OF ILLINOIS LIBRARY AT URBANA-CHAMPA1GN BIOLOGY FIEL Botany NEW SERIES, NO. 27 PTERIDOPHYTA OF PERU Part IV 17. Dryopteridaceae Rolla M. Tryon Robert G. Stolze With the collaboration of: John T. Mickel Robbin C. Moran "Mr 21 February 28, 1991 Publication 1424 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Information for Contributors to Fieldiana General: / warily a journal for Field Museum staff members and research associates, although nsidered as space permits. i65.00 per printed page or fraction thereof. Payment of at least 50% of page which reduces the publication time. Contributions from staff, research authors will be considered for publication regardless of ability to pay page charges, however, the full i'.ed authors of unsolicited manuscripts. 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FIELDIANA Botany NEW SERIES, NO. 27 PTERIDOPHYTA OF PERU Part IV 17. Dryopteridaceae Rolla M. Tryon Harvard University Herbaria 22 Divinity Avenue Cambridge, Massachusetts 02138 and Department of Biology University of South Florida Tampa, Florida 33620-5150 Robert G. Stolze Associate Curator Department of Botany Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 With the collaboration of: John T. Mickel, New York Botanical Garden and Robbin C. Moran, Missouri Botanical Garden Accepted August 10, 1990 Published February 28, 1991 Publication 1424 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1 99 1 Field Museum of Natural History Library of Congress Catalog Card Number: 90-85755 ISSN 00 15-0746 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents List of Illustrations INTRODUCTION ACKNOWLEDGMENTS . . 17. DRYOPTERIDACEAE Ctenitis . . 1 1 2 5 Megalastrum 11 Triplophyllum 19 Tectaria 21 Cyclopeltis 29 Rumohra 31 Lastreopsis 31 Dryopteris 35 Cyrtomium 38 Didymochlaena 40 Stigmatopteris 42 Cyclodium 47 Polystichum 49 Olfersia 55 Polybotrya 57 Diplazium 65 Athyrium 88 Hemidictyum 90 Cystopteris 92 Woodsia 94 Oleandra 96 Bolbitis 98 Lomariopsis 105 Lomagramma 109 Elaphoglossum Ill Peltapteris 167 MAP OF PERU 171 INDEX TO NAMES . .172 1. Ctenitis: C. submarginalis; C. ampla . . 6 2. Megalastrum biseriale 12 3. Triplophyllum funestum var. funest um 20 4. Tectaria incisa var. incisa 22 5. Cyclopeltis semicordata 28 6. Rumohra adiantiformis 30 7. Lastreopsis: L. effusa; L. killipii 32 8. Dryopteris: D. paleacea; D. denticulata 34 9. Cyrtomium dubium 39 10. Didymochlaena truncatula 41 1 1 . Stigmatopteris: S. heterophlebia; S. pel- lucidopunctata 43 12. Cyclodium: C. trianae; C. meniscioides var. meniscioides 46 1 3. Polystichum: P. pycnolepis; P. platyphyl- lum; P. montevidense var. montevi- dense 50 14. Olfersia cervina 56 15. Polybotrya: P. fractiserialis; P. osmun- dacea; P. polybotryoides 58 16. Diplazium: D. cristatum; D. roemeri- anum; D. striatum 66 1 7. Athyrium dombeyi 89 18. Hemidictyum marginatum 91 19. Cystopteris fragilis 93 20. Woodsia montevidensis 95 21. Oleandra: O. articulata; O. lehmannii . . 97 22. Bolbitis: B. serrata; B. serratifolia; B. lindigii 99 23. Lomariopsis: L. fendleri; L. japurensis; L. nigropaleata 104 24. Lomagramma guianensis 108 25. Elaphoglossum: E. paleaceum; E. eri- naceum; E. piloselloides; E. squamipes ..110 26. Peltapteris peltata f. peltata 168 111 PTERIDOPHYTA OF PERU Part IV 17. Dryopteridaceae Introduction This fourth part of the "Pteridophyta of Peru" is devoted exclusively to the family Dryopterida- ceae. The general style, typography, form of ci- tations, and so forth follow the previously pub- lished parts I and II. These matters are adequately dealt with in part I (Fieldiana: Botany, n.s. No. 20, 1989), and it is not necessary to repeat them here. Stigmatopteris, Olfersia, and Polybotrya have been contributed by Robbin C. Moran, and Elaph- oglossum and Peltapteris have been contributed by John T. Mickel. Other genera have been a joint effort of Rolla M. Tryon and Robert G. Stolze, each critically reviewing the treatments prepared by the other. The Dryopteridaceae are the largest family of Pteridophyta in Peru, with 26 genera and about 200 species. Many of the genera are taxonomically difficult because of considerable variation and the lack of recent critical work on the Andean ele- ments that would provide guidance. Among these may be mentioned Ctenitis, Megalastrum, Tec- tana, Polystichum, Diplazium, and Elaphoglos- sum. In these genera there is clearly a maximum of variation in the Andes from Bolivia north to Colombia, and also extensive intergradation be- tween many of the variations of a species. The taxonomy is often less complicated to the east and north, especially in Mexico and Central America. Accordingly, a number of commonly recognized species are placed as synonyms for Peru or they are maintained with some doubt. Although the treatments for Peru are based on all materials from South America, or from tropical America, there is need for monographic study of many genera, so that the species characters are better known and intergradation and its geographic basis are more fully understood. Type collections from Peru are mentioned in the nomenclature but are not repeated in the spec- imen citations. They are, however, included in the Peru range and ecology. The nomenclature of the genera and species is not intended to be complete. It includes all names based on Peru material and other names that are considered useful to mention. C. V. Morton (Amer. Fern J. 62: 57-64. 1972) has argued that the Cryptogamae Vasculares Qui- tenses of Sodiro was published over a period of years, most of it in the Anales Univ. Quito, 1890- 1895, and a small part in the Anales Univ. Central Rep. Ecuador, 1897. He proposed that the book was not published until 1897. This conclusion may be correct, although Luis Mille, who was certainly in a position to know, indicated (Nov. Rec. Crypt. Vase. Ecuador, page 1 and page 5, 1927) that the book was published in 1893. We accept that date and consequently only one of the names cited for the Pteridophyta was published earlier in the An- ales: Acrostichum setigerum Sodiro, 1890 = Elaphoglossum setigerum (Sodiro) Diels. Abbreviations of periodicals generally follow the system of Botanico-Periodicum-Huntianum (1 968), while those of books and authors generally follow the system of Taxonomic Literature (TL- 2, 1976 et seq.) The acronyms for herbaria follow Index Herbariorum and are also provided below. Acknowledgments The authors are extremely grateful to Dr. John T. Mickel (Elaphoglossum, Peltapteris) and Dr. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. Robbin C. Moran (Olfersia, Polybotrya, Stigma- topteris) for contributing treatments of genera that are their particular specialties. Their having lent their expertise to this Flora has provided our best understanding of the species of these genera in Peru. We would like to extend special thanks to Dr. Abundio Sagastegui (HUT) and Blanca Leon (USM) for their invaluable assistance in preparing loans and arranging for packing and shipment of spec- imens from those two important Peruvian her- baria. The original drawings were contributed by Field Museum scientific illustrators Zorica Dabich and Clara L. Richardson, and by volunteer illus- trator Julia A. Liesse. Their careful study, pains- taking efforts, cooperation, and patience have pro- duced the splendid drawings which add so much to the descriptions. We are also grateful to Dr. David S. Harrington (Polystichurri) and Dr. Alan R. Smith (Cyclodium) for their critical suggestions on treatments of these two genera. Thanks are also extended to reviewers of the manuscript for their valuable comments. We are also grateful to the officers of the follow- ing institutions for granting loans of their material or allowing us to examine specimens in their her- baria: Herbarium Amazonense, Universidad Na- tional de la Amazonia Peruana, Iquitos, Peru (AMAZ); Herbarium Jutlandicum, Aarhus Univer- sitet, Denmark (AAU); Botanischer Garten und Bo- tanisches Museum, Berlin-Dahlem, Berlin (B); British Museum (Natural History), London (BM); Herbario Nacional Colombiano, Museo de His- toria Natural, Bogota, Colombia (COL); Royal Bo- tanic Garden, Edinburgh (E); Field Museum of Natural History, Chicago (F); Harvard University Herbaria, Cambridge, Mass.— most Gray Herbar- ium (GH), some Arnold Arboretum (A); Herbarium Truxillense, Universidad Nacional de Trujillo, Trujillo, Peru (HUT); Royal Botanic Gardens, Kew, England (K); Herbario Nacional de Bolivia, Uni- versidad Mayor de San Andres, La Paz, Bolivia (LPB); Missouri Botanical Garden, St. Louis (MO); New York Botanical Garden, New York (NY); Mu- seum National d'Histoire Naturelle, Paris (P); Her- bario del Institute de Ciencias Naturales, Univer- sidad Central, Quito, Ecuador (Q); Institute de Ciencias, Pontificia Universidad Catolica del Ec- uador, Quito, Ecuador (QCA); University of Cali- fornia, Berkeley (uc); Institute Botanico, Caracas, Venezuela (VEN); United States National Herbar- ium, Smithsonian Institution, Washington, D.C. (us); Herbario San Marcos, Universidad Nacional Mayor de San Marcos, Lima, Peru (USM); and Na- turhistorisches 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. Family 17. DRYOPTERIDACEAE Dryopteridaceae Herter, Rev. Sudam. Bot. 9: 15. 1949; description from Baker in Mart., Fl. brasil. 1(2): 458, Polypodiaceae, Tribe Aspi- deae. TYPE: Dryopteris Adans. Stem stout to slender, often branched, erect, rarely arborescent, decumbent to long-creeping or scandent, usually indurated, bearing scales. Leaves ca. 1 cm to 3 m long, circinate in vernation, mono- morphic to dimorphic, entire to usually pinnate, rarely to flabellate, glabrous, pubescent, or scaly abaxially. Petiole lacking stipules, not articulate to the stem. Veins usually free, to anastomosing. Sori abaxial, round to very elongate, at the tip of a vein, or along it, indusiate or exindusiate, or the spo- rangia generally distributed over the surface of the fertile segments; sometimes paraphysate, usually not. Sporangia with a 2- or 3-rowed stalk and a vertical or nearly vertical annulus which is inter- rupted by the stalk. The Dryopteridaceae are a family of perhaps 1,000 species and over 50 genera. There are 26 genera in Peru. Is is characterized by monolete spores and a chromosome number based on n = 40 or 41. The groups of genera are sometimes considered as subfamilies or families. Tryon and Tryon (1982) treated the Peruvian genera as in the following synopsis. Key to Tribes of Dryopteridaceae in Peru a. Sporangia in sori, or sporangia acrostichoid and then the fertile and sterile leaves very strongly dimorphic, the fertile with much reduced green tissue and soon withering after the spores are shed . b FIELDIANA: BOTANY b. Petiole continuous c c. Three vascular bundles, or more, near the base of the petiole Tribe Dryopterideae, genera I-XV c. Two vascular bundles near the base of the petiole .... Tribe Physematieae, genera XVI-XX b. Petiole jointed Tribe Oleandreae, genus XXI Sporangia acrostichoid and the fertile and sterile leaves more or less dimorphic, the fertile with green tissue and more or less persistent Tribe Bolbitideae, genera XXII-XXVI The key to genera has been simplified, as much segments) may have a different configuration on as possible, by including in a genus only the char- the adaxial (upper) side from that of the pinna- acters of species known from Peru or those likely rachis. to occur there. For example, the species of Wood- sia with an articulate petiole are only far north of Peru and so the genus is keyed out as having a Reference continuous petiole. In addition, genera are some- times keyed out more than once to avoid long TRYON, R. M., AND A. F. TRYON. 1982. Dryop- headings. Especially in species with decompound teridaceae, pp. 454-627, in Ferns and allied leaves, the costa (an axis bearing adnate or joined plants, Springer- Verlag, New York. Key to Genera of Dryopteridaceae a. Sporangia borne in well-defined sori, sometimes along the veins b b. Indusium scalelike and attached at the base and arching over the sorus, or attached around the base of the receptacle, or elongate and attached at the center or on one side c c. Indusium scalelike and attached at the base and arching over the sorus, or attached around the base of the receptacle d d. Indusium scalelike and attached at the base and arching over the sorus XIX. Cystopteris d. Indusium attached around the base of the receptacle XX. Woodsia c. Indusium elongate, attached at the center or usually along one side e e. Lamina 2-pinnate, the pinnules dimidiate, 3 or more vascular bundles near the base of the petiole X. Didymochlaena e. Lamina entire to 5-pinnate, if 2-pinnate then the pinnules nearly equilateral; 2 vascular bundles near the base of the petiole f f. Veins free, except as the tips are connected by a marginal strand, or veins copiously anastomosing and the lamina entire to pinnatisect, at least beyond the basal pinnae . . . g g. Sori often on both sides of a vein (especially on the basal veins of a vein-system) and then each sorus and indusium separate distally; lobes and ultimate segments (when present) entire, crenulate, or slightly and obtusely dentate; lamina entire to 3-pinnate- pinnatifid XVI. Diplazium g. Sori single on a vein, sometimes partly on the other side and then the sorus and indusium continuous distally; ultimate segments linear or strongly and sharply dentate; lamina 2-pinnate to 5-pinnate XVII. Athyrium f. Veins freely anastomosing toward the margin; lamina fully 1 -pinnate XVIII. Hemidictyum b. Indusium peltate or with a sinus and nearly orbicular to reniform, or indusium fugacious, nearly obsolete, or truly absent h h. Petiole articulate (jointed) or pinnae articulate to the rachis i i. Pinnae articulate, entire, with a large, basal, basiscopic auricle that overlies the rachis . . . V. Cyclopeltis i. Petiole articulate; lamina entire XXI. Oleandra TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 3 h. Petiole continuous (not articulate); pinnae (when present) continuous with the rachis j j. Adaxial side of the costa raised (axes of lower order, e.g., pinna-rachis or rachis, may be sulcate adaxially) k k. Veins anastomosing, or if free then the lamina pinnatifid (2-pinnatifid to 3-pinnatifid) IV. Tectaria k. Veins free and the lamina fully pinnate 1 1. Adaxial side of the pinna-rachis raised, or if sulcate then the groove not open to admit the groove of the rachis m m. Stem decumbent to erect; leaves caespitose; scales of the costae and rachis ample to abundant n n. Trichomes on the adaxial side of the axes reddish brown, with 2—4 cells, less than 0.4 mm long, catenate and usually with a blunt tip when dry; vein tips slender; basal basiscopic vein of distal pinnules arising from the costule (or very close to its juncture with the costa) I. Ctenitis n. Trichomes on the adaxial side of the axes whitish, usually with more than 4 cells, 0.5 mm or more long, terete or flat and with a pointed tip when dry; vein tips clavate; basal basiscopic vein of the distal pinnules definitely arising from the costa II. Megalastrum m. Stem long-creeping, bearing leaves at intervals; scales of the costae and rachis few or none III. Triplophyllum 1. Adaxial side of the pinna-rachis sulcate, the groove continuous with that of the rachis o o. Adaxial side of the rachis with a central ridge and a glabrous groove on each side VI. Rumohra o. Adaxial side of the rachis with lateral ridges on each side of a pubescent groove VII. Lastreopsis j. Adaxial side of the costa sulcate or nearly flat (there may be a ridge on each side of the groove) p p. Sterile and fertile leaves strongly dimorphic; each meristele of the stem surrounded by a dark sclerenchymatous sheath; indusium absent XV. Polybotrya p. Sterile and fertile leaves monomorphic or nearly so; each meristele not enclosed by a dark sclerenchymatous sheath; indusium often present q q. Lamina tissue usually pellucid-punctate (the "dots" especially visible with transmitted light), or if the tissue (rarely) lacking visible "dots" then the lamina 1 -pinnate, the indusium absent, and the pinna-base better developed on the basiscopic side XI. Stigmatopteris q. Lamina tissue uniform, not pellucid-punctate, the lamina entire to 5-pinnate, if 1 -pinnate the indusium present, or if absent then the pinna-base better developed on the acroscopic side r r. Veins mostly to wholly free; lamina pinnatifid (2-pinnatifid to 3-pinnatifid) and indusium present, or veins anastomosing and orbicular indusia often present and these with a sinus IV. Tectaria r. Veins mostly or wholly free and the lamina pinnate, or veins anastomosing and orbicular indusia (if present) peltate s s. Petiole with 2 vascular bundles near its base XVI. Diplazium s. Petiole with 3 or more vascular bundles near its base t t. Lamina 2- to 3-pinnate; ultimate segments sharply dentate, or at least the apex mucronate, or if the apex only slightly so then the margins strongly revolute XIII. Polystichum t. Lamina 1- to 5-pinnate; if 2- to 3-pinnate then the ultimate segments with nearly flat margins that are smooth to obtusely or subacutely dentate . . . u u. Veins free or mostly free v v. Indusium with a sinus, reniform to more or less orbicular, persistent VIII. Dryopteris FIELDIANA: BOTANY v. Indusium peltate, orbicular, or if with a sinus then fugacious XII. Cyclodium u. Veins anastomosing w w. Indusium absent, or if present then peltate, the pinnae coriaceous with a notably thickened margin IX. Cyrtomium w. Indusium peltate, pinnae herbaceous, the margins not thickened . . XII. Cyclodium a. Sporangia distributed over the surface of the fertile lamina or fertile segments (acrostichoid); fertile and sterile leaves dimorphic, usually strongly so x x. Pinnae articulate (jointed) y y. Veins free; paraphyses lacking XXIII. Lomariopsis y. Veins anastomosing; usually a few paraphyses scattered among the sporangia XXIV. Lomagramma x. Pinnae, when present, continuous with the rachis z z. Most pinnae or pinna-segments 3 cm long or longer, or the lamina entire and the veins anastomosing with included free veins aa aa. Veins free and close, simple or forked only near the costa; the vein tips connected by a marginal strand; sterile lamina 1 -pinnate, with the terminal segment conform with the lateral ones (pinnae) XIV. Olfersia aa. Veins free or anastomosing, a marginal strand absent, or if present then the veins fully anastomosing; sterile lamina with a pinnatifid apex to rarely with a conform apical segment bb bb. Fertile lamina more complex than the sterile, 1 -pinnate-pinnatifid or more complex, or if 1 -pinnate then with a marginal strand in the sterile segments; veins usually free XV. Polybotrya bb. Fertile and sterile lamina of equal complexity, 1 -pinnate or less complex, or if 1 -pinnate- pinnatifid then so only in the basal portions of the lamina; veins anastomosing .... XXII. Bolbitis z. Pinnae or pinna-segments ca. 0.5-1.5 cm long, or the lamina entire and the veins free, or if anastomosing then without included free veinlets cc cc. Sterile lamina simple, entire, pinnately veined, or in one rare species deeply pedately lobed XXV. Elaphoglossum cc. Sterile lamina flabellate or pinnate, or if entire then flabellately veined XXVI. Peltapteris I. Ctenitis Ctenitis (C. Chr.) C. Chr. in Verd., Man. pterid. 543. 1938. Figure 1. Dryopteris subgenus Ctenitis C. Chr., Biol. Arbejder tilegnede Eug. Warming 77. 191 1. TYPE: Dryop- teris Ctenitis (Link) Kuntze (Aspidium Ctenitis Link) = Ctenitis distans (Brack.) Ching. Plants terrestrial or sometimes epipetric. Stem decumbent to erect, provided with usually many, fibrous roots, the petiole base with a dense cluster of clathrate scales, these 6-30 mm long (although shorter and less abundant in C. nigrovenid). Leaves monomorphic or essentially so, ca. 0.2-2.5 m long, caespitose, not articulate to the stem. Lamina 1 -pinnate-pinnatifid to 4-pinnate, reduced to a pinnatifid apex, provided especially on axes with scales and "Ctenitis hairs," the latter being reddish brown, unbranched, articulate trichomes, with 2- 4 cells and 0.1-0.4 mm long. Rachis not sulcate or, if so, the adaxial groove not open to admit the groove of the pinna stalk, the adaxial ridges (if any) not continuous with the ridge of the pinna axis, scales ample to abundant. Veins free (or anas- tomosing in a few Old World species), their tips slender and usually terminating at or near the seg- ment margin, basal basiscopic vein of distal pin- nules arising from the costule or very close to its juncture with the costa. Sori roundish, borne along the veins or at their tips, receptacle somewhat el- evated, lacking paraphyses. Indusia reniform, or nearly circular and attached at the center, with a narrow sinus, persistent, fugacious, or lacking. Spores ellipsoidal, monolete, surface saccate or TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 1cm 1mm FIG. 1 . Ctenitis submarginalis: a. stem and portion of petiole; b, portion of lamina; c, base of pinna, abaxial side. Ctenitis ampla: d, portion of pinna segment, abaxial side; e, marginal trichome. (a, c from Macbride 4241, F, b from Penland & Summers 262, Ecuador, F, d, e, from Killip & Smith 24072, F.) FIELDIANA: BOTANY echinate, in some species with broad folds that might appear winglike under the light microscope. Ctenitis is an essentially pantropical genus of 70-80 species, with about half of these in the Neo- tropics. Its species grow primarily on the floor of rain forests, at lower or middle elevations, and are characterized by large decompound leaves and small, roundish sori with (when present) circular to reniform indusia. The genus has been confused with the closely related Lastreopsis, but it can be readily separated by the configuration of the ad- axial axes. In Lastreopsis ridges are continuous with the ridges on the axes of the next order above or below; but in Ctenitis these ridges are lacking or, when present, not continuous onto adjacent axes. Christensen's revisions (1913,1 920) provid- ed great insight into the characters and relation- ships of the genus, but much more work is still required to produce a satisfactory analysis of the many American species. Recently a number of species were removed from Ctenitis and placed in the genera Triplophyllum and Megalastrum. The evidence supporting this separation is even stronger than that which prompted the separation of Lastreopsis. For com- parison of these features, see discussions under treatments of Triplophyllum and Megalastrum. References CHRISTENSEN, C. 1913. A monograph of the ge- nus Dryopteris, part 1. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 7, 10: 55-282. CHRISTENSEN, C. 1920. A monograph of the ge- nus Dryopteris, part 2. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8, 6: 3-132. Key to Species of Ctenitis a. Lamina 1 -pinnate-pinnatifid, at least above the basal pinnae b b. Ultimate segments (at least of proximal pinnae) crenate or crenate-serrate, lacking marginal tri- chomes; veins of crenate segments often 1 -forked 4. C. nigrovenia b. Ultimate segments entire, with minute trichomes scattered to abundant on their margins (at least in segment sinuses); veins unbranched c c. Indusia pubescent, subpersistent; tissue between veins minutely pubescent abaxially and/or adaxially; basal veins terminating at segment margin, well above the sinus 3. C. microchlaena c. Indusia lacking, or minute and fugacious; tissue between veins glabrous (glandular in C. re- fulgens); basal veins terminating short of segment margin or reaching it at or near the sinus d d. Pinnae incised %-y2(-2/3) to the costa, with minute yellow or reddish glands scattered on (and often between) the veins abaxially; rachis and costa scales entire, not or obscurely clathrate, often with a sub-bullate base 1. C. refulgens d. Pinnae incised % or more to the costa, eglandular; rachis and costa scales remotely dentic- ulate, conspicuously clathrate, more or less flat throughout 2. C. submarginalis a. Lamina 2-pinnate-pinnatifid to 4-pinnate e e. Indusia conspicuous and subpersistent; veins and costules eglandular or with conspicuous reddish glands 5. C. ampla e. Indusia lacking, or minute and fugacious; veins and (often) costules with minute yellow glands . . 6. C. sloanei Ctenitis refulgens (Mett.) (credited in error to C. Chr. by) Vareschi, Flora Venezuela 1 : 404. 1969. Phegopteris refulgens Mett., Ann. Sci. Nat. Hot., ser. 5, 2: 240. 1864. LECTOTYPE (designated by C. Chr., p. 89. 1913): Guyana (as British Guiana), Schomburgk 1183 (B!; frag., B!). Mettenius cited four collections, two were cited as the type by Christensen, one of them, Schomburgk 1183, was also cited by Mettenius. Dryopteris refulgens (Mett.) C. Chr., Index fil. 288. 1905. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. Dryopteris refulgens var. peruviana C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd. Ser. 7, 10: 90. 1913. SYNTYPES: Peru, Monte Campana prope Tarapoto (San Martin), Spruce 4657 (B, L,); Peru, In Monte Guayrapurima, prope Tarapoto (San Martin), Spruce 4712 (w); Peru, prope Tabalosas inter urbem Moyobamba et flu- vium Rio Huallaga, Stiibel 1097 (B). ISOSYN- TYPES: Spruce 4657 (BM!, GH!, P!); Spruce 4712 (MO!, us!). Plants terrestrial. Stem erect, this and the petiole base densely clothed with orange to reddish brown, linear scales, these obscurely clathrate, about 1 cm long, with margins entire. Leaves caespitose, to 1 m long and 30 cm broad, the axes adaxially and the segment margins amply provided with "Cten- itis hairs." Lamina 1 -pinnate-pinnatifid, the tissue between the veins glabrous, or usually with scat- tered, yellow to reddish glands, the rachis and cos- tae amply provided abaxially with orange to red- dish brown, filiform, entire scales, these not or obscurely clathrate, and usually with a sub-bullate base. Pinnae incised %-!4(-2/3) to the costa, the segment entire, or sometimes serrulate at the ob- tuse or truncate apex. Veins simple, 5-8(-10) pairs per segment, typically reaching the segment mar- gin, but 1-4 basal ones terminating at or near the segment sinus (rarely merging to form a costal areole). Sori mostly inframedial between costule and margin. Indusia lacking. In and at edges of wet forests, 135-900 m, San Martin, Loreto, Huanuco. Southern Mexico (Chiapas); French Guiana to Colombia; Peru; Brazil. Christensen distinguished his var. peruviana from C. refulgens on the basis of several quanti- tative characters: pinnae less deeply incised, fewer veins running to the sinus and basal ones some- times united. He also said that the costa scales were fewer, small, and ovate. However, there are no such scales on the syntypes examined, and none of the other variable characters are correlated. Costa and rachis scales are described above as usually having a sub-bullate base. Although some of the scales are flat throughout as in the closely related C. submarginalis, the majority have their basal portion incurved so that the edges meet; hence the scale is essentially filiform with a narrowly cylindrical base, yet not with the expanded, in- flated appearance which is commonly termed bul- late. San Martin: Prov. Lamas, Alonso de Alvarado, J. Schunke V. 5941 (F). Loreto: Puerto Arturo, Rio Hua- llaga below Yurimaguas, Killip & Smith 27821 (us). Huanuco: Prov. Pachitea, Dist. Puerto Inca, D. Smith 1299 (GH, MO). 2. Ctenitis submarginalis (Langsd. & Fisch.) Ching, Sunyatsenia 5: 250. 1940. Figure la-c. Polypodium submarginale Langsd. & Fisch., Icon. fil. 12, t. 13. 1810. TYPE: Brazil, "insula Cathari- nae," Langsdorff(ho\otype, LE). Nephrodium tarapotense Hooker, Sp. fil. 4: 107. 1862. TYPE: Peru, "in Monte Campana prope Taro- poto" (San Martin), Spruce "conf. 4016" (holo- type, K!). Type not to be confused with Spruce 4016 (K, P) which is C. microchlaena. Nephrodium lagerheimii Sodiro, Crypt, vase. Quit. 252. 1893. TYPE: Ecuador, "Las pendientes occiden- tales del Tungurahua," Lagerheim (holotype, not located; isotypes, c, s!). Dryopteris submarginalis (Langsd. & Fisch.) C. Chr., Index fil. 296. 1905. Dryopteris submarginalis var. tarapotensis (Hooker) C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Na- turvidensk. Afd., ser. 7, 10: 98. 1913. Dryopteris submarginalis var. lagerheimii (Sodiro) C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Na- turvidensk. Afd., ser. 7, 10: 98. 1913. Plants terrestrial. Stem erect to decumbent, this and the petiole base densely clothed with orange or reddish brown, clathrate, linear to lanceolate scales, these 1-2 cm long, with margins entire or sparsely denticulate. Leaves caespitose, to 1 m long and 30 cm broad, the axes adaxially and the seg- ment margins abundantly provided with "Ctenitis hairs." Lamina 1 -pinnate-pinnatisect, the tissue between the veins glabrous, the rachis and costae amply provided abaxially with orange or brown, clathrate, flat, linear to filiform, remotely dentic- ulate scales. Pinnae (at least proximal ones) deeply incised to within 1-2 mm of the costa, the seg- ments entire, or serrulate at the obtuse to subacute apex. Veins simple, 8-15 pairs per segment, typ- ically reaching the segment margin, basal ones ter- minating at or near the segment sinus (or often ending short of the sinus). Sori inframedial to su- pramedial between costule and margin. Indusia lacking, or very rarely minute and fugacious. In deep, wet forests, on rocky slopes, 1 200-1 500 m, Amazonas, San Martin, and Huanuco. United States (Florida); Hispaniola; southern Mexico to Panama; Venezuela and Colombia south to Argentina and Uruguay. This is a variable species that was separated by Christensen (1913) into six infraspecific taxa. He based distinctions on shape of segments and num- ber of their veins, indusia present or lacking, po- FIELDIANA: BOTANY sition of son, and the tissue between veins gla- brous or glandular-pubescent. These characters appear to be too variable and uncorrelated for the recognition of taxa. Soral position, for example, varies considerably, even on the same lamina, from inframedial to supramedial. Often son form an elongated "V", being remote from the costule near the segment base but crowding it toward the apex. Specimens in southern South America tend to have longer, narrower segments, with acute tips and more veins; elsewhere segments are relatively broad and obtuse, yet these tendencies are too inconsistent to merit even infraspecific status. Amazonas: Prov. Bongara, Dist. Sipabamba, along Quebrada Fortuna, Young & Eisenberg 322 (MO, uc). Huanuco: Rio Huallaga Canon, below Rio Santo Do- mingo, Macbride 4241 (F, us). "[I]n Panatahuac Pro- vincia," Ruiz 68 (B). 3. Ctenitis microchlaena (Fee) Stolze, comb. nov. Aspidium microchlaena Fee, Mem. foug. 8: 102. 1857. TYPE: Mexico, Orizaba, Schaffner 459 (holo- type, presumably P, but not found; isotype, K!; photos, F & GH of K). Dryopteris microchlaena (Fee) C. Chr., Index fil. 278. 1905. Aspidium karstenii A. Braun, Ind. sem. hort. Berol. app. 3. 1867 (also published in Ann. Sci. Nat. Bot. [5] 9: 379. 1868). TYPE: Cultivated Hort. Berol. from spores of plant collected in Venezue- la, Karsten (B; frag., BM!; isotype, K!; photos, BM of B, F & GH of K). Dryopteris karstenii (A. Braun) C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 7, 10: 98. 1913. Southern Mexico; Costa Rica; Colombia; Ven- zuela; Peru. Alan Smith (Pteridophytes of Venezuela, an an- notated list. 1985, ined.) placed Dryopteris kar- stenii as a synonym of C. microchlaena of Costa Rica and Colombia. There are minor characters that differ, in the original descriptions of both spe- cies, such as glabrous, caducous indusia, and gla- brous laminar tissue in the latter species, vs. per- sistent, ciliate indusia and puberulent tissue in D. karstenii. However, an examination of type and other material of each indicates that Smith was correct. There is variation throughout the range in relative size and pubescence of indusia, and relative abundance of laminar pubescence, but none of this correlates with any other diagnostic feature. Amazonas: Prov. Bagua, along Rio Utcubamba, on Cerro Tapir, Hutchison 1482 (uc, us). San Martin: Prov. Lamas, Dist. Lamas, near Rio Chupisena, Belshaw 3503 (F, GH, MO, uc, us). Monte Guayrapurima, near Tara- poto, Spruce 4016 (K, P); this is a much different plant than Spruce "conf. 4016" (K), which is the type ofNeph- rodium tarapotense (= C. sub marginal is). 4. Ctenitis nigrovenia (Christ) Copel., Gen. fil. 1 24. 1947. Nephrodium nigrovenium Christ, Bot. Gaz. (Craw- fordsville) 20: 545. 1895. TYPE: Honduras, San- ta Barbara, San Pedro Sula, Thieme (holotype, P?; isotype, us!). Dryopteris nigrovenia (Christ) C. Chr., Index fil. 279. 1905. Plants terrestrial. Stem erect, this and petiole base densely clothed with orange, clathrate, linear to lanceolate scales, these to 1 cm long, with mar- gins entire or sparsely and minutely denticulate. Leaves caespitose, to 90 cm long and 25 cm broad, the margins and the axes on both sides abundantly provided with "Ctenitis hairs." Lamina 1-pinnate- pinnatisect, minutely glandular-pubescent on one or both of the sides, the rachis and costae provided with filiform to lanceolate scales like those of the stem. Pinnae deeply incised to within 1-2 mm of the costa. Veins simple, rarely 1 -forked, basal ones reaching the margins well beyond the segment si- nus. Sori medial to supramedial between costule and margin. Indusia orange to reddish brown, more or less pubescent, commonly persistent. On floor of wet, dark forests, ca. 1400 m, Ama- zonas and San Martin. Plants terrestrial. Stem decumbent to erect, this and the petiole abundantly provided with dark brown, clathrate, lanceolate scales, these to 4-7 mm long, subentire. Leaves caespitose, to 70 cm long and 20 cm broad, the margins glabrous, the axes adaxially provided with ''''Ctenitis hairs." Lamina 1 -pinnate-pinnatisect, abaxially provided (usually sparsely) with minute yellow or orange glands, the rachis and costae amply provided with reddish brown or blackish, linear to filiform scales, these 1-3 mm long with an expanded or sub-bul- late base. Pinnae incised near or to the costa, ul- timate segments entire or proximal ones crenate- serrate. Veins simple, or those of crenate segments usually 1 -forked, basal ones reaching the margin well beyond the segment sinus. Sori medial to in- framedial between costule and margin. Indusia minute, reddish brown, glabrous, and (in Peru) subpersistent. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. Thus far represented in Peru by one specimen: on shady river bank, 890 m, San Martin. Southern Mexico to Panama; Trinidad; Vene- zuela; Colombia to Peru. Although C. nigrovenia grows from southern Mexico to Peru, nowhere has it been collected in abundance. Distinctive features are the sparsely glandular abaxial surface and the lack of marginal trichomes. The great majority of species in the genus are amply provided with " Ctenitis hairs" on segment margins, especially at the sinus. The small indusium is usually persistent in South American representatives of this species, but is often fugacious in those from Central America. San Martin: Near Moyobamba, banks of Mayo River, Woytkowski 35268 (uc). 5. Ctenitis ampla (Willd.) Ching, Sunyatsenia 5: 240. 1940. Figure Id. Polypodium amplum Willd., Sp. pi. ed. 4, 5: 207. 1810. TYPE: "America meridionale prope Caripe" (Venezuela), Humboldt (holotype, B, Herb. Willd. 19722; photos, F, GH), not Dryopteris ampla of authors (= C. sloanei (Sprengel) Morton). Aspidium catocarpum Kunze, Linnaea 9: 95. 1834. TYPE: Peru, between Cassapi and Pampayacu (Huanuco), Poeppig, coll. July 1829 (holotype, LZ, destroyed). Aspidium nemophilum Kunze, Linnaea 9: 95. 1834. TYPE: Peru (Huanuco), Pampayacu, Poeppig, coll. July, 1 829 (holotype, LZ, destroyed). Dryopteris catocarpa (Kunze) Kuntze, Revis. gen. pi. 2: 812. 1891. Dryopteris nemophila (Kunze) C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8. 6: 57. 1920. Ctenitis nemophila (Kunze) Ching, Sunyatsenia 5: 250. 1940. Ctenitis catocarpa (Kunze) Morton, Fieldiana, Bot. 28: 12. 1951. Plants terrestrial. Stem erect, this and the petiole base densely clothed with orange to reddish brown, linear to filiform scales, these subclathrate, 1.5- 2.5 cm long, with margins entire. Leaves caespi- tose, to 2.5 m long and 80 cm broad, the axes adaxially and the margins amply provided with ^Ctenitis hairs," and abaxially with orange to red- dish brown, lanceolate, clathrate scales. Lamina deltoid, 2-pinnate-pinnatifid to (at base) nearly 4-pinnate, veins and costules abaxially provided with reddish, cylindric to elliptic glands, or eglan- dular. Veins simple to 1 -forked in the ultimate segments. Indusia large or small, but conspicuous and persistent, often with elliptic, reddish glands, or short-pubescent or glandular-pubescent. In dry, open woods or montane rainforests, on slopes, or in ravines, 380-2100 m, Amazonas to Junin and Ucayali. Venezuela; Colombia to Argentina. This is part of a species complex that has long been confused; for detailed discussion, see C. sloanei below. It is possible that C. ampla, as treat- ed here, contains two separate entities. A number of specimens in Peru, and one seen from Bolivia, have large, reddish brown, quite persistent indu- sia, these lacking marginal trichomes, but with several reddish brown, elliptic glands scattered on the surface. These glands are also conspicuous along the veins and costules abaxially. Typical C. ampla, occurring throughout the range, including Peru, has smaller indusia which are usually lighter in color, with short-pubescent or glandular-pubes- cent indusia. However, elliptic, reddish brown glands are lacking on the segments and indusia. Since the taxonomy of the entire C. sloanei/ ampla complex is in need of revision, it seems unwise to create another species or variety at this time. Amazonas: Prov. Chachapoyas, Ingenio-Pomacocha, Lopez et al. 431 1 (GH, MO, USM). San Martin: Tarapoto, Spruce 3942 (K, us). Huanuco: Muna, Bryan 530 (F, GH, us). Pasco: Prov. Oxapampa, Canyon de Huancabamba, Leon 666 (F, USM). Junin: La Merced, Killip & Smith 24072 (F, GH, us). Ucayali (as Loreto): Prov. Coronel Portillo, Boqueron, Ferreyra 16057, in part (USM). 6. Ctenitis sloanei (Sprengel) Morton, Amer. Fern J. 59: 66. 1969. Polypodium sloanei Sprengel, Syst. veg. ed. 16, 4: 59. 1827. TYPE: "In sylvis densioribus humidis Cu- bae ad Cahoba," Poeppig (holotype, LZ, de- stroyed; frag., BM!; isotypes, B!, HBG, L, P!; frag., us!; photo, us of HBG). Ctenitis ampla of authors, not Polypodium amplum Willd. 1810, = C. ampla (Willd.) Ching. Plants terrestrial. Stem erect, this and the petiole base densely clothed with orange to (en masse) reddish brown, linear to filiform scales, these sub- clathrate, 1.5-3 cm long, with entire margins. Leaves caespitose, to 1.5m long and 50 cm broad, the axes adaxially and the margins amply provided with "Ctenitis hairs," and abaxially with orange to reddish brown, lanceolate, clathrate scales. Lamina deltoid, 2-pinnate-pinnatifid to (at base) nearly 4-pinnate, provided abaxially on veins, and usually costules, with minute (to 0.5 mm) yellow- ish, cylindrical glands. Veins simple to 1 -forked in the ultimate segments. Indusia lacking, or oc- casionally minute and fugacious, short-pubescent. 10 FIELDIANA: BOTANY In lowland forests, 100-400 m, Cuzco and Ma- dre de Dios. United States (Florida); West Indies; southern Mexico (Chiapas); Nicaragua to Panama; Vene- zuela; Colombia; Ecuador; Peru. This is one of a variable complex of neotropical species whose relationships Christensen (1920) and Morton (Amer. Fern J. 59: 66. 1969) tried to clar- ify. Despite their efforts a good revision is still needed. Only a comparison of types against mass collections will suffice to delineate species and as- certain the correct application of names. There are few good characters with which to separate C. am- pla, C. sloanei, C. catocarpa, and C. nemophila, and it is not surprising that Christensen was un- certain whether all were conspecific, or were close- ly related taxa with few intermediates. Separation of C. ampla and C. sloanei has been attempted on a suite of characters: color and shape of scales, presence or lack of glands and indusia, shape of apex and stalk length of pinnules. Examination of specimens from many areas of the Neotropics in- dicates that perhaps only the indusia and laminar glands are reliable and relatively constant, but even these are at times questionable characters. Throughout the species complex indusia may be large and persistent to small and fugacious or lack- ing, and even persistent indusia can be folded and inconspicuous within a large sorus of fully ex- panded sporangia. Large, red glands on the veins can easily be seen under low magnification in many specimens of C. ampla, but the smaller yellowish ones blend with the tissue in C. sloanei and frequently can be lo- cated only with diligence and higher (30 x ) mag- nification. As treated here, C. sloanei is wide- spread in the Neotropics, yet it is rare in Peru. Ctenitis ampla is rather common in Peru, but not especially abundant elsewhere. Ctenitis sloanei tends to prefer lower elevations throughout its range (less than 500 m in the two Peruvian collections), whereas C. ampla is mostly, although not always, found above 800 m. Cuzco: Prov. La Convention, Rio Apurimac below San Martin, Davis et al. 1332 (F). Madre de Dios: Prov. Manu, Parque National del Manii, Foster 9802 (F, MO). II. Megalastrum Megalastrum Holttum, Gard. Bull. Straits Set- tlem. 39: 161. 1986. TYPE: Megalastrum vil- losum (L.) Holttum, (Polypodium villosum L.). Figure 2. Plants terrestrial. Stem decumbent to erect, oc- casionally subarborescent to 2 m tall, provided with many fibrous roots, the petiole usually with a dense cluster of clathrate scales 6-30 mm long. Leaves monomorphic, ca. 0.5-3.0 m long, caes- pitose, not articulate to the stem. Lamina 1-pin- nate-pinnatifid to 4-pinnate-pinnatifid, reduced to a pinnatifid apex, often provided on the margins and/or abaxial surface with "Ctenitis hairs," the latter being reddish brown, articulate trichomes with 2-4 cells and 0.1-0.4 mm long, but the ad- axial axes (except in one Brazilian species) bearing trichomes with usually more than 4 cells and typ- ically over 0.5 mm long, these drying terete or flat, and with pointed tips. Rachis not sulcate or, if so, the adaxial groove not open to admit the groove of the pinna stalk, the adaxial ridges (if any) not continuous with the ridge of the pinna axis, scales ample to abundant. Veins free, their tips conspic- uously clavate, ending short of the segment mar- gin, basal basiscopic vein of distal pinnules arising from the costa. Sori roundish, borne along the veins, receptacle somewhat elevated, lacking pa- raphyses. Indusia reniform, or nearly circular and attached at the narrow sinus, or lacking. Spores monolete, roundish to ellipsoidal, echinate or cris- tate. Megalastrum is a tropical genus of 45-50 spe- cies, most of these in America. It was separated from Ctenitis by Holttum (1986), a decision sub- stantiated by Smith and Moran (1987) in a paper that added new combinations for 39 species. The different trichomes on the adaxial axes is the most effective character for distinguishing the two gen- era (see key to genera of Dryopteridaceae), but there are a number of other corroborating features. In Megalastrum, the clavate vein tips terminate obviously short of the segment margin, whereas in Ctenitis the tips are not or scarcely enlarged and terminate at or very near the margin. In Ctenitis the sporangia stalks, and sometimes indusia and veins, bear unicellular, glandular trichomes, but these are lacking in Megalastrum. Although the two genera now may be effectively separated as outlined above, many problems still exist in delineating their components. Both con- tain variable species complexes in which taxa are distinguished merely by length and abundance of indument. Detailed examination of specimens throughout the Neotropics will very likely prove that a number of taxa currently recognized as spe- cies are merely varieties or forms of others. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 11 FIG. 2. Megalastrum biseriale: a, habit; b, ultimate segment, adaxial side; c, ultimate segment, abaxial side, (a, c from J. Schunke A227, us; b from Soukup 1032, F.) 12 FIELDIANA: BOTANY References CHRISTENSEN, C. 1913. A monograph of the ge- nus Dryopteris, part 1. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 7, 10: 55-282. CHRISTENSEN, C. 1920. A monograph of the ge- nus Dryopteris, part 2. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8, 6: 3-132. HOLTTUM, R. E. 1 986. Studies in the fern-genera allied to Tectaria Cav. 6. A conspectus of genera in the Old World regarded as related to Tectaria. Gard. Bull. Straits Settlem., 39: 153-167. SMITH, A. R., AND R. C. MORAN. 1987. New combinations in Megalastrum (Dryopterida- ceae). Amer. Fern J., 77: 124-130. Key to Species of Megalastrum a. Lamina 1 -pinnate-pinnatifid to (proximally) 2-pinnate; pinnules entire to crenate, or proximal ones shallowly lobed b b. Trichomes on the abaxial side of axes and veins 1-2 mm long, moderate to abundant; trichomes on segment margins to 1 mm long; stem scales often with narrow blackish margins 6. M. hirsutosetosum b. Trichomes on the abaxial side of axes sparse, 0.1-0.8 mm long, or lacking, or if abundant and to 1 mm long, then mixed with many other trichomes ca. 0.2 mm long; trichomes on segment margins less than 0.3 mm long; stem scale margins not blackish c c. Lamina chartaceous; most veins obscure; tissue on abaxial side commonly with minute (0. 1 mm), cylindrical glands d d. Sori mostly inframedial; rachis scales filiform, to 6 mm long; trichomes lacking or sparse on the axes abaxially 1 . M. honestum d. Sori mostly supramedial; rachis scales lanceolate, 2-3 mm long; trichomes abundant on the axes abaxially 2. M. yungense c. Lamina firm-herbaceous; most veins evident; tissue eglandular e e. Rachis scales scattered, 1-1.5 mm long 4. M. platylobum e. Rachis scales moderate to abundant, 2-4 mm long f f. Axes abaxially densely covered with trichomes to 0.2 mm long; abaxial surface of lamina abundantly provided with minute trichomes 3. M. microsorum f. Axes abaxially glabrous, or with scattered trichomes to 0.6 mm long; abaxial surface glabrous 5. M. biseriale a. Lamina 2-pinnate-pinnatisect to 4-pinnate-pinnatifid; pinnules regularly pinnatisect to 1 -pinnate or more complex g g. Indusia large, persistent 10. M. andicola g. Indusia lacking h h. Lamina short-pubescent, the trichomes (at least abaxially) 0.1-0.5 mm long i i. Costules, costae, and rachis abaxially glabrous, or the trichomes scattered and varying in length from 0.3 to 0.6 mm long 7. M. subincisum i. Costules, costae, and usually the rachis, densely and regularly puberulent abaxially, the trichomes 0.05-0.2 mm long 8. M. vastum h. Lamina abaxially long-pilose, especially on costae and veins, the trichomes moderate to abun- dant, mostly 0.7-1.5 mm long 9. M. pulverulentum 1 . Megalastrum honestum (Kunze) Smith & Mor- an, Amer. Fern J. 77: 128. 1987. Polypodium honestum Kunze, Linnaea 9: 49. 1834. TYPE: Peru (Huanuco), Pampayaco (Pampaya- cu), Poeppig 22 (holotype, LZ, destroyed). Polypodium fibrillosum Baker, Syn. fil. 307. 1867. TYPE: Peru (San Martin), Tarapoto, Spruce 47 42 (holotype, K!; isotypes, BM!, BR, GH!; photo, us of BR). Dryopteris fibrillosa (Baker) C. Chr., Index fil. 264. 1905. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 13 Dryopteris honesta (Kunze) C. Chr., Index fil. 271. 1905. Ctenitis honesta (Kunze) R. & A. Tryon, Rhodora 84: 127. 1982. Plants terrestrial. Stem erect, this and the petiole base densely clothed with orange, linear scales, these obscurely clathrate, to 1.5 cm long, the mar- gins entire or rarely with a few, minute, scattered setae. Leaves to 1.2 m long and 40 cm broad, the axes on the abaxial side sparsely provided with trichomes to 0.3 mm long or, more commonly, glabrous. Lamina chartaceous, the tissue and veins abaxially often with minute, scattered, yellow to reddish cylindrical glands (ca. 0.1 mm), 1-pinnate- pinnatisect, to 2-pinnate near the base, the rachis and costae abaxially provided with abundant, red- dish brown, filiform, setulose scales, these 2-6 mm long (to 4 mm on costae). Pinnae incised nearly or quite to the costae, the ultimate segments entire to crenate, with scattered marginal trichomes 0. 1- 0.2 mm long. Veins obscure. Sori mostly infra- medial. Indusia lacking. In forests, 400-1700 m, San Martin, Huanuco, Junin, Madre de Dios. Peru; Bolivia. In this and Megalastrum yungense the tissue is so dense that most veins can usually be seen only by transmitted light. The lamina of other species in the complex is much thinner and veins are mostly quite evident. Axes on the abaxial side typically lack trichomes, although filiform scales are quite abundant. On only one of all Peruvian collections examined (Ucayali, cited below) was a sparse covering of minute trichomes detected. Huanuco: Pampayacu, Kanehira 179 (us). La Divi- soria, 25 km NE of Tingo Maria, Moran 3707 (USM). Junin: Schunke Hacienda above San Ramon, Killip & Smith 24659 (us). Chanchamayo Valley, C. Schunke 957 (F). Ucayali: Fundo Chela, Sinchono (as Loreto or Hua- nuco), Aguilar, Aug. 3, 1948 (GH, USM). Madre de Dios: Prov. Manu, Rio Sotileja, Parque Nacional Manu, Foster etal. 11598(F). 2. Megalastrum yungense (Christ & Rosenst.) Smith & Moran, Amer. Fern J. 77: 1 29. 1987. Dryopteris yungensis Christ & Rosenst., Repert. Spec. Nov. Regni Veg. 5: 234. 1908. TYPE: Bolivia, Sirapuya near Yanacachi, 2000 m, Buchtien 493 (holotype, P!; isotype, us). Plants terrestrial. Stem erect, this and the petiole base densely clothed with brown (sometimes mot- tled with black), linear scales, these scarcely or obscurely clathrate, ca. 6 mm long, the margins setulose. Leaves to 80 cm long and 25 cm broad, the axes on the abaxial side amply provided with trichomes 0.2-0.6 mm long. Lamina chartaceous, the tissue and veins abaxially often with minute, scattered, cylindrical glands (ca. 0.1 mm), 1-pin- nate-pinnatisect, to 2-pinnate near the base, the rachis and costae abaxially amply provided with reddish brown, setulose, lanceolate scales, these 2-3 mm long (less on costae). Pinnae incised near- ly or quite to the costae, the ultimate segments entire, obtuse, with scattered marginal trichomes 0.1 mm long. Veins obscure. Sori mostly supra- medial. Indusia lacking. Thus far known in Peru from one collection, in high montane forest, 1 800 m, San Martin. Peru; Bolivia. Rachis scales in this species complex are typi- cally filiform, many times longer than broad; but those of M. yungense are lanceolate to narrow- deltoid. Scales of the stem and petiole base are also distinctive, at least in the specimen from Peru: essentially dark brown, but with scattered black cells or groups of cells. Because the type consists only of a leaf broken off above the stem, it is not possible to ascertain if this color pattern is con- sistent. San Martin: Prov. Rioja, Pedro Ruiz-Moyobamba Rd., km 390, D. Smith 4424 (uc). 3. Megalastrum microsorum (Kuntze) Stolze, comb. nov. NephrodiummicrosorumHoo]/ieT,Sp.fil.4: 106. 1862, nom. illeg., not Endl. 1833. TYPE: Ecuador, at the foot of Chimborazo, Spruce (holotype, K!; frag., BM!; photos, F & GH of K). Dryopteris microsora Kuntze, Revis. gen. pi. 2: 813. 1891, nom. nov. for Nephr odium microsorum Hooker and with the same type. Dryopteris leptosora C. Chr., Index fil. 274. 1905, nom. nov. for Nephrodium microsorum Hooker and with the same type. Megalastrum leptosorum (C. Chr.) Smith & Moran, Amer. Fern J. 77: 128. 1987. Plants terrestrial. Stem erect, this and the petiole base densely clothed with orange to brown, linear scales, these narrowly clathrate, to 2 cm long, the margins conspicuously setose, the setae often bi- furcate. Leaves to 1 m long and 30 cm broad, the axes on the abaxial side abundantly provided with whitish, pluricellular trichomes 0.1-0.2 mm long. 14 FIELDIANA: BOTANY Lamina firm-herbaceous, the abaxial surface abundantly and minutely pubescent, 1-pinnate- pinnatisect to 2-pinnate, the rachis and costae abaxially provided with abundant reddish brown, filiform, setulose scales, these 2-4 mm long. Pin- nae incised nearly or quite to the costae, the ul- timate segments entire to crenate, with scattered marginal trichomes ca. 0.1 mm long. Veins evi- dent, although sometimes indistinct. Sori supra- medial to somewhat inframedial. Indusia lacking. Thus far known in Peru from the single speci- men cited below: beneath a moist rock wall, 2800 m, Junin. Ecuador; Peru. Megalastrum microsorum can be distinguished from its nearest relatives by the abundant, though minute, pubescence of the abaxial surface, on and between the veins. An interesting feature is the character of the stem, petiole, and (sometimes) rachis scales. These are conspicuously setulose as in many species of the genus, but the setae are commonly bifid at the apex. This condition also has been observed on petiole scales of M. platy- lobum, although only rarely. Junin: Paucartambo, Woytkowski 6742 (us). 4. Megalastrum platylobum (Baker) Smith & Moran, Amer. Fern J. 77: 128. 1987. Polypodium platylobum Baker, Syn. fil. 307. 1867. TYPE: Mt. Guayrapurima, near Tarapoto (San Martin), Spruce 4656 (holotype, K!; isotypes, BM!, K!, P!). Polypodium tarapotense Baker, Syn. fil. ed. 2:505. 1 874. TYPE: based on one sheet of type of P. platylo- bum (K.!, see discussion below). Dryopteris platyloba (Baker) C. Chr., Index fil. 285. 1905. Dryopteris tarapotensis (Baker) C. Chr., Index fil. 297. 1905. Plants terrestrial. Stem erect, this and the petiole base densely clothed with orange to brown, linear scales, these narrowly and obscurely clathrate, to 1 cm long, the margins setulose, the setae occa- sionally bifid at apex. Leaves to 1.5 m long and 40 cm broad, the axes on the abaxial side densely covered with whitish trichomes of mixed lengths, many ca. 1 mm long, and many more only 0.1 mm long. Lamina firm-herbaceous, 1-pinnate- pinnatisect, or 2-pinnate near the base, occasion- ally bearing some scattered, minute (0.05 mm) trichomes on the tissue abaxially, the rachis abax- ially provided with scattered, reddish brown, fi- liform scales, these 1-1.5 mm long and with se- tulose margins. Pinnae incised nearly or quite to the costae, the ultimate segments entire to crenate or rarely shallowly lobed, their margins amply cil- iate with trichomes ca. 0.5 mm long. Veins evi- dent. Sori supramedial. Indusia lacking. Thus far known in Peru only from the type col- lections, San Martin. Venezuela; Peru. At Kew there are five sheets of Spruce 4656. Three, from Hooker's herbarium, represent the holotype of Polypodium platylobum; the two oth- ers, in another folder and marked "sheet 1 " and "sheet 2," are isotypes. After having described P. platylobum, Baker seven years later described P. tarapotense, based on one of the sheets of Spruce 4656, which contains the distal portion of the leaf, plus the petiole of a species of Cyatheaceae (Cnem- idaria speciosal). For a full discussion, see C. Christensen's monograph (1913, p. 110). Megalastrum platylobum can be further distin- guished from closely related species by the mixed trichomes on the axes abaxially, especially on the costae: many spreading pluricellular ones ca. 1 mm long, plus a dense covering of 1 (-2)-cellular ones ca. 0.1 mm long. Axes trichomes of M. hirsuto- setosum are all long (1-2 mm), and those of others in the species complex are short (commonly 0.3- 0.5 mm). 5. Megalastrum biseriale (Baker) Smith & Mo- ran, Amer. Fern J. 77: 127. 1987. Figure 2. Polypodium biseriale Baker, Syn. fil. 309. 1867. TYPE: Ecuador, Mt. Tungurahua, Spruce (holotype, K!). Dryopteris biserialis (Baker) C. Chr., Index fil. 254. 1905. Ctenitis biserialis (Baker) Lell., Fern gaz. 11: 108. 1975. Plants terrestrial. Stem erect, this and the petiole base densely provided with orange or light brown, linear scales, these obscurely clathrate, to 1.5 cm long, the margins setulose. Leaves to 1 m long and 30 cm broad, the axes on the abaxial side sparsely provided with whitish trichomes 0.2-0.6 mm long, or glabrate. Lamina firm-herbaceous, the tissue glabrous and eglandular, 1 -pinnate-pinnatisect to 2-pinnate, the rachis and costae abundantly pro- vided on the abaxial side with reddish brown, fi- liform scales, these 3-4 mm long (on costae 1-2 mm), margins remotely setulose. Pinnae incised nearly or quite to the costae, ultimate segments TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 15 entire to crenate or shallowly lobed, with scattered marginal trichomes 0. 1 mm long. Veins evident. Sori mostly inframedial. Indusia lacking. In forests and wooded ravines, 600-1800 m, Pasco, Junin, Madre de Dios. Panama; Colombia; Ecuador; Peru. Pasco: San Nicolas, Pichis Trail (as Junin), Killip & Smith 26021 (us). Junin: Chanchamayo Valley, C. Schunke 45, 87 (F, us). Schunke Hacienda, above San Ramon, C. Schunke A227 (us). La Merced, Chancha- mayo, Soukup 1032 (F). Madre de Dios: Prov. Manu, Atalaya, Foster & Wachter 7423 (F, MO). 6. Megalastrum hirsutosetosum (Hieron.) Smith & Moran, Amer. Fern J. 77: 128. 1987. Dryopteris hirsute- setosa Hieron., Hedwigia 46: 343, t. 6. 1907. TYPE: Ecuador, plateau above All- payacu between Banos and Jivaria de Pintuc, Stu- bel 903 (holotype, B!; frag. & photo, BM). Ctenitis hirsuto-setosa (Hieron.) Lell., Proc. Biol. Soc. Wash. 89: 709. 1977. Plants terrestrial. Stem erect, provided at apex with a dense cluster of deep orange to reddish brown scales, these obscurely clathrate, to 1 cm long, with narrow, often blackish, densely setulose margins. Leaves caespitose, to 1.2 m long and 30 cm broad, the axes and veins on both sides mod- erately to abundantly provided with rigid, pluri- cellular trichomes 1-2 mm long. Lamina 1-pin- nate-pinnatisect to 2-pinnate, the axes lacking scales, or the rachis with a few, scattered, filiform ones. Pinnae incised nearly or quite to the costae, the ultimate segments subentire, crenate or very shallowly lobed, the margins with abundant spreading trichomes to 1 mm long. Veins indis- tinct to obscure, 1 -several-forked. Sori usually me- dial between costule and margin. Indusia lacking. In forests, 1000-1500 m, Amazonas, Huanuco, Pasco, Junin, Ucayali. Colombia; Ecuador; Peru. This can be easily distinguished from all the less dissected species of Megalastrum by the abundant, spreading trichomes which clothe the axes and veins, abaxially and adaxially. There is a single specimen from San Martin (Knapp & Mallet 7022, MO) that closely resembles M. hirsutosetosum, in that the axes and margins have abundant, long trichomes. However, the tissue is glandular abax- ially, and the axes are conspicuously filiform-scaly. The few scattered sporangia all appear to be bar- ren, so the plant is probably a hybrid, perhaps involving M. platylobum. j-v Amazonas: Prov. Bagua, 12 km E of La Peca, Barbour 2499 (F, MO). Huanuco: Prov. Leoncio Prado, Tingo Maria, Aguilar 25 (USM). Pampayacu, Kanehira 184 (GH). Pasco: Pichis Trail, San Nicolas (as Junin), Killip & Smith 26037 (us). Junin: La Merced, Chanchamayo, C. Schunke 22 (uc). Ucayali: Prov. Coronel Portillo (as Loreto), Sin- chono, between Tingo Maria and Pucallpa, Aguilar 865 (USM). 7. Megalastrum subincisum (Willd.) Smith & Moran, Amer. Fern J. 77: 129. 1987. Polypodium subincisum Willd., Sp. pi. ed. 4, 5: 202. 1810. TYPE: Venezuela, Caracas, Bredemeyer (holotype, B, Herb. Willd. 19701; photos, F, us). Dryopteris subincisa (Willd.) Urban, Symb. antill. 4: 19. 1903. Ctenitis subincisa (Willd.) Ching, Sunyatsenia 5: 250. 1940. Plants terrestrial. Stem erect, this and the petiole base densely clothed with bright brown to casta- neous, linear to filiform scales, these clathrate, 1.5- 2.5 cm long, the margins setose. Leaves to 3 m long and 1.2m broad, the axes adaxially provided with a dense covering of terete, curved trichomes mostly 0.5-0.8 mm long. Lamina 2-pinnate-pin- natisect to 3-pinnate-pinnatifid, the costules, cos- tae, and rachis abaxially glabrous, or with tri- chomes scattered and varying in length from 0.3 to 0.6 mm long, the scales of the rachis and costae dark brown, clathrate, linear or filiform (from a sometimes dilated base) and mostly 2-6 cells wide, the margins commonly setose. Pinnae with tissue between the veins typically glabrous abaxially, but occasionally sparsely to abundantly puberulent. Sori medial to inframedial on the segments. In- dusia lacking. In rain forests and wooded ravines, 500-2 1 00 m, Amazonas, San Martin, Junin, Cuzco. West Indies; southern Mexico to Panama; Ven- ezuela; Colombia to Bolivia. This is one of the most widely distributed spe- cies in the genus, and is part of a complex that is rather variable in laminar indument. This group includes M. vastum from Peru and the Andes and M. spectabile (Kaulf.) Smith & Moran from south- ern South America. Christensen stated (1920, p. 68) that M. vastum might be merely a variety of M. subincisa; the other species appears to differ in no greater degree. The only diagnostic features, as 16 FIELDIANA: BOTANY noted in the key, are those of size and abundance of trichomes and shape and margin of scales, and there is some variability even in these. Although pubescence is generally confined to the axes and veins on the above species, some speci- mens are sparsely to abundantly puberulent on the tissue between the veins. This condition has been noted in both M. subincisum and M. vastum. The entire species complex is in need of study through- out its range; meanwhile the two species are main- tained as distinct for purposes of this treatment. Amazonas: Prov. Bagua, valley of Rio Maranon above Cascadas de Mayasi, Wurdack 1851 (us). San Martin: "In Monte Campana, prope Tarapoto," Spruce 4340 (K). Junin: Schunke Hacienda, above San Ramon, C. Schunke A225 (us). Cuzco: Prov. Urubamba, along Rio Urubam- ba near town of Machu Picchu, Tryon & Try on 5412 (F, GH). Prov. Paucartambo, Kosnipata-Pilcopata, Vargas 11284(GH). . Megalastrum vastum (Kunze) Smith & Moran, Amer. Fern J. 77: 129. 1987. Polypodium vastum Kunze, Linnaea 9: 50. 1834. TYPE: Peru (Huanuco), "inter Pampayaco (Pam- payacu) et Cocheros et ad Ventanilla de Cassapi," Poeppig 217 (holotype, B!; isotypes, B!, L; photos, F & us of L). Dryopteris vasta (Kunze) Hieron., Hedwigia 46: 347. 1907. Dryopteris mollicoma C. Chr., Kongel. Danske Vi- densk. Selsk. Skr. Naturvidensk. Afd. ser. 8, 6: 75. 1920. TYPE: Ecuador, "in silv. suband. ori- ent." Oyacachi, Sodlro (holotype, P; frag., BM; iso- type, A!; photo, F of BM). Ctenitis mollicoma (C. Chr.) Ching, Sunyatsenia 5: 250. 1940. Megalastrum mollicomum (C. Chr.) Smith & Moran, Amer. Fern J. 77: 128. 1987. Plants terrestrial. Stem erect, this and the petiole base densely clothed with bright brown to casta- neous, lanceolate to filiform scales, these clathrate, 1-3 cm long, the margins denticulate to setulose. Leaves to 2 m long and 0.8 m broad, the axes adaxially provided with curved trichomes mostly 0.5-0.8 mm long or glabrescent. Lamina 2-pin- nate-pinnatisect to (proximally) 3-pinnate-pin- natifid, the costules, costae, and (often) the rachis densely and regularly puberulent abaxially, the tri- chomes 0.05-0.2 mm long, the scales of the rachis and costae dark brown, clathrate or subclathrate, lanceolate to filiform and mostly 3-10 cells wide, the margins entire to denticulate or sometimes setose. Pinnae with tissue between the veins typ- ically glabrous abaxially, but sometimes sparsely to abundantly puberulent, or occasionally hirsute, adaxially. Sori typically medial. Indusia lacking. Dense forests, 750-1850 m, Amazonas, Hua- nuco, Junin, and Ayacucho. Costa Rica; Colombia to Bolivia; Paraguay. Killip & Smith 22779, cited below, differs from typical M. vastum in the dense puberulence abax- ially on the tissue between the veins. In the M. subincisum complex, some scattered trichomes are often evident on the laminar tissue, but never so conspicuously as seen on this particular specimen. This character should be considered, along with other conditions of indument, in future studies of the various species complexes of both Ctenitis and Megalastrum. Probably M. vastum should be recognized mere- ly as a variety of M. subincisum, under which see further discussion. Only the type and a few other collections have been found in Peru; the species seems to be most abundant in Ecuador. Amazonas: Prov. Bagua, 1 2 mi E of La Peca, Barbour 2504 (F, MO). Huanuco: Cushi, Macbride 4846, 4855 (F, us). Junin: Prov. Chanchamayo, Chilpex, 26 km S of San Ramon, Smith & Palacios 2639 (F, MO). Ayacucho: "Aina" (Ayna), between Huanta and Rio Apurimac, Kil- lip & Smith 22779 (us). 9. Megalastrum pulverulentum (Poiret) Smith & Moran, Amer. Fern J. 77: 129. 1987. Polypodium pulverulentum Poiret in Lam., Encycl. 5: 555. 1804. TYPE: Plumier, Traite foug. Amer. t. 34. 1705, based on a plant from Hispaniola. Polypodium karstenianum Klotzsch, Linnaea 20: 390. 1847. TYPE: "Colombia" (Venezuela), Karsten II, 3 (holotype, B; probable isotype, HBG; photo, F & us of HBG). Dryopteris karsteniana (Klotzsch) Hieron., Hedwigia 46: 348. 1907. Dryopteris pulverulenta (Poiret) C. Chr., in Urban, Symb. antill. 9: 305. 1925. Ctenitis pulverulenta (Poiret) Copel., Gen. fil. 124. 1947. Ctenitis karsteniana (Klotzsch) Vareschi, Flora Ven- ezuela 1: 405. 1969. Plants terrestrial. Stem erect, arborescent, in Peru to 1 m tall, this and the petiole base densely clothed with orange to brown, linear scales, these sub- clathrate, 1-2.5 cm long, the margins entire to remotely setulose. Leaves to 3 m long and 1.5 m broad, the axes on both sides moderately to abun- dantly pilose. Lamina 3-pinnate to nearly 4-pin- nate-pinnatifid, the rachis deciduously pilose and with scattered scales 1-2 mm long, these with con- TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 17 spicuously setose margins. Pinnae commonly 2-pinnate-pinnatisect, but basal ones often basi- scopically enlarged and more highly dissected, the costae and veins moderately to abundantly pro- vided with spreading pluricellular trichomes mostly 0.7-1.5 mm long or a little shorter adaxially. Sori inframedial, medial, or rarely supramedial. In- dusia lacking. In forests, 540-1800(-2700) m; San Martin, Cuzco. Greater Antilles; southern Mexico to Costa Rica; Venezuela; Colombia to Bolivia. This, like M. hirsutosetosum, is distinguished by the long (1-2 mm), spreading trichomes on the axes and veins. However, leaves of the latter are never more than 2-pinnate, and scales of the pet- iole base have blackish, densely setulose margins. Leaves of M. pulverulentum are often 4-pinnate- pinnatisect at base, and the petiole scales are con- colorous, with subentire to remotely denticulate margins. The few specimens found thus far in Peru are var. pulverulentum, a common variety widely dis- tributed throughout the Neotropics. Variety hey- dei (C. Chr.) Smith & Moran, with essentially gla- brous laminae, is confined to Guatemala (or possibly also in Ecuador). Young & Leon 4958, cited below, appears to be intermediate between M. pulverulentum and the Central American M. pansamalense (C. Chr.) Smith & Moran, the latter differing chiefly in its denser covering of long trichomes and its broad, entire laminar scales. The Young & Leon collec- tion has much smaller leaves, and abundant lan- ceolate to ovate scales with entire margins, but is similar in every other character to M. pulverulen- tum. Although fully mature, the cited specimens have leaves only 50 cm long and 18 cm broad, whereas both of the above species have leaves from 1 to 3 m long. It is possible that Young & Leon 4958 (2650-2750 m) represents a depauperate specimen or a high elevation variant; it is uncom- mon to find M. pulverulentum above 1500 m. San Martin: Tarapoto, Spruce 47 18 (BM, K, p). Prov. Mariscal Caceres, Rio Abiseo National Park, Young & Leon 4958 (F, GH). Cuzco: San Miguel, Urubamba Val- ley, Cook & Gilbert 1128 (us). Prov. Paucartambo, Ha- cienda Villa Carmen, Vargas 14685 (GH). Department unknown: Peru, undesignated locality, Soukup 296 (F). 10. Megalastrum andicola (C. Chr.) Smith & Moran, Amer. Fern J. 77: 127. 1987. Plants terrestrial. Stem erect, this and the petiole base densely clothed with orange to brown, linear to lanceolate scales, these subclathrate, 1-2.5 cm long, the margins setulose. Leaves to 1.5 m long and 40 cm broad, the axes on both sides amply to abundantly pubescent. Lamina 2-pinnate-pinnati- sect to 3-pinnate-pinnatifid, the rachis and costae sparsely to amply scaly, the scales orange to brown, linear, 4-5 mm long, their margins denticulate to subentire. Pinnae mostly 1 -pinnate-pinnatisect, but basal ones more deeply divided and enlarged at the base basiscopically, the ultimate segments mi- nutely pubescent on both sides on the veins, and sometimes on the intervening tissue. Indusia per- sistent, light to dark brown, 1 mm or more in diameter, often glandulose. This is part of a variable complex of neotropical species that is in need of further study. It is one of the few indusiate species of Megalastrum, and the only indusiate one in Peru. A number of va- rieties and forms have been described, based chief- ly on differences in length and abundance of in- dument. Megalastrum andicola is probably only varietally distinct from the West Indian M. vil- losum (L.) Holttum; the latter apparently differs only in its larger size, greater dissection of lamina, and in the fewer and early deciduous scales on the axes. Pending future study of the species complex, M. andicola and one other form are tentatively recognized in Peru. Key to Forms of Megalastrum andicola a. Trichomes on axes, veins, and leaf tissue ca. 1 mm long lOa. f. andicola a. Trichomes on axes, veins, and leaf tissue 0.2-0.3 mm long lOb. f. lehmannianum 18 FIELDIANA: BOTANY lOa. Megalastrum andicola f. andicola Nephrodium villosum var. opacum Hieron., Bot. Jahrb. Syst. 34: 446. 1904, LECTOTYPE (designated here): Colombia, Fusagasuga, Lindig 159 (B). Nephrodium villosum f. spruceanum Hieron., Bot. Jahrb. Syst. 34: 446. 1904. LECTOTYPE (des- ignated here): Ecuador, Spruce 5295 (B!; isotypes, c, K!). Dryopteris andicola C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd. ser. 8, 6: 88. 1920. TYPE: Ecuador, Spruce 5295 (holotype, B!; iso- types, c, K!). Dryopteris andicola f. spruceana (Hieron.) C. Chr., reference as above, p. 89. Ctenitis andicola (C. Chr.) Ching, Sunyatsenia 5: 250. 1940. In rain forests and cloud forests, 750-2300 m, Amazonas, San Martin, Junin, and Pasco. Colombia; Ecuador; Peru. Amazonas: Prov. Bagua, 12 mi E of La Peca, Barbour 2587 (F, uc). Prov. Bongara, hills 1-5 km SSE of Yam- brasbamba, Wurdack 1030 (GH, us). San Martin: Tara- poto, LI. Williams 5985 (F). Pasco: Oxapampa, Soukup 3354 (GH, us). Junin: Prov. Tarma, Valle del Rio Chan- chamayo, Esposto 663 (USM). lOb. Megalastrum andicola f. lehmannianum (Hieron.) Stolze, comb. nov. Nephrodium villosum f. lehmannianum Hieron., Bot. Jahrb. Syst. 34: 446. 1904. LECTOTYPE (des- ignated here): Colombia, Lehmann 7369 (holo- type, B; isolectotypes, K!, us!). Dryopteris andicola f. lehmanniana (Hieron.) C. Chr., Bot. Jahrb. Syst. 34: 89. 1904. Thus far represented in Peru by the single col- lection cited below: Podocarpus forest, 1 880-1 950 m, Junin. Colombia; Ecuador; Peru. Hieronymus originally distinguished his new form by size of indusia and number of sori on a segment. It has been seen that these characters are too variable and inconsistent to be of taxonomic value. The only other difference observed between f. lehmannianum and f. andicola is the length of trichomes on the lamina, but at least this character is consistent and conspicuous. The tissue, veins, and minor axes of f. andicola are densely covered on both sides with whitish, spreading trichomes ca. 1 mm long. In f. lehmannianum equivalent axes are as densely pubescent, but the trichomes are never more than 0.3 mm long, and those on the tissue abaxially are noticeably less abundant. Junin: Prov. Chanchamayo, Rio Rondayacu, 45 km from San Ramon, D. Smith el al. 2615 (F, GH, MO). III. Triplophyllum Triplophyllum Holttum, Kew Bull. 41: 239. 1986. TYPE: Triplophyllum protensum (Sw.) Holt- tum (Aspidium protensum Sw.). Figure 3. Plants terrestrial. Stem creeping, sparsely to moderately provided with narrow, nonclathrate scales, these commonly 1—4 mm long. Leaves 1 -pinnate-pinnatifid to nearly 4-pinnate (as to bas- al portion), borne at approximate to subdistant intervals, monomorphic, to ca. 1 .6 m long and 0.8 m broad, long-petiolate, not articulate to the stem. Lamina deltoid-pentagonal (to somewhat elongate in a few Old World species), the basal pinnae con- spicuously larger than adjacent ones, and more than half as long as the rest of the lamina, provided especially on the axes with scales and ''''Ctenitis hairs," the latter being reddish brown, un- branched, articulate trichomes, with 2-4 cells and 0.1-0.4 mm long. Rachis not sulcate or, if slightly so, the adaxial groove not open to admit the groove of the pinna stalk, the adaxial ridges (if any) not continuous with the ridge of the pinna axis, scales few or lacking. Veins free, or in a few species rarely anastomosing, but without free veins in the are- oles. Sori roundish, borne along the veins or at their tips, receptacle somewhat elevated, lacking paraphyses. Indusia reniform (in Peru species) or rarely lacking. Spores monolete, roundish to ellip- soidal, the surface bearing thin, winglike ridges. Triplophyllum is a genus of 20 tropical species, five of them in America, with but a single variety occurring in Peru, and one species perhaps to be expected. The more obvious characters separating it from Ctenitis are seen in the key to genera of Dryopteridaceae. In addition, Ctenitis often has cylindric glands on indusia, and on axes and tissue between the veins. If glands are present on the lamina or indusia in Triplophyllum, they are spherical. According to Holttum ( 1 986) there are two oth- er varieties of T.funestum: var. perpilosum Holtt. of Colombia, with many trichomes between the veins on both surfaces, and var. hirsutum Holtt. of Guyana, with trichomes to 1 mm long on the axes. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 19 5 cm FIG. 3. Triplophyllum funestum var. funestum: a, habit; b, petiole scale; c, ultimate segment, abaxial side (a b from Petelot s.n., Brazil, F; c from Klug 174, F.) 20 FIELDIANA: BOTANY Reference HOLTTUM, R. E. 1986. Studies in the fern-genera allied to Tectaria Cav., 5. Triplophyllum, a new genus of Africa and America, Kew Bull., 41: 237-260. 1 . Triplophyllum funestum (Kunze) Holttum var. funestum, Kew Bull. 41: 256. 1986. Figure 3. Aspidiumfunestum Kunze, Linnaea 9: 96: 1 834. TYPE: "ad Ega, Brasiliae, lect. 1 832," Poeppig(ho\otype, presumably w). Nephrodium funestum (Kunze) Hooker, Sp. fil. 4: 129, t. 259. 1862. Dryopteris protensa var.funesta (Kunze) C. Chr., Kon- gel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8, 6: 91: 1920. Ctenitis protensa (Sw.) Ching, var. funesta (Kunze) Proctor, Rhodora 63: 34. 1961. Plants terrestrial. Stem creeping, this and the petiole and rachis sparsely to moderately provided with reddish brown, lanceolate scales, these non- clathrate, 1—4 mm long. Leaves approximate to somewhat spaced along the stem, to 1 m long and 50 cm broad, with ''''Ctenitis hairs" ample on the axes and few on the veins and segment margins. Lamina deltoid-pentagonal, 1 -pinnate-pinnatifid to 2-pinnate-pinnatifid, but 3-pinnate-pinnatifid as to the greatly enlarged basal pinnae. Central pinnae commonly 1 -pinnate-pinnatifid, the basal pinnules more strongly developed acroscopically, ultimate segments obtuse, the basal pinnae often nearly as large as the rest of the lamina, much more strongly developed basiscopically. Veins branched in the ultimate segments, terminating at or near the margin, with slender tips. Sori borne near the segment margins. Indusia orange to red- dish brown, conspicuous, persistent, with minute pluricellular trichomes. In wet soil of rain forests, sea level to 700 m, San Martin, Loreto, Pasco, and Madre de Dios. Puerto Rico; Lesser Antilles; Nicaragua to Pan- ama; Trinidad; the Guianas to Colombia, and southward to Brazil and Bolivia. Some specimens from Peru approach T. acuti- lobum Holttum, a large species of Brazil, with bas- al pinnae 44 cm long or more, axes adaxially gla- brous, and segments and lobes mostly acute. Allard 22003 and Vargas 18629 (cited below) lack ad- axial trichomes and have some segments acute, but both have small leaves with basal pinnae about 1 5 cm long. Other specimens examined have one or the other of these features, but not both. Ob- viously these characters are too variable to serve as a basis for the further division of var. funestum. San Martin: On ridge east of Tingo Maria, Allard 22003 (GH, us). Loreto: Mishuyacu, near Iquitos, King 174 (F, us). Prov. Maynas, Rio Momon, Sounders 1365, 1367, 1368 (GH). Pasco: Prov. Oxapampa, Quebrada Castilla on Omaiz River, Leon & Young 1020 (GH). Ma- dre de Dios: Prov. Tambopata, Tambopata Nature Re- serve, Harbour 4921, 5189 (F). Prov. Tambopata Vargas 18629 (GH). Comments Triplophyllum dicksonioides (Fee) Holttum, Kew Bull. 4: 257. 1986. Aspidium dicksonioides Fee, Crypt vase. Bresil 1 : 143, t. 49. 1869. TYPE: Northern Brazil, Rio Negro near San Gabriel, Spruce 2129 (holotype, not lo- cated; isotypes, BM, K), cited in error (fide Chris- tensen) as Glaziou 2129. Dryopteris protensa (Sw.) C. Chr. var. dicksonioides (Fee) C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8, 6: 93: 1920. This species occurs from the Guianas to Colom- bia, and in northwest Brazil, so there is good rea- son to believe it may be found in Amazonian Peru, perhaps in Loreto where T. funestum grows. It is distinguished from the latter by the tiny, spherical glands on the indusia and lamina, by the sori, which are mostly apical on the veins, and by the more highly dissected lamina (up to 4-pinnate) with much smaller ultimate segments. In contrast, T. funestum var. funestum is minutely pubescent on the veins and indusia, but eglandular; sori are seldom apical on the veins, and the lamina is 1 -pinnate-pinnatifid to (in basal pinnae) 3-pin- nate-pinnatifid. IV. Tectaria Tectaria Cav., Anales Hist. Nat. 1: 115. 1799. TYPE: Tectaria trifoliata (L.) Cav. (1802) (Polypodium trifoliatum L.). Figure 4. Aspidium Sw., J. Bot. (Schrader) 1800(2): 4, 29. 1802, nom. superfl. for Tectaria and with the same type. Stem erect or decumbent, usually stout, to mod- erately long-creeping and slender, bearing scales. Leaves ca. 1 0 cm to 2 m long, petiole continuous. Lamina simple and entire to deeply lobed, or to TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 21 FIG. 4. Tectaria incisa var. incisa: a, portion of stem and petiole; b, lamina; c, portion of pinna, abaxial side, (a from Tryon & Tryon 5206, F, b from Alfaro s.n., Costa Rica, F, c from Moran 3653, F.) 22 FIELDIANA: BOTANY 3-pinnate-pinnatifid, glabrous, scaly, or pubes- cent, usually monomorphic, or dimorphic with the fertile usually more erect, longer and less expanded than the sterile, pinnae not articulate, adaxial side of the costa raised. Veins usually anastomosing, usually with included free veinlets, rarely free. Sori (in Peru) usually roundish, borne on the veins and covered by reniform to peltate (rarely some elon- gate) indusia, or exindusiate and roundish or rarely some elongate, not paraphysate (elsewhere the son may be linear, rarely on a marginal projection or the sporangia partly acrostichoid). Spores ellip- soidal, monolete, ridged, cristate or echinate. Tectaria is a large genus of about 150 species, some 20 of them in America and six in Peru. It is pantropical and sometimes subtropical. The sorus and lamina architecture is unusually variable in the genus, and several segregate genera have been recognized on the basis of these variations. Tryon and Tryon (1982) list the ones that occur in Central America, the Greater Antilles, and northern South America. Tectaria is a difficult genus in the Andes because of a lack of critical studies, the uncertainty of the application of some names, the probability of un- recognized hybrids, and the lack of cytological in- formation. This treatment, based largely on Pe- ruvian materials, will need to be refined, especially as field studies of populations and cytological in- formation on the different morphological kinds become available. The cytological complexities of American Tectaria, with diploids, triploids, and tetraploids, are partly indicated by Jermy and Walker (1985 and references). Soral variation is shown in Mexia 8217, MO (T. antioquoiand) and Smith et al. 1709, MO (T. li- zarzaburui); in both, the sorus and the indusia vary from roundish to reniform to quite elongate. References JERMY, A. C., AND T. G. WALKER. 1985. Cyto- taxonomic studies of the ferns of Trinidad. Bull. Brit. Mus. (Nat. Hist.) Bot., 13: 133-286. MORTON, C. V. 1966. The Mexican species of Tectaria. Amer. Fern J., 56: 120-137. TRYON, R. M., AND A. F. TRYON. 1 982. Tectaria, pp. 470-48 1 , in Ferns and allied plants, Spring- er-Verlag, New York. Key to Species of Tectaria a. Veins mostly or wholly free 1 . T. brauniana a. Veins fully anastomosing, except for included free veinlets b b. Lamina 1 -pinnate or more complex, or the rachis fully alate, or if the lamina simple then with a pair of basal lobes and an abrupt base c c. Sori of the lateral pinnae or large pinna-segments in 2 series between the costules, each series extending from the costa to the margin in a usually regular line, or a third series present only near the costa; sori indusiate, most or all of the indusia with a sinus, often attached laterally d d. Lamina simple, with 2 basal lobes, or with a single pair of stalked or sessile pinnae, or with 2 or more such pairs and then the basal pinnae entire to shallowly lobed or pinnatifid, these and the usually entire pinnae distally mostly with obtuse lobes 2. T. incisa d. Lamina with 2 or more pairs of stalked or sessile pinnae, the basal pinnae deeply pinnatifid or more complex, with 2 or more large, basiscopic lobes, these pinnae and distal ones mostly regularly lobed with acute to acuminate, sometimes subacute or obtuse lobes 3. T. lizarzaburui c. Sori of the lateral pinnae or large pinna-segments in 3-6, usually 4 or 5, series between the costules, each series extending from the costa toward the margin in a regular or sometimes irregular line; sori indusiate or not e e. Basal pinnae stalked or sessile, the rachis immediately above them not alate; sori indusiate, or most exindusiate, or a few small indusia present; proliferous buds sometimes present at the base of the pinnae 4. T. antioquoiana e. Basal pinnae joined to the pair above by the alate rachis; sori exindusiate; proliferous buds absent 5. T. draconoptera b. Lamina entire, long-decurrent at the base; sori exindusiate 6. T. plantaginea TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 23 1 . Tectaria brauniana (Karsten) C. Chr., Index fil. suppl. 3: 177. 1934. Aspidium braunianum Karsten, Fl. columb. 1: 63, /. 31. 1859. TYPE: Colombia, "Villavizencio" (Vi- llavicencio). Karsten (not located). Stem rather long-creeping. Leaves ca. 25-70 cm long. Lamina mostly 2-pinnatifid, to 3-pinnatifid at the base, gradually reduced at the apex, pu- bescent on the axes on both sides but especially abaxially; basal pinna-segments pinnatifid, more developed on the basiscopic side; lateral pinna- segments obtusely to sometimes acutely lobed; proliferous buds absent. Veins mostly to usually wholly free. Leaf-tissue glabrous to somewhat pu- bescent. Sori indusiate, in a single series between the costules and margin. Indusium with a broad to narrow sinus, usually glabrous. Primary forests, wet ravines, one label indicates that the plant was an epiphyte, 250-800 m, Huan- uco, Cuzco, and Madre de Dios. Costa Rica to Bolivia. The essentially free venation is distinctive for this species, as is the 2-3 -pinnatifid lamina archi- tecture. Huanuco: Ca. 1 5 km N of Tingo Maria, Woytkowski 1363 (uc). Cuzco: Prov. Paucartambo, Cosnipata Valley, N of Patria, Wachteret al. 196 (F). Madre de Dios: Prov. Tambopata, Dist. Tambopata, Yung 21 (MO, uc). 2. Tectaria incisa Cav., Descr. pi. 249. 1802. TYPE: Puerto Rico, comm. (and probably collected by) Ventenant (holotype, MA, seen by C. Chr. [Dansk Bot. Ark. 9(3): 14. 1937]). Figure 4a-c. Aspidium martinicense Sprengel, Anleit. Kenntn. Gew. 3: 133. 1804. TYPE (cited by Morton, 1966): Dominican Republic, Poiteau (not located). Plu- mier, Traite foug. Amer, t. 145, also cited by Sprengel, is this species. Tectaria martinicensis (Sprengel) Copel., Philipp. J. Sci. 2C:410. 1907. Stem ascending to erect. Leaves ca. 20 cm to 2 m long. Lamina simple, with 2 basal lobes, or with 2 basal entire pinnae and an entire terminal seg- ment, or usually with 2 or more stalked or sessile pinnae, then the basal pinnae entire or usually with 1 large, basiscopic lobe on 1 or both of the pinnae, or with 2 lobes on 1 or both of the pinnae, or less often with 2 or more large, basal lobes and smaller lobes beyond on both sides; lateral pinnae usually entire, sometimes lobed, the stalked terminal seg- ment entire to variously lobed; proliferous buds sometimes present at or near the base of the pin- nae, or only 1 bud present; the stalked terminal segment and some distal pinnae may be long-de- current on the rachis. Veins fully anastomosing, except for the included free veinlets. Leaf-tissue glabrous to very short-pubescent to pilose on both surfaces. Sori indusiate, the indusia mostly or all with a sinus, often attached laterally, in 2 series between costules from the costa to the margin, or a third series present only near the costa, in usually regular lines. Indusium glabrous or sometimes shortly glandular-pubescent on the margin and surface. In primary forests, in cloud forests, on hillsides, in ravines, often among rocks, along roadsides, and sometimes on wet, shady cliffs, 100-2100 m, Cajamarca south to Madre de Dios. Tropical America. In Peru Tectaria incisa is a highly variable spe- cies in the architecture of the basal pinnae of the lamina and of the stalked terminal segment, in the presence or absence of proliferous buds at or near the base of the pinnae, and in the pubescence or lack of it on the leaf tissue. Population studies, including information on developmental stages, as well as on cytology and hybridization are needed to determine the basis for the extreme variation and the evolutionary validity of some of the vari- ations. The characters employed to define taxa in this complex all vary from one extreme to another, and sometimes a single collection will consist of more than one named taxon. For these reasons the species Tectaria incisa is treated broadly. The principal variations with their names are distin- guished below and by our annotations. This treat- ment serves to organize the variation of the species so that it may be studied more readily, and at the same time avoids the formal recognition of groups that may not be evolutionary units. Although these groups are not recognized as taxa in Peru, in regions to the north the circumstances may be different. a. Typical Tectaria incisa has both basal pinnae, or at least one of them, with a large, basiscopic lobe, the two sides beyond are entire or nearly so, the stalked terminal segment is usually lobed, of- ten with two, sometimes one, large basal lobe(s), the pinnae have the leaf-tissue glabrous or essen- tially so above and beneath, small plants may have the lamina with two basal lobes, or with two basal 24 FIELDIANA: BOTANY pinnae and a large terminal segment. Specimens have been annotated as Tectaria incisa. This variation (var. incisa) occurs throughout the range of the species in Peru, except for Caja- marca, where it is not yet known. Amazonas: Prov. Bagua, Rio Maranon, above Cas- cadas de Mayasi, Wurdack 1861 (us). San Martin: Cha- zuta, Rio Huallaga, Klug 4038 (F, GH, MO). Loreto: Prov. Maynas, 10 km S of Iquitos, Tryon & Tryon 5206 (F, GH, us). Above Pongo de Manseriche, Mexia 6343 (F, GH, MO, uc, us). Huanuco: Prov. Huanuco, near conflu- ence of Rio Huallaga and Rio Cayumba, Mexia 8304 (BM, F, GH, MO, uc, us). Cerca Tingo Maria, Aguilar 307 (USM). Pasco: Yapas, Pichis Trail (as Junin), Killip & Smith 25584 (us). Junin: Prov. Tarma, 4 km N of La Merced, Tryon & Tryon 5443 (GH, us). Ucayali: Prov. Coronel Portillo, Bosque Nacional von Humboldt, Vas- quez 3882 (F, MO). Ayacucho: Near Kimpitiriki, Killip & Smith 22975 (GH, us). Cuzco: Prov. La Convention, Rio Apurimac, above Boca de Tigre rapids, Davis et al. 1315, 1 317 (F, GH). Madre de Dies: Parque Nacional del Manu, Cocha Cashu Station, Foster & Terborgh 6613 (GH). b. Plants with leaves that are similar to typical T. incisa may have the leaf tissue pubescent above and beneath. These have sometimes been named Tectaria incisa var. pilosa (Fee) Morton. The in- dument varies from pilose to very short-pubescent and from quite dense to very sparse. Specimens have been annotated as Tectaria incisa-pilose. They are scattered through the range of the species and do not seem to represent a taxon. This variation ranges in Peru from Cajamarca south to Madre de Dios. A few collections are the following: Cajamarca: Woytkowski 6905 (us). Huanuco: Allard 21868 (us), 27577 (us), Ferreyra 10238 (GH). Loreto: Wurdack2146 (GH), Mexia 6354 (GH, MO, uc, us). Pasco: Killip & Smith 26516 (F, us), Leon 307 (USM). Madre de Dios: Chavez 1846 (MO). c. Proliferous buds are present at or near the base of the pinnae on some or all leaves of a plant. This variation has been named Tectaria incisa f. vivipara (Jenm.) Morton, and as a species, Tectaria vivipara Jermy & Walker. There may be tight, scaly buds or the buds may have developed into small plantlets; sometimes there is a single bud on the lamina. This variation is sporadic in its distribu- tion and does not seem to be a taxon. Collections have been annotated as Tectaria incisa-vivipa- rous. In Peru it is known from Loreto, Huanuco, Pas- co, and Madre de Dios. The following document the occurrence of this variation in Peru: Loreto: J. Schunke V. 2688 (F, GH, us), Mexia 6250 (F, GH, MO, uc, us). Huanuco: Gentry et al. 41435 (MO). Pasco: (as Junin), Killip & Smith 26690 (F, us). Madre de Dios: Foster et al. 3401 (F), Nunez 5728 (MO). d. The basal pinnae, and sometimes those above, are definitely lobed beyond the one or more large, basal basiscopic lobes, and the stalked terminal segment is lobed. This variation has been named Tectaria incisa ssp. transiens Morton, and has been treated as a species, Tectaria transiens (Morton) A. R. Sm. The leaf-tissue may be pilose above and beneath or glabrous. It does not seem to merit recognition as a taxon in Peru. Specimens have been annotated as Tectaria incisa "transiens." It occurs from Cajamarca south to Madre de Dios. The following represent this variation: Cajamarca: Soukup 3813 (us). Amazonas: Barbour 2496 (MO). Loreto: /. Schunke 325 (F, GH, us). Madre de Dios: Foster & Wachter 7420 (F, MO). e. All of the pinnae and the stalked terminal segment may be entire or nearly so, and proliferous buds usually occur near or at the base of the pin- nae, or a single bud may be present. This is Tec- taria andina (Sodiro) C. Chr. The leaf-tissue is glabrous on both surfaces. It occurs in Ecuador and Peru and may represent a taxon. However, it seems to intergrade with typical Tectaria incisa and so is not treated formally here. Specimens have been annotated as Tectaria incisa "andina." It is known from Amazonas, Loreto, and Hua- nuco in Peru. A few of the collections are: Amazonas: Soukup 6611 (GH), Wurdack 1823 (GH, us). Loreto: Mexia 6131a, in part: (GH, us). Huanuco: Skog et al. 5141 (us), Gentry et al. 36148 (MO). 3. Tectaria lizarzaburui (Sodiro) C. Chr., Index fil. suppl. 3: 181. 1934. Nephrodium lizarzaburui Sodiro, Recens. crypt, vase. Quit. 55. 1883. TYPE: Ecuador, Atacazo, Ha- cienda Bolona, Sodiro (holotype, not located). Ec- uador, Sodiro (K.!), photo GH may be authentic, it is not wholly typical. Stem ascending to erect. Leaves to 2.5 m long. Lamina mostly 1 -pinnate, more complex at the base; terminal stalked segment acutely lobed on each side, with two large, acuminate basal lobes, somewhat decurrent on the rachis (cuneate at the base); basal pinnae usually deeply pinnatifid on the basiscopic side with 2 to several large, acu- TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 25 minate lobes, sometimes to 2-pinnate-pinnatifid, the acroscopic side mostly acutely to subacutely lobed; lateral pinnae mostly regularly, acuminate- ly to obtusely lobed; proliferous buds absent. Veins fully anastomosing, except for included free vein- lets. Leaf-tissue glabrous to sometimes short-pu- bescent abaxially. Sori indusiate, in 2 series be- tween the costules from the costa to the margin, in usually regular lines, or a third series present only near the costa. Indusium with a sinus, often attached laterally, glabrous to slightly pubescent on the surface, the edge glabrous to ciliate. In primary forests, in cloud forests, and in sec- ondary growth, 1000-2000 m, Amazonas south to Ayacucho. Venezuela and Colombia, south to Peru. This species is close to Tectaria incisa, especially to variation "d" with the basal pinnae lobed be- yond the large, basal lobe. That variant of T. incisa differs, however, in having obtuse lobes and the leaf is usually 1 m or less long. Tectaria lizarza- burui has most lobes acuminate to subacute and a large leaf to 2.5 m long. Cajamarca: Prov. Santa Cruz, 3.5 km ENE of Mon- teseco, Santisteban & Guevara 130 (F). Amazonas: Prov. Bagua, 12 km E of La Peca, Harbour 2692 (F, MO, uc). Prov. Bongara, 5 km N of N end of Lake Pomacocha, on road to Rioja, Hutchison & Wright 6787 (GH, uc, us). Pasco: Prov. Oxapampa, Rio El Tungui, Smith et al. 1709 (MO). Junin: Above San Ramon, C. SchunkeA217 (us), Killip & Smith 24639 (F, us). Chanchamayo Valley, C. Schunke 36 (F, us), 576, 950 (F). Ayacucho: Prov. La Mar, eastern Massif of the Cordillera Central, Dudley 11907 (GH). 4. Tectaria antioquoiana (Baker) C. Chr., Index fil. suppl. 3: 177. 1934. Nephrodium sodiroi Baker, J. Bot. 15: 16. 1877. TYPE: Andes of Ecuador, Sodiro (holotype, Ecuador, fl. Pilaton, Sodiro, K!; photo, GH), nom. rejec. (see discussion). Nephrodium antioquoianum Baker, J. Bot. 19: 205. 1881. TYPE: Colombia, Antioquia, Kalbreyer 1806 (holotype, K!; photo, GH). The epithet al- tered to antioquianum by Baker, Ann. Bot. (Lon- don) 5: 329. 1891. Polypodium haynaldii Sodiro, Recens. crypt, vase. Quit. 61. 1883. TYPE: Ecuador, San Miguel de los Colorados, Sodiro (holotype, not located; iso- type, K!; photo, GH). Tectaria sodiroi (Baker) Maxon, Proc. Biol. Soc. Wash. 43: 88. 1930, nom. rejec. Tectaria haynaldii (Sodiro) C. Chr., Index fil. suppl. 3: 180. 1934. Stem ascending to erect. Leaves ca. 50 cm to 1 .5 m long. Lamina 1 -pinnate, with 1 to 2 pairs of stalked or sessile pinnae; stalked terminal segment and adnate large pinna-segments, when present, long-decurrent on the rachis, the stalked terminal segment usually lobed, sometimes entire; basal pinnae entire to usually with 1, rarely 2, large, basal, basiscopic lobes, not connected to the pair above by an alate rachis; proliferous buds rarely present, usually absent. Veins fully anastomosing, except for included free veinlets. Leaf-tissue gla- brous above and beneath. Sori usually indusiate, with a sinus, rarely a few peltate, in 3-6 series between costules from the costa toward the mar- gin, or well beyond the costa, in regular or often irregular lines. Indusium glabrous, rarely some very small indusia present. In dense forests, primary forests, somewhat open or shrubby woods, often on hillsides or in ravines, 100-1500 m, Amazonas south to Cuzco. Colombia south to Peru. This species differs from the next, Tectaria dra- conoptera, in not having the rachis alate to the basal pair of pinnae, and from Tectaria incisa in having three to six, usually four or five, series of sori between adjacent costules. Tectaria sodiroi has been commonly applied to this species, and we have annotated sheets with this name. However, Dr. Robbin Moran has pointed out to us that the holotype (seen by him at K) consists of a part of a stem with lomariopsid vascular pattern and an attached sterile leaf of Bolbitis nicotianifolia (Sw.) Alston. There is also a fertile pinna of Tectaria sodiroi. Since the fertile pinna provides an inadequate holotype, the name Tectaria sodiroi is rejected. Amazonas: Prov. Bagua, below Montenegro, 1 8 km E of Olmos, Hutchison & Wright 3649 (GH, MO, uc, us). Prov. Bagua, Montenegro-Chiriaco, Sagdstegui 5922 (GH). San Martin: Zepelacio, near Moyobamba, Klug 3666 (GH, MO, us). Loreto: Santa Rosa, below Yurima- guas, Killip & Smith 28985 (us). Prov. Maynas, Yano- mono, Vdsquez & Jaramillo 4100 (MO). Huanuco: Prov. Leoncio Prado, E de Tingo Maria, /. Schunke V. 10169 (F, MO, uc). Prov. Huanuco, Dist. Churubamba, Mexia 8217 (BM, F, GH, MO, uc, us). Tingo Maria (as San Martin), Allard 21564, 21565 (us). Pasco: Prov. Oxapampa, Rio Alto Iscozacin, Foster & d'Achille 10062 (F). Cuzco: Prov. Paucartambo, Atalaya, Foster et al. 3074 (GH). Prov. La Convention, Rio Mapitunuari, Dudley 11428 (GH, MO). 5. Tectaria draconoptera (D. C. Eaton) Copel., Philipp. J. Sci. 2 C: 410. 1907. 26 FIELDIANA: BOTANY Aspidium draconopterum D. C. Eaton, Mem. Amer. Acad. Arts n.s. 8: 211. 1860. TYPE: "Turbo in sinu Uraba, Novae Granadae" (Colombia), A. Schott 19 (not located). Stem ascending to erect. Leaves ca. 80 cm to 1 .5 m long. Lamina deeply pinnatisect, the large pin- na-segments all connected by the broadly to nar- rowly alate rachis; basal pinna-segments entire or with 1 large, basal, basiscopic lobe; lateral pinna- lobes entire; proliferous buds absent. Veins fully anastomosing, except for included free veinlets. Leaf-tissue glabrous above and beneath. Sori ex- indusiate, in 4-7 series between costules from the costa to near the margin, the series in regular or mostly irregular lines. In primary forests and in forested ravines, 350- 700 m, Amazonas south to Cuzco and Madre de Dios. Central America; Colombia, Ecuador, and Peru. Tectaria draconoptera is a distinctive species, with very many small, exindusiate sori, and with the basal pinnae connected to the pair above by the alate rachis. Amazonas: Valley of Rio Maranon, above Cascadas de Mayasi, Wurdack 1862 (us). Loreto: Veradero de Ma- zan, Rio Amazonas to Rio Napo, Croat 19530 (F, MO). Huanuco: Prov. Leoncio Prado, E of Tingo Maria, /. Schunke V. 10171 (F, MO, us). Pasco: Prov. Oxapampa, Paujil, Leon 296 (USM). Junin: 2 km S of Satipo, Solomon 3274 (MO, uc). Ucayali: Vicinity of Aguaytia (as Loreto), Croat 209 19 (MO). Cuzco: Prov. Paucartambo, Hacienda Villa Carmen, Vargas 14682 (GH). Prov. Paucartambo, Cosnipata, Vargas 15779 (GH). Rio Alto, Urubamba, Bites 1760 (us). Madre de Dios: Parque Nacional del Manu, Foster & Terborgh 6549 (F). 6. Tectaria plantaginea (Jacq.) Maxon, Contr. U.S. Natl. Herb. 10: 494. 1908. Polypodium plantagineum Jacq., Collectanea 2: 104, t. 3,f. 1. 1788. TYPE: Martinique, probably Jac- quin (holotype, probably w or BM, not seen). The illustration is definitive. Stem short-creeping. Leaves ca. 20-60 cm long. Lamina entire, long-decurrent at the base; a pro- liferous bud present at the apex of the lamina. Veins fully anastomosing, except for included free veinlets. Leaf-tissue glabrous above and beneath. Sori exindusiate or rarely indusiate, in 2 series between the costules, or in 3 or 4 series from the costa to the margin, the lines regular when in 2 series, often irregular and only near the costa when in 3 or 4 series. In ravines and on stream banks in primary wet forests, 350-750 m, San Martin, and Loreto, south to Huanuco and Madre de Dios. Central America and West Indies, south to Peru and Brazil. This is a very distinctive species with an entire lamina that is long-decurrent at its base and with an apical bud. The species is usually exindusiate, but in the Guianas T. plantaginea var. macrocarpa (Fee) Morton is indusiate. Tectaria plantaginea var. conjluens Morton, with especially the lower sori confluent and the sporangia on an elongate receptacle, seems of sporadic occurrence and not a taxon. San Martin: Prov. Mariscal Caceres, Palo Blanco, above Rio Tocache, Plowman & Schunke 7451 (F). Prov. Mariscal Caceres, Nuevo Progreso, J. Schunke V. 3135 (F, GH, us). Prov. Mariscal Caceres, Santa Rosa de Mi- shollo, J. Schunke V, 6813 (F, us). Loreto: Balsapuerto, Killip & Smith 28535 (F, GH, us). Huanuco: Tingo Maria (as San Martin), Allard 20884, 20887, 20894, 21550 (us). Madre de Dios: Prov. Manu, Casa Erika, Rio alto Madre de Dios, Foster & Baldeon 12887 (F). Comments Two species have been excluded from the Pe- ruvian Tectaria flora. Tectaria heracleifolia (Willd.) Undenv., Bull. Tor- rey Bot. Club 33: 200. 1906. Aspidium heracleifolium Willd., Sp. pi. ed. 4, 5: 217. 1810. LECTOTYPE (designated by Underw., Bull. Torrey Bot. Club 33: 200. 1906): Hispaniola, Plu- mier, Traite foug. Amer. /. 147, 1705. This species occurs in southern Florida, the West Indies, Mexico and Central America, and northern South America. It evidently does not occur in Peru. It has a centrally attached, peltate indusium and in this character differs from other species in America. While a few Peruvian collections have some peltate indusia, some to most of the indusia have a sinus and the specimens are referred to other species. Tectaria trifoliata (L.) Cav., Descr. pi. 249. 1802. Polypodium trifoliatum L., Sp. pi. 1087. 1753. LEC- TOTYPE (designated by Underw., Bull. Torrey TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 27 FIG. 5. Cyclopeltis semicordata: a, habit; b, pinna base, abaxial side. (From Stolze, Ferns & fern allies of Guatemala, 1981.) 28 FIELDIANA: BOTANY Bot. Club 33: 199. 1906): Martinique, Plumier, Traite foug. Amer. /. 148. 1705. This species occurs in the West Indies, and in Surinam, Venezuela, and northern Brazil. The in- dusia have a sinus and the sori are in several series between costules. It differs from Tectaria antio- quoiana, to which materials may key out, in hav- ing the lamina only lobed, or if pinnate then the terminal stalked segment is abrupt at the base rath- er than strongly decurrent. Two names of uncertain application are based on Peruvian collections. While they both probably represent Tectaria antioquoiana, accurate identi- fication depends on a study of authentic materials: Tectaria kunzei (Hieron.) C. Chr., Index fil. suppl. 3: 181. 1934. is based on Aspidium kunzei Hieron., Hedwigia 46: 353, 15 July, 1907., which in turn is based on Aspidium macrophyllum var. decurrens Kunze (not Tectaria decurrens (Presl) Copel., 6 Nov. 1907). Aspidium macrophyllum var. decurrens Kunze, Linnaea 9: 89. 1834 is based on two collections of Poeppig: Diar. 1138 from Pam- payaco, and Diar. 2288 from Prov. Maynas. The latter is evidently the type collection, because Diar. 1138 was segregated as a distinct species by Presl in 1851, leaving Diar. 2288 to represent var. de- currens. Tectaria poeppigii (Presl) C. Chr., Index fil. suppl. 3: 183. 1934, is based on Aspidium poeppigii Presl, Epim. Bot. 62. 1851. Presl cites only Pampayaco, Poeppig, which, from the locality, must be Poeppig 1138. V. Cyclopeltis Cyclopeltis John Sm., Companion, Bot. Mag. 72, III, 2: 36 (first of two pages). 1846. TYPE: Cyclopeltis semicordata (Sw.) John Sm. (Poly- podium semicordatum Sw.). Figure 5a-b. Stem decumbent, rather stout, short-creeping, bearing scales. Leaves ca. 20 cm to 1.5 m long, petiole continuous. Lamina 1 -pinnate, the pinnae entire or nearly so, nearly glabrous to somewhat scaly and pubescent, monomorphic, pinnae artic- ulate, adaxial side of the costa raised. Veins free or slightly anastomosing. Sori roundish, on the veins in 1-3 series on each side of the costa, not paraphysate, covered by fugacious to persistent, peltate indusia. Spores spheroidal, monolete, with prominent folds, more or less spinulose. Cyclopeltis is a tropical genus of four to six spe- cies, with one of them in American and in Peru. 1. Cyclopeltis semicordata (Sw.) John Sm., Com- panion Bot. Mag. vol. 72, III, 2: 36 (first of two pages). 1846. Figure 5a-b. Polypodiwn semicordatum Sw., Prodr. 132. 1788. TYPE: Jamaica, Swartz (not seen); "Jamaica, ex Vahl" B!, Herb. Willd. 19742, photo GH is doubt- less authentic. Stem short-creeping, the apex densely scaly with soft, long, linear, brown scales. Leaves mostly 50 cm to 1 m long, the petiole more or less persistently scaly especially toward the base. Lamina 1 -pin- nate, usually with a conform apical segment, pin- nae entire or rarely slightly lobed, sessile or nearly so, auriculate at the base of the basiscopic side, the large auricle overlapping the rachis, articulate to the rachis and deciduous with age. Veins free or slightly anastomosing. Sori borne in 2 series (rarely 1 or 3) on each side of the costa, indusium peltate. Primary forests, hillside forests, thickets, partly disturbed forests, and river banks, 100-750 m, mostly 400 m or lower, Amazonas to Madre de Dios. Southern Mexico and the Greater Antilles, south to Bolivia and the Amazon Basin of Brazil. This is a very distinctive species, with a 1 -pin- nate lamina, articulate pinnae with a large basal, basiscopic auricle, and peltate indusia. The veins are usually free but sometimes they are casually anastomosing. Amazonas: Prov. Bagua, 3-5 km above mouth of Rio Santiago, Wurdack 2180 (F, GH, us). E of Huampami, Rio Cenepa, Berlin 301 (MO). San Martin: Prov. Maris- cal Caceres, Dist. Campanilla, Mashuyaca, J. Schunke V. 4218 (F, GH, MO, us). Juan Jui, Klug4166 (MO). Loreto: Gamitanacocha, Rio Mazan, C. Schunke 194 (F, GH, uc, us). Mouth of Rio Santiago, above Pongo de Manseriche, Mexia 6112 (F, GH, MO, uc, us). Huanuco: Prov. Pachi- tea, Bosque Nacional de Iparia, J. Schunke V. 1247 (F, GH, us). Prov. Huanuco, Tulumayo, cerca a Tingo Maria, Ferreyra 2174 (GH, USM). Prov. Tingo Maria, Aldave & Fernandez 5603 (HUT). Pasco: Quillasu, Soukup 3328 (F, GH). Junin: Prov. Tarma, La Merced, Cerrate 2828 (F, GH). Ucayali: Prov. Coronel Portillo (as Loreto), Bos- que Nacional de Iparia, J. Schunke V. 2673 (F, GH, us). Prov. Coronel Portillo, Bosque von Humboldt, Young & Salazar 1017 (MO). Ayacucho: Near Kimpitiriki, Rio Apurimac, Killip & Smith 22881 (GH, us). Madre de Dios: Parque Nacional de Manu, Cocha Casha, Foster TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 29 2 cm 2 cm FIG. 6. Rumohra adiantiformis: a, habit; b, portion of rachis with bases of costae, adaxial side; c, pinnule, abaxial side. (From Anderson et al. 35820, Brazil, F.) 30 FIELDIANA: BOTANY & Terborgh 6636, 6665 (F). Prov. Tambopata, Rio Ma- dre de Dios, Alfaro 1851 (MO). VI. Rumohra Rumohra Raddi, Opusc. Sci. 3: 290. 1819. TYPE: Rumohra aspidioides Raddi = Rumohra adiantiformis (Forster) Ching. Figure 6a-c. Stem rather stout, long-creeping, bearing scales. Leaves ca. 10 cm to 1 m long, petiole continuous. Lamina 2-pinnate-pinnatifid to 4-pinnate, gla- brous or somewhat scaly, monomorphic, adaxial side of the rachis with a central ridge and glabrous grooves on each side, pinnae not articulate. Pinna- rachis sulcate on the adaxial side, the groove con- tinuous with that of the rachis. Veins free. Sori roundish, on the veins or at the vein tips, not paraphysate, covered by peltate indusia that, with the slender central stalk, is often fugacious. Spores rather ellipsoidal, monolete, with short to long ridges, or saccate. Rumohra is a small genus consisting of the wide- ly spread R. adiantiformis and a few other species. In America there is the former species and R. ber- teriana (Colla) Duek & Rodriguez of the Juan Fer- nandez Islands. 1 . Rumohra adiantiformis (Forster) Ching, Sinen- sia 5: 70. 1934. Figure 6a-c. Polypodium adiantiforme Forster, Prodr. 82. 1786. TYPE: "Ins. austral." G. Forster (holotype or iso- type, BM, GOET). Stem long-creeping, the leaves spaced, densely covered with brown, broad, thin scales. Leaves to ca. 1 m long, the petiole usually scaly, especially toward the base. Lamina 2-pinnate-pinnatifid to 4-pinnate, glabrous or somewhat scaly, coria- ceous, the basal pinnae the largest. Sori roundish, indusium peltate, often fugacious. On rocks near river, ca. 2400 m, Amazonas. Bermuda, south to southern Chile and the Falk- land Islands; Old World. Rumohra adiantiformis is uncommon in the Andes. It is distinguished from species of other genera by the peltate indusia, stalked pinnae, and the entire to obtusely dentate ultimate segments. If the indusia are absent, the adaxial side of the rachis, with a central ridge and a groove on each side, is distinctive. The adaxial side of the penul- timate segments has the costa prominently raised, rather than flattened or grooved as in Dryopteris. Amazonas: Prov. Chachapoyas, Rio Ventilla, 1-2 km W of Molinopampa, Wurdack 1472 (GH, us). VII. Lastreopsis Lastreopsis Ching, Bull. Fan Mem. Instil. Biol., Bot. 8:157.1938. TYPE: Lastreopsis recedens (Moore) Ching (Lastrea recedens Moore) = Lastreopsis tenera (R. Br.) Tindale. Figure 7a-b. Stem erect to short- or long-creeping, usually moderately stout, bearing scales. Leaves ca. 50 cm to 2 m or rarely 3 m long, petiole continuous. Lamina 2-pinnate to 5-pinnate-pinnatifid, pubes- cent, glandular-pubescent, glandular, and often scaly, monomorphic, adaxial side of the rachis with a ridge on each side of a pubescent groove, pinnae not articulate, the pinna-rachis sulcate adaxially, the groove continuous with that of the rachis. Veins free. Sori roundish, borne on the veins or on the vein tips, not paraphysate, covered by reniform, rarely subpeltate indusia or exindusiate. Spores rather ellipsoidal, monolete, strongly ridged, or saccate. Lastreopsis is a genus of about 35 species, with five in America and two in Peru. It is pantropical and south temperate in its distribution. It is readily separated from Dryopteris, Stigmatopteris, and Cyclodium by the raised costa on the adaxial side. Those other genera, which may have some species that resemble some of Lastreopsis, all have the costa grooved on the adaxial side. Reference TINDALE, M. D. 1 965. A monograph of the genus Lastreopsis Ching. Contr. N.S.W. Natl. Herb., 3: 249-339. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 31 FIG. 7. Lastreopsis effusa ssp. divergens: a, portion of lamina. Lastreopsis killipii: b, portion of rachis and pinna base, adaxial side; c, tertiary segment, abaxial side, (a from R. Williams 1273, Bolivia, GH; b, c from Little 9401, Colombia, F.) FIELDIANA: BOTANY Key to Species of Lastreopsis a. Lamina catadromic, at least above the basal pinnae; sori exindusiate; rachis glabrous or nearly so 1 . L. of f usa a. Lamina anadromic throughout; sori indusiate; rachis prominently scaly 2. L. killipii 1 . Lastreopsis effusa (Sw.) Tindale, Viet. Nat. 73: 184. 1957. Figure 7a. Polypodium effusum Sw., Prodr. 134. 1788. LEC- TOTYPE (designated by Tindale, 1957): Jamai- ca, Swartz(ho\oiype,Bl,Herb. Willd. 1 97 24; pho- to GH; isotypes, c, UPS). Stem decumbent, short-creeping, to nearly erect, bearing brown, lanceolate scales. Leaves ca. 1-2.5 m long; the petiole glabrous, or slightly scaly, es- pecially toward the base. Lamina 3 -pinnate to 4-pinnate-pinnatifid, catadromic, at least above the basal pinnae, the apex gradually reduced, usu- ally somewhat pubescent and/or glandular, es- pecially on the adaxial side of the axes; usually bearing a proliferous bud toward the apex of the rachis; pinnae stalked, the basal the largest. Veins free. Sori borne on the veins, exindusiate. Dense forests, disturbed forests, and on moun- tain slopes, 135-1 800 m, Cajamarca to Cuzco and Madre de Dios. Tropical America. Of the four more or less sympatric subspecies recognized by Tindale (1957), only ssp. divergens (Willd.) Tindale is in Peru. Lastreopsis effusa is one the few species of the genus that has the lamina usually with a scaly bud toward the apex of the rachis. Dryopteris killipii Maxon, Amer. Fern J. 18: 4. 1928. TYPE: Panama, Chiriqui, W of El Boquete, Killip 5360 (holotype, us; isotype, GH!). Stem short-creeping, decumbent, bearing brown, narrow scales. Leaves ca. 2-5 m long; the petiole rather persistently scaly, with mostly brownish, narrow scales. Lamina 4-pinnate to nearly 5-pin- nate, anadromic, the apex gradually reduced, var- iously scaly, pubescent, and/or glandular, especial- ly on the axes; lacking a proliferous bud; pinnae stalked, the basal the largest. Veins free. Sori borne on the veins, covered by a reniform, persistent indusium. Dense forests and cloud forests, 2080-2700 m, Cuzco. Costa Rica, Panama, Colombia, and Peru. The leaves are up to 3.5 or even 5 m long. Cuzco: Prov. La Convencion, altura de Pintobamba, Vargas 3552 (us). Prov. La Convencion, Cordillera Vil- cabamba, Dudley 11243 (GH, us). Comments Except for the widespread Lastreopsis effusa, other species of South America are not commonly collected, but two of them may be found in Peru. Cajamarca: Prov. Hualgayoc, Soukup 3812 (F, us). Prov. Santa Cruz, Sagdstegui et al. 12391 (F, GH, HUT). Loreto: Prov. Maynas, Rio Nanay, across from Bellavis- ta, McDaniel & Rimachi 18794 (GH, MO). Santa Rosa, below Yurimaguas, Killip & Smith 28733, 28853 (us). Huanuco: Pampayacu, Rio Chinchao, Macbride 5036 (F, us). Pasco: Prov. Oxapampa, Valle del Palcazu, Leon 679 (F). Junin: Above San Ramon, C. Schunke A226 (us). Chanchamayo Valley, C. Schunke 50, 90 (F, us). Prov. Satipo, Leon 186 (MO, USM). Cuzco: Prov. La Con- vencion, Rio Apurimac, Davis et al. 1310 (F). Madre de Dios: Prov. Manu, Cocha Cashu uplands, Nunez 5815 (MO). 2. Lastreopsis killipii (Maxon) Tindale, Viet. Nat. 73: 185. 1957. Figure 7b-c. Lastreopsis amplissima (Presl) Tindale, Viet. Nat. 73: 185. 1957. Polystichum amplissimum Presl, Epim. Bot. 58. 1851. SYNTYPES: Brazil, Sellow; Brazil, Serra d'Es- trella, Beyrich (neither located); ISOSYNTYPE: Sellow (B). This species has a wholly anadromic lamina with indusiate sori and is closely related to L. killipii. It has the ultimate segments mostly sharply point- ed to mucronate apically, while L. killipii has them mostly obtuse to subacute. This species ranges from Venezuela south to southeastern Brazil and Par- TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 33 FIG. 8. Dryopteris paleacea: a, habit; b, pinna segment, abaxial side; Dryopteris denticulata: c, lamina, (a, b from Killip & Smith 18824, Colombia, F, c adapted from Stolze. Ferns & fern allies of Guatemala, 1981.) 34 FIELDIANA: BOTANY aguay; it is also in Bolivia and may be expected to grow in southern Peru. Lastreopsis exculta (Mett.) Tindale, Viet. Nat. 73: 185. 1957. Aspidium excultum Mett., Uber einige Farngatt. IV. Phegopt. Aspid. 69, t.!7,f. 9. 1859. (Abh. SenckenbergNaturf. Ges. 2: 353. 1858. TYPE: Venezuela, Caracas, Moritz 433 (holotype, B?; isotype, GH!). This species has a catadromic lamina, indusia, and a short-pubescent costa adaxially. It grows in Mexico south to northern South America and Ec- uador and probably also grows in northern Peru. VIII. Dryopteris Dryopteris Adans., Fam. plantes 2: 20. 551. 1763, nom. conserv. TYPE: Dryopteris filix- mas (L.) Schott (Polypodium filix- mas L.). Figure 8. Arachniodes Blume, Enum. pi. Javae 24 1 . 1 828. TYPE: Arachniodes aspidioides Blume = IDryopteris ar- istata (Forster) Kuntze. Lastrea subsection Ipolystichopsis John Sm., Hist. fil. 217. 1875. LECTOTYPE (designated by C. Chr., Index fil. xxi. 1906): Lastrea pubescens(Sv/.) Presl (Polypodium pubescens (L.) = Dryopteris pubes- cens (L.) Kuntze. Dryopteris subgenus Polystichopsis (John Sm.) Ch. Chr., Danske Vidensk. Selsk. Skrift. Nat. Math. VII, 6(1): 101. 1920. Polystichopsis (John Sm.) Holtt., Flora Malaya 2 (Ferns Malaya): 484. 1955. Byrsopteris Morton, Amer. Fern J. 50: 149. 1960. TYPE: Byrsopteris aristata (Forster) Morton (Polypodium aristatum Forster) = Dryopteris ar- istata (Forster) Kuntze. Stem decumbent, small or stout, or erect to ca. 30 cm tall, rarely long-creeping, bearing scales. Leaves ca. 1 0 cm to 2 m long, petiole continuous. Lamina 1 -pinnate, usually 2-pinnate-pinnatifid, to rarely 6-pinnate, glabrous, glandular, pubescent, or scaly, monomorphic or slightly dimorphic, ad- axial side of the costa sulcate to nearly flat. Veins free. Sori roundish, borne on the veins or on the vein tips, not paraphysate, covered by usually ren- iform, rarely subpeltate, indusia, or exindusiate. Spores rather ellipsoidal, monolete, prominently ridged, saccate, or cristate, often spinulose. Dryopteris is a large genus of about 1 50 species. It is worldwide in its distribution. The center of species diversity is in China and adjacent regions. There are about 25 species in America, and five in Peru. Dryopteris sensu stricto is closely allied to the sometimes recognized genera Arachniodes and Polystichopsis. Some species, usually placed in one of the three groups, are divergent in their char- acters toward another group. The whole assem- blage is closely allied and relations of a number of species are unclear. For these reasons all species are treated here in the single genus Dryopteris. Two subgenera may be recognized: Dryopteris subgenus Dryopteris has the stem with internal glands, the lamina often catadromic, and the basal pinnae not, or not much, enlarged. Peruvian spe- cies are Dryopteris patula, D. paleacea, and D. saffordii. Dryopteris subgenus Polystichopsis (in- cluding Arachniodes) has the stem lacking internal glands, the lamina anadromic, and the basal pin- nae enlarged. Peruvian species are Dryopteris den- ticulata and D. ochropteroides. Reference TRYON, R. M., AND A. F. TRYON. 1982. Dryop- teris, pp. 496-509, in Ferns and allied plants, Springer- Verlag, New York. Key to Species of Dryopteris a. Rachis definitely scaly with long, narrow, dark brown or darker scales, these grading into fibrils; pinnae sessile or nearly so, deeply pinnatifid to usually pinnatisect; secondary segments approximate, nearly entire 1 • D- paleacea a. Rachis with a few scattered scales, or none, long-pubescent, minutely glandular, or glabrous . . . . b b. Lamina minutely glandular, especially on the rachis, at the base of the pinnae, on the pinna- rachis, costa, and usually elsewhere c c. Pinnae stalked, or rarely short-stalked, the basal ones not or hardly reduced, usually the largest on the lamina; basal secondary segments on the central pinnae narrowed at the base, or stalked, TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 35 usually connected to the next pair of segments by a narrow wing of the pinna-rachis 2. D. patula c. Pinnae sessile or nearly so, the basal ones reduced, usually much reduced, shorter than the central pinnae; basal secondary segments on the central pinnae usually joined to the next pair of segments by their decurrent base 3. D. saffordii b. Lamina eglandular, glabrous, or with a few fibrils, or long-pubescent, especially on the axes d d. Rachis and other axes glabrous or with a few fibrils 4. D. denticulata d. Rachis and other axes long-pubescent, especially on the abaxial side, or partly so 5. D. ochropteroides 1. Dryopteris paleacea (Sw.) Hand.-Mazz., Ver- handl. Zool.-Bot. Gesells. Wein 58: (100). 1908. Figure 8a-b. Aspidium paleaceum Sw., Syn. fil. 52. 1806. TYPE: Peru, Lagasca misit (holotype, s, Herb. Swartz; photos, GH, us). Aspidium parallelogramma Kunze, Linnaea 13: 146. 1839. SYNTYPES: Mexico, Hegewisch, Mexico, Karwinsky (not located). Dryopteris parallelogramma (Kunze) Alston, Amer. Fern J. 47: 92. 1957. Stem erect, stout. Leaves borne in a crown, ca. 30 cm to 1.5m long. Lamina 1 -pinnate-pinnatifid to usually 1 -pinnate-pinnatisect, more or less scaly, especially on the rachis, which is rather densely scaly, with mostly long, narrow, reddish brown to dark brown or partly blackish scales abaxially and often fibrillose adaxially. Pinnae sessile or nearly so, the basal not much shorter than the central ones; the secondary segments approximate, nearly entire to somewhat dentate. In dense forests or thickets, disturbed forests, in moist ravines, on wet, shaded road banks, and at the base of rocks, 2400-4000 m, Cajamarca to Puno. Mexico and Greater Antilles, south to Argentina and southern Brazil; primarily montane. The correct taxonomy of this species is not clear. There may be one, essentially pantropical, species, or there may be two, one American and the other Old World. The American element is here treated as a separate species, pending a definitive reso- lution of this problem. The correct nomenclature depends upon the in- terpretation of the publication of Aspidium pale- aceum by Don (Prodr. fl. nepal. 4. 1825). If this is a new species (Alston, Amer. Fern J. 47: 91-92. 1957; A. R. Smith and Fraser-Jenkins, Taxon 31: 326-329. 1982) then the correct name for the col- lective species is Dryopteris wallichiana (Sprengel) Hylander. This is also the correct name for the Old World element, as a species, while the Amer- ican element is Dryopteris parallelogramma (Kunze) Alston. If Don's name is a later use of Aspidium pale- aceum Sw. (the conclusion adopted here; also by Fraser-Jenkins, Taxon 29: 610-611. 1980) then the correct name for the collective species is Dryopteris paleacea (Sw.) Hand.-Mazz. This is also the correct name for the American element, as a species, while the Old World element is Dryopteris wallichiana (Sprengel) Hylander. Cajamarca: Prov. San Miguel, Vista Alegre, Llatas Q. 1574 (F). Prov. Celendin, entre Celendin y Cajamarca, Lopez et al. 4462 (GH, HUT). Amazonas: Prov. Chacha- poyas, Cerros Calla Calla, Hutchison & Wright 5753 (F, GH, MO, uc, us). Prov. Chachapoyas, Puma-urcu, SE of Chachapoyas, Wurdack 559 (F, GH, uc, us). La Libertad: Prov. Pataz, above Pampa Rosas, Young 3 105 (F). Prov. Santiago de Chuco, Dist. Cachicadan, Sounders 884 (F, GH). Between Huamachuco and Cajabamba, Correll & Smith P925 (GH). San Martin: Dist. Huallaga, valley of Rio Apisoncho, Hamilton & Holligan 925 (uc). Prov. Mariscal Caceres, Rio Abiseo National Park, Young & Leon 4982 (F). Huanuco: Mito, Macbride & Featherstone 1512 (F, us). Prov. Huanuco, 32 km from Huanuco on Huanuco-La Union Road, Smith et al. 2187 (MO). Junin: Carpapata, above Huacapistana, Killip & Smith 24465 (F, us). Huancavelica: Entre Huschocolpa y Surcubamba, Tovar 4237 (GH). Ayacucho: Prov. La Mar, Dudley 9051 (GH). Prov. La Mar, between El Tambo and Ayna, Plow- man & Davis 4684 (GH). Apurimac: Dist. Abancay, Sounders 761 (GH). Cuzco: Machu Picchu, Peyton & Pey- ton 759 (GH, MO). Prov. La Convention, Dist. Vilcabam- ba, Davis et al. 7279 (F, GH). Puno: Prov. Carabaya, entre Ayapata y Kahualluyac, Vargas 10735 (GH). 2. Dryopteris patula (Sw.) Underw., Our native ferns, ed. 4, 117. 1893. Aspidium patulum Sw., Kongl. Vetensk. Acad. Handl. 1817: 64. TYPE: Brazil, Minas Gerais, Freyreiss (holotype, s, seen by C. Chr. Monogr. genus Dryopteris 1: 19. 1913). 36 FIELDIANA: BOTANY Stem erect or nearly so, rather stout. Leaves borne in a cluster, ca. 25 cm to 1 m long. Lamina 2-pinnate to 3-pinnate-pinnatifid, minutely glan- dular, especially on the rachis, at the base of the pinnae, on the pinna-rachis, and often elsewhere, the rachis sometimes with scattered light brown scales. Pinnae stalked, rarely short-stalked to near- ly sessile, the basal not or hardly reduced, usually the largest, secondary segments usually spaced, usually shallowly to sometimes deeply lobed or more complex. In woods, among shaded rocks, and on rocky slopes, 2000-2700 m, Huanuco, Junin, and Cuz- co. Mexico and the Greater Antilles, south to Ar- gentina and southern Brazil; primarily montane. This species has a more complex lamina than Dryopteris saffordii. The basal segments of the cen- tral pinnae are narrowed at the base, or stalked and connected to the next pair of segments by a narrow wing along the pinna-rachis. Huanuco: Chinchao to Puente Durand, Coronado 82 (us). Mima, Bryan 419 (F). Yanano, Macbride 3821 (F, us). Junin: Carpapata, Kunkel 603 (GH). Cuzco: Machu Picchu, Leon 453 (F, GH, HUT), Coronado 95 (uc, us). Prov. Paucartambo, Pillawata, Vargas 16688 (GH). 3. Dryopteris saffordii C. Chr., Amer. Fern J. 1: 94. 1911. TYPE: Peru, mountains back of Lima, Arroya Railroad, W. E. Safford 994 (holotype, us; photo, GH; isotype, us). Stem usually small, decumbent to erect. Leaves borne in a cluster, ca. 15-60 cm long. Lamina 1 -pinnate-pinnatifid to rarely 2-pinnate-pinnati- fid, minutely glandular, especially on the rachis, at the base of the pinnae, on the pinna-rachis, the costa, and often elsewhere, the rachis sometimes with scattered light brown scales. Pinnae sessile or nearly so, the basal ones much reduced, secondary segments approximate or spaced, nearly entire to lobed or rarely pinnatifid. Brushy, rocky slopes, open rocky places, at the base of rocks, 400-4000 m (400-600 m on Loma Lachay, otherwise at 2200-4000 m), Cajamarca to Puno. Peru and Bolivia. Dryopteris saffordii is very close to D. patula and might better be treated as a subspecies or va- riety of it. Soukup 2961, Huancayo, and Fiebrig 3133, in part, Bolivia (GH) have large leaves with the lamina 2-pinnate-pinnatifid and nearly sessile pinnae, the basal definitely reduced. These collec- tions are rather intermediate between the two species. Apparently intermediate specimens, how- ever, are few and its seems best to maintain two species at the present time. Cajamarca: Prov. Contumaza, Guzmango, Sagdstegui 2938 (F, GH). 29 km from Cajamarca on road to Chilete, Correll & Smith P849 (GH), P852 (GH, us). Prov. Caja- marca, cumbre El Galivan, Ferreyra 3276 (USM). La Li- bertad: Prov. Otuzco, Huaranchal, Lopez et al. 2663 (GH). Between Huamachuco and Cajabamba, Correll & Smith P922 (GH). Prov. Sanchez, Carrion, Sagdstegui 9445 (HUT). Ancash: Prov. Bolognesi, E of Huasta, Cer- rate 2460 (GH, USM). Prov. Bolognesi, arriba de Chiqui- an, Cerrate 1550 (GH). Lima: Prov. Chancay, Loma de Lachay (as Luchay), Tryon & Tryon 5417 (F, GH, us), Coronado 22 (GH, uc, us). Prov. Huarochiri, Dist. Surco, Saunders 657 (F, GH, us). Junin: Huancayo, Soukup 2961 (F, GH, MO, us). Cuzco: Prov. Urubamba, Chupani, Var- gas 11125 (GH). Puno: Near Puno, Soukup 92 (F). 4. Dryopteris denticulata (Sw.) Kuntze, Revis. gen. pi. 2: 812. 1891. Figure 8c. Polypodium denticulatum Sw., Prodr. 134. 1788. TYPE: Jamaica, Swartz (holotype, s, seen by C. Chr., Monogr. genus Dryopteris 2: 113. 1920). Arachniodes denticulata (Sw.) Ching, Acta Bot. Sinica 10: 260. 1962. Stem decumbent to erect, small to rather stout. Leaves borne in a cluster, ca. 25 cm to 1 m long. Lamina 3 -pinnate-pinnatifid to 4-pinnate-pinnati- fid, or 5 -pinnate, glabrous or glabrate. Pinnae stalked, the basal ones enlarged, the longest, sec- ondary segments 1 -pinnate or more complex. Moist forests, and roadside banks, 2000-3000 m, Cajamarca, south to Puno. Southern Mexico and the Greater Antilles, south to northern South America and Bolivia; south- eastern Brazil. The lamina is very strongly and wholly anad- romic, and the ultimate segments are mostly sharply dentate, especially the apical ones. Cajamarca: Prov. Cutervo, Llipa, Mostacero et al. 1749 (F, HUT). Amazonas: Prov. Chachapoyas, 5 km below Chachapoyas, Wurdack 778 (F, GH), 779 (F, USM). San Martin: Leon 2144, 2162 (F). Prov. Bongara, El Ingenio- Pomacochas, Sagdstegui 5968 (GH). Huanuco: Cushi, Bryan 685 (F). Pasco: Prov. Oxapampa, Rio San Alberto valley, E of Oxapampa, Smith & Pretel 7598 (F). Puno: Sandia, Vargas 11859 (on). TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 37 5. Dryopteris ochropteroides (Baker) C. Chr., In- dex fil. 280. 1905. Nephrodium ochropteroides Baker, Ann. Bot. (Lon- don) 5: 325. 1891. TYPE: Jamaica, Fox's Gap, Hart (holotype, K; isotype, u). Polystichopsis ochropteroides (Baker) Morton, Amer. Fern J. 50: 155. 1960. Arachniodes ochropteroides (Baker) Lell., Amer. Fern J. 77: 101. 1988. Stem rather slender and long-creeping in small plants, becoming rather stout and decumbent. Leaves clustered to spaced, ca. 40 cm to 1 m long. Lamina 3-pinnate to 4-pinnate-pinnatifid, with the axes long-pubescent, especially on the abaxial side, usually densely so but sometimes the pubescence scattered. Pinnae stalked, the basal ones enlarged and the longest, secondary segments 1 -pinnate or more complex. Dense cloud forests, ca. 1290 m, Huanuco; a single collection from Peru. Jamaica, Surinam, Venezuela, and Peru; range uncertain. The lamina is definitely anadromic and the ul- timate segment are subacute to acute. This species is separated from the similar Dryopteris macrostegia (Hooker) O. Kuntze, of Venezuela, Colombia, and northern Brazil, by the axes of the lamina that are long-pubescent. The latter species has the axes of the lamina glabrous. The pubescence of Dryopteris ochropteroides may be scattered and sparse. Both species are seldom collected and additional material is needed in or- der to understand their relation. long, petiole continuous. Lamina 1 -pinnate, or rarely entire, or 2-pinnate, glabrate to more or less scaly, monomorphic, adaxial side of the costa sul- cate, pinnae not articulate, more or less inequi- lateral at the base. Veins free to regularly anas- tomosing and with or without included free veinlets. Sori roundish, borne on the veins or at the vein tips, in 2 to several (rarely 1) series on each side of the costa, not paraphysate, covered by persistent to fugacious, peltate indusia or rarely exindusiate. Spores ellipsoidal to spheroidal, monolete, ridged, rugose or saccate. Cyrtomium is a genus of about 25 species, most of them in eastern Asia. In America the nine or ten species are concentrated in Mexico and Central America. A single native species, C. dubium, oc- curs in Peru. Although the predominantly free-veined Amer- ican species are sometimes recognized as a distinct genus Phanerophlebia, the two groups are not wholly distinctive. References CHING, R. C. 1936. On the genus Cyrtomium Presl. Bull. Chinese Bot. Soc., 2: 87-106. CHRISTENSEN, C. 1930. The genus Cyrtomium. Amer. Fern J., 20: 41-52. MAXON, W. R. 1912. Notes on the North Amer- ican species of Phanerophlebia. Bull. Torrey Bot. Club, 39: 23-28. UNDERWOOD, L. M. 1899. American ferns, II. The genus Phanerophlebia. Bull. Torrey Bot. Club, 26: 205-216. Huanuco: Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13062 (GH, us). 1. Cyrtomium dubium (Karsten) R. & A. Tryon, Rhodora 83: 134. 1981. Figure 9a-b. IX. Cyrtomium Cyrtomium Presl, Tent, pterid. 86. 1836. TYPE: Cyrtomium falcatum (L. f.) Presl (Polypodium falcatum L. f). Figure 9. Phanerophlebia Presl, Tent, pterid. 84. 1836, cited as a synonym of Cyrtomium by Moore, Index fil. Ixxxii. 1857. TYPE: Phanerophlebia nobilis (Schlect. & Cham.) Presl (Aspidium nobile Schlect. & Cham.) = Cyrtomium nobile (Schlect. & Cham.) Moore. Stem decumbent, short, rather stout, to nearly erect, bearing scales. Leaves ca. 15 cm to 1.5 m Phegopteris dubia Karsten, Fl. columb. 1: 169, t. 84. 1861. TYPE: Colombia, Bogota, Tequendama, Karsten (not located). Phegopteris dictyophylla Kuhn, Linnaea 36: 106. 1869. TYPE: Ecuador, Spruce 5263 (holotype, B?; iso- type, as Spruce 5265, p, Herb. Bonaparte. Not Spruce 5263, P, Herb. Bonaparte which is Polys- tichum Bonapartii Rosenst.; holotype, p, Herb. Bonaparte; isotype, GH!). Polystichum dubium (Karsten) Diels, Nat. Pflanzen- fam. 1(4): 194. 1899. Stem decumbent, very short-creeping, to erect, bearing dark brown, lustrous scales. Leaves ca. 30 cm to 1 m long, the petiole usually sparingly scaly. Lamina usually 1 -pinnate, to rarely partly or fully 38 FIELDIANA: BOTANY FIG. 9. Cyrtomium dubium: a, habit; b, portion of pinna, abaxial side, (a from Peyton & Peyton 1275, GH; b from Killip & Smith 20366. GH.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 39 2-pinnate, the apex gradually reduced, pinnae usu- ally entire or nearly so, herbaceous, usually rather sharply dentate, often with a usually prominent basal auricle on the acroscopic side, veins mostly free to freely anastomosing, with or without in- cluded free veinlets. Sori in usually 3 (2 to 4) series on each side of the costa or costule, exindusiate. Dense hillside forests, scrubby forests, some- times among rocks, or rarely a low epiphyte, 1450- 3400 m, Cajamarca to Cuzco. Costa Rica south to Bolivia. Cyrtomium dubium is a morphologically rather isolated species that is not clearly placed in either Polystichum or Cyrtomium. It has, however, the usually anastomosing veins and multiseriate sori of the latter genus and seems best placed there. It is a variable species in lamina architecture and venation. Spruce 5263 is evidently a mixed collection of 2 species and Spruce 5265 is a mix- ture of 1 -pinnate and 2-pinnate Cyrtomium du- bium. The pinnae vary from usually entire with an obtuse to acuminate apex, to more or less serrately lobed, or the pinnae may be basally mostly 1 -pinnate or fully 1 -pinnate, as in Barbour 2875. The venation varies from fully anastomosing, ex- cept at the margins, to mostly free. Although some of this variation may be related to hybridization, especially with species of Polystichum, spores ap- pear to be normal and the lamina architecture is regular. The 1 -pinnate and 2-pinnate lamina forms are therefore considered to be within the normal range of variation of the species, as in various species of Lindsaea. Cajamarca: Prov. Cutervo, San Andres, Lopez & Sa- gdstegui 5401 (GH, HUT). Amazonas: E of La Peca, Bar- bour 2875 (MO). San Martin: Between Mirador and La Playa, Young & Leon 4927 (F, USM). Huanuco: Mima, Bryan 554 (F, us). Huacachi, near Mima, Macbride4l26 (F, us). Junin: Carpapata, above Huacapistana, Killip & Smith 24464 (BM, F, us). Ayacucho: Between Huanta and Rio Apurimac, Killip & Smith 22362 (us). Cuzco: Uru- bamba, Machu Picchu, Peyton & Peyton 1328 (GH, MO), 1504 (MO). Quillabamba, Santa Teresa, Peyton & Peyton 1275 (GH, MO). Prov. Paucartambo, Pillahuata, Aldave 5021 (HUT). Comments Cyrtomium falcatum (L. f.) Presl, Tent, pterid. 86. 1836. Polypodium falcatum L. f., Suppl. pi. syst. veg. ed. 13, 446. 1782. TYPE: Japan, Nagasaki, Thunberg (holotype, LINN?; isotype, UPS). This is a commonly cultivated species of eastern Asia. It has been collected on the sea-cliffs near Lima and it may be adventive elsewhere in Peru. It diners from Cyrtomium dubium in being in- dusiate, and having very coriaceous pinnae with thickened margins. Lima: Prov. Lima, Magdalena del Mar, cerca a Lima, Julio 14, 1942, Zuniga (GH). X. Didymochlaena Didymochlaena Desv., Ges. Naturf. Freunde Ber- lin Mag. Neuesten Entdeck. Gesammten Na- turk. 5: 303. 1811. TYPE: Didymochlaena sinuosa Desv. = Didymochlaena truncatula (Sw.) John Sm. Figure lOa-c. Stem decumbent or usually erect, rather stout, bearing scales. Leaves ca. 75 cm to 2 m or rarely 3 m long, petiole continuous. Lamina 2-pinnate, nearly glabrous to scaly and pubescent, mono- morphic, pinnules dimidiate. Veins free. Sori elongate, borne on the veins, not paraphysate, cov- ered by elongate indusia on each side of a vein. Spores ellipsoidal to spheroidal, monolete, with short ridges or saccate, spinulose. Didymochlaena is a monotypic genus of pan- tropical distribution. The single species is variable, especially in its lamina architecture. 1. Didymochlaena truncatula (Sw.) John Sm., J. Bot. (Hooker) 4: 196. 1841. Figure lOa-c. Adiantum lunulatum Houtt., Nat. Hist. 2, 14: 209. t. 100, f. 1. 1783, not Burm. 1768. TYPE: uncer- tain, the figure may serve for identification. Aspidium truncatulum Sw., J. Bot. (Schrader) 1800(2): 36. 1802, nom. nov. for Adiantum lunulatum Houtt., and with the same type. Didymochlaena lunulata Desv., Mem. Soc. Linn. Par- is 6: 282. 1827, nom. superfl. tor Aspidium trun- catulum Sw., and with the same type. Stem stout, bearing dense, linear, brown scales. Leaves to 2 m or rarely 3 m long, the petiole mod- erately scaly to usually densely so toward the base. Lamina 2-pinnate, the ultimate segments (pin- nules) short-stalked, dimidiate, entire, obtuse, gla- 40 FIELDIANA: BOTANY FIG. 10. Didymochlaena truncatula: a, stem and portion of lamina; b, pinnules, abaxial side; c, portion of costule with pinnule bases, adaxial side. (From Stolze, Ferns & fern allies of Guatemala. 1981.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 41 brous or slightly scaly, the rachis with long pro- cesses at the base of the pinnae on the adaxial side. Sori covered by an elongate indusium, on each side of a vein, and continuous distally. Wet forests and rain forests, 1 1 0-900 m, Ama- zonas to Cuzco and Madre de Dios. Tropical America; Old World. This is a highly distinctive species, unusual in its pantropical distribution. In Peru, it has been most commonly collected in Loreto. Amazonas: Rio Maranon above Cascadas de Mayasi, Wurdack 2015 (GH, us). Serrania de Bagua, above La Peca, Harbour 2390 (MO). San Martin: Prov. Mariscal Caceres, Tocache Nuevo, /. Schunke V. 6905 (F, MO, us). Prov. Mariscal Caceres, Cerro Santa Cruz, /. Schunke V. 8036 (MO, USM). Prov. Rioja, Pedro Ruiz-Moyabam- ba, D. Smith 4430 (MO). Loreto: Rio Itaya, Soledad, Killip & Smith 29768 (F, GH, us). Prov. Maynas, Yan- amona, McDaniel 16195 (GH). Alto Amazonas, Andoas, Vdsquez 4400 (MO). Huanuco: Tingo Maria, Stork & Morion 9473 (F, uc, us). Pasco: Prov. Oxapampa, vicin- ity of Chequitavo, D. Smith 5263 (MO). Junin: Chan- chamayo Valley, C. Schunke 8 (F, us). Ucayali: Prov. Coronel Portillo, between La Divisoria and El Boqueron, Plowman & Kennedy 5768 (F, GH). Cuzco: Prov. Quis- pichanchi, Punkiri, Vargas 15408, 16108 (GH). Madre de Dios: Prov. Manu, Shintuya, Vargas 17837 (GH), Fos- ter et al. 3143 (GH, us). Plants terrestrial. Stem short-creeping to erect. Lamina 1 -pinnate to 2-pinnate-pinnatifid, taper- ing to a pinnatifid apex, glabrous on both surfaces, with internal, pellucid (sometimes drying black) punctate glands. Pinnae commonly linear to nar- rowly oblong, the apices long-acuminate and ser- rate. Rachis and costae stramineous to brown, scaly, the scales thin, flaccid, usually ending in a papil- lose-glandular tip. Groove of the costae pubescent within, the trichomes less than 0. 1 mm long. Veins free to irregularly anastomosing, curved-ascend- ing, ending behind the margin in a clavate tip. Sori round or slightly oblong near the costae. Indusium absent, but some species with an indusiumlike scale. The genus contains about 25 species, with dis- tribution from southern Mexico to southeastern Brazil and the Antilles. The most distinctive fea- ture of Stigmatopteris is the internal punctate glands. These glands are most easily seen when the leaf is held between the eye and a light, but are also visible under a microscope (oblique light) as tiny, raised bumps. References XI. Stigmatopteris Contributed by Robbin C. Moran Stigmatopteris C. Chr., Bot. Tidsskr., 29: 292. 1909. TYPE: Polypodium flavopunctatum Kaulf. = Stigmatopteris rotundata (Willd.) C. Chr. (Aspidium rotundatum Willd.). Figure 11. Dryopteris subgenus Stigmatopteris (C. Chr.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvi- densk. Afd., ser. 7, 10: 73. 1913. CHRISTENSEN, C. 1 909. On Stigmatopteris, a new genus of ferns with a review of its species. Bot. Tidssk., 29: 291-304. CHRISTENSEN, C. 1913. A monograph of the ge- nus Dryopteris, part 1. The tropical American pinnatifid-bipinnatifid species. Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 7, 10: 53-282. CHRISTENSEN, C. 1 920. A monograph of the ge- nus Dryopteris, part 2. The tropical American bipinnate-decompound species. Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8, 6: 1-132. Key to Species of Stigmatopteris a. Veins free b. Lamina 1-pinnate-pinnatifid FIG. 1 1 . Stigmatopteris heterophlebia: a, habit; b, portion of pinna, abaxial side. Stigmatopteris pellucidopunc- tata: c, portions of pinna segments, adaxial side, (a, b from Moran 3165, Costa Rica, F, c from Brant et al. 1679, Colombia, F.) 42 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 43 a. c. Pinnae cut ca. % to the costa 1 . S. pellucidopunctata c. Pinnae entire or cut less than V4 to the costa 2. S. longicaudata b. Lamina 2-pinnate-pinnatifid 3. S. lechleri Veins anastomosing d d. Rachis and costae densely puberulent 4. S. heterophlebia d. Rachis and costae lacking trichomes 5. S. opaca 1. Stigmatopteris pellucidopunctata (C. Chr.) C. Chr., Dot. Tidsskr. 29: 304. 1909. Figure lie. Polypodium macrophyllum Hooker, Sp. fil. 4: 241. 1 862, nom. illeg., not (Blume) Mett. 1 856. TYPE: Peru, San Martin, Mt. Guayrapurima, August 1856, Spruce 4720 (holotype, K.!; isotypes, BM!, o!, P!, us!; photos, F, GH, MO, NY, all of BM). Dryopteris pellucido-punctata C. Chr., Index fil. 283. 1905, nom. nov. for P. macrophyllum Hooker and with the same type. Lamina 1 -pinnate-pinnatifid. Pinnae 20-40 cm long and 2.5-6 cm broad, 15-20 pairs, cut ca. % to the costa, sessile or the basal ones short-stalked, the stalk 2-7 mm long. Segments 4-10 mm broad, serrate at the apex, the margins entire, serrate or lobed. Rachis and costae stramineous or brown, scaly, the scales ovate, lanceolate or filiform. Veins free, 7-14 per segment, unbranched or 1 -forked with a short acroscopic branch bearing the sorus. Sori round. Wet forests, 200-600 m, Amazonas, San Mar- tin, Huanuco, Pasco, and Madre de Dios. Colombia; Ecuador; Peru; Bolivia. This species closely resembles S. ichtiosma (So- diro) C. Chr., a species known only from the west- ern cordillera of Ecuador but which eventually may be found in Peru. It differs from S. pellucido- punctata by the presence of a small, scalelike in- dusium associated with the sorus and having only lanceolate scales on the axes (i.e., it lacks filiform scales). Amazonas: Prov. de Bagua, Quebrada Tambillo (be- low km 280 of Maranon road), valley of Rio Maranon above Cascadas de Mayasi, Wurdack 2002 (GH, us, USM). Huanuco: Fundo Chela, Sinchono, Aguilar 927 (USM). Pasco: Paujil, near Puerto Bermudez, Leon 311 (USM). Madre de Dios: Prov. Manu, Atalaya, vicinity of Ha- cienda Amazonia, 2-3 km W of village, Foster & Wach- ter 7437 (MO, USM). 2. Stigmatopteris longicaudata (Liebm.) C. Chr., Bot. Tidsskr. 29: 300. 1909. Polypodium longicaudatum Liebm., Kongel. Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 5, 1: 209. 1849. LECTOTYPE (designated by A. R. Smith, Fl. Chiapas, part 2:213. 1981): Mexico, Veracruz, Baranca de Huitamalco, Liebmann Flora Mex. 737 (c- Folio Herbarium; ISOLECTOTYPE, K!). Dryopteris longicaudata (Liebm.) Maxon, Contr. U.S. Natl. Herb. 13: 18. 1909. Lamina 1 -pinnate. Pinnae mostly 14-20 cm long and 2.0-2.5 cm broad, linear to narrowly lanceo- late, 1 2-20 pairs, the margins entire or with cre- nate lobes cut less than V4 to the costae, the upper pinnae adnate to the rachis with a conspicuous decurrent tapering basal wing, lower pinnae free, subsessile or stalked, the stalk 2-5 mm long. Ra- chis and costae glabrous to densely scaly, the scales ovate to lanceolate, or (in Venezuela, Peru, and Bolivia) linear and fibrillose. Veins free, 3-6 per segment. Sori round, filiform scales present among the sporangia in plants from Venezuela, Peru, and Bolivia. Wet forests, 220-2 100(-2500) m, San Martin, Huanuco, Junin, Ucayali, Cuzco, and Madre de Dios. Southern Mexico; Guatemala; Costa Rica; Pan- ama; Venezuela; Peru; Bolivia. This species is distinctive in its decurrent, ta- pering wings from the pinna bases, which are usu- ally confluent in the distal V2 to !/3 of the lamina. San Martin: Mt. Guayrapurima, near Tarapoto, Spruce 4012 (BM, G, K, P). Huanuco: Tingo Maria, Tryon & Tryon 5330 (BM, F, GH). Junin: E of Quimiri Bridge, near La Merced, Killip & Smith 23906 (NY, us). Ridge E of Tingo Maria, Allard 22558 (us). Ucayali: Prov. Coronel Portillo (as Loreto), Sinchono, cerca de la Divisoria, Aguilar 861 (GH, USM). Sinchono (as Loreto), entre Tingo Maria y Pucallpa, Aguilar 865 (GH). Cuzco: Prov. Pau- cartambo, Cosnipata Valley, Rio Tono, first foothill ridge on road N of Patria, Wachter et al. 200 (F). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, 10-15 km NNW of Shintuya, transect to ridgetop, Foster et al. 10908 (F). Parque Nacional del Manu, Cocha Cashu Biological Station, Foster P-84-91 (F). 44 FIELDIANA: BOTANY 3. Stigmatopteris lechleri (Mett.) C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8, 6: 28. 1920. Phegopteris lechleri Mett., Fil. lechl. 2: 25. 1859. TYPE: Peru, Puno, "St. Gaban" (San Gaban), Lechler 2497 (holotype, B!; frag., BM!). Polypodium punctatum Hooker, Sp. fil. 4: 262, 1862, nom. illeg., not (L.) Swartz, 1802. LECTOTYPE (designated by Christensen, Kongel. Danske Vi- densk. Selsk. Skr. Naturvidensk. Afd., ser. 7, 10: 79. 1913): Peru, San Martin, Tarapoto, Mt. Guayrapurima, August 1856, Spruce 4719 (K.!; ISOLECTOTYPES, G!, P!; photos, F, MO of K). Polypodium prasinum Baker, Syn. fil. 312. 1867, nom. nov. for Polypodium punctatum Hooker and with the same type. Dryopteris prasina (Baker) C. Chr., Index fil. 285. 1905. Stigmatopteris prasina (Baker) C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Adf., ser. 7, 10: 79. 1913. Stigmatopteris ecuadorensis C. Chr., Kongel. Danske Vidensk. Selsk. Skr. Naturvidensk. Afd., ser. 8, 6: 29. 1920. TYPE: Ecuador (Andes of Quito), Sodiro (holotype, B!; isotypes, BM!, P). Lamina 2-pinnate-pinnatifid. Pinnae 25-40 cm long and 5-15 cm broad, widest at the base. Pin- nules 4-10 cm long and 1.0-2.3 cm broad, nar- rowly lanceolate-triangular, sessile to broadly ad- nate and decurrent, the lobes oblong to rectangular, entire to serrate apically. Rachis and costae stra- mineous or brown, scaly, the scales linear or nar- rowly triangular, tortuous, fibrillose. Veins un- branched or rarely (in the larger segments) branched, nearly reaching the margin. Sori round. Wet forests, 200-1700 m, San Martin, Huanu- co, Pasco, and Puno. Costa Rica; Venezuela; Colombia; Ecuador; Peru. This species, with its 2-pinnate-pinnatifid lam- ina, is the most finely divided in the genus. Huanuco: 25 km NE of Tingo Maria, La Divisoria, Moran 3696 (MO, USM). Pasco: Prov. Oxapampa, Gran Pajonal, north of Chequitavo, D. Smith 5082 (MO). 4. Stigmatopteris heterophlebia (Baker) R. C. Moran, comb. nov. Figure lla-b. Polypodium heterophlebium Baker, J. Bot. 22: 363. 1884. TYPE: Costa Rica, Prov. unknown, La- guna, 305 m, Harrison 59 (holotype, K!). Dryopteris heterophlebia (Baker) C. Chr., Index fil. 270. 1905. Lamina 1 -pinnate. Pinnae 10-16(-18) cm long and 2. 5-3. 5(— 4.5) cm broad, widest at or near the base, the margins entire to crenate, the base adnate and decurrent to or nearly to the next pinna pair, free pinnae pairs 1-3, the basal pinnae usually somewhat reduced, widest near the middle, stalked, the stalk 2—4 mm long. Rachis and costae tan, light brown, or stramineous, puberulent and scaly, the hairs less than 0. 1 mm long, erect, capitate-glan- dular (when fresh), the scales to 3 mm long, lan- ceolate, not fibrillose. Veins anastomosing irreg- ularly. Sori discrete or confluent where the veins anastomose. Wet forests, 100-1900 m, Loreto. Nicaragua to Peru. Differing from the closely related S. alloeoptera (Kunze) C. Chr., which grows in Venezuela and Colombia, by the glandular hairs on the axes and fewer pinna pairs. Loreto: Santa Rosa, lower Rio Huallaga below Yuri- maguas, Killip & Smith 28984 (NY). 5. Stigmatopteris opaca (Baker) C. Chr., Kongel. Danske Vidensk. Selsk. Skr., Naturvidensk. Afd., ser. 7, 10: 78. 1913. Meniscium opacum Baker, J. Bot. 166. 1877. TYPE: Ecuador, Andes of Quito, Sodiro 54/3 (holotype, K!; photos, BM, GH, us, all of K). Polypodium oligophlebium Baker, Syn. fil. ed. 2, 506. 1874, nom. illeg., not Kunze, 1850. TYPE: Peru, San Martin, Tarapoto, Cerro de Guayrapurima, Spruce 4653 (holotype, K!). Dryopteris christii C. Chr., Index fil. 257. 1905, nom. nov. for Meniscium opacum, and with the same type, not Dryopteris opaca (Don) C. Chr. 1905. Dryopteris paucinervata C. Chr., Index fil. 283. 1905, nom. nov. for Polypodium oligophlebium Baker, and with the same type. Lamina 1 -pinnate, opaque, the internal glands not visible. Pinnae 12-27 cm long and 3-5 cm broad, widest at or just above the base, the margins entire to crenate, the base sessile, the upper pinnae with a short decurrent base, the basal pinnae near- ly equalling the above pinnae, widest near the mid- dle, sessile. Rachis and costae lacking hairs, scaly, the scales lanceolate to linear or amorphous. Veins anastomosing, those of the sterile leaf with a single excurrent vein projecting from the apex where the two lateral veins join. Sori round, oblong, or ar- cuate. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 45 2mm ""Oaf 5cm FIG. 1 2. Cyclodium trianae: a, lamina; b, stem and petiole base. Cyclodium meniscioides var. meniscioides: c, lamina apex; d. fertile pinna; e, sori. (a, b from Ellenberg 3257, Ecuador, GH; c, d from Fiebrig 6344, Paraguay, F, e from Schunke V. 5610, F.) 46 FIELDIANA: BOTANY Wet forests, 250-600 m. Ecuador and Peru. In Peru known only from Spruce's type collec- tion ofPolypodium oligophlebium from San Mar- tin. XII. Cyclodium Cyclodium Presl, Tent, pterid. 85. 1836. LEC- TOTYPE (designated by John Sm., Hist. fil. 203. 1875): Aspidium confertum Kaulf. = Cy- clodium meniscioides (Willd.) Presl. Figure 12a-b. Peltochlaena Fee, (Mem. foug. 5) Gen. fil. 289. 1852, nom. provis., illegit. Stem usually rather short-creeping and moder- ately stout, to long-creeping (to 5 m when climb- ing), bearing scales. Leaves ca. 30 cm to 2 m long, petiole continuous. Lamina 1 -pinnate to 2-pin- nate-pinnatifid or rarely 3 -pinnate at the base, or rarely simple, usually slightly short-pubescent or scaly, monomorphic to dimorphic and then the fertile more erect, longer, and with less expanded segments. Veins free or anastomosing, with or usu- ally without included free veinlets. Sori roundish, borne on the veins, not paraphysate, covered by often fugacious indusia which are peltate or or- bicular-reniform with a narrow sinus. Spores el- lipsoidal to spheroidal, monolete, slightly to strongly ridged, usually finely papillate. Cyclodium is a tropical American genus of ten species. It was formerly treated in Stigmatopteris (Tryon & Tryon, 1982), but the revision by Smith (1986) clearly shows that it is distinct from, and perhaps not closely related to, that genus. There are two species in Peru, and an additional one may occur there (see Comments). This treatment has been adapted from that of Smith (1986) and has been prepared with the col- laboration of Dr. A. R. Smith. References SMITH, A. R. 1986. Revision of the Neotropical fern genus Cyclodium. Amer. Fern J., 76: 56- 98. TRYON, R. M., AND A. F. TRYON. 1982. Stig- matopteris, pp. 519-524 in Ferns and allied plants, Springer- Verlag, New York. Key to Species of Cyclodium a. Veins all free; lamina 1 -pinnate-pinnatifid to 3-pinnate at the base, the apex gradually reduced . . . l.C. trianae a. Veins regularly anastomosing; lamina 1 -pinnate, with a conform (or nearly) apical segment . 2. C. meniscioides 1. Cyclodium trianae (Mett.) A. R. Sm., Amer. Fern J. 76: 92. 1986. Figure 12a-b. Aspidium trianae Mett., Ann. Sci. Nat. Bot. 5, 2: 243. 1864. TYPE: Colombia, Prov. Barbacoas (Nari- no), via de Tuquerres, Triana "32" (holotype, B; isotype, BM; both seen by A. R. Smith; photos of BM at uc, us). Nephrodium firmifolium Baker, Syn. fil., ed. 2, 501. 1874. TYPE: Peru (San Martin), Mt. Guayapu- rima, near Tarapoto, Spruce 4662 (holotype, K; isotypes BM, BR, p; all seen by A. R. Smith). Terrestrial, stem creeping. Leaves monomor- phic or only slightly dimorphic, gradually reduced at the apex. Lamina 1 -pinnate-pinnatifid to usu- ally 2-pinnate or 2-pinnate-pinnatifid, or 3-pin- nate only at the base. Veins free. Sori with orbic- ular-reniform indusia with a narrow sinus, or rarely peltate. In dense primary forests or partly cleared for- ests, 300-900 m, Lambayeque and Loreto, south to Junin. Panama, south to Peru. Var. trianae occurs throughout the range of the species. In South America it is generally confined to the Amazonian side of the Andes. Var. cho- coense A. R. Sm. is only in Panama and the Pacific side of the Andes in Colombia. The only repre- sentative in Peru is var. trianae. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 47 Lambayeque: Puerto Nazareth, 5 km from Olmos, El- lenberg 3449 (GH). Loreto: Balsapuerto, Killip & Smith 28540 (GH, NY, us). Pasco: Prov. Oxapampa, Rio Pal- cazu, cerca de Iscozacin, Leon 712 (GH). Prov. Oxapam- pa, Palcazu valley, Iscozacin, Foster 9498 (MO). Junin: Prov. Satipo, above Pichanaki, Leon 216 (GH). the fertile then more erect and with shorter and narrower pinnae. Lamina 1 -pinnate or rarely sim- ple, with a conform (or nearly) apical segment, the pinnae entire to strongly crenulate. Veins anas- tomosing, usually with included free veinlets. Sori with peltate indusia. 2. Cyclodium meniscioides (Willd.) Presl, Tent, pterid. 85. 1836. Terrestrial or epiphytic and short-climbing to 2 m. Leaves subdimorphic to strongly dimorphic, Trinidad; and widely distributed in tropical South America. In addition to the two varieties in Peru, there is var. rigidissimum (C. Chr.) A. R. Sm., which rarely occurs in Guyana and perhaps in Venezuela. Key to Varieties a. Margins of the pinnae entire, sinuate, crenulate to crenate 2a. var. meniscioides a. Margins of the pinnae, especially toward the apex of the sterile ones, sharply serrate 2b. var. paludosum 2a. Cyclodium meniscioides var. meniscioides. Figure 12c-e. Aspidium meniscioides Willd., Sp. pi. ed. 4, 5: 218. 1810. TYPE: Brazil, Hoffmannsegg (holotype, B!, Herb. Willd. 19737; photo, GH). Stigmatopteris meniscioides (Willd.) Kramer, Proc. Kon. Nederl. Akad. Wetensch. C, 71: 521. 1968. In dense primary forests, in swampy forests where sometimes epiphytic, or in partly disturbed forests, rarely on fallen logs or epiphytic to 2 m, 100-650 m, Amazonas south to Madre de Dios. Range of the species. Amazonas: Near Pongo Mori, Rio Comain, Berlin 949 (MO, uc). Prov. Bagua, 34 km NE of Chiriaco, Barbour 4401 A (MO). San Martin: Prov. Mariscal Caceres, To- cache Nuevo, J. Schunke V. 5610 (F, us). 23 km S of Nuevo San Martin, Gentry et al. 37573 (MO). Loreto: 7 km SW of Iquitos, Croat 18579 (GH, MO, uc). Above Pongo de Manseriche, Mexia 6 193 a (GH, uc, us). Mishu- yacu, near Iquitos, Klug 241 (F, us). Pasco: Puerto Ber- mudez (as Junin), Killip & Smith 26566 (us). Prov. Oxapampa, Palcazu valley, D. Smith 3733 (MO, uc). Ucayali: Vicinity of Aguaytia, Mathias & Taylor 5138 (F). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10704 (F). 2b. Cyclodium meniscioides var. paludosum (Morton) A. R. Sm., Amer. Fern J. 76: 87. 1986. Dryopteris paludosa Morton, Bull. Torrey Bot. Club 66: 50. 1939. TYPE: Colombia, Antioquia, Puer- to Berrio, Pennell 3723 (holotype, NY; frag., us, both seen by A. R. Smith). Stigmatopteris paludosa (Morton) R. & A. Tryon, Rhodora83: 136. 1981. 160 m, Loreto. Colombia and Peru. Loreto: Bersalles-Iquitos, Vargas 11476 (GH). Comments Cyclodium guianense (Klotzsch) Gomez, Phyto- Iogia60: 371. 1986. Aspidium guianense Klotzsch, Linnaea 20: 364. 1847. TYPE: "British Guiana," Schomburgk 1157 (ho- lotype; B?; isotypes, K, uc, seen by A. R. Smith). Stigmatopteris guianensis (Klotzsch) C. Chr. Ind. fil suppl. 3: 174. 1934. This species occurs in Trinidad, northern South America, and in the Amazon basin of Colombia, and may occur in Peru. It has all of the veins free, as in C. trianae, but has a 1 -pinnate lamina rather than a 2-pinnate to 2-pinnate-pinnatifid one, and peltate indusia rather than orbicular-reniform ones, these with a narrow sinus. Only rarely does C. trianae have peltate indusia. 48 FIELDIANA: BOTANY XIII. Polystichum Polystichum Roth, Tent. fl. germ. 3: 31, 69. 1799. TYPE: Polystichum lonchitis (L.) Roth (Poly- podium lonchitis L.). Figure 13. Plecosorus Fee, (Mem. fam. foug. 5) Gen. fil.: 150. 1852. TYPE: Plecosorus mexicanus Fee = Ple- cosorus speciosissimus (Kunze) Moore = Polys- tichum speciosissimum Kunze) R. & A. Tryon. Stem decumbent to erect, often small, to stout, bearing stramineous to blackish scales. Leaves 5 cm to often 1 m or rarely 3 m long, petiole con- tinuous. Lamina usually 1 -pinnate, 2-pinnate, or to 3-pinnate-pinnatifid, rarely entire to pinnatifid, more or less scaly, often densely so, monomorphic or rarely dimorphic, adaxial side of the costa sul- cate to nearly flat. Veins free. Sori round, borne on the veins, not paraphysate, covered by per- sistent to fugacious, peltate indusia, or (in all Pe- ruvian species) exindusiate. Spores ellipsoidal to spheroidal, monolete, saccate, cristate, or ridged, often spinulose and frequently perforate. Polystichum is a large, nearly worldwide genus of about 1 50 species. There are perhaps 25 species in America and four in Peru. The species of An- dean Polystichum are not well known, and the following treatment is tentative. Species of Polys- tichum are evidently variable genetically and cy- tologically and they also vary in different envi- ronments. Population studies, cytological studies, and broad morphological comparisons are needed in order to recognize and define the taxa (Bar- rington, 1985). Currently unrecognized hybrids may be an important element in the apparent in- tergradation of taxa. Although characters of the scales of the leaf axes are emphasized in the key to species by Smith (1985), among others, these have been found to be usually variable in Peru and are not generally employed in this treatment. However, these struc- tures need further investigation and may be more important than now considered. Among the synonyms, only those names based on Peruvian material and those that are in general use are included. The names of many species rec- ognized, especially from Ecuador, Bolivia, and southeastern Brazil, have not otherwise been in- cluded. Although Christensen (Index fil. 727. 1906) in- dicated that the first publication of the generic name Polystichum was in Arch. Bot. (Leipzig) 2( 1 ). 1 799, Taxonomic Literature-2 indicates that Roth's publication was in 1 799 while Arch. Bot. (Leipzig) 2(1) was in 1800. This treatment has been prepared with the per- spective and critical collaboration of Dr. David S. Barrington, who has seen some of the types cited. The key will be adequate for the identification of many complete collections. However, in some cases, a concordance of the majority of the char- acters mentioned may be necessary for identifi- cation, and rarely materials may represent a vari- ation that is not accounted for in the key. References BARRINGTON, D. S. 1985. The present evolu- tionary and taxonomic status of the fern genus Polystichum: The 1984 Botanical Society of America Pteridophyte Section Symposium. Amer. Fern J., 75: 22-28. SMITH, A. R. 1985. Pteridophytes of Venezuela, an annotated list, pp. 1 89-1 92, published by the author, Department of Botany, University of California, Berkeley. TRYON, R. M., AND A. F. TRYON. 1982. Polys- tichum, pp. 524-538, in Ferns and allied plants, Springer- Verlag, New York. Key to Species of Polystichum a. Petiole with scales few or not persistent above its base, varying to densely scaly, scales mostly small, to ca. 0.5 cm long, elongate, rarely larger and broad; rachis with flattish scales and/or fibrils; pinnules with revolute margins or plane (flat) b b. A scaly bud usually present in the axil of a pinna near the apex of the lamina, or more than one present; apical portion of the lamina usually prolonged, with a 1 -pinnate portion 5-10 cm long with pinnae mostly 1-2.5 cm long, or longer 2. P. platyphyllum TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 49 FIG. 13. Polystichum pycnolepis: a, habit. Polystichum platyphyllwn: b, habit; c, distal pinnae, with proliferous bud; Polystichum montevidense var. montevidense: d, pinnule, (a from Madison 1034, GH; b, c from Macbride 4034, F, d from Soukup 4329, F.) 50 FIELDIANA: BOTANY b. Apex of the lamina lacking a scaly bud, usually gradually reduced, if prolonged the 1 -pinnate portion short with most pinnae less than 1 cm long c c. Pinnules definitely mucronate, usually plane (flat), or if the margins revolute then the apical mucro plane; lamina 2-pinnate; if the lamina less than 8 cm broad then the basal pinnae reflexed and/or the lamina apex prolonged, the apex of the pinnae acute to attenuate; or if the lamina over 8 cm broad then the basal pinnae patent or reflexed 1 . P. montevidense c. Pinnules sometimes definitely mucronate and plane, usually not or hardly mucronate, often with the margins and apical mucro revolute; lamina 2-pinnate to 3-pinnate; the lamina usually less than 8 cm broad and the basal pinnae patent or ascending, the lamina apex not or hardly prolonged, and the apex of the pinnae obtuse to subacute; or if the lamina over 8 cm broad, then the basal pinnae ascending 3. P. orbiculatum a. Petiole with scales usually persistent above its base, mostly large, ca. 1-2 cm long and sometimes to 1 cm broad; rachis, at least at the base, usually with some brown or darker, involute or twisted scales, these often also on the petiole; pinnules with revolute margins; apex of the lamina lacking a scaly bud 4 P. pycnolepis 1 . Polystichum montevidense (Sprengel) Rosenst., Hedwigia46: 111. 1906. Leaves ca. 10 cm to 1.5 m long. Petiole scales sparingly persistent or persistent, especially at or near the base, or petiole nearly devoid of scales; scales brown to dark brown or darker, less than 1 cm long or rarely longer. Lamina 2-pinnate to rare- ly 3-pinnate at the base of some pinnae; ca. 2—40 cm broad; the apex lacking a bud, gradually to sometimes rather abruptly reduced, or prolonged. Rachis bearing a few to many light brown to dark brown fibrils and narrow scales, or sometimes broader ones, or nearly devoid of scales. Pinnae mostly patent, the apex acute to usually attenuate and deeply pinnatifid to the tip; basal pinnae re- duced or not, patent or reflexed; pinnules plane (flat) or nearly so. Key to Varieties of Polystichum montevidense a. Leaves ca. 50 cm to 1.5 m, mostly 60-80 cm long; lamina ca. 10 to 40 cm broad, mostly 2 to 2'/2 times as long as broad, the apex gradually to sometimes rather abruptly reduced, the portion beyond the 2-pinnate portion shorter than the longest pinna; basal pinnae not or slightly reduced, patent or somewhat ascending la. var. montevidense a. Leaves ca. 10-90 cm, mostly ca. 30 cm long; lamina ca. 2-1 1 cm, mostly ca. 6 cm broad, mostly 3 to 4 times as long as broad, the apex usually prolonged, the portion beyond the 2-pinnate portion longer than the longest pinna; basal pinnae reduced and/or reflexed 1 b. var. nudicaule 1 a. Polystichum montevidense var. montevidense. Figure 13d. Polypodium montevidense Sprengel, Syst. veg., ed. 16, 4: 59. 1827. TYPE: Uruguay, Monte Video, Sello (not located). Polystichum lehmannii Hieron., Hot. Jahrb. Syst. 34: 452. 1904. TYPE: Colombia, Prov. Cauca, Los Motilones, Lehmann 3674 (holotype: B; isotype, us!). Polystichum mexiae Copel., Univ. Calif. Publ. Bot. 19: 299. 1941. TYPE: Ecuador, Prov. Pichincha, Nono to Mindo, Mexia 7679 (holotype, uc; isotypes, F!, GH!). In dry rocky places and on open hillsides, more often in forests in rocky places, and in cloud for- ests, 1800-3600 m, Piura to Puno. Venezuela and Colombia, south to Bolivia, east through Argentina to Uruguay and southeast Bra- zil; possibly in Central America. Recent studies have usually confined Polysti- chum montevidense to northwestern Argentina and eastward to southeastern Brazil. However, Peru materials are not separable, and the species is best regarded as an Andean one, with an extension east- ward. Intergradation between var. montevidense TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 51 and var. nudicaule indicates that the two are not distinct species. In spite of the several characters presented in the key, there is considerable overlap between the two taxa. Plants with the leaves especially scaly on the rachis and pinna-rachises have been referred to Polystichum montevidense var. squamulosum (Hi- eron.) Hieron., but this seems appearance to be a part of the normal variation of the species in Peru. Polystichum yungense Rosenst. of Bolivia is prob- ably a large, well-developed variation of this spe- cies. Piura: Ayabaca, Soukup 4329 (F, us). Cajamarca: Prov. Contumaza, Cascas-Contumaza, Lopez et al. 9109 (GH). Prov. Hualgayoc, Monte Seco, Soukup 3816 (F, us). Prov. San Miguel, Cerro Quillon, Mostacero et al. 1293 (F, GH). Amazonas: Prov. Bagua, SE of La Peca, Barbour 3696 (uc), 3984 (MO). San Martin: Lamas, Bel- shaw 3428 (GH, uc, us). Prov. Mariscal Caceres, Rio Abiseo National Park, Young & Leon 4586 (F). Huanuco: Mito, Macbride & Featherstone 1698 (F, GH, us). Yana- no, Macbride 3827 (F, us). Pasco: Prov. Oxapampa, San Alberto, van der Werff et al. 8433 (MO, uc). Ayacucho: Between Huanta and Rio Apurimac, Killip & Smith 23239 (GH). Cuzco: Prov. Urubamba, Chupani, Vargas 11 126 (GH). Prov. La Convention, between Rumichurco and Alcobamba, Davis et al. 1227 (F, GH). Puno: Prov. San- dia, entre Sandia y Cuyocuyo, Ferreyra 16818 (GH, USM). Ib. Polystichum montevidense var. nudicaule (Rosenst.) Tryon, comb. nov. Polystichum nudicaule Rosenst., Repert. Spec. Nov. Regni Veg. 11: 56. 1912. SYNTYPES: Bolivia, Unduavi, 3400 m, Buchtien 2657, 2658, 2659, 2662 (not located); ISOSYNTYPES, Buchtien 2657 (us), 2655 (uc!, us), 2659 (us). Rocky crevices, on cliffs, at base of rocks, and in boggy places in heath, 1 800-4000 m, Piura to Puno. Venezuela and Colombia, south to Bolivia; per- haps in southeastern Brazil and Chile. This variety is distinguished especially by its prolonged lamina apex and the usually reflexed and reduced basal pinnae. It intergrades with var. montevidense and perhaps represents a variation of it growing under less favorable conditions. Po- lystichum wolfii Hieron. from Bolivia is probably this variety. A few collections have some leaves on the stem with closely spaced and nearly entire pinnules and also some that have widely spaced and rather la- ciniate pinnules. Piura: Prov. Huancabamba, above Cachaque on road to Huancabamba, Hutchison 1643 (GH, uc). Cajamarca: Prov. Cajamarca, Cajamarca-Bambamarca, Smith & Vdsquez 3477 (uc). Prov. Contumaza, Las Achiras, Sa- gdstegui 3887 (GH, HUT). Amazonas: Prov. Chachapoyas, NNE of Diosan, Wurdack 1596 (F, GH, uc, us). La Liber- tad: Prov. Santiago de Chuco, Laguna El Toro, D. Smith 2300 (GH). San Martin: Rio Abiseo National Park, Young & Leon 4401 (F). Huanuco: Cani, Macbride 3459 (F, us). Tambo de Vaca, Macbride 4420 (F, us). Lima: Prov. Huarochiri, Dist. San Mateo, Sounders 324 (F, uc). Rui- nas de Cantamarca, Coronado 297 (GH, uc). Junin: In- cahuasi, Soukup 3177 (F). Near Huancayo, Killip & Smith 23366 (us). Ayacucho: Between Huanta and Rio Apu- rimac, Killip & Smith 22254 (us). Apurimac: Prov. An- dahuaylas, Dist. Chincheros, Sounders 737 (GH). Prov. Abancay, Curahuaci, Marin 1961 (F). Cuzco: Prov. Cuz- co, Saxihuaman, Cuzco, Tryon & Tryon 5359 (GH, us). Taucca, Davis et al. 1540 (F). Puno: Granja Salcedo, Soukup 66 (F). Have, Soukup 872 (F). 2. Polystichum platyphyllum (Willd.) Presl, Tent, pterid. 84. 1836. Figure 13b-c. Aspidium platyphyllum Willd., Sp. pi., ed. 4, 5: 255. 1810. TYPE: Venezuela, Caracas, Bredemeyer (holotype; B!, Herb. Willd. 19879; photo, GH). Leaves ca. 40 cm to 1.5 m long. Petiole scales sparingly or not persistent, light brown to brown, or especially near the base dark brown; less than 1 cm long, rarely longer. Lamina 2-pinnate, or rarely nearly 3 -pinnate at the base of some pinnae, ca. 10-30 cm broad; the apex with the axil of a pinna bearing a scaly bud, or more than 1 axil bearing a bud, the apex usually prolonged, very gradually, or sometimes abruptly reduced. Rachis bearing mostly few, sometimes many, light brown to brown fibrils and narrow scales, some broader scales also often present. Pinnae patent, apex acute or to attenuate and deeply pinnatifid to the tip; basal pinnae the longest to somewhat reduced, usually slightly ascending or patent, rarely re- flexed; pinnules plane (flat). In primary forests, on steep forested slopes, in ravines and along stream banks, and on open, rocky, moist road banks, 600-2800 m, Cajamarca and Amazonas to Puno. Southern Mexico and Central America; Jamai- ca; Venezuela and Colombia, south to northwest- ern Argentina and east to southeastern Brazil. Nearly all leaves of this species have a bud near the apex of the lamina, and perhaps all plants have at least one leaf with a laminar bud. In a collection 52 FIELDIANA: BOTANY from a large population (Try on & Try on 5404, GH) all leaves except one very small one have a bud. In old leaves, this bud may develop into a small plantlet, with roots and small leaves. Other dis- tinctive characters of this species are mentioned in the key. Cajamarca: Prov. Santa Cruz, Monte Seco, Sagdstegui et al. 12407 (F, MO). Prov. Hualgayoc, Hacienda Taulis, Hutchison & Bismark 6331 (F, GH, MO, uc, us). Ama- zonas: Prov. Chachapoyas, Quebrada Molino, Wurdack 771 (F, GH, uc, us). Prov. Bagua, 12 km E of La Peca, Barbour 2400 (F, MO, uc). San Martin: Near Tarapoto, Spruce 3965 (BM, GH, us). Between Tarapoto and Yu- rimaguas, Croat 58100 (F). Huanuco: Mima, Bryan 548 (F, GH, us), Macbride 4034 (F, GH, us). Prov. Huanuco, near Riachuela Contalagua, Mexia 8297 (F, GH, MO, uc, us). Pasco: Pichis Trail (as Junin), between San Nicolas and Azupizu, Killip & Smith 26120 (us). Prov. Oxapam- pa, 5 km SE of Oxapampa, D. Smith 2915 (GH). Junin: Chanchamayo valley, C. Schunke 13 (F, us). Near La Merced, Killip & Smith 23911 (GH, us). Prov. Chancha- mayo, cerca a Monobamba, Fernandez & Vargas 204 (USM). Ayacucho: Between Huanta and Rio Apurimac, Killip & Smith 22443 (us). Apurimac: Prov. Andahuay- las, Alavarado (USM). Cuzco: Prov. Urubamba, near town of Machu Picchu, Tryon & Tryon 5404 (BM, F, GH, uc, us). Prov. Urubamba, camino a Winayhuayna (Huinay- huayna), Chavez 3440 (GH, MO). Puno: Prov. Carabaya, Vargas 17529 (GH). 3. Polystichum orbiculatum (Desv.) Remy, in Gay, Hist. Fis. Pol. Chile, Bot. (Fl. Chilena) 6:515. 1853 (as Polystichum orbiculare by Christ, Ark. f. Bot. 4[12]: 3. 1905). Leaves ca. 1 5 cm to 1 m or more long. Petiole scales persistent or sparingly so; usually brown to dark brown near the base and brown to whitish beyond; scales less than 1 cm long, rarely longer. Lamina 2-pinnate to 3-pinnate, or rarely small ones 1 -pinnate-pinnatifid, ca. 2-25 cm broad; the apex lacking a bud, shortly acute to gradually re- duced and attenuate. Rachis bearing a few to many brown to whitish scales, these broad or sometimes narrow and grading into fibrils. Pinnae mostly as- cending, less often patent, the apex obtuse to less often subacute; basal pinnae somewhat reduced or not, mostly patent, sometimes ascending; pinnules plane (flat) or usually with revolute margins and apex. The new names of Pteridophyta in Gay, Flora Chilena 6: 470-549 have been variously attributed to Gay, to Fee, to Remy, to Fee & Remy, and to Remy & Fee. All of the new nomenclature is ev- idently to be attributed to Remy (as Remy, in Gay. Fl. Chil. 6:) and we have followed this citation. In his Cat. method, chron. public. A. L. A. Fee, 1 863, Fee does not mention any contribution to the Flo- ra Chilena of Gay, and in 1858, Sturm in Abhandl. natur.-hist. ges. Nurnberg 2: 151-202 (reprint 1- 52) nine of the 1 1 new species are credited to Remy. Looser in Rev. Universit. (Santiago, Chile) 1 5(7): 694-717. 1930, credits the treatments of pterido- phytes to Remy, although in later publications, on Blechnum for example, he cites species authors as Remy & Fee. It seems that the original treatment was prepared by Remy and that Fee had some editorial role with the manuscript. Key to Varieties of Polystichum orbiculatum a. Lamina usually 2-pinnate, rarely 1 -pinnate-pinnatifid or 3-pinnate only at the base of some pinnae 3a. var. orbiculatum a. Lamina 3-pinnate well beyond the base of the pinnae 3b. var. boboense i. Polystichum orbiculatum var. orbiculatum Aspidium orbiculatum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Na- turk. 5: 321. 1811. TYPE: "America australi" (holotype, P!, Herb. Desvaux; photo, GH, us). Nephrodium polyphyllum Presl, Reliq. haenk. 1: 37. 1825. TYPE: Peru, Haenke (holotype, PR! or PRC; photo, BM of PR). Nephrodium trapezoides Presl, Reliq. haenk. 1: 37, /. 6,f. 1. 1825. TYPE: Peru, Haenke (holotype, PR or PRC?; isotype, K!). Polypodium rigidum Hooker & Grev., Icon. fil. /. 163. 1830, not Lam. & DC. 1805. TYPE: "Andium TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 53 Peruvianorum" (Ecuador), Monte Pichincha, Jameson (holotype, K!). Polystichum polyphyllum (Presl) Presl, Tent, pterid. 83. 1836. Polystichum haenkeanum Presl, Tent, pterid. 83. 1 836, nom. nov. for Nephrodium trapezoides Presl, not Polystichum trapezoides (Sw.) Presl, and with the same type. Jamesonia paleacea Kunze, Hot. Zeit. 1844: 739, not Polystichum paleaceum (Borkh.) Schwarz, 1949. TYPE: Venezuela, Prov. Merida (also "Cara- cas"), Linden 505 (holotype, B; isotypes, E!, GH!, K!; photos, GH of BR and BM may be this species, or they and the holotype may be P. pycnolepis). Polypodium saxatileK\o\zsch, Linnaea 20: 383. 1847. TYPE: Peru, Herb. Ruiz 65 (holotype, B?). Hi- eronymus (Hedwigia, 46: 358. 1907) treats this as a variety of Polystichum orbiculatum. Polypodium crenatodentatum Klotzsch, Linnaea 20: 384, 1847. TYPE: Peru, Ruiz 73 (holotype, B?) Hieronymus (op. cit., p. 357) treats this as a va- riety of Polystichum orbiculatum. Commonly growing in rocky, shaded places, at the edge of rocks, or on cliffs, on sandstone and on limestone, less often on grassy or shrubby hill- sides, or at high altitudes in the puna, 2700-4900 m, mostly 3200-^*000 m, Lambayeque to Puno. Southern Mexico; Costa Rica; Venezuela and Colombia south to Bolivia. Polystichum orbiculatum is a highly variable species with most of the more distinctive varia- tions included in var. orbiculatum. One extreme, especially characteristic of the puna, has erect leaves with strongly ascending pinnae that are closely set and the pinnules have strongly revolute margins. Another extreme has spreading leaves with mostly patent and spaced pinnae and the pinnules are flat. Another variation has the lamina relatively broad and the pinnae and pinnules sharply acute to acu- minate. These all intergrade to an extent that im- plies they are variations within one taxon. The label on Soukup & Guillen 5386 (us): "Iqui- tos" (Loreto) is undoubtedly an error. The species grows at much higher altitudes than the ca. 100 m of Iquitos. Lambayeque: 7 km NW of Incahuasi, Sillon A Skill- man 4110(r, GH, uc). Cajamarca: Prov. Hualgayoc, cer- ca a Hualgayoc, Ferreyra 8539 (GH, USM). Prov. Con- tumaza, Casabamba. Sagdstegui et al. 10721 (F). Amazonas: NNE of Diosan. Wurdack 1597 (F, GH, uc, us). La Libertad: Prov. Santiago de Chuco, above Cachi- cadin. Stork & Morton 9976 (F, uc, us). Prov. Pataz, entre Retamas y La Paccha, Lopez & Sagdstegui 3592 (GH). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Young & Leon 4402 (F). Ancash: Prov. Pallasca. Conzuzo, Lopez 1165 (GH, us). Prov. Yungay, above Yungay, Tryon A Tryon 6557 (F, GH). Lima; Be- tween Parac and Toncuyo, Coronado 310 (GH, uc). Prov. Huarochiri, Mina Perdida, Cerrate 2040 (USM). Pasco: Puna de Huaron, Humbert 30894 (GH). Prov. Pasco, Huayllay, Boeke 1086 (AAU, MO). Jonin: Near Huancayo, Killip A Smith 23363 (GH, us). Ca. 1 8 km SSE of Tarma, Iltis et al. 129 (GH, uc, us). Huancav elica: Prov. H uan- cavelica. westward from Huancavelica, Stork A Horton 10851 (F, uc, us). Prov. Tayacaja, arriba de Toca, Tovar 2018 (GH, USM). Ayacucho: Pumaccuanca pass, West 3658 (uc). Prov. Huanta, Ayacucho-Huanta, Sounders 727 (GH). Apurimac: Bosques de Ampay, Vargas 1061 (GH). Cuzco: Prov. Urubamba. Machu Picchu, Peyton A Pey- ton 1055 (GH, MO). Prov. Espinar, Hacienda Chachachi. Vargas 10812 (GH). Arequipa: Cailloma, Mutter A Mul- ler 2151 (GH). Puno: Salcedo, Soukup 275 (F, GH, uc). Capachica Peninsula, Tutin 1071 (A, BM). 3b. Polystichum orbiculatum var. boboense i Hi- eron.) Tryon, comb. nov. Polystichum boboense Hieron., Hedwigia 46: 358. /. 8,f. 25. 25a. 1907. TYPE: Ecuador, Rio Bobo, near Tulcan. Stubel 353, in part (holotype, B). Polystichum sodiroi Christ, Ark. f. BoL 4( 1 2): 4. 1 905, TYPE: Ecuador, Quito, Sodiro (not located). In ravines, 2800-3000 m, Piura. Ecuador and Peru. This variety is the most distinctive among the variations of Polystichum orbiculatum. The lam- ina is fully 3-pinnate and often the leaves are large, up to 1 mm or more long. Intermediates between the two varieties, with the lamina 3-pinnate only at or near the base of some pinnae, are present throughout most of the range of var. orbiculatum. Piura: Prov. Huancabamba, above Huancabamba. Hutchison 1608 (uc). Prov. Huancabamba, Huanca- bamba-Cuello del Indio, Sagdstegui et al. 8256 (F, uc; at MO this collection is toward var. boboense). 4. Polystichum pycnolepis (Klotzsch) Moore, In- dex fil. 92. 1858. Figure 13a. Aspidium pycnolepis Klotzsch, Linnaea 20: 365. 1847. SYNTYPES: Venezuela, Merida, Paramo de Cu- lata, Aforitz 295, Moritz 296; Karasten "Coll. II, a, b". (B?); ISOSYNTYPES: Moritz 295 (E!), (BM!, E!, K!; photo, GH of P), Moritz s.n. (GH!). Aspidium gelidum Klotzsch, Linnaea 20: 365. 1847. TYPE: Venezuela, Merida, Paramo de Culata. Moritz 294 (holotype, B?; isotypes, BM, K!; photo, GH of BM). Polypodium cochleatum Klotzsch, Linnaea 20: 388. 1847. TYPE: Peru, Ruiz 63 (holotype, B?). Polystichum gelidum (Klotzsch) Fee, (Mem. foug. 5) Gen. fil. 278. 1852. Polystichum cochleatum (Klotzsch) Hieron., Jahrb. BoL Syst. 34: 452. 1904. 54 FIELDIANA: BOTANY Leaves ca. 35 cm to 1 m long. Petiole scales more or less persistent, especially toward the base, dark brown to atropurpureous, or some light brown, often many ca. 2 cm long and to 1 cm broad. Lamina 2-pinnate, very rarely nearly 3-pinnate at the base of some pinnae, ca. 8-20 cm broad, rarely 5-7 cm broad; the apex lacking a bud, gradually to rather abruptly reduced. Rachis bearing usually very many brown fibrils and brown or darker, long, often involute or twisted, narrow scales, some lanceolate scales and broader ones usually also present. Pinnae patent to strongly as- cending, the apex subacute to acuminate and deep- ly pinnatifid to the tip; basal pinnae reduced or not, usually ascending, sometimes patent or re- flexed; pinnules with revolute to strongly revolute margins, the tip usually revolute. Most commonly growing on open, rocky hill- sides, in rock crevices, and at the edge of large rocks, also on grassy or shrubby slopes and in heath lands, 1 700 to usually 3000-4400 m, Ama- zonas to Puno. Venezuela and Colombia south to Bolivia. Aspidium pycnolepis is evidently the correct ba- sionym, among those names proposed by Klotzsch in the same publication. Aspidium gelidum was reduced to Phegopteris pycnolepis by Mettenius (Abh. SenckenbergNaturf. Ges. 2; 295. 1858; Uber einige Farngattugen, 4; 1 1. 1859) and Polypodium cochleatum (as Phegopteris cochleata Mett.) was reduced to Polypodium pycnolepis by Hooker (Sp. fil. 4: 248. 1862). This species is characterized by dense, brown fibrils on the rachis and narrow, involute or twist- ed, usually brown or darker scales, especially to- ward the apex of the petiole and on the rachis. The usually large petiole scales are persistent. Amazonas: Prov. Chachapoyas, Cerro Campanario, Wurdack 1589 (F, GH, uc, us). La Libertad: Between Huamachuco and Cajambamba, Correll & Smith P924 (GH). Huanuco: Tambo de Vaca, Macbride 4369 (F, GH, us). Mito, Macbride & Featherstone 1699 (F, us). Apo- rimac: Prov. Abancay, Dist. Abancay, Sounders 762 (GH). Cuzco: Prov. Calca, Vargas 15583 (GH). Prov. Urubam- ba, Chincheros, Taucca, Davis et al. 1572 (F). Puno: Prov. Sandia. S of Limbani, Metcalf 30436 (F, GH, MO, uc, us). Prov. Huancane, Occa Pampa, Shepard 74 (GH). Comments Polystichum muricatum (L.) Fee, (Mem foug. 5) Gen fil. 278. 1852. Polypodium muricatum L., Sp. pi. 1093. 1753. TYPE: Haiti, Petiver, Pteri-graphia Amer., no. 53, /. 1, f. 6. 1712. (evidently a poor copy of Plumier, Traite foug. Amer. t. 39). Polystichum moritzianum (Klotzsch) Hieron., Hed- wigia46: 354. 1907. This species has been commonly recorded from Peru; however, the collections are mostly either P. platyphyllum or P. montevidense. Polystichum muricatum is a species of Mexico and Central America, the Greater Antilles, and the northern part of South America. It is characterized by large leaves, to 1 m or more long, attenuate pinnae that are deeply pin- natifid to the tip, flat pinnules, large, and brown or darker petiole scales that are often short-ciliate, and usually persistent well above the base, and especially by the rachis bearing only fibrils and very narrow scales, rather than some broad scales. The species is commonly indusiate, although Smith (1985) indicates that in Venezuela it is both in- dusiate and exindusiate. XIV. Olfersia Contributed by Robbin C. Moran Olfersia Raddi, Opusc. Sci. 3: 283, /. lib. 1819. TYPE: Olfersia corcovadensis Raddi = O. cer- vina (L.) Kunze. Figure 14. Plants terrestrial or low-climbing. Stem short- creeping, in cross-section with the meristeles ar- ranged irregularly and each one not surrounded by a dark sclerenchymatous sheath. Sterile and fertile leaves strongly dimorphic. Sterile lamina pinnate with a conform terminal pinna. Veins forking near or at the base, parallel, ca. 1 mm apart, connected by a submarginal vein (this some- times difficult to see in dried material). Fertile leaves 2-pinnate, rarely 1 -pinnate. Son exindu- siate, linear to oblong. Spores monolete, echinu- late, ca. 50 microns long. Olfersia is a monotypic genus that occurs from southern Mexico to southeastern Brazil and in the Antilles. It has been included in Polybotrya be- cause of its strongly dimorphic leaves and creep- ing, scaly stem; however, Olfersia differs in stem anatomy, venation, and shape of the leaf apex. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 55 FIG. 14. Olfersia cervina: a, stem; b, apex of sterile leaf; c, portion of fertile leaf; d, portion of pinna, abaxial side, (a from Pabst 4771, Brazil, F, b, c from H. H. Smith 1075, Colombia, F; d from Madison et al, 4616, Ecuador, F.) 56 FIELDIANA: BOTANY Reference XV. Polybotrya MORAN, R. C. 1986. The neotropical fern genus Olfersia. Amer. Fern. J., 76: 161-178. 1 . Olfersia cervina (L.) Kunze, Flora 7: 3 1 2. 1 824. Figure 14. Osmunda cervina L., Sp. pi. 2: 1065. 1753. LECTO- TYPE (designated by Proctor, Flora Lesser An- tilles 223. 1977): Plumier, Traite foug. Amer. t. 154. 1705, illustrating a plant from Martinique. Olfersia corcovadensis Raddi, Opusc. Sci. 3: 283, t. lib. 1819. TYPE: Brazil, Rio de Janeiro, Mt. Corcovado, Raddi (holotype, presumably n). Polybotrya cervina (L.) Kaulf, Enum. fil. 55. 1824. Stem scales to 2.5 cm long and 0.1 cm broad, linear, golden-brown, entire. Sterile leaves 0.5-1 .2 m long and 0.3-0.5 m broad, subcoriaceous, gla- brous. Petiole shorter than the lamina, scaly at base. Pinnae 15-30 cm long and 3-8 cm broad, ovate-lanceolate to lanceolate, 4-12 pairs, short- stalked, entire, the basiscopic side excavate. Ra- chis and costae not or only faintly grooved, gla- brous. Fertile leaves produced freely on terrestrial as well as scandent stems, more erect than the sterile leaves and taller due to the longer petiole, soon withering after the spores are shed. Petiole 40-85 cm long. Pinnae 7-20 cm long, subdistant. Pinnules 0.5-1 cm long and 0.1-0.3 cm broad, pectinately arranged, broadly adnate and joined by a narrow wing of tissue. Shaded forests, ravines, swamps, mostly terres- trial or on dead logs, only occasionally climbing, 0-2000 m, Amazonas, Huanuco, Cuzco, Madre de Dios, Puno. Entirely neotropical, West Indies; southern Mexico south to Bolivia and southeastern Brazil. Amazonas: Prov. Bagua, 43 km by road NE of Chi- riaco, Barbour 4514 (MO). Huanuco: Ridge E of Tingo Maria, Allard 22301 (GH, us). Cuzco: Prov. La Conven- tion, Dudley 10324 (GH). Madre de Dios: Pantiacolla, serrania across Rio Alto Madre de Dios from Shintuya, Gentry et al. 27366 (MO). Puno: Prov. Carabaya, Vargas 18936 (GH). Contributed by Robbin C. Moran Polybotrya Willd., Sp. pi., ed. 4, 5: 99. 1810. TYPE: Polybotrya osmundacea Willd. Figure 15. Soromanes Fee, Mem. foug. 2: 16. 1845. TYPE: So- romanes serratifolium Fee = Polybotrya serrati- folia (Fee) Klotzsch. Plants hemiepiphytic or (in 2 species) terrestrial. Stem 1-3 cm thick, densely scaly, long-creeping in the hemiepiphytic species, short-creeping in the terrestrial ones, in cross-section with 4-10 circu- larly arranged meristeles, each surrounded by a dark sclerenchymatous sheath. Sterile and fertile leaves strongly dimorphic. Sterile lamina 1— 4-pin- nate, the apex pinnatifid or (in P. polybotryoides) subconform. Veins free or (in 4 of the 1 -pinnate species) anastomosing. Indusia absent. Sporangial capsules glabrous or (in P. pubens) setose. Spores monolete, echinate. Polybotrya contains 35 species and is entirely neotropical, occurring from Mexico to southeast- ern Brazil and the West Indies. Its center of di- versity is in the Andes, where 23 species occur, 12 of them in Peru, with four more expected there. The genus is divided into three subgenera, all of them in Peru. Subgenus Sorbifolia Moran (1987a), type, P. sorbifolia Kuhn, is represented in Peru by species 1. P. fractiserialis and 2. P. crassirhizoma. Subgenus Soromanes (Fee) Moran (1987a), type, P. serratifolia (Fee) Klotzsch, is rep- resented in Peru by 3. P. polybotryoides, 4. P. sub- erecta, and 5. P. andina. The other species, 6-16, belong to subgenus Polybotrya. The genus is readily identified by its strong ster- ile-fertile leaf dimorphy, unique stem cross-sec- tion, and (in most species) hemiepiphytic habit. References MORAN, R. C. 1987a. Monograph of the Neo- tropical fern genus Polybotrya (Dryopterida- ceae). Bull. Illinois Nat. Hist. Surv., 34: 1-138. MORAN, R. C. 1987b. Sterile-fertile leaf dimor- phy and evolution of soral types in Polybotrya (Dryopteridaceae). Syst. Bot., 12: 617-628. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 57 58 FIELDIANA: BOTANY Key to Species of Polybotrya a. Sterile lamina 1 -pinnate b b. Veins of sterile pinnae free (subgenus Sorbifolid) b c. Plants terrestrial; stem scales brown, concolorous; fertile pinnules commonly round or oblong, usually less than 1 cm long 1 . P. fractiserialis c. Plants hemiepiphytic; stem scales orange or reddish brown, with a dark central stripe and lighter borders; fertile pinnules usually more than 1 cm long 2. P. crassirhizoma b. Veins anastomosing (subgenus Soromanes) d d. Lamina apex a terminal segment similar to the lateral pinnae 3. P. polybotryoides d. Lamina apex gradually reduced, evenly pinnatifid, unlike the lateral pinnae e e. Sterile lamina glabrous or rarely pubescent abaxially, the trichomes 0.1-0.3 mm long, colorless, erect, 1-3-celled; pinnae 4-5 cm broad, 6-13 pairs 4. P. suberecta e. Sterile lamina densely pubescent abaxially, the trichomes 0.5-1.2 mm long, tawny, spread- ing, 5-12-celled; pinnae 5.7-10 cm broad, 4-7 pairs 5. P. andina a. Sterile lamina 1 -pinnate-pinnatifid to 4-pinnate (subgenus Polybotrya) f f. Pinnules of medial pinnae anadromous, i.e., the acroscopic pinnule is closest to the rachis . . . .g g. Stem scales yellowish to dark golden brown, long, tortuous, and intricate; the basal basiscopic veinlet often arising between the main pinnate groups of veins 6. P. altescandens g. Stem scales dark brown, rather straight, spreading or ascending; the basal basiscopic veinlet not arising between the main pinnate groups of veins 7. P. osmundacea f. Pinnules of medial pinnae catadromous, i.e., the basiscopic pinnule is closest to the rachis . . h h. Axes pilose, the trichomes 1-2.5 mm long, acicular i i. Stem scales usually denticulate, translucent, cream-colored to castaneous; lamina broadest at the base or nearly so, the petiole more than 15 cm long; fertile pinnules caudate, 3-15 mm broad; sporangial capsules setose 8. P. pubens i. Stem scales entire, opaque, dark brown; lamina base tapered to a short (less than 4 cm long) petiole; fertile pinnules 1-3 mm broad, botryoid; sporangial capsules glabrous 9. P. glandulosa h. Axes glabrous or, if pubescent, the trichomes less than 1 mm long j j. Tertiary segments or lobes of the sterile lamina 0.5-1.5 mm wide and with one vein .... 10. P. lechleriana j. Tertiary segments or lobes of the sterile lamina more than 2 mm broad and with more than one vein k k. Stem scales yellowish or dark golden brown, membranous, translucent, often long, tor- tuous and intricate, the margins erose to denticulate 1 1 . Lamina pubescent on both surfaces; veins prominulous adaxially 1 1 . P. puberulenta 1 . Lamina glabrous on both surfaces; veins not prominulous m m. Adaxial groove of costa pubescent within; basal basiscopic veinlet often arising between the main pinnate groups of veins; pinnae pinnatifid in the distal portion 6. P. altescandens m. Adaxial groove of costa glabrous within; basal basiscopic veinlet arising from the costule; pinnae divided nearly to the apex 12. P. aequatoriana k. Stem scales dull brown, thick, opaque, spreading to curved and ascending, the margins entire or nearly so n n. Sterile lamina mostly 2-pinnate-pinnatifid; tertiary segments or lobes entire or nearly so o o. Leaf margins sparsely and minutely ciliate; plants usually growing at 0-600 m . 13. P. caudata FIG. 15. Polybotrya fractiserialis: a, habit, fertile and sterile leaf. Polybotrya osmundacea: b, sterile pinna. P. polybotryoides: c, portion of pinna, abaxial side, (a from Irwin et al. 54784, Surinam, F, b from Schunke V. 5785, F, c from Moran 3593, Ecuador, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 59 o. Leaf margins glabrous; plants growing at 1000-2100 m 14. P. appressa Sterile lamina mostly 3-pinnate-pinnatifid; tertiary segments lobed p p. Basal tertiary segments less than 2.5(-3) times longer than broad 15. P. alfredii p. Basal tertiary segments 3-6 times longer than broad q q. Costa evenly puberulent abaxially; groove of the costules on the adaxial side truncated by the ridges of the costal groove and therefore not decurrent, glabrous or nearly so within 1 6. P. hickeyi q. Costa glabrous or with only scattered trichomes abaxially; groove of the costules on the adaxial side decurrent into the groove of the costa, filled with reddish to brownish trichomes . . 7. P. osmundacea 1 . Polybotrya fractiserialis (Baker) John Sm., Hist, fil. 133. 1875. Figure 15a. Acrostichum fractiseriale Baker, Syn. fil. 414. 1868. TYPE: Peru, San Martin, Mt. Campana, Spruce 4337 (holotype, K; color slide, MO of K; photos, GH, US Of K.). Acrostichum plumbicaule Baker, Syn. fil. 413. 1868. TYPE: Peru, San Martin, Tarapoto, Spruce 4090 (holotype, K; photo, GH; photo and frag., P!, us!). Polybotrya plumbicaulis (Baker) John Sm., Hist. fil. 133. 1875. Plants terrestrial. Stem scales dull brown, opaque, the margins entire or rarely denticulate. Sterile lamina 1 -pinnate, glabrous. Pinnae (11-) 15-25(-28) cm long and (2.1-)2.5-4(-4.5) cm broad. Axes glabrous, or rarely with fine pubes- cence, the trichomes 0.1-0.2 mm long, adaxial grooves shallow, glabrous. Veins free. Fertile lam- ina 2-pinnate. Pinnules 1-3 mm broad, round, oblong or linear, pectinately arranged. Wooded talus slopes, usually along rocky streams, 200-1 500 m, Amazonas, San Martin and Loreto to Cuzco and Madre de Dios. The range of P. fractiserialis consists of two widely separated regions, the eastern slopes of the Andes from Ecuador and Bolivia, and the Guian- as. Amazonas: Prov. Bagua, Valley of Rio Maranon above Cascadas de Mayasi near Campamento Sta. Montenegro (Kms. 280-284 of Maranon road), Wurdack 1854 (us). San Martin: Prov. Lamas, between Tarapoto and Moy- obamba, ca. 10 km NW ofTabalosas, Croat 51154 (MO). Loreto: Above Pongo de Manseriche, Mexia 6246a (uc, us). Aguaytia, Croat 21014 (MO). Huanuco: Prov. Huan- uco, Tingo Maria, Tryon & Tryon 5221 (GH). Previsto, antes de Aguaytia, Aguilar 947 (USM). Pasco: Iscozacin, Foster 7937 (F, USM). Prov. Oxapampa, Palcazu, Rio Alto Iscozacin, Foster 9996 (F). Junin: Prov. Satipo, Pi- chanaki, Leon 226 (USM). Chanchamayo Valley, C. Schunke 165 (F, us). Cuzco: Prov. Paucartambo, Vargas 11280 (GH). Madre de Dios: Prov. Manu, town of Ata- laya, 2-3 km W of village, Foster 7455 (F, MO, USM). Pantiacolla, serrania across Rio Alto Madre de Dios from Shintuya, Gentry et al. 27365 (MO). Valle de Marcapata, Herrera 1200 (us). 2. Polybotrya crassirhizoma Lellinger, Amer. Fern J. 62: 49, / 1, 8. 1972. TYPE: Peru, Loreto, Gamitanacocha, Rio Mazan, /. Schunke 268 (holotype, us!; isotypes, F!, GH!, NY!, uc!). Polybotrya macbridei Lellinger, Amer. Fern J. 62: 5 1, / 2, 9. 1972. TYPE: Peru, Junin, Schunke Hacienda, La Merced, Macbride 5602 (holotype, us!; isotype, F!). Plants hemiepiphytic. Stem scales dull orange or reddish brown, rarely shiny and dark brown, with a dark central stripe and lighter borders, the margins denticulate to erose. Sterile lamina 1 -pin- nate, lanceolate, glabrous. Pinnae ( 1 1-) 1 5-20(-24) cm long and 2-4 cm broad, the margins crenate to dentate. Axes sparsely scaly and pubescent, the trichomes ca. 0.1 mm long, colorless, subulate, adaxial grooves glabrous within. Veins free, often with a single basiscopic veinlet arising from the costa between the main 1 -pinnate groups. Fertile lamina 2-pinnate, occasionally 2-pinnate-pinnati- fid in large leaves. Pinnues 1-3 mm broad, oblong to linear, pectinately arranged. Wet forests, 100-1500 m, Loreto, Huanuco, Pasco, and Junin. Colombia to Bolivia and western Brazil; most frequent and abundant in Amazonia, but also on the eastern slopes of the Andes. Loreto: Prov. Maynas, Quebrada Yanomona, Explo- rama tourist camp, Rio Amazonas above the mouth of 60 FIELDIANA: BOTANY Rio Napo, Moran 3640, 3641 (F, Q, QCA). Huanuco: Prov. Pachitea, Rio Pozuzo, Foster 9284 (F, MO, USM). Pasco: Prov. Paucartambo, Cosnipata Valley, Rio Tono, road N of Patria to first foothill ridge, Foster 10627 (F). Junin: Chanchamayo Valley, C. Schunke 157, 158 (F). Schunke Hacienda, above San Ramon, Killip & Smith 24605 (NY, us). 3. Polybotrya polybotryoides (Baker) Christ, Bull. Herb. Boissier, ser. 2, 1: 70. 1901. Figure 15c. Acrostichum polybotryoides Baker, J. Bot. 19: 207. 1881. TYPE: Colombia, Norte de Santander, Ocana, Kalbreyer 1254 (holotype, K; color slide, MO of K; photos, GH, MO of K). Acrostichum juglandifolium Baker, J. Bot. 19: 207. 1881, nom. illeg., not Kaulfuss, 1 824. TYPE: Co- lombia, Antioquia, Kalbreyer 1798 (holotype, K; color slide, MO of K.; photos, GH, MO of K). Polybotrya juglandifolia Christ, Bull. Herb. Boissier, ser. 2, 4: 965. 1905. nom. nov. tor Acrostichum juglandifolium Baker, and with the same type. Polybotrya kalbreyeri C. Chr., Index fil. 504. 1906, nom. super/1, for Polybotrya juglandifolia Christ. Plants hemiepiphytic. Stem scales lustrous, golden yellow when fresh, turning yellow-brown to purple-brown upon drying, ascending with spreading tips, the margins denticulate. Sterile lamina 1 -pinnate, the tissue between the veins gla- brous on both surfaces. Pinnae 1 2-20(-30) cm long and 3-6(-8) cm broad, 5-12 pairs, the margins entire, crenate or broadly serrate. Axes glabrous or nearly so, adaxial grooves glabrous within. Veins anastomosing, the tips connected by a faint intra- marginal connecting strand. Fertile leaves 2-pin- nate. Pinnules linear, entire or lobed toward the base. Wet forests, 0-1400(-1850)m, Huanuco, Pasco. Southern Mexico to Central Peru. Huanuco: SW slope of Rio Llullapichis watershed, as- cent of Cerros del Sira, Dudley 13290D (GH). Pasco: Prov. Oxapampa, Abra los Mellizos, 4-8 km from Enen- as, Skog et al. 5036 (us). Plants hemiepiphytic. Stem scales dark brown, linear, stiff and ascending to spreading, denticu- late. Sterile lamina 1 -pinnate, glabrous or rarely with uncinate trichomes abaxially. Pinnae (10-)1 5- 21(-25) cm long and (3-)4-5(-6.5) cm broad, 6- 13 pairs, the margins entire or shallowly serrate. Axes glabrous or rarely pubescent, the trichomes less than 0. 1 mm long, uncinate, the adaxial grooves glabrous within. Veins anastomosing, the tips oc- casionally uniting to form a discontinuous sub- marginal strand. Fertile leaves 2-pinnate to 2-pin- nate-pinnatifid. Pinnules linear, entire, or lobed at the base, pectinately arranged. Wet forests, 600-1800 m, Junin. Colombia to Peru. Junin: Prov. Tarma, Chanchamayo, Esposto 10928 (USM). 5. Polybotrya andina C. Chr., Index fil. 504. 1 906, nom. nov. for Acrostichum insigne Baker, not Fee, 1872-73, and with the same type. Acrostichum insigne Baker, J. Bot. 15: 167. 1877, nom. illeg., not Fee, 1872-1873. TYPE: Ecuador, Pi- chincha, "Andes of Quito," Sodiro (holotype, K; photo, GH, us of K; isotypes, AAU!, GH!, Q!, uc!). Plants hemiepiphytic. Stem scales dark casta- neous, linear, shiny or dull, opaque, denticulate. Sterile lamina 1 -pinnate, the apex pinnatifid, pu- bescent abaxially, the trichomes spreading, tawny, uncinate, 4-10-celled. Pinnae 17-33 cm long and 5.7-10 cm broad, 4-7 pairs, the margins entire. Axes pubescent, the adaxial grooves glabrous. Veins anastomosing. Fertile lamina 2-pinnate. Pinnules linear, pectinately arranged, entire to lob- ulate. Wet forests, 1 000-1 200 m, not known from Peru but expected to occur there. Ecuador. 4. Polybotrya suberecta (Baker) C. Chr., Index fil. 506. 1906. Acrostichum suberectum Baker, J. Bot. 19:207. 1881. TYPE: Colombia, Antioquia, Kalbreyer 1877 (holotype, K; color slide, MO of K; photos, GH, us ofK). Acrostichum hackelianum Sodiro, Crypt Vase. Quit. 491. 1893. TYPE: Ecuador, Pichincha, banks of the Rio Pilaton, Sodiro (holotype, Q!)- 6. Polybotrya altescandens C. Chr., Index fil. 7. 1905, nom. nov. tor Acrostichum chrysolepis Sodiro, not Fee, 1 869, and with the same type. Acrostichum chrysolepis Sodiro, Crypt vase. Quit. 485. 1893, nom. illeg., (not Fee, 1869). TYPE: Ecua- dor, Pichincha, "ad Pilaton-Toachi," Sodiro (ho- lotype, P!). TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 61 Plants hemiepiphytic. Stem scales yellow to dark golden brown, linear, mostly concolorous, shiny, long, tortuous and intricate, denticulate, the base cordate and darkened around point of attachment. Sterile lamina to 2-pinnate-pinnatifid or rarely 3-pinnate at the base, lanceolate, glabrous. Pinnae 14_25(-35) cm long and 3-10(-17) cm broad, not deeply cut, pinnatifid above the base in the distal portions. Pinnules of the medial pinnae 6-10 cm long and 1-3.5 cm broad, catadromous, sessile. Axes pubescent and sparsely scaly abaxially, the trichomes 0.1-0.2 mm long, erect, the scales scat- tered, denticulate, adaxial grooves slightly pubes- cent within. Veins free, with the basal basiscopic veinlet often arising between the main pinnate groups of veins. Fertile lamina 3-pinnate. Pinnules narrowly oblong to linear, pinnate, with the sori obovate or oblong. Wet forests, (800-) 1200-2500 m, Loreto, Hua- nuco. Colombia; Ecuador; Peru. Loreto: Prov. Coronel Portillo, Dist. Padre Abad, divi- soria cerca al Rio Chino, /. Schunke 10200 (MO). Hua- nuco: La Divisoria, ca. 25 km NE of Tingo Maria, Moran 3688 (F, G, GH, MO, USM). 7. Polybotrya osmundacea Willd., Sp. pi. ed. 4, 5: 99. 1810. TYPE: Venezuela, Monagas, Cari- pe, Humboldt 459a (fertile), 459b (sterile) (ho- lotype, B, Herb. Willd. 19507-1, 19507-2; photos, F, GH). Figure 15b. Acrostichum osmundaceum (Willd.) Hooker, Sp. fil. 5: 246. 1864. Plants hemiepiphytic. Stem scales variable, with plants from Amazonia tending to have thick, opaque, dark brown, subentire, somewhat squar- rose scales and plants from elsewhere tending to have shiny, ascending, spreading scales, often with a dark central stripe and lighter borders, margins erose to denticulate. Sterile lamina 2-pinnate-pin- natifid to 3-pinnate-pinnatifid, deltate, ovate, or lanceolate, the tissue between the veins glabrous or rarely glandular, the glands round, sessile, res- inous, the margins glabrous. Pinnae 20-50 cm long and 10-25 cm broad, anadromous, deeply cut to the apex. Pinnules of the medial pinnae to 1 4 cm long and 4 cm broad, anadromous, or sometimes in plants from Amazonia, catadromous, the base prolonged acroscopically, stalked, the stalk 1-5 mm long. Tertiary segments lobed. Axes glabrous or pubescent abaxially, the trichomes 0.2-0.5 mm long, whitish, the scales absent or few and incon- spicuous, adaxial grooves pubescent within. Veins free. Fertile lamina to 3-pinnate-pinnatifid. Pin- nules lanceolate, pinnate. Wet forests, 0-2100 m, San Martin and Loreto to Cuzco and Madre de Dios. One of the most common and widespread spe- cies of Polybotrya, from Guatemala to French Gui- ana, Bolivia and northern Brazil; Antilles. San Martin: Camino a Pushurumbo, 7-8 km E del puente de Palo Blanco, Mariscal Caceres, Tocache Nue- vo, J. Schunke 5785 (COL, NY, us). Loreto: Prov. Maynas, Peter Jensen's Explorama Lodge, 50 mi downriver from Iquitos, Moran 3642 (AMAZ, F, USM). Huanuco: Tingo Maria, Allard 21609, 21997 (us). Pasco: Pichis Trail, Yapas, Killip & Smith 25452 (NY, us). Prov. Oxapampa, Palcazu, Rio Alto Iscozacin, Ozuz, Foster 9994 (F). Ju- nin: E of Quimiri bridge, near La Merced, Killip & Smith 23991 (F, NY, us). Ucayali: Rio Aguaytia above mouth of Quebrada Yurac-Yacu, Croat 20851 (MO). Cuzco: Prov. Paucartambo, arriba de Tono, cerca de Patria, Fernan- dez et al. 1013 (F). Madre de Dios: Prov. Manu, Vargas 17743 (GH). 8. Polybotrya pubens Mart., Icon. pi. crypt. 87, /. 25. 1834. LECTOTYPE (designated here): Brazil, ad flum. Amazonas prope Serpa, Mar- tius (holotype, M!). Polybotrya pubens Kunze, Linnaea 9: 22. 1 834, not Martius, 1834. TYPE: Peru, Loreto, Prov. May- nas, ad Yurimaguas, Poeppig (holotype, P!, one of the syntypes of P. pubens Mart.). Polybotrya decorata Lellinger, Amer. Fern J. 62: 54, / 5, 6, 11. 1972. TYPE: Peru, Loreto, Iquitos, Killip & Smith 26955 (holotype, us!; isotype, NY!). Plants hemiepiphytic. Stem scales variable, on large mature stems whitish to coffee-brown, con- colorous, ovate, membranous, denticulate, or on young or terrestrial stems narrowly lanceolate, cas- taneous. Sterile lamina mostly 1 -pinnate-pin- natifid, the base occasionally 2-pinnate-pinnatifid, pubescent throughout or rarely glabrous adaxially, the trichomes 1-2 mm long, acicular, whitish. Pin- nae 9-19 cm long and 2.5-3.5 cm broad, often with the lowest pinnae having a few free proximal segments or pinnules. Pinnules or segments catad- romous, oblong, slightly falcate. Veins free. Axes conspicuously pubescent, adaxial grooves pubes- cent. Fertile lamina mostly 1 -pinnate-pinnatifid, sometimes 2-pinnate at base. Pinnae mostly en- tire, long-caudate. Sporangial capsules setose. 62 FIELDIANA: BOTANY Wet forests, primarily in Amazonia, 0-1400 m, Amazonas, San Martin, Loreto, Pasco, Junin, Ma- dre de Dios. Colombia to Bolivia; western Brazil. This species is characterized by its relatively uncut lamina, long, dense, whitish pubescence, and long-caudate fertile pinnae. In addition, it is the only species of Polybotrya with setose sporangial capsules. The name Polybotrya pubens was pub- lished twice in the same year. The name of Martius is accepted as correct, since Kunze took his name from a specimen annotated by Martius. Amazonas: Ridge crest of Quebrada Chuivi (above Km 278 of Maranon road), valley of Rio Maranon, near Cascadas de Mayasi, Wurdack 1933 (us). San Martin: Monte Campana, Spruce 4740 (photo, GH, us). Camino a Pushurumbo, 7-8 km E del Puente de Palo Blanco, M ari seal Caceres, Tocache Nuevo, J. Schunke 5789 (NY, us). Loreto: Above Tamishuyacu, Croat 19771 (AMAZ, MO). Pasco: Prov. Oxapampa, Iscozacin, Foster 7858 (F, USM). Junin: E of Quimiri bridge, near La Merced, Killip & Smith 23196 (NY, us). Puerto Bermudez, Killip & Smith 26543 (NY, us). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, 10-15 km NNW of Shintuya, Foster 10821 (F). 9. Polybotrya glandulosa Kuhn, Linnaea 36: 65. 1869. TYPE: Brazil, Amazonas, San Gabriel, Spruce (holotype, B!). Polybotrya subelliptica Lellinger, Amer. Fern J. 62: 56. 1972. TYPE: Peru, Loreto, Mishuyacu, near Iquitos, Klug 1390 (holotype; us!; isotypes, F!, NY!). Plants hemiepiphytic. Stem scales dull brown, thick, opaque, concolorous to bicolorous, curved- appressed, the margins entire, the base expanded, the dorsal surface sometimes with a medial groove. Sterile lamina 2-pinnate-pinnatifid, subelliptic, gradually tapered to a short, less than 4 cm long, petiole, pilose on both surfaces. Pinnae 14-16 cm long and 3-3.5(-4) cm broad, sessile or nearly so. Pinnules of the medial pinnae 15-20 cm long and 5-7 cm broad, catadromous, crenate or lobed. Axes pilose, the trichomes 1-2.5 mm long, whitish, acic- ular, adaxial grooves pilose. Veins free. Fertile lamina 2-pinnate-pinnatifid, botryoid. Wet lowland forests, 1 00 m, Loreto. Southern Venezuela; northeastern Peru; north- western Brazil. This species is known from only three collec- tions (in Peru only from the type of P. subelliptica) and is apparently endemic to the northeastern Amazon basin. It is easily recognized by its lam- ina, which gradually tapers to a short, less than 4 cm long, petiole, and the long-pilose pubescence. 10. Polybotrya lechleriana Mett., Fil. lechl. 1:1: 4. /. /,/ 7-5. 1854. TYPE: Peru, Puno, San Gaban, Lechler 2156 (holotype, LZ destroyed; isotypes, B!, K, L!; frag., F!, us!; photo, GH of K). Acrostichum lechlerianum (Mett.) Hooker, Sp. fil. 5: 246. 1864, nom. illeg. (not Mettenius, 1856). Plants hemiepiphytic. Stem scales lanceolate, cream to dull brown, thin, appressed, the margins denticulate to entire. Sterile lamina 3-4 pinnate, lanceolate to elliptic, pubescent on both surfaces and especially so along the veins. Pinnae (7-) 1 0- 15 cm long and (2-)3-7(-12) cm broad, deeply and finely cut to the apex. Pinnules of the medial pinnae 1.5-5 cm long and 1-2 cm broad, sessile or nearly so. Ultimate segments 0.5-1 mm broad, single- veined, falcate. Axes pubescent, the junc- tures often provided with a brown, ovate scale, adaxial grooves pubescent or glabrous. Fertile lamina 3-4-pinnate. Pinnules oblong, botryoid. Wet forests, (1 00-) 1000- 1500 m, San Martin, Junin, Cuzco, Puno. Colombia to Bolivia; Guyana (Mt. Roraima). This species has the most finely divided lamina of any in Polybotrya. The narrow, 1 -nerved ulti- mate segments readily distinguish the species. San Martin: Mt. Guayrapurima, near Tarapoto, Spruce 4744 (L, P, frag., us). Junin: Prov. Satipo, Mapiri, ca. 12 km SW of Chequitavo, Smith 61713 (MO, USM). Cuzco: Prov. La Convention, Cordillera Vilcabamba, Camp 2.5, Dudley 10325 (GH). Puno: La Pampa, Rio Tavara, Wat- kins (us). 11. Polybotrya puberulenta R. C. Moran, Bull. Illinois Nat. Hist. Surv. 34: 90, / 41. 1987. TYPE: Ecuador, Napo, Cordillera Oriental, camino Baeza-Tena, 5 km S de Cosanga, Mo- ran 3528 (holotype, F!; isotypes, COL!, GH!, LPB!, MO!, NY!, Q!, OCA!, uc!, us!, YEN!). Plants hemiepiphytic. Stem scales yellow to sor- did orange, darker in the center with light yellow borders, membranous, spreading, erose to dentic- ulate. Sterile lamina to 2-pinnate-pinnatisect, del- tate to broadly ovate, puberulent on both surfaces, TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 63 the trichomes ca. 0. 1 mm long, spreading. Pinnae to 34 cm long and 1 8 cm broad, deeply cut to the apex. Pinnules of the medial pinnae to 10 cm long and 3 cm broad, catadromous, the basiscopic mar- gin thickened, decurrent on the costa. Axes pu- berulent, adaxial grooves pubescent within. Veins free, prominulous adaxially. Fertile leaves 4-pin- nate. Pinnules lanceolate, pinnate, botryoid. Wet forests, 1 100-1 500 m, not known from Peru but expected to occur there. Ecuador; Bolivia. 12. Polybotrya aequatoriana R. C. Moran, Bull. Illinois Nat. Hist. Surv. 34: 94, / 43. 1987. TYPE: Ecuador, Napo, Cordillera Oriental, camino Baeza-Tena, 34 km S de Baeza, Mo- ran 3512 (holotype; F!; isotypes, AAU!, COL!, GH!, LPB!, MO!, NY!, Q!, QCA!, uc!). Plants hemiepiphytic. Stem scales yellowish or- ange, membranous, long, tortuous and intricate, the margins minutely denticulate. Sterile lamina 2-pinnate-pinnatifid, lanceolate, moderately to slightly pubescent below and often glandular, the trichomes 0.1-0.3 mm long, fine, appressed, the glands resinous, round. Pinnae 25-35 cm long and 1 3-1 8(-20) cm broad, divided nearly to the apex, prolonged acroscopically, catadromous, the base stalked, the stalk 1-3 mm long, the margins slight- ly thickened and lighter colored, often revolute upon drying. Axes pubescent and scaly below, the trichomes 0. 1-0.2 mm long, erect, the scales lin- ;ar, flexuous, dark, often appressed, adaxial grooves pubescent within. Veins free, the basal basiscopic veinlet arising from the costule. Fertile leaves 3-pinnate. Pinnules linear, pinnate, botryoid. Cloud forests, 2100-2400 m, not known from Peru but expected to occur there. Ecuador; Bolivia. 13. Polybotrya caudata Kunze, Linnaea 9: 23. 1834. TYPE: Peru, Huanuco, Pampayacu, Poeppig (B!). Olfersia caudata (Kunze) Kunze, Linnaea 21: 206. 1848. Acrostichum caudatum (Kunze) Hooker, Sp. fil. 5: 244. 1864, nom. illeg., not Hooker, 1840. Plants hemiepiphytic. Stem scales dull brown, opaque, appressed, entire, the base elevated, thick- ened, curved. Sterile lamina 2-pinnate-pinnatifid (rarely 3-pinnate at the base), ovate to lanceolate, the tissue between the veins pubescent or (rarely) glabrous, the trichomes 0.2-1 .5 mm long, acicular, the margins minutely and sparsely ciliate. Pinnae up to 20-45 (-60) cm long and 7-20(-30) cm broad. Pinnules of medial pinnae 7-15 cm long and 1-3 cm broad, catadromous, the base slightly pro- longed acroscopically. Tertiary segments or lobes entire or nearly so. Axes glabrous or pubescent to various degrees, the trichomes 0.2-1.5 mm long, acicular, whitish, adaxial grooves usually pubes- cent within, often densely so at the junctures. Veins free. Fertile lamina 2-pinnate. Pinnules 0.4-1 cm broad, linear-caudate, entire or lobed at the base. Wet forests, 0-1900 m, Amazonas and Loreto to Cuzco and Puno. Eastern and southern Mexico to French Guiana, northern Brazil and Bolivia. Amazonas: Prov. Bagua, left bank of Rio Maranon opposite Quebrada Mirana (opposite Km 277 of Ma- ranon road above Cascadas de Mayasi), Wurdack 201 1 (GH, us, USM). San Martin: San Roque, LI. Williams 7681 (F, us). Loreto: Prov. Maynas, Peter Jensen's Ex- plorama Lodge, 50 mi downriver from Iquitos at Yano- mona Ck. Moran 3663 (AMAZ, USM). Huanuco: 10 km S of Tingo Maria, Stork & Morton 9509 (F, uc, us). Pasco: Puerto Bermudez, Killip & Smith 26637 (NY, us). Madre de Dios: Parque Nacional de Manu, Cocha Cashu Bio- logical Station, M. Foster P-84-42 (F). Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, 5-10 km NNW of Shintuya, Palotoa Control Post, Foster et al. 11045 (F). Cuzco: Prov. La Convention, at Camp Zero, Dudley 11513 (GH, us). Puno: San Gaban, Lechler 2321, 2329 (B). 14. Polybotrya appressa R. C. Moran, Bull. Il- linois Nat. Hist. Surv. 34: 94, / 44. 1987. TYPE: Ecuador, Napo, Cordillera Oriental, camino Baeza-Tena, 34 km al sur de Baeza, Moran 3586 (holotype, F!; isotypes, GH!, MO!, NY!, Q!, QCA!, us!). Plants hemiepiphytic. Stem scales dull brown, thick, opaque, entire, appressed or curved and as- cending. Sterile lamina mostly 2-pinnate-pinnati- fid, dark green adaxially, lighter below, both sur- faces and the margins glabrous. Pinnae to 37 cm long and 18 cm broad. Pinnules of the medial pinnae catadromous, the base prolonged acroscop- ically, stalked, the stalk 1-5 mm long. Tertiary segments entire or nearly so. Axes evenly pubes- cent abaxially, the trichomes less than 0. 1 mm long, the scales absent or few and inconspicuous, 64 FIELDIANA: BOTANY linear to lanceolate, appressed, light brown, ad- axial grooves sparsely pubescent within. Veins free. Fertile lamina unknown. Cloud forests, 1000-2250 m, Cajamarca. Ecuador and Peru. Cajamarca: Lower edge of Cutervo National Park, 10- 1 5 km N of San Andres de Cutervo, Gentry et al. 61498 (MO). 15. Polybotrya alfredii Brade, Bradea 1: 12, t. 1, f. 2. 1969. TYPE: Costa Rica, Prov. San Jose, Tablazo, Brade & Brade 98 (holotype, HB; iso- type, NY!). Plants hemiepiphytic. Stem scales dull brown, opaque to translucent, spreading, the margins den- ticulate or more, commonly entire. Sterile lamina 2-pinnate-pinnatifid to 4-pinnate, ovate, the tissue between the veins glabrous. Pinnae ( 1 5-)20-45 cm long and 10-22 cm broad, deeply cut to the apex. Pinnules of the medial pinnae up to 12 cm long and 5 cm broad, catadromous, the base stalked, the stalk 1-3 mm long. Tertiary segments lobed. Axes pubescent abaxially, the trichomes 0.1-0.2 mm long, usually tawny, adaxial grooves pubes- cent within. Veins free. Fertile leaves 3-pinnate- pinnatifid. Pinnules lanceolate, pinnate. Wet forests, 700-1900 m, Huanuco. Costa Rica to Bolivia. A thick mucilage often invests the petiole bases and distal portion of the stem. This mucilage dis- appears after drying and leaves no vestige of its former presence. Huanuco: SW slope of the Rio Lullapichis watershed on the ascent of Cerro del Sira, Dudley 13258, 18265 A (GH). 1 6. Polybotrya hickeyi R. C. Moran, Bull. Illinois Nat. Hist. Surv. 34: 88, / 40. 1987. TYPE: Bolivia, Cochabamba, Prov. Chapare, road from Cochabamba to Villa Tunari, Hickey 801 (holotype, GH!; isotype, MU). Plants hemiepiphytic. Stem scales dark choco- late-brown, mostly opaque, concolorous, or the apex with thinner, lighter borders, the margins denticulate to entire, the base thickened, attached across its full width. Sterile lamina to 3-pinnate- pinnatifid, deltate, the abaxial surface sightly pu- berulent, the trichomes up to 0. 1 mm long, cylin- drical, appressed. Pinnae to 46 cm long and 25 cm broad, deeply cut to the apex. Pinnules of the me- dial pinnae 5-10 cm long and 2-4 cm broad, ca- tadromous, the base slightly prolonged acroscop- ically, the basiscopic margins thickened and decurrent on the costa. Tertiary segments lobed. Axes puberulent and scaly, the trichomes ca. 0. 1 mm (?) long, erect, the scales appressed, denticu- late, flexuose, adaxial groove glabrous or nearly so within, not prominent. Fertile lamina 4-pinnate. Pinnules lanceolate to oblong, pinnate, botryoid. Wet forests, 1700-2400 m, Pasco. Colombia; Peru; Bolivia. Pasco: Prov. Oxapampa, Canyon de Huanca- bamba, Leon 622 (GH). Comments Polybotrya fulvastrigosa Christ, Bull. Herb. Bois- sier, ser. 2, 1: 70. 1901. TYPE: Peru, Loreto, Cerro de Canchahuaya, Huber 1448. I have not seen the type and am uncertain from the description to what species this name belongs. Polybotrya lomarioides Mett., Fil. lechl. 2: 5. 1 858. TYPE: Peru, Puno, San Gaban, Lechler. I have not seen the type and am uncertain from the description if this name applies to a species of Polybotrya or to another genus. Polybotrya nutans Kunze, Linnaea 9: 24. 1834. TYPE: Peru, "Sylvae flor. Peruv. ad Pam- payaco . . . ," Poeppig, in July 1829 (B!, P!). The fertile leaf of the type specimen is a Poly- botrya, but I do not know which species. The ster- ile leaf of the type specimen is a tree fern, perhaps a species of Trichipteris. XVI. Diplazium Diplazium Sw., J. Bot. (Schrader) 1800 (2): 61. 1 802. TYPE: Diplazium plantagineum (L.) Sw. (Hemionitis plantaginea Sm., Asplenium plantagineum L., nom. super jl. for A. plan- TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 65 FIELDIANA: BOTANY taginifolium L.) = Diplazium plantaginifoli- um (L.) Urban. Figure 16. Anisogonium Presl, Tent, pterid. 115. 1836. TYPE: Anisogonium fraxinifolium (Presl) Presl = Dipla- zium fraxinifolium Presl. Diplazium subgenus Anisogonium (Presl) C. Chr., In- dex fil. xxxi. 1906. Plants terrestrial, rarely epipetric or epiphytic. Stem commonly erect, in some species to 1 m tall, often decumbent, rarely long-creeping, sparsely to abundantly scaly, with not or scarcely clathrate scales, and usually bearing many long fibrous roots. Leaves essentially monomorphic, ca. 25 cm to 3 m long, caespitose to (occasionally) subdistant, not articulate to the stem. Lamina simple and entire to 3-pinnate-pinnatifid, glabrous to sparsely pu- bescent or somewhat scaly on abaxial axes, on some species bearing a proliferous bud distally on the rachis, axes adaxially sulcate and usually am- ply and minutely puberulent within the sulci. Ra- chis deeply sulcate adaxially, the costae less so but the edges of the sulci often thin and raised, inter- rupted and enlarged at the junction with the axis of the next order, in a few species appearing as perpendicular, herbaceous wings. Veins free, or in a few species anastomosing and the areoles with- out included veinlets. Sori elongate, elliptic to (more commonly) linear, borne on one or often both sides of the vein (diplazioid), receptacle scarcely or slightly raised, lacking paraphyses. In- dusia commonly very narrow, often delicate and hyaline, attached along the vein, sometimes shriv- eling at maturity, rarely fugacious or lacking. Spo- rangia on stalks with 2 or 3 files of cells, annular cells (12-) 15-20. Spores more or less ellipsoidal, monolete, laesura with winglike folds, the surface smooth, or papillate to echinate. Diplazium is a pantropical genus of over 300 species, and although a number of regional treat- ments have appeared in recent years, it is sorely in need of monographic work. Little has been done to delineate true relationships of species or infra- generic groups. The following key, as in previous floristic works, artificially separates many species according to degree of laminar dissection. Al- though this character frequently is helpful in key- ing out certain taxa, there is evidence to suggest it is a poor indicator of species relationships. Char- acters that more accurately note true affinities seem to be those of venation, indument, and perhaps soriation. There is ample evidence of hybridization within the genus in Peru, especially in the species with decompound leaves. A number of intermediate specimens have been examined, many with abor- tive sporangia. A relatively small group of species with areolate venation and unusual stem scales perhaps merits recognition as subgenus Anisogonium. At least one species (D. lechleri) is distinguished by otherwise free veins connected by an inframarginal vein. Certain species complexes are characterized by coarse, rigid scales on stem and axes, others have thin, flaccid ones, and still others entirely lack lam- inar scales. Minute, 1-2-celled trichomes occur on rachis and costae in varying frequency and posi- tion, but this feature is more variable and hence probably less reliable as a group indicator than are scales. Sori borne "back-to-back" on both acroscopic and basiscopic sides of a vein were termed dipla- zioid by early authors, and this has become the trademark of the genus. There are rare examples of this in Asplenium. These sori appear more com- monly in some species than others, but the fre- quency of their occurrence seems not to delineate groups as well as the color, texture, and shape of indusia. More critical study of sori and indusia would be advisable in future studies of the genus. References PROCTOR, G. R. 1985. Diplazium, pp. 392-409, in Ferns of Jamaica. Brit. Mus. (Nat. Hist.). London. SMITH, A. R. 1976. Diplazium delitescens and the Neotropical species of Asplenium sect. Hy- menasplenium. Amer. Fern. J., 66: 1 16-120. STOLZE, R. G. 1 98 1 . Diplazium, pp. 1 82-1 99, in Ferns and fern allies of Guatemala, Part II, Poly- podiaceae. Fieldiana, Bot. n. s., 6. TRYON, R. M., AND A. F. TRYON. 1982. Dipla- zium, pp. 543-554, in Ferns and allied plants, Springer- Verlag, New York. FIG. 16. Diplazium cristatum: a, habit, D. roemerianum: b, apex of lamina. Diplazium striatum: c, base of pinna segment; d, portion of rachis and pinnae bases, adaxial side, (a from Schunke V. 2733, F, b from Marling et al. 19758, Ecuador, F; c from /. Schunke 295, F, d from LI. Williams 702, F.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 67 Key to Species of Diplazium a. Veins copiously anastomosing b b. Leaves pinnatifid to pinnatisect, often pinnate at base, conspicuously petiolate 29. D. pinnatifidum b. Leaves simple, subentire, petiole short or lacking 30. D. praestans a. Veins free or, in D. lechleri, connected only at their tips by a slightly inframarginal vein c c. Leaves simple, subentire 28. D. plantaginifolium c. Leaves pinnate to decompound d d. Pinnae deeply pinnatifid to nearly 3-pinnate, or if subentire to shallowly lobed then conspic- uously inequilateral at base (cuneate or excavate basiscopically, truncate and strongly produced and often auriculate acroscopically) e e. Pinnae, at least several proximal pairs, 1-3-pinnate f f. Costae essentially glabrous abaxially, or sometimes with spreading trichomes 0.4-0.5 mm long or filiform scales, but never regularly and minutely puberulent g g. Pinnae and pinnules strongly reflexed; axes flexuous 13. D. flexuosum g. Pinnae and pinnules spreading or ascending; axes essentially straight (costae occa- sionally somewhat sinuous near apex) h h. Scales of axes, especially of costae and costules, frequent to abundant, conspicu- ously setose i i. Scales of axes bicolorous, the marginal cells and teeth castaneous to blackish, the teeth mostly bifid; sori linear, mostly 3-4 mm long; pinnae remote 9. D. remotum i. Scales of axes concolorous, the marginal teeth not bifid; sori elliptic, less than 2 mm long; pinnae approximate 7. D. moritzianum h. Scales of the axes, if any, with margins entire (rarely remotely denticulate) . . . . j j. Indusia thick, bicolorous, dark brown, but blackish near line of attachment 8. D. bicolor j. Indusia thin, essentially concolorous throughout k k. Lamina 2-pinnate, pinnules of larger pinnae crenate to lobed, cut halfway or less to the costules, free portion of segment (if any) commonly as broad as long 3. D. ambiguum k. Lamina 2-3-pinnate-pinnatisect, pinnules of larger pinnae cut % or more to the costule (or tertiary segments free and again pinnatifid), free portion of segment 1 .5-4 times as long as broad 1 1. Costules conspicuously alate adaxially, the wings often interrupted at segment bases and expanded into cristate lamellae; indusia margins erose- ciliolate 5. D. buchtienii 1 . Costules not or slightly alate, the wings (if any) not or scarcely expanded; indusia subentire (in D. hians irregularly splitting at maturity) m m. Indusia tumid (inflated); veins commonly 4 or 5 pairs per ultimate segment; free portion of ultimate segments about 1.5 times as long as broad 1 . D. hians m. Indusia flat; veins 6-8 pairs in larger segments; free portion of ulti- mate segments 2-4 times as long as broad n n. Free portion of ultimate segments twice as long as broad; costae and costules either moderately scaly or slightly pubescent with spreading trichomes about 0.4 mm. long 2. D. divergens n. Free porton of ultimate segments 3—4 times as long as broad; costae and costules completely lacking indument . 6. D. vastum f. Costae moderately to abundantly puberulent on abaxial side with minute trichomes, these 0. 1-0.2 (0.3) mm long and sometimes partly obscured by scurf o o. Tissue between the veins on abaxial surface minutely puberulent 10. D. expansum o. Tissue between the veins glabrous p 68 FIELDIANA: BOTANY p. Lamina (2-)3-pinnate-pinnatisect; sori less than 2 mm long; pinnules 1.5-4(-5) cm long 4. D. alienum p. Lamina 2-pinnate (pinnules entire to deeply lobed); sori 4-10 mm long; larger pinnules (4-)5-12 cm long q q. Pinnae with 12-16 pairs of pinnules; larger pinnules 4-6 times as long as broad, entire or crenate-serrate; indusia dark brown with a blackish base 11. D. venulosum q. Pinnae with less than 8 free pairs of pinnules; larger pinnules 3-3.5 times as long as broad, most of them deeply lobed to pinnatifid (sometimes subentire); indusia light brown 1 2. D. tungurahuae e. Pinnae subentire to pinnatisect (rarely fully pinnate at the base of basal pinnae) r r. Pinnae commonly truncate and subequilateral at base, or sometimes more strongly pro- duced basiscopically s s. Proliferous buds commonly in the axils of distal pinnae; proximal pinnae deltoid or subdeltoid, larger ones 9-22 cm broad, their ultimate segments with attenuate or acuminate apices 14. D. macrophyllum s. Proliferous buds lacking; proximal pinnae ovate, elliptic or oblong (rarely deltoid- lanceolate), larger ones 3-5.5(-7) cm broad, their ultimate segments obtuse to acute t t. Ultimate segments of proximal pinnae narrowly acute, their margins crenate to shallowly lobed; mature leaves 75 cm long and 22 cm broad 15. D. subobtusum t. Ultimate segments obtuse, occasionally subacute, their margins subentire to ser- rulate; mature leaves 1-2 m long and 30-50 cm broad u u. Indusia vestigial or lacking; sori mostly 1.5-2.5 mm long; lamina chartaceous to subcoriaceous; ultimate segments of larger pinnae more than 20 pairs .... 16. D. lindbergii u. Indusia persistent; sori mostly 3-7 mm long; lamina firm-membranaceous; ultimate segments of pinnae 15 pairs or fewer 17. D. stria turn r. Pinnae (most of them) conspicuously inequilateral at base, cuneate or excavate basiscopi- cally, truncate and much more strongly produced (often auriculate) acroscopically . . v v. Pinna base attenuate or excavate basiscopically, opposed acroscopically by a greatly elongated auricle; basiscopic pinna margin subentire, crenate, or shallowly lobed (sometimes pinnatifid on the basal pinnae) 21. D. bombonasae v. Pinna base cuneate to shallowly or deeply lobed basiscopically, opposed acroscopically by a slightly enlarged basal lobe; basiscopic margins of (at least) proximal pinnae deeply lobed to pinnatisect w w. Rachis bearing a proliferous bud on the adaxial side towards the apex; free pinnae 16-20 pairs; proximal pinnae subequilateral at base 18. D. caracasanum w. Rachis lacking proliferous buds; free pinnae 8-15 pairs; proximal pinnae (and others) conspicuously inequilateral x x. Proximal pinnae incised nearly to costa, their basal acroscopic segments free and usually short-stalked; indusium usually with a very narrow, whitish margin; scales of stem apex somewhat flaccid, dull medium-brown, 3—4 mm long . . 19. D. stuebelianum x. Proximal pinnae lobed % to % to the costa, their ultimate segments never free; indusium concolorous; scales of stem apex coarse, castaneous to blackish, often sublustrous, 1-3 mm long 20. D. cristatum d. Pinnae entire to crenate, or lobed V4 or less to costa, essentially subequilateral at base y y. Lamina gradually reduced to a pinnatifid apex z z. Pinnatifid apical portion of leaf about !/2 the length of the lamina; free pinnae 2-4 pairs 25. D. paucijugum z. Pinnatifid apical portion of leaf '/5 (or commonly less than) the length of the lamina; free pinnae on mature leaves 7-25 pairs aa TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 69 y. aa. Rachis and costae sparsely to moderately provided on abaxial side with appressed (often filiform) scales; a proliferous bud often borne distally on the rachis 24. D. celtidifolium aa. Rachis and costae lacking scales; proliferous bud lacking bb bb. Pinnae 11-17 pairs, most of them lobed about L4 to the costa, acute at apex, broadest near the center 22. D. cuneifolium bb. Pinnae commonly less than 10 pairs, with entire to shallowly crenate margins and acuminate apex, broadest at or near the base 23. D. grandifolium Lamina terminating abruptly in a conform or subconform apical segment (or rarely non- conform: with 1 or 2 basal lobes) cc cc. Veins 1-2 forked, not connected by an inframarginal vein; sori arching, the indusia thin, mostly borne on one side of a vein 26. D. roemerianum cc. Veins simple, or paired at the costa, connected at their tips by a slightly inframarginal vein; sori straight, the indusia firm, rigid, mostly double 27. D. lechleri 1. Diplazium hians Klotzsch, Linnaea 20: 361. 1847. TYPE: Venezuela, Merida, Moritz 289, in part (holotype, B, frag., us!; isotypes, BM!, K!, frags., us! of BM & K; photos, us of BM & K). Plants terrestrial, rarely on bases of tree trunks. Stem erect, this and the very base of the petiole provided with blackish, lanceolate scales to 1.5 cm long. Leaves to 2 m long, 2-pinnate-pinnatifid. Lamina chartaceous or subcoriaceous, to 80 cm broad, glabrous abaxially, the costae and costules moderately to abundantly scaly. Pinnae contigu- ous, patent or ascending, short-stalked or sessile, costae on abaxial side with scales orange to cas- taneous, filiform to lanceolate or ovate and atten- uate. Pinnules moderately to deeply lobed, cos- tules adaxially with parallel, cartilaginous ribs, abaxially scaly as on the costae. Ultimate segments 8-10 pairs, obtuse, subentire, the free portion about 1.5 times as long as broad. Veins free, commonly 4 or 5 pairs on a segment. Sori elliptic, all except basal ones less than 2 mm long. Indusia thin, gray- brown, tumid (inflated) and subentire until ma- turity, then usually irregularly splitting, with only a narrow portion persistent. In forests, 1200-3100 m, San Martin, Pasco, Junin. Venezuela; Colombia; Ecuador; Peru; Brazil. This species has been poorly understood— it is often a name under which several species have been placed. The type collection is a mixture; Mo- ritz 289 at Paris is a new species called D. mor- itzianum from Colonia Tovar, Edo. Aragua; the other isotypes are from Edo. Merida. With D. hians probably should be included D. bogotense (Kar- sten) Hieron. of Colombia, and perhaps also D. altissimum (Jenm.) C. Chr. from the Greater An- tilles. The species included as D. hians in Proctor's "Ferns of Jamaica" (1985, p. 405) is something else, for the minor axes are described as puberu- lous, with scales few or lacking. Furthermore, Field Museum specimens from the Greater Antilles that are annotated as D. hians have linear, persistent indusia (not elliptic, tumid, and evanescent). Taxa allied with this species are in great need of revision. Also see D. divergens for further discussion. San Martin: Prov. Mariscal Caceres, Rio Abiseo, Leon & Young 2164 (F, uc, USM). Pasco: Prov. Oxapampa, Dist. Oxapampa, Rio Alberto, Leon 649 (USM). Prov. Oxapampa, 4-5 km N of Mallampampa, D. Smith & Canne 5804 (MO, uc). Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8479 (MO, uc). Junin: Carpapata, above Huacapistana, Killip & Smith 24462 (GH, us). Chanchamayo Valley, C. Schunke 144, 147, 166 (F). 2. Diplazium divergens Rosenst., Repert. Spec. Nov. Regni Veg. 12: 471. 1913. TYPE: Bo- livia, Yungas, Polo-Polo, near Coroico, Buch- tien 3393 (holotype, B?; isotypes GH!, P!, us!; photos, F & us of P). Plants terrestrial. Stem erect, this and the petiole base sparsely scaly. Leaves to 2 m long, 2-pinnate- pinnatifid. Lamina firm-membranaceous, to 80 cm broad, the costae and costules moderately scaly, the scales orange, filiform to linear and attenuate, glabrous, or rarely the costae with scattered spreading trichomes about 0.4 mm long. Pinnae contiguous to subdistant, proximal ones conspic- uously petiolate. Pinnules deeply pinnatisect. Ul- 70 FIELDIANA: BOTANY timate segments 12-15 pairs, obtuse, free portion twice as long as broad. Veins free, commonly 6- 8 pairs on a segment. Sori linear, most of them 1.5-2.5 mm long. Indusia thin, gray-brown, flat, subentire. In forests, 1000-2100 m, Huanuco, Pasco, Ju- nin. Peru and Bolivia. This rare species is closely related to, and often confused with, D. hians, but the lamina is thinner in texture, ultimate segments are more numerous and relatively narrower, and the indusia are linear and flat, not elliptic and tumid. The Bryan and Smith specimens cited below are intermediate be- tween the two in most characters. They have abor- tive sporangia and are apparently hybrids. The Macbride collection seems to be a hybrid involv- ing D. divergens and a pubescent species, as it has few scales on the axes, but has some spreading trichomes on the costae abaxially, and also has many abortive sporangia. Hunuco: Muna, Bryan 550 (F, us), Macbride 4036 (F, GH, us). Junin: Chanchamayo Valley, C. Schunke 44, 708 (F). Pasco: Prov. Oxapampa, Rio El Tunqui, D. Smith et al. 1721 (F, MO). 3. Diplazium ambiguum Raddi, Opusc. Sci. 3: 292. 1819. TYPE: Brazil, Mandiocca, Raddi (ho- lotype, PI?; isotype, FI; photo, us of FI). Athyrium ambiguum (Raddi) Milde, Bot. Zeit. (Berlin) 28: 350. 1870. Diplazium ambiguum var. pubescens Rosenst., Hed- wigia 46: 108. 1906. TYPE: Brazil, Estado Santa Catharina, Blumenau, Passo Mansa, Haerchen 96 (holotype, s; possible isotype, us). Plants terrestrial. Stem erect, this and the petiole base scaly, the scales firm, dark brown to blackish, lanceolate or linear-lanceolate, the margins sub- entire. Leaves to 1.4 m long, 2-pinnate. Lamina firm-herbaceous to chartaceous, to 70 cm broad, abaxially glabrous, or the costae and costules sparsely to moderately pubescent with spreading, articulate trichomes, these mostly 0.4-0.5 mm long. Pinnae approximate, or proximal ones subdistant, patent or slightly ascending, stalked, costae sparse- ly scaly (rarely naked), the scales light brown, ovate and attenuate to filiform, their margins entire. Pin- nules sessile, often partially adnate, crenate or lobed about halfway to the costule, costules abaxially scaly as on the costae, adaxially with low, parallel, herbaceous or cartilaginous wings, these some- times interrupted and expanded near segment bas- es, but not produced into conspicuous lamellae. Ultimate segments (if any) with free portion com- monly broader than long. Veins free, 3-6 pairs to a segment. Sori linear, commonly 3-5 mm long. Indusia linear, persistent, thin, light or dark brown, flat, the margins subentire or, very rarely, erose- ciliolate. In forests, 1 50-2400 m, Amazonas and Loreto to Ayacucho. Venezuela and Colombia, south to Brazil and Paraguay. Diplazium ambiguum is a fitting name for the specimens assigned to this species. There is some question that it is correctly applied, since the type has not been examined. However, all specimens so determined from Peru match the simple orig- inal description and the excellent set of six isotype photos at us. It is distinguished from the other decompound species in that leaves are essentially 2-pinnate. The pinnules of larger pinnae are ses- sile, many of them at least partly adnate, and are crenate or dissected halfway or less to the costule. Other decompound species are at least deeply 2-pinnate-pinnatisect and either glabrous on the axes abaxially or regularly and minutely puberu- lent, with stout, rigid 1-3-celled trichomes mostly 0.1-0.2 mm long. Most specimens of D. ambig- uum are glabrous abaxially, but occasionally the costae and costules are sparsely to moderately pu- bescent with spreading, delicate, multicellular tri- chomes about 0.5 mm long. This species is most closely related to D. dipla- zioides (Klotzsch & Karsten) Alston, of northern South America, Central America, and the Lesser Antilles, but the latter differs in that the pinnules are cut nearly or quite to the costule, the sori are much shorter, the indusia are always erose-cilio- late, and the axes are never abaxially pubescent. Diplazium ambiguum further lives up to its name in the characters of indusia and indument. Mar- gins of the indusia are predominantly entire but occasionally are erose-ciliolate, as in D. dipla- zioides. Axes are most commonly glabrous abax- ially, but sometimes they are pubescent with spreading, multicellular trichomes. (On Spruce 4344 from Monte Campana, San Martin, axes of different leaves vary from glabrous to pubescent.) Further uncertainty is introduced by Rosen- stock's D. ambiguum var. pubescens, which was said to differ from the typical in the glandular- puberulent axes and fimbriate indusia. Although TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 71 the type has not been seen, a topotype, Haerchen 35a (F), does have erose-ciliolate indusia, but the axes are glabrous abaxially. Costae are puberulent adaxially, which perhaps is what Rosenstock was referring to, but this condition is quite common throughout the genus, so the character is of little importance. Obviously this species and its allies are very much in need of further study, throughout their range. Amazonas: Mouth of Rio Santiago, Tessmann 4390 (us). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Quebrada Saule Chico, /. Schunke V. 4362 (F, GH, us). Monte Campana, near Tarapoto, Spruce" 4 344" (BM, BR, K, w). Mt. Guayrapurima, Spruce "4344" (p). Loreto: Prov. Maynas, Yanamono Tourist Camp, van der Werffet al 9855, 9868 (MO). Pasco: Oxapampa, 4- 5 km N of Mallampampa, D. Smith & Canne 5801 (MO). Junin: Prov. Chanchamayo, Chilpes, D. Smith & Pa- lacios 2637 (F, MO). Ayacucho: "Aina" (Ayna), between Huanta and Rio Apurimac, Killip & Smith 22727 (us). 4. Diplazium alienum (Mett.) Hieron., Hedwigia 59: 336. 1918. Asplenium alienum Mett., Fil. lechl. 2: 1 8. 1 859. LEC- TOTYPE (designated here): Peru (Puno), San "Gavan" (Gaban), Lechler 2320 (B!; ISOLEC- TOTYPE, P!; photos, BM & F of B, GH & us of P.). PARATYPE: Lechler 2173 (B!). Asplenium fuscopubescens Hooker, Sp. fil. 3: 264. 1860. LECTOTYPE (designated here): Peru, (San Mar- tin), Monte Campana, near Tarapoto, Spruce 4759 (K!; ISOLECTOTYPES, BM!, P!; photos, GH & us of BM). Schlim 69 of Colombia, also cited by Hooker, is Diplazium gracilescens. Diplazium fuscopubescens (Hooker) Moore, Index fil. 329. 1861. Diplazium lehmannii Hieron., Bot. Jahrb. Syst. 34: 458. 1904. TYPE: Colombia, "prope Ricaurte ad rivulum Cuaiquer," Lehmann 5064 (holotype, B; frag., BM!; isotypes, F!, us!). Plants terrestrial, or occasionally growing on the base of tree trunks. Stem erect, this and the petiole base provided with a few broad, brown scales. Leaves to 2 m long, 2-3-pinnate-pinnatisect. Lam- ina firm-membranaceous, to 50 cm broad, the cos- tae, and often the costules and rachis, moderately to abundantly puberulent abaxially, the trichomes spreading, septate, 0.1-0.3 mm long. Pinnae trun- cate and subequilateral at the base. Pinnules 1 0- 20 pairs on larger pinnae, 1.5-4(-5) cm long, the costules adaxially alate, the wings perpendicular to the lamina and often expanded into crestlike lamellae. Ultimate segments 8-12 pairs, those of larger pinnae usually free, lobed to pinnatisect. Veins free. Sori less than 2 mm long, usually single, occasionally diplazioid. Indusia light or (more commonly) dark brown, subentire to lightly erose or ciliolate, persistent. In dense, wet forests and wooded valleys, 700- 2700 m, San Martin to Cuzco and Puno. Colombia; Ecuador; Peru; Bolivia. Hooker and Baker (Syn. fil. 242. 1867) treated this as conspecific with Asplenium sandwichianum (= Diplazium sandwichianum (Presl) Diels) of Ha- waii. The type of the latter has not been examined, but herbarium specimens so determined are less dissected and puberulent, and costules are scarce- ly winged. The disjunction is also unlikely. Diplazium buchtienii is possibly synonymous; see treatment of the latter for further discussion. A distinctive feature of this species and D. alienum is the conspicuous wings on the adaxial side of the costules, which are membranaceous and borne perpendicular to the lamina surface. Often these are expanded, near the bases of tertiary segments, into crestlike lamellae similar to those in certain species groups of Hymenophyllum. Huanuco: Slope of Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13252 (GH), 13253 (MO, us). Pasco: Prov. Oxapampa, Palcazu Valley, Rio San Jose, D. Smith 3949 (MO, uc). Junin: Prov. Satipo, Pichanaki, Leon 230 (USM). Cuzco: La Convencion, Valle San Mi- guel, Bues 2147 (GH, us). Prov. La Convencion, Cordil- lera Vilcabamba, Dudley 10326B (GH). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10702 (F). 5. Diplazium buchtienii Rosenst., Repert. Spec. Nov. Regni Veg. 6:312.1 909. TYPE: Bolivia, San Carlos, near Mapiri, Buchtien 1136 (ho- lotype, B?; isotype, us!). Plants terrestrial. Stem erect, this and the very base of the petiole provided with broad, firm, cas- taneous to blackish scales. Leaves 60-1 50 cm long, 2-3-pinnate-pinnatisect. Lamina firm-membra- naceous to chartaceous, to 40 cm broad, essentially glabrous abaxially, scales lacking, or rare and scat- tered. Pinnae approximate, slightly ascending, ses- sile, or proximal ones short-stalked. Pinnules 1 0- 1 6 pairs per pinna, perpendicular to the costa, the costules adaxially alate, the wings often interrupt- ed at segment bases and there expanded into crest- like lamellae. Tertiary segments 8-15 pairs, entire and adnate to free and again pinnatifid. Veins free, 4(5) pairs on a segment, or less. Sori elliptic, less 72 FIELDIANA: BOTANY than 2 mm long. Indusia light or dark brown, thin, erose-ciliolate. In wet forests, 1200-2500 m, Huanuco to Cuz- co. Peru; Bolivia. This species probably should be included with D. alienum, as it seems to differ from the latter primarily in its glabrous, rather than regularly pu- berulent, axes on the abaxial side. Although the occurrence of laminar trichomes and/or scales is presumed to be a strong diagnostic character in Diplazium, it seems to be less consistent in some species groups. Monographic study is needed. Huanuco: Prov. Huanuco, km 452, Lima-Tingo Ma- ria road, Young & Sullivan 557 (MO, uc). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachanga, van der Werff et al. 8456 (MO, uc). Junin: Chanchamayo Valley, C. Schunke 49, 143, 171 (F). Cuzco: Prov. Pau- cartambo, Valle de Pillahuata, Herrera 1612 (us). In dense forests, 500-1800 m, San Martin, Ju- nin, Cuzco. Colombia; Ecuador; Peru. Among the decompound species of the genus, this species is marked by its naked lamina, which is dark green adaxially and quite pale green abax- ially, the stramineous axes, the stalked, subdistant pinnae and pinnules, and the ascending, com- monly obtuse, ultimate segments with parallel sides. Peyton & King 1422 from Prov. La Convention in Cuzco (GH, MO) is a hybrid involving D. vastum. The lamina has the typical light green color abax- ially, and the pinnules have numerous, narrow, ascending segments, but the axes are amply pro- vided with attenuate, spreading scales, and the in- dusia are short and elliptic. The sporangia are all abortive. Junin: Prov. Tarma, Chanchamayo, Esposto (Cat. No. 11011, USM), C. Schunke 79, 447, 843 (F, us). Cuzco: Prov. Paucartambo, Hacienda Villa Carmen, Vargas 14681 (GH). 6. Diplazium vastum (Mett.) Diels, Nat. Pflanzen- fam. 1(4): 228. 1899. Asplenium vastum Mett., Ann. Sci. Nat. Bot., ser. 5, 2: 237. 1864. TYPE: Colombia, Bogota, Alto Tri- go, Lindig 349 (holotype, B, frag., us!; iso types, BM, K!, P!; photos, F, GH, us of P). Diplazium tarapotense Rosenst., Repert. Spec. Nov. Regni Veg. 7: 295. 1909. TYPE: Peru, San Mar- tin, Tarapoto, Mt. Campana, Spruce 4682 (ho- lotype, P!; isotypes, BM!, GH!, K!; frag., us! of BM; photos, GH & us of P). Plants terrestrial. Stem not seen, probably erect. Leaves huge, probably to 2 m long, 2-pinnate- pinnatifid, occasionally 3-pinnate as to base of larger pinnae. Lamina firm-membranaceous to chartaceous, to 80 cm broad, pale green and gla- brous abaxially, lacking scales, dark green adaxi- ally. Pinnae subdistant, somewhat ascending, long- stalked. Pinnules incised nearly or quite to the costule, numerous, subdistant, proximal ones short-stalked, costules adaxially with low, thick- ened wings, these often interrupted and slightly expanded near segment bases, but not produced into lamellae. Ultimate segments 10—15 pairs, nar- row-oblong, with parallel sides, commonly obtuse, ascending, entire or occasionally crenulate, larger ones 3—4 times as long as broad. Veins free, 6-8 pairs on larger segments. Sori linear, 2-3 mm long. Indusia thin, light brown, subentire, persistent. 7. Diplazium moritzianum Stolze, sp. nov. Squamae caulis nigellae, ovato-lanceolatae, margini- bus integris vel denticulatis; folium usque ad 2 m Ion- gum, 2-pinnato-pinnatisectum; lamina in pagina abax- iali glabra, axibus moderate vel abundanter paleaceis; pinnae contiguae, sessiles vel petiolulatae; costao et cos- tulae squamis fuscis vel castaneis, dentatis vel setosis; segmenta ultima 6-10-jugata, obtusa, saepe crenato-ser- rata; venae liberae, 3-5-jugatae; sori elliptic!, minus quam 2 mm longi; indusium tenue, plerumque planum, eroso- ciliatum. Plants terrestrial. Stem erect to ascending, this and the petiole base scaly, the scales coarse, black- ish, sublustrous, ovate-lanceolate, the margins en- tire to remotely denticulate. Leaves to 2 m long, 2-pinnate-pinnatisect. Lamina chartaceous, to 70 cm broad, abaxially glabrous, the axes moderately to abundantly scaly. Pinnae approximate, patent or ascending, sessile, or proximal one stalked, cos- tae with scales dark brown or castaneous, narrow, attenuate, the margins conspicuously dentate to setose. Pinnules deeply incised, often nearly to the costule, costules abaxially scaly as on the costae, adaxially with low herbaceous parallel wings, these often interrupted and expanded near segment bas- es, but not produced into conspicuous lamellae. Ultimate segments 6-10 pairs, obtuse, larger ones crenate-serrate and about twice as long as broad. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 73 Veins free, 3-5 pairs to a segment. Sori broadly elliptic, less than 2 mm long. Indusia persistent, thin, light or dark brown, flat or occasionally slightly tumid, the margins erose-ciliolate. TYPE— Venezuela, Colonia Tovar, Merit z 289, in part (holotype, P!, 2 sheets; frag., F!; photos, F & GH of P). Montane rain forests, 2600-3100 m, San Mar- tin, Huanuco, Pasco. Venezuela; Peru; southeastern Brazil (Lanna 1708 [F]). This has the same type number as D. hians, but type specimens of the latter are designated as Me- rida, "Colombia" [Venezuela], whereas the two sheets at Paris are designated as Colonia Tovar, which is in Edo. Aragua. The Paris specimens rep- resent a very different species, with scales on axes conspicuously dentate to setose, the ultimate seg- ments relatively narrower and usually crenate-ser- rate, and the indusia commonly flat, with margins erose-ciliolate. In comparison, D. hians has scales with entire margins, ultimate segments broader and subentire, and indusia strongly tumid. Diplazium moritzianum is more closely related to D. pedatum Klotzsch of Venezuela, which also has indusia with fimbriate margins. Among other characters, the latter differs from D. moritzianum in the very narrow, often subulate and entire rhi- zome and petiole scales. San Martin: Prov. Mariscal Caceres, near Mirador, Rio Abiseo National Park, Leon 2164 (F, USM). Huanuco: Carpish Divide, Sandeman 5117 (K). Pasco: Prov. Oxa- pampa, Rio San Alberto Valley, D. Smith & Pretel8033 (GH, MO). 8. Diplazium bicolor Stolze, sp. nov. Folium 2-pinnato-pinnatifidum, usque ad 1.5 m Ion- gum; lamina glabra, squamis absentibus; pinnae subdis- tantes, petiolulatae; pinnulae numerosae, subdistantes, ad basin truncatae, ad apicem attenuatae; segmenta ul- tima lata, obtusa; venae liberae, 4-5-jugatae; sori linea- res; indusia crassa, integra, bicoloria, fusca et nigella. Plants terrestrial. Stem erect to decumbent, pro- vided with stout, appressed, ovate-lanceolate scales, these blackish brown, to 1.5 cm long, with margins entire. Leaves to 1.5 m long, 2-pinnate- pinnatifid, petiole essentially glabrous and not scaly. Lamina chartaceous, to 70 cm broad, gla- brous except for sparse, minute puberulence in the sulci of axes adaxially. Pinnae subdistant, some- what ascending, stalked. Pinnules pinnatifid, nu- merous, subdistant, gradually tapering from a truncate base to a pinnatifid, attenuate apex, cos- tules adaxially with low, herbaceous wings per- pendicular to the plane of the lamina. Ultimate segments about 1 0 pairs, obtuse, the free portion often nearly as broad as long. Veins free, 4-5 pairs on a segment. Sori linear, commonly extending from the midrib nearly to the segment margin, at least the basal acroscopic ones diplazioid. Indusia thick, the margins entire, bicolorous, essentially dark brown, but blackish near the line of attach- ment, commonly persistent. TYPE— Peru, Pasco, Prov. Oxapampa, Dist. Oxapampa, Rio San Alberto, Leon 643 (holotype, F!; iso types, GH!, USM). Wet forests, 1200-2000 m, Pasco, Junin. Peru; Bolivia (Beck 3059 [F]). This is one of a few species in Peru, including D. celtidifolium, with thick, bicolorous indusia. Indusia in the genus are typically thin-textured and delicate, and may be light to dark brown, whereas in D. bicolor they are thicker, quite firm, and dark brown, becoming blackish in a line along the point of attachment to the vein. Also characteristic is the "herringbone" soral pattern. The four or five pairs of sori are straight, ascending and crowded, and nearly fill the space between midrib and seg- ment margin. Junin: Chanchamayo Valley, C. Schunke 662 "A"' (F). 9. Diplazium remotum Fee, Crypt, vase. Bresil, 1 : 8l,t.24,f. 1. 1869. TYPE: Brazil, Glaziou 2332 (holotype, P). Plants terrestrial. Stem erect, to 50 cm long, stout, subarborescent, provided with coarse, dark brown to blackish scales to 1 cm long. Leaves to 2 m long, 2-pinnate-pinnatifid. Lamina charta- ceous, to 70 cm broad, glabrous, or axes minutely and sparsely puberulent abaxially, axes also mod- erately to abundantly scaly, the scales mostly bi- colorous, the main body light brown, the margins castaneous to blackish and conspicuously dentate, the dark, elongated, teeth commonly bifid. Pinnae remote, stalked, patent to slightly ascending. Pin- nules 8-12 free pairs (on larger pinnae), deeply incised. Ultimate segments 8-12 pairs, mostly ob- tuse and entire, or the apices crenulate-serrate. Veins free. Sori mostly 3—4 mm long and single, or those on basal veins diplazioid. Indusia linear, 74 FIELDIANA: BOTANY bicolorous, brown or gray-brown, but blackish near point of attachment, firm, persistent, margins sub- entire or slightly erose. In dense, wet forests, often on stream banks, 1 700-3000 m, Amazonas to Cuzco. Venezuela; Peru; Brazil. The scales of the axes are most distinctive. They are the same as those of the putative areolate sub- genus A nisogonium, i.e., with dark brown to black- ish margins contrasting with the lighter brown of the main body of the scale, and elongated, setalike, bifid, marginal teeth. The type has not been ex- amined, and the protologue mentions nothing of these singular scales. However, Fee's illustration does indicate the abundantly paleaceous axes, and an enlarged drawing of a scale depicts the long, marginal, apparently dark, teeth. Furthermore, Dr. Badre of the Laboratoire de Phanerogamic at Paris has kindly examined the holotype and confirmed the presence of these scales. Probably conspecific is D. rostratum Fee, of Bra- zil, described in the same publication and illus- trated as Figure 2 of the same plate. The former is described as having pinnule apices entire, the segment apices rostrate, and the rachis bicanalicu- late, whereas in D. remotum the pinnule apices are supposed to be serrate, the segment apices dentate, and the rachis simply canaliculate. Dr. Badre has stated (in litt.) that the scales of the holotype (Gla- ziou 2331) are identical to those of D. remotum. Skog & Skog 5210 (us), from Cuzco, apparently is a hybrid involving D. remotum and another species. The specimen has proliferous buds scat- tered all over the lamina, dentate scales (with teeth that are not always dark or bifid), and axes often puberulent abaxially. Most sporangia are abortive. Amazonas: Prov. Bagua, 20 km E of La Peca, Barbour 2763 (F, MO, uc). San Martin: Prov. Mariscal Caceres, Rio Abiseo National Park, Leon 2160 (F). Pasco: Prov. Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8457 (MO). Cuzco: Prov. La Convention, Cordillera Vilcabamba, Dudley 10460 (GH, MO). Prov. Paucartambo, Valle de Pillahuata, Herrera 1623 (us in part). 10. Diplazium expansum Willd., Sp. pi. ed. 4, 5: 354. 1 8 1 0. TYPE: Venezuela, Caracas, Brede- meyer (holotype, B!, Herb. Willd. 19948; pho- tos, F, GH, US). Asplenium expansum (Willd.) Presl, Reliq. haenk. 1 : 46. 1825. Allantodia asplenioides Kunze, Linnaea 9: 72. 1834. TYPE: Peru, Huanuco, Cuchero, Poeppig, Aug. 1829 (holotype, w; isotypes, B!, BM; prob. isotype, MO!; photos, BM, of w; GH, p & us of BM). Diplazium asplenioides (Kunze) Presl, Tent, pterid 114. 1836. Athyrium expansum (Willd.) Milde, Bot. Zeit. 28: 353. 1870, not Moore, 1860. Diplazium bonapartii Rosenst., Repert. Spec. Nov. Regni Veg. 7: 295. 1909. TYPE: Peru (San Mar- tin), Monte "Guayrapurina" (Guayrapurima), Spruce 4683 (holotype, P!; frag., us!; photos, F, GH, us). Plants terrestrial. Stem erect, this and the petiole naked, or with a few dark brown scales. Leaves to 2 m long, 2-pinnate-pinnatifid. Lamina firm- membranaceous, to 70 cm broad, gradually re- duced to a pinnatifid apex, minutely puberulent on costae, costules, veins, and leaf tissue abaxially, trichomes of the axes reddish brown, septate, 0. 1- 0.3 mm long, those between the veins mostly uni- cellular and whitish. Pinnae truncate and subequi- lateral at base, mostly stalked. Pinnules 10 pairs or more on larger pinnae, deeply incised (some basal ones nearly to the costule), truncate and sub- equilateral at base. Ultimate segments 8-12 pairs, mostly obtuse or truncate. Veins free. Sori mostly 2 mm long or less and single, or those of basal veins diplazioid. Indusia linear, thin, dark brown, erose-ciliolate, subpersistent. In dense, wet forests, 100-2000 m, Loreto and San Martin to Cuzco. Greater Antilles; southern Mexico; Guatemala; Venezuela; Colombia to Peru; Brazil. This is distinguished from most species in the genus by the abundant minute trichomes on the tissue between the veins abaxially. Diplazium bon- apartii has numerous, rather than sparse, ap- pressed scales on the costules, but is otherwise identical, therefore conspecific. Diplazium mela- nosorum (Sodiro) C. Chr., of Ecuador and Colom- bia, is similar, but the lamina is subcoriaceous, and indusia are broadly elliptic instead of linear. San Martin: Near Tarapoto, Spruce 4124 (BM, K). Lo- reto: Prov. Maynas, 50 mi from Iquitos, Moran 3655 (MO). Huanuco: Prov. Huanuco, Rio Huallaga, above Rio Cayumba, Mexia 8316 (K, us). Junin: Prov. Tarma, Chanchamayo, Esposto (USM), C. Schunke 35, 43, 499 (F). Ucayali: Prov. Coronel Portillo, Boqueron, Ferreyra 16057 (GH, USM in part). Cuzco: Prov. Quispicanchi, Valle de Marcapata, Herrera 1595 (us). 11. Diplazium venulosum (Baker) Diels, Nat. Pflanzenfam. 1(4): 226, 228. 1899. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 75 Asplenium venulosum Baker, Syn. fil. 238. 1 867. TYPE: Ecuador, Mt. Tungurahua, Spruce 5343 (holo- type, K.!; isotypes, BM!, P!, w!; photos, F & GH of K). Plants terrestrial. Stem erect, provided with co- pious, linear, brown scales, these to 1.5 cm long, their margins remotely ciliolate or denticulate. Leaves to 3 m long, 2-pinnate, the petiole densely scaly as on the stem, but marginal teeth of scales elongated, more numerous and crowded. Lamina chartaceous, to 85 cm broad, often with a prolif- erous bud distally on the rachis, indument of the axes abaxially a mixture of minute trichomes, scurf (appressed squamules), and filiform brown, den- tate scales. Pinnae narrowly acute at apex, sub- equilateral and truncate at base, mostly stalked, patent or slightly ascending. Pinnules remote, en- tire to crenate-serrate, free ones of larger pinnae 12-16 pairs, larger ones 6-12 cm long, 4-6 times as long as broad, costules adaxially provided with low wings of tissue perpendicular to the plane of the lamina, but these not abruptly expanded or cristate. Veins free. Sori linear, 5-10 mm long. Indusia dark brown, becoming blackish at base, firm, persistent, the margins subentire. Rare, in dense, wet forests, usually along stream banks, 1700-1900 m, Amazonas and San Martin. Colombia; Ecuador; Peru. In addition to the key characters, this species also can be separated from D. tungurahuae by the dense covering of linear, attenuate, tortuous, scales on the petiole. Few species ofDiplazium have such an abundance of petiole scales. The few (if any) scales on the petiole base of D. tungurahuae are broad, rigid and appressed, with margins suben- tire. Amazonas: Prov. Bagua, Cordillera Colan SE of La Peca, Barbour 4189 (USM). San Martin: Prov. Rioja, Venceremos, D. Smith 4456 (F, MO, uc). 1 2. Diplazium tungurahuae (Sodiro) C. Chr., Ind. fil. suppl. 1: 28. 1913. Asplenium tungurahuae Sodiro, Anal. Univ. Central Quito 22: 97 (Sert. fl. Ecuad. 2: 20) 1908. TYPE: Ecuador, Tungurahua, Sodiro, in 1 904 (holotype, Q?; isotype, P!; frag., us!; photos, GH & us of P). Asplenium crassifolium Sodiro, Anal. Univ. Central Quito 22: 97 (Sert. fl. Ecuad. 2: 21) 1908. TYPE: Ecuador, Chillanes, Mt. Chimborazo, Sodiro, in 1891 (holotype, Q?; isotype, P!; photos, F, GH & us of P). Diplazium crassifolium (Sodiro) C. Chr., Ind. fil. suppl. 1: 26. 1913. Plants terrestrial. Stem erect or decumbent, pro- vided at apex with coarse, dark brown scales about 1 cm long. Leaves to 1.5 m long, 2-pinnate. Lam- ina chartaceous, to 50 cm broad, often with pro- liferous buds on rachis or costae, indument of the axes abaxially a mixture of minute, 1- or 2-celled trichomes, broad, appressed scales, and scurf (ap- pressed squamules). Pinnae subequilateral and truncate at base, mostly stalked. Pinnules suben- tire to pinnatifid, sessile to short-stalked, broadest at the base, free ones 8 pairs or fewer, larger ones 4-9 cm long, 3-3.5 times as long as broad, costules adaxially provided with low wings of tissue per- pendicular to the plane of the lamina, but these not abruptly expanded or cristate. Veins free. Sori linear, commonly 4-6 mm long. Indusia light brown, persistent, with margins subentire or light- ly erose. In dense, wet forests, often in ravines or along stream banks, 100-2300 m, Cajamarca, Amazo- nas, and Loreto to Junin. Colombia; Ecuador; Peru. This species is a coarse fern, the broad pinnae and pinnules mostly triangular from a truncate base. As in most of the puberulent group of de- compound species, the trichomes on the costae and costules abaxially are minute (in D. tungura- huae mostly 0.1 mm long and 1-2-celled). The minor axes also have few to many broad, ap- pressed scales, and the costa and rachis also often have a sparse to ample covering of appressed squa- mules, that sometimes partially obscure the tri- chomes. Not all specimens examined have prolif- erous buds, but on those that do the buds often are on the costae as well as the rachis. This may be only a variant of D. striatum, under which see further discussion. Cajamarca: Prov. Cutervo, road to San Andres Na- tional Park, Lopez & Sagastegui 5430 (GH, HUT, uc). Amazonas: Prov. Bongara, Shillac, D. Smith & Vasquez 4886 (MO, uc). Loreto: Balsapuerto, lower Rio Huallaga basin, Killip & Smith 28506 (us). Huanuco: Fundo Chela, Sinchono, Aguilar 921 (USM). Tingo Maria, Tryon & Tryon 5283 (BM, F, GH, us). Pasco: Prov. Oxapampa, Cordillera de San Matias, D. Smith 2004 (F, MO). Junin: Chancha- mayo Valley, C. Schunke 960 (F, us). 13. Diplazium flexuosum Presl, Tent, pterid. 1 14. 1836, nom. nov. for Asplenium flexuosum Presl and with the same type. 76 FIELDIANA: BOTANY Asplenium flexuosum Presl, Reliq. haenk. 1: 46, t. 7, f. 1. 1825, not Schrader, 1818. TYPE: Peru, "in vallibus Cordillerarum," Haenke (holotype, PR or PRC; isotype, K). Athyrium flexuosum (Presl) Milde, Hot. Zeit. (Berlin) 28: 353. 1870. Diplazium preslianum C. Chr., Index fil. 237. 1905, nom. super/1, for Diplazium flexusosum Presl and with the same type. Plants terrestrial, probably clambering. Stem decumbent, provided at the apex with coarse, blackish scales to 6 mm long. Leaves to 2 m long, 2-pinnate-pinnatisect. Lamina chartaceous, essen- tially glabrous, with some scattered trichomes on the rachis, but not regularly puberulent, the axes conspicuously flexuous. Pinnae strongly reflexed, proximal ones long-stalked, to 60 cm long. Pin- nules strongly reflexed, lobed to pinnatisect, in- equilateral at base, broadly cuneate acroscopically, truncate and more strongly produced basiscopi- cally. Veins free. Sori linear, 2-4 mm long, those of basal veins mostly diplazioid, the rest usually single. Indusia light brown or gray-brown, per- sistent, the margins entire to lightly erose. In forests, leaves probably clambering on shrubs, about 1 600 m, Huanuco and Ucayali. Rare and endemic. This cannot be confused with any other species of the genus in Peru. The alternate pinnae and pinnules are strongly reflexed and abruptly bent at the axils, which results in a conspicuous, flex- uous aspect to the axes. Huanuco: Pampayacu, Kanehira 181 (GH, us). Ucayali (as Loreto): Prov. Coronel Portillo, NE of the pass at La Divisoria, Skoget al. 5151 (F, us). Department unknown: Matthews 1818 (BM, K, us), Poeppig, in 1829 (K, p). 14. Diplazium macrophyllum Desv., Prodr. 280. 1827. TYPE: "Hab. in America calidiori," collector and locality not cited, but possibly Dombey or Poeppig, Peru (holotype, p; pho- tos, GH, us). Asplenium desvauxii Mett., Abh. Senckenberg. Na- turf. Ges. 3: 225, / 4, 1859, nom. nov. for Dipla- zium macrophyllum Desv., not Asplenium mac- rophyllum Mett. 1856. Asplenium procerum Sodiro, Anal. Univ. Central Qui- to 22: 96 (Sert. Fl. Ecuad. 2: 19). 1908. TYPE: Ecuador "in silvis subtropic. val. Nanegal," So- diro, 1 89 1 (holotype, Q?; isotype, P!, 3 sheets; pho- tos, F, GH). Plants terrestrial. Stem erect, provided at apex with sublustrous, brown scales, these 8-14 mm long. Leaves to 2.5 m long, 1-pinnate-pinnatisect (at least proximally), petiole sparsely scaly toward base. Lamina to nearly 1 m broad, terminating abruptly in a pinnatifid apex, essentially glabrous (rachis rarely sparsely puberulent), but sparsely provided abaxially on costae and rachis with dull, light brown scales, these filiform or attenuate, mostly flaccid and appressed, and distal pinna ax- ils commonly provided abaxially with proliferous buds. Pinnae subequilateral, proximal ones del- toid or subdeltoid, stalked, incised nearly (rarely quite) to the costa, distal ones subentire, larger pinnae 9-22 cm broad, their ultimate segments with attenuate or acuminate apices. Veins free, pinnately branched in the segments. Sori single, or often diplazioid on the basal acroscopic vein branch. Indusia commonly persistent, light to dark brown, or often blackish near their attachment to the vein. In dense, wet forests, often in ravines and along stream and river banks, 400-1800 m, Amazonas to Cuzco. Venezuela; Colombia to Bolivia. Specimens of D. macrophyllum are often deter- mined as D. costale (Sw.) Presl of Jamaica, which differs in the darker, larger, very abundant scales on the costae, the deeply serrate segment apices, and the short (ca. 5 mm) sori, which extend only halfway or less to the segment margin. There are several other taxa in this species complex, which is in need of more detailed examination: Dipla- zium appolinaris Fee (Lesser Antilles) belongs here, as well as D. oxylobum Sodiro (Ecuador). The lat- ter, with short sori, appears to be intermediate between D. macrophyllum and D. costale. A collection from San Martin, Knapp & Alcorn 7749 (F, MO), is similar to D. macrophyllum in every way except for its exceedingly dense indu- ment. Leaves of the latter are essential glabrous, and with a few scales, but in this specimen the petiole and rachis are covered with attenuate, brown scales with setulose margins, these to 2 cm long toward the stem, becoming shorter and less abundant in the distal portion of the lamina. In addition, the rachis distally and the costae and veins on both sides are beset with copious septate trichomes up to 1 mm long. A number of shorter trichomes also are borne adaxially on the tissue between the veins. Although the specimen is ro- bust (leaf 1.5 m long), it is sterile, and may be merely a monstrous form or hybrid. At any rate, TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 77 it is inappropriate here to describe a new form or variety on the basis of a single, sterile specimen. Ama/onas: Prov. Bagua, valley of Rio Maranon above Cascadas de Mayasi, Wurdack 1860 (us). San Martin: Mt. Campana, near Tarapoto, Spruce 4336 (BM, K, p). Huanuco: Prov. Huanuco, Dist. Churubamba, Mt. Santo Toribio, Mexia 8249 (BM, F, GH, MO, uc, us). Pasco: Prov. Oxapampa, Palcazu Valley, Iscozacin, Foster et al. 7881 (F, MO). Junin: Schunke Hacienda, above San Ramon, C. Schunke A- 194 (GH, us). Ucayali (as Loreto): Along Rio Aguaytia above mouth of Quebrada Yurac- Yacu, Croat 20888 (F, MO, uc). Cuzco: Prov. La Con- vention, Cordillera Vilcabamba, Rio Klause, Dudley 10182 (GH, us). Puno: Near "San Gavan" (San Gaban), Lechler 2158 (K, p). 1 5. Diplazium subobtusum Rosenst., Repert. Spec. Nov. Regni Veg. 7: 296. 1909. TYPE: Peru, San Martin, Mt. Guayrapurima (as Ecuador, "Monte Guayrapurina"), Spruce 4019 (ho- lotype, P!; frag., us!; photos, F, GH, us). Plants apparently terrestrial. Stem erect, pro- vided at apex with a few broad, castaneous scales. Leaves 75 cm long, 1 -pinna te-pinnatisect above the 2-pinnate basal pinnae, petiole moderately pu- bescent, lacking scales. Lamina 22 cm broad, grad- ually reduced to a pinnatifid apex, axes and veins moderately puberulent abaxially with spreading, flexuous trichomes to 0.3 mm long, the interven- ing tissue also with scattered, shorter trichomes, costae abaxially with a few scattered, appressed, light brown scales, rachis lacking proliferous buds. Pinnae subequilateral, short-stalked, incised near- ly (the basal pair quite) to the costa, the ultimate segments of proximal pinnae narrowly acute, their margins crenate to shallowly lobed. Veins free. Sori single, or frequently diplazioid. Indusia per- sistent, thin, light brown, the margins entire to slightly erose. Known thus far in Peru only from the type col- lection, San Martin. Costa Rica; Panama; Peru. This is one of the questionable species related to D. striatum, from which it differs principally in the great degree of pinna dissection, a feature that is variable in the latter species. Apparently D. sub- obtusum has not been collected again in Peru since the type, nearly 130 years ago. It may be merely a variant ofD. striatum. However, no matter how deeply lobed the pinnae of the latter, the ultimate segments are essentially entire, with obtuse to sub- truncate apices (rarely subacute). Most of the seg- ments in proximal pinnae of D. subobtusum are not only narrowly acute, but their margins are con- spicuously lobed. In this respect, they are some- what intermediate between D. striatum and D. macrophyllum. Furthermore, the leaf is only 75 cm long and 22 cm broad, whereas most leaves of D. striatum are 1-2 m long. Tentatively, then, D. subobtusum is maintained here as distinct. 16. Diplazium lindbergii (Mett.) Christ, Prim. Fl. Costar. 3: 27. 1901. Asplenium lindbergii Mett., Ann. Sci. Nat. Bot. ser. 5, 2: 236. 1864. LECTOTYPE (designated by Lei- linger, Proc. Biol. Soc. Wash. 89: 707. 1977): Bra- zil, Caldes, Lindberg 543 (B). Plants terrestrial. Stem erect or decumbent, sparsely scaly. Leaves to 1.5 m long, 1-pinnate- pinnatisect, petiole dark reddish brown to atro- purpureous, toward the base provided with dull brown, linear or lanceolate scales about 1.5 cm long. Lamina chartaceous to subcoriaceous, to 40 cm broad, generally reduced to a pinnatifid apex, some sparse, filiform scales on the apex, glabrous except for some scattered, septate trichomes abax- ially on the costa and segment midribs, lacking proliferous buds. Pinnae subequilateral, linear or linear-lanceolate, subsessile or short-stalked, in- cised % to % (or rarely quite) to the costa, costae awned adaxially near the base of costules, ultimate segments more than 20 pairs on larger pinnae, obtuse to truncate, often strongly revolute. Veins free. Sori 1 .5-2.5 mm long (slightly longer on basal veins), often diplazioid. Indusia vestigial or lack- ing. Rare in Peru, in wet forests, 1 800-2000 m, San Martin, Pasco, and Cuzco. Southern Mexico to Panama; Venezuela and Colombia, south to Bolivia and Brazil. In this species, the raised, adaxial edges of costae are strongly produced, appearing usually as per- pendicular, herbaceous wings. Where these are in- terrupted near the costule bases, the ends com- monly separate from the costa, as short, subacute awns. A similar condition is evident in D. stria- tum, but the wings are less pronounced and the awns are less conspicuous or lacking in that spe- cies. Several other neotropical species probably should be included here, among them D. grande 78 FIELDIANA: BOTANY (Baker) C. Chr. (Colombia, Ecuador), D. indura- tum Diels (Costa Rica), and D. subnudum (Kar- sten) Alston (Colombia). Each has been separated from D. lindbergii chiefly on the degree of pinna dissection. In Diplazium, this highly variable fea- ture is of little value when unsupported by other characters. Specimens of D. lindbergii are often determined as D. brasiliense Rosenst., which is similar in as- pect and texture. However, the type of the latter (Brazil, Haerchen 91, s!) has broader and fewer segments (to 1 2 pairs), the indusia are subpersis- tent, and the axes are minutely, but densely, pu- berulent abaxially. San Martin: Prov. Rioja, Venceremos, D. Smith & Vasquez 4995 (F, MO, uc). Pasco: Prov. Oxapampa, SE of Oxapampa, D. Smith 2909 (F, MO). Cuzco: Prov. Pau- cartambo, Valle de Pilcopata, Herrera 1623 (us). 17. Diplazium striatum (L.) Presl, Tent, pterid. 114. 1836. Figure 16c-d. Asplenium striatum L., Sp. pi. 2: 1082. 1753. TYPE: Petiver, Pter. Amer. t. 3, f. 3, 4, copied from Plumier, Traite foug. Amer. /. 18, 19, based on a specimen from Martinique. Diplazium obtusum Desv., Prodr. 281. 1827. TYPE: "Crescit in Peruvia" ("Habitat in America cali- diori" on label), collector unknown (holotype, P!; photos, BM, GH, us). Diplazium tabalosense Hieron., Hedwigia 47: 214. 1 908. TYPE: Peru (San Martin), above Tabalosos on the way to Moyobamba, Huallaga River val- ley, Stubel 1089 (holotype, B!; photos, BM, F, OH, us). Plants terrestrial. Stem erect, near the apex sparsely provided with dark brown scales to 1 cm long. Leaves to 2 m long, 1 -pinnate-pinnatifid to nearly 2-pinnate, petiole brown at base, lighter brown to stramineous distally, scaly at base. Lam- ina firm-membranaceous, to 50 cm broad, lacking proliferous buds, reduced to a pinnatifid apex, abaxially glabrous to minute puberulent, and axes sparsely to moderately provided with appressed, attenuate scales. Pinnae (2.5-)3.5-5(-6) cm broad, lanceolate or deltoid-lanceolate, sessile to short- stalked, incised V2 to % (or sometimes quite) to the costa, costae with perpendicular herbaceous wings adaxially, these interrupted at base of costules, there sometimes ending as short awns. Ultimate segments 10-15 pairs or fewer, their apices obtuse to subtruncate (rarely subacute), their margins plane. Sori mostly 3-7 mm long, often diplazioid. Indusia thin, light brown, often shriveling (but persistent) at maturity. In dense forests, often in wet ravines or on river banks, 100-1800 m, Cajamarca, Amazonas, and Loreto south to Cuzco and Madre de Dios. West Indies; southern Mexico to Panama; Ven- ezuela and Colombia to Bolivia. This species varies considerably in leaf dissec- tion. Further study may prove that a number of related taxa are conspecific. Larger pinnae of D. striatum vary from 2 cm broad and cut halfway to the costa, to 6 cm broad and dissected nearly or quite to the costa. Narrower pinnae are more common toward the north of the range, whereas specimens from Peru typically have broader and deeply dissected pinnae, as in the type of D. ta- balosense. The leaves of D. tungurahuae are even more highly dissected. It is similar in most other characters and may better be considered a variety of D. striatum. In fact, some sheets of Schunke 295 (below) are intermediate between the two. A species with narrower, less dissected pinnae is D. angelipolitanum Rosenst. (Colombia), which probably should be included with D. striatum. Cajamarca: Prov. Cutervo, Dist. Cujillo, Tambillo, Jelski 1053 (GH, us). Amazonas: Prov. Bagua, between Aramango and Montenegro, Lopez et al. 4223 (GH, HUT). San Martin: Mt. Campana, near Tarapoto, Spruce 4685 (K, P, w). Loreto: Mouth of Rio Santiago, Pongo de Man- seriche, Mexia 6729 (B, F, GH, K., MO, uc, us). Gamitan- acocha, Rio Mazan, /. Schunke 295 (F, GH, uc, USM). Pasco: Pozuzo, Hacienda Ballisteros, Bryan 697, 698 (F). Junin: Chanchamayo Valley, C. Schunke 1344 (F). Cuz- co: Prov. Paucartambo, Villa Carmen, Vargas 11246, 14683 (GH). Madre de Dios: Parque Nacional del Manu, M. Foster P-84-57, P-84-86 (uc). 18. Diplazium caracasanum (Willd.) Moore, In- dex fil. 324. 1861. Asplenium caracasanum Willd., Sp. pi. ed. 4, 5: 338. 1810. TYPE: Venezuela, Caracas, Bredemeyer (holotype, B!; Herb. Willd. 19919; photos, F, GH, us). Asplenium striatum var. caracasanum (Willd.) D. C. Eaton, Mem. Amer. Acad. Arts 8: 206. 1860. Diplazium shepherdii var. proliferum Rosenst., Re- pert. Spec. Nov. Regni Veg. 7: 294. 1909. SYN- TYPES: Peru (San Martin), "in silvis secus flu- men Mayo," Spruce 4755 (B!, GH!, K!, P!); Spruce 4785 (BM!, P!, us!). Plants terrestrial. Stem erect, provided at apex with a few medium to dark brown scales, these TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 79 coarse, often sublustrous, 3-6 mm long. Leaves 50-90 cm long, 1 -pinnate-pinnatifid, petiole na- ked or with a few dark brown scales at base. Lam- ina 10-20 cm broad, gradually reduced to a pin- natifid apex, glabrous and with scales rare or lacking, bearing a proliferous bud adaxially on the rachis toward the apex. Pinnae 1 6-20 pairs, all but the distal ones cut 34 or nearly to the costa, mostly inequilateral at base, but proximal ones subequi- lateral, segments subacute, larger ones 2-3 times as long as broad. Veins free, pinnately branched in the segments. Sori single, or sometimes dipla- zioid on the basal acroscopic vein branch. Indusia thin-textured, dull brown, but sometimes with a very fine white margin. In forests, usually along streams or in ravines, ca. 700 m, San Martin, Huanuco. Lesser Antilles; Venezuela; Peru. Some taxa in the D. cristatum complex may not merit species status, as they seem to differ only in the degree of pinna dissection. D. caracasanum, however, is distinguished by additional characters, as noted in the key. Furthermore, indusia of the latter often bear a fine white line along the margin, as in the next species, D. stuebelianum. This char- acter, a very rare one in the genus, was not noted in the original descriptions of the two species. Huanuco: Prov. Huanuco, near Tingo Maria, Tryon & Tryon 5230 (BM, F, GH, uc, us). 19. Diplazium stuebelianum (Hieron.) Stolze, stat. et comb. nov. Diplazium shepherdii var. stuebeliana Hieron., Hed- wigia 47: 212. 1908. LECTOTYPE (designated here): Peru (Amazonas?), between Pacasmayo and Moyobamba, Stiibel 70726 (B!; photos, F, GH). PARATYPE: Ecuador, between Banos and Ji- varia de Pintuc, valle Pastaza, Stiibel 986 (B!; pho- tos, F, GH). Plants terrestrial. Stem erect to decumbent, pro- vided at apex with dull, medium brown scales, these somewhat flaccid, 3-4 mm long. Leaves 42- 58 cm long, 1 -pinnate-pinnatisect, petiole naked or with a few dark brown scales at base. Lamina 8-16 cm broad, gradually reduced to a pinnatifid apex, glabrous and without scales, proliferous buds lacking. Pinnae 10-15 pairs, strongly inequilateral at base, proximal ones (at least) incised nearly to the costa, their basal acroscopic segments free and usually short-stalked. Veins free, pinnately branched in the segments. Sori single, or sometimes dipla- zioid on the basal acroscopic vein branch. Indusia thin-textured, dull brown, with a very fine white margin. In forest and thickets, 1200-1700 m, Amazo- nas?, Junin, Ucayali. Ecuador; Peru; Bolivia. Like several other taxa in the species complex (e.g., D. lonchophyllum and D. werckleanum), D. stuebelianum differs from D. cristatum in degree of pinna dissection. However, unlike these others, D. stuebelianum is also distinguished by some good qualitative characters. Scales at the stem apex in D. cristatum and nearest allies are coarse, blackish, sublustrous, and usually less than 3 mm long, whereas in D. stuebelianum they are nearly flaccid, dull brown, and 3-4 mm long. The indusium color is also a very subtle but distinctive character. In fully expanded indusia, there is a very fine, chalky white line along the margin. Indusia of D. crista- tum are concolorous, although a white-edged mar- gin is often present in D. caracasanum. Junin: Schunke Hacienda, above San Ramon, C. Schunke A- 189 (us). La Merced, Chanchamayo, C. Schunke (A, P). Chanchamayo Valley, C. Schunke 112 (us), 729 (F, us). Ucayali: Prov. Coronel Portillo, Sin- chono, Aguilar 892 (GH, USM). 20. Diplazium cristatum (Desr.) Alston, J. Hot. 74: 173. 1936. Figure 16a. Meniscium cristatum Desr. in Lam., Encycl. 4: 94. 1797. TYPE: Martinique, Joseph Martin (holo- type, P!, Herb. Lamarck). Asplenium arboreum Willd., Sp. pi. ed. 4, 5: 320. 1810. TYPE: Venezuela, Caracas, Bredemeyer (holo- type, B!, Herb. Willd. 19892-1; photos, F, GH). Asplenium denticulosum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Na- turk. 5: 323. 1811 (not Gaud. 1827). TYPE: "In America calidiore," Dombey (holotype; P!; photo, GH). Asplenium shepherdii Sprengel, Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 10:231.1821. TYPE: Jamaica, Henry Shepherd (holotype, LZ, destroyed; isotype, PH). Diplazium shepherdii (Sprengel) Link, Hort. berol. 2: 70: 1833. Diplazium arboreum (Willd.) Presl, Tent, pterid. 1 14. 1836. Diplazium denticulosum (Desv.) C. Chr., Index fil. 231. 1905. Plants terrestrial. Stem erect, provided at the apex witih a few castaneous to blackish scales, 80 FIELDIANA: BOTANY these coarse, often somewhat lustrous, 1-3 mm long. Leaves 25-80 cm long, 1-pinnate-pinnatifid, petiole naked or with a few dark brown scales at base. Lamina to 30 cm broad, gradually reduced to a pinnatifid apex, glabrous and lacking scales, proliferous buds absent (or very rarely present). Pinnae 8-12 free pairs, mostly lobed V4 to % to the costa (rarely more deeply), inequilateral at base, i.e., cuneate basiscopically, truncate acroscopical- ly, the basal acroscopic lobe or segment much more strongly produced than the opposing basiscopic one, segments obtuse, 1 or 2 times as long as broad. Veins free, pinnately branched in the segments. Sori commonly diplazioid on the basal acroscopic vein branch, single on the others. Indusia thin- textured, dull brown, the margin subentire. In dense wet or dry forests, often in ravines or along stream banks, 300-1700 m, Amazonas and Loreto to Ayacucho and Madre de Dios. West Indies; Mexico to Colombia and Vene- zuela, south to Argentina and Paraguay. This and near allies form a neotropical species complex that is as taxonomically difficult as that ofAsplenium auritum/ cuspidatum. Because of the variability of leaf dissection it has been divided into a number of species, varieties, and forms. The group is characterized by glabrous, medium-sized, pinnate to pinnate-pinnatisect leaves that taper to a pinnatifid apex, pinnae with conspicuously in- equilateral base (i.e., excavate to cuneate basi- scopically, truncate and much more strongly pro- duced acroscopically), and stem apices bearing only a few, small, coarse, blackish scales. With pinnae deeply lobed to pinnatifid, D. cristatum is at the center of the complex. Taxa with entire to shal- lowly lobed pinnae are D. werckleanum Christ (Mesoamerica), D. unilobum (Poir.) Hieron. (West Indies), and D. bombonasae. The species D. lon- chophyllum Kunze (Mesoamerica) and D. drepa- nolobium A. R. Smith (southern Mexico) have pinnae dissected nearly or quite to the costa. Ul- timately, monographic treatment will probably determine that most of these names should be synonyms of D. cristatum. Two specimens of the latter seen from Peru could easily key out to D. werckleanum, and several others to D. loncho- phyllum in the keys of Stolze (1981) and other recent neotropical Floras. See D. bombonasae and D. stuebelianum for further comparisons. C. Schunke 500 and 944 (F) from Junin, with proximal pinnae deeply dissected, are practically identical to Central American specimens of D. lon- chophyllum. Both specimens also bear proliferous buds distally on the rachis, a condition extremely rare in D. cristatum but a diagnostic feature in D. caracasanum. Amazonas: Prov. Bagua, 12 km E of La Peca, Barbour 2500 (MO, uc). San Martin: River Mayo, near Tarapoto, Spruce 4758 (BM, GH, K, us). Loreto: "Flumen Maranon inf. fluv. Huallaga," Spruce 3911 (K). Pasco: Pozuzo, Hacienda Ballisteros, Bryan 691, 693 (F). Junin: Colonia Perene, Killip & Smith 24925 (F, GH, us). Ucayali: Prov. Coronel Portillo, Dist. Iparia, Bosque Nacional de Imparia, /. Schunke V. 2733 (F, GH, us). Ayacucho: Prov. San Miguel-La Mar, Teresita, Barron (USM). Madre de Dios: Rio Manu, Cocha Cashu Station, Parque Nacional del Manu, Foster et al. 7197 (F). 2 1 . Diplazium bombonasae Rosenst, Repert. Spec. Nov. Regni Veg. 7. 294. 1909. TYPE: Ec- uador, Rio Bombonasa, Spruce (holotype, P!, Herb. Bonaparte 10027; photos, GH & us). Plants terrestrial. Stem erect to decumbent, pro- vided at the apex with a few dark brown to blackish scales, these coarse, sometimes sublustrous, 2-3 mm long. Leaves 35-65 cm long, 1 -pinnate, pet- iole gray-brown, with a few dark brown scales at the base. Lamina to 1 2 cm broad, firm-herbaceous to chartaceous, gray-green, gradually reduced to a subattenuate, pinnatifid apex, essentially glabrous and lacking scales, lacking proliferous buds. Pin- nae 15-24 pairs, approximate to subdistant, fal- cate, attenuate, conspicuously inequilateral at base (attenuate to excavate basiscopically, truncate ac- roscopically and with a pronounced basal auricle), basiscopic margins beyond the basal auricle sub- entire, crenate, or shallowly lobed, or sometimes pinnatifid as to basal pinnae. Veins free, com- monly 1-3-forked. Sori single or diplazioid, borne on the acroscopic branch of each vein. Indusia thin-textured, dull brown, the margins subentire to erose. In forests, on stream banks or on slopes of ra- vines, 180-600 m, Loreto, Huanuco, Madre de Dios. Ecuador; Peru; Bolivia. As in other taxa related to D. cristatum, pinna dissection in D. bombonasae can vary widely. Be- yond the enlarged basal auricle, pinna margins may be essentially entire throughout the lamina (as in the type), or shallowly to deeply lobed. In deeply lobed pinnae the acroscopic auricle is sometimes incised nearly to the costa. South American specimens are sometimes iden- tified as D. unilobum (Poir.) Hieron. of the West TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 81 Indies, a similar taxon with subentire, but strongly auriculate, pinnae. However, stem scales of the latter are 2-3 times longer, the lamina is lighter in color (yellow-green) and often subcoriaceous, and the petiole and rachis are stramineous. Al- though monographic study may prove that both taxa are only variants of D. cristatum, D. bom- bonasae is tentatively maintained here as a distinct species. Loreto: Near Rio Santiago above Pongo de Manse- riche, Mexia 6217 (BM, GH, K, MO, uc). Huanuco: Prov. Leoncio Prado, Dist. Rupa Rupa, 5 km from Tingo Ma- ria, J. Schunke V. 3267 (F, GH, us). Madre de Dios: Prov. Manu, Atalaya, vicinity of Hacienda Amazonia, Foster & Wachter 7445 (F, MO). 22. Diplazium cuneifolium Rosenst, Repert. Spec. Nov. Regni Veg. 12: 470. 1913. TYPE: Bo- livia, North Yungas, Polo-Polo near Coroico, Buchtien 3390 (holotype, s; isotypes, GH!, P!, us!; photo, BM of s, F, & us of P). Plants terrestrial. Stem erect, provided with coarse, broad, dark gray-brown to blackish scales about 5 mm long. Leaves to 80 cm long, 1 -pinnate. Lamina to 24 cm broad, membranaceous, grad- ually reduced to a short, pinnatind apex, lacking scales, sparsely and minutely puberulent on the axes and adjacent tissue abaxially, costae (toward the base) and rachis moderately and minutely pu- berulent within the adaxial sulci. Rachis deeply sulcate adaxially, lacking proliferous buds. Pinnae 11-17 pairs, well-spaced, at least proximal ones short-stalked, broadest near the center, narrowly acute at apex, narrowly to broadly cuneate at the subequilateral base, margins lobed about one- quarter to costa, larger pinnae 10-13 cm long and 2-3 cm broad. Veins free, pinnately branched. Sori borne on most of the vein branches, extending from the costa nearly to the pinna margin, usually diplazioid on the basal branches, the other sori usually single. Indusia thin, dull brown, margin subentire to erose or irregularly fimbriate. In forests, 700-1000 m, Junin, Cuzco, Madre de Dios. Peru and Bolivia. This is a rarely collected species, probably con- fined to lower mountain slopes of Bolivia and the southern half of Peru. In the type collection, pin- nae in the proximal half of the lamina have rather narrowly cuneate bases, but in all other specimens examined the pinna bases are much more broadly cuneate. Considering the variability typical of oth- er species of Diplazium, it seems likely that the narrowly cuneate base seen in the type will be a character more exceptional than representative. Junin: Prov. Satipo, San Francisco de Satipo, Solomon 3315 (F, MO). Prov. Satipo, Reserva Forestal de Universi- dad Central near Satipo, van der Werff et al. 8633 (MO, uc). Cuzco: Prov. La Convention, Palma Real, Vargas 17296 (GH). Madre de Dios: Prov. Manu, Atalaya, vi- cinity of Hacienda Amazonia, Foster & Wachter 7447 (F). 23. Diplazium grandifolium (Sw.) Sw. var. andi- cola Stolze, var. nov. Varietas haec a varietate typica differt indusiis 0.2- 0.4 mm latis, tenuibus, brunneolis, planis (nee usque ad 1 mm latis, nee crassis, nee bicoloribus, nee involutis in maturis). Plants terrestrial. Stem erect or decumbent, pro- vided with coarse, broad, blackish scales about 5 mm long. Leaves to 1.2 m long, 1 -pinnate. Lamina to 28 cm broad, firm-herbaceous to chartaceous, gradually to abruptly reduced to a short, pinnatifid apex, lacking scales, glabrous to minutely puber- ulent on the axes abaxially, rachis and bases of costae puberulent adaxially within the sulci. Ra- chis deeply sulcate adaxially, lacking proliferous buds. Pinnae commonly 7-10 pairs, well-spaced, not or shortly stalked, broadest at or near the base, acuminate at the apex, broadly cuneate to truncate at the subequilateral base, margins entire to broad- ly crenulate, larger pinnae 8-15 cm long and 2- 3.5 cm broad. Veins free, pinnately branched. Sori borne on most of the vein branches, extending from the costa to a few millimeters from the mar- gin, usually diplazioid on the basal acroscopic branch, the other sori single or diplazioid. Indusia dull light brown, flat, about 0.2-0.4 mm broad, the margins subentire to erose. TYPE— Peru, Loreto, Gamitanacocha, Rio Ma- zan, J. Schunke 281 (holotype, us!; isotypes, F!, GH!, USM!). In deep, wet forests, often in ravines, 100-1500 m, San Martin and Loreto to Madre de Dios and Puno. Venezuela; Ecuador; Peru; Bolivia; western Bra- zil. This variety differs from D. grandifolium var. grandifolium in the delicate, narrow, dull light brown indusia that do not change essentially in color and form, whereas developing and mature 82 FIELDIANA: BOTANY indusia of var. grandifolium are strikingly differ- ent. Immature sporangia are protected by a broad (often to 1 mm) bicolorous indusium, dark brown to blackish and very firm at its attachment along the vein, but thin and often hyaline toward the margin. As sporangia mature, the indusium rolls back tightly to reveal a dark, usually lustrous, un- derside. This is nearly identical with the indusium of D. celtidifolium, whereas that of var. andicola may be compared with D. centripetale (Baker) Maxon, of Venezuela, Ecuador and the West In- dies. Diplazium grandifolium var. grandifolium is common in the West Indies, and from Central America to Colombia, whereas var. andicola is primarily Andean, although a single specimen has been seen from Brazil, near the Peru-Bolivia bor- der. The ranges overlap in Venezuela and Ecuador, where a few specimens of each have been found. San Martin: Prov. Mariscal Caceres, Tocache Nuevo, /. Schunke V. 8489 (F, MO). Loreto: Santa Rosa, lower Rio Huallaga below Yurimaguas, Killip & Smith 28911 (us). Huanuco: Prov. Huanuco, Tingo Maria, Tryon & Tryon 5224 (BM, GH, us). Pasco (as Junin): Cahuapanas, on Rio Pichis, Killip & Smith 26781 (us). Junin: Chan- chamayo Valley, C. Schunke 142, 451 (F, us). Ucayali: Prov. Coronel Portillo, Dist. Iparia, Bosque Nacional de Iparia, /. Schunke V. 2687 (F, GH, us). Madre de Dios: Prov. Tambopata, Tambopata Nature Reserve, Barbour 4963 (F, MO, uc). Puno: Prov. Carabaya, San Gaban, Vargas 18934 (GH). 24. Diplazium celtidifolium Kunze, Bot. Zeit. (Berlin) 3: 285. 1845. TYPE: Venezuela, Ca- racas, Linden 544 (holotype LZ, destroyed; isotypes, FI, K!, P!; photos, s of FI & K). Plants terrestrial. Stem erect to decumbent, this and petiole base provided with coarse, dark brown, linear or linear- lanceolate scales 4-14 mm long. Leaves 1-2 m long, 1 -pinnate. Lamina 30-60 cm broad, herbaceous to subcoriaceous, terminating in a short, pinnatifid apex, the axes glabrous or densely puberulent and sparsely to amply provid- ed on the abaxial side with flaccid, appressed, brown scales, these mostly linear to filiform, es- pecially abundant on the costae. Rachis deeply sulcate adaxially, sometimes bearing proliferous buds at the bases of distal pinnae. Pinnae 8-12 pairs, well spaced, conspicuously stalked, long- acuminate at apex, broadly cuneate to truncate at the subequilateral base, margins entire to shallow- ly crenate-dentate, larger pinnae 12-26 cm long and 2.5-7 cm broad. Veins free, pinnately branched. Sori borne on most of the vein branch- es, extending from the costa to about 5 mm from the margin, always diplazioid on the basal acro- scopic vein, the other sori single or diplazioid. Indusia bicolorous, brown and blackish, at ma- turity rolled back to the blackish, often lustrous base. Two varieties are recognized here. Key to Varieties a. Rachis and costae glabrous; leaves to 2 m long and 60 cm broad, chartaceous to subcoriaceous; larger pinnae 1 7-26 cm long, 4-7 cm broad 24a. var. celtidifolium a. Rachis and costae minutely but densely puberulent; leaves to 1 m long and 35 cm broad, herbaceous; larger pinnae 12-17 cm long, 2.5-3.5 cm broad 24b. var. puberulum 24a. Diplazium celtidifolium var. celtidifolium. Diplazium callipteris Fee, (Mem. foug. 5) Gen. fil. 2 1 4. 1852. PROBABLE TYPE: Venezuela, Funck & Schlim 233, in 1845-1846 (holotype, not located; isotype, BM!); originally cited in error as Linden 233 from Cuba (see discussion above). Asplenium callipteris (Fee) Baker, Syn. fil. 231. 1867. Asplenium celtidifolium (Kunze) Baker, Syn. fil. 232. 1867. Athyrium celtidifolium (Kunze) Milde, Bot. Zeit. (Ber- lin) 28: 353. 1870. In wet forests, 500-1050 m, Amazonas, San Martin, Puno. Trinidad; French Guiana; Surinam; Venezuela; Colombia; Peru; Brazil. There has been confusion in the nomenclature, synonyms, and typification of names associated with D. celtidifolium. Apparently Fee erred when citing the type of D. callipteris as Linden 233 from Cuba. During studies of Diplazium in Peru, Lin- den 233 could not be located. However, there is a TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 83 Funck & Schlim 233 from Venezuela (BM), marked as an isotype, that matches the original description in every way. Linden, half-brother to Schlim, col- lected with him in Venezuela, and it is possible names and numbers became mixed, bringing about Fee's mistake in citing the type. Therefore, if fur- ther search does not produce Linden 233, the Brit- ish Museum specimen must be considered at least an isotype. Furthermore, since this specimen does not differ significantly from the type of D. celti- difolium, the two names must be synonymous. A number of specimens from the Lesser Antilles are found in herbaria identified as D. callipteris. However, all of these are apparently D. legalloi Proctor, a similar species with coarser leaves, usu- ally conspicuously crenate pinna margins, and del- icate, light brown indusia. Amazonas: Prov. Bagua, Montenegro-Chiriaco, Sa- gdstegui 5923 (GH). San Martin: Mt. Guayrapurima, Spruce 4760 (K, p, us). Puno: Prov. Carabaya, San Ga- ban, Vargas 18933 (GH). 24b. Diplazium celtidifolium var. puberulum Stolze, var. nov. Varietas haec a varietate typica differt foliis usque ad 1 m longis et 35 cm latis, herbaceis, pinnis grandioribus 12-17 cm longis et 2.5-3.5 cm latis, rachidi et costis dense puberulis. Leaves to 1 m long and 35 cm broad, herbaceous (not chartaceous to subcoriaceous). Larger Pinnae 12-17 cm long and 2.5-3.5 cm broad. Rachis and costae densely puberulent on both sides. TYPE— Peru, Pasco (as Junin), Pichis Trail, "Ya- pas" [Yapaz], dense forest, 1350-1600 m, Killip & Smith 25508 (holotype, us!; isotype, GH!; pho- tos, F & GH of us). Thus far known only from the type, Pasco, and one paratype, cited below. Although the most obvious difference between this and var. celtidifolium is in the smaller and thinner-textured leaves, this is not merely a de- pauperate form of D. celtidifolium, for it differs also in the puberulent axes. In the typical variety, there is no laminar indument other than the scales on the rachis and costae abaxially; but in var. pu- berulum the axes are densely covered on both sides with minute trichomes about 0.1-0.2 mm long. Such trichomes are found with greater or lesser frequency in others of the species complex, notably such taxa as D. grandifolium, D. centripetale (Ba- ker) Maxon of the West Indies and northern South America, and D. eggersii Sodiro of Ecuador (the latter most likely conspecific with D. centripetale, see Comments). This is a character that needs more attention in future monographic work on the ge- nus. Pasco (as Junin): Pichis Trail, "Yapas" [Yapaz], Killip & Smith 25457 (us). 25. Diplazium paucijugum Stolze, sp. nov. Caulis paleis crassis, nigellis; folium usque ad 65 cm longum et 24 cm latum, 1-pinnatum, longipetiolatum, parte apicali pinnatifida cerca dimidia longiores quam lamina; rachis et costa in pagina adaxiali paleis flaccidis, appressis, linearibus vel nliformibus; pinnae 2-4-juga- tae, apicibus acuminatis, basibus subaequilateribus, truncatis vel subcordatis; venae liberae, pinnatiramosae; indusia crassa, bicoloria, brunnea et nigella, maturitatate involuta. Plants terrestrial. Stem erect to decumbent, pro- vided with coarse, broad, blackish scales about 5 cm long. Leaves to 65 cm long, 1 -pinnate, on a petiole nearly the length of the lamina. Lamina to 24 cm broad, firm-herbaceous, the pinnatifid api- cal portion of the leaf about half the total length of the lamina, essentially glabrous, but axes sparse- ly puberulent abaxially, to glabrate, also provided on the abaxial side with flaccid, appressed, brown scales, these mostly linear to filiform, especially abundant on the costae. Rachis deeply sulcate adaxially, lacking proliferous buds. Free pinnae 2- 4 pairs at base of lamina, approximate to subdis- tant, acuminate at apex, truncate to subcordate at the subequilateral base, basal pair short-stalked, margins entire, or basal pinnae broadly crenate, larger pinnae 10-13 cm long, 3.5-4 cm broad. Veins free, pinnately branched. Sori borne mostly on the basal acroscopic vein branches, extending from near the costa to about 5 mm short of the margin, mostly single, occasionally diplazioid. Indusia coarse, bicolorous, brown and blackish, at matu- rity rolled back to the blackish base. TYPE— Peru, "In monte Campana, prope Tara- poto, Peruviae Orientalis" (Dept. San Martin), Spruce 4339 (holotype, K!; isotype, P!; photos, F & GH Of K & P). Thus far known only from the type and one other collection from Huanuco, in jungle on ridge east of Tingo Maria, 625-1 100 m, cited below. This species is distinguished from others in Peru by the unusual configuration of the lamina. On a 84 FIELDIANA: BOTANY petiole nearly as long as the lamina are borne a few pairs of discrete pinnae, and the rest (half or more) of the lamina is gradually reduced to a pro- longed, pinnatifid apex. A similar condition occurs in D. riedelianum (Kuhn) C. Chr. of Brazil and (probably a synonym) D. verapax (Donn.-Sm.) Hi- eron. of Mesoamerica. However, those taxa have narrower laminae, much smaller pinnae with cu- neate bases and long-tapering tips, and several proliferous buds in the axils of proximal pinnae. Diplazium paucijugum is closely related to D. celtidifolium, especially in the narrow, flaccid scales on the abaxial axes and the glabrous adaxial axes. However, it differs from the latter species es- pecially in the unusual lamina shape, the few and smaller pinnae, the lack of proliferous buds and by the minutely puberulent (to glabrate) axes abax- ially. A proliferous bud is often found distally on the rachis in D. celtidifolium, and axes are totally lacking in indument except for the abaxial scales on rachis and costae. Huanuco (as San Martin): E of Tingo Maria, Allard 22334 (GH, us). 26. Diplazium roemerianum (Kunze) Presl, Tent, pterid. 113. 1836. Figure 16b. Asplenium roemerianum Kunze, Linnaea 9: 62. 1834. TYPE: Peru, "in argillosis humidis ad Pampaya- co" (Pampayacu, Huanuco), Poeppig 166, July, 1829 (holotype, B!; isotypes, B!, K!, L, P!; photos, F, GH, MO & us of L). Asplenium Jlavescens Mett., Ann. Sci. Nat. Bot., ser. 5, 2: 234. 1864, sp. nov. based on Hooker, Fil. exot. /. 100. 1859, not Asplenium juglandifolium Lam. TYPE: Cultivated at Kew from a Venezu- elan plant collected by Wagener, this evidently illustrated in /. 100 (holotype, K; isotype, BM!). PARATYPE: Venezuela, Colonia Tovar, Fendler 498 (K). ISOPARATYPE: (GH!). Sloane, Hist. Jam. "Filix maxima . . ." 82, t. 371 is excluded. Jen- man, Bull. Bot. Dept. Jamaica n.s. 1(5): 77. 1894 says that the Sloane specimen at BM is a sterile leaf of Acrostichum cervinum (= Olfersia cervina). Diplazium flavescens (Mett.) Christ, Farnkr. Erde 217. 1897. 2-8 pairs, well-spaced, short-stalked, acuminate at apex, broadly to narrowly cuneate at the sub- equilateral base, the margins entire to crenulate- serrate. Veins 1 -forked, free, the branching vein- lets spreading from the costa at 50- to 70-degree angles and gently arching to the pinna margin. Sori mostly borne on the acroscopic vein branch, thus gently arched like the veinlet. Indusia borne most- ly along one side (occasionally on both sides) of a veinlet, light to reddish brown, the margins erose to subentire. In wet forests, 700-2300 m, Loreto and Ama- zonas south to Cuzco. Greater Antilles; Guadeloupe; Venezuela; Co- lombia; Ecuador; Peru; Brazil. This and D. flavescens have been separated on the basis of relative number of pinnae and serrate vs. entire pinna margins: typical D. flavescens with 6-8 pairs of pinnae and crenulate margins; D. roe- merianum with 1-2 pairs of pinnae and entire margins. The two characters are inconsistent, and there are no other significant differences. Pinnae with entire margins are often serrate toward the apex, and when many specimens are compared throughout the range it becomes apparent that margins are highly variable, whether on laminae with few or many pinnae. It would appear that the type of D. Jlavescens simply represents a more ro- bust form of the species. The Colombian Asplenium caucense Karsten and A. ocanniense Karsten also surely belong here; however the Central American taxon once clas- sified as D. flavescens var. proliferum Christ has been recognized (Stolze, 1981) as a quite distinct species, D. obscurum Christ. Amazonas: Prov. Bagua, 12 km E of La Peca, Barbour 2647 (F, MO, uc). San Martin: Eastern Peru, near Tarapo- to, Spruce 4674 (BM, P). Loreto: Sierra del Pongo, Mexia 627 3a (GH, K., uc, us). Huanuco: Fundo Chela, Sinchono, Aguilar 920 (USM). Pasco: Pozuzo, Hacienda Ballisteros, Bryan 672 (F, us). Junin: Chanchamayo Valley, C. Schunke 8 (P), 72 (F, us), 73, 802 (F). Cuzco: Prov. La Convencion, Guayanay, Vargas 13238 (GH). Plants terrestrial. Stem erect or decumbent, this and the petiole base provided with 3-7 mm long, blackish, linear to lanceolate, subentire scales. Leaves to over 1 m long, 1 -pinnate, long-petiolate. Lamina to 35 cm broad, firm-herbaceous, termi- nating abruptly in a nearly conform apical segment (this occasionally with a large basal lobe), glabrous, with scattered filiform scales on the rachis. Pinnae 27. Diplazium lechleri (Mett.) Moore, Index fil. 141. 1859. Asplenium lechleri Mett., Fil. lechl. 1: 16, /. 2. 1856. TYPE: Peru (Puno), "St. Gavan" (San Gaban), "in sylvis montanis," Lechler 2269a (holotype, B; isotypes, K!, L; photos, F, GH & us of L, us of K). TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 85 Plants terrestrial. Stem erect or decumbent, this and the petiole base scaly, the scales 7-12 cm long, dark brown, rigid, linear to lanceolate, attenuate, subentire. Leaves to 2 m long, 1 -pinnate, long-pet- iolate. Lamina to 35 cm broad, chartaceous to subcoriaceous, terminating abruptly in a conform apical segment, subglabrous, but axes (and some- times the laminar surface) sparsely provided on abaxial side with filiform scales that grade into dark pluricellular trichomes. Pinnae 6-9 pairs, subdistant, short-stalked, acuminate at apex, rounded to cuneate at the subequilateral base (al- though basal pinnae sometimes narrow-cuneate at the basiscopic base). Veins simple, or paired at the costa, straight, spreading at broad (75- to 80-de- gree) angles, free, except connected at their tips by a slightly inframarginal vein. Sori straight, crowd- ed, and strictly parallel, mostly diplazioid. Indusia dark brown, firm and persistent. San Martin to Puno. Costa Rica; Surinam; Venezuela; Colombia; Peru; Brazil. This species is easily distinguished by an apical segment that is nearly identical to the lateral pin- nae, the crowded, straight sori borne at nearly right angles to the costa, and the mostly simple veins that are free almost to the margin, where they join an inframarginal vein. San Martin: Mount Guayrapurima, Spruce 4687 (K, p). Huanuco: SW slope of Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13085, 13381 (GH). Junin: Pichis Trail, Porvenir, Killip & Smith 25922 (us). Cuzco: Prov. Paucartambo, Cosnipata Valley, Rio Tono, Wachter et al. 166 (F). Madre de Dies: Prov. Manu, Cerro de Pantiacolla, Rio Palotoa, Foster et al. 107 05 -A (F). 28. Dipla/ium plantaginifolium (L.) Urban, Symb. antill. 4: 31. 1903. Asplenium plantaginifolium L., Syst. nat. ed. 10, 2: 1323. 1759. TYPE: P. Browne, Jamaica (not lo- cated). Proctor, Ferns of Jamaica, p. 394. 1985, designated a (necessary?) neotype: Maxon 1949, Jamaica, St. Catharine, Mt. Diablo (us!; isoneo- type, BM!). Asplenium plantagineum L., Sp. pi. ed. 2: 1537. 1763, nom. superfl. for A. plantaginifolium L. and with the same type. Diplazium plantagineum (L.) Sw., J. Bot. (Schrader) 1800(2). 62. 1802. Plants terrestrial. Stem erect or decumbent, this and the petiole base sparsely provided with deep reddish brown or blackish scales, these linear or narrow-deltoid, 1-3 mm long. Leaves to 60 cm long, simple, entire to sinuate, or crenate-serrate apically, rarely bearing a proliferous bud at base of lamina. Petiole nearly as long as, or longer than, the lamina. Lamina to 6 cm broad, glabrous, lan- ceolate to oblong-lanceolate, acuminate at apex, broadly cuneate to truncate at base. Veins spread- ing from the costa at 50-75 degrees, with 2-4 pairs of strongly ascending branches running parallel with the primary vein, to the leaf margin. Sori (many of them) extending nearly from rachis to margin and borne on both sides of a vein. Indusia very narrow, delicate and often partly deciduous. Southern Mexico to Panama; West Indies; Ven- ezuela; Peru; Bolivia; Brazil. In rain forests, 800-1800 m, Junin. Only a few American species of Diplazium have simple leaves, two of which occur in Peru; but they are easily distinguished by the characters used in the key. Junin: East of Quimiri Bridge, near La Merced, Killip & Smith 23971 (GH, us). Chanchamayo Valley, C. Schunke 457 (F). 29. Diplazium pinnatifidum Kunze, Linnaea 9: 72. 1834. TYPE: Peru (Huanuco), "In crepidi- nibus aquosis, lutosis, umbrosissimus," Cu- chero, Poeppig diar. 1149 July, 1829 (holo- type; w!; probably isotype, K!; photo, BM of w). Anisogonium pinnatifidum (Kunze) Presl, Tent, pte- rid. 116. 1836. Asplenium kunzei Mett., Fil. hort. hot. Lips. 74. 1 856, nom. nov. for Diplazium pinnatifidum Kunze, and with the same type. Not Asplenium pinnatifidum Nutt. 1818. Plants terrestrial. Stem erect or decumbent, this and the petiole base sparsely paleate, the scales 3- 4 mm long, dull brown, ovate or lanceolate, com- monly subentire, but sometimes with castaneous to blackish marginal cells that develop elongated teeth that are bifid at apex. Leaves to 1 m long, mature ones pinnatifid or pinnatisect, often pin- nate at base, long-petiolate, juvenile ones suben- tire to shallowly lobed. Lamina to 28 cm broad, coriaceous, tapering gradually to a pinnatifid apex, abruptly reduced at base, cut deeply to the rachis, the rachis and costae usually concealed abaxially by copious, amorphous, brown scales, and some- times with a few, spreading, dark-toothed ones like 86 FIELDIANA: BOTANY those of the stem. Segments 3-7 pairs, joined by an acute to broadly rounded sinus (or the basal 1- 2 pairs sometimes fully discrete), the apex obtuse to acute, the surfaces glabrous. Veins copiously anastomosing from costa to margin. Sori dipla- zioid, commonly branching to follow the veins. Indusia dull brown, narrow, erose to fimbriate, very delicate and early deciduous. In forests, in wet, shady places, often on hillsides and ravine banks, 250-2500 m, Amazonas and Loreto to Cuzco and Madre de Dios. Colombia; Ecuador; Peru; Bolivia. The protologue ofDiplazium pinnatifidum con- tains the exact citation of locality and habitat as on the type label (above), except that the former reads "Pampayaco" instead of Cuchero. Both sites are in Huanuco. The two areolate species of Diplazium in Peru not only are easily distinguished by the characters in the key; they differ also in their scales. In D. praestans these are filiform and entire; in D. pin- natifidum they are broad and often conspicuously dentate, with dark setiform teeth. The areolate ve- nation and these singular scales indicate close re- lationship with D. aberrans Maxon & Morton and D. pactile Lell. (Colombia and Ecuador) and D. chimborazense (Baker) Christ and D. macrodic- tyon (Baker) Diels (Ecuador). Some authors be- lieve the characters help define a natural species group or subgenus: Diplazium subgenus Aniso- gonium. Diplazium pactile differs from D. pinnatifidum in the thin-textured and pubescent leaf tissue; D. chimborazense has larger, thinner leaves, attenu- ate pinnae, and the veins merge only toward the pinna margin. Diplazium aberrans differs in its simple, oblanceolate leaves, with petiole short or lacking; D. macrodictyon has stellate trichomes on the abaxial surface and veins. Another taxon, D. fuscum (Baker) C. Chr. of Ecuador, seems to be merely a robust form of D. pinnatifidum; an iso- type and two authentic specimens at Kew are like the latter in every respect, except that their larger leaves have six or seven discrete pinnae, one pair of which has lobed margins. Amazonas: Prov. Bagua, E of La Peca, Barbour 2588, 2814 (F, MO, uc). San Martin: Mt. Guayrapurima, near Tarapoto, Spruce 4686 (B, BM, GH, p). Loreto: Pumayacu, between Balsapuerto and Moyobamba, Klug3218 (F, GH, K, MO, us). Huanuco: Tingo Maria, Tryon & Tryon 5280 (BM, F, GH, uc, us). Pasco: Prov. Oxapampa, Puerto La- guna, D. Smith 8448 (uc). Junin: Chanchamayo Valley, C. Schunke 515 (F, us). Ucayali: Prov. Coronel Portillo, Distr. Calleria, J. Schunke V. 3080 (F, GH, us). Cuzco: Santa Isabel, Valle Cosnipata, Scolnik 930 (us). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10718 (F). 30. Diplazium praestans (Copel.) Morton, Contr. U.S. Natl. Herb. 38: 41. 1967. Athyrium praestans Copel., Amer. Fern J. 38: 132. 1948. TYPE: San Martin, Chazuta, Rio Huallaga, Klug 4002 (holotype, us!; isotypes, F!, GH!, MO!, P!, uc!). Plants terrestrial. Stem erect or decumbent, oc- casionally short-creeping, provided with medium to dark brown, filiform scales, these 2-3 mm long and only several cells broad, their margins entire. Leaves to 35 cm long, simple, entire to broadly sinuate, lacking a proliferous bud. Petiole 0-5 cm long, with scales like those of the stem, but longer and broader. Lamina to 9 cm broad, glabrous, elliptic to (commonly) oblanceolate, subacute at apex, long-attenuate at base. Veins copiously anas- tomosing. Sori linear, gently arching, up to 5 cm long, mostly single (rarely double). Indusia nar- row, firm, persistent. In dense forests, in wet places in deep shade, often along banks of rivers and streams, 100-900 m, San Martin and Loreto to Junin and Madre de Dios. Peru; Bolivia; Amazonian Brazil. Another species with simple leaves and copi- ously anastomosing veins is Diplazium aberrans Maxon & Morton of Colombia and Ecuador, but this differs from D. praestans in its larger (to 60 cm) leaves and longer and narrower areoles. It differs even more significantly in its peculiar scales, which are borne along the costa, and sometimes veins, abaxially, as well as on the stem and petiole. These scales (at least of the stem) have castaneous to blackish marginal cells that develop elongated, bifid teeth. Scales on D. praestans (confined to the stem and petiole) are entire and concolorous. For further discussion of these scales see D. pinnati- fidum. San Martin: Prov. Mariscal Caceres, Distr. Campani- lla, Quebrada de Mashuyacu, J. Schunke V. 4250 (F, GH, us). Loreto: Rio San Alejandro, Woytkowski 5118 (GH, MO, uc, us). Huanuco: Prov. Leoncio Prado, Rio Mon- zon, near Bella, Plowman 5875 (GH). Junin: Rio Pinedo, N of La Merced, Killip & Smith 23621 (GH, us). Ucayali: Prov. Coronel Portillo, Distr. Iparia, Bosque Nacional de Iparia, J. Schunke V. 2752 (GH, us). Madre de Dios: TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 87 Prov. Manu, Parque Nacional Manu, R. Foster 11722 (F). Comments Diplazium centripetale (Baker) Maxon, Pterido- phyt. Porto Rico 441. 1926. Asplenium centripetale Baker, Syn. fil. ed. 2: 490. 1874. LECTOTYPE (designated by Proctor, Ferns of Jamaica): Jamaica, Macfayden (K). Asplenium eggersii Sodiro, Crypt, vase. Quit. 188. 1 893. TYPE: Ecuador, between Bodegas and Bal- sapamba, Sodiro, (holotype, not located; probable isotype, P!; frag., us!; photos, F & us of p). Diplazium eggersii (Sodiro) C. Chr., Index fil. 231. 1905. This is very similar to D. celtidifolium var. pu- berulum, especially in the pinna size and the pu- berulent axes. It occurs in the West Indies, Ven- ezuela, and Ecuador, and might be expected in Peru. The most obvious differences from the latter species are in the characters of sori and pinnae. In D. centripetale, sori extend only halfway to % to the pinna margin, and indusia are thin, flat, and dull light brown throughout. The 15-25 pairs of pinnae are commonly sessile. In D. celtidifolium, sori extend from the costa to within 5 mm of the pinna margin, and indusia are firm, bicolorous (brown distally and blackish proximally), and at maturity are rolled back to the blackish, often lus- trous, base. There are only 8-12 pairs of pinnae, most of which are conspicuously stalked. In the protologue, the type of Asplenium eggersii was said to be collected in Ecuador, between Bo- degas and Balsapamba. Thus far no specimen has been found exactly designating this locality. How- ever, there are two sheets at Paris collected by Sodiro in January, 1891 "in reg. tropic, inter Bo- degas et Pisagua," one of which is inscribed in Sodiro's hand as "Asplen. (Dipl.) eggersii, Nov. Sp." Given the confusion that usually attends So- diro specimens, descriptions, and type localities, this is likely to be an isotype (if not the holotype) of the name. In any case, these specimens at Paris, and other Ecuador collections made in adjacent areas and determined as D. eggersii, quite match specimens of D. centripetale from the West Indies. Key to Species of Athyrium a. Lamina 2-pinnate to 2-pinnate-pinnatifid; ultimate segments rather sharply dentate 1 . A. dombeyi a. Lamina 4- to 5-pinnate; ultimate segments entire and elongate, bifid, or lobed . . 2. A. ferulae-emu XVII. Athyrium Athyrium Roth, Tent. fl. germ. 3: 58. 1 799. TYPE: Athyrium filix-femina (L.) Roth (Polypodium filix-femina L.). Figure 17. Stem usually decumbent to nearly erect, usually short-creeping and moderately stout, or long- creeping and slender, bearing scales, these not or obscurely clathrate, brown, concolorous. Leaves ca. 20 cm to 2 m long, petiole continuous. Lamina 1 -pinnate-pinnatifid to 5-pinnate, nearly glabrate, somewhat scaly, or slightly pubescent or glandular, monomorphic or nearly so. Veins free. Sori round- ish or usually elongate on one side of a vein, or sometimes also on the other side distally, not pa- raphysate, usually covered by reniform to elongate or hook-shaped indusia, or essentially exindusiate. Spores rather ellipsoidal, monolete, variously ridged. Athyrium is a large genus of about 100 species, most of them in eastern and southeastern Asia. There are two species in the Andes and in Peru. The genus is technically separated from Dipla- zium by its chromosome number of n = 40 or multiples, and the indusia either only on one side of a vein, or if on the other side then only distally and continuous around the apex of the sorus. Di- plazium has a chromosome number of n = 41 and multiples, and usually some sori are on both sides of a vein and distinct distally. Athyrium ferulaceum and related species of Cen- tral America are placed here on the basis of the chromosome number of n = 40 (Gomez, Atti 1st. Bot. Univ. Pavia 7: 30. 1971, as "Athyrium bra- dearum") (= Diplazium bradeorum), and the in- dusia that are commonly only on one side of a vein, or rarely partly on the other side and con- tinuous distally. The group is of uncertain affinity and the spores resemble species of Diplazium more than those of Athyrium. However, it seems closer to Athyrium in its chromosome number and more obvious characters. 88 FIELDIANA: BOTANY FIG. 17. Athyrium dombeyi: a, habit; b, pinnule, abaxial side. (From Tryon & Tryon 6029, Columbia, GH.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 89 1. Athyrium dombeyi Desv., Mem. Soc. Linn. Paris 6: 266. 1827. (as dombei). TYPE: "Pe- ruvia (Herb. Mus. Paris)," presumably Peru, Dombey (holotype, P). Figure 17a-b. Stem stout, decumbent to erect, bearing brown to dark brown, elongate scales. Leaves ca. 20 cm to 1 m long, the petiole mostly glabrous except for usually persistent scales near the base. Lamina 2-pinnate to 2-pinnate-pinnatisect, glabrous or very slightly scaly, pinnae sessile to usually short- stalked, the basal usually somewhat reduced, ul- timate segments rather sharply dentate. Sori more or less elongate, on one side of a vein, rarely some on the other side distally, covered by an indusium of similar extent. In ravines, on damp banks, along streams, and at the edge of aqueducts, 2500-3300 m, Lamba- yeque to Cuzco. In South America southward to southeastern Brazil and Argentina; perhaps also northward to Central America, Mexico, and Hispaniola. Athyrium dombeyi is a member of the Athyrium filix-femina complex which is mostly temperate and boreal in distribution and it is perhaps a va- riety of that species: Athyrium filix-femina var. dombeyi (Desv.) Hieron. Lambayeque: Prov. Ferrenafe, 4 km NW of Incahuasi, Dillon & Skillman 4157 (F, GH). Cajamarca: Prov. Ca- jamarca, Dist. Encanada, Sanchez 318 (GH). Amazonas: Prov. Bagua, E of La Peca, Barbour 2759 (F). La Libertad: Prov. Otuzco, above Agalpampa, Sounders 894 (GH), 595 (F, GH). San Martin: Dist. Huallaga, 30 km above Jucusbamba, Hamilton & Holligan 929 (us). Prov. Mar- iscal Caceres, between Mirador and La Playa, Young & Leon 4932 (F). Ancash: Above Yungay on road to Huan- ganuco, Correll & Smith P963 (GH). Huanuco: Mima, Macbride 4327 (F, us). Cani, Macbride 3404 (F, us). Cuz- co: Prov. Paucartambo, vicinity of Achirani, Vargas 11139(F,vc). pinnae sessile to short-stalked, the basal the larg- est, ultimate segments entire and elongate, bifid, or lobed. Sori elongate, on one side of a vein, rarely some on the other side distally, covered by an indusium of similar extent. A single collection known from Peru: in humus, 1450 m, Cuzco. Elsewhere in forests, on ravine banks, and rarely on sandstone cliffs, 100-2700 m. Central America south to Peru. Cuzco: Prov. La Convention, Choquellowanca, Var- gas 12928 (GH). XVIII. Hemidictyum Hemidictyum Presl, Tent, pterid. 1 10. 1836. TYPE: Hemidictyum marginatum (L.) Presl (Asple- nium marginatum L.). Figure 18. Stem erect to decumbent, stout, bearing scales. Leaves ca. 1 to 3 m (or more) long, petiole con- tinuous. Lamina 1 -pinnate, the pinnae entire, gla- brous. Veins free near the costas, fully anasto- mosing toward the margin, without included free veinlets, the vein ends connected by a marginal vascular strand. Sori elongate along the veins, not paraphysate, covered by elongate indusia. Spores rather ellipsoidal, monolete, prominently ridged or saccate. Hemidictyum is a monotypic, morphologically and cytologically isolated genus of the American tropics. The chromosome number of n = 31 may raise doubts as to its proper inclusion in the Dryopteridaceae, which have n = 40 or 4 1 . How- ever, in other characters, the alliance of the genus seems to be with that family and with the Tribe Physematieae. 2. Athyrium ferulaceum (Hooker) Christ, Bull. Herb. Boissier, ser. 2, 4: 968. 1904. Asplenium ferulaceum Hooker, Sp. fil. 3: 216. 1860. SYNTYPES: Colombia (New Grenada), Hartweg 1519; Ecuador, Quito, Jameson (both K). Diplazium ferulaceum (Hooker) Lell., Proc. Biol. Soc. Wash. 98: 376. 1985. Stem small to stout, decumbent to erect, bearing broad, brown scales, especially at the apex. Leaves ca. 30 cm to 1 m long, the petiole essentially gla- brous. Lamina 4-5 pinnate, glabrous or nearly so, 1. Hemidictyum marginatum (L.) Presl, pterid. 111. 1836. Figure 18a-c. Tent. Asplenium marginatum L., Sp. pi. 1082. 1753. TYPE: Either LINN 1250.20 or Petiver, Pteri-graph. Amer. /. 12, f. 2, both of which are this species, the latter based on Plumier, Traite foug. Amer. t. 106. Stem apex densely scaly, with brown, linear scales. Leaves ca. 1 to 3 m (or more) long, the petiole glabrous or somewhat scaly near the base. Lamina 1 -pinnate, with a conform apical segment, 90 FIELDIANA: BOTANY FIG. 18. Hemidictyum marginatum: a, leaf apex; b, portion of pinna, abaxial side; c, pinna margin, adaxial side. (From Stolze, Ferns & fern allies of Guatemala, 1981.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 91 pinnae large, entire, thin. Veins with the ends con- nected by a nearly marginal vascular strand. Sori elongate, often very long, borne along part of a free vein, toward the costa from the anastomosing veins, indusium of similar extent, very thin. Steep banks in forests, in dense forests and in cloud forests, 260-1 700 m, Amazonas to Ucayali. Southern Mexico and the Greater Antilles, south to Bolivia and southeastern Brazil. Amazonas: Prov. Bagua, Quebrada Mirana, Rio Ma- ranon, above Cascadas de Mayasi, Wurdack 1890 (GH, us). Prov. Bagua, 12 km E of La Peca, Barbour 2582 (MO). San Martin: Tarapoto, Spruce 4783 (GH, us). Cha- zuta, Rio Huallaga, Klug 3932 (GH, us). Loreto: Between Rio Amazonas and Rio Napo, Croat 19556 (MO). Hua- nuco: Prov. Huanuco, Tingo Maria, Tryon & Tryon 5254 (GH, us). Prov. Huanuco, Gasa, Stork & Morton 9874 (F, us). Pasco: Pichis Trail, Enenas, (as Junin), Killip & Smith 25765 (F, us). Prov. Oxapampa, Paujil, Leon 285 (USM). Prov. Oxapampa, Villa Rica, van der Werffet al. 8304 (MO). Junin: Near La Merced, Killip & Smith 23875 (F, GH, us). Near Perene Bridge, Killip & Smith 25291 (F, us). Ucayali: Prov. Coronel Portillo, La Divisoria, (as Loreto), /. Schunke V. 10199 (F, MO, us). Madre de Dios: Prov. Manu, Cocha Cashu uplands, Nunez 5841 (MO). XIX. Cystopteris Cystopteris Bernh., Neues J. Bot. (Schrader) 1(2): 26. 1806, nom. conserv. TYPE: Cystopteris fragilis (L.) Bernh. (Polypodium fragile L.). Figure 19. Stem short and decumbent to rather long-creep- ing, bearing scales. Leaves ca. 10-80 cm long, petiole continuous. Lamina 2-pinnate to 3-pin- nate-pinnatifid, glabrous, glandular or slightly pu- bescent, monomorphic. Veins free. Sori round, borne on the veins, not paraphysate, more or less covered by scalelike to half-cup shaped indusia. Spores rather ellipsoidal, monolete, echinate, ver- rucate, slightly rugose, or with inflated processes. Cystopteris is a small genus of about six species, nearly worldwide in distribution, with a single spe- cies, C. fragilis, in the American tropics. The An- dean Cystopteris is highly variable and may even- tually be classified as various taxa representing different cytotypes. At this time it seems best to treat it as a single species. BlasdelFs interpretation of variation in C. fragilis as being due to wide- spread hybridization seems to lack evidence and merit; accordingly the morphological variations are treated as within the natural variation of one species. There are two subgenera, Cystopteris (including C. fragilis) and submenus Acystopteris(Nakai) Bias- dell, the latter often recognized as a genus. Reference BLASDELL, R. F. 1963. A monographic study of the fern genus Cystopteris. Mem. Torrey Bot. Club, 21(4): 1-102. 1. Cystopteris fragilis (L.) Bernh., Neues J. Bot. (Schrader) 1(2):27, t. 2, f. 9. 1806. Figure 19a-b. Polypodium fragile L., Sp. pi. 1091. 1753. TYPE: Plunkenet, Phytographia t. 180, f. 5. 1691. Cystopteris translucens Desv., Mem. Soc. Linn. Paris 6: 264. 1827. TYPE: Peru, (holotype, P, Herb. Desvaux; photos, GH, us). Athyrium fumaroides Presl, Reliq. haenk. 1: 39, /. 6, f. 2. 1825. TYPE: Peru, Haenke (holotype, PR or PRC). Stem scales light to dark brown, usually rather broad. Leaves ca. 10-50 cm long, the petiole usu- ally glabrous, sometimes slightly scaly. Lamina 1-pinnate-pinnatisect to 3-pinnate-pinnatifid, gradually reduced at the apex, essentially glabrous, pinnae nearly sessile to short-stalked, the basal somewhat reduced or not. Veins free. Sori round- ish, borne on the veins, covered by a basally at- tached, scalelike indusium, its apex often some- what elongate. Usually in rocky, locally moist places: on ledges, in crevices of rocks, on rock walls, and on rocky roadside banks, also along streams, on shrubby hillsides, and banks of irrigation ditches, rarely in wet forests or rarely epiphytic, 300—4500 m, Ca- jamarca to Puno. This is a widely distributed spe- cies in Peru, especially at 3000-3500 m; the lowest altitude for the species, 300-700 m, is on the Lo- mas de Atiquipa, Department of Arequipa. Widely distributed in both hemispheres; pri- marily montane in the tropics. Cajamarca: Arriba de Incahuasi, Sagdstegui et al. 12894 (F, GH, HUT). Prov. Contumaza, Lleden, Sagdstegui 9392 (MO). Amazonas: Rio Ventilla, near Molinopampa, Wur- dack 1548 (GH, us). La Libertad: Prov. Santiago de Chu- co, Motil to Shorey, Sounders 891 (F, GH). Prov. Sanchez Carrion, Huayllides, D. Smith 2275 (MO). Ancash: Prov. 92 FIELDIANA: BOTANY FIG. 19. Cystopteris fragilis: a, habit; b, pinnule, abaxial side. (Adapted from Stolze, Ferns & fern allies of Guatemala, 1981.) TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 93 Huaylas, Huascaran National Park, Smith et al. 9786 (F). Prov. Bolognesi, Chiquian, Mostacero et al. 1342 (MO). Huancuco: Chaglla, Macbride 3638 (F, GH, us). Lima: Viso, Macbride & Featherstone 613 (F, us). Prov. Canta, Dist. Huamantanga, Sounders 1149 (f, GH). Prov. Yauyos, Tupe, Cerrate 1071 (USM). Pasco: Prov. Oxa- pampa, Cordillera de Yanachaga, van der Werff et al. 8470 (MO, uc). Junin: Prov. Tarma, 5 km SW of Hu- acapistana, Tryon & Tryon 5432 (GH, us). Huancavelica: Entre Conaica y Manta, Tovar827 (GH). Ayacucho: Nas- ca to Puquio, Correll & Smith PI 56 (GH). Apurimac: Prov. Abancay, Vargas 2296 (uc). Cuzco: Prov. Cuzco, between Cuzco and Sagsaywaman, Stevens 22070 (F, MO). Machu Picchu, Leon 465 (USM). Yucay, Coronado 144 (GH). Arequipa: Lomas de Atiquipa, Coronado 37 (GH, uc, us); ca. 100 km N of Arequipa, Treacy 694 (GH). Puno: 10 km from Puno, Correll & Smith PI 87 (GH). XX. Woodsia Woodsia R. Br., Prodr. 158. obs. IV sub Alsophila. 18 10, as Woodia; also and corrected in Trans. Linn. Soc. London 1 1 : 1 7 1-1 74. 1 8 1 5. TYPE: Woodsia ilvensis (L.) R. Br. (Acrostichum il- vense L.). Figure 20. Stem decumbent to erect, usually small and short-creeping, bearing scales. Leaves ca. 5-50 cm long, the petiole joined in a few (not South Amer- ican) species. Lamina 1 -pinnate or 1 -pinnate-pin- natisect or rarely to 2-pinnate-pinnatifid, glabrous, glandular, scaly, and/or pubescent. Veins free. Sori roundish, borne on the veins, not paraphysate, more or less covered by an indusium surrounding the receptacle that is globose, or of a few large segments, or of a few trichomelike cilia. Spores ellipsoidal to spheroidal, monolete, with short to long winglike ridges and somewhat echinate. Woodsia is mostly a circumboreal genus of about 25 species, with a single species in South America and Peru. Cheilanthes crenata Kunze, Linnaea 9: 84. 1834. TYPE: Peru, (Huanuco), Huanuco, Poeppig in 1830 (holotype, LZ, destroyed; isotype, not locat- ed). W oodsia peruviana Hooker, Sp. fil. 1 : 6 1 , /. 2 IB. 1 844. TYPE: Peru, (Lima), Huamantanga, Mathews 602 (holotype, K; isotype, GH!). Woodsia crenata (Kunze) Hieron., Bot. Jahrb. Syst. 34: 440. 1904. Stem with light brown to dark brown, usually broad scales. Leaves ca. 8-40 cm long, the petiole glabrous or often slightly; scaly, especially at the base. Lamina 1-pinnate-pinnatifid to usually 1 -pinnate-pinnatisect, rarely 2-pinnate-pinnatifid, gradually reduced at the apex, glandular-pubes- cent and pubescent, pinnae sessile or nearly so, the basal ones reduced. Veins free. Sori roundish, borne on a vein, at first covered by a nearly glob- ular indusium that later splits into segments. In rocky places, on cliffs, in crevices of rocks, at the bases of large rocks, or on Inca walls, also on brushy rocky slopes, on pastured hillsides, grassy slopes, or stream banks, rarely in wet woods or on wooded hillsides, 200-4300 m, Lambayeque to Arequipa and Puno. The species grows at 200-600 m on coastal lomas and usually at 3000—4000 m in the Altiplano. Hispaniola; Venezuela and Colombia, south to Argentina and to southeastern Brazil; southern Af- rica. Woodsia motevidensis is a highly variable spe- cies in the size of the lamina and the extent to which the margin is flat or revolute. At high al- titudes, especially in exposed sites, the leaves may be less than 1 0 cm long and the ultimate segments have revolute margins. Especially on lomas, which are mostly cloudy and foggy during the growing season, the leaves may be to 40 cm long and the ultimate segments of the lamina are expanded and flat. Reference BROWN, D. F. M. 1964. A monographic study of the fern genus Woodsia. Beih. Nova Hed- wigia, 16: 1-154. 1 . Woodsia montevidensis (Sprengel) Hieron., Bot. Jahrb. Syst. 22: 363. 1896. Figure 20a-d. Dicksonia montevidensis Sprengel, Syst. veg. ed. 16, 4: 122. 1827. TYPE: Uruguay, "Monte Video," Sello (a specimen at B is authentic). Lambayeque: 42 km from Olmos on road to Jaen, Correll & Smith P816 (GH). Cajamarca: Prov. Contu- maza, arriba de Contumaza, Lopez et al. 37 13 (F, GH). Prov. Contumaza, alrededores de Guzmango, Sagdste- gui 9675 (MO). La Libertad: Lomas de Viru, Coronado 284 (GH, uc), Lopez et al. 3638 (GH). Ancash: Lomas de Mongon, Coronado 299 (GH). Cerro al E de Chiquian, Ferreyra 7343 (GH). Huanuco: Piedra Grande, Macbride 3671 (F, GH, us). Muna, Woytkowski 5252 (GH, MO). Lima: Loma de Amancaes, Tryon & Tryon 5215 (F, GH, us), Coronado 12 (MO). Matucana, Macbride & Feath- erstone 427 (F, us). Junin: Near Huancayo, Killip & Smith 22127 (GH, us). Huacapistana, Cerrate 2884 (GH). Huan- cayo, Soukup 3549 (MO). Huancavelica: Entre Colca- bamba y Paucarbamba, Tovar 2008 (GH). Ayacucho: Be- 94 FIELDIANA: BOTANY a \ FIG. 20. Woodsia montevidensis: a, habit; b, stem scale; c, pinna segment, abaxial side. (From Bryan 375, F.) tween Huanta and Rio Apurimac, Killip & Smith 22485 Amparaes, Nunez 6642, 6649 (MO). Slopes of Sicuani, (GH, us). Apurimac: Prov. Aymaraes, Dist. Cotarosi, Vargas 9831 (F, GH, uc). Arequipa: Arequipa, Pennell Saunders 775 (GH). Cuzco: Machu Picchu, Tryon & Tryon 13197 (GH, us). Puno: Puno, Graja Salcedo, Mexia 7781 5402 (GH, us), Coronado 96 (GH, uc, us). Prov. Calca, (GH, uc, us). Chucuito, Coronado 165 (GH, uc). TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 95 XXI. Oleandra Oleandra Cav., Anales Hist. Nat. 1:115. 1799. TYPE: Oleandra neriiformis Cav. Figure 21. Stem erect and branching, or climbing and long- creeping, slender, to 5 m long, bearing scales. Leaves ca. 10-50 cm long, petiole articulate. Lam- ina simple, entire, glabrate, pubescent, and/or scaly, monomorphic or very rarely dimorphic. Veins free. Sori roundish, borne on the veins, not paraphy- sate, covered by a reniform to peltate indusium. Spores rather spheroidal, monolete, more or less prominently ridged and echinate. Oleandra is a pantropical genus of about 35 spe- cies. There are at least six species in America and three in Peru. It is one of the few ferns that have some species with an erect, branched stem that results in a shrublike habit. The petiole is articulate and the portion of it below the joint is called a phyllopodium. Reference MAXON, W. R. 1914. The American species of Oleandra. Contr. U.S. Natl. Herb., 17: 392-398. Key to Species of Oleandra a. Stem widely creeping, its scales spreading; phyllopodia mostly 5-30 mm long, naked, like the petiole above the joint; lamina and indusia glabrous 1. O. articulata a. Stem ascending, erect or climbing, its scales closely appressed b b. Lamina glabrous or with some scales, mostly 1-2 cm wide, rarely some to 2.5 cm or less than 1 cm wide; phyllopodia mostly 10-20 cm long, or some as short as 3 mm, or as long as 30 mm, naked, like the petiole above the joint; indusia glabrous 2. O. lehmannii b. Lamina pubescent, at least in part, especially on or near the costa, mostly 3-3.5 cm wide, rarely some 2 cm or more than 3.5 cm wide; phyllopodia 1-3 mm long, or some to 6 mm long, at first scaly, like the stem; indusia slightly pubescent on the surface and with ciliate margins 3. O. pilosa 1. Oleandra articulata (Sw.) Presl, Tent, pterid. 78. 1836. Figure 21a-b. Aspidium articulation Sw., J. Bot. (Schrader) 1800(2): 38. 1 802. TYPE: Martinique, Plumier, Traite foug. Amer. t. 136. Aspidium nodosum Willd., Sp. pi. ed. 4, 5: 21 1. 1810, nom. superfl. for Aspidium articulatum and with the same type. Oleandra nodosa Presl, Tent, pterid. 78. 1836, nom. nov. for Aspidium nodosum Willd. and with the same type. Stem widely creeping, with spreading, usually widely spreading, scales. Phyllopodia 5-30 mm long, slender, without scales. Lamina glabrous, ca. 15-40 cm long and 2.5-8 cm wide. Indusia gla- brous. Epiphytic in dense forests or cloud forests, 1 330 to ca. 1 700 m, Huanuco, pasco, and Junin. Guatemala to Panama; West Indies; northern South America south to Peru and northern Bra- zil; Cocos and Galapagos Islands. Rarely the scales of the stem are mostly only slightly spreading, rather than widely spreading. Maxon chose Plumier t. 136, the only element included in the original publication of Aspidium articulatum Sw., as the type of the name. He in- vestigated material from Mauritius, included in Aspidium articulatum later by Swartz (Syn. fil. 42, 236. 1805) in the Swartz herbarium, Stockholm, and decided that it could not be type material of Aspidium articulatum. Morton (Amer. Fern J. 58: 105-107. 1968) reached a contrary conclusion and applied the name Aspidium nodosum to the Amer- ican species and Aspidium articulatum to the spe- cies of Africa and adjacent islands. Joncheere (Taxon 18: 538-541. 1969) argued persuasively that Maxon's original typification was correct and we confidently agree with his conclusion. The cor- rect name for the African species is Oleandra dis- tenta Kunze (Pic.-Ser., Webbia 20: 755. 1965). Huanuco: Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13243 (GH). Pasco: Pichis Train, Enenas (as Junin), Killip & Smith 25756 (F, NY, us). Prov. Oxa- 96 FIELDIANA: BOTANY rid •• t* i , s.mm, 2cm FIG. 21. Oleandra articulata: a, habit; b, portion of lamina, abaxial side. Oleandra lehmannii: c, stem and phyllopodium. (a, b from Ollgaard & Balslev 9085, Ecuador, F; c from D. Smith 5395, F.) pampa, Cordillera San Matias, Leon 313 (GH, USM). Junin: Above San Ramon, Killip & Smith 24878 (NY, us). 2. Oleandra lehmannii Maxon, Contr. U.S. Natl. Herb. 17:395. 1 9 14. TYPE: Colombia, Antioquia, above Amalfi, Lehmann XLII (holotype, us!; iso- type, K!; photo, GH). Figure 21c. Stem ascending to erect and climbing, with closely appressed scales. Phyllopodia (3-)10- 20(-30) mm long, mostly slender, without scales. Lamina glabrous or with some scales, ca. 10-30 cm long and l-2(-2.5) cm, rarely less than 1 cm, broad. Indusia glabrous. Epiphytic or rarely terrestrial in rich organic matter, montane rainforests, elfin forests, rarely in Sphagnum bogs, 1480-2500 m, Huanuco and Pas- co. Colombia south to Peru; northern Brazil. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 97 Huanuco: Rio Llullapichis watershed, ascent of Cerros del Sira, Dudley 13306B, 13453, 13489 (OH), 1 3554 (GH, us). Pasco: Pichis Trail, Dos de Mayo (as Junin), Killip & Smith 25870 (F, GH, NY, us). Prov. Oxapampa, 20 km W of Oxapampa, D. Smith 5395 (GH). Chontabamba, Ledn et al. 968 (F, GH). capata, de Lantreppe (NY). Prov. Paucartambo, San Pe- dro, Vargas 6801 (uc, us). Puno: Tatanara, Lechler2539 (B, K). Comments 3. Oleandra pilosa Hooker, Gen. fil. t. 45B. 1840. TYPE: "British Guiana," Berbice, Schom- burgk 416 (holotype, K!; photo, GH). Aspidium pendulum Splitg. Tijdschr. Natuurl. Gesch. Physiol. 7: 412, 1840, not Raddi, 1819. SYN- TYPES: Surinam, Berlijn plantation, Splitgerber (L; photo, GH); Schomburgk 416 (K, L; photo GH ofK). Oleandra micans Kunze, Bot. Zeit. (Berlin) 9: 346. 1851. TYPE: Peru, (San Martin), Mission To- cache, (sterile), PoeppigM. Aug. 1830(Z)/ar. 7955) (holotype, LZ, destroyed). Data from Linnaea 9: 89. 1834. Tracing in B! Herb. Mett. of "Oleandra micans Kunze, Peruvia Poeppig." Stem ascending to erect and climbing, with closely appressed scales. Phyllopodia l-3(-6) mm long, stout, at first scaly. Lamina pubescent, es- pecially on or near the costa, ca. 10-40 cm long, and (2-)3-3.5(-6) cm broad. Indusia slightly pu- bescent on the surface and with ciliate margins. Epiphytic in forests, 325-1000 m, San Martin to Puno. Trinidad and northern South America, south to Peru. Oleandra pilosa and O. lehmannii are the only species in Peru with appressed scales on the stem. The variation in these species makes it impossible to determine the identity of the type of Oleandra micans from the original description and the trac- ing at B. No isotype has been seen. However, col- lections of O. pilosa in Peru are from 325-1000 m, while those of O. lehmannii are from 1480- 2500 m. Since Tocache is at an altitude of about 500 m, O. micans is treated as a synonym of O. pilosa. Huanuco: Tingo Maria (as San Martin), Allard 21215, 21999 (GH). Prov. Leoncio Prado, E of Tingo Maria, J. Schunke V. 5194 (GH, MO, us). Pasco: Prov. Oxapampa, Palcazu valley, D. Smith 3807 (GH). Cuzco: Rio Mar- Oleandra hirta Brack., U.S. Expl. exped. 16 (fil.): 214. 1854. Atlas, t. 29. 1855. TYPE: Brazil, Organ Mountains, U.S. Expl. Exped. (us). This species is found primarily in Brazil, but it has been collected in Bolivia (Tate 1152, NY) and may occur in southern Peru. The stem and scales are like those of O. articulata but the lamina is pubescent and the indusia are usually slightly pu- bescent on the surface or somewhat ciliate. The shape of the lamina is variable, as it is in O. pilosa and other species of the genus. It may be gradually attenuate basally and more or less acuminate api- cally (type of O. hirta), or it may vary to narrowly cuneate to broadly rounded basally and acute to caudate apically. XXII. Bolbitis Bolbitis Schott, Gen. fil. 1. 13 (fasc. 3). 1835. TYPE: Bolbitis serratifolia (Kaulf.) Schott (Acrosti- chum serratifolium Kaulf.). Figure 22. Plants terrestrial to epipetric, sometimes be- coming hemiepiphytic. Stem short- to long-creep- ing, or scandent well up tree trunks, slender to rather stout, moderately to densely provided with light brown to blackish, subclathrate scales and with few to many fibrous roots. Leaves strongly to weakly dimorphic, ca. 25 cm to 2 m long, ap- proximate to widely spaced, petiole continuous with the stem. Lamina simple and entire to 1 -pinnate-pinnatifid, sometimes imparipinnate, the pinnae not articulate to the rachis, glabrate to somewhat scaly, the rachis or apical segment often bearing a proliferous bud. Veins areolate, with or without included free veinlets, or (in a few species not in Peru) free. Sporangia commonly borne over the abaxial surface of the segment, lacking pa- FIG. 22. Bolbitis serrata: a, habit; b, portion of pinna, abaxial side. Bolbitis serratifolia: c, portion of pinna, abaxial side. B. lindigii: d: portion of pinna, abaxial side, (a, b from Killip & Smith 28495, us; c from C. Schunke 161, F, d from Killip & Smith 25184, F.) 98 FIELDIANA: BOTANY TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 99 raphyses. Indusium lacking. Spores monolete, more or less ellipsoidal. Bolbitis is a pantropical genus of 44 species, in- cluding 1 4 in America. It is allied to Lomagramma and Lomariopsis, especially in the large, dimor- phic leaves, which are borne on a stem that some- times climbs well up the trunks of trees, and the acrostichoid pattern of sporangia. Venation is typ- ically areolate, although veins of several Asian spe- cies are free. It differs from both allied genera in that the pinnae are not articulate to the rachis. Species in Peru can be further distinguished from Lomariopsis by their anastomosing veins. This treatment is adapted in large part from the mono- graph of Hennipman (1977). Reference HENNIPMAN, E. 1977. A monograph of the fern genus Bolbitis (Lomariopsidaceae). Leiden Bot., ser., 2: 1-331. Key to Species of Bolbitis a. Sterile lamina pinnate and gradually reduced to a nonconform, usually pinnatifid, apex 1 . B. aliena a. Sterile lamina simple, or pinnate and abruptly reduced to a discrete (usually conform) apical segment b b. Free veinlets abundant, spreading in various directions within the areoles c c. Sterile lamina lacking proliferous buds; adult sterile lamina commonly with an apical segment and 3-8 pinnae (very rarely simple or with 2 pinnae) 4. B. nicotianifolia c. Sterile lamina either with a subterminal or an axillary proliferous bud; adult sterile lamina simple, subternate or ternate d d. A proliferous bud borne in the pinna axils (or on a pinna stalk); sterile lamina ternate or subternate; pinnae often to 20 cm long and 12 cm broad 5. B. oligarchica d. A proliferous bud borne at or near the lamina apex; sterile leaf usually simple, occasionally subternate; pinnae (when present) to 8 cm long and 4 cm broad 7. B. pandurifolia b. Free veinlets lacking within the areoles or, when present, directed either distally (toward the pinna margin) or proximally (toward the costa), but not or rarely in various directions e e. Pinnae with only costal areoles (or rarely 1 or 2 more produced beyond the costal one); lateral pinnae commonly 4-6, less than 2 cm broad; rachis distinctly alate throughout 2. B. serrata e. Pinnae amply areolate beyond the costal areole; lateral pinnae of mature leaves 8-36, 2-5 cm broad; rachis nonalate, or scarcely alate toward the apex f f. Areoles mostly isomorphic and slightly elongated, free included veinlets lacking or rare; veins commonly distinct and abaxially raised; stem scales mostly 8-20 mm long and linear (sometimes caducous) 6. B. lindigii f. Areoles mostly heteromorphic and rather elongated, free included veinlets lacking to fre- quent and directed distally; secondary and tertiary veins becoming indistinct between pri- mary ones, not or scarcely raised; stem scales 2-5 mm long, ovate to broadly lanceolate . 3. B. serratifolia 1. Bolbitis aliena (Sw.) Alston, Bull. Misc. In- form. 1932: 310. Acrostichum alienum Sw., J. Bot. (Schrader) 1800 (2): 13. 1802. TYPE: Collector unknown, Jamaica (holotype, s, Herb. Swartz; isotype, s, Herb. Swartz). Gynnopteris aliena (Sw.) Presl, Tent, pterid. 244. 1836. Leptochilus alienus (Sw.) C. Chr., Bot. Tidsskr. 26: 285. 1904. Plants terrestrial or epipetric. Stem long-creep- ing. Sterile leaves well spaced, pinnate or deeply pinnatisect, 25-90 cm long, 14—30 cm broad. Lamina broadest at or near the base, reduced grad- ually to a deltate, pinnatifid apex. Rachis broadly alate throughout, or nonalate toward base, lacking proliferous buds. Pinnae commonly 3-6 pairs, ad- nate, or proximal ones stalked, entire to shallowly lobed, or basal pair with a greatly expanded basal 100 FIELDIANA: BOTANY lobe. Veins copiously areolate, lacking included free veinlets. Fertile leaves commonly smaller than the sterile, with pinnae less deeply lobed. In rain forests and wooded ravines, creeping on earth or rocks or rocky cliffs, 100-1000 m, Ama- zonas to Pasco and Ucayali. Southern Mexico; Honduras; Nicaragua; West Indies; Venezuela and Colombia, southward to Bolivia. This should not be confused with other species in Peru, as it is the only one lacking a discrete apical segment. It is sometimes confused with B. portoricensis (Sprengel) Hennipman, a similar species with a pinnatifid apex whose range extends to Ecuador. However, the areoles of B. portori- censis have included free veinlets, whereas these are lacking in B. aliena. Amazonas: Prov. Bagua, Rio Maranon opposite Que- brada Mirana, Wurdack 2033 (GH, us). San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Schunke V. 3891 (F, GH, MO, us). Huanuco: Prov. Huanuco, Tingo Maria, Tryon & Tryon 5281 (F, GH, uc, us). Pasco: Prov. Oxa- pampa, Palcazu, Rio Alto Iscozacin, Foster & d'Achille 10129 (F). Ucayali: Km 86 on Pucallpa-Tingo Maria road, along trail to Arboretum of Bosque von Humboldt Experimental Station, D. Smith 1225 (MO). 2. Bolbitis serrata (Kuhn) C. Chr., Index fil. suppl. 3: 50. 1934. Figure 22a-b. Chrysodium serratum Kuhn, Linnaea 36: 63. 1869. TYPE: Peru, San Martin, near Tarapoto, Spruce 4123 (holotype, B; isotypes, BM!, GH!, K!, NY, p, us!; photos, F & GH of us). Acrostichwn serratum (Kuhn) Baker, Syn. fil., ed. 2: 524. 1874, nom. illeg. (not Poiret, 1810). Leptochilus serratus (Kuhn) C. Chr., Bot. Tidsskr. 26: 289. 1904. Plants terrestrial or epipetric. Stem long-creep- ing, slender, to 5 mm thick. Sterile leaves lacking a proliferous bud, subdistant, pinnate, to 40 cm long and 1 0 cm broad. Lamina with 4-6(-8) pin- nae and a conform apical segment, this narrow- lanceolate or narrow-elliptic and with a narrowly cuneate base. Rachis distinctly alate throughout, the wing with a cartilaginous margin. Pinnae 0.8- 2(-2.3) cm broad, short-stalked, the margin entire to crenate-serrate, the base narrow-cuneate. Veins areolate only along the costa, or rarely with 1 or 2 more areoles beyond the costal one, the areoles lacking included free veinlets. Fertile leaves com- monly longer than the sterile, the lamina smaller, but the petiole much longer. In rain forests, creeping among wet rocks or on soil, 100-550 m, San Martin and Loreto. Colombia; Ecuador; Peru. The smaller leaves of B. serrata, with fewer pin- nae and more slender stems, usually distinguish it from B. lindigii and B. serratifolia, but the vena- tion pattern is a more diagnostic character. In most pinnae there is merely a costal areole, but some- times one or two areoles may be produced beyond this. None of the areoles bear included free vein- lets. The veins of B. lindigii regularly anastomose, are commonly distinct and raised abaxially, and the areoles beyond the costal one are nearly iso- morphic (pentagonal or hexagonal, not greatly elongated), they lack included free veinlets, or the rare ones are minute and often directed toward the costa. Veins of B. serratifolia rather freely anas- tomose, but are not or scarcely raised, and sec- ondary and tertiary veins become indistinct be- tween the primary ones. The areoles are quite irregular in shape and many are greatly elongated. Free included veinlets may be frequent to sparse or lacking, but when present they are directed to- ward the pinna margin. San Martin: Tarapoto, near road to Yurimaguas, Mar- tin & Plowman 1815 (GH, us). Loreto: Balsapuerto, lower Rio Huallaga basin, Killip & Smith 28495 (F, GH, NY, us). Prov. Alto Amazonas, above Pongo de Manseriche, Wurdack 2110 (F, GH, NY, s, u, uc, us). 3. Bolbitis serratifolia (Kaulf.) Schott, Gen. fil. t. 13. 1835. Figure 22c. Acrostichum serratifolia Kaulf., Enum. fil. 66. 1824. TYPE: Brazil, Rio de Janeiro (B?, LE?). Poecilopteris crenata Presl, Epim. bot. 174. 1851. TYPE: Brazil, near Rio de Janeiro, Mikan (ho- lotype, PR or PRC). Leptochilus serratifolius (Kaulf.) C. Chr., Bot. Tidsskr. 26: 289. 1904. Leptochilus crenatus (Presl) C. Chr., Index fil., suppl. 1: 48. 1913. Bolbitis crenata (Presl) C. Chr., Index fil., suppl. 3: 47. 1934. Plants terrestrial, epipetric, or hemiepiphytic. Stem short- to long-creeing, sometimes low-scan- dent, to 1 .5 cm thick, sparsely provided with ovate or lanceolate scales 2-5 mm long. Sterile leaves occasionally with a proliferous bud (or primor- dium) subapically on the apical segment (or pin- nae), approximate on the stem, pinnate, to 1.2 m long and 30 cm broad, the rachis, costae and veins sparsely provided abaxially with castaneous, fila- mentous scales. Lamina with 8-36 pinnae and a TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 101 conform or subconform apical segment, this broadly lanceolate and with a broadly cuneate or rounded base. Rachis not alate, or only scarcely so near the apex. Pinnae 2—4 cm broad, short- stalked, the margin entire to crenate-serrate. Pri- mary veins distinct, but secondary and tertiary ones usually indistinct between the primary ones, not or scarcely raised, amply anastomosing, with a broad costal areole and many heteromorphic and often elongated areoles beyond, free included vein- lets lacking, or few to many and then directed distally (toward pinna margin). Fertile leaves sub- equal to the sterile but the petiole proportionately longer and the pinnae much shorter and narrower. In dense forests or wooded ravines, on moist ground, often low-scandent on bases of tree trunks, occasionally on rocks, 150-1600 m, San Martin and Loreto south to Ayacucho and Cuzco. Southern Mexico (Chiapas); Costa Rica; Pana- ma; Surinam to Colombia, southward to Brazil and Argentina. This species bears a superficial resemblance to B. lindigii. See treatments of the latter and B. ser- rata for detailed comparison. San Martin: Near Tarapoto, Spruce 4736 (BR, G, GH, K., P). Loreto: Puerto Arturo, below Yurimaguas, Killip & Smith 27797 (F, us). Huanuco: Prov. Huanuco, Tingo Maria, Tryon & Tryon 5234 (F, GH, us). Junin: Schunke Hacienda above San Ramon, C. Schunke A-215 (GH, us). Ayacucho: Estrella, between Huanta and Rio Apu- rimac, Killip & Smith 22626 (F, us). Cuzco: Prov. La Convention, Cordillera Vilcabamba, Rio Klause, Dud- ley 10187 (GH). 4. Bolbitis nicotianifolia (Sw.) Alston, Bull. Misc. Inform. 1932: 310. Acrostichum nicotianifolium Sw., Syn. fil. 13. 199. 1806. TYPE: Virgin Islands, St. Thomas, Ven- tentat (holotype, s). Gymnopteris nicotianifolia (Sw.) Presl, Tent, pterid. 244. 1836. Leptochilus nicotianifolius (Sw.) C. Chr., Bot. Tidsskr. 26: 285. 1904. Leptochilus killipii Maxon, Amer. Fern J. 21: 138. 1931. TYPE: Panama, above Juan Diaz, Killip 2778 (holotype, us!; isotypes, B, P, s). Bolbitis killipii '(Maxon) Lell., Fern Gaz. 11: 107. 1975. Plants terrestrial or hemiepiphytic. Stem short- to long-creeping or scandent, moderately provided with lanceolate, brown to blackish scales 3-5 mm long. Sterile leaves lacking a proliferous bud, ap- proximate to subdistant on the stem, to 1.2m long and 40 cm broad. Lamina pinnate (rarely simple or subternate), with 3-8 pinnae and a conform or somewhat enlarged apical segment, this elliptic and with a cuneate or shortly decurrent base. Rachis nonalate. Pinnae commonly to 28 cm long and 8 cm broad, subsessile or short-stalked, the margin essentially entire, the base cuneate or often obtuse basiscopically. Veins copiously areolate, with many free included veinlets spreading in various direc- tions. Fertile leaves pinnate (rarely simple), sim- ilar to sterile ones, but with fewer and smaller pinnae. In rain forests or in wooded ravines, on wet ground or on bases of tree trunks, 100-540 m, San Martin, Loreto, Huanuco, Cuzco, Madre de Dios, and Puno. Guatemala to Panama; West Indies; the Guia- nas to Colombia and south to Peru. The rare specimens with simple leaves or only with two pinnae might be confused with B. oli- garchica or B. pandurifolia. See treatments of these two species for further discussion. Bolbitis nicotianifolia and B. oligarchica are very closely related and may hybridize, as suggested by Hennipman (1977). There is a specimen from Ta- rapoto, San Martin, LI. Williams 6064 (us), con- taining a portion of a sterile leaf which seems to be intermediate between the two species. Although there are nine pinnae (typical of B. nicotianifolia), these are more strongly produced basiscopically and only 1 1/2 times as long as broad (characteristic of B. oligarchica). Furthermore, there is a prolif- erous bud in the axil of a subapical pinna, a con- dition common in the latter species, but not in the former. Both species have been found in San Mar- tin. San Martin: "In monte Guayrapurima," Spruce 4636 (K). Loreto: Salinas, Rio Mazan, Schunke 372 (F, GH, uc, us). Huanuco: Prov. Pachitea, Comunidad Nativa Santa Marta, D. Smith 1244 (MO). Cuzco: Prov. Paucartambo, Hacienda Villa Carmen, Vargas 14680 (GH). Madre de Dios: Rio Manu, Cocha Cashu Station, Foster & Ter- borgh 6622 (F). Puno: Prov. Carabaya, Hacienda Pal- mera, Vargas 16131 (GH). 5. Bolbitis oligarchica (Baker) Hennipman, Amer. Fern J. 65: 30. 1975. Acrostichum oligarchicum Baker, Syn. fil. 418. 1868. LECTOTYPE (designated by Maxon, Amer. Fern J. 21: 139. 1931): Peru, San Martin, Mt. Guay- rapurima, near Tarapoto, Spruce 4737 (K; photo, us). 102 FIELDIANA: BOTANY Leptochilus oligarchicus (Baker) C. Chr., Bot. Tidsskr. 26: 285. 1904. HypoderrisstuebeliiHieron., Hedwigia46: 323. 1907. TYPE: Ecuador, Tungurahua, Stiibel 906 (B). Leptochilus bradeorum Rosenst., Repert. Spec. Nov. Regni Veg. 9: 70. 1910. TYPE: Costa Rica, La Palma, Brade& Brade(ho\otype, B?; isotypes, NY, uc!). Leptochilus stuebelii (Hieron.) Maxon, Proc. Biol. Soc. Wash. 46: 142. 1933. Bolbitis bradeorum (Rosenst.) C. Chr., Index fil. suppl. 3: 47. 1934. Bolbitis stuebelii (Hieron.) C. Chr., Index fil. suppl. 3: 50. 1934. Plants terrestrial. Stem stout, short-creeping, moderately provided with lanceolate scales 3-5 mm long. Sterile leaves approximate, often crowd- ed, ternate, 30-90 cm long. Lamina with a pair of subopposite pinnae and a greatly enlarged apical segment, this broadly elliptic to rhomboid, often to 40 cm long and 30 cm broad, the base cuneate to short-decurrent. Rachis bearing at the axils (or on stalks) of pinnae a proliferous bud. Pinnae often to 20 cm long and 1 2 cm broad, subsessile or short- stalked, essentially entire, the base cuneate and usually more strongly produced basiscopically. Veins copiously areolate, with many included free veinlets spreading in various directions. Fertile leaves always ternate, much smaller than the ster- ile, but with longer petioles, the pinnae with longer stalks and more widely spaced from the apical segment. In rainforests, on wet, often rocky, ground, 400- 1 700 m, Amazonas to Junin and Ucayali. Costa Rica; Colombia to Bolivia. Three of the species of Bolbitis occurring in Peru have free veinlets included within the areoles and spreading in various directions: B. nicotianifolia, B. oligarchica, and B. pandurifolia. The other spe- cies in Peru either lack free veinlets or have them directed always toward the pinna margin, or al- ways toward the costa. It is interesting that the leaves of these three species tend to consist of a large terminal segment subtended by a pair of somewhat to greatly reduced pinnae. This condi- tion is found but rarely in B. nicotianifolia, oc- casionally in B. pandurifolia, and always in B. oli- garchica. Amazonas: Prov. Bagua, valley of Rio Maranon above Cascadas de Mayasi, Wurdack 1896 (F, GH, uc, us). San Martin: Valley of Rio H uallaga, road between Tocache Nuevo and Juanjui, Croat 58073 (MO). Huanuco: Prov. Leoncio Prado, Dist. Rupa Rupa, near Cerro Quemado, Schunke V. 10161 (F, MO, us). Pasco (as Junin): Pichis Trail, Santa Rosa, Killip & Smith 26174 (GH, us). Junin: "Prov. Satipa-Pichanaki" (Dist. Pichanaqui in Prov. Chanchamayo?), Rodal del Projecto Peruano-Aleman, Leon 224 (USM). Ucayali (as Loreto): Prov. Coronel Por- tillo, Dist. Padre Abad, Boqueron de Padre Abad, Schunke V. 3073 (F, GH, us). 6. Bolbitis lindigii (Mett.) C. Chr., Index fil. suppl. 3: 48. 1934. Figure 22d. Chrysodium lindigii Mett., Ann. Sci. Nat. Bot., ser. 5, 2: 205. 1864. TYPE: Colombia, Boyaca, Muzo, Lindig 258 (holotype, B; isotypes, BM, K, p). Acrostichum lindigii (Mett.) Baker, Syn. fil. 423. 1 868, nom, illeg. (not Karsten, 1858). Leptochilus lindigii (Mett.) C. Chr., Bot. Tidsskr. 26: 290. 1904. Plants hemiepiphytic. Stem long-creeping, scan- dent high on tree trunks, to 2 cm thick, abundantly provided with linear scales 8-20 mm long (these sometimes caducous). Sterile leaves lacking a pro- liferous bud, commonly widely spaced, pinnate, to 1 m long and 30 cm broad. Lamina with 16- 36 pinnae and a conform or subconform apical segment, this broadly lanceolate and with a cu- neate base. Rachis not alate, or only scarcely so near the apex. Pinnae 2-5 cm broad, short-stalked, the margin crenate-serrate to crenately lobed. Veins commonly distinct and raised abaxially, regularly anastomosing, with a broad costal areole and many nearly isomorphic areoles beyond, free included veinlets lacking or very rare and then usually di- rected proximally (toward the costa). Fertile leaves shorter than the sterile and the pinnae much short- er and narrower. Hemiepiphytic, scandent to 23 m on tree trunks, in deep rain forests, 100-1200 m, San Martin and Loreto to Cuzco and Madre de Dios. Costa Rica southward to Bolivia; Brazil. This species, with its large leaves, numerous pinnae, and a conform apical segment is some- times confused with B. serratifolia. However, the latter is only low-scandent (if at all), whereas stems of B. lindigii often climb on tree trunks, sometimes to a height of 23 m. The two also are easily sep- arated by the venation pattern, which is discussed in detail under B. serrata. San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, Puerto Pizana, Schunke V. 6897 (MO, uc). Lore- to: Gamitanacocha, Rio Mazan, J. Schunke 333 (F, GH, NY, s, us, USM). Huanuco: Prov. Pachitea, Dist. Puerto Inca, Bosque National de Iparia, Schunke V. 2846 (F). Pasco: Oxapampa, Palcazu Valley, Iscozacin, Foster et TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 103 2 mm FIG. 23. Lomariopsis fendleri: a, habit. Lomariopsis japurensis: b, portion of rachis and pinna base, adaxial side. Lomariopsis nigropaleata: c, portion of rachis and pinna base, abaxial side; d, scale from petiole base, (a from Killip & Smith 27792, F, b from LI. Williams 5262, F, c, d from Schunke V. 2843, F.) 104 FIELDIANA: BOTANY al. 7886 (F, MO). Junin: Chanchamayo Valley, C. Schunke 846, 854 (F). Cuzco: Prov. La Convention, near Rio Mapitunuari and Hacienda Luisiana, Dudley 11511, 11515 (GH). Madre de Dios: Tambopata, Albergue "Cuz- co Amazonico," Leon 827 (F, MO). (as Junin): Pichis trail, San Nicolas, Killip & Smith 26039 (NY, us). 7. Bolbitis pandurifolia (Hooker) C. Chr. Index fil. suppl. 3: 49. 1934. Gymnopteris pandurifolia Hooker, Sec. cent, ferns t. 87. 1861. TYPE: Peru, San Martin, Mt. Guay- rapurima near Tarapoto, Spruce 4741 (holotype, K; isotypes, BM, BR, P; photos, F & us of BM). Acrostichum pandurifolium (Hooker) Hooker, Sp. fil. 5: 271. 1864. Leptochilus pandurifolius (Hooker) C. Chr., Hot. Tidsskr. 26: 286. 1904. Plants terrestrial or low-scandent. Stem stout, short-creeping, moderately provided with lanceo- late, attenuate scales 2—4 cm long. Sterile leaves approximate and often crowded, simple or sub- ternate, to 60 cm long and 20 cm broad, essentially glabrous, bearing a proliferous bud at or near the apex. Lamina simple, broadly oblong-lanceolate, or often with a pair of greatly reduced, subopposite pinnae, in simple leaves the base truncate to cor- date, and in ternate leaves the large apical segment usually cuneate at base. Pinnae (when present) to about 8 cm long and 4 cm broad, subsessile or short-stalked, elliptic, essentially entire, the base cuneate. Veins copiously areolate, with included free veinlets spreading in various directions. Fer- tile leaves about as long as the sterile, but petiole proportionately much longer and lamina shorter. In dense rain forests, on wet ground or occa- sionally scandent on bases of tree trunks, 450- 1 100 m, Amazonas, San Martin, and Pasco. Ecuador and Peru. The lamina of this species is frequently simple, or sometimes it consists of a large apical segment subtended by a pair of much-reduced pinnae. Rarely, leaves of B. nicotianifolia may be simple and then the base of the lamina is cuneate to de- current. In contrast, the simple lamina of B. pan- durifolia is truncate to subcordate at the base. Amazonas: Prov. Bagua, valley of Rio Maranon above Cascadas de Mayasi, Wurdack 1897 (GH, NY, us). Pasco: Key to Species of Lomariopsis a. Sterile pinnae of adult leaves 20-30 pairs, larger ones 1-1 .8 cm broad; fertile pinnae 1 .5-4 mm broad; rachis broadly or narrowly alate throughout, at least in the distal half 1 . L. fendleri XXIII. Lomariopsis Lomariopsis Fee, Mem. Foug. 2: 10. 1845. TYPE: Lomariopsis sorbifolia (L.) Fee (Acrostichum sorbifolium L.). Figure 23. Plants terrestrial (occasionally epipetric), usu- ally becoming scandent-epiphytic. Stem long- creeping or scandent, to ca. 15m long, slender to stout, bearing scales and few to many fibrous roots. Leaves moderately to strongly dimorphic (fertile pinnae narrower than the sterile), ca. 0.2-1.5 m long, closely to (usually) widely spaced on the stem. Lamina 1 -pinnate, imparipinnate, glabrate to slightly and minutely scaly, the pinnae entire to crenate-serrate (or on juvenile leaves deeply sin- uate or dentate), articulate to the rachis, the apical segment continuous (or in three West Indian spe- cies articulate). Veins free, simple or forked. Spo- rangia borne over the abaxial surface of the pin- nae. Indusia and paraphyses lacking. Spores monolete, ellipsoidal. The genus Lomariopsis is essentially pantropi- cal, containing about 45 species, with perhaps 1 5 in the Neotropics. It is characterized by the dor- siventral and usually scandent stem, the 1 -pinnate, dimorphic lamina, and the articulate, free-veined pinnae with acrostichoid sori. Its closest allies in Peru, with which it might be confused, are Loma- gramma and Bolbitis. References HOLTTUM, R. E. 1932. On Stenochlaena, Lo- mariopsis and Teratophyllum in the Malayan region. Card. Bull. Straits Settlem., 5: 245-313. HOLTTUM, R. E. 1940. New species of Loma- riopsis. Bull. Misc. Inform., 1939: 613-628. UNDERWOOD, L. M. 1907. American ferns, 7: The American species of Stenochlaena, Bull. Torrey Bot. Club, 38: 591-603. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 105 Sterile pinnae of adult leaves 2-12 pairs (or leaves rarely simple in #2 & #3), larger ones 3-9 cm broad; fertile pinnae 7-20 mm broad; rachis narrowly alate only near the apex b b. Scales of stem and petiole base moderate to abundant, 3-12 mm long, linear to lanceolate, essentially concolorous, tawny to brownish black c c. Larger sterile pinnae 3-5 cm broad, sessile or short-stalked (0-2 mm), apex acuminate, base commonly obtuse to broadly cuneate; larger fertile pinnae 8-16 mm broad, sessile or subsessile 2. L. japurensis c. Larger sterile pinnae 6-9 cm broad, on stalks (3-)4-15 mm long, apex acute to subacute, base narrow-cuneate; larger fertile pinnae 1 6-20 mm broad, the stalks 3-5 mm long 3. L. latipinna b. Scales of stem and petiole base sparse and scattered, l-2(-3) mm long, ovate or broadly lanceolate, mostly bicolorous, lustrous black with a ferruginous border 4. L. nigropaleata 1 . Lomariopsis fendleri D. C. Eaton, Mem. Amer. Acad. Arts, n.s. 8: 195. 1860. TYPE: Vene- zuela ("prope Coloniam Tovar"), Fendler335 (holotype, YU; isotypes, OH!, K!). Figure 23a. Lomariopsis vestita Fourn., Bull. Soc. Bot. France 19: 250. 1872. TYPE: Nicaragua, Chontales, Levy 476 (holotype, p; isotype, K!). Stenochlaena fendleri (D. C. Eaton) Underw., Bull. Torrey Bot. Club 33: 595. 1907. Stenochlaena vestita (Fourn.) Underw., Bull. Torrey Bot. Club 33: 600. 1907. Stem stout, provided with stramineous to taw- ny, lanceolate, attenuate scales, these to ca. 1 cm long and with ciliate margins. Sterile leaves to 65 cm long and 20 cm broad, petiole 1-10(-12) cm long, with scales like those of the stem. Lamina strongly to moderately reduced at the base, those of adult leaves with (18-)20-30 pairs of alternate pinnae. Rachis broadly to narrowly alate through- out, or at least in the distal half. Pinnae of adult lamina 1-1.8 cm broad, sessile, broadest at or near the base, margins entire to crenulate or serrulate (in juvenile leaves often deeply sinuate to dentate), subcordate, truncate, obtuse or (very rarely) broadly cuneate at base, apices acuminate to long-tapering. Fertile leaves usually shorter than sterile ones, but with proportionately longer petioles. Pinnae 1.5- 4 mm broad, sessile or subsessile. In dense forests, scandent on tree trunks, 1 30- 1300 m, San Martin, Loreto, Junin, Cuzco, and Madre de Dios. Southern Mexico to Panama; Venezuela; Co- lombia to Bolivia. This is very similar to the West Indian L. sor- bifolia (L.) Fee, which differs in its short-stalked, fertile pinnae and its fewer (15-20 pairs) sterile pinnae; the pinnae of the latter are commonly broadest near the center and have a cuneate base. Pinnae of L. fendleri are broadest at or near the obtuse or truncate base. However, pinna bases in L. fendleri are occasionally broadly cuneate, so this character should be used in combination with oth- er features. Stenochlaena angusta Underw., from Colom- bia, is probably to be included within L. fendleri. Two of the isotypes examined, H. H. Smith 1051, Colombia (F, us), are virtually indistinguishable from L. fendleri, except that the sterile pinnae are broadly cuneate at base. A paratype from Peru cited by Underwood (Spruce 1556) is simply a sterile juvenile plant of L. fendleri, with the pinna bases broadly cuneate rather than truncate. In the L. sorbifolia complex, juvenile leaves of- ten are quite different from adult ones. Conspic- uous wings, each up to 1.5 mm broad, are usually borne on the petiole as well as on the rachis, pinna margins may be broadly and deeply dentate to sinuate, and pinna bases often may be broadly cuneate. For these reasons, many South American collections containing only juvenile leaves have been erroneously identified as L. sorbifolia. San Martin: Prov. Mariscal Caceres, Dist. Tocache Nuevo, J. Schunke V. 3569 (F, us), 7609 (F, MO, uc). Tingo Maria, Allard 20889 (GH, us). Loreto: Lower Rio Huallaga, LI. Williams 5080 (F, us), 5052 (F). Junin: East of Quimiri Bridge, near La Merced, Killip & Smith 23908 (F). Cuzco: Prov. La Convention, along Rio Mapitunu- ari, Dudley 10189 (GH). Madre de Dios: Prov. Manu, Parque Nacional de Manu, Rio Manu, Foster et al. 6847 (F). 2. Lomariopsis japurensis (Mart.) John Sm., Hist, fil. 140. 1875. Figure 23b. Acrostichum japurensis Mart., Icon. pi. crypt. 86, t. 24, 1834. TYPE: Colombia (as Brazil), Rio Ja- 106 FIELDIANA: BOTANY pura, Martins (holotype, M?; isotype, L; photos, GH & us). Acrostichum phlebodes Kunze, Linnaea 9: 33. 1835. TYPE: Peru, Maynas, Poeppig, Diar. 2326 (ho- lotype, LZ, destroyed; isotype, B?). Stenochlaena japurensis (Mart.) Griseb., Fl. Brit. W. Ind. 676. 1864. Stem stout, moderately to abundantly provided with essentially concolorous, ferruginous to brownish black scales, these appressed to spread- ing, 3-5(-6) mm long, linear to narrow-lanceolate, their margins subentire to ciliolate. Sterile leaves to 1 m long and 30 cm broad, the petiole 10-30 cm long, with scales like those of the stem, but larger (to 10 mm long). Lamina pinnate (or in juvenile leaves often simple), strongly to moder- ately reduced at base, with (0)1-12 pairs of alter- nate pinnae. Rachis narrowly (if at all) alate only toward the apex. Pinnae (larger ones of adult lam- inae) 3-5 cm broad, sessile or with stalks less than 2 mm, lanceolate or elliptic-lanceolate, margins entire, commonly obtuse or broadly cuneate at base, the apices acuminate. Fertile leaves usually shorter than sterile ones. Pinnae 8-16 mm broad, sessile or subsessile. In lowland rain forests and wooded ravines, scandent (to 12 m) on tree trunks, 100-700 m, San Martin and Loreto to Ayacucho and Madre de Dios. Guatemala to Panama; Trinidad to Colombia, south to Brazil and Bolivia. This species has been confused with L. eryth- rodes (Kunze) Fee of Brazil, which is similar in many laminar characteristics, but which has light- er colored stem and petiole scales and narrower fertile pinnae, these less than 7 mm broad. Lo- mariopsis nigropaleata is even more closely relat- ed, and may be merely a geographic variant of L. japurensis. See treatment of the latter for further discussion. One specimen of L. japurensis from Loreto (Gentry et al. 29748, MO) has more nar- rowly cuneate pinnae (as in L. nigropaleata), and many of the stem scales lighter in color (like L. erythrodes). This is a prime example of the vari- ability of characters in this species complex. San Martin: Prov. Mariscal Caceres, Tocache Nuevo, Quebrada de Cachiyacu, J. Schunke V. 8518 (F, uc). Loreto: Gamitanacocha, Rio Mazan, J. Schunke 306 (F, GH, uc, us, USM). Huanuco: Pachitea, Codo de Pozuzo, Foster 9376 (F, MO). Ayacucho: Rio Apurimac Valley, near Kimpitiriki, Killip & Smith 22871 (us). Cuzco: Prov. La Convencion, near Hacienda Luisiana, Dudley 11571 (GH). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10756 (F). 3. Lomariopsis latipinna Stolze, sp. nov. Folia dimorpha; folium sterile pinnatum, raro sim- plex, usque ad 60 cm longum et 50 cm latum; rhachis prope apicem anguste alatae; paleae petiolae 6-12 mm longae, fulvae vel aurantiaceae, non nitidae, margine ciliolato; pinnae steriles (0)1-4 jugae, maiores 6-9 cm latae, ellipticae vel elliptico-lanceolatae, ad basin anguste cuneatae, apicibus acutis vel subacutis, marginibus in- tegris, petiolulis (3)4-15 mm longis; pinnae fertiles ma- iores 16-20 mm latae, petiolulis 3-5 mm longis. Stem stout, moderately to abundantly provided with concolorous, tawny to dull orange scales, these appressed to spreading, 6-8 mm long, lanceolate, their margins ciliolate. Sterile leaves to 60 cm long and 50 cm broad, the petiole 7-20 cm long, with scales like those of the stem but longer (8-12 mm) and broader. Lamina pinnate (or rarely simple), not or scarcely reduced at base, with (0)1-4 pairs of alternate pinnae. Rachis narrowly alate (if at all) only toward the apex. Pinnae (larger ones of adult laminae) 6-9 cm broad, on stalks (3-)4-15 mm long, elliptic or elliptic-lanceolate, margins entire, narrow-cuneate at base, the apices acute or subacute. Fertile leaves shorter than sterile ones. Pinnae 16-20 mm broad, their stalks 3-5 mm long. TYPE— Peru, Junin, Prov. Chanchamayo, "Pi- chanaki" (Dist. Pichanaqui), van der Werff et al. 8626 (holotype, uc!; photo, F). In lowland rain forests, scandent on tree trunks, 300-700 m, Lambayeque, Pasco, Junin, and Ma- dre de Dios. Apparently endemic. This species and L. nigropaleata both have broad, stalked, cuneate pinnae, but those of the latter species are abruptly narrowed to a sharply pointed apex. Pinnae of L. latipinna are even broader (to 9 cm) and gradually taper to an acute or subacute apex. Stem and petiole scales of the two species also differ markedly: those of L. ni- gropaleata are scattered, tiny and bicolorous, whereas those of L. latipinna (especially at the petiole base) are abundant, larger, and concolor- ous. The lamina of juvenile leaves is occasionally simple in both L. japurensis and L. latipinna. In- sufficient juvenile material has been seen to be able to establish diagnostic characters, but it appears that the simple lamina of L. japurensis is rarely more than 5 cm broad, while in L. latipinna the simple lamina is 8-10 cm broad. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 107 FIG. 24. Lomagramma guianensis: a, portion of stem and petiole, adaxial side; b, portion of sterile leaf; c, rachis of sterile leaf; d, portion of pinna, abaxial side; e, portion of fertile leaf; f, apex of fertile pinna, adaxial side, (a from Soukup & Lopez 4615, GH; b, d, e, f from Schunke V. 2846, F; c from Schmalz 40, Brazil, F.) Lambayeque: Puerto Nazareth, 5 km from Olmos, El- lenberg 3452 (GH). Amazonas: Bagua, forest behind Par- celacion Monterrico, Knapp & Alcorn 7592 (MO). Pasco: Prov. Oxapampa, Puerto Laguna, D. Smith 8412 (MO?, uc). Madre de Dios: Prov. Manu, Parque Nacional del Manu, Cocha Cashu Biological Station, M. Foster P-84- 112, P-84-119(uc). 4. Lomariopsis nigropaleata Holttum, Bull. Misc. Inform. 1939; 618. 1940. TYPE: Peru, Lore- to, Florida, Rio Putumayo, Klug 2082 (ho- lotype, us!; isotypes, BM, F!, GH!, K, NY). Fig- ures 23c-d. Stem stout, sparsely provided with mostly bi- colorous scales, these lustrous black with a fer- ruginous, entire or sparsely ciliolate margin, 1-2 mm long, ovate or broadly lanceolate. Sterile leaves to 1 m long and 30 cm broad, the petiole to 25 cm long, with scattered, appressed scales like those of the stem. Lamina pinnate, somewhat reduced at base, with 4— 10 pairs of pinnae. Rachis narrowly alate only toward the apex. Pinnae (larger ones of adult laminae) 3.5-5.5 cm broad, at least proximal ones on stalks (2-)3-6 mm long, elliptic to oblong- lanceolate, the margins entire to obscurely crenate 108 FIELDIANA: BOTANY or sinuate, cuneate at base, the apices abruptly narrowed to a mucronate or caudate tip. Fertile leaves usually shorter than the sterile ones. Pinnae 7-16 mm broad, on stalks 2-5 mm long. In lowland rain forests and wooded ravines, scandent to 6 m on tree trunks, 100-750 m, San Martin and Loreto to Madre de Dios. Peru; Bolivia; western Brazil. This and L. japurensis are very closely related. The most effective means of distinguishing them is the character of the stem and petiole scales. Those of L. nigropaleata are 1 or 2 mm long, with a usually lustrous black center and ferruginous margin, and are tightly appressed and widely scat- tered on the stem and the base of the petiole. In L. japurensis the scales of the stem are typically 3-5 mm long, rather uniformly dull and reddish or blackish brown, becoming much more numer- ous on the petiole base, where they are usually spreading and up to 1 cm long. Other characters are usually helpful in separat- ing the two species: lengths of pinna stalks typically differ as indicated in the key; and pinna base is commonly narrowly cuneate in L. nigropaleata but obtuse to broadly cuneate in L. japurensis. However, these features are occasionally incon- sistent, which suggests that L. nigropaleata may be merely a variant of L. japurensis. San Martin: Tarapoto, Spruce 4738 (c, K, P, us). Lore- to: Balsapuerto, lower Rio Huallaga basin, Killip & Smith 28594 (GH, NY, us). Huanuco: Prov. Pachitea, Distr. Honoria, Bosque Nacional de Iparia, J. Schunke V. 1288, 137 1 (F, GH, us). Pasco: Prov. Oxapampa, between Is- cozacin and Villa America, D. Smith 2835 (F, MO). Junin: Prov. Chanchamayo, "Pichanaki" (Dist. Pichanaqui), van der Werffet al. 8616 (MO, uc). Ucayali (as Loreto): Prov. Coronel Portillo, Bosque Nacional von Humboldt, Diaz et al. 729 (MO, us). Madre de Dios: Prov. Tam- bopata, Rio Piedras, Vargas 18631, 18637 (GH). XXIV. Lomagramma Lomagramma John Sm., J. Bot. (Hooker) 4:152. 1841. TYPE: Lomagramma pteroides John Sm. Figure 24. Plants terrestrial or sometimes epipetric, usually becoming scandent-epiphytic. Stem long-creeping or scandent to ca. 20 m long, rather slender, bear- ing scales and few to many fibrous roots. Leaves dimorphic (fertile segments narrower than the sterile), ca. 25 cm to 1.5 m long, widely spaced. Lamina 1 -pinnate or rarely 2-pinnate, usually im- paripinnate, the pinnae articulate to the rachis, glabrate to slightly scaly. Veins anastomosing but without included free veinlets. Sporangia borne over the abaxial surface (acrostichoid) or some- times on the margin and extending around it onto the edge of the adaxial surface, mixed with usually few paraphyses. Indusium lacking. Spores mono- lete, more or less ellipsoidal to spheroidal. Lomagramma is a tropical genus of about 20 species, but only L. guianensis occurs in the Neo- tropics. The genus is characterized by a scandent- epiphytic, dorsiventral stem, a 1- to 2-pinnate, usually imparipinnate lamina, articulate pinnae, and anastomosing veins. Although L. guianensis was formerly treated as a species of Bolbitis, it was excluded from that genus by Hennipman in his monograph (1977). References HENNIPMAN, E. 1977. A monograph of the fern genus Bolbitis (Lomariopsidaceae). Leiden Bot. Sen, 2:313-314. HOLTTUM, R. E. 1937. The genus Lomagramma. Gard. Bull. Straits Settlem., 9: 190-221. TRYON, R. M., AND A. F. TRYON. 1982. Loma- gramma, pp. 6 1 3-1 6, in Ferns and allied plants, Springer- Verlag, New York. 1 . Lomagramma guianensis (Aubl.) Ching, Amer. Fern J. 22: 17. 1932. Figure 24. Polypodium guianense Aubl., Hist. pi. Guiane 2: 962. 1 775. TYPE: Aublet, French Guiana (isotype, BM; photos, GH, us). Leptochilus guianensis (Aubl.) C. Chr., Bot. Tidsskr. 26: 288. 1904. Bolbitis guianensis (Aubl.) Vareschi, Flora Venezuela 1: 376. 1969 (credited to Kramer, Acta Bot. Neerl. 3: 486. 1954, which is invalid; Art. 33.2). Stem 2-6 mm thick, sparsely scaly, the scales 1-2 mm long, linear-lanceolate, blackish, obscure- ly clathrate. Leaves 1 -pinnate, with 10-14 pairs of articulate pinnae and a conform or subconform nonarticulate terminal segment, petiole 8-18 cm long, adult leaves 25-60 cm long, 1 4-20 cm broad, but fertile ones much smaller. Rachis sparsely pro- vided with filiform, tortuous, castaneous scales. Sterile pinnae to 15 cm long and 3 cm broad, subsessile, or proximal ones short-stalked, lanceo- late, the apices acute to attenuate, the margins subentire to (distally) crenulate-serrate. Veins TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 109 FIG. 25. Elaphoglossum paleaceum: a, habit, stem with fertile and sterile leaf; b, scales on abaxial surface. Elaphoglossum erinaceum: c, scales of leaf margin. Elaphoglossum piloselloides: d, habit, stem with fertile and sterile leaf; e, scales on abaxial surface. Elaphoglossum squamipes: f, habit, stem with fertile and sterile leaf; g, scales on abaxial surface. (Adapted from Stolze, Ferns and fern allies of Guatemala, 1981.) 110 FIELDIANA: BOTANY anastomosing throughout, but the tips commonly free at the pinna margin. Fertile pinnae linear, 3- 10 cm long, 0.2-0.5 cm broad, the tips obtuse or acute, the slender, tawny or light brown paraph- yses scattered sparsely among the sporangia. Terrestrial, commonly becoming scandent on tree trunks, in lowland forests, 100-900 m, Loreto and Huanuco to Madre de Dios. Greater Antilles (except Jamaica); Colombia to the Guianas, south to Bolivia and Brazil; Argen- tina. Plants initially are terrestrial; later they become scandent on trees but retain their terrestrial con- nection. Stems often creep for long distances, and are slender along the ground but stouter on trees. Fertile leaves most commonly occur on the scan- dent stems and are often most numerous toward the stem apex. Loreto: Pena Blanca on Rio Ataya, Killip & Smith 29674 (F, us). Huanuco: Prov. Pachitea, Dist. Honoria, Bosque Nacional de Iparia, Schunke V. 1372 (F, GH, us). Pasco: Puerto Bermudez (as Junin), Killip & Smith 26572 (F, us). Prov. Oxapampa, Quebrada Castilla, along Rio Omaiz, Leon & Young 1064 (F, USM). Pichis Trail, Santa Rosa (as Junin), Killip & Smith 26158 (GH, us). Ucayali: Prov. Padre Abad, Bosque Nacional von Humboldt, Narite 3 (USM). Madre de Dios: Maldonado, El Pilar, Lopez & Soukup 4583, 4613 (us). XXV. Elaphoglossum Contributed by John T. Mickel Elaphoglossum John Sm., J. Bot. (Hooker) 4: 148. 1841, nom. conserv. TYPE: Elaphoglossum conforme John Sm. Figure 25. Epiphytic, terrestrial, or epipetric. Stem com- pact to long-creeping, horizontal, rarely erect, slen- der to stout (1-15 mm in diameter). Stem scales orange to black, basally attached or peltate, entire to dentate or ciliate. Leaves monomorphic or usu- ally more or less dimorphic, 2-200 cm long, erect, spreading or pendent, simple or rarely pedate. Pet- iole glabrous or scaly, sometimes also with minute glandular trichomes, long or very short, base often darker (phyllopodium) with abscission at its upper demarcation rather than at the stem. Lamina lin- ear to ovate or oblanceolate, apex acuminate or caudate to obtuse, base rounded (rarely cordate) to long-attenuate or decurrent; costa sulcate adax- ially, usually with scales similar to those of the petiole abaxially. Veins generally free, rarely anas- tomosing or with a marginal commissural vein, simple to twice-forked, ending near the margin, those ending well short of the margin usually ter- minating in conspicuous hydathodes. Lamina scales generally differing from those of the stem or petiole, abundant to lacking, often greatly re- duced and appearing as stellate trichomes. Fertile leaves longer or shorter than the sterile ones but generally with narrowed laminae and proportion- ally longer petioles. Fertile lamina completely cov- ered abaxially with sporangia (acrostichoid sori). Sporangia long-stalked, the annulus erect, inter- rupted by the stalk; paraphyses (in the form of intersporangial scales) present in some species, but generally lacking. Spores bilateral, monolete, most with high crests or low ridges, but some echinate or verruculate without ridges or crests. There are probably well over 600 species in Ela- phoglossum, more than three-fourths of them oc- curring in tropical America. There are 1 2 1 species recognized in Peru, although our knowledge of them is far from complete. Several are known from only one or a few specimens. The genus is very difficult taxonomically; it has not been adequately treated with a usable subgeneric breakdown until recently. Variation of the characters, such as plant size, lamina form, scale color, and scale type, is not fully understood in terms of species delimi- tation. The characters lie mostly in the scales of the stem and lamina. The fertile leaves add characters of relative size, intersporangial scales, and spore details, but virtually all the species can be iden- tified on the basis of vegetative material alone. Unfortunately, collectors are hesitant to collect sterile material and thus many records have gone uncollected. The large number of new species de- scribed here is a reflection of the enormous di- versity within the genus and gives an indication of our heretofore poor knowledge of this fascinat- ing group. In the following descriptions, the stem diameter is given excluding the scale covering. The stem scales may be appressed or widely spreading and might too greatly distort the stem diameter mea- surements were they included. In the key and de- scriptions, the terms "leaves" and "laminae," when not specifically referring to fertile leaves and lam- inae, refer to sterile ones, which are much more abundant than the fertile. Petiole bases are differ- entiated into evident phyllopodia in some but not all species. Although the veins seem to run to the margin in some species, they end just short of it, leaving a pale, thin margin 0.5-1 mm wide, which TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. Ill is often difficult to distinguish, especially in very coriaceous or heavily indumented leaves. This margin is more readily distinguished in the fertile leaves where it remains sterile in contrast to the acrostichoid sorus of the rest of the lower surface. Vein angles and intervein distances are measured at midleaf, halfway between the costa and margin. The scales of the stem and lamina are generally quite distinct from one another, and on the petiole they intergrade or in some cases remain distinct and occur together. The lamina scales, although basically the same type on upper and lower sur- faces, are more highly dissected abaxially. In some this means longer teeth, but in more extreme cases they are reduced to stellate trichomes or even to resinous dots. In subglabrous leaves, the lamina scales are reduced further in size to minute stellate trichomes (trichomidia) that are visible only with a lens. On the fertile lamina, the scales adaxially are similar to those of the sterile lamina, but abax- ially the scales are generally limited to the costa and in only a few species are there scales among the sporangia. Another type of indument is mi- nute, erect glandular trichomes, which are found in varying degrees on the petiole and occasionally on the lamina in E. lindenii, E. erinaceum, E. tam- billense, E. rubellum, E. haynaldii, and their rel- atives. In most cases Elaphoglossum is distinct in its simple lamina, only rarely becoming pedate or crested, and generally has free veins and acrosti- choid sori. This treatment contains numerous citations from Christ's "Monographic des Genus Elaphoglos- sum," which appeared in the journal Denkschr. Schweiz. Naturf. Ges. 36. 1899. In the interest of brevity the publication is cited throughout as Monogr. Elaphoglossum. References GOMEZ, L. D. 1972. The first Peruvian record of Elaphoglossum cardenasii Wagner. Amer. Fern J., 62: 95. MICKEL,J. T. 1985. Elaphoglossum, pp. 76-107, in A. R. Smith, ed., Pteridophytes of Venezuela, an annotated list, published by the author, Berkeley, California. MICKEL, J. T., AND L. ATEHORTUA G. 1980. Sub- division of the genus Elaphoglossum. Amer. Fern J., 70: 47-68. Key to Species of Elaphoglossum a. Sterile leaf bearing linear-lanceolate scales that are usually inrolled to appear hairlike (subulate), at least on petiole but generally throughout the leaf, and/or with veins ending in hydathodes b b. Hydathodes lacking; leaves (25-)35 + cm long c c. Petiole lacking; lamina narrowly oblanceolate; lamina scales orange d d. Stem scales slightly sinuous; sterile lamina 12-56 x 1.7-6.0 cm; lamina apex acuminate to long-caudate; spore ridges smooth, surrounding a perforated area 97. E. raywaense d. Stem scales conspicuously flexuous; sterile lamina 45-95 x 7.7-9.5 cm; lamina apex acu- minate to sharply cuspidate; spore ridges short-spiny, surrounding an area with fine spines 3. E. amazonicum c. Petiole present; lamina narrowly elliptic to lanceolate (rarely oblanceolate); lamina scales black to dark brown e e. Lamina scales very sparse, deciduous 93. E. propinquum e. Lamina scales conspicuous f f. Lamina base attenuate g g. Lamina narrowly oblanceolate, 4.6-6.3 cm broad; petiole scales dense, 4-5 mm long 1 2. E. blepharoglottis g. Lamina linear-elliptic, 1.4-1.7 cm broad; petiole scales sparse, 1-2 mm long 44. E. hystrix f. Lamina base rounded to subcordate h h. Lamina base rounded; 700-3325 m elev 28. E. erinaceum h. Lamina base truncate to subcordate; 200-3200 m elev 11. E. barbatum 112 FIELDIANA: BOTANY b. Hydathodes present, or if lacking, leaves only 6-1 5 cm long i i. Stem long-creeping, petioles 0.5-3 cm apart j j. Lamina oblong-ovate, 6.5-12 cm broad, lamina base deeply cordate . . . 86. E. pascoense j. Lamina elliptic to linear-elliptic, 1.2-4.8 cm broad; lamina base cuneate to truncate . . . k k. Lamina scales dark brown, curved; leaves 6-12 cm long 13. E. camptolepis k. Lamina scales orange to dark brown, straight; leaves 14-40 cm long 1 1. Stem 1-2 mm in diameter m m. Leaves 8-12 cm long, 0.6-1.3 cm broad; petiole 0.5-0.7 mm in diameter; fertile lamina 1 .5-2.5 cm long 35. E. gracillimum m. Leaves 30—47 cm long, 1.8-3.0(4.8) cm broad; petiole 1.4—1.7 mm in diameter; fertile lamina 1 1-13 cm long 50. E. latevagans 1. Stem 3-4 mm in diameter n n. Sterile lamina chartaceous, margin crenulate, base truncate; petiole ca. '/2 the sterile leaf length, 0.8-1.0 mm in diameter; petiole scales patent, dark brown 105. E. simulans n. Sterile lamina subcoriaceous, margin entire, base rounded; petiole %-'/3 the sterile leaf length, 1-1.5 mm in diameter; petiole scales appressed to ascending, orange 82. E. pachyrrhizum i. Stem short-creeping, petioles less than 0.5 cm apart o o. Leaves 1 5-70 cm long, or if smaller, lamina scales 3-5 mm long p p. Leaves nearly sessile, petiole nearly lacking or winged; lamina with white patches between veins q q. Lamina linear-oblanceolate; petiole essentially lacking 19. E. zebrinum q. Lamina elliptic to oblanceolate; petiole winged to base 2. E. alipes p. Leaves distinctly petiolate; lamina lacking regular white patches between veins r r. Petiole V4 or more of total leaf length; lamina lanceolate to elliptic, (2)3-10 cm broad (rarely linear), 1.1-1.6 cm broad; lamina and petiole scales linear-lanceolate to lan- ceolate, plane, rarely subulate s s. Leaves 70-1 10 cm long, lamina 8.0-12.8 cm broad, with subulate scales 10. E. bakeri s. Leaves 16-55(-75) cm long; lamina 2.2-7.8 cm wide, with lanceolate or linear- lanceolate scales (rarely subulate in E. rufum) t t. Lamina 3-8 cm broad u u. Lamina with sparse scales ca. 1 mm long; leaves 32-75 cm long v v. Petiole black, especially of the fertile leaves 15. E. castaneum v. Petiole of both sterile and fertile leaves dull gray-green 85. E. papillosum u. Lamina with abundant scales 2-3 mm long; leaves 22-30 cm long w w. Lamina elliptic, never proliferous; intersporangial scales black I.E. albescens w. Lamina lanceolate, usually proliferous at apex; intersporangial scales orange-tan x x. Scales of petiole and lamina margin subulate; petiole scales spreading, abundant; intersporangial scales sparse to lacking . 102. E. rufum x. Scales of petiole and lamina margin lanceolate; petiole scales ap- pressed, scattered; intersporangial scales abundant 117. E. wardiae t. Lamina 2.0-2.4 cm broad Y y. Sterile lamina rounded at base 24. E. diversifrons y. Sterile lamina attenuate at both ends z z. Petioles pale; lamina scales orange-tan, scattered ... 80. E. oxyglossum z. Petioles black; lamina scales dark red-brown, very sparse 68. E. moyeri TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 113 r. Lamina linear to narrowly elliptic, 0.6-3.2 cm broad; petiole scales subulate or lan- ceolate a2 a2. Lamina scales (other than costal) less than 1 mm long, entire b2 b2. Petiole scales subulate to lanceolate, spreading; stem scales linear-lanceolate, spreading c2 c2. Petiole and costal scales dark red-brown, subulate . . 30. E. eximium c2. Petiole and costal scales tan, lanceolate 84. E. palorense b2. Petiole and stem scales ovate, appressed d2 d2. Lamina undivided, entire 103. E. russelliae d2. Lamina pedately lobed (5-7 lobes) 14. E. cardenasii a2. Lamina scales over 2 mm long, denticulate e2 e2. Lamina scales 3-6 mm long; fertile lamina round to oblong, obtuse; fertile leaf much shorter than the sterile f2 f2. Lamina apex acuminate to acute-cuspidate, 1.3-2.4 cm wide; base cu- neate; glandular trichomes to 0.5 mm long; 1500-2450 m elev 39. E. haynaldii f2. Lamina apex acute to obtuse, 2. 1-3.2 cm wide; base rounded; glandular trichomes ca. 0.2 mm long; 400-500 m elev 100. E. rubellum e2. Lamina scales 1-2 mm long; fertile lamina lanceolate to long-oblong, about equal to sterile leaf or longer g2 g2. Lamina 1.7-2.4(3.6) cm wide; fertile lamina truncate at base, 1.8-2.3 cm wide 104. E. setigerum g2. Lamina 0.9-1.6 cm wide; fertile lamina rounded at base, 0.6-1.6 cm wide h2 h2. Leaves 28-38 cm long; lamina apex acuminate; petiole ca. V4 the leaf length; stem scales 3-5 mm long; veins 2-3 mm apart; lamina scales subulate, dark brown 92. E. poeppigianum h2. Leaves 1 1-21(32) cm long; lamina apex acute to obtuse-apiculate; petiole V3-l/2 the leaf length; stem scales to 13 mm long; veins 1 mm apart; lamina scales linear, orange-tan 96. E. quitense o. Leaves 2-16 cm long; lamina scales 1-2(3) mm long i2 i2. Lamina ovate to lanceolate, 1.4-2.7 cm wide, cuspidate or rarely acuminate j2 j2. Lamina truncate to subcordate at base; petiole and lamina conspicuously scaly . . 90. E. pilosius j2. Lamina broadly cuneate to rounded at base; petiole and lamina sparsely scaly to glabrous k2 k2. Lamina sparsely scaly, minute glandular trichomes sparse ... 56. E. lindenii k2. Lamina lacking scales but with abundant minute glandular trichomes, especially at lamina base and upper petiole 109. E. tambillense i2. Lamina linear to spatulate, 0.4-1 .0 cm wide, obtuse 12 12. Lamina linear-elliptic; leaves 7-13 cm long 42. E. horridulum 12. Lamina obovate-oblanceolate to narrowly elliptic; leaves 2— 4(-8) cm long . . . m2 m2. Fertile lamina narrowly oblanceolate, much longer than broad, often inrolled, base cuneate 46. E. hieracioides m2. Fertile lamina orbicular to ovate, flat or conduplicate, base rounded, rarely cuneate n2 n2. Leaves 2-4(6) cm long; sterile lamina spatulate; fertile lamina rounded, strongly conduplicate, with distinctly darker scales than those of sterile leaf; hydathodes very inconspicuous to lacking .... 89. E. piloselloides n2. Leaves 7-15 cm long; sterile lamina elliptic to oblong; fertile lamina orbicular, flat, with scales same color as those of sterile lamina; hydathodes evident 38. E. hayesii 114 FIELDI ANA: BOTANY a. Sterile leaf densely scaly to virtually glabrous; scales various (broadly lanceolate or ovate, ciliate, stellate, round-peltate) but not subulate; hydathodes lacking (except in E. mathewsii and E. hartwegii) o2 o2. Scales, stellate trichomes, or glandular dots present on lamina and petiole; if lamina scales sparse, lamina chartaceous, not coriaceous p2 p2. Lamina scales limited to margin and midvein, laminar surface lacking any form of scale (including resinous dots) q2 q2. Lamina scales golden, round, entire; petiole V4-V3 the sterile leaf length; sterile lamina cuspidate 2 1 . E. decoratum q2. Lamina scales tan-orange, lanceolate, deeply dentate; petiole VS-V6 the sterile leaf length; sterile lamina broadly obtuse 35. E. obtusum p2. Lamina scales not limited to margin and costa, with some form of scale, although perhaps very sparse, on the laminar surface r2 r2. Abaxial surface with stellate trichomes and/or resinous dots s2 s2. Abaxial surface with stellate trichomes, sometimes mixed with resinous dots, or with small, loosely attached scales with resinous dots t2 t2. Abaxial surface with scales, stellate trichomes rare or lacking u2 u2. Stem compact; stem scales reddish black, indurated, ascending; petiole V3 the sterile leaf length 70. E. nastukiae u2. Stem long-creeping; stem scales black, thin, recurved; petiole '/2-3/s the sterile leaf length v2 v2. Leaves 2-3.5 cm long, ovate-acuminate; lamina scales lanceolate, long- ciliate 45. E. jucundum v2. Leaves 15-35 cm long, narrow-elliptic; lamina scales ovate, linear or linear-lanceolate, entire or ciliate only at scale base w2 w2. Lamina scales ovate (or ovate-deltate to ovate-lanceolate), entire; petiole scales imbricate 95. E. punae w2. Lamina scales linear adaxially, linear-lanceolate abaxially, short- ciliate at scale base; petiole scales scattered 59. E. longius t2. Abaxial lamina surface with stellate trichomes (rarely just resinous dots; see couplet y2) x2 x2. Scales of petiole and costa (and adaxial lamina surface) round, fimbriate; those of petiole and abaxial costa often bicolorous (dark-centered with white fringe) y2 y2. Abaxial surface with stellate trichomes 110. E. tectum y2. Abaxial surface with resinous dots 107. E. stenophyllum x2. Scales of petiole and lamina (especially abaxial costa) lanceolate to roundish or stellate, neither conspicuously round and fimbriate nor bicolorous, though some may be dark and araneiform z2 z2. Lamina narrowly elliptic, 6-19 mm wide; costa and petiole with some dark appressed araneiform scales 6. E. angustius 7.2. Lamina linear to narrow-elliptic, 4-6(-9) mm wide; costa and petiole scales stellate to lanceolate, lacking dense, araneiform scales .... a3 a3 . Lamina and petiole scales with noticeable scale body, adaxial scales peltate, round to ovate; abaxial scales all stellate; petiole with linear-lanceolate scales 94. E. pumilio a3. Lamina and petiole scales on both surfaces all stellate b3 b3. Adaxial lamina indument strictly stellate trichomes; fertile leaf shorter than sterile; petiole ca. 0.5 mm diameter, !/3-!/2 sterile leaf length; sterile leaves 9-13(-30) cm long, 4-20 mm broad; terrestrial, at 1500-3650 m 1 12. E. tenuiculum b3. Adaxial lamina indument with slight scale body in center; TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 115 fertile leaf longer than sterile; petiole 0.6-0.9 mm diameter, ca. '/s the sterile leaf length; sterile leaves 7.5-10 cm long, 2.5- 3 mm broad; epiphytic, at 900-1300 m 46. E. killipii s2. Abaxial lamina surface with resinous dots, lacking scales and stellate trichomes c3 c3. Lamina elliptic, 60-75 cm long, 3.4-4.6 cm wide; stem compact 1 9. E. craspedotum c3. Lamina linear to narrowly elliptic, 12-44 (to 78 in E. ciliatum) cm long, 0.8- 2.3(-3.0) cm wide; stem short-creeping d3 d3. Lamina subsessile, V30-Vs the sterile leaf length; veins conspicuous, 2 mm apart 72. E. nigrescens d3. Lamina distinctly petiolate, V4-2/3 the sterile leaf length; veins inconspicu- ous, ca. 1 mm apart e3 e3. Lamina long-cuspidate or acuminate (rarely obtuse); adaxial lamina surface with scattered to abundant hair-toothed scales 88. E. petiolosum e3. Lamina narrowly obtuse to acute; adaxial lamina scales entire to erose or with only short teeth f3 f3. Lamina nearly scaleless above with only scattered 1 mm scales, especially near margin; 400-2650 m elev g3 g3. Stem long-creeping, naked, glutinous; phyllopodia long; epi- phytic at 400-2650 m 17. E. ciliatum g3. Stem short-creeping; phyllopodia short; terrestrial at 2050- 3200 m 43. E. huacsaro O. Lamina scaly adaxially, usually densely so, scales delta te-lanceo- late, often bicolorous, ca. 2 mm long; 3200-5100 m elev. . . . h3 h3. Lamina scales entire to erose 37. E. hartwegii h3. Lamina scales denticulate, ciliate, at least at base 62. E. mathewsii r2. Abaxial surface with round to lanceolate scales (often glabrous in E. mathewsii, E. hartwegii, and E. litanum), lacking stellate trichomes and resinous dots i3 i3. Abaxial surface with round, ciliate, peltate scales; stem scales weakly dentate to entire j3 j 3 . Scales of abaxial surface white, irregular, round, peltate to ovate, variably dentate to ciliate; petiole and abaxial costa scales pale, concolorous; lamina narrowly cuneate at base, apex long-acuminate; stem compact, petioles approximate . . 47. E. laminarioides j3. Scales of abaxial surface mostly reddish, scattered, dark; petiole and abaxial costal scales black with white, fimbriate margin; lamina broadly cuneate at base, apex cuspidate to acuminate; stem creeping, petioles 0.5-2 cm apart 20. E. cuspidatum i3. Abaxial surface with lanceolate, cilio-denticulate scales; adaxial the same or gla- brous; stem scales strongly cilio-denticulate k3 k3. Lamina linear, 30—45 cm long, 0.4-0.6(-1.0) cm wide 13 13. Abaxial lamina scales long-ciliate (cilia longer than scale body is wide), appearing as a dense, woolly mat; lamina 5-10 mm wide 16. E. chloodes 13. Abaxial lamina scales short-ciliate (cilia much shorter than scale body is wide), appressed, not a woolly mat; lamina 4-5 mm wide 115. E. vittarioides k3. Lamina elliptic to narrowly elliptic, 1-6 cm broad m3 m3. Lamina scales undivided (entire, erose to denticulate), not long-ciliate along scale margin, although sometimes ciliate at base, generally dark with pale margin n3 116 FIELDIANA: BOTANY n3. Stem long-creeping; abaxial lamina surface densely scaly 99. E. rosenstockii n3. Stem short-creeping; abaxial lamina surface lacking scales See couplet h3 m3. Lamina scale margins ciliate or coarsely dentate o3 o3. Stem long-creeping, petioles mostly 5-20 mm apart p3 p3. Lamina scales dense, imbricate, surface not or barely visible; lamina lanceolate, long-acuminate 55. E. leprosum p3. Lamina scales scattered to sparse, surface visible; lamina elliptic to linear-elliptic, apex obtuse to acuminate q3 q3. Leaves 55-86 cm long, 3.5-5.6 cm broad; abaxial lamina lacking scales other than sparse, lanceolate ones along costa; lamina margin with dense orange scales 0.5-1 mm long . . 5. E. amplum q3. Leaves less than 53 cm long, mostly less than 35 cm long, 2.0-4. 1 cm broad; abaxial lamina surface scaly, margin not densely orange-scaly r3 r3. Stem 2-3 mm in diameter s3 s3. Stem scales strongly dentate . . 53. E. laxisquama s3. Stem scales entire to sparsely denticulate t3 t3. Phyllopodia 2-3.8 cm long; lamina apex acu- minate; petiole and costal scales linear-lanceo- late, brown to tan, not sclerotic 32. E. fortipes t3. Phyllopodia 1 .2-2.2 cm long; lamina apex acute to obtuse; petiole and costal scales black, scle- rotic, usually appressed ... 116. E. vulcanicum r3. Stem 1-2 mm in diameter u3 u3. Lamina 0.4-0.6 cm wide, leaf 8-16 cm long; abaxial surface scaly, scales overlapping, curled, ciliate-den- tate 111. E. tenue u3. Lamina 1.0-2.5 cm wide, leaf 14-34 cm long; ab- axial surface glabrous or glandular v3 v3. Lamina abundantly glandular abaxially; lami- na apex acuminate; petiole !/2-3/5 the sterile leaf length See couplet u2 v3. Lamina eglandular; lamina apex acute to ob- tuse; petiole V3-V2 the sterile leaf length 116. E. vulcanicum o3. Stem compact, leaves approximate, 0-2 mm apart w3 w3. Lamina apex acuminate, or if only acute, stem scales with widely spreading, long, hairlike teeth x3 x3. Stem scales maroon, linear, 12-20 mm long y3 y3. Lamina scales imbricate, appressed; lamina 25-36 cm long; petiole scales appressed to ascending; 1 100-2750 m elev 29. E. erythrolepis y3. Lamina scales loosely arranged; lamina (30-)52-66 cm long; petiole scales widely spreading; 3400-3750 m elev. 101. E. ruficomus x3. Stem scales orange to black, linear-lanceolate to lanceolate, 1-6 mm long, often with hairlike teeth z3 z3. Petiole V^o-'/eC/i) the sterile leaf length a4 a4. Lamina apex abruptly long-acuminate; stem scales TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 117 1 mm long, brown, entire, somewhat resinous; stem 2 mm in diameter 63. E. megalurum a4. Lamina gradually diminishing at apex; stem scales 4-7 mm long, orange to orange-tan or dark brown, long-ciliate or entire; stem 5-10 mm in diameter b4 b4. Stem scales dark reddish brown, very indu- rated, entire; stem ca. 5 mm in diameter; sterile lamina glabrous abaxially, costa with minute, ciliate scales, glabrescent adaxially except for dense, cilio-denticulate, 1 mm long, pale scales at margin 58. E. litanum b4. Stem scales orange to orange-tan, thin, long- ciliate; stem ca. 10 mm in diameter; lamina densely ciliate-scaly on both surfaces .... c4 c4. Lamina apex acute, the scales lanceolate, 6-12 cells wide; petiole terete 91. E. plumosum c4. Lamina apex acuminate, the scales skele- tonized, 3 cells wide, petiole flattened . . 9. E. auricomum z3. Petiole %-% sterile leaf length or, if shorter, at least some petioles of the same plant longer d4 d4. Stem scales orange to black, entire or weakly pro- vided with lax, hairlike teeth e4 e4. Stem scales with lax, tan trichomes; lamina scales appressed 79. E. orbignyanum e4. Stem scales entire; lamina scales loose 40. E. hickenii d4. Stem scales black, with stiff, black, hairlike teeth f4 f4. Lamina elliptic; costal scales orange 83. E. paleaceum f4. Lamina linear-elliptic; costal scales black . . g4 g4. Lamina scales dense; costal scales lanceo- late, 2-3 mm long .... 98. E. rimbachii g4. Lamina scales scattered; costal scales arachnidoid, 0.5-1 mm long 36. E. guamaniatfsm w3. Lamina apex obtuse to acute, stem scales entire to weakly dentate or with long-ascending hairlike teeth h4 h4. Lamina ( 1 0-) 1 4—33 mm broad, with orange to maroon scales i4 i4. Abaxial lamina scales black, appressed 7. E. atropunctatum i4. Abaxial lamina scales orange (or black only on costa), loosely arranged J4 j4. Leaves 65-70 cm long, 5.5-6 cm broad; fertile leaf linear 22. E. dichroum j4. Leaves 1 7—47 cm long, 1 .4-2.7 cm broad; fertile leaf elliptic k4 k4. Larger petiole scales long-ciliate 26. E. engelii 118 FIELDIANA: BOTANY k4. Larger petiole scales erose to very short-ciliate 14 14. Lamina scales scattered, greatly reduced, sometimes nearly to stellate trichomes; cos- tal scales black; petiole scales black with white margin 76. E. oculatum 14. Lamina scales abundant; petiole and costal scales concolorous, orange-tan 69. E. muscosum h4. Lamina 4-1 2 mm wide, or if broader (to 22 mm), with black stem scales m4 m4. Stem scales castaneous to dark red-brown n4 n4. Lamina 4-7 mm wide; costal and laminar scales linear-lanceolate, laminar ones with hairlike teeth much longer than scale body is wide 113. E. tomentellum n4. Lamina 6-1 2 mm wide; costal scales ovate to lan- ceolate, ciliate, black, lightly appressed, the hair- like teeth about half as long as scale body is wide 49. E. lasioglottis m4. Stem scales black o4 o4. Lamina narrowly elliptic, 8-15 mm broad; lami- nar scales erect, nearly round with cilia longer than scale body 48. E. lanatum o4. Lamina elliptic, 12-22 mm wide; laminar scales appressed, imbricate, lanceolate, erose to short- ciliate 73. E. nivosum o2. Scales lacking on lamina and petiole except rarely some minute, stellate trichomes or a few scales at petiole base; if lamina scales generally distributed abaxially or along margin, lamina very coriaceous; stem and lamina scales not regularly dentate, only occasional processes or weak trichomes along margin p4 p4. Stem 4-10 mm in diameter q4 q4. Lamina nearly sessile, petiole less than !/,0 the sterile leaf length r4 r4. Lamina coriaceous, with scales, especially toward lamina base and petiole; fertile lamina 3-3.6 cm wide 60. E. lurid urn r4. Lamina chartaceous, this and petiole lacking scales; fertile lamina 2-2.8 cm wide s4 s4. Lamina 4.0-5.2 cm wide, apex acute; petiole of sterile leaf ca. 1 cm long; stem scales orange; fertile lamina 2 cm wide 71. E. nidiformis s4. Lamina 1 .9-2.4 cm broad, apex acuminate; petiole of sterile leaf 2-3(-6) cm long; stem scales dark brown; fertile lamina 2.3-2.8 cm broad 52. E. lawyerae q4. Lamina elliptic, with distinct petiole '/8 or more the sterile leaf length t4 t4. Lamina surface abaxially with conspicuous reddish brown, skeletonized scales; lam- ina very coriaceous, blue-green adaxially when fresh 66. E. metallicum t4. Lamina surface abaxially lacking conspicuous scales; lamina chartaceous to coria- ceous, not distinctly blue-green when fresh u4 u4. Stem erect, a slender trunk 30-90 cm long; petiole !/8-% the sterile leaf length; lamina oblanceolate 54. E. lechlerianum u4. Stem horizontal; petiole V4 or more the sterile leaf length; lamina elliptic or linear v4 v4. Stem long-creeping; lamina punctate with glands or stellate trichomidia . . w4 TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 119 w4. Abaxial lamina surface with stellate trichomidia; leaves 60-75 cm long; fertile lamina linear 118. E. williamsiorum w4. Abaxial lamina surface with glandular dots, sometimes with adhering spores making them appear black; leaves 25—4 1 cm long; fertile lamina elliptic, similar in size and shape to the sterile 64. E. meladenium v4. Stem compact to moderately creeping x4 x4. Lamina linear, 1.8-2.1 cm wide; stem scales orange and on petiole beyond phyllopodia 114. E. velongum x4. Lamina elliptic to ovate-lanceolate, 4.9-15 cm wide; stem scales dull brown to blackish, rarely orange, not covering phyllopodia y4 y4. Lamina ovate-lanceolate, 9-1 5 cm wide, abaxial surface with fine, dissected, hairlike scales along costa .... 81. E. pachyphyllum y4. Lamina elliptic, 4.9-9.8 cm wide, abaxial surface glabrous or with occasional lanceolate scales along costa near lamina base 51. E. latifolium p4. Stem l-3(— 4) mm in diameter z4 z4. Stem long-creeping, petioles often over 1 cm apart a5 a5. Stem 1-1.5 mm in diameter; stem and petiole scales tan; leaves 2-11 cm long; phyllopodia lacking b5 b5. Lamina lanceolate-ovate to lanceolate, oblanceolate, or suborbicular, apex ob- tuse c5 c5. Lamina lanceolate to ovate-lanceolate; leaves 7-12 cm long; abaxial lamina scales inconspicuous, tan, not sclerotic 106. E. squamipes c5. Lamina ovate to oblanceolate or suborbicular, 1 .3-2.4(-3.5) cm long, with dark brown to black sclerotic scales, especially abaxially 18. E. concinnum b5. Lamina elliptic to narrowly lanceolate, apex acuminate d5 d5. Leaves (9-)21-24 cm long, 1.4-2.0 cm wide; petiole scales appressed to ascending, 3—4 mm long; lamina apex acuminate; lamina scales scattered 27. E. ensiforme d5. Leaves 7-12 cm long, 0.7-1.1 cm wide; petiole scales widely spreading, 2-3 mm long; lamina apex acute to acuminate, lamina scales sparse, mostly along costa 6 1 . E. macilentum a5. Stem (1.5-)2-3 mm in diameter; stem scales black to tan, sparse; leaves 15-38 cm long; phyllopodia present e5 e5. Stem scales lustrous, strongly spreading, blackish brown, not sclerotic . . . . f5 f5. Lamina lanceolate, 28-48 cm wide 65. E. melancholicum f5. Lamina linear-elliptic, 10-13 cm wide 8. E. atrosquamatum e5. Stem scales dull to lustrous, orange-tan to black, obviously sclerotic, not strong- ly spreading (except in E. patinii) g5 g5. Stem scales lanceolate, mostly 2-3 mm long, orange-tan to dark brown, thin or sclerotic; leaf apex acute to acuminate; 1200-3500 m elev. . . h5 h5. Sterile lamina ovate, 5.5-6.5 cm broad 78. E. oophyllum h5. Sterile lamina elliptic to lanceolate or oblanceolate, 1.7-5.5 cm broad i5 i5 . Sterile lamina linear-elliptic, 1 . 7-2.0 cm broad; stem scales orange- tan, widely spreading 87. E. patinii i5. Sterile lamina elliptic, lanceolate or oblanceolate, 2.2-5.5 cm broad; stem scales orange-tan to black, appressed or slightly ascending J5 j 5 . Abaxial lamina surface with black, coarsely dentate scales and/ or conspicuous, black, stellate trichomidia 77. E. odontolepis 120 FIELDIANA: BOTANY j5. Abaxial lamina surface glabrous or with minute trichomidia inconspicuous to the naked eye 34. E. glossophyllum g5. Stem scales deltate to lanceolate, 0.5-1.5 mm long, black, sclerotic; leaf apex obtuse or long-acuminate; 300-2200 m elev k5 k5. Lamina narrowly elliptic, chartaceous, apex long-acuminate, base at- tenuate 4. E. amphioxys k5. Lamina ovate to lanceolate or oblong, coriaceous, apex obtuse to acute, base narrowly to broadly cuneate 57. E. lingua z4. Stem short-creeping or compact; petioles generally 1-5 mm or less apart, rarely 1 cm apart 15 15. Lamina very broadly lanceolate, 50-100 cm long, (5.5)9-1 5 cm broad; lamina base usually broadly rounded 81. E. pachyphyllum 15. Lamina linear, elliptic or oblanceolate, less than 42 cm long, 5 cm broad; lamina base cuneate to attenuate m5 m5. Lamina apex broadly obtuse; lamina obovate, margin with dissected orange scales 74. E. obovatum m5. Lamina apex narrowly obtuse to acuminate or cuspidate; lamina elliptic to linear or narrowly oblanceolate; scales lacking to lightly distributed abaxially, not concentrated on margin n5 n5. Lamina margin much thickened or inrolled, lamina lanceolate-acuminate to obovate-obtuse-cuspidate, lustrous, glabrous, or with scattered stellate trichomidia; stem scales orange, linear, with irregular processes 108. E. styriacum n5. Lamina margin not thickened, dull below; lamina linear to elliptic or oblanceolate o5 o5. Lamina linear or linear-elliptic, 5-10(16) mm wide p5 p5. Stem scales concolorous, black to dark brown, often with mar- ginal row of light brown cells; lamina glabrous 32. E. glabellum p5. Stem scales tan or with varying degrees of black streaks; lamina minutely punctate, rarely with small stellate trichomes like tufts of wool 67. E. minutum o5. Lamina elliptic or oblanceolate, (16-)20-48 mm wide q5 q5. Lamina oblanceolate; leaf nearly sessile, petiole ca. V20 the leaf length 31. E. flaccid um q5. Lamina elliptic; leaf distinctly petiolate; petiole V5-V2 the leaf length r5 r5. Stem scales reddish orange; lamina abruptly decurrent; pet- iole about half the sterile leaf length 23. E. discolor r5. Stem scales dark brown; lamina abruptly to gradually nar- rowed at base s5 s5. Lamina elliptic to broadly so, 2.7-4.8(-7.5) cm broad, abruptly (to gradually) narrowed and slightly decurrent at base; petiole V4-V3 the sterile leaf length 25. E. elegantipes s5. Lamina narrowly elliptic, 1.6-2.5 cm broad, gradually tapering at base; petiole !/5 the sterile leaf length 32. E. glabellum 1 . Elaphoglossum albescens (Sodiro) Christ, de Lloa, Sodiro (not located); cerca de Chillanos, Monogr. Elaphoglossum 1 23. / 68. 1 899. Sodiro - Acrostichum albescens Sodiro, Recens. crypt, vase. Stem creeping, 3-4 mm in diameter, with white Quit. 75. 1883. SYNTYPES: Ecuador, en el valle blotches on it, the petiole and the lamina scales TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 121 linear, lustrous, blackish maroon, 2-3 mm long, entire or with sparse teeth. Phy Ho podia lacking. Leaves slightly apart, 16—40 cm long, 2.2-3.7 cm broad. Petiole !/3-2/3 the sterile leaf length, scales linear, dark (or with pale tip), lustrous, to 4 mm long, minutely serrulate, appressed to ascending. Lamina elliptic, chartaceous, apex acuminate to caudate, base broadly cuneate to rounded. Veins obscure, ca. 1 mm apart, at 70-80 degree angle. Hydathodes present. Lamina scales linear, brown, lustrous, to 3 mm long, scattered on laminar sur- face, to 5 mm on margin, more abundant on mar- gin. Fertile leaves longer than the sterile, petiole 4/5 the fertile leaf length, scales black and with more pronounced teeth, scales black on costa and mixed with the sporangia. Terrestrial and epiphytic in wet forest, 1700- 3450 m, Amazonas, San Martin, Huanuco, Cuzco. Colombia; Ecuador; Peru. Although the material cited here is relatively uniform, there are a few specimens that do not fit well into this species. For example, Young 1950 (USM) (San Martin: Mariscal Caceres) has the pet- iole subglabrous but with scattered black scales, the lamina is adaxially glabrous and abaxially with scales scattered, black, lanceolate, 2-3 mm long. Further, Plowman & Davis 4802 (GH) (Cuzco: La Convencion) has petiole with scales dark, more or less appressed, the leaf is very large (to 64 cm long, 9.5 cm broad) with scales dark, 1.5-2 mm long. These may represent distinct species. There are additional variations from Ecuador. More study is needed of E. albescens throughout its range to determine how many taxa might be recognized in this complex. Amazonas: Chachapoyas, Cerros Calla Calla, on road to Leimebamba, Hutchinson & Wright 5797 (F, GH in part, MO, P, uc, us). San Martin: Mariscal Caceres, NW corner of Rio Abiseo Nat. Park, Young 4462 (USM). Huanuco: Muna, Bryan 549 (F). Cuzco: La Convencion, Valle de Santa Ana, above Quillabamba, Plowman & Davis 4802 (GH). 2. Elaphoglossum alipes Mickel, sp. nov. Inter species squamis subulatis provisis stipite alato unicum. Stem short-creeping, 2-3 mm in diameter, scales linear-deltate, lustrous, blackish maroon, 1.5-2 mm long, entire. Phyllopodia lacking. Leaves 1-2 mm apart, 18-26 cm long, 2.7-3.9 cm broad. Pet- iole lacking. Lamina oblanceolate, membranous, apex acute, base narrowed to a wing about V3 the sterile leaf length. Veins evident, free, 1.5-2 mm apart, at 70 degree angle. Hydathodes present. Lamina scales scattered on both surfaces, subulate, tan, 2 mm long on surfaces, 3 mm long on margin. Fertile leaves unknown. TYPE— Peru, Ucayali (as Loreto), along Rio Aguetia (Aguaytia) above mouth of Quebrada Yu- rac-Yacu, 2 Oct 1972, Croat 20857 (holotype, uc!). Endemic. Epiphytic in wet forest, no elevation given, Ucayali. The winged petiole, subulate scales, and white lamina patches distinguish this species. Thus far known only from the type. 3. Elaphoglossum amazonicum Atehortua ex Mickel, sp. nov. Ab affinibus rhizomate crasso, rhizomatis paleis flex- uosis, laminis sterilibus latioribus apice cuspidatis, spo- risque spinulosis differt. Stem compact, horizontal to erect, 1-2 cm in diameter, scales linear-lanceolate, fulvous, con- colorous, to 15 mm long, lustrous, 0.5 mm broad, crispate to very flexuous toward the tip. Phyllo- podia present, 1.5 cm long, dark brown, aero- phores bilateral to the phyllopodium like a long strip, and visible even in dry specimens but totally covered with dense mass of scales. Leaves fascic- ulate, 45-95 cm long, 7.5-10 cm broad. Petiole of sterile leaf 2-10 cm long, conspicuously winged, stout, terete but abaxially deeply canaliculate, 0.4 mm in diameter, densely scaly, scales subulate, 0.8 mm long, fulvous to brown-ferrugineous to- ward the costa, also with minute, short, resinous, capitate, glandular trichomes on the petiole and costa. Lamina oblanceolate, chartaceous to sub- coriaceous, apex acuminate to abruptly long-cus- pidate, base attenuate. Veins 1-2 mm apart, at 45 degree angle to costa. Hyathodes lacking; costa prominent on the abaxial side and densely scaly. Lamina scales lacking (at least in dry specimens) except on the costa and margin, but lamina densely covered with appressed, yellow, branched, glan- dular trichomes, which are usually oriented to- ward the margin, margin hyaline, densely covered with 2-3 rows of fine, orange to yellow, subulate scales similar in size and shape to those that cover the costa, and mixed with erect, capitate, glandular trichomes. Fertile leaf shorter than the sterile but 122 FIELDIANA: BOTANY with a longer petiole (3-8.5 cm long), lamina nar- rowly lanceolate, 25-50 cm long, 2.3-4.0 cm broad, apex acuminate, base attenuate, the petiole be- coming winged proximally; intersporangial scales lacking. Spores 25.4-31.5 pm long with narrow, short, spiculate ridges. TYPE— Peru, San Martin, Zepelacio, near Moyo- bamba, Oct-Nov 1933, Klug 3330 (holotype, us!; isotypes, BM!, MO!, NY!). This species is very closely related to E. ray- waense and E. latwn (Mickel) Atehortua ex Mick- el, comb. nov. (E. apodum (Kaulf.) Smith var. la- turn Mickel, Amer. Fern J. 69: 100. 1979), but differs in the flexuous scales, large size, cuspidate lamina apex, stout, fibrous petiole, and finely spi- nulose spores. Underwood annotated specimens of this species (e.g., R. S. Williams 1027, NY) as a new species of Elaphoglossum, using the epithet williamsii, but did not publish the name. There is now an E. williamsii Vareschi (in sect. Elaphoglossum, sub- sect. Pachyglossd), so another epithet was needed. San Martin: Prope Tarapoto, in monte Campana, Pe- ruvia orientalis, Spruce 4639 (BM). Junin: Jauja, Satipo, Monte Alto margen del rio, Ridoutt 11562 (us). Cuzco: Ccochayoc, Biies 1737 (us). 4. Elaphoglossum amphioxys Mickel, sp. nov. Rhizomate gracillimo squamis scleroticis nigris induto necnon lamina utrinque attenuata nuda inter affines sin- gula. Stem long-creeping, 1.5-2 mm in diameter, scales scattered, ovate to lanceolate, dull to lus- trous, sclerotic, orange to black, ca. 1 mm long, subentire. Phyllopodia present. Leaves 1-3 cm apart, 18-31 cm long, 2.0-4.3 cm broad. Petiole about '/a the sterile leaf length, glabrous. Lamina lanceolate to narrowly elliptic, chartaceous, apex long-acuminate, base attenuate. Veins obscure, free, 1 mm apart, at ca. 80 degree angle. Hydathodes lacking. Lamina scales lacking, but surface punc- tate with resinous dots or substellate trichomes. Fertile leaves equal to or slightly shorter than the sterile, petiole V2-3/5 the leaf length, lamina nar- rowly lanceolate; intersporangial scales lacking. TYPE— Peru, Ucayali (as Loreto), Chacra de Cesar Vela SE of granja del Sr. Parrera (Aguaytia), Coronel Portillo, Padre Abad, J. Schunke V. 5493b (holotype, NY!). Endemic. Epiphytic in wet forests, 295-800 m, Amazonas, Ucayali, Huanuco, Junin, Cuzco, Ma- dre de Dios. Elaphoglossum amphioxys is allied to E. lingua but is distinct by the slender lamina that is atten- uate at both ends. Amazonas: Chachapoyas, cerro Puma Urco, Soukup 4089 (us). Huanuco: SW slope of Rio Llullapichis wa- tershed, Dudley 13331 (us). Junin: Prov. Tarma, Tarma- La Merced road, Skog et al. 5021 (us). Cuzco: Paucar- tambo, Cosnipata Valley, Rio Tono, ridge on road N of Patria, Wachter 168 (F, GH). Madre de Dios: Manu, Cerro de Pantiacolla, Rio Palotoa, 10-15 km NNW of Shin- tuya, Foster 10755 (F, GH). 5. Elaphoglossum amplum Mickel, sp. nov. Rhizomate elongate repenti necnon lamina magna marginem versus squamulis minutis induta distinguen- da. Stem compact, horizontal, 2-5 mm in diameter, scales linear-lanceolate, lustrous, dark brown, 1- 2 mm long, with short, patent teeth. Phyllopodia evident but hidden by scales. Leaves 0.5-1 cm apart, 55-86 cm long, 3.5-5.6 cm broad. Petiole about '/2 the sterile leaf length, scales linear, dark brown to tan, appressed to ascending, 2-3 mm long, with short, patent teeth, lamina narrowly el- liptic, chartaceous, apex acuminate, base broadly cuneate. Veins evident, free, 1.5-2 mm apart, at ca. 80 degree angle. Hydathodes lacking. Lamina scales abaxially scattered, round-peltate, ciliate, margin with dense orange scales 0.5-1 mm long, costa scales appressed, tan, dentate, 0.5-2 mm long, adaxial surface punctate with minute stellate trich- omidia. Fertile leaves longer than the sterile, pet- iole % the fertile leaf length, lamina linear, 1 .8- 2.0 cm broad; intersporangial scales lacking. JYPE— Peru, Cuzco, Quillabamba, Santa Tere- sa, Mandornilloc, 0.5 km W of La Playa, Peyton & Peyton 1246 (holotype, GH!). Endemic. Terrestrial in wet forests, 2390-2700 m, Huanuco, Pasco, Cuzco. In the creeping stem and long leaves it super- ficially resembles E. williamsiorum but differs from that species in having dentate, recurved stem scales, marginal dentate lamina scales, and linear fertile lamina. Huanuco: Muna, Bryan 547 a (F). Pasco: Oxapampa, SW of Oxapampa on road to Maria Teresa & Llaupi, Foster 7617 (F). Cuzco: Urubamba, Machu Picchu, 0.5 TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 123 km N of union of Sayacmarca & Aobamba Rivers, Pey- ton & Peyton 1460 (GH). 6. Elaphoglossum angustius Mickel, sp. nov. Ab E. tenuiculo stipite paleis nigris araneiformibus obsito laminaque majori diversa. Stem short-creeping, 2-4 mm in diameter, scales linear, blackish maroon to castaneous with tor- tuous hair tip, ca. 3 mm long, subentire. Phyllo- podia distinct. Leaves fasciculate, 1 5-46 cm long, 0.6-1.9 cm broad. Petiole V6-V3 the sterile leaf length, scales 1-2 mm long, some ascending, these sparse, linear-lanceolate, dark, sparsely dentate, ca. 2 mm long, mostly stellate and appressed, also some dark, appressed and arachnoid. Lamina nar- rowly elliptic, apex acuminate to caudate, base cuneate. Veins obscure, free, ca. 1 mm apart, at 60-70 degree angle. Hydathodes lacking. Abaxial lamina scales stellate, scattered, those on adaxial surface often with slight body or surface glabres- cent. Fertile leaves longer than the sterile, petiole %-2/3 the fertile leaf length, lamina narrower than the sterile; intersporangial scales lacking. TYPE— Peru, San Martin, Zepelacio, near Moyo- bamba, Klug3503 (holotype, NY!; isotypes, F!, GH!, K!). Endemic. Epiphytic in wet forests, (100) 900- 2300 m, Amazonas, San Martin, Loreto, Lima, Junin, Ucayali, Cuzco. This is related to E. tenuiculum but differs in the larger leaves, petiole and abaxial costa often with black araneiform scales, and the scales of the adaxial lamina surface with a slight scale body, not strictly stellate trichomes. Occasionally the stellate trichomes are gland-based, as in J. Schunke 369 and Saunders 1247. Amazonas: Mendoza, Woytkowski 8141 (GH). Loreto: Gamitanacocha, J. M. Schunke 14264 (us). Maynas, Gamitanacocha, Rio Mazan, J. M. Schunke 369 (F, GH, NY, uc). Lima: Canta, Huamantanga, 4 km on road to Huamantanga from Lima-Canta road, Saunders 1247 (F). Canta, 1 km on same road, Saunders 1238 (GH). Junin: Chanchamayo valley, C. Schunke 1349 (F), 1453 (F). Chanchamayo, C. Schunke Aug. 1908 (F, GH). San Ramon, Killip & Smith 24772 (NY). Ucayali (as Loreto); Rio Aguaytia, Croat 20904 (uc). Cuzco: Machu Picchu, road to ruins, Leon 460 (F). Rio Marcapato, 60 km above Quincemil, Madison 1010, in part (GH), 70/0 (GH). 7. Elaphoglossum atropunctatum Mickel, sp. nov. Ab E. oculato laminae facie abaxiali paleis appressis nigris obsita abstans. Stem compact, horizontal, ca. 5 mm in diam- eter, scales linear-lanceolate, dull orange, with dark center, 4-7 mm long, dentate. Phyllopodia evi- dent, often under scales. Leaves approximate, 1 8- 30 cm long, 2.0-3.2 cm broad. Petiole !/4-2/5 the sterile leaf length, with scales ovate-lanceolate, erose-denticulate, orange-tan scales, 3-5 mm long, and also some smaller, appressed scales with dark center. Lamina narrowly elliptic, coriaceous, apex obtuse, base broadly cuneate. Veins obscure, free, ca. 1 mm apart, at ca. 70 degree angle. Hydathodes lacking. Lamina scales abaxially ovate to lanceo- late, black, with pale hair-teeth, 1 mm long, scat- tered to dense, with round peltate ones ca. 0.5 mm long between the larger ones, adaxially ovate to lanceolate, white, cilio-denticulate, 1-2 mm long. Fertile leaves unknown. TYPE— Peru, Pasco, Oxapampa, 2-4 km N of Mallampampa, D. Smith & Canne 5835 (holotype, MO!). Endemic. Epiphytic or terrestrial on organic matter in wet forest, 2200-2400 m, Huanuco, Pas- co, Junin. This species is distinct from E. engelii by the erose-denticulate petiole scales, and from E. mus- cosum and E. oculatum in the dense, dark, ap- pressed scales on the abaxial lamina surface. Huanuco: Carpish, Coronado 74 (us). Pasco: Oxapam- pa, 2-4 km N of Mallampampa, D. Smith & Canne 5861 (MO). Junin: Prov. Tarma, Tarma-La Merced, Skog el al. 5023 (us). 8. Elaphoglossum atrosquamatum Mickel, Brit- tonia 39: 315. 1987. TYPE: Venezuela, Ta- chira, faldas debajo del Paramo de Tama, cer- ca de la frontera Colombo-Venezolana, Steyermark, Dunsterville & Dunsterville 98348-A (holotype, NY!; isotype, MO!). Stem long-creeping, ca. 2 mm in diameter, scales spreading, linear-lanceolate, blackish brown, very indurated, lustrous, 6-8 mm long, margin entire. Phyllopodia present. Leaves 0.5-1.5 cm distant, sterile ones 5-20.5 cm long, 1.0-1.3 cm wide. Pet- iole 0/5) y3-V2 the sterile leaf length, scales sparse 124 FIELDIANA: BOTANY to lacking (deciduous), spreading, lanceolate, dark brown, lustrous, 2-4 mm long, margin entire or with a few long hairlike processes. Lamina nar- rowly elliptic, coriaceous, apex acuminate, base narrowly cuneate, margin distinctly revolute. Veins mostly obscure, 1 mm apart, at 70-80 degree an- gle. Hydathodes lacking. Abaxial lamina scales scattered, linear-deltate, black to brown, lustrous, 0.5-2 mm long, often hastate, margin subentire or with long, hairlike processes and surface with or- ange stellate trichomidia, adaxial surface glabrous or with costal scales sparse, black, as on abaxial surface. Fertile leaves about equal to the sterile in size and shape, petiole '/j-V^ the fertile leaf length; lamina scales present only along the costa. Terrestrial (epiphytic and epipetric in Venezue- la) in wet forest, 3500-4000 m, San Martin, An- cash. Venezuela; Colombia; Peru. The slender, long-creeping stem with recurved, black scales and the narrowly elliptic lamina with scattered black scales along the costa distinguish this species. San Martin: Mariscal Caceres, NW corner of Rio Abi- seo Nat. Park, Chochos, Young & Leon 4687 (USM). Ancash: Carhuaz, Huascaran Nat. Park, Quebrada Ishin- ca, D. Smith & Buddensiek 11257 (F, HUT, NY). 9. Elaphoglossum auricomum (Kunze) Moore, In- dex fil. 7. 1857. Acrostichum auricomum Kunze, Linnaea 9: 28. 1834. TYPE: Peru, Huanuco, Pampayacu, Poeppig, July 1829 (holotype, LZ, destroyed; isotype, P!; photo, us of P). Acrostichum dadotrichum Sodiro, Sert. fl. ecuad. 1: 9. 1905. TYPE: Ecuador, Nanegal, Sodiro, Sept. 1901 (holotype?, P!; isotypes, B!, P!, us!). Stem short-creeping, 3-5 mm in diameter, scales linear-lanceolate, orange, 4-7 mm long, margin with hairlike teeth. Phyllopodia inconspicuous. Leaves fasciculate, 19-27 cm long, 1.4-1.6 cm broad. Petiole nearly lacking, ca. 1 cm long, dense- ly clothed with spreading, orange scales 4 mm long. Lamina narrowly elliptic, chartaceous, apex acu- minate, base narrowly cuneate. Veins obscure, free, 1-2 mm apart, at 60-70 degree angle. Hydathodes lacking. Lamina scales orange, liberally distrib- uted but not overlapping, linear, 1-4 mm long with long, hairlike teeth, the teeth at least half as long as the scale, some scales reduced to nearly stellate trichomes. Fertile leaves slightly shorter than the sterile, petiole ca. '/3 the fertile leaf length, with a few stellate trichomes on the costa abaxially, lam- ina narrower than the sterile; intersporangial scales lacking. Epiphytic in wet forests, 850 m, Huanuco. Mexico to Costa Rica; Colombia to Bolivia; Hispaniola. Huanuco (as San Martin): Tingo Maria, Allard 21359 (us). 1 0. Elaphoglossum bakeri (Sodiro) Christ, Mono- gr. Elaphoglossum 132. 1899. Acrostichum bakeri Sodiro, Recens. crypt, vase. Quit. 77. 1883. SYNTYPES: Ecuador, Atacazo cerca de Canzacoto en la orilla del rio Yamboya, So- diro, (not located); Ecuador, mas abajo hasta en la zona tropical, Sodiro (not located). Stem short-creeping, 4-12 mm in diameter, scales linear-lanceolate, light brown, lustrous, to 7 mm long. Phyllopodia lacking. Leaves fascicu- late, 70-1 10 cm long, 8.0-12.8 cm broad. Petiole about '/j the sterile leaf length, with scales abun- dant, widely spreading, subulate, brown, lustrous, sparsely serrulate. Lamina narrowly lanceolate, chartaceous, apex short cuspidate, base broadly cuneate to rounded. Veins evident, free, ca. 2 mm apart, at ca. 80 degree angle. Hydathodes distinct. Lamina scales castaneous, on costa abundant, widely spreading as on petiole, 2—4 mm long, on both surfaces and margin scattered, 1-2 mm long. Fertile leaves longer than the sterile, petiole ca. 2/3 the fertile leaf length, lamina slightly narrower than the sterile, scales appressed on costa, and scattered among the sporangia. Terrestrial in wet forest, 2000 m, Amazonas. Costa Rica; Ecuador; Peru. Amazonas: Prov. Bagua, ca. 20 km E of La Peca, Bar- hour 2815 (F, uc). 11. Elaphoglossum barbatum (Karsten) Hieron., Bot. Jahrb. Syst. 34: 553. 1904. Acrostichum barbatum Karsten, Fl. Columb. 2: 155. t. 181. 1869. TYPE: Colombia, Bogota, Lindig 111 (not located). TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 125 Elaphoglossum lindbergii var. truncatum Rosenst., Repert. Spec. Nov. Regni Veg. 25: 63. 1928. TYPE: Bolivia, Hacienda Simaco sobre el camino a Tipuani, Buchtien 5166 (isotype, us!). Stem horizontal, compact, 8-16 mm in diam- eter, scales linear, lustrous, castaneous, 4-7 mm long, with irregular teeth near scale apex. Phyl- lopodia lacking. Leaves fasciculate, 35-54 cm long, 4.1-10.4 cm broad. Petiole V3-V2 the sterile leaf length, with scales spreading, blackish-maroon, ca. 5 mm long, and with abundant, erect, minute glands and appressed stellate trichomidia. Lamina lan- ceolate, chartaceous, apex acuminate to caudate, base truncate to subcordate. Veins barely evident, free, ca. 1 mm apart, at 70 degree angle. Hyda- thodes lacking. Lamina scales limited to costa, 4- 5 mm long, and margin, 2-3 mm long, adaxially only on margin, both surfaces with stellate tricho- midia. Fertile leaves slightly shorter than the ster- ile, the petiole ca. V2 the fertile leaf length, the lamina nearly as broad as the sterile, base rounded to subtruncate; intersporangial scales lacking. Epiphyte in wet forests, 2050-3250 m, Ama- zonas, San Martin, Huanuco, Cuzco. Colombia; Ecuador; Peru. Elaphoglossum barbatum is distinct from E. er- inaceum by the truncate base and many tricho- midia, even onto the upper petiole. Amazonas: Bagua, E of La Peca, Barbour2757 (F). San Martin: Mariscal Caceres, Rio Abiseo Nat. Park, Young 3394 (NY), Young & Leon 5006 (USM). Huanuco: Hu- acachi, near Mima, Macbride 4690 (F). Cuzco: Urubam- ba, Machu Picchu, Vargas 16822 (GH), Cook & Gilbert 849, S52(us). 1 2. Elaphoglossum blepharoglottis Mickel, sp. nov. Elaphoglosso erinaceo affinis, ab ea lamina anguste elliptica basi anguste cuneata, costa adaxialiter glabra, abaxialiter squamis subulatis nigris dense induta, mar- gine squamis appressis 1-1.5 mm longis nigris ciliata, squamisque rhizomatis fusco-aurantiacis crispis diversa. Stem compact, 6-10 mm in diameter, scales lin- ear-lanceolate, orange-brown, crispate, entire, ca. 10 mm long. Phyllopodia indistinct. Leaves fas- ciculate, 46-68 cm long, 4.6-6.3 cm broad. Petiole ca. '/<, the sterile leaf length, scales black, subulate, 4-6 mm long, dense, patent. Lamina narrowly el- liptic, chartaceous, apex acuminate, base attenu- ate. Veins evident, 1-1.5 mm apart, at ca. 75 de- gree angle. Hydathodes lacking. Lamina scales limited to abaxial costa, black, subulate, spread- ing, 2-3 mm long, and margin, black, subulate, to 1 mm long. Fertile leaves much shorter than the sterile (30 cm long), petiole about !/3 the fertile leaf length, lamina similar in shape to the sterile but narrower (ca. 2.4 cm); intersporangial scales lack- ing. TYPE— Peru, Huanuco, Muna, 23 May-4 June 1923, Bryan 534 (holotype, F!). Endemic. Epiphytic in dry woods, 2150 m, Huanuco. Elaphoglossum blepharoglottis is in the E. eri- naceum complex, with dark subulate scales and no hydathodes, but is distinct in the dense scales of the petiole and abaxial costa. Thus far known only from the type. 13. Elaphoglossum camptolepis Mickel, sp. nov. Rhizomate longe repenti necnon lamina lanceolata squamis fuscis curvatis instructa inter affines determi- nanda. Stem long-creeping, 1-1.5 mm in diameter, scales linear, lustrous, spreading, recurved, black- ish-maroon, with short, sparse (often curved) teeth. Phyllopodia present, often indistinct or covered with scales. Leaves 1-3 cm apart, 6-12 cm long, 1.6-2.4 cm broad. Petiole about V4 the sterile leaf length, with scales linear, blackish maroon, lus- trous, spreading and recurved, sparsely serrulate, 1-2 mm long. Lamina lanceolate, chartaceous, apex acute to acuminate, base rounded. Veins indis- tinct, free, 1-2 mm apart, at ca. 70 degree angle. Hydathodes present but inconspicuous, as a pit, but not dark. Lamina scales on both surfaces as on petiole, strongly curved, serrulate, more con- centrated on costa, straighter and longer on upper surface, 1-2 mm long abaxially, 2-3 mm long adaxially, abaxial surface also with resinous dots. Fertile leaves not seen. TYPE— Peru, San Martin, Mariscal Caceres, Rio Abiseo Nat. Park, Puerta del Monte, Young 1913 (holotype, NY!; isotype, USM!). Endemic. Terrestrial(?), above timberline, 3500 m, San Martin. Thus far known only from the type. 14. Elaphoglossum cardenasii Wagner, Bull. Tor- rey Bot. Club 81: 62. 1954. TYPE: Bolivia, 126 FIELDIANA: BOTANY Dept. Cochabamba, Prov. Chapare, about Km. 1 20 Cochabamba to Chimore, Cardenas 795 (holotype, GH!). Stem short-creeping, 4-10 mm in diameter, scales flat, appressed, ovate-acuminate, dull brown, 2-3 mm long, entire. Phyllopodia lacking. Leaves approximate, 33-39 cm long. Petiole '/2-2/3 the ster- ile leaf length, with scales light brown, appressed, 1-2 mm long, overlapping in lower part, scattered distally. Lamina pedately lobed, 1 4-20 cm broad, chartaceous, apex acuminate, 2-3 times basally divided lateral lobe at base of each side of lamina; margin slightly undulate, shallowly crenulate. Veins evident, free, 1-2 mm apart, at ca. 40-60 degree angle. Hydathodes evident. Lamina scales deltate adaxially, 0.2-0.5 mm long, abaxially consisting of squamules to stellate trichomidia, costal scales lanceolate, 0.5-1 mm long. Fertile leaves shorter than the sterile, petiole 4/5-9/i0 the fertile leaf length, the lamina similar in architecture but smaller than the sterile lamina, 4-8 cm long, 5-10 mm broad; intersporangial scales lacking. ceolate to elliptic or ovate-lanceolate, chartaceous, apex acuminate, base cuneate to subtruncate. Veins inconspicuous, free, ca. 1 mm apart, at 70-80 de- gree angle. Hydathodes conspicuous. Lamina scales on both surfaces sparse, 0.5-1 mm long, awl- shaped, dark, lustrous. Fertile leaves longer than the sterile, petiole % the fertile leaf length, lamina narrowly lanceolate; intersporangial scales lack- ing. Terrestrial in wet forests, (7 50-) 19 50-3 300 m, San Martin, Huanuco, Pasco, Cuzco. Costa Rica; Panama; Colombia; Ecuador; Peru; Bolivia. San Martin: Mariscal Caceres, NW corner of Rio Abi- seo Nat. Park, Puerta de Monte, Young & Leon 4463 (USM). Huanuco: Tambo de Vaca, Macbride 4451 (F, us). Pasco: Prov. Oxapampa, Oxapampa, van der Werff8610 (uc). Prov. Oxapampa, San Alberto, van der Werff8430 (uc) Oxapampa, Rio San Alberto, Abra Esperanza, Fos- ter & d'Achille 10294 (F, GH). Cuzco: Paucartambo, at Km 1 42 on Paucartambo-Manu road, Manu Nat. Park, Skog & Skog 5203 (NY, us). Epiphytic in wet forests, ca. 3050 m, Cuzco. Peru; Bolivia. This is the best known of the dissected forms of Elaphoglossum (excluding Peltapteris). It is ap- parently fixed genetically as it is found in several localities. Other kinds of dissection (cresting, cru- ciform) are found in several species, but not with any consistency. A pedate leaf has been found in E. simulans, which see. I have not seen the Pe- ruvian specimen, and conceivably it is a misiden- tified specimen of E. simulans. Cuzco: Alturas de Sicre, Biies (cuz). 1 5. Elaphoglossum castaneum (Baker) Diels, Nat. Pflanzenfam. 1(4): 333. 1899. Acrostichum castaneum Baker, J. Linn. Soc. Bot. 15: 166. 1 877. TYPE: Ecuador in sylv. vulc. Corazon, a Bango, Sodiro, Aug 1873 (holotype, K; isotypes, B!, s!, us!). Stem moderately creeping, 3-5 mm in diameter, scales brown to black, linear-lanceolate, lustrous, ca. 2 mm long, older parts of stem often with scattered, arched scales less than 1 mm long. Phyl- lopodia lacking. Leaves slightly apart, 27-55 cm long, 3.0-7.8 cm broad. Petiole V3-2/3 the sterile leaf length, glabrous except at base. Lamina lan- 16. Elaphoglossum c hi nodes Mickel, sp. nov. Lamina lineari abaxialiter squamis aurantiacis inter- textis indua recognoscenda. Stem short-creeping, 2-3 mm in diameter, scales linear-lanceolate, lustrous, brown, ca. 3 mm long, entire. Phyllopodia distinct. Leaves 1-2 mm apart, 30-45 cm long, 0.8-1.0 cm broad. Petiole V4-V3 the sterile leaf length, with dark brown, linear, lustrous, entire scales, at petiole base ascending, 2-3 mm long, distally appressed, tan, 0.5-1 mm long, entire. Lamina linear, chartaceous, apex acu- minate, base attenuate. Veins evident, free, 1 mm apart, at ca. 70 degree angle. Hydathodes lacking. Lamina scales abaxially matted, white, peltate, round, dentate, ca. 0.5 mm across, membranous, most delicate in the genus, costal scales scattered, dark or tan, appressed as on petiole, adaxially scat- tered, white, flaky, ovate, entire, especially near margin. Fertile leaves unknown. TYPE— Peru, Ayacucho, Ccarrapa, between Huanta & Rio Apurimac, Killip & Smith 22391 (holotype, NY!; isotype, us!). Endemic. On moist banks and rotten logs in wet forests, 1500-2380 m, Amazonas, Ayacucho, Cuzco. This forms a complex with E. eatonianum (E. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 127 G. Britt.) C. Chr., of Ecuador, and E. vittarioides, all having a linear leaf, linear, lustrous brown stem scales, and dark-streaked costal scales. Amazonas: Bongara, 4 km N of Pomacochas on road to Rioja, Knapp et at 7487 (MO). Cuzco: Quillabamba, Santa Teresa, 0.5 km W of La Playa, Peyton & Peyton 1149(Gn). 17. Elaphoglossum ciliatum (Presl) Moore, Index fil. 8: 353.1857, based on Olfersia ciliata Presl. Acrostichum ciliatum Presl, Reliq. haenk. 1: 15. 1825, not. Desv. 1811. TYPE: Peru, ad Huanuco, Haenke s.n. (holotype, PR or PRC). Olfersia ciliata Presl, Tent. Pterid. 234. 1836, nom. nov. (Art. 72) for A. ciliatum Presl, not Desv. Acrostichum preslianum Fee, Mem. foug. 2: 46. 1845, nom. nov. for Acrostichum ciliatum Presl, not Desv. Elaphoglossum preslianum (Fee) Christ, Monogr. Ela- phoglossum 95. 1899. Stem long-creeping, ca. 2-3 mm in diameter, glutinous, scales lacking except for a few resinous brown ones 1 mm long. Phyllopodia 2-4 cm long. Leaves fasciculate, 1-10 mm apart, 26-78 cm long, 1.2-2.3 cm broad. Petiole V3 the sterile leaf length, with sparse brown scales, these widely spreading, 1 mm long, entire. Lamina linear-elliptic, char- taceous to subcoriaceous, apex acute to acuminate, base cuneate. Veins obscure, free, 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales as on petiole: sparse, small, brown, entire, ovate, on abaxial costa and margin, often glan- dular-punctate below. Fertile leaf longer than the sterile, petiole % the leaf length, the lamina linear, 6-8 mm wide; intersporangial scales lacking. Epiphytic and terrestrial in wet forests, 400- 2650 m, Huanuco, Junin, Pasco, Ucayali, Cuzco. Huanuco: Carpish Divide, C. Landeman 5157 (K). Leoncio Prado, La Divisoria, Plantation Azul, Ridoutt, 3 Aug. 1942 (USM). Cushi, Macbride 4812 (F, us). Muna, Bryan 532 (F, us). Pasco: Prov. Oxapampa, Abra los Mellizos, Skog et al. 5042 (us). Junin: Tarma, Chan- chamayo, Esposto 686 (GH). Pichis Trail, Enenas, Killip & Smith 25668 (NY, us). Chanchamayo Valley, C. Schunke 136 (F, us). Schunke Hacienda, above San Ra- mon, C. Schunke A 145 (us). Tarma, Agua Dulce, Woyt- kowski 35494 (uc). Utcuyacu, Woytkowski 494 (us). Ucayali (as Loreto): Coronel Portillo, cerca a la Divi- soria, Ridoutt (USM: 12470), 3 Aug. 1942 (GH). Cuzco: Paucartambo, Sta. Isabel, Kosnipata, C. Vargas 23014 (GH). 18. Elaphoglossum concinnum Mickel, sp. nov. Ab E. squamipedi frondibus minoribus, lamina squamis obscurioribus vestitis distal. Stem long-creeping, 0.5-0.8 mm in diameter, scales linear-lanceolate, orange-tan, ca. 3 mm long, entire. Phyllopodia lacking. Leaves 1-5 mm apart, 1.3-2.6 (3.5) cm long, 0.5-0.9 cm broad. Petiole %-3/5 the sterile leaf length, with tan scales widely spreading, 1-2 mm long, entire. Lamina ovate and suborbicular to oblanceolate, subcoriaceous, apex obtuse, base narrowly to broadly cuneate. Veins obscure, free, 1 mm apart, at 60-70 degree angle. Hydathodes lacking. Lamina scales dark brown to black, sclerotic, scattered, especially abaxially, lin- ear-deltate, subentire, with sparse irregular teeth, adaxial ones fewer, paler. Fertile leaves longer than the sterile, petiole % the leaf length, lamina ovate- deltate to suborbicular; intersporangial scales black, sclerotic. TYPE— Peru, San Martin, Mariscal Caceres, Rio Abiseo Nat. Park, near Las Papayas ruins, Young 130 1 (holotype, NY!). Endemic. Epiphytic in wet forests, 2600-3750 m, Amazonas, San Martin. Amazonas: Prov. Bongara, 2-4 km WSW of Poma- cocha, Wurdack 874 (us). San Martin: Mariscal Caceres, trail between La Playa camp & Papayas camp, Rio Abi- seo Nat. Park, Young & Leon 4960 (USM). 19. Elaphoglossum craspedotum Copel., Univ. Calif. Publ. Bot. 19: 303. t. 63. 1941. TYPE: Peru, Huanuco, Dist. Churubamba, Hacienda Mercedes, Mexia 8177 (holotype, uc!; iso- types, F!, GH!, us!). Stem compact, horizontal, ca. 8 mm in diam- eter, scales linear-lanceolate, lustrous, dark red- brown, 4—6 mm long, entire. Phyllopodia evident, ca. 4 cm long. Leaves fasciculate, 60-75 cm long, 3.4—4.6 cm broad. Petiole '/5 the sterile leaf length, with tan to orange scales appressed to spreading, 1-2 mm long, with very thin, sparse hair- teeth. Lamina narrowly elliptic, chartaceous, apex acu- minate, base attenuate. Veins evident, free, ca. 1 mm apart, at 70-80 degree angle. Hydathodes lacking. Lamina scales adaxially white, substellate, scattered, ca. 0.2-0.5 mm long, abaxial laminar surface punctate, glandular, costal scales pale, nearly round, ciliate, 0.5 mm long. Fertile leaf 128 FIELDIANA: BOTANY shorter than the sterile, petiole 3/5 the fertile leaf length, the lamina linear, 10-13 mm broad; in- tersporangial scales lacking. Endemic. Epiphytic in wet forest, 1200 m, Huanuco. Thus far known only from the type. 20. Elaphoglossum cuspidatum (Willd.) Moore, Index fil. xvi. 1857. Acrostichum cuspidatum Willd., Sp. pi. ed. 4, 5: 106. 1810. TYPE: Venezuela, Caracas, Bredemeyer (holotype, B!, Herb. Willd. 19516). Stem short-creeping, ca. 4-8 mm in diameter, scales linear-lanceolate, lustrous, black-brown, 2- 3 mm long, entire. Phyllopodia distinct. Leaves 1- 10 mm apart, 28-75 cm long, 1.9-4.6 cm broad. Petiole l/3-V2 the sterile leaf length, with ovate to lanceolate scales, these black with white, ciliate margin, appressed, 1-3 mm long. Lamina narrow- ly elliptic, chartaceous, apex cuspidate (to acu- minate), base broadly to narrowly cuneate. Veins obscure, free, 1 mm apart, at ca. 80 degree angle. Hydathodes lacking. Lamina scales orange, abax- ially consisting of a felt-like mass of round, peltate ciliate scales 0.5 mm long and ovate-lanceolate scales 1 mm long, the larger ones with maroon center, costal scales ovate to ovate-lanceolate, red- black with pale ciliate margin as on petiole, lamina adaxially glabrous or with scales scattered to dense, round, peltate, ciliate, pale, ca. 0.7 mm in diam- eter. Fertile leaves shorter than the sterile, petiole 2/3 the fertile leaf length, the lamina slightly nar- rower than the sterile; intersporangial scales lack- ing. Epiphytic, rarely terrestrial in wet forests, (600) 1200-3800 m, Piura to Amazonas, south to Cuz- co. Costa Rica; Venezuela to Bolivia. Piura: Huancabamba, Loma redonda (Sapalache- Chinjuela), Sagdstegui 10193 (NY, HUT, uc), Ayabaca, Soukup 4315 (us). Cajamarca: Chota, La Palma-Dist. Conchan, A. Diaz & Laos 5730 (USM). Chota, Chota- Tacabamba road, D. Smith & Vasquez M. 3556 (uc). Amazonas: Bongara, 3 km S Pomacocha, Wurdack 1002 (F, GH, NY). NW of Jumbilla, Soukup 5245 (GH). La Libertad: Otuzco, Huaranchal, Sagdstegui 0203 (GH). San Martin: Lamas, Alonso de Alverado, San Juan de Pacayzapa, /. Schunke V. 5891 (F). Palo Blanco, J. Schunke V. 5673 (F). Zepelacio, near Moyobamba, Klug 3503a (us). Ancash: Yungay, Quebrada Llanganuco, Perez 99 (USM). Pasco: Oxapampa, Soukup 3355 (F, GH, us). Oxapampa, near Villa Rica, Chacra del Sr. Simon Es- pilco, van der Werff et al. 8317 (MO). Junin: Chancha- mayo Valley, C. Schunke 173 (F). Tarma, Cumbre Ya- cunay ridge above La Merced, Hutchinson 1978 (F, GH, uc, us). Tarma, Agua Dulce, Woytkowski 35498 (uc). Ucayali (as Loreto): Coronel Portillo, La Divisoria, Fer- reyra 1049 (GH). Cuzco: Prov. La Convention, Quello- huno-Chirumbia, C. Vargas 11377 (GH). La Conven- cion, NE from Hacienda Luisiana & Apurimac River (Cordillera Vilcabamba), Dudley 1 1 147 (GH). Urubam- ba, Machu Picchu, Sounders 462A (F). Los Palmitos, Cabecero Rio Karibeni, Biies 1959 (us). 21. Elaphoglossum decoratum (Kunze) Moore, Index, fil. 8. 1857. Acrostichum decoratum Kunze, Linnaea 9: 25. 1835. TYPE: Peru, Huanuco, Pampayaco (Pampaya- cu), PoeppigDiar. 1 134, July 1829 (holotype, LZ, destroyed; isotype, MO!) Stem short-creeping, to 6 mm in diameter, scales linear, orange, crispate, to 1 6 mm long, entire or with very sparse, small, irregular teeth. Phyllo- podia lacking. Leaves fasciculate, to 70 cm long, 1 2 cm broad. Petiole V4-V3 the sterile leaf length, scales orange, ovate, spreading, with obtuse tip, to 1 0 mm long. Lamina elliptic, apex cuspidate, base broadly cuneate; texture thin. Veins evident, free (with rare anastomoses), ca. 2 mm apart, at ca. 70 degree angle. Hydathodes lacking. Lamina scales limited to the costa and margin, forming a con- tinuous band of overlapping, cordate, golden scales on the margin, elliptic on the costa. Fertile leaves rare, shorter than the sterile ones, margins lacking scales; intersporangial scales lacking. Epiphytic in wet forests, 800-1 500 m, Huanuco, Cuzco. Guatemala; Costa Rica; Panama; West Indies; Colombia and Venezuela to Bolivia. Huanuco: Rio Llullapichis watershed, Cerros del Sira, Dudley 13008 (GH). Sira mountains, Seidenschwarz 442/1 (us, GH). Cuzco: Paucartambo, Cosnipata Valley, Rio Tono, Wachter 132 (F). 22. Elaphoglossum dichroum Mickel, sp. nov. Stipitis costaeque squamis dentatis bicoloribus, squamisque dentatis laminae marginem versus congestis praestans. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 129 Stem compact, 5 mm in diameter, scales linear, dull, tan with blackish streaks, ca. 10 mm long, entire. Phyllopodia present. Leaves approximate, 65-70 cm long, 5.5-6.0 cm broad. Petiole '/3-2/5 the sterile leaf length, scales black-centered with white margin, larger ones ovate-lanceolate, 2-6 mm long, slightly spreading, smaller ones 1 mm long, appressed. Lamina narrowly elliptic, sub- coriaceous, apex acute, base truncate. Veins evi- dent, 1 mm apart, at 75 degree angle. Hydathodes lacking. Lamina surface adaxially glabrescent, with concentration of 1 mm hair-toothed scales at mar- gin, abaxially glabrous, indistinctly gland-dotted, or with scattered stellate trichomes, the scales re- duced to 1-3 rays, costal scales 1-2 mm long, erose- denticulate, black-centered with white margin. Fertile leaves shorter than the sterile, petiole 3/5 the leaf length, lamina linear, 1 cm broad; interspor- angial scales abundant, ciliate. TYPE— Peru, San Martin, Mariscal Caceres, Par- que Nac. Rio Abiseo, Puerta del Monte, Leon & Young 1510 (holotype, USM!). Endemic. Epiphytic in wet forests, elevation un- known, San Martin. This species is closely allied to E. oculatum, as evidenced by the marginal concentration of scales, ciliate intersporangial scales, bicolorous petiole and costal scales, the nearly naked lamina surfaces with scales reduced to stellate trichomes, but it is much larger, the apex is acute rather than obtuse, and the fertile lamina is linear and shorter than the sterile. Thus far known only from the type. 23. Elaphoglossum discolor (Kuhn) C. Chr., In- dex fil. 306. 1905. Acrostichum discolor Kuhn, Linnaea 36: 53. 1869. SYNTYPES: Brazil, San Gabriel ad Rio Negro, Spruce 2309, 2245; Brazil, Panure, Spruce 2869 (none located, all B?). Stem compact, horizontal, ca. 3-4 mm in di- ameter, scales linear-lanceolate, reddish-orange, ca. 3 mm long, with irregular weak teeth. Phyl- lopodia evident. Leaves fasciculate, ca. 30 cm long, 4.0-4.5 cm broad. Petiole V2 the leaf length, with orange to blackish, lanceolate to linear scales to trichomidia, scales ascending, 2-3 mm long, with irregular teeth. Lamina elliptic, chartaceous, apex acuminate, base cuneate and abruptly decurrent. Veins evident, free, 1 mm apart, at ca. 80 degree angle. Hydathodes lacking. Lamina scales linear with long, sparse teeth on both sides of lamina, 1- 1.5 mm long. Fertile leaves shorter than the sterile, petiole 3/5 the fertile leaf length, the lamina nar- rowly elliptic, 2 cm broad, acute; intersporangial scales lacking. Epiphytic and terrestrial, low forest on white sand and wet forests, 150-980 m, San Martin, Loreto, Pasco, Junin. Venezuela; Colombia; Ecuador; Peru; Brazil. Elaphoglossum discolor can be distinguished by the short stem, its scales linear and reddish-orange with irregular processes, the petiole scales scat- tered, lamina scales scattered, linear, flexuous, dis- sected, and the fertile leaf longer than the sterile. San Martin: Rioja, near Rioja and Naranjos, Knapp et al. 7444 (uc). Loreto: Maynas, Iquitos, Carretera de Varillal km 6 de Quisto Cocha al caserio de Varillal, Rimachi Y. 7844 (NY). Prov. Maynas, Distr. Iquitos, Puerto Almendras, van der Werffet al 9788, 9819, 9820 (uc). Prov. Requena, Jenaro Herrera, van der Werffet al. 10001 (uc). Vicinity of Iquitos, Revilla 3758 (NY), 4322 (uc). Pasco: Oxapampa, Palcazu valley, D. Smith 3852 (uc). Junin: East of Quimiri Bridge near La Mer- ced, Killip & Smith 23850 (us). 24. Elaphoglossum diversifrons C. Chr., Index fil. Suppl. 1:42. 1913. Acrostichum diversifolium Sodiro, Sert. fl. ecuad. II. 32. 1908, not Blume, 1828. SYNTYPES: Ecua- dor, in silvis suband. vulc. Corazon Sodiro; Ata- catzo, Sodiro (P!, s!); "itemque in loc. praeruptis opacis prope Quito," Sodiro (P!). Stem short-creeping, 1.5-2 mm in diameter, scales linear, lustrous, castaneous, 3—4 mm long, denticulate. Phyllopodia lacking. Leaves approx- imate, 15-24 cm long, 2.0-2.4 cm broad. Petiole V3-l/2 the sterile leaf length, scales castaneous, lus- trous, subulate, patent, denticulate, 1-2 mm long. Lamina elliptic, chartaceous, apex acuminate, base broadly cuneate to rounded. Veins evident, 2-2.5 mm apart, at ca. 60-degree angle. Hydathodes present. Lamina scales evenly scattered on both surfaces, subulate, castaneous, 1-2 mm long. Fer- tile leaves slightly longer than the sterile, petiole about % the leaf length, lamina lanceolate; inter- sporangial scales lacking. Epiphytic in wet forests, 2700 m, Huancavelica. Ecuador; Peru. Huancavelica: Tayacaja, Marcavalle, Tovar4756 (GH). 130 FIELDIANA: BOTANY 25. Elaphoglossum elegantipes Mickel, sp. nov. Rhizomatis squamae fuscae cum stipite gracile notulas diagnosticas proferunt. Stem compact, horizontal, 3-4 mm in diameter, scales linear-lanceolate, dull dark brown, 3-5 mm long, with hairlike processes. Phyllopodia present. Leaves fasciculate, 24—42 cm long, 2.7—4.8 cm broad. Petiole l/4-l/3 the sterile leaf length, glabrous. Lamina elliptic, chartaceous, apex long-acumi- nate, base attenuate. Veins evident, free, 1 mm apart, at 70-80 degree angle. Hydathodes lacking. Lamina indument lacking ahaxialK . but occasion- ally some minute, orange, stellate trichomidia adaxially. Fertile leaves longer or slightly shorter than the sterile, petiole 3/5-% the fertile leaf length, lamina slender, 1.4-2.0 cm broad; intersporangial scales lacking. TYPE— Peru, Cuzco, Paucartambo, Cosfiipata Valley, Rio Tono, N. of Patria, Wachter 138 (ho- lotype, F!; isotype, GH!), Endemic. Epiphytic, rarely terrestrial, in wet forests, 300-2400 m, Amazonas, Huanuco, Pasco, Junin, Ucayali, Cuzco, Madre de Dios, Puno. Amazonas: Bagua, ca. 20 km E of La Peca, Barbour 2766 (uc). Huanuco: SW slope of Rio Llullapichis wa- tershed on ascent of Cerros del Sira, Dudley 13134 (GH). Carpish, Ellenberg 3931 (GH). Pasco: Oxapampa, Pal- cazu valley, between Iscozacin & Villa America, D. Smith 3852 (MO). Oxapampa, Gran Pajonal, 2-3 km N of Che- quitavo, D. Smith 5087 (MO, uc). Oxapampa, 4-5 km N of Mallampampa, D. Smith & Canne 5795 (MO). 2- 4 km N of Mallampampa, D. Smith & Canne 5862 (MO). Oxapampa, Palcazu, Rio Alto Iscozacin, Ozuz. Foster 9950 (F). Chontabamba, Abra "La Suiza", camino al Rio Chontabamba, Leon et al. 979 (F). Junin: La Merced- Chanchamayo, Soukup 1104 (F). Satipo, Pichanaki, Leon 229 (USM). Chanchamayo Valley, C. Schunke 174 (F). Ucayali (as Loreto): Coronel Portillo, Chacra de Cesar Vela, J. Schunke V. 5493 (F). Cuzco: Paucartambo, Cosnipata Valley, Rio Tono, N of Patria, Wachter 169 (F, GH). Madre de Dios: Manu, Cerro de Pantiacolla, Rio Palotoa, 10-15 km NNW of Shintuya, Foster 10752 (F). Puno: La Pampa, Rio Tavara, Watkins in 1916 (us). 26. Elaphoglossum engelii (Karsten) Christ, Monogr. Elaphoglossum 81. 1899. Acrosti- chum engelii Karsten, Fl. Columb. 1 : 1 1 8, t. 59. 1860. TYPE: Colombia, cordillera Meri- densis, Karsten (holotype, B?). Stem compact, horizontal, 5-8 mm in diameter, scales linear, lustrous, maroon, 8-15 mm long, entire to irregularly hair-toothed. Phyllopodia present but hidden by scales. Leaves fasciculate, 17-47 cm long, 1.4-2.7 cm broad. Petiole V3-V2 the sterile leaf length, scales orange, appressed, 3- 7 mm long, ciliate. Lamina narrowly elliptic, co- riaceous, apex obtuse, base broadly cuneate to rounded. Veins obscure, free, 1 mm apart, at ca. 70 degree angle. Hydathodes lacking. Lamina scales abaxially lanceolate, orange, ciliate, 2-3 mm long, dense to loose, costal scales broader, ovate-lan- ceolate, orange to dark centered, 3-4 mm long, adaxially same as below. Fertile leaves longer than the sterile, petiole 2/3 the fertile leaf length, lamina narrower than the sterile; intersporangial scales lacking (costal scales ovate-lanceolate, ciliate, darker center plus small round ciliate ones). Terrestrial, epipetric, and epiphytic, in erica- ceous heath, shrubby grassland, puna, rocky road- banks, talus slopes, elfin forest, and cloud forests, (1800-)2800-4600 m, Piura to Amazonas, south- ward to Ayacucho and Puno. Elaphoglossum engelii is in the E. muscosum group, differing from others in that complex by the petiole scales being long-ciliate rather than erose-denticulate. A name sometimes used for this species is E. denticulatum Ruiz & Pavon. I have not found the reference or description (the epithet would be under Acrostichum), but if it does apply, it would have priority. Piura: Huancabamba, above Canchaque on road to Huancabamba, Hutchinson 1651 (uc). Lambayeque: Ferrenafe, ca. 7 km NW of Incahuasi, Dillon & Skillman 4116 (F, GH). Ferrenafe, Incahuasi, Laguna Temlami- nara, Sagdstegui 12780 (uc). Cajamarca: Cajabamba, Cajabamba-Luchubamba, Sagastegui 11200 (GH, NY). Amazonas: Chachapoyas, Cerros Calla Calla, above Lei- mebamba on road to Balsas, Hutchinson & Wright 5567 (F, GH, NY, uc, us). Entre donila y Cohechan, Soukup 4133 (F). La Libertad: Sanchez, Carrion, road from Tru- jillo to Huamachuco, Dimmitt 1135 (NY). San Martin: Mariscal Caceres, Rio Abiseo Nat. Park, Young & Wat- son 3508 (NY). Ancash: Huari, Huascaran Nat. Park, D. Smith et al. 12382 (F, NY). Yungay, Quebrada de Llan- ganuco, Perez 85 (USM). Huanuco: Mito, Bryan 370 (F). Huanuco, Carpish, Coronado 74 (GH, uc). Junin: Chan- chamayo, Mina Pichita, above San Ramon, van der Werff et al. 8658 (MO, uc). Ayacucho: Pampalca between Huan- ta & Rio Apurimac, Killip & Smith 23283 (NY, us). Apurimac: 5 km N of Huancarama, West 3801 (uc). Cuzco: Urubamba, Machu Picchu, in Llactapampa on Aobamba River, Peyton & Peyton 815 (GH, MO). Pilla- huata, Cerro de Cusilluyoc, Pennell 14125 (GH, NY, us). Puno: Sandia, S of Limbani, Metcalf 30453 (GH), also Ruiz 4570 (53 crossed out) (us ex Herb. Lambertii). 27. Elaphoglossum ensiforme Mickel, sp. nov. Ab E. lloensi squamis stipitis laminaeque pallidis et lamina graciliori apicem versus gradatim atenuata distat. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 131 Stem long-creeping, 1-1.5 mm in diameter, scales linear-lanceolate, dull tan, 3-5 mm long, entire. Phyllopodia lacking. Leaves fasciculate, (9-)21-26 cm long, 1.4-2.0 cm broad. Petiole 2/5- 3/5 the sterile leaf length, with orange scales ap- pressed to ascending, 3-4 mm long, subentire with occasional irregular teeth. Lamina narrowly ellip- tic, chartaceous, apex acuminate, base narrowly cuneate. Veins evident, free, 1-1.3 mm apart, at ca. 60 degree angle. Hydathodes lacking. Lamina scales scattered, linear-deltate, tan, 1-2 mm long, hastate abaxially, sparse adaxially. Fertile leaves not known. TYPE— Peru, Cuzco, Urubamba, Machu Picchu, in Llactapampa, a small pampa below Palcay on the Aobamba River, Peyton & Peyton 814 (holo- type, MO!). Endemic. Epiphytic in wet forests, 3300 m, Cuz- co. Elaphoglossum license (Hooker) Moore (Ven- ezuela, Colombia, Ecuador) has lamina propor- tionally broader, with more caudate apex, lamina scales darker, and petiole scales with black base. Cuzco: Urubamba, Machu Picchu, in Llactapampa, a small pampa below Palcay on the Aobamba River, Pey- ton & Peyton 811 (MO). 28. Elaphoglossum erinaceum (Fee) Moore. Index fil. 9. 1857. Figure 25c. Acrostichum erinaceum Fee, Mem. foug. 2: 41. 1845. SYNTYPES: Guadaloupe, riviere de Saint-Louis, de Thiouville (P!); Brazil, Gaudichaud; Mexico, Karwinsky, in 1827. Elaphoglossum erinaceum var. boliviensis Rosenst., Repert. Spec. Nov. Regni Veg. 12: 476. 1913. TYPE: Bolivia, Polo-Polo bei Coroico, Nordyun- gas, Buchtien 3470 (isotype, us!). Stem short creeping, 10-20 mm in diameter, scales linear, castaneous to orange, to 1 7 mm long, plane to crispate, entire or with very irregular teeth near the tip. Phyllopodia lacking. Leaves fascicu- late, 30-52 cm long, 3.7-9.2 cm broad. Petiole '/3- '/2 the sterile leaf length, scales dark reddish brown, subulate, widely spreading, to 5 mm long, also with minute erect glandular trichomes on petiole and mid vein. Lamina lanceolate to elliptic, apex acuminate to caudate, base rounded (rarely cu- neate). Veins evident, free, 1-1.5 mm apart, at 70- 80-degree angle. Hydathodes lacking. Lamina scales subulate, dark reddish brown, often ap- pearing black, mostly on the midvein and spread- ing from the margin, also with minute, simple, branched, or stellate trichomidia on the abaxial surface. Fertile leaves slightly shorter than the ster- ile, petiole l/T,-l/2 the fertile leaf length, lamina nar- rowly elliptic, with scattered dark subulate scales on the abaxial costa, intersporangial scales lacking. Epiphytic, rarely terrestrial, in wet forests, 700- 3325 m, Cajamarca, Amazonas, San Martin, Huanuco, Madre de Dios. Mexico to Panama; West Indies; Venezuela and Colombia to Bolivia and Brazil. Elaphoglossum erinaceum belongs to a very per- plexing group. The Peruvian collections are some- what larger and have more rounded lamina bases than the Mexican and West Indian material, gen- erally stout petioles (stouter than in Central Amer- ica), 2.5 mm thick, and broad laminae and longer fertile leaves. Cajamarca: Colasay, Woytkowski 7019 (us). Amazo- nas: Bagua, E of La Peca, Barbour 2528 (MO), 2757 (uc). Serrania de Bagua, 1 7 km E of La Peca, Gentry et al. 22977 (us). San Martin: Mariscal Caceres, Chochos val- ley, NW corner Rio Abiseo Nat. Park, Young 263 3 (NY). Young & Watson 3468 (NY). Huanuco: Huacachi, Es- tacion near Muna, Macbride 4690 (us). Ucayali (as Lore- to): Coronel Portillo, Tingo Maria-Pucallpa road, Skog et al. 5154 (us). Madre de Dios: Manu, Cerro de Pan- tiacolla, Rio Palotoa, R. Foster 10849 (F). Department unknown: C. Schunke, Aug 1908 (uc). 29. Elaphoglossum erythrolepis (Fee) Moore, In- dex fil. 9. 1857. Acrostichum erythrolepis Fee, Mem. foug. 2: 60. 1845. TYPE: Peru, Dombey (holotype, P!). Stem compact, horizontal, ca. 10 mm in di- ameter, scales linear, often forked into hairlike branches, lustrous, maroon, ca. 20 mm long, with scattered stiff hair-teeth. Phyllopodia distinct. Leaves fasciculate, 25-36 cm long, 1.4-2.2 cm broad. Petiole '/3-2/5 the leaf length, with light to dark reddish brown dense scales, these ascending, lanceolate, 3-5 mm long, ciliate. Lamina narrowly elliptic, coriaceous, apex acuminate, base cuneate. Veins not evident, free, 1 mm apart, at ca. 70- degree angle. Hydathodes lacking. Lamina scales abaxially dense, lanceolate, 1 .5-2 mm long, ciliate, reddish, costal scales 3-4 mm long, more ascend- ing, adaxially pale, ovate to lanceolate, longer and more ascending on costa, to glabrescent. Fertile 132 FIELDIANA: BOTANY leaves longer than the sterile, petiole V2 the leaf length, lamina as large as the sterile; intersporan- gial scales lacking. Terrestrial and epipetric, 2900 m, Huanuco, Ju- nin, Apurimac. Ecuador; Peru. Huanuco: Mito, Macbride & Featherstone 1392 (F, in part). Pachitea, Huapalla 2027 (USM). Along highway between Huanuco y Tingo Maria, at Rio Tulca, Croat 57899 (F). Chinchao, entre Huanuco y Tingo Maria, Ferreyra 16981 (GH). Junin: Manto to Yaupi, Woyt- kowski 6551 (GH). Tarma, Carpapata, Cerrate 2775 (GH). Huacapistana, Sandeman 4549 (K), Killip & Smith 24501, 24175 (NY). Apurimac: Abancay, 11-12 km E of Aban- cay, Hutchinson 1734 (uc). 30. Elaphoglossum eximium (Mett.) Christ, Monogr. Elaphoglossum. 107. 1899. Acrostichum eximium Mett., Ann. Sci. Nat. Bot. 5, 2: 199. 1864. TYPE: Venezuela, Colonia Tovar, Moritz 419 (isotype, us!). Stem short-creeping, ca. 3-4 mm in diameter, scales brown to castaneous, lustrous, linear, entire, 2-4 mm long. Phyllopodia lacking. Leaves fascic- ulate, to 44 cm long, 1.3 cm broad. Petiole short, Vio-'/ie the sterile leaf length, densely clothed with short, brown, subulate scales. Lamina linear, tex- ture thin, apex acuminate, base attenuate, margin slightly crenulate. Hydathodes evident. Lamina scales conspicuous but sparse on costa, smaller on margin, mostly between crenulations, minute punctae inconspicuous on lamina surfaces. Fertile leaves shorter than the sterile, petiole % the fertile leaf length, lamina smooth-margined, lanceolate to narrowly elliptic, apex apiculate, base cuneate to rounded; intersporangial scales lacking. Epiphytic or terrestrial in wet forests, 1 800-2320 m, San Martin, Huanuco, Cuzco. Mexico to Panama; Colombia; Venezuela; Ec- uador; Peru. Peruvian material differs from plants of Vene- zuela and Central America in having the fertile lamina base cuneate or rounded rather than trun- cate to subcordate and may represent a distinct taxon. San Martin: Rioja, Venceremos, D. Smith 4434 (uc). Huanuco: Huacachi, near Muna, Macbride 4131 (F, us). Cuzco: La Convention, Huayopata, Peyton & Peyton 885 (GH). 3 1 . Elaphoglossum flaccidum (Fee) Moore, Index fil. 356. 1862. Acrostichum flaccidum Fee, Mem. foug. 2: 35. /. 7,f. 2. 1 845. SYNTYPES: Guyana, Oyapoc, Leprieur, in 1834 (P!), Schomburgk 448 (B!); S. Yago de Cuba, Linden 2058 (P!); Guadalupe, Lezarede, I'Herminier (P!); Martinique, Heraud. Stem compact, horizontal, ca. 3-4 mm in di- ameter, scales linear-lanceolate, lustrous, blackish brown, 3-4 mm long, entire, somewhat deciduous. Phyllopodia evident. Leaves fasciculate, 23-37 cm long, 2.3-3.6 cm broad. Petiole ca. !/20 the sterile leaf length, with orange-tan scales appressed, 1-2 mm long, with weak hairlike teeth. Lamina nar- rowly elliptic, chartaceous, apex acuminate, base narrowly cuneate. Veins evident, free, 1-1.5 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales few, like those of petiole at costa base. Fertile leaves shorter than the sterile, petiole ca. '/2 the fertile leaf length, the lamina narrowly elliptic: intersporangial scales lacking. Epiphytic in wet forests, 600-1300 m, Ama- zonas, San Martin, Huanuco, Pasco, Junin, Madre de Dios. West Indies; Guianas; Venezuela: Colombia to Bolivia. Amazonas: Prov. Bagua, Quebrada Tambillo, Wur- dack 1989 (us). San Martin: Mariscal Caceres, Palo Blanco, al oeste del Puente, Tocache Nuevo, J. Schunke V. 5656 (F), Plowman 7469 (F). Tingo Maria, jungle E of Tingo Maria, Allard 20601, 20941 (us), 21388 (GH, us). Mishuyacu near Iquitos, Klug 1461 in part (F). Huanuco: Leoncio Prado, Rupa Rupa, al este de Tingo Maria, J. Schunke V. 5192 (F, uc). Pasco: Oxapampa, Quebrada Castilla on the Omaiz river, Leon & Young 1022 (F). Junin: Pichis Trail, San Nicolas Azupizu, Killip & Smith 26122 (us). Madre de Dios: Manu, Cerro de Pantiacolla, Rio Palotoa 10-15 km NW of Shintuya, Foster 10752 (GH), 10826 (F, GH). 32. Elaphoglossum fortipes Mickel, sp. nov. Rhizomate elongato repenti phyllopodiisque longis- simis praestans. Stem long-creeping, 2-3 mm in diameter, scales linear-lanceolate, lustrous, dark brown, 3-4 mm long, entire. Phyllopodia evident, 2-3.8 cm long. Leaves 1-2 cm apart, 12-15 cm long, 1.1-1.4 cm broad. Petiole '/4-2/5 the sterile leaf length, with brown scales ascending to widely spreading, 2-2.5 TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 133 mm long, with minute teeth. Lamina very nar- rowly elliptic, chartaceous, apex acuminate, base cuneate. Veins evident, free, 1 mm apart, at 70- 80-degree angle. Hydathodes lacking. Lamina scales on abaxial costa deltate-lanceolate, 1 mm long, adaxially and abaxially scattered, 0.5-1 mm long, orange, deltate to deltate-lanceolate, dentate. Fertile leaves unknown. TYPE— Peru (Cuzco), Puente de Nantto, Valle de Lares, Biles 1875 (us). Habitat unknown, 2000 m, Cuzco. Thus far known only from the type. 33. Elaphoglossum glabellum John Sm., London J. Bot. 1: 197. 1842. TYPE: British Guiana, Schomburgk 447 (holotype, BM!; isotype, us!). Acrostichum glabellum (John Sm.) Klotzsch, Linnaea 20:421. 1847. Stem short to moderately creeping, 2-3 mm in diameter, scales lanceolate, lustrous, dark brown to black, with pale brown base, margin, and tip, 1-2 mm long, entire or with a few weak teeth, scales tending to be somewhat deciduous, often leaving a partially naked stem. Phyllo podia dis- tinct, 4-8 mm long. Leaves spaced slightly apart, to 33 cm long, 0.5-1.0 (1.6) cm broad, the margin usually incurved. Petiole Vto-V5 the sterile leaf length, naked or with minute appressed scales. Lamina linear-elliptic, gradually acuminate at both ends, coriaceous. Veins obscure, free, ca. 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales lacking. Fertile leaves slightly short- er than the sterile, the petiole !/3-'/2 the fertile leaf length, the lamina ca. 1 0 mm broad; intersporan- gial scales lacking. Epiphytic in wet forests, 130-1800 m, Loreto, Huanuco, Junin, Cuzco, Madre de Dios. Mexico to Panama: West Indies; Trinidad; Guy- ana; Surinam; Venezuela; Colombia to Bolivia. Loreto: Maynas, Distr. Iquitos, Puerto Almendras, van der Werff9800 (uc). Maynas, Varillal, carretera a Nauta, Vdsquez 1211 (MO). Huanuco: SW slope of Rio Llullap- ichis watershed on ascent of Cerros del Sira, Dudley 13131 (GH). Junin: East of Quimiri Bridge, near La Merced, Killip & Smith 23993 (GH, NY, us). Satipo, Pichanaki, rodal del Proyecto Peruano-Aleman, Leon 239 (USM). Chanchamayo Valley, C. Schunke 115, 172 (F). Schunke Hacienda, above San Ramon, C. Schunke A 144, A147 (us). Cuzco: La Convencion, Cordillera Vilcabamba, near Rio Apurimac and Hacienda Luisiana, Dudley 10061 (GH). Madre de Dios: Manu, Cerro de Pantiacolla, Rio Palotoa, 10-15 km NNW of Shintuya, Foster 10691 (F). 34. Elaphoglossum glossophyllum Hieron., Hed- wigia 44: 180. 1905, nom. nov. for E. lin- guaeforme Hieron. 1 904, not Moore, 1857. Elaphoglossum linguaeforme Hieron., Bot. Jahrb. Syst. 34: 542. 1904, not Moore. 1857. SYNTYPES: Colombia, above Popayan, Lehmann 6944 (B!, us!), 6943 (not located); Antioquia, near Sonson, Lehmann 7582 (not located). Stem long-creeping, ca. 2-3 mm in diameter, scales lanceolate to ovate-lanceolate or deltate, dull orange or with lustrous black streaks, 4-7 mm long, with hairlike processes. Phyllopodia present. Leaves mostly 1-3 cm distant, 1 6-54 cm long, 2.2- 5.0 cm broad. Petiole V3-V2 the sterile leaf length, glabrous. Lamina lanceolate, coriaceous, apex acute to acuminate, base broadly cuneate to rounded. Veins obscure, free, 1 mm apart, at ca. 80-degree angle. Hydathodes lacking. Lamina scales lacking, or surfaces with stellate trichomidia to glabrous. Fertile leaf longer than the sterile, petiole !^-2/3 the leaf length, the lamina acute to obtuse, nearly equal to the sterile in size; intersporangial scales lacking. Epiphytic in wet forests, 2200-3500 m, Caja- marca, Amazonas, San Martin, Huanuco, Pasco, Ayacucho, Cuzco. Colombia; Ecuador; Peru. Cajamarca: San Miguel, Taulis Playa (Calquis), Mos- tacero et al. 1192 (F, NY). Chota, Chota-Tacabamba road, 14 km from Chota, D. Smith & Vdsquez 3554 (F, MO, uc). Amazonas: Chachapoyas, Cerros Calla Calla, E side, 1 9 km above Leimebamba on road to Balsas, Hutchison & Wright 5547 (F, GH, NY, uc). Chachapoyas, Cerros Calla Calla, W side, above Balsas on road to Leimebam- ba, Hutchison & Wright 5810 (F, GH, MO, NY, uc, us). San Martin: Mariscal Caceres, NW corner of Rio Abiseo Nat. Park, Chochas, Young & Leon 4735 (USM). Huanu- co: Cani, Pueblo 7 mi NE at Mito, Macbride 3403 (F). Carpish Pass, 84 km from Tingo Maria, Allard 20985 (us). Pasco: Oxapampa, Cumbre de la cordillera San Gutardo, Leon 521 (USM). Ayacucho: LaMar, eastern Massif of Cordillera Central, Dudley 12027 (GH, us). Cuzco: La Convencion, grasslands at camp 6, cordillera Vilcabamba, Dudley 11041 (GH). Cabecera del Karibeni, Biies 1 960 (us). 35. Elaphoglossum gracillimum Mickel, sp. nov. Rhizomate pergracili, lamina lineari tenui, frondisque stipite elongata et lamina parva recognita. 134 FIELDIANA: BOTANY Stem long-creeping, 1-1.5 mm in diameter, scales linear, lustrous, castaneous, to 4 mm long, entire to minutely and sparsely serrulate. Phyllo- podia lacking. Leaves ca. 1 cm apart, 8-1 2 cm long, 0.6-1.3 cm wide. Petiole '/8-% the sterile leaf length, with scales castaneous, widely spreading, 1-2 mm long, lanceolate, finely serrulate toward apex. Lamina linear-lanceolate, chartaceous, apex acuminate, base broadly cuneate to rounded. Veins evident, free, 1.5-2 mm apart, at 60-70-degree angle. Hydathodes distinct. Lamina scales subu- late, castaneous, scattered, 1 mm long on both surfaces, also punctate with stellate trichomidia. Fertile leaves slightly shorter than the sterile, pet- iole ca. 4/5 the fertile leaf length, lamina elliptic, apex rounded, base rounded to broadly cuneate, scales of petiole, adaxial surface and abaxial costa ovate, smaller (0.5 mm long), and darker (maroon) than on sterile. TYPE— Peru, Pasco, Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8441 (holotype, NY!; isotypes, MO!, uc!). Endemic. Epipetric on wet rocks in wet forest, 2300-2500 m, Pasco. Thus far known only from the type collection. 3 6 . Elaphoglossum guamanianum (Sodiro) C. Chr. , Index fil. suppl. 1: 42. 1913. Acrostichum guamanianum Sodiro, Sert. fl. ecuad. 2. 35. 1908. TYPE: Ecuador, "in pascuis andin. an- dium orient, in m. Guamani," Sodiro, Jan. 1900 (holotype, P!; isotype, B!). Stem compact, horizontal, 4-5 mm in diameter (ca. 1 cm with scales and phyllopodia), scales lin- ear-deltate, lustrous, dark red-brown, 1.5-2 mm long, with stiffhairlike teeth. Phyllopodia present. Leaves approximate, 17-35 cm long, 1.0-1.7 cm broad. Petiole VS-V4 the sterile leaf length, scales scattered, much reduced, to 1 mm long, with long hairlike teeth, dark red-brown, lustrous, ap- pressed. Lamina linear-elliptic, chartaceous, apex acuminate, base narrowly cuneate. Veins obscure, ca. 1 mm apart, at 70-degree angle. Hydathodes lacking. Lamina scales adaxially lacking, abaxially loosely clothed with scales 0.5-1 mm long, orange- tan, lanceolate, with long hairlike teeth, costal scales scattered, dark, as on petiole. Fertile leaves slightly shorter than the sterile, petiole ca. '/3 the leaf length, lamina only slightly narrower than the sterile, apex acute; intersporangial scales lacking. Epiphytic in wet forests, 1 700 m, Amazonas. Ecuador; Peru. Peruvian specimens match in that costa scales are small and black, the laminar scales are small (0.5-1 mm long), but lamina is wider in Ecuador- can material (3.5-4.1 cm vs. 1.4-1.7 cm). Ela- phoglossum guamanianum is similar to E. pale- aceum in the black, stiffly dentate stem scales and long-ciliate laminar scales, but it differs in the cos- tal scales being black, the laminar scales small, and the scales are much smaller overall. Amazonas: Prov. Bagua, 12 km E of La Peca, Barbour 2564 (F, MO, NY, uc). 37. Elaphoglossum hartwegii. (Fee) Moore, Index fil. xvi. 1857. Acrostichum hartwegii Fee, Mem. foug. 2: 53, /. 9, / 2. 1845. TYPE: "Quito, Popayan, Bogota," Hart- weg 1486 (FI, Herb. Webb, & G, Herb. Delessert). Stem long-creeping, 1-1.5 mm in diameter, scales linear-lanceolate, castaneous, lustrous with markedly paler margins, with sagittate base, en- tire, ca. 3 mm long. Phyllopodia distinct. Leaves often 1 cm apart, to 22 cm long, 1.2 cm broad. Petiole '/2-% the sterile leaf length, scales mostly 1-2 mm long, brown with pale margin, appressed to slightly spreading, entire to slightly dentate. Lamina linear-lanceolate, apex acuminate to acute or obtuse, base cuneate. Veins inconspicuous, free, 0.8-1 mm apart, at 60-70-degree angle. Hyda- thodes present but inconspicuous, usually hidden under scales. Lamina scales of adaxial surface gen- erally abundant but not overlapping, lanceolate to deltate, peltate, brown with pale margins, adaxial surface often essentially glabrous. Fertile leaves slightly taller than the sterile, petiole 2/3-% the fer- tile leaf length, lamina similar in size and shape to the sterile; intersporangial scales lacking. Terrestrial and epipetric in paramos, 4000-4700 m, Junin, Madre de Dios. Mexico; Guatemala; Costa Rica; Colombia to Bolivia. This is often mistaken for E. mathewsii, from which it can be distinguished by the broader, erose lamina scales without ciliate base. The two occa- sionally grow together but are distinct. It is even more closely allied to E. rosenstockii, which has similar scales, but has a wider creeping stem and scales on the abaxial lamina surface. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 135 Junin: Mount La Juntay, near Huancayo, Killip & Smith 22053 (GH [mixed with mathewsii], NY). Huaytapallana, Tiller 122 (USM). Madre de Dios: Pinasniocj, Pantiacolla Pass, Cook & Gilbert 1792 (us). 38. Elaphoglossum hayesii (Kuhn) Maxon, Proc. Biol. Soc. Wash. 46: 105. 1933. Acrostichum hayesii Kuhn, Linnaea 36: 43. 1869. TYPE: Panama, Sutton-Hayes 30, (holotype B!) Similar to E. piloselloides except: lamina thin, oblong, rounded at base, with hydathodes appar- ent; fertile lamina orbicular. Terrestrial or epipetric on dripping cliffs, 400- 1 500 m, San Martin, Junin. Panama; Venezuela; Colombia; Peru. San Martin: Boqueron Pass, 92 km from Tingo Maria on highway to Pucallpa, Allard2221, in part (us). Junin: Chanchamayo Valley, C. Schunke318 (F). 39. Elaphoglossum haynaldii (Sodiro) Losch, Mitt. Dot. Staatssamml. Munchen 1: 21. 1950. Acrostichum haynaldii Sodiro, Recens. crypt, vase. Quit. 80. 1883. TYPE: Ecuador, Los Colorados, Sodiro (not located). Stem short-creeping, ca. 5 mm in diameter, scales lanceolate, dark orange to maroon, ca. 5 mm long, entire or with irregular teeth. Phyllopodia lacking. Leaves fasciculate, to 40 cm long, 1 .3-2.4 cm broad. Petiole ca. V4 the sterile leaf length, with scales many, spreading, reddish, subulate, 4-6 mm long; petiole also with minute erect glandular trichomes ca. 0.2 mm long. Lamina linear-elliptic, charta- ceous, apex acuminate to acute-cuspidate, base cu- neate. Veins obscure, free, 1.5-2 mm apart, at ca. 60-degree angle. Hydathodes evident. Lamina scales subulate, reddish, conspicuous, 4-6 mm long, uniformly and liberally distributed over the surfaces, margin and costa, lamina also with mi- nute, erect, glandular trichomes. Fertile leaves un- known, but expected to be much shorter than the sterile, lamina orbicular to oblong; intersporangial scales lacking. Epipetric in wet forests, 1500-2450 m, Ayacu- cho, Cuzco. Ecuador; Peru. This differs from E. siliquoides (Jenm.) C. Chr. (of Jamaica) in being epipetric, and having res- inous petiole and stem, shorter lamina scales, con- spicuous hydathodes, and slender stem. Ayacucho: Ccarrapa, between Huanta & Rio Apuri- mac, Killip & Smith 22392 (F, NY, us). Cuzco: Paucar- tambo, Manu Nat. Park, Skog & Skog 5205 (us). Pau- cartambo, Pillawata, Paso del Aguila, Vargas 22993 (GH). 40. Elaphoglossum hickenii (Sodiro) C. Chr., In- dex fil. suppl. 1:42. 1913. Acrostichum hickenii Sodiro, Sert. fl. ecuad. 1: 35. 1908. TYPE: Ecuador, in silv. suband. vulc. Tun- gurahua, Sodiro, Dec. 1904 (isotypes, s!, us!). Stem compact, 4-6 mm in diameter, scales lin- ear-lanceolate, lustrous, castaneous to black-ma- roon, 3—4 mm long, entire. Phyllopodia present, obscured by scales. Leaves approximate, 48-64 cm long, 2.5-4. 1 cm broad. Petiole V2 the leaf length, scales moderately dense, appressed to ascending, lanceolate to linear-lanceolate, orange-tan, some with dark red tip or center, erose to very short- ciliate. Lamina narrowly elliptic, chartaceous, apex acuminate, base broadly cuneate. Veins ca. 1 mm apart, at 70-degree angle, hidden by scales. Hy- dathodes lacking. Lamina scales adaxially sparse to lacking, ovate-lanceolate to lanceolate, orange- tan, short-ciliate; abaxially ovate-lanceolate to lanceolate, orange, ciliate, 1-2 mm long; costal scales orange, some with maroon center, 1-2 mm long, the larger ones erose to short-ciliate. Fertile leaves nearly equalling the sterile in length, petiole % the leaf length, lamina slightly narrower than the sterile, apex obtuse, base subtruncate; costal scales linear-lanceolate, ciliate, red-centered, in- tersporangial scales lacking. Terrestrial on steep, rocky slopes, ca. 1 800-3000 m, Apurimac, Huanuco, Cuzco. Ecuador; Peru. Elaphoglossum hickenii is similar to E. cuspi- datum except: abaxial lamina scales ovate to lan- ceolate, loosely arranged, not appressed, the costal scales lanceolate, dark red-centered. It closely re- sembles E. orbignyanum but differs in the abaxial lamina scales being loosely arranged and the stem scales nonciliate. In general appearance it may also be taken for E. laminarioides, but the petiole and lamina scales are lanceolate rather than round. The epithet was originally published as "hike- 136 FIELDIANA: BOTANY nil" and so combined by Christensen, but Mille (Nov. Recens. 28. 1928) added the "c". The spell- ing is mixed on type specimens; e.g., an isotype (s) is labelled "hikenii" but with a "c" inserted in the same hand. There is no reference to the name in the protologue, but it is likely named for the Argentine botanist Cristobal M. Hicken. \ pun mac: Abancay, Bosque Ampay, Hocking 36 (USM). Huanuco: Mito, Macbride & Featherstone 1392 (us). Near Huanuco, Calderon, 8 June 1967 (USM). Cuzco: Paucar- tambo, Chacapampa, Vargas 9880 (uc). Machu Picchu, Ferreyra 27070 (GH), Sounders 462A (K), Tryon & Tryon 5397 (GH), D'Arcy 13771 (MO). 4 1 . Elaphoglossum hieracioides Mickel, Brittonia 39: 320. 1987. TYPE: Venezuela, Aragua, To- var, Fendler 268 (holotype, us!; isotypes, GH!, MO!). Similar to E. piloselloides except: fertile lamina narrowly oblanceolate, much longer than broad, often inrolled, base cuneate. Terrestrial on clay banks and in open woods, 750-2300 m, Amazonas, San Martin, Huanuco, Pasco, Junin, Ayacucho. Venezuela; Colombia; Peru. Amazonas: Chachapoyas, slopes of Cano Santa Lucia E of Chachapoyas, Wurdack 743 (F, GH, NY, us). San Martin: Boqueron Pass, 92 km from Tingo Maria on highway to Pucallpa, Allard 2221, in part (us). Huanuco: Muna, Bryan 469 (F, GH). Pasco: Oxapampa, around Villa Rica, van der Werff et al. 8277 (MO, uc). Junin: Chanchamayo Valley, C. Schunke 474 (F). Ayacucho: Ayna, between Huanta & Rio Apurimac, Killip & Smith 23130 (F, GH, NY, us). 42. Elaphoglossum horrid ulum (Kaulf.) John Sm . . Hot. Voy. Herald 232. 1854. Acrostichum horridulum Kaulf, Enum. fil. 58. 1824. TYPE: Brazil, Otto (holotype, not located). Stem compact, horizontal to ascending, ca. 2 mm in diameter, scales linear-lanceolate, entire, orange, lustrous, 3-5 mm long. Phyllopodia lack- ing. Leaves fasciculate, 7-13 cm long, 0.5-0.9 cm broad. Petiole V3-V2 the sterile leaf length, with scales subulate, orange to maroon, ca. 3 mm long, widely spreading, and with scattered minute erect glandular trichomes. Lamina narrowly to linear elliptic, chartaceous, apex acute, base narrowly cu- neate. Veins obscure, free, 1.5-2 mm apart, at ca. 30-degree angle. Hydathodes evident. Lamina scales subulate, tan to dull orange, evenly distrib- uted on both surfaces, and with minute, incon- spicuous, gland-tipped trichomes. Fertile leaves 2/3 as long to slightly surpassing the sterile in length, petiole 3/4-7/8 the fertile leaf length, lamina elliptic, apex obtuse, base broadly cuneate; intersporangial scales lacking. Epipetric in wet forests, ca. 450 m, San Martin, Junin. Venezuela; Peru; Bolivia; Brazil. The Brazilian material is often somewhat small- er but there seems to be no substantial difference. San Martin: Lamas, along Rio Curiyacu, 8 km above San Antonio, Belshaw 3609 (F, GH, NY, uc, us). Near Tarapoto, Spruce 4040 (GH, NY, us). Junin: Chancha- mayo Valley, C. Schunke 474 (us). 43. Elaphoglossum huacsaro (Ruiz) Christ, Monogr. Elaphoglossum 96. 1899. Acrostichum huacsaro Ruiz, Mem. sobre la legitima Calaguala, 57. 1805. TYPE: Peru, Ruiz 54 (B!, BR!; photos, F, us). Acrostichum calaguala Klotzsch, Linnaea 20: 421. 1847. SYNTYPES; Peru, in Andium nemoribus, Ruiz 54 (B!, BR!); Colombia, Paramo de Mucute, Moritz 315 (not located). Elaphoglossum calaguala (Klotzsch) Moore, Index fil. 7. 1857. Stem short creeping to ascending, 3-5 (9) mm in diameter, scales linear-lanceolate, black, lus- trous, entire, 1-2 mm long. Phyllopodia indistinct. Leaves fasciculate, 14-31 cm long, 0.8-1.9 cm broad. Petiole '/4-!/3 the sterile leaf length, with scales ovate, dark to pale, often pale with dark sclerotic tip, 1 mm long, entire, mostly appressed, a few spreading. Lamina narrowly elliptic, char- taceous, apex acute to obtuse, base narrowly cu- neate. Veins obscure, free, ca. 1 mm apart, at ca. 60-degree angle. Hydathodes lacking. Lamina scales reduced to resinous dots on both surfaces, scales along lamina margin ovate and entire, ca. 1 mm long, some scattered on costa abaxially. Fertile leaves far exceeding the sterile in length, but lamina narrower, petiole nearly as long as the entire sterile leaf; intersporangial scales lacking. Epiphytic in wet forests, 2050-3200 m, Caja- marca and Amazonas south to Huancavelica and Cuzco. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 137 Mexico to Panama; West Indies; Colombia to Bolivia. Huanuco: Cuchero, Poeppig, July 1829 (photo of pho- to at BM). Junin: Villa Amoretti, Kunkel 535 (GH). Cajamarca: San Miguel, Niepos, Quiroz 1539 (F). A ma/onas: Chachapoyas, Cerros Calla Calla, 5 km above Leimebamba, Hutchison & Wright 4818 (F, NY, uc, us). Chachapoyas, cerros Calla Calla, west side, 45 km above Balsas, Hutchison & Wright 5799 (uc). Cordillera de Calla Calla, Balsas-Leimebamba road, Duncan 2607 (uc). San Martin: Huallaga, Valley of Rio Apisoncho, Ham- ilton & Holligan 901 (K), 90S(uc). Mariscal Caceres, near La Playa camp, Rio Abiseo Nat. Park, Young & Leon 4950 (USM). Huanuco: W of Carpish, Stork & Horton 9892 (uc). Huanuco, Carpish Km 453, Plowman 6066 (GH). Huanuco-Tingo Maria road, Luteyn 5453 (uc). Pasco: Oxapampa, Canyon de Huancabamba, Leon 618 (F). Oxapampa, border of Yanachaga Nat. Park, Leon et al. 955 (F). Junin: Carpapata above Huacapistana, Killip & Smith 2441 1 (NY), 24414 (uc, us). Huanca velica: Prov. Tayacaja, Quebrada, Stork & Horton 10312 (us). Aya- cucho: Condorcunca, Barron 16 May 1964 (USM). Apu- rimac: E of Abancay, road to Cuzco, Hutchison 1733 (uc). Cuzco: Paucartambo, between Pillawata and Patria (Kosnipata Valley), Plowman & Davis 4939 (GH). Dist. Vilcabamba, Valle San Miguel Convencion, Yungacal- marica, Bues 2056 (us). 44. Elaphoglossum hystrix (Kunze) Moore, Index fil. 10. 1857. Acrostichum hystrix Kunze, Linnaea 9: 26. 1834. TYPE: Peru, Huanuco, Pampayaco, Poeppig (ho- lotype, LZ, destroyed; isotype, B?). Stem lacking, probably compact, scales linear- lanceolate, lustrous, castaneous, 3-4 mm long, en- tire. Ph yllopodia present. Leaves approximate, 30- 33 cm long, 1 .4-1 .8 cm broad. Petiole about '/5 the sterile leaf length, scales scattered, dark red-brown, subulate, patent, ca. 3 mm long, entire, and also with scattered, 2-3 -celled, erect, glandular tri- chomes ca. 0.1 mm long. Lamina linear, charta- ceous, apex acuminate, base cuneate. Veins evi- dent, 1-1.7 mm apart, at 70-80-degree angle. H ydathodes lacking. Lamina scales adaxially scat- tered, red-brown, subulate, 1-1.5 mm long on sur- face, 1.5-2 mm long on margin, 1.5-2 mm long on costa, most abundant on margin and costa; abaxially abundant on costa, sparse on surface, and also with scattered, linear trichomidia ca. 0.2 mm long. Fertile leaves unknown. Endemic. Habitat unknown but presumably epiphytic in wet forests. Huanuco, Junin. Similar to E. erinaceum except: lamina linear elliptic, 1 .4-1 .8 cm broad, long-acuminate at apex, narrow at base. 45. Elaphoglossum jucundum Mickel, sp. nov. Rhizomate repenti squamis patulis fuscis obsito et lamina parva apice cuspidata abaxialiter glandulosa squamisque dentatis instructa notabile. Stem wide-creeping, ca. 1 mm in diameter, scales linear, lustrous, dark reddish brown, 3-4 mm long, spreading and recurved, sparsely denticulate and with a few short cilia at scale base. Phyllopodia present. Leaves 3-5 mm apart, 1.5-3.5 cm long, 0.6-1.1 cm broad. Petiole ca. V3 the sterile leaf length, with scales linear-lanceolate, dark brown to orange, lustrous, 1.5-2 mm long, denticulate, ciliate at scale base, spreading, and with resinous dots. Lamina ovate-lanceolate, chartaceous, apex acuminate, base rounded. Veins obscure, ca. 1 mm apart, at ca. 60-degree angle. Hydathodes lacking. Lamina scales curved, ciliate, 1-1.5 mm long, mostly orange, some dark red-brown along abaxial costa, more abundant abaxially, with resinous dots on both surfaces. Fertile leaves unknown. TYPE— Peru, Choquezuffa, al pie del Nevado, Bues 1379 (holotype, us!). Endemic. Epiphytic in wet forests, ca. 900 m, Cuzco. Thus far known only from the type. 46. Elaphoglossum killipii Mickel, sp. nov. Ab E. tenuiculo fronde fertile steriles longe superanti, habitu epiphytico, regionis inferiosis incola diversum. Stem short-creeping, 1.5-2 mm in diameter, scales linear, with twisted hairlike tip, lustrous, castaneous, 1-1.5 mm long, entire. Phyllopodia present. Leaves approximate, 75-10 cm long, 2.5- 3 cm broad. Petiole about Vs the sterile leaf length, with dense scales appearing as stellate trichomes with resinous base. Lamina linear, chartaceous, apex acuminate, base attenuate. Veins not visible. Hydathodes lacking. Lamina scales appearing as stellate or substellate trichomes, adaxially white to orange, abaxially more abundant, orange, with resinous base. Fertile leaves longer than the sterile (to twice as long), petiole ca. 2/5 the leaf length, lamina broader than the sterile (5-6 mm broad); intersporangial scales lacking. 138 FIELDIANA: BOTANY TYPE— Peru, Junin, San Ramon, Killip & Smith 24772 (holotype, us!). Endemic. Epiphytic in wet forests, 900-1 300 m, Junin. This is close to E. tenuiculum but distinct in the substellate trichomes rather than being strictly stellate, the fertile leaf much longer than the sterile, plants epiphytic rather than terrestrial, and oc- curring at lower elevation (900-1300 vs. 1500- 3000 m). 47. Elaphoglossum laminarioides (Fee) Moore, Index fil. 10. 1857. Acrostichum laminarioides Fee, Mem. foug. 2: 57. /. 12. 1 845. TYPE: French Guiana, Leprieur in 1834 (holotype, P!). Stem short-creeping, ca. 3-5 mm in diameter, scales linear-lanceolate, lustrous, dark brown to black, 3-5 mm long, with short teeth. Phyllopodia distinct. Leaves fasciculate, 32-104 cm long, 4.1- 5.4 cm broad. Petiole about 2/5 the sterile leaf length, with pale, appressed, round, peltate, ciliate scales ca. 1 mm long. Lamina narrowly elliptic, charta- ceous, apex acuminate to cuspidate, base attenu- ate. Veins obscure, free, 1 mm apart, at ca. 80- degree angle. Hydathodes lacking. Lamina scales on both surfaces round, peltate, ciliate, white, 1 mm in diameter. Fertile leaves shorter than the sterile, petiole '/3 the fertile leaf length, the lamina linear, 9-12 mm wide; intersporangial scales lack- ing. Epiphyte in wet forests, 600-1300 m, San Mar- tin, Junin, Ayacucho, Cuzco. Venezuela; Ecuador; Peru. This plant has white, round, peltate, ciliate scales on the petiole and the lamina surfaces. San Martin: Mariscal Caceres, Palo Blanco, al oeste del Puente, Tocache Nuevo, J. Schunke V. 5673 (NY). Tingo Maria, Allard21586 (GH, us). Junin: E of Quimiri Bridge, near La Merced, Killip & Smith 23960 (NY, us). Prov. Jauja, Valle de Sanibeni, Ridoutt 12262 (us). Aya- cucho: Carrapa, between Huanta and Rio Apurimac, Kil- lip & Smith 22424 (NY). Cuzco: Machu Picchu, Coronado 109 (uc). ab E. nervoso laminae squamis erectis longe ciliatis dis- tantior. Stem compact, horizontal, 4-7 mm in diameter, scales linear-lanceolate, lustrous, black, 3-5 mm long, dentate, at least toward base. Phyllopodia present but hidden by scales. Leaves fasciculate, 15-38 cm long, 0.8-1.5 cm broad. Petiole '/3-'/2 the leaf length, scales orange, appressed, round, erose to short-ciliate, 1 mm in diameter, also with dark, ascending, lanceolate, erose scales 2-3 mm long. Lamina narrowly elliptic, chartaceous, apex obtuse, base narrowly to broadly cuneate. Veins obscure, free, 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales adaxially round to lanceolate, white-membranous (silvery), 1-2 mm long, erose, abaxially the dense, erect scales more or less round with long cilia, longer than scale body, costal scales broad-ovate, white, dark-cen- tered, ovate, 1-2 mm long, adaxially round to lan- ceolate, white-membranous (silvery), 1-2 mm long, erose. Fertile leaves longer than the sterile, petiole 2/3-3/4 the fertile leaf length, lamina about equal the sterile in size and shape; intersporangial scales lacking, costa with ovate, white (dark-centered) ciliate scales 1-2 mm long. TYPE— Peru, Apurimac, Abancay, E of Aban- cay, road to Cuzco, Hutchison 1732 (holotype, NY!; isotypes, F!, GH!, uc!, us!). Endemic. Terrestrial on brushy roadcuts and steep slopes, 2050-3600 m, Amazonas, Lima, Ju- nin, Ayacucho, Apurimac, Cuzco. This is variable in the degree of ciliation of the laminar scales; i.e., the scale body ranges from substantial to only very slight. Amazonas: Chachapoyas, Cerros Calla Calla above Leimebamba, Hutchison & Wright 48 ISA (NY, uc). Alrededores de Leimebamba, Lopez el al. 4377 (GH). Lima: Canta, Huamantanga, Sounders 1253 (GH). Junin: Tarma, between Palca and Carpapata, Stork 10959 (uc). Huancayo, arriba de Huancayok, Tovar 2792 (GH). Aya- cucho: Ccarrapa, between Huanta & Rio Apurimac, Kil- lip & Smith 22282 (NY). Apurimac: Abancay, laderas altos de Abancay, Vargas C. 16596 (GH). Cuzco: Paruro, Vargas C. 7914 (GH). Department unknown: Ruiz & Pa- von (us ex Herb. Kew). 48. Elaphoglossum lanatum Mickel, sp. nov. Laminae squamis appressis et costae obscuris ovato- lanceolatis erosis inferne dissectis ab E. deorso diversa, 49. Elaphoglossum lasioglottis Mickel, sp. nov. Ab E. nigrocostato lamina apice obtusa costaeque squamis latioribus dilutius nigris dissimile. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 139 Stem compact, horizontal, 2-5 mm in diameter, scales linear, lustrous, castaneous to dark red- brown, 3-5 mm long, with sparse, sometimes de- ciduous hair-teeth. Phyllopodia evident. Leaves fasciculate, 8-25 cm long, 0.6-1.2 cm broad. Pet- iole V3-V2 the sterile leaf length, with dark red- brown to black scales appressed to spreading, 1— 2 mm long, dentate, appearing resinous. Lamina narrowly elliptic, subcoriaceous, apex obtuse to acute, base cuneate. Veins obscure, free, 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales abaxially ovate-lanceolate, orange, ciliate, costa larger, lanceolate, red-brown, short- ciliate, adaxially pale orange, dense, round to lan- ceolate, 1-2 mm long, cilio-denticulate. Fertile leaves slightly shorter or longer than the sterile, petiole ca. % the fertile leaf length, lamina about same size and shape as the sterile lamina; inter- sporangial scales lacking. TYPE— Peru, La Libertad, Santiago de Chuco, Chota-Shorey, Sagdstegui 11102 (holotype, NY!; isotype, GH!). Epipetric in damp banks, rock crevices, and along streams, 2800-3250(-4200) m, Piura, Cajamarca, La Libertad, Ancash, Huancavelica. Colombia to Bolivia. This is closely related to E. nigrocostatum Mick- el, of Venezuela. Piura: Huancabamba, above Canchaque on road to Huancabamba, Hutchison 1644 (F, NY, uc). Cajamarca: San Miguel, Cerro Quillon (Agua Blanca), Mostacero L. et al. 1299 (F). La Libertad: Otuzco, Otuzco, above Agal- pampa on road Trujillo-Quiruvilca, Saunders 893 (F, GH). 3 km west of Huamachuco, Correll&E. Smith P933 (GH). Ancash: Yungay, Huascaran Nat. Park, Quebrada Parla, D. Smith et al. 9173 (NY). Huancavelica: Tayacaja, 1 km before Huari, Saunders 1 142 (F, GH). 50. Elaphoglossum latevagans Mickel, sp. nov. Rhizomate longe repenti squamis linearibus nigris in- duto necnon lamina lanceolata hydathodiis squamisque linearibus badiis provisa distinguibilis. Stem long-creeping, 2-3 mm in diameter, often with some white patches, scales linear-lanceolate, lustrous, maroon, 1.5-2 mm long, entire. Phyl- lopodia lacking. Leaves distant, mostly 5-20 mm apart, 30-47 cm long, 1.8-3.0(4.8) cm broad. Pet- iole about '/-> the sterile leaf length, scales scattered, linear, dark brown to tan, appressed to ascending, 2-3 mm long, more and darker distally, and ex- tending onto lamina, those of costa with dark base or center, those on lamina pale, on both surfaces, 2-5 mm long. Lamina lanceolate to narrowly el- liptic, chartaceous, apex acuminate to caudate, base narrowly to broadly cuneate. Veins evident, free, ca. 1 mm apart, at ca. 60-degree angle. Hydathodes distinct. Lamina scales abaxially scattered, linear, tan, entire, 3 mm long, those on costa with reddish brown sclerotic streaks, adaxially similar but 2 mm long and lacking streaks on costal scales. Fer- tile leaves approximately equalling sterile ones in length, petiole about 2/3 the fertile leaf length, lam- ina lanceolate, dark scales on costa and pale ones among sporangia. TYPE— Peru, Amazonas, Chachapoyas, Cerros Calla Calla, Hutchison & Wright 5808 (holotype us!; isotypes, F!, GH!, NY!, uc!). Endemic. No habitat data; 3100 m, Amazonas. The paratype has a broader lamina (to 4.8 cm) than does the type material. Amazonas: Chachapoyas, Cerros Calla Calla, Hutch- ison & Wright 5808A (GH). 51. Elaphoglossum latifolium (Sw.) John Sm., London J. Bot. 1: 197. 1842. Acrostichum latifolium Sw., Prodr. 128. 1788. TYPE: Jamaica, Swartz (holotype, s). Stem short to moderately creeping, 4-10 mm in diameter, scales linear-lanceolate, orange to dull tan or dirty brown to slightly blackish, occasion- ally somewhat lustrous, to 8 mm long, entire or with occasional hairlike processes. Phyllopodia distinct. Leaves slightly to distinctly spaced, to 1 28 cm long, 4.9-9.8 cm broad. Petiole usually V4-V3 the sterile leaf length, glabrous. Lamina linear-lan- ceolate, coriaceous, apex acuminate, base cuneate to broadly cuneate. Veins obscure, free, about 1 mm apart, at 70-80-degree angle. Hydathodes lacking. Lamina surfaces usually naked, but some- times minute stellate black trichomidia on the ab- axial surface. Fertile leaves about equal to the ster- ile ones in dimensions but sometimes slightly narrower or with longer petiole; intersporangial scales lacking. Epiphytic in wet forests, 550-3800 m, Caja- marca, Amazonas, La Libertad, San Martin, Lo- reto, Huanuco, Pasco, Junin, Cuzco. Mexico to Panama; West Indies; Venezuela and Colombia to Bolivia and Brazil. 140 FIELDIANA: BOTANY This is placed in E. latifolium in the broad sense. In the narrow sense it is probably limited to the West Indies, but the complex is taxonomically very difficult. The Peruvian material has rather large leaves, with moderately creeping stems. Cajamarca: Contumaza, bosque de Cachil (Cascas- Contumaza), Lopez M. et al. 9112 (F, GH). Amazonas: Chachapoyas, Cerros Calla Calla, 45 km above Balsas, midway on road to Leimebamba, Hutchison & Wright 5796 (F, GH, NY, uc, us). La Libertad: Otuzco, Hua- ranchal, Sagastegui A. 0199(GH). San Martin: Mariscal Caceres, Tocache Nuevo, Cerro Sinsin, 15 km W of Tocache, Plowman & Schunke V. 11463 (F). Loreto: Mi- shuyacu near Iquitos, Klug 1461 (F). Huanuco: Muna, Bryan 541 (F). Prov. Huanuco, Mirador, cerca a Hua- nuco, Ridoutt, Feb. 1940 (GH). Pasco: Oxapampa, Gran Pajonal, 2-3 km N of Chequitavo, D. Smith 5087 (NY). Junin: E of Quimiri Bridge near La Merced, Killip & Smith 23850 (NY). Cuzco: Pillahuata, Cerro de Cusilluy- oc, Pennell 14027 (GH, NY). 52. Elaphoglossum lawyerae Mickel, sp. nov. Rhizomate compacto, frondibus subsessilibus lineari- ellipiticis glabris distinguendum. Named for Jean Lawyer, who was a volunteer in my office for fifteen years, helping in matters of curation, research, and the American Fern Society. Stem compact, ca. 5 mm in diameter, scales lanceolate, black, dull, thin, 3-4 mm long, with occasional irregular teeth. Phyllopodia present. Leaves approximate, 34-42 cm long, 1.8-2.4 cm broad. Petiole V15-VW (!4) the sterile leaf length, glabrous. Lamina linear-elliptic, firmly charta- ceous, apex acuminate, base attenuate. Veins ev- ident, 1-1.5 mm apart, at ca. 70-degree angle. Hy- dathodes lacking. Lamina scales lacking, but sparse, stellate trichomidia on both surfaces. Fertile leaves nearly equalling the sterile in length, petiole ca. '/3 the leaf length, lamina broader than the sterile (2.8 cm); intersporangial scales lacking. TYPE— Peru, Pasco (as Junin), Pichis Trail, be- tween San Nicolas and Azupizu, Killip & Smith 26122 (holotype, NY!). Endemic. Epiphytic in wet forests, 650-900 m, Pasco. Thus far known only from the type. 53. Elaphoglossum laxisquama Mickel, sp. nov. Lamina magna laxe squamata necnon stipite hirta in- signis. Stem moderately to short-creeping, to 20 cm long, 2-3 mm in diameter, scales linear-lanceolate, lustrous, red-black, 5-7 mm long, with hair-tip and divergent hair-teeth. Phyllopodia evident. Leaves approximate, to 5 mm apart, 25-53 cm long, 2.0-4.1 cm broad. Petiole V3-V2 the sterile leaf length, with tan to black-streaked scales widely spreading, 3-5 mm long, with hairlike teeth. Lam- ina narrowly elliptic, coriaceous, apex obtuse, base broadly cuneate to rounded or truncate. Veins ob- scure, free, 1 mm apart, at 80-degree angle. Hy- dathodes lacking. Lamina scales abaxially ovate to lanceolate, 1-1.5 mm long with long, hairlike teeth, scattered, concentrated at margin, orange to black, costal scales 2-3 mm long, dense, adaxial surface glabrescent or with scattered scales, these 1-2 mm long, orange. Fertile leaves longer than the sterile, petiole % the leaf length, lamina nar- rower; intersporangial scales lacking. TYPE— Peru, San Martin, Mariscal Caceres, Puerta del Monte, Rio Abiseo Nat. Park, Young 1741 (holotype, NY!; isotype, USM!). Endemic. Terrestrial and epiphytic in wet for- ests, 3300-3500 m, La Libertad, San Martin, Pas- co, Cuzco. La Libertad: Pataz, Chirimachay, Pataz, Young 2960 (NY, USM). San Martin: Mariscal Caceres, forest patch above timberline, Puerta del Monte, Rio Abiseo Nat. Park, Young 1847, 1951 (NY). Mariscal Caceres, Cho- chos, Rio Abiseo Nat. Park, Young 2536 (NY). Mariscal Caceres, Puerto del Monte, Rio Abiseo Nat. Park, Young 1926, 1995 (USM). Pasco: Oxapampa, Huancabamba, Sta. Barbara, above Lanturachi, Foster 10429 (F, GH). Cuzco: La Convention, Cordillera Vilcabamba, 28 km walking distance NE from Hacienda Luisiana & Apu- rimac River, Dudley 11188 (GH). 54. Elaphoglossum lechlerianum (Mett.) Moore, Index fil. 11. 1857. Acrostichum lechlerianum Mett., Fil. lechl. 1:3.1 856. TYPE: Peru, Puno, Tatanara, Lechler 2522 (iso- types, BR!, s!; photo, us of BR). Stem creeping, up tree, vertical, ca. 4-6 mm in diameter, scales linear-lanceolate, lustrous, or- ange, 4-6 mm long, with a few low teeth distally and an occasional hairlike process. Phyllopodia present. Leaves fasciculate, 26-52 cm long, 2.2- 4.2 cm broad. Petiole '/8-'/4 the sterile leaf length, with orange scales scattered, appressed, ovate to lanceolate, 1-2 mm long, entire. Lamina narrowly elliptic, chartaceous, apex acuminate, base atten- TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 141 uate. Veins evident, free, 1-1.5 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales abaxially reduced to minute, stellate tricho- midia, a few appressed scales on the costa base, adaxially smaller and sparser. Fertile leaves about equal the sterile, petiole V3-V2 the fertile leaf length, the lamina narrow linear-elliptic to linear-oblan- ceolate, 1.0-2.8 cm broad; intersporangial scales lacking. Terrestrial and epiphytic in wet forests, 1 760- 2770 m, Amazonas, Huanuco, Pasco, Cuzco, Puno. Ecuador; Peru. Amazonas: Bagua, 25 km E of La Peca, Barbour 2954 (MO). Bagua, 17 km E of La Peca, Gentry et al. 22977 pp (MO). Huanuco: SW slope of Rio Llullapichis water- shed on ascent of Cerros del Sira, Dudley 13328 (GH). Prov. Huanuco, Carpish, C. Vargas C. 5424 (uc). Pasco: Chontabamba, Abra "La Suiza," camino al Rio Chon- tabamba, Leon et al. 978 (F). Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8427 (MO, uc). Cuzco: La Convencion, Huayopata, 6 km from In- catambo on Lucumayo River, Peyton & Peyton 1557 (MO). La Convencion, Cordillera Vilcabamba, camp 1- '/2, Dudley 10342 (GH). La Convencion, 15 km walking NE from Hacienda Luisiana & Apurimac River, Cor- dillera Vilcabamba, Dudley 1 1248 (GH). La Convencion, Huayopata, Puncuyoj, 10 km SW of Incatambo, Peyton & King 1413 (GH). Machu Picchu, Cook & Gilbert 863 (us). Los Palmitos, Cabecera del Rio Karibeni, Biies 1962 (us). 55. Elaphoglossum leprosum (Kuhn) Christ, Monogr. Elaphoglossum 119. 1899. Acrostichum leprosum Kuhn, Linnaea 36: 48. 1869. TYPE: Peru (Junin), Tarma, Phillipi (holotype, B!). Stem creeping, ca. 2 mm in diameter, scales linear, lustrous, dark brown, 4-6 mm long, with short, patent teeth. Phyllopodia evident but ob- scured by scales. Leaves 2-15 mm apart, 16-20 cm long, 1.8-2.1 cm wide. Petiole '/3-2/5 the sterile leaf length, with tan, black-centered scales widely speading, 3 mm long, with short patent teeth, also with appressed, peltate, ciliate scales to 1 mm long. Lamina lanceolate, chartaceous, apex long-acu- minate, base broadly cuneate. Veins obscure, free, ca. 1 mm apart, at 70-80-degree angle. Hyda- thodes lacking. Lamina scales abaxially dense, most of them lanceolate, orange, ciliate, 2 mm long, costal scales (some) sclerotic in part, plus a few lanceolate scales with sclerotic patches, adaxially white, lanceolate, ciliate. Fertile leaves longer than the sterile, petiole !/2 the fertile leaf length, lamina similar to the sterile except slightly narrower; in- tersporangial scales lacking. Endemic. Epiphytic in wet forests, 3950 m, Huanuco, Junin. Huanuco: Tarn bo de Vaca, Macbride 4413 (F). 56. Elaphoglossum lindenii (Fee) Moore, Index fil. xvi. 1862. Acrostichum lindenii Fee, Mem, foug. 2: 48, t. 18, f. 3. 1845. TYPE: uncertain, four syntypes origi- nally cited. Stem short-creeping to ascending, ca. 3 mm in diameter, scales linear, maroon, to 8 mm long, with short teeth. Phyllopodia lacking. Leaves fas- ciculate, 7-13 cm long, 1.4-2.4 cm broad. Petiole !/2-3/4 the sterile leaf length, the scales orange, su- bulate, widely spreading, 2-3 mm long and also with a few minute erect glandular trichomes. Lam- ina ovate to widely elliptic, chartaceous, apex cus- pidate, base rounded or subcordate. Veins evident, free, ca. 2 mm apart, at 60-70-degree angle. Hy- dathodes distinct. Lamina scales subulate, orange to maroon, scattered on both surfaces, 2-3 mm long. Fertile leaves slightly shorter than the sterile, petiole to 4/5 the fertile leaf length, the lamina apex obtuse, base truncate; intersporangial scales lack- ing. Epipetric or terrestrial in wet forests, 3700-3840 m, Cajamarca, Ancash, Cuzco. Mexico to Panama; Venezuela and Colombia to Chile. Cajamacra: Prov. San Miguel, El Tingo, Sagdstegui 8824 (uc). Ancash: Yungay, Huascaran Nat. Park, D. Smith et al. 10470 A (NY) Prov. Yungay, Yungay, Llan- ganuco, Sounders 521 (F, uc). Cuzco: Paucartambo, Huaisampillo, Vargas C. 9963 (uc). 57. Elaphoglossum lingua (Raddi) Brack., U.S. Expl. Exped. 16: 74. 1854. Acrostichum lingua Raddi, Opusc. Sci. 3: 283. 1819. TYPE: Brazil, Raddi (FI or PI). Stem long-creeping, ca. 2 mm in diameter, scales sparse, ovate to lanceolate, black, sclerotic, lus- trous, 1-2 mm long, with cilia, especially at base. 142 FIELDIANA: BOTANY Phyllopodia present but not always dark. Leaves distant, mostly 1-3 cm apart, 9-30 cm long, 3.1- 5.2 cm broad. Petiole %-% the sterile leaf length, glabrous or with sparse black or black-tipped scales near petiole base. Lamina ovate to lanceolate or oblong, coriaceous, apex obtuse to acute, base nar- rowly to broadly cuneate. Veins obscure, free, 1 mm apart, at ca. 70-80-degree angle. Hydathodes lacking. Lamina scales lacking, or reduced abax- ially to stellate trichomidia. Fertile leaves about equal the sterile in length, petiole '/2-2/3 the fertile leaf length, the lamina narrowly oblong; inter- sporangial scales lacking. Epiphytic in wet forests, 550-1900 m, San Mar- tin, Huanuco, Pasco, Junin, Cuzco. Costa Rica; Panama; West Indies; Venezuela; Colombia to Bolivia and Brazil. San Martin: Mariscal Caceres, Tocache Nuevo, Cerro Sinsin, 1 5 km W of Tocache Nuevo along road to Puerto Pizana, Plowman & J. Schunke V. 11460 (F, NY). Hua- nuco: Dist. Churubamba, Puente Durand to Exito, Mex- ia 82 3a (uc). SW slope of Rio Llullapichis watershed on ascent of Cerros del Sira between Laguna & Pelogroso, Dudley 13205 (GH). Pachitea, slopes in logging area across Rio Huancabamba from town Pozuzo near bridge of Oxapampa-Pozuzo road, Skog et al. 5095 (NY). Cani, Macbride 3403, in part (us). Pasco: Oxapampa, Oxa- pampa to villa Rica, Skog et al. 5046b (us). Oxapampa, Quebrada Castilla on the Omaiz river, Leon & Young 1069 (F). Junin: Satipo, Pichanaki, rodal del proyecto Peruano-Aleman, Leon 231 (USM). Chanchamayo Val- ley, C. Schunke 473, 1441, 1465 (F). E of Quimiri Bridge near La Merced, Killip & Smith 23992 (NY, us). Pichis Trail, Porvenir, Killip & Smith 25916 (NY, us), 25698 (us). Cuzco: Paucartambo, entre Montanesa & Patria, C Vargas C. 16890 (GH). 58. Elaphoglossum litanum (Sodiro) C. Chr., In- dex fil. 310. 1905. Acrostichum litanum Sodiro, Sert. fl. ecuad. 10. 1905. TYPE: Ecuador, "ad fl. Lita in prov. Esmeral- das," Sodiro, Aug 1904 (holotype, P!; isotypes B!, pi). Stem compact, 5-8 mm in diameter, scales lin- ear-lanceolate, lustrous, dark red-brown, ca. 5 mm long, entire. Phyllopodia not evident, perhaps very short and obscured by crowded, stout petiole bas- es. Leaves approximate, 48-60 cm long, 2.5—4.0 cm broad. Petiole about '/10 the sterile leaf length, scales ovate-lanceolate, pale, dull, appressed, 2-3 mm long, erose. Lamina linear-oblanceolate, char- taceous, apex acute-acuminate, base attenuate. Veins evident, 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales adaxially de- ciduous except for fringe of 1 mm, pale, cilio- denticulate scales at margin; abaxially glabrous ex- cept for much-reduced fimbriate scales on costa. Fertile leaves slightly shorter than the sterile, pet- iole ca. 2/5 the leaf length, lamina obtuse, slightly narrower than the sterile, adaxial surface densely clothed with pale, round to lanceolate, cilio-den- ticulate, 1 mm long scales; intersporangial scales lacking. Endemic. Epiphytic in wet forests, 1800-1900 m, San Martin. This closely resembles E. subciliatum Rosenst. of Costa Rica, but that has a linear, fertile lamina. San Martin: Rioja, Pedro Ruiz-Moyobamba road, D. Smith 4503 (uc). 59. Elaphoglossum longius Mickel, sp. nov. Ab E. tenui stipite laminaque longioribus, laminaque ipsa squamis basi ciliolatis dorsaliter obsita semovenda. Stem long-creeping, 1 mm in diameter, scales linear, lustrous, dark red-brown, 2-3 mm long, minutely and sparsely dentate. Phyllopodia pres- ent. Leaves 1-1.5 cm apart, 18-34 cm long, 1.6- 2.5 cm broad. Petiole ca. 3/5 the sterile leaf length, scales scattered, linear, orange-tan, 2-3 mm long, minutely denticulate, and smaller ones appressed, more dissected. Lamina narrowly elliptic, char- taceous, apex long-attenuate, base cuneate. Veins evident, 1.5 mm apart, at ca. 70-degree angle. Hy- dathodes lacking. Lamina scales scattered on both surfaces, linear-lanceolate-deltate adaxially, abax- ially lanceolate, ciliate-denticulate with glandular dots. Fertile leaves nearly equalling the sterile in length, petiole 3/5 the leaf length, lamina nearly equalling the sterile in size and shape; interspor- angial scales lacking. TYPE— Peru, Cuzco, Abra de Malaga, 15 km toward Quillabamba, Ellenberg 4863 (holotype, GH!). Endemic. Epiphytic in mossy elfin forests, 3740 m, Cuzco. Similar to E. tenue except petiole 1 0-1 7 cm long, scales scattered, leaves longer, 22-34 cm, 1.7-2.5 cm broad, lamina scales abaxially deltate-lanceo- late, ciliate-based, gland-based, adaxially linear- TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 143 lanceolate, scattered; fertile lamina same shape and nearly same size as sterile. 60. Elaphoglossum luridum (Fee) Christ, Monogr. Elaphoglossum 33. 1899. Acrostichum luridum Fee, Mem. foug. 2: 35. t. 19, f. 1. 1845. SYNTYPES: "Guiana" (Guyana), Oya- poc, Leprieur 56 (P!), 385 (P!); "British Guiana" (Guyana), Melinon 373 (P!), Schomburgk 450 (BM!). Acrostichum schomburgkii Fee, Mem. foug. 2: 32. t. 8,f. 2. 1845. TYPE: "British Guiana" (Guyana), Schomburgk 450 (holotype, P!). Elaphoglossum schomburgkii (Fee) Moore, Index fil. 14. 1857. Stem compact, horizontal, ca. 6-10 mm in di- ameter, scales linear-lanceolate, lustrous, brown to blackish brown, 7-12 mm long, with hairlike processes. Phyllopodia present. Leaves fasciculate, 26-78 cm long, 5.0-7.6 cm broad. Petiole about '/10 the sterile leaf length, with black scales ap- pressed to spreading, 3-6 mm long, some further reduced and skeletonized with irregular hairlike processes. Lamina oblanceolate to broadly elliptic, coriaceous, apex acuminate, base narrowly cune- ate. Veins obscure, free, 1-1.5 mm part, at 80- degree angle. Hydathodes lacking. Lamina abax- ially glabrous, adaxially with black skeletonized scales, especially toward lamina base and reduced to stellate trichomidia. Fertile leaves shorter than the sterile, petiole 2/5 the fertile leaf length, the lamina oblanceolate to elliptic, 3-3.5 cm wide; intersporangial scales lacking. Epiphytic in wet forests, 100-1 200 m, San Mar- tin, Loreto, Pasco, Junin, Ucayali, Madre de Dios. Costa Rica; Panama; West Indies; Trinidad; Guianas; Venezuela; Colombia to Bolivia and Brazil. San Martin: Mariscal Caceres, Palo Blanco, at oeste del Puente, Tocache Nuevo, /. Schunke V. 5759 (NY). Tingo Maria, Allard 21396 (us). Loreto: Maynas, Napo, environs of Rio Santa Maria, on trail S of Secoya village of Vencedor, King 501 (F). Pasco: Oxapampa, Gran Pa- jonal, trail to Shumahuani from Chequitavo, D. Smith 5218 (MO, NY, uc). Junin: La Merced, Killip & Smith 24013 (NU, us). Ucayali: Aguaytia, Ridoutt 13091 (us). Madre de Dios: Mania, Cerro de Pantiacolla, Rio Pa- lotoa, 10-15 km NNW of Shintuya, Foster 10754 (F). 6 1 . Elaphoglossum macilentum Mickel, sp. nov. Ab E. squamipedi lamina sterili anguste elliptica et lamina fertili lineari-elliptica, ulterius stipitis squamis lanceolatis semota. Stem long-creeping, ca. 1 mm in diameter, scales linear-lanceolate, dull orange-tan, 2-4 mm long, entire. Phyllopodia lacking. Leaves 3-14 mm apart, 7-12 cm long, 0.7-1.1 cm broad. Petiole V3-V2 the sterile leaf length, with tan-orange scales widely spreading, ovate-lanceolate, 2-3 mm long, sub- entire, with sparse irregular teeth. Lamina narrow- ly elliptic, chartaceous, apex acute to acuminate, base narrowly cuneate. Veins evident, free, 1 mm apart, at 40-60 degree angle. Hydathodes lacking. Lamina scales very sparse, mostly along abaxial costa. Fertile leaves slightly longer than the sterile, petiole % the leaf length, lamina linear-elliptic, 1.8-2.5 cm long, 4-5 mm broad; intersporangial scales lacking. TYPE— Peru, Madre de Dios, Prov. Manu, Parque Nac. Manu, Rio Manu, Rio Cumerjali, Foster 11978 (holotype, F!). Endemic. On log in wet forest, 350-450 m, Ma- dre de Dios. Similar to E. squamipes except: leaves generally more slender, petiole scales lanceolate to ovate- lanceolate rather than ovate, sterile lamina nar- rowly elliptic instead of ovate-lanceolate, apex acute to acuminate, base cuneate, fertile lamina linear-elliptic, lamina scales very sparse, vein an- gle more acute. Elaphoglossum macilentum differs from E. license (Hooker) Moore (Venezuela, Co- lombia, Ecuador) in the petiole scales appressed, not spreading, and the lamina scales tan, not dark. Thus far known only from the type. 62. Elaphoglossum mathewsii (Fee) Moore, Index fil. 12. 1857. Acrostichum mathewsii Fee, Mem. foug. 2: 54, t. 2,f. 2. 1845. TYPE: Peru, Mathews (holotype, pre- sumably P; frag., B!). Elaphoglossum longipes Brack., U.S. Expl. exped., Fi- lic. 16: 70. t. 9,f. 2. 1854. TYPE: Peru, Andes, Banos, Brackenridge (holotype, us!). Stem short- to moderately creeping, 2-3 mm in diameter, scales linear-lanceolate, dark red-brown to red-black, lustrous, denticulate, ca. 3 mm long. Phyllopodia distinct. Leaves spaced, to 1 cm apart, 12-32 cm long, 1.0-1.8 cm broad. Petiole '/2-2/3 the sterile leaf length, the scales mostly 1-3 mm long, lustrous brown, sclerotic, with pale margin, appressed to slightly spreading, dentate to ciliate near base. Lamina narrowly elliptic, subcoria- ceous, apex acuminate to acute or obtuse, base cuneate. Veins inconspicuous, free, 0.8-1 mm 144 FIELDIANA: BOTANY apart, 60-70-degree angle. Hydathodes present but inconspicuous. Lamina scales of adaxial surface generally abundant and usually overlapping, lan- ceolate to deltate, peltate, brown with pale mar- gins, abaxial surface often essentially glabrous, sometimes glandular, occasionally with scattered scales, these ciliate, especially toward base. Fertile leaves slightly longer than the sterile, petiole %-% the fertile leaf length; scales sparse on abaxial cos- ta, lacking among the sporangia. Terrestrial or epipetric, 3200-5100 m, Caja- marca south to Huancavelica and Madre de Dios. Mexico to Costa Rica; Venezuela and Colombia to Chile. I include here E. longipes, which may be a dis- tinct taxon. However, the only difference seems to be the very long petiole and linear lamina. Elaphoglossum mathewsii specimens: Cajamarca: Hualgayoc, Soukup & Carmons 5015 (GH). Prov. Caja- marca, Encanada, Komolka, Sanchez 193 (GH). Caja- marca, 28 km NW of Cajamarca on road to Pedernal, Ugent & Ugent 5461 (GH). La Libertad: Otuzco, Angulo 1701 (GH). Bolivar, arriba de Bolivar, Lopez & Sagd- stegui 1620 (NY), 3221 (GH). Ancash: Carhuaz, Huas- caran Nat. Park, Quebrada Ishinca, D. Smith & Bud- densiek 11265 (NY). Huanuco: Mito, Bryan 202 (F). Tambo de Vaca, Macbride 4387 (F, us), Bryan 654, 699 (F). Lima: Canta, Carhuai, Acleto 219 (USM). Junin: La Quinua, MacBride & Featherstone 2026 (F, us). Huan- cayo, 1 1 km from Huancayo towards Chameseria, Soun- ders 652 (GH). Huancavelica: Tayacaja, Pampas, Gutte 1074 (USM). Ayacucho: 14 km NE of La Quinua, Luteyn 6339 (NY). Cuzco: Urubamba, trail from Ch'uso to Cuz- co, 2 hrs walk from Chinchero, King et al. 215 (F, NY). Madre de Dios: Pinasniocj, Pantiacolla Pass, Cook & Gilbert 1839 (us). Elaphoglossum longipes specimens: Pasco: Valle del Paucartambo, hacienda Churu, Herrera 1643 (us). Ju- nin: Huancayo, Soukup 2009 (GH). Cuzco: Saxaihuaman hills, Herrera 159 (us). Paucartambo, Hacienda Chura, Herrera 273 (GH, us). Urubamba, Chincheros, S. King et al. 289 (F, NY). Quispicanchi, Marcapata, Vargas 13447 (GH). Prov. Cuzco, San Jeronimo, v.d. Hoogte & Roersch 274 (F). 63. Elaphoglossum megalurtim Mickel, sp. nov. Elaphoglossum auricomum quoad laminae squamas dissectas et stipitem abbreviatum simulans, sed stipitis squamae latiores et laminae apice caudatae sunt. Stem short creeping, ca. 2 mm in diameter, scales lanceolate, lustrous, brown, ca. 1 mm long, entire, more or less resinous. Phyllopodia evident. Leaves approximate, 21-30 cm long, 1.9-2.4 cm broad. Petiole !/6 the sterile leaf length, with tan scales appressed to spreading, 1-2 mm long, cilio-den- ticulate. Lamina narrowly elliptic, chartaceous, apex cuspidate to caudate, base cuneate. Veins ev- ident, free, 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales abaxially and adaxially substellate, adaxially glabrescent with age, abaxial costal scales 1.5-2 mm long, pale, lanceo- late, ciliate. Fertile leaves not known. TYPE— Peru, Pasco, Oxapampa, Quebrada Cas- tilla on the Omaiz river, Leon & Young 1071 (ho- lotype, F!). Endemic. Epiphyte in wet forests, elevation un- known, Pasco. Thus far known only from the type. 64. Elaphoglossum meladenium Mickel, sp. nov. Ab E. latifolio lamina lineari et stipitis squamis au- rantiacis sparsis abstat. Stem creeping, ca. 4 mm in diameter, scales lanceolate, lustrous, orange, tinged dark brown with age, 7-9 mm long, entire. Phyllopodia present. Leaves 2-8 mm apart, 25-41 cm long, 2.5-4.0 cm broad. Petiole 2/5-3/5 the sterile leaf length, with scattered orange scales widely spreading toward petiole base, these 4-6 mm long, entire, the lamina surface also with glandular dots, and with spores adhering to the scales to give an appearance of black dots. Lamina lanceolate to elliptic, charta- ceous, apex acute, base broadly cuneate. Veins ob- scure, free, 1 mm apart, at ca. 80-degree angle. Hydathodes lacking. Lamina scales on both sur- faces with black dots, dark with spores attached plus occasional 1-2 mm ovate scales along costa. Fertile leaves slightly longer than the sterile, pet- iole 3/5-3/4 the fertile leaf length, lamina narrower than the sterile, 1 .9-2.3 cm broad; intersporangial scales lacking. TYPE— Peru, Ancash, Yungay, alrededores La- guna Llanganuco, Mostacero L. et al. 1408 (ho- lotype, NY!; isotypes, F!, HUT!). Endemic. Terrestrial and epipetric in wet for- ests, 3750-4100 m, Ancash, Cuzco. Ancash: Huari, Huascaran Nat. Park, slopes & valley of Laguna Ichicpotrero, D. Smith et al. 12380 (NY). Cuz- co: Calca, Amparaes, SW from Cuzco, Nunez 6726 (MO). 65. Elaphoglossum melancholicum Vareschi, Act. Bot. Venez. 1: 109. /. JOB. 1966. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 145 TYPE: Venezuela, Merida, Paramo Monsalves, Vareschi 2217 (holotype, YEN). Stem wide-creeping, branched, ca. 2-4 mm in diameter, scales linear-lanceolate, lustrous, black, sclerotic, 3-5 mm long, with sparse, irregular teeth. Phyllopodia evident. Leaves 1.5-4 cm distant, (7)18-50 cm long, (0.9)2.8^.8 cm wide. Petiole 04)'/3-'/2 the sterile leaf length, glabrous. Lamina lanceolate to elliptic, coriaceous, apex acuminate, base cuneate to rounded. Veins evident, free, 1 mm apart, at 70-80-degree angle. Hydathodes lacking. Lamina scales on abaxial surface scat- tered, linear to linear-dentate, black, sclerotic, ca. 1 cm long; slightly longer on costa, adaxially with a few by the costa. Fertile leaves slightly longer than the sterile, petiole ('/»)2/3 the fertile leaf length, the lamina about the same shape and width as the sterile or with truncate base; intersporangial scales lacking. Epiphytic and epipetric, 3000-3400 m, La Li- bertad, San Martin, Huanuco, Junin. Venezuela; Colombia; Ecuador; Peru. Elaphoglossum megarhizon Rosenst. of Bolivia is similar, and if they prove to be the same speces, that would be the correct name. Piura: Huancabamba, above Canchaque, Hutchison 1636 (uc). La Libertad: Pataz, Puerta del Monte, Paso La Sabana, Lopez & Sagdstegui 3464 (GH). Pataz, Young 2985 (NY, USM). San Martin: Mariscal Caceres, Rio Abi- seo Nat. Park, Puerta del Monte, Young 1682 (NY). Huanuco: Tingo Maria, Allard 1049 (us). Tambo de Vaca, Bryan 652 (F, us). Junin: Conception, Comas, km 39, road Concepcion-Satipo, Sounders 1049 (GH). Cuzco: Machu Picchu, Soukup 178 (F). 66. Elaphoglossum met a Hi cum Mickel, sp. nov. Lamina vivide azurea utraque facie aequibiliter squa- mulis parvis dissectis nigris obsita singularis. Stem short-creeping, ca. 4-6 mm in diameter, scales linear-lanceolate, lustrous, orange tinged with dark brown, 9-14 mm long, with hairlike pro- cesses. Phyllopodia present. Leaves 3-6 mm dis- tant, 22-67 cm long, 3.7-5.5 cm wide. Petiole '/3-'/2 tne sterile leaf length, with black scales ap- pressed, skeletonized, 1 mm long, with long irreg- ular, processes. Lamina lanceolate to broadly el- liptic, coriaceous, margin inrolled, apex acute to obtuse, base broadly cuneate. Veins evident, free, 1-1.5 mm apart, at ca. 30-degree angle. Hyda- thodes lacking. Lamina scales abaxially with scat- tered skeletonized, reddish-brown scales 1 mm long, adaxially fewer, glabrescent. Fertile leaves shorter than the sterile, petiole !/2 the fertile leaf length, the lamina narrow, 2. 1 cm wide, narrowly lanceolate; intersporangial scales lacking. TYPE— Peru, San Martin, Zepelacio, near Moyo- bamba, Klug3636 (holotype, NY!; isotypes, F!, GH!, K!, MO!, us!). Endemic. Epiphytic (?) in wet forests, 750-1 100 m, San Martin, Huanuco. San Martin: Tarapoto, LI. Williams 5986 (F). Hua- nuco: SW slope Rio Llullapichis watershed on ascent of Cerros del Sira, Dudley 13169 (GH), 13264 (GH). 67. Elaphoglossum minutum (Fee) Moore, Index fil. 12. 1857. Acrostichum minutum Fee, Mem. foug. 2: 39. /. 10, f. 3. 1845. TYPE: Brazil, near Goyaz, Pohl (holo- type, w). Acrostichum unitum Fee, Mem. foug. 2: 44. 1845. SYNTYPES: Peru, Dombey (holotype, P!, photo, us); Mexico, Mt. Orizaba, Galeotti?; "Martini- ca," Desvaux (P?). Acrostichum leptophyllum Fee, Mem. foug. 2: 45. t. 17, f. 1. 1845, not Lam. & DC, 1805. TYPE: Brazil, Blanche! 548 (holotype, P!). Elaphoglossum leptophyllum (Fee) Moore, Index fil. 11. 1857. Stem short- to moderately creeping, 2-3 mm in diameter, scales ovate to (ovate-lanceolate) linear- lanceolate, orange or tan with varying degrees of dark, shiny, sclerotic streaking, ca. 3 mm long, entire or with occasional small, irregular hairlike processes. Phyllopodia distinct. Leaves spaced 1- 8 mm apart, 6-38 cm long, 0.6-1.4 cm broad. Petiole V4-V2 the sterile leaf length, glabrous or with tan, ovate scales. Lamina linear-elliptic, apex acute to obtuse, base attenuate. Veins obscure, free, ca. 1 mm apart, at 60-70-degree angle. Hydathodes lacking. Lamina scales sparse, minute, stellate hairs or dots on abaxial surface, rare on adaxial surface. Fertile leaves longer than the sterile, petiole ('/3-)V2-3/5 the leaf length, the lamina similar to the sterile in size and shape; intersporangial scales lacking. Terrestrial in wet forests, cliffs, grassy slopes, and roadsides, (800-)2400-3700 m, Piura to Amazonas, south to Huancavelica and Puno. Costa Rica; Panama; Jamaica; Hispaniola; Ven- ezuela and Colombia to Bolivia. 146 FIELDIANA: BOTANY Piura: Huancabamba, Mitopampa (Huancabamba- Cuello del Indio), Sagdstegui 8238 (F, HUT, NY, uc). Lam- bayeque: Ferrenafe, 7 km NW of Incahuasi near Cerro Punamachay on trail to Laguna Hualtaco, Dillon & Skill- man 4116A (F). Cajamarca: Contumaza, Tambo La Lima (Cascas-Contumaza), Lopez M. 9029 (F, GH, uc). Ama- zonas: Chachapoyas, Cerros Calla Calla, 45 km above Balsas, road to Leimebamba, Hutchison & Wright 5806 (F, GH, NY, uc). La Libertad: Trujillo, Cerro Campana, Sagdstegui 12917 (F). San Martin: Mariscal Caceres, NW corner of Rio Abiseo Nat. Park, Young & Leon 4685 (USM). Ancash: Huaraz, Huascaran Nat. Park, Quebrada Shallap, D. Smith et al. 10777 (NY). Huanuco: Mito, Bryan 369 (F). Lima: San Bartolome, Bosque de Zarate, Huarochiri, Leon 907 (F, NY). Pasco: Border Prov. Oxa- pampa and Pasco, San Cotardo, van der Werffet al. 8526 (uc). Junin: Huancayo, Kunkel 441 (GH). Huancavelica: Tayacaja, Quebrada SE of Salcabamba, Stork & Horton 10312 (F). Ayacucho: Pampalca, between Huanta & Apu- rimac, Killip & Smith 23267 (NY). Cuzco: "Pillahuata," Cerro de Cusilluyoc, Pennell 141 12 (GH, NY, us). Puno: Sandia, S of Limbani, Metcalf 30437 (uc, us). The following specimens are flocculose abaxi- ally, with sparse, dissected scales resembling tufts of wool. They occur at 1000-2900 m, are on lime- stone or epiphytic. Cajamarca: 25 km E of Cajamarca, Dillon et al. 2892 (F). Pasco: Oxapampa, Canyon de Huancabamba, fundo La Esperanza, Leon 619 (F). Cuzco: La Convention, be- tween Santa Teresa and Chaullay, Nunez & Motocanchi 8789 (MO). Some specimens are larger than the usual, with longer petiole (ca. V2 the leaf length), wider, obtuse lamina, and nearly naked stern. They may repre- sent E. affine (Mart. & Gal.) Moore. These are: Cajamarca: Contumaza, Pampa de la Sal, Sagdstegui 10743 (F, NY). Ancash: Carhuaz, Huascaran Nat. Park, Quebrada Ishinca, Smith & Buddinsiek 11258 (NY). 68. Elaphoglossum moyeri Mickel, sp. nov. Ab E. papilloso lamina anguste elliptica necnon stipite gracili diversum. Named for Dr. Arden Moyer, who is a volunteer in my office, helping with curatorial and research projects. Stem compact, ca. 4 mm in diameter, scales linear-lanceolate, lustrous, dark red-brown, very indurated, entire, ca. 1 mm long. Leaves approx- imate, 22-23 cm long, 1.8-2.5 cm broad. Petiole about '/5 the leaf length, glabrous. Lamina narrowly elliptic, chartaceous, apex long-acuminate, base narrowly cuneate. Veins evident, 1 mm apart, at ca. 70-degree angle. Hydathodes present but in- distinct. Lamina scales abaxially linear, sclerotic, dark red-brown, ca. 1 mm long, sparse along ab- axial costa at maturity, younger leaves with scales scattered on both surfaces. Fertile leaves about equal to or slightly shorter than the sterile, petiole '/2-3/5 the leaf length, lamina narrowly elliptic but narrower than the sterile, 11-17 mm broad; in- tersporangial scales lacking. TYPE— Peru, Pasco, Oxapampa, San Alberto, Cordillera de Yanachaga, van der Werffet al. 8429 (holotype, uc!; isotype, MO!). Endemic. Epiphytic in cloud forest, 2300-2500 m, Pasco. Thus far known only from the type collection. 69. Elaphoglossum muscosum (Sw.) Moore, Index fil. 12. 1857. Acrostichum muscosum Sw., Prodr. 128. 1788. TYPE: Jamaica, Swartz (holotype, s; isotype, B, Herb. Willd. 19523; photo, GH of B). Stem compact, to 12 mm in diameter, scales linear, reddish orange, to 1 3 mm long, with short teeth. Phyllopodia inconspicuous, hidden by scales. Leaves approximate, 1 1-22 cm long, 2.1-2.5 cm broad. Petiole '/3-2/5 the leaf length, scales linear, light orange, erose-denticulate, to 6 mm long, spreading. Lamina narrowly elliptic, subcoria- ceous, apex obtuse, base broadly cuneate to round- ed. Veins obscure, ca. 1 mm apart, at ca. 60-degree angle. Hydathodes lacking. Lamina scales of ad- axial surface white, or with slightly darker centers, peltate, ovate-lanceolate, long-toothed, abundant, on abaxial surface longer, darker orange, less pel- tate. Fertile leaves longer than the sterile, petiole ca. 3/5 the leaf length, lamina slightly narrower than the sterile; intersporangial scales ciliate. Epiphytic in wet forests, 3000 m, Huancavelica, Cuzco. Mexico to Panama; West Indies; Colombia to Peru. This is distinguished by the erose-denticulate scales on the petiole and costa, and the concol- orous lamina scales. Huancavelica: Tayacaja, Ampurco, Tovar 3772 (GH). Cuzco: Machu Picchu, Soukup "177", March 1936 (F). TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 147 70. Elaphoglossum nastukiae Mickel, sp. nov. Ab E. huacsaro lamina adaxialiter densius squamosa abaxialiter glandulosa et squamulis reductis ciliolatis ob- sita discriminatum. Named for Ruth Nastuk, who is a volunteer in my office and the New York Botanical Garden greenhouses. Stem short-creeping, ca. 5 mm in diameter, scales linear-lanceolate, lustrous, black, sclerotic, sub- entire, 2-3 mm long, with sparse, very short teeth. Phyllopodia present. Leaves slightly apart, 1 8-23 cm long, 1 .0-1 .5 cm broad. Petiole ca. '/3 the sterile leaf length, scales subentire, spreading, 2-3 mm long, dark or orange with dark tip, distally smaller and paler, also with very small, roundish, pale, appressed scales. Lamina narrowly elliptic, char- taceous, apex acuminate, base narrowly cuneate. Veins evident, ca. 0.8 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales adaxi- ally abundant, touching but not much overlap- ping, ovate or round, entire to erose, 0.5-1.5 mm long; abaxially with reduced, scattered, fimbriate scales and resin dots, costal scales as on petiole. Fertile leaves longer than the sterile, petiole ca. 3/5 the leaf length, lamina slightly narrower than the sterile; intersporangial scales lacking. TYPE— Peru, Huancavelica, Tayacaja, quebrada SE of Salcabamba, Stork & Morton 10312 (holo- type, uc!). Endemic. Terrestrial "in mossy loam under for- est cover," 3300 m, Huancavelica. Thus far known only from the type. 71. Elaphoglossum nidiformis Mickel, sp. nov. sterile, petiole V2 the fertile leaf length, lamina nar- rowly oblanceolate; intersporangial scales lacking. TYPE— Peru, Madre de Dios, Manu, Cerro de Pantiacolla, Rio Palotoa, 1 0-1 5 km NNW of Shin- tuya, Foster 10891 (holotype, F!). Endemic. Epiphytic in wet forests, 700-1000 m, Madre de Dios. Thus far known only from the type. 72. Elaphoglossum nigrescens (Hooker) Diels, Nat. Pflanzenfam. 1(4): 332. 1899. Acrostichum nigrescens Hooker, Sp. fil. 5: 214. 1864. TYPE: Venezuela (as British Guiana), Roraima, Schomburgk (holotype, K!). Stem short-creeping, 1-2 mm in diameter, scales lacking, stem glutinous. Phyllopodia present, short, ca. 5 mm long. Leaves approximate, 19—40 cm long, 1.1-2.0 cm broad. Petiole V30-V5 the sterile leaf length, glabrous. Lamina linear-elliptic, mem- branous to chartaceous, apex acute to acuminate, base attenuate. Veins evident, free, 1.5-2 mm apart, occasionally elongate laterally at tips but not join- ing, at ca. 60 degree angle. Hydathodes lacking. Lamina scales essentially lacking except on costa and margin, surface glandular-punctate abaxially, costa scales occasional, ovate, brown, under 1 mm long, margin scales occasional, dissected, brown, under 1 mm long. Fertile leaves about equal to the sterile in length, petiole 2/3 the leaf length, lamina linear, 3-5 mm broad; intersporangial scales lack- ing. Epiphytic in wet forests, 100-1200 m, Loreto, Junin, Madre de Dios. Venezuela; Colombia; Ecuador; Peru. Rhizomate valido squamis aurantiacis induto et fronde sessile oblanceolato chartacea nuda distincta. Stem compact, horizontal, ca. 10 mm in di- ameter, scales ovate-lanceolate, lustrous, orange, 4-6 mm long, subentire with sparse irregular teeth. Phyllopodia evident. Leaves fasciculate, 30-37 cm long, 4.0-5.0 cm broad. Petiole essentially lacking, to 1 cm long, with orange, appressed to spreading scales, 4-5 mm long, subentire. Lamina oblanceo- late, chartaceous, apex acute, base narrowly cu- neate. Veins evident, about 1.5 mm apart, at 70- degree angle. Hydathodes lacking. Lamina scales lacking, but abaxial surface with scattered, minute, stellate trichomidia. Fertile leaves longer than the Loreto: Prov. Maynas, Dist. Napo, bank of Rio Santa Maria, King 452 (F, NY). Junin: Satipo, Gran Pajonal, trail to Pajonal Panquerete, S of Chequitavo, D. Smith 5133 (NY, uc). Madre de Dios: Manu, Cerro de Pantia- colla, Foster 10792 (F, GH). 73. Elaphoglossum nivosum (Kunze) Mickel, comb. nov. Acrostichum nivosum Kunze, Bot. Zeit. (Berlin) 1845: 281. TYPE: Venezuela, Caracas, Moritz (holo- type, LZ, destroyed). Stem compact, 4-8 mm in diameter, scales lin- ear-lanceolate, lustrous, sclerotic, black, ca. 5 mm 148 FIELDIANA: BOTANY long, entire. Phyllopodia inconspicuous. Leaves fasciculate (7-) 17-30 cm long, 1.2-2.2 cm broad. Petiole y3-V2 the leaf length, scales with dark center, pale margin, dense, imbricate, ascending, 2-3 mm long, erose. Lamina narrowly elliptic, chartaceous, apex acute to obtuse, base cuneate. Veins obscure, free, 1 mm apart, at ca. 70-degree angle. Hyda- thodes lacking. Lamina scales abaxially densely imbricate, reddish-orange, erose to short cilio- denticulate, 2-3 mm long, costal scales dark, scle- rotic centered, adaxially white, round to ovate, peltate, scattered. Fertile leaves longer than the sterile, petiole 2/3 the fertile leaf length, lamina nar- row, ca. 1 .2 cm broad, costal scales black-centered, cilia longer than the scale body width; interspor- angial scales lacking. Terrestrial on rocky slopes and clay banks, 2600- 3900 m, Cajamarca, La Libertad, Ayacucho, Ju- nin. Venezuela; Ecuador; Peru; Bolivia. Cajamarca: Prov. Cajamarca, Cerro Tunazorco, Sun- chubamba, Cabanillas et al., 26 Aug 1974 (us). Contu- maza, La Herilla Gumango, Sagdstegui et al. 6461 (NY) 9676 (uc). La Libertad: Santiago de Chuco, Cachicadan, Stork & Morton 9951 (F, us). Bolivar, Chomparen, Lopez & Sagdstegui 3203 (GH). Ayacucho: Ccarrapa, between Huanta & Rio Apurimac, Killip & Smith 22284 (NY). Junin: Tarma, Palca, 2 km beyond Yanamayo, Tarma to La Merced road, Saunders 707 (F, GH). Paucartambo, Woytkowski 6688 (us), 6690 (us). 74. Elaphoglossum obovatum Mickel, Brittonia 39: 322. 1987. TYPE: Venezuela, Amazonas, Cerro Yapacana, Steyermark & Bunting 103199 (holotype, us!; isotype, NY!). Stem compact, 3—4 mm in diameter, scales lin- ear, orange, lustrous, 3^4 mm long, margin entire or with a few long hairlike processes. Phyllopodia present. Leaves fasciculate, 6-11 cm long, 2-4 cm broad. Petiole V5-V3 the sterile leaf length, the scales scattered, appressed to spreading, linear, deeply lacerate, orange to brown, lustrous, 2-4 mm long. Lamina obovate, coriaceous, apex broadly round- ed, base cuneate. Veins obscure, free, 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lam- ina scales abaxially scattered, concentrated at mar- gin, skeletonized, orange to brown, dull, ca. 2 mm long, adaxially subglabrous. Fertile leaves shorter (about V2) than the sterile, petiole about V2 the fer- tile leaf length, the lamina ovate, apex obtuse, base rounded, narrower than the sterile lamina; inter- sporangial scales lacking. Epiphytic in wet forests, 130-140 m, Loreto. Venezuela; Peru. The fertile leaf in the only fertile Peru specimen (Rimachi 7613) is slightly different from that of the Venezuelan material; it is ovate with rounded base rather than cuneate. Loreto: Maynas, Dist. Iquitos, Carretera de Santo To- mas, Rimachi Y.7613 (NY). Prov. Requena, Jenaro He- rrera, van der Werff et al. 10029 (uc). Prov. Maynas, Mishana, van der Werff 10177 (uc). 75. Elaphoglossum obtusum Mickel, sp. nov. Ab E. ornato laminae costa ac margine squamis den- ticulatis provisis recognita. Stem compact, horizontal, ca. 4 mm in diam- eter, scales linear-lanceolate, lustrous, orange, 4- 7 mm long, with hairlike teeth. Phyllopodia ca. 4 mm long, but not dark. Leaves fasciculate, 1 5-23 cm long, 3.2-5.5 cm broad. Petiole Vg-V6 the sterile leaf length, with orange scales widely spreading, 3-7 mm long, 1-1.8 mm broad, with long, hairlike teeth. Lamina oblanceolate, chartaceous, apex broadly obtuse, base nearly to broadly cuneate. Veins evident, free, 1-1 .5 mm apart, ca. 70-degree angle. Hydathodes lacking. Lamina scales as on petiole but narrower, on lower surface limited to costa and margin, remaining surface completely naked, upper surface with scales scattered. Fertile leaves about equal the sterile in length, petiole 2/5 the leaf length, lamina much narrower, ca. 1.3 cm broad, narrowly elliptic, apex obtuse, base cune- ate; intersporangial scales lacking. TYPE— Peru, San Martin, Palo Blanco, al oeste del Puente, Mariscal Caceres, Tocache Nuevo, J. Schunke V. 5757 (holotype, NY!; isotype, F!). Endemic. Epiphytic in wet forests, 700-800 m, San Martin. This resembles Venezuelan material cited as E. ornatum (Kuhn) Christ (Smith, p. 99) in the den- tate scales on costa and margin, the remaining lamina naked, and the oblanceolate lamina. How- ever, the Venezuelan plant narrows to an acute apex rather than being broadly obtuse. The cited Venezuelan material is probably not E. ornatum, which differs, according to the original description, in the size and form of the lamina and the laminar scales. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 149 76. Elaphoglossum oculatum Mickel, sp. nov. Ab affini E. muscoso lamina utraque facie nuda, costa abaxialiter squamis parvis rotundis albis nigro-oculatis ciliolatis induta, laminaeque margine squamis imbricatis dentatis albis ciliata praestans. Stem compact, ca. 5 mm in diameter, scales linear-lanceolate, maroon to black, 5-8 mm long, toothed. Phyllopodia present, hidden under scales. Leaves approximate, 18-31 cm long, 1.8-3.9 cm broad. Petiole %-2/5 the sterile leaf length, with scales ovate-lanceolate, erose-denticulate, 2-3 mm long, black or dark orange with white margin, smaller scales appressed. Veins evident, 1 mm apart, at 65-70-degree angle. Hydathodes lacking. Lamina scales lacking on surface abaxially, with marginal scales cilio-denticulate, ca. 1 mm long, adaxial surface glabrous or with greatly reduced scales or nearly substellate trichomes, costal scales round, black with white margins, cilio-denticulate, 0.5-1 mm long. Fertile leaves longer than the ster- ile, petiole about V2 the leaf length, lamina slightly narrower than the sterile; intersporangial scales abundant, ciliate. TYPE — Peru, Piura, Huancabamba, above Huancabamba, road to Canchaque, Hutchison 1629 (holotype, uc!; isotype, GH!). Endemic. Epiphytic in wet forests, 1900-3100 m, Piura, Junin, Cuzco. Junin: Tarma, Agua Dulce, Woytkowski 35433 (MO, uc). Cuzco: Machu Picchu, Soukup "176", March 1936 (F). 77. Elaphoglossum odontolepis Mickel, sp. nov. Rhizomate longe repenti laminaque coriacea secus costam squamis obscuris hinc inde dentatis provisa praestans. Stem long-creeping, ca. 3 mm in diameter, scales linear-lanceolate, lustrous, orange to black-brown, 3-5 mm long, hastate, with occasional hairlike processes. Phyllopodia present. Leaves 1-3 cm apart, 32-50 cm long, 3.4-5.5 cm broad. Petiole ca. '/3 the sterile leaf length, glabrous. Lamina nar- rowly oblanceolate to narrowly lanceolate, coria- ceous, apex acuminate, base attenuate. Veins ob- scure, free, 1 mm apart, at ca. 80-degree angle. Hydathodes lacking. Lamina scales abaxially scat- tered, especially near lamina base and along costa, 2-3 mm long, hastate, black, with irregular pro- cesses, and with black punctae, also adaxially black- punctate and stellate-punctate. Fertile leaves near- ly equalling the sterile in length, petiole about V3 the leaf length, lamina nearly equal to the sterile in size and shape; intersporangial scales scattered, black, sclerotic, with irregular teeth. TYPE— Peru, Cuzco, Urubamba, Machu Picchu, E of Phuyupatamarca overlooking Urubamba River and Winay Huayna, Peyton & Peyton 356 (holotype, MO!). Endemic. Epiphytic in wet forests, 3200-4000 m, San Martin, Cuzco. San Martin: Mariscal Caceres, NW corner of Rio Abi- seo Nat. Park, Chochos, forest on edge of Laguna de Chochos, Young & Leon 4866 (USM). Cuzco: La Con- vention, Huayopata 7 km from Incatambo, S side of Lucumayo River, Peyton & Peyton 958 (GH). 8 km SE of Machu Picchu, Whiteman, Aug 1980 (NY). 78. Elaphoglossum oophyllum Mickel, sp. nov. Rhizoma longe repens et lamina magna ovato-lanceo- lata squamis minutis nigris conspersa diagnoscenda. Stem long creeping, ca. 3 mm in diameter, scales linear-lanceolate, dull orange-tan, 3-4 mm long, with hairlike processes. Phyllopodia present. Leaves 5-10 mm apart, 23-32 cm long, 5.5-6.6 cm broad. Petiole about !/2 the sterile leaf length, scales black, linear-lanceolate, appressed to spreading, 2—4 mm long, with irregular hairlike processes. Lamina ovate to ovate-lanceolate, char- taceous, apex acute, base rounded. Veins obscure, free, 1 mm apart, at 70-80-degree angle. Hyda- thodes lacking. Lamina scales abaxially reduced to orange stellate trichomidia, along costa scales black, linear-lanceolate, with irregular hairlike processes, adaxially glabrous. Fertile leaves longer than the sterile, petiole 2/3-% the fertile leaf length, lamina narrowly elliptic; intersporangial scales lacking. TYPE— Peru, Cuzco, La Convention, Ichiquia- to, Vargas C. 22337 (holotype, GH!, isotype, GH!). Endemic. Epiphytic(?) in wet forests, 1000 m, Cuzco. Thus far known only from the type collection. 79. Elaphoglossum orbignyanum (Fee) Moore, Index fil. xvi. 1857. 150 FIELDIANA: BOTANY Acrostichum orbignyanum Fee, Mem. foug. 2: 56. /. 13, f. 2. 1845. TYPE: Bolivia, Yungas, Chupe, Marancel del Monte, Orbigny 138 (holotype, P!). Stem compact, 6-12 mm in diameter, scales lan- ceolate-deltate, black, very indurated, lustrous, ca. 5 mm long, with weak, tan cilia along margin. Phyllopodia present but hidden by scales. Leaves approximate, (30-)75-100 cm long, 4-7 cm broad. Petiole about 2/5 the leaf length, scales dense, ap- pressed to ascending, lanceolate to linear-lanceo- late, ciliate, orange, some red-centered, to 7 mm long at petiole base, but mostly 2-3 mm long. Lamina elliptic, chartaceous, apex gradually to abruptly acuminate, base broadly cuneate. Veins ca. 1 mm apart, at 80-degree angle, obscured by scales. Hydathodes lacking. Lamina scales adax- ially barely touching one another, lanceolate to ovate-lanceolate, 1-3 mm long, orange, ciliate, abaxially densely imbricate, ciliate, orange, often red-centered, 1-3 mm long, costal scales lanceo- late to linear-lanceolate, 2-4 mm long, red-cen- tered, appressed. Fertile leaves nearly equalling the sterile in length, petiole % the leaf length, lam- ina elliptic, narrower than the sterile; interspor- angial scales lacking. Terrestrial in wet forests, (680-) 1350-3200 m, San Martin, Junin, Cuzco. Venezuela; Colombia; Peru; Bolivia. This differs from E. hickenii in that the latter has stem scales nonciliate, petiole and costal scales erose to short-ciliate, and lamina narrower (2.5- 4.1 cm). San Martin: San Roque, LI. Williams 7418 (us). Ju- nin: Colonia Perene, Killip & Smith 24954 (NY, uc, us). Carapata, above Huacapistana, Killip & Smith 24413 (NY). Chanchamayo Valley, C. Schunke 134, 135 (us). Tarma, Agua Dulce, Woytkowski 35438 (MO, uc). Cuzco: Machu Picchu, Coronado 141 (uc). Prov. Cuzco, Ribera del Rio Urubamba, Angulo 1773 (GH). Machu Picchu, Soukup "175" (F). San Miguel, Urubamba Valley, Cook 6 Gilbert 11 59 (us). 80. Elaphoglossum oxyglossum Mickel, sp. nov. Ab E. zebrino, cui arete afnnis, rhizomate compacto et laminae squamis brevioribus abstat. Stem short-creeping, ca. 3 mm in diameter, with white patches on stem and petiole bases, scales lanceolate, maroon-black, lustrous, 0.5-1 mm long, entire. Phyllopodia lacking. Leaves to ca. 2 mm apart, 55-65 cm long, 2.0-2.5 cm broad. Petiole about !/2 the sterile leaf length, with sparse linear- lanceolate, appressed, blackish scales ca. 1 mm long. Lamina linear-elliptic, chartaceous, apex acuminate, base cuneate. Veins evident, free, ca. 1.5 mm apart, at 60-70-degree angle. Hydathodes distinct. Lamina scales lacking adaxially except for sparse, tan ones 1.5 mm long, along margin near hydathodes, abaxially sparse, small, lanceolate, dark to pale, 0.5-1 mm long on costa and lamina. Fertile leaves slightly shorter than the sterile, pet- iole % the fertile leaf length, lamina apex acute, base cuneate, scales 1-1.5 mm long, maroon on costa; intersporangial scales pale. TYPE— Peru, Pasco, Oxapampa, road from Oxa- pampa to Villa Rica, Skog et al. 5099 (holotype, us!). Endemic. Epiphytic in wet forest, ca. 1500- 2150(-3400) m, San Martin, Huanuco, Pasco. San Martin: Mariscal Caceres, Young 3783 (USM). Huanuco: Huacachi, near Muna, Macbride 4129 (F, us). Muna, Bryan 538 (F). Cushi, Macbride 4818 (F, us). 8 1 . Elaphoglossum pachyphyllum (Kunze) C. Chr., Index fil. 312. 1905. Acrostichum pachyphyllum Kunze, Linnaea 9: 26. 1834. TYPE: Peru, Pampayacu, Poeppig, July 1829 (holotype, LZ, destroyed; isotype, P!; photos, F!, NY!). Hymenodium kunzeanum Fee, Mem. foug. 2: 90, t. 58. 1845, based on Acrostichum pachyphyllum Kunze and with the same type. Stem short-creeping, (3) 6-10 mm in diameter, scales linear-lanceolate, dull brown, 10-20 mm long, with hairlike processes. Phyllopodia present. Leaves 1-2.5 cm distant, 50-100 cm long, 0.9-1.5 cm broad. Petiole V3-V2 the leaf length, scales at base scattered, orange, spreading, 5-10 mm long, with hairlike processes, scales distally reduced, ob- scure, finely skeletonized, appressed. Lamina broadly lanceolate, coriaceous, apex acute to acu- minate, base truncate to broadly cuneate. Veins barely visible, anastomosing, 1 mm apart, at 80- degree angle. Hydathodes lacking. Lamina scales lacking except for finely skeletonized, orange ap- pressed ones along costa. Fertile leaves shorter than the sterile, petiole V2 the leaf length, the lamina similar in shape to the sterile; intersporangial scales lacking. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 151 Endemic. Epiphytic in wet forests, 680-2700 m, Amazonas, San Martin, Huanuco, Junin, Cuzco, Madre de Dios. Amazonas: 12-18 km E of La Peca in Serrania de Bagua, Gentry et al. 22886 (MO). San Martin: Mariscal Caceres, Rio Abiseo Nat. Park, Young & Leon 5021 (USM). Huanuco: Pampayacu, Kanehira 126 (GH). Junin: Chanchamayo Valley, C. Schunke 175 (F). La Merced, Hacienda Schunke, Macbride 5773 (F). Colonia Perene, Killip & Smith 24952 (F, GH, NY, us). Cuzco: Prov. Pau- cartambo, Valle de Pillahuata, Herrera 1615 (us). Madre de Dios: Manu, Cerro de Pantiacolla, Rio Palotoa 10- 15 km NNW of Shintuya, Foster 10889A (F, GH). 82. Elaphoglossum pachyrrhizum Mickel, sp. nov. Rhizomate valido longe repenti, lamina lineari ob- scure venosa necnon laminae squamis lanceolatis badiis ab affinibus remota. Stem long-creeping, to 4 mm in diameter, scales tan, lanceolate, 3-5 mm long, entire. Phyllopodia lacking. Leaves distant, 18-30 cm long, 1.3-1.7 cm broad. Petiole V4-l/3 the leaf length, scales im- bricate, lanceolate, tan to brown, 1-2 mm long, entire, appressed to somewhat spreading. Lamina linear-elliptic, apex acuminate, base cuneate. Veins indistinct, free, 1-2 mm apart, at 50-70-degree angle. Hydathodes evident. Lamina scales brown, lustrous, lanceolate, 1-2 mm long, mostly along costa, deciduous on abaxial surface but with min- ute punctae or stellate trichomidia, those of the margin entire, ca. 1 mm long, mostly appressed. Fertile leaves shorter than the sterile, petiole about 2/3 the fertile leaf length, lamina apex acute, base subtruncate or rounded; intersporangial scales lacking. TYPE— Peru, Amazonas, Bagua, Cordillera Co- lon, SE of La Peca, Barbour 4111 (holotype, MO!). Endemic. Terrestrial in wet forests, 2350-2450 m, Amazonas. Thus far known only from the type. 83. Elaphoglossum paleaceum (Hooker & Grev.) Sledge, Bull. Brit. Mus. (Nat. Hist.) Bot. 4(2): 95. 1967. Figure 25a-b. Acrostichum paleaceum Hooker & Grev., Icon. fil. 2: t. 235, Alph. Index et Syst. Index. 1831. TYPE: the original figure, drawn from Madeira, Lowe (not located). Acrostichum vestitum Hooker & Grev., Icon. fil. 2: text for/. 235. 1831, not Schlecht.& Cham. 1830. Stem compact, horizontal, 2-6 mm in diameter, scales linear-lanceolate, brown to black and opaque, lustrous, ca. 5 mm long, margin with long, stiff, hairlike teeth. Phyllopodia distinct. Leaves ap- proximate, 10-28 cm long, 0.8-1.6 cm broad. Pet- iole V4-l/3 the sterile leaf length, densely to sparsely clothed with spreading, black or orange scales, these often with black teeth and tip, to 3 mm long. Lam- ina narrowly elliptic, apex acuminate, base broad- ly to narrowly cuneate. Veins obscure, free, ca. 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales abundant, overlapping but loose, lanceolate, ca. 2 mm long, long hair-toothed, usually those of the petiole and costa dark. Fertile leaves shorter than the sterile, to 20 cm long, pet- iole V2-2/3 the fertile leaf length, but with same lam- ina shape or more obtuse; intersporangial scales lacking. Epiphytic and terrestrial on rocky banks in wet forests, 950-3950 m, Piura to Amazonas, south to Cuzco and Puno. Mexico to Panama; West Indies; Venezuela and Colombia to Bolivia and Brazil; Azores and Ma- deira. There is considerable variation in this species, and I take a rather broad view of it here. Most of the specimens agree with the measurements cited above. These may match E. plicatum (Cav.) C. Chr., in which case that is the correct name, but closer study is needed. Plants with larger, broader leaves (26-61 cm long, 2.4—4.1 cm broad), often with a caudate apex (examples cited below with asterisk), may be E. molle (Sodiro) C. Chr. (Type from Ecuador; isotype, s!). Piura: Huancabamba, above Canchaque on the road to Huancabamba, Hutchison 1648 (GH). Cajamarca: Co- losay, Woytkowski 7020 (MO). Hualgayoc, Soukup-Car- mona Fa5234* (us). Amazonas: Chachapoyas, Cerros Calla Calla, 1 9 km above Leimebamba on road to Balsas, Hutchison & Wright 5571 (F, GH, NY, uc, us). San Mar- tin: Mariscal Caceres, Chochos, NW corner Rio Abiseo Nat. Park, Young 3592* (NY). Rioja, Pedro Ruiz-Moyo- bamba road, D. Smith 4355* (uc). Huanuco: Rio Llulla- pichis watershed on the ascent of Cerros del Sira, Dudley 13340A* (GH). Tambo de Vaca, Bryan 649* (F, GH). Muna, Macbride 4301* (us). Pasco: Oxapampa, Abra los Mellizos, Skog et al. 5046a (us). Cuzco: Urubamba, ruins of Machu Picchu, Tryon & Tryon 5397 (GH). Cor- dillera Vilcabamba, 28 km NE of Hda. Luisiana and Apurimac River, Dudley 11188 (us). Montana de Colca, Valle de Lares, Sues 1794 (us). Puno: Carabaya, valle de San Gaban, Hda. Quillabamba a Pte. Arica, Vargas C. 18918* (GH). 152 FIELDIANA: BOTANY 84. Elaphoglossum palorense Rosenst., Repert. Spec. Nov. Regni Veg. 7: 149. 1909. TYPE: Ecuador, Cordillera oriental ad flumen Pa- lora, Rimbach 66 (holotype, s!). Stem short-creeping, ca. 4 mm in diameter, scales linear-lanceolate, lustrous, castaneous, 2-3 mm long, entire. Phyllopodia lacking. Leaves approx- imate to only slightly apart, (15-)35-42 cm long, 0.8-1.1 cm broad. Petiole ca. '/10 the leaf length, scales linear, orange-tan, 2-3 mm long, ca. 0.4 mm broad, patent. Lamina linear, chartaceous, apex acuminate, base long-attenuate, margin entire to barely crenulate. Veins evident, 1.5-2 mm apart, at ca. 65-degree angle. Hydathodes present but indistinct. Lamina scales linear, orange-tan, scat- tered on abaxial costa, to 1 mm long, very sparse on abaxial surface and margin, to 0.5 mm long. Fertile leaves shorter than the sterile, petiole % the leaf length, lamina narrowly elliptic, apex apicu- late, base broadly cuneate, 0.9-1.1 cm broad; in- tersporangial scales lacking. Epiphytic in wet forests, 1450-1800 m, San Martin, Huanuco. Ecuador; Peru. Similar to E. eximium except: petiole and lam- ina scales not subulate and dark, but tan-orange, linear, not punctate abaxially; fertile leaf more nar- rowly elliptic, the lamina 6-7.5 cm long, 0.9-1.1 cm broad, and with cuneate base. San Martin: Rioja, Venceremos, D. Smith 4434 (F). Huanuco: SW slope of Rio LLullapichis watershed on the ascent of Cerros del Sira, Dudley 13198 (GH). 85. Elaphoglossum papillosum (Baker) Christ, Monogr. Elaphoglossum. 130. 1899. Acrostichum papillosum Baker. J. Hot. 15: 167. 1877. TYPE: Ecuador, Andes of Quito, Sodiro (holo- type, K). Stem short-creeping, 4 mm in diameter, scales black, lanceolate-deltate, 1-2 mm long. Phyllo- podia lacking. Leaves approximate, 34—38 cm long, 5.2-5.7 cm broad. Petiole '/3 the sterile leaf length, scales scattered, appressed, black-brown, lanceo- late, to 1 mm long. Lamina elliptic, chartaceous, apex acuminate, base cuneate. Veins distinct, free, ca. 1.5 mm apart, at ca. 70-degree angle. Hyda- thodes evident. Lamina scales on both surfaces brown-black, lanceolate, 0.5 mm long, scattered adaxially, sparse abaxially. Fertile leaves longer, lamina oblanceolate, apex acute, base attenuate, petiole 3/5 the fertile leaf length; intersporangial scales lacking. Epiphytic or terrestrial in wet forests, 2650-2750 m, San Martin. Costa Rica; Panama; Venezuela and Colombia to Bolivia. Similar to E. castaneum except: petiole of fertile leaf gray-green (dried), similar to those of sterile leaves, not black; laminar scales and petiole scales short, lanceolate, dark, not subulate and toothed; stem scales black, not castaneous. San Martin: Prov. Mariscal Caceres, Rio Abiseo Nat. Park, Young & Leon 5007 (USM). 86. Elaphoglossum pascoense R. Tryon. Amer. Fern J. 74. 108. 1984. TYPE: Peru, Oxapam- pa, road between Oxapampa and Villa Rica, Foster 9127 (holotype, GH!; isotypes, MO!, NY!). Stem long-creeping, 5-7 mm in diameter, scales dark, appressed, resinous, dark brown, entire, 1- 2 mm long, sparse except at apex. Phyllopodia lacking. Leaves 28-54 cm long, 6.5-12 cm broad. Petioles distant, long-decurrent on the stem, V2-2/3 the sterile leaf length, scales abundant, appressed, tan, ovate to linear-lanceolate, finely ciliolate. Lamina oblong-ovate, apex caudate, base deeply cordate to sagittate. Veins distinct, free, ca. 2 mm apart, at 75-80-degree angle. Hydathodes evident. Lamina scales sparse, lanceolate, light brown, den- ticulate, but broader, more dense, minutely cilio- late, and appressed on costa. Fertile leaves equall- ing the sterile in length, petiole 3/4 the fertile leaf length, lamina lanceolate, 1 2 cm long, 2 cm broad, apex acuminate, base truncate; intersporangial scales elongate, brownish, ciliate. Endemic. Terrestrial in wet forests, 2100-2400 m, Pasco. Thus far known only from the type collection. 87. Elaphoglossum patinii (Baker) Christ, Mon- ogr. Elaphoglossum 58. 1899. Acrostichum patinii Baker, in Hooker & Baker, Syn. fil. ed. 2: 519. 1874. SYNTYPES: Andes of New Granada, Patin (K); Peru, Cuzco, Pintobamba, Pearce(BM; photo, us). TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 153 Stem short-creeping, 2-3 mm in diameter, scales tan-orange, linear-lanceolate, spreading, dull, 6-8 mm long, 1-1.5 mm wide, entire. Phyllopodia present, hidden among scales. Leaves 1-1.5 cm apart, 25-28 cm long, 1.7-2.0 cm broad. Petiole '/is-'/io the leaf length, glabrous. Lamina linear- elliptic, coriaceous, apex narrowly obtuse, base narrowly cuneate. Lamina scales abaxially sparse, black, sclerotic, hastate, ca. 1 mm long, adaxially lacking. Fertile leaves shorter than the sterile, pet- iole !/5 the leaf length, lamina broader than the sterile, 2.0-2.4 cm broad, apex narrowly obtuse, base rounded; intersporangial scales lacking. Epiphytic in wet forests, elevation unknown, Cuzco. Colombia; Peru. Possibly distinct from the Colombian syntype, in which case the Patin collection should be des- ignated the lectotype and the Peru specimen re- named. It resembles E. velongum in the linear, coriaceous, glabrous lamina and orange stem scales, but E. patinii has longer, more spreading stem scales, more slender petiole, and lacks petiole scales. 88. Elaphoglossum petiolosum (Desv.) Moore, In- dex fil. 12. 1857. Acrostichum petiolosum Desv., Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Na- turk. 5: 309. 1811. TYPE: Peru, /. de Jussieu (holotype, P!). Acrostichum acuminatum Poir. in Lam., Encycl. Suppl. 1: 120, Sept. 1810, (not Willd. 1810). TYPE: Peru, J. de Jussieu (holotype, P). Acrostichum caudatum Hooker, Icon. pi. t. 215. 1 840, not Cav. 1802. TYPE: Colombia, Andes, Pilz- hum, Jameson (holotype, K!). Elaphoglossum caudatum (Hooker) Moore, Index fil. 7. 1857. Elaphoglossum glutinosum Christ. Monogr. Elapho- glossum 120. 1899. TYPE: Andes of Quito, Ec- uador, Spruce 5614 (isotype, B!). Stem short-creeping, 3-4 mm in diameter, scales linear, lustrous, red-black, 6-10 mm long, entire or with very sparse teeth (1-2 per scale). Phyllo- podia distinct. Leaves 1-5 mm apart, 15-44 cm long, 1.8-3.0 cm broad. Petiole '/2-2/3 the sterile leaf length, with orange scales appressed to spread- ing, 2 mm long, with hairlike teeth, also glandular trichomes, especially at base of petiole. Lamina lanceolate to elliptic, subcoriaceous, apex long- cuspidate, base rounded to broadly cuneate. Veins evident, free, ca 1 mm apart, at 70-80-degree an- gle. Hydathodes lacking. Lamina scales lanceolate, cilio-denticulate, 1 mm long, scattered on adaxial surface, abaxially glabrous except for glandular dots, costa and margin with scales 2 mm long. Fertile leaves nearly equalling the sterile in length, petiole 2/3 the fertile leaf length, lamina similar to sterile in size and shape, shorter-cuspidate, con- duplicate; intersporangial scales lacking. Epiphytic in wet forests, 3050-3950 m, Ama- zonas, La Libertad, San Martin, Huanuco, (Ju- nin?), Cuzco. Colombia to Bolivia. Amazonas: Chachapoyas, Cerros Calla Calla, 26 km above Leimebamba on road to Balsas, Hutchison & Wright 6977 (F, GH, NY, uc, us). Prov. Chachapoyas, S side of Molinopampa-Diosan pass, Wurdack 1610 (us). La Libertad: Pataz, Puerta del Monte, Paso La Sabana, Lopez & Sagdstegui 3465 (GH). San Martin: Mariscal Caceres, forest patch above timberline, Puerta del Mon- te, Young 1619 (NY). Mariscal Caceres, N side of Cho- chos Valley, NW corner of Rio Abiseo Nat. Park, Young 3660 (USM). Mariscal Caceres, small forest patch above timberline, Chochos, Young 2281 (NY). Mariscal Cace- res, Puerta del Monte, NW corner of Rio Abiseo Nat. Park, Young & Leon 4918 (USM). Dist. Huallaga, Valley of Rio Apisoncho above Jucusbamba, Hamilton & Hol- ligan 504, 300 (K). Huanuco: Tambo de Vaca, Bryan 653 (F, us). Cuzco: Lucumayo Valley, Cook & Gilbert 1343 (us). Department unknown (probably Junin): Between Arma (Tarma?) & Yanama (Yanano?), Bingham 2060 (us). 89. Elaphoglossum piloselloides (Presl) Moore, Index fil. 13. 1857. Figure 25d-e. Acrostichum piloselloides Presl. Reliq. haenk. 1: 14, /. 2,f. 1. 1825. TYPE: Hab. in montanis Peruviae, Haenke (holotype, PR or PRC). Stem erect or ascending, ca. 2 mm in diameter, scales linear, orange, entire, 3-6 mm long. Phyl- lopodia lacking. Leaves fasciculate, 2-4 (8) cm long, 0.4—1.0 cm broad. Petiole slender, usually 2/3-% the sterile leaf length, moderately clothed with tan to orange, very narrow, subulate scales 2-4 mm long. Lamina spatulate, obovate-lanceolate to nar- rowly elliptic, apex obtuse, base broadly cuneate. Veins obscure, free, ca. 1 mm apart, at 40-50- degree angle. Hydathodes present but inconspic- uous. Lamina scales somewhat subulate (slightly rolled at base) tan to dull orange, uniformly and moderately covering the lamina surfaces. Fertile leaves equal to or longer than the sterile, more erect, petiole ca. % the fertile leaf length, lamina smaller than the sterile, often folded in half, spatu- late to nearly round, scales of lamina and upper 154 FIELDIANA: BOTANY petiole dark brown to black, lustrous, scale bases not inrolled but spreading and denticulate; inter- sporangial scales lacking. Terrestrial or epipetric, on rock cliffs, open woods, clay banks, dripping cliffs, 750-2600 m, Piura to Cajamarca, south to Huancavelica and Cuzco. Mexico to Panama; West Indies; Suriname; Venezuela; Colombia to Chile. This differs slightly from E. spatulatum (Bory) Moore of Bourbon Island and Africa and often is placed under that name. Piura: Huancabamba, just below summit of Abra Por- culla Pass, Mesones-Muro Hwy., Hutchison 1385 (uc, us). Lambayeque: 46 km from Olmos on road to Jaen, Correll & Smith P823 (us). Cajamarca: Jaen, Granadi- llas, in Quebrada Granadillas above Tabaconas, 1 8 km SE Huancabamba, Fosberg 27832 (us). La Libertad: Otuzco, Huaranchal, Lopez et al. 2692 (GH). Huanuco: Mito, steep banks on NW slopes, Macbride 3281 (F, us). Pasco: Oxapampa, Palcazu, Foster 10230 (F). Junin: Tar- ma, La Merced road, 46 km from Tarma, D. Smith & Canne 5942 (F, uc). Huancavelica: Tayacaja, near Sal- cabamba, Tovar 3592 (GH, USM). Ayacucho: Ccarrapa, between Huanta & Rio Apurimac, Killip & Smith 22431 (NY, us). Cuzco: San Miguel, Urubamba Valley, Cook & Gilbert 1181 (us). 90. Elaphoglossum pilosius Mickel, Brittonia 39: 324. 1987. TYPE: Venezuela, Merida, trail leading from La Negrita to the Boqueron of the Quebrada de las Canas, Luteyn 6139 (ho- lotype, NY!). Stem compact, ca. 4 mm in diameter, scales linear-lanceolate, dark reddish-brown, lustrous, 6- 8 mm long, denticulate. Phyllopodia lacking. Leaves fasciculate, (6)9-16 cm long, 1.4-2.7 cm broad. Petiole '/2-2/3 the sterile leaf length, with scales patent, subulate, reddish-brown, lustrous, 2-3 mm long, denticulate, mixed with minute glandular trichomes. Lamina lanceolate to ovate- lanceolate or deltate-lanceolate, chartaceous, apex cuspidate, base rounded or truncate. Veins evi- dent, free, 1 mm apart, at 60-75-degree angle. Hy- dathodes present. Lamina scales scattered, subu- late, reddish-brown, lustrous, 2-3 mm long on adaxial surface, costa and margin, 1-1.5 mm long on abaxial surface. Fertile leaves longer than the sterile, petiole % the fertile leaf length, lamina ob- long, apex obtuse, base rounded to truncate, often folded along costa (conduplicate), costal scales dense, subulate, but intersporangial scales lacking. Terrestrial or epipetric on roadbanks and rocky slopes in grassy shrubland, 2900-4000 m, Caja- marca, Ancash. Costa Rica; Panama; Venezuela; Colombia to Peru. Elaphoglossum pilosius appears to be distinct from E. lindenii on the basis of its greater pubes- cence and rounded to truncate lamina base, but more extensive population samples are needed to settle the matter. Cajamarca: San Miguel, El Tingo, Sagdstegui 8824 (F). Ancash: Yungay, 25 km E of Yungay, Edwin & J. Schunke V. 3810 (F, GH, us). Yungay, Huascaran Nat. Park, Que- brada Ranincuray, D. Smith et al. 10470b (NY). 9 1 . Elaphoglossum plumosum (Fee) Moore, Index fil. 364. 1862. Acrostichum plumosum Fee, Mem. foug. 2: 54. t. 20, f. 1. 1845. TYPE: Guyana (as French Guiana), Schomburgk 446 (isotype, BM!). Stem compact, horizontal, to 10 mm in diam- eter, scales linear-lanceolate, pale, ca. 5 mm long, margins with long, hairlike teeth. Phyllopodia dis- tinct. Leaves fasciculate, 16-36 cm long, 2.2-3.7 cm broad. Petiole '/10-% the sterile leaf length, densely clothed with spreading orange scales sim- ilar to those of the stem but broader, scales to 5 mm long. Lamina narrowly elliptic to narrowly oblanceolate, apex acute to obtuse, base acumi- nate. Veins obscure, free, ca. 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales abundant, overlapping, lanceolate, ca. 2-3 mm long, orange, long hair-toothed, completely covering abaxial side of leaf and sporadic to lack- ing on adaxial side, denser along costa. Fertile leaves about equal the sterile in length, petiole about '/2 the fertile leaf length, lamina ca. 2 cm broad, lanceolate, scales along abaxial costa, in- tersporangial scales lacking. Epiphytic in wet forests, 120-150 m, Loreto. Guianas; Venezuela; Colombia; Ecuador; Peru; Brazil. Loreto: Maynas, Rio Nanay, Caseria Mishana, 30 km SW of Iquitos, Foster 4461 (F). Maynas, Dist. Iquitos, Carretera de Varillal, km 6 de Quisto Cocha al caserio de Varillal, Rimachi Y. 7845 (NY). Maynas, Iquitos, road beyond Quista Cocha, McDaniel 13593 (GH). Vic. Iqui- tos, Revilla 4317 (uc). TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 155 92. Elaphoglossum poeppigianum (Fee) Moore, Index fil. 13. 1857. Acrostichum villosum var. poeppigianum Fee, Mem. foug. 2: 50, t. 20, f. 2. 1845. TYPE: Peru, ad Pampayacu, Poeppig 190 (holotype, P!; isotype, B, uc!). Acrostichum poeppigianum (Fee) Fee, Gen. fil. 43. 1852. Stem compact, horizontal, ca. 3 mm in diam- eter, scales linear, brown, lustrous, 3-5 mm long, entire. Phyllopodia lacking. Leaves fasciculate, 28- 37 cm long, 11-16 cm broad. Petiole %(Vio) the sterile leaf length, with scales abundant, orange- brown, widely spreading, subulate, slightly toothed, 2-4 mm long. Lamina linear-elliptic, chartaceous, apex acuminate, base cuneate to attenuate. Veins distinct, free, 2-3 mm apart, at 60-degree angle. Hydathodes evident. Lamina scales scattered, red- dish orange, subulate, slightly toothed, 1-2 mm long on lamina and margin, 2-3 mm long on costa. Fertile leaves about equal the sterile in length, petiole % the fertile leaf length, lamina elliptic, apex apiculate, base rounded; intersporangial scales lacking. Endemic. Terrestrial in wet forests, 2085-2700 m, Huanuco, Pasco, Junin. The van der Werff specimen most closely re- sembles the type; the Ellenberg collection differs in having an attenuate lamina base and very short petiole. Huanuco: Carpish, Ellenberg 3930 (GH). Carpish, Sandeman 5 157 a (K). Pasco: Prov. Oxapampa, San Al- berto, Cordillera de Yanachaga, van der Werff et al. 8450 (MO, uc). Junin: La Merced-Chanchamayo, Soukup 1017 (F). Department unknown: Toccachillo, Jelski 1078 (P). 93. Elaphoglossum propinquum (Kuhn) Christ, Monogr. Elaphoglossum. 115. 1899. Acrostichum propinquum Kuhn, Linnaea 36: 45. 1869. TYPE: Peru, near Agapata, Lechler 2007 (holo- type, B!; isotype, P!; photos, F & us of P). Stem compact, to 8 mm in diameter, scales cas- taneous, stiff to crispate, to 12 mm long. Phyllo- podia lacking. Leaves fasciculate, 15-39 cm long, 2.6-4.5 cm broad. Petiole stramineous, l/3-V2 the sterile leaf length, with scales subulate, spreading, sparse to lacking, petiole also with a few minute, erect, glandular trichomes. Lamina narrowly lan- ceolate, chartaceous to subcoriaceous, apex cau- date, base broadly cuneate to rounded. Veins ev- ident, free, 1.5-2 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales black, subu- late, mostly along the margin, generally early de- ciduous, mature specimens wtih only a few scales or scales totally lacking, lamina also with minute, stellate trichomidia. Fertile leaves shorter than the sterile, petiole ca. 2/3 the leaf length, lamina the same shape as the sterile but slightly narrower; intersporangial scales lacking. Endemic. Epiphytic or epipetric in wet forests, 1500-3000 m, Amazonas, Huancavelica, Cuzco, Puno. Amazonas: Bagua, Cordillera Colon near La Peca, Bar- hour 4175 (MO). Huancavelica: Tayacaja, Chuspi, Tovar 2036 (GH, USM). Cuzco: Calahuala, Machu Picchu, Cook & Gilbert 881 (us). Torontoy, Herrera 1314 (us). 94. Elaphoglossum pumilio Mickel, sp. nov. Ab E. tenuiculo lamina adaxialiter squamis peltatis, abaxialiter squamis stellatis cum nonnullis ovato-resi- noso-sclerotico-ciliatis induta diversa. Stem short-creeping, ca. 2 mm in diameter, scales linear-lanceolate, lustrous, castaneous to dark brown, 2-3 mm long, entire. Phyllopodia distinct. Leaves 1-2 mm apart, (5)1 1-27 cm long, 0.4—0.6 cm broad. Petiole !/3-4/5 the sterile leaf length, with castaneous scales ascending to spreading, resinous, 1-2 mm long, with sparse irregular short teeth, the scales mixed with glands and peltate, ciliate, pale scales. Lamina linear, chartaceous, apex acumi- nate to obtuse, base cuneate. Veins barely visible, free, ca. 1 mm apart, at 60-70-degree angle. Hy- dathodes lacking. Lamina scales abaxially reduced to stellate trichomes with a slight scale body, both surfaces with scattered peltate, round to ovate- lanceolate, red-brown, resinous, ciliate scales; cos- ta abaxially with scattered lanceolate lustrous scales. Fertile leaves equal to or slightly longer than the sterile in length, petiole %-'/2 the fertile leaf length, the lamina narrow, 0.3-0.5 cm broad; intersporangial scales lacking. TYPE— Peru, Huanuco, Yanano, Macbride 4943 (holotype, NY!; isotypes, F!, us!). Endemic. Terrestrial on roadsides and rocky clay banks, 1850-2800 m, Cajamarca, La Libertad, Huanuco, Cuzco. This resembles small specimens of E. tenuicu- lum but apparently it lacks strictly stellate tri- 156 FIELDIANA: BOTANY chomes, having round, peltate, ciliate scales that can appear as such. Cajamarca: Contumaza, alrededores de Guzmango, Sagdstegui 9044 (HUT, MO). Contumaza, Contumaza- Cascas, Sagdstegui et al. 6503 (GH). Contumaza, Bosque de Cachil (Cascas-Contumaza), Lopez et al. 9063 (GH, us). Prov. Cajamarca, La Posada, (Las Quinuas-Hua- tum), Sagdstegui 10134 (MO, uc). La Libertad: Pataz, Leon & Young 1089 in part (USM). Cuzco: Hacienda Ayasbamba, Vargas 365a (GH). 95. Elaphoglossum punae Mickel, sp. nov. Ab E. tenui stipitis squamis imbricatis, lamina latiori, laminaeque squamis ovatis basi-glandulosis segreganda. Stem long-creeping, ca. 2 mm in diameter, scales linear, lustrous, dark red-brown, ca. 3 mm long, minutely denticulate, recurved. Phyllopodia pres- ent. Leaves 1-2.5 cm apart, 10-32 cm long, 1.1- 1.6 cm broad. Petiole !/2-3/5 the leaf length, scales to 3 mm long at petiole base, distally ca. 1 mm long, longer ones linear-lanceolate and spreading, shorter ones lanceolate, appressed, brown, entire. Lamina narrowly elliptic, chartaceous, apex long- acuminate, base cuneate. Veins obscure, 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales ovate-deltate, to 1.3 mm adaxially; abaxially 0.5-0.8 mm long, scattered, with con- spicuous, glandular dots. Fertile leaves unknown. TYPE— Peru, Ayacucho, La Mar, Cordillera Central between Tambo San Miguel, Ayna & Ha- cienda Luisiana, Dudley 12031 (holotype, us!; iso- type, GH!). Endemic. Terrestrial in grassland above tim- berline on exposed summit ridges, 3400-3600 m, Ayacucho. Similar to E. tenue except: petiole scales im- bricate, lamina broader (1-1.7 cm) and longer (to 3 1 cm), abundantly glandular, scales mostly ovate to deltate, ca. 1 mm long, costal and laminar scales often touching, adaxially also glandular, the scales more lanceolate. Thus far known only from the type collection. 96. Elaphoglossum quitense (Baker) C. Chr., In- dex fil. 314. 1905. Acrostichum quitense Baker, Ann. Bot. 5: 493. 1891. TYPE: Ecuador, Andes, Sodiro (isotype, P!). Stem compact, horizontal, 3-5 mm in diameter, scales linear, lustrous, orange brown, to 13 mm long, entire. Phyllopodia lacking. Leaves fascicu- late, 1 1-21 cm long, 0.9-1.4 cm broad. Petiole '/3- '/2 the sterile leaf length, scales tan, linear, patent, some curved, lax, to 4 mm long, with some minute erect glandular trichomes. Lamina linear-ellipti- cal, chartaceous, apex acute to obtuse, apiculate, base narrowly cuneate. Veins obscure, free, ca. 1 mm apart, at ca. 60-degree angle. Hydathodes dis- tinct. Lamina scales abundant, narrow, not over- lapping laterally, 1-2 mm long, pale, tan to white, adaxial surface glabrescent or with scales longer than on abaxial surface. Fertile leaves barely short- er to somewhat longer than the sterile, petiole 2/3- % the fertile leaf length, lamina oblong, shorter than but as broad as the sterile, apex acute to obtuse, base rounded; perhaps with a few inter- sporangial scales. Epiphytic in wet forests, 2500-3265 m, Caja- marca, La Libertad, San Martin, Huanuco, Junin. Colombia; Ecuador; Peru. Cajamarca: Celendin, Gelig, Mostacero et al. 0899 (F, GH). Contumaza, Lopez et al. 3727 (F, GH, NY). Callan- jaya, Diaz & Vdsquez 634 (USM). La Libertad: W of Huamachuco, Correll & E. Smith P938 (GH). San Mar- tin: Huallaga. Valley Rio Apisoncho, Hamilton & Hol- ligan 904 (us). Huanuco: Cerro de Pasco, Ellenberg4012 (GH). Mito, Bryan 389 (F). Junin: Huancayo, Kunkel420, in part (GH). 97. Elaphoglossum raywaense (Jenm.) Alston, Bol. Soc. Brot. 2, 32: 24. 1958. Acrostichum raywaense Jenm., Ferns Brit. W. Ind. 341. 1909. TYPE: Guyana, region of Mt. Raywa, on upper parts of Isorooroo River, Jenman (ho- lotype, NY!; isotype, NY!). Acrostichum apodum var. sprucei Baker, in Mart., Fl. bras. 1(2): 578. 1870. TYPE: Brazil, Prov. Alto Amazonas, in sylvis catingas prope S. Gabriel da Cachoeira, Spruce 2186 (not located). Stem compact, horizontal, 0.3-1 cm in diame- ter, scales linear-lanceolate, yellow-brown to fer- rugineous, lustrous and sometimes resinous to- ward the tip, 5-8 mm long, undulate, entire. Phyllopodia present. Leaves fasciculate, 18-56 cm long, 2.2-6 cm broad. Petiole almost lacking, 0.5- 1.0 cm long, covered by dense subulate scales, 6 mm long, golden-orange, densely spreading on petiole and base of costa, 4-6 mm long, also with short, minute, resinous, brown, capitate glandular TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 157 trichomes on petiole and both faces of costa. Lam- ina oblanceolate, chartaceous, apex acuminate to long-caudate, base attenuate. Veins evident, free, ca. 0.8 mm apart, at 45-degree angle. Hydathodes lacking. Lamina scales on costa abundant, subu- late, orange to orange-brown, mixed with many short, minute, resinous, capitate glandular tri- chomes, laminar scales brown-orange to yellow- orange, 0.5-1.5 mm long, spreading mostly on the adaxial surface, lamina also densely covered with short, appressed, resinous, capitate glandular tri- chomes which are usually oriented toward the margin, margin densely covered by 2 or more rows of fine subulate scales, similar in size and color to those on the costa. Fertile leaves shorter than the sterile, petiole about equal to that of the sterile leaf, lamina narrower than the sterile, lanceolate to linear-lanceolate, apex acuminate, base atten- uate, petiole alate; intersporangial scales lacking. Epiphytic in wet forests, 275-1200 m, Ama- zonas, San Martin, Loreto, Huanuco, Pasco, Ju- nin, Ucayali, Madre de Dios. Guianas; Venezuela; Ecuador; Peru; Brazil. Ama/onas: Bagua, Rio Maranon, Wurdack 1858 (us), 7959 (F, GH, NY, uc). San Martin: Mariscal Caceres, Dist. Tocache Nuevo, /. Schunke V. 13751 (MO, NY). Tingo Maria, Allard 20888 (us). Loreto: Balsapuerto, Rio Hua- llaga basin, Killip & Smith 28498 (NY, us). Maynas, Al- pahuayo, van der Werffet al. 10264 (uc). Huanuco: be- tween Huanuco and Pampayacu, Kanehira 151 (GH, us). Pasco: Oxapampa, Palcazu, Rio Alto Iscozacin, Foster & d'Achille 10076 (F). Junin: Oxapampa, Shiringamazu, D. Smith & Salick 8346 (uc). Cahuapanas, on Rio Pichis, Killip & Smith 26782 (us). Ucayali (as Loreto): Aguaytia, /. Schunke V. 5495 (F, NY). Madre de Dios: Manu, Cerro de Pantiacolla, Rio Palotoa, Foster et al. 10693 (F, GH). 98. Elaphoglossum rimbachii (Sodiro) Christ, Monogr. Elaphoglossum 126./ 70. 1899. Acrostichum rimbachii Sodiro, Crypt, vase. Quit. 478. 1893. TYPE: Ecuador, Prov. Cuenca, entre Su- rrucucho y Quinoas, Rimbach (not located). Elaphoglossum pichinchae Christ, Monogr. Elapho- glossum 68. 1899. SYNTYPES: Ecuador, Pichin- cha, Sodiro (P!); Colombia, Azufral, Anden von Pasto, Columb., Andre 3224 and 5552 (not lo- cated). Stem compact, 4-5 mm in diameter, scales lin- ear-lanceolate, lustrous, dark maroon to black, 4- 6 mm long, with stiff hairlike teeth (teeth decid- uous with age). Phyllopodia lacking. Leaves ap- proximate, 25-105 cm long, 1.0-2.2 cm broad. Petiole '/j-'/j the leaf length, scales dense to scat- tered, black, ca. 1 mm long, slightly spreading, long hair-toothed. Lamina linear-elliptic, chartaceous, apex acuminate to narrowly obtuse, base narrowly cuneate. Veins obscure, hidden by scales, ca. 1 mm apart, black, sclerotic, 1-3 mm long. Fertile leaves nearly equalling the sterile in length, petiole '/3-14 the leaf length, lamina slightly narrower than the sterile; intersporangial scales present or lacking. Epiphytic, less commonly terrestrial, in wet for- ests, 2200-3500 m, San Martin, Huanuco, Pasco. Ecuador; Peru. San Martin: Mariscal Caceres, Chochos, NW corner of Rio Abiseo Nat. Park, Young & Leon 4569 (USM), Young 2190 (NY), 2247 (USM). Huanuco: Muna, Mac- bride 4301 (F, GH, NY, us). Tambo de Vaca, Bryan 651 (F, GH), 677 (F). Pasco: Oxapampa, 2-4 km N of Mallam- pampa, D. Smith & Canne 5863 (MO). 99. Elaphoglossum rosenstockii Rosenst., Repert. Spec. Nov. Regni Veg. 4: 6. 1907. TYPE: Ec- uador, Mt. Cubilin, Rimbach 8, (holotype, s!; isotype, us!). Elaphoglossum pseudohirtum Rosenst., Meded. Rijks. Herb. Leiden no. 19: 23. 1913. TYPE: Bolivia, in valle Corani, Herzog 2165a (not located). Stem long-creeping, 2-3 mm in diameter, scales linear-lanceolate, lustrous, dark brown, 2-5 mm long, entire. Phyllopodia evident. Leaves 1-3 cm apart, 24-35 cm long, 1.2-1.6 cm broad. Petiole V2-3/5 the sterile leaf length, with orange scales ap- pressed to ascending, 1.5-3 mm long, erose. Lam- ina narrowly elliptic, chartaceous, apex obtuse, base cuneate. Veins obscure, free, 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales abaxially dense, imbricate, orange to tinged with reddish, sclerotic, lanceolate to linear-lan- ceolate, erose-denticulate, 1-1.5 mm long, costal scales 2 mm long, wider than on the surfaces, erose, adaxially scattered (dense at first), ovate to lan- ceolate, erose, white, membranous. Fertile leaves about equal the sterile in length, the petiole 3/5 the fertile leaf length, the lamina about equal the ster- ile in size and shape; intersporangial scales bico- lorous, lanceolate, denticulate, 1.5 mm long. Terrestrial in patches on open, mossy, rocky places, 3400-3950 m, Cajamarca, Amazonas, An- cash, Huanuco. 158 FIELDIANA: BOTANY Cajamarca: Contumaza, Pampa de la Sal, Sagdstegui 10745 (F). Amazonas: Calla Calla Pequena, Clements 2023 (NY, USM). Ancash: Huari, Huascaran Nat. Park, D. Smith et al. 12537 (NY). Huanuco: Tambo de Vaca, Macbride 4450 (F). 100. Elaphoglossum rubellum Mickel, sp. nov. Ab E. siliquoidi statura minore et lamina apice acuta vel obtusa dispar. Stem short-creeping, 2-3 mm in diameter, scales lanceolate, brown, with sparse, irregular teeth, contorted, zigzag, ca. 4 mm long. Phyllopodia lacking. Leaves fasciculate, to 30 cm long, 2. 1-3.2 cm broad. Petiole V5-Vj the leaf length, with scales abundant, reddish to blackish, subulate, mostly 4- 8 mm long, widely spreading, often curved, lax, petiole also with erect glandular trichomes (3- celled, gland-tipped, to 0.5 mm long). Lamina nar- rowly elliptic, chartaceous, apex obtuse, base trun- cate to rounded. Veins obscure, free, 1.5-2 mm apart, at 50-60-degree angle. Hydathodes evident. Lamina scales subulate, reddish, long and con- spicuous, mostly 3-5 mm long, lax, uniformly and liberally distributed over the surface, margin and costa, lamina also with erect glandular trichomes as on petiole (3 cells, 0.2-0.5 mm long). Fertile leaves much shorter than the sterile, usually about V2 the length, lamina orbicular to elliptic, apex obtuse, base rounded, scales on petiole, adaxial lamina surface, and abaxial costa, but interspor- angial scales lacking. TYPE— Peru, Ucayali (as Loreto), Coronel Por- tillo, Boqueron del Padre Abad, Skog et al. 5127 (holotype, NY!; isotype, us!). Endemic. Epipetric on wet shady calcareous cliffs, 400-470 m, Ucayali. This closely resembles E. siliquoides of Jamaica in the very long, reddish, hairlike scales, short rounded fertile leaves, and relatively long, glan- dular trichomes, but differs in the smaller size, the acute to rounded lamina apex, and being epipetric rather than epiphytic. Ucayali: Coronel Portillo, Dist. Padre Abad, J. Schunke V. 3064 (F, us). 101. Elaphoglossum ruficomus Mickel, sp. nov. Rhizomatis squamis linearibus ramosis purpureis, lamina laxe squamosa et stipite hirto recognita. Stem compact, horizontal, ca. 10 mm in di- ameter, scales linear, long-attenuate, lustrous, ma- roon, 12-20 mm long, entire. Phyllopodia indis- tinct. Leaves fasciculate, (30-)52-66 cm long, 2.5- 3.6 cm broad. Petiole about V2 the sterile leaf length, with orange scales widely spreading, 8-12 mm long, ciliate, others 2-3 mm long, appressed. Lamina narrowly elliptic, chartaceous to coriaceous, apex acute, base broadly cuneate to rounded. Veins ev- ident, ca. 1 mm apart, at ca. 60-degree angle. Hy- dathodes lacking. Lamina scales abaxially 1-3 mm long, orange, lanceolate to ovate, imbricate to scat- tered, costa scales 3-4 mm long, linear-lanceolate, spreading, adaxially scattered, orange, 1-2 mm long, ciliate, lanceolate. Fertile leaves shorter than the sterile, petiole '/2 the fertile leaf length, lamina narrower than the sterile (2.0 cm broad); inter- sporangial scales lacking or present. TYPE— Peru, Cajamarca, Celendin, Sendamal (Ruta a Celendin), Sagdstegui 12091 (holotype, NY!). Endemic. Epiphytic in wet forest or epipetric on brushy slopes, 3050-4200 m, Cajamarca, San Martin, Ancash. San Martin: Mariscal Caceres, Puenta del Monte, Young 1740 (USM). Ancash: Yungay, Quebrada Llanga- nuco, Perez 84 (USM). Yungay, Huascaran Nat. Park, D. Smith et al. 9174 (NY). 102. Elaphoglossum rufum Mickel, Brittonia 37: 277. 1985. TYPE: Peru, Dept. San Martin, Lamas, along Rio Curiyacu, 8 km above San Antonio, Belshaw 3601 (holotype, us!; photo, NY!; isotypes, GH!, uc!). Stem short-creeping, 3-4 mm in diameter, scales linear-lanceolate, ferrugineous, 4-6 mm long, en- tire, tortuous toward tip. Phyllopodia lacking. Leaves 31-50 cm long, 4.5-7 cm broad. Petiole ca. i/5_'/3 the sterile leaf length, with scales abun- dant, spreading, subulate, somewhat denticulate, rufous, 2-3 mm long. Lamina lanceolate, charta- ceous, generally with a proliferous bud in the re- tuse apex, otherwise acute to acuminate, base at- tenuate. Veins distinct to indistinct, free, 1 mm apart, at 60-70 degree angle. Hydathodes evident. Lamina scales sparse to moderately abundant on both sides, spreading, lanceolate to subulate, 2-3 mm long, denticulate, rufous, denser along costa and margin. Fertile leaves about same length as sterile, petiole ca. 5/8-% the fertile leaf length, lam- TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 159 ina lanceolate to narrowly lanceolate, apex obtuse, base cuneate; intersporangial scales sparse to ab- sent, lanceolate, entire to denticulate, spreading, denser along costa and margin, 1-2 mm long. Endemic. Epiphytic in wet forests, 425-2250 m, Cajamarca, San Martin. Cajamarca: Prov. Cutervo, San Andres, Lopez & Sa- gdstegui 5428 (GH). 103. Elaphoglossum russelliae Mickel, sp. nov. Ab E. cardenasii non nisi lamina integerrima stipi- teque breviore diversa. Named for Ruth Russell, who has been a vol- unteer in the New York Botanical Garden fern herbarium for 17 years. Stem short-creeping, ca. 4 mm in diameter, scales flat, ovate-acuminate, 2-3 mm long, appressed, dark brown, sparse. Phyllopodia lacking. Leaves fasciculate, to 55 cm long, 2.2-3.0 cm broad. Pet- iole about V3 the sterile leaf length, sparsely clothed with tan, ovate scales 1-3 mm long, mostly ap- pressed. Lamina linear-elliptic, thin, apex acu- minate, base narrowly to broadly cuneate, margin crenulate. Veins evident, free, 1-2 mm apart, at 60-70-degree angle. Hydathodes evident. Lamina scales abaxially on costa tan, lanceolate, ca. 1 mm long, occasional on margin between crenulations, laminar surfaces with minute stellate trichomidia. Fertile leaves shorter than sterile, petiole about % the fertile leaf length, lamina smooth-margined, lanceolate, apex acuminate, base truncate to sub- cordate; intersporangial scales lacking. TYPE— Peru, Cajamarca, Cutervo, Parque Na- tional de Cutervo, Diaz & Osores 2587 (holotype, NY!). Endemic. Epiphytic in dry woods, ca. 2150m, Cajamarca, Huanuco. This is probably the precursor of the bizarrely divided E. cardenasii, differing only in the dissec- tion and shorter petiole. Huanuco: Mima, Bryan 531 (F), 533 (F). 1 04. Elaphoglossum setigerum (Sodiro) Diels, Nat. Pflanzenfam. 1(4): 333. 1899. Acrostichum setigerum Sodiro, Anales Univ. Quito 4: 174. 1890. TYPE: Ecuador, Cordillera occidental near Nono, Sodiro (holotype, not located). Stem compact, horizontal, to 6 mm in diameter, scales linear, reddish orange, to 10 mm long, en- tire. Phyllopodia lacking. Leaves fasciculate, 18- 28 cm long, 1.7-2.4(-3.6) cm broad. Petiole V4-2/5 the sterile leaf length, with scales reddish-orange, spreading, subulate, entire, 2-3 mm long. Lamina narrowly elliptic, chartaceous, apex acuminate to cuspidate, base rounded. Veins distinct, free, 1.5- 2.5 mm apart, at 60-70-degree angle. Hydathodes evident. Lamina scales scattered, reddish-orange, subulate, slightly dentate, denser at margin, spreading, ca. 2 mm long on costa and margin, 1 mm long on surface, fewer scales on the adaxial surface. Fertile leaves slightly shorter than the ster- ile, petiole 2/3-% the fertile leaf length, lamina lan- ceolate, apex acute, base truncate to cordate; in- tersporangial scales lacking. Epiphytic(?) in wet forests, 3100 m, Amazonas. Ecuador; Peru. This species closely resembles E. crinipes C. Chr. of Mesoamerica and Hispaniola, and may prove to be the same. Amazonas: Prov. Chachapoyas, Cerros Calla Calla, Hutchison & Wright 5797, in part (GH), 5795 (uc). 105. Elaphoglossum simulans Mickel, sp. nov. Ab affini E. eximio stipite longiori et lamina latiori base truncata diversa. Stem creeping, ca. 5 mm in diameter, stem scales flat, appressed, ovate-lanceolate, brown, entire, 2- 3 mm long. Phyllopodia lacking. Leaves 5-10 mm apart, 30—40 cm long, 1.5-2.2 cm broad. Petiole about !/2 the sterile leaf length, with scales scat- tered, light brown, tightly appressed at petiole base, distally patent, 1-3 mm long, orange-tan, dense, linear-lanceolate, some subulate. Lamina linear or pedately divided into three arms, thin-textured, apex acuminate, base broadly cuneate to truncate, margin crenulate. Veins evident, 1.5-2 mm apart, at ca. 70-degree angle. Hydathodes evident. Lam- ina scales on costa scattered, orange-tan, subulate, 2-3 mm long, spreading; adaxially scales reduced to sparse squamules 0.3-0.5 mm long, abaxially to stellate trichomidia. Fertile leaves unknown. 160 FIELDIANA: BOTANY TYPE— Peru, Cuzco, Urubamba, Machu Picchu, Peyton & Peyton 1316 (holotype, GH!). Endemic. Terrestrial in wet forests, 2560-2810 m, Cuzco. This is like E. eximium but in E. simulans the stem is long-creeping, the petiole longer, the lam- ina broader and more truncate at base, there are stellate trichomidia on the lamina, costal scales are spreading, the margin is crenulate, and veins are 2-2.5 mm apart. I have not seen the one specimen of E. carden- asii reported from Peru (Gomez, 1972). There should be no question of its determination, it being so distinct in its pedate form, but the discovery of the same form of dissection in E. simulans lends some question to the record of E. cardenasii. Cuzco: Urubamba, Machu Picchu, Peyton & Peyton 1316b (GH). group with long-creeping stems and broad, pale stem and petiole scales. It is very closely related to Peltapteris, which justifiably could be placed in Elaphoglossum, differing from E. squamipes et off. only in leaf dissection. Amazonas: Chachapoyas, upper slopes of Puma-urcu ESE of Chachapoyas, Wurdack 692 (F, NY, uc, us). Cha- chapoyas, Cerros Calla Calla, 1 0 km above Leimebamba on road to Balsas, Hutchison & Bennett 4753 (F, GH, NY, uc). Chachapoyas, Cerros Calla Calla, 45 km above Bal- sas midway on road to Leimebamba, Hutchison & Wright 5825 (F, GH, NY, uc, us). San Martin: Mariscal Caceres, Rio Abiseo Nat. Park, above timberline, Puerta del Monte, Young 1997 (NY). Mariscal Caceres, Parque Nac. Rio Abiseo, valle Pampa de Cuy Leon & Young 1280 (USM). Huanuco: Mito, Macbride 3359 (F, us). Muna, Bryan 526 (F, us), Macbride 4024 (F, us). Pasco: Oxa- pampa, Oxapampa- Villa Rica Road, 7 km from road- head, Smith & Alban 5571 (NY). Oxapampa, San Alberto, van der Werff8459 (uc). 106. Elaphoglossum squamipes (Hooker) Moore, Index fil. 15. 1857. Figure 25f-g. Acrostichum squamipes Hooker, Icon. pi. 1. 197. 1837. TYPE: Peru. Chachapoyas, Mathews (holotype, K!). Stem long-creeping, ca. 1 mm in diameter, scales ovate to linear-lanceolate, dull orange-tan, entire, 3-6 mm long. Phyllopodia lacking. Leaves 3-20 mm distant, 5-14 cm long, 1 . 1-2.2 cm broad. Pet- iole slender, V2-2/3 the sterile leaf length, moderately clothed with ovate, dull orange scales, these 2-3 mm long. Lamina ovate to lanceolate, subcoria- ceous, apex obtuse, base broadly cuneate to round- ed. Veins obscure, free, ca. 1 mm apart, at 60-70- degree angle. Hydathodes lacking. Lamina scales sparse, small, inconspicuous, 1-2 mm long, linear, often coarsely and irregularly dentate, especially toward the scale base, more frequent on the ab- axial surface. Fertile leaves equal to or slightly longer than the sterile leaves, petiole ca. % the fertile leaf length, lamina ovate to lanceolate, smaller than the sterile with a few dark brown to black scales along the abaxial costa and among the sporangia. Epiphytic and epipetric in wet forests, 2100- 3300 m, Amazonas, San Martin, Huanuco, Pasco. Mexico to Panama; Venezuela and Colombia to Bolivia. Elaphoglossum squamipes represents a distinct 107. Elaphoglossum stenophyllum (Sodiro) Diels, Nat. Pfanzenfam. 1(4): 333. 1899. Acrostichum stenophyllum Sodiro, Crypt, vase. Quit. 468. 1893. TYPE: Ecuador, Sancullan, Sodiro, Dec. 1889 (isotype, us!). Similar to E. tectum in its round, white, peltate scales on the adaxial surface and round, ciliate scales on the petiole, but the abaxial surface in E. stenophyllum lacks stellate trichomes, having in- stead glandular dots below and scattered linear- lanceolate scales on the costa. Terrestrial on shrubby slopes, 2750-3100 m, Amazonas, Huanuco. Ecuador; Peru. Amazonas: Chachapoyas, Cerros Calla Calla, Hutch- ison & Wright 5805 (uc). 108. Elaphoglossum styriacum Mickel, Brittonia 39: 326. 1987. TYPE: Venezuela, Bolivar, Chimanta Massif, Steyermark 75347 (holo- type, MO!). Stem compact, horizontal, ca. 3-4 mm in di- ameter, scales linear-lanceolate, orange, 2-3 mm long, entire or with occasional processes. Phyllo- podia present. Leaves fasciculate, 19-37 cm long, 2.5-4.9 cm broad. Petiole V10-V6 the sterile leaf length, glabrous. Lamina oblanceolate to elliptic, coriaceous, margin distinctly thickened, apex acu- TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 161 minate to obtuse, subcuspidate, base narrowly cu- neate-decurrent. Veins inconspicuous, free, 1 mm apart, at 5 5-6 5 -degree angle. Hydathodes lacking. Lamina scales essentially lacking abaxially, lamina often lustrous, with scattered, dark, stellate tricho- midia, adaxially glabrous. Fertile leaves equal to or slightly shorter than the sterile, petiole 2/5-V2 the fertile leaf length, the lamina narrower than the sterile, 13-16 mm broad; intersporangial scales lacking. Epiphytic and terrestrial in wet forests, 100-770 m, Amazonas, San Martin, Loreto, Pasco. Venezuela; Peru. This species is rather variable in its lamina shape: elliptic to oblanceolate, with apex obtuse to acu- minate. The abaxial surface is usually lustrous and has scattered to sparse stellate trichomidia. The stem scales persist onto the phyllopodia, and are linear, orange, with irregular processes. Amazonas: Bagua, along roadside from Chiriaco to Puente Venezuela, 43 km NE of Chiriaco, Barbour 4464 (MO). San Martin: Mariscal Caceres, Granja Santa Ines, 4 km arriba de Tocache Nuevo, /. Schunke V.3651 (F, uc). Road between Moyobamba & Chachapoyas, E of Naranjos at Rio Naranjos, Croat 58155 (F). Loreto: Mi- shuyacu, near Iquitos, Klug 1461 (NY, us). Pasco: Oxa- pampa, Valle del Palcazu, Rio Palcazu, cerca de Iscoza- cin, Leon 7 18 (F). 109. Elaphoglossum tambillense (Hooker) Moore, Index fil. 15. 1857. Acrostichum tambillense Hooker, Icon. pi. t. 656. 1 844. TYPE: Ecuador, Tambillo, Jameson (holotype, K!; isotypes, B!, P!). Stem compact, horizontal, to 10 mm in diam- eter, scales linear-lanceolate, reddish brown, to 7 mm long, entire. Phyllopodia lacking. Leaves fas- ciculate, 7-14 cm long, 1.4-2.3 cm broad. Petiole 2/5-3/5 the sterile leaf length, with scales lacking, but with minute gland-tipped trichomes present. Lam- ina ovate-lanceolate, chartaceous, apex cuspidate to acuminate, base truncate to rounded. Veins ob- scure, free, ca. 1 mm apart, at 70-80-degree angle. Hydathodes evident. Lamina scales lacking but gland-tipped trichomes present, especially at base near costa and near margin, mostly on abaxial surface. Fertile leaves shorter than the sterile, pet- iole 3/5 the fertile leaf length, lamina similar to sterile in shape but smaller, apex acute; inter- sporangial scales lacking. Grassy cliffs, upland gorge, 2000-2800 m, Hua- nuco, Cuzco. Ecuador; Peru. Huanuco: Mito, Macbride 3373 (F, us). Cuzco: Toron- toy, Herrera 1326 (us). 110. Elaphoglossum tectum (Willd.) Moore, In- dex fil. 15. 1857. Acrostichum tectum Willd., Sp. pi. ed. 4, 5: 102. 1810. TYPE: Venezuela, Caripe, Humboldt (holotype, B!, Herb. Willd. 195201; isotypes, LE!, P!). Acrostichum elongatum Kunze, Linnaea 9: 31. 1834. TYPE: Peru, ad Pampayacu, 1829, Herb. Kunze. (holotype, LZ, destroyed; isotype, B?) Elaphoglossum elongatum (Kunze) Moore, Index fil. 9. 1857. Stem short- to moderately creeping, 2-4 mm in diameter, scales linear-lanceolate, dark reddish- black, sclerotic, lustrous, 3-5 mm long, with tor- tuous tip, entire, or with sparse pale weak hairlike teeth. Phyllopodia indistinct, usually hidden by scales. Leaves fasciculate to slightly spaced, to 3 mm apart, 15-44 cm long, 1.4-2.8 cm broad. Pet- iole V3-V2 the sterile leaf length, with scales ap- pressed, peltate, round with dark center and short hairlike teeth, often mixed with scattered dark scales like those of stem. Lamina linear-elliptic, apex long-acuminate, base narrowly cuneate. Veins obscure, free, mostly 0.7-1 mm apart, at ca. 80- degree angle. Hydathodes lacking. Lamina scales on adaxial surface white, appressed, round, pel- tate, dentate to ciliate, becoming glabrous with age, on abaxial surface reduced to stellate tri- chomes. Fertile leaves about as long as the sterile or slightly longer, petiole usually !/2-2/3 the fertile leaf length, lamina narrower than the sterile; in- tersporangial scales lacking. Terrestrial on rocks and walls, in woods and fields, 1800-2900 m, Cajamarca and Amazonas south to Ayacucho and Cuzco. Mexico to Panama; West Indies; Venezuela and Colombia to Bolivia and Brazil. This species is readily distinguished by the round, peltate scales on the petiole and adaxial lamina surface and stellate trichomes on the abaxial sur- face. Cajamarca: Celendin, Gelig, Mostacero 866 (HUT, MO, uc). Amazonas: Chachapoyas, Cano Santa Lucia, Wur- dack 735 (F, GH, NY, uc, us). Cerros Calla Calla, Hutch- 162 FIELDIANA: BOTANY ison & Wright 48 18 A (us). San Martin: Tingo Maria, Allard 22205 (us). Huanuco: Prov. Huanuco, Chinchao, entre Huanuco y Tingo Maria, Ferreyra 16941 (GH). Mito, Macbride & Featherstone 1391 (us). Junin: La Merced Chanchamayo, Soukup 1099 (F). Prov. Tarma, between Palea & Carpapata, Stork 10959 (K). Huacap- istana, Killip & Smith 24177, 24319 (us). Ayacucho: Ccarrapa between Huanta & Rio Apurimac, Killip & Smith 22424 (GH). Cuzco: Machu Picchu, road to ruins. Leon 460 (GH). Machu Picchu to San Miguel, Urubamba, Vargas 2066 (us). 111. Elaphoglossum tenue Mickel, sp. nov. Rhizomate tenuissimo squamis recurvis induto nec- non lamina parva gracili infra laxe squamata notabilis. Stem long creeping, ca. 1-2 mm in diameter, scales linear, lustrous, dark brown, ca. 5 mm long, recurved, with sparse small teeth. Phyllopodia ev- ident, ca. 5 mm long. Leaves 1-2 cm apart, 8-16 cm long, 0.4-0.6 cm broad. Petiole about '/2 the sterile leaf length, with orange scales scattered, ap- pressed to spreading, 1 mm long, with small teeth. Lamina linear-elliptic, chartaceous, apex acumi- nate, base narrowly cuneate. Veins evident, free, 1 mm apart, at 60-70-degree angle. Hydathodes lacking. Lamina scales abaxially linear-lanceolate, 2 mm long, orange, cilio-denticulate, often con- torted, with glandular base, adaxially scattered, lanceolate, dentate near scale base, mostly on cos- ta. Fertile leaves not known. TYPE— Peru, Cuzco, La Convention, Cordillera Vilcabamba. Dudley 11192 (holotype, GH). Endemic. Epiphytic in wet forests, 3330-3410 m, Cuzco. Thus far known only from the type. 1 12. Elaphoglossum tenuiculum (Fee) Baker, Ann. Bot. (London) 5: 491. 1891. Acrostichum tenuiculum Fee, Mem. foug. 10: 6, /. 29, f. 2. 1865. TYPE: Venezuela, Tovar, Fendler272 (holotype, P!). Stem short-creeping, ca. 2 mm in diameter, scales linear-lanceolate, lustrous, castaneous to brown, ca. 2 mm long, entire. Phyllopodia distinct. Leaves 1-2 mm apart, 9-13 (30) cm long, 0.4-2.0 cm broad. Petiole l/3-V2 the sterile leaf length, with sparse stellate trichomes and glandular dots, oc- casionally with sparse scales near base. Lamina linear, chartaceous, apex acuminate, base cuneate. Veins barely visible, free, ca. 1 mm apart, at 60- 70-degree angle. Hydathodes lacking. Lamina scales lacking, stellate trichomes or resinous dots on abaxial surface, those on the adaxial surface with slight scale body. Fertile leaves shorter than the sterile, petiole about '/2 the fertile leaf length, lamina narrower than the sterile, 5 mm broad; intersporangial scales lacking. Terrestrial on wet banks and cliffs, 1 500-3650 m, Amazonas, La Libertad, Lima, Junin, Cuzco. Costa Rica; Panama; Venezuela; Colombia to Bolivia and Brazil. There is considerable variation in the degree of pubescent vs. glandular dots. Amazonas: Chachapoyas, Quebrada Molina 5 km be- low Chachapoyas, Wurdack 653 (F, GH, uc, us). La Li- bertad: 3 km W of Huamachuco, Correll & Smith P936 (GH). Lima: Canta, 4 km up road Huamantanga from Lima-Canta road, Sounders 1247 (GH). Junin: Carpa- pata, 27 km from Tarma, Gentry et al. 39772 (F). Car- papata, above Huacapistana, Killip & Smith 244 10 (us). Cuzco: Paucartambo, Manu Nat. Park, Skog & Skog 5197 (us). Paucartambo, Paso del Aguila, Pillawata, Var- gas C. 22996 (GH). Rio Marcapato, 60 km above Quin- cemil, Madison 1010 pp (GH). Hacienda Huy-Huy, Valle de "Graves" Bties 1823 (us). 113. Elaphoglossum tomentellum Mickel, sp. nov. Lamina parva gracili squamis multo dissectis auran- tiacis induta, necnon costae squamis lineari-lanceolatis ab affinibus diversa. Stem compact, horizontal, ca. 2 mm in diam- eter, scales linear, lustrous, castaneous, ca. 5 mm long, subentire, glandular. Phyllopodia evident. Leaves fasciculate, 9-16 cm long, 0.4-0.7 cm broad. Petiole !/2 the sterile leaf length, scales orange, ap- pressed to spreading, ca. 2 mm long, subentire or irregularly sparsely dentate. Lamina narrowly el- liptic, chartaceous, apex acute to acuminate, base narrowly cuneate. Veins obscure, 1 mm apart, at ca. 70-degree angle. Hydathodes lacking. Lamina scales abaxially orange, much reduced to linear or ovate with cilia longer than the scale body, gland- based, costal scales linear, reddish, dentate, ca. 2 mm long; adaxial scales round to lanceolate, white, ciliate at base, ca. 1 mm long, or orange and dense when young, 1-2 mm long. Fertile leaves not known. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 163 TYPE— Peru, Cajamarca, Contumaza, Cerro Campanulas, Guzmango, Sagdstegui 2993 (ho- lotype, GH!). Rocky slopes, 3050 m, Cajamarca. Thus far known only from the type. 1 14. Elaphoglossum velongum Mickel, sp. nov. Ab E. latifolio lamina linear! et stipitis squamis au- rantiacis sparsis abstat. Stem compact, horizontal, 4-7 mm in diameter, scales linear-lanceolate, lustrous, orange, 7-10 mm long, with hairlike processes. Phyllopodia present. Leaves fasciculate, 27-30 cm long, 1.8-2.1 cm broad. Petiole V4 the sterile leaf length, at base with orange scales as on stem, reduced distally nearly to stellate trichomes, scales appressed to spread- ing. Lamina linear, subcoriaceous, apex acumi- nate, base narrowly cuneate. Veins obscure, free, 1 mm apart, at 70-80-degree angle. Hydathodes lacking. Lamina scales sparse, orange or white, highly dissected, almost tuft-like (these nearly re- duced to stellate trichomes with long lax arms). Fertile leaves longer than the sterile, petiole V2 the leaf length, lamina similar to the sterile in size and shape; intersporangial scales lacking. TYPE— Peru, Amazonas, Leimebamba, Woyt- kowski 7842 (holotype, GH!). Endemic. Terrestrial in wet forests, 2400-2800 m, Cajamarca, Amazonas. Cajamarca: Celendin, La Ranca-Gelig, Sagdstegui 12133 (NY). Amazonas: Cordillera de Calla Calla; Balsas- Leimebamba road, Duncan 2607 (us). Prov. Chacha- poyas, Quebrada Molino, Wurdack 655 (us), 656 (us). 115. Elaphoglossum vittarioides Mickel, sp. nov. Ab E. eatoniano Rhizomatis squamis ciliatis, costae squamis brevioribus, necnon lamina adaxialiter squa- mulosa recedens. Stem compact, 2-3 mm in diameter, scales lan- ceolate-deltate, lustrous, black, very indurated, 2- 3 mm long, with lax, tan, marginal trichomes. Phyllopodia present but very short (5-8 mm). Leaves approximate, 21-35 cm long, 4-6 mm broad. Petiole '/,0-% the sterile leaf length, scales scattered, appressed, black, with white teeth, to 2 mm long at petiole base, distally smaller and with pale scale margin. Lamina linear, coriaceous, apex long-acuminate, base attenuate. Veins obscure. Hydathodes lacking. Lamina scales adaxially scat- tered, orange-tan, lanceolate, 1-1.5 mm long, sub- entire to very short-ciliate, some of those on costa with dark center, slightly imbricate to glabrescent, abaxially tightly appressed, orange or white with orange center, deltate, subentire to short-ciliate, to 1 mm long, costal scales linear-lanceolate, black, 1-1.2 mm long, distally with broader white mar- gin, subentire to short-ciliate. Fertile leaves un- known. TYPE— Peru, Cuzco, Paucartambo, Sta. Isabel, Vargas 1306 (holotype, uc!). Endemic. Epiphytic in wet forests, 1 1 50 m, Cuz- co. This forms a complex with E. eatonianum and E. chloodes, which see for further discussion. Thus far known only from the type. 116. Elaphoglossum vulcanicum Christ, Monogr. Elaphoglossum 131. 1899. AcrostichumfurfuraceumBaker,].Bot. 15: 166. 1877. not Kuhn, 1869. TYPE: Ecuador, Andes of Quito, Sodiro (holotype, K). Stem widely creeping, ca. 2-3 mm in diameter, scales linear, lustrous, dark red-brown, 2-6 mm long, entire to sparsely denticulate. Phyllopodia distinct, ca. 2 cm long. Leaves ca. 1 cm apart, 10- 27 cm long, 1.5-2.6 cm broad. Petiole V3-V2 the sterile leaf length, scales black, sclerotic, appressed to spreading, ovate to lanceolate, 1-3 mm long, dentate. Lamina narrowly oblong, subcoriaceous, apex obtuse, base cuneate. Veins evident, free, 1 mm apart, at 60-70-degree angle. Hydathodes lacking. Lamina scales of costa black, ciliate, 1 mm long, deltate to lanceolate, scales on adaxial and abaxial surfaces scattered, orange (abaxially) or white (adaxially), margin with concentrated scales often black or flecked with black. Fertile leaves longer than the sterile, petiole % the fertile leaf length, lamina linear, 3-6 mm broad, inrolled; intersporangial scales lacking. Epiphytic in wet forests, 2450-3600 m, San Martin, Huanuco, Cuzco. Ecuador; Peru; Bolivia. Peyton & Peyton 945 has petiole scales more spreading than usual. 164 FIELDIANA: BOTANY San Martin: Mariscal Caceres, forest patch isolated above timberline, Chochos NW corner of Rio Abiseo Nat. Park, Young & Leon 4651 (USM). Huanuco: Mima, trail to Tambo de Vaca, Macbride 4304 (F). Cushi, trail to Tambo de Vaca, Bryan 623 (F). Cuzco: La Conven- tion, Huayopata, 8 km from Incatambo, Peyton & Pey- ton 945 (GH). Paucartambo, cordillera de 3 cruces, Var- gas C. 12241 (GH). Rio Calzuda, Huadquina, Biies 1266 (us). 1 17. Elaphoglossum wardiae Mickel, Brittonia 37: 277. 1985. TYPE: Bolivia, Dpto. Cochabam- ba, Prov. de Chapare, road to San Onofre, M. S. Foster 79-179 (holotype, uc!). Stem short-creeping, 4 mm in diameter, scales dark brown, lustrous, narrowly deltate, crispate, irregularly serrate, 2-3 mm long. Phyllopodia lack- ing. Leaves to 1 cm apart, 31-36 cm long, 4.0-5.7 cm broad. Petiole V4-V3 the sterile leaf length, with scales abundant, appressed to spreading, lanceo- late, peltate, tan, 2-3 mm long, minutely serrulate. Lamina lanceolate, chartaceous, apex notched with bud, base broadly cuneate. Veins distinct, free, 2- 3 mm apart, at ca. 70-degree angle. Hydathodes evident. Lamina scales absent to sparse on both sides, peltate, deltate to lanceolate, somewhat ap- pressed, tan, 1-2 mm long, minutely serrulate, sometimes more dense along margin. Fertile leaves nearly equal to the sterile in length, petiole %-V5 the fertile leaf length, lamina lanceolate, apex apic- ulate, base broadly cuneate; intersporangial scales abundant, peltate, ovate to lanceolate, finely ser- rulate, tan, 1-2 mm long, somewhat appressed on costa. Epiphytic in wet forest, 700-1700 m, Huanuco, Madre de Dios. Peru; Bolivia. Huanuco: La Divisoria, NE of Tingo Maria on road to Pucallpa, Moran 3703, 3705 (MO). Madre de Dios: Prov. Manu, Cerro de Pantiacolla, Foster et al. 10890 (GH). 118. Elaphoglossum williamsiorum, Mickel, sp. nov. Rhizomate longe repenti et frondibus elongatis tricho- midiis stellatis minutis sparsis (minutim stellato-punc- tatis) notanda. Named for Ethelyn and Victor Williams, who have long been supporters of the New York Bo- tanical Garden fern program. Ethelyn has been the primary fern propagator, by spores, for the Garden for the past 1 6 years. Stem long-creeping, 4-6 mm in diameter, scales lanceolate, lustrous, brown to black, ca. 3 mm long, entire. Phyllopodia present. Leaves ca. 3 cm apart, 75 cm long, 5.0-5.7 cm broad. Petiole 2/5 the sterile leaf length, glabrous. Lamina narrowly elliptic, chartaceous, apex lacking, base attenuate. Veins obscure, free, 1 mm apart, at 70-80-degree angle. Hydathodes lacking. Lamina scales lacking adaxially, lamina abaxially with black stellate- punctate indument. Fertile leaves not seen. TYPE— Peru, Cuzco, Urubamba, Machu Picchu, 0.5 km N of union of Sayacmarca & Aobamba Rivers, Peyton & Peyton 1491 (GH). Endemic. Terrestrial in wet forests, 2390 m, Amazonas, Cuzco. This superficially resembles E. amplum, which see. Amazonas: Bongara, above Pedro Ruiz on road to Pomacochas, Knapp & Alcorn 7550 (uc). Prov. Chacha- poyas, slopes of Puma-arcu SE of Chachapoyas, Wur- dack 554 (us). 119. Elaphoglossum zebrinum Mickel, sp. nov. Ab E. oblanceolato statura minori laminisque angustis diversa. Stem short-creeping, 2-4 mm in diameter, scales mostly lacking, some small ones 1-2 mm long near apex, black, lustrous, linear-lanceolate, entire. Phyllopodia lacking. Leaves fasciculate, 32-47 cm long, 2.3-3.4 cm broad. Petiole nearly lacking, to Vio the sterile leaf length, glabrous but with white blotches on petiole and lamina (mostly between the veins). Lamina linear-oblanceolate, charta- ceous, costa strongly keeled, apex acuminate, base long-attenuate. Veins obscure, free, 1.5-2 mm apart, at ca. 70-degree angle. Hydathodes distinct. Lamina scales sparse on both surfaces, subulate, brown, generally less than 1 mm long. Fertile leaves slightly shorter than the sterile, petiole about V2 the fertile leaf length; intersporangial scales lack- ing. TYPE— Peru, Amazonas, Bagua, Valley Rio Ma- rafion, Wurdack 1859 (holotype, us!). Epiphytic in wet forests, 295-1000 m, Lamba- yeque, Amazonas, Madre de Dios. Colombia; Peru. TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 165 Lambayeque: Puerto Nazareth, near Olmos, Ellenberg 1427 (GH). Madre de Dios: Manu, Cerro de Pantiacolla, *io Palotoa, NNW of Shintuya, R. Foster 10639 (F), 10891 (GH). Comments At the time the manuscript was being delivered lo the printer, two more new species were discov- ered. Although it was impractical at this point to revise the key and order of species, they are in- cluded here as addenda. Elaphoglossum pattersoniae Mickel, Amer. Fern J. 80: 110. 1990. TYPE: Peru, Pasco, Prov. Oxapampa, Oxapampa- Villa Rica road, D. Smith & Alban 5590 (holotype, NY!; isotype, MO!). Stem compact, horizontal with ascending apex, scales linear, lustrous, dark red-brown, ca. 4 mm long, denticulate, ascending. Phyllopodia present. Leaves fasciculate, 6-1 6 cm long, 1 .2-1 .8 cm broad. Petiole !/3 the sterile leaf length, scales 1-2 mm long, linear-lanceolate, ascending to spreading, dark red-brown, lustrous, cilio-denticulate. Lamina el- liptic, chartaceous, apex obtuse to broadly acute, base cuneate. Veins at 55-60-degree angle. Hy- dathodes lacking. Lamina surface adaxially with scattered orange-tan, stellate to lanceolate cilio- denticulate scales, to 0.5 mm long, costal scales more abundant and lustrous red-brown, those on abaxial surface more sparse, smaller, and costal scales mostly orange-tan. Fertile leaves longer than the sterile, petiole 2/3 the leaf length, similar to sterile but narrower, ca. 8 mm broad; scales of petiole and adaxial lamina surface mostly sub- stellate, lustrous, dark red-brown, appressed; in- tersporangial scales lacking. Terrestrial in high montane rain forest, 2120 m. Elaphoglossum pattersoniae is closest in Peru to E. guamanianum, but differs from that in its smaller size, acute-obtuse lamina apex, lack of dark-arachnoid scales on the costa abaxially, and fewer laminar scales. Known only from the type. Elaphoglossum potomogeton Mickel, Amer. Fern J. 80: 112. 1990. TYPE: Peru, Junin, Prov. Chanchamayo, Chilpez, ca. 26 km S of San Ramon, D. Smith & Palacios 2653 (holotype, NY!; isotype, MO). Stem long-creeping, 1-1.5 mm in diameter, scales linear, lustrous, dark red-brown, ca. 3 mm long, entire, slightly recurved. Phyllopodia pres- ent, covered by scales. Leaves 0.5-1.5 cm apart, 13-17 cm long, 1.7-2.2 cm broad. Petiole 2/5-'/2 the sterile leaf length, scales 2-3 mm long, linear- lanceolate, spreading and recurved to appressed distally on the petiole, tan to sclerotic, red-brown, entire. Lamina elliptic, chartaceous, apex obtuse, base broadly cuneate. Veins obscure, ca. 1 mm apart, at 65-degree angle. Hydathodes lacking. Lamina scales ovate to ovate-deltate, fimbriate- denticulate, ca. 0.5 mm long, sparse abaxially, abaxially and adaxially concentrated at the mar- gin, to 1 mm long, lamina surfaces with scattered glandular dots. Fertile leaves longer than the ster- ile, petiole ca. % the frond length, similar to the sterile in apex and base but narrower, ca. 1 cm broad; intersporangial scales lacking. Epiphytic, primary high montane forest domi- nated by Podocarpus, Cedrela, and Juglans neo- tropica, 1720-1850 m. Elaphoglossum potomogeton resembles E. pu- nae and E. longius in the slender, long-creeping stem with red-brown recurved scales and the lam- ina surface with glandular dots. It is distinct from those species in its obtuse lamina apex and laminar scales concentrated at the lamina margin. Known only from the type. The following names, based on Peru material collected by Poeppig, undoubtedly pertain to spe- cies of Elaphoglossum. However, they cannot be placed without reference to the original specimens, which have not been located. Acrostichum adenolepis Kunze, Linnaea 9: 27. 1834. TYPE: Peru, Pampayacu, Poeppig, Jul. 1829 (holotype, not located). Acrostichum calophyllum Kunze, Linnaea 9: 27. 1834. TYPE: Peru, Pampayacu, Poeppig (ho- lotype, not located). Acrostichum curvans Kunze, Linnaea 9: 30. 1834. TYPE: Peru, Pampayacu, Poeppig diar. 1115, Jul. 1829 (holotype, not located). Acrostichum dissimile Kunze, Linnaea 9: 28. 1834. TYPE: Peru, Cassapi, Poeppig, Jul. 1829 (ho- lotype, not located). 166 FIELDIANA: BOTANY XXVI. Peltapteris Contributed by John T. Mickel. Peltapteris Link, Fil. spec. 147. 1841. TYPE: Acrostichum peltatum (Sw.) Sw. = Peltapteris peltata (Sw.) Morton. Figure 26. Epiphytic. Stem long-creeping, slender, scaly. Leaves small, dimorphic. Petiole scaly. Lamina flabellate to pinnate, undivided to 4 times divided; if divided, segments linear, glabrous, chartaceous. Fertile lamina round, cordate or two-lobed, spo- rangia covering abaxial surface; spores bilateral. Five species in tropical America, one wide- spread, others limited. We are of two minds regarding the recognition of Peltapteris as a genus distinct from Elapho- glossum. On the one hand, its leaf architecture is strongly dissected and is easily recognized but tends to be lost in the large number of species in Ela- phoglossum. On the other hand, its anatomy, ii dument, chemistry, and spores are nearly identic to those of E. squamipes, and it is clear that lei architecture is the only character separating th two taxa. Rhipidopteris Fee, Mem. foug. 2: 14. 1845, nom. illeg. References GOMEZ P., L. D. 1975. Contribuciones a la pt ridologia costarricense. VI. El genero Peltaptet Link in Costa Rica. Brenesia, 6: 25-31. MICKEL, J. T. 1980. Relationships of the di sected elaphoglossoid ferns. Brittonia, 32: 1 0< 117. MICKEL, J.T., AND L.ATEHORTUAG. 1980. Sul division of the genus Elaphoglossum. Amer. Fei J., 70: 47-68. MORTON, C. V. 1955. Notes on Elaphoglossw III. The publication of Elaphoglossum ar Rhipidopteris. Amer. Fern J., 45: 1 1-14. UNDERWOOD, L. M. 1905. A much-named fer Torreya, 5: 88. Key to Species of Peltapteris a. Sterile lamina flabellate, either entire or dissected; fertile lamina orbicular to notched at apex . . 2. P. pelta a. Sterile lamina pinnately divided, the pinnae linear, entire or dichotomously divided; fertile lamii linear b. Pinnae with 1 vein per segment, ca. 1 mm broad 1 . P. moor b. Pinnae often with more than 1 vein per segment, 2-3. 1 mm broad 3. P. peruviai 1. Peltapteris moorei (E. G. Britt.) Gomez, Bre- nesia 6: 29. 1975. Acrostichum moorei E. G. Britt., Mem. Torrey Bot. Club 4: 273. 1895. TYPE: Bolivia, Yungas, Bang 558 (holotype, NY!; isotype, us!). Elaphoglossum moorei (E. G. Britt.) Christ, Bull. Herb. Boissier2, 3: 148. 1903. Elaphoglossum bangii Christ, Monogr. Elaphoglos- sum 99. 1899. TYPE: Bolivia, Yungas, Bang 558 (holotype, presumably p). Microstaphyla bangii (Christ) Hieron., Bot. Jahrb. Syst. 34: 539. 1904. Rhipidopteris rusbyi Christ, Farnkr. der Erde 46. 1 897. TYPE: Bolivia, Yungas, Bang 558 (holotype, pre- sumably P). Microstaphyla moorei (E. G. Britt.) Underw., Torre 5: 88. 1905. Stem long-creeping, ca. 1 mm in diameter, seal lanceolate, orange-tan, 1-2 mm long, entire. Phj lopodia lacking. Leaves 3-10 mm apart, 6-14 c long, 1 .5-2.9 cm broad. Petiole y3-V2 the leaf lengt scales orange-tan, ascending, ca. 1 mm long, e: tire. Lamina narrowly lanceolate, chartaceous, ap< acuminate, base truncate, pinnate, the pinnae ei tire (linear) to furcate, segments 1-1.5 mm broa Veins obscure, free, 1 per segment. Hydathod lacking. Lamina scales scattered, orange-tan, Hi ear-lanceolate-deltate, more commonly abaxiall TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 16 FIG. 26. Peltapteris peltata f. peltata: a, habit; b, fertile lamina, abaxial side; c, fertile lamina, adaxial side. (From Stolze, Ferns & fern allies of Guatemala, 1981.) 168 FIELDIANA: BOTANY Fertile leaves shorter than the sterile, petiole % the leaf length, lamina linear, entire; intersporangial scales lacking. Epiphytic in wet forests, 1750-2400 m, Cuzco. Cuzco: Cerro Chuyapi, BiiesA41 (NY). La Convention, alrededores de Tunguimayo (Itma), Vargas C. 10669 (GH). La Convention, Cordillera Vilcabamba, Knox's Cascade, Dudley 10490 (GH). 2. Peltapteris peltata (Sw.) Morton, Amer. Fern J. 45: 13. 1955. Acrostichum peltatum (Sw.) Sw., J. Bot. (Schrad.) 1 80 (2): 11. 1802. Rhipidopteris peltata (Sw.) Fee, Mem. foug. 2: 78. 1 84! Elaphoglossum peltatum (Sw.) Urban, Symb. antil 4: 60. 1903. Stem long-creeping, slender, ca. 1 mm in di ameter, scales lanceolate, tan. Leaves dimorphic sterile ones 3-15 cm long, distant. Petiole aboi % the sterile leaf length, sparsely scaly. Lamin flabellate, 2.5-5 cm broad, undivided to 5 time dichotomously divided, surfaces with a few smal tan scales. Fertile leaves longer than the sterile an nearly undivided, usually 2-lobed, 5-20 mm broac Osmunda peltata Sw., Prodr. 127. 1788. TYPE: Ja- maica, Swart z (holotype, s). Key to Forms a. Sterile lamina 4-5 times dichotomously divided 2a. f. peltat a. Sterile lamina entire or dichotomously divided into two parts 2b. f. flabellat 2a. Peltapteris peltata f. peltata. Figure 26. Sterile lamina mostly 4-5 times dichotomously divided, the segments linear, 0.5-1.5 mm broad. Epiphytic in wet forests, 400-1400 m, Ama- zonas, San Martin, Pasco, Junin, Ucayali. Mexico to Panama; West Indies; Guianas; Ven- ezuela and Colombia to Peru. Amazonas: Bagua, Montenegro-Chiriaco, Sagdstegui 5932 (GH). Huampami & Shaim, Berlin 445 (us). San Martin: Boqueron Pass, 92 km from Tingo Maria on highway to Pucallpa, Allard 22122 (GH, us). Pasco (as Junin): Pichis Trail, San Nicolas, Killip & Smith 26016 (GH, NY). Junin: Schunke Hacienda, above San Ramon, C. Schunke A 149 (us). Hacienda Schunke, La Merced, Macbride 5809 (us). Chanchamayo Valley, C. Schunke 460 (us). Ucayali: Coronel Portillo (as Loreto), Padre Abad, Boqueron de Padre Abad, /. Schunke V. 3047 (GH, NY, us). Coronel Portillo, Km 209 entre Tingo Ma- ria y Pucallpa, Aguaytia, Ridoutt, 31 July 1943 (GH). Coronel Portillo (as Huanuco), Fundo Chela, Sinchono, Aguilar 918 (USM). 2b. Peltapteris peltata f. flabellata (Willd.) Go- mez, Brenesia 6: 28. 1975. Acrostichum flabellatum Willd., Sp. pi. 5: 1 10. 1810. TYPE: America meridionali, Humboldt & Bon- pland (holotype, B!, Herb. Willd. 19530). Acrostichum flabellatum var. sphenophyllum Kunz Linnaea 9: 32. 1834. TYPE: Peru, P oeppig (h< lotype, LZ, destroyed; isotype, PR; frag., NY!). Acrostichum sphenophyllum (Kunze) Kunze, Analeci pteridogr. 11, t. 7. 1837. Rhipidopteris flabellata (Willd.) Fee, Mem. foug., '. 78, 1845. Rhipidopteris sphenophylla (Kunze) Fee, Mem. fouj 2: 79. 1845. Elaphoglossum peltatum f. flabellatum (Willd.) Mid el, Brittonia32: 116. 1980. Similar to f. peltata except in f. flabellata th sterile lamina is less divided, either entire or i two lobes. Epiphytic in wet forests, 700-2200 m, Am£ zonas to Cuzco and Madre de Dios. Costa Rica; Panama; Venezuela and Colombi to Peru. Amazonas: Mendoza, Woytkowski 8297 (GH). Sa Martin: In monte Campana prope Tarapoto, Spruce 463 (NY, us). Pasco: Oxapampa, Cordillera San Matias, Led 323 (USM). Prov. Oxapampa, Abra los Mellizos, Skog < al. 5039 (us). Junin: Villa Amoretti, near La Mercec Kunkel 632 (GH). Pichis trail, Porvenir, Killip & Smit 25903 (us). Ayacucho: Ayna, between Huanta & Ri Apurimac, Killip & Smith 22781 (NY, us). Ccarraps between Huanta & Rio Apurimac, Killip & Smith 2320 (GH, NY, us). Cuzco: Paucartambo, Pillawata, Yana mayo-Tombomayo, Vargas C. 16733 (GH). Prov. Pat cartambo, San Pedro a San Isabel, Vargas C. 6786 (us TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 16' Rio Caracol, Valle de San Miguel, La Convention, Biies This closely resembles P. moorei in general ar- 20/6 (us). Madre de Dies: Manu, Cerro de Pantiacolla, chitecture, but has the pinnae notched and forked, Rio Palotoa, 10-1 5 km NNW of Shintuya, Foster 10894 often whh more than Qne vdn per segment Endemic. Epiphytic in wet forests, ca. 3050 m, Cuzco. Thus far known only from the type. 3. Peltaptens peruviana Gomez, Rev. Biol. Trop. 18: 217. 1971. TYPE: Peru, Cuzco, Alturas de Sicre, Biies 1572 (holotype, us!; isotype, cuz). 170 FIELDIANA: BOTANY Colombia 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. Tumbes Piura Lambayeque Cajamarca Amazonas La Libertad San Martin Loreto Ancash Huanuco Lima Pasco JunTn Ucayali lea Huancavelica Ayacucho Apurimac Cuzco Madre de Dios Arequipa Puno Moquegua Tacna Chile TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 17 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 accepted name or synonym occurs. Acrostichum acuminatum 154 adenolepis 166 albescens 121 alienum 100 apodum var. sprucei 157 auricomum 125 bakeri 125 barbatum 125 calaguala 137 calophyllum 166 castanewn 127 caudatutn 64 caudatum Hooker, 1 840. 1 54 caudatum Hooker, 1864. 64 cervinum 85 chrysolepis 6 1 ciliatum 128 cladotrichum 125 curvans 166 cuspidatum 129 decoratum 129 denticulatum 131 discolor 130 dissimile 166 diversifolium 130 elongatum 162 engelii 131 erinaceum 132 erythrolepis 132 eximium 133 flabellatum 169 var. stenophyllum 169 Jlaccidum 133 fractiseriale 60 furfuraceum 164 glabellum 134 guamanianum 135 hackelianum 61 hartwegii 135 hayesii 136 haynaldii 136 hickenii 136 horridulum 137 huacsaro 137 Ay«r/;c 138 ilvense 94 insigne 6 1 japurense 107 juglandifolium 61 laminarioides 139 latifolium 140 lechlerianum 141 lechlerianum (Melt.) Hooker 63 leprosum 142 leptophyllum 146 lindenii 142 lindigii 103 lingua 142 litanum 143 luridum 144 mathewsii 144 minutum 146 moorei 167 muscosum 147 nicotianifolium 102 nigrescens 148 nivosum 148 oligarchicum 102 orbignyanum 151 osmundaceum 62 pachyphyllum 151 paleaceum 152 pandurifolium 105 papillosum 152 patinii 153 peltatum 169 petiolosum 154 phlebodes 107 piloselloides 154 plumbicaule 60 plumosum 155 poeppigianum 156 poly hot ryoides 6 1 preslianum 128 propinquum 156 quitense 157 raywaense 157 rimbachii 158 schomburgkii 144 serratifolium 101 serratum 101 setigerum 160 sorbifolium 105 sphenophyllum 169 squamipes 161 stenopyllum 161 suberectwn 61 tambillense 162 tectum 162 tenuiculum 163 unitum 146 villosum var. poeppigianum 156 Adiantum lunulatum 40 Allantodia asplenioides 75 Anisogonium 67 fraxinifolium 67 pinnatifidum 89 Arachniodes 35 aspidioides 35 denticulata 37 ochropteroides 38 Aspidium 21 articulatum 96 braunianum 24 catocarpum 10 confertum 47 ctenitis 5 dicksonioides 21 draconopterum 27 excultum 35 funestum 21 gelidum 54 guianense 48 heracleifolium 27 karstenii 9 kunzei 29 macrophyllum var. decurrens 29 martinicensis 24 meniscioides 48 microchlaena 9 nemophilum 10 «0Me 38 nodosum 96 orbiculatum 53 paleaceum 36 parallelogramma 36 patulum 36 pendulum 98 platyphylum 52 poeppigii 29 protensum 19 pycnolepis 54 rotundatum 42 trianae 47 truncatulum 40 Asplenium alienum 12 arboreum 80 callipteris 83 caracasanum 79 caucense 85 celtidifolium 83 centripetale 87 crassifolium 76 denticulosum 80 desvauxii 11 eggersii 87 expansum 75 ferulaceum 90 flavescens 85 flexuosum 11 fuscopubescens 12 juglandifolium 85 kunzei 87 fec/zfcn 85 lindbergii 78 macrophyllum 11 marginatum 90 ocanniense 85 pinnatifidum 87 172 FIELDIANA: BOTANY plantagineum 86 plantaginifolium 86 procerum 77 roemerianum 85 sandwichianum 72 shepherdii 80 striatum 79 var. caracasanum 79 tungurahuae 76 vastum 73 venulosum 76 Athyrium 88 ambiguum 71 "bradearum" 88 celtidifolium 83 dombei 90 dombeyi 90 expansion 75 ferulaceum 90 filix-femina 88 var. dombeyi 90 flexuosum 77 fumaroides 92 praestans 87 Bolbitis 98 aliena 100 bradeorum 103 crenata 101 guianensis 109 ft////?// 102 lindigii 103 nicotianifolia 102 oligarchica 102 pandurifolia 105 portoricensis 101 serrata 101 serratifolia 101 stuebelii 103 Byrsopteris 35 aristata 35 Cheilanthes crenata 94 Chrysodium lindigii 103 serratum 101 Cnemidaria speciosa 15 Ctenitis 5 ampla of authors 10 ampla (Willd.) Ching 10 andicola 19 biserialis 15 catocarpa 10 distans 5 hirsute- setosa 16 honesta 14 karsteniana 17 microchlaena 9 mollicoma 17 nemophila 10 nigrovenia 9 var. funesta 2 1 pulverulenta 17 refulgens 7 sloanei 10 subincisa 10 submarginalis 8 Cyclodium 47 guianense 48 meniscioides 48 var. meniscioides 48 var. paludosum 48 trianae 47 var. chocoense 47 var. trianae 47 Cyclopeltis 29 semicordata 29 Cyrtomium 38 dubium 38 falcatum 40 nobile 38 Cystopteris 92 subg. Acystopteris 92 subg. Cystopteris 92 fragilis 92 translucens 92 Dicksonia montevidensis 94 Didymochlaena 40 lunulata 40 sinuosa 40 truncatula 40 Diplazium 65 subg. Anisogonium 67 aberrans 87 alienum 72 altissimum 70 ambiguum 71 var. pubescens 1 1 angelipolitanum 79 appolinaris 77 arboreum 80 asplenioides 75 bicolor 74 bogotense 70 bombonasae 8 1 bonapartii 75 bradeorum 88 brasiliense 79 buchtienii 72 callipteris 83 caracasanum 79 celtidifolium 83 var. celtidifolium 83 var. puberulum 84 centripetale 88 chimborazense 87 costale 77 crassifolium 76 cristatum 80 cuneifolium 82 delitescens 67 denticulosum 80 diplazioides 7 1 divergens 70 drepanolobium 8 1 eggersii 88 expansum 75 ferulaceum 90 flavescens 85 var. proliferum 85 flexuosum 76 fraxinifolium 67 fuscopubescens 72 fuscum 87 gracilescens 72 grande 78 grandifolium 82 var. andicola 82 var. grandifolium 82 hians 70 induratum 78 lechleri 85 legalloi 84 lehmannii 72 lindbergii 78 lonchophyllum 80 macrodictyon 87 macrophyllum 77 melanosorum 75 moritzianum 73 obscurum 85 obtusum 79 oxylobum 77 pactile 87 paucijugum 84 pedatum 74 pinnatifidum 86 plantagineum 86 plantaginifolium 86 praestans 87 preslianum 76 remotum 74 riedelianum 85 roemerianum 85 rostratum 75 sandwichianum 72 shepherdii 80 var. proliferum 79 var. stuebeliana 80 striatum 79 stuebelianum 80 subnudum 78 subobtusum 78 tabalosense 79 tarapotense 73 tungurahuae 76 unilobum 81 vastum 73 venulosum 75 verapax 85 werckleanum 80 Dryopteridaceae 2 Tribe Bolbitideae 3 Tribe Dryopterideae 3 Tribe Oleandreae 3 Tribe Physematieae 3 Dryopteris 35 subg. Ctenitis 5 subg. Dryopteris 35 subg. Polystichopsis 35 TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 17 subg. Stigmatopteris 42 ampla 10 andicola 19 f. lehmanniana 19 f. spruceana 19 arista ta 35 biserialis 15 catocarpa 10 christii 45 ctenitis 5 denticulate 37 fibrillosa 13 filix-mas 35 heterophlebia 45 hirsuto-setosa 16 honesta 14 karsteniana 17 karstenii 9 A://///)// 33 leptosora 14 longicaudata 44 macrostegia 38 microchlaena 9 microsora 14 mollicoma 17 nemophila 10 nigrovenia 9 ochropteroides 38 opaca 45 paleacea 36 paludosa 48 parallelogramma 36 patula 36 paucinervata 45 pellucido-punctata 44 platyloba 15 prasinna 45 protensa 21 var. dicksonioides 2 1 var.funesta 21 pubescens 35 pulverulenta 17 refulgens 1 var. peruviana 8 safTordii 37 subincisa 16 submarginalis 8 var. lagerheimii 8 var. tarapotensis 8 tarapotensis 15 vo«a 17 wallichiana 36 yungensis 14 Elaphoglossum 1 1 1 sect. Elaphoglossum subsect. Pachyglossa 123 a$/«e 147 albescens 121 alipes 122 amazonicum 122 amphioxys 123 amplum 123 angustius 124 apodum var. towra 123 atropunctatum 124 atrosquamatum 124 auricomum 125 bakeri 125 bangii 167 barbatum 125 blepharoglottis 126 calaguala 137 camptolepis 126 cardenasii 126 castaneum 127 caudatum 154 chloodes 127 ciliatum 128 concinnum 128 conforme 1 1 1 craspedotum 128 crinipes 160 cuspidatum 129 decoratum 129 denticulatum 131 dichroum 129 discolor 130 diversifrons 130 eatonianum 127 elegantipes 131 elongatum 162 engelii 131 ensiforme 131 erinaceum 132 var. boliviensis 132 erythrolepis 132 eximium 133 flaccidum 133 fortipes 133 glabellum 134 glossophyllum 134 glutinosum 154 gracillimum 134 guamanianum 135 hartwegii 135 hayesii 136 haynaldii 136 hickenii 136 hieracioides 137 "hikenii" 137 horridulum 137 huacsaro 137 hystrix 138 j urn ml urn 138 killipii 138 laminarioides 139 lanatum 139 lasioglottis 139 latevagans 140 latifolium 140 latum 123 lawyerae 141 la \isquama 141 lechlerianum 141 leprosum 142 leptophyllum 146 lindbergii var. truncatum 126 lindenii 142 lingua 142 linguaeforme 134 litanum 143 license 132 longipes 144 longius 143 luridum 144 mad 1 en turn 144 mathewsii 144 megalurum 145 megarhizon 146 m c lade ilium 145 melancholicum 145 metallicum 146 minutum 146 molle 152 moorei 167 moyeri 147 muscosum 147 nastukiae 148 nidiformis 148 nigrescens 148 nigrocostatum 140 nivosum 148 obovatum 149 obtusum 149 oculatum 150 odontolepis 150 oophyllum 150 orbignyanum 150 ornatum 149 oxyglossum 151 pachyphyllum 151 pachyrrhizum 152 paleaceum 152 palorense 153 papillosum 153 pascoense 153 patinii 153 pattersoniae 166 peltatum 169 f. flabellatum 169 petiolosum 154 pichinchae 158 piloselloides 154 pilosius 155 plicatum 152 plumosum 155 poeppigianum 156 potomogeton 166 preslianum 128 propinquum 156 pseudohirtum 158 p ii mil io 156 punae 157 quitense 157 raywaense 157 rimbachii 158 rosenstockii 158 rubellum 159 ruficomus 159 rufum 159 russelliae 160 schomburgkii 144 setigerum 160 siliquoides 136 174 FIELDIANA: BOTANY simulans 160 spatulatum 155 squamipes 161 stenophyllum 161 styriacum 161 subciliatum 143 tambillense 162 tectum 162 tenue 163 tenuiculum 163 tomentellum 163 velongum 164 vittarioides 164 vulcanicum 164 wardiae 165 williamsii 123 williamsiorum 165 zebrinum 165 Gymnopteris aliena 100 nicotianifolia 102 pandurifolia 105 Hemidictyum 90 marginatum 90 Hemionitis plantaginea 85 Hymenodiwn kunzeanum 151 Hypoderris stuebelii 103 Jamesonia paleacea 54 Lastrea subsect. IPolystichopsis 35 pubescens 31 recedens 31 Lastreopsis 3 1 amplissima 33 effusa 33 ssp. divergens 33 exculta 35 killipii 33 recedens 31 tenera 3 1 Leptochilus alienus 100 bradeorum 103 crenatus 101 guianensis 109 killipii 102 lindigii 103 nicotianifolius 102 oligarchicus 103 pandurifolins 105 serratifolius 101 serratus 101 stuebelii 103 Lomagramma 109 guianensis 109 Lomariopsis 105 erythrodes 107 fendleri 106 japurensis 106 latipinna 107 nigropaleata 108 sorbifolia 105 vestita 106 Megalastrum 1 1 andicola 18 f. andicola 19 f. loh m a n nia nu m 19 biseriale 15 hirsutosetosum 16 honestum 13 leptosorum 14 microsorum 14 mollicomum 17 pansamalense 18 platylobum 15 pulverulentum 17 var. heydei 18 var. pulverulentum 18 spectabile 16 subincisum 16 vastum 17 villosum 1 1 yungense 14 Meniscium cristatum 80 opacum 45 Microstaphyla bangii 167 moorei 167 Nephrodium antioquoianum 26 firmifolium 47 funestum 21 lagerheimii 8 lizarzaburui 25 microsorum 14 nigrovenium 9 ochropteroides 38 polyphyllum 53 sodiroi 26 tarapotense 8 trapezoides 53 villosum 18 var. opacum 19 f. lehmannianum 19 f. spruceanum 19 Oleandra 96 articulata 96 distenta 96 hirta 98 lehmannii 97 micans 98 neriiformis 96 nodosa 96 pilosa 98 Olfersia 55 caudata 64 cervina 57 ciliata 128 corcovadensis 57 Osmunda cervina 57 peltata 169 Peltapteris 167 moorei 167 peltata 169 f. flabellata 169 f. peltata 169 peruviana 170 Peltochlaena 47 Phanerophlebia 38 nobilis 38 Phegopteris cochleata 55 dictyophylla 38 dubia 38 lechleri 45 pycnolepis 55 refulgens 1 Plecosorus 49 mexicanus 49 speciosissimus 49 Poecilopteris crenata 101 Polybotrya 57 subg. Polybotrya 57 subg. Sorbifolia 57 subg. Soromanes 57 aequatoriana 64 alfredii 65 altescandens 6 1 andina 6 1 appressa 64 caudata 64 cervina 57 crassirhizoma 60 decor ata 62 fractiserialis 60 fulvastrigosa 65 glandulosa 63 hickeyi 65 juglandifolia 61 kalbreyeri 6 1 lechleriana 63 lomarioides 65 macbridei 60 nutans 65 osmundacea 62 plumbicaulis 60 polybotryoides 6 1 pubens Kunze 62 pubens Martius 62 puberulenta 63 serratifolia 57 sorbifolia 57 subelliptica 63 suberecta 6 1 TRYON & STOLZE: PTERIDOPHYTA OF PERU. IV. 17 Polypodiaceae Tribe Aspidiae 2 Polypodium adiantiforme 3 1 amplum 10 aristatum 35 biseriale 15 cochleatum 54 crenatodentatum 54 denticulalum 37 effusum 33 falcatum 40 fibrillosum 13 fiJix-femina 88 filix-mas 35 flavopunctatum 42 /ra#/7