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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>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.
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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
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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).
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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