blades, shorter petioles, and grad-
ually reduced proximal pinnae. It also completely
lacks stellate trichomes.
Huanuco: Rio Bella, 7 km de Tingo Maria, Soukup
3091 (F, MO).
and with a confluent, pinnatifid apex. Bud often
present in axil of distal pinna. Rachis with a mix-
ture of acicular trichomes 0. 1-0.3 mm and furcate
or stellate ones 0.1 mm. Pinnae 5-10(-12) pairs,
often short-stalked to 3 mm, 8-15 x 1.5-3 cm,
incised V3—V2 their width, proximal ones narrowed
at the base and lacking auricles, lowermost spread-
ing or slightly deflexed. Segments mostly 3.5-7
mm wide, rounded or truncate at tip. Veins mostly
6-13 pairs per segment, simple, proximal pair from
adjacent segments connivent at or just below the
sinus (and then at an acute angle), rarely the low-
ermost pair united at an obtuse angle with an
excurrent vein. Indument abaxially on costae and
veins of sparse to moderately dense, acicular tri-
chomes mostly ca. 0.1-0.2(-0.3) mm, sometimes
also with furcate trichomes at base of costae, lam-
inar tissue glabrous, veins and laminar tissue gla-
brous adaxially. Sori medial, indusia brownish,
setose; sporangia glabrous or with setae on spo-
rangial stalks.
Lowland rain forests, 100-400 m, Loreto,
Huanuco, Ucayali, Cuzco, and Madre de Dios.
Hispaniola; Lesser Antilles; Trinidad and To-
bago; Guianas; Amazonian Brazil; Ecuador to Bo-
livia.
Schunke 2686 differs in lacking a bud in the axil
of a distal pinna, having fewer (5 or 6) pinna pairs,
and in having the lowermost pair of veins united
below the sinus with an excurrent vein to 5 mm.
68. Thelypteris abrupta (Desv.) Proctor, Rhodora
61: 306. 1959 [I960].
Polypodium abruptum Desv., Mem. Soc. Linn. Paris
6: 239. 1827. TYPE: Antilles, collector not stated
(holotype, P, Herb. Desvaux).
Goniopteris pyramidata Fee, Mem. Fam. Foug. 11:
61, /. 16, f. 2. 1866. TYPE: Guadeloupe, L'Her-
minier (holotype, P; isotypes, B, P).
Dryopteris pyramidata (Fee) Maxon, Contr. U.S. Natl.
Herb. 10: 489. 1908.
Stem short-creeping to suberect, 5-7 mm in di-
ameter, or caudex up to 1.5 cm in diameter, scales
light brown, dull, ovate-lanceolate, with stellate
and furcate trichomes 0. 1 mm. Leaves few, clus-
tered or not, 45-90 cm long, monomorphic or
slightly dimorphic with fertile longer-petioled and
with slightly contracted pinnae. Petiole 20-50 cm
x 1.5-4 mm, tan to brown, glabrescent or with
acicular and stellate trichomes 0. 1 mm. Lamina
herbaceous, not verrucose, 1 -pinnate-pinnatifid
Loreto: Prov. Maynas, Dtto. Alto Nanay, near Santa
Maria de Nanay, Simpson & Schunke 725 (F, GH). Prov.
Maynas, Yanamono Explorama Tourist Camp, halfway
between Indiana and mouth of Rio Napo, van der Werff
et al. 9864 (MO), 9963 (MO, uc). Lower Rio Huallaga,
Puerto Arturo, Yurimaguas, LI. Williams 5029 (F). Hua-
nuco: Prov. Pachitea, Dist. Honoria, Bosque Nacional
de Iparia, Rio Pachitea, Schunke V. 1375 (F, in part; this
no. also T. tristis). Ucayali: Prov. Coronel Portillo, Dist.
Iparia, Quebrada de Tabacoa, Rio Ucayali, Schunke 2686
(F, GH). Prov. Coronel Portillo, Dist. Calleria, Bosque
Nacional Alexander von Humboldt, (as Loreto), Schunke
V. 10411 (MO, uc). Cuzco: Prov. Paucartambo, Atalaya-
Carbon, Vargas 13429 (GH). Madre de Dios: Prov. Tam-
bopata, ca. 30 km SSW of Puerto Maldonado, Barbour
4997 (MO, uc), 5141 (F, MO, uc, USM), 5315 (F, MO, USM),
5727 (MO, uc). Tambopata, Albergue, "Cuzco Amazoni-
ca," Leon 885 (F, USM).
69. Thelypteris lugubriformis (Rosenst.) R. Tryon,
Rhodora 69: 7. 1967.
Dryopteris lugubriformis Rosenst., Repert. Spec. Nov.
Regni Veg. 7: 299. 1909. TYPE: Peru, [San Mar-
58
FIELDIANA: BOTANY
tin], Tarapoto, Spruce 4749 (holotype, P!; frag.,
us).
Goniopteris eggersii (Hieron.) Alston, Bull. Jard. Bot.
Etat27: 57. 1957.
Stem short-creeping, 0.8-1.5 cm in diameter, or
suberect with caudex up to 2.5 cm in diameter,
scales not seen. Leaves few, 90-120 cm long,
monomorphic. Petiole 40-60 cm x 3-7 mm, tan
to brown, with dense stellate trichomes 0.1-0.2
mm or glabrescent. Lamina thick-herbaceous to
subcoriaceous, not verrucose, 1-pinnate-pinnati-
fid with a confluent, pinnatifid apex or the apex
subhastate and more or less similar to the lateral
pinnae. Bud usually present in axil of distal pinna.
Rachis with a mixture of stout, acicular trichomes
to 1.5 mm and more numerous furcate or stellate
ones 0.1-0.3 mm. Pinnae 11-15 pairs, sessile or
stalked to 3 mm, 12-16 x 2.5-3.5 cm, incised V2-
V3 their width, lowermost slightly narrowed at the
base and lacking auricles, spreading or slightly de-
flexed. Segments mostly 4-6 mm wide, rounded
or truncate at tip. Aerophores peglike, to 0.5 mm
long at pinna bases. Veins mostly 12-18 pairs per
segment, simple, proximal pair from adjacent seg-
ments connivent at or just below the sinus, the
next 1-2 pairs running to sinus. Indument abaxi-
ally on costae, veins, and laminar tissue of nu-
merous acicular trichomes mostly 0.2-1.5 mm,
the costae also with furcate trichomes 0. 1-0.2 mm,
laminar tissue adaxially with numerous adpressed
trichomes 0.2-0.4 mm. Sori inframedial to me-
dial, indusia brownish, densely setose; sporangia
with setae on sporangial stalks.
Lowland rain forests, 100-800 m, San Martin,
Loreto, Pasco, Cuzco, and Madre de Dios.
Colombia to Bolivia.
Loreto: Ca. 50 mi downriver from Iquitos on Amazon
River at Peter Jensen's Explorama lodge, Moran 3650
(MO, uc, USM). Yanomono, Explorama Tourist Camp,
Rio Amazonas, Gentry et al. 36667 (MO, USM). Soledad,
on Rio Itaya, Killip & Smith 29559 (NY). Pasco: Puerto
Bermudez, Killip & Smith 26649 (NY). Cuzco: Prov. La
Convention, from Hda. Luisiana and Apurimac River
via cam pa trail through Polies Carpo's property, Dudley
11568 (GH). Madre de Dios: Prov. Manu, Manu Park,
Cocha Cashu uplands, Nunez 6116 (MO).
70. Thelypteris eggersii (Hieron.) Reed, Phyto-
logia 17: 274. 1968.
Nephrodium eggersii Hieron., Bot. Jahrb. Syst. 34:
441. 1904. TYPE: Ecuador, El Recreo, Eggers
15319 (holotype, B; photo, us).
Dryopteris eggersii (Hieron.) C. Chr., Index fil. 263.
1905.
Stem suberect to erect, caudex 1.5-3 cm in di-
ameter, scales light brown, appressed, tan, densely
stellate-pubescent. Leaves few, (50-)80-120(-160)
cm long, monomorphic. Petiole (20-)35-60 cm x
3-5 mm, brownish, with dense, stellate trichomes
0.1 mm. Lamina chartaceous, not verrucose,
1 -pinnate-pinnatifid, the apex confluent and pin-
natifid. Bud usually present in axil of distal pinna.
Rachis with dense, mostly stellate trichomes ca.
0. 1-0. 1 5 mm. Pinnae 1 5-20 pairs, sessile or stalked
to 2 mm, mostly 10-16 x 1.5-2.5 cm, incised %-
%(-%) their width, lowermost spreading, narrowed
at the base with several pairs of reduced segments,
auricles lacking. Segments mostly 3-5 mm wide,
truncate to acutish at tip. Veins mostly 9-12 pairs
per segment, proximal pair from adjacent seg-
ments connivent or at running to sinus. Indument
abaxially on costae of very dense, simple, furcate,
and stellate trichomes 0. 1-0.3 mm, veins and lam-
inar tissue with acicular trichomes 0.1-0.4 mm,
veins and laminar tissue adaxially glabrous or with
sparse trichomes toward margin. Sori medial, in-
dusia large, to 1 mm in diameter, reddish brown,
densely setose; sporangia glabrous or with setae
on sporangial stalks.
Montane rain forests, 1000 m, Junin.
Costa Rica to Panama; Colombia to Peru.
Among Peruvian Thelypteris, this is most sim-
ilar to T. biformata and differs by the very dense,
rather uniformly short stellate pubescence on the
costae abaxially. In addition, the lamina is more
gradually tapering distally, never abruptly re-
duced, and the number of lateral pinna pairs is
larger. Peruvian material differs from that of Cen-
tral America in the longer and more numerous
unbranched trichomes on costae and veins abax-
ially.
Junin: Chanchamayo valley, C. Schunke 56, 80, 84,
85 (all F).
71. Thelypteris erythrothrix A. R. Sm., sp. nov.
A speciebus ceteris subg. Goniopteridis distinguenda
laminae apice confluente pinnatifido, trichomatibus
atrorubris stellatis furcatis acicularibusque 0.1-0.5 mm
longis ad rhachim et costas abaxialiter dispositis, lamina
glabra adaxialiter, soris exindusiatis.
Stem suberect, caudex ca. 1 cm in diameter,
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
59
scales brown, glabrous or with a few stellate tri-
chomes 0.1 mm. Leaves few, 45-60 cm long,
monomorphic. Petiole 15-30 cm x 2 mm, tan,
glabrescent or with sparse stellate trichomes 0.1
mm. Lamina chartaceous, not verrucose, 1-pin-
nate-pinnatifid, the apex confluent and pinnatifid.
Bud lacking in axil of distal pinna. Rachis abaxi-
ally with moderate, red, mostly furcate and stellate
trichomes ca. 0.15-0.5 mm, adaxially the un-
branched trichomes deep red, more numerous, up
to 1 mm. Pinnae 10-12 pairs, sessile or stalked to
1 mm, mostly 6-9 x 1.6-2.2 cm, incised ca. V2
their width, lowermost spreading or slightly de-
flexed, with 1-2 pairs of reduced segments, auri-
cles lacking. Segments mostly 3—4 mm wide, trun-
cate to acute at tip. Veins mostly 8-1 1 pairs per
segment, proximal pair (or 1 V2 pairs) from adjacent
segments connivent at or just below sinus. Indu-
ment abaxially on costae and costules of sparse to
moderately dense, hyaline to often red, furcate and
stellate trichomes 0.2-0.5 mm, veins and laminar
tissue glabrous or with sparse hyaline acicular tri-
chomes 0.1-0.2 mm, adaxially with veins and
laminar tissue glabrous. Sori inframedial to me-
dial, indusia lacking; sporangia glabrous.
TYPE— Peru, Amazonas, Prov. Bagua, Cordille-
ra Colan SE of La Peca, Third camp, Barbour 3894
(holotype, MO!).
Epiphytic on tree bole and on rocky outcrop-
ping, 2100-2400 m, Amazonas, known only from
the type.
The deep red, stellate, furcate, and simple tri-
chomes on both sides of the rachis suggest that
this is related to T. curta (Christ) Reed (Costa Rica
to Ecuador and Venezuela), T. killipii A. R. Sm.
& Lellinger (Colombia and Ecuador), and T. peri-
pae (Sodiro) Reed (Ecuador and Colombia); T.
biformata may also be related. From all of these
species, T. erythrothrix differs in the absence of
indusia, sparser pubescence on rachis, costae, and
lamina abaxially, and glabrous lamina adaxially.
The elevation is unusually high for any species of
subg. Goniopteris.
72. Thelypteris biformata (Rosenst.) R. Tryon,
Rhodora69: 5. 1967.
Dryopteris biformata Rosenst., Repert. Spec. Nov.
Regni Veg. 7: 300. 1909. TYPE: Peru, (San Mar-
tin), nearTarapoto, ad rivulum Cachi-yacu, Spruce
4037 (holotype, P!; isotypes, BM!, K!; frag., us!).
1-3 cm in diameter, scales dark brown or brown,
densely stellate-pubescent. Leaves few, mostly 50-
100(-125) cm long, monomorphic or often sub-
dimorphic with fertile ones longer-petioled and
with contracted pinnae. Petiole 25-60 cm x 3-6
mm, tan to brownish, with dense, stellate tri-
chomes 0. 1 mm. Lamina herbaceous, not verru-
cose, 1 -pinnate-pinnatifid, the apex abruptly re-
duced and pinnalike, confluent with proximal
pinnae or often more or less free. Bud usually pre-
sent in axil of distal pinna. Rachis with dense,
stellate trichomes ca. 0.1-0.3 mm and longer hy-
aline to often red acicular ones to 1 .5 mm. Pinnae
9-16 pairs, sessile or stalked up to 5 mm, mostly
8-20 x 1.6-4 cm, incised 2/3-4/5 their width, low-
ermost spreading, narrowed at the base with sev-
eral pairs of reduced segments, auricles lacking.
Segments mostly 3-6 mm wide, rounded to acu-
tish at tip. Veins 10—22 pairs per segment, prox-
imal 1-2 pairs from adjacent segments connivent
at or running to sinus. Indument abaxially on cos-
tae of dense acicular trichomes to 1.5 mm and
more numerous short acicular, furcate, and stellate
ones 0.1-0.2 mm, veins and laminar tissue gla-
brous or with acicular trichomes 0. 1-0.5 mm, veins
and laminar tissue adaxially glabrous or with sparse
trichomes toward margin. Sori medial, indusia
reddish brown, setose; sporangia glabrous.
Lowland and occasionally montane rain forests,
1 00-1 700 m, Amazonas, San Martin, Loreto, Pas-
co, and Junin.
Ecuador; Peru; Amazonian Brazil.
This is a common and rather variable species
in Peru, and it probably hybridizes with other spe-
cies, e.g., T. tristis.
Amazonas: Prov. Bagua, along Quebrada Tambillo,
valley of Rio Maranon above Cascadas de Mayasi, Wur-
dack 1988 (GH, us, USM). Loreto: Balsapuerto, lower Rio
Huallaga basin, Killip & Smith 28532 (NY). Mishuyacu,
near Iquitos, Klug 312 (F, NY). Prov. Maynas, Dist. Iqui-
tos, Rio Momon, Puesto San Miguel, McDaniel 17316
(F, GH, MO, USM). Gamitanacocha, Rio Mazan, Schunke
15 (F, GH, NY, uc). Prov. Maynas, Yanamono, Explora-
ma Tourist Camp on Rio Amazonas between Indiana
and mouth of Rio Napo, Gentry et al. 29125 (F, MO, uc),
36667 (uc). Pasco: Pichis Trail, Yapas, (as Junin), Killip
& Smith 25533 (NY). Junin: Schunke Hacienda, above
San Ramon, Killip & Smith 24724 (NY). Cahuapanas, on
Rio Pichis, Killip & Smith 26785 (NY).
73. Thelypteris tristis (Kunze) R. Tryon, Rho-
dora69: 8. 1967.
Stem suberect to erect, caudex up to 1 0 cm long, Polypodium triste Kunze, Linnaea 9: 47. 1 834. TYPE:
60
FIELDIANA: BOTANY
Peru, Huallaga ad Mission Tocache, Poeppig(ho-
lotype, LZ, destroyed; presumed isotypes, B, K!).
Dryopteris tristis (Kunze) Kuntze, Rev. gen. pi. 2: 8 14.
1891.
Goniopteris tristis (Kunze) Brade, Bradea 1:217.1 972.
Stem short-creeping, to 1 cm in diameter, scales
brown, somewhat shining, with furcate or stellate
trichomes 0. 1 mm. Leaves few, mostly 60-1 75 cm
long, monomorphic or very slightly dimorphic with
fertile ones longer-petioled and with slightly con-
tracted pinnae. Petiole 20-90 cm x 3-7 mm, tan
to brownish, glabrous or with sparse stellate tri-
chomes 0. 1 mm. Lamina chartaceous, not verru-
cose, 1 -pinnate-pinnatifid, the apex abruptly re-
duced and pinnalike, more or less free. Bud usually
present in axil of distal pinna. Rachis glabrous or
with stellate trichomes 0.1-0.2 mm, sometimes
also with longer acicular ones to 1.5 mm. Pinnae
5-16 pairs, sessile or stalked up to 5 mm, mostly
15-25(-35) x 2-^(-6) cm, incised '/2-3/4 their width,
lowermost spreading or slightly ascending, nar-
rowed at the base with several pairs of reduced
segments and often a cuneate wing, auricles lack-
ing. Segments mostly 4-8 mm wide, often subfal-
cate, rounded to acutish at tip. Veins 9-22 pairs
per segment, proximal 2-3 pairs from adjacent
segments connivent at or running to sinus. Indu-
ment abaxially on costae lacking or of sparse to
moderately dense acicular trichomes 0.1-1.5 mm
and fewer furcate and stellate ones 0.1-0.2 mm,
veins glabrous or sometimes with acicular tri-
chomes, laminar tissue glabrous, veins and lami-
nar tissue glabrous adaxially or the veins occa-
sionally with scattered trichomes. Sori inframedial
to medial, often confluent at maturity, indusia small
to medium-sized, apparently lacking in a few spec-
imens, brownish, usually with trichomes 0.1-0.5
mm; sporangia glabrous.
Lowland and lower montane rain forests, 1 00-
500(-1200) m, Amazonas, San Martin, Loreto,
Huanuco, Ucayali, Ayacucho, Cuzco, and Madre
de Dios.
Panama; Colombia to French Guiana and Bo-
livia; Amazonian Brazil.
This is the most common member of subg. Go-
niopteris in Peru, with over 45 collections seen. It
is extremely variable in pubescence (glabrous or
with short to very long trichomes greater than 1
mm), size of indusia (minute to moderately sized,
seemingly absent in a few specimens), and pres-
ence or absence of buds in the axils of distal pin-
nae. Probably, T. tristis hybridizes with several
other species.
Amazonas: Prov. Bagua, Rio Utcumbamba on Cerro
Tapur, above Hda. Misqui, ca. 40 km S of Bagua Grande,
Hutchison 1487 (GH, uc). San Martin: San Martin, 5-
10 km NE of Shapaja on road to Cazuta, along Rio
Huallaga, Knapp & Mallet 6918 (F, MO). Loreto: Above
Pongo de Manseriche, mouth of Rio Santiago, Mexia
6111 (BH, F, G, GB, GH, MO, NY, UC, z). Huanuco: Dist.
Honoria, Bosque Nacional de Iparia, a lo largo del Rio
Pachitea cerca del campamento Miel de Abeja, Schunke
V. 1375 (F, GH). Ucayali: Bosque Nacional von Hum-
boldt, Km 88, Pucallpa-Tingo Maria road, Gentry et al.
36384 (MO, uc). Ayacucho: Rio Apurimac Valley, near
Kimpitiriki, Killip & Smith 22904 (NY). Cuzco: Prov. La
Convention, Rio Apurimac, below San Martin, Davis et
al. 1323 (GH). Madre de Dios: Prov. Tambopata, Puerto
Arturo, Vargas 18773 (GH).
74. Thelypteris juruensis (C. Chr.) R. Tryon &
Conant, Acta Amazonica 5: 33. 1975.
Dryopteris juruensis C. Chr., Kongel. Danske Vidensk.
Selsk. Skr., Naturvidensk. Afd., ser. 7, 10: 256.
1913. TYPE: Brazil, Amazonas, bei Bom Fim,
Rio Jurua, Ule 5325 (holotype, B; isotype, P).
Goniopteris juruensis (C. Chr.) Brade, Bradea 1: 216.
1972.
Stem short-creeping to suberect, caudex 0.8-2
cm in diameter, scales brown, with numerous stel-
late trichomes 0.1 mm. Leaves few, mostly 25-80
cm long, monomorphic or very slightly dimorphic
with fertile ones longer-petioled. Petiole 1 2-45 cm
x 1.5-4 mm, tan to brownish, glabrous or with
sparse stellate trichomes 0.1 mm. Lamina char-
taceous, verrucose, 1 -pinnate-pinnatifid, the apex
abruptly reduced and pinnalike, more or less free.
Bud(s) usually present in axil(s) of distal pinna(e).
Rachis glabrous or with sparse to moderately dense
furcate and stellate trichomes 0.1 mm. Pinnae
mostly 3-7 pairs, sessile or stalked up to 3 mm,
mostly 8-16 x 2-3 cm, crenate to incised up to
!/3 their width, all narrowed at the base with several
pairs of reduced segments, auricles lacking, low-
ermost spreading or slightly ascending. Segments
mostly 4-6 mm wide, rounded to truncate at tip.
Veins mostly 6-12 pairs per segment, proximal 2-
3 pairs from adjacent segments connivent at or
just below sinus or running to sinus, distal vein of
proximal pair arising from costa. Indument abax-
ially on costae lacking or of sparse acicular tri-
chomes 0.1 mm, occasionally with a few stellate
or furcate trichomes 0. 1 mm, veins and laminar
tissue glabrous on both sides. Sori inframedial,
indusia small or apparently lacking, brownish, gla-
brous or with trichomes 0. 1 mm; sporangia gla-
brous.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
61
Lowland rain forests, 100-350 m, Loreto and
Madre de Dios.
Ecuador; Peru; Bolivia; Amazonian Brazil;
French Guiana.
This is one of the most commonly collected ferns
in Loreto and Madre de Dios, wtih 25 collections
seen, but apparently rare or uncollected elsewhere
in Peru. Two collections from Ecuador, where it
was unknown until recently (Smith, 1983), are
Lawesson et al. 43361 (AAU) and Foster 3800 (us).
Loreto: Near mouth of Rio Napo, Croat 20209 (F, MO,
uc, USM). Yurimaguas, lower Rio Huallaga, Killip & Smith
27665. 27697 (NY). Madre de Dios: Prov. Tambopata,
ca. 30 air km SSW of Puerto Maldonado, Tambopata
Nature Reserve, Barbour 5130 (F, MO, uc, USM). Parque
Nacional del Manu, Cocha Cashu Biological Station, M.
Foster P-84-59 (uc), P-84-71 (MO).
75. Thelypteris tryonorum A. R. Sm., sp. nov.
A speciebus ceteris subg. Goniopteridis distinguenda
laminae apice libero pinnaformi, pinnis lateralibus vulgo
5-9-jugis incisis ad plus minusve tertiam partem lati-
tudinis basi angustatis, gemmis vel plantulis ex pinna-
rum distalium axillis ortis, 3-6-paribus venarum ex seg-
mentis contiguis ad sinum vel sub sinu conniventibus,
trichomatibus plerumque acicularibus simplicibus vel
nonnullis furcatis, 0. 1-0.2 mm ad costas abaxialiter dis-
positis, soris exindusiatis.
Stem suberect to erect, caudex 1-2 cm in di-
ameter, scales brown, with sparse stellate tri-
chomes 0.1 mm. Leaves few, mostly 75-100 cm
long, monomorphic or slightly dimorphic with fer-
tile ones longer-petioled and with slightly nar-
rowed pinnae. Petiole 35-55 cm x 3-5 mm, tan
to brownish, glabrescent or with sparse to mod-
erate stellate trichomes 0.1 mm. Lamina char-
taceous to subcoriaceous, not verrucose, 1-pin-
nate-pinnatifid, the apex abruptly reduced and
pinnalike, free. Bud(s) and sometimes plantlets
usually present in axil(s) of distal pinna(e). Rachis
with moderately to dense furcate and stellate tri-
chomes 0.1-0.2 mm. Pinnae mostly 5-9 pairs,
stalked up to 5 mm, mostly 12-20 x 3^4.5 cm,
incised ca. '/3 their width, all narrowed at the base
with several pairs of reduced segments, auricles
lacking, lowermost spreading or slightly ascend-
ing. Segments mostly 5-7 mm wide, truncate to
acute at the subfalcate tip. Veins mostly 12-18
pairs per segment, proximal 3-6 pairs from ad-
jacent segments connivent at or below sinus, distal
vein of proximal pair arising from costa. Indument
abaxially on costae, costules, and sometimes veins
of moderately dense to dense acicular and a few
furcate trichomes 0.1-0.2 mm, also with light
brown linear to lanceolate scales to 2 mm, laminar
tissue glabrous, veins and laminar tissue adaxially
glabrous. Sori inframedial, indusia lacking; spo-
rangia glabrous.
TYPE— Peru, Huanuco, Tingo Maria, Tryon &
Tryon 5248 (holotype, GH!; isotype, F!, USM!).
Hillsides and rocky slopes in wet montane for-
ests, 600-1500 m, Huanuco, Pasco, and Junin.
Known only from Peru.
Huanuco: Fundo Chela, Aguilar 936 (GH). Tingo Mar-
ia, Cueva de las Pavas, Aldave & Fernandez 5573 (GH).
60° slope at Rio Huallaga at Tingo Maria, Croat 21049
(MO), 21064 (MO, uc). Tingo Maria to Pucallpa, Ellen-
berg 3830 (GH). Pasco: Prov. Oxapampa, Valle del Pal-
cazu, Rio Cacazu, Cacazu, Leon 685 (F, USM). Junin: La
Merced, Killip & Smith 23717 (NY). Chanchamayo Val-
ley, C. Schunke 141 (F). La Merced, Chanchamayo,
Schunke (Rosenst. exs. 29) (P). La Merced-Chancha-
mayo, Soukup 1049 (F).
76. Thelypteris biolleyi (Christ) Proctor, Bull. Inst.
Jamaica, Sci. Ser. 5: 58. 1953. Figure 6.
Nephrodium nemorale Sodiro, Crypt, vase. Quit. 267.
1893. TYPE: Ecuador, cerca de San Miguel de
los Colorados, Sodiro (possible type material, P;
photo, us; frag., us!).
Aspidium biolleyi Christ in Pittier, Prim. fl. costaric.
3: 31. 1901. TYPE: Costa Rica, Tuis, pres Tur-
rialba, Pittier 11243 (holotype, P!; isotype, us!).
Dryopteris asterothrix Rosenst., Repert. Spec. Nov.
Regni Veg. 7: 305. 1909. TYPE: Peru, (San Mar-
tin), secus rivulum Cachi-yacu, Spruce 4659 (ho-
lotype, P; isotypes, w!; frag., us).
Thelypteris nemoralis (Sodiro) R. Tryon, Rhodora 69:
7. 196 7 (notching, 1936).
Thelypteris sodiroi Reed, Phytologia 17: 314. 1968. A
nom. nov. for T. nemoralis (Sodiro) R. Tryon.
Goniopteris biolleyi (Christ) Pic.-Ser., Webbia 31:251.
1977.
Stem short-creeping to erect, caudex 2-3 cm in
diameter, scales brownish, with numerous stellate
trichomes 0.1 mm. Leaves few, 75-175 cm long,
monomorphic. Petiole 40-70 cm x 3-5 mm, tan
to brownish, glabrescent or with stellate trichomes
0.1 mm. Lamina herbaceous, often verrucose,
1 -pinnate-pinnatifid, the apex gradually reduced,
pinnatifid, confluent. Bud absent in axil of distal
pinna. Rachis with moderately dense to often
dense, stellate trichomes 0.1-0.2 mm. Pinnae
mostly 20-30 pairs, sessile or stalked to 1 mm,
mostly 12-22 x 1.8-3 cm, incised ca. '/2-2/3 their
width, lowermost spreading or slightly ascending,
narrowed at the base with several pairs of reduced
62
FIELDIANA: BOTANY
segments, sometimes with a cuneate wing, auricles
lacking. Segments mostly 4-6 mm wide, rounded
to truncate at tip. Veins mostly 12-15 pairs per
segment, proximal pair from adjacent segments
usually uniting below sinus at 45-90° with an ex-
current vein to sinus, next 1-2 pairs running to
sinus. Indument abaxially on costae, costules, veins,
and laminar tissue of moderately dense to dense
stellate trichomes mostly 0.1 mm, adaxially the
veins and laminar tissue with appressed, stellate
trichomes 0.1 mm and fewer acicular ones to 0.8
mm on veins. Sori inframedial, exindusiate or with
a minute indusium; sporangia glabrous or with a
few stellate trichomes 0.1 mm.
Lowland and montane rain forests, 200-1500
m, San Martin, Loreto, and Junin.
Jamaica; southern Mexico; Guatemala to Pan-
ama; Colombia to Peru, Venezuela; Brazil.
In most other parts of the range this species has
anchor-shaped trichomes on the abaxial costae and
sometimes on the lamina tissue; such trichomes
are lacking in Peruvian specimens.
San Martin: Prope Tarapoto, Spruce 4659 (w). Loreto:
Above Pongo de Manseriche, mouth of Rio Santiago,
Mexia 6114 (BH, F, GB, GH, MO, NY, uc). Junin: Prov.
Tarma, E tributary of Rio Tullumayo along path to set-
tlement "Tirol," ca. 3 km SE of San Ramon, Iltis & lltis
248 (GH). Chanchamayo valley, C. Schunke 818 (F).
77. Thelypteris ancyriothrix (Rosenst.) A. R. Sm.,
Fl. Ecuador 18: 140. 1983.
Dryopteris ancyriothrix Rosenst., Repert. Spec. Nov.
Regni Veg. 7: 305. 1909. TYPE: Peru, (San Mar-
tin), in monte Guayrapurima, Spruce 4748 (ho-
lotype, P!; 2 pinnae from type, us!).
Stem suberect, scales brownish, with numerous
stellate trichomes 0.1 mm. Leaves few, 100-150
cm long, monomorphic. Petiole 50-75 cm x 3-5
mm, stramineous to tan, with anchor-shaped and
acicular trichomes to 0.5 mm. Lamina herbaceous,
often verrucose, 1 -pinnate-pinnatifid, the apex
gradually reduced, pinnatifid, confluent. Bud ab-
sent in axil of distal pinna. Rachis with moderately
dense to often dense anchor-shaped trichomes to
0.5 mm. Pinnae sessile or short-stalked, 15-20 x
2-3 cm, incised ca. % their width, lowermost
spreading or slightly ascending, narrowed at the
base with several pairs of reduced segments ta-
pering to a long-cuneate wing, auricles lacking.
Segments mostly 4-6 mm wide, rounded to trun-
cate at tip. Veins mostly 10-16 pairs per segment,
proximal pair from adjacent segments connivent
at sinus or uniting below sinus at an obtuse angle
with an excurrent vein to sinus, next pair running
to sinus. Indument abaxially on costae, costules,
veins, and laminar tissue of moderately dense to
dense anchor-shaped trichomes mostly 0.3-0.6
mm, also with a few longer acicular trichomes,
veins and laminar tissue adaxially with scattered
to numerous appressed, stellate trichomes 0. 1 mm.
Sori medial, exindusiate; sporangia glabrous.
Lowland rain forests, swampy sites, 1 30-260 m,
San Martin, Loreto, and Madre de Dios.
Ecuador and Peru.
San Martin: Mt. Guayrapurima, Spruce 4748 (P, us).
Loreto: Prov. Maynas, Yanamono Explorama Tourist
Camp, halfway between Indiana and mouth of Rio Napo,
van der Werffet al. 9908 (MO, uc). Madre de Dios: Prov.
Tambopata, ca. 30 km or 70-80 river km SSW of Puerto
Maldonado at effluence of Rio La Torre (Rio D'Orbi-
gny)/Rio Tambopata, Barbour 5326 (MO, uc).
78. Thelypteris schunkei A. R. Sm., sp. nov.
Inter species subg. Goniopteridis, T. urbanii (Sodiro)
A. R. Sm. proxima trichomatibus utrinque appressis ses-
silibus stellatis et indusiis stellato-pubescentibus, sed dif-
fert pinnis truncatis basi, pinnis proximalibus sessilibus
vel brevistipitatis, pinnis crenatis vel vix lobatis mar-
gine, (3-)4-6 paribus venarum ex segmentis contiguis
sub sinu conjunctis.
Stem suberect, caudex 1.5-3 cm in diameter,
scales light brown, with numerous appressed, stel-
late trichomes 0.1 mm. Leaves few, 70-160 cm
long, monomorphic. Petiole 30-80 cm x 3-8 mm,
tan, glabrescent or with stellate trichomes ca. 0. 1
mm. Lamina herbaceous, often verrucose, 1 -pin-
nate with pinnae shallowly lobed, the apex abrupt-
ly reduced, similar to lateral pinnae, nearly or quite
free. Bud absent in axil of distal pinna. Rachis with
moderately dense to often dense stellate trichomes
0. 1-0. 1 5 mm. Pinnae mostly 9-1 3 pairs, sessile or
stalked to 3 mm, mostly 15-20 x 2.8-4.5 cm,
crenately incised to lobed to ca. V4 their width,
lowermost spreading, not narrowed at the base,
truncate, auricles lacking. Segments mostly 5-8
mm wide, rounded to truncate at tip. Veins mostly
(8-) 10- 14 pairs per segment, proximal (3-)4-6
pairs from adjacent segments usually uniting be-
low sinus or the more distal ones connivent to a
common vein that runs to sinus. Indument abax-
ially on costae, costules, veins, and laminar tissue
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
63
of moderately dense to dense, stellate trichomes
0. 1-0. 1 5 mm, branches of trichomes appressed to
somewhat ascending, veins and laminar tissue
adaxially with appressed, stellate trichomes 0.1
mm, acicular trichomes absent or nearly so
throughout plant. Sori medial, indusiate, indusia
with dense stellate trichomes; sporangia often with
a few stellate trichomes 0. 1 mm.
TYPE— Peru, Junin, Chanchamayo valley, C.
Schunke 54 (holotype, F!).
Lower montane rain forests, 300-900 m, San
Martin and Junin.
This species is closely related to T. urbanii (So-
diro) A. R. Sm., from Nicaragua to Ecuador, but
differs in the truncate (vs. broadly to narrowly
cuneate) pinna bases, nearly sessile or short-stalked
proximal pinnae, crenate- margined or very shal-
lowly lobed pinnae, and greater number of pairs
of anastomosing veins. It is also very close to T.
pennata, but differs by the lighter green lamina,
sessile (vs. many short-stalked), stellate trichomes
on costae abaxially, arms of stellate trichomes of-
ten appressed (vs. often ascending), lowermost
veins uniting at an obtuse (vs. acute) angle, veins
all arising from costules (vs. lowermost one some-
times from costa), and more shallow-lobed pinnae
with margin crenate or lobed less than l/4 the dis-
tance to costa.
San Martin: Prov. San Martin, 4 mi E of Tarapoto,
Woytkowski 3521 1 (MO, uc). Prov. Lamas, Dist. Lamas,
2-4 km N of San Antonio, along Rio Cumbasa, Belshaw
3537 (GH, uc). Junin: La Merced, Chanchamayo, Sou-
kup 1062 (F). Chanchamayo Valley, C. Schunke 675 (F).
79. Thelypteris pennata (Poiret) Morton, Contr.
U.S. Natl. Herb. 38: 64. 1967.
Polypodium pennatum Poiret in Lam., Encycl. 5: 535.
9 Jan 1 804. TYPE: "Amer. Mend.," collector not
stated (holotype, P).
Polypodium megalodus Schkuhr, 24. Kl. Linn. Pfl.-
Syst. [Krypt. Gew.] 1: 24, /. 19b. 1804. TYPE:
Uncertain.
Dryoptehs megalodus (Schkuhr) Urban, Symb. antill.
4: 21. 1903.
Thelypteris megalodus (Schkuhr) Proctor, Bull. Inst.
Jamaica, Sci. Ser. 5: 61. 1953.
Goniopteris pennata (Poiret) Pic.-Ser., Webbia 31: 252
1977.
Stem suberect to erect, scales brown, with nu-
merous appressed, stellate trichomes 0.1 mm.
Leaves few, 60-100(-125) cm long, monomor-
phic. Petiole 30-50 cm x 3-5 mm, brownish, gla-
brescent or with stellate trichomes ca. 0.1 mm,
with a few persistent scales in proximal third.
Lamina herbaceous, verrucose, 1 -pinnate-pinnati-
fid, the apex abruptly reduced, similar to lateral
pinnae but more triangular, nearly or quite free.
Bud absent in axil of distal pinna. Rac his with
dense, stellate trichomes 0.1-0.15 mm. Pinnae
mostly 5-8(-10) pairs, stalked to 5 mm, mostly
14-18(-25) x 3-5 cm, shallowly lobed V3-V2 their
width, lowermost spreading, narrowed and broad-
ly cuneate at the base, auricles lacking. Segments
mostly 5-8 mm wide, rounded to truncate at tip.
Veins mostly 9-14 pairs per segment, proximal
pair from adjacent segments uniting below sinus
with next l-2(-3) pairs running to a common vein
or to sinus. Indument abaxially on costae, costules,
veins, and laminar tissue of moderately dense to
dense, stellate trichomes 0.1-0.15 mm, branches
of trichomes appressed to ascending, veins and
laminar tissue adaxially with appressed, stellate
trichomes 0.1 mm, acicular trichomes absent or
nearly so throughout plant. Sori medial, indusiate,
indusia with stellate trichomes; sporangia often
with a few stellate trichomes 0.1 mm.
Lowland rain forests, 100-200 m, Amazonas
and Loreto.
Antilles; Guianas; Venezuela; Colombia to Peru.
Lamina color in dried specimens of T. pennata
is dark gray-green, in marked contrast to the grass-
green or yellow-green leaves of T. schunkei.
Amazonas: La Poza, Rio Santiago, Berlin 3601 (MO).
Loreto: Prov. Maynas, ca. 10 km SW of Iquitos at zoo-
logical park, Moran 3669 (MO, uc). Prov. Maynas, ca.
50 mi downriver from Iquitos on Rio Amazonas at Jen-
sen's Explorama lodge, Moran 3725 (uc). Prov. Maynas,
Alpahuayo, ca. 26 km along Iquitos-"Nanto" (Nauta?)
road, van der Werff et al. 10229 (uc). San Antonio on
Rio Itaya, Killip & Smith 29437 (F). Soledad, on Rio
Itaya, Killip & Smith 29721 (F).
80. Thelypteris tetragona (Sw.) Small, Ferns s.e.
States 256. 1938.
Polypodium tetragonum Sw., Prod. 132. 1788. TYPE:
Jamaica, Swartz (holotype, s).
Goniopteris tetragona (Sw.) Presl, Tent, pterid. 183.
1836.
Dryopteris tetragona (Sw.) Urban, Symb. antill. 4: 20.
1903, not Kuntze( 1891).
Stem short-creeping, scales brown, with stellate
trichomes 0. 1 mm. Leaves few, mostly 50-100 cm
long, subdimorphic with the fertile longer-petioled
64
FIELDIANA: BOTANY
and with contracted pinnae. Petiole mostly 25-50
cm x 2—4 mm, tan, glabrescent or with stellate
trichomes ca. 0.1 mm. Lamina herbaceous, not
verrucose, 1 -pinnate-pinnatifid, the apex abruptly
reduced, similar to lateral pinnae, nearly or quite
free. Bud absent in axil of distal pinna. Rachis with
sparse to moderately dense, stellate and acicular
trichomes 0. 1-0. 1 5 mm. Pinnae mostly 6-10 pairs,
sessile, 7-15 x 1.5-2.5 cm, pinnatifid V2-2/3 their
width, lowermost spreading or deflexed, narrowed
with several pairs of reduced segments at the base,
auricles lacking. Segments mostly 3-5 mm wide,
rounded to truncate at tip. Veins mostly 6-10 pairs
per segment, proximal pair from adjacent seg-
ments united below sinus with an excurrent vein
to sinus, next pair running to or just above sinus.
Indument abaxially on costae of sparse to mod-
erately dense, mostly acicular and a few furcate
trichomes 0.1-0.3 mm, veins and laminar tissue
glabrous on both sides. Sori inframedial, exin-
dusiate; sporangia with acicular trichomes 0. 1 mm.
Edges of lowland forest, elevation not known,
rare in Peru, department unknown.
Eastern and southern Mexico to Panama; Flor-
ida; Antilles; Colombia to Peru and Surinam.
Department unknown: Without locality data, Soukup
1163, 1 164 (both F).
81. Thelypteris poiteana (Bory) Proctor, Bull. Inst.
Jamaica, Sci. Ser. 5: 63. 1953.
Polypodium crenatum Sw., Prod. 132. 1788, not
Forssk. (1775). TYPE: Jamaica, Swartz (holo-
type, s).
Lastrea poiteana Bory, Diet, class. 9: 233. 1825. TYPE:
"Guiane" [presumably French Guiana], com-
muniquee par Poiteau (presumably p, not found).
Dryopteris poiteana (Bory) Urban, Symb. antill. 4: 20.
1903.
Goniopteris poiteana (Bory) Ching, Sunyatsenia 5: 239.
1940.
Stem short-creeping, ca. 5-10 mm in diameter,
scales brown, with furcate and stellate trichomes
0.1 mm. Leaves few, mostly 50-100 cm long,
monomorphic or nearly so. Petiole mostly 25-50
x 2-5 mm, stramineous to tan, glabrescent or with
stalked-stellate trichomes 0.1-0.3 mm most nu-
merous in adaxial groove, sometimes also with a
few longer acicular trichomes. Lamina herbaceous,
not verrucose, 1 -pinnate, the apex abruptly re-
duced, free, similar to lateral pinnae. Bud(s) some-
times present in axil(s) of distal or proximal pin-
na(e). Rachis usually with moderately dense to
dense, stellate, furcate, and acicular trichomes
mostly 0.1-0.5 mm, sometimes glabrescent. Pin-
nae mostly 2-6 pairs, sessile or short-stalked to 2
mm, 9-15(-20) x 2.5-4.5(-6) cm, crenate to very
shallowly lobed to ca. 2 mm, lowermost spreading
or slightly ascending, narrowed at the base, auri-
cles lacking. Crenations mostly 3-6 mm wide,
rounded to truncate. Veins mostly 6-10 pairs per
vein group, proximal 3-5 pairs from adjacent
groups united below sinus with a common excur-
rent vein running to sinus. Indument abaxially on
costae, veins, and laminar tissue of moderately
dense to dense, mostly acicular trichomes mostly
0.2-1 mm, veins and laminar tissue adaxially sim-
ilarly vested. Sori medial, exindusiate; sporangia
with acicular trichomes 0.1-0.3 mm.
Margins of lowland rain forests, ca. 200 m, in-
frequent in Peru: Loreto and Huanuco.
Southern Mexico to Panama; Antilles; Colom-
bia to the Guianas and Peru; northern Brazil; Ga-
lapagos.
Throughout the range of T. poiteana, buds are
consistently borne in axils of proximal pinnae, a
unique placement in subg. Goniopteris, but com-
mon in species of subg. Meniscium. However,
Mexia 6367 (GH) has them in axils of distal pinnae,
as in many species of Goniopteris.
Loreto: Washintsa and vicinity, Rio Huasaga, Lewis
et al. 11747 (MO). Above Pongo de Manseriche, mouth
of Rio Santiago, Mexia 6367 (G, GH, MO, NY, uc). Lower
Rio Nanay, LI. Williams 371 (F). Huanuco: Tingo Maria,
Valle de Huallaga, Ridoutt (GH, USM).
82. Thelypteris clypeata (Morton) Kramer, Acta
Bot. Neerl. 18: 141. 1969.
Dryopteris clypeata Morton, Bull. Torrey Bot. Club
66: 52. 1 939. LECTOTYPE (chosen here): Pan-
ama, Puerto Obaldia, San Bias coast, Pittier 4309
(us 679433!; isolectotypes, us!).
Stem short-creeping to suberect, ca. 5-10 mm
in diameter, scales brown, with acicular trichomes
0.1 mm. Leaves few, to ca. 90 cm long, mono-
morphic or the fertile slightly longer-stalked and
with narrower pinnae. Petiole to ca. 60 cm x 2-
4 mm, tan, glabrous or glabrescent. Lamina char-
taceous, not verrucose, 1 -pinnate, apex similar to
lateral pinnae, free. Bud(s) absent. Rachis usually
glabrescent or with sparse acicular trichomes 0. 1
mm. Pinnae 3-5 pairs, stalked to 5 mm, mostly
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
65
10-20 x 3-6 cm, cuneate at base, entire to sub-
repand or very shallowly crenate, proximal ones
slightly ascending, broadly cuneate at the base,
auricles lacking, apex cuspidate or acuminate. Veins
mostly 10-17 pairs per vein group, regularly unit-
ing with veins from adjacent vein groups to form
rows of areoles between costa and margin (veins
meniscioid), secondary veins straight to subsig-
moid. Indument abaxially lacking or of sparse acic-
ular trichomes 0. 1 mm or less on costae and cos-
tules, veins and laminar tissue glabrous on both
sides, stellate and furcate hairs apparently lacking
everywhere. Sori medial, with vaulted indusia to
1.5 mm in diameter, bearing trichomes 0.1 mm;
sporangia glabrous.
Lowland rain forest, 100 m, Ucayali.
Panama, Peru.
This peculiar species seems transitional between
subgenera Goniopteris and Meniscium. It is ab-
errant in the former in the complete lack of furcate
and stellate trichomes; from Meniscium, it differs
from all known species by the very large indusia.
Ucayali: Rodal Semillero INIA, near Km 96, Pucall-
pa-Tingo Maria road, (as Loreto), Maas et al. 4591 (u,
2 sheets, USM).
The following collections, most of them incom-
plete, do not appear to match any known species
of subg. Goniopteris; they may represent unde-
scribed species or perhaps hybrids. Additional col-
lections are necessary before adequate descriptions
and diagnoses can be written.
Berlin 201 (F, MO, uc)— Amazonas, Rio Cenepa,
300 m. Perhaps most similar to T. biformata, but
the costal trichomes are all very short (0.2 mm or
less) and ascending.
Velarde 5452 (GH)— Junin, Pueblo Pardo, 700
m. Closest to T. lugubriformis but lacking aero-
phores, adaxial lamina lacking trichomes between
veins, texture thinner.
Davis et al. 1314 (GH, 3 sheets)— Cuzco, prov.
La Convention, Rio Apurimac, 20 min below
Puerto Capito above the Boca de Tigre rapids, 600
m. Most similar to T. tristis but differing in the
narrower pinnae, lamina distally gradually re-
duced (conform apical pinna lacking).
Croat 20491 (uc)— Loreto, Rio Tacsha Curaray.
Most similar to T. biformata but the costal tri-
chomes abaxially mostly stellate, 0. 1-0.2 mm, less
dense; pinnae ca. 3 cm broad with segments subfal-
cate, acute.
III'. Thelypteris subg. Meniscium.
Thelypteris subg. Meniscium (Schreber) Reed,
Phytologia 17: 254. 1968. Figure 7.
Meniscium Schreber, Gen. pi. (Linn.), ed. 8 (Schreber)
2: 757. TYPE: Polypodium reticulatum L. = The-
lypteris reticulata (L.) Proctor.
Lamina 1 -pinnate, with entire, undulate, cren-
ulate, or serrate pinnae, rarely the lamina simple;
proximal pinnae the longest or nearly so, distal
pinnae gradually to abruptly reduced with a con-
form or subconform apical pinna; aerophores ab-
sent; buds sometimes in axils of proximal pinnae.
Veins regularly anastomosing in pairs, with the
cross- veins forming regular areoles in 4-25 rows
between costa and margin, usually with a single
excurrent, generally free veinlet arising at the point
of fusion of the cross veins. Indument, if present,
of acicular trichomes, these sometimes curved. Son
on the cross-veins, oblong or linear, straight or
usually arcuate, infrequently in pairs between main
lateral veins, rarely the sporangia acrostichoid (also
arising between the veins), indusia always absent;
sporangia glabrous or setose from the stalk or cap-
sule; receptacular glands present in a few species.
Spores with prominent winglike ridges, surface pa-
pillate or reticulate, x = 36.
Subgenus Meniscium comprises about 20 trop-
ical American species, from Florida, the Antilles,
and southern Mexico to northern Argentina and
Paraguay. In general the species occur in lowland
and montane rain forests, often along streams.
Several species often grow in swampy sites. The
distinctive venation found in subg. Meniscium also
occurs in a few species of subg. Goniopteris and
in many Paleotropical species of Pronephrium. It
seems likely that this venation pattern has evolved
independently in Old and New World species
groups but is probably an indication of close af-
finity between Goniopteris and Meniscium.
References
MAXON, W. R., AND C. V. MORTON. 1938. The
American species of Dryopteris subgenus Me-
niscium. Bull. Torrey Bot. Club 65: 347-376.
MORTON, C. V. 1971. The proper disposition of
Meniscium macrophyllum Kunze. Amer. Fern
J. 61: 17-20.
66
FIELDIANA: BOTANY
Key to Species of subg. Meniscium
a. Lamina simple, undivided 83. T. gigantea
a. Lamina 1 -pinnate b
b. Fertile pinnae with the sporangia appearing acrostichoid; sterile pinnae commonly more than 5
cm wide c
c. Pinnae margins entire; sporangia arising from cross- veins and laminar tissue
84. T. macrophylla
c. Pinna margins often undulate; sporangia arising only from cross-veins d
d. Sori with trichomes (sporangiasters) from receptacle or sporangial stalks; pinnae usually
more than 4 cm wide 85. T. chrysodioides
d. Sori with receptacle and sporangial stalks glabrous; pinnae 1.5-3 (-4) cm wide
9 1 . T. salzmannii
b. Fertile pinnae with the sporangia in distinct sori confined to the arcuate cross-veins, or the sori
sometimes confluent at maturity; sterile pinnae 1-8 cm wide e
e. Pinnae uncinate-serrate, at least toward apex f
f. Distal pinnae greatly reduced; pinnae gradually attenuate from a broad base, mostly stalked;
veins and usually lamina strongly pubescent abaxially 89. T. serrata
f. Distal pinnae not or only slightly shortened; pinnae narrowly oblong, rather abruptly acumi-
nate-caudate at apex, mostly sessile; veins and lamina glabrous abaxially
90. T. consobrina
e. Pinnae undulate, crenulate, or usually subentire g
g. Sporangial stalks lacking trichomes h
h. Tubular yellow to orange glands borne on the receptacle or from the sporangial stalks
i
i. Pinnae rounded to truncate at base, 5-1 5 lateral pairs; areoles 1 3-20-seriate between
costa and margin; leaves usually more than 1 m long 97. T. andreana
i. Pinnae cuneate at the base, 2-5 lateral pairs; areoles 7-10-seriate between costa and
margin; leaves less than 1 m long 98. T. arcana
h. Tubular glands lacking from sorus j
j. Sori often biseriate between main lateral veins; pinnae 3-5 lateral pairs, broadly
elliptic, cuneate at the base; distal pinnae strongly adnate and decurrent
96. T. lingulata
j. Sori nearly always uniseriate between main lateral veins; pinnae usually more than
5 lateral pairs, lanceolate or narrowly elliptic, truncate or cuneate at the base; distal
pinnae not adnate k
k. Areoles of sterile pinnae about as broad as long, the secondary veins straight or
nearly so 1
1. Pinnae entirely glabrous, the proximal ones subcordate, sessile
94. T. ensiformis
1. Pinnae sparsely pubescent on the costae and veins, the proximal ones cuneate
at base, long-stalked 95. T. falcata
k. Areoles of sterile pinnae much shorter than broad, the secondary veins curving,
sometimes subsigmoid m
m. Costae, veins, and lamina abaxially with numerous short-stipitate glands 0. 1
mm, trichomes entirely lacking 88. T. maxoniana
m. Costae, veins, and lamina abaxially lacking glands, trichomes present at least
on costae and veins n
n. Pinnae narrowly cuneate at the base, usually less than 2(-2.5) cm wide;
costae abaxially with appressed, curved trichomes; areoles 4-10(-13)-se-
riate 93. T. angustifolia
n. Pinnae broadly cuneate, rounded, or truncate at the base, 2-4 cm wide;
costae abaxially with mostly spreading, crispate or straight trichomes; ar-
eoles mostly 8-1 6-seriate o
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III. 67
o. Secondary veins (cross-veins) nearly straight, forming more or less rect-
angular areoles; costal trichomes dense; fertile pinnae with the sporangia
in distinct sori 92. T. lancea
o. Secondary veins curved or subsigmoid, areoles not rectangular; costal
trichomes moderately dense; fertile pinnae with the sporangia appearing
acrostichoid 91. T. salzmannii
Sporangial stalks often bearing an acicular trichome, or sterile sporangia (sporangiasters)
with such trichomes p
p. Trichomes of costae and lamina abaxially appressed; pinnae 4-7 cm wide
86. T. membranacea
p. Trichomes of costae and lamina abaxially spreading; pinnae 2-8 cm wide q
q. Pinnae mostly 2-3(-4) cm wide 87. T. arborescens
q. Pinnae 3.5-8 cm wide 85. T. chrysodioides
83. Thelypteris gigantea (Mett.) R. Tryon, Rho-
dora69:6. 1967.
Meniscium giganteum Mett., Fil. Lechl. 1:19. 1856.
TYPE: Peru, (Puno), "St. Gavan" (San Gaban),
Lechler 2292 (holotype, B; isotype, K).
Dryopteris gigantea (Mett.) C. Chr., Index fil. 267.
1905 (notKuntze, 1891).
Dryopteris simplicifrons C. Chr., Index fil. 486. 1906,
nom. nov. for Dryopteris gigantea (Mett.) C. Chr.,
not Kuntze.
Stem short-creeping and then ascending to erect.
Leaves few, mostly 40-1 30 cm long, subdimorphic
with fertile longer-petioled and with narrower
blade. Petiole to 60 cm x 3-5 mm, brown, with
scattered persistent lanceolate brown scales and
dense, thin, appressed trichomes. Lamina simple,
ovate to elliptic, mostly 30-70 x 7-15 cm, base
rounded to cuneate, margin entire to crenate, apex
acute to acuminate. Bud lacking. Rachis with dense,
appressed trichomes like those of petiole. Veins
from rachis ca. 8-10 mm apart on sterile blades,
giving rise to more or less straight secondary veins
that unite and give rise to an excurrent veinlet,
these veinlets partially or completely bisecting the
areole, daughter areoles rhomboid; areoles in rows
of 25-35 between rachis and margin. Indument
abaxially of numerous adpressed, thin trichomes
ca. 0.3-0.5 mm long on veins, laminar tissue gla-
brous on both sides. Sori linear-arcuate along nearly
the entire length of secondary veins; sporangial
stalks with long-stalked, red to orange, spherical
glands.
In lower montane rain forests, terrestrial or on
wet rocks, 600-1100 m, San Martin, Huanuco,
Pasco, Cuzco, and Madre de Dios.
Nicaragua to Panama; Colombia to Peru.
San Martin: Prov. Mariscal Caceres, Dist. Campani-
lla, 7.4 km N of Pulcache, Plowman & Schunke V. 11595
(F, USM). E of Tingo Maria, Allard 21575 (GH), 22297
(GH). Huanuco: Prov. Huanuco, Tingo Maria, Tryon &
Tryon 5282 (F, GH, USM). Pasco: Pichis Trail, San Ni-
colas, (as Junin), Killip & Smith 26018 (GH, us). Cuzco:
Quince Mil, Lockwood 569 (GH). Madre de Dios: Prov.
Manu, Cerro de Pantiacolla, Rio Palotoa, 10-15 km
NNW of Shintuya, R. Foster et al. 10902 (F).
84. Thelypteris macrophylla (Kunze) Morton,
Amer. Fern J. 61: 17. 1971.
Meniscium macrophyllum Kunze, Flora 22, Beibl. 1:
44. 1839. TYPE: Brazil, Bahia, Serra do Mar,
Martins 363 (isotypes, BR, K, NY).
Meniscium guyanense Fee, (Mem. fam. foug. 5) Gen.
fil. 224. 1852. TYPE: French Guiana, ad amnes
Conana et Gaberet, Leprieur (holotype, P?).
Acrostichum fendleri Baker, J. Bot. 25: 100. 1887.
TYPE: Trinidad, Fendler 88 (holotype, K; iso-
types, NY, uc!, us).
Dryopteris macrophylla (Kunze) C. Chr., Index fil.
suppl. 1: 35. 1913.
Dryopteris anceps Maxon, Contr. U.S. Natl. Herb. 24:
62. 1922. Based on Acrostichum fendleri Baker.
Bolbitis macrophylla (Kunze) Maxon & Morton, Bull.
Torrey Bot. Club 65: 375. 1938.
Thelypteris guyanensis (Fee) Morton, Amer. Fern J.
61: 19. 1971.
Stem creeping. Leaves few, mostly 1-2 m long,
subdimorphic to dimorphic, the fertile with longer
petioles and contracted, more numerous pinnae.
Petiole mostly 50-65 cm x 4-10 mm, tan to
brownish, glabrescent. Lamina 1 -pinnate, the ster-
ile with l-3(-6) pairs of pinnae and a larger ter-
minal one, fertile pinnae ca. 4-7 lateral pairs; ster-
ile pinnae up to 35 x 5-12 cm, short-petiolulate
to sessile or sometimes slightly adnate to rachis,
the base rounded to cuneate, margin entire or sub-
repand, apex acuminate; fertile pinnae ca. 8-20 x
68
FIELDIANA: BOTANY
2-5 cm, stalked to 1 cm. Bud lacking. Rachis gla-
brous or glabrescent. Veins arising from costae ca.
3-5 per 3 cm on sterile pinnae, 8-15 per 3 cm on
fertile pinnae, secondary veins straight or nearly
so, uniting at an obtuse angle and giving rise to an
excurrent veinlet, these veinlets mostly free and
partially bisecting the areole; sterile areoles in rows
of 16-30 between costa and pinna margin. Indu-
ment on sterile blades abaxially lacking or of sparse,
thin trichomes 0.1-0.5 mm on costae and veins,
laminar tissue usually glabrous on both sides; in-
dument of fertile blades abaxially sparsely to de-
cidedly pubescent with thin trichomes 0. 1-0.5 mm
long on costae, veins, and laminar tissue, adaxially
with fewer similar trichomes to glabrescent. Spo-
rangia appearing to cover the lamina (acrosti-
choid) on fully fertile blades; sporangial stalks gla-
brous (ours) or with minute trichomes 0.1 mm
arising from capsule.
Lowland rain forests, 400-500 m, Loreto and
Pasco.
Trinidad and Tobago; Guianas; eastern Vene-
zuela; Ecuador; Peru; northern Brazil.
Because of it acrostichoid sporangia, this species
was thought until rather recently to belong to Bol-
bitis, but it undoubtedly belongs in subg. Menis-
cium (Morton, 1971). It may be closely related to
Thelypteris chrysodioides.
Loreto: Above Pongo de Manseriche, ca. 1 km from
mouth of Rio Santiago, Mexia 6202 (F, GH, MO, uc, us),
6296 (GH). Quebrada Tahuayo above Tamishiyaco, Croat
19735 (F, GH, MO, uc, USM). Altura Tula Pichco on Rio
Napo, Croat 20309 (F, GH, MO, uc). Pasco: Prov. Oxa-
pampa, Palcazu Valley, Iscozacin, R. Foster 9496 (MO,
USM). Prov. Oxapampa, Palcazu, Rio Alto Iscozacin, Ozuz
to Rio Lobo, R. Foster & d'Achille 10068 (F, USM).
85. Thelypteris chrysodioides (Fee) Morton var.
goyazensis (Maxon & Morton) Morton, Contr.
U.S. Natl. Herb. 38: 51. 1967.
Dryopteris chrysodioides (Fee) Maxon & Morton var.
goyazensis Maxon & Morton, Bull. Torrey Bot.
Club 65: 374. 1938. TYPE: Brazil, Est. Goyaz,
Rio Corumba, Glaziou 22631 (holotype, NY; iso-
types, c, F!, uc!).
Stem unknown, probably creeping. Leaves
mostly 70-200 cm long, monomorphic or subdi-
morphic, fertile leaves with longer petiole and con-
tracted pinnae. Petiole up to 100 cm x 15 mm,
tan, glabrescent. Lamina 1 -pinnate, with up to ca.
10 pairs of lateral pinnae and a similar but slightly
smaller terminal one; pinnae 18—40 x 3.5-8 cm,
stalked up to ca. 5 mm, the base rounded to cu-
neate, margin subentire to usually subrepand or
crenate, apex acuminate. Bud lacking. Rachis with
sparse trichomes or glabrescent. Veins arising from
costae ca. 5-9 per 3 cm, secondary veins straight
or subsigmoid, uniting at an obtuse to acute angle
and giving rise to an excurrent veinlet, these vein-
lets free or completely bisecting the areole; sterile
areoles in rows of 1 5-28 between costa and pinna
margin. Indument of sterile blades abaxially of
moderately dense to occasionally dense, spreading
trichomes mostly 0.1-0.3(-0.4) mm on costae,
veins, and laminar tissue, lamina adaxially nearly
or quite glabrous. Sori elongate-arcuate on the
cross-veins, sometimes confluent at maturity,
mixed with filamentous structures (sporangias-
ters?) bearing 1-3 trichomes; sporangial stalks gla-
brous or with trichomes to 0.2 mm.
Lowland forests, 100-260 m, San Martin, Lo-
reto, and Madre de Dios.
Guyana; Brazil; Ecuador to Bolivia.
Klug 3970 and Ridoutt are largely sterile and so
are determined only provisionally. Sterile fronds
of this species seem to be much less pubescent
abaxially than the fertile ones. In venation and
length and shape of sori, this species is similar to
T. membranacea, which differs in having ap-
pressed trichomes on costae and lamina abaxially.
This species is now known from Ecuador: Napo,
Rio Lagarto Cocha, near Redondo Cocha, 1 90 m,
Lawesson et al. 44383 (AAU).
San Martin: Chaquta, Rio Huallaga, Klug 3970 (GH,
MO, NY, USM). Loreto: Mishuyacu, near Iquitos, Klug 200
(F, NY), 1533 (F, NY). Junin: Prov. Jauja, Satipo, Ridoutt
[USM 1 1661] (GH). Madre de Dios: Prov. Tambopata, 30
air km SSW Puerto Maldonado at effluence Rio La Tor-
re/Rio Tambopata, Barbour 5182 (MO, uc).
86. Thelypteris membranacea (Mett.) R. Tryon,
Rhodora69: 7. 1967.
Phegopteris membranacea Mett., Fil. lechl. 2: 22. 1859.
LECTOTYPE (chosen by Maxon & Morton, 1 938,
p. 366): Peru, (Puno), prope Azangaro, Lechler
1785 (isolectotype, GH!; photo, GH of K).
Nephrodium lechleri Hieron., Bot. Jahrb. Syst. 34: 448.
1904, nom. superfl. for Phegopteris membranacea
Mett. and with the same type.
Dryopteris lechleri (Hieron.) C. Chr., Index fil. 274.
1905, nom. superfl.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
69
Stem ascending? Leaves mostly 100-150 cm
long, monomorphic. Petiole up to 100 cm x 8
mm, brownish, pubescent with thin short tri-
chomes to 0.3 mm. Lamina 1 -pinnate, with ca. 4-
6 pairs of lateral pinnae and a similar terminal
one; pinnae 1 8-30 x 4-7 cm, sessile or stalked to
ca. 5 mm, the base rounded or truncate, margin
subentire or obscurely crenulate, apex acuminate.
Bud(s) present in axil of proximal pinna(e). Rachis
with often dense, more or less appressed, thin,
crispate trichomes. Veins arising from costae of
fertile pinnae ca. 8-10 per 3 cm, secondary veins
straight, uniting at an obtuse angle and giving rise
to an excurrent veinlet, these veinlets free or com-
pletely bisecting the areole into two equal squares;
sterile areoles in rows of ca. 1 8-20 between costa
and pinna margin. Indument on sterile blades
abaxially of moderately dense to dense, thin, ap-
pressed trichomes mostly 0.2-0.4 mm on costae,
veins, and laminar tissue, lamina adaxially nearly
or quite glabrous. Sori elongate, straight or slightly
arcuate on the cross-veins, not confluent at ma-
turity, mixed with filamentous structures (spo-
rangiasters?) bearing 1-3 trichomes; sporangial
stalks glabrous or with trichomes to 0.2 mm.
Lowland rain forests, 250-650 m, San Martin,
Cuzco, and Puno.
Colombia to Peru; Venezuelan specimens at-
tributed to this species seem to represent an un-
described species.
This species is aptly named, being thinner in
texture than most other species of subg. Menis-
cium. It differs further in the peculiar, appressed,
and thin pubescence (a character it shares with T.
giganted) and in the straight (rather than curved)
sori. The Klug collections cited by Maxon and
Morton (1938) under this species are referred here
to T. chrysodioides.
San Martin: Mt. Campana, prope Tarapoto, Spruce
4645 (K, not seen, us). Cuzco: Prov. Quispicanchi, entre
Inambari y Quince Mil, Vargas 16501 (GH).
87. Thelypteris arborescens (Willd.) Morton,
Contr. U.S. Natl. Herb. 38: 50. 1967.
Meniscium arborescens Willd., Sp. pi. ed. 4. 5: 133.
1810. TYPE: Venezuela, Humboldt & Bonpland
(holotype, B, Herb. Willd. 195 76; microfiche, uc).
Phegopteris mollis Mett., Ann. Sci. Nat. Hot., V, 2:
242. 1864. TYPE: Colombia, Triana (holotype,
B?; isotype, BM!).
Dryopteris permollis Maxon & Morton, Bull. Torrey
Bot. Club 65: 372. 1938. Based on Phegopteris
mollis Mett., not Dryopteris mollis (Jacq.) Hieron.
Stem short-creeping to ascending. Leaves most-
ly 70-150 cm long, monomorphic or nearly so.
Petiole 30-70 cm x 5-8 mm, stramineous to tan,
glabrescent to pubescent with trichomes to 0.3 mm.
Lamina 1 -pinnate, with ca. 10-25 pairs of lateral
pinnae and a similar terminal one; pinnae mostly
12-22 x 2-3.5 cm, sessile or lowermost stalked
to ca. 2(-5) mm, the base rounded or truncate,
margin subentire or obscurely crenulate, apex acute
to acuminate. Bud(s) absent. Rachis with often
sparse to moderate, spreading trichomes. Veins
arising from costae of fertile pinnae ca. 12-19 per
3 cm, secondary veins straight (fertile) to arcuate
or subsinuate (sterile), uniting at an obtuse (fertile)
to acute (sterile) angle and giving rise to usually
free excurrent veinlet; sterile areoles in rows of ca.
8-15 between costa and pinna margin. Indument
abaxially of moderately dense to dense, spreading
trichomes mostly 0.2-0.5 mm on costae, veins,
and laminar tissue, lamina adaxially nearly or quite
glabrous except along costae. Sori oblong, straight
or slightly arcuate on the cross- veins, sometimes
confluent at maturity, mixed with trichomes like
those of axes; sporangial stalks glabrous or with
trichomes to 0.2 mm.
In lowland and montane forests, along streams,
100-1 100 m, San Martin, Loreto, Huanuco, Pas-
co, and Puno.
Honduras to Panama; Colombia to Venezuela
and Bolivia; Brazil.
The differences between this species and T. lon-
gifolia (Desv.) R. Tryon are slight and the two
should perhaps be combined. Thelypteris longi-
folia is said (by Maxon & Morton, 1938) to differ
by the cuneate pinna bases, longer stalked pinnae,
and shorter and less dense pubescence. I see nearly
a continuum between the extremes.
San Martin: NW of San Martin, Rioja, Rio Negro,
Soukup 5279 (GH). Rioja, NW of San Martin, Rio Negro,
Soukup 5221 (GH). Loreto: Quebrada de Nauta, along
Rio Maranon, Croat 17530 (MO). Brillo Nuevo, Ya-
guasyacu River, Dist. Pebas, ca. 1 50 km ENE of Iquitos,
Treacy & Alcorn 253A (F). Prov. Loreto, Nauta, Rio
Maranon above mouth of Rio Ucayali, Gentry et al.
29975 (MO). Prov. Alto Amazonas, Yurimaguas, ca. 30
km towards Tarapoto, Hormia 2086 (H). Huanuco: Pam-
payacu, Kanehira 129 (GH, us). Pasco: Prov. Oxapampa,
Iscozacin, near confluence of Rio Palcazu and Rio Is-
cozacin, D. Smith 1952 (MO). Puno: Prov. Sandia, be-
tween Rio Azata-Colorado, Nunez & Munoz 5347 (MO).
70
FIELDIANA: BOTANY
88. Thelypteris maxoniana A. R. Sm., sp. nov.
Figure 7.
Dryopteris desvauxii f. glandulosa Maxon & Morton,
Bull. Torrey Bot. Club 65. 372. 1938. TYPE: Bra-
zil, Sao Paulo, Morro das Pedra, Brade 5753 (ho-
lotype, NY).
Thelypteris longifolia R. Tryon f. glandulosa (Maxon
& Morton) Morton, Contr. U.S. Natl. Herb. 38:
52. 1967.
A speciebus ceteris subg. Meniscii distinguenda tri-
chomatibus ad laminam abaxialiter nullis, et glandibus
abundantibus luteolis brevistipitatis 0.1 mm longis ad
costas et venas et spatia intervenas.
Stem short-creeping to ascending. Leaves most-
ly 80-150 cm long, monomorphic or nearly so.
Petiole 30-90 cm x 5-8 mm, stramineous to tan,
glabrous. Lamina 1 -pinnate, with ca. 10-20 pairs
of lateral pinnae and a similar terminal one; pinnae
mostly (12-)20-35 x (i_)2-3(-4) cm, sessile or
lowermost stalked to ca. 5 mm, the base cuneate
to rounded or truncate, margin subentire or ob-
scurely crenulate, apex acute. Bud(s) absent. Ra-
chis glabrous or sparsely glandular. Veins arising
from costae of fertile pinnae ca. 8-13 per 3 cm,
secondary veins straight (fertile) to arcuate or sub-
sinuate (sterile), uniting at an obtuse (fertile) to
acute (sterile) angle and giving rise to usually free
excurrent veinlet; areoles in rows of ca. 8-15 be-
tween costa and pinna margin. Indument abaxially
of moderately dense to dense, light yellowish, short-
stipitate glands 0. 1 mm long on costae, veins, and
laminar tissue, lamina adaxially glabrous, even on
costae. Son oblong, straight or arcuate on the cross-
veins, sometimes confluent at maturity, lacking
trichomes; sporangial stalks glabrous.
TYPE— Peru, Prov. Maynas, Quistococha, vicin-
ity of Iquitos, Gentry 20751 (holotype, MO!; iso-
type, uc!).
In swamps, along roadsides and streams, often
on white sand, 100-130 m, Loreto.
Colombia; Bolivia; probably Brazil.
It is uncertain whether the type off. glandulosa,
from southern Brazil, is really conspecific with the
Peruvian plant. In any case, the epithet glandulosa
cannot be transferred to Thelypteris because of the
preexisting T. glandulosa (Desv.) Proctor. Thus, I
give this species a new description and type.
This is actually one of the most easily recogniz-
able species of subg. Meniscium, and it certainly
warrants more than form status given it by Maxon
and Morton. It is easily distinguished from T. lon-
gifolia by the complete lack of trichomes on the
lamina abaxially. It is possible that its closest rel-
ative is indeed that species because of the general
similarity in size, venation, and pinna shape. The-
lypteris maxoniana seems to be one of the com-
monest menisciums in Amazonian Peru.
Loreto: 7 km NE of Rio Nanay, Croat 18312 (MO). 7
km NE of Puerto Almendra at Rio Nanay, Croat 18333
(F, MO, uc, USM). 17 km SW of Iquitos, Croat 18435
(MO, uc). Quebrada Tahuayo, above Tamishiyaco, Croat
19759 (MO). Prov. Maynas, Dist. Iquitos, Rio Nanay,
trail between Rio Mazan and Picuruyacu, Hickok 613
(GH). Mishuyacu, near Iquitos, Klug 509 (F, us), 7255
(F, NY). Prov. Maynas, San Juan, Midler & Muller 2517
[USM 14231] (GH). Prov. Maynas, Dist. Iquitos, Puerto
Almendra, Rio Nanay, Revilla 1249 (MO). Vicinity of
Iquitos, Revilla 3515 (MO). Prov. Maynas, Quistacocha,
Sagdstegui & Aldave 5794 (GH). Prov. Maynas, 3 km S
of Iquitos, Tryon & Tryon 5164 (GH, USM).
89. Thelypteris serrata (Cav.) Alston, Kew Bull.
1932: 309. 1932.
Meniscium serratum Cav., Descr. pi. 548. 1 802. TYPE:
Cuba, Havana, Guio (holotype, MA).
Dryopteris serrata (Cav.) C. Chr., Index fil. 29 1 . 1 905.
Stem short-creeping to ascending. Leaves most-
ly 100-200 cm long, monomorphic or subdi-
morphic, the fertile with generally contracted pin-
nae. Petiole 30-90 cm x 5-18 mm, stramineous
to tan, glabrous or glabrescent. Lamina 1 -pinnate,
with ca. 15-25 pairs of gradually shortened lateral
pinnae and a small, lanceolate, terminal one; pin-
nae mostly 12-25 x 2-3.5(-4.5) cm, sessile or
lowermost stalked to ca. 4 mm, the base rounded
or truncate, margin serrate or uncinate-serrate, apex
acute. Bud(s) absent or occasionally borne at base
of proximal pinnae. Rachis with sparse to dense
trichomes. Veins arising from costae of fertile pin-
nae ca. 8-13 per 3 cm, secondary veins straight
(fertile) to arcuate or subsinuate (sterile), uniting
at an obtuse (fertile) to acute (sterile) angle and
giving rise to usually free excurrent veinlet; areoles
in rows of ca. 10-15 between costa and pinna mar-
gin. Indument abaxially of moderately dense to
dense, spreading or curved trichomes mostly 0. 1 5-
0.3 mm on costae, veins, and sometimes laminar
tissue, lamina adaxially glabrous except along cos-
tae. Sori oblong to linear, straight or arcuate on
the cross- veins, often somewhat confluent at ma-
turity, lacking trichomes; sporangial stalks gla-
brous.
Marshes, swamps, roadside ditches, along
streams, sometimes in full sun, 100-900(-1650)
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
71
2mm
'I
5cm
FIG. 7 Subgenus Meniscium. Thelypteris maxoniana: a, leaf apex; b, portion of pinna, adaxial side; c, portion of
pinna, abaxial side. (From Croat 18435, uc.)
72
FIELDIANA: BOTANY
m, Amazonas, Loreto, Junin, Cuzco, and Madre
de Dios.
Florida; Antilles; Mexico to Panama; Colombia
to Guianas and Bolivia; Brazil; northern Argen-
tina; Paraguay.
Amazonas: Prov. Bagua, Chiriaco to Puente Venezue-
la, 3.9 km NE of Chiriaco, Barbour 4336 (MO, USM).
Loreto: Iquitos, Killip & Smith 27036 (F, GH, us). Mis-
huyacu, near Iquitos, Klug 431 (F, us). Rio Paranapura
above Yurimaguas, Croat 17942 (MO). Pasco: Oxapam-
pa, Rio Iscozacin tributary of Rio Palcazu, Knapp et al.
7835 (MO). Junin: Satyro alrededores, Leon 200 (MO).
Cuzco: Prov. Paucartambo, Kosnipata-Pilcopata, Var-
gas 14749 (GH). Quispicanchi, Fortaleza a Quince Mil,
Vargas 16539 (GH). Prov. La Convencion, ca. halfway
from Luisiana and Rio Apurimac to Camp 1, Dudley
10157 (GH). Madre de Dios: Prov. Manu, Sintuya, Alfaro
827 (MO). Prov. Tambopata, Dist. Tambopata, jet. Rio
La Torre and Rio Tambopata, Young 1 17 (MO, uc).
90. Thelypteris consobrina (Maxon & Morton) R.
Tryon, Rhodora 69: 5. 1967.
Dryopteris consobrina Maxon & Morton, Bull. Torrey
Bot. Club 65: 356. 1938. TYPE: Peru, Junin, near
La Merced, Killip & Smith 24087 (holotype, us!,
3 sheets; isotype, NY).
Stem creeping. Leaves mostly 100-250 cm long,
subdimorphic, the fertile with contracted pinnae.
Petiole up to 120 cm x 8-15 mm, stramineous to
tan, glabrous or glabrescent. Lamina 1 -pinnate,
with ca. 8-12 pairs of lateral pinnae and a similar,
large terminal one; pinnae mostly 20-35 x 4-7
cm, sessile or lowermost stalked to ca. 4 mm, the
base rounded or truncate, margin serrate or un-
cinate-serrate, apex acute to acuminate. Bud(s) ab-
sent or occasionally borne at base of proximal pin-
nae. Rachis glabrous or nearly so. Veins arising
from costae of fertile pinnae ca. 8-13 per 3 cm,
secondary veins straight (fertile) to slightly arcuate
(sterile), uniting at a generally obtuse angle and
giving rise to usually free excurrent veinlet; areoles
in rows of ca. 1 5-25 between costa and pinna mar-
gin. Indument abaxially lacking or of sparse tri-
chomes mostly 0.1-0.2 mm on costae, trichomes
more numerous on fertile costae, veins and lam-
inar tissue glabrous, lamina adaxially glabrous ex-
cept along costae. Sori oblong to linear, straight
or arcuate on the cross- veins, crowded at maturity,
lacking trichomes; sporangial stalks glabrous.
Lowland and montane forests, along streams and
edges of forests, 400-2000 m, Huanuco, Pasco,
Junin, and Ayacucho.
Known only from Peru.
Huanuco: Jingo Maria to Pucallpa, Ellenberg
3818 (GH). Along Rio Monzon, a few miles above
Tingo Maria, Hodge (GH, uc). Huacachi, estacion
near Mufia, Macbride4170 (F). Rio Huallaga, above
Rio Cayumba, Mexia 8315 (F, GH, MO, uc). Pasco:
Prov. Oxapampa, Gran Pajonal, vicinity of Che-
quitave, D. Smith 5261 (MO, uc). Junin: La Mer-
ced, Chanchamayo, C. Schunke (Rosenst. exsicc.
17) (A, uc). Ayacucho: Rio Apurimac Valley, near
Kimpitiriki, Killip & Smith 22945 (F).
9 1 . Thelypteris salzmannii (Fee) Morton, Los An-
geles County Mus. Contr. Sci. 35: 7. 1960.
Meniscium salzmannii Fee, (Mem. foug. 5) Gen. fil.
223. 1852. TYPE: Brazil, Bahia, Salzmann (iso-
types, B, c, NY).
Dryopteris salzmannii (Fee) Maxon & Morton, Bull.
Torrey Bot. Club 65: 357. 1938.
Stem short-creeping to suberect. Leaves mostly
100-180 cm long, subdimorphic, the fertile longer
and with contracted pinnae. Petiole up to 110 cm
x 8-15 mm, stramineous to tan, glabrescent to
moderately pubescent. Lamina 1 -pinnate, with ca.
10-22 pairs of progressively shorter lateral pinnae
and a similar, generally small terminal one; pinnae
mostly 12-22 x i.5_3(-4) cm, often strongly as-
cending ca. 45° from rachis, sessile or lowermost
stalked to ca. 2 mm, the base rounded or truncate,
margin entire to crenulate, apex acute to acumi-
nate. Bud(s) often borne at base of proximal pin-
nae. Rachis glabrescent or pubescent. Veins arising
from costae of fertile pinnae ca. 12-18 per 3 cm,
secondary veins generally arcuate or subsigmoid,
uniting at an obtuse to generally acute angle and
giving rise to usually free excurrent veinlet; areoles
in rows of ca. 8-16 between costa and pinna mar-
gin. Indument abaxially of sparse to moderately
dense trichomes mostly 0.1-0.3 mm on costae,
trichomes more numerous on fertile costae, veins
and laminar tissue glabrous, lamina adaxially gla-
brous except along costae. Sori oblong, generally
arcuate on the cross- veins, crowded and often be-
coming confluent at maturity with blades appear-
ing acrostichoid, lacking trichomes; sporangial
stalks glabrous.
Lowland and montane forests, along streams,
swamps, and thickets, 450-1200 m, Amazonas,
San Martin, Huanuco, Pasco, Cuzco, and Madre
de Dios.
Venezuela and Colombia to Bolivia; Brazil.
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
73
Amazonas: Prov. Bagua, valley of Rio Maranon above
Cascadas de Mayasi near Campamento STte. [sic] Mon-
tenegro, Wurdack 1829 (GH, USM). San Martin: Zepe-
lacio, near Moyobamba, Klug 3354 (F, GH, MO). Huanu-
co: Pampayacu, Hacienda at mouth of Rio Chinchao,
Macbride 5046 (F, GH, us). Pasco: Outskirts of Pozuzo,
near Rio Pozuzo, Gentry et al. 40089 (MO, uc). Prov.
Oxapampa, Gran Pajonal, via Chequitavo, D. Smith 5282
(MO, uc). Cuzco: Tumbuimayo-Quispicanchi, Vargas
11788 (GH). Madre de Dies: Prov. Manu, Hda. Defenza,
Vargas 15205 (GH).
92. Thelypteris lancea A. R. Sm., sp. nov.
Ex affinitate T. angustifolii (Willd.) Proctor et speci-
erum affinium subg. Meniscii sed distinguenda trichoma-
tibus patentibus rectis ad costas abaxialiter, pinnis steri-
libus usque ad 3.5 cm latis, pinnis proximalibus sessilibus
vel brevistipitatis usque ad 2 mm basi rotundatis vel
truncatis.
Stem creeping. Leaves ca. 100 cm long, subdi-
morphic, the fertile with contracted pinnae. Pet-
iole up to 60 cm x 8 mm, tan to brown, glabrescent
to moderately pubescent. Lamina 1 -pinnate, with
ca. 10-16 pairs of lateral pinnae and a similar
terminal one; sterile pinnae up to ca. 22 x 3.5 cm,
fertile to ca. 15 x 1.5 cm, sessile or lowermost
stalked to ca. 2 mm, the base rounded or truncate,
margin entire (sterile) to crenulate (fertile), apex
acuminate. Bud(s) not seen. Rachis glabrescent or
pubescent. Veins arising from costae of fertile pin-
nae ca. 1 3 per 3 cm, secondary veins straight (fer-
tile) to arcuate or subsigmoid (sterile), uniting at
an obtuse (fertile) to acute (sterile) angle and giving
rise to usually free excurrent veinlet; areoles in
rows of ca. 9-15 between costa and pinna margin.
Indument abaxially of moderately dense, patent
trichomes mostly 0.1-0.2 mm on costae, tri-
chomes more numerous on fertile costae, veins
and laminar tissue glabrous, lamina glabrous
adaxially except along costae. Sori arcuate on the
cross-veins, crowded and subconfluent at matu-
rity, lacking trichomes; sporangial stalks glabrous.
TYPE— Peru, Pasco, Prov. Oxapampa, Palcazu,
Rio Alto Iscozacin, Ozuz to Rio Lobo, R. Foster
& d'Achille 10061 (holotype, F!; isotype, USM!).
Along forested trail, 400-500 m, Pasco.
Peru; Bolivia.
This is most similar and probably most closely
related to T. angustifolia, from which it differs in
the spreading (vs. curved and appressed) tri-
chomes on the abaxial costae, the broader sterile
pinnae, and the lanceate pinnae that are not nar-
rowly cuneate at the base. A duplicate of the para-
type cited below was determined as T. serrata by
Maxon and Morton (1938), but the uc specimen
resembles that species only remotely.
Paratype: Bolivia, Dept. Santa Cruz, Prov. Sara,
Rio Japacani, 400 m, Steinbach 7500 (uc!).
93. Thelypteris angustifolia (Willd.) Proctor, Bull.
Inst. Jamaica, Sci. Ser. 5: 57. 1953.
Polypodium salicifolium Vahl, Eclog. amer. 3:51. 1 807.
(not Thelypteris salidfolia (Hooker) Reed, 1968).
TYPE: Montserrat, Ryan (holotype, c).
Meniscium angustifolium Willd., Sp. pi. ed. 4. 5: 133.
1810. TYPE: Venezuela, Caracas, Bredemeyer
(holotype, B, Herb. Willd. 1 95 74; microfiche, uc).
Dryopteris angustifolia (Willd.) Urban, Symb. antill.
4: 21. 1903.
Stem short- to long-creeping. Leaves ca. 30-
100(-130) cm long, subdimorphic, the fertile with
contracted pinnae. Petiole 15-50(-85) cm x 3-10
mm, tan to stramineous, glabrescent to moder-
ately pubescent. Lamina 1 -pinnate, with (6-)8-20
pairs of lateral pinnae and a similar terminal one;
pinnae(5-)10-15(-18) x (0.5-)l-2(-2.5) cm, ses-
sile or lowermost stalked to ca. 5 mm, the base
cuneate, margin entire (sterile) to crenulate (fer-
tile), apex acuminate. Bud(s) not seen. Rachis usu-
ally pubescent with falcate or crispate trichomes.
Veins arising from costae of fertile pinnae ca. 11-
18 per 3 cm, secondary veins straight (fertile) to
arcuate or subsigmoid (sterile), uniting at an ob-
tuse to acute angle and giving rise to usually free
excurrent veinlet; areoles in rows of ca. 4-10(-13)
between costa and pinna margin. Indument abax-
ially of sparse to usually moderately dense, falcate-
ascending to crispate trichomes mostly 0. 1-0.2 mm
on costae, especially of fertile blades, veins and
laminar tissue glabrous, lamina glabrous adaxially
except along costae. Sori arcuate on the cross- veins,
crowded and subconfluent at maturity, lacking tri-
chomes; sporangial stalks glabrous.
In lowland and montane rain forests, along
streams and on wet rocks, possibly sometimes a
rheophyte, 200-1200 m, Amazonas, San Martin,
Loreto, Huanuco, Pasco, Junin, Ucayali, and Cuz-
co.
Antilles; southern Mexico to Panama; Colom-
bia and Venezuela to Bolivia; southern Brazil; Par-
aguay.
Amazonas: Prov. Bagua, Chiriaco to Puente Venezue-
la, 43 km NE of Chiriaco, Barbour 4399 (MO, USM). San
Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo,
74
FIELDIANA: BOTANY
Puerto Pizana, Schunke 4665 (F, GH, MO, USM). Chazuta,
Rio Huallaga, King 4035 (F, GH, MO, uc). Lamas, Km
47.9 of Tarapoto-Yurimaguas road, Knapp & Mallet 8473
(F, MO). Tarapoto, LI. Williams 6136 (F). Loreto: Above
Pongo de Manseriche, Rio Santiago, Mexia 6167 (BH, F,
GH, MO, uc, z). Huanuco: Prov. Huanuco, Tingo Maria,
Tryon & Tryon 5338 (GH, USM). Pasco: Prov. Oxapampa,
Gran Pajonal, vicinity of Chequitavo, D. Smith 5311 (F,
MO, uc). Junin: Rio Paucartambo Valley, near Perene
Bridge, Killip & Smith 25260 (F, us). Ucayali: Coronel
Portillo, Pucallpa-Lima Hwy, Km 85 (as Loreto),
McDaniel 13943 (GH, USM). Cuzco: Prov. Paucartambo,
Pilcopata, Vargas 11 640 (an).
94. Thelypteris ensiformis (C. Chr.) R. Tryon,
Rhodora 69: 5. 1967.
Dryopteris ensiformis C. Chr., Kongel. Danske Vi-
densk. Selsk. Skr., Naturvidensk. Afd., ser. 7, 10:
269, / 46. 1913. TYPE: Costa Rica, La Palma,
Tonduz 12533 (holotype, P; isotype, us!).
Stem creeping. Leaves ca. 200-300 cm long,
monomorphic. Petiole ca. 130-1 70 cm x 1-2 cm,
tan to stramineous, glabrous. Lamina 1 -pinnate,
with up to ca. 10 pairs of lateral pinnae and a
similar terminal one; pinnae to 25-40 x 4-6 cm,
sessile, the base truncate to rounded, margin entire
to crenulate, apex acute. Bud(s) not seen. Rachis
glabrous. Veins arising from costae of fertile pin-
nae ca. 10 per 3 cm, secondary veins straight or
slightly curved, uniting at an obtuse angle and giv-
ing rise to usually free excurrent veinlet; areoles
in rows of ca. 1 2-1 5 between costa and pinna mar-
gin, about as long as broad. Indument abaxially
lacking, veins and laminar tissue glabrous, lamina
adaxially glabrous except along costae. Sori ar-
cuate on the cross- veins, distinct at maturity, lack-
ing trichomes; sporangial stalks glabrous.
In montane rain forest, 1400-1700 m, Junin.
Costa Rica; Venezuela; Ecuador; Peru.
This rare species has also recently been found
in Ecuador: Napo, Foster 85- 137 '-A (uc).
Junin: Schunke Hacienda, above San Ramon, Killip
& Smith 24651 (F, NY, us).
95. Thelypteris falcata (Liebm.) R. Tryon, Rho-
dora 69: 6. 1967.
Afeniscium falcatum Liebm., Kongel. Danske Vi-
densk. Selsk. Skr., Naturvidensk. Afd., V, 1: 183.
1849. LECTOTYPE (chosen by Smith, Fl. Chia-
pas 2: 234. 1981): Mexico, Oaxaca, Distr. Chi-
nantla, Liebmann Fl. Mex. 680 (c!; isotypes, K!,
us!).
Meniscium jurgensenii Fee, (Mem. foug. 5) Gen. fil.
223. 1852. TYPE: Mexico, Jurgensen 917 (ho-
lotype, not found at P).
Dryopteris falcata (Liebm.) C. Chr., Kongel. Danske
Vidensk. Selsk. Skr., Naturvidensk. Afd., ser. 7,
10: 270. 1913, not Kuntze, 1891.
Dryopteris jurgensenii (Fee) Maxon & Morton, Bull.
Torrey Bot. Club 65: 360. 1938.
Stem creeping to suberect. Leaves mostly ca.
100-250 cm long, monomorphic. Petiole up to ca.
130 cm x 1 cm, tan to stramineous, glabrous.
Lamina 1 -pinnate, with mostly 8-12 pairs of lat-
eral pinnae and a similar terminal one; pinnae to
ca. 30(-40) x 3-5 cm, the proximal ones stalked
to 5-30 mm, the base cuneate, margin entire to
crenulate, apex acuminate. Bud(s) not seen. Rachis
glabrous or sparsely pubescent. Veins arising from
costae of fertile pinnae ca. 8-12 per 3 cm, sec-
ondary veins straight or slightly subsigmoid, unit-
ing at an obtuse angle and giving rise to an ex-
current veinlet that is free or often completely
bisects the areole into two subequal rhomboid
halves; areoles in rows of mostly 10-15 between
costa and pinna margin, about as long as broad.
Indument abaxially of sparse trichomes 2-3 mm
long, veins and laminar tissue glabrous, lamina
glabrous adaxially except along costae. Sori oblong
to arcuate on the cross- veins, distinct at maturity,
lacking trichomes; sporangial stalks glabrous.
In montane rain forests, 800-1600 m, Pasco,
Junin, and Cuzco.
Cuba; southern Mexico to Panama; Colombia
to Bolivia.
Pasco: Pichis trail, Yapas (as Junin), Killip & Smith
25570 (NY, us). Junin: E of Quimiri Bridge, near La
Merced, Killip & Smith 23890 (F, NY, us). Cuzco: Prov.
La Convention, Abra de Ichiquiato, Vargas 14486 (GH).
96. Thelypteris lingulata (C. Chr.) Morton, Contr.
U.S. Natl. Herb. 38: 43. 1967.
Dryopteris lingulata C. Chr., Kongel. Danske Vidensk.
Selsk. Skr., Naturvidensk. Afd., ser. 7, 10: 271.
1913. TYPE: Costa Rica, vallee du Rio Hondo,
pres Madre de Dios, Pittier 10349 (holotype, P).
Stem creeping. Leaves mostly ca. 150-200 cm
long, monomorphic. Petiole up to ca. 110 cm x
1 cm, tan to stramineous, glabrous. Lamina
1 -pinnate, with mostly 3-5 pairs of elliptic lateral
pinnae and a similar terminal one; pinnae 20-35
x 5-10 cm, the proximal ones stalked to l-2(-4)
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
75
i, the base cuneate, margin entire to crenate,
ex acminate. Bud(s) not seen. Rachis glabrous.
;ins arising from costae of fertile pinnae ca. 4-
per 3 cm, secondary veins straight (fertile) to
bsigmoid (sterile), uniting at an obtuse angle and
/ing rise to an excurrent usually free veinlet;
soles of fertile pinnae 2-3 times broader than
ig, in rows of mostly 1 7-22 between costa and
nna margin. Indument abaxially lacking or of
arse trichomes 0.1 mm on costae, veins and
•ninar tissue glabrous, lamina adaxially gla-
ous. Sori oblong to arcuate on the cross- veins,
often biseriate between main lateral veins and
z sori round to oblong, in either case not con-
ent over the lamina at maturity, lacking tri-
omes; sporangial stalks glabrous.
Lowland rain forests, 100-500 m, Loreto and
izco.
Nicaragua to Panama; Colombia to Peru.
Loreto: San Antonio, on Rio Itaya, Killip & Smith
378 (NY, us). Cuzco: Prov. Quispicanchi, entre Machi-
i e Inambari, Vargas 16459 (GH).
. Thelypteris andreana (Sodiro) Morton, Contr.
U.S. Natl. Herb. 38: 50. 1967.
Meniscium andreanum Sodiro, Recens. crypt, vase.
Quit. 71.1883. TYPE: Ecuador, Rio Toachi, near
Santo Domingo, Sodiro (not located).
Dryopteris andreana (Sodiro) C. Chr., Index fil. 252.
1905.
Stem short-creeping. Leaves mostly ca. 1 00-250
i long, monomorphic or often subdimorphic.
dole up to ca. 100 cm x 5-10 mm, tan to stra-
neous, glabrous. Lamina 1 -pinnate, with mostly
1 5 pairs of elliptic lateral pinnae and a similar
minal one; pinnae mostly 20-30(-40) x 4-7
i, proximal ones sessile or nearly so, the base
imded to truncate, margin entire to crenulate,
ex acuminate. Bud(s) not seen. Rachis glabrous.
;ins arising from costae of fertile pinnae ca. 6-
per 3 cm, secondary veins nearly straight, unit-
l at an obtuse angle and giving rise to a usually
e excurrent veinlet; areoles of fertile pinnae about
long as broad, in rows of mostly 13-20 between
sta and pinna margin. Indument abaxially usu-
y lacking, lamina adaxially glabrous. Sori ob-
ig to arcuate on the cross-veins, infrequently
seriate between main lateral veins and the sori
round to oblong, never completely confluent over
the lamina at maturity, lacking trichomes; recep-
tacle with yellow to orange, opaque, tubular glands;
sporangial stalks glabrous.
Montane rain forests, 900 m, San Martin and
Loreto.
Nicaragua to Panama; Colombia to Peru.
San Martin: Prov. Lamas, Alonso de Alvarado, San
Juan de Pacayzapa, al este del Puente (Carretera a Moyo-
bamba), Schunke V. 5943 (F). Loreto: Varadero de Ma-
zan from Rio Amazonas to Rio Napo, Croat 195 14 A
(MO).
98. Thelypteris arcana (Maxon & Morton) Mor-
ton, Contr. U.S. Natl. Herb. 38: 42. 1967.
Dryopteris arcana Maxon & Morton, Bull. Torrey Bot.
Club 65: 352, t. 11. 1938. TYPE: Ecuador, Napo,
trail from Tena to Napo, Mexia 7174 (holotype,
us!; isotypes, F!, GH!, uc!, us!).
Stem short-creeping. Leaves 70-100 cm long,
subdimorphic. Petiole up to ca. 60 cm x 6 mm,
tan to stramineous, glabrous. Lamina 1 -pinnate,
with mostly 2-5 pairs of elliptic lateral pinnae and
a similar terminal one; pinnae mostly 10-20 x
2.5-5.5 cm (fertile 1.5-2.5 cm wide), the proximal
ones stalked 3-10 mm, the base cuneate, margin
entire to crenulate, apex acuminate. Bud(s) not
seen. Rachis glabrous. Veins arising from costae
of fertile pinnae ca. 13-18 per 3 cm, secondary
veins nearly straight (sterile subsigmoid), uniting
at an obtuse angle and giving rise to an excurrent
usually free veinlet; areoles of fertile pinnae about
as long as broad, in rows of mostly 7-10 between
costa and pinna margin. Indument abaxially lack-
ing or of scattered orangish tubular glands along
veins, lamina adaxially glabrous. Sori oblong to
arcuate on the cross- veins, not confluent over the
lamina at maturity; receptacle with orangish,
stalked, tubular glands; sporangial stalks glabrous.
Lowland rain forests, 100-850 m, Loreto and
Huanuco.
Ecuador to Bolivia.
Loreto: Varadero de Mazan from Rio Amazonas to
Rio Napo, Croat 19514 (MO, uc). Huanuco: Prov. Pachi-
tea, Dist. Puerto Inca, ca. 14 km from a point across Rio
Pachitea from Puerto Inca, D. Smith 1274 (MO).
FIELDIANA: BOTANY
Colombia
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
Tumbes
Piura
Lambayeque
Cajamarca
Amazonas
La Libertad
San Martin
Loreto
Ancash
Huanuco
Lima
Pasco
Junih
Ucayali
lea
Huancavelica
Ayacucho
ApurTfnac
Cuzco
Madre de Dios
Arequipa
Puno
Moquegua
Tacna
Chile
DEPARTMENTS OF PERU
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
77
Index to Names
Accepted names are in roman type, synonyms are in italics, and new names are in boldface. A page
number is provided for the principal place, or the only place, where the name occurs.
Acrostichum
fendleri 68
thelypteris 6
Amauropelta 9
breutelii 9
cheilanthoides 34
concinna 29
deflexa 30
diplazioides 14
oligocarpa 15
opposita 32
pilosula 19
rivulorum 32
rudis 26
thomsonii 22
Amphineuron 39
opulentum 43
Aspidium
abruption 46
atrorubens 24
biolleyi 62
cheilanthoides 34
coarctatum 32
conterminum 32
extensum 43
gongylodes 40
hispidulum 4 1
incanum 52
macrourum 44
navarrense 15
opulentum 43
pilosulum 19
pusillum 17
scalare 15
semihastatum 57
sprengelii 3 1
stipulare 44
uliginosum 3
Bolbitis 69
macrophylla 68
Ceterach
aspidioides 13
Christella 39
dentata 42
hispidula 4 1
parasitica 39
Ctenitis 3
Cyclosorus 39
dentatus 42
gongylodes 40
Dryopteris 2
subg. Steiropteris 46
anceps 68
ancyriothrix 63
andreana 76
angustifolia 74
arcana 76
aspidioides 13
var. subhastata 13
assurgens 30
asterothrix 62
atrorubens 24
balbisii 3 1
bangii 42
biformata 60
boqueronensis 26
brachyodus 49
brachypus 18
brausei 28
canadasii 2 1
caucaensis 28
cheilanthoides 34
var. resinosofoetida 34
chrysodioides
var. goyazensis 69
clypeata 65
coarctata 32
columbiana 15
comosa 49
concinna 29
consobrina 73
contermina 32
corazonensis 27
decussata 49
<fe/kxa 30
deltoidea 46
densa 34
densiloba 52
dentata 42
desvauxii
f. glandulosa 1 1
diplazioides 14
dumetorum 21
eggersii 59
engelii 26
ensiformis 75
euchlora 25
extensa 43
falcata 75
/wrva 19
gigantea 68
glandulosa
var. brachyodus 49
glandulosolanosa 20
incana 52
jamesonii 56
jurgensenii 75
juruensis 6 1
laevigata 24
lechleri 69
leprieurii 5 1
leucothrix 32
limaensis 20
lindigii 30
lingulata 75
linkiana 14
lomatosora 15
lugubriformis 58
macbridei 19
macrophylla 68
macrotis 57
var. nephrodioides 56
mapiriensis 49
megalodus 64
mercurii 3 1
m///« 28
mollis 42
multiformis 35
nitens 35
oligocarpa 15
oligophylla
var. aequatorialis 45
var. kunzeana 46
pachyrhachis
var. bogotensis 3 1
parasitica
var. glanduligera 4 1
patens 44
pavoniana 23
var. contracta 23
permollis 70
peruviana 23
phacelothrix 22
pilosohispida 27
pilosula 19
poiteana 65
ptarmiciformis 17
pteroidea 25
pusilla 17
pyramidata 58
quadrangularis 4 1
resinosofoetida 34
retrorsa 21
rimbachii 19
rivulorum 32
ruiziana 35
salzmannii 73
scalaris 16
sellensis 3 1
semihastata 51
serrata 71
simplicifrons 68
sprengelii 3 1
stuebelii 22
subandina 20
supina 26
supralineata 52
tetragona 64
thelypteris 6
78
FIELDIANA: BOTANY
thomsonii 22
tremula 9
tristis 6 1
uliginosa 3
valdepilosa 50
warmingii 56
Glaphyropteris 48
decussata 49
Goniopteris 52
biolleyi 62
crenata 52
eggersii 59
juruensis 6 1
pennata 64
poiteana 65
pyramidal a 58
tetragona 64
/rw/w 6 1
Grammitis
linkiana 14
Gymnogramma
diplazioides 14
polypodioides 14
Lastrea
cheilanthoides 34
mfms 35
poeppigiana 57
poiteana 65
scabriuscula 44
Lastreopsis 3
Macrothelypteris 3
torresiana 3
Megalastrum 3
Meniscium 66
andreanum 76
angustifolium 74
arborescens 70
falcatum 75
giganteum 68
guyanense 68
jurgensenii 75
macrophyllum 68
salzmannii 73
serratum 71
Nephrodium
"carazanense" 21
brachypus 18
canadasii 21
caucaense 28
conspersum 42
corazonense 27
crassipes 3 1
deflexum 30
eggersii 59
gardnerianum 52
jamesonii 56
kunzeanum 46
7 69
leprieurii 5 1
longipilosum 14
macradenium 21
macrotis 57
nemorale 62
pilosohispidum 27
quadrangulare 4 1
resinosofoetidum 34
retrorsum 27
schizotis 44
supinum 26
valdepilosum 50
Phegopteris
laevigata 24
membranacea 69
mo//w 70
Polypodium
abruption 58
balbisii 3 1
brachyodus 49
concinnum 29
crenatwn 65
cross ii 23
decussatum 49
dentatwn 41
euchlorum 25
megalodus 64
35
oligocarpum 15
oppositum 32
patens 44
pavonianum 23
pennatum 64
pteroideum 25
reticulatum 66
rivulorum 32
rufum 20
ruizianum 35
salicifolium 74
tetragonum 64
thomsonii 22
tottum 40
Polystichum
torresianum 3
Pronephrium 66
Pteris
interrupta 40
Steiropteris 46
gardneriana 52
incana 52
leprieurii 5 1
valdepilosa 50
Thelypteridaceae 2
Thelypteris 5
sect. Blennocaulon 35
sect. Glaphyropteris 46
sect. Lepidoneuron 9
sect. Macrothelypteris 3
sect. Phacelothrix 36
sect. Steiropteris 46
sect. Uncinella 29
subg. Amauropelta 9
subg. Cyclosorus 39
subg. Goniopteris 52
subg. Macrothelypteris 3
subg. Meniscium 66
subg. Steiropteris 46
subg. Thelypteris 6
abrupta 58
aequatorialis 45
ancyriothrix 63
andicola 16
andreana 76
angustifolia 74
arborea 24
arborescens 70
arcana 76
arenosa 33
argentina 21
arrecta 36
aspidioides 13
assurgens 30
atrorubens 24
balbisii 31
biformata 60
biolleyi 62
brachyodus 50
brachypus 18
brausei 28
canadasii 21
caucaensis 28
cheilanthoides 34
var. resinosofoetida 34
chrysodioides
var. goyazensis 69
clivalis 45
clypeata 65
coarctata 32
comosa 49
comptula 23
concinna 29
confluens 6
consobrina 73
conspersa 42
contermina 32
corazonensis 27
ctenitoides 37
curta 60
decussata 48
var. decussata 49
var. mapiriensis 49
var. velutina 48
deflexa 30
deltoidea 46
demissa 21
densa 34
dentata 4 1
depilata 45
diplazioides 14
dudleyi 33
dumetorum 27
TRYON & STOLZE: PTERIDOPHYTA OF PERU. III.
79
eggersii 59
enigmatica 16
ensiformis 75
erythrothrix 59
euchlora 25
extensa 43
exuta 37
falcata 75
frigida 18
funckii 18
furfuracea 34
furva 19
gardneriana 52
gigantea 68
glandulosa
var. brachyodus 49
var. glandulosa 50
var. longipilosa 50
glandulosolanosa 20
gongylodes 40
grandis
var. aequatorialis 45
var. grandis 46
var. kunzeana 46
var. pallescens 46
guyanensis 68
hispidula 4 1
var. inconstans 41
var. versicolor 4 1
hutchisonii 29
interrupta 40
invisa
var. kunzeana 46
jamesonii 56
juruensis 61
killipii 60
lacvigata 24
lancea 74
leoniae 18
leprieurii 51
var. glandifera 52
var. incana 52
var. leprieurii 5 1
leucothrix 32
limaensis 20
limbata 9
lindigii 30
lingulata 75
linkiana 14
lomatosora 15
longifolia 70
f. glandulosa 1 1
longipilosa 14
loretensis 30
lugubriformis 58
macbridei 19
macrophylla 68
macrotis 57
mapiriensis 49
maxoniana 71
megalodus 64
membranacea 69
mercurii 3 1
micula 33
millet 28
multiformis 35
navarrensis 15
nemoralis 62
nitens 35
oligocarpa 15
oligophlebia 3
opposita 32
opulenta 43
pachyrhachis 3 1
var. bogotensis 31
var. sprucei 3 1
palustris 6
parasitica 39
patens 43
var. dissimilis 44
var. patens 44
var. scabriuscula 44
var. smithiana 44
pavoniana 23
pennata 64
pennellii 50
peripae 60
peruviana 23
phacelothrix 22
pilosohispida 27
pilosula 19
pinnatifida 56
poiteana 65
proboscidea 36
ptarmiciformis 17
pteroidea 25
pusilla 17
quadrangularis 41
var. quadrangularis 4 1
var. repens 4 1
resinosofoetida 34
reticulata 66
retrorsa 27
rudis 25
rufa20
ruiziana 35
salicifolia 74
salzmannii 73
scalaris 15
schunkei 63
semihastata 57
serrata 71
sodiroi 62
sprengelii 3 1
subandina 20
supina 26
tetragona 64
thomsonii 22
torresiana 3
totta 40
tristis 60
tryonorum 62
urbanii 64
valdepilosa 50
Thelypteroideae 3
Trigonospora 2
80
FIELDIANA: BOTANY
Other Fieldiana: Botany Titles Available
FERNS AND FERN ALLIES OF GUATEMALA. Part I. Ophioglossaceae through Cyathe;<-
Robert G. Stolze. Fieldiana: B< 1976. 130 pages, 25 illus.
Publication 1246, $12.00
FERNS AND FERN ALLIES OF GUATEMALA. Part II. Polypodiaceae. By Robert C vvith
John T. Mickel and Alan R. Smith. Fieldiana n.s.. no. 6, 1981. 522 pages, 80 illus.
Publication 1317, $55.00
FERNS AND FERN ALLIES OF GUATEMALA. Part III. Marsileaceae, Salviniaceae, and the I
Allies (Including a Comprehensive Index to Parts I, II, and III). By Robert G. Stolze with Benjamin
011gaard and R. James Hickey. Fieldiana n.s.. no. 12, 1983. 91 pages, 10 illus.
Publication 1349, $10.00
PTERIDOPHYTA OF PERU. Part I. 1. Ophioglossaceae- 12. Cyatheaceae. By Rolla M. Tryon and
Robert G. Stolze. Fieldiana: Botany, n.s., no. 20, 1989. 145 pages, 24 illus., map.
Publication 1397, $27.00
PTERIDOPHYTA OF PERU. Part II. 13. Pteridaceac-15. Dennstaedtiaceae. By Rolla M. Tryon and
Robert G. Stolze. Fieldiana: Botany, n.s., no. 22, 1989. 128 pages, 30 illus., map.
Publication 1403, $23.00
PTERIDOPHYTA OF PERU. Part IV. 17. Dryopteridaceae. By Rolla M. Tryon and Robert G. Stolze,
with the collaboration of John T. Mickel and Robbin C. Mon-r 1991.
176 pages, 26 illus., map.
Publication 1424, $33.00
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