UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN NATURAL HIST. SURVEY FIELD Botany NEW SERIES NO. 20 UBMW PTERIDOPHYTA OF PERU Parti 1. Ophioglossaceae-12. Cyatheaceae Rolla M. Tryon Robert G. Stolzc January 31. 1989 Publication 1397 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY nation diana ' THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. FIELDIANA Botany NEW SERIES, NO. 20 PTERIDOPHYTA OF PERU Parti 1. Ophioglossaceae-12. Cyatheaceae Rolla M. Tryon Gray Herbarium Harvard University 22 Divinity Avenue Cambridge, Massachusetts 02138 Robert G. Stolze Collection Manager, Fern Herbarium Department of Botany Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 Accepted February 19, 1988 Published January 31, 1989 Publication 1397 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1989 Field Museum of Natural History Library of Congress Catalog Card Number: 88-80741 ISSN 00 15-0746 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents List of Illustrations INTRODUCTION 1 1 . ACKNOWLEDGMENTS 2 2. KEY TO FAMILIES OF PTERIDOPHYTA IN PERU 3 3. 1 . OPHIOGLOSSACEAE 5 4. Botrychium 6 5. Ophioglossum 8 2. MARATTIACEAE 13 6. Marattia 13 Danaea 15 7. 3. OSMUNDACEAE 20 8. Osmunda 21 9. 4. SCHIZAEACEAE 23 10. Anemia 24 Lygodium 30 11. Schizaea 33 5. GLEICHENIACEAE 37 12. Gleichenia 37 Dicranopteris 46 13. 6. HYMENOPHYLLACEAE 49 14. Hymenophyllum 50 15. Trichomanes 76 16. 7. LOXOMATACEAE 98 17. Loxsomopsis 99 8. PLAGIOGYRIACEAE 99 18. Plagiogyria 101 19. 9. DlCKSONIACEAE 101 Culcita 103 20. Dicksonia 105 21. 10. LOPHOSORIACEAE 107 22. Lophosoria 107 23. 11. METAXYACEAE 109 24. Metaxya 109 1 2. CYATHEACEAE Ill Sphaeropteris 112 Alsophila 116 Nephelea 118 Trichipteris 1 20 Cyathea 129 Cnemidaria 1 36 MAP OF PERU 140 INDEX TO NAMES . .141 Botrychium virginianum 7 Ophioglossum: O. palmatum; O. crotal- ophoroides; O. reticulatum 10 Marattia laevis 14 Danaea: D. moritziana; D. nodosa .... 16 Osmunda: O. regalis var. spectabilis; O. cinnamomea 22 Anemia: A. pastinacaria; A. phylli- tidis 26 Lygodium: L. venustum; L. volubile ... 31 Schizaea elegans 35 Gleichenia bifida 40 Dicranopteris: D. pectinata; D. flex- uosa 48 Hymenophyllum: H. fucoides var. fu- coides\ H. polyanthos; H. crispum 51 Trichomanes: T. radicans; T. pinnatum; T. hymenoides 77 Loxsomopsis pearcei 100 Plagiogyria semicordata 102 Culcita coniifolia 104 Dicksonia sellowiana 106 Lophosoria quadripinnata var. quadri- pinnata 108 Metaxya rostrata 110 Sphaeropteris: S. elongata; S. quindiu- ensis 113 Alsophila engelii 117 Nephelea cuspidata 119 Trichipteris pubescens 121 Cyathea caracasana var. boliviensis ... 1 34 Cnemidaria speciosa 137 111 PTERIDOPHYTA OF PERU Parti 1. Ophioglossaceae-12. Cyatheaceae Introduction The pteridophytes form a large and conspicuous element of the Peruvian flora, including 96 genera and about 1 ,000 species. Peru, which encompasses one of the world's richest biotas, occupies a central position in the Andes, and accordingly a knowl- edge of its flora is basic to understanding the plant life of the Andes as a whole. The Andes form a largely tropical mountain chain with an essentially north and south orientation, which is of special significance to its biogeography and the underlying processes of speciation. A portion of the Ferns of Peru was published by Rolla M. Tryon in 1964 (Contr. Gray Herb., vol. 194), with the encouragement and active sup- port of Theodore K. Just of Field Museum of Nat- ural History. A commitment was made at that time to complete the project after undertaking a major work on the genera of pteridophytes in America. The present work is a cooperative proj- ect of Harvard University Herbaria and Field Mu- seum of Natural History to present an account of all of the Pteridophyta of Peru. This is particularly important in relation to the Flora of Peru, initiated at Field Museum over 50 years ago, and thus far encompassing some 8,500 published pages. It is the only comprehensive treatment of Pacific coast- al, Andean, and Amazonian plants, and is closer to completion than any flora of the larger South American countries. The taxonomy is based not only on Peruvian materials, but includes an assessment of the An- dean species of a genus and, where appropriate, of all tropical American species. Most of the work will be accomplished by the authors, according to their knowledge of particular genera, but several treatments will be contributed by specialists cur- rently involved in monographic studies. The re- sults will be published in five parts, divided as equally as practicable, beginning with the present volume on the Ophioglossaceae through Cyathea- ceae. Studies are based primarily on the collections at Field Museum, Harvard University Herbaria, and United States National Herbarium (Smith- sonian Institution), but specimens at many other United States and European institutions have been examined. The extensive collections made under the current Flora of Peru project, a joint under- taking of Field Museum and Missouri Botanical Garden, are fully utilized. A close collaboration has also been established with Universidad Na- cional de Trujillo, Trujillo, Peru, and Museo de Historia Natural "Javier Prado" de Universidad Nacional Mayor de San Marcos, Lima, Peru, in- cluding loans of their specimens. 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 authentic material. Original drawings illustrate the diagnostic features of each genus and, where possible, some of the species. In addition, a number of plates published in Fieldiana: Botany, Ferns and fern allies of Gua- temala, have been used for species occurring in Peru. Voucher specimens cited in the legends are from Peru unless otherwise indicated. Prior to 1 944 the Department of Pasco was a part of Junin, and until recently Ucayali was a part of Loreto. An attempt has been made to ac- TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. count for these changes. This is not difficult when labels cite towns and provinces; however, on older specimens the labels often contain only sketchy data, making it impossible to determine in which part of Loreto or Junin a plant was collected. The map of Peru at the end of the text shows the de- partments and indicates the sequence of the col- lection citations. In the present work, Part I, the treatments of the Ophioglossaceae through the Plagiogyriaceae have been prepared by Robert G. Stolze, and those of the Dicksoniaceae through the Cyatheaceae, by Rolla M. Tryon. However, each author has re- viewed and edited the manuscript of the other, to the extent that the treatments are a joint effort. New names published here are indicated by boldface in the Index to Names. Collections have been cited from each department of Peru from which material has been seen, and further collec- tions are sometimes cited to include other her- baria. In general, all collections seen are cited for rare species, and a selection is cited for common species. Type collections, mentioned in the syn- onymy, are not repeated in the specimen citations, although they are included in the Peruvian range and ecology. 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. Appended to some of the generic treatments are portions of text labeled Comments. Herein are in- cluded species to be expected in Peru, names based on Peruvian material but of uncertain application, excluded species (erroneous reports that may have special significance), and cultivated species that are possibly adventive. Abbreviations of periodical publications gen- erally follow the system of Botanico-Periodicum- Huntianum (1968), and abbreviations of authors' names and of books generally follow TL-2 Taxo- nomic Literature by Stafleu and Cowan (1976 et seq.). Acknowledgments The authors are indebted to Dr. Michael Dillon (F), Dr. Abundio Sagastegui (HUT), and Blanca Leon (USM) for their invaluable assistance in pre- paring loans and arranging for the packing and shipment of specimens from the two Peruvian her- baria. The original drawings were contributed by Field Museum scientific illustrators, Zorica Da- bich and Clara L. Richardson, and by volunteer illustrators, Julia A. Liesse, Rosemarie Seitz, and Lisa M. Thorns. To these fine artists we wish to express our sincere appreciation, not only for their painstaking work, but also for their patience and understanding. Special thanks are extended to sev- eral persons who have been particularly helpful to us, either by providing special aid with the ex- amination of specimens at their institutions, or through stimulating discussions or correspon- dence concerning problems with various genera: Dr. David Lellinger (US), Dr. Alan R. Smith (UC), Dr. Henk van der Werff (MO), Dr. W. H. Wagner (MICH), and Dr. Paulo Windisch (Dept. Botanico, Rio Preto, Brazil). We also appreciate comments on the manuscript by several reviewers. 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: Botanischer Garten und Botanisches Museum, Berlin-Dahlem, Berlin (B); British Mu- seum (Natural History), London (BM); Jardin Bo- tanique National de Belgique, Meise (BR); Royal Botanic Garden, Edinburgh (E); Field Museum of Natural History, Chicago (F); Harvard University, Cambridge, Mass. — most Gray Herbarium (GH), some Arnold Arboretum (A); Institut fur Allge- meine Botanik und Botanischer Garten der Uni- versitat, Hamburg (HBG); Herbarium Truxil- lense, Universidad Nacional de Trujillo, Trujillo, Peru (HUT); Royal Botanic Gardens, Kew, En- gland (K); Rijksherbarium, Leiden, The Nether- lands (L); Botanische Staatssammlung, Munich (M); University of Michigan, Ann Arbor (MICH); Missouri Botanical Garden, St. Louis (MO); New York Botanical Garden, New York (NY); Museum National d'Histoire Naturelle, Paris (P); Univer- sity of California, Berkeley (UC); United States National Herbarium, Smithsonian Institution, Washington, D.C. (US); Museo de Historia Nat- ural "Javier Prado" de Universidad Nacional Mayor de San Marcos, Lima, Peru (USM); and Naturhistorisches Museum, Vienna (W). 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. FIELDIANA: BOTANY Key to Families of PTERIDOPHYTA in Peru The following key is restricted to the families that occur in South America and to the genera that are known from Peru or are likely to occur there. Several families are keyed out more than once in order to simplify the headings and to provide for more accurate identification. a. Two or more sporangia joined in a synangium b b. Synangium single in the axil of a bifid, bractlike leaf; aerial stem slender, green, dichotomously forked 24. Psilotaceae b. Two elongate synangia at the apex of a spike, or many synangia on the abaxial surface of a leaf; stem subterranean c c. Two elongate synangia at the apex of a spike; stem small 1 . Ophioglossaceae c. Many synangia on the abaxial surface of a leaf; stem stout to massive 2. Marattiaceae a. Sporangia separate d d. Sporangia borne on the inner side of a peltate sporangiophore in an apical strobilus; leaves much reduced, forming a sheath at the apex of each elongate, ridged interode Goint) of the stem 25. Equisetaceae d. Sporangia single in or near the axil of a leaf, or on 1 lobe of a 2-lobed leaf, or several to many sporangia borne on a leaf: at the margin, on the abaxial surface, on a specialized portion, or on a specialized leaf e e. A single sporangium borne in or near the axil of a leaf f f. Plants homosporous; leaves lacking a ligule 26. Lycopodiaceae f. Plants heterosporous, with megasporangia and microsporangia; leaves with a ligule g g. Leaves less than 1 cm long, borne along an elongate stem, fertile leaves in an apical strobilus 27. Selaginellaceae g. Leaves 2 cm long or usually longer, clustered at the apex of a compact to slightly elongate stem; all leaves usually fertile 28. Isoetaceae e. Several to many sporangia borne on a leaf, or a single sporangium borne on 1 lobe of a 2- lobed leaf h h. Plants heterosporous; the leaf bearing megasporangia and/or microsporangia, these enclosed in small, specialized structures i i. Plant with stem rooted in wet soil or underwater; leaves filiform, or with 4 leaflets at the apex of the petiole 22. Marsileaceae i. Plant floating on water; the floating leaves entire, oblong to suborbicular, or unequally 2-lobed with 1 lobe submerged 23. Salviniaceae h. Plants homosporous, the isomorphic sporangia exposed at the margin or on the abaxial surface of a leaf, or on a specialized portion of a leaf, or on a specialized leaf, sometimes enclosed prior to maturity within an indusium j j. Sporangia sessile or subsessile, or with a stalk of 4 or more rows of cells, annulus absent, or if present then apical, lateral, or oblique, not interrupted by the stalk k k. Annulus lacking, or sporangia with a poorly differentiated annulus 1 1. Sporangia lacking an annulus, borne on a specialized fertile branch of a leaf; spores lacking chlorophyll 1 . Ophioglossaceae 1. Sporangia with a poorly differentiated lateral annulus, borne on fertile pinnae; spores with chlorophyll (green) 3. Osmundaceae k. Sporangia with a well-differentiated apical to oblique annulus m m. Sporangia on the abaxial surface of a fertile portion of a leaf, distantly attached in a cluster or single on a vein, or in wholly fertile panicles . . 4. Schizaeaceae m. Sporangia contiguous on the receptacle of marginal or abaxial sori n n. Sporangia in marginal sori o o. Leaf very thin, 1-few cells thick, translucent, lacking stomata; stem pro- tostelic 6. Hymenophyllaceae o. Leaf thickened, with stomata; stem siphonostelic or dictyostelic p TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. p. Stem long-creeping, with scattered, short, stiff trichomes; receptacle elongate 7. Loxomataceae p. Stem massive, short-creeping to arborescent, with a dense mass of very long, soft trichomes; receptacle short or globose ... 9. Dicksoniaceae n. Sporangia in abaxial sori q q. Stem and leaves lacking evident indument, a mucilaginous secretion (flaky when dry) sometimes present; leaves strongly dimorphic 8. Plagiogyriaceae q. Stem and usually the leaves with evident trichomes and/or scales; leaves monomorphic to somewhat dimorphic r r. Stem slender, long-creeping, subterranean, freely branched; leaf usually partly pseudodichotomously branched with axillary arrested or dormant buds 5. Gleicheniaceae r. Stem stout to massive, more or less epigeous or arborescent, sparingly if at all branched; leaves wholly pinnately branched s s. Stem and petiole bearing scales, trichomes may be present or absent 12. Cyatheaceae s. Stem and petiole bearing only trichomes t t. Lamina 2-pinnate-pinnatifid to 3-pinnate-pinnatisect 10. Lophosoriaceae t. Lamina 1 -pinnate 11. Metaxyaceae j. Sporangia with a 1- to 3 -rowed stalk; the annulus vertical or nearly so, interrupted by the stalk u u. Petiole articulate at or very near the stem 21. Polypodiaceae u. Petiole continuous with the stem, or articulate well above the base of the petiole . . . v v. Stem scales clathrate w w. Spores monolete x x. Lamina entire; indusium lacking 14. Vittariaceae x. Lamina pinnatifid or more complex, or entire and the sori indusiate . . y y. Spores with chlorophyll (green), or lamina more or less dichotomous (furcate) 21. Polypodiaceae y. Spores lacking chlorophyll (not green) and lamina entire or pinnately branched z z. Sori abaxial; indusia, when present, circular to reniform, with a some- times narrow sinus 17. Dryopteridaceae z. Sori abaxial, indusiate, the indusia elongate, or sori nearly marginal 18. Aspleniaceae w. Spores trilete aa aa. Sori very long, near and parallel to the margin, or following the anasto- mosing veins, or in scattered groups 14. Vittariaceae aa. Sori roundish or united in a short line close to the costa 21. Polypodiaceae v. Stem scales not clathrate, or stem with trichomes bb bb. Spores trilete, or monolete and the sori marginal cc cc. Indusia absent dd dd. Stem with trichomes only 13. Pteridaceae dd. Stem with scales ee ee. Sori elongate along the veins, or sporangia acrostichoid 13. Pteridaceae ee. Sori roundish or united in a short line close to the costa 21. Polypodiaceae cc. Indusia present ff ff. Stem with trichomes, or with scales and an abaxial indusium present . 15. Dennstaedtiaceae FIELDIANA: BOTANY ff. Stem with scales and an abaxial indusium absent gg gg. Spores trilete 13. Pteridaceae gg. Spores monolete 15. Dennstaedtiaceae bb. Spores monolete and the sori or sporangia abaxial hh hh. Sori elongate, adjacent to and parallel to the segment axis 20. Blechnaceae hh. Sori roundish, or elongate, and most or all of them neither adjacent to nor parallel to the segment axis, or sporangia acrostichoid ii ii. Pinna stalks, if present, continuous with the rachis jj jj. Spores with chlorophyll (green); lamina or main veins more or less dichotomous (furcate) 21. Polypodiaceae jj . Spores lacking chlorophyll (not green) and lamina pinnately veined or branched kk kk. Petiole, at least basally, with 2 vascular bundles 11 11. Lamina with unicellular, acicular, or variously branched trichomes, or if glabrate then indusia absent 16. Thelypteridaceae 11. Lamina lacking trichomes, or with multicellular ones, or with minute, unicellular, blunt trichomes; indusia present 17. Dryopteridaceae kk. Petiole, at least basally, with 3 or more vascular bundles . . 17. Dryopteridaceae ii. Pinnae articulate to the rachis mm mm. Sporangia acrostichoid, or in indusiate sori and the lamina 2- pinnate, or 1 -pinnate and the pinnae with a large, basal, bas- iscopic auricle 17. Dryopteridaceae mm. Sporangia in indusiate sori, lamina 1 -pinnate, the pinnae cor- date, or the basal, basiscopic side less developed . 19. Davalliaceae Family 1: OPHIOGLOSSACEAE Ophioglossaceae C. Agardh, Aphor. bot. 113.1 822. TYPE: Ophioglossum L. Stem erect or prostrate, fleshy, lacking indu- ment, or pubescent-scaly at apex. Leaves erect or folded, not or scarcely circinate in vernation, en- tire to pinnate or subpalmate or dichotomous, commonly partially dimorphic, the sterile leaf with an expanded lamina, the fertile one with sporangia borne on a special branch (or branches) arising from the base of or below the sterile lamina. Veins free or anastomosing. Sporangia sessile or subses- sile, separate or laterally joined in a synangium, lacking an annulus. Spores trilete, lacking chlo- rophyll, from 1,500-15,000 in each sporangium. This is the most primitive of all living fern fam- ilies and contains three genera and 50 to 60 species. Botrychium and Ophioglossum occur in Peru and are virtually cosmopolitan, with representatives in subarctic as well as tropical regions. The Ophiog- lossaceae are distinctive in the fleshy, mycorrhizal roots lacking root hairs, the erect or folded (rather than circinate) vernation, and the massive spo- rangia with an extremely high spore capacity. Some of the species tend to have very wide, and often disjunct, distributions. References CLAUSEN, R. T. 1938. A monograph of the Ophioglossaceae. Mem. Torrey Bot. Club, 19(2): 1-177. UNDERWOOD, L. M., AND R. C. BENEDICT. 1909. Ophioglossaceae, in N. Amer. fl., 16: 3-13. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. Key to Genera of Ophioglossaceae a. Sterile lamina ternately decompound; veins free; sporangia borne in a panicle, separate from each other and not immersed in the fertile segment I. Botrychium a. Sterile lamina entire or palmately or digitately lobed; veins anastomosing; sporangia joined laterally in a synangial spike II. Ophioglossum I. Botrychium Botrychium Sw., J. Hot. (Schrader) 1800(2): 8, 110. 1802. TYPE: Botrychium lunaria (L.) Sw. (Osmunda lunaria L.). Figure 1. Plants terrestrial, very rarely epiphytic. Stem erect, short, lacking indument. Leaves solitary or few, 2-70 cm long, glabrous to pubescent, bud of the next leaf wholly or partly enclosed by the ex- panded stipular base of the current leaf. Sterile lamina sessile to long-stalked, pinnatifid to de- compound. Veins free. Fertile branch paniculate, racemose or spicate, arising at or below the base of the lamina. Sporangia protruding from the sur- face of the fertile branch, not immersed in the tissue. The genus contains about 25 species, with dis- tribution in temperate to tropical regions in both hemispheres, but is poorly represented in tropical America. Within the four subgenera of Botrychium only four species of Sceptridium have been found in the Neotropics, and but a single species of sub- genus Osmundopteris (B. virginianum). The pau- city of species in the Neotropics may be partly a natural phenomenon, but the problem may be also attributed to poor and sparse collections, as ferns of this genus often go undetected. Increased col- lecting efforts in North America have produced more species, and similar results may be attained through greater efforts in the tropics. Collectors should be aware that when a single plant is found in the field there are likely to be many more of that and other species nearby. References BUTTERS, F. K. 1917. Botrychium virginianum and its American varieties. Rhodora, 19: 207. MILDE, J. 1869. Botrychiorum monographia. Verh. Zool. Bot. Ges. Wien, 19: 55-190. Key to Species of Botrychium a. Fertile leaf with the sterile lamina long-stalked; leaf bud completely enclosed by the sheath of the current leaf 1 . B. schafTneri a. Fertile leaf with the sterile lamina sessile or nearly so; leaf bud partly exposed within the sheath of the current leaf 2. B. virginianum 1. Botrychium schafTneri Underw., Bull. Torrey Bot. Club 30: 51. 1903. TYPE: Mexico, San Luis Potosi, Schaffner 9 (holotype, NY; iso- type, K). Fertile plants 12-30 cm tall (sometimes to 50 cm outside Peru), the common stalk to 4 cm long, the leaf bud completely enclosed by the sheath of the current leaf. Sterile lamina of fertile leaf long- stalked (3-8 cm), fleshy, ternate, to 3-pinnate-pin- natifid, 3-10 cm long, 6-10 cm broad. Ultimate segments 0.3-0.7 cm long, mostly obtuse to sub- acute, separated by usually narrow to broadly U- shaped sinuses, the margins entire to crenate, very rarely serrate. Terrestrial, in high scrub forests and open mead- ows, 2350-3600 m, Amazonas, Cuzco, Madre de Dios(?). Mexico; Honduras south to Argentina. This is often difficult to distinguish from B. un- derwoodianum (Guatemala to Venezuela), which is usually a much larger fern with stalks of sterile laminae up to 28 cm long. However, Mexican plants of B. schqffneri seem to rival the latter in size, and the more reliable diagnostic features ap- pear to occur in the ultimate segments. Those of B. underwoodianum are 0.7-1 .4 cm long with mar- FIELDIANA: BOTANY FIG. 1. Botrychium virginianum: a, lamina with fertile branch; b, stem and part of petiole; c, part of fertile pinna, (a from Seller 865, El Salvador, GH; b-c from Lyonnet 896, Mexico, GH.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. gins typically serrate, whereas in B. schaffneri they are usually less than 0.7 cm long and entire to crenate (only rarely serrate). The taxonomy of the species complex requires detailed analysis. Amazonas: Chachapoyas, along Rio Ventilla, 1-2 km W of Molinopampa. Wurdack 1528 (us). Middle eastern Calla Calla slopes, km 41 1-416 of Leimebamba-Balsas road, Wurdack 1319 (us). Cuzco: Urubamba, Machu Picchu, slopes of Altillero, Peyton & Peyton 504 (MO). Dept. Unknown: "Pinasniocj, Panticalla Pass," Cook & Gilbert 1845 (us) (this may be on Cerro de Pantiacolla, Dept. Madre de Dios, near the Cuzco border). 2. Botrychium virginianum (L.) Sw., J. Bot. (Schrader) 1800(2): 111. 1802. Figure 1. Osmunda virginiana L., Sp. pi. 1064. 1753. TYPE: "America," Kalm (LINN 1244.3). Osmunda cicutaria Savigny in Lam., Encycl. 4: 650. 1797. TYPE: based on plants collected by Plu- mier in Hispaniola, Traite foug. Amer. 136, t. 159. 1705. Botrychium cicutarium (Savigny) Sw., Syn. fil. 171. 1806. Botrychium virginianum var. mexicanum Grev. & Hooker, Bot. Misc. 3: 223. 1833 (as B. virginicum beta mexicanum). SYNTYPES: Mexico, Vera- cruz, Chamisso s.n.; "Rigla," Veitch (both E? or K?). Fertile plants ca. 25-70 cm tall, the common stalk 1 9-40 cm long, the leaf bud partly exposed within the sheath of the current leaf. Sterile lamina of fertile leaf sessile or short-stalked (to 5 mm), membranous, subternate, to 3-pinnate-pinnatifid, to 20 cm long and 25 cm broad. Ultimate segments 0.5-0.9 cm long, joined by narrow, acute sinuses, elliptic to obovate, truncate, rounded or subacute, their apices variously dentate or lacerate. Forests and wooded canyons, 1800-2930 m, Cajamarca, Amazonas, Huanuco, Pasco, Junin, Cuzco. Essentially cosmopolitan. In the Neotropics from Mexico south to Bolivia and Brazil, and in the Greater Antilles. Botrychium virginianum has been treated by earlier authors as several species or as one species with several varieties or subspecies. Variants have been distinguished primarily by relative length of the fertile branch or degree of dissection of the sterile lamina. However, closer examination of more recent and numerous collections throughout the range of the species indicates that these are variable characters and not wholly correlated with geography. Proctor (Ferns of Jamaica, 1985) has observed that "... the almost complete intergra- dation of the alleged distinguishing characters ren- ders even varietal recognition difficult to uphold." We concur with this judgment. Cajamarca: Prov. Celendin, canyon of Rio Maranon above Balsas, Hutchison & Wright 5358 (uc). Amazonas: Prov. Chachapoyas, Cerros Calla Calla, 5 km above Lei- mebamba, Hutchison & Wright 4885 (uc). Huanuco: Mito, in canyon, Bryan 387 (F). Muna, in dry woods, Bryan 537 (F). Pasco: Prov. Oxapampa, 5 km SE of Oxapampa, D. Smith 2897 (MO). Junin: Huacapistana, dense forest, Killip & Smith 24308 (us). Cuzco: Prov. Calca, Vilcabamba, Vargas 3907 (us). II. Ophioglossum Ophioglossum L., Sp. pi. 1062. 1753. TYPE: Ophioglossum vulgatum L. Figure 2. Cheiroglossa Presl, Suppl. tent, pterid. 56. 1846. TYPE: Cheiroglossa palmata (L.) Presl = Ophioglossum palmatum L. Plants terrestrial or epiphytic. Stem erect, glo- bose or short-prostrate, small, lacking indument, or filiform-scaly at apex. Leaves solitary or several, 1-75 cm long (or to more than 1.5 m in the Old World O. pendulum), glabrous, the eroded stipular base of the expanded leaf not enclosing the bud of the next leaf. Sterile lamina sessile or short-stalked, entire, or palmately or digitately lobed. Veins anastomosing, often with free, included veinlets. Fertile branch or branches borne on or below the base of the sterile lamina. Sporangia joined lat- erally in a synangial spike. Ophioglossum is a genus of 25-30 species scat- tered in temperate to tropic regions around the world. Plants are often small and inconspicuous and are commonly concealed in grassy turf or her- baceous growth; consequently, current collection records probably do not represent true distribution patterns. Although six species are recognized here from Peru, several others might be expected in the future. Reference WAGNER, W. H., ET AL. 1984. Ophioglossum el- lipticum in Louisiana and the taxonomy of O. nudicaule. Castanea, 49: 99-1 10. FIELDIANA: BOTANY Key to Species of Ophioglossum a. Sterile lamina deeply and irregularly palmately lobed; fertile spikes several; plants epiphytic 1 . O. palmatum a. Sterile lamina entire; fertile spike solitary; plants terrestrial b b. Stem globose 5. O. crotalophoroides b. Stem cylindrical, sometimes swollen, but never globose c c. Sterile lamina stalked, the common stalk and stalks of fertile and sterile segments hypogeous and scarious 4. O. scariosum c. Sterile lamina sessile or nearly so, the common stalk and stalks of sterile and fertile segments mostly epigeous, not or scarcely scarious d d. Primary areoles each enclosing several smaller secondary areoles and/or some free veinlets; base of sterile lamina cordate, truncate or abruptly cuneate 2. O. reticulatum d. Primary areoles not enclosing secondary ones or free veinlets; base of sterile lamina narrowly cuneate to attenuate e e. Sterile lamina elliptic or oblong, 1.5-3 times as long as broad, cuneate at base; plants of higher elevations (over 2500 m) 3. O. nudicaule var. tenerum e. Sterile lamina linear-lanceolate to narrow-oblanceolate, (4-)4.5-8 times as long as broad, long-attenuate at base; plants of lower elevations (under 700 m) 6. O. lusitanicum ssp. coriaceum 1 . Ophioglossum palmatum 1 ... Sp. pi. 1063. 1 753. TYPE: based on illustration of plant from Haiti, Plumier, Traite foug. Amer., /. 163. 1705. Figure 2a. Cheiroglossa palmata (L.) Presl, Suppl. tent, pterid. 57. 1846. Plants epiphytic. Stem stout, elongate, abun- dantly provided at apex with tawny to orange scales which are hairlike for most of their length. Leaf (15-)20-75 cm long, with common stalk (5-) 10- 40 cm long. Sterile lamina deeply and irregularly palmately lobed, obdeltoid, narrowly cuneate at base, lacking a distinct midvein or pale median band of tissue. Veins distinct, the primary areoles large, often enclosing smaller secondary areoles and occasionally a few free veinlets. Fertile seg- ments (l-)3-l 2, borne on or below the base of the lamina, the fertile spikes 0.5-6 cm long, on stalks 0.2-2 cm long. Suberect or pendent from trunks or branches of trees, in cloud forests, elfin forests, and shaded ravines, 1000-2450 m, Amazonas, Huanuco, Pas- co, Cuzco. Southern Florida; West Indies; southern Mexico to Peru and Brazil; scattered in Old World tropics. Amazonas: Prov. Bagua, E of La Peca, Barbour 2512, 2860, 2940 (MO). Prov. Chachapoyas, along Rio Ventilla 1-2 km W of Molinopampa, Wurdack 1478 (F, GH, NY, uc, us). Huanuco: East of Tingo Maria (as San Martin) in deep ravine, Allard21580 (us). Pasco: Prov. Oxapam- pa, 1800 m, van der \Verff8352 (MO). Cuzco: Prov. La Convention, Tunquimayo, Idma, Vargas 10681 (MICH, MO). 2. Ophioglossum reticulatum L., Sp. pi. 1063. 1753. LECTOTYPE (designated by Proctor, Flora Lesser Antilles 2: 43. 1977): Plumier, Traite foug. Amer., /. 164, based on specimen from Haiti. Fig- ure 2c-d. Ophioglossum petiolatum Hooker, Exot. fl. 1: 56. 1823. TYPE: based on plants from West Indies sent to Liverpool Botanic Garden and cultivated there (not located). Ophioglossum peruvianum Presl, Suppl. tent, pterid. 52. 1845. TYPE: Peru, Poeppig(noi located). Plants terrestrial. Stem cylindrical, lacking in- dument. Leaf (5-) 10-40 cm long, common stalk mostly epigeous, 3-20 cm long. Sterile lamina membranaceous, 2-12 cm long, 1.5-5 cm broad, sessile or subsessile, ovate to nearly circular, apex obtuse to acute, base cordate to truncate or (oc- casionally) abruptly cuneate. Veins distinct to in- distinct, primary areoles enclosing a few smaller secondary areoles which are scarcely differentiated from the primary ones, free included veinlets sev- eral to numerous. Fertile segment solitary, borne at base of sterile lamina, on mature leaves some- what to greatly exceeding the lamina, fertile spike 1.4-4 cm long, on a stalk 3.5-14.5 cm long. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 3 mm 3cm 5mm FIG. 2. Ophioglossum palmatum: a, habit. Ophioglossum crotalophoroides: b, habit. Ophioglossum reticulatum: c, venation of sterile leaf; d, fertile spike, (a from Allard 21580, us; b from Vargas 3978, us; c-d from Tryon & Tryon 5419, F.) 10 FIELDIANA: BOTANY From high, rocky meadows to the open forests of middle elevations, and the grassy soil of coastal lomas, 400-3800 m, Amazonas south to Cuzco. Widely distributed in tropical America; Old World. Ophioglossum reticulatum is variable in the shape and base of the sterile lamina and in the frequency of secondary areoles and free, included veinlets. These characters intergrade freely with those of the often recognized O. petiolatum, so that it is impractical to separate the two as distinct taxa. Thus circumscribed, O. reticulatum is perhaps the most ubiquitous of any species in the genus. Amazonas: Prov. Bongara, Shilla. Young & Eisenberg 460 (MO). La Libertad: Prov. Pataz, Pumatambo, Puerta del Monte. Lope: & Sagdstegui 3438 (GH). San Martin: Monte Campana, near Tarapoto, Spruce 4709 (p, us). Lima: Lomas de Quilmana, Coronado 27 (us). Prov. Chancay, Lomas de Lachay, Coronado & Velarde 15 (uc, us); Tryon & Tryon 5419 (GH). Pasco: Prov. Oxapampa, Chontabamba, D. Smith & Brack 3056 (F, MO). Ayac- ucho: Aina, between Huanta and Rio Apurimac, Killip & Smith 22809 (GH, NY, us). Apurimac: Prov. Abancay, Cachora, Huillcayoc, Vargas 9096 (MICH, uc). Cuzco: Prov. Urubamba, Machu Picchu, Leon 452 (GH, USM); Tryon & Tryon 5400 (GH, us), 5419 (GH). Madre de Dios: Prov. Manu, Parque Nacional Manu, Foster et al. 11486 (F). and sedges, 2800-4000 m, Cajamarca, Lima, Cuz- co. Colombia; Peru; Brazil. In spite of Clausen's work (1938), considerable study is still needed to understand the variation within O. nudicaule. The Peruvian plants here identified cannot be clearly placed within any of the varieties delineated by Clausen; but, aside from the fact that their fertile segments do not rise too conspicuously beyond the sterile lamina, they seem to fit best under var. tenerum. Diagnostic features of O. nudicaule are so few and variable that it is often difficult to discern which are innate differ- ences and which are merely induced by rigorous habitat or attributed to degree of maturity of the individual plant. It is likely that this and other varieties accepted by Clausen are not supportable, as evidenced by the studies of Wagner et al. ( 1 984), which suggest that they "all should be subsumed under the single binomial, O. nudicaule." Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui et al. 6412 (GH, HUT). Lima: Prov. Huarochiri, Dist. de Mariatana, Cueva Mortero, Cerrateet al. 4812 (GH, USM). Cuzco: Prov. Acomayo, Quenco Grande, near Acomayo, Vargas 9752 (F, GH, K, ucp.p., us). Prov. Espinar, Yauri, Vargas 5619 (MICH). Prov. Urubamba, Vargas 14122 (GH). 3. Ophioglossum nudicaule L. f. var. tenerum (Mett.) Clausen, Mem. Torrey Bot. Club 19(2): 151. 1938. Ophioglossum ypanemense Mart., Icon. pi. crypt. 39, 1 30, /. 73. 1 834. TYPE: based on specimens from "Ypanema," Brazil. Ophioglossum tenerum Mett. in Prantl, Ber. Deutsch. Bot. Ges. 1: 352. 1883. TYPE: United States, Georgia, collector undesignated (holotype, B). Plants terrestrial. Stem cylindrical to somewhat swollen, lacking indument. Leaf 2.5-7 cm long, common stalk mostly epigeous, 0-4 cm long. Ster- ile lamina chartaceous to somewhat carnose, 0.8- 2.5 cm long, 0.3-1 .0 cm broad, 1.5-3 times as long as broad, sessile, elliptical to subovate, obtuse or subacute, base cuneate. Veins indistinct to ob- scure, areoles small, not enclosing secondary ar- eoles or (except rarely) free veinlets. Fertile seg- ment solitary, borne at base of sterile lamina, on mature leaves somewhat exceeding the lamina, fertile spike 0.5-1 .2 cm long, on a stalk 0.8-2. 5(-4) cm long. In high rocky meadows and slopes, with grasses 4. Ophioglossum scariosum Clausen, Mem. Tor- rey Bot. Club 19(2): 153. 1938. TYPE: Peru, Junin, vicinity of Oroya, Kalenborn 125 (ho- lotype, us!; isotypes, GH!, MO!, NY). Plants terrestrial. Stem broadly elongate, 4-6 mm thick, lacking indument. Leaf 3-6 cm long, with common stalk and bases of fertile and sterile stalks hypogeous and scarious, common stalk 0.3- 0.8 cm long. Sterile lamina 1-1.5 cm broad, ob- long-ovate to suborbicular, apex apiculate, at base truncate, reduced abruptly to an attenuate stalk 0.8-1.2 cm long. Veins indistinct to nearly ob- scure, primary areoles usually enclosing some sec- ondary ones as well as free veinlets. Fertile seg- ment solitary, borne at the base of the stalk of the sterile lamina, on mature leaves somewhat to greatly exceeding the lamina, fertile spike 0.6-1.2 cm long, on a stalk 1.3-4 cm long. In open, grassy areas, 2840-3850 m, Junin, Cuz- co, Puno. Peru and Bolivia. Leaves of this rare fern grow with proximal por- tions of leaves embedded in the soil. This is evi- TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 11 dent even in dried material, as the scarious com- mon stalk and proximal portions of the stalks of fertile and sterile segments contrast sharply with the dark color of the rest of the leaf. Cuzco: Prov. Urubamba, Vargas 9165 (MICH). Puno: Lake Titicaca, Bishop 1994 (us). 5. Ophioglossum crotalophoroides Walter, Flora Caroliniana 256. 1788. TYPE: Carolina, Walter (holotype, BM?). Figure 2b. Plants terrestrial. Stem globose, 4-12 mm in diameter, lacking indument. Leaf 3-15 cm long, with common stalk 0.5-4 cm long. Sterile lamina ovate to deltoid, acute or rarely obtuse, cordate or truncate and abruptly attenuate at base, often lon- gitudinally folded. Veins distinct or indistinct, pri- mary areoles not or rarely enclosing secondary ones or free veinlets. Fertile segment solitary, borne at the base of the sterile lamina; in mature leaves the fertile segments somewhat to greatly exceeding the lamina, fertile spike 0.5-1.5 cm long, on a stalk to 10 cm long. On moist, open, grassy slopes, 600-3900 m, Huanuco, Lima, Junin, Apurimac, Cuzco. Southern United States; Mexico to Honduras; Colombia and Venezuela south to Argentina and Chile. Among the diminutive species of Ophioglos- sum, a few plants are found with stems which are somewhat swollen. However, only in O. crotalo- phoroides is the stem so conspicuously globose. With this should probably be included O. opacum Carmichael, from Tristan da Cunha. It is said to differ from the former merely in the somewhat shorter fertile segment and slightly smaller globose stem. Huanuco: Prov. Huanuco, Huanuco-Tingo Maria road, from Bandera Blanca to Carpish Tunnel, Luteyn & Lu- teyn 5464 (NY). Lima: Prov. Lima, Lomas de Lurin, Ferreyra 9566 p.p. (USM). Junin: Huaytapata. near Ha- cienda Conocancha, Tiller 14 (USM). Apurimac: Prov. Abancay, Cachora, Vargas 9054 (MICH p.p., MO). Cuzco: Prov. Calca, Vilcabamba, Vargas 3978 (us). Prov. Can- as, slopes of San Andres de Checca, Vargas 11010 (F, uc). 6. Ophioglossum lusitanicum L. ssp. coriaceum (A. Cunn.) Clausen, Mem. Torrey Bot. Club 19(2): 161. 1938. Ophioglossum coriaceum A. Cunn., Companion Bot. Mag. 2: 361. 1836. TYPE: based on specimens from Matauri, New Zealand (not located). Plants terrestrial. Stem cylindrical to somewhat swollen, lacking indument. Leaf 4-8(-9) cm long, common stalk mostly epigeous, 0.5-2.5 cm long. Sterile lamina chartaceous to somewhat carnose, 1.5—4 cm long, 0.3-0.6 cm broad (ca. 4.5-8 times as long as broad), sessile, linear-lanceolate to nar- row-oblanceolate, apex acute or subacute, base narrowly cuneate to (typically) long-attenuate. Veins indistinct to obscure, areoles commonly long and narrow, not enclosing secondary areoles or free veinlets. Fertile segment solitary, borne at base of sterile lamina, on mature leaves somewhat to conspicuously exceeding the lamina, fertile spike 0.4-1.7 cm long, on a stalk 1-7 cm long. In soil pockets or on thin soil over rocks, in Peru possibly confined to the coastal lomas in the vi- cinity of Lima, 400-650 m. Peru; Bolivia; Chile; New Caledonia; Australia; New Zealand. In Peru, this is typically a very delicate plant, with long, narrow, sterile leaves and a fragile, fil- iform, fertile stalk, and has thus far been found only at low elevations. However, we have exam- ined some Bolivian plants of this subspecies, from elevations to 3000 m, which are more variable: sometimes more robust, with stouter fertile stalks, and with sterile laminae broader and less attenuate to base and apex. Subspecies lusitanicum, from Europe, Asia, and Africa, has obtuse sterile lam- inae with larger and fewer areoles. Subspecies cal- ifornicum (Prantl) Clausen (California & Mexico) has laminae larger and broader, with less attenuate bases than in ssp. coriaceum and, according to Clausen (1938), perhaps does not merit distinc- tion. As with the O. nudicaule complex (q.v.), far more study is needed to clarify the taxonomy, at both inter- and intraspecific levels. Lima: Prov. Chancay, Lomas de Lachay, Coronado & Velarde 16 (GH, uc, us); Tryon & Tryon 5414 (GH p.p., us). Lomas de Quilmana, Coronado 27 (uc, us p.p.). Lomas de Lurin. Ferreyra 9566 (GH, USM p.p.). Lomas de Atocongo, Grant 7509 (GH). Comments Ophioglossum ellipticum Hooker & Grev., Icon, fil. 1, t. 40A. 1831. TYPE: illustration, based 12 FIELDIANA: BOTANY on specimens collected in Demerara, "British Guiana," Parker. This perhaps may be expected in Peru, as its range is thus far reported from Guatemala to Costa Rica, and the Guianas southward to Brazil and (possibly) Bolivia. Although usually a smaller plant than Ophioglossum reticulatum, the aspect of the sterile lamina is similar in that the primary areoles contain a number of secondary areoles and usually some free veinlets. However, the secondary ar- eoles are much more numerous in O. ellipticum, and the walls are thinner and far less distinct than those of the primary ones. Additionally, laminae in O. ellipticum have a somewhat distinct midvein and a pale, median band of tissue which persists to within a short distance of the apex. It is most closely related to O. nudicaule, in fact, the plants heretofore recognized as O. ellipticum may be merely O. nudicaule with larger and more fully developed leaves (see Wagner et al., 1984). Family 2: MARATTIACEAE Marattiaceae Bercht. & J. S. Presl, Prir. rostlin 1 : 272. 1820. TYPE: Marattia Sw. Danaeaceae Agardh, Aphor. hot. 117. 1822. TYPE: Danaea Sm. Stem fleshy, stout to massive, creeping or erect, sometimes scaly. Leaves moderate in size to huge (3 m or more), circinate in vernation, 1 -pinnate to pinnately compound or rarely simple or pal- mate, monomorphic to somewhat dimorphic. Pet- iole with an expanded, stipular base, there swollen and sometimes subarticulate, frequently also with several swollen and darkened nodes throughout its length. Pinnae commonly borne in opposite pairs from swollen, darkened nodes on the rachis. Veins free or (in Christensenia) reticulate. Spo- rangia borne on the abaxial surface of leaves, with thickened walls, lacking an annulus, separate and contiguous or fused into indurated synangia and opening by terminal pores. Spores ellipsoidal and monolete or trilete, rugose or echinate, very nu- merous (1,500-1,700) in each sporangium. Of the seven genera (about 1 50 species) in Mar- attiaceae, two are represented in Peru: Danaea, confined to the Neotropics, and Marattia, pan- tropical. Next to Ophioglossaceae, it is considered to be the most primitive of the living fern families and can be easily recognized by the character of the sporangia being fused into thick, elongate, dou- ble-rowed synangia. References DEVRIESE, W. H.,ANDP. HARTING. 1853. Mon- ographic des Marattiacees, Leiden, Nether- lands. TRYON, R. M., AND A. F. TRYON. 1982. Mar- attiaceae, pp. 39-50, in Ferns and allied plants, Springer- Verlag, New York. UNDERWOOD, L. M. 1909. Marattiaceae, in N. Amer. fl. 16: 15-23. Key to Genera of Marattiaceae a. Leaves 1-3-pinnate or more, monomorphic; synangia scattered, borne near the ends of veins I. Marattia a. Leaves 1 -pinnate, somewhat dimorphic (fertile ones commonly longer and narrower and with smaller pinnae); synangia nearly covering the abaxial surface of fertile pinnae II. Danaea I. Marattia Marattia Sw., Prodr. 128. 1788. TYPE: M. alata Sw. Figure 3. Plants terrestrial. Leaves coarse, to ca. 3(-4) m long, 2-4-pinnate, often ternate in architecture, monomorphic. Lamina with expanded tissue es- sentially glabrous, but in some species sparsely to moderately scaly or with a few trichomes on the axes abaxially. Penultimate divisions with axes commonly broadly alate, the wings narrowing abruptly at the base of each segment. Veins free. Sporangia in two rows, opening by longitudinal slits, fused into bivalvate, ovoid synangia, which are borne near the ends of veins on a receptacle, TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 13 FIG. 3. Marattia laevis: a, portion of lamina; b, segment base, with synangia. (a from Macbride 4817, F, b from Haught 6145, Colombia, F.) 14 FIELDIANA: BOTANY sessile or (in M. laevis) on short, thickened stalks. Spores ellipsoid, monolete. This is a pantropical genus of about 40 species, commonly occurring in dense, wet forests. It is readily separated from other Neotropical genera by the large, decompound, subternate leaves with stipular bases, and the distinctive ovoid synangia. A single species is known in Peru. 1. Marat tia laevis Sm., Plantarum Icones Inedi- tae ... 2: /. 47. 1790. TYPE: as "Polypodium pedicellata, in argento pedunculat(o?), He Dominique (Dominican Republic, not Do- minica), Thiery" (holotype, LINN). Figure 3. Marattia alata sensu Raddi, PI. bras. nov. gen. 1 : 74, t. 83-84. 1825. Marattia Kaulfussii Hooker, Gen. fil., t. 26 (as to note). 1839. TYPE: Brazil, 1816, Cunningham s.n. (holotype, BM!; photo, F!). Eupodium Kaulfussii (Hooker) Hooker, Sec. cent, ferns, /. 95. 1860. Marattia alata Sw. var. laevis (Sm.) Farw., Amer. Midi. Naturalist 12: 308. 1931. Leaves 2—4 m long, commonly 3-pinnate-pin- natifid, at least as to base of lamina. Axes (at least minor ones) sparsely to moderately provided on abaxial side with filiform to lanceolate scales, these orange to light brown, flaccid, 0.5-3 mm long, often caducous. Costae of penultimate segments bearing on the adaxial side 1 -several well-spaced, often appressed, awns, these 0.2-1 mm long (or on older leaves sometimes withering or broken away near their bases). Synangia 8-16(-18)-lo- cular, mature ones obviously stipitate, the stalks stout, to 0.7 mm long. Spores echinate. Occurring in dense forests, principally along the Cordilleras Central and Oriental, in Amazonas, Huanuco, Pasco, Junin, and Cuzco, 1400-2400 m. Costa Rica; Cuba; Hispaniola; Colombia; Ven- ezuela; Brazil; Ecuador; Peru; Bolivia. In various herbaria, M. laevis has been confused with M. alata, a species from the Greater Antilles. Although leaf outline is similar in each, M. laevis is easily distinguished by its stalked synangia and awned penultimate axes, these characters are unique in the genus, at least in New World species. There has been some confusion as to the identity of M. laevis, beginning with Smith's original de- scription, which mentioned nothing about the dis- tinctive stalked synangia and adaxial awns. Kaul- fuss (Enum. fil., p. 32, 1824) mentioned seeing some Brazilian specimens of M. laevis with "soros ... stipitati," upon which character Hooker (fol- lowing Smith's unpublished description) partially based the new species M. Kaulfussii (1839). Ob- viously Hooker had seen the type of M. laevis, and supposed it lacked the stalked synangia; therefore, he surmised that the Brazilian collections of Kaul- fuss represented a new species. Finally in 1936 Alston (J. Bot., p. 174) combined the two names, but failed to specify reasons for doing so. In turn- ing to the British Museum for clarification of the matter, this reply was received from Miss Jose- phine M. Camus: "I have examined the (type) specimen in the Linnaean Herbarium. It does in- deed have synangia with very elongate receptacles. The original label bears the name Polypodium ped- icellata in Linnaeus fil.'s hand. The 'awns' are pres- ent on the upper surface of the pinnule axes... ." Thus, there seems to be no question that the fern considered to be M. Kaulfussii for nearly 1 50 years is M. laevis. Amazonas: Prov. Bagua, Cordillera Colon SE of La Peca, Barbour 3900 (MO). Huanuco: Churubamba, Trail Puente Durand to Exito, Mt. Santo Toribio, Mexia 8250 (F, GH, MICH, NY, uc, us). Pasco: Cushi, Macbride 4817 (F, GH, us). Junin: Schunke Hacienda above San Ramon, Killip & Smith 24540 (NY, us). Pichis Trail, Enenas, Killip & Smith 25738 (GH, NY, us). Chanchamayo Valley, C. Schunke 671 (F). Cuzco: La Convention, above Knox's Cascade, Dudley 10619 (GH, MO). II. Danaea Danaea Sm., Mem. Acad. Roy. Sci. (Turin) 5: 420. 1793, nom. consent., not Allioni, 1785. TYPE: Danaea nodosa (L.) Sm. (Acrostichum nodos- um L.). Figure 4. Plants terrestrial, with stems creeping to decum- bent or erect. Leaves to 2 m long, 1 -pinnate (or simple in 2 species outside Peru), monomorphic to somewhat dimorphic (fertile ones commonly longer and narrower and with smaller pinnae). Pet- iole (in most species) nodose, scaly, bearing sti- pules at the base. Rachis nodose, nonalate to con- spicuously alate at least distally, glabrous or more often with appressed scales. Lamina with a distinct apical segment, or this replaced by a proliferous bud, provided with scales abaxially, those of the laminar tissue minute and scattered. Sterile pinnae opposite or subopposite, straight to subfalcate, ses- sile or short-stalked, entire and often undulate, or serrate near the apex. Veins free. Fertile pinnae similar to the sterile ones, but commonly reduced TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 15 FIG. 4. Danaea moritziana: a, habit. Danaea nodosa: b, apex of sterile lamina; c, portion of fertile pinna with synangia. (a from Tryon & Tryon 5278, F, b-c from Wurdack 1934, F.) 16 FIELDIANA: BOTANY in size, and with vegetative tissue nearly lacking. Sporangia nearly covering the abaxial surface, co- alescing into much elongated, indurated synangia and opening by terminal pores. Spores monolete, the surface prominently echinate. Danaea is confined to the Neotropics. With its nodose leaf axes and the distinctive synangia near- ly covering the abaxial surface of simple, usually opposite, pinnae, it should not be confused with other Peruvian genera. However, distinguishing species is another matter. To date there has been no comprehensive study of the genus, and many of the characters used by Underwood to separate taxa are highly suspect: e.g., length of fertile pin- nae, number of veins per centimeter, veins paired in origin versus distinctly forked. Species previ- ously thought of as distinct in Central or South America may be synonymous with some in the West Indies. Although about 35 species have been recognized, 20-25 is a more realistic total to expect once a greatly needed revision has been under- taken. Six species from Peru are recognized in the following treatment. References PROCTOR, G. R. 1977. Danaea, pp. 45-49, in Howard, R. A., ed., Flora of the Lesser Antilles: Leeward and Windward Islands, Vol. 2, Pteri- dophyta, Arnold Arboretum, Harvard Univer- sity, Jamaica Plain, Mass. UNDERWOOD, L. M. 1 902. A review of the genus Danaea. Bull. Torrey Hot. Club, 29: 669-679. Key to Species of Danaea a. Larger sterile pinnae (5-)6-40 cm long; larger fertile pinnae 4-25 cm long b b. Sterile pinnae 2-6 pairs; fertile pinnae 3-5 pairs 2. D. elliptica b. Sterile pinnae (7-)8-16 pairs; fertile pinnae 7-25 pairs c c. Petiole of mature plants lacking nodes; apices of sterile pinnae entire (very rarely serrulate); larger sterile pinnae (2.5-)3-6.5 cm broad 1. D. nodosa c. Petiole of mature plants with 1-3 nodes; apices of sterile pinnae serrate; larger sterile pinnae 1 .2-2.8 cm broad d d. Sterile pinnae broadly oblanceolate, broadest above the middle, abruptly acuminate at the apex; veins predominantly simple, occasionally paired in origin or forked 3. D. oblanceolata d. Sterile pinnae lanceolate to oblong-lanceolate, broadest near or below the middle, tapering gradually to an acuminate or attenuate apex; veins predominantly forked, only occasionally simple or paired at the base 4. D. moritziana a. Larger sterile pinnae 1.5-3.5(-4) cm long; larger fertile pinnae 1.1-1.7 cm long e e. Sterile pinnae strongly inequilateral at base, narrow and rounded basiscopically, much broader and abruptly cuneate to truncate acroscopically; veins predominantly forked, some of them simple; petiole nodes 1-3; terminal segment of sterile lamina equaling or longer than the pinnae, lacking a proliferous bud 5. D. humilis e. Sterile pinnae subequilateral at the truncate base; veins simple, rarely forked; petiole nodes lacking; terminal segment of sterile lamina much shorter than larger pinnae, or replaced by a proliferous bud 6. D. trichomanoides 1 . Danaea nodosa (L.) Sm., Mem. Acad. Roy. Sci. Danaea grandifolia Underw., N. Amer. fl. 16: 18. 1909. (Turin) 5: 420. 1793 Figure 4b-c TYPE: Colombia, Valparaiso, Santa Marta, //. H. Smith 992 (holotype, NY; isotypes, BM, F!, GH!, MO!, us!). Acrostichum nodosum L., Sp. pi. 1070. 1753. LEC- TOTYPE (designated by Proctor, Flora Lesser Antilles 2: 48. 1977): Plumier, Traite foug. Amer., Sterile leaves to 2 m long and 60 cm broad, /. 108, based on specimen from Martinique or imparipinnate; petiole lacking nodes, sparsely sca- Dan^ealongifolia Desv., Ges. Naturf. Freunde Berlin * or abundantly so at base, the scales mostly ap- Mag. Neuesten Entdeck. Gesammten Naturk. 5: pressed, dark brown, peltate, nearly circular and 307. 1811. TYPE: "in antillis" (p). ca. 1 mm in diameter or to 3 mm long, margins TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 17 often fimbriate; rachis not or scarcely alate, sparse- ly scaly; pinnae 7-16 pairs, subsessile to short- stalked, elliptic to oblanceolate, larger ones 20-40 cm long and (2.5-)3-6.5 cm broad, base cuneate, apex long-acuminate, margins entire (very rarely serrulate along the acuminate tip), abaxial surface and veins sparsely provided with minute (ca. 0. 1 mm) scales, these appressed, nearly circular, dark or reddish brown, commonly erose to stellate; veins mostly paired or 1 -forked near the base. Fertile pinnae 7-14 pairs, 12-25 cm long, 1.5-2.5 cm broad, subsessile to short-stalked, the apex acute to acuminate. Terrestrial, in dense forests, 100-2100 m, Ama- zonas and Loreto to Cuzco and Madre de Dios. Southern Mexico to Panama; West Indies; Co- lombia; Venezuela; Surinam; Ecuador; Peru; Bra- zil. Danaea grandifolia is said to differ from D. no- dosa in the broader sterile pinnae and more broad- ly cuneate bases of fertile pinnae. The latter char- acter is not consistent, and the former is hardly sufficient reason for separation even at infraspe- cific level. There is also some question as to the merit of separating D. nodosa and D. elliptica. While the characters of nodose or non-nodose pet- ioles and pinnae number usually serve to distin- guish the two, even these have not been reliable consistently in specimens seen from Peru. Further study is needed. Ama/onas: E of Huampami, near Pedro Alberto, Ber- lin 1524 (MO). San Martin: Prov. Rioja, km 390, Pedro Ruiz-Moyobamba road. D. Smith & Vasquez 4726 (F, MO). Loreto: Balsapuerto (lower Rio Huallaga basin), Kil- lip & Smith 28410 (GH, us). Huanuco: Prov. Pachitea, Dist. Honoria, en frente a Tournavista, al este del Rio Pachitea, J. Schunke 1800 (F, GH). Pasco: Puerto Ber- mudez (as Junin), Killip & Smith 2651 1 (NY, us). Junin: Prov. Satipo, near Satipo, van der Werffet al. 8644 (MO). Cuzco: Prov. Quispicanchi, Punkiri Rio Arriba, Vargas 16121 (GH). Madre de Dios: Prov. Tambopata, SSW of Puerto Maldonado, Barbour 5178 (F, GH, MO). 2. Danaea elliptica Sm. in Rees, Cycl. 1 1 : Danaea No. 2. 1808. LECTOTYPE (designated by Proctor, Flora Lesser Antilles 2: 48. 1977): Sloane Herb. 1: 85, based on plant from Mt. Diablo, Jamaica (BM). Danaea elliptica var. crispula Rosenst., Repert. Spec. Nov. Regni Veg. 7: 310. 1909. TYPE: Peru, San Martin, "In montibus secus flumen Mayo, prope Tarapoto," Spruce 4770 (holotype, K; isotypes, GH!, L, P!, uc!. us!; photo, F of L!). Sterile leaves to 1 m long and 30 cm broad, imparipinnate; petiole with 1-5 nodes, sparsely scaly, the scales appressed, dark brown, amor- phous, peltate, often fimbriate, commonly less than 2 mm long; rachis nonalate to narrowly so distally (or alate throughout in juvenile leaves), sparsely scaly; pinnae 2-6 pairs, subsessile to short-stalked, more or less elliptic, 8-18 cm long, (2.5-)3— 4.5 cm broad, base cuneate, apex acute to acuminate, margins entire and plane to somewhat undulate, abaxial surface and veins very sparsely provided with minute (ca. 0. 1 mm) scales, these appressed, nearly circular, dark or reddish brown, commonly erose to stellate; veins mostly paired or 1 -forked near the base. Fertile pinnae 3-5 pairs, 5-14 cm long, (l-)1.7-2.8(-3) cm broad, short-stalked, the apex acute to acuminate. In dense forests, principally along the Cordillera Oriental, Amazonas to Junin, 125-1600 m. Southern Mexico to Panama; West Indies; the Guianas to Colombia and south to Brazil and Peru. There is no justification in recognizing Rosen- stock's var. crispula as a separate entity; suppos- edly it differed from the typical in having one less pair of pinnae, these broadest at or above the mid- dle, with margins more crisped. All fall within the natural variation of this species. Amazonas: Prov. de Bagua, valley of Rio Maranon near Cascadas de Mayasi, Wurdack 1934 (GH, us). San Martin: On ridge in jungle E of Tingo Maria, Allard 21381 (GH, us). Loreto: Fierro Cano a km 4 del Centre Forestal Jenaro Herrera, Spichiger et al. 1439 (MO). Huanuco: Dist. Churubamba, Hacienda Mercedes, trail to Balsa-playa, Mexia 817 Oa (uc). Pasco: Prov. Oxa- pampa, Palcazu Valley, Iscozacin, Foster 9502 (MO). Junin: Pichis Trail, Yapas, Killip & Smith 25507 (GH, NY, US). 3. Danaea oblanceolata Stolze, Amer. Fern J. 77: 33. 1 987. TYPE: Peru, Dept. Pasco (as Junin), Cahuapanas, on Rio Pichis, Killip & Smith 26777 (holotype, us!; frag., F!). Sterile leaves 40-50 cm long, 13-18 cm broad, apical segment (on mature leaves) replaced by a proliferous bud; petiole with 1-2 nodes, moder- ately to abundantly scaly; rachis narrowly alate; pinnae 10-12 pairs, mostly short-stalked, oblong to (more commonly) broadly oblanceolate, larger ones 7-11 cm long and 2-2.8 cm broad, inequi- lateral at base, narrow and rounded to cordate basiscopically, broader and cuneate acroscopical- ly, terminating abruptly in an acuminate and ser- 18 FIELDIANA: BOTANY rate apex, abaxial surface amply provided with minute, dark brown scales; veins commonly sim- ple, but sometimes paired at origin or forked. Fer- tile pinnae 12-14 pairs, larger ones 7-8 cm long and 0.8-1 cm broad, short-stalked, the apex ob- tuse. Terrestrial in dense forests, 0-500 m, thus far known only from Peru: Pasco and Ucayali. This is perhaps most closely related to D. alata Sm., of the West Indies and Venezuela, especially in that both species have predominantly simple veins. However, in D. oblanceolata pinnae are fewer and relatively shorter and broader, and most are broadest well above the middle, where the margins then bend abruptly to a short-acuminate apex. In D. alata, as in all members of the D. moritziana complex, pinnae are broadest at or near the middle, from whence they taper gradually to a moderately acuminate or attenuate apex. Pasco: Oxapampa, Palcazu Valley, Iscozacin, ./?. Foster 9466 (MO), 10049 (F). Ucayali: Vicinity of Aguaytia, Croat 20938 (MO). 4. Danaea moritziana Presl, Abh. Konigl. Bohm Ges. Wiss. 5(4): 35. 1845. TYPE: Venezuela, Colonia Tovar, Moritz 257 (holotype, PR?; is- otype, L; frag., us!; photo, us). Figure 4a. Sterile leaves to 1.2 m long and 35 cm broad (somewhat broader than fertile ones), imparipin- nate, or sometimes with the apical segment re- placed by a proliferous bud; petiole with 1-3 nodes, sparsely scaly; rachis scarcely alate, or slightly so in younger plants; pinnae 8-16 pairs, subsessile to short-stalked, elliptic to oblong-lanceolate, larger ones (5-)6-20 cm long and 1 .2-2.2 cm broad, ine- quilateral at base, narrow and rounded to cordate basiscopically, broader and cuneate to subtruncate acroscopically, gradually tapering to an acuminate or long-attenuate apex, margin subentire but con- spicuously serrate at apex, veins (and often the tissue between the veins) amply provided with mi- nute scales; veins predominantly forked at or near the base, occasionally simple, or paired at origin. Fertile pinnae 8-15 pairs, larger ones 4-11 cm long, 0.6-0.8 cm broad, subsessile or short-stalked, the apex acute to apiculate. Terrestrial, in dense forests, often along streams or in ravines, 300-2300 m, Cajamarca to Loreto, south to Cuzco and Madre de Dios. Colombia; Venezuela; Peru. Except for Danaea nodosa, this is the most widely distributed representative of Danaea in Peru. Moreover it is likely that the Central American D. cuspidata Liebm. also belongs here, along with one or two West Indian species. A number of taxa with pinnae under 2.5 cm broad were separated by Un- derwood (1902), merely on the degree of forking and spacing of veins, an inconsistent character cor- related rarely or not at all by other features, hence a comprehensive revision of the genus should prove many of these to be synonymous. Peruvian spec- imens of D. moritziana apparently are more vari- able than in other areas, sometimes having smaller and fewer pinnae as in Venezuela, sometimes with more and larger pinnae as in "D. cuspidata" of Costa Rica. A few very large specimens from Tin- go Maria have pinnae with rather glossy under- surfaces and extremely attenuate apices. Very closely related to this is D. alata Sm. of the Lesser Antilles, Trinidad, and Venezuela. Although D. alata tends to have more numerous pinnae than D. moritziana, the only consistent difference be- tween the two is that of simple versus forked veins. To further add to the confusion, some specimens of D. moritziana in Peru have been incorrectly determined as D. stenophylla Kunze, which is merely a synonym for D. alata. Underwood (1902) described both of the latter as having unforked veins and distinguished them principally by the degree of crowding of veins, and that D. stenophylla was a "small" plant. Later (1909) he reversed himself by describing D. steno- phylla as a "rather tall plant" and as having mostly forked veins. Obviously he had not seen the type collection, for the isotype (2 sheets) at Kew con- tains leaves to nearly a meter long, with veins obviously simple (only rarely forked), differing from D. alata in no way that we can perceive. We cannot be certain which West Indian plants with forked veins were seen by Underwood, and later by Proctor (1977), but perhaps they are D. ja- maicensis. Cajamarca: Tambillo, Jelski 1073 (P). Amazonas: Bag- ua, ca. 25 km E of La Peca, Barbour 2969 (MO). San Martin: Monte Campana, Tarapoto, Spruce 4711 (P). Loreto: Rio Corrientes, Cachuela, McDaniel 11198 (F, GH, MO). Huanuco: Tingo Maria, Tryon & Tryon 5278 (F, GH, uc, us). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8434 (MO). Junin: Chanchamayo Valley, C. Schunke 156 (F, us). Cuzco: Prov. La Convention, ca. 10 km from Hacienda Luisiana, Dudley 10412 (GH). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa 10-15 km NNW of Shintuya, Foster 10750 (F). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 19 5. Danaea humilis Moore, Index fil. 286. 1861. LECTOTYPE (designated by Lellinger in Proc. Biol. Soc. Wash. 89: 710. 1987): Peru, Dept. San Martin, Tarapoto, Spruce 4769 (lecto- type, K; isolectotypes, BM!, GH!, L!, P!, us!). Sterile leaves 20-34 cm long, 3.5-7 cm broad, larger than the fertile ones, imparipinnate, the ter- minal segment equaling or longer than the largest pinnae, with or without a proliferous bud; petiole with 1-3 nodes, amply to abundantly scaly; lamina slightly shorter than the petiole, the tissue between the veins abundantly provided with minute scales; rachis conspicuously alate, the margins of wings sometimes revolute, but not crispate; pinnae 14— 24 pairs, oblong-elliptic, subfalcate, 2.2-3.5(-4) cm long, 0.6-1(-1.4) cm broad, strongly inequilateral at base, narrow and rounded basiscopically, much broader and abruptly cuneate to truncate acros- copically, apex acute to subacuminate and usually serrulate, margin subentire, essentially plane; veins predominantly forked above the base, yet many of them simple. Fertile lamina scarcely alate along rachis; pinnae 12-25 pairs, 1.1-1.4 cm long, 0.2- 0.3 cm broad, subsessile to short-stalked, mostly acute to subapiculate. In Peru known only from the type collection: San Martin. Rare, terrestrial, in rain forests, 1000-1400 m, Venezuela; Colombia; Ecuador, Peru; Bolivia. This and Danaea trichomanoides are among the smallest species of the genus, with even the larger sterile pinnae commonly less than 3 cm long. Oth- er than size, however, they share few other similar features, as evidenced by the characters listed in the key. 6. Danaea trichomanoides Moore, Index fil. 288. 1861. TYPE: Peru, "in monte Guayrapurima, prope Tarapoto" (San Martin), Aug. 1856, Spruce 4710 (holotype, K!; frag., NY!; photos, F, GH; isotypes, BM!, P!). Sterile leaves 10-12 cm long, 3—4 cm broad, shorter than the fertile ones, with a terminal seg- ment much shorter than the larger pinnae, or re- placed by a proliferous bud; petiole lacking nodes, abundantly scaly; lamina 2-4 times longer than the petiole, the tissue between veins essentially lacking scales; rachis broadly crispate-alate; pin- nae 1 1-15 pairs, oblong, 1.5-2.2 cm long, 0.5-0.9 cm broad, subequilateral at the truncate base, apex rounded to subacute, margin slightly crispate; veins predominantly simple, rarely forked. Fertile lam- ina with rachis scarcely alate; pinnae 1.4-1.7 cm long, 0.3-0.4 cm broad, obviously stalked, apex rounded. Thus far known only from the type collection. When Moore published the description of the species, taken from a Spruce manuscript, he noted that pinnae were "unequal at the base." Perhaps this conclusion was drawn from observing the cris- pate margins of some of the pinnae which were partially folded on drying. However, study of the two type specimens reveals that the bases of most pinnae are subequally truncate and approximately as broad on either side of the costa. This is in marked contrast with most species of Danaea, whose bases are commonly broader and less abruptly tapered on the acroscopic side. Danaea trichomanoides is comparable to D. wendlandii Reichb. f., from Costa Rica, for the two are similar in size, and have nodeless petioles and crispate pinnae, with the presence (usually) of a terminal bud. However, the latter has pinnae with inequilateral bases, veins predominantly forking, and rachis wing plane or revolute (not crispate). Family 3: OSMUNDACEAE Osmundaceae Bercht. & J. S. Presl, Prir. rostlin 1: 272. 1820. TYPE: Osmunda L. Stem massive, woody, decumbent to erect, lack- ing indument, with hard, fibrous roots. Leaves cir- cinate in vernation, densely caespitose, pinnate to pinnately compound, monomorphic to (in Os- munda) partly or fully dimorphic. Petiole with an expanded, stipular base, not articulate to the stem. Veins free. Sporangia not in definite sori, borne abaxially along the segments or (in Osmunda) completely replacing the vegetative tissue of the lamina or of only some of the pinnae, exindusiate, short-stalked, with walls 1 cell thick, the annulus lacking or of only a few thickened cells near the distal end. Spores uniform, green, 1 20-5 1 2 in each sporangium. The three genera of the Osmundaceae are rep- resented by 1 5-20 species. Osmunda is nearly cos- mopolitan, while Leptopteris and Todea are con- 20 FIELDIANA: BOTANY fined to the Old World. This is a very old fern family; only Ophioglossaceae and Marattiaceae are considered to be more primitive. Like the other leptosporangiate members of the Filicales, Os- mundaceae have sporangial walls only one cell thick, yet the family is similar to the Eusporan- giatae in that the annulus is lacking or is composed of only a few thickened cells. References BENEDICT, R. C. 1909. Osmundaceae, in N. Amer. fl., 16: 27-28. TRYON, R. M., AND A. F. TRYON. 1982. Os- mundaceae, pp. 50-57, in Ferns and allied plants, Springer- Verlag, New York. I. Osmunda Osmunda L., Sp. pi. 1063. 1 753. TYPE: O. regalis L. Figure 5. Plants terrestrial. Leaves coarse, 1-2 m or long- er, 1-2-pinnate, completely or partially dimor- phic, the fertile ones (or portions of them) lacking green leaf tissue. Sterile lamina commonly gla- brous at maturity, but the axes often sparsely to moderately provided with trichomes. Veins free, at least 1 -forked. Sporangia relatively large, de- veloping simultaneously, commonly borne in clusters on the segments of the fertile leaves. Spores tetrahedral-globose, trilete, rugose or crested with slender echinate processes. This is a genus of about 1 0 species which gen- erally grow in swampy areas in temperate and tropical regions of both hemispheres. Two species occur in Peru. Key to Species of Osmunda a. Sterile lamina 1-pinnate-pinnatisect; ultimate segments entire, the veins commonly 1 -forked; fertile lamina nonfoliaceous throughout 1 . O. cinnamomea a. Sterile lamina 2-pinnate; ultimate segments serrulate, the veins commonly 2-forked; fertile lamina nonfoliaceous and fertile only in the distal portion 2. O. regalis var. spectabilis 1. Osmunda cinnamomea L., Sp. pi. 1066. 1753. TYPE: United States, Maryland, Kalm (ho- lotype, LINN 1 244. 1 2; photo, F). Figure 5c-d. Leaves completely dimorphic, vegetative tissue essentially lacking in fertile ones. Petiole glabrous or moderately provided with tortuous, filiform, pluricellular, reddish brown trichomes, sheathed at the base. Sterile lamina 1-pinnate-pinnatifid, tissue glabrous, but with filiform trichomes as on the petiole appearing especially at bases of pinnae, rachis narrowly alate adaxially; pinnae to 10 cm long, spreading to strongly ascending, subopposite, articulate with age at the rachis, deeply incised to form broad, subfalcate, obtuse or subacute seg- ments, the segment margins subentire and bearing short, filiform trichomes; veins 1 -forked. Fertile lamina nearly 2-pinnate, the pinnae more strongly ascending than the sterile ones, with dark, filiform trichomes abundantly interspersed among the crowded sporangia. In wet places, usually in sphagnum swamps, 1 700-2800 m, Cajamarca, Amazonas, and Pasco. Eastern and central United States and Canada; West Indies; southern Mexico to Honduras; Costa Rica; Colombia; Venezuela; Brazil; Peru; Para- guay; southeast Asia. Some earlier authors recognized this in the Neo- tropics as var. imbricala (Kunze) Milde, distin- guishing it from North American plants on the basis of darker-colored tomentum and thicker lam- ina. We find little justification for the separation. Cajamarca: Jaen, Quebrada de Pajonal, above Taba- conas, 19 km ESE Huancabamba, Fosberg 27795 (MICH, NY). Amazonas: Chachapoyas, Jalca zone, 3-6 km W of Molinopampa, Wurdack 1404 (us). Pasco: (as Junin) Pichis trail, Enenas, Killip & Smith 25708 (NY, us). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 21 FIG. 5. Osmunda regalis var. spectabilis: a, habit; b, sterile ultimate segment. Osmunda cinnamomea: c, habit; d, sterile ultimate segment. (From Stolze, Ferns and fern allies of Guatemala, 1976.) 22 FIELDIANA: BOTANY 2. Osmunda regalis L. var. spectabilis (Willd.) A. Gray, Manual ed. 2: 600. 1856. Figure 5a-b. Osmunda spectabilis Willd., Sp. pi. 5: 98. 1810. TYPE: United States, Pennsylvania, Muhlenberg (holo- type, B, Herb. Willd. 19504; photo, GH). Osmunda palustris Schrader, Gott. Gel. Anz. 866. 1824. TYPE: not located. Osmunda regalis ssp. palustris (Schrader) Love & Love, Taxon 26: 324. 1977. length of their stalks. The characters are too in- significant and inconsistent to merit such a dis- tinction. Amazonas: Mendoza, on rocky wall of ravine, Woyl- kowski 8253 (GH, MO, us). Prov. Bongara, W & S margins of Laguna Pomacocha, Wurdack 896 (us). Pasco: Prov. Oxapampa, Canyon de Huancabamba, Leon 612 (GH). Cuzco: Empalizada, Biies 1705 (us). Leaves partly dimorphic, fertile ones with fertile and sterile portions combined on the same lamina, the fertile pinnae commonly distal and essentially lacking vegetative tissue. Petiole glabrous at ma- turity, sheathed at the base. Sterile lamina 2-pin- nate, chartaceous to subcoriaceous, tissue gla- brous, but a few tortuous, pluricellular, reddish brown trichomes often clustered at the base of pinnules, the rachis nonalate; pinnae to 20 cm long, strongly ascending, subopposite, short- stalked, articulate with age at the rachis, costae very narrowly alate (at least distally) on adaxial side; pinnules distant, subopposite, short-stalked, articulate with age at the costa, oblong to narrow- elliptic, obtuse to subacute, margins serrulate. Veins mostly 2-forked. Fertile pinnae 2(-3)-pinnate, more strongly ascending than the sterile ones, the crowd- ed sporangia almost completely replacing the leaf tissue. In damp places near lakes or streams, occasion- ally among rocks, 1300-2200 m, Amazonas, Pas- co, Cuzco. Eastern and central United States and Canada; West Indies; southern Mexico; Guatemala; Hon- duras; Costa Rica; Colombia; Venezuela; Brazil; Ecuador, Peru; Paraguay; Uruguay. Variety regalis, which occurs in the Old World, is said to differ in its broader leaves, narrower panicles, and more numerous blackish to casta- neous trichomes persistent along the rachis. Al- though specimens we have seen from both the New and Old World do not seem to differ strongly or consistently enough to warrant separation, we maintain the traditional varietal distinction for purposes of this treatment. Earlier workers pro- posed yet another variant, ssp. palustris, for South American plants, based on size of segments and Family 4: SCHIZAEACEAE Schizaeaceae Kaulf., Wesen farrenkr. 119. 1827. TYPE: Schizaea Sm. Stem erect to decumbent, usually small and sometimes branched, or long-creeping, slender and freely branched, provided with trichomes or (in Mohria of Africa) with scales. Leaves circinate in vernation, entire or filiform, or dichotomous, or pinnate, glabrous to pubescent or (in Mohria) with scales, partially or wholly dimorphic. Sporangia borne abaxially on slightly to strongly modified portions of the leaf, separate, or crowded on each side of a vein, or in loose clusters on wholly fertile panicles, sessile, or with a short, many-rowed stalk, and an apical annulus. Spores tetrahedral-globose and trilete, or ellipsoidal and monolete, lacking chlorophyll. This is an old family, with the four extant genera so highly diverse in habit and form that some authors prefer to segregate them into distinct fam- ilies. Mohria is confined to Africa and Madagas- car; Anemia is in America, Africa, and southern India; the other two genera are pantropic. All ex- cept Mohria occur throughout tropical America and are all well represented in Peru. References LELLINGER, D. B. 1969. Schizaeaceae, in The botany of the Guayana Highland— Part VIII. Mem. New York Bot. Gard., 18: 1-11. MAXON, W. R. 1909. Schizaeaceae, in N. Amer. fl., 16: 31-52. Key to Genera of Schizaeaceae a. Leaves erect or suberect; pinnae lacking arrested buds in branch axils; sporangia not covered by a laminar flange b TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 23 b. Leaf pinnate or pinnately decompound, rarely pinnatifid I. Anemia b. Leaf filiform or with dichotomous veins or branches III. Schizaea a. Leaves vinelike, spreading and scandent; pinnae short-stalked, with an arrested bud in the axil of the first, pseudodichotomous branch; sporangia each covered by a laminar flange . . II. Lygodium I. Anemia Anemia Sw., Syn. fil. 6, 155. 1806, nom. conserv. (sometimes as "Aneimia"). TYPE: Anemia phyllitidis (L.) Sw. (Osmunda phyllitidis L.). Figure 6. Coptophyllum Gardn., London J. Bot. 1: 133. 1842. TYPE: Coptophyllum buniifolium Gardn. = Ane- mia buniifolia (Gardn.) Moore. Anemirhiza J. Sm., Seemann, Bot. Voy. Herald 243. 1854. TYPE: Anemirhiza adiantifolia (L.) J. Sm. (Osmunda adiantifolia L.) = Anemia adiantifolia (L.) Sw. Plants terrestrial. Stem decumbent to creeping, rarely erect, bearing short to long trichomes and fibrous roots. Leaves ca. 1-75 cm long, erect, par- tially (as in Peruvian species) dimorphic, with a pair of fertile pinnae at the base of or below the sterile lamina, or with similar leaves longer and more erect than the wholly sterile leaves, or wholly dimorphic, the whole leaf either sterile or fertile. Sterile lamina 1-2-pinnate (rarely pinnatifid); veins free, or rarely anastomosing without included free veinlets. Sporangia borne on fertile segments which are reduced to mere axes or have narrow borders of laminar tissue along the axes. Spores tetrahe- dral-globose, trilete. The genus contains about 80 species, which oc- cur in tropical to subtropical regions of both hemi- spheres, predominantly in the Neotropics. Species limits are not always clearly drawn, as diagnostic features are often quite variable, such as: relative length of fertile pinnae and sterile portion of lam- ina; color of rhizome indument; degree of dissec- tion of pinnae. Color of trichomes is particularly difficult to assess, inasmuch as each observer sees colors in different ways, and definitions of color terms are usually difficult to interpret. Rhizome indument in Anemia generally grades from orange to reddish orange to red to brownish red; in many specimens the distinctions are clear, but in others they are somewhat inconstant. Hybridization is not uncommon in the genus and must be taken into consideration when attempting identification of collections. The following key will be more ef- fective if attention is given to combinations of characters, whenever possible. Reference MICKEL, J. T. 1962. A monographic study of the fern genus Anemia, subgenus Coptophyllum. Iowa State Coll. J. Sci., 36: 349-482. Key to Species of Anemia a. Fertile pinnae borne 3 mm or more below the base of the sterile lamina; ultimate fertile segments with narrow (but distinct) borders of laminar tissue along the axes b b. Sterile pinnae crenate to pinnatisect, not cut entirely to the costa (except rarely as to proximal lobes of basal pinnae) 2. A. villosa b. Sterile pinnae 1-2-pinnate c c. Apex of stem ascending or erect, covered with petiole bases; fertile pinnae commonly shorter than the sterile portion of the lamina d d. Petiole slender (less than 1 mm thick), dark brown; fertile pinnae suberect, with stalk 0.3- 0.8 cm long 1 . A. clinata d. Petiole stout (1-2 mm thick), commonly yellow; fertile pinnae erect, with stalk 1.5-5 cm long 3. A. flexuosa c. Apex of stem horizontal, exposed beyond the most recent petiole bases; fertile pinnae commonly longer than sterile portion of the lamina e e. Lamina hirsute to glabrate, petiole commonly glabrate and atropurpureous (rarely light brown to yellow) 4. A. ferruginea 24 FIELDIANA: BOTANY e. Lamina and petiole conspicuously lanate, petiole yellow 5. A. myriophylla Fertile pinnae borne at base of sterile lamina, or less than 1 mm below it; ultimate fertile segments with laminar tissue lacking or essentially so f f. Veins free; stalk of fertile pinnae usually 2-4 times the length of the panicle; sterile pinnae less than 4 cm long g g. Petiole (at least the fertile) 8-25 cm long; lamina gradually reduced to a pinnatind apex (or rarely terminating in a subconform lanceolate segment); apex of sterile pinnae acute or subacute h h. Sterile pinnae mostly deeply incised, the segments commonly narrow and cuneate 6. A. hirsute h. Sterile pinnae crenulate or denticulate, or rarely with a few deep lobes on several proximal pinnae 7. A. pastinacaria g. Petiole less than 6 cm long; lamina abruptly terminating in a broadly obovate or obdeltoid apical segment; apex of sterile pinnae broadly rounded 8. A. oblongifolia f. Veins anastomosing; stalk of fertile pinnae usually equaling or shorter than the panicle; sterile pinnae (larger ones) 5-14 cm long 9. A. phyllitidis 1. Anemia din at a Mickel, Amer. Fern J. 56: 58. 1966. TYPE: Peru, Junin, along Rio Perene, Killip & Smith 21594 (holotype, us!; isotypes, F!, NY!). Stem with apex ascending and thickly covered with petiole bases, densely provided with orange trichomes. Leaves 10-25 cm long, 4-9 cm broad, pilose to hirsute, the suberect fertile pinnae borne 8-16 mm below the sterile portion of the lamina; petiole dark brown, less than 1 mm thick. Sterile lamina 1 -pinnate-pinnatisect to 2-pinnate, with 5- 1 1 pairs of pinnae; pinnae dissected nearly or quite to the costa, the ultimate segments joined at the base or discrete, but broadly adnate, not stalked; veins free. Fertile pinnae suberect, usually shorter than the sterile portion of the lamina, short-pe- tiolulate (stalk 0.3-0.8 cm long), the ultimate fer- tile segments with narrow bands of laminar tissue on each side of the axes. In thickets and deep forests, 600-1 300 m, Junin. Panama; Colombia and Venezuela to Bolivia and western Brazil. One of the distinguishing features of A clinata is the suberect or slightly spreading position of the fertile pinnae, which are rigidly erect in most species of Anemia. It is also one of the smaller and more delicate species, with slender, wiry petioles less than 1 mm thick. Junin: Near La Merced, E of Quimiri Bridge, Killip & Smith 23591 (NY, us). Colonia Perene, Killip & Smith 25036 (F, NY, us). La Merced, Kunkel 564 (OH). Chan- chamayo Valley, C. Schunke 78 (F, us). 2. Anemia villosa Willd., Sp. pi. ed. 4, 5: 92. 1 8 1 0. TYPE: "America meridionale," Humboldt & Bonpland (holotype, B, Herb. Willd. 19496; photos, GH, us). Venezuela, Prov. Cumana, "Between Catuaro and Cariaco": fide HBK., Nov. gen. sp. fol. ed. 1: 26. 1815. Stem with apex ascending and thickly covered with petiole bases, or the apex horizontal and ex- posed beyond the most recent petioles, densely provided with orange to brownish red trichomes. Leaves 10-55 cm long, villous to hirsute, the erect fertile pinnae borne 3-18 mm below the base of the sterile portion of the lamina; petiole yellow to light brown, 1-2 mm thick. Sterile lamina 1 -pin- nate to pinnate-pinnatisect, with 7-19 pairs of pin- nae; pinnae with rounded lobes, not cut entirely to the costa (except rarely as to proximal lobes of basal pinnae); veins free. Fertile pinnae erect, slightly shorter than to (more typically) somewhat longer than the sterile portion of the lamina, their stalks 1-5 cm long, the ultimate fertile segments with narrow bands of laminar tissue on each side of the axes. On rocky, open slopes, 900-2000 m, Cajamarca, Amazonas, San Martin, Ayacucho, Cuzco. Surinam to Colombia, south to Peru; east coast of Brazil. This appears to differ only quantitatively from Anemia flexuosa, the latter with sterile pinnae more strongly dissected and, typically, with fertile pin- nae shorter than the sterile portion of the lamina. Fertile pinnae of A. villosa are commonly longer TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 25 FIG. 6. Anemia paslinacaria: a, habit; b, apex of fertile pinna. Anemia phyllitidis: c, sterile pinna, (a-b from Irwin et al. 24260, Brazil, p, c from Stork & Morton 9432, F.) 26 FIELDIANA: BOTANY than the lamina, but occasionally equal or exceed it. Evidently these are the reasons why Prantl con- sidered it merely a variety of A. flexuosa. See Mickel (1962) for more detailed discussion. Cajamarca: Prov. Jaen, San Patricio, Chontali, Chi- moy65 (USM). Amazonas: Prov. Chachapoyas, Cano Santa Lucia, Wurdack 599 (F, GH, NY, uc, us). San Martin: Tarapoto, in monte Campana, Spruce 4708 (GH, NY, p, us). Ayacucho: Prov. La Mar, Aina, Ldpez & Soukup 15.088 (GH). Cuzco: Prov. La Convencion, Urusaiwa, Vargas 22349 (GH). 3. Anemia flexuosa (Savigny) Sw., Syn. fil. 156. 1806. Osmunda flexuosa Savigny in Lam., Encycl. 4: 652. 1797. TYPE: based on undesignated specimen in Jussieu herbarium (not located). Anemia flexuosa var. setosa Prantl, Unters. Morph. Gefasskrypt. 2: 95. 1881. SYNTYPES: Brazil, Lagoa Santa, Warming (not located): Peru, Cu- chero, Poeppig (not located). ISOSYNTYPE: Poeppig (L; photos, GH, us). Stem with apex ascending and thickly covered with petiole bases, densely provided with orange or reddish orange trichomes. Leaves 18-65 cm long, hirsute or glabrate, the erect fertile pinnae borne 5-25 mm below the base of the sterile por- tion of the lamina; petiole yellow to light brown, 1-2 mm thick. Sterile lamina 2-pinnate (some- times 2-pinnate-pinnatifid), with 6-12 pairs of pinnae; pinnae (many of them) divided to the costa into broadly adnate, obtuse pinnules; veins free. Fertile pinnae erect, commonly shorter than the sterile portion of the lamina (rarely equal to it or slightly longer), their stalks 1.5-5 cm long, the ultimate fertile segments with narrow bands of laminar tissue on each side of the axes. In thickets and open woods or on moist rocky slopes or clay banks, 900-3000 m, Amazonas, Huanuco, Junin, Ayacucho, Cuzco. Surinam to Colombia, south to Bolivia. Many Peruvian specimens of Anemia flexuosa are to be found in various herbaria; especially abundant are collections from Huanuco, Junin, and Cuzco. This is a highly variable species, par- ticularly in the dissection of sterile laminae, some- times 2-pinnate-pinnatifid, typically 2-pinnate, occasionally approaching the 1 -pinnate-pinnatifid condition of A. villosa (q.v.) and thus is often con- fused with it. As in some other species of Anemia, this tends to hybridize; in fact there was a hybrid found in Cuzco: A. flexuosa x A. phyllitidis, Var- gas 19836, Prov. La Convencion, alt. 1650 m (GH). Leaves are much thinner in texture than typical A. flexuosa, with some anastomosing veins, and fertile pinnules are long-petiolulate. Amazonas: Rodriguez de Mendoza, Soukup 5023 (us). Huanuco: Vilcabamba, on Rio Chinchao, Macbride4985 (F, NY, us), Stork & Morton 9876 (F, uc, us). Junin: Huacapistana, Killip & Smith 24152, 24324 (F, NY, us). Ayacucho: Ccarrapa, between Huanta and R;o Apuri- mac, Killip & Smith 22319 (F, GH, NY, us). Cuzco: Prov. La Convencion, Valley of the Sambray, Mexia 8038 (F, GH, MO, uc). 4. Anemia ferruginea HBK., Nov. gen. sp. 1: 32. 1815, var. ferruginea. TYPE: Venezuela, Prov. Cumana, "prope Guardia de San Augustin," Humboldt & Bonpland (holotype, P?). Stem with apex horizontal, and exposed beyond the most recent petioles, densely provided with red or brownish red trichomes. Leaves 10-55 cm long, the erect fertile pinnae borne 3-7 mm below the base of the sterile portion of the lamina; petiole commonly atropurpureous (rarely yellowish), gla- brate, 1-2 mm thick. Sterile lamina 2-pinnate, hirsute to subglabrous, with 8-12 pairs of pinnae; pinnae dissected to the costa, the pinnules mostly discrete, adnate at base, acute or subacute at apex, sometimes those of proximal pinnae shallowly pinnatifid; veins free. Fertile pinnae erect, com- monly longer than the sterile portion of the lamina, their stalks 1 .5-6 cm long, the ultimate fertile seg- ments with narrow bands of laminar tissue on each side of the axes. In sunny places, open woods and clearings, on rocky slopes and clay banks, 850-2200 m, San Martin, Huancavelica, Cuzco. Honduras; Guyana to Colombia, south along the Andes to Bolivia and Brazil. The variety ahenobarba (Christ) Mickel, from Brazil, differs in its more deeply dissected pinnules with acute to acuminate tips. Both are difficult to distinguish from Anemia tomentosa (Savigny) Sw., especially var. anthriscifolia (Schrader) Mickel, from northern Argentina, Bolivia, Paraguay, and southeastern Brazil. See Mickel (1962) for further comments. San Martin: Tarapoto, Spruce 4044 (GH, NY, p, us). Huancavelica: Prov. Tayacaja, Dist. Huachocolpa, near Quintabamba, Tovar 4157, 4693 (GH). Cuzco: Prov. Convencion, Valley of the Sambray, Mexia 8038 (NY, us). Prov. Convencion, Hacienda Sahuayaco, Vargas 1656 (GH, MO). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 27 5. Anemia myriophylla Christ, Bull. Herb. Bois- sier 2(7): 793. 1907. TYPE: Bolivia, Padcaya, Fiebrig 2541 (isotype, p). Stem with apex horizontal and exposed beyond the most recent petioles, densely provided with reddish to orange trichomes. Leaves 20-50 cm long, the erect fertile pinnae borne 3-8 mm below the base of the sterile portion of the lamina; petiole yellow, conspicuously lanate, 1-2 mm thick. Ster- ile lamina 2- to nearly 3-pinnate, conspicuously lanate, with 7-12 pairs of pinnae; pinnae with pin- nules cut deeply or quite to the costule, the tips of ultimate segments commonly broadly rounded; veins free. Fertile pinnae erect, commonly longer than the sterile portion of the lamina, their stalks 1.5-5 cm long, the ultimate fertile segments with narrow bands of marginal tissue on each side of the axes. On open slopes and rocky banks, 2000-2200 m, Cajamarca, Amazonas. Peru; Bolivia; Argentina. Anemia myriophylla has a more highly divided leaf than any Anemia in Peru. This and the densely lanate lamina and petiole make it rather easy to recognize. Cajamarca: Between San Marcos and Cajabamba, Correll & Smith P911 (GH, us). Prov. Celendin, Sagds- legui et al. 8457 (F, HUT, MO, uc). Amazonas: W of Chachapoyas on road to Gaelic. Hutchison & Bennett 4519 (F, GH, MO, NY, uc, us). 6. Anemia hirsuta (L.) Sw., Syn. fil. 155. 1806. Osmunda hirsuta L., Sp. pi. 1064. 1753. LECTO- TYPE (designated by Lellinger in Proc. Biol. Soc. Wash. 98: 387. 1985): Plumier, Traite foug. Amer., /. 762, based on a Plumier collection from His- paniola. Anemia hirsuta var. humboldtiana Hieron., Bot. Jahrb. Syst. 34: 566. 1905. LECTOTYPE (designated by Lellinger in Proc. Biol. Soc. Wash. 98: 366. 1985): Venezuela, border Edo. Sucre- Monagas, Hum- boldt 459 (B, Herb. Willd. 19495-2; photos, GH, us). Stem short-creeping, densely provided with or- ange trichomes. Leaves to 35 cm long and 5 cm broad, glabrate or somewhat villous on axes and lamina, the erect, fertile pinnae borne at the base of the sterile portion of the lamina; petiole (at least on fertile leaves) 8-1 8 cm long. Sterile lamina pin- nate-pinnatifid to nearly 2-pinnate, with 7-1 2 pairs of pinnae, gradually tapering to a pinnatifid apex; pinnae to 2.5 cm long and 0.8 cm broad, com- monly inequilateral at base, truncate acroscopi- cally and cuneate basiscopically, obliquely incised into linear or narrowly cuneate segments, obtuse to subacute at apex; veins free. Fertile pinnae to 18 cm long, the stalk (1.5-)2-3 times the length of the panicle, the ultimate fertile segments essen- tially lacking laminar tissue. In dry, grassy areas, on slopes or mesas, 1 200- 2 1 50 m, Huanuco, Cuzco. Mexico to Panama; Greater Antilles; Trinidad & Tobago; Venezuela and Colombia south to Bo- livia and Brazil. This is scarcely distinct from Anemia pastina- caria, essentially differing only by the characters noted in the key. More detailed studies are needed to clarify relationships of the several variable taxa of the A. hirsuta group which, furthermore, appear to hybridize readily with a number of other species of Anemia throughout the entire range. For ex- ample, there is a specimen of A. pastinacaria at Paris (San Martin, Tarapoto, Spruce 4134) which also contains a very large, highly dissected leaf with abortive spores— possibly a hybrid between A. hirsuta and A. phyllitidis. Huanuco: Muna, Bryan 427 (F, GH). In grass steppe, Woytkowski 219 (uc). Cuzco: Quillabamba, Coronado 124 (uc). Dept. Unknown: Mat hews 3299 (us). 7. Anemia pastinacaria Prantl, Unters. Morph. Gefasskrypt. 2: 110. 1881. LECTOTYPE (designated by Lellinger in Proc. Biol. Soc. Wash. 98: 367. 1 985): Venezuela, "in convalli del Tigre," Moritz 26 (B; probable isolecto- type, GH!). Figure 6a-b. This differs from Anemia hirsuta essentially only in the following characters: Leaves to 45 cm long and 7 cm broad. Sterile lamina gradually tapering to a pinnatifid apex, but occasionally with a dis- tinct, nonconform apical segment; pinnae to 3.5(-5) cm long and 1 .5(-2) cm broad, margins denticulate or crenate, or rarely with a few deep lobes on sev- eral proximal pinnae. On dry, rocky slopes or in forest clearings, 750- 2000 m, San Martin, Loreto, Huanuco, Junin, Ay- acucho, Cuzco. Mexico to Panama; Cuba; Jamaica; Trinidad; Surinam to Colombia, south to Bolivia and Brazil. This and Anemia hirsuta differ only in their rel- ative size and depth of dissection of pinnae, and 28 FIELDIANA: BOTANY a few specimens in Peru tend to be intermediate even in these characters. See A. hirsuta for further discussion. San Martin: Tarapoto, Spruce 4 134 (p in part). Loreto: Yanayacu, Biies 1037 (us). Huanuco: Yanano, Macbride 3816 (F, us). Junin: Prov. Huancayo, Pariahuanca, Tovar 7948a (USM). Ayacucho: Aina, between Huanta and Rio Apurimac, Killip & Smith 22614 (F, NY, us). Cuzco: Prov. Convencion, Hacienda Sahuayaco, Vargas 1655 (GH), 1657 (GH, MO). 8. Anemia oblongifolia (Cav.) Sw., Syn. fil. 156. 1806. Osmunda oblongifolia Cav., Icon. 6: 69, /. 592, / 2. 1801. TYPE: Panama, Nee (holotype, MA), ver- ified by C. Christensen [Dansk. Bot. Ark. 9(3): 31. 1937]. Osmunda humilis Cav., Icon. 6: 69, /. 592, f. 3. 1801. TYPE: Panama, Taboga Island, Nee(nol located); earlier presumed at MA, but not seen by C. Chr. [Dansk. Bot. Ark. 9(3): 31. 1937]. Anemia humilis (Cav.) Sw., Syn. fil. 156. 1806. Anemia oblongifolia var. humilis (Cav.) Hooker & Baker, Syn. fil. 431. 1868. Stem decumbent, rather densely provided with orange trichomes. Leaves to 30 cm long and 3.5 cm broad, sparsely to densely villous on axes and lamina, the erect fertile pinnae borne at the base of the sterile portion of the lamina; petiole com- monly less than 6 cm long. Sterile lamina 1 -pin- nate, with 2-9 pairs of pinnae, abruptly terminat- ing in an obovate or obdeltoid apical segment which is often as broad as long; pinnae to 1.8 cm long and 0.7 cm broad, strongly inequilateral at base, truncate or rounded acroscopically and cuneate basiscopically, broadly rounded at apex, the mar- gins crenulate or subentire; veins free. Fertile pin- nae to 1 8 cm long, the stalk 2—4 times the length of the panicle, the ultimate fertile segments essen- tially lacking laminar tissue. In forests, on rocky slopes, 1000-1 100 m, Cuz- co. Mexico to Panama; Venezuela to Brazil and Bo- livia. Anemia oblongifolia is rather distinctive in its coarse pinnae which are broadly rounded at their tips and often nearly as broad as long. Sterile leaves are commonly very short-petiolate (often subses- sile), and typically are so densely caespitose that they give the appearance of a rosette. Some authors have separated A. humilis on the strength of its villous sterile pinnae with crenulate margins; but these characters are too variable and inconsistent to be of real value. We tentatively maintain it here as a synonym, although we have not seen the type to fully substantiate this. Cuzco: Echarate, Camino de Sahuayacu, Bites 845 (us). Ccochayoc, Biies 1712, 1719 (us). Prov. Convencion, Valle de Lucumayo-Pistipata, Vargas 4176 (MO, uc, us). 9. Anemia phyllitidis (L.) Sw., Syn. fil. 155. 1806. Figure 6c. Osmunda phyllitidis L., Sp. pi. 1064. 1753. TYPE: based on illustration of Hispaniola plant in Plu- mier, Traite foug. Amer., /. 156. 1705. Anemia haenkei Presl, Reliq. haenk. 1: 74. 1825. TYPE: Peru, "in vallibus Cordillerarum Peru- viae," Haenke (holotype, PRC?). Stem decumbent (rarely short-creeping), rather densely clothed with orange or brownish tri- chomes. Leaves to 80 cm long and 20 cm broad, the erect fertile pinnae borne at the base of the sterile portion of the lamina or less than 1 mm below it; petiole (at least on fertile leaves) ca. 10- 35 cm long. Sterile lamina 1 -pinnate, with 3-7 pairs of pinnae and a conform apical one; pinnae (larger ones) 5-14 cm long and 1.3-3 cm broad, subequilateral, ovate to lanceolate, acute to acu- minate, the margins crenulate to dentate (or rarely the basal pair lobed); veins copiously anastomos- ing. Fertile pinnae to 40 cm long, the stalk equaling or (commonly) shorter than the panicle; the ulti- mate fertile segments essentially lacking laminar tissue. Moist, often rocky slopes and ravine banks, in and at edges of forests, 700-1700 m, Amazonas to Loreto, south to Huancavelica and Cuzco. Mexico to Panama; Greater Antilles; Trinidad; Venezuela and Colombia to Argentina and Uru- guay. This is one of the most common species of Ane- mia in Peru and, with its large, subentire pinnae and reticulate venation, is the easiest to recognize. Amazonas: Prov. Bagua, S of Bagua Grande on Rio Utcubamba, Hutchison 1479 (uc, us). San Martin: Ze- pelacio, near Moyobamba, Klug 3440 (F, MO, NY, uc, us). Loreto: Yanayacu, Biies 2005 (us). Huanuco: Prov. Huanuco, Puente Durand, Stork & Morton 9432 (F, GH, MO, uc, us). Pasco: Oxapampa, Soukup 2361 (F, GH). Junin: Yucapata, Woytkowski 6642 (GH, MO, us). Huan- cavelica: Prov. Tayacaja, SE of Tintay, Tovar 46 11 (GH). Ayacucho: Aina, between Huanta and Rio Apurimac, Killip & Smith 22719 (NY, us). Cuzco: Prov. La Con- TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 29 vencion, upper valley of Rio Sanbray, Mexia 8056a, (F, GH, uc, us). Comments Anemia hispida Kunze, Linnaea 9: 20. 1834. TYPE: "In fruticetis ad Chibangata fl. Pe- ruv.." 1829, Poeppig diar. 1165 (LZ? de- stroyed, B?, w?). Apparently known thus far only from the type collection. Kunze's plant was described as having: creeping rhizome; pinnate, hispid leaves; fertile pinnae twice as long as the sterile lamina; pinnae remote, pat- ent, sessile, dimidiate, oblong, obtuse, with mar- gins crenate or subincised. He likened it to A nemia repens Raddi (= A. hirsuta1?), but having pinnae less incised. Assuming it belongs to subg. Anemia (unsubstantiated by the description), it could be A. pastinacaria, but we cannot be certain without examining the type. II. Lygodium Lygodium Sw., J. Bot. (Schrader) 1800(2): 106. 1 802, nom. consent TYPE: Lygodium scan- dens (L.) Sw. (Ophioglossum scandens L.). Figure 7. Plants terrestrial. Stem slender, short- to long- creeping, provided with short trichomes. Leaves 1-10 m long, vinelike, indeterminate, spreading and scandent, partially dimorphic, the fertile por- tions somewhat contracted with marginal fertile lobes, pinnate, glabrous to somewhat pubescent. Pinnae short-stalked, pseudodichotomously branched with an arrested bud in the axil, each primary pinna-branch radiately lobed or branched or pinnate. Veins reticulate or (in Peruvian species) free. Sporangia borne separately on marginal lobes of a pinna segment or on a wholly fertile segment, each covered by a laminar outgrowth (flange). Spores tetrahedral-globose, trilete. Lygodium is a natural group, easily distin- guished within the family by its long leaves and vinelike and scrambling habit, often climbing to considerable heights by twining around the small- er branches of shrubs and trees. It is widely dis- tributed, primarily pantropic, with several extra- tropical species both in the New and Old World. There are 30-35 species, but the number could be somewhat reduced by further study, as some ap- pear to intergrade rather freely. Three species are recognized in Peru. References DUEK, J. J. 1978. FeddesRepert., 89:411^23. TRYON, R. M., AND A. F. TRYON. 1982. Lygo- dium, pp. 69-76, in Ferns and allied plants, Springer- Verlag, New York. Key to Species of Lygodium a. Primary pinna-branch pinnate, the pinnules stalked, except sometimes near the apex of the primary branch b b. Pinnules (at least fertile ones) diminishing in size toward the apex of the primary branch, mostly expanded at base into lobes, the lobes often discrete or even stalked 1 . L. venustum b. Pinnules subequal, not expanded at base, subentire, very rarely some basal ones lobed at the base 2. L. volubile a. Primary pinna-branch radiate, the ultimate segments deeply lobed, joined at the base . 3. L. radiatum 1. Lygodium venustum Sw., J. Bot. (Schrader) 1801(2): 303. 1803. TYPE: Jakob Breyne, Cent. I, t. 96. 1678, not "Brazil, Breynius" (designated by Proctor, Flora Lesser Antilles 2: 51. 1977), a specimen which probably does not exist. Figure 7a. Pinna-stalks 2-10 mm long, moderately to densely pubescent with tawny to orange pluricel- lular trichomes. Primary pinna-branches pinnate to (rarely) 2-pinnate, sessile to short-stalked, broadly to narrowly deltoid. Fertile pinnules 9- 2 1 , alternate and commonly widely spaced, all but the distal ones stalked, stalk often nodose at its juncture with pinnule base, sparsely to abundantly pilose on axes, veins and (occasionally) the leaf tissue, larger ones 2.5-5 cm long, gradually di- minishing in size toward the apex of the primary 30 FIELDIANA: BOTANY FIG. 7. Lygodium venustum: a, primary axis and pair of pinnae. Lygodium volubile: b, pinna; c, base of pinnule, (a from Irwin et al. 21469, Brazil, F, b from Jeanpert s.n., Brazil, F, c from L. B. Smith 1352, Brazil, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 31 branch, most of them expanded at base into lobes, the lobes rounded to cordate or subpalmate. Ster- ile pinnules similar to the fertile, but often larger and not much reduced toward the apex of the pri- mary branch. Veins free, 1 -several times-forked, prominulous. Along roadsides and stream banks, in fields or in thickets and open forests, usually twining over low shrubs, 100-1000 m, San Martin, Loreto, Huanuco, Junin, Ucayali, Cuzco. Tropical America. This species is widely distributed throughout the Neotropics. A number of other species have been confused with it or should be included within it. For detailed discussions see Smith, Flora of Chia- pas: Pteridophytes, Calif. Acad. Sci. p. 145. 1981, and Stolze, Ferns & Fern Allies of Guatemala: Part I, Fieldiana, Bot. 39: 38. 1976. The closely related Lygodium volubile readily can be distinguished by the fewer, larger, and subequal pinnules on each pinna-branch; i.e., the distal ones are nearly or quite as large as the proximal ones. Pinnules (at least fertile ones) of L. venustum gradually dimin- ish in size and shape so that subapical ones are often one-third to one-half the size of those near the pinna base. The Breyne plate was cited by Swartz and is clearly this species. San Martin: Dist. San Martin, 2 km NW of Tarapoto, Belshaw 3353 (F, GH, MO, NY, uc, us). Juan Jui, Alto Rio Huallaga. Klug 4218 (F, GH, MO, NY, uc, us). Loreto: Puerto Arturo, lower Rio Huallaga below Yurimaguas, Killip & Smith 27847 (NY, us). Huanuco: Prov. Leoncio Prado, Huachipa, Plowman 5957 (F, GH). Junin: La Merced, Killip & Smith 23379 (F, NY, us). Ucayali: Pu- callpa (as Loreto), Soukup 3049 (F). Cuzco: Santa Ana, Cook & Gilbert 1614 (us). 2. Lygodium volubile Sw., J. Bot. (Schrader) 1801(2): 304. 1803. TYPE: Jamaica, Swartz (holotype, S-PA; photo, us). Figure 7b-c. Lygodium micansj. W. Sturm in Mart., Fl. bras. 1(2): 178. 1859. TYPE: "British Guiana," Schom- burgk 399 (holotype, B). Pinna-stalks 0.5-3 mm long, essentially gla- brous. Primary pinna-branches pinnate or (very rarely) 2-pinnate at base, the pinnae stalked (1.5- 5 cm), broadly oblong. Fertile pinnules 4-10, al- ternate and widely spaced, conspicuously stalked, nodose and often articulate at juncture with the pinnule base, glabrous to sparsely pilose on axes and veins (especially at bases of sporangia), 6-14 cm long, proximal and distal ones subequal, broadly cuneate to truncate at base, only rarely lobed at base. Sterile pinnules similar to the fertile, but often somewhat larger. Veins free, several times-forked, prominulous. In wet, often inundated, lowland forests and open fields, scandent on shrubs and low trees, often along stream and river banks, sea level to 250 m, Ama- zonas, San Martin, Loreto, Madre de Dios. Southern Mexico; Guatemala; Belize; Panama; Greater Antilles; Trinidad; Venezuela to the Guianas, south to Argentina and Brazil. This and Lygodium micans have been separated on the basis of several, variable, quantitative char- acters, although these may appear more distinct in certain regions outside South America: relative size and shape of ultimate segments and their bas- es; degree of pubescence on abaxial surface; angle of veins. None of these are consistently correlated in the many Peruvian specimens examined. Amazonas: Quebrada Huampami, Kayap 1066 (MO). San Martin: Prov. San Martin, Laguna Sauce, Ramirez & Sotero 092-85 (F, GH, HUT). Loreto: Rio Tacsha Cur- aray, Croat 20394 (F, GH, MO, NY). Gamitanacocha, Rio Mazan, J. Schunke 234 (F, GH, NY, uc, us). Madre de Dios: El Pilar, Lopez & Soukup 4601 (GH). 3. Lygodium radiatum Prantl, Unters. Morph. Gefasskrypt. 2: 66. 1881. TYPE: based on L. digitatum D. C. Eaton. Lygodium digitatum D. C. Eaton, Mem. Amer. Acad. Arts n.s. 2, 8: 217. 1860 (not Presl, 1825). LEC- TOTYPE (inferentially designated by Duek, Feddes Repert. 89: 417. 1978): Panama, Gatun, Hayes 25 (YU; isolectotype, us!). Pinna-stalks reduced to short (0.5 mm) projec- tions along the primary axis (or sometimes ob- solete). Primary pinna-branches long-stalked (1.5- 4 cm), the pinnae radiate (subpalmate), divided nearly to the base into (2)3-7 ultimate segments. Fertile ultimate segments linear-lanceolate to nar- rowly oblanceolate, 5-15 cm long and 1.5-2 cm broad, the margins serrate, glabrous, or sparsely pubescent along the midrib. Sterile segments sim- ilar to the fertile, but often somewhat longer (to 25 cm). Veins free, mostly 1- or 2-forked, prom- inulous. Rare, in wet forests, often along stream and river banks, scandent on shrubs and small trees, 135- 850 m, Amazonas, Loreto, Pasco, Ucayali. Panama; Colombia; Ecuador; Peru. 32 FIELDIANA: BOTANY Amazonas: Al lado de Huampami, Kayap 1217 (MO, us). Loreto: Santa Rosa, lower Rio Huallaga below Yu- rimaguas, Killip & Smith 28937 (F, GH, NY, us). Puerto Melendez, below Pongo de Manseriche, Tessmann 4735 (NY). Pasco: Prov. Oxapampa, west side of Cordillera de San Matias, D. Smith 2011 (F, MO). Ucayali: Prov. Co- ronel Portillo, km 89 Carretera Federico Basadre, C. Vdsquez 1 (USM). Comments Lygodium oligostachyum (Willd.) Desv., Mem. Soc. Linn. Paris 6: 205. 1827. Hydroglossum oligostachyum Willd., Sp. pi. ed. 4, 5: 81. 1810. TYPE: Plumier, Traite foug. Amer., t. 92. 1705 (based on specimens from Haiti, near Lake Miragoan). This is similar in general aspect to Lygodium venustum, but differs in the conspicuously 2-pin- nate pinnae and the strongly flexuous (zigzag) and more delicate primary and secondary pinna rach- ises. Although L. oligostachyum has been consid- ered by most authors to be confined to Hispaniola, Duek (1978) assigned to it a broad distribution (West Indies; parts of Central and South America) and cited two specimens at Prague from Peru: Poeppigand Cuming, neither of which he had ex- amined. Under L. oligostachyum he also placed L. mexicanum Presl and two varieties of L. po- lymorphyum (Cav.) HBK. (illeg.); however, plants usually identified under either name have usually turned out to be L. venustum. We have not seen the Poeppig and Cuming collections in question, but they are probably L. venustum and not L. oli- gostachyum as suggested. All specimens of the lat- ter which we have examined in various herbaria have been collected in Hispaniola, so it is unlikely to be found in Peru. dichotoma (L.) Sm. (Acrostichum dichoto- mum L.). Figure 8. Lophidium Rich., Actes Soc. Hist. Nat. Paris 1: 114. 1 792. TYPE: Lophidium latifolium Rich. = Schi- zaea elegans (Vahl.) Sw. Actinostachys Wall., Numerical list of plants in East Indies Company Museum, 1. 1829, description from R. Br., Prod. 162. 1810. TYPE: Actinosta- chys digitata (L.) Wall. (Acrostichum digitatum L.) = Schizaea digitata (L.) Sw. Plants terrestrial. Stem erect or ascending, rath- er densely provided with septate trichomes and with slender, fibrous roots. Leaves ca. 5-50 cm long, caespitose, long-petiolate, glabrous or with scattered, small trichomes, slightly to partially or wholly dimorphic, fertile ones with elongate fertile segments borne pinnately or subdigitately at the apex of laminar axes which are simple or dichot- omously branched. Lamina filiform, grasslike and scarcely or not at all foliaceous, or fusiform to flabelliform (in general outline) and foliaceous. Veins free. Sporangia crowded in 1 or more rows on the scarcely foliaceous segments (sporangio- phores). Spores ellipsoidal, monolete. Schizaea sometimes has been divided into sev- eral genera or subgenera on the basis of the more expanded, foliaceous lamina (Lophidium) or the subdigitate (Actinostachys) versus pinnate (Schi- zaea) fertile branches, as well as some differences in the gametophytes. Nevertheless, the groups are quite closely allied and are here treated as a single genus. About 30 species occur in tropical or ex- tratropical regions of both hemispheres, five of which are found in Peru. Reference TRYON, R. M., AND A. F. TRYON. 1982. Schi- zaea, pp. 76-83, in Ferns and allied plants, Springer- Verlag, New York. III. Schizaea Schizaea Sm., Mem. Acad. Roy. Sci. (Turin) 5: 419. 1793, nom. conserv. TYPE: Schizaea Key to Species of Schizaea a. Leaves with lamina obviously foliaceous, flabelliform in general outline, the divisions linear-oblong to obovate 1 . S. elegans a. Leaves not or scarcely foliaceous, but filiform and grasslike or, if flabelliform in outline, then the divisions filiform b b. Sporangiophore pinnatifid, its segments bearing sporangia in a single row on each side of the vein . .c TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 33 c. Leaves 0-2 times dichotomous, monomorphic; axis of the sporangiophore straight to slightly curved at maturity d d. Leaves 7-10 cm long, unbranched; sporangiophores with 5-7 pairs of ultimate segments 2. S. pusilla d. Leaves (12-) 15-50 cm long, simple to once or twice acutely and dichotomously branched; sporangiophores with (10-) 12-25 pairs of ultimate segments 3. S. incurvata c. Leaves 3-6 times dichotomous, dimorphic; axis of the sporangiophore strongly curved at maturity 4. S. poeppigiana b. Sporangiophore subdigitate, its segments bearing sporangia in 2 or more (sometimes indistinct) rows on each side of the vein 5. S. pennula 1. Schizaea elegans (Vahl) Sw., J. Bot. (Schrader) 1800(2): 103. 1 80 1 . Figure 8. Acrostichum elegans Vahl, Symb. hot. 2: 104, t. 50. 1791. TYPE: Trinidad, von Rohr (c). Schizaea flabellum Mart., Icon. pi. crypt. 115, /. 55. 1834. TYPE: Brazil, Prov. Rio Negro, Martins (holotype, BR; isotype, B; photo, us of B). Lophidium elegans (Vahl) Presl, Suppl. tent, pterid. 77. 1845. Lophidium flabellum (Mart.) Presl, Suppl. tent, pterid. 77. 1845. Leaves 20-80 cm long, petiole 15-65 cm long, 1-2.5 mm in diameter, subterete, obtusely angled abaxially. Lamina subcoriaceous, essentially gla- brous, flabelliform (in general outline), 1 -several times-dichotomously forked or cleft, the divisions oblong to obovate, lateral margins entire, distal ones strongly and sharply lacerate. Veins dichot- omously forked within the ultimate divisions. Sporangiophores borne along the distal margins of the lamina, pinnately branched at the tips of the marginal lacerations. Sporangia in a single row on each side of the vein. In humus on forest floor, or in sandy or rocky soil on wooded slopes and ridges, 130-2200 m, Amazonas, San Martin, Loreto, Huanuco, Pasco, Junin, Cuzco. Southern Mexico to Panama; Trinidad to Co- lombia, and south to Bolivia and Brazil. Several varieties have been recognized, based upon number and shape of laminar divisions, characters which do not appear to merit distinc- tion. Amazonas: Mendoza, Woytkowski8211 (GH, MO). San Martin: Zepelacio, near Moyobamba, Klug 3415 (F, GH, MO, NY, us). Loreto: Balsapuerto, Klug 2885 (F, GH, MO, NY, us). Huanuco: Prov. Huanuco, Tingo Maria, Tryon & Tryon 5292 (BM, GH, NY, u, uc, us, USM). Pasco: Prov. Oxapampa, Cordillera San Matias, Leon 324 (F, GH). Junin: La Merced, Hacienda Schunke, Macbride 5601 (F, us). Cuzco: Valle de Urubamba, Machu Picchu, Her- rera 3299 (us). 2. Schizaea pusilla Pursh, Fl. Amer. sept. 2: 657. 1814. TYPE: United States, New Jersey, Bur- lington Co., Quaker Bridge, Pursh? (not lo- cated). Leaves unbranched, 7-10 cm long, linear, grass- like, with expanded lamina essentially lacking, pet- iole 0.4—0.5 mm broad, rather flattened, or sub- terete at base, sterile and fertile leaves erect and straight or slightly flexuous, fertile ones apically bearing a pinnatifid, conduplicate, scarcely folia- ceous sporangiophore. Sporangiophore 0.5-0.7 cm long, with 5-7 pairs of linear segments, abundantly provided on the margins and among the sporangia with deep orange to reddish brown, flexuous tri- chomes. Sporangia in a single row of 5-7 on each side of the vein. Thus far known only from Pasco, "plant colo- nizing sandy, wet landslide; shrubland on white sandstone, 2700-2800 m." United States (New Jersey, New York); Canada (Newfoundland, Nova Scotia). Heretofore, Schizaea pusilla has been known to occur only in a few scattered locations in north- eastern North America. The recent collection in Pasco duplicates the northern plants in every fea- ture (including spores) except that the sterile leaves are straight to slightly flexuous instead of con- spicuously curling. The great disjunction in dis- tribution is at first startling until one understands the problems of encountering this very inconspic- uous fern. This and the other diminutive species of Schizaea (like those of Ophioglossaceae and Hymenophyllaceae) are very difficult to detect. Hence true distribution patterns may never come to light. It is most closely allied to S. incurvata, under which see further discussion. Less closely related is the Old World S.fistulosa Labill. and its variety australis (Gaud.) Fosb. from Chile and Ar- gentina, which differ in the lack of pubescence on 34 FIELDIANA: BOTANY FIG. 8. Schizaea elegans: a, habit; b, sporangiophore; c, sporangiophore segment, abaxial side; d, sporangiophore segment, adaxial side. (From Stolze, Ferns and fern allies of Guatemala, 1976.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 35 the sporangiophore and in the erose-lacerate mar- gins of the fertile ultimate segments. Pasco: Oxapampa, Cordillera de Yanachaga, Cerro Pajonal, Foster 9065 (F, GH, MO, us). 3. Schizaea incurvata Schkuhr, 24 Kl. Linn. Pfl.- Syst. 1: 138, t. 137. 1809. TYPE: "British Guiana, habitat in India Occidentali circa Es- sequebum," Gartner (not located). Leaves simple to 1 - or 2-forked at an acute angle (branches subparallel), mature ones ( 1 2-) 1 5-50 cm long, linear, grasslike, with an expanded lamina essentially lacking, petiole 0.6-0.8 mm in diam- eter, convex abaxially, shallowly sulcate adaxially, somewhat flattened, or terete at base, sterile and fertile leaves straight or slightly flexuous, fertile ones apically bearing a pinnatifid, conduplicate, scarcely foliaceous sporangiophore. Sporangio- phore 1.2-3 cm long, with (10-) 12-2 5 pairs of linear segments, abundantly provided on the mar- gins and among the sporangia with pale to light brown, flexuous trichomes. Sporangia in a single row of (8-) 10- 18 on each side of the vein. Thus far known from a single collection (dept. unknown) in Peru; elsewhere in open woods or clearings and savannas, in sand or sandy or rocky soil, sea level to 400 m. Surinam to Venezuela and northern Brazil; Peru. Of the many specimens seen by us in various herbaria, only one was (possibly) collected in Peru: Poeppig (NY). The label is worded simply "In Pe- ruvia, diar. Kunze, 1834." However, Kunze (Lin- naea9: 19. 1834) lists only one species of Schizaea from Peru (41. 5". dichotomd), which is S. poep- pigiana. Since there is no "Diar." number on the label of the New York sheet, the data are suspect. That the collecting site is correctly des- ignated is also problematical, for the known range of the species is somewhat remote from Peru. However, as noted in the discussion of 5. pusilla (q.v.), the grasslike appearance of several species of Schizaea masks them well in any savanna-type vegetation, so that more specimens may yet be discovered throughout the Neotropics after dili- gent search, especially in Peruvian Amazonia in sandy areas. Schizaea pusilla is similar to S. in- curvata, both in general habit and pubescence of the sporangiophore, but the two are easily sepa- rated by the characters noted in the key. 4. Schizaea poeppigiana J. W. Sturm in Mart., Fl. bras. 1(2): 181. 1859. SYNTYPES: Peru, "ad Ventanilla de Cassapillo, 1829," Poeppig (w?); "British Guiana, in montibus Canuku," Rich. Schomburgk 1189(ei). Lophidium poeppigianum (J. W. Sturm) Underw., N. Amer. fl. 16: 38. 1909. Leaves dichotomous, 12-40 cm long, dimor- phic; petiole 10-30 cm long, ca. 1 mm broad, con- vex abaxially, flattened to concave adaxially, sparsely to amply pilose. Sterile lamina circular in outline (often appearing conduplicate and fla- belliform after pressing), 5-8 times dichotomously forked, the divisions linear, scarcely or slightly foliaceous. Fertile lamina on somewhat longer pet- ioles than the sterile, 3 or 4 times dichotomous, the divisions linear, nonfoliaceous, bearing spo- rangiophores at their apices. Sporangiophores 1.5- 2 cm long, with 12-25 pairs of linear segments, amply provided on the segment margins and among the sporangia with pale to light brown, flexuous trichomes. Sporangia in a single row of 1 5-25 on each side of the vein. Thus far known in Peru only from one of the syn types. Greater Antilles; Mexico (Chiapas); Costa Rica to Venezuela and Guyana; Peru; Bolivia. 5. Schizaea pennula Sw., Syn. fil. 150, 379. 1806. TYPE: "America meridionalis ... habitat in America calidiore" (holotype, S-PA). Actinostachys pennula (Sw.) Hooker, Gen. fil., /. Ilia. 1842. Leaves ca. 1 2-50 cm long, unbranched, linear, grasslike, with expanded lamina essentially lack- ing; petiole 1-1.8 mm broad, triquetrous and the faces sulcate, often flattened apically and subterete at base, sterile and fertile leaves erect and straight or slightly flexuous, fertile ones apically bearing a subdigitate, scarcely foliaceous sporangiophore. Sporangiophore 1-4 cm long, with 6-14 linear, strongly ascending segments, these essentially with entire and glabrous margins, and with abundant, flexuous, reddish brown trichomes interspersed among the sporangia. Sporangia in 2 or more (sometimes indistinct) crowded rows on each side of the vein. 36 FIELDIANA: BOTANY In sand and sandy soil of open forests and clear- ing, sea level to 260 m, San Martin, Loreto. Costa Rica; Puerto Rico; Guadeloupe; Trinidad; Surinam to Colombia, south to Bolivia and Brazil. San Martin: Chazuta, Rio Huallaga, Klug 3986 (F, GH, MO, NY, uc, us). Loreto: San Juan, vicinity of Iquitos, Asplund 14414 (F, p, NY, us). Dist. Iquitos, Maynas, Que- brada Shushuna, Rimachi 3995 (F, NY), 4870 (NY). Family 5: GLEICHENIACEAE Gleicheniaceae (R. Br.) Presl, Reliq. haenk. 1 : 70. 1825, as Order GleicheniaeR. Br., Prod. 160. 1811, as Tribe of Filices. TYPE: Gleichenia Sm. Terrestrial. Stem long-creeping, bearing tri- chomes and/or scales. Leaves 20 cm long, erect, to over 5 m long and scandent or trailing, often forming dense thickets, circinate in vernation, monomorphic, once or several times pseudodi- chotomously branched (rarely simple or with sev- eral 2-pinnate pinnae), indeterminate, with a per- manently arrested bud at the fork of its branches, or the lamina partially pinnately branched, the rachis (and sometimes pinna-rachis) with a pe- riodically dormant bud between the last developed branches, a pair of accessory foliaceous segments sometimes present at the base of otherwise usually naked axes, in addition to stipule-like segments borne within the forks. Lamina bearing scales and/ or trichomes. Veins free. Penultimate segments pinnatisect (rarely pinnate) pectinate. Sporangia borne in exindusiate sori on the abaxial surface of ultimate segments, with a short many-rowed stalk of cells and a central to oblique or nearly apical annulus not interrupted by the stalk. Spores monolete or trilete, lacking chlorophyll, 1 20 to ca. 800 in each sporangium. The pseudodichotomous branching of leaves and the usually pectinate penultimate segments make this one of the most distinctive of fern families. These ferns frequently are found in dry, open areas and species with larger leaves sometimes spread over low shrubs to form dense thickets. Approx- imately 1 20 species have been recognized by var- ious authors in as many as eight genera. The fol- lowing treatment is based generally on the classification of Holttum (1957). Two genera are recognized in Peru. References HOLTTUM, R. E. 1957. Florae Malesianae Prae- cursores XVI. On the taxonomic subdivision of the Gleicheniaceae... . Reinwardtia, 4: 257-280. MAXON, W. R. 1909. Gleicheniaceae, in N. Amer. fl., 16: 53-63. NAKAI, T. 1950. A new classification of Glei- cheniales. Bull. Natl. Sci. Mus., 29: 1-71. UNDERWOOD, L. M. 1907. American ferns VIII. A preliminary review of the North American Gleicheniaceae. Bull. Torrey Bot. Club, 34: 243- 262. Key to Genera of Gleicheniaceae a. Axils and axillary buds with scales; veins simple to 1 -forked; sporangia 2-4(-5) per sorus I. Gleichenia a. Axils and axillary buds with trichomes; veins 2-4-forked; sporangia ca. 8-15 per sorus II. Dicranopteris I. Gleichenia Gleichenia Sm., Mem. Acad. Roy. Sci. (Turin) 5: 419. 1793. TYPE: G. polypodioides (L.) Sm. (Onoclea polypodioides L.). Figure 9. Sticherus Presl, Tent, pterid. 51. 1836. TYPE: S. lae- vigatus (Willd.) Presl (Mertensia laevigata Willd.) = Gleichenia truncata (Willd.) Sprengel. Diplopterygium (Diels) Nakai, Bull. Natl. Sci. Mus. 29: 47. 1950. Gleichenia section Diplopterygium Diels, Nat. Pflanz. 1(4): 353. 1900. TYPE: Glei- chenia glauca (Houtt.) Hooker (Polypodium glaucum Houtt.). Stem provided with setose or short-ciliate scales, and sometimes stellate trichomes. Leaves once or several times pseudodichotomously branched (but simple in G. simplex, or with several 2-pinnate pinnae in G. bancroftii), with the penultimate seg- TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 37 ments in diverging, subequal, usually pectinate 100 species. The three subgenera, Gleichenia, Di- pairs. Axils of forks bearing a dense tuft of setose plopterygium, and Mertensia, are treated as genera to ciliate (very rarely entire) scales, these often by some authors, flanked by reduced, stipule-like appendages. Lam- ina, at least abaxially, provided with setose to cil- iate scales or stellate trichomes. Veins simple to Reference 1 -forked. Sori commonly paraphysate. Sporangia 2-4(-5) per sorus. CHING, R. C. 1940. On the genus Gleichenia Smith. Sunyatsenia, 5(4): 269-288. Gleichenia is a pantropical genus of more than Key to Species of Gleichenia a. Leaves simple (or very rarely with a single fork), deeply pinnatisect to pinnate; lamina linear 2. G. simplex a. Leaves pseudodichotomously forked, or with 1-3 pairs of 2-pinnate pinnae; lamina broad, never linear b b. Pinnae 2-pinnate, not forked; axillary scales with margins entire 1 . G. bancroftii b. Pinnae once to several times pseudodichotomous; axillary scales conspicuously ciliate or setose (or entire in G. nitiduld) c c. Ultimate segments moderately to densely tomentose on midrib and/or veins abaxially, the whitish to orange tomentum often obscuring the segment tissue (sometimes thinning in age) d d. Larger penultimate segments 1.4-2.2 cm broad; larger ultimate segments 1.2 cm long or less; pinnae usually 3-4 times-forked; branches strongly ascending, usually tightly crowded and subparallel with adjacent ones 3. G. pennigera d. Larger penultimate segments 2.5-8 cm broad; larger ultimate segments 1.5-4.5 cm long; pinnae 1 - or 2-forked; branches spreading to moderately ascending, not crowded e e. Sori inframedial, crowding or touching the segment midrib; ultimate segments (larger ones) 1.8-3 mm broad beyond the dilated base, usually strongly revolute 4. G. bifida e. Sori medial, rarely crowding the midrib; ultimate segments (larger ones) (3-)3.5— 4.5 mm broad, plane to moderately revolute 5. G. tomentosa c. Ultimate segments naked to scaly abaxially, never tomentose (if trichomes present, these scattered, short, or rigid) f f. Penultimate segments less than 2 cm broad; ultimate segments 2-3.5(-4) times longer than broad g g. Scales ample to abundant on costae and veins abaxially 6. G. revoluta g. Scales essentially lacking on lamina h h. Axillary scales broad, ciliate; primary axis conspicuously tuberculate to muricate . . 7. G. tuberculata h. Axillary scales narrow and rigid, entire or (rarely) sparingly dentate or short-setose; primary axis smooth 8. G. nitidula f. Penultimate segments (at least larger ones) 2.2-8 cm broad; ultimate segments (4-)5-12 times longer than broad i i. Midribs of ultimate segments naked 9. G. lechleri i. Midribs of ultimate segments conspicuously scaly, at least abaxially j j. Ultimate segments (most of them) remote, separated by a space once or twice their width; sori medial to supramedial 10. G. remota j. Ultimate segments approximate, contiguous at their bases; sori commonly inframedial (or some of them medial) k k. Costa scales commonly less than 1 mm long; midrib scales deltoid, the scale body 5 or more cells wide; axillary scales 0.5-1.5 mm long, with whitish or pale orange, lax cilia 11. G. peruviana 38 FIELDIANA: BOTANY k. Costa scales 1.5-3 mm long; midrib scales filiform, the scale body 1-3 cells wide; axillary scales 2-3 mm long, with dark, rigid setae 1 1. Penultimate segments 4-9 cm broad; margins of ultimate segments plane to slightly revolute; veins not or slightly raised 12. G. longipinnata 1. Penultimate segments 2.2-3.2 cm broad; margins of ultimate segments mod- erately to strongly revolute; veins strongly raised abaxially . . 1 3. G. rubiginosa 1. Gleichenia bum-mini Hooker, Sp. til. 1: 5. 1844. TYPE: Jamaica, Bancroft (holotype, K). Mertensia bancroftii (Hooker) Kunze, Linnaea 18: 307. 1844. Dicranopteris bancroftii (Hooker) Underw., Bull. Tor- rey Bot. Club 34: 252. 1907. Hicriopteris bancroftii (Hooker) Ching, Sunyatsenia 5: 278. 1940. Diplopterygium bancroftii (Hooker) A. R. Smith, Amer. Fern J. 70: 26. 1980. Leaves with 1-3 pairs of 2-pinnate pinnae, these to 1.5 m long and 40 cm broad, the axes naked or with scattered, linear, or filiform scales. Axillary scales lanceolate to ovate, long-acuminate to at- tenuate, yellowish to light brown, with entire mar- gins. Penultimate segments (pinnules) very nu- merous, crowded, at nearly right angles to the costa, pectinate, cut nearly or quite to the costa. Ultimate segments strongly revolute, midribs abaxially often provided with scattered, filiform scales. Sori in- framedial. In open forests, commonly on slopes or ravine banks, 2400-2800 m, infrequent in Peru: Huan- uco, Pasco, Cuzco. Mexico to Panama; West Indies; Colombia; Venezuela; Ecuador; Peru; Bolivia. Huanuco: Huanuco, within 5 km of Carpish, Tryon & Tryon 5316 (F, OH, uc, us). Pasco: Border Prov. Oxa- pampa and Pasco, van der Werffet al. 8604 (MO). Cuzco: La Convencion, Valle de San Miguel, Biies 2126 (us). 2. Gleichenia simplex (Desv.) Hooker, Icon, pi., /. 92. 1837. Mertensia simplex Desv., Dictionaire sciences natu- relles 62(2), /. 91. 1827; Mem. Soc. Linn. Paris 6: 200. 1 827. TYPE: Peru, collector unknown (ho- lotype, P; photo, us). Dicranopteris simplex (Desv.) Maxon, Contr. U.S. Natl. Herb. 24: 50. 1922. Sticherus simplex (Desv.) Ching, Sunyatsenia 5: 285. 1940. Leaves simple (very rarely with a single fork), deeply pinnatisect to pinnate, linear, 20—50 cm long and less than 3 cm broad. Scales of the petiole essentially lacking, but abundant on the rachis abaxially, these lanceolate, attenuate, orange to reddish brown, the margins long-ciliate. Ultimate segments contiguous at the very base or (especially proximal ones) discrete and almost their own width apart, strongly revolute, on the abaxial side slightly to conspicuously pruinose and scaly, scales along the midrib similar to those of the rachis, but to- ward the segment apex grading to pluricellular, stellate trichomes. Sori medial to inframedial. Road cuts, banks and open rocky slopes, (1900-)2700-3800 m, Piura to Cajamarca, south to Huancavelica and Cuzco. Colombia to Bolivia. There has been some confusion as to publication of the name Mertensia simplex. In the Diet. Sci. Nat. 1 827 there is no text, but the plate with anal- ysis constitutes a valid publication. A description was given at approximately the same time by Des- vaux in Mem. Soc. Linn. Paris 6: 200. 1827. Piura: Prov. Huancabamba, road to Canchaque, Hutchison 1617 (F, GH, NY, uc). Lambayeque: Dist. In- cahuasi, Laguna Tembladera, Sagdsteguiet al. 12774 (F, HUT, MO). Cajamarca: Prov. San Miguel, El Tingo, Sa- gdstegui et al. 9517 (F, MO, NY). Amazonas: Prov. Chach- apoyas, banks above swamp on summit of Cerros de Calla-Calla, Wurdack 1212 (F, GH, NY, us). La Libertad: Prov. Pataz, Huaylillas, Puerta del Monte, Lopez & Sa- gdstegui 3457 (GH). San Martin: Dist. Huallaga, Valley of Rio Apisoncho, Hamilton & Holligan 540 (us). An- cash: Prov. Huari, Huascaran National Park, Quebrada Pachachaca, D. Smith et al. 12564 (F, MO). Huanuco: Yanano, stony clay bank, Macbride 4925 (F, GH, us). Huancavelica: Prov. Tayacaja, Chuspi-Tocas, between Colcabamba and Paucarbamba, Tovar 2066 (GH, USM). Cuzco: La Convencion, Biies 2078 (GH, us). 3. Gleichenia pennigera (Mart.) Moore, Index fil. 381. 1862. Mertensia pennigera Mart., Icon. pi. crypt. 130, t. 59, f. 1. 1834. TYPE: Brazil, "Prov. Minarum, in Serra de S. Geraldo," Martius (M; photo, us). Dicranopteris pennigera (Mart.) Maxon, Contr. U.S. Natl. Herb. 24: 48. 1922. Sticherus penniger (Mart.) Copel., Gen. fil. 27. 1947. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 39 FIG. 9. Gleichenia bifida: a, habit; b, leaf axil with bud; c, ultimate segment, (a from Schunke V. 5976, F, b from Tryon & Tryon 6605, Brazil, F, c from Luteyn & Luteyn 6735, Ecuador, F.) 40 FIELDIANA: BOTANY Leaves pseudodichotomously forked. Pinnae (2-)3— 4-forked, the branches (at least in Peruvian specimens) strongly ascending, often tightly crowded and subparallel with adjacent ones. Ax- illary scales lanceolate, orange, the margins co- piously beset with long, tortuous pale or whitish cilia. Larger penultimate segments 1.4-2.2 cm broad, the costae abaxially provided with a mix- ture of orange or whitish long-ciliate scales and a scattered whitish tomentum, adaxially naked or sparsely to amply white-tomentose. Ultimate seg- ments slightly to strongly revolute, larger ones to 1.2 cm long, densely covered abaxially with whit- ish tomentum (often thinning in age). Sori infra- medial, mostly crowding or touching the midrib. Open places in forests, on slopes or stream banks, 1000-2100 m, Loreto?, Huanuco, Pasco, Junin, Cuzco. Colombia; Venezuela; Peru; Bolivia; Brazil. Special note should be made of the branching pattern throughout the range of this species. Branches may be very strongly ascending, thus quite crowded and even subparallel with each oth- er; or they may diverge at broader angles, as seen in the type. Thus far, all specimens examined from Peru exhibit the crowded pattern, which might tempt one to consider separation at subspecific level. However, this difference appears uncorre- lated with other characters, and specimens, at least outside Peru, seem to grade freely from one pattern to the other. Loreto?: " Altura de Salarayacu" (Sarayacu?), Biies 847 (us). Huanuco: Between Huanuco and Pampayacu, Ka- nehira 153 (GH, us). Pasco: Prov. Oxapampa, Gran Pa- jonal, D. Smith 5073 (F, MO). Junin: Concepcion, Her- rera 142 (us). Cuzco: Valle de Lares, Hda. Huy-huy, Biies 1826 (us). Prov. Convention, Lucumayo, Vargas 4221 (us). 4. Gleichenia bifida (Willd.) Sprengel, Syst. veg. 4: 27. 1827. Figure 9. Mertensia bifida Willd., Kongl. Vetensk. Acad. Nya Handl. 25: 168. 1804. TYPE: Venezuela, Czn- cas,Bredemeyer(ho\olype,B,Herb. Willd. 19468; photo, GH). Dicranopteris bifida (Willd.) Maxon, N. Amer. fl. 16: 60. 1909. Gleichenia mathewsii Hooker, Sp. fil. 1 : 9. 1 844. TYPE: Peru, Mathews 1092, in part (holotype, K; photo, us). Mertensia mathewsii (Hooker) Fee, Mem. Foug. 1 1: 122. 1866. Sticherus bifidus (Willd.) Ching, Sunyatsenia 5: 282. 1940. Sticherus mathewsii (Hooker) Nakai, Bull. Nat. Sci. Mus. 29: 22. 1950. Leaves pseudodichotomously forked. Pinnae 1- or 2-forked, the branches spreading to moderately ascending. Axillary scales lanceolate to ovate, at- tenuate, light brown to pale orange, the margins amply ciliate. Larger penultimate segments (2.5-)3- 7 cm broad, the costa on the abaxial side amply to abundantly provided with orange, long-ciliate scales, on the adaxial naked or sometimes with sparse whitish tomentum. Ultimate segments moderately to (typically) strongly revolute, larger ones 1 5-35 mm long and 1 .8-3 mm broad (beyond the dilated base), densely covered abaxially with orange to whitish tomentum (sometimes thinning in age), the midribs also sparsely provided with tawny to whitish, filiform, ciliate scales. Sori in- framedial, mostly crowding or touching the mid- rib. In forests, thickets and clearings, often on banks and slopes, 150-1800 m (very rarely to 2200 m) Piura to Loreto, south to Madre de Dios and Puno. Mexico to Panama; West Indies; Colombia and Venezuela south to Bolivia and Paraguay. Piura: Prov. Huancabamba, slopes of Cerro La Viuda, Sagdsteguiet al. 8 209 p.p. (NY). Antazonas: Prov. Bagua, along roadside, 37 km NE of Chiriaco, Barbour 4486 (MO). San Martin: Zepelacio, near Moyobamba, Klug 3458 (F, GH, MO, NY, us). Loreto: Pumayacu, between Balsapuerto and Moyobamba, Klug 3242 (F, GH, MO, NY, us). Huanuco: Dist. Churubamba, trail Cotirarda to Mercedes, Mexia 8191 (F, GH, MICH, MO, NY, uc, us). Pasco: Prov. Oxapampa; Palcazii, Rio Alto Iscozacin, Foster & d'Achille 10098 (F). Junin: Huacapistana, be- tween Tarma & San Ramon, Ferreyra 286 (GH, USM). Ucayali: Vicinity of Aguaytia, on steep slopes along Rio Aguaytia, Croat 21003 (F, MO, uc). Ayacucho: Ca. 25 km SW from Hacienda Luisiana and Rio Apurimac, ca. 25 km from Tambo, Dudley 11887 (GH). Cuzco: Prov. Paucartambo, near Asuncion, West 7122 (MICH, uc). Madre de Dios: Prov. Manii, Rio Inambari, Chavez 1057 (MO). Puno: San Gaban to Ollachea, Dillon et al. 1243 (F, MO, NY). 5. Gleichenia tomentosa (Sw.) Sprengel, Syst. veg. 4: 27. 1827. Mertensia tomentosa Sw., Kongl. Vetensk. Acad. Nya. Handl. 25: 1 77, /. 5J. 4. 1 804. TYPE: Peru, Herb. Cavanilles (holotype, s; isotype, p?; photos, us of S&P). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 41 Mertensia velata Kunze, Linnaea 9: 15. 1834. TYPE: Peru, Huanuco, Pampayacu, July 1829, Poeppig (diar. 1117) (holotype, B?; isotype, M!; probable isotype, P!; frag., us!; photos, us of M & p). Gleichenia velata (Kunze) Mett., Fil. hort. bot. Lips. 113. 1856. Gleichenia buchtienii Christ & Rosenst., Repert. Spec. Nov. Regni Veg. 5: 229. 1908. TYPE: Bolivia, "Yungas australis, Sirupaya prope Yanacachi," Buchtien 496 (holotype, P!; isotypes, F!, P!, s). Dicranopteris velata (Kunze) Maxon, Contr. U.S. Natl. Herb. 24: 50. 1922. Sticherus buchtienii (Christ & Rosenst.) Copel., Gen. fil. 28. 1947. Sticherus velatus (Kunze) Copel., Gen. fil. 28. 1947. Leaves pseudodichotomously forked. Pinnae 1 - or 2-forked, the branches spreading to moderately ascending. Axillary scales lanceolate to ovate, at- tenuate, deep to pale orange, or castaneous at base, the margins amply long-ciliate. Larger penultimate segments 4-7 cm broad, the costa abundantly pro- vided abaxially with orange, long-ciliate scales, and adaxially with orange to whitish filiform scales which often grade into a pale tomentum (this sometimes caducous). Ultimate segments plane to moderately revolute, larger ones 20—45 mm long and (3-)3.5-4.5 mm broad above the dilated base, densely covered abaxially with orange to whitish tomentum (this sometimes thinning in age), the midribs provided also with orange, ciliate scales. Sori mostly medial, rarely, if ever, crowding the midrib. In cloud or elfin forests, usually in open sites on ridges, slopes or ravine banks, 1 500-2700 m, Ca- jamarca, Amazonas, San Martin, Huanuco, Pasco, Cuzco. Colombia; Venezuela; Ecuador; Peru; Bolivia. Much confusion has attended circumscription of the densely tomentose species of Gleichenia, which include G. bifida as well as those combined here under G. tomentosa. We have been unable to obtain the type specimens of the latter, but type photos (us) indicate clearly enough that ultimate segments are too broad and soral lines appear too far from midribs to be G. bifida. The photos, along with Swartz's original description ( 1 804) of Mer- tensia tomentosa and his subsequent Latin de- scription (Syn. fil. 164, p. 392, 1806) make it ev- ident that G. tomentosa and G. velata are the same. All this confusion within the species complex is warranted, for G. bifida and G. tomentosa do not differ strongly. As seen in the key, the former has narrower, more strongly revolute ultimate seg- ments, and soral lines usually are closer to the midrib; also the plants seem to occur at lower elevations in Peru: 150-1800 m (rarely to 2200 m) as compared with 1 500-2700 m in G. tomen- tosa. Color of scales on the buds and major axes affords another, but less consistent, diagnostic fea- ture; in both species these are pale to deep orange, but in G. tomentosa they are often liberally inter- mixed with castaneous ones. A few specimens have been seen which are intermediate, thus emphasiz- ing the close relationship of the two species. For example, the only collection of G. tomentosa (cited below) from Cajamarca has segments strongly re- volute and 3 mm broad or less, but sori are medial and scales frequently are castaneous at base. Cajamarca: Prov. Cutervo, La Achira (San Andres- Socota), growing over bushes, Lopez & Sagdstegui 5464 (GH, HUT). Amazonas: Prov. Bagua, Cordillera Colan SE of La Peca, Barbour 3980 (MO). San Martin: "In monte Campana prope Tarapoto," Spruce 4707 (p, us). Huan- uco: Playapampa, steep banks, Macbride 4509 (F, us). Pasco: Cushi, trail to Tambo de Vaca, Bryan 687 (F). Cuzco: La Convention, ca. 12 km NE from Hacienda Luisiana and Rio Apurimac, Dudley 10545 (GH, us). 6. Gleichenia revoluta HBK., Nov. gen. sp. 1: 29. 1815. TYPE: Ecuador, Saraguru, Humboldt & Bonpland (holotype, p; isotype, B!; frag., us of B; photos, us of B & p). Mertensia revoluta (HBK.) Desv., Mem. Soc. Linn. Paris 6: 200. 1827. Mertensia pruinosa Mart., Icon. pi. crypt. 109. 1834. TYPE: Brazil, Minas Gerais, Freyreiss (probable holotype, M!; isotype, s; photos, F & us of M). Gleichenia pruinosa (Mart.) Mett., Ann. Mus. Bot. Lugduno-Batavum 1: 49. 1863. Gleichenia affinis Kuhn, Linnaea 36: 167. 1869. TYPE: Peru, Dept. Puno, San Gaban, Lechler 2265 (ho- lotype, B!; photo, us). Dicranopteris affinis (Kuhn) Maxon, Contr. U.S. Natl. Herb. 24: 47. 1922. Dicranopteris pruinosa (Mart.) Maxon, Contr. U.S. Natl. Herb. 24: 49. 1922. Sticherus pruinosus (Mart.) Ching, Sunyatsenia 5: 284. 1940. Sticherus revolutus (HBK.) Ching, Sunyatsenia 5: 285. 1940. Sticherus affinis (Kuhn) Nakai, Bull. Natl. Sci. Mus. 29: 13. 1950. Leaves pseudodichotomously forked. Pinnae 2- 4-forked. Axillary scales lanceolate to ovate, ±lax, thin-textured, orange to reddish brown, the mar- gins amply ciliate. Larger penultimate segments I - 1.8(-2) cm broad, the costa on the abaxial side amply to copiously provided with orange to red- dish brown, long-ciliate scales, on the adaxial side 42 FIELDIANA: BOTANY the costa naked or filiform-scaly. Ultimate seg- ments moderately to strongly revolute, larger ones 2-3.5(-4) times longer than broad, the midrib and veins scaly as on the costa, those of the veins often filiform or grading to tortuous, stellate trichomes, the veins not or scarcely raised. Sori medial to supramedial. In elfin forests, clearings, and along roadsides, 1 770-3400 m, Piura to Amazonas, south to Cuzco and Puno. Costa Rica & Panama; Colombia; Venezuela; Ecuador; Peru; Brazil; Bolivia. This species is locally abundant in middle to upper elevations and is sometimes confused with Gleichenia bifida, probably because of the often copious indument borne on the abaxial surfaces. These scales, especially on the veins, often become filiform and even grade into stellate trichomes; however, the surfaces are never truly tomentose, as in G. bifida. Also, penultimate segments of the latter are much broader, with linear ultimate seg- ments, and the sori crowd the midrib, whereas in G. revoluta sori are medial to supramedial, most of them quite remote from the midrib. Further- more, G. bifida grows at lower elevations. It is likely that G. revoluta occurs also in Costa Rica and Panama [as G. costaricensis (Underw.) C. Chr.] and in Bolivia [as G. boliviensis(Maxon & Morton) Lell.]. Probably G. subandina Sodiro and G. hy- poleuca Sodiro of Ecuador are also synonymous with G. revoluta. Piura: Prov. Huancabamba, above Huancabamba, road to Canchaque, Hutchinson 1618 (F, OH, uc, us). Caja- marca: Prov. Cutervo, alrededores de Cutervo, Ldpez & Sagdstegui 5314 (GH, MO). Amazonas: Prov. Chacha- poyas, slopes of Puma-urcu SE of Chachapoyas, Wur- dack 552 (F, GH, NY). La Libertad: Prov. Pataz, Puerta del Monte, Paso La Sabana, Lopez & Sagdstegui 3463 (GH). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8477 (MO). Huanuco: Playapampa, in sphagnum montana, Macbride 4508 (F, GH, NY, us). Cuzco: Cordillera Vilcabamba, 23 km NE from Hacienda Luisiana, Dudley 11133 (GH, us). Puno: Prov. Sandia, near Limbani, Metcalf 30539 (MO, us; atypical in narrower segments and darker, narrower scales). 7. Gleichenia tuberculata Kuhn, Linnaea 36: 166. 1869. TYPE: Peru, Dept. Puno, Tatanara, Lechler2572 (holotype, B!; isotype, B!; photos, F & MICH of B). Sticherus tuberculatus (Kuhn) Nakai, Bull. Natl. Sci. Mus. 29: 30. 1950. Leaves pseudodichotomously forked, petiole and rachis conspicuously tuberculate to muricate. Pin- nae 2-3-forked. Axillary scales closely imbricate, ovate to lanceolate, castaneous to reddish brown, the margins amply ciliate. Larger penultimate seg- ments 1 .3-1.5 cm broad, pruinose abaxially, scales essentially lacking. Ultimate segments moderately revolute, larger ones ca. 3 times longer than broad. Sori supramedial. Thus far represented only by the type collection, from Puno. This species appears to differ from Gleichenia revoluta only in the lack of scales on the laminar surface and in the tuberculate to mur- icate primary axis. However, the specimens were taken from a plant apparently quite advanced in age, for few sporangia remain on the segments, and it is possible that any scales once borne on the axes may have fallen away as well. Density of indument varies within some species of Gleichen- ia, depending upon maturity. Furthermore, tu- berculate primary axes are not uncommon in the genus, although never so conspicuously so as in G. tuberculata. The species is maintained as dis- tinct here, but with reservations. 8. Gleichenia nitidula Rosenst., Repert. Spec. Nov. Regni Veg. 10: 275. 1912. TYPE: Costa Rica, San Carlos, Brade & Brade 503 (holotype, not located; isotypes, NY, s). Sticherus nitidulus (Rosenst.) Copel., Gen. fil. 28. 1 947. Leaves pseudodichotomously forked, petiole and rachis not or scarcely tuberculate. Pinnae 1-3- forked. Axillary scales not or slightly imbricate, linear or linear-lanceolate, with subcaudate tips, deep reddish brown, the margins entire or (rarely) sparingly dentate or short-setose. Larger penulti- mate segments 1.4-1.8 cm broad, not or scarcely pruinose abaxially, scales lacking. Ultimate seg- ments slightly revolute, larger ones ca. 3 times longer than broad. Scandent on wet, shady banks, in forests, 1 600- 1900 m, San Martin, Pasco. Costa Rica; Panama; Ecuador, Peru. The species is notable for its disjunct distribu- tion, and has rarely been collected in South Amer- ica. Distinctive characteristics are the lack of scales on the narrow penultimate segments, and the sparse, rigid axillary scales with entire margins. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. Apparently represented thus far in Peru by the follow- ing two collections: San Martin: "Prope Tarapoto, in monte Guayrapurima," Spruce 4018 (K). Pasco: (as Jun- in) Pichis Trail, Enenas, dense forest, Killip & Smith 25777 (F). 9. Gleichenia lechleri Kuhn, Linnaea 36: 167. 1869. TYPE: Peru, "Tabina," Lechler 2040 (holotype, B!; isotypes, K, L; frag., us!; photos, F, GH, us of B). Gleichenia yungensis Rosenst., Repert. Spec. Nov. Regni Veg. 5: 228. 1908. TYPE: Bolivia, Yungas, Unduavi, 3300 m, Buchtien 902 (holotype, s; is- otypes, uc!, us!). Dicranopteris yungensis (Rosenst.) Maxon, Contr. U.S. Natl. Herb. 24: 50. 1922. Sticherus yungensis (Rosenst.) Copel., Gen. fil. 28. 1947. Sticherus lechleri (Kuhn) Nakai, Bull. Natl. Sci. Mus. 29: 21. 1950. Leaves pseudodichotomously forked, with fo- liaceous, often pinnatifid, appendages borne with- in each axil (especially proximally). Pinnae 1-3- forked. Axillary scales ovate to linear-lanceolate, orange to reddish brown, attenuate, the margins short-ciliate. Penultimate segments (larger ones) 2.5-6 cm broad, costae sparsely to amply scaly abaxially, the scales orange, linear or linear-lan- ceolate, long-attenuate, the margins somewhat short-ciliate. Ultimate segments plane or slightly revolute, commonly pruinose abaxially, larger ones 12-28 mm long and 2.5-4.5 mm broad, the mid- ribs lacking scales. Sori commonly supramedial. In mountain forests, commonly on exposed ridges, (1 1 00-) 1 800-3000 m, Huanuco, Cuzco. Trinidad; Colombia; Ecuador; Peru; Bolivia. This species is characterized by lack of midrib scales, commonly pruinose lamina, plane or slight- ly revolute ultimate segments, and the suprame- dial sori. With it should be included Dicranopteris brittonii Maxon of Trinidad, and probably also G. leucocarpa Sod. of Ecuador. Although we have not seen the type of the latter, a type fragment and photo (us) reveal no significant differences be- tween the two species. Huanuco: Carpish, Coronado 78 (us). Cuzco: Machu Picchu, Valle del Urubamba, Herrera 3286 (us). Cerro de Cusilluyoc, forest along Rio Pilahuata, Pennell 13940 (F, us). Prov. Paucartambo, "Pillawata," Yanamayo- Tambomayo, Vargas 16702, 16705 (GH). 10. Gleichenia remota (Kaulf.) Sprengel, Syst. veg. 4: 27. 1927. Mertensia remota Kaulf., Enum. fil. 39. 1824. TYPE: Brazil, Ilha de Sta. Catarina, Chamisso (holotype, LZ destroyed; isotype, LE?). Dicranopteris remota (Kaulf.) Maxon, Contr. U.S. Natl. Herb. 24: 50. 1922. Leaves pseudodichotomously forked, with fo- liaceous appendages usually borne within each axil (especially proximally). Pinnae 1 - or 2-forked. Ax- illary scales lanceolate to ovate, attenuate, rigid, 1-1.5 mm long, often somewhat convex, casta- neous to reddish brown, lustrous, not or slightly imbricate, their margins short-setose, or rarely en- tire or with a few short cilia at base. Penultimate segments (larger ones) 4-9 cm broad, costae abax- ially provided with scattered castaneous or dark reddish brown scales less than 1 mm long, these ovate to lanceolate, with setose to short-ciliate margins, adaxially virtually naked (or occasionally with scattered patches of arachnoid scurf). Ulti- mate segments remote, most of them separated by a space once or twice their width, slightly to strong- ly revolute, not or slightly pruinose abaxially, larg- er ones 20-50 mm long and 1.7-2.5 mm broad, the midribs amply to copiously scaly on abaxial side, the scales deltoid, orange to tawny, with long, pale cilia on the margins, or toward segment tips sometimes becoming substellate. Veins often raised. Sori medial to supramedial. At edges of forests, roadsides, and on rocky banks, 350-2000 m, Amazonas, San Martin, Huanuco. Costa Rica; Panama; Cuba; Trinidad; Colombia to the Guianas; Ecuador; Peru; Bolivia; Brazil (Amazonas, Ilha Sta. Catarina). This and Gleichenia longipinnata are superfi- cially similar in the great size of their penultimate segments (sometimes to 50 cm long and 9 cm broad) and the extremely long and narrow ultimate segments. However, besides the characters noted in the key, G. remota can also be sharply distin- guished by the laminar scales. Those within the axils are relatively short (1-1.5 mm) and broad, while the costal scales are even shorter (less than 1 mm) and broader. In G. longipinnata, both the axillary scales and costal scales are linear to fili- form, the former 2-3 mm long and the latter 1.5- 2 mm long. 44 FIELDIANA: BOTANY Ama/onas: Prov. Bagua, roadside, 37 km NE of Chi- riaco, Barbour 4485 (MO), 4486 (USM). San Martin: Tar- apoto-Yurimaguas Hwy., km 39, McDaniel 14209 (GH, MO). Tarapoto, Carretera Tarapoto-Yurimaguas, Ri- machi 5163, 5251 (MO, NY). Huanuco: Monzon, con- fluencia con Huallaga, cerca de Tingo Maria, Ferreyra 10047 (GH). 11. Gleichenia peruviana (Maxon) 1 ell.. Amer. Fern J. 74: 57. 1984. Dicranopteris peruviana Maxon, Amer. Fern J. 33: 133. 1943. TYPE: Peru, Huanuco, Playapampa, ca. 2700 m, Macbride 4510 (holotype, F!; isotype, us!). Leaves pseudodichotomously forked, with fo- liaceous appendages often borne within some of the more proximal axils. Pinnae 1-3-forked. Ax- illary scales ovate or lance-ovate, attenuate, 0.5- 1.5 mm long, castaneous to reddish brown, closely imbricate, their margins amply ciliate, the cilia lax, short and whitish to pale orange. Penultimate seg- ments (larger ones) 2.5^4(-5) cm broad, costae on abaxial side amply to copiously scaly, the scales lustrous castaneous to reddish brown, ovate to lan- ceolate, acuminate or attenuate, commonly less than 1 mm long, their margins short-ciliate, costae on adaxial side sparsely to amply provided with tortuous trichomes or filiform scales. Ultimate seg- ments crowded, contiguous at their bases, slightly to moderately revolute, not or slightly pruinose, larger ones 1 2-24 mm long and 1 .5-2.5 mm broad (beyond the dilated base), abaxially the midribs and (usually) veins minutely scaly, ciliate- or se- tose-scaly, the scales often becoming filiform or grading into stellate, whitish trichomes. Veins often raised. Sori mostly inframedial. In thickets, open forests and ravines, often on exposed rocky ridges, 2000-3400 m, La Libertad, Huanuco, Pasco, Cuzco. Apparently confined to Peru, Cordilleras Cen- tral and Oriental. La Libertad: Prov. Pataz, Pampa de Huayno-huincho, Huaylillas, Ldpez & Sagdstegui 3517 (GH, HUT). Huan- uco: Wet, dense jungle, 12 mi S of Panao, Macbride & Featherstone 2217 (F). Pasco: Prov. Oxapampa, Dist. Oxapampa, Rio San Alberto, Foster et al. 10306 (F); Leon 641 (F, GH); D. Smith & Pretel 7594 (F, MO). Cuzco: Prov. La Convention, on open exposed ridge, Dudley 10710 (GH, MO). 12. Gleichenia longipinnata Hooker, Sp. til. 1: 9. 1844. TYPE: Surinam, Hostmann 238 (ho- lotype, K; isotypes, BM, K; frag., us!; photo, us ofK). Mertensia longipinnata (Hooker) Klotzsch, Linnaea 18: 537. 1844. Dicranopteris longipinnata (Hooker) Maxon, Contr. U.S. Nail. Herb. 24:48. 1922. Sticherus longipinnatus (Hooker) Nakai, Bull. Nat. Sci. Mus. 29: 21. 1950. Leaves pseudodichotomously forked, occasion- ally bearing foliaceous appendages within the ax- ils. Pinnae commonly 1 -forked. Axillary scales linear to filiform, 2-3 mm long, lustrous reddish brown to deep orange, not or rarely tightly im- bricate, the margins with dark, rigid, ascending setae. Penultimate segments (larger ones) 4-9 cm broad, costae on abaxial side amply to abundantly scaly, the scales 1.5-2 mm long, lustrous reddish brown, setose and filiform or substellate, on adax- ial side often appressed filiform-scaly. Ultimate segments approximate, contiguous at their bases (very rarely some proximal ones discrete and somewhat apart), plane to slightly revolute, not or slightly pruinose, larger ones 25-45 mm long and 2.5-3.5 mm broad, abaxially bearing filiform or substellate scales on midrib and veins and often some pale, delicate trichomes at the very margin. Veins not or slightly prominulous. Sori mostly raised. In forests, on slopes and ridges, 300-1600 m (reported on Dudley 10448 label as also seen at 3300 m, but probably mistaken for G. peruviana), Amazonas, Loreto, Pasco, Huanuco, Cuzco. Surinam; Peru; Brazil. Despite obvious differences, this and Gleichenia rubiginosa share a distinctive type of laminar scale not found in other species in Peru. The scales along the costae of penultimate divisions, abaxially, are typically long and narrow, the marginal cell walls deep reddish brown and bearing rigid dark setae. Along midrib and veins, these grade into filiform scales often only two to three cells wide (the setae then much longer than the width of the scale body) and thence into delicate, pluricellular trichomes. The Costa Rican G. mellifera Christ also bears this kind of laminar scale, and although we have not seen the type, it probably belongs here. Amazonas: Prov. Bagua, valley of Rio Maranon above Cascadas de Mayasi, Wurdack 2049 (F, NY, uc, us). Huanuco: SW slope of Rio Llullapichis watershed, Dud- ley 13193 (GH). Ucayali: Prov. Coronel Portillo, Padre Abad, Chacra de Cesar Vela, J. Schunke 5465, (F, GH, MO, NY, us). Pasco: Prov. Oxapampa, Valle de Palcazii, TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 45 Foster 4504 (F). Cuzco: Prov. Convention, Cordillera Vilcabamba, Dudley 10448 (GH). 13. Gleichenia rubiginosa Mett., Ann. Sci. Nat. Bot. 5, 2: 267. 1864. TYPE: Colombia, Puente Nacional, 1650 m, Lindig 71 (holotype, B!; isotype, P; frag., us!). Gleichenia rubiginosa f. virescens Hieron., Bot. Jahrb. Syst. 34: 561. 1905. SYNTYPE: Peru, Matthews 1092 (us). Dicranopteris rubiginosa (Mett.) Maxon, Contr. U.S. Natl. Herb. 24: 50. 1922. Sticherus rubiginosus (Mett.) Nakai, Bull. Nat. Sci. Mus. 29: 28. 1950. Leaves pseudodichotomously forked, rarely bearing foliaceous appendages in the axils. Pinnae 1- or 2-forked. Axillary scales linear or linear- lanceolate, attenuate, 2-3 mm long, sublustrous, reddish brown, not or loosely imbricate, the mar- gins with dark, rigid, ascending setae. Penultimate segments (larger ones) 2.2-3.2 cm broad, costae on abaxial side copiously scaly, the scales 2-3 mm long, deep orange, sublustrous, lanceolate or lin- ear-lanceolate and attenuate, with setose margins, on adaxial side naked or occasionally sparsely ar- achnoid-scaly. Ultimate segments approximate, contiguous at their bases, moderately to strongly revolute, pruinose or not, larger ones 12-18 mm long, (2-)2.5-3 mm broad, abaxially bearing fili- form and setose to substellate scales on midrib and veins, and sometimes some delicate trichomes at the very margin. Veins raised abaxially. Sori inframedial or occasionally medial. In wet forests and thickets, 1 100-2850 m, Piura, Huanuco, Pasco, Ucayali, Cuzco, Puno. Colombia; Venezuela; Ecuador, Peru. Although easily distinguished from G. longipin- nata (q.v.), this is apparently closely related to the latter by virtue of the distinctive laminar scales. Piura: Prov. Huancabamba, Dist. Sondor, subiendo al Cerro La Viuda, Sagdstegui et al. 8209 p.p. (NY). Huanuco: Prov. Huanuco, Carpish, between Huanuco and Tingo Maria, Ferreyra 10009 (GH, USM). Huanuco- Tingo Maria road, S of Chinchayo, Gentry et al. 19322 (F, MO). Pasco: Prov. Oxapampa, road between Oxapam- pa and Paucartambo, D. Smith 1592 (F, MO). Ucayali: Prov. Coronel Portillo, Cordillera Azul, km 43 Tingo- Maria-Pucallpa Road, Young & Sullivan 737 (F, MO). Cuzco: Valle San Miguel, La Convention, Bites 2061 (us). Puno: Prov. Carabaya, San Gaban Carretera, Var- gas 18877 (GH). II. Dicranopteris Dicranopteris Bernh., Neues J. Bot. 1(2): 38. ("1806") 1805, nom. nov. for Mertensia Willd. (not Roth), and with same type. Figure 10. Mertensia Willd., Kongl. Vetensk. Acad. Nya Handl. 25:165.1 804, illeg. (not Roth 1 790). TYPE: Mer- tensia dichotoma (Murray) Willd. (Polypodium dichotomum Murray), Dicranopteris dichotoma (Murray) Bernh. = Dicranopteris linearis (Burm. f.) Underw. Stem provided with pluricellular trichomes, scales lacking. Leaves 1—4 (rarely to 20) m long, pseudodichotomously branched, with 1 -several pairs of opposite pinnae which again typically branch 1 or more times in opposite pairs, or with several pinnae which branch unequally. Axils of forks each bearing a dense tuft of trichomes and often a pair of reduced, stipule-like appendages. Lamina glabrous, or pubescent with simple or stel- late trichomes, scales lacking. Veins 2— 4-forked. Sori lacking paraphyses. Sporangia ca. 8-15 per sorus. This, like Gleichenia, is an essentially pantrop- ical genus. The two are readily distinguished by the character of their indument: that of Gleichenia being scales and trichomes, while scales are com- pletely lacking in Dicranopteris. Three species of the latter are recognized in Peru. Key to Species of Dicranopteris a. Leaves with 1 -several opposite pairs of stalked pinnae, which again branch 1 or more times; ultimate segments contiguous at base; lamina abaxially glabrous or moderately pubescent with orange, stellate trichomes (rarely tomentose) b b. Pinna-forks branching in subequal pairs; accessory, mostly reduced, leafy segments usually borne in pairs at the base of each fork (these in addition to stipule-like segments borne within the forks); spores trilete 1 . D. flexuosa b. Pinna-forks obviously branching unequally; accessory leafy segments lacking at the base of each 46 FIELDIANA: BOTANY fork (not to be confused with stipule-like segments often borne within the forks); spores monolete 3. D. pectinate a. Leaves 1 -forked, each branch consisting of a single, sessile, pinnate (to distally pinnatisect) pinna; ultimate segments mostly well-spaced, not contiguous; lamina abaxially with dense, reddish brown tomentum . 2. D. nervosa 1. Dicranopteris flexuosa (Schrader) Underw., Bull. Torrey Bot. Club 34: 254. 1907. Figure lOb-c. Mertensia flexuosa Schrader, Gott. Gel. Anz. 863. 1 824. TYPE: Brazil, Maximilian Prinz Neuwied s.n. (holotype, LZ? destroyed; isotype, M?). Mertensia rigida Kunze, Linnaea 9: 16. 1834. TYPE: Peru, "Chibangata," Poeppig 1153 (holotype, LZ destroyed; isotype, p). Gleichenia flexuosa (Schrader) Men., Ann. Mus. Bot. Lugduno-Batavum 1: 50. 1863. Gleichenia rigida (Kunze) Bommer & Christ, Bull. Soc. Roy. Bot. Belgique 35: 174. 1896, not G. rigida J. Sm. Leaves with 1-several opposite pairs of stalked pinnae, these repeatedly pseudodichotomous, with all subsequent branches forking in subequal pairs (i.e., each of the stalks subequal in length and di- verging at about the same angle), and a pair of reduced, accessory, pectinate segments commonly produced at the base of each (but rarely the ulti- mate) fork. Axils bearing a tuft of stout, rigid, castaneous trichomes and a pair of reduced sti- pule-like segments. Pinnae and their branches long- stalked, the axes and laminar surfaces glabrous, the forked, penultimate segments sessile, to 30 cm long and 6 cm broad. Ultimate segments contig- uous at base, to 30 mm long and 4 mm broad, coriaceous, glaucous abaxially, the margins strong- ly revolute. Spores trilete. On ridges and open slopes, at edges of forests, 750-2750 m, San Martin, Huanuco, Ucayali, Cuz- co, Puno. West Indies; southern Mexico to Panama, S to Brazil, Bolivia, and Paraguay (some isolated col- lections in the United States from coastal Ala- bama). This species is very closely related to Dicran- opteris linearis (Burm.) Underw. of the Paleotrop- ics. Suzanne Roth, who is working on a revision of the Neotropical Gleicheniaceae, has suggested (in litt.) that the two are probably conspecific. If so, D. linearis has priority. San Martin: NW of San Martin, Rioja, Rio Negro, Soukup 5156 (GH). Huanuco: Vilcabamba, hacienda on Rio Chinchao, Macbride 5010 (F, NY, us). Ucayali: Prov. Coronel Portillo, Obeteni, E ridge of basin, Chrostowski 66-13 (uc). Cuzco: Prov. Convention, Choquallo, Var- gas 8153 (MO, uc). Puno: Prov. Sandia, Asalaya, Vargas 14835 (OH). 2. Dicranopteris nervosa (Kaulf.) Maxon, Contr. U.S. Natl. Herb. 24: 49. 1922. Mertensia nervosa Kaulf., Enum. fil. 37. 1824. TYPE: Brazil, Sta. Catarina Is., Chamisso (holotype, not located, LZ? destroyed; isotype, LE?, s?). Gleichenia nervosa (Kaulf.) Sprengel, Syst. veg. 4: 25. 1827. Leaves 1 -forked, each branch consisting of a sin- gle, sessile, pinnate (to distally pinnatisect) pinna. Axils bearing a dense tuft of long, lax, reddish brown trichomes and usually a pair of reduced stipule-like appendages. Pinnae (larger ones) 1 5- 40 cm long, 5-12 cm broad, the axes and laminar surfaces abaxially covered with a dense reddish brown tomentum (this sometimes gray to whitish in age). Ultimate segments well-spaced, not con- tiguous at base, to 60 mm long and 3.5 mm broad, coriaceous, glaucous abaxially, the margins strong- ly revolute. Spores trilete. Thus far represented in Peru by a single collection: Puno: Region of Rio Inambari, trail from Aricoma Pass to Santo Domingo, 1800 m, McCarroll 125 (MICH). Venezuela?; Peru; Bolivia; Brazil. This is linked to Dicranopteris schomburgkiana (Sturm) Morton and D. seminuda (KJotzsch) Max- on by the tomentose abaxial surfaces. The three are usually separated by size of segments, color of tomentum, and other quantitative features; how- ever, D. nervosa appears to be further distin- guished by the delicate, somewhat flexuous axil- lary trichomes and the single pair of sessile pinnae. The related species have either stout, rigid axillary trichomes, or stalked, branching pinnae, or both. Whether these characters are sufficient to consti- tute specific or subspecific distinction is open to question. Further study of the species complex is needed. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 47 FIG. 10. Dicranopteris pectinata: a, primary axis and one pinna. Dicranopteris flexuosa: b, lamina, habit; c, axil of primary fork, (a from Belshaw 3435, F, b-c from Britton 7186, Cuba, F.) 48 FIELDIANA: BOTANY 3. Dicranopteris pectinate (Willd). Underw., Bull. Torrey Hot. Club 34: 260. 1907. Figure lOa. MertensiapectinataWMd., Kongl. Vetensk. Acad. Nya. Handl. 25: 168. 1804. TYPE: Venezuela, Cara- cas, Bredemeyer(hololype,B,Herb. Willd. 19465; photo, GH). Gleichenia pectinata (Willd.) Presl, Reliq. haenk. 1 : 71. 1825. Leaves with 1-several opposite pairs of stalked pinnae, these each bearing several to many un- equal branches which are again unequally branched or which terminate in a pair of penultimate seg- ments (i.e., each of the stalks unequal in length and diverging at much different angles), lacking accessory leafy segments at the base of the forks. Axils bearing a tuft of orange to reddish brown, curved to flexuous trichomes and often a pair of reduced stipule-like segments (at least within the proximal fork). Pinnae and their branches long- stalked, the axes commonly glabrous, the midribs and veins glabrous or, more often, sparsely to am- ply provided on the abaxial side with orange, stel- late trichomes, the forked, penultimate segments sessile, to 23 cm long and 5 cm broad. Ultimate segments contiguous at base, to 25 mm long and 5 mm broad, chartaceous, glaucous abaxially, the margins scarcely to moderately revolute. Spores monolete. Forest clearings and thickets, along roadsides and on open banks, 100-2000 m, Amazonas to Loreto, south to Cuzco and Puno. Degree of pubescence on abaxial surfaces varies greatly in this species. In Central America and the West Indies the lamina is commonly glabrous, or with a few orange, stellate trichomes scattered along the veins; but in South America, especially in Peru, some pubescence is to be expected— varying from sparse to copious— and on a few specimens we have examined it is so dense as to nearly obscure the sori. It was perhaps one of these tomentose plants on which Hieronymus based his Gleichenia (Dicranopteris) flexuosa f. monstrosa (see discus- sion below under Comments). Amazonas: Prov. Bagua, along roadside from Chiriaco to Puente Venezuela, Barbour 4488 (MO). San Martin: Prov. Lamas, Dist. Lamas, Belshaw 3435 (F, GH, MICH, MO, NY, uc, us). Loreto: Mishuyacu, near Iquitos, Klug 850 (F, NY, us). Huanuco: Prov. Huanuco, Dist. Chu- rubamba, Pampa Hermosa, Mexia 8147 (F, GH, MICH, MO, NY, uc, us). Pasco: Prov. Oxapampa, road between Puente Paucartambo and Oxapampa, D. Smith 1472 (F, MO). Junin: La Merced, thickets, Killip & Smith 23804 (F, GH, NY, us). Ucayali: Prov. Coronel Portillo, road Tingo Maria to Pucallpa, Ridoutt s.n. (GH). Cuzco: Prov. Paucartambo, near Asuncion, West 7121 (MICH p.p., MO, vcp.p.). Comments Dicranopteris flexuosa f. monstrosa (Hieron.) Na- kai, Bull. Natl. Sci. Mus. 29: 60. 1950. Gleichenia flexuosa f. monstrosa Hieron. Hedwigia 48: 289. 1909. TYPE: Peru, Amazonas, Quebra- da de Santa Lucia near Chachapoyas, Stiibel 1070 (holotype, not located). This supposed form of Dicranopteris flexuosa was described from a plant which was rusty-to- mentose on the juvenile segments of more distal pinnae. We have not seen evidence of tomentum on D. flexuosa, either on mature or juvenile plants, but it is possible this could represent a form of the species, or perhaps a hybrid. It is more likely that it belongs to the complex of tomentose species which are discussed above in the treatment of D. nervosa, particularly D. seminuda of Venezuela, which has the general aspect of D. flexuosa. Another likely possibility is that it may be a densely pu- bescent variant of D. pectinata (q.v.), although Hi- eronymus certainly should have mentioned the distinctive branching pattern of the latter if this were the case. Pending examination of the type, it is fruitless to conjecture further. Family 6: HYMENOPHYLLACEAE Hymenophyllaceae Link, Handbuch 3: 36. 1833. TYPE: Hymenophyllum Sm. Stem erect to decumbent, often slender and long- creeping, usually bearing scattered, short tri- chomes. Leaves commonly circinate in vernation, entire or pinnatifid to decompound, typically monomorphic, or rarely dimorphic in Trichom- anes, glabrous or pubescent, lamina very thin (often 1 cell thick), lacking stomates. Veins free, or in a few species reticulate. Sporangia borne in margin- al sori on a short to elongate receptacle, enclosed within a bivalvate or tubular indusium, with a short stalk and an oblique annulus not interrupted by the stalk. Spores trilete, tetrahedral-globose to spheroidal, commonly with chlorophyll. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 49 The Hymenophyllaceae are a natural and dis- tinctive family, with representatives found in trop- ical to wet temperate regions nearly throughout the world. Most of the nearly 600 species have the lamina only one cell thick, which has earned them the common name "filmy fern." The filmy leaves and the marginal sori shaped like a cup or pouch make them readily recognizable as a family, but through the years there have been great differences of opinion as to its subdivision. Various authors have split it into as many as 42 genera, whereas but two are recognized here. The term marginate is used in this family to describe an axis of the leaf that has a very narrow band of green tissue on each side. Frequently this band is scarcely raised beyond the surface of the axis and is hardly discernible except for the color contrast with the axis. It is a rudimentary wing and represents the ultimate reduction of the con- ditions elsewhere described in the text as "broadly alate" or "narrowly alate." The publications of Presl on the family have been cited according to the first publication, with the date and page number of the separately issued publication. Later publication with different pa- gination in the Abh. Konigl. Bohm Ges. Wiss. is not mentioned. References COPELAND, E. B. 1938. Genera Hymenophylla- cearum. Philipp. J. Sci., 67: 1-1 10. DIEM, J., AND J. LICHTENSTEIN. 1959. Las Hy- menofilaceas del area Argentina-Chilena del sud. Darwiniana, 11: 61 1-760. LELLINGER, D. B. 1984. Hymenophyllaceae, in Botany of the Guayana Highland— Part X. Mem. New York Bot. Gard., 38: 9-38. MORTON, C. V. 1 968. The genera, subgenera and sections of the Hymenophyllaceae. Contr. U.S. Natl. Herb., 38: 153-214. PRANTL, K. 1875. Die Hymenophyllaceen. Un- ters. Morph. Gefasskrypt., 1: 1-73. PRESL, K. B. 1843. Hymenophyllaceae, pp. 1- 70 (from Abh. Konigl. Bohm. Ges. Wiss., 5). STOLZE, R. G. 1976. Hymenophyllaceae, Ferns and fern allies of Guatemala. Fieldiana, Bot., 39: 51-90. TRYON, R. M., AND A. F. TRYON. 1982. Hy- menophyllaceae, pp. 97-1 24, in Ferns and allied plants, Springer- Verlag, New York. Key to Genera of Hymenophyllaceae a. Indusium bivalvate, the valves '/2 or more the length of the indusium (rarely less); receptacle not or rarely and slightly exserted; venation anadromous; ultimate segments sometimes serrate, lacking false veins I. Hymenophyllum a. Indusium tubular and entire, or bilabiate, rarely bivalvate (and then the valves not more than '/a the length of the indusium; receptacle commonly exserted at maturity, often strongly so; venation anad- romous or catadromous; ultimate segments not serrate, with false veins sometimes present . . II. Trichomanes I. Hymenophyllum Hymenophyilum Sm., Mem. Acad. Roy. Sci. (Tur- in) 5: 418. 1793. LECTOTYPE (designated by Presl, Hymenophyllaceae 31. 1843): Hy- menophyllum tunbridgense (L.) Sm. Figure 11. Leptocionium Presl, Hymenophyllaceae 26. 1843. TYPE: Leptocionium dicranotrichum Presl = Hy- menophyllum dicranotrichum (Presl) Sadeb. Hymenophyllum subg. Leptocionium (Presl) Christ, Farnkr. Erde 20. 1897. Sphaerocionium Presl, Hymenophyllaceae 33. 1843. TYPE: Sphaerocionium hirsutum (L.) Presl (Tri- chomanes hirsutum L.) = Hymenophyllum hir- sutum (L.) Sw. Hymenophyllum subg. Sphaerocionium (Presl) C. Chr., Index fil. Suppl. 3: 5. 1934. Mecodium(Cope\.)CopeL, Philipp. J. Sci. 67: 17. 1938. Hymenophyllum subg. Mecodium Copel., Philipp. J. Sci. 64: 93. 1937. TYPE: Hymenophyllum po- lyanthos (Sw.) Sw. Plants epiphytic, occasionally epipetric or ter- restrial. Stem filamentous, long-creeping, usually bearing scattered trichomes and small, delicate roots. Leaves monomorphic, petiolate, commonly 1-20(-50) cm long or, in H. speciosum, to nearly 2 m long. Lamina simple and entire (outside Peru) or pinnatifid to decompound, glabrous or sparsely to densely pubescent. Ultimate segments entire to 50 FIELDIANA: BOTANY FIG. 1 1. Hymenophyllum fucoides \ar.fucoides: a, habit; b, pinna with son. Hymenophyllum polyanthos: c, pinna with sori. Hymenophyllum crispum: d, pinna with sori. (a from Seller 295, El Salvador, F, b from Plowman 6074, F, c from Schunke 462, F, d from Wurdack 1510, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 51 serrate, bearing usually a single vein. Veins free (reticulate in a few Old World species), anadro- mous, false veins absent. Sori terminal on the veins. 1 ml usiiim bivalvate, the valves Vz or more the length of the indusium (rarely less), the sporangia borne on an elongate, sometimes subglobose, receptacle. Receptacle not or rarely exserted beyond the mouth of the indusium. Nearly 300 species of Hymenophyllum are found throughout tropical regions of the world, although a few occur in wet subtropical and temperate areas. While most are epiphytes, several species have been reported from wet clay or rocky banks in dense forests. A number of genera and/or subgen- era have been segregated by various authors, but in the following treatment Peruvian species are aligned into three well-defined subgenera: Hy- menophyllum, Mecodium, and Leptocionium. A good revision of the latter was done by Morton ( 1 947) under the name Sphaerocionium, but this and others still require careful study, and some of the species concepts are only provisionally treated below. Although there is no attempt here at a for- mal subgeneric classification, the species are so designated in the key and their descriptions follow in natural order. To facilitate use of the key to species, some ex- planation of the diagnostic characters may be help- ful here, especially in subg. Leptocionium. Caution is recommended when using the couplets that sep- arate those species having trichomes borne only on veins and margins of segments (Morton's sub- section Ciliata) from those in which trichomes are borne also on the segment surface between the veins and margins (his subsection Lanata). In a few species trichomes between the veins may be sparse— present only in areas toward the base of segments. Care must be exercised that these are not overlooked. Conversely, there are several species with stellate trichomes so dense on veins and margins, and with stalks so long, that they may at first appear to spring from the intervening tissue. Furthermore, certain groups of species in subg. Hymenophyllum and Leptocionium are distin- guished by the presence of lamellae: accessory green wings or crests of tissue that emerge from the veins or costae, but not on the same plane as the lamina. Typically these lamellae are made conspicuous by their size, shape, or abundance, but in H. loba- toalatum they are small and scattered, and in H. plumosum they are not at all foliose, but instead are low and inconspicuous, and usually obscured by the dense, matted tomentum covering the en- tire lamina. In H. mirificum they could be mis- taken for trichomes, since many of them, although four to six cells wide at the base, abruptly become uniseriate and grade to a filiform tip. Reference MORTON, C. V. 1947. The American species of Hymenophyllum sect. Sphaerocionium. Contr. U.S. Natl. Herb., 29: 139-201. Key to Species of Hymenophyllum a. Ultimate segments conspicuously serrate, glabrous, or rarely the serrations tipped by a short, pluri- cellular trichome [subg. Hymenophyllum] b b. Costae and veins lacking lamellae c c. Pinnae subequilateral at least beyond the base; sori (most of them) borne in the same plane as the lamina, only occasionally arcuate; petiole usually more than 0.3 mm in diameter 1 . H. fucoides c. Pinnae dimidiate (i.e., secondary segments all borne on acroscopic side); sori (most of them) strongly arcuate, thus arranged nearly perpendicular to the plane of the lamina; petiole less than 0.3 mm in diameter 2. H. peltatum b. Costae or veins (at least adaxially) bearing few to many lamellae not in the plane of the lamina, these several to many cells wide at base, becoming uniseriate toward apex d d. Petiole nonalate; rachis flexuous, nonalate, or slightly so to marginate distally; lamellae scattered and inconspicuous 3. H. mirificum d. Petiole broadly alate, often nearly to base; rachis straight, broadly alate throughout; lamellae frequent and conspicuous 4. H. lamellatum a. Ultimate segments entire, sometimes undulate to crispate, but never serrate, or rarely minutely denticulate and then the teeth tipped by unicellular or stellate trichomes e 52 FIELDIANA: BOTANY e. Lamina glabrous; ultimate segments with margins entire (although often crispate) [subg. Mecodium] f f. Leaves 1.5-4(-5) cm long; pinnae 3-6 pairs; rachis strongly flexuous to fractiflex 5. H. apiculatum f. Leaves (mature ones) 6-30 cm long; pinnae 8-30 pairs; rachis straight or slightly (rarely strongly) flexuous g g. Mature indusia ovoid, elliptical, or rhomboid, most of them somewhat to markedly longer than broad, apex obtuse to acute, base narrowly to broadly cuneate (occasionally rounded) and slightly to deeply immersed in the segment tissue; soriferous segment tips not or slightly constricted, as broad as (or slightly narrower than) mature indusia h h. Mature indusia broad- or narrow-ovoid, broadest toward the rounded or broadly cuneate base, slightly immersed in the segment tissue; receptacle filiform to narrow-cylindrical, short, not or rarely exserted i i. Petiole marginate or alate near apex or throughout (wings sometimes partly deciduous); rachis alate throughout; leaves commonly less than 15 cm long; growing at 100- 2000(-2400) m 6. H. polyanthos i. Petiole neither alate nor marginate; rachis commonly nonalate at base, at least on one side; leaves (mature ones) commonly 14-30 cm long; growing at 2100-3150 m . . . . 7. H. mathewsii h. Mature indusia rhomboid, elliptic, or conical, broadest at or beyond the middle, the base narrow-cuneate, deeply (often halfway) immersed in the segment tissue; receptacle fili- form, long, often exserted 8. H. trichomanoides g. Mature indusia subglobose, circular in outline to broader than long, the apex rounded, the base rounded or subtruncate, not or scarcely immersed in the segment tissue; soriferous segment tips somewhat to greatly constricted, narrower than the mature indusia (usually so much so that indusia appear pedicellate) j j. Sporangia (5-)6-20 per sorus; petiole marginate or alate, at least distally (wing sometimes deciduous); rachis alate throughout k k. Indusium apex erose; 1 -several elongated pinnae intermixed among normal ones . . 9. H. ferax k. Indusium apex entire; elongated pinnae rare or lacking among normal ones 10. H. myriocarpum j. Sporangia l-4(-5) per sorus; petiole neither marginate nor alate; rachis commonly non- alate toward base (or discontinuously alate on either side due to decurrent pinna bases) 1 1 . H. u lid u I a 1 11 ii i e. Lamina pubescent (at least on margins or veins), often densely so; ultimate segments entire or rarely minutely denticulate [subg. Leptocionium} 1 1. Trichomes lacking on tissue between the veins m m. Petiole slender, 0.2-0.3 mm in diameter n n. Rachis nonalate at least in the proximal portion, or weakly and irregularly alate due to the decurrent pinna bases; petiole nonalate, or weakly alate on one side due to the decurrent pinna base o o. Pinnae (at least proximal ones) short-stalked, abundantly to densely pubescent on veins and margins p p. Costae regularly and conspicuously alate throughout on both sides; pinnae reg- ularly diminishing in length toward lamina apex, larger ones with 4-9 pairs of segments 12. H. molle p. Costae partially nonalate, or discontinuously alate due to the decurrent bases of segments; pinnae often irregular in length, some of them greatly elongate, normal ones with 2-3(-4) pairs of segments 13. H. trichophyllum o. Pinnae sessile or adnate, sparsely to moderately pubescent on veins and margins . Q q. Marginal trichomes mostly forked at base or 2-forked (a few simple or stellate), petiole trichomes simple or forked; leaves determinate 14. H. elegans TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 53 q. Marginal and petiole trichomes mostly stellate; leaves indeterminate 1 5. H. adiantoides n. Rachis regularly and conspicuously alate throughout; petiole alate on both sides, at least at its apex r r. Segments essentially plane; trichomes of segment margins mostly stellate or 2-forked 16. H. hirsutum r. Segments undulate-crispate; trichomes of segment margins simple or forked from base 1 7. H. crispum m. Petiole (of mature leaves) stout, 0.4-1 .5 mm in diameter s s. Rachis conspicuously alate throughout; petiole usually alate just below lamina; trichomes lacking on adaxial side of lamina t t. Ultimate segments and rachis wing slightly to strongly undulate; stellate trichomes lacking on petiole 1 8. H. valvatum t. Ultimate segments and rachis wing not or scarcely undulate; stellate trichomes fre- quent on petiole among the simple and forked ones 19. H. microcarpum s. Rachis nonalate, at least in the proximal portion; petiole nonalate; trichomes present on axes and veins both abaxially and adaxially 20. H. ruizianum 1. Trichomes (few to many) borne on the segment surface between the veins as well as on the veins and margins u u. Veins lacking lamellae v v. Petiole slender, 0.2-0.3 mm in diameter (or sometimes to 0.4 mm in H. amabile) . . . w w. Pinnae or primary segments simple, entire; leaves to 7 cm long ...21. H. simplex w. Pinnae or primary segments lobed to pinnatisect; leaves over 12 cm long (except sometimes less in H. fragile) x x. Lamina moderately to abundantly pubescent, but veins and tissue always clearly visible y y. Rachis broadly alate throughout; leaves determinate; petiole trichomes simple to forked and often stellate 22. H. fragile y. Rachis nonalate (sometimes alate distally); leaves indeterminate; petiole tri- chomes simple or forked, not or rarely stellate 23. H. elegantulum x. Lamina densely tomentose, the veins and tissue often obscured 24. H. amabile v. Petiole (of mature leaves) stout, 0.4-1.0 mm in diameter z z. Lamina densely tomentose, the veins and tissue mostly obscured by the tomentum; rachis nonalate throughout aa aa. Trichomes on rachis subsessile or short-stalked, very tightly appressed; larger pinnae (2.5-)3-8 cm long, with 6-14 pairs of segments 25. H. speciosum aa. Trichomes on rachis short- to long-stalked, spreading; larger pinnae 1-2.3 cm long, with 4-7 pairs of segments 26. H. karstenianum z. Lamina moderately to abundantly pilose, but surfaces not obscured by the trichomes; rachis commonly alate, at least distally, only occasionally nonalate bb bb. Petiole (8-)10-15 cm long, 0.7-1.2 mm in diameter; lamina ovate to ovate- lanceolate, 8-15 cm broad, not or slightly reduced at base .... 27. H. lindenii bb. Petiole 2-7 cm long, 0.4-0.6(-0.7) mm in diameter; lamina linear, 3-8 cm broad, often strongly and gradually reduced toward base 28. H. plumieri u. Veins bearing conspicuous lamellae (or, in H. plumosum, the outgrowths low and incon- spicuous, these and the veins obscured by dense tomentum) cc cc. Rachis nonalate nearly or wholly throughout dd dd. Lamellae minute and inconspicuous, these and the veins obscured by dense to- mentum 29. H. plumosum dd. Lamellae broad and conspicuous, especially abaxially (if surface densely tomentose, most lamellae still usually visible among the trichomes) ee ee. Trichomes frequent, but not forming a dense cover; lamellae conspicuous abax- ially, sometimes less so adaxially 30. H. multialatum 54 FIELDIANA: BOTANY ee. Trichomes densely covering (sometimes obscuring) tissue and veins; lamellae sometimes lacking adaxially 31. H. tomentosum cc. Rachis alate nearly or wholly throughout ff ff. Petiole rather stout, 0.4-0.7 mm in diameter; rachis strongly and regularly alate, at least above the base; leaves 9-60 cm long; pinnae (larger ones) with 6-24 segments gg gg. Pinnae mostly narrow-triangular, narrowly acute to attenuate, larger ones 20- 45 mm long, with (6-)7-l 2 pairs of segments hh hh. Lamellae on abaxial side mostly consisting of scattered flanges, about as broad as long, lacking adaxially 32. H. lobatoalatum hh. Lamellae on abaxial side abundant, conspicuous, elongate, often nearly the length of the vein, sometimes less abundant and conspicuous adaxially . . 33. H. pyramidatum gg. Pinnae oblong to broadly triangular, obtuse or subacute, larger ones 6-20 mm long, with 3-6(-8) pairs of segments 34. H. verecundum ff. Petiole slender, 0. 1 0-0. 1 5 mm in diameter; rachis weakly and irregularly alate due to the decurrent pinna bases; leaves less than 7 cm long; pinnae with 2-4 segments 35. H. tarapotense 1. Hymenophyllum fucoides (Sw.) Sw., J. Hot. (Schrader) 1800(2): 99. 1802. Leaves 4-25(-30) cm long, 1 .5-8 cm broad. Pet- iole 0.3-0.8 mm in diameter, nonalate or margin- ate to alate distally, glabrous to sparsely or mod- erately pubescent with catenate trichomes. Lamina broadly or narrowly oblong, not or rather abruptly reduced at base, 3-4-pinnatisect. Rachis broadly to narrowly alate throughout, or nonalate just at the base, sparsely to moderately provided with flexuous, catenate trichomes. Pinnae 3-20(-25) pairs, adnate, or proximal 1-2 pairs short-stalked, subequilateral except at base, there truncate ac- roscopically, cuneate basiscopically, with 2-8 pairs of pinnatisect to 2-pinnatisect segments. Sori 1-8 per pinna, not or scarcely immersed in the segment tissue, most of them borne in the same plane as the lamina, not or only occasionally arcuate. In- dusia narrowly to broadly oblong or elliptic, the apex obtuse to subacute and entire to laciniate, receptacle narrow-cylindrical or fusiform to ovoid, not exserted or occasionally slightly so. The species occurs in the West Indies; southern Mexico to Panama; Venezuela and Colombia to Brazil and Bolivia. This species has been misunderstood, both in its nomenclature and taxonomy, since it was first recognized by Swartz in 1788. The source of the greatest confusion is a supposed type collection (BM) of Hymenophyllum cristatum Hooker & Grev. This sheet contains two specimens of//, fucoides, while pinned to it is an exact copy of/. 148, Icon, fil. 1829, the illustration of//, cristatum, complete with globose receptacle and conspicuous lamellae on the veins, which is an Ecuadorean species of a different section, Buesia (Morton) Morton. This led to another error, for the subsequent illustration H. fucoides in Hooker's "Century of Ferns" (1854) was evidently produced from this mixed sheet, thus misleading later investigators about some of the principal diagnostic features (for a full discus- sion see Stolze, Amer. Fern J. 77: 137-140. 1987). A second source of confusion has been the high- ly variable nature of the species, which has prompted the recognition of a number of related taxa at various levels over the years. Morpholog- ical features are rather constant in the West Indian specimens, where leaves are quite small and com- pact; but variability increases in Central America, where some leaves are larger and more elongate, and apices of indusia are deeply dentate. Even greater differences are found in South American plants, especially in Peru, where variability is at its peak. It is likely that part of this may be due to the effects of ecological niches and altitudinal levels, but this kind of analysis must wait for a much- needed revision of this part of the genus. For purposes of this treatment //. fucoides has been segregated into four varieties, a key to which is provided here. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 55 Key to Varieties a. Rachis marginate to narrow-alate, or if broadly alate then alae essentially plane, ultimate segments plane b b. Apex of indusium entire, erose or denticulate c c. Rachis narrow-alate, or sometimes nonalate at base; petiole nonalate, or occasionally marginate near lamina base; basal pinnae often short-stalked; sori rarely more than 3 to a pinna ...... la. var. fucoides c. Rachis broadly alate throughout, each wing broader than the rachis; petiole usually alate distally, at least near lamina base; basal and other pinnae adnate; sori often 5-8 to a pinna Ib. var. calodictyon b. Apex of indusium spinulose-dentate to laciniate Ic. var. pedicellatum a. Rachis broadly alate its entire length, alae and/or ultimate segments often undulate and their margins discontinuously conduplicate Id. var. chachapoyense la. Hymenophyllum fucoides var. fucoides. Fig- ure lla-b. Trichomanes fucoides Sw., Prodr. 136. 1788. TYPE: Jamaica, Swartz (holotype, s; isotypes, B, BM; phOtOS, F & GH Of B, US of BM). Leptocionium fucoides (Sw.) Presl, Hymenophyllaceae 27. 1843. Meringium fucoides (Sw.) Copel., Philipp. J. Sci. 67: 45. 1938. Petiole (0.3-)0.4— 0.8 mm in diameter, nonalate, occasionally marginate near apex. Rachis narrow- alate or marginate, at least distally, alae plane. Pinnae adnate, or 1-2 proximal pairs short-stalked, ultimate segments plane. Sori l-3(-4) pairs to a pinna, apex of indusia entire to denticulate, rarely dentate. On trunks and branches of trees in wet forest, rarely on wet rock walls, 1 700-3300 m, Amazonas, San Martin, Huanuco to Puno. West Indies; southern Mexico to Panama; Ven- ezuela and Colombia to Brazil and Bolivia. The most diminutive of these ferns scarcely dif- fer from H. tunbridgense (L.) J. Sm., the type of the genus, which is said to differ in its smaller leaves and in its strongly arched sori. The latter species supposedly occurs in the West Indies and in Europe and Africa, but it may be only another component in this complex. Some authors place it, with H. peltatum and a few others, in sect. Hymenophyllum, distinguished principally by the strongly arched sori, which are borne nearly per- pendicular to the plane of the lamina. In Old World specimens especially, this character is inconstant and is but one of the problems which needs closer scrutiny when the group is revised. Amazonas: Trail east of La Peca in Serrania de Bagua, Gentry et al. 23034 (F, MO, us). San Martin: Venceremos, near Amazonas border, Gentry et al. 45402 (MO). Huan- uco: Cani, NE of Mito, Bryan 206, 384 (F); Macbride 3397 (B, F, us). Cuzco: Cerro Chuyapi, Bues ASS (GH, us), A56 (us). Puno: Sandia, Weberbauer 713 (B). Ib. Hymenophyllum fucoides var. calodictyon (Bosch) Stolze, comb. & stat. nov. Hymenophyllum calodictyon Bosch, Ned. Kruidk. Arch. 5(3): 172. 1863. TYPE: Peru, Ruiz 84 (ho- lotype, B!). Petiole 0.5-0.8 mm in diameter, conspicuously alate to marginate distally. Rachis broadly alate throughout, each of the alae plane and much wider than the rachis. Pinnae all adnate, with ultimate segments plane. Sori ( l-)2-8 pairs to a pinna, apex of indusia entire. On trees in wet forests, Cajamarca and Ama- zonas south to Cuzco, 2100-3300 m. Ecuador; Peru. In addition to the characters mentioned in the key, var. calodictyon commonly may be distin- guished from the other varieties by the broader segments, membranaceous tissue, and the lighter, yellow-green color. Cajamarca: Prov. Cutervo, Dist. San Andres, Agua Fria, A. Diaz et al. 5340 (USM). Amazonas: Prov. Bagua, Cordillera Colan SE of La Peca, Barbour 3586 (F, MO, USM). Prov. Bongara, near Pomacocha, Wurdack 865 (F, GH, uc, us). Huanuco: Prov. Huanuco, Carpish, Plow- man 6074 (F, GH, us). Pasco: Prov. Oxapampa, E of Abra Cantarizu, Skog et al. 5108 (us). Cuzco: Prov. Paucar- tambo, Dist. Marcachea, near Achirani, Vargas 11142 (F, uc, us). 56 FIELDIANA: BOTANY Ic. Hymenophyllum fucoides var. pedicellatum (Klotzsch) Hieron., Bot. Jahrb. Syst. 34: 435. 1904. Hymenophyllum pedicellatum Klotzsch, Linnaea 20: 439. 1847. TYPE: Venezuela, Merida, Moritz 346 (holotype, B; isotypes, BR, us in part!). Leptocionium pedicellatum (Klotzsch) Fourn., Bull. Soc. Bot. France 19: 249. 1872. Petiole 0.3-0.5 mm in diameter, nonalate. Rachis broadly to narrowly alate, alae plane. Pin- nae adnate, or basal pair short-stalked, ultimate segments plane. Sori 1-3 pairs to a pinna, apex of indusia laciniate to spinulose-dentate. On trees in wet forests, 500-2400 m, Amazonas. Venezuela; Colombia; Ecuador; Peru. This may be synonymous with H. peruvianum, which see under Comments. It is likely that //. ectocarpon Fee, of Central America and the Lesser Antilles, should be included here also. Amazonas: Prov. Bagua, Cordillera Colan, SE of La Peca, Barbour 3598 (F, MO, USM), 3976 (F, MO). Prov. Bagua, Montenegro-Chiriaco, Sagdstegui 5931 (GH). Id. Hymenophyllum fucoides var. chachapoyense Stolze, var. nov. Differt a var. fucoide characteribus sequentibus: alae rhachidis latae et saepe undulatae; segmenta ultima saepe undulata. Petiole 0.5-0.9 mm in diameter, commonly alate distally, at least at the apex. Rachis broadly alate throughout, the alae often undulate. Pinnae ad- nate, the ultimate segments often undulate and their margins discontinuously conduplicate. Sori commonly 3-10 to a pinna, apex of indusia entire. TYPE— Peru, Dept. Amazonas, Prov. Chacha- poyas, Cerros de Calla Calla, 18 km above Lei- mebamba on road to Balsas, 3100 m, Hutchison & Wright 5659 (holotype, GH!; isotype, uc!). On trunks and branches of trees, in wet forests and moist, wooded ravines, 2900-3700 m, Ama- zonas, Huanuco. Colombia (Dept. del Valle); Ecuador (Prov. Car- chi & Napo); Peru. This is a robust variety of higher elevations, similar to var. calodictyon in its stout petioles and larger leaves (to 25 cm long, occasionally longer). In its undulate to crispate rachis wings and ulti- mate segments, var. chachapoyense resembles Hy- menophyllum tortuosum Hooker & Grev. of Chile; however, alae and segments of the latter are much more strongly crispate and are setose-serrate as well. Amazonas: Prov. Chachapoyas, Cerros de Calla Calla, 26 km above Leimebamba on road to Balsas, Hutchison & Wright 6987 in part (F, GH, uc). Huanuco: Tambo de Vaca, Macbride 4460 (F, us). 2. Hymenophyllum peltatum (Poir.) Desv., Mem. Soc. Linn. Paris 6: 333. 1827. Trichomanes peltatum Poir. in Lam., Encycl. 8: 76. 1808. TYPE: Mauritius, "Ile-de-France, Bory de Saint-Vincent (V.s. in Herb, du Petit-Thouars)" (not located). Leaves 2.5-8 cm long, 1-2 cm broad. Petiole 0.15-0.3 mm in diameter, nonalate, glabrous. Lamina narrowly oblong to lanceolate, slightly re- duced at base, 2-3-pinnatisect. Rachis nonalate, occasionally marginate distally, essentially gla- brous. Pinnae 4-10 pairs, adnate, or 1-2 proximal pairs short-stalked, dimidiate (i.e., the 1-4 sec- ondary segments all bome on acroscopic side). Sori 1 or 2 per pinna, not immersed in the segment tissue, most of them strongly arcuate, thus ar- ranged nearly perpendicular to the plane of the lamina. Indusia subspheroid to ovoid, the apex obtuse, entire, receptacle narrow-cylindrical, not or rarely exserted. On rocky slopes and cliffs, 2800-4700 m, Junin, Cuzco, Madre de Dios, Puno. Peru; Bolivia; Argentina; Chile; Europe; Africa. Diem and Lichtenstein (1959) reported that H. peltatum is highly variable in southern South America. They recognized six varieties in Argen- tina and/or Chile and also consider that it is also present in one form or another in areas of the Old World. Apparently only var. peltatum occurs in Peru. It is a much more delicate fern than H. fu- coides and can be easily distinguished from the latter by the strongly dimidiate pinnae. Most sori are rather conspicuously arched out of the plane of the lamina, a character which Morton (1968) used to separate sect. Hymenophyllum from others in subg. Hymenophyllum. This feature is not con- stant in some other species, notably H. tunbridg- TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 57 ense (L.) Sm., and its value is questionable as a diagnostic character above the level of species. Junin: Prov. Huancayo, Dist. Huancayo, Huaytapal- lana. Sounders 1 167 (GH). Cuzco: "Ccarcco" (Ccorca?), Biies 1395 (us). ?Madre de Dios: Pinasniocj, "Panticalla Pass" (Pantiacolla?), Cook & Gilbert 1868, 1870 (us). Puno: Prov. Carabaya, Ayapata, Vargas 10705 (GH). 3. Hymenophyllum mirificum Morton, Bot. Gaz. (Crawfordsville) 93: 338. 1932. TYPE: Peru, Cuzco, Prov. La Convencion, Vilcabamba, Biies 1600 (holotype, us!; photo, F). Buesia mirifica (Morton) Copel., Philipp. J. Sci. 67: 48. 1938. Leaves 16-38 cm long, (5-) 7-1 4 cm broad. Pet- iole 4.5-7 cm long, (0.4— )0. 5-0. 8 mm in diameter, nonalate or, rarely, very narrowly so or marginate at apex, sparsely provided with brownish, flex- uous, catenate trichomes. Lamina broadly or nar- rowly oblong, not or abruptly reduced at base, deeply 3-4-pinnatisect, sparsely and inconspic- uously lamellate adaxially. Rachis strongly flex- uous, nonalate, or weakly alate toward the apex, sparsely provided with flexuous, catenate tri- chomes and (adaxially) also with widely scattered lamellae, these usually 4—6 cells wide at base, abruptly becoming uniseriate and often grading to a sharp, filiform tip. Pinnae 1 0-20 pairs, proximal ones subdistant and short-stalked, distal ones ap- proximate and adnate, costae flexuous and nar- rowly alate, with 6-10 pairs of 2-pinnatisect sec- ondary segments, ultimate segments long and narrow, glabrous, plane, the margins serrate, la- mellae inconspicuous and often widely scattered along veins adaxially. Sori several to many, es- pecially on distal pinnae, not or scarcely immersed in the segment tissue, lying essentially in the plane of the lamina. Indusia subspherical to ovoid, the margins subentire, receptacle subglobose, not exserted. Endemic. In damp ravines and on trees, in wet forests, 2800-3650 m, Cuzco. This is closely related to Hymenophyllum cris- tatum Hooker & Grev. of Ecuador, which has the same general aspect and habit. Both species have elongate, pendent leaves with flexuous rachises, subspherical to ovoid indusia, subglobose recep- tacles, and lamellae on axes and veins. But H. mirificum has larger pinnae, rachis wings essen- tially lacking, lamellae sparse and inconspicuous, and apices of indusia entire to denticulate. Rach- ises in H. cristatum are obviously alate through- out, lamellae are numerous and conspicuous, and indusia are deeply dentate to laciniate at apex. For further discussion about the confused identity of H. cristatum, see H. fucoides. Cuzco: Prov. La Convencion, Vilcabamba, Biies 1592, 1593, 2104 (us). Prov. Paucartambo, Acanacu, Vargas 29 (GH); West 7144 (uc, us). Prov. Urubamba, Puyu- pata-Yuncapata, Vargas 2922 (MO, us). 4. Hymenophyllum lamella turn Stolze, sp. nov. Folia 7-1 5 cm longa, 2.5-4.5(-6) cm lata. Petiolus 1 .5- 5 cm longus, 0.35-0.7 mm diametro, conspicue alatus. Lamina 2-pinnatisecta vel 3-pinnatisecta, conspicue la- mellata. Rhachis recta, late alata, lamellis conspicuis in- structa. Pinnae 9-13-jugatae, confertae vel imbricatae, adnatae, l-2.5(-3) cm longae, costis et venis lamellatis praeditae. Segmenta saepe ultima undulata. marginibus serratis. Indusia sphaeroidea vel ovoidea, brevipedicel- lata, marginibus integris, receptaculis subglobosis vel ovoideis. Leaves 7-15 cm long, 2.5-4.5(-6) cm broad. Petiole 1.5-5 cm long, 0.37-0.7 mm in diameter, conspicuously alate well toward, or quite to, base, sparsely provided with brownish, flexuous, caten- ate trichomes. Lamina oblong-lanceolate, not or scarcely reduced at base, 2-3-pinnatisect, con- spicuously lamellate adaxially. Rachis straight, broadly alate throughout, sparsely to amply pu- bescent with flexuous, catenate trichomes and also provided with conspicuous scalelike lamellae, these several to many cells wide at base, long-acuminate and often grading to a sharp, filiform tip. Pinnae 9-13 pairs, crowded to imbricate, adnate at base, l-2.5(-3) cm long, adaxially lamellate on costae and veins as on the rachis, with 3-6 pairs of 2- pinnatisect or (distally) pinnatisect segments, ul- timate segments glabrous, usually slightly to strongly undulate, the margins serrate, often con- duplicate. Sori few, commonly 1 per pinna on the basal acroscopic segment, not immersed in the segment tissue, lying essentially in the plane of the lamina. Indusia subspherical to ovoid, the margins entire, receptacle subglobose or ovoid, not or slightly exserted. TYPE— Peru, Huanuco, Prov. Huanuco, trail from S entrance of Carpish tunnel to crest of ridge, Luteyn & Luteyn 5474 (holotype, us!; isotype, NY!). Endemic. In wet forests, on tree trunks and branches, 2750-3000 m, Huanuco, Cuzco, Puno. 58 FIELDIANA: BOTANY Some of the specimens cited below were earlier suspected of being new species. On the sheets of Bites A 17 & A20 (us) are Morton's penciled re- marks, "sect. Buesia, n. sp.?"; and Lechler 2568 (B) is annotated by Mettenius, "//. latipes Mett." Apparently Mettenius intended to publish, but a search of the literature has revealed nothing under this name. In its small size, compact lamina, and conspicuously alate axes, this is the most distinc- tive species in sect. Buesia. Cuzco: Alturas de Sicre, Biies 1562 (us). Valley of Rio Urubamba, Biies A- 17, A-20 (us). Prov. La Convention, at Klaus' Folly, above Camp 5, Dudley 1 WOO (GH). Puno: Tatanara, Lechler 2568 (B). fewer pinnae, but the character of its sori is dif- ferent. The narrowly cuneate base of the indusium is deeply immersed in the segment tip, whereas in H. polyanthos the indusium base is broadly cu- neate to obtuse and only slightly immersed in the tissue. San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Cerro Sinsin, Plowman & Schunke 11478 (F). Loreto: Mishuyacu, near Iquitos, Klug 1518 (F, NY, us). Huanuco: Environs of Jingo Maria, Aguilar 304 (uc). Tingo Maria (as San Martin), Allard 20741, 21485 (GH, us). Pasco: Puerto Bermudez (as Junin), Killip & Smith 26415 (F, us). Cuzco: Valle de Lares, alturas del Rio Lachac, Biies 1813a (us). 5. Hymenophyllum apiculatum Kuhn, Linnaea 35: 391. 1868. TYPE: Venezuela, Edo. Aragua, Colonia Tovar, Fendler 32 (holotype, B; iso- type, us). Hymenophyllum dendritis Rosenst., Repert. Spec. Nov. Regni Veg. 6: 308. 1909. TYPE: Bolivia, San Car- los near Mapiri, Buchtien 1093 (holotype, B?; is- otype, us!). Mecodium dendritis (Rosenst.) Copel., Philipp. J. Sci. 67: 26. 1938. Mecodium apiculatum (Kuhn) Vareschi, Flora Ven- ezuela, Caracas 1: 198. 1969. Leaves 1.5-4(-5) cm long, 0.8-1.8 cm broad. Petiole 0.3-1 .5 cm long, 0. 1-0.25 mm in diameter, marginate to narrow-alate, glabrous or with a few scattered trichomes, especially at very base. Lam- ina ovate or deltoid-ovate, not or scarcely reduced at base, 2-3-pinnatifid, glabrous. Rachis glabrous, strongly flexuous to fractiflex, alate throughout. Pinnae 3-6 pairs, adnate, larger ones 0.5-1.2 cm long, bearing 1—4 pairs of secondary segments. Ul- timate segments entire, plane, or occasionally the margins involute or conduplicate. Sori 1-4 to a pinna, usually confined to apical portion of lam- ina. Indusia rhomboid or ovoid, apex entire and obtuse to subacute, with broadly cuneate base, about halfway immersed in the segment tissue, receptacle filiform, not exserted, bearing 6-12 spo- rangia. In dense forests on branches and trunks of trees and on fallen logs, 100-1 500(-2000) m, San Mar- tin, Loreto, Huanuco, Pasco, Cuzco. Venezuela; Colombia; Peru; Bolivia. This would appear to be simply a reduced form of Hymenophyllum polyanthos; however, it not only has smaller leaves, more delicate petioles, and 6. Hymenophyllum polyanthos (Sw.) Sw., J. Bot. (Schrader) 1800(2): 102. 1802. Figure lie. Trichomanes polyanthos S\v., Prodr. 137. 1788. TYPE: Jamaica, Swartz (holotype, s; isotypes, B, BM; photOS, US Of S, F & GH Of B). Mecodium polyanthos (Sw.) Copel., Philipp. J. Sci. 67: 19. 1938. Mecodium mexiae Copel., Univ. Calif. Publ. Bot. 19: 294, /. 48. 1941. TYPE: Peru, Huanuco, near confluence of Rio Huallaga, Mexia 8282 (holo- type, uc!; isotypes, B!, F!, MO!). Hymenophyllum mexiae (Copel.) Morton, Contr. U.S. Natl. Herb. 38: 173. 1968. Leaves (6-)8-15 cm long, 2-5(-6) cm broad. Petiole 0.5-5 cm long, 0.3-0.6 mm in diameter, alate near the apex or throughout, glabrous. Lam- ina ovate or elliptic, scarcely reduced at base (or proximal 1-2 pairs of pinnae slightly reduced), 3- 4-pinnatisect, glabrous. Rachis glabrous, straight or slightly flexuous, alate throughout. Pinnae 8- 14 pairs, essentially adnate, but not strongly over- lapping the rachis, larger ones 1 -4 cm long, bearing 2-many secondary segments. Ultimate segments entire, plane or occasionally a few with condupli- cate segment margins, larger ones commonly more than 1 mm broad, the free portion relatively short, usually about twice as long as broad. Sori 2-many per pinna. Indusia relatively small, usually about as broad as the segment apex, ovoid, elliptic, or lanceolate, broadest toward the base, apex obtuse or subacute, the broadly cuneate base slightly im- mersed in the segment tissue, margins entire, re- ceptacle filiform to narrowly cylindrical or fusi- form, not exserted, bearing 6-15 sporangia. In dense forests, on fallen logs, and on trunks of trees and tree ferns, 1 00-2000(-2400) m, Ama- zonas, Loreto, San Martin south to Puno. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 59 Pantropical. This is one of the most widely distributed ferns of tropical regions around the world and with its many close relatives represents one of the more taxonomically difficult species complexes. Com- ponents of the group each have only a few distin- guishing characters and these are often inconstant. Some of the features seem to vary more or less in different geographic areas, so that species concepts in each of these regions vary proportionately. The group has long been misunderstood; consequently, in most herbaria, a number of specimens may be filed incorrectly under Hymenophyllum polyan- thos. For purposes of this treatment, H. polyanthos includes H. mexiae, which differs only in its small- er leaves. Leaves of H. apiculatum are also very small, but this species (q.v). is characterized by other, quantitative, features, and so should be con- sidered distinct. Hymenophyllum mathewsii may be merely a more luxuriant, higher elevation form of H. polyanthos, with larger leaves and sori, and the axes less distinctly alate. Another species of the subgenus occurring in Peru is H. myriocarpum, which differs principally in shape and character of the indusia: in most specimens sori appear to be pedicellate, for the apices of ultimate segments are strongly constricted; furthermore, mature indusia are subglobose, or often noticeably broader than long. This contrasts with the essentially ovoid in- dusia of//, polyanthos et al., commonly somewhat longer than broad and slightly to strongly im- mersed in the segment tips, which are not or scarcely constricted. While this character is prob- ably the most reliable one in subg. Mecodium in Peru, unfortunately even it is not 1 00% constant, for on occasional specimens indusia may vary somewhat from one shape to another. Conse- quently, most species in the subgenus must be sep- arated by suites of characters. Ultimately, taxo- nomic problems will be solved only by thorough study of species groups throughout their entire range. Two other characters are often helpful in sep- arating H. polyanthos (at least in Peru) from H. myriocarpum— particularly useful where sterile specimens are involved. Pinnae of H. myriocar- pum commonly have basal segments strongly overlapping the rachis. In H. polyanthos pinnae are adnate and basal segments often crowd or touch the rachis, but rarely do some of them overlap. Also, H. polyanthos occurs at lower elevations in Peru, 100-2000(-2400) m; whereas H. myriocar- pum is found at ( 1 700-)2000-4200 m. Amazonas: Prov. Bagua, 1 2 mi E of La Peca, Harbour 2466 (uc in part). San Martin: Ravine E of Tingo Maria, Allard 21446 (GH, us). Prov. Mariscal Caceres, Tocache Nuevo, J. Schunke 5693 (F). Loreto: Sierra del Pongo, Mexia 6291a (GH, uc). Huanuco: Prov. Huanuco, Cerro Cucharas, Woytkowski 1152(cH). Prov. Huamalies, Rio Monzon, 20 km W of Tingo Maria, Tryon & Tryon 5299 (GH). Pasco: Prov. Oxapampa, Cordillera San Matias, Leon 317 (F, GH, USM). Pichis Trail, Enenas (as Junin), Killip & Smith 25651 (F, NY, us). Junin: La Merced, E of Quimiri Bridge, Killip & Smith 24022 (F, NY, us). Madre de Dies: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, Foster et al. 10683 (F). Cuzco: Prov. Paucartam- bo, Kosnipata, Vargas 17860 (GH). Puno: San Gaban, Lechler 3327 (B). 7. Hymenophyllum mathewsii Bosch, Ned. Kruidk. Arch. 5: 162. 1863. SYNTYPES: "Peruvia," Mathews (B!; isosyntype, p; prob- able isosyntype, B!; photos, F, GH, & us of p). Ecuador, Quito, Cuming (B). Leaves (6-) 1 4-30 cm long, 1.5-4 cm broad. Pet- iole 2.5-6(-8) cm long, 0.35-0.5 mm in diameter, nonalate, glabrous. Lamina linear-lanceolate to oblanceolate, rather abruptly reduced or strongly and gradually reduced at base, 2-3-pinnatisect, glabrous. Rachis glabrous, alate throughout except usually nonalate at base, or there narrow-alate on only one side. Pinnae 14-18 pairs, adnate and often overlapping the rachis, or basal ones short-stalked, larger ones (1.5-)2-3.5 cm long, bearing 4-8 pairs of secondary segments. Ultimate segments entire, plane or occasionally undulate, or a few with mar- gins conduplicate. Sori 4-many per pinna. Indusia relatively large, usually slightly broader than the segment apex, ovoid, apex obtuse to subacute, the broadly cuneate base slightly immersed in the seg- ment tissue, margins entire, receptacle short and filiform, bearing 6-12 sporangia. Cloud forests, commonly on trunks and branch- es of trees, 2 1 00-3 1 50 m, Huanuco, Pasco, Cuzco. Ecuador; Peru. Hutchison 1751 (cited below) has delicate, di- minutive leaves only 6-8 cm long; this depauper- ate condition probably results from the unusual habitat, "in cave." This species is very closely re- lated to Hymenophyllum polyanthos (q.v.) and may not merit distinction at the species level. Normal specimens of//, mathewsii tend to have larger sori, in relation to the segment apex, and longer, pen- dent leaves, with no trace of tissue along the petiole and, commonly, none between the proximal two pinnae. Leaves of H. polyanthos are smaller and 60 FIELDIANA: BOTANY more compact, erect or arching, with rachis dis- tinctly alate. The petiole is marginate or alate throughout, or at least distally. Often the wing is deciduous upon drying, but traces can usually be found by close scrutiny. Huanuco: Prov. Huanuco, Carpish, Plowman 6074A (F). Pasco: Prov. Oxapampa, San Alberto, Yanachaga, van der Werffel al. 8437 (MO). Cuzco: Huadquina, Bues 1346 (us). Prov. La Convention, Cochapata, Valle de San Miguel, Bues 2176 (us). Prov. Urubamba, summit of Huayna Picchu, Hutchison 1751 (atypical) (F, uc). 8. Hymenophyllum trichomanoides Bosch, Ned. Kruidk. Arch. 5(3): 158. 1863. SYNTYPES: Peru, San Martin, Tarapoto, Spruce 4696 (isosyntypes, GH!, NY!, P!, us!). Colombia, Moritz (BM?), Schomburgk(BM?). Ecuador, Pi- chincha, Quito, Cuming (not located). "Brit- ish Guiana," Schomburgk (BM?). Mecodium trichomanoides (Bosch) Pic.-Ser., Webbia 28: 469. 1973. Leaves (9-)l 1-26 cm long, 4-6(-8) cm broad. Petiole 5-12 cm long, 0.4-0.8 cm in diameter, marginate to alate distally (rarely nonalate), with a few scattered trichomes. Lamina broadly or nar- rowly ovate (1 or 2 pairs of proximal pinnae com- monly somewhat reduced), 2-3-pinnatisect, gla- brous. Rachis glabrous, slightly flexuous, narrowly to broadly alate throughout, the wing margin sometimes conduplicate. Pinnae 7-14 pairs, ad- nate and overlapping the rachis, or 1-2 proximal pairs short-stalked, larger ones 2.2-4 cm long, bearing 4-8 pairs of secondary segments. Ultimate segments entire, plane, or some of them with in- volute margins, larger ones commonly less than 1 mm broad, the free portion linear, usually 2.5-4 times as long as broad. Sori 6-12 per pinna. In- dusia relatively small, usually about as broad as the segment apex, rhomboid, elliptical or conical, broadest at or beyond the middle, apex acute to obtuse, the sharply cuneate base deeply (often half- way) immersed in the segment tissue, margins en- tire, receptacle long, filiform, often exserted, bear- ing 6-10 sporangia. In dense, wet forests, on trunks of trees, 300- 3000 m, San Martin to Cuzco. Surinam to Colombia, south to Bolivia. Besides the characters used in the key, Hymen- ophyllum trichomanoides can often be distin- guished by the narrower ultimate segments. Larger ones are commonly 0.6-0.9 mm broad and linear. Other species of subg. Mecodium have relatively broader ultimate segments, most of them over 1 mm broad, or if not, then only about twice as long as broad. This is a rather reliable way to identify sterile specimens. Hymenophyllum trichomanoides differs sharply from all other species of subg. Mecodium in Peru by the narrowly cuneate sori which are deeply im- mersed in the segment tissue. Indusia are broadest at or beyond the middle and, at least in Peru, are long and narrow: either narrowly elliptic with sub- acute apices, or conical, with rounded apices. All specimens seen from Peru clearly match the Spruce syntype. Some specimens examined from Colom- bia and Venezuela have shorter and broader in- dusia and much shorter petioles, and it is possible that these represent a variant. However, this has not been substantiated by comparison with the other syntypes from northern South America, which we have not seen. With H. trichomanoides probably should be included H. trianae Hieron. of Colombia. A syntype of the latter, Lehmann 7410 (us), matches Spruce 4696, except that re- ceptacles are not exserted. Huanuco: Cerros del Sira, SW slope of Rio Llullapichis watershed, Dudley 13375 (GH). Pasco: Prov. Oxapampa, Palcazu Valley, Cabeza de Mono, D. Smith 3756 (F, MO). Junin: Schunke Hacienda, above San Ramon, Killip & Smith 24844 (F, GH, NY, us). Cuzco: Sahuayacu, Bues 804, 817 (us). 9. Hymenophyllum ferax Bosch, Ned. Kruidk. Arch 4: 392. 1 859. TYPE: Venezuela, Merida, Funck & Schlim 1578 (holotype, L?; isotype, p; photo, us of P). Mecodium ferax (Bosch) Copel., Philipp. J. Sci. 67: 25. 1938. Leaves 16-38 cm long, 4-8 cm broad. Petiole 2-9 cm long, 0.4-0.9 cm in diameter, alate toward apex (wings sometimes deciduous), essentially gla- brous. Lamina ovate to narrow-elliptic, scarcely to somewhat reduced at base (1-several proximal pairs of pinnae reduced), 3-pinnatisect, glabrous. Rachis glabrous, straight or slightly flexuous, alate throughout, rachis wings and those of other axes sometimes conduplicate. Pinnae 1 2-24 pairs, ad- nate, and basal segments overlapping the rachis, larger ones 2.5-5 cm long, bearing 7-10 pairs of TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 61 secondary segments, some elongate pinnae inter- mixed with normal ones. Ultimate segments en- tire, plane to slightly undulate. Sori 8-many per pinna. Indusia relatively large, subrotund, circular in outline or broader than long, apex erose, base rounded, not immersed in the greatly constricted segment apex, receptacle short, filiform, bearing 7-12 sporangia. On trees in wet forests, 2400-3300 m, Cuzco. Venezuela; Colombia to Peru. This is most closely related to Hymenophyllum myriocarpum var. endiviifolium, from which it dif- fers in the erose indusia tips and the presence of one to several unusually elongate pinnae which are scattered among the normal ones— the latter a condition similar to that of H. undulatum. A remarkably huge specimen, Lechler 2420 (B; not other 2420 at B, F, K: H. elegans), was collected in Peru at an undesignated site. This is apparently a monstrous form of H. ferax. It matches this species in every way, including the erose indusia, but the leaf is 85 cm long and the petiole nearly 2 mm in diameter. It is by far the stoutest specimen of Hymenophyllaceae we have seen and belies the appellation "filmy fern." Cuzco: Vilcabamba, Bues 1601 (us). Prov. Paucartam- bo, Pilahuata, Vargas 4912 (us). Dept. Unknown: Lech- ler 2420 (B in part). 10. Hymenophyllum myriocarpum Hooker, Sp. fil. 1: 106, /. 37 d. 1844. TYPE: Colombia, Hart- weg 1530 (holotype, K!; isotypes, BM, p; photo, F of P). Leaves 7-28 cm long, 2.5-5(-6) cm broad. Pet- iole 1.5-8(-10) cm long, 0.3-0.7 mm in diameter, marginate to alate throughout or at least distally (wings sometimes deciduous), essentially gla- brous. Lamina lanceolate to ovate, elliptic, or ob- lanceolate, scarcely to strongly reduced at base, 2- 3-pinnatisect, glabrous. Rachis glabrous, straight or slightly flexuous, alate throughout, rachis wings and those of other axes often conduplicate. Pinnae 8-30 pairs, adnate and basal segments strongly overlapping the rachis, larger ones 1-4 cm long, bearing 5-10 pairs of secondary segments. Ulti- mate segments entire, plane to undulate, often with conduplicate margins. Sori 5-many per pinna. In- dusia subglobose or broader than long, much broader than (and not at all immersed in) the strongly constricted segment apex, rounded at base, rounded and entire at apex, receptacle short to nearly obsolete, filiform, bearing 8-20 sporangia. This, like H. polyanthos (q.v.), is a widespread and greatly misunderstood species which is sorely in need of detailed study. It seems to consist of several, not highly distinctive components which are recognized as varieties. With var. myriocar- pum might possibly be included H. axillare Sw. of the West Indies and H. andinum Bosch of Ec- uador. Of all the names, H. axillare has priority. The species occurs in Mexico and Central Amer- ica and in South America from Venezuela and Colombia to Bolivia. Key to Varieties a. Lamina elliptic to oblanceolate, broadest at or beyond the middle; several or many proximal pinnae somewhat to greatly reduced lOa. var. myriocarpum a. Lamina ovate to lanceolate, broadest near the base; 1 or 2 proximal pinnae (if any) slightly reduced b b. Larger leaves 7-1 5 cm long; pinnae commonly patent and strongly imbricate lOb. var. nigrescens b. Larger leaves ( 1 2-) 1 5-28 cm long; pinnae (most of them) ascending and contiguous or only slightly imbricate . lOc. var. endiviifolium 1 Oa. Hymenophyllum myriocarpum Hooker, var. myriocarpum. Mecodium myriocarpum (Hooker) Copel., Philipp. J. Sci. 67: 25. 1938. Leaves (9-) 12-28 cm long. Lamina elliptic, ob- long or oblanceolate, broadest at or beyond the middle, several proximal pinnae somewhat to greatly reduced. Rachis and ultimate segments plane, some of their margins occasional condu- plicate. Sori 10-20 per pinna. 62 FIELDIANA: BOTANY In wet forests, usually pendent from tree trunks and branches, or on wet, mossy rocks, rarely on the forest floor, (1 700-)2000-3200 m, Cajamarca, Amazonas, Huanuco to Cuzco. Mexico to Costa Rica; Venezuela; Colombia to Bolivia. Cajamarca: Prov. Hualgayoc, Hacienda Taulis, Hutchison & Bismarck 6415 (F, GH [as Lambayeque], MO, uc, us). Amazonas: Prov. Bongara, WSW of Po- macocha, Wurdack 866 (GH, us). Huanuco: Mito, Bryan 206a (F). Pasco: Enenas (as Junin), Pichis Trail, Killip & Smith 25700 (F, NY, us). Cuzco: Prov. Urubamba, along Rio Urubamba near town of Machu Picchu, Tryon & Tryon 5410 (F, GH, us). lOb. Hymenophyllum myriocarpum var. nigres- cens (Liebm.) Stolze, stat. & comb. nov. Hymenophyllum nigrescens Liebm., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd. 5,1: 292. 1849. LECTOTYPE (designated by A. R. Smith, Flora of Chiapas 2: 133. 1981): Mexico, Puebla, Chinantla, Liebmann 537 (lectotype, c; isolec- totypes, BM, K, P). Sphaerocionium nigricans KJotzsch, Linnaea 18: 536. 1 844, not Hymenophyllum nigricans Colla, 1836. SYNTYPES: Peru, Dombey 87 (syntype, u!; iso- syntype, B); Venezuela, Aragua, Colonia Tovar, Moritz 268 (syntype, B; isosyntype, in part, K!), 268b (syntype, B; isosyntype, BM). Hymenophyllum nigricans (KJotzsch) Kunze, Bot. Zeit. (Berlin) 244. 1847. Hymenophyllum nigricans (KJotzsch) Copel., Phillip. J. Sci. 67: 25. 1938. Leaves 7-1 5 cm long. Lamina ovate. Rachis and ultimate segments plane, their margins frequently conduplicate. Sori 3-15 per pinna. Erect or arching from tree trunks or branches, rarely from wet rocks, in thickets and wet forests, 1750-3400 m, Amazonas to Cuzco. Southern Mexico to Costa Rica; Venezuela; Co- lombia to Bolivia. Sterile specimens can easily be confused with H. polyanthos, but the latter occurs at lower ele- vations, and laminae are less compact. See dis- cussion of H. polyanthos for further comments. Amazonas: Prov. Chachapoyas, Cerros de Calla Calla above Leimebamba, Hutchison & Wright 6987 (in part: F, GH, uc). San Martin: Prov. Rioja, Pedro Ruiz-Moy- obamba Road, D. Smith 4376 (F, MO). Pasco: Prov. Ox- apampa, Abra los Mellizos, 4-8 km from Enenas, Skog et al. 5033 (vs). Junin: Huacapistana, Killip & Smith 24164 (us). Ucayali: Plantation Azul, Ridoutt (USM). Prov. Coronel Portillo (as Loreto), NE of pass at La Divisoria, Skoget al. 5157 (us). Cuzco: Valley of Urubamba, Sues A18,A19(us). lOc. Hymenophyllum myriocarpum var. endivi- ifolium (Desv.) Stolze, stat. & comb. nov. Hymenophyllum endiviifolium Desv., Mem. Soc. Linn. Paris 6: 334. 1827. TYPE: "Peruvia" (holotype, p; photos, GH, uc, us). Hymenophyllum multiflorum Rosenst., Meded. Rijks- Herb. 19: 5. 1913. TYPE: Bolivia, Comarapa, Herzog 1951 (holotype, B; isotype, us!). Mecodium multiflorum (Rosenst.) Copel., Philipp. J. Sci. 67: 25. 1938. Mecodium endiviifolium (Desv.) Pic. -Sen, Webbia 28: 469. 1973. Leaves (12-) 15-28 cm long. Lamina ovate or ovate-lanceolate. Rachis and ultimate segments slightly to strongly undulate, their margins often conduplicate. Sori 1 6-many per pinna. On trees and wet banks in deep forest, 2400- 4200 m, Amazonas, Huanuco, Huancavelica, Cuzco. Colombia; Ecuador; Peru; Bolivia. Amazonas: Prov. Chachapoyas, Calla Calla slopes near km 4 1 5-4 1 8 of Leimebamba-Balsas road, Wurdack 1750 (us, USM). Huanuco: Prov. Huanuco, Mirador, road Aco- mayo to Chanchao, Mexia 7761 (F, GH, K, MO, uc, us). Huancavelica: Prov. Tayacaja, Chuspi-Tocas, between Colcabamba and Paucarbamba, Tovar 2050 (GH, USM). Cuzco: Prov. Paucartambo, Dist. Marcachea, near Achirani, Vargas 11143 (F, K. in part, uc). 11. Hymenophyllum undulatum (Sw.) Sw., J. Bot. (Schrader) 1800(2): 101. 1802. Leaves 6-35(-40) cm long, 0.7-2.5 cm broad. Petiole 0.5-3(-5) cm long, 0.15-0.3 mm in di- ameter, nonalate, essentially glabrous. Lamina lin- ear to narrow-elliptic, often strongly and gradually reduced at base, 2-3-pinnatisect, glabrous. Rachis glabrous, marginate to crispate-alate, commonly nonalate toward base, or discontinuously alate on alternate sides due to the long-decurrent pinna bases. Pinnae 1 6-many pairs, adnate, commonly less than 2 cm long, but often a few of them greatly elongate and lamina-like, normal ones bearing 3- 8(-10) pairs of secondary segments. Ultimate seg- ments entire, plane, or the margins conduplicate in drying, to undulate or markedly crispate. Sori 3-many per pinna. Indusia nearly circular, but on mature son most of them broader than long, not TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 63 or scarcely immersed in the segment tissue, com- monly truncate at base, the margins entire and plane to crispate, receptacle very short, filiform, bearing 1^4(-5) sporangia. This species varies greatly as to length and shape of lamina and margins of rachis wings and ulti- mate segments. In rocky habitats leaves usually tend to be less than 10 cm long and narrowly el- liptic, whereas the flaccid, linear leaves pendent from tree branches sometimes reach 40 cm in length. Several species have been described based on whether the tissue is plane or undulate or strongly crispate. However, comparison of indi- viduals throughout the entire range seems to in- dicate that few of these characters are very con- sistent. Only the conspicuously crispate variants seem to merit recognition, as indicated in the fol- lowing key. Key to Varieties a. Ultimate segments and rachis wings plane to undulate, margins often discontinuously conduplicate in drying; rachis nonalate in the proximal '/3 to l/2 or nearly throughout 1 la. var. undulatum a. Ultimate segments and rachis wings markedly crispate, often so strongly as to appear dentate; rachis commonly alate except near lamina base, or wings sometimes lacking in the proximal 'A or lk of the rachis . ..lib. var. fendlerianum 1 1 a. H ymenoph y Hum undulatum var. undulatum. TrichomanesundulatumSw.,Prodr. 137. 1788. TYPE: Jamaica, Swartz (holotype, s; isotypes, B, Herb. Willd. 20238, BM!, P; photos, F & us of s). Hymenophyllum reniforme Hooker, Sp. fil. 1: 110, /. 38c. 1844. TYPE: Peru, Mathews 1783 (holo- type, K.!; isotypes, B!, F!, K!, P!; photos, F & us of P). Hymenophyllum rimbachii Sodiro, Anal. Univ. Quito 13: 47 (Crypt, vase. quit. 33). 1893. TYPE: Ec- uador, Prov. Azuay, Rimbach (holotype, not lo- cated; isotype, us!). Mecodium undulatum (Sw.) Copel., Philipp. J. Sci. 67: 26. 1938. In wet forests on rock cliffs or pendent from tree trunks and branches, 1500-4100 m, Huanuco to Puno. Greater Antilles; Guadeloupe; southern Mexico to Panama; Surinam to Colombia, south to Bo- livia; southeast Brazil. Hymenophyllum reniforme and H. rimbachii were based on depauperate specimens in which ultimate segments are mostly plane but with their margins conduplicate. Thus the segments appear to be much narrower than typical H. undulatum. Some of this folding of the margins appears to be a product of drying and some of it merely part of the natural variation of the species. Huanuco: Tambo de Vaca, Bryan 624 (F, us). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8482 (MO, uc). Junin: Prov. Tarma, Yanango Mountains, Weberbauer 2135 (B). Cuzco: Pau- cartambo, slopes of Pilahuata, Vargas 4909 (MO, us). Puno: Sandia, Weberbauer 1293 (B). 1 1 b. Hymenophyllum undulatum var. fendlerian- um (J. W. Sturm) Stolze, comb. & stat. nov. Hymenophyllum fendlerianum J. W. Sturm in Mart., Fl. bras. 1(2): 291. 1859. LECTOTYPE (desig- nated by Lellinger, Mem. New York Dot. Gard. 38: 12. 1984): Venezuela, Edo. Aragua, Colonia Tovar, Fendler 35 (us; isolectotype, BR; photo, us of BR). Hymenophyllum contortum Bosch, Ned. Kruidk. Arch. 5(3): 170. 1863. TYPE: Costa Rica, Aguacate, Hoffman (B). Hymenophyllum polycarpum Kuhn, Linnaea 35: 391. 1868. TYPE: "Peruvia" (holotype, B!; photo, F). Mecodium fendlerianum (J. W. Sturm) Copel., Phil- ipp. J. Sci. 67: 26. 1938. Mecodium contortum (Bosch) Copel., Philipp. J. Sci. 67: 26. 1938. In forests on wet, rocky cliffs, or pendent from tree trunks and branches, 1250-2900 m, Ama- zonas, Huanuco, Junin, Puno. Hispaniola; southern Mexico; Guatemala; Cos- ta Rica; Surinam to Colombia, south to Bolivia; southeastern Brazil. Amazonas: Prov. Bagua, Cordillera Colan SE of La Peca, Barbour 3539, 3738 p.p. (MO). Huanuco: Cushi, trail to Tambo de Vaca, Bryan 684 (F, us). Playapampa, Macbride 4498 (F, us). Junin: Schunke Hacienda, above San Ramon, Killip & Smith 24843 (NY, us). Puno: Prov. 64 FIELDIANA: BOTANY Carabaya, road from San Gaban to Macusani, Maas et al. 6109 (MO). Dept. Unknown: Without location, Dom- bey (P). 12. Hymenophyllum molle Morton, Contr. U.S. Natl. Herb. 29: 149. 1947. TYPE: Peru, Cuz- co, Vilcabamba, Achiyayoc Inca ruins, Biies 2102 (holotype, us!; isotype, GH!). Sphaerocionium molle (Morton) Pic.-Ser., Webbia 28: 471. 1973. Leaves indeterminate, to 35 cm long and 3.5 cm broad. Petiole 3-6 cm long, 0.2-0.3 mm in di- ameter, nonalate, sparsely pubescent with simple to (rarely) forked trichomes, or glabrate. Lamina 2-pinnatisect or nearly 2-pinnate, abundantly pu- bescent, the rachis commonly nonalate through- out. Pinnae short-stalked (at least proximal ones), cut deeply or nearly to costa, costae regularly and conspicuously alate on both sides, the 4-9 pairs of segments simple or bifid, entire, plane, veins and margins abundantly or densely pubescent with stellate trichomes, but trichomes lacking between veins and margins. Indusia slightly broader than long, often broader than the segment tips. Endemic. In forests, 2200-3600 m, Cuzco. Cuzco: Huadquina, Biies 708 (us). Prov. La Conven- cion, Biies 2072, 2115 (us). Prov. Paucartambo, Vargas 3636 (GH). 13. Hymenophyilum trichophyllum HBK., Nov. gen. sp. 1: 27. 1815. Leaves indeterminate, to 25 cm long and 3 cm broad. Petiole 0.5-4 cm long, 0.2-0.25 mm in di- ameter, nonalate, sparsely pubescent with simple to stellate trichomes, or glabrate. Lamina 2-pin- nate-pinnatisect, abundantly pubescent, the rachis commonly nonalate throughout. Pinnae short- stalked (at least proximal ones), cut deeply or quite to costa, costae partially nonalate or discontin- uously alate by the decurrent bases of segments, normal pinnae with 2-3(-4) pairs of segments (but often a few of them greatly elongate and equaling the lamina in size and shape), the segments (or pinnules) simple or bifid, plane to undulate or cris- pate, veins and margins abundantly to densely pu- bescent with stellate trichomes, but trichomes lacking between veins and margins. Indusia about as broad as long and nearly as broad as the segment tips. The species occurs in Guatemala; Costa Rica; Panama; Guyana to Venezuela and south to Bo- livia. The leaves of this delicate fern sometimes in- tertwine to form mats on the trunks of trees. A singular feature is the irregular character of the pinnae: often, but not always, one to several pin- nae become greatly elongate and once again pin- nate-pinnatisect, each of these sometimes equaling the length of the leaf on which it is borne. Of the several varieties of Hymenophyllum trichophyl- lum, two are recognized in Peru. Key to Varieties a. Ultimate segments plane to slightly undulate 13a. var. trichophyllum a. Ultimate segments conspicuously undulate to crispate 13b. var. buesii 13a. Hymenophyllum trichophyllum var. tricho- phyllum. TYPE: Venezuela, Cumana, be- tween Cocollar and Guardia de San Augustin, Humboldt (holotype, P). Hymenophyllum procerum Bosch, Ned. Kruidk. Arch. 4: 409. 1859, nom. nud. Sphaerocionium trichophyllum (HBK.) Copel., Phil- ipp. J. Sci. 67: 32. 1938. In Peru, thus far known only from one collec- tion: on shaded rocks, Puno. Elsewhere, in wet forests, on rocks, clay banks or trees, 2000-3800 m; range the same as the species. The name H. procerum was based on H. pul- chellum sensu Mett., for which there was no de- scription. The specimen probably intended as the type is the only collection of var. trichophyllum thus far known from Peru. It closely matches other specimens of var. trichophyllum and is cited here. Puno: San Gaban, Lechler 2250 (B, BR, F, P). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 65 13b. Hymenophyllum trichophyllum var. buesii Morton, Contr. U.S. Natl. Herb. 29: 1 52. 1 947. TYPE: Peru, Cuzco, Prov. La Convention, Abra Mirador, Bues 2076 (holotype, us!; is- otype, GH!). Thus far known only from the type collection, 3150m. Although Morton distinguished his var. buesii from var. trichophyllum solely on the obviously undulate segments, it also appears that the former has less densely pubescent laminae. However, un- til more specimens are collected it remains to be seen if either of the differences are sufficiently sig- nificant to merit even varietal separation. 14. Hymenophyllum elegans Sprengel, Syst. veg. 4: 133. 1827. TYPE: Brazil, Sellow (holotype, LZ destroyed; isotype, B!). Hymenophyllum tenerrimum Bosch, Ned. Kruidk. Arch. 5: 185. 1863. TYPE: Peru, San Martin, Tarapoto, "in monte Campana," Spruce 4702 (holotype, K!; isotype, BM!). Sphaerocionium tenerrimum (Bosch) Pic.-Ser., Web- bia28:472. 1973. Sphaerocionium elegans (Sprengel) Copel., Philipp. J. Sci. 67: 32. 1938. Leaves determinate, to 20 cm long and 2 cm broad. Petiole 0.5-2 cm long, 0.15-0.2 mm in di- ameter, nonalate, sparsely pubescent with simple or forked trichomes. Lamina pinnate-pinnatisect, sparsely to moderately pubescent, the rachis non- alate, or narrowly and irregularly alate distally or throughout due to the decurrent bases of pinnae. Pinnae sessile or adnate, pinnatifid or pinnatisect, the l-2(-3) pairs of segments, simple or (basal ones) bifid, entire, plane to slightly undulate, mar- gins sparsely to moderately pubescent, with tri- chomes highly variable: simple to 2-3-forked at base, or the branches again forked; trichomes rare on veins and lacking between veins and margins. Indusia slightly broader than long, about as broad as the segment tips. On shaded rocks, San Martin, Puno, and from two other collections, localities unspecified, 3000 m. Elsewhere, in wet forests, on rocks, clay banks or tree trunks, 800-2400 m; Guatemala; Costa Rica; Panama; West Indies; Venezuela; Colombia; Peru; Bolivia; Brazil. Hymenophyllum tenerrimum was said to differ only by lack of trichomes on the veins, and by having marginal trichomes mostly three-forked at the base. However, in H. elegans, trichomes are only occasional to rare or lacking on the veins, and marginal trichomes are highly variable, often in- cluding many which are three-forked at base. The isotype of H. tenerrimum has occasional tri- chomes on the veins, and a number of marginal ones that are simple or one-forked. In fact, both taxa may be only a variant of//, lineare (Sw.) Sw., of Central America and the West Indies, which also lacks trichomes on veins, but which appears to have more pinna segments and simpler and less variable marginal trichomes. Further study is needed. San Martin: Tarapoto, "in monte Campana," Spruce 4700, 4701 (K). Puno: San Gaban, Lechler (B). Dept. Unknown: Lechler (NY, us), Lechler 2420 (B, F, K; another 2420 at B is H. ferax). 15. Hymenophyllum adiantoides Bosch, Ned. Kruidk. Arch. 5: 184. 1863. TYPE: Peru, San Martin, Tarapoto, Spruce (holotype, L; prob- able isotypes, B!, GH!, K!, L!, NY, us; photos, GH & us of L). Sphaerocionium adiantoides (Bosch) Pic.-Ser., Web- bia28:470. 1973. Leaves indeterminate, to 1 2 cm long and 2 cm broad. Petiole 1-2 cm long, 0.2-0.25 mm in di- ameter, nonalate, sparsely pubescent with (mostly) stellate trichomes. Lamina pinnate-pinnatisect, sparsely to moderately pubescent, the rachis non- alate near the base, narrowly alate distally or ir- regularly alate due to the decurrent bases of pin- nae. Pinnae adnate, pinnatifid, or pinnatisect, the 3-4 pairs of segments simple or bifid, entire, slight- ly undulate, veins and margins sparsely to mod- erately pubescent with (mostly) stellate trichomes, but trichomes lacking between veins and margins. Indusia as long as or slightly longer than broad, about as broad as the segment tips. Apparently known only from the type collec- tion. With this probably should be included Hy- menophyllum sprucei Baker, under which see Comments. 16. Hymenophyllum hirsutum (L.) Sw., J. Bot. (Schrader) 1800(2): 99. 1802. 66 FIELDIANA: BOTANY Trichomanes hirsutum L., Sp. pi. 2: 1098. 1753. TYPE: based on t. 50b of Plumier, Traite foug. Amer., apparently representing a plant from the West Indies. Trichomanes ciliatum Sw., Prodr. 136. 1788. TYPE: Jamaica, Swartz (holotype, s; isotype, B; photo, us of s). Hymenophyllum ciliatum (Sw.) Sw., J. Bot. (Schrader) 1800(2): 100. 1802. Sphaerocionium ciliatum (Sw.) Presl, Hymenophyl- laceae 34. 1843. Sphaerocionium hirsutum (Sw.) Presl, Hymenophyl- Iaceae34. 1843. Leaves determinate, to 12 cm long and 4 cm broad. Petiole 0.3-2.3 cm long, 0.2-0.3 mm in diameter, strongly alate (often nearly to base), sparsely provided with stellate trichomes. Lamina 2-pinnatisect, sparsely to moderately pubescent, the rachis regularly and conspicuously alate throughout. Pinnae 6-15 pairs, ovate or oblong- ovate, deeply incised, the 2-5 pairs of segments plane, simple to bifid, the veins and margins pu- bescent with stellate or (on the margins) 2-forked trichomes, but trichomes lacking between veins and margins. Indusia as long as or (usually) longer than broad, somewhat broader than the segment tips. In dense, wet forests, on branches and trunks of trees, 500-2000 m, Loreto, Amazonas to Puno. Mexico to Panama; West Indies; Guianas to Venezuela, south to Bolivia and Brazil. Amazonas: Prov. Bagua, E of La Peca, Barbour 2753 (MO). San Martin: Tarapoto, Monte Campana, Spruce 4698 (K). Loreto: Balsapuerto, lower Rio Huallaga basin, Killip & Smith 28539 (F, GH, NY, us). Huanuco: Prov. Huanuco, Dist. Churubamba, crest of Santo Toribio, Mexia 8145 (B, F p.p., GH, us). Pasco: Prov. Oxapampa, W side of Cordillera de San Matias, D. Smith 2054 (MO). Cuzco: Prov. La Convencion, Valle San Miguel, San Ja- cinto, Bues 2139 (us). Puno: San Gaban, Lechler 2296 (B). 1 7. Hymenophyllum crispum HBK., Nov. gen. sp. 1: 26. 1815. TYPE: Venezuela, Silla de Ca- racas, Humboldt & Bonpland (holotype, p; photos, GH & us of P). Figure lid. Sphaerocionium crispum (HBK.) Klotzsch, Linnaea 1: 537. 1844. Leaves determinate, to 20 cm long and 4 cm broad. Petiole 0.5-5 cm long, 0.1-0.3 mm in di- ameter, alate near apex, sparsely provided with delicate, simple or 1 -forked trichomes. Lamina pinnate to nearly 3-pinnate, sparsely pubescent, the rachis regularly and conspicuously alate throughout, the wing strongly undulate or undu- late-crispate. Pinnae 8-20 pairs, simple (distally) or bifid to nearly 2-pinnate, the 1-8 pairs of seg- ments undulate or crispate, the veins and margins sparsely pubescent, marginal trichomes simple or 1 -forked, but trichomes lacking between veins and margins. Indusia as long as or (usually) longer than broad, Oiten broader than the segment tips. The species occurs in Jamaica, Mexico, Gua- temala, Costa Rica, Venezuela, Colombia, Peru, Bolivia, and Brazil. Hymenophyllum crispum might be confused with H. valvatum, another species in the subgenus with undulate segments. However, the latter is larger and coarser with a much thicker petiole, and the rachis wing and segments are not as strongly un- dulate. Two varieties are recognized here. Key to Varieties a. Pinnae simple (distally) to trifid or, if pinnatisect, the secondary segments 1-2 pairs, simple to bifid 1 7a. var. crispum a. Pinnae (larger ones) 2-pinnatisect, the secondary segments 4-8 pairs, pinnatisect 1 7b. var. bipinnatisectum 1 7a. Hymenophyllum crispum var. crispum. Fig- ure lid. In forests, on trees or wet banks or cliffs, 2 1 50- 3000 m, Amazonas, Cuzco, Puno. Range the same as the species. Amazonas: Prov. Chachapoyas, along Rio Ventilla, W of Molinopampa, Wurdack 1510 (F, GH, uc, us). Cuzco: Rio Lachac, Sues 1813 (GH, us). Prov. Urubamba, Ma- chu Picchu, lit is et al. 1061 (GH, uc, us). Puno: Sandia, Weberbauer 797 (B). 1 7h. Hymenophyllum crispum var. bipinnatisec- tum Morton, Contr. U.S. Natl. Herb. 29: 161. 1947. TYPE: Peru, Cuzco, Alturas de Sieve, TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 67 Bties 1580 (holotype, us!; isotype, GH!; photo, us). Endemic. Thus far known only from the type and one other collection from Peru: Cuzco: Alturas de Sieve, Bues 1528 (us). 18. Hymenophyllum valvatum Hooker & Grev., Icon. fil. 2, /. 279. 1831. TYPE: Ecuador, for- ests of Esmeraldas, on trunks of trees, Jame- son (holotype, K; possible isotype, K!). Hymenophyllum crispatulum Bosch, Ned. Kruidk. Arch. 4: 412. 1859. TYPE: Peru, Tatanara (Puno), Lechler 2530 (holotype, L?; isotypes, B!, P!; pho- tos, GH & us of P). Hymenophyllum platylobum Bosch, Ned. Kruidk. Arch. 5:189.1863. TYPE: Peru, Dept. Puno, San Gaban, Lechler 2489 (apparently cited in error originally as 2498) (holotype, P; isotype, F!; photo, us! of P). Sphaerocionium valvatum (Hooker & Grev.) Copel., Philipp. J. Sci. 67: 31. 1938. Leaves determinate, to 20 cm longhand 6 cm broad. Petiole of mature leaves (2-)3-lOcm long, 0.4-1.0 mm in diameter, commonly alate just be- low lamina base, pubescent with simple or forked trichomes or glabrate. Lamina pinnate-pinnatisect to nearly 2-pinnate-pinnatifid, sparsely to mod- erately pubescent, the rachis alate throughout, the wing somewhat undulate. Pinnae 8-1 5 pairs, ovate- to elliptic-lanceolate, incised deeply to the costa into bifid or pinnatifid pinnules, ultimate segments slightly to strongly undulate, the veins (abaxially) and margins sparsely to amply pubescent, but tri- chomes lacking on tissue between veins and mar- gins. Indusia commonly longer than broad, and slightly longer than the segment tips. On tree trunks in wet forests, 900-1800 m, Huanuco, Junin, Puno. Lesser Antilles; Guyana to Colombia, south to Bolivia. It is with some reluctance that this is maintained as distinct from Hymenophyllum microcarpum, from which it seems to differ only in the undulate ultimate segments and (sometimes) rachis wings, and the character of the petiole trichomes. Even these features are variable, as only in a few spec- imens examined are the segments and rachis wings markedly undulate to crispate— most are only moderately so. Conversely, in a number of spec- imens determined to be H. microcarpum, the ul- timate segments are only slightly undulate. A per- fect intermediate is the isotype of H. platylobum, which is placed here in synonymy with H. val- vatum. The petiole trichomes of this specimen match those of//, microcarpum, but the segments and rachis wings are scarcely or only slightly un- dulate. Other features considered by Morton and subsequent workers seem not to be significant, at least in specimens from Peru. Furthermore, both species may be only robust forms of H. hirsutum, which seems to differ chiefly in its smaller lamina and more delicate petiole. The entire complex is in need of detailed study. Huanuco: Rio Llullapichis watershed, Cerros del Sira, Dudley 13339 (GH). Junin: Chanchamayo Valley, C. Schunke 1491, 1559 (F, us). Puno: "St. Gavan" (San Gaban), Lechler (K). Prov. Sandia, Chunchusmayo, We- berbauer 1237 (B). 19. Hymenophyllum microcarpum Desv., Mem. Soc. Linn. Paris 6: 333. 1827. TYPE: His- paniola (holotype, P; photo, uc). Hymenophyllum beyrichianum Kunze, Linnaea 9: 108. 1834. TYPE: Peru, Huanuco, Pampayacu, Poep- pig (holotype, LZ destroyed). Mecodium microcarpum (Desv.) Copel., Philipp. J. Sci. 67: 25. 1938. Sphaerocionium microcarpum (Desv.) Copel., Phil- ipp. J. Sci. 67: 34. 1938. Leaves determinate, to 30 cm long and 10 cm broad. Petiole of mature leaves (3-)5-15 cm long, 0.6-1.0 mm in diameter, broadly alate just below lamina base, pubescent with simple, forked, and stellate trichomes or glabrate. Lamina pinnate- pinnatisect to nearly 2-pinnate-pinnatifid, sparse- ly to moderately pubescent, the rachis conspicu- ously alate throughout. Pinnae 10-16 pairs, ovate- to elliptic-lanceolate, incised deeply or nearly to the costa into bifid or pinnatifid pinnules, the ul- timate segments entire or deeply bifid, the veins (abaxially) and margins sparsely to moderately pu- bescent, but trichomes lacking on tissue between the veins and margins. Indusia commonly longer than broad, and slightly broader than the segment tips. On tree trunks and clay banks in wet forests, 1 500-2700 m, Amazonas, Huanuco to Cuzco. Greater Antilles; Guatemala to Panama; Guy- ana to Colombia, south to Bolivia and Brazil. 68 FIELDIANA: BOTANY With this probably might be included H. val- vatum, under which see for further discussion. Amazonas: Prov. Bagua, E of La Peca, Barbour 2547 (MO). Huanuco: Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13290B (MO). Pasco: Prov. Ox- apampa, Palmazu, van der Werff et al. 8410 (MO, uc). Junin: Chanchamayo Valley, C. Schunke 463 (f, us). Cuzco: Prov. La Convention, Bites 2018, 2035 (us), 2034 (GH). Prov. La Convention, Alturas de Pintobamba, Vargas 3294 (MO, us). 20. Hymenophyllum ruizianum (Klotzsch) Kunze, Hot. Zeit. (Berlin) 1847: 199. 1847. Sphaerocionium ruizianum Klotzsch, Linnaea 18: 535. 1 844. TYPE: Peru, Ruiz 85 (holotype, B!; isotype, MA?, P; photos, GH & us of P). Leaves determinate, to 45 cm long and 12 cm broad. Petiole of mature leaves (7-) 1 0-20 cm long, 0.7-1.5 mm in diameter, nonalate, densely pu- bescent with mostly simple or forked trichomes. Lamina 2-3-pinnatifid, sparsely to amply pubes- cent on both sides of the lamina, the rachis alate distally. Pinnae commonly patent or their tips drooping, 10-18 pairs, ovate-lanceolate, incised nearly to the costa into pinnatifid or 2-pinnatifid pinnules, the ultimate segments entire or bifid, the veins and margins sparsely to moderately pubes- cent, but trichomes lacking on tissue between veins and margins. Indusia varying from broader than long to longer than broad, commonly slightly broader than the segment tips. In wet forests, on tree trunks and banks of rav- ines, and in sphagnum on forest floor, 1000-3300 m, Amazonas, Huanuco, Pasco, Cuzco, Puno. Colombia; Ecuador; Peru; Bolivia. A singular feature of this species is the tendency for the long, flexible pinnae to droop gracefully at their tips, a character which is usually preserved in dried specimens. Perhaps its nearest relative is H. trapezoidale Liebm., from Mexico and north- ern South America, which is a much smaller fern with less dissected pinnae which are rather strong- ly ascending (never drooping) and with more del- icate, merely sparsely pubescent, petioles. Amazonas: Prov. Chachapoyas, above Leimebamba, Hutchison & Wright 5553 (F, GH, MO, uc, us). Huanuco: Playapampa, Macbride 4516 (F, us). Pasco: Prov. Oxa- pampa, San Alberto, Cordillera de Yanachaga, van der Werff et al. 8423 (MO), 8439, 8491 (MO, uc). Cuzco: Prov. La Convention, Dudley 10790 (F, GH, MO). Puno: San Gaban, Lechler 2260 (B). 21. Hymenophyllum simplex Morton, Contr. U.S. Natl. Herb. 29. 171. 1947. TYPE: Peru, Huanuco, Tambillo, Jelski 897 (us!). Sphaerocionium simplex (Morton) Pic.-Ser., Webbia 28: 472. 1973. Leaves determinate, fragile and small, 5-7 cm long. Petiole slender, 0.1-0.15 mm in diameter, nonalate, glabrate or sparsely pubescent with sim- ple to stellate trichomes. Lamina simply pinnate or pinnatisect, moderately pubescent, the rachis narrowly alate distally, nonalate at base or in the proximal half. Pinnae few, ascending, simple, lin- ear, entire, the tissue and veins amply provided with orange, stellate trichomes. Veins lacking ac- cessory wings. Indusia about as broad as long, usu- ally narrower than the pinna tips. Endemic. Besides the type (Huanuco) thus far known only from two other collections: Cajamarca and San Martin. Habitat and elevation not re- corded. This delicate little fern is distinctive among the species of subg. Leptocionium in its few, ascend- ing, linear, entire pinnae. Although we now know it only from two Peruvian collections, its small, inconspicuous leaves make it difficult to detect. It is hoped that more diligent search will uncover additional specimens throughout Peru and, per- haps, in other areas of South America as well. Cajamarca: Cutervo, Jelski 914 (P, us). San Martin: Tarapoto, Spruce (K), collection is without number, but with notation "conf. 4048"; #4048 is type number of H. tarapotense. 22. Hymenophyllum fragile (Hedwig) Morton, Contr. U.S. Natl. Herb. 29: 172. 1947. Trichomanes fragile Hedwig, Fil. gen. sp., t. 18. 1802. TYPE: based on illustration from plant collected in "America meridionalis." Sphaerocionium fragile (Hedwig) Pic.-Ser., Webbia 28: 471. 1973. Leaves determinate, (3-)6-18 cm long, 1.5-2.5 cm broad. Petiole ca. 0.2 mm in diameter, sparsely to moderately pubescent with simple, forked, or often stellate trichomes. Lamina pinnate-pinna- tisect, moderately pubescent, the rachis broadly and consistently alate throughout. Pinnae (3-)5- 1 2 pairs, deeply 1 -forked (distally) to pinnatisect, the 1-2 pairs of segments entire, the tissue and TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 69 veins moderately provided with orange, short- stalked, stellate trichomes. Veins lacking accessory wings. Indusia about as broad as long, usually nar- rower than the ultimate segments, the receptacle not exserted. In dense forests, on tree trunks or mossy banks, 800-2700 m, Pasco, Junin, Ucayali, Cuzco. Mexico to Panama; Greater Antilles; Venezuela and Colombia to Bolivia and Brazil. This may be confused with Hymenophyllum ele- gantulum, under which see for further discussion. Pasco: Oxapampa, Soukup 2650 (GH). Junin: East of Quimiri Bridge, near La Merced, Killip & Smith 23834 (NY, us). Ucayali (as Loreto): Prov. Coronel Portillo, NE of pass at La Divisoria, Skog et al. 5156 (us). Cuzco: Prov. La Convencion, Pintobamba, Vargas 3262 (MO, us). 23. Hymenophyllum elegantulum Bosch, Ned. Kruidk. Arch. 4: 408. 1859. LECTOTYPE (designated as type by Morton, Contr. U.S. Natl. Herb. 29: 170. 1947): based on t. 33a (Hooker, Sp. fil. 1 . 1 844), which in turn, ac- cording to Morton, was based on Ecuador, Pillzhum, Jameson 366 (K.?; probable isolec- totype, NY!). Hymenophyllum pulchellum sensu Hooker, Sp. fil. 1, /. 33a. 1844, not Schlecht. & Cham. 1830. Sphaerocionium elegantulum (Bosch) Copel., Philipp. J. Sci. 67: 32. 1938. Leaves indeterminate, 12-35 cm long, 2-4 cm broad. Petiole ca. 0.2 mm in diameter, glabrate or sparsely pubescent with simple or forked tri- chomes. Lamina pinnate-pinnatisect, amply pu- bescent, the rachis flexuous, nonalate, or some- times alate in the distal portion. Pinnae subdistant to remote, ascending, 8-24 pairs, deeply pinnati- sect, the 4-6 pairs of segments 1-2-forked or bifid, the tissue and veins moderately provided with or- ange, sessile to short-stalked, stellate trichomes. Veins lacking accessory wings. Indusia about as broad as long, and usually as broad as the ultimate segments. On tree trunks, in forests and wet clearings, 3000- 3600 m, Amazonas, Cuzco, Puno. Mexico to Panama; Greater Antilles; Venezuela and Colombia south to Peru and Bolivia. Some leaves of Hymenophyllum elegantulum vary somewhat in the character of the rachis wing, as in the case of Biies 720, cited below. Some of the laminae have the rachis narrowly winged well into the proximal half, and these specimens might be confused with H. fragile. However, the alate condition in these atypical leaves is irregular and mostly due to the decurrent bases of pinnae, whereas in H. fragile the wings are very conspic- uous and of nearly consistent width throughout. Ama/onas: Prov. Bagua, Cordillera Colan NE of La Peca, Barbour 3415 (F, MO). Cuzco: Michihuanuncca, Biies 720 (us). Puno: Prov. Carabaya, Ayapata, Vargas 1071 7 (GH). 24. Hymenophyllum amabile Morton, J. Wash. Acad. Sci. 22: 63. 1932. TYPE: Peru, Cuzco, Prov. La Convencion, Huadquina, Bites 715 (holotype, us!; isotype, GH!; photo, F of us). Leaves indeterminate, to 40 cm long and 4 cm broad. Petiole 0.2-0.3(-0.4) cm in diameter, non- alate, sparsely pubescent with mostly stellate tri- chomes. Lamina pinnate-pinnatisect, densely to- mentose, the rachis nonalate throughout, or rarely slightly alate due to decurrent bases of pinnae. Pinnae numerous, oblong or ovate-oblong, obtuse, often incised nearly to the costa, the 4-8 pairs of segments entire to forked, ultimate divisions en- tire, the tissue and veins often obscured by a dense tomentum of orange, stalked, stellate or bistellate trichomes. Veins lacking accessory wings. Indusia broader than long, about as broad as the segment tips, the receptacle very short, not exserted. Pendent from tree trunks and branches in wet forests, 2600-4200 m, Huanuco, Cuzco, Puno. Ecuador and Peru. Huanuco: Tambo de Vaca, Bryan 628 (F, us). Cuzco: Prov. La Convencion, Vilcabamba, Biies 2108, 2109, 2110 (us). Prov. La Convencion, Loma Grande, Biies 2148 (F, GH, uc, us). Urubamba, Machu Picchu, Peyton & Peyton 1 104 (MO). Puno: Sandia, Weberbauer 796 (B). 25. Hymenophyllum speciosum Bosch, Ned. Kruidk. Arch. 5: 181. 1863. LECTOTYPE (designated as type by Morton, Contr. U.S. Natl. Herb. 29: 175. 1947): Peru, Puno, San Gaban, on tree trunks, Lechler 2246 (L?; iso- lectotypes, B!, F!, p; probable isolectotype, NY!; photos, GH, us of P). Hymenophyllum spectabile Kuhn, Linnaea 35: 392. 1868. TYPE: Bolivia, Yungas, D'Orbigny 175 (holotype, B?; isotype, p; photo, us of P). 70 FIELDIANA: BOTANY Sphaerocionium spectabile (Kuhn) Copel., Philipp. J. Sci. 67:31. 1938. Leaves indeterminate, mature ones 30-180 cm long, 4-9 cm broad. Petiole (of mature leaves) 0.5- 0.7 mm in diameter, nonalate, glabrate or sparsely pubescent with simple to stellate trichomes. Lam- ina pinnate-pinnatifid or pinnate-pinnatisect, densely tomentose, the rachis nonalate through- out, commonly flexuous, with a tightly appressed covering of subsessile or short-stalked, stellate tri- chomes. Pinnae numerous, larger ones (2.5-)3-8 cm long, lanceolate or linear-lanceolate, often at- tenuate, incised deeply or nearly to the costa, the 6-14 pairs of segments shallowly to deeply bifid, the ultimate divisions entire, the tissue and veins mostly obscured by a dense tomentum of orange (aging whitish), short-stalked, stellate trichomes. Veins lacking accessory wings. Indusia about as broad as long and often broader than the segment tips, the receptacle not exserted. Pendent from trees or clay banks in wet forests, 1800-3350 m, Amazonas, Pasco, Cuzco, Puno. Peru and Bolivia. This might better be treated as a variety ofHy- menophyllum karstenianum, as the two differ only quantitatively, e.g., appressed versus spreading to- mentum, strongly versus slightly flexuous rachis, relative size and number of segments of pinnae. Amazonas: Prov. Chachapoyas, Hutchison & Wright 5569 (F,GH, MO, uc, us); Sagastegui 7453 (F, HUT). Pasco: Prov. Oxapampa, Cordillera San Gutardo, Leon et al. 534(usM). Cuzco: Vallede Lares, Biies 1808, 1817, 1821, 1931 (us). Valle de San Miguel, La Convention, Biies 2178 (GH, MO, us). 26. Hymenophyllum karstenianum J. W. Sturm, Bot. Zeit. (Berlin) 17: 298. 1859. LECTO- TYPE (designated as type by Morton, Contr. U.S. Natl. Herb. 29: 176. 1947): Venezuela, Merida, Moritz 381 p.p. (BR). Sphaerocionium karstenianum (J. W. Sturm) Pic.-Ser., Webbia28:471. 1973. Leaves indeterminate, mature ones 15-50 cm long, 2-3.5(-4) cm broad. Petiole 0.4-0.5 mm in diameter, nonalate, glabrate or sparsely pubescent with simple to stellate trichomes. Lamina pinnate- pinnatifid to -pinnatisect, densely tomentose, the rachis nonalate throughout, straight or slightly flexuous, abundantly to densely covered with short- to long-stalked stellate trichomes, these mostly spreading, not tightly appressed. Pinnae numer- ous, larger ones 1-2.3 cm long, lanceolate or lin- ear-lanceolate, acute to subattenuate, shallowly or deeply incised, the 4-7 pairs of segments entire to bifid, the tissue and veins mostly obscured by a dense tomentum or orange to tawny, short-stalked, stellate trichomes. Veins lacking accessory wings. Indusia commonly as broad as long, and as broad as the segment tips, the receptacle not exserted. Pendent from trees or clay banks in wet forests, 700-1900 m throughout the range, but thus far apparently known in Peru only from one collection cited below. Venezuela; Colombia; Peru. This and Hymenophyllum speciosum are very closely related. See discussion of the latter for ad- ditional comments. San Martin: Near Tarapoto, in Monte Campana, Spruce 4694 (BR, us). 27. Hymenophyllum lindenii Hooker, Sp. I'll. 1: 94, t. 34c. 1844. TYPE: Venezuela, Caracas, Linden 173 (holotype, K!; isotype, P; photos, F of K, us of P). Sphaerocionium lindenii (Hooker) Vareschi, Flora Venezuela 1: 217. 1969. Leaves indeterminate, to 35 cm long and 8-15 cm broad. Petiole (8-) 10-1 5 cm long, 0.7-1 .2 mm in diameter, commonly nonalate, pubescent with simple to stellate trichomes. Lamina ovate or ovate- lanceolate, not or slightly reduced at base, nearly 2-pinnate-pinnatisect, moderately pilose, the rachis abundantly stellate-pubescent, scarcely alate, or alate distally (or sometimes to base). Pinnae patent to ascending, 14-20 pairs, 4-9 cm long, acuminate to subattenuate, incised nearly to the costa into 8- 14 deeply pinnatifid pinnules, the ultimate seg- ments entire to deeply bifid, rarely undulate or crispate, tissue between the veins sparsely provid- ed with scattered, sessile or stalked, stellate tri- chomes. Veins lacking accessory wings. Indusia broader than long, and about as broad as the seg- ment tips, receptacle not exserted. Pendent from tree trunks in wet forests, 2300- 2450 m, Amazonas, Puno. Venezuela; Colombia; Ecuador(?); Peru. Hymenophyllum lindenii is one of the most ro- TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 71 bust species in the subgenus, the long, stout petiole often measuring a full millimeter in diameter. The presence of a rachis wing, usually a good diagnostic feature in most species of subg. Leptocionium, is not consistent in this or in the closely related H. plumieri. The rachis in H. lindenii is occasionally alate to the base of the lamina; more commonly the wing is present in the distal half but sometimes it is nearly lacking throughout. The two Lechler specimens cited below are most atypical, being broadly alate even well down the petiole, and with the wings very strongly crispate. The latter feature apparently was what prompted Mettenius's orig- inal annotation on this sheet as "//. valvatum." Lechler 2489 is a mixed collection, as the other four specimens of 2489 at Berlin are isotypes of H. platylobum (= H. valvatum). Amazonas: Prov. Bagua, Cordillera Colan, Barbour 3554 (MO, USM), 3602, 3751 (MO). Puno: near San Gaban, Lechler 2489 (u). Tatanara, Lechler 2563 (B, K). 28. Hymenophyllum plumieri Hooker & Grev., Icon. fil. 2, /. 123. 1829. TYPE: Ecuador, western side of Pichincha, Jameson (holo- type, K.!; photo, F). Hymenophyllum interruptum Kunze, Linnaea 9: 107. 1834. TYPE: Peru, Huanuco, Pampayacu, Poep- pig 1104 (holotype, w?; isotype, HBG). Sphaerocionium plumieri (Hooker & Grev.) Presl, Hy- menophyllaceae 34. 1 843. Sphaerocionium interruptum (Kunze) Presl, Hymen- ophyllaceae 34. 1843. Leaves indeterminate, to 50 cm long and 3-8 cm broad. Petiole 2-7 cm long, 0.4-0.6(-0.7) mm in diameter, nonalate (or sometimes alate just at the apex), pubescent with mostly forked to stellate trichomes. Lamina linear, usually strongly and gradually reduced at base, often interruptedly fer- tile (e.g., groups of fertile pinnae separated by some sterile ones), pinnate-pinnatifid to -pinnatisect, moderately pilose, the rachis alate throughout, or occasionally alate only distally or not at all. Pinnae patent to ascending, 1 5-many pairs, 1 .5-6 cm long, acute to subattenuate, the 5-12 pairs of segments simple and entire to deeply bifid, the tissue be- tween the veins sparsely to moderately provided with sessile or stalked, stellate trichomes. Veins lacking accessory wings. Indusia about as broad as long or usually broader, as broad as the segment tips. Pendent from tree trunks and clay banks in wet forests, 600-2200 m, San Martin to Puno. Venezuela and Colombia to Peru and Bolivia. It is not possible to separate this satisfactorily from Hymenophyllum interruptum, which was said to differ in its groups of fertile pinnae interrupted by a few sterile ones, by its rachis not alate toward the base, and by variations in lamina base and in marginal trichomes. In a number of specimens examined during this study (identified as H. in- terruptum) fertile pinnae are continuous, and rach- ises are fully to partially alate or (occasionally) nonalate throughout. The entire species complex, which also involves H. dependens Morton and H. superbum Morton from northern South America, needs closer examination. All may be conspecif- ic. San Martin: In monte Guayrapurima, prope Tara- poto, Spruce 4025 (K, P). Huanuco: Rio Llullapichis wa- tershed, ascent of Cerros del Sira, Dudley 13014 (GH, us), 13204, 13377 (GH). Junin: Colonia del Perene, Killip & Smith 24955 (F, GH, NY, us). Cuzco: Prov. Paucartam- bo, Pilahuata, Vargas 4914 (MO, us), 76756, 76767 (GH). Madre de Dios: Prov. Manu, Shintuya, Chavez 861 (MO). Puno: Prov. Carabaya, Vargas 17562 (GH). 29. Hymenophyllum plumosum Kaulf., Enum. fil. 267. 1824. TYPE: Brazil, Chamisso (holo- type, LZ destroyed; isotype, LE?). Sphaerocionium plumosum (Kaulf.) Copel., Philipp. J. Sci. 67: 30. 1938. Leaves indeterminate, to 75 cm long and 6 cm broad. Petiole 0.3-0.7 mm in diameter, nonalate, pubescent with simple to stellate trichomes or gla- brate. Lamina pinnate-pinnatifid to rarely nearly 2-pinnate, densely appressed-tomentose, the rach- is nonalate throughout. Pinnae numerous, lanceo- late, pinnatifid to pinnatisect, or rarely incised to the costa, the ultimate segments entire (some prox- imal ones bifid), the tissue and veins completely obscured by a tomentum of sessile to short-stalked, stellate trichomes. Veins (especially abaxially) with low, inconspicuous lamellae. Indusia about as broad as long, usually slightly broader than the segment tips. Epiphyte on trunks or branches in dense, wet forests, or occasionally pendent from wet, clay banks, 550-3100 m, Huanuco, Junin, Cuzco, Madre de Dios, Puno. Costa Rica to Colombia; Peru; Bolivia; Brazil. 72 FIELDIANA: BOTANY The lamellae along the veins in this species are scarcely "wings" in the sense of being broad and thin flanges of tissue. Instead, they are commonly long, low outgrowths borne along the abaxial or (sometimes) adaxial sides of the veins contiguous to, but not in the same plane with, the segment tissue. They are often difficult to discern, since the veins and laminar surface are so densely tomen- tose. The best way to observe them is on the older (proximal) pinnae where some of the indument may have fallen away. For further discussion see also Hymenophyllum tomentosum. Easily confused with Hymenophyllum plumo- sum are H. karstenianum and H. speciosum, which have no lamellae on the veins, but are similar to the former in most other features, including the dense matting of tomentum obscuring the laminar surface. Extreme care and patience must be ex- ercised to determine whether the low outgrowths are present or lacking. In fact, the real importance of the presence or absence of lamellae bears more careful analysis in the future. During this study of Peruvian species of Hymenophyllum, it has been noted that size and frequency of these outgrowths, on one or both surfaces, may be more variable than earlier assumed, and thus may be of much less taxonomic significance. Huanuco: Vilcabamba, Hacienda on Rio Chinchao, Macbride 5146 (F, us). Junin: Schunke Hacienda above San Ramon, Killip & Smith 24840 (F, GH, us). Cuzco: Prov. Paucartambo, above Tambomayo, West 7108 (MO, uc). Madre de Dies: Prov. Mania, Shintuya, Chavez 860 (MO). Puno: San Gaban, Lechler 2264 (B). 30. Hymenophyllum mult ialat urn Morton, Contr. U.S. Natl. Herb. 29: 185. 1947. TYPE: Peru, Cuzco, Prov. La Convencion, Alturas de Sieve, Biies 1575 (holotype, us!; frag., GH!). Sphaerocionium multialatum (Morton) Pic.-Ser., Webbia28: 471. 1973. Leaves indeterminate, to 50 cm long, 3-8(-15) cm broad. Petiole 0.4-0.7 mm in diameter, non- alate, sparsely hirsute with mostly simple tri- chomes. Lamina pinnate-pinnatisect, hirsute, the rachis nonalate throughout. Pinnae numerous, narrow-oblong or -deltoid, pinnatisect, sometimes irregularly and greatly elongate, the ultimate seg- ments commonly bifid, the tissue and veins mod- erately provided with sessile to short-stalked ses- sile trichomes. Veins bearing (especially abaxially) irregularly interrupted, conspicuous lamellae. In- dusia about as broad as long, equaling or slightly broader than the segment tips. Epiphyte in dense, wet forests, 2000-3200 m, Amazonas, Huanuco, Pasco, Cuzco. Colombia; Ecuador; Peru. Shape of lamina and pinnae can be variable and irregular in this species. Typically the leaves are linear, with numerous pinnae 3 or 4 cm long, yet at times, scattered along the rachis, there may be a few greatly elongated, attenuate pinnae, occa- sionally reaching 1 5 cm in length. Although Hy- menophyllum multialatum has been thus far re- ported from four departments in Peru, the great majority are represented by Biies's collections in Cuzco. For further discussion see H. tomentosum. Amazonas: Prov. Bagua, Cordillera Colan, Barbour 3740 (F, MO). Huanuco: Prov. Huanuco, 5 km from Car- pish, Tryon & Tryon 5318-B (GH). Pasco: Prov. Oxa- pampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8495 (MO). Cuzco: Valley of Rio Urubamba, Biies A- 16 (F, GH, us). Montana de Calca, Biies 1916 (GH, us), 1919, 1923, 1932 (us). Prov. La Convencion, Cor- dillera Vilcabamba, Dudley 1 1 137 (GH, us). 31. Hymenophyllum tomentosum Kunze, Lin- naea 9: 107. 1834. TYPE: Peru, Huanuco, Pampayacu, 1829, Poeppig Diar. 1134 (ho- lotype, w?; isotype, BM?; frag., B!; probable isotype, MO!; photo, us of MO). Sphaerocionium tomentosum (Kunze) Presl, Hymen- ophyllaceae 34. 1843. Hymenophyllum fusagasugense J. W. Sturm, Bot. Zeit. (Berlin) 1859: 297. 1859 (as "fusugasugense"). TYPE: Colombia, Cundinamarca, "Fusugasuga" (Fusagazuga) Karsten (not located). Hymenophyllum fusagasugense var. aberrans Mor- ton, Contr. U.S. Natl. Herb. 29: 187. 1947 (as "fusugasugense"). TYPE: Venezuela, Merida, Tabay, Gehriger 584 (holotype, us!; isotypes, F!, GH!). Leaves indeterminate, to 50 cm long and 2.5 cm broad. Petiole 0.5-0.6 mm in diameter, nonalate, sparsely pubescent with mostly simple trichomes. Lamina pinnate-pinnatifid, densely hirsute, the rachis nonalate throughout. Pinnae numerous, ob- long or narrow-deltoid, pinnatifid (sometimes deeply so), the ultimate segments simple or (oc- casionally) bifid, the tissue and veins densely cov- ered with (and often obscured by) sessile to short- stalked trichomes. Veins abaxially bearing irreg- ularly interrupted, conspicuous winglike lamellae, TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 73 these wings much smaller or sometimes lacking adaxially. Indusia about as broad as long, equaling or slightly broader than the segment tips. On steep banks and tree trunks, 2750-3100 m, Amazonas, Huanuco. Colombia; Venezuela; Peru. This is distinguished from Hymenophyllum multialatum and H. plumosum only by the char- acters used in the key. All three appear to be alike, except for the size and frequency of the lamellae along the veins and by the relative density of pu- bescence. Of these, H. plumosum (q.v.) is the most distinct, in that the lamellae are merely low and inconspicuous outgrowths and the trichomes are so densely matted as to completely obscure the veins and tissue. In H. tomentosum the indument is also densely matted, but the lamellae are so large and conspicuous that many of them emerge through the tomentum. Morton (1947) chose to separate H. fusagasugense and var. aberrans on the pres- ence or absence of these processes adaxially. How- ever, the size or lack of lamellae on the adaxial side varies considerably in the species, and the character is difficult to observe due to the dense surface tomentum. To prove that the lamellae are completely absent on a given specimen would re- quire scraping away all the tomentum of each leaf. A thorough study of the species complex through- out its range is needed. Amazonas: Prov. Chachapoyas, Cerros de Calla Calla, 19 km above Leimebamba, Hutchison & Wright 5570 (uc). Prov. Chachapoyas, near km 422, Balsas-Leime- bamba road, Wurdack 1249 (F, GH, uc, us). 32. Hymenophyllum lobatoalatum Klotzsch, Lin- naea 20: 438. 1847. TYPE: Peru, Panatahua, Ruiz 83 (holotype, B!; isotype, MA?). Sphaerocionium lobatoalatum (Klotzsch) Pic.-Ser., Webbia28:471. 1973. Leaves indeterminate, to 60 cm long and 8 cm broad. Petiole 0.4-0.7 mm in diameter, alate, if at all, only near base of lamina, pubescent with simple to stellate trichomes or glabrate. Lamina pinnate-pinnatifid, pubescent with stellate tri- chomes, the rachis strongly alate throughout. Pin- nae numerous, larger ones 20-45 mm long, 7-1 1 mm broad at base, most of them narrow-trian- gular, the apex narrowly acute to subattenuate, deeply pinnatind, the 7-12 pairs of ultimate seg- ments bifid, the tissue and veins moderately pro- vided with light or dark brown, short-stalked, stel- late trichomes. Veins abaxially bearing small, scattered lamellae, these about as broad as long, but lacking adaxially. Indusia slightly longer than broad, usually as broad as the segment tips, re- ceptacle not exserted. In wet forests, pendent from trunks and branch- es of trees, 600-1300 m, Loreto and Huanuco. Colombia; Ecuador; Peru. There is little, beyond the key characters, to distinguish this from Hymenophyllum pyrami- datum. Leaves of the latter are perhaps more ro- bust, with relatively narrower pinnae and much denser pubescence. Both also share nearly the same range, and more study may prove them to be con- specific. Also very closely related is H. verecun- dum, which is more easily separated by its much lighter colored trichomes and shorter, relatively broader and more crowded pinnae. But even these characters sometimes tend to be intermediate. Loreto: Pumayucu, between Balsapuerto and Moy- obamba, Klug 3248 (B, F, GH, MO, us). Huanuco: Prov. Huanuco, Dist. Churubamba, Mt. Santo Toribio, Mexia 8258 (B, F, GH, uc, us). 33. Hymenophyllum pyramidatum Desv., Mem. Soc. Linn. Paris 6: 332. 1827. TYPE: "habitat in America" (possibly a Dombey collection from Peru) (holotype, P; photos, GH, us). Sphaerocionium pyramidatum (Desv.) Copel., Phil- ipp. J. Sci. 67: 30. 1938. Leaves indeterminate, to 65 cm long and 6 cm broad. Petiole 0.5-0.7 mm in diameter, often alate 1-2 cm near the apex, pubescent with simple to stellate trichomes. Lamina pinnate-pinnatifid, densely pubescent with simple to stellate tri- chomes, the rachis alate throughout. Pinnae nu- merous, larger ones 20-45 mm long, 6-1 1 mm broad at base, most of them narrow-triangular, the apex narrowly acute to attenuate, deeply lobed to pinnatifid, the 6-12 pairs of ultimate segments simple to bifid, the tissue and veins usually co- piously provided with orange to dark brown (rare- ly tawny), subsessile to short-stalked, stellate trichomes. Veins abaxially with numerous, con- spicuous, elongate, winglike lamellae (often nearly the length of the vein), these sometimes less con- spicuous and abundant on the adaxial surface. In- dusia slightly longer than broad, usually as broad as the segment tips, receptacle not exserted. 74 FIELDIANA: BOTANY In dense, wet forests, pendent from stumps, tree trunks, and branches, and on wet banks, 600-1 800 m, along Cordillera Central from San Martin to Cuzco. Colombia; Peru; Bolivia. This belongs to a close-knit complex of species involving Hymenophyllum verecundum and H. lobatoalatum. See discussion of the latter for fur- ther comments. San Martin: Tingo Maria, Allard 20818 (us). Huan- uco: Pampayacu. Kanehira 123 (F, GH, K, us). Pasco: Prov. Oxapampa, 4-8 km from Enenas, Skoget al. 5077 (us). Junin: Chanchamayo Valley, C. Schunke 466, 926 (F. us), 7055 (F). Cuzco: Prov. Paucartambo, between Mistiana and Keros, Vargas 7380 p.p. (MO). 34. Hymenophyllum verecundum Morton, Contr. U.S. Natl. Herb. 29: 183. 1947. TYPE: Peru, Huanuco, Churubamba, Mexia 8143-A (ho- lotype, us!; isotypes, GH!, MO!, uc!). Leaves indeterminate, to 50 cm long and 4 cm broad. Petiole 0.4-0.6 mm in diameter, alate, if at all, just below the lamina, glabrate or slightly pubescent with simple to stellate trichomes. Lam- ina pinnate-pinnatifid, densely pubescent with stellate trichomes, the rachis alate throughout, or sometimes nonalate at base. Pinnae numerous, larger ones 6-20 mm long, 3-7 mm broad at base, oblong to very broadly triangular, the apex obtuse to subacute, pinnatifid, the 3-6(-8) pairs of ulti- mate segments simple to retuse or occasionally bifid, the tissue and veins copiously provided with whitish to tawny, subsessile or short-stalked, stel- late trichomes. Veins abaxially with numerous conspicuous, elongate, winglike lamellae, these less conspicuous and abundant on the adaxial surface. Indusia about as long as broad and usually as broad as the segment tips, receptacle not exserted. In dense, wet forests, pendent from tree trunks, branches, and wet banks, (5 50-) 1500-2 700 m, Amazonas, Huanuco, Junin, Cuzco, Madre de Dios, Puno. Colombia; Peru; Bolivia. This is closely related to Hymenophyllum pyr- amidatum and H. lobatoalatum. See discussion of the latter for further comments. (us). Madre de Dios: Prov. Manii, Shintuya, Chavez 833 p.p. (MO). Puno: Prov. Sandia, E of Oconeque, Hodge 6090 (F, GH, us). 35. Hymenophyllum tarapotense Stolze, sp. nov. Folia usque ad 6 cm longa et 1.6 cm lata. Petiolus 1- 2 cm loneus. 0.1-0.15 mm diametro. Lamina pinnato- pinnatifida. Rhachis versus basin non alata. Pinnae 6- 12-jugatae, bifidae vel pinnatifidae, usque ad 1 cm lon- gae, pinnae basales admodum reductae, marginibus, venis et interveniis trichomatibus stellatis instructis. Venae lamellis cristiformibus numerosis praeditae. Leaves indeterminate, to 6 cm long and 1.6 cm broad. Petiole 1-2 cm long, 0.1-0.15 mm in di- ameter, nonalate, sparsely provided with simple trichomes. Lamina pinnate-pinnatifid, pubescent with stellate trichomes, the rachis weakly alate dis- tally, nonalate toward the base. Pinnae 6-12 pairs, bifid, or distal ones pinnatifid with 3-4 segments, larger ones 1 cm long, basal ones often reduced to mere auricles, the margins, veins, and intervening tissue moderately to amply provided with tawny trichomes. Veins bearing numerous, conspicuous accessory foliar crests not in the plane of the lam- ina, these less frequent on the adaxial surface. In- dusia about as long as broad, as broad as the seg- ment tips, receptacle minute, not exserted. TYPE— Peru, Tarapoto (Dept. San Martin) in monte Guayrapurima, Aug. 1856, Spruce 4048 (holotype, K!; photo, F). Another Spruce specimen at K, with notation "conf. 4048," is H. simplex. Known only from the type. This tiny fern is unquestionably the smallest of the species in subg. Leptocionium which bear ac- cessory tissue on their veins. These foliar processes are not long, low wings as seen in Hymenophyllum pyramidatum, but are separate, crestlike flanges of tissue similar to those frequently seen in more delicate specimens of//, verecundum. Superficially //. tarapotense closely resembles H. fragile, in its small size, delicate petiole, and reduced, often bi- fid, pinnae. It may seem strange that this distinc- tive species has not been described before, nor collected again in over 1 00 years. However, many of these diminutive members of the genus are very easily overlooked by collectors. Amazonas: Prov. Bagua. Cordillera Colan, Barbour 3740 (MO in part). Huanuco: Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13370 (GH, us). Junin: Pichis Trail. Porvenir, Killip & Smith 25948 (F, GH, us). Cuzco: Hacienda Pintobamba, Chaupimayo, Biies 1943 Comments Hymenophyllum peruvianum Hooker & Grev., Icon. fil. 2, /. 208. 1831. TYPE: Ecuador (as TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 75 Peru), Prov. Esmeraldas, Jameson (holotype, K). The original drawing seems to illustrate most of the features characteristic of Hymenophyllum fu- coides var. pedicellatum, except that the apex of the indusium is deeply dentate rather than lacin- iate. It is likely that the two taxa are synonymous, but we have not seen the type to verify this. The problem is but one of many in the H. fucoides complex begging solution through a complete re- vision of the group. Hymenophyllum poeppigianum Presl, Hymeno- phyllaceae 54. 1843. TYPE: Peru, Huanuco, Pampayacu, Poeppig (holotype, PR; probable isotype, P; photos, GH & us of P). Some authors have placed this with Hymeno- phyllum polyanthos. The type has not been seen, but judging from the protologue and photos of a probable isotype (P) the species is more likely to be H. myriocarpum. The protologue refers to: the indusia "ovate-subrotund"; the rachis alate throughout; the petiole alate toward the apex; the lamina linear-lanceolate with the base acute. The description of the indusia could conceivably fit either H. polyanthos or H. myriocarpum, but that of the lamina (as well as the photo of the possible isotype) more closely match H. myricarpum or H. undulatum. However, the petiole and (usually) the rachis base of H. undulatum are nonalate. Hymenophyllum sprucei Baker in Hooker & Bak- er, Syn. fil. 65. 1 867. TYPE: Peru, San Martin, Tarapoto, Spruce (not located). Morton (p. 155, 1947) considered that this is the same as Hymenophyllum adiantoides, and there is nothing in the description to argue otherwise. It is possible that Baker was unaware of the publi- cation of the latter species, and even that he based his new species on the same specimen cited earlier by van den Bosch. Specimens (B, GH, K, L, NY, us) thought to be probable isotypes of H. adiantoides are those of Spruce from Tarapoto. They bear the notation "conf. 4700" and are conceivably the ones on which Baker based his "//. sprucei." Hymenophyllum trifidum Hooker & Grev., Icon, fil. 2, 1. 196. 1 83 1 . TYPE: Ecuador (as "Peru"), Esmeraldas, Jameson (E?). In the original description the type was cited as having come from Peru, but Esmeraldas is an Ecu- adorean province in which Jameson did much of his collecting. Yet Hymenophyllum trifidum still might be expected in Peru, as it could be merely a form of H. elegans. From the description and plate it appears to be very similar to the latter, but until the type is located the question must remain unanswered. II. Trichomanes Trichomanes L., Sp. pi. 1097. 1753. TYPE: Tri- chomanes scandens L. Figure 12. Didymoglossum Desv., Mem. Soc. Linn. Paris 6: 330. 1827. TYPE: Didymoglossum muscoides (Sw.) Desv. (Trichomanes muscoides Sw.) = Trichom- anes hymenoides Hedwig. Trichomanes subg. Didymoglossum (Desv.) C. Chr., Index fil. xiv. 1906. Trichomanes subg. Achomanes Presl, Hymenophyl- laceae 15. 1843. TYPE: Trichomanes crispum L. Trichomanes subg. Pachychaetum Presl, Hymeno- phyllaceae 16. 1843. TYPE: Trichomanes rigi- dum Sw. Homoeotes Presl, Gefassbiindel Farm, 23. 1847. TYPE: Homoeotes heterophylla Presl = Trichom- anes humboldtii Lell. Plants epiphytic, occasionally epipetric or ter- restrial. Stem stout and erect or decumbent to long- creeping and filiform, bearing scattered to abun- dant trichomes and delicate to large, fibrous roots or (especially in subg. Didymoglossum) lacking roots. Leaves monomorphic or (less commonly) dimorphic, 0.5—40 cm or longer. Lamina entire to nearly 5 -pinnate, subsessile to petiolate, with pet- iole commonly shorter than the lamina, glabrous or with trichomes borne mostly on costae and margins. Veins free or, in Trichomanes diversi- frons, reticulate near the lamina margin, pinnate (anadromous or catadromous) or sometimes fla- bellate, false veins also present in some species, these mostly parallel, but in a few species perpen- dicular to the true veins. Sori terminal on the veins. Indusium mostly tubular, the mouth bilabiate or entire and often flaring, or very rarely nearly bi- valved, the sporangia borne on an elongate recep- tacle. Receptacle commonly exserted beyond the mouth of the indusium, often greatly so. There are about 300 species in the genus, oc- curring in tropical to subtropical regions of both hemispheres. As in Hymenophyllum, most are 76 FIELDIANA: BOTANY FIG. 12. Trichomanes radicans: a, habit; b, pinna with sori. Trichomanes pinnatum: c, portion of pinna with false veins and sori. Trichomanes hymenoides: d, apical portion of leaf, (a from Acosta-Solis 13782, Ecuador, F, b from Malhias & Taylor 5075, F, c from Barbour 5194, F, d from Wurdack 1513, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 77 found as epiphytes in dark, wet forests, but a num- ber of species may be found on wet, rocky cliffs or occasionally on clay banks. Also as in Hymen- ophyllum, Trichomanes has been divided by pre- vious authors into a number of genera and/or sub- genera. Only one genus is recognized here, with the key and species order reflecting as nearly as possible four of the subgenera. The monotypic subg. Cardiomanes (Presl) Christ occurs in New Zea- land. "False veins" may be found on laminae in two of the subgenera. These are long sclerenchymatous strands which can be observed as faint to rather distinct lines, parallel or perpendicular to the true veins. They may be connected to the true veins but are much more commonly free from them. Their position and frequency are important char- acters in recognizing some of the species. As in Hymenophyllum, veins of several species of Tri- chomanes bear crestlike lamellae, perpendicular to the plane of the lamina. One of these, T. mar- tiusii, occurs in Peru. The group of Trichomanes crispum L. in subg. Achomanes is the most taxonomically difficult in the genus, and a few of these species occurring in Peru may eventually be found to merit only infra- specific status. Some of the difficulties are very likely the result of yet unsolved questions involv- ing hybridization. References BOER, J. G. W. 1962. The new world species of Trichomanes sect. Didymoglossum and Micro- gonium. Acta Hot. Neerl., 11: 277-330. WINDISCH, P. G. 1977. Synopsis of the genus Trichomanes, subgenus Achomanes. Ph.D. the- sis, Department of Biology, Harvard Univer- sity, Cambridge, Mass., 198 pages. Key to Species of Trichomanes a. Venation anadromous, never flabellate; false veins lacking b b. Stem long-creeping; leaves well-spaced to remote [subg. Trichomanes] c c. Stems relatively stout, 0.7-2.2 mm in diameter; mature leaves 15-50 cm long d d. Pinnae 2-3-pinnatisect; stem amply to densely provided with castaneous trichomes; ultimate segments narrow, commonly linear e e. Indusium narrow-cylindrical or -elliptic, often subfusiform, the apex not or scarcely expanded; petiole (1— )4— 15 cm long; elevation 0-500 m 1. T. radicans e. Indusium conical, apex expanded into a wide-flaring mouth; petiole 0.3-3 cm long; elevation (400-)700-3200 m 2. T. collariatum d. Pinnae pinnatifid or pinnatisect (sometimes 2-pinnatifid as to basal segments); stem with trichomes sparse to widely scattered; ultimate segments relatively short and broad, not linear 3. T. rupestre c. Stems filiform, 0.2-0.5 mm in diameter, leaves 3-15(-18) cm long f f. Rachis alate throughout; pinnae commonly adnate, the costae alate to base (rarely nonalate just at base) g g. Tube of indusium broadly conical, 1-1.5 times as long as broad, mouth not or slightly flaring; tissue of most ultimate segments with elongate, narrow folds parallel to the veins 4. T. pyxidiferum g. Tube of indusium narrowly cylindrical; 2—4 times as long as broad, mouth abruptly and widely flaring; tissue of segments essentially plane (margin sometimes undulate) 5. T. diaphanum f. Rachis nonalate nearly to the apex; pinnae stalked, the costae not alate at base h h. Costae alate, except not below the basal pinnule; ultimate segments 0.5-0.8 mm broad; sori 1-5 per pinna; indusia commonly alate, the wings several cells wide 6. T. angustatum h. Costae nonalate except near the apex; ultimate segments 0.1-0.4 mm broad; sori rarely more than 1 per pinna; indusia not or scarcely alate, wings (if any) 1 cell wide 7. T. capillaceum b. Stem erect, or occasionally short-creeping; leaves caespitose to (occasionally) approximate [subg. Pachychaetum] i 78 FIELDIANA: BOTANY i. Tissue of segments opaque, the cells small and occluded, not evident when observed with magnification of 8-12 x; leaves (mature ones) 15-90 cm long; petiole (0.6-)0.7-3.5 mm in diameter j j. Rachis obtusely quadrangular, alate (often broadly so) at least in the distal half; pinnae adnate, the costae alate; lamina usually more than 1 cell thick, at least away from margins 8. T. elegans j. Rachis terete, nonalate, or marginate to slightly alate near apex; pinnae (at least proximal ones) short-stalked, costae not alate at base; lamina essentially 1 cell thick throughout . . . 9. T. rigidum i. Tissue of segments translucent, the cells large and clear, quite evident with magnification of 8-12 x; leaves commonly less than 15 cm long; petiole 0.3-0.6 mm in diameter 10. T. cellulosum a. Venation at least partly catadromous (or sometimes flabellate); false veins present or lacking . . . k k. False veins present, these submarginal or borne between and parallel with the true veins [subg. Didymoglossum] 1 1. Leaves with simple to stellate, commonly dark, trichomes along the margins; indusia distinctly bilabiate and the lips commonly dark-margined m m. Venation flabellate, a percurrent costa lacking 11. T. punctatum ssp. sphenoides m. Venation pinnate, or if subflabellate, the costa essentially percurrent n n. Mature leaves entire, or distally lobed; stellate trichomes borne all along the margin 1 2. T. angustifrons n. Mature leaves 1-2-pinnatifid; stellate trichomes confined to segment sinuses, with simple or bifid trichomes borne on outer margin o o. Son protruding from the leaf tissue, or slightly immersed at base; false veinlets extending toward the lamina margin, but rarely running parallel to it p p. False veinlets sparse, scattered; lips of indusium mostly broader than long; cells of lamina mostly isodiametric 13. T. hymenoides p. False veinlets commonly abundant; lips of indusium mostly as long as or longer than broad; cells of lamina mostly elongate 14. T. reptans o. Sori partly to deeply immersed in the leaf tissue, or at least broadly alate; false veinlets (many of them) parallel to and very near the margin 15. T. krausii 1. Leaves with margins glabrous or provided with circular scalelike processes (these sometimes deciduous in age); indusia not or scarcely bilabiate, the mouth not dark-margined q q. Lamina lacking paired, circular scalelike processes, but bearing a peripheral, submarginal false vein; venation essentially pinnate 16. T. kapplerianum q. Lamina bearing numerous paired, circular, scalelike processes all along the margin (these sometimes deciduous in age), but lacking a submarginal false vein; venation flabellate . . . 1 7. T. membranaceum k. False veins lacking or, in T. pinnatum, perpendicular to the true veins (some short, veinlike lines in sect. Lacostea); [subg. Achomanes] r r. Lamina trichomes stellate or forked from the base; stem filiform, 0.3-0.4 mm in diameter 18. T. polypodioides r. Lamina trichomes simple or lacking; stem relatively stout and wiry, 0.5-1.2 mm in diameter s s. Stem long-creeping up tree trunks, leaves subdistant to remote; lamina appressed to tree trunks by means of rhizoids on axes t t. Lamina linear, subentire to lobed, 1-1.5 cm broad; indusia mouth rather widely flaring 1 9. T. tanaicum t. Lamina oblong, pinnatisect or nearly pinnate, mature ones ( 1 .8-)2-9 cm broad; indusium mouth not or slightly flaring u u. Segments of sterile laminae entire or shallowly crenate; veins simple, not forked; indusium mouth not expanded; rare in Peru 20. T. tuerckheimii u. Segments of sterile laminae deeply crenate or crenate-serrate; veins (some or many) 1 -forked; indusium mouth slightly flaring; locally common 21. T. ankersii TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 79 s. Stem erect to decumbent and the leaves crowded to caespitose, or if in some species long- creeping and the leaves subdistant, then the lamina not appressed to tree trunks by rhizoids on leaf axes v v. Leaves dimorphic w w. Stem erect or decumbent; leaves crowded to caespitose; fertile leaves simple and essentially entire, or without laminar tissue and the sori completely free x x. Fertile leaves with lamina entire to pinnatisect; sori fully immersed in tissue . . y y. Sterile lamina with 4-8 veins issuing from the rachis between adjacent costae; mouths of indusia neither flaring nor recurved, strongly indented between the vein branches 23. T. diversifrons y. Sterile lamina with 2(-3) veins issuing from the rachis between adjacent costae; mouths of indusia slightly flaring and recurved, but not indented between the vein branches 24. T. trollii x. Fertile leaves without laminar tissue; sori completely free . . . 25. T. botryoides w. Stem wiry, long- or short-creeping; leaves subdistant to remote; fertile leaves with lamina foliose and deeply pinnatisect to nearly pinnate 28. T. humboldtii v. Leaves monomorphic z z. Lamina 3-4-pinnatisect; petiole conspicuously alate throughout, the alae broader than the petiole quite or nearly to base 22. T. bicorne z. Lamina not more than 1 -pinnate-pinnatisect; petiole nonalate in the proximal half or throughout, or very narrowly alate in the proximal portion aa aa. Lamina abruptly terminating in a conform apical segment or the rachis prolonged, flagellate, and proliferous at tip; indusia fully exserted from segment margin, often stalked bb bb. False veins copious and perpendicular to the true veins; marginal vein strong, continuous; pinna margins sharply serrate to spinulose . .26. T. pinnatum bb. False veins rare or lacking; marginal vein lacking or faint and discontinuous, pinna margins commonly obtusely serrate 27. T. hostmannianum aa. Lamina gradually reduced to a pinnatifid apex, the rachis not prolonged and flagellate; indusia at least partially immersed in the laminar tissue, wholly or partially alate on each side cc cc. Pinnae regularly and deeply lobed to pinnatisect dd dd. Pinnae lobed to pinnatifid; lamina l-2(-2.5) cm broad, not at all obscured by the moderate pubescence 29. T. crinitum dd. Pinnae pinnatisect (rarely the segments again lobed); lamina (2.5-)3-12 cm broad, partially to completely obscured by the dense covering of trichomes 31. T. lucens cc. Pinnae entire to serrate or crenate (or rarely a few with some irregularly scattered lobes) ee ee. Veins bearing crestlike lamellae on abaxial side; pinnae in proximal por- tion of lamina deflexed 30. T. martiusii ee. Veins lacking lamellae; pinnae patent or a few basal ones deflexed . . ff ff. Petiole conspicuously alate halfway or more to the stem; stem erect to decumbent, leaves contiguous to caespitose .... 32. T. pellucens ff. Petiole not alate, or scarcely so near apex due to decurrent lamina base; stem short-creeping or decumbent, the leaves subdistant to con- tiguous (rarely caespitose) gg gg. Rachis trichomes on abaxial side predominantly dark brown, stout, rigid, most of them terete toward their base . . 33. T. plumosum gg. Rachis trichomes tawny to orange, delicate, tortuous, the cells flattened hh hh. Lamina linear or linear-lanceolate, mature ones commonly 5-12 times as long as broad; mature leaves (10-) 15-50 cm 80 FIELDIANA: BOTANY long; rachis abaxially provided with trichomes predominantly unicellular beyond the basal one, occasionally with 2-3 cells 34. T. cristatum hh. Lamina oblong- to ovate-lanceolate, 3-4 times as long as broad; mature leaves 5-15 cm long; rachis abaxially provided with trichomes of 2-8 cells beyond basal one, as well as with many unicellular ones 35. T. vandenboschii 1. Trichomanes radicans Sw., J. Hot. (Schrader) 1800(2): 97. 1802. TYPE: Jamaica, Swartz (holotype, s; photo, us). Figure 12a-b. Trichomanes kunzeanum Hooker, Sp. fil. 1: 127. 1844. SYNTYPES: Peru, Huanuco, Pampayacu, Poep- pig 1132 (K!; photo, us); Peru, Junin, Pangoa, Mathews 1088 (K.!; photos. A, us); Venezuela, Merida, Linden 176 (K.!; isosyntypes, BM, n). Trichomanes brachyblastos Kuhn, Linnaea 35: 388. 1868. TYPE: Peru, San Martin, Tarapoto, Monte Guayrapurima, Spruce 4703 (holotype, B?; iso- types, BM!, BR!, GH!, K!, w!). Vandenboschia radicans (Sw.) Copel., Philipp. J. Sci. 67: 54. 1938. Trichomanes radicans var. kunzeanum (Hooker) Duek & Lell., Amer. Fern J. 68: 120. 1978. Stem long-creeping, relatively stout, 1-2.2 mm in diameter, amply to densely covered with cas- taneous trichomes. Leaves remote, 20-50 cm long, axes often provided with scattered septate tri- chomes abaxially. Petiole (l-)4-15 cm long, 0.8- 2 mm in diameter, nonalate to marginate or alate (often the wings deciduous). Lamina 2-3-pinna- tisect, ovate or deltoid-ovate and scarcely reduced at base, or occasionally elliptic and somewhat strongly reduced toward base. Rachis narrowly to broadly alate. Pinnae mostly 2-pinnatisect, slight- ly to strongly adnate to the rachis. Ultimate seg- ments commonly narrow, often linear, with simple or 1 -forked veins, false veinlets lacking. Venation anadromous. Indusia very slender, cylindrical or elliptic, rarely subconical, often subfusiform, not or scarcely alate, the wings of tissue (if any) 1- 2(-3) cells wide, apex truncate, not expanded, re- ceptacle short- or long-exserted. Hemiepiphytic on tree trunks in dense, wet for- ests, (400-)700-3200 m, San Martin to Puno. Mexico to Panama; West Indies; Guianas to Co- lombia and southward to Brazil and Paraguay; Old World. Trichomanes radicans, T. collariatum, and T. rupestre form a close-knit species complex which is in need of closer study. The three are only pro- visionally maintained here at the species level. In Central America T. radicans and T. collariatum are more distinct, e.g., leaves of the latter are con- sistently reduced to a very short and broadly alate petiole; whereas in T. radicans leaves are not or scarcely reduced at base, and petioles are much longer and not or very narrowly alate or marginate. These characters are also somewhat diagnostic in South America, but are much more variable, and are therefore not thoroughly reliable. In Peru, often the two species can be positively separated only by the soral features. San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, J. Schunke 4361 (F, us). Huanuco: Near Tingo Maria, confluence of Monzon and Huallaga Rivers, Stork & Norton 9501 (F, GH, MO, uc, us). Pasco: Paujil near Puerto Bermudez, Leon 288 (GH, USM). Junin: Chan- chamayo Valley, C. Schunke 459 (F, us). Ucayali: Prov. Coronel Portillo, Sinchono, between Tingo Maria and Pucallpa, Aguilar 883 (GH). Ayacucho: Prov. La Mar, between Tambo San Miguel, Ayna and Hacienda Lu- isiana, Dudley 11904 (GH). Cuzco: Prov. Paucartambo, bosques de Keros, Vargas 7382 (uc). Puno: Prov. Car- abaya, Hacienda Palmera, Vargas 16137 (GH). 2. Trichomanes collariatum Bosch, Ned. Kruidk. Arch 4: 368. 1859. TYPE: Mexico, Tabasco, Linden (holotype, L?; probable isotype, K; photo, us of K). Stem long-creeping, relatively stout, 1-1.7 mm in diameter, amply to densely covered with cas- taneous trichomes. Leaves remote, 1 5-38(-45) cm long, axes often provided with scattered, septate trichomes abaxially. Petiole 0.3-3 cm long, 0.7- 1.5 mm in diameter, broadly alate nearly or quite throughout. Lamina 2-3-pinnatisect, oblong or oblong-lanceolate, often abruptly reduced at base. Rachis commonly broad-alate throughout. Pinnae mostly 2-pinnatisect, slightly to strongly adnate to TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 81 rachis. Ultimate segments commonly narrow, often linear, with simple or forked veins, false veinlets lacking. Venation anadromous. Indusia conical, gradually widening from base to a conspicuously flaring mouth, alate, the wings of tissue on each side 4-6 cells wide, receptacle long-exserted. Hemiepiphyte on tree trunks in wet forests, sea level to 500 m, San Martin, Loreto, Madre de Dios. Southern Mexico to Panama; British Guiana to Colombia, south to Peru; Brazil. This is not greatly distinct from T. radicans, under which see further discussion. San Martin: Prov. Mariscal Caceres, Dist. Campan- illa, left margin of Rio Huallaga, J. Schunke 4120 (F, GH, us). Loreto: Soledad, on Rio Ilaya, Killip & Smith 29739 (us). Madre de Dios: Prov. Tambopata, Lago Tres Chim- badas, Barbour 5669 (MO). Parque Nacional del Manu, Cocha Cashu Biological Station, M. S. Foster P-84-4 (uc), P-84-67 (MO); R. Foster 6868 (F). 3. Trichomanes rupestre (Raddi) Bosch, Ned. Kruidk. Arch. 4: 370. 1859. Hymenophyllum rupestre Raddi, PI. Bras. nov. gen. 1 : 67, /. 80. 1 825. TYPE: Brazil, Raddi (holotype, FI?; isotype, K.; photos, A & us of K). Stem long-creeping, stout, 0.7-1.2 mm in di- ameter, provided with a few scattered to sparse castaneous trichomes. Leaves remote, mature ones 15-30 cm long, axes sometimes provided with scattered, septate trichomes on abaxial side. Pet- iole 0.2-2 cm long, 0.6-0.8 mm in diameter, par- tially alate. Lamina commonly 1-pinnate-pinna- tifid, oblong or elliptic-oblong. Rachis broadly alate throughout. Pinnae pinnatifid or pinnatisect, or rarely 2-pinnatifid as to basal segments, strongly adnate to rachis. Ultimate segments relatively short and broad, not linear, with simple or forked veins, false veinlets lacking. Venation anadromous. In- dusia conical, gradually widening from base to a slightly flaring mouth, narrowly alate on each side, receptacle short- to long-exserted. Creeping on tree trunks or wet banks in dense, wet forests, 1 100-1600 m, Huanuco and Ucayali. Venezuela and Colombia; Peru; Bolivia; Brazil (also reported from Costa Rica by Alan Smith, in. litt). This is not as common as Trichomanes radicans or T. collariatum and is midway between them in shape of indusia. The indusium in the latter is conical, with a conspicuous and widely flaring mouth; in T. rupestre it is narrower, with mouth only slightly flaring, and in T. radicans the indu- sium is even more slender, often narrowing slight- ly, with the apex truncate, not or scarcely spread- ing. See T. radicans for further discussion. Huanuco: Prov. Huanuco, Dist. Churubamba, Cotir- arda, Mexia 8221 (F, GH, MO, uc, us). Tingo Maria, Selva Real, Morrow 11132 (GH). Ucayali: Prov. Coronel Por- tillo (as Loreto), Aguaytia, Ridoutt 13082 (USM). 4. Trichomanes pyxidiferum L., Sp.pl. 1098. 1753. TYPE: based on Plumier, Traite foug. Amer., /. 50E, 1705, illustrating a plant from Haiti. Vandenboschia pyxidifera (L.) Copel., Philipp. J. Sci. 67: 53. 1938. Stem long-creeping, filiform, 0.4-0.5 mm in di- ameter (excluding indument), densely covered with blackish trichomes. Leaves well-spaced to remote, 3-12 cm long, axes sometimes provided abaxially with scattered, minute trichomes. Petiole 1-3 cm long, 0.3-0.6 mm in diameter, flattened, or terete at base, narrowly to broadly alate distally or throughout, the wings plane to somewhat crispate. Lamina 2-3-pinnatifid. Rachis alate throughout, the wings commonly undulate to crispate. Pinnae 4-10 pairs, strongly adnate to the rachis (the costae alate). Ultimate segments linear, margins plane to undulate, tissue commonly with elongate, narrow folds parallel to the veins, false veinlets lacking. Venation anadromous. Sori 1 -several per pinna. Indusia broadly conical, the tube 1-1.5 times as long as broad, not or scarcely bilabiate, the mouth not or slightly flaring, receptacle commonly long- exserted. On trees in wet forests, 900-1 200 m, Amazonas, San Martin. Pantropical. A distinctive feature of Trichomanes pyxidifer- um is the rather sharp, longitudinal folding of seg- ment tissue parallel to the veins. Other related species have plane segments, or some with the margins undulate, but none exhibit this unique character. Perhaps it was this which prompted Morton's incorrect observation (1968) of false veins in the species, since the sharper folds of tissue sometimes cause dark shadow lines in the seg- ments. However, in careful study of specimens throughout the range, no leaves were seen with the 82 FIELDIANA: BOTANY sclerenchymatous strands which produce these "false veins." The species thus far is represented in Peru by only two collections, even though its full range extends throughout tropical regions of the world. Amazonas: Prov. Chachapoyas, .la/an (Ingenio- Chachapoyas), Ldpe: el al. 4264 (GH, HUT). San Martin: Tarapoto, Spruce 4761 (BR, OH, K, L, us). 5. Trichomanes diaphanum HBK., Nov. gen. sp. 1:25 (fol. 1 6). 1 8 1 6. TYPE: Venezuela, Hum- boldt & Bonpland (holotype, p; isotype, B!). Trichomanes leptophyllum Bosch, Ned. Kruidk. Arch. 4: 363. 1859 (not A. Cunn., 1836), nom. illeg. TYPE: based on T. pyxidiferum Hooker & Grev. Trichomanes hymenophylloides Bosch, Ned. Kruidk. Arch. 5: 209. 1863, nom. nov. for T. leptophyllum Bosch and type based on that name. Vandenboschia diaphana (HBK.) Copel., Philipp. J. Sci. 67: 53. 1938. Vandenboschia hymenophylloides (Bosch) Copel., Philipp. J. Sci. 67: 53/1938. Stem long-creeping, filiform, 0.2-0.5 mm in di- ameter (excluding indument), sparsely to abun- dantly provided with castaneous trichomes. Leaves well-spaced to remote, 5-15(-18) cm long, axes glabrous, or the rachis sometimes abaxially pro- vided with scattered, minute trichomes. Petiole 0.5-6 cm long, 0.3-0.8 mm in diameter, flattened, or terete at base, nonalate, or narrowly to broadly alate for most of its length, the wings plane or occasionally undulate. Lamina essentially 3-pin- natifid. Rachis alate throughout, the wings narrow to broad, plane to undulate or occasionally cris- pate. Pinnae 4-12 pairs, adnate to the rachis, the costae alate (rarely nonalate at base). Ultimate seg- ments linear, 0.6-0.9 mm broad, plane, not folded, false veinlets lacking. Venation anadromous. Sori 1-10 per pinna. Indusia salverform, the alate tube narrowly cylindrical, 2-4 times as long as broad, abruptly expanding into a broad-flaring mouth, receptacle commonly long-exserted. On trees, moist banks and (rarely) wet rocks, in deep forests 700-2900 m, Amazonas and Loreto to Puno. Southern Mexico to Panama; West Indies; Trin- idad; the Guianas to Colombia, south to Peru and Brazil. Previous authors have separated Trichomanes hymenophylloides on characters such as number of sori per pinna, width of tissue wings flanking axes and indusia, crowding of pinnae and dissec- tion of lamina. Each of these may appear to be important in individual specimens, but when many collections are examined throughout the range of distribution, it becomes evident that they are quite variable and uncorrelated. Amazonas: Prov. Bagua, Cordillera Colan SE of La Peca, Harbour 3 904 (MO). I .ore to: Sierra del Pongo, Mex- ia 6288B (GH, uc, us). Pasco: Prov. Oxapampa, Dist. Oxapampa, road Oxapampa to Villa Rica, B. Ledn 660 (USM in part). Junin: La Merced, E of Quimiri Bridge, Killip & Smith 24006 (F), 24023 (GH, us). Cuzco: Prov. La Convencion, "El Dorado," Vargas 3518 (GH). Puno: San Gaban, Lechler (L). 6. Trichomanes angustatum Carm., Trans. Linn. Soc. London 12: 513. 1818. TYPE: Tristan da Cunha, Carmichael (holotype, K!; isotype, BM!). Trichomanes tenerum Sprengel, Syst. veg. 4: 1 29. 1 827. TYPE: Brazil, collector not stated (holotype, LZ destroyed). Vandenboschia tenera (Sprengel) Copel., Philipp. J. Sci. 67: 53. 1941. Vandenboschia angustata (Carm.) Copel., Philipp. J. Sci. 73: 466. 1941. Stem long-creeping, filiform, 0.2-0.3 mm in di- ameter, amply provided with castaneous tri- chomes. Leaves well-spaced to remote, 4-14(-18) cm long, glabrous. Petiole 0.5-5 cm long, 0.2-0.4 mm in diameter, terete, nonalate. Lamina 2-pin- nate-pinnatifid to 4-pinnate, linear-lanceolate to ovate-lanceolate, abruptly or gradually reduced at base. Rachis nonalate, or scarcely alate near the apex. Pinnae 6-many pairs, short-stalked, the cos- tae alate except below the basal pinnule. Ultimate segments linear, 0.5-0.8 mm broad, each bearing a single vein, false veinlets lacking. Venation anad- romous. Sori 1-5 per pinna. Indusia narrowly fun- nelform to salverform, the tube 2-4 times as long as broad, the mouth flaring, margins commonly alate, with wings several cells wide, receptacle long- exserted. In deep forests, on trees, wet cliffs and banks, and on decaying logs, 600-2800 m, Cajamarca and San Martin to Ayacucho and Cuzco. Southern Mexico to Honduras; Greater Antilles; Venezuela and Colombia to NW Argentina and Brazil. This is very closely related to Trichomanes cap- illaceum, under which see further discussion. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 83 Cajamarca: Colasay, Woytkowski 7002 (us). San Mar- tin: Monte Campana, near Tarapoto, Spruce 4705 (BR, K, NY). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8428 (MO, uc). Junin: Carpapata, above Huacapistana, Killip & Smith 24492 (F, uc, us). Ayacucho: Ccarrapa, between Huanta and Rio Apurimac, Killip & Smith 22406 (F, GH, us). Cuzco: Valley of Urubamba, Machu-Picchu, base of Huayna Picchu, Iltis el al. 1064 (GH, MO, uc, us). 7. Trichomanescapillaceum L., Sp. pi. 1099. 1753. LECTOTYPE (designated by Proctor, Ferns of Jamaica, British Museum, London, p. 109. 1985): Plumier, Traite foug. Amer., t. 99D, based on a Plumier collection from Haiti. Vandenboschia capillacea (L.) Copel., Philipp. J. Sci. 67: 53. 1938. This differs from Trichomanes angustatum only in the characters used in the key. In deep forests, on tree trunks, 800-1100 m, Pasco and Cuzco. Southern Mexico to Panama; Greater Antilles; Venezuela and Colombia to Peru. With this might possibly be included Trichom- anes angustatum, but the two are separated here provisionally, pending much needed monographic revision. Typically, T. capillaceum has very little expanded tissue along the axes, and even the ul- timate segments are rarely more than 0.3 mm broad. Indusia are not or scarcely alate, and son are solitary on each fertile pinna. Conversely, be- yond the first pinnule in T. angustatum, the costae and costules are obviously alate throughout, and ultimate segments are often nearly 1 mm broad. The tissue also forms wings on each side of the indusia, and sori are frequently four or five per pinna. Even some of the characters listed in the key are subject to variation in certain geographic regions, and intermediates are found frequently. Never- theless there are areas in which only one or the other occur. Only Trichomanes angustatum oc- curs in Bolivia and southern Brazil, whereas in Central America it has not been found south of Guatemala, a region where T. capillaceum is rath- er common. In Peru, the latter is rare, and even the few specimens located are not typically skeletal in their appearance, whereas Trichomanes angus- tatum is rather widespread and the specimens rather typical. One specimen from Cuzco, Vargas 8653 (MO), is exactly intermediate between the two; significantly, Cuzco is one of the two de- partments in which both species are known to oc- cur. Therefore, based only on collections from Peru, one would be tempted to combine the two species, but a thorough study of the characters of both taxa throughout their range is necessary before further classification is attempted. Pasco: Pichis Trail, San Nicolas (as Junin), Killip & Smith 25990 (F, GH, us). Cuzco: Prov. Paucartambo, Cosnipata Valley, Rio Tono, Wachter et al. 199 (F). Trichomanes elegans Rich., Actes Soc. Hist. Nat. Paris 1: 114. 1792. TYPE: French Guiana, Le Blond (P) (not T. elegans Rudge, 1805). Trichomanes prieurii Kunze, Analecta Pteridogr. 48. 1837. SYNTYPES: French Guiana, Leprieur (LZ destroyed; isosyntype, F!); Brazil, Amazonas, Rio Japura, Martius (M); British Guiana, Rudge (K). Trichomanes opacum Bosch, Ned. Kruidk. Arch. 5(2): 1 75. 1 86 1 . TYPE: Peru, Puno, San Gaban, Lech- ler 2175 (holotype, L?; isotype, P!; photo, F, frag., L!). Davalliopsis elegans (Rich.) Copel., Philipp. J. Sci. 67: 82. 1938. Terrestrial, rarely epiphytic or epipetric. Stem erect or rarely short-creeping. Leaves caespitose to occasionally approximate, 20-90 cm long, rachis, costae, and often costules amply to abundantly provided with castaneous, pluricellular trichomes on abaxial side. Petiole 10-40 cm long, 1-3.5 mm in diameter, usually alate distally. Lamina 2-3- pinnate-pinnatifid, deltoid, opaque, usually more than 1 cell thick, at least away from the margins, the cells small and occluded, not evident even when magnified at 10-15x. Rachis obtusely quadran- gular, alate (often broadly so) at least in distal half. Pinnae adnate, the costae alate to base. Ultimate segments laciniate, with linear lobes, each of these bearing a single vein, false veins lacking. Venation anadromous. Sori usually bent down away from the plane of the lamina. Indusia subconical to somewhat urceolate, not bilabiate, the mouth truncate or slightly (rarely strongly) flaring, recep- tacle short- to long-exserted. In dense, wet forests or wet, wooded ravines, on ground, or rarely on tree trunks or wet rocks, sea level to 1300 m, Amazonas and Loreto south to Madre de Dios and Puno. Lesser Antilles and Trinidad; Nicaragua to Pan- ama; Guianas to Colombia and south to Bolivia and Brazil. 84 FIELDIANA: BOTANY With its large, blue green leaves and laciniate pinnae, this is one of the most beautiful species in the family. Its most distinctive features are the usually strongly arching sori (bent downward out of the plane of the lamina) and the relatively thick tissue. Although the lamina in Hymenophyllaceae is typically only one cell thick, the tissue in Tri- chomanes elegans is often several cells thick, es- pecially away from the segment margins. The broadly alate rachis traditionally has been used to separate this from T. rigidum (with which it often grows), and although usually it is a good diagnostic feature, occasional specimens of T. elegans can be found throughout the range in which rachis alae are very narrow or lacking in the proximal portion. Consequently, supporting key characters should be employed for accurate identification. Amazonas: Prov. Bagua, left bank of Rio Maranon opposite Quebrada Mirana, Wurdack 2031 (F, GH, us). San Martin: Prov. Mariscal Caceres, Tocache Nuevo, J. Schunke4855 (F, GH, us), 6955 (F, MO in part, us). Loreto: Sierra del Pongo, Mexia 1 195 (F, GH, MO, us). Huanuco: Prov. Pachitea, Dist. Honoria, /. Schunke 2259 (F, GH). Pasco: Prov. Oxapampa, Valle del Palcazu, Iscozacin, Le6n 692 (F, GH). Junin: Puerto Bermudez, Killip & Smith 26537 (GH, us). Ucayali: Prov. Coronel Portillo (as Lor- eto), Padre Abad, J. Schunke 5393 (F, us). Cuzco: Bajada de Tocate, Biies 1748 (us). Madre de Dios: Prov. Tam- bopata, Rio Tambopata Nature Reserve, Barbour 5165 (MO). Puno: Prov. Carabaya, San Gaban, Vargas 18864 (GH). 9. Trichomanes rigidum Swartz, Prodr. 137. 1788. TYPE: Jamaica, Swartz (holotype, s; isotype, B, Herb. Willd. 20202- 7; photos, us of s, GH & us of B). Selenodesmium rigidum (Sw.) Copel., Philipp. J. Sci. 67:81. 1938. Terrestrial, rarely epiphytic or epipetric. Stem erect or occasionally short-creeping. Leaves caes- pitose to sometimes approximate, 1 5-35 cm long, lamina glabrous, the axes glabrous, or rarely the rachis or petiole base with a few, scattered tri- chomes. Petiole 5-16 cm long, 0.6-1.6 mm in di- ameter, nonalate. Lamina 3-4-pinnate-pinnatifid, deltoid, opaque, 1 cell thick, but cells small and occluded, not evident even when magnified at 1 0- 1 5 x . Rachis terete, nonalate, or marginate to slightly alate toward apex. Pinnae (at least prox- imal ones) short-stalked and the costae not alate at base. Ultimate segments with linear lobes, each of these bearing a single vein, false veins lacking. Venation anadromous. Sori (most of them) borne essentially in the plane of the lamina. Indusia sub- conical, not bilabiate, the mouth truncate, not or scarcely flaring, receptacle commonly long-exsert- ed. On slopes and ridges of dense forests and in wooded ravines, occasionally on wet rocks or cliffs, rarely on bases of tree trunks, 350-2500 m, Ca- jamarca and San Martin, south to Cuzco and Madre de Dios. Southern Mexico to Panama; West Indies; South America, to Bolivia and Brazil; probably Old World tropics. This is sometimes confused with Trichomanes elegans, with which it often grows. For compari- son, see discussion under the latter species. Cajamarca: Tambillo, Jelski 891 (us). San Martin: Tarapoto, in monte Guayrapurima, Spruce 4047 (BR, K). Huanuco: East of Tingo Maria (as San Martin), Allard 22356 (GH, us). Pasco: Oxapampa, Cordillera San Ma- tias, Leon 326a (USM). Junin: La Merced, Hacienda Schunke, Macbride 5633 (F, us). Ucayali: Vicinity of Aguaytia, along Rio Aguaytia (as Loreto), Croat 20927 (MO). Cuzco: Prov. Urubamba, base of Huayna Picchu, Iltis et al. 1062 (GH). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster 10903 (F). 10. Trichomanes cellulosum Klotzsch, Linnaea 18: 531. 1844. TYPE: "British Guiana," (Guy- ana). Kanuku Mountains, Rich. Schomburgk 1186 (holotype, B!; photo, F; isotypes, B! 2 sheets, BM!). Epiphytic, terrestrial, or sometimes epipetric. Stem erect to short-creeping. Leaves caespitose to (occasionally) approximate, 6-15 cm long, gla- brous. Petiole 1-7 cm long, 0.3-0.6 mm in di- ameter, narrowly alate at least distally. Lamina 4- 5-pinnate, lanceolate to ovate, translucent, the cells large and clear and quite evident when magnified at 8-12x. Rachis terete, alate throughout. Pinna divisions skeleton-like, the costae, costules, and ultimate segments with bands of tissue only 1- several rows wide. Ultimate segments linear, 0.3- 0.7 mm broad, each bearing a single vein, false veins lacking. Venation anadromous. Indusia sub- conical to somewhat urceolate, not bilabiate, the mouth truncate or slightly flaring, receptacle short- to long-exserted. In wet forests, on trees or on the forest floor, sometimes on wet rocks, 300-1 300 m, Amazonas, Loreto, Huanuco. Surinam to Colombia; Peru; Brazil. With this perhaps should be included Trichom- TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 85 anes sprucei Baker of northern South America, which is probably only a more robust form, with broader segments and broader bands of tissue along the axes. Amazonas: Prov. Bagua. roadside from Chiriaco to Puente Venezuela, Barbour 4455 (MO, USM). Loreto: Pu- mayacu, between Balsapuerto and Moyobamba, Klug 3192 (F, GH, MO, us). Tierra Doble on Rio Nanay, LI. Williams 1057 (F, us). Huanuco: Rio Llullapichis wa- tershed, ascent of Cerros del Sira, Dudley 13032 (GH), Wolfe 12352 (GH, us). 1 1 . Trichomanes punctatum Poiret ssp. sphen- oides (Kunze) Boer, Acta Bot. Neerl. 1 1: 301. 1962. Trichomanes sphenoides Kunze, Farrnkrauter 216, /. 88, f. 2. \ 840. TYPE: Peru, Cuchero (Dept. Huan- uco), Poeppig (holotype, w!; isotype, MO!). Didymoglossum sphenoides (Kunze) Presl, Hymeno- phyllaceae 23. 1843. Stem long-creeping, filiform. Leaves approxi- mate or widely spaced, to 1.5 cm long and 1 cm broad, subsessile or short-petiolate. Lamina cir- cular to obovate or spathulate, margin entire to irregularly lobed, a costa lacking or, in some fertile laminae, distinct only in the proximal portion, gla- brous except for the dark, stellate, marginal tri- chomes. Venation flabellate, the veins crowded, with false veinlets parallel to them. Sori several, or sometimes solitary, partially immersed in the lamina tissue, commonly borne between lobes, thus rarely extending beyond the outline circumscribed by the lobe apices. Indusia narrow-cylindrical, bil- abiate, the lips mostly wide-flaring and narrowly dark-margined, receptacle not or scarcely exsert- ed. In dense, wet forests and wooded ravines, on tree trunks, 100-1 100 m, Loreto to Pasco. Guatemala; Costa Rica; Panama; Greater An- tilles; Venezuela; Colombia to Bolivia. Boer (1962) recognized four subspecies of Tri- chomanes punctatum, and only this one occurs in Peru. Others differ from ssp. sphenoides in having more distinctly lobed laminae and/or larger and more broadly dark-margined indusium lips. Loreto: Maynas. Iquitos, Ushpa-Cana across from Rio Itaya, McDaniel 11381 (GH, MO). Huanuco: Prov. Huan- uco, near confluence of Rio Cayumba with Huallaga, Mexia 8275 (F, MO, us). Pasco: Prov. Oxapampa, Valle de Palcazu, Cacazu, Leon 681 (F). Puerto Bermudez (as Junin), Killip & Smith 26592 (us). 1 2. Trichomanes angustifrons (Fee) Boer, Fl. Neth. Antill. I (Pterid.): 17. 1962. Didymoglossum angustifrons Fee, Mem. foug. 11: 113, t. 28, f. 5. 1866. TYPE: Guadeloupe, I'Herminier (holotype, P!). Stem long-creeping, filiform. Leaves approxi- mate or well-spaced, 0.5-1.5 cm long, to 0.6 cm broad, subsessile or short-petiolate. Lamina nar- rowly or broadly oblong, entire, or distally lobed, costa percurrent, glabrous except for the dark, stel- late trichomes borne along the margin. Venation catadromous, pinnate, with false veinlets sparsely to amply distributed between the true veins. Sori solitary at lamina apex, or few and terminating each distal lobe, somewhat to deeply immersed in the tissue. Indusia narrow-conical, bilabiate, the lips mostly wide-flaring and narrowly dark-mar- gined, receptacle commonly exserted. Thus far represented in Peru by a single speci- men from Huanuco, on ridge in jungle, 850 m; elsewhere apparently always epiphytic. Costa Rica and Cocos Island; West Indies; Trin- idad and Tobago; the Guianas to Brazil; Peru; Paraguay (reported from Chiapas, Mexico by Alan Smith, in litt.). Collections of this species are rather sparse, ex- cept from Brazil and the West Indies; yet it is surely a much more common plant than the spec- imens would indicate. Since it is such a tiny and inconspicuous fern, it certainly has been over- looked and probably is not accurately represented in herbaria. Huanuco (as San Martin): E of Tingo Maria, Allard 21383a (us). 1 3. Trichomanes hymenoides Hedwig, Fil. gen. sp., /. 3,f. 3. 1799. LECTOTYPE (designated by Boer, Acta Bot. Neerl. 11: 306. 1962): Hed- wig, t. 3, f. 3, probably based on a Swartz specimen from Jamaica. Figure 12d. Trichomanes muscoides Sw., J. Bot. (Schrader) 1 800(2): 95. 1802. TYPE: Jamaica, Swartz (holotype, S-PA). Didymoglossum hymenoides (Hedwig) Desv., Mem. Soc. Linn. Paris 6: 330. 1827. Didymoglossum muscoides (Sw.) Desv. Mem. Soc. Linn. Paris 6: 330. 1827. Stem long-creeping, densely covered with dark trichomes that extend onto the petioles. Leaves 86 FIELDIANA: BOTANY well-spaced, (0.5-)l-3.5 cm long, to 1.5 cm broad, subsessile or short-petiolate. Lamina obovate to elliptic, 1-pinnatifid or rarely 2-pinnatifid, the cos- ta percurrent (but juvenile laminae often subren- iform and merely lobed and the costa indistinct distally), glabrous except for the dark, marginal trichomes which are stellate in sinuses and simple to bifid on outer margin, laminar cells commonly isodiametric. Venation catadromous, commonly pinnate (occasionally subflabellate in juvenile leaves), false veinlets sparse, scattered between true veins, extending toward lamina margin but not parallel to it. Sori 1-several borne near lamina apex, conspicuously exserted, or only slightly im- mersed at base, not or very narrowly alate. Indusia narrow-funnelform, bilabiate, but the lips quite short, commonly broader than long, and usually (not always) dark-margined, receptacle long- exserted. In forests on trees, or rarely on rocky or sandy soil, 700-2400 m, Amazonas, Loreto. Southern Mexico to Panama; West Indies; Trin- idad; Venezuela and Colombia, south to Argentina and Uruguay. Juvenile leaves are often nearly reniform and shallowly lobed, with veins subflabellate and the costae sometimes indistinct toward the apex. Hence they can be confused with Trichomanes puncta- tum. However, fertile leaves are pinnatifid, with stellate trichomes borne only in the sinuses, and son strongly protrude from the leaf tissue. In T. punctatum ssp. sphenoides sori are partially im- mersed, and marginal trichomes are consistently stellate. Amazonas: Prov. Chachapoyas, Rio Ventilla W of Molinopampa, Wurdack 1513 (F, OH, us). Loreto: Sierra del Pongo, Mexia 6289B (GH, us). 14. Trichomanes reptans Sw., Prodr. 136. 1788. TYPE: Jamaica, Swartz (holotype, S-PA). Didymoglossum reptans (Sw.) Presl, Hymenophylla- ceae 23. 1843. Stem long-creeping, densely covered with dark trichomes that extend onto the petioles. Leaves well-spaced, 2-9 cm long, 1—4 cm broad, subsessile to short-petiolate. Lamina lanceolate to ovate or elliptic, 1-pinnatifid or rarely 2-pinnatifid, the rachis distinct throughout, glabrous except for the dark, marginal trichomes which are stellate in si- nuses and simple to bifid on outer margins, lam- inar cells mostly elongate. Venation catadromous, pinnate, false veinlets commonly abundant, ex- tending toward lamina margin but not (or very rarely) parallel to it. Sori 1 or a few, borne near lamina apex, conspicuously protruding from the leaf tissue, or only slightly immersed at base, not or scarcely alate. Indusia narrow-funnelform, bil- abiate, the lips dark-margined, and sometimes flaring, most of them as long as or longer than broad, receptacle short- to long-exserted. On wet rocks in forests, 1900-2750 m, Ama- zonas, Huanuco. Southern Mexico to Panama; Greater Antilles; Venezuela; Colombia; Ecuador; Peru; Brazil; Ar- gentina. Although found thus far in Peru only on wet rocks, Trichomanes reptans may be expected also on trees, since it frequently has been reported as an epiphyte throughout much of its range. Amazonas: Prov. Bagua, Cordillera Colan SE of La Peca, Harbour 3903 (MO, USM). Huanuco: Mito, Bryan 392 (F). 15. Trichomanes krausii Hooker & Grev., Icon, fil. 2, /. 149. 1831. TYPE: Dominica, Kraus (holotype, E). Didymoglossum krausii (Hooker & Grev.) Presl, Hy- menophyllaceae 23. 1843. Stem long-creeping, densely covered with dark trichomes which extend onto the petioles. Leaves well-spaced, 2-8 cm long, 1-3 cm broad, subsessile to short-petiolate. Lamina narrow-ovate to -ellip- tic or obovate, 1 -pinnatisect to 2-pinnatifid, the rachis distinct throughout and often abaxially dark- pubescent at least near the base, the lamina pro- vided with simple to bifid trichomes, but with stellate ones in the segment sinuses. Venation ca- tadromous, pinnate, false veinlets seldom numer- ous, many of them parallel to and very near the lamina margin. Sori 1 or a few, terminating the segments, partly to deeply immersed in the seg- ment tissue, the protruding portion narrowly or broadly alate. Indusia salverform, bilabiate, the lips commonly dark-margined and flaring, recep- tacle short- to long-exserted. Found near 800 m, San Martin, Huanuco. There are no other label data on the only two collections TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 87 found thus far in Peru, but Trichomanes krausii occurs in forests on trees or wet rocks elsewhere throughout its range. United States (Florida); Mexico to Panama; West Indies; Trinidad; the Guianas to Colombia, south to Paraguay and Argentina. Besides the characters used in the key, Trichom- anes krausii often can be distinguished from closely related species by the dark trichomes that fre- quently extend up the petiole onto the abaxial side of the lower rachis. Rachises of the other species are essentially glabrous. San Martin: Tarapoto, Spruce 3991 (BR, w). Huanuco: Fundo Chela, Sinchono, Rio Chino, Aguilar 912 (USM). 16. Trichomanes kapplerianum J. W. Sturm, Fl. bras. 1(2): 276. 1859. TYPE: Surinam, near Station Victoria, Kappler 1760 (holotype, w!). Hemiphlebium kapplerianum (J. W. Sturm) Prantl, Hymenophyllaceae 46. 1875. Trichomanes ekmanii Boer, Acta Bot. Neerl. 11: 319. 1962. TYPE: Dominican Republic, La Cumbre, Cordillera Central, Ekman H- 14342 (holotype, u; isotypes, F!, G, GH!, MO!, s, uc, us!). Stem long-creeping, filiform, sparsely to abun- dantly provided with brown trichomes which ex- tend onto the petioles. Leaves well-spaced, to 3 cm long and 1.5 cm broad, subsessile or short - petiolate. Lamina obovate or oblong (or in juve- nile ones often circular), base rounded, cordate or cuneate, entire or distally lobed, with a distinct costa at least in the proximal portion, glabrous, or sparsely pubescent basally along the costa. Vena- tion catadromous, essentially pinnate (but some- times subflabellate distally), false veins ample to abundant, parallel to the true veins and with a submarginal, continuous or sometimes interrupt- ed false vein around the perimeter of the lamina. Sori several to ample near lamina apex or on the lobes, fully immersed in the tissue. Indusia fun- nelform, not bilabiate, the mouth expanded, but not dark-margined, receptacle short- or long- exserted. In dense forests and wooded ravines, on tree trunks (also on rocks, outside Peru), sea level to 1000 m, San Martin, Loreto, Huanuco, Puno. Guatemala to Panama; West Indies; the Guian- as to Venezuela; Peru; Brazil; Bolivia. Detailed study of many specimens throughout the Neotropics indicates that Trichomanes ek- manii is not distinct. The latter was said by Boer (1962) to differ in the uninterrupted submarginal false vein, bordered on the outside by cubical cells, whereas the submarginal vein of T. kapplerianum was said to be sometimes interrupted and bor- dered by tangentially lengthened cells. These fea- tures prove to be minor ones that are quite vari- able. In careful study of many specimens, including types, one can nearly always find some interrup- tions in submarginal false veins; furthermore, shape of the border cells is seldom constant. Many leaves on the type specimen of T. kapplerianum, for ex- ample, have nearly as many cubical cells as tan- gentially lengthened ones, and some cells are even radially lengthened (supposedly a distinguishing feature of T. hookeri Presl, of the Greater Antilles). Additionally, most of the type specimens of T. ekmanii have at least a few leaves with somewhat interrupted submarginal veins. There is probably no justification for separating either species from T. hookeri, which Boer distinguished on similar characters. However, the type of the latter has not been examined and T. kapplerianum is provision- ally maintained here as distinct. San Martin: Near Tarapoto, Spruce 4762 (BR, w). Lor- eto: San Antonio, on Rio Ataya, Killip & Smith 29522 (F, GH). Prov. Maynas, on the Amazon, 50 miles down- river from Iquitos, Moran 3661 (F). Huanuco: E of Tingo Maria (as San Martin), Allard 20912 (us). Puno: Near San "Gavan" (Gaban), Lechler 2297 (BR). 17. Trichomanes membranaceum L., Sp. pi. 1097. 1753. TYPE: "America," without specific lo- cality (holotype, LINN 1253.1). Lecanium membranaceum (L.) Presl, Hymenophyl- laceae 12. 1843. Didymoglossum membranaceum (L.) Vareschi, Flora Venezuela 1: 222. 1969. Stem long-creeping, slender, densely covered with dark brown trichomes which extend onto the short petioles. Leaves well-spaced, 2-6 cm long and often nearly as broad, subsessile. Lamina near- ly circular and entire, or spathulate and incised into regular lobes, lacking a distinct midrib, gla- brous, but bearing along the margin numerous, paired, circular, scalelike processes (these some- times deciduous in age). Venation flabellate, the veins repeatedly dichotomous, false veinlets abun- dant and parallel with the true veins, but a con- 88 FIELDIANA: BOTANY tinuous submarginal false vein lacking. Sori sev- eral to many on vein tips toward the lamina apex, partially to fully immersed in the tissue. Indusia narrow-funnelform, not or scarcely bilabiate, the mouth neither flaring nor dark-margined, recep- tacle short- or long-exserted. In deep forests or wooded ravines, on tree trunks or wet rocks or cliffs, sea level to 850 m, Ama- zonas, Loreto, Huanuco. Southern Mexico to Panama; West Indies; the Guianas to Colombia, south to Bolivia. Amazonas: Prov. Bagua, Rio Maranon opposite Que- brada Mirana, Wurdack 2040 (F, us). Loreto: Dist. Tigre, Rio Tigre near Tigre, McDaniel & Rimachi 18531 (GH). Pongo de Manseriche, Mexia 6198 (F, OH, MO, us). Huanuco: E of Tingo Maria (as San Martin), Allard 21549 (GH, us). 18. Trichomanes polypodioides L., Sp. pi. 1098. 1753. NEOTYPE (designated by Proctor, Flora Lesser Antilles 92. 1977): Montserrat, Proctor 19068 (A). Trichomanes poeppigii Presl, Hymenophyllaceae 4 1 . 1843. TYPE: "Habitat in Peruvia," Poeppig(ho- lotype, w?). Stem long-creeping, filiform, sparsely provided with scattered brown trichomes. Leaves well- spaced, 4-12(-18) cm long, 1-3 cm broad, sub- sessile or short-petiolate. Lamina linear- or ob- long-lanceolate, deeply lobed to pinnatifid, the axes, veins and margin provided with dark brown tri- chomes that are stellate or forked from the base. Venation catadromous, lacking false veins. Sori 1-few at or near segment apex, fully immersed in the segment tissue. Indusia salverform, neither bil- abiate nor dark-margined, the mouth flaring, re- ceptacle long-exserted. In deep forests, on tree trunks, very rarely on wet clay banks, 350-2300 m, Loreto and Huanuco to Cuzco and Madre de Dios. Southern Mexico to Panama; West Indies; Trin- idad; South America, to Brazil and Uruguay. Smaller specimens could be mistaken for Tri- chomanes reptans or other related species of subg. Didymoglossum, which are similar in their deli- cate, long-creeping stems, small, pinnatifid leaves, marginal stellate trichomes, and general aspect. However T. polypodioides lacks the false veinlets common in that subgenus, and trichomes are sparsely scattered along the stem, unlike the dense, blackish indument found on stems of T. reptans and its relatives. Loreto: Sierra del Pongo, Mexia 6228 (GH, MO, uc, us). Huanuco: E of Tingo Maria (as San Martin), Allard 21391 (GH, us). Pasco: Pichis Trail, Yapas (as Junin), Killip & Smith 25543 (F, GH), 25555 (F). Junin: Schunke Hacienda above San Ramon, C. Schunke A240 (GH, us). Cuzco: Altura del Rio Tocate, Bues 1744 (us). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster el al. 10713 (F). 19. Trichomanes tanaicum J. W. Sturm in Mart., Fl. bras. 1(2): 260. 1859. TYPE: Brazil, Para, Rio Acara, Spruce 410 (holotype, B; possible isotypes ["Spruce 44"], K.!, us; photo, A of K). Lacostea tanaica (J. W. Sturm) Prantl, Unters. Morph. Gefasskrypt. 1: 50. 1875. Trichomanes ankersii Hooker & Grev. var. tanaicum (Sturm) Sadebeck in Engler & Prantl, Nat. Pflan- zenfam. 1(4): 105. 1899. Stem long-creeping, stout and wiry, 0.5-0.8 mm in diameter, amply provided with brown rhizoids which extend onto the rachis and adhere to tree trunks. Leaves subdistant to remote, mature ones 3-15 cm long, 0.7-1.5 cm broad, subsessile or scarcely petiolate. Lamina linear, subentire or lobed (at base occasionally incised nearly to the rachis), the rachis abaxially often provided with rhizoids, otherwise glabrous. Venation catadromous, veins free, 1 -several times-forked in the lobes, false veins lacking. Sori 1 or a few, terminating the lobes, not or scarcely immersed in the segment tissue. In- dusia salverform, not or scarcely bilabiate, the mouth rather widely flaring, not dark-margined, receptacle scarcely to long-exserted. In deep forests, commonly hemiepiphytic on tree trunks, 100-130 m, Loreto, Ucayali. Venezuela; Colombia; Brazil; Amazonian Peru. The lamina in Trichomanes tanaicum is com- monly lobed less than one-third to the rachis, but rarely (as in Klug 92) some of the leaves are deeply pinnatifid. Nevertheless, the narrow laminae with their linear outline distinguish them from other species in section Lacostea. Loreto: Mishuyacu, near Iquitos, Klug 92 (F, us). Prov. Maynas, Dist. Nanay, Santa Maria de Nanay, J. Schunke V. 2449 (F, GH). Ucayali: Prov. Coronel Portillo (as Lor- eto), Rio Mazan, J. Schunke 300 (F, GH, uc, us, USM). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 89 20. Trichomanes tuerckheimii Christ, Hedwigia 44: 361. 1905. LECTOTYPE (designated here): Guatemala, Alia Verapaz, Cubilquitz, von Tuerckheim 8348 (lectotype, P!; isolectotypes, K., P!). LECTOPARATYPE: Peru, Loreto, near Leticia, Ule 6228 (K!, P!). Stem long-creeping, relatively stout and wiry, 0.5-1.2 mm in diameter, sparsely to abundantly provided with brown rhizoids that extend onto the leaf axes and adhere to tree trunks. Leaves sub- distant to remote, larger ones (9-)12-20(-30) cm long, 2-9 cm broad, subsessile. Lamina broadly oblong, pinnatisect, the rachis, costae and (some- times) veins and margins abaxially provided with simple, reddish brown trichomes that adhere to tree trunks, free portion of larger segments 2.8- 4.5 cm long and 0.8-1.2 cm broad, entire to shal- lowly crenate. Venation catadromous, veins free, simple, false veins lacking, but often some very short, darkened lines scattered between the veins. Sori few to several on the pinna-lobes, not or scarcely immersed in the segment tissue. Indusia tubular, neither bilabiate nor dark-margined, mouth not expanded, receptacle commonly long- exserted. Thus far known in Peru only from the lecto- paratype, cited above, on tree trunks; elsewhere hemiepiphytic, in forests on tree trunks, sea level to 500 m. Southern Mexico to Panama; Surinam to Co- lombia; Amazonian Peru. Included on the sheet with the syntype at Paris is a fragment of a 2-pinnatifid leaf with nearly laciniate lobes, which might have been mounted here by mistake. It resembles the more highly di- vided specimens which have been described as Trichomanes pedicellatum and T. subsessile. Re- lationships with these are discussed under T. an- kersii. (Also described there are the dark lines in the lamina which might be mistaken for the false veins of sect. Didymoglossum.) 21. Trichomanes ankersii Hooker & Grev., Icon, fil. 2, t. 201. 1831. TYPE: Guyana, Demerara, Ankers (holotype, K; photo, us). Stem long-creeping, relatively stout and wiry, 0.5-1.2 mm in diameter, sparsely to abundantly provided with brown rhizoids that extend onto the leaf axes and adhere to tree trunks. Leaves sub- distant to remote, larger ones 5-18 cm long, (1.8-)2-6 cm broad, subsessile. Lamina oblong, pinnatisect, or pinnate to a broadly winged rachis, the rachis, costae, and (sometimes) veins abaxially provided with simple, reddish brown trichomes that adhere to tree trunks, free portion of larger segments 1.5-2.8(-3) cm long and 0.5-0.8 cm broad, deeply crenate or crenate-serrate. Venation at least partly catadromous, veins free, commonly simple, but some- to many-forked, false veins lacking, but often some very short, darkened lines scattered between the veins. Sori few to several on the pinna lobes, not or scarcely immersed in the segment tissue. Indusia tubular, neither bilabiate nor dark-margined, mouth scarcely to slightly ex- panded, receptacle commonly long-exserted. In forests, commonly hemiepiphytic, tightly ap- pressed to tree trunks, sea level to 1 000 m, Loreto, Huanuco, Pasco, Cuzco. The Guianas to Colombia, south to Brazil and Bolivia. This, Trichomanes tuerckheimii, and T. tanai- cum, all members of sect. Lacostea, are commonly hemiepiphytic. Their leaves usually have the dis- tinctive habit of being tightly appressed to tree trunks. They cling to the bark by means of rust- colored, prehensile trichomes that are abundant along the stems, rachises, and in T. ankersii and T. tuerckheimii, on midribs and veins. Some col- lectors' labels describe them as "plastered to the bark of trees." Another character peculiar to the three species (though very rare in T. tanaicum) is the presence of very short, dark lines seen scattered between and parallel with the veins. They some- what resemble the false veins of sect. Didymo- glossum, except that they are not lengthy scler- enchymatous strands. Rather, they merely appear to be a series of (usually) two or three constricted and occluded cells that are mostly found toward the segment margin. Very closely related to Trichomanes ankersii are T. pedicellatum Desv. and T. subsessile Splitg., of northern South America and the West Indies. These appear to differ only in their more highly divided sterile lamina, which is 2- to 3-pinnatifid. Further study may prove them all to be conspecific. Loreto: Mishuyacu, near Iquitos, Klug 149, 1135 (F, us). Sierra del Pongo, Mexia 6289 (GH, MO, uc, us). Huanuco: 5 km NE of Tingo Maria, Stork & Horton 9518 (F, GH, uc, us). Pasco: Prov. Oxapampa, Valle del Palcazu, Leon 706 (F, GH). Cuzco: Prov. Quispicanchi, near Inambari, Vargas 16460 (GH). 90 FIELDIANA: BOTANY 22. Trichomanes bicorne Hooker, Icon, pi., t. 892. 1854. LECTOTYPE (designated here): Brazil, Amazonas, Barra do Rio Negro, Spruce 1 178 (lectotype, K!; isolectotype, us!). LECTO- PARATYPE: Brazil, Amazonas, Sao Gabriel, Spruce 2334 (K!, us). Ptilophyllum bicorne (Hooker) Prantl, Unters. Morph. Gefasskrypt. 1: 48. 1875. Stem stout, decumbent or erect, abundantly provided with lustrous, castaneous, pluricellular trichomes. Leaves monomorphic, crowded or caespitose, 3-10 cm long, 3-4-pinnatisect, the axes (and sometimes veins) sparsely provided on the abaxial side with simple, castaneous, pluricellular trichomes. Petiole 0.5-4 cm long, conspicuously alate throughout, the alae broader than the petiole quite or nearly to base. Lamina ovate, the axes broadly alate, alae and segments often undulate. Venation catadromous, veins free, false veins lack- ing. Sori immersed in the segment tissue. Indusia short-tubular, borne in the forks of veins, each of which extend well beyond the indusial mouth, the mouth not dark-margined, receptacle long-exsert- ed. In dense forests, epiphytic, or on wet humus or decaying logs, sea level to 1 50 m, Loreto. A lectoparatype (Spruce 2334) is mounted on the same sheet with the lectotype at Kew. How- ever, there are other specimens bearing this Spruce number that represent the type collection of Tri- chomanes spruceanum Hooker, a closely related species with dimorphic leaves which occurs in northern South America. One sheet (BM) is the isotype of T. spruceanum. Another, presumably the holotype, is at Kew. The species is aptly named for its sori, which are borne at segment apices in the forks of veins. The veinlets, bordered on the outside with narrow wings of tissue, each extend well beyond the mouths of indusia and are often curved, thus appearing like two "horns." Loreto: Vicinity of Lago Llanchama near Rio Nanay, Croat 18705 (F, MO). Mishuyacu, near Iquitos, Killip & Smith 29988 (F, us). Maynas, Dist. Iquitos, Cacerio Mis- hana, Rimachi 1256 (GH, MO). 23. Trichomanes diversifrons (Bory) Sadebeck in Engler & Prantl, Nat. Pflanzenfam. 1(4): 108. 1899. Trichomanes elegans Rudge, PI. Guian. 24, /. 35. 1805, nom. illeg. (not L. C. Rich., 1792, or Poiret, 1808). Hvmenostachys diversifrons Bory, Diet, class, hist. nat. 8: 462. 1825. TYPE: "Guiane," Poiteau (holo- type, P?; isotype, L; photos, GH & us of L). Feea diversifrons (Bory) Copel., Phillip. J. Sci. 67: 74. 1938. Stem stout, erect. Leaves crowded to caespitose, 10-35 cm long, strongly dimorphic, sterile ones elliptic or lanceolate, deeply pinnatisect, short-pet- iolate, fertile ones commonly longer, linear, sub- entire, long-petiolate. Sterile lamina (2-)3-8 cm broad, deeply divided into subfalcate segments 4- 7 mm broad, the rachis abaxially provided with dark, simple trichomes and often flagellate and proliferous at the tip. Venation catadromous, a few to many veins anastomosing toward the segment margin, 4-8 of them issuing from the rachis be- tween adjacent costae, false veins lacking. Fertile lamina simple, essentially entire, with sori ar- ranged in a nearly continuous line along each mar- gin, fully immersed in tissue in the forks of veins. Indusia subconical, not bilabiate, the mouth nei- ther flaring nor recurved, strongly indented be- tween the vein tips, receptacle short- to long- exserted at maturity. Terrestrial in wet forests, in wooded ravines and on ridges and banks, 1 50-1 300 m, Amazonas and Loreto to Junin, Madre de Dios. Southern Mexico to Panama; the Guianas to Colombia, south to Brazil and Bolivia. This is often confused with Trichomanes trollii, under which see further discussion. Amazonas: Prov. Bagua, Quebrada Tambillo, valley of Rio Maranon above Cascadas de Mayasi, Wurdack 2057 (GH, us). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Santa Rosa de Mishollo, J. Schunke V. 6815 (F, MO, uc). Loreto: Prov. Maynas, Brillo Nuevo and vicinity, Plowman et al. 6829 (GH). Huanuco: Tingo Maria, Tryon & Tryon 5277 (F, GH, us). Pasco: Prov. Oxapampa, Cordillera San Matias, Ledn 320 (F, USM). Junin: Cahuapanas, on Rio Pichis, Killip & Smith 26762 (F, GH, us). Madre de Dios: Prov. Manu, Cerro de Pan- tiacolla NNW of Shintuya, Foster et al. 10948 (F). 24. Trichomanes trollii Bergdolt, Flora 127: 256, 264, t. 3. 1933. TYPE: Bolivia, Dept. La Paz, San Carlos (Mapiri), Troll 2531 (holotype, B; frag., us!). Feea trollii (Bergdolt) Vareschi, Flora Venezuela 1 : 247. 1969. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 91 Stem decumbent or erect. Leaves crowded to caespitose, 10-25 cm long, strongly dimorphic, sterile ones elliptic, pectinate, short-petiolate, fer- tile ones commonly longer, linear, subentire, long- petiolate. Sterile lamina 1.5-4.5 cm broad, cut nearly to the rachis into subfalcate segments 1.5- 3(— 4) mm broad, the rachis abaxially provided with simple, dark brown trichomes and often fla- gellate and proliferous at the tip. Venation catad- romous, a few to many veins anastomosing toward the segment margin, 2(-3) of them issuing from the rachis between adjacent costae, false veins lacking. Fertile lamina simple, essentially entire, with sori arranged in a nearly continuous line along each margin, fully immersed in tissue in the forks of veins. Indusia subconical, not bilabiate, the mouth very slightly flaring and recurved, but not indented between the vein tips, receptacle strongly exserted at maturity. Terrestrial in dense, wet forests, often in wet ravines or along stream banks, 350-850 m, Ama- zonas to Pasco. Surinam to Colombia; Peru; Bolivia. This is closely related to the more robust Tri- chomanes diversifrons. Besides the characters in the key the latter can usually be recognized by its broader lamina (to 8 cm) and segments (4-7 mm), these being separated from the rachis by a con- spicuous wing ca. 2 mm broad on each side. The lamina of T. trollii is rarely more than 4 cm broad and are cut nearly to the rachis into segments which are commonly 3 cm broad or less. Additionally, the fertile lamina of T. diversifrons has dentate margins, due to the apices of the indusia which are deeply indented between the vein forks. In- dusial mouths of T. trollii are typically flush with the vein tips, so that the leaf margins are essentially entire throughout. Amazonas: Prov. Bagua. along roadside from Chiriaco to Puente Venezuela, Barbour 4461 (MO in part). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Palo Blanco. Plowman & Schunke 7431 (F). Huanuco: Ascent of Cerros del Sira. Wolfe 12263 A (GH, MO, us). Pasco: Prov. Oxapampa. W side of Cordillera de San Matias, D. Smith 2001 (MO). 25. Trichomanes botryoides Kaulf, Enum. fil. 263. 1824. TYPE: French Guiana, Poiteau (holo- type?, P). Stem erect. Leaves crowded to caespitose, 4-12 cm long, strongly dimorphic, sterile ones 1 -pin- nate, elliptic-lanceolate, short-petiolate, fertile ones commonly longer and long-petiolate, sori subdis- tichous on the nonfoliaceous axis. Sterile lamina 1.2-3 cm broad, the pinnae oblong-lanceolate, subentire to serrate, 3—4 mm broad, the petiole and rachis with scattered, brown, pluricellular tri- chomes, the rachis often flagellate and proliferous at the tip. Venation catadromous, the veins free, commonly 1 -forked, false veins lacking. Fertile leaves bearing numerous, stalked or subsessile, as- cending sori on each side of the primary axis. In- dusia urceolate, scarcely or not bilabiate, the mouth not flaring, receptacle strongly exserted at matu- rity. Thus far known in Peru from a single collection, roadside along a sandy, rocky stream bank, 350- 700 m, Amazonas. Elsewhere terrestrial or epipetric, commonly near banks of streams or rivers, 250-800 m; Panama; the Guianas and Colombia. The small size, strongly dimorphic leaves, and nonfoliaceous fertile leaves easily distinguish this from others in the genus. Amazonas: Prov. Bagua, ca. 40-43 km NE of Chiriaco, Barbour 4524 (MO, USM). 26. Trichomanes pinnatum Hedwig, Fil. gen. sp., /. 4,f. 1. 1799. LECTOTYPE (designated by Proctor, Flora Lesser Antilles. 89. 1977): Hedwig, /. 4,f. 1, based on specimen allegedly from Jamaica. Figure 12c. Trichomanes pennatum Kaulf., Enum. fil. 264. 1824. TYPE: French Guiana, without collector (holo- type, LZ destroyed). Neurophyllum pinnatum (Hedwig) Presl, Hymeno- phyllaceae 19. 1843. Stem short-creeping to decumbent, provided with dark brown or blackish pluricellular tri- chomes. Leaves essentially monomorphic (but fer- tile ones often somewhat larger, with longer pet- ioles), approximate or subcaespitose, to 70 cm long, 1 -pinnate, the axes sparsely to amply provided on the abaxial side with castaneous trichomes. Petiole about as long as the lamina, nonalate (sometimes marginate toward the lamina). Lamina ovate or subdeltoid, terminating in a conform apical seg- ment, or the rachis prolonged, flagellate and pro- liferous at the tip, the tissue thin and translucent, glabrous. Venation catadromous, the veins free, 92 FIELDIANA: BOTANY except connected at the tips by a continuous mar- ginal vein, false veins copious and perpendicular to true veins. Sori arranged in a nearly uninter- rupted line along pinna margins at the tips of veins, not immersed in the tissue, often stalked. Indusia narrowly tubular, not or scarcely bilabiate nor dark margined, mouth not flaring, receptacle long- exserted. Terrestrial, in deep forests or wooded ravines, in humus or clay banks of streams, 150-1000 m, Amazonas and Loreto to Junin and Madre de Dios. West Indies and Trinidad; Mexico to Panama and south to Brazil and Bolivia. Further differences between this and Trichom- anes hostmannianum are reviewed below in dis- cussion of that species. Amazonas: Prov. Bagua, valley of Rio Maranon near Cascadas de Mayasi, Wurdack 1939 (GH, us). San Mar- tin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Gran- ja Santa Ines, J. Schunke V. 3654 (F, GH). Loreto: 1 7 km SW of Iquitos on road to Puerto Almendra, Croat 18475 (F, MO, uc). Huanuco: Prov. Pachitea, Dist. Honoria, Bosque Nacional de Iparia, J. Schunke V. 1333 (F, GH, us). Pasco: Prov. Oxapampa, Palcazu Valley, Cabeza de Mono, D. Smith 3741 (F, MO). Junin: E of Quimiri Bridge, near La Merced, Killip & Smith 23924 (F, us). Ucayali: Along trail to Arboretum of Bosque von Humboldt Ex- perimental Station, km 86 of Pucallpa-Tingo Maria road, D. Smith 1226 (F, MO). Madre de Dios: Prov. Tambo- pata, Vargas 18625, 18711 (GH). 27. Trichomanes hostmannianum (Klotzsch) Kunze, Bot. Zeit. (Berlin) 5: 352. 1847. Neurophyllum hostmannianum Klotzsch, Linnaea 1 8: 532. 1844. TYPE: Surinam, Hostmann 75 (ho- lotype, B; isotypes, K, NY, u, us!; photos, GH & us ofK). Odontomanes hostmannianum (Klotzsch) Presl, Epi- mel. hot. 21. 1849. Ptilophyllum hostmannianum (Klotzsch) Prantl, Un- ters. Morph. Gefasskrypt. 1: 49. 1875. Stem decumbent to erect, provided with dark brown pluricellular trichomes. Leaves essentially monomorphic (but fertile ones often larger, with longer petioles), caespitose, to 30 cm long, 1 -pin- nate, the axes sparsely provided on the abaxial side with brown trichomes. Petiole commonly as long as or longer than the lamina, not alate, or narrowly so toward the lamina. Lamina ovate or subdeltoid, terminating in a subconform apical segment, or the rachis prolonged, flagellate and proliferous at the tip, the tissue relatively thick, commonly obscure, glabrous, pinna margins ser- rate, the serrations obtuse to subacute. Venation catadromous, the veins free, a lateral marginal vein faint and discontinuous or (more commonly) lack- ing, false veins lacking or a few very rare and faint ones perpendicular to true veins. Sori arranged in a nearly uninterrupted line along pinna margins at tips of veins, not immersed in the tissue, fre- quently stalked. Indusia conical to tubular, neither bilabiate nor dark-margined, mouth not flaring, receptacle exserted. Terrestrial in dark, wet forests and wooded rav- ines, often in frequently inundated areas along streams, sea level to 450 m, Amazonas, San Mar- tin, Loreto. Surinam to Colombia; Amazonian Brazil and Peru. This and the closely related Trichomanes pin- natum are sometimes confused in herbaria. Be- sides the differences noted in the key they also differ in size and in leaf texture. Leaves of T. host- mannianum do not exceed 30 cm, pinnae are rare- ly more than 6 cm long and 1 cm broad, and tissue of mature leaves is commonly dull green and ob- scure. Leaves of T. pinnatum often are 70 cm long, with pinnae to 1 5 cm long and 2 cm broad, and tissue is typically thin and translucent, so that the false veins are easily visible with little magnifi- cation. Contrary to previous reports, false veins do oc- cur in T. hostmannianum. They are infrequent and scattered and so faint as to be overlooked, or un- detectable, in the thick leaf tissue. However, in immature leaves or in the rare thin-textured ones, high magnification sometimes reveals a few or sev- eral of them, perpendicular to the true veins as in T. pinnatum. As this character is so difficult to observe, it is rather ineffective in delineating T. hostmannianum from species in other sections of the subgenus; yet it serves to further establish its position in sect. Neurophyllum (Presl) Moore, along with T. pinnatum. Morton ( 1 968) placed it outside the section because of the supposed lack of false veinlets. Amazonas: Nazareth, Osgood 25 (F, us). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, near Granja San Ysabel, J. Schunke V. 10313 (F, MO). Loreto: Rio Tacsha Curaray, Croat 20403 (F, MO, uc). Dist. Iqui- tos, trail through Versailles, Mexia 6500 (GH, MO, uc, us). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 93 28. Trichomanes humboldtii Lell., Mem. New York Bot. Gard. 38: 35. 1984, nom. nov. for Trichomanes heterophyllum Willd. and with the same type. Trichomanes heterophyllum Willd., Sp. pi. ed. 4, 5: 503. 1810, nom. illeg., not T. heterophyllum (Sm.) Poir. 1808. TYPE: Brazil, Amazonas, Javita, Humboldt (holotype, B, Herb. Willd. 20210; pho- tos, F, GH). Homoeotes heterophylla Presl, Gefassbiindel Farm 24. 1847, nom. nov., based on Trichomanes hetero- phyllum Willd. and with the same type. Feea humboldtii Bosch, Ned. Kruidk. Arch. 4: 347. 1859, nom. nov., based on Trichomanes hetero- phyllum Willd. and with the same type. Feea heterophylla Copel., Philipp. J. Sci. 67: 73. 1938, nom. nov., based on Trichomanes heterophyllum Willd. and with the same type. Stem wiry, long- to short-creeping. Leaves deep- ly pinnatisect to nearly pinnate, subdistant or re- mote, to 22 cm long, dimorphic, sterile ones short- petiolate, to 3 cm broad, fertile ones long-petio- late. far exceeding the sterile ones and 1 cm broad or less, the petiole essentially nonalate, sparsely or amply provided with castaneous, pluricellular tri- chomes. Sterile lamina oblong or oblong-elliptic, commonly equaling or longer than the petiole, the apex pinnatifid. Venation catadromous, veins free and dichotomously branched, false veins lacking. Fertile lamina linear, much shorter than the pet- iole, with a few sori terminal on each pinna and fully immersed in the tissue. Indusia subconical or broadly tubular, not bilabiate, the mouth not (or scarcely) flaring nor dark-margined, the recep- tacle long-exserted. Terrestrial, in open to dense forests, commonly in wet, sandy places, sea level to 1400 m, San Martin, Loreto. Venezuela; Colombia; Amazonian Peru and Brazil. There has been some confusion with nomen- clature of this species. Lellinger's (1984) citing of Trichomanes humboldtii "Bosch" (1858) was in error, since there was no such name; instead, it was "Feea humboldtif that Bosch had published. However, Lellinger's use of 'T. humboldtii"'' is le- gitimate, as a nom. nov. (Art. 72); therefore this name is correct for the species, with Lellinger as the author. San Martin: Rioja, Soukup 5055 (us). Rio Negro, Woytkowski 6211 (GH, MO, us). Loreto: Prov. Maynas, Rio Nanay, Mishana, Foster & Foster 4103 (GH); McDanielet al. 22121 (F). Mishuyacu, near Iquitos, Klug 1246 (F, us). 29. Trichomanes crinitum Sw., Prodr. 136. 1788. TYPE: Jamaica, Swartz (holotype, SET; iso- types, BM!, s; photos, us of BM & s). Ragatelus crinitus (Sw.) Presl, Hymenophyllaceae 16. 1843. Stem erect. Leaves monomorphic, caespitose, 5-25 cm long, l-2(-2.5) cm broad, moderately to amply pubescent, the trichomes delicate, un- branched, unicellular (or rarely pluricellular) be- yond a short, enlarged, basal cell. Petiole nonalate, or narrowly so toward the lamina. Lamina linear, cut nearly or quite to the rachis into deeply lobed to pinnatifid, patent, pinnae, gradually reduced to a pinnatifid apex, lamellae lacking on veins. Ve- nation catadromous, the veins free, false veins lacking. Sori terminating the veins on the apical part of the pinnae. Indusia tubular, deeply im- mersed in the segment tissue or, if partially im- mersed, broadly alate on each side, the mouth bilabiate and somewhat flaring, the lips not dark- margined, receptacle exserted. Apparently known thus far from two collections in Peru, in damp elfin forest, on tree trunks, ca. 1850 m, Huanuco. Elsewhere epiphytic or epi- petric, 700-2300 m. West Indies; Costa Rica; Venezuela; Colombia; Ecuador; Peru. Trichomes in many species of subg. Achomanes are pluricellular, with two to several greatly elon- gate cells, these often springing from a very short, widened basal cell. In species most closely related to Trichomanes crinitum, these are often mixed with a number of unicellular (above the basal cell) trichomes. However, T. crinitum differs from its Peruvian relatives in having all, or nearly all, of its laminar trichomes unicellular above the basal cell. Huanuco: SW slope of Rio Llullapichis watershed, on the ascent of Cerros del Sira, Dudley 13525 (GH), 13545 (GH, us). 30. Trichomanes martiusii Presl, Hymenophyl- laceae 36. 1843. TYPE: a renaming of T. pi- losum (sensu Martius in part, Icon. pi. crypt. 104, t. 68 right) presumably based on "Brazil, 94 FIELDIANA: BOTANY Arara-coara" (Araracuara, on Rio Caqueta, now Colombia), Martius (holotype, M; iso- types, BR, L; photo, us of L). Trichomanes plumula Presl, Hymenophyllaceae 15, 36. 1843. TYPE: based on Martius in part. Icon. pi. crypt. 104, /. 68 left, specimens probably as for T. martiusii. Ptilophyllum martiusii (Presl) Prantl, Unters. Morph. Gefasskrypt. 1: 48. 1875. Stem erect to decumbent. Leaves monomor- phic, crowded to caespitose, 1 0-40 cm long, (2-)3- 8 cm broad, densely (on axes) to amply pubescent, the trichomes unbranched, unicellular beyond the short basal cell, or pluricellular on petiole and rachis. Petiole nonalate. Lamina narrowly elliptic or lanceolate, gradually reduced to a pinnatifid apex, cut nearly or quite to the rachis into linear pinnae, the proximal ones commonly deflexed, subentire to crenulate or serrate, or rarely with a few, scattered lobes, with numerous, crestlike la- mellae borne on the veins abaxially. Venation ca- tadromous, the veins free, false veins lacking. Sori terminating the veins on the apical part of seg- ments. Indusia tubular or narrow-conical, deeply immersed in the segment tissue, the mouth trun- cate, not dark-margined, sometimes apparently bilabiate due to narrow wings of tissue extending beyond the mouth on two sides, receptacle com- monly exserted. On tree trunks in forests, or on clay banks along roads and streams, sea level to 150 m, thus far found only in Loreto, but locally common there. Surinam to Colombia; Peru; Amazonian Brazil. The shape of the indusium is somewhat vari- able. The mouth is commonly truncate and not at all bilabiate, but frequently the wings of tissue along the sides extend beyond the mouth, giving it a "horned" appearance, much like Trichomanes bi- corne. This is a distinctive species in Peru, because of the crestlike lamellae borne on many of the veins on the abaxial side, perpendicular to the plane of the lamina. These processes are similar to those occurring in several species of Hymeno- phyllum. On some of the more densely hirsute specimens of T. martiusii, the lamellae are par- tially obscured by the indument and thus they have been mistaken for other species in subg. Achomanes, particularly T. pilosum Raddi, of Bo- livia and southern South America. Loreto: Prov. Maynas, Rio Nanay, behind Mishana, Gentry & Revilla 20704 (MO, uc, USM). Mishayucu, near Iquitos, Klug 196, 51 1 (F, us). Prov. Maynas, 10 km S of Iquitos, Tryon & Tryon 5179 (GH, us). Timbuchi on Rio Nanay, LI. Williams 957 (F, us). 31. Trichomanes lucens Sw., Prodr. 136. 1788. TYPE: Jamaica, Swartz (holotype, SET; iso- type, BM; frag., B; photos, us of SBT & BM), not T. lucens Hooker & Grev., 1827. Trichomanes lambertianum Hooker, Sp. fil. 1: 139, /. 4 IB. 1846. TYPE: Peru, Huanuco, Pillao, Ruiz & Pavon (holotype, K; isotype, B; photos, GH & us of K). Ptilophyllum lambertianum (Hooker) Prantl, Unters. Morph. Gefasskrypt. 1: 48. 1875. Stem erect to decumbent. Leaves monomor- phic, crowded to caespitose, 10-60 cm long, (2.5-)3-12 cm broad, amply to densely pubescent (indument often obscuring the laminar surface), the trichomes unbranched, unicellular and pluri- cellular beyond the scarcely evident basal cell. Pet- iole nonalate. Lamina linear to lanceolate, pin- nate-pinnatifid (occasionally 2-pinnate-pinnatifid toward base), gradually reduced to a pinnatifid apex, pinnae narrow-deltoid, mostly ascending, lacking lamellae. Venation catadromous, the veins free, false veins lacking. Sori commonly terminal on ultimate segments near apex of pinnae. Indusia tubular, deeply immersed in the segment tissue, the mouth scarcely or slightly flaring, not dark- margined, sometimes apparently bilabiate due to the narrow wings of tissue extending beyond the mouth on two sides, receptacle commonly exsert- ed. In forests and wooded ravines, pendent from trees or wet rock crevices, or in sphagnum on sandy banks, 2150-3500 m, Cajamarca, Amazonas, Huanuco, Pasco, Cuzco. Jamaica; Costa Rica; Venezuela and Colombia, south to Bolivia; Brazil. With its l-(2-)pinnate-pinnatifid leaves and the copious, long trichomes that often obscure the lamina surface, this should not be mistaken for any other species of subg. Achomanes. Cajamarca: Prov. Jaen, Paso de Huascarai, 15 km SE of Huancabamba, Fosberg 27842 (us). Amazonas: Prov. Chachapoyas, Cerros de Calla Calla above Leimebamba, Hutchison & Wright 5568 (F, GH, MO, uc, us). San Mar- tin: Prov. Mariscal Caceres, Rio Abiseo National Park, Young & Ledn 4471 (F). Huanuco: Playapampa, Mac- bride 4499 (F, GH, us). Pasco: Prov. Oxapampa, Cordi- llera Yanachaga, Cerro Pajonal, Foster 9047 (F, MO). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 95 Cuzco: Prov. La Convention, Cordillera Vilcabamba, Dudley 10789 (F, GH, MO). 32. Trichomanes pellucens Kunze, Linnaea 9: 104. 1 834. TYPE: Peru, Tocache, upper Rio Hual- laga, Poeppig (holotype, LZ destroyed; iso- types, BR!, w; possible isotype, L!; photos, GH, HB & us of BR). The epithet T. pellucidum was used by Presl (Hymenophyllaceae 15. 1843), but later he indicated this was an error for T. pellucens Kunze (Epimel. bot. 16. 1851). Ptilophyllum pellucens (Kunze) Prantl, Unters. Morph. Gefasskrypt. 48. 1875. Stem erect to decumbent. Leaves monomor- phic, close, contiguous or caespitose, 8-30 cm long, 3-7 cm broad, sparsely to moderately pubescent, the trichomes unbranched, light to dark brown, those of the veins and margins unicellular beyond a very short basal cell, those of primary axes 1-3- celled beyond the basal cell. Petiole rather con- spicuously alate halfway or more to the stem. Lam- ina narrowly or broadly lanceolate, gradually re- duced to a pinnatifid apex, cut nearly or quite to the rachis into adnate, linear or linear-oblong, pat- ent pinnae (or basal ones deflexed), their margins subentire to crenate, lacking lamellae. Venation catadromous, the veins free, false veins lacking. Sori borne at tips of veins in the forks, near and at segment apices. Indusia tubular, deeply im- mersed in the segment tissue, the mouth not bil- abiate, flaring, or dark-margined, receptacle com- monly exserted. Epiphytic in forests, or terrestrial, most com- monly in sandy soil in ravines or on stream banks, 1 50-800 m, Amazonas, San Martin, Cuzco, Madre de Dios. Venezuela; Colombia; Ecuador; Peru; Brazil. Amazonas: Prov. Bagua, NE of Chiriaco, Barbour 4461 (MO in part), 4462 (USM), 4575, 4518 (MO). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Palo Blanco, Plowman & J. Schunke V. 7434 (F); J. Schunke V. 5722 (F, MO). Cuzco: Prov. Paucartambo, Cosnipata Valley, Rio Tono, Wachter el al. 134 (F). 33. Trichomanes plumosum Kunze, Linnaea 9: 104. 1834. TYPE: Peru, "in sylvis montanis ad Pampayaco" (Pampayacu, Dept. Huanu- co), Poeppig diar. 1107 (holotype, B; isotypes, BR!, K; photo, BM of BR). Trichomanes elatum Desv., Mem Soc. Linn. Paris 6(3): 327. 1827 (not Forst., 1786, or Bosch, 1861). TYPE: "habitat in America calidiori," without collector (holotype, P; photo, GH). Stem short-creeping to decumbent. Leaves monomorphic, subdistant to contiguous, (rarely caespitose), mature ones 1 5-50 cm long, 3-8 cm broad, moderately to amply pubescent, the tri- chomes unbranched, those of the veins and mar- gins unicellular beyond a very short basal cell, those of the rachis (especially abaxially) predom- inantly dark brown, 1-5-celled above basal cell, stout, rigid and spreading, most of them terete toward their base. Petiole essentially nonalate, am- ply provided with dark brown trichomes. Lamina lanceolate, often broadly so, gradually reduced to a pinnatifid apex, cut nearly to the rachis into linear to narrow-deltoid or -oblong segments, these patent (or the basal ones deflexed), lacking lamel- lae, the margins subentire to crenate or serrate, often undulate. Venation catadromous, the veins free, false ones lacking. Sori borne near and at segment apices, at tips of veins, sometimes flanked by vein forks. Indusia tubular or narrow-conical, deeply to fully immersed in the segment tissue, the mouth neither bilabiate nor dark-margined, not or scarcely flaring, receptacle exserted. In dense, wet forests, occasionally epiphytic, but commonly on wet sandy or mossy soil, 550-2100 m, San Martin to Puno. Ecuador; Peru; Bolivia; Brazil. This and Trichomanes cristatum are easily con- fused. Some collections from Peru are interme- diate between the two, and there is evidence of hybridization involving these and other closely re- lated species. The indument on the abaxial side of the rachis seems to be the best character for dis- tinction, as indicated in the key. In T. cristatum most of these trichomes are tawny to orange, rath- er long, delicate and tortuous, with their usually four to six cells greatly flattened. Those of T. plu- mosum are predominantly dark brown (some blackish), shorter, stout and rigid, and terete at least toward the base. Careful examination of spec- imens is necessary to view these trichomes, as those on the adaxial side of the rachis differ little, being primarily light brown, even tawny, on both species. Curiously (and unfortunately) most of the speci- mens examined for this study had been mounted with abaxial side down, so that it was very difficult to determine the character of trichomes on the other side. Besides differences in indument, T. plu- 96 FIELDIANA: BOTANY mosum can usually be distinguished from T. cris- tatum (at least in Peru) by lamina shape: that of the latter is commonly linear, rarely more than 4 cm broad, whereas the lamina of T. plumosum is typically broadly lanceolate, usually more than 4 cm broad. San Martin: Tarapoto, Spruce 4766 (BR, w). Huanuco: SW slope of Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13051 (GH), 1 3128 (GH, us), 1 3512 (GH). Pasco: Prov. Oxapampa, 1 9 km W of Oxapampa, D. Smith 2209 (MO). Junin: La Merced, Macbride 5634 (F, GH). Cuzco: La Convencion, Biies 2028, 2140 (us). Madre de Dies: Prov. Manu, Shintuya, Chavez 804 (MO). Puno: Prov. Carabaya, San Gaban, Vargas 18875 (GH). Dept. Unknown: "ad saxa humida prope Sangari," Lechler 2548 (BR, K, w 3 sheets); although this is the type number of T. undulatum (= T. vandenboschii), these specimens rep- resent part of a mixed collection). 34. Trichomanes cristatum Kaulf., Enum. fil. 265. 1824. TYPE: Brazil, collector undesignated (holotype, LZ destroyed). Trichomanes sellowianum Presl, Hymenophyllaceae 15, 37. 1843. TYPE: "Brasilia," Sellow 197 (ho- lotype, PR?; possible isotypes, 2 unnumbered Sel- low sheets from "Brasilia," B!, K!). Stem short-creeping to decumbent, rarely erect. Leaves monomorphic, subdistant to contiguous, mature ones 15-50 cm long, 2-4(-5) cm broad, moderately to rather densely pubescent, the tri- chomes unbranched, those of the veins and mar- gins unicellular beyond a very short basal cell, those of the rachis abaxially tawny to orange, del- icate, most of them long and tortuous and with the cells flattened, predominantly unicellular be- yond the basal one (occasionally with 2-3 cells). Petiole essentially nonalate, amply provided with long, orange to tawny trichomes, or these dark brown toward the stem. Lamina linear to linear- lanceolate, commonly 5-12 times as long as broad, gradually reduced to a pinnatifid apex, cut nearly to the rachis into linear or narrowly oblong seg- ments, these patent (or basal ones deflexed), lack- ing lamellae, the margins subentire to crenulate, often undulate. Venation catadromous, the veins free, false veins lacking. Sori borne near and at segment apices, at tips of veins, often flanked by the vein forks. Indusia tubular or narrow-conical, deeply to fully immersed in the segment tissue, the mouth not dark-margined, not or slightly flar- ing, not bilabiate but sometimes appearing so due to slight projections of tissue along the flanking vein forks, receptacle exserted. Terrestrial, or occasionally on tree trunks, com- monly in wet forests on sandy soil or in open sphagnum bogs, sea level to 2000 m, San Martin and Loreto to Puno. Venezuela and Colombia, south to Brazil and Argentina. This species is very easily confused with Tri- chomanes plumosum; characters are discussed above in the treatment of the latter. Trichomanes cristatum and T. plumosum share nearly the same distribution and are often found growing together. In Peru, at least, there are frequent intermediates, which indicates a strong probability of hybridiza- tion. These problems are not confined to species of Peru but also occur in other taxa of the group of T. crispum L. throughout the Neotropics, as pointed out by Windisch (1988). Obviously there is a great need for careful analysis of plants in situ as well as further caryological study before ac- curate relationships are understood. San Martin: Rio Negro, Woytkowski 6210 (GH, MO, uc, us). Loreto: Prov. Maynas, Dist. Iquitos, Rio Nanay, McDaniel & Rimachi 18924 (F, MO). Huanuco: Tingo Maria (as San Martin), Allard 21585 (GH, us), 27559 (us). Pasco: Prov. Oxapampa, Palcazu, van der Werff el at. 8376 (MO, uc). Junin: E of Quimiri Bridge near La Merced, Killip & Smith 23952 (F, us). Cuzco: Prov. La Convencion, Valle de Santa Ana, Herrera 3008 (us). Puno: Prov. Sandia, Vargas 1 1832 (GH). 35. Trichomanes vandenboschii Windisch, Bra- dea 5(4): 57. 1988, nom. nov. for T. undula- tum Bosch. Trichomanes undulatum Bosch, Ned. Kruidk. Arch. 5(2): 147. 1861, nom. itleg. (not Swartz, 1788). LECTOTYPE (designated by Windisch, Bradea 5(4): 57. 1988): "Peruvia, prope Sangari" (Dept. Puno, Prov. Carabaya, Dist. Ayapata) Lechler 2548 (L; isolectotypes, BR!, P; photos, HB oft, GH of BR, GH & us of P). Specimens of Lechler 2548, a mixed collection at BR, K, and w (3 sheets), are not type material, but are instead T. plumosum. LECTOPARATYPES: "Peruvia, Tatanara" (Dept. Puno, Prov. Carabaya, Dist. Ayapata), Lechler 2503. 2571 (L?, P?). Stem short-creeping. Leaves monomorphic, ap- proximate to subdistant, 5-15 cm long, 1.8-3 cm broad, moderately to densely pubescent, the tri- chomes unbranched, those of the veins and mar- gins unicellular beyond a very short basal cell, TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 97 those of the rachis tawny to orange, delicate, many of them long and tortuous and with the cells flat- tened, frequently with 2-8 cells beyond basal one, as well as with many unicellular ones. Petiole es- sentially nonalate, sparsely provided with short, light to dark brown trichomes. Lamina oblong- to ovate-lanceolate, 3-4 times as long as broad, grad- ually reduced to a pinnatifid apex, cut nearly or quite to the rachis into narrowly oblong segments, these patent (or basal ones deflexed), lacking la- mellae, the margins entire to crenulate, often un- dulate. Venation catadromous, the veins free, false veins lacking. Sori commonly borne at segment apices, at tips of veins, often flanked by the vein forks. Indusia urceolate, fully immersed in the seg- ment tissue, the mouth somewhat flaring, not dark- margined, not or scarcely bilabiate, receptacle exserted at maturity. In deep, wet forests, on tree trunks or in wet, boggy places, rarely in crevices of wet rocks, 750- 1 800 m, Cuzco. Venezuela; Colombia; Ecuador; Peru; Bolivia; northern Brazil. This is closely related to Trichomanes cristatum, from which it is distinguished primarily by its smaller size and relatively broader lamina. Besides the key characters, it can usually be separated from both T. cristatum and T. plumosum by its some- what flaring indusium mouth. That of the other two species is scarcely or not at all expanded. The confused identities of species within this complex is illustrated by the fact that the number of the type collection is shared with specimens of T. plu- mosum, as noted above in the citation of types. Cuzco: Maranura, Beatriz, Bites 895 (us). Prov. La Convencion, Sahuayaco, Vargas 6288 (GH). Prov. La Convencion, Choquello, Vargas 8184 (uc). Prov. Pau- cartambo, Atalaya, Valle Kosnipata, Vargas 23155 (OH). chomes (to 4 mm), reflexed pinnae in the proximal portion of the lamina, and the petiole very nar- rowly alate halfway to the stem. Windisch (in litt.) stated that he had seen one specimen (AAU) from Peru (Loreto, Prov. Maynas, Plowman et al. 6633). Although this specimen has not been examined, duplicates (F, GH) are scarcely distinct from T. cris- tatum, as rachis trichomes are less than 2 mm long and proximal pinnae are patent or just a few of them reflexed. Only a few petioles are obscurely alate to marginate. The species is found in Trin- idad, the Guianas, Venezuela, Colombia, and Bra- zil, and future collections of this species complex from Peru need to be examined carefully to de- termine if T. accedens truly occurs in Peru. Trichomanes delicatum Bosch, Ned. Kruidk. Arch. 5(2): 145. 1861. TYPE: Ecuador, Quito, Cum- ing 21 (holotype, B; frag., L). This is closely related to T. crinitum in that the trichomes are unicellular above the short basal cell. It differs from that species in the alate petiole, the more shallowly lobed pinnae, and the glabrous margins of the indusia. It is to be expected in Peru, since it has been found in Colombia, Ecuador, and Bolivia. Trichomanes haenkeanum Presl, Hymenophylla- ceae 15, 36, 65. 1843. TYPE: Peru, mountains of Huanuco, Haenke (holotype, PR?). Trichomanes crispum var. haenkeanum C. Chr., In- dex fil. 641. 1905. In the protologue, Presl indicated a close rela- tionship with Trichomanes crispum. Characters used in the description certainly align T. haen- keanum with this group of species but are insuf- ficient to place it precisely. Until the type is located and compared with other species in the complex, nothing more can be determined. Comments Trichomanes accedens Presl, Epimel. bot. 14: 1851. SYNTYPES: Guyana, Rich. Schomburgk 27 1 (B; photos, GH, us); Hooker & Grev., Icon, fil., t. 12, based on a Guilding collection from St. Vincent (specimen not seen by Presl). This belongs to the Trichomanes crispum com- plex and is allied to T. cristatum. It differs from the latter primarily in its predominantly erect stem with contiguous leaves, much longer rachis tri- FamilyT: LOXOMATACEAE Loxomataceae Presl, Gefassbiindel Farm 31.1 847, as Loxsomaceae. TYPE: Loxoma Cunn. (often altered to Loxsoma). Stem long-creeping, slender to rather stout, branched, densely pubescent with stiff trichomes enlarged at base. Leaves to 5 m long, pinnate, commonly pubescent abaxially, circinate in ver- nation. Petiole lacking stipules. Sporangia borne in marginal sori on an elongate receptacle, within 98 FIELDIANA: BOTANY a more or less urceolate indusium, short-stalked (ca. 6 rows of cells), with an oblique annulus not interrupted by the stalk. This small family contains two genera: Loxoma of New Zealand, and the Neotropical Loxsomop- sis. I. Loxsomopsis Loxsomopsis Christ, Bull. Herb. Boissier II. 4: 399. 1904. TYPE: Loxsomopsis costaricensis Christ. Figure 13. Terrestrial. Stem bearing scattered fibrous roots and abundant, rigid, lustrous, dark trichomes which are enlarged at the base. Leaves monomorphic, well-spaced on the stem, 2-pinnate-pinnatifid to nearly 3-pinnate. Lamina subcoriaceous, glabrous adaxially, glabrous to pubescent abaxially. Veins free. Sorus marginal, paraphysate, the indusium narrow-cyathiform to urceolate, the rim entire, the elongate receptacle exserted. Spores trilete, glo- bose-tetrahedral, the surface coarsely tuberculate. The genus is probably represented by a single species. However, two very closely related species have been described (from Costa Rica and Bolivia, respectively), based on supposed differences in size of leaf, color of petiole, and shape of pinnae and indusia. Two others have been separated on the basis of other variable and doubtfully significant characters. Observations of a number of speci- mens throughout the range suggest that such char- acters vary with age of the plant and maturity of sori, and they demonstrate no significant corre- lation. glaucous abaxially. Petiole dark brown to atro- purpureous, sublustrous. Lamina subdeltoid, sparsely to amply provided on surface, veins and costules with light to dark brown, tortuous, septate trichomes, glabrous adaxially. Pinnae commonly well-spaced, subdeltoid, somewhat or slightly re- duced at base, with pinnules strongly ascending, segments and venation catadromous. Veins pin- nately branched, prominulous. Indusia urceolate to (especially at maturity) narrow-cyathiform. Scandent or trailing, or arching to pendent from clay banks, in elfin forests, often on exposed ridges, 2200-3400 m, Huanuco, Pasco, Cuzco. Ecuador and Peru. Range of the species perhaps should be extended to include Costa Rica and Bolivia. Specimens de- scribed as Loxsomopsis costaricensis Christ differ little, if at all, from L. pearcei. The type (Werckle & Brune 279, Costa Rica) has not been seen, but the original description and illustrations suggest the two species are synonymous. Loxsomopsis no- tabilis Slosson (type from Bolivia, R. S. Williams 1303) is supposedly distinguished by its glaucous abaxial surface and by the larger pinnae being greatly reduced at base. Two sheets of isotype (us) seem to illustrate this clearly, and yet another is- otype (P) is only slightly glaucous and pinnae are not very strongly reduced at base. Lack of consis- tency in all these features suggests that there is but one species of Loxsomopsis. Huanuco: SW slope of Rio Llullapichis watershed, as- cent of Cerros del Sira, Dudley 13420 (GH). Playapampa, Macbride4521 (F, GH. us). Pasco: Prov. Oxapampa, Cord. San Gutardo, Ledn 532 (USM). Cuzco: Valle San Miguel, La Convention, Bues 2119 (us). Prov. La Convention, Cordillera de Vilcabamba, Dudley 10713 (GH, MO). 1 . Loxsomopsis pearcei (Baker) Maxon, Proc. Biol. Soc. Wash. 46: 105. 1933. Figure 13. Dicksonia pearcei Baker, Ann. Bot. (London) 5: 197. 1891. TYPE: Ecuador, "Eastern Andes, 8000- 9000 ft.," Pearce 251 (holotype, K!; photo, us). Loxsomopsis lehmannii Hieron.. Bot. Jahrb. Syst. 34: 435. 1904. TYPE: Ecuador, prope Chinguinda, Cordillera Oriental de Sigsig, 1 800-2500 m, Leh- mann 5061 (holotype, LZ destroyed; isotype, us!). Dennstaedtia pearcei (Baker) C. Chr., Index fil. 218. 1905. Leaves 0.4-5 m long, 2-pinnate-pinnatifid to nearly 3-pinnate, sometimes inconspicuously Family 8: PLAGIOGYRIACEAE Plagiogyriaceae Bower, Ann. Bot. (London) 40: 484. 1926. TYPE: Plagiogyria (Kunze) Mett. Stem stout, erect to decumbent, indurated, bear- ing stiff, fibrous roots, essentially lacking indu- ment. Leaves pinnate, circinate in vernation, di- morphic. Petiole expanded basally into indurated stipules, commonly with a double row of aero- phores at base (these rarely discernible in dried plants). Sporangia exindusiate, not paraphysate, commonly covering the abaxial surface of fertile TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 99 FIG. 1 3. Loxsomopsis pearcei: a, habit; b, portion of fertile pinna. (From Prieto P-259, Ecuador, F.) 100 FIELDIANA: BOTANY pinnae, long-stalked (4-6 rows of cells), with an oblique annulus not interrupted by the stalk. Spores trilete, tetrahedral-globose, the surface irregularly tuberculate. The family consists of a single genus essentially confined to wet, montane areas in Asia, Austra- lasia, and tropical America. These ferns resemble some species ofBlechnum because the sterile leaves arise obliquely from the thick rhizome in a circular pattern, from the center of which spring several dimorphic, fertile leaves. Stems in Plagiogyria, however, lack indument whereas those of Blech- num are scaly, an easily observed character which immediately distinguishes the former from simi- lar-appearing Blechnum species. I. Plagiogyria Plagiogyria (Kunze) Mett., Abh. Senckenberg Na- turf. Ges. 2: 265. 1858. TYPE: Lomaria eu- phlebia Kunze = Plagiogyria euphlebia (Kunze) Mett. Figure 14. Terrestrial. Leaves to 2 m long, pinnatisect to 1 -pinnate, dimorphic (fertile ones erect, common- ly with longer petioles and narrower, constricted pinnae). Sterile pinnae subentire to serrulate or biserrate. Veins free, simple, paired at or near the base, or forked. Fertile pinnae subentire to erose, the margins at first slightly to strongly reflexed to protect developing sporangia, later spreading, or sometimes so strongly retroflexed that both edges touch on the adaxial side. The most recent studies on Plagiogyria have recognized about 50 species in the genus; however, these have been based on a number of mostly overlapping or quantitative characters, e.g., degree of serration on pinna margins, relative length of basal pinnae, degree of branching of veins, none of which seem to be consistent or significant. Therefore, we believe the actual number of species in Plagiogyria is more likely to be nearer 1 5. Based on study of type collections and on Peruvian spec- imens attributed to various species, we have con- cluded that there is only one somewhat variable species in Peru. References COPELAND, E. B. 1929. The fern genus Plagio- gyria. Philipp. J. Sci., 38: 377-417. LELLINGER, D. B. 1971. The American species of Plagiogyria sect. Carinatae. Amer. Fern J., 61: 110-118. 1 . Plagiogyria semicordata (Presl) Christ, Farnkr. Erde. 176. 1897. Figure 14. Lomaridium semicordatum Presl, Epimel. hot. 155. 1849. TYPE: "In sylvis Columbia," without col- lector (holotype, PRC!). Plagiogyria costaricensis Kuhn, Linnaea 36: 149. 1869. TYPE: Costa Rica, Volcan Barba, Wendland 1066 (holotype, c?; drawing, B!). Plagiogyria latifolia Copel., Philipp. J. Sci. 38: 411. 1929. TYPE: Peru, Huanuco, Cani, 7 mi NE of Mito, ca. 2600 m, Macbride 3432 (holotype, us!; isotypes, F!, OH!). Plagiogyria denticulata Copel., Philipp. J. Sci. 38: 4 1 2. TYPE: Bolivia, Dept. Santa Cruz, alt. 2600 m, Herzog 1954 (holotype, us!). Petiole and rachis stramineous, the former brown at base, 8-30 cm long. Sterile lamina to 60 cm long and 18 cm broad, 1 -pinnate to deeply pin- natisect, lanceolate-elliptic, tapering to a pinna- tifid apex, gradually reduced to base, where pinnae are '/2-2/j as long as central ones; pinnae contiguous at base with adjacent ones, or widely spaced, 3- 10 cm long, 0.5-1.0 cm broad, margins serrulate to biserrate, apex acute to short-attenuate; veins commonly 1-2-forked, or a few simple or paired at the costa. Fertile lamina slightly longer and nar- rower than the sterile; pinnae very widely spaced, commonly 3-4 mm broad, margins strongly re- troflexed at maturity. At edges of forests, ravines, on shaded clay and rocks banks, 1800-3600 m, Cajamarca, Amazon- as, San Martin, Huanuco. Cuba; Jamaica; Mexico; Guatemala; Costa Rica to Peru and Bolivia. Cajamarca: Hualgayoc, Soukup & Carmona Fa.5008 (us). Amazonas: Prov. Chachapoyas, Summit of Puma- Urcu SE of Chachapoyas, Wurdack 1159(F, GH, NY, uc, us). San Martin: Prov. Mariscal Caceres, Puerta del Monte, Young 1980 (USM). Huanuco: Yanano, Macbride 3830 (F, GH, us). Family 9: DICKSONIACEAE Dicksoniaceae Bower, Origin land fl. 591. 1908, as Dicksonieae. TYPE: Dicksonia L'Her. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 101 5cm FIG. 14. Plagiogyria semicordata: a-b, habit; c, apex of fertile leaf; d, apex of fertile pinna, (a from Cuatrecasas 5465, Colombia, F, b from Wurdack 1159, F, c-d from Little 9184, Colombia, F.) 102 FIELDIANA: BOTANY Stem stout to massive, usually not branched, prostrate to decumbent to erect and then often arborescent, indurated, densely covered with usu- ally long trichomes. Leaves usually large, ca. 1-3 m long, circinate in vernation, monomorphic to dimorphic (the fertile nearly lacking green tissue and more complex than the sterile), pinnate, gla- brous to pubescent. Petiole lacking stipules, not articulate to the stem. Veins free. Sori marginal, usually paraphysate with slender trichomes, en- closed within a usually firm and partly green adax- ial indusium and a thinner, light brown abaxial indusium, these separate or basally or fully joined. Sporangia with a 4-6-rowed stalk and a complete, oblique annulus. The Dicksoniaceae are a family of five genera and 35—40 species. Two genera are in Peru. The family, with the possible exception of Cys- todium of Malaysia, is natural and distinctive. It has been united with the Cyatheaceae, but that family differs in having scales on the stem and an abaxial sorus and the two are of uncertain affinity. Reference TRYON, R. M., AND A. F. TRYON. 1982. Dick- soniaceae, pp. 138-155, in Ferns and allied plants, Springer- Verlag, New York. Key to Genera of Dicksoniaceae a. Lamina 4-5-pinnate-pinnatifid, broadest at the base, its tertiary axes grooved on the adaxial side I. Culcita a. Lamina 2-3-pinnate-pinnatifid (in America), reduced at the base, its tertiary axes ridged (not grooved) on the adaxial side . II. Dicksonia I. Culcita Culcita Presl, Tent, pterid. 135. 1836. TYPE: Cul- cita macrocarpa Presl. Figure 15. Terrestrial. Stem prostrate, decumbent or rarely erect and to 3 m tall, stout. Leaves monomorphic, to ca. 3 m long. Lamina 4-5-pinnate or slightly more complex, the tertiary axes grooved adaxially, veins free. Sori marginal, the adaxial indusium joined basally to the thinner abaxial indusium, or the two separate. Spores tetrahedral-globose, tri- lete, nearly smooth, rugulose, or tuberculate. Culcita coniifolia of tropical America and C. macrocarpa of the Canary Islands, Azores, and Spain form subg. Culcita. The other five species, of Malaysia to Samoa and Australia, comprise the subgenus (or, better, the genus) Calochlaena Max- on. 1 . Culcita coniifolia (Hooker) Maxon, Annual Rep. Smithsonian 1911: 488. 1 9 1 2; J. Wash. Acad. Sci. 12:456. 1922. Figure 15. Dicksonia coniifolia Hooker, Sp. fil. 1: 70, t. 24A. 1 844. TYPE: Venezuela, (Dist. Federal), Caracas, Linden 538 (holotype, K; isotype, BR; photo, GH). Stem prostrate to 1 m tall, the apex with few leaves. Leaves to 3 m long, the petiole with a dense covering of trichomes at the base, often longer than the lamina. Lamina deltoid, to 4- or 5-pinnate- pinnatifid at the base, the major segments stalked, thinly pubescent, or glabrous with age, sterile ul- timate segments with bluntly acute lobes. Sorus single on each ultimate lobe. In wet elfin forests at ca. 2400-3100 m, Ama- zonas south to Cuzco. Southern Mexico, Central America, Greater Antilles, Venezuela and Colombia south to Peru; Mt. Itatiaia, Brazil. Culcita coniifolia is a distinctive species and should be mistaken for no others. Although of wide distribution, it has been rarely collected in Peru. Ama/onas: Prov. Chachapoyas, 18 km above Lei- mebamba on road to Balsas, Cerros de Calla Calla, Hutchison & Wright 5687 (F, GH). Along Rio Ventilla, 1-2 km W of Molinopampa, Wurdack 1468 (GH, USM). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 103 FIG. 1 5. Culcita coniifolia: a, stem and part of petiole; b, pinna; c, rachis and base of pinna, adaxial side; d, fertile ultimate segment, (a from Macbride 4519, r, b-d from Lent 732, Costa Rica, F.) 104 FIELDIANA: BOTANY Huanuco: Playapampa, Macbride 4519 (F). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8436 (MO). Cuzco: El Dorado, Rio Uru- bamba valley, Aug. 12, 1941, Bites (GH). Prov. La Con- vention, Dudley 10712 (GH). II. Dicksonia Dicksonia L'Her., Sert. angl. 30. 1788. TYPE: Dicksonia arborescens L'Her. Figure 16. Terrestrial. Stem erect, to 10 m tall, or decum- bent at the base, usually massive. Leaves mono- morphic to dimorphic (the fertile nearly lacking green tissue and more complex than the sterile), to ca. 3.5 m long. Lamina 2-pinnate-pinnatifid to 4-pinnate, reduced at the base, tertiary axes adax- ially ridged (not grooved), veins free. Sori mar- ginal, the adaxial indusium joined basally to the thinner abaxial indusium. Spores tetrahedral-glo- bose, trilete, granulate or reticulate. Dicksonia is a genus of about 20 species, from Malaysia to Samoa, St. Helena, tropical America, and the Juan Fernandez Islands. Reference STOLZE, R. G. 1 976. Dicksonia, in Ferns and fern allies of Guatemala. Part I. Fieldiana, Bot., 39: 99-102. Key to Species of Dicksonia a. Tertiary segments of the larger fertile pinnae each with few to several sori 1 . D. sellowiana a. Tertiary (ultimate) segments of the larger fertile pinnae each with a single sorus ... 2. D. stuebelii 1. Dicksonia sellowiana Hooker, Sp. til. 1: 67. 1844. TYPE: Brazil, Sellow (lectotype desig- nated here, K; isolectotype, HBG; photos, GH, us of HBG; lectoparatype, Brazil, Miers, K). Figure 16. Balantium karstenianum Klotzsch, Linnaea 20: 444. 1847. TYPE: "Columbia," Karsten, no. 9 (Coll. II) (holotype, B?; isotype, HBG; photo, GH of HBG). Dicksonia gigantea Karsten, Fl. columb. 2: 1 77, 1. 193. 1 869. TYPE: Colombia, (Cundinamarca), Andes of Bogota, Guadeloupe, Karsten (not located) (Karsten, HBG; photo, GH, may be authentic). Dicksonia karsteniana (Klotzsch) Moore, Index fil. 190(1860), 313(1861). Dicksonia spruceana Kuhn, Linnaea 36: 153. 1869. TYPE: Peru, (San Martin), Tarapoto, Spruce 4728 (holotype, B; isotypes, A!, BM, GH!; photos, GH, MO, uc of BM). Stem to 10 m tall, enclosed, at least basally in dense fibrous roots, with persistent leaf bases, to ca. 25 cm in diameter, bearing many leaves in a crown. Leaves to 3 m long, the petiole very short, densely covered with long trichomes. Lamina 2- 3-pinnate-pinnatifid, the secondary and tertiary segments nearly sessile, glabrous to thinly pubes- cent abaxially, sterile lobes bluntly acute to acute, the margin flat or curved. Sori few to several on the tertiary segments of the larger fertile pinnae. In wet woods or cloud forests, at 1 550-2400 m, Cajamarca and Amazonas, south to Cuzco. Southern Mexico, Central America, Venezuela, and Colombia south to Bolivia, Paraguay, Uru- guay and southeastern Brazil. Dicksonia sellowiana is a widespread and vari- able species. Several segregates are sometimes rec- ognized but these evidently represent minor vari- ations. The other American species are D. stuebelii of Peru and D. berteriana (Colla) C. Chr. of the Juan Fernandez Islands. Cajamarca: Prov. Cutervo, La Pucarilla, Ldpez & Sa- gdstegui 5457 (GH, HUT). Amazonas: Prov. Bagua, ca. 20 km E of La Peca, Harbour 2870 (MO). Huanuco: Cerros del Sira, Rio Llullapichis watershed, Dudley 13254 (F, GH, MO), 75259, 13373 (GH). Pasco: Oxapampa (as Jun- in), Soukup 2334 (F, GH). Prov. Oxapampa, Canyon de Huancabamba, Ledn 614 (F). Cuzco: Prov. La Conven- tion, Cordillera Vilcabamba, Dudley 10439, 10603, 11308(GH). 2. Dicksonia stuebelii Hieron., Hedwigia 45: 228, /. 72, / /. 1906. TYPE: Peru, (Amazonas), Tambo Ventilla, Pascomayo to Moyobamba, Stiibel 1076 (holotype, B). Similar to Dicksonia sellowiana, but the lamina is 2-pinnate-pinnatifid to 2-pinnate-pinnatisect, TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 105 FIG. 16. Dicksonia sellowiana: a, 2 pinnae; b, fertile ultimate segments, abaxial side; c, secondary axis, adaxial side, with ultimate segments, (a from Reiss 61, Brazil, F, b from Killip & Smith 18095, Colombia, F, c from Holm- Nielsen el al. 5645, Ecuador, F.) 106 FIELDIANA: BOTANY the larger fertile pinnae have the tertiary (ultimate) segments bearing a single sorus, and the sterile margins are recurved. In the Jalca zone and in wet sandy soil, ca. 2200- 3400 m, Amazonas. Endemic in northern Peru. This is evidently a rare and local species. It is unusual in growing within the range of the related Dicksonia sellowiana. Amazonas: Cerro de Fraijaco (Huaui-Huni), NE of Tambo de Ventilla. Pennell 15855 (GH). Prov. Chach- apoyas, 3-6 km W of Molinopampa, Wurdack 1410 (GH). Cerro Yama-uma, above Taulia, 12-15 km SE of Molinopampa, Wurdack 1679 (F, GH, uc, us, USM). Family 10: LOPHOSORIACEAE Lophosoriaceae Pic.-Ser., Webbia 24: 700. 1970. TYPE: Lophosoria Presl. Stem sometimes branched, massive, decumbent to erect and arborescent, indurated, densely cov- ered by long trichomes. Leaves usually large, ca. 1-5 m long, circinate in vernation, monomorphic, pinnate, more or less pubescent. Petiole lacking stipules, not articulate to the stem. Veins free. Sori exindusiate, on the abaxial surface of the segments, paraphysate with slender trichomes. Sporangia with a short, 6-rowed stalk and a complete, oblique annulus. The family Lophosoriaceae contains a single, American genus. Reference I. Lophosoria Lophosoria Presl, Gefassbiindel Farm 36. 1847. TYPE: Lophosoria pruinata (Sw.) Presl = Lo- phosoria quadripinnata (Gmelin) C. Chr. Fig- ure 17. Terrestrial. Leaves rarely 0.25 m , usually 2-3 m, sometimes to 5 m long. Lamina 2-pinnate- pinnatifid to 3-pinnate-pinnatisect, segments often glaucous abaxially and slightly to densely pubes- cent. Sori round, borne on the veins, the receptacle hardly elevated. Spores trilete, tetrahedral-glo- bose, with a large equatorial flange, coarsely tu- berculate to rugose. A monotypic genus, ranging from Mexico and the Greater Antilles, south to Bolivia and Brazil, also in southern Argentina and Chile, and the Juan Fernandez Islands. Lophosoria quadripinnata is a widely distrib- uted and distinctive species of the cloud forest zone. One high altitude variety that is local in Ecuador and Peru may be recognized. 1. Lophosoria quadripinnata (Gmelin) C. Chr., Skottsb. Nat. hist. Juan Fernand. 2: 16. 1920. Stem apex with few leaves. Petiole about as long as the lamina or somewhat shorter, with 3 con- voluted vascular bundles. Lamina very narrowly lanceolate to usually broadly ovate, to 3-pinnate- pinnatisect, with sessile or short-stalked segments, very sparsely to densely pubescent and often glau- cous abaxially, ultimate segments obtuse to sub- acute. Sori single on a fertile vein. Range of the genus. TRYON, R. M., AND A. F. TRYON. 1982. Lopho- soriaceae, pp. 156-161, in Ferns and allied plants, Springer- Verlag, New York. Key to Varieties a. Leaf ca. 2-4 rn long, pinnae spaced, patent, the central ones ca. 0.5-1.0 m long la. var. quadripinnata a. Leaf ca. 0.25-1 m long, pinnae imbricate, ascending, the central ones ca. 0.05-0.20 m long . Ib. var. contracta TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 107 FIG. 1 7. Lophosoria quadripinnata var. quadripinnata: a, stem and portion of petiole; b, pinna; c, base of fertile segment, (a from Wurdack 1752, F, b-c from Stork & Morton 10348, F.) 108 FIELDIANA: BOTANY 1 a. Lophosoria quadripinnata var. quadripinnata. Figure 17. Polypodium glaucum Sw., Prodr. 1 34. 1 788, not Houtt. 1 783. TYPE: Jamaica, Swam (holotype, B!, Herb. Willd. 19723; photo, GH, designated as type by Hieronymus, it may be the holotype or an iso- type). Polypodium quadripinnatum Gmelin, Syst. nat. 2(2): 1314. 1791, nom. nov. for Polypodium glaucum Sw. (not Houtt.) and with the same type. Polypodium pruinatum Sw., J. Bot. (Schrader) 1800(2): 29. 1802, nom. nov. for Polypodium glaucum Sw. (not Houtt.) and with the same type. Alsophila pruinata (Sw.) Kunze, Linnaea 9: 99. 1834. Lophosoria pruinata (Sw.) Presl, Gefassbiindel Farm 37. 1847. Alsophila quadripinnata (Gmelin) C. Chr., Index fil. 44. 1905. In wet forests, forest borders, on brushy slopes, in cloud forests and elfin forests, 700-3100 m, Cajamarca and Amazonas, south to Puno. Range of the genus. Throughout most of mountainous tropical America, this variety is a typical element of the cloud forest zone. Cajamarca: Prov. Celendin, Agua Colorado, Sanchez 206 (GH). Amazonas: Cerros de Calla Calla, above Lei- mebamba, Hutchison & Wright 5686 (F, GH, uc, us). Cerro Puma Urco, Chachapoyas, Soukup 4086 (F, us). San Martin: Monte Campana, Tarapoto, Spruce 4248 (GH, us). Huanuco: Panao, Bryan 407 (F, GH). Playapam- pa, Macbride 4860 (F, us). Pasco: Quillasu, Soukup 3290 (F, GH, uc, us). Yapas, Pichis Trail (as Junin), Killip & Smith 25456 (F, GH, us). Junin: Huacapistana, Ferreyra 3654 (USM). Ucayali: La Divisoria, Aguilar 849 (GH). Huancavelica: Prov. Tayacaja, Surcubamba, Stork & Horton 10348 (F, GH, uc, us). Cuzco: Prov. La Conven- tion, Dudley 10883 (F, GH). Pilahuata, Cerro de Cusil- luyoc, Pennell 13938 (GH, us). Puno: Prov. Sandia, San- dia to Chunchusmayo, Weberbauer 1333 (USM). 1 b. Lophosoria quadripinnata var. contracta (Hi- eron.) R. & A. Tryon, Rhodora 84: 1 26. 1 982. Alsophila contracta Hieron., Hedwigia 45: 236, /. 75, / 8. 1906. SYNTYPES: Peru, (Amazonas), near Inez and Calle-calle, Stitbel 1067 (B!; frag., GH!); (Amazonas), near Challuayacu and Tambo Cen- tamala, Stubel 1066 (B!). Wet, shrubby areas, 2800-3500 m, in Amazon- as and La Libertad. Ecuador and Peru. The var. contracta is evidently a high-altitude ecotype. The following collection is intermediate between the two varieties: Peru, Amazonas, Puma- Urcu, SE of Chachapoyas, Wurdack 791 (us). Amazonas: Prov. Chachapoyas, Pomacocha, Lopez el al. 4392 (GH). Prov. Chachapoyas, Cerros de Calla Calla slopes, Wurdack 1752 (GH, us, USM). Prov. Chachapoyas, between Leimebamba and Calla-Calla, Smith & I'dsquez 4978 (GH). La Libertad: Prov. Bolivar, cerca Nevado de Cajamarquilla, Ferreyra 1349 (GH, USM). Family 11: METAXYACEAE Metaxyaceae Pic.-Ser., Webbia 24: 701. TYPE: Metaxya Presl. 1970. Stem rather stout, sometimes branched, pros- trate to nearly erect, indurated, with dense, long trichomes, especially toward the apex. Leaves usu- ally large, to 2 m long, circinate in vernation, monomorphic, pinnate, glabrate. Petiole lacking stipules, not articulate to the stem. Veins free. Sori exindusiate, on the abaxial surface of the pinnae, paraphysate with slender trichomes. Sporangia with a 4-rowed stalk and a complete, slightly oblique annulus. The family Metaxyaceae contains a single, American genus. Reference TRYON, R. M., AND A. F. TRYON. 1982. Metax- yaceae, pp. 162-165, in Ferns and allied plants, Springer- Verlag, New York. I. Metaxya Metaxya Presl, Tent, pterid. 59. 1836. TYPE: Me- taxya rostrata (HBK.) Presl (Aspidium rostra- turn HBK.). Figure 18. Terrestrial or rarely on tree bases. Leaves to ca. 2 m long. Petiole usually with short trichomes at the very base. Lamina 1 -pinnate, with simple pin- nae, veins free. Sori roundish to elongate, the re- ceptacle nearly flat. Spores globose, trilete, gran- ulate. A monotypic genus ranging from southern Mex- ico and Central America; Guadeloupe; Trinidad; and French Guiana west to Colombia and south to Bolivia; Amazonian Brazil. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 109 FIG. 18. Metaxya rostrata: a, habit; b, portion of fertile pinna, abaxial side. (From Wurdack 1872, F.) 110 FIELDIANA: BOTANY 1. Metaxya rostrata (HBK.) Presl, Tent, pterid. 60. 1836. Figure 18. Polypodium rostratum Willd., Sp. pi. ed. 4, 5: 193. 1810, not Burm. 1 768. TYPE: Venezuela, Javita, Humboldt 966 (holotype, B!, Herb. Willd. 19691; photo, GH). Aspidium rostratum HBK., Nov. gen. sp. 1: 12. 1815, nom. nov. for Polypodium rostratum Willd. (not Burm.) and with the same type. Alsophila blechnoides Hooker, Sp. fil. 1: 35. 1844, nom. superfl. for Alsophila rostrata (HBK.) Mart. and with the same type. Stem prostrate, the apex with few leaves. Petiole about as long as the lamina, with 1 convoluted vascular bundle. Lamina narrowly lanceolate to ovate, 1 -pinnate, with a conform terminal pinna, the pinnae stalked, glabrous or nearly so abaxially, entire to finely serrate, the serrate apex acuminate to caudate. Sori 1-3 on each fertile vein. Woods and dense forests, usually in sandy, moist, well-drained soil, 100-800 m, Amazonas to Cuzco and Madre de Dios. Range of the genus. Metaxya is unusual in having the 1-pinnate- pinnatifid leaves of juvenile plants more complex than the 1 -pinnate leaves of adult plants. Amazonas: Prov. Bagua, Rio Maranon, above Cas- cadas de Mayasi, Wurdack 1872 (us, USM). San Martin: Prov. Mariscal Caceres, Isla de Pucunuchu, J. Schunke 4783 (F, GH). Prov. Mariscal Caceres, Rio Sion, J. Schunke 3524 (F, GH, us). Loreto: Iquitos, Killip & Smith 27489 (GH, us). Huanuco: Prov. Pachitea, Bosque Nacional de Iparia, J. Schunke 1620, 1833 (F, GH, us). Pasco: Prov. Oxapampa, Cordillera San Matias, Leon 332 (USM). Ucayali: vicinity of Aguaytia, Croat 20966 (us). Prov. Coronel Portillo, Bosque Nacional Humboldt, Vdsquez 3910 (F, MO). Cuzco: Prov. Quispicanchi, Vargas 16462 (GH). Prov. Paucartambo, Vargas 1 1245 (GH). Madre de Dios: Prov. Tambopata, Tambopata Nature Reserve, Barbour5191 (F). Family 12: CYATHEACEAE Cyatheaceae Kaulf., Wesen Farrenkr. 119. 1827. TYPE: Cyathea Smith. Stem usually massive, erect, arborescent, un- branched, very rarely slender and scandent, to small, decumbent and short-creeping, indurated, bearing scales and sometimes spines. Leaves usu- ally large, ca. 1-4 m long, circinate in vernation, monomorphic to rarely dimorphic, pinnate or very rarely entire, glabrous, pubescent and (or) scaly abaxially. Petiole lacking stipules, not articulate to the stem. Veins free or rarely reticulate. Sori ex- indusiate or indusiate, on the abaxial surface of the segments, usually paraphysate with slender tri- chomes or rarely with enlarged ones. Sporangia with a 4-rowed stalk and a complete, oblique an- nulus. The Cyatheaceae are a family of six genera and a few hundred species. All of the genera occur in Peru. This treatment has been adapted from the lit- erature cited under the family and the genera. There are at least 1 9 species, especially of Ecuador but also of Bolivia, that may well also occur in Peru, but these are too numerous to mention. Characters of all parts of the leaf are useful and often necessary for the accurate identification of the genera and species of Cyatheaceae. Of special importance are characters of the petiole scales. However, the scales may be abraded on the petiole of a mature leaf and their characters are best seen on the croziers. The height and diameter of the arborescent stem vary in a species depending upon the age of the plant and the growing conditions. Accordingly, these characters as well as other quantitative ones are usually not included in the species descriptions. Several special terms that are used in the Cy- atheaceae are the following. PETIOLE SPINES— Corticinate: The spine is an ex- tension of the cortex of the petiole; each bears, at least when young, a scale at its apex which is rather blunt when the scale falls off. Squaminate: The spine is not an extension of the petiole; it breaks, or can be broken, from the petiole at its base. It has a sharp apex that does not bear a scale. These spines have evidently evolved from petiole scales. PETIOLE SCALES— Conform: All cells, except those that may be borne on the edge, are similar in ori- entation, shape, and usually in size and color. Marginate: With a narrow to broad margin of cells different from those in the central portion of the scale in orientation, size, and usually in shape and color. Body: All of the scale except for cilia, teeth, or setae borne on the edge. Margins: The cellularly differentiated areas on each side of the central por- tion of marginate scales. Concolorous: All of the scale of one color or nearly so. Bicolorous: The margins definitely of lighter color than the central portion. Concordantly bicolorous: The lighter col- or confined to the differentiated margins and the darker color to the central portion of elongate cells. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. Ill Discordantly bicolorous: The lighter color of the margins extends into part of the central portion of elongate cells. PETIOLE SCURF— Large scales: The scales are small, but definitely larger than most of the scurf which is composed of minute scales and tri- chomes. INDUSIA— Hemitelioid: Attached at the base and often at the sides of the receptacle, but not com- pletely surrounding it. Meniscoid: Completely sur- rounding the receptacle, nearly flat or with slightly upturned edges. Cyathiform: Cup-shaped. Urceo- late: Urn-shaped. Subsphaeropteroid: Enclosing the sporangia except for a rather small opening at the apex. Sphaeropteroid: Completely enclosing the sporangia and with an apical umbo. References TRYON, R. 1970. The classification of the Cy- atheaceae. Contr. Gray Herb., 200: 2-53. . 1986. Cyatheaceae, in Harling, G., and L. Anderson, eds., Flora of Ecuador, 27: 17-56. TRYON, R. M., AND A. F. TRYON. 1982. Cy- atheaceae, pp. 166-212, in Ferns and allied plants, Springer- Verlag, New York. Key to Genera of Cyatheaceae a. Petiole scales conform I. Sphaeropteris a. Petiole scales marginate b b. Petiole scales with a dark, opaque apical seta c c. Petiole lacking spines, or with corticinate spines that bear, at least when young, a scale at the apex, other petiole scales almost appressed to the surface and attached at a usually pseudopeltate base; croziers lacking spines II. Alsophila c. Petiole with squaminate spines, many of these large, black, with a sharp apex; petiole scales patent, attached at a usually narrowed base; croziers with large, black, sharp spines III. Nephelea b. Petiole scales lacking a differentiated apical seta, the apex rounded to filamentous d d. Indusium absent IV. Trichipteris d. Indusium present e e. Spores lacking large pores, sometimes with variously distributed small pits or pores; veins free, the basal ones of adjacent segments extending to the margin above the sinus, rarely a few costal areolae present V. Cyathea e. Spores with three large equatorial pores, often also with smaller pits or pores; veins anas- tomosing to form costal areolae, or if free then the basal veins of adjacent segments connivent to the sinus . . VI. Cnemidaria I. Sphaeropteris Sphaeropteris Bernh., J. Bot. (Schrader) 1800(2): 122. 1802. TYPE: Sphaeropteris medullaris (Forster) Bernh. (Polypodium medullare Fors- ter). Figure 19. Terrestrial. Stem stout, erect and arborescent or rarely short-decumbent, lacking spines. Leaves monomorphic to slightly dimorphic, with scales, especially on the croziers and the base of the pet- iole, that are conform and with or without a dark apical seta. Petiole lacking spines or with corticin- ate spines. Lamina 1 -pinnate to 3-pinnate-pin- natifid, veins free or rarely partially anastomosing without included free veinlets. Sori round, often borne at the fork of a vein, exindusiate or with a hemitelioid or sphaeropteroid indusium. Spores tetrahedral-globose, trilete, echinate, porate, ver- ruca te or nearly smooth. Sphaeropteris occurs in tropical America, India and southeastern Asia to New Zealand and Pit- cairn Island. Among the 120 species, there are 23 in America and seven in Peru. References TRYON, R. 1971. The American tree ferns allied to Sphaeropteris horrida. Rhodora, 73: 1-19. WINDISCH, P. G. 1977. Synopsis of the Genus 112 FIELDIANA: BOTANY FIG. 19. Sphaeropteris elongata: a, portion of pinna. Sphaeropteris quindiuensis: b, portion of fertile segment; c, portion of petiole scale, (a from Soukup 1651, F, b from Soukup 2336, F, c from Plowman 6062, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 113 Sphaeropteris (Cyatheaceae) with a revision of the neotropical exindusiate species. Bot. Jahrb. Syst, 98: 176-198. — . 1978. Sphaeropteris (Cyatheaceae), the systematics of the group of Sphaeropteris hir- suta. Mem. New York Bot. Gard., 29: 2-22. Key to Species of Sphaeropteris a. Petiole scales without a dark apical seta and without dark marginal setae, rarely the margins ciliate toward the apex [subg. Sclephropteris Windisch] b b. Veins free c c. Sori exindusiate 1 . S. aterrima c. Sori indusiate, the indusium sometimes inconspicuous d d. Sori with a small, hemitelioid (flabellate) indusium e e. Costa and costules with whitish, fimbriate or highly dissected scales abaxially 2. S. rufescens e. Costa and costules lacking scales or with scales various, but not whitish, fimbriate, or highly dissected 3. S. macrosora Sori with a sphaeropteroid indusium 5. S. atahuallpa b. Veins regularly anastomosing, especially along the costa, sori with a hemitelioid indusium .... 4. S. bradei Petiole scales with a dark apical seta and dark marginal setae [subg. Sphaeropteris] f f. Sori exindusiate 6. S. elongata f. Sori with a sphaeropteroid indusium 7. S. quindiuensis d. 1 . Sphaeropteris aterrima (Hooker) Tryon, Contr. Gray Herb. 200:20. 1970. Ahophila aterrima Hooker, Syn. fil. 38. 1866. TYPE: Peru, (San Martin), Tarapoto, Spruce 4713 (ho- lotype, K!; isotypes, P!, us!). Cyathea aterrima (Hooker) Domin, Pterid. 262. 1 929. Petiole without spines or sparingly short-acu- leate, with very narrow scales intergrading to tri- chomes, these rather dense, especially toward the base of the petiole, straw-colored to light brown, lacking a dark apical seta and dark marginal setae, sometimes ciliate. Lamina 1-2-pinnate-pinnatifid, pinnules long-pubescent and sometimes with a few flattish scales abaxially, veins free. Sori with pa- raphyses about as long as the sporangia, exindu- siate. This species has been collected only in San Mar- tin at mid-elevations. Colombia and Peru. Sphaeropteris aterrima is notable for its petiole indument which consists of scales intergrading to trichomes. San Martin: Monte Morro de Moyobamba, Stiibel /77J(B, NY). 2. Sphaeropteris rufescens (Kuhn) Windisch, Bradea 1: 372. 1973. Hemitelia rufescens Kuhn, Linnaea 36: 159. 1869. TYPE: Peru, (San Martin), Monte Guairapurima, Tarapoto, Spruce 4727 (holotype, B!; isotype, K.; photo, GH of K). Cyathea rufescens (Kuhn) Domin, Pterid. 264. 1 929. Petiole muricate, pubescent, scales brown with lighter margins to wholly light colored, the edges toothed. Lamina 2-pinnate-pinnatifid, with tri- chomes, bullate scales, and long whitish, fimbriate or highly dissected scales abaxially, veins free. Sori usually at the fork of a vein, paraphyses longer than the sporangia, indusium hemitelioid (flabel- late). Known only from the type collection from San Martin. Endemic to Peru. The whitish fimbriate scales beneath and the small indusium make this a distinctive species. 3. Sphaeropteris macrosora (Baker) Windisch, Bradea 1: 372. 1973. 114 FIELDIANA: BOTANY Alsophila macrosora Baker, Timehri 5: 211. 1886. TYPE: Venezuela, (Bolivar), Ml. Roraima, 1m Thurn 87 (holotype, K!; isotype, us!). Cyathea macrosora (Baker) Domin, Pterid. 263. 1929. Petiole glabrous to sparingly pubescent, nearly smooth to muricate with the scales whitish to brownish and with lighter margins, these some- times broad, or most of the scales wholly whitish, lacking a dark apical seta and dark marginal setae, sometimes ciliolate, especially apically. Lamina 2- pinnate-pinnatifid, the pinnules glabrous, pubes- cent and (or) slightly scaly abaxially, the scales various, but not whitish, fimbriate, or highly dis- sected. Sori with paraphyses longer than the spo- rangia, indusium hemitelioid (broadly flabellate to reduced). At 400-500 m, Pasco. Guyana and Brazil, west to Colombia and Peru. Sphaeropteris macrosora is represented in Peru by var. reginae Windisch which occurs throughout the range of the species. Variety macrosora is in British Guiana and Venezuela and var. vaupensis Windisch is in Colombia and Venezuela. Pasco: Prov. Oxapampa, Palcazu, Foster & d'Achille 10168 (F). 4. Sphaeropteris bradei Windisch, Bradea 1: 372. 1973. TYPE: Colombia, Vaupes, Cerro Mitu, Schultes, Raffauf& Soejarto 2422 9 (holotype, GH!). Petiole sparingly aculeate, with rather broad scales, these usually dense, especially toward the base of the petiole, shining, dark brown, lacking a dark apical seta and dark marginal setae, usually finely short-ciliate. Lamina 2-pinnate-pinnatifid, pinnules with scattered, short, often crispate tri- chomes and sometimes a few bullate scales abax- ially, veins regularly anastomosing, especially along the costa. Sori with paraphyses shorter than the sporangia, indusium hemitelioid, usually 2-lobed. Primary forests, 130-200 m, San Martin and Loreto. Colombia, Ecuador, and Peru. The regularly anastomosing veins of Sphaer- opteris bradei are unique among the American Cy- atheaceae, except for species of the genus Cnem- idaria. San Martin: Prov. Lamas. Caserio Bonilla, km. 75 of Tarapoto-Yurimaguas road, Knapp & Mallet 7141 (MO). Loreto: Mishuyacu, near Iquitos, Klug 1546 (F, NY, us). Estacion Biologica Callicebus. Rio Nanay, 2 horas rio urn ha de Iquitos, Vasque: el al. 649 (GH). Near mouth of Rio Napo, Croat 20194 (F, uc). 5. Sphaeropteris atahuallpa Tryon, Rhodora 74: 442. 1972. TYPE: Peru, Amazonas, Prov. Chachapoyas, Cerros de Calla Calla, above Balsas on road to Leimebamba, Hutchison & Wright 6922 (holotype, GH!; isotypes, MO, uc). Petiole sparingly very short-aculeate, with large, broad scales to 6 cm long, these dense well above the base of the petiole, brownish white, lacking a dark apical seta and dark marginal setae, sparsely and finely ciliate, especially toward the apex. Lam- ina 2-pinnate-pinnatifid to 2-pinnate-pinnatisect, pinnules with prominent whitish, flattish scales abaxially, veins free. Sori with paraphyses about as long as the sporangia, indusia sphaeropteroid. This species has been collected in Peru only at ca. 3000-3300 m in the Department of Ama- zonas. Ecuador and Peru. Sphaeropteris atahuallpa is a distinctive and at- tractive species. The croziers are completely en- veloped by large, whitish scales and these form a dense covering to the petiole. Some of the scales are up to 6 cm long and several cells thick at their base. Amazonas: Summit of Puma-Urcu, SE of Chachapoy- as, Wurdack 1153 (GH, us). Prov. Chachapoyas, Calla- Calla, Smith & Vasquez 5012 (GH). 6. Sphaeropteris elongata (Hooker) Tryon, Contr. Gray Herb. 200: 20. 1970. Figure 19a. Alsophila elongata Hooker, Sp. fit. 1 : 43. 1 844. TYPE: Colombia, Hartweg 1528 (holotype, K!; isotype, LD), not Cyathea elongata Karsten. Alsophila poeppigii Hooker, Sp. fil. 1 : 43. 1 844. TYPE: Peru, 1829, Poeppig (holotype, K!), a synonym of Sphaeropteris elongata by Windisch, Bot. Jahrb. Syst. 98: 189. 1977. Cyathea poeppigii (Hooker) Domin, Pterid. 263. 1929. Petiole aculeate, with rather narrow scales, these usually sparingly persistent, whitish to light brown or with a darker base, with a dark apical seta and dark marginal setae. Lamina 2-pinnate-pinnatifid TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 115 to 2-pinnate-pinnatisect, pinnules sparingly to def- initely pubescent and sometimes with a few flattish scales abaxially, veins free. Sori with persistent paraphyses mostly longer than the sporangia, ex- indusiate. In primary forests, rain forests, and on open hillsides, at 450-2000 m, Amazonas and Loreto, south to Puno. Costa Rica south to Bolivia; also in southeastern Brazil. Sphaeropteris elongata is the most widely dis- tributed species of the genus in America and con- sequently is a variable species. Alsophila poeppigii was described on the basis of a plant with unusu- ally coriaceous leaves. Specimens without petiole scales may be similar to those of Trichipteris con- jugata, but that species has sessile rather than stalked pinnae. Amazonas: Prov. Bagua, Chiriaco to Puente Venezue- la. Barbour 4385 (MO, USM). San Martin: Prope Tara- poto. Spruce 4722 (GH). Loreto: Cerca a Pongo, entre Yurimaguas y Tarapoto, Ferreyra 17474 (GH, USM). Huanuco: Huamalies, Valle de Monzon, Weberbauer 3471 (USM). Chinchao, Ferreyra 16950 (GH, USM). Junin: La Merced, Soukup 1061 (F). Cerca a San Ramon, Cer- raie 2870 (GH, USM). 10 km SW of San Ramon, Tryon & Tryon 5445 (F, GH). Lcayali: 10 km NE of Aguaytia, Gentry el al. 41422 (F, MO, USM). Ayacucho: Prov. La Mar, Hacienda Santa Rosa, Dudley 11697 (GH). Ayna, between Huanta and Rio Apurimac, Killip & Smith 22716 (NY, us). Puno: Prov. Carabaya, Puente Inambari, Var- gas 18431 (GH). 7. Sphaeropteris quindiuensis (Karsten) Tryon, Contr. Gray Herb. 200: 20. 1970. Figure 19b- c. Cyathea Quindiuensis Karsten, Linnaea 28: 454. 1 857. TYPE: Colombia, (Tolima), between Rios Mag- dalena and Cauca, Karsten (not located). (Toli- ma), Paramo Quindio, Karsten (Herb. Mett. B! is authentic). Petiole without spines, with long, narrow scales, these dense, usually well above the base of the petiole, whitish to shining brown, with a dark api- cal seta and dark marginal setae. Lamina 2-pin- nate-pinnatifid to 2-pinnate-pinnatisect, pinnules with a few to many bullate scales, especially on the costules of fertile segments, and often some trichomes abaxially. Sori with paraphyses about as long as the sporangia, indusium sphaeropteroid. On steep mountain slopes, in rain forests and cloud forests, 1700-2500 m, Amazonas south to Cuzco. Colombia south to Bolivia. This species is unique among those of Peru in having some of the scales at the top of the crozier with the marginal teeth retrorse at the scale apex but changing to antrorse below. Amazonas: Prov. Bagua, Cordillera Colin, SE of La Peca, Barbour 3749 (MO). Huanuco: Prov. Leoncio Pra- do, Cordillera Azul, Gentry et al. 29573 (GH). Playapam- pa, Macbride 4856 (F). Pasco: Oxapampa (as Junin), Sou- kup 2336 (F, GH). Prov. Oxapampa, road Oxapampa- Paucartambo, Smith & Pretel 1643 (F, MO). Junin: Prov. Chanchamayo, ca. 26 km S of San Ramon, Smith & Palacios 2649 (F, MO). Cuzco: Prov. La Convention, Cor- dillera Vilcabamba, Dudley 11313 (GH). II. Alsophila Alsophila R. Br., Prodr. 158. 1810. TYPE: Also- phila australis R. Br. Figure 20. Terrestrial. Stem stout, erect and arborescent or very rarely short and decumbent or climbing, lack- ing spines or these rarely present. Leaves mono- morphic or sometimes dimorphic, with scales, es- pecially on the crozier and petiole base, that are marginate and with a usually dark apical seta. Pet- iole lacking spines or with corticinate spines. Lam- ina entire to 4-pinnate, veins free. Sori round, often at the fork of a vein, exindusiate or with a hem- itelioid to sphaeropteroid indusium. Spores tet- rahedral-globose, trilete, prominently ridged. Alsophila is a pantropical genus of ca. 230 species with 1 3 of them in America, mostly in the Greater Antilles, and a single one in Peru. In the treatment of Conant (1983), the first 13 species are Alsophila and the others are treated here in the next genus, Nephelea. Reference CONANT, D. S. 1983. A revision of the genus Alsophila (Cyatheaceae) in the Americas. J. Ar- nold Arbor., 64: 333-382. 1 . Alsophila engelii Tryon, Contr. Gray Herb. 200: 29. 1970, nom. nov. for Cyathea elongata 116 FIELDIANA: BOTANY 4 cm 3 mm FIG. 20. Alsophila engelii: a, pinna; b, ultimate segment and son; c, apex of petiole scale. (From Dudley 11944, OH.) Karsten, not Alsophila elongata Hooker. Fig- Petiole smooth to tuberculate, brown, bearing ure 20. mostly blackish to light brown scales with 1 apical seta. Lamina 2-pinnate-pinnatifid, gradually ta- pering to the apex, rachis light brown to brown, Cyathea elongata Karsten, Fl. Columb. 2: 1 59. 1 869. with usua"y numerous> sma11 dissected scales, pin- TYPE: Venezuela, (Merida), Merida, Engel 138 nae sessile or nearly so. Indusia deeply cyathiform (holotype, B!). to sphaeropteroid, glabrous. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 117 Montane rain forests, 1 700-1 900 m, Amazonas, San Martin, and Ayacucho. Venezuela and Colombia, south to Peru. Amazonas: Prov. Bongara, Shilla, Young & Eisenberg 442 (uc). San Martin: Prov. Rioja, Pedro Ruiz-Moy- obamba, D. Smith 4454 (GH). Ayacucho: Prov. La Mar, along Inca trail between Huanhuachayo and Punccu, Cordillera Central, Dudley 11944 (GH). Comments Alsophila paucifolia Baker of Colombia and Ec- uador may be found in Peru. It differs from A. engelii in having a 1-pinnate-pinnatifid lamina. Al- sophila capensis (L. f.) John Sm. ssp. polypodioides (Sw.) Conant of southeastern Brazil may also occur in Peru. It is characterized by the highly dissected aphlebiae at the base of the petiole. Terrestrial. Stem stout, erect and arborescent, sometimes branched, with spines. Leaves mono- morphic, with scales, especially on the croziers and petiole base, that are marginate and with a dark apical seta. Petiole with squaminate spines. Lam- ina 1-3-pinnate-pinnatifid, veins free. Sori round, usually at the fork of a vein, exindusiate or with a hemitelioid to sphaeropteroid indusium. Spores tetrahedral-globose, trilete, with prominent short to long ridges. Nephelea is a tropical American genus of 17 species, with three in Peru. It ranges from southern Mexico and the West Indies, south to northern Argentina and southern Brazil, centering in the Greater Antilles where nine species occur. The squaminate spines on the petiole are dis- tinctive of Nephelea. They are blackish, have a very sharp tip, and at least on old leaves they may be broken off at their base. III. Nephelea Nephelea Tryon, Contr. Gray Herb. 200: 37.1 970. TYPE: Nephelea polystichoides (Christ) Tryon = (Cyathea polystichoides Christ). Figure 21. Reference GASTONY, G. J. 1973. A revision of the fern genus Nephelea. Contr. Gray Herb., 203: 81- 148. Key to Species of Nephelea a. Veins with stellate trichomes or squamules abaxially b b. Indusium glabrous; pinna-rachises not green-alate between the more distal, sessile pinnules; petiole scales on the abaxial side lacking lateral setae 1 . N. erinacea b. Indusium with whitish trichomes; pinna-rachises usually (sometimes narrowly) green-alate be- tween the more distal, sessile pinnules; petiole scales on the abaxial side with dark lateral setae 2. N. cuspidata a. Veins lacking indument abaxially, or present and not stellate; pinna-rachises green-alate between the more distal, sessile pinnules; petiole scales lacking lateral setae, indusium with trichomes .3. N. incana 1 . Nephelea erinacea (Karsten) Tryon, Contr. Gray Herb. 200: 40. 1970. Cyathea erinacea Karsten, Linnaea 28: 453. 1857. TYPE: Venezuela, (Merida), Merida, 2000 m, Karsten (holotype, not located; isotype, B!). Alsophila erinacea (Karsten) Conant, J. Arnold Arbor. 64: 371. 1983. Scales on the abaxial side of the petiole with a dark apical seta and without lateral setae. Lamina usually coriaceous, predominantly 2-pinnate-pin- natifid, usually abruptly reduced to a pinna-like apex, pinna-rachises not green-alate between the more distal sessile pinnules, veins with minute, whitish, stellate squamules abaxially. Indusia cy- athiform to urceolate or rarely subsphaeropteroid, glabrous to densely scaly. In forests at ca. 1200-2000 m, Amazonas to Huanuco and Pasco. Costa Rica south to Venezuela, Colombia, and Bolivia. 118 FIELDIANA: BOTANY FIG. 2 1 . Nephelea cuspidata: a. basal portion of pinna; b, expanding crozier, c, ultimate segments and sori; d, apex of petiole scale, (a from Cuatrecasas 14843, Colombia, F, b, d from Davis 1109, Bolivia, F, c from Rimachi 481, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 119 Nephelea erinacea var. purpurascens (Sodiro) Gastony has the indusium densely and persistently scaly, rather than glabrous to sparsely scaly as in var. erinacea. It is presently known only from a few volcanoes in Ecuador. Amazonas: Prov. Bongara, 5 km N of N end of lake Pomacocha, on road to Rioja, Hutchison & Wright 6803 (GH, uc, USM). Rio Cenepa, 10 km E of Huampami, Berlin 219 (F, MO). Huanuco: Macbride 4843 (F, us). Pasco: Prov. Oxapampa, trail to summit of Cordillera Yanachaga, D. Smith et al 7817, 7851 (F, MO). Prov. Oxapampa, Rio San Alberto, Smith & Pretel 7942 (USM). 2. Nephelea cuspidate (Kunze) Tryon, Contr. Gray Herb. 200: 40. 1970. Figure 21. Cyathea cuspidata Kunze, Linnaea 9: 101. 1834. TYPE: Peru, (Loreto), Prov. Maynas, Feb. 1831, Poeppig diar. 2286 (holotype, LZ destroyed; iso- types, B!, P; photo, GH of p). Alsophila cuspidata (Kunze) Conant, J. Arnold Arbor. 64: 371. 1983. Scales on the abaxial side of the petiole with a dark apical seta and usually with numerous, small- er apical and lateral setae. Lamina usually papyr- aceous, predominantly 2-pinnate-pinnatifid, re- duced to a pinna-like apex, pinna-rachises green- alate between the more distal sessile pinnules, veins with stellate trichomes or stellate squamules abax- ially. Indusia rarely cyathiform, to urceolate to sphaeropteroid, glabrous or with stellate indu- ment. In low-elevation rain forests and less often in cloud forests, usually at 1 50-900 m, rarely to 2200 m. Amazonas and Loreto south to Puno. Nicaragua to Panama, northern South America, south to Bolivia and Paraguay. Nephelea cuspidata is the most widely distrib- uted species of the genus. Among Peruvian species it is distinctive in having the scales of the petiole, especially on the abaxial side, with several dark apical setae and dark marginal setae. Amazonas: Rio Cenepa, ca. 5 km E of Chavez Val- divia, Berlin 2066 (F, MO, uc). Loreto: Near mouth of Rio Santiago, above Pongo de Manseriche, Mexia 6197 (F, GH, MO, NY, uc, us). Alto Rio Itaya, LI. Williams 3288 (F, GH, NY, us). Huanuco: Tingo Maria (as San Martin), Allard 20618 (GH). Junin: E of Quimiri Bridge, near La Merced, Killip & Smith 23986 (F, NY, us). Ucay- ali: Prov. Coronel Portillo, Cordillera Azul, Young & Sullivan 705 (F, MO). Ayacucho: Estrella, between Huanta and Rio Apurimac, Killip & Smith 22654 (GH). Cuzco: Prov. La Convention, Rio Apurimac, above Boca del Tigre rapids, Davis et al. 1304 (F, GH). Madre de Dios: Prov. Tambopata, SSW of Puerto Maldonado, Barbour 5297 (F, MO). Puno: Prov. Carabaya, San Gaban, Puente Arica, Vargas 1891 9 (GH). 3. Nephelea incana (Karsten) Gastony, Contr. Gray Herb. 203: 137. 1973. Cyathea incana Karsten, Fl. Columb. 1: 75, t. 37. 1860. TYPE: Colombia, (Cundinamarca), Andes of Bogota, 2500 m, Lindig(nol located). Alsophila incana (Karsten) Conant, J. Arnold Arbor. 64: 371. 1983. Scales on the abaxial side of the petiole with a dark apical seta and without lateral setae, or rarely with an additional apical seta and lateral setae. Lamina rigidly papyraceous, 2-pinnate-pinnatifid, abruptly reduced to a pinna-like apex, pinna-rach- ises green-alate between the more distal sessile pinnules, veins glabrous abaxially, or with occa- sional trichomes. Indusia meniscoid to subsphaer- opteroid, pubescent with rather soft and often cris- pate trichomes or trichomoid processes. In primary forests at ca. 800-2400 m, Caja- marca and Amazonas south to Junin and Ucay- ali. Colombia south to northwestern Argentina. Cajamarca: Prov. Hualgayoc, Soukup 3810 (F). Prov. Cutervo, San Andres, Lopez et al. 6684 (GH, HUT). Ama- zonas: Laguna Pomacocha, NW of Jumbilla, Soukup 2560a (GH). Pasco: Prov. Oxapampa, Oxapampa, Leon 502 (USM). Junin: Esposto 670 (USM). Prov. Tarma, ca. 3 km SE of San Ramon, Iltis & Iltis 249 (GH, USM). Ucayali: Prov. Coronel Portillo, Cordillera Azul, Young & Sullivan 668 (F, MO). IV. Trichipteris Trichipteris Presl, Delic. prag. 1: 172. 1822. TYPE: Trichipteris excelsa Presl = Trichipteris cor- covadensis (Raddi) Copel. Figure 22. Terrestrial. Stem stout, erect and arborescent or rarely decumbent, lacking spines. Leaves mono- morphic, with scales especially on the croziers and petiole base that are marginate and lack a dark, apical seta. Petiole lacking spines or with corticin- ate spines. Lamina 1 -pinnate to 3-pinnate-pin- natifid, veins free. Sori round, often at the fork of a vein, exindusiate. Spores tetrahedral-globose, trilete, porate, verrucate, or finely echinate. 120 FIELDIANA: BOTANY FIG. 22. Trichipteris pubescens: a, apical portion of lamina; b, fertile ultimate segments; c, portion of segment, showing a sorus and a receptacle; d, portion of petiole scale, (a from Buchtien 5304, Bolivia, F, b-c from Killip & Smith 24871, F, d from Ollgaard et al. 9106, Ecuador, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 121 An American genus of 55 species, from Mexico References and the West Indies south to northern Argentina and southern Brazil. Fifteen species are in Peru, HARRINGTON, D. S. 1978. A revision of the genus many formerly treated in the genus Alsophila be- Trichipteris. Contr. Gray Herb., 208: 3-91. cause of the exindusiate sori. RIBA, R. 1969 (1967). Revision monografica del The name is sometimes incorrectly spelled as complejo Alsophila Swartziana Mart. (Cyathea- Trichopteris. The International Code of Botanical ceae). Ann. Inst. Biol. Univ. Nac. Auton. Mex- Nomenclature, Art. 73.8 confines the use of an ico, ser. bot., 38(1): 61-100. incorrect compounding form to epithets, thus ex- cluding generic names. The original spelling Tri- chipteris should be maintained. Key to Species of Trichipteris a. Lamina 1 -pinnate-pinnatifid b b. Pinnae sessile to short-stalked c c. Petiole 15-30 cm long; lamina not or not much reduced at the base 10. T. pubescens c. Petiole 3-6 cm long; lamina much reduced at the base 11. T. phegopteroides b. Pinnae, especially below the lamina apex, very long-stalked 4. T. latevagans a. Lamina 2-pinnate or more complex d d. Lamina mostly 3-pinnate-pinnatifid; pinnules abundantly short-pubescent adaxially, or petiole scales with a narrow, dark center stripe and broad white margins e e. Leaf ca. 1.5 m long, pinnules with many small, bullate scales abaxially, abundantly short- pubescent adaxially; petiole scales nearly concolorous, brown with very narrow, lighter margins 7. T. flava e. Leaf ca. 4 m long or more, pinnules with a few flattish scales abaxially, glabrate or with a few long trichomes adaxially; petiole scales with a narrow dark center stripe and broad whitish margins 9. T. serpens d. Lamina 2-pinnate-shallowly lobed to 2-pinnate-pinnatisect; if rarely partly 3-pinnate-pinnatifid then sparingly long-pubescent adaxially and the petiole scales with narrow, lighter margins . . . f f. Lamina abruptly reduced at the apex g g. Abaxial and lateral sides of the pinna-rachises lacking trichomes to (usually) sparingly pubescent with usually brown trichomes to 1 mm long h h. Fertile and sterile veins simple; or if rarely forked (T. procerd) then the ultimate lobes entire and the paraphyses much shorter than the sporangia i i. Petiole aculeate, pinnules usually sessile, basal veins end above a sinus, paraphyses much shorter than the sporangia 1 . T. procera i. Petiole muricate to tuberculate, pinnules stalked, basal veins connivent to a sinus, paraphyses longer than the sporangia 13. T. lechleri h. Fertile and sterile veins forked; ultimate lobes usually crenulate; paraphyses longer than the sporangia 5. T. nigra g. Abaxial and lateral sides of the pinna-rachises usually densely pubescent with usually whitish trichomes mostly 1.5-2 mm long 6. T. pilosissima f. Lamina gradually reduced at the apex j j. Pinna-rachises with few to several sharp spines 12. T. microdonta j. Pinna-rachises lacking spines k k. Pinnae and pinnules mostly long-stalked; ultimate segments subobtuse to acuminate or apiculate 3. T. kalbreyeri k. Pinnae mostly sessile to short-stalked; pinnules sessile to short-stalked or rarely long- stalked and then the ultimate segments obtuse 1 1. Pinnules with prominent, large, flattish scales abaxially; paramo and subparamo (elfin forest) 8. T. frigida 122 FIELDIANA: BOTANY Pinnules glabrate or pubescent abaxially or somewhat scaly with bullate or small flattish scales; montane forest and rain forest m m. Paraphyses shorter than the sporangia; petiole scales brown to dark brown, nearly concolorous; central and basal pinnae short- to long-stalked .... 2. T. nigripes m. Paraphyses longer than the sporangia n n. Petiole scales light brown to dark brown, with lighter margins; tertiary (ulti- mate) segments apically entire to coarsely toothed; central and basal pinnae short- to long-stalked (rarely sessile) 6. T. pilosissima n. Petiole scales dark brown to atropurpureous, usually with broad lighter mar- gins, or whitish to light brown and concolorous; tertiary (usually ultimate) segments apically crenulate or denticulate; central and basal pinnae sessile o o. Petiole with long trichomes, soon glabrous, the caducous trichomes leaving a smooth petiole surface 14. T. conjugate o. Petiole with long trichomes, these more or less persistent, when deciduous leaving a hard base and a scabrous petiole surface 15. T. tryonorum 1 . Trichipteris procera (Willd.) Tryon, Contr. Gray Herb. 200: 46. 1970. Polypodium procerum Willd., Sp. pi. ed. 4, 5: 206. 1810. TYPE: Brazil, Hoffmannsegg (holotype, B!, Herb. Willd. 19717). Polypodium pungens Willd., Sp. pi. ed. 4, 5: 206. 1810, a synonym of Trichipteris procera (Willd.) Tryon by Harrington, Contr. Gray Herb. 208: 23. 1978. Alsophila procera (Willd.) Desv., Mem. Soc. Linn. Paris 6: 319. 1827. Alsophila dombeyi Desv., Mem. Soc. Linn. Paris 6: 320. 1827. TYPE: Peru, (Huanuco), Cochero, Dombey (holotype, Herb. Desv. P!; isotype, P!). Alsophila infesta Kunze, Linnaea 9: 98. 1834. TYPE: Peru, (San Martin), Tocache, Rio Huallaga, Poep- pig (holotype, LZ destroyed; isotype, B!). Alsophila armigera Kunze, Linnaea 9: 98. 1 834. TYPE: Peru, (Huanuco), Ventanilla de Cassapi, Jul. 1829, Poeppig (holotype, LZ destroyed; isotypes, MO!, P!; photos, GH, us of P). Alsophila pycnocarpa Kunze, Linnaea 9: 97. 1834. TYPE: Peru, (Huanuco), Pampayacu, Jul. 1829, Poeppig (holotype, LZ destroyed; isotype, Poeppig 201, B!). Alsophila peruviana Klotzsch, Linnaea 20: 441. 1847. TYPE: Peru, (Junin), Tarma, Ruiz Herb. 66 (ho- lotype, B!). Alsophila pterorachis Baker, Syn. fil., ed. 2, 456. 1874. TYPE: Peru, (San Martin), Tarapoto, Spruce 47 17 (holotype, K.!). Alsophila floribunda Baker, Syn. fil., ed. 2, 458. 1874. TYPE: Peru, (San Martin), Cerro Campana, Spruce 4715 (holotype, K; isotype, P!). Cyathea pungens (Willd.) Domin, Pterid. 263. 1929. Cyathea willdenowiana Domin, Acta Bot. Bohemica 9: 171. 1930, nom. nov. for Polypodium procerum Willd., not Cyathea procera Brause. Trichipteris infesta (Kunze) Tryon, Contr. Gray Herb. 200: 45. 1970. Trichipteris dombeyi (Desv.) Barr., Contr. Gray Herb. 208: 27. 1978. Petiole lacking trichomes, the scales dark brown or with whitish margins. Lamina 2-pinnate-pin- natifid, abruptly reduced at the apex, central and basal pinnae short- to long-stalked, their pinnules sessile to short-stalked, abaxial and lateral sides of the pinna-rachises sparingly short-aculeate or lacking spines, glabrate or with scattered to rather numerous brownish trichomes. Pinnules obtuse to acuminate, glabrate or with some small scales and (or) trichomes abaxially, ultimate segments obtuse to subacute, fertile veins simple or rarely forked. Paraphyses much shorter than the sporangia. In original rain forests or cloud forests, often in secondary forests or in other partly disturbed vege- tation, 100-1500 m, Amazonas and San Martin south to Puno. West Indies; South America south to northern Brazil, Peru and Bolivia. Trichipteris dombeyi, although recognized by Barrington, is very close to T. procera and is here included within it. Trichipteris procera has an ex- tensive distribution and is quite variable as the many synonyms described from Peru imply. Amazonas: Prov. Bagua. Rio Maranon, above Cas- cadas de Mayasi, Wurdack 1825 (GH). San Martin: Cam- pana, near Tarapoto, Spruce 4323bis (GH). Prov. Rioja, Rioja, Ferreyra 18455 (USM). Loreto: Gamitanacocha, Rio Mazan, J. Schunke 269 (F, GH, NY, uc). Iquitos, Mexia 6497 (F, GH, MO, uc, us). Huanuco: Tingo Maria, Tryon & Tryon 5256 (F, GH). Prov. Huanuco, near con- fluence of Rio Cayumba with Huallaga, Mexia 8293 (F, GH, MO, NY, uc, us). Pasco: Prov. Oxapampa, Enenas, Soukup 6706 (GH). Prov. Oxapampa, Rio Palcazu, Ledn 702 (F). Junin: Above San Ramon, Killip & Smith 24541 (F, GH, NY, us). Ucayali: Prov. Coronel Portillo, Dist. Padre Abad (as Loreto), J. Schunke 9852 (MO). Cuzco: TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 123 Prov. Paucartambo, Kosnipata-Santa Ines, Vargas 11318 (GH). Madre de Dios: Pantiacolla, Gentry el al. 2735 1 (uc). Puno: Prov. Carabaya, Hacienda Palmares, Vargas 16145 (GH). Alsophila podophylla Baker, J. Bot. 19: 202. 1881. TYPE: Colombia, Antioquia, Kalbreyer 1375 (holotype, K; isotype, B!). Cyathea kalbreyeri (Baker) Domin, Pterid. 262. 1929. 2. Trichipteris nigripes (C. Chr.) Barr., Rhodora 78: 4. 1976. Alsophila nigripes C. Chr., Index fil. 45. 1905, nom. nov. for Alsophila melanopus Hooker (not Hassk.) and with the same type. Alsophila melanopus Hooker, Syn. fil. 37. 1866 (not Hassk., 1855). TYPE: Ecuador, Chimborazo, Spruce 5742 (holotype, K!; isotype, P!). Cyathea nigripes (C. Chr.) Domin, Pterid. 263. 1929. Petiole lacking trichomes, the scales brown to dark brown, nearly concolorous. Lamina 2-pin- nate-pinnatifid to 2-pinnate-pinnatisect, gradually reduced at the apex, central and basal pinnae short- to long-stalked, their pinnules sessile to nearly short-stalked, pinna-rachises lacking spines, the abaxial and lateral sides glabrous or with a few to sometimes many light brown to brown, rarely whitish trichomes. Pinnules obtuse to attenuate, glabrous or with short to rather long, mostly scat- tered trichomes abaxially, ultimate segments ob- tuse to acute, fertile veins simple or usually forked. Paraphyses shorter than the sporangia. In primary forests and somewhat disturbed for- ests, wet ravines, 200-800 m, Amazonas, San Martin and Loreto south to Madre de Dios. Costa Rica south to Peru. Recent collections have shown that Trichipteris nigripes var. brunnescens of Colombia and Ec- uador, with well-developed petiole spines, pubes- cent pinna-rachises, and sessile pinnules, is only one extreme within a spectrum of variation. Amazonas: Prov. Bagua, Dist. Senepa, Tillett 672-115 (GH). San Martin: Prov. Mariscal Caceres, Puente Palo Blanco, Rio Tocache, Plowman et al. 1 1354 (GH). Loreto: Above Pongo de Manseriche, Mexia6191 (F, GH). Huan- uco: Hills E of Tingo Maria, Croat 21153 (MO). Ucayali: Km 86 on Pucallpa-Tingo Maria road, Smith et al. 1 179 (GH). Cuzco: Prov. Paucartambo, Cosnipata valley, Wachler et al. 221 (F, GH). Madre de Dios: Prov. Tam- bopata. SSW of Puerto Maldonado, Barbour 4934 (MO). Petiole lacking trichomes, the scales atropur- pureous or with lighter margins. Lamina 2-pin- nate-pinnatifid, rather gradually reduced at the apex, pinnae and most of the pinnules long-stalked, pinna-rachises lacking spines, the abaxial and lat- eral sides glabrous or with some short, brownish trichomes. Pinnules acute to acuminate, glabrate abaxially, ultimate segments subobtuse, acute, acuminate or subapiculate, fertile veins simple or forked. Paraphyses shorter than the sporangia. In forests, 1200-1600 m, San Martin. Colombia south to Bolivia. San Martin: Zepelacio, near Moyobamba, Klug 3256 (F, GH). Monte Campana, near Tarapoto, Spruce 4330 (GH, NY). 4. Trichipteris latevagans (Baker) Tryon, Contr. Gray Herb. 200: 45. 1970. Alsophila latevagans Baker, J. Bot. 19: 203. 1881. TYPE: Colombia, Antioquia, Kalbreyer 1327 (holotype, K). Cyathea latevagans (Baker) Domin, Pterid. 262. 1929. Petiole lacking trichomes, the scales at the base brown, nearly concolorous. Lamina 1-pinnate- pinnatifid, gradually reduced at the apex, pinnae very long-stalked, especially the central and lower ones, their apex acute. Pinnules glabrous or some- what pubescent on the margins, obtuse to sub- acute, fertile veins forked. Paraphyses shorter than to longer than the sporangia. In pajonal vegetation, 2200 m, San Martin. Colombia and Peru. The very long-stalked pinnatifid pinnae and the atropurpureous petiole and rachis are distinctive features of this species. San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba, D. Smith 4799 (GH). 3. Trichipteris kalbreyeri (Baker) Tryon, Contr. Gray Herb. 200: 45. 1970. 5. Trichipteris nigra (Mart.) Tryon, Contr. Gray Herb. 200:46. 1970. Alsophila kalbreyeri Baker, Summary new ferns 9. 1892, nom. nov. for Alsophila podophylla Baker (not Hooker) and with the same type. Alsophila nigra Mart.. Icon. pi. crypt. 71.1 834. TYPE: Brazil, Prov. Rio Negro, Rio Japura, Martius (ho- lotype, not located: isotypes, B!, K). 124 FIELDIANA: BOTANY Alsophila lasiosora Kuhn, Linnaea 36: 157. 1869. TYPE: Peru, Spruce 4349 (holotype, B; isotypes, GH!, P!, us!). Alsophila tarapotensis Rosenst., Repert. Spec. Nov. Regni Veg. 7: 291. 1909. TYPE: Peru, (San Mar- tin), Tarapoto, Spruce 4349 (holotype, P!; iso- types, GH!, us!). Cyathea lasiosora (Kuhn) Domin, Pterid. 262. 1929. Cyathea primaeva Domin, Pterid. 263. 1929, nom. nov. for Alsophila nigra Mart., not Cyathea nigra Fourn. Alsophila killipii Maxon, Amer. Fern J. 32: 58. 1942. TYPE: Peru, Loreto, between Yurimaguas and Balsapuerto, Killip & Smith 28133 (holotype, us!; isotype, F!; numerous paratypes cited). Trichipteris lasiosora (Kuhn) Tryon, Contr. Gray Herb. 200: 45. 1970. Petiole lacking trichomes, the scales brown or dark brown with lighter margins. Lamina 2-pin- nate-pinnatifid to 2-pinnate-pinnatisect, abruptly reduced at the apex, pinnae short- to long-stalked, pinna-rachises lacking spines, the abaxial and lat- eral sides glabrate to somewhat pubescent, some- times with rather numerous, long, mostly brown to light brown trichomes. Pinnules acuminate to attenuate, with usually scattered, short to mod- erately long trichomes and some small scales abax- ially, ultimate segments obtuse to acute, fertile veins forked. Paraphyses longer than the sporan- gia. In wet primary forests, 100-1 100 m, Amazonas south to Madre de Dios. Dudley 13532, from Huanuco, is a juvenile plant, probably of Trichip- teris nigra; it was collected in elfin forest at 1850 m. Venezuela and northern Brazil, Colombia south to Bolivia. The few collections with long trichomes on the abaxial and lateral sides of the pinna-rachises and the abaxial side of the segments suggest that this species may not be wholly distinct from Trichip- teris pilosissima. Amazonas: Prov. Bagua, Chiriaco to Puente Venezue- la, Barbour 4465 (MO). San Martin: Prov. Mariscal Cac- eres, Tocache Nuevo, J. Schunke V. 4781 (F, MO, us), 5644 (GH). Loreto: Sierra del Pongo, Mexia 6270 (GH, uc). Prov. Maynas, Mishana, Rio Nanay, Gentry et al. 21109 (F, MO). Huanuco: Tingo Maria (as San Martin), Allard 20607 (GH, us). Pasco: Prov. Oxapampa, Valle del Palcazu, Ledn 703 (GH). Prov. Oxapampa, Rio Is- cozacin, Gentry et al. 41911 (USM). Junin: Santa Rosa, Pichis Trail, Killip & Smith 26169 (GH, NY). Ucayali: La Divisoria (as Huanuco), Gentry et al. 18874 (USM). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10703 (F, GH). 6. Trichipteris pilosissima (Baker) Barr., Contr. Gray Herb. 208: 76. 1978. Alsophila pilosissima Baker, Syn. fil., ed. 2, 457. 1874. TYPE: Peru, (San Martin), Mt. Campana, Spruce 4322 (holotype, K!; isotype, BR; photos, GH of K, GH, us of BR). Cyathea pilosissima (Baker) Domin, Pterid. 262. 1929. Petiole sometimes with abundant short to mod- erately long trichomes, the scales light brown to dark brown with lighter, sometimes nearly whit- ish, margins. Lamina 2-pinnate-pinnatifid to 2- pinnate-pinnatisect, gradually to abruptly reduced at the apex, central and basal pinnae short- to long- stalked, rarely sessile, their pinnules, especially to- ward the base, sessile to usually short-stalked, pin- na-rachises lacking spines, the abaxial and lateral sides with abundant long, mostly whitish tri- chomes. Pinnules obtuse to attenuate, with long, mostly whitish trichomes abaxially, ultimate seg- ments obtuse to subacute, the apex entire to coarsely toothed, fertile veins simple or usually forked. Paraphyses longer than the sporangia. In primary and secondary forests, rarely in sea- sonally inundated forests, 50-800 m, Tumbes, Amazonas, and Loreto, south to Cuzco and Madre de Dios. Panama south to Peru. This species is close to Trichipteris nigra, and their relationship requires further study. At the present time it is maintained as a distinct species, although there seems to be intergradation with T. nigra. Tumbes: Cerro Alegria, Canchaya 5169 (GH). Ama- zonas: Prov. Bagua, Quebrada Chuivi, Wurdack 1927 (GH). San Martin: Prov. Mariscal Caceres, Tocache, Plowman & Kennedy 5790 (F, GH). Loreto: Prov. May- nas, Mishana, Ldpez et al. 8677 (HUT). Prov. Maynas, Rio Ampiyacu, Plowman et al. 6601 (F, GH, us). Huan- uco: E of Tingo Maria, Croat 21153 (MO, uc). Pasco: Prov. Oxapampa, near confluence of Rio Palcazu and Rio Iscozacin, Smith & Franzen 1921 (F, MO). Junin: Near La Merced, Killip & Smith 23922 (F). Ucayali: Vicinity of Aguaytia (as Loreto), Croat 20952 (F, MO, uc). Cuzco: Prov. Paucartambo, Cosnipata valley, Wachterel al. 156 (F, GH). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10701 (F, GH). 7. Trichipteris flava Tryon, sp. nov. Petiolus squamis brunneis vel atrobrunneis. Lamina ca. 1 m longa tripinnata vel triptnnato-pinnatifida. Pin- nulae acutae supra breviter pubescentes subter squa- mulis multis pallide brunneis. Soris deest. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 125 Petiole lacking trichomes, the scales brown, con- colorous or with very narrow lighter margins. Lamina 3-pinnate-pinnatifid nearly throughout, gradually reduced at the apex, pinnae moderately long-stalked, pinna-rachises lacking spines, the abaxial and lateral sides with scattered, short to long trichomes. Pinnules short-stalked, acute, with many small, light brown, bullatc scales abaxially, ultimate segments mostly acute, many falcate. Sori absent. HOLOTYPE— Peru, Dept. Huanuco, above Chin- chao, Tingo Maria to Chinchao, 1 Aug. 1965, D. Soejarto 1432 (GH 2 sheets). Only known from the type collection from an exposed hillside at 2500 m in Huanuco. Endemic to Peru. The stem is reported on the label as slender and ca. 75 cm tall, the leaves ca. 1.5 m long, and the cut stem exuding yellow sap. Although the collec- tion is sterile, it very probably represents a species of Trichipteris. The lamina complexity and the abundant short-pubescence on the upper surface of the segments amply distinguish the species. 8. Trichipteris frigida (Karsten) Tryon, Contr. Gray Herb. 200:45. 1970. Alsophila frigida Karsten, Fl. Columb. 1 : 61, t. 30. 1859. TYPE: Colombia, (Cundinamarca), Andes of Bogota, 2600 m, Karsten (holotype, not locat- ed; isotype, B!). Cyathea frigida (Karsten) Domin, Pterid. 262. 1929. Petiole lacking trichomes, the scales light to dark brown usually with rather narrow lighter margins. Lamina 2-pinnate-pinnatifid, rarely partly 3-pin- nate-pinnatifid, gradually reduced at the apex, pin- nae sessile to long-stalked, pinna-rachises lacking spines, the abaxial and lateral sides with large flat- tish scales or their hard bases, with scattered brown to light brown trichomes or often with dense, ca- ducous, slender, matted, whitish trichomes or strongly dissected scales. Pinnules sessile to short- stalked, obtuse to rarely attenuate, with promi- nent, large flattish scales abaxially, ultimate seg- ments obtuse to subacute, fertile veins forked. Pa- raphyses shorter than the sporangia or of the same length. Growing in moist scrub forests, 2500-3500 m, Amazonas south to Junin. Venezuela and Colombia south to Peru. Trichipteris frigida typically grows at higher al- titudes than other species of the genus. Amazonas: Prov. Chachapoyas, Molinopampa-Dios- an pass, Wurdack 1654 (F, GH, us). Huanuco: Cerros al sudoestede Monzon, Weberbauer 3389(\jstA). Huanuco- Tingo Maria road, near Carpish, Gentry & Smith 44863 (F, MO). Pasco: Prov. Oxapampa, Santa Barbara, D. Smith 8172 (F, MO). San Gotardo, van der Werff et al. 8562 (MO, uc). Junin: Huacapistana, Weberbauer2272(?, USM). 9. Trichipteris serpens Tryon, sp. nov. Petiolus deest. Pedum circinatum sine trichomata, squamis centralibus fuscatis angustis marginatis late niv- eis. Lamina ca. 3-4 m longa tripinnato-pinnatifida pinnis pinnulisque longe petiolulatis. Pinnulae acuminatae sub- ter squamis paucis brunneis complanatis. Venae fertiles furcatae. Paraphyses longiores quam sporangia. Petiole lacking trichomes, the scales with a dark, narrow central stripe and broad whitish margins (characters from croziers). Lamina 3-pinnate-pin- natifid nearly throughout, apex absent, pinnae long- to very long-stalked, pinna-rachises lacking spines, the abaxial and lateral sides somewhat appressed pubescent and scaly. Pinnules short- to long- stalked, acuminate, glabrate or with a few large, flattish, brown scales abaxially, fertile veins forked. Paraphyses longer than the sporangia. TYPE— Peru, Dept. Cuzco, Prov. La Conven- tion, Cordillera Vilcabamba, 10 July 1968, T. R. Dudley 10949 (holotype, NA 2 sheets; isotype, GH 2 sheets). Known only from the type collection in elfin forest at 2900 m in Cuzco. Endemic to Peru. The leaves of this new species are unusual: they are 4-6 m long, or more, and trail or are scandent over low vegetation. The scales of the croziers, which will in part become petiole scales, are quite different from those of the related Trichipteris fri- gida. 10. Trichipteris pubescens (Baker) Tryon, Contr. Gray Herb. 200: 46. 1970. Figure 22. Alsophila pubescens Baker, Syn. fil. 449. 1868. TYPE: Peru, (San Martin), Mt. Guayrapurima, Tara- poto, Spruce 4712 (holotype, K; isotypes, OH!, NY!, P!). Cyathea bipinnatifida (Baker) Domin, Pterid. 262. 1929. 126 FIELDIANA: BOTANY Cyathea pubens Domin, Pterid. 263. 1929, nom. nov. for Alsophila pubescens Baker, not Cyathea pu- bescens Kuhn. Petiole lacking spines, the scales light brown with lighter margins. Lamina 1-pinnate-pinnati- fid, gradually reduced at the apex, pinnae sessile to short-stalked, with abundant, short to moder- ately long trichomes abaxially or rarely these few, ultimate segments obtuse, fertile veins simple or forked. Paraphyses as long as the sporangia. Growing at 350-1750 m, in San Martin and (probably) Puno. Endemic to Peru. This species has a small leafless than 1 m long and a very short petiole ca. 3-6 cm long; the basal pinnae are much reduced. San Martin: Mt. Guayrapurima, near Tarapoto, Spruce 4028 (GH, NY). Prov. Riqja, Pedro Ruiz-Moyobamba, D. Smith 4452 (GH). Puno(?): Lechler (F). In montane rain forests, cloud forests, or ceja de la montana, 850-2200 m, Amazonas south to Puno. Guyana, Venezuela, and Colombia south to Bo- livia. This species and the next are the only ones in Peru with a 1-pinnate-pinnatifid lamina. Trichip- teris pubescens is widely distributed and, although variable, is characterized by the features men- tioned in the key. Amazonas: Prov. Bagua. La Peca, Barbour 2764 (MO, uc), 3987 (F, MO). San Martin: Prov. Rioja, Pedro Ruiz- Moyobamba, D. Smith 4798 (GH). Huanuco: Cerros del Sira, Rio Llullapichis watershed, Dudley 13371, 13413, 13538 (GH). Pasco: Prov. Oxapampa, Palcazu, D. Smith 8553 (F, MO). Junin: Above San Ramon, Killip & Smith 24871 (F, us). Ucayali: Cerca a La Divisoria (as Loreto), Ferreyra 1071 (GH, USM). Ayacucho: Prov. La Mar, be- tween Huanhuachayo and Punccu, Cordillera Central, Dudley 11941 (GH). Cuzco: Prov. La Convention, Rio Mapitunuari, Dudley 10159B (GH). Puno: San Gaban, Lechler 2190 (K). Prov. Sandia, San Juan del Oro, Fer- reyra 16678, 16701 (GH, USM), 16704 (GH). 11. Trichipteris phegopteroides (Hooker) Tryon, Contr. Gray Herb. 200: 46. 1970. Alsophila phegopteroides Hooker, Syn. fil. 32. 1865. TYPE: Peru, (San Martin), Tarapoto, Spruce 4020 (holotype, K; isotypes, P!, us!). Cyathea phegopteroides (Hooker) Domin, Pterid. 263. 1929. Petiole pubescent, also bearing dark brown scales with lighter brown margins. Lamina 1-pinnate- pinnatifid, gradually reduced at the apex, pinnae sessile, rachis scaly and densely pubescent with long trichomes. Pinnule-segments obtuse, pubes- cent adaxially and abaxially, fertile veins forked. Paraphyses shorter than the sporangia. 12. Trichipteris microdonta (Desv.) Tryon, Contr. Gray Herb. 200: 46. 1970. Polypodium microdontum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Na- turk. 5: 319. 1811. TYPE: "America australis" (holotype, Herb. Desv. P!; photos, GH, us). Alsophila microdonta (Desv.) Desv., Mem. Soc. Linn. Paris 6: 319. 1827. Cyathea microdonta (Desv.) Domin, Pterid. 263. 1929. Petiole lacking trichomes, the scales light brown to dark brown, concolorous. Lamina 2-pinnate- pinnatifid to 2-pinnate-pinnatisect, gradually re- duced at the apex central and basal pinnae short- to long-stalked, their pinnules sessile to short- stalked, pinna-rachises with few to several sharp spines, their abaxial and lateral sides glabrate or with usually numerous short to moderately long trichomes. Pinnules acute to attenuate, with few to usually many, short to usually long trichomes abaxially, ultimate segments obtuse to acute, fer- tile veins forked. Paraphyses longer than the spo- rangia or the same length. In primary forests or disturbed forests, 100-750 m, San Martin, Loreto, and Cuzco. Mexico and the Greater Antilles, south to Peru and Brazil. The sharp spines on the pinna-rachises are a distinctive feature of Trichipteris microdonta. Rarely, they may be few and rather short, as in Luna 895. San Martin: Near Tarapoto, Spruce 4726 (NY). Loreto: Mishuyacu, near Iquitos, Killip & Smith 29876 (F, GH, NY). Quistococha, 12 km from Iquitos, Luna 895 (F). Rio Itaya. Iquitos, Tryon & Tryon 5172 (F, GH). Cuzco: Prov. Quispicanchi. Quince Mil, Vargas 15340 (GH). 13. Trichipteris lechleri (Mett.) Tryon, Contr. Gray Herb. 200:45. 1970. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 127 Alsophila lechleri Men., Fil. lechl. 2: 28. 1859. TYPE: Peru, (Puno), Tatanara, Lechler 2532 (holotype, Herb. Mett. B!; photo, GH). Alsophila w/e; Christ, Hedwigia 44: 367. 1905. TYPE: Peru, (Amazonas), Cerro Ponasa, Ule 6901 (ho- lotype, P?; isotypes, B!, L; photos, GH of B, us of L). Cyathea subtropica Domin, Pterid. 263. 1 929, nom. nov. for Alsophila lechleri Mett., not Cyathea lech- leri Mett. Cyathea ulei (Christ) Domin, Acta Hot. Bohemica 9: 168. 1930. Petiole lacking trichomes, the scales dark brown to atropurpureous with rather broad, lighter mar- gins. Lamina 2-pinnate-shallowly lobed to 2-pin- nate-pinnatifid, abruptly reduced at the apex, cen- tral and basal pinnae long-stalked, their pinnules, especially toward the base, long-stalked, pinna- rachises lacking spines, abaxial and lateral sides glabrate. Pinnules acuminate to attenuate, glabrate abaxially, lobes or ultimate segments more or less obtuse, fertile veins simple. Paraphyses longer than the sporangia. Wet forests and cloud forests, 500-1300 m, Amazonas, Huanuco, Junin, and Puno. Venezuela and Colombia south to Bolivia. Trichipteris lechleri usually has the pinnules rather shallowly pinnatifid or lobed and the basal veins on an ultimate segment are connivent to a sinus. Huanuco: Cerros del Sira, Rio Llullapichis watershed, Dudley 13047, 13214, 13220 (GH). Junin: Prov. Satipo, S of Chequitavo, D. Smith 5130 (F). 14. Trichipteris conjugata (Hooker) Tryon, Contr. Gray Herb. 200: 45. 1970. Alsophila conjugata Hooker, Syn. fil. 37. 1 866. TYPE: Ecuador, Chimborazo, Spruce 4745 as published, 5745 is correct (holotype, K; frag., us!). Petiole with long trichomes, soon glabrous, the caducous trichomes leaving a smooth surface, the scales dark brown to atropurpureous with usually rather broad, light brown to whitish margins, or light brown to whitish and concolorous, the mar- gins slightly or not dark-denticulate apically. Lam- ina 2-pinnate-pinnatifid to 2-pinnate-pinnatisect, gradually reduced at the apex, pinnae sessile, pin- na-rachises lacking spines, the abaxial and lateral sides with usually abundant, long, whitish to brownish trichomes. Pinnules sessile, acuminate to mostly attenuate, with rather sparse to usually abundant long trichomes abaxially, ultimate seg- ments obtuse to acute, the apex denticulate, fertile veins forked. Paraphyses longer than the sporan- gia. In montane forests, 700-2000 m, Huanuco south to Cuzco. Venezuela and Colombia south to Bolivia. Trichipteris conjugata and the next species, T. tryonorum, are members of a group of some 1 3 species with dark denticulate margins on the pet- iole scales. The dark denticulate margins, how- ever, are often absent in the two Peruvian species. Huanuco: Prov. Leoncio Prado, Young & Sullivan 867 (F, MO). Prov. Huanuco, Yuracyaca, Ridoutt in 1 943 (GH, us, USM). Junin: Above San Ramon, Killip & Smith 24643 (F, GH, us). Ucayali: Prov. Coronel Portillo, La Divisoria, Aguilar 848 (F, GH). Ayacucho: Prov. La Mar, between Huanhuachayo and Punccu, Dudley 1 1943 (F, GH). Cuz- co: Prov. Urubamba, near town of Machu Picchu, Rio Urubamba, Tryon & Tryon 541 1 (GH, USM). Machu Pic- chu, Iltis et al. 1024 (GH, uc). 15. Trichipteris tryonorum (Riba) Tryon, Contr. Gray Herb. 200:46. 1970. Alsophila tryonorum Riba, Rhodora 69: 66. 1967. TYPE: Colombia, Cundinamarca, Cuesta "Fu- sugasuga," (Fusagasuga), Cuatrecasas 8036 (ho- lotype, us!). Petiole with long trichomes, these persistent or deciduous and then leaving a hard base and a sca- brous petiole surface, the scales dark brown to atropurpureous, usually with broad, lighter mar- gins, or whitish to light brown and concolorous, the margins slightly or not dark denticulate. Lam- ina 2-pinnate-pinnatifid to 2-pinnate-pinnatisect, gradually reduced at the apex, pinnae sessile, pin- na-rachises lacking spines, the abaxial and lateral sides with usually abundant long, whitish to brownish trichomes. Pinnules sessile, acute to usu- ally acuminate or attenuate, with usually abundant long trichomes and often some whitish bullate scales abaxially, ultimate segments acute to ob- tuse, the apex crenulate to coarsely denticulate, fertile veins forked. Paraphyses longer than the sporangia. Growing at 200-2400 m, Amazonas and Pasco. Venezuela, Colombia, south to northern Peru. This species is especially distinguished from Tri- chipteris conjugata by the persistent trichomes on the petiole, or their hard bases that provide a sca- 128 FIELDIANA: BOTANY brous surface when they fall off. Trichipteris con- jugata has caducous trichomes on the petiole that leave a smooth surface. Amazonas: Prov. Bagua, Cordillera de Colan, SE of La Peca, Barbour 3750 (MO). Pasco: Prov. Oxapampa, Rio Yamaquizu valley, D. Smith 4207 (OH). V. Cyathea Cyathea Sm., Mem. Acad. Roy. Sci. Turin 5: 416. 1 793. TYPE: Cyathea arborea (L.) Sm. (Poly- podium arboreum L.). Figure 23. Hemitelia R. Br., Prodr. 158. 1810. TYPE: Hemitelia multiflora (Sm.) Sprengel = Cyathea multiflora Sm. Terrestrial. Stem stout, erect and arborescent, rarely very slender and scandent or short, lacking spines. Leaves monomorphic to somewhat di- morphic, with scales, especially on the croziers and the petiole base, that are marginate and lack a dark apical seta. Petiole lacking spines or with corticin- ate spines. Lamina 1-4-pinnate, veins free or rare- ly anastomosing without included veinlets. Sori round, often at the fork of a vein, with a hemi- telioid to sphaeropteroid indusium. Spores tetra- hedral-globose, trilete, nearly smooth, porate, ver- rucate or minutely echinate. Cyathea is a tropical American genus of ca. 40 species, with 14 of them in Peru. The genus is sometimes treated in a broad sense to include nearly all of the family Cyatheaceae. Here it is restricted to a natural American group. Reference TRYON, R. 1976. A revision of the genus Cy- athea. Contr. Gray Herb., 206: 19-98. Key to Species of Cyathea a. Indusia hemitelioid b b. Sori costal 3. C. vilhelmii b. Sori medial to submarginal c c. Basal veins straight or nearly so, ending above a sinus d d. Ultimate segments with simple fertile veins and usually entire (rarely coarsely dentate), or with forked fertile veins and coarsely dentate 1 . C. multiflora d. Ultimate segments with forked fertile veins, entire or finely dentate to strongly crenate . . 2. C. andina c. Basal veins curved, connivent to a sinus, sometimes forming costal areolae ... 4. C. petiolata a. Indusia sphaeropteroid, sometimes evanescent and then the apical umbo or a basal remnant is usually present e e. Petiole scales either whitish to light brownish and concolorous or nearly so, or discordantly bicolorous with whitish to brownish margins; petiole scurf whitish to light brown, or rarely brown at the base of the petiole f f. Lamina 2-pinnate-pinnatifid or 3-pinnate only at the base of the pinnae g g. Large scales of the petiole scurf flattish h h. Pinnae and pinnules mostly long-stalked 5. C. divergens h. Pinnae and pinnules sessile or nearly so 7. C. ruiziana g. Large scales of the petiole scurf mostly, or many of them, crested 6. C. pallescens f. Lamina 3-4-pinnate nearly throughout i i. Tertiary segments entire or basally 2-lobed; bullate scales on the abaxial side of the pinnule- rachis dark brown 8. C. microphylla i. Tertiary segments with up to usually 5 or more segments or lobes; bullate scales on the abaxial side of the pinnule-rachis light brown 9. C. multisegmenta e. Petiole scales either brown and nearly concolorous, or (sometimes narrowly) concordantly bico- lorous with light brown to brown margins; petiole scurf brown or absent j j. Lamina 2-pinnate-pinnatifid or 3-pinnate only at the base of central and basal pinnae; larger ultimate segments usually ca. 1 0 mm long k k. Petiole scales light brown to brown, concolorous to narrowly bicolorous; pinnules usually TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 129 with few or no scales abaxially, abundantly to moderately long- pubescent 10. C. delgadii k. Petiole scales with a reddish brown to usually dark reddish brown or atropurpureous body, or with dark areas or streaks, bicolorous with sometimes narrow margins 1 1. Petiole aculeate to abundantly, rarely sparsely, muricate, with abundant, sometimes caducous scurf 11. C. caracasana 1. Petiole nearly or quite smooth, or with some scattered tubercles, scurf absent or sparse m m. Pinnules sessile to short-stalked 12. C. lechleri m. Pinnules long-stalked 13. C. ebenina Lamina 3-pinnate nearly throughout; larger ultimate segments ca. 5 mm long 14. C. dudleyi 1. Cyathea multiflora Sm., Mem. Acad. Roy. Sci. (Turin) 5: 416. 1793. TYPE: "Amer. merid.," R. Shakespeare (holotype, Herb. Banks BM!; photos, GH, NY, us). Hemitelia multiflora (Sm.) Sprengel, Syst. veg. 4: 1 26. 1827. Petiole aculeate or less often sparingly aculeate, the scales light brown to dark brown, nearly con- colorous to usually, sometimes narrowly, concor- dantly bicolorous, scurf dense, often caducous, the large scales flattish. Lamina 2-pinnate-pinnatifid to 2-pinnate-pinnatisect, pinnules sessile to short- stalked, glabrate to usually slightly to prominently long-pubescent abaxially and usually with a few scales, ultimate segments entire or coarsely den- tate, fertile veins simple or forked. Sori medial to nearly submarginal, indusia hemitelioid. In dense forests, 100-800 m, Amazonas south to Cuzco and Madre de Dios. Belize south to Bolivia; northern Brazil. Although close to the next species, Cyathea mul- tiflora is evidently distinct. In addition to the char- acters mentioned in the key, the small indusium is entire. Amazonas: Laguna Pomacocha, NW of Jumbilla, Sou- kup 5260B (GH). Loreto: Gamitanacocha, Rio Mazan, J. Schunke 1 17 (GH, MO, NY, uc, USM). San Antonio, Rio Itaya, Killip & Smith 29379 (F, NY, us). Junin: E of Quimiri Bridge, near La Merced, Killip & Smith 23979 (F, NY, us). Cuzco: Cerca de Atalaya, Vargas 16274 (GH). Entre Quince Mil y San Lorenzo, Vargas 11754 (GH). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10721 (F). 2. Cyathea andina (Karsten) Domin, Pterid. 263. 1929. Hemitelia andina Karsten, Linnaea 28: 452. 1856. TYPE: Colombia, "Santa Martha," 2500 m, Kar- sten (not located; authentic specimen, Karsten, Herb. Men. B!; photo, GH!). Petiole slightly muricate to usually sparingly aculeate, the scales light brown, or partly whitish, to dark brown and concolorous, or usually nar- rowly concordantly or discordantly bicolorous, scurf dense, often caducous, the large scales flat- tish. Lamina 2-pinnate-pinnatifid to 2-pinnate- pinnatisect, pinnules nearly sessile to rather long- stalked, glabrate, slightly scaly or pubescent abax- ially, fertile veins forked, rarely simple. Sori me- dial to submarginal, indusia hemitelioid. In dense forests and forest borders 20-1300 m, Loreto south to Puno. Greater Antilles, French Guiana west to Colom- bia and south to Bolivia. Cyathea andina is close to C. multiflora, and the two species have a similar range in the Andes. The indusium of C. andina is usually larger than in C. multiflora and it commonly splits at maturity into two segments. Loreto: Yanamona, Rio Amazonas above mouth of Rio Napo, Gentry et al. 36580 (MO). Pasco: Prov. Ox- apampa, Gran Pajonal, N of Chequitavo, D. Smith 5088 (GH). Junin: E of Quimiri Bridge, near La Merced, Killip & Smith 23995 (GH, NY, us). Prov. Tarma, Esposto 659 (USM). Ucayali: La Divisoria, Ferreyra 1694 (us). Prov. Coronel Portillo, Obeteni, Chrostowski 66-14 (uc). Madre de Dios: Prov. Tambopata, Rio Tambopata, Barbour 4775 (F, MO). Puno: Prov. Carabaya, Vargas 17536 (GH). 3. Cyathea vilhelmii Domin, Pterid. 264. 1929, nom. nov. for Hemitelia lechleri Mett., not Cyathea lechleri Mett. 130 FIELDIANA: BOTANY Hemitelia lechleri Mett., Fil. lechl. 2: 28. 1859. LEC- TOTYPE (designated by Tryon, Contr. Gray Herb. 206: 41. 1976): Peru, (Puno), Tatanara, Lechler2654 (holotype. Herb. Mett. B!; frag., GH!; lectoparatype, Peru, Tatanara, Lechler2650, Herb. Mett. B!). Petiole lacking, scales and scurf unknown. Lam- ina 3-pinnate-pinnatifid, pinnules long-stalked, glabrate abaxially, ultimate segments subentire to slightly toothed, fertile veins forked. Sori costal, indusia hemitelioid, small, nearly concealed by the sporangia. Only known from the two Lechler collections from Puno. Endemic to Peru. 4. Cyathea petiolata (Hooker) Tryon, Contr. Gray Herb. 206: 42. 1976. Hemitelia petiolata Hooker, Sp. fil. 1: 31, t. 26. 1844. TYPE: Isthmus of Panama, Dr. Sinclair (hole- type, K). Cyathea panamensis Domin, Pterid. 264. 1929, nom. super/1. Intended as a nom. nov. for Hemitelia petiolata Hooker, not John Sm., 1841, but the latter is invalid. Petiole nearly smooth, the scales brown to dark brown, uniformly so or in patches, discordantly bicolorous, with whitish to light brown margins, scurf rather sparse and caducous, large scales flat- tish when present. Lamina 2-pinnate-pinnatifid, pinnules long-stalked, glabrate abaxially, ultimate segments toothed toward the apex, fertile veins simple or forked. Sori submarginal, indusia hem- itelioid. Montane forests and cloud forests, to 1750 m, Huanuco and Cuzco. Panama south to Peru. Unusual features of this species are the basal veins that are connivent to a sinus where they sometimes form costal areolae. Huanuco: Tingo Maria (as San Martin), Allard 22360 (GH) is probably a juvenile plant. Cuzco: Prov. La Con- vention, Cordillera de Vilcabamba, Dudley 10616 (GH). 5. Cyathea divergens var. divergens Cyathea divergens Kunze, Linnaea 9: 1 00. 1 834. TYPE: Peru, (Huanuco), Pampayacu, Jul. 1 829, Poeppig (Diar. 1163) (holotype, LZ destroyed; authentic specimen, Poeppig 2 19 (Diar. 1152), B!, P!; photo. GH of P). Cyathea equestris Kunze, Linnaea 9: 100. 1834. TYPE: Peru, (Huanuco), Cerro de Cristobal, Pampayacu, 1829, Poeppig (holotype. LZ destroyed; frag., K). Petiole short-aculeate to aculeate, the scales very light brown to dark brown, predominantly dis- cordantly bicolorous with lighter margins, scurf dense, usually persistent, the large scales Hattish. Lamina 2-pinnate-pinnatifid to 2-pinnate-pinna- tisect, pinnules short- to long-stalked, glabrous, slightly pubescent or with a few scales abaxially, ultimate segments subentire, slightly denticulate or shallowly crenate, fertile veins forked. Sori cos- tal or nearly so, or rarely medial, indusia sphaer- opteroid. Montane forests and cloud forests, 1 200-2000 m, San Martin south to Cuzco. The var. divergens occurs from Costa Rica and Panama; Guyana west to Colombia and south to Peru. Two varieties are recognized by Tryon (1976). Only var. divergens occurs in South America; the other, var. tuerckheimii, is confined to Mexico and Guatemala. In Ecuador and Peru, var. divergens usually has very long pinnule stalks. San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba, Smith & Vdsquez 4637 (GH). Huanuco: Pampayacu, Ka- nehira 120 (GH). Cushi, Macbride 4819(r). Junin: Prov. Satipo, Gran Pajonal, S of Chequitavo, D. Smith 5123 (GH). Cuzco: Sahuayaco, Chaupiorcco, Biies 840 (us). 6. Cyathea pallescens (Sodiro) Domin, Pterid. 263. 1929. Alsophila pallescens Sodiro, Recens. crypt, vase. Quit. 20. 1883. TYPE: Ecuador, Bosques de Nanegal, Sodiro (holotype, not located; authentic speci- men, Sodiro, P!; photo, GH). Petiole muricate to aculeate, the scales whitish and concolorous to brown or rarely dark brown and discordantly bicolorous, with lighter margins, scurf dense, mostly persistent, the large scales crested or rarely absent. Lamina 2-pinnate-pin- natifid to 2-pinnate-pinnatisect, pinnules sessile or nearly so, glabrate to pubescent and (or) scaly abaxially, ultimate segments subentire to crenate or rarely lobed, fertile veins simple or forked. Sori costal to nearly medial, indusia sphaeropteroid. TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 131 On steep slopes of ceja de la montana, in cloud forests and in grasslands, less often in secondary growth, 1 500-3300 m, Amazonas, Pasco, Ucayali, and Cuzco. Colombia south to Bolivia. Although this is a variable species, it is char- acterized by the crested large scales of the petiole scurf. These are expanded into two or more planes and usually dissected at the apex. The indusium is rarely subsphaeropteroid or less developed. Amazonas: Prov. Bagua, Cordillera Colan, SE of La Peca, Barbour 3590 (F, MO), 3722 (MO). Pasco: Prov. Oxapampa, Rio San Alberto, Foster et al. 10285 (F, GH). Ucayali: Prov. Coronel Portillo, Cordillera Azul, Young & Sullivan 762 (F, MO). Cuzco: Prov. La Convention, antes de San Luis, Vargas 22780 (GH). Prov. La Con- vention, Dudley 10950B, 11030 (GH), 11149 (GH, us). 7. Cyathea ruiziana Klotzsch, Linnaea 20: 439. 1847. TYPE: "In peruviae andium nemori- bus," Ruiz 72 (holotype, B!; isotype, us!). Petiole aculeate, the scales with a dark center, discordantly bicolorous with whitish margins, scurf dense, rather persistent, the large scales flattish. Lamina 2-pinnate-pinnatifid to 2-pinnate-pinna- tisect, pinnules sessile or nearly so, rather scaly abaxially, ultimate segments subentire, fertile veins forked. Sori medial, indusia sphaeropteroid. In montane forests, ca. 2000 m, Huanuco. Endemic to Peru. Cyathea ruiziana has the large scales of the pet- iole scurf unusually well-developed and 3-4 mm long: they are whitish and mostly concolorous. Huanuco: Huacachi, near Muna, Macbride 4135 (F, us). The Ruiz collection also bears the name of Pana- tahuas, formerly a province west of Huanuco. 8. Cyathea microphylla Melt., Fil. lechl. 1: 23, /. 3,f. 1-6. 1856. TYPE: Peru, (Puno), St. Ga- ban (Rio San Gaban), Lechler 2160 (LZ de- stroyed; authentic specimen, Lechler 2569, Herb. Mett. B!; duplicate, F! (frag.), P!; photo, GH of P). Petiole smooth, the scales whitish to light brown, broad, concolorous, scurf rather dense, small, more or less persistent. Lamina 3-pinnate to 3-pinnate- pinnatifid, pinnules sessile, scaly abaxially, ulti- mate segments entire, fertile veins simple. Sori subcostal, indusia sphaeropteroid. In thickets and forests, 2100-2800 m, Cuzco and Puno. Endemic to Peru. The type collection, Lechler 2160, is clearly a mixture. It is cited by Mettenius under Cyathea microphylla and also under Cyathea schanschin (= C. delgadii). There is a specimen at Paris of the collection correctly identified as the latter species. Cyathea microphylla is a rare species with small leaves to ca. 75 cm long. Cuzco: Prov. La Convention, Dudley 10943 (GH). Puno: Valle Grande, Sandia, Vargas 11858 (GH). 9. Cyathea multisegmenta Try on, Contr. Gray Herb. 206: 65. 1976. TYPE: Peru, Cuzco, Prov. La Convention, Dudley 11326 (holotype, NA!; isotypes, GH!, us!). Petiole nearly smooth, the scales whitish to light brown, broad, concolorous, scurf dense, persis- tent, the large scales flattish. Lamina 3-pinnate- pinnatifid to 4-pinnate, pinnules sessile, pubes- cent, and scaly abaxially, ultimate segments entire to lobed, fertile veins simple or forked. Sori sub- costal, indusia sphaeropteroid. In dense cloud forest, 1 800 m, Cuzco. Endemic to Peru. Known only from the type collection. The leaves are noted as 4 m or more long. 10. Cyathea delgadii Sternb., Vers. Fl. Vorwelt 1: 47, /. B. 1820. TYPE: Brazil, Goias, Gancho General Delgado, via ad Caldas Novas, Pohl (holotype, PRC; frag., GH!). Cyathea schanschin Mart., Icon. pi. crypt. 77, /. 54. 1834. TYPE: Brazil, Sao Paulo, Martius (holo- type, M?; isotype, BM!; photo, GH). Cyathea oligocarpa Kunze, Linnaea 9: 101. 1834. TYPE: Peru, (Huanuco), Pampayacu, Jul. 1829, Poeppig Diar. 1101 (holotype, LZ destroyed; is- otypes, Poeppig 218, B!, MO!, P!). Cyathea pilosa Baker, Syn. fil., ed. 2, 19. 1874. TYPE: Peru, (San Martin), Tarapoto, Spruce 4729 (ho- lotype, K; isotypes, BM!, GH!, P!; photos, GH, uc, us of K, GH of P). Petiole slightly muricate to aculeate, the scales light brown to brown, nearly concolorous or nar- 132 FIELDIANA: BOTANY rowly concordantly bicolorous, scurf sometimes dense, usually caducous, large scales flattish. Lam- ina 2-pinnate-pinnatifid to 3-pinnate at the base of the central and basal pinnae, pinnules sessile to short-stalked, abundantly to moderately long-pu- bescent abaxially, sometimes with few long tri- chomes or few to several concolorous scales, ul- timate segments entire to partly lobed, fertile veins forked or rarely simple. Sori costal or nearly so, indusia sphaeropteroid. In montane forests, primary or open forests, or cloud forests, 850-2880 m, Amazonas, San Mar- tin, Huanuco, Pasco, and Cuzco. Costa Rica and Panama; northern South Amer- ica south to Bolivia and southeastern Brazil. Cyathea delgadii is characterized by the long trichomes abaxially, which are usually abundant and the rather few, if any, scales. It is one of the most widely distributed species of Cyathea. Amazonas: Prov. Bagua, Cordillera Colan, SE of La Peca, Barbour 3612 (F, MO, USM). San Martin: Rioja, Soukup 5223 (GH). Huanuco: Tingo Maria, Tryon & Tryon 5219 (F, GH, us). Cerros del Sira, Rio Llullapichis wa- tershed, Dudley 13070 (GH). Pasco: Prov. Oxapampa, Yanachaga National Park, Leon 1014 (GH). Cuzco: Prov. La Convention, Dudley 10058, 10268 (GH). Prov. Uru- bamba, camino a Huinayhuayna, Chavez 3442 (MO). 1 1. Cyathea caracasana (KJotzsch) Domin, Pter- id. 262. 1929. Alsophila caracasana KJotzsch, Linnaea 1 8: 54 1 . 1 844. TYPE: Venezuela, (Dist. Federal), Caracas, Mor- itz 117 (holotype, not located; isotypes, GH!, P!). Petiole muricate to aculeate, the scales predom- inantly dark reddish brown to atropurpureous, concordantly, sometimes narrowly, bicolorous with lighter margins, scurf dense, persistent or ca- ducous, the large scales flattish. Lamina 2-pinnate- pinnatifid to 2-pinnate-pinnatisect, pinnules ses- sile to short-stalked or rarely long-stalked, gla- brous, pubescent and (or) scaly abaxially, some- times densely so, ultimate segments subentire to crenate, fertile veins forked, rarely simple. Sori costal to medial, indusia sphaeropteroid. The species occurs in the Greater Antilles; Costa Rica; and Venezuela and Colombia south to Bo- livia. Cyathea caracasana is the most variable species among the American Cyatheaceae. Five varieties were recognized by Tryon (1976) in order to fa- cilitate the study of this complex. In addition to the two varieties in Peru, var. maxonii is endemic to Costa Rica, var. caracasana is in the Greater Antilles and northern South America, and var. chimborazensis is in Venezuela, Colombia, and Ecuador. Key to Varieties a. Pinnules usually tapering to the apex from beyond the middle, abaxially usually with abundant scales and trichomes 1 la. var. boliviensis a. Pinnules, especially toward the base of the central and basal pinnae, tapering to the apex from the base, indument abaxially sparse or absent lib. var. meridensis 1 la. Cyathea caracasana var. boliviensis (Ro- senst.) Tryon, Contr. Gray Herb. 206: 77. 1976. Figure 23. Cyathea mexicana var. boliviensis Rosenst., Repert. Spec. Nov. Regni Veg. 25: 56. 1928. TYPE: Bo- livia, Hacienda Simaco, above Tipuani, Buchtien 5140 (holotype, not located; isotypes, F!, GH!, NY!, us!). In wooded ravines, mountain forests, and es- pecially in cloud forests, 1 300-2800 m, Amazonas south to Puno. Venezuela and Colombia south to Bolivia. Amazonas: Prov. Chachapoyas, entre Ingenio y Po- macocha, Ldpez et al. 4312 (GH, HUT). Prov. Chacha- poyas, Molinopampa-Diosan pass, Wurdack 1653 (F, GH, us). Huanuco: Within 5 km of Carpish, Tryon & Tryon 5326 (F, GH, NY, uc, us). Pasco: Prov. Oxapampa, Rio San Alberto valley. Smith & Pretel 8003 (F, MO). Oxapampa (as Junin), Soukup 2335 (F, GH). Ucayali: Prov. Coronel Portillo, La Divisoria, (as Loreto), Fer- reyra 1074 (GH, us), 7696 (us, USM). Huancavelica: Prov. Tayacaja, entre Huachocolpa y Tintay, Tovar 4201 (GH). Cuzco: Prov. Paucartambo, near Santa Isabel, Pilahuata to Patria, Plowman & Davis 4988 (GH, USM). Prov. Uru- bamba. Puente Ruinas, Machu Picchu, Iltis et al. 1025 (GH, uc, us). Puno: Cerca a San Juan del Oro, Valle del Alto Tambopata, Ferreyra 16684 (GH, USM). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 133 X — — ^^ri ^^ri^T^fc^li ^^ '*^z^r ~<^s *'/•': FIG. 23. Cyathea caracasana \ar. boliviensis: a, rachis and base of pinna; b, sori (one opened); c, portion of petiole scale. (From Wurdack 1653, F.) 134 FIELDIANA: BOTANY lib. Cyathea caracasana var. meridensis (Kar- sten) Tryon, Contr. Gray Herb. 206: 79. 1 976. Cyathea meridensis Karsten, Fl. Columb. 2: 161, t. 184. 1869. TYPE: Venezuela, (Merida), Merida, 2000 m, Karsten (holotype, not located; Karsten's illustration is probably taken from the type col- lection). In forests, 1800-2300 m, Amazonas, south to Junin. Venezuela, Colombia, Ecuador, and Peru. Amazonas: Prov. Bongara, Pomacocha a Yambras- bamba, Ferreyra 15243 (GH). Serrania de Bagua, E of La Peca, Gentry et al. 22897 (MO). San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba, D. Smith 4401 (GH). Huanuco: Prov. Leoncio Prado, Sullivan & Young 1178 (USM). Pas- co: Prov. Oxapampa, Rio San Alberto valley, Smith & Pretel 7999 (F, MO). Junin: Prov. Chanchamayo, Chilpez, S of San Ramon, Smith & Palacios 2641 (F, MO). 12. Cyathea lechleri Mett., Fil. lechl. 2: 32. 1859. TYPE: Peru, (Puno), St. Gaban (Rio San Ga- ban) Lechler (holotype, LZ destroyed; authen- tic specimen, Lechler 2309, Herb. Mett. B!). Cyathea castanea Baker, Syn. fil. ed. 2, 451. 1874. TYPE: Peru, (San Martin), Tarapoto, Spruce 47 23 (holotype, K; isotypes, GH!, MO!, P!, us!; photos, GH, US Of P). Petiole nearly smooth, the scales reddish brown to atropurpureous, concordantly bicolorous with lighter margins, scurf essentially absent. Lamina 2-pinnate-pinnatind to 2-pinnate-pinnatisect, pin- nules sessile to short-stalked, glabrate abaxially, ultimate segments subentire to slightly crenate, fertile veins forked. Sori medial, indusia sphaer- opteroid. In wet forests and cloud forests, ca. 600-1500 m, San Martin, Huanuco, Pasco, and Puno. Venezuela, Peru, and Bolivia. San Martin: 36 km NE of Tarapoto, Rio Cainarache, Gentry et al. 37928 (GH, MO). Huanuco: Cerros del Sira, Rio Llullapichis watershed, Dudley 1 3007, 13213, 13262 (GH). Pasco: Prov. Oxapampa, W side of Cordillera de San Matias, D. Smith 2046 (F, MO). Petiole smooth or with a few scattered tubercles, the scales reddish brown to atropurpureous, con- cordantly bicolorous with lighter margins, scurf absent. Lamina 2-pinnate-pinnatind, pinnules long-stalked, glabrous to slightly pubescent abax- ially, ultimate segments subentire to shallowly crenate, fertile veins simple or forked. Sori more or less medial, indusia sphaeropteroid. Growing at 2100 m, Amazonas. Venezuela and Colombia, south to Peru. This species is unique in the genus in the char- acters of the basal segments of the pinnules being decurrent onto the pinnule-stalk which is of a strongly contrasting lighter color than the dark pinna-rachis. Amazonas: Prov. Bongara, N end of lake Pomacocha, Hutchison & Wright 6814 (GH, NY, us). 14. Cyathea dudleyi Tryon, Contr. Gray Herb. 206: 85. 1976. TYPE: Peru, Cuzco, Prov. La Convencion, Cordillera Vilcabamba, Dudley 10867B (holotype, GH!; paratypes, same lo- cality, Dudley 10738, 10867 GH!). Petiole nearly smooth, the scales brown to dark brown, nearly concolorous, scurf absent. Lamina 3-pinnate nearly throughout, pinnules sessile to short-stalked, somewhat scaly and pubescent abaxially, ultimate segments entire, fertile veins simple. Sori subcostal, indusia sphaeropteroid. Known only from five collections, in dwarf for- ests or wet, dense cloud forests, 2100-2700 m, Pasco and Cuzco. Endemic to Peru. Cyathea dudleyi has a more complex lamina than the related C. lechleri and the concolorous petiole scales distinguish it from the somewhat smaller C. microphylla with bicolorous scales. Pasco: San Gotardo, border of Prov. Oxapampa and Pasco, van der Werff el al. 8589 (MO, uc). Prov. Oxa- pampa, 20 km W of Oxapampa. D. Smith 5375 (GH). Comments 13. Cyathea ebenina Karsten, Linnaea 28: 461. 1856. TYPE: Venezuela (Aragua), between Caracas and Puerto Cabello, Karsten (holo- type, not located; isotype, Caracas, Karsten, B!). Two collections from Dept. Amazonas, Prov. Bagua, Cordillera Colan, Barbour 3748 and 3920 (MO), may be Cyathea fulva (Mart. & Gal.) Fee. The materials are not wholly adequate for certain TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 135 identification and the species is not treated under Cyathea. It is known in Ecuador and may well occur in northern Peru. Specimens of Cyathea fulva will key out near C. delgadii, from which it differs in having few or no scales or trichomes on the segments abaxially, or with some trichomes and also some scales. Cy- athea delgadii has few or no scales on the abaxial side of the segments, and the abaxial side is mod- erately to usually abundantly long-pubescent. In addition, Cyathea fulva has some to many of the large scales of the petiole scurf crested, while in C. delgadii they are flattish. VI. Cnemidaria Cnemidaria Presl, Tent, pterid. 56. 1836. TYPE: Cnemidaria speciosa Presl. Figure 24. Terrestrial. Stem rather small to stout, ascend- ing to erect, usually hardly arborescent, rarely to 3 m tall. Leaves monomorphic, with scales, es- pecially on the croziers and petiole base, that are marginate and lack a dark apical seta. Petiole smooth, muricate, or with corticinate spines. Lam- ina 1 -pinnate to 1 -pinnate-pinnatisect, veins free and the basal ones connivent to a sinus, or usually regularly anastomosing without included free veinlets, especially along the costa. Sori round, borne on the veins, rarely at a fork, with a hem- itelioid or meniscoid indusium. Spores tetrahe- dral-globose, trilete, nearly smooth, with three large equatorial pores and often smaller ones. A tropical American genus of 25 species, with five of them in Peru. Species of Cnemidaria were formerly placed in Hemitelia because of the small indusia. The genus is characterized by a lack of trichomes on the adax- ial side of the costae and costules and by the spores with three large equatorial pores. All of the Pe- ruvian species have regularly anastomosing veins, except for C. uleana, which sometimes has free veins. Reference STOLZE, R. 1974. A taxonomic revision of the genus Cnemidaria (Cyatheaceae). Fieldiana, Bot., 37: 1-98. Key to Species of Cnemidaria a. Pinnae (excluding the basal pair and reduced apical ones) deeply pinnatifid or pinnatisect; the segment sinuses extending % or more to the costa b b. Lamina gradually reduced to a nonconform pinnatifid apex; rachis not alate; petiole scales bi- colorous, dark brown with broad to narrow whitish margins c c. Basal basiscopic veins commonly arising from the costule or from its junction with the costa; rachis and petiole muricate to spiny; scales of costae and costules, if present, deep brown . . . l.C. horrida c. Basal basiscopic veins, especially toward the basal and apical portions of pinnae, commonly arising from the costa; rachis smooth, petiole smooth or tuberculate; scales of costae and costules whitish 2. C. uleana b. Lamina abruptly reduced to a conform or subconform apical segment similar to the lateral pinnae; rachis conspicuously alate, especially distally; petiole scales predominantly whitish 3. C. alatissima a. Pinnae (excluding the basal pair and reduced apical ones) subentire to shallowly pinnatifid; the segment sinuses, if present, extending less than halfway to the costa d d. Pinnae shallowly and obtusely lobed to shallowly pinnatifid; indusia more or less semicircular, attached on the costular side of the receptacle 4. C. speciosa d. Pinnae subentire to broadly and coarsely serrate; indusia circular, completely surrounding the receptacle 5. C. nervosa 136 FIELDIANA: BOTANY FIG. 24. Cnemidaria speciosa: a. part of rachis, with pinnae; b, pinna midrib, venation pattern, and sori; c, petiole scale, (a from Tryon & Tryon 5260, F, b-c from Buchtien 5224, Bolivia, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 137 1. Cnemidaria horrida (L.) Presl, Tent, pterid. 57. 1836. Polypodium horridum L., Sp. pi. 1092. 1753. TYPE: Plumier descript. pi. Amer., /. 4. 1693 (Traite foug. Amer., /. 8. 1705). Hemitelia horrida (L.) R. Br., Prodr. 158. 1810. Petiole muricate, or more often with stout spines to 5 mm long, the scales, usually only at the base of the petiole, bicolorous, dark brown with narrow whitish margins. Lamina gradually reduced to a nonconform pinnatind apex, rachis not alate, mur- icate or often spiny. Pinnae incised % or more to the costa, the costa and costules rarely with dull brown scales abaxially, basal basiscopic veins usu- ally arising from the costule. Soral lines supra- medial to submarginal between the costule and segment margin, indusia more or less semicircular, subentire to erose. In forest, on stream banks, and mountain slopes, 300-1 500 m, Amazonas and Loreto, south to Cuz- co. Greater Antilles; Costa Rica; Venezuela and Co- lombia south to Peru. Cnemidaria horrida is the most widely distrib- uted species of the genus and has the greatest range of altitude. It is also the largest species with trunks sometimes to 3 m tall and 7 cm in diameter. Amazonas: Prov. Bagua, Rio Maranon, above Cas- cadas de Mayasi, Wurdack 1991 (us). San Martin: Near Tarapoto, Spruce 3943 (GH, us). San Roque, LI. Wil- liams 7156 (F, NY, us). Loreto: Pumayacu, between Bal- sapuerto and Moyobamba, Klug 3182 (F, GH, MO, NY, us). Pasco: Prov. Oxapampa, Rio Palcazu, Smith & Sal- ick 8377 (MO). Cuzco: Prov. Paucartambo, Villa Carmen, Kosnipata-Pilcopata, Vargas 77269(GH). 2. Cnemidaria uleana (Samp.) Tryon var. uleana. Cnemidaria uleana (Samp.) Tryon, Contr. Gray Herb. 200: 52. 1970. Hemitelia uleana Samp., Bol. Mus. Nac. Rio de Ja- neiro 1:65.1 923. TYPE: Brazil, (Rio de Janeiro), Perto de Nova Friburgo, Alta da Serra, Vie (ho- lotype, R!; photos, F, GH). Petiole smooth to slightly muricate, the scales bicolorous, dark brown with broad whitish mar- gins. Lamina gradually reduced to a nonconform pinnatind apex, rachis not alate, essentially smooth. Pinnae incised %-% to the costa, the costa and costules usually with broad, pale yellowish to whit- ish scales abaxially, basal basiscopic veins, espe- cially toward the basal and apical portions of the pinnae, commonly arising from the costa. Soral lines medial to supramedial between the costule and segment margin, indusia semicircular, sub- entire to several-lobed. In cloud forests, 1700-2200 m, Cuzco. Southern Peru and southeastern Brazil. Cnemidaria uleana is unique in the genus in that anastomosing of the basal veins may be present or absent. All other Peruvian species have costal areolae. Specimens of var. uleana with anasto- mosing veins can be separated from Cnemidaria horrida by the characters in the key. The species ranges from Colombia to Peru, and in southeastern Brazil. The collections from Peru are var. uleana (also in Brazil), with a gradually reduced pinnatifid lamina apex and usually with costal areolae. The var. abitaguensis (Domin) Stolze occurs in Colombia and Ecuador, with a conform or subconform apical segment and free veins. Cuzco: Prov. La Convention, Cordillera Vilcabamba, Dudley 10459 (GH, NA), 706/7 (GH), 77J72 (GH, NA). 3. Cnemidaria alatissima Stolze, Fieldiana, Bot. 37: 55. 1974. TYPE: Peru, Huanuco, Rio Llullapichis watershed, Dudley 13282 (holo- type, GH! 2 sheets; photo, F; isotype, NA!). Petiole with minute spines, thickly covered near the base by large, ovate, whitish scales. Lamina abruptly reduced to a subconform apical segment, rachis with a green wing 1-2 mm wide on each side, smooth. Pinnae incised %-% to the costa, the costa and costules lacking scales abaxially, basal basiscopic veins usually arising from the costule. Soral lines about medial between the costule and segment margin, indusia often rudimentary, usu- ally less than semicircular, subentire. In dense cloud forests, ca. 1 500 m, Huanuco. Endemic to Peru. This species is characterized by the abundant white petiole scales and the broad green alate rach- is, the wings extending down onto the petiole. It is known only from the type collection and one juvenile plant. Huanuco: Rio Llullapichis watershed, Dudley 13281 (GH, us), a juvenile plant. 138 FIELDIANA: BOTANY 4. Cnemidaria speciosa Presl, Tent, pterid. t. 1, f. 16-17. 1836. TYPE: Peru, (Huanuco), Pam- payacu, Jul. 1829, Poeppig 221 (Diar. 1144) (holotype, PR or PRC; frag., NY!, us!; isotypes, B!, BR, P!). Figure 24. Hemitelia subincisa Kunze, Bot. Zeit. (Berlin) 2: 296. 1844, nom. nov. for Cnemidaria speciosa Presl, not Hemitelia speciosa (Willd.) Kaulf. Petiole smooth or rarely tuberculate at the base, the scales lanceolate to ovate, bicolorous, whitish with a brown central stripe. Lamina abruptly re- duced to a conform or subconform apical segment, rachis not alate, essentially smooth. Pinnae very deeply crenate to lobed less than '/2 to the costa, the costa and costules rarely with dull, brown scales abaxially, basal basiscopic veins mostly arising from the costule or its base. Soral lines supra- medial between the costule and segment margin, indusia more or less semicircular, rather large. In forests and at forest borders, along stream banks, and on mountain slopes, 1 1 5-1900 m, San Martin and Loreto south to Puno. Peru and Bolivia. Three collections from Loreto, Varadero de Ma- zan, (Rio Amazonas to Rio Napo), Croat 19500 (MO, uc), 79574 (F, MO) and 20797 (MO, uc) have a few broad whitish scales on the costae abaxially, rather than few small brown ones or none. These may represent a distinct variety of the species. San Martin: Prov. Mariscal Caceres, Palo Blanco, To- cache Nuevo, J. Schunke V. 5682 (F, GH, NY). Loreto: Prov. Maynas, Varadero, Vdsquez 721 (MO). Huanuco: Prov. Huanuco, Tingo Maria, Tryon & Tryon 5260 (F, GH, u, us). Pasco: Yapas, Pichis Trail, Killip & Smith 25563 (GH, NY, us). Prov. Oxapampa, Palcazu, Cerro de Pantiacolla, Foster 10907 (F). Junin: Near La Merced, Killip & Smith 23889 (NY, us). Chanchamayo valley, C Schunke 52 (F). Ucayali: Prov. Coronel Portillo, Padre Abad, J. Schunke V. 5477 (F, GH, us). Aguaytia, Hua- palla 2470 (USM). Cuzco: Prov. Quispicanchi, entre In- ambari y Quince Mil, Vargas 16502 (GH). Madre de Dies: Prov. Manu, Vargas 17738 (GH). Puno: Prov. Car- abaya, San Gaban, Vargas 18863 (GH). 5. Cnemidaria nervosa (Maxon) Tryon, Contr. Gray Herb. 200: 52. 1970. Hemitelia nervosa Maxon, J. Wash. Acad. Sci. 34: 309. 1944. TYPE: Peru, Loreto, mouth of Rio Santiago, Mexia 6291 (holotype, us! 3 sheets; is- otypes, F!, GH!, NY!, uc!). Petiole smooth to muricate, the scales, mostly near the base, lanceolate or linear-lanceolate, bi- colorous, brown with narrow whitish margins. Lamina abruptly reduced to a conform or subcon- form apical segment, rachis not alate, smooth. Pin- nae subentire to broadly serrate, the costa and pri- mary veins lacking scales abaxially, basal basiscopic vein arising from a primary vein. Soral lines be- tween the primary veins, indusia commonly cir- cular, completely surrounding the receptacle, en- tire to lobed. Rain forests, 300—450 m, Amazonas and Lor- eto. Ecuador and Peru. The circular indusium is a character in the genus shared only by Cnemidaria cocleana of Panama. Cnemidaria nervosa is a rare fern, represented by one collection from Ecuador and two from Peru. Amazonas: Prov. Bagua. valley of the Rio Maranon, Wurdack 2059 (NY, us). TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 139 Colombia 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. Tumbes Piura Lambayeque Cajamarca Amazonas La Libertad San Martin Loreto Ancash Hulnuco Lima Pasco Junlh Ucayali lea Huancavelica Ayacucho ApurTmac Cuzco Madre de Dios Arequipa Puno Moquegua Tacna Bolivia Chile DEPARTMENTS OF PERU 140 FIELDIANA: BOTANY Index to Names Accepted names are in roman type, synonyms are in italics, and new names are in boldface. A page number is provided for the principal place, or the only place, where the name occurs. Acrostichum dichotomum 33 digit at um 33 elegans 34 nodosum 15 Actinostachys 33 digit at a 33 pennula 36 Alsophila 1 16 armigera 123 aterrima 1 14 a usual is 1 16 blechnoides 1 1 1 capon sis 1 18 ssp. polypodioides 1 1 8 caracasana 133 conjugata 128 contracta 109 cuspidata 120 dombeyi 123 elongata 1 1 5 engelii 1 1 6 erinacea 1 1 8 floribunda 123 frigida 126 incana 120 infesta 123 kalbreyeri 124 killipii 125 lasiosora 125 latevagans 124 fecAter/ 128 macrosora \ 1 5 melanopus 124 microdonta 127 mgra 124 nigripes 124 pallescens 131 paucifolia 1 18 peruviana 123 phegopteroides 127 pilosissima 125 podophylla 124 poeppigii 1 1 5 procera 123 pruinata 109 pterorachis 123 pubescens 126 pycnocarpa 123 quadripinnata 109 rostrata 1 1 1 sprucei 1 1 5 swartziana 122 tarapotensis 125 tryonorum 128 «/« 128 Aneimia 24 Anemia 24 subg. Coptophyllum 24 adiantifolia 24 buniifolia 24 Anemia clinata 25 ferruginea 27 var. ahenobarba 27 var. ferruginea 27 flexuosa 27 x phyllitidis 27 var. setosa 27 haenkei 29 hirsuta 28 var. humboldtiana 28 hispida 30 humilis 29 myriophylla 28 oblongifolia 29 var. humilis 29 pastinacaria 28 phyllitidis 29 repens 30 tomentosa 27 villosa 25 var. anthriscifolia 27 Anemirhiza 24 adiantifolia 24 Aspidium rostratum 1 1 1 Balantium karstenianum 105 Blechnum 101 Botrychium 6 subg. Osmundopteris 6 subg. Sceptridium 6 cicutarium 8 lunaria 6 schaffneri 6 underwoodianum 6 virginianum 8 var. mexicanum 8 virginicum 8 fotfa mexicanum 8 /foes/a mirifica 58 Cheiroglossa 8 palmata 9 Christensenia 13 Cnemidaria 136 alatissima 138 cocleana 139 horrida 138 nervosa 139 speciosa 139 uleana 138 var. abitaguensis 1 38 var. uleana 138 Coptophyllum 24 buniifolium 24 Culcita 103 subg. Calochlaena 103 coniifolia 103 macrocarpa 103 Cyathea 129 andina 130 arborea 129 aterrima 1 14 bipinnatifida 126 caracasana 133 var. boliviensis 133 var. caracasana 1 33 var. chimborazensis 133 var. maxonii 133 var. meridensis 135 castanea 135 cuspidata 120 delgadii 132 divergens 131 var. divergens 1 3 1 var. tuerckheimii 131 dudleyi 135 ebenina 135 elongata 1 1 7 equestris 131 erinacea 1 1 8 frigida 126 fulva 135 incana 120 kalbreyeri 124 lasiosora 125 latevagans 124 lechleri 135 macrosora 1 1 5 meridensis 135 mexicana 133 var. boliviensis 1 33 microdonta 127 microphylla 132 multiflora 130 multisegmenta 132 nigra 125 nigripes 124 oligocarpa 132 oreites 1 1 5 pallescens 131 panamensis 131 petiolata 131 phegopteroides 127 /7/7o.sa 132 pilosissima 125 poeppigii \ 1 5 polystichoides 1 1 8 primaeva 125 procera 123 pubens 127 pubescens 127 pungens 123 quindiuensis 1 1 6 rufescens 1 14 rui /i ana 132 TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 141 Cyathea schanschin 132 sprucei 1 1 5 subtropica 128 ulei 128 vilhelmii 130 willdenowiana 123 Cyatheaceae 1 1 1 Cystodium 103 Danaea 15 cuspidata 19 elliptica 18 var. crispula 18 grandifolia 17 humilis 20 jamaicensis 19 longifolia 17 moritziana 19 nodosa 17 oblanceolata 18 stenophylla 19 trichomanoides 20 wendlandii 20 Danaeaceae 13 Davalliopsis elegans 84 Dennstaedtia pearcei 99 Dicksonia 105 arborescens 105 berteriana 105 coniifolia 103 gigantea 105 karsteniana 105 pearcei 99 sellowiana 105 spruceana 105 stuebelii 105 Dicksoniaceae 101 Dicranopteris 46 affinis 42 bancroftii 39 bifida 41 brittonii 44 dichotoma 46 flexuosa 47 f. monstrosa 49 linearis 47 longipinnata 45 nervosa 47 pectinala 49 pennigera 39 peruviana 45 pruinosa 42 remold 44 mbiginosa 46 schomburgkiana 47 seminuda 47 simplex 39 velata 42 yungensis 44 Didymoglossum 76 angustifrons 86 hymenoides 86 Didymoglossum krausii 87 membranaceum 88 muscoides 86 reptans 87 sphenoides 86 Diplopterygium 37 bancroftii 39 Eupodium 15 kaulfussii 15 diversifrons 9 1 heterophylla 94 humboldtii 94 91 Gleichenia 37 sect. Diplopterygium 37 o$n/s 42 bancroftii 39 bifida 41 boliviensis 43 buchtienii 42 coslaricensis 43 flexuosa 47 f. monstrosa 49 glauca 37 hypoleuca 43 lechleri 44 leucocarpa 44 longipinnata 45 mafAwsn 41 mellifera 45 nervosa 47 nitidula 43 pectinata 49 pennigera 39 peruviana 45 polypodioides 37 pruinosa 42 remota 44 revoluta 42 r/£/rffl 47 rubiginosa 46 f. virescens 46 simplex 39 subandina 43 tomentosa 4 1 truncata 37 tuberculata 43 ve/a/a 42 yungensis 44 Gleicheniaceae 37 Hemiphlebium kapplerianum 88 Hemitelia 129 andina 130 horrida 138 fccAferi 131 multiflora 130 Hemitelia nervosa 139 petiolata 131 rufescens 1 1 4 speciosa 139 subincisa 139 uleana 138 Hicriopteris bancroftii 39 Homoeotes 76 heterophylla 94 Hydroglossum 33 oligostachyum 33 Hymenophyllaceae 49 Hymenophyllum 50 sect. Buesia 59 sect. Hymenophyllum 56 sect. Sphaerocionium 52 subg. Hymenophyllum 52 subg. Leptocionium 50 subg. Mecodium 50 subg. Sphaerocionium 50 adiantoides 66 amabile 70 andinum 62 apiculatum 59 axillare 62 beyrichianum 68 calodictyon 56 ciliatum 67 contortum 64 crispatulum 68 crispum 67 var. bipinnatisectum 67 var. crispum 67 cristatum 55 dendritis 59 dependens 72 dicranotrichum 50 ectocarpon 57 elegans 66 elegantulum 70 endiviifolium 63 fendlerianum 64 ferax 61 fragile 69 fucoides 55 var. calodictyon 56 var. chachapoyense 57 var. fucoides 56 var. pedicellatum 57 fusagasugense 73 var. aberrans 73 fusugasugense 73 hirsutum 66 interruptum 72 karstenianum 71 1. 1 n H' I la in in 58 /ar/pes 59 lindenii 7 1 lineare 66 lobatoalatum 74 mathewsii 60 mexiae 59 microcarpum 68 mirificum 58 142 FIELDIANA: BOTANY Hymenophyllum molle 65 multialatum 73 multiflorum 63 myriocarpum 62 var. endiviifolium 63 var. myriocarpum 62 var. nigrescens 63 nigrescens 63 nigricans 63 nigricans 63 pedicellatum 57 peltatum 57 peruvianum 75 platylobum 68 plumieri 72 plumosum 72 poeppigianum 76 polyanthos 59 polycarpum 64 procerum 65 pulchellum Hooker 70 pulchellum Mett. 65 pyramidatum 74 reniforme 64 rimbachii 64 ruizianum 69 rupestre 82 simplex 69 speciosum 70 spectabile 70 sprucei 76 superbum 72 tarapotense 75 tenerrimum 66 tomentosum 73 tortuosum 57 trapezoidale 69 trianae 61 trichomanoides 61 trichophyllum 65 var. buesii 66 var. trichophyllum 65 trifidum 76 tunbridgense 50 undulatum 63 var. fendlerianum 64 var. undulatum 64 valvatum 68 verecundum 75 Hymenostachys diversifrons 9 1 Lacostea tanaica 89 Lecanium membranaceum 88 Leptocionium 50 dicranotrichum 50 fucoides 56 pedicellatum 57 Leptopteris 20 Lomaria euphlebia 101 Lomaridium semicordatum 101 Lophidium 33 elegans 34 flabellum 34 latifolium 33 poeppigianum 36 Lophosoria 107 pruinata 109 quadripinnata 107 var. contracta 109 var. quadripinnata 109 Lophosoriaceae 107 Loxoma 98 Loxomataceae 98 Loxsoma 98 Loxsomopsis 99 costaricensis 99 lehmannii 99 notabilis 99 pearcei 99 Lygodium 30 digitatum 32 mexicanum 33 micans 32 oligostachyum 33 polymorphyum 33 radiatum 32 scandens 30 venustum 30 volubile 32 Marattia 13 alata 15 var. laevis 1 5 alata 15 kaulfussii 15 laevis 15 Marattiaceae 13 Mecodium 50 apiculatum 59 contortum 64 dendritis 59 endiviifolium 63 fendlerianum 64 ferax 6 1 mexiae 59 microcarpum 68 multiflorum 63 myriocarpum 62 polyanthos 59 trichomanoides 6 1 undulatum 64 Meringium fucoides 56 Mertensia 46 bancroftii 39 Mertensia pectinata 49 pennigera 39 pruinosa 42 remota 44 revoluta 42 simplex 37 tomentosa 4 1 dichotoma 46 flexuosa 47 laevigata 37 longipinnata 45 mathewsii 4 1 nervosa 47 Metaxya 109 rostrata 1 1 1 Metaxyaceae 109 Mohria 23 Nephelea 118 cuspidata 120 erinacea 1 1 8 var. erinacea 1 1 8 var. purpurascens 120 incana 120 polystichoides 1 1 8 Neuromanes pinnatum 92 Neurophyllum hostmannianum 93 pinnatum 92 Odontomanes hostmannianum 93 Onoclea polypodioides 37 Ophioglossaceae 5 Ophioglossum 8 coriaceum 12 crotalophoroides 12 ellipticum 12 lusitanicum 12 ssp. californicum 12 ssp. coriaceum 12 ssp. lusitanicum 12 nudicaule 1 1 var. tenerum 1 1 opacum 12 palmatum 9 pendulum 8 peruvianum 9 petiolatum 9 reticulatum 9 scandens 30 scariosum 1 1 tenerum 1 1 vulgatum 8 ypanemense 1 1 Osmunda 21 adiantifolia 24 cicutaria 8 cinnamomea 21 var. imbricata 21 flexuosa 27 hirsuta 28 humilis 29 lunaria 6 TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 143 Osmunda oblongifolia 29 palustris 23 phyllitidis 29 regalis 23 ssp. palustris 23 var. regalis 23 var. spectabilis 23 spectabilis 23 virginiana 8 Osmundaceae 20 Plagiogyria 101 sect. Carinatae 101 costaricensis 101 denticulata 101 euphlebia 101 latifolia 101 semicordata 101 Plagiogyriaceae 99 Polypodium arboreum 129 dicholomum 46 glaucum 109 horridum 138 medullare 1 1 2 microdontum 127 pedicellata 15 procerum 123 pruinatum 109 pungens 123 quadripinnatum 109 rostratum 1 1 1 Ptilophyllum bicorne 9 1 hostmannianum 93 lambertianum 95 martiusii 95 pellucens 96 Ragatelus crinitus 94 Schizaea 33 dichotoma 33 digitata 33 elegans 34 tistulosa 34 var. australis 34 flabellum 34 incurvata 36 pennula 36 poeppigiana 36 pusilla 34 Schizaeaceae 23 Selenodesmium rigidum 85 Sphaerocionium 50 adiantoides 66 ciliatum 67 crispum 67 elegans 66 elegantulum 70 fragile 69 hirsutum 67 Sphaerocionium interruptum 72 karstenianum 71 lindenii 1 1 lobatoalatum 74 microcarpum 68 multialatum 73 nigricans 63 plumieri 72 plumosum 72 pyramidatum 74 ruizianum 69 simplex 69 spectabile 1 1 tenerrimum 66 tomentosum 73 trichophyllum 65 valvatum 68 Sphaeropteris 1 1 2 subg. Sclephropteris 1 14 subg. Sphaeropteris 1 14 atahuallpa 1 1 5 aterrima 1 14 bradei 115 elongata 1 1 5 hirsuta 1 14 horrida 112 macrosora 1 1 4 var. macrosora 1 1 4 var. reginae 1 1 5 var. vaupensis 1 1 5 medullaris 1 1 2 quindiuensis 1 16 rufescens 1 14 Sticherus 37 affinis 42 bifidus 4 1 buchtienii 42 laevigatus 37 lechleri 44 longipinnatus 45 mathewsii 4 1 nitidulus 43 penniger 39 pruinosus 42 revolutus 42 rubiginosus 46 simplex 39 tuberculatus 43 velatus 42 yungensis 44 Todea 20 Trichipteris 120 conjugata 128 corcovadensis 120 dombeyi 123 excelsa 120 flava 125 frigida 126 infest a 123 kalbreyeri 124 lasiosora 125 latevagans 124 Trichipteris lechleri 127 microdonta 127 nigra 124 nigripes 124 var. brunnescens 124 phegopteroides 127 pilosissima 125 procera 123 pubescens 126 serpens 126 tryonorum 128 Trichomanes 76 sect. Didymoglossum 90 sect. Lacostea 89 sect. Microgonium 78 sect. Neurophyllum 93 subg. Achomanes 76 subg. Cardiomanes 78 subg. Didymoglossum 76 subg. Pachychaetum 76 subg. Trichomanes 78 accedens 98 angustatum 83 angustifrons 86 ankersii 90 var. tanaicum 89 bicorne 9 1 botryoides 92 brachyblastos 8 1 capillaceum 84 cellulosum 85 ciliatum 67 collariatum 81 crinitum 94 crispum 76 var. haenkeanum 98 cristatum 97 delicatum 98 diaphanum 83 diversifrons 9 1 ekmanii 88 elatum 96 elegans 84 elegans 9 1 fragile 69 fucoides 56 haenkeanum 98 heterophyllum 94 hirsutum 67 hookeri 88 hostmannianum 93 humboldtii 94 hymenoides 86 hymenophylloides 83 kapplerianum 88 krausii 87 kunzeanum 81 lambertianum 95 leptophyllum 83 lucens 95 martiusii 94 membranaceum 88 muscoides 76 opacum 84 pedicellatum 90 144 FIELDIANA: BOTANY Trichomanes Trichomanes Trichomanes pellucens 96 pyxidiferum 82 tuerckheimii 90 pellucidum 96 radicans 8 1 undulatum 97 peltatum 57 var. kunzeanum 8 1 vandenboschii 97 pennatum 92 reptans 87 Trichopteris 122 pilosum 95 rigidum 85 pilosum 94 rupestre 82 pinnatum 92 scandens 76 Vandenboschia plumosum 96 sellowianum 97 angustata 83 plumula 95 sphenoides 86 capillacea 84 poeppigii 89 spruceanum 91 diaphana 83 polyanthos 59 sprucei 86 hymenophylloides 83 polypodioides 89 subsessile 90 pyxidifera 82 prieurii 84 tanaicum 89 radicans 8 1 punctatum 86 tenerum 83 tenera 83 ssp. sphenoides 86 trollii 9 1 TRYON & STOLZE: PTERIDOPHYTA OF PERU. I. 145 281 5/93 Other Fieldiana: Botany Titles Available Publication 1246, SI 0.50 Publication 13* Publication 1349. UNIVERSITY OF ILLINOIS URBANA 580 5FBN.S COOS FIELDIANA BOTANY CHGO 20 1988 30112017564094 UNIVERSITY OF ILLINOIS-URBANA